The Fungi Which Cause Plant Diseases

THE FUNGI WHICH CAUSE PLANT DISEASE
BY
F. L.
STEVENS,
Ph. D.
PROFESSOR OF VEGETABLE PATHOLOGY AND DEAN, COLLEGE OF AGRICULTURE AND MECHANIC ARTS, MAYAGUEZ, PORTO RICO. FORMERLY OF THE NORTH CAROLINA COLLEGE OF AGRICULTURE ALSO FORMERLY PRESIDENT OF THE AMERICAN PHYTOPATHOLOGICAL SOCIETY
Nnn f nrk
THE MACMILLAN COMPANY

1913
All rights reaerved
Reprinted with the permission of Mr. Frank D.
Murphy
LTD.
JOHNSON REPRINT CORPORATION

Ill Fifth Avenue,
JOHNSON REPRINT COMPANY
New
York, N.Y. 10003
Berkeley Square House, London, W.I
COPTRIOHT, 1913
By
the MACMILLAN COMPANY

Published November, 1913
Set up and electrotyped.
^J-W;
First reprinting,
1966, Johnson Reprint Corporation
Printed in the United States of America
GARDEIi
TO
MY WIFE ADELINE CHAPMAN STEVENS

IN
ACKNOWLEDGMENT
OF
HELP,
ENCOURAGEMENT
AND INSPIRATION
PREFACE
This volume
is
intended to introduce to the student the more
important cryptogamic parasites affecting economic plants in the United States, with sufficient keys and descriptions to enable their identification. Technical description of each division, order,
family, genus and species when important is given unless the essential characters are to be clearly inferred from preceding keys or text. Gross descriptions of the host as diseased, i, e., of the
disease
itself, have been avoided since such are to be found in "Diseases of Economic Plants." Effort has been made to avoid Abundant duplication of matter contained in that volume.
citations to the
more important papers are given, sufficient, it is believed, to put the student in touch with the literature of the
While
subject.
many parasites not yet known in the United States are mentioned, especially the more important ones or those which are likely to invade America, no attempt has been made to
briefly
list all
of these.
Non-parasitic groups closely related to those that
are parasitic have been introduced in the keys merely to give a larger perspective to the student.
Effort has been
genus that
is
of
to give at least one illustration of each importance in the United States.
made
The author
is
indebted for descriptions, keys,
etc.,
to the
various standard works.
Those which have been drawn upon most largely are Saccardo's Sylloge Fungorum, Die NatUrlichen
Pflanzenfamilien of Engler
&
Prantl, Clinton's Ustilaginales of
North America, Clement's Genera of Fungi, and Minnesota Mushrooms, Plowright's British Uredineae and Ustilagineae, Arthur and Murrill each in North American Flora. The author wishes also to express thanks for suggestions and
criticism of the manuscript to T.
H. Macbride, who read the por-
tion
on Myxomycetes;
J. J.
Davis, Phycomycetes; L. R. Jones

vii
viii
PREFACE
and T. J. Burrill, Bacteria; G. M. Reed, Perisporiales; G. P. Clinton, Ustilaginales; J. L. Siieldon, Ascomycetes in part; D. Reddick, Ascomycetes in part; J. C. Arthur, Uredinales; F. D. Heald, Fungi Imperfecti in part; F. C. Stewart, Fungi Imperfecti in part;
H. Metcalf, Basidiomycetes in part; to Mrs. Flora W. Patterson for aid in securing descriptions otherwise unobtainable; to Dr. Marshall Avery Howe for assistance with the glossary; to Messrs. Norton, Rosenkranz and Fawcett, for aid in proofreading and in preparation of the manuscript, though no responsibiUty for error attaches to those who have so kindly
aided.
It is probable,
owing to the present unsatisfactory condition of
of the fungi, loose and imperfect description of species, disregard of generic limitation, lack of knowledge regarding the limits of specific variation, influence of environment, biologic host
taxonomy
relations, etc., that
many
of the species treated in the text are
has, however, attempted so far as possible to reflect the facts as they appear in the light of present knowledge and has deemed it more useful to err on the side of conservatism
untenable.
The author
than to attempt to reduce the apparent number of species by consolidation without full and complete evidence as to the real identity
of the species in question.
F. L. Stevens.
Mayagijez, Porto Rico.
CONTENTS
PAGE
Introduction
Division Division
I,
Myxomycetes
5
13
II, Schizomycetes Bibliography of Introduction, Myxomycetes and Schizomycetes
53
Division
III, Eumycetes Class Phycomycetes Bibliography of Phycomycetes Class Ascomycetes Bibliography of Ascomycetes
59 G5
103
113
288 298 466 475 666 678 681 697
Class Basidiomycetes Bibliography of Basidiomycetes Fungi Imperfecti Bibliography of Fungi Imperfecti Bibliography of Books and Periodicals Glossary Index
iz
INTRODUCTION
The principal non-flowering vegetable parasites which cause plant diseases belong to three divisions: the Slime Molds (Myxomycetes) ; the Bacteria (Schizomycetes) ; and the True Fungi
(Eumycetes including the Phycomycetes). The term fungi, in the broad sense, is often used to include all three of these divisions. All are devoid of chlorophyll and therefore all differ from the green
plants in the essential ways which result from this deficiency. Transpiration, respiration, and true assimilation are the same as with the green plants, but photosynthesis or starch manufacture cannot be accomplished by them. Sunlight being thus useless to
them
no
directly they can live in the dark as well as the light.
Having
foods from inorganic matter these organisms are limited to such nutriment as they can obtain from plants or animals which have elaborated it; that is, they must
ability to elaborate their
own
have organic foods for their sustenance. The fungi have acquired various food habits and adapted themselves to different methods of nutrition. Some are nearly omnivorous and can subsist upon almost any decaying tissue or upon
Others thrive only or solutions rich with organic debris. special substances, as for example, some particular plant or animal, the host, perhaps only upon some particular part of that
soils
upon
tfiat prey upon living things are Those living upon dead things are saprophytes. No hard and fast line can be drawn between these two classes. An organism which is usually a saprophyte may live upon a dead member of some plant, gradually encroach upon the still living part and thus become partially a parasite. Again there
plant or animal.
called
The organisms
parasites.
are times in the history of a plant

difficult to tell
when
life
ebbs so low that
it is
the living from the dead.

1
The pulp
of the apple
ripe, a resting seed, the cells of the potato tuber in winter, are undoubtedly alive, yet their activity is so little that many organisms can gain a foothold upon these stages of the plant
when
that
ence.
cannot do so at more vigorous periods of
their
exist-
Tubeuf
are parasites, but
ranks as hemi-parasites those organisms that usually may sometimes become saprophytic, and as
hemi-saprophytes such as are usually parasitic, but may excepThese distinctions are of little tionally become saprophytic. import, other than to bring out clearly that each species has its
own
limits as to food requirements.
be thought that these parasites and saprophytes have always been dependent organisms. The true fungi for example are best to be regarded as degraded descendants of algse, in which ancestors they once possessed chlorophyll and could prepare their own food from mineral matter by the aid of sunIt is hardly to
light.
No discussion of the general metabolic processes of the fungi is here necessary further than to indicate that among the products of their activity there are various excretions and secretions, which
bear important relations to parasitism. Thus certain fungi growing in artificial culture produce enzymes or organic ferments
capable of softening and dissolving cellulose, also toxins, poisons which are capable of killing the cells of the host plant. Such enzymes and toxins are numerous and their bearing upon parasitism
is
obvious.
They enable the
parasite to
kill
adjacent
cells
and then to effect an entrance through the cell walls to the protoplasm and other nutrients contained within the
of the host
cell.
of the parasite, or secretions produced by it, often abnormal growth responses from the host. These take very diverse forms, either the undergrowth or overgrowth, hypertrophy, of single cells or tissues, or even the excessive development of large plant parts as in the case of the witches' brooms, and the "double flowering" of the dewberry. The probable relations of the groups under consideration to the
calls forth
The presence
other
members
of the Thallophyta are suggested in the following
scheme.

Bacteria, Schizomycetes.
CVANOPHYCE^,
Blue-green Algae.
Myxomycetes, Slime Fungi.

PERIDINE.E, Dinoflagellates.
Conjugates.
DlATOME^. (Conjugate,
Heterocont^.
^"^^^CHLOROPHYCEiE, Green
I
Algae.
"<;;harace^, Stoneworts.
Algae.
^^^RhodophycevE, Red
^^^^"Phycomycetes* Alga-like Fungi.

^^^Phjeophyce^,
Key
Brown
Algae.
^^îÊuMYCETES,
Fungi.
to the three Divisions important as plant parasites:
Vegetative body a multinucleate naked Plasmodium Division I. Myxomycetes,
p. 5.
Vegetative body a single-walled cell, nucleus absent or not of the form typical in the other fungi, reproduction by fission (by conidia in a few non-parasitic forms).. .Division II. Schizomycetes, p. 13.
Not
as above: Vegetative body usually filamentous, reproduction by various means Division III. Eumycetes, p. 59.
DIVISION
FUNGI"-**
MYXOMYCETES, SLIME MOLDS, SLIME

(p.
3)
These are the lowest organisms considered by the botanist, and partake so much of the nature of both animals and plants that their position has long been debated. Their affinities are with the lowest living things, on the boundary between the animal and the vegetable kingdom, and sometimes more attention is accorded them by the zoologist than by the botanist.
The distinctive character of this group is that the vegetative condition consists either of distinct amoeboid cells or of a mass of
naked protoplasm, the Plasmodium, composed of numerous
units,
cell
at the completion of the free vegetative stage, produce numerous walled spores either free or in sporangia of various forms. The spores upon germination
each unwalled.
The plasmodia,
produce either zoospores or amoeboid bodies which multiply and unite to form either new plasmodia or pseudoplasmodia. The slime molds consist of three orders:
Key
Parasitic
to Orders of Myxomycetes
1.
Plasmodiophorales,
Acrasiales
p. 5.
Saprophytic
Vegetative phase of free amoebse
2. 3.
Vegetative phase plasmodial
Myxogastrales,
p. 9.
The
Acrasiales contain
some
five
genera and ten species purely
saprophytic.
Plasmodiophorales
Intracellular
parasites;
vegetative
stage
plasmodial;
spores
formed by the simultaneous breaking up of the Plasmodium into an indefinite number of independent cells.
5

The Plasmodiophorales appear
to include all of the true para-
sites of
the Myxomycetes.
Key to Geneba of
Spores
free, spherical
Plasmodiophorales
1.
Plasmodiophora,
p. 6.
Spores united into groups Spores in groups of four

Spores in larger groups Spores forming a hollow sphere. ... Spores forming a spongy spore-ball
2.
Tetramyxa,
p. 8.
3. 4.
Sorosphaera, p.
8.
Spongospora,
p. 8.
Plasmodiophora Woronin
This genus
of plants,
is
parasitic in the living

filling
the plasmodia
the
cells
parenchyma of the roots and causing galls at

of the
the point of attack. Europe and America.

P. brassicse Wor.^"*'
There are three species

200-203, 208
genus in
^^^^ j^j^g ^ggj^
known
as a parasite
and recent work indicates that other ^^^ families, as the Umbelliferae and cucurbs, are also susceptible. The parasitised cells especially, and the adjacent cells as well, are stimulated to enormous overgrowth; this hypertrophy resulton the
crucifers generally
ing in a characteristic root "clubbing." Study of diseased sections shows that the medullary rays and cortex are abnormally thick (hypertrophy and hyperplasia) and
many
of their cells are parasitized.
pressed by creased proportionately.

less
the parasite and the xylem
Sclerenchyma cells are supis reduced and phloem inof stored starch
is
The amount
much
than in normal tissues. Infection does not appear to pass from cell to cell but groups of diseased cells are thought to arise from repeated division of a cell
after its infection.
In the enlarged host cells the protoplasm appears abnormally dense and fine grained. Eventually the whole lumen of the cell is occupied by the crowded, amoeboid, individuals, each uninucleate
and unwalled, and still distinct from the later fuse into a Plasmodium the nuclei
These individuals which enlarge and undergo simultaneous mitotic division. Still later the mass divides into uninuclear segments each of which matures to a spore 1.6 m
other.
of in diameter, covered
by a
thin,
smooth, colorless membrane.
The decay of the host liberates the spores in the soil. Their germination may be readily studied upon a microscope slide where in from five to twenty-four hours uninucleate zoospores
are produced.
The zoospores
are differentiated into an inner
Fio.
cabbage cells occupied by the unicellular parasite; 5, later stage, parasite many-nucleate; 10, host cell full of spores; 11, germinating
1.
P.
10
brassicse: 3,
11
spores.
After Lotsy.
granular part and an outer hyaline part, the hyaloplasm, which may extend to form pseudopodia, thus giving the cell an amoeboid movement in addition to that due to the single long cilium. Infection by these swarm spores is supposed to occur through the
root hairs though the
mode
of
primary infection
is
not definitely
8
knovMi.
Seedlings raised in soil inoculated with chopped roots bearing the disease become badly diseased as do also seedlings
upon which
infected water
is
is
poured.
^
P. humili Kirk
mentioned by Kirk
^
;
as the cause of club
root of hops in New Zealand. P. vitis Viala & Sauvageau
P. calif ornica Viala

^^
P. orchidis Massee
and P. tomato Abbey
& Sauvageau have been reported

;
as the causes of serious diseases but their relation to the diseases
and even
tioned.12-14
their
identity as actual organisms
is
seriously ques-
Tetramyxa Goebel grows upon water

Sorosphaera Schroter
plants, notably Ruppia.^
(p. 6)
Parasitic in the parenchyma of living plants; spores ellipticwedge shaped, forming a hollow, spherical spore ball. One species is found upon Veronica f a second species has been reported upon tea.^'' '^^ on the roots S. graminis Schwartz is reported by Schwartz and other grasses where it caused nodules much resembling of Poa
those of nematodes.
Spongospora Brunchorst
(p. 6)
Similar to Sorosphaera but the spores forming a spore ball
with open reticulations. S. subterranea (Wallr.)
Lag.^^''^ causes the
powdery scab
of
potatoes in Great Britain, Europe and South America. It has ^^ been closely studied by Osborne who shows it to appear first in
the tuber
cells
as a uninucleate
myxamceba which
ultimately
develops into a multinucleate amoeboid Plasmodium. Sorolpidium Nemec is a new genus with the species.
S. betas
Several
Nemec which little known
is
on beets. ^^^
genera, kin to the above, attack algse,
fungi, pollen, etc.
Pseudomonas radicicola, the legume tubercle organism has been by some placed in this order under the name Phytomjoca leguminosarum.
Myxogastrales (p. 5) This order comprises some forty-seven genera and four hundred species of great variety and beauty. The Plasmodium,
which varies from a millimeter or
less
(
to several decimeters in diameter, produces either flat encrusted masses of

spores, aethalia, or develops
nJ,/
"\
\
"^^^ Ih^^'^-'A
_
spores in
/^
îst;/
sporangia which show some superficial resemblance to very small puffballs, The interior of the sporanFig. 2.
<^X
Zi*^.
gium
like
is
structure,
often permeated by a threadthe capillitium. They
^'tu^-^^S'lX^ôSt^
^î^.
After Macbnde.
are not parasites but occasionally injure plants
by overgrowing
them.
Key to
externally
Families of Myxogastrales
Spores not enclosed in a sporangium, borne
upon the
fruiting bodies.
..
1.
Ceratiomyxaceae.
Spores enclosed in a sporangium

Capillitium wanting, or very poorly de-
veloped Periderm of uniform thickness, rupturing irregularly

2.
Liceaceae.
Periderm of unequal thickness Periderm with a subapical thin
line,
3.
opening by an operculum Periderm unequally thick above, the

thin portions evanescent, leaving a network formed by the thicker
Orcadellaceae.
portions Capillitium well developed
4.
Cribrariaceae.
Calcareous deposits absent, or rarely present in the periderm

Capillitium of hollow, usually sculptured threads; spores light colored
o.
Trichiaceae.
Capillitium
smooth and usually much branched threads:

of
solid,
spores dark colored Fruiting bodies aethalioid or in-
10

definite, walls
fraying
out
into
poorly defined, a pseudo6.
capillitium
Reticulariaceae.
Sporangia definite, true capillitium more or less prominent

Fruiting bodies separate sporangia with columella and abimdant
capillitium
7.
Brefeldiaceae.
8.
Stemonitaceae.
Calcareous deposits present Capillitium not calcareous

9. Didymiaceae, p. 10. Capillitium simple 10. Spumariaceae, p. 11. Capillitium more intricate Fructification calcareous throughout 11. Physaraceae, p. 11.
Didymiaceae
Fructification of separate sporangia or plasmodiocarps, periderm simple or double, the outer calcareous; columella present or absent; capillitial threads thin, colorless or violet, arising from the base of the sporangium or passing from the columella to the peri-
derm, usually without calcareous deposits, which
if
present are
violet.
very small crystals; spores in mass black, spore walls
Key to Genera
of Didymiaceae
Calcareous deposits in the form of stellate crystals, frosting the surface
1.
Didymium,
Didenna.
p. 10.
Calcareous deposits not stellate. Calcareous deposits forming a superficial crust Calcareous deposits forming large superficial
scales
2.
3.
Lepidoderma.
Didymium
Schroter
Sporangia distinct, stipitate, sessile or even plasmodiocarpous, never sethalioid; the peridium thin, irregular in dehiscence, covered with a more or
less dense coating of calcareous crystals; columella more frequently present; capillitium of delicate threads, simple or sparingly branched, extending from the columella to the
peridial wall.
D. daedalium. B.
culture. ^^
&
Br.
is
occasionally injurious to melons in
11
Spumariaceae
(p. 10)
Sporangia separate or sethalioid; calcarious deposit in the periderm or columella, never in the capillitium; capillitium radiating from various points of the columella, branching and anastomosing
to form a network, the ultimate branchlets of which support the
periderm.
Key to Genera
Fructification of ordinary sporangia
Fructification sethalioid
of Spumariaceae
1.
Diachea.
2.
Spumaria,
p. 11.
Spumaria Persoon
Fructification a^thalioid, consisting generally of large cushion-
shaped masses covered without by a white foam-like crust; within, composed of numerous tubular sporangia, developed from a com-
mon
hypothallus, irregularly branched, contorted and more or less confluent; the peridial wall thin, delicate, frosted with stellate
lime crystals, which
mark
in section the boundaries of the several
sporangia; capillitium of delicate threads, generally only slightly
branched, terminating in the sporangial wall, marked with occasional swellings or thickenings. S. alba (Bui.) D. C. Like all other
members
its
of the order
fre-
the present species
is
not a parasite but
sethalia are
^^ and other plants in quently produced upon grass, strawberries such abundance as to cause more or less serious injury. The sporangia are fused into a large sethalium which is white or cream-
colored,
from
to 7 cm. long
and
half as thick.
(p. 10)
Physaraceae
Key to Genera
Fructification sethalioid
of Physaraceae
1.
Fxiligo, p. 12.
Fructification plasmodiocarpous or of distinct
sporangia
Peridium without lime

Plasmodiocarpous
Sporangia distinct Peridium calcareous, more or
less throughout CapilUtium calcareous throughout
2. 3.
Cienkowskia.
Leocarpus.
4.
Badhamia.
12

Capillitium in part hyaline Sporangium vaselike, or more or less tubular
Opening irregularly Opening by a lid

Sporangia various, dehiscence irregular Capillitium evenly branched; the calcareous nodes small, fusiform
Capillitium intricate
5.
Physarella.
6.
Craterium.
7.
Tilmadoche.
8.
Physarum,
12.
The species of Fuligo produce very large yellowish plasmodia which change to yellowish or brownish aethalia. Some are credited
with damage similar to that of the preceding species. ^^
Physarum Persoon
Sporangia plasmodiocarpous, sethalioid or distinct; the peridium usually simple, sometimes double, irregularly dehiscent, more or
less definitely calcareous;
capillitium a uni-
form irregular
the
nodes,
peridial wall.
net, dilated
and calcareous at
all
adherent
on
sides
to
the
P. cinereum (Batsch), Pers., the species most commonly reported as injurious, forms
its
tiny sessile, gray sporangia in great numbers on living plants,^"' '^^ often smother-
Fig.
3. Phy sporangium. Macbride.
them. The peridium is lime charged as a r u m ing After are also the nodes of the capillitium. The spores are brown or violet, and warty. P. bivalve F. has been noted as injuring young bean plants. ^^
Dendrophagus globosus Toumey was reported by Tourney

as the probable cause of crown gall, but such relation ful (p. 36). It is said to be closely related to Physarum.
is
^^
doubt-
DIVISION
II
(p, 3)
BACTERIA, SCHIZOMYCETES"'"-^'
Bacteria are extremely minute, unicellular organisms, which in

outline present three primary forms each of great simplicity,
namely the spheres (cocci), the straight rods (bacteria), the curved rods (spirilli).
In addition to these forms
^^
which
of
comprise
the
vast
0'
majority of
known species bacteria there are also

fila-
ô
Fig.
''
bacteria consisting of
4. The
three
mentous bodies,
either sim-
'''^''''' ^' '^^'
' ^p^^'^'^"
type forms of bacteria; ^^''' ^^-
In both structure and physbacteria are allied to the vegetable kingdom and in it iology most closely to the blue green algse. Bacteria are inconceivably small. Most of the spherical bacteria
pie or branched, attached or free.
within the limits of from 0.5 to 1.5/1 in diameter. Among the rod and curved bacteria the length in most species is between 1 and 1.5 n, the diameter between 0.5 and 1 n. Among the
fall
B. megatherium, 2.5 x 10 m; Clostridium butyriand Spirillum volutans, 13 to 50 n long. Among /z; the smallest is Spirillum parvum 0.1-0.3 ju in diameter and Pseudolargest species
is
cum, 3 X 10
monas
indigofera 0.06 x 0.18
fi.
It is practically impossible to conceive these dimensions.
An
these
illustration
may
is about 87.5 m thick. It would therefore take about 200 bacteria of ordiFiG. 5. This dot ia nary size or 400 moderately small or 20 very 1 mm. in diameter, -^ j^^^g^j^ j^^^.^^ ^^^^ pj^^^j ^^j ^^ ^^^^j ^^ ^q^^j 1571 ordinary bacteria the thickness of this paper. It would take (1 X 2 fi) end to end to reach around the circumference of a dot
aid the imagination. words are printed
The paper on which
13
14

in diameter.
(Fig. 5.)
if
mm.
placed side by side or 785,400
500 to reach across it; and 392,700 placed on end to cover its area, and
about 500,000,000 to fill a cube the edge of which is 1 miUimeter, making no allowance for lost space of the interstices. Considerably
more than 500,000,000,000 bacteria of this size would find room enough to move about in a space of one cubic centimeter.
The
typical
mode
of increase
among
except among sion of one cell, the mother

Fig. 6.
the sheath bacteria

cell,
bacteria
the only
mode
is
by
fission or direct divi-
into two, the daughter cells.
course
at
all,
The rapidity with which fission can proceed depends of upon conditions of environment, ranging from no growth
due to
cold, lack of nutriment, presence of inhibiting sub-
stance, to a
maximum
that varies with the species.
For bacteria
Fig.
6.
Diagram
^LA A UUIU
illustrating the fission of bacteria, bacilli
and
cocci.
After Novy.
in general
under very favorable surroundings, with proper temof food, from 20 to 40 minutes may be reckoned as a generation. In 24 hours, with the divisions once each hour, the progeny of one germ will be 16,777,216; with
perature and abundance
divisions each 30 minutes
If cell division
it will
be (16,777,216)^.
be in one direction only and the resulting daughIf cell ter cells remain undisturbed, a thread-like row results. cells adhere in groups, division be in two planes, and the resulting If the division be tablets of 8, 16, and 64 will occur frequently.
in three planes
The
and their cells adhere, packets result. structure of the bacterium cell owing to its minuteness
The most enduring portion of is yet very incompletely known. the vegetative cell is the cell wall. This is surrounded by a layer, the capsule and bears the flagella. The number of the flagella
and
their position varies in different species.
Some
none, some one, two, or many.
They may be
species have at the ends, polar,
THE
FUx\GI
WHICH CAUSE PLANT DISEASE

surface,
15
or scattered over
the whole
diffuse
or
peritrichiate.
is
They
are the organs of locomotion.
Within the wall
the pro-
toplast consisting of a peripheral layer, inner strands, imbedded granules and vacuoles bearing cell sap. The existence of a nucleus
comparable to that in higher plants is a much controverted point. Spores; Typically a bacterial spore consists of a highly refractive, ovoid, walled body within the mother cell. This body possesses
high resistance to ordinary stains, a great tenacity against decolorizing if it be once
stained, a higher resist-
ance against adverse temperatures and adverse

conditions generally than
<EIII
OiM3^^KDO^
do vegetative
cells,
and
SEZHi^
Fin. 7. Spores of bacteria showing their positiou within the cells. After Frost & McCampbell.
finally the ability to germinate and thus aid in
perpetuating the species. While the absolute number of bacterial

that form spores
species
is
large,
comparatively they are few.
They
are
most frequently met among the rod forms, and are rare among the spirilla and cocci. None are known among the important
plant pathogens. In the simplest cases of spore formation, the protoplasm becomes more dense in some part of the mother cell, the remaining
protoplasm of the
cell is
drawn around the denser mass, and the

whole resulting dense region becomes
enclosed within a special wall.
in this process nearly all the
Usually protoplasm
is
of the
mother
cell,
the sporangium,
cells
used.
The mother
formation
Fig.
8.
Spore
may
during spore remain of the normal
bacteria.
formation in After Fischer.
vegetative size and shape; they may take on (B. subtilis) or abandon (B.
megatherium) the habit of thread formation. Bacteria of many species become swollen at the point where the spore develops, Figs. 7 and 9; be this in one end (Vibrio rugula) or in the middle (B. inflatus). The swelling at the end is very common, giving rise
16

and
characteristic
to the peculiar
form known as "Nail Head" or

all
"Drum
Stick" bacteria.
In nearly
species of the Eubacteria
the spores are solitary. There are three modes of spore germination.
The most com-
mon, polar germination, consists in a rupture of one pole of the spore and the development of a normal vegetative cell through
the opening. The second mode, equatorial, Fig. 9, consists in a rupture in the side instead of the end of the spore. The third mode,
'If 4
cells,
FiQ.
9.
Spores of bacteria, showing bispored

germination.
After Prazmowski,
De Bary and Koch.
spore formation and spore
absorption, consists in a direct development of the whole spore into a vegetative cell. In suitable environment germination may
occur immediately after spore formation; able it may be delayed for many years.
if
conditions be unsuit-
Under certain conditions most bacteria undergo abnormal changes in form becoming elongated, branched, swollen, bulged,
curved, or variously,
termed involution forms.
Such are usually irregularly, distorted. They are in most cases due to unfavor-
able conditions of temperature and nutriment, and the bacteria resume their normal form when again in normal environment. The branched forms found in root tubercles after the period of luxuriant growth has passed, and the branched thread-like growth of the bacterium of human tuberculosis upon artificial media, are
by many regarded
as involution forms.
17
Constancy of Species. Bacteria in nature and under artificial conditions remain true to species. There may be variation from
generation to generation as among all other plants or animals of the world, and by the slow process of evolution, a species may in
many
generations become modified, leading eventually to
new
races, varieties,
and possibly
species.
That one
directly and suddenly to another, much into a species of another genus, is not to be credited. Marked variation is brought about in many species by change in tempera-
species can change less a species of one genus
changes in size, form, sporulation, flagellation, virulence, chromogenesis, fermentative power, grouping, etc. These changes belong to the life cycle of the species and
ture, food,
oxygen supply,
etc.,
occur as reactions to the environment.

Bacteria were discovered by Loewenhoek in 1683. That they do not originate spontaneously was shown by Pasteur in 1860-4. The first disease producing bacteria were recognized in anthrax by Pollander & Davaine in 1849; and the first definite proof that bacteria actually cause animal disease was made by Koch Avith
anthrax in 1875-1878.
cribed to bacteria
The
first
was the pear
plant disease to be definitely asblight by Burrill in 1879. The
invention of the cotton plug, Schroeder & Dusch, 1853, the gelamethod of plating for the isolation of species, Koch, 1881, and the use of stains, Weigert, 1875, were practically necessary
tine
prerequisites to
any considerable advance
in bacteriology.
For
long it was contended, especially by European bacteriologists, that bacteria do not cause plant diseases but most convincing
proof to the contrary was adduced by E. F. Smith. Entrance to the host plant is made in various ways, very often
through wounds, particularly wounds caused by insects, through roots, stomata, water pores, through delicate tissues as blossoms,
etc.
Once
in the tissue, bacteria
may
of the vessels, intercellular spaces or dissolved by the aid of enzymes.
migrate rapidly by means more slowly through cavities
In all there are some thirty-six well recognized Classification. genera embracing twelve hundred or thirteen hundred purported This number \vill doubtless be greatly despecies of bacteria.
creased
when the organisms have been
well studied,
that
many
so-called species are not really distinct.
by finding The number
18

many
additions of forms not as yet
will also of course receive
known.
No
system of
classification
can yet be said to have general
now in vogue will undoubtedly undergo minor changes and perhaps changes in fundamental
acceptance and
conception.
all classifications
The system of Migula" meets probably with most favor. With the omission of genera of little import pathologically, and
with the introduction of the order Myxobacteriales,
it is
as follows
SCHIZOMYCETES
three directions of space.
tion.
(p.
3)
Fission plants, without phycochrome, dividing in one,
two
or
Reproduction by vegetative multiplicaResting stages, endospores, exist in many species. Motility

flagella in
by means of
many
genera.
Key to Orders,
rein
Families,
and Genera of Schizomycetes

Order
I.
Cells without sulphur or bacterio-purpu-
Eubacteriales.
Coccaceae, p. 21.
Cells in free condition gobular; in di-
vision
somewhat
elliptical
I.
Nonflagellate Division in only one direction, cells single, in pairs, or chains
1.
Streptococcus.
Division in two directions;
cells
2.
may remain
in plates
cells
Micrococcus,
p. 21.
Division in three directions
may
Flagellate
remain
in
bale-like
3.
packets
Division in two directions
Sarcina.
4.
.
Planococcus.
Planosarcina.
Division in three directions.

Cells
5.
long
tion
or
short,
cylindrical,
straight, division in
one direcII.
6.
Bacteriaceae, p. 21.
Nonflagellate
Flagellate Flagella diffuse Flagella polar
Bacterium,
p. 21.
7. 8.
Bacillus, p. 37.
Pseudomonas,
p. 22.

Cells spirally
19
curved or representing part of a spiral, division in one direction

.
.
III.
Spirillaceae.
Cells cylindric in sheathed threads.
Cells with sulphur Motile rods in pseudoplasmodial masses

in a gelatinous matrix,
IV. Chlamydobacteriaceae. Order II. Thiobacteriales.
and forming
Order
III.
highly developed cysts
Myxobacteriales.
The
twenty-five genera in
species of families 3 and 4 and of orders II and III, some all, are so far as is known, unimportant as
regards plant disease. All of the known plant pathogens belong to one or other of the first two families of the Eubacteriales. Each
of these families contains several
dangerous parasites upon aniBacillus typhosus, Spirillum cholerae-asiaticse. Bacterium tuberculosis.

mals,
e.
g.,
The
specific
characters of bacteria are
chiefly
chemical or
physiological and rest in the relation of the forms to oxygen, gelatine liquefaction, fermentation of various sugars, acid production,
relation to nitrogenous compounds, chromogenesis, etc. To enable brief expression of these characters the Society of
'
'
^^^
American Bacteriologists endorses the following numerical

tem.*
sys-
A Numerical
100.
System of Recording the Salient Characters of an

Organism.
(Group Number)
200.
10.
Endospores produced Endospores not produced Aerobic (Strict)

Facultative anaerobic
20. 30.
1.
Anaerobic
(Strict)
2.
Gelatine liquefied Gelatine not liquefied
0.1
0.2
0.3
.0.4
*
Acid and gas from dextrose Acid without gas from dextrose No acid from dextrose No growth with dextrose
This
will
be found useful as a quick method of showing close relationships

is
inside the genus, but
not a sufficient characterization of any organism.
20

01

The
to the
21
plant pathogens as yet known, with few exceptions, belong two genera Pseudomonas and Bacillus between which they
are about equally divided. In the earlier days of bacteriology and to
some extent
in recent
days, bacteria have been seen in diseased plant tissues and have been placed by their observers in one genus or another and cited
as the causes of the diseases in question but without actual evidence that they cause the diseases and very often without any real evidence as to the genus to which the bacteria belonged. It is
of course usually impossible to identify such forms be dropped from consideration.
and they must
Coccaceae
(p.
18)
No
been
representative of this family parasitic upon plants has yet Micrococcus tritici Pril -* reliably recorded in America.
upon wheat in England is probably in reality Bacillus prodigiosus and not pathogenic. Micrococcus phytophthorus Frank "'. -^ reported as a cause of potato rot and also associated with potato
black-leg is perhaps in reality identical with Bacillus phytophthorus Appel. Micrococcus nuclei Roze, M. imperiatoris Roze, M. flavidus Roze, M. albidus Roze, M. delacourianus Roze and
^^ as the causes of vapellucidus Roze are assigned by Roze rious potato troubles in Europe, and M. populi Del.-^ is said to be
M.
the cause of canker on Populus.
Bacteriaceae
(p. 18)
Bacterium Ehrenberg
(p.
18)
their lack of
These non-motile forms, perhaps owing to
power
of locomotion, are comparatively rare as plant pathogens. Bact. briosianum Pav. is given as the cause of rotting of tomato fruit and distortion of vegetative parts in Italy. ^^ It is
described also on Vanilla.^^

Bact. montemartinii Pav.
of Wisteria.
1^-
is
described as the cause of a canker
Bact. mori B.
&
L.
is
said to cause leaf
and branch spots on
mulberry.
^^^
22

Bact. teutlium Metcalf.^^
(Group number 222. 220 .)

/i,
A
3 X
short rod with rounded ends, 1.5 x 0.8

1
before division
m; non-motile,
no
flagella seen;
no spores; Gram-positive;
agar colonies round, thin, not viscid, porcelaneous to transparent, seldom over 0.5 (x. No liquefaction. Broth clouded, precipitate
thin or none, no pelUcle. Milk not coagulated. T. D. P. 45, 10 min. Opt. 17. Aerobic, no gas. Beets diseased by this organism were honeycombed with pockets
filled
with a viscous
tissue
fiuid,
The vascular
was not
a practically pure bacterial culture. rotted. Inoculation by pricking the
bacterial exudate into healthy beets resulted in typical disease. Pure cultures isolated by use of cane-sugar-agar gave similar results.
Three weeks
after inoculation the exudate-forming pockets
were
is
typically developed.
Surface inoculation failed and there
no
evidence that the organism can infect except through wounds. No rotting followed inoculation on potato, white turnip, radish,
tomato, or apple.
Bact. pini Vuill.^ as their cause.
was found
in tissue of pine galls
and regarded
fici Cav.^^'* is reported as the cause of a disease of figs. Bact. scabigenum Busse & v. Faber is described as the cause of scab of sugar-beets in Germany.^^
Bact.
Pseudomonas Migula
(p. 18)
Short or long rods motile by polar flagella, fig. 10, whose number varies from one to ten but is most commonly one. Endospores are sometimes present. The cells in some species adhere to form short chains. The basis of separation into species is the
growth upon gelatine, character of the colonies, chromogenesis and numerous other cultural characters.^-' ^^' ^^
of
Something over seventy-nine species are known, at least fifteen which cause diseases in plants, some of them very serious. Many other species occur in water, soil and manure, while others
are suspected animal pathogens.
One prominent group of plant pathogens, ^^ the yellow Pseudomonas group, contains, according to Smith, Ps. campestris, Ps.
phaseoli, Ps. hyacinthi, Ps. stewarti, Ps. juglandis, Ps. vascularum, Ps. dianthi, Ps. amaranti, Ps. malvacearum. These, he says,

agree in the following particulars:
23
They are yellow rod-shaped organisms of medium size, straight or slightly crooked with rounded ends. The segments multiply by fission, after elongation. They are generally less than 1 yu in diameter. The segments
occur singly, in pairs or in fours joined end to end, or in clumpy masses of variable size (zoogloese), more rarely they are united
into long chains or into filaments in which no septa are visible. Endospores have not been observed. The segments are motile by
means
of one polar flagellum which is generally several times as as the rod, and may be wavy or straight when stained. The long species grow readily on all of the ordinary culture media, but so
far as
is
definitely
known
all
are strictly aerobic.
None
are gas
The yellow producers. They do not reduce nitrates to nitrites. color appears to be a lipochrome and in the different species varies from deep orange and buff-yellow, through pure chrome and canary-yellow, to primrose-yellow and paler tints. Ps. aeruginosus Del. possibly identical with Ps. flourescensputridus FlUgge is the cause of a leaf and stem disease of tobacco
in France."*^
Ps. avenae Manns, (Group

\vith
fj,
number
jj..
111.2223032.)
short rod
x 1 to 2 round ends, 0.5 to 1 Actively motile, generally by one polar fiagellum, occasionally by two or three. Gram What seem to be endonegative. spores are found in old cultures. On
agar stroke, growth very slow, filiform, rather flat, glistening; margin
smooth, opaque to opalescent; nonchromogenic. Liquefaction occurs on gelatine in seven to twelve days.
Agar coloround with surface nies, amorphous, No gas in smooth, edges entire.
is
Broth
slowly clouded.
dextrose, saccharose, lactose, maltose, or glycerine. Ammonia and intlol
Fig.
lO- Ps-
avenae.
After
not formed.
Nitrates reduced to nitrites.
T. D. P.
10 min.,
in 1909,^^
60, Opt. 20 to 30.
This organism was isolated and described by

as the cause of a serious oat blight.
Manns
Inoculations with
it
alone by
24
hypodermic injection produced only limited lesions but similar inoculations with a mixed culture of Ps. avense and Bacillus avenae
Manns, moreover, noticed that the produced typical disease. virulence of the Pseudomonas decreased when kept in culture free
from the Bacillus, also that in the disease as it occurs in nature His conclusion is that the these two organisms are associated. Pseudomonas is the active parasite and that the Bacillus is an important, perhaps a necessary symbiont.
Fig. 11.
1
Showing
2, six
effert of inoculation of Ps. campestris into
cabbage plants.
and
weeks after inoculation.
No.
3,
check plant uninoculated.
Nos. After
Russell.
Infection in nature is chiefly stomatal by spattering rain. Soaking of seed in suspensions of bacteria did not produce the disease. Inoculations on wheat failed, though from one variety of blighted wheat. Extra Square Head, the typical organism was isolated. Inoculations on corn made during wet weather produced lesions which spread rapidly and the organism was re-isolated. Barley is said by Manns to be susceptible and what he believes to be the same disease occurs on blue grass and timothy.
Ps. campestris (Pam.) E. F. Sm.
(Group number 211.333151.)

/x;
rod-shaped, motile, organism generally 0.7 to 3.0 x 0.4 to 0.5

pale.
color dull waxy-yellow to canary-yellow, occasionally brighter or
more
One
polar flagellum; no spores known.
Aerobic but
25
not a gas or acid producer, gelatine liquefied. Cavities are formed around the bundles but the organism seems to be only feebly A brown pigment is produced in the destructive to cellulose. Growth host plants and on steamed cruciferous substrata. rapid on steamed potato cylinders at room temperatures, without Growth feeble at 7, rapid at 17 to 19, odor or brown pigment. luxuriant at 21 to 2G, very feeble at 37 to 38 and ceases at 40. T. D. P., 10 min., 51.
Fig. 12. Ps. campestris. Soction of a cabbage loaf parallel to the surface and near the margin, showing the result of infection through the water-pores. After
Smith.
It is closely related to Ps. hyacinthi
from which
it
differs chiefly
its
in its pathogenic properties, its duller yellow color
and
higher
thermal death point.
troublesome upon cabbage, turnips, cauliflowers, coUards; and a very large number of cruciferous hosts, both cultivated and wild are susceptible. It enters the host
It is
plant through the vascular system which becomes decidedly brown. This organism was first isolated by Pammel ^ (see also ^^) from rutabagas and yellow turnips in 1892; grcen-hou.se inoculations with pure cultures were made in scalpel wounds, which were then
26
sealed with wax.
The plants showed rot in a few days and the actual causal relation of the organism was thus established. Con^'^ from puncture inoculafirmatory evidence was gained by Russell
tions
in
cabbage and
It
cauliflower petioles. was further shown
by
E. F. Smith
^^
that the
cabbage and turnip organisms are identical
and that the
bacteria,
by solution
lose,
of the cellu-
produce pits and
holes through the walls of the host cells resulting
eventually
in
large cavities.
Infection was shown ^^ and by by Russell 3' 39 Smith to be chiefly
through the water pores or wounds through
made by
bacteria
insect
insects;
the
being air or borne and de-
rived largely from infected soil. After entering the plant the bacteria multiply rapFig. 13.
Ps. campestris; cross-section

root.
idly,
of a turnip
and
migrate
in
every
direction
of the veins.
by
After Smith.
means
Studies of Harding, Stewart and Prucha ^ (see also) ^- showed that it can survive the winter on the seed and thus infect seedlings. Ps. destructans Potter '^^ is described as an uniflagellate organism
causing a destructive soft rot of turnips and beets in England. Doubt has been thrown upon its identity by the work of Harding
and Morse
^^
and
of Jones
^^
who found supposedly
authentic
cultures to bear peretrichiate rather than polar flagella.
See
p. 42,

Ps. dianthi Arth.
27
&
Boll.''
Though
originally reported as the

is
probable cause of carnation leaf spot, this organism as a saprophyte.
now regarded
medium
Ps. fluorescens (Flugge) Mig. Straight and curved rods of size in chains of two or several members. Cells 0.68 x
fi. Spores not seen. Flagella 3-6 polar. Gelatine liquefied; surrounding medium colored greenish-yollow; Gram negative. Milk not coagulated. Indol weak. Bouillon,
1.17-1.86
turbid, fluorescent. This organism or
two
varieties of
sponsible for a decay of celery. numbers in the decayed tissue;
by Barlow held reThe organism was found in large was isolated and typical rot was
it
are
'^^
induced by inoculation of pure cultures upon

stems.
It is also credited
sterilized celery
with two distinct types of tobacco disease in one of them on seed, the other on the growing plant. France, *' has claimed that both of the varieties, Ps. Recently Griffon
fluorescens liquefaciens and Ps. putrida are capable of producing wet rot of various vegetables, carrots, rutabagas, tobacco, tomatoes, melons,
and that the
latter
organism
is
identical with Ps.
Eeruginosus.
It is also held that B. brassicsevorus
and B.
''"
cauli-
vorus are forms of Ps. fluorescens.

Ps. fluorescens exitiosus v. Hall
rot of
Iris.
is
said
by van Hall
to cause
Ps. hyacinthi (Wak.) E. F. Sm., is a serious pest of hyacinths in the Netherlands but has not yet been recorded in America.''^
is medium sized rod with rounded ends, measuring in the host 0.8-1.2 x 0.4-0.6 n; actively motile by one long polar flagellum;
It
sporiferous; liquefies gelatine slowly; aerobic; no gas. It produces indol. Does not Fio. i4.Ps. hyacinthi. ^^^""^ ^"'*^grow at 37. Opt. 28 to 30, T. D. P. 10
non
min., 47.5.
It is
wound
forming a bright yellow slime and

pestris
parasite which grows in the vessels is closely related to Ps. cam-
and
Ps. phaseoli.
Ps.
iridis V.
Hall
is
described
by van Hall
as the cause of
^^
decay
of shoots
and rhizomes
of Iris.
Ps. juglandis Pierce.
(Group number -11.
51-.)
rod
28
1-2 X 0.5

n,
flagellum.
with rounded ends, actively motile by one long polar Bright chrome-yellow in growth; disastatic ferment
No gas; aerobic. It was isolated from diseased nuts, and twigs of English walnut in California in 1901. Inleaves, The oculations by spraying demonstrated its pathogenicity.
present.
organism is closely related to Ps. campestris but is distinguished from it by the abundant bright yellow pigment produced upon the surface of extracts of leaves of walnut, magnolia, fig, castor bean and loquat.
Ps. leguminiperdus (v. Oven.) Stev.,^^ said to be distinct from Ps. phaseoli, occurs on peas and other legumes. It was isolated, cultivated and inoculations made. Ps. levistici Osterw.^^ occurs on Levisticum. Ps. maculicolum (McC.) Stev. (Group number 211.3332023.) A short rod, forming long chains in some media. Ends rounded. Size from leaf 1.5 to 2.4 fi by 0.8 to 0.9 fj.; in 24-hour beef-agar culture, 1.5 to 3 Ai by 0.9 n. No spores, actively motile, one to
five polar flagella
tile
two to three times the length

media.
of the rod.
Mo-
in
most
artificial
Involution forms in alkaline beef
Pseudo-zooglcese in Uschinsky's solution. Gram negaStains readily with carbol fuchsin and with an alcoholic solution of gentian violet.
bouillon.
tive.
Agar plate colonies visible on the second day as tiny white specks, in three to four days, 1 to 3 mm. in diameter, white,
round, smooth,
crinkled,
flat,
shining,
and translucent, edges
entire,
with
age dull to dirty white, slightly irregular, edges undulate, slightly
and with
indistinct radiating marginal lines.
Buried
colonies small, lens-shaped. Agar streak cultures white, margins slightly undulate. Beef bouillon clouds in twenty-four hours. Growth best at surface
where a white
layer, not a true pellicle, is formed.
No
zoogloeae.
No
rim.
Gelatine stab cultures liquefied in eight to ten days.
Growth
Slight
from surface
craterif orm slight, white, granular precipitate.

;
green fluorescence.
production feeble.
separation into curd and whey. T. D. P. 46. Opt. 24-25. Max. 29.
it
No
Indol
Min.
below
0.
Isolated from cauliflower leaves on which
forms brownish to
THE
FUiXGI

mm.
in diameter.^^
29
purplish-gray spots 1-3

this host also
is
on cabbage was proved by inoculation.

Sm.^'*'
^^' '^
Pathogenicity on Its entrance
stomatal.
This yellow organism, resembles Ps. campestris but its slime is more translucent on potato and it does not attack the cabbage. It was
pathogenic on cotton,
Ps. malvacearum, E. F.
much
by Smith and made by spraying a suspension of a young agar culture of the organism upon cotton leaves and bolls. No description has
grown
in pure culture
successful inoculations were
been published.
number 212.3332133.)
1.2-2.4 X 0.5-0.8 m;
Ps. medicaginis Sackett.^^ (Group A short rod,

filaments 20.2spores;
long.
actively motile with 1-4 bipolar flagella; capsules and zoogloea none. Agar streak
/i
37.2
No
filiform,
later
echinulate,
ghstening,
smooth, translucent, grayish-white; no gelatine liquefaction; bouillon

slightly turbid, pellicle
sediment
scant.
on third day, Milk unchanged.

Fig. 15.
Agar
colonies
round,
No gas or indol. + 15 to +18 Fuller's
gra5dsh-white. Optimum reaction

scale.
Ps. malvacearum.
Early stage of stomatal infection in angular leaf-spot of cotton. After Smith.
T. D. P.
.
No diastase, invertase, 49-50, 10 min. Opt. 28-30. Aerobic zymase, rennit or pepsin. It occurs as a pathogen on alfalfa and issues in clouds visible to
the naked eye from small pieces of tissue of the diseased stem or leaf when mounted in water on the sHde. These clouds under the
thick.
high power resolve into actively motile rods, relatively short and The bacteria are also found in practically pure culture in
the exudate which oozes from the diseased tissue as a clear viscous
and collects in drops or spreads over the stem. Sackett with pure culture inoculations produced the typical disease and re-isolated the organism with unchanged characters. Re-inoculated
liquid
30
it

again caused disease.
inoculations by hundred per cent infection. scarification or puncture gave one Controls remained undiseased. Infection, stomatal or water pore, was also secured through the apparently unbroken epidermis.
More than a hundred
The
virulence of the or-
ganism was retained after It five months on agar.

is
believed that the usual

of
mode
infection
is
through rifts in the epidermis due to frost and that the germ is wind-borne
from infected
Ps.
F.
soil.
Sm.)
michiganense (E. Stev. (Group
number
Rod
Fig.
-22. 252-.) short with rounded

/x-
ends, 0.35-0.4 x 0.8-1.0
16. Ps. medicagini.s; 48-hour agar-culture, showing formation of filaments. After Sackett.
motility Seen from gêms. Flagclla apparently
No
round.
polar but not seen distinctly. Agar colonies pale-yellow, smooth, Agar stab canary-yellow, opaque, viscid. Bouillon moderately clouded, a moderate slimy precipitate; no rim or pellicle. Gelatine not liquefied.
The organism was
described
by Smith
^^
as the cause of a
stem
disease of tomatoes in Michigan. No fungi were seen but bacteria were present in great numbers in the bundles also in cavities in
the pith and bark.
The organism was isolated and the disease was both by pure culture inoculations and by crude inoculaproduced tions, using an impure inoculum. The disease caused is less rapid in development than that caused by B. solanacearum and less
browning of the infected tissue occurs. Rod Ps. mori (B. & L.) Stev. (Group number 222. 202-.) with rounded ends, 1.8-4.5 x 0.9-1.3 fx, mostly 3.6 x 1.-2 fx; motile by one, sometimes two polar flagella. No spores. PseudoAgar Agar colonies round, smooth, flat. zooglcese present. Gelatine stab filiform, no streaks spreading, flat, dull-white.
liquefaction.
Milk not coagulated.
No
gas.
T. D. P. 51.5.

"^
31
In 1894 Boyer and Lambert produced successful inoculations on mulberries with an organism to which they gave the above
name, but without description.

^^ In 1908 E. F. Smith, plated out, from blighted mulberry
leaves
in
collected
a Georgia, white species
with
which
he
made numerous
infections on
leaves
both stems and of mul-
berry.
From
cultures
these
Smith
the
supplied
in
description
quoted
above.
1
part
re-
The
which Bacillus cubonianus ^^ has to

at
i
on
this mulberry
disease
is
un-
Fig.
17.
Ps.
A
aud surface colonies by
medicaginis; agar colonics 7 days old, deep reflected light. After Sackett.
known.
Ps. phaseoli E. F. Sm.
short round-ended rod, wax-yellow
Milk coagulates, and the whey slowly separates without acidity; gelatine liquefies slowly. Growth feeble at 37, none at 40. T. D. P. 10 min.,
to chrome; motile; anaerobic.
49.5.
A starch enzyme is produced and the middle lamella also dissolved. This organism is pathogenic to beans and
some
related legumes
and
it
is
closely related to
The bean was noted and asAfter Smith. cribed to bacteria by Beach ^^ and ]:)y Hal^ sted in 1892, and the organism was described by E. F. Smith in 1897 ^^ after it had been grown in pure culture and successful inoculations had been made.
Fig. 18.
Ps.
Ps. hyacinthi
and
Pg. campestris.
phaseoli
disease, occasioned by
32

Ps. pruni, E. F. Sm.
The organism resembles

it
Ps. campestris
growth on potato and which it converts into a by its behavior in Uschnisky's solution It consists of small rods, motile by one to several viscid fluid. Gelatine not liquefied. Casein T. D. P. 51. polar flagella.
but
is
distinguished from
by
its
feebler
slowly precipitated and later redissolved. No gas. ^^ The bacteria enter through the stomata of the Japanese plum; cause small watery spots on green fruit and leaves, and finally the
Fig. 19. pruni.
Earliest stage of
The
fruit spot on green plums, due to Ps. bacteria entered through the stoma. After Smith.
death of the affected tissue. In earliest disease they are limited to the substomatal space but gradually they invade more distant Wounds are not necessary to infection. It seems to have tissue. been seen first on the peach in 1903 by O'Gara in Georgia and in ^^ the same year by Clinton in Connecticut. Rorer by numerous
cultures
sponsible for
cross inoculations proved this same organism rea leaf, tvng and fruit disease of peaches. In the twig the bacteria were present in great numbers in the bast. Ps. radicicola (Bey.) Moore.^^ The legume root-tubercle or-
and
33
ganism, by some regarded as a parasite, though beneficial, and byothers regarded as a mutuaHstic symbiont will not be discussed
here owing to
its beneficial
character.
Fig. 20.
Part of sweet-corn stem parasitized by Ps. stewarti.
After Smith.
Ps. savastanoi (E. F. Sm.) Stev. A rod with rounded ends, nonsolitary or in short chains, 1.2-3 x 0.4-0.8 ju; motile; aerobic;
Standard agar, sporing; flagella 1-several, often 2-4, polar. surface colonies, white, small, circular, smooth 1.5-3 mm. at three days, edge entire; bouillon thinly clouded, precipitate slight, white, no rim or
peUicle.
On
gelatine
no liquefaction; colonies white,
round, erose, margin pale.
34

From
swellings known as olive tubercles on Californian olive ^ which is in part branches, E. F. Smith isolated this organism
Ps. oleae-tuberculosis
and which may bear
relation to several other
olive bacteria previously described in Europe.
The organism when
inoculated by puncture into young olive
Later it was reshoots produced the characteristic tubercle. isolated from these artificially produced tubercles and used in a
second series of inoculations which gave a second crop of tubercles. Controls showed no infection and healed promptly. The oleander
was not susceptible to
infection.
full
Smith's results are not in
accord with
much
of the
European
work on the olive tubercle. Ps. sesami Malk. causes disease on sesame ^^. Ps. stewarti, E. F. Sm. A medium size rod, 0.5 - 0.9 with rounded ends, and 1 polar flagellum. Buff-yellow
;u
-2
}i,
to
chrome
or ochre color; non-liquofying; does not separate casein in milk. T. D. P. 10 min., 53.
Agar
cular,
colonies
subcir-
becoming lobate;
bouillon rendered tur-
bid
with yellow-white
precipitate.
No
gas.
The
was
bacterial
corn
blight of this organism

first
described
Stewart in 1897 " and

attributed to bacteria.
scribed
in
by
The organism was deby E. F. Smith

1890
^s
ture furnished
Fig. 21.
from a culby Stew-
Various forms of Ps. stewarti, grown on potato agar; a and b are typical forms. After Stewart.
art.
Definite proof
of
by
the
inoculation
causal relation of this

particular
organism to the disease was adduced
in
1902
by
" Some plants showed sprinkling bacteria upon the leaves.^^' constitutional symptoms during the first month, most of typical
them
in
two or three months when the plants were
several feet

high.
35
tures of
In these plants the vessels become plugged with pure culPs. stewarti from tip to base. Small holes filled with
yellow slime appeared later in the parenchyma. Wounds were entirely unnecessary to infection, though the vessels are the primary
seat of disease.
Ps. syringae v. Hall tumefaciens Ps.
"^
causes disease of Syringa and other plants.
(S.
&
T.)
Stev.^^"^
(Group
number
212.2322023.) Vegetative cells taken directly from a gall usually 0.6 to 1.0 M X 1.2 to 1.5 MWhen grown on agar for two days 2.5 to 3 m x 0.7 to 0.8 m or
occasionally wider.
of one,
Endospores not observed. Motile by means sometimes two or three terminal fiagella; viscid on agar but capsules not demonstrated. Readily stained in ordinary basic
anilin stains;
up
negative. Agar surface colonies usually in from four to six daj^s at 25,
Gram
come
white, smooth, circular; margin even,
semi-transparent, maximum 2 to 4 mm. Agar streak; growth size

shining,
moderate,
filiform.
On
sterile
potato
rapid, in one cylinders or two days covering the entire surface

of
growth more
the cylinder;
grayish,
smooth wet-glistendarker
^vith
ing, slimy to viscid, odorless; potato
cylinder
age,
Fig.
never yellow.
white,
Gelatine colonies dense,

small,
22. Fiagella of Ps. tumefavarious stains.
circular,
medium not
stained.
non-liquefying, In beef broth
ciens,
After
Smith.
clouding often absent or inconspicuous, rim of gelatinous threads

present, also more or less of pelUcle; in young cultures very delicate suspended short filaments, best seen on shaking. Milk coagu-
whey begins only after several days; litmus milk gradually blued, then reduced. Cohn's solution, growi^h scanty or absent, medium non-fluorescent.
lation delayed; extrusion of
gas produced; organism aerobic in its tendencies; nitrates not reduced. Indol produced in small quantity, slowly. Slight
toleration for citric, malic, and acetic acids. Toleration for alkali slight. Optimum reaction between +12 and +24, Fuller's
scale.
No
T. D. P. 51.
Opt. between 25 and 28 Max. =^37. Growth
36
occurs at
0. Milk, bouillon, dextrose peptone water with calcium carbonate are the best media for long continued growth.
recommended as quick tests for differential appearance of colonies on +15 agar plates made from the tumors; young agar stroke cultures; behavior in milk and litmus milk; growth on potato; behavior in Cohn's
following are
The
purposes.
Time
of
solution; stringy ring
and suspended filaments
in peptonized beef
bouillon; inoculations into young, rapidly growing daisy shoots or into growing sugar-beet roots.
The organism is readily plated from young sound galls, i. e., those not fissured or decayed. In galls on the Paris daisy (Chrysanthemum frutescens) these
bacteria were found in small numbers. By plating they were obtained in pure culture and puncture inoculations repeatedly resulted in the characteristic gall. From these the organism was reisolated and the disease again produced, thus giving conclusive
evidence that the organism
is
the actual cause of the
gall.
Swell-
ings began four or five days after inoculation and in a month they were well developed though they continued to enlarge for several
months, reaching a size of 2-5 cm. in diameter. Tumor-producing Schizomycetes have also been isolated from over-growths on plants belonging to many widely separated Natural galls have been families (Compositae to Salicacese).
studied on Chrysanthemum, peach, apple, rose, quince, honeysuckle, Arbutus, cotton, poplar, chestnut, alfalfa, grape, hop, beet, The organisms from salsify, turnip, parsnip, lettuce, and willow.
these sources are closely alike on various culture media, and many of them are readily cross-inoculable, e. g., daisy to peach, radish,
grape,
sugar-beet,
hop; peach
sugar-beet, poplar;
to daisy.
hop to
to daisy, apple. Pelargonium, daisy, tomato, sugar-beet; grape to
almond, sugar-beet; poplar to cactus, oleander, sugar-beet; willow With eight of these organisms tumors have been produced on sound specimens of the species from which obtained.
Some
cross-inoculate
more
readily than others,
and there are
also
slight cultural differences.
Thus,
it
is
probable that there are
several races of the gall-forming organisms varying more or less in amount of virulence and in adaptability to various hosts. In
general
it is
said that
all
plants susceptible to crown galls,
i.
e.,
37
those on which the galls have been found in nature, are susceptible
Hard gall, hairy root, and soft gall due to infectious bacteria. As tentative hypotheses Smith assumes either: (1) That the hairy root organism while resembling the crown gall organism is not identical with it; or (2) That they are the same, and that if infection takes place in a certain group of cells an ordinary gall
to artificial cross inoculation.
are also
all
will develop, while

i.
if other special groups of cells arc first invaded, the root anlagc, then a cluster of the fleshy roots will develop. Some of his inoculation experiments point to the latter conclusion.
e.,
'^' Ps. vascularum (Cobb) E. F. Sm. "' cane, filling the bundles with a yellow slime. ported in America.
is
parasitic
It has not
on sugar been re-
Ps. sp. indet.
1-3 polar
flagella,
was
short rod, 2-4 x 1-1.5 p., actively motile by isolated from diseased spots on the larger
veins and petioles of beet leaves by Brown."^ The organism was successfully inoculated in pure culture, disease produced, and the organism reisolated. It is infective as well for lettuce, sweet
pepper, nasturtium, egg plant and bean. Agar colonics are creamy white, thin, circular, turning the surrounding agar j'cllow-grcen
in
bouillon
three days. Gelatine is Hquefied; litmus milk is clouded. Opt. 28.
turns blue;
Ps. sp. indet.

flagella
was
isolated
short rod, 2-4 n long, motile from diseased nasturtiums
by 1-3
polar
leaves
by
Jamiesson.^''
(Tropeolum) Pure culture inoculations induced typical
clouds bouillon; produces on agar small, round, bluish-white colonies; liquefies gelatine and does not proIt is pathogenic duce gas. Opt. 25. T. D. P. 49-50, 10 min.
disease.
The organism
also for sweet-pea, lettuce, pepper, sugar-beet
and bean.
Bacillus Cohn. (p. 18.)
This genus not polar, flagella.
differs
from Pseudomonas only in its Endospores are often present.
peritrichiate,
Of the four
to to
more species nineteen at least are known be plant pathogens. Numerous animal pathogens also belong
hundred and
fifty or
this genus, notably B. typhosus, B. pes'as.
B. ampelopsorae Trev. is said to cause grape galls in Europe, but the evidence is by no means conclusive. Cf. B. uvae.
38

B. amylovorus (Burr.)
Bacillus
in
De
Toni.
(Group number 221.

ix,
.)
broth,
0.9-1.5
x 0.7-1.0
flagella
longer
when
older.
Gram
positive;
no capsule;
several;
no spores; broth
clouded, pellicle slight.
Gelatine shows slow, crateriform liquefaction. Agar, buried colonies white, surface
colonies elevated, circular wet-shining, margin Milk coagulated in three-fourths of
irregular.
a day, later digested to a pasty condition. Opt. 25-30. T. D. P. 43.7, 10 min. Facultative anaerobe.
Indol produced; no gas; no pig-
ment.
Fig. 23.
B. amylomultiplj'-
Burrill in 1877.'-'
Bacteria were noted in blighted pear twigs by ^^ In 1880 he ^* demonstrated
vo rus,
ing by fission. After Whctzel and Stewart.
the communicability of the disease by introducing the bacterial exudate into healthy pear
trees as well as into
This constitutes the

tributed to bacteria.
in 1884
^^
apple and quince trees. plant disease definitely atBurrill's results were confirmed by Arthur
first
case of
by one hundred and twenty-one puncture
inoculations, using the exudate, also a bacterial suspension from diseased twigs. He further demon-
strated the susceptibility of Juneberry and hawthorn. Usually the disease appeared about a
week
after inoculation.
Attempted raspberry and
grape inoculations failed. Arthur placed the whole matter on a firm foundation by passing the bacteria through a long series of artificial cultures and then by inoculations,
showing that they were capable of causing the blight.^^' ^^ He further demonstrated that the bacterial exudate from the tree, when freed of
bacteria
24. Claw Fig from bee's foot with blight on bacteria and about it
showing the
relative
zel art.
size.
The
could not produce disease. results of an extensive study of the bacteria on
by
filtration,
After Whetand Stew-
media; of their morphology and stain reactions were published by Arthur in 1886.^^ Bacteria were shown to penetrate twigs 3-4 dm. beyond their area of visible
various
effect.207
In 1902 Jones
^^
isolated
an organism from blighted plum
trees.

This he demonstrated
bj'
39
culture
and
cross inoculation in fruits to
be identical with the pear blight organism, though inoculations in plum twigs did not give disease, presumably due to the high resistance of this plant. Similarly Paddock has shown this organism to attack the apricot.^ Detmers has reported what she regarded
as this blight caused by this Bacillus on blackberries.-'^ Other hosts are hawthorn, shad bush, mountain ash.
By
inoculations with pure cultures of the apple body-blight
bacteria, blight upon twigs and blossoms was in 1906,^^ thus proving the identity of these
produced by Whetzel two forms of disease,
an identity assorted
first
by
Burrill.^^
Fig.
25.^B.
aroideis.
After Townsend.
B. apii (Brizi.) Mig.^^
which
by
is reported as the cause of a celery rot, possibly identical with a bacterial rot reported earlier Halsted.^^ is
B. araliavorous Uyeda, described on Ginseng in Korea is perhaps also the cause of soft rot of Ginseng in America.^ The organism was isolated and studied by Uyeda who made inoculations.
Pseudomonas
aralise
and Bacillus koraiensis were

3=*
also
com-
monly
present in the Oriental disease.^^
B. aroideae
Town.
(Group Number 221.2223022.)
This organism was described in 1904 as the cause of soft rot of ^^ calla corms and leaves. The bacteria were present in almost
pure culture in affected tissue and by puncture inoculation in pure culture produced the typical disease in a few days. Townsend regarded the organism as distinct from B. carotovorus, B. oleraceae, B.
hyacinthi septicus and Pseudomonas de-
40

Harding and Morse, however, believe
See
p. 42.
it
sttuctans.
specifically
identical with B. carotovorus.
B. atrosepticus v. Hall,^^ was isolated from ducts of potatoes

affected with black leg.
B. avenae, Manns.^^
avense.
This
is
the symbiont of Pseudomonas
See
p. 23.
short,
(Group number 222.2223532.) A very actively motile bacillus, rod-shaped with rounded ends, 0.75 to 1 x 1.5 to 2 ;u.
Fig. 26.
Plate culture of B. avenae, on nutrient glucose agar,

four days at 30''C.
After Manns.
negative; endospores not observed; flagella many, diffuse, long, undulate; growth on agar stroke rapid, filiform, white, glistening, later somewhat dull, margin smooth, growth rather
Gram
opaque, turning yellow third day; gelatine not liquefied; broth clouded and on the second day showing heavy yellow precipitate; milk coagulated at end of two weeks with extrusion of whey;
agar colonies round, entire, surface smooth, slightly raised. No gas in dextrose, saccharose, lactose, maltose, or glycerins. Indol production moderate; nitrates reduced to nitrites. T. D. P. 10
min., 60; Opt. 20-30.
41
B. betae Mig. is reported as the cause of gummosis of beet.^ B. brassicaevorus Del. isolated from diseased cabbage ^"^ is per-
haps identical with Pseudomonas fluorescens. See page 27. B. carotovorus Jones. (Group number 221.1113022.) From
agar 1-2 days old as
short or long rods, in short or long chains.
0.7-1.0 X 1.5-5
jjL,
com-
monly
flagella
0.8
x2
No
n; ends
rounded.
spore;
peritri-
2-10,
chiate;
no
capsule;
Gram negative. White

on
all
media.
Agar
opaque
In gela-
slope filiform to spreading, glistening,
to opalescent.
tine stab; liquefaction
crateriform
dibuliform.
to infun-
Broth
thin
clouded,
to
peUicle
sediment flocculent; milk coaguabsent,

lated.
Agar
colonies,
round, smooth, entire

to
undulate,
or
amorFig. 27.
phous
lactose
granular.
in dextrose,
B.
carotovuru.s wedging apart cells of

carrot.
tlie
Some gas
After Smith.
and saccharose, nitrates reduced
to nitrites; indol feeble.
Opt. 25-30. A considerable number of cultivated plants suffer soft rot from the attacks of a non-chromogenic liquefying bacillus. Among the
plants so affected are cabbage, turnips and other crucifers; parsnip,
potato, celery, tomato, Jerusalem rhubarb, onion and iris. artichoke, asparagus, In 1901, Jones reported an organism isolated from rotting carrots
carrot,
T. D. P. 48-50.
mangel,
sugar-beet,
It disorganized tissue which he named B. carotovorus.'''-' '^ by solution of the middle lamella; and infection into wounds led
42
to decay of roots of carrot, parsnip, turnip, radish, salsify, of onion bulbs, hyacinth corms, cabbage heads, celery stalks and
fruits of
tomato, pepper and egg plant. Jones found no decay produced in young carrot or parsnip plants, fruits of orange, banana,
apple, pear, cauliflower head,* Irish potato tuber, beet root or tomato stems. ^^ Infection
^\
^__^
iv\
did
not occur
unless
the
epidermis was broken. The rotten mass was always soft, wet, and exuded a
liquid
teria.
clouded
^^
with
bac-
Jones
FiG. 28.
in 1909
n
made an
B. carotovorus.
After Jones.
extensive study of the cytolitic
enzyme
oi
this germ.
.i-
This enzyme was separated by heat, filtration, formalin, phenol, thymol, chloroform, diffusion, alcohol, and its conditions of production and action investigated. Heating the enzyme to 60 inhibited its activity to a marked degree; higher than 63 inhibited it entirely; chloroform, thymol and phenol did not retard its action.
was suffered through alcoholic precipitation and dried enzyme remained active for fully two Its effect was greatest at 42, less at 32 and 48. No years. diastatic action was observable. In 1909 Harding and Morse, ^^ from an extended study of some
loss
No
resolution.
The
of
liquefying soft-rot bacilli forty-three pathogenic strains (including B. carotovorus, B. olera;ce8e, B. omnivorus, B. aroidese and what Potter regarded as Pseudomonas destructans) from six different vegetables, con-
12,000 cultures of non-chromogenic,
some
clude that unless later studies of the pathogenicity of these cultures shall offer a basis for subdividing them, there is no apparent
reason
why they
should not
all
be considered as somewhat variant
members
of a single botanical species.
This conception would lead to the abandonment of the supposed

species mentioned above and the recognition of all oi them under their oldest described form, B. carotovorus Jones, which in our
*
Harding and Stewart
later
showed that
it
is
capable of rotting cauli-
flower.
43
present knowledge seems certainly to be the most wide spread, common and destructive of the soft rot bacteria. Some, perhaps
much,
of the rot of crucifers generally

is
monas campestris
Harding
thought to be due to Pseudocaused by B. carotovorus. See probably
&
Morse.^^
Pril. & D(>1. has been reported as the cause of also as a parasite on a large number spots on grapes under glass, of other plants among them Pelargonium, potato, begonia, clemIt is later stated that this is probably really a variety of atis.
B. caulivorus,
Ps. putrifaciens liquefaciens.
B. cepivorus Del. (possibly a Bacterium)

bulbs.si
is
recorded on onion
B. coli (Esch.) Mig. or an organism indistinguishable from
it is
held by Johnston
rot.
capable of causing rot of soft tissues of the cocoanut plant and is perhaps responsible for cocoanut bud
B. cubonianus Mace, was originally described as the cause of mulberry disease (cf. Ps. mori). This organism, or at least one that was regarded as indistinguishable from it, has been mentioned as the cause of a disease of hemp.^^ B. cypripedii Hori is a medium sized slender, non-sporulating form with four flagella.^^^
B. delphini E. F.
^^^
Sm.
This
is
a motile, gray-white, nitrate-
reducing, non-liquefying organism. On agar young colonies small, Grows well at 30, not at all at 37.5. T. D. circular, wrinkled.
P. 48^9.1.
The cause of stomatal infection of larkspur resulting in sunken black spots on leaves and stems. ^"^ B. elegans Hegyi is reported on lupine.
^'^'^
B. dahliae Hori B.
&
Bakis
is
on dahlia.
^^^
Stedman was reported by Stedman ^^ as the cause of cotton-boll soft rot in Alabama; much doubt, however, remains as to its actual identity and causal relation. It was degossypini
scribed as a short, straight, spore-forming motile bacillus;
.75
fi;
1 .5
aerobic; non-liquefying
(?).
gummis Comes, has by some been held mosis or mal new of the grape vine ^"^ though
B.
idea.
responsible for gumothers discredit this
44

B. haria Hori
&
Miy.
is
a parasite of the Japanese basket

is
willow.
^^'^
B. hyacinthi septicus Heinz/'" white rot of the hyacinth.
recorded as the cause of a soft
B. lactuca? Vogl. is said to cause a lettuce disease.'^^ B. lycopersici Hegyi has been described as the cause of a rot
of tomatoes.^^^
B. maculicola Del.
11^
is
regarded as the cause of a tobacco leaf

M.^^'is recorded in England on potatoes.
flagella
spot.
B. melanogenus P.
&
B. melonis Giddings."^ An actively motile bacillus, O.G-0.9 x 1-1.7

peritrichiate;
/z;
4-6
no
spores.
Gram negative. Broth strongly clouded,

no
pellicle
or
ring,
ip..,^
slight sediment. Agar stroke slimy, glistening
translucent;
colonies
round or amoeboid.
In
gelatine stab liquefaction infundibuliform in
two days.
agulated
with
.
Milk coabun-
dant gas.
slight.
Nitrite pro;
duction abundant indol

T.
D.
P.
49-
Opt. vegetables rotted were
50.
30.
The
muskmelon,
rot,
citron, car-
potato,
beet,
cu-
cumber and
turnip.
In the soft rot caused
by this organism in muskmelons, motile

bacteria were observed in abundance by Giddings in 1907. Plating gave pure cultures which by inoculation tests were shown to be
those of the causal organism of the rot. Decay is produced by solution of the middle lamella by enzymic action, the remainder The bacteria are thus of the walls withstanding the attack.

strictly intercellular.
45
Wound
in
inoculations in
muskmelon generally
from three to seven days. Similar inoculagave complete decay tion of citron and cucumbers resulted in decay, though inoculation into squash did not. No decay of musk-
melon followed applications to unbroken surfaces.

B.
-22.1
of the bacteria
mycoides
8-.)
Fliigge.
(Group number
Rods
thick, 0.95 x 1.6-2.4 m, usually in
long threads, sporiferous. Spores elliptical, 1.3-1.48 X 0.7-0.9 mm. Gelatine colonies
white with mycelium-like outgrowths; gelatine

liquefied.
Pellicle
formed in broth.
Gram
positive.
This
common
soil
organism has been held
^^^ responsible for a disease of beets. B. nicotianae Uyeda is ascribed as the
^^^ which cause of a tobacco wilt in Japan ^^^' resembles that caused by B. solaclosely nacearum in America.
The bacillus is 1-1.2 x 0.5-0.7 ^ with rounded ends, actively motile by peritriSpores are produced. A fiagella. complete physiological study is to be found in the articles above cited. Bacillus oleae (Arc.) Trev. (Group numchiate
ber -22.333-0.) C. O. Smith describes the organism as a motile rod with rounded
ends, 1.5-2.5 x 0.5-0.6
ji.
On
agar slant
liquefacFig. 30.
growth
tion.
thin, gray-white, spreading; colonies
circular, whitish.
On
gelatine
no
Milk not coagulated.
Distribution of
fiagella not stated. In oleander tubercles on leaves and twigs, and in oUve tubercles C. O. Smith ^^^ found
Cultures of B. mclonis on silicate jelly slants, 12 days' growth, After Gid(30 C.)
dings.
bacteria which he regards as this species. Upon puncture inoculation in both olive and oleander, tubercles were produced. Controls
were not diseased.
The organism was
reisolated
from the
46

produced knot with unchanged characters. E. F. ^ do not agree with those of C. 0. Smith. (See
savanastoi.)
120-122
artificially
Smith's results
Pseudomonas
(Group number 221.1113022.) This organism was studied by Harrison in 1901 in Canada where it was found associated with a soft rot of cauliflower, cabbages and In the rotting tissue it was always present; it was isoturnips. lated, and upon inoculation and cross inoculation characteristic The organism was reisolated in unchanged infection followed.
B. oleracese Harr.
character.
filtration,
The chemical products

also
of the
bacillus,
produced the characteristic tissue changes.
secured by Sec-
tions of diseased tissue
showed the bacteria
in the intercellular
spaces, occupying the position of the middle lamella which was softened and eventually dissolved by the bacterial enzymes.
^^ from their extensive studies conclude Harding and Morse that this form is identical with B. carotovorus. See p. 42. ^"^ as the cause of B. omnivorus v. Hall is described by van Hall a soft rot of iris shoots and rhizomes. According to Harding & Morse ^^ it does not present characters sufficient to distinguish it from B. carotovorus. See p. 42. A species closely related to B. omnivorus is described by Uyeda ^-"^ as the cause of a disease of Zingiber. The organism was isolated and studied and the disease produced by inoculation with pure culture. ^-^ as the cause of an B. oncidii (Pegl.) Stev. is mentioned
orchid leaf spot. B. oryzaB Vogl. has been mentioned as the possible cause of
brusone
B.
^'^
of rice.
phytophthorus Appel.
(Group number 221.21230.)
non-sporiferous rod, 0.6-0.8 x 1.5-2.5 n, actively motile by perGram negative. It rots potatoes, cucumbers, itrichiate flagella.
aerobic or a facultative anaerobe; grayish white on agar; surface colonies round, smooth; gelatine liquefaction moderate bouillon clouded; no indol; no gas. Nitrate changed to nitrite.
etc.; is
-
Milk coagulated and casein precipitated. Opt. 28-30. T. D. P. 47. ^-^ of Berlin as the chief cause of It was described by Appel The description given above is by E. F. Smith ^-^ potato black-leg. and was made from Appel's organism. Smith also isolated it from potatoes grown in Maine and in Virginia.

It is closely related to
47
but
is
not identical with B. solanisaprus
and B.
on the poplar ^-^. B. pseudarabinus R. G. Sm.^^" is capable of producing on inoculation a crimson-red gum in the vessels of sugar cane and is perhaps responsible for a disease showing this symptom.
B. rosarum Scalia is the name given to a very imperfectly described organism said, on scant evidence, to be the cause of rose tumors or crown galls. ^^
atrosepticus.^'^ B. populi Brizi is said to cause galls
Malkoff in infection experiments caused a with this organism. disease of sesame B. solanacearum E. F. Sm. (Group number 212.333-8 .)
B. sesami Malk.''^
A medium
Motile,
sized,
easily
stained, strictly
aerobic
bacillus
with
1.5
/x.
rounded ends; about 13^-3 times longer than broad; 0.5 x

sluggish
or
diffuse.
active; fiagella long, Spores not known. Zoogloea occur in liquid media as small, white flecks or as surface rings. It grows well at 20-30. Milk is saponified with
no casein precipitation or
acidity.
Gelatine not liquefied.

first
Agar
surface colonies, dirty-white. Agar streaks yellowish to brownish-white, then brown.
On
dirty-white, later potato as on agar,
but darker, with substratum and fluid browned. cane sugar, lactose, maltose or dextrose.
No
gas from
The
plants
J.JII
disease caused by this bacillus upon tomato and other was early studied by Halsted ^î-iss ^^j^^ perhaps by-Bur134-135 Halsted made inoculations which produced the disease
but he did not use pure cultures.
The
first
complete account of
the causal organism was given by E. F. Smith ^^^' ^^^ in 1896. In its hosts the bacillus is found in the pith, in the xylem which
is browned, and more rarely in the bark. From the cut ends of infected ducts bacteria exude as a viscid ooze and the diseased ducts
may be traced to
to leaf.
From
great distances through the plant, even from root the bundles the organism later invades other tissues.
Needle prick inoculations in tomatoes and potatoes with pure cultures, were followed after several weeks (tomato) by typical
disease.
Inoculations in Irish potato resulted similarly, though in
parenchyma and bark were eventually invaded, and the tuber was reached through its stem end and rotted. In South Carolina, Smith noted the disease on egg plants and crude cross
this host the
48

made
to tomato.
inoculations were
Smith demonstrated
experi-
mentally the efficiency of the potato beetle in transmitting the

disease.
The
and
disease
was described
for tobacco
by Stevens
^^^
and Stevens
Sackett.^39
in 1909 though pepper gave negative results when inoculated with this bacillus. In addition to the above hosts it is known to grow upon Datura, Solanum nigrum, Phy sails and Petunia. B. solanicola Del. was reported as the cause of a potato stem
disease.^'*^
Smith
Successful inoculations were reported upon tobacco by E, F. " in his earlier trials tobacco and
A rod, B. solaniperda Mig. (Group number 121. .) 2.5-4 X 0.7-0.8 n, with rounded ends, often in long chains; actively motile spores present. Agar colonies dirty-white gelatine liquefied.
;
;
This was shown by Kramer in 1890
^^^
to be the cause of soft rot of potatoes.
The organism
was grown in pure culture and inoculated on potatoes

producing the characterisdecay. The germs enter through the lenticels, contic
sume the
sugar, then
cell
at-
tack the intercellular substances and the

wall.
Later the albuminous substances are destroyed. B. solanisaprus Harr.
(Group number 221.212-0_ \ 143

Fig. 31.- -Surface colonics of B. solanisaprus.
bacillus with
rounded
m,
After Harrison.
ends,
1.5-4
0.6-0.9
variable in culture; no capsule; actively motile by 5-15+ peritrichiate flagella; no spores seen. Gram negative. Gelatine colonies, punctiform 0.25 mm. at two days; gelatine stab filiform. Liquefaction noticeable on the thirty-fifth day. Agar colonies punctiform at two days, 1-5 mm., gray-white, slimy, flat. Bouillon turbid
with fine sediment; ring, and thin band present; milk curdled.

Gas only
in
49
mannite and lactose. Nitrate reduced to nitrite. T. D. P. 54, 10 min. It was found constantly associated with a type of potato disease which Harrison regarded as distinct from black-leg and from the
Opt. 25-28.
by B. was isolated repeatedly

disease caused
solanacearum.
It
from diseased tubers, stems and leaf veins and occurred in practically
pure culture
in
freshly infected tissue.
The organisms first appeared in the ducts and thence invaded

the
sue,
surrounding
tis-
dissolving middle lamellae
the
producing
and cavities.
solanisaprus,
of pure cultures into healthy Fig. 32. B. plants produced characteristic lesions
Inoculations
from
agar
24
hours.
After Harrison.
enzymic action
slices of potato.
and the organism was reisolated. Characteristic was observed on placing precipitated enzyme on
B. sorghi Burr."* Rods 0.5-1 (usually 0.7) x 1-3 (usually M, cylindrical or oval, motile, spore-bearing, non-liquefying. Colonies on agar, white to pearly. In broth with a white smooth
1.5)
membrane.
The
by
bacillus
Burrill
and
this
was recognized as the cause of a sorghum blight view was confirmed by Kellerman & Swingle
through
"^ inoculation experiments. " causes B. spongiosus A. & R. gummosis of cherry in Ger-
many.
B. subtUis (Ehr.) Cohn. Sporiferous. Straight rods, often united in threads, 0.7 x 2-8 /x. Spores central or lying near one pole; germination equatorial.
Flagella,
6-8,
peritrichiate;
gelatine liquefied; gelatine colonies
50
bordered by numerous fine filamentous outgrowths. Growth on slant agar gray. It is reported as the cause of vegetable rot.^'*^
B. tabacivorus Del.
is
recorded on tobacco stems.
B. tabificans Del.^^ which perhaps belongs to the genus Bacterium is reported as the cause of a beet disease in France.
B. tracheiphUus, E.
F.
Sm.
(Group number 222.
03-.)
THE FUNGI WHICH

Bacillus 1.2-2.5 x 0.5-0.7
cultures.
fx,
CAUSP:
PLANT DISEASE
51
variable, actively motile in

peritrichiate.
young
Capsulated,
no spores,
No
liquefaction. On agar thin, smooth, milk-white. No or facultative anaerobic. Milk not curdled. T. D. P. 43, 10 min.
gelatine gas, aerobic
This pathogen was first reported by E. F. Smith without de^^^ and more fully in 1895.^^ It is found filling scription in 1893
the vessels of cucurbits, (musk melons and cucumbers) affected with wilt. Smith produced the disease artificially by puncture
inoculations on the blades of leaves with the white sticky fluid from infected veins. The inoculated plants showed symptoms of
wilt after four days and sixteen days later the ducts of the vine were found to be plugged with bacteria. The organism was then The cultures thus isolated from this artificially infected plant. obtained were carried by transfers over winter and in December were used successfully to
infect
cucumber
plants.
%
*
^^
,
'
_i__
Control plants were never
\^ ,5;
%
diseased.
growth
of the
is
The ready
organism
in
1
'
.-*'
^
J
*
'*
^ *^
**
*
*
'
the vessels
attributed to
Tv-.^!/
o^î
'*-*"'
V^
the alkalinity of the latter
the failure to grow in the parenchyma is attributed

to
its acidity.
^, ,,
v.
-"-^^
B. uvae Cug. & Mac. is reported as causing injury

to
It
T
^j,*
'^**''L-c,
^*
C*'
^
^*-
C'V^
^^^r*
young grape
is
clusters.
^^^
\^
^^
%*
perhaps identical with

(Flugge)
B. ampelopsorae. B. vulgatus
This organism is Mig. found as small thick rods with rounded ends, or is
often paired or
-in
Fig. 34.
B. tracheiphilus.
After Smith.
chains of four; sporiferous.
Gelatine colonies
dirty-white. round, liquefaction rapid. ^^^ It has been shown capable of causing rot of various vegetables. B. zeae Burr, is the name applied to a bacillus isolated from
Growth on agar
diseased corn plants
by
Burrill in 1887-1889.i^3'
^^^
It
is
often
52
cited as the cause of a bacterial corn disease but the evidence of
causal relation as well as the identity of the germ are not clear. B. zinzgiberi Uyeda causes a disease of Zinzibar,-^ B. sac-
chari
,
and B. glangae are on sugar cane as the possible cause

58
of
sereh.
An organism called Clostridium persicae-tuberculosis by Cavara ^^^ is mentioned as cause of knot on peach trees. Less known bacterial plant diseases. The literature abounds in references to what are regarded as cases of plant bacteriose, cases which as yet rest upon very incomplete evidence. In many of these bacteria are found in abundance in the diseased tissue
but pathogenicity has not been proved by inoculation nor pure cultures made. Among such incompletely studied diseases may
be mentioned those of geranium; i^^"!^^ celery, ^^^ onion, ^^^ orchard grass, ^^^ lettuce, cucumber, (one lettuce disease is due to a motile rod-shaped organism cultured and inoculated
^^
'
but not named, ^^'*) strawberries, ^^''"^^^ mulberry, ^^^ hemp,^^ calceolaria.
^^^
There are also several obscure bacterial beet diseases; another ^^^ a decay of apples said by cabbage rot due to Pseudomonas; ^^^ Prillieux to be due to a Bacillus; the blossom-end-rot of tomatoes which is perhaps bacterial; ^^^ a cyclamen leaf spot; ^^^ a juniper dis^^ a pine gall; ^^" an ash bark disease; ^^^ and an ash canker; ^^^ ease; a salsify rot; ^^^ a carnation an ivy canker; ^^'^ a grape disease;
^^^ and a banana disease; ^^^ a gummosis of tobacco; ^^* a spot; ^^^ also perhaps the serious widespread disease of tobacco seedlings; mosaic disease of tobacco and an orchid gummosis. ^^
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Jour.
Am. Pub. H.
1896, p. 76, Torr. Bot. CI. Assn., Jan. 1898: 60;
Recommendation
for the
study of Bacteria.
210
See also Rept. Soc. Am. Bact. Meeting of 1907. Pavarino, L., Rend. d. r. Ac. d. Lincei, Classe Scienze, 20: 161, 1911.
DIVISION
EUMYCETES. TRUE FUNGI
The Vegetative Body
consists of a
III
'" '' '' '' "' "*'
(p. 3)
mycelium.
is devoid of chlorophyll and typically or less branched filament of apical growth, the This mycelium may be cut into cells by partitions
more
(septa) or may be continuous, i. e., without septa. The cells of the septate mycelium do not differ essentially from typical plant cells except in the absence of chlorophyll. They consist of masses
of protoplasm, the protoplasts, bearing vacuoles less rich in oils, acids, ^
and are more or
gums, alkaloids, sugars,
resins,
etc.,
coloring
matter,
in
varying
amount and kind

the
particular
with
species of the
and condition
fungus.
is
The
protoplast
covered
cellulose
by a
cell
wall which
of
consists
though
often
of
special quality
known
as fungous cellulose.
The
protoplast bears
Fig. 35.- -Showing a septate
cells.
one or in some fungi
two or more
cells,
mycelium within host After Stevens and Hall.
nuclei.
of the protoplasm, the
The vacuolation
teristic.
mode
of branching of the
their color, dimensions, etc., are in
some cases quite charac-
In one class, the Phycomycetes, the active vegetative mycelium possesses no septa except such as serve to cut off the sexual or other reproductive organs or such as are found in senility. The
59
60

is
protoplasm
therefore continuous throughout the whole plant
body and may be regarded as constituting one cell though it may be of great extent and bear very numerous nuclei. Such multinucleate
cells, coenocytes, the walls omitted.
may
be regarded as
cell
complexes with
In one comparatively small order, the Chytridiales, there is often no filamentous mycelium and the vegetative body consists merely of a globular, irregularly spherical or amoeboid cell. Such
in
forms are thought by some mycologists to be degenerate, to have remote time possessed a mycelium which has been lost owing to
the present simple
mode
of life of the fungus, the needs of

^
which
no longer
call for
a filamentous body, while others
find here
primitive forms of Phycomycetes, and trace their phylogenetic connection with the higher orders of the class.
Reproduction.
Most mycelia, if cut in bits and placed in suitable environment, continue to grow, soon equaling the parent mycelium in size if abundant nourishment obtains. Bits
Vegetative.
of diseased tissue, bearing
stitute
persal.
mycelium, thus con-
ready means
of multiplication
and
dis-
Asexual Spores. A spore is a special cell set aside to reproduce the plant. An asexual spore is a spore not produced by a sexual process.
Manifold forms of asexual spores
the fungi.
exist
among
In some of the simplest cases, budlike out-growths (gemmae) appear on the mycelium; or portions of the mycelium itself are cut
off
by
partitions
and the protoplasm
inside
gathers into a mass and protects itself by a firmer wall than that of the mycelium, chlaFig.
mydospores. In other cases special branches, 36. One form hyphae, are set apart for the purpose of bearing of conidium. t/- ji p r j.tj.* r If the spores are cut on from the tip of Oidium. After spores.
j.
^Yie branch they are known as conidia or conidioand the branch bearing them is a conidiophore. Conidia spores, may be borne singly or in false clusters caused by the youngest pushing the older conidia aside; frequently they are produced in
Bioiotti.

chains, catenulate, Fig. 36,
either simple,
61
owing to the development of one spore below another before the elder spore is shed. Conidia may be
or
of one cell, or compound, composed of two In compound spores each cell is at least potentially a spore and can germinate under favorable conditions and per-
composed
more
cells.
petuate the species. In many compound spores the germinating function is sacrificed by one or more of their component cells. Conidiophores may consist of loosely branching, rather long
hyphae, or they
distinct
spots.
may
Fig.
spore 371.
spots
be short, innate, and in close clusters forming bearing
Such
sporiferous naked are

vuli.
when
acer-
called
Often the conid-
iophores are roofed over
with
net-work
of
woven fungous threads thus constituting a
special
spore-bearing
structure, the pycnidium.
Conidiobe solitary or grow together in bundles or branch loosely as

Figs. 37, 335.
phores
may
in Fig. 383.
The basidium, Fig. 38, a special kind of sporophore bearing at its ,, n r apex usually tour, or
is
Fig.
37. Conidia borne
in a pycnidium. Quaintance and Shear.
After
two, small projections, sterigmata, each of w^hich produces one spore, for distinction called a basidiospore.
Some
fungi bear the spores loose inside of the swollen tips of
sporophores as in Fig. 68. The spore bearing structure is then called a sporangium and its stalk a sporangiophore. The ascus is another spore bearing structure. In it the spores are borne very
much
ber,
as they are in the sporangium but usually of definite

2,
num-
1,
4,
8,
16, etc., eight
being the most
common number.
Asci
may
be naked or covered, scattered or collected in groups.
62

covered, the chamber in which they are borne is called a when on an open disk the disk is called an
life spores are classed as: 1. restto tide over unfavorable conditions,
When
perithecium, Fig. 39;
apothecium, Fig. 101. According to their length of

ing spores whose function
is
hence the
in
common name
"winter
"Summer
spore," and contradistinction: 2.
spores" which produced in abundance in warm weather,

are
germinate immediately, and can ordinarily live
but a short time. In some species the

spores that are to func-
r-sk
Fig. 38.
tion in water possess cilia, and the power of motion.
Basidia
of various ages.
After Schenck.
These are zoospores or
swarm
At sporing time many kinds
for the bearing of spores.
spores, Fig. 44.
produce special structures The fungous threads interweave to

of fungi
form a
firm, or
parenchyma.
even a densely solid, mass and constitute a false Such are the stalks and caps of the mushrooms and
cross of the shelving toadstools, the skin of the puff ball, etc. section of such a structure appears much as a true parenchyma, a longitudinal section shows it to be merely a mass of interwoven
fungous threads. Sexual Spores are formed by the union of sexual elements, gametes. They are most conspicuous among the Oomycetes where the antheridium carries the sperms into the oogonium, fertilizes the obsphere and produces an oospore. Figs. 53-55. As a rule the sexual spores are produced toward the end of the
vegetative period of the fungus.
earlier
The
asexual spores are produced
and
for
a longer period.
Sexual spores are commonly
resting spores.
Germination of spores. Under suitable environment mature spores germinate and eventually give rise to vegetative bodies
63
The most usual mode is for the similar to that of the parent. to rise directly from the spore. In other instances the mycelium spores produce zoospores which migrate, come to rest, then develop
a mycelium. In still other cases a short mycelium, promycelium, is formed and from this small conidia, sporidia, are made. Figs. 217, 240. These conidia give direct rise to the mycelium.
Spores of some species may by gemmation lead a more or prolonged existence without return to the mycelial stage.
less
Heat and Moisture Relation.
Like
all
living
things these
The organisms cannot develop without heat and moisture. Some necessary degree of each varies with different species.
qermimr/on
Fig. 39.
A perithecium with
asci.
After Reddick.
species are strictly aquatic, and must be surrounded with water; others can grow in comparatively dry situations. Generally
speaking, however, dampness favors fungous development, and the growth of most fungi is more vigorous in a damp atmosphere than in a drier one. Similarly moderate warmth, as that of summer
heat, favors fungous growth. Humidity and warmth combined are proverbial as producers of mold and mildew. So conspicuous is the coincidence of these conditions with fungous growth, that
minds of many a warm damp air is the cause rather than the condition of fungous development. Respiration with the fungi as with other plants and animals consists in oxidation, involving intake and consumption of oxygen
in the
accompanied by the giving
off of
carbon dioxide
antl water,
and
64
since
is

no photosynthesis occurs,
this process
is
never masked as
it
in the case of the chlorophyll-bearing plants.
In nutrition requirements there is great diversity; but in all must be taken from some organic source. Starch, cellulose and kindred compounds are frequent sources of sugar,
cases carbon
the carbon food supply. Nitrogenous foods are, generally speaking, not required in such abundance by the Eumycetes as by the bacteria and advantage may frequently be taken of this fact in
in nitrogen, in
from bacteria by growing them on media poor which case the fungi often outgrow the bacteria. The color of the fungi is determined largely by the constitution ^' *' ^ of the media upon which they grow.^*
isolating the fungi
Many
life
fungi exhibit a peculiar heteroecism, that

is
is,
part of their
cycle passed through upon one host, part of it upon another even of very distant botanical kinship. Thus among the host, rusts; in one instance part of the life cycle is upon the apple, the remainder upon the cedar tree. Fungi also exhibit polymorphism, i. e., in one stage they exhibit one spore form and in anIn this way other stage another spore form totally different.
several apparently quite distinct types of spores structures may belong to the same species.
Classification of Fungi.'
'^'
and sporiferous
^^' ^^'
selves constitute a very large group

of
The true fungi in themmade up of diverse forms, many

^'''
^^
which are as yet

is
little
known.
fication
impossible until
Any satisfactory system of classimuch more knowledge is gotten regard-
ing their morphology, cytology, life histories and especially their relations to their hosts. According to present knowledge they comprise very
numerous
species distributed in three classes as follows:
Key
Mycelium
stage
to Classes of Eumycetes
Class
1.
continuous in vegetative
Phycomycetes,
Ascomycetes,
p. 65.
Mycelium septate
Spores in asci Spores on basidia*
Class
2.
p. 113.
Class 3. Basidiomycetes, p. 298.
Not
as above; spores on conidiophores, naked, or in pycnidia; or spores quite unknown
Fungi Imperfecti,
is
p. 475.
In the rusts and smuts the promycehum
regarded as a basidium.

Class
65
I.
Phycomycetes, Alga-like Fungi

are characterized
(p.
64)
by the absence of septa in the mycelium except in sporing branches, where they occur to cut off the spore-bearing cells or the gametangia, and in old filaments. The body is multi-nucleate and sexual spores as well as
asexual ones are usually, though not always, produced. Some of the Phycomycetes live in water and possess zoospores, others are These parasitic on land plants and bear conidia or sporangia. may germinate either by germ tubes or by zoospores. The characteristic fertilization consists of
The Phycomycetes
a union of two gametes which
may
(isogamy) or unlike (heterogamy). If the sexual organs are unlike the receptacle which bears the sexual spores is called the oogonium, its eggs before fertilization oospheres,
be
like in character
and the spores oospores. The receptacle bearing the fertilizing gamete is the antheridium, and the fertilizing elements are the sperms. The sperms may be motile and swim or creep into the oogonium or the antheridium may develop a tube leading In into the oogonium through which the fertilizing nuclei pass. some forms which, by their sexual or asexual spores, show relation to the Phycomycetes the mycelium is wanting and the vegetative body is reduced to a single spherical or amoeboid cell, which frequently lives in a purely parasitic manner entirely imbedded in
the protoplasts of
strictest
its
host.
This
mode
of
life
constitutes the
its
kind of parasitism inasmuch as the fungus derives nourishment from the still living host cell.
Key
Sexual
spores
to Orders of Phycomycetes
present
when
mous
sporangia only
heterogaSubclass
I.
Obmycetes,
p. 66.
Conidia absent; sexual spores and zoo-
Mycelium poorly developed, frequently

reduced to a single
cell
cell,
Fruiting mycelium a single
or a
group
of cells in a sorus, forming
either asexual
resting spores or sporangia from the entire proto-
plasmic mass
1.
Chytridiales, p. 66.
66

Fruiting
mycelium
cells
multicellular,
some
forming sporangia,
others producing
gametes and
2.
oospores
Ancylistidiales.
Mycelium
well developed Fertilization by motile sperms. ...

Fertilization
idial
3.
Monoblepharidiales.
Saprolegniales, p. 74.
through an anthertube
4.
5.
Conidia present Sexual spores isogamous, formed by the union of similar gametes.. Subclass Asexual spores several, in sporangia. Asexual spores sohtary, conidia
.
Peronosporales,
p. 77.
II.
6.
7.
Zygomycetes, p. 101. Mucorales, p. 102. Entomophthorales, p. 107.
Of these orders the Ancylistidiales which are parasitic upon Algse, and the Monoblepharidiales which are saprophytic will not
be considered further.
Subclass Oomycetes
(p. 65)
In the Oomycetes there is pronounced difference between the male and female sexual organs. The oogonium is comparatively large, and contains one or more large passive eggs (oospheres), which are fertilized by sperms, differentiated or not, which either
swim
to the
oogonium by
cilia,
creep to
it,
or are carried to
it
by
Oospores are in some species produced frequently and abundantly while in others they are entirely unknown. The asexual reproduction is by either conidia or sporangia.
a fertilizing tube.
Chytridiales
(p.
65)
The members
of this order are the simplest of of
any
of the
Phy-
are single, more or less globose, undifcomycetes. Many ferentiated cells, others have a more or less prominent haustoria-
them
mycelium, while but few have any approach to a true mycedevelopment. Most are intracellular parasites; a few of the more highly developed genera are intercellular parasites. With few exceptions reproduction is entirely asexual, all spores being formed directly from the vegetative cell. Zoosporangia and thicklike
lial
67
one or two
species.
walled resting spores are produced. The zoospores have cither There are over forty genera and two hundred cilia.
The majority
and
of the species are
inconspicuous parasites
of algse
some genera, like Synchytrium and Urophlyctis, produce conspicuous sori and even cause hyperinfusoria; but
trophy of land plants.
Key to Families of
Chytridiales
Spores all asexual, or rarely formed by the union of free-swimming gametes
Mycelium none
Sporangia solitary Sporangia grouped into
1.
sori
2.
Olpidiaceae, p. 67. Synchytriaceae, p. 69.
Mycelium present Mycelium of delicate, evanescent haustoria-like strands
Mycelium
minal
limited,
sporangia
ter3.
Rhizidiaceae.
Mycelium extended, sporangia

minal or intercalary
ter4.
5.
Cladochytriaceae,
p. 72.
Mycelium permanent hyphse Spores both sexual and ase.xual

Gametes hetrogamous Gametes isogamous
of
Hypochytriaceae.
Oochytriaceae, p. 73.
Zygochytriaceae.
G.
7.
Four only
algae
of these families
higher plants in
have parasitic representatives on the others being chiefly parasitic on America,
and
infusoria.
Olpidiaceae
This family which contains the simplest members of the order has no mycelium; the entire plant body consists of a single more or less globular or elliptic cell which never divides, but at maturity forms either a zoosporangium or an asexual resting spore which
after a period of rest gives rise to
are endobiotic.
of
swarm spores. The family contains some forty
All the species species but few
which are of economic importance.
68

Key
to Geneea of Olpidiaceae
1.
Vegetative body amoeboid Vegetative body of definite form Sporangia free in the cells of the host
Sporangial membrane evanescent
Reessia.
very
firm,
delicate,
2.
Sphaerita.
Sporangial
ing
membrane
by a
swarm
spores escaping
definite open-
Sporangium globular or ellipsoid Sporangium with only one or two

openings
Swarm
spores uniciliate
3. 4.
5.
6.
Vegetative cells globose or subglobose
Olpidium,
p. 68.
Vegetative
cells stellate
Asterocystis, p. 69.
Olpidiopsis.
Swarm
spores biciliate
Sporangium ^^^th several openings Sporangium elongate

Sporangial membrane united to the wall
the host
cell
Pleotrachelus.
Ectrogella.
7.
of
8.
Pleolpidium.
Olpidium
Braun
In this genus a single swarm spore invades the cell of the host and develops in its proLater a cell toplasm.
wall forms
tative
and the vegeinto
body changes
a zoosporangium which develops a neck. This

reaches to
of the host
the
outside
even though the fungus be developed several cells below the

surface.
Fig.
The
this
uniciliate
40. O.
cell; left,
brassier; right, three sporangia in resting spores. After Woronin.
ZOOSporeS
through
paSS OUt
neck
tO
make
their escape.
Thick-walled resting spores are also formed.
69
There are some twenty-five species most of which hvc as parasites on algse, worms, pollen grains, etc. O. brassicae (Wor.) Dang.^ is parasitic on quite young cabbage seedlings, sometimes infecting cells deeply seated in the host. The same or a nearly related species also attacks tobacco and
several weeds.
Sporangia solitary or several in each infected host cell, globular; zoospores numerous, globose, uniciliate; resting spores globose, with a wrinkled epispore which gives them more or less of a starlike
appearance.
Fig. 40.
Asterocystis de
Wildeman
d. cell
(p.
68)
There
is
a single species, A. radicis

its stellate
Wild.^ which differs from

of the
Olpidium in
vegetative
and the absence
tube
for the escape of the zoospores, this being accomplished by the breaking away of the tissues of the host. The fungus attacks the
roots of various plants, notably flax, Brassica
and other
crucifers,
Plantago, Veronica and numerous grasses, producing chlorosis. It has not been reported from America. A Chytridiaceous fungus of unknown genus thought to stand
near the Olpidiaceae and Synchytriaceae has been described by Home ^ as the cause of an Irish potato disease.
Synchytriaceae
(p. 67)
cell
The
infecting
zoospore invades the host

still
parasitic
upon the
living protoplasm.
Hypertrophy
and becomes of this and
adjacent host cells is usually induced, resulting in the formation of a small gall around the infected cell. This gall is often colored and bears a superficial resemblance to a rust sorus. The parasite In enlarges until it occupies nearly the whole of the host cell.
S>Tichytrium the one nucleus then enlarges and divides to produce ^' ^"' ^^' ^" The whole mass then divides very numerous nuclei.
into segments regarded as sporangia, and each sporangium divides into numerous uninucleate parts, each of which develops into a
In some species development is arrested before the zoospore. division of the primary nucleus and the protoplast becomes spherical, invests itself with a thick wall and becomes a resting
70
spore.
this

(Fig. 42.)
After a more or less protracted period of rest
produces zoospores.
The family includes some fifty species, all of which, except two small genera, are parasitic upon land plants.
Key
to
Genera of
Synchytriaceae
Zoosporangia formed by direct division of the entire plasma of the young fruiting
body.
Swarm
sporangia completely filling the host cell, membrane united to the

wall of the host
cell
cell 2. 3.
1.
Rozella.
Swarm sporangia lying free in the host

Parasitic on algse
Parasitic
Woronina.
Woroniella.
on land plants Zoosporangia formed by division of an initial cell to form a sorus of sporangial cells. Sporangia formed directly, from the full-
grown plant body

Sporangia formed by the division of a thinwalled mother cell after its escape
4.
Synchytrium,
p. 70.
from the plant body
5.
Pycnochytrium,
p. 72.
Synchytrium de Bary
& Woronin
Upon reaching maturity the plant body develops directly into a sporangial sorus. Both zoosporangia and winter spores present.
Fig. 41.
Showing nucleus
(Schilb.)
in
bynchytrium.
After Stevens.
S.
endobioticum
disease of the potato,
was
Perc, the cause of a very serious wart originally described as Chrysophlyctis
endobioticum by Schilberszky ^^ and transferred to Synchytrium by Percival.^^ It invaded America about 1909.^^^ It was reported from Africa by Zimmermann.^^

In
71
summer
eter are found in
the resting spores which average about 52 n in diamabundance in the host cells near the surface, few
in the outer layer, more below down to the sixth or eighth
row
spore
of
cells.
Each
contains
resting several hun-
dred roundish zoospores which measure 2-2.5 /x. In spring

the resting spores germinate,
Fig. 42. A, section showing sporangia or sporocysts; B, zo6si)ores, ciliated and amoeboid. After Percival.
numerous pear-shaped which at first swim with a jerky motion but soon become amoeboid.
freeing
uniciliate zoospores,
The summer sporangia may germinate without protracted
rest,
and also give rise to zoospores. Another type of sporangium consists of thin sacs, produced singly or two to five in a sorus,
each bearing numerous zoospores somewhat smaller than those
from the
first type of sporangia. zoospores, says Percival, enter the potato apparently in the amoeboid state in bud tissue of rhizomes and in the "eyes" of
The
young
tubers.
Usually only one zoospore enters each
cell
but
Crushed sporangia produced occasionally more may do so. characteristic warts in three to four days when placed on susceptible parts.
Successful inoculations were also
Crompton.^^ grown tumors vary in size from that of a pea to a hen's egg, and represent metamorphosed branch systems.
full
made by Salmon and The cytology has been studied by Percival. ^^ The
S. vaccinii
Thomas
^^"^^
is
the cause of a
disease of the cranberry and related hosts. It forms numerous, small, reddish galls in which,
deeply embedded, are the sori. S. papillatum Farl.^ occurs on Alfilaria in

California.
Fig.
43. Gail of S. vaccinii. After
Other species of Synchytrium are found upon
Shear.
dandelion, CEnothera, Geranium, Amphicarpa, Ornithogalum, clover, elm, etc., but as yet are not of economic importance in America.
72
Pycnochytrium Schroter
(p. 70)
Only resting spores are known. In germination their protoplasmic contents emerges and forms a sporangial sorus. P. anemones (D. C.) Schr. is common on various species of Anemone; P. globosum (Schr.) Schr. on the violet, blackberry,
maple,
etc.
None
of the species are of
any considerable economic
importance.
Cladochytriaceae
(p. 67)
cells of
branching mycelium runs through or between the
the host drawing nourishment from many cells. Sporangia are either apical or intercalary and contain uniciliate zoospores.
Resting spores are also produced.
There are about a half dozen
genera and some thirty
species.
Key
Resting spores
onlj^
to Genera of Cladochytriaceae
1.
Swarm
spores only
known known
first ciliate,
Physoderma.
Intracellular
and endophytic
becoming
2.
Swarm
Swarm
spores at
amoeboid
spores not becoming amoeboid
3.
Cladochytrium, p. 72. Pyroctonium, p. 73.
Living free
among
the hosts
4.
5.
Sporangia opening by a pore Sporangia opening by a lid
Amoebochytrium. Nowakowskiella.
Cladochytrium Nowakowski-^
The genus contains about ten species of intercellular parasites with branched mycelial threads. The zoosporangium is globose, and opens by a distinct mouth which develops a tube for the escape of the zoospores much as does Olpidium. Resting spores are not known.
73
The most important species are C. tenue Nowak. on Acorus and Iris; C. graminis Busg. on various grasses, C. violae Berlese on violets.-^ C. viticulum Pru.^ and C. mori Pru.-'* have been described on grape and mulberry, but further study is very desirable. C. brassicae E. & B.-'' is described from dead leaves of cabbage. C. caespitis G.
& M.^^ occurs in France on Lolium. Pyroctonium sphaericum Pru.-^ was reported in 1894 as the cause of wheat disease in France but has not since been
found.
Oochytriaceae
(p.
67)
cell or
The
plant body
is
either
an undifferentiated
a well de-
veloped mycelium; reproduction by means of asexual swarm spores and sexual resting spores. Of the three genera only one
is
of
economic importance.
Key
to Genera of Oochytriaceae
1.
Mycelium entirely lacking Mycelium present Mycelium producing a
Diplophysa.
single gametangium Mycelium producing several gametangia
2.
Polyphagus.
Urophlyctis,
p. 73.
3.
Urophlyctis Schroter
Mycelium endophytic, producing zoosporangia on the

of the host
uniciliate.
surface
and
thick- walled oospores within the tissues; zoospores The genus contains some half dozen species all of
-^
which are parasitic on higher plants.

causes "beet root leperoides (Sacc. & Trab.) Magnus -^' tumor," in North Africa and Western Europe. The rootlets of
XJ.
the upper portion of the root are attacked and develop tumorous growths, sometimes as large as a walnut. The infection is super-
74
ficial

and does not extend to the
of the spores is the typical
fleshy tap root.
The develop-
ment
method
for the genus, the an-
theridium persisting at the base of the oogonium and retaining its hyphal connection, while the oogonium becomes free just before
conjugation.
The oospores
are
subglobose, depressed on one side, smooth, brown, 45-50 x 30 n.

(Wallr.) Schr., a related species occurs on closely
U. pulposa
the aerial portions of Chenopo-
dium and
Atriplex.
U. alfalf Mag.^' ^"' "^' ^î' ^"^ causes a crown gall of alfalfa in
America and Europe.

ease
is
The
dis-
FiG.
Mag., a forms closely species, on the leaves and petioles of small, glassy, globose pustules various species of clover in Europe. U. hemispherica (Speg.) Syd.^^ in South America, U. kriegeriana Mag.^^ in Europe and U. pluriannulata (B. & C.) Farl.în America form Synchytrium-like galls on various umbelliferous genera. All may belong to the same species. U. major Schr. and U. rubsaameri Mag. infect respectively the leaves and the roots of
formation;
Schroter.
d,
44. Urophiyctis pulposa. a, sporangium; h, zoospores; c, oospore

'
quite similar to that dezoo- scribed above for the beet
mature oospores. After
U.
trifolii
(Pass.),
related
Rumex.
Saprolegniales
Asexual reproduction
large
is
(p.
66)
mainly by
biciliated spores
formed
in
in sporangia of various shapes. often apogamous, are produced in most genera,
numbers
Sexual spores, much after the
fashion of those of the Peronosporales except that
more than one
oospore The order consists of

or saprophytes
is
frequently formed
in
one oogonium. ^^"
fifty or
more
species,
upon aquatic organisms. One causes serious disease in young fish. Achlya
There are three families:
mostly parasites species of the genus

Key to
75
Families of Saprolegniales
Vegetative mycelium of thick tubular hyphae; aquatic; zoosporangia cylindrical not much thicker than the mycelium
Filaments uniform, not constricted Filaments constricted regularly

Vegetative mycelium of thin hyphse, mostly
parasitic or saprophytic
1.
Saprolegniaceae.
2.
Leptomitaceae.
on plant
tis-
sues; zoosporangia
much broader than

3.
the mycelium, mostly globular
Pythiaceae, p. 75.
Dictyuchus Leitgeb.
This genus of the Saprolegniaceae contains the only parasite
genus in the first two families. Sporangia cylindric or clavate, swarm-spores becoming walled within the sporangium and emerging singly through its lateral walls. The genus is usually saprophytic but, D. monosporus ^"^^ Leit. is said by Halsted to be a serious hyacinth enemy. The other members are mainly on dead or diseased insects or other animals that are in water or are on diseased algae or in watersHme.
"^
Pythiaceae
This family shows affinity with both the Peronosporales and the Saprolegniales and is sometimes classed with the one, sometimes with the other. It consists of three genera and about twenty
species characterized by a mycelium of very delicate hyphae which show no differentiation into sterile and fertile regions. The species are either aquatic or terrestial; in the latter case they are
When of aerial soil fungi that grow to maturity upon seedlings. habit the sporangia become conidial in character, that is, they are detached from the hypha before the discharge of the zoospores.
Zoosporangia elongate Zoosporangia spherical or oval, not linear Zoospores formed outside of the zoosporangia
2.
1.
Nematosporangium.
Pythium,
p. 76.
Zoospores formed within

rangia
the
zoospo3.
Pythiacystis, p. 77.
76

^^
Pythium Pringsheim
(p. 75)
The mycelium
tissue as fine,
found in abundance in and about the infected branched continuous threads. These, in the terrestial
is
Fig. 45.
Cucumber seedlings.
mycehum
itself.
Pots
5, 6,
Control.
and 8 inoculated with Pythium. After Atkinson.
Pot
7,
species, bear conidia
on branches which are of the same character
as the
The
of the wall
conidia germinate either by a rupture or by the formation of a process

is
beak-like
through which the

it
protoplasm
extruded, after which
becomes differentiated into zoospores. Gemmae, very like the conidia in appearance, are also produced. The oogonia are quite like the conidia
and gemmae
in structure
but develop
oospores within. The oogonium is at first multinucleate but as the oosphere Fig. 46. Fertilization in Pythium, showing oogonium, matures all of the nuclei except one
antheridium,
oospore,
cf
peri-
plasm
nuclei.
and the
and
After Miyaki.
migrate toward the periphery, the periplasm, or degenerate in the ooplasm, re-
sulting at maturit}^ in an uninucleate egg. This is fertilized by one nucleus from the antheridium. No sperm is differentiated,

and the contents
77
of the anthoridium are carried over to the egg
by a
fertilizing
tube.
Members
of
the
genus are aggressively parasitic only under most favorable environmental conditions
of heat
and moisture.
sixteen species are
Some
P. de
baryanum
Hesse,
known. is most comOff."
mon
^^'^^
as the cause of
"Damping
/x;
Zoosporangia or "conidia" globose to

elliptic,
usually papillate, 20-25
gemmae
form and size; oospores globose, smooth, 15-18 /x. hyaline, P. intermedium de Bary, causes a
similar in
Fi(!. 47.
P. citriophora; dn(if
vclopmciit
s\v;irnispores
"damping
gracile
off"
of
fern
rot
prothalia,^^
of
^^
P.
from
sporaiiKiii.
After
Schenck,
palmivorum Butler, a palm

Pithiacystis,
P. ginger; disease in India
Smith and Smith.

3C.
:> 1
Smith
&
Smith
(p. 75)
The sporangiophore
sporangia sympodially.
is
delicate; septate;
and bears numerous

biciliate zoospores
These produce many

internally.
No
oospores have been

is
seen.
Only one species
known.
^^
P. citriophora Sm.
Parasitic
&
Sm.^^-
on lemons, the
the
sterile
rind; inhabiting spores normally formed in the soil near infected fruits sporangia ovate
;
mycelium
Fig. 48. Sporangiophores and spoAfter rangia of Pythiacystis.
or lemon-shaped, papillate, 20-60 x
Smith and Smith.
averaging 35 x 50 n, borne sympodially; zoospores 10-16 /x.
30-90
jjL,
at first elongate,
This was
first
in California.
becoming rounded and bearing two lateral cilia. ^^ on rotting lemons noted by Smith and Smith Infection by pure cultures proved that the fungus
''''
was the true cause
of the rot.
Peronosporales
(p. 66)
These fungi constitute an order characterized by a richly developed, branching, non-septate, usually coarse, mycelium of
78

The mycehal threads
cells
strictly parasitic habit.
wander between the host
in most genera and draw nutriment from them by
short branches, sucking organs (haustoria), (Fig. 49) of various forms, which penetrate into the victimized cell. In one genus
only, Phytophthora, does the mycelium grow directly through cells. Two kinds of spores are produced, sexual and asexual. The
sexual spores result from the union of two unlike gametes, the egg (oosphere) and
sperm, borne respectively in the oogonium and antheridium. Each oogonium bears a
Fertilization is accomsolitary oosphere. plished by means of a tube from the anther-
idium and penetrating into the oogonium.

FiG. 49.
After sistant, and usually require a long time to Peronospora. ^P^reach maturity. They are, therefore, often In germinating the sexual spores procalled "resting spores."
,
Haustoria
The
of a
.
sexual
,
spores
,,
are
thick walled,
.
re,
,.
duce either germ tubes or develop directly into zoosporangia. The asexual spores are conidia. They are borne on conidiowhich arise from the mycelium and which may be short
phores
or long, simple or branched, subepidermal or superficial accordin various genera ing to the habit of the species. The conidia three methods, (1) a germ tube is sent out by
germinate by the conidium,
of the spore (2) the entire protoplasmic contents a germ tube, or passes outside the spore wall and then forms breaks up into zoospores. (3) the conidium by internal division
Key
to Families of Peronosporales
Conidiophores, short, thick, subepidermal, conidia catenulate

Conidiophores, longer, superficial, simple or branched, conidia not catenulate
1-
Albuginaceae,
p. 78.
2.
Peronosporaceae,
p. 82.
Albuginaceae
There is a single genus. Albugo (Persoon) Roussell. This genus of about fifteen species is entirely parasitic upon flowering plants.
THE FUXGl WHICH CAUSE PLANT DISEASE
79
causing the "white rusts." The conidia are borne in white bUstcr-Uke sori under the raised and finally ruptured epidermis of the host. The conidiophores are short, club-shaped, arranged
Fig. 50.
Albugo. A, section through a sorus showing epidermis, conidia, conidiophores and mycelium; B, conidiophores and conidia; C, mycelium and haustoria. After Bergen and Davis.
alusters; the spores are borne in basipetal succession and remain attached in rather long chains unless disturbed. The mycelium is very fine, intercellular and penetrates the cells by globular haustoria. The rudimentary oogonium is multinucleate and filled with uniform proto-
in
As the oogonium grows older plasm. the protoplasm within differentiates

two parts, the inner part of dense protoplasm, the oosphere, and the outer part less dense, the periplasm.^^
into
Figs. 51, 53, 54. During this process the nuclei enlarge, undergo one or two
mitoses,
all
Fig. 54,
and
in
some
species
Fig.
the nuclei except one pass to the In other the periplasm. species
multmucleate at maturity. After discharging sperms. Stevens. The latter type is fertilized by numerous nuclei from the antheridium, the former by a single nu^cleus.^^'^'*' After fertilization the oosphere matures to an
oosphere
is
..1
51. Multiple fertilization

Antheridial tube
ij.'
in A. bliti.
oospore.
80
The globular oospores fall into two classes; ^^ first tuberculate or ridged; second, reticulated. These are illustrated in
Fig. 52.
aw.w.
07.
Fig. 52.
Oospores
4.
panduranae.
platensis.
9.
of Albugo. 1. A. Candida. 2. A. tropica. 3. A. ipomoesDA. Icpigoni. 5. A. swertia;. 6. A. tragopogonis. 7. A. bliti. 8. A. A. occidentalis. 10. A. portulacae. After Wilson.
The conidia in germination usually produce several ovate zoospores with two unequal, lateral cilia. After a brief period of motility they became walled and produced germ tubes capable of
infecting susceptible hosts.
The oospores
after a period of rest

germinate in a similar manner.
81
Conidia germinate freely only if they are chilled.^" A. Candida (Pers.) Roussel. '^ Sori on all parts of the host except the roots, white or rarely light-yellow, prominent and rather deepseated, variable in size
and shape, often confluent and frequently
producing marked distortion of the host; conidiophores hyaline, clavate, about 35^0 x 15-17 n; conidia, globular, hyaline, with
/x; oospores, much less common than conidia, usually confined to stems and fruits, chocolate-colored.
uniformly thin walls, 15-18
Fig.
54.a. bliti. showing differ-
entiation
of
ooplasm
Fig.
bliti, young oogonium and antheridium show-
and
Fig. 55.
53. A.
periplasm, the nuclei in mitos is.
A. bliti, antheridium showing the multinucleate
After
ing nuclei.
After Stevens.
Stevens.
tube. vens.
After
Ste-
40-55 fjL; epispore thick, verrucose, or with low blunt ridges which are often confluent and irregularly branched. This is the most widely distributed and most common species
of the genus.
It occurs
of cruciferous hosts,
and often gives
throughout the world on a large number rise to very pronounced
Practically all cultivated crucifers, cabbage, In radish, turnip, etc., are subject to attacks of this fungus. the caper and mignonette are attacked by the same Europe
hypertrophy.
It has been reported in New York on Tropoeolum.'*^ A. ipomoeae-panduranae (Schw.) Sw."*^' ^^" Sori amphigenous or caulicolous, white or light yellow, prominent, superficial, 0.5-
species.
20 mm., rounded, marked distortions
often
confluent
and
frequently
producing
clavate.
of the host; conidiophores hyaline,
82
unequally curved at base, 15 x 30 /z; conidia hyaline; shortcylindric, all alike or the terminal more rounded, 14-20 x 12-18 û;
the
nounced.
equatorial thickening, usually very proOosporic sori separate from the conidial, caulicolous, rarely on petioles, 1-2 x 5-6 cm. or even more, causing marked
membrane with an
distortion; oospores light yellowish-brown, papillate or with irregular, curved ridges.
25-55 n;
epispore
Common
vulaceae,
morning glory, moon causing but little damage.
throughout the world on various species of Convolflower, sweet potato, etc., although
A. occidentalis G. W. W., reported by has been collected but once.
Pammel
''
on the beet
bliti (Biv.)
A. portulaceae (D. C.) Kze. on purslane ^^ and A. Kze."*'- occur on Amaranthus and related plants.
A. tragopogonis (D. C.) S. F. G.^^'
'^^
caulicolous, prominent, deep-seated, white verulent, rounded or elongate, 1-3 x 1-8
Sori hypophyllous or or yellowish, pul-
mm;
conidiophores
/x;
hyaUne, clavate, about 12-15 x 40-50

less
fx;
conidia, 12-15 x 18-22
light yellow or hyaline, short-cylindric, the terminal larger
and
angular than the others, membrane with an equatorial thickening; oospores produced in stems and leaves, dark brown or almost black at maturity, opaque, 44-68 û, epispore reticulate,
wing bearing papillate tubercles at its angles. cosmopolitan species of less economic importance in America than in Europe attacking a wide range of hosts of the Compositae. Salsify is the chief economic host.
areolae 2 n;
Peronosporaceae
(p. 78)
The members of this family, producing the diseases commonly known as the "downy mildews," have been long known and much studied. They contain many important plant pathogens. The
globular oospores are in general indistinguishable from those of the Albuginacese but the conidiophores are quite different from
In those of that family, being aerial instead of subepidemal. most cases they are branching and tree-like, Fig. 63, but in a
few genera they are short. The oospore in such genera as have been studied (Peronospora ^^ Sclerospora ^^) is formed as in Albugo resulting when mature in an uninucleate egg surrounded by a
83
periplasm bearing the degenerate supernumerary nuclei. Ferhave an uninucleate tilization is as in the Albugos that
ggg
41, 44, 52, 53
The family has suffered many revisions of classification and much renaming of genera. Plasmopara and Peronospora are
especially rich in a
masquerade
of names."*^'
'^*'^^'
Key
to Genera of Peronosporaceae
;
Conidiophores scorpioid-cymosely branched

conidia germinating
by zoospores
or
1.
Phytophthora,
p. 84.
Conidiophores
monopodially dichotomously branched.

simple,
Conidiophores simple or monopodially
branched; conidia germinating by zoospores or by a plasma

Conidiophores branched
simple
or
irregularly
2.
Kawakamia,
p. 89.
Conidiophores regularly branched Conidiophores with the main axis

indurate, the
lateral
branches
3.
reduced and basidia-like
Basidiophora,
p. 89.
main the with Conidiophores axis not indurate, the lateral branches developed normally.
Conidiophores
sparingly
fugacious,
stout,
branched; permanently united

wall of the oogonium
oospore to the
4.
Sclerospora, p. 89.
Conidiophores persistent, slender, usually freely branched; oospore free from the wall of
Branches Branches
the oogonium of the
conidiophore
5.
apically obtuse
Plasmopara,
p. 90.
the conidiophore apically acute

of
6.
Peronoplasmopara,p.93.
Conidiophores dichotomously branched; conidia germinating by a germ tube. Conidiophores with subapical disk-like enlargements from which the ul-
84

timate branchlets arise radially;
germ tube produced from the apex

of the conidia
7.
Bremia,
p. 95.
Conidiophores without subapical enlargements;

conidia
germinating
8.
from the side
Peronospora,
83)
p. 95.
Phytophthora de Bary
(p.
This genus is of especial interest on account of its one exceedingly destructive representative, P. infestans, which occupies an historic position in phytopathology as one of the earliest of parafungi to receive study in any way complete or adequate; study moreover which did much to turn attention and interest
sitic
toward plant pathology.
A distinctive character is that the conidiophores have irregular thickenings below the apparently lateral conidia. The conidiophore is at first simple and bears a single apical conidium, after the production of which a lateral branch arises below the conidium
and grows on
appearance.
in
such a
way
as to give the
is,
first
conidium a
lateral
This process
in
some
species, repeated until a
large scorpioid
cyme is produced. The genus contains seven or eight species, all parasitic.
is much branched, non-septate, the conidiophores arise singly or in hyaline; groups from the stomata, or break through
The mycelium
the epidermis; conidia oval, papillate; zoospores oval, biciliate, escaping by rupture of the papilla; oospores, when present, with the epispore more or less ridged.
Fig. 56.
tails
branched *below, apparently simple above but really one to many times cymosely conidia oval or elliptic, papillate, 35-50 x 20-24 n; branched; germination by about fifteen zoospores. Oogonia in the seed
of P.
Structural de
P. phaseoli Thax.^^"^^ Mycelium well developed, intracellular; conidiophores single or in clusters from the stomata, simple or
phaseoli
After Thaxter.
coats or cotyledons of seeds, rarely in the pods, thin walled, slightly folded; subspherical 23-28 n; oospores spherical or
85
86

smooth, moderately thick walls, hyaline or It was described on lima beans in 1889. ix.
-
subspherical with
light yellow,
18-26
The methods
of infection
were studied by Sturgis
^"
who showed
that spores are carried to the basal portion of the style and ovary by visiting insects. Oospores were described and extensive artifirst grew the on corn-meal-agar, and other media, on which oospores were produced in abundance. The species is unique within the genus on account of the single conidia which are borne at the apex of apparently simple conidiophores but subtended by several enlargements of the kind so ficial
culture experiments
made by Clinton ^^ who
fungus successfully in pure culture
characteristic of the genus.
P. infestans (Mont.) de Bary.^"'
^'"'^
^46-148
Mycelium
well developed, probably perennial; conidiophores
single or in groups of 2-4

six to sixteen zoospores.
branched; conidia 27-30 x 15-20
from the stomata; scorpiose-cymosely ix, ovoid, germinating by about
On
tomato,^^ this species

^^,====3.
diseased solanaceous hosts, particularly the potato and is very destructive. It was first described
in
1845 as a Botrytis and
/^j^<~Px\'
C-fe^'"^'
^^s since been the subject

of
!^
many
The
"*
\l^^-i;-^^/7
extensive papers. are conidiophores

sides
\^^^^/^
X'^rS
abundant on the lower

invasion
line.
.\
^"^jl^^^î^v
/^P^^''
-^ /'^.
'/
of infected leaves near the
\^^s>-^'^"Y^
\ \?T
The myce-
"'^^
/ll3^-
\mm
^^^^^
^"
^^:
Fig.
58.
V
\
of
migrates between the piercing them with

existence of oospores
'^^
haustoria.
^%i^^ji^ '/\ \ and mature oospores Young

After Clinton.
^""^^
The
is
a
x
mUch
j.i,
Controverted
structures
j. j.
p. phaseoli.
pomt;
the
re-
as oospores probably belonged to some other Jones ^^ found peculiar thick- walled bodies, fungus. Recently somewhat resembUng oospores, in undoubtedly pure cultures of
ported by Smith
P. infestans.
are oospores is not known. Clinton that he, in pure cultures, has obtained "absolutely perfect oogonia, antheridia and even oospores-" The
Whether they
^^
has recently announced
THE FUXGI WHICH CAUSE PLANT DISEASE

oval, flattened biciliate zoospores
87
swim about, come

tube.
which emerge from the conidia, to rest, develop a wall, then produce a germ Direct germination by a germ tube also occurs rarely. In-
FiG.
59.
P.
infostans;
5,
formation;
section showing conidiophores aiul 1, germination of a conidia. After Scribner.
conidia-
is brought about by the germ tube, either by penetrating through stomata or directly through the epidermis. The walls and contents of parasitized cells are browned. When this fungus is alone on the tubers dry rot is induced, but invasion
fection
of
into a disagreeable
numerous saprophytic fungi and bacteria usually turns this wet rot. Tuber infection occurs largely from
88

soil
conidia washed into the
by
rain; possibly
sometimes by the
mycelium migrating by
way
of the stem.
The fungus was extensively studied by Jones in pure culture and a decided difference in luxuriance of growth was observed on blocks cut from different varieties of potatoes, Fig. 57. The mode of hibernation is not thoroughly known but undoubtedly hibernation occurs in part in live mycelium in infected tubers. The conidia are short-lived, especially when dry.
P. omnivora de Bary. Conidiophores simple or branched; conidia ovoid or lemon-shaped, 50-60 or even 90 x 35-40 fx, germinating by as many as fiftj'^ zoospores; oospores smoothish or
wrinkled, light-brown, transparent, 24-30 /x. This species which includes forms previously described as P. cactorum (Lebert &
Cohn) Schr., P. fagi Hartig, and P. sempervivi Schenk is found upon seedlings of some fifteen families ranging from Pinacse to the It is of considerable economic importance higher Angiosperms.
in
Europe
especially in the seed beds of the forester.
Recently it been found on ginseng in Japan and the United States.^^

has
The same fungus

destructive
"^^
is
credited with
of
rotting
apples
'^^
and pears in Europe and with two wide-spread tropical causing diseases, the cocoa pod rot and a palm disease. From the studies of de Bary ^^ and from the nature of the more recent outbreaks credited to this fungus it appears that P. omnivora is a composite species
Fig. 60. Formation of swarm-spore of Phytophthora. After Smith.
which
gated.
will
eventually
be
segre-
Coleman " already been begun. of India as P. omnivora var. arecae while Maubfecting fungus lanc^^ has gone further and described the cocoa disease as P. faberi. See also " ^^"
species,
lilac in
segregation has has described the palm in-
Indeed
P. syringae recently described by Klebahn is a closely related which is very destructive in the propagating beds of the
Germany.
THE FUNGI WHICH CAUSE

P. agaves Gan.^^ occurs on the P. nicotiana v. B. d H.^
is
PLAX'I'
DISEASE
89
Agave
in
Mexico.
but culture work shows

host as
it
it
also closely related to P. omnivora, to be rather fastidious in its choice of
attacks only tobacco seedlings. P. calocasiae Rac. occurs on Calocasia antiquorum in the Orient.
An
undescribed species on Castor
is
also reported.**^
Kawakamia Miyabi
Mycelium
in
(p.
83)
slender, copiously branched; conidiophores single or
more from the stomata, simple or sometimes branched, but branches never arising near the conidia. irregularly Conidia usually upon a slender pedicel cell, lemon-shaped, obgroups
of 2-5 or
tusely tipped, contents and wall colorless, germination normally by zoospores; zoospores oval, flattened and laterally biciliate;
oospores spherical, smooth.
A single species, K. cyperi (M. & I.) Miyabe,^^ which was introduced from Japan into Texas in imported plants of a sedge, Cyperus tegetiformis. The species is very destructive in Japan. Both conidia and oospores were produced in the Texan material.^Basidiophora Roze
&
Cornu
(p.
83)
culti-
B. entospora R. & C. occurs on species of Erigeron and vated aster in Europe and America.
Sclerospora Schroter
(p.
83)
This genus ponderance of
all other Peronosporales in the prethese are the conspicuous stage, while oospores; the conidiophores and conidia are few, small and evanescent. There are about five species.
differs
from
its
Mycelium much branched, with small

conidiophores
fugaceous,
erect,
vesicular
haustoria;
solitary
low and
groups of two or three, stocky, sparsely branched, the branches

or
in
also stocky; conidia elliptic or globose-elliptic, hyaline,
smooth;
oospores globose, intramycelial, the epispore broA\Ti, irregularly wrinkled, permanently united to the persistent wall of the oogo-
nium.
90

S. graminicola (Sacc.) Schr.,''-'
''^
infectsleaves and inflorescences,
the oospores causing marked distortion of the latter and rapid disintegration of the former; conidiophores 100 x 10-12 (x, conidia
20 X 15-18 m; oogonium wall thick, 4-12 n, at maturity 30-60 /z in diameter, reddish-brown; oospore pale-brown, 26-36 fx. The conidial phase is not prominent, while the oospores by their
disintegrating effect upon the leaves of the host, render the plants quite conspicuous and closely simulate the habit of a brown smut.
Fig. 61.
S. graminicola.
Conidiophores and conidia germinating

;
Fig. 62.
S. Kraniinicola,
oo-
conidia
spores.
zooAfter Butler.
and
gonium, oospore and antheridium in section. After Stevens.
On
millet (Setaria italica), pearl millet, fox tail
and corn;
tassels
in India
economic importance.^^ S. macrospora Sacc. has been reported in wheat in Italy and the United States.^^' ^^
of considerable
com
and on
Conidia unknown; embedded firmly in the tissue of the host, not causing oogonia
fi.
disintegration as in S. graminicola; oospores light yellow, smooth,
60-65
Plasmopara.
Schroter
(p.
83)
134
The
this
tree-like,
branching conidiophores, Fig. 63, are
common
to
genus, Peronospora, Peronoplasmopara and Bremia, and unlike the conidiophores of Phytophthora they are completely formed before they begin to bear spores.
Mycelium branched; haustoria
simple;
conidiophores
erect.

solitary or fasciculate,
91
from the stomata
of the host,
monopodi-
ally branched, the branches arising at right angles to the main axis, as do also the secondary branches (at least never appearing
truly dichotomous) the ultimate branches apically obtuse; conidia globose to ovoid, hyaline or smoky, germinating by zoospores or
the entire protoplasmic mass escaping and then sending out a
germ tube; oospore globose yellowish-brown, the epispore variously wrinkled sometimes appearing somewhat reticulate; oogonium persistent, but free from the oospore. ^^' "^' '''' ''' first collected in P. viticola (B. & C.) B. & d T.,''1834 by Schweinitz and regarded as a Botrytis was first published in 18515 Hypophyllous, caulicolous, or on young fruits, covering the
infected areas with a white
;
downy growth; on
the leaves epiphyl-
lous discoloration yellowish on the
fruit often causing
a brown rot
without producing conidia; conidiophores fasciculate, 250-850 x 5-8 fi, 4-5 times branched, the ultimate branchlets about 8 fi long; conidia ovate-elliptic, very variable in size, 9-12 x 12-30 n;
oospores 30-35
epispore brown, wrinkled, or almost smooth; oogonium thin-walled, hyaline or light yellowish-brown. The mycelium is found in all diseased tissues except the xylem.
ju,
The conidiophores
issue
from stomata.
The
conidia germinate
readily in water, producing in about three-fourths of an hour biciliate zoospores. These after fifteen to twenty minutes activity
cease motion, round off, become walled, then germinate by a tube. This bores through the epidermis and develops into the internal mycelium. Infection is almost exclusively from the lower side of
Oospores are much more rare than conidia but are often found in autumn, sometimes two hundred to a square millimeter of
the leaf .^^
Though hibernation is doubtless chiefly by oospores has been shown that the mycelium can perennate in old wood, and even form oospores therein. The fungus is dependent on
leaf surface.
it
abundant moisture.
P. nivea (Ung.) Schr. attacks various species of umbellifers including the parsnip and carrot. It has been reported in America only from the region of San Francisco. P. halstedii (Farl.) B. & d T. This form is quite variable and should perhaps be separated
92
into several distinct species.
It is limited to the Compositse, Helianthus and Madia being the only hosts of economic impor-
tance.
Hypophyllous; conidiophores fasciculate, slender, 300-750 m, 3-5 times branched, ultimate branchlets 8-15 fx long, verticillate
Fig. 63. P. viticola. A, seotion of a leaf with conidiophores emerging from a stoma; C, formation of swarm spores; D, formation of oospores. After Millardet.
below the apex of the branching axis which is frequently swollen and ganglion-like; conidia oval or elliptic, 18-30 x 14-25 /x; oospores 30-32 ix, epispore yellowish-brown, somewhat wrinkled. P. ribicola (Schr.) Schr. grows on various species of currants in Europe and America but is probably of but slight economic
importance.

P. obducens (Schr.)
93
Sclir. occurs on Imputiens, both wild and North America, Europe and Asia. P. pygmea (Ung.) Schr. on various Kanunculacese, including Aconitum in Europe and cultivated Hepaticas in America,^'' is
cultivated, in
of little
Peronoplasmopara
(Berlese) Clinton (p. 83)
There are three species which have been variously designated as Peronospora, Plasmopara, Pscudoplasmopara and Peronoplasmopara. The genus combines colored conidia and zoosporic germination with a type of conidiophores intermediate between those of
Peronospora and Plasmopara.

small, usually simple; conidiophores pseudo-monopodially branched, the ultimate branchlets acute, the primary arising at acute angles; conidia colored,
elliptic,
Mycelium much branched, haustoria
thin-walled,
conspicuously papillate both apically and basally; oospores smooth or roughened; oogonium thin-walled. P. celtidis (Waite) Cl.^^ is unique in the family as the only
species infecting dicotyledonous trees. It occurs on hackberry in the region about Chesapeake Bay, also in Japan. P. humuli Miy. & Taka ^- causes a serious hop disease in Japan.
It
has recently been found by Davis P. cubensis (B. & C), Cl.^^'^^' ^^^
^^
on wild hops
in Wisconsin.
Hypophyllous, rarely amphigenous; discoloration of the host yellowish, or water-soaked; conidiophores 1-2 rarely more from a
stoma, 180-400 x 5-9
fi,
3-4, rarely 2-5 times branched, the ultifj.
mate branchlets recurved;

14-25
fi;
apically acute, 5-20 long; conidia gray, brownish or smoky, ovoid to ellipsoid, papillate, 20-40 x
oospores spherical, yellowish, warty-papillate, 30-43

in the
fi,
decaying leaves. The mj^celium abounds in the spongy parenchyma. The conidiophores emerge through stomata, or rarely directly through the cuticle, near the invasion line of the fungus. Fresh conidia
maturing
germinate in water in two to four hours forming flatfish zoospores with one anterior and one posterior cilium. The zoospores later
become spherical, walled and develop a germ tube. These germ tubes enter the host through the stomata or directly through the cuticle from cither above or below. Moist weather is favorable to
94
the fungus in that conidia are produced more abundantly and retain their power of germination longer when moist. Disease
spots appear two or three days after infection; conidia or ten days after infection.
same nine
The
species
is
perennial in Florida
^'^
and spreads northward as
the season advances, reaching Ohio and
New York by late summer
Fig. 64. P. cubensis: .3. Conidiophore wjth young and old conidia. 5. Conidium. 6. Conidium germinating. 18. Infection through 11. Zoospores. a stoma. After Clinton.
or early autumn.^^ For a series of years after its discovery it was not well known even scientifically, its first serious outbreak being
and appeared in Japan about the same time The oospores have is now known to be almost cosmopolitan. been found only by Rostewzew and have not been seen in America.
about
1889.^^
It
wide range of wild and cultivated cucurbits
is
infected,
among
THE FUNGI WHICH CAUSE PLAXT DISEASE
95
them the pumpkin, squash, cucumber, muskmelon, watermelon, ^"^ gourd, in fact according to the work of Selby any cucurbit apUable to attack. CHnton infected muskmelons with spores pears
produced on cucumber. The fungus cucumbers raised under glass.
is
especially prevalent on
Bremia Kegel
(p.
84)
As in Peronospora except that just below the ends of the conidiophore branches there are pronounced swellings from which spring radially a number of short branches each
The bearing an ovate, papillate conidium. conidia germinate by apical germ tubes.
There
is only one species. B. lactucae Kegel is found on lettuce and several other Compositse.^^ It is more in-
jurious in
Europe than in America. Hypophyllous or amphigenous,
causing
fig.
discoloration, then wilting of the host; conidiophores produced singly but in great abun-
65. B. lactucse. ^^^^^ Tubeuf.

fx;
dance,
much branched;
fi,
small, 26-35
light
conidia ovate, 16-22 x 15-20 brown, the epispore wrinkled.

52
oospores
Peronospora Corda
(p.
84)
This genus of some sixty species contains several aggressive Its conidiophores are much like those of Plasmospara parasites. but with more tendency to dichotomous branching and to more
filiform, simple or branched; conidiophores dichotomously 2-10 times branched at acute angles, ultimate branchlets acute, more or less reflexed;
graceful habit; the apices are acute. Mycelium well developed, haustoria
conidia hyaline or colored, papillate, germinating directly
by lateral
germ tubes; oospores globose,

smooth.
P. parasitica (Pers.)
reticulate, tuberculate, wrinkled or
De
Bary.^^
This
is
often associated with
Albugo Candida, giving it the appearance of a parasite on that fungus. Almost all species of Cruciferae are subject to attack,
among them cabbage,
cauliflower, radish, collards, turnips, horse-
96
radish,

and others
of
minor economic importance.
It
is
cos-
mopoHtan in distribution. The fungus covers any green part
of the host with a dense white
growth, often causing hypertrophy especially in oospore formation; conidiophores 200-300 x 10-12 ii, bushy branched, stout,
deliquescent,
with 5-8 main

branchlets
branches,
slender,
each from
3-7 times
curved, usually arising at acute angles, about 12-15 x 2-3 ju; conidia broadly elliptic, bluntish, often becoming globose, about 12-22 x
branched,
ultimate
more
or
less
24-27
;u,
hyaline
or
globose,
less.
yellow-brown,
very light; oospore 26-45 ix, epispore

thick, color-
smooth or wrinkled; oogonium
P. efifusa (Grev.) Rab. causes a serious disease of spinach. ^"^ It also occurs on a wide range of weeds of the Chenopodiaceae.
The
all
species was formerly made to include the effusae forms of the genus so that
literature
abounds with references to
it
on
Viola, Plantago,
Polygonum,
etc.
Hypophyllous, causing yellowish or brownof
mass of conidiophores a violet cast; conidiophores 150-400 x 7-9 ju, much branched, the ultimate branches
ish discolorations, the
at right angles, usually recurved, 8-15 x 3-4 ii] conidia ellipsoid to globose 17-18 x
P. cnusa on Fig. G6. After Halspinach.

^ ^
22-24 30-40
less
jx,
violet or
i i
u,
epispore
smoky; oospores globose, light brown, more or
regularly
wrmkled;
oogonmm thm,
brown.
was first described as aBotrytis in 1841. was noted in America in 1872 by Taylor,^*^^ later by Trelease ^^ and by many others. ^^ A very complete description was given ^^^ in 1904 under the name P. schleideniana. by Whetzel The conidia in mass present a purplish tint. The conidioP. schleideni Ung.^^
It
phores usually emerge singly through the stomata. The slender, branched haustoria abound in the parasitized part often with their ends wrapped around the nuclei. In water the conidia
97
germinate directly to form an infective tube (Fig. G7) which grows into the stomata. According to Whetzel conidia retain their germinating power only a few hours. Shipley believed them
viable for a
much
longer
time.'"''
Fertilization occurs
much
as in
P. parasitica (Fig.
G7) and the sexual spores, which abound,
serve for hibernation.
The}^
may
live several years.

etc.
It is found on onion, garlic, (Allium sps.) everywhere, covering leaves with a dense growth; conidiophores, 3-6 times branched, 300-700 x 12-15 fx; branches 2-5, scattered, ultimate
branchlets subulate, 15-20 /x, more or less recurved; conidia large, obovate to pyriform, basally papillate, 45-58 x 20-25 ju, the membrane violet; oospore globose, light-brown, about 30 fi,
epispore smooth or slightly wrinkled. P. sparsa Berk, is parasitic on roses
pest in Europe,
and constitutes a serious though not so common in America. Hypophyllous, with a whitish growth; conidiophores about
^^
9 times branched, the ultimate branchlets refiexed; conidia subpale gray. P. trifoliorum de Bary. Hypophyllous, forming a dense grayish or dirty- white growth over the host; conidiophores slender, 360elliptic,
600 X 9-11 fi, 6-8 times branched at acute angles, the primary branches rather erect, the secondary more spreading, flexuose, more
or less recurved, ultimate branchlets at right or obtuse angles,
straight, subulate, 7-12 x 7-3 n; conidia globose to broadly elliptic,
15-20 x 18-36 n, violet; oospores globose, 24-30 brown, smooth.

It causes serious loss to clover in
/t,
epispore light
genera also suffer.

in
America by
its
Europe. Species of related has assumed a role of importance Recently attacks upon Alfalfa ^^ on which it occurs from
it
New York
to California.
It differs from P. viciae in the branching of the conidiophores, the lighter color of the spot and fungus, and the smooth oospores. P. viciae Berk. Hypophyllous or caulicolous, covering the host
with a grayish-violet growth, epiphyllous discolorations yellowish or inconspicuous; conidiophores fasciculate, 300-700 x 9-11 n,
5-8 times branched, the main branches arising at acute angles,

erect, the ultimate subequal, slightly flexuose, arising at right or
obtuse angles, the lateral recurved, 10-17 x 2-3 n; conidia
elliptic
98
11
14
Fig. 67. p. schlcideni. 11. Mycelial threads between the large conductive cells of the leaf; (a) the mycelial thread; (b, b) branched or coiled haustoria; (c) branched haustorium wrapped about the nucleus. 1.3. Young conidiophorcs, (a, a) turning toward the stoma, (b) (c) haustorium wrapped about the nucleus of the 14. Mature conidiophore (a) with mature conidia, (c, c) epidermal cell. 15. Oospores, (a) mature oospore (d) germ tube of conidium entering stoma. with old antheridiiim, (d) still attached; (b) mature oospore still inclosed in the After Whetzel. old wall of the oogonium.
;
;
99
or obovoid, 15-20 x 21-28 n, light-violet; oospores small, 25-30 n,
epispore yellowish-brown, with low, broad reticulations, areolae
about 8 ju; oogonium thin, fugaceous, 32-40 /x. This fungus on ^'i('ia and related genera is sometimes quite
Fig. 68.
A sporangium with a columella (Mucor).

After Sachs.
serious,
particularly
on vetch and peas
in
Europe, Asia and
America.
P. violae de Bary; on cultivated violets and the pansy in Europe and America,^^ forming discolored spots; foUicolous or caulicolous,
with a pale violet growth, conidiophores fasciculate, short, 2-7 times dichotomously branched; ultimate branchlets short, subulate, reflexed; conidia elliptic, short, apiculate, 20-22 x 15-18 ti,
violet.
100
P. dipsaci Tul. on teasel and Scabiosa in Europe and America and P. violacea Berk, on the flowers of species of Scabiosa in Europe
kills
are quite distinct from the preceding; P. schachtii Fcl. on beets seedlings in Europe. P. linarias Fcl. is on digitalis; P. cytisi
Fig. 69.
Sporophores
Brefeld,
in the
Zygomycptes.
After
De
Bary,
Cunningham, Schroter.
on species of Cytisus in Europe; P. arborescens (Berk.) de Bary on poppies, especially garden seedlings, in Europe and
Rost.^^''
''-
Asia.
Species of less importance are: P. rubi Rab. on various species of
Rubus
in
Europe and
America; P. fragariae R. & C, usually cited as a synonym of P. potentillse de Bary, on the strawberry in France and America;
P. trichomata Mas.^^'^' "^ the cause of a root rot of Colocasia
THE FU\GI WHICH CAUSE PLAXT DISEASE

in the
101
West
and upon non-economic Primulaceae
Indies; P. Candida Fcl. on the primrose in Europe in America; P. maydis Rac.^^^

[Its
the cause of a disease of corn in Java.
identity with Sclero-
spora graminicola is suggested by the recent studies of that species by Butler.] P. vincae Schr. on Vinca minor in Europe; P. myosotidis
de Bary on several species
of
forget-me-not
and
related
;
genera in Europe and America P. cannabina Otth. on hemp in
y<^
Europe and Japan; P. conglomerata Fcl. upon alfilaria in Europe; P. ficariae Tul. on various species of Ranunculus both in the old and the new world; P. antirrhini Schr. on the snapdragon and related
hosts in Europe; P. nicotianae Speg.^^^ on various ornamental
species of Nicotiana in South Fig. 70. Mucor: zygospore formation.
America and California; P. valeria mellae Fcl. in Europe on Valerianella; P. Valerianae Trail on Valerian; P. dianthi de Bary on species of Dianthus in Europe; P. corallae Tranz. on Campanula in Europe; P. jaapiana "^ on rhubarb in Europe; P. phoenixae Tap. on Phcenix ^^'^ and an undetermined species on Para rubber. Mycelophagus castaneae Man,^^^ is an imperfectly described form which may belong either to the present group or to the A serious disease of the chestnut in France is Chytridiales.
charged to
it.
'^"'^^'"
^"'^'^'^
Zygomycetes
This group of fungi
its
(p.
66)
is readily distinguished from the Oomycetes sexual organs, when these are present. In the isogamous by absence of sexual organs the general type of sporangium is usually
sufficient
mark
of distinction for those
who
are even but slightly
acquainted with the two groups. Tho mycelium, if young, serves to indicate relationship to the Phycomycetes. Older mycelium is
often septate and would lead the
unwary into
errors of classification.
102

in sporangia or are borne as conidia. usually with a columella. The spore-bearing
Asexual spores are either
The sporangium
ing, Fig. 69.
is
stalks exhibit the widest diversity in shape
and form
of branch-
Sexual spores (zygotes) are produced through the union of two gametangia. (Fig. 70.) Though the cytology of zygote formation has not been completely studied it seems clear that the
like
is multi-nucleate as in Albugo two uniting elements are ccenogametes.
fertilization
^^"^
bliti
and that the
Key
to Orders of Zygomycetes
Asexual spores borne in sporangia which in some genera are reduced to

conidia-like bodies
1.
Mucorales,
p. 102.
Asexual spores true conidia borne singly

at the apex of the conidiophores
.
...
2.
Entomophthorales,
p. 107.
Mucorales
This order
is
(p. 66)
genera and one hundred
comprised mainly of saprophytes, about twenty fifty species; but includes a few forms which prey upon vegetation in a very
low ebb
of
life,
as cells of ripe fruit,
species which are of especial interest as they grow upon other fungi. The sporangial
tubers, etc.,
and a few
stage
is exceedingly common; the zygosporic much less so, very rare in ^-" the case of some species. Blakeslee
has
Fig.
shown
that
in
some
species,
71. Phypomycetes .showing zygosporic lines at regions of contact between + and strains. After Blakeslee.
though the two uniting sexual organs are to all appearances alike, the plants
are
m
.
reality dioecious; that a
,.
,.
branch
^^^^ ^^^
^j.^^^ ^,^^^^^ produce sexual
organs that will unite with other sexual organs produced upon the same plant. IVIoreover, there appears to be a differentiation of sex in that one plant,
which
may
provisionally be re-
103
garclcd as the male, unites freely with another plant, provisionally the female, but this male plant refuses to unite with any other
plant which is capable of uniting with the female and all plants that can unite with the male refuse to unite with the females. In
some
tive
species the plants of one sex
show a more luxuriant vegeta-
growth than do plants
of the other sex.
Key
to Families of Mucorales.
Asexual spores in typical sporangia, although

in some genera few-spored Sporangium with columella; zygospores
nakctl or thinly covered with outgrowths of the suspensor
1.
Mucoraceae,
p. 103.
Sporangium without a columella; zygospores closely covered by hyphae ... Asexual sporangia monosporic and conidialike, sometimes accompanied by larger
polysporic sporangia Sporangia of two kinds, polysporic and
2.
Mortierellaceae.
monosporic
Sporangia
3.
Choanephoraceae,
Chaetocladiaceae.
p. 106.
monosporic; parasitic on other genera of Mucorales

all
.
4. 5.
Sporangia simulating chains of conidia.
Piptocephalidaceae.
Of these families the second and

while the fourth
is
fifth
parasitic
upon other members
are pure saprophytes, of the order.
Mucoraceae
Mycelial threads all alike or of two kinds, one aerial, the other buried in the substratum, coenocytic during growth but septate at maturity; reproduction by asexual spores borne in sporangia
and
])y
zygospores formed by the union of equal gametes; spor-
angiophores, simple or branched; sporangia variable, typically with a columella, and many spores but in some genera some of
the sporangia are few-spored and without columellas; zygospores variable, smooth or spiny, borne on short branches of the myce-
Hum.
104

Key to Subfamilies and Genera
of Mucoraceae
Sporangial
membrane cuticularized and permanent above, thin and fugaceous be-
low Subfamily I. Piloboleae. Sporangiophore of equal size throughout; Pilaira. spore mass not forcibly discharged
.
. .
Sporangiophore swollen beneath the spo-
rangium;
spore
mass
thin
forcibly
dis-
charged at maturity
Sporangial membrane
antl
Pilobolus, p. 105.
fugaceous
II.
throughout
Sporangia
all
similar
Subfamily
Mucoreae.
Mycelium
gion Aerial
differentiated into a colorless
vegetative and a colored aerial re-
mycelium
stoloniferous, zygoth(>
spores formed in
substratum
1.
Sporangiophores arising from the nodes
Rhizopus,
Absidia.
p. 105.
Sporangiophores arising from the

internodes
Aerial
2.
mycelium
not
stoloniferous;
zygospores aerial
Sporangiophores simple dichotomously Sporangiophores branched
3.
Spinellus.
4.
Syzygites.
Mycelium imdiif erent iated Mycelium gray or brown; suspensors

smooth
Sporangiophores simple Sporangiophores variously branched Sporangia borne apically on the
sporangiophore branches
antl
its
5.
Mucor,
p. 106-
Zygospores formed from equal
gametes Zygospores formed from

equal gametes
0.
Calyptromyces.
un7.
Zygorhynchus.
Sporangia
only on the lateral, circinate branches of the sporangiophore
borne
THE FUNGI WHICH CAUSE PLAXT DISEASE

Sporangia globular; columolla not constricted
Sporangia pear-shaped mella constricted
;
105
8.
Circinella.
colu9.
Pirella.
Mycelium
Sporangia
spored
of
metallic; suspensors spiny 10.
Phycomyces.
two
kinds,
many-spored;
the
primary secondary fewSubfamily
the
III.
Thamnidieae.
Pilobolus crystallinus (Wigg.) Tode, a form with beautiful crystalline sporangia on yellowish, evanescent sporangiophores has
been frequently noted as injuring or smudging chrysanthemum, i'-i"i-rose and other leaves by its profuse discharge of spoIt is not, however, a parasite. rangia.
Of the other genera the only ones
of interest regarding plant
disease are Rhizopus and Mucor. The others are saprophytes found on a great variety of substances, manure, fungi, and many other kinds of organic matter,
Rhizopus Ehrenberg
(p.
104)
The sporangium
sporangia are
all
vâll
of
is not cutinized, and falls away. The one kind and with columellas. The sporan-
giophore
;
is
never
dichoto-
mous zygotes are found in the mycelium. The suspensor is

without outgrowths.
rophytes.
Aerial R. nigricans Ehr. at maturity chocomycelium
''
.
Twelve
or fifteen species, chiefly sap-
^ Fig.
late-colored;
rhizoids
1
numerp
ous; sporangiophores lasciculate, erect,
Diagram showing mycelium and sporophorcs. After Coulter, Barnes and Cowles.
72.
_, Rhizopus.
.
_^.
aseptate; sporangia globose, l)lackish-olive, granular; columella hemispheric; spores gray to brown, subglobose or irregular, 11-14 n; zygospore 150-200 n, epispore with rounded warts, black. This is the cause of soft rot of stored vegetal)les, particularly of sweet potatoes,^-âlso of Irish potatoes,^^^ apples
it
causes death of squash blossoms
^-^
and
is
and pears; destructive to barley
106
during malting. It is distinctly a wound parasite and is unable to force entrance through a sound epidermis. The richly branched mycelium which varies from very thin and hyaline to thick, coarse and slightly fuscous, is found throughout
the rotten portion of the hast.
tive
After a period of luxuriant vegetaair,
growth hyphae protrude to the
first
through existing
ruptures in the epidermis, later by rifts forced by the fungus itself. Sporangiophores then form in dense bush-like growths,
each sporangiophore bearing one terminal sporangium. The sporangia are at first white, later black and contain very numerous ^-^ spores. Spore formation has been closely studied by Swingle.
Aerial stolon-like liyphge reach out in various directions and at their points of contact with some solid develop holdfasts (Fig. 72)
and a new
in Fig. 70.
cluster of sporangiophores.
Zygotes are produced
by union
of
two mycelial
tips as
is
shown
Orton
^-^
Irish potato
inoculated pure cultures of this fungus on sterile raw and induced typical decay. He also noted that there
was a
difference in the rate of decay produced by strains of Rhizopus derived from different sources and that the most rapid decay of potatoes was caused by strains taken from rotting potatoes. R. necans Mas.^-" causes decay of lily bulbs in Japan. R. schizans Mas. is cited as the cause of split-stone in peach. ^-^
Mucor
Mycelium
all of
Linnaeus
(p.
104)
one
kind, buried in the
substratum or grow-
ing over its surface; sporangiophores scattered or not, simple or branched; sporangia globose; columella cylindric, pyriform or clavate; spores numerous, variable; zygospores globose, smooth or
warty.
Some thirty species, chiefly saprophytes. M. mucedo L. is destructive to beech nuts
in winter.
M.
fruits.
pyriformis Fisch and
M. racemosus
Fes. cause decay of
Choanephoraceae
(p.
103)
Mycehum parasitic on living plants; sporangia of two kinds; macrosporangia globose, columella small, spin}', spores few, on
107
simple or branched, erect sporangiophores; microsporangia clavatc, one-spored simulating conidia and borne in heads on the enlarged apices of umbellately branched sporangiophores; zygospores as
in IMucoracae.
single genus, with three species.
A. Alull occur on blossoms
Choanephora infundibulifera (Curry) Sacc. and C. americana hi India and South America.
A third species, C. cucurbitarum (B. & Br.) Thaxter, is the cause of decay of cucurbits especially pumpkins, in the eastern and southern states. ^-^
Entomophthorales
This order
(p.
66)
is predominately one parasitic on insects. Some are known, only four of which are plant parasites. fifty species
Asexual reproduction is chiefly by conidia, apically borne and for the most part forcibly ejected from their stalks at maturity.
Key to
Families of Entomophthorales
.
Endozoic parasites (lusecta, Arachnoidea)
1.
Entomophthoraceae.
Basidiobolaceae,
p. 107.
Endophytic or saprophytic
2.
Basidiobolaceae
This family
is
characterized chiefly
by
its
habitat.
Septa are
numerous
in the vegetative
mycelium.
of Basidiobolaceae
Key to Genera
Intracellular parasites, the
mycelium greatly
1.
reduced
Saprophytes, or parasites on higher fungi, the mycelium well developed.
Completoria,
j).
108.
Conidia produced directly from an un-
swoUen conidiophore.
higher fungi
Parasites on
2.
Conidiobolus.
Basidiobolus.
Conidia cut
off
from the
ajx^x of
a swelling
3.
of the conidiophore.
Saprophytic...
108
of the one species given below these are not on higher plants. parasitic Completoria complens Lohde is parasitic upon fern prothallia.^^ Vegetative body compact, of oval or curved branches in a single host cell, extending to other cells by slender tubes. Resting spores 10 to 20, formed in the host cell. Propagation by non-motile
With the exception
conidia, 15-25
fi,
in diameter.
BIBLIOCillAPHY OF PIIYCOMYCETES *
(pp. 59-108)
1
Atkinson, G. F., Ann.

Stevens, F. L.
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2
3
'
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J. G.,
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8 9
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1"
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12
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IJ,:
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Zimmermann, E., Nat. Zeit f. Forst u. Land. 8: 320, 1910. Salmon, E. S. & Crompton, T. E., Wye Ag. Coll. R. Ec. Myc. 109,
Thomas, Insect
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I.
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"
i
Halsted, B. D., N. " Shear, C. L., B. P.
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Farlow,
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2' 2"
Berlese, A. N., Riv. Path. Veg. 7: 167, 1901.
C. R.
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222, 1894.
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"
2
=^ 2s 2^
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d. Fr. 26:
B.
My.
C. R. 119: 108, 1894. Magnus P. Ann. Bot. 11: 92, 1897. Massee, Bull. Kew Garden, 1906.
'
Magnus, P. Ber. Deutsch. Bot. Ges. 20: 291, 1902. Sydow. Ann. Myc. 1: 517, 1904. " Farlow, W. G., Rhodora 10: 9, 1908.
'1
*See footnote, page

109
53.
110
33 34 35
3

Atkinson, G. F., N. Y. (Cornell) B. 94, 1895. Mass. Agr. Exp. Sta. R. 8: 220, 1890.
Butler, E.
37
38
Miyaki, Ann. Bot. 15: 653, 1901. J., Mem. Dept. Agric. India, Botan. Scr. 15: 86-91, 1907. Halsted, B. D., N. J. R. 13.
J.,
Butler, E.
R. Pusa. 10: 44, 1909.
Smith, E. H. & Smith, R. E., Bot. Gaz. 4S: 215, 1909. Smith, R. E., Cal. B. 190.
39
41
Stevens, F. L., Bot. Gaz. 32: 77, 1901.
Stevens, F. L., Bot. Gaz. 28: 149, 1899. Davis, B. M., Bot. Gaz. 29: 297, 1900.
44
45
Wager, H., Ann. Bot. 10: 295, 1896. Wilson, G. W., Torr. Bull. 84: 61, 1907. 4 Melhus, I. E., Sc. 33: 156, 1911. " Halsted, B. D., N. J. R. 11: 350, 1890. 48 Stewart, F. C., N. Y. (Geneva) B. 328, 1910. 45 Halsted, B. D., N. J. B. 76: 1890.
Pammell, L. H., la. B. 15: 236, 1891. " Halsted, B. D., N. J. R. 15: 355, 1894.
" Ruhland,
53
Diss., 1903.
54
Stevens, F. L., Bot. Gaz. 34: 420, 1902. Wilson, G. W., Torr. Bull. 34: 387, 1907.
"
59
J. Myc. 13: 205, 1907. Clinton, G. P., Ct. R. 329, 1904. " Berlese, A. N., Riv. d. Pat. Vcg. 9:
58
5''
"0
1, 1900; 10: 185, 1902, Bot. Gaz. U: 273, 1889. Thaxter, R., Thaxter, R., Ct. R. (State) Sta. 167, 1899, 1890. Scribner, F. L., D. Agr. R. 337, 1888.
i 82 83
G., N. Y. (Cornell) Bui. 113: 249, 1896. Sturgis, Bot. Gaz. 25: 191, 1898. Clinton, G. P., Ct. R. 278, 1905.
Lodeman, E.
94
65
88
87
Smith, R. E., Cal. B. 175, 1906. Smith, W. G., Card. Chron. 1875.
G., Quar. Jour. Mic. Sc. 15: 1875. G., Diseases of Crops, 1884. Jones, L. R., Sc. 29: 271, 1909. Clinton, G. P., Sc. 33: 746, 1911.
Smith, Smith,
W. W.
88 83 ">
Clinton, G. P., Ct. R. 362, 1904; also R. 304, 1905.
" Whetzel, H. H., Sc. 31: 790, 1910. " Osterwalde, A., C. Bak. 15: 434, 1906. " Bubak, Fr., Zeit. 20: 257, 1910.
BlBLlOGRArilV OF PHVCOMVCETES
^^de Bary, A., Bot. Zeit. 587, 1881.
HI
"
"'
Colenuui, L.
C,
INIaubUinc, L'Agr. Prat. d.
:Mycol. Bull. 2: Dcpt. Agiic. Mysore State, 1910. Pays Cliauds 79: 315, 1909.
Kidley, II. X., Agr. B. Straits tt Fed. Malcy Sts. 10: 70, 1911. " Petch, T., Circ. and Agr. J. Pvuy. Bot. Gard. Ceylon 5: 143, 1910. " Gandary, G., Mem. Y. Rev. See. Cient "Antonio Alzate" 23: 293,
1909.
o
Meded, Lands. Plant. Batavia 15: 1896.

Butler, E. J., Kept. Agr. Research Inst. Pusa 10: 45, 1909-1910. Patterson, F. and Charles V. K., B. P. I. 171: 1910.
'
8'
Kawakamia, a new genus belonging to Peronosporacea; on Cyperus tcgetiformis. With a postscript by Dr. Kingo Miyabe, 1904.
s-"
5
8 8^
Butler, E. J., Mem. Dept. Agric. India, 2: No. 1, 1907. Cugini, G. and Traverso, G. B., Staz. sperim. Agr. Ital. 35: 46, 1903. Peglion, C. Bale. 28: 580, 1910.
Berkeley,
J.,
Hort. Soc. Lond. 6: 289, 1851.

u. F.
8 83
Dept. Agr. R. 96, 1886.
Appel
Waite,
&
Stewart, F.
Richm, Ber d. Kais. Biol Ans. f. L. C, X. Y. (Geneva) B. 32S: 352.

B., Journ.
Heft,
8,
1908.
"
92
M.
Myc.
7: 105, 1902.
Sci. 1: 1909.
Miyabe, K., Trans. Sappora Acad.

J. J.,
" Davis, "

95 9
9'
Science, 31: 752, 1910. Clinton, G. P., Ct. R. 336: 1904, 1905.
Rostewzew, Ann. Inst. Agron. Moscow, 0: 47 and Flora 92: 405, 1903. Clinton, G. P., Ct. R. 23: 277, 1899.
H., Fla. R. 30, 1900. Garrison, S. C. B. 116: 7, 1905. 9 Halsted, B. D., Bot. Gaz. 1/,: 149, 1889. lo" Farlow, W. G., Bot. Gaz. 14: 187, 1889.
Hume, H.
Orton
&
'!
i2
Selby, A. D., Bot. Gaz. 27: 67, 1909. Stewart, F. C, X. Y. (Geneva) B. 110: 158, 1897.
w'
1" "^
io
Arthur,
J.
C, X. Y. (Geneva)
R.
/,:
253, 1885.
Halsted, B. D., X. J. B. 70.
Whetzel, H. H., N. Y. (Cornell) B. 21S: 1904. Taylor, T. R., D. Agr. 193, 1S72.
i'Trelease,
i8 '"9
Wm.,
Trans. Wis. Acad. Sc. G:
7,
1881-1884.
Wis. R. 16: 34, 1883. Shipley, A., B. 19: Miss. Kcw. 1887.
Stewart, F. 305: 394, 1908.

"'
""
C,
French, G. T.,
1892.
&
Wilson, T. K., B. X. Y. (Geneva)
Rostrup, Zeit. 2:
1,
112
1= 1' 1"
1'
i

Magnus,
P.,
Hedw.
149, 1892.
Massee, G., Jour. Linn. Soc. Bot. 2^: 48, 1887. Barrett, O. W., R. Porto Rico 398, 1904.
Raciborski, M., Ber. d. Deut. Bot. Ges. 15: 475, 1897. Spegazzini, C., Rev. Argent. Hist. Nat. 1: 36, 1891. " Magnus, P., Ber. d. Deut. Bot. Ges. 28: 250, 1910.
18
1'
Taplin, W. H., Amer. Florist 21: 587. C. R. Acad. Sci. Paris, 136: 472, 1906.
Blakeslee, A. F., Proc. Acad. Art. & Sci. ^0: 1904. Halsted, B. D., Amer. Flor. 13: 117.
20
21
" Stewart, F. C, N. Y. (Geneva) B. 328: 342. " Halsted, B. D., N. J. B. 76: 1890. " Orton, W. A., Sc. 29: 916, 1909. " Kirk, T. W., N. Zeal. D. Agr. R. 77: 1909.
2s
"
28 29
Swingle, D. B., B. P. Kew Bull. 871, 1897.
I.
37: 1903.
30
Rept. Mic. Vio., N. S. Wales, 1909. Thaxter, R., Rhodora 99: 1903. Atkinson, G. F., N. Y. (Cornell) B.
9I^:
252: 1895, also Bot. Gaz.
19: 47, 1894.
Gussow, Ottawa B. 63, 1909. Trow, A. H., Ann. Bot. 18: 541, 1904. " Idem, 15: 269, 1901. " Rosenberg, 0., Bihand till K. Svens Vet. Akad. Handl. 28: 35 Gruber E., Ber. d. Deut. Bot. Gaz. 19: 51, 1901.
'2
3
31
10, 1903.
3^
38
Edgerton, C. W., La. B. 126: 1911. Smith, E. G., Sc. 30: 211, 1909. McCallum, W. B., Ariz. R. 583, 1909.
Halsted, B. D., N. J. R. 1893, 393. Stevens, F. L., Bot. Gaz. 38: 300, 1904. Bubak, Fr., C. B. 8: 817, 1902.
39
^0
" Magnus, P., C. " Farlow, W. G., " Scribner, F. L., Stewart, F. C.,
Bak. 9: 895, 1902.

Bus. Inst. 1: 415, 1871. D. Agr. R. 96, 1886 ond 88, 1887.
Eustace, H.
J.
&
Sirrine, F. A.,
N. Y. (Geneva) B.
2U:
1903.
J.,
Morse, W. " L.
Jones,
Me. B. 169:
1909.
R., Vt. B. 72: 1899.
*8 Stewart, F. C., Eustace, H. J. and Sirrine, F. A., N. Y. (Geneva) B. 26^: 1904. 1^9 Farlow, W. G. B. Bussey, Inst. 415, 1876.
ASCOMYCETES
2. 7. 19. 26, 46. 62. 63.
62
(p.
64)
The distinguishing mark of this group is the ascus. This in its typical form is shown in Fig. 73, as a long, slender or club-shaped sac in which the spores are borne. The number of spores in the ascus is usually definite and is commonly of the series, 1, 2, 4, 8,
16, 32, 64, etc., the
vary
in
in
size,
color,
most common number being 8. The spores shape, markings and septation. The asci
layer, con-
most genera are arranged in a definite group, a stituting the hymenium which may be either concave, convex, or flat. Between the asci in the
hymenium
are often found slender hyphal threads
of various form, the paraphyses, Fig. 73. The hymenium may be borne in or upon
i\
threads, the stroma, or upon a very tenuous substratum, the subicu lum, or without any definite subascal structure.
firm substratum of
woven
The stromata vary widely
in character, size, tex-
ture, color, surface, form, etc.
The mycelium
and
this
is
usually abundant, branched
septate, the septation readily distinguishing
group from the Phycomycetes. In many mycelium weaves together into a false parenchyma and constitutes relatively large
species the
pj^,
spore-bearing structures.
Fig. 74.
of
73._port ion a hymenium

asci
The
if
showinK
ter
and
Af-
ascigerous organ, ascocarp, or ascoma,

is
paraphyses.
saucer-shaped and open

if
an apothecium,
Chamberlain.
closed a perithecium, Fig. 144. ascigerous layer covers the exterior surface.
Fig. 92;
In other cases, the

Fig. 74.
On the boundary lines between the Ascomycetes and other groups

are fungi which do not present the typical Ascomycete picture
but which are regarded as probably belonging to the group, i. e., transition forms between this and other groups. Among such are
113
114
forms in which the asci are without either stroma or covering, (Protodiscales, p. 125); others in which the asci are not even in
groups but are scattered irregularly throughout the ascocarp (Aspergillales, p. 164); and still others with the asci neither in
One regular groups nor covered (Protoascomycetes, p. 119). further deviation from the typical form occurs in the Hemiascomy-
FiLi. 74.
The large ascocarp of the murel.
After Freeman.
which possess a sporangium-like structure resembling that of the typical Zygomycete; but a mycelium like that of the typical Ascomycetes. This is by many regarded as the transition form bridging the gap between and indicating the kinship of these two groups; a view strongly supported by the existence of very similar
cetes
sexual processes in the two groups. Besides the ascus the Ascomycetes possess
many
other kinds of
THE
Frxr.i
wiimi cause plant disease
115
These may be rows on simple or branched conidiophores. The conidiophores may be single or variously grouped in columns or layers. Figs. 352, 378, 382. In some instances they are very
reproductive struetures in the form of conidia.
in
borne singly or
.an
err.
Fig. 75.
Sphserotheca castagnei. Fertilization and development of the perithecium. Og= oogonium, an = antheridium, st= stalk-cell. 6 as the aseogonium derived from the oogonium. After Harper.
form sporogenous cushions below the epidermis or again they may be borne inside of a hollow structure, the pycnidium, which covers them. Chlamydospores are also found.
short, innate; again they are long, loose or fioccose. emerge through stomata singly or in tufts or they may
They may
One
or
several
distinct
types of
to
A.
sporification
may
belong
one
species of Ascomycete. These different forms of spores may appear
simultaneously on the same mycelium or the}' may follow in definite

succession regulated in environment, or
by the changes
Boudiera. Six sets of Fiu. 76. sexual organs. After Claussen.
'n/
more
again one or forms belonging to the life history of the fungus may be omitted for long intervals to appear only as the result of stimuli of which little is yet known.
of the spore
The
conidia and chhimydospores are asexual spores.
Sexuality
116
A.n
/
's/^;
V'
in the great majority of
Ascomy-
cetes
has not been investigated;
but
'7^
in
some
in
hLh(K%
known
at
species fertilization is to occur; in many species,
least
form similar to that

it is
shown by the Phycomycetes,

absent, probably having been
lost
T<^ \
/ -^
77. Later
by degeneration
modified.
or else very
much
Fig.
stage showing asci and ascophores. After Clausscn.
of the Discomycetes one or more carpogonia and fertilization is through a tri-
In
some
is
there
chogyne by spermatia; a mode often met among the lichens. InPyronema,^Fig. 78, the carpogonium is multi-nucleate and it is fertilized by a multi-nucleate antheridium through a trichogyne. Fuase
asc
Fig. 78. oogonium, t= trichogyne. Pyronema eonfluen.s. A. the .sex organs, og B. fertilization stage in section through young apothecium, asc=asci, asf asfilament. After Harper. cogenous
E
=
=
si
on
cetes.
is probably in pairs as in Albugo bliti of the PhycomyIn Boudiera a very similar relation is found. Figs. 76, 77. In some Perisporiales ^ an uninucleate oogonium is fertilized by
of nuclei
'"
an uninucleate antheridium.
Fig. 75.

The oogonium
after fertilization gives rise to
of
117
less
a more or
simple in
the ascogenous hyphae, very in some Discomycetes, which produces Erysiphaceae, very complex the asci. The sterile parts of the ascocarp, the paraphyses and enveloping structures, arise from parts below the oogonium and
complicated system
antheridium.
The very young ascus usually receives two nuclei from the parent strand of the ascogenous hypha. These nuclei unite giving the
?-?>^'
'A'<?
Fig. 79.
Tip of ascus
showing
of Erysiphe
delimitation of ascospore from asco-
plasm
rays.
by astral
After Harper.
118
Hemiascomycetes
There
is
(p.
117)
a single order, the Protomycetales, which contains about
twenty-five species. MyceUum filamentous, branched, septate; conidia present; asci sporangia-like, containing numerous spores, terminal, naked or covered with a hyphal felt; in some species
known
to originate from the fertilization of an oogonium.
Protomycetales
Key to
Asci naked
Asci long, tubular
Families of Protomycetales
1.
Ascoideaceae,
Asci elliptic or globular Asci more or less covered by hyphse
2. 3.
Protomycetaceae, Monascaceae.
p. 118.
Of these families the

rophytic.
first is
found
in slime flux; the last
is
sap-
Protomycetaceae
asci intercalary or terminal, large, dearrested before spores are formed; a process which is velopment completed only after a period of rest.
Mycelium prominent;
Key
to Genera of Protomycetaceae
1.
Parasitic, intercellular in living plants
Protomyces,
p. 118.
Saprophytic,
building
hemispheric sporing
2.
masses
Endogone.
Protomyces Unger
Asci thick walled; after a long period of rest forming a large
mass
of elliptic spores which conjugate in pairs, then germinate immediately by a germ tube. This genus is sometimes placed with the Phycomycetes.^^
119
P. macrosporus Ung. Asci globose to elliptic, 40-80 x 35-60 n; membrane yellowish, up to 5 /i in thickness, contents colorless; spores elongate-ellipsoid,
2-3 X
It
/x.
produces small
galls,
which are at
first
watery looking, then
Fig.
81. .-1, mycelium and Protomyces. young ascus; E, ascus with mature spores.
After
De Bary.
brown, upon the leaves and stems of various economic and noneconomic Umbelliferae. P. P. pachydermus Thiim. affects carrots and dandelions.
rhizobius Trail, grows on Poa annua in Scotland. species are found on wild plants.
Several other
Subclass
Protoascomycetes
(p.
117)
There
is
a single order, the Saccharomycetales, with about

often undeveloped asci isolated or formed at different
;
seventy species.
Mycelium
points on the mycelium, mainly 4-spored; spores asexual reproduction by gemmation or by conidia.
unicellular;
120
Key
to Families op Saccharomycetales
Vegetative cells single or looselyattached in irregular colonies, mycelium not usually developed,
asci isolated,
not differentiated
1.
from vegetative cells Vegetative cells forming a mycelium,

asci
Saccharomycetaceae,
p. 120.
terminal,
or
intercalary,
2.
differentiated
from mycelium..
Endomycetacese,
p. 122.
The first family, the yeasts, to which belong the majority of the species of the order, is of prime importance in fermentation. A
Fig. 82.
Yeast plant-bodies, showing After budding and spomlation.
Coulter and Rees.
few species are known to cause animal diseases; others are found associated with the slime fluxes.
SaccharomycetaceaB
separate or few together, never truly filamenasci globose to elliptic, 1 to 8-spored; growing typically in sugary or starchy materials.
Vegetative
cells
tous, propagating
by buds;
121
Key to Genera
cells
of Saccharomycetaceee
Vegetative cells globose, ovoid, pyriform, etc.

Vegetative
increasing
by budding;
asci typically 3 to 4 spored.
Spores globose or ovoid. Spores
upon germination
cells.
forming
typical yeast
Ascus formation preceded by the

conjugation of like gametes.
.
1.
Zygosaccharomyces.
Ascus formation not preceded by

the conjugation of gametes. Spore membrane single Spore membrane double Spores upon germination forming a
poorl^^ developed proraycelium. Spores pileiform or limoniform, costate
4. 5. 2. 3.
Saccharomyces, p. Saccharomycopsis.
121.
Saccharomycodes.
Willia.
Spores hemispheric, angular or irregular

in form,
upon germination forming

6.
;
an extended promycelium
Vegetative
8-spored
cells
Pichia.
increasing
by
fission
asci
7.
Schizosaccharomyces.
Vegetative
cells elongate,
cylindric; spores
filiform,
Asci 1-spored Asci 8-spored
8. 9.
Monospora. Nematospora,
p. 122.
Saccharomyces Meyen
Vegetative cells globose, ellipsoid, ovate, pyriform, etc., reproducing by budding and remaining attached in short, simple or branched pseudo-mycelial groups, at length separating; asci
globose, ellipsoid, or cylindric, 1 to 4-spored (typically 3 to 4spored), single or in chains; spores globose to ellipsoid, continuous.
Many
species, chiefly saprophytes.
S. croci
Roze
is
described as the cause of a crocus disease.'*

plants suffering from blight a yeast
From sorghum
by
Radais.^
was
isolated
This when inoculated in pure culture into healthy
plants produced the characteristic lesions and effects.
122

Nematospora Peglion
(p.
121)
Colonies (in culture) disciform;
cells
elongate; asci cylindric,
8-spored; spores filiform, continuous, long-ciliate, hyaline.
Monotypic. N. coryli Pegl.,^ the cause of malformation of the hazel nut in Italy, is a peculiar fungus with what appears to be asci containing eight long slender flagellated spores.
Endomycetaceae
(p.
120)
Mycelium usually well developed, often producing a luxuriant growth, multiseptate; asci borne singly on branches, or intercalary, 4 to 8-spored; spores one-celled; conidia produced apically,
unicellular.
Key
Mucorales
to Genera of Endomycetaceae
parasitic
Mycelium poorly developed, Mycelium

well developed
on
1.
Podocapsa.
Asci formed after conjugation of a pair of

spirally
entwined branches
2.
Eremascus.
Asci formed asexually, produced terminally, rarely intercalary. Asci 4-spored Asci 8-spored
3. 4.
Endomyces,
Oleina.
p. 122.
Endomyces Rees
Mycelium
well developed, byssoid;
asci
borne singly on the ends of short lateral

branches, globose spores continuous.
to
pyriform,
4-spored,
'J
X':<^
The members
tionable
of this
genus are of ques-
importance
are
commonly found
parasites. in sap exuding
as
Some
from
^'sJli'E'SsTel
later
form in these,
they, together with other fungi present, set up a fermentation the products of which prevent the wound
tree
re-
wounds" where
from healing and result in injury. One species has been ported in America as an active parasite on apples.

E. mali Lewis
^
123
developed, multiseptate; conidia formed on short conidiophores or on the ends of short germ tubes, averaging 3 X 8 /x; no yeast-like budding; asci usually
Mycelium
well
on short
lateral branches,
11-14
/x
in diameter;
ascospores sphseroidal, slightly elongate, 4.5 x 5.5 n with thickened places on the walls, brown
when mature.
,
Figs. 83, 84.

,
1
^^^'
1
84. E.
mali.
Typical manner Lewis isolated the fungus from decayed spots ' beanng conidia T on apples by plate cultures. Inoculations proved on agar. After ^^^^' that it is capable of causing a slow decay without the aid of other fungi. An extensive cultural study as well as a considerable cytological study was made.
,
E. decipiens (Tul.) Rees
is
parasitic
on Armillaria; E. parasitica
Fayod on Tricholoma."'
^^
Euascomycetes
This
is
(p.
117)
species, with great variety of
an extraordinarily large group comprising some 16,000 size, color and shape of plant body.
still
Most
of ther. are saprophytes,
many
are parasites either in
their ascigei'ous or their conidial stages of development.
The twelve
orders
may
be recognized by the following key.
Key
nium, no ascoma
to Orders of Euascomycetes
Asci approximate in an indefinite hyme1.
Protodiscales, p. 125.
Asci grouped in a definite ascoma Asci collected in a flattened, concave or

closed ascoma, often bordered
distinct layer
by a
Ascoma
at maturity open
and more or
Discomycetes Ascoma open from the first, clavate or

convex, pitted, or gyrose
2.
less cup-like.
Helvellales, p. 130=
Ascoma
at
first closed,
opening early,
3.
without special covering, more

or less fleshy
Pezizales, p. 133.
124

Ascoma opening
tardily, enclosed
by
a tough covering which becomes torn open at the maturity of the
spores
Ascoma
roundish, opening
by
stel4.
late or radiating fissures
Phacidiales, p. 154. Hysteriales, p. 159.
Ascoma
Ascoma
elongate,
opening by a
5.
longitudinal fissure at maturity closed and tuber-
like, subterranean, Asci collected in a cylindric or globose
6.
Tuberales.
perithecium
Perithecia sessile, solitary and free, or united and embedded in a stroma
Asci arranged at different levels in the perithecium Asci arising from a
7.
Aspergillales, p. 164.
common
level
Mycelium
superficial,
perithecia
scattered, globose
and without
8.
apparent
ostiole, or flattened
and
ostiolate
superficial, peri-
Perisporiales, p. 170.
Mycelium nearly
Perithecia
thecia ostiolate
and stroma
(if
presp. 195.
ent) fleshy or
membranous,
9.
(if
bright colored Perithecia and stroma
Hypocreales,
pres-
ent) hardened, rarely
mem-
branous, dark colored
Wall
of perithecia scarcely distinguishable from the

10.
stroma
Perithecia with distinct wall, free or embedded in the
Dothidiales, p. 215.
stroma
Perithecium borne on a short pedicel; microscopic fungi parasitic on
insects
11. Sphaeriales, p. 221.
12.
Laboulbeniales.
Of these all contain plant parasites with two exceptions; the Tuberales, which bear underground tuber-like ascocarps, some of
125
these prized as table delicacies, and the Laboulbeniales, an order rich in species which are all parasitic upon insects.
Protodiscales
(p.
123)
The 4-8
which
to
arises directly
many-spored asci form a flat pali.sade-like hymenium from the mycelium; paraphyses none; spores,
one-celled, elliptical or round.
Key
Parasitic
to Families of Protodiscales
1.
Exoascaceae,
p. 125.
Saprophytic
2.
Ascocorticiaceae.
Of these families the second contains only one genus and two species found in bark. The first family is aggressively parasitic.
Exoascaceae
This
is
8-10. 292,
327
the most simple of the parasitic Ascomycetes, definitely recognizable as such, and is comparable with the Exobasidiales
among
the Basidiomycetes.
are parasitic
All the
and many of species them very injurious. The mycelium, which can be distinguished from
that of other fungi by
its
cells
of
very irregular size and shape, wanders between the host cells (intraf^""^ cellular in one species), or is some//
times
limited
to
the
region
asci
below the
cuticle.
The
develop
just Fiq. 85.Exoascus showing myceHum and asoi. After Atkinson.
form on a mycelial network under the epidermis, or the cuticle, or on the ends of hyphaj arising from below the epidermal cells. They are exposed by the rupture of the cuticle or
in a palisade
epidermis and contain four to eight hyaline, oval, one-celled spores. These by budding, while still in the ascus, may produce numerous secondary spores, conidia, which give the impression of a many-spored ascus. The ascospores also bud freely The primary-ascus-nucleus arises from in nutritive solutions.
125
fusion of

two nuclei
spore-nuclei arise
as is general among the Ascomycetes. The repeated mitoses of the primary nucleus. by Affected leaves, fruit and twigs become swollen and much disthe torted; wrinkled, curled, arched, puckered. In woody twigs
mycelium often induces
unnatural, profuse, tufted branching resulting in "witches brooms" though such structures
often
tion
arise from irritadue to other causes. Many attempts have
Fig. 86.
Taphrina showing mitoses in the young aecus leading to the development of sporenuclei. After Ikeno.
been made to arrange the

species in natural genera;
some based on the num-
354 6. ^^^ others largely on the biologic grounds of anber of ascospores,*' ^ whose classification nual or perennial mycelium.^ Giesenhagen
is
followed here, recognizes two genera, Exoascus being merged
into Taphrina.
Key
to Genera of Exoascaceae
Asci cylindric, clavate or abbreviate-cylinof dric, produced above the epidermis
the host
Asci saccate, in epidermis
1.
Taphrina,
p. 126.
2.
Magnusiella.
Taphrina Fries
or perennial; asci 4 to 8-spored, or by germination of the ascospores, multispored, borne on the surface of blisters and other hypertrophied areas, cylindric to clavate, or a modifica-
Mycelium annual
tion thereof.
^ Of this genus Giesenhagen recognizes four which are arranged in three subgenera. of species
series
Subgenus
1.
Taphrinopsis,ône
series (Filicina)
The asci are slender clavate, narrowed at each end, rounded above, broadest in the upper fourth. Parasitic on ferns. None of the five species is of economic importance.
127
Subgenus.
2.
Eutaphrina,
one series (Betula)

On
Asci broadly cylindric, rarely contracted at the base or from the middle down, truncate above and sometimes in-sunken.
Amentacese, chiefly Betula, Alnus, Ostrya, Carpinus, Quercus, Populus. Of the twenty-four species of this series but few are
of importance.
T. coerulescens (D.
& M.)
Tul.^^
Annual, producing blisters

asci elongate,
on the leaves
conidia.
of oak, the sporing surface bluish;

/z;
broadly cylindric, 55-78 x 18-24
spores breaking
up
into
On
various species of Quercus in Europe and America.
T. ulmi (Fcl.) Joh., on the elm; T. aurea (Pers.) Fries on the leaves of Populus and T. johonsonii Sad. on the fertile aments of the aspen are among the more important remaining species of the
series.
Subgenus
3.
Exoascus,
two
series
Asci clavate, normally cylindric or more or less abbreviated. Asci slender, clavate, narrowed (1) Prunus series on Rosacese.
below, broadest in their upper fourth, varying through mediate forms to narrowly cylindric.
(2)
all inter-
on Sapindacse, Anacardiacea;, etc. Asci broadly cylindric, short, rounded or truncate. The more important economic species of the genus belong to the Prunus series. T. deformans (Fcl.) Tul.^- '-^ '^
tEscuIus series,
The irregular vegetative mycelium devoid of haustoria grows in the leaf parenchyma and petiole and in the cortex of branches.
distributive mycelium lies close beneath the epidermal cells of twigs and in the pith and extends some distance through the twig. Fig. 87. Branches arise from the vegetative mycelium, penetrate between the epidermal cells to the cuticle and then branch freely
to form a network of short distended cells beneath the cuticle.
This
is the hymenium, a layer of ascogenous cells. These cells elongate perpendicularly to the host's surface. Fig. 85, rupture the cuticle, and form a plush-like layer. The protoplasmic con-
128
tents crowds toward the tips of these cells and a basal septum cuts off the ascus proper from the stalk cell, Fig. 88. The spores then
form within the ascus.
The ascospores may bud

or after
either before
extrusion
from the ascus, producing conidia, which
may
themselves bud
indefinitely,
producing
secondary, tertiary, In this etc., crops.

condition the conidia
strongly
resemble
On the yeast host plant ascospores

cells.
germinate
tubes,
by
germ
which are cae
pab
of
infecting
proper
attempts
nidia.
is
hosts.
No
success has
rewarded
secure
co-
to
germ tubes from

chiefly
Leaf infection
external
;
rarely
in
internal
from
mycelium perennating
curs
the twigs. It ocwhen the leaf is

Infected
are
very young.
leaves
Fig.
1,
thickened
87. T. deformans. distributive hypha; 5, vegetative hyphae; 9, sporiferous hyphse. After
and
the
broadened
tissues
and
stiff
are
Pierce.
and coriaceous.
The
palisade cells increase in size and

phyll.
number and
lose their chloro-
Blistering and reddening of the leaves follows. Asci clavate, 25-40 x 8-11 /x; spores 8, subglobose or oval, 3-4 }j.. On the peach in Europe, North America, China, Japan,
Algeria and South Africa. T. pruni (Fcl.) Tul.^' ^^
is
found in Europe and North America
129
on plum and wild cherry, causing "plum pockets." The ovary is The mycelium after bud infection pervades the mesocarp which hypertrophies and alone produces a much
the seat of attack.
enlarged fruit, usually with entire sacrifice of the other fruit parts. Asci are formed over the diseased surface much as in the last species. The mycelium is perennial in the bast and grows out into the new shoots and
buds each spring.

fection also
In-
reaches
other shoots and trees
by means
of the spores.
Ascus elongate-cylindric, 30-60 x 8-15 /x; spores 8, globose 4-5 /z.

Perennial.
produces
T.cerasi(Fcl.)Sad.S'i^ the witches

effect
broom
It
is
upon
in
culti-
vated and wild cherries.
common
rare
;
Eu-
rope,
in
America.
clavate
Perennial
asci
ii;
30-50 X 7-10
8,
forming
M.
spores conidia in
the ascus,
oval,
6-9 x
nMrrr
fig. ss.
-T. deformans.
Young and
old asci.
5-7
After Pierce.
avium, P. cerasus, etc. in North America and Europe. T. mirabilis (Atk.) Gies.^' ^^ grows on leaf buds and twigs of Prunus angustifolia, P. hortulana, P. americana in North America. Perennial sporing on the fruits and tips of branches of the host
;
;
On Prunus
asci subcylindric, blunt above,
25^5
x 8-10
ix;
spores
8,
ovate.
in
T. longipes (Atk.) Gies.

Perennial; sporing on
not,
is
on Prunus americana
fruits; asci cylindric,
ii.
North
America, causing plum pockets.^
young
8,
truncate or
ii; spores T. rhizipes (Atk.) Gies.
30-40 X 7-10
globose or ovate, 3-4

^
Known
ing pockets on Japanese plums;
only in North America, causprobably of wider distribution.
130
Perennial; asci cylindric, or club-shaped, 30-40 x 8-10 n, appearing to have basal rhizoids; spores 8, globose. T. communis (Sad.) Gies.^ Perennial in branches; sporing on immature fruits; asci clavate, 24-45 x 6-10 fi; spores 8, elliptic, 5 X 3-4 II, often producing conidia.
On Prunus americana, P. maritima, P. North America.

T.
institiae
nigra,
and P. pumila,
in
(Sad.) Job.
institia,
P.
domestica,
Forming witches brooms on Prunus and P. pennsylvanica in Europe and
America.^
Perennial; sporing on the under side of the leaf; asci clavate to
cylindric,
25-30 x 8-10
(Atk.)
/x;
spores 8, not rarely producing conidia,
globose, 3.5 m-
T. decipiens America.^
Gies.
On Prunus americana
in
North
Perennial; sporing on under surfaces of leaves; asci irregularly

clavate, often almost cylindric, 20-40 x 10 into conidia.
/x;
spores breaking
up
T. buUata (Fcl.) Tul.
On
Annual;
asci clavate,
pear and Japanese quince. 36-40 x 8-9 ^c; spores 8, often forming
conidia, globose, about 5 /*. T. farlowii (Sad.) Gies.^ls
found on Prunus serotina in America;
T. minor Sad. on leaves of Prunus chamaecerasus and P. cerasus, in Germany and England. It has recently caused considerable
damage
in
South England.
T. bassei Fab. causes witches broom of cacao in Kamerun.

T. rostrupiana (Sad.) Gies. is on Prunus spinosa; T. crataegi (Fcl.) Sad. on Crataegus oxycantha. T. maculans Butler
Butler.-^^
is reported on Tumeric and Zinzibar by T. theobromae Ritzema Bos. is reported as injurious to
the cacao tree.
Many other species of Taphrina of minor importance occur upon

alder, poplar (Populus), Carpinus, birch, elm, maple,
hawthorn,
oak and numerous other
hosts.
Helvellales
(p.
123)
Ascoma
fleshy, separable into a definite

is
hymenium
and
of asci
and
paraphyses and a stroma which
usually large
stalk-like;

fertile
131
portion more or less cap-like; hymenium free from the first " asci or covered with a thin, evanescent veil; cylindric, opening
by an
apical pore; spores ellipsoid, colorless or light yellow, smooth, or in one genus echinulate.
Key
Ascocarp stalked
Fertile
to Families of Helvellales
portion
clavate
or
capitate;
asci
1.
opening by an irregular slit Fertile portion pileatc; asci opening by a
Geoglossaceae,
Helvellaceae.
p. 131.
lid.
2. 3.
Ascocarp
sessile
Rhizinaceae, p. 132.
The majority of the species of this order are saprophytes, the only parasites being of the first and third families. Of the second family many of the species are edible and some are very large.
Geoglossaceae
Key to Tribes and Genera of

Ascoma
clavate or spatulate, ascigerous portion usually more or less compressed, rarely subglobose.
. . .
Geoglossaceae
Tribe
I.
Geoglosseae.
Ascoma
Spores
clavate, fertile portion at
most
or
only slightly decurrent

small,
elliptic,
cylindric
;
continuous fusiform, bright colored
plants
1.
Mitrula, p. 132.
Spores long, elliptic to cylindric, 3 to many-septate at maturity
Hymenium Hymenium
bright colored black or blackish
2.
Microglossum.
Corynetes.
Spores hyaline
Spores brown
3. 4.
Geoglossum.
Ascoma
or fan-shaped, asdecurrent on the stipe cigerous portion Ascigerous only on one side of the
spatulate
stem
Ascigqrous on both sides the stem
5.
Hemiglossum.
Neolecta.
Spathularia.
Spores globose
6.
7.
Spores elongate
Ascoma
genus
stalked, capitate or pileate, in one

sessile
II.
Cudonieae.
132

Mitrula sclerotiorum Rost.^* which causes a disease of alfalfa in
Denmark is the only pathogen of the family. The infected plants die and later the roots and stems become
Fig. 89. Mitrula. B, habit sketch; F, asci. After Schroter.
filled
lie
with
black
sclerotia
which
dormant about a
year.
resuming growth they become covered by light red eleva-
Upon
tions,
which bear small
light red ascocarps.
Rhizinaceae
(p. 131)
Key
Spores
elliptic or
to Genera of Rhizinaceae
spindle-shaped
1.
Without rhizoid-like structures With rhizoid-like structures

Spores globose
Psilopezia.
2.
Rhizina, p. 132.
3.
Sphaerosoma.
Only one genus, Rhizina, causes

disease.
Rhizina
species
is
some eight recognized by its crustFries
with
formed,
root-like
side.
sessile,
ascophore with outgrowths from the lower

fiat
Fig.
90.
spored, opening
celled, hyaline;
is
by a
Asci cylindrical, 8lid; spores oneIt
paraphyses many.
often purely saprophytic, growing

is
in burned-over spots in forests. "^ R. inflata (Schiiff) Quel.'^'
counted as the cause of serious root

diseases
of
Fig. 90.
forest
trees,
especially
Europe. The fungus also occurs in Asia and America. R. undulata causes death of fir seedlings. '^^
conifers, in
Rhizina inflata. B, ascoearp from below; C, asci and paraphyses. After Schroter and Tulasne.
133
Pezizales
(p.
123)
In this order unlike the last, the hymenium is at first enclosed but soon becomes exposed. The apothecia at maturity are typically disc or saucer-shaped (Fig. 101) or sometimes deeper, as They vary from a size barely cup, beaker or pitcher-shaped. visible up to 8-10 cm. in diameter. Some are stalked, more often they are sessile. In consistency they vary from fleshy or even gelatinous to horny. Paraphyses are present and may unite over the asci to form a covering, the epithecium. The apothecium may be differentiated into two layers; the upper bearing the asci is the
hypothecium, the lower the peridium.

are formed.
In some cases sclerotia
species possess conidiospores as well as ascospores, borne either on hyphse or in pycnidia. The great majority are saprophytes, a few are parasitic. There are some three thou-
Many
sand
species.
Key to
Xo
lichenoid thallus
Families of Pezizales
algal cells
and no
Ascocarps free, solitary or cespitose Ascocarps fleshy or waxy, rarely gelatinous; ends of paraphyses free Peridium and hypothecium without
distinct lines of junction
Ascoma open from the
beginning,
convex; peridium wanting or
Ascoma concave
poorly developed at first; a fleshy
1.
Pyronemaceae.
peridium present.
Asci forming a uniform stratum, at maturity not projecting
.
2.
Pezizaceae.
Asci projecting from the ascoma

at maturity Peridium forming a more or
3.
Ascobolaceae.
less dif-
ferentiated
membrane.
elongate,
parallel
Peridium
of
pseudo-parcnchymatous, hyaline,
thin-walled
cells
4.
Helotiaceae, p. 134.
134

Peridium
firm, of
roundish or angu5.
lar, pesudo-parenchymatous, mostly dark, thick-walled cells
Mollisiaceae, p. 146.
Ascocarps leathery, horny or cartilaginous ends of the paraphyses united

;
an epithecium Peridium wanting or poorly developed Peridium well developed, mostly leathery or horny Ascccarps free from the beginning,
into
6.
Celidiaceae.
dish or plate-shaped, never enclosed by a membrane
7.
Patellariaceae.
Ascocarps at first embedded in a matrix, then erumpent, urceolate or cup-shaped, at first en-
closed in a
membrane which
8.
disappears later
Cenangiaceae, p. 150.
Ascocarps borne on a highly developed stringy or globoid stroma
Ascocarps at the ends of the branches of a cord-like stroma

Ascocarps embedded in the upper portion of a globoid stroma
Lichenoid thallus more or
less
9.
Cordieritidacese.
10.
Cyttariaceae.
prominent,
11.
algal cells typically present, asci disap-
pearing early, disk with a mazsedium.
Caliciaceae, p. 153.
A-scobolaceae are pure saproPeziacese, on organic matter in the ground or on rotting wood. The phytes Patellariaceae are largely, and the Celidiaceae are nearly all, parasitic
The Pyronemacese,
and
on
lichens.
The
Cordieritidaceae of four species, possessing
a stony stroma, are unimportant.
The
Cyttariaceae, of one genus,
and some
six species, are limited to the
southern hemisphere where
they grow on branches of the beech.
Helotiaceae
(p.
is
133)
In members of this family there

peridium.
first
The apothecia
a distinctly differentiated are usually fleshy or waxy, superficial,
closed, later opening; the
Asci 8-spored.
paraphyses form no epithecium. Spores round to thread-shaped, one to 8-celled,

hyaline.
135
Some
of the genera are
among
the most serious of
plant pathogens.
About one thousand
species.
Key to Genera of
Helotiaceae
Ascocarps fleshy, fleshy- waxy, thick or membranous

Ascocarps fleshy-waxy, brittle when fresh,
leathery
when dry
I.
Sarcoscypheae.
Ascocarps felty hairy externally Ascocarps covered with bristle-like hairs

externally
1.
Sarcoscypha.
Pilocratera.
2.
Ascocarps naked Ascocarps springing from a sclerotium

Ascocarps
not
springing
3.
Sclerotinia, p. 136.
from
sclerotium
Spores 1-celled
Substratum green Substratum uncolored

Spores at length 2 to 4-celled. ... Ascocarps waxy, thick, tough or membranous
4.
5.
Chlorosplenium,
Ciboria.
p. 144.
6.
Rutstroemia.
Ascocarps externally hairy Ascocarps resting on an extended

arachnoid mycelium
Spores 1-celled Spores becoming several-celled ... Ascocarps without arachnoid mycelium
II.
Trichopezizeae.
7. 8.
Eriopeziza.
Arachnopeziza.
Spores globose
9.
Lachnellula.
Spores ellipsoid or elongate Disk surrounded by black hairs. 10. Desmazierella.
Disk smooth
Paraphyses obtuse at the apex Walls of ascoma delicate;
spores mostly 1-celled, rarely 2-celled al ma-
Walls
turity of
11.
Dasyscypha,
p. 144.
ascoma
thick;
spores 2-celled at
turity
ma12.
Lachnella,
p. 145.
136

Paraphyses lancet-shaped at
apex
Spores 1-celled
13.
Lachnum.
Erinella.
Spores
at
length
several14.
celled
Ascocarps naked
Spores globose
Spores ellipsoid or fusiform
III.
15.
Helotieae.
Pitya.
Spores 1-celled
Border of disk smooth
16.
17.
Hymenoscypha,
Cyathicula.
p. 146.
Border of disk toothed

Spores at length 2 to 4-celled
Ascocarps
sessile,
rarely
com18.
pressed at base
Belonium.
Ascocarps stalked, or at least compressed like a stalk
Walls of ascoma waxy; stem short and dehcate

Walls of ascoma waxy, thick;
19.
Belonioscypha.
stem thick
Spores filiform
20.
Helotium.
Gorgoniceps. Pocillum.
Ascocarps
sessile
21.
Ascocarps stalked Ascocarps gelatinous

gristly,
22.
horny when
IV. Ombrophileae.
dry
Of these genera only the five given below have parasitic representatives of economic importance, while only one to two others are The rest grow as saprophytes on rotting wood and parasitic. debris in the soil. organic
Sclerotinia Fuckel (p. 135)
of
This genus contains several very important pathogens, some them preying upon a wide range of hosts and causing great loss. striking feature of the genus is the sclerotium which is black and
borne within the host tissue or upon its surface. From the sclerotia after a more or less protracted period the apothecia develop. These
are disc-shaped and stalked. The asci are 8-spored; spores elliptical or fusiform, unicellular, hyaline, straight or curved. Some species
137
possess Botrytis forms (sec pp. 141 and 578), others Monilia (see pp. 139 and 558) forms of conidial fructification. In addition to
may be gonidia, which appear to be functionless conidia. In some species degenerate, there is no known spore form except that in the
these there
ascus.
S. ledi
Now.
is
of especial interest as the
one
fungus outside of the Uredinales that exhibits

heteroecism.^"^-
forms found upon separate hosts and presenting slight differences under the micro-
Many
no microscopic differences, have been named as separate species. Only long careful culture studies and inoculation experiments will determine which of these species are valid, where more segregation, where more
scope, often even
aggregation
is
needed.
Fig. 91.
S. urnula,
moniliform conidia
association of Botrytis or Monilia conidial forms with Sclerotinia, in the same host,
The mere
with
disjunctors.
After Woronin.
has repeatedly led to the assumption that such forms were geneticSuch assumptions are not warranted Only the most ally connected and most complete evidence justify such conclusions. careful study
.
contains some fifty species which are divided into two Stromatinia Boud., forming sclerotia in the fruits of subgenera; the host; Eusclerotinia Rehm forming sclerotia in or on stems and leaves of the host. When conidia are known those of Stromatinia are of the Monilia
The genus
type and those of Eusclerotinia of the Botrytis type. Each group contains important economic species. 291-295 S. fructigena, S. cinerea and S. laxa.-o- î. 25.
These forms are perpetuated

ascus-forms are
chiefly
by
their conidia.
The
much
less often seen.
When the conidia fall upon the peach, the mycelium develops and penetrates even the sound skin, then rapidly induces a brown rot. The mycelium within the tissue is septate, much branched, and light brown in color. It soon proceeds to form a subepidermal layer and from this the hyphae arise in dusty tufts of Monilia-form conidiophores and conidia (Fig. 92). The earlier conidia are thin-
138
walled and short lived, the later ones thicker walled and more
enduring. After some weeks these tufts cease forming and disappear. The mycelium within the fruit persists, turns olivaceous and forms
large irregular sclerotioid masses may produce fresh conidia.
which on the following spring

fruits
These
sclerotioid
(mummified)
under suitable conditions
Fig. 92. Sclerotinia on plum, a, section showing a spore pustule and chains of conidia; b, part of a spore-chain; c, spores germinating; d, a and ascophores; e, an ascophore; /, ascus; g, mature spores. After plum
mummy
Longyear.
blossom time of the host, can also produce demonstrated by Norton. ^^ These apothecia develop in large numbers from old fruits half buried in soil, and send forth ascospores to aid in infection. The ascospores germinate readily in water and it was proved by Norton that they give rise to a mycelium which produces the characteristic Monilia. Inoculation of ascospores on fruit and leaves also gave positive results in two or three days. The flowers, and through them the twigs, are also invaded by the mycelium which seeks Shot-hole effect is produced on chiefly the cambium and bast.
in nature, usually at
apothecia, a fact
first
leaves of peach
and cherry (Whetzel
^^).
Infection
is
frequently
through minute wounds.^*
139
On the apple the fungus shows two different modes of development. In some cases the myceUum accumulates under the epidermis without producing spores, becomes dark colored and also causes a darkening of the contents of the host cells, which results in a black spot giving rise to the name black In other cases -^ the mycelium produces a rot.
brown
rot
and abundant
circles
conidial
tufts,
ar-
ranged in concentric
of infection.
around the point

has long been
The form on pomaceous
fruits
regarded as identical with that on stone fruits; but recently, at least in Europe, they have been
distinguished
on cultural and morphological

as separate species,
Fig. 93 -S. furkellattachment ana. organ. After Istvanffi.
grounds
(see also^^'-^,
the most distinctive character perhaps being the color of the mass of conidia. In a similar
way
S. laxa
Ad &
Ruhl.
is
set aside as a distinct species infecting
only apricots.^^^
American mycologists are inclined to doubt the distinctness of the species on drupes and pomes in this country. S. fructigena (Pers.) Schr.
Apothecia from sclerotia produced either on mummied fruits, 0.5-3 cm. high, stem dark brown, disk lighter, 5-8 or even 15 mm. in diameter; asci 125-215 x 7-10 /x; spores ellipsoidal, 10-15 x 5-8 n. Conidia (=Monilia fructigena Pers.). Coin or
nidiophores covering the fruits of the host with a dense mold-like growth of light
brownish-yellow or ochraceous color; spores averaging 20.9 x 12.1 /z. On stone and
Fig. 94.
Typical conid-
pome
fruits, especially
the latter.
iophore and eonidia of the Botrytis-fonn of S. fuckeliana. After Smith.
S. cinerea (Bon.)
Wor.
S.
Apothecia and asci similar to those of fructigena, Conidia (=Monilia cinerea
Bon.). Conidiophores covering the fruits with a dense grayish mold-like growth; spores averaging 12.1 x 8.8 On stone and pome fruits, especially the former.
jti.
140
S. linhartiana P.
&
D.^^
is
reported on quince in France.
S.
mespili Schell on medlar.

seaveri,
is
S. seaveri,
Rehm., conidia =Monilia

22
S. padi
on Prunus serotina.^^^ Wor. is found on Prunus padus and

e.,
Castanea.''
It possesses
junctors,
i.
a Monilia-form conidial stage with typical disspindle-shaped cellulose bodies between the conidia,
facilitate
which easily break across to
the separation of the conidia.
Fig. 95.
S.
Sclerotia produced in artificial cullibertiana. ture. After Stevens and Hall.
Wor. is found on cranberry. It is unique in that the spores in each ascus are larger than the others. The half of conidial stage is a Monilia.
S. oxycocci
S.
fuckeUana (De Bary)
Fcl.^^
Apothecia
in clusters of 2-3
from
sclerotia in the leaves, rarely
n across, stem 10-11 x 6-7 /x. slender; ascospores Conidia (=Botrytis cinerea Pers., B. vulgaris Fr.). Conidiophores simple or branched, forming dense gray tufts; conidia subin the fruits, of the host, yellowish-brown, 0.5-4
globose,
Fig. 94.
usually minutely apiculate,
almost hyaline,
10-12
/x.

It
141
leaves, fruit
causes a rot of the grape, much dreaded in Europe, attacking and stem. The fungus can persist long as a sapro-
phyte in the conidial condition. Scleratia are borne within the On germination they may either produce the affected tissues. conidia directly or form apothecia. Both ascospores and conidia are capable of infecting the grape but infection is much more
certain from a vigorous
mycehum
(see S. libcrtiana, p. 142).
Attachment organs, c. f. Fig. 93, which consist of close branchclusters and seem to be induced by contact of a mycelial tip with any hard substance are present in abundance. Both toxins and
digestive
enzymes are produced.-^'
Botrytis douglasii on pine is perhaps identical with the conidial form of the last fungus (see p. 140) as may also be the Botrytis of
Ward's Lily Disease; ^ the Botrytis causing disease of gooseberries ^^ and many others that have been named as distinct
species of Botrytis. S. galanthina Ludwig, close kin to S. fuckeliana, attacks snowdrops. S. rhododendri Fisch. occurs on Rhododendron.
The former
been adduced.
of these
two
is
of Botrytis galanthina Berk.
&
supposed to be the ascigerous form Br. but no conclusive proof has
S. libertiana Fcl.^^
Sclerotia
from a few millimeters up to 3 cm.
in length, black;
apothecia scattered, pale, 4-8 asci cylindric, 130-135 x 8-10
mm.
ju,
or
more broad, stem
slender;
apically
very
slightly
bluish;
spores
ellipsoid,
usually minutely guttulate, 4-6.5 /i; paraphyses clavate.
9-13
This fungus affects numerous hosts. Among the most important on which it
causes serious disease are
ginseng,^^
^^' ^^' ^^
lettuce,^''
Fiu. 'M. S. libcrti;ina. Sclerotia and apothecia. After Stevens and Hall.
carrot, potato, parsley, hemp, rape, various bulbs, zinnia,
cucumber,^-"
petunia, etc. The white mycelium is found superficially and within the host, especially at places where moisture is retained, as between leaves, at leaf axils, etc.,
also within plant cavities. Microscopically it consists of long Within cells branching in a rather characteristic way, Fig. 97.
142
the host's tissue the hyphal threads are thicker, richer in proto-
plasm, more septate, and than outside of the host.
much more branched and crooked Aerial hyphal filaments when they
touch a solid repeatedly branch in close compact fashion forming the attachment organs.
At the exhaustion
of the food supply
and the consequent term-
ination of the vegetative period the mycelium becomes very dense in spots and within these clumps of mycelium the sclerotium
forms; at
They
first white, later pink, finally smooth and black (Fig. 95). are often found in the leaf axils (lettuce), in the pith of
stems
(carrot), etc.
Under
on un-
some
conditions, as
suitable
nutrient media, gonidia are produced.
The sclerotia can germinate at once or remain

Fig. 97. Mycelium showing septation aud branching. After Stevens and Hall.
dormant
several
for
,
one, perhaps
mmation they send
...
years.
,
On
i
gerc forth
. i
from
soil
1 to 35 negatively geotropic sprouts which grow to the surface unless that be more than about 5 cm. distant. On
^^
reaching the light the apex of the sprout begins to thicken and soon develops its apothecium; at first invertedconidial, soon flat, and finally somewhat revolute. Changes in atmospheric humidity cause
the discharge of ascospores in white clouds. The ascospores germinate readily but the re-
such small vigor that it If the ascospore is incapable of parasitism. where it can maintain a saprophytic germinates
sulting
mycelium
is
of
Fig.
98. S.
asci
liber-
...
mycelium paraphyses. After Stevens and Hall. the mycelium may become parasitic, Both ascospores and mycelium are comparatively short-lived. The mycelium can migrate but a short distance over soil. No form of conidia except the apparently functionless gonidia is
life
is
until a vigorous
,.,
,.
developed then
',
tiana,
and
produced.
The fungus may be

is
cultivated easily
upon almost any

a
medium, corn-meal-agar
especially suitable. It has been repeatedly claimed that this fungus possesses

Botrytis conidial stage but
this.^^'
143
tlie
results of
much
careful
work deny
Recent
is
tests
by Westerdijk
'"
indicate the
absence of such
biologic specialization in regard to hosts as
Erysiphe and elsewhere. Oud. & Kon.^^' ''^ parasitize; the leaves and stems of tobacco. It is
found
in the
S. nicotianae
possibly identical with S. libertiana.

S. trifoUorum Erik.^'^"^^
In general this resembles S. libertiana with which it is by some regarded as idenevidence has, however, not been adduced to prove them the same. The sclerotia, varying in size from that of a mustard seed to a pea, are found in the detical; sufficient
cayed
tissue, or as larger flat surface sclero-
No conidia except the functionless gonidia.

tia.
Unknown on
S.
clover.
bulborum
'^^
Rehm
which
(Wak.) is very
Fig. 9U. Cultures of sclerotinia from tobacco on potato agar, showing sclerotia. After Clinton.
similar to S. trifoliorum
and without known conidia grows on hyacinth, crocus, scilla and tulip. Cross infections between hyacinth and clover have not, however, been successful and the species may be distinct.
sterile
form,
Sclerotium
tuliparum,
culti-
inosum.
co
aenig/, ascus;
found on the tulip S. tuberosa Fcl. vatcd anemones.
may
is
also belong here.
found on wild and
n FdTa.^*Âf ter
Bre-
Rehm and
them
riae
Several other species of the genus, among S. alni Maul, S. bctulaB Wor., S. aucupa-
Ludw,
S.
crategi
Magn., are
found on
Ericaceae, Betulacese, Rosaceae, Gramineae, etc., but they are not of sufficient economic importance to warrant further notice here.
144
Chlorosplenium Fries
(p.
135)
Ascoma mostly
aggregated, small, stalked, smooth without,
green; asci cylindric, 8-spored; spores elongate, 1-celled, guttulate,

hyaline; paraphyses linear. The genus consists of some ten species only one of which interest here.
C. aeruginosum (Oed.) d Not.
is
of
The apothecia and mycelium
are verdigris-green as
is
also the
Fig. 101.
D. wilkomii. A, natural size and single apothecium enlarged; B, an aseus. After Lindau.
wood penetrated by
phytic but
may
The fungus appears to be mainly saproit. be partially parasitic. Fig. 100.

Dasyscypha
Fries (p. 135)
is a genus of some one hundred fifty species, mostly saprobut sometimes parasitic on twigs. The apothecia are small, phytic
This
short-stalked or sessile,
disk,
waxy
or
membranous, bright colored

spores
ellipsoid
in the
with mostly simple hairs on the outside and margin.

or
clavate,
Asci
cylindrical
8-spored;
or
fusiform,
hyaline, 1-celled, rarely 2-celled, sometimes guttulate; paraphyses blunt, needle-like.
145
D. willkommii Hart.^^ causes a serious European laroli disease and affects also the pine and fir. The stromata appear as yellowish-white pustules on the bark soon after its death. Here hyaline conidia are produced on the open surface or in cavities. Apothecia 2-5 mm. broad appear The ascospores can infect wounds: the conidia seem to be later.
functionless.
The myceintercellular
lium spreads through the

sieve
spaces,
pith.
tubes,
and xylem
to the
i\
Apothecia short-stalked,
yellowish without, orange within; asci 120 x 9 /z;
spores
18-25
5-6
/x;
paraphyses
the ascus.
longer
than
''^
D. resinaria
a wound
like the
fects.
Rehm
in its
is
parasite
much
efFig. 102.
above
Lachnclla.
F, habit sketch G, ascus

;
and paraphyses.
After
Rehm.
on and larch but sometimes also on pine, both in Europe spruce and America. Ascophores upon cankers on branches and trunk of the host, very similar to those of the preceding species but with more evident stipe and paler disk; spores very minute, subglobose, 3 x 2-2.5 m;
It occurs chiefly
conidia 2 x
/z.
D. calyciformis
(d Wild.)
fir
Rehm
and
D. subtilissima (Sacc.) on
larch;
occurs on several conifers; D. abietis Sacc. on Picea.
Lachnella Fries
This
sessile
is
(p.
135)
and the spores usually 2-celled at maturity, and rows in the ascus. There are about forty species.
similar to the last genus but with the apothecia usually in two
The apothecia are brown L. pini Brun.^^ injures pine twigs. outside; the disc reddish-yellow with a white margin. Ascoma short-stipitate, 5 mm. in diameter, pale brown; disk light
146

fji;
orange-red with a pale margin; asci 109 x 8-9.5

6.5-8.5
fx,
spores 19-20 x
hyaline.
Hymenoscypha
Fries (p. 136)
This genus of over two hundred species is mainly saprophytic, one species only in its conidial stage being parasitic.
Ascoma
smooth;
spores
sessile
or short-stipitate, usually
hyaline;
enlarged,
conidial
asci cylindric to globoid, 8-spored;
elliptic,
blunt to pointed,
apically
in
paraphyses
hyaline.
filamentose,
P.
H. tumulenta
stage
as
&
D.^*'
its
Endoconidium, affects rye grain causing it to shrivel and assume poisonous The conidia are borne enproperties.
dogenously in the terminal branches of the hyphae and escape through an opening in the end of the branch.
Fig.
103.
Hymenoscyless globose,
Mollisiaceae
(p.
134)
or sunken in
first
pha. J, habit sketch; K, ascus and paraphyses. After Rehm.
Ascocarp free from the
first
the substratum and later erumpent, at
more or
becoming flattened;
asci 8-spored,
opening
by a slit; spores hyaline, 1 Above four hundred species.
to many-celled; paraphyses slender.
Key
to Genera of Mollisiaceae
I.
Ascocarp fleshy, waxy, rarely membranous. Ascocarps not sunken in the substratum Ascocarps on a visible, often radiate
Mollisiefle.
mycelium
Spores
elongate,
often
fusiform,
."
1-celled
filiform,
1.
many-celled Spores Ascocarps not seated on a visible celium

Spores 1-celled Spores spherical Spores elongate
2.
Tapesia. Trichobelonitun.
my-
3. 4.
Mollisiella.
Mollisia.

Spores becoming Spores elongate
2-cellccl
5. 6. 7.
147
Niptera.
filiform, 4-celled. ...
Belonidium.
Belonopsis.
Spores filiform many-celled Ascocarps at first sunken in the substra-
tum, later erumpcnt Ascocarps bright colored, only slightly

erumi)ent Spores eUispoid or elongate, rounded,
1-celled
8.
Pseudopeziza,
Fabraea,
p. 147.
Spores becoming many-celled
9.
p. 149.
Ascocarps
dark
colored,
at
length
strongly erumpent Spores ellipsoid or fusiform, 1-celled
Ascocarps bristly externally and on the margin Ascocarps externally smooth, the
10.
Pirottaea.
margin at most merely shredded

Spores
11.
Pyrenopeziza.
many-celled
by transverse
12. Beloniella.
septa
Ascocarps gelatinous
gristly,
horny when
II.
dry
Callorieae.
Of
this large
number
of
genera only two are important patho-
gens, several of the others are parasitic on non-economic hosts while others are saprophytic chiefly on decaying woody parts.
Pseudopeziza Fuckel
The genus comprises some ten species, all parasitic on leaves, them upon economic plants causing serious disease. The very small apothecium develops subepidermally breaking
several of
through only at maturity, light colored; spores 1-celled, hyaUne, in two ranks in the ascus; paraphyses somewhat stout, hyaline. Conidial forms are found in Gloeosporium, CoUetotrichum and Marssonia.
P. medicaginis (Lib.) Sacc.^'" ^The epiphyllous apothecia are in the older leaf spots, subepidermal at first but eventually breaking through.
Apothecia saucer-shaped, light colored, fleshy;
asci
clavate;
148
.spores hyaline,
10-14 ii long; paraphyses numerous, filiform. A Phyllosticta thought to be its conidial stage has been reported.^' On dead spots in leaves of alfalfa and black medick.
P. trifolu (Bernh.) Fcl. This is closely related to, perhaps identical with, the last species.
Sporonema (Sphaeronaema) phacidioides Desm.

its
is
conidial
form.
supposed to be This coalfalfa.
nidial stage has not however,
C3
been observed on
Ascocarps mostly epiphyllous, on dead spots, averaging

0.5
mm. broad, yellowish or brownish; spores elUptic 10Fig. 104. P, trifolii. Ascus and paraphyses; germinating spores. After Ches- 14 X 5-6 At.
Conidia in cup-shaped pycnidia which are numerous, small, light brown; disk cinnamoncolored; conidia ovoid-oblong, 5 m, bi-guttulate.
ter.
P. tracheiphila Miiller-Thurgau
^*
is
found upon the grape in

Conidia
(
Europe.
P. salicis (Tul.) Pot. occurs on Salix.
salicis).
=Gloeosporium
P.
ribis, Kleb.^^-^^
Apothecia appear in the spring on dead leaves of the previous

season; saucer-shaped, fleshy,
stalked;
asci
somewhat
spores hyaline, ovoid; paraphyses simple or branched,

clavate,
slightly clavate, rarely septate.
Conidial phase (=Gloeosporium ribis) on the leaves of the host forming an abundant amphigenous infection; acervuli stromatic; conidiospores
commonly
/x,
19 X 7 M) varying from 12-24 x 5-9 escaping in gelatinous masses.
Fig.
105,p. ribis. Apothecium in section. After Klebahn.
On
red and white currants less comin
monly on black currants and gooseberries both

America.
Europe and
The
ascigerous
'""'
Klebahn
in
stage of this fungus was demonstrated by 1906 to be genetically connected with what had been

earlier
149
known
as Gloeosporium ribis (Lib.)
Mont.
& Desm.
Old
filter
leaves bearing the latter fungus were wintered out-doors in
paper and in the spring were found with this aseigerous stage. ascospores were isolated, grown in pure culture and typical The ascospores also infected the host conidia were produced.
The
leaves
successfully
The
conidial stage
is
producing there the typical Gloeosporium. the only one ordinarily seen. The acervuli
are subepidermal elevating the epidermis to form a pustule which eventually ruptures and allows the spores to escape as a gelatinous whitish or flesh-colored mass. The spores are curved and usually
larger at one
end than at the other,
Fabraea Saccardo
This
is
(p.
147)
a genus of some ten species of small leaf parasites which much resemble Pseudopeziza but differ from it in its 2 to
4-celled spores.
F. maculata (Lev.)
The
perfect stage
is
AikJ'^ common on pear and quince
leaves which
have wintered naturally. 8-spored asci may be

in small elliptical areas.
When
seen
such leaves are wet the white
crowding through the surface
The
apothecium
is
paraphysate; the
Fig. 106.
spores hyaline and 2-celled. Conidial form ( = Entomo-
sporium maculatum) on leaves
and
fruits;
acervuli,
black,
subepidermal,
the
breaking away
spore
epidermis to expose the

hyaline
in
4-cells
18-20
mass; X 12
spores
/x,
maculata. 1, acervulus of stage in section; 3, spores. After South worth.

conidial
F.
cluster, the lateral cells smaller, depressed; stipe filiform 20 x 0.75 n; the other cells with long setae.
Atkinson ^^ proved the connection of the aseigerous with the conidial form by cultivating the conidia from the ascospores. The conidial form is very common and destructive on pear and quince leaves and fruit. The mycelium which abounds in the diseased spot is hyaline when young, dark when old. It collects to form a
150
On this the spores are produced on short erect conidiophores, Fig. 106; eventually the cuticle ruptures and the spores are shed. The spores germinate by a tube which
thin subcuticular stroma.
arises
from near the base
of a bristle.
F. mespili (Sor.) Atk. on medlar with the conidial form Entomosporium mespili (D. C.) Sacc. is perhaps identical with the above.
There are only minor and uncertain differences
in the conidial
Sorauer by inoculation with conidiospores produced on stage.^^ pear typical spots which bore mature pustules after an interval of
about a month.
Fries.
He
referred the fungus to the genus, Stigmatea
See p. 243.
Cenangiaceae
(p.
134)
Ascoma at first buried, later erumpent, on a stroma, dark, with a rounded or elongate disk; asci 8-spored; spores long or filiform, 1 to many-celled, often muriform, hyaline or dark; paraphyses branched forming a complete epithecium. About two hundred fifty species. Key to Subfamilies and Genera of
Ascocarps
coriaceous,
fresh
first
Cenangiaceae
corneous
or
waxy
I.
when
Dennateae.
Ascocarps without a stroma, at mersed.

Spores 1-celled
im-
Ascocarps externally bright colored,
downy
Ascocarps externally dark Ascocarps smooth; spores hyaline.
Ascocarps downy; spores colored. Spores 2 to 4-celled, elongate
Spores hyaline
.
1.
Velutaria.
2. 3.
Cenangium,
p. 151.
Schweinitzia.
Spores always 2-celled; ascocarp
smooth
Spores 2
to
4-celled;
4.
Cenangella.
ascocarps
5.
externally Spores at length brown or black Disk elongate with a thick rim.
downy
Crumenula.
Tryblidiella.
6.
Disk rounded
Rim Rim
thin; spores 2-celled
7.
.
Pseudotryblidium.
Rhytidopeziza.
involute; spores 4-celled.
8.

Spores many-celled, filiform Ascocarps springing from a more or developed stroma Spores
9.
151
Gordonia.
less
8, not sprouting in the ascus. Spores sprouting in the asci which become filled with conidia
. .
10.
Dermatea,
Tympanis.
Bulgarieae.
p. 152.
11.
II.
Ascocarps gelatinous when fresh Ascocarps sessile or stalked, with smooth,

saucer-shaped disc
Spores 1-celled, round
12.
Pulparia.
Spores 1-celled, elongate
Ascocarps
sile,
soft, gelatinous inside, ses-
thin
soft,
13. Bulgariella.
Ascocarps
thick
gelatinous, stalked,
14. Bulgaria, p. 152.
15.
Ascocarps watery gelatinous

Spores 2-celled Spores
Sarcosoma.
unequally 2-celled rounded

16.
.
at the ends
Spores elongate, acute at the ends.
17.
18. 19.
Paryphedria. Sorokina.
Spores filiform
Spores muriform Ascocarps with convolute tremelliform
discs
Holwaya. Sarcomyces.
Spores 1-celled, hyaline

Spores muriform, blackish
20.
21.
Haematomyces. Haematomyxa.
so far as known saprothat further study will reveal some of phytes though probable them as weak parasites or possibly as destructive ones.
With few exceptions these genera are

it is
Cenangium
Fries (p. 150)
Parasitic or saprophytic chiefly in bark, the apothecium developing subepidermally and later breaking through to the surface; sessile, light colored without, dark within; asci cylindric-globoid,
1 or rarely 2-celled, hyaline or brown, in one row; paraphyses colored. About seventy species. C. abietis (Pers.) Rehm.^^ has caused serious epidemics upon
8-spored; spores ellipsoid,
pine in Europe and America.
Ascoma dark-brown, erumpent,

10-12 X 5-7
M.
clustered;
spores
ellipsoid,
152

Conidia ( = Brunchorstia destuens Erikss.) in pycnidia which are partially embedded in the host,
the
smaller
simple,
the
larger compound, 1-2

in diam.; spores
mm.
30-40 x
Fig. 107.
Cenangium, habit sketch,

paraphyses.
After Tulasne.
asci
and
II, tapering-rounded at each end, 2 to 5-septate. A second conidial phase
= Dothichiza
ferruginosa
Sacc.) has simple spores, C. vitesia occurs in conidial
form as Fuckelia on Ribes.

Fries (p. 151)
Dermatea
genus of over sixty species some of them parasitic. species conidia in pycnidia are known.
Ascocarps scattered or clustered, black or brown; asci small, thickwalled,
lipsoid
In
or
many
not,
stromate,
sessile
8 or 4-spored; spores elor spindleform, 1-celled, 4

to
6-celled,
becoming
brown,
2-ranked; paraphyses septate, apically enlarged and colored.

(Pers.) Rehm. is wound parasite oh the hornbeam and beech; D. cinnamomea (Pers.) Rehm. on oaks; D. acerina
D. carpinea
Karst,
on maple (Acer pseudo;
platanus)
its
all
in Europe.
D. prunastri (Pers.) Fr., with conidial form Sphseronema spurium Fr. is found on Bark various species of Prunus, in Europe and America.
Bulgaria Fries
(p.
Fig. 108. ermatea. ^, habit sketch; C, ascus and paraphyses. After Tulasne and Rehm.
of
151)
gelatinous apothecium is rather large and dark colored; asci 4 to 8-spored; spores 1-celled, elongate, brown.
The
There
is
one species worthy of mention.
153
B. polymorpha (Ocd.) Wett."-' ""^ is a common saprophyte on bark. It is said to sometimes become parasitic. Ascocarps black,
stipitate; disk scarcely
cupped, ranging up to 4 cm. in diameter
although usually smaller.
Caliciaceae (p. 134)
Stroma
without
more
algal
or
less
thalloid,
cells,
often
with or rudimentary and
inconspicuous; ascoma more or less globoid, the apex of the ascus dissolvstipitate;
the spores are matured, thus the hyaline unripened spores to esallowing cape and mature afterwards.
ing
Q O Q
Q
I
before
This
lichens
less
small
family
(less
than
one
hun-
dred twenty-five species)

of
contains
the only
phytopathological importance, unon

2,
\
pallida
pear on
ered.
the foliose lichens which sometimes appoorly kept fruit trees be consid-
Grape Root. Ascus. After
Massee.
Key to the Gexer.\ of

Ascoma with a long
stalk
Caliciacese
Spores spherical, or subspherical Spores colorless or only

Spores elongated, septate
slightlj' colored.
1.
Spores brown or brownish
2.
Coniocybe, p. 153. Chaenotheca.
Spores elongate
elliptic or egg-shaped, usually two-celled Spores elliptic to sj)indlc-form, 4 to
3.
Calicium.
8-cclled
4.
Stenocybe,
Ascoma
short stalked
5.
Spores 2-celled Spores globose, 1-cellcd
Acolium.
Sphinetrina.
G.
Coniocybe pallida (Pers.) Fr. is generally distributed throughout Europe and America, commonly on the bark of various forest trees and upon the crown and roots of the grape. The parasitic
154
THE
PÛNGI

22
The entire height of the asnature of the fungus is in doubt. cocarp is 2 mm.; head white, then grayish brown; asci cyhndric, 8-spored; spore tinged with brown, 4-5 /x in diameter. The species
&
as a pathogen is usually referred to as Roesleria hypogsea Thiim ^^ folPass, and given a place in the Geoglossacese; but Durand
Fig. 109.
lows Schroter in excluding the species from that family.
Phacidiales
(p. 124)
This order, comprising some six hundred species only a few of which are pathogens, is characterized as follows: mycelium well
multiseptate; ascocarps fleshy or the substratum or in a stroma, rounded or stellate, for a long time enclosed in a tough covering which at maturity becomes torn; paraphrases usually longer
developed,
leathery,
much branched,
or
free
sunken
in
than the
asci,
much branched, forming an

Key
epithecium.
to Families of Phacidiales
1.
Ascocarps soft, fleshy, bright colored Ascocarps leathery or carbonous, always black
Stictidaceae, p. 154.
Ascocarps at
first
sunken, later strongly

2.
erumpent, hypothecium tliick Ascocarps remaining sunken in the substratum, hypothecium thin, poorly
Tryblidiaceae, p. 155.
developed
3.
Phacidiaceae, p. 155.
Stictidaceae
The members
hundred
fifty
of this family
(about twenty genera and two
usually considered saprophytes, although one species of Stictis has recently been described as a
species)
are
parasite.
Stictis
Persoon
Perithecia sunken, pilose, at length erumpent; asci eylindric,
containing eight filiform, multiseptate spores; paraphyses filiform,

richly branched apically. or more species of the
155
Of
the
seventy
only one, S. panizzei d Not., originally described from fallen olive leaves in Italy, has been
genus
charged with producing disease.^'' It has within the last few years become very destructive in Italy.
The
Tryblidiaceae,
species,
with
are
six
genera
and
some seventy
likewise
chiefly
saprophytes with the possible exception of the two genera HeterosphKria and Scleroderris.
The former
the
of
latter
occurs
on umbellifers
the
perfect
while
stage
may
contain
certain
currant
and
gooseberry
fungi (Mastomyces and Fuckelia) of Europe as well as a European parasite of the willow.
Fig. no. Stictis. D, habit sketch, E, ascus and paraphyses. After
Phacidiaceae
(p.
154)
less
erumpent, disk-like or elongate, single or grouped, leathery or carbonous, black, firm, opening by lobes or rifts.
Apothecia sunken, more or
Rehm
Key to Genera of
Phacidiaceae
Apothecia not inseparably united to the substratum

Spores elongate, hyaline, 1-celled
I. 1.
Pseudophacidieae.
Pseudophacidium.
Spores elongate, spindle-form or

multicellular.
filiform,
Spores elongate to filiform, not muri-
form
Apothecia rounded, opening by a rounded mouth
Spores
elongate or spindle-form paraphyses, none
2.
Dothiora,
p.
156
Spores elongate-globoid, 2-celled;
paraphyses present
3.
Rhagadolobium.
156

Spores
needle-like;
paraphyses
4.
present
Coccophacidium.
Clithris, p. 157.
Apothecia elongate; opening by a

slit
5.
Spores elongate, muriform, with pa-
raphyses Apothecia firmly united to the substratum Apothecia separate, no stroma
6.
.
Pseudographis.
Phacidieae.
II.
Spores ellipsoid or globoid, Spores 1-celled
to 4-celled
Apothecia rounded Paraphyses not forming an epithecium
7.
Phacidium,
p. 157.
Paraphyses forming an epithecium

Apothecia
opening
irregular,
8.
Trochila, p. 157.
elongate,
irregular
9.
by
an
mouth
Spores 2 to 4-celled
Spores hyaline
Cryptomyces,
p. 158.
Apothecia rounded, spores 2 to

4-celled
10.
Sphaeropeziza.
Apothecia
2-celled
elongate,
spores
11. 12.
Schizothyrium.
Keithia.
Spores brown
Spores
filiform
or
needle-like,
to
13.
many-celled
Coccomyces.
Apothecia collected on a stroma, opening

elongate
Spores 1-celled, hyahne Spores ovate Spores filiform or needle-like Spores 2-celled Spores hyaline Spores brown
14. 15.
Pseudorhytisma. Rhytisma, p. 158,

Marchalia.
Cocconia.
16. 17.
Dothiora Fries
(p. 155)
There are about ten wood-inhabiting species. Ascocarp at first sunken in the substratum, later irregularly erumpent; disk black;
asci
clavate,
8-spored; spores elongate or spindle-form,
many-

celled
157
or muriform,
hyaline
or
slightly
yellowish;
paraphyses
wanting.
D. virgultorum Ft. grows on birch.
Clithris Fries (p. 156)
A small genus of about twenty species found on wood and bark; mainly saprophytes. Ascoma sunken, then erumpent, elongate, with lip-like margins, dark colored; asci clavate,
8-spored, often blunt pointed; spores linear or
spindle-shaped,
multicellular;
paraphyses
fili-
form, coiled apically, hyaline.

C. quercina (Pers.) Rehm. is found on oak branches and is perhaps identical with C. aureus Mass. on willows. C. juniperus is found on liv-
ing juniper.
Phacidium
Fries (p. 156)

chiefly
Over seventy species

herbaceous
stems.
on
leaves
or
*
and
paraphyses.
^
soon
.,.,,,. becommg
Ascoma
lenticular,
rift; asci
single,
flattened, Fig. in. Clithris. Ascus with spores
breaking
open
by
an irregular
^^
'
clavate, 8-spored; spores
ovate or spindle-shaped, hyaline, 1-celled; paraphyses thread-like, Conidial form probably in part =Phyllachora. hyaline. P. infestans Karst. is a parasite on pine leaves.
Trochila Fries
(p.
156)
Perithecia sunken and closed, later erumpent, black, leathery;

asci clavate 8-spored; spores long, hyaline,
1-celled;
paraphyses
filamentose forming an epithelium. Fig. 112. T. popularum Desm. is thought by Potebnia
gcrton
-^^
to
be the ascigerous form

is
of
and EdMarssonia castagnei

-^"^
D.
& M.
the ascigerous form of Gloeosporium para-
T. craterium
doxum.
See p. 541.
158
Cryptomyces Greville
(p. 156)
genus of some ten species living on large black blotches.
wood
or leaves, forming
Ascoma sunken
1-celled,
in the substratum, flattened,
erumpent, irregu-
lar in outline, coal black; asci clavate, 8-spored; spores elongate,
paraphyses
(Fr.)
filifoim.
Rehm is a parasite on willow and dogwood twigs in Europe and America, forming large carbonous areas under the bark.
C.
maximus
Rhytisma
Fries (p. 156)
belong about twenty-five species which cause very conspicuous, though but slightly injurious, black leaf-spots. The spots which are white within, are due to sclerotial
cushions formed in the host tissue.
l/ll
To Rhytisma
of
the
leaf
occurs
in
the
infected
part.
Thickening One-
celled
conidia
(Melasmia form) are abundantly
produced
in pycnidia early in the season, followed
by sclerotium formation. Much

^nwvii
later, usually well into winter or the following spring, the apothecia
Fig.
several species of which the asco-spores are una n d %araphy- known have been referred here,
112.
Tro-
appear.
Besides the asco-spore-producing forms
ses.
After
Ascoma on a
is
sclerotial stromatic layer,
which
opening by
black above, white within; ascocarps elongate, a lip-like slit; asci clavate, often blunt pointed,
8-spored; spores filiform or needlelike, hyaline, mostly 1-celled, lying parallel and lengthwise of the ascus; paraphyses filiform,
hyaline, often arched above. R. acerinum (Pers.) Fr,
The spot is at first yellow and thickened and in this stage bears The apothecia numerous conidia upon short conidiophores. ripen in spring and rupture by numerous irregular fissures which follow the ridges of the wrinkled surface. Klebahn secured infection by ascospores resulting in three weeks in yellow spots and in eight weeks in conidiospores. The conidia are supposed to aid in
159
spreading the fungus during the summer though they have not yet actually been observed to germinate or to cause infection. Apothecia radiatcly arranged on the stroma which is about
0.5-1.5
cm.
across;
asci
120-130 X 9-10 m; spores 65-80 x 1.5-3 m; large,

paraphyses numerous, curved or hooked. Conidia
(
in-
Me
acerina Lev.) preceding the asci, producing numerous

small,
hyaline,
in
l-ccUed
spores
an extended hyspecies
of Fig.
m^mm^ >âr vior
;#^'^r
^'^^'^-^^^
>?j-
menial layer. On various

maple,
apparently
consist-
113. R. acerinum. F, conidial layer; E, ascus and paraphyses. After Tulasne.
ing of races since in different localities the host differs without a crossing over of the fungus. R. punctatum (Pers.) Fr. also occurs on maple, especially Acer
pseudoplantanus.
its
It
may
be distinguished from the preceding by
small, speck-like stromata. R. salicinum (Pers.) Fr. is
America.
It is quite similar in external
found on willow in Europe and appearance to R. acerina
except for the smaller average size of the spots. R. symmetricum Mull, is another willow inhabiting species.
The apothecia
are amphigenous and are said to mature in autumn on the still live leaves. Other species are common especially on various Ericaceae and Coniferse in Europe and America.
Hysteriales
(p.
124)
Small species with elongated, black, covered apothecia which open by a long narrow slit exposing the hymenium asci 8-spored
;
spores usually long and slender. Some few are leaf parasites but most are wood saprophytes. Pycnidia are found in some species.
The order
serves as a bridge between the Discomycetes Pyrenomycetes. About four hundred species.
and the
160
Key
to Families of Hysteriales
Ascocarps immersed; walls of the ascocarps connate with the membranous covering
1.
Hypodermataceae, p,
160.
Ascocarps at
walls free
first
immersed, erumpent,
.
Walls membranous or coriaceous, black Walls thick, almost corky, gray or

black
2.
Dichaenaceae, p. 162.
Ostropaceae.
3.
Ascocarps from the first free Walls carbonous, black; shield-shaped,

round,
linear
oval
or
more
or
commonly
4.
Hysteriaceae,
p. 163.
Walls
horny, ascocarps vertical, clavate
membranous
brown,
5.
Acrospermaceae.
The
third
and
fifth families
contain no pathogens.
Hypodermataceae
Ascocarp flattened, rounded or elongate, rarely branched, united to the substratum; opening by a slit; asci 4 to 8-spored; paraphyses apically branched, the branches forming an epithecium,
or
hooked or crimped.
About
fifty
species,
chiefly
saprophytes.
Key
Spores 1-celled or
to Genera of Hypodermataceae
Spores elongate, rather broad

l^y
cross walls 2 to
many-celled
Spores 1-celled Asci 8-sporcd, spores spindle-form
Spores hyaline
1.
Henriquesia,
Farlowiella.
Spores brown Asci 4-spored, spores hyaline Spores 2-celled, h3âline Apothecium black
2. 3.
Hypodennella,
p. 161.
4. 5.
Hypoderma,
Angelinia.
p. 161.
Apothecium red

Spores 4 to many-celled, spindle-form
Spores 4-celled, mostly hyaline Spores 4 to many-celled, brown Spores niuriform, hyaline Spores
filiform, 1-celled
G.
161
Gloniella.
7.
Rhytidhysterium.
Hysteropsis,
p. 161.
p. 161.
8.
9.
Lophodermium,
Of these genera only four are important here.
Hypodermella Tubeuf
This
differs
(p.
160)
from the next genus
in its
spores; asci 4-spored. Two species, both European H. larius Tub. affects larch needles in Europe. ^^
pyriform unicellular and economic.
H. sulcigena Link
is
on pine needles.
(p.
Hypoderma De CandoUe
160)
Apothecia oblong, opening through a thin black cover by a

long fissure; asci 8-spored; spores cylindrical or fusiform, 2-celled at maturity; paraphyses hooked at the end.
H. desmazieri Duby,^^^ on pine needles

rope.
in
America and Euspores

hyaline,
leaf
Amphigenous;
linear-elliptic,
asci broadly" clavate, obtuse and 2-rowed.
sessile;
H.
fall in
laricis,
H.
strobicola,
H.
pinicola,
produce premature
various conifers.
Hysteropsis
Rehm
Asci clavate, 8-spored; spores hyaline, muriform; paraphyses
branched, forming an epithecium. H. brasiliensis occurs on cacao trees.
Lophodermium Chevall
Spores long, thread-like, continuous; conidiospores in pycnidia. L. pinastri (Schr.) Chev. occurs in Europe ^~ and America on Pinus sylvestris especially on young plants causing the leaves to
fall.
The
first
pearing until the
year pycnidia only are formed, the asci not apsecond year.^^
162

leaf,
Ascocarps scattered on the
shining black, up to
mm.
long; asci clavate, 8-spored; spores nearly as long as the ascus,
90-120 X
L.
1.5
/i.
Conidia cyhndric, hyaUne,

1
;u.
continuous, 6-8 x
brachysporum Rost.
stalked,
Perithecia epiphyllous; asci cylindric, shortapex rounded, 120 x 20-25 fi, 8-
spored;
spores
paraphyses
oblong,
bacillar,
apex
1-rowed,
hyaline,
^^^
28-30
curved; x
9-10
It
yu.
is
common on
Several
pine leaves. other species are parasitic
upon
various conifers,
rium (Hart.) Europe and America; L. nervisequum (D. C.) Rehm, on fir leaves, a very destructive
Fig. 114. L. pinastri. H^ habit sketch; J, ascus and paraphj-ses.
among them: L. macrospoRehm, on spruce leaves, in
European
d
species;
the
Septoria pini Fuckel;
After
pines; L.
larch.
Not, on juniper Europe and America; L. gilvum Rost., on abietis Rost., on spruce leaves; L. laricinum Duby, on
Rehm.
last four species are
pycnidial stage is L. juniperinum (Fr.) leaves and twigs in
The
European.
160)
Dichaenaceae
(p.
This family contains the single genus Dichaena.
Dichaena Fries
Apothecia grouped in rounded spots; at sunken, then erumpent, rounded or elongate, dark brown; asci irregularly pyriform, 4
first
^m\i'
mrih\Mk
êsg,^t^}]l'-^\:])^^^^
to 8-spored; spores ellipsoidal, at first 1-celled, ^^^^fesn^^y^ at maturity multicellular; paraphyses filiform, îq. ii5. Hysterium Some seven species are found upon various macrosporum. Af^ ^
ter Hartig.
trees.
D. quercina Fr. causes rough black patches on bark of young oaks in Europe and America; D. faginea Fr., a similar effect on
beech.

Hysteriaceae
Ascocarps
free,
163
(p.
160)
seated upon the substratum,

straight,
elongate or linear,
curved or even
fila-
branched, disk-form, boat-shaped or band-like,

black; asci usually 8-spored; paraphyses mentous, often forming an epithecium.
About fourteen genera and some two hundred fifty species, many but poorly known. Several genera contain plant patho., r gens, but they are not often of economic mi,
.
^^-
n^-
D.
quer-
1-,
cina.
Ascus and
paraphyses. After
^"^^
portance.
Key
Ascoma
to Genera of Hysteriaceae
linear, flattened, broadly sessile Spores ellipsoid or spindle-shaped, many-
celled
Spores 1-celled, 16 in each ascus
1.
Cyclostomella.
Spores
2-celled,
sometimes
4-celled,
ellipsoid or elongate
Spores hj'aline Asci 8-spored, spores 2 to 4-celled Paraphyses scarcely branched.

.
2.
Aulographum.
Glonium.
Hariotia.
Paraphyses forming an epithecium

Asci many-spored, spores 2-celled
ing fungi
.
3.
4.
Spores colored, 2-celled; leaf infect-
Paraphyses present
Ascoma seated on a cottony

stroma
5.
Lembosia.
Ascoma
radial,
on a
8
circular
stroma
Spores 2-celled, ascus
ularly circular
in
each
6.
Parmularia.
Hysterostomella.
Paraphyses absent, stroma
irreg7.
Spores 4 to 8-celled, elongate or spindle-
form
Spores
hyaline,
celled
spindle-form,
48.
Hysteroglonium.
164

Spores
brown,
elongate,
to
9.
8-cellecl
Spores elongate, muriform Ascoma boat or band-shaped, not

celled
10.
sessile
Hysterium, Hysterographium,
p. 164.
Spores spindle-formed, brown, many-
Spores 4 to 8-celled; asci 8-spored.

Spores many-celled; asci 4-spored.
Spores filamentose, hyaline or yellow
11. 12.
13.
Mytilidium. Ostreion.
Lophium.
Actidium.
Ascoma
stellate
14.
Hysterographium Corda
Asci clavate, 8-spored; spores muriform, dark colored
mature; paraphyses branched forming an epithecium.

seventy species.
when About
H. fraxini
(Pers.)
de Not. occurs on Oleaceae, particularly on
Fig.
17.
fraxini.
H. Ascus
Fig. 118.
and paraphyses. After Rehm.
Gymnoascus, sexual organs.

After Dale.
the ash, perhaps only as a saprophyte.
It is
found both
in
Europe
and America.
Aspergillales
(p.
124)
The
asci are
Aspergillales
are
clearly
distinguished
Ascomycetes by the possession
of a closed ascocarp in
from the other which the
The forms with the
not collected in a hymenium but are irregularly scattered. least developed peridium are evidently related
165
to the Endomycetacesc; the forms with a more highly developed pcridium, to the Pyrenomycetes, particularly to the Perisporiales. Conidial forms are usually present, indeed in many cases they
which
preponderate almost to the entire exclusion of the ascigerous form may be seen only under very exceptional conditions.
Sexual reproduction has been demonstrated in several families. In the Gymnoascacese (Dale ^^ and Eidam ^^) there are usually two twisted branches (Fig. 118) which conjugate. These branches
are multinucleate at the time of fusion.
The ascogonium
de-
described on pages 116In the Aspergillacea? similar sexual organs are formed but 117. parthenogenesis or a much reduced form of fertilization is often
velops from
this fertilization
much
as
is
met.
In
all,
the species
number two hundred
fifty or
more.
Key to Families of
Aspergillales
.
Pcridium made up of loose floccose hyphse Pcridium compact, closed Ascocarps mostly small, not subterranean sessile without mostly Ascocarps stroma; peridia remaining closed.
.
Gymnoascaceae.
2.
Aspergillacese, p. 166.
Ascocarps mostly stalked peridia opening at maturity by lobes, or ir;
regularly
3.
Onygenaceae.
Ascocarps sessile, the spores issue in columnar masses from the gobletAscocarps
shaped peridia sessile on a small stroma.
4.
.
Trichocomaceae.
Myriangiaceae,
p. 170.
5.
Ascocarps mostly enlarged, tuberous, subterranean.
Peridium
clearly
distinct
;
from
the
walls of the ascocarp spore masses powdery at maturity
6.
Elaphomycetaceae.
Peridium not clearly limited, continuous with the walls of the ascocarp;
spore masses never powder)'
7.
Terfeziaceae.
Of thpse the second and
fifth families
only contain pathogens.

fifteen species are
The Gymnoascacese
of five genera
and some
found on manure, and other organic matter.
The
third
and fourth
166

the third on hoofs, horn, etc.; the less subter-
families are monogeneric;

sixth
is
subterranean and the Terfeziacese more or
ranean.
Aspergillaceae
aseocarp, in
(p.
165)
The is a small spherical many or tuber-shaped body, usually indehiscent, rarely opening by a pore. The spherical or pyriform asci bear from 2 to 8 spores which may be from 1 to many-celled. The aseocarp is in some genera
forms but rarely seen,
provided with appendages which strongly resemble these of the Erysiphacese (Microascus). Conidia are produced in great abundance.
In Aspergillus and Penicillium fertilization is said by some observers to be accomplished by conjugation of a straight oogonium with a spirally coiled antheridium, this act resulting in an
Dale ' (see also Fraser and in one species of Aspergillus Chambers ^^) which she studied, though sexual organs were often present, and predicates a reduced form of sexuality consisting of fusion of the nuclei of the ascogonium with each other.
ascogenous hypha.
Recent work denies such fusion
of
Key to Genera of
Aspergillaceae
Spores l-celled Perithecium flask-shaped, beaked or papillate

1.
Microascus.
Perithecium not beaked
Perithecium with hair-like appendages peridium compact, mostly dark

colored
Appendages straight
a hairy
felt
hairs or forming
2. 3.
Cephalotheca.
Appendages apically coiled hairs. ... Perithecium unappendaged peridium

;
Magnusia.
membranous
or fleshy
Conidia borne directly on the mycelium

Chlamj^dospores borne in chains. Chlamydospores borne singly
.
4. 5.
Thielavia, p. 167. Rostrella, p. 168.

Conidia borne on distinct conidiophores Conidia borne
conidioat
right
6.
167
singly;
phores
angles
branching
in chains
Aphanoascus.
Conidia borne
Conidiophores
sinijÎc,
aggre7.
gated into bundles
EmericeUa.
Conidiophores enlarged apically

bearing
numerous
sterig-
mata
Sterigmata simple
St<?rigmata branched
8. 9.
Aspergillus, p. 168.
Sterigmatocystis.
Conidiophores,
sympodially
10. Eurotiopsis.
branched
Conidiophores bushy branched Conidiophores
Conidiophores
single,
peri11.
thecia sessile
in bundles, api-
Penicillium, p. 169.
cal cells swollen, thecia stalked
peri12. Penicilliopsis.
stel-
Spores 2-celled; peridium at maturity

late
13.
Testudina.
Of the thirteen genera and some one hundred to two hundred

species only four of the genera are of interest here. occur on rotting leaves, manure, etc.
^--''
The
others
Thielavia Zopf
(p.
166)
T. basicola (B. and Br.) Zopf. This, the one species of the genus, is on the boundary between the Aspergillales and the Perisporiales and is classed by some with
the one, by some with the other order. The ascocarps, which are the form less
commonly
seen, are
round, brown, completely closed and have no appendages. The asexual spores are of two kinds. First: hyaline conidia produced
endogenously within "pistol-formed" conidiophores from the ends of which they are expelled. Second: short cylindrical conidia or chlamydospores with a thick brown wall; borne in series of
168

These
three to six on the ends of hyaHne branches, Fig. 119. conidia fall apart as they age.
The hyaline conidia preponderate in early disease, giving the surface of the root a mildewed appearance; the dark conidia preponderate root with
later,
covering the black coating.

is
Finally, after the host
dead,
the ascocarps appear.
The
lium
tissue.
delicate hyaline
wanders
mycethrough the
affected root disorganizing its
The
superficial
myceasci
lium
is
lightly tinted.
Perithecia 80-100
ovate,
ticular,
n;
8-spored;
spores
len-
vacuolate,
1-celled,
chocolate-colored,
8-12
4-5
25-65 fx long; conidia hyaline about 10-20 X 4-5 /x. In Europe and Eastern North America on Aralia quinquefolia. Begonia rubra, Begonia sp., Catalpa speciosa. Cyclamen sp., Gossypium herbaceum, Linaria canadensis, Lupinus angustifolius, L. albus, L. luteus, L. thermis, Nasturtium armoracea, Nemophila auriculata, Nicotiana tabacum, N. rustica, Onobrychis cristagalli, Oxalis stricta,
Fig. 119. Thielavia basicola, showing two conidial forms and ascus and ascospores. After Van Hook and Zopf.
#^<x.fl.-^j*5.
O^t^^
chlamydospores in chains, at maturity separaabout ting, short-cylindric, '5~8 X 12 yu; the entire group
ju;
Phaseolus vulgaris, P. multiflorus, Pisum sativum, Senecio elegans, Trifolium repens, Trigonella coerulea, Vigna sinensis and Viola odorata.
Rostrella coffeae
coffee in Java.^^
Zimm.
is
described as the cause of canker of
Aspergillus Micheli
(p.
167)
are small, spherical, indehiscent, smooth bodies which at maturity are entirely filled with 8-spored asci; spores
The ascocarps
THE
FUxXGI
109
The conidiophores, which serve better to characterize l-coUcd. the genus, are swollen at the end, and bear numerous sterigmata (Fig. 120) on which the spores are borne basipetally in chains.
Sclerotia are sometimes formed.
The members
of the
genus are
all
saprophytes but some of them
Fig. 121.
Penicillium,
a,
A
Fig.
120.
gillus,
showing
Aspern i d i o-
phore;
s,
a conidioproducing
c o
phore.
King.
After
chains of conidia, c; three spores more a g n i fi e d. highly After Longycar.
on
cause injury to fruit in the tropics; for example, A. ficuum, Reich. figs; A. phcenicis Pat. & Del. on dates.
Penicillium Link
'*'
(p.
167)
The ascocarp
It
is
may 8-spored. intervention of a sclerotial stage. serves to distinguish the genus by
as in the last genus, with the asci 4 to develop directly from the mycelium or with the
much
its
The characteristic conidiophore mode of branching. Fig. 121.
Instead of being apically swollen as in the preceding genus it branches repeatedly, the branches bearing terminal sterigmata and giving the conidiophore the appearance of a brush; hence the
name.
For species see page 573.
170
Myriangiaceae
(p.
165)
Perithecia numerous upon or in a stroma; asci in a pseudoparenchymatous substance witiiin the perithecium; spores muri-
form.
Key to Genera
Stroma Stroma
of Myriangiaceae
1.
valsoid, perithecia superficial. ...

effused, perithecia
Myriangium.
Myriangiella, p. 170.
300
immersed.
...
2.
Myrangiella orbicularis Zimm. parasitizes coffee in Java

62
Pyrenomycetes
Tlie four following orders are usually grouped together as the Pyrenomycetes; separated from the preceding forms by their closed ascocarp with the asci arranged in a hymenium. They constitute a vast assemblage of more than ten thousand species, the large majority saprophytic and unimportant except in the general
economy
as scavengers.
Perisporiales
(p.
124)
The present
order
is
characterized
by
its
almost universal
parasitic habit, the evident mycelium and the globoid perithecia without ostioles, or in one family flattened, ostiolate perithecia.
The mycelium
conspicuous.
is
superficial
upon the host and frequently quite
Key to Families of
Perithecia
Perisporiales
mostly
spherical,
imperforate
1.
Mycelium white;
perithecia with append-
ages External mycelium dark colored or wanting, perithecia without true appendages, but
Erysiphaceae, p. 171.
sometimes surrounded by
2.
appendage-like hyphae
Perithecia
late
Perisporiaceae, p. 189.
flattened,
shield-shaped,
ostio3.
Micro thyriaceae,
p. 195.

"'''''
'
171
Erysiphace^
This family on account of
pUcity of structure,
(p.
170)
its
abundance everywhere,
its
sim-
possession of typical ascigcrous and conidial stages forms a favorite type for introductory study of the
its
and
Ascomycetes.
Its
members
are easy of recognition, form-
ing a coating of white conidia,
conidiophores and mycelium upon the surface of its hosts
and giving them an appearance much as though they had

been lightly dusted with flour. Later in the season the white
patchesliberally
are
more
or
less
sprinkled
with
leading
the
to
Fig. 122. I, E. graminis, showing branohing haustoria. 33, Phyllactinia, intercellular hyphsD. After Smith.
black
the
perithecia
common name powdery An important list mildew.

of
the economic forms and their hosts has been published by
Halsted."
The mycelium except

is
in Phyllactinia
is
is
entirely superficial.
It
usually quite hyaline and

It
uninucleate.
cells
branched, septate and its cells fastens to the host and penetrates its epidermal
Figs. 122, 123.
(1) (2)
by uninucleate haustoria which by their various lobings aid

those those
in specific characterization.
may be grouped in three general classes; arising directly from the lower surface of the mycelium;
Haustoria
arising at the side of the
(3) arising from more mycelium. The relation of the haustoria to the host cells has been extensively studied by Smith. "^ The conidia arise in basipetal succession on simple scattered
as small semicircular processes; or less deeply-lobed lateral swellings of the
mycelium
conidiophores
(Fig.
129);
are
hyaline,
oval
or
barrel-shaped,
size
smooth, 1-cclled. Neger has shown that they vary greatly in with nutrition conditions.^"
air,
Conidia germinate readily at once in dry air, better in humid Haustoria are producing from one to three germ tubes.
172
THE
FUxNGI
formed at once and the mycelium develops to a more or less circular colony, producing new conidia in a few days. Artificial inoculations on susceptible plants, using conidia, usually result within two to five days in typical mildew spots.
Neger,^
who
shown that
light hastens the
studied the germination of conidia extensively has growth of the germ tubes, which in
many cases are negatively phototropic. Contact stimulus leads to the growth of appres-=/^
123. Erysiphe
soria.
The
what
rîG.
perithecia are subspherical, often some-
when young, dark to black and reticulated when mature; toria. After Salmon are without ostiole but are provided with of various types. Figs. 130, 133-136, which give main appendages characters to mark the genera. The appendages serve by hygroflattened, white to yellow
showing lobcd
haiis-
movements to aid in the distribution of the fungus.^^ The ascospores become free after dissolution of the perithecium by weathering. The asci are either solitary or quite numerous within
scopic
the perithecium and bear two to eight hyaline spores each. The conidia are short-lived summer spores. The perithecia
mature more slowly and constitute the hibernating condition. In some instances the ascus-form is unknown;
the fungus
is
then
classified
solely
by
its
conidial stage
and
falls
under the form genus
Oidium
(see p. 569.)
^-
In Sphaerotheca
an antheridial and an
oogonidial branch, each uninucleate, are de-
nium
its
veloped, and cut off by septa. enlarges; the antheridium

'
The oogolengthens,
_
fig.
124.Alcogoniumwith
five nuclei.
nucleus divides, and a septum is run in ^ separating the stalk cell from the antheri-
\"be After Harper.
dium.
The sperm
nucleus
enters
the
oogonium
and fuses
with the oogonial nucleus. Simultaneous with fertilization occurs, from the stalk cell of the oogonium, the development of a sterile system of enveloping threads which surround and protect the fertilized oogonium and eventually mature into the
sporocarp.
The
fertilized
versely producing a series of
oogonium divides several times transcells, one of which is binucleate.
THE FUNGI WHICH

This binucleate one ascus
cell
CAUSP]
PLANT DISEASE
173
after fusion of its nuclei develops into the
The ascus nucleus by of the genus. the spore nuclei and the spores are cut out of the periplasm by reflexion of the astral rays. In Erysiphe^^ the oogonium and antheridium arise in a very
characteristic
division gives rise to
similar way, the oogonium being somewhat curved. Fertilization After is also similar consisting of the union of two gametic nuclei.
fertilization the
oospore nucleus divides and the oogonium de-
velops into a short bent tube, which contains from five to eight nuclei. Septa now appear cutting off cells, some uninucleate, some with two or more nuclei.
The ascogenous hyphse develop

two
a knot and soon divide into or three cells each and
give rise to the asci which are in the beginning binucleate.

the OOgOIn Phyllactmia and fertilinium, antheridium
84
î''-
,r tû .12^Pnyllactinia, male and female branches; uninucleate oogonium and ^"theridium. After Harper.
zations are as in Erysiphe, though the
oogonium may be quite

the anther-
curved so as to
idium.
make almost a complete turn around
Fig. 125.
After fertilization the antheridium degenerates and enveloping protective hyphse arise both from the oogonium and the antheridium
stalk cells.
The oogonium becomes
three to five nucleate and
develops to a row of cells of which the penultinate cell has more than one nucleus. The ascigerous hyphae arise from this binucleate
cell,
perhaps also from other
cells of
the
series,
become
septate and form the asci either terminally, laterally or interThe young ascus is binucleate, fusion follows and the calary. spores develop as in the preceding genera.
The family
and eleven
hundred
dred species.
contains, according to Salmon,
forty-nine species
varieties, according to
Saccardo more than one hunThese are parasitic on some one thousand five
hosts,
some
of
them upon economic
plants
is
and
of
serious harmfulness.
The matter
cult,
owing
of delimiting species to intergrading forms.
and even genera

This question
is
often
diffi-
complicated
174
still

further
by
biologic specialization such that forms
which are
in-
distinguishable under the microscope show in inoculation tests different abilities regarding host infection. Thus Neger,^^ Salmon/^* ^^
Reed,^^ and others have shown that spores borne on a particular host are capable of infecting only that host or in other cases only
nearly related species of the same host genus. Forms which can pass from one genus to another are less common. Forms morphologically distinct are regarded as separate species. Differentiations within such species, regarding the species of host plant which they
parasitize, give rise to ''biologic species" or "biologic varieties."
Reed writes of these biologic forms thus "So far as investigated, Erysiphe cichoracearum, is the only one with doubtful exceptions, shown to be capable of infecting plants belonging to more than one genus." "There are other cases where the mildew is limited closely to plants of a single genus," and "Several cases are recorded where the mildew from one species will not infect other species of the same genus. Most of these claims, however, rest on insufficient data." Some morphological species show a very wide range of hosts;
^'^
:
one species, Phyllactinia corylea is known on forty-eight genera in twenty-seven families, others are limited to single genera or to single species of host plant. Two, three, and even five species
are recorded for
some
species of host.
Geographically the Erysiphacese are widely distributed, practically of world distribution, but they are more abundant in the
temperate zones than elsewhere. A pycnidium-bearing parasite, Cicinnobolus, p. 494, is quite frequently found on the mycelium and conidiophores of the Erysiphacese.
Owing to the extreme variability of the perithecial characters and the almost promiscuity of host selection this family presents a most difficult problem to the taxonomist who must either segregate or
"lump"
species.
No middle ground seems open at present.
Key
to Subfamilies and Genera of Erysiphaceae
Mycelium wholly external to the tissues of the

host plant, usually sending haustoria into the epidermal cells only, perithecial
appendages various, more or
less flaccid
I.
Erysipheae.

Perithccial
175
appendages
indeterminate,
ir-
similar to the mycelium, simple or
regularly branched Perithecia containing a single ascus. Perithecia containing several asci
1.
Sphaerotheca,
p. 175.
2.
Erysiphe,
j).
177.
Perithccial appendages determinate Appendages hooked or coiled at the
apex
3.
Uncinula,
p. 180.
Appendages dichotomous at the apex

Perithecia containing a single ascus. Perithecia containing several asci ...
4. 5.
Podosphaera,
Microsphaera,
p. 182. p. 185.
Mycelium with
special intercellular
haus-
toria-bearing branches which enter the host by the stomata; perithccial apII. Phyllactinieae. pendages rigid, with a bulbous base.
.
single genus
6.
Phyllactinia, p. 187.
Sphaerotheca, Leveille
Perithecia subglobose; appendages floccose, brown or hyaline, spreading horizontally and often interwoven with the mycelium,
simple or vaguely branched, frequently obsolete; ascus single, Five species, according to Salmon; 8-spored. Engler and Prantl give fourteen.
S.
humuU
(D. C.) Burr.^s-^^
Amphigenous; mycelium usually evanescent; perithecia usually somewhat gregarious, but varying from scattered to cespitose,
58-120 n in diameter; cells small, averaging 15 n; appendages few or numerous, usually long, often exceeding nine times the diameter of the perithecium, more or less straight, septate, dark
brown throughout:
white or even obsolete.
variations are, short, flexuose, pearly-brown, Ascus broadly-elliptic to subglobose,

ju;
rarely abruptly stalked, 45-90 x 50-72 rarely larger, averaging 22 x 15 n.
spores 20-25 x 12-18
m,
Conidia
= Oidium
fragariae) ovate, white,
membrane smooth.
Salmon
has shown that subjecting the conidia of this variety
two hours, increases their germinating power. from the hop, on hop, Potentilla and Spirea he Sowing ascospores secured infection only on the hop.^^ Conidia from hop infected hop but not Spirea.
to low temperature, 0
176

species
is
The
cosmopolitan and among
its
numerous hosts are
the economic genera Dipsacus, Fragaria, Humulus, Phlox, Pyrus,
Rosa, Ribes, Rubus, Scabiosa, Spirea and Viola. It is a common rose mildew of America and England and
especially destructive on the strawberry. S. humili var. fuliginea. (Schl.) Sal.
is
also
Perithecia usually smaller than in the

in diameter, wall usually harder
last,
sometimes only 50
/j.
and more
brittle, cells larger, irfx;
regularly shaped, averaging 25
appendages usually short, pale brown; spores 20-25 x 12-15 /x. Throughout Europe, Asia and North America. is It recorded on Arnica,
Calendula,
Gaillardia,
Coreopsis,
Fragaria,
Impatiens,
Phlox,
Taraxacum, Verbena, Viola, and several other noneconomic genera.

Scabiosa,
Fig. 126.S. mors-uvse, a perithecium discharging its single ascus which After Longcontains eight spores.
^^^^'
S. pannosa (Wallr.) Lev.^--^^ Mycelium persistent, forming dense satmy patches on the
,
.
stem, calyx, petiole, and rarely

at
first
on
shiny white, then becoming gray, buff or rarely brown; perithecia more or less (usually completely) immersed in the persistent mycelium, globose to pyriform, 85-120 yu
leaves,
about 100 ;u; cells obscure, about 10 wide; appendages few, often obsolete, very short, tortuous, pale brown, septate; ascus broadly-oblong to globose, 88-115 fi, averaging 100 X 60-75 spores 20-27 x 12-15 mConidia ( = Oidium leucoconium) ovoid, 20-30 x 13-16 /x,
in diameter, usually
/jl
fj.;
hyaline; conidiophores short.
The
is
Hosts: peach and rose; cosmopolitan. conidia are very common on the rose, but the perithecia
are rare.
What
often passes for this species on roses in America

^
in reality S. humili.
S.
mors-uve (Schw.) B.
at
first
&
C'-^'
is
The mycelium
white,
exceptional
among the
Erysiphese

in that
it
177
later becomes quite brown. It is found in closely felted on stems and fruit. Pcrithecia begin to form in June. patches Amphigenous; mycelium persistent, at maturity forming dense pannose patches of brownish hyphae; perithecia gregarious, more or less immersed in the persistent mycelium, subglobose, 76-110 cells large, at first well defined, then becoming Id in diameter; 10-25 n wide; appendages usually few or even obsolete, obscure, pale-brown, short, rarely longer, up to five times the diameter of
the perithecium, tortuous; ascus elliptic-oblong to subglobose, 70-92, rarely 100 x 50-62 mI spores 20-25 x 12-15 /x. On wild and cultivated species of Ribcs in America; recently
introduced into Europe where it is very destructive. S. lanestris Hark, occurs on various species of oaks in the United
States.
Erysiphe Hedwig
(p.
175)
pendages
Perithecia globose, or slightly depressed, rarely concave; apfloccose, simple or irregularly branched, sometimes
obsolete, usually
more or
less similar to
the mycelium and inter-
woven with it; " Salmon recognizes

E. polygoni D. C.^^
asci several, 2 to 8-spored.
eight species; Engler
and Prantl/ twenty.
Amphigenous; mycelium very variable, persistent, thin, effused and arachnoid, rarely thick, or more often evanescent; perithecia
gregarious or scattered, usually rather small, averaging 90 m, hut ranging from 65 to 180 m;
wide; appendusually distinct, 10-15 variable in number and length, ages very few or many, distinct or more or less intercells
/j.
woven with the mycelium, brown

''
'
or colorless; ^ ,^Fig. 127. E.

-r.
,
polygoni,
the asci. After Salas 22, variable in shape and size, usually small and ovate, with or without a short stalk, 46-72 x 30-45 fi; spores 3-8 rarely 2,
asci
2-8 or rarely as
many
19-25 x 9-14
M(
Conidiophores
hyaline. One of the
= Oidium balsamii) medium;
conidia ovate,
and turnip.
commonest species, especially destructive to the pea It was studied by Salmon on one hundred ninety host
178
more
species belonging to eighty-nine genera; one hundred forty-six hosts, some doubtful, are reported. Among the economic
host genera are Adorîs, Alyssum, Anemone, Aquilegia, Brassica, Calendula, Catalpa, Clematis, Cucumis (?), Cucurbita (?), Dahlia,
Daucus, Delphinium, Diervilla, Dipsacus, Fagopyrum, Lupinus, Lycopersicum, Medicago, Pseonia, Phaseolus, Pisum, Tragopogon, TrifoUum, Verbena, Vicia, Scabiosa, Symphytum, Valeriana. This is the most variable species of this genus varying widely in its every character. It includes several species which have by some been set aside as distinct, e. g., E. martii, E. umbelliferarum and
E. hriodendri.
of
Salmon ^^ found that the conidia this form grown on Trifolium
pratense were unable to infect other species of Trifolium.

E. cichoracearum D. C.^
Amphigenous; mycehum usually

evanescent, rarely persistent, white or sometimes pink; perithecia gregarious
or
scattered,
jj.;
80-140
or
rarely 180
cells variable,
often
very distinct, 10-20 n; appendages

variable in
Fig.
number and
,
size,
some
shade of brown; asci usually nuand spores. After Salmon. r^ ^ merous, about 10-15, but varymg from 4 to 36, variable in size and shape, narrowly ovate or subcylindric to broadly-ovate, more or less stalked, 58-90 x 30-35 At; spores 2, rarely 3, 20-28 x 12-20 fx.
asci
,
-,
128. e. cichoracearum,
-,
Conidiophores ( = Oidium ambrosise Thum), short; conidia minute, elliptic, white, 4-5 x 7-5.3 fx. The species is quite variable sometimes closely approaching E. polygoni.
The hosts are very numerous, among them Borago, Calendula, Centaurea, Cichorium, Clematis, Cucurbita, Dahha, Helianthus, Humulus, Mentha, Nicotiana,
Cosmopolitan.
being:
Phlox, Tragopogon, Valeriana, Verbena,
Symphytum.
It
is
of
especial import on composites and cucurbits. Reed ^^ has made very extensive culture studies
of this species
179
and concludes that the same form of ''Erysiphe cichoracearum D. C, occurs on at least eleven species of the cucurbits, belonging
to seven genera, infection occurring in these cases in fifty per cent or more of the trials. Six other species were also infected, but in
a smaller percentage of cases.
...
It is also plain that
the biologic
form
of
Erysiphe cichoracearum, occurring on so

is
many
cucurbits
of
this
not entirely confined to the
Out of fifty-four species leaves of Plantago rugelii, a species belonging to

one family.
the Plantaginacese, which were inoculated, ten Furthermore out of ten became infected.
. . .
leaves of squash seedlings, inoculated with conidia and the from plantain, six became infected
.
. .
Helianthus annuus, was infected in thirty-five per cent of the trials in which conidia from the squash were sown on leaves of seedThe cucurbit mildew could not be lings.
sunflower,
. . .
transferred to asters and goldenrods nor was the mildew occurring on these in nature able to infect the squash.
Neither the aster mildew nor

able to
infect
the
cucurbit mildew proved

caesia.
goldenrod, Solidago on this host able to infect asters or squashes." E. taurica Lev. is found in Europe, North
Africa and Asia on
Nor was the mildew
Capparis, Cicer, Clematis

Fig._
1
and various other hosts. E. graminis D. C.

n 1-Usually epiphyllous, rarely amphigenous; mycelium more or less persistent, forming scattered
TT
11
129.-E. graconidial
minis, stage. ^'
After Sal-
first white, then brown or gray; perithecia large, usually about 200 n, scattered or gregarious, cells obscure; appendages rudimentary, few or numerous, very short, pale browTi; asci numerous, 9-30, cylindric to ovate-oblong, more
patches, at
135-280
/x,
or less long-pedicellate, 70-108 x 25-40 m; spores 8, rarely 4, 20-23 X 10-13 /x, seldom produced on the living host plant.
monilioides) with a grayish cast; coniconidia ovoid, white or sordid, 25-30 x 8-10 mdiophores medium It is found on a large number of species of the Gramineae in-
Conidial form
= Oidium
tall ;
180
eluding species of Avena, Festuca, Hordeum, Phleum, Poa, Saccharum, Secale, and Triticum. The asci are peculiar in that they usually contain undifferentiated granular protoplasm, not spores, though in some cases the ^ found that after a few spores, normally 8, are present. Wolff days in water the undifferentiated ascoplasm developed spores which proceeded to normal germination. This species on grasses shows no morphological differences, yet inoculation tests have revealed in it numerous biologic varieties. Reed^^ summarizes the results of his own work together with that of Marchal ^^^ and Salmon ^"^ as follows "So far as tested, all species of A vena are susceptible to the oat mildew. All species of Triticum are likewise susceptible to the wheat mildew. We find, however, that certain varieties of Triticum dicoccum are practically immune to the wheat mildew. Other varieties of this same species are entirely susceptible. Some species of Hordeum are immune to the barley mildew, and the same seems to be true of certain species of Secale with reference to the rye mildew. "To these general statements there are two possible exceptions. Marchal states that the oat mildew will infect Arrhenatherum elatius. Salmon, however, obtained a negative result with the
:
oat mildew on this grass. way. The other exception
The evidence
is
is
not conclusive either
that, according to Salmon, conidia
from wheat can infect Hordeum silvaticum. "It would seem then that under normal conditions there are well-defined forms of Erysiphe graminis occurring respectively on the species of each of the four cereals." It is thought that some hosts may act as bridging species and enable the parasite to pass from one host to another to which it
could not pass directly.
Uncinula Leveille
(p. 175)
Perithecia globose to globose-depressed; appendages simple or rarely once or twice dichotomously forked, uncinate at the apex, usually colorless, rarely dark brown at base or throughout; asci
several, 2 to 8-spored.
There are eighteen or twenty
species.

U. necator (Schw.)
Burr.io^*
i"^'
181
^"^
Amphigenous; mycelilim subpersistent; perithecia usually epiphyllous, occasionally hypophyllous or on the inflorescence, more or less scattered, 70-128 n; cells distinct, rather irregular in shape, 10-20 fi; appendages very variable in number and
length, 7-32, rarely
up to
40,
to 4-times the diameter of the
perithecium, septate, thin walled, light or dark amber-brown basally, rarely branched, asci 4-6 rarely up to 9, broadly-ovate or
ovate-oblong to subglobose, with or without a short stalk, 50-60 x 30-40 /x; spores
4-7, 18-25 X 10-12 /x. Conidial form ( = Oidium tuckeri), conidiophores short; conidia elliptic, oblong,
or obtusely rounded, 2 to 3-catenulate,
hyaline, 25-30 x 15-17
/z.
Ampelopsis and Actinidia. One of the worst pests of the family.
Hosts
Vitis,
ingly septate.
The mycelium is thin walled and sparThe haustoria arise from

lateral
swellings of the hyphae, the epidermis with a filamenpenetrate tous projection and swell within the host
cell
lobed
to a bladder-like body.
cells
The
paraI.
and later the neighboring ones turn brown and die.

sitized
germmate T air or m water, sendmg to several germ tubes.

,
The
conidia
piQ 130. U. necator. II. Perithecium showing/, appendages, and a, asci. IV. in moist Group of asci removed readily from perithecium emitr iL f
. .
forth from one
ting
s,
ascospores.
After
^'^'
The perithecia are found well developed as early as June or July in the United States and are rather evenly scattered over the affected surfaces. Bioletti ^^^ says that a period of warm moist
weather which favors luxuriant mycelial growth, followed by sudden lowering of temperature to about 50 F., favors their most
rapid formation.
their
They
are at
first
hyaline, later brown.
After
during winter, the appendages develop as outgrowths from the outer walls. During ^"^ winter the appendages break off. failed to secure Galloway
definite, usually
form and walls become
germination of ascospores earlier than February or March, but
182
had been exposed to the weather until spring and were then placed in a hanging drop culture afforded spores, some of which grew though many of them burst as they emerged from the perithecium. Ascospores are known to have remained
perithecia which
viable for at least eighteen months. ^^
No
successful infections
were made from ascospores.
Though
rarely.
not found in Europe until 1892

It appears that in their
perithecia are frequently found in America they were ^^ and are now found there but
absence the fungus hibernates in
Fia. 131.
necator. Photomicrographs of perithecia on surface of leaf. A, Magnified 8 times. B, Magnified 35 times. After Bioietti.
U.
.1
specially resistant cells of the mycelium 330 knotty swellings near the haustoria.
which develop within
U. salicis (D. C.) Wint. on willow and poplar in Europe, Asia, and America, U. aceris (D. C.) Sacc. and TJ. circinata C. &. P. on maple are common species. U. flexuosa Pk. occurs on tEscuIus. and elm, U. clandestina (Biv.) Schr. on elm, U. prunastri (D. C.) Sacc. on species of Prunus, especially P. spinosa in Europe. U. mori Miy. is on Morus in Japan.^**^ Several other species of small importance affect numerous hosts.
Podosphaera Kunze
(p. 175)
Perithecia globose or globose-depressed; ascus solitary, subglobose, 8-spored; appendages equatorial or apical, dark-brown
or colorless, dichotomously
branched at the apex, branches simple

and two
1
183
and knob-shaped; appendages rarely of one set apical, brown, rigid, unbranched or rarely to 2-times dichotomous at the apex, the other set basal, short,
straight or swollen
kinds,
flexuous, frequently obsolete.
Salmon
recognizes four species; Engler
and Prantl
seven,
P. oxyacanthae (D. C.)
De Bary
i'
^"^
Amphigenous; mycelium
variable,
persistent in thin patches
or evanescent; perithecia scattered or more or less gregarious, subglobose, 64-90 n; cells 10-18 /x; appendages spreading more
Fig. 132.
tips; h,
a, perithecium showing the appendages with the one large ascus containing eight spores; c, the summer a spore germinating in water. After Longyear. spore-form; d,
P. oxyacanthae.
or
less,
equatorial, variable in
number and
length,
from 4-30
in
number and from 3^-6
or even
10-times the diameter of the
perithecium, usually unequal in length, dark brown for more than half their length from the base, apex 2 to 4-times dichotomously
branched, branches usually short and equal, ultimate branches rounded, swollen, and more or less knob-shaped, Fig. 133; ascus
broadly obovate, or subglobose, 58-90 x 45-75 6, 18-30 X 10-17 M-
fi;
spores
8,
rarely
Conidia
= Oidium
cratsegi).
Salmon
finds the species very variable
separate species P. tridactyla and P. myrtillina as authors. On some hosts perithecia are rare. It
but cannot set aside as is done by some

is
thought that
Pyrus,
the mycelium remains alive over winter. Hosts: Amelanchier, Crataegus, Diospyros,
Prunus,
184
Spirea and Vaccinium. Especially damaging to cherry and apple. Throughout the northern hemisphere.
P. tridactyla (Wal.)
De Bary
is
considered
by Salmon
^^
as a
variety of the last species. other species of Prunus and of Spirea. Similar to the preceding in habit and general character but differing in more critical characFig. 133.
Hosts: Plum and
P. oxycanappendage
After
Sal-
thae,
tips.
Perithecia 70-105 n; cells 10-15 ju; appendages 2-8 usually 4, 1 to 8-times the
ters.
mon.
diameter of the perithecium, apical in origin, more or less erect, apically 3-5 or 6-times
dichotomously branched, primary branches usually more or less elongate, sometimes slightly recurved; asci globose or subglobose, 60-78 X 60-70 /x; spores 8, 20-30 x 13-15 n. Chiefly European but found also in Asia and America.
P. leucotricha (E.
&
E.) Salm.
persistent,
thin,
Mycelium amphigenous,
effused;
perithecia
densely gregarious, rarely more or less scattered, 75-96 n, subglobose, cells 10-16 fj,; appendages of two kinds, one set apical
the other basal; apical appendages 3-11 in number, more or less widely spreading, or erect-fasciculate, 4 to 7-times the diameter
of the perithecium, apex undivided and blunt or rarely once or twice dichotomously branched, brown basally; basal appendages nearly obsolete or well developed, short, tor-
tuous, pale brown, simple or irregularly branched; ascus oblong to subglobose, 55-70 x 44-50 m
spores 22-26 x 12-14
fx,
crowded in the ascus.

ellipsoid, trun-
Conidia
= Oidium farinosum):
28-30 x 12 /i. Primarily American but occurring in Europe and Japan. A most serious pest of the apple. This and P. oxyacanthse, the apple mildews of
cate, hyaline,
America, have been variously treated by writers so that the literature presents an almost inextricable tangle as has been pointed out by Pam-
^^^"^
134. trie ha, aptips.
pendage
After Salmon.
and by Stewart,^^^ Podosphaera oxyacanthse being frequently reported instead of P. leucotricha. Sphaerotheca mali and Podosphsera oxyacanthse have also been much confused, due
mel
^^^
185
to similarity of habit and the frequent abnormal development of the appendages, so that the published references are not always
reliable.
Microsphaera L^vielle
(p.
175)
Perithecia globose to subglobose; asci several, 2 to 8-spored-, appendages not interwoven with the mycelium, branched in a
at the apex, usually dichotomously and often very ornately, rarely undivided or merely once dichotomous. there are thirteen species; Engler and According to Salmon
definite
manner
Prantl
recognize thirty.
grossulariae (Wal.) L6v. Epiphyllous or amphigenous;
M.
mycelium evanescent
or
sub-
persistent; perithecia scattered or densely aggregated, globose-
depressed, 65-130 n; cells 14-20 n; appendages 5-22, colorless, 1-1
times the diameter of the perithecium, 4 to 5-times closely dichoto-
mously branched, branches of first and second order very short, all
fiq.
i35._m.
grossularijE,
append-
age tips. After Salmon, segments deeply divided, tips not recurved; asci 4-10, broadly ovate or oblong, usually with a very short stalk, 46-62 x 28-38 m; spores 4-6, rarely 3, 20-28 x 12-16 li. On five species of Ribes and two of Sambucus. This is the common European gooseberry-mildew, which is not common in America except on the elder.
M.
berberidis (D. C.) L^v. occurs on the barberry in Europe

alni (Wal.) Salm.
and Asia.
M.
persistent; perithecia scattered to gregarious, globose-depressed, very variable in size, usually small, 66-110 n, or even up to 135 n; cells 10-15 n wide; appendages variable in number (4-26) and length, ^/^ to
Amphigenous; mycelium evanescent or
23^ times the diameter of the perithecium, more or less rigid, colorless throughout or amber-brown at base, apex variously
186
(but not always) more or less closely 3 to 6-times dichotomously branched, tips of ultimate branches regularly and distinctly recurved; asci 3-8, ovat3 to ovate-globose, 42-70 x 32-50 m, usually but not always short stalked; 4 to
8-spored; spores 18-23 x 10-12 /x. This species is the most variable of
the Erysiphese showing large latitude in number of spores in the ascus, in length,
color
Fig.
and branching
i j
of appendages, in
136. M.
tips.
appendage size of perithecia. It occurs upon very After Salmon. rr-ii numerous hosts. 1 he economic ones on
alni,
it is most common are: Syringa, Lonicera, Alnus, Betula, Quercus, Carya, Castanea, Juglans, Platanus. It is confined to the northern hemisphere.
which
Salmon recognizes in addition to the typical form six varieties. Those of economic importance are: (a) extensa (C. & P.) Salm., a robust form on various American
species of oaks;
(b)
oaks
calocladophora (Atk.) Salm., also a robust form on American but having pseudo-trichotomously branched appendages
and large spores;

(c) vaccinii (Schw.) Salm., in America on Catalpa and various genera of Ericaceae is a small-spored, long-appendaged form. It
includes
(d)
M elevata on Catalpa;
&
P.
lonicerae (D. C.) Salm.,
on species
of Lonicera in Europe.
M.
diffusa C.
Amphigenous; mycelium persistent, thin and effused, or subpersistent and forming vague patches, or quite evanescent; perithecia scattered or gregarious, globose-depressed, very variable in
size,
55-126
ii
in diameter, averaging
90-100
jx,
cells
10-20 m
wide; appendages very variable in number and length, 4-30, or rarely crowded and as many as 50, 13^ to 7-times the diameter of the perithecium, smooth, aseptate or 1 to 3-septate in the lower half,
colorless or pale
brown towards the
base, flaccid
when
long, thin-
walled above, becoming thick-walled towards the base, apex 3 to 5times dichotomously or subdichotomously divided, branching diffuse
and irregular, branches of the higher orders sub-nodulose, often apparently lateral, tips of ultimate branches not recurved;
187
asci 4-9, 48-60 x 28-30 /x, ovate-oblong with a very short stalk; spores 3-6, usually 4, 18-22 x 9-11. Hosts: Desmodium, Glycyrrhiza, Lespedeza, Phaseolus, Sym-
has recently been described as a species injurious to the beet. It is said to resemble E. polygon! but that cross inoculation between the beet and clover could not be made.
phoricarpos. M. betae Vanha
^^^
M.
ferruginea
Erik,
is
found on cultivated Verbenas
^^^
in
Sweden.
M.
and
euphorbiae (Pk.) B.
&
C. occurs on various hosts in America
Asia, including Astragalus, Colutea, Cuphea and Euphorbia. Its only economic importance is as the cause of a disease of the
AC
and cowpea ^^^ on which it is very common. Amphigenous; mycelium usually subgeniculate; perithecia gregarious in fioccose patches or scattered, 85-145 ju, rarely 180 yi, cells 10-15 m; appendages 7-28, usually narrow, more or less flexuose and nodose, 2.5 to 8 times the diameter of the perithecium,
roselle
colorless above, 3 to 4-times
irregular
and
lax; asci 4-13, rarely

ii;
dichotomously branched, branching up to 26, ovate or ovate-oblong,

spores usually
4,
short-stalked, 48-66 x 26-35
rarely 3, 5 or
6,
16-21 x 10-12
II.
Phyllactinia Leveill^
(p.
175)
Perithecia large, globose-depressed to lenticular; asci
many, 2
or 3-spored; appendages equatorial, rigid, acicular, with a bulbous base; apex of perithecium with a mass of densely crowded
branched outgrowths. Typical epidermal haustoria are not produced but the mycelium sends special branches through the stomata into the intercellular spaces of the leaf.^^^ These branches attain some length and constitute a limited internal mycelium, a character that is considered
by some
as of sufficient importance to set the genus apart in a separate family. The internal mycelium gives off haustoria which The appendages exhibit strikpenetrate cells of the mesophyll.
ing hygroscopic
movements and
is
aid in dissemination.
recognized by Salmon. P. corylea (Pers.) Karst. Hypophyllous or rarely amphigenous; mycelium
Only one species
evanescent
188
or more or less persistent; perithecia usually scattered, rarely gregarious, 140-270 n, rarely up to 350 /i; cells rather obscure,
15-20
fi;
the apical outgrowth becomes mucilaginous attaching

it
the perithecium firmly to places where
may
fall;
appendages
Fig.
1. Natural size, on corylea. Perithecium enlarged. 3. Two asci. 6. Co4. Three spores. 5. Conidia-bearing hyphae. nidium germinating. After Anderson.
137.
Phyllactinia
leaf.
2.
chestnut
to 3-times the diameter of the perithecium; asci 5-45, subcylindric to ovate-oblong, 60-105 x 25-40 fx, more or less stalked, 2, rarely 3-spored; spores 30-42 x 16-25 /x.
5-18, equatorial,
Conidia
clavate.
(=Ovulariopsis)
acrogenous,
solitary,
hyaline,
sub-
Magnolia, Liriodendron, Berberis, Xanthoxylum, Ilex, Celastrus, Acer, Desmodium, Crataegus, Heuchera, Ribes, Hamamelis, Fraxinus, Asclepias, Catalpa, Cornus, Ulmus, Betula, Alnus, Corylus,
On
Ostrya, Carpinus, Quercus, Castanea, Fagus and Typha.
189
Perisporiaceae
Aerial
(p.
170)
mycelium covering the substratum with a dark growth, rarely absent, usually astromate. Perithecia on the mycelial threads or on a stroma, black, more or less globose, without opening or appendages, although in some genera (Meliola, etc.) mycelial outgrowths from the base of the pcrithecium simulate aj)pendages.
Asci elongate, numerous; spores various; paraphyses none. Chiefly parasites, although several genera are saprophytes. About three hundred species.
forms are known.
Aside from ascospores, in some species conidia of one or several These may be borne in pycnidia or uncovered
on hyphae.
Apiosporium
is
especially rich in the
number
of its
conidial forms.
Key to Genera of
Spores 1-celled Spores not curved
Perisporiaceae
Spores hyaline Spores brownish Spores curved, green

Spores 2-celled
Spores,
at
least
1.
Anixia.
2. 3.
Orbicula.
Pseudomeliola.
when immature, ap4. Zopfiella.
pendaged Spores not appcndaged Perithecia borne on the

Spores smooth
Aerial Aerial
aerial
mycelium
Spores not enlarging after maturity
mycelium prominent. ... mycelium none, or poorly

cylindric-clavatc;
sites
5.
Dimerosporiuni,p. 191.
developed
Asci Asci
para6.
Parodiella.
saccate,
large;
sapro7. 8.
phytes
Spores finely cchinulate
Spores enlarging after maturity ... Perithecia borne on a hairy stroma..
.
Zopfia.
Marchaliella.
9.
Richonia.
Lasiobotrys,
p. 191.
10.
Spores 3 or more colled Aerial mycelium none or poorly developed
190

Spores with cross walls only Spores elongate to oylindric
Spores 4-celled, saprophytes Spores 4 to 8-celled; parasites. ... Spores needle-formed

Spores muriform Spores brown
11. 12. 13.
Perisporium.
Schenckiella.
Hyaloderma.
14. Cleistotheca.
Spores hyaline
Aerial
15.
Saccardia.
mycelium prominent
Spores with cross walls only
Spores hyaline
Saprophytic
Parasitic
16.
Scorias.
17.
Zukalia, p. 191.
Spores brown
Perithecia without apparent ap-
pendages
Perithecia
rounded,
elongate,
opening
18.
irregularly
Antennaria,
p. 192.
Perithecia
clavate,
slits.
.
opening by regular
Perithecia
19.
Apiosporium,
p. 191.
appearing
to
have
20. Limacinia, p. 193.
appendages
Stromatic
Not stromatic
(Some species of Meliola have muriform spores) Spores muriform Spores with an appendage at each end
Spores not appendaged
Subicle crustose
21. Meliola, p. 193.
22. Ceratocarpia.
23.
Capnodium,
p. 192.
Subicle radiate
24. Pleomeliola, p. 193.
The genera
coatings than
leaves, fruit
of interest as
covering, shading
pathogens induce disease rather by and smothering leaves with dense sooty-black
by
parasitizing their hosts.
They
upon
are not strictly
speaking parasites but live saprophytically upon the surfaces of
and twigs often subsisting exudations, the so called "honey dew."
insects or insect

Dimerosporium Fuckel
(p.
191
189)
Perithecia depressed-globose, membrano-carbonous; asci clavate

to ovate, 8-spored; spores 2-celled, hyaline or brownish; mycelium abundant, dark, forming a film and often bearing conidia on
conidiophores.
D. mangiferum Sacc. does some harm to the mango. D. pulchrum, Sacc. grows upon the leaves of several woody plants, such as privet, Lonicera, Carpinus and
Cornus.
Conidia=Sarcinella heterospora. D. coUinsii (Schw.) Thiim., forms witches
brooms on the service berry.

Lasiobotrys Kunze (p. 189) \f /
'
^ ^. Pjq 138. Dimerospo"o'nidTa^ G?ascus!

^^^^^ Winter.
Perithecia
superficial,
black, aggregated
in
minute, globose, botryose fashion, stro-
cylindric, 8-spored; spores oblong, 2-celled, hyaline. species L. lonicerae Kze. forms dark coatings on honeysuckle leaves in Europe, North Africa and Siberia but does Uttle
mate;
asci
The one
or no harm.
Zukalia Saccardo
This genus
its
is
(p.
190)
like
Meliola except in
is
its
hyaline spores and in
perithecium.
Z.
stuhlmanniana
on seedling cocoanuts and other palms.

(p.
Apiosporium Kunze
nous
190)
Perithecia superficial, minute, globose to pyriform, membraor carbonous; asci ovate to clavate, 8-spored; spores
globose to oblong, hyaUne; paraphyses none.
Conidia=Torula,
Fumago, Chsetophoma,
Several forms are
of
etc.
known
to constitute sooty coatings
on leaves
plants, subsisting on insect secretions. The specific limitations in the genus have not been satisfactorily worked out
woody
owing to the comparative rarity of the ascigerous stages. A. salicinum. (Pers.) Kze. is common on leaves of many species
of
woody
plants.
Perithecia brownish, gregarious, globoid-oblong,
composed
of
192

;
cells as in the Erysiphaceae spores ovate, guttulate, hyaline, 10 X 8 m; conidia of various kinds, formed from the bases of the perithecia, (a) multicellular macroconidia, (b) unicellular micro-
minute
conidia, (c)
gemmae.
in Brazil. reported on grape Various species also occur on numerous woody and herbaceous plants which are infected with aphids or upon which their "honey
A. brasiliense
Noack
is
^^^
dew"
falls.
differs but little from Apiosporium. A. pithyophila Nees. occurs on leaves of fir; A. elaeophila Mont.
Antennaria Link
Fig. 139.
Apiosporium aalicinum.
After Anderson.
on the Olive; A. setosa Zimm. on coffee; A. monly on green house plants; A. piniphilum
footi B.
Fcl.
&
fir.
D. com-
on
Capnodium Mont.
This
C.
is
(p.
190)
of similar habit
easily distinguished
from genera
by
its
muriform
C.
spores.
quercinum Pers. occurs on oak; C. taxi S. & R. on Taxus; foedum Sacc. on Oleander; C. coffeae Del. on coffee; C. tiliae

Fcl. on Tilia; C. and America.
citri
193
B.
&
P.
on leaves of
citrus fruits in
Europe
on various
trees in
C. stellatum Born, and C. guajavae Born, cause sooty mold trees in the tropics;'"'*"' C. corticolum McAlp. on citrous
South Wales ^^^ and Australia; C. javanicum Zimm., C. meridionale Arnaud is on Oleander, oak, and on coffee. ^"' C. olea Arnaud ^^^ on olive in France. olive, in Europe; Limacinia tangensis P. Henn. is on the mango and cocoanut in
New
^^^
Africa.
Pleomeliola hyphaenes P. Henn.

Africa.
is
on leaves
of
Hyphaene
in
Meliola Fries
Perithecia
(p.
190)
globose, surrounded by dichotomously branched which resemble the appendages of the Erysiphacese; asci hyphae
short,
broad,
spores oblong,
rarely
to 8-spored; 2 to 5-septate,
muriform;
paraphyses
none.
This is a genus of over one hundred thirty species, whose mycelium grows superficially upon leaves and twigs.
M.
camelliae (Catt.) Sacc. oc-
curs on Camellia.
Mycelium,
copious,
black,
bearing various sporing bodies;

perithecia black, spherical, 80150 II., containing several 8-
spored asci; spores 16-18 x 45

olivaceous,
4-celled.
/x,
n,
Stylo-
spores ovoid, 5
hyaline, borne
in flask-shaped pycnidia
which
may
high;
be as
much
as 1 or 2
mm.
M. caniclH.'E. Fiu. 140. and spores. 4, other

nidium.
pycnidia globose resemthe perithecia but smaller, bling

spores.
3, pyonidium form of pyc5, pcrithecium, ascus and asAfter Webber. cospores.
containing spherical spores of about the same size as the styloChlamydospores are also formed by the breaking up of
194

Fumago
camelliae Catt.
is
the mycelium.
species.
a conidial form of this
is found on Citrus forming a sooty black on "honey dew," following principally certain insects as Aleyrodes, Ceraplastes, Dactylopius, and Aphis. The
M.
penzigi Sacc.^^^"^^^
It subsists
mold.
species
is
The hyphae
superficial,
quite similar to the preceding. are from olive-green to dark
are connected into a compact
membrane.
brown and when old The fungus is entirely
possessing, however, small knob-like projections for attachment and large discs (hyphopodia). Reproduction is by conidia, pycnidia, stylospores and perithecia.
Webber
says:
"Several forms of conidia are produced, some being but slight modifications of the common cells of the mycelium, while others
are compound spores. Pycnidia are small, spherical black reproductive bodies, about 40 fi in diameter, and are usually present in considerable numbers in the mycelium. They may be readily
seen with a strong magnifying hand lens, but cannot be definitely distinguished from perithecia or the young stages of the stylospores.
Stylospores are borne in conceptacles, which in their simplest form resemble flasks with long drawn-out necks. Frequently, however,
much branched, and as they project from 1 to 2 mm. bethe mycelium they form quite a conspicuous part of the yond fungus. They are easily recognized with the unaided eye, and can
they are
be seen with considerable distinctness with a hand
lens.
Perithecia
are black, spherical reproductive bodies closely resembling pycnidia,
from which they can not be distinguished with a hand
lens.
How-
ever, they are larger, being eighty micro millimeters in diameter. Each perithecium contains several asci and each of these bears
Some of the investigators who have studied eight ascospores. this disease have failed to find perithecia, and only twice has the writer found them in his examination of material from Florida.
various reproductive bodies other than perithecia, particthe conidia and stylospores, are developed in great abunularly dance."
"The
M.
ing
niessleanea Wint.
insects
is
common on Rhododendron.
Several entomogenous f ungi ^^^"^ '^ have been found which by prey-
upon those
which secrete honey dew, lessen the injury

from all sooty molds. and Sphaerostilbe.
195
''''
Among
these are the genera Aschersonia
Microthyriaceae
(p.
170)
Mycelium superficial, dark; perithecia superficial, separate, shield-shaped, unappendaged, black, membranous to carbonous, formed of radiating chains of cells; asci 4 to 8-spored, short; paraphyses usually present. A family of over twenty genera and more than three hundred species, chiefly poorly understood.
Only two species have been noted as serious economic pathogens; Scolecopeltis aeruginea Zimm. and Microthyrium coffae both on coffee in Africa. The genera of the Ascomycetes which remain to be treated, and which are separated from those preceding by the possession of an ostiole, are by some known under the name Pjrrenomycetes. Cf. p. 170. There are three orders, the Hypocreales, Dothidiales and Sphseriales.
Hypocreales
(p.
124)
chief character separating this order from other Pyrenomyand cetes is the brighter color yellow, purple, scarlet, red, etc.
The
the more tender texture of
its perithecia,
soft, fleshy,
cottony,
from that of the preceding orders in the possession of a distinct opening, ostiole, for the exit of spores. Perithecia globose to cylindric or flask-shaped, free on the substratum (rarely subepidermal) or united by a common matrix,
wall
patellate or effused. The perithecium also differs
which varies from a cottony subiculum to a distinct fleshy stroma, membranous or at least not truly carbonous; asci cylindric, clavate or subovoid, mostly 4 to 8-spored but often becoming 16-spored by the separation of each original spore into two globose
or subglobose cells; spores simple or globose to filiform.
compound, hyaline or
colored,
Conidia are usually produced freely, each genus usually possessing at least one form of free-borne conidia, while in some genera several different kinds of conidia are found. Pycnidia are rare.
Often the ascigerous stage is nearly suppressed and rare while one or more of the conidial forms predominates.
196
Such form genera as Verticillium, Tubercularia, Sphacelia, Sphserostilbe and Isaria are connected with the Hypocreales. The order includes some sixty genera, and over eight hundred species. Of these only a half dozen genera contain important plant parasites, another half dozen genera, parasites of less importance. The rest are saprophytes, insect parasites, etc., of no economic
significance.
Opinion
differs as to
the characters which should be
made the
stress
basis for subdivision of this family,
whether to throw main
upon the structure

the spores.
of the perithecium or
7
upon the character

of
the order contains a smgle family, Hypowhich may be divided into six subfamilies. Accordcreaceae,^-^'^-^ ing to a more recent treatment of the American members of the
Following Lindau
group by Seaver
'
'
two famihes and four
tribes are recognized.
Lindau's tribes Hyponectrieae, Hypomyceteae, and Melanosporeae are united with a part of Nectriese under the last name while the
remaining genera, referred by Lindau to this

tribe Creonectrese.
tribe, constitute
the
These
tribes constitute the family Nectriaceae.
The remaining
same
tribes,
Hypocreese and Clavicipiteae with about the
limits constitute the family Hypocreacese.
Key to Tribes
Perithecia at
later
first
of Hypocreaceae
sunken
in the substratum,
1.
erumpent Perithecia not sunken in the substratum; stroma present or absent Stroma cottony, never fleshy; perithecia immersed in the stroma, or borne on
its
Hyponectrieae.
surface
fleshy or
2.
Hypomyceteae.
Stroma
wanting
Spores dark colored; perithecia free on the substratum (in some species of Melanospora with a cottony
stroma) scattered Spores hyaline, yellow or red Perithecia without a stroma, or on a

fleshy
3.
Melanosporeae.
stroma
4.
Nectrieae, p. 197.

Perithecia sunken in a fleshy stroma
197
Spores not filiform; j)erithecia iialf or entirely sunken in the

stroma, and distinct from
it.
.
5.
Hypocreeae,
p. 198.
Spores
filiform;
perithecia
in
comthe
dis6.
pletely
embedded
stroma and not clearly tinct from it
Clavicipiteae, p. 199.
The
first tribe
third contain but one each.
contains no parasitic genera while the second and Of the Hypomyceteae, the genus
(p. 200) is set off from the others by its 2-celled hyaline Of the Melanosporeae fusiform spores, and its cottony stroma. the genus Melanospora (p. 200) is distinguished by the long beaks
Hypomyces
of its flask-shaped perithecia,
which are brown rather than black,
and
its
brown
2-celled spores.
Keys to the Genera of
Nectrieae, Hypocreeae
(p. 196)
and
Clavicipiteae
Tribe IV. Nectrieae

Conidiophores not of the Stilbum type
Spores elongate, 1-celled; perithecia free on the substratum stroma none
;
Spores not appendaged Perithecia yellow or red

Asci cylindric; ostiole concolorous with the perithccium Asci
clavate-cylindric
;
1.
Nectriella.
ostiole
2. 3. 4.
darker than the perithecium.. Perithecia violet or blue Spores appendiculate Spores elongate, 2 to many-celled Spores with cross walls only Spores 2-cellcd Asci 8-spored; often with 1-celled, conidia formed in the ascus
Thelocarpon.
Lisiella.
Eleutheromyces.
Perithecium yellow or red

Spores hj'aline
5. 6.
7.
Nectria, p. 201.
Spores brown Perithecium blue or violet
Neocosmospora,
Lisea.
p. 205.
198

Asci many-spored
Perithecium
vated
fleshy,
ostiole ele8.
Metanectria.
Perithecium hard, ostiole sunken

Spores 2 to many-celled Spores not appendiculate
9.
Cyanocephalium.
Perithecium bright colored, not

blue
10. Calonectria, p. 205. 11.
Perithecium blue or violet

Spores appendiculate, 4-celled Perithecia clavate, ostiole wartlike
Gibberella, p. 206.
12.
Paranectria.
Perithecia flask-shaped,
ostiole
13.
elongate
Lecythium.
Spores muriform Perithecium

blue
bright
colored,
not
14.
. .
Pleonectria,
p. 207.
Perithecia dark colored or blue.
15. Pleogibberella.
Spores filiform
Perithecia fleshy, bright colored Perithecia horny, brown
16.
17.
Ophionectria,
p. 207.
Barya.
Conidiophores of the Stilbum type, stroma wanting

Spores 2-celled Spores 4-celled
18. 19.
Sphaerostilbe, p. 207. Stilbonectria.
Spores muriform
20.
Megalonectria.
197)
Tribe V. Hypocreeae
Stroma sunken
to
it,
(p.
in the
substratum or grown
21. Polystigma, p. 207. 22. Valsonectria, p. 208.
23. Cesatiella.
usually free later
Spores 1-celled
Spores 2-celled Spores several-celled by cross walls Spores muriform Spores hyaline Spores olive-brown Stroma from the first separable from the
24. Thyronectria. 25. Mattirolia.
substratum
Spores 1-celled Spores 2-celled Cells of the spores separating in the ascus
26. Selinia.

Stroma Stroma
ascus
patellato or effuse erect, simple or branched.
27.
. .
199
28.
Hypocrea, Podocrea.
p. 209.
Cells of the spores not separating in the
Stroma Stroma
patellate or effuse
erect,
29.
Hypocreopsis.
branched
not
30. Corallomyces.
Spores 3 to many-celled
Stroma bright or dark

conidia-bearing
colored,
31.
Broomella.
Stroma dark, green

conidia
or
black,
with
32. 33. 34.
Conidia of two kinds
Loculistroma,
p. 215.
p. 211.
Secondary conidia absent Spores muriform

Tribe VI. Clavicipiteae
Aciculosporium,
Uleomyces.
(p.
197)
Stroma effused Stroma forming a sheath about the host Stroma flat, tuberculate, or disk-shaped Stroma not conidia-bearing Stroma thick, usually light colored. Stroma thin, black Stroma with the inner portion conidia. .
35. Epichloe, p. 210.
36. Hypocrella. 37. Dothiochloe, p. 210.
bearing
38. Echinodothis, p. 211.
Stroma erect Stroma small, saccate, membranous Stroma large, erect, with distinct sterile and fertile portions, the latter often
knob-like
39.
Oomyces.
Stroma formed in the bodies of insects and spiders, or in subterranean

fungi
40.
in the inflorescence of
etc.,
Cordyceps.
Stroma formed
Glumaccae,
spores continuous
41. Balansia, p. 209.
Stroma not growing from a sclerotium Stroma growing from a sclerotium

after a period of rest Asci preceded by conidia Asci preceded by smut-like chla-
42. Claviceps, p. 211.

f
43. Ustilaginoidea, p. 213.
mydospores
44. Ustilagmoidella,p. 114.
200
Hypomyces
Fries (p. 197)

of considerable
Stroma an effused cottony subiculum, often
extent; perithecia numerous, usually thickly scattered and immersed in the subiculum, rarely superficial; asci cylindric, 8-spored;
spores fusoid or fusiform,

usually
apiculate,
rarely
blunt, 2-celled, hyaline; conidial phase variable.
species
This genus of some forty contains but few
saprophytes, the majority
being parasitic, chiefly on the larger fungi. The genus
economic interest only affecting mushrooms, though one species, H. hyacinth! has been found
is
of
as
causing
secondary
19
infec-
Fig.
141. Hypomyces ochraceus. thecia; C, asci and spores; D, spores; E, conidia; F, chlamydospores. After Tulasne.
tion in onions,*' following a bacterial trouble, and

follows
another; H. solani Reinke a similar disease
on potatoes.
longing to various
Chlamydospores and conidiospores develop, beform genera as Verticillium, Mycogone, Fuligo,
Diplocladium, Dictylium, Sepedonium, Blastotrichum. Allied to this genus are probably Mycogone rosea and
niciosa,
M.
per-
which are destructive enemies of mushroom
culture.
Melanospora Corda
(p.
197)
Perithecia superficial, without a stroma, globose-pyriform or flask-shaped, with a long neck which is usually clothed at the
tip
with a fringe of hairs, perithecia often hairy; asci broadly clavate, 4 to 8-spored; spores 1-celled, brown to brownish-black.
The genus
saprophjrtes.
contains
some
is
forty
species,
mostly
common
^"^
M. damnosa
(Sacc.) Lin.
serious
on wheat and
rye.^-^'

M. stysanophora Mat.
Dematophora glomerata,
is
201
said to be
cf. p.
an ascigerous stage of 230, so injurious to the grape.

(p.
Nectria Fries
197)
Stroma absent or tubercular, fleshy, bright colored; perithecia on or in the stroma or among cottony hyphse, globose or ovate, walls fleshy, yellow, red or brown, smooth or
single, or gregarious,
hairy; ostiole papillate or not; asci cylindric or clavate, 8-spored; spores elongate blunt or pointed,
hyaline, rarely red, 2-celled, forming conidia in the ascus; paraphyses
usually none.
form Tubercugenera Cephalosporium, laria, Fusarium, Spicaria, Fusidium and Chsetostroma. Much doubt
conidial stages occur the
exists as to specific limitations,
As
and
as to the
life
histories of the species.
Some two hundred fifty species have p^^, i42.-Melanospora. K, perithecium; L, asci; M, spores. After been described. Several are credLindau. ited with causing serious diseases, most of them occurring as wourid parasites and unable to effect entrance into sound tissue. Other species are pure saprophytes and harmless. The genus Nectria is divided into seven sub-genera, which are
frequently given generic rank, as follows
:
Key
Spores smooth
Perithecia
to Subgenera of Nectria
smooth Stroma fleshy Stroma a cottony subiculum Stroma usually absent; perithecia
scattered
1.
Eunectria,
p. 202.
2.
Hyphonectria.
Dialonectria, Lasionectria.
p.
3.
205.
Perithecia hairy Perithecia scaly
4. 5.
6.
. . .
Lepidonectria.
Spores tuberculatc
Cosmonectria.
Phaeonectria.
Spores appearing striated, golden brown
7.
202

of
The majority
economic species belong to the

Eunectria
(p.
first
subgenus.
201)
N. cinnabarina (Tode) Fr. Stroma erumpent, tubercular, at first pinkish or yellowish-red, darker with age, 1-2 mm. high and broad; perithecia almost globose, the ostiole rather prominent,
first
becoming
slightly collapsed, at
bright cinnabar-red, darker with age, granular, 375-400 ju in diameter; asci clavate, 50-90 x 7-12 n; spores mostly 2-seriate, ends elongate, elliptic obtuse,
slightly
curved,
12-20
4-6
n;
paraphyses delicate. Tubercularia vulgaris borne on the stroma is the conidial stage.
Conidiophores aggregated into tubercular masses each 50-100
ti long; conidia on short lateral branches, elliptic, hyaline, 4-6 x 2 p..
The wood
,
closely
septate
delicate
hyphse grow rapidly through the

Fig. perithecia in stroma, ascospores issuing in After cirri; germinating spores.
143. N. cinnabarina,
or bark, penetrating nearly every Cell and turning the wood , , ^^ ^ r black and COilectmg to form Stro,
^^""^^s-
mata on
fall
or in
the bark.
These
or spring break through the epidermis and produce warty, gray to pink, excrescences, which at first bear profuse conidia both terminally and laterally on short stalks and later
stromata in
dark-red ascigerous structures; though the latter are much less common and are often absent. The fungus is said to be unable to
affect living
cambium and
cortex.
found saprophytically on many decayed woody plants that have been frost killed, and parasitically on pear, Tilia, iEsculus, China berry, Betula, Ribes, Acer, Carya, Morus, Prunus, Quercus,
It is
germinated spores on a cut branch; the mycelium spread to and killed the main stem; tubercles appeared and during the following year perithecia developed on these tubercles. In America the species has attracted attention on the currant ^.'^' ^^ in which host the mycelium invades chiefly the
Ulmus,
etc.
Mayer
^^^

cambium.
parasitic.^-^
203
On
this
host,
however,
it
is
now
said to be non-
Durand,^-^ culturing the conidial form on sterile currant stems, observed the formation of tubercles with abundant conidia after about fourteen days. On agar conidia were produced directly from single hyphae without any stroma. Perithecia were found in the
field
on the tubercles with the conidia in February. N. ditissima Tul. Stroma light colored; perithecia cespitose, densely and irregu-
larly clustered, or rarely scattered, ovate, ostiole prominent, bright
red,
smooth or roughened;
asci cylindric to clavate,
80-90 x 8-10
/x;
spores fusoid, 12-16 x 4-5 ix. The unicellular microconidia are followed by falcate, multicellular, macroconidia (Fusidium candidum), which are borne on
pale stromatic cushions.
Common
on dicotyledonous
trees, especially beech, oak, hazel,
ash, alder, maple, lime, apple
and dogwood, where
it is
usually a
especially
wound parasite, particularly common after hail. well known from Europe and has more recently
tion in America.
It
is
attracted atten-
The mycelium does not usually advance more then one centimeter in each year. It is believed that it can travel within the wood and break through the cambium and cortex at points some distance from the place of original infection, thus producing new spots. Very minute conidia produced in the bark aid in tissue deWhite conidial (Fusidium) stromata appear near composition.
the periphery of affected spots and here, too, in groups or scattered, appear the deep red perithecia.
N. cucurbitula Sacc.
Perithecial clusters erumpent, often irregular in form, 1-2
mm.
in diameter; perithecia densely clustered, bright red, ovate, with
a prominent ostiole, rarely collapsing; asci cylindric to clavate 75-100 X 6-8 n; spores at first crowded and partially 2-seriate,
finally
becoming
1 -seriate,
lying obliquely in the ascus, broad,
fusoid, rarely subelliptic, 14-16

Its hosts are spruce,
fir,
x 5-7
p..
pine and other conifers in Europe and

parasite, often follo\ving hail.
North America.
The fungus
is
usually a
wound
204
Germ tubes from ascospores or conidia enter the cortex and develop a rich myceHum in the sieve tubes and soft host. This advances most rapidly during the dormant period of the bast. White or yellow stromata the size of a pinhead appear and bear numerous conidia. Later come the red perithecia whose ascospores ripen
in winter or spring.
Fig.
Rab. stroma compact; perithecia subCespitose, of perithecia. globose, smooth; ostiole papillate; asci subclavate, After Halsted. gQ-lOO X 15; spores elongate or fusoid, hyaline, On currant. 1-septate, 18-20 x 5-6 mm. N. ipomoeae Hals.
ribis (Tode)
N.
144. N.
ipo-
Perithecia clustered, ovate, roughened, red; asci cylindricclavate; spores elliptic; conidial phase (Fusarium) appearing as a white mold-like covering of the host; conidia several-celled, falcate.
Halsted
^-^
inoculated sterilized egg-plant stems with the Nectria

asci-
spores and the Fusarium form developed, followed by the
gerous stage. The Nectrias found upon egg-plant rise to the Fusarial stage. and sweet potato, morphologically alike, were proved by cross
inoculations to be identical.
Ascospores in hanging drop were also seen to give
N. rousselliana Tul. and N. pandani Tul. are parasitic on Buxus AIT and Pandanus respectively, the former with the conidial stage.
Volutella buxi.
N. solani Ren. & Bert, is said by Massee to be the ascigerous 22 form of Fusarium solani. Perithecia crowded on a stroma, minute, conic-globose, smooth, blood-red; asci clavate; spores hyaline, 8-9 x
paraph yses slender, tips strongly clavate. Conidia ( = Fusarium solani) hyalirie, 3 to 5-septate, fusiform, 15^0 x 5-8 m, but very
5
ju;
variable,
borne on erect, simple or branched

1
Fig.
J.
COnidlophoreS.
145. n. ipomcese, the Fusarium stage. After
Halsted. Zimm. is on cacao and vanilla; N. bainii Mas. N. amerunensis A. & Str. and N. diversispora ^^ The three latter Petch. are reported parasitic on cacao pods. names are probably synonyms of the first.
N.
cofiffeicola
205
tivated \'anda:
N. vandae Wah. and N. goroshankiniana (Wah.) grow on culN. theobromae Alu.ss., })r()l)al)ly identical with
is
N. striatospora Zimm.,
geri
found on cacao trunks as
is
also
N. jun-
Henn.
is
N. bulbicola. Henn.
Ficus.
on orchids and N. gigantispora Zimm. on
Dialonectria
(p.
201)
is
N. graminicola B.
nivale
in
is
^^^
& B., the conidial stage of which destructive to winter wheat and rye
bogoriensis
Fusarium
Bern and N. luteopilosa Zimm. and N. fruticola Zimm. on coffee; ^^^ N. theobromicola Mass. on Theobroma. Neocosmospora E. F. Smith was reported by Smith ^^^ as the ascigerous form of Fusarium pjQ i46. n. ipoascus. moese, an vasinfectum and consequently as the cause of o, c, a, germination of ascospores. Afwilt diseases. Recent work by many serious ^^^' ^^* and by Butler "^ has shown Higgins that in all probabilitj^ there is no genetic connection between these forms and that the fungus unHer discussion is merely a
N.
vanillae
Europe. Less known are N.
Zimm. on
vanilla;
harmless saprophyte.
Calonectria
(p.
198)
Perithecia free, often closely gregarious, true stroma wanting but perithecia often surrounded by a radiate, white mycelium which may simulate a stroma; perithecia globose to ovate, red or yellow; asci elongate, 8-spored; spores elongate, more than
2-celIed.
About
sixty species.
C. pyrochroa (Desm.) Sacc, has been reported parasitic on Platanus. Its conidial stage is Fusarium .platani. C. flavida Mass. is in the West Indies on cacao causing
canker.
C.
cremea Zimm. with Spicaria
colorans,
Corymbomyces
albus,
206

^"
Clanostachys theobromse fruits and stems of cacao.

C. bahiensis
is
probably as
its
conidial stages,
is
on
really
Hem. reported in South America on cacao stems an Anthostomella; C. gigaspora Mass. is found on
sugar-cane.
Gibberella Saccardo
(p.
198)
or occasionally scattered
Stromata tuberculate, more on or surrounding the stroma;
or less effused; perithecia cespitose

asci clavate,
8-spored; spores fusoid, 4 to many-celled, hyaline; conidial phase a Fusarium,
Of the thirteen species but few are parasitic. G. saubinetii (Durieu & Mont.) Sacc. '^^' '''
Perithecia gregarious, leathery membranous, verrucose, ovate,
subpedicellate, bluish, papillate,
200-300 X 170-220
12
spores
m; asci
oblong
clavate, acuminate, 60-76 x 10ju;
one or obliquely
two-ranked, fusiform, curved or straight, acute, 4-celled, 18-24 x
4-5
FiG. 147. G. saubinetii; 2, Fusarium spores, 5, the asci. After Selby.
fji;
mycelium
/ t-i
effused, crus-
tose, white to rose Colored.
Ooor
nidia
= lûsanum)
solitary,
clustered, fusiform, curved, acute or apiculate, 5-septate, hyaline, 24-40 x 5 M-
Many species of Fusarium, e. g., F. culmorum, F. avenaceum, F. hordei, F. heterosporum, have been referred to this ascigerous ^^ stage. Spherical stylospores are also reported.
The mycelium and the
conidial stages often coat the grains
and
heads of cereals with red or pink. Perithecia are rare as shining dark dots on the grains in the late season. The Fusarium stage also is said to cause a clover and alfalfa disease and the fungus by inoculation and culture is shown to be identical on wheat, clover,
barley, rye, spelt,
emmer, and
oat.
It
is
carried
from season to
season on infected seed and causes large loss of young plants. Doubt as to the relationship of the Fusarial forms mentioned with
the ascigerous stages has been raised by the work of Appel and WoUenweber. See also Fusarium (p. 646).
207
G. cerealis Pass., the cause of a serious wheat disease in Italy '" may be identical with the last species. G. moricola Ces. & d. Not. grows on Morus.
Pleonectria Saccardo
(p.
198)
Perithecia cespitose or separate, globose, pale, papillate; asci
8-spored; spores many-septate, muriform, hyaline. P. berolinensis Sacc, which occurs on various
species
of
wild and cultivated currants both in Europe and America has been reported by Durand ^" as associated with a currant trouble
in
New
York.
P. cofifeicola
Zimm. attacks
coffee.
Ophionectria Saccardo
(p.
198)
su-
Stroma globose, tubercular, depressed or none; perithecia

perficial,
clustered or scattered; asci cylindric to clavate, 2 to 8-spored; spores 4 to many-celled, fusoid to subfiliform, hyaline or subhyaline.
sects
About fourteen species. O. coccicola E. & V. attacks scale inand is said also to cause gummosis of oranges. ^^^ O. foliicola Zimm. is found on cofifee.
Sphaerostilbe Tulasne
(p.
198)
stalk with a globose or conical head; perithecia bright colored, membranous, globose, subglobose or ovate; asci cylindric or subcylindric, 8-spored; spores 2-celled, eUiptic or
subelliptic, hyaline.
Stroma a slender
Conidial phase Stilbum, Atractium or MicroS.
cera.
Some twenty
species.
repens B.
&
Br. in India causes a
root disease of Hevea^^^ and arrowroot.

S. flavida Mass.
causes disease of coffee in tropical America.
Polystigma
De CandoUe
(p.
198)
fleshy, effused, red or reddish-brown, growing on leaves; perithecia sunken, only the ostiole being above the surface, thin,
Stroma
leathery, hyaline; asci elongate, clavate, 8-spored; spores ellipsoid,

1-celled, hyaline.
Three
species.
208
P. ruba (Pers.) D. C. causes reddish spots on the leaves of

first bearing pycnidia with filiform hooked, con(Libertella rubra) tinuous conidia. Perithecia produced on old
Prunus.
Stroma
at
bearing ellipsoid to elongate asci; 10-13 x 6 /x> smooth. spores The invaded leaf tissue is colored by the mycelium which bears a reddish oil. Nuleaves,
merous
diseased
perithecia
are
immersed
in
the
area and, opening to the surface,
Fig.
148. p. rubrum.
extrude spores which seem incapable of inDuring winter the stroma darkens, turns hard and produces the perithecia and
fecting.
D, asci; E, conidia. After Fisch.
Ascogonium and trichogyne-like have been described.^'*'* organs P. ochraceum (Wahl.) Sacc. occurs on Prunus padus.
ascospores.
Valsonectria Spegazzini
(p.
198)
Stroma thin, cushion-shaped, under the bark of the host; perithecia similar to those of Valsa, sunken in the stroma, the beak
erumpent,
light
red;
asci
cylindric,
8-
spored; spores 2-celled, hyaline or
brown. genus of but three species which differ from Valsa chiefly in their
red color.
V. parasitica (Murr.) Rehm.^^^' ^^^' Pustules numerous, erumpent, at

first
yellow, changing to
brown
at Fig.
maturity; perithecia usually ten to
twenty in number, closely clustered,
149. Showing a pycnidium of Valsonectria and the manner iu which the spores issue from it. After Murrill.
flask-shaped, deeply embedded in the stroma in the inner bark, scarcely visible to the unaided eye; necks long, slender, curved, with thick black walls and rather prominent ostiola; asci oblongclavate, 45-50 x 9 m, 8-spored; spores usually biseriate, hyaline, oblong, rounded at the ends, often slightly constricted, uniseptate,
9-10 X 4-5
/x.
Summer
spores very minute,
x 2-3
ix,
pale-
209
yellowish, cylindrical, slightly curved, discharged in twisted threads as in Cytospora.
This fungus, originally described as Diaporthe parasitica, is a The mycelium grows through serious parasite on the chestnut. the inner bark in all directions from the initial wound at which infection occurred, eventually girdling the part. affected. The perithecia appear in abundance
The wood
upon
is
also
or in cracks
of the bark, extruding their spores in greenish to yellow threads.
Hypocrea
Fries (p. 199)
Stroma subglobose to patellate, fleshy or subfleshy; perithecia entirely immersed, subglobose to ovate, the necks slightly protruding; asci cylindric, originally 8-spored, spores breaking each into two so that the asci at maturity contain sixteen hyaline
spores.
About one hundred ten species. H. ceretriformis Berk, occurs on the bamboo H. sacchari on sugar cane.
Balansia Spegazzini
in
Tonkin;
(p. 199)^^^
Sclerotium composite, formed of the affected parts of the host embedded in a well developed mass of fungous tissue; stroma
arising from the sclerotium, stipitate and capitate or sessile, pul-
vinate, obovate, discoid, or separated from the sclerotium as
soon as the latter
is
mature, sur-
face slightly papillate from the projecting ostiola of the im-
mersed scattered perithecia; asci 8-spored; paraphyses none. Conidia, when known, an Ephelis and preceding the stroma.
curs
B. hypoxylon (Pk.) Atk. ocon various grasses, chiefly
in the southern
United States.
Fig.
B. claviceps Speg. infests Setaria and Pennisetum in tropical lands.
B. hypoxylon, section of 150. pscudosclcrotium and one stroma showing pi'rithecia, stom, leaf elements and an ascus. After Atkinson.
The^ remaining species, chiefly of

inhabiting.
warm
regions, are mostly grass
210

Dothichloe Atkinson
^^^
(p.
199)
Stroma
thin,
hard when dry, black, especially the outer portion,
lighter within, effuse, pulvinate, disciform or armilla-form, partly
or entirely surrounding the host; perithecia crowded, confluent with the stroma, but the thin walls of distinctive structure, im-
mersed, the apex projecting; asci cylindric, 8-spored; spores filiform, septate at maturity, and eventually separating at the septa into short seg-
ments.
Like the preceding genus, both species D. atramentosa (B. & C.) Atk. and D.
Atk. are grass inhabitors of regions of the United States. The former is the commoner species with a
aristidae
warm
wider range of hosts.

Epichloe (Fries) Tul.
(p.
199)
Stroma
effused, subfleshy, at first pale,
becoming bright orange, sheathing the host; perithecia immersed or with the
ostiola
protruding;
asci
cylindric,
8-
spored; spores filiform, many-celled. Of some nine species only one is important. E. tjrphina (Pers.) Tul. Stroma efat first pale, becoming bright orange, forming sheaths 2-5 cm. long A, habit around stems of various grasses, often the inflorescence; perithecia asco" destroying
fused,
Bre- thickh'^ Scattered, partially or entirely
Fig.
151. Epichloe.
L^cS^bâfcX'A
After Winter, spore. feld and Lindau.
im-
the stroma, soit, membranous, concolorous with the stroma, the ostiole rather prominent; asci very long; spores almost as long as the ascus, closely fasciculate, multiseptate, about 2 fx in diameter; conidia elliptic, hyaline,
mersed
r,
4-5 X 3
ju,
preceding the perithecia on the stroma.
Many grasses are affected, often to serious extent. The mycelium

shows first as a yellowish cobwebby growth surrounding the leaf sheath and soon develops a conidial stroma. Later the stroma

turns
layer.
211
to
orange-color
and the
perithecia
appear,
"^
forming a
Echinodothis Atkinson
(p.
199)
Stromata subfleshy or corky, light-colored, pulvinate to subglobose or irregular in form, often constricted at the base, sometimes entirely surrounding the host, consisting of several layers of
different consistency; perithecia superficial, scattered, subcylindric, sessile, giving an echinulate appearance to the stroma; asci cylindric, 8-spored; spores linear, septate, at
length separating at the
septa into short segments.
Two
species, parasitic
on grasses
in the
warmer parts
of the
western hemisphere. E. tuberiformis (Berk.
&
Stromata subglobose,
or divided, seated
Rav.) Atk.^ê cm. or more in diameter, entire, lobed,

or
upon the reed
upon the leaf-sheath and
fas-
tened by a whitish mycelium consisting of radiating threads which are sometimes tinged yellowish-brown; substance leathery or corky,
consisting of three layers, an inner layer white to pinkish, an intermediate layer light ochraceous and an outer layer cinnamon; stroma externally dark brownish becoming black; conidiophores needle-shaped; conidia ovoid to fusoid, 3^ x 7-10 ix; perithecia entirely superficial in small clusters or evenly distributed over
the exposed surface of the stroma, subconic in form, giving the whole stroma a spiny appearance, clothed except the apex with a
dense covering of minute ^threads which are at first whitish, becoming cinnamon colored, the naked apex becoming black, about 0.3 X 1 mm. asci cylindric, with a swelling at the apex, very large,
;
spores nearly as long as the ascus, hyaline or slightly yellowish, many-septate, the joints 15 x 4-5 /x. On Arundinaria in the Southern States.
(j.;
475-750 x 14-20
Asciculosporium take Miy.'^^ forms witches' brooms on bamboo

in Japan.
It is closely related to Dusiella
and Epichloe.
199)
Claviceps Tulasne
(p.
Sclerotium formed within the hypertrophied tissues of the ovary of the host, succeeding the conidial stage which is a Sphacelia; stroma erect, with a long sterile base and a fertile,
212
usually knot-like head; perithecia closely scattered, sunken in the stroma with only the ostiole protruding, flask-shaped, the walls scarcely distinguishable from the stroma; asci cylindric,
Fig. 152.
C. purpurea. D, Sphacclia stage; E, germinated sclerotia; G, section of stroma; H, section of a perithecium; J, ascus with spores. After
Tulasne.
Some twelve or fifteen 8-spored; spores hyaline, continuous. species are recorded all affecting the ovaries of the Graminese.
C. purpurea (Fr.) Tul.^^^
Sclerotium elongate, more or less curved, and resembling a much enlarged grain, after a period of rest producing few or many, clustered or scattered stromata which are 0.5-1.5 cm, high; spore

60-70
dric,
fi.
213
the
long.
Conidia
= Sphacelia segetum) produced on

is
grain before the sclerotium

1-celled
formed, conidiophores short, cylin-
arranged in a compact palisade, bearing small, oval, hyaline, conidia. Hosts, rye, wheat, oats and numerous other
is
grasses.
Infection of the ovary at blooming time
followed by complete
possession and consumption of the ovarial tissue by the mycelium, and by considerable development of stroma beyond the ovary.
On
its distal
the external much-folded part of this stroma, particularly at end, are borne layers of conidiophores and numerous
is
conidia and a sweet fluid
exuded.
The
conidia, carried
by
in-
Lator the stroma, losing a large sects, of the distal region, rounds off to a definite sclerotium, smooth, part firm, blue to black in color, and several times larger than the
infection.
till the following season, the sclerotium gives rise to several stalked, capitate, perithecial stromata. The perithecia are arranged around peripherally, the
spread summer
normal grain of the host plant. After a period of rest, usually lasting
ostioles protruding and giving the head a rough appearance. The sclerotium constitutes the ergot of pharmacy and contains a
powerful alkaloid capable of causing animal disease if eaten. This species appears to be differentiated into a number of
^'*^ biologic races.
Europe and America, being
C. microcephala (Wal.) Tul. infects numerous grasses both in especially destructive to blue grass.
Two species C. paspali S. reported on Paspalum.^^^
&
H. and C.
rolfsii S.
&
H. have been
Ustilaginoidea Brefeld
(p.
199)
150
Sclerotium formed in the grain of the host, resembling supera smut sorus, in the center composed of closely interwoven hyphae, externally the hyphse are parallel, radiating towards the
ficially
periphery and bearing echinulate, globose, greenish conidia; stroma with a long sterile stem and a fertile head; perithecia immersed
in the
stroma as
Two
in Claviceps; asci are known, one species
ascigerous stage,
and spores also as in Claviceps. on Setaria which produces an the other on rice, the ascigerous stage of which
214
is

is
not known but which
placed in this genus on account of the
similarity of its conidial stage with that of the other species.
ical,
U. virens (Cke.) Tak. Ascigerous stage unknown, sclerotia spherabout 5 mm. in diameter; conidia spherical, at first smoothwalled, hyaline, at maturity echinulate and olive green,
4-6
fx.
The
short
thick
walled
hyphse of the interior of the

sclerotium
are
.
closely
in'
terwoven to a false tissue. T? ..IFig. i-o"^!--^ 1o3. U. virens; a, spores germinated m water; b, germinated in bouillon. After toWard the periphery they become parallel and are directed radially. Here a yellow layer is produced and spores are formed laterally on the hyphae. When mature the spores are in mass dark olive-green and form an outer green layer on the
sclerotium.
tive
The
spores germinate in water, producing a vegeta-
mycelium which bears secondary spores and somewhat resembles the mycelium of the Ustilaginales.^^^ Successful inoculations have not been made.
Ustilaginoidella Essed (p. 199)
a genus erected by Essed ^- to receive the species U. musaeperda, which he regards as the cause of the "Panama
This
is
disease" of bananas, at least as it occurs in Suriname. Sclerotia similar to those of Ustilaginoidea are found; chlamydospores and conidia obtain, among the latter are some of marked
Fusarium type; others are
in pycnldia.
U. oedipigera Essed is also described by Essed ^^ as the cause of another less important banana disease in Suriname and Columbia; a disease accompanied by hypertrophy of the base of the stem and leading to the common name "bigie footoe." This fungus differs from the last in its 1 to 2 to 3-celled conidia. This species U. graminicola Essed causes a rice disease. ^^ differs but slightly from the two preceding. Chlamydospores
smaller, conidia
to 5-celled.
215
LocuHstroraa Patterson
&
Charles
^^-
(p.
199)
Stromata upright,
sessile, at
the nodes of the host, fleshy, soft,
green or black, containing conidial chambers in which are produced hyaline filiform conidia and on the outer surface of which
are borne Cladosporium-like conidia; perithecia scattered, partly
immersed, ostiolate; asci clavate, cylindric, 8-spored; spores fusiform, 3 to many-septate, olivaceous, biseptate; paraphyses none.
There is only one species known. L. bambusae. P. & C.^^^ Stromata 1 cm. long by 2 mm. in diameter; perithecia almost spherical, 125 x 100 m; asci 45-50 x 9-10 n; spores 22 x 4. 5-5 n; primary conidia 14-16 x 0.75-1 n; borne in chambers on basidia, 8 x 0.5 ju; secondary conidia external, 1 to 3-celled, borne on
external olivaceous hyphae. It causes a witches' broom of
bamboo (Phyllostachys
sp.), in
China.
fully
The Infection probably occurs in the terminal node. developed sclerotia-like structures, resembling those of
Claviceps, are dark green to black when mature, and consist of a central hyaline sclerotial tissue in which are many round conidial chambers. Perithecia develop from the peripheral
layer.
Dothidiales
(p.
124)
only one family the Dothidiaceae. in the substratum, septate, at length forming a thick, dense, very dark stroma in which the perithecia are sunken and with which their walls are completely fused, rarely
is
There
Mycelium developed
partly free; asci borne from the base of the perithecium; paraphyses present or none.
species and more from the last order in their They firm black sclerotium-like stromata which are usually pale to white
Dothidiaceffi contain
The
some four hundred

differ
than twenty-four genera.

within.
The perithecia are usually grouped together in great numbers in the external layer of the stroma, sunken in its undifferentiated body. Conidia of various forms are present.
216
Key
Stromata at
free
first
to Genera of Dothidiaceae
later
sunken
more
or less
Perithecia standing free on the stroma; spores at maturity, 4-celled, dark

Perithecia almost completely
1.
Montagnellao
embedded
in the stroma Stromata variable, more or less irregular in outline but never elongate
Spores 1-celled
Spores hyaline Asci typically borne at the

base of the perithecium
Asci 8-spored
Spores eUipsoid
Perithecia few
Perithecia numerous.
.
2.
3.
Mazzantia.
Bagnisiella.
Spores filiform Asci many-spored

Asci
4.
5.
Ophiodothis.
Myriogenospora.
borne laterally at the

equator of the perithecium, spores ellipsoid.
.
.
6.
Spores brown
7.
Diachora, p. 217. Auerswaldia.
Spores 2-celled Spores hyaline Spores ovate

Spores needle-like
8. 9.
Plowrightia,
p. 217=
Rosenscheldia.
Roussoella.
Spores colored Cells of the spore similar. ...

Cells of the spore dissimilar.
.
10.
11.
Dothidea,
p. 220.
Spores several-celled Spores with cross walls only

Spores hyaline, 4-celled Spores colored, multicellular Spores muriform Spores hyaline
Spores colored
14. 15. 12. 13.
Darwiniella.
Homostegia.
Curreyella.
Curreya.
Stromata elongate, linear or lanceolate

Spores hyaline
Spores 1-celled Spores 2-celled Spores 4 to 8-celled, fusiform.
16.
217
Scirrhiella.
17.
.
.
Scirrhia.
18.
Monographus.
Rhopographus.
Spores colored, multicellular, fusi-
form Stromata sunken, permanently united to the epidermis and substratum

Spores 1-celled
19.
20. Phyllachora, p. 220.
Spores 2-celled
Spores of similar
cells
21. Dothidella, p. 221.
Spores of dissimilar cells Stromata from the first superficial
22.
Munkiella.
Hyalodothis.
Stromata encrusted, widely spreading. Stromata cushion-shaped, limited
23.
24. Schweinitziella.
Of these genera only
five are of interest as plant
pathogens.
The majority contain only
saprophytes.
Diachora Muller
(p.
216)
The genus
only as
is
easily recognized
by
its
peculiarity of bearing asci
an equatorial band instead of on the floor of the perithecia, a character unique

cetes.
among
the Pyrenomy-
D. onobrychidis (D. C.) Mull, is reported as causing black spots on

leaves of sainfoin and Lathyrus in
Europe.
Plowrightia Saccardo
(p.
216)
p,^ 154.-0!" onobrychidis. E, conidial stage ;F,ascocarp and asci.
After Muller.
Stromata formed within the tissues

Of the host plant, erumpcnt, tuber-
cular or cushion-shaped, depressed or

elevated, smooth, later frequently wrinkled, white within; asci cylindric, 8-spored; spores ovate, 2-celled, hyaline or light green;
conidial forms Cladosporium,
Dematium,
etc.
218

They
are distinguished from
Some twenty species are known. Dothidia by the hyaline spores.
P. morbosa. b, magnified section of a knot showing the Fig. 155. perithecia; c, conidiophores and conidia; d, section of a perithecium showing numerous asci, one of which is shown more highly magnified at e; f. several of the two-celled ascospores germinating in water. After Longyear.
P. morbosa (Schw.) Sacc^^^'^^''
''^^
Stromata elongate, cushion-shaped, rarely tubercular, up to 2 or
219
3 dm. long; perithecia scattered, often entirely suppressed; asci about 120/1 long; spores variously arranged in the ascus, 16-20 x
8-10 fi, ovate, the cells usually unequal; paraphyses Conidia (= Cladosporium sp.) produced upon greenish areas on the young
stromata; conidiophores erect, flexuose, septate, simple, 40-60 x 4-5 n; conidia borne singly at the apex of the conidiophore, obovate, unicellular, light brown,
filiform.
about 6-8 X 2-5
/z.
Hosts: Cultivated sour cherry and plum, wild red and yellow plum,
red
Chickasaw plum, choke cherry, wild cherry and wild black cherry.
Fig.
Found only in America. The mycelium invades the cambium

of twigs
morbosa; host showing opportunity for lodgement for spores in a

156.
P.
'"S
m
At
this
crotch.
After Lodeman.
and from it grows outward into the bark region causing the bark elements to overgrow and the twig to swell slightly during the first summer. With the renewed growth of the following spring the swelling proceeds rapidly. During May to June the
mycelium ruptures the bark which gous pseudoparenchyma is formed. appear, forming a velvety growth
is
soon
lost
and a dense fun-
From
this the conidiophores
of olivaceous color.
period the knot consists largely of a fungous stroma with an admixture of bark elements and even some wood cells.
Later in the season conidiophores cease to form and the knot turns to a black hard stroma. Perithecia now become easily visible in this black stroma and in January or later the asci mature. Farlow has described " stylospores " (a form named Hendersonula morbosa by Saccardo the connection of which to P. morbosa is
in
some doubt) and spermogonia and pycnidia.
Humphrey
^^^
from ascospores, in artificial media, raised a pycnidial form which seemed to be distinct from any of these. That the fungus is the actual cause of the black knot was first demonstrated by Farlow ^^^ in 1876, though the fungus was described as early as
1821 by Schweintiz.i^s Lodeman ^""^ considered that infection
existing at crotches of the tree.
Fig. 156.
is
favored by cracks
220

is
found in Ribes twigs and on birch. Both are European.. virgultorum (Fr.) P. agaves occurs on the maguey. ^^'^ Dothidea Fries, distinguished from Plowrightia by its colored spores, contains some twenty-five species which occur on twigs of
P. ribesia (Pers.) Sacc. P.
Sacc.
Sambucus, Rosa, Buxus, Betula, juniperus, Quercus and many

other
woody
plants.
D. rosae Fries, is common as the supposed cause of a rose tumor. D. noxia Ruhl. causes an oak twig disease in Germany. ^^^
Phyllachora Nitschke
(p.
217)
of
Stroma sunken, united to the parenchyma and epidermis

the host
leaf, rarely
erumpent, encrusted, usually jet-black; perithecia sunken in the stroma, rather numerous, with more or less distinct ostioles; asci cylindric,
8-spored; spores ellipsoid or ovate, 1-celled, hyaline or yellowish; paraphyses present.
tropical, are
More than two hundred species, largely known on a wide range of hosts.
P. graminis (Pers.) Fcl.
All are leaf parasites.
size
Stromata variable in and form, causing conspicuous black spots

of the host; perithecia
on leaves
7-8
tiolate; asci short-pedicillate, cylindric,

(x;
immersed, os70-80 x
spores obliquely uniseriate, ovoid, hya8-12 X 4-5 n; paraphyses filiform. No conidia are known. FiG. 157. P. gramThis fungus occurs on many grasses and inis. B, stroma in section; C, an ascus and spores. After sedges with slight injury to them.
line,
Winter.
P. pomigena
(Schw.) Sacc. produces black
spots, scarcely ever
above 5
in
mm.
in diameter,
on apples, especially
Little
is
the
Newton known of the

P.
trifolii
Pippin,
species.
the eastern
United States.
(Pers.) Fcl. causes small black spots 1
mm.
or less
in diameter
on clover leaves; 8-10 x 5-6 hyaline,

/jl.
asci cylindric; spores uniseriate, oval,
Conidia
= Polythrincium
trifolii)
precede the asci on the stro-
221
mata; conidiophores wavy or zigzag, erect, simple, black, conidia obovate, 1-septate, constricted, pale olivaceous, 20-24 x 9-10 /z. The conidial form is very common on various species of clover
in
Europe and America while the ascosporic stage
is
mentioned
^'' and Clevenger.^^^ only by Cooke P. cynodontis (Sace.) Niess. on Cynodon, P. poae (Fcl.) Sacc. on Poa and P. dapazioides (Desm.) Nke. on
Box and Rhododendron are European, P. makrospora Zimm. occurs on Durio

linus;
zibel-
P. sorghi v. Hoh. on Sorghum vulgare.^^Dothidella Spegazzini differs from Phyllachora
having 2-celled hyaline spores, the cells un , size, ihere are over .f., equal mty species of the genus. Epiphyllous, subrotund confluent,
in
,
m
.
Fig. 158. D. betulina. Asci and ^^^^^ winter.^'
convex, grayish-black, on white spots; ostiole granular; asci cylindric, short-stipitate, 60-70 x S n; spores obD. ulmi Duv.^^ Colong, ovate oblong, hyaline, 10-15 x 5 /i.
nidia=Septoria ulmi and Piggatia astroidea. On elm in Europe and America. Other species are D. thoracella (Rostr.) Sacc. on
Sedum, in Europe, D. betulina and Asia.
(Fries) Sacc.
on Betula in Europe
Sphaeriales
(p.
124)
Mycelium
chiefly confined to the
substratum; perithecia vari-
able, usually globose, with a
or smooth, free borne on or sunken in a stroma; asci borne basally, variable in size, opening by a pore; spores variable, globose, ovate to elongate
or filiform, hyaline or colored; paraphyses usually present; conidial forms various.
more or less elongated ostiole, hairy on the substratum, more or less deeply sunken, or
crustaceous structure.
The stromata may vary from a delicate hyphal weft to a firm The pycnidia are mostly carbonous, black
brittle.
and
Conidia of many forms are present and often conthe only truly parasitic form of the fungus; the ascigerous form developing only after the death of the part of the host involved.
stitute
222
The order is very large, embracing according to Engler some eighteen families and over six thousand species.
&
Prantl
Key to Famiues
of Sphaeriales
Perithecia free, either without a stroma, partly seated in a loose mass of mycelium, or sessile above an imperfect stroma Walls of the perithecia thin and memasci soon disappearing branous; Perithecia always superficial, with
copious tufts of hair at the mouth Perithecia usually sunken, with only short hairs about the mouth
1.
Chaetomiaceae.
Sordariaceae, p. 224.
2.
Walls of the perithecia coriaceous or carbonous

Perithecia either entirely free, or with the base slightly sunken in the
substratum or stromatic layer
Stroma wanting or only thread-like

or tomentose
Mouths Mouths
of the perithecia mostly in the form of short papilla;. ...

of the perithecia
3.
Sphaeriaceae, p. 225.
more
or
.
less elongate, often hair-like.
4.
Ceratostomataceae,
p. 232.
Stroma present Stromata mostly well developed,

indefinite; perithecia in close irregular masses, never flasklike of funnel-like at the
apex
5.
Cucurbitariaceae,
p. 234.
Stromata
lar
sharp-bordered; perithecia in rows or in regusmall,
shaped
Perithecia
in
rounded masses, flaskwith funnel-shaped

6.
mouths more or
Coryneliaceae.
less
deeply simken
the substratum at base, free

of the perithecia circular in
7.
above
Mouths
outline
Amphisphaeriaceae.
THE FLNGI WHICH CAUSE PLANT DISEASE

Mouths
of
223
the perithecia laterally

8.
compressed Perithecia without a stroma, and sunken
Lophiostomataceae.
in
the substratum, or with a stroma Stromata none; perithecia rarely united above by a black tissue (clypeus)
Asci not thickened at the apex, mostly

projecting at maturity Walls of the perithecium thin,
cori-
aceous;
mouth mostly
short or
plane Asci chnging together in fascicles,
[p.
9.
235.
without paraphyses Asci not fasciculate; with paraphyses Walls of the perithecia carbonous or
thick coriaceous;
Mycosphaerellaceae,
Pleosporaceee, p. 250.
10.
spores
large,
.
mostly enveloped by gelatine. Asci usually thickened apically, opening
11.
Massariaceae, p. 262.
by a pore;
perithecia usually
12.
13.
p. 263.
beaked
Perithecia without a clypeus Perithecia with a clypeus
Perithecia
firmly imbedded in a stroma, the mouths only projecting, or becoming free by the breaking away of the
Gnomoniaceae,
p. 276.
Clypeosphaeriaceae,
outer stromatic layers
Stromata fused with the substratum

Conidia produced in pycnidia Conidia developed from a flattened
surface
14. Valsaceae, p. 277.
15.
Melanconidaceae,
p. 279.
Stromata formed almost wholly ened fungal hypha?

Spores
small,
cylindric,
of hard-
1-celled,
mostly curved, hyaline or yellowish-brown

Spores rather large,
to many-celled, hyaline or brown, conidia mostly in cavities in the stroma
1
16.
Diatrypaceae, p. 281.
17.
Melogrammataceae,
p. 282.
rarely 2-celled, 1-celled, Spores blackish-brown. Conidia devel-
224

oped on the upper surface of the young stroma
18. Xylariaceae, p. 284.
Families Nos.
animals.
1,
6,
7,
8,
17 are saprophytes on plants and
Sordariaceae
(p.
222)
Perithecia superficial or deeply sunken in the substratum, often erumpent at maturity, thin and membranous to coriaceous, slightly transparent to black and opaque; stroma usually absent, if present the perithecia immersed in it with projecting papilliform beaks;
asci
usually
very delicate, cylindric, 8-spored; spores usually
dark-colored; paraphyses abundant. A small order, chiefly dung inhabiting.
Key to Genera of
Spores continuous
Sordariacese
Without a stroma
Neck Neck
of the perithecium hairy

of the perithecium with black
1.
Sordaria.
spines
2.
With a stroma Spores 2 or more

Spores 2-celled
3.
Acanthorhynchus, Hypocopra.
p. 224.
celled
Spores hyaline
Spores dark-brown Spores 4 to many-celled
4. 5.
Bovilla.
Delitschia.
Stroma absent Stroma present

Spores muriform; stroma present
6.
7. 8.
Sporormia.
Sporormiella.
Pleophragmia.
^^^
Acanthorhynchus Shear
Perithecia scattered, submembranous, buried, beaked, the beak with non-septate spines; asci opening by an apical pore; paraphyses present, septate; spores continuous, brownish-yellow. There is a single species, A. vaccinii Sh.^^^
Amphigenous: perithecia subglobose to flask-shaped, scarcely

/z in diameter, neck stout, exserted, ^l^-Û the length of the perithecium; spines 50-70 x 8-9 ix; asci subelliptic
erumpent, 120-200
to
somewhat
clavate, subsessile, 120-155 x 24-44
/x;
spores oblong-

elliptic,
225
surrounded by a mucilaginous layer, 24-32 x 12-18 jj.; paraphyses exceeding the asci. The mycelium produces rot of cranberries, also leaf spots, but the fructification of the fungus is rarely found in nature except on
Fig. 160.
Acan-
thorhynchus; a germinating aaFig. 159. A single perithecium

of A. vaccinii taken from a pure culture on corn meal. After Shear.
cospore bearing the peculiar
appressorium,
17,
view
from
above. Shear.
After
old fallen leaves.

thecia.
When on
In culture, however, it produces abundant perithe leaf the perithecia are subepidermal and
are sparsely scattered over the lower surface.
No
conidial or
pycnidial form is known. Remarkable appressoria are produced by the germ tubes from the spores, Fig. 160.
Sphaeriaceae
(p.
222)
which they are more or

pendaged.
Perithecia single or clustered, free or with a false stroma in less sunken; walls leathery, horny or woody;
ostiole rarely elongate, usually papillate; spores frequently ap-
The family
ostioles.
is
distinguished
by
its free
perithecia with papillate

species.
It contains
about seven hundred
Key
and hairy beneath
Spores
1
to Genera of Sphaeriaceae
Perithecia hairy above, rarely

or 2-celled
smooth above
Perithecia thin, cuticulate or leathery Spores 1-ccllcd; asci apically thick-
ened
1.
Niesslia.
226

Spores 2-celled; thickened
Perithecia thick,
asci
not
apically
2.
Coleroa,
p. 227.
leathery or carbon-
ous
Spores hyahne, sometimes becoming
or 2-cellecl
3. 4. 5.
brown,
Spores elHpsoid
Spores cylindric, bent
Trichosphaeria, p. 228.
Leptospora.
Spores dark colored, 2-celled

Spores more than 2-celled
Perithecia thin, leathery or cuticularized
Neopeckia.
6.
Acanthostigma,
p. 229.
Perithecia thick, carbonous or woody Spores 4-celled, the two middle cells
brown, the end

Spores
cells
hyaline.
..
7.
ChaetosphaBria.
many-celled, hyaline or brown
concolorous,
8.
Spores spindle-form
Spores elongate-cylindric
Perithecia
Herpotrichia, p. 229.
Lasiosphaeria.
9.
smooth
tuberculate
2
or
irregularly
Perithecia
thickened
Spores
Spores
ellipsoid,
to
many-celled,
10. Bertia.
hyaline
spindle-form,
4
to
11-celled,
hyaline
11.
Stuartella.
Spores muriform, dark Perithecia not tuberculate
12.
Crotonocarpia.
Spores 1-celled, dark Spores with hyaline appendages on each end; perithecia thick, leathery
13.
Bombardia.
Spores unappendaged, carbonous Spores 2 to many-celled
perithecia
14. Rosellinia, p. 230.
Perithecia thin, leathery; spores 2celled

15.
Lizonia.
Perithecia
thick,
leathery
or
car-
bonous, brittle
Spores ellipsoid
Spores 2-celled

Spores hyaline, somotiinos hccoiniiijJ!
227
brown
Ki. 17.
IS.
Melanopsamma.
Thaxteria.
Sorothelia.
Spores hyaline to green Spores dark-colored
Spores 3 to many-celled
Spores hyaline Spores dark-colored Spores elongate, spindle-form, hyaline,
19.
Zignoella.
20.
Melanomma.
Bombardiastrum
many-celled
21.
Coleroa Fries
(p.
226)
Perithecia free, small, globose, flask-shaped; asci 8-spored; spores
Fig. 161.
C.
chsetomium.
C, perithecia; D, asci.
After Lindau and Winter.
ovate, 2-celled, hyaline, green or golden-brown; paraphyses poorly
developed.
Comdia=Exosporium. This genus, of some thirteen

is
quite similar to Venturia.
The
species all of which are parasitic, chief economic species are C.
chaetomium (Kze.) Rab. (Conidia=Exosporium rubinus) on Rubus in Europe and C. sacchari v. B. d H., on sugar cane in Java.^^^
228
Trichosphaeria Fuckel
Perithecia usually free, globose,
(p.
226)
or
woody
carbonous, hairy,
ostiole flat or papillate; asci-cylindric,8-spored; spores 1 to 2-celled,
hyaline; paraphyses present. There are some forty species, mainly saprophytes. T. sacchari Mass.^^^' ^^^
Perithecia broadly ovate, dark-brown, beset with brown hairs; spores elongate-ellipsoid, 1-celled; the conidial forms are various
Fig. 162.
Trichosphaeria.
stage.
E, habit sketch; G, conidial After Lindau, Winter and Brefeld.
their genetic connection is by no means certain. (1) (=Coniothyrium megalospora) Pycnidia 1-3, on a dark-colored, parenchymatous stroma; conidia elongate, straight or curved, brownish, 12 X 5 iu, (2) The macroconidia (=Thielaviopsis ethaceticus) see p. 596, are often found forming intensely black, velvety (3) Microlayers lining cracks and cavities in diseased canes. Their conidia produced on the surface in Oidium-like chains. connection with this fungus is disputed and uncertain,^^^ It is a sugar cane parasite.
and
229
Acanthostigma de Notaris
(p.
226)
Perithecia free, globose or ovate, very small; walls leathery,

black, beset with
stiff bristles, ostiolc
short; asci
usually cylindric, rarely
ovate, 8-spored; spores spindle-shaped, multicellular by cross walls, hyaline;
paraphyses few or none. There are some thirty
species,
mostly saprophytes.
A. parasiticum (Hart.) Sacc.^^''"^^^ Perithecia globose, minute, with
rigid divergent
hairs,
0.1-0.25
/j,
mm.
early
in
diameter;
asci
50
long,
disappearing; spores fusoid, straight or curved, smoky, 15-20 /t, continuous

or 2 to 3-septate. Common on leaves of Abies, Tsuga and other conifers in Europe and
America. The hyaline mycelium grows on the lower sides of branches and onto the leaves killing them
and matting them
to the branches.
Fig. 1G3. Peiithecium of A. Trichosphocria parasiticum, showing ostiole, bristles, asci, paraphyses and spores. After Hartig.
The
mycelial cushions later turn brownish and eventually very
small perithecia form on them.
Herpotrichia Fuckel
(p.
226)
Perithecia superficial, globose or subglobose, texture firm, coriaceous to subcarbonous, hairy or smooth, ostiole papillate or not; asci oblong to clavate; spores fusiform, 2 or many-celled,
hyaline or brown; paraphyses none.
The species, numbering about twenty-five and growing on woody plants, are mostly saprophytes. H. nigra Hart.^^^ Mycelium dark-brown, widely spreading, haustoria slender,
lighter in color; perithecia globose,
asci elongate,
76-100 x 12
/x;
in diameter; 1-3 septate. spores constricted,
dark, 0.3
mm.
230
THE FUNGI W HICH CAUSE PLANT DISEASE

in
Europe on branches of Larix, Abies, Juniperus, damage. The dark-bro\\Ti myceUum over the plant, killing and matting the leaves. grows
spruce and
pine, doing great
Common
Rosellinia Cesati
&
de Notaris
(p.
226)
Perithecia superficial, but often with the bases more or less sunken in the substratum, coriaceous or car-
bonous;
not;
brittle, spherical or ovate,
bristly or
spores elliptic, oblong or fusiform, 1-celled, brown or black; paraphyses fusiform. Conidia of the type of
cylindric,
asci
8-spored;
Fig. 1G4.
HerpotriB,
ascus;
chia.
Coremium, Sporotrichum, etc. In most cases the active parasitic stage occurs on roots and consists of a vigorous white mycelium, which remains for a long time sterile, developing large branching and inter.
C, spore. ^"*^^'
After lacmg rhizomorphs (Dematophora) which later become brown. These resemble somewhat, but
/-r^.
are distinguishable from, the rhizomorphs of Armillaria mellea; again, they are Rhizoctonia-like. There are over one hundred seventy species, mostly saprophytic.
R. necatrix (Hart.) Berl.^-
^'^
A destructive fungus, long known as Dematophora necatrix, possesses a white mycelium which invades the small roots, thence passes to larger ones, extending in trees through the cambium and wood to the trunk, occasionally rupturing the bark and producing white floccose tufts. Sclerotia of one or more kinds are produced in the bark and often give rise to conidia on tufted conidiophores in a Coremium-like layer (Fig. 165). The mycelium, when old, turns brown and produces large branching, interlacing rhizomophic strands which spread to the soil, or wind about the
roots.
sterile or
In some instances the connection of the ascigerous with the conidial stages is well established; in others the asci
is
have been found but rarely and the evidence of genetic connection not complete. It is probable that some fungi reported as Dematophora do not in reality belong to Rosellinia.
The fungus
attacks nearly
all
kinds of plants.
THE FUNGI VVmCIl CAUSE PLANT DISEASE

Perithecia wore found
231
and by Prillieux ^^^ on old wood, long dead from such attack. These belong to the genus Rosellinia and are believed to present the ascigerous form of Dema-
by Viala
^^^
tophora necatrix. Similar claims of relationship of this fungus to several other genera have been made and its actual position cannot be considered
as
established
with
KA
?.>-
certainty.
R. massinkii Sacc.
Perithecia sparse, globose
or
depressed,
At;
165
8
IX]
asci cylindric,
carbonous, 54 x
liptic,
spores dark-brown, el1-rowed, 10 x 5 /i.
It is reported
by Halsted
on hyacinth bulbs.
R. bothrina B.
the cause
disease.
of
&
Br.
is
tea root
Pseudodematophora
closely allied to the
above
forms
rens
roots.
is
described
by Beh-
^'^
on diseased grape
quercina
Hart,
is
R.
Fig.
parasitic on roots and stems of Aôung oaks, producing a
1G.5. R. necatrix. 4. corcniiuni and conidia; 5, perithecia extruding spores; 6, asci and paraphyses. After Hartig, Prillieux and Viala.
Rhizoctonia-like mycelium, at first white, later brown. Perithecia are usually abundant. Black sclerotia the size of a pin head are
also present superficially.
R. radiciperda Mas. closely allied to R. necatrix, affects a large
number
potato.
of hosts,
among them
apple, pear, peach, cabbage,
and
An undetermined
berry
disease.'^''
species of this genus
is
said to cause a cran-
Shear, however, berry diseases, did not find it. R. aquila (Fr.) d. Not. injures Morus.
in his extensive studies of cran-
Its conidial
form
is
232
Sporotrichum fuscum. R. ligniaria (Grev.) Nke. occurs on ash R. echinata Mas. is reported on "all kinds of Dicotyledonous shrubs and herbs."
trees.
Melonomma
Fcl. in the species
M. henriquesianum
Bros.
&
Roum. is parasitic on cacao stems. M. glumarum Miy. is on rice.^"^

Ceratostomataceae
(p.
222)
The fungi of this family are very similar to the Sphseriaceae, but are distinguished by less pronouncedly carbonous perithecia which may be merely membranous, and open by an elongate,
beak-like
ostiole.
It
is
a family of only about one hundred
twenty-five species, chiefly saprophytes.
Key to Genera of
Spores
1 -celled
Ceratostomatacese
Spores hyaline Spores brown
1.
Ceratostomella,
p. 232.
2.
Ceratostoma.
Lentomita.
Spores 2-celled
Spores hyaline Spores dark-colored Perithecia on a cottony stroma
Perithecia not on a cottony stroma.
.
.
3.
4.
5.
Rhynchomeliola.
Rhynchostoma.
Spores many-celled Spores with cross walls only Spores elongate, 4 to many-celled, hyaline or
brown
many-celled,
usually
6.
Ceratosphaeria.
Spores
filiform,
hyaline Perithecia erect, astromatic Perithecia horizontal in stromatic
7.
Ophioceras.
nodules
Spores muriform
8. 9.
Cyanospora,
p. 233.
Rhamphoria.
Ceratostomella Saccardo
Perithecia superficial, firm; asci ovate, 8-spored, disappearing
early; spores elongate, blunt or pointed, 1-celled, hyaline. About An extensive study of the genus was made by thirty species.

Hedgcock
^^^
233
who recognizes several species as discoloring lumber. 13 C. pilifera (Fr.) Wint. has been described in detail by von ^ Schrenk as the cause of a blue color in pine wood.'^
Cyanospora Heald
& Wolf
(p.
232)
Perithecia solitary or in clusters of

nodules, immersed, horizontal;
ostiole
two or three on stromatic lateral, neck short; asci
Fig.
Fig. 166.
C.
thecium,
pilifera peri- Fig. 167. asci and tion of
C. albicedrae.
168. C. albicedrte. Upper part of an ascus
Sec-
showing
thickened
spores.
Schrenk.
After
von
a perithecium in its stroma. After Heald and Wolf.
apical wall and coiled After Heald spores.
and Wolf.
apically thick-
slender, linear, surrounded
by a gelatinous matrix,
ened; spores filiform, multiseptate, hyaline.
single species. C. albicedrae Heald
&
Wolf.
Stroma on bark or wood of the host, varying from gray on the bark to black on wood, lenticular, 1-2 mm. long, solitary or clustered; perithecia 825-1200 x 260-400 n; asci 700-1100 x 8-10 m;
spores 600-1000 x 3 m; paraphyses numerous, continuous, in diameter.
1
/x
The fungus
is
described in detail by Heald and Wolf
^^^
as caus-
234
ing whitening of the mountain cedar (Sabina sabinoides) from Texas to Central Mexico. The seat of infection is the younger
twigs and the young trees, especially where in shade. may kill the entire trees.
The
disease
Cucurbitariaceae
(p.
222)
Perithecia clustered, immersed at first, then erumpent, seated on a stroma, leathery to carbonous; paraphyses present. The species numbering about one hundred fifty are mostly saprophytes.
Key to Genera of
Spores 1-celled Asci 8-spored
Spores large, green
Cucurbitariaceae
1.
Bizzozeria.
Spores suuiU, hyaline Asci many-spored

Spores 2 or more-celled
2.
Nitschkia.
Fracchiaea.
3.
Spores 2-cclled
Perithecia bristly, spore walls hyaline. Perithecia smooth, spore walls brown.
.
4.
Gibbera,
Otthia.
p. 234.
5.
6. 7.
Spores more than 2-celled Spores muriform
Gibberidea.
Cucurbitaria, p. 234.
Gibbera Fries
179
Perithecia cespitose on a superficial, thick, Demataceous, conidiabearing, carbonous, fragile, bristly stroma; ostiole papillate; asci
cylindric, 8-spored; spores oblong, elliptic, hyaline, uniseriate. The genus contains some half dozen species, one of which
G.
vaccinii (Sow.) Fr. occurs
on Vaccinium
vaccinii.
in
Europe. The
conidial
form
is
Helminthosporium
Fig. 169.
Cucurbitaria Gray
Perithecia cespitose or
more
rarely gregarious
on a crustaceous
stroma covered by Demataceous hyphse, spherical, glabrous, black, coriaceous; asci cylindric, 8-spored; spores uniseriate, oblong or elliptic, muriform, brownish, paraphyses present.
THE
FU-NDi WlilCii CAl SK
I'LA.X T
DlriEASE
235
of
Over seventy species, several of whicli are parasitic but noii" importance in America. C. laburni Pers. is on branches of Cytisus;
C. sorbi Karst on Sorbus twigs; C. pityophila (Kze.) d Not. on various conif(>r twigs; C. berberidis (Pers.) Gray on Rerberis;
C. elongata (Fr.) Grev. on Robinia; C. piceae Brothwick, on Picea.
Mycosphaerellaceae
Perithecia
(p.
223)
subcuticular, finally or leathery,
mostly subei)idcrmal,
rarely
more
or less
erumpent or even
superficial,
membranous
Fig. 1G9. Gibbera vaccinii.
An
asms. After Winter.
Fig. 170. Cucurbitaria berberidis. G, habit sketch H, ascus. After Lindau

;
and Winter.
fragile; asci fasciculate, 8-spored; spores variable, septate, rarely
muriform, hyaline to dark-brown; paraphyses none. This family of over seven hundred species contains
rophytes and several very important parasites.
many
sap-
Key to Genera of
Spores
1
Mycosphaerellaceae
to 2-cclled
Spores hyaline or green Spores 1-cellcd or not clearly 2-ccllod Perithecia very small, on a basal
growth of thick branched hyplia^
1.
Ascospora,
p.
236.
236

Perithecia
without
such
basal
growth
Spores typically 1-celled Spores usually unequally 2-celled
Spores 2-celled Perithecia produced
2.
.
Massalongiella.
3.
Guignardia,
p. 237.
on
living
4.
plants Perithecia appearing after the death

of the host
Stigmatea,
p. 243.
5.
Mycosphaerella,
p. 243.
Spores dark-colored
Spores 1-celled
6. Miillerella.
Spores 2-celled
Lichen-inhabiting
7.
8.
Tichothecium.
Phaeosphaerella.
Not
lichen-inhabiting
Spores several-celled, hyaUne Spores elongate, with cross walls only

Spores 2 to 4-celled on lichens Spores 4-celled with a cottony subicu; ;
9.
Pharcidia, p. 250.
lum
Spores many-celled Spores muriform
10.
Sydowia.
11. Sphaerulina, p. 250.
12. Pleosph3erulina,p.250.
Ascospora Fries
(p.
235)
Perithecia borne on a subiculum of thick, brown, much-branched
hyphse, globoid, black, carbonous; asci clavate,

clustered, 8-spored, small; spores 1-celled, hya-
171
A~^
Asci.
paraphyses none. half a dozen species, one of which is ^^^ ^^^^ ^y Vuillemin to be the ascigerous form
line;
About
himantia.
mosis.
of
CorjTieum
beyerinckii,
wound
parasite
common on
drupaceous
trees
is
causing
gum-
Cultural evidence of this relationship
lacking, but his
hypothesis may be tentatively assumed. A. beyerinckii Vuil. Perithecia black, depressed-globose, apapil-
100-130 /i in diameter; spores ends obtuse, continuous, hyaline, guttulate, 15 x 5elliptic-fusoid,

late; ostiole indistinct or absent,
7 n.
Conidia,
1.
(=PhylIosticta beyerinckii) pycnidia globoid with
hyaline spores.
THE
Conidia,
FUxXGI
237
2. (=Cor}Tieuin beyerinckii) conidiophores short, from a minute subepidermal stroma; conidia single, crowded, On elliptic-oblong, 1 to 5-septate, brown, about 36 x 15 /x. hosts. drupaceous In spots on the bark the mycelium is often sterile, but when it becomes old distinct pustules usually show in a well developed subepidermal stromatic tissue and from these pustules, as they rupture the epidermis, the conidiophores are produced. Conidia usually abound on the surface of twigs which have borne affected leaves. They germinate readily and produce either a sooty super-
Fio. 172.
Section through a Coryneum pustule on peach.

After Smith.
ficial
mold
or
if
on new bark enter the host tissue and induce
spotting. The conidial stage (Coryneum) of the fungus was grown in arti^^^ ficial culture by Smith but no ascigefous stage corresponding with that of Vuillemin was found.
fruits
A. geographicum (D. C.) Desm. is common on leaves of and A. padi Grev. defoliates cherries in Europe.
pome
Guignardia Viala
& Ravaz
(p.
236)
Perithccia sunken, globoid or flattened, black, leathery; ostiole flattened or papillate; asci clavate, 8-spored; spores elhpsoid or fusiform, hyaline, somewhat arched, 1 or 2-celled; paraphyses
none.
238

Some
are impor-
Over one hundred thirty species are known.

tant parasites. Conidial forms are found in
Phoma and
'^'""^
Phyllosticta.
G. bidwellii (E.) V.
asci
&
R.^'
Perithecia minute, globose, subepidermal, erumpent, perforate; clavate-cylindric, obtuse, 60-70 x 10-13 fx; spores elliptic to
/x.
oblong, continuous, 12-17 x 43^-5
Conidia
= Phoma
uvicola, Phyllosticta labruscae,

n,
Nsemospora
elliptic.
ampelicida) borne in pycnidia 180 x 180
subepidermal,
8(76CO6p0re5 '^
âa"
mqermm/zo/r
Fig. 173. Diagrammatic section of a perithecium conGermination of a spore at the taining ascospores. right. After Reddick.
thick-walled; conidiophores short, simple; conidia ovate to elliptic, 8-10 X 7-8 n. Filiform microconidia ("spermatia") are borne in
flask-shaped pycnidia 0.1-0.2 x 0.45-0.46 n.

successively in the genera Sphseria, Lsestadia and Guignardia. Physalospora, CO An extensive synonomy is given by E. Rose who concludes
rt
The fungus has been placed
that the
It is
ampelicida. green parts of Vitis and Ampelopsis, the ascigerous stage common only on the mummified fruits. Perithecia were first found in 1880 by Dr. Bidwell in New Jersey.
name should be G.
all
found on
They
are abundant on berries, which have wintered out doors. Reddick admirably describes the development of the spots as
follows:

On the
leaves the
239
first evidence of the spot is the sH^ht blanching one of the smaller areola of the leaf. Soon the blanching extends to adjacent arcoliE, and if an areola is entered it is usually entirely involved. The small veinlets form the margin of the spot so that the outline is finely crenulate. By the time
of a single
the spot
is .3
to
.4
mm.
in
diameter
it is
The margin,
while sharply defined,
has a cinereous appearance. not changed in color. By
the time the spot is 1 mm. in diameter, the margin appears as a black line, while the remainder of the spot is grayish-brown. A
Spores...^
0cy=
Germinated spores
Fig. 174. Diagrammatic section through a pyciiidium, showing how the spores are produced and how they germinate. After Reddick.
the margin is a brownish band and the browTi gradually extends inward until the whole spot is covered. As soon as the browTi band attains some width the blackish line on the margin
little later
to be seen again. A second wave of deeper brown may pass across the spot but sometimes it does not get entirely across and thus leaves a marginal band of a deeper bro^\^l than the central
is
disc.
Spots vary in size from
in general are 3 to 5
mm.
destroyed but this is by the spot has attained full size pycnidia protrude from under the cuticle and either dot the entire surface of the spot \\'ith minute
specks or are more often confined to a more or less concentric ring. The different shades of color are apparent on the under side of
1 mm. up to 8 mm. in tliamcter, but or larger. Occasionally the whole leaf is the coalescence of many spots. When
240
the leaf on such varieties as have leaves which are smooth beneath.
The pyncidia, however, have never been seen of the leaf in our varieties.
on the under
side
On
stems,
tendrils,
spot in its first
peduncles, petioles and leaf veins the appearance is a small darkened depression which
soon becomes very black. On a cane the lesion rarely extends more than a quarter of the way round, while on a tendril or leaf
petiole
it
may extend from half to
all of
the
way
round.
On shoots,
the lesions never extend so
deep as to cut off the sap supply, but on petioles this

occasionally happens, rarely so on peduncles, and quite commonly so on pedicels and
tendrils.
The
first
indication
of
is
Black Rot on the berry the appearance at some

of
point
Fig. 175. Section of a pustule showing microconidia. After Longyear.
small
circular
1
blanched spot, scarcely

in diameter.
mm.
The blanching
is so slight as to be detected only by careful observation. It rapidly becomes more apparent and has a whitish appearance, the contrast becomes more ap-
gin.
parent by the appearance of a brownish line at the marThe whitish center increases in size and the brownish or
reddish-brown ring increases in diameter as well as in width and is quite evident when the spot is 2 mm. in diameter. When the spot is 3 mm. in diameter the ring is one-half mm. in width and enough darker to give a bird's eye effect (a light circular disc with an encircling darker band). The spot rapidly increases
more it may be 6 to 8 mm. in diamband nearly 2 mm. in width. After five hours more, the spot is 8 or 9 mm. in diameter and there begins to appear an outer darker band and an inner lighter brown one which have in some cases a much lighter line between them. The aureole is thus composed of two or three bands or rings. Eighteen hours later, the spot is 1 cm. or more in diameter, is distinctly flattened, and numerous minute brown specks appear on the
in size so that in twelve hours
eter,
and the
encircling

white center of the spot.
241
In
five
hours more they are so numerous
as to give a blackish appearance. In New York, Reddick found that the asci begin to ripen in May and continue to mature throughout the summer being still abundant
in October.
The
asci swell in
water often to twice the length
given above; spores are forcibly ejected from the asci at maturity, being thro\vTi to a height of
There is at one end of the ascoa hyaline vesicle which probably aids in spore fixing it to the host.^^^ They germinate but slowly, requiring from thirty-six to forty-eight hours to show germ tubes. Reddick determined the incubation period on fruit as from eight to twenty-one days and found that only
2 to 4 cm.
tender leaves
The berry
has
fallen.
is
still growing are susceptible. susceptible even after the calyx The pycnidial spores are said by
^^^
show a hyaline appendage others by careful study fail to some

to
1 . I
though
it.^^'^
find
These spores often live over winter. ^^ The ... T microconidia which develop m pycnidia similar to those of the macroconidia do not occur so abundantly early in the season as they do later and seem to be mainly limited to the
1 1

Fig-
176. g.
bid-
-1
wellii; ^5,
ture
nearly maascus with
-'
''
...
Iscospores-' ;2^germinating ascospores.

29,
same with
pressoria.
apAfter
fruits.
occur and
seen.
Sporeless pycnidia, pycnosclerotia, also may eventually develop into perithecia.
Conidia on
hyphse of questionable relationship to the fungus are
sometimes
Reddick
1.
secured pure cultures in the following ways. In poured plate dilution of asci; some twenty days were
^^^
required.
inverting a plate of sterile agar over a bunch of mature The ejected ascospores thus clung floating on water. to the agar and gave pure cultures in ten days.
2.
By
mummies
3.
4.
aseptic transfer of the mycelium. aseptic transfer of pycnospores. Artificial infections have been reported in
By By
Europe from both

in-
conidia and ascospores: Reddick,
who made many thousand
242
oculations

under
all
conceivable conditions, failed utterly of posi^^^
tive results.
From
the Caucasus Prillieux and Delacroix
have described a
Guignardia causing a black rot of grapes which is regarded as distinct from the usual American form, differing both in the perithecial and conidial stages. This is called G. baccae (Cav.) Jacz.
Its conidial
form Phoma reniformis eventually covers the whole

berry with pustules.
are described.
Two
^''
kinds of pycnidia
G. vaccinii
Sh.^^^'
Perithecia on
young
fruit or flowers, sub-
epidermal, globose, walls thick,

asci
^
carbonous;
/^^^
Fig.
^^
177. A
vertical sec-
oTcu'ignardi'avaccinii!
showing
Shear.
asci.
After
60-80 n long; spores elliptic or subrhomboidal, hyaline, becoming tinted. Conidia (=Phyllosticta) borne in pycnidia similar to the perithecia but thinner- walled, 100-120 Mi conidia hyaline, obovoid, 10.513 5 ^ 5-6 '^ On Vaccinium. In the decaying berries all sporing forms
clavate,
,jl.
of
the fungus are rare though in the softened tissues fungous hyphae abound. Transferred to culture media these hyphse grow
readily
and produce spores abundantly.

conidial form
is
The
thetical
common
rare.
in artificial culture; the peri-
Pycnidia on leaves are suband are quite abundant. The epidermal, usually hypophyllous, spores at maturity issue in coils from the ostiole. The fungus was studied extensively in artificial culture by Shear, wet sterilized cornmeal proving a most suitable medium. Pycnidia appeared in four to eight days after inoculation and spores were mature at twelve to eighteen days. Both pycnidia and perithecia
form comparatively
were obtained in pure cultures. The rarity of cultures able to produce perithecia is explained by Shear on the assumption "that
there
is
some inherent
potentiality in the
mycelium
of the fungus
it
in certain strains, races, or generations
which causes
to produce
the ascogenous stage whenever conditions for its growth are favorable, i. e., on favorable culture media without special reference to
their exact composition or
environment or on the leaves of
its
nat-
ural host."
Conclusive infection experiments have not been made.

G. theae Born. G.
leaves.
'^^
243
^rows on tea leaves.
(Laestadia)
It is
The perithecia develop on box probably saprophytic although sometimes considered

buxi Desm.
a parasite.
Stigmatea Fries
Perithecia
(p.
236)
thin,
subepidermal,
or
subcuticular,
black;
asci
oblong, subsessile, 8-spored; spores ovoid-ellipsoid, 2-celled, yellowish or hyaline; paraphyses present. The
ascigerous
stage of
two
species
of
Entomo-
to belong to sporium are said by Lindau this genus. Atkinson, however, places them in the genus Fabrea, see p. 149. S. juniperi (Desm.) Wint., on living leaves of Juniperus in Europe and America and on ' Fig.
. .
. .
, 1<8.
. fetigmatea.
Sequoia in California.
Perithecia scattered, lenticular or subhemispheric, rough,
Asci
and
spores.
in diameter, asci rounded and obtuse 200-300 above, abruptly tapering below into a short stipe, 60-70 x 20 ju; spores ovate-lanceolate, unequally 2-celled, yellowish-hyaline, 16fj.
25 X 6-8
n.
S. alni occurs
on alder leaves
in
Europe.
Mycosphaerella Johans.
(p.
236)
Perithecia subepidermal, suberumpent, globose-lenticular, thin, membranous, ostiole depressed or short papillate; asci cylindric to clavate, 8-spored; spores hyaline or greenish, ellipsoid, 2-celled;
five hundred species formerly known as It is often Sphserella contains several serious plant pathogens. found in its conidial forms as: Ramularia, Ascochyta, Septoria,
paraphyses none. This large genus of over
Phleospora, Cercospora, Ovularia, Cylindrosporium, Phyllosticta, Graphiothecium, Phoma, Diplodia or Soptoglocum. In many cases the relationship of the ascigerous and conidial forms is as yet but imperfectly known. The perithecia are usually found late in the season, often only on leaves that have borne the conidial stage in the summer and have then wintered.
244

fragariae (Tul.) Lin.^^^
M.
Perithecia on leaves, are produced late in the season, globose,
subepidermal, membranous, black, thin-walled; asci few, clavate,
Fig. 179. Mycosphserella fragariae. b, conidiophores bursting through the epidermis; c, arising from apex of a pycnidium; d, summer spores, one germinating; e, section of a spermogonium; /, section of peritheeium; g, ascus containing eight two-celled spores. After Longyear.
8-spored, 40 n long; spores hyaline, 2-celled, 15 X 3-4 At.
with acute
tips,
Conidia (=Ramularia tulasnei) abundant in early summer on reddish spots, stromatic, conidiophores simple; conidia elliptic 20-40 X 3-5 fi, 2 to 3-celled. On Fragaria.
The life history was first studied in 1863 by the Tulasne brothers under the name Stigmatea. The generic name was changed to Sphaerella in 1882 and later to Mycosphserella.
245
The slender mycelium pervades the diseased areas disorganizing the host cells and resulting in reddish coloring of the sap. Observations of Dudley ^^^ indicate that the mycelium or portions of it can remain alive over winter in the host tissue ready to produce
abundant conidia in the spring. The most abundant conidial stage is the Ramularia-form (Fig. 179) which abounds all summer. Sowings of these conidia, under conditions of humid atmosphere, result in characteristic Toward winter sclerotial spots in from ten to eighteen days. bodies are formed from the mycelium. These in culture dishes have been seen to produce the typical summer conidia. Some of these sclerotia-like bodies have been reported as "spermogonia," bearing numerous "spermatia" 1 x 3 m- Perithecia abound in autumn. These are larger than the spermogonia and are usually embedded in the leaf tissue, though they sometimes appear superficially. Conidiophores are often borne directly on the perithecium
wall.
resulting
Ascospores germinate within the ascus. From the mycelium from ascospores Dudley ^^^ observed the formation of
typical
summer
conidia.
^^^
M.
grossulariae (Fr.) Perithecia hypophyllous,

ostiole, black;
gregarious,
spherical,
asci short-pedunculate, clavate,
with minute 55-66 x 8-12 n;
spores fusoid, filiform, curved or straight, uniseptate, hyaline,
26-35 X 3-4 n. It has been reported on the gooseberry associated with Cercospora angulata and Septoria ribis. M. rubina (Pk.) Jacz.ôo
late,
Perithecia minute, gregarious, submembranous, obscurely papilsubglobose or depressed, erumpent, black; asci cylindric,
70-80 x 10-12 )u; spores oblong, obtuse, uniseptate, constricted in the middle, 15 x 6-7 n, upper cell broadest. generally Conidia ( = Phoma) are associated with the perithecia and are
subsessile,
supposed to be genetically connected with them as spore form ( =Coniothyrium).
is
also a second
The
species
is
held responsible for bluish-black spots on raspis
berry canes.
M.
cerasella Aderh.^"^
reported as the perithecial stage of

cherry.
Cercospora cerasella
common on
246

Conidia
(
247
= Diplo(lina
n.
citriillina)
Pycnidia similar to the peri-
thecia, spores 2-celled, hyaline, straight or curved,

cylindric,
more
or less
10-18 x 3-5
The fungus was isolated in pure culture by Grossenbacher ^"^ from muskmelons by direct transfer of diseased tissue to potato
agar. Inoculations from these cultures proved the fungus capable of entering healthy uninjured tissue, the
disease showing about six days after inoculation. The brownish pycnidia origi-
nate
from
an
extensive
subepidermal,
much-branched, brownish mycelium but soon break through and M. sentiiiu. A, FiG. 182. r ^^Tl appear almost superfacial. When moisperithecium and asci. Af^^^ Klebahn. tened, spores issue in coils. Darker periBoth thecia, nearly superficial, are found on old diseased spots.
partially cortical,
,
,
ascospores and conidia are capable of causing infection.

lations
Inocu-
on pumpkin and watermelon gave positive results; these on cucumber. West Indian gherkin, squash, pumpkin, and gourd were negative. The same fungus has been reported as cause of
canker of tomatoes. -''^
M.
tabifica (P.
&
D.) Johns.ô^-z^"
Perithecia rounded, brown; asci oblong-clavate, 8-spored; spores hyaline, upper cell larger, 21 x 7.5 /x.
3.5
Pycnidia (=Phoma) subglobose; conidia fi, escaping as a gelatinous cirrus.
elliptic, hyaline,
5-7 x
This conidial form, common on beets causing leaf spot throughout the summer, is said by Prillieux and Delacroix to be connected with M. tabifica the perithecial form, which is found upon the
dead
petioles at the
this connection
end of the season. Convincing evidence "^^ of seems wanting. The conidial stage ^^ is variously
beta,
known
tabifica.
as
Phoma
Phoma
The Phoma-form from stems and
sphserosperma, Phyllosticta rotten roots and the
^^" Phyllosticta-forms from leaves were both studied by Hedgcock in pure cultures on many media and many inoculations were made,
leading to the conclusion that the are identical.

all
Phoma and
the Phyllosticta
M.
tulasnei Jacz.-"
Perithecia subglobose,
minute; asci cylindric fusoid; spores
248

cell in
oblong, rather pointed, upper than the others, 28 x 6.5 fi.
the ascus somewhat larger
Conidia of two kinds, (1) (=Cladosporium herbarum) tufts forming a velvety blackish-olive, effused patch, conidiophores erect, septate, rarely branched, often nodose or keeled;
dense,
conidia often in chains of 2 or

3-septate,
3,
subcylindric pale-olive,
to
10-15x4-7
Sacc.)
Hyphae
cladosporioidies erect, simple, bearing apically or laterally a tuft

/x.
(2)
= Hormodendrum
M. citrullina, A. pycnidium (Diplodia) in secFig. 183. After Grossention, B, perithecium; C, ascus and spores. bacher.
of small, eUiptic, continuous, chains.
brown conidia
in simple or
branched
Europe, being especially injurious to cereals after a rainy season preceded by a drought and is found also parasitic on pea, apple, raspberry, cycad, agave and as a saprophyte almost anywhere.
It is the cause of serious disease in
M.
stage alone
stratiformans Cobb, affects sugar cane. The perithecial ^^^ is known. Further study is desirable.
gossypina (Cke.) Er.^^^-si^ Perithecia ovate, blackish, partly immersed, 60-70 x 65-91 fx, asci subcylindric, 8-10 x 40-45 /i; spores elliptic to fusoid, constricted at the septum, 3-4 x 15-18
At.
M.
Conidia
(=Cercospora gossypina);
hyphse
flexuose,
brown.
249
120-150 ^l high; conidia attenuate above, 5 to 7-septate, hyaline, 70-100 X 3 M. On cotton. The intercellular mycelium is irregular, branched, septate, and produces tuberculate stromata from which the brownish hyphae
arise.
The
perithecia,
much
less
common,
are partly
immersed
in old leaves.
M.
morifolia (Fcl.) Lin. in
its conidial stages,
Cylindrosporium
trees.
mori and Septogloeum mori, affects Morus. M. maculiformis (Pers.) Schr. grows on
many
Especially
common
are
its
conidial stages Cylindrosporium castinicolum
and
Phyllosticta maculiformis. M. rosigena E. & E. -'8-219
Amphigenous on reddish-brown, purple-bordered spots which
mm. in diameter; perithecia thickly scattered over the spots, minute, 60-75 ju, partly erumpent, black; asci subclavate to oblong, 25-30 x 8-10 /x; spores biseriate, clavateoblong, hyaline, 1-septate, 10-12 x 2 /i, ends subacute.
are about 3-4
It causes leaf spots of rose in
(
America.
M. brassicaecola = Phyllosticta brassicsecola) grows on cabbage. M. punctiformis Pers. produces leaf spot on oak, lime, hazel; M. fagi Auser. on beech; M. pinifolia Due. on pine leaves; M. abietis (Rost.) Lin. a leaf disease of balsam.^^ M. taxi Cke. grows on yew; M. hedericola Desm. on Hedera leaves; M. gibelliana Pass, on Citrus leaves; M. vitis Fcl. on grape leaves; M. elasticae Kr. ^-^ on Ficus elastica. M. cydoniaB Vogl. ^^^ on quince is probably identical with M. sentina on pear and apple. M. ulmi Kleb. occurs on elm with its conidial forms, a Phleo^^'^
spora and Phyllosticta bellunensis. M. comedens Pass, is on the same host.
M.
larcina Hart, and
its
conidial
form Leptostroma larcinum
affect larch, causing defoliation.
M. M.
Icfifgreni
N. on oranges and
M.
is
coffeae
N. on
coffee are
tropical forms.
populi Schr. (=Septoria populi)
on Populus.^*^
250

pinodes Berk
M.
12
ii;
&
Blox.
/x;
Perithecia numerous, 100-140
asci oblong-cylindric,
58-62 x
spores 2-rowed, 14-16 x

pisi),
5.
Pycnidia (=Septoria
3.5 n, 1 to 3-septate,
with large
ostiole; spores
leaves.^"^'
^^
35-45 x 3-
On pea stem and
M. primulae is on primrose; M. tamarind! on tamarinds in Africa. M. cinxia Sacc. is on lilies, causing leaf blight; M. fusca Pass, on the gladiolus; M. coffeicola on coffee in Mexico. M. shiraina Miy. and M. hondai Miy. are on rice. M. convexula (Sch.) Rand.
pent, 100-200
asci
Perithecia hypophyllous, gregarious or scattered, finally erumju in diameter, papillate at maturity; no paraphyses;
fasciculate,
54-100 x 9-11
ix, /t.
8-spored;
spores allantoid,
1-septate, hyaline, 13-27 x 3.5-5.5
leaf spot on pecans.^^^ undetermined species of Mycosphaerella has been reported on the grape by Rathay.^"^
Forming a
An
Many
other species are
known on
ferns, cereals,
lilies,
and va-
rious trees
and
herbs.
In the genus Pharcidia. P. orzae Miy. is on rice.^*^^ In Sphaerulina the species Sphaerulina taxi Mass. is injurious on
yew
leaves.
Pleosphaerulina Passer
(p.
236)
Perithecia subepidermal, erumpent, small, globoid or lenticular,

black; asci 8-spored, clavate; spores muriform, hyaline; paraphyses none.
P. briosiana Pol. causes a leaf disease of alfalfa in Italy.
Pleosporaceae
(p.
223)
Perithecia sunken, at length erumpent, or from the first more or less free, membranous or coriaceous, usually papillate; asci
clavate-cylindric, double-walled; spores variable, but usually colored, oblong, fusoid or elliptic; paraphyses present. An order of some nineteen hundred species most of which are
saprophjês, although several are parasites, some of considerable
importance.
251
Key to Genera of
Pleosporaceae
Spores l-ccUed Spores with blackish appendages, elongate, hyaline

1.
Urospora.
Spores unappendagcd
Spores elongate, hyaline or light yel-
low
Spores elongate, fusoid, hyaline; tips bent
2.
Physalospora,
p. 252.
3.
Therrya.
Spores 2-celled Spores with the 2

size
cells
very unequal in
Upper
Basal
cell
the smaller; parasitic on

4.
...
5.
Riccia
cell
Arcangelia.
the smaller; saprophytes.

cells
Apiospora.
Spores with both

Perithecia
about equal
spores
hairy;
hyaline
or
6.
brown
Perithecia smooth
Venturia, p. 253.
Spores hyaline Spores brown Perithecia not stromatic

Perithecia borne on a stroma
7.
Didymella,
p. 255.
8. 9.
Didymosphaeria,p.256.
Gibbellina, p. 256.
Spores more than 2-celled Spores elongate, with cross walls only Spores appendaged Spores clavate, 4 to 6-celled, brown, the basal cell hyaline long-
appendaged
Spores
filiform,
10.
Rabentischia.
Dilophia, p. 257.
many-celled,
with
11.
filiform
appendages
Spores not appendaged Spores fusoid or elongate, blunt, never

filiform or separating into cells Spores elongate, 3 to many-celled,
hyaline or brown Spores with a thick, dark-brown epispore and a thin hyaline
endospore, 4-celled,
cl-
hpsoid
12.
Chitonospora.
252

Spores not as above, elongate 3 to many-celled hyaline or
brown
Perithecia hairy Perithecia smooth
13.
Pocosphaeria.
Spores hyahne Spores

yellow
or
14.
Metasphaeria,
Leptosphaeria,
p. 257.
dark15.
brown
Spores fusoid, 7 to many-celled, the central cell enlarged and brown, the rest hyaline Spores fusoid, up to 30-celled hyaline or
p. 257.
16.
Heptameria.
Saccardoella.
brown
separating into
17.
Spores
filiform, often
cells
Perithecia hairy Perithecia smooth/
18.
19.
Ophiochaeta.
Ophiobolus,
p. 259.
Spores muriform Asci 8-spored

Spores appendaged Spores not appendaged
Perithecia hairy Perithecia
20.
Delacourea.
21.
22.
Pyrenophora,
Pleospora,
p.
262.
smooth
p. 259.
Asci 16-spored
23.
Capronia.
Physalospora Niessl.
Perithecia
(p.
251)
subglobose,
covered,
membranous, or coriaceous, black, with the ostiole erumasci clavate-cylindric spores ovoid or oblong, continuous, hyaline or subhya-
pent;
paraphyses present. This genus contains over one hundred thirty species, a
line;
few
"
of
which are parasitic on

leaves.
^
184. Physalospora
ascus.
twigs and
Perithecia and
VV inter.
Some
spe-
FiG.
After
^ pOSSeSS a Gloeosporium ^ as the conidial form.

gjes
P. gregaria and
its
conidial
stages
Tetradia salicicola and

Macrodcndrophoma
Ireland.
224
253
oziers in
salicicola cause black cankers
on
found on Picea; P. cattleyae Maub. & Las. in its conidial form, Gloeosporium macropus--'^ parasitizes Cattleya. P. laburni Bon. is on Cytisus. P. woronini JNI. & F. is described as causing a disease of grapes
P. abietina P.
D.^^^
is
&
in the Caucasus.
^^^
P. vanillae
Zimm.
is
on
vanilla;
P. fallaciosa Sacc. on banana leaves.
Venturia Cesati
&
de Notaris
(p.
251)
Perithecia superficial or erumpent, bristly, ostiolate,
membra-
nous, dark colored; asci sessile or short stipitate, ovate or saccate; spores oblong to ovoid elliptic, hyaline or yellowish; paraphyses
usually none.
The conidial stages in some cases belong to the form genus Fusicladium and constitute the parasitic portion of the life history
form usually being limited to old or wintered parts of the host. There are over fifty species, several of which cause diseases.
of the fungus, the ascigerous
V- Aderh. pinna
230, 312, 313, 350 '
Perithecia
gregarious,
smooth
or bristly, globoid, 120-160
/x;
asci cylindric; spores
unequally 2-celled, yellowish-green, 14-20 x
5-8 MConidia (=Fusicladium pirinum) effused, velvety, blackish-olive, conidiophores short, wavy or knotted, thick- walled; conidia ovate fusoid, olive, becoming 1-septate with age, 28-30 x 7-9
/jl.
found on the pear wintering in perithecial form on leaves, and in conidial form, or as mycefium on twigs. V. inaequalis (Cke.) Aderh. (=V. pomi [Fries] Winter). Perithecia globose, short-necked, 20-160 smooth or bristly above; asci cylindric, 40-70 n long; spores yellowish-green, unequally 2-celled, upper cell shorter and broader, 11-15 x 4 -8 /i. Conidia ( = Fusicladium dendriticum) effused, velvety, forming dendritic patches of compact masses of erect closely septate brown mycelium; conidiophores closely septate, brown, 50-60 x 4-6 wavy or nodulose; conidia solitary, terminal, obdavate,
It is
ij.,
fjL,
254
Fig. 185. V. inoequalis. A, portion of a section through a scab spot on apple; 6, spreading under and lifting the cuticle, a; c, partly disorganized cells of the apple; e, healthy cells of the apple. B, two conidiophores with summer spores /. C, spores germinating. Z), portion of a section showing a perithecium and asci. E, two asci, each containing 8 two-celled spores, three of which are shown at F. After Longyear.

yellowish-olive, continuous
255
when young but
at
length septate,
30 X 7-9
IX.
Its hosts are apple and other pomaceous fruits except the pear. Conidia of special form have been known under the name Napicladium soraueri.
The two last conidial forms have been long regarded as identical and are found in literature as Fusicladium dendriticum. The
both cases grows subepidermally in the the epidermis and forming subepidermal killing Stromatal stromata from which conidiophores are produced.
olive-green
leaf
mycelium
in
and
fruit
development
is
also said to be often subcuticular, resulting in a
separation of the cuticle
from the epidermis. conidia are produced apically on short stalks and as each conidium is cut off the conidiophore grows forward, leaving scars
The
number to the conidia produced. Pycnidia have been reported on the mycelium in twigs in winter.^^^ Perithecia first form on the lower leaf surface in October and
equal in
in April. They are most abundant when protected by sod or piles of leaves, and appear as small black pustules often on grayish spots. Their connection with the conidial stage was
mature
shown by Aderhold -^^ and confirmed by Clinton. -^^ The fungus from apple was cultured on apple-leaf-agar by Clinton. Pure colonies developed in 4 to 5 days and infection was secured on leaves. Cultures from ascospores gave rise to typical conidia. V. crategi Aderh. occurs on Crataegus. V. cerasi Aderh. ( = Fusicladium cerasi) is found on cherries.
first
Aderholt
^^^
demonstrated the connection between the ascigerous
and
conidial forms.
V. ditricha (Fr.) Karst. ( = Fusicladium bctulse) is found on = birches; V. tremulae Aderh. ( Fusicladium tremulse) on aspen; V. fraxini Aderh. ( = Fusicladium fraxini) on ash; V. inaequalis var. cinerascens Lin. (= Fusicladium orbiculatum)
on Sorbus.
Didymella Saccardo
(p.
251)
Perithecia covered, mcml)ranous, globose-depressed, minutely papillate; black; asci cylindric or clavate* spores ellipsoid or
ovate, 2-celled, hyaline; paraphyses none.
256

is
terest since
Of the some one hundred twenty species D. citri N. it forms cankers on orange trees in Brazil.
Didymosphaeria Fuckel
(p.
of in-
251)
Perithecia immersed, later erumpent; asci cylindric to clavate, 8-spored; spores elliptical to ovate, 2-ceUed, brown.
This genus
spores.
differs
from Didymella
cfhiefly in
the dark-colored
It contains
occasional parasitic
some one hundred twenty species and has representatives on leaves and twigs.
Fig. Fig. 18G. Didymella. A, ascus; B, hymenium of a pvTnidium. After Brefeld.
187.
Didymo-
Fig.
188.
DiloAfter
sphaeria.
C, an aso II i d i ocus; D, phore and oonidia. After Brefeld.
p h i a graminis. J, ascus; K,
spore.
Winter.
D. sphaeroides (Pers.) Fr. D. catalpae.^^^

leaf,
is
on Populus leaves
in
Europe.
Perithecia very small, scattered, embedded in the tissue of the pyriform to nearly spherical, varying in width from 48-104 n
in
and
asci 8-spored, cylindrical, usually
depth from 64-140 n; ostiole broadly conical, erumpent; somewhat curved; paraphyses
few or wanting; spores oblong-elliptical, hyaline or yellowish, uniseptate, constricted in the middle, 9.6-13 x 3-4 /x. On Catalpa. D. populina VuilL, causes death of poplars in Europe.-^^
D. epidermidis
Fr.
is
found on Berberis, Sambucus and

(p.
Salix.
Gibbellina Passerina
251)
Stromata black, sunken in the substratum, formed of thin, closely interwoven hyphae; perithecia sunken in the stromata, globose;
THE
FUNCJl W
HIGH CAUSE PLANT DISEASE
257
asci elongate-globoid; spores elongate, 2-ceIIed,
brown; paraphyses
present. Genus of one species. G. cerealis Pers. causes a serious grain disease in Europe, es^^^ pecially of wheat in Italy.
Dilophia Saccardo
(p.
251)
Perithecia sunken, not erurapent, delicate, dark-colored, ostiole

papillate; asci long-cylindric; spores elongate-fusiform to filiform,
end appendaged, the appendages hyaline, the or yellow. Fig. 188. spores hyaline There are three species, one of which D. graminis (Fcl.) Sacc. The conidial form Diloparasitizes rye and wheat in Europe.
multicellular, each
phospora graminis Desm.
is
especially
common.
(p.
Metasphaeria Saccardo
252)
Perithecia clavate, sunken in a stroma, at first covered; leathery, dark, with ostiole asci cy lindric to clavate, 8-spored spores ellipsoid,
;
elongate, blunt or appendaged, 3 to many-celled; paraphyses filiform.
M.
albescens Thum.
is
on
rice in
Japan.
(p.
Leptosphaeria Cesati
Perithecia at
first
&
de Notaris
252)
subepidermal, at last more or less erumpent,
subglobose, coriaceo-membranous, globose, ostiole usually papillate; asci subcylindric; spores ovoid, oblong or fusoid, two or more
septate, olivaceous, yellowish or brown. There are about five hundred species,
conidial forms
many of which in the embrace Cercospora, Phoma, Hendersonia, Sporidesmium, Soptoria, Coniothyrium or Cladosporium.
L. coniothyrium (Fcl.) Sacc.^^^'
^^'^
Perithecia gregarious, subepidermal, depressed, globose, black;

papillate, erumpent; asci cylindric, stipitate, 8-spored, 66-96 X 4-6 fjL] spores 1-rowed, oblong, 3-septate, constricted, fuscous, 10-15 X 3.5-4 ju.
ostiole
Pycnidia (= Coniothyrium fuckelii), similar to perithecia; spores ovate, continuous, fuscous. It occurs on black and red raspberries and numerous other hosts. Stewart ^" verified the assumed identity of the conidial form
with this ascigerous fungus by pure culture studies.
258

tritici
^^^
(Gar.) Pass (=Pleospora tritici). On wheat.^^ Perithecia innate, globose, black, papillate; asci clavate, short-
stipitate,
8-spored; paraphyses filiform, 48-50 x 15-16
/x;
spores
2-seriate, round, fusoid, 3-septate, constricted, pale,

5.5.
18-19 x 4.2-
L. herpotrichoides d. Not.^^^ parasitizes rye causing the stalks

i
to break at the nodes;
FiQ. 180. Cross-section of raspberry bark showing two perithecia of L. coniothyriuni at the top, A, and two pycnidia of Coniothyriuni fuckelii, at the bottom, B. 4. An ascus of L. coniothyrium. 5. Spores of L. coniothyrium. After Stewart.
L. sacchari V. B. d H. occurs on sugar-cane. L. napi (Fcl.) Sacc. (=Sporidesmium exitiosum)
is
found on
rape; L. phlogis Bos. (=Septoria phlogis) on Phlox; L. circinans (Fcl.) Sacc. kills alfalfa roots, potato, clover, beets
and other hosts;
~^^
L. vitigena Schul. occurs on grape tendrils; L. stictoides Sacc. on Liriodendron; L. rhododendri
on Rho-
dodendron; L. iwamotoi Miy. on
rice;

L. taxicola R. K. L.
is
259
on Taxiis canadensis;
Its conidial
vagabunda
Sacc. spots linden branches.

tihse.-^^
form
perhaps
Phoma
Ophiobolus Riess
Perithecia scattered, subglobose,
fusiform, hyaline or yellowish.
(p.
252)
submembranous, covered or
asci cylindric; spores
suberumpent, ostiole papillate or elongate;
X400
Fig. 190.
Ophioasms;
After
X400
bolus. B, C, spore.
Lin(l;ui
ter.
and Win-
Fig. 191. Pleospoia from passion-fruit. The spores are just beginning to germinate, the end cells starting first. After Cobb.
X60
genus of some one hundred twenty-five species. O. graminis Sacc. and O. herpotrichus Sacc. occur on grasses and are quite injurious in Europe.^^^ O. oryzeae Miy. is found on rice.^''^
Pleospora Rabenhorst
Perithecia covered at
(p.
252)
membranous,
first, later more or less erumpent, usually black, globose; asci oblong to clavate; spores elon-
gate or ovate, muriform; paraphyses present.
260

as
Macrosporium, Alternaria, Cladosporium, Sporidesmium, Phoma, Helminthosporium. There are over two hundred twenty-five species, mostly saprophytic. Many conidial forms whose connection to this genus have not yet been definitely proved probably belong to it and are
in
Conidia occur
many
instances parasites.
^"^"^
P. tropeoli Hals, is reported as the cause of disease of the cultivated Nasturtium.

Perithecia pyriform, 140-160 /x; asci oval, one-sided, spores hyaline or very light-olivaceous, 25-35 x 6-8 fx. The Alternaria-form was grown from the ascospores by Halsted
and from the Alternaria
spores, grown in pure culture, perithecia were obtained in about twelve days. P. albicans Fcl. occurs on chicory as Phoma albicans; P. hyacinthi Sor. on hyacinths with its conidia as Cladosporium
fasciculare; P.
hesperidearum Cotton,
in its conidial form, Spori-
desmium pyriforme, causes a black mold on oranges. P. herbarum (Pers.) Rab. (conidia= Macrosporium commune)
is
common
saprophyte which sometimes becomes parasitic.
P. pisi (Sow.) Fcl.^'^ is found on the garden pea; Perithecia and spores as in P. herbarum but spores more narrow. P. ulmi. Fr. causes an elm leaf spot. P. infectoria Fcl. a com-
mon
saprophyte, parasitizes tobacco. P. oryzae Miy. is on rice; P. negundinis Oud. is injurious to nursery stock of
^'*
P. putrefaciens (Fcl.) Fr. (conidia= Sporidesmium)
Negundo; is on carrots.
Pleosporae on grains.^^^' Several species of Pleospora with their attendant conidial forms
Helminthosporium and Alternaria are known on various grains and grasses. Cross inoculation experiments have shown here
of
biologic
specialization similar to that encountered among the Erysiphese, in that conidia or ascospores from one host usually
give negative results on host species other than that on which they ^''^ grew. Thus Diedicke says the Pleospora of Bromus cannot
be grown on Triticum repens nor on cultivated barley or oats. Helminthosporium was formerly thought to be the conidial stage of all of these grain Pleosporas, but recent work of Diedicke shows
that one form which he regards as P. trichostoma (Fr.) Wint.

possesses an Alternaria conidial form.
261
Following Diedicke, the
forms given below would be recognized.

P. bromi Died.
Perithecia brown, hairy; asci 189-288 x 34-59 n, saccate, thinwalled; spores 2-seriate, golden-brown, 4-celled, 48-83 x 19-33 /x.
150 X 13-20
P.
Conidia (=Helminthosporium bromi) on brownish spots, 108On Bromus. n, 5 to 7-celled, dark colored.
gramineum Died.
Conidia (=Helminthosporium gramineum); conidiophores short,

subflexuose, light-brown; conidia solitary, elongate-cylindric, 4 to 7-celled, 15-19 p, wide and of variable length.
The mycelium invades the tissue causing long brown spots. These later become covered with an abundance of conidiophores which emerge through the stomata. Potter also reports in22 vasion and complete occupation of ovaries by the mycelium.
Sclerotia-like bodies are
formed on leaves and stems. They were first

seen
in
artificial
cul-
tures of the fungus
by
Ravn
since
^^^
and have been

in
found
244).
nature
(Noack
The conidiospores
have been shown to be long-lived, and spring
infection begins largely Fig. 192. p. trichostoma. 1, group of iisci 2, a single spore at the apex of an ascus. After carried from conidia
,
Diedicke.
winter on seed. Extensive study was made of the conidial form by Ravn who found the mycelium to be of two kinds, one aerial and hyaline, the other strict and dark. It grew well on acid or neutral media.
over
Careful infection experiments (Ravn) proved the pathogenicity

of
H. graminum
for barley
but showed
it
incapable of infecting
oats, rye or wheat.
regards the disease produced by H. gramineum as often not local, in that the mycelium may invade the growing general,
points, resulting in infection of all the leaves.
Ravn
262
P. tritici-repentis Died, is found on Triticum repens (=Agropyron repens.) Conidia=Helminthosporium tritici repentis. P. trichostoma (Fr.) Wint. ( = Pyrenophora trichostoma (Fr.)
Sacc.2^2
Perithecia gregarious, innate, conical, black, ostiole surrounded by black hairs, which are simple, septate, 6-8 ix in circumference
;
asci clavate
300 x 40
/x;
spores broadly oblong, obtuse, unequally
4 to 6-septate, muriform, brownish, 52 x 20 m; paraphyses branched. On rye with the conidial form =Alternaria trichostoma Died.
In the present state of our knowledge little is to be gained by recognition of these purely "biologic species," and all the forms may be grouped under the name P. trichostoma, recognizing the
fact that
it
shows biologic
differentiation.
Two
hypothetical forms P. teres Died, and P. avense Died,
pertain to Helminthosporiums of corresponding names.
Massariaceae
(p.
223)
Stroma none; perithecia separate, sunken, not erumpent, opening by a small pore, leathery or carbonous, compact; spores usually surrounded by a jelly-like substance; paraphyses present. This family of ten genera and about one hundred twenty-five
species contains only one parasite of interest.
Key to Genera
Spores
1 -celled
of Massariaceae
Spores not surrounded by a jelly-like substance Spores surrounded by a jelly-like substance
Enchnoa.
Pseudomassaria.
2.
Spores several-celled Spores not muriform Spores hyaline or yellow

Spores ellipsoid to spindle-shaped,
several-celled, hyaline Spores spindle-formed, curved, 3 to 4celled,
3.
Massarina.
Ophiomassaria.
Charrinia,
p. 263.
yellow
4.
Spores 2 to 4-celled, elongate, hyaline

5.

Spores brown
Spores 2-cellcd
Pefithocia scattered
irre}z;ularl\'.
.
.
263
G.
7.
Perithecia in circular clusters
Phorcys. Massariovalsa.
Spores more than 2-celled Spores ellipsoid to spindle-shaped,
many-celled Spores cylindric, bent, 8-cclled.

Spores muriform
8.
.
.
Massaria,
p. 263.
9.
Cladosphaeria.
10.
Pleomassaria.
of the tea plant. The Charrinia is said l\v Viala & Ravaz ^'^' to contain the ascigegenus rous form of Coniothyrium diplodiella (Speg.) Sacc.
Massaria theicola Fetch invades the ducts
Gnomoniaceae
(p.
223)
Perithecia sunken, with an elongate, cylindric, beak-like ostiole, rarely with a papillate one; leathery or membranous, rarely borne
on a stroma;
asci
mostly thickened apically and opening by a pore;

fifty species;
spores hyaline; paraphyses usually absent.
family of about one hundred
four genera con-
tain important pathogens.
Key
to Genera of Gnomoniaceae
Spores l-celled Mouth of the perithccium short Asci cylindric, 8-spored

Asci clavate, 2-8porcd Mouth of the perithccium elongate, beaklike
1.
2.
Phomatospora. Geminispora.
Mouth
of the perithccium Asci many-sporcd Asci 8-spored
straight
3.
Ditopella, p. 264.
Spores ellipsoid or fusoid
clypeus present
4.
5.
Mamiania.
Glomerella,
Cryptoderis.
p. 264.
p.
Stroma present Stroma absent

Spores elongate fusoid, or filiform Mouth of the perithccium recurved.
. .
6. 7. 8.
Gnomoniella,
273.
Camptosphaeria.
264
Spores 2 or more-celled Asci 8-spored
Spores elongate, 2 to 4-celled

Spores fusiform, curved, 2-celled Asci many-spored; spores elongate,
2-celled
9.
Gnomonia,
p. 274.
10.
Hendersonia.
Perithecia beaked Perithecia not beaked
11.
Rehmiella.
Rehmiellopsis,
p. 276.
12.
Ditopella de Notaris
(p.
263)
Perithecia corticolous, covered, globose or

ostiole
somewhat depressed,
oblong
suberumpent;
asci subclavate, polysporous; spores
or fusoid, continuous, subhjâline; paraphyses none. D. ditopa (Fr.) Schr. causes death of oak twigs in Europe; D. fusarispora d. Not., occurs on alder in Europe.
Glomerella Spaulding
& von
Schrenk
2^2. 342
(p
263)
Perithecia cespitose, membranous, dark brown, rostrate, of a lighter color at the apex in early stages, flask-shaped, hairy, on
or immersed in a stroma; asci sessile, clavate; spores 8, hyaline, oblong, 1-celled, slightly curved, elliptic; paraphyses usually none.
Conidia=in part Colletotrichum and Gloeosporium. This genus was first described by Stoneman, from perithecia obtained from cultures of the conidia,^'*'^ as Gnomoniopsis. On account of preoccupation it was renamed Glomerella by Spaulding and von Schrenk ^^^ in 1903. Studies by Shear have shown that there is much variation in pure line cultures both from ascospores and from conidiospores.'^^ This leads to great uncertainty as to specific
limitations as will
become apparent
in the paragraphs below.
The The
conidial forms are very common and are usually parasitic. Sometimes they ascigerous stages are comparatively rare.
are found in nature; again only in artificial culture. Some forms known to be ascigerous may in one culture yield abundant perithecia while other cultures of the
refuse to bear asci at
all.
same fungus may
persistently
G. rufomaculans (Berk.) S. & S.2^259 Perithecia on decaying fruits, subspherical,
more
or less grouped;

asci
265
12-
subclavate, fugaceous, 55-70

(
ju;
ascospores allantoic!,
22 X 3-5 n; conidial stage

small
= Gloeosporium
rufomaculans) with
sori, developing in more or less concentric circles, usually soon rupturing and pushing out spores in small pinkish masses;
spores
hyaline
to
greenish,
chiefly oblong, unicellular 10-
28 X 3.5-7
fjL.
The
Rev.
as a
conidial
first
fungus was
stage of this described by
M.
J.
Berkeley in 1854
It was later Septoria. transferred to the form genus
Gloeosporium
under
which
Fig.
name
to
it
the literature pertaining is largely to be lOUnd.
193. 7. pcrithecium
of G.
rufomacu-
lans showing asci in situ; 6, asci showAfter Spaulding and von ing detail.
See Southworth.2^0
The
as-
^'^'"^-
-'"^ in 1902 and the fungus cigerous stage was found by Clinton described as a Gnomoniopsis. In 1903, it was given the present
A bibliography of some one hundred eighty titles is given by Spaulding and von Schrenk.'''^ The conidia germinating on apples send germ tubes through the skin, usually through wounds, occasionally through a sound
name.
surface.-'^
The
mycelium
rapidly,
grows
subepidermally,
branching
and intracellularly, absorbing the sugar and other nutrients present, and resulting in brown discoloration of cells and dissolution of their connecintercellularly
tion with neighboring

is
cells.
The mycelium
Fig.
brg'eTm?ating'spores'^ After Spaulding and von Schrenk.
hyaline but later, especially in the 194. g. cactorum. stromata, it may be quite dark. Acervuli ôou appear, oftcu in conccntric rings, liftfirst
ing the epidermis with their palisades of

.
Ihe latter, at first hyaline, conidiophores. later olivaceous, bear the numerous conidia, which are pinkish, rarely cream-colored, in mass. In germination the conidia be-
,.
rr-M
i-
come uniseptate and
often on the tips of the
young mycelium
develop the dark thick-walled irregularly shaped spore-like structures, so common on the sporelings of the Melanconiales. These
266

by Hasselbring
^^^
structures are regarded

well.
as organs of attachment
to aid in infection, though they doubtless serve other purposes as

first obtained by Clinton who abundance on artificial media from sowings of coniThe typical Gloeosporium diospores taken from pure cultures. stage was also grown from ascospores. Perithecia were also found in pure cultures on apple agar by Spaulding and von Schrenk. They appeared in black knotted masses of mycelium which were often 4-5 mm. in diameter, the perithecia varying from one to many in each such stroma. The asci were evanescent, disappearing soon after the spores matured. That this fungus is the cause of a limb canker was suggested by Simpson's discovery of the canker in July, 1902 and was definitely
Perithecia of this species were

in
^^^
grew them
proved by Spaulding and von Schrenck,-^*
and by
Blair
year.
^^^
Burrill
in the
and same
In
tion
canker
the
in
formalive
mycelium
the
grows
bark, killing it the cambium.
and
The
cankers
Fig.
195.
are
thought
perhaps
the
G.
rufonmculans,
Note septa and appressoria. von Schrenk.
germinating conidia. After Spaulding and
to
be comparatively
lived,
short
surviving
third
only
between fruit and twigs year. Reciprocal have proved the fungus in the two cases to be identical. Conidia and ascospores develop on both fruit and twigs. The fungus has been repeatedly grown in pure culture on numerous media by many investigators and many inoculations with
inoculations
of the fungus to the apple rot
conidia into both fruit and twigs have proved the causal relation and twig canker. Inoculations from
ascosporic material have given the same results. That the spores may be insect-borne was shown
that they
may
also travel
by Clinton; -^^ on the wind was shown by Burrill.
-''^
THE FUNGI WPIICH CAUSE PLANT DISEASE

The mycelium
fruit.25^
2G7
hibernates in
hmb
cankers and in mummified
It
is
impossible morphologically to distinguish the conidial
stages of manj^ species of Gloeosporium and Colletotrichum growing on a great variety of hosts, and much inoculation work has
been done to ascertain the relationships existing between these ^^ in Dr. Halsted's laboratory forms. Thus the author made inoculations as indicated in Fig. 367. Southworth cross inoculated a Gloeosporium from grape to apple and from apple to Stoneman from grape; -^~ Even such quince to apple.
many
cultures give
difference
little
evidence of
between these forms
and
it usually is impossible to distinguish between the conidial forms on either morphological
or biological grounds.
Some group under Glomeralla rufomaculans as its conidial

forms,
what
G.
were
formerly
known
genum,
as Gloeosporium fructi-
rufomaculans,
Iseticolor.
G.
versicolor
and G.
Further studies of the ascigerous stages have led to consolidation rather than to segregation of
ascigerous
ella,
species.
Thus an
Glomer-
Fig. 196.
Plate culture of G. rufoniacushowing

perithecia-bearing After Spaulding and von
lans
stage,
masses. Schrenk.
culture
was obtained in pure from the following

258
conidial
forms
by
Shear
and
Wood:
G. rufomaculans from grape, G. fructigenum from apple, G. sps. from cranberry, G. elasticae from Ficus (see p. 544) a Gloeosporium from Gleditschia, one from Ginkgo, Colletotrichum gossypii from cotton (see p. 271) and C. lindemuthianum. (See
547) from bean. These authors after careful study of these perithecia and cultures conclude that: "in the present state of
p.
268
our knowledge, it may be best to regard the various forms we have studied as varieties of one species." Among the hosts of G. rufomaculans may probably be num-
bered at least apple, grape, pear, quince, peach, tomato, egg^^^ and cherry. -^^ plant, pepper, sweet pea
G. rufomaculans var. cyclaminis P.
&
C.-''^
Perithecia densely gregarious, indefinite, light-colored, around

spots,
brown, membranous, subglobose or distinctly rostrate, ostiolate

;
asci clavate-
apex pointed, 5065 X 8-9 /*; spores oblong to elliptic, 16-18 x 4-4.5 /x.
cylindric,
Con
chum)
;
dia
= Collet otriamphilarge;
linear,
acervuli
genous, conidia
brownish,
oblong
to
obovate, straight, or slightly curved, ends round, 12-15 x
4-5 n; conidiophores long,

slender;
rigid.
setae
free,
short,
This variety
is
reported
on greenhouse Cyclamens,
causing leaf spotting. Mature perithecia were found
Fig. 197. G. rufomaculans. Pustules apple, enlarged. After Spaulding and
on von
Schrenk.
Cultures on the leaves. from the ascospores gave a
Colletotrichum as
nidi al
in
the
co-
form and a similar Colletotrichum collected from the leaves
pure culture gave the Glomerella. G. cingulata (Atk.) S. & S.
Perithecia cespitose, stromate, dark-brown, flask-shaped, memshortly rostrate, more or less hairy; branous, 250-320 X 150 asci clavate, 64-16 /*; spores hyaline, elliptic, slightly curved,
fj.,
20-28 X 5-7 fx. Conidia (=Gloeosporium
cingulatum); acervuli 100-150 rupturing the epidermis, in age black; conidiophores numerous.
ijl,

crowded, simple, hyaline; conidia oblong to curved, basally pointed, 10-20 x 5-7 n.
elliptic,
209
straight or
This was first described in conidial form as a GlcEosporium by Atkinson "^ on privet as cause of cankers. The fungus was isolated and gro^\^l in pure culture. Later perithecia were obtained in
the pure
cultures.^'*^
& E.) S. & S. Perithecia cespitose, thinly membranous, dark-brown, pyriform, hairy; asci clavate; spores slightly curved, elUptic, 12-18 x 4-6 /i.
G. piperata (E.
Fig.
a,
e,
198. Diagrammatic section through acervulus of G. rufomaculans. parenchyma, h, cuticle, c, subhymenial fungous layer, d, conidiophores, After Clinton. spores, 6, conidiophores and conidia in detail.
Conidia (=Gloeosporium piperatum) on circular or oval spots;

acervuli pustular, concentrically arranged, conidia 12-23 x 5-6 ^t.^^^ The ascigerous stage was grown from pure cultures of the conidia
the perithecia appearing taken from pepper by Miss Stoneman about a month after inoculation. Typical conidia were also se-
^'*"
cured from ascospore sowings. G. cincta. (B. & C.) S. & S.-^
Perithecia
180-280
/z,
flask-shaped,
membranous,
yn;
cespitose;
asci clavate, truncate or obtuse,
65-70
spores
elliptic,
curved,
5-20 X 3
M-''^
Acervuh erumpent; conidia (=Colletotrichum cinctum) 12-15 x
270 3-4

n,
elliptic,
guttulate; sete present, but almost obscured

'^'^
by
the spore mass.
from ascigerous stage was demonstrated by Stoneman pure culture studies. The conidial stage was described by Halsted ~^^ as the cause of a blighting of orchid leaves (Sobralia) in
The
New
Jersey.
Various hosts are orchids, Sarracenia, rubber plant, Dracaena "^ and Anthurium.-^-
Fig.
199.
G. rufoniaculans, acervulus showing conidia, conidiophores and setae. After Hasselbring.

S.
G. rubicola (Ston.)
&
S.
Perithecia quite similar to those of G. piperata and G. cinta but lacking the apical tuft of hair and rather larger in size.
Conidia
(=Colletotrichum
brown patches on the upper surface
rubicolum) forming large, darkof the leaf; sori small, dark,

^'^^
suberumpent; conidia oblong, elliptic, 12.5 x 6 /i. The conidial form on red raspberry was shown by Stoneman by pure culture studies to possess this ascigerous stage. G. psidii (Del.) Shel.-*''''-26^
Perithecia 200-300
n,
spherical, rarely distinctly beaked; asci

cylindric to broadly clavate, blunt, 45-55 curved, continuous, granular, 13-15 x 5-6 /z.
271
spores
defi-
9-10
^l^,
Conidia
= (!loeosporium
fj.;
psidii), acervuli
subepidermal on
nite spots, 90-120
4-5
ju;
conidia
elliptic,
conidiophores hyaline, cylindric, 15-18 x oval, hyaline, 10-13 x 4-6 /i.
Artificial culture studies
ascigerous stage.
-^^ demonstrated the by Sheldon -^^' was made of the growth on Extensive study
Fig. 200.
G. piperata,
tion.
99, perithecium external and in sec100, asci in detail. After Stoneman.
Two distinct forms of conidia apple-agar, apples, plums, etc. were observed, one on loose hypha?, the other in acervuli. The
species should probably be regarded as a variety of G.
rufo-
maculans.
on the guave. G. gossypii (South.) Edg. Perithecia distinct or crowded, very abundant, covered, dark brown to black, subglobose to pyriform, 80-120 x 100-160 /i, beak up to 60 n long; asci numerous, clavate, 55-70 x 10-14 n;
It occurs
272
spores elliptic, hyaline, rarely curved, 12-20 x 5-8 n; paraphyses
long and slender, very abundant. Conidia ( = Colletotrichum gossypii), acervuli erumpent, conidiophores colorless, longer than the spores, 12-28 x 5 fx; conidia
irregularly oblong, hyaline or flesh-colored in mass; setse single or tufted, dark at base,
colorless
The
scribed
above, straight, rarely branched. conidial stage of this fungus was de^"^
and independby Southworth ^^^' -^^' ^^^' ^'-^ on cotton. ently by Atkinson The ascigerous stage was first seen by Shear & Wood ^^^ in artificial culture and by them regarded as probably a variety of G.
rufomaculans. Since these studies Edgerton -^ from examination of perithecia developed naturally in the open, has proposed
it
as a separate species.
The mycelium
Fig. 201. G. gossypii. Section of young boll,
is
richly
branched and sep-
showing
the fungus penetrating the hull
but sometimes slightly smoky. It grows between and in the host cells which are often filled with it, causing
tate, usually hyaline
j and brownmg. ^^^ show Studies by Atkinson and by Barre ^^^t in case of diseased bolls the mycelium
collapse, loss of chlorophyll,
in
ySung'^sled.'^Spores are being produced
upon the outer portion of the hull and upon the surface of the young seed coat,
After Barre.
may
..
.
its
mner
extend through the pericarp, sporing on .1 ,, j.i, Wall; extend thence to the seeds;
.
j.
the
cells of
the
lint.
penetrate and grow in them. Fig. 201, and in Barre has shoAAm that even the endosperm
and cotyledons may be invaded, Fig. 201, and spores produced upon them while \\dthin the seed coats. Such seeds and lint may appear outwardly as though perfectly normal. The conidia are formed in acervuli, subtended by stromata. Setae, from few to many increasing with age of the acervulus, are present and conidia are occasionally found on them. In germination conidia usually develop one, sometimes two septa and produce dark chlamydospores. Acervuli are common on bolls, ' less so and smaller on leaves and stems.
The
perithecia as found in the field
by Edgerton
in Louisiana
273
were usually entirely embedded, with the beaks only protruding and were often numerous and crowded. Cultural evidence that Edgerton's specimens were actually genetically connected with
the cotton anthracnose are wanting. The fungus has been repeatedly studied in pure culture and numerous inoculations have thoroughly proved its pathogenicity,
the disease usually showing within a few days after inoculation, though
sometimes
incubation
is
delayed
much
longer.
is
Infection of stems
often at a
wound such
as a leaf scar; or on leaves at some point of weakness.
Cotyledons and young plants are

especially
susceptible.
is
On
at
bolls
Fig.
infection
common
the line
202. G.
gossypii,
'"^
^l^^Baire cording to Barre, there is evidence that the fungus may destroy the contents of the boll before it shows upon the outside. Barre showed that 44% of flowers
of dehiscence of the carpels.
Ac-
D, and E, fun''" "* ^^^'^'
that received spores within ten hours after opening produced diseased bolls; but inoculations by spraying produced no results on
they were three-fourths grown. Seed from a field that bore 35% infected bolls gave on germination, 12% of infected seedlings, the disease appearing upon cotyledons or hypocotyls even before they unfolded. Atkinson ^^ found that conidia five months old were alive, but that at seven months they failed to germinate. Barre also found the conidia and the mycelium of the fungus to be comparatively short lived. G. atrocarpi Del. on Atrocarpus leaves has been described as
bolls after
a perfect stage of Gloeosporium atrocarpi Del. A fungus on Cattleya ^^' ^^^ described by Maublanc
as a Physalospora should perhaps be considered as
&
Lasnier
a Glomerella.
Gnomoniella Saccardo
(p.
263)
dric,
Perithecia sunken and usually remaining so, with a long cylinerumpent ostiole, leathery, black; asci ellipsoid or fusoid,
274

apically thickened
and opening by a pore;

hyaline; paraphyses
spore none.
elliptic,
1-celled,
Fig. 203. G. tubiforAfter mis, perithecia.
This genus of some twenty-five species contains G. tubiformis (Tode) Sacc. which is said to be the ascigerous stage of Lepto-
Winter.
thyrium alneum Sacc. growing on Alder.
Two
other species, G. fimbriata and G. coryli are found on hornbeam and hazel respectively.
Gnomonia
ostiole
Cesati
&
de Notaris
(p.
264)
Perithecia covered, or erumpent, submembranous, glabrous, more or less elongate; asci ellipsoid or fusoid, apically thick-
ened, opening by a pore; spores elongate, hyaline, 2 to 4-celled; paraphyses none.
There are some sixty species. Fusicoccum, Myxosporium, Sporonema, Gloeosporium, Marssonia, Asteroma, Leptothyrium occur in some species as the conidial form. The ascigerous form usually
follows as a saprophyte after the parasitic conidial stage.
G. veneta (Sacc.
Perithecia
&
Speg.) Kleb.^^^^
s'^^,
323, 335
immersed, subglobose or slightly flattened,
150-
200
short, rostrate; asci long-clavate, 48-60 x 12-15 /x, generally bent at right angles at the base, apically very thick, opening
fi,
by a pore; spores 14-19 x

the upper
cell longer.
4-5, straight
or slightly curved, unequally 2-celled,
Conidia
habit.
(1)
variable
(
in habitat, and = Gloeosporium nervise-
quum)
300 />;
acervuli
subcuticular
100-
conidiophores short, conidia oozing out in a creamy-white mass, hyaline, ellipsoid, 10-14 x 4-6 /x,
pointed at one end and rounded at the other. (2) ( =G. platani) acervuli sub-
Fig.
204.
G.
epidermal, conidiophores long; conidia as above. (3) ( =Discula platani = Myxosporium valsoideum) forming minute, subepidermal, erumpent pustules on twigs; conidia
elliptic to
cium.
veneta, peritheAfter Edgerton.
oblong, hyaline, 8-14 x 4-6
/x;
(4)
=Sporonema
platani
275
= Fuscicoccum veronense). Pycnidia formed on old leaves on the ground, crumpent, subcuticular, brown, 200-300 ju; conidia numerous, oblong, ovoid to f usoid, 7-1 1 x 3-4 /x.
form on sycamore and oak, first described in 1848, and young branches, the mycelium checking the sap-flow and causing death of surrounding tissue. A stroma is formed on the outer layers of the mesophyll and from this arise
conidial
is
The
common on
leaves
the
short
conidiophores
to constitute
the
acervulus.
-^^ gave experiments by Tavel Other infection experiments negative have also been unsatisfactory.
Infection
results.
The
Klebahn
ascigerous form was first found by ^'^ on old leaves on which it ma-
'^^^J spores.^ After'
Ed-
tured about Christmas time.
While the co-
gerton.
nidia are uniform in shape four as stated above.
modes
of
development are found,
Pure cultures from all the spore forms were compared by Edger^~confirming Klebahn's conclusion as to their identity. Cul-^^ tures by Stoneman showed the forms on sycamore and oak to be the same. G. leptostyla (Fr.) Ces. & d. Not. Perithecia conic, short-beaked; asci subclavate, 45-65 x 10-12
ton
ju;
spores
f usoid,
(=Marssonia
rounded;
curved, 18-22 x 4 Acervuli juglandis).
/i,
hyaline.
Conidial phase
gregarious,
n,
hypophyllous,
pointed above, truncate below, greenish. The connection between the conidial and ascigerous forms was
obovoid,
1-septate,
conidia
8-10 x 4-5
demonstrated by Klebahn ^^^ by pure cultures and by ascosporic infection. The conidial form is common on walnut leaves; especially severe on the butter-nut (Juglans cinerea) often defoliating this host in mid-summer. G. quercus-ilicis Berl. occurs on oak leaves in Italy. G. erythrostoma Auer. is the cause of a disease of cherry leaves
in
Europe; G. padicola Kleb. is the ascigerous stage of Asteroma padi which is widely distributed in Europe on Prunus.
-'*''
'^''^
276
G. oryzae Miy. occurs on rice.'^'^ G. rubi Rehm may occasionally cause disease of blackberry
canes.-'*
Rehmiellopsis Bubak
& Kabat
(p.
264)
Similar to Rehmiella except that the perithecia are not beaked and the pycnidia do not have a definite opening.
R. bohemica Bub.
curs as a parasite on
&
fir
Kab.; (conidia=Phoma bohemica)^*^ ocneedles.
Clypeosphaeriaceae
(p.
223)
Perithecia immersed, astromatic or with a pseudostroma built of hyphse which, with the adjacent substratum, forms a thin cly-
peus that
beaked, paraphyses usually present. A small family chiefly saprophytes.
usually evident only above; ostiole short to longerumpent, walls mostly carbonous to membranous;
is
Key to Genera
Spores 1-celled
of Clypeosphaeriaceae
Perithecia soft-membranous, spores hyaline or brown

Perithecia leathery; spores Spores more than one-celled
1.
Trabutia.
brown
2.
Anthostomella,
p. 276.
Spores with cross walls only Spores cylindric, ellipsoid or fusiform Spores hyaline,
1
to 3-septate
3.
Hypospila.
Clypeosphaeria.
Phaeopeltosphaeria.
Spores brown Spores elongate Spores

fusiform,
4.
more
than
45.
septate, sometimes muriform.
Spores filiform, hyaline to yellow Spores muriform Spores ovate, brown

Spores short, fusiform, hyaline
6.
Linospora.
Peltosphaeria. Isothea.
7.
8.
Anthostomella Saccardo
Mycelium fusing with the upper

form a
thin, black,
surface of the substratum to rounded pseudostroma; perithecia sunken, sub-

globose,
277
with
short,
conical
ostiolc,
walls
black, carbonous to leathery; asci cylindric, 8-spored; spores elliptic, continuous, brown,
unappendaged paraphyses usually present. Over one hundred species, chiefly sapro;
phytes. A. sullae
leaf spot
Montem. occurs
sulla.-^^
as the cause of a
Fig.
on
206. A.
des-
A. bohiensis (Hmp.) Speg. is on cacao; A. destruens Sh. on cranberry;
truens.
8, pcrithe-
spores;
A.
coflfeae
Desm. on
coffee. ^^^'
'^^
cium; 9, ascus; 10, 11, germinating spore. After Shear.
Valsaceae
(p.
223)
Stroma effused, subglobose, conic, or pulvinate, often indefinite; perithecia sunken in the stroma, scattered or clustered, black, leathery; asci cylindric or clavate; paraphyses usually present.
Over one thousand
species, chiefly saprophytic.
Conidia are
present on hyphse or in pycnidia.
Key to Genera op
Spores 1-celIed
Valsaceae
Spores cylindric or ellipsoid, with a brown
membrane
Spores ellipsoid, curved or not, with a
hyaline membrane Spores more than I-celled Spores with cross walls only
I.
Anthostoma.
Valsa, p. 278.
2.
Spores hyaline
Spores unappendaged
Spores ellipsoid or fusoid 2 to 4celled
3.
Diaporthe,
Vialaea.
p. 278.
Spores elongate, fusoid, constricted in the middle
4.
Spores appendaged, I appendage at each end and 2 or 3 in the middle

5.
Caudospora.
Spores brown Spores 2-cellcd, ellipsoid

Spores many-celled, fusoid
6. 7.
Rhynchostoma.
Kalmusia.
278
Spores muriform
Stroma effused
Spores hyaline Spores colored
8.
9.
Thyridella.
Thyridium.
Fenestella.
Stroma none or pulvinate
10.
Valsa Fries
Perithecia on a
ostiole
(p.
277)
erumpent,
or less definite stroma, immersed, the black, firm; asci globose to cylindric, often
more
long-pedunculate; spores
1 -celled,
rarely 2-celled,
cylindric, rounded,
line or light-brown;
hya-
paraphyses none. V. leucostoma (Pers.)

tp
22, 229, 280
Fiu. 207.
Valsa.
C. asci.
A, habit sketch; B, perithecia; After Tulasne.
vex,
Stroma strongly con2-3 mm., whitish
and granular within, outer layer coriaceous; perithecia immersed; asci fusoid-clavate, subsessile, 35-45 x 7-8 /x; spores biseriate, allantoid, hyaline, slightly curved, 9-12 x 2-2.5 n.
rubescens); stromate, erumpent, reddish; On pome and stone fruits throughout conidia allantoid, 4 fi. Australia and America causing the disease known as Europe, " ^^^ who worked dieback." The fungus was studied by Rolfs -"^'
(
Conidia
= Cytospora
out
its life cycle.
V. oxystoma Rehm. occurs on Alnus in Europe; V. (Eutypa) caulivora Rehm. affects Hevea.
V. ambiens Fr. is on the apple in Europe. V. (Eutypella) prunastri (Pers.) Fr. is the cause of serious diseases of apples, plums, etc., in England.
V. (Eutypa)
in the tropics
erumpens Mas. is reported as a wound parasite on Ficus, and cacao.

Diaporthe Nitschke
(p.
277)
Stroma very
variable, usually definite; perithecia
membranous
subcoriaceous, generally pale-cinereous within, with a cylindric or filiform beak; asci fusoid; spores fusoid to subelliptic, 2-celled,

hyaline,
279
raphyses
appendaged or not; panone. Conidia=Phoma,

etc.
Cytospora,
D. taleola (Fr.) Sacc. Stroma cortical, definite, depressed, pulvinate, 2-4 mm., covered; perithecia few, 4-10, buried, their ostioles converging, erumpent in a small light-colored disk; asci
cylindric,
elliptic,
n, spores uniseptate, constricted, with
120-140 x 10-12
appendages,
setaceous
15-22
8-9
It
(ji.
the
cortex
causes canker on oak, killing over large areas. A
year later the cushion-like stromata

appear.
The mycelium
penetrates
o
Fig.
both wood and bark, probably enterin section; C, lasne. ing through wounds. D. albocarnis E. & E. on Cornus is destructive.
208. Diaporthe.
asci.
B, stroma, After Tu-
D. ambigua and D. sarmentella are on pear and hop, D. strumella on a wide range of hosts, in conidial form as Phoma.
Melanconidaceae
(p.
223)
small family of less than two hundred species contains only

in the
four parasitic genera.
Stroma pulvinate, sunken; perithecia sunken

the
stroma,
mouth erumpent;
asci cylindric or clavate;
paraphyses present.
Key to the Genera

Spores
1 -celled, hyaline Spores ellipsoid or short-fusiform Spores elongate-cylindric, curved
of Melanconidaceae
I.
Cryptosporella, p. 280.
2.
Cryptospora.
Spores 2-celled
Spores hyaline Conidia in pyenidia; 1-cellcd, hyaline. Conidia not in pyenidia, dark brown.
Spores brown
3.
4.
5.
Valsaria.
Melanconis, p. 281. Melanconiella.
280
Spores more than 2-celled

Spores hyaline
Spores elongate, multicellular
6. 7.
Calospora,
Holstiella.
p. 280.
Spores fusiform, multicellular Spores brown
Spores elongate, 8 or 4-spored
multicellular;
asci
8.
Pseudovalsa,
Titania.
p. 281.
Spores long-cylindric, very large, asci

l-spored
9.
Calospora Saccardo
One
species,
is
trouble,^^
C. vanillae Mas., reported as causing a Vanilla perhaps identical with Gloeosporium vanillae C. & M.
Cryptosporella Tulasne
(p.
279)
Stroma valsoid, pustuliform, covered; perithecia embedded, subcircinate, with converging necks united in an erumpent disk; asci cylindric to globoid; spores
elongate, cyHndric, hyaline, 1-celled. C. anomala (Pk.) Sacc^^^- -^
Pustules
prominent,
2-5
mm.,
penetrating the wood and generally having a thin black
erumpent;
crust beneath them, disk convex or

slightly depressed, cinereous to black
;
crowded, deeply embedded in the stroma, often elonperithecia

gate, ostioles scattered, black; asci
Fig. 209. C. anomala. 31, stroma and perithecia; 32, an ascus; 33, After Humphrey. spores.
short, broad, fugaceous; spores

line, elliptic,
hya-
Common
simple, 7-8 /i. on hazel and filbert in
America, causing the destruction of the tops while the roots re-
main
alive.
C. viticola Sh.324
Pycnidia (=Fusicoccum) with labyrinthiform chambers, ostiolate but frequently rupturing. Spores hyaline, continuous, of two forms 2. Long, slender, in the same cavity. 1. Subfusoid, 7.5 x 2-5 fi.
curved, 18-30 x 1-1.5
/x.
Perithecia buried in irregular pulvinate

stromata, beak exserted; asci 60-72 x 7-8
septate,
/x;
281
wavy; ascospores
Fig. 210.
subelliptic, hyaline, continuous,
paraphyses slender, 11-15 x

as the cause of
4-6
tx.
The
conidial stage
was described by Reddick

^^^
necrosis of grape vines
though he has since stated that the amount of damage due to this disease is not so great as at first
thought.
^^^ ascigerous form in pure culture in the hands of Shear gave rise to the typical conidial form, identical with that grown from pure cultures of the pycnospores.
The
Melanconis Tulasne
(p.
279)
Stroma valsoid, seated in the substratum, partially orumpent; perithecia clavate, immersed, with long cylindric beak; asci cjdindric, long-clavate, 8-spored; spores ellipsoid to elongate, hyaline.
About twenty species; chiefly saprophytes. M. modonia Tul. in its conidial form (=Fusicoccum
pernicio^^^
sum) causes a serious disease of the chestnut in Europe.^^^' Pseudovalsa longipes (Tul.) Sacc. is parasitic on oak.
Diatrypaceae
(p.
223)
or pulvinate, built of thick hyphae, under the at length prumpent, bearing both asci and conidia or peridium, present only with the conidia; perithecia sunken in the stroma or
Stroma effused
superficial, ostiolate; asci usually
thickened apically; 4 to 8 or
many-spored; spores usually continuous, small, cylindric, curved.
About one hundred seventy-five species. One parasitic genus occurs on cherry and plum.
Key to Tribes and Genera of

Stroma absent from ascosporic stage
Asci 8 (rarely 4)-spored
Diatrypaceae
Calosphaerieae.
I.
Spores 1-celled Spores 2-celled

Asci many-spored
1.
Calosphaeria, p. 282.
2.
3.
Cacosphaeria.
Coronophora.
Diatrypeae.
Stroma present
in the ascosporic stage
II.
282
CalosphaBria.
Tulasne
(p
281)
clustered, ostiole
Perithecia astromate, free or on the inner bark, scattered or more or less elongate; asci clavate, fasciculate;
spores small, cylindric, curved, hyaline, continuous; paraphyses longer than the asci, stout lanceolate, evanescent.
About
thirty-five species chiefly saprophytes. C. princeps Tul. Perithecia on the inner bark in orbicular or elliptic groups, gen-
erally densely crowded, globose,
smooth and shining, necks
long,
Fig.
210. Cryptoviticola.
sporella
Asci
and
raphyses. Shear.
paAfter
Fig. 211.
Calosphajria
B,
perithecia; lasne.
princeps. A, group of conidial stroma. After Tu-
decumbent, flexuose, spores 5-6 x 1-5 yu.
cylindric,
erumpent;
asci
12-26 x 4
)u,
On
plum, cherry, peach and even pomaceous
trees.
Melogrammataceae
(p.
223)
Stroma usually pulvinate, rarely effused, hemispheric, subperidial then erumpent and more or less superficial; perithecia sunken in the stroma; conidia occur in acervuli on the surface of
the young stromata, or in pycnidia. A small family of about one hundred twenty-five species, only one genus of which contains important pathogens.
283
Key
Spores
1 -celled
to Genera of Melogrammataceae
Spores roundish ellipsoid, asci long fusi-
form
Spores ellipsoid or ovate, asci clavate. Spores 2 or more-celled Spores with cross walls only Spores 2-celled
...
1.
Gibelia.
2.
Botryosphaeria, p. 283.
Spores hyaline
Paraphyses present
3.
Endothia.
Myrmaeciella.
Paraphyses absent
Spores brown Spores more than 2-celled, eUipsoid to
filiform
4.
5.
Myrmaecium.
Spores hyaline many-celled

Spores hyaline 3-celled
Spores brown Spores muriform
6.
Sillia.
7. 8.
Melanops,
Berlesiella.
p. 284.
Melogramma,
p. 284.
9.
Botryosphaeria Cesati
&
at
de Notaris
first
Stroma pulvinate, black; perithecia

remaining so or becoming more or less prominent,
usually small, globose, ostiole
sunken
in the stroma,
inconspicuous, papilliform; asci clavate; spores

elliptic
to
oval,
hyaline,
continuous; paraphyses
present.
B. ribis G. &. Dug.^s^
Stromata black, more or

pulvinate, outer surface botryose, 1-4 mm. in ^^grliomQ+/^T. noiioUir O Q tvitvi Uiameter, USUany Z-6 mm.,
less
^^
212.
Botryosphferia.
more
B. stroma in sec-
and
surrounded
by the
îon; C, part of pcrithecium in section. After Tulasne.
and pycnidium
fissured periderm, regularly scattered or in
or less definite,
longitudinal rows or elongated stromata.
Perithecia
somewhat
284
top-shaped, with papillate ostioles and usually projecting, sometimes practically superficial. Few to many in a stroma and usually
interspersed
among
IX,
pycnidia; 175-250
jx
in width.
filiform
Asci clavate,
paraphyses. Spores fusoid, continuous, hyaline, 16-23 x 5-7 y.. Pycnidia of the compound stylosporic form, Dothiorella, are borne the same or similar stromata; spores fusoid, continuous, hyaline,
80-120 X
17-20
and with numerous
18-31 X 4.5-8
)U.
phoma,
are
embedded
Pycnidia of the simple stylosporic form, Macroin the outer bark under the much-raised
primary cortex of young shoots, depressed globular, 175-250 mm. wide; spores fusoid, hyaUne, continuous, 16-25 x 4.5-7.5 n. The cause of a blight of canes of currants. The fungus was first noted in sterile form by Fairchild.-^^ Its history was first fully worked out by Grossenbacher & Duggar.^^^
Extensive inoculation experiments and pure culture studies definitely established its pathogenicity.
B. dothidae Ces.
cultivated roses.
&
is
d.
Not. causes epidemics of disease among
B. gregaria Sacc.
injurious
on
^villows in Europe.^^^
(p.
Melanops Fuckel
283)
asci
Stroma lens-shaped, black; perithecia sunken;

brown.
elongate,
8-spored; spores elongate, 3-celled, hyaline; paraphyses elongate,

^^^ the conidial stage of some members of According to Shear, this genus is a Sphseropsis which is indistinguishable from S. vitic-
ola
and
S.
malorum.
henriquetii Br.
Melogramma
&
Cav.
(p.
is
parasitic
on cork oak.
Xylariaceae
224)
Stroma variable, usually free but often more or less sunken in the matrix, either upright and often branched or horizontal, effused, crustaceous, pulvinate, globose or hemispheric, black or
becoming black, usually woody or carbonous; perithecia peripheral,
immersed, leathery or carbonous, black;
asci
cylindric or
cylindric-clavate,
8-spored; spores continuous, brown or
black,
fusiform or ellipsoid, paraphyses present or absent. A family of over five hundred species.
285
Key to Genera
of Xylariaceae
Stroma encrusted, shield-form, globose or

hemispheric, without a sterile base. ... Conidial layer beneath the surface of the
I.
Hypoxyleae.
stroma, erumpent Conidial layer free from the Stroma encrusted
1.
Nummularia,
p. 285.
first
Spores 1-cellcd Spores 2-ceIled
2. 3.
Bolinia.
Camarops.
Stroma
discoid
to
hemispheric,
en-
crusted together
Young stroma
fleshy,
covered
by
4.
conidia, at length carbonous ...
Ustulina, p. 286.
Stroma carbonous or woody from the

first
Stroma without concentric layers. Stroma with concentric layers ....

Stroma
branched, clavate or cylindric, with a sterile base
of these genera are saprophytic
erect, simple or
5.
Hypoxylon.
Daldinia.
6.
II.
Xylarieae.
Most
on wood or bark.
Nummularia Tulasne
Stroma
orbicular,
cupulate or discoid, becoming black, mar-
ginate; perithecia monostichous, peripheral, immersed; asci cylindric; spores subelliptic, continuous, dark.
The genus contains

injurious.
forty species.
Only one
is
recorded as
N. discreta (Schw.) Tul. Stroma erumpent, orbicular, 2-4 mm., cupulate, with a thick
raised margin; ovate, cylindric, nearly 1 mm. long, abruptly contracted above into a short neck; asci 110-120 x 10-12 ju; spores
subglobose, nearly hyaline, then opaque,

filiform.
10-12 n; paraphyses
This fungus is usually a saprophyte but has been reported by Hasselbring as a serious parasite on the apple in Illinois. ^^ The mycelium grows more rapidly in the wood than in the bark,
286
first the parenchyma cells and medullary rays. The young stromata appear under the bark bearing when young small The stromata later turn hard and black and unicellular conidia.
attacking
N. discreta, B, stroma and perithecia, C, a Figs. 213-214. perithecium, D. asci and spores. After Hasselbring.
in
the upper layers bear numerous flask-shaped perithecia with
long necks, Figs. 213-214.
Ustulina Tulasne
(p.
285)
Stroma
at first superficial, subeffuse, rather thick, determinate,
clothed with a pulverulent cinereous conidial hymenium, finally more or less hollow; rigid, carbonous, black, bare and generally

perithecia immersed,
large,
287
papillate-ostiolate;
asci
pedicellate,
8-spored; spores ovoid-fusiform; paraphyses present. A genus of about ten species, chiefly saprophytes. U. zonata Lev. is the cause of the commonest root disease of
tea and
is
common
also
on Hevea.
BIBLIOGRAPHY OF ASCOMYCETES*
1
2 ^
Harper, R. A., Ann. Bot. U: 321, 1900. Clausen, P., Bot. Zeit. 63: 1905. Harper, R. A., Carnegie Inst. Pub., Oct. 1905. Roze, E., C. R. Acad. Sc. Paris, 125: 780.
Radais, M., Bot. Gaz. 28: 65, 1899. Peglion v., C. Bak. 7: 754, 1901. Lewis, C. E., Me. B. 178: 1910.
*
8 6 ' 8
'
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& Smith, R. E., Mass. R. 57, 1901, Larsen, L. D. H., Sug. PI. Assn. B. 10.
3M
297
3"
Aderhold, R., Landw. Jahr. 25: 875, 1896 and 29: 541, 1900. Brooks, F. J., Ann. Bot. 24: 285, 1910.
"^Prillieux, E. 1893.
and Delacroix,
G., Bui. Soc.
M.
d.
France, 9: 269,
^" Bull. Soc. My. d. Fr. 14: 24, 1898. 3" Brefeld, Unt. 9.
BAsiDioMYCETEs
This class
typically
is is
(p.
ei)!^.^:!!;^:^;":^:'?'"'
distinguished from all others by its basidium, which a sporophore bearing on its distal end short stalks,
the sterigmata, usually four,
on which are borne spores, basidiospores, one on the tip of each sterigma. Fig. In the great ma215.
jority of genera the basidia are typical and are clearly
recognizable as such. In many of the
lower
basidiomycetes the basidia deviate somewhat from the

typical form.
Thus
in the
Hemibasidii,
fungi,
the smut
the basidia are not
typical in -that they always

arise from chlamydospores, not directly from the myFig. 215.
The typical basidium with mata and spores in different stages

velopment.
After
sterigof de-
ceUum,
Figs. 217, 231,
and
De Bary.
these often
that they maj' produce more than the normal number
of
four sporidia and
from
lateral,
not terminal
sterigmata. The basidia in the large group of rust fungi are also atypical. The mycelium of the Basidiomycetes is septate and branched, and is always well developed. It is often found invading cells
several meters from the sporogenous structures and frequently weaves together to form rhizomorphs. Peculiar cell connections known as clamp connections, or knee
joints. Fig. 287, are often found.
The
basidia in
many
genera are
298
299
borne on large complex sporophores composed of the mycelial threads interwoven to form a false
The spores may parenchyma. germinate by tubes or by budding.
Typical
tirely
sexuality
seems
organs,
en-
wanting, even rudimentary

sexual
cer-
or
vestigial
tainly recognizable, have not been found. The group is supposed in

this regard, to represent the results
of
extreme simplification; the sexual organs to have long ago disfunctionally as fertilization.
Fig.
216.
Ustilago
now
development.
spores showing After Do Bary.
appeared and the simple nuclear fusions that
exist to serve
Key
to the Subclasses of Basidiomycetes
Chlamydospores at maturity free in a sorus, produced intercalary, from the mycelium basidiosporcs borne on a promycelium and simulating
;
conidia
1.
Hemibasidii,
p. 299.
Chlamydospores absent or when present borne on definite stalks Basidia septate, arising from a resting spore or borne directly on
a hymenium
Basidia nonseptate, borne on a hy2.
Protobasidii,
p.
323.
menium
3.
Eubasidii, p. 393.
Hemibasidii
The Hemibasidii contain one

Ustilaginales
Parasitic fungi,
order.
46. 47. 124. 126-129. 131. 137
smut producers, mycehum consisting
of hyaline,
somewhat
septate, branched, mostly intercellular filaments, practically limited to the interior of the host; at maturity often
300
disappearing partially or wholly through gelatinization; fertile mycelium compacting into masses and giving rise to numerous chlamcontents. Conidia rarely develop on Sori prominent, usually forming dusty or agglutinated spore-masses that break out in definite places on the host or more rarely remain permanently embedded in the tissues.
its
ydospores formed from the exterior of the host.
Spores (chlamydospores) light to dark colored, single, in pairs, or in spore-balls, the latter often composed in part of sterile cells. The Ustilaginales are all parasites on higher flowering plants. The vegetative mycelium is mostly inconspicuous and is often
Fig. 217.
Ustilago.
2,
tosis; 5, witii
4 nuclei;
pruinycelium with nucleus in miAfter Harper. 6, with conidia.
evidence of
distributed very widely in the host plant without giving external It sends its presence until time of spore formation.
variously formed botryose or spherical haustoria into the host At time of maturity of the fungus, the mycelium develops cells. in great abundance at certain special places in the host, often in
the ovary, leading to the development of large mycelial structures in the place of the host tissue.
The chlamydospores develop directly from the vegetative mycelium; new and numerous transverse cell-walls are formed; the
resulting short cells swell,
round
off
and become coated with a
gelatinous envelope.
This later disappears and the spores develop a new, thick, usually dark, double wall which is variously marked.

The chlamydo.spores may be simple
part sterile the genera below.
less
301
in in
or compound, fertile or and marked as described and are variously shaped at once or after a
The chlamydospores may germinate

protracted rest interval.
more
or
In germination in water or nutrient solution (manure water, etc.) a short tube is protruded, the promycelium, this differing in character in the two famiHes, Figs. 217,
231.
From
the promycelium of most species there develop conidia,
The promycelium is (often called sporidia) 1-12 or even more. as homologous wnth the basidium of the other basidioregarded
mycetes and the conidia as basidiospores.
The
and
conidia in suitable nutrient solutions often undergo repeated
indefinite
budding
is
Fusion of conidia
Fig. 218.
closely simulating yeast not uncommon.
cells in
appearance.
in suitable plant parts
Conidia finding lodgment under suitable

rise to
pa^^^
^JS^T^
environmental conditions give

infection.
fection
points at which incan occur are very diverse

species
The
with different
and
will
be
*^
Fig. 218.
fusing;
considered under the separate species below. r-,, ,. 1The vegetative cells are binucieate in TiUetia, multinucleate in the Usti,
Ustilago.
19,
17, ponidia promyceiiai cells
^^^^^ ^'f'Verii^Sêl!
'"
The young chlamydowere shown by Dangeard ^^^' in the case of Doassansia, spores Entyloma, Ustilago and Urocystis to be binucieate. These two nuclei, according to Dangeard, later fuse rendering the mature In germination the one nucleus passes into spore uninucleate. the promycelium, then divides mitotically Fig. 217, 2. A second
laginacea}.^'^^^
division gives four nuclei (Fig. 217, 5) the spore nuclei.'*
panying nuclear fusion,
In the fusions of smut conidia Federly has found an accomin salsify smut, while Lutman finds similar
is
fusion in the conjugating promyceiiai cells of oat smut.^ Whether or not these nuclear fusions represent a sexual act
controverted point. There are according to Clinton about four hundred species in America.^' ^"'^
much
302
Key to
Families of Ustilaginales
lateral
1.
Promycelium usually with sporidia

at septa
Ustilaginaceae, p. 302.
Tilletiaceae, p. 314.
Promycelium
sporidia
with
clustered
terminal
2.
Ustilaginaceae
forming exposed dusty or agglutinated sporeGermination of chlamydospores by means of septate promycelia which give rise to terminal and lateral sporidia or else
Sori
usually
masses.
to infection-threads.
Key to Genera of
Spores single
Sori dusty at maturity
Ustilaginaceae
definite false membrane With false membrane of definite fungous
Without
1.
Ustilago, p. 303.
cells
2.
Sphacelotheca,
p. 310.
Sori agglutinated at maturity
Firmly agglutinated into conspicuous

tubercular nodules
3.
Melanopsichium.
Cintractia.
Developed around a central columella

(rarely dusty) Spores chiefly in pairs
4.
Sori agglutinated (on leaves)
5.
Schizonella.
Sori dusty (inside peduncles) Spores in balls of more than two

Sori dusty or granular Spore-balls often evanescent;
6.
Mykosyrinx.
spores
7.
olive-brown or black-brown
Spore-balls rather permanent; spores yellowish or reddish, with markings
Sorosporium,
p. 312.
only on free surface

Spore-balls quite permanent; spores adhering by folds or thickenings of outer coat
8.
Thecaphora,
p. 313.
9.
Tolyposporium, p. 313

Sori asslutinated
(variable) Spore-balls thick-walled spores
303
composed
of
10.
Tolyposporella.
Spore-balls with peripheral spores central sterile cells
and
11. Testicularia.
important plants.
Of these genera numbers three to eleven inclusive occur on unAmong them are: Polygonum, Rynchospora,
Psilocary, Cyperus, Carex, Luzula, Juncus, Fimbrystylis, Cissis; various unimportant grasses, members of the Carduacese, Fabacese,
Eriocaulacese.
Nyctaginacese, Amarantaceae, Cyperacese, Dracenacese, and The most important genera are Ustilago and
Sphacelotheca.
Ustilago (Persoon) Roussel
Sori
(p.
302)
on various parts of the
hosts, at
maturity forming dusty
spore masses, usually dark colored; spores single, produced irregularly in the fertile mycelial threads which early entirely disappear through gelatinization, small to medium in size; germination by
means
of a septate
or with sporidia
sporidia in nutrient solutions

fashion.
promycelium producing only infection-threads formed terminally and laterally near the septa; water usually germinate into infection-threads but in
multiply indefinitely,
yeast4
About two hundred species, seventy-two of which are given by Clinton ^ as occurring in America. Besides the species discussed below
many
others occurring upon grasses or plants of minor value are omitted.
other
U. avenae (Pers.) Jens.^^.
ne,
ii7. 124, 125
Sori in spikelets, rarely in leaves, forming a dusty olive-brown spore-mass, about 6-12 mm.
Fia.
219. U. ayeAfter
*'^*'^'
long by half as wide, usually rather completely
in water.
destroying
eventually becoming dissipated; spores lighter colored on one side, subspherical to spherical though often elongate, minutely echinulate, 5-9 ju in length,
floral parts,
widespread on oats.
The fungus was known by the name Ustilago
as early as 1552
304
THE rUNGI WHICH CAUSE PLANT DISEASE
and was called U. avenae in 1591. The species of Ustilago on oats, wheat and barley were considered identical until Jensen ^ showed that they are not intercommunicable. Wolff ^ showed that seedlings can be infected through the first sheath Brefeld ^ studying infection more closely found it to be leaf. accomplished by germ tubes from sporidia and that plants are free from infection after the growing leaves have pushed one
centimeter through the sheath leaf. The mycelium, after infection, grows through the plant until blooming time when it seeks the ovaries and the enclosing glumes in which it forms a mycelial
mass, which soon changes into spores. In nutrient solutions the conidia bud indefinitely, while on the host plant they produce
infecting hyphae.
Germination was
first
studied
by
Prevost.^
ily in
It occurs
read-
water, a well de-
veloped
resulting
Fig. 219.
promycelium in about
twenty-four
are
hours. The sporidia
elliptical.
mostly narrowly Fusion of

is
sporidia
common.
are
The
promycelia
usually four-celled
occasionally
especially base.
and
branch, the near

'''
[i''
U. crameri Korn.-^Sori in the spikelets,

of
infecting
spike,
all
the
ovate,
in
about
length,
int
2-4
FiG. 220.
mm.
Much
^^'^^-
Growing
enlarged,
point of the stem of barley. showing smut mycelium. After ^^^
chiefly destroying
rl ana K Dasai
1
parts,
spores reddish-brown,
chiefly ovoid to subspherical
though occasionally more elongate
and
irregular,
smooth, with usually pitted contents, chiefly 8-11 n
in length.

The promycelium
duced.
is
305
much branched but no

affects the ovaries of
sporidia are proSetaria.
The smut commonly
Panicum and
In America it has been collected on millet in several states. U. crus-gaUi T. & E.i"^
Sori often encircling stems at nodes or at the juncture of the
both stem and leaves, prominent, often by a tough hispid membrane which upon rupture discloses an olive-brown dusty spore-mass; spores ovoid to spherical, occasionally more elongate, rather bluntly echinulate or even verruculose, chiefly 10-14 /x in length.
inflorescence, infecting
nodular, one to several centimeters in length, protected
On Panicum crus-galli throughout the United States. U. bulgarica Bub. is on Sorghum vulgare. European. U. medians Bieden, on barley, is closely like U. hordei.^'*
U. scorzonerae (A.
&
S.) Schr.
on Scorzonera
is
very close to
U. tragopogonis-pratensis. U. sacchari Rab.^"^

in diameter, olive-brown or rufous, epispore thick,
Spore-mass black, spores globose or angularly globose, 8-18 smooth.
/x
On
world.
sugar-cane throughout the tropics, especially in the old
damage.
In Java this fungus has been reported as the cause of serious Barrett observed it in Trinidad, where the damage was
less extensive.
The leaves especially the young ones which have not yet separated from each other are the parts affected. From the upper part of the affected cane, as a rule, no secondary shoots arise, and those
which do
arise from the lower part become infected in their turn. discolored whip-like structure at the end of an attacked cane becomes dusty and black and contains the spores of the fungus.
The
U. hordei (Pers.) K.
&
S.-^-
''^' '^'
Sori in spikelets, forming an adhering purple-black spore-mass, about 6-10 mm. in length, covered rather permanently by the trans-
parent basal parts of the glumes; spores lighter colored on one side, usually subspherical or spherical, smooth, 5-9 ju, the most elongate rarely 9-11 /x in length. Common on barley.
This was
first
recognized as distinct from the oat smut in 1591
306

Lobelius.^
by
description.^^
Persoon in 1801 first gave a definitely recognizable In 1888 the species was separated from the other
smut on
barley.^^
by one, rarely two, tubes, usually 4-celled, and produce abundant sporidia; these increase by budding, produce germ tubes, or fuse with each other. U. levis (K. & S.) Mag.24- "^ Sori in spikelets, forming a black-brown adhering spore-mass,
^ N
The
spores germinate freely in water
sometimes small and entirely concealed by the glumes but usually evident and destroying inner and basal parts; spores lighter colored on one side,
subspherical to spherical or rarely elongate, smooth, 5-9 fx, the most elongate rarely 11 m in length. On oats throughout America and Europe, prob-
ably more
it differs XJ.
common than
difficult to distinguish
chiefly in its
records show as it is very from U. avense from which smooth granular spores.
FiG.
221. U.
le-
germination in modified C o h n 8 Afsolution.

vis,
'
macrospora Desm. and glumes, generally showing as linear striae, but often more or less merged, at first covered by the epidermis, but this later rupturing and disclosing black-brown dusty lines of spores; spores medium to dark reddish-brown, chiefly ovoid to spherical or occasionally somewhat irregular and elongate, coarsely verrucose, at
Sori in leaves
circumference usually showing the projections as tinted blunt scale-like appendages, sometimes even semi-reticulate,
ter Clinton.
12-19 M in length.
On
U. nuda (Jens.) K.
various species of Agropyron in Europe and America. & S.^^. ne
6-10
Sori in spikelets, forming a dusty olive-brown spore-mass, about mm. long by half as wide, temporarily protected by a thin
dissipated leaving the naked rachis colored on one side, minutely echinulate, behind; spores lighter in length. subspherical to spherical or occasionally elongate, 5-9
membrane which soon becomes
/j,
In Europe and America. This smut on barley is distinguishable from the covered smut, U. hordei, by its olive-green spore-mass and by its early shedding of spores. As a rule, each spikelet, ex-

cept the
307
In
awn and
rachis
is
entirely transformed into smut.
water and in nutrient solutions the spores germinate by a single promycelium, 1 to 3-septate, and often branched, but without
That infection is floral in loose smut of both wheat and was first shown by Maddox ^^ and the fact was corrobbarley The myand Hecke.^*"' orated by Wakagawa,^^ Brefeld celium has been demonstrated in the embryo by Broili.^^^ The spores falling between the glumes germinate, penetrate the ovary wall, and into the growing point of the embryo. The mycelium here lies dormant until the seed germinates, when it
sporidia.
^'^
^'^
grows, keeping pace with the growing point throughout the season and finally invading the ovaries to produce its spores.
The
infection of the pistil, the penetration of the integuments
and the nucellus and embryo sac was followed in microtome sections by Lang.^^^ The embryo was reached by the mycelium some four weeks after infection of the pistil. In resting grains the mycelium is abundant in the scutellum as well as in
all
embryo parts except the roots. Cross inoculation by Freeman and Johnson
^^
from barley to wheat and the reverse gave negative results. The optimum time for infection has been determined as the period of full bloom.
U. perennans Rost.^^^' "^
basal
more or less destroying the and inner parts, sometimes even running down on pedicels, oblong, about 3-8 mm. in
Sori in spikelets,
'germination 'm
ûtio*i?
length, with dusty, olive-brown spore masses; mycelium perennial in perennial parts of host;
Kellerman ^ se.
After and
spores chiefly subspherical or spherical, occasionally ovate to ellipsoidal, usually lighter colored on one side, more or less
minutely echinulate, especially on the lighter

length.
side,
5-8 n
in
On
the tall oat grass throughout its range. U. rabenhorstiana Kiihn occurs on several species of Panicum. U. tritici (Pers.) Rost.^^- î". 12-1, 125. 128
8-12
Sori in spikelets, forming a dusty olive-browm spore-mass, about mm. long by half as wide, usually entirely destroying floral
308
parts and eventually becoming dissipated and leaving behind only the naked rachis; spores lighter colored on one side, usually subspherical to spherical, occasionally elongate, minutely echinulate
especially
on the
lighter side,
5-9
ix
in length.
On wheat
delicate
where-
ever cultivated.
The smut mass

Infection
is floral
is
covered at
first
by a very
membrane.
as described for U. nuda.
The
7-septate,
spores germinate in water by a long 2 to 3, or even 6 to promycelium, often curved. In nutrient solutions the
Fig. 223. U.
;i4.
zeae,
25.
^^-
stages in spore development.
After Knowles.
promycelium branches profusely but sporidia are few or are entirely absent.
''''' '''U. zese (Beck.) Ung.i^- ""' '''-'-' '^^' Sori on any part of the corn plant usually prominent, forming irregular sweUings from a few millimeters to over a decimeter in
diameter, at
first
protected by a sort of false white

cells
membrane
composed
of
plant
and semi-gelatinized fungous threads,
soon rupturing and disclosing a reddish-brown spore-mass; spores ellipsoidal to spherical or rarely more irregular, prominently though rather bluntly echinulate, 8-11 fi the most elongate 15 /i
in length.
The
was
first
germina.tion of the spores, which occurs but poorly in water, studied by Kiihn ^^ in 1857. In 1874 Kiihn saw the pene-
germ tubes through the epidermis of the corn plant. Brefeld showed that the spores germinate well in nutrient solutions and that secondary spores are formed also that corn can be
tration of the
;
infected
by the sporidia
at
any point on
its
surface above ground

when the
is
30d
infection
tissues are soft

host.^^
and
actively growing;
and that
local
on the
It is now known that the chlamydospores are capable of germination without hibernation and that they remain viable one,
two,
years.
perhaps
It
more was shown

in
by
that
Brefeld
1895
chlamydospores produce conidia

in the air freely.
is
the
It
these,
air-borne,
from spores arising on the ground, manure, etc., which are

chiefly responsible for
infection.
They must
reach the plant on a

susceptible part and under suitable conditions
of
moisture.
The germ tubes from

the conidia penetrate the epidermis, grow
through
the
cells,
or
between
Fig.
223,
with an irregular mycelium which branches
Fig. 224. U. zese. 1, germination after three days in water; 2, similar but in air showing air sporidia. After Clinton.
profusely and calls forth great hypertrophy of the surrounding host tissue. In sporing, the mycelium forms a great number of
short, slender, irregular branches which make up a close tangled network in the diseased tissue. These slender branches swell, gelatinize, and portions of them round off as spores. Fig. 223.
U. striaeformls (West.) Niess.-^' ^-'^ Sori in leaves, sheaths and rarely in the inflorescence, from short to linear, often extending, apparently by terminal fusion, for
several centimeters, also occasionally fusing laterally to cover most of the leaf; at first covered by the epidermis but this is soon rup-
tured and dusty
brown
to black, linear
masses of spores become
310
scattered

and the leaves become shredded; spores usually
ellip-
soidal to spherical, occasionally irregular, prominently echinulate, chiefly 9-14 /x in length.

It
appears to be perennial.
is
The
spores germinate sparsely.
The promycelium
conidia.
long, branched, septate,
and produces no and
On numerous
species of
species of grass, including red top, timothy
Poa and Festuca throughout Europe and America.

;
Species of less importance, not all found in America are: U. schiriana Hem. which attacks bamboo ^^
European on rye; possibly a Tilletia. U. esculenta P. Hen. which causes swellings on Zizania which
U. secalis Rab.
is
are eaten in the orient; U. vaillantii Tul. in the sexual organs of the Liliacese;
U. panici-miliacei (Pers.) Wint. on Panicum miliaceum; U. tragopogi-pratensis (Pers.) Wint. on the flowers of Trago-
pogon; U. cruenta Kiihn, widespread in Europe on sorghum; U. violacea (Pers.) Fcl. on the anthers of various members of
the Caryophyllaceae; U. tulipae Wint. on tulips and related hosts; U. vrieseana Vuill. on eucalyptus roots, a very doubtful species; U. sphaerogena Burr, on Panicum crus-galli.
The fungus described

a Sterigmatocystis as and U. ficuum Reich on
is
as U. fischeri Pers.
is
also U. phcenicis
from Italy on corn Corda on date fruits
figs.
Sphacelotheca
De Bary
(p.
302)
Sori usually in the inflorescence, often limited to the ovaries,
provided with a definite, more or less temporary, false membrane, covering a dusty spore-mass; and a central columella, usually
formed
is
chiefly of the host plant's tissues.
The
false
membrane
which are
composed largely or
entirely of sterile fungous cells
hyaline or slightly tinted, oblong to spherical, and usually
more
or less firmly bound together; spores single, usually reddish-brown, developed in a somewhat centripetal manner as in Cintractia,
small to
medium
in size; germination as in Ustilago.
Sixteen species are recorded
by Clinton
for America.
Of these

only three are of economic importance. genus is not separated from Ustilago.
311
By Engler and
^^sise.
PrantI, the
Sphacelotheca sorghi (Lk.)
Cl.^'^'
^^'
in
Sori usually in the ovaries or stamens forming oblong to ovate bodies 3-12 mm. in length (rarely fusing the very young spikelets into irregular forms), protected for
some time by a false membrane upon the rupture of which the

olive-brown
tinct
spore-mass
of
scattered, leaving
becomes naked the displant tissue. the membrane

extent
into
columella
to
The
sterile cells of
break
groups,
up
some
oblong to subspherical, chiefly 7-18 / in length;

hyaline,
On C'o'i On
subspherical to spherical, smooth, contents often granular, in diameter. 5.5 8.5 spores
hVd^.
!^lfe
e."^
iJL
On Johnson grass and sorghum throughout the world. The young pistil and usually the stamens as
well are displaced
myceUum,
the
by the fungous two being often
blended together. The spores germinate readily in water, either
when
fresh or a year old,
showing
Fig.
225.
S.
b,
sorghi,
cross-section
through base of young infected
papillae in
from three to ten hours.

is
body (ovary),
celium,
a, false
The promyceUum
of its cells
2 to 3-septate and from the ends of one or more
of epidermal cells
and
mature
c,
sterile spores, b ',
membrane myimAfter
mature spores,
Clinton.
columella.
narrow tubes appear.
joints."
with the adjacent cell, forming the "buckle Either infection tubes or sporidia may also arise from the promycelium. Infection is possible only with young plants.
These
later fuse
lar,
grows rapidly into long irregumm. thick, which run through and between the cells. It is most abundant in the parenchyma, advancing especially through the pith region with the growth of the host. The young ovaries and stamens are eventually reached
hyaline, thin-walled threads 2-4
The mycelium
in the host plant
312
and the mycelium there develops richly under the epidermis. The cells remain sterile and constitute the membrane; the imier gelatinize and develop into spores.
outer
(Kuhn) Cl.'^' Sori very prominent forming irregular masses including more or
less
S. reiliana
''''
'''
''''
'''
of
the
entire
panicle,
usually
first
5-15 cm. in length; often at

tected
false
by the
leaf-sheath.
A
the
prowhitish
membrane
encloses
black-
brown spore-mass and the
ray-like re-
mains of the peduncles or columellas. In time it becomes ruptured and the

spores scattered. Sterile cells are also Scattered in groups through the sporeGerFig. 226.-S. reiliana. mination in water. After mass, chiefly subspherical, 7-15 n in Hitchcock and Norton. diameter; spores somewhat opaque, to spherical or occasionally ovoid or slightly chiefly subspherical
,
angled, minutely but abundantly verruculose, 9-14 n in length. Thig is a cosmopolitan but comparatively rare form on corn,
affecting the ovaries.
In germination It occurs also on sorghum. a 3 to 4-celled, often branched, promycelium is formed and conidia
are produced.
S. diplospora (E. & E.) CI. is found on Panicum crus-gaUi related grasses in the lower Mississippi Valley.
and
Sorosporium Rudolphi
(p.
302)
Sori in various parts of the host, forming dusty, dark colored spore-masses; spore-balls of medium size composed of numerous fertile cells, often rather
loosely united pletely
and frequently at maturity comspores

usually
olive
separating;
of
or
size; germination reddish-brown, similar to that of Ustilago; sometimes with elongate germ thread and no sporidia.
medium
Several species are parasitic on the coarser range grasses. S. consanguineum E. & E.,
S. everhartii Ell.
Fig. 227. Sorosporium. Spore mass. After Dietel.
&
Gall.,
most important.
S. dianthi
and S. ellisii Winter, are probably the Rab. is found on Dianthus.
313
Thecaphora Fingerhuth
(p.
302)
Sori in various parts of the host, often as indefinite masses in the floral parts or forming rather firm pustules on the stem, at ma-
turity with a dusty spore-
mass; spore-balls composed of few to many fertile cells,

of small to large size; rather
permanently united; spores usually yellowish or reddish,
smooth on contiguous sides but usually marked on the
Fig.
228. Thecaphora, spore

*^^-
ball ^^t^"" ^'"^f^^^-
gerraina-
free surface; germination, so far as known, by means of a single sporidium at the tip of the elongate septate promycelium. A small genus of slight economic importance. T. deformans Dur. & M.^-''' '-^
Sori in the seeds, showing
when the legumes
are broken open
as reddish-brown, dusty spore-masses which destroy most of the seeds; spore-balls reddish-brown, ovoid to spherical, rather firm,
composed of 3-25 (usually 7-12) spores, chiefly 27-60 ix in length; spores in optical section triangular to polygonal or when free irregular oblong, free surface with papillae that sometimes vary to
spiny processes, 15-25 m, chiefly 15-20 m in length. On a large number of Leguminous hosts, including species of Vicia, Lathyrus, Lupinus, Trifolium, etc., in widely scattered regions of both the old and the new world.
Tolyposporium Woronin
(p.
302)
Sori usually in the inflorescence, especially the ovary, forming granular spore-masses at maturity; spore-balls dark-colored, of
numerous spores permanently united, germination about as

Ustilago.
in
A
T.
genus of about ten species.
buUatum
Schr.i^s. i54
Sori in ovaries, infecting occasional ones, ovate,

in length,
about 3-5
mm.
covered with a thin, greenish, smooth membrane, upon of which the black granular spore-mass becomes scattered; rupture
314

spore-balls black, opaque, oblong to spherical or polyhedral, usually containing one hundred or more
firmly agglutinated spores, chiefly 50-180 M in length; spores from
nearly
hyaline,
to
light
reddishor
less
brown,
outer
coat
more
Fig.
229.
Tolyposporium.
folded in ridges, often spiny, ovoid or to subspherical polyhedral, or rarely 12 /x in chiefly 7-10
jjl
Spore
ball germination.
After Brefeld.
length.
On ranicum
/-^
-r
n-
crus-galli
the
,i
United States east of the Rocky Mountains also in Europe. T. filiferum and T. volkensii, occur on sorghum in Africa.
TiUetiacese
Sori either forming dusty
(p.
302)
embedded
in the tissues.
erumpent spore-masses or permanently Germination by means of a short promy-
celium which usually gives rise to a terminal cluster of elongate sporidia, that, with or without fusing in pairs, produce similar
or dissimilar secondary sporidia or germinate directly into infection
threads.
The American Tilletiacese embrace nine genera and about one hundred twenty-five species.
Key
Spores single
Sori dusty at maturity
to Genera of Tiiletiaceae
Spores without a conspicuous hyaline
appendage
Spores with an elongate hyaline appendSori
1.
Tilletia, p. 315.
age permanently embedded in the tissues
2.
Neovossia.
Sori definite, small

Sori indefinite, large
3.
4.
Entyloma, p. 320. Melanotaenium.
Spores in balls
Sori dusty; spore-balls with sterile cor-
tex
5.
Urocystis, p. 318.

Sori
315
rather
tissues
permanently embedded
without
sterile
in
Spore-balls
cortex
of
6.
Spore-balls consisting entirely dark-colored spores.
Tuburcinia.
Spore-balls consisting of light-colored
spores Spore-balls with or without central

sterile cells
7.
Burrillia.
Spore-balls with central network of filaments Spore-balls with sterile cortex
8.
9.
Tracya. Doassansia,
p. 322.
Neovossia occurs on Phragmites; Tuburcinia on Convallariacese, Primula, Trientalis and Geranium in Russia; Burrillia on Limnan-
themum, Echinodorus and

Tilletia
Sagittaria;
Tracya on Spirodela.
Tulasne "
(p.
314)
Sori in various parts of the hosts, usually in the ovaries, forming dusty spore-masses; spores single and usually formed singly in
the ends of the mycelial threads
which disappear more or less completely through gelatinization, germination usually by a short promycelium which bears a terminal cluster of elongate
sporidia that in nutrient solutions, with or without fusing in
pairs,
may
give rise to a consec-
siderable
mycelium bearing
closely
in
ondary
air-sporidia.
The genus
Ustilago
spores
resembles
its
except
larger Fig. 230. T.
fcetens,
spores.
Photo-
micrograph. and mode of germination. Twenty-two American species are listed by Clinton. Only three are of economic importance. T. pancicii Bub. & Ran. is reported on barley heads in Servia.^^ T. giomerulata. Coco. & Mor. is a doubtful species on alfalfa.
After Clinton.
316

(B.
T. foetens.
Sori
less
&
C.) Trel.^^'
i^^-
120. 135
ovate or oblong, 5-8 mm. in length, more or concealed by the glumes, all or only part of the ovaries of
in ovaries,
a spike infected; spores light to dark-brown, oblong to chiefly subspherical or spherical, occasionally
pecially
when young, smooth,

fx
somewhat angular, foetid, eschiefly 16-22 n, the most elongate
rarely 28
in length.
On wheat wherever
Kiihn
^^
grown. found that infection occurs as
in
oats in the very
young plants. From the infection point the mycelium approaches the growing point and follows the development of
its
host, sending its branches into each
spikelet
develops a close knot and in the ends of the threads and in the
ovules.
it
and Here
finally
into
the
growing
short
branches
the
spores
form.
The
spores germinate by a rather long, continuous, thick promycelium on the tip of
which a crown of long slender conidia develops.
The
and often
Fig.
sporidia soon become arched fuse in pairs; they develop inWint.^^S
fection threads.
231. T.
faêns.
A.
T.
triticl (Beij.)
Sori in ovaries, ovate to oblong, 5-8 mm. fnTiôri^fa; TsZldfa which have united. One in length, more or less concealed by the
has produced a secondary and this is sporidium at
hyaline, subthread^ ^Âfter Freeman spherical, with medium-thin wall, smaller and Stedman. than the fertile cells which are chiefly
i,
<.
i-
glumes;
sterile
Cells
few,
subspherical,
about
/x
to dark-brown, with winged reticulations 2-4 n wide, and 16-22 î in diameter. high by
light
On wheat
everywhere. Experiments have shown this distinct from T. foetans which
it
closely resembles except for its reticulate spores.
T. texana Long: CW^^ Sori in ovaries, ovoid or oblong, about 3-5

or less hidden
mm. in length, more by enveloping glumes, forming a somicwhat aggluspore-mass;

sterile
cells
tinated
light-reddish-brown
not very

numorous, cells very
h,yaline,
317
with very
tliick,
light colored, orange-yellow
often lamellate walls; fertile appearing as if immature,
chiefly subspherical or spherical, with prominent conical tubercles which extend out 2-3 to the hjâline envelope, chiefly 19-25 /x in diameter (inchuling envelope.)
fj.
On Hordeum nodosum
T. hordei
in Texas.
an Asiatic form on Hordeum. T. secalis (Cda.) Kiihn. occurs on rye in Europe.^^^

is
Kcke
T. horrida Tak.^^'
iî-
no
Sori in the ovaries
more
or less destroying them, completely
Fig.
2.S2.
Tilletia
moist
air,
tiitici. A. Two spores Kerminatcd in promyccliuni and coiiidia, several of which have
fused in pairs. Secondary conidia at C. B. Sjiores germinated in water, proniycelia elongate, septate. Tlie proover into the younger cells. After Tubeuf. toplasm passes
concealed by envelojjing glumes; spores usually present in different stages of development, the mature spores almost opaque, chiefly
subspherical to spherical, with very coarse hyaline or slightly tinted, somewhat curved, scales which show at the circumference of the spore as a band about 2-4 fx wide and on its top as polyg-
onal areas 2-3

in length.
fx
and often at one
across; hyaline membrane more or less evident side in a short thread-like projection, 22-33 n
Cross sections of stems l)earing smutted heads reveal the mycelium in the chlorophyll parenchyma between the fibrous tissue.-
On
rice in
America and Asia.
318

Urocystis Rabenhorst
^"^^
(p.
314)
Sori usually in the leaves or stems, occasionally in other parts,
producing dark-colored, usually dusty, spore-masses; spore-balls permanent, composed of an enveloping cortex of tinted sterile cells and usually one to
Fig.
233. Spore several interior fertile

;
cells;
fertile cells generally
After dark-colored germination often by a short promyceThaxter. jj^^ ^^^^^ ^^^_

puia'.
duces terminally-grouped sporidia;
these give
rise
to
similar
secondary sporidia or to infection-threads.
Besides
the
forms
discussed
below, foreign species are listed
on Anemone,
Liliacese, Gladiolus,
^''
Primula, etc. U. cepulffi Frost.2"'
" i^^
Sori in leaves, forming isolated pustules or affecting them for
the greater part of their length
and breadth, sometimes occurring at their bases, in the bulbs. Upon rupture of the covering
membrane a dusty black-brown

spore-mass appears; spore-balls ovoid to spherical, 17-25 fx in length; sterile cells tinted, ovoid
to spherical, small, rather
com-
covering spores, usually 4-8 fx in length; fertile cells reddish-brown, ovoid to

pletely
spherical, usually
1,
the
ball, chiefly
12-16
fx
rarely 2 in Fig. 234. Spores of U. cepulce germiin length. nating. X, sporidia; 23, germinating
conidium.
After Tliaxter.
On AUium.
The
first
American description
of the fungus
was by Farlow
^^
in 1876.
A second thorough paper was from Thaxter in 1889.^^ The mycelium grows between the host cells. At maturity lateral
319
outgrowths appear from the hyphae at various points. One of these assumes a somewhat spherical form and matures to the fertile spore, while the other branch or branches grow around it, Fig. 235, branching and dividing into joints which eventually round off to
form
the
sterile
exterior
cells.
Spores
are
known
^^ to live in soil for at least twelve years. period of rest is necessary before they can
=^
germinate.
In germination the central spore
produces a single short hypha, commonly branched, on which the conidia are borne terminally and laterally.
ranean.
Fig. 234.
Experiments
is
by Thaxter indicate that infection

U. occulta (Wal.)
Sori
in
subter-
RaW^'
'''
leaves,
especially
in
the sheaths,
striae
culms and inflorescence, forming linear
Fig. 235. Successive stages in formation of spore balls. Af-
ter Tha.xter. usually of great length and often merged into a continuous stratum of dusty, reddish-black, spore-balls; sporein length; sterile cells balls oblong to subspherical, 16-32
jjl
often incompletely covering the spores, hyaline or yellowish, subspherical to oblong, usually with distended and uniformly thick-
ened walls;
fertile
cells
1
often flattened, smooth,
reddish-brown, oblong to subspherical, to 4 in a ball, 11-18 /i in length.
On
The seat
though
rye wherever cultivated, though not common in America. of spore formation is most often on the stems or sheaths,
all aerial
parts of the plant are susceptible. In the vegetative parts the fungus is commonly found in the tissue between the vascular bundles.
U. violae (Sow.) F. de W.^^s,

Sori
i43. loo
on stems, rootstocks, petioles and leaves forming prominent irregular swellings often sevFig. 236. Spore balls of U. oc- eral centimeters in length, rather permanently c u 1 a -'^^ter covered by the host tissues but upon rupture Th disclosing black-brown spore-masses; spore-balls
1
reddish-brown, rather irregular, oblong to subspherical, chiefly
28-55 n in length;
sterile cells yellowish-tinted
with age, 6-10 m

in length.
in length; fertile cells light reddish-brown, ovoid to spherical or
polyhedral, chiefly 4-8 in a ball, mostly 11-15
fj.
320

violets.
On
In America
it
has been reported in Canada, Min-
nesota and Utah.
U. anemones (Pers.) Wint.^^^ occurs on various species of culaceae in both the old and new world.
Ranun-
U. agrop)rri (Preu.) Schr.^ Sori in various parts, commonly in leaves, forming striae, which may be distinct or cover the surface of the leaf; at first lead-colored
and protected by the epidermis but soon rupturing and

fj.
scatter-
ing the reddish-brown spores; spore-balls oblong to subspherical, in length; sterile cells hyaUne to yellowish, oblong to 16-32
subspherical, usually completely covering the fertile cells, outer wall thin and by collapsing giving a ridged effect to the covering; spores 1 or 2, rarely 3 or 4 in a ball, reddish-brown, oblong to subspherical, often flattened, smooth, 11-18
/x
in length.
On Agropyron and some

United States and Europe. U. colchici (Schl.) Rab.^^^
other coarse grasses throughout the
On
various species of Liliaceae but

is
not on hosts of economic importance in America. U. italica Speg. probably not a true smut,
acorns, chestnuts
injurious to
and the seeds of the white Species of less importance or non-American U. gladioli (Req.) Sm. on Gladiolus; U. ornithogali Korn. on Ornithogalum; U. kemetiana Mag. in pansy ovaries; U. primulicola Mag. on primrose flowers.
fir.-^
are:
Entyloma De Bary
^^-'
^^^
(p.
314)
Sori usually foliar, generally forming discolored but not distorted areas, permanently embedded in the tissues; spores single, pro-
duced terminally or intercalary in the mycelium which does not entirely disappear through gelatinization, free (sometimes irregularly adhering through pressure), hyaline to yellowish or reddishyellow, rarely dark-colored, germination by a short promycelium bearing a terminal group of sporidia which usually conjugate in pairs and produce secondary sporidia or infection-threads; sporidia often formed by germination of the spores in situ, the promycelium
protruding through the stomata.
Twenty American
species are recorded.
321
Foreign species are on Papaver, Ranunculus, Delphinium, Calendula, Thalictrum and several other hosts. E. betiphilum Bub. is described on beet seed capsules;^''
E. lephroideum for the same host in France; E. calendulae (Oud.) de B. on Calendula, E. crastophilum Sacc.^ Sori in leaves, subcircular to linear, about 0.25-2
mm.
in length,
usually distinct
though occasionally merged, black, long covered the epidermis; spores dark-brown, tightly packed and adhering by
E. cllisii, rhlamydosporcs germinating within Fig. 237. the leaf tissue, sporidia superficial. After Halsted.
more
or
less, chiefly
ovoid to spherical or angled through pressure,

ju
rather thick-walled, 8-14
in length.
Poa, Phleum, Agrostis and other grasses in Europe and America.

E. irregulare Joh. occurs on species of
On
Poa
in
Europe and
America;
mon
E. polysporum (Pk.) Farl. on various hosts including the comsunflower.

E.
ellisii Hals.'î
Sori in leaves, forming pale white spots, indefinitely limited, subconfluent; spores hyaline or slightly yellowish, clustered in the
intercellular spaces
beneath the stomata, spherical, thick-walled, (2-5 fx) chiefly 16-20 m but varying from 11 to 25 m in diameter; conidia hypophyllous, abundant, acicular, small, 10-14 fx by less
than
1
/x.
On
spinach.
New
^^^
Jersey.'^^'
The chlamydospores germinate in

and bear the sporidia on
the stomata, presenting
situ
tufts of promycelia
beneath the stomata which emerge through
much
the appearance of a Hyphomycete.
322
E. australe Speg. Sori foliar, forming spots, yellowish to eventually dark, usually 0.5-6 mm. in length; spores light to reddish-yellow, ovoid to
spherical or slightly angled, chiefly 10-16
linear,
/i
in
length; conidia
somewhat curved, usually 30-55 x 1-2 fx. Common and destructive on many species of Physalis and on Solanum, especially on some of the cultivated forms throughout the Americas and in Africa. E. fuscum Schr.^^^
Sori in leaves, about 2-6
mm.
or
by confluence much
larger,
spores light yellow to chestnut-brown, provided (especially when young) with a conspicuously swollen gelatinous envelope, smooth,
13-19 m in length; the hypophyllous matted outgrowths usually show few conidia which are- fusiform, single-celled or septate, 10-22 X 3 M.
chiefly
It occurs
on Papaver
in
Europe and Eastern North America. and

irregular areas,
E. nymphaeae (Cunn.) Set.^^'' Sori in leaves, forming variable
usually
with age reddishbrown, scattered or confluent; spores hyaline, ovoid to subspherical, usually apiculate and with the remains of the hypha as a basal
side, yellowish or
most prominent on the under
appendix, smooth or under an immersion lens minutely verrucuin length; conidia not observed but spores said to lose, 10-14 germinate in situ.
fjL
On
leaves of various water
lilies
in
both the old and new world.
Doassansia Cornu."^' "
(p.
315)
Sori in various parts of the host, usually

in the leaves, rather permanently embedded in the tissues; spore-balls conspicuous, per-
manent,
layer
FiG. 238. -Doassansia. Part section through a spore ball showing
sterile
and a
cells
i
consisting central
of
distinct
cortical
mass
of fertile cells en-
^'^^^y ^^^'""^ ^^^ interior, or
with the inneri.
most
cclls
Supplanted by parenchymatous
i
and
fertile cells,
OF hyphal threads; spores hyalme or
,i
After Dietel.
germination often in
yellowlsh, with smooth, usually thin, walls; situ, by means of a short promycelium which
323
gives rise to a terminal group of elongate sporidia, these often
bearing secondary and even tertiary groups. The only species of this genus which occur on economic plants are D. gossypii Lagerh.^'- on cotton in Ecuador and D. niesslii
de Toni (Niess) Schr. on Butomus. The following genera, which are usually referred doubtfully to the Ustilaginales will be found under "Genera of Unknown Affinity" page 663.
Graphiola Poit. on various palms. Bornetina M. & V. on Vitis.

Protobasidii
(p.
Schinzia Nag. on Solanum.
299)
The
three orders which belong to this group are characterized

basidia.
by septate
Key to Orders of
Basidia with cross walls
Basidia arising from chlamydospores, Life cycle polymorphic. Parasites
Protobasidii
1.
Uredinales,
p. 323.
Basidia not arising from chlamydospores Not polymorphic. Gelatinous sapro-
phytes Basidia with lengthwise partitions, gelatinous saprophytes
2.
Auriculariales, p. 392.
3.
Tremellales.
Uredinales"' "' "'

ferns
*'
"'
^^'^-
^^-^^^' ^'^' ^^^-^^^
Small fungi, mostly microscopic, parasitic in the tissues of and seed plants. Mycelium much branched, septate, and
Spores borne in
sori
with haustoria.
below the surface of the host,
or rarely single within the host. Sori naked, enclosed by peridia or paraphyses, or embedded in a thin stroma. Spores of five mor-
phological sorts, not all present in every genus; minute, thin-walled, without surface sculpturing,
small, smooth, of
(1) basidiospores,
(2)
pycniospores,
function, (3) seciospores, verrucosely sculptured, borne in chains, (4) urediniospores, echinulately or
* Arthur's terminology involving the words pycnium, spcium, uredinium, telium and derivatives from these words, will be followed in the treatment of
unknown
this order.
324
verrucosely sculptured, borne singly, or sometimes in chains, (5) teliospores, smooth or variously sculptured but not echinulate,
borne singly or in chains. In every species the mycelium eventually gives rise to teliospores, which produce in germination four basidia, either remaining within the spore-cell or borne in
the air on a short proraycelium, each basidium supporting a single, stalked or sessile basidiospore.
some two thousand species, constituting the many of them living on cultivated plants of high Its members are strict, value, is of great economic significance. which in no stage of the life except in the obligate, parasites promycelial stage can develop other than on the living host. The complexities of the life histories of the species, with their five distinct spore forms, inhabiting at different seasonal periods two
of
The order
"rust" fungi,
or
plants, renders the order

difficult
even three different host both
and exceedingly
history of the
in-
teresting.
The
life
most
complete of these fungi be stated as follows:

I.
may
O.
iEcia
(aecidia)
and
pycnia (often
called
spermo-
gonia or pycnidia). The mycelium arising from a basidiospore invades the host plant,
and vegetates
ficient
0&
Fig. 239.
until vigor sufspore formation is attained, meantime often pro-
to
^cium and pycnium.

Tavel.
After
ducing local spotting, hypertrophy, or other injury to the host. The mycelium then de-
velops a stroma which produces spore beds (son) and ruptures the epidermis. These sori are usually deeply sunken in the host and cup-shaped and take the
common name
aecidia.
The sporophores
"cluster cups," Fig. 239, technically aecia or arise from a hyphal plexus at the base
in acropetal sue-
of the
cup and the spores are borne catenulate

cession.
325
The whole
cup
layer of the
is usually red or yellow. The outer consists of a palisade of sterile sporousually
structure
phores bearing
sterile cells
and constitutes the peridium.
The
aeciospores are usually nearly globular, or angular by compression, reddish and rough and sometimes bear germ pores. They are capable of germination at once and on germination give rise to germ
tubes which
may infect susceptible hosts, leading to a mycelium. This in turn again produces sori which in some species may be secia, in others telia, but in most species, uredinia.
Associated with the secia, occasionally with other spore forms, but never borne alone, are minute pycnia with sporophores arising from their walls and bases. These bear unicellular pycnioSterile hairs usually protrude from the ostioles. The whole structure in gross appearance is much like the pycnidium of Phoma or Phyllosticta but it is reddish or orange in color. " " These pycnia were formerly often spoken of as spermogonia and the spores as "spermatia," due to the thought that they stood for degenerated male organs; a view supported by the fact that the spores were not observed to germinate. Germination has now been observed and there is no longer reason to regard them
spores.
"^''^
as sexual organs.
II.
Uredinia
of
(uredo-sori).
The
seciospores
may
infect
the
same
plant produced the seciospores (autoespecies or plants of an entirely different species (heteroecious). cious)
that
The mycelium produced by the

host; usually remains
local,
seciospore develops within the
and causes spotting.
When
it
has
attained sufficient vigor and age, usually after about two weeks, it produces a sub-epidermal hyphal plexus from which arises a bed
of sporophores
which bear
unicellular, hyaline to
brown, nearly
globose, thin-walled, usually echinulate or rough spores, each with from 2 to 10 germ-pores variously placed. These are the ure-
diniospores borne in uredinia (uredo-sori). They may germinate at once producing a germ tube which develops to a mycelium.
These spores falling on susceptible tissues, by infection, usually stomatal, continue the production of uredinia and spread the disease. The urediniospores are usually short-lived and function
to spread
summer
infection.
They continue
to form throughout
the growing season.
326
In a few species ^^^' ^^^ there are what are known as amphispores or resting forms of urediniospores provided with thickened walls. They have colorless contents and pedicels more persistent than
those of the usual urediniospore. III. Telia (teleuto-sori). Toward the latter part of the growing seasons another kind of spore appears, often in the same
sorus with the urediniospore
and from
of
the
same mycelium.
varies
in
It
is
various
forms in different genera, one or morecelled,
shape,
thickness
of
marking, color, etc., but is uniform in the character of the germination which is very different from
wall,
surface
that of any of the other rust-spores.

teliospore germination, typically of the teliospore sends forth one germ tube. These tubes soon cease
In
each
cell
growth and by septation become
4-celled.
Each
cell
then sends out a short branch
round or
(sterigma) on which there develops one oval, 1-celled, thin-walled spore,

in
this
the basidiospore, often called the sporidium.
group
Fig.
240 Germination of teliospores of P. asparagi
After Smith
Morphologically the promycelium is a basidium bearing its four sterigmata and four basidiospores. Relationship is thus shown on the one hand to the Ustilagi-
nales, on the other hand to the Auriculaan assumption that is borne out by cytological evidence. Deviations from the typical mode of germination are found in several genera mentioned below (e. g., Coleosporium). Basidiospores germinate immediately by germ tubes which on suitable hosts give rise again to secia and pycnia or in some species to other spore forms completing the life cycle. The most complex lile cycle is thus seen to comprise pycniospores, seciospores, urediniospores, teliospores and basidiospores. For brevity the first four stages are commonly designated by
riales,
the following symbols:

0. Pycnia or pycnial stage 1. ^cia or ajcial stage
II.
327
Uredinia or uredinia stage
III. Telia or telial stage
The
spores in
all
cases, except those of the basidiospores
and
Fig. 241.
Anlphi^^pores, urediniosporcs and teliospores of Pucciriia vexans.

After Holway.
pycniospores arise by direct conversion of a mycelial cell into a spore, i. e., they arc chlamydospores. Mesospore is a term applied to occasional unicellular forms of
teliospores found in Puccinia
and related genera which do not have unicellular teliospores. usually As has been said the pycniospores seem to be functionless though by some it is thought that they do function but that man has yet failed to find the conditions under which they readily germinate and cause infection. The acial stage appearing first, and thus commonly in the spring, is often called the "spring stage."
328

an early stage to propagate and spread the fungus. uredinia often called the "summer stage" constitute the
It serves as
The
phase usually of longest duration and of most injury. Its function is preeminently to multiply and spread the fungus. The telia, often called the "winter stage," usually, but not always, constitute the resting, hibernating stage. In many instances the teliospores must rest over winter before they are capable of
is based primarily on the teUospores. of the spore forms discussed above are typical of many species there are many other species which do not possess all of these forms or indeed which may possess only one spore form.
germination.
Classification
While
all five
for convenience groups the rusts, according to the Schroter spore forms that they show, under the following type names though it must be recognized that such grouping is purely artificial and does not necessarily bring together closely related species.
Eu-type Brachy-type
Opsis-type
0, 0, 0,
I,
II,
II,
III III
III
present;
present; present;
present;
omitted.
omitted.
I,
II
Hemi-type
Micro-type
Lepto-type
II,
III
O,
omitted.
only III only III
present; germination only after
a resting period. present; germination without a

resting period.
As examples
of the
Eu-type, Puccinia asparagi,
above we have the following: 0, 1, II, and III,

II,
I,
all
on Asparon on
thistle.
salsify.
agus.
Brachy-type, Puccinia suaveolens, 0, Opsis-type, Puccinia tragopogonis, O,
and and
and
III, all
III, all
Hemi-type, Uromyces caryophyllinus,

II,
III,
both on Dianthus.
Micro-type, Puccinia ribis. Lepto-type, Puccinia malvacea-
III,
on Ribes. on hollyhock.
rum,
III,
Hundreds
of the hemi-types will doubtless be revealed
by study
to be heteroecious eu-types.

Heteroecism.^-
i.
329
e., all
plant.
examples just given are autoecious, forms are found on the same species of host spore In many other rusts, however, heteroecism prevails, i. e.,
All of the
known
one stage of the fungus is found on one species of host and another stage upon another host; rarely three host plants are involved in the cycle. Aside from the rusts only one other fungus (Sclerotinia
ledi) is
known
to
show heteroecism.
Heteroecism has been experimentally proved in some one hundred and fifty cases and may be assumed to exist in many hundreds of cases not yet investigated.
Examples
of heteroecism are as follows:
Eu-type, Puccinia graminis,

" "
sorghi,
Stages 0, Berberis
I.
Stages
II, III.
Wheat
Corn Pea
rubigo-vera, Boraginaceas Oxalis
Opsis-type,
Uromyces pisi, Euphorbia Gymnosporangium macropus. Apple
Red
cedar (III)
It
frequently
is
happens that part

of the life cycle
passed upon a mo-
nocotyledonous
plant,
the remain-
der upon a dicotyledon. In such
event
often
it
is
the
II
more and
III stages that are
on
the monocoty\V
ledon while the O, I stages are on the

dicotyledon; examples of this are af-
forded
in
the nurusts of
merous
F'^- -4-Urediniospore of P. asparagi germinating on surface of plant, and separate spores. After Smith.
grasses, sedges
and
rushes.
In one group the pycnia and the
330
secia are on pines (Peridermium), while the other stages are on dicotyledons. In the Gymnosporangiums the pycnial and secial stages are on Rosacese; the telial on Juniperus and its kin. While a few general rules can be worked out concerning host relations there are many exceptions and to know one stage of a heteroecious rust generally gives little or no clue to what its
complementary host may be. The mycelium of the rusts is usually intercellular and local though in a few instances it is extensive and even perennial in the host. It is abundantly branched, closely septate, gives off haustoria and usually bears numerous oil drops which lend a yellow or orange color. Irritation by the mycelium often induces marked hypertrophy or even witches' brooms or other deformation of the host. Hypertrophy is most common with the secia but may result from the telia as
^^
I ),
well, as is conspicu'
ously shown in the
J)\
genus angium.
habit
GymnosporIn some
the host
instances the whole

of
plant is altered by the presence of the
mycelium so as to
render
it
almost un-
FiG. 243.
Cross-section, showing infection

of P. asparagi.
recognizable, e. g., the secium of Urofrom spore rnyoes ^

_
After Smith.
oisi '^
On Eu-
phorbia.
seldom killed by the mycelium, which abstracts its food supply from the carbohydrates and other nutrients of the cell sap without direct injury to the protoplasm, though ultimately there is serious effect upon both growth of the host
cells are
The host
and
its
seed production.
ISO' 189'
31^31^
Cytology.1^9'
Dangeard"4 and Sappin-Trouffy
^^^^
showed that the mycelial cells of the rusts are binucleate, a condition which begins just below the secium. The origin and significance of this condition
is
of
much
interest.
THE
FUxNGI
331
In all of the rusts so far investigated that have an aecium or primary iiredinium there is in the aicio-mycelium or the primary uredinio-myceliima a fusion of uninucleated cells, gametes. This
after long delay;
not, however, followed by a nuclear fusion until but the two nuclei remain in the fusion cell and when this cell divides both nuclei divide mitotically and simultaneously but still independently of each other (conjugate division). This process continues through the secial sporophores, or uredinial
cellular fusion
is
sporophores, and in the production of the spores, with the result
"^^jC.l
^^'r'/^%.--C.m.
C.nv.j.
Fig. 244.
i--h S h o w n g
i
Fig.
245.
Gymnospoclavariaj-
Fig. 246. nuclear
Conjugate
division
in
conjugate nuclei and degenerating cells in chain conidiospore of ^cidium. After Sappin-Trouffy.
rangium,
forme, mitosis of a nucleus in the promycelium. After Blackmail.
G.\Tnnosporangium clavaria?forme showing four chromatin
masses.
After
Blackman.
that the
cells of all of
these are binucleate.
The conjugate
and
divi-
sion continues further through the uredinia
until teliospore
formation occurs, the whole intervening series of cells being binuPrior to the formation of the promycelium and in the cleate.
teliospore the nuclei unite, reducing the cells again to
cleate condition.
an uninu-
In rusts which have only teliospores the binucleate condition begins somewhere in the mycelium from which the teliospores
arise.
It
is
generally held that the cellular fusion
is
a sexual act
with long delayed fusion of the sexual nuclei: and consequently that the uninucleate phase is the gametophyte; that the be-
332
ginning of the binucleate condition marks the origin of the sporophyte.

Biologic specialization iphales occurs also in the Uredinales.
of
^''^'
1^2.174. 309
mu^h
as
is
There are
found in the Erysmany species, each
which
is
hosts the fungus
found on a large number of hosts. Upon its numerous may show no morphological variation, yet at-
Fig. FiG. 247. Conjugate nuclear division in cells of P u c c i n i a podophylli.
248.
Diagram-
matic representation
of fusion of nuclei in the teliospore. After Delacroix and
After Christman.
Maublanc.
tempts to inoculate from one host to another
may uniformly give negative results. It further often occurs that one stage, e. g., the secia of a species may grow upon only one host while the uredinia or telia may grow upon many different species of hosts;
and in such cases that seciospores which have arisen on host X, from infection with spores from host A, are capable of infecting host A and that host only; while seciospores which have arisen on host X, by infection with spores from host B, are capable of infecting host B and only this host; and so on for numerous forms. Yet the uredinia and telia of these different races may be mor-
PLAxNT DISEASE
333
phologically inseparable as are also their seeia

their
when grown upon
common
excellent
host.
An
in the
example
of such biologic specialization is offered
common
pine Peridermium.
yEcia
may
be produced upon
the pine by sowing of Coleosporium teliospores from Senccio, Campanula, Pulsatilla, etc., but the seciospores which develop on the pine are capable of infecting only hosts from
those species
of
which the
^'^^
telio-
spores were taken.

Similarly Eriksson
has de-
termined that though rusts from many grains can infect

barberry, the seciospores there produced are not capable
the
infecting plants of species other than those from which

of
the fungus was derived, or at most they can infect but a
very limited number of species. A further complication arises from the facts obtained through
experiments
tries,
FiG. 249. Urediniospores in Rubus showAfter Blacking nuclear conditions.
in
various
coun-
man.
is
which have shown that what

consist of a large
apparently the same species
which behave differently in different geographic areas. The stem rusts of wheat and barley, for instance, are very similar, interchanging hosts easily and being capable of transfer to various grasses in this country, though in Sweden the stem rust of wheat goes with difficulty to barley and rye, while the stem rusts of barley and
may
number
of strains or varieties
rye interchange hosts very easily. Owing to the prominence of its author and
ture a
its
place in litera-
word may be given

i9o-i'J2.
to the usually discredited
mycoplasm
theory
312
^f Eriksson.
This affirms the existence in the
cells of wheat grain of an intimate mixture of rust protoplasm and host protoplasm. This mycoplasm may rest thus for months. the host-cell nucleus becomes digested and the fungous Finally
plasm develops to a mycelium which proceeds to invade the sur-
334
rounding tissues of the seedling as these develop on germination

of the seed.
Infection
rusts
Experiments.
their
by observing
life
Since the method of studying the histories in the laboratory where
they are under complete control of the observer has assumed such prominence of late years the technique deserves notice. The first step is to find associated in the field the secia and other stages of a rust in such way as to suggest relationship between
Material of the rust

are also
lected in the
till
two forms hitherto unknown to be connected. is then collected and healthy host plants removed to the laboratory. If the teliospores are col-
fall they are kept out of doors in cheese cloth bags germination time in the spring. Whether collected in spring or fall the viability of the spores must be tested by sowing in a
hanging drop of water. If germination is plentiful then the infecis made. First the suspected alternate host is sprayed with water to give the spores proper conditions for germination, then masses of spores are placed directly on the plant by a scalpel and a bell jar is placed over the plant to assure a humid atmosphere. In from five to eight days yellow spots should indicate where the infection has taken place and in a short time pycnia and secia or other sori follow. In all infection work it is
tion experiment
imperative to know that the plants used be not already infected in the field from another source.
The
is
aecium
is
by some regarded as a structure whose function

^
to
^''^ restore vigor to the rust fungus.
Freeman and Johnson

of
On the other hand, found that in j&fty-two generations
was no apparent diminution

generation.
the fungus, without the intervention of aecia or telia there in the vitality of the uredinial
telial
Form Genera. The

stage of the rust fungus
and
is
stage is regarded as the highest the one on which classification is
often based.
Thus an aecium, uredinium, cseoma, etc., that is to possess a teUal form is regarded as part of the species indicated by its teliospore, e. g., iEcidium berberidis being part
known
of Puccinia graminis has no specific identity but is regarded as a stage of P. graminis. There are numerous uredinia, secia and other non-telial forms

of which the telial stage
for
is
335
not yet known. It beconxes necesconvenience of reference, to have sary names by which to designate these forms. For this purpose the form-genera /EcicUum Caôma, Peridermium, Roestclia and Uredo are recognized. We group these under the heading Uredinales
the
present,
for
Imperfecti.
Darluca and TubercuHna, two imperfect fungi, are often found growing as parasites upon the rust fungi.
Key to Families of Uredinales

Teliospores in germination becoming
i-celled,
compacted
laterally
into
waxy
layers; walls of the

1.
spores weakly gelatinous

Teliospores germinating celium
Coleosporiaceae, p. 335.
by a promy-
Teliospores
into a crust or
solitary
compacted laterally column (rarely

within
the tissues);
2.
walls of the spores firm Teliospores free or fascicled; walls

of the spores firm or with an outer hygroscopic layer covered by cuticle
Melampsoraceae,
p. 340.
3.
4.
Teliospores
unknown
Pucciniaceae, p. 353. Uredinales Imperfecti,
p. 389.
Coleosporiaceae
Teliospores united in a one or two-layered waxy cushion, sessile or borne on a broad sac-like stalk and then at the beginning
2-celled.
cells
Each
original spore-cell divides to four super-imposed
from each of which a simple sterigma emerges.
This bears a
large basidiospore.
is the peculiar mode of basidiothe 4-celled promycelium being formed within spore production, the spore.
The most important character
The family is of stage on conifers.
little
economic importance except in
its secial
336
Key
to Genera of Coleosporiaceae.
1.
Basidiospore spindle-shaped Basidiospore ellipsoid Teliospores in a single layer

Teliospore layer strongly arched, forming minute knobs Teliospore in a flat crust Teliospores in a double
Ochropsora,
p. 336.
2.
3.
Mikronegeria. Coleosporium,
Chrysopsora.
Trichopsora.
p. 336.
layer,
long4.
5.
stalked
Teliospores in a columnar mass
Ochropsora Dietel
Urediniospores solitary. Teliospores in a waxy crust, loosely united, originally 1 -celled, later 4-celled, each cell bearing a single basidiospore on
III.
II.
a simple sterigma. 0. sorbi (Oud.) Diet.

1.
iEciospores {=JE. leucospermum) on anemone.
II
and
III.
Urediniospores on Sorbus and Spirea.
Coleosporium Levielle
0. Pj^cnia flattish, linear, dehiscent
filaments.
I
by a
slit,
without ostiolar
(=Peridermium). vEcia erumpent, definite. Peridium colorwith verrucosa walls. Spores globose to oblong, with colorless walls, the outer part formed of densely packed, deciduous tubercles.
less
II.
Uredinia
erumpent,
definite,
Avithout
peridium.
Spores
catenulate, globoid to oblong, pulverulent; wall colorless, closely verrucose, pores obscure.
III. Telia indehiscent
what
indefinite,
except through weathering, waxy, someusually roundish. Spores sessile, 1-celled (by
;
early division of the contents appearing 4-celled) colorless, thickened and gelatinous at apex.
wall smooth,
The genus is usually heteroecious. Arthur lists some twentyfour species for America. There are many biologic forms, morphologically indistinguish-
THE FUNGI WHICH CAU8E PLANT DISEASE

The aecial stage is found able yet not inter-inoculable. leaves of conifers, the telia on a large variety of hosts. C. ipomoeae (Schw.) Burr.
and
II.
I.
337
on
Unknown.
hypophyllous,
Uredinia
widely
scattered
or
somewhat
clustered, 0.25-1
mm.
across, early naked, orange-yellow fading to
white, ruptured epidermis usually inconspicuous; spores ellipsoid, 13-21 X 18-27 fi, more or less angular and irregular; wall thin,
1-1.5
III.
^l,
often confluent, pulor less across, deep reddish-orange fading to paleyellow; spores with wall swelling 20^0 jx above; contents orangeyellow fading to colorless, oblong, or slightly clavate, 19-23 x
vinate, 0.5
closely and noticeably vefrucose. TeUa hypophyllous, widely scattered,
mm.
rounded or obtuse at both ends. on various Ipomoeas and their kin among them morn^^^ ing glory and sweet potato.
60-80
/i,
Common
C. solidaginis (Schw.) Thiim."' 0. Pycnia amphigenous, scattered, numerous, originating between mesophyll and cortical layer, noticeable, 0.3-0.5 mm. wide
'''"'"
by 0.5-0.8 mm. long, dehiscent by a longitudinal slit, low-conoidal, 80-100 M high. 1 ( = Peridermium acicolum). Êcia from a limited mycelium, amphigenous, numerous, scattered on discolored spots occupying part of a leaf, erumpent from longislits, tongue-shaped, 0.5-1 mm. long by 0.5-0.7 mm. high; peridium
tudinal
rupturing irregularly, moderately firm,

white, cells overlapping, 35-45 ^l long, not much narrower, walls transversely
striate, inner coarsely verrucose, thick,
outer less rough and somewhat 11on or thmner; spores ellipsoid, 20-25 x 28-40 m; wall colorless, closely and
5-6
IX,
FiG. 250.
Stages
O. and
Coieosporium
I. of solidaginis
'
"
rdi'*''vf teT Wnton^
coarsely verrucose with deciduous tubercles which are directed away from a
up one
side,
thick,
smooth spot extending 2-3 n on the smooth spot, increasing to
5-6 n on the opposite side, including the tubercles. II. Uredinia hypophyllous, rarely also epiphyllous, irregularly
338
somewhat gregarious and crowded, 0.3-0.5 mm. soon naked, yellow or orange-yellow, ruptured epidermis across, inconspicuous; spores ellipsoid or globoid, 17-22 by 20-30 /x; wall rather thin, 1-2 n, closely and strongly verrucose; contents
scattered, or at first
orange-yellow
III. Telia
when
fresh, fading to colorless.
irregularly or sometimes elevated, 0.3-0.5 mm. across, reddish-orange; spores with wall swelling 30-40 n thick above; contents orange-yellow fading to colorless, terete, 15-23 x 55-
hypophyllous,
scattered
crowded and confluent,
slightly
rounded or obtuse at both ends; basidiospores globoid or about 12 x 18 /i, orange-yellow. I. ^cia on Pinus rigida. II and III. Uredinia and telia on Aster, Solidago and cultivated aster (Callistephenis) widespread and common. The connection between the stages was demonstrated by inoculations by
80
M,
elliptical,
CHnton.196.
197
C. senecionis (Schum.) Fries.
0. Pycnia amphigenous, scattered, numerous, originating between mesophyll and cortical layer, noticeable, 0.2-0.3 mm. wide,
0.5-1
slit,
mm.
long, dehiscent
by a
longitudinal
70-100 M high.
.
I JEc'm (=Peridermium oblongisporium) from a limited mycelium, amphigenous, bullate,

1
tongue-shaped,
1-2
mm.
long,
0.7ir-
mm.
high, whitish; peridium rupturing

of
regularly, fragile, white, cells overlapping, outer
and inner walls

smooth,
Fig. 251.
same
thickness,
3^
n,
outer
spores wall
inner moderately
ellipsoid,
' '
verrucose;
num senecionis, showing germination

of
Coieospoowng
.
broadly
,
17-24
u,
.
by 28-36
n,
colorless, thick,
3-4
densely verrucose with
teliospores.
prominent elongate
jj^
papillae.
Uredinia hypophyllous, thickly scattered,
about 0.5
mm.
across; early naked, bright orange-yellow fading
to pale-yellow, ruptured epidermis evident; spores ellipticalgloboid or obovate-globoid, 17-21 by 20-27 ix; wall thin, 11.5 evenly but not densely verrucose, with low papillae.
fji,
III. Telia hypo-phyllous, scattered, often confluent, small, 0.3
mm.
across, brilliant orange-yellow fading to pale orange-yellow;
339
spores with wall swelling 15-25 /x thick above; contents orangeyellow fading to pale-yellow, clavate or clavate-oblong, 16-20 by
60-83
I.
fi,
rounded at both ends or narrowed below.

sylvestris.
telia
^cia on Pinus
and
III.
is
II
Uredinia and
reported
same fungus
also
on Senecio. What may be this on cultivated Cineraria. ^^^ The
teliospores hibernate in their dark-red sori producing promycelia The sporidia bring about spring infection of the in the spring.
The connection
/-,
pine leaves and young twigs, later resulting in pycnia and aecia. of the forms was established by Wolff in 1872.
C. pini Gall. 0. Pycnia unknown, probably wanting. III. Telia amphigenous, on yellow spots, usually near the tips of the leaves, long covered by the epidermis, 1-5 mm. long, or when confluent up to 10 mm. or more, reddish-orange fading to
..--,,,
49, 199
ruptured epidermis inconspicuous; ju above, and soon disappearing upon exposure; contents orange-yellow fading to nearly colorless, clavate, slender, 13-20 by 60-100 ix, acute or rounded
pale-yellow
dirty- white,
or
teliospores with
walls swelling 30-50
below, sides wavy or irregular. as the type of a distmct genus, is set apart by Arthur based on the absence of spore forms other than the Gallowaya,
above, This
much narrowed
teliospores.
It causes serious leaf loss
on Pinus virginiana.
C. campanulae (Pers.) Lev.-^*^ 0. Pycnia amphigenous, scattered, numerous, originating between mesophyll and cortical layer, noticeable, large, 0.2-0.4 mm.
wide, 1-2
high.
rostrupi). Mcia from a limited mycelium, amphigenous, scattered, 1-3 on discolored spots, buUate, tongueI
(
mm.
long, dehiscent
by a longitudinal
slit,
90-110
fx
= Peridermium
shaped, large, 1-3
mm.
long, 0.7-1.5
mm.
high, yellow, fading to
white; peridium rupturing irregularly, fragile, white, cells overlapping, outer and inner walls same thickness, about 4-6 /x, outer
smooth, inner moderately verrucose; spores broadly ellipsoid or globoid, 17-22 by 22-31 n; wall colorless, thin, 2-3.5 /x, densely
verrucose, with prominent, elongate papillae. II. Uredinia hypophyllous, scattered, often confluent, 0.5-1
mm.
340
across,
soon naked, orange-red fading to white, ruptured epidermis evident; spores ellipsoid, 18-23 by 20-30 n; wall thin, 1-1.5 n, densely verrucose, with prominent, elongate papillae.
III. Telia
hypophyllous, scattered, often confluent, small, 0.2-
0.5
mm. across, slightly elevated, blood-red, fading to pale brownish-
yellow; spores with wall swelling 15-25 fi thick above; contents orange-red fading to nearly colorless, cylindrical or clavate-oblong, 17-24 by 55-85 ii, rounded or obtuse at each end.
and I on Pinus rigida. and III on Campanula and kin. There are numerous other species of
II
less
importance.
Melampsoraceae
Telia forming a
(p.
335)
more
or less definite crust or column; teliospores
compacted
sessile;
laterally into layers or rarely solitary in the tissues,
The family is
do
wall firm or rarely with a gelatinous layer. of little importance. Its uredinial
and
telial
stages
slight injury
on poplars and willows.
Key
Telia indehiscent.
Sori
all
to Genera of Melampsoraceae. com-
subcuticular;
teliospores
pacted in dense layers to form a crust; aecia when present without a perid-
ium; uredinia when present without a peridium or with an imperfect one of paraphyses Teliospores in a single layer; uredinia with spores and paraphyses intermixed
Teliospores in
1.
Melampsora,
p. 342.
more than one layer

2.
Uredinia with peripheral paraphyses only Uredinia without paraphyses Pycnia subcuticular, other sori subepidermal, or the telia within the epider-
Physopella, p. 345.
3.
Bubakia.
mal
cells
or between the mesophyll
cells;
uredinia
when
present with a
peridium

Teliosporcs approximating in a single layer within or beneath the epider-
341
mis;
urediniosporcs
globoid
to
oblong Walls of
the
teliospores
colored
Urediniosporcs echinulate throughout Urediniosporcs echinulate except at the apex
4.
Pucciniastrum,
p. 346.
5.
Melampsoridium,p.347.
Melampsorella, Hyalopsora.
Uredinopsis.
p. 348.
Walls
of
the
teliospores
colorless
0.
Urediniosporcs echinulate Urediniosporcs vcrrucose

Teliosporcs solitary within the mcsophyll; urediniosporcs pointed ....
Telia erumpent, sori
all
7.
8.
subepidermal
laterally;
secia
Teliospores
compacted
when present with

ium,
rupturing
flattened pcrid-
apically;
urcdinia
when present with a dehcate peridium and catenulate spores With all spore forms in life cycle With tclia and pycnia only
Proniycelium of the ordinary type. Promycelial cells changing directly
9.
Melampsoropsis,
p. '349.
10.
Chrysomyxa,
p. 350.
to basidiosporcs 11. Barclay ella. Teliospores often adhering and extruded

in long columns; secia when present with inflated pcridium, dehiscence
circumscissile; urcdinia
when
present,
wdth pcridium, spores borne singly on

pedicels.
Teliosporcs 1-ccllcd Telia naked

Telia forming columns Teliospores firmly united side-
wise and endwise
12.
Cronartium,
p. 350.
Teliosporcs loosely united laterally, separating in disks .... 13. Alveolaria.

Telia not extruded
Wall brownish, thick Wall colorless, thin
14.
Baeodromus.
Cerotelium.
15.
342

Wall
slightly colored
16.
Cionothrix.
Telia with a peridium Telia half projecting
above the
17.
host surface
Telia sunken in the tissue of the
Dietelia.
host
Teliospores 2-celled
18.
Endophyllum,
Pucciniosita.
p. 353.
Peridium present Peridium none
19.
20.
Didymopsora.
Melampsora Castaigne
0. Pycnia half spherical.
1.
(p.
340)
^cia
of cseoma-type,
no peridium or paraphyses.
II.
Urediniospores solitary,
membrane
III.
colorless.
1-celled,
Teliospores
in
flat
more,
crusts.
irregularly
rarely limited
The
zation
Basidiospores spherical. question of biologic specialiis
this genus.
especially complicated in The uredinial and telial
stages occur in abundance on wil-
lows and poplars, the
secial
stage on
a wide range of plants embracing
gymnosperms, monocotyledons and

dicotyledons.
M.
lini
D. C.
0. Pycnia amphigenous, numerous, scattered, inconspicuous, subflattened pale-yellow, or lens-shaped, 100-175 n globoid in diameter, 65-95 /x high; spores
epidermal,
Fig. 252.
Pucciuiastrum goeppertianum, showing germinating ellipsoid,

teliospores.
2-3 by 3-4
/x.
Êcia chiefly hypophyllous, nu0.2-0.4 mm. across, bright orangemerous, scattered, rounded,
1.
After Hartig.
and mesophyll, soon naked, ruptured epidermis evident; spores globoid, 19-27 x 21-28 fi.; wall colorless, thin, about 1 n, finely and evenly verrucose, with distinct papillae, pores not evident.
yellow, conspicuous, formed between epidermis
343
II. Uredinia amphigenous and eaulicolous, scattered or somewhat gregarious, often crowded, round or on stems elongate,
0.3-0.5
mm.
across, soon naked, reddish-yellow fading to nearly
white, pulverulent, ruptured epidermis noticeable; spores broadly elliptical or obovate, 13-18 x 15-25 ix, wall colorless, rather thin,
2 n, evenly and finely verrucose, with low papillae, pores equatorial, obscure; paraphyses intermixed with the spores, capitate,
large,
5-22 x 40-05
/x,
smooth, wall thick.
amphigenous and caulicolous, scattered, often conround or elongate, 0.2-0.5 mm. across, slightly elevated, reddish-brown becoming blackish; spores subepidermal, appressed into a single layer, prismatic, 1-celled, 10-20 x 42-50 n; wall brown, smooth, thin, about 1 yu, not thickened above. Autoecious on flax. Sometimes very injurious.-''^ M. medusae Thiim.
III. Telia
fluent,
().
Pycnia chiefly epiphyllous, scattered or somewhat gregarious,

punctiform,
pale-yellow,
minute,
inconspicuous,
subcuticular,
hemispherical, 40-80 ii in diameter, half as high. I. iEcia chiefly hypophyllous, scattered or somewhat gregarious, small, 0.1-0.3 mm. broad, round or oblong, pale-yellow
fading to white, inconspicuous, formed between epidermis and mesophyll, soon naked, pulverulent, ruptured epidermis noticeable; aeciospores globoid, 17-22 by 17-24 /x; wall colorless, thick,
2.5-3
n,
minutely verrucose, with minute crowded
papillae,
pores
indistinct.
II.
Uredinia amphigenous, or only hypophyllous, scattered,
somewhat pulto pale brownish-yellow, ruptured verulent, orange-yelfow, fading epidermis usually inconspicuous; urediniospores ellipsoid or
roundish, small, 0.2-0.4
across, early naked,
mm.
by 22-30 /x, usually flattened laterally; up to 10 /x on the flattened sides, sparsely and evenly verrucose, with fine papilla3, except on the flattened sides which are smooth; paraphyses numerous, intermixed with the spores, capitate, smooth, 40-65 n long, head 14-25 ii broad, wall thick, 3-6 ix, peripheral paraphyses thirmer-walled and more
obovate-ellipsoid, 15-18 wall colorless, 2.5-3 ^ or
clavate.
III.
Telia
amphigenous or only hypophyllous, scattered or

irregularly
somewhat
confluent,
roundish,
small,
0.2-0.4
mm.
344
across,
slightly elevated, light reddish-brown, becoming deep chocolate-brown, subepidermal; teliospores prismatic, 12-15 by 30-45 î; wall smooth, cinnamon-brown, uniformly thin, 1 fx.
and
on Larix,
II
and
III
on Populus.
Common
on
all
species of Populus and often doing serious defoliation of the trees.
damage by
its
early
M.
Salix
all
bigelowii Thiim. with O and I on Larix and II and III on It occurs on practically quite similar to the preceding. of willow. species
is
Other species not found in America are:
M.
1
alUi-fragUis Kleb.^'i
on Allium vineale and A. sativum. II and III on Sahx.

allii-salicis albae Kleb.^^^
M.
I
on Allium. II and III on Willow.
M.
I
allii-populina Kleb.^^^
on Allium. II and III on Populus. M. klebahni Bub. I on Corydalis. II and III on Populus.
M.
I
larici-pentandrae Kleb.^^^
Salix.
on Larix. II and III on
M.
I
larici
populina Kleb.^^^
on Larix. II and III on Populus.

pinitorqua Rost.^^^
(
M.
I
=Cseoma pinitorquum).
to
structive
leaves.
pine seedlings.
The Cseoma-stage is quite deThe teliospores grow on Populus
M.
I
repentis Plow.
orchidis); on Orchis. ribesii-viminalis Kleb.^^^
II II
(=Cseoma
on Ribes.
and Illon
Salix.
M.
I
and III on
Salix.
M.
I
rostrupii Wagn.^^^ on Mercurialis. II and III on Populus.

M. saxifragarum
I
345
(D. C.) Schr.
and
111
on Saxifrages,
Physopella Arthur
(p.
340)
telia recognized,
Cycle of development imperfectly known; only urodinia and both subepidermal. Uredinia erumpent, definite,
roundish, pulverulent, encircled by more or less clavate paraphyses which are often united at their bases, or wholly, into a pseudo-
peridium opening by a central pore. Urediniospores borne singly on pedicels, obovate-globoid or ellipsoid; wall pale-yellow, echinulate or rarely verrucose, pores obscure. Telia indehiscent, forming lenticular masses, two or spores 1-celled; walls smooth.
more
cells
thick at center.
Telio-
P. vitis (Thum.) Arth." II. Uredinia hypophyllous, scattered thickly over wide areas,
mm. or less across, soon naked, arising between epidermis and mesophyll, surrounded by numerous incurved paround, minute, 0.1
raphyses, pulverulent, pale-yellow, fading to dirty white, ruptured epidermis inconspicuous; urediniospores broadly ellipsoid or obovate, 13-17
by 18-27 n; wall nearly colorless, thin, 1 /x, minutely and rather closely echinulate, pores obscure; paraphyses hyphoid, curved and irregular, 6-10 n thick, 30-60 n long, wall uniformly
thin, 1
/i,
yellowish.
hypophyllous, scattered thickly over large areas, roundish, minute, 0.1-0.2 mm. across, indehiscent, 3 to 4-cells thick; teliospores ovoid, 12-15 by 20-30 n, wall smooth, nearly
III. Telia
colorless, thin, 1
/x
or
less.
On
grape leaves in Southern United States
and West
Indies.
Also in South America and Japan. P. fici (Cast.) Arth."

II.
Uredinia hypophyllous, scattered thickly over large areas,
roundish, usually small, 0.1-0.3

arising
mm.
across, or rarely larger, bullate,
between epidermis and mesophyll, tardily dehiscent by central rupture, encircled by delicate, evanescent paraphj^ses, pulverulent, pale cinnamon-brown, ruptured epidermis overarching or wall pale-yellow, erect; spores obovate-globoid, 14-20 by 18-27 thin, 1-1.5 M, sharply and rather sparsely echinulate, pores ob/jl;
346

m
scure; paraphyses hyphoid, very delicate, collapsing, 60-80 long, wall colorless, very thin, slightly thickened at apex, 1 mIII.
II.
Teha, unknown.
On
fig
and osage orange.

Pucciniastrum Otth.
(p.
341)
secia,
Heteroecious.
The
uredinia and
telia,
cycle of development includes pycnia, with distinct alternating phases.
0. Pycnia subcuticular, low-conoidal, without ostiolar filaments. Peridium delicate, verrucose 1. Mcia, erumpent, cyhndrical.
on inner surface. Spores is thinner and smooth.

II.
ellipsoid,
verrucose except one side which
Uredinia barely protruding through the epidermis, dehisPeridium hemispherical, delicate, cells cent by a central pore.
orifice. Spores borne singly on pedicels, obovate to wall colorless, echinulate, pores indistinct. ellipsoid; III. Telia indehiscent, forming more or less evident layers in
longer at
the epidermal
cells or
immediately beneath the epidermis.
Spores
oblong or prismatic, 2 to 4-celled planes; wall smooth, colored.
by
vertical partitions in
two
Arthur*'
tant.
lists
nine American species but none are very impor-
P. hydrangeae (B.
&
C.) Arth.
O
II.
and
I.
Unknown.
Uredinia hypophyllous, scattered, round, small, 0.1-0.2 mm. across, dark-yellow fading to pale-yellow, ruptured epidermis inconspicuous, dehiscent by a central pore; peridium hemispherical, delicate, cells
small, cuboid, walls uniformly thin,
1-1.5 n,
ii, barely pointed, walls thin, smooth; spores broadly elliptical or obovate, 12-18 X 16-24 n; wall nearly colorless, thin, 1-1.5 m, sparsely and
ostiolar cells slightly or not elongate,
10-16
strongly echinulate. III. Telia amphigenous,
or
chiefly
confluent into small angular groups,

raised, reddish-brown; spores
epiphyllous, 0.3-0.8 mm.
effused,
or
across,
not
forming a single layer within the or sometimes between the epidermis and mesophyll, globoid, 22-28 x 24-28 /x, wall dark cinnamon-brown, uniformly thin, 1.5-2 fx.
epidermal
cells,

It
is
347
which
I.
it
found in the uredinial and -"may be quite serious.

(J.
telial
stages on
Hydrangea on
P. goeppertianum
(
Kiihn.) Kleb.
Ill on Vaccinium.
It
=^-E.
columnare) on Abies leaves.

stage
is
The
aecial
the destructive form.
has been found
Fig. 253.
Melampsoridium, section through germinating

telium.
After Tulasne.
^'
but a few times in America, "^'

Fig. 252. P. pustulatum (Pers.) Diet.
lobii.)
while the
telial
stage
is
common.
(=P. abieti-chamaenerii, P. epi-
and
1
on Abies.
II
and
III
on Epilobium.
P. padi (Kze.
(
&
Schm.) Diet.
II
= ^. strobilinum) on fir. and III on Prunus padus.

is
P. myrtilli (Schm.) Arth.
found
in the uredinial
and
telial
stages on various Vacciniums.
Melampsoridium Klebahn
(p.
341)
0. Pycnia flattened-conoidal, without ostiolar filaments.

1.
JEc'ia
erumpent, subcylindrical.
Peridium regularly dehis-
Spores ellipsoid to globoid; wall colorless, thin, verrucose except one side which is thinner and smooth. Peridium firm, dehiscent II. Uredinia somewhat erumpent.
cent, cells rhomboidal.
by central pore;
peridial cells isodiametric, those of orifice pro-
348
longed into sharp points. Spores borne singly on pedicels, ellipsoid; wall colorless, echinulate, pores indistinct. III. Telia indehiscent, forming evident layers immediately beneath the epidermis. Spores oblong or prismatic, 1-celled;
wall smooth, slightly colorjed. M. betulae (Schiim.) Arth. occurs, Betula.
and
on
larch, II
and III on
Melampsorella Schroter
(p.
341)
0. Pycnia hemispherical, without ostiolar filaments. 1. ^cia erumpent, definite, oblong, buUate. Peridium colorless,
with thin-walled
thin, verrucose,
II.
cells, ^ciospores without smooth spot.
ellipsoid; wall colorless,
Uredinia barely protruding through the epidermis, dehiscent
central pore. Peridium hemispherical, delicate, cells slightly or not enlarged at orifice. Urediniospores borne singly on pedicels,
by a
obovate to
scure.
III.
ellipsoid; wâll slightly colored, echinulate, pores ob-
Telia effused, indehiscent. Teliospores globoid to ellipsoid, 1-celled; wall smooth, colorless, thin.
M.
elatina (A.
& S.)
Arth.-O"*
0. Pycnia epiphyllous, few, scattered, punctiform, inconspicuous, subcuticular, not extending much into walls of epidermis,
depressed-hemispherical, small, 100-130 n broad, 40-50 // high. 1. ^cia from a perennial mycelium, dwarfing the young shoots,
and forming witches' brooms, hypophyllous, forming two irregular lines, deep-seated, wholly dropping out of the substratum
roundish or irregularly oblong, large, 0.5-1 mm. across, bladdery, soon open by falling away of the upper part; peridium colorless, dehiscence irregular, cells Avith thin inner and
at maturity,
outer walls; seciospores broadly ellipsoid, or nearly globoid, 1418 X lG-28 m; wall colorless, thin, 1-1.5 /x, closely and rather
finely verrucose.
II.
Uredinia
amphigenous, scattered
or
somewhat grouped,
orange-red when fresh, paleyellow when dry; peridium hemispherical, dehiscent by a small central orifice, cells elongate at sides, polygonal above, inner and
small, round, 0.1-0.4
across,
mm.
outer walls same thickness; urediniospores ellipsoid or obovoid,
THE FUNGI WHICH
CAUrfE
PLANT DISEASE
ju;
349
sparsely
12-18 X 16-30 m; walls pale-ycUow, rather thin, 1-1.5

echinulate with short conical points. III. Telia hypophyllous, on whitish
or pale reddish spots; teliosporcs within tlu; epidermal cells, 1-celled, short-cylindrical or polygonal, 13-20 m broad; wall colorless, smooth, thin.
I
(
=Peridermium elatinum) on
and
III
fir
causing swelling, cankers and
witches' brooms.
II
on various members
possess
significance,
of the pink family.
All
is
stages
perennating mycelium.
The
secial
stage
of
most economic
various sizes.
of the witches'
The
iccia
producing witches' brooms of are formed only on the deformed needles
brooms.
(p.
Melampsoropsis (Schroter) Arthur

Cj^cle of
341)
development includes pycnia, aecia, uredinia and telia, with distinct alternating phases; heteroecious. Pycnia and other
sori
subepidermal. 0. Pycnia deep-seated,

1.
somewhat erumpent,
and transversely
flask-shaped.
firm,
Mcia erumpent,
flattened laterally.
Peridium
outer
wall of cells greatly thickened
striate, inner wall
smooth.
II.
^-Eciospores ellipsoid to globoid; wall colorless, coarsely verrucose with deciduous tubercles.
Uredinia erumpent, pulverulent.
Peridium very
delicate,
Urediniospores catenulate, globoid to lanceolate; wall colorless, verrucose with somewhat deciduous tubercles, pores ol)scure.
evanescent,
III.
sometimes
wanting.
Telia erumpent, definite, roundish,
waxy becoming
vel-
vety.
Teliospores catenulate,
1-celied,
oblong or cuboid; wall
colorless, thin,
smooth.
pycnial and Picea excelsa.
M. rhododendri
secial
(D. C.) Arth. Uredinial and telial stages on Rhododendrons;

stages
(
= iEcidium abietinum) on
The pycnia appear on fir leaves in spring and about a month sec'ia. The Bciospores germinate upon the Rhododendron. The mycelium perennates in its evergreen leaves and
later the
produces the uredinial and

serves for dissemination.
leaves.
telial
stages, the
former of which
The
basidiospores infect the
young
fir
350
Chrysomyxa Unger
III. Teliospores
(p.
341)
formed
of
a series of superimposed cells, of which the lower are sterile,

or slightly elevated, orange or reddish,
forming
flat
waxy,
crusts.
Germina-
tion of the teliospore
by a
each
cell,
promycelium which produces mostly
from
Fig. 254.
Melampsoropsis rhododendri, section through telium. After De Bary.
four basidiospores. C. abietis (Wal.) Ung.

T'dlo ^^^^^
/->riKr
^^^^
'
Tf ^^
(r^r-r^^
^"^"^^
leaves
and
spots on spruce the basidiospores seem able to infect the same host.
yellow
European.
Cronartium Fries
(p.
341)
0. Pycnia deep-seated, broad and flat. (=Peridermium). iEcia erumpent, inflated.
Peridium
mem-
branous, rupturing at the sides rather than above, 2-4 cells thick, outer surface smooth, inner verrucose. Spores ellipsoid; wall
colorless,
coarsely verrucose with deciduous tubercles, except a
smooth spot on one side. II. Uredinia somewhat erumpent.
Peridium moderately
firm,
rupturing above, upper part evanescent; peridial cells isodiametric. Spores borne singly on pedicels, globoid to ellipsoid wall nearly or
;
quite colorless, echinulate, pores obscure. III. Telia erumpent, at first arising from the uredinia, the catenulate spores adhering to form a much extended, cylindrical or
filiform
column, horny when dry.
Spores oblong to fusiform,
1-celled; wall slightly colored, thin, smooth. Five American species are recognized by Arthur.
All
known aecial stages are Peridermiums on stems of conifers. ~'' C. ribicola F. de WaV'^--^'-
0. Pycnia caulicolous, scattered, honey-yellow, forming minute, bladdery swellings. Spores hyaline, ovoid to elliptical, 1.9-4.7 ti.

I
351
(=Pcridermium
strobi).
JEc'ia caulicolous,
causing fusiform
swellings of the stem, rounded to elongate; peridium inflated, rupSpores ellipsoid to ovoid, turing at sides, thick, membranous.
18-20 X 22-23
thick.
II.
m,
wall
colorless,
/i
elongate smooth spot, 2-2.5
thick,
coarsely verrucosc except on on smooth spot 3-3.5 n
Uredinia hypophyllous, thickly scattered in groups, round,
pustular, 0.1-0.3
mm., at
first
bright yellow; peridia delicate.
^^9
Fig. 255.
Cronartium.
A, uredinium;
J5,
telium.
After Tubeuf.
Spores ellipsoid to obovate, 14-22 x 19-3.5 2-3 ju thick, sparsely and sharply echinulate.
III.
/x,
wall
colorless,
Telial
columns hypophyllous,
cylindrical, 125-150
^ thick,
up to 2 mm.
ish;
long, curved, bright orange-yellow,
colorless,
spores oblong or cylindrical, smooth, rather thick, 2-3
becoming brown8-12 x 30-60 /z; wall nearly
/x.
Heteroecious O, I, on white pine, Pinus cembra and several other 5-leaved species; II and III on currant and gooseberry and
several other species of Ribes.
The telial stage was first noted in Geneva, N. Y., in 1900.-*^ The rust is now known in some nine states. It has been known in
Europe since 1854. Its though the telial stage
generic connections of
most serious in its aecial stage, abundant and conspicuous. The very the forms was proved by Klebahn in 1888
effects are
is
is
by inoculations. The mycelium
is
doubtfully perennial in Ribes and certainly
so in the bark of the pine.
352
C. comptoniaB Arth.
I
(=Peridermium pyriforme) on Pinus trunks.
Ill
on Comp-
tonia.
The Peridermium
it
is perennial in the trunks of the pine where does considerable injury. Clinton sowed aeciospores from pine
Fig.
256.
Cronartium
Pinus.
comptoniai (Peridermium) on After Clinton.
on Comptonia and
to appear.
^^^
in
about twelve days the uredinia began
C. quercus (Brond.) Schr. Heteroecious I (= Peridermium

oak.
Successful inoculations were
first
cerebrum)
on pine.
^^^
Ill
on
reported by Shear,
later
by
Arthur and Hfedgcock.^^'' formed on pine trees.
Globoid swellings 5-25 cm. across are
C. asclepiadeum (Wil.) Fries. Heteroecious I (= Peridermium

tris.
cornui)
on
Pinus
silves-
II
and
III on
Cynachum,
is
Paeonia, Gentiana
and several other

kills
hosts.
European.
perennial in pine twigs and gradually
The mycelium
them.
353
Endophyllum
Leviell^ (p. 342)
The
cycle of development includes only pycnia
and
telia,
both
subepidermal. 0. Pycnia deep-seated, somewhat erumpent, flask-shaped, with

ostiolar filaments.
III. Telia bullate, definite, round, pulverulent.
Peridium evan-
spores but flattened. Spores catenulate or without order, 1-celled, globoid to ellipseemingly compacted
escent, cells resemljling
soid; wall colored,
medium
thick, verrucose.
E. sempervivi (Alb.
Pycnial and telial lium perennial in the host.
Schw.) D. By. stages on species of Sempervivum.
&
Myce-
Pucciniaceae
Teliospores
(p.
335)
stalked (stalk sometimes short or evanescent) with several cells in a row or several united to form a parasol-like head on a compound stalk; separate or gelatinousembedded. Basidiospores formed from promycelia. Êcia with
1-celled or
or without peridia. Urediniospores solitary. This is the largest and most important family of the order, inlesting numerous valuable agricultural plants and causing enorspecies are manifold and the complexities owing to heteroecism and biologic specialization are very polymorphism,
loss.
mous
The
great.
Key to Genera of
Pucciniaceae
Teliospores united into a head on compound pedicles, or several sessile or stalked on
common simple pedicel; sori subcuticular or subepidermal; urodinia when

a
present without peridium or encircling
paraphyses. Teliospores united into a head on a com-
pound pedicel Teliospores free, 1-4 on a simple pedicel,

all
1.
Ravenelia,
but one lateral

2.
Teliospores flattened laterally
Dicheirinia.
354

Teliospores
flattened
above and be3.
low
Teliospores not flattened, but urediniospores flattened laterally Teliospores free, 2-8 at apex of a
stalk
Pileolaria.
4.
Hemileia,
p. 355.
common
5. 6.
With all spore forms With pycnia and teha only

Teliospores not borne on a or united into heads.
Tranzschelia,
Polythelis.
p. 356.
common
pedicel,
Tehospore wall with a more or dent gelatinous layer.

Teliospores
less evi-
with
evident
gelatinous
7.
layer, pores lateral
Phragmopyxis.
Uropyxis.
Tehospores 2-celled Teliospores with obscure

layer, pores apical.
8.
gelatinous
Teliospores with appendaged pedicels

9.
Prospodium.
Nephlyctis.
Tehospores
pedicels
without
appendaged
10.
Teliospore wall without gelatinous layer Pycnia subcuticular, other sori sub-
epidermal secia when present without peridium; uredinia when present without peridium, but usually
;
with encircling paraphyses. Teliospores mostly tuberculate, the

pores more than one Teliospores 1-celled Teliospores with 3 or
and
lateral
11.
Trachyspora.
more
cells
clustered at the apex of the
pedicel
12.
.
Tehospores more than 3-celled Teliospores with more than three

cells hneally arranged Teliospores mostly smooth, the pores
13.
Triphragmium, p. 358. Sphaerophragmium.
14.
Phragmidium,
p. 358.
one in a
cell
and
apical.
15.
Spirechina.

Tcliospores 2-cclled Teliosporos 3 or more-celled. Without uredinia
IG.
355
Gymnoconia,
p. 359.
17.
18.
Without
Sori
all
secia
Xenodochus, p. 361. Kuehneola, p. 361.
subopiderinal; secia
when
present with a peridiuni; uredinia when present with no
peridium or rarely with encircling paraphyses.
Teliospores embedded in a more or less gelatinous matrix Teliospores not embedded in a

. .
19.
Gymnosporangium,
p. 361.
gelatinous matrix. Teliospores colorless Teliospores colored
20.
Eriosporangium.
Teliospores 1-celled Teliospores 2-celled
21.
Uromyces,
p. 371.
22. Puccinia, p. 375.
Hemileia Berkley
& Brown
(p.
354)
Cycle of development imperfectly knowTi; only uredinia and both subepidermal. II. Uredinia formed beneath the stomata, erumpent, without peridium or paraphyses, spores borne singly on short pedicels,
telia recognized,
arise from a protruding hymenium of agglutinated hyphse, obovate, laterally flattened and dorsiventral wall pale-yellow, smooth on ventral side, papillose on dorsal side, pores obscure or absent.
which
III. Telia replacing

cels,
the uredinia.
Spores borne singly on pedi"^
1-celled, napiform; wall nearly or
quite colorless, smooth. H. vastatrix Berk. & Br.

II.
Hypophyllous,
thickly
scattered,
xbrtSK]OL^
or rarely
somewhat
circinate,
very small,
projecting
i
^^p^
trix,
.i,
about 0.1
to
,
mm.
across, light-orange fading ^^^
.57.-Hemileia vasta-
pale-yellow,
.
pulverulent,
1
1
through stomata and rarely rupturmg
trliosporc.
uredinium; c, After Diet el
and Ward.
the epidermis; spores bilateral, slightly obovate, flattened on the ventral side, 20-28 by 30-40 n; wall pale-yellow, 1-1.5 /x thick, rather thickly and very coarsely papil-
358
lose

on dorsal side with
IX
Ijluntly
pointed tubercles 2-4 n long,
smooth, pores obscure. III. Hypophyllous, arising from uredinia, thickly scattered, very small, about 0.1 mm. across, pale-yellow; sjôres napiform
or globoid,
1-1.5
in diameter, ventral side
ingly
somewhat umbonate above; wall pale-yellow or seemcolorless, thin, 1 /x, slightly if any thicker above, smooth;
and
is re-
pedicel h^âline, one-half to once length of spore, slender. It constitutes a serious coffee parasite in the orient
ported also from Porto Rico. H. woodii K. & C. is a serious coffee parasite and occurs also on Vanguieria edulis. H. oncidii Clriff & Maub. is on cultivated Oncidiums in France.
Tranzschelia Arthur
(p.
354)
Cycle of development includes pycnia, vec'ia, in-edinia and telia, with alternating phases; autoccious or heteroecious. Pycnia subcuticular, other sori subepidermal.
0. 1. Pycnia depressed-conical or hemispherical; hymenium flat. Êcia erumpent, cylindrical, Peridium dehiscent at apex, becoming recurved. Êciospores globoid; wall colored, finely verrucose.
Uredinia erumpent, definite, without peridium. Uredinioborne singly on pedicels, with paraphyses intermixed, obovoid, somewhat narrowed at both ends; wall colored, usually
II.
spores
paler below, echinulate; pores equatorial. III. Telia erumpent, definite, pulverulent, without peridium. Teliospores forming heads or balls by being attached by short,
fragile pedicels to a
common
stalk,
which
is
short and incon-
spicuous, 2-celled by transverse septum, cells rounded falling apart, wall colored, verrucose.
and
easily
T. punctata (Pers.) Arth.^''

I
-"'' '-^^' -^3
(=^cidium punctatum).
whole
of
Peridia uniformly scattered over
foliage, hypophyllous, flat, semi-immersed, with torn yellowish edges. Spores subglobose, pale yellowishbrown, 15-24 n in diameter. Pycnia scattered, blackish, puncti-
the
the
form.
II.
often
Uredinia light-brown, small, round, crowded, pulverulent, confluent. Spores ovate or subpyriform, apex darker,
357
thickened, l)luntly conical, closely echinulate, brown, 20-35 x 12-1 G n, mixed with numerous capitate brownish paraphyses.
Telia pulverulent, dark-l)rown, almost black. Spores consisting of two spherical cells, flattened at their point of union, the lower cell often being smaller and paler. Epispore uniformly
III.
I'lG.
258.
piuii-tatu,
un.'(_liiii(ASi;urrri.
Alter llolwux
studded with short stout spines. 30-45 x 17-25 yu Pedicels short, colorless. Spores and I on Hepatica and Anemone. Heteroecious II and III on Prunus sps., peach, almond, plum, cherry, aprithick, chestnut-brown, thickly
:
cot.
Widely distributed in North America, Europe and Asia and apparently introduced into Australia about 1883. The jEcial stage is perennial. Urediniospores have also been shown to remain viable
over winter.
The
this
peculiar character of the urediniospores has
sometimes led
fungus to be mistaken for a Uromyces. In 1904, Tranzschel -^^ made cultures of the a^cial stage from Anemone on various Prunaceous hosts. Arthur made similar
inoculation from Hepatica in 1906.
358
Triphragrnium Link
Teliospores 3-celled, one basal,
or
tvvô
(p.
354)
apical,
each
cell
with one
more germ
T. ulmariae
tubes.
Schm. occurs on Ulmaria
in
England and at one
station in America.
Phragmidium Link
0. Pycnia present.
1.
(p.
354)
iEciospores in basipetal chains.
The
first
two spore forms
Fig.
260. Phragmidium bulbosum,
Fig. 259.
Triphragrnium
ulniaii^f, g.i rui-
teliospore germinating. After Tulasne.
nating teliospore.
After Tulasno.
are in pulverulent sori, surrounded
by clavate or
capitate, hyaline
paraphyses.
IL Urediniospores
single.
in. Teliospores separate, pedicellate, consisting of from three to ten superimposed cells, the uppermost of which has a single
apical
germ
iBcial
The
pore, the others about four each, placed laterally. stage is a Cseoma but with a border of incurved pa-
'HIE
FUXGi WHICH cau.se plant disease
359
raphyses. The unicellular urediniospores are similarly surrounded, and bear numerous germ pores. The genus is limited to Rosaceous hosts and its species are autoecious. ^^ on roses as Eight American forms are recognized by Arthur
follows:
P.
montivagum
Arth., P. discifiorum (Tode) James, P. ameri-
Fio. 261.
Urcdinio- and tfjliosporcs uf; 1. P. americanum; 2. P. rosie-setigera;; 3. P. rosae calif ornicse; 4. P. rosa.'-arkansanae; 5. P. montivagum; 6. P. disciflorum. After Arthur.
canum Diet., P. rosae-setigerae Diei., P. rosae-californicae Diet., P. rosae-arkansanae Diet., P. subcorticinum (Schr.) Went, and P. rosae-acicularis Sire.
They
are mostly on wild roses
and
is
of but little
economic im-
portance. P. violaceum (Schul.) Went,
often serious on
Rubus
in
Europe.
P. rubi-idaei (Pens.) Wint. is found on raspberries in Europe. P. speciosum Fr. on rose has been separated by Arthur as
Earlea speciosa on account of

cel, its
its
large
compact caulicolous
telia
non-gelatinous teliospore pediand the absence of uredinia.

355)
Gymnoconia Lagerheim
0. Pycnia conic.
I
(
(p.
=Ca?oma), peridia and paraphyses none.

is
Spores as in Puccinia. This genus bears a superficial resemblance to Puccinia but

III.
easily distinguished
by
its
naked
secial sori.
360 G.
I

interstitialis (Schl.)
Lag.
272-274. 211
0. Pycnia glandular,
numerous mostly epiphyllous.
hypophyllous, sori irregular, confluent; spores orange-red, globose to elliptic, epispore thin, 18-35 x 12-24.
nitens).
III. Telia
(=Caeoma
more or
less angular,
hypophyllous, few, sparse, cinnamon-brown; spores 36-45 x 22-27 n, pedicel short or wanting.
in United States,
Autoecious, on raspberries and blackberries, wild and cultivated, Canada, Europe and Asia.
The pycnial stage appears first in spring giving to the leaves and stems a glandular appearance. About two or three weeks
Fig. 202.
-G.
interstitialis,
cseoma sorus.
After
Newcomb.
later the ajcial stage is visible
on the lower surface of the leaves;
the epidermis soon ruptures and the orange beds of spores show.
The pycnia
are then fully developed. The affected plants are stunted and are unproductive but are not killed. The fungous mycelium is intercellular, growing rapidly into formative
much
tissues
and perennating
-^^
in the
woody
shoots.
The
knob-like
and often lie against the nuclei. The is especially abundant in the pith near the bundles. mycelium The fficiospores may germinate at once and infect susceptible hosts. The teliospore which is less conspicuous and therefore
haustoria penetrate the
cells
rarely seen
is
of the
Puccinia type.
The
telia
appear in July

and August, usually hypoi>hyllous, and the and inconspicuous.
Artificial infection of
sori are
361
very small
stage
Rubus with the
spores of the
Caoma
^^^ gave rise to the telial form, demonstrating by Tranzschel the identity of the two. Cultures were also made by Clinton
about the same time.
Xenodochus Schlecht
iEciospores
catenulate;
(p.
355)
teliospores
uredinia wanting;
short-
pedicelled, several celled in linear arrangement.
X. carbonarium Schl., autoecious on Sanguisorba in Europe.
Kuehneola Magnus
(p.
355)
iEcia wanting; uredinia pulvinate, telia similar to Phragmidium but with smooth spores with the germ pores apical. K. uredinis (Lk.) Arth.
II. (=Uredo muelleri.) Uredinia lemon-yellow, minute dots; spores globose to elliptic, about 26 //, hyaline, slightly verrucose. III. Telia solitary, pale, 250-500 fx broad; spores 5 to 6 to
12-celled, epispore hyaline, cells
17-47 x 15-26 n; basidiospores
8.5-9.5
fx.
The
t'.iem
telia are pale yellowish-white, thus readily distinguishing
from other Rubus rusts. The uredinia are common and sometimes The sori are small and scattered.
injurious
on Rubus.
in British
K. gossypii (Lagerh.) Arth. is reported on cotton Guiana,-^^ also Florida, Cuba and Porto Rico.
Gymnosporangium Hedwig
f/-06-2ii.
213. 217
(p
secia
355)
telia,
Cycle of development including pycnia,

distinct alternating phases; heteroecious
and
with
and
autoecious.
Pycnia
and other
sori
subepidermal.
O. Pycnia deep-seated, usually globoid, generally prominent and conspicuous, at first honey yellow, usually becoming blackish,
globose or flattened-globose, with ostiolar filaments.
I
(
=Rocstelia) erumpent, at
first
cylindric.
Peridium dingy
white, usually elongated into a tubular form,
membranous, tending
362
to rupture by longitudinal slits along the sides; peridial cells imbricate and often articulated, occasionally hygroscopic, outer walls smooth, rather thin, inner walls smooth, verruculose, verrucosa,
rugose, or spinulose.
nate barren
cells,
Êciospores in basipetal chains with alterenclosed in a peridium, globoid to broadly ellip-
soid; wall colored, verrucose, usually with
numerous, scattered,
evident germ pores. III. Telia erumpent, naked, usually definite, variously shaped, gelatinous and elastic at maturity, expanding considerably when moistened. Teliospores chiefly 2-celled, in some species 3, 4, or
Fig. 263.
Gymnosporangium, spore masses just emerging.

After Heald.
by transverse septa; walls colored, of various smooth; pores usually two in each cell, sometimes, 1, 3,
5-celled,
thickness, or 4, vari-
ously arranged; pedicels hyaline, elastic, usually of considerable length, cylindric, rarely carotiform, walls thick, the outer portion
and becoming gelatinous to form a which the spores appear embedded.

swelling
jelly-like
matryx
in
All of the species agree in possessing the same spore forms, pycnia, aecia, and telia which appear in the same sequence in the different species; also, in the fact with two exceptions, that
the secia grow on pomaceous plants and the telia on Juniperus

(with few exceptions).

The
a^ciospores are
363
borne
in
acia which rest in orange or
Pycnia abound. The yellow spots often strongly thickened. with its thick peridium is erumpent and projects to aecium
some distance above

the host surface, this character giving rise
to the separate form-
genus, Ra'stelia.
peridial
The
whicii
margin
is
may
be
lacerate
or
in
fimbriate
specific
tion.
used
characteriza-
The
spores are
secia.
borne and function as

in
ordinary
They bear
pores.
several germ
iEciospores nate at once
germi-
and
if
they
fall
upon suitable
coniferous hosts bring
about infection.
the
often
leaf
The
mycelium penetrating
or
branch
large
induces
hypertrophy. In spring in moist
weather
spores
are
j
the
telio-
spore
posecl
found in masses com,
01
the
are
spores,
which
USUallv
Fig. 264. Gymnosporangium teliosporcs. a, G. clavariseforme; h, G. glohosuni; c, G. nuicropus; d, G. iiidus-avis; c, G. nelsoni;/, G. clavipcs. After King.
and of their long gelatinous pedicels. usually bears several germ pores near the through one of which the tube emerges. The teliospores germinate immediately in situ by
orange or yellow,
Each
cell
septum
typical
4-celled promycelia
and four basidiospores are produced on each
promycelium.
364
The basidiospores are capable of infecting only the appropriate alternate host and that when the parts are still young and
tender.
An abnormal development of germ tubes instead of the usual promycelium has been reported in some instances. According to Lloyd & Ridgway ^^^ several crops of basidiospores are produced in one season. The various species usually make good subjects with which to
study infection. The teliospore masses placed in water soon become covered with basidiospores. Suspensions of these in water
applied to susceptible hosts usually give positive results readily. G. juniperi-virginianae Lk. ~^^' '-''' ^^^(=G. macropus) Schu. 0. Pycnia epiphyllous.
1. Êcia (=Roestelia pyrata) chiefly hypophyllous, usually in annular groups, on thickened discolored spots, at first cylindric,
0.1-0.4
mm.
in
diameter;
peridium splitting extremely

early,
becoming fimbriate to
the base, strongly revolute; cells peridial usually seen
only in side view, long and narrow, 10-16 x 65-100 /x,
becoming much curved when wet, inner and side walls

rather sparsely rugose
with
ridges extending half way across the side walls; secio-
spores
wall
globoid
ellipsoid,
or broadly 16-24 x 21-31 fi,
light
ju
chestnut-brown,
finely
2-3
c'ose.
thick,
verru-
Fig. 265.
G. juniperi-virginiana', aecia.
After Heald.
III.
Telia
or
or
globoid
appearing on reniform galls
5-30
eter,
mm.
more
in
diam-
in
evenly disposed, cylindric or cylindric-acuminate, 1.5-3 mm. diameter by 10-20 mm. long, golden-brown; teliospores 2-celled,
ellipsoid,
rhombic-oval or narrowly
15-21 x 42-65 m; slightly or
365
not constricted at the septum, wall pale cinnamon-brown, thin, about 1 m; pedicel cylindric, 3-5 ^ in diameter; pores two in each cell near the septum. I. JEcisb on apple both wild and cultivated. III. Telia on Juniperus virginiana and J. barbadensis. Destructive, particularly in East and South. Sporidia are matured in twelve to twenty-four hours after the spore-masses expand by moisture and as soon as the sori begin to dry they are carried away by wind and on suitable hosts infect through the
cell
walls
by
appresoria.
Two or three crops of sporidia
one season but the first crop is largest. ^^^ Each crop result in a corresponding crop of aecia. The stage on apple fruits shows as pale-yellow spots of pinhead size about seven
may may
arise in
to ten days after infection. The spots finally become orangecolored and in a few weeks the pycnia appear as black specks. On leaves hypophyllous cushions 0.5-1 cm. in diameter form on the
spots and bear the scia, the mature tubes of which are split and recurved giving a stellate appearance. Êciospores pass back to
the cedar in
summer and cause

when the
infection.
The mycelium
^^'
here
remains practically dormant according to Heald

lowing spring
tehal galls first
until the fol-
become
visible.
These
galls grow throughout the summer, mature in the fall, and give The mycelium is rise to the teliospores during the next spring.
thus seen to be biennial.
G. clavarigeforme (Jacq.) D. C.
I.
'ô'^'
ôs-
2u
Mcia hypophyllous,
in small
fructicolous
crowded
groups 2-3
times in larger groups on densely aggregated on the fruits and occupying part or
surface, cylindric, 0.7-1.5
or caulicolous, usually across on the leaf blades, somethe veins, petioles and twigs, often
mm.
all of
the
mm.
high by 0.3-0.5
mm.
in diameter;
ing; pcridial cells long
peridium soon becoming lacerate, usually to base, erect or spreadand narrow, often becoming curved when
n, linear
wet, linear in face view, 18-30 x 80-13

wall
or linear-oblong
in side view, 15-25 n thick, outer wall 1-2 n thick, smooth, inner
and side walls 5-7 n thick, rather coarsely verrucose with roundish or irregular papilla) of varying sizes; aeciospores globoid, 21-27 X 25-30 fx, wall light cinnamon-brown, 2.5-3.5 ju thick,
moderately verrucose.
366
III. Telia caulicolous, appearing on long fusiform swellings of various sized branches, numerous, scattered, or sometimes aggre-
gated, cylindric, or slightly compressed, 5-10 mm. long by 0.8-1.5 mm. in diameter, acutish, or sometimes forked at the apex,
brownish-yellow; teliospores 2-celled, lanceolate, 13-20 x 40-80

above,
n,
occasionally longer, rounded or narrowed
usually narrowed below, very slightly or not at all constricted at the septum, wall golden-yellow, thin, about
1
n; pores 2 in
each
cell,
near the sep-
tum.
I.
iEcia on Crataegus spp., Amelanchier,
Fig.
266.
GymnosporanAfter
Aronia, Cotoneaster, Cydonia, and Pyrus. III. Telia on Juniperus communis, J.
mPtinytd[ofpo*'
Richards.
oxycedrus, and J. sibirica. Spindle-shaped swelUngs occur on Juniper branches. Cylindric spore-masses ooze through rifts
in the bark. Jl^ciospores shed in June germinate at once on Juniper twigs and result in the following year in swellings which often Jater cause death. In spring the spore-masses emerge and the teliospores germinate in situ. Upon the Rosaceous hosts spots
appear eight to fourteen days after infection. Kienitz-Gerloff reports the occasional formation of a germ tube instead of a promycelium. This is, however, to be regarded as an abnormal
condition.
G. globosum Farl.
O. and
I.
ô^- 213. 215. 216
^cia
chiefly
hypophyllous and crowded irregularly
or rarely in approximately annular groups 2-7 mm. across, cylindric, 1.5-3 mm. high by 0.1-0.2 mm. in diameter; peridium soon
splitting in the
upper part, becoming reticulate half way to base; peridial cells seen in both face and side views, broadly lanceolate in face view, 15-23 x 60-90 n, linear rhomboid in side view, 13-19 n
thick, outer wall
about
1.5
^t
thick, smooth, inner
and
side walls
II, thick, rather densely rugose with ridge-like papillae of varying length; seciospores globoid or broadly ellipsoid, 15-19 x 18-25 IX, wall light chestnut-brown, 1.5-2 n thick, finely verru-
3-5
cose.
III. Telia caulicolous,
appearing on irregular globoid, gall-like
367
excrescences 3-25 mm. in diameter, unevenly disposed, often separated by the scars of the sori of previous seasons, tongue or wedge-shaped, 1.5-3 mm. broad by 2-5 mm. long at the base and
6-12 mm. high, chestnut-brown; teliospores 2-celled, ellipsoid, 16-21 X 37-48 M) somewhat narrowed above and below, slightly
at the septum, wall pale cinnamon-brown, 1-2 thick; pores 2 in each cell, near the septum. I. ^cia on apple, pear, Crataegus, quince, mountain ash.
constricted
III. Telia on Juniperus virginiana and J. barbadensis. Common and widely distributed in eastern America. The telial galls are from 0.5 to 2.5 cm. in diameter, very irIn late spring dark-brown spore-masses, later yellowregular.
orange, 0.5 to 2.5 cm. long appear. The Roestelia spots are 0.5-1.0 cm. across.
Pycnia blackish
hypophyllous spots, long, slender, soon splitting and becoming fimbriate. Mesospores occur The seciospores germinate on the cedar. The occasionally. mycelium stimulates the hosts to extra formation of parenchy-
above.
The
secia
are on thickened
mateous
tissue.
G. juniperinum (L.) Mart. I. ^cia (=Roestelia penicillata [Pers.] Fries.) hypophyllous, in annular or crowded groups, 2-5 mm. across on large thickened
discolored spots, at
high, 0.5-1 mm. in fimbriate to base and diameter; peridium soon becoming finely somewhat twisted or incurved; peridial cells usually seen only in
first cylindric,
0.5-1.5
mm.
medium
rhomboid, very thick, 30-35 x 60-90 fx, outer wall thin, 2-3 n, smooth, inner wall medium thick, 7-10 fx, side walls very coarsely rugose with thick, somewhat rugose, irregular ridges, roundish or elongate ridge-like papillae interspersed; seciospores globoid, very large, 28-35 x 30-45 fi, wall
side view,
chestnut-brown, thick, 3-5

III. Telia
n,
rather finely verrucose.
on hemispheric swellings (1-4 cm. long) breaking forth along the sides of the larger branches, or on subglobose galls (1.5-2 cm. in diameter) on the smaller branches, applanate, indefinite, usually of considerable size, often
caulicolous,
appearing
covering
the
whole hyjiertrophied area, sometimes becoming

teliospores 2-celled,
m,
patelliform
ellipsoid,
when expanded, chocolate-brown;
18-28 x 42-61
usually slightly narrowed both above
368

slightly or not constricted at the septum, wall
and below,
cinnamon
brown, 1-1.5 /x, thick; pores usually 3 in upper cell. 1 apical, 2 near the septum, in the lower cell 2 pores near the septum.
on apple and mountain ash. on Juniperus communis and J. sibirica. In Europe. The teliospores occur on both twigs and leaves. Marked deformation is caused by this stage on leaves and petioles.
I
and
III. Telia
G. clavipes C. & P. ^^ ( =*G. germinale [Schw.] Kern). I. ^cia (=Rcestelia aurantiaca) on stems and fruits, crowded
on hypertrophied areas of various size on the twigs and peduncles, occupying part or nearly all of the surface of the fruits, cylindric, 1.5-3 mm. high by 0.3-0.5 mm. in diameter; peridium whitish, becoming coarsely lacerate, sometimes to base, erect or spreading; peridial cells seen in both face and side views, poh^gonal-ovate or polygonal-oblong in face view, 19-39 x 45-95 /i, rhomboid in side view, 25-40 /x, thick, outer wall moderately thick, 3-5 n, inner wall
very thick, 13-23 fx, coarsely verrucose with loosely set, large, irregularly branched papillae, side walls verrucose on inner half similar to inner wall; seciospores globoid, large, 31-32 x 24-39
.
/jl,
wall pale yellow, thick, 3-4.5
/x,
rather coarsely verrucose with
crowded
slightly irregular papillae.
III. Telia caulicolous,
usuall}^ aggregated, roundish,
appearing on slight fusiform swellings, 1-4 mm. across, often confluent,
hemispheric,
ellipsoid,
1-3
mm,
18-26 x 35-51
high, orange-brown; teliospores 2-celled, /i, roundish or somewhat acutish above,
obtuse below, slightly or not constricted at the septum, wall yellowish, 1-2 n thick, slightl}^ thicker at the apex; pedicles carotiform, 9-19 /x in diameter near the spore; pores one in each apical in the upper, near the pedicel in the lower.
I.
cell,
iEcia
on
Amelanchier,
Aronia,
Crataegus,
Cydonia, and
apple.
III. Telia
on Juniperus communis and G. cornutum (Pers.) Arth.

rather
J. sibirica.
uncommon
species with I (=Roestelia cornuta [Pers.]
on Sorbus spp. and III on Juniperus communis and J. sibirica. Ranging from New York to Wisconsin and northward; also in the mountains of Wyoming and Colorado: Europe. Of no considerable economic importance in America.
Fries)

G. eUisii (Berk.) Farl.
369
Êcium unknown. III. Telia on Cupressus thyoides. Probably of very small economic importance. G. transformans (Ellis) Kern. (=Roestelia transformans
I.
Ellis).
iEcia on Pyrus arbutifolia, which is of no economic importance. Confined to a small area from Massachusetts to New
I.
Jersey.
III. Telia
unknown.
-^' î^
G. nidus-avis Thax.
I. vEcia amphigenous, especiallj^ fructicolous, cylindric, 2^ mm. high by 0.4-6.7 mm. in diameter; peridium soon becoming irregu-
larly lacerate usually to base, slightly spreading; peridial cells, seen in both face and side views, lanceolate in face view, 15-23 x 55-88 n; linear in side view, 14-18 n, thick, outer wall 1-.5 n
thick, smooth, inner and side walls 5-7 m thick, coarsely rugose with narrow ridges, with shorter, often roundish papillae interspersed; seciospores globoid or broadly ellipsoid, 18-23 x 23-28 n,
wall cinnamon-bro\vn, rather thick, 2.5-4 n, very finely verrucose, appearing almost smooth when wet.
III. Telia
cauUcolous, often dwarfing the young shoots and
brooms, usually causing a reversion of the leaves to the juvenile form, sometimes appearing on isolated areas on the larger branches and producing gradual
enlargements, solitary or rarely confluent, of variable size and shape, roundish to oval on the young shoots, 1-2 mm. across, oval
causing birds' nest distortions, or witches'
on the woody branches, 1.5-3 mm. wide by long, pulvinate when young, becoming hemispheric, dark reddish-brown; teliospores 2-cellcd, ellipsoid, 16-23 x 39-55 ju,
to nearly elliptic
2-7
mm.
wall pale cinnamon-brown, rather thin, 1-1.5 n, very slightly thicker at apex; pores one in a cell, apical. Mycelium perennial in leaves, branches or trunks of Juniperus virginiana very commonly inducing a "bird's nest" distortion.
I.
Mc'ia on Amelanchier
III. Telia
and quince. on Juniperus virginiana.
G. sabinae (Dicks) Wint. O and I (=Roestelia cancellata), on pear in Europe. III. Telia on several Junipers.
370
The telial mycelium is perennial and causes swellings. From these in spring ooze the gelatinous, transparent spore-masses. The mycelium in Juniperus causes increase in wood-bast and
rind, thickened twisted tracheids, increase in
number and
found
in the
thick-
ness of the medullary rays.

itself.
No mycelium
is
wood
G. biseptatum
I.
Ell.
hypophyllous, usually in groups of 2-8, rarely solitary, borne in gall-like pyriform protuberances 1-1.5 mm. in diameter by 1.5-3 mm. high, cyUndric, 0.5-0.8 mm. in diameter by 2-4 mm. high; peridium soon becoming finely
JEcia
(=RoesteUa botryapites)
cancellate, not dehiscent at apex; peridial cells cylindric, hyphallike, 9-14 n in diameter by 145-190 n long, often irregularly bent,
and side walls of equal thickness, about 1.5-2 n, whole surface smooth; seciospores globoid, small, 15-17 x 16-22 n, wall dark cinnamon-brown, rather thick, 2.5-3 fx, moderately
outer, inner,
verrucose.
III. Telia caulicolous, appearing on fusiform swellings, scattered, oval or irregular, about 1.5-3 mm. wide by 2-7 mm. long, often confluent, hemispheric, chestnut-brown; teliospores 2 to 4-celled,
n, usually rounded above, somewhat narrowed beconstricted at the septa, wall pale-yellow, 1.1-5 fx low, slightly thick, pores 2 in each cell, near the septa.
13-19 X 35-77
I.
JEcia.
III. Telia
on Amelanchier, on Chamaecyparis.
in small
G. nelsoni Arthur. JEcia hypophyllous and fructicolous, usually groups 1-2 mm. across, cylindric, 2-4 mm. high by 0.2-
0.3
mm.
in diameter; peridium whitish, dehiscent at
apex and
also rupturing
more
or less along the sides; peridial cells seen in
both face and side view, 18-35 x 75-115 n, linear rhomboid in side view, 16-35 fi, thick, outer wall rather thin, 1.5-2 n, smooth,
inner and side walls rather thick, 7-12
/x,
verruculose; seciospores globoid, 19-26 x 21-29 brown, 2-3 thick, finely verrucose.
fjL
evenly and densely wall chestnut/jl,
III. Telia caulicolous, appearing on firm, woody, globose galls 0.5-5 cm. in diameter, unevenly disposed, densely aggregated or often separated by the scars of the sori of previous seasons, ir-
regularly flattened, about 1-1.5
mm.
broad by 1-5
mm.
long at the

base by 3-4
371
mm.
high, often confluent, Ught chestnut-brown; telio-
spores 2-celled, narrowly ellipsoid, 18-26 x 50-65 /x, narrowed at both ends, slightly constricted at the septum wall pale cinnamonbrown, 1-1.5 thick; pores two in each cell, near the sep;
fjL
tum.
I.
iEcia on Amelanchier, Peraphyllum, quince and pear.
on Juniperus spp. Range; Alberta, south to Colorado and Arizona. G. japonicum ^yd. '^^^ I. JEc'm (=R. koreaensis), on Pear. III. Telia on Juniperus. This form has been imported into America.
III. Telia
G. torminali-juniperinum (Ed.) Fischer. This species has its a?cial stage on species of Sorbus and its telia on Juniperus in Europe. It is closely related to G. cornutum of the northern part of our own continent, and of
Europe. G. yamadae Miyabe.

found.
It infests the apple
Only the aecia of this species have been and various other species of Malus in
Japan.
Uromyces Link
(p.
355)
0. Pycnia spherical with minute ostioles. 1. Êcia with peridia, spores without pores.
II. Urediniospores generally with many germ pores, unicellular, spherical, ellipsoid or variously shaped, usually rough.
III. Teliospores
unicellular, pedicellate,
with an apical germ
pore.
The
unicellular teliospores
their single
may be
germ
distinguished from urediniopore,

also
spores by thicker walls
apical
usually
by
their
and absence
of the roughness so characteristic of
urediniospores.
The genus is a very large one, with hundreds of species, which exhibit heteroecism, autoecism, biologic specialization and the various
types regarding spore forms that are noted on pages 324-327. U. appendiculatus (Pers.) Lev.-'^
I.
yEciospores angularly globose, whitish, slightly punctulate.
372

/x.
17-32 X 14-20
X
II,
Urediniospores pale-brown, aculeolate, 24-33

elliptical
16-20
n.
III. Teliospores
or
subglobose,
.smooth,
dark-brown, apex much thickened, with a small, hyaline, wartlike papilla, 26-35 x 20-26 /x-
An
autoecious
eu-type.
On
Phaseoliis,
Dolichos
and other
related legumes.
The
sori
usually appear late
in the season
on
leaves, rarely
on
stems and pods. The mycelium is Great difference in local.

varietal susceptil)ility is noted. U. pisi (Pers.) de B.'^
Leaf Tissue
Fig. 267.
Diagrammatic seclion of ureof U. appendiculatus.
I.
^r,
=yEcidium cyparissia?).
j.j.
i
dinium
After
Êcia Scattered over the whole
xi,
leaf surface. Peridia cup-shaped, with whitish edges. Spores subglobose or polygonal, orange, finely verrucose, 17-26 ^t in diameter. II. Uredinia roundish, scattered or crowded, cinnamon-brown.
W^*'*^^^-
Spores
subglobose
or
elongate,
jêllowish-
brown, echinulate, 1720 X 20-25 MIII. Telia
roundish
blackish.
Epidermis of Leaf
or
elliptical,
Spores
shortly
subglobose
or
elliptical, finely
but
closely
punctate,
apex only
slightly
Leaf
Younq Spore
thickened, 20-30 x 17Pedicels long, 20 n.

colorless, fragile.
Tissue
heteroecious
euin
type not America:

II
found
Fig. 268. Diagrammatic section of teliospores of U. appendiculatus. After Whetzel.
and I on Euphorbia. and III on Lathyrus, Pisum,

aecial
Vicia.
it is
The
stage dwarfs the host in which
perennial
373
U. fabae (Pers.) De B.--" This is an autoecious eu-type which causes a rust of the broad bean, vetches, peas. U. trifolii (Hed.) Lev.--^'-^ An autoecious eu-type.
iEcia in circular clusters, on pallid spots. Peridia shortly cylindric, flattish, on the stems in elongated groups; edges whitish, torn. Spores sul)globose or irregular, finely verrucose, paleI.
orange, 14-23 n in diameter.

II.
Uredinia pale-brown, rounded, scattered, surrounded by the
Fig.
Fiu. 2(39. I loniyccs appendiculatus, tcliospore gcr-
2t<j. Urediniospore and tclioof Uiomyspore Afccs trifolii.
minating.
After Tulasne.
ter
Cobb.
torn epidermis.
torial
Spores round or ovate, with three or four equagerm pores, echinulate, brown, 20-26 x 18-20 juIII. Telia small, rounded, almost black, long covered by the
epidermis. Spores globose, elliptical or subpj^riform, with wartlike incrassations on their summits, smooth, dark-brown, 22-30 x
15-20
/x.
Pedicels long, deciduous.
Cosmopolitan on white, crimson and alsike clovers. Stages and I are most common on Trifolium rcpens, least common on T. incarnatum. Pycnia appear in early spring or even in winter. The seciospores germinate readily in water and give infections which
give rise to urediniospores in about two weeks. Urediniospores ma}'^ be produced throughout the summer and may even survive
the
winter.
sori
separate
Teliospores arc produced in the uredinia or in The teliospores by infection late in the season.
374
give rise to the pycnial and secial stages. Considerable distortion arises in parts affected by either stage. U. fallens (Desm.) Kern.^^^ A form on crimson, zig-zag and red clover often confused with the last species.
and
II.
unknown.
Urediniospores with four to six scattered germ pores. III. Teliospores similar to those of U. trifolii.
U. medicaginis Pass.
O
II.
and
I.
Pycnia and
secia as in
U.
pisi.
Uredinia chestnut-brown, spores globose to

light-brown.
elliptic,
17-
23
tx,
Fig. 271.
Uredo stage of U.
betae.
After Scribner.
III. Telia
dark-brown, spores ovate-elliptic or pyriform 18-28 x
14-20.
heteroecious eu-type.
II and-'III
on
alfalfa
is
U. minor Schr.
on Euphorbia; in Europe, Europe and America. an autcecious opsis-type, I and III on TrifoI,
and
clovers in
autcecious eu-type; on members In the United States of the genus Beta both wild and cultivated. observed only in California. Recorded in Europe, Africa, Australia.
'
lium montanum. Bfi U. betae (Pers.) Tul.
9Q1
An
U. kuhnei Krug. occurs on sugar cane.^^^ U. dactylidis Otth. is a heteroecious eu-type; II and III on
Phleum, I on Ranunculus, in Europe. U. poae Rab. is a heteroecious eu-type; Ficaria; II and III on Poa.
on Ranunculus and

U. caryophyUinus (Schr.) Wint.226-229 I. iEcia on Euphorbia in Europe. II. Uredinia sparse, confluent on stems, spores round, or oblong, 40 x 17-28 n, light-brown.
III. Teliospores globose, irregular or ovoid,
375
elliptic
apex thickened 2335 X 15-22 mm., pedicel 4-10 ix. II and III on cultivated carnations and several other members
of the genus Dianthus. I on Euphorbia gerardiana. It has been known in Europe since 1789 but was not noted in the United States
until 1890 when it was found by Taft at Lansing, Mich. It soon invaded the whole country causing great loss. There is large racial
difference in host susceptibility.
urediniospores germinate readily in water and serve to propagate the fungus. Studies of the effects of toxic substances
The
upon these have been made by Stevens

aecial
^^
stage has recently been recognized
and by Stewart.^-'^ The ^^3 as M. by Fischer
euphorbise-gerardianae.
Less important species are: U. ervi (Wallr.) Plow, an autoecious eu-type on Vicia in Europe; U. erythronii (D. C) Pass, an opsistype occasional on cultivated Lilium in Europe. U. ficariae Schw.
is on Ficaria; U. pallidus Niess. a lepto-type on Cytisus; U. scillarum (Grev.) Wint. a micro-type on Scilla and Muscari. U. jaf-^^ frini Del. is reported on vanilla; U. colchici Mas. on Colchinum
speciosum in Europe.
Puccinia Persoon
0,
I,
(p.
355)
II,
as in
Uromyces.
separate,
pedicellate,
cells
cell
III. Teliospores
consisting of
two superimposed
with a germ pore.

piercing
its
The
superior
produced in flat sori, each of which is provided has its germ pore, as a rule,
apex; in the inferior or lower the
germ pore
is
placed
immediately below the septum. Mesospores (p. 327) are not rare. They are merely teliospores with the lower cell wanting, and function as teliospores. Some one thousand two hundred twenty-six species are enumerated by Sydow presenting great diversity heteroecism and biologic variation.
spore relation,
376

is
P. cerasi Ces.
a hemi-type on cherries in Southern Europe.

III
P. ribis-caricis Kleb.
differentiates five species of Puccinia on Ribes belonging to the Ribis-Carex group. These are P. pringsheimiana P. ribis-pseudocyperi, P. ribis nigri-acutse, (I.=^'E. grossulariae.)
on Ribes. Klebahn -^^

I
II
and
on Carex.
and P. magnusii. P. asparagi D. C.^s^-^ô I. Peridia in elongated patches upon the stems and larger branches, short, edges erect, toothed. Spores orange-yellow,
P. ribis nigri-paniculatse
round, very finely echinulate, 15-26 diameter.
/x
in
II. Uredinia brown, flat, small, long covered by the epidermis. Spores irregu-
larly
late.
round or oval, 18-25 X 20-30
clear-brov^Ti, echinuM-
III. Telia
black-brown, compact, pul-
elongate or rounded, scattered. Spores oblong or clavate, base rounded, apex thickened, darker, central convinate,
striction
slight
or
absent,
deep chest/jl.
nut-brown, 35-50 X 15-25

Fig. 272. St'Ctioii through black nist pustule, showing teliospores of P. asAfter Smith. paragi.
Pedicels
persistent, colorless or brownish, as long
as or longer than the spores. An autoecious eu-type on Asparagus, cultivated and wild. The fungus has
been kno^vn in Europe since 1805 but did not attract attention "^^ in the United States until 1896 in New Jersey when it began ^^^ its devastating westward migration across the country reaching California in 1900 or 1901.
The secial stage appears in early spring; the aeciospores may germinate at once or if dry remain viable for several weeks, their germ tubes penetrating the host in most cases stomatally. The uredinia appear in early summer soon after or with the secial
wind borne, distribute the fungus. The uredinioremain viable a few months when dry. The telial stage spores
stage and,
appears late in the season and germinates only after hibernation. Unicellular spores, mesospores, are sometimes met.

P. bullata (Pers.) Schr.
is
377
a brachy-puccinia which
its
is
autoecious
on on
celery, parsley,
dill
and other umbellifers.

uredinial
P. apii (Wallr.) Cda. also occurs in

celery.
is
and
telial
stages
P. castagnei Thiim
recorded for celery in France.
Rud. is a hemi-type on cultivated onions. P. porri Sow. is an autoecious rust which is sometimes deP.
allii (I).
C.)
structive to onions in
.^
Europe. P. endiviae Pass.-ôccurs on

Italy P.
.^<^=p^^
o '"'n moa9^yr
C)
endive in
and America.
phragmitis
on
circular
Fig. 273.
Schum.^^^--^^
I
(=iE. rubellum).
Peridia
red spots 0.5-1.5 cm.

in diameter, shallow,
Cross-section of
secia of P. asparagi.
After Smith.
edges white, torn. in diameter.

II.
Spores white, subglobose, echmulate, 15-16 n
without paraphyses. 25-35 X 15-23 m-
Uredinia rather large, dark brown, elliptical, pulverulent, Spores ovate or elliptical, echinulate, brown,
III. Telia large, long, sooty black, thick, often confluent. Spores elliptical, rounded at both ends, markedly constricted in the middle, dark blackish-brown, smooth, 45-65 x 16-25 fx. Pedicels very long, 15(>-200 x 5-8 3'ellowish, firmly attached.
/jl,
Heteroecious; I on
mitis.
Rumex and
Found only
rarely in America,^'*^'
rhubarb, II and III on Phrag^^^ except in the middle
west.
P. cyani (Schl.) Pass, is on cultivated Centaurea. P. tragopogonis (Pers.) Cda.
shortly
I.
^cia on the whole plant

cylindrical, at
first
leaves, stems, bracts, receptacles
mammiform,
peridia with whitish,

/x,
torn edges.
Spores rounded, verrucose, orange-yellow, 18-27

/i
sometimes as much as 35
long.
Mycelium
diffused throughout
the host-plant. III. Telia brown, few, small, scattered, elliptical or elongate.
378
long
covered by the epidermis. Spores broadly oval, often almost globose, slightly constricted, apex not thickened, thickly verrucose, brown, 26-48 x 30-35 ju. Pedicels short, colorless,
deciduous.
MyceUum
The
locaUzed.
An
.opsis-type
on cultivated Tragopogon.
unknown.
variable.
teliospores are often unicellular
Urediniospores are and are very

is
P. taraxaci Plow,
is
common on
dandelion.
P. cichorii Pass,
is
hemi-type on Cichorium. P.
isiacae
on Phragmitis
thought to be
Fig. 274.
P. graminis, telium and germinating

teliospore.
After Carleton.
the
is
telial
stage of
M.
brassicae
on cabbage.
^^^
P. fagopyri Barcl.
found on buckwheat. P. menthae Pers. ^

I.
torn; principally
rarely
Êcia with peridia immersed, fiat, opening irregularly, edges on the stems, which are much swollen, more
on concave spots on the
leaves.
Spores subglobose or
polygonal, coarsely granular, pale-yellowish, 17-26 x 26-35 mII. Uredinia small, roundish, soon pulverulent and confluent, cinnamon-brown. Spores irregularly rounded or ovate, echinulate,
pale-brown, 17-28 x 14-19
n.
roundish, pulverulent. Spores elliptical, oval, or subglobose, central constriction slight or absent, apex with a hyaline or pale-brown papilla, verrucose, deep-brown, 26-35 X 19-23 ix. Pedicels long, delicate, colorless.
III.
TeUa black-brown,
An
autcecious eu-type on
many
mints.

P. graminis Pers.
Tk
379
rt
.
'
166-182, 246, 306, 322
...
I {=JE. berberidis). Spots generally circular, thick, swollen, Pcridia cylindrical, with whitish reddish above, yellow below. torn edges. Spores subglobose, smooth, orange-yellow, 15-25 mII. Uredinia orange-red, linear, but often confluent, forming very long lines on the stems and sheaths, pulverulent. Spores elliptical, ovate, or pyriform, with four very marked, nearly
equatorial
germ
pores, echinulate, orange-yellow,
25-38 x 15-20
m-
naked, linear, generally forming lines on the sheaths and stems, often confluent. Spores fusiform or clavate, constricted in the middle, generally attenuated below, apex much
III. Telial persistent,
thickened (9-10 fi), rounded or pointed, smooth, chestnut-brown, 35-65 X 15-20 m- Pedicels long, persistent, yellowish-brown. O and I on Berberis and Mahonia. II and III on Avena, Hordeum, Secale, Triticum and nearly Of great importance on wheat in the Great fifty other grasses.
and along the Ohio. This fungus was the subject of the classic researches of de ^^^ Bary begun in 1865 and has since repeatedly served as the basis of fundamental investigations in parasitism, cytology and biologic
Plains
specialization.
cereal infection
seciospores can bring about seems to have been shown as early as 1816. ^^ Inoculations in the reverse order were made in 1865. ^^^ Extensive
That the barberry
studies
by Eriksson
^^"^
are interpreted
species
was formerly regarded as one

logic
by him to show that what must be separated on bio-
grounds into several races which he finally erects as species, though others do not agree that their rank should be specific. These are: P. graminis secalis. P. graminis avenae. P. graminis
tritici.
P. graminis
airae,
P. graminis poae.
P. phlei-pratensis.
These words from Butler and
Hayman
^^
show the complexity
of the status of these biologic forms.

it has become more and more established that which are capable like these rusts of living on parasitic fungi, several hosts, tend to develop 'races' on their different hostspecies, marked off from each other by definite characters. Sometimes the characters are such as are capable of being detected
"Of
late years
microscopically.
ance, identical,
Usually, however, the fungi are, to
all
appearof
and
differences only appear
when
their
manner
380
The chief of these is the incapacity of a life is carefully studied. Such forms as race to attack the host-plants of another race. are thus outwardly identical but which show a constant difference
in their
mode
of life are
known
as "biological" species or as/ormce
speciales.
"A
specialized
fixed'
form
is
sharply
of the
according as
considered to be 'sharply fixed' or 'not it is wholly incapable, or sometimes
capable, of attacking the host-plants of the other specialized forms
Thus the P. graminis of wheat (P. graminis not sharply fixed, for it can attack barley, rye, f. sp. Tritici) The P. graminis found on grasses of the genus &c., sometimes.
same fungus.
is
Agrostis (P. graminis f. sp. Agrostis) is sharply fixed, for it attacks this genus only and does not pass to the other grasses on which
it
has been tried.
fixed forms, such as the P. graminis be entirely incapable of attacking some of the In other species which bear other forms of the same fungus. words a form may be not sharply fixed in regard to some host-
"But even the not sharply
of wheat,
may
of this occurs in India.
plants and sharply fixed in regard to others. A striking instance P. graminis can be divided amongst others
on wheat (/. sp. Tritici), rye and barley (/. sp. Secalis), and oats (/. sp. Avenoe). The/, sp. Tritici can attack barley sometimes, and did so in four out of sixteen of our inoculations, but it does not, in India at least, attack oats. Hence it is sharply fixed in regard to oats and not sharply fixed in regard to barley. The /. sp. Secalis on barley also does not pass to oats, but infected wheat doubtfully in two out of sixteen inoculations. These two forms are common in India, and the practical bearing of their
into races
not passing to oats is considerable, for the /. sp. Avenoe has not yet been observed in this country." The mycelium branches intercellularly and bears small haus-
which penetrate the cells. In the barberry it is local. The epiphyllous pycnia appear first followed soon by the mainly hypophyllous secia. The flask-shaped pycnia at maturity bear numerous pycniospores and exserted paraphyses. Their hyphse are orange-tinted, due to a coloring matter in the protoplasm or
toria
later in the cell walls.
The aecium
originates in the lower region of the mesophyll

from a hyphal weft.
spores every alternate
381
chains of
The
fertile
branches give
rise to
which atrophies. The outer row of sporophores and potential spores remains sterile to form the peridium. When young the acium is immersed and globular, at maturity erumpent and forms an open cup. These spores
cell of
germinate by a tube capable upon proper hosts of stomatal infection and following this of producing the uredinium.
Urediniospores are produced throughout the season even through the winter under proper climatic conditions. They also remain
viable for weeks
^^^'
^'^'^
and doubtless serve hibernation purposes.

teliospores,
^'^^
Teliospores
arise
later in
the season in the uredinia or in
separate
telia.
Unicellular
mesospores,
are
oc-
TeUospores germinate best after normal outdoor hibernation, producing the typical 4-celled promycelium,
casionally seen.
long stcrigmata and solitary basidiospores. If under water the usual promycelium becomes abnormal and resembles a germ
The aecial stage may not occur under certain climatic conditions, and the uredinia alone perpetuate the fungus. -^^' -^-' -'^' ^^ It therefore follows that eradication of the barberry as was attempted by legislative enactment in 1660 in Europe and in 1728 and 1755 in Connecticut and Massachusetts -^^ does not exterminate the rust
^^'^
(see also
^^^'
~^).
Basidiospores were shown by
De
Bary,
^^^
confirmed by
Ward
-^^
and Eriksson, to be incapable

ficient
of infecting
wheat
leaves.
Suf-
such attempts have, however, not been

^^
made on young
tissue.
succeeded in securing germination of pycniospores but the resulting mycelium soon died and infection was not attained. The same author holds that seciospores may remain viable about a month, the urediniospores a much shorter time. Still hibernation by urediniospores is possible where climatic relaJaczewski
tions allow the formation of
^^^
new
uredinia during the winter.

'''' ^â- 306
P. rubigo-vera (D. C.) Wint."'^'
asperifoUum, Pers). Spots large, generally circular, disPeridia flat, broad, with torn white colored, generally crowded.
edges.
II.
I(=^.
Spores subglobose, verrucose, orange-yellow, 20-25 ju. Uredinia oblong or linear, scattered, yellow, pulverulent.
382
Spores mostly round or ovate, echinulate, with three or four germ pores, yellow, 20-30 x 17-24 mIII. Telia small, oval, or linear, black, covered by epidermis, surrounded by a thick bed of brown paraphyses. Spores ob-
long or elongate, cuneiform, slightly constricted, the lower cell generally attenuated, apex thickened, truncate or often obliquely conical. Spores smooth, brown, variable in size, 40-60 x 15-20 mPedicels short.
and I on BoraginacesB. Heteroecious; II and III on rye. The teliospores germinate as soon as mature.
Fig. 275.
P. rubigo-vera, section
After Bolley.
of uredinium.
P. triticina Erik,
of all rusts of the
is
the most
common and
widely distributed
India. ^^
It
United States and is a serious wheat pest in ordinarily shows only the uredinial stage. The telio-
spores germinate the following spring after a resting period. Coextensive with wheat culture.^"^ Epidemics are frequent. ^' ^' ^^ (see also ^''^) has shown it capable of hibernation Bolley
by
urediniospores and by live winter mycelium and it has further been shown that the spores themselves can survive freezing in ice. The
secial
and two races. The name P. dispersa is also used to cover the same two species. P. rubigo-vera tritici on wheat and P. rubigo-vera secalis on rye.
This species
stage can be entirely omitted. is combined with P. triticina
by Carleton
^^^
treated as
383
The secial stage of the former of these is not known. Its uredinia survive the severest winters even so far north as the Dakotas.
P. coronata Cda.i^^,
248. 306
I (=iE. rhamni). Peridia often on very large orange sweUings, causing great distortions on the leaves and peduncles, cylindrical, with whitish torn edges. Spores subglobose, very finely verrucose,
orange-yellow, 15-25 x 12-18 ix. II. Uredinia orange, pulverulent, elongated or linear, often confluent. Spores globose or ovate, with three or four germ pores,
confluent, long covered by the epidermis. Spores subcylindrical or cuneiform, attenuated below, constriction slight or absent, apex truncate.
echinulate, orange-yellow, 20-28 x 15-20 n. III. Telia often linear, persistent, black,
Fig. 276.
P. coronata, various teliospore forms.

six or
n.
After Bolley.
somewhat thickened, with

brown, 40-60 x 12-20
Heteroecious
II
;
seven curved blunt processes,
Pedicels short, thick.
I, on Rhamnus frangula. and III on various grasses but not on oats. From this form as earlier understood Klebahn has separated P. coronifera Kleb. on evidence derived from inoculations, and made the latter to include these forms with the aecial stage on Rhamnus cathartica and the uredinial and telial stages on Avena, LoUum, Festuca, Holchus, Alopecurus and Glyceria. P. coronifera has been still further divided by Eriksson into eight biologic forms and P. coronata into three such forms. ^'^^ P. glumarum (Schm.) Er. & Hu.-^^' ^^ is widely distributed on wheat, rye, barley and a few other grasses in India and Europe but is not known in America.^"^ Its secia are not known. By some this is regarded as a race of P. rubigo-vera. Both uredinia and teliospores have been reported in the pericarp of
grains.^
384
P. simplex (Korn.) Er.

I.
&
He.
Unknown.
II and III on barley in Europe and seemingly of recent introduction into the United States.^^ One of the least important of the grain rusts. Mesospores are
common.
P. sorghi Schw."'"'
Peridia hypophyllous, rarely amphigenous, I {=M. oxalidis). crowded, concentric, epispore smooth, 24-28 juII. Uredinia amphigenous, numerous, often confluent; spores globose to ovate, 23-30 x 22-26 mm., slightly verrucose.
Fig. 277.
Puccinia sorghi.
After Scribner.
III. Telia
obtuse, constricted.
amphigenous, black. Spores ovate-oblong or clavateEpispore thick, 28-45 x 12-17 n, smooth,

II
pedicel long, 5 n, persistent. and I on Oxalis. Hetercecious.
and III on Zea.
Of
little
The
it is
relation of the secial stage

5
believed,
was demonstrated by Arthur; ^^^ however, that hibernation is largely by the uredinioC.
spores.
P. purpurea dark-brown.
II.
Amphigenous, spot purplish,
sori
irregular,
Urediniospores ovate, 35 x 25-30 m, smooth, brown. brown, long-pedicellate, 40-45 X 22-25 ju. On Sorghum in Southern United States and West
III. Teliospores elongate, ovate,
Indies.

P. phlei-pratensis E.
I.
385
&
H.^'s-êi,
.ios-sos
^-Ec'ia
II.
Uredinia 1-2
i)robably on Berbcris, but rarely formed. mm. long on leaves and stems, confluent in or
lines 10
mm.
more
long, yellow-brown; spores oblong, pyriform,

fx.
spiny, 18-27 x 15-19

III.
Mycelium
sheaths
perennial.
and stems, 2-5 mm. long or dark-bro\m to black, open or partly more, confluent, narrow,
Telia
in
leaves,
Ficj. L'7S.
1*.
malvacearum.
Alter Holway.
Spores fusiform or club-shaped, medially constricted, chestnut-brown, apically thickened, 38-42 x 14-16 /x.
closed.
II and III on timothy grass. This species is closely related to P. graminis and probably a derivate from it, but it does not seem capable of infecting the bar^^^ berry under ordinary conditions.^^^' Inoculation experiments with timothy rust at Washington, D. C, show that it can be transferred easily to various grasses. Similar It is not a results have been obtained by Eriksson in Europe.
well fixed species
and by using bridging hosts
it
can be made to
386

it
transfer to various cereals which
will
not attack directly.

is
That
such transfers take place in nature to some extent

P.
probable.^^^
poarum Niess occurs on
bluegrass.
P. malvacearum Mont.^^^^^^
III. Telia grayish-brown, compact, round, pulvinate, elongate on the stems, scattered, seldom confluent, pale reddishbrown. Spores fusiform, attenuated at both extremities, apex
sometimes rounded, constriction slight or absent, apical thickening Pedicels firm, slight, smooth, yellow-brown, 35-75 x 15-25 sometimes measuring 120 n. long, A lepto-puccinia on three species of Althea, seven of Malva, two
(jl.
of
Malope; particularly serious on the hollyhock. A native of Chili, it was first known as a pest in Australia; soon afterward in Europe. It seems to have entered the United States sometime prior to 1886 and is now almost universal. The teliospores germinate immediately in suitable environment, mainly from the apical cell, or may remain alive over winter and originate the
spring infection. The mycelium also hibernates in young leaves. Mesospores are common. 3 to 4-celled teliospores are also met.
P. heterogena Lag. is also described on hollyhock from^"^ South America. P. chrysanthemi Roze.^^^"^''^
II.
Uredinia chocolate-brown, single or in
circular groups, hypophyllous, rarely epiphyl-
Spores spherical to pyriform. Membrane spiny and with three germ pores, 17-27 x 24-32 M.
lous.
III. Telia
dark-brown hypophyllous.
Telio-
spores rarely in uredinia, dark, obtuse, apex
thickened,
membrane
n.
25 x
35^3
thick, finely spiny, 20Pedicel 1-13^ times the spore
length.
urediniospores like the other urediniogpores in all other respects but 2-celled are a habit unique with this rust. In many places urediniofound;
ally
FiQ. 279. p. helianthi, uredinio- <and teliospores. After Cobb.
On
cultivated
Chrysanthemum.
Occasion-
may be produced continuously and teliospores be but rarely thus in America only urediniospores have been found. It was seen,
spores

first
387
country.
seen in America in 1896 (Mass.) and soon spread over the Numerous inoculation trials go to show that it is inde-
pendent of the other rusts

P. arenariae Wint.^^
III. Telia
common on
nearly related Compositse.^^^
compact, pulvinate, roundish, scattered, often circinate. Spores broadly fusiform or pyriform, summits pointed or rounded, often thickened, base rounded or attenuated, slightly constricted, smooth, pale yellowish-brown, 30-50 x 10-20 n.
Pedicels hyaline, colorless, as long as the spores.
A lepto-puccinia common on Dianthus. P. helianthi Schw.

0. Pycnia clustered. 1. iEcia in orbicular spots; peridial margins pale, torn; spores
orange, rarely whitish. II. Urcdinia minute, round, chestnut-brown, spores globose to
ovate, 22-26 x 17-22 fi, minutely spiny. III. Telia round, dark-brown to black; spores rounded at base,
slightly constricted,
38-50 x 20-27
fx,
smooth; pedicel hyaline,
equal to or longer than the spores. Autoecious on numerous species of Helianthus, probably divisible into numerous biologic forms. Imported from America to
Europe. Arthur ^^ used fifteen species of Helianthus on which to sow the teliospores of Puccinia helianthi produced on three species. The
on page 388. In the course of three years' work with this species sixty sowings were made.
results are given in table I
"Looking over the table it will be seen that each set of spores grew upon the species of host from which derived, but not upon the other two species, except that spores from H. Icetijlorus sown on H. mollis gave a tardy showing of pycnia, without further development. Also each set of spores grew luxuriantly upon H. annuus, and each made a feeble growth upon H. tomentosus, but on
other species they either failed to infect or made a feeble growth, with the single exception that spores from H. Icetiflorus grew well on H. scaberrimus.^' P. Helianthi thus affords an example of a
all
single species
having
many
races, for
which H. annuus acts as a
bridging host.
388
Table
I *
RESULTS OP INOCULATIONS OF HELIANTHUS RUST
THE FUNGI WPIICH CAUSE PLANT DISEASE
389
P. gentianae Strauss is a ou-puccinia on many species of cultivated gentians. P. gladioli Cast occurs on gladiolus. P. granulans Kale. & Cke. is on cultivated Pelargoniums in France i^^"
P. tulipae Sclir. on tulips; P. scillae Lk. on Scilla; P. schroeteri Pass, on Narcissus in Europe. P. pazschkei Diet, is a lepto-
puccinia on cultivated saxifrages in Europe.
P. horiana Hen.
Fil;.
280. p.
dianthi.
After Holway.
is
destructive on
Chrysanthemums
is
in Japan.-"^
P. iridis (D. C.)

of
Iris.
Duby,
in the
hemi-puccinia,
its
found
on many species
is
P. cannae Hen. in
uredinial stage
destructive to
is
West
Indies.
II
P. persistans Plow,
III
Cannas heteroecious. I on
on Agropyron. P. asteris Duby. is a very common lepto-puccinia on various asters. P. anemonesvirginianae Schw. is a lepto-puccinia common on anemone.
Thalictrum.
and
Key
Spores catenulate Peridium absent
to Uredinales Imperfecti
(p.
335)
1.
Caeoma,
p. 390.
Peridium present Toothed, body cui>shapcd

Fimbriate, body elongate
Irregularly split
2.
3. 4. 5.
iEcidium,
p. 390.
Roestelia, p. 391.
Spores not catenulate
Peridermium, p. Uredo, p. 392.
390.
390
^cidium Persoon
(p.
389)
Spores surrounded by a cup-shaped peridium; produced catenulate. in basipetal series. Germination as in Uredo.
The
cinia
species are very
numerous and belong

of the
in the
main
to Puc-
and Uromyces.
Most
forms of economic interest are
found under these genera. A few others of occasional economic bearing whose telial stage has not yet been recognized are given
below.
is perhaps identical with Puccinia See p. 378. A. tuberculatum E. & K.^^^ is reported as destructive on the poppy mallow. A. pelargonii Thlim. occurs A. cinA. otogense Lindsay on Clematis.^" on geraniums;
A. brassicse Mont, on Brassica
isiacse.
^'^''
namomi
Rac.
is
serious
on the cinnamon tree in Java.

(p.
CaBoma Link
389)
Sori without a peridium, accompanied by pycnia, with or without paraphyses, produced in chains. Germination as in Uredo. The forms are mostly stages of Melampsora, Phragmidium or
their kin.
Those
of
economic interest are found under Gymno-
conia and Melampsora.
Peridermium
Pycnia truncate-conic.
Levielle (p. 389)
Peridia caulicolous or foliicolous, erumpent, saccate to tubular, lacerate-dehiscent, spores catenulate or at maturity appearing
solitary,
globose to elliptic or oblong, polyhedral
by
pressure,
yellowish-brown.
The
secial
Epispore always verrucose-reticulate. stages of Coleosporium, Cronartium, Pucciniastrum,
Melampsorella and Chrysomyxa. The peridia usually extend conspicuously above the host surface,
by weathering. grow on the Coniferse, most of them on Pinus on both leaves, branches and bark. On the leaves the secia are much of the type shown in Fig. 256. When on the woody parts great distortion may be caused by the perennial fungus and much injury result to the wood (see Cronartium quercus, p. 352). The mycelium may live intercellularly in rind, bast and wood
irregularly
All of the species
and rupture

of pine
391
for years causing swellings of are either subcuticular or subepidermal and the Pycnia twigs. pycniospores often issue in a sweetish liquid, ^cia occur as
and continues to extend
wrinkled sacs emerging from the bark of the swollen places and bear spores perennially.
key to some thirty species is given by Arthur & Kem.^^^ So far as it relates to the distribution of the Peridermiums to
their telial genera it is as follows
:
Key to
Pycnia subcuticular
iEcia cylindrical Mcia, tongue-shaped
Species of Peridennium
Pucciniastrum.
Melampsorella, Melampsoridium.
Pycnia subepidermal .^cial peridia one cell thick On Pinus
On On
Picea
Coleosporium. Melampsoropsis.
Uredinopsis.
cell
Abies
thick
Pycnia subcorticular ^cial peridia more than one
Cronartium.
telial
Such forms as are

stage
is
of
economic interest and of which the
known
are discussed under Coleosporium, Cronartium,
Melampsorella, Melampsoropsis and Pucciniastrum, Several other forms are found on pine, spruce and Tsuga.
Roestelia Rebentisch (p. 389)
0. Pycnia spherical or cup-
formed.
iEcia with strongly dethick-walled periveloped
1.
^<^
dium, flask-shaped or cylindric; spores globose,

1-celled,
Fig. 281.
R. pyrata, cups showing peridial

cells.
After King.
brown to yellow, catenulate, with The forms are the ajcial stages
occur mostly on Rosaceous hosts.
several evident
of
germ pores. Gymnosporangiums and

will
The economic forms
be
found under Gymnosporangium.
392
Uredo Persoon
(p.
389)
Spores produced singly on the terminal ends of mycelial hyphae. Germination by a germ-tube which does not produce basidiospores, but enters the host-plant through
the stomata.
These forms are in the main discussed under their telial genera. U. orchidis Wint. and U. satyrii Mass.
are in the leaves of cultivated orchids.
U.tropaeoli Desm. is found onTropseolum; U. arachidis Lag. the peanut; ^'-^ U. aurantiaca
Mont, on Oncidium.2'^ u. autumnalis Diet, on Chrysanthemums in ^^^ and U. kuhnii (Kr.) Nak. on Japan
sugar cane in Java.
The
irregular
Auriculariales (p. 323)
Mycelium septate, forming a gelatinous, and expanded or capitate sporo-
carp;
hymenium
variable, densely beset
ou each segment of which is FiG.2S2.-Variousbasidiaof with basidia, the lower basidiomycetcs, bornc a long sterigma, with its single ^
1,
auricularias
2,
tremellas
with longitudinal divisions; spore.
di4"ed^Sdtasklil"."
The
Auriculariales are mostly saproof little
phytic and
They embrace some fifty species in two families and are chiefly of interest on account of the form of their basidia Fig. 282, which
shows relationship both to the Ustilaginales and to the orders to
follow.
Key
to Families of Auriculariales
1.
Hymenium gymnocarpous Hymenium angiocarpous

Auriculariaceae
2.
Auriculariaceae, p. 392. Pilacraceae.
Key
to Tribes or Genera of Auriculariaceae
Sporocarp arising from a ton-like base of

mycelial threads
I.
Stypinelleae.

Basidia free on the end of ihc hyplue without saccate cell
Basidia subtended
b}^
393
1.
a saccate
auriform
cell
2.
Stypinella, p. 393. Saccoblastia.

Platyglceeae.
Sporocarps crustaceous
S])orocarps
gelatinous,
II.
or
capIII.
shaped
Auricularieae, p. 393.
In tribe
III, Auricularieop, there is
a single genus, Aiiricularia.
Cap more or less cup-shaped or ear-like, jelly-like l)ut firm when wet, horny when dry, the hymenium often veined or folded, but without tooth. The name refers to the cup-like form.
A. auricula-judiae (L.) Schr.
is
is
a very
common
saprophyte which
occasionally parasitic on elder, elm, and mulberry in Europe. In tribe I, few cases of parasitism of any importance are reported.
Stypinella
mompa
(Tan.) Lin.
is
found on the roots of mulberry
in Japan.
Eubasidii
(p.
299)
The Eubasidii represent the higher development of the basidiaThe basidia, fungi and contain the majority of the species. the typical clulj-shaped undivided stalks, bear usually four,
sometimes two,
])or
is
or eight unicellular spores on a like numand are mostly arranged in hymenia. There great diversity in the form and size of the sporophore from
six,
of sterigmata
an almost unorganized mycelial microscopic weft to the large complex structures of the toad stools and puff balls. Conidia and chlamydospores while occasionally present are much less common than in the preceding groups or orders. The cells of the sporophore in many forms investigated are binucleate ^^ in other forms they are multinucleate.
;
origin of the binucleate condition often antedates the formation of the sporophore and may occur far back in the mycelium,
^'" perhaps as far back as the germinating basidiospore itself.^'"' In the basidial layer, however, even of those forms with multi^^'
The
nucleate vegetative cells, the nuclei are reduced to two so that the general statement is permissible that in the hymenial layer From such cells of the Basidiomycetes the cells are binucleate.
two
nuclei
wander into the basidium primordium where they

This
fuse to one, reducing this cell to a uninucleate condition.
394
nucleus

bj^
two mitoses gives
rise to four nuclei
which wander
through the sterigmata into the spores and constitute the four
basidiospore nuclei.
The
significance of this
phenomenon
is
followed
by
division,
which
of fusion in the basidium wide spread and apparently the
Fig. 283. Stages in the development of the basidium (Agaricus); original binucleate condition, followed (E-F) by fusion, and subsequent mitosis N-R, resulting in four spore nuclei. After Wager.
dominant typical phenomenon among the Basidiomycetes including both high forms, Agarics," and low forms, Dacryomycetes,^^ the Uredinales ^^^' ^^' ^^^' ^^^' ^^^ and even the Gasteromycetes (Maire),^^^ is much debated. By some it is regarded as a very much modified type of fertilization, a view to which support is lent by the fact that in some of these fungi, perhaps all, the nuclei multiply by a process of conjugate division. Thus the two nuclei found in the young basidium, although belonging to the same cell may in ancestry be very distantly related.

Key to Orders of
Gelatinous fungi with forked basidia Basidia clavate, undivided
395
Eubasidii
1.
Dacryomycetales.
Hyinenium without stroma,
parasites,
2.
basidia free, strict Stroma usually well developed, fleshy, coriaceous, leathery or woody
Exobasidiales,
p. 396.
Spores arising from basidia which form
a distinct membranous hymenium which is naked at maturity, and

frequently covers the surface of gills, pores or spines (Hymenomycetes)
3.
Agaricales, p. 398.
Spores arising from basidia enclosed in a

definite peridium (Gasteromycetes.) Spores borne in a more or less deli-
quescent gleba which is at enclosed in a peridium, but maturity elevated on stipe

until
first
is
at
4.
Phallales, p. 462.
Spores remaining within the peridium
maturity
Basidia united into a

ular cavities
hymenium
of irreg-
which hnes the walls
Hymenial
cavities
remaining
together in the peridium, their boundaries mostly
disappearing at maturity Fleshy imtil the maturity of the spores, capillitium
none
Fleshy when young, at maturity filled with dust-like
spore-masses mixed with
capillitium (puff balls)
. .
5.
Hymenogastrales.
6.
Lycoperdales,
p. 464.
Hymenial
cavities separating at
maturity from the cup-like
peridium (bird-nest fungi)

Basidia
7.
Nidulariales.
uniformly
distributed
the peridium or forming skein-like masses
through
8.
Sclerodermatales.
396
The Dacryomycetales include forms with a gelatinous sporophore. They are mostly small, inconspicuous saprophytes, common on decaying wood, leaves, etc. The Hymenogastrales are puff-ball forms, and are very numerous and of very diverse structure. None have been reported as parasitic. The Nidulariales is a small order The comprising the curious bird-nest fungi, all saprophytes.
ranean, and not
Sclerodermatales are thick-skirmed puff known to be parasitic.
balls,
mostly subter-
Exobasidiaies
Strictly parasitic, the
(p.
395)
mycelium penetrating the host and usually causing marked hypertrophy; hymenium unaccompanied by fleshy
sporocarp, consisting only of the closely-crowded, clavate basidia which break through the epidermis of the host.
The basidia Vjear four, rarely five or six sterigmata and spores. The spores are mostly curved. Conidia are also found in some species. The basidiospores germinate with a germ tube which produces fine sterigmata and secondary spores capable of budding. The hymenial cells are binucleate, the two nuclei of the basidial cell
This divides mitotically givfusing into one basidium-nucleus. rise to the spore nuclei. ing This order among the basidia fungi is analogous to the Exoascales
among
the ascus fungi.
There are two genera and some
twenty-five species.
Key
to Genera of Exobasidiaies.
1.
Basidia 6-spored; not gall producers Basidia 4-spored; producing galls
2.
Microstroma, Exobasidium,
p. 396.
p. 396.
Microstroma
Niessl. contains only three species of
which
M. album (Desm.) Sacc. is on oak; M. juglandis (Ber.) Sacc. on Juglans and

Exobasidium Woronin
Hicoria.
Mycelium penetrating the host and causing distinct hypertrophy, hymenium subcuticular, erumpent, basidia 4-spored,
spores elongate. There are some twenty species, mostly on
members
of the

Ericaceic.
397
Cultural work ami studies in infection are needed bel)e

^^
fore species can
properly delimited. E. vaccinii (Fcl.) Wor. occurs on Vaccinium vitis idsea, forming large blisters on the leaves,
rarely on petioles and stems, discoloration red or purple. The fungus appears as a
white bloom on the under

surface of the
leaf;
spores
narrowly 1-2 M.
E.
fusiform,
^^
5-8
Richards
who
studied
and E. andromedffi from inoculations

vaccinii
concludes:
"Aside from the form of

the
distortion,
E.
vaccinii
and E. andromedae cannot
The well be distinguished. former can produce the same form of distortion on both
and the
on
not
Gaylussacia and Andromeda latter has been made
to produce a similar growth
Andromeda.
these
differ.
is
Microforms
scopically
do
Fig. 284.
The
natural
conclusion
species of
that these two
Exobasidium are one and the same and the form producing large bag-like distortions on Andromeda should be considered a form of E. vaccinii."
Exohatîdiuni androniedae on Andromeda, showing host cells, mycelium, basidia and spores. After Richards.
E. oxycocci
cinii,
Rost causes
greatf^r
distorting
young twigs and
leaves;
hypertrophy than E. vacspores 14-17 x 30 m;
smaller conidia often present. The mycelium infests the leaves and stems of the cranberry.^" IVIorphologically the species agrees Infection experiments are needed. closely with E. vaccinii.
E. vexans
Mas
^^
causes a serious disease on tea.
E. andromedae
398
Pk., E. rhododendri Cram., E. japonicum shirai and E. peckii Hal. are reported on Rhododendron and Andromeda;
E. azaleas Pk. and several other species on various Rhododendrons; E. vitis Prill, was noted in France on the grape; E. lauri (Borg) Geyl. is on Laurus.
^^
E.
cinnamomi Petch on cinnamon

Agaricales
(p.
in Ceylon.
'' '
395)
" " "
"
This
is
The mycelium grows
a very large order of over eleven thousand species. to long distances over or through the supporting
-c --ta
nutrient
often
me-
dium,
resistant
forming
conspicuous long-1 i v e d
rhizomorphic
~i
strands or sheets, sometimes developing sclerotia
or again
appearing
as a
mere
floccose weft.
The
simple
basidia bear four
spores, in rare cases two, six or eight.
Other forms of conidia are found in some species
and chlamydospores
be borne either ex-
may
Fig. 285. An agaric (Amanita) sporophore showing parts; c, pileus; m, c, striated margin; g, gills; a, annulus; s, stem; v, volva; me, mycelium. After Peck.
w^-
rac
ternally on the sporophore, in the hymenium,
or inside of the sporo-
phore
tissue.
In
the
lowest forms the basidia
arise directly from the mycelium without the formation of any definite sporophore but in most species the sporophore is highly complex, consisting of
large, stalked or sessile,
mushroom, menium. Fig. 286,

stools,
etc.)
lies;
pseudoparenchymatous structures (toadon special surfaces of which, the hycovering

gills
or spines or lining pits or
pores.
The
general relation of the basidia to the
hymenium and
the

is shown in Figs. 285, by the character of the sporophore,
399
sporophore
286.
Families are delimited
distribution of the hymenial
surfaces, presence of cystidia, size
and color of spores, and other more minor points.
l(X3
-4W
In germination the spore pro- /? "^3^^^V>7^'^9c^ duces a germ tube which develops vV^^-\---^^
directly into a mycelium.
In
many ^
is
species
the
young
mycelium
^''*^^ ^-^
îLitTz^^-^
^^-^_?<>)s<
~'
"
conidia-bearing.
Cytologically the group conforms to the general description given on
"^^"""r^Z^"
pages 393, 394.
The
Agaricales are chiefly of in-
terest to pathologists as wood fungi though in a comparatively few in-
stances they are found on herbs.
Upon wood they may do harm.

First,
case death
as root parasites, in which may follow through in-
absorption or anas causes of Second, chorage. heart rots leading to weakness and eventual overthrow of the tree.
Third,
terference with
cambium
as parasites of sap wood, or bark leading to death
of a part of the host
and often
its
Fig. 286.
Cross
section of the
gill
complete In many instances
loss.
draws
its
subsistence
the fungus from host
showing basidia, sterigmata and spores, also a cystidium stretching from one gill to the next.
After Buller.
viewpoint can start their career on a host plant as saprophytes and after attaining a stage of vigorous vegetative growth become truly parasitic. In
cells not actually alive and hence strictly speaking saprophytes. Nevertheless, since their ultimate effect tree is to cause disease or death, from the practical these fungi are pathogenic. Many species, moreover,
they are
upon the
most instances they are wound parasites, which cannot gain

cess to the inner portions of the host
ac-
through uninjured tissue
400

^
.
iS C3
g
tt>
T3.2
o
-a
"H
-0 03 3 a S
3 O O O
^
c o
"I-
J2
83
5 3
-f^
03
:!^ "3
m
q; r;
m G
en
13
o
c8
<^
T3
J7a
Cj
Ml
(-1
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01
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p g O <D S). 0; Ti G c o 03 u. fe n c bc3 . C3 ^ G ^

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fij
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2o3
CO

but must
hail,
401
make entrance
througli
some wound,
as tliose due to
wind, snow, insects, men and other animals, etc., which exposes the inner bark, cambium, sap wood or heart wood without its natural outer
may cause the disappearance of substances,^^^ e. g., Fomes igniarius consumes the tannin, or the mycelium
may
secrete enzymes which penetrate the host to long distances. These may dissolve first one component of the cell, e. g., the lignin, next the
fig. 288. choid of
protecting tissues. Within the tissues the mycelium
Tra
pine
decomposed by
Trainetcs pini. T h o primary
wall dissolved as far as aa; in the lower part the sec-
most
lignified
residue,
the middle
lamella,
re-
sulting in dissolution of the tissue.
In other cases the parts of the cell walls other than the middle lamella are first affected and soon shrink resulting
Fig. 289.
ondary and
ter-
in cracks.
istic color
Some
fungi cause character-
tiary layers are onlj- (A cellulose;

c,
myce-
changes particularly in those cell walls which are rich in carbon. Parasitism in this
lium
e.
holes at d and
tig.
making
After Har-
group growing
is
old
since
on
wood
are
good examples of agarics found as early as in the
Tertiary
period.'*^
These fungi spread to new hosts by spores borne in various ways; by insects (Trametes radiciperda) animals, wind (Polyporus pinicola) etc., or in a purely
vegetative
manner by the mycelium which
in the
rhizomorphs (Armillaria mellea) through the ground to considerable distances.^^ An excellent summary of the early history of our
Fui. 289. Pine tracheid acted
form
of
travels
knowledge
t> BuUer.
of
wood destroying
./
fungi o
is
given by t, j
338
upon by Polyporus
schweiceUulosc hâ^ been extracted
The number
^^'^^
of species of Agaricales
which
afis
^^^^ plants in the
ways mentioned above
walls
chiefly
has""ca^Js'e"d cracks. After Hartig.
very great but in many instances research in this field has not yet revealed the true relation existing between the fungi and the woody plants upon
which
they are found growing; whether they '", occur as parasites or as saprophytes; whether
'
-^
actually injurious or not.
The
species given below are mainly
402
regarded as actually injurious. If more questionable cases were to be included the number would be increased several fold.
Key
to Families of Agaricales
Basidia loosely aggregated on a mold-like or arachnoid base, formed from loose

floccose hyphae Basidia closely aggregated, forming a com1.
Hypochnaceae,
p. 402.
pact layer
Hymenium smooth
Sporocarp effused, resupinate or rarely pileate, usually not fleshy
Sporocarp clavate, the upper portion only sporogenous, usually fleshy.
.
2.
Thelephoraceae,
p. 405.
3.
Clavariaceae, p. 412.
Hymenium variously folded or pitted Hymenium with teeth, tubercles or

tooth-like plates
which are sporo4.
genous
HydnacesB,
p. 413.
Hymenium
lining pores
Pores not easily separating from the

pileus,
which
is
leathery, corky or
commonly punky
5.
Polyporaceae,
Boletaceae,
p. 416.
Pores readily separating from the

pileus
which
is
fleshy
6.
p. 440.
Hymenium
covering
the
surface
of
7.
radiating plates
Agaricacese, p. 442.
Hypochnaceae
indefinite, of loosely floccose hyphae; the basidia clavate, loosely aggregated into an ill-defined hymenium.
Sporophore poorly developed and often
woven
In the simplicity of the sporogenous structures the members group approach the Hyphomycetes from which they are separated only by their sporophores which are of the nature of basidia rather than of ordinary conidiophores.
of the
small family of some half dozen genera and sixty species.

Key
to Genera of Hypochnaceae
403
Spores colorless, smooth, rarely granular Basidia with two sterigmata Basidia circinate
Basidia not circinate Basidia pyriform, beaked Basidia clavate, not beaked
1.
Helicobasidium.
2.
3.
Urobasidium. Matruchotia.
Basidia with 2-4 rarely 6 sterigmata Basidia with numerous sterigmata Sterigmata small
4.
Hypochnus,
p. 403.
5.
Sterigmata large
Spores colored, mostly spiny
6.
7.
Aureobasidium, Pachysterigma. Tomentella.
p.
405.
Hypochnus Ehrenberg.
Floccose or fungoid, rarely thinly fleshy, spreading over the substratum; basidia clavate; spores colorless, smooth or minutely
granular.
This genus which contains half the species of the family,
is
Fig. 290.
H. ochroleucus sporogenous reticulum prior to spore formation. 8 basidia, sterigmata, and spores. After Stevens and Hall.
it
with difficulty distinguished from Corticium from which in the character of its hymenium.
differs
H. ochroleucus N. **~^^ Sporogenous reticulum of a very close, irregular net work of hyphse variable in thickness; basidia scattered,
404
clavate, swollen; sterigmata 4; spores oblong, slightly flattened on the side adjacent to the companion spores, tapering slightly at each end, 4.7-5.8 x 10.5-11.6 n. A migratory mycehum is
present, covering twigs
rhizomorphs
white, later buff,
and leaves with a brown felty growth; about 5.8 n, septate. Sclerotia
are also found.
The long cottony rhizomorphic strands extend along the twigs, up the petioles and in places aggregate to form brown sclerotia,
Fig. 291.
Hypochnus,
Fig.
of
semi-diagrammatic section showing development of hymenium and basidia, with nuclear conditions. After Harper.
292. Mycelium Hypochnus show-
ing clamp connections. After Harper.
which are especially abundant near the terminal buds. On the ^^ leaves Stevens and Hall describe a loose network from which the basidia arise. Fig. 295. The species is found on apple, pear,
*""'
lilac,
quince,
Vibernum and probably other
hosts,
and
is
widely
distributed.
H. cucumeris Frank.
Fungus gray or brown; basidia elongate, bearing 4 sterigmata; ^"' ^^ in 1883. spores ovoid hyaline. Reported on cucumbers H. solani P. & D. is said to be a parasite of potatoes.^^ It is
probably identical with Corticium vagum solani.
See
p. 407.
thp:
fungi which cause plant disease

""
405
live
H. theae Bern, occurs on
tea;
H. filamentosus Pat. on
leaves of CaryophyllaceiE and Amaryllidaceffi in Quito; H. fuciformis (Berk.) McAlp on grasses in Australia. An undetermined species of Hypochnus was studied by Eustace^^ as the cause of rot of stored apples.
its
Artificial inoculations
proved
parasitism, though
it
was unable
to
make entrance through

m-
sound surfaces.
The
spores are hyaline, smooth, usually obovate, 4-5.5 x 2.5-3.5
Aureobasidium Viala
&
Boyer
^^
(p.
403)
or less
basidia
The fungus body consists of delicate, floccose, more webby masses of much-branched, septate, golden hyphse; with numerous sterigmata; spores cylindric.
single species, A. vitis, V.

Italy^^'
&
B., occurs
on grape roots
in
France and
^"
Thelephoraceae
(p.
402)
(rarely fleshy,
Sporocarp leathery or membranous, punky) resupinate or pileate, simple or

spine-like cystidia.
corky or
compound; hymenophore
smooth, warty or wrinkled; basidia numerous, interspersed with

This is a very large family, but of only a few are parasites.
its
eleven hundred species
Key to Genera
of Thelephoraceae
Hj'mcnophore without cj'stidia Hyiueuophore entirely resupinate

Spore membrane colorless Contents colorless
Spores sessile Basidia with 2 sterigmata. Basidia with 4 sterigmata.
1.
Cerocorticium.
Corticium,
]).
2. 3.
406.
Basidia without .sterigmata. Spores stalked
Protocoronospora,p.409.
4.
5.
Michenera.
Aleurodiscus.
Contents colored
Spore membrane colored Hymenophore soon gelatinous.
...
6.
7.
Aldridgea.
Hymenophore fleshy-leathery. ... Hymenophore partially free, shelving

Context of several layers
Coniophora.
8.
Stereum,
p. 409.
406

Context of only one layer
Hymenophore
leathery
Hymenium not ribbed Hymenium almost smooth

with warts
Basidia continuous
or
9.
Thelephora,
p. 410.
p. 411.
Basidia septate
10. 11.
Hymenophore smooth
Septobasidium, Hypolyssus.
Cladoderris.
Beccariella.
Hymenium
with ribs
12. 13.
Ribs becoming warty Ribs with warty spines
Hymenophore not leathery Hymenophore gelatinous-fleshy Hymenophore membranous, rarely fleshy or fleshy-leathery
14.
Phlebophora.
Hymenium Hymenium
exterior
to
the
15.
hymenophore
inside
Craterellus.
the
hysoli-
menophore Hymenophores mostly

tary
Hymenophore
Hymenophore
attached
sessile
or
16.
laterally stipitate
Cyphella.
centrally
17.
Discocyphella.
Solenia.
Hymenophores
grouped
closely
18.
Hymenophore with
cystidia
cell
Cystidia of a single
Cystidia unbranched Hymenophore of a single layer
Hymenophore resupinate Hymenophore laterally

stalked
19.
Peniophora.
Skepperia.
short20.
Hymenophore
Cystidia of several
of several layers.
21. 22.
Hymenochaete,
Asterostroma. Bonia.
405)
p. 411.
Cystidia stellate-branched
cells
23.
Corticium Persoon
(p.
Hymenophore homogeneous in structure, membranous, leathery or fleshy, almost waxy, rarely approaching gelatinous; hymenium

arising
407
immediately from the mycelium, smooth or mimitely warty; basidia clavate, with four sterigmata; spores small, globose or ellipsoid, with a smooth colorless membrane.
genus of some two hundred
fifty species,
mostly wood inhab-
iting.
One
in its sterile
species possesses a mycelium form as a Rhizoctonia.
which has long been known
Corticium
vagum
solani Burt.
^'^''' ^^^
Hymenophore, white when sporing, poorly developed, of loosely interwoven hyphse; basidia short, cylindric or oblong; spores some-
Fig. 293. C. vagum solani Rhizoctonia stage. After Duggar.
FiG.
294.
C.
basidia, spores.
vagum-solani, and sterigmata After Rolfs.
what
elliptic,
often irregular in outline, 9-15 x 6-13
/x.
mycelium (= Rhizoctonia solani Rhizoctonia violacea) turning yellowish with age, and branching approximately at
Sterile
^^
right angles; often forming sclerotia-like tufts with short, broad cells more or less triangular which function as chlamydospores.
to black sclerotial structures, a few millimeters in diameter, consisting of coarse, broad, short-celled hyphse of peculiar and characteristic branching also occur freely, both in nature and
in culture, Fig. 293.
Brown
These
cells
seem capable
of functioning as
chlamydospores.
The hymenophore
consists of a dark
network of hyphse which
changes to grayish-white when sporing. It frequently entirely surrounds the green stems of the host near the ground. The tips of
spores germinate mycelium. The relation which the various Rhizoctonias which have been described on numerous hosts may bear to the one species under
readily, developing into typical Rhizoctonia
the outermost hyphse are sterigmatate.
The
408

is
discussion
needed.
Some
problematic. Much culture and inoculation work is of the various hosts upon which a Rhizoctonia
apparently closely allied to that of Corticium thus far been found in America are
:
vagum
solani
have
Sugar-beet,
radish,
bean, carrot, cabbage, cotton, lettuce, potato, sweet potato, pumpkin, watermelon, garden pea, corn,
Solanum verbascifolium, egg plant, pig-weed, spiny pig-weed, Heterotheca subaxillaris, Richardia, Crotalaria, Cyperus rotundus, Heterotheca lamarckii and Phytolacca decandra,
purslane,
Picea, Pseudotsuga, carnation and alfalfa. mycelium was noted in Europe on potato many years ago; its existence in America has been kno\vn since 1890 (Duggar ^^). Its identity with the genus Corticium was demonstrated in 1904 both Vjy observing the connection between the myceby Rolfs lium and the basidia on young potato plants and by culture of the typical Rhizoctonia stage from the basidiospores. The parasitism of the organism was proved by inoculations made with pure
Pinus
sps.,
The
sterile
'^^
culture
by
Rolfs.
''^
The sterile mycelium (Rhizoctonia) occurs in two forms on the potato, a light-colored actively parasitic form usually somewhat deep in the affected tubers and a darker mycelium growing
superficially
on the host or over the

branching
is
soil.
In
artificial culture
the
manner
of
typical,
the young branches running
nearly parallel to the main thread and bearing slight constrictions

at their bases.
key to the species
in
France
is
given by Bourdot and Gol-
zin.i"
C. laetum (Karst.) Bres.
salmon-colored, soon fading to a dirty-white; which are nodose, septate, irregular, 4-10 ju, basidia clavate, 35-50 x 7-12 /x; spores oblong ovate, subdepressed on one side, hyaline, 10-14 x 6-8 fx. On fig and apple in Louisiana, and in Europe and in the Northern United States on Alnus, and Corylus. It causes the limb blight ^' ^^^ of the fig, gaining entrance through dead twigs. While the fungus is usually a saprophyte, once it gains entrance to the host it follows down the branch, covering it \\'ith its bright salmoncolored fructification and causing sudden wilting and dying of the
first
Plant body at
context, of hyphse

leaves.
409
The cambium
is
spreads rapidly but

in
layer is the seat of the disease. The fungus not a serious pathogen except in rainy periods
midsummer.
C. javanicum (Hen.) S.
causes disease of coffee and tea; ^^ C. comedens C. dendriticum Hen. parasitizes orange stems;
S.
"''
&
(Nees) Fr. occurs on oak as a

S.
&
Syd. injures many Curt, occurs on cacao.
&
parasite; C. zimmermannii '^ C. lilacino-fuscum Berk, tropical trees;
wound
C. chrysanthemi Plow,
is
reported as the cause of death of
cultivated
chrysanthemum
in
England.
Protocoronospora Atkinson
&
Edgerton
(p.
405)
as in Corticium, except for the basidia which bear 4-8 oblong or elliptic spores. P. nigricans Atk. & Edg.^^ forms narrow elongate spots on vetch pods, stems and leaves. Spot, oblique on the pods, 2-5 x 1-2 mm.,
Genus
sessile,
at first white or with a purple border, later black; subhymenial layer subepidermal two or three cell layers thick; basidia clavate, to subcjdindric, 20-30 x 6-8 ix; spores sessile, pale-pink in mass, oblong to subelliptic, hyaline, smooth, gi'anular, continuous, or
1-septate in germination, straight or curved. Ithaca, N. Y., associated with Ascochyta.
Found on vetch
at
Stereum Persoon
leathery or
(p.
405)
woody, persistent, of several layers, Hymenophore sometimes perennial, laterally or centrally attached; hymenium smooth. A genus of about twô hundred fifty species chiefly wood inhabiting, but a few grow in humus. S. hirsutum (Willd.) Pers.
ish; the
Hymenophore leathery, firm, expanded, wrinkled, hairy, yellowhymenium yellowish, smooth. It causes a rot of oak in which the wood appears white-spotted
S.
in cross section.
quercinum Potter ,^^
S. frustulosum (Pers.) Fries,

trees, is
found on oak in Europe. though sometimes found on living confined to dead wood. It causes a speckled rot of oak
is
wood.^^
Fig. 295.
410
S. purpureum Pers. Hymenophore expanded,
leathery, arched, grayish-white; hy-
meniiim smooth, purple. This species is constantly associated with an English and Canadian disease of drupaceous and pomaceous trees, manifest by a
Fig. 295.
Oak
timber rotted by Stereum frustulosum.
The
After
lighter colored, irregular, small bodies are sporophores.
von Schrenk and Spaulding.

silvering of the leaves, death of branches
and
finally of the tree.
The
S.
causal agency of the fimgus has not been fully established.^^

in distribution.
Cosmopolitan
rugosum
Fr. parasitizes the cherry laurel.
Thelephora Ehrenberg
(p.
406)
variable
in
Hymenophore
sessile or pileate,
leathery,
context
similar,
form,
even or more commonly plicate; hymenium confined to the lower surface or extending all over the hymenophore, smooth or uneven, sometimes warty; basidia numerous,
clavate; spores elongate,
membrane often
dull
brown, and granular.
411
T. laciniata Pers. injures various trees by its leathery incrustations, T. galactina Fr.
The
Resupinate, broadly effused, encrusted, smooth, milky in color. root rot on oak is in type much like that caused by Arraillaria
Fig. 29G.
Telephora laciniata.
After Clonicutd.
mellea.
It also causes
States.^''
a root rot of apple trees throughout the
Central
HymenochaetaB noxia Berk, is a practically onmivorous fungus attacking hevea, cacao, tea, dadap, castilloa, Caravonica cotton, bread fruit, camphor, throughout the eastern tropics.
Septobasidium Pat.
(p.
406)
As Thelephora but with septate
basidia.
412

Pat.*'"^
S. pedicillata (Schw.)
Resupinate,
effused,
byssoid,
subcompact,
light
cinnamon-
yellow to white, hymenium smooth. On oak, palmetto, tupelo, apple, etc.
Cosmopolitan.
Clavariaceae
(p.
402)
leathery, cartilaginous or waxy, cylinbranched often quite large and conspicuous hymenium with cystidia; basidia clavate, with 1 to 4 sterigmata;
Hymenophore
fleshy,
dric-clavate, simple or
spores elliptic or fusiform, hyaline. There are about five hundred species.
sitic.
One genus only
is
para-
Key to Genera of
Hymenophore
Basidia with
small, simple
1
Clavariaceae
or 2 sterigmata
1.
Spores colored
Spores hyaline
Baumanniella.
Hymenophore expanded above

cap, basidia with
1
into a
.
.
Hymenophore
sterigma.. clavate, basidia with
2.
Gloeocephala.
Pistillaria.
2 sterigmata Basidia with 4 sterigmata Hymenophore clavate or filiform..
3.
4. 5.
Typhula,
p. 412.
Hymenophore
Hj^menophore
rarely simple
capitate, hollow. .*
large,
Physalacria.
usually
branched,
Hymenophore mostly round, branches

never
leaf-like
6.
Hymenophore fleshy Hymenophore not fleshy Hymenophore cartilaginous or horny. Hymenophore leathery and hairy ... Hymenophore leafdike
Clavaria.
7.
Pterula.
8. 9.
Lachnocladium.
Sparassis.
Typhula graminum Karst. has been reported as injuring wheat. H^^menophore fleshy or waxy, delicate, simple or rarely branched, filiform or cylindric, clavate; spores colorless. Sometimes forming
sclerotia.
Fig. 297.
is
T. variabilis Riess.
regarded as a parasite of beets.
THE
FUx\GI
413
Hydnaceae
(p.
402)
Sporophore variable
in texture, cu-
ticular, leathery, corky, felty, fleshy or
woody;
free
and
stipitate,
shelving
tooth-like
or resupinate; the thorny, spiny or
hymenium warty,
with
plates; basidia usually 4-spored, rarely
1-spored.
Over
very
five
hundred
in
species,
mostly
Fig.
limited
their
geographical
297.
and chiefly epixylous, some are humus-loving. although

distribution,
Typhula
variabilis,
M,
and
habit sketch; n, basidium spores. After Winter.
Key
to
Genera of Hydnaceae
Sporophore annual Hymenium without a subiculum Hymenium with a subiculum.
1.
Mucronella.
Hymenium
with folds or wrinkles

2. 3.
Crest of the folds entire Crest of the folds incised
Phlebia.
Lopharia.
Hymenium
with granules or warts

fleshy
firm,
4.
Granules penicillate, multifid
Hymenoj^hore
KneiflBella.
Hymenophore
Granules simple
not fleshy.
...
5.
Odontia.
Hymenium
granular
porose,
reticulate,
6.
Asterodon.
Hymenium
warts
with obtuse cylindric

7.
Radulum.
Grandinia.
Hymenium
warts
with globose hollowed

8.
Hymenium
with more or
less
subulate
teeth or spines Pileus clavaria-like

Pileus not clavaria-like
9.
Hericium.
Teeth
free, mostly fleshy Teeth rounded.
414

Spores hyaline
10.
Hydnum,
Phaeodon.
p. 414.
Spores colored
11.
12.
Teeth lanimeliform Teeth connected at base, coriaceous Cystidia none

Cystidia present
Sistotrema.
Irpex, p. 415.
13. 14.
Hydnochaete.
Sporophore perennial, punky or woody Upper surface smooth, or sulcate
15.
Upper surface zonatc
16.
Echinodontium, p.415. Steccherinum, p. 416.
Hydnum
Linnaeus
in form, resupinate; pileus, shelving, or
Sporophore cuticular, leathery, corky, woody or fleshy, variable bushy branched; hymenium beset with pointed spines; basidia with 4 sterigmata; spores hya-
line.
The
species of this genus,
between two hundred
fifty
and three
hundred, are mostly saprophytes but a few are true

parasites
on woody plants. H. erinaceus Bul.^^
Cap 5-30 cm.
wide, white,
then yellowish or somewhat brownish, the branches form-
head covered stem fleshy; short and stout, 2-8 cm. long
ing
dense
with
teeth,
and
ing;
thick, or entirely lack-
Fig. 298.
Fruiting body of Hydnum erinaAfter von Schrenk
ceus in a hollow log. and Spaulding.
cm. long, crowded slender, densely spores globose, clear, 5-6 /xteeth
3-10
The name
refers to the ap-
pearance of the head.

It is
the cause of a white rot on
many
deciduous
trees, chiefly
oaks.
filled
The
rotted
wood
is
soft
and mushy.
Numerous
large holes
with masses o^ light yellowish fluffy mycelium occur in the heart- wood. Sporophores are often absent on the rotted tree.
H. septentrionale
Fr.^^
Sporophores in bracket-like clusters, up to 20-30 cm. wide by
415
50-80 cm. long, creamy white in color, texture at first fleshy, becoming more fibrous; pileus often 3 cm. thick, upper surface almost plain, slightly scaly, all pilei united behind, teeth slender,
often 12
mm.
long.
On
sugar maple, beech, etc., causing rot of the heart-wood. H. diversidens Fr." causes white rot of oak and beech in Europe.
H. schiedermayeri
Heuff,*"^ injures
apple trees in Europe.
Irpex Fries
(p.
414)
Sporophore shelving or resupinate, hymenium on the lower side, from the first toothed; teeth firm, subcoriaceous, acute, continuous
Fiu.
li'J'J.
1.
flavus.
H, habit sketch.
After Hennings.
with the pileus, arranged in rows or reticulately, basally widened and lamellate or even fa void; basidia 4-spored. I. fusco-violaceus (Schrad) Fr.^^ is a wound parasite on pine in
Europe.
I.
flavus Klotsch
I.
is
injurious to the Para rubber, cloves
and
coffee;
I.
destruens to tea.
Fr.,
paradoxus (Schrad)
according to Glazan,^ causes timber
rot.
Echinodontium
Similar to
Ellis
&
Everhart
(p.
414)
Hydnum
but differing in perennial habit; pileus,
smooth, woody; cystidia bearing spines. '^ E. tinctorium E. & E.^^' is the only species. Spines brown, 1 cm. long, 13/2~2 mm. broad; cystidia subconic, reddish-brown, 20-30 x 6-7 /*.
416

living trunks of Tsuga,
On
Pseudotsuga
^''^
and Abies
in north-
western North America.
Steccherinum
S. F.
Gray
(p.
414)
Perennial, pileate, sulcate, zonate, radiately subrugose; teeth
wide, irregular.
S. ballouii Banker is the single economic species. Campanulate to subdimidiate, more or less intricate, sessile, decurrent to pendent, 1-4 x 1-5 cm. laterally connate up to 10 cm.;
surface velutinous
when young,
often licheniferous at base, dark
olive-brown, drying gray-brown in older parts and seal-brown in younger; margin obtuse, seal brown; substance thin, 1-2 mm.,
of
two
layers, the
upper harder, somewhat
brittle,
dark brown,
lower softer and lighter colored; hymenium colliculose, goldenyellow, fading to buff or cream; teeth variable, subterete to diform,
confluent, papalloid to elongate, usually obtuse, tips brownish, 1-5 X 0.5-1 mm. irregularly distributed; spores hjâline broadly elliptic to subglobose, 7-7.2 x 5.5-6.5 fj,.
"^ Chamsecyparis in New Jersey .^^ According to Ballou this fungus is devastating the forests of swamp cedar in New Jersey. As it grows only in the tops of the tree and dies with the host, the dead sporophores soon disappearing, it is a species not easily
On
observed.
Polyporaceae (402)
or
Sporophore annual or perennial; context fleshy, tough, corky woody; hj^menium poroid or lamelloid, fleshy to woody, rarely
edible but they are as bracket forms.
gelatinous.
The sporophores are sometimes fleshy, even more commonly hard and woody, occurring
Fig. 310,
on tree trunks.
Key
narrow
form
to Genera of Polyporaceae
Pores reduced to shallow pits separated by

ridges, folds or reticulations.
.
I.
Merulieae,
Polyporeae.
p.
418.
Pores well developed, variable in size and

II.
Sporophore, at least in part gelatinous

Sporophore
417
more
or
less
gelatinous
1.
throughout
Laschia.
Sporophore leathery above, the pores

gelatinous
2.
Gloeoporus.
Sporophore leathery, corky or punky, never gelatinous. Pores minute and rounded or large and angular
Sporophore
ing
resupinate,
never
shelv3.
Poria,
p. 418.
Sporophore normally
Pores
pileate, only ac-
cidently resupinate usuallj'^ small or
medium
sized,
and round
Substance of the pileus not continuing between the pores
Sporophore at
hardening
first fleshy,
then
4.
Polyporus,
p. 418.
Sporophore
annual
from
the
first
leathery or spongy usually

5.
first
Polystictus, p. 426.
Sporophore from the

or
less
more
6.
corky
or
punky,
usually perennial Substance of the pileus continued
Femes,
p. 428.
Pores usually large,
between the pores hexagonal or labyrinthiform rarely bounded
7.
Trametes,
p. 437.
by large plates
Pores hexagonal
Stipe lateral; pores elongate.
Sessile; pores regular
...
8. 9.
Favolus,
p.
439.
Hexagonia.
Pores labyrinthine, or replaced by

plates
Sporophore
sessile
Hymenium
labyrinthine,
be10.
coming irpiciform
Daedalea,
p. 439.
Hymenium
lamellate, not be11.
coming irpiciform
Sporophore stipitate, concentrically furrowed
Lenzites, p. 439.
12.
Cyclomyces.
418

of the tribe Meruliese
is
Merulius lachrymans
violets.
said to parasitize
Poria Persoon
(p.
417)
Sporophore entirely resupinate, often widely extended, the base leathery to punky, pores small, rounded, covering almost the
entire surface.
genus of almost three hundred species.

^^
P. laevigata Fr. causes a white rot of the birch. P. vaporaria (Pers.) Fr. is a wound parasite on coniferous trees
especially
common on spruce and

Pers.^^
fir
causing a brown rot of the sap
wood. P. subacida
.Sporophore effused, determinate; margin
l)ubescent, white; pores minute, subrotund, 2-6 mm. oblique, odor subacrid. common saprophyte on deciduous and conif-
Irregular especially, pine, hemlock, and spruce. form within the diseased wood and become lined with a tough felt of hyphse, yellow on the inner side,
erous
trees
cavities
P. hypolaterita Berk, causes a tea disease in Ceylon.^^ P. vineta Berk, is reported as causing a rot of Hevea in Ceylon.^
Polyporus (Micheli) Paulet
(p.
417)
Sporophore usually annual; simple or compound, rather thick, fleshy, leathery or corky, stipitate or shelving, pores developing
from the base toward the margin. Grading into Polystictus on the one hand and approaching Fomes on the other. There are about five hundred species. P. obtusus Berk."' ^^ Pileus somewhat imbricate, large and spongy, at length indurate, dimidiate, sessile, often ungulate, 5-7 x 10-15 x 3-5 cm.; surface
hirtose, azonate, smooth, sordid-white to or fulvous; margin very thick and rounded, sterile, entire, concolorous; context spongy-fibrous, white, indurate with
spongy-tomentose,
isabelline
age especially below, 1-2 cm. thick; tubes very long, 2-3 cm., white to isabelline within, mouths large, irregular, often sinuous, 1-2 mm. broad, edges thin, fimbriate-dentate to slightly lacerate, white to isabelline, at length bay and resinous in appearance; spores globose, smooth, hyaline, 6-8 n; hypha? hyaline, 6 n; cystidia none.

It causes a heart-rot of living oaks, occurring as a
419
and invading the sap wood when decay also found on black locust.
P. sulpnureus (Bui.) Fr.
fk
is
wound parasite well advanced. It is
11-
/T)
Ti
66, 67, 78, 74, 79, 80 ' ' '

' '
Hymenophore cespitose-multiplex, 30-60 cm. broad; pileus cheesy, not becoming rigid, reniform, very broad, more or less
stipitate,
5-15 x 7-20 x
;
0.5-1 cm. surface finely
tomentose to glabrous, rugose, anoderm, subzonate at times, varying from lemon-yellow to orange, fading out
with age; margin thin,
fertile, concolorous,
subzonate,
finely
to-
m e n to s e,
rarely
lobed;
undulate, context
fragile
cheesy,
very
when
dried
dry, yellow
when
fresh, usually
white in
Fig. 300. Polyporus sulphurous. Scattered fruit bodies on living oak. After Atkinson.
specimens,
homogenous, 3-7
mm.
thick;
tubes annual,
thin,
2-3
mm.
long, sulphur-yellow within;

irregular,
mouths minute, angular,

lacerate,
somewhat
3^
to
a mm., edges very
sulphur-yellow, \\ath color fairly permanent in dried specimens; spores ovoid, smooth or finely papillate, hyaline, 6-8 x 3-5 mIt is
common
trees, conifers
especially
as a cause of red heart-rot of forest and shade and deciduous, and also does damage in the orchard, on cherry, apple and pear, and in the forest to oak, chest-
nut, poplar, maple, walnut, butternut, alder, locust, ash, pine,
hemlock, larch.
is
The decayed wood resembles a mass characterized by radia;! or concentric
of
red-brown charcoal and
forms thin leathery sheets. Round gonidia are often formed within filled with the fungus. the wood.
P. squamosus (Huds.) Fr.
'
cracks in which the fungus In dicotyledons the vessels become
^^"
^^
Sporophore of immense size, reaching 50 cm. in breadth and 3 cm.
420

found
in imbricated
in thickness, iisuall}^
masses projecting from
the trunks of hving trees.
Pileus subcircular
and umbilicate when
young, soon lêcoming flabelliform and explanate; surface ochraceous to fulvous, covered with broad, appressed, darker scales
Fig. 301.
Polyporus squamosus.
After Clements.
which are \evy
close together in
young specimens; margin
in-
volute, thin, entire; context fleshy-tough, juicy, milk-white; very thick, odor strong; tubes decurrent, white or j)ale yellowish, very
mouths large, alveolar, 1 mm. or more in diameter, edges thin at maturity, toothed at an early age, becoming lacerate: spores broadly ovoid, smooth, hyaline, 5 x 12 ju; stipe excentric
short,
to lateral, obese, reticulate above, clothed at the base with short,
421
dark brown or black, velvety tomentum, often reduced, variable

in length.
The mycelium causes white

trees, particularly
rot of nut, ornamental
and
fruit
maple, pear, oak, elm, walnut, linden, willow,
ash, })irch, chestnut, beech,
growing on dead parts of living
trees.
rapidly along the wood vessels and often bear clamp connections. 1 R A beautiful biological study has been publisiied by I'uller
1 '
The hyphae advance most
who states that a single sporophore may produce 11,112,500,000 spores and that "the number produced by a single fungus from a single tree in the course of a year may, therefore, be some fifty
times the population of the globe." He showed the following enzymes to be present in the sporophore:
laccase,
tyrosinase,
amylase,
emulsin,
protease,
lipase,
Pectase, maltase, invertase, trehalase and cytase were not found; It is evident, however, that the myrennetase,
and coagulase.
wood produces cytase and possibly hadromase. P. hispidus Bui. Pileus thick, compact, fleshy to spongy, dimidiate, sometimes imbricate, compressed-ungulate, 7-10 x 10-15 x 3-5 cm.; surface
celium in
hirsute, ferruginous to fulvous, azonate, smooth; margin obtuse, velvety; context spongy-corky, somewhat fragile when dry, ferruginous to fulvous, blackening with age, 1-1.5 cm. thick; tubes
slender,
about
2-3 to a
mm.
1 cm. long, ferruginous within, mouths angular, ferruginous to bay, blackening with age, edges thin,
very
fragile, lacerate;
spores broadly ovoid, smooth, thick-walled,
deep-ferruginous, 2-guttulate, 5-6 x 7-8 /x. It is common on all kinds of deciduous trees, often injuring fruit trees, especially the apple.
P. giganteus (Pers.) Fr. has been reported as injurious to the

oak.
P. glivus Fr.
is
common saprophyte on
deciduous trees and in
some
cases
may
be parasitic.
P. dryophilus Berk.
14 X 2-3 cm.
Pileus thick, unequal, unguliform, subimbricate, rigid, 7-8 x 10surface hoary-flavous to ferruginous-fulvous, becom;
ing scabrous and bay with age; margin thick, usually obtuse, sterile, pallid, entire or undulate: context ferruginous to fulvous,
422
zonate, shining, 3-10 mm. thick; tubes slender, concolorous with the context, about 1 cm. long, mouths regular, angular, 2-3 to a mm., glistening, whitish-isabelline to dark-fulvous, edges thin,
Fig. 302. Decomposition of spruce-timber by Polyporus borealis. a, a tracheid containing a strong mycelial growth and a brownish yellow fluid which has originated in a medullary ray; at h and c the mycelium is still brownish. At d and e the walls have become attenuated and perforated, the filaments delicate; at / the pits are almost destroyed; at g and h only fragments of the walls remain." The various stages in the destruction of the bordered pits are to be followed from i to r; at i the bordered pit is still intact; at k the walls of the lenticular space have been largely dissolved, their inner boundary being marked by a circle; at I one side of the bordered pit has been entirely dissolved; at m. and n one sees a series of pits which have retained a much-attenuated wall on one side only namely, on that which is provided with the closing membrane. In makBetween o and r ing the section a crack has been formed in this wall. both walls of the pits are found to be wholly or partially dissolved, only at p and q has the thickened portion of the closing membrane been preserved; at d the spiral structure of both cell-walls is distinctly recognizable. These walls when united form the common wall of the tracheid; at t hyphse are seen traversing the tracheids horizontally. After Hartig.
entire to toothed; spores subglobose, smooth, deep-ferruginous,
6-7
At;
cystidia scanty
and
short; hyphae deep-ferruginous.
It causes a disease of oaks.
P. fruticum B.
oleander in Cuba.
&
C. occurs on living twigs of the orange and

P. borealis (Wahl.) Fr."'
Pilcus
sessile,
^^
423
subimhricate,
dimidiate to flabelliform,
often
narrowly attached, spongy to corky, very tough, moist and juicy when fresh, 5-8 x 8-12 x 2-4 cm.; surface uneven, soft and spongy,
entire,
hirtose-tomentose, azonate, white to yellowish; margin thin, white, somewhat discolored on drying: context fibrous-coriaceous above, fibrous-
woody below, white, 0.5-1.5 cm. thick; tubes 4-8 mm. long, white to pallid within,
mouths angular, irregular, somewhat radiately elongate, sinuous at times, 1-2 to a mm., stuffed when young, edges thin,
white to ochraceous, dentate to lacerate; spores ovoid, smooth, hyaline, 5-6 x 3-4 /x;
hyphffi 6-7
fi;
cystidia none.
Fig.
rcalis,
303. Polyporus Ik)hymenium with

After At-
pine, spruce, hemlock, balsam pine, as a wound parasite or as a saproetc., phyte on dead trees producing a white rot.
On
sinuous pores.
kinson.
The mycelium
ad-
vances
At certain longitudinally, radially tangentially. it is very abundant and forms cords in the channels formed stages by the fungous enzyme. Later these cords disappear. The yoimg mycelium is stout and yellow, later it is more delicate. Dissolution of the cells begins at the lumen and proceeds outward, the middle lamella
and
persisting last.
P. dryadeus Fr
66
Sporophore very
large, sessile, dimidiate,
rarely circular, usually imbricate, applanate or depressed above, convex below, fleshy to
Fig. 304.
realis,
Polyporu.s
l)o-
hymenium with
After
spongy-corky, rather fragile when dry, 1530 X 25-65 x 3-5 cm.; surface very uneven,
rounded pores.
Atkinson.
azonate,
opaque,
hoary-isabelline,
anoderm
weepshining and bay; margin ing; context thick, zonate, subglistening, ferruginous-isabelline to fulvous, 2.5-4 cm. thick; tubes grayish-umbrinous to fulvous
within, 5-15
to very .thinly encrusted, thick, pallid, entire to undulate,
sub-
mm.
young, becoming
long, slender, very fragile, mouths whitish when somewhat resinous in appearance and finally
424

first
bay-brown, at
minute, circular, becoming angular, 4 to a
thin, fimbriate to lacerate, deeply splitting and with age: spores subglobose, smooth, 9-10 x 7-8 ju, separating the outer wall hyaline, the inner membrane brown; cystidia 15-
mm., edges
35 X 5-9
P.
/x.
It causes rot of
oak wood in America and Europe.

^^^
amarus
Hedg.^^-
Pileus soft
when
old, ungulate, often spuriously stipitate
and spongy when young, becoming hard and chalky from knot-holes, fre-
quently large, 5-11 x 10-20 x 6-12 cm.; surface pubescent when young, rimose and chalky when old, at first buff, becoming tan and often blotched with brown when older; margin obtuse, frequently having an outer band of darker brown, often slightly furrowed; context creamy-yellow to tan-colored, usually darker in
outer layers
when
old,
4-8 cm. thick; tubes not
stratified,
brown
within, cylindric, 0.5-3 cm. in length, shorter next the margin, mouths circular or slightly irregular, 1-3 to a mm., yellow-green during growth, turning bro-Ti when bruised or old, becoming
lacerate; spores hyaline or slightly tinged with brown, smooth,
ovoid, 3-4 X 5-8 n, nucleated; cystidia none. The cause of "pin rot" or peckiness of incense cedar.
P. schweinitzii
Fr.^^'
'^
Pileus spongy, circular, varying to dimidiate or irregular, 15-20 cm. broad, 0.5-2 cm. thick; surface setose-hispid to strigosetomentose and scrupose in zones, ochraceous-ferruginous to fulvous-castaneous or darker, quite uneven, somewhat sulcate, ob-
scurely zonate; margin yellow, rather thick, sterile: context very soft and spongy, fragile when drj^ sometimes indurate with age,
flavous-ferruginous to fulvous, 0.3-0.7 mm. thick; tubes short, 2-5 mm. long, flavous within, mouths large, irregular, averaging 1 mm. in diameter, edges thin, becoming lacerate, ochraceous-
olivaceous to fuliginous, rose-tinted when young and fresh, quickly changing to dark-red when bruised spores ovoid, hyaline 7-8 x 3-4
:
stipe central to lateral or obsolete, very irregular, tubercular or very short, resembling the pileus in surface and substance.
/x:
On coniferous trees especially spruce, fir, pine, larch, arbor vitffi, entering through the root system and extending up the trunk, causing heart-rot. The tracheids exhibit spiral cracks and fissures

due to the slirinking
of the walls.
425
is
Fig. 289.
Diseased wood
dry.
yellowish and of cheesy consistency; brittle P. betulinus (Bui.) Fr."' ^3

Pileus
fleshy
when
to
corky,
compressed-ungulate,
convex above,
plane below, attached by a short
umbo
behind, varying to bell-
shaped when hanging from horizontal trunks, 5-30 x 5-20 x 2-5 cm.; surface smoky, covered with a thin, separating pellicle, glabrous, devoid of markings, cracking with age; margin velvety, concolorous, obtuse, projecting nearly a centimeter beyond the
426

fleshy-tough, elastic, homogeneous, 3 cm. thick,
hymenium: context
milk-white; tubes 0.5 cm. long, 2-3 to a mm., sodden-white, separated from the context by a thin pink layer; mouths very irregular, dissepiments thicker than the pores, obtuse, entire, crumbling
in age, leaving the smooth, white context; spores white, cylindrical, curved, 4-5 n in length. The mycelium penetrates
away
lignified cell walls entering the living cells
and causing death.
On birch
by Fomes
causes a decay of the sap fomentarius.

it
wood
similar to that caused
P.adustus (Wild.)
Fr.
is
common saprophyte of deciduous trees.

(p.
Polystictus Fries-
417)
Sporophore leathery, usually thin; pores developing from the center to the circumference of the hymenophore. The thicker
forms are quite close to some species of Polyporus. About four hundred fifty species.
P. versicolor (L.)
Fr.^''
^^
Pileus densely imbricate, very thin, sessile, dimidiate, conchate, 2-4 X 3-7 X 0.1-0.2 cm.; surface smooth, velvety, shining, marked with conspicuous, glabrous zones of various colors, mostly latericeous,
bay or black; margin
thin, sterile, entire; context thin,
mem-
branous, fibrous, white; tubes punctiform, less than 1 mm. long, white to isabelline within, mouths circular to angular, regular,
mm., edges thick and entire, becoming thin and dentate, white, glistening, at length opaque-isabelline or slightly umbrinous: spores allantoid, smooth, hyaline, 4-6 x 1-2 n;
even, 4-5 to a
hyphae 2-6 ix; cystidia none. Von Schrenk regards this as strictly a saprophyte except when on catalpa, where it causes a heart-rot. It is common on almost
any kind
soft
its action becomes straw-colored and finally and pithy. Both cellulose and lignin are dissolved. P. sanguineus (L.) Fr. & P. cinnabarinus (Jacq.) Fr. are saprophytes on dead parts of live trees.
of wood. Catalpa wood under
P. velutinus (Pers.) Fr.
is
common saprophj^te Avhich is perhaps
sometimes
parasitic.
is
P. occidentalis Klachb.
indicus in the
recorded as a parasite on Pterocarpus
Malay
peninsula.^^

P. pergamenus
Fr.^^
427
Pileus exceedingly variable, sessile or affixed by a short tubercle, dimidiate to flabelliform, broadly or narrowly attached, 2-5 x 2-6 X 0.1-0.3 cm.; surface finely villose-tomentose, smooth, white
or
slightly
yellowish,
marked with a few narrow indistinct latericeous or bay zones; margin thin, sterile, entire to
lobed; context very thin, white, fibrous; tubes 13
mm.
long, white to dis-
mouths somewhat irregangular, 3-4 to a mm., ular,

colored
within,
usually becoming form at an early
irpici-
stage,
edges acute, dentate, becoming lacerate, white to

yellowish
or
umbrinous
Fig. 306.
:
Polystictus pergamenus.
Clements.
After
spores smooth, hyaline. It causes a sap wood rot of practically all species of deciduous trees, often on dead trees, less frequently on living trees which
have been severely injured. In general the rotten that produced by P. versicolor; microscopically the fungus attacks chiefly the lignin.
P. hirsutus Fr.
Pileus confluent-effused,
wood resembles
it
is
seen that
more or less imbricate, sessile, dimiapplanate, corky-leathery, rather thick, flexible or rigid, 3-5 X 5-8 X 0.3-0.8 cm. surface conspicuously hirsute, isabelline to
diate,
;
cinereous, concentrically furrowed
and zoned; margin at length
thin, often fuliginous, sterile, finely strigose-tomentose, entire or
undulate: context white, thin, fibrous, spongy above, 1-4 mm. thick; tul)es white, 1-2 mm. long, mouths circular to angular, 4 to a mm., quite regular, edges thin, firm, tough, entire, white
to yellowish or umbrinous; spores smooth, hyaline, cylindrical,
slightly curved, 2.5-3 n.
It is
wound
parasite of the
Mountain
Ash.^
428
Fomes
Sporoi)hore
sessile,
Fries (p. 417)
ungulate or applanate; surface varnished, encrusted, sulcate, vinose, or anoderm, rarely zonate; context corky to punky; tubes cylindric, stratiose; spores smooth, hyaline
or brown.
A genus of some three hundred species. 78 79 /T F.ignianus (L.) Gill. 55, 67

.
/"I'll
Pileus woody, ungulate, sessile, 6-7 x 8-10 x 5-12 cm.; surface smooth, encrusted, opaque, velvety to glabrous, ferruginous to
Fig. 307.
Fomes
igniarius,
from maple.
After Atkinson.
fuscous,
becoming rimose with age; margin obtuse,
sterile,
fer-
ruginous to hoary, tomentose; context woody, distinctly zonate, ferruginous to fulvous, 2-3 cm. thick; tubes evenly stratified, 2-4 mm. long each season, fulvous, whitish-stufTed in age, mouths
minute, 3-4 to a mm., edges obtuse, ferruginous to fulwhen young: spores globose, smooth, hyaline, 6-7 fx; spines 10-25 x 5-6 lt is the cause of a white heart-rot, is one of the most widely
circular,
vous, hoary
/Li-
distributed forms of
wound
parasites
and occurs on more
species
of broad-leaf trees than
any other similar fungus. Among its hosts are beech, oak, apple, peach, willow, aspen, the maples, birch,
butternut, walnut, oak, hickory, alder.
The
first
sporophores usually appear at the point of
initial

infection.
429
it
The mycelium grows mainly
in the heart
wood but
may gain entrance through the sap wood or encroach upon the sap wood from the heart wood. Its growth may continue after
the death of the host.
In early stages
it
follows the medullary
FiQ. 308.
A dead
rius.
beech tree with sporophores of F. fomentaAfter von Schrenk and Spaulding.
rays.
it
The completely rotted wood is white to light yellow and in the mycelium abounds in the large vessels and the medullary rays. The walls of the affected wood cells are thin and the middle
lamella
is
often wholly lacking, due to solution of the lignin.
F. fomentarius (L.) Fr."' Pileus hard, woody, ungulate, concave below, 7-9 x 8-10 x 3-10 cm. surface finely tomentose to glabrous, isabelline to avellaneous
;
"
430
and
finally black and shining with age, zonate, sulcate, hornyencrusted; margin obtuse, velvety, isabelline to fulvous; context
punky, homogeneous, ferruginous to fulvous, conidia-bearing, 3-5 mm. thick; tubes indistinctly stratified, not separated by layers of context,
3-5
mm. long each season,
avellaneous to umbrinous
to a mm.; circular, whitish-stuffed when young, to avellaneous, turning dark edges obtuse, entire, grayish-white when bruised: spores globose, smooth, very light brown, 3-4 /x;
within, mouths
3^
hyphae brown, 7-8 m; cystidia none. sap
The mycelium kills the cambium and causes a white rot of the wood of deciduous trees, especially beech, birch, elm, maple. The wholly rotted wood is soft, and spongy, light yellow and
its separate fibers. F. everhartii (E. & G.) ^''' ^^ ( = Pyropolyporus prserimosa). Pileus woody, dimidiate, ungulate, broadly attached behind, 6-10 X 6-15 X 3-8 cm.; surface glabrous, slightly encrusted, deeply
crumbles into
sulcate,
in age;
not polished, gray to brownish-black, elightly rimose margin obtuse, covered with ferruginous tomentum, be-
coming gray and glabrous: context corky to woody, repeatedly

zoned, fulvous in dried specimens, 2-3 cm. thick; tubes evenly stratified, 0.5-1 cm. long each season, fulvous, mouths circular,
4 to a mm., edges rather thin, entire, ferruginous to fulvous, glistening, the hymenium becoming much cracked in age: spores
globose, smooth, ferruginous, 3-4.5 larger at the base, 15-25 x 6-10 /x.
(jl;
spines abundant, pointed,
On
black oaks, and walnuts
causing a rot almost indistin-
guishable from that caused by F. igniarius. often grows into the living sap wood.
F.
The mycelium
cameus
Nees."^'
Pileus woody, dimidiate, varying
from conchate to ungulate

x 6-8
x. 0.5-3
often imbricate
and longitudinally
effused, 2-4
cm.;
surface rugose, subfasciate, slightly sulcate, rosy or flesh-colored, becoming gray or black with age; margin acute, becoming obtuse,
undulate; context floccose-fibrose to corky, 0.2-2 cm. thick; tubes indistinctly stratose, 1-2 mm. rose-colored, long each season, mouths circular, 3-4 to a mm., edges obtuse,
sterile,
pallid, often
concolorous; spores ellipsoid, smooth, thick-walled, sub hyaline,

3.5 x 6
Ai.

On
431
spruce and
red cedar and arbor vitae causing pockets, also on dead fir. The cellulose is almost all removed from the
affected cells of the heart wood.
The mycelium
is
scant and
when
pale and with numerous clamps. It extends horizontally through the tracheids, giving off lateral branches. None is found in the sap wood.
young
is
annosus (Fr.) Cke. (=Trametes, radiciperda R. Hartig). Pileus woody, dimidiate, very irregular, conchate to applanate, 10-13 X 5-8 X 0.5-2 cm.; surface at first velvety, rugose, anoderm,
F.
'
light brown, becoming thinly encrusted, zonate, and finally black with age; margin pallid, acute, becoming thicker; context softcorky to woody, white, 0.3-0.5 cm. thick; tubes unevenly stratified,
2-8 mm, long each season, white, mouths subcircular to irregular, 3-4 to a mm., edges rather thin, entire, firm, white, unchanging: spores subglobose or ellipsoid, smooth, hyaline, 5-6 x 4-5 n.
fir and various deciduous trees, described by Hartig^^ most dangerous of all conifer parasites. It is not so plentiful in America as in Europe. The sporophores appear near or on the roots, between the bark scales, where the white felted delicate mycelium also occurs.
On
pine,
as the
spores, carried presumably by rodents, germinate upon the bark of roots; the mycelium penetrates to the living cortex, forces its way into the wood and follows up the stem and down the root. The parenchyma cells are killed and browned; the wood becomes
violet,
The
later
brownish-yellow.
pierce the walls.
lumen and
The hyphse The lignified
travel
in
the
cell-
first, later the middle lamella disappears. root system may become involved and the death of the tree result. F. juniperinus (v. Sch.) S. & Sy}^ Pileus woody, ungulate, 3-5 x 5-8 x 5-7 cm. surface tomentose,
;
parts are dissolved Eventually the whole
deeply sulcate, ferruginous to gray, at length rough and grayishblack; margin obtuse, velvety, melleous or ferruginous to hoary:
context corky to woody, reddish-fulvous,
0.5-1 cm. thick; tubes
indistinctly stratified, 0.5-1 cm. long each season, melleous within, reddish-fulvous in the older layers, mouths circular to angular,
2-3 to a mm., edges rather thin, entire, even, melleous: spores

reddish-brown, smooth; spines blunt, only slightly projecting. On red cedar.
432
In the holes caused by the fungus in the heart-wood is found a velvety mass of reddish-yellow mycelium, glistening with colorless liquid and holding masses of reddish-brown wood fiber. Long white fibers of cellulose with the lignin removed project into the
cavities
from the ends.
Structural change begins soon after the mycelium enters a cell lumen. The primary lamella becomes granular and is dissolved a lignin-splitting enzyme, the secondary lamella becomes white by
and the
cells fall apart.
The mycelium
in all directions.
in
extends lengthwise.
newly invaded tissue is nearly hyaline and Within the tracheids branches are given off
is
The sporophore appears after decomposition advanced.
considerably
F. laracis (Jacq.) Murr.^^^ Pileus firm, at length fragile, ungulate to cylindical, 3-8 x 5-10 X 4-20 cm.; surface anoderm, powdery, white or slightly yellow-
concentrically sulcate, becoming slightly encrusted, tuberculose and rimose; margin obtuse, concolorous: context soft, tough, at
ish,
length friable, chalk-white or slightly yellowish, very bitter, with the odor of fresh meal, 1-3 cm. thick; tubes evenly stratified, concolorous, 5-10 mm. long each season, mouths circular to
angular, 3-4 to a mm., edges thin, fragile, white, l)ecoming discolored and lacerate, wearing away with age spores ovoid, smooth, hyaline, 4-5 ix; hyphse 5 m; cystidia none.
:
A wound
America.
parasite of the larch, pine
and spruce
in
Europe and
F. ribis (Schw.) Gill.89 Pileus tough, corky, Ijecoming rigid, conchate, laterally connate, 3-5 X 5-10 X 0.7-1.5 cm.; surface rough, velvety, anoderm, indistinctly zoned,
and
slightly encrusted with age;
ferruginous to umbrinous, becoming glabrous margin undulate to lobed, fer-
ruginous,
furrowed:
context
punky, fulvous,
3-5
mm.
thick;
tubes indistinctly stratified, 1-2 mm. long each season, fulvous, mouths circular, 5-6 to a mm., edges rather thin, entire, ferruginous
to fulvous,
hoary when young: spores globose or subglobose, pale yellowish-brown, smooth, 3-4 x 3 /i; hyphse 2.5 m; cystidia none.
THE
This
I'UNGl ^^1IICH
CAUSE PLANT DISEASE
433
is a wound parasite on the heart wood of sassafras and is found on roots and stems of various shrubby plants including The fungus fills the large vessels rose, gooseberry and currant. and tracheids with a brown mycelium and dissolves the entire
also
wall locally. F. fulvus (Scop) Gill.'^^ Pileus wood}^ triquetrous, rarely ungulate, thick and broadly attached behind, 1-3 x. 5-7 x 3-8 cm.; surface smooth, very
slightly
sulcate,
velvety,
ferruginous,
becoming
homy and
glabrous and finally nearly black with age; margin subobtuse, ferruginous, velvety; context woody, fulvous, 1-2 cm. thick;
tubes evenly stratified, 2-3 mm. long each season, fulvous, mouths circular, 3 to a mm., edges obtuse, entire, ferruginous to fulvous;
spores globose, compressed on one side, hyaline, 5.5-6 x 4.5-5 n; spines fulvous, 15-20 x 7-9 /x; hypha; 2.5 On plum, birch and other trees.
/x.
The decayed wood

F. fulvus oleae Lin. F. nigricans
Fr.^"'
is
is
is
red-brown and crumbles when crushed. injurious on olive in Italy. very similar to F. igniarius from which it
differs chiefly in the
black upper surface and the bluish or blackish ^^ hymenial surface of the sporophores. Murrill regards it as a
it
variety of F. igniarius. As a wound parasite
causes a reddish-brown heart-rot of
deciduous
trees, especially of willow, birch, poplar, beech.
F. lucidus (Fr.) Bon. causes a cocoanut root-rot. F. fraxinophUus (Pk.) Sacc.^^' ^^
Pileus woody, subtriangular, compressed-ungulate, usually decurrent, 5-10 x 6-12 x 2-4 cm.; surface white, pulverulent or finely tomentose, concentrically sulcate, becoming gray or black
and rimose with age; margin tumid, white or yellowish, velvety to the touch; context corky to woody, zonate, isabelline, 0.5-1 cm.
mm. long each white when young, concolorous with the context in the season, older layers, mouths white, subcircular, 2 to a mm., edges obtuse;
thick; tubes evenly but indistinctly stratified, 2-4
spores broadly ellipsoid, smooth, hyaline, thin-walled, 6-7 x 7-8 fx; hyphse light yellowish-brown, 10-12 fi; cystidia none.
It causes a heart-rot of
trunk and branches of species of ash.

is
The
starch in the host cells
lost early
by
diastatic action in
434
advance of the fungus, the nearest hyphae of which may be several milUmeters distant, and is replaced by a decomposition product. The mycelium advances through the medullary rays and spreads
through spring and summer bands, abstracting the lignin; the middle lamella dissolves and the cells fall apart. Completely rotted wood is straw-colored, very soft, non-resistant. The young hyphae are very fine and require an immersion less for observation. Clamp connections are frequent. The sporophore appears after the destruction of the wood is considerably advanced. F. hartigii All. is very similar to,, if not identical with,
F. igniarius. It produces a white rot of
firs
and spruces.
The mycelium
is
yellowish with numerous branches which may fill the cavities of the bordered pits of the tracheids. The middle lamella is eventually dissolved, later the irmer walls. F. robiniae (Murr.) S. & Sy. ^^' ^^
A large fungus with dark rimose surface and tawny hjonenium. Pileus hard, woody, dimidiate, ungulate to applanate, 5-25 x 5-50 X 2-12 cm.; surface velvety, smooth, soon becoming very rimose
and roughened, fulvous to purplish-black, at length dull-black, deeply and broadly concentrically sulcate; margin rounded,
context hard, woody, concentrically banded, 1-3 cm. thick, fulvous; tubes stratose, 0.15-0.5 cm. long, 50 a mm., fulvous, mouths subcircular, edges entire, equaling the
velvety, fulvous;
tubes in thickness: spores subglobose, smooth, thin-walled, ferruginous, copious, 4-5 At; cystidia none. On black locust causing heart-rot, arising from wound infection
The very hard wood becomes a soft, yellow to brown mass, spongy when wet. The decay extends out in radical
of living trees.
lines
from the
center, along the large medullary rays, killing the

lignin is first dissolved,
cambium and bark on reaching them. The

later the cellulose.
F.
The fungus ceases growth on the death of its host. marmoratus Berk. (=F. fasciatus [Sw.] Cooke.)
Pileus hard, woody, dimidiate, applanate to ungulate, convex above, 7-10 x 8-15 x 2-6 cm.; surface finely tomentose, at length
glabrous, concentrically sulcate, at first mole-colored, changing to umbrinous, and finally avellaneous with black fasciations;
435
margin acute to obtuse, isabelline, sterile, undulate or entire; context punky, thin, ferruginous to fulvous, zonate, 3-5 mm. thick, tubes indistinctly stratified, 5-10 mm. long each season, avellaneous within, mouths circular, minute, 4-5 to a mm. edges
obtuse avellaneous to umbrinous, becoming darker
spores subglobose, smooth, light brown, 5-7 cystidia none.
/x;
when
bruised:
hyphae brown, 4-6/i;
On
water oak and orange in Florida,^ especially abundant on
the former.
F. sessilis (Murr.) Sacc.
A variable fungus with wrinkled varnished cap and acute margin, found on decaying deciduous trees. Pileus corky to woody, dimidiate, sessile or stipitate, imbricate or connate at times, conchate to fan-shaped, thickest behind, thin at the margin, 5-15 x 7-25 X 1-3 cm.; surface glabrous,
laccate,
shining,
radiate-
rugose,
cate,
concentrically sulreddishyellow to
chestnut, at length opaque,
dark-brown usually marked

near the margin with alter-
bay and tawny zones; margin usually very thin and acute, often curved
nating
rarely
downward, often undulate, becoming truncate,

:
white, at length concolorous context soft-corky or woody,

radiate-fibrous,
cally
concentri-
banded, ochraceoustubes 0.52 cm. fulvous;

long, 3-5 to a
mm., brown
circular
Fig. 309.
of
F. pinicola growing on dctul trunk

After von Schrenk.
western hemlock.
within,
mouths
or
angular, white or grayish-brown, edges thin, entire: spores ovoid, obtuse at the summit, attenuate and truncate at the base, verrucose, yellowish-brown, 9-11 x 6-8 n; stipe laterally attached,
usually ascending, irregularly cylindrical, 1-4 x 0.&-1.5 cm., resembling the pileus in color, surface and substance, often obsolete.
436
It

occurs on oak and maple
^^
as a
wound
parasite, destroying
bark
and cambium.
This
and
related
species
are
usually
saprophytic. F. pinicola (Fr.) Cke.^^ Pileus corky to woody, ungulate, 8-15 x 12-40 x 6-10 cm.; surface glabrous, sulcate, reddish-bro^Ti to gray or black, often
resinous; margin at first acute to tumid, pallid, becoming yellowish or reddish-chestnut: context woody, pallid, 0.5-1 cm. thick; tubes
distinctly stratified, 3-5
mm.
long each season, white to isabelline,
Fig. 310.
Fomes applanatus.
After Clements.
mouths circular, 3-5 to a mm., edges obtuse, white to cream-colored; spores ovoid, smooth, hyaline, 6 n; hyphse 8 /x; cystidia none. It occurs on conifers; pine, hemlock, spruce, balsam, larch, etc.,
more rarely on beech, birch and maple, as a wound parasite of The sporophores are often absent until after the heart wood.
death of the host.
The
tracheids bear
many
holes.
The wood
carbonizes, the cellulose is destroyed and sheets of mycelium form, particularly \^^thin the space occupied by the medullary rays and
Fig. 309. F. applanatus (Pers.) Wallr. Pileus hard, woody, dimidiate, applanate, 6-15 x 8-30 x 1-4 cm.; surface milk-white to gray or umbrinous, glabrous, concentrically
in tangential crevices.
'
"
sulcate,
encrusted,
fasciate with
ol:)scure
lines,
condia-bearing,

usuall}'
437
brownish during the growing season from the covering of conidia; margin ol)tuse, broadly sterile, white or slightly eremeous,
entire to undulate: context corkj^ usually rather hard, zonate, fulvous to bay, 5-10 mm. thick, thinner with age; tubes very
evenly stratified, separated by thin layers of context, 5-10 mm. long each season, avellaneous to umbrinous within, mouths circular,
5 to a mm., whitish-stuffed when young, edges obtuse, entire, white or slightly yellowish to umbrinous, quickly changing color
when
bruised: spores ovoid, smooth or very slightly roughened, pale yellowish-bro'WTi, truncate at the base, 7-8 x 5-C ju^^ It is described by Heald as the cause of rot of both heart and
sap wood of living cotton-wood trees. The invaded medullary rays first lose their starch by digestion. Next the lignin is dissolved, then the cellulose.
Von Schrenk regards this fungus as a saprophyte since it grows usually only on outer sap wood that is dead and so far as he observed, it does not cause a true disease.
F. ulmarius Fr. is injurious to elm. F. semitosus Berk, causes root rot of
F. australis Fr.
is
Hevea
in India.
wound
parasite on Acacia in Ceylon.

Fries (p. 417)
Trametes
Sporophore annual, rarely perennial, sessile; context homogeneous, coriaceous to corky, extending between the tubes, which
are circular or irregular. There are about one hundred forty-five species. T^:^: /T'U \ T? 66, 74, 78, 79 . pini ( 1 hore) hr.
'
Pileus hard, woody, typically ungulate, conchate or efTusedreflexed in varieties, often imbricate, 5-8 x 7-12 x 5-8 cm., smaller
in varieties; surface
very rough, deeply sulcate, tomentose, tawnybrown, becoming rimose and almost black with age; margin rounded or acute, tomentose, ferruginous to ta\vny-cinnamon, entire,
specimens: context soft-corky to indurate, homogeneous, ferruginous, 5-10 mm. thick, thinner in small specimens; tubes stratified, white to avellaneous within, becoming ferrugisterile in large
nous at maturity and
in the older layers, 5
mm.
long each season,

circular or
much
shorter in thin specimens, mouths irregular, daedaleoid, often radially elongate, averaging 1 to a
mm., edges
438

when young,
at
ferruginous to grayish-umbrinous, glistening

thin,
rather
entire;
spores
subglobose, smooth,
hyaline
maturity,
becoming brownish with age, 5-6 x 3-4 ii) spines abundant, short, 25-35 X 4-6 /i. It occurs on pine, spruce, larch, hemlock, and fir as a wound parasite of the heart wood; it is also on willow in Europe and
America.
The
faces,
'
spores are wind-borne and, lodging on unprotected sur-
develop a mycelium which grows both up and down, spreadmost rapidly in a longitudinal direction, or horizontally following an annual ring. The fungous enzyme first dissolves the lignin leaving the individual tracheids free and of nearly pure cellulose.
ing
cellulose is later dissolved, resulting in holes in the wood. found on most of the conifers of the United States as a saprophyte. The wood becomes white-spotted. In late stages of decay the entire wood is full of small holes which are lined with a white
It is
The
fungous felt. T. ribinophila Murr.
is
perhaps a parasite on the black locust. T. these Zimm. cause a rootrot of tea in India.^
T. suaveolens (L.) Fr.

Pileus
large,
subimbricate,
dimidiate, sessile, convex above, plane or concave below, 4-6 x 5-12 x 1-3 cm. surface smooth,
;
anoderm, azonate, finely villosetomentose to nearly glabrous,

white to pale-isabelline
thick,
sterile,
;
margin
context
entire:
white,
3ii.~Favoius europseus. ^^^'^^^y^-
punky-corky,
1-2
cm.
fresh.
thick, very fragrant

Fig.
when
with the odor of anise; tubes 515 mm. long, white within,
mouths
circular, 2 to a mm., edges at first very thick, white, entire, becoming thinner and often blackish with age: spores oblong-ovoid,
/x;
subsinuate, smooth, hyaline, 8-9 x 3-5 none.
hyphse 7
p.;
cystidia
On
willow.
THE FUNGI WHICH
CAUiSE
PLANT DISEASE
439
Favolus Fries
(p.
417)
Sporophore leathery, fleshy, or coriaceous, laterally stipitate; hymenium with large elongated pores which may even become
lamellate, Fig. 311.
A
is
genus of some seventy species.
F. europaeus Fr. is a European parasite of fruit also common in America.
and nut
trees
it
Daedalea Persoon
(p.
417)
epixylous, usually large and annual, sessile, to ungulate; surface anoderm, glabrous, often zonate: applanate context white, wood-colored or brown, rigid, woody, tough or
Hymenophore
punky: hymenium normally labyrinthiform, but varying to lamellate and porose in some species: spores smooth, hyaline. About seventy-six species. Fig. 312. D. quercina (L.) Pers.
Pileus
corky,
rigid,
dimidiate,
in section,
sessile,
imbricate,
applanate,
;
convex below, triangular

isabelline-avellaneous
to
6-12 x 9-20 x 2-4 cm. surface cinereous or smoky-black with age,
slightly sulcate, zonate at times, tuberculose to colliculose in the
margin usually thin, pallid, glabrous; context homogeneous, 5-7 mm. thick; tubes labyrinthiform, becoming nearly lamellate with age in some specimens,
older
portions;
isabelline, soft-corky,
1-2 cm. long, 1-2 mm. broad, chalk-white or discolored within, edges obtuse, entire, ochraceous to avellaneous. Common on oak and chestnut, often on living trees but
growing only on the dead wood.

Lenzites Fries
(p.
417)
Hymenophore small, annual, epixylous, sessile, conchate; surface anoderm, usually zonate and tomentose: context white or brown, coriaceous, flexible; hymenium lamellate, the radiating
gill-like
dissepiments connected transversely at times, especially

Fig. 313.
when young: spores smooth, hyaline. About seventy-five species.
440

on
firs.
L. abietina (Bui.) Fr. occurs
L. sepiaria (Wulf.) Fr.
has been reported as a parasite on pine, spruce, etc., but recent work of Spaulding ^^ shows it to be merely a saprophyte. L. cor-
rugata Klot, L. vialis Pk. and L. betulina
(L.) Fr. are
common
F:g. 312.
Daedalea quercina on oak.
After von Schrenk and Spaulding.
s9.prophytes on deciduous trees; perhaps also parasitic; L. varie-
gata Fr. on beech and poplar.
Boletaceae
Sporophores
fleshy,
(p.
402)
capitate,
rarely actually sessile;
hymenium on the under
centrally or laterally stipitate, surface only, of
tubes which separate readily from the pileus and are united to each other or only closely approximated.
A family of less than three hundred species.

Key
Pores separate tubes
to Genera of Boletaceae
I.
Pores adnata to each other
Boletineae.
Fistulineae.
Fistulina, p. 441,
II.
Sporophore more or
less fleshy
1.
Sporophore leathery Tubes with a central papilla Tubes without papillae
2.
3.
Theleporus. Porothelium.
441
Fistulina
BuUer
(p.
440)
Sporophore
face at
first
flesh}',
laterally
short-stipitate,
hymenial
sur-
granular, then each granule ])ecoming a tube; these are approximate but not united; spores browTi in mass. Fig. 314.
A genus of a half dozen species.
Fig. 313.
Lenzites betulina.
After Clements.
F. hepatica (Scha.) Fr.
Cap 8-20 cm.

shelf-like,
more
wide, bright-red or red-brown, liver-shaped to or less lobed, smooth, more or less sticky when
wet; flesh containing reddish fibers; stem short, lateral and almost wanting, or sometimes long excentric; tubes pale to yellowish or
442
pinkish;
It is

spores
yellowish
to
pinkish,
ellipsoid,
5-7 x 3-4
fx.
found on oak and chestnut.
Agaricacese
Sporophore usually
(p. 402)^^
fleshy, rarely coriaceous or leathery, stip-
itate or shelving; stipe variable in
development, lateral or central,
Fia. 314.
Fistulina hepatica.
After Clements.
annulate or not, the entire young sporophore often volvate at first; hymenium lamellate, the lamellae usually free, rarely anastomosing, sometimes dichotomous, rarely reduced to ridges or
slight folds.
A family of over twelve

Key
Hymenium
like,
hundred
species.
to Tribes of Agaricaceae
with the lamelliB ridge or foldI.
imperfectly developed
Cantharelleae,
p. 443.
Hymenium with normally

Lamellse sometimes
developed
gills
anastomosing, forming meshes Lamellse not anastomosing

(in
and
II.
Paxillese.
Lamellae and often the cap deliquescent
Montagnites withering,)
III.
Coprineae.
Lamellse not deliquescent Lamellse thick and fleshy, becoming
waxy
IV. Hygrophoreae.

Lamellae not fleshy or waxy Substance of the pileus of two kinds of hyphse, one thick,
443
tubular and in bundles, the

other
thin,
single
and
fre-
quently lactiferous Substance of the pileus of only one kind of hyphae
V. Lactarieae.
Sporophore at maturity leathery

or corky, persistent, rarely fleshy
Lamellae
at
maturity
split
lengthwise Lamellae at maturity

sphtting
VL
not
Schizophyllese,
p. 444.
VII. Merasmieae, p. 445.

fleshy,
Sporophore at maturity
finally putrescent
VIII. Agariceae,
p. 448.
Cantharelleae
This tribe
ceiB.
is
(p.
442)
The hymenial characters
characterized by its low ridge or fold-like lamellae. indicate an approach to the Thelephora-
Key to Genera
Hymenium on the upper side Hymenium on the under side Hymenium with thin veins
Veins anastomosing Veins not anastomosing
of Cantharelleae
1.
of the pileus. of the pileus.
Rimbachia.
2.
Campanella.
Arrhenia.
3.
Hymenium
with thick folds

of
Substance
the
pileus
leathery,
4.
tough Substance of the

leathery
Trogia,
p. 444.
pileus
thin,
soft-
Sporophore
sessile,
at
first
saucer5.
shaped
Sporophore
laterally
Leptopus.
stalked,
fan6. 7.
shaped Substance of the pileus fleshy
Leptoglossum.
Cantharellus.
444
THE
FUxNGI
Trogia Fries
(p.
443)
persistent, sessile, irreg-
Cap
thin, leathery, or
^^^_
membranous,
ular;
folds,
'
hymenium
of
branched
//^F^'^'^^^^^^^^^S^
the branches chiefly mar-
jfc^^^ffj^^^^;^^^^^-
There are some seven
species.
^^^^^'^[(r-:;^::;^^SmMji^^^^^
wf^ "-%.g^^FiG.
::^-ag:rTss^^-^^<
Sporophore 1-2 cm. broad, beaker-formed or irregular margined; ^ yellow or Orange without. ^ ^ rarely whitish, with hne hairs;
' '
.
315. Trogia faginea. After Henmngs.
ribs
concolorous, dichotomous;
colorless.
spores
cylindric,
x 1-1.5
/x,
smooth,
It injures birch, beech, hazel, etc.
Schizophylleae
(p.
443)
cleft gills
Sporophore, leathery, persistent, the

margins.
species.
group
of
but four genera and
less
with recurved than a score of
Key
Sporophore leathery,
Stipe central
to Genera of Schizophylleae
1.
sessile
Schizophyllum,
p.
444.
Sporophore fleshy or membranous, stipitate
Cap Cap
thin,
membranous
2. 4.
fleshy
Rhacophyllus. (Edemansiella.
Pterophyllus.
Stipe lateral
3.
Schizophyllum Fries
Cap woolly, upturned, sessile, epixylous; gills cleft, the gins recoiled; texture leatller5^ About twelve species. Fig. 316.
S. alneum (L.) Schr. Cap 1-4 cm. wide, white
mar-
or gray-woolly, upturned, attached
excentrically, irregularly saucer-shaped* to purplish spores subglobose, 2-3 fx.

;
stem lacking;
gills
grayish

It parasitizes
445
sugar cane, horse chestnut, chestnut, mulberry
and orange.
Fig. 31G.
S.
alneum.
After Clements.
Marasmieae
Pileus tough, leathery, thin,
fleshy, reviving after
(p.
443)
membranous, or rarely somewhat
drying with the return of moisture.

fifty species.
About
five
hundred
Key to Genera
Gills leathery-horny; spores black Gills leathery; spores h3'aline
of Marasmieae
1.
Anthracophyllum.
Pileus not distinct from the stipe; sporo-
phore trumpet-shaped
Gills forked,
Gills
edge blunt with a thin edge Gills toothed on the margin

with an even margin
2.
Xerotus.
3.
Lentinus,
p.
445.
Gills
4.
Panus,
p.
446.
Pileus distinct from the stipe
Annulus wanting Pileus firm and dry

Pileus
5.
6. 7.
Marasmius,
Heliomyces.
p. 446.
somewhat gelatinous
Annulus present
Merasroiopsis.
Lentinus Fries
Sporophore trumpet-shaped,
leathery,
About
pileus and stipe not distinct, central or lateral, gills toothed; spores white. pileus three hundred forty species.
446

(Bui.)
L. conchatus
L. lepideus
^^^
Schr.
is
found on birch, poplar, aspen.
Fr.
on
pine, birch, etc.
447
Fig. 318.
Marasmius plicatus.
After Fulton.
448
plicatus Wak. Pileus submembranous, convex or subcampanulate, glabrous, sulcate-striate, chestnut or light wine-colored; gills rather distant,
white,
M.
downy
basally attached; stipe slender, below.

^'^^
glabrous above,
white
Wakker
plicatus
iana.^"^
Marasmius parasitism of sugar cane was first described by ^^ in 1895 later by Howard. In these cases M. sacchari
it
or varieties of
were
identified.
In 1908 Fulton described
M.
Wak.
as the cause of serious sugar cane troubles in LouisThis fungus which exists first as a saprophyte resides
soil
primarily in the
from which
it
grows over the stools and
eventually penetrates living tissue, destroys many roots and smothers the developing buds. The white mycelium is found
cementing the lower
leaf sheaths to the cane.
It is
probable that
several species are concerned.
M. M.
sacchari
Wak. occurs on sugar cane

is
hawiiensis Cobb,
in the oriental tropics. as associated with the preceding reported
species in Hawaii. M. semiustus B. & C. affects the stems, peduncles and inflorescence of the banana. M. equicrinis MilU. Banc,
causes horse-hair blight of cacao and

lar disease of the tea plant
M. sarmentosus
Fr. a simi-
and
of forest trees in India.
Agariceae
This tribe contains
all
(p.
443)
is
the
gill
fungi and
characterized
by a
fleshy, putrescent sporophore; gills weak, easily broken, not deliquescent, without milky juice. It is The genera are conveniently the largest tribe of the family. as black, brown, rusty, pink or red, and white-spored grouped
fleshy, rarely tough or leathery,
forms.
None
of the black-spored species are
known
as parasites.
Amaurosporeae (brown-spored
series)
With a volva
at base
1.
Chitonia.
Without a volva.
Veil remaining on the stem as
Gills free Gills
an annulus
2. 3.
from the stem united with the stem
Agaricus.
Stropharia.

Veil remaining attached to the margin of
449
the pileus, often not apparent in very

old specimens
Veil inconspicuous or wanting
Gills free
Gills
5. G. 4.
Hypholoma,
Pilosace,
p. 450.
decurrent
Deconica.
Gills
adnata or sinuate
7. 8.
Margin of pileus incurved when young Margin of pileus always straight. ...
Phaeosporeae
(Rusty-spored
Psilocybe, Psathyra.
p. 451.
series)
Annulus continuous
Veil single, forming the annulus
\'cil
1.
Pholiota,
p. 452.
forming the annulus and deciduous scales on the pileus

double,
2.
Rozites.
Annulus arachnoid, filamentous or evanescent, often not apparent in old speci-
mens
Gills
adnate; terrestrial.
..
3. 4.
Cortinarius.
Gills decurrent; epiphytal
Flammula,
p. 452.
Annulus wanting
Gills decurrent; stipe
with a cartilaginous
5.
rind
Gills
Tubaria.
not decurrent
Stipe fleshy
Without a volva
Pileus fibrinous or silky Pileus smooth and viscid
6. 7.
Inocybe.
Hebeloma.
Locellina.
With a volva
Stipe with a cartilaginous rind
8.
Margin of pileus incurved young Margin of pileus straight

Pileus viscid;
gills free
gills
when
9.
Naucoria.
PluteoluSo
'
10.
11.
Pileus not viscid;
attached.
..
Galera.
Rhodosporeae
Stipe lateral
(Pink-spored
series)
12.
Claudopus.
Volvaria,
p. 452.
Stipe central.
Volva present; annulus wanting
13.
450

14.
Volva absent; annulus present Volva and annulus both wanting Gills free from the stipe Gills adnate or sinuate
Stipe fleshy Stipe with a cartilaginous rind Pileus torn into scales
Pileus papillose, subcampanulate
Gills decurrent
.
Annularia.
Pluteus,
15.
p.
45
1.
IG.
Entoloma.
Leptonia.
17.
18.
Nolanea.
Clitopilus.
on the stipe
19.
Stipe fleshy Stipe with a cartilaginous rind
20. Eccilia.
series)
Leucosporeae (White-spored
Stipe lateral, or none Stipe central
21.
Pleurotus,
p. 454.
Volva and annulus both present Volva present; annulus none Volva absent; annulus present Gills free from the stipe
Gills united to the stipe
22. 23.
Amanita.
Amantiopsis.
24. Lepotia.
Pileus usually smooth Pileus floccose
25. Armillaria, p. 455.
26. Costinellus.
Volva and annulus both absent Gills decurrent on the stipe

Stipe fleshy
Stipe with a cartilaginous rind
Gills adnate, stipe
27. Clitocybe, p. 457.
28.
Omphalia.
CoUybia,
p. 458.
with a cartilaginous
29.
rind
Gills sinuate
Stipe fleshy
Stipe with a cartilaginous rind Pileus membranous, more or less
striate
30.
Tricholoma,
p.
460.
31.
Mycena,
p. 460.
Pileus very thin, without a pellicle
32. Hiatula.
Hypholoma
Fleshy;
gills
Fries (p. 449)
up
into fragments
attached; annulus imperfect, or none; veil breaking which are more or less persistent on the margin
species.
of the cap.
About seventy

H. appendiculatum
(Bui.) Karst.
is
451
perhaps parasitic, occurring
at the bases of living trees. H. fasciculare (Huds.) Fr,
is
said to
grow
parasitically
upon
Fig. 319.
Hypoloma
rot.
^'^^
appendiculatum.
After Clements.
roots, causing a
white
It is
mentioned as a parasite on raspalso possibly parasitic
berry roots in Australia. H. lateritium (Scha.) Schr.
is
on
trees.
Psilocybe Fries
(p.
449)
Pileus smooth, margin at first incurved; gills and spores at length brownish or purplish; stipe cartilaginous, hollow or stuffed, veil absent or rudimentary.
About forty species. Fig. 320. P. spadicea (Scha.) Fr. is a weak
wound
parasite
on various
to'
woody
plants.
P. henningsii Jung, is said to be occasionally injurious winter grains ^^ in Europe.
452
Pholiota Fries
Pileus symmetrical,
Fig. 321. P. aurivilla
(p.
449)
fleshy,
more
gills
or
less
thick,
with a
veil
which forms an amiulus;

(Bat.)
adnate, becoming rusty at maturity.
Quel,
and P. squarrosa
trees,
Miill.
occur on
deciduous
apple.
especially
on
the
P.
parasitic
spectabilis Fr. on oaks.
is
occasionally
is
P. mutabilis (Scha.) Quel,

parasite
a root
on
trees.
P. adiposa Fries.
sometimes disappear when old, convex stem 5-15 cm. by 1-2 cm., above and darker, scaly below the more or less imyellow, paler
Fig. 320.
Psilocybe pennata. After Cooke.
Cap mediiun, 5-10 cm. wide, yellow, very sticky when moist, with spreading or erect rust-brown scales which
to plane;
perfect tufted ring, solid or stuffed; gills adnate, yellowish to rustcolored, broad, croAvded; spores rust-colored, elliptic, 7-8 x 5 ju.
The name may
refer to the sticky cap. Chiefly a saprophyte, occasionally on living trees, both deciduous trees and conifers, as a wound parasite.
P. destruens Brand, occurs on poplar; P. cervinus Scha. on
various trees.
Flammula
Pileus fleshy, margin at
well
Fries (p. 449)
first
marked by the bright yellow About sixty species.

is
incurved; stipe fleshy, fibrous, or orange colored cap.
^"^^ probably a root parasite. F. penetrans Fr. and F. spumosa Fr. are regarded by Cavara
F. alnicola Fr.
^^
as root parasites of forest trees.
Volvaria Fries (p. 449)

Fleshy;
gills
free, white, later
pink; spores ellipsoid, smooth;

annulus none; volva present. Easily distinguished from pink-spored genera by the volva. Fig. 322.
all
453
other
About
thirty-six species.
Fig. 321.
Pholiota adiposa.
After Clements.
V. bombycina (Scha.) Quel. Cap large, 8-25 cm. wide, all white and silky, more rarely some-
what
scaly, hemispheric or bell-shaped to convex;
stem 8-12 cm.

elliptic,
by 1-2
6-7 X 4
cm., white, smooth, tapering upward, solid, volva large

gills
and spreading;
/i.
free,
It is often parasitic
salmon-pink, crowded, spores on various trees.
454

Pluteus Fries
(p.
450)
gills
Pileus fleshy, regular; separating easily from the stipe;

free;
volva and annulus both absent; spores elliptic. P. cervinus Scha. Fig. 323.
Cap
large,
5-16 cm. wide, usually
some shade
yellowish
to
of browTi,
from grayish or
blackish-brown, more or less fibrous or hairy on the disk, some-
times sticky, convex or plane; stem 715 cm. by 3^-1 cm., brownish, smooth
or
black-hairy,
solid;
gills
free,
pink,
broad;
spores
pink,
ju-
rarely
greenish,
globoid, 7-8 X 5-6
A common
saprophyte which
is
oc-
casionally parasitic.
Pleurotus Fries
(p.
450)
stipitate.
Pileus laterally sessile, or excentrically Fig. 324.
genus of about two hundred
fifty
species.
P. ostreatus Jacq.
Cap
Fig. 322.
cina.
large,
7-24 cm. wide, white, gray

less scaly in
Volvaria
or tan,
age,
smooth or more or
boniby After Clements.
convex or plane, shelf
or shell-
shaped, more or less lobed and torn at the margin; stem short and lateral, or none, white, solid, more or less hairy at base; gills long-decurrent, connected by veins on the
stem, white or yellowish; spores
elliptic,
8-10 x 4-5
n.
Common
berry, etc.
on deciduous
is
trees,
mainly saprophytic.
P. salignus Schrad.
P. ulmarius Bui.
often parasitic on willow, poplar, mul-
Cap large, 8-15 cm. wide, white, whitish or tan, often brownish toward the center, smooth, often cracked, usually convex, sometimes plane; stem long and stout, often nearly central, 5-12 cm. by 2-3 cm., white or tan, smooth or hairy toward the base, solid.
THE
elastic,
FUx\GI

gills
455
often curved;
annexed or sinuate, whitish, broad,
close; spores globose, 5-6 fi. Parasitic on elm and maple or usually a saprophyte.
456
but sometimes thin or even lacking; gills touching broadly or running down the stem, whitish or yellowish; spores elliptic or rounded, 7-10 fi.
This
trees,
tralia.^"^
is
common wound
parasite of conifers
and deciduous
Aus-
causing a root-rot.
It also causes a potato disease in

is
The abundant mycelium
white and extends a meter
BB9^^Î
THE
FUxXCU
457
The sporophores
or on the bark.
are borne in clusters in
autumn on the ground
The spores, sown in plum decoction, develop a mycelium which soon produces rhizomorphs. These advancing give off delicate hyphffi which may penetrate into the host. The mycelium spreads
Fig. 325.
Armillaria mellea.
.\fter
Clements.
most rapidly through the medullary rays and from them into other
tissue elements.
A. fuscipes Fetch causes a root disease of Acacia in Ceylon. A. mucida (Schrad.) Quel, is reported as a wound parasite of the
beech.
CUtocybe Fries
Pileus
(p.
450)
first
more or
less fleshy,
gills
margin at
decurrent.
incurved; stipe fleshy,
often becoming hollow;
About ninety
species.
Fig. 427.
458
C. parasitica Wil.^"^
Growing
in dense clusters; pileus
6-8 cm., convex or umbonate, usually minutely scaly, mottled

buff to yellow-brown in color;
gills
paler,
first
becoming mottled,
decurrent;
at
noticeably
stipe
1
10-16
cm.
high,
solid,
up
to
cm.
thick,
usually
curved, darker than the pileus; black rhizomorphs present.

It differs
from
Armillaria
mellea in having
no annulus,
clus-
and
ters.
in
growing in denser
causes
The fungus
by
Fig. 32U.
root-
rot very similar to that caused
Armillaria
mellea.
There
-Rhizomorphs of A. nifUea. After Freeman.
are present typical subcortical strands, mostly between the
cortex and
cambium and some-
times
subterranean black rhizomorphs adhering close to the cortex of the roots.

characteristic
Fungous branches enter the wood

rays and there
is
chiefly
later rapid vertical

cell
through the medullary growth through the vessels
and
tracheids.
The
forming loops around the nucleus.
contents are destroyed, the hyphae often The sporophores occur in

is
groups at the base of the tree after the disease
well developed.
An
extensive bibliography
is
given by
Wilcox.^"''
Collybia Fries
Pileus thin, fleshy, margin at
first
(p.
450)
incurved; stipe cartilaginous.

Fig. 329.
About two hundred seventy-five

C. velutipes Curt.
species.
Cap 2-8 cm. wide, yellow-brown or reddish brown, rarely paler except toward the margin, smooth, very sticky when moist, convex to plane or somewhat recurved, often excentric or irregular
through pressure.
459
Fig. 327.
Clitocybe parasitica.
After Wilcox
1 la.
328.
Clitocybe
parasitica,
mycelium entering medullary
ray.
After Wilcox.
460

^^ as the saprophyte which is reported by Stewart of death of the horse-chestnut. It has also been reported in Europe as a
A common
probable
cause
parasite.
Tricholoma Fries
Stout
(p.
450)
stipe
and
fleshy,
rnd
pileus of the
same subor
stance; gills sinuate nate. Fig. 330.
ad-
T. rutilans Scha. occurs on pine roots; T. saponaceum Fr. on various tree roots.
Fig.
329. Collybia
velutipes.
After
Mycena
Fries (p. 450)
Lloyd.
Small; pileus usually bell-
or less striate, shaped, rarely umbilicate, at first with the straight margin applied to the stipe; gills only slightly toothed, not decurrent or only so by a tooth; stipe slender, cartilaginous, usually
membranous and more
hollow.
Fig. 331.
A
M.
genus of some three

species.
hundred
epipterygia Scop. Five to ten cm. high;

pileus 1-2 cm. broad, viscid when moist, ovate to
conic
later
tuse,
or
campanulate,
more expanded, obthe margin striate,

minutely
age
sometimes
toothed, grayish, in often reddish; stipe 2
mm.
Fig.
gills
thick, flexuous or straight
330.Tricholoma. After Lloyd.
with soft hairs at the base;
decurrent by a small tooth,
varying in color from whitish through gray to a tinge of blue or red. Usually a saprophyte, but injurious to various kinds of trees.
Widely distributed
in the
North temperate zone.
461
462
PhaUales
gleba, which contains a series these lined by a hymenium of
(p.
395)
Mycelium rhizomorphic; sporophore with a

closely
fertile portion, the of labyrinthine spore chambers,
approximated basidia, the parenchymatous, spongy and elastic in texture, forming a receptacle which varies in size and form in the different genera. Except in Rhizogaster the young sporophore is volvate, and at the bursting of the volva immediately assumes its mature
supporting tissue
size
and form.
order of less than
fifty species of interesting,
An
foul odor
most disagreeable
fungi.
At present they
yet from their are regarded
as mainly saprophytes.
Key to
Families of Phallales
Receptacle stipitate, tubular or cylindric, capitate, with the gleba external

1.
Phallacese, p. 462.
Receptacle
branched,
closed
latticed
or
irregularly
sessile or stalked;
gleba en2.
by the receptacle
Clathraceae, p. 463.
Phallaceae
Key
to Genera of Phallaceae
Gleba borne directly on the upper portion of the stem; no special pileus Gleba smooth, even Gleba papillate or uneven Gleba covered by a rudimentary network Gleba borne on the outer surface of a special
pileus Pileus even, rugose, or reticulate Veil poorly developed or none
1.
Mutinus.
Jansia.
2.
3.
Floccomutinus.
4.
Phallus, p. 463.
Veil well developed Surface of the pileus regularly reticulate

5.
Dictyophora,
Clautriavia.
p. 463.
Surface of the pileus irregularly folded and convoluted

Pileus lamellate
6. 7.
Itajahya.

Phallus Linnaeus
(p.
463
4G2)
Stipe cylindric, even, bearing at the apex a smooth, rugose, or reticulate pileus. Less than ten species are known. The following are of economic importance.
P. impudicus L.
This
is
species of the
one of the most widely distributed genus, but not so common in

in
America as
Europe.
It
is
reported
^'
as the
cause of a root disease of the grape in Hungary. P. rubicundus Bosc.

as
has described a disease of sugar cane due to Ithyphallus coralloides. Lloyd, "however, refers the causal fungus to the present F
,
.
Cobb
^^^
\/;
g.
332.
Phallus
After
species,
smce
"
lie
considers
that
,1
11
all
the
ii
red
impudicus. Lloyd,
forms of
Phallus" constitute a single species.
Dictyophora Desvaux
(p.
462)
^^^ as species of this genus is suspected by Cobb causal fungi in a root disease of sugar cane in Hawaii.
one of
tlie
Clathraceae
(p.
462)
The
receptacle consists of a series of
arms which are
either
spreading, erect, or latticed.
Key
to Genera of Clathraceae
Receptacle of free arms, or lobes at the
sum1.
mit of the stipe Stipe columnar, arms free

Stipe enlarged upwards Limb of the receptacle with suberect
lobes
Lysurus.
2.
Anthurus.
Aseroe.
Limb
of the receptacle with radiating

3.
lobes
Receptacle of simple, erect, columns, apically imited and fertile only on their inner
side
Sessile
4. 5.
Stalked
Laternea, p. 464. Pseudocolus.
464
Receptacle clathrate or latticed

Sessile
6.
Clathrus.
Stalked
Receptacle a simple net

Stipe simple Borne on a series of columns which
are united basally into a hollow
7.
Simblum.
tube
Receptacle with the network covered with knot-like projections
8.
Colus.
9.
Klachbrennera.
Latemea Turpin
Receptacle
sessile,
(p.
463)
of
upright,
convergent
only on
columns, apically the inner surface.

L.
united
and
fertile
columnata (Bosc.) Ness,

^^^
is
recorded
by
as one of the fungi of the root disease of sugar cane in Hawaii. The species is rather
Cobb
common
in the
America, the
West
Southern United States, South Indies and Hawaii.

(p.
Lycoperdales
FiG. 333
395)
;
Later-
nea columnata. After Lloyd.
Mycelium arachnoid to rhizomorphic sporophores from the first appearing as small balls '^
.
.
of
which enlarge to maturity, gleba internal at maturity, becoming a powdery spore-mass; base the sporophore sterile peridium double or single, parenchyma;
tous, separating into flakes or breaking regularlj''; fertile hypha;, persistent in the spore mass as a capillitium which is usually at-
tached to the columella.
single family Lycoperdaceae \Yith species
which are usually
saprophytes.
Key
Outer peridium
Capillitium
to Genera of Lycoperdaceae
fragile,
more or
less
decidu-
ous, often warty, spiny or scaly

of
an even thickness, not
branched

Sporophore with a pronounced
persistent base
sterile
PLAxNT DISEASE
465
1.
Lycoperdon,
p. 465.
Sporophore
sterile
without
base
pronounced
.
Inner pcridium opening irregularly. Inner peridium opening by a basal pore, the outer peridium breakhalf
2.
Globaria.
ing equatorially and the upper with the attached inner

3.
peridium forcefully ejected Capillitium free, short-branched with

pointed ends Sporophore with a pronounced persistent sterile base
Catastoma.
4.
Bovistella.
Sporophore
without
pronounced
sterile base Inner peridium papery, opening by an
apical
mouth
thick,
5.
Bovista.
Inner
peridium
breaking
. .
irregularly, capillitium spiny.
6.
Mycenastrum.
Outer peridium splitting into star-like reflexed, persistent segments Inner peridium opening by a single mouth
Inner peridium opening by several mouths
7.
Geaster.
8.
Myriostoma.
Lycoperdon Tournefort
Sessile,
sterile
with a pronounced
Aflg^*=^
^M
î^''-
base;
peridium
thin,
^^}
^HB^
^fc
^JMSmf^:
opening regularly by an apical perforation, smooth, warty or

spiny; spore-mass and capillitium filling the interior of the
^MMBt^fiMi WWiBy ^y ^'
^"^
IHBf^îK.
ll^W^^Bt
'-'?'^'
*<.*
sporophore
phffi.
wdth
echinulate
spores and even, simple hyL.
^^
..,.,1
gemmatum
in
Bat.
^^'^
is
refir
ported by Cavara
trees
on
Italy, sending Fig. 334. Lycoperdon gemmatum. After Lloyd. rhizomorphic mycelial strands cambium and bark causing the destruction of both. through
its
BIBLIOGRAPHY OF BASIDIOMYCETES
Lutman, B.
F., Sc. 31: 747, 1910.
' '
* 6 6
'
Dangeard, P., Ser. L. Bot. 3: 240, 1892. Harper, R. A., Am. Nat. 44: 544. Harper, R. A., Trans. Wis. Acad. Sci. 12: 483, 1899. Clinton, G. P., Proc. Boston, Soc. Nat. Hist. 31: 1904.
Wolff, R., Der Brand d. Getreides, Halle, 1874. Brefeld, 0., Unt. Heft 2, also Heft, 11: 1895. Jensen, J. L., Les. Charbon des C^râles, July 4, 1889.
>
'"
Prevost, B., Memoir, Paris 29: 1807. Lobelius, Icones stirpium, 36.
" " " "

18
Persoon, Synopsis method fung: 224.

Jensen,
J. L.,
Cm
Koen, Brand,
56, 1885.
Maddox, F., D. of Ag. Tasmania, 1895, and 1897. Wakagawa, S., Bull. Imp. Cent. Ag. St. Japan 1: 1907. Brefeld, O., Klub. d. Landw. z. Berlin, 466, 1903.
Hecke, Zeit. f Land. Vers, in Oestr. 1904. Hecke, Ber. d. deut. Bot. Ges. 23: 250. Freeman, E. M. and Johnson, E. C, B. P.
.
"
18
i
I.
152: 1909.
Arthur,
2"
Kuhn,
Stuart, W., Ind. R. 12: 84, 1900. Hedwigia, 2: 5, 1858.

J.
C,
" "
2"
Brefeld, 0., Unt. Aus d. Gesamat d. Myk. Heft. 11: 1895. Hori, S., Bull. Imp. Cent. Sta. Japan 1: 73, 1905.
Clinton, G. P., Clinton, G. P.,
111.
111.
Bui. 47: 1897. Bui. 57: 1900.

545.
" Bubak, F., Zeit. Land Vers. Ost. 12: " Anderson, P. A., S. C. B. 41: 1899.
"
Farlow,
W.
G., Mass. St. Bd. Agr. R. 24: 164, 1877.
Thaxter, R., Conn. R. 1899: 129, 1890. = Ncger, F. W., Tharand, Fors. Jahr. 60: 222. 3 Bubak, Zeit. Land. Vers. St. Oest. 477, 1901. '1 Halsted, B. D., Bull. Torn Bot. Club 17: 95, 1890.
32
''
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FUNGI IMPERFECTI
species of fungus
(p.
64)
In the preceding pages it has been repeatedly evident that one may have two, even several different types of in the Erysiphales the perithecial form and the conidial; spores;
and conidia; iii the Sphaeriales the form and several conidial forms; in the Basidiomycetes ascigerous the basidial form and various conidial forms; in the Uredinales spring and summer stages and teliospores. In comparatively few instances among the many thousand species of fungi are all of the different spore forms belonging to the species known to man. In very many cases the lower or conidial forms are known without any higher spore form (ascigerous, basidial, or sexual form), being known to be genetically connected with them, though it seems very probable, reasoning by analogy, that these conidial forms really constitute part of the life cycle of some fungus which embraces also a higher form of spore. It is probable, indeed certain, that some of these conidial forms at present possess also higher, as yet unknown, forms of fructification. It is likewise probable that
in the Peronosporales oospores
in
many
cases the conidial form, though

in its
it
does not
now
possess
any higher spore form, did
not remote phylogeny possess
such forms; indeed that all of them are phylogenetically related to fungi which produced one of the higher types of spores. In some cases even in the absence of the higher spore it is possible
to refer the fungus to its proper order as for example is the case with the conidial forms of the Peronosporales, the summer or spring forms of the Uredinales, or the Oospora forms of the Erysiphales.
Regarding
ence
is
many thousands
of other conidial
forms such
refer-
impossible or hazardous, since from the conidial form the form of the higher spore can be inferred with only a small degree of
accuracy or not at
all. For example, the conidial form known as Gloeosporium in the higher form of some of its-species proves to be a Glomerella, in other cases a Pseudopeziza; some Fusariums prove to belong to the life cycle of Nectria, others to that of Neocos* mospora, etc.
475
47G
In plant pathology and in systematic mycology it becomes necessary to classify, for convenience of reference and designation, these multitudinous conidial forms of which the higher spore
unknown, which may exist now or which may have more or less remote past. From analogy it is probable that most of them pertain to the Ascomycetes, though a few may find place among other classes. This whole group of forms, which is characterized chiefly by the
form
is
as yet
existed only in the
imperfection of our knowledge of them, the name Fungi Imperf ecti.
is
classed together under
The Fungi Imperfecti

orders, families,
are in a temporary way divided into and species as are other fungi, with full genera
recognition of the fact that future research will result in many cases in the disclosure of higher spore forms and the consequent removal of species to their proper place in the general scheme of
classification.
Recognizing the tentative nature of the genera in the Fungi Imperfecti these are spoken of as "form-genera." Pathologically, the Fungi Imperfecti are of high importance, often
occurring on leaves, stems, fruit, wood, bark, etc., as active parasites, though very many are also saprophytes. Upon leaves they are particularly common causing diseased areas known as "leaf spots."
The Fungi
Imperfecti display three principal types of fructifi-
cation, pycnidia, acervuli
and hyphae.
Pycnidia are more or less spherical, hollow sporocarps on the inside of which conidia are borne on stalks (conidiophores) arising
from the base or base and
sides.
Figs. 349, 354.
The pycnidium
may be of various colors though it is most commonly black or dark; it may be superficial or imbedded, and with or without a beak
The opening for the escape of the spores (ostiole) may (rostrum) be narrow, or wanting or it may be very large, round, irregular,
.
The walls vary from extremely delicate to very thick, smooth or variously provided with hairs, spines, etc. As need arises, it is common to speak of micro-pycnidia, and macro-pycnidia. Pycnidia with very small spores are sometimes
etc.
called
spermogonia especially
if
the spores do not germinate, a cusas a pycnidium without

its wall.
tom to be deprecated. The acervulus may be regarded

It consists of a close
477
bed
of short conidiophores.
Figs. 371, 379.
Acervuli
ficial
may be small or large, subepidermal, subcortical or superand may or may not be provided with hairs (setae) Fig. 370,
An
acervulus with a well marked basal stroma
of various kinds.
is is
known
as a sporodochium. Fig. 435. If the sporodochium stalk markedly developed the structure becomes a coremium. It is
sometimes quite difficult to distinguish between a pycnidium with an extremely large ostiole, or one with a very thin wall, and the acervulus. For such purposes thin longitudinal sections are
most
useful.
Hyphae are conidiophores which grow free for some distance above their supporting substratum and in more loose form than in
the acervuli, so that the terminal parts at least stand out as separate threads. Figs. 383, 384, 396, 410.
short, or long and much branched. the hyphae are very short and closely crowded to form a sporogenous cushion the condition of an acervulus is approached
The hyphse may be simple and
When
and confusion
arises.
conidia borne in the pycnidia, acervuli or on the hyphae are of as various forms and types as is well conceivable and are made
The
the chief basis for subdivision of orders into form-genera. They may be simple or compound, of almost any color, and may be borne
in bisipetal succession in chains, or solitary, or in
groups at the
apices of the conidiophores.
The
artificial
following scheme of Saccardo presents the confessedly groups into which conidia may for convenience be divided.
Scheme of Spore Sections.

Amerosporse: spores
1
-celled,
not
stellate, spiral or filiform.
Hyalosporse: spores hyaline or clear, globose to oblong, continuous. Phaeospora?: spores dark, yellow to black, globose to oblong, continuous.
Didymospora;: spores, 2-celled. Hyalodidymse: spores hyaline,
2-celled.
Phseodidymse: spores dark, 2-celled.
Phragmosporaj: spores 3 to many-celled by cross septa.
Hyalophragmite: spores hyaline, 3 to many-celled.

Phaeophragmia;: spores dark, 3 to many-celled.
478

spores septate,
Dictyosporse:
both crosswise and lengthwise,
i.
e.,
muriform.
Hyalodictyae spores hyaUne, muriform. Phseodictyae: spores dark, muriform.
:
Scolecosporae: spores needle-shaped to filiform, continuous or septate. Helicosporse: spores spirally twisted, hyaline or dark, continuous or
septate.
Staurosporae: spores stellate or radiate, hyaline or dark, continuous or septate.
The mode of bearing spores and the color of the fungus both of which it is seen are made the basis of classification have been shown ^ ^ by Stevens and Hall and others to depend largely on environwhile the septation of the spores, also a fundamental charment, acter in present classification, depends often on the age of the Many spores are imicellular until spores or on other factors. but then become typically 2-celled; e. g., germination begins Gloeosporium. Such conditions have led to much inaccuracy in description and doubtless to undue multiplication of formspecies.
It
has been quite customary, probably to some extent excusably

describe as
form-species when no form-species presame host or its near botanical kin could be regarded as identical with it. Thus a Septoria found on Vitis would ordinarily be regarded as new unless some of the Septorias already described on soir\e of the Vitaceae seemed to be the same, even though indistinguishable from dozens of Septorias on other families of plants. This course has led to enormous multiplicaso, to
new a
viously described for the
tion of so-called species in these form-genera giving rise to such form-genera as Septoria, Cercospora, and Phyllosticta with species
numbering more than 900, 500, 800,
respectively.
The
species
condition
is
much
as
is
depicted by Cobb:^ "Is a fungus
newly found on a peach? Call it new and name it pruni. Same genus on the grape name it ampelinum. On the apple? New, call it mali. On banana? Christen it musas. What next?
Sparrow
in a pear tree. Passer pyri?"

of the
e. g.,
Many
founded,
form-genera are purely
artificial
not
at
all
well
separated from Phyllosticta only by the supposed inability of the latter to grow on structures other than
is
Phoma

leaves, a distinction
It is
479
evident that
much
which has been shown to be quite untenable.^ careful study by cultures and cross in-
is needed to reveal the true status in these Fungi. Since the conception of species is here most loose the form species given below must be regarded as purely tentative. The names are
oculations
to serve merely as handles for convenience in treating of the various parasites and in only comparatively few instances do
they signify that they are really species. In many cases forms appearing under two or more names may prove eventually to be identical while in other cases forms may need to be subdivided.
Key
to Orders of Fungi Imperfect!

1.
Conidia produced in pycnidia Conidia not in pycnidia
Spheeropsidales,
p. 479.
Hyphse innate within the matrix Hyphae somewhat superficial, often
2.
Melanconiales,
p. 537.
floccose 3. Moniliales, p. 564.

cells
4.
Conidia or other special reproductive
unknown
Mycelia
sterilia, p. 659.
The
Sphaeropsidales
slits,
Conidia in pycnidia which open by pores or resembling the perithecia of the Ascomycetes.
superficially
The
many
etc.
of
Spha?ropsidales are preeminently leaf-spotting fungi though them grow on fruit or stems causing blight, rot, cankers
The
weak
vitality,
vast majority are saprophytes or parasitic on tissues of but not a few are active parasites.
Key to
Families of Sphaeropsidales
Pycnidia globose, conic, or lenticular Pycnidia membranous, carbonous or

coriaceous, black
1.
Sphaerioidaceae, p. 480.
Pycnidia fleshy or waxy, light colored.
2.
Nectrioidaceae, p. 526.
Pycnidia more or
less dimidiate, irregular

3. 4.
or shield-shaped, black Pycnidia cup-shaped or patelliform, black.
Leptostromataceae,
Excipulaceae,
p. 528.
p. 533.
480
The
Sphserioidaceae (p. 479)
Pycnidia globose, ovate, or clavate, leathery to carbonous, black or dark brown, opening by a pore, superficial, erumpent or covered; stroma present or absent; conidia variable in form,
color,
and
division.
is
The family
below.
subdivided according to
its
spores as indicated
Key
falcate,
1
to Sections of Sphaerioidaceae
straight
Conidia globose to elongate,

to many-celled
or
Conidia
gate
1-celled, globose,
ovate or elonI.
1.
Amerosporae.
Hyalosporae, p. 480. Phaeosporse, p. 500.
Conidia hyaline Conidia colored Conidia 2-celled, ovate to elongate. Conidia hyaline Conidia colored
Conidia 3 to many-celled, by transverse
septa, elongate
...
2.
II.
3.
4.
Didymosporae. Hyalodidymae, p. 505. Phaeodidymae, p. 509.
III.
5. 6.
.
Conidia hyaline Conidia colored

Conidia muriform, ovate to elongate. Conidia hyaline
Phragmosporae. Hyalophragmiae, p. 513. Phaeophragmiae, p. 514.

Hyalodictyas.
Phaeodictyae, p. 516.
9.
IV. Dictyosporae.
7.
Conidia colored
8.
Conidia
filiform, 1 to
many-celled, hyaline
or colored
V.
coiled,
1
Scolecosporae,
p. 517.
Conidia cylindric, spirally

Conidia
stellate,
1
to
many-celled, hyaline or colored

to many-celled, hyaline
VI. 10. Helicosporae.

VII. 11. Staurosporae.
colored
Sphaerioidaceae-Hyalosporae
Spores hyaline, 1-celled, spherical,
elliptical or long.
Key to Genera
op Sphaerioidaceae-HyalosporaB
Stroma none; pycnidia separate. Pycnidia smooth

Conidia borne singly

Conidia unappendaged
Pycnidia free in the substratum; subiculum none Pycnidia not beaked, opening by a
pore, or irregularly
481
Not growing on other fungi Pycnidia opening by a regular pore Pycnidia more or less sunken in the
substratum
Pycnidia globose,
etc.
not spindle-
shaped Pycnidia borne on dark colored

spots, or on wood, globose Conidiophores simple or nearly
so
Pycnidia rather large Pycnidia at first covered, then erumpent

Spores under 15
At
On
leaves only Not on leaves
1.
2. 3.
Phyllosticta, p. 483. Phoma, p. 490.
Spores over 15 m Pycnidia from the

ficial
Macrophoma,
Aposphaeria,
p. 493.
first
super4.
p. 494.
Pycnidia very small, closely aggregated on dark spots
on the
leaf
5. 6.
Asterostomella.
Pycnidia scattered,
cylindric
superficial,
Dendrophoma,
p. 494.
circular, conidia elongate or

7.
;
Crocicreas.
Pycnidia globose
circinate
conidiophores
8.
Pyrenotrichum.
Glutinium.
Pycnidia
dric
horizontal,
free,
cylin9.
Pycnidia
opening
irregularly,
or
operculate
Spores globose
10.
Mycogala.
Spores elongate or ellipsoid
Pycnidia operculate
11.
Piptostomum.
Pycnidia opening irregularly
482

Spores pointed Spores blunt
12.
Sclerotiopsis.
13.
14. 15.
Plenodomus.
Cicinnobolus,
p.
On Erysiphaceae Pycnidia beaked Pycnidia on a subiculum

Conidia ovate or elongate
494.
Sphaeronema,
p. 494.
Subiculum
of simple hyphse
16.
.
Pycnidia free
Byssocystis.
Pycnidia sunken in the subiculum.
17. 18.
Subiculum radiate
Conidia Y-shaped
Conidia appendaged Conidia in chains
Conidial chains separate and simple ... Conidial chains connected, often forming a net
Chaetophoma, p. 495. Asteroma, p. 496.

Ypsilonia.
19.
20.
Neottiospora.
21. Sircoccus.
22.
Pecia.
Pycnidia appendaged or hairy
Appendages simple
Pycnidia with short simple tubercles;
conidia irregular in outline
23.
Muricularia.
Pycnidia
with
long
bristles;
conidia
regular
Bristles septate, usually covering the
entire
pycnidium,
conidia
cylin24.
dric fusoid, usually curved Bristles
Vennicularia, p. 496.
usually only at the apex; conidia ovate, elongate or cylin25.
dric, straight
Pyrenochaeta,
p. 497.
apex Pycnidia stromatic, superficial or sunken Pycnidia single on the stroma Pycnidia with a single chamber
Conidiophores filiform
Appendages
stellate at the
26. Staurochseta.
27.
Phomopsis,
p. 493.
Conidiophores indistinct or absent
Stroma indistinct Stroma rather well developed Pycnidia typically with more than one chamber
Pycnidia with well developed stroma, free or buried Pycnidia with separate mouths
28.
29.
Plenodomus.
Sclerophoma.
12. Sclerotiopsis.

Pycnidia single on the stroma Pycnidia several on each stroma
30.
483
Dothiopsis.
Conidia separate from each other

Pycnidia scattered irregularly
Stroma sharply defined, globose, etc. Pycnidial chambers appearing as enlargements from without ... Pycnidial chambers not as above Stroma valsoid
Conidia straight Conidia large, fusiform
31.
Anthracoderma.
32.
Fusicoccum,
p. 498.
Conidia small, ovate, clavate or

cylindric
Conidia ovate or clavate, very

small
33. Cytosporella, p. 498.
Conidia larger, ovate, or elongate
Pycnidia superficial or subsuperficial
Conidiophores simple Conidiophores branched.

Pycnidia deep seated Conidia allantoid
34.
...
Dothiorella, p. 499.
35. Dothiorellina.
36. 37.
Rabenhorstia.
Cytospora,
p. 499.
Stroma pulvinate Stroma indefinite, on black spots on

the host plant
38. Fuckelia, p. 500.
39. Placosphseria.
Pycnidia regularly arranged on the stroma around a sterile center.

.
.
40.
41.
Conidia adhering basally in fours Pycnidia on each stroma with a com-
Lamyella. Gamosporella.
mon
ostiole
Stroma globose or flask-shaped; conidia

curved
42. Torsellia.
Stroma conic-truncate, conidia elongate

cylindric, straight
43.
Ceuthospora,
p. 500.
Stroma
thin, effuse; conidia
curved to
44.
allantoid
Plagiorhabdus,
p. 500.
Phyllosticta Persoon (p. 481)
Pycnidia immersed, erumpent or with the beak piercing the
484
epidermis, lenticular to globose, thin membranous, opening by a pore; conidia small, ovate to elongate, continuous, hyaline or green; conidiophore short or almost obsolete.
On
leaves.
In part =Guignardia, Valsonectria, Mycosphaerella. The genus is a very large one of some eight hundred forms, few of which have been adequately studied. It differs from Phoma only in
that
it is
foliicolous while
Phoma
is
caulicolous, a distinction
which
Fig. 335.
P. solitaria. 1, spction through apple; 4, spores from apple blotch showing appendages; 6-7, germinating After Scott 9, mycelium from corn-meal cultures. spores; and Rorer.
not consistently maintained limitation (see p. 478).

is
and which
is
untenable for generic
The fungus produces

leaf tissue
unless
by killing or weakening the The spots are circular or subcircular, mycelium. rendered angular by obstruction by veins, and the pycnidia
leaf spots
with
its
may
leaf forbids.
usually be seen with a lens in old spots unless the color of the Similar effects follow on fruits.
P. ampelopsidis E. & M. on Ampelopsis is probably identical with P. labruscae =Guignardia bidwellii. See p. 238. P. bellunensis Mart, on elm = Mycosphaerella ulmi. See p. 249. P. brassicae (Carr.) See p. 249. siaecola.^
West on cabbage,
etc.
= Mycosphaerella brasSee
P. labruscae Thiim. on the grape =Guignardia bidwellii.

p.
238
485
P. tabifica Prill is perhaps identical with Mycospharella tabifica, though Potebnia^ questions this. See p. 247. P. maculiformis (Pers.) Sacc. on chestnut = My cosphairella m.aculiformis.
See
p. 249.
E.^'
P. soUtaria E.
&
Perithecia minute, immersed, the ostiole only erumpent; conidia broadly elliptic, 8-10 x 5-6 /x, surrounded by a mucilaginous sheath.
It
is
On
the fruit
the cause of apple fruit blotch and of cankers and leaf spots. ^ it was first reported by Clinton in 1902. The fruit
spots
characteristic fringed appearance owing to the advance of the mycelium which is limited to the outerunequal most fruit cells. In the fruit the pycnidia develop subepidermally.
show a
486
P. circumscissa Cke.
Amphigenous; spots orbicular, reddish-brown, at length deciduous; pycnidia scattered, minute; conidia elliptic, 8 x 2 Spots and shot holes are formed on drupaceous hosts.
)u.
P. prunicola Sacc.^Spots subcircular, epiphyllous, sordid-brownish or ochraceous,
margin subconcolorous; pycnidia scattered, punctiform; conidia ovoid to ellipsoid, 5 x 3 ju.

It is
and
P. P. P. P.
Australia.
foimd on Prunus, causing leaf spots in Europe, America Scurf is also produced on apple bark. armenicola Far. is associated with an apricot fruit disease. grossulariae Sacc. grows on Ribes grossularia. fragaricola D. & R. is widespread in Europe on the strawberry. vitis Sacc. and P. succedanea (Pass.) All. are found on grape
in Europe.
P. vialae R.
&
P. bizzozeriana Mass. in
G. also parasitizes the grape. Hungary produces a grape disease
superficially resembling black rot.^^
P. putrefaciens Sh. occurs on cranberry. P. oleae Pet. and P. insulata Mont, cause leaf spots on the olive. P. cannabinis Kirch forms spots on hemp leaves; P. humuli Sacc.
minute, black; spots small rounded, whitish with a purple margin; conidia ellipsoid, 5 X 2fx. Leaf spots are produced on the sweet potato.
P. nicotiana E.
P. bataticola E.
& Speg. on the hop. & M. Pycnidia scattered,
&
E.
Spots brown, reddish, zonate; pycnidia 200 n, black; conidia 3.5-5 X 1.5 /x. It causes leaf spots of tobacco;^* P. tabaci Pass, also occurs on Nicotiana.
P. medicaginis (Fcl.) Sacc. occurs on alfalfa; ^^ P. japonica Miy. and P. miuria Miy. parasitize
P. betas Oud.
rice.
^^
Spots grayish-ochre, large and irregular; pycnidia epiphyllous, minute, densely clustered, bro\\Tiish, subimmersed; conidia elliptic, 5-6 X 3 M.
mentioned by Stewart as the cause of leaf spots of beets. P. malkoffi Bub. causes cotton leaf spots in Bulgaria. P. coffeicola Del. and P. comoensis Del. are on coffee;
It is
^'^
THE
FUNCJI
487
P. hevea
P. cinnamoni Del. on cinnamon leaves; Zimm. on Para rubber. P. hortorum

'^
Speg.^**'
Spots circular, indefinite, fulvous, gray in the center, amphigenous, at last falling away; pycnidia in the center of the spot,
minute, 80-90 fx, globose-lenticular, thin, membranous, dull fusco-olivaceous; conidia elliptic to ovoid, rounded at the ends, 4-6 X 2-2.5 MIt
produces spots on leaves and fruit of egg plant in Europe
and America.
P. chenopodii Sacc.^ Spots irregular, scattered or confluent,
ochraceous, fuscous n; conida oblong-
margined; pycnidia lenticular, punctiform, 50

elliptic,
5 X 3 /i. A leaf spot is produced on spinach. P. apii Hals.^^ forms brown spots on leaves of celery; pycnidia punctiform, black; conidia elliptic to ovate oblong.
P. phaseolina Sacc.
Spots irregularly scattered, subcircular, 2-10 mm., deep rusty brown, becoming lighter in center and darker margined; pycnidia
scattered, 70-90 n; conidia ovoid oblong, 4-6 x 2-2.5
It causes spotting of
/x.
bean and cowpea.^^^
P. cucurbitacearum Sacc.
tiform, 80-100
leaves.
" 21-23
"^
Spots epiphyllous or amphigenous, sordid, whitish; pycnida punclenticular; conidia oblong, 5-6 x 214 M> curved.
iJL,
On muskmelon, cucumbers and

P. citrullina Chester
is
other cucurbs, spotting the
on melons, ^** P. maculicola Hals.^^ produces spots in Dracaena and related

also reported
plants.
P. hedericola Dur.
& M. and
^''^
P. hederacea (Arc.) All. cause
spots on Hedera leaves,-'^' P. rosae Desm. and P. argillaceae Bres. occur on roses.
P. rosarum Pass, causes a black spot on roses in
New
South
Wales.
P. althaeina Sacc.^^
Spots
lenticular,
On
with a dark brown margin; pycnidia few, ochraceous; conidia ovate-oblong, 6-7 x 3-4 ju. hollyhock in Italy, France and America.
irregular,
90
fi,
488
P. idaecola Cke. forms spots on cultivated species of Sida and

Hibiscus.
P. dianthi West, grows on Dianthus leaves. P. primulaecola Desm. occut-s on Primula leaves.^^
yllous,
Amphigenous; spots large, white, light margined; pycnidia epiphnumerous, prominent, globose, black; conidia subglobose.
^^
P. violae Desm.-^'
Amphigenous, spots white, round; pycnidia numerous, minute, brown; spores minute, subcylindric, 10 n long. Common, causing leaf spots on pansy and violet. P. hydrangeae E. & E.^^ Spots 1.5-1 cm. or more, rusty brown, margin narrow, raised, at first shaded with purple; pycnidia epiphyllous, lenticular, 100-115 n; conidia oblong, 10-12 x 2.5-3.5 ju.
On Hydrangea
causing leaf spots. P. syringae West, is common on lilac. P. halstedii E. & E.
Amphigenous, spots roundish, reddish-brown, J4~lVi cm. conix, immersed; conidia broadly f usoid-oblong, 15-20 x 5-7 m- Causing a leaf spot
centrically zonate, pycnidia few, lenticular, 100-150
of the lilac.
P. cruenta (Fr.) Kick.
Spots subcircular, reddish, becoming paler in the center; pycnidia gregarious or scattered, globose-lenticular, dark olivaceous; conidia
ovate-oblong, 14-16 x 5.5-6. 5 )n;conidiophores,cylindric, 10-12x4^. It causes leaf spots of cultivated Solomon's Seal.
P. cyclaminis Brun. occurs on cyclamen; P. digitalis Bell on digitalis. P. chrysanthemi E.
&
D.^^
n.
Spots purplish-brown, pycnidia 80-100 fi; conidia 4-5 x 2.5-3 It causes leaf spots on cultivated chrysanthemums.
P. leucanthemi Speg.
is
occasionally found in spots of chrysan-
themum
lily
leaves.
is
P. richardiae Hals.^^
common
as a leaf spotter of the calla
but has not been satisfactorily described.
P. opuntiae Sacc. & Speg. occurs on various of the Cactacese; P. liliicola Sacc. on lily; P. vincae minoris B.
&
K. on Vinca minor;

P. pteridis Hals, causes blighting of cultivated Pteris; P. narcissi Aderh. of narcissus.^"
'^
489
P. cavarae Trinch. produces white spots on leaves of Anthurium.^' P. dracaenae causes spots on Dracaena leaves.
P. funckia Hals.
Pycnidia 75-150
fi,
straw colored.
The cause of P. dammarae
leaf spots of cultivated Funkias.^'^

is
found on
Dammara
in Italy;
P. nobilis Thiim. on Laurus.

P. ulmicola Sacc.
Spots indefinite, ochraceous, margin concolorous; pycnidia gregarious, punctiform, 70-80 n, lenticular; conidia oblong ellipsoid,
6-3
fjL.
It infests
elm
leaves.^'*
P. acericola C.
&
E.32
Spots irregular, fuscous, brown margined; pycnidia densely scattered on the central part of the spot, subepidermal, flaskshaped, dark brown, 120 m; conidia ovate, 8-9 x 5-6 /x.
It causes serious leaf spotting of
maples throughout the United
States.
P. aceris Sacc. forms small spots on maple leaves. P. paviae Desm,
Spots indeterminate, reddish, lighter margined; pycnidia epiphyllous, black; conidia cylindric-elliptic, 11-12 n long. It is said to be common on Êsculus.^'*
P. sphaeropsidea E,
&
E.^^
Epiphyllous; spots reddish-bro^îi, margin lighter, scattered or confluent, 1-2 cm.; pycnidia scattered, immersed, punctiform, erumpent above, subepidermal; conidia globose to broadly ellipsoid, hyaline, 12-15.5
It
x 8-10
n.
causes serious leaf spotting of chestnuts throughout the

tiliae
United States.
P.
Sacc.
&
Speg.
is
P.
minima
P, catalpae
(B. & C.) E. & M.
E.
found on Tilia. is on Negundo.
Spots, rounded, brown, 3-6 mm., often confluent; pycnidia subcuticular, small, black, scattered, 112 x 84; conidia ovate, 5-7 x 2.5-4.5 fi. It causes leaf spots on Catalpa.^*
K. on ash; P.
P. magnoliae Sacc. causes leaf spots on Magnolia; P. viridis E. ilicina Sacc. on the cork oak.
&
490

unidentified species has been reported on watermelon ^^ as Halstead mentions also an undeteroats.^^
An
cause of considerable injury.
mined species on
6-7
fjL,
Pycnidia 150-250
/i;
spores 12-18 x
pyriform.
Phoma
The genus
It
it is
(Fries)
Desmaziere
(p.
481)
as at present recorded contains over 1200 forms.
is
indistinguishable from Phyllosticta (see p. 484) except that caulivorous. Several species are regarded as conidial forms
of Diaporthe, Mycosphserella, etc.
P. reniformis on grape =Guignardia bidwellii. See p. 238. P. albicans Rob. & Desm. on chicory = Pleospora albicans.
See
p. 260.
P. betae Fr. on beet = Mycosphserella tabifica. See p. 247. P. bohemica Bub. & Kab. on fir tree needles =Rehmielliopsis.
See
P.
p. 276.
ambigua
(Nitz.)
Sacc. on pear
= Diaporthe ambigua.
See
p. 279.
P.
sarmentella
Sacc.
on
hop = Diaporthe sarmentella.
See
p. 279. P. persicae Sacc.
Pycnidia scattered to gregarious, globose lenticular, /e- /s mm. conidia oblong ovoid, 8-3 x 2 n, conidiophores cylindro-conical,
equal in length.
It
produces constriction and death of peach twigs.^^
P. mail S.
&
S.^'-
38
Pycnidia gregarious, subcuticular, depressed, ostiole erumpent; conidia oblong-fusoid, 2-3 x 5-8 (x. It attacks the wood of young apple trees and also causes a decay of the fruit.
P. cydoniae Sacc.^^
Pycnidia subgregarious, depressed, ostiole obtuse or erumpent, conidia elliptic oblong, 8-9 ix long; conidiophores short. A form causing rot of quince fruit was provisionally referred
to this species
by Halsted.
P. limonis
Thum. &
Boll.
P.
citri
Sacc.
and P. aurantiorum
(Rab.) Sacc. occur on citrous fruits;
THE FUXGl WHICH CAUSE

P.
rLANi' DISEASE
491
pomarum
Thiiin. on poinaceous fruits in Europe.
P. myxiae Far. is associated with an apricot fruit spot. P. omnivora McA. is described as the cause of Australian wither
tip of the
common P. mororum
tralia.'*^
while to P. citricarpa McA orange Australian citrous fruit disease.

*
is
attributed another
Sacc.
is
on Morus.
causes a disease of grape berries in Ausis
P. tuberculata
McA.
P. lophiostomoides Sacc.
cereals.
common and
perhaps parasitic on
wheat
P. hennebergii Kiihn produces brown spots on the glumes of ''antl leads to some injury to the grain. P. solanicola P. & D.''^ causes a disease of potato ^^^ stems in
France.
P. solani Hals.^^
Pycnidia innate, depressed, oblong; conidia oblong.
On egg plant causing damping-off of seedlings. P. subcircinata E. & E.

Pycnidia black, 70-90 n; conidia 5-6 x 2-2.5
Spots are produced on bean pods. P. sanguinolenta Rost.^^
/x.
Pycnidia scattered, subglobose; conidia ellipsoid, 4-6 x 1.5-3 a slime which gives the spore-mass a violet-red fj.; surrounded by
color.
As the cause
Jersey. P. oleraceae
of a rot of carrot roots
it
has been reported in
New
Sacc.''""''^
the tissues, J<4-K
globose depressed, papillate, sunken in mm.; conidia oblong, subcylindric, medially constricted, apically obtuse, 5-6 x 2 ;u. Manns notes this fungus causing a serious cabbage disease in
Pycnidia
scattered,
Ohio. The pycnidia are sparse on oval sunken diseased areas on the stems, and bacterial invasion follows soon in leaves, cambium and xylem. The cambium is rapidly destroyed and the
plant collapses.
Bos
^^
and Quanjer
'^^
have demonstrated the

in
pathogenicity of the fungus. P. napobrassicae Rost. causes
mark;
^^^^
also recently reported
rot of mangolds from Canada. '^^
Den-
492
P.
P. apiicola Speg. is recorded on celery.^brassicae Thiim. on cabbage is probably identical

is
with
P. oleracese.
P. roumii Fron.
Africa. ^^
said to cause a serious cotton disease in
P. batatae E.
&
H.^''
Pycnidia blackish, gregarious, immersed; conidia terete, ovoid; The cause of dry rot of sweet potaconidiophores slender.
toes.
P. chrysanthemi Vogl.
is
found on leaves of chrysanthemum
causing them to wilt. P. malvacearum West
is
noted on European hollyhocks;

;
P. devastatrix Berk, on cultivated lobelias
Fig. 337.
P. oleracea.
A, showing pycnidia with spores streaming out; B, section; C, spores. After Manns.
P. dahliae Berk, on stems and flowers of Dahlia.

P. cyclamenae Hals, is given as the cause of Cyclamen leaf spots but without ample description.
P. oleandrina Del.
is
on the
rose, laurel, etc.

fir
P. pithya Sacc. seems to be parasitic on the

striction
causing con-
and death
of twigs.
P. strobi (B. & Br.) Sacc. is prevalent on white pine in Europe. P. strobilinum P. & C. is closely related to the above. P. sordida Dur. & M. occurs on Carpinus. P. ribesia Sacc.
fusoid, 10 X
3^
ju,
hyaline.
Pycnidia collected, erumpent, spores oblongIn branches of Grossularise.
Several undetermined species have been reported,
among them

one on snapdragon,^"* another on Clematis
^''^
493
roots,*^
and one on ap-
ple
" Phomopsis Saccardo As

in
(p.
482)
P. aloeapercrassae Trinch. the aloe in Italy.
Phoma, but with hooked conidiophores. A small genus. is reported on scapes and flowers of
P. stewartii Pk.
Perithecia gregarious,
commonly occupying grayish
or bro\Mi
spots, thin, subcutaneous, at length

"2.
erumpent, depressed, minute,

first; filiform,
/i,
curved, second; oblong or subfusiform, hyaline, commonly binucleate, 8-12 x 2-3 n; sporophores slender, equal to or shorter than the spores.
Iz- I mm. broad, black; spores of two kinds, fiexuous or uncinate, hyaline, 16-25 x 1-1.5
The fungus with its filiform spores only was noted as a parasite on Cosmos by Halsted who referred to it as a species of Phlyctaena.^' ^^ It has been noted in New York by Stewart, and is destructive both in the greenhouse and in the open.
Macrophoma
Berlese
&
Voglino
(p.
481)
As in Phoma, but the ostiole of the pycnidium not papillate, and the pore smaller; conidia over 15 î long; conidiophores simple, short or filiform.
About one hundred seventy-five
species.
M.
hennebergii (Kilhn) Berl.
&
Vogl. causes a serious disease
on wheat in Sweden.^^ The fungus which appears in the Uterature as

Pk.
is
M.
curvispora
in reality
p. 542,
is
and
Gloeosporium malicorticis, see that referred to as M. malorum

corticolum.
Myxosporium
M.
vestita Prill.
&
Del. attacks
See p. 546. cacao in
^0^^
,, Fig. 338. M. curvispora. spores. Af^^ ^ ^^'
Ecuador. M. dalmatica (Thiim.) ^ '

olive;
M. taxi B. & M. abietis M. & H. is associated with a fir disease; M. manihotis Hem. is on cassava; M. nag on Hydrangeas; M. helicinia Magnag on ivy.
Taxus;
V. ^ parasitizes the V. attacks the leaves of
B.&
ligustica
Mag-
494

reniformis (V.
M.
Italy
&
R.) Car.
is
reported on grapes in Algiers,
and Russia.
Aposphseria Berkley
(p.
481)
Pycnidia globose, carbonous, with a papillate ostiole, erumpent or superficial; conidia elongate to globose; conidiophores
very short or absent.
One hundred
and
species are recognized.

species
An undetermined
was found by Stevens
-^
in
New York
New
Jersey in 1892, causing diseased spots on strawberry
leaves.
Dendrophoma Saccardo
(p.
481)
Pycnidia superficial or subepidermal and erumpent, carbonous; ostiole papillate; conidia elongate; conidiophores branched.
genus of some
fifty
species,
chiefly
sapro-
phytes.
D. marconii Cav. occurs on hemp stems; D. con- Fig. 339. DenCav. on leaves of Convallaria majalis; n^dôp^'orês D. valsispora Penz on living lemon leaves. ^"^^j. ^"j'^'gl cêr. Cicinnobolus Ehrenberg is frequently met as a parasite on the mycelium of the Erysiphales. Macrodendrophoma salicicola on Salix = Physalospora gregaria. See p. 252.
vallariae
Sphaeronema
Fries (p. 482)
Pycnidia superficial or not, p3Tiform, cylindric or globose, rostrum long; conidia ovate or elongate. Some seventy-five species, chiefly saprophytes, have been described.
S.
phacidioides Desm. on clover = Pseudopeziza
trifolii.
See
p. 148.
S.
fimbriatum (E.
&
H.) Sacc.^^'
n,
^^
Pycnidia globose,
100-200
surrounded by septate, hyaline
495
hyphse, rostrum, 20-30 ^ long, apically fimbriate; conidia globoseelliptic,
5-9
/x.
The fungus grows
in the
black spots in the skin.
The
sweet potato producing dark, almost tissue below becomes olive-green.
The dark mycelium
is
found penetrating
through and between cells of the diseased area where numerous olivaceous
conidia
are
also
present.
The
elon-
gated beaks of the pycnidia rise like a small forest from the surface of the
potato.
mycelium is and with nudark, abundantly septate merous oil globules. Long multiseptate
artificial
In
culture the
.,
conidiophores ^\^th light arise from the medium.

hyaline
ently
colored
tips
fiq.
340.s. fimbriatum;
1,
From
340,
these
conidia
are
produced,
Fig.
apparOliva-
pycnidium sending forth spores; 2, hyaline conidia; o, oHve conidia. After Haisted
endogenously.
ceus, globose to elliptical. Fig. 340, conidia are
formed within the
medium on branches of the mycelium in much the same manner. The pycnidia develop in about nine days after inoculation and the conidia are extruded from the fimbriate mouth of the long rosr
trum.
Inoculations
proved
the
pathogenicity
of
the
organism,
typical black rot appearing in about three wrecks after infection.
S. S.
S.
adiposum Butler causes a black rot of sugar cane. pomarum Sh. is on cranberr3^ spurium (Fr.) Sacc. on Prunus is often reported as Dematium
is
prunastri. S. oryzae Miy.
on
rice.^^
Chaetophoma Cooke
(p.
482)
Pycnidia superficial, very small, on a subiculum of interwoven hyphae; conidia ovate or elliptic, very small.
Some forty species, chiefly American. C. glumarum Miy. parasitizes rice in Japan. '^
496
THE FUNGT WHICH CAUSE PLANT DISEASE
Asteroma De Candolle
(p.
482)
hyphae; conidia ovate or short cylindric.

p. 274.
Pycnidia very small, globose, erumpent, often on a mass of In part =Gnomonia. See
species chiefly parasitic.
About forty
p. 275.
A. padi (D. C.)
Grev. on
Prunus=Gnomonia
padicola.
See
A. geographicum (D. C.) Desm. occurs on various species of
Pomaceae;
A. punctiforme Berk, on the rose; A. stuhlmanni Hen. on bananas and pineapples in Africa. A. codiaei All. is said to be a serious parasite of Codiaeum.^^
Vermicularia Fries
(p.
482)
Pycnidia superficial, or erumpent, globose depressed, to globose leathery or carbonous, black, ostiolate or not, beset with rather long, stiff, septate, dark colored bristles; conidia
clavate,
cylindric-fusoid, often curved.
Some
V.
one
hundred
thirty
dematium
(Pers.) Fr.
/^, /
Pycnidia erumpent, superficial, 80-120 n, conic, then depressed,

often
confluent, black, spines pale at the ends, 150-200 x 5 /i; conidia cylindric-elongate, 20 x 4-6 n, apically rounded, curved.
Fig. 341.
occasionally causes asR, hyphse. After Reed. paragus disease .^^ In Europe it is reported as the cause of much loss to the ginseng crop. On The fungus was this plant it produces a stem anthracnose.
V. dematium. C, a nearly mature "perithecium; " S, spores;
Commonly
fungus
a saprophyte, this
isolated
and
its
cultural characters studied
by Reed.^^
V. trichella Fr.
Pycnidia ovate, small, black, spines long, at the apex of the pycnidium; conidia fusoid, curved, pointed, 16-25 x 4-5 /x.

On
living parts of
plants.^^
497
many
fruit hosts, as well as ivy
and other
woody
V. melicae Fcl. grows on Melica; V. microchaeta Pass, on camellia.
V. circinans Berk.^^'
^^
Spots orbicular; pycnidia arranged concentrically, small, setae long; conidia oblong, curved, obtuse. On onions the fungus appears as small black dots on the scales.
These later become encircled by rings of black pycnidia. Stoneman found no true pycnidium; this would indicate relationship of the organism with Volutella rather than with Vermicularia. V. varians Due. is described by Ducomet as the cause of a scablike disease of tomato and potato.^ V. subeffigiirata Schw. Pycnidia large, scattered, dark, subelevated; spines unequal. V. telephii Karst.î
On
carnation leaves.
Pycnidia scattered, erumpent, superficial, spherical, dark, 100150 n; conidia fusoid bacilliform, acutely curved, 22-32 X 4 M.
On
leaves
and stems
of cultivated
is
Sedums.
-'
reported by Halsted as causing unsightly spots on Dracaena. V. denudata Schw. A Vermicularia referred
to as probably this species is reported as damaging to Kentucky blue grass in Dakota.^*^ V. polygoni-virginica Schw. is reported by Reed & Cooley on rhubarb. ^^^
V. concentrica Lev.
K
'
'^
_
berbciidis,
cha?ta
e'oSa.^ Af";r a?lescher.
An undetermined
species
is
reported as injurious to the potato.^

(p.
Pyrenochaeta de Notaris
Pycnidia
black,
482)
globose-clavate,
ostiolate;
erumpent,
leathery
bristly,
conidia ovate,
or carbonous, elongate or cylindric;
conidiophores branched.
A genus of some thirty species. P. phloxidis Mas. is common just above ground on living stems of Phlox causing cankers.
P. ferox Sacc. P. oryzae
found on potato stems. ^^ Miyake occurs on rice in Japan.

is
498

Fusicoccum Corda
(p.
483)
Stroma subepidermal, several-chambered, erumpent, leathery, black; conidia fusoid, straight and usually large. Some forty species, several of which are regarded as conidial forms of Diaporthe and Gnomonia. F. veronense Massal on Sycamore and Oak=Gnomonia veneta.
See
p. 274.
F. viticolum Red.
on grape =Cryptosporella
viticola.
See p. 280.
Fig. 343.
F. viticolum, compound
raphyses.
pycnidium, germinating spores, paAfter Reddick.
F. amygdali Del. causes a spot disease of
almond twigs
in
Europe.
F. bulgarium Bub.
Austria.^^
is
described as the cause of a grape disease in
F.
perniciosum
See
Briosi
p. 281.
&
Farm,
on
chestnut=Melanconis
modonia Tul.
Cytosporella Saccardo
(p.
483)
Stroma tuberculate or cushion-form, immersed, then erumpent,

leathery, black, lighter within; conidia clavate or ovate, usually quite small. Some twenty-five species.

C. cerei, Poll, is oii Ceieus; C. citri JVIaynag. on oranges; C. damnosa Pet. on pine; all in Italy. C. persicae Scliw. is reported on young peach branches.
499
Cytospora Ehrenberg
(p.
483)
Stroma
chambers;
superficial
or
erumpent,
tubercular,
with
irregular
conidia
elongate
allantoid.
Ascigerous forms belonging to Valsa are
known.
Some two hundred

saprophytes.
C.
species,
chiefly
palmarum Cke.
is
on palms.
C. ceratophora Sacc. is the suspected cause of a blight of Japanese chestnuts.^^
SecFig. 344. Cytospora. tion through a stroma. After Chester.
C. acerina Aderh. causes disease of Acer in Europe.^" C. sacchari Butler is found on sugar cane in Bengal.
Dothiorellina Bubak with the one species D. tankoffiii Bub. has ^^ recently been described as the cause of disease of the mulberry.
Dothiorella Saccardo
(p.
483)
Pycnidia erumpent, on a stroma, leathery, ostiole papillate or

not; conidia ovate or elongate.
Some seventy
D.
ribis (Fcl.)
Sacc, on a wide range of hosts =Diaporthe stru-
mella.
See
p. 279.
Fig.
34.5.
D.
mori.
.V,
section of .stroma,
(J,
conidio-
phores and conidia.
After AUescher.
D. mori Berl. and D. populi Sacc. are perhaps parasitic on Morus and Populus respectively. An unidentified species is reported by Duggar on currant as the cause of cane blight. Inoculations using the conidia have
produced the disease.
See also p. 283.
500
Fuckelia Bonordin
(p.
483)
Stroma erumpent, globose-pulvinate,

elliptic.
substipitate, dark without,
lighter within, with several angular pycnidial locules; conidia
single species F. ribis
form of Cenangium
vitesia.
Bon. on currants in Europe See p. 151.

483)
is
a conidial
Ceuthospora Grevielle
(p.
Stroma
coalescing, erumpent, cushion-shaped, leathery,

all
many-
chambers opening by a common pore; conidia elongate cylindric, mostly straight.

chambered,
Some
twenty-five species, chiefly saprophytes.
Fig.
346. C.
c
cattleyae, a
py
nid
u m.
After
Delacroix.
Fig. 347. Plagiorhabdus oxycocci on cranberry. After Shear.
C. cofifeicola Del. C. cattleyae Sacc.

^^^
is
of questionable parasitism
on
coffee;
&
Syd. on orchids.
Plagiorhabdus oxycocci Shear has

berry.
been
reported on cran-
Sphaerioidaceae
Conidia
Phaeosporae
(p.
480)
1-celled, dark, globose,
ovoid or oblong.
Key
to Genera of Sphaeropsioidacese
Phaeosporae
Pycnidia separate Pycnidia without mycelium or subicle Pycnidia smooth, not hairy
Conidia
in chains, globose
1.
Sirothecium.

Conidia not
Pycnidia
in
501
chains
spheroid Pycnidia beaked Pycnidia not beaketl

sessile,
2.
Naemosphaera.
spindle-form, with both ends light colored. Conidia globose to elliptic

.
Conidia
3.
Hypocenia.
Pycnidia
larly
opening
irregu4.
Harknessia.
Pycnidia opening by a regular ostiole
Conidia large, ovate to

elliptic
").
Sphaeropsis,
p. 501.
Conidia very small, globose to ellipsoid. ...

Pycnidia stipitate, clavate Pycnidia Pycnidia hairy or setose with distinct mycelium
subicle
in a
6. 7.
Coniothyrium, Levieuxia.
Chaetomella.
p.
503.
8.
or
Pycnidia astomous,
Pycnidia perforate
dark subicle.
9.
Capnodiastrum.
Cicinnobella.
10.
Pycnidia cespitose or in a stroma Pycnidia in dense erumpent clusters
11.
Haplosporella.
Pycnidia not as
above,
in
definite
stroma
Stroma applanate or effuse, foliicolous Stroma dot-like, discoid or hemi.
12.
Discomycopsis.
spheric
Stroma dot-like, immersed Stroma discoid to hemispheric Stroma discoid; spores large Stroma pulvinate; spores minute,
catenulate
13.
Melanconiopsis.
Nothopatella.
Cytoplea.
14.
15.
;
Stroma
hemispheric
pycnidia
16.
circinate
Weinmannodora.
Sphaeropsis Leveill^
Pycnidia immersed, erumpent, globoid, black, leathery,

conidiophores rod-like.
mem-
branous, with the ostiole papillate; conidia ovate or elongate,
502

species several of
About two hundred

pathogens.
S.
them important plant
malorum Pk. " ^'^ ""^^ Mycelium sooty-brown; pycnidia erumpent,
usually surrounded
by broken epidermis, apically somewhat depressed; conidia oblong elliptic, brown, usually about twice as long as broad, 22-32 x 10-14 fx, varying in size with host and part attacked. On apple, pear, quince, hawthorn; on twigs causing canker or blight; on fruit causing rot and on leaves causing spots.
I'iG.
348.
S.
malorum,
the
b,
dark colored mycelial

of
threads
among
cells
the
fruit;
d,
pycnidium, which has pushed through the epidermis, c, and is giving off dark colored spores, e; B, mature spores germinating in water. After Longyear.
one of the common causes of pomaceous fruit rots and United States. Its occurrence in leaf spots was noted in 1898,^^ and in 1902 Clinton ^ recognized it as their cause. Cultures were obtained from diseased leaf spots by Scott & Rorer ^^
This
is
of leaf spot in the
and by inoculations the

definitely proved.
ability of the
fungus to cause spots was
This fungus was reported by Paddock ^^ as the probable cause of apple twig blight and canker and cross inoculation between
twigs and fruit proved the identity of the fungus on these two parts. The mycelium is very dark or olivaceous and abounds in the
rotten pulp of affected fruit, also in diseased bark,
and
is
even

present in
tissue.
503
wood though extending but
sparingly into
woody
A pycnidial fungus agreeing witli S. malorum morphologically has been shown by Shear *' to be a eonidial form of the ascigerous
fungus Melanops (=Botryosphseria), see p. 284. S. pseudodiplodia (Man.) G. & M. ^^' ^^ causes an apple disease
in
Europe.
S. S.
mori
ulmi
Berl. parasitizes
S.
Morus;
R. the elm; S. magnoliae Magnag. the Magnolia in Italy;

S.
&
japonicum Miy.
rice in
Japan.
^^
S. vincaB S.
& W.
Pycnidia gregarious or scattered, globose, immersed, black, small, 2G0-300 /x; ostiole papillate, erumpent; conidia ovate, ovate-oblong or subpyriform, 17-28 x 10-14 ju. On Vinca.^^
Many
other forms are recorded on various hosts but their

is
parasitism
questionable.
Coniothyrium Corda
(p.
501)
Pycnidia subcortical, erumpent or not, globose or depressed, ostiole papillate, black, leathery to carbonous; spores small, ellipsoid, conidiophore reduced or absent.
More than one hundred

C. pyriana (Sacc.) Shel. regarded as their cause.^
is
fifty species.
common on
apple leaf spots but
is
not
C. concentricum (Desm.) Sacc. occurs on Yucca, Dracaena, etc. C. tumefaciens Gus.*^ is described as the cause of a rose canker.
C. melastorum (Berk.) Sacc.^^ C. fuckelii Sacc.^^
is
on sugar cane.
Pycnidia superficial, scattered, dark, 180-200 n, globosedepressed; conidia numerous, globose to short-elliptic, 2.4-5 x 2-3.5 AX.
The European form is reported on dead and dying branches and a form closely allied to it, probably identical, has been studied in New York as tlie cause of a raspberry cane-blight. This fungus and no other was present and typical disease followed inoculation. The organism was recovered in pure culture. Both new and old canes died within two months after inoculation.
504
This

is
a conidial form of Leptosphseria coniothyrium.
See
p. 257.^^^
studied
The same fungus was reported by Stevens & Hall^^ and was by O'Gara^^ and determined by inoculation and cross
and apple
rot.
inoculating, using pure cultures, to be the cause of rose
canker and apple fruit
C. diplodiella (Speg.) Sacc.^"
Pycnidia minute, subcuticular, erumpent, browTi, 100-150 n; conidia ovoid to elliptic, 7-11 x 5.5 n; conidiophores simple or branched, hyaline, filiform. This is the cause of a v/hite rot of grapes and has been reported
Fici.
349.
C. diplodiella, section through

After Scribner.
pycnidium.
by
See
Viali
& Ravez
as belonging to the ascigerous genus Carrinia.^

of
p. 263.
Though probably
in Italy in 1878.
first
American
it
In 1887
origin it was first recognized caused alarm in France and it was

is
in the affected pulp
noted in America in the same year. The mycelium and sometimes upon the seeds.
abundant
Peduncles
are often killed.
The pycnidia
is
are subcuticular,
first
pink, then
white, later brown. C. scabrum McA.

in Australia.
the cause of black scurf of citrous fruit

coffee in Java.
C. coffeae C.
Zimm. is on vagabundum Sacc.
causes premature
fall of
leaves of goose-
berries.
C. japonicum Miy., C. brevisporum Miy. ^^ Miy. are found on rice in Japan.
and C. anomale

C. wernsdorfSae C. hellebori C.
505
Kock
is
occurs on roses.
& M.
found on hellebore.
Sphaerioidaceae
Hyalodidymae
(p.
480)
Conidia hyaline, 1-septate, ovoid, ellipsoid or
oblonfj;.
Key
to Genera of Sphaerioidaceae
Hyalodidymae
Pycnidia separate Pycnidia not beaked

Pycnidia in discolored areas, maculicole Pycnidia immersed, then erumpent,
perforate
Conidia muticate Conidia with set* at the apex.

Pycnidia superficial, astomous Pycnidia not maculicole Pycnidia hairy
...
1.
Ascochyta,
Robillarda.
p. 506.
2.
3.
Pucciniospora.
4.
Didymochaeta.
Pycnidia smooth Conidia with an appendage at each
end
Conidia with
1
or
more
bristles
5.
Darluca.
Tiarospora.
Conidia with cap-like appendages

6.
Conidia muticate
Conidiophores 1-spored Pycnidia without subicle
Pycnidia on a cobwebby subicle, phyllogenous Conidiophores several to manyS.
7.
Diplodina,
p. 509.
Actinonema,
Cystotricha.
p.
508
spored Pycnidia beaked Pycnidia in a stroma
9.
10.
Rhynchophoma.
Stroma effuse Stroma consisting
of
two
distinct
11. 12.
layers Stroma of a single layer
Thoracella.
Placosphaerella.
Stroma verruciform Stroma superficial Stroma erumpent
13.
Patzschkeella.
14.
Cytodiplospora.
506

Ascochyta Libert
(p.
505)
Usually producing definite spots; pycnidia globose-lenticular,

ostiolate; conidia ovate.
About two hundred
fifty species.
A. pisi Lib.2i' î- 92=Mycosphserella pinodes.^^^ See p. 250. Spots variable in size, roundish, yellowish with brown margin;
pycnidia centrally located, black, of angular cells, 5-7 ix; ostiole rounded, surface reddish brown; conidia
slightly
constricted at the septum, oblong, 12-16 X 4-6 n; exuded spore-mass brown.
peas, beans, vetch, Cercis, etc. The pycnidia are visible on the dead areas of the
On
stems, leaves, pods or seeds. hibernates in affected seeds,
The mycelium
reduces their
germinating power and

Fig. 350. a. citri, After Sacspores. cardo.
the fungus over to the succeeding crop. ^' boltshauseri Sacc. on bean in Switcarries
zerland
.
^^
is
closely
"^
related
to
the
last
species.
A. armoraciae Fcl.
A.
ellisii
is on horse radish, causing leaf spots; Thiim. on grape; A. brassicaB Thiim. on cabbage, forming large dull patches;
often quite injurious. A. rhei E. & E.
Spores finally constricted and 1-septate, 7-12 x 3.5-4 On rhubarb.

A. viciae Lib.
/x,
hyaline.
red;
Epiphyllous; spots roundish, reddish, margin elevated, orange pycnidia minute, clustered, black, 90-100 /z; conidia
/x;
oblong-ovate, obtuse, slightly constricted, 12-15 x 4-5 mass white. On Vicia.
exuded
A. nicotianae Pass.^
Spots between the veins, irregularly scattered, brown; conidia oblong ovate, constricted at the septum. On tobacco.
A. parasitica Faut.
Spots whitish; epiphyllous; pycnidia small, black. liptic, 3-4 X 6-10 iu.
Conidia
el-

This
is
507
^^
found associated with rust sori on malvaceous hosts. '^' A. polemonii Br. & Cav. occurs on Polcmonium.
A. piniperda Lin. is parasitic on fir leaves. A. aquilegiae Roum. spots columbine leaves. A. beticola P. & D. is on beet leaves; A. orobi Sacc. on sainfoin; and A. oryzae Catt. on rice in Italy. A. lactucae Rost. is on lettuce;
A. aesculi Bub.
&
Kab. on îculus
in
Europe;
A. pallida Bub. & Kab. on Acer in Europe; A. pruni Bub. & Kab. on the cherry in Europe; A. populicola Bub. & Kab. on the Silver Poplar in Europe; A. dianthi Berk, on Dianthus and other pinks; A. violae Sacc. causing spots on violet leaves; A. digitalis Fcl. on digitalis; A. iridis Oud. on
Iris.
^^
A. juglandis Bolt, causes spots on leaves of Juglans; A. aspidistrae Mas. on Aspidistra.

A. fragariae Sacc. Perithecia partly immersed, black, 100-125 to cylindric, constricted, 14-27 x 4-5.5 fx.
/x;
conidia fusiform
This was reported by Dudley ^^ as occurring in injurious form near Rochester, N. Y., causing spots, at first red, later brown,
on strawberry leaves.
A. primulae Wail.^^
globose,
Epiphyllous; pycnidia on discolored spots, scattered, depressed 100-110 fi, pale brown, papillate ostiolate; conidia cy-
lindric, obtuse,
5-6 x 2-2.5
/x.
On
Primula.
A. chrysanthemi Stev.^^
Pycnidia few, immersed, early erumpent, single or scattered,

hemispheric, amber-colored, 100-200 n; ostiole central, small, often raised by a neck, dark-bordered; conidia oblong, straight or irregular, 3-6.2 x 10-20 n, apically obtuse, septum often obscure,
sometimes more than one; not constricted till germination. chrysanthemums. A. medicaginis Bres.
It causes blighting of ray flowers of
apiculate, pale,
Spots small, angular, pale, clustered; pycnidia sublenticular, becoming black, 200 x 100 n, context parenchy-
matous; conidia oblong, obtuse, scarcely constricted, 10-12 x 4-
508
4.5 n.
According to Stewart, French & Wilson, spots are caused on alfalfa.^^ The American form is distinct from the European and has been described under the name A. imperfecta Pk.'*'*^
A. lycopersici Brum. Spots red or brown, large, rounded or irregular; pycnidia sparse, minute, black; conidia oblong, constructed, 8-10 x 2.5 m-
Spots are produced on leaves and fruits of egg plant. A. caulicola Lau. causes injury to Melilotus alba.^
A. cookei
Mas.
is
A. corticola
McA.
reported on Sweet William. is the cause of lemon bark-blotch in Australia,
killing the trees.^^
A. graminicola Sacc. occurs on grasses and grains;
Fig. 351.
Actinoneiua
mycelium;
iob;i;.
o,
4, spores.
branching strand of superficial After Southworth.
A. manihotes Hen. on cassava in Africa.
A. tremulae Sacc. occurs on aspen; A. melutispora B. & Br. on ash.
Actinonema
Fries (p. 505)
Pycnidia very small, not ostiolate, with a radiating mycelial
growth on the surface

conidiophores.
of the
host; conidia elongate,

leaf parasites.
on short
genus of about fifteen species, chiefly

'*'
A. rosae (Lib.) Fr.^^'
Spots rounded or irregular, black or purple, epiphyllous, often
509
confluent, murginalh' fimbriate, the radiating fibers arachnoid, white, distinctly branched; pycnidia tuberculariform, scattered or confluent, black; conidia oblong, constricted, 18-20 x 5. n;
conidiophores short.
This fungus was

leaves.
first
described in 1826.
is is
It is
common on
rose
The mycelium
in part subcuticular,
visible
in part deeper.
The
this
subcuticular part
through the
cuticle,
consisting
of radiate strands each
composed
of several parallel
hypha. From
mycelium branches penetrate deep into the leaf. The dark is due to discoloration of the contents of the diseased cells; the mycelium itself having little or no color.
color of the leaf spots
A.
tiliae All.
causes defoliation of Tilia.
A. fagicola All. occurs on beech leaves; A. fraxani All. on ash.
Diplodina Westendorp
Pycnidia immersed
It differs
(p.
505)
ostiole
or
erumpent, globose;
papillate,
black, small; spores elongate.
from Diplodia only

species chiefly
in the hyaline spores.
About eighty
saprophytes.
D.
citrullina
(C.
O. Sm.)
Gres. on cucurbs=Mycosph8erella ciruUina. See p. 246.
D. castaneae P.
jures
loss.i2
&
D.
in-
Fig.
chestnut
leaves,
and
shoots
in
D. castanca;. pycnidium 352. After Delacroix. spores.
and
causes cankers on the
France,
resulting in serious
D. parasitica (Hart.) Prill, occurs on the basal leaves of young shoots of spruce causing defoliation. D. salicina C. & M. causes tips of willows to die.
D. corticola A.
&
S. is
found on cacao
in Africa.
Sphaerioidaceae-Phaeodidymae
Conidia dark, 1-septate, ovoid to oblong.
(p.
480)
510

Key
to Sphaerioidaceae-PhaBodidymae
Pycnidia separate
Pycnidia beaked Pycnidia hairy

1.
Rhynchodiplodia,
Pellioniella.
p. 510.
Pycnidia smooth Pycnidia not beaked

Pycnidia hairy
2.
3.
Chaetodiplodia,
p. 510.
Pycnidia smooth Conidia with a mucous layer, very large Conidia without a mucous layer
Pycnidia erumpent, muticate
conidia
4.
Macrodiplodia.
Conidia less than 15 m long. Conidia 15 m or more long.

Pycnidia superficial, lignicole. Pycnidia cespitose or in a stroma
Pycnidia cespitose Pycnidia in a stroma
5. 6.
Microdiplodia,
Diplodia,
Diplodiella,
p. 510.
p. 511.
p. 512.
7.
8.
Botryodiplodia,
p.
513.
Pycnidia and subicle enclosed in a hemispheric stroma Pycnidia
9.
Lasiodiplodia,
Diplodiopsis.
p. 513.
without
subicle,
in
a
10.
globose stroma
Rhynchodiplodia Briosi
&
Farneti
Pycnidia rostrate, pilose; conidia oblong. single species, R. citri B. & F., causes disease of the lemon.
Chaetodiplodia Karsten
Pycnidia erumpent, globose, ostiolate, black, carbonous, hairy or bristly; conidia elongate. A genus of about ten species, chiefly saprophytes.
C. vanillas
membrano-
Zimm.
is
on
vanilla.
Microdiplodia Allescher
ous, globose or depressed,
Pycnidia subcuticular, erumpent, membranous to subcarbonminutel}"^ ostiolate; conidia ovoid to

/jl.)
oblong, small, (under 15

More than twenty-five
species, chiefly saprophytes. anthurii Trinch. occurs on Antliurium.
511
M.
Diplodia Fries
p3'-cnidia
(p.
510)
black,
immersed,
erumpent,
ellipsoid
carbonous,
or
usually
ostiolate-papillate;
conidia
ovate;
conidiophores
needle-shaped, simple, hyaline. Over four hundred fifty species,
many
of
them saprophytes.
D. zese (Schw.) Lev-^^^-^"^ On ears and stalks of corn, pycnidia borne on the husks, cobs, stalks and rarely the grains, gregarious, small, lenticular to flaskshaped or
irregular, papillate; conidia elliptic, straight or curved, constricted or not, 25-30 xQ It occurs as the cause of a very serious dry rot of ear corn. The actual growing mycelium is hyaline and much branched.
jj,.
Pycnidia in the cob are principally on the scales which surround the inner ends of the kernels and are set in a dense mass of white
mycelium.
the
On dead
stalks
pycnidia form below

rind,
the
particularly
at
the
nodes,
breaking
through during the following summer, and extruding

the spores in
cirri.
The fungus was studied
were made using pure cul- Fig. 353. Pycnidia of Diplodia, from corn kerAfter Burrill and Barrett. nel. tures. Spores placed under the husk or in the silk, or sprayed upon plants in suspensions, resulted in disease.
extensively by Burrill & Barrett ^^ and inoculations
Smith and Hedges '"^ report that infection is often by way of the root system, the mycelium reaching the grains through the stem and from the cob. D. macrospora Ea.
gate, irregularly clavate, curved or constricted,
Pycnidia scattered, large, erumpent, carbonous; conidia elon70-80 x 6-8 m-
512

is
This
responsible for a corn
mold
similar to that caused
by
the last species. ^^
Other parasitic species are: D. oryzae Miy. on rice;
D. cerasorum Fcl. on cherries; D. aurantii Catt. on oranges; D. mori West, on Morus; D. gongrogena Temme on Populus in Germany; D. sapinea (Fr.) Fcl. on conifers; D. pinea Kick, on pine leaves in Europe; D. coffeicola Zimm. on coffee; D. perseana Del. on the avocado. D. opuntiae Sacc. is sometimes a serious pest of the cactus. D. citricola McA. occurs in Australia on lemon twigs, stems and green fruit. ^"^ D. destruens McA. is on orange and lemon leaves in Australia; ^* D. heteroclita D. & M. on Citrus in Algiers.^"* D. cacaoicola Hen. does much injury to cacao and sugar
cane
^''^
in the
West
Indies.
of citrus fruits in the
D. natalensis Ev. causes a serious black rot

Transvaal;
^"^
Pycnidia scattered, covered, later erumpent, black; papillate 150-180 n; spores elliptical, 1-septate, not constricted, dark, 24 X 15 n, exospore with striated bands.
Diplodia which cannot be distinguished from this was studied Fawcett and Burger and is reported as the cause of gumby mosis of peach and orange in Florida.-^^ Pure culture inoculations and cross inoculation showed the same fungus able to cause the disease on both hosts. D. rapax Mas. is the cause of a stem disease of Para rubber. D. epicocos Cke. grows on the coconut and an undetermined
species attacks ripe pineapples.
Diplodiella Karsten (p. 510)
Pycnidia superficial, globose, ostiolate papillate, black, smooth, rather carbonous; conidia elliptic.
About twenty-five
D. oryzae Miy.
is
species, chiefly
rice.
saprophytes on wood.
found on
513
Botryodiplodia Saccardo
(p.
510)
Pycnidia botryose-confluent, erumpent, stromatic, membranocarbonous, black, usually ostiolate-papillate; conidia elongate or
ovate.
Over thirty
species of this genus is given by Butler as the ^"^ probable cause of a coconut palm disease in India.
An unnamed
Lasiodiplodia EUis
&
Everhart
(p.
510)
Pycnidia collected on a stroma, covered with a brown mycelium, paraphyses among the conidiophores. Otherwise as in Diplodia.
Two
species,
both parasites.
L. tubericoIaE.
AE.""
Pycnidia globose, 250-305 n; stromatic mass about 1 mm. in diameter; conidia elliptic, 18-22 x 11-14 /x, not constricted;
conidiophores short;
ju
paraphthe
Fig.
354.
yses
It
45-55
long,
overtopping
conidia.
Lasiodiplodia
tu-
was found on sweet potatoes from Java which were brought to the
Perithecium, paraphyses and spores. After Clendenin.
bericola.
Louisiana Experiment Station in 1894. L. theobromae (Pat.) G. & M. is a wound parasite of Hevea
Sphaerioidaceae-Hyalophragmiae
(p.
480)
Conidia hyaline, 2 to many-septate, oblong to fusoid.
Key to Genera of SphaBrioidaceae-Hyalophragmiae

Pycnidia more or
Subicle none
less
globose
Conidia appendaged at apex

Setaj
1
1
.
Kellermania.
Bartalinia.
Setse 3
2.
ol4

Conidia muticate
3.
4.
Subicle present, dark, phyllogenous
Stagonospora, p. 514. Asterostomidium.
Pycnidia elongate to cylindric
5.
Mastomyces,
p. 514.
Stagonospora Saccardo
;^
Pycnidia superficial or erumpent,

globose,
ostiolate-papillate,
c^:^^9 .ajT^ ^ '^
black,
co-
membranous
or
subcarbonous;
nidia elongate, 3 or more-celled.
^
^^^^
^~~^
Fig. 355.
^^'^
Over one hundred
species, chiefly
"^^
l'"^
*^^ V7r77=^'-*^ \T?

'^f^TrS^
C, pycD, spores,
saprophytes; differing from Hendersonia onh^ in the hyaline conidia. S. carpathica Bsouml.
Stagonospora.
in section. ^ ^'
Spots circular, 1-3 mm., light brown with a narrow darker border:
nidium
pycmdia 120-180
in
i-
i r.rx
n,->
fj,]
comclia escapmg
slightl}'^
constricted.^^
curved, 14-28 x 4 ;u, It causes leaf spots on alfalfa. S. iridis Mass. occurs on iris.
gelatinous mass, straight or 2 to 5-celled, frequently slightly
Mastomyces Mont.
Pycnidia gregarious, separate,
erumpent, elongate, papillate-
ostiolate; conidia fusiform, 3-septate.
There are two species, one of which, M. friesii Mont., is probably the conidial form of Scleroderris ribesia, see p. 155, the cause of a relatively unimportant currant disease of Europe.
Sphaerioidaceae-Phaeophragmiae
(p.
480)
Conidia hyaline, 2 to several-septate, oblong to f usoid.
Key
to Genera of Sphaerioidaceae-Phseophragmise
Pycnidia .separate Pycnidia not beaked

Conidia free from each other
Conidia muticate
Pycnidia papillate or subastomous Pycnidia with flattened base
1.
Macrobatis.

Pycnidia globose, without flattened base but on a stellate
superficial subiclc
2.
515
Couturea.
Pycnidia
without
subiclc,
crumpcnt
Pycnidia hairy Pycnidia smooth Pycnidia opening widely, with an
3. 4.
Wojnowicia. Hendersonia,
p. 515.
operculum
Pycnidia superficial, dark, hairy Pycnidia immersed, pale, smooth Conidia appendagcd
5. 6.
Angiopoma.
Lichenopsis.
Cryptostictis,
p.
Conidia
Conidia
1-ciliate at
1-ciliate at
each end
base
7. 8.
516.
Urohendersoria.
Santiella.
Conidia with a round or cup-like appendage at each end
9.
Conidia united in groups Conidia united into a fascicle

Conidia stellately united Pycnidia beaked
Pycnidia in a stroma
10.
11. 12.
Eriosporina. Prostheir*'-m.
13.
Pseudographium. Hendersonula, p. 516.
Hendersonia Berkley
Pycnidia
immersed,
erumpent
or
not,
globose with a papillate ostiole or depressed, membranous or subcarbonoiis; conidia elon-
gate or fusoid, 2 to many-septate.
Some two hundred

sitic species.
fifty
species,
chiefly
saprophytic, although there are several para-
H. mall Thiim.
large. 1111 pycnidia disciform, 1-1 on brow^msh, orbicular, scattered, black,
,,
1
Epiphyllous;
Fig. 356.
Hondc^-
sonia, hymcnium and spores. After

Starback.
violet-margined spots; conidia clavate, apex rounded, base somewhat acute, not constricted, 12-14 x 4-5
/x.
On
H. H. H.
leaves of apple in Europe and America. piricola Sacc. is on pear; H. cydonae C.

acicola
&
E. on quince;
M. &
T. causes a pine leaf disease. ^^^
coffege Del.
is
on
coffee;
516
H. oryzae Miy. on rice; H. notha Sacc. & Br. on Juniperus H. togniniana Poll, on Cycas. H. foliicola (Berk.) FcL^^^
leaves;
Pycnidia epiphyllous, brownish-black, subglobose, subelliptie or irregular; conidia elliptic to clavate, obtuse, 3 to 5-septate; conidiophores filiform, radiating.
On
Juniperus and Pine.
Cryptostictis Fuckel (p. 515)
erumpent, globose or depressed, conidia elongate, 2 or more septate, ostiolate;

Pycnidia
subapically appendiculate with a long hyaline
bristle.
It differs
from Hendersonia
chiefly in
spore characters. A small genus of eleven species, chiefly sapro357. Cryptos- phytes. ^^' ^' cynosbati (Fcl.) Sacc. and tif StaTback C. caudata (Preu.) Sacc. occur on the rose, the former on the fruit and branches forming wounds.
FiG.
Hendersonula Speg. Stroma black,
(p.
515)
irregular; ostioles punctiform; spores ellipsoid,
several-celled, colored.
In part =Plowrightia.
A form
on the plum=Plowrightia morbosa.

Sphaerioidaceae-Phaeodictyae
See
p. 218.
(p.
480)
Conidia dark, muriform, oblong to ovoid, rarely radiate or

cruciate.
Key
to Genera of Sphaerioidaceae-Phaeodictyae
Pycnidia separate Conidia not reticulate Pycnidia on bark, erumpent Pycnidia on wood, superficial Conidia reticulate Pycnidia merely locules in a stroma
1.
Camarosporium,
Cytosporium, Endobotrya. Dichomera.
p.
517.
2. 3.
4.

Camarosporium Schulzcr von Miiggenburg
Pycnidia
globose,
(p.
517
516)
immersed,
erumj)ent,
to
ostiolate,
papillate,
carbonous
separate, or
submembranous;
conidia ovate
fusiform,
muriform, with 2 to
saprophytes. C. fissum
roses.
many
cross walls.
species,
Over one hundred twenty

(Pers.)
chiefly
Star,
causes
injury
to
C. viticola (Cke.
&
T\/r
H.) Sacc.
is
reported on
grape
.
mon
Sacc. on MorUS.
CamaroFig. 358. sporiuni, spores. After Starback.

(p.
Sphaerioidaceae-Scolecosporae
480)
Conidia hyaline or light colored, elongate-fusoid, rod-shaped or filiform, continuous or septate.
Key
to Genera of Sphaerioidaceae-Scolecosporae
Pycnidia separate Pycnidia membranous or carbonous Pycnidia superficial
Pycnidia hairy Conidia single on the conidiophores Conidia ternate on the conidiophores
1.
Trichocollonema.
2.
Gamospora.
Cornularia.
Pycnidia smooth Pycnidia beaked Pycnidia not beaked

Conidia not expelled
3.
Conidia usually expelled in a ball in a ball.

.
4.
5.
Collonema.
Septorella.
p. 518.
ycnidia immersed or erumpent Pycnidia hairy, maculicole
6.
Trichoseptoria,
Pycnidia smooth Pycnidia beaked Pycnidia not beaked

Pycnidia
maculicole,
7.
Sphaerographium.
mainly
8.
9.
phyllogenous Conidia hyaline
Septoria, p. 518.
Conidia colored
Phaeoseptoria,
p. 525.
518

Pycnidia not maculicole
Pycnidia
complete
at
top,
10.
usually papillate Pycnidia more or less incomplete at top
Rhabdospora,
p. 525.
Pycnidia gaping, showing a gelatinous spore mass.

Pycnidia not exposing a gelatinous
11.
Gelatinosporium.
mass
12.
.
Pycnidia
foliicole
Phleospora,
Phlyctaena.
p. 525.
Pycnidia rami-caulicole.
Pycnidia cespitose
13.
Pycnidia suberose, incomplete, often pale

14.
15.
Pycnidia merely gregarious Pycnidia in a stroma
Micropera. Micula.
Conidia 4 to 6-fasciculate on a ccnidio-
phore Conidia separate

Conidia setose-penicillate Conidia muticate
16.
Eriospora.
17.
Bilophospora,
p. 525.
Stroma superficial, setose Stroma erumpent or immersed

in the stroma, conidia hyaline Pycnidia as locules in the stroma, conidia colored
18.
Septodothideopsis.
Pycnidia distinct
19.
Cytosporina,
p. 526.
20.
Septosporiella.
Trichoseptoria Cavara
(p.
517)
Pycnidia separate, erumpent, on spots, membranous, hairy; conidia needle-shaped, septate. A single species. T. alpei Cav.^^^ is reported by Cavara as
injurious to
lemon
fruits in Italy.
Septoria Fries
(p.
517)
spots, globose lenticular,
Pycnidia immersed, usually on

ostiolate,
leaf
membranous, black; conidia narrowly elongate

species, all parasitic,
to
fili-
form, multiseptate, hyaline, conidiophores very short.
Over nine hundred

non-economic hosts.
several of
considerable economic importance but most of
them of them occurring on
519
In part=Mycosphscrella, LeptosphaTia. is a very large one similar to Phoma and Phyllosticta except in its spore form and in the ostiole which is frequently
The genus
Septoria and Phleospora are distinguished onh' by large. the lesser development of the walls of the latter and many species which in early stages pass as Phleospora would in older stages be
very
classed as Septoria.
Septoria and llhabdospora are distinguished only

of the host affected,
by the part
in these
stem or
leaf,
and many forms
two
genera are undoubtedly identical. S. pisi West, is on peas.

S. piricola
Desm. on pear and apple = My cosphserella
sentina.
See
p. 246.
S. populi
S. phlogis Sacc. p. 258. S. ribis Desm.^^^
Desm. on Populus=Mycosphserella populi. See p. 250. & Speg. on Phlox=Leptosph8eria phlogis. See
liypophyllous; spots small, irregular, bounded by the leaf veins,
brownish-purple;
pycnidia
in-
nate, minute, convex, brownishblack; cirri in mass reddish; co-
nidia elongate,
linear,
curved.
50 n long.
On
tion.
gooseberry and currant,
causing leaf spots and defoliaS. aciculosa E.
& E.
Fig.
359. S.
^p^^^-
libis,
Pycnidia innate to superficial,
a pyciuduun ^^^'^ Longyear.
ai,d
grouped, minute, amphigenous; conidia needle-shaped, continuous, 15-20 X 0.75 M.

It
is
found on the strawberry.
S. fragariae
Desm.
brown, with reddish-brown margin; pycnidia minute, innate, prominent, brownish; cirri white;
Epii)hyllous; spots suborl)icular,
conidia cylindric, obtuse, 3-septate.
Perhaps =Mycosphserella
fragariffi.
See
wild,
p. 244.
On
spots.
strawberry,
cultivated
and
forming circular
leaf
520
S. cerasina Pk.^"
Spots scattered or confluent, minute, subangular, brown or reddish-brown; pycnidia pale, collapsing; conidia filiform, straight
or curved, 50-75 S. pruni E.^^^
/i
long.
On
cherry.
Spots dark brown, dry, subrotund, soon breaking out, 1-3 mm. pycnidia brown, immersed, 60 m; conidia linear, obtuse, 4 to 6-septate, 30-50 x 2 /x. On plum.
;
Pass, and S. sicula Penz, occur on citrus; glaucescens Trab. on the mandarin; S. loefgreni N. on oranges in Brazil;
S. S.
limonum
S.
ampelina B.
&
C. on the grape.
is
S. longispora Sh. (not Miy.)
found on the cranberry.^^^
S.
graminum
Desm.^^"^^^
Spots slightly elongate, pale, fuscous-margined, limited by the

leaf veins;
slender,
straight 55-75 X 1-1.3 M.
pycnidia seriate or scattered, brownish; conidia very or curved, non-septate, but multiguttulate,
This is a frequent saprophyte or weak parasite on wheat, oats and numerous wild grasses. Under some conditions it becomes an injurious parasite, especially upon winter wheat. S. tritici Desm. is closely like S. graminum. ^^^ It is associated with Leptosphseria tritici on wheat. See
p. 258.
S. S.
glumarum Pass, is also found on wheat. nodorum Berk, occurs, particularly at the
nodes, on the
same
host.
is on wheat and rye leaf sheaths; on leaves of oat. S. longispora Miy. (not Shear) and S. curvula Miy. are on
S. secalina Jancz. S. avenae Frank,
rice.
S. betae
West.
Spots pale brown, white in the center, brownish-margined;

pycnidia epiphyllous, minute, black, prominent; conidia cylindric, straight or curved, white in mass.
It
was noted by Humphrey
^^^
causing a beet leaf spot.
S. citruUi E.
&
E.
Spots small, round, white, scattered; pycnidia mostly solitary,
521
one in the center of each spot, but slightly prominent; conidia cylindric or clavate-cylindric, 10-25 x 1.5-2 ju.
On watermelon
S.
leaves.
cucurbitacearum Sacc. is also on cucurbits. ^^ S. cannabina Pk. is on hemp producing leaf spots. S. nicotianae Pat. is reported from France as the cause of tobacco
leaf spotting.^^'^
S. dolichi B.
&
C.
Spots white, with a broad, light brown margin; conidia straight,

subfusiform, 3-septate, 40 n.
S. medicaginis
On
cowpeas.^^^
Rob.
& Desm.
is
on
alfalfa.
Spots whitish, angulate-subcircular, confluent; pycnidia lenticular, 70-90 m; conidia slender, vermiform, tortuous, 60-70 x 1 ju,
septate. S. petroselini
Desm.
amphigenous; pycnidia epiphyllous,
promi-
Spots brown, in age white,

minute,
olivaceous,
nent; conidia filiform, straight or curved, 35^0 x 1-2 n. On

parsley. S. petroselini
apii
Br.
&
Cav.25.
122-126
This
leaves
Italy
common and
fungus
first
very decelery
structive
on
was
described in
America
Cavara and in was early noted ^22 and Halsted.^'^ by Chester The pycnidia are abundant in ^ t, riG. 360. b. petroselini apu. Pycnidium v J the leaf spots and m the case showing spores oozing through the ostiole.
by
it
o,
celery they are found scattered over the blanched petioles.

of
Til
stored
.1
After Jensen.
Essentially
it is
only
a host variety.
S. lycopersici Speg.^^'
^^^'
^^^
Spots large,
cinereous,
often confluent
and covering the
entire leaf, sordid
subindeterminate; pycnidia scattered, hypophyllous,
lenticular-hemispheric, prominent, membranous; conidia elongate, cylindric, 70-110 x 3.3 m, pluriseptate.
522

It was noted in of leaf spots of tomato. Jersey and in Ohio in 1896.^-^ It is one of the serious
The cause
about 1893
New
^-^
tomato
pests,
S. lactucae Pass.''^^
entire
Spots irregular, brownish, angulate, sometimes destroying the punctiform, scattered, 90 /x in leaf, pycnidia minute,
diameter, conidia filiform, straight or curved, 25-
30 X 1.7-2
/x.
On
lettuce.
S. consimilis E.
&
M.^^
Distinguishable from the preceding by the more

indefinite spots, slightly larger pycnidia (90-100 /x) and longer conidia (30-45 x 2-2.5 fx). It causes
brown spots on
Fig.
lettuce leaves.
36i.-Spore3
.
S. armoraciae Sacc.^^
of S. lycopcrsici.
After Reed.
form, grouped
Spots irregular, ochraceous; pycnidia puncti^ / nr. the center ot the spot, oO mJ
i
m
.
conidia filiform, curved, 15-20 x 2-2.5 ix, 1 to 3-septate. horseradish causing leaf spot. ^^^ S. antirrhini Desm. attacks the snapdragon severely.
S. rosae S.
On
Desm.
is
on
rose;
hydrangeas Bizz. on cultivated hydrangea; S. iridis C. Mass. on Iris. S. cyclaminis Dur. & M. on cyclamen. S. sedi West.
Epiphyllous; spots circular, gray to gray-brown; pycnidia minute, numerous, brown, scattered, erumpent; conidia cylindric, straight or curved, 5-guttulate; cirri white. On cultivated Sedum.
S. hederae
West,
is
S. rostrupii Sacc.
&
on Hedera; Syd. and S. varians

^^
Jaff.
on chrysanthe-
mum;
S.
as
is
also:
Spots ochraceous, dark margined; pycnidia epiphyllous, punctiform; conidia 40-50 x 2.5-2 n,
chrysanthemella Cav.^^'
obscurely septate. It causes damping off of chrysanthemum cuttings and spotting of the leaves.
^'^^
S. dianthi Desm.^^
Spots yellowish, oblong, roundish or irregular; pycnidia globose,
523
depressed, brownish-black; cirri white; conidia elongate, cylindric, curved, obtuse, 30-45 x 4 mIt is the cause of a common and injurious leaf spot of the carnation.
The pycnidia
are visible as dark specks
on the blanched
background
of the spot.
S. azaleae Vogl.
Spots reddish-yellow; pycnidia amphigenous, immersed, globose

to depressed, black; conidia oblong cylindric, filiform, straight or curved, 1 to 3 or more septate, constricted slightly at the septa,
12-18 X 1.5-2.5
/x;
conidiophores cylindric, short, 3-5 m long.
On
Azalea.
S. divaricatae E.
&
E.
Spots whitish, amphigenous, confluent, purple-bordered; pycnidia numerous, epiphyllous, lenticular, 100-120 m. dull black; conidia 18-30 x 0.75-1 ix, nearly straight, non-septate, finely
guttulate.
-^^ frequently injures cultivated phlox. S. narcissi Cass, is on Narcissus.
It
S. exotica Speg.
is
on Veronicas
in cultivation.
S. fairmanii E.
&
E.
Spots amphigenous, scattered, subangular, 3-4 mm. dark brown, limited by the veins, with a narrow dark margin; pycnidia epiphyllous, scattered, rather numerous, black, subprominent, 100-112 n; conidia filiform, slightly curved, guttulate, 30-45 x
1.5-2
M.
^^ hollyhock leaves.
It parasitizes
S. parasitica Fau.
cylindric,
Spots amphigenous, white; pycnidia punctiform, innate; conidia 30-40 x 3.5-4 m-
conidia are broader than in the preceding species and the gross appearance is quite different. It is found associated v/ith rust sori on hollyhock. ^^
S. helianthi E.
The
&
K.
with a yellowish elevated margin; pycnidia epiphyllous, immersed, brown, collapsing, 105 /x; conidia linear-filiform, 3 to 5-scptate, 30-70 x 2-3 /x-^^^
Spots brown, definite, 2.5-7.5
mm.
On
sunflower leaves.
S. majalis Aderh. causes a leaf spot of lily-of-the-valley;
524

In leaf spots of oleander.
S. oleandrina Sacc. S. veronicae
Desm.
amphigenous, small, subrotund, brownish or grayish, becoming white, border umbrinous; pycnidia epiphyllous, gloSpots
bose, prominent, pale brownish-black; conidia elongate, slender, straight or flexuose.
It is parasitic on cultivated Veronicas.^"* S. caraganae Hen. is on Caragana.
S. ochroleuca B.
&
C.
Spots scattered, suborbicular, pale, brown margined; pycnidia

central, minute, scattered, hypophyllous, pale, collapsing; conidia
fusoid-filiform, curved, continuous or 1-septate, 25
ju
long.
In leaf spots on chestnut.

S. castanicola
Desm. and
S. castanea Lev. are
on chestnut;
S. nigro-maculans Thiim. on walnuts and horse-chestnut; S. aesculi Lib. and S. hippocastani B. & Br. on horse-
chestnut.
S. pseudoplatini R.
&
D. occurs on sycamore and maple;
S. fraxani
Desm. on ash;
on Cercis;
S. cercidis Fr. S.
tiliae
West, on Tilia; S. curvata (R. & B.) Sacc. on Robinia leaves.

S.
spadicea P.
&
C.^^^
causes
common
twig blight of
pine.
late, becoming slightly erumpent browning needles, scattered, membranous, fuscous-olivaceous, subimmersed, 190-225 /x in diameter. Spores hyaline, cylindrical, slightly curved or flexuous, apex acute, 1-septate, rarely constricted at septum, 3-4 x 30-45 n. Conidio-
Pycnidia not spot-forming,

of
on inner surface
phores short. S. ulmariae Oud.
Pycnidia minute, immersed, spores cylindric, curved, hyaline, continuous, 5.0 x 2.5 m- On Spirea. S. cornicola Desm.
Spot orbicular, margin dark purple; pycnidia epiphyllous, few, black; spores cylindric, curved, 35-40 x 2-2.5 n, obsolete 2 to 4septate, hyaline. On Cornus. S. parasitica Hart, is found
lateral shoots.
on young spruce buds
killing the
525
Phaeoseptoria Miyabe
(p.
517)
As
in Septoria but with colored conidia. P. oryzae Miy. is on rice in Japan.
Rhabdospora Montaigne
(p.
518)
Pycnidia innate, erumpent, globose or depressed, brown or

black; conidia as in Septoria. Similar to Septoria, but on stems.
and R. cofifeae Del. are on & S. on cacao; R. oxycocci Sh. on cranberry.

R.
cofifeicola Del.
coffee;
R. theobromae A.
R. rubi E. Pycnidia black, subglobose, innate, erumpent, scattered, 100-195 m; conidia linear, curved, 3 to 4-septate, 4045 X 3 /x. On blackberry.
Phleospora Wallroth
(p.
518)
Pycnidia innate, imperfectly developed, and chiefly formed of modified host tissue; conidia elongate-fusoid, thick, 2 to manyseptate.
About twenty-five species of leaf parasites. This genus closely approaches the Melanconiales in structure. Several forms have been shown to be allied to Mycosphaerella, e. g.,
P. ulmi to
M.
ulmi.
P. mori (Lev.) Sacc. on Morus=Cylindrosporium

cosphsella.
mori=Myof
P. moricola (Pass). Sacc. on Morus is a conidial form Septogloeum mori. P. aceris Lib. is found on maple and sycamore leaves; P. oxycanthae Desm. on hawthorn leaves; P. caraganae Jacz. on Caragana.^^^
Dilophospora Desmazieres
(p.
518)
unicellular, with hair-like
Pycnidia globose, ostiolate, usually stromatic; conidia cylindric, appendages at each end. In part = Dilophia. D. graminis Desm. = Dilophia graminis. See p. 257.
526
Cytosporina Saccardo
(p.
518)
Stroma valsoid, cushion-formed or tubercular; pycnidia sunken, the ostiole erumpent; conidia fiUform, curved, 1-celled.
Twenty
These
species of bark
and wood inhabiting
fungi.
forms of the Valsacese. C. ribis Miy.^^^ occurs on currant and -gooseberry bushes in Holland attacking the cortex, later the wood, and killing the
are, in part at least, conidial
shoots.
Nectrioidaceae
(p.
479)
Pycnidia fleshy or waxy, light colored, globose, rarely cup-
shaped or hysterioid; stroma present or absent; conidia various,

usually hyalin^. This group contains
fungi Aschersonia
some twenty-five genera none
of
which are
serious plant pathogens.
Some
are conidial forms of the ascigerous
and Polystigma.
Key
to Subfamilies and Groups of Nectrioidaceae

I.
Pycnidia globose, ostiolate Conidia 1-celled
Zythieae.
p. 526.
Hyaline
1.
Hyalosporae,
Phaeosporae.
Dark
colored
2.
Conidia two-celled hyaline Conidia 3 to several-celled, hyaline

Elliptic to fusoid
3.
Hyalodidymiae.
4.
5.
Hyalophragmiae.
Scolecosporae.
Olluleae.
Bacillar to filiform
Pycnidia cupulate or hysterioid
II.
Zythieae-Hyalosporae
Conidia hyaline, continuous, ovoid to
elliptic.
Key to Genera of
Pycnidia separate Pycnidia smooth Pycnidia beakless Conidia in chains
Zythiaceae-Hyalosporse
1..
Sirozythia.

Conidia not in chains
Pycnidia on creeping hyphse Pycnidia without mycehum
2.
527
Eurotiopsis.
Conidia spiny or Conidia spiny
ciliate
3. cilia
Roumegueriella.
Ciliospora.
Conidia with several
at
4.
apex Conidia smooth

Pycnidia single-walled Pycnidia more or
late
less papil-
5.
Zythia,
p. 527.
Pycnidia
with cratcriform
6. 7.
ostiole
Libertiella.
Pycnidia cup-shaped Pycnidia with outer circumscissile wall
Lemalis.
Dichlaena.
8.
Pycnidia beaked.
9.
Sphaeronemella.
Pycnidia hairy or spiny Pycnidia densely

1-cellcd seta?
beset
with
conoid
10.
Muricularia.
Pycnidia with slender bristles or hairs Hairs fasciculate

Hairs separate Hairs everywhere but at the apex Hairs only around the wide ostiole
11.
CoUacystis.
Chaetozythia,
12.
13.
Pseudozythia.
Pycnidia cespitose or in a stroma

Pycnidia cespitose, beaked;
chains
conidia
in
14.
Treleasiella.
Pycnidia in a stroma
Stroma more or
Conidia fusoid
less
pulvinate
15. 16.
Aschersonia.
Conidia globose
Munkia.
Hypocreodendron.
Stroma
fruticose,
branched;
conidia
17.
bacillar
Zythia Fries
Pycnidia erumpent or superficial, globose, with more or less evident papillate ostioles, white or bright colored; conidia ovate
or elongate.
Some twenty
species.
528

Is said to cause a
Z. fragariae Laib.
strawberry disease.
(p.
Leptostromataceae
Pycnidia membranous
479)
or carbonous, black,
more or
less dis-
tinctly dimidiate, scutiform, astomous, ostiolate or cleft, or superficial. Over two hundred species.
erumpent
Key
Conidia 1-cellcd
to Sections of Leptostromataceae.
Hyaline Colored
Conidia 2-celled
1.
Hyalosporae,
Phaeosporae,
p. 528.
2.
p. 531.
Hyaline Colored
Conidia 3 to several-celled
3. 4.
Hyalodydimae. Phaeodidymae.
Hyalophragmiae,
Phaeopharagraiae.
Scolecosporae,
p. 532.
Hyaline Colored
Conidia
1
5.
6.
p. 531.
to several-celled, filiform
7.
Leptostromataceae-Hyalosporae
Conidia hyaline,
1-celled, globose to ovoid.
Key
to Genera of Leptostromataceae-HyalosporflB
Pycnidia separate Pycnidia astomous or variously perforate, but not cleft
Conidiophores lacking Pycnidia on a subicle

Subicle of fumaginous hyphse Subicle of broad fibers
1.
2.
Eriothyrium. Trichopeltulum.
Pycnidia without subicle Conidia muticate
Pycnidia stellately divided

cleft
or
3.
Actinothecium.
Pycnidia depressed-clypeate, not

stellate
4.
5. 6.
Leptothyrium,
Tracyella.
Piggotia,
p. 529.
Conidia setulose at each end

Conidiophores present, cyhndric
p. 530.

Pycnidia more or
wise
less clearly cleft length-
529
Pycnidia elongate or lanceolate

Pycnidia subcircular Pycnidia in a stroma
7. 8.
Leptostroma,
p. 530.
Labrella, p. 530.
Stroma phyllogenous Stroma growing on animal
9.
Melasmia,
p. 530.
hairs
10.
Trichophila.
Leptothyrium Kunze
&
Schweinitz
(p.
528)
Pycnidia superficial or erumpent, dimidiate, scutiform, membrano-carbonous, black, coalescing or scattered, ostiole variable,
structure cellular; conidia ovoid-oblong to fusoid.
Some one hundred species. In part=Gnomonia and Gnomoniella.

L.
alneum
tubiformis.
L.
_,
1*
.
(Lev.) Sacc. Seep. 274.
on alder = Gnomoniella
^^^
1
111 ! black spots pycnidia differentiated in late winter, 25-100 fi; conidia el.
pomi (M.
&
F.) SaCC.^^^'
n
Fm. 362.-LJptothyrium oxyCOCCI.

spores.
^^''
;
Four
ormmg mniute superncial

12-14 X 2-3
is
ix.
After
liptic,
The mycelium
of the spots breaks
away
and probably functions
The fungus
as a reproductive body. common in sterile form on pomaceous fruits but
the sporing stages are rarely found. L. periclymeni Desm., L. acerinum Ktz. and L. buxi Pass, are
on Lonicera, acer and box respectively.

L. oxycocci Sh.
^^'^
Pycnidia
black,
dimidiate, amphigenous, scattered, subcoriaceous to coriaceous, irregularly subglobose,
subepidermal, erumpent, rupturing irregularly; conidia subfusoid, sometimes slightly curved,

jo-îi~r^ôa^
Fig.
pseudoseptate, 10-15 x 2.5-3
/z;
conidiophores
363.-ê"pto- simple, straight, tapering, slightly longer than thyrium oxycocci. ^k '^'^^ pp,p;f1| COniQia. Section of a
pycnidium, showing its dimidiate char-
Qn
L.
Cranberry.
. .
actor. ^^"
After
macrothecium
Fcl.
is
said
to cause a
strawberry disease.
L. peonae Br. & Cav. is on peony leaves. L. parasiticum Poll, is on Cereus stems in Italy.
530

species has been reported as a "fly speck" on
An undetermined
cabbage.
^^^
Piggotia Berkley
& Brown
(p.
528)
Pycnidia applanate, inequilateral, thin-membranous, later with a stellate cap which is thrown off forcibly; conidia elongate or
cylindric.
A genus
of less
than ten
species.
P. astroidea Berk.
P. fraxini B.
&
Br. parasitizes elm leaves.
&
C.
fx
Perithecia hypophyllous; spores oblong, 5-7
long.
On
ash,
causing leaf spot.
Leptostroma Fries
(p.
529)
Pycnidia dimidiate, subsuperficial, applanate, elongate, black,
more
In part = Hysteriaceae.
L. larcinum Fcl.
or less hysterioid; conidia ovate, elongate or allantoid. There are some sixty species.
on larch =Mycosphserella
larcina.
See
p. 249.
L. piricola B. & S. occurs on the pear; L. punctiforme Wallr. on willow.
LabreUa Fries
(p.
529)
Pycnidia black, round, often indefinite; spores long, fusiform,

or spherical, hyaline, continuous. L. piricola Bres. & Sacc. is on pear leaves; L. coryli (Desm. & Rob.) Sacc. on Corylus.
Melasmia
Levielle (p. 429)
Pycnidia dimidiate, carbonous, black, often on an effused black

stroma; conidia allantoid. In part=Rhytisma. Over twenty species. M. acerina Lev. is the conidial form of Rhytisma acerinum. See p. 158. M. punctata S. & R. and M. salicina Lev. of the two
corresponding Rhytismas.
See
p. 158.
531
Leptostromataceae-Phaeosporae
Conidia globose to oblong,
1-celled, dark.
(p.
528)
Key to Genera
of Leptostromataceae-Phaeosporae
Pycnidia separate Pycnidia on a dark subiclc, radiatcly dehiscent

1.
Asterostomella.
Pycnidia not on a subiclc Conidia conglobate, verrucose
2.
Conidia not conglobate, smooth Pycnidia in a stroma
3.
Discomycopsella. Pirostoma, p. 531.
Stroma membranous
Pj'cnidia distinct, exserted
4.
Peltostroma.
Pycnidia merely locules, immersed.
...
5.
Lasmenia.
Poropeltis.
Stroma carbonous; immersed
locules
many,
6.
Pirostoma Fries
Pycnidia separate, shield-shaped, rounded or elongate, leathery, conidia globose to ellipsoid.
genus of
less
than
five species.
P. farnetianum Poll, occurs on Pandanus in Italy.
LeptostromataceaB-Hyalophragmiae
(p.
528)
Conidia oblong to fusoid, hyaline, 2 to several-septate.
Key
to Genera of Leptostromataceae-Hyalophragmiae
Pycnidia a.stomous or ostiolatc, not cleft Conidia muticate; pycnidia with creeping
hyphoe Conidia ciliate
1.
Asterothyrium.
Discosia,
p. 531. p. 532.
Conidia fusoid, 1-ciliate at each end. Conidia cruciate, each arm 1-ciliate.
Pycnidia rimose dehiscent
2. 3.
Entomosporium,
Cystothyrium.
4.
Discosia pini Heald has been reported as a parasite on pine

hypocotyls.
532
Entomosporium
4-celled, cruciate,
Levielle (p. 531)
Pycnidia depressed, sul)globose, not ostiolate, black; conidia each arm 1-ciliate.
A genus of
three species of parasites.
In part=Fabrea.
E, mespili (D. C.) Sacc. =F. mespih. See p. 150. E. maculatum Rev. on pear and quince =F. raaculata.
p. 149.
See
E. thumenii (Cke.) Sacc. occurs on hawthorn.
Leptostromataceae-Scolecosporae
Conidia usually hyaline, linear or
filiform,
(p.
528)
continuous or septate.
Key to Genera of
Leptostromataceae-Scolecosporae
Pycnidia astomous or opening variously Pycnidia with a round ostiole; conidia

catenulate
1.
Crandallia.
Pycnidia astomous or irregularly dehiscent
Pycnidia with radiate-fimbriate margin

2.
Actinothyrium.
Pycnidia not radiate-fimbriate

Pycnidia of two kinds, small simple,
and
large
and
loculate
3.
Brunchorstia,
p. 532,
Pycnidia of
one kind Conidia muticate

Pycnidia corrugate, not hairy; conidia not separating
4.
Melophia.
Chaetopeltis.
Giulia.
Pycnidia hairy; conidia separating into joints

5.
Conidia
ciliate-penicillate
at
6.
apex
Pycnidia elongate, longitudinally
cleft
7.
Leptostromella,
p. 533.
Brunchorstia Eriksson
singly, the larger as
Pycnidia erumpent, irregular in form, the smaller occurring chambers in a stroma, opening by an irregular
See
p. 151.
pore; conidia filiform, septate. A single species. B. destruens Erik, on pine=Cenangium abietis.
533
Leptostromella Saccardo
(p.
532)
Pycnidia at first covered, at maturity apparently superficial, depressed convex, subcarbonous, dark colored; conidia bacillar or filiform, continuous or septate.
of
About twenty species, chiefly no economic importance.

L. elasticae E.
&
E.
or less el-
Spots large,
liptic,
more
whitish,
sordid,
purplish
pj^
margined; pycnidia epiphyllous,

hysterioid,
364.-Lcptostro~
elastics.
0.5-0.7
mm.;
longi-
After Stone and Smith.
4-5
tudinally dehiscent; conidia oblong, hyaline, continuous, 12-15 x /x; conidiophores 12-15 x 3-4 fi; obtuse, subolivaceous.
The cause
of leaf spots of Ficus elastica.^^^
Excipulaceae
(p.
479)
late or hysterioid, at first
Pycnidia membranous to carbonous, black, cup-shaped, patelmore or less spherical but at length widely
open, erumpent or superficial, glabrous or hairy.
Key to
Sections of Excipulaceae
Conidia globose to fusoid, continuous
Hyaline Colored
Conidia 1-septate, hyaline Conidia 2 to several-septate
I.
Hyalosporae,
Phaeosporae.
p.
533.
II.
III.
Hyalodidymae,
p. 536.
Hyaline Colored
Conidia filiform or bacillar
IV. Hyalophragmiae. V. Phaeophragmiae.
VI. Scolecosporae, p. 536.
Excipulaceae-Hyalosporae
Conidia
hjâline,
continuous, globose to oblong.
534
Key to Genera of
Excipulaceae-Hyalosporae
Pycnidia pilose or setose Conidia muticate; pycnidia cupulate. Conidia ciliate; pycnidia cupulate
...
1.
Amerosporium,
p. 534.
Conidia
Conidia several-ciliate at apex 1-ciliate at each end

less
2.
3.
Polynema. Dinemasporium,
p. 535.
Pycnidia smooth or nearly so Pycnidia more or ciform

cup-shaped, or dis-
Pycnidia black
Pycnidia composed of conglutinate dark hyphse Pycnidia with cellular context Pycnidia cup-like when mature, sometimes obconoid
4.
Godroniella.
Conidiophores simple Pycnidia cup-shaped

Pycnidia terete-conic Conidiophores branched Pycnidia subglobose, disciform or verruciform
5. 6.
7.
Excipula. Catinula.
Heteropatella.
Pycnidia subglobose, irregularly

dehiscent and collabent ....
8.
Dothichiza,
p. 535.
Pycnidia disciform, often imperfect and covered by

epidermis
9.
Discula,
p. 535.
Pycnidia verruciform; mucose-involute

Pycnidia purple
conidia
10. 11.
Agyriellopsis.
Lemalis.
Psilospora.
Pycnidia hysterioid or valvately gaping Pycnidia widely hysterioid
12.
Pycnidia valvately gaping Conidiophores typically branched.

Conidiophores simple or none
..
13.
14.
Sporonema, p. Pleococcum.
535.
Amerosporium Spegazzini
Pj'cnidia subcupulate, setulose, conidia cylindric to ellipsoid.
Some twenty-five species, chiefly saprophytes. A. oeconomicum E. & T.
535
Spots orbicular, 2-6 mm., white above with a reddish border, mostly entirely reddish below; p3^cnidia epiphyllous, erumpent, conic-hemispheric, broadly perforate above, beset with straight, spreading, grayish-black, septate bristles, 100-150 x 4 ju; conidia
ol)long-fusoid, 18-27 x 4
/x.
\'ery
common on cowpea
leaves in circular spots, with dark
pycnidia in concentric circles
on white background.^^^
L^vielle (p. 534)
Dinemasporium
Pycnidia cupuliform, superficial, black, with conidia elongate or allantoid, with apical spines.
dark
bristles;
Some
thirty species, chiefly saprophytes.

is
D. oryzae Miy.
on
rice.^^
Dothichiza Libert
(p.
534)
disculate, irregularly
Pycnidia erumpent, roundish, somewhat
dehiscent; conidia elongate or cylindric. In part About eleven species, chiefly saprophytes.
=Cenangium.
D. populea
S.
&
B. parasitizes poplar. ^^
Discula Saccardo
(p.
534)
Pycnidia disciform-patellate, imperfectly differentiated the substratum; conidia ellipsoid, elongate or cylindric. Some twenty-five or thirty species, chiefly saprophytes. D. platani (Pk.) Sacc.=Gnomonia veneta. See p. 274.
from
Sporonema Desmazieres
closed, then
(p.
534)
Pycnidia subepidermal, erumpent, at first opening radiately; conidia ovate
or cylindric. ^' Some sixteen species, chiefly saprophytes. S. platani Baum on Platanus=Gnomonia Fig. 365. Sporonema oxycocpi on cranberry a o'7/ veneta. See p. 274. leaf. After Shear.
J.
S. phacidioides
folii.
Desm.=Pseudopeziza
tri-
See
p. 148.
S. oxycocci Sh."^ Pycnidia amphigenous, excipuliform, thickened at the base,
636
gradually disappearing above, immersed, erumpent, depressedglobose, gregarious or scattered, 50-100 ijl, sometimes collapsing rupturing irregularly by a slit or triangular split; conidia cylindric,
straight, 17-19
x 3-4 n; conidiophores simple, oblong to subglobose,
about 34 the length of the spore, or less. S. pulvinatum Sh. is also on cranberry.
On
cranberry.
Excipulacese-Hyalodidymae
(p.
533)
Conidia hyaline, 1-septate, oblong or fusoid.
Key to Genera
Pycnidia discoid or patellate
Pycnidia discoid, veiled;
simple
of Excipulaceae-Hyalodidymae
conidiophores
1.
Discella, p. 536.
Pycnidia patellate, subsuperficial; conidiophores branched

Pycnidia hysterioid or irregularly gaping
2.
Pseudopatella.
Pycnidia hysterioid, elongate Pycnidia globose, then irregularly gaping;

conidia catenulate
3.
Scaphidium.
Siropatella.
4.
Discella Berkley
& Broome
Pycnidia disco-patellate, imperfectly formed; conidia fusoid or oblong. Some twelve or fifteen species, chiefly saprophytes. D. cacaoicola A. & S. is on cacao in Africa.
The Excipulaceae-hyalophragmise, Excipulaceae-phaeophragmiaB contain no important parasites.

Excipulaceae-Scolecosporae
Conidia typically hyaline, bacillar or
septate.
(p.
533)
continuous or
filiform,
Key
to Genera of Excipulaceae-Scolecosporae
Pycnidia separate Conidia separating at the joints Conidia not separating Pycnidia discoid, covered, erumpent,
1.
Schizothyrella.
margin lacerate; conidia
filiform.
2.
Protostegia.

Pycnidla mostly cupulato, not lacerate;
conidia
537
hamate
3.
Oncospora.
Ephelis,
p. 537.
Pycnidia in a stroma Pycnidia pezizoid

4.
Pycnidia superficial
5.
Pseudocenangium.
Ephelis Fries
Stroma
in
black, subcffused, sclerotioid; pycnidia pezizoid, sunken the stroma; conidia cylindric to filiform.
species, chiefly of no economic importance. a conidial form of Balansia. See p. 209.
Some seven
It is
Melanconiales
(p.
479)
Mycelium internal; true pycnidia never developed, the conidiophores form a stratum; strata typically bearing conidia in acervuli which are immersed or erumpent, black or light colored, waxy,
corneous or even submembranous, accompanied by setae or not;
conidia variable.
The common name caused by a member of
"
anthracnose"
is
applied to any disease
this order.
A single family Melanconiaceae which contains about forty-five genera and over twelve hundred species.
Key to Sections of Melancomaceae
Conidia globose to elongate Conidia continuous
Hyaline Colored Conidia 1-scptate Hyaline Colored

Conidia 2 to
I.
II.
Hyalosporae, p. 538. Phaeosporae, p. 553.
III.
Hyalodidymae,
p. 555.
IV. Phaeodidymae, p. 556.

septate
many
Hyaline Colored
Conidia muriform, dark Conidia long-cylindric to filiform Conidia stellate
V. Hyalophragmiae,
VI. Phaeophragmiae, VII. Phaeodictyae.
p. 556. p. 557.
VIII. Scolecosporae, p. 561.
IX. Staurosporae.
538
Melanconiaceae-Hyalosporae
Conidia hyaline,
(p.
537)
1-celled, globose to oblong, rarely dilute colored.
Key
to Genera of Melanconiaceae-Hyalosporae
Conidia muticate
Masses, or acervuli, not setose Conidia not catenulate
Conidia not allantoid
Masses bright colored, subtremelloid

1.
Hainesia.
Masses
black, rarely bright colored, waxy or horny Growing, for the most part, on
gray
to
leaves or fruits
2.
Gloeosporium,
p. 539.
Growing usually on twigs

or shrubs
of trees
3.
Masses black,
Conidia in chains
discoid,
horny
4.
5.
Myxosporium, p. Melanostroma.
Nsemospora,
546.
Conidia allantoid
p. 547.
Masses
hard
oblong,
hysterioid,
dark,
6.
Hypodermium,
Myxosporella.
p. 547.
Masses discoid, pulvinate or conoid Masses bright colored, soft Masses dark to black
Conidiophores repeatedly branched
7.
Masses discoid; conidiophores dichotomous Masses depressed-pulvinate

;
8.
Blennoria.
conidiophores verticillate
9.
Agyriella.
Conidiophores simple
Masses
scutellate,
olive
or
10.
ashen
Myxormia.
Bloxamia.
Masses truncate, black below, pale above Masses setose at margin; conidiophores
short, fasciculate
11.
12. Colletotrichuni,p. 547.
Conidia aristate with a branched awn at
apex
13. Pestalozziella.
539
GlcBosporium and CoUetotrichum are prominent in pathology as the "anthracnosc fungi" and cause many important diseases. The two genera, separated only by the occurrence or non-occurrence of
setae,
contain
many
species
of these genera to the other
which have been transferred from one on this character, which is to some ex-
tent a variable one depending upon the supporting medium, conditions of growth and the particular strain of the fungus under observation.
Many form-species have been described solely on a basis of the hosts affected. Subsequent culture study, and cross inoculation has often failed to sustain these species so that many forms
that were formerly considered as distinct are now grouped under one name. No satisfactory disposition of these forms can be made
imtil their ascigerous stages are
known and compared and
their
biologic relations investigated.
Such work as has been done (see page 267) leads rather to consolidation than to segregation of species. For sake of clearness and convenience, mention is made below
of
many form
species of these
two genera under
their old names,
though the evidence now is that in consolidated with other species.

Gfoeosporium Desmazieres
many
instances they should be
&
Montaigne
(p.
538)
Conidial layer subepidermal, disciform or pulvinate, usually erumpent, pale or fuscous; conidia ovate, rarely oblong; conidio-
phores needle-shaped. In part=Glomerella,
Pseudopeziza, Gnomoniella,
Gnomonia,
Trochila, Physalospora, Calospora. There are over three hundred species of parasites,
many
of
them very important pathogens. The spores in germination commonly form dark colored, thick-walled chlamydospores and usuall}^ become 1-septate.
G. rufomaculans (Berk.) Thiim. on a large variety of hosts= Glomerella rufomaculans. See p. 264. G. melengonea E. & H. is reported on egg-plant fruits in New Jersey;^* G. orbiculare Berk, on cucurbs; G. fructigenum Berk, on many fruits.
G.
laeticolor
Berk, on peaches and apples and G. versicolor
540
P
S m
o
60 bO
->^
O " rt w sj; a
-^pq
..
w^
Ot-I
ftoT
4)
Q,
r*;;
3 3-?^
11
g-.
2 S S o "p^.a ft ^ Q iC? 03 ...

03
CD
<-*>;
THE FUNGI WHICH CAUSE PLAxNT DISEASE
541
& C. on peaches are probably rufomaculans.

B.
all
identical with Glomerella
G.
salicis West, on Salix=Pseudopeziza salicis. See p. 148. G. piperatum E. & E. on pepper=Glomerella piperata. See
p. 269.
G. cylindrospermum (Bon.) Sacc. on Alnus = Gnomoniella tubiSee

p. 274.
formis.
G. vanillae Cke. on orchids =Calospora vanillsc. See p. 280. G. macropus Sacc. on Cattleya=Physalospora cattleya. See
p. 253.
& C. on ore bids = Glomerella cincta. See p. 269. G. paradoxum (de Not.) Fcl. on Hedera=Trochila craterium. See p. 157. G, cingulatum Atk. on privet = Glomerella cingulata. See
G. cinctum B.
p. 268.
G. psidii Del. on guava=Glomerella psidii. See p. 270. G. atrocarpi Del. on Atrocarpus= Glomerella atrocarpi.
p. 273.
See
G. nervisequum
See
p. 274.
(Fcl.)
Sacc.
on sycamore =Gnomonia veneta.

ribis.
G. ribis (Lib.)
p. 148.
M. & D. on Ribes=Pseudopeziza
on grapes
in Australia.
See
G. bicolor
M.
Cal. occurs
G. ampelophagum (Pass.)
ters gray;
Sacc.^^^"^^^
Spots subcircular often confluent, from cortex of the berry, cenmargin dark or red. Acervuli subepidermal, minute, col-
lected; conidia oblong, ellipsoid or ovoid, 5-6 x 2-3 fx, hyaline. Small dark spots are produced on fruit, leaf or cane of grape.
These
later enlarge
and show white centers with dark or even red
The mycelium lies just below the epidermis. On shoots the cambium is killed and cankers develop. Two kinds of spores have been found by Viala and Pacottet^^^ one very small and linear,
borders.
the other larger and in Phoma-like pycnidia. eastern United States and Europe.
Common
in the
G. depressum Penz. is on Citrus. G. spegazzinii Sacc; and G. intermedium Sacc. grow on Citrus fruits; G. citri Mas., G. hendersonii B. & Br. on oranges causing
leaf scorch in
England, and Trinidad.
542

G.
G. variabile Laii. grows on Ribes alpinum. Oud. is described as a currant parasite. G. malicorticis Cor.^^
outline;
curvatum
Caulicolous; spots brownish, slightly depressed, irregular in acervuli minute, erumpent; conidia elliptic, curved,
or greenish-tinged, hyaline granular, 24 x 6 ju.
On
apples in northwestern
United States. been reported

connected.
^^^
Neofabrea has
as
genetically
re-
G.
cydoniae
Mont,
is
corded as a parasite on the

quince.
G. musarum C. & M. is a common wound parasite on

bananas.
Acervuli innate; erumpent, gregarious, rose-tinged; conidia
elongate-ellipsoid, ends
rounded, 10-12 x 4-5
ular.
fi,
gran-
variety, importatum,
Fig. 367.
malicorticis; a, acervulus; E, germinating spore. After Cordley.
G.
is
also recognized.
G. diospyri E.
Acervuli
&
E.
dis-
on yellowish
colored areas, innate, erumpent, epiphyllous, on the leaf veins, minute, tuberculiform, pale; conidia ovate, granular, 6-14 x 5-7 n.
On persimmon.
G. fragariae (Lib.) Mont.
Spots
indeterminate,
red,
epiphyllous;
acervuli
applanate,
rugulose, black; conidia cylindric, 4 to 5-guttulate. On strawberries but not usually troublesome.
G. amygdalinum
Brizi.^^^
occurs on the green fruit of the al-
mond
in Italy.
"'^^"^^^
G. venetum
Speg.^^'
Caulicolous or foliicolous; spots orbicular or elliptic, border raised, darker, 2-3 mm. in diameter; conidia oblong, elliptic, 5-7 x 3 Ai, in mass amber-colored.
543
The fungus occurs on all aerial parts of the raspberry and is wide-spread in Europe and America causing serious disease. On canes small purple spots first show near the ground, enlarge and soon develop ashen centers. The leaf spots are small, often
scarcely
1
mm.
in diameter.
^^^'
^'^'^
G. mangif erae Hen. is found on mango leaves in Cuba and other West Indian Islands. G. olivarum d'Alm. parasitizes olive fruit in Europe. G. minus Sh. is on cranberry. ^^^ G. myrtilli All. is injurious to Vaccinium myrtillus. G. cofifeanum Del. occurs on coffee in Java; G. pestis Mass. on yam leaves in Fiji. G. trifolii Pk.iî"i^2
Spots subcuticular, brown, suborbicular, concentrically zonate; conidia oblong to cylindric, obtuse, 15-23 x 4-6.3 n.
as the
The fungus was first observed in America and what was regarded same was later seen in Europe as the cause of dying of stems
of clover.
and leaves
G. caulivorum Kirch. ^^^

Caulicolous, spots forming long dark streaks, more or less sunken, blackish-bordered; acervuli minute; conidia curved, more or less
pointed, 12-22 x 3-5 n. This was said by Kirchner (see ^^^) to be the cause of the more serious European anthracnose affecting stem, fruit and leaf of
clover.
Fulton
^^"^
in 1910 reported
it
in
America and showed that

in
pure cultures of the fungus readily produced infection in
even on un wounded succulent parts when have been known to live twelve months. G. morianum Sacc. is on alfalfa. G. medicaginis E. & K.
wounds or humid air. The conidia
Acervuli scattered, innate, blackish, rather large, visible on both sides the leaf, opening below; conidia oblong, cylindric, granular,
subhyaline, more or less narrowed at the middle, 15-20 x 3-4 fx. On withered leaves and stems of alfalfa, defoliating the lower part of the stem.
G. manihotis Hen. is found on Cassava in Africa. G. lagenarium (Pass.) Sacc. on cucumbers is probably identical with Colletotrichum lindemuthianum.
544
G. concentricum Grev. causes spotting of leaves of cabbage,

cauliflower, etc.
G. cattleyae (P. & D.) Sacc. grows on leaves of Cattleya; G. dianthi Cke. on carnation in England. G. clematidis Sor. is found on cultivated clematis stems ;^^'^ G. rosae Hals.-^ is reported by Halsted on rose canes causing
injury similar to that of G. venetum on the raspberry.
G. mezerei Cke.
is
on Daphne.
G. oncidii
G. affine Sacc. grows on various orchids and Hoya. Oud. on Oncidium.
G. euphorbiae Hals, is on clusters and stems of spurge.'^ G. stanhopeicola Hen., G. laeliae Hen. and G. pallidum Karst. & Har. are on orchids; G. helicis (Desm.) Oud. on English ivy. G. cactorum Ston. occurs on a number of species of cacti; G. beyrodtii Klitz on Vanda; G. opuntiae E. & E. on Opuntia; G, elasticae C. & M. on Ficus; G. bruneum Fetch, and G. alborubrum Fetch, on Hevea. G. rhodendendri Br. & Cav. forms yellow spots on various species of Rhododendron. G. aquilegiae Thiim is on Aquilegia. G. violae B. & Br.^^
Spots pale, becoming whitish; acervuli very thin, solitary; conidia yellowish. Causing leaf spots on cultivated violets. G. cytisi B. & Br. is on laburnum;
G. G. G. G.
bidgoodii Cke. is on Oncidium; pelargonii C. & M. on Felargonium.

crotonis Del. occurs on Codiseum.
soraurianum
All. also
on Codiaum leaves and described as
a dangerous parasite is perhaps identical with G. crotonis. G. anthuriophilum Trinch. causes spots on Anthurium leaves.
G. begoniae Magnag. is on begonias in Italy. G. fagicolum. Fass. is widespread and destructive to beech
in
Germany.
G. umbrinellum B. & Br. is found on Quercus; G. inconspicuum Cav. on elm; G. theae-sinensis Miy. and G. theae Zimm. on tea
in
Japan and
Africa.

G.
Palms.
allescheri
545
Bres.
and
G.
nanoti
P.
&
D.
occur
on
G. tiliae Oud. is a serious parasite on the twigs, petioles and leaves on the linden in Denmark. G. tiliaceum (AU.)^''^ said to be distinct from the above occurs in Germany on Tilia. G. juglandis (Lib.) Mont, causes a common, and serious leaf
blight of the butternut.
^^^
G. fagi (D.
Spots
&
R.)
West
is
on Fagus;
above, olivaceous, vitreous beneath; acervuli small, prominent, honey-colored; conidia oblong minutely 1 to 3-guttulate; conidiophores ovate, 15-20 x 7-8
subcircular,
fuscous
ij.,
fasciculate, cylinclric, fuscous.
and
G. apocryptum E. ^^ of box elder.
&
E. causes a nursery disease of maples
Acervuli numerous, minute, mostly hypophyllous, on dead areas

of the leaf; conidia very variable in size, to narrowly elliptic.
5-12 x 23^-5
ju,
oblong
G. betularum E.
& M.
Spots rounded, 2-3 mm., blackish margined; acervuli amphigenous, brownish, 120-140 n, becoming cupulate; conidia hyaline, obovate, 9-10 x 5-G
fx,.
It is
common on leaves of American birches. Other common species on deciduous trees are
G. G. G. G. G. G. G. G.
tremulae (Lib.) Pass, on Populus;
betulinum West, on beech; alneum West, on alder; carpini (Lib.) Desm. on Carpinus; coryli (Desm) Sacc. on Corylus; quercinum West, on oak; nervicolum Massal on oak. kawakamii Miy. is found on Paulownia
in
Japan causing
witches' brooms.
G. saccharini E.
&
E.
ju,
Acervuli minute, numerous; spores oblong-fusoid, 6-7 x 1.5-3 hyaline, continuous. On maple.
G. caryae E.
Spots
&
D. =Gnomonia
1-2
caryse.'*'*^
suborbicular,
cm.,
margin
subdefinite;
acervuli
546
hypophyllous, numerous, 75-150 n; spores allantoid, continuous, 7-10 X 1.5-2 On Carya. G. berberidis Cke.
fjL.
Hypophyllous;
5 X 3
/x-
acervuli
collected,
numerous; spores
ovoid,
Barberry. G. tamarindi Hem. is on tamarinds, in Africa. G. canadense E. & E.
On
Spots amphigenous, center pale, border brownish; acervuli few.
Fig. 368.
M. corticolum, on apple twig.

Edgerton.
After
180-200 ju; spores ovate-oblong, hyaline, 10-14 x 3.5-4.5. On white oak.
Myxosporium Link
conidia ovate,
(p.
538)
Acervuli immersed or superficial, indefinite, pallid or reddish;

hyaline or pale,
species,
conidiophores slender-cylindric.
Some seventy
some
See
All.
of
which are important pathogens.
In part=Gnomonia.
p. 274.
M. valsoideum
See
p. 274.
(Sacc.)
on sycamore = Gnomon ia veneta.
M.
corticolum Edg.
Acervuli erumpent, originating under several layers of cortex, 1-2 mm. in diameter, scattered over the diseased area; conidia
straight or curved, cylindric, very densely granular, 18-36 x 6-9
H,
oozing out of the pores in white
cirri;
conidiophores very
short.
Very similar to an immature Sphseropsis malorum but considered distinct by Stewart and his associates.^* It forms bark cankers in pear and apple in America.
547
M. loiigisporum Edg.'"' Acervuli erumpent, subcorticular, variable in size up to 1.5 mm., scattered over the host in poorly defined rows; conidia straight
or curved, 30-48 x 12-15 phores very short.
/x,
oozing out in white
cirri;
conidio-
On
twigs of Liriodendron.
Other parasitic species are: M. piri Fcl. on pear; M. mali Bres. on apple; M. abietinum Rost. on conifers; M. devastans Rost. on beech; M. lanceola S. & R. on oak; M. carneum Lib. on beech twigs.
Naemaspora Persoon
Acervuli
subgelatinous,
indefinite,
(p.
538)
colored;
bright
conidia
allantoid, short, with a bristle at each end. N. crocea (Bon.) Sacc. is reported by Massee as the cause of
die-back of peach shoots in England, ^^^
Hypodermium Link
oblong, catenulate.
(p.
538)
Acervuli subcuticular, erumpent, elongate, black; conidia ovate-
H. orchidearum Cke.
is
on Cymbidium.
Colletotrichuni Corda (p. 538)
with long black

of setse, a
Acervuli innate erumpent, discoid or elongate, dark, surrounded setae; conidia terete to fusoid; conidiophores short.
The genus is distinguished from Gla?osporium by the presence somewhat unreliable character. See p. 539.
In part=Glomerella and Pseudopeziza.
See pp. 264, 147.
Some eighty
species, several of
them very important plant path-
ogens. C. gossypii Sout. on cotton =Glomerella gossypii. See p. 271. C. cincta Ston. on orchids=Glomerella cincta. See p. 269.
C. nibicolum E.
&
E. on red raspberry=Glomerella rubicolor.
See
p. 270. C. lindemuthianum (Sacc.
& Magnus)
Briosi
&
Cavara.^^'
^^'
164-167
548
scattered, surrounded
Spots subelliptic to irregular, depressed, brownish; acervuli by a few not very conspicuous black seise;
conidia oblong, ends rounded, straight or curved, 15-19 x 3.5-5.5

n; conidiophores cylindric, simple,
45-55
fx.
This fungus, generally known as C. lindemuthianum,

inmracnose CanKfr
i
is
accord-
SeedCoat
Starcll
Grains
C. lindemuthianum. Showing relation of -the fungus Fig. 369. to the tissues of the bean. To the left above is a diagram of a section across a bean pod through a canker. The drawing below is a much enlarged largely diagrammatic view of a portion of this same section. It shows how the mycelial threads of the fungus may penetrate the seed coat and enter the starchy tissue of the seed, there to remain dormant until the following season. On the left is a spore germinating and penetrating the epidermis. To the right a magnified view of spores, one germinated. After Whetzel.
ing to the cultural studies of Shear & a variety of Glomerella rufomaculans.
Wood
See
(see p. 267),
probably
p. 264.
On
the bean
it
attacks stems, leaves, cotyledons, or the pods,
producing sunken spots of dead tissue which bear the numerous pink acervuli. It has been shown that the mycelium on the fruit may penetrate through the pericarp and into the seeds beneath
and there hibernate.

C. lindemuthianum.
leaves C. lagenarium (Pers.) E. & H. ^^^ is probably identical with It is described as the cause of spots on fruit
and stems
of cucumbers, watermelons, squash,
pumpkins
^^^
and
citron.
all parts.
C. oligochaetum Cav. grows on cucurbs, attacking
Probably =C. lindemuthianum.

C. carica S.
549
&
/x,
H.^-"'
'^^
becoming black, hemispherical, numerous, bearing 1-12 (or often 0) long, slender, irregular setae which are dark throughout, acute, rigid, septate, 2-6 x 22brown,
small, 85-250
Acervuli
conidia regular, oblong, obtuse, 3.5-6.6 x 8.7-20; conidiophores slender, 1-2 x 45 /x, hyaline. It is the cause of a decay of figs in the United States.
n,
106
C.
ampelinum Cav.
is
on grape
leaves.
C. gloeosporioides Penz. Acervuli sparse or scarcely gregarious, subepidermal, erumpent, dark, cylindric, setae continuous or few-septate, dark colored,
40-90 X 5-6
fjL,
conidia cylindric, straight, 16-28 x 4-6 n; co-
Fl(j.
370.
curica.
;"),
((jiiidiosiiDics,
produced on germ tubes.
spores ;ind s(.'t;i', 4, black bodies After Stevens & Hall.
nidiophores
densely
fasciculate,
cylindric,
rounded
apically,
tenuous, 18-25 x 4-5 n. It causes "wither tip" of orange, pomelo and lemon,^^*^^^ spots on citrus leaves, lime canker, and anthracnose of stem and flower with great financial loss in Florida, West Indies, South America,
Australia,
Malta and many other
localities.
In "wither tip" the fungus enters through the terminal bud or from leaves. On lemons attack is through bruises. Acervuli are
found on
leaf, twig or fruit, breaking through the epidermis. Cross inoculation on the various hosts and with the different forms
of the disease
demonstrated the identity of all. Smith ^^^ has questioned the casual relation of this fungus as regards "wither-
tip" as it occurs in California. C. falcatum Went.

Acervuli poorly defined, setae irregularly arranged, cuspidate,
550

^,
fx; conidiophores hyaline to fuscous. This is believed to be the chief cause of the red rot of sugar^^^ It was reported in the United States by Edgerton,^^^ cane.
100-200 X 4
brownish; coniclia falcate, 25 x 4
ovoid, 20 X 8
n.,
^'^ Inoculation experiments indicate that it by Stevens. distinct from C. lineola, on sorghum and Johnson grass, which
also
is it
resembles morphologically. C. cereale Manns. ^"
setse
Spots circular to ovoid, 30 mm.; acervuli dark brown, or black; few or many, dark brown to black, at base 6-8 m thick, continuous or 1 to 2-septate; tapering to a length of 60-120
/jl,
Fici.
371.
C. cereale, acervulus showing mycelium,

iospores and spores.
setse,
conid-
After Manns.
conidia 18-26
x 3-4
n,
spindle to boat-shaped,
2 to several-
guttulate; conidiophores, very short, 12-6 x 1-2 mThis fungus is parasitic on the roots, stems, blades and spikes of rye, wheat, oats, barley, emmer, orchard grass, timothy, blue
grass
and
chess.
The
shrivelling of the grain.
disease causes a premature ripening and Superficially the diseased heads present
the same appearance as those attached by scab (Fusariose) but no pink over-growth is present, nor is the presence of the disease
always apparent, as it was found on numerous heads of grain which appeared to be healthy. IVIorphological studies indicate that all the hosts mentioned above are affected by the same fungus. This was confirmed by cross inoculations in the case of wheat and
emmer.

C.
trifolii Ikiin.''*"'^"
551
Spots dark, depressed; acervuli erumpent, scattered or gregarious; conidia straight, ends rounded, 3-4 x 11-13 /x; conidiophores cylindric or fusoid, hyaline; setae few or many, continuous or uniseptate, dark, paler apically, 4-7 x 39-62 n, sinuous, or
nodose.
It occurs as
an anthracnose producer on stems, rarely on

alfalfa causing
leaves,
of clover
and
it is
pearance
much
like
very serious injury. In general apGloeosporium caulivorum.
C. spinaciae E.
&
H.-^
Spots roundish, dirty-white, or greenish, 2-4 mm., with a slightly raised border; acervuli amphigenous, punctiform, 40-75 (x, with 3-12 erect or spreading bristle-like setse, 60-75 x 4-4.5 n, sub-
bulbous at base, subhyaline, subacute above, dark brown below; conidia subfalcate, fusoid, 2 to 4-guttulate, 14-20 x 2.5-3 n, ends
subacute; conidiophores short.
It
produces blotches on spinach leaves.
C.
phomoides
(Sacc.) Ches.^^'
'^' '''-^'''' ^^^
Spots depressed, circular, slightly discolored, center black, 5-10 ram., later irregular and confluent; acervuli abundant, densely gregarious, rusty brown to black, applanate, 95-150 m; seta) abundant, fuliginous, generally curved, septate, 65-112 /x; conidia oblong, 16-24 x 4 ends subacute; conidiophores short, slender, 30-40 m high, arising from a well developed stroma. On tomato. This is a common cause of ripe rot of tomatoes. The fungus was
fj.,
studied in culture by
different
Stoneman
^^
who
reported
it
as
somewhat
from G. rufomaculans, though Edgerton ^^ thought the ^^^ throws apple and tomato forms the same. Work by Gueguin doubt on the American form on tomato being identical with the Chester ^'^^' **- has European form known as G. phomoides. reported what he regarded as the last species as setigerous.
C. nigrum, E.
&
H.^^'
^^
Spots blackish, depressed; decaying; acervuli numerous, superficial; setse numerous, slender, setse pointed; conidia oblong.
This form which appears quite different from G. piperitum was

described from
New
Jersey by Halsted.
552
C.

malvarum
Br.
&
Casp.^^^
(=C.
althaea.)
Epiphyllous and caulicolous; spots brown, sunken; acervuli erumpent; setae dark brown, abundant, 1 or 2-septate, usually colorless below, 60-109 x 3-5 n, appearing after the conidiophores which are colorless, cylindric, tapering slightly and apically rounded, slightly longer than the conidia; conidia irregular, oblong,
granular, flesh-colored in mass, 1 1-28 x o fx. ^^^ It is described by Southworth as the cause of anthracnose of
the hollyhock. The fungus closely resembles C. lindemuthianum but cross inoculations on the bean failed to produce disease though
on hollyhock inoculations succeeded

C.
easily.
schizanthi Jensen
&
Stewart was found on greenhouse
Schizanthus plants in Ithaca.^^^

C. agaves Sacc.
Spots pale; acervuli conic; setse few, 90-100 x 5-6 m, brownish ochraceous, 2 to 3-septate, conidia 22-26 x 4-5 ju; conidiophore '^^ subramose, fuscous at base. On sisil hemp and agave.
C. bletiae Hals,
is
on
Bletia.
C. violae-tricoloris R. G. Sm.^^^^^^
more or
definite,
Spots pale-yellow on leaves. Dead areas on petals occur with less deformity of blossom. Spots at first orbicular and
later
confluent and irregular, acervuli numerous,
50-
often confluent; stroma usually poorly developed; setse fx, mostly single or in pairs, 20-70 ju, deep brown, 1 to 2-septate, tapering gradually to a point; conidia oblong or slightl}^ curved,
150
ends blunt, 20 x 5 n; conidiophores short, hyaline. It causes spotting of pansy leaves in several states and leads
to failure to bloom.
C. anthurii Del. occurs on Anthurium; C. luxificum H. & D. on cacao in the West Indies; C. elasticae (C. & M.) Koo. on Ficus. C. primulae Hals.-^
is
reported as causing a leaf disease of the
primrose. C. kentiae Hals,

C.
is
on palms.^^
ju,
omnivorum
Hals.
falcate;
Spots dry, irregular in outline; conidia 20-28 x 3-5

setse elongate, acute, black.
On
Aspedistra and other plants.-^

C. camelliae ]\Ias.
is
553
on
tea.
C. antirrhini Stew.'^^
3-10 mm, stroma well developed; conidia numerous, crowded; 16-21 X 4 /i, straight or curved, ends rounded; conidiophores short; setse abundant, dark brown, 50-100 fx, simple, mostly straight and tapering uniformly to a subacute point. It attacks the stems and leaves of the cultivated snapdragon producing sunken spots. C. cyclamenae Hals, is on Cyclamen; C. dracaenae Hals, on Dracaena; C. coffeanum N. is on coffee; ^^^ C. heveas Fetch, on Hevea in India; C. cradwickii Banc, on cacao pods; " C. brachytrichum Del. on cacao leaves; C. theobromae A. & S. and C. theobromicolum Del. are on
Spots depressed,
elliptic or orbicular, often confluent,
acervuli
cacao.
C. incarnatum
Zimm.
is
on Hevea and vanilla;
C.
macrosporum
Sacc. on vanilla;
C. poUaccii ]Ma;ynag. on Japanese loquot in Italy; C. hedericola Lau. on Hedera.
Undetermined species of Colletotrichum have been reported on asparagus. ^^ carnation, ^^ pear,^^ and many other hosts.
Melanconiaceae-Phaeosporae
(p.
537)
Conidia dark, continuous, globose to oblong or fusoid.
EIey to
Genera of Melanconiaceae-Phaeosporae
Conidia solitary on the conidiophores Conidia globose or oblong Conidia fusoid, often arcuate
Conidiophores not swollen at base. Conidioj)horcs swollen at base
...
1.
Melanconium,
Cryptomela. Basiascum.
p. 554.
2.
3.
Conidia in chains
Conidial chains separate Conidial chains in a mucose head
4.
5.
TruUula, p. 554. Thyrsidium.
554
Melanconium Link
(p.
553)
Acervuli subcutaneous, conic or discoid, black; conidia elongate to globose-oblong, erumpent in black masses; conidiophore simple.
In part=Ti'ichosphaeria.
See
p. 228.
More than one hundred
fifty species.
M.
fuligineum
S.
&
V. Cav.^''
first
Acervuli scattered or gregarious, at
gray-cinereous, then
brownish, subepidermal, erumpent; conidia ovoid to ellipsoid,

inequilateral, acute,
filiform,
It
9-12 x 4-6 n, olive, guttulate; conidiophores from a well developed stroma. causes the important bitter rot of ripening grapes, especially the white varieties, occurring also on shoots and peduncles. Acervuli appear on the
surface of the rotted berries.
The myce^^^
lium penetrates even to the seeds. Southworth suggested in 1891

this
that
and Gloeosporium
^^'^
are congeneric but

generically distinct. what he regarded as
^t
,
r*^^^^^^*^
Atkinson
culture
who
studied the fungus in pure
i^^^d^^^^
Fig.
thinks
^^^
them
372. m. fuligineum.
After Atkinson.
Noack
.,
,
the typical
-itvtpi-M.
wlio studied
fuligineum says that
-^
it
is
preceded by and belongs to a Gloeosporium-form. M. pandani Lev. is a common parasite on Pandanus in greenhouses, killing the branches. M. sacchari Mass.^-^
Acervuli numerous, collected in indeterminate pallid orbicular spots; conidia cylindric, 10-15 x 3-4 /x, straight or sHghtly curved,
olivaceous, smooth.
In leaves, sheaths and culms of sugar cane.

rind disease.
The cause
of the
TruUula Cesati
(p.
553)
depressed,
Acervuli subcuticular, erumpent, discoid-pulvinate, or conicalblack; conidia oblong-cylindric, catenulate; conidiospecies, of trifling
is
phores long, bacillar, simple or branched.
Some twenty
T. vanillae Hen.
on vanilla
in Africa.
THE FUNGI
WPIICII
CAUSE PLANT DISEASE
555
Melanconiaceae-Hyalodidymae
Conidia
hyaliiie, 1-septate,
(p,
537)
ovoid to fusoid.
Key to Genera of Melanconiaceae-Hyalodidymae

Conidia inutic
Saprophytic, on stems and fruits
1.
Biogenous, typically on leaves Conidia 3 to 4-ciliate at each end
2.
3.
Septomyxa. Marssonia,
p.
555.
Gloesporiella.
Marssonia Fisch
Acervuli globose-discoid, pale, conidia ovate to elongate.
part In
=Gnomonia, Trochila, Pseudopeziza. Some seventy-five species, all leaf parasites,
several of eco-
nomic importance.
M.
See
castagnei (D.
&
M.) Sace. on Populus = Trochila popularum.

tt
p. 157.
juglandis (T.ib.) Sacc. on walnut leptostyla. See p. 275.
M.
=Gnomonia
O^ MM.
r\(]
M.
populi (Lib.) Sacc.
W^ Jf
.
(J
fluent,
'
Spots suborbicular, epiphyllous, separate or conbrown, darker margined, acervuli convex ^ Fig.
'
. .
.
ti
^..3.
.,_.,
M.per,,
to applanate, fulvous; conidia obovate to subpyriform, 20 x 12 n, constricted at the septum,

straight or curved.
It
is
forans,
spore^sf
conidio^
A ft"
Selby.
common on
leaves of Populus as the cause of blighting

INIag. is
of lateral twigs.
It is injurious in nurseries.-^
M. M. M. M.
panattoniana (Bed.)
secalis (Oiid.)
found on lettuce in Italy;
Mag. on
rye;
martini
E. on oak; potentillae (Desm.) Fisch as the cause of a disease of the

S.
fall
&
strawberry in Europe.
M. M.
rosae Trail causes premature
of rose leaves.
perforans, E.
&
E.^-^
Spots small; irregular, 1-2 mm., pale, soon deciduous; acervuli
556

100-120
/x,
or
by confluence
larger;
conidia abundant, clavate or wedgeshaped, 11-15 X 2.5-3 n.
The cause
of
leaf
perforations of
X^
f] I
rs
lettuce.
M.
violae (Pass.) Sacc.^^^
.^ "
^ Fig.
374.
y
^~^y)
Discoloration
acervuli
of
the
host
slight;
conidiophores. After Jones and Giddings.
Spores and
M ar sk o nia
violte.
numerous, scattered, small; conidia curved, or straight, 15-18 x 5-6 At, septum usually excentric. Small spots are produced on violet ^
leaveS.
jyj^
medicaginis Vors.
(p.
is
OH
alfalfa.
Melanconiaceae-Phaeodidymae
Conidia dark, 1-septate, ovoid to fusoid.
537)
Key to Genera
Conidia solitary Conidia muticate Conidia
isthmi
1
of Melanconiaceae-Phaeodidymae
1.
Didymosporium,
Neobarclaya.
BuUaria.
p. 556.
to 3-ciliate at apex
2.
Conidia catenulate, connected by hyaline

3.
Didymosporium Nees
Acervuli rounded or elongate, covered, erumpent; conidia elongate or fusoid. A genus of less than twenty species. D. salicinum Vuill is on poplar.
Melanconiaceae-Hyalophragmiae
(p.
537)
Conidia hyaline, 2 to several-septate, oblong to fusoid or clavate.
Key to Genera of Melanconiaceae-Hyalophragmiae

Conidia separate Conidia muticate
Conidia oblong or fusoid, masses usually

pale
1.
Septogloeum,
p.
557.

Conidia long-clavatc masses dark. ... 1 to scveral-ciliate, usually at the
;
557
2.
Rhopalidium.
Pestalozzina.
Conidia
apex Conidia united at base into a radiate or

late
3.
stel4.
group
Prosthemiella.
Septogloeum Saccardo
(p.
556)
Acervuli very small, subepidermal, erumpent, pallid; conidia oblong. Some twentyfive species of leaf parasites.
S.
hartigianum Sacc. attacks maple, killing
very young twigs. S. ulmi Fr. is found on elm leaves =Phyllachora ulmi.
S.
(?)
mori Lev. on mulberry = My cosphserella

cydoniae
mori. (?)
S.
(Mont.)
Pegl.
is
on
the
quince;
S. manihotis
Zimm. on manihot
is
in Java.
'675.-
S. arachidis Rac.
seriously injurious to Fig.

is
Septoacerinum,
the peanut in Java.^^^ S. profusum E. & E.

S. fraxini Hark,
gloeum
found on Corylus;
on
ash.
conidiosporea and conidia. After Saccardo.
Melanconiaceae-Phaeophragmiae
(p.
537)
Conidia dark, 2 to several-septate, oblong to cylindric.
Key to Genera
Conidia mutic
of Melanconiaceae-phaeophragmiaB.
Conidia separate, not in chains Conidia oblong or elongate, not stellate Conidia curved-attenuate,
line-rostrate
i.
e.,
hya-
Conidia dark, except the hyaline beak
1.
Scolecosporium.
Conidia with 2 inner

others clear
cells
opaque,
2.
Toxosporium,
p. 558.
558

Conidia oblong, not rostrate Conidia cirrhose protruded. ......
3. 4.
Stilbospora.
Conidia not protruded Conidia stellate-lobed, lobes severalseptate
Coryneum,
p. 560.
5.
Asterosporium
Seiridium.
Seiridiella.
Conidia in chains
Conidia connected with filiform isthmi Conidia chains without isthrai

Conidia
ciliate ciliate at
6. 7.
Conidia
apex alone
8.
9.
Conidia
1-ciliate
Monochaetia,
Pestalozzia,
p. 558.
Conidia several-ciliate
p. 558.
Conidia
1-ciliate at
each end
10.
Hyaloceras.
Toxosporium Vuillemin
(p.
557)
Acervuli sublenticular, erumpent, scattered, minute, black; conidia curved, beaked at each end, central cells dark, apical hyaline;
conidiophores short, simple. T. abietinum Vuill.^^^ causes drying out of Abies leaves in
Europe.
Monochaetia Saccardo
As
seta.
in Pestalozzia except that the conidia bear only a single
About
sixty species,
leaf spots
is common on Castanea, causing large, with the acervuli showing in somewhat
M. pachyspora Bubak.
circular,
dead
concentric circles.
Pestalozzia de Notaris
Acervuli subcutaneous, erumpent, discoid or pulvinate, black; cells hyaline, with several
conidia elongate, colored or the end hyaline setse on each end.
genus of over two hundred species of various habit, some of
considerable economic importance. P. hartigii Tub. causes disease of tree and shrub seedlings in nurseries, constricting the stem just above the soil and resulting
in death.

P. funerea Desm.
559
Acervuli scattered, punctiform, blackish, subepidermal, arum'
stroma depressed, white; pent; conidia oblong, 5-cellecl, fusoid, constricted at the septa, the three
central
cells
fuscous,
the
others
hyaline, 22-32 x 6-8 m, with 2-5 recurved hyaline spines, 10-15 x 0.7-1 n; conidiophores short, 5-9 x
1-1.5
M-
found on various conifers Fig. 376. p. funeiea. A.. Spores. is a common B. A young mycelium. C. Hypha of an older mycelium. After Reed. saprophyte. In America it causes a stem spot or anthracnose of ginseng,^"* girdling the petioles. The culture characters were studied by Reed.^^^
It is
causing disease and
P. guepini Desm. var. vaccinii Sh.^^^ Acervuli minute, punctiform, convex,
black,
subepidermal,
el-
erumpent; conidia
liptic
and somewhat
about unequilateral, 20 fi long; central cells

dark, the two end cells hyaline, the apical cell
with 3-4 filiform
setse
22-35 /i long, the basal with a short hyaline appendage, 6-12 ju. Common on fallen
and
leaves of cranberries, associated with

It
rot of the berries.

Fig. 377.
Pestalozzia guepini vaccinii; a, a conidium having an apical appendage with three branches; causing a serious disa conidium having an apical appendage with b, four branches; c, a germinating conidium; d, a ger- ease, also on Camellia, minating conidium sending out two germ tubes.
is
common
on
tea
After Shear.
Magnolia,
Citrus,
Rhododendron.
P. uvicola Speg.^^^
Acervuli globose,
lenticular,
black,
subepidermal, erumpent,
560

fx;
300-400
conidia fusiform, 5-celled, the three median olivaceousfi,
fuscous, the others hyaUne, 35 x 8-10 X 1 ju, superior group 8-10 x I n.

It is described as the cause of rot of
inferior
appendage 25-30
grape berries and of a leaf
spot of the vine. P. aloea Trinch. occurs on aloe in Italy; P. clusiae Griff. & Mont, on Clusia leaves in France; P. richardiae Hals, sometimes disfigures calla leaves.
P. tumefaciens Hen. is found on Abies causing galls. P. gongrogena Temme causes galls and cankers on willow; P. fuscescens Sor. parasitizes cultivated Corypha; P. fuscescens var. sacchari
Wak.
S.
is
on sugar
S.,
cane.^'
On palms
occur P. palmicola P. phoenicis Grev.
&
P.
palmarum
Cke., and
P. inquinans C. & Hark is on eucalyptus in California; P. stictica B. & C. on sycamore and linden; P. suffocata E. & E. and P. discosioides E. & E. on roses.
P. lupini Sor. on Lupinus cotjdedons.-^"
Coryneum Nees.
(p.
558)
Acervuli discoid or pulvinate, subcutaneous, erumpent, black,
compact; conidia oblong to fusoid.
Some
C.
seventy-five species.
modonia (Sacc.) Griff. modonia Tul. See p. 281.
& Maub.
on chestnut =Melanconis
C. juniperinum E. on juniper =Exosporium juniperinum.^^ C. beyerinckii Oud.
Acervuli minute, punctiform, black, gregarious; conidia borne
on a parenchymatous pulvinate stroma,

septa or not,
stipitate, oblong, ovate,
light olivaceous, 3 to many-septate, slightly constricted at the

cells not all of the same size. Ascospora beyerinckii is said by Vuillemin to be the ascigerous See p. 236. It is reported as injurious to peaches, causstage.
ing blight
and shot-hole
in Africa.
in California
It is close to
^"
and peaches
and to apricots, cherries and perhaps identical with
Clasterosporium carpophilum. The acervuli occur as black specks near the centers of the leaf
561
spots but since diseased tissue soon drops out they are often difficult to find. Bark spots are often sterile. Late in the season spores
abound scattered on the surfaces

places, as near leaf scars.
of twigs, especially at
rough
with a mycelium and

twig and produce dead
kill
Spores near bud scales penetrate them the buds or if on bark they enter the
spots.
C. foliicolum Fuckel.-o^
erumpent; conidia ellipsoid-oblong, 17 x 6-7
Spots epiphyllous, ochraceous, indefinite; acervuli punctiform, n, 3-septate, con-
FiG. 378.
C. beijerinckii.
Longitudinal section of a pustule. After Smith.
stricted
at
the septa,
olivaceous,
ju-
lower
cell
subhyaline, stalk
subhyaline, 15-20 x 1.25
It is present as a saprophyte on apple leaf spots and causes cankers on twigs and branches. C. camelliae Mas. is reported on cultivated Camellia.
C. mori
Namura
causes mulberry twig blight in India.^"*
The Melanconiaceae-Hyalodictyae and Melanconiaceae-Phaeodictyae containing
only three genera have no parasitic species of
importance.
Melanconiaceae-Scolecosporse
(p.
537)
Conidia cylindric, filiform or suballantoid, hyaline, mostly continuous.
562

Key
to Genera of Melanconiaceae-Scolecosporse
1.
Conidia allantoid Conidia bacillar to filiform Conidia fasciculate at the apex of the
conidiophores Conidia solitary
Naemospora.
2.
Trichodytes.
Masses white or pale, foliicolous conidia

;
filiform
3.
Cylindrosporium, p. 562.
Masses gray or dark, usually ramicole;

conidia falcate
4.
Cryptosporium,
p. 564.
Masses
bright-colored, conidia falcate
saprophytic
5.
Libertella, p. 564.
Cylindrosporium Unger
Acervuli subepidermal, white or pallid, disciform or subeffuse; conidia filiform, continuous, hyaline, straight or curved.
parasites,
About one hundred species of several of them of con-
siderable economic importance. C. mori Berl. on Morus=Mycosphserella morifolia.
See
p. 249.
C.
castanicolum
See
p. 249.
(Desm.)
Berl.
on Castanea=Mycosphserella maculiformis.
C. padi Karst.'^--io
C. padi, section of acer FiG. 379. vulus. After Pammel.
Hypophyllous; spots angular, subfuscous; acervuli subepidermal causconidia elevations; curved, ing
cylindric,
48-60 x
ju;
conidio-
phores minute, produced in great abundance. This is the cause of the most common, familiar, widespread and destructive shot-hole disease of the cherry and plum. When on
The the peduncles the fruits are dwarfed and ripen unevenly. diseased leaf tissue usually falls away, resulting in "shot-holes."
Acervuli abound.
Arthur found a
asci
Phoma
in
the same pycnidia.
associated with this fungus and later He believed these forms all con-

nec.'ted
563
duced.
C.
but confirmation of such conclu.sion has not been adConnection with Coccomyces has also been reported. ""^^ C. tubeufiana All., also on Prunus is closely like C. padi.
pomi
Brooks.211-212
Acervuli pallid, subeffuse, at first subepidermal, then erumpent; conidia granular, filiform, straight or flexuose, 15-80 x 2-2.5 n.
It
is
reported
of
as
common from
and Michivarieties,
all
New Hampshire
gan on apples
to Virginia
almost
causing small, dry, sunken, brown fruit spots; a disease which had hitherto
been referred to a variety of causes.
The fungus was studied

culture
After artififial cultures. Brooks. proved by inoculations. C. chrysanthemi E. & D.^^ Spots subindefinite, 1 cm. or more broad, black; acervuli innate, amphigenous, 100-170 /x; conidia fusoid straight, 50-100 x 3-4. 5/z.
media and
its
numerous F pathogenicity was

in
I li.
380.
ponii.
Cylindrosporium Sporo formation in
The fungus causes dark blotches on the leaves of chrysanthemum.

C. clematidis E.
&
E.
y.;
Spots amphigenous, reddish-brown, round or subangular, 1-3
acervuli comparatively few, epiphyllous, immersed, scattered; conidia fusoid-linear, 75-80 x 2.5-3 m, somewhat curved, exuding in a
white mass.
It causes leaf spot of Clematis.'^
Later
is
described
a variety jackmanii which differs from the species in the acervuli not being confined to the spots, which are less definite, and exuding a black mass of spores which are themselves hyaline.
C.
hamuli E.
&
E.
Spots amphigenous, mostly hypophyllous, small, angular, limited by the veinlets, rusty brown; acervuli minute, black, amphigenous; conidia nearly cylindric, granular, 40-50 x 3 /x, oozing out in a white mass.
The cause
of leaf spots of hops.^^
Other parasitic forms are: C. saccharinum E. & E. on maple; C. colchici Sacc. on Colchicum; C. inconspicuum Wint. on lily;
564
C. orni (Pass.) Pegl. and C. viridis E. & E. on ash; C. quercus Sor. on oak;
An
and
jury.
C. cercosporoides E. & E. on tulip tree. unidentified Cylindrosporium is reported from
New
Jersey
in-
New York
as causing spots of
tomato with considerable
It is possible that this
was
in reality Septoria lycopersici.-^'
^^
Cryptosporium Kunze
(p.
562)
Acervuli discoid-conic, covered by a peridium, erumpent, forming a pseudo-pycnidium from the substratum; conidia fusoidfalcate, large, continuous, typically stipitate.
Some
forty species, chiefly saprophytes. C. leptostromiforme Kiihn forms black stromata on lupine. C. minimum Lau. is the cause of a rose stem disease and of
cankers.
Libertella Desmaziere (p. 562)
Acervuli covered,
irregularly
and tardily erumpent, conidia
filiform, falcate, elongate, continuous.
Some twenty
L. rubra Bon.
L. ulcerata
on Prunus =Polystigma rubrum. See p. 208, Mas. causes cankers on fig trees in greenhouses.^^*
Moniliales
(p.
479)
The Moniliales differ from the Sphaeropsidales in the absence of the pycnidium and from the Melanconiales in their somewhat loose, separate hyphse, not innate and closely aggregated
There are genera on the boundaries between these orders which are difficult to place, as for example Coryneum, some species of which are often put in Helminthosporium; Vermicularia which sometimes is confounded with
as in the Melanconiales.
Volutella, etc.
one of very great diversity and contains a multitude of forms. Many are only saprophytes while some are aggressive
The order
is
parasites.
5G5
Key
Hyphae
in
to Families of Moniliales
loose
more
or less
cottony
bright
I.
;
masses
Hyphae and conidia

colored
clear
or
Moniliacese,
p. 565.
Hyphse and conidia typically both dark

one or the other always dark
II.
Dematiaceae,
p. 594.
Hyphae compactly united or forming a globose to cylindric body which is

often stalked
Hyphal
body
i.
cylindric
e.,
to
capitate,
stalked,
a synnema or coryIII. less
mium
Hyphal body more or
sessile,
i.
Stilbaceae, p. 632.
globose,
e.,
a sporodochium
IV. Tuberculariaceae, p. 638.
Moniliaceae
HyphsD hyaline or bright colored, more or less fragile, lax, not cohering in fascicles; conidia concolorous, hyaline or bright colored.
Key
cylindric
to Sections of Moniliaceae
Conidia globose, ovate, oblong or short-
Conidia continous Conidia two-celled
1.
2. 3. 4.
Conidia three or more-celled

Conidia muriform
Amerosporae, p. 565. Didymosporae, p. 585. Phragmosporae, p. 588.

Dictyosporae, Helicosporae.
p. 592.
Conidia
cylindric,
spiral
or
convolute,
5.
cells.
.
usually septate
Conidia of several stellately grouped
6.
Staurosporae,
p. 593.
Moniliaceae-Amerosporae
Conidia continuous, globose or ovoid to short cylindric.
Key to Groups
of Moniliaceae-Amerosporae
little dif-
Hyphae very short, or obsolete, ferent from the conidia

Conidia not in chains Conidia in chains
1.
2.
Chromosporieae, p. 566. Oosporeae, p. 567.
566

distinct
HyphJB elongate and

conidia
from the
branched,
3.
Conidiophores simple or
apically swollen
little
Conidia solitary Conidia in heads Conidia not

in chains
Hartigielleae, p. 570.
4.
5.
Cephalosporieae, p. 570.
Aspergilleae, p, 572.
Conidia in chains
Conidiophores much-branched, conidia not in heads
Conidia borne irregularly on simple or branched but not inflated or

verticillate conidiophores
6.
Botrytideae, p. 574.
Verticilliae, p. 583.
Conidia
borne
on
verticillately
7.
branched conidiophores Conidiophores with inflated nodes upon which clusters of conidia are borne
8.
Gonatobotrytideae.
Chromosporieae
Hyphae short or
(p.
565)
obsolete, conidia not in chains.
Key
to Genera of Chromosporieae
Conidia solitary, at least not capitate Saprophytic Hyphae almost none
Conidia separate Conidia joined in twos or threes, not catenulate
1.
Chromosporium.
Selenotila.
2. 3.
Hyphae very Entomogenous Phytogenous
short, branched, septate
Coccospora.
4.
Massospora.
In fungi Conidia ovoid, smooth Conidia globose, vcrrucose
5. 6.
Myceliophthora,
Coccosporella.
p.
567.
In leaves, hyphae vermiform-tortuous Conidia capitate; hyphae lacking; biophilous
7.
Ophiocladium,
Glomerularia.
p. 567.
8.
567
Myceliophthora Costant
(p.
566)
Sterile hyphse creeping, simple or somewhat branched, bearing conidia at the apex; conidia solitary,
acrogenous, ovoid.
^^Jil
single species
M.
lutea Costant.
__
causes disease of
cultivated
mush-
^V/P^//^"^
rooms
m France
- ^ "*
.
^m O^^^^^^r^^^^TT^
(p.
Ophiocladium Cavara
Fertile
566)
torellip-
hypha?
fasciculate,
^ "X*
^^â^DCÛ^^''^^"^ \\\\ irr^

After Cavara.
tuose;
tic.
conidia
acrogenous,
Fio.381. O. hordeiaccrvulusand
spon.'s.
Two species of
which one, O. hordei Cav., parasitizes barley.
Oosporeae
Hyphse
(p.
5G5)
short or obsolete, conidia in chains.
Key
to Genera of Oosporeae
Conidial chains arising in the hyphae Conidial branches simple, arcuate Conidial
1.
Malbranchea.
Glycophila.
branches
dichotomous,
not
2.
arcuate
Conidial chains arising at the apex of the
hyphse Conidia globose,

Conidia
all
elliptic,
or ovate
of equal size
.
.
Sterile Sterile
hyphse very short or none.
3.
OSspora,
p. 568.
hyphae evident, rarely none Conidia not connected by an

isthmus
Growing within the substratum Haustoria none

Haustoria present Growing on the surface of living
plants
4.
5.
Monilia,
p. 568.
Oidiopsis.
6. 7.
8. 9.
Oidium,
p. 569.
Conidia connected by an isthmus. Apical conidium larger Conidia fusoid, acute at each end
Paepalopsis.
Halobyssus. Fusidium, p. 568.
568
Conidia cylindric or cuboid
Hyphse nearly
dric
obsolete; conidia cylin10.
Cylindrium.
Hyphse
distinctly present
11.
Conidia cylindric Conidia cuboid
12.
Polyscytalum. Geotrichum.
Oospora Wallroth
Fertile
(p. 567)
slender;
hyphae short, sparingly branched, conidia catenulate, globose to ovoid, hyaline or dilute-colored.
Over one hundred species, chiefly saprophytes. O. scabies Thax.î^- 217. 219-224
Vegetative hyphse rarely
1
ju
thick, curved
ir-
regularly, septate or falsely septate, branching; aerial hyphse at first white, then grayish,
evanescent,
spores. Isolation
Fig. 382. Oospora After Saccardo.
breaking
up
into
bacteria-like
and inoculation in pure culture by Thaxter showed this to be the cause of the common American form of scab of Irish potato,
firs
beet, turnip, etc.
O. abietum Oud. causes defoliation of
in Europe.^^^
Fusidium Link
Hyphse which are
colored.
(p. 567)
short, simple, not well differentiated
from the conidia
fusoid, catenulate, acute at the ends, hyaline or dilute-
In part =Nectria.
See
p. 201.
A poorly understood genus of over 50 species.

F.
candidum Link, on various
trees =Nectria ditissima.
See
p. 203.
Monilia Persoon
(p.
567)
Hyphse erect, branched, forming a dense mycelial felt, which produces numerous conidiophores; conidia catenulate, hyaline or In part=Sclerotinia. See light-colored, ovate or lemon shaped.
p. 136.
Some
fifty species.

fructigena Pers. on fructigcna. See p. 139,
569
M.
stone and
pome
fruits =Sclerotinia
M.
cinerea Bon. on stone and
pome
fruits =Sclerotinia cinerea.
See p, 137. M. cinerea Bon. on Prunus padus=Sclerotinia padi. See p. 140. M. cinerea Bon. on Vaccinium oxycoccus=Sclerotinia oxycocci. See
p. 140.
M. M. M.
laxa A.
& R. on apricots =Sclerotiiiia laxa. See p. 137. linhartiana Sacc. on medlars and quinces " =Sclerotinia
See
p. 141.
linhartiana.
crategi Diedicke on
Crataegus =Sclerotinia
crategi.
^^^
See
See
p. 143.
M.
p.
seaveri Reade on Prunus =Sclerotinia

fimicola Cast.
seaveri.'*^^'
140.
M.
& Matr.
is
a parasite
of
mushrooms, Clitocybe
and Pleurotus.
Oidium Link
(p.
567)
On
the surface of living leaves;

fifty species.
erect, simple conidiophores
hj'-phae branched, white, bearing with catenulate, ovoid conidia.
About
These conidial fungi in the main belong to the Erysiphales though some forms are placed in Oidium which clearly do not
belong to that ascigerous order. Salmon states that there are some forty-four apparently Erysiphaceous Oidiums listed; but that twenty-five of these grow on plants known to be the hosts
of ascus bearing Erysiphacese.
O. fragariae Harz.=S. humuli. See p. 175. O. leucoconium Desm.=S. pannosa. See p. 176.
Seep. 181. Grogn.=P. oxyacanth. See p. 183. farinosum Cke.=P. leucotricha. See p. 184. The following may also be mentioned O. erysiphoides Fr. on hop, clover, cucumber, etc.; O. chrysanthemi Rab. on chrysanthemums;
crataegi
:
O. O. O. O. O. O.
balsamii Mont. = E. polygoni. See p. 177. ambrosiaB Thum.=E. cichoracearum. See

monilioides Link. = E. graminis.
p. 178.
See
p. 179.
tuckeriBerk.=U. necator.
570
O. O. O. O.

mespilinum Thiim on Mespilus; tabaci Thlim on tobacco; verbenae T. & B. on Verbenas.
quercinum Thiim has been reported as the cause of much injury to oaks in Europe since 1907. The identity of the Oidium causing the epidemic is, however, much questioned. By some it is
regarded as a stage of Microsphsera alni, by others it under a separate name as O. alphitoides G. & M.^-^* ^^^
Hartigielleae (p. 566)
is
set
up
One genus, Hartigiella Sydow. The species H. laricis (Hart.) Syd. causes fall of needles of larch.^^^
Cephalosporieae
(p.
566)
Hyphse elongate; conidia
in heads.
Key to Genera of
Cephalosporieae
Conidia globose or oblong Conidia sessile on the head or nearly so

Fertile hyphse inflated at
apex
Apical vesicle globose-inflated Conidia sessile, not mucus-covered

Vesicle verrucose or muriculate
Fertile hyphse simple
Fertile hyphse sigmoid,
1.
(Edocephalum.
much
2.
...
.
branched
Vesicle hexagonally areolate.
3. 4.
Sigmoideomyces. Rhopalomyces.
Gliocephalus.
Conidia on stalks, mucus-covered

Apical vesicle clavate or lobed
Vesicle
disk-shaped,
stellate5.
6.
lobed
Vesicle clavate or subpalmate. ... Fertile hyphse not inflated at apex
Coronella.
Buseella.
Conidial head covered with

mucus
7.
Hyalopus.
Gliobotrys.
hyphse branches at tip Conidial head without mucus

Fertile hyphse
Fertile
with
verticillate
8.
with one head

9.
Conidia not separating
Papulospora.

Conidia separating
571
Head elongate Head globose or

Sterile Sterile
10.
Doratomyces.
Haplotrichum.
slightly clavate
11.
hyphae scanty
hyphse long, bent
decum12.
Cephalosporium,
p. 571.
Fertile hyphaj with 2 to several heads
Conidia Conidia
upright
in
on
verticillate
13.
conidiophores
Ccemansiella.
more
definite
heads
Fertile hyphse simple, with 3 to
several heads of conidia
on
14.
spines
Fertile
Botryosporium,
p. 571.
hyphse several
little
times
to
15.
3-fid
Trichoderma.
Conidia borne on
stalks or sterig-
mata
Fertile hyphae simple Fertile hyphse verticillate branched.
.
16.
Corethropsis.
17. Spicularia.
Conidia short cylindric Conidia without mucus

Conidia covered with mucus
18. 19.
CylindrocephaLum. Acontium.
Cephalosporium Corda
Hyphse creeping, conidiophores short, erect, not apically swollen. Conidia spherical or ovate, hyaline or slightly colored. The small spored condial forms often associated with Fusarium
(microconidia) belong to this form-genus.
Botryosporium Corda
Hyphse assurgent, simple or forked, elongate, irregularly laterally branched; fertile branches simple, with three or more short apical branches which end in heads of conidia; conidia globose to
ovate.
A genus of only about ten species. B. diffusum (Grev.) Cda. has been reported as parasitic on Casuarina; B. pulchrum Cda. on wheat
and
radish.
B. longibrachiatum (Oud.) Maire on various green-house plants.
672
Aspergilleae (p. 566)
Hyphae
well developed; conidia in heads, catenulate.
Key
to Genera of Aspergilleae
apex
Fertile hyphse inflated at
Fertile hyphse simple or nearly so
Sterigmata of apical vesicle none or

simple Conidia terminal on sterigmata Conidia lateral and terminal on
1.
Aspergillus,
p. 572.
ster2. 3.
igmata Sterigmata verticillately branched. ... Fertile hyphse dichotomous, branches

curved
Dimargaris.
Sterigmatocystis, p. 573.
Dispira.
4.
Fertile hyphse little or not at all inflated
Fertile hyphse verticillately

tip
branched at
Tips equally verticillate; conidia doli-
form
Tips unequally
globoid
verticillate;
5.
Amblyosporium.
conidia
Conidia without mucus

Conidiophores slender Conidiophores swollen, conidia en* closed in mucus.
.
6.
Penicillium,
p. 573.
7. 8.
Gliodcladium,
Briarea.
p. 574.
Fertile hyphse not verticillate at tip
Aspergillus (Micheli) Link
Hyphae effused, creeping; conidiophores erect, simple, capitate; conidia catenulate; sterigmata none or indistinguishable from the
conidia.
conidia are often found, the asci but rarely. A. fumigatus Brizi, is held responsible by Brizi^^^ for pathological changes in wheat seed which rendered them incapable of germination.
The
A. ficuum (Hen.)
figs.
Weh. and A.
phoenicis (Cda.) Lind. occur on
573
Sterigmatocystis Cramer
(p.
572)
As
in Aspergillus
but with the sterigmata branched in whorls
at the apex.
S. niger
Van
Tiegh.
slender, conidiophores erect, 800-1000 x 11-16 /x, thickwalled, hyaline or dark above, the apical globose swelling black;
HyphsB
basidia 40
/i
long, radiately arranged, sterigmata obclavate, 8-
Fig.
lus.
384.
AspergilStrass-
From
of
FiQ.
383. B. pulckrum.
Corda.
After
burger et
Book
fx,
Text Botany.
al.,
10 n long; conidia globose, 3.4-4.5
minutely verrucose,
violet-
brown, catenulate. On tobacco. S. ficuum (Reich.) Hen.

Conidia globose, 3.8 n, dark violet, thick-walled, smooth. In the fruits of figs in Asia Minor and United States.
S. luteo-nigra Lutz.
is
injurious to cacao in the tropics.
Penicillium Link
(p.
572)
Hyphffi creeping; conidiophores erect, apicallj' irregularly verticillate-penicillately branched; conidia catenulate, spherical, or
574

hyaline or variously colored.
elliptical,
For the ascigerous stage
see page 167.
The conidial stages of many Penicilliums have been given searching investigation and comparative study by Thom.^^* P. glaucum Lk.
Hyphse effused, creeping, septate, interwoven, white, conidiophores penicillate, branches single or in pairs, erect, forked; conidia
globose to broadl}^ elliptic, smooth, hyaline, with a tinge of green, 4 ix. It is the cause of rot of ripe oranges, lemons,
apples, etc.
P. italicum Weh.
is
described as a
22
wound
Fig.
385. Penicillium.
After Bref eld.
It is very parasite on oranges by Massee. similar in appearance to P. glaucum but a little greener; conidia elliptic-oblong, 7-9 x 4 /x.
i.
t. j- * /t^ \ o P. digitatum (Fr.) bacc. Similar to the preceding species in habit but the conidia are
white in mass, 4-G

to P. glaucum. P. olivaceum
//.
Often associated with, and similar in effects

is
Weh.
found on citrous
is
fruits;
P. luteum Zuk. on apple.
An
undetermined species
reported as the cause of a white dry
rot of sweet potatoes. ^'^'^
Various other species of the genus occur on fruits and vegetables

causing their decay.
Gliocladium Corda
(p.
572)
as in
Hyphse effused, spreading; conidiophores and conidia Penicillium but the conidia surrounded by a mass of mucus.
genus of onl}^ about ten species. agaricinum C. &. M. arrests growth and breaks the
pilei of
mushrooms.
Botrytideae
(p.
566)
Conidiophores elongate, simple or branched but not inflated, and the branches not verticillate; conidia borne variously, globose
or ovate to elliptic.

Key
to Genera of Botrytideae
575
Conidia smooth or scarcely roughened

Saprophjlic or apparently
parasites
so, often real
Conidia typically plcurogynous

Fertile hypha3 2 to several-furcate.. Fertile hypha) simple or nearly so
.
1.
Haplaria.
Conidia globose or ellipsoid Conidia short cylindric

Conidia acrogenous or plcurogynous Some intermediate joints of the hypha)
2. 3.
Acladium.
Cylindrotrichum.
swollen
and
denticulate,
4.
conidia-bearing
Physospora.
Intermediate joints equal

Conidia-bearing hypha; of two sorts, the upright alone denticulate Conidia-bearing hypha? of one sort
Fertile hyphae simple or nearly so
5.
Blastomyces.
Hyphae
not denticulate; conidia solitary

crust-like
6.
.
.
Hyphae forming a
stratum
Hypha) loose, cobwebby. Hyphaj denticulate; conidia usually grouped Hyphae everywhere denticulate,
7.
Hyphoderma. Acremonium,
p.
577.
bearing
conidia
8.
only at tip Hyphai denticulate or proliferous at tip alone
Xenopus.
Apex Apex
denticulate,
many9.
sporcd
inflated-ampulli-
Rhinotrichum.
Olpitrichum.
form, 1-spored
Fertile hyphae
10.
branched
Conidia globose to ovoid
Both
sterile
and
fertile
hypha; procumbent
Sterile
lar
hypha? intracellu11.
Hartigiella.
576

Sterile
hyphae superficial
Fertile hyphse vaguely
branched
Conidia acro-pleuroConidia
gynous on
12.
Sporotrichum,
p.
577.
one-
sided
sympo13.
dium
Fertile hyphso dichotomous; conidia ac-
Monopodium.
rogenous on spinelike branches

Fertile hyphse erect or as-
14.
Langloisula.
cending Conidia soUtary acrogenous

Fertile
hyphse spinybranched at apex. not Fertile hyphis

spiny-branched. Conidia loosely grouped about the apex Conidia not involved
.
15.
Plectothrix.
16.
Monosporium.
in
mucus
17.
Conidia on inflated
muriculate apices
Phymatotrichum.
Botrytis, p. 578.
Apices not muriculate or inflated.

18.
Conidia
Conidia
fusoid
involved
to
in
19.
mucus
cylindric
Fertile hyphse
Tolypomyria.
mostly pro20. Sporotrichella.
cumbent
Fertile hyphse erect or as-
cending Conidia fusoid
on
the
21.
upper side of curved branches
Martensella.
Conida acrogenous
Conidia-bearing branches terete.
. .
22.
Cylindrophora.

Conidia-bcaring branches ellipsoid
577
23.
Cylindrodendrum.
Biogenous Conidia smooth

Catenulatc
Solitary
24. 25. 26.
Ovularia,
p. 582.
p. 582.
Ovulariopsis,
Conidia densely spiny Conidia muricate or tuberculose-stellate
Ramulaspera.
Conidia globose Conidia merely muricate
Hyphae loose, cobwebby Hyphae woven into a subgelatinous

pellicle
27.
Sepedonium.
28. Pellicularia, p. ;382.
Conidia setose at apex as well as muricate

29.
30.
Chaetoconidium.
Asterophora.
Conidia tuberculose-stellate
Acremonium Link
conidia
or
(p.
575)
Hyphae subsimple, procumbent; conidiophores simple, short;

solitary,
hyaline
light
colored,
oval to ellipsoid.
genus of some ten species.

^'^'
.|
An undetermined species is recorded by ^-^ as causing disease of cucumHumphrey

bers in Massachusetts.
VjjjM
^^lP{\lfA
I'
Sporotrichum Link
(p.
576)
Fig. 386.
Acremonium.
Hyphae widely spreading, conidiophores simple, short; conidia solitary or in groups on separate sterigmata, ovoid or subglobose. Over one hundred twenty-five species are described, most which are saprophytes.
S. poae Pk.
of
much branched;
^^^^' Saccardo.
Hyphae creeping, interwoven, branched, continuous or sparingly

p., hyaline, forming a loose cottony stratum; conidia of two kinds; microconidia, globose or broadly ovate, 4-12 /x; macroconidia abundant, elongate elliptic to ovate elliptic, 1 rarely 2-septate, about three or four times as
septate, variable in thickness, 2.5-6
large as the microconidia.
578

The form
that
it
is
an atypical one
kinds
is
in of
produces two spores; one kind which

septate
usually-
Heald ^-^ also Stewart and Hodgkiss -^^ have described it as the
cause of bud rot of carnations, while
the latter authors also mention
it
in connection with a disease known " silver top" of June grass in as Fig. pos. 11, Hyphabear- ^^hich the ^ panicles wither as they 3S7.--|. -^ ing comdiophores and macroconidia.
13,
Hypha
nidiophores and After Heald.
bearing comicroconidia.
expand, though the authors express
doubt as to
, ,
its
actual causal relation
be the carrier of the spores.
to the disease. A mite appears to Cultural studies and cross-inoculation

identical.
showed the fungus form on the two hosts to be

Botrytis (Micheli) Link (p. 576)
Hyphse creeping; conidiophores simple or more or

dendritic
less
markedly
branched, erect, branches various, thin and apically thick and obtuse or cristate; conidia variously grouped pointed, at the apex of the branches, never in true heads, continuous,
globose, elliptic or oblong, hyaline or light colored. In part =Sclerotinia. See p. 136.
of some two hundred or more species, several of them economic importance. This form-genus contains manj^ parasites on various hosts. In some instances they are known to include ascigerous stages, (Sclerotinia), in their life cycle; in others no such relation is known, though it has often been assumed on quite untenable grounds. Specific limitations are but poorly understood and the relations between the various forms and between these forms and the asof great
A genus
some instances the same

manifest impossibility, B. cinerea.)
cigerous stages are in a state of much confusion c. conidial stage is claimed
f.
(p. 137).
In
by
different in-
vestigators as belonging to
(e.
g.,
ascigerous species, a S. fuckeliana and S. libertiana with

distinct
two

The more prominent forms
nizing that some of them 231-243 Ti r> B. cinerea Pers.
79
as described are given below,
ret
og-
may
be co-specifie.
Hyphae slender, constricted at septa, gregarious, simple or sparsely branched, erect, cinereous, conidia globose, pale. A form which occurs on the grape is usually referred to S. fuckeliana though there is not entire agreement on this point.
On
and produces dense tufts sclerotia form within the
the grape the Botrytis develops its mycelium in the berries of conidia over their surfaces. The
fruits.
Leaves and
canes are also affected, (see p. 140). On the lily Ward^^^ in a classic study demonstrated the parasitism of the fungus showaction to be dependent upon toxins and enzymes. The type in this case deviated someing
its
what from the usual B. cinerea

spores were a
little
in that its
larger than is usual, l)ut it nevertheless seemed to be this species. No
ascigerous stage
was found.
On Cyclamen and Primula Wehmer^^^ reports a similar case.-'*'' The fungus has also
often been reported on the cultivated gera^ *
f^g.
Bailey,-^^ and many others have reported a greyish mold on the leaves due to a Botrytis which is often cited as B. cinerea though it appears to form no ascigerous stage. The affected part of the leaf collapses and is covered with a conspicuous growth of the conidiophores and conidia. Small sclerotia are produced in consideral^le abundance when on artificial media but they, on germination, bear clusters, dense bushy tufts, of conidiophores. That this form on lettuce
mum. On lettuce Humphrey,
38s. b. cinerea. After Smith.
^"
Jones,
^^^
ever produces ascophores is doubtful. Smith describes a case in which linden stems l:eginning at the ground were parasitized by B. cinerea. The bark appeared to be
first
affected
and
sclerotia
formed
in the
cortical
parenchyma.
Similar cases are on record regarding the rose, various conifers, grape (Brizi) Aesculus, Prunus. In all of these cases the Botrytis seems to be B. cinerea and without ascophores.
580

of the power of B. cinerea to infect growing have been made with the conclusion that it is a weak parasite
Numerous
tissue
studies
^^^
and that to become aggressively parasitic it must first develop a vigorous mycelium saprophytically. Attempts to immunize plants against its attack have been made with partial success.^^^
Extensive studies of the condition of sclerotia and conidia formation
He
by B. cinerea (from grape) were made by Reidemeister.^'*' concludes that sclerotia form on all media favorable to growth
They are on an average 5-8 mm. in diameter, smaller under conditions of poor nutriment, high osmotic pressure or strong transpiration. They are often found in concentric rings
of the fungus.
and
induced by the presence of various agents Conidia are formed under condition of energetic transpiration and on media of high osmotic tension. Conidia and sclerotia vary inversely in production. Appressoria develop on all media where sclerotia grow and are favored by
their formation
is
which inhibit groAvth.
substances which inhibit growth. Conidia are suppressed by conditions favoring the formation of appressoria.
B. depraedens Cke. is a pest of the sycamore. B. fascicularis (Cda.) Sacc.^^
Cespitose, minute, brownish; hyphse erect, fasciculate, flexuose, brown, or semi-pellucid, branches hyaline; conidia in subglobose, white heads, oblong, large. A mold of egg-plants is attributed to this fungus by Halsted.
An undetermined Botrytis and another on Ribes.^^^

B. parasitica Cav.
is
reported on carnation
by Atkinson
Hyphse
large, short-pedicillate,
cinereous, sparse, erect, inflated at base; conidia ovate, on short branches, heads umbellate, hya-
line or dilute, cinereous,
16-20 x 10-13
^^
/x.
This
in Italy
is
said
by Halsted
to be the
cularly the bulbs, in
and
is
Jersey. It widespread and destructive.
New
form found on lilies, partiwas first reported by Cavara
B. poeoniae Oud. 23. 245, 246 Mycelium in the parenchyma of the host, hyphse erect, 0.25mm. high, protruding through the stomata, branches spirally
TPIE
FUNGI
WlllCri
CAUSE PLANT DISEASE
581
arranged, simple, or branched; conidia, numerous, in heads 12-15 /i across, oblong or ovate-oblong, lG-18 x 7-7.5 /x, hyaline or dilute colored.
It is reported as the cause of considerable injury to peonies in
different parts of the

sclerotia are
It is
United States. The greenish-black flat found inside the stems. as parasitic on young peony stems and mentioned by Bos
~'^'^
on
lily-of-the-valley
and
lilac.
248
B. longibrachiata Oud."2.
White, sparse, cespitose, 1.5-2 branched;

fertile
and
It
mm. high; mycelium creeping, hyphse, verticillate, hyaline, much branched apically inflated; conidia numerous, hyaline, oval, 4.5-5.5 x
as the cause of
2.5 m.
was reported by Thaxter ^-^ bacco in the curing house. The

veins.
stem rot of to-
affected stems are covered with
white velvety patches of mycelium which soon spread to the

^^^ reported by AderhOld on ferns. This is perhaps a form of B. cinerea. B. douglassi Tub. on fir may be B. cinerea and=Sclerotinia
The same fungus
is
fuckeliana.
See
p. 140.
B. citricola Brizi, closely related to B. cinerea, attacks oranges
and
citrus fruits resulting in
mummies.
B. diospiri Brizi attacks the persimmon fruit near time of ripening and prevents maturity. B. patula S. & Ber.^ô
Cespitose, minute, greyish-white, spots cottony, suborbicular;

fertile
hyphse assurgent, continuous,
filiform,
late; conidia large, globose or globose-elliptic.
branching, panicu30 fi in diameter,
On raspberries. B. infestans (Hazsl.) Sacc. is common on hemp in Europe. It sometimes is associated with Sclerotinia libertiana which has led
light yellow.
some
to
assume
its
connection with that fungus; but no such
genetic relation is probable. B. galanthina Lud. occurs on snowdrops. It is said to belong to Sclerotinia galanthina, see p. 141, but the asci have not been seen.
Undetermined species are reported on carnations, Ribes and

mangold.^^^
682
Ovularia Saccardo
(p.
577)
Hyphse, simple, or sparingly branched, erect, apically simple or dendritically branched; conidia globose or
ovoid, solitary, rarely in short chains.
Over seventy-five species, all parasites. O. necans (Pass.) Sacc. produces spots on quince leaves in Italy and France; O. canaegricola Hen. on economic species of
Fia. 389. Ovularia. After Sorakin.
Rumex
O. O.
interstitialis
Q. armoracise Fcl. on horseradish; B. & Br. and O. primulana Thlim. on primrose
leaves;
on Vicia; Sacc. on Primula; O. corcellensis

viciae (Frank.) Sacc.
O. alnicola Cke. on Alnus; O. rosea Fcl. on willow; O. villiana Mag. on lemons; O. syringae Berk, on lilac. O. citri B. & F. causes the white rust of lemons in Sicily.^^^ A lemon disease in Australia has been credited by McAlpine
to O. citri McAlpine. O. medicaginis Br.
^^^
& Cav. is on alfalfa; O. exigua (W. Sm.) Sacc. on clover.

Ovulariopsis Patouillard
&
Harlot
(p. 577)
Similar to Ovularia, except in the solitary, acrogenous, sub-
clavate conidia.
hyphse creeping, conidiophores erect, simple, septate, at with a single 1-celled hyaline, subclavate conidium. apex A small genus of leaf parasites, in part conidia of the Erysiphacese (Phyllactinia). See p. 187.
Sterile
O. ulmorica Del. causes a mildew of mulberries.-^"*

Pellicularia
Cooke
(p. 577)
Hyphse creeping, branched, septate, in a subgelatinous pellicle, conidia sessile.
583
A single species. P. koleroga Cke'. causes a coffee leaf rot in ^^-^ as causing serious leaf India and has been reported by Fawcett blight of the same host in the West Indies.
Verticillieae (p. 566)
Conidia acrogenous, on verticillate branches of the conidiophore.
Key
chains
to Genera of Verticillieae
Conidia solitary or loosely grouped, not in

Conidia-bcaring branches very short,
pulliform
am1.
Pachybasium.
Conidia-bearing branches terete or longer Conidia globose to ovoid
Tips of branches clavate, in twos

rectangularly
2.
Verticilliopsis, p. 584.
Tips cf branches not as above Conidia conglutinate into a stra-
tum
Conidia not conglutinate Conidia separating readily from
the tips
separating with ficulty from the tips Conidia cylindric or elongate
3.
Corymbomyces,
p. 584.
4.
Verticillium, p. 584.
Conidia
dif5.
Cladobotyrum.
Conidia-bearing branches or sporo-
phores 1-sporcd
Sporophores straight Sporophores uncinate Sporophores several-sporcd
6.
7.
Acrocylindrium. Uncigera.
Sporophorc
inflated verrucose
at
8.
apex Sporophore incurved, with seriate conidia below

Conidia capitate or dcncely spicate, not in
chains
Calcarisporium.
9.
Coemansia.
Conidia
sessile
Conidia capitate, involved in mucus

Fertile hyphae
smooth
10. 11.
Acrostalagmus,
Gloeosphaera.
p. 584.
Fertile hyphae asperate
584

Conidia
densely
spirally
spicate
at
12.
apices Conidia on small stalks
Clonostachys.
13.
Sceptromyces.
Conidia in chains
14. Spicaria, p. 585.
Verticillium Nees (p. 583)
Hyphse creeping; conidiophores erect, verticellately branched; conidia borne singly at the apex of the branchlets, globose-ovoid, hyaline or light colored.
A genus of some seventy-five species, which are in the main conidia of various species of Hypocreales. See p. 196. V. albo-atrum McA. is a weak wound parasite of the potato.
Verticilliopsis
Cast
(p.
583)
Fertile hyphse with verticillate branches, 2 or 3 at each node; fertile branches clavate; conidia in heads, surrounded by a slime.
monotypic genus. V. infestans Cast, infects
mushrooms
in culture.
Corymbomyces Appel & Strunk

Sterile
(p.
583)
hyphse creeping;
fertile
hyphse erect, septate, dichoto-
mous corymbose;
conidia ellipsoid, clustered in gelatinous masses at the apex of the branches. A single species, C. albus Appel & Strunk. on cacao in Africa.
Acrostalagmus Corda
(p.
583)
creeping;
septate,
Hyphse
phores
conidiorichly
erect,
^^-'-''^^^''nf
branched; conidia borne in slimy beads on the enlarged end of the secondary
verticillately
branches.
About
Fig. 390. Cross-section of a vascular bundle. Acrostalagmus threads in the
vessels.
fifteen species.
^' ^^^JUS Preu.
After
Van Hook.
HyphSB
,
CespitOSe, effuse, slcn,
subangular, contmuous or septate, conidiophores, 200-220 x 1.7-2 /x, erect; fertile branches continuous, straight or curved; conidia in spherical heads, 9-10 /i
der,

in diameter,
585
numerous, minute,
elliptic
oblong, 3.3-3.4 x 1-1.5
fj.,
hyaline.
It
and
causes a wilt of ginseng. The vascular bundles are yellowed the ducts plugged by the
I
mycelium. Entrance is apparently through the leaf scars. The fungus
? //*
Nk
was
isolated
by Van Hook
made.
~'^''
^^
and
cultural
article
studies
In a later
cHf
Rankin
has discussed what
appears to be this fungus under the
name
A. panax. A. vilmorinii Gue.^''^'^^^ causes a
disease of
China
asters
and a
species
closely related, one of cacao fruits.
Spicaria Harz (p. 584)
Hyphse
erect,
creeping;
much branched;
conidiophores conidia apical,
fig.
391. Spore-beurins
stalks of
catenulate, ovate or elongate, hyaline or dilute colored.
^pôêraL^bo/nîn headsând^r^
of slime.
species. S. solani Hart, is said to produce effects like those of Fusarium solani.^^^
About ten
.,
held together at first by a coat After Van Hook.
on the potato much

is
S. colorans v. Hall, the cause of cacao cankers conidial stage of Calonectria cremea. See p. 205.
probably a
Moniliaceae-Didymosporeae
(p.
565)
Conidia hyaline, or bright colored, 1-septate, ovoid oblong or short fusoid.
Key
to Genera of Moniliaceae-DidymosporesB.

Saprophytic or on fungi Conidia smooth
Fertile hyphae simple or nearly so
Hypha) inflated at apex or joints Hyphse denticulate, inflated at

apex; conidia fusoid
1.
Diplorhinotrichum.
586

Hyphse inflated at both apex and joints Hyphai not
inflated
3.
2.
Arthrobotrys.
Haplariopsis.
Conidia spirally pleurogynous Conidia solitary, acrogenous

or capitate
Conidia capitate at apex. Conidia solitary at apex

Fertile hyphse long
4.
Cephalothecium,
Trichothecium.
p. 586.
5. 6.
Fertile hyphse very short
Didymopsis.
Diplosporium. Diplocladium.
Cylindrocladium.
Fertile hyphse
branched
7.
Branching irregular Branching verticillate Branching dichotomous;
8.
sterig9.
mata subternate
Conidia echinulate; conidial
equal
Biophilous Conidia obliquely beaked
cells
un10.
Mycogone,
p. 587.
11.
Rhynchosporium, p. 587.
Conidia not beaked
Hyphse mostly simple, not

twisted
spirally
12.
Hyphse simple,
Conidia catenulate
spirally twisted.
13.
Didymaria, p. 587. Bostrichonema.

Hormiactis.
Fertile hyphse simple, short

Fertile hyphse verticillately branched.
.
14. 15.
Didymocladium.
Cephalothecium Corda
Hyphse prostrate conidiophores erect, simple, septate, conidia apical, subcapitate, oblong to pyriform, hyaline.
;
Five species, chiefly saprophytes.

C.
roseum
Cda.^'ô^ês
Cespitose in sul^rotund, rose colored spots, fading with age, byssoid; hyphae
Fig.
392.Spores
of
''^^''''
Eustace
Cepha- creeping, branched; conidiophores erect, ^^^^'^ continuous, hyaline; conidia simple, oblong-ovate, constricted at the septum,
capitate, light rose.

It is often
found following apple scab gaining entrance through
587
the injured cuticle and causing rot. A ring of pink conidiopliores and conidia is formed around the margin of the scab. InocuUition tests showed the fungus unable to penetrate through sound cuticle
though
it
readily
made entrance through

living
woinids.
It
has been
occasionally reported on
twigs and leaves and
as a
it
common
America
1899 as
saprophyte has long been kno\\ii.
The
first
it
account of
in in
was from
New York
'^
in
1902 though
"^^
was described
injuring pears in
Germany.
Mycogone Link
HyphcD
(p.
586)
intricately branched; conidiophores
short, lateral; conidia unequally 2-celled, the
upper larger, echinulate. There are about fifteen species of mycogenous fungi which are probably conidial stages of Hy-
pomyces.
See p. 200.
-^^
M.
host;
perniciosa Mag.""*
White throughout, byssoid, deforming the

conidiophores
less
/i.
short;
conidia
solitary,
more or
It
is
pyriform, almost colorless, 17Fig. 393. Mycogone, conidiophores and conidia. After Saccardo.
22 X 9-12
reported by Mrs. Patterson as the cause of a mushroom disease in America. A verticilliurn conidial stage was present but
no ascigerous form.
M.
rosea Link, also occurs on mushrooms.
Rhynchosporium Heinsen
(p.
586)
On leaf spots; hyphffi filiform, hyaline, creeping, septate; conidiophores erect, with incurved branches, hyaline, apically denticulate; conidia short-cylindric, with a short oblique beak, medially
septate, hyaline.
monotypic genus.
R. graminicola Hein. occurs on rye, wheat, and barley in Europe.
Didymaria Corda
(p.
586)
Conidiophores simple, conidia borne apically, ovate, hyaline.
About twenty species of leaf parasites. D. prunicola Cav. produces spots on plum
leaves.
588
Moniliaceae-Phragmosporae
(p.
565)
Conidia hyaline or bright colored, 2 to several-septate, oblong, fusoid or elongate.
Key to Genera
Fertile hyphse very short
of Moniliaceae-Phragmosporae
Uttle dif-
ferent
and from the conidia
Conidia in chains, cylindric or oblong. Conidia not in chains
1.
Septocylindrium,
p. 589.
Sporophore
inflated
3-celled,
upper
cell
much
2.
Milowia.
Sporophore not
obsolete
inflated,
sometimes
Conidia
ciliate at
apex and upper

3.
septum
Conidia not
ciliate
Mastigosporium,
p.
590.
Hypha5 lacking; conidia not aggregate

4.
Fusoma,
p. 590.
Hyphse
distinct;
conidia
ag-
gregate
Conidia in mucose glomerules Conidia in fascicles, not mu.
5.
Rotaea.
Paraspora.
cose
Fertile hyphse manifest
6.
and
distinct
from
the conidia
Saprophytic Conidia solitary or at least not capitate

Sterile
Sterile
hyphse lacking
7. 8.
Dactylella.
hyphae abundant Fertile hyphse branched
Monacrosporium.
Dactylium. Blastotrichum.
Cephaliophora.
Dactylaria.
Hyphse Hyphse
verticillately branched.
9.
irregularly branched.
10.
Conidia capitate
Fertile hyphse vesiculose at tip.
.
.
11.
Fertile hyphse not swollen
Hyphse simple,
sterile lacking
12.

Hyphsc
Parasitic
vcrticillatc;
sterile
589
hy13.
phsD present
Mucrosporium.
Conidia
toic!,
mucose-conglobate, often continuous
allan14.
AUantospora.
Conidia not nuicose-conglobate Conidia ovatc-cylindric or elongate, often catenulate

15.
Ramularia,
p. 590.
Conidia obclavate-pyriform
16. Piricularia, p. 591.
Septocylindrium Bonardin
(p.
588)
Conidiophores very short, scarcely distinct from the conidia, or in parasitic species distinct but short and inflated or denticulately sublo]:)ate at the apex; conidia oblong or cylindrical, one
to many-septate, catenulate, the chains often branched. About thirty species, a few of them of economic
tance.
S. areola (Atk.) P.
impor-
&
C.^^^-
'''
Spots amphigenous, pale, becoming darker in age, 1 to 10 mm., by the veins of the leaf, conidiophores amphigenous, fasciculate, subangular, limited
several
nodose, branched or not, times septate,
hyaline, 25-75 x 4-7 n; conidia oblong, usually abruptly pointed at the
ends, catenulate or not,
14-30 X 4-5
Leaf
/i,
hyaline.
spots are duced on cotton.
pro-
The
^^''-
394. S.
areola.
After Atkinson.
conidia and stalks are so abundant on the undersides of spots as to give them a frosted appearance.
S.
rufomaculans (Pk.) P.
&
C.
leaf,
Spots numerous or confluent and even covering the entire
reddish; conidiophores very short, hypophyllous, cespitose; conidia catenulate, variable, ellipsoid-oblong to cylindric, hyaline, 8-16 x
3-4
At.
It is
somewhat
injurious
on buckwheat
in America.
^^
590

Aderh -'^^ is reported by Aderhold as the cause Prunus and Pyrus.
of
S. radicicolum
death of roots of
Mastigosporium Riess
Conidiophores short,
(p.
588)
stipitate, continuous,
soid, large, 3-septate, hyaline,
hyaUne, conidia fuwith apical and subapical bristles.
small genus in part =Dilophia.
Fusoma Corda
Mycelium
(p.
588)
obsolete or poorly developed; conidia innate, fusiform
separate. F. parasiticum Percival, causes a wilt of hops.^^^
Ramularia Unger
Conidiophores
fasciculate,
(p.
589)
simple
or
with
short,
scattered
branchlets, often flexuose, nodulose, or denticulate towards the apex, hyaline or light colored; conidia acrogenous or acropleuro-
gynous on the denticulations, hyaline, sometimes subcatenulate,

oblong, cylindric, typically many-septate. About three hundred species. In part =Mycosph8erella. p. 243.
See
R. tulasnei Sacc. on strawberry =Mycosph8erella fragariae.

p. 244.
See
R. armoraciae
Fcl.^^
Spots amphigenous, subochraceous becoming gray; conidiophores fasciculate, continuous, subsimple, 40-50 x
2.5-3
fx;
conidia rod-shaped, obtuse, hyaline,

fx.
15-20 X 3-4
On
horseradish causing leaf spots. R. taraxaci Karst.
Fig. 395.
ciffi.
R.
Hypophyllous, spots purple-margined, hyphse 35-45 X 2-3 n, spores bacillar, simple, straight, hyaline, 18-30 x 2-3 n. armora,.
Sac-
After
On
dandelion.
^^''^-
R. spinaciae Nip.
is
on spinach;
R. betas Rost. on beet; R. necator Mas. on cacao;

R. geranii (West.) Fcl. on cultivated geraniums R. primulae TliUm.
Spots
rounded-angular,
subochraceous,
591
emarginate;
conidio-
phores amphigenous, 50-GO x 5 ix, continuous, somewhat denticulate, rarely branched; conidia fusoid-cylindric, 20-30 x 3-6 /n, continuous or 1-septate. On Primula.-^ R. lactea (Desm.) Sacc. is on violets;
R. heraclei (Oud.) Sacc. on cultivated Heracleums; R. onobrychidis P. & D. on leaves of sainfoin. R. cynarae Sacc. causes loss of artichokes in France and Africa. R. coleosporii Sacc. is on sweet potato in Porto Rico.^^^ R. modesta Sacc. is recorded for the strawberry.
R. narcissi Chit, and R. vallambrosae Br.
of leaves
&
Cav. cause disease
and stalks
of Narcissus.-
kills
R. goeldiana Sacc.
twigs of coffee.
Piricularia Sacc. (p. 589)
Conidiophores simple, rarely branched, conidia obclavate to pyriform, 2 to manyseptate, solitary acrogenous, hyaline. small genus of parasites.
P. grisea (Cke.) Sacc.^^^'-^^ produces pallid or water-soaked, spots on culms
and
leaves,
with age greyish; conidioof
phores in clusters
two or
five
from
the stomata, simple or rarely sparingly branched, greyish, septate; conidia single,
terminal in scorpioid cymes, ovate,
2-septate, 24-29 x 10-12 n.

It causes death of rice plant tissue and the disease called "blast." If affected
Piiicularia grisea. Fig. 306. a, conidiophores. h, germinating spores. After Fulton.
leaves or stalks be placed in a
damp
at-
about a day a delicate greyish fungus, the sporing mycelium appears.
mosphere
for
The fungus grows

the
well in culture
and applied to the
rice plants gives rise to
typical disease spots.
This species was originally described on Digitaria sanguinalis and the form on rice has been called
592
P. oryzae; but morphological characters and inoculation experiments indicate their identity on various other grasses. P. caudata A. & S. occurs on cacao.
To
the Moniliaceae-scolecosporae belongs only one genus:
Cercosporella Saccardo
filiform,
Hyaline throughout; conidiophores simple or branched; conidia many-septate. Distinguished from Cercospora only in
color.
The genus contains some seventy
species
of parasites.
C. persicae Sacc.^^*
Conidiophores cespitose, on discolored areas,

filiform,
2 to 3-branched, continuous; conidia

/x,
40-60 X 1-5
torulose.
in
The
conidia develop
abundance on the
lower sides of leaf spots of peach causing a frosty mildew.

C. narcissi Boud. occurs on Narcissus; C. inconspicuus (Wint. & Hohn) on lily.
C. pastinacae Karst.
.
u.
per-
Spots amphigenous, f uscus or whitish

filiform,
conidia
After
Sac-
somewhat curved,
fi.
slender, attenuate,
50-90 X 2
On parsnip and
other Umbelliferse.
C. albo-maculans E.
&
E.
Spots orbicular, white, dark-margined, conidiophores amphigenous, cespitose, 8-12 x 2 ju, hyaline, continuous; conidia cylindric, 40-68 X 2-2.5 fx, straight or curved, 3-septate. A common cause of pale spots on turnip leaves.
Moniliaceae-Dictyosporeae
or cubic.
(p.
565)
Conidia hyaline or bright colored, muriform, globose, ovoid
Key to Genera of
Saprophytic
Moniliaceae-Dictyosporeae
Hyphae much-branched; conidia

or globose, cells uniform
elliptic
1.
Stemphyliopsis.

Hyphce little-branched; conidia six-lobed and sarcinffforni, central cell larger,
colored, lobes hyaline
2.
593
Synthetospora.
Parasitic
3.
Hyalodema,
p. 593.
evansii Mag., recently described

Africa.
This group contains but one important pathogen, Hyalodema on Zizyphus in by Magnus
-'''
The Moniliaceae-Helicosporae contain no important

Moniliaceae-Staurosporae
Conidia hyaline or bright colored,
septate or continuous.
(p.
parasites.
565)
stellate, radiate or forked,
Key
to Genera of Moniliaceae-Staurosporae
1.
Hyphae lacking; conidia trident-shaped. ... Hyphse present Conidia globose to cylindric, permanently
attached to 2 or 3 divergent sterig-
Tridentaria.
mata
Conidia themselves stellate or radiate
Conidia bilobate-forked lobes
;
2.
Tetracladium.
parallel,
3.
contiguous Conidia narrowly digitate Conidia 3 to 4-radiate
4.
Pedilospora. Prismaria.
Titaea, p. 593.
Conidia
ciliate at
the apex
5.
Conidia muticate
Conidia 3-radiate
Conidia 4-radiate
Fertile hyphse
6.
Trinacrium.
Fertile hyphse
very short, simple branched

Titeae Saccardo
7. 8.
Tetracium. Lemonniera.
Conidiophores simple, continuous; conidia subradiately 4 to 5celled, the cells
unequal in
size.
small genus of little economic importance. T. maxilliformis Rost. has been found on the roots of clover in
Denmark. ^^^
Oi/4
THE FUNGI \MiICH CAUSE PLANT DISEASE
Dematiaceae
(p.
565)
Hj'phae dark or black, cobweb])y, loose, usually rigid, not cohering in definite fascicles; conidia typically dark and concolorous,
but sometimes the hyphse are dark and conidia clear, or the conidia dark and the h>'ph8e clear. This family parallels the Moniliacese
and certain intermediate forms must be sought
in both.
Key
Conidia globose,
cylindric
to Sections of Dematiaceae
oblong or short
1.
ovate,
Conidia continuous
Conidia 2-celled
Conidia 3 or more-celled
2. 3. 4.
5.
Amerosporae, p. 594. Didymosporse, p. 601. Phragmosporae, p. 608.

Dictyosporae,
Scolecosporae,
p. 615.
Conidia muriform
Conidia long, filiform or vermicular Conidia cylindric, spiral or convolute, typically septate
p. 625.
6.
Helicosporae.
Conidia of several stellately grouped
cells.
7.
Staurosporae.
D ematiaceae-Amerosporae
Conidia continuous, globose to oblong.
Key
to Subfamilies of Dematiaceae-AmerosporaB.
Conidiophores very short, scarcely distinguishable from the mycelium
Conidia borne singly Conidia catenulate Conidia in heads or racemes

Conidiophores manifest and distinct from the myceUum and spores
1.
Coniosporieae, p. 595.
Toruleae,
p. 595.
2.
3.
Echinobotryeae.
Conidia dark, rarely subhyaline Conidia not in chains Conidia capitate Conidia verticillate, or at least
eral
4. Periconiese, p. 597.
lat5.
Anthrinieae.
Conidia
borne
singly
on
short
6.
lateral
branches
Trichosporieae, p. 598.

Conidia solitary, acrogcnou.s Conidia in chains
7.
595
p. GOO.
Monotosporeae,
Haplographieae,
8.
p. GOO.
Conidia hyaline Conidia acrogcnous on short hetcromorconidiophorcs at the lower part or bases of erect hy]:)luTO Conidia single or catenulate
phic
9.
Sarcapodieae.
Myxotrichelleae.
Conidia
m heads
10.
Conidia on hyphie of the same kind Conidia in heads Conidia in chains
11. Stachylidieae.
12.
Chalarieae.
Conidiosporieae
(p.
594)
Conidia not catenulate, conidiophores short.
Key to Genera
Conidia spherical, Conidia elongate
elliptic or discoid
of Coniosporieae
1.
Coniosporium,
Fusella.
p. 595.
2.
Coniosporium Link
globose, ovoid or discoid, borne
line conidiophores.
Hyphse very much reduced; conidia dark, on short hyaeighty-five species, chiefly saprophytes.
^"Â
, .
About
sporiiim. After
Saccardo.
C. onobrychidis Mag. occurs on sainfoin; C. filicinum Host, on Pteris and other ferns.
Toruleae
(p.
594)
Conidia in chains.
Key
Conidia of two
ate
sorts,
to Genera of Toruleae
macroconidia catenul1.
Microconidia glomerate Microconidia internal, catenulate
Heterobotrys.
Thielaviopsis, p. oC6.
2.
Conidia
all alike
Hyphai dark
Chains breaking up readily Conidia globose or ovoid
3.
Torula,
p. 597.
596

Conidia clavate
4.
Gongromeriza.
Gyroceras.
Chains breaking up with Chains curved
difficulty
5. 6. 7.
Chains straight or nearly so
Hormiscium.
Monilochaetes,
p. 597.
Hyphse hyaline
"^
Thielaviopsis Went.
(p.
595)
Hyphse creeping, subhyaline; conidiophores simple, septate; conidia of two kinds; macroconidia catenulate, ovate, fuscous;
Fig. 399. T. ethaceticus. and After Wakker
Fig.
400. Torula.
Saccardo.
After
Went.
microconidia cylindric, hyaline, catenulate within the conidiophore. In part=Trichosph8eria.
Only two species, both of economic importance. T. paradoxa (d. Seyn) v. Hohn (=Chalara paradoxa.) Macroconidia 16-19 x 10-12 n; microconidia 10-15 x 3.5-5 mIt is the cause of a pineapple rot, in which role it was first described in 1886; and of a sugar cane disease. '^^ In addition to micro and macrospores the fungus possesses a pycnidial form. With variation of the substratum the spores vary
597
considerably from the typical. In disinfection tests Patterson and Charles showed the macrospores to be considerably more resistant than the microspores, also that fumigation kills superficial
spores and spores placed in incisions in the fruit. T. podocarpi Pet. is known from Podocarpus roots.^^^
Torula Persoon
(p.
595)
Hyphae decumbent; conidiophores short, scarcely different from the conidia, which are catenulate, breaking away singly or in groups, dark to black, oblong to fusoid, smooth or roughened. Some one hundred fifty species, chiefly saprophytes.
T. exitiosa d. Seyn is said to cause much injury to chestnuts. T. sphaerella Cke. causes a sooty mold of coffee.
MonUochaetes
Ellis
&
Halsted
^^
(p.
596)
like;
Hyphae brown; conidiophores obsolete or very short, conidia conidia in chains, moniliform, dark; some chains interspersed
with larger conidia.
Monotypic and poorly known.
M.
infuscans
Ell.
&
Hals.^^''
The mycelium grows subepidermally in sweet potato causing discoloration and withering. The conidiophores
from the surface bearing
their simple chains of conidia.
roots
arise
Periconieae
Conidia dark, capitate.
(p.
594)
Key to Genera of
Fertile hyphce simple,
Periconieae
but often with short
apical branches
HyphE with apical branches or conidiophores

Parasitic
1.
Periconiella.
Saprophytic Apex with

phores
heterogeneous
conidio-
Apex
swollen; conidiophores 3 to 4
2.
Haplobasidium.
598
THE
FUx\GI

conidiophores
3.
Apex not
swollen;
many
Apex short-branched, rarely simple Apex short-branched or simple Apex not swollen Apex swollen Apex capitate-branched; branches
2
to
3-furcate
Stachybotrys.
4.
5.
Periconia,
p. 598.
Stachybotryella.
and
spine6.
bearing Hyphse without apical branches or conidiophores
Cephalotrichum.
Conidia globose Conidia boat-shaped

dark-ringed Conidia fusoid,
line
7.
Trichobotrys.
curved;
hypha;
8.
Camptoum.
Acrotheca.
sometimes
subhya9.
Fertile
hyphse branched below the apex Hyphse forked below apex; conidia oblong
10.
Synsporium.
Hyphse repeatedly dichotomous; conidia

globose or elliptic
11.
Dicyma.
Periconia Bonordin
-^^
creeping, or obsolete; conidiophores simple, dark, apically fertile; conidia globose, fuscous, solitary on short sterigmata. Fig. 401.
Hyphse
Some
forty species chiefly saprophytes.
Trichosporieae
(p.
594)
Conidia dark, borne singly on short lateral branches.
Key
to Genera of Trichosporieae
Hyphse loose, typically saprophytic Hyphse vesiculose-inflated here and there

Conidia-bearing vesicles pleurogynous. Conidia-bearing vesicles acrogenous ... Hyphse not vesiculose-inflated
Fertile hyphse erect
1.
(Edemium.
Cystophora.
2.

Branches circinate at apex; conidia mesogcnous, muricatc Branches
spirally
599
3.
Acrospeira.
Streptothrix,
5C9.
twisted;
conidia
4. p.
exogenous
Hypha;
simple branches
or
with
straight
5.
Virgaria.
All hyphiE
more or
less
creeping
Branches curved or lash-like. Branches not curved Conidia spiny, rarely smooth Conidia smooth
Conidia
sessile
6.
Campsotrichum.
7.
Zygodesmus,
p. 599.
8.
Trichosporium.
Conidia on stalks Conidia on tooth-like sterig-
mata
Conidia on jar-like stalks .... Hyphse forming a crust, parasitic
9.
Rhinocladium.
Basisporium. Glenospora.
10.
11.
Streptothrix Corda
Conidiophores erect, monopodially branched, the branches spirally coiled; conidia apical or lateral, single, sessile or with short sterigmata,
dark colored.
small genus.
S. dassonvillei Broc-Ros. is
noted as the cause of mold of grain and fodder.337
Zygodesmus Corda
ter branched, light or
Hyphie and conidiophores creeping, the latdark colored, here and
there irregularly inflated, septate at the swellings; conidia globose or ovate, muricate, rarely
smooth, on short sterigmata or on basidia-like branches of the sterigmata.
Fig. 401.
Periconia.
Some
After Saccardo.
fifty species, chiefly non-parasitic. Z. albidus E. & H.^^
Ilalsted describes a disease characterized
by a
floury coating
on
violet leaves
and ascribes
it
to this species.
600
Monotosporeae
Conidia dark,
solitary, acrogenous.
(p.
595)
Key
Sterile
to Genera of Monotosporeae
hyphiB lacking
Fertile hyphse short
and
fascicled at
1.
base
Fertile hypliEe longer, separate
Sterile
Hadrotrichum.
2.
Monotospora.
Nigrospora. Sporoglena. Acremoniella,
hyphse present Conidia with a loose hyaline membrane Conidia without a membrane
Conidia with a large shining drop. Conidia without a shining drop
.
3.
4.
5.
p. 600.
Acremoniella Saccardo
Hyphse creeping, simple or ramose, hyaline or colored; conidiophores simple,

short,
subbulbous below;
conidia globose to ovoid, fuscous.
About a dozen
species.
A. occulta Cav.^^" forms brownish-yellow flakes on the stems of cereals;
A.
Italy.
Fig.
verrucosa
Togn.
on
wheat
in
402^cremonieUa.
After Bonorden.
Haplographicae
(p.
595)
Conidia dark, catenulate.
Key
Sterile
to Genera of Haplographieae.
hypha?
all
creeping or obsolete
Fertile hyphae simple, not
branched at tip Chains of conidia lateral 1. Dematium. Chains terminal

Conidia without isthmi
Conidia connected
isthmi
2.
Catenularia.
by
cylindric
3.
Prophytroma.
Fertile hypha?
branched
4.
Hypha) dendroid
Hormodendrum,
p. 601.

Hyphse capitate branched at tip. sterile hypha> erect and mixed witli
. .
.
001
5.
Haplographium.
Hormiactella.
Some
the fertile
6.
Hormodendrum Bonarden
Hyphse creeping; conidiophores
dendritically branched;
(p.
GOO)
erect, septate,
brown, variously
conidia catenulate on
ovoid,
the
branches,
globose,
olivaceous
to
fuscous.
About a dozen species. H. hordei Bruhne on barley stems and

leaves often reduces the yield. Spots brown, scattered over the entire leaf or
confluent, oblong; hyphae simple, septate; coH. hordei .. Fig. 403. 111 ., nidia various, cylmdric, rounded or subattenAfter Bruhne. uate, or ellipsoid to subglobose, verrucose.
.
Dematiaceae-Didymosporae
(p.
594)
Conidial 1-celled, dark, rarely hyaline, ovoid or oblong.
Key
the conidia
to Genera of Dematiaceae-Didymosporae
different
Hyphse very short or scarcely

Conidia not
in chains
from
Hyphse lacking
Hyphffi present, circinate Conidia in chains
1.
2.
Dicoccum, p. 602. Cycloconium, p. G02.

Bispora.
3.
Hyphai
distinctly different
from the conidia
Conidia smooth, muticate Conidia not capitate Conidia more or

first
less catenulate
at
Hypha? and conidia biform, the

latter 1-celled or continuous,
dark or hyaline
4.
Epochnium.
Cladotrichum.
Hypha> and conidia uniform Hyphai here and there inflated
5.
602

Hyphee not inflated Hyphse erect; conidia longcatenulate
6.
Diplococcium,
p. 603.
Hyphaj somewhat decumbent;

conidia
short-catenulate
7.
or finally solitary Conidia not catenulate
Cladosporium,
p. 606.
Hyphae
beautifully
flexuose8.
torulose
Polythrincium,
p. 603.
HypliEB not torulose or flexuose Hyphae inflated at tip, branched
9.
Pseudobeltrania.
Hyphse
not
inflated,
little
usually
short and
branched
1
Conidia merely acrogenous

septate
or 210.
Fusicladium,
p. 606.
Conidiophores rather long,

multiseptate
11.
.
Conidia acro-pleurogenous ..
12.
13.
Passalora, p. 607. Scolecotrichum, p. 607.
Conidia capitate
Conidia muriculate or Conidia muriculate Conidia
ciliate
Cordana.
Trichocladium.
Beltrania.
ciliate
14.
sterile
apex; fertile hyphee intermixed
at
and
15.
Dicoccum Corda
(p.
601)
Hyphae creeping, chiefly very short, simple; conidia elongate or short-clavate, dark. About a dozen species.
D. rosae Bon. produces spots on rose leaves.
Cycloconium Castaigne
(p.
601)
Hyphae
in the walls of the epidermis,
dichotomous branched,
very fugacious, black; conidia ovoid, solitary. There is one species:

C. oleaginum Cast. Mjêlium circinate, fugacious, black; conidia sessile, ovoid, yellow-green. It forms blotches on olive leaves and on peduncles of the fruit
in Ital}'
and France and
is
somewhat
injurious in California.
^^*
603
Diplococcium Grove
Conidiophores
erect,
(p.
602)
olivaceous;
septate,
branched,
conidia
catenulate, 2-ccllcd.
Fig. 404.
Cycloconium
is
FiG.405.-Clodosporiuincucumerinum, mycfUuni, a hyphal knot, conidiot)horo
and spores.
phrey.
After
Hum-
oleaginum. After Boyer.
D. conjunctum (Bon) Sacc. 458 geranium

,.r,v^;i,rv^
reported as a parasite of the
Cladosporium Link
(p.
602)
Hyphre decumbent, intricately-branched, olivaceous; conidia globose to ovoid, greenish. In part=MycosphaTella. See p. 243.
Some one hundred seventy-five species, many of them of economic importance. C. fasciculate Fr. on hyacinth = PI eospora hyacinthi. See p. 260. C. herbarum (Pcrs.) Lk. on many hosts =]Mycosphaerella tulasnei.
See p. 247.
604
C.

herbarum
^""^
(Pers.) Lk. var. citricolum.^^^'
2-2
Fawcett
The fungus was grown

resulting in
recognizes this as the cause of scaly bark of Citrus. in pure culture and inoculations were made
from forty to sixty days in typical spots. the fungus was re-isolated. C. cucumerinum E. & A.^^' ^s^-^s?
Effused, maculose; in
olivaceous,
From
these
forming
spots
mass greyish-brown, changing to dark on fruits; condiophores cespitose,
sparingly septate, simple, denticulate, pale; conidia ovoid, lemonshaped or fusoid; olivaceous, 10-13 x 3-4 ju. It causes watery
spots on cucumber leaves, also decayed spots in fruit. C. elegans Penz. is the cause of disease on various Icinds of
oranges in Italy. with the next.

C. citri Mas.283.
This species
288-291
is
in the literature
much
confused
Conidiophores tufted, erect, branched, septate, brown, 30-75 X 2-4 n; conidia fusiform, dusky, continuous, or 1 to 3-septate, 8-9 X 2.5-4 fi.
The cause
elos.
of scab
on lemons, sour oranges, satsumas and pom-
It
was grown
in artificial culture
by Fawcett.
C. carpophilum Thum.-^' =92-294. 459 Spots orbicular, often confluent, blackish-green, forming circles; conidiophores erect, simple, sinuous, septate; conidia
ovate,
obtuse,
continuous or
1-septate,
10-12 x
4-6 /x. This

of
is
the cause of the widely distributed scab
peach, plum, nectarine, apricot, cherry. The deep olive-brown hyphse are found intermingled with the hairs of the peach. The disease was first
The fungus was cultured in Austria in 1877. and inoculations were made by Chester.^^ carpophilum.' After Cobb. jj^ ^^le twig the fungus breaks the cuticle from the layers below and its hyphse project through cracks. Upon the
noted
leaf it causes shot holes.
k*i*
C. sicophilum Far. attacks C. fulvum Cke.32
fig fruits.
Conidiophores densely crowded rupturing the cuticle, sparingly branched, septate, nodulose, bearing a few conidia near the apex;
605
conidia elliptic-oblong, 1-septate, translucent, tawny, 10-20 x 4-
6m.
The
hyphffi are
abundant on the lower
sides of
tomato
leaves,
forming a mold, varying from whitish to purplish in color. It causes serious disease in Europe and America. C. condylonema Pass, is found on leaves of Prunus causing
spot and curl.
C. bigarardia is on Citrus. C. macrocarpum Preu.-
leaf
Subeffuse,
black; conidiophores subfasciculate, simple, some-
FiG.
407. C. fulvum. After Southworth.
what
fiexuose,
brown; conidia oblong, oblong-ovate, 2 to several
septate, obtuse, pale brown. On spinach leaves in New Jersey, causing disease. C. graminum Cda.
Clusters minute, irregular, scattered, greyish-brown; conidio-
phores
nidia
distinct,
erect,
simple,
concolorous,
continuous
nodulose-flexuose, to several-celled,
brown; corounded or
oblong.
It is
wheat
florets
reported that this fungus was commonly present on sterile and that inoculation with it increased such sterility
slightly.
C. oryzae S. & Sy. is on rice; C. orchidis C. & M. on Oncidium; C. pisi Cu. & Ma. on Pisum.
606
Spots
C. peoniae Pass. large, chestnut
brown,
hyphse
short,
simple;
spores
On peony. various, long, 1 to 2-septate. C. epiphyllum Mart, is on oak, sycamore, poplar, etc.; C. juglandis Cke. on walnut;
C. scribnerianum Cav. on beech
;
C. hypophyllum Fcl. on elm; C. tuberum Cke. on sweet potato tubers; C. scabies Cke. on tomato and cucumber;
C. oxycocci Sh. C. zeae Pk.
Sterile
on cranberry.
;
oblong, 4 X 20
hyphse hyaline, sub-cutaneous, erumpent spores M, continuous or 1 to 3-septate. In immature corn grains.
C. brunneo-atrum
elliptic-
McA.
is
on orange leaves and young shoots

in
in Australia;
C. javanicum
Wak. on sugar cane
Java causing root molds.

(p.
Polythrincium Kunze
& Schmidt
602)
Conidiophores erect, fasciculate,

lose,
black, simple;
regularly flexuose or toruconidia acrogenous, obo-
void.
Monotypic.
p. 220.
Inpart=Phyllachara.
See
P.
folii.
trifolii
Kze. on clover =Phyllachora
tri-
See p. 220.
Fusicladium Bonardin
jr
FiG.
(p.
602)
sparco-
Conidiophores
short,
erect,
straight,
ingly Septate, subfasciculate, After Corda. nidia ovoid or subclavate, 1-septate, acrogenous, solitary or paired.
trifolii.
408. p.
olivaceous;
continuous
or
In part=Venturia and Phyllachora. Over forty species, several pafhogenic. F. fraxini Aderh. on Ash. = V. fraxini. See
'
220 2S3
p. 255.
F. saliciperdum (All. See p. 255.
&
Pvb.) Land, on SaIix=V. chlorospora.
F. cerasi (Rab.) Sacc. on cherry, peach, =V. cerasi.
See p. 255.
607
F. pirinum (Lib.) Fcl. on pear=V. pirinia. See p. 253. F, dendriticum (Wal.) Fcl. on pomaceous fruits=V. inaiqualis.
See
See
F.
p. 253.
F. orbiculatum
p. 255.
Thiim on Sorbus=V.
insequalis var. cinerascens.
depressum (B. & Br.) Sacc. on Umbelliferffi=Phyllachora. F. betulae Aderh. on birch=V. ditricha. See p. 255.
F. tremulae Fr. F. fagopyri Oud. is found F. lini Sor. on Linum.
on aspen =V. tremulse. See on buckwheat;
p. 255.
F. eriobotryae Cav. attacks leaves of Eriobotrys.^' F. destruens Pk.
Conidiophores short, 20-50 n, fasciculate, continuous or 1 to 2-septate, basally, colored, clusters slightly olive-green; conidia acrogenous, continuous or 1-septate, subcatenulate, ellipsoid to oblong, colored, 7-20 x 5-7 n.
On
F.
oats.
effusum Wint.^^^
Spots minute, rounded, rarely efTused, confluent, smoky; conidiophores erect, simple or slightly branched, septate, torulose, brownish, lighter above, 100-140 x 4 /x; conidia oblong fusoid to
rhomboid, continuous or uniseptate, light fuscous, subtruncate, 17-24 X 5.5-7

fji.
pecan scab affecting the leaves, stems and nuts. on vanilla. An undetermined species is the cause of a black canker of Hevea.
It constitutes the
F. vanillae Zim.
is
Passalora Fries
&
Montaigne
(p.
602)
olive;
Conidiophores
filiform,
intricate multiseptate,
conidia
oblong to fusoid, acrogenous. A small genus quite similar to Fusicladium except for the pluriseptate conidiophores. P. bacilligera M. & F.
and P. microsperma
Fcl. occur
on Alnus.
Scolecotrichum Kunze
&
Schmidt
olive;
(p.
602)
subfasciculate,
conidia oblong or
ovate, pleurogenous or acrogenous.
A genus of some
thirty species very similar to Fusicladium.
608
S.

graminis Fcl.
Spots foliicolous, elongate, ochraceous; conidiophores densely fasciculate, filiform, simple, sinuose, 90-100 x
6-8
p.,
subcontinuous; conidia fusoid-obclavate,

ju,
35-45 X 8-10
It is
uniseptate, olive-bro^Ti.
common, causing leaf spots on grasses, especially on Avena and Phleum. It is described on the latter by Trelease.^^^ The mycelium collects below the stomata and pushes
its tuft of
S.
Fig.
hyphae through them. melophthorum P. & D. parasitizes melons

in France; ^^
is
and cucumbers
409. S. gramiAfter
Sacnis.
S. fraxini Pass, S. iridis F.
cardo.
&
R.
is
on ash. on Iris;
S.
S. avenae Erik, is
musae on banana. on
oats.
Dematiaceae-Phragmosporae
cylindric or vermicular.
(p.
594)
Conidia 2 to many-celled, dark, rarely light or hyaline, ovoid to
Key
to Genera of Dematiaceae-PhragmosporaB
Fertile hyphse very short or little different
from the conidia

Conidia not
in chains
Conidia muticate
Conidia united at base, fasciculate,

cylindric
1.
Cryptocoryneum.
Conidia separate Conidia straight ovoid to cylindric Conidia solitary
2. 3.
Clasterosporium,
Stigmina,
p. 610.
p. 609.
Conidia in bundles
Conidia fusoid-falcate
4.
Fusariella.
Conidia cuspidate or setose Hyphse dichotomous and broadened

at apex Hyphae not dichotomous
5.
Urosporium.
or broad6.
ened
Conidia
in chains
Ceratophorum,
p. 610.

Conidia not connected by isthmi Conidia connected by isthmi
Fertile hyphse distinctly different
7.
609
8.
Septonema. Polydesmus.
from
the conidia
Conidia solitary or nearly so, acrogenous for the most part Conidia muticate
9.
Conidia echinulate
Conidia smooth
10.
Helminthospora. Heterosporium, p. 610.

Ophiotrichum.
Hyphse creeping, radiate Hyphse short, ascending or

erect,
11.
conidia
ovoid
to
12.
oblong
Napicladium,
p. 611.
Hyphse
longer, rigid; conidia
ovoid to elongate
Conidia ovoid Conidia elongate
13.
14.
Hyphse
flexuous, pannose. ...
15.
Brachysporium. Helminthosporium, Drepanospora.
p. 611.
Conidia 1 to 3-ciliate at apex Conidia verticillate or capitate Hyphse dark Conidia acrogenous, forming a
16.
Camposporium.
head
Hyphse simple Hyphse branched at the apex

Conidia
pleurogenous,
verticillate
17.
.
Acrothecium.
Atractina.
18.
some-
what
Hyphse
rostrate
and naked at
19.
.
apex
Rhynchomyces.
Spondylocladium,
Neomichelia.
p. 614.
Hyphse not rostrate at apex. Hyphse hyaline or bright colored, apex denticulate
Conidia catenulate as a rule
20.
21.
Conidia arising from the interior of

the hyphse IV. Sporoschismeae.
Conidia
arising from the apex, sometimes solitary
V. Dendryphieae,
(p.
p. 615.
Clasterosporium Schweinitz
608)
Hyphae creeping, here and there swollen, erect, bearing 2 to several-septate, solitary, apical conidia.
610
genus of some seventy-five species. glomerulosum Sacc. on Juniperus leaves is often reported as Sporodesmium glomerulosum. '"' ^^' ^'^ C. carpophilum (Lev.) Aderh.'^' Aderhold by inocuC.
showed this fungus capable of causing prunaceous hosts though C. herbarium did not do so. Effuse, hyphse simple or short-branched, densely aggregated, septate, conidia elongate-fusoid, obtuse, 4 to 5-septate, slightly conlations, properly controlled,
gummosis
of
stricted at the septa. It is commonly seen as the cause of a broAvn spot on peaches. Spores do not appear in the young spots but are found
sparingly in older brown areas. Pure* culture inoculations by Stewart

FiQ.
peach twigs resulted in blackening 410. Stigmina. mosis.
on and gumalso de-
^^
After Saccardo.
^ amygdalcarum
(Pass.)
Sacc.
is
scribed
on rosaceous hosts. It is perhaps identical with C. carpophilum and may be connected with Pleospora vulgaris.^^^ C. putrefaciens (Fcl.) Sacc. causes spots on leaves of the sugarStigmina Saccardo
beet.
(p.
608)
Hyphae epiphyllous; conidiophores very short or obsolete; conidia ovate or elongate, 3 or more-celled, aggregated. S. briosiana Far. causes disease of apricots in Europe.
Ceratophorum Saccardo
(p.
608)
short,
erect;
HyphB creeping, scant; conidiophores
conidia
fusoid or cylindric, 2 to many-septate, dark or reddish-brown. A small genus. C.
setosum Kirch,
is
found on leaves and shoots of young plants

^^
of Cytisus, etc., in greenhouses;
C. ulmicolum E.
&
K. on Ulmus
leaves.
Heterosporium Klotzsch
(p.
609)
Hyphse subcespitose, smoothish, often branched; conidia oblong, 2 to several-septate, smoothish to granular or echinulate-
genus of forty species or more.

H. echinulatum
="^
611
(Berk.) Ckc."-i' Spots gregarious, on fuscous areas; conidiophores fasciculate from a stromatic base, 150-200 x 8 rarely shorter, flexuoseij.,
nodose, fuliginous; conidia at the nodes, o])long-cylindric, rounded at the ends, 2 to 3-septate, 40-50 x 15-lG /z, slightly constricted,
roughened, brownish. It causes a destructive mold on carnation leaves and stems.

first
The
H.
epidemic Avas noted by Sorauer gracile (Wal.) Sacc. was determined to be the cause of disin Berlin in 1883.
^"'^
ease of Ins, Narcissus and other Monocotyledons. H. variable Cke.ôe
Conidiophores flexuose, slender, more or
less
nodulose at the
septa; conidia cylindric oblong, 2 to 4-septate, minutely warted,
20-25 X 7-10
fjL,
pale olive.
On
spinach.
Other parasitic species are: H. ornithogali Klotz. on Liliaceae; H. laricis C. & M. on larch leaves;
H. auriculi Mas. on cultivated Auricula; no H. syringae Oud. on lilac leaves. H. minutulum C. & M. causes disease of hops.
Napicladium von Thiimen
(p.
609)
Conidiophores short, subfasciculate, smoothish; conidia acrogenous, solitary, large, oblong, 2 to many-septate, smoothish. A small genus.
N. janseanum Rac. is on rice. N. soraueri is a form of Venturia insequalis with somewhat atypical napiform spores. See p. 253.
Helminthosporium Link
Conidiophores erect,
(p.
609)
rigid, subsimple, fuscous; conidia fusoid or cylindric, pluriseptate, fuscous, smooth. to elongate-clavate In part=Pleospora. See p. 259.
Al)out two hundred species; several are important pathogens,

others saprophytes.
The
species
show
biologic differentiation into races similar to
612
that exhibited in the Erysiphacese, though morphologically they may be inseparable.
Fig. 411.
Helminthosporium
phoree and spores.
gramineum.
After King.
Conidio-
H. gramineum (Rab.)
See
p. 261.
Erik,
on grasses =Peleospora gramineum.

See p. 260.
H. trlchostoma=Pleospora trichostoma.
Fig. 412.
Helminthosporium
and
spores.
teres. Conidiophores After King.
H. teres Sacc.
Spots oblong, olive, amphigenous; conidiophores fasciculate, often crooked and nodulose, septate, brown, 100-130 x 12 ju;

conidia acrogenous, straight, cylindrical, ends rounded, 4
septate,
613
to 5-
not
constricted,
dark
olive-bro\vn,
100-115
14-
18 M-
On
oats and barley.

Ei.
H. avenae
Similar to H. teres, but the conidiophores scattered, 150-200 x 9-12 n, septate, brown; conidia cylindric, brownish, 4 to 6-septate,
80-100 X 15-16
M-
On
oats.
The conidia of the two last species infect grains and seedlings. The conidia spread the disease from the early infection centers
to other parts of the plants but the
mycelium remains local. H. bromi Died, on Bromus=Pleospora bromi, see p. 261. H. tritici-repentis Died.=Pleospora tritici-repentis, see p. 262. H. sativum (P.) K. & B.ô^
Mycelium branched, septate; conidiophores fasciculate, fuscous, brown, septate, 8-10 /x wide, sometimes swollen between the septa; conidia solitary, apical, dark brown, 6 to 11-septate, 105130 X 15-20
M.
The cause
of
Saskatchewan.
a destructive late blight of barley from Iowa to The disease manifests itself by dark colored,
elongate spots on the leaves. It also occurs on the glumes
and
spikelets,
sometimes even
re-
penetrating the grains.
ported
H. sorokinianum Sacc. is on wheat and rye H.

tritici
in
Russia;
Africa;
H.
are
Hen. on wheat in sigmoideum Cav.

Italy; while several
on
rice in
species
recorded on bamPass.
Fig.
413. Helminthosporium sativum from barley, spores and conidiophores.
boo.
After King.
H. turcinum
late, septate,
Spots, large, dry, brownish; conidiophores, gregarious to fascicu-
150-180 x 6-9 is, pale olive, apex almost hyaline, often nodulose; conidia spindle-shaped, acute, 5 to 8-septate,
pale olive, 80-140 x 20-26
It produces spots
fx.
on corn and sorghum
in
Europe and America.
H. inconspicuum C.
&
E."-
308-3io
614

brown; conidia
Conidiophores elongate, septate, nodose, pale lanceolate, 3 to 5-septate, 80-120 x 20 ii, smooth.
It
has been reported on sweet corn from Long Island by Stewart. H. gramineum, H. turcinum and H. incon-
spicuum are closely related, possibly identical. Johnson ^' concludes that H. gramineum with
its
ascosporic stage includes Piricularia grizea,
P. oryzse,
Helminthsporium oryzse and H. tur-
cinum.
H.
inaequalis Sh.
Sterile
bro^\^l;
hyphse
effuse,
much
erect,
branched, dark
septate,
conidiophores
variable
in length,
^'tho^poriiTm^^'^în-
6-8
ju
in diameter; conidia
both
ter-
^^^^^ ^^^
lateral,
more
or less curved,
3 to
n.
ffiqualis.
After
5-celled, thick-walled,
brown, 23-32 x 11-14
On
cranberry.
H. heveae Fetch, is on Para rubber; H. theae Bernard on tea in India; H. iberidis Poll, on Iberis and H. lunariae
in Italy.
Poll,
on Lunaria, both
Spondylocladium Martius
(p.
609)
erect,
Hyphae creeping, septate; conidiophores

conidia
a srticillate,
simple,
rigid;
fusoid, usually 3-celled, brownish.
small genus.
S. atrovirens Harz.^*
Conidiophores solitary or clustered, cj'lindric, septate, dingy, up to 400 n high; conidia elongate, ovate, apex 5 to 7-septate, concolorous with the conidnarrowed,
olive or brownish,
iophores, 30-50
X 6-9
M-
On
potatoes this fungus causes blackish to olive
spots soon depressed, 2-3 cm. across, which are beset with small black sclerotia and followed by dry
^^^ the sclerotia According to Appel & Laubert develop whorls of conidiophores. The species is said to occur in the British Isles, the Continent
rot.
"atrovi re~ns! After Massee.
and
the
in America.^^-
Its sterile
mycelium has been described under
name
Phellomyces.^^^
615
Dendryphium Wallroth
(p.
009)
Hyphae creeping or obsolete; eonidiophores erect, with short apical branches; conidia cylindric, 2 to many-septate, catenulate,
brown.
Some thirty or more species. D. comosumWal. is the cause of a cucumber

Dematiaceae-Dictyosporae
Conidia dark, rarely
light,
leaf spot in
England.
(p.
594)
muriform, globose to oblong.
Key
the conidia
to Genera of Dematiaceae-Dictyosporae.
different
Hyphse very short or scarcely

from
I.
Micronemeae.
Conidia not appendaged Conidia irregularly muriform

sarciniform
or
Conidia with a conic point at each

side
1.
Oncopodium.
Conidia without conic points Conidia globose to oblong

Conidia ovoid to oblong, loose Conidia globose to ovoid, aggregated
2.
Sporodesmium,p. 616.
Stigmella.
3.
Conidia sarcinseform, often coalescent

4.
Coniothecium,
p. 617.
Conidia as
if
composed
cells
of parallel
chains of
Chains of conidia never separating

5. 6.
Dictyosporium.
Speira.
Chains of conidia separating Conidia corniculate at apex Conidia in chains, often asperate or with
isthmi
7.
Tetraploa.
8.
Hypha?
distinctly different
Conidia of
from the conidia the same form
II.
Sirodesmium. Macronemeae.
Conidia not in chains or capitate Conidia bearing little conidia on their

surfaces
9.
Xenosporium.
616

Conidia single
Hyphse
alike
Conidia cruciate-divided, verrucose

10.
Tetracoccosporium.
Conidia
muriform,
typically
11.
smooth Hyphae decumbent Hyphse erect or ascending

Conidia
globose,
Stemphylium,
p. 617.
pleuro-
gynous Conidia around the apex of the hyphse Conidia conglobate around the base Conidia ovoid to oblong,
mostly acrogenous Conidiophores somewhat
lax, colored
12.
Coccosporium.
Trichaegum.
13.
14.
Macrosporiuni,
p. 618.
Conidiophores dark
rigid,
very
15.
Mystrosporium, p.
620.
Hyphse
of
sterile,
kinds, shorter fertile
two
longer
16.
Conidia capitate Conidia catenulate
17.
Septosporium, p. 620. Dactylosporium.
Hyphse velvety, Hyphse
erect,
subsimple;
18.
conidia caudate
crustose, various; conidia 2-
Alternaria,
p. 621.
celled; conidia-like ganglia sar-
cinseform
19.
Fumago,
p. 624.
Conidia of two forms, dark sarcinseform
and subhyaline
falcate
20. Sarcinella, p. 625.
Sporodesmium
Link.^^^ (p. 615)
Mycelium and conidiophores poorly developed;

fuligineus.
conidia ovoid
oblong, subsessile or short-stalked, rather large, clathrate-septate,
Over eighty
S.
p.
species.
piriforme
Cda.
on
oranges =Pleospora
hesperidearum,
p. 258.
260.
S. exitiosum
Kiihn on crucifers=Leptosphaeria napi,
THE FUNGI WHICPI CAUSE PLANT DISEASE

S. exitiosum var. solani
617
Schenck
is
is
reported as the cause of a
potato disease. S. solani-varians
Vanha
the cause of
potato disease in Europe, the foUage bearing brown spots and finally dying in a manner resembling death caused by Phytophthora.
Cladosporium and pycnidial forms are said

to exist.
S.
mucosum
Sacc.
holt
on cucumber
scorzonerae
fruit
was reported by Aderand leaves causing

causes a salsify
disease.
S.
Aderh.
stem and leaf disease. ^^'* Other parasitic species are: S. melongena Thiim. on egg plant; S. dolichopus Pass, on potato leaves
Italy; S. ignobile Karst.
S. S.
in
Fig. 416.
Sporodesmium
After
Sac-
antiquum.
cardo.
on asparagus; putrefaciens Fcl. on beet; brassicse Mas. on Brassica in Bengal.
Coniothecium Corda
(p.
615)
Hyphse obsolete or poorly developed; conidia
gemmiform
Over
in origin, variously septate.
fifty species of
very simple parasitic or

resembling apple common in Tas-
saprophytic fungi. C. chomatosporum Cda.
scab in
its effect is
noted as
mania^^^ and Australia.^'*^
Stemphylium Walroth
(p.
616)
Conidiophores
decumbent, intricately
branched, hyaline or smoky; conidia acrogenous, ovoid to subglobose, 2 to many-muriform''Tium '7?teTfa^: septate, fuligineus.
cardo.
Over thirty
species.
S.
ericoctonum B.
&
deB.
is
parasitic
on Erica
in green-houses.
618
S. citri Pa.
&
Ch.^^
Vegetative mycelium long, hyaline, becoming dark, 4 )u in diameter, septate; conidiophores short; conidia dark l)rown, subglobose to oblong, apiculate, irregularly muriform, 20-30 x 1215 M, usually in chains of three.
Arizona.
This was found associated with an end-rot of oranges from Inoculated in pure culture in oranges the fungus deIt is
veloped well.
S. tritici Pa.
perhaps the cause of the disease.

closely
Hyphae irregularly branched; conidiophores

4-5
fjL
septate,
in diameter; conidia catenulate, irregular, usually clavate,

/x,
constricted slightly at the septa, 24-35 x 12-15 It fuligineus, isthmus short, 3-4 fx in diameter.
is
vermiculate, described as
the cause of floret sterility of wheat.
'
Macrosporium
Fries (p. 616)

less
Conidiophores fasciculate, erect or not, more or
branched*
colored; conidia usually apical, elongate or globose, dark-colored. In part=Pleospora. See p. 259.
About one hundred eighty species, many of them saprophytes while others are important pathogens. M. commune Rab.=M. sarcinula parasiticum Thilm. on various grasses =Pleospora herbarium. ^^^' ^^^ See p. 260. This is reported by Thaxter ^^^ as the common black mold which
follows Peronospora on the onion and which occurs often also on onions not so diseased, being especially common on the seed stalks.
It is usually associated
with injured plants and
may
be important
only as a
wound
parasite.
M.
porri E.^^^
Effuse, fuligineus; hyphse short, simple, subfasciculate; conidia
elongate-clavate,
basally
attenuate,
multiseptate,
150-180
12-20
M.
It is
common on
seed onions, less
common on market
in all directions
onions.
finally
The dark mycelium penetrates the host
and
produces stromata below the stomata and sends up short hyphcs. M. alliorum C. & M. is also on onion; ^^
M. hurculeum E. & M. Amphigenous on rounded, grey
spots; conidiophores erect, ces-

pitose, flexuose,
619
brown, few septate, 70-80 x 5 /x; conidia brown, multiseptate, clavate, 200-225 x 21-26 mIt causes leaf spots on turnips, horse radish and other crucifers. M. brassicae Berk.
clavate, antenna^form, 5 to x 12-14 /x. 11-septate, 50-60 It is a common cause of black mold on cabbage, collards and other crucifers.
Mycelium inconspicuous, conidia
M. ramulosum Sacc. is on celery. M. catalpae E. & M. ^' On brownish spots; conidiophores brown,
curved, nodose, 8 to
12-septate, erect, amphigenous, 90-135 x 6 m; conidia brown, obovate to pyriform, submuriform, 27-51 x 15-27 n. Producing leaf spots on Catalpa in company with Phyllosticta
catalpae.
M. nobile Vize. is on Dianthus. M. iridis C. and E. and M. aductum Mas. are on iris; M. cheiranthi (Lib.) Fr. on Cheiranthus. M. tabacinum E. & E.^^^ causes thin, white amphigenous spots, 2-3 mm. with a narrow dark border; conidiophores effused, 35-45 x
3-4
12
n,
M, sessile
septate and torulose above; conidia obovate, 15-25 x 10or short stipitate, usually 3-septate.
It is
concentric, rusty brown, amphigenous spots, 3 to 5 mm. in diameter; conidiophores effused, amphigenous, slender, 40-70 x 3-4 fi, septate, often contracted at the septa, erect and more or
less
M. On
reported to cause white leaf spots on tobacco. longipes E. & E.
above; conidia clavate, 40-50 x 15-20 m, 3 to 7-septate, attenuate below into a distinct stipe. On tobacco. M. sarciniforme Cav. is reported by Walkoff ^^^ on red clover
torulose
in
Clermany where it causes the leaves to dry and die. nigricanthium Atk.^-^ Amphigenous; condiophores subfasciculate or scattered nodose, septate, olive-brown, 50-140 x 6-7 n; conidia olive-brown, constricted about the middle, rostrate at one side of the apex, 1822 X 36-50 M- On cotton.
M.
M. cucumerinum
E.
&
E.^"'^^'*
Epiphyllous on orbicular, subconfluent, brownish spots, 3-4
620

mm.
5-6
in diameter;
conidiophores fasciculate or
1
solitary, subgeniculate,
ix;
to 3-septate, 35-50 x
slender-stipitate,
conidia
clavate,
to
constricted, submuriform, 30-75 X 15-25 m; pedicel 25-35 /x long. On leaves, stems and fruits of cantaloupes.
8-septate,
somewhat
M. cladosporioides Desm. onion and many other hosts.

M. M. M. M. M. M.
is
on
beet, lettuce,
verrucosum Lutz. occurs on cacao; gramineum Cke.^-'' on sugar cane. uvarum Thiim. is reported on Vitis; violae Poll, on violets in Italy;
saponariae Pk. on Saponaria. macalpinianum S. &. Sy. is injurious to
Fig.
418. Macro-
sporium cucumerinum on canteloupe. Chester.
After
Pelargonium.
Mystrosporium Corda
(p.
616)
Conidiophores simple or sparingly branched, short, septate, fuscous, rigid; conidia elliptic, subglobose or oblong, pluriseptate,
muriform, dark, usually solitary, acrogenous.
Some twenty
species.
M. abrodens
Nebr.
is
described as the cause of a very serious
grain disease in France.
M. aductum Mas. M. alliorum Berk,
injures Iris bulbs;
forms dark spots on onion.
Septosporium Cda.
(p.
616)
sterile
intermixed with longer
hyphse;
conidia ovoid to pyriform, fuscous. A small genus.

S.
heterosporium E.
&
G.
Spots scattered, confluent or not, rusty brown, 0.5-1 cm. in diameter, conidiophores hypophyllous, fasciculate from the sto-
mata;
conidia variable, oblong cylindric, constricted septa, 20-40 x 5-7 m, separating into gemmae. ^" on the wild grape in California. Reported in 1888
at
the
621
Alternaria Nees.
(p.
616)
short;
Conidiophores fasciculate, erect, sub-simple, clavate-lageniform, septate, muriform, catenulate. In part=Pleospora. See p. 259.
conidia
Some
thirty or
more
species,
many
of
pronounced economic im-
portance. A. sp. on Tropoeolum=Pleospora tropceoli. See p. 260. A. trichostoma Died, on barley =Pleospora trichostoma.
p. 260. A. forsythiae Harter.^-^
See
Hyphse
cespitose,
n.
amphigenous; spot concentric zonate: conidia
18-60 X 10 X 16.5
It causes subcircular leaf spots
on cultivated Forsythia.
A. brassicae (Berk.) Sacc.
Conidiophores short, continuous, short-branched, apically equal, conidia elongate, fusoid, clavate, 60-80 x 14-18 n, 6 to 8muriform-septate, olivaceous.
On
crucifers.
A.
brassicae
(Berk.)
Sacc.
var. phaseoli Brun. occurs beans in Italy.
on
A. cucurbitae Let.^^^
^^^
may
be identical with A. brassicae.

It
was
in
.
noted
by Thaxcausing
Fig.
ter
Connecticut
.
blight of melons.
The black
419.a.
mold
IS
copious
the older
in-
,1
violte, germinating spores. After Dorsett.
circular spots.
Pure cultures were obtained and successful
oculations were
made on normal uninjured melon
leaves.
It is also reported leaf spots in Ohio,
by Selby^-^ as the probable cause of muskmelon and it is a common source of troubles on various
is
=^29
cruciferous hosts.
A. tenuis Nees.^^ A. violae 0.
reported
by Behrens on tobacco
seedlings.
&
D.
Conidiophores erect, pale-olive, septate, simple, 25-30 x 4 /i, conidia in chains at or near the apex of the conidiophore, clavately
622
flask-shaped, strongly constricted at the septa, olive, 40-60 x
10-17
M.
Circular leaf-spots are produced on violets. Spores are found on the spots only when conditions are most favorable, i. e., in a humid
air.
The
parasitism of the fungus was demonstrated
by
inocula-
tion with spores on living leaves in distilled water. A. panax Whet.-^^
Spots amphigenous,
circular,
becoming dingy white with a
Fig. 421.
A. dianthi.
3,
Mycelium showing
420. Altern a r i a. Spores and spore-bearing stalks.

ter
branching and septation. 4< Showing mycelium below stoma and hyphse emerging through the stoma. 5, Showing catenulate spores as borne upon hyphae. 6, Spores showing shape, septation and catenulation.
Af-
van Hook.
young cluster of hyphae. 8, An older cluster of hyphse. After Stevens and Hall.
7,
reddish-brown margin, covering half the leaflet or less; hyphse brown, septate, 5-7 n in diameter; conidiophores erect, tufted, somewhat irregular, especially at the tips, brown, septate, 100120 X 5-6 m; conidia brown, in chains of 5 or 45-65 X 15-20 ju.
6, elliptic
to oblong,
On ginseng causing leaf blight. A. dianthi S. & H.^î

Spots epiphyllous, ashen-white,
definite, circular.
Conidiophores
cespitose from stomata, amphigenous, dark-brown, 1 to 4-septate, erect, 1-25 from a stoma; conidia 26-123 x 10-20 n, clavate,
623
tapering, obtuse, basally dark-brown, slightly constricted at the septa, 5 to 9 times cross-septate and 0-5 times longitudinally
septate. It causes injury on carnation leaves and stems. ^'^""^ A. solani (E. & M.) Jones & Grout. '''' '''
Spots brown, circular to

nous,
irregularly
elliptic,
scattered
over
concentrically zonate, amphigethe leaf surface; mycelium
Fio. 422.
A.
solani, 2, spores
potato
leaf; ^,
germinating and penetrating the living showing catenulation of spores. After Jones.
light-brown; conidiophores erect, septate, 50-90 x 8-9 fx; conidia obclavate, brown, 145-370 x 16-18 n with 5 to 10 transverse
septa, longitudinal septa few, conidia terminating in a very long hyaline, septate beak 3^ the length of the conidium or longer.
and tomaand is widely prevalent. It was first described in 1882 in America but is now knoAvn to be widely destructive.^"''' On potatoes it was first recorded by Galloway in 1891. In 1891 also Chester ^^^ and Galloway ^^^ proved its pathogenicity by inoculations on
It causes early blight, a leaf spot disease of potatoes
toes,^^^
624
tomato and potato, the spots appeared

inoculation.
in eight or ten days after Jones, using pure cultures, confirmed the conclusions of Chester and Galloway, the disease spots appearing as early as the third to fifth day after inoculation on vigorous uninjured
The mycelium grows luxuriantly within the leaf but spores do not usually form until after the death of the supporting tissues when the conidiophores emerge through the stomata or by rupturOften no spores are formed and rarely are ing the epidermis. many present. The mycelium may live a year or more and resume
leaves.
sporulation the following season. A. fasciculata (C. & E.) Jones
&
Grout.i^^'
^^^'
^'"''
^'^
becoming almost black, Conidiophores light darker than the vegetative hypha? but like them echinulate, 30-40 X 4-5 ij.; conidia concolorous with the conidiophores, 35-66
or dark-brown,
X 16-20
n, obclavate, 3 to 6 times cross-septate, 1 to 2 longitudinal septa, apical cell hyaline. This fungus is associated as a saproph\i:e with the blossomend-rot of tomatoes and also causes a serious decay of the ripened
fruit.
tains a
The literature of the disease is rather voluminous and connumber of sjTionyms, among them Macrcsporium tomato.
M.
also
lycopersici, ]M. rugosa,
M.
fasciculata.
Altemaria solani has
been credited with

be identical.^^^
fici
this disease
and indeed the two species
may
A.
Far.
is
on
figs
A. tabacinum Hori on tobacco; A. vitis Cav, on Vitis.
An undetermined Altemaria accompanied by a Macrosporium was constantly found in Nevadillo bianco olives which were
shrivelled, particularly at the apex.
These fungi were regarded as the cause
of the disease.^*^
Fumago Persoon
(p.
616)
Hyphse decumbent, intricate, frequently pseudo-stromatic, black; conidiophores, erect, branched; conidia ovoid, oblong or sarcinseform, 1 to 2-septate.
A small genus, chiefly conidial forms of See pp. 192, 193.
Capnodium and
Meliola.

F.
625
See
camelliae
Cat.
on various hosts =Meliola
camelliae.
p. 193.
Sarcinella Saccardo (p. 616)
much
Hypha) decumbent, septate, branched, dark; conidiophores reduced; conidia of two kinds: 1, dark packet-like; 2, subhyahne falcate. Both are intermixed.
small genus chiefly conidial forms of Dimerosporium.
See
p. 190. S. heterospora Sacc.
on various hosts = Dimerosporium pul-
chrum.
See
p. 191.
Dematiaceae-Scolecosporae
(p.
594)
Conidia dark or subhyaline, vermiform or filamentose, multiseptate.
There
is
only one genus.
Cercospora Fries
Conidiophores variable, almost obsolete or well developed, simple or branched; conidia vermiform or filiform, straight or cm'ved, multiseptate, subhyaline to dark.
The genus
In part =Mycosph8erella. See p. 243. is a very large one, some seven hundred species, and
chiefly causing spots are often blanched and are rendered ashen colored in the centers by the presence of the dark hyphie.
leaf spotting.
contains very
many aggressive, important parasites,

The
The hyphae
borne.
are usually geniculate at the point of spore production, Fig. 427, and thus old hyphae bear traces of spores previously
C. cerasella Sacc. on cherries =Mycosph8erella cerasella. p. 245. C. gossypina Cke. on cotton =Mycospherella gossypina.
p. 248. C. circumscissa Sacc.
See
See
Spots amphigenous, circular, pallid, dry, deciduous; conidiophores fasciculate, nodulose, brownish, simple; conidia acicular, narrowed apically, attenuate, tinged brown, 50 x 3.5-4 /x.
626

various species of Prunus this causes leaf holes. on the almond.^^^
On
It is reported as especially serious
C. boUeana (Thurn.) Sacc.
Hypophyllous, spots subfuscous to olivaceous; conidiophores fasciculate, filiform, 50-80 x 5-6 n, non-septate, fuscous; conidia terete, fusoid, 35-40 x 7-8 /i, apically obtuse, somewhat constricted, 1 to 5-septate, olive-green.
On
figs causing leaf spotting.^"*^ C. viticola (Ces.) Sacc.
Spots
amphigenous, subcircular to irregular, 2-10

fasciculate, filiform, septate,
mm.
in
diameter, ochraceous, emarginate;
conidiophores erect, densely
50-200 x 4-5
n,
straight,
somewhat
denticulate,
ochraceous;
somewhat atelongate-obclavate, 3 to 4-septate, 50-70 x 7-8 n, o\\\etenuate, brown.^^^

conidia
It is
apparently an unimportant parasite

leaves.
is
on grape
C.
C. rubi Sacc,
on Rubus;
is
fumosa
musae
Pass, on leaves of Citrus fruits.
C. moricola Cke. C.
common on
on
mulberry;
leaves
in
Zimm.
banana
Java.
Fig. 423.
6.
C. concors.
C. roesleri Sacc. occurs in Europe, causing

i
.
Hyphaj emerging late mjury to the grape. through a stoma and p antn\]afa Winf 113 ^' anguiaia vvmt.
twining about hair. After Jones and
j.i
Spots roundish, angulato, whitish to cinereous, margined, 1-3 mm. in diameter, often
confluent; conidiophores hypophyllous, fasciculate, erect, straight or only slightly flexuose, simple, brownish, few septate, 78-105 x 5 n; conidia filiform-obclavate, long attenuate, hyaline, 7 to 16septate,
80-170 x 3.5
/x.
On
the currant.
C. oryzae Miy.^^ is on rice in Japan. C. concors (Casp.) Sacc^^^- 345
definite;
erect,
Spots amphigenous, pale above, whitish beneath, rounded, inconidiophores fasciculate or single from the stomata,
brown,
septate,
simple,
40-80 m high;
conidia
single,

apically variable in form, ovate to elongate, curved,
1
627
to 5-septate,
subhyaline, 15-90 x 4-G n. In America this potato parasite was noted in Vermont in 1905
and study
In
of lierbarium material revealed

it
two
earlier collections.
Germany
was known
in 1854
and
it
has been seen in
many
parts of Europe since, sometimes in epidemic form.^^^ Conidia are abundant on the spots on stalks emerging from the stomata. The superior and inferior hyphae differ considerably
in length
within the
and branching. Brown bead-like chlamydospores form leaf. The mycelium is strictly intercellular. The fungus
Fig.
424. C.
nicoti-
anae.
After Jones.
C. nicotianae, spores germinating and Fia. 425. entering stomata. After Jones.
was studied
lations
in artificial culture by Jones & Pomeroy^^^ and inocuwere made, diseased spots appearing about three weeks after inoculation by spraying with suspensions of spores.
C. nicotianae E,
&
E.
Spots amphigenous, pale, becoming white, with a narrow and inconspicuous reddish border, 2-5 mm. in diameter, conidiophores
amphigenous, tufted, brown, septate, 2 or 3-times geniculate above, simple or sparingly branched, septate, 75-100 x 4-5 /x; conidia slender, slightly curved, multiseptate, 40-75 x 3-3.5 m, hyaline.
On
tobacco
it
abundant near the center

C. raciborskii S.
causes leaf spots.^'^ The sporiferous hyphjE are of the disease spots.
&
Sy. on tobacco in Java
and
Australia,^''^ is
near relative of C. nicotianae.
628
C. apii Fr.
ter,
Spots amphigenous, subcircular, pale-brown, 4-6 mm. in diamewith a more or less definite elevated margin; conidiophores hy-
pophylous, light-brown, fasciculate, continuous or 1 or 2-septate, subundulate,
40-60 X 4-5 ju; conidia hyaline obclavate, or almost cylindric, 3 to 10-septate, slender, 50-80 X 4 M. A serious leaf spot is produced on celery,
parsnips, etc.^^"'^^^ C. beticola Sacc.^^.
352
bordered,
Spots amphigenous, brownish, purplebecoming ashy centered; co-
nidiophores fasciculate, short, simple, erect, flavous, 35-55 x 4-5 n; conidia elongate,
filiform
obclavate,
hyaline,
multiseptate,
75-200 X 3.5-4 mThis fungus, described in 1873, causes a very serious disease of beet producing
spots on the leaves. It is common and deThe structive in America and Europe.
conidiophores usually, though not always, emerge from the sto-
mata from a few-celled stroma and are amphigenous.
They vary
and
If in
in
length
septation
with age.
atmosphere
become
spores
infest
Fig.
After Fig. 426. C. apii. Duggar and Bailey. spots hoary, due to the large number of Each cell of the spore is present.
humid
the
The germ capable of germination. the host through the stomata

of
tubes
Pure
427. Fertile hy- cultures

After
cured by mycelium produces dense matted colonies of deep olive color and a greenish-grey aerial growth but no conidia. Found also on Spinach in Texas.
C. beticola.
phae and spores of

Duggar.
the fungus may readily be seHere the the usual methods.

C. flagelliformis E.
629
&
H.
Spots aniphigenous, indefinite, yellowish; conidia very long,

curved, tapering. The cause of spinach leaf spots. C. citruUina Cke.
Epiphyllous, spots orbicular, 2-4
mm.
in diameter, white with a
purple margin; conidiophores elongate, terete, pale olivaceous, conidia very long, attenuate above, few-septate, hyaline, 120-140 X 3 /i.
The cause
of leaf spots
on watermelon.^^'
C. cucurbitae E.
&
E.^''
Spots amphigenous, rounded, subochraceous, becoming thin and white, 1 to 4 mm. in diameter, border slightly raised; conidiophores
tufted,
4 /x, continuous, subgeniculate conidia linear clavate, 100-120 x 3-4 n, above, apically obtuse;
olive-brown,
hyaline, septate.
70-80 x
On cucumbers
curbitacearum.
in
America, associated with Phyllosticta cu-
C. melonis Cke. grows
and
set
New
Zealand.
What
on cucumbers and melons in England is probably the same fungus has been
^^-^ as a new genus Corynespora. up by Giissow
C. armoraciae Sacc.
Spots amphigenous, pale; conidiophores short, simple, 30^0 X 5 ju; conidia rod-shaped, cuspidate, 100-120 x 5 m, hyaline,
multiseptate. On horseradish.
C. bloxami B.
C. personata (B.
in
& Br. occurs on Brassica. & C.) E.

mm.
or
Spots hypophyllous, small, brown, orbicular, 2-4
more
diameter; conidiophores densely tufted, short, brown, continuous; conidia clavate, pale-brown, about 3 to 4-septate, 30-
50 x 5-6
M-
On
the peanut in the Southern United States and
West
Indies.^^^
C. cruenta Sacc.
Spots amphigenous, indefinite, reddish conidiophores, subf asci;
culate, simple, subdenticulate, light olivaceous; conidia obclavate,
curved, 60-80 x 4 ceous.
ju,
subacute, 6 to 7-septate, hyaline or oliva-
630

in America,
On cowpea and bean

damage.
Usually causing but slight
C. vignae Rac. (not E. cowpea in Java.^^* C. medicaginis E.
&
E.)
is
described as injurious to the
&
E.
Spots amphigenous, smoky to black, 0.5-5 mm. in diameter, orbicular, indefinite; conidiophores subhy aline, becoming brownish, continuous, geniculate, 35-45 x 4-5 m; conidia cylindricf usoid,
3 to 6-septatc, 40-60 x 3 n.
On alfalfa and
crimson clover. ^^^
&
& Cav, is found on sulla leaves; C. saccharii Br. d. H. C. longipes Butler, C. acerosum D. and C. kopkei Krug. are on sugar-cane. H., C. vaginae C. capparidis Sacc. is found on caper.
C. ariminensis Br.
C. asparagi Sacc. & C. caulicola Wint. affect asparagus. C. malkoffi Bubak causes an anise disease in Sadova. C. these v. Br. d. H. occurs on tea in India; C. violae Sacc.
Spots amphigenous, rounded, bleached; conidiophores short, greyish, 30-35 x 4 /x; conidia long and slender, rodshaped, multiseptate, hyaline, 150-200 x 3.5 /z.
simple,
It
produces a violet leaf spot.-
C. althaeina Sacc. occurs on hollyhock.
Spots
fasciculate, slender,
amphigenous, brown, 2-4 mm. broad; conidiophores 40 x 5 /x, few-septate, olive brown; conidia apical, cylindric, to obclavate or broadly fusoid, straight, 40-60 x 5 n, apically obtuse,
2 to 5-septatc, hyaline. C. kellermanii Bub.
Fig.
428. C.
rosEDcola.
Spots amphigenous, irregular, angular, olivebrown, up to 1 cm. across; conidiophores fasciculate, slender, few-septate, 150 ^ x 4-5 olive-brown; conidia fiUform, 50-150 x 4-5
/jl,
/j.,
After Soutiiworth.
It also occurs
on hollyhock and
5 ^^ 15-septate, straight or curved, hyaline. is nearly related to C. malvarum
Sacc.
C. rosicola Pass.^
Spots ochraceous,
fuscous-margined,
2-3
mm.
in
diameter;
THE FUNGI WHICH CAQSE PLANT DISEASE
631
conidiophores cespitosc, small, densely gregarious, fuliginous, subcontinuous, 20-40 x 3-5 î, conidia cylindric, straight, short, 30-50 X 3.5-5 subfuscous, 2 to 4-septate.
fjL,
On
roses.
C. hypophylla Cav.
species.
on
roses in
Europe
is
very
like
the preceding
C. omphacodes E. & H, and C. phlogina Peck, are the causes of rather unimportant leaf spots of cultivated phlox, C. neriella Sacc. is on oleander.
C. sordida Sacc. produces leaf spots C. angreci Roum. is on orchids; C. cheiranthi Sacc. on Cheiranthus. C. brunkii E.
zonale.)
and
defoliates
Tecoma.
&
G.
is
reported on the geranium (Pelargonium
C. resedae Fcl.^^^
Spots punctiform, greyish; conidiophores fasciculate, simple, continuous or few-septate, 50-70 x 4-5 fx, fuscous; conidia apical to linear, obclavate, 4 to 5-septate, hyaline, 100-140 x 2.5-3 /xSpots are caused on the mignonette and the plants are blighted.
The hyphfE appear through the stomata.

C. odontoglossi P. & D. occurs C. unicolor S. & P. on hly. C. richardiaecola Atk.^^
on cultivated Odontoglossum;
Spots amphigenous, black, with small white centers, subcircular, 2-6 mm. broad; conidiophores fasciculate, light-brown with a
reddish tinge, becoming reddish-brown, erect or apically flexuose, denticulate, 30-80 x 5 m; conidia hyaline, obclavate, 4 to 10
or
more
septate, 50-100 x 3-4

lily.
jj,.
On
calla
C. microsora Sacc.^^
Spots amphigenous, minute, brown, gregarious; conidiophores subfasciculate from a tubercular stroma, short, continuous, subolivaceous, 20-30 X 3 //; conidia filiform, stricted at the septa, olivaceous, 35-45 x 3.5 It causes spotting and defoliation of Tilia.
3 to 5-septate,
/x.
con-
C. cercidicola E.
Spots amphigenous, dull grey above, rusty-brown ^ beneath, with a blackish-brown raised border; conidiophores amphigenous,
632

brown, 90-114 x 3.5-4
n,
fasciculate,
subgeniculate above; conidia
oblong, clavate, faintly 3-septate, 30-40 x 5-7 /x. It seriously injures the Japanese red-bud and occurs also on the
American
species.^^'^
C. acerina Hartig is C. sequoiae E. & E.
on maple seedlings.
Large compact olivaceous tufts are formed on languid leaves; conidiophores ferruginous, brown, abruptly bent, subnodose, toothed, sparingly septate, 50-70 x 4-5 fx; conidia oblong, becoming clavate, 40-70 x 6 /x, concolorous with the hyphse, 3 to 5-septate,
constricted at the septa. It is said to seriously interfere with the growth of Sequoia in the eastern states. C. halstedii E.
&
E.
Spots hypophyllous, indefinite, brownish to olivaceous, 2^ mm. across; conidiophores few-septate, 100-150 x 5-7 n, undulate or
crisped; conidia obclavate, 65-80 x 5-7 n, 3-septate, constricted.
It
somewhat
produces blotches on pecan leaves and causes partial de-
fohation.289
Stilbaceae (p. 565)

Sterile
hyphae creeping, scanty;
fertile
hyphse collected into a
stalk-like or stroma-like fascicle, bearing conidia at the top,
more
rarely along the sides, pale, bright-colored or dark.
Key
Hyphse
to Sections op Stilbaceae
hyaline
or
light
I.
and
conidia
colored
Hyalostilbeae.
Conidia globose,
1-celled
elliptic or
oblong
1.
2-celIed
2. 3. 4.
3 to several-celled
Amerosporae, p. 633. Didymosporae. Phragmosporae.

Helicosporae. Phaeostilbese.
Conidia
filiform, coiled
Hyphse or conidia dark

Conidia globose,
elliptic,
II.
or oblong,
p. 635.
With
cross walls only

5.
1-celled
Amerosporae,

2-cclled
G.
633
3 or more-celled
7.
8.
Didymosporae. Phragmosporae, 637.

Dictyosporae. StaurosporaB.
Murifonn
Conidia of a stellatcly arranged group of
cells
9.
Hyalostibeae-Amerosporae
Bright or continuous.
light-colored,
(p.
632)
elliptic
conidia
globose,
or
oblong,
Key to Genera of Hyalostibeae-Amerosporae

Conidial part distinctly capitate or at least terminal
Conidia not in chains Head of conidia not gaping or ting above
split-
Head not spiny

Conidiophores of head normal Conidia covered with mucus
Synnema monocephalous
Conidiophores
verticillate
dendroid-
Without distinct mata With obpiriform mata

verticillate
sterig1.
Dendrostilbella.
sterig2.
Pirobasidium.
Stilbella, p. 635.
Conidiophores not dendroid3.
Synnema polycephalous
Capitula
on
on
extremely
4.
short branches
Polycephalum.
Tilachlidium.
spreading Capitula subulate branches ...
5.
on Capitula branches
Conidia without mucus
erect
6.
Corallodendron.
Synnema monocephalous
Conidiophores
twisted
spirally
7.
Martindalia.
634

Conidiophores more or
straight
less
Conidia
rhombic
or
8.
biconic
Rhombostilbella,
Ciliciopodium.
p.
635
Conidia globose to fusoid

9.
Synnema polycephalous
Terrestrial, large, 1-2 cm.;
conidia ovoid
10.
Macrostilbum.
Not
terrestrial, small;
co.
nidia elongate-ovate.
11.
Chondromyces.
Conidiophores conidium-like, septate;
monocephalous
radiating
spic-
12. Atractiella.
Head spiny with

ules
Spicules conic, granulate
13.
Actiniceps.
Spicules
with
many
persistent
curved
14.
branches at middle
Heterocephalum.
Head
of
conidia
below,
15. Pilacre.
splitting
above
conidia
Conidia in chains
Synnema with conidia above; without mucus
Synnema not pubescent Synnema pubescent Synnema with conidia below; with mucus
16. 17.
Coremium,
p. 635.
Lasioderma.
Microspatha.
conidia
18.
Conidial part cylindric or long-clavate Conidia more or less equally scattered
Sterigmata denticulate, branched ... Sterigmata none or simple Conidia in lateral heads or racemes
Conidia
bate
in
19.
Cladosterigma.
20. Isaria, p. 635.
racemes;
synnema
lo-
21.
Peribotryum.
Conidia in heads
Conidiophores with lateral nodes, usually escaping through the
stomata
Conidiophores without nodes, usually
22.
Helostroma.
entomophilous
23. Gibellula.
635
Stilbella
Lindau
(p.
G33)
Hyphse forming a corcmium which is capitate above; conidiophoros borne on the cap; conidia small, often enclosed in slime. Over one hundred species chiefly saprophytes. (Commonly
known
it
as Stilbum but the type of the genus being a
hymenomycete
was renamed.)
S. flavida (Cke.) Kohl, causes S. theae Bern, S.
is
a serious coffee disease.
on tea in India.
nanum Mas.
S. populi
on poplar =Mycosph8erella populi.

Rhombostilbella Zimmermann
causes the thread blight of tea. See p. 250.

(p.
634)
Synnemata
verticillate-stilbiform; conidia rhomboid to biconic,
acute, without mucus. Monotypic. R. rosae Zimm. is found on Liberian
coffee.^'^
Coremium Link
Coremium
fertile;
(p.
634)
cylindric, apically enlarged
and
conidia
very
small,
catenulate.
small genus. In part =Rosellinia and Penicillium. See p. 230.

Isaria Persoon (p. 634)
Stromata
tile
erect, clavate or branched, ferthroughout, hairy; conidia small, globose
to ellipsoid, hyaline.
Over one hundred

enous.
I.
species, chiefly
entomog-
fuciformis Berk,
is
reported from Engits
land
and Australia
graminiperda B.
forming
stromata
Fig.
429.
on the inflorescences of Festuca.

I.
Coremium
glaucum
Corda.
After
& M.
also causes con-
^^ siderable injury to grasses in Australia.
Phaeostilbeae-Amerosporae
(p.
632)
Dark
conidia continuous, globose to elongate.
636
Key to Genera of
Phaeostilbeae-AmerosporaB
Synnema setose Synnema naked

Conidia asperate, on minute basidia. Conidia smooth
,
.
1.
Saccardsea.
Basidiella.
2.
Synnema Synnema
carnose, racemose-branched fibrous or corneous, not
3.
Stilbothamnium.
racemose
Conidiophore lageniform
4.
Ceratocladium.
Conidiophore lacking, at least not

lageniform
Synnema
tic
stalked, fibrous
ellip5.
Conidia dark, globose to
Sporocybe.
Conidia hyaline Conidia ovoid to oblong. Conidia elongate or falcate.

. .
6. 7.
Graphium. Harpographium,
Glutinium.
Trichurus.
Synnema
Conidia in chains
sessile,
corneous
8.
Synnema setose Synnema not setose

Stalk branched above
9.
10.
Stemmaria.
Stalk simple or nearly so
Capitule loose
Base of synnema subequal; usually on stems Base of synnema perithecioid; usually on leaves
Capitule compact Conidia globose
11.
Stysanus,
p. 636.
12.
Graphiothecium.
Conidia echinulate
13.
Harpocephalum.
Heydenia.
Briosia,
Conidia smooth
Conidia pleurogenous Conidia acrogenous
14. 15.
Conidia ovoid to oblong
16.
Antromycopsis.
Stysanus Corda
Stromata
erect, cylindro-clavate, dark, rigid conidia in

;
an oblong
THE
PÛNGI
G37
or subglobose panicle,
line.
ovoid, lemon-shaped or fusoid, subhya-
Some
twenty-five species.
See Fig. 430.

in Italy;
in Ireland.
S. veronicae Pers. occurs
on cultivated Veronicas
S. ulmariae INi'W. S. stemonitis
on Spirea
Cda. causes a brown rot of potatoes in storage.

(p.
Phaeostilbeae-Phragmosporae
G33)
Conidia 3 to several-celled, oblong to cylindric, dark or hyaline.
Key
to Genera of Phaeostilbeae-Phragmosporae
Conidia capitate
Synnema simple Synnema black;

tate
conidia densely capi1.
Arthrobotryum.
Isariopsis, p. 637.
Synnema
fuscous
or
pale;
conidia
2.
loosely capitate.
Synnema dendroid-branched Conidia not capitate

Conidia catenulate
3.
Xylocladium.
4.
Dendrographium.
Conidia not catenulate

Stalk fibrous
Synnema simple
or branched conidia
;
acro-pleurogenous
5.
Podosporium.
Negeriella.
Synnema branched;
gcnous
Stalk parenchyma-like
conidia
acro6.
Conidia pleurogenous, on a disk. Conidia acrogenous
...
7. 8.
Riccoa.
Podosporiella.
Isariopsis Fries
Slender, dark or subhyaline, cylindric hyphse laxly aggregated; conidia in a lax panicle or head, cylindric or clavate. See Fig. 431. I. griseola Sacc.
Spots hypophyllous, ochraceous; coremium stipitate, dense. 200 X 30-40 composed of filiform hyphce; conidia borne on reflexed ends of the hypha?, cylindric-fusoid, curved, 50-60 the
iJi,
X 7-8
n,
grey,
to 3-septate, constricted.
It causes disease of beans.
638
Tuberculariaceae
(p.
565)
Hyphse compacted into a globose,
discoid, or verruciform body,
Fig. 430.
Stysanus. After Saccardo.
Fig. 431. Isariopsis. After Saccardo.
the sporodochium; sporodochia typically

latinous, white, bright-colored or
sessile,
waxy
or subgeetc.
dark to black. In part = Nectria, Claviceps and Hymenoscypha,
See
pp. 14G, 201, 211.
Key to
Sections of Tuberculariaceae
I.
Hyphce and conidia hyaline or bright-colored

Conidia globooe to fusoid or falcate Conidia continuous
Mucedinese.
1.
Conidia 1-septate Conidia 2 to many-septate

Conidia muriform
2. 3.
Amerosporeae, p. 639. Didymosporae. Phragmosporae, p. 645.

Dictyosporae.
4.
5. 6.
Conidia spirally coiled Conidia forked or cruciate
Helicosporae.
Staurosporae.
Hyphse
olive to
brown
or black; conidia
II.
concolorous, rarely
hyahne
Dematieae.
Conidia globose to elongate Conidia continuous
7.
8.J
Conidia 1-septate
Amerosporae, p. 654. Didymosporae.

Conidia 2 to many-scptatc Conidia muriform
Conidia
filiform, hyaline
9.
639
p. 657.
Phragmosporae,
Dictyosporae,
Helicosporae.
10.
p. G58.
11. 12.
13.
Scolecosporae.
Conidia spirally twisted Conidia angulose-stellatc
Staurosporae.
Tuberculariaceae-Mucedineae-Amerosporae
(p.
638)
Conidia hyaline, or bright-colored, continuous, globose to fusoid;

hyphaB hyaline.
Key
to Genera of Tuberculariaceae-Mucedineae-Amerosporeae
Sporodochia smooth or nearly so

Conidiophores normal Conidia muticate
Conidia not covered with mucus Conidia not acrogenous-capitate Sporodochium girt by a heterogenods
cup Sporodochium without a heterogenous cup

Conidia not catenulate or scarcely so Conidia escaping from interior of
1.
Patellina.
hyphse Conidiophores branched

Conidiophores simple Conidia arising on outside of hyphse
2.
3.
Endoconidium,
Trichotheca.
p. 641.
Conidiophores lacking Conidia large, pellucid
Conidia globose Conidia oval
4.
5.
Sphaerosporium.
Diaphanium.
Pactilia.
Conidia small, not pellucid

Conidiophores present Conidia plcurogcnous
pleurogenous Conidia globose
or acro-
6.
7.
8.
Beniowskia.
Tubercularia,
Fusicolla.
p. 642.
Conidia ovoid to oblong Conidia fusoid to cylindric Conidia acrogenous
9.
640

Conidiophores verrucose Conidiophores not verrucose
Uredinicolous
10.
Dacrymycella.
Tuberculina.
11.
Not
uredinicolous
Sporodochia globose Conidia globose; conidiophores short Conidia ovoid conidiophores

;
12.
.^gerita.
branched
Sporodochia pulvinate Conidia acicular
Conidia terete-oblong
13.
Granularia.
14.
Kmetia.
Bactridiopsis.
15. 16.
17.
Sporodochia disk-shaped Sporodochia cupulate Sporodochia verruciform

effuse
Hymenula. Hyphostereum.
or
Conidiophores simple Conidiophores radiate, united at base

Conidiophores not arising from a cellular mass.
.
18.
Clinoconidium.
Ustilaginoidea,
p. 643.
Conidiophores arising
a cellular mass
from
19.
Conidiophores not united or

radiate
20. Sphacelia, p. 643.
dendroid
21.
Dendrodochium,
p. 643.
Conidia
in chains
Conidia covered with mucus. ... 22. CoUodochium.

Conidia without mucus Conidia globose Conidia hyaline Conidia blue
23. 24.
SphaerocoUa.
Sporoderma.
Conidia
elliptic to
oblong
disk-shaped,
25.
Sporodochium
orange-red
Necator,
p. 643.
Sporodochium
whitish
subglobose,
26. Patouillardia.
Conidia cylindric
Sporodochium dilated
stalked
above,
27. Bizzozeriella.

Sporodochia globose to verruciform
G41
Sporodochia gelatinous, sessile Sporodochia not gelatinous,

short-stalked
28.
CylindrocoUa.
Sphaeridium.
29.
Conidia
acrogenous-capitate; dochia turbinate
sporo.'30.
Cephalodochium.
Thecospora.
Epidochiopsis.
Conidia covered with mucus
Sporodochium globose, hardened

Sporodochia verruciform or subeffuse.
Sporodochia discoid Conidia ciliatc
Conidia Conidia
1-ciliate
.
31.
32. Illosporium, p. 643. 33.
at base only
34. Stigmatella.
1-ciliate at
each end each end

conidiainternal
35. Thozetia.
36.
Conidia 7 to
Conidiophores
8-ciliate at
Chaetospermum.
Scoriomyces.
with
bcaring areoles
37.
ciliate
Sporodochia setulose,
or uniformly
woolly Sporodochia woolly or setulose
Sporodochia
late
setulose;
conidia
catenu38. Periola.
Sporodochia woolly or velvety; conidia

capitate
Conidia globose Conidia oblong. ...
39.
40.
Dacryodochium. Lachnodochium.
Sporodochia
ciliate at the margin Sporophores none; conidia coacervate
41. Volutellaria.
Sporophores distinct Conidia in chains

Conidiophores below
6-ciliate
42. Volutina.
above, united
43. Guelichia.
Conidiophores not
ciliate
or united. ...
44. Volutella, p. 644.
Endoconidium
Prillieux
&
Delacroix
(p.
639)
Sporodochia pulvinate, white; conidiophores hyaline, racemose; conidia hyaline, rounded, formed within the conidiophore and
escaping apically. A small genus, chiefly saprophytes.
642
E.

temulentum
P.
&
D. =Hymenoscypha temulenta.
See
p. 146.
Tubercularia Tode
(p.
639)
sessile
Sporodochium tubercular or
smooth, rarely with
wart-like,
or
subsessile,
bristles, usually reddish;
conidiophores very
Fig.
432. Hyphaa
Fig. 433. T. fici, sporodochium, showing setaj and conidial formation. After Edgerton.
of Tubercularia, bearing couidia.
After Durand.
slender, usually branched; conidia apical, ovate to elongate part =Nectria. See p. 201.
In
Over one hundred species, chiefly saprophytes. T. vulgaris Tode =Nectria cinnabarina. See p. 202.
T.
fici
361
Edg.7^'
light pink, variable in size
Sporodochia scattered or gregarious, superficial or subcuticular, up to 3-4 x 1-1.5 mm., smooth, irregular
in
outline; conidiophores crowded, hyaline, 20-27 x 1-2 n; conidia small, clear, elliptic to oval, regular in size, 5-7 x 2.5-5 ft;
THE FUNGI WHICPI CAUSE PLANT DISEASE

setse scattered or
643
abundant, variously placed, straight or curved, or subhyaline, septate, papillose, 60-90 x 4-6 /Lihyaline lt is the cause of a fig canker.
Tuberculina Saccardo
Several species occur in sori of the Uredinales. Ustilaginoidea Brefeld, a small genus of Ascomycetes, one species of which, U. virens, on rice is Icnown only in the conidial stage.
See
p. 214.
Sphacelia L^viell^
(p.
640)
Sporodochia planose,
effuse,
stromate or sclerotioid; conidio-
phores short, simple, filiform; conidia apical, ovate. A small genus, chiefly conidia of Claviceps and related genera. S. segetum Lev. =Claviceps purpurea. See p. 212.
S. typhina (Pers.) Sacc.
=Epichloe typhina.
(p.
See p. 210,
640)
Dendrodochium Bonardin
Sporodochium pulvinate or verruciform, white or light-colored; conidiophores verticillate, branched; conidia acrogenous, ovoid to
oblong.
A genus of about forty species. D. lycopersici March is found on tomatoes

Necator Massee
(p.
in Belgium.^^''
640)
Sporodochium
erumpent,
small,
elliptic,
slightly
convex,
becoming
orange-red; conidia oblong or
catenulate, contents orange.

^^^
Monotypic. N. decretus Mas.
is
a dangerous parasite of
(p.
coffee, tea, etc.^^^'
Illosporium Martins
641)
Sporodochia wart-like, pulvinate or subeffuse, white or lightcolored, subgelatinous or
globose, sigmoid, variable,
waxy; conidiophores variable; conidia embedded in mucous. There are some
forty species.
I.
maltfoliorum Shel.
Spots suborbicular or coalescing and becoming irregular, brown
644

and with a small gray spot near the
center,
or mottled with gray
5-15
mm.
in diameter; sporodochia hypophyllous, minute, gelat-
inous, yellow-amber, becoming black, spherical, becoming discoid or irregular, 150-160 n in diameter; conidio-
phores branched; conidia oblong, 1-3.5 x 4 ju. ^^^ It is said by Sheldon to be one of the most
common and destructive causes of leaf spots of the apple often resulting in nearly complete deIn the centers of the leaf spots other foliation.
Fig.
spots bearing other species of fungi are often found, leading to the thought that perhaps the Illosporium in such cases results from secondary infection in the wounds made by the earlier fungus. The sporodochia are hyporium maculicola.
After Saccardo.
434. lUospo-
phyllous, often hidden
by the normal pubescence
of the leaf.
Volutella
Tode
(p.
641)
Sporodochia discoid, regular, margin ciliate, sessile or stipitate; conidiophores usually simple; conidia ovoid to oblong.
Some seventy
species.
V. leucotricha Atk.
Sporodochia convex-discoid, white to pale flesh-color;

filiform, few-septate,
;
setse few,
subhyaline conidiophores densely fasciculate,
filiform; conidia oblong.
On cuttings in greenhouses. V. fructi S. & H.

Spots on the
circles, fruit, circular;
sporodochia, numerous in concentric
subcuticular,
erumpent, elevated 200250 n, 150-400 n in

diameter;
black;
mycelium
distributed
to
3Fig. 435.
setae
throughout the sporodochium,

septate,
black, acute,
V.
fructi.
Sporodochia in section.
smooth,
At;
After Stevens and Hall.
100-400
5-8
co-
nidiophores
elongate,
hyaline,
simple,
25-35
/x;
conidia
smooth, oblong-fusoid to falcate-fusoid, hyaline or sub-olivaceous, 17-23 X 2.5-3.5 n.
THE FUNGI \MI1CH CAUSE PLANT DISEASE

It is the cause of a drj' rot of apples.**^'
^^^
645
V. dianthi (Hal.) Atk.-i It is the cause of anthracnose of carnation.
The
acervuli are conspicuous with black setse.

is
V. concentrica Hals,
reported by Halsted as the cause of leaf
spots of Bletia.-^
Tuberculariaceae-Mucedineae-Phragmosporae
colored, fusoid to falcate, rarely short
of Fusarium.
(p.
G38)
Hyphffi hyaline; conidia 2 to several-septate, hyaline or bright-
and simple
in
some
species
Key
to Genera of Tuberculariaceae-Mucedineae-Phragmosporae
..
.
Conidia somewhat catcnulate, cylindric. Conidia rarely catenulate

gelatinous
1.
DiscocoUa.
Conidia cruciately 4-celled; sporodochium

2.
Sarcinodochium.
Conidia not cruciate

Conidiophores short, simple Conidia very large, terete-oblong.
Conidia doliform
3. 4.
Bactridium.
Pithomyces.
Conidiophores more or
key-like
less
branched
5.
Conidiophores dichotomous; conidia
Heliscus.
Conidiophores usually verticillately

branched,
cate,
conidia usually
fal-
sometimes oblong Sporodochium gelatinous
6. 7.
Pionnotes,
p. 645.
p. 646.
Sporodochium waxy
or byssoid ...
Fusarium,
Pionnotes Fries
firm, orange, pulvinate or lobed; or branched; conidia rather large, conidiophores fasciculate, simple fu.soid to cylindric, curved.
Sporodochium gelatinous, then
Only twelve or fifteen species, chiefly saprophytes. P. betae Mas. occurs on mangels and beets and according to Massee is probably identical with P. rhizophila which attacks
stored Dahlia roots and potatoes.
646

Fusarium Link
(p.
645)
Sporodochium pulvinate, or subeffused; conidiophores branched; conidia terminal, solitary, fusiform or falcate, more or less curved,
pluriseptate.
In part=Nectria, Neocosmospora,
205, 206.
Gibberella.
See pp. 201,
This
is
a large genus, (some four hundred species have been dewill
scribed) though future study names to synonymy.
undoubtedly relegate
many
Many of the species

of plants
and by stoppage
are destructive parasites, invading the ducts of the water-supply causing the class of
diseases
etc.
known as "wilts." Others induce rot, spotting, cankers, Taken as a whole the genus is one of the most injurious with
live normally as encroach upon living roots of susceptible
which plant pathology has to do. It seems probable that some of the forms that
saprophytes in
plants
soil
may
when these
are available.
In nature the spores typical of this form-genus are borne in sporodochia, coremia or acervuli and are crescent-shaped or fusoid. The same mycelium that produces these structures often, indeed
and smaller conidia scattered (=Cephalosporium). These two forms are called macroconidia and microconidia respectively. The microconidia are regarded by Appel & Wollenwelêr ^^^ as depauperate macroconidia. Frequently chlamydospores form in the mycelium; either terminal or intercalary. Sclerotia are also not uncommon. Undoubted species of Fusarium have been shown to belong to several different Hypocrealous ascomycetes, while still more have as yet revealed no ascomycete connection.
usually,
produces
also
similar
on
single hyphse
Biologic specialization has been found, in that forms morphologically indistinguishable are frequently incapable of cross inoculation onto other than their usual hosts.
Fusarium grows marked differences
well in culture
in
and the species often show on various media, particularly in growth

is
the colors that are developed. As with the anthracnoses much study
here needed to throw
647
/
/
648
light
on the inter-relation of the various species and their hosts. ^*^^ have made an extensive study of several Apple & Wollenweber
ground for a monograph. They conclude that important characters are the forms of the conidia, especially their bases, apices, and degree of curvature and septation (see Fig. 436); the color of the mycelium and spores;
species to lay the
in delimiting species
the presence or absence of chlamydospores.

valueless
many
earlier descriptions
substituting
They cast aside as new diagnoses
and new names. According names should stand.
to their conception, the following
commutatum
F. solani (Mart.) Sacc. Sacc.
=Fusisporium solani Mart. =Fusarium
F. martii A.
& W.
=Fusisporium solani Mart.
F. coeruleum Lib.
F.
F.
=Fusarium solani. discolor A. & W. =Fusarium solani. rubiginosum A. & W. =Fusarium solani.
sulphureum
(Schl.)
F. discolor var.
A.
& W. =Fusarium
sul-
phureum
F.
F.
Schlecht.
subulatum A.
& W.
not Schl.
metachroum A. & W.
F. orthoceras A.
F. theobromae A. F.
F.
F.
& W. =F. oxysporum Sm. & Sw. & Struk. wilkom.mii Lin. =F. bacilligerum B. & Br. falcatum A. & W. =F. vasinfectum pisi Schk. gibbosum A. & W.
It will
be noted that several of the species mentioned below
are here involved.

F. platani
Mont. =Calonectria pyrochroa.

F. rubi Wint.
See
p. 205.
Mycelium white, becoming pink, especially abundant on the flowers; conidia elongate, 1 to 8-septate, variable in size and form,
_ _ ^ FiG. 437. Section
.
of
straight or curved, 14-30 x 3-3.5 ^' not conu,

'
f"
ovary showing my- stricted.

pe/ ami ovdie.*^\fter
Coolc
^^^' ^^^
ound
this
fungus in diseased
double-
^^-
buds
of dewberries
and by inoculation dem-
onstrated that it is responsible for witches-broom, blossom, and similar abnormal growths of this plant.

F.
649
described by Aderhold ^' as fatal to flower buds of cherry before they open, a conclusion supported by inoculation experiments. The disease in general appearance re-
gemmiperda Aderh.
is
sembles sclerotiniose.
F.
rhizogenum
P.
&
C.
ram. wide, dense, convex, white or whitish, hyphae densely interwoven, septate, subramose; conidia
Sporodochia
superficial, 1 to 2
oblong, roundish, 1-septate, 70 x 4 /i. It was originally described as a parasite on apple roots in Nebraska ^^' and is mentioned by Aderhold ^ as the cause of death of
roots of apple and cherry trees in Europe. The mycelium grows within the roots and gummosis of the wood occurs. A Cephalo-
sporium form
is
known,
also chlamj^dospores.
is
F. putrefaciens Osterw.^^^'^^^ decay of pomaceous fruits. F.
said
by Osterwalder
^^^
to cause
cubense E.
F.
Sm. was
isolated
from bananas affected with
Inoculation showed the fungus capable of growing through the bundles for long distances.^^^
blight.
F. limonis Bri.^' 3".
437
Sporodochia gregarious, confluent, white; hyphae spreading, branched, septate; conidiophores erect, with alternate or opposite branches; conidia variable, acrogenous, continuous to 3-septate,
oblong to fusiform, curved, pointed, slightly constricted, 26-27 x
2.4-2.8
fx.
This fungus
the
fruits
is
Mal-di-gomma
held to be contributory to, or foot-rot of citrous
if
not responsible
for,
which is known practically wherever these fruits are cultivated.

F.
culmorum (W. Sm.)
Sacc^^"-
385
Reddish-yellow, gelatinous, effuse; hyphse few-septate, tortuous; fertile, short, continuous; conidia fusoid-falcate, 3 to 5-septate,
28-32 X 6-8 n on wheat.

affects chaff
Fig.
438. F. culmorum.
After Chester,
The fungus
and
seed,
first
appearing as a whitening of the upper halves of the glumes followed later by a pink color. The glumes become cemented together and the whole head may be involved. The grains are of
light
weight and are often covered with the fungus.
Chester
650
showed that the mycelium penetrates the seed and may even consume it entirely. F. sp. occurs on raspberry.^^^
F. moniliforme Shel.
salmon-pink; conidiophores simple or with opposite branches; microconidia continuous, oblong-ovoid,
Sporodochium subeffuse,
Fig. 439. Fuârium on corn. After

Burrill
rett.
and
Bar-
F. vasinfectum. A. MacroFig. 440. conidia. B. Portion of a hypha. C. A germinating macroconidium.
After Reed.
moniliform, 6-10 n long; macroconidia falcate, acute,

3-septate, 25-40 n long. It causes molding of corn.^"
usually
Several other undetermined species have been isolated from
corn on which they occur as the cause of dry rot of the grain. ^^ A fusarium on banana is by Essed referred to Ustilaginoidella. See p. 214.
F. vasinfectum Atk.^^"^^^
Hyphse
at maturity yellowish, 2-4 n in diameter; conidia borne

singly;
microconidia
oval,
con-
tinuous; macroconidia falcate, 2 to 3-septate, 1-2 x 2-4 /x.
Fig. 441. F. vasinfectum. D. MiE. Chlamydospores. croconidia.
Atkinson on cotton
i
.
^^^
first
described this
Plants
it
-i
and okra in which r j the WaS found plugging i.i,

i
After Reed.
ducts
in
mycelium here was 2-4 n

seen within the ducts.
tions with these
with its mycelium. The diameter and microconidia were
Pure cultures were obtained and inoculaon plants already injured by Pythium resulted in

ascigerous stage was said Neocosmospora and the many wilts caused various authors who follow
infection.
651
The
^^ to be a by Smith by Fusarium have l)y
Smith
been
reported
as
Neocosmospora
though
without real evidence that
they are such. See

page 205. Recent studies of Hig-
and Butler ingins dicate that the Fusarium

of
^^^
^^^
the
Neocosmospora
is
a saprophyte and that the Fusariums parasitic in the

wilt
diseases
in
are
as
yet
unknown
form.
ascigerous
The Fusarium parasitic on cotton is believed to Fig. ,. .J -.i.ij. be identical with that on okra but distinct biologically
1 ,
442.
F.
of veins.
if
vasinfectum, showing thrombosis After Atkinson.
not morphologically from that
of
watermelon.
F. vasinfectum var. tracheiphila E. F. Sm.^^^ This form on cowpea, which appears to be morphologically identical with F. vasinfectum is not capable of infecting cotton. F.
wilt.
niveum E.
F. Sm.^^'
it is
^^-'^^'*
is
the cause of the watermelon

as identical with this
Morphologically fungus regarded by Reed
like F.
vasinfectum.
^^' ^^^
was
also
described as causing wilt of ginseng. F. vasinfectum var. pisi v. Hall has been described as a variety ^^^ affecting the pea.^^^'
F.
udum
Butler on pigeon pea in India

is
is
closely related to
this last fungus.
F. aurantiacum (Lk.) Sacc. on stems, leaves and fruits. F. oxysporum Schl."''^-^' ^^^
recorded for cucurbs occurring
Sporodochia convex, subverrucose,
rose,
erumpent, confluent;
b52
conidia on short conidiophores; microconidia continuous, elliptic; macroconidia falcate-fusoid, 3 to 4-septate, 40-60 x 7-8 mSmith and Swingle ^^^ mention 11 described species of Fusarium
recorded by Saccardo for the Irish potato,
viz.
Fusarium oxysporum
Sacc,
F.
Schl.,
F.
(Fusisporium) solani (Mart.)

Mart.,
F.
(Fusisporium)
solani-tuberosa
didymum
Harting, F. solani Schl., F. (Fusisporium) roseolum (B. &. B.) Sacc, F. violaceum Fcl., F. ca^ruleum (Lib.) Sacc, F. diplosporum
C.
&
E., F.
commutatum Sacc,
F. pestis Sorauer, F. seruginosum
Delacroix, F. acuminatum E. & E., F. affine Fautr. & Lamb, all of which they tentatively regard as synonyms, attributing such
differences as
have been noted in descriptions to variations in the environment under which the fungus was growing when described.
disease caused
is
The potato
common
over a considerable portion
and is variously known as "bundle blackening," "stem rot," "dry end rot," and "dry rot." The fungus grows readily on manj^ culture media, showing large variation with the environment. It is aerobic and tolerates large amounts of malic, citric and tartaric acids.
of the United States
F. acuminatum E. & E. Sporodochia gregarious, minute, whitish or flesh-colored; conidia falcate, attenate, 3 to 5 or 6-septate, not constricted.
Described by Stewart New York.
^^^
as causing a girdling of potato stems in
F, roseum-lupini-alba Sacc.
Sporodochia pulvinate, minute, confluent, cinnabarine; conidiophores variable, long, slender, branched, branches nodulose, fusoid; conidia fusoid falcate, 45-55 x 4 ^t, 4 to 6-septate. It
causes spots on leaves and pods of lupines and attacks the seeds,
inducing
rot.
is
F. cucurbitariae Sacc.
on cucumbers
in Queensland.
F. solani (Mart.) Sacc.
Sporodochia globose, irregular, white; conidiophores branched; conidia fusoid-falcate, 3 to 5-septate, 40-60 x 7-8 n, subhyaline.
Clinton,^^^ also
Wehmer
^^^
and
others,
have shown
It
this to
be
the cause of "dry end rot tical with F. oxysporum.

F. pestis Sor.
is
"
of stored potatoes.
may
be iden-
given by Sorauer as the cause of "black-leg"

(see p.
653
F.
46)
of potatoes;
it
is
oxy-
sporum. F. erubescens A.
&
"'''"
v.
Ov. produces small black sunken spots
on green and
toes
in
ripe
tomare-
Germany
sulting finally in
fication.
mummiby was
Parasitism
of
means
enzymes
demonstrated.
F. lycopersici Sacc.^^"''-
Sporodochia as in F. oxy-
sporum; conidia falcate, acute, 25-30 x 3.5-4 fi,

hyaline to yellowish. It is the cause of a towilt or "sleeping After Bolley. Fig. 443. F. liiii. disease" resulting from invasion of the ducts. Conidia of two kinds are produced, Fusarium and Diplocladium. Infection is subterranean.
mato
nearly related disease differing chiefly in the fact that the fungus does not reach far above ground has been described by
Smith.'"'-
The fungus
in
both cases
is

F.
oxysporum.
F. lini
Boll.'*3
erumcream pent, compact,

Sporodochia
to
flesh-colored
;
co-
nidiophores
nidia
JîG. 444.
short,
cofusi-
much-branched;
3-septate,
F. lini, sketch, showing the mode of attack upon a voung root tip of a seedhng flax plant. After Bolley.
form, slightly
to
falcate,
n.
curved 27-38 x
3-3.5
A serious widespread
flax wilt is caused.
The mj^celium develops
luxuriantly from bits of diseased stem laid in sterile Petri dishes and grows well in culture media. Normally a soil saprophyte, it
invades the roots, grows through the veins, plugs the ducts and causes death. The sporodochia are found abundantly on the bases
654

The
seeds.
of diseased plants.
particularly on the
spores abound on all diseased parts, Infection experiments have demonsaid to cause a
rot of
strated
F.
its
pathogenicity.
is
tabacivorum Del.
is
tobacco in
France.
economic importance on cabbage. ^^ Inoculations of an undetermined species of Fusarium in pure

F. brassicae Thiim.
of
culture into soil also resulted in infection of
83%
of the
cabbage
plants grown therein. F. decemcellulare Brick and F. theobromae Lutz. occur on

cacao.
F. ricini (Ber.) Bizz. F.
is
injurious to the castor
oil
plant.
incarnatum (Desm.) Sacc. is reported as the probable cause of an aster wilt or blight in Europe. '^"^ An undetermined ^' ^^ and species is also reported on China aster by Galloway
others.
species of Fusarium on carnation leaves following in rust was reported by Stewart ^^ and a wilt disease or stem rot of carnation was studied by Sturgis.^"^ He found the Fusarium in the affected plants, it was isolated and inoculated into the soil around the roots of carnations producing disease in several insori
stances. F. pelargonii Crou. is described from geraniums.^^ F. dianthi P. & D.'*^^ on Dianthus cuttings, is a wound parasite, following insect injury. F. violae Wolf.
Infected areas dark, sunken; sporodochia within the host; conidia fusiform-falcate, 28-38 x 4-6 ii, 3 to 5 times septate; hyphse hyaline, 4-7 /x in diameter, irregularly branched. It causes
a disease of roots and stems of pans3^ F. pini is believed to be the species responsible for a disease of
pine seedlings.'*^F. blasticola Host, causes death of conifer seedlings in Europe.
Tuberculariaceae-Dematiae-Ainerosporae
(p. .638)
Hyphse olive to brown or black; conidia continuous, rarely hyaline globose to elongate, sometimes unequal.
655
KIey to
Genera of Tuberculariaceae-Dematiae-Amerosporae

Sporodochia not setose
Conidiophorcs lacking
Lichenicolous
1.
Spilomium.
Not
lichenicolous
Sporodochia
gelatinous;
conidia
2.
globose, vesiculosa
Myriophysa.
Sporodochia not gelatinous Sporodochia hemispheric, with a stratum of conidia

Sporodochia
ate
disk-like,
3.
Spermodermia.
Sclerodiscus.
p. 656.
applan4.
Conidiophorcs present
Sporodochia thick, tremelloid Sporodochia not tremelloid with a slender Conidiophorcs

apical appendage;
5.
Epidochium,
conidia glo6.
bose
Bonplandiella.
Conidiophorcs not appendaged Conidia globose
Sporodochia
layers
cellular,
uniform
7.
Epicoccum,
Triplicaria.
p. 656.
Sporodochia of three hyphal

8.
Conidia ovoid to bacillar
Conidiophorcs bacillar; sporodochia subdiscoid

9.
Hymenopsis.
No brown
base
radiate hyphse at
10.
Strumella,
p. 656.
Brown
base
radiate
hyphae at
11.
Astrodochilum.
Sporodochia ciliate or with exserted hyphse Sporodochia with loose exserted conidiophores, verruciform
12.
Trichostroma.
Sporodochia
setse
margins
with
hairs
or
Seta?
dark
13.
Chaetostroma,
p. 656.
Setai or hairs white
14.
Myrothecium.
656
Conidia in chains
Conidiophores lacking
15.
Exosporina,
p, 656.
Conidiophores present
Sporodochia globose
16.
17.
Sphaeromyces.
Sporodochia
stellate
Actinomma.
Epidochium
Fries (p. 655)
Sporodochium thick, tremelloid, subglobose or wart-form, black or pallid, erumpent; sporophores filiform, equal or apicallj^ swollen; conidia ovoid, oblong or pyriform, solitary or catenulate.
Some
fifteen species.
is
E. oryzae Miy.
found
on
rice.
Epicoccum Link
j^^
(p.
655)
^wHU ^P
'^'^!^'-'^^
Sporodochia globose or convex, cellular, dark; conidiophores very short; conidia globose.
Some
fifty species.
is
SCif^^^2l'-^'S
ef^^^^fii^
^' hyolopes Miy.
on
rice.
1^
.
Strumella Saccardo
(p.
655)
'""-^Ji!^^
-,
..^ iHG. 445.
^ Epicoccum.
Sporodochia wart-shaped; conidiophores branched conidia ovate, often somewhat bent.

; '
After Saccardo.
Some
is
fifteen species.
S. sacchari Cke.
found on sugar cane.^^^
Chaetostroma Corda
(p.
655)
Sori dark or cushion-form black bordered with black hairs;
spores elliptical, fusiform or rarely almost spherical. C. buxi Corda on Box=Nectria rousseliana. See p. 204. C. cliviae Oud. causes blotches on Clivia.
Exosporina Oudemaus
Sporodochia
erumpent;
conidia
catenulate,
homomorphic,
continuous, greenish. Monotypic. E. laricis Oud. is parasitic on larch leaves in Europe.
G57
Tuberculariaciae-Dematieae-Phragmosporae
(p.
639)
Hyphae dark; conidia usually colored, 2 to several-septate, oblong to cylindric.
Key to Genera of Tuberculariaceae-Dematiae-Phragmosporae.

Conidia in chains; sporodochium discoid. Conidia not in chains Conidia 1-cihate at each end
1.
Trimmatostroma,
Ciliofusarium.
p. 657.
2.
Conidia muticate
Sporodochium hairy Sporodochium smooth

Conidia laterally proliferate and joined in bundles Conidia
3.
Excipularia.
4.
Amallospora.
not
proliferate
and
.
united
Sporodochia convex-pulvinate.
or clavate
5.
Exosporium,
p. 658.
Sporodochia vertically cylindric

6.
Listeromyces.
Trimmotostroma Corda
Sporodochia pulvinate, compact, bearing a layer of conidiophores; conidia oblong, often curved, 2 to 8-septate, catenulate
brown.
genus of a half dozen species.
T. abietina Doh.î''
perennial; sporodochia foliicolus or caulicolus, conidiophores subhyaline, or tinged with olive-browTi, 4.5 X 20-30 fx, septate, sparsely branched, bearing the conidia
Mycelium
diffuse;
terminally;
conidia catenulate, very variable, dark olivaceous-
brown, slightly roughened, usually oblong, spherical, straight or inequilateral, continuous, spherical, 5 n, or 2 to 5-celled and 5-6
X 8-16
n,
not constricted, rarely muriform, 5 x 10

firs
^t.
On
white and balsam
the mycelium makes are produced till the second year after infection the more readily overlooked.
perennial habit of the pest a persistent one and as no conidia

in
its
Canada.
The
presence
is
658
Exposorium Link
(p.
657)
Sporodochia convex, compact; conidiophores dark, simple, densely compacted; conidia single, oblong to cylindric, pluriseptate.
Some
See pp. 227, 236. E. juniperinum (E.) Jacz. = Coryneum juniperinum. See p. 236. E. laricinum Mas. is found on living larch twigs.
E.
tiliae
twenty-five species. In part = Coleroa and Coryneum.
Lk. grows on young shoots of Tilia.
E. palmivorum Sacc.^^^
Fig. 446.
E. palmivorum.
5, spores.
3, a sporodochium, After Trelease.
Spots amphigenous, minute, suborbicular, 1-3
mm.
in diameter,
sporodochia superficial, densely gregarious, punctiform, black; 30 x 60-80 ju; conidiophores oblong, continuous, reddish olive, 5-6 x 14-16 /x, conidia borne singly,
scattered;
fusoid, straight or curved, apically obtuse or acute, basally obtuse,
brown,
8 to 10-septate, not constricted, olive-brown, ends paler, 8-9 x
80-90
fx.
On
palms, especially species of Phoenix in America. E. presii Bub. on species of Phcenix in Europe is very similar
to the preceding species. In the Tuber culariacese-Dematieae-Dictyosporse Th3nrococcum sirakoffi Bubak forms black tubercles under bark of mulberry
and
kills
the twigs.^*^
659
Mycelia-Sterilia
(p.
479)
Numerous forms are known merely us sterile myeelia. They or may not make sclerotia. In several instances these sterile forms are so aggressive as to warrant classing them among the worst of plant pathogens. Until more is known of them it becomes necessary to arrange and name them, for convenience of reference,
may
in a purely artificial
manner.
Mycelia-Sterilia.
Key to form Genera of

Tubercle-like
Tubercles connected with Tubercles without

fibrils
fibrils
1.
Rhizoctonia,
Acinula.
p. 659.
Cortex discrete Cortex not discrete
2.
3.
Sclerotium,
p. 660.
Maculiform Black stromata in leaves and stems Pseudo stromata in cortex

Root-like
4.
5.
Ectostroma.
Phellomyces.
Filaments rigid, broad, terete or depressed, dark, white within Filaments

rigid, capilliform,
6.
Rhizomorpha.
Capillaria.
dark, closely
7.
adhering Clavariform; filaments terete, vertical, simple or branched
8.
Anthina.
Cobwebby
or byssoid
Cespitose
interwoven, joined in bundles

black
soft,
primary
hyphse
9.
Ozonium,
p. 661.
Cespitose interwoven, hyphse not fasciculate,
10.
Rhacodium.
evanescent, white or pale Adpresscd, creeping, dendritic, while to brownish, not forming a continuous
Cobwebby,
11.
Hypha.
membrane
Membrane-like; densely interwoven, forming a continuous subcrose or coriaceous
12.
Himantia.
membrane
Rhizoctonia
Sclerotia variable
in
13.
Xylostroma,
p. 663.
De CandoUe
homj^-fleshy
;
form,
cortex thin,
mem-
660
branous, persistent, inseparable; formed among and connected by the mycelial threads. There are about a dozen so-called species, some of them very
important plant pathogens. See pp. 407, 408. R. betae Kiihn and R. solani Kiihn =Corticium vagum solani, as does also part of what has been referred to as R. violacea. R. medicaginis D. C. (Tul.);i^ (see also ^^^' ''^-'''). Hyphae subtomentose, on the cambium of the host, forming a
membrane
or fasciculate strands, covering the host in time with a
violet coating; sclerotia reddish-violet.
On
cium
alfalfa in
Europe and America.^^^

has studied this form and the form allied to Corti-
Duggar who
(pp. 407, 408) regards the
who has also "The fungus appears upon

hyphae composed of
cells
two as distinct though Giissow ^- studied both pronounces them the same. Duggar says,
the root as a close weft of violet-colored
less
more or
uniform in diameter.
]\Ior-
phologically it bears no resemblance to the sterile stage of Corticium." This form is found on alfalfa, asparagus, beet, and possibly other plants.
Leptosphseria has been reported as its ascigerous stage though the evidence of such connection is not conclusive.
R. crocorum D. C.
is
a form which
kills
the corms of saffron.
R. strobi Scholz
injuring
is
the
name given to
a form described as seriously
young pine
trees in Austria.
^"^
R. subepigea Ber. is destructive on the roots of coffee. '*^^ A Rhizoctonia of undetermined species has been found on buck-
wheat in the United
States."*-^
Sclerotium Tode
(p.
659)
Sclerotia roundish or irregular in form, cartilaginous-fleshy, not connected by mycelial threads; cortex thin, membranous, in-
separable.
Over 200
species
have been described.

5. 438
S. rolfsii Sacc.2i'
Sclerotia small, brown, about the size of a mustard seed. This sterile fimgus possesses a very aggressive mycelium which under favorable conditions of moisture grows on almost anything living or dead, producing a dense white cotton-like mass of threads.

Soon the
sclerotia
661
form as mustard-secd-liko bodies. They are abundance on all media but neither these structures nor the mj^celium have yet been seen to bear spores of any kind. The fungus was first studied l:)y Ilalsted ^^ and later by many others. It was described and named by Saccardo from specimens communicated by Stevens.
produced
in great
cepivorum Berk. Minute, spherical, gregarious, black.

S.
It
is
found on various
species of Allium, causing rot.
Fig. 447.
-S. rolfsii,
sclerotia.
After Halsted.
S. rhizoides Auer.^-^
Subglobose, at
first white-villose,
then smooth, black, rugose.

It causes considerable
Calamagrostis and other grasses. injury to the hay crop in Europe.
On
S. tuliparum Klebahn,'*-^ S. tulipae Lib. and S. bulborum Wak. are found on tulips, and other bulbs. A relation to Sclerotinia is
usually assumed but has not been demonstrated. See p. 136. S. oryzae Catt. is found on rice in Japan and Italy.
Ozonium
Link.
(p.
659)
or byssoid, cespitosc, primary hyphse fasciculate.
Cobwebby
hyphse densely interwoven,
Some twelve
species.
662

Sh.'
O. omnivorum
^-^'^^
Mycelium
dirty yellow; sometimes whitish
when young, growforming strands
ing in the vascular bundles of the host; hyphsB
and spreading from them, producing a rather dense arachnoid layer on the surface of the host and bearing 1 to 4 branches arising
1'
IG.
448.
S. rolfsii, sterile
mycelium growing on
carrot.
After Stevens and Hall.
and growing
3 to 5
at right angles from the same point near the ends, in diameter, tapering toward the ends. /x It causes root rot on almost any kind of plant including among its hosts a large variety of trees. The first description was by
Pammel
in a
Texas Bulletin; a
later
one was by Shear.
The fungus
destroys the smaller rootlets, cortex of older roots and invades the vascular system and medullary rays, resulting in wilt and death.
It
may
superficially.
be seen as dirty yellowish strands or as a thin weft Sclerotia-like bodies appear on the roots often at

lenticels.
663
mycelium is not typically are hyaline. The fungus was in early studies difficult to isolate but Atkinson '-^ in 1893 obtained pure cultures by rinsing the diseased roots in
associated to form strands and
its cells
Inside of the host tissue the
distilled water, cutting in small pieces

sterile
and placing on
sterile filter
sand in a moist chamber. In a few days the paper lying on strands grew over the paper onto sterilized slides. Bits of sterilized cotton-root were then placed in contact with the advancing hypha. Soon the new culture thus secured could be transferred at will.
slight acidity retards bacterial
growth and renders
isolation of
the
Ozonium
less difficult.
In culture sclerotia about 3
mm.
in
diameter, whitish and woolly, later brown, appear.
Xylostroma Tode
This occurs, forming thick,
It is regarded as the
(p.
659)
cracks of timber.
es-
felt-like layers, in
mycelium
of various
Hymenomycetes,
pecially Fomes.
Fungi of Unknown
Affinity
The following imperfectly known genera do not fit readily into the scheme of classification and are all in need of careful study.
Acrocystis Ellis
;
&
Halsted
Monotypic though technically a nomen nudem, the illustrations arc recognizable. Examination of the original material shows Saccardo's reference to the Mucorales to be untenable.
A. batatae E.
&
H.^^
Hyphpe intercellular, branched, producing enlarged cysts at the ends of branches, the nature of these unknown; enlarged, intramycelial swellings contain numerous rounded conidia. It is described as the cause of soil rot of sweet potatoes.
Graphiola Poit
Mycelium within the host; fruiting body rotund, carbonous duplex, the outer layer of interwoven branched hyphae, firm, in-
66i
and sterile fasciculate hyphse; fertile hyphse with short branches bearing the spore-raother-cells which divide
terior softer, of fertile
into
two globose or angular spores; germination by a
filiform
conidia-bearing mycelium.
A genus
of seven species, chiefly on palms. G. phoenicis (Moug.) Poit.^^^' '^^ Sprodochiiim 1-1.5 x 500 n; exoperidium homy, black, inner
peridium membranous, hyaline; spore-mass yellow; spores globose or elliptic, 3-6 n, with a thick, smooth, hyaline wall.
On
the date and other palms throughout the world.
665
G. phoenicis. FiQ. 449. II, sporifcrous organ. Ill, section of the same. After Stone and Smith.
BIBLIOGRAPHY OF FUNGI IMPERFECTI*

Stevens, F. L. and Hall, J. G., Bot. Gaz. 45; ^Leininger, H., C. Bak. 29: 4, 1911.
I 3
1,
1909.
*
5
6
->
Cobb, N. A., D. Agr. New So. Wales, Miss. Pub. 666: 21. Hedgcock, G., Myc. 10: 2, 1904. Kirk, T. W., N. Z. R. 348, 1906. Potebnia, A., Ann. Myc. 8: 58, 1910.
Soott,
Scott,
Quaintance, A. C, F. B. 283: 14, 1907. " Clinton, G. P., 111. B. 69: 1902. 10 Stewart, F. C, N. Y. (Geneva) R. 1J^: 545, 1895.
II 12
W. M. and W. M. and
Rorer, J. B., B. P.
I.
B.
lU:
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13
Scott, W. M. and Rorer, J. B., B. P. I. B. 121: 1908. Stevens, F. L., N. C. B. 196: 54, 1907. Ann. Inst. Cent. Amp. Roy. Hong. 3: 167, 1905.
"
6
i
Selby, A. D., 0. B. 2U: 445, 1910. Briesig, Bui. Min. Ag. Ind. & Cen.
Rome,
1910.
Miyaki, Bot. Mag. Tokyo 23: 1909. " Stewart, F. C, N. Y. (Geneva) B. 328: 1911. i Halsted, B. D., N. J. R. 12: 279, 1891.
20
"7c?em. 19, i^.- 355, 1890. Halsted, B. D., N. J. B. 70:
9,
1890.
21
" "
2"
Halsted, B. D., N. J. R. 1I^: 355, 1893. Selby, A. D., 0. B. 105: 222, 1899. Patterson and Charles, B. P. I. B. 171.
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"
28
Humphrey,
J. E., Zeit. 3:
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360, 1893.
" Gueguin,
'
B., S.
Fr. 18: 312, 1902.
Aderhold. R., C. Bak.
"1
="2
6: 620, 1900. Trinchieri, R., Bui. Ort. Bot. R. Univ. Napoli 2: 409, 1909. U. S. D. Agr. R. 88.
"
Stewart, F.
C, (Geneva) R.
*
15: 456, 1896.
See footnote, page 53.
666
BIBLIOGRAPHY OF FUNGI IMPERFECTI

'*
667
^0
S. Dept. Agr. R. 87. Chester, F. D., Del. R. 5: 75, 1892. Selby, A. D., 0. B. 92: 233, 1898.
U.
" Lewis, C.
3
'
E., Sc. 31: 752, 1910. Lewis, C. E., Me. B. 170: 1909. Plalsted, B. D.,"N. J. B. 91: 1892.
40 41
McAlpine, Dept. Agric. jMclbourne, 132, 1899. McAlpine: Fungi of the Vine in AustraUa.
S.
Frank, Zcit. 3: 28, 1893. " PrilHeux and Delacroix, B.

44
4=^
My.
Fr. G: 178, 1890.
Rostrup,
E., Zeit. 4: 195, 1894.

F., Sc. 32: 726, 1910. F.,
4
47 4 43
Manns, T. Manns, T.
Mycologia
1: 28, 1911.
Bos. Zeit. 16: 257, 1906.
Quanjer, Zeit. 17: 259, 1907.
'
Rostrup, E., Zeit. 4: 322, 1894. Rostrup, E., Tid. f. Landok. R. 5:

Kirk, T. W.,
11, 330, 1891.

'
"
62
New
Zealand, D. Ag. R. 13: 410, 1905.
" "
Klebahn, H., Zeit. 20: 1, 1910. Fron, G., Bull. Trim. Soc. Myc. France 25: 66. Halsted, B. D., N. J. B. 76: 25, 1890.
Stewart, F.
C, N. Y. (Geneva)
B. 179, 1900.
" Arthur, "

68 6" 6
N. Y., (Geneva) R. 3: 383, 1884. Deidicke, H., Ann. Myc. 9: 8, 1911. Steward, F. C., Rolfs, F. M., and Hall, F. H., N. Y. B. 191: 298, 1900.
J. C.,
Cordlcy, A. B., Ore. Bui. 60: 1900.
Paddock, W., N. Y. (Geneva) B. 163: 203, 1899.

Zeit.
"Eriksson,
2
63
"4 "6
'
1:29, 1891.
Halsted, B. D. and Fairchild, D. G., J. AUescher, A., Zeit. 5: 276, 1895.
Myc.
7: 1891.
Reed, H.
S.,
Mo.
B. 69: 1905.
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Doherty,
Rept.
M. W., Bot. Gaz. 30: 400, Mo. Bot. Gard. 9: 159, 1898.
1900.
Sci. i4- 899, 1901.
^18
Duggar, B. M., N. Y. (Cornell) B. 163, 1899. Stone and Smith, Mass. R. 67: 1897.
Heald, F. D., Phytop. 1: 103, 1911.
Gussow, H.
T., Zeit. 16: 135, 1906.
S., Earle, Stout, A. B., Sc. 33: 156, 1911. "^ Klebahn, H., Jahr. d. Hamb. Wiss. Ans. S2.
Stevens, F. L. and Wilson, W. G., Sc. 33: 943, 1911. < Scholz, E., Verh. K. K. Zool. Bot. Ges. Wien, 47: 541. ^" Selby, A. D., 0. B. 92. <24 Bertoni, W. S., Rev. Agr. Cien. Apt. Paraguay i; 211. 5 F. Ala. B. lOS: 1900.
'
4: Pammel, H., Shear, C. L. and Miles, G. F., B. P. I. B. 102: 39, 1907. 3 Galloway, B. T., and Woods, A. F., Y. B. 248, 1896. "4 Stewart, F. C. and Blodgett, F. H., N. Y. (Geneva) B. 167: 283. "6 Johnston, T. H., Agr. Gaz. N. S. Wales 21: 563. " Stevens, F. L. and Hall, J. G., N. C. R. 31: 72, 1909. *" Swingle, W. T., and Webber, H. J., V. P. P. B. 8: 32, 1896. "8 Fulton, H. R., La. B. 101: 1908.
Shear, C. L., Bull. Tor. Bot. Club 34: 305, 1907. Atkinson, G. F., Bot. Gaz. IS: 16, 1893. " Pammel, L. H., Tex. R. 2: 61, 1889. i L. B. Tex. 1888.
<28
"6 Porto Rico R. 449: 1903. Arthur, J. C, N. Y. (Geneva) R. 4: 250, 1885.

"1
^2
677
Laubert, R., Gartcnflora 59: 409, 1910.

Chester, F. D. Del, 6: 111, 1893.
"'
'*'
Peck, N. Y. St. Mus. Nat. Hist. B. 157: 21. Bot. Gaz. 54: 231.
Cordlcy, Ore. Sta. Bienn. Crop. Kept.
^ Sc. 37: 638.
"
1913. <"
*^8
&
Hort. R. 1911-12, 1S7:
Stone, Ann, Myc. 10: 564, also Mclhus, Phytop. 3: 56. Stewart, F. C. N. Y. (Geneva) B. 328, 387. "'Ann. Myc. 10: 491.
^s"
Gussow, Canada Exp. Farms. R. 1912, 202. Ann. Myc. G: 112. ^"Peck, N. Y. Mus. R. 32: 41.
^s'
"'Chester, F. D., Del. B. 70: 13.
Gussow,
Zeit. 16: 10.
*"
Heald, F. D., Mycol. 1: 215, 1909. Wehnier, C, C. Bak. 3: 646, 1897. ' Percival, J., Jour. Southeast Agr. Col. Wye, 81: 1902. 8 Chifflat, J., Jour. Soc. Nat. Hort. France, Ser. 4, 8, 348. Arthur, J. C, Ind. B. 19: 5, 1889.
SOME OF THE MOST USEFUL BOOKS

1.
Buller Researches on Fungi.
2.
A.
3.
Biology of the Fungi MyceClarendon Press, 1887. G. Delacroix and A. Maublanc: Maladies des Plantes Cultiv6es.
tozoa
De Bary: Comparative Morphology &
&
Bacteria.
J.
4.
5.
B. BaiUiere
&
Fils, 1909.
Ducomet: Pathologie V6g6tale, Chas. Amat, 1908. B. M. Duggar, Fungous Diseases of Plants. Ginn
A.
Engler:
1907.
&
Co., 1909.
6.
Syllabus
der
Pflan/;onfamihen,
Gebriider Borntraeger,
T.
Engler and Prant: Naturliche Pflanzenfamilien,

1897.
Wm.
Engelmann,
8.
W.
9.
G. Farlow: Bibliographical Index to N. American Fungi, Carnegie Inst, of Washington, 1905. Farlow & Seymour: Host Index of the Fungi of the U. S., Cambridge,
1888.
10. A. B.
Frank: Die PilzparasitSren Krankheiten der Pflanzen.
Ed.
11.
12.
Trewendt, 1896. E. M. Freeman: Minnesota Plant Diseases, Pioneer Press, 1905. K. Goebel: Outlines of Classification & Special Morphology of Plants, Clarendon Press, 1887.
R. Hartig: The Diseases of Trees. 14. R. Hartig: Lehrbuch d. Pflanzenkrankheiten, Julius Springer, 1900. 15. Ideta: Text-book of Plant Diseases, Japanese.
13.
16. 0.
Kirchner: Die Krankheiten und Beschadigungen unserer landwirtschaftlichen Kulturpflanzen.
Eugen Ulmer,
1906.
17.
Ernst
Kiister:
Pathologische
d.
Pflanzenanatomie,
Gustav Fischer,
1903.
18.
Franz Lafar: Handbuch

1904-1907.
Technischen Mykologie, Gustav Fischer,

d.
19.
Lindau
in
P.
Sorauer:
Handbuch
Pflanzenkrankheiten.
Paul
Parey, 1908.
20.
21. Lotsy:
Lindau and Sydow, Thesaurus Literaturse Mycologicse. Vortrage liber botanische Stammesgeschichte, Gustav Fischer,
1907.
678
SOME OF THE MOST USEFUL BOOKS

22. Geo.
679
Massee: Diseases of Cultivated Plants and Trees.

Massee: Text-book of Fungi, Duckworth Maladies des plantes agricoles.
Duckworth
<fe
Co., 1910.
23. Geo.
&
Co., 1906.
24. E. Prillieux:
26. P. A. Saccardo: Sylloge
Fungorum, Pa via; R. Fricdlander
&
Sohn
1882.
26. Stevens 27.
28.
and Hall: Diseases of Economic Plants, Macmillan Co., 1910. Tubcuf and Smith: Diseases of Plants Induced by Cryptogamic Parasites, Longmans, Green & Co., 1897. L. M. Underwood: Moulds, Mildews and Mushrooms. Henry Holt &
Co., 1899.
29.
30.
31.
H. Marshall Ward: Disease in Plants. Macmillan & Co., 1901. Strasburgcr, Noll, Schenck and Karsten: Text-book of Botany, 1908. F. D. Chester: Manual of Determinative Bacteriology. Macmillan &
Co., 1901.
32.
H.
W. Conn:
1909.
Agricultural Bacteriology.
P. Blakiston's
Son
&
Co.,
33. A. Fischer: 34. C. Flugge:
35.
36.
Vorlesungen ueber Bakterien, Gustav Fischer, 1903. Die Mikroorganismen. F. C. W. Vogel, 1896. E. 0. Jordan: General Bacteriology. W. B. Saunders Co., 1908. J. G. Lipman: Bacteria in Relation to Country Life. Macmillan
Co., 1908.
&
37.
38.
W. Migula: System der

Miquel
et
Bakterien. Gustav Fischer, 1897. Cambier: Traits de Bacteriologie Pure et Appliquê.
Masson
39. E. F.
et Cie, 1902.
Smith: Bacteria in Relation to Plant Diseases.
Carnegie In-
stitution, Sept., 1905.
41. Geo.
The Mycetozoa, 1895. Massee: Monograph of the Myxogastres, 1892. 42. H. Macbride: North American Slime-Moulds. Macmillan, 1899. 43. Rostafinski: Sluzowce Monografia, 1875.
40. A. Lister: 44. Torrend:
Les Myxomyc^tes.
Brotevia 7:
5,
177;
pi.
1-9, also sepa-
rate, 1908.
45. Plowright, British Uredinese 46. Salmon,
&
Ustilaginese.
Monograph
of the Erysiphacese.
47. Clinton, G. P.,

48. Murrill,
North American Flora,
Ustilaginales.
etc.
49. Arthur, 50. Atkinson, G. F.,
W. A., North American Flora, Polyporaceae, J. C, North American Flora, Urediniales.

Mushrooms.
51. Clements, The Genera of Fungi. 52. Ellis and Everhart, North American Pyrenomycetes.
680

Kryptogamen Flora von Deutschland. Handbook of the Fungous Diseases
of Trees, Trans,
53. Rabenhorst,
54. Bancroft, K.,
of
West Indian
Plants, 1910.
55. Hartig,
The Diseases
by Sommerville.
56. McAlpine, Rusts of Australia. 57. Jaczewski, A. A., Phytopathology (Russian). 58. Noel Deer: Sugar Cane. Norman Rodger, Manchester, 1911.
59. 60. Sj'dow,
61.
Klebahn, H., Die Wirtwechselnden Rostpilze, 1894 Monographia Uredinearum, 1904.
Holway, E. W. D., North American Uredineae, 1905. 62. Seaver,, North American Flora, 3: Pt 1. 63. Viala, R., Les Maladies de la Vigne.
64. P. Hariot,
Les Uredinees, Paris, 1908.
Periodicals of use to the Phytopathologist

Phytopathology. Annales Mycologici.
Centralblatt fur Bakteriologie U. Paristenkunde II. Abt. HoUrung's Jahresbericht u. d. Gebeit der Pflanzenkrankheiten. Experiment Station Record.
Zeitschrift
fiir
Pflanzenkrankheiten.
Rivista di Patologia vegetale.

Hed^vigia.
Mycologia, formerly Journal of Mycology.

Practische Blatter
fiir
Pflanzenschutz.
Just's Botanischer Jahresbericht.
Bulletin Trimestriel de la Soci6ti6 Mycologique de France. For Bibliographies of special articles, see pages 53, 109, 288, 466, 666.
GLOSSARY
A, privative.
Acervulus
Acicular.
(i).
Signifying without. A small cluster, tuft of mycelium bearing spores. Slender or needle-shaped.
Acrogenous. Growing at the apex. Produced in a succession towards the apex. Adnate. Attached the whole length. .^ciospore. .^cidiospore. A spore formed in an aecium. .^cium (a), .^cidum (a). A special form of sorus in the Uridinales. Aerial. Living above the surface of the ground or water.
Acropetal.
Aerobic.
ASrobiotic.
.^thalioid.
Requiring oxygen. Like an sethalium.
iEthalium (a). A compound sporiferous body formed from a large combination of plasmodia. Agglutinated. Agglutinate. Glued together.
Aggregate. Collected together. Sausage-shaped, crescent-shaped with rounded ends. Alveola (ae). Cavities or pits on the surface. Alveolate. Pitted like
Aggregated.
Allantoid.
a honeycomb. Alveolar. Pertaining to or resembling Alveolae. Amoeboid. Like an amoeba, exhibiting creeping movement by pseudopodia.
Amorphous. Shapeless. Amphigenous. Growing

ticular surface.
all
round an object, not restricted to any par-
Ampulliform. Swollen out. Flask-shape. Anastomosing. Uniting and forming a network. Angiocarpous. Invested by some covering. Annulate. Ring-shaped or with a ring. Annulus. A ring-like portion of the ruptured marginal
expansion of the pileus. Anoderm. Without a skin.
veil, after
the
Antheridium (a). In fungi the male sexual organ. At the point of any structure. Apical. Apicula. A sharp and short, but not stiff point.
681
682
GLOSSARY
Showing apogamy.
(a).
An ascocarp in which the while the asci are maturing. Appendages. Processes of any kind.
Appendiculate. Furnished with appendages. Appiculate. Furnished with an appicula.
Apogamous. Apothecium
hymenium
lies
exposed
Lying flat for the whole length. Organs of attachment of germinating Approximate. Close together, but not united. Arachnoid. Like a cobweb. Arcuate. Curved. Areola (ae). A space marked out on a surface.
Appressed.
Appresoria.
Aristate.
parasites.
Awned.
Armilla.
bracelet^like
frill.
Armilla-fonn.
Ascigerous.
Armilla-like.
asci.
Bearing
Ascocarp.
sporocarp producing
asci.
Ascogenous. Producing asci. In ascomycetous fungi, the cell or group of cells ferAscogonium. tilized by a sexual act. Ascoma. Receptacle and hymenium of the larger fungi. Ascoplasm. Protoplasm of the ascus. Ascus (i). A large cell in the ascocarp in which spores are developed,
usually eight.
Aseptate. Without cross-divisions. Asexual. Destitute of male and female organs. Asperate. Rough with hairs or points
Attenuate.
Tapered.
Ear-shaped.
Auriform.
Autoecious.
parasite which runs
its
whole course on a single host.
Avellaneous.
Drab, hazel, hazel-nut-brown.
B
Bacillar.
Bacilliform.
of.
Rod- or club-shaped.
Basal.
At the base
Basidiospore.
Basidium.
Basipetal.
The
spore acrogenously abjointed upon a basidium. mother-cell from which spores are acrogenously
in the direction of the base.
abjointed.
Growth
Bay.
Reddish brown or chestnut color. Biogenous. Growing on living organisms.
GLOSSARY
Racemose. Botryose. BuUate. Blistered or puckered.
Byssoid.
Fla.x-likc or cottony.
683
Calcareous.
Cancellate,
Capillitium.
Chalk-white, chalky.
Bell-shaped.
Latticed, as in Clathrus. Sterile thread-like tubes or fibers,
Campanulate.
mixed with the spores
within a sporangium. Capitate. Having a head.
Carbonous.
Carbonaceous.
(a).
Dark
colored.
Consisting chiefly of sub-
stances in which carbon predominates.
Carpogonium
Cartilagenous.
Part of a procarp resulting in a sporocarp after
fertilization.
Castaneous.
Catenulate.
a chain.
Hard and tough. Chestnut-colored.

Formed
of parts united or linked
Concatenate.
as in
Caulicolous.
Living on stems.
Cespitose. Growing in tufts. Chlamydospore. A spore having a very thick membrance. Chromogenesis. Color production. Chromogenic. Chromogenous. Color-producing.
Ciliate.
Fringed with hairs.

(a).
Cilium
Vibratile whip-like processes zoospores and similar bodies move.
of
protoplasm
by which
Cinereous.
Circinate.
Cineraceous.
Circinnate.
Ashy.
Circumscissile.
Coiled into a ring or partially so. Dehiscing as if cut circularly around.
Cirrhose.
Clathrate.
Cirrose.
Latticed.
Cirrhous.
Having a
cirrhus or tendril.
Clavate.
Clypeate.
Clypeus.
Coenocyte.
Club-shaped, thickened towards the apex. Buckler or shield-shaped, having a clypeus. A buckler or shield-shaped tissue around the mouth of a
cell.
perithecium. A multinucleate
ColIai)sing.
CoUabent.
Colliculose.
With
little
round elevations.
Columella.
Sterile axile
body within a sporangium.
684
Columnar.
GLOSSARY
Having the form
of a column.
Compound. Similar parts aggregated into a common Con or Com. In Latin compounds signifying with.
Conchate.
Shell-shaped.
whole.
Concolorous. Of one color. Confluent. Blended into one.

Conglobate.
Conidiophore.
Conidiospore.
Collected into a ball.
A sporophore bearing a
Same
as conidium.
conidium.
Conidium
(a).
Dust-like
spores
like
usually produced
directly
from the
hyphte.
Conjugation. Union of two Connate. United.

Constricted.
gametes to form a zygote.
Context.
Drawn together; contracted. The flesh of a mushroom and the corresponding substance
in
other pileate fungi. Continuous. The reverse of interrupted. Convoluted. Convolute. Rolled round.
Coremium.
Coriaceous.
The name
Leathery.
of a genus of fungi, derived
from a Greek word
meaning broom.
Corneous.
Cortex.
Cortical.
Of horny texture. The bark or rind. The peridium

Relating to the cortex. Ribbed.
of Fungi.
Costate.
Crateriform.
Cristate.
Globet or cup-shaped.
Crested.
Cruciate.
Cross-shaped.
Crust-like.
Crustose.
Cuboid.
Cupulate.
Cuticle.
Resembling a cube.
With a
cupule.
like a small cup.
Cupuliform.
Cuticulate.
Shaped
The outermost
Having a
skin.
cuticle.
Cyme.
Cyst.
Cylindric. Cylindrical. Elongated, with a circular cross-section. Cluster of determinate or centrifugal type, especially a broad and flattened one.
sac or cavity.
Cystidium (a). Large, one-celled, sometimes inflated bodies, projecting beyond the basidia and paraphyses of the hymenium of Agarics. Cytolitic. A ferment which dissolves the cell-wall.
GLOSSARY
685
D
Deciduous.
Decumbent.
Decurrent.
Definite.
Falling in season. Reclining with the
summit ascending.
Running down.
Dehiscence.
Precise; of a certain number. The mode of opening.
Deliquescent. Dissolving or melting away. Dendritic. Having a branched appearance. Dendroid. Tree-like in form, or branching.
Minutely toothed. Sunk down, flattened. Depressed. Determinate. Definite.

Denticulate.
Di.
Two
or double.
Dichotomous.
Forked.
Dichotomy. Forking in pairs. Difform. Of double form, irregular.

Diffuse.
Widely or loosely spreading. Fingered: compound. As in the Horse Chestnut leaf. Dimidiate. Halved, as when half an organ is so much smaller than the
Digitate.
Disciform.
Discoid.
Disculate.
other as to seem wanting. Flat and circular.
Resembling a
disk.
Having a
disk.
Disjunctors.
Spindle-shaped cellulose connections between conidia.
Dissepiment. A partition. Dolifonn. Barrel-shaped.
E, Ex. Privative in Latin compounds. Echinulate. Having small prickles. Effuse. Expanded. Ellipsoid. Ellipsoidal. Elliptic. Like an
ellipse.
Embedded.
Surrounded in. Endogenous. Produced within. Endophyte. Growing inside another plant. Endophytic. As an endophyte. Endospores. Spores formed endogenously. Endozoic. Living inside an animal.
Entire.
With even margin.
686
GLOSSARY
Entomogenous. On insects. Enzyme. An unorganized or soluble" ferment. upon ". Epi. In Greek compounds to mean Epiphyllous. Growing on leaves.
Epispore.
Outer coat of a spore.
Epithecium.
Epixylous.
The
surface of the fructifying disk.
Growing on wood.
Breaking through. Soon disappearing.
Wart-like.
Erumpent.
Evanescent.
Excipuliform.
Exospore. The
outer covering of the spore.
Explanate. Spread out fiat. Exserted. Protruding beyond.
F
Facultative. Falcate.
Occasional, incidental as opposed to obligate.
Sickle-shaped. A little bimdle. Fasciclie,
Fasciculate.
In clusters or bundles.
Rust-colored.
Favoid.
Like a honeycomb.
Ferruginous. Femigenous. Ferrugineous. Fibrinous. Fibrillose. Furnished with fibers.
Filamentous. Of free hyphae which are at most loosely interwoven but without forming bodies of definite shape and outline.
Filiform.
Thread-shaped.
Fimbriate.
Fission.
With the margin bordered by long slender
processes.
Splitting.
Shaped as a fan. Limp, flabby. Flagellate. Provided with whip-like processes. Flagellum (a). Whip-like process of protoplasm of a swarnispore. Flavous. Nearly pure yellow.
Flabelliform.
Flaccid.
Succulent. Fleshy. Flexuose. Flexuous.

Flocci.
Bent alternately in opposite
directions, zigzag.
Locks hke
soft hair or wool.
Floccose.
Flocculent.
Bearing flocci. Diminutive of Floccose.
Fluorescence.
Fluorescent.
Foetid.
The property
of diminishing refrangibility.
Exhibiting fluorescence.
Fetid, stinking.
GLOSSARY
Foliar.
687
Leafy or
leaf-like.
On
leaf.
Foliicolous.
On
leaves.
Free.
Not adhering.
Living on fruit. Living on shrubs.
Fructicolous.
Fruticolous.
Fruticose.
Fugacious.
Fuligineus.
Shrubby. Soon perishing.

Fuliginous.
Sooty, or soot-colored.
Fulvous.
Furcate.
Yellow, tawny.
Fumaginous.
Smoky;
sooty.
Forked.
Fuscous. Dusky, too brown for a gray. Fusiform. Thick but tapering towards each end. Fusoid. Somewhat fusiform.
G
Gametangiixm (a). A differentiated cavity, which produces gametes. Gamete. A sexual protoplasmic body. Gemma (ae). A young bud. Gemmation. Budding.
Gemmiform. Bud-shaped. Gill. The plates or lamellae

Glabrous.
Gleba.
Globoid.
of
an Agaric.
in the Phallales.
Without
Rounded.
hair.
The
gelatinous spore
mass
Globose. Nearly spherical. Glomerate. Agglomerate, collected into heads. Granular. Composed of grains. Gregarious. Growing in company. Associated but not matted. Guttulate. Resernbling drops, with drops.
Gymnocarpous. Naked fruited. Gyrose. Curved backward and forward
in turn.
Hamate. Hooked at the tip. Haustorium (a). Special branch of a filamentous mycelium serving as an organ of attachment and suction.
Heteroecism.
Heteroecious.
Condition of a hetcroecious parasite. Passing its stages on more than one host.
Heterogamy.
With gametes not uniform.
688
GLOSSARY
Heteromorphic. Heteromorphous. Variation from normal structure, as having organs differing in length; dimorphic.
Hirtose.
Hirtus.
Bristly.
Hairy; hirsute.
fine
Hispid.
Hoary.
Host.
Hyaline.
Gray from
A plant which
pubescence. nourishes a parasite.

or clear non-granular portion of pro-
Colorless or translucent.
Hyaloplasm.
toplasm.
The hyaline matrix
Hymenium
(a).
An
aggregation of spore mother-cells in a continuous
layer on a sporophore.
Hymenophore. That part which bears the hymenium. Hypha (ae). The thread-like vegetative part of a fungus.
Hyphoid. Resembling hyphae. Hypertrophy. An abnormal enlargement Hypophyllous. Situated under a leaf.
of
an organ.
Hypopodium
Hypothallus.
(a).
The stalk or support. The marginal outgrowth
of
hyphae often strand-like,
from the
thallus.
Hypothecium. hymenium.
Hysterioid.
layer
of hyphal
tissue
immediately beneath the
Elongated boat-shaped, resembling the genus Hysterium.
I,
n, HI.
Symbols
for the stages of the rusts, see p. 324, 326.

tiles
Imbricate.
Overlapping as the
on a
roof.
Immersed.
Below the surface. Imperforate. Without an opening. Incrassation. Thickened growth. Indehiscent. Not opening along regular lines. Indeterminate. Not terminated definitely.
Indurate.
Innate.
Hardened.
Shaped like a funnel. Born on the apex of the support. Imbedded. Intercalary. Growth which is not apical but between the apex and the
Infundibuliform.
base.
Intercellular.
Intracellular.
Between
Inside a
cells.
cell.
Intramycelial. Within the mycelium. Involute. Enwrapped, having the edges of the leaves rolled inwards.
Irpiciform.
Having teeth resembling those
in Irpex.
GLOSSARY
Isabelline.
689
dirty
tawny
tint.
Isogamous. Used for those plants which produce like gametes. Isogamy. Conjugation of two gametes of similar form.
Keeled.
Carinate.
Labyrinthifonn. Marked by sinuous Lacerate. Torn, or irregularly cleft. Lactiferous. Latex bearing.
lines.
Lamella
(ae).
The
gills of
Agaricales.
Lamellate. Lamelloid.
Made up
of thin plates.
Lamelliform.
In the shape of a plate or scale.
Resembling lamellae. Shaped like a Florence flask. Lanceolate. Narrow, tapering to each end.
Lageniform.
Latericious.
Latticed.
Lateritious.
Brick-red.
Cross-barred.
Lax.
Loose, distant. Lenticular. Shaped like a double convex lens.
Lichenoid.
Lignicole.
Irregularly lobed as lichens.
Growing on wood. Limoniform. Lemon-shaped. Linear. Narrow, several times longer than wide. Lipochrome. A yellow pigment. Lobate. Lobed. Divided into or bearing lobes.
Locule.
Loculus.
cell
or cavity.
is
Lumen.
The space which

cell.
bounded by the walls
of
an organ, as the
central cavity of a
M
Macro.
Mega.
In Greek compounds to signify large.

spots.
Maculicole.
On
Mammiform.
Marginate.
acter.
Breast-shaped.
Broad-brimmed, furnished with a margin
of distinct char-
Matrix. The body on which a Fungus or Lichen grows. Melleus. Melleous. Like honey.
690
GLOSSARY
Membranaceous.
Thin and semi-transparent,
like
Membranous.
fine
membrance.
Medullary. Relating to the pith, pithy. Micro. To signify small, little. Microsporangium (a). A sporangium which produces microspores. Mon. In Greek compounds to signify one. Monopodium (a). An axis which continues to grow at the apex
in
the direction of previous growth, while lateral structures of like kind are produced beneath it in acropetal succession.
Monosporic. Bearing one spore. Monostichous. In a single vertical row.

Slimy. Latin element signifying many or much. Muricate. Rough with short hard excrescences.
Mucosa.
Multi.
Muriculate. Diminutive of Muricate. Muriform. With cells resembling bricks nal and transverse septa. Muticous. Muticate. Pointless, blunt.
in
a wall, with both longitudi-
Mycelium. Vegetative portion more hyphse.
of thallus of fungi
composed
of one or
Myxamceba
(ae).
Swarm-cells with purely amoeboid creeping motion.
N
Nodose. Nodule.
Knotty or knobby.
A small knot or rounded body.
A symbol for the pycnial stage of the rusts. Ob. As a prefix meaning inversely or oppositely. Obese. Excessively f^t; fleshy.
O.
Obligate. Obsolete.
Necessary, essential.
Comp. Facultative. Wanting or rudimentary.

Ocher-colored, yellow with a tinge of red.
Ochraceous.
Olivaceous.
The
color of a ripe olive.
Oogonium. Female sexual organ, containing one or more oospheres. Oosphere. Naked mass of protoplasm which, after fertilization, develops
into the oospore.
Oospore. Immediate product of fertilization of oosphere. Opalescent. Reflecting an iridescent light. Operculate. Furnished with a lid.
GLOSSARY
Operculum.
division.
691
line
lid
or cover
which separates by a transverse
of
Opt.
Ostiolate. Ostiole.
Abbreviation for Optimum. Bearing an ostiole.
An opening or mouth. Broadly elliptic. Ovate. Shaped like a longitudinal section of a hen's egg. Ovoid. Resembling an egg.
Oval.
Pannose.
Papilla
(ae).
Felt-like.
Soft superficial protuberances.
Having papillae. Papilliform. Shaped like a papilla. Papilloid. Resembling a small nipple. Paraphysate. With paraphyses.
Papillate.
Paraphyses. gams.
Parasite.
living
Sterile filaments occurring in the fructification of crypto-
An organism living on or organism (the host).

Shaped
like a patella.
in
and at the expense
of another
Patellate.
Patelliform.
Like a small dish, circular and rimmed.

support.
Pedicel.
The
Pedicellate.
Pellicle.
Borne on a
pedicel.
A small skin;
Wholly or
Like a
a delicate superficial membrane.
Pellucid.
partially transparent.
brush. Pencil-shaped. Pierced through. Peridium. The outer enveloping coat of a sporangium. Periplasm. The protoplasm in the oogonium and which does not share in conjugation.
Penicillate.
little
Perforate.
the antheridium
Perithecium.
Peritrichiate.
rounded, oval, pyriform or beaked structure in which
asci are borne.
With hairs from all of surface. Remaining till the part which bears it is wholly matured. Phycochrome. The coloring matter of brown Algse. Phyllogenous. Growing upon leaves. Phytogenous. Growing on plants. Pileate. Having the form of a cap.
Persistent.
Pileiform.
Pileus.
Pileus-shapcd.
Cap.
The dome-shaped
part of a sporophore.
692
Pilose.
GLOSSARY
Pilous.
Plane.
Hairy, having soft and distinct hairs.
Planose.
asymmetrical sporangium of the Myxogastres. of naked plurinucleated protoplasm exhibiting amoeboid motion. Pleurogenous. Growing from the sides.
Plasmodiocarp.
An
Plasmodium.
Body
Plexus.
Plicate.
Polar.
A network. Folded into plaits usually lengthwise. Relating to the poles of an organ.
Polymorphous. With several or various forms, variable
spored.
Polymorphic.
Polysporic.
as to habit.
Many
Porcelaneous.
Poroid.
Like porcelain.
Resembling pores. Containing pores. Proliferous. Bearing offshoots. Promycelium. Short and short-lived product of tube-germination of a spore which adjoins acrogenously a small number of spores
Porose.
(sporidia) unlike the mother-spore
and then
dies
off.
In prefix signifying false, counterfeit, spurious. Pulverulent. Powdered, as if dusted over.

Pulvinate.
Pseudo.
Cushion-shaped. In the form of a point or dot. Punctulate. Marked with small points.
Punctiform.
Pustular.
Blister-like, bearing blisters.
Pustule.
A pimple or blister.
Having slight blister-like Becoming rotten.
elevations.
Pustuliform.
Putrescent.
Pycnidium (a). A variously shaped cavity resembling a pyrenocarp and containing conidia.
Spores borne in pycnia. of the Uridinales; see pp. 324-326. Pycnium. Pycnosclerotia. Sclerotia bearing pycnidia.
Pycniospore.
A structure
Pycnospores. Spores from pycnidia. Pyriform. Piriform. Resembling a pear in shape.
Radiate.
Ramicole.
Spreading from or arranged around a Growing on branches.
common
center.
Ramose.
Branched.
Receptacle.
That part which bears one
or
more organs.
GLOSSARY
Reniform.
Resupinate.
Reticulate.
693
Kidnoy-shapcd.
Without a
pilcus.
Revolute.
Rhizoid.
Netted, like network. Rolled back from the margin or apex.

root-like structure.
Rhizomorph. A root-like branched strand of mycelial hyphse. Rhomboidal. Approaching a rhombic outline. Rimose. Rimous. Cracked.
Rostrate.
With a beak.
Rostrum. Any beak-like extension. Rufous. Reddish. Rugose. Rugous. Covered with wrinkles.
Saccate.
Bag-shaped.
Saprophyte. Sarcinaeform.
Having the form
plant living on dead organic substance. of the genus Sarcina.

to the touch.
Scabrous.
Sclerotioid.
Rough
Like a sclerotium.
Sclerotium.
Scopulate.
A compact mass of hyphae in dormant state. Broom-Uke or brush-like. Scorpioid. With the main axis coiled like the tail of a scorpion.
Scrupose. Jagged, rough. Scutiform. Buckler-shaped,.
Septate.
Septum
Seriate.
Sessile.
Seta(ae).
(a).
Divided by a partition. Any kind of partition.

series.
In a
Destitute of a stalk.
bristle or bristle-shaped
Bristle-like.
body.
Setaceous.
Setose.
Setulose.
Bristly, beset with bristles.
Resembling a fine bristle. In the form of a buckler; clypeatc, peltate, or scutate. Sigmoid. Doubly curved in opposite directions, like the Greek sigma. Simple. Of one piece or series, opposed to compound. Sinuous. Sinuose. Sinuate. Withadeep wavy margin. Sorus (i). Heap, or aggregation; a heap of spores.
Shield-shaped.
Spatulate.
Like a druggist's spatula.
Sperm.
male reproductive
(a).
cell.
Spermatum
Male non-motile gamete, sometimes erronously used
for various conidia-
694
Sphseroidal.
GLOSSARY
Somewhat
spherical.
Spindleform.
Spindle-shaped, fusiform.
Sporangiophore. A sporophore bearing a sporangium. Sporangium. Sac producing spores endogenously.

Spore.
single cell which becomes free directly into a new plant.
and
is
capable of developing
Sporidium. Diminutive of spore, especially applied to the spores produced on promycelia. Sporocarp. A many-celled body serving for the formation of spores.
Sporodochium.
Sporogenous.
Stellate.
The
sporiferous
apparatus
in
fungi
belonging
to
the Tuberculariales.
Producing spores.
Star-shaped or radiating like the points of a star. (ta) . A stalk-like branch of a basidium bearing a spore. Sterigma
Stipe.
A general
A
term
for stalk.
Stipitate.
Having a
stipe.
Stolon.
sucker or runner.
Stoloniferous.
Stratose.
Striate.
Bearing stolons. In distinct layers. Marked with fine longitudinal parallel
lines.
Strigose.
With sharp-pointed appressed
straight
in
and
stiff
hairs
or
bristles.
Stroma
(ta).
cushion-like body, on
or
which the perithecia are
immersed.
Stromatic.
Stuffed.
Pertaining to or resembling a stroma.
Solid, farctate.
Stylospore.
A spore borne on a filament.

;
Sub.
or below; in compounds usually implies an approach to the condition designated somewhat or slightly. Subiculum. Subicle. A felted or byssoid basal stratum of hyphse.
Under
Subulate.
Sulcate.
Awl-shaped.
Grooved or furrowed.
Superficial.
On
the surface.
Suspensor.
club-shaped or conical portion of hypha adjoining a

axis
gamete-cell.
Sympodium.
An
made up
of the bases of a
number
of successive
axes arising as branches in succession one from the other. Syn. Signifies adhesion or growing together.
Synema.
A column of
combined filaments.
GLOSSARY
695
Abbreviation for Thermal-death-point. A resting spore of Uridinales on germination producing a promycelium. Teleuto-stage. Stage producing a teleutospore.
Teleutospore.
T. D. P.
Telium.
Terete.
A sorus of the Uredinalcs; see pp. 324-326. Circular in transverse section.

In threes.
Ternate.
Thalloid.
Having the nature or form of a thallus. Thallus. A vegetative body without differentiation into stem and leaf. Tomentose. Densely pubescent with matted wool, or short hairs. Tortuous. Bent or twisted in different directions.
Torulose.
Irregularly bending.
Somewhat moniliform with swollen
portions.
Tremellifonn.
Tremelloid.
Gelatinous in texture.
Jelly-like in substance or appearance.
Trichogyne.
Trident.
The
receptive filament of the female organ.

teeth.
Having three
Triquetrous. Three-edged, with three salient angles. Truncate. As though cut off at the end.
Tubercular.
Tuberculate.
Having
tubercles, or like a tubercle.
Beset with knobby projections or excrescences.
Tubular. Cylindrical and hollow. Turaid. Inflated, swollen.
U
Umbellate.
Umbilicate.
Having the
Navel-like.
boss.
inflorescence in umbels.
Umbo.
Umbrinaceous. Umbrinous. raw umber. Uncinate. Hooked.

Ungulate.
Umbrinus.
Umbrinose.
The
color of
Having claws or
Hoof-shaped.
hoofs.
Unguliform.
Uni. In composition, one, or single. Urceolate. Urceolar. Pitcher-like, hollow and contracted at the
mouth
an urn. Uredinium. A sorus

like
in the Uredinales, see pp. 324-326.
696
GLOSSARY
Vacuolate.
Valsoid.
Possessing vacuoles.
Resembling Valsa. Valvate. Opening by valves. Vegetative. Growing.

Veil.
special envelope in Agaricales within
which the growth
of the
sporophore takes place. Velutinous. Velvety, due to a coating of fine soft hairs. Vermicular. Venniculate. Worm-shaped, thickened and bent in places. Verruciform. Wart-shaped.
Verrucose.
Verticillate.
Verrucous. Whorled.
Verruculose.
Full of warts.
Warty.
Vesicular.
Villi.
-Composed
villi.
of vessels.
Long weak
Bearing Vinous.
hairs.
Villous.
Vinose.
Viscid.
Wine
colored.
Volva.
Sticky from a tenacious coating or secretion. A covering, the sac enclosing the Agaric sporophore.
Zonate.
Zooglceae.
Marked
circularly.
colony embedded in a gelatinous substance.
Zoosporangia.
Sporangia which produce zoospores.
Zoospore.
motile spore.
INDEX
(Boldface figures refer to headings, italic figures to illustrations)
Actiniceps, 634 Actinida, 181

Abies, 229, 230, 347, 391, 416, 558,
Actinomma, 656
Actinonema, 505, 508 Fagicola, 509 F'raxani, 509 Rosa;, 508, 508 Tilia), 509 Actinothecium, 528 Actinothyrium, 532
Adonis, 178 iEcidium, 324, 334, 335, 389, 390
560
Absidia, 104
Acacia, 436, 457
Acanthorhynchus, 224 Vaccinii, 224, 225 Acanthostigma, 226, 229 Parasiticum, 229, 229 Acer, 188, 202, 499, 507, 529 Pseudoplatanus, 152, 159 Aciculosporium, 199
Acinula, 659
Acladium, 575 Acolium, 153 Aconitum, 93
Acontium, 571 Acorus, 73, 320

Acrasiales, 5
Abietinum, 349 Asperifolium, 381 Berberidis, 334, 378 Brassica;, 378, 390 Cinnamomi, 390 Columnare, 347 Cyparissia;, 372
EuphorbifE-gerardiana;, 375
Grossularia;, 376
Acremoniella, 600, 600

Occulta, 600
Leucospermum, 336
Mespili, 371
Verrucosa, 600
Acremonium, 575, 577, 577

Acrocylindrium, 583 Acrocystis, 663
Batata;, 663
Otogense, 390
Oxalidis, 384
Pclargonii,
390 Rhamni, 382
Acrospeira, 599
Acrospermaceaj, 160 Acrostalagmus, 583, 584, 584

Albus, 584, 585
Rubollum, 377 Strobilinum, 347 Tuberculatum, 390 Êgerita, 640

^sculus, 182, 202, 489, 507, 579 Agaric, 394
Agaricaceae, 402, 442
Panax, 585
Vilraorinii,
585
Acrotheca, 598
Acrothecium, 609 Actidium, 164
Key
697
to,
442
Agaricales, 395, 397
698
Agaricales,
INDEX
Key
to,
402
Alternaria, Violae, 621, 6S1

Vitis,
Agariceae, 443, 448
624
Agaricus, 448, 455
Althea, 386
Agave, 89, 248, 552 Agropyron, 306, 320, 389 Repens, 262 Agrostis, 321, 380 Agyriella, 538 Agyriellopsis, 534 Albuginacese, 78, 82 Albugo, 78, 79, 82 Bliti, 79, 81, 82, 102-116 Candida, 81, 95
Ipomoese-pandurana;, 81
Occidentalis, 82
Portulaceae, 82
Alyssum, 178 Alveolaria, 341 Amallospora, 657 Amanita, 398, 450 Amantiopsis, 450 Amarantaceaj, 303 Amaranthus, 82 Amaryllidacese, 405 Amblyosporium, 572
Amelanchier, 183, 306, 368, 369, 370, 371
Amentaceae, 127
Amerosporse, 633, 635 Key to, 636 Amerosporium, 534
Tragopogonis, 82
Alder, 130, 203, 243, 264, 274, 419,
428, 545 Aldridgea, 405
Aleurodiscus, 404, 405 Aleyrodes, 194

Alfalfa, 29, 36, 97, 132, 148, 206, 250,
fficonomicum, 534 Amocbochytrium, 72 Ampelopsis, 181, 238, 484
Amphicarpa, 71
Amphisphseriaceae, 222
258, 315, 408, 486, 508, 514, 521,
Amphisporc, 327
Anacardiacese, 127
Ancylistidiales, 66
543, 551, 556, 582, 630, 660

Alfilaria, 71, 101
Alga-like Fungi,
3,
65
AUantospora, 589
Allium, 97, 318, 344, 661
Andromeda, 398 Anemone, 72, 143,

389
178, 318, 336, 357,
Almond,
607
36, 357, 498, 542,
626
Angelinia, 160
Alnus, 186, 188, 278, 408, 541, 582,

Aloe, 493, 560
Alternaria, 260, 261, 616, 621
Angiopoma, 515 Anise, 630

Anixia, 189
Annularia, 450
621 Cucurbitse, 621 Dianthii, 622, 632 Fasciculata, 624 Fici, 624 Forsythise, 621 Panax, 622 Phaseoli, 621 Solani, 623, 6S3, 624 Tabacinum, 624 Tenuis, 621 Trichostoma, 262, 621
Brassicse,
Antennaria, 190, 192 Elaeophila, 192

Footi, 192
Piniphilum, 192
Pityophila, 192
Setosa, 192
Anthina, 657
Anthostoma, 277
Anthostomella, 203, 276 Bohiensis, 277
-
-Coffese, 277
Destruens, 277, 277
INDEX
Anthostomella, SuUae, 277
Artichoke, 41, 591 Arundinaria, 211
Aschersonia, 195, 527
693
Anthracoderma, 483 Anthracophyllum, 445 Anthrinieie, 594
Asciculosporium Take, 211

Asclepias, 188
Anthurium, 270, 489, 511, 544, 552 Anthurus, 4G3 Antromycopsis, 630
Aphanoascus, 167 Aphis, 194 Apiospora, 251
Apiosporiuni, 190, 191, 192
Brasiliense, 192
Ascobolacca;, 133, 134
Ascochyta, 243, 409, 505, 606 ^sculi, 507 Aquilcgiu', 507

Armoracia;, 506
Aspidistras,
507
Beticola, 507
Salicinum, 191, 192 Aposphacria, 481, 494
Boltshauseri, 506
Apple, 36, 38, 52, 88, 105, 122, 123,

139, 184, 203, 220, 231, 246, 248,
Brassies, 506 Caulicola, 508
249, 255, 265, 267, 268, 271, 278,

285, 329, 365, 367, 368, 371, 404,
405, 408, 412, 415, 419, 421, 428, 452, 485, 486, 490, 492, 502, 503, 504, 515, 519, 539, 540, 542, 546,
547, 561, 563, 574, 586, 617, 644, 645, 649
Rot, 266
Twig Canker, 266

Appressoria, 266
Apricot, 139, 357, 486, 491, 560, 569,
604, 610 Aquilegia, 178, 544
Arachnopeziza, 135 Aralia, 168
Arbor
Vitse, 424,
431
Arbutus, 36
Archangelia, 251
Armillaria, 123, 450, 456
Fuscipes, 457
Mcllea, 230, 410, 411, 455, 457,
458 Mucida, 457

Arnica, 176
Aronia, 366, 368
Arrhenatherum, 180 Arrhenia, 443 Arrowroot, 207 Arthrobotrys, 586 Arthrobotryum, 637
Chrysanthemi, 507 Cookei, 508 Corticola, 508 Digitalis, 507 Dianthi, 507 Ellisii, 506 Fragariaj, 507 Graminicola, 508 Imperfecta, 508 Iridis, 507 Juglandis, 507 Lactucse, 507 Lycopersici, 508 Manihotes, 508 Medicaginis, 507 Melutispora, 508 Nicotiana), 506 Orobi, 507 Pallida, 507 Parasitica, 506 Piniperda, 507 Pisi, 506 Polemonii, 507 Populicola, 507 Primula;, 507 Pruni, 507 Rhei, 506 Tremula;, 508 Vicia;, 503 Violffi, 507
Ascocorticiaceae, 125
700
Ascoideaceae, 118
INDEX
Atractina, 609
Ascomycetes,
64,
113
Key
to,
117
Atractium, 207 Atriplex, 74

Atrocarpus, 273, 541
Ascospora, 235, 236
560 Geographicum, 237 Himantia, 236 Padi, 237 Aseroe, 463

Beijerinckii, 236,
Attachment organs,
Auerswaldia, 216 Aulographum, 163
141, 142
Ash, 52, 164, 203, 232, 255, 284, 419

421, 433, 509, 524, 530, 557, 564,
603, 608
Aureobasidium, 403, 405 Vitis, 405 Auricula, 611

Auricularia, 393
Auricula judia;, 393

Auriculariacea;, 392
Asparagus, 41, 328, 376, 553, 617, 630, 660 Aspedistra, 507, 552
Key
Key
to,
392
Auriculariales, 323, 326, 392
Aspen, 255, 428, 446, 607 Aspergillacea), 165, 166 Key to, 166
Aspergillales, 114, 124, 164, 167
Key
to,
165
Aspergillese, 566, 572,
572
392 393 Autoecious, 329 Avena, 608 Avocado, 512 Azalea, 523
to,
Auricularieaj,
Key
573
to,
572
Aspergillus, 166, 167, 168, 169, 572,
B
Bacillus, 18, 21, 37
Ficuum, 169, 572 Fumigatus, 572

Phoenicis, 169, 572
Ampelopsorse, 51, 37
Aster, 89, 179, 289, 328, 585, 654
Amylovorus, 3S, 38
Anthracis, 37
Apii,
Asterocystis, 68, 69
Radicis, 69
39
Asterodon, 413
Araliavorus, 39
Aroidea;, 39, 39, 42
Asteroma, 274, 482, 496 Codisei, 496 Geographicum, 496 Padi, 275, 496 Punctiforme, 496 Stuhlmanni, 496 Asterophora, 577 Asterosporium, 558
Asterostomella, 481, 531
Atrosepticus, 40, 47
Avense, 24, 40, 40

Betse, 41
Brassicaevorus, 27, 41
Caratovorus, 39, 40, 41, 42, 46 Caulivorus, 27, 43

Cepivorus, 43
Coll, 43 Cubonianus, 31, 43 Cypripedii, 43 Dahha;, 43 Delphini, 43
Asterostomidium, 514 Asterostroma, 406 Asteroth5Tium, 531

Astragalus, 187
Astrodochilum, 655 Atractiella, 634
Elegans, 43
Glangse, 52
INDEX
Bacillus, Gossypini, 43
701
702
Basidiobolus, 107
INDEX
Bird nest fungi, 395 Bispora, 601
Bizzozeria, 234
Bizzozeriella,
Basidiomycetes, 64, 298, 475 Key to, 299

Basidiophora, 83, 89
640
Entospora, 89 Basisporium, 599
Blackberry, 276, 360, 525

Black-leg, 652
Baumanniella, 411, 412
Bean,
12, 31, 37, 86, 267, 373, 408,
Blastomyces, 575 Blastotrichum, 200, 588

Blennoria, 538
Bletia, 552,
487, 491, 506, 540, 548, 621, 630, 637 Beccariella, 406
645
Beech, 106, 134, 152, 162, 203, 249,

415, 421, 427, 429, 430, 433, 436,
440, 444, 445, 509, 544, 545, 547,
Blossoms, 107, 108 Bloxamia, 538
Blue grass, 213, 385, 497, 550

Blue-green Alga;, 3
Boletacese, 402, 440
60o
Beet, 8, 22, 26, 36, 37, 41, 44, 45, 50,
52, 73, 82, 100, 187, ;247, 258, 321,
412, 486, 490, 507, 526, 568, 590, 610, 617, 620, 628, 645, 660
Begonia, 43, 168, 544 Belanioscypha, 136
440 440 Bolinia, 285 Bombardia, 226
Key
to,
Boletineae,
Belonidium, 147 Beloniella, 147
Belonium, 136 Belonopsis, 147 Beltrania, 602

Beniowskia, 639
Berberis, 185, 188, 235, 256, 329, 379,
Bombardiastrum, 227 Bonia, 406 Bonplandiella, 655 Boraginacese, 329, 382

Borago, 178 Bornetina, 323
Bostrichonema, 586
Botryodiplodia, 510, 513
Botryospha^ria, 283, 283, 503
Dothidae, 284
384
Berlesiella,
283
Bertia, 226
Gregaria, 284
Ribis, 283
Beta, 374
Betula, 130, 157, 186, 188, 202, 220,
221, 255, 348 Betulacea}, 143
Bibliography of introduction, mycetes and Bacteria, 53
Myxo-
Ascomycetes, 288 Books, 678 Basidiomycetes, 466 Fungi Imperfecti, 667 Periodicals, 680 Phycomycetes, 109
Biologic
species
Botryosporium, 571 Diffusum, 571 Longibrachiatum, 571 Pulchrum, 571 Botrytideae, 566, 574, 575 Key to, 575
Botrytis, 86, 91, 96, 137, 142, 576, 578
Cinerea, 140, 578, 578, 581

Citricola, 581
Depraedens, 580 Diospyri, 581
and
specialization,
174, 260, 262, 332, 380, 611,
640
Douglasii, 141, 681 Fascicularis, 580
Birch, 418, 421, 426, 428, 430, 433,

436, 444, 446, 545, 607
Galanthina, 141, 581

Infestans, 581
INDEX
Eotrytis, Longibrachiata, 581
703
278, 409, 411, 448, 493, 509, 512,

525, 536, 552, 553, 573, 584, 585, 590, 592, 620
580 Parasitica, 580 Patulu, 581 Vulgaris, 140 Boudiera, 115, 116 Bovilla, 224 Bovista, 465 Bovistella, 465
Paeoniae,
Cacospha;ria, 281
Cactus, 36, 488, 512, 544

Ca;otna, 334, 335, 342, 358, 361, 389,
390
Nitcns, 360
Orchidis, 344
Box, Buxus, 204, 220, 221, 243, 529, 656 Box elder, 545
Pinitorquum, 344
Calamagrostis, 661
Calcarisporium, 583
Calceolaria, 52
Brachysporium, 609 Brachy-type, 328

Brassica, 69, 178, 390, 617, 629
Calendula, 176, 178, 321 CaliciacesD, 134, 153
Bread
fruit,
411
Key
to,
153
Brefeldiacea?, 10
Calicium, 153
Calistephenis, 338
Calla, 39, 488, 500, 631
Calloriea?, 147
Bremia, 84, 90, 95

Lactucae, 95, 95
Briarea, 572
Briosia, 636
Calocasia, 89
Bromus, 260, 261, 613

Broomella, 199 Brown alga), 3
Brunchorstia, 632
Calonectria, 198, 205

Bahiensis, 206
Destruens, 151, 532 Bubakia, 340
Buckwheat, 378, 589, 607, 660

Bulb, 141, 661
Bulgaria, 151, 152
Cremea, 205, 585 Flavida, 205 Gigaspora, 206 Platani, 206 Pyrochroa, 206, 648 Calosphseria, 281, 282 Princeps, 282, 282
Calospha?riea;, 281
Polymorpha, 162
Bulgarieae, 151
Bulgariella, 151
Calospora, 280, 539 Vanilla, 280, 541
556 315 Buseella, 570 Butomus, 323

Bullaria,
Burillia,
Calyptromyces, 104 Camarops, 285
Camarosporium, 516, 517, 617 Fissum, 617

Viticola,
Butternut, 275, 419, 428, 545

Byssocystis, 482
Mori, 517 517
Camellia, 497, 559, 561
Campanclla, 443
Cabbage,
7, 24, 25, 26, 29, 41, 42, 46,
52, 69, 73, 81, 95, 231,
249, 378,
408, 484, 491, 492, 530, 544, 619,
503, 654
Campanula, 101, 333, 340 Camphor, 411 Camposporium, 609 Campsotrichum, 599
Camptosphaeria, 263
Cacao, 130, 204, 205, 203, 232, 277,
704
Camptoum, 598 Canker, 266, 349 Canna, 389 Canteloupe, 620
Cantharelleae, 442, 443
INDEX
Catastoraa, 465
Catenularia, 600 Catinula, 534
Cattleya, 253, 273, 541, 544
Caudospora, 277
Cauliflower,
25,
443 Cantharellus, 443 Caper, 81, 630 Capillaria, 659 Capnodiastrum, 501
Key
to,
26,
28,
46,
95,
544
Caulocassia, 100
Cedar, 234
Capnodium,
Citri,
190, 192,
624
incense, 424 Celastrus, 188
193
Celery, 27, 39, 41, 42, 52, 377, 487,

492, 521, 619, Celidiacea;, 134
Citricolum, 193
Coffeaj, 192
628
Foedum, 192
Guajavse, 192 Javanicum, 193
Meridionale, 193
Olea, 193
Cenangella, 150 Cenangiacea;, 134, 150 Key to, 150
Cenangium,
Vitesia,
150, 151,
152
Abietis, 151,
532
Quercinum, 192
Stellatum, 193
Taxi, 192
Tiliaj,
500
192
Centaurea, 178, 377 Cephaliophora, 588 Cephalodochium, 641

Cephalosporiea;, 566, 570
Capparis, 179
Capronia, 252 Capsule, 14
Key
to,
570
Caragana, 524, 525 Caravonica, 411 Carduacese, 303 Carex, 303, 376
Carnation, 27, 52, 375, 408, 497, 523,
544, 553, 578, 580, 581, 611, 623,
645, 654
Carpinus, 130, 188, 191, 492, 545

Carrinia, 504
Carrot, 27, 41, 42, 44, 91, 119, 141,

142, 260, 408, 491 Carya, 186, 202, 546
Caryophyllacea;, 310, 405
Cassava, 493, 543

Castanea, 140, 186, 188, 558, 562 Castilloa, 411
Castor plant, 89, 654

Casuarina, 571
Catalpa, 168, 178, 186, 188, 256, 426,
489, 619
Cephalosporium, 201, 571, 646, 649 Cephalotheca, 166 Cephalothecium, 586 Roseum, 586, 586 Cephalotrichum, 598 Ceraplastes, 194 Ceratiomyxacese, 9 Ceratocarpia, 190 Ceratocladium, 630 Ceratophorum, 608, 610 Setosum, 610 Ulmicolum, 610 Ceratosphaeria, 232 Ceratostoma, 232 Ceratostomatacese, 222, 232 Key to, 232 Ceratostomella, 232 Pilifera, 233, 233 Cercis, 506, 524 Cercospora, 243, 257, 478, 625 Acerina, 632
INDEX
Cercospora, Acciosum, 630
AUha'iiui, 630
705
Cercospora, Rocslcri, 626

Ro-sicola, 630,
6J0
Angreci, 631
Rubi, 626
Sacchari, 630
Sequoia,',
626 628 Arimincnsis, 630 Armorac'i;p, 629 Asparagi, 630

An{j;ulata, 245,
Apii, 628,
632
Sordida, 631
Thoa;, 630
Unicolor, 631
Vagina,",
Beticola, 628,
628
630
Bloxami, 629 Bolloana, 626

Brunkii, 631
Vignae, 630
Viola>,
630
Viticola, 626
Capparidis, 630
Cerasella, 245, 625
Cercosporella, 592
Cercidicola, 631
Cheiranthi, 631 Circumscissa, 625
Albo-maculans, 692 Inconspicuus, 592 Narcissi, 592

Pastinacaj, 692
Persica?,
629 Concors, 626, 626 Cruenta, 629 CucurbitsD, 629 Flagelliformis, 629 Fumosa, 626 Gossypina, 248, 625 Halstedii, 632 Hypophylla, 631 Kellermanii, 630 Kopkei, 630 Longipes, 630 Malkoffi, 630 Malvarum, 630 IMedicaginis, 630 Melonis, 629 Microsora, 631 Moricola, 626 Musffi, 626 Neriella, 631
Citrullina,

592
Cereal, 248, 250, 385, 491, GOO
Cereus, 499, 529 Cerocorticiuni, 405
Cerotelium, 341
Cesatiella, 198
Ceuthospora, 483, 500

Cattleya, 500,
Coffeicola, 500
MO
Cha;notheca, 153
Chajtocladiaceai, 103
Chcctoconidium, 577 Chatodiplodia, 510

VaniUa;, 510 Cha;tomella, 501
Chffitomiacea;, 222 Chajtopcltis, 532
Cha;tophoma, 191, 482, 496 Glumarum, 496
Nicotians, 627, 627 Odontoglossi, 631
Cha^tospermum, 641 Chstosphajria, 226

Chffitostroma, 201, 655, 666
Omphacodcs, 631
Oryzai, 626
Buxi, 656
CUvia;, 656
Personata, 629
Phlogina, 631
Raciborskii, 627
Chsetozythia, 527
Reseda, 631
Richardiaecola, 631
Chalara paradoxa, 506 Chalariea;, 595 Chamajcyparis, 370, 416

Characeae, 3
706
Charrinia, 262, 263
INDEX
Ciliospora, 527
Cheiranthus, 619, 631 Chenopodiaceae, 96
Cineraria, 339
Cinnamon, 390, 398, 487

Cintractia, 302, 310
Chenopodium, 74
Cherry, 49, 129, 138, 184, 219, 237,
245, 255, 268, 275, 282, 357, 376, 419, 507, 512, 520, 560, 562, 604,
606, 625, 649
Cionothrix, 342
Circinella, 105
Cissis,
303
44,
45,
Citron, Citrus Fruits,
193,
Cherry
laurel,
410
194, 249, 490, 491, 504, 512, 520,
Chess, 550
540, 541, 548, 559, 574, 581, 604, 605, 626, 649 Citysus, 100
Chestnut, 36, 101, 209, 281, 321, 419,

420, 439, 442, 445, 489, 498, 499, 509, 524, 560, 597
Cladobotyrum, 583
Cladochytriaceae, 67, 72
Chicory, 490
China berry, 202 Chitonia, 448

Chitonospora, 251
Key
to,
72
Chlamydobacteriaceae, 19
Cladochytrium, 72 Brassicse, 73 Csespitus, 73

Graminis, 73 Mori, 73
Chlamydospores, 60
Chlorophycea?, 3
Chlorosplenium, 135, 144 iEruginosum, 143, 144
Tenue, 73 Violaj, 73
Viticolum, 73 Cladoderris, 406
Cladosphaeria, 233
Choanephora, 107 Americana, 107 Cucurbitarum, 107 Infundibulifera, 107

Choanephoraceae, 103, 106, 107 Chondromyces, 634
Cladosporium, 603, 617
217,
219,
257,
602,
Bigarardia, 605
Chromosporium, 566 Chrysanthemum, 36,

589
105, 386, 389,
392, 409, 488, 492, 507, 522, 563,
Chrysomyxa, 341, 350 Abietis, 350

Chrysophlyctis Endobiotica, 70, 71 Chrysopsora, 336
Chytridiales, 60, 65, 66, 101
Key
Cicer, 179
to,
67
Ciboria, 135
Cichorium, 178, 378 Cicinnobella, 501

Cicinnobolus, 174, 482, 494
Cienkowskia, 11 Ciliciopodium, 634

Ciliofusarium, 657
Brunneo-atrum, 606 Carpophilum, 604, GOJ^. Citri, 604 Condylonema, 605 Cucumerinum, 603, 604 Elegans, 604 Epiphyllum, 606 Fasciculare, 260, 603 Fulvum, 604, 605 Graminum, 605 Herbarum, 248, 603 Citricolum, 604 Hypophyllum, 606 Javanicum, 606 Juglandis, 606 Macrocarpum, 605 Orchidis, 605 Oryzse, 605
INDEX
Cladosporium, Oxycocci, 606 Pconia\ GOG Pisi, 605 Scabies, 606
Scribncrianum, 606 Sicophiluni, 604
Clove, 415
707
Clover, 71, 74, 97, 143, 187, 206, 220,

221, 258, 373, 374, 494, 543, 551,
569, 582, 593, 606, 019, 630
Clusia, 560
Tuberum, 606
Zcaj,
Clypeosphairia, 270 Clypeosphx-riacoa', 223, 276
606
Key
to,
276
Cladosterigma, 634 Cladotrichum, 601
Coccaceae, 18, 21 Cocci, 13
Clanostachys theobroma;, 206 Clasterosporium, 608, 609 Amygdalearum, 610 Carpophilum, 5G0, 610 Glomerulosurn, 610 Putrefaciens, 610 Clathraceaj, 4C2, 463 Key to, 463 Clathrus, 464 Claudopus, 449 Clautriavia, 462 Clavaria, 412 Clavariacea;, 402, 412 Key to, 412
Claviceps, 199, 211, 211, 213, 215, 643
Coccomyces, 156, 563

Cocconia, 156
Coccophacidium, 156
Coccospora, 566
Coccosporella, 566
Coccosporium, 616 Cocoa, 88
Cocoanut, 513
43,
191,
193,
433,
512,
Codiaeum, 496, 544 Coemansia, 583

Cccmansiella, 571
Coffee, 168, 170, 192, 193, 195, 205,
207, 249, 250, 277, 356, 409, 415,
486, 500, 504, 512, 515, 525, 543,

553, 583, 591, 597, 635, 643, 660
Microcephala, 213
Paspali, 213
Purpurea, 212, 643

Rolfsii,
213
199
Colchicum, 375, 563 Coleosporiacea;, 335 Key to, 336

Coleosporium, 326, 333, 336, 390, 391 Campanula?, 339
Ipwmcea', 337
Pini,
Clavicipitea;, 196, 197
Key
to,
Cleistotheca, 190
Clematis, 43, 178, 179, 390, 492, 544,
339
563
Clinoconidium, 640 Clithris, 156, 157, 157
Aureus, 157 Juniperus, 157
Quercina, 157 Clitocybe, 450, 457, 455, 569
Parasitica,
Clitopilus,
Senecionis, 338,
Solidaginis, 337,
338 337
Coleroa, 226, 227, t;58
Chaitomium, 227, 227

Sacchari, 227
Collacystis, 527
CoUard, 25, 95, 619

Colletotrichum,
147, 264, 267, 268, 538, 539, 540, 547
458
450
Clivia, 656
Clonostachys, 584 Clostridium Butyricum, 13

Persicae tuberculosis, 63
Agaves, 562 Althasa;, 552
Ampelinum, 549
Anthurii, 552
708
Colletotrichum, Antirrhini, 553
Bletiae,
INDEX
Colutea, 187
552
Brachytrichum, 553 Camolliae, 553

Carica, 549, 5J^9
Cereale, 550, 550
Completoria, 107 Complens, 108
Composite, 82, 92, Comptonia, 352

Conidiobolus, 107
95, 178,
386
Cincta, 269, 647
Conifers, 132, 145, 159, 161, 162, 203.

229, 235, 350, 390, 417, 418, 423,
Coffeanum, 563 Cradwickii, 653 Cyclamens, 653 Dracaense, 553 Elasticse, 552 Falcatum, 549
Gloeosporioides, 549
431, 436, 452, 512, 547, 559, 579,
654 Coniocybe, 153 Pallida, 153 Coniophora, 405

ConiosporieiE, 594, 595
Gossypii, 267, 272, 547
Hedericola, 553
Heveae, 553
Key
to,
595
Coniosporium, 595, 595 Filicinum, 595

Onobrychidis, 595 Coniothecium, 615 Chomatosporum, 617
Incarnatum, 553 Kentiai, 552 Lagenarium, 648 Lineola, 550
Lindemuthianum, 267, 543, 647,
Anomale,
54s Luxificum, 552
Macrosporum, 553 Malvarum, 552 Nigrum, 561

Oligochajtum, 648
Coniothyrium, 245, 257, 501, 603 504 Brevisporum, 504 Coffea;, 504 Conccntricum, 503
Diplodiella, 263, 504, 50^
Fuckelii, 257, 258, 503 Hellebori, 505
Omnivorum, 662
Phoraoides, 551 Piperitum, 551
Pollaccii,
553
Japonicum, 228 Megalospora, Melastorum, 503 Pyriana, 503

Scabrum, 504
Tumefaciens, 503 Vagabundum, 504 Wernsdorffia;, 505
Conjugatae, 3
504
Primula;, 562
Rubicolum, 270, 547 Schizanthi, 552 Spinaciffi, 551 Theobromae, 563 Theobromicolum, 553 Trifolii, 551 Viote-tricoloris, 562 CoUodochium, 640 Collonema, 517 Collybia, 450, 468
Vclutipes, 468, JfiO
Conjugate division, 321, 331, 332 Convallaria, 494 Convolvulaceae, 82 Coprinese, 442 Corallodendron, 633
Corallomyces, 199
Cordana, 602
Cordieritidacese, 134
Columbine, 507 Colus, 464
Cordyceps, 199
INDEX
Coremium, 230,
Coreopsis, 176
634, 635, 635
709
Cranberry, 71, 140, 225, 231, 267, 277,

397, 486, 495, 500, 520, 525, 529, 536, 543, 559, 606, 614
Corethropsis, 571
Coronella, 570
Crandallia, 532
CratiCKus,
130, 367, 368, 569 183, 188, 255,
Corn, 34, 52, 90, 101, 308, 310, 312,

329, 408, 511, 512, 606, 613, 614,
366,
650
Cornularia, 517
Cratcrellus, 406
Craterium, 12
Creonectrieae, 196
Cribrariaceae, 9
Cornus, 188, 191, 279, 524 Coronophora, 281 Corpyha, 560

Corticium, 403, 405, 406, 409
Crocicreas, 481
Crocus, 121, 143
Chrysanthemi, 409 Comedens, 409 Dendriticum, 409 Javanicum, 409 Lajtum, 408 Lilacino-fuscum, 409
Vagum-solani, 660
Cortinarius, 449
404,
Cronartium, 341, 350, 351, 390, 391 Asclcpiadeum, 352 Comptonia?, 352, 35^ Querous, 352, 390 Ribicola, 350
Crotalaria, 408
4C6,
407,
Crotonocarpia, 226 Crown gall, 12

Crucifcr, 6, 25, 41, 43, 69, 81, 95, 616, 619, 621
Zimmermannii, 409
Corydalis, 344
Corylus, 188, 408, 530, 545, 557
Corymbomyces,
583, 584, 584
Albus, 205, 584

Coryneliacese, 222
Crumenula, 150 Cryptocoryneum, 608 Cryptoderis, 263 Cryptomela, 553

Cryptomyces,
156, 158
Corynespora, 629 Corynetes, 131
Maximus, 158
Cryptospora, 279
Cryptosporclla, 279, 280
Coryneum, 558, 560, 564, 658

Beijcrinckii, 236, 237, 560, 5(11
Anomala, 280
498 Cryptosporium, 562, 564 Loptostroniifornie, 564 Minimum, 564
Viticola, 280, 2S2,
Cryjitostictis, 515, 516,
Camellise, 561
FoUiicolum, 561
Juniperinum, 560, 658 Modonia, 560 Mori, 561

Cosmonectria, 201 Cosmos, 493
Costinellus, 450
516
Caudata, 516 Cynosbati, 516
Cucumber,
604,
44, 45, 46, 51, 52, 76, 95,
Cotton, 29, 36, 43, 249, 267, 272, 303,

361, 408, 411, 486, 492, 547, 589, 619, 625, 650, 651
141, 247, 404, 487, 543, 548, 560,
606,
608,
615.
617,
629,
652
Cucumis, 178 Cucurb, 6, 51, 94, 95,
Cucurbita, 178
107-, 178, 179, 487, 509, 521, 539, 548, 051
Cotton-wood, 437 Cotoneaster, 366 Couturea, 515
Cowpea,
187, 487. 521, 535, 651
710
Cucurbitaria, 234
Berberdis, 235, 235
INDEX
Cylindrosporium, Orni, 564 Padi, 562, 662, 563
Elongata, 235
Pomi, 563, 56S

Quercus, 564
Laburni, 235 Picete, 235

Pityophila, 235
Sorbi, 235 Cucurbitariaceffi, 222, 234
Saccharinum, 563 Tubeufianum, 563 Viridis, 564

Cylindrotrichum, 575 Cymbridium, 547
Key
to,
234
Cudonieae, 131 Cuphea, 187
Cynachum, 352
Cynodon, 221
Cyperaceaj, 303
Cupressus, 369
Currant, 92, 148, 155, 203, 207, 284,

351, 433, 499, 500, 519, 542, G26
Cyperus, 89, 303, 408 Cyphella, 406
Currya, 216 Curryella, 216 Cuttings, 644
Cystophora, 598
Cytisus, 235, 253, 375, 610
Cyanocephalium, 198 Cyanophycese, 3 Cyanospora, 232, 233 Albicedra;, 233, 2SS Cyathicula, 136 Cycad, 248, 516
Cytodiplospora, 505 Cytoplea, 501
Cytospora, 209, 279, 483, 499, 499 Acerina, 499
Ceratophora, 499 Palmarum, 499
Rubescens, 278
Sacchari, 499
Cyclamen, 553, 579
52, 168, 268, 488, 492, 522,
Cytosporella, 483, 498

Cerei, 499
Citri,
Cycloconium, 601, 602, 603 Oleaginum, 603 Cyclomyces, 417 Cyclostomella, 163 Cydonia, 366, 368 Cylindrium, 568 Cylindrocephalum, 571 Cylindrocladium, 586 Cylindrocolla, 641 Cylindrodendrum, 577 Cylindrophora, 576
Cylindrosporium, 243, 562, 562 Castanicolum, 249, 562 Cercosporoides, 564 Chrysanthemi, 563 Clematidis, 563
499
Damnosa, 499
Persies, 499
Cytosporina, 518, 526, 526 Ribis, 526
Cytosporium, 516 Cystothyrium, 531 Cystotricha, 505

Cyttariacese, 134
D
Dacromycetalcs, 395, 396 Dacryodochium, 641
Dacrymycella, 640
Dactylaria, 588
Dactylella, 588
Jackmani, 563 Colchici, 563 Humuli, 563 Inconspicuum, 563 Mori, 249, 525, 562
Dactylium, 588
Dactylopias, 194
Dactylosporium, 616
INDEX
Dadap, 411
Dajdalea, 417, 439
711
Quercina, 439, It.'fi Dahlia, 43, 178, 492, 045 Daisy, 36

Daldinia, 285
Dendrographium, 037 Dendrophagus Globosus, 12 Dcndrophoma, 481, 494, JÔJ^ Convallaria?, 494 Marconii, 494 Valsispora, 494
Dendrostilbclla, 033
Dammara, 489 Damping Off, 77

Dandelion, 71, 119, 378, 590 Daphne, 544
Darluca, 335, 505 Darwinella, 216
Dendryphica;, 009, 015
Dendryphium, 615 Cornosum, 615 Dermatea, 151, 152, /J2

Accrina, 152
Dasyscypha, 135, 144,

Abietis, 145
1I^J^
Carpinea, 152
Cinnamomea, 152
Prunastri, 152
Calyciformis, 145 Resinaria, 145

Subtilissima, 145
Dermateae, 150
Desmazierella, 135 Desmodium, 187, 188
Willkommii, lU, 145 Date, 169, 310 Datura, 48 Daucus, 178 Deconica, 449 Delacourea, 252 Delitschia, 224 Delphinium, 178, 321 Dematiacea;, 565, 594 Key to, 594 Amerosporae, 594, 599 Key to, 599 r Dictyospora), 594, 615 Key to, 615
Didymospora?, 599, 601
Dewberry, 648
Diachnse, 11
Diachora, 216, 217
Onobrychidis, 217
Dialonectria, 201, 205
Dianthus,
101, 312, 488, 507, 619, 654
328, 375, 387,
Diaphanium, 639
Diaporthc, 277, 278, 279, 490, 498 Albocamis, 279
Ambigua, 279, 490

Parasitica, 209
Sarmentella, 279, 490
Strumella, 279, 499

Taleola, 279
Key
to, 601
Heliocospora;, 594
Diatomeae, 3
Diatrypacese, 223, 281
Phragmospora?, 594, 608 Key to, 608

Saturosporae, 594
Scolecosporae, 594, 625
Key
to,
281
Diatrypcae, 281
Dichaenaccae, 160, 162
625 Dcmatium, 217, 600 Prunastri, 495 Dematophora, 230 Glomerata, 201
Key
to,
Dichea, 162
Faginea, 162 Quercina, 162, 16S
Dichcirinia, 353
Dichla-na, 527
Necatrix, 230,
i?Si
Dichomcra, 510
Dendrodochium, 640, 643 Lycopersici, 643
Dicoccum, 601, 602

Rosa;, 602
712
Dicotyledones, 203
INDEX
Diplocladium, 200, 586, 653 Diplococcium, 602, 603 Conjunctum, 603
Diplodia, 243, 510, 511,
511, 511,
Dictyophora, 462, 453 Dictyosporse, 633 Dictyosporium, 615

Dictyuchus, 75
513
Aurantii, 512
Monosporus, 75 Dicyma, 598 Diderma, 10 Didymaria, 586, 587, 587 Didymella, 251, 255, 256
256 Didymiacese, 10 Key to, 10 Didymium, 9, 10
Citri,
Cacaoicola, 512
Dsedalium, 10 Didymochseta, 505 Didymocladium, 586 Didymopsis, 586 Didyraopsora, 342
Didymosphseria, 251, 256, 256 Catalpa;, 256 Epidermidis, 256

Populina, 256
Spheroides, 256 Didymosporse, 633
Didymosporium, 556, 556

Salicinum, 556
Cerasorum, 512 512 Coffeicola, 512 Destruens, 512 Epicocos, 512 Gongrogena, 512 Heteroclita, 512 Macrospora, 511 Mori, 512 Natalensis, 512 Opuntise, 512 Oryzeae, 512 Perseana, 512 Pinea, 512 Rapax, 512 Sapinea, 512 ZejE, 511 Diplodiella, 510, 512, 612 Oryzse, 512
Citricola,
Diplodina, 247, 505, 509, 509

Castanese, 509, 509
Citrullina, 247,
Dieback, 278 Diervilla, 178 Dietelia, 342

Digitalis, 100, 488, 507 Digitaria, 591
Dill,
509
Corticola, 509
Parasitica, 509
Salicina,
609
377
Diplodiopsis, 510
Dilophia, 251, 257, 525, 590
Diplophysa, 73
Diplorhinotrichum, 585
Graminis, 256, 257, 525 Dilophospora, 518, 525, 525

Graminis, 257, 525
Dimargaris, 572
Diplosporium, 586 Dipsacus, 176, 178 Discella, 536, 536

Cacaoicola, 636 Discocyphella, 406
Dimerosporium, 189, 191 Collinsii, 191 Mangiferum, 191

Pulchrum, 191, ini, 625 Dinemasporium, 534, 535, 535 Oryzse, 535
Dinoflagellates, 3
Discomycetes, 116, 117, 123, 159

Discomycopsella, 531 Discomycopsis, 501
Discosia, 531
Pini,
531
Diospyrus, 183
Discula, 534, 535, 535
INDEX
Discula, Platiini, 274, 535
Dispira, 572
713
Drupe, 139, 236, 237, 410, 486

Durio, 221
Dusiella, 211
Ditopella, 263, 264
Ditopa, 264 Fusispora, 264

Doassansia, 301, 315, 322, 322
Gossypii, 323
Dyctilium, 200
E
Earlea spiciosum, 359
Eccilia,
322 Dogwood, 158, 203 Dolichos, 373 Doratorayces, 571 Doscocolla, 645
Niesslii,
450
Echinobotryese, 594 Echinodontiurn, 414, 415
Dothichiza, 534, 535, 536 Ferruginosa, 152
Tinctorium, 415
Populea, 535
Dothidiacea;, 216
Key
to,
216
210
Dothichloe, 210
Aristida;,
Echinodorus, 315 Echinodothis, 199, 211 Tuberiformis, 211 Ectostroma, 657 Ectrogella, 68
Egg
Atramentosa, 210 Dothidea, 216, 218, 220 Noxia, 220 Rosa?, 220 Dothidella, 219, 221 Betulina, 221 Thoracella, 221 Ulmi, 221
Dothidiaceaj, 215
plant, 37, 42, 47, 204, 268, 408, 487, 491, 508, 539, 540, 580, 617
Elaphomycetaceae, 165 Elder, 185, 393 Eleutheromyces, 197
Elm,
71, 127, 130, 182, 221, 249, 260,
393, 421, 430, 437, 455, 484, 489, 503, 530, 544, 557, 603 Emericella, 167
Emmer,
206, 550
Key
to,
216
Dothidiales, 124, 195, 216
Dothiochloe, 199 Dothiopsis, 483
Enchnoa, 262 Endive, 377 Endobotrya, 516

Endoconidium, 146, 639, 641, 641
Dothiora, 155, 156
Temulentum, 642
Endogone, 118 Endomyces, 122
Decipiens, 123
Virgultorum, 157
Dothiorclla, 284, 483, 499
Mori, 499, 499 Populi, 499

Ribis,
Mali, 122, 123, 123

Parasitica, 123 EndomycetaceiB, 120, 122, 165 Key to, 122 Endophyllum, 342, 353 Sempcrvivi, 353 Endothia, 283 Entoloma, 450
499
Dothiorellina, 483, 499

Tankoffii, 499
Double Blossom, 648 Downy Mildew, 82

Dracaena, 270, 487,
489, 497,
503,
553
Draca;nacea;, 303
Entomogenous
fungi, 194
Drcpanospora, 609
Entomophthoracca?, 107
714
Entomophthorales, 66, 102, 107
INDEX
Erysiphe, Cichoracearum, 174, 178,
17S, 569
Key
to,
107
Entomosporium, 243, 531, 532, 532 Maculatum, 149, 532
Graminis, 171, 179, 179, 569 Liriodendri, 178

Martii, 178
MespiU, 150, 532 Thumenii, 532 Entyloma, 314, 320 Australe, 322 Betiphilum, 321 Calendulse, 321 Crastophilum, 321 Ellisii, 321, 321 Fuscum, 322 Irregulare, 321 Lephroideum, 321 Polysporum, 321 Enzymes, 2 Ephelis, 209, 537
EpichlcE, 199, 210, 210, 211
Polygoni, 177, 177, 178, 187 Taurica, 179 Umbelliferarum, 178

Erysipheae, 260
Euascomycetes, 117, 123 Key to, 123

Eubacteriales, 18
Eubasidii, 299 Eubasidiomycetes, 394 Key to, 394
Eucalyptus, 310, 560 Eumycetes, 1, 3, 59
Key
to,
64
Eunectria, 201, 202
Typhina, 210, 643 Epicoccum, 655, 656, 6S6 Hyolopes, 656

Epidochiopsis, 641
Euphorbia, 187, 329, 330, 372, 374, 375

Eurotiopsis, 167, 527
Eusclerotinia, 137
Epidochium, 655, 656 Oryzse, 656 Epilobium, 347 Epochnium, 601 Eremascus, 122 Ergot, 213 Erica, 617
Ericaceae, 143, 159, 186,
Eutaphrina, 127
Eutypa, 278 Eu-type, 328 Excipula, 534

Excipulaceae, 479, 533
to, 533 Hyalodidymae, 533, 537 Key to, 536 Hyalophragmiae, 533, 536 Hyalosporae, 533 Key to, 534 Phaeophragmise, 533, 536 Phaeosporae, 533 Scolecosporae, 533, 536 Key to, 536 Excipularia, 657 Exoascaceae, 125 Key to, 126 Exoascales, 396 Exoascus, 125, 127
Key
397
Erigeron, 89
Erinella, 136
Eriobotrys, 607
Eriocaulacese, 303
Eriopeziza, 135
Eriospora, 518
Eriosporangium, 355 Eriosporina, 515 Eriothyrium, 528

Erysiphacese, 117, 166, 170, 171, 176,
192
Key
Erysiphe,
175,
to,
174
172,
173,
Erysiphales, 332, 475, 494, 569

117,
143, 166,
Exobasidiales, 125, 395, 396
177
Key to, 396 Exobasidium, 396
INDEX
Exobasidium, Andromeda, 396, 397 Azalcae, 398 Cinnamomi, 398 Japonicuin, 398 Lauri, 398 Oxy cocci, 397 Peckii, 398 Rhododendri, 398 Vaccinii, 397 Vexans, 397 Vitis, 398 Exosporina, 656, 656 Laricis, 656 Exosporium, 227, 657, 658 Juniperinum, 560, 658 Laricinum, 658
715
438, 410, 465, 490, 492, 493, 507, 568, 581, 057 Fission, 14
Fistulina, 4-10,
441
Ilepatica, 441,
Fistulinea?,
442
440
Flagella, 14
Flammula, 449, 452 Alnicola, 452 Penetrans, 452 Spumosa, 452

Flax, 69, 343, 653
Floccomutinus, 462 Fodder, 599
Palmivorum, 658, 658 Preslii, 658

Tilia;,
Rubinus, 227 658
Fomes, 417, 418, 663 Annosus, 431 Applanatus, 436, 436 Australis, 437 Carneus, 430 Everhartii, 430
Fomentarius, 426, 429, 429 Fraxinophilus, 433
Fulvus, 433
Fabaceae, 303
Fabrsea, 147, 243
Fulvus
olea;,
433
Hartigii,
434
Malculata, 149, 149, 532

Mespili, 150, 532
Igniarius, 401, 428, 428, 429, 433,
434
Juniperinus, 431
Laricis,
Fagopyrum, 178
Fagus, 188, 545 Farlowiella, 160
Favolus, 417, 439
Australis,
432
Lucidus, 433
Marmoratus, 434
Nigricans, 433
438
Europa?us, 438, 439

Fenestrella, 278
Ferments, organic, 2 Fern, 77, 108, 126, 250, 581, 595

Festuca, 180, 310, 383, 635
Ficaria,
375
436 432 Robinia;, 434 Semitosus, 437 Scssilis, 435 Ulmarius, 437
Pinicola, 435,
Ribis,
Ficus, 205, 249, 267, 278, 533, 544,
552
Fig, 22, 169, 310, 346, 408, 549, 564,
Forget-me-not, 101 Form Genera, 476 Forsythia, 621
572, 573, 604, 624, 626, 643

Filbert,
Fox
Tail,
90
280
145, 162,
192,
Fracchisea, 234
Fimbrystylia, 303
Fir,
132,
203,
276,
Fragaria, 176, 244 Fraxinus, 188

Fruits, 106
320, 347, 349, 418, 424, 431, 434,
716
Fuckelia, 152, 155, 483, 500
Ribis, 500
INDEX
Fusarium, Nivale, 205 Niveum, 651
Fuligo, 11, 12, 200
Oxysporum, 648, 651, 652, 653

Pelargonii, 654
Pestis,
Pini,
Fumago,
Fungi,
1,
191, 616, 624
Camellise, 194, 625
652 654
Classification, 64
Platani, 205, 648
Imperfecti, 64, 475
Key
Slime.
to,
479
Putrefaciens, 649 Rhizogenum, 649

Ricini, 654 Roseolum, 652
True, 1, Funkia, 489

Fusariella,
See Mycomycetes. 2, 59
Roseum-lupini-alba, 662
608
Fusarium, 201, 204, 203, 475, 571, 645, 646 Acuminatum, 652 .^ruginosum, 652 Affine, 652 Aurantiacum, 651 Avenaceum, 206 Blasticola, 654 Brassica;, 654 Cocruleum, 647, 648, 652 Commutatum, 648, 652 Cubense, 649 Cucurbitaria?, 652
Rubi, 648 Rubiginosum, 647, 648 Solani, 204, 585, 648, 652 Subulatum, 648 Tabacivorum, 654 Theobromaj, 648, 654 Udum, 651
Vasinfectum, 205, 650, 650, 651 Pisi, 648, 651
Tracheiphila, 651
Violaceum, 652 Violae, 654
Wilkommii, 648
Fusella, 595
Culmorum,
206, 649, 649
Fusicladium, 253, 602, 606

Betulse, 255,
Decemcellulare, 654
Dianthi, 654
647, 652 Diplosporum, 652 Discolor, 648 Sulphureum, 648 Erubescens, 653 Falcatum, 648 Gemmiperda, 649 Gibbosum, 648 Heterosporum, 206 Hordei, 206 Incarnatum, 654 Limonis, 649 Lini, 653, 653 Lycopersici, 653 Martii, 648 Metachroum, 648 Moniliforme, 650
607
Cerasi, 255, 606
Didymium,
Dendriticum, 253, 255, 607 Depressum, 607 Destruens, 607 Effusum, 607 Eriobotrys, 607
Fagopyri, 607
Fraxini, 255, 606
Lini, 607 Orbiculatum, 255, 607 Pirinum, 253, 607 Saliciperdum, 606
Tremulae, 255, 607

Vanillse,
607
Fusicoccum, 274, 280, 483, 498 Amygdali, 498 Bulgarium, 498 Perniciosum, 281, 498
INDEX
Fusicoccum, Vcronense, 275, 498 Viticoluin, 498, 49S Fusicolla, 639 Fusidium, 201, 203, 5G7, 568 Candidum, 568 Fusisporium solani, 648 Fusoma, 588, 590 Parasiticum, 690
Ginkgo, 267
717
Ginseng, 30, 88, 141, 496, 559, 585,

622, 6,51
Giulia, 532
Gladiolus, 250, 318, 320, 389

Gleditschia, 267
Glcnospora, 590
Gliobotrys, 570
Glioccphalus, 570 Gliocladium, 574
Galera, 449
Agaricinum, 574 Gliodcladium, 572, 574 Globaria, 465

Gla'ocephala, 412
Gloeoporus, 417
Gloeospha;ra, 583
Gallowaya, 339 Gamospora, 517 Gamosporolla, 483 Gaphiothecium, 630 Garden-pea, 408
Garlic, 97
Gloeosporium, 147, 252, 264, 266, 267, 269, 274, 475, 478, 538, 539, 547
Affine,
544
Gasteromycetes, 395 Gaylussacia, 397

Geaster, 465
Alborubrum, 544 Alle^cheri, 545 Alneum, 545
Gelatinosporium, 518 Geminispora, 263
Ampelophagum, 541 Amygdalinum, 542

Anthuriophilum, 544 Apocryptuni, 545
Aquiligia?,
Gemmse, 60
Gentiana, 352, 389
Geoglossaceae, 131, 154 Key to, 131
Geoglossese, 131
544
Atrocarpi, 273, 541

Begonia?, 644
Bcrberidis, 546 Betularuni, 545
Gcoglossum, 131 Geotrichum, 568 Geranium, 52, 315, 390, 591, 603, 631, 654 Gherkin, 247 Gibbellina, 251, 256 Cerealis, 256 Gibellula, 634 Gibbera, 234 Vaccinii, 234, 235 Gibberella, 198, 206, 646 Cerealis, 207 Moricola, 207 Saubinettii, 206, 206 Gibberidca, 234 Gibelia, 283 Ginger, 77
Betulinum, 646 Beyrodtii, 544 Bicolor, 541 Bidgoodii, 544 Bruneum, 544 Cactorum, 544
Canadense, 646
Carpini, 545
Carya;, 545
Cattleya?, 544
Caulivonim, 543 Cinctum, 541 Cingulatum, 268, 541 Citri, 541 Clematidia, 644
718
Gloeosporium, Coffeanum, 543 Concentricum, 544
INDEX
Gloeosporium, Pestis, 543 Piperatum, 269, 541
Platani, 274
Psidii, 271,
Coryli, 545
Crotonis, 544
641
Curvatum, 542
Cydonise, 542 Cylindrospefmum, 641 Cytisi, 544 Depressum, 541 Dianthi, 544 Diospyri, 542 Elastica;, 267, 544 Euphobiae, 544 Fagi, 545 Fagicolum, 544 Fragaria, 542 Fructigenum, 267, 539 Helicis, 544
Quercinum, 545 Rhododendri, 544 Ribis, 148, 541 Rosse, 544

Rufomaculans, 265, 2S5, 267, 539 Saccharini, 545 Salicis, 148, 541 Soraurianum, 544 Spegazzinii, 541 Stanhopeicola, 544 Tamarindi, 546 These, 544 Sinensis, 544 Tiliaceum, 546 TiliB, 546 Tremulae, 545 Trifolii, 543 Umbrinellum, 544 Vanilla;, 280, 541 Variabile, 542 Venetum, 542 Versicolor, 267, 539 Violae, 544 Glcesporiella, 555 Glomerella, 263, 264, 475, 539, 547 Atrocarpi, 273, 541 Cactorum, 265 Cincta, 269, 270, 541, 547 Cingulata, 268, 541
Gossypii,
267,
Hendersonii, 541
Inconspicuum, 544 Intermedium, 541 Juglandis, 545 Kawakami, 545 Laeliae, 544 Laeticolor, 267, 539 Lagenarium, 543 Macropus, 253, 541
Malicorticis, 493, 542,
6^2
Mangiferaj, 543
Manihotis, 543
Medicaginis, 543
Melengonea, 539 Mezerei, 544 Minus, 543 Morianum, 543 Musarum, 542
271,
272,
273,
547
Piperata, 269, 270, 271, 541
Psidii, 270,
Importatum, 642 543 Nanoti, 646 Nervicolum, 545 Nervisequum, 274, 641 Olivarum, 543 Opuntiae, 544 Pallidum, 544
Myrtilli,
541
Rubicolor, 270, 547
Rufomaculans,
541, 548
264,
265,
266,
267, 268, 269, 270, 271, 272, 539,
Paradoxicum, 157, 541 Pelargonii, 544
Cyclaminis, 268 Glomerularia, 566 Gloniella, 161
Glonium, 163
INDEX
Glumacese, 199
Glutinium, 481, 630 Clyceria, 383
Glycophila, 567 Glycyrrhiza, 187
719
Graphiola, 323, 663

Phcenicis, 664, 665 Graphiothccium, 243 Graphium, 630
Grass,
211,
8,
11,
24,
69,
73,
209, 210, 260,
Gnomonia, 264, 274, 529, 539

Carya^, 545
213,
220,
259,
303,
310, 312, 320, 321, 329, 333, 379,
Erythrostoma, 275 Leptostyla, 275, 555 Oryza?, 276 Padicola, 275, 496
383, 385, 405, 508, 520, 612, 635,
661
Green
Algse, 3 Grossularia;, 493
Qucrcus Illicis, 275 Rubi, 276

Veneta, 274, 274, 275, 498, 535, 541, 546
Guava, 271, 541 Guelichia, 641

Guignardia, 236, 237, 484
Gnomoniacea?, 223, 263 Key to, 263

Gnomoniclla, 263, 273, 539 Coryli, 274
Fimbriata, 274
Ampelicida, 238 Bacca;, 242

Bidwellii,
238,
238,
239,
241,
484, 490
Buxi, 243 Thea;, 243

529,
Vaccinii, 242, 242 Gymnoascacese, 105 Gymnoascus, 164
Tubiformis, 541
274,
274,
Gnomoniopsis, 264, 265 Godroniella, 534 Golden Rod, 179 Gonatobotrytidiae, 566 Gongromeriza, 596
Gooseberry, 141, 148, 155, 185, 245, 351, 433, 504, 519, 526
Gymnoconia, 355, 359, 390 Interstitialis, 360, 360
Gymnosporangium,
362, 391
330,
355,
361,
Biseptatum, 370
Clavaria;forme, 331,
363, 366,
Gordonia, 150
Gorgonicepes, 136
366
Clavipes, 363, 368
Gossypium, 168 Gourd, 95, 247

Grain, 257, 260, 333, 384, 451, 508,
599, 620 Grallardia, 176, 178
Cornutum, 308, 371

Ellisii,
369
Germinale, 368
Graminea;, 143, 179 Grandinia, 413

Granularia, 640
Globosum, 363, 366 Japonicum, 371 Juniperinum, 367

Juniperi-virginianaj,
329,
363,
364
Nelsoni, 363, 370
Grape, 36, 37, 43, 51, 52, 73, 141, 148,

153, 192, 201, 231, 242, 249, 250,
Nidus-avis, 363, 369
253, 258, 267, 268, 281, 345, 398,

405, 463, 484, 486, 490, 491, 494,
498, 504, 506, 517, 520, 540, 541,
Sabina, 369
Terminali-juniperinum, 371 Transformans, 369
549,
554,
560,
579,
580,
620,
Yamadaj, 371
Gyroceras, 596
626
720
INDEX
Helminthoaporium, Sorokinianum, 613 Teres, 612, 613 Thea;, 614 Trichostoma, 612 Tritici, 613 Tritici Repentis, 262, 613 Turcicum, 613, 614 Vaccinii, 234 Helostroma, 634 Helotiacea;, 133, 134 Key to, 135 Helotieae, 136
Helotium, 136
Helvellacea?, 131
Helvellales, 123, 130
Hackberry, 93 Hadrotrichum, 600 Haematomyces, 151
Haematomyxa, 151
Hainesia, 538
Halobyssus, 567
Hamamelis, 188 Haplaria, 575 Haplariopsis, 586

Haplobasidium, 597
Haplographiea;, 595, 600
Key to, 600 Haplographium, 601

Haplosporella, 501
Haplotrichum, 571
Hariotia, 163
Key
to,
131
Harknessia, 501
Hemiascomycetes, 114, 117, 118 Hemibasidii, 298, 299
Harpocephalum, 630 Harpographium, 630 Hartigiella, 570, 575 Laricis, 570 Hartigiellese, 566, 570
Hemiglossum, 131 Hemileia, 354, 355 Oncidii, 356

Vastatrix, 355, 355
Hawthorn,
38, 39, 130, 502,
525
Woodii, 356 Hemi-parasites, 2
Hazel, 122, 203, 249, 274, 280, 444
Hebeloma, 449
Hedera, 249, 487, 522, 541, 553 Helianthus, 92, 178, 179, 388
Helicobasidium, 403
Heliomyces, 445 Heliscus, 645

Hellebore, 505
Helminthospora, 609 Helminthosporium, 260, 564. 609, 611 Avenaj, 613 Bromi, 261, 613 Gramineum, 261, 612, 612, 614 Hevea?, 614 Iberidis, 614
614 Inconspicuum, 613, 614 Lunarise, 614 Oryzae, 614 Sativum, 613, 613 Sigmoideum, 613
Inajqualis, 614,
Hemi-saprophytes, 2 Hemi-type, 328 Hemlock, 418, 419, 423, 435, 436, 438 Hemp, 52, 101, 141, 486, 494, 521, 58J Hendersonia, 257, 264, 516, 616 Acicola, 515 Coffeae, 515 Cydonae, 515 Foliicola, 516 Mali, 515 Notha, 516 Oryzae, 516 Piricola, 515 Togniniana, 515, 616 Hendersonula, 515, 516 Morbosa, 219 Henriquesia, 160 Hepatica, 93, 357 Heptameria, 252 Heracleum, 591 Hericium, 413
INDEX
Herpotrichia, 226, 229, 230
721
Nigra, 229
Heterobotrys, 595 Heterocephalum, 634

Heterocontae, 3 Hetercecism, 64, 329
Heteropatella, 534 Heterosphaeria, 155
Horse Chestnut, 445, 460, 524 Horse Radish, 95, 96, 506, 522, 582, 590, 619, 629 Hoya, 544
Humulus,
176, 178
Hyacinth, 27, 42, 44, 75, 143, 231, 260, 603

Hyaloceras, 558
Heterosporiuin, 609, 610

Auriculi, 611
Hyalodema, 593 Hyaloderma, ISO

Hyalodothis, 217
Echinulatum, 611 Gracile, 611 Laricis, 611 Minutulum, 611 Ornithogali, 611 Syringa?, 611 Variabile, 611
Heterotheca, 408 Heuchera, 188
Hyalopus, 570 Hyajospora, 341 Hydnaceae, 402, 413 Key to, 413 Hydnochaete, 414
Hydnum,
414, 414 Diversidens, 415
Hevea, 207, 278, 287, 411, 418, 437,

513, 544, 553, 607
Erinaceus, 414, 414 Schiedermaycri, 415

Septentrionale, 414 Hydrangea, 347, 488, 493, 522 Hygrophorese, 442 Hymenium, 113 Hymenochajtae, 406
Hexagonia, 417 Heydenia, 630 Hiatula, 450

Hibiscus, 488
Hickory, 428
Hicoria, 396
Noxia, 411
Himantia, 657 Holchus, 383

Hollyhock, 328, 386, 487, 492, 523, 552, 630 Hoist iella, 280
Hymenogast rales, 396 Hymenomycetes, 394

Hymenopsis, 655
Hymenoscypha,
136, 146, 146
Holwaya, 151 Homostegia, 216 Honey dew, 190 Honeysuckle, 36, 191 Hop, 8, 36, 93, 175, 279, 486, 490, 5G3, 569, 590, 611 Hordeum, 180, 317, 379 Hormiactella, 601 Hormiactis, 586 Hormiscium, 596 Hormodendrum, 600, 631 Cladosporioides, 248
Hordci, 601, 601
Temulenta, 146, 642 Hymenula, 640 Hypha, 657

Hyphaene, 193
Hyphoderma, 575 Hypholoma, 449, 450

Appendiculatum, 451, 4^1 Fasciculare, 451 Lateritium, 451 Hyphomycete, 321, 402, 663 Hyphostereum, 640
Hypocenia, 501 Hypochnaceae, 402 Key to, 403
Hornbeam,
152, 274
Hypochnus, 403, 406 Cucumeria, 404
722
Hypochnus, Filamintosus, 404 Fuciformis, 404 Ochroleucus, 403 Solani, 404 Theae, 404
Hypochytriaceae, 67
INDEX
Hysterium, 164 Hysteroglonium, 163 Hysterographium, 164
Fraxini, 164, 164 Hysterostomella, 163
Hypocopra, 224 Hypocrea, 209, 199 Ceretiformis, 209 Sacchari, 209

Hypocreaceae, 196 Key to, 196
Iberis,
Ilex,
614
188
Illosporium, 641, 643
Hypocreales, 124, 195, 584 Hypocreeaj, 197
Key
to,
198
Maculicola, 644, 644 Malifoliorum, 643 Impatiens, 93, 176
Hypocrella, 199
Hypocreodendron, 527
Hypocreopsis, 199
Inocybe, 449 Ipomeae, 337

Iris, 27, 11, 46, 73,
389, 507, 514, 522
Hypoderma, 161
Desmazieri, 161
Laricis, 161
608,611,619,620
Irish Potato, 105, 106 Irpex, 414, 415
Pinicola, 161
Destruens, 415
Flavus, 415, 415
Fusco-violaceus, 416
Strobicola, 161
Hypodermataceae, 160 Key to, 160
Hypodermella, 160, 161

Laricis, 161
Paradoxus, 415 Isaria, 196, 634, 635 Fuciformis, 635
Sulicigena, 161
Hypodermium,
538, 547
Orchidearum, 547 Hypolyssus, 405

Ilypomyces,, 197, 200, 200 Hyacinthi, 200
Solani,
Graminiperda, 635 638 Griseola, 637 Isothea, 276 Itajahya, 462

Isariopsis, 637,
Ivy, 152, 493, 497, 544
200
Hypomyceteae, 196, 197 Hyponectria, 201 Hyponectrieae, 196

Hypospila, 276 Hypoxyleae, 285
Jansia, 462 Johnson Grass, 311
Hypoxylon, 285
Hysteriaceae, 160, 163, 530 Key to, 163
Hysteriales, 124, 159
Juglans, 186, 275, 396, 507 Juncus, 303 Juneberry, 38
June Grass, 578

Juniper, 52, 157, 162, 220, 230, 243, 330, 560
Key
to,
160
Hysteriopsis, 161
Brasiliensis,
Juniperus,
161
362, 365, 366, 369, 370, 371, 516, 610
367,
368
INDEX
Lasiospha^ria, 226
723
Lasmenia, 531
Kalmusia, 277
Latcrnca, 463, 464
Kawakamia,
83,
89
Columnata, 464, ^C4

Lathyrus, 217, 313, 372 Laurel, 492
Laurel, Cherry, 409
Cypcri, 89 Kcithia, 156

Kc'llermania, 513 Klachbrennera, 4G4 Kinetia, 640 Kncifficlla, 413
Laurus, 398, 489
Kuchneola, 355, 361 Gossypii, 361 Uredinis, 361
Lccythium, 198 Legume, 28, 31, 32, 313, 373 Lemahs, 521, 534 Lembosia, 163
Lemon,
77, 494, 508, 510, 512, 518, 540, 549, 574, 582, 604
Lcmonniera, 593
Lentinus, 445
Conchatus, 446
Laboulbeniales, 124
Labrella, 529, 530
Coryli, 529, 530
Lepideus, 446,
Variegata, 446
^6
530 Laburnum, 544

Piricola,
Lentomita, 232 Lenzites, 417, 439

Abietina, 440
Betulina, 440,
Lachnclla, 135, 145, Pini, 145

Lachncllula, 135
11^5
Ul
Corrugata, 440
Sepiaria,
440
Lachnocladium, 412 Lachnodochium, 641
Variegata, 440
ViaUs, 440
Lachnum, 136
Lactarica), 423,
443
Leocarpus, 11 Lepiderma, 10
Lepidonectria, 201
Lcpotia, 450 Leptoglossum, 443 Leptomitaceae, 75
Laestadia, 238
Buxi, 243
Lamyella, 483 Langloisula, 576

Larch, 145, 161, 162, 249, 348, 419,
424, 432, 436, 438, 530, 570, 611,
Lcptonia, 450
Lcptopus, 443
Leptospha;ria, 252, 257, 519, 660
Circinans, 258
656, 658
Larix, 230, 344
Larkspur, 43 Laschia, 417

Lasiobotrys, 189, 191 Lonicera), 191
Lasioderma, 634
Lasiodiplodia, 510, 613
Thoobromff, 513
Tubericola, 513 Lasionectria, 201
Coniothyrium, 257, 258, 504 Herpotrichoides, 258 Iwamotoi, 258 Napi, 258, 616 Phlogis, 258, 519 Rhododendri, 258 Sacchari, 258 Stictoides, 258 Taxicola, 259
724
Leptosphseria, Tritici, 258, 520
INDEX
Lilac, 88, 404, 488, 581, 582, 611
Liliaceai, 310, 318,
Vagabunda, 259
258 Leptospora, 226 Leptostroma, 529, 530 Larcinum, 249, 530 Piricola, 530 Punctiforme, 530
Vitigena;,
320
Lilium, 375
Lily, 106, 141, 250, 488, 563, 579, 580,
592, 631
Lily-of-the-valley, 523, 581
Limacinia, 190
Leptostromataceae, 479, 528
Key
to,
528
Hyalodydimse, 528 Hyalosporse, 528 Key to, 528

HyalophragmiEe, 528, 531 Key to, 531 Phseodidymse, 528 Phaiopharagmise, 528
Phseosporse, 528, 531
Tangensis, 193 Lime, 203, 249 Limnanthemum, 315 Linaria, 168
Linden, 259, 421, 545, 560 Linospora, 276
Linum, 607
Liriodendron, 188, 258, 547 Lisea, 197
Lisiella,
197
Listeromyces, 657
Lizonia, 226 Lobelia, 492
Locellina, 449
Key
to,
531
Scolecosporae, 528, 532
532 Leptostromella, 532, 533 Elastica?, 533, 533
Key
to,
Loculistroma, 199, 216
Bambusa;, 215
Locust, 419, 434
Leptothyrium, 274, 528, 529 Acerinum, 529 Alneum, 274, 529

Buxi, 529
Locust Black, 438 Lolium, 73, 383

Lonicera, 186, 191, 529
Macrothecium, 529 Oxy cocci, 529, 529 Parasiticum, 529 Peonse, 529 Periclynieni, 529 Pomi, 529 Lepto-type, 328
Lespedeza, 187
Lettuce, 36, 37, 44, 52, 95, 141, 142,
408, 507, 522, 555, 556, 579, 620 Levieuxia, 501
Lopharia, 413 Lophiostomatacese, 223 Lophium, 164
Lophodermium, 161
Abietis, 162
Brachysporum, 162 Gilvum, 162 Juniperinum, 162 Laricinum, 162
Levisticum, 28
Libertella, 562, 564
Rubra, 208, 564 Ulcerata, 564

Libertiella,
527
Macrosporum, 162, 162 Nervisequum, 162 Pinastri, 161, 162 Loquot, 553 Lunaria, 614 Lupine, 560, 564, 652 Lupinus, 43, 178, 313
Albus, 168
Augustifolius, 168
Liceaccaj, 9
Lichenopsis, 515
Lichens, 134
Luteus, 168
INDEX
Lupinus, Thermis, 168 Luzula, 303
Lycopordaceac, 464
725
Key
to,
464
Lycoperdales, 305, 464
Lycoperdon, 465
Macrosporium, Nobilo, 619 Porri, 618 Ilamulosum, 619 Rugosa, 624 Saponaria;, 620 Sarciniformo, 619
Jf65
Ciommatum, 465,
Lycopersicum, 178 Lysurus, 463
Sarciiuila-parasiticuni,
618
M
Macrodendrophoma
494
Macrobatis, 514 Macrodiplodia, 510
Salicicola,
Tabacimim, 619 Tomato, 624 Uvarum, 620 Vcrrucosum, 620 Viola, 620 Maorostilbum, 634
253,
Madia, 92 Magnolia, 188, 503, 559 Magnusia, 166

Magnusiella, 126
Macrophoma,
Abietis,
284, 481, 493
493
Curvispora, 493, 493
Dalmatica, 493
Hclicinia, 493
Maguey, 220 Mahonia, 379 Malbranchea, 567 Mal-di-gomma, 649 Malope, 386
Malus, 371 Malva, 386
Malvaceae, 507
Hennebergii, 493
Ligustica, 493
Malorum, 493
Manihotis, 493
Reniformis, 494 Taxi, 493
Vestita, 493
Macrosporium, 616, 618 Aductum, 619 Alliorum, 618 Brassica;, 619 Catalpa?, 619 Cheiranthi, 619 Cladosporioides, 620 Commune, 260, 618
Mamiania, 263 Mandarin, 520 Mangel, 41, 645 Mango, 191, 193, 543 Mangold, 491, 581 Manihot, 557 Maple, 72, 130, 152,
159,
182, 203,
419, 421, 428, 430, 436, 455, 489,
524, 525, 545, 557, 563, 632
Marasmiea;, 443, 445 Key to, 445
Cucumerinum,
619, 620
Fasciculata, 624
Gramineum, 620
Herculeum, 618 Iridis, 619 Longipcs, 619 Lycopersici, 624
Marasmius, 445, 446 Equicrinus, 448 Ilawiiensis, 448 Plicatus, U7, 448 Sacchari, 448 Sarmentosus, 448 Semiustus, 448
Marchalia, 156
Marchaliclla, 189
Macalpinianum, 620 Nigricanthium, 619
Marssonia, 147, 274, 556
726
Marssonia, Castagnei, 157, 555 Juglandis, 275, 555
Martini, 555 Medicaginis, 556
INDEX
Melampsora, Saxifragarum, 346 Melampsoraceae, 335, 340 Key to, 340
Melampsorella, 341, 348, 390, 391 Elatina, 348
Panattoniana, 555
Perforans, 555
Populi, 555
Potentillaj,
Melampsoridium, 341, 347, 3^7, 391 Betulse, 348

Melampsoropsis, 341, 349, 350, 391 Rhododendri, 349 Melanconiaceaj, 537
to, 537 Hyalodidymae, 538, 555 Key to, 555 Hyalophragmia;, 538, 556 Key to, 556 Hyalosporse, 538 Key to, 538 Phseodictya?, 537, 561 Phaeodidymaj, 537, 556 Key to, 556 PhseophragmiP, 537, 557 Key to, 557 Phaeospora;, 537, 553 Key to, 553 Scolecosporae, 537, 561 Key to, 562 Staurosporse, 537 Melanconiales, 265, 525, 537, 564 Melanconidacese, 223, 279 Key to, 279 Melanconiella, 279 Melanconiopsis, 501 Melanconis, 279, 281 Modonia, 281, 498, 560 Melanconium, 553, 554
555
RosiB, 555
Secalis,
555
Viola?, 556,
556
Key
Martensella, 576
Martindalia, 633 Massalongiella, 236
Massaria, 263
Theicola, 263 Massariaceae, 223, 262
Key
to,
262
Massarina, 263
Massariovalsa, 263
Massospora, 566 Mastigosporium, 588, 590 Mastomyces, 155, 514 Friesii, 514 Matrouchotia, 403 Mattirolia, 198 Mazzantia, 216 Medicago, 178 Medick, 148 Medlar, 140, 150, 569 Megalonectria, 198 Melanconiales, 479, 527 Melampsora, 340, 342, 390 Allii-fragilis, 344 Allii-populina, 344 Allii-salicis albae, 344 Bigelowii, 344 Klebahni, 344 Larici-pentandra?, 344 Larici-populina, 344 Lini, 342 Medusae, 343 Pinitorqua, 344 Repentis, 344 Ribesii-viminalis, 344 Rostrupii, 344
Fuligineum, 554, Pandani, 554

Sacchari, 554
554-
Melanomma, 227, 232 Glumarum, 232

Henriquesianum, 231 Melanops, 283, 284, 503 Melanopsamma, 227 Melanopsichium, 302
Melanospora, 196, 197, 200, 201 Damnosa, 200
INDEX
Mclanospora, Stysanophora, 201 Melanosporea;, 196, 197 Melanostroma, 538 Mclanotitnium, 314 Mclasmia, 158, 529, 530 Acerina, 159, 530
Punctata, 630
Salicina,
IMelica,
727
Microdiplodia, 510
Anthurii, 511 Microglossuni, 131
Micropera, 518 Microspatha, 634

Microspha,'ra, 175, 185
Alni, 185, isa,
570
530
497
Melilotus, 508
iMeliola, 189, 190, 191, 193, 193,
024
Calocladophora, 106 Extensa, 186 Lonicera?, 186 Vaccinii, 186

Bcrbcridis, 185
Beta;,
Camellia?, 193, 193, G25
Niesslcanca, 194
Ponzigi, 194
187
Diffusa, 186
IVIelogramma, 283
Hcnriquetii, 284
Elevata, 186 Euphorbia;, 187

Fcrruginea, 187
Grossularia-, 185, 185
Mclogrammatacea;, 223, 282 Key to, 283

Melon, 10, 27, 487, 608, 621, 629 Melophia, 532
Mentha, 178
Merasmieaj, 443, 446 Key to, 445
Microstroma, 396 Album, 396 Juglandis, 396

Microthyriacea?, 170, 195
Merasmiopsis, 445 Mercurialls, 344

Meruliea?, 410, 418
Microthyrium Coffa;, 195 Micro-type, 328

Micula, 518
Mcrulius Lacrymans, 418 Mesospore, 327, 375, 384 Mespilus, 570 Metanectria, 198 Mctasphseria, 252, 257 Albescens, 257 Michenera, 405 Micothyriaceai, 170 Microascus, 166 Microcera, 207 Micrococcus, 18, 21 Albidus, 21 Delacourianus, 21 FlaviduR, 21 Imperatoris, 21 Nuclei, 21 Pellucidus, 21 Phytophthorus, 21 Populi, 21 Tritici, 21
Mignonette, 81, 631 Mikronegeria, 336 Millet, 90

Milowia, 588
Mitrula, 131, 132
Sclerotiorum, 132
Mohonia, 379
Molds, Slime. Mollisia, 146
Sec Myxomycctes.
Mollisiacea;, 134, 146
Key
to,
146
Mollisiclla, 146
Monacrosporium, 588
Monascacea;, 118
Monilia, 137, 138, 140, 567, 668
Cinerea, 139, 569 Crategi, 669
Finiicola,
669
Fructigena, 139, 569
Laxa, 569
Linhartiana, 569
728
Monilia, Seaveri, 140, 569 Moniliaceaj Scolecospora;, 592
Moniliaceae, 565
INDEX
Mucor, Pyriformis, 106 Racemosus, 106
Mucoracea;, 103, 107 Key to, 104
Kry
to,
565
Amt^jsporse, 565 Key to, 565

Chromosporiea}, 565 Key to, 566
Dictyospora;, 565, 592
Mucorales, 66, 102 Key to, 103
Mucoreae, 104 Mucronella, 413
Mucrosporium, 589
Mulberry, 21, 31, 43, 52, 73, 393,
445, 454, 499, 557, 561, 582, 626,
Key
to,
592
585
Didymosporffi, 565, 585
Key
-
to,
658
MuUerella, 236 Munkia, 527 Munkiella, 217
Muricularia, 482, 527
-Helicosporse, 565, 593

Oosoporeffi, 565, 567
Key
-
to,
567
Phragmosporse, 565, 588 Key to, 588 Staurospora?, 565, 593 Key to, 593 Moniliales, 464, 479, 554
Muscari, 375
Mushroom,
584, 587
200, 398, 567, 569, 574,
Muskmelon,
44, 51, 95, 247,
487
Key
to,
465
Monilochajtes, 596, 597

Infuscans, 597
Pachyspora,
Monoblepharidiales, 66 Monochaetia, 558
Mutinus, 462 Mycelia Sterilia, 479, 659 Key to, 659 Myceliophthora, 566, 567 Lutea, 567
558
Monocotyledones, 611 Monographus, 217 Monopodium, 576
Mycelophagus Castaneae, 101 Mycena, 450, 460, 461 Epipterygia, 460 Mycenastrum, 465
Mycogala, 481 Mycogone, 200, 586, 587, 587 Perniciosa, 200, 587 Rosea, 200, 587 Mycoplasm Theory, 333
Mycospha;reIla, 236, 243, 484, 490,
519, 525
Abietis,
Monospore, 121 Monosporium, 576 Monotospora, 600

Monotosporeae, 595, 600
Key
to,
600
Montagnella, 216 Montagnites, 442 Moon Flower, 82 Morel, 114
249 484
Brassicaecola, 249,
Morning Glory,
Morus,
491,
182,
82,
337
207,
512,
Cerasella, 245, 625
Mortierellacese, 103
Cinxia, 250
231,
517,
202,
503,
249,
525,
Citrullina, 246, 2^8,
509
499,
562
Mountain Ash, 39, 367, 368, 427 Mucor, 90, 101, 104, 105, 106 Mucedo, 106
249 Coffeieola, 250 Comedens, 249 Convexula, 250 Cydoniae, 249

Coffeae,
INDEX
MycosphsDrella, Elasticac, 249
FaKi, 249
Fragariac, 244, 244, 519,
729
Mystrosporium, Alliorium, 620 Myrothccium, ()55
590
iMytilidium, 164
Fusca, 250
Gibelliana, 249
Myxobacterialcs, 11 Myxogastralcs, 5, 9
Gossypina, 248, 625 Grossulariae, 246

Hedericola, 249
Key Key
to,
9
1, 3,
Myxomycetes,
to,
Hondai, 250
Laricina, 249, 530
Locfgreni, 249
Myxormia, 538
Myxosporella, 538
Maculiformis, 249, 485, 562 Mori, 557

Morifolia, 249, 562
Pinifolia,
Myxosporium, 274, 538, 546 Abietinum, 547
Cameuni, 547
Corticolum, 493, S46, 546
249
Devastans, 547
Lanceola, 547
Pinodes, 250, 506

Populi, 249, 519, 635 Primulse, 250
Punctiformis, 219
Rosigena, 249
Rubina, 245
Sentina, 246, 246, 247, 249, 519
Shiraina, 250 Stratiformans, 248
Tabifica, 247, 485, 490
Longisporum, 547 Mali, 547 Piri, 547 Valsoideum, 274, 546 Myxotrichelleae, 595
N
Naemosphaera, 501
Najrnospora, 122, 538, 547, 562
Tamarindi, 250 Taxi, 249 Tulasnei, 247, 603 Ulmi, 249, 484 Vitis, 249
Mycosphaerellaceae, 223, 236
Ampelicida, 238 Coryli, 122

Crocea, 547 Napicladium, 609, 611 Janseanum, 611 Soraueri, 255, 611
Narcissus, 389, 489, 523, 591, 592, 611
Key
to,
235
IMykosyrinx, 302 IMyrangiclla, 170

Orbicularis, 170
Myrangium, 170
Myriangiacea?, 165, 170 Key to, 170
Nasturtium, 37, 168, 260 Naucoria, 449 Necator, 640, 643

Decretus, 643 Necrosis, 281
Nectarine, 604
Nectria, 197, 201, 475, 646
Myriogenospora, 216 Myriostoma, 4G5 Myrmseciclla, 283 MyrmEBcium, 283
Myroiphysa, 655 Mystrosporium, 616, 620 Abrodens, 620 Aductum, 620
Bainii,
Amerunensis, 204 204 Bogorionsis, 205 Bulbicola, 205
Cinnabarina, 202, 202, 642
730
Nectria, Coffeicola, 204
INDEX
Neovossia, 314, 315 Nephlyctis, 354
Nicotinia, 101, 168, 178, 486
Nidulariales, 396
Niesslia,
Cucurbitula, 203
Ditissima, 203, 568
Diversispora, 204
Fruticola, 205
225
Gigantispora, 205
Goroshankianna, 205 Graminicola, 205 Ipomoea;, 204, 204, 205 Jungeri, 205 Luteopilosa, 205 Pandani, 204 Ribis, 204
Rousselliana, 204, 656
Solani, 204
Nigrospora, 600 Niptera, 147

Nitschkia, 234
Nolanea, 450
Nothopatella, 501
Nowakowskiella, 72 Nummularia, 285

Discreta, 285, 286 Nyctaginaceae, 303
Striatospora, 205
Theobromae, 205 Theobromicola, 205 Vandffi, 205 Vanillse, 205 Nectriacese, 196
Nectriese, 196
O
Oak, 130, 152, 157, 162, 177, 186, 192,
193, 203, 220, 231, 249, 264, 275,
279, 281, 352, 396, 409, 410, 411,

414, 415, 419, 421, 422, 424, 428, 430, 434, 436, 439, 440, 442, 452, 489, 498, 545, 546, 547, 555, 564,
570, 608
Key
to,
197
Nectriella, 197
Nectrioidaceae, 479, 526

to,
Key
526
Oat, 23, 206, 213, 260, 301, 304, 305,

306, 380, 383, 490, 520, 550, 607,
Hyalodidymise, 526
Hyalophragmiae, 526 Hyalosporse, 526 Key to, 526 Olluleae, 526 Phaeosporse, 526
Scolecosporse, 526
Zythiese, 526
608
Oat Grass, 307

Ochropsora, 336 Sorbi, 336 Odontia, 413
Odor^toglossum, 631 CEdemansiella, 444
Negeriella, 637
Negundo, 260, 489 Nematospora, 121 Nematosporangium, 75 Nemophila Auriculata, 168 Neobarclaya, 556
Neocosmospora, 197, 205, 475, 646,
651
Neolecta, 131
ffidemium, 598 CEdocephalum, 570

(Enothera, 71 Oidiopsis, 567
Neomichelia, 609 Neopeckia, 226 Neottiospora, 482
Oidium, 60, 172, 567, 569 Alphitoides, 570 Ambrosia;, 178, 569 Balsamii, 177, 569 Chrysanthemi, 569 Crsetaegi, 183, 569 Erysiphoides, 569 Farinosum, 184, 569
INDEX
Oidium, Fragariic, 175, 569 Lcucoconium, 176, 569 Mcspilinum, 570 Monilioidcs, 179, 569 Qucrcinum, 570 Tabaci, 570
Tuckeri, ISl, 669
Verbena;, 570
731
Okra, 650, 651 Oleaceae, 164

Oleander, 36, 45, 192, 193, 422, 524, 631
Oleina, 122
Olive, 34, 45, 155, 192, 193, 433, 486,
Ophiocladium, 566, 567 Ilordii, 567, 567 Ophiodothis, 216 ()l)hiorna.s.saria, 262 Oi)hionectria, 198, 207 Coccicola, 207 Foliicola, 207 Ophiotrichurn, 609 Opsis-type, 328 Opuntia, 544
Orange, 207, 249, 256, 260, 409, 422, 435, 445
Orbicula, 189
Orcadellacesc, 9
493, 543, 602, 624
Orchard Grass,
52,
550
Olopccurus, 383
Olpidiacea, 67, 69 Key to, 68
Olpidiopsis, 68
Orchid, 46, 52, 205, 270, 392, 500,

541, 544, 547, 631 Orchis, 344
Olpidium, 68, 69, 72 Brassicae, 68, 69

Olpitrichum, 575 Onibrophilea;, 136
Ornithogalum, 71, 320 Osage Orange, 346

Ostreion, 164
Ostropacese, 160
Ostrya, 188
Otthia, 234 Ovularia, 243, 577, 582, 582
Alnicola, 582
Omphalia, 450 Oncidium, 356, 392, 544, 605 Oncopodium, 615 Oncospora, 537
Onion, 41, 42, 43, 52, 97, 200, 377,
491, 497, 499, 512, 520, 541, 549, 574, 581,
604,
606,
ArmoracitE, 582
Canaigricola, 582
Citri,
582
616,
618,
Corccllensis, 582
620
Onobrychis, 168 Onj'genacese, 165
Oochytriacea?, 67, 75
Exigua, 582
Interstitialis,
582
Medicaginis, 582
Key
to,
73
65, 66, 101
Oomyces, 199 Oomycetes, 62,
Oospora, 475, 567, 568, 568 Abietum, 568

Scabies, 568
Necans, 582 Primulana, 582 Rosea, 582 Syringa^, 582 Vicise, 582
Villiana,
582
Ovulariopsis, 188, 577, 682
Ophiobolus, 252, 259, 259
Ulmorica, 682
Oxalis, 168, 329,
Graminis, 259
Herpotrichus, 269
Oryzese, 259
384
Ozier, 253
Ophiocoras, 232
Ophiochaita, 252
Ozonium, 657, 661 Onanivorum, 662
732
INDEX
502, 515, 519, 530, 540, 546, 547,
Pachybasium, 583 Pachysterigma, 403 Pactilia, 639 Pajonia, 176, 178, 352 Paepalopsis, 567
Palm, 77, 88, 191, 323, 499, 545, 552, 560, 658, 664 Palmetto, 412 Pandanus, 204, 531, 554 Panicum, 305, 307, 310, 312, 314 Pansy, 99, 320, 488, 552, 654 Panus, 445, 446 Stipicus, 446 Papaver, 321, 322 Papulospora, 570 Paranectria, 198 Para Rubber, 101, 415, 487, 512, 614 Paraspora, 588
Parmularia, 163 Parodiella, 189
Parsley, 141, 377, 521
553, 607 Pearl Millet, 90
Pecan, 250, 607, 632 Pecia, 482

Pedilospora, 593
Pelargonium, 36, 43, 389, 544, 620, 631

Pellicularia, 382, 577,
682
Koleroga, 583
510 276 Peltostroma, 531

Pellioniella,
Peltospha:;ria,
Penicilliopsis, 167
Penicillium, 166, 167, 169, 169, 572,

573, 635
Digitatum, 574 Glaucum, 574 Italicum, 574 Luteum, 574
Olivaceum, 574 Peniophora, 406 Pennisetum, 209

Peony, 178, 529, 581, 606
Pepper, 37, 42, 268, 269, 5^0, 541
Parsnip, 36, 41, 42, 91, 592, 628 Paryphedria, 151
Paspalum, 213
Passalora, 602, 607
607 Microsperma, 607 Patellariaceae, 134 Patellina, 639 Patouillardia, 640 Patzschkeella, 505 Paulownia, 545 Paxillea>, 442
Bacilligera,
Peraphyllum, 371 Peribotryum, 634 Periconia, 598 Periconieae, 594, 597 Key to, 597 Pcriconiella, 597
Pea, 28, 99, 177, 248, 250, 260, 329, 373, 506, 519, 651
Peach, 36, 53,
106,
128,
137,
138,
176, 231, 268, 282, 357, 428, 485,
490, 499, 512, 539, 540, 541, 547, 560, 592, 604, 606
Peridermium, 330, 333, 335, 336, 350, 389, 390 Acicolum, 337, 387 Cerebrum, 352 Cornui, 352 Elatinum, 349 Oblongisporium, 338 Pyriforme, 352 Rostrupi, 339 Strobi, 351
Peridinese, 3
Peanut, 392, 557, 629

Pear, 38, 105, 130, 149, 202, 231, 246,
249, 253, 255, 268, 367, 369, 371,
Peridium, 325 Periola, 641

Perisporiaceaj, 170, 189
404, 419, 421, 246, 279, 485, 490,
Key
to,
189
INDEX
Perisporiales,
116,
124,
165, 167,
733
66, 74,
Pcronosporales,
75,
77,
170
475
to,
Key
170
Key
to,
78
Perisporium, 189 Perithecium, 62, 63
Persimmon, 540, 581 Pestalozzia, 558

Ala>a, 560
Perono plasm opara, 83, 90, 93

Celtidis, 93
Clusix, 560
Discosioides, 560
Cubensis, 93, 94 Humuli, 93
Peronospora, 78, 82, 84, 90, 93, 95, 618

Antirrhini, 101
Funerea, 559, 559 Fuscesccns, 560

Sacchari, 560 Gongrogena, 560 Guepini, 559, 559 Hartigii, 558 Inquinans, 560 Lupini, 560 Palmarum, 560 Palmicola, 560 Pha-nicis, 560 Richardise, 560 Stictica, 560 Suffocata, 560 Tumefaciens, 560 Uvicola, 559 Pestalozziella, 538 Pestalozzina, 557
Arborescens, 100 Candida, 101
Cannabina, 101 Conglomerata, 101

Corolla;, 101
Cytisi, 100
Dianthi, 101
Dipsaci, 100
Effusa, 96, 96
Ficarise,
101
Fragarise, 100
Jaapiana, 101 Linariaj, 100
Maydis, 101
Myosotidis, 101 Nicotianae, 101
Parasitica, 95, 97
Petunia, 48, 141

Pezizaceae, 133, 134
Pezizales, 123, 133
Phocnixse, 101
Potentillse,
Key Key
98
to,
133
100
Phacidiaceae, 154, 155

to,
Rubi, 100
Schachtii, 100
Schleideni, 96,
155
154
Phacidiales, 124, 154
Key
to,
Schleideniana, 96 Sparsa, 97 Trichomata, 100 Trifoliorum, 97 Valeriana;, 101

Valerianella, 101
Viciae,
Phacidieae, 156
Phacidium, 156, 157 Infestans, 157 Phaconectria, 201 Phaeodon, 414

Phaeonectria, 201
Phaeopeltosphacria, 276 Phaeophyceae, 3
97
VincE, 101
Violacea, 100
Violae,
Phacoseptoria, 517, 525

Oryzae, 525
Phaeosphacriclla,
99
Peronosporaccae, 78, 82 Key to, 83
236
Phallaceae, 462
734
462 462 Phallus, 462, 463 Impudicus, 463 Rubicundus, 463 Pharcidia, 236, 250 Oryza?, 250
Phallales, 395,
INDEX
Phoma,
Citricarpa, 491 Cyclamenae, 492
Key
to,
Cydonse, 490 Dahhse, 492

Devastatrix, 492
Hennebergii, 491 Limonis, 490
Phaseolus, 178, 187, 372 Multiflorus, 168

Vulgaris, 168
Lophiostomoides, 491
Phellomyces, 614, 657

Phlebia, 413
Phlebophora, 406
Phleospora,
243, 249, 518,
519,
MaU, 490 Malvacearum, 492 Mororum, 491 Myxiae, 491 Napobrassicae, 491
Oleandrina, 492
Oleracea, 491, 492
Persicse,
526
Aceris, 525
490
Caraganae, 525 Mori, 525 Moricola, 525 Oxycanthop, 625
Pithya, 492
Pomarum, 491
Reniformis, 242, 490
Ribesia, 492 Roumii, 492
Phleum, 180, 321, 374, 608

Phlox, 176, 178, 258, 497, 519, 523, 631
Phlyctsena, 493, 518
Phcenix, 101, 658 Pholiota, 449, 452
Sanguinolenta, 491
Sarmentella, 490
Solani,
491
Solanicola, 491
Sordida, 492
Adiposa, 452, /^53

Aurivella, 452
Sphserosperma, 247 Strobi, 492

Strobilinum, 492
Subcircinata, 491
Tiliffi,
Cervinus, 452
Destruens, 452
Mutabilis, 462
Spectabilis,
259
452
Tuberculata, 491 Uvicola, 238
Squarrosa, 452
Phoma,
532
238, 243, 245, 247, 257, 279,
Phomatospora, 263 Phomopsis, 482, 493

Alccapercrassae, 493
325, 478, 481, 484, 490, 493, 519,
Stewart ii, 493

Phorcys, 263
Albicans, 260, 490

Aleracea, 491, 492
Ambigua, 490
Apiicola, 492
Batatae, 492
Beta?, 247,
490
Phragmidium, 354, 358, 390 Americanum, 359, 359 Bulbosum, 358 Disciflorum, 359, 359 Montivagum, 359, 359
Rosa?-acicularis, 359
Bohemica, 276, 490 Brassicffi, 492 Phoma, Chrysanthemi, 492
Rosae-arkansana?, 359, 359

RosEe-californicae, 369,
359
INDEX
Phragmidium, 359
Rubi-idrei,
Rosae-setigerae,
735
359,
Phyllosticta, Cavarrr, 489
Chcnopodii, 487
359 Sppciosum, 359 Subcorticium, 359 Violaceurii, 359 Phragmites, 315, 377, 378 Phragmopyxis, 354 Phragmosporaj, 033, 037 Key to, 637
Chrysanthemi, 488 Cinnaniorii, 487 Circumscissa, 486 Citrullina, 487 Coffeicola, 486 Coma>n.sis, 486 Cruenta, 488
Cucurbitacearum, 487, 629 Cyclaminis, 488 Dammara?, 489 Dianthi, 488 Digitalis, 488 Dracaena;, 489 Fragaricola, 486 Funckia, 489 Grossularia?, 486 Halstedii, 488 Hederacca, 487 Hedericola, 487 Hevea, 487 Hortorum, 487 Humuli, 486
Hydrangese, 488 Ida^cola, 488 Ilicina, 489
Insulata, 486
Phycomyces, 105 Phycomycetes, 1,

113, 116, 118
3, 59, 64, 65,
101,
Key
to,
65
Phyllachora, 157, 217, 220, 221, 600, 007
Cynodontis, 221 Dapazioides, 221 Graminis, 220, 220 Makrospora, 221

Poaj,
221
Pomigena, 220 Sorghi, 221 Trifolii, 220, 006 Ulmi, 557

171,
Phyllactinia,
173,
175,
187,
582
Corylea, 174, 187, 188 Phyllostachys, 215
Phyllosticta, 148, 238, 242, 243, 325,
476, 481, 483, 490, 519
Japonica, 486
Labrusca?, 238, 484
Acericola, 489
Aceris, 489
Althffiina,
487
Lcucanthemi, 488 Liliicola, 488 Limitata, 485 Maculicola, 487

Maculiformis, 249, 485
Magnolia?, 489
Malkcffi, 486
Ampelopsidis, 484 Apii, 487

Argillacea, 487
Armenicola, 486
Bataticola, 486
Bellunensis, 249, 484
Beta?,
Medicaginis, 486
Minima, 489
Miuria, 486
Narcissi,
486
489
Bizzozeriana, 486
Brassicae,
484
Brassicaecola, 249
Nicotiana, 486
Cannabinis, 486 Catalpae, 489
489 486 Opuntia\ 488 Paviae, 489

Nobilis,
Olcir,
736
Phyllosticta, Persicae, 485
INDEX
Physospora, 575 Phytolacca, 408
Phaseolina, 487
Piricola,
Pirina,
485 485
Phytomyxa Leguminosarum, 8
Phytophthora, 78, 83, 84, 88, 90, 617 Agaves, 89 Cactorum, 88 Calocasise, 89 Faberi, 88 Fagi, 88 Infestans, 84, 85, 86, 87 Nicotianae, 89 Omnivora, 88, 89 Areca;, 88 Phaseoli, 84, 84, 86 Sempervivi, 88 Syringae, 88
Picea, 145, 235, 253, 349, 391, 408
Pichia, 121
Primulicola, 488
Prunicola, 486
Pteridis,
489
Putrefaciens, 486
Richardiae,
Rosaj, 487
488
Rosarum, 487
Solitaria, 4^4, 485, 485 Sphaeropsidea, 489
Succedanea, 486 Syringae, 488 Tabaci, 486 Tabifica, 247, 485 Tilia3, 489 Ulmicola, 489
Vialaj,
Piggotia, 528, 530
486
Astroidea, 221, 530

Fraxini, 530
Vincae-minoris, 488
Violae,
488 Viridis, 489 Vitis, 486
Pigweed, 408 Pilacre, 634

Pilaira,
104
Phymatotrichum, 576 Physalacria, 412 Physalis, 48, 322
Pileotaria,
354
Piloboleae, 104
Pilobolus, 104
Physalospora, 238, 251, 252, 252, 273,
539
Abietina, 253 Cattleyse, 253, 541
Fallaciosa, 253
Crystallinus, 105 Pilocratera, 135

Pilosace, 449
Pinacese, 88
Pine, 22, 52, 141, 145, 151, 157, 161,

162, 203, 230, 233, 249, 330, 333,
Gregaria, 252, 404 Laburni, 253

Vanillse,
351, 352, 391, 401, 415, 418, 419,
253
423, 424, 432, 431, 436, 438, 440,

446, 454, 460, 492, 499, 512, 515, 516, 524, 531, 532, 654, 660
Woronini, 253 Physaraceae, 10, 11
Key
to, 11
Pineapple, 496, 512, 596

Pink, 349, 507
Pinus, 161, 337, 338, 339, 340, 351,
352, 390, 408 Pionnotes, 645
Betae,
Physarella, 12
Physarum,
12 Bivalve, 12 Cinereum, 12
12,
Physoderma, 72
Physopella, 340, 345
Fici,
645
Vitis,
345 345
Rhizophila, 645 PiptocephalidaceE, 103
Piptostomum, 481
INDEX
Prrella,
737
105
Pleonectria, 19S, 207

Berolinensis, 207
Coffcicola, 207
Piricularia, 589,
591 Caudata, 592

Grisea, 591, 501, G14 Oryza), 592, G14
Ploophragriiia, 224
Pleospha-rulina, 236, 250
Pirobasidium, 633 Pirostoma, 531
Briosiana, 250
Plcospora, 252, 259, 259, 611, 618

Albicans, 260, 269, 490
Farnetianum, 531
Pirotta;a, 147
Pistillaria,
Avena>, 262
412
Bromi, 261, 613
Pisum, 1G8, 178, 372, 605 Pithorayces, 645 Pitya, 136

Placosphajria, 483
Placosphaerella, 505
Gramineum, 261, 612 Herbarum, 260, 618

Hesperidcaniin, 260, 616 Hyacinthi, 260, 603
Infectoria,
260
Plagiorhabdus, 483, 500

Oxycocci, 500, 500 Planococcus, 18 Planosarcina, 18
Plantago, 69, 96, 179 Plasmodiophora, 6
Brassica), 6, 7
Negundinis, 260 Oryzse, 260

Pisi,
260
Putrefaciens, 260
Teres, 262
Tritici,
Trichostoma, 260, 262, 612, 621 258
Califomica, 8 Humili, 8
Orchidis, 8
Tritici-repentis, 262, 613 Tropocoli, 260, 621
Ulmi, 260
Vulgaris, 610
Tomati, 8
Vitis,
Pleosporacese, 223, 250
Plasmodiophorales, 5 Key to, 6
Key
to,
251
Pleotrachelus, 68
Plasmopara, 82, 83, 90, 93, 95 Halstedii, 91 Nivea, 91 Obducens, 93 Pygmsa, 93 Ribicola, 92 Viticola, 91, 92 Platanus, 186, 205, 535 PlatyglcEea;, 392 Plectothrix, 576 Plenodomus, 482 Pleococcum, 534
Pleogibberella, 198
Pleurotus, 450, 454, 569

AtroccErulcus, 455
Corticatus, 455
Mitis, 455
Nidulans, 455
Ostreatus, 454, 456
Salignus, 454
Ulmarius, 454
Plowrightia, 216, 217
Agaves, 220
Morbosa, 218, 218, 219, 516 Ribcsia, 220 Virgultorum, 220
Pleolpidium, 68
Pleomassaria, 263 Plcomeliola, 190
Plum,
32, 38, 129, 138, 184, 219, 271,
278, 282, 357, 433, 516, 520, 562,
586, 604
Hyphaenes, 193
Pockets, 129
738
Pluteolus, 449 Pluteus, 450, 454
INDEX
Polystictus, Pergaraenus, 426, 4^6
Cervinus, 454, 455
Sanguineus, 425 Velutinus, 425

Versicolor, 425, 427
Poa,
8, 119, 180,
221, 310, 321, 375
Pocillum, 136
Pocosphaeria, 252
Polystigma, 198, 207 Ochraceum, 208
Podocapsa, 122 Podocarpus, 597 Podocrea, 199

Podosphajra, 175, 182
Leucotricha, 184, 569 Myrtillina, 183
Rubra, 208, 208, 564 Polythelis, 354

Polythrincium, 602, 606
Trifolii, 220, 606, 606 Pomelo, 549, 604 Pomes, 139, 237, 255, 278, 282, 362,
Oxyacantha;, 183, 183, 184, 569

Tridactyla, 183, 184
Podosporiella, 637
410, 491, 496, 502, 529, 569, C07,
649
Poplar, 21, 36, 47, 130, 182, 256, 340,
342, 419, 433, 440, 446, 454, 507, 535, 556, 606
Podosporium, 637 Polemonium, 507 Polycephalum, 633 Polyscytalum, 568 Polydesmus, 609 Polygonum, 96, 303 Polymorphism, 64 Polynema, 534 Polyphagus, 73 Polyporacese, 402, 416 Key to, 416 Polyporese, 416 Polyporus, 417, 418, 426 Adustus, 426 Amarus, 422
Betulinus, 425, 4^5
Borealis, 432, 423,
Poppy, 100 Mallow, 390

Populus, 127, 130, 249, 344, 499, 512,
519, 545, 555 Poria, 418
Hypolaterita, 418
LfEvigata, 418
Subacida, 418
Vaporaria, 418 Vineta, 418
Poropeltis, 531
Porothelium, 440
Potato,
8, 21, 40, 41, 43, 44, 46, 47,
48, 49, 69, 70, 86,
141, 200, 231,
4^3
258, 404, 408, 456, 491, 497, 568, 583, 584, 591, 614, 616, 617, 623, 624, 627, 637, 645, 652, 653
Beetle, 48
Potentilla, 175
Dryadeus, 423 Dryophillus, 421 Fruticum, 422 Giganteus, 421 Glivus, 421 Hispidus, 421 Obtusus, 418
Schweinitzii, 400, 401, 424
Powdery Mildew, 171

Primrose, 101, 250, 320, 552, 582
Primula, 315, 318, 488, 507, 579, 582, 591

Primulacese, 101 Prismaria, 593
Privet, 191, 269, 541
Squamosus, 419, 4~0
Sulphureus, 419, 419
Polystictus, 417, 418, 426
Cinnabarinus, 425
Hirsutus, 426
Occidentalis, 425
Promycelium, 63, 300 Prophytroma, 600 Prospodium, 354
INDEX
ProsthemioUa, 557 Prosthemium, 515
Protoba.sidii, 299,
739
Iridi.s,
PscudotnoniLs,
27
Juglandis, 27
Protoascomycetes, 114, 117, 119 323
LcKiiminijuTdus, 28
Levistici,
28 29
Key
to,
323
Maculicolurii, 28
Protocoronospora, 405, 409

Nigricans, 409
Protodiscales, 114, 123, 125
Malvarc'urum, 22,
Miciiig.uicnsc, 30
2!),
Medicaginis, 29, SO, 31
Key
to,
125
Mori, 30, 43
Olex'-tubcrculosis, 34
Protomyces, 118, 119 Macrosporus, 119 Pachydornnis, 119 Rhizobius, 119 Protomycetaceaj, 118 Key to, 118 Protomycetales, 118 Key to, 118 Protostegia, 536
Prunus, 129, 130, 140, 152, 182, 183,
184, 202, 208, 275, 347, 357, 486,
Phaseoli, 22, 27, 28, 31, 31
Pruni, 32, 32
Putridus, 23 Putrifaciens Liqucfacicns, 43

Kadicicola, 8, 32 Savastanoi, 33, 4G
Sesami, 34
Sps. Indet, 37
Stewart i, 22, 33, 34, 34 Syringa), 35
495, 496, 563, 564, 569, 579, 605, 610, 626
Psathyra, 449
Pseudobeltrania, 602 Pseudocenangium, 537 Pseudocolus, 463 Pseudodematophora, 231
Tumefacions, 3.5, 36 Vascularum, 37 Pseudopatella, 530

Pseudopcziza,
547,
147, 149,
475,
539,
555
Medicaginis, 147 Ribis, 148, 541

Salicis, 148, 541 Tracheiphila, 148
Trifolii, 148, 148, 494,
Pseudographis, 156
Pseudographium, 515 Pseudomassaria, 262

Pseudomeliola, 189
535
Pseudomonas,
18, 21,
22
Pseudophacidea;, 155 Pseudophacidium, 155
iEruginosus, 23, 27
Amaranti, 22 Araliae, 39
Avense, 23, 23, 40
Pseudoplasmopara, 93 Pscudorhytisma, 156 Pscudotryblidium, 150

Pseudotsuga, 408, 416 Pseudovalsa, 2S0. 281 Longipos, 281
Pseudozythia, 527 Psilocan', 303
Psilocybe, 449, 461
Ilcnningsii,
Campestris, 22, 2J,, 24, 25, 26, 28, 29, 31, 32, 43
Destructans, 26, 39, 42 Dianthi, 22, 27 Fluoresccns, 27, 41
Exitiosus, 27
451
Liqucfacicns, 27
Pennata,
Psilopczia, 132
-{^-^
Putrida, 27 Hyacinthi, 22, 25, 27, 28, 31

Indigofera, 13
Spadicca, 461
Psilospora, 534
740
595 Pterocarpus Indicus, 426
Pteris, 489,
INDEX
Puccinia, Pazschkei, 389
Persistens, 389 Phlei-pratensis, 384
Pterophyllus, 444
Pterula, 411 Puccinia, 355, 359, 3G1, 375, 390
Allii,
Phragmitis, 377
377
377
Poarum, 386 Podophylli, 332

Porri,
Anemones-virginianse, 389
Apii,
377
Pringsheimiana, 376
Arenarise, 387
Purpurea, 384
Ribis, 328
caricis,
Asparagi, 326, 328, 329, 330,

376, 376
376
389 Bullata, 377 Canna?, 389 Castagnei, 377 Cerasi, 376

Asteris,
Nigri-acuta^, 376
paniculata?, 376
Pseudocyperi, 376
Chrysanthemi, 386 Cichorii, 378

Convallarise-digraphidis, 388
Rubigovera, 329, 381, 383 Secalis, 382 Tritici, 382 Schrceteri, 389 Scilte, 389
Simplex, 383
Sorghi, 329, 384, 384
Coronata, 382, 383 Coronifera, 383

Cyani, 377
Dianthi, 389
Dispersa, 382
Endivise, 377
Suaveolens, 328
Taraxici, 378
Tragopogonis, 328, 377 Triticina, 382

Tulips, 389 Vexans, 327
Viola;,
Fagopyri, 378
Gentianae, 389
Gladioli, 389
388
353
Glumarum, 383
Graminis, 329, 385
AiriB,
Pucciniacese, 335, 353
334,
378,
37.9,
Key
to,
379
Pucciniastrum, 341, 346, 390, 391 Abieti-chamsenerii, 347

Epilobii, 347
Avenae, 379
Phlei-pratensis, 379
379 Secalis, 379 Tritici, 379 Granularis, 389 Helianthi, 386, 387 Heterogena, 386 Horiana, 389 Iridis, 389 Isiacae, 378, 390 Magnusii, 376
Poa;,
Goeppertianum, 342, 347 Hydrangese, 346 Myrtelli, 347 Padi, 347 Pustulatum, 347 Pucciniosita, 342 Purciniospora, 505 Puff-balls, 395
Pulparia, 151
Pulsatilla,
333
95, 107, 247, 408,
Pumpkin,
548
Malvacearum, 328, 385, 386 Menthae, 378
Purslane, 82, 408
Pycnidium, 61
LNDEX
Pycnochytrium, 70, 72 Anoinonos, 72
Globosuni, 72 Pyrenochajta, 482, 497
Fcrox, 497
Oryza', 497
Phlogis, 497
741
Ramularia, Bcta>, 590 f\)I(?os{K)rii, 591

Cynara-, 591 Gcranii, 591
Pyrcnomycetes, 150, 105, 170, 217

Pyrcnopezizu, 147
li)5,
Galdiana, 591 591 Lactoa, 591 Modcsta, 591 Narcissi, 591 Necator, 590
Ilcraclii,
Pyrenophora, 252, 262 Trichostoma, 262 Pyrenotrichum, 481 Pyroctonum, 72 Sphajricum, 73 Pyroneina, 116 Pyroncmacca;, 133, 134
PyropoIyporou.s Prarimosa, 430
Onobryoliidis, 691 Primula-. 591

Spinaciii',
590
Taraxaci, 590
Tulasnci, 244, 590
Vallambrosa', 591
Ramulaspcra, 577
Ranunculacca?, 93, 320
Pyrus, 17G, 183, 3G6, SCO

Arbutifolia, 369
Ranunculus, 101, 321, 375 Rape, 141, 258

Raspberry, 38, 245, 248, 257, 258,
270, 359, 3G0, 451, 503, 543, 544, 547, 581, 650
Pythiacca\ 75
Pythiacystis, 75, 77, 77
Citrojihthora, 77
Pythium, 75, 76, 76, 050 do Baryanurn, 77 Gracile, 77

Intcrmediuni, 77
Ravenclia, 353 Red Alga;, 3
Red Bud, 632 Red Cedar, 329, 431

Redtop, 310
Reessia, 68
Palmivoruin, 77
Rehmiella, 264 276
Q
Quercus 127, 186, 188, 202, 220, 544
Quince, 36, 38, 130, 140, 149, 249,
207, 208, 307, 309, 371, 404, 490, 502, 515, 540, 542, 557, 509, 582
Rehmiollopsis, 264, 276, 490
Bohemica, 276
Rcticulariacex, 10
R
Rabenhorstia, 483 Rabcntischia, 251
Radish, 30, 42, 81, 95, 408, 571 Radulum, 413
Rhabdospora, 518, 519, 626 Coffca?, 519, 525 Coffeicola, 519, 525 Oxycocci, 519, 525 Kubi, 519, 625 Thcobrom.'e, 519. 526
Rhacodium, 657 Rhacophyllus, 444 Rhagadolobium, 155 Rhaniphoria, 232
Ramularia, 243, 245, 589, 590

Armoraciae, 590, 590
Rhamus, 383 Rhinocladium, 599 Rhinotrichum, 575
742
Rhizidiaceae, 67
INDEX
328, 344, 351, 376, 486, 541, 542,
580, 581
Rhizina, 132
Inflata, 132,
132
Riccia, 251
Undulata, 132
Rhizinaceae, 131, 132
Rice, 46, 213, 214, 232, 250, 257, 258,

259, 260, 276, 317, 486, 495, 497,
Key
659
to,
132
503, 504, 507, 512, 516, 520, 525, 535, 591, 605, 611, 613, 626, 643, 656, 661
Rhizoctonia, 230, 231, 407, 408, 657,

Betse, 660 Crocorum, 660 Medicaginis, 660 Solani, 407, 660
Richardia, 408 Richonia, 189

Riccoa, 637 Rimbachia, 443 Robillarda, 505 Robinia, 235, 524
Roesleria Hypogaea, 154
Strobi, 660
Subepigea, 660
Violacea, 407, 660 Rhizogaster, 462
Roesteha, 335, 361, 363, 389, 391
Rhizomorpha, 659
Rhizopus, 104, 105, 105 Necans, 106 Nigricans, 105
Schizans, 106
Aurantica, 368
Botryapites, 370 Cancellata, 369
Cornuta, 368 Cydoniae, 371

221, 258,
Koreaensis, 371
Penicillata, 367
Rhododendron,
141,
194,
349, 398, 544, 559 Rhodophycea^, 3
Rhombostilbella, 634, 635

Rosae, 635
Pyrata, 364, S91 Transformans, 369 Rosa, 36, 47, 97, 105, 176, 220, 249,
284, 359, 433, 487, 492, 503, 504, 505, 509, 516, 517, 522, 544, 555, 560, 5G4, 602, 631
Rhopalidium, 557 Rhopographus, 217 Rhopalomyces, 570 Rhubarb, 41, 101, 377, 497, 506 Rhynchodiplodia, 510 Citri, 510 Rhynchomeliola, 232 Rhynchomyces, 609 Rhynchophoma, 505 Rhynchosporium, 586, 587 Graminicola, 587 Rhynchostoma, 232, 277
Rhytidhysterium, 161 Rhytidopeziza, 150 Rhytisma, 156, 158
Rosacea?,
127,
143,
330,
359,
391,
610
Roselle, 187
Rosellinia, 226, 230, 635
Aquila, 230, 231
Bothrina, 231 Echinata, 232

Ligniaria, 232
Massinkii, 231 Necatrix, 230, 231
Quercina, 231
Acerinum, 158, 159, 530 Punctatum, 159, 530 Salicinum, 159, 530 Symmetricum, 159
Ribes, 152, 176, 185, 188, 202, 220,
Radiciperda, 231 Rosenscheldia, 216

Rostrella, 166
Coffea;,
168
Rotaea, 588 Rozites, 449
INDEX
Roumegiiericlla, 527
Saprolegniales, 66, 74, 75
743
RoussccUa, 21C
Rozella, 70
Key
to,
75
Rubber
Rubus,
plant, 270
39, 72, 100, 17G, 227, 333, 359,
Sarcapodica;, 595 Sarcina, 18

Sarcinella, 191, GIG,
625
3G1, G26
Rumex,
74, 377,
582
Sarcinodoohiurn, G45 Sarcomyces, 151

Sarcoso3-pha, 135
Ruppia, 8 Rush, 329
Rust Fungi, 64, 298, 324 Rutabaga, 27

Rutstrocmia, 135
Sarcoscyphea;, 135 Sarcosoma, 151

Sarracenia, 270
Sassafras, 433 Satsuma, 604
Rye, 14G, 180, 200, 20G, 213, 257,

258, 2G2, 305, 310, 317, 319, 333, 380, 382, 520, 550, 555, 587, G13
Saxifrage, 345, 389
Scabiosa, 100, 176, 178
Kynchospora, 303
S
Sabina, 234
Saccarda?a, G30
Scaphidium, 536 Sceptromyces, 584 Schenckiella, 189 Schinzia, 323 Schizanthus, 552
Schizomycetes, 1, Key to, 18 SchizonoUa, 302
3, 13,
18
Sacrardia, 190
Saccardoclla, 252
Schizophyllea;, 443, 444
Saccharomycetacea;, 120
Key
to,
444
Key
to, 121
Schizophyllum, 444
Saccharomycps, 121 Croci^ 121

Saccharomycetales, 119 Key to, 120
Alneum, 444,
44-^
Schizosaccharomyces, 121 Schizothyrella, 536

Schizothyrium, 156 Schweinitzia, 150
Schweinitziella, 217
Scilla, 143, 375,
Saccharomycodes, 121 Saccharomycopsis, 121 Saccharum, 180 Saccoblastia, 393 Saffron, GGO Sagittaria, 315
Sainfoin, 217, 507, 591, 595
Salix, 148, 25G, 344, 494, 541,
Salsify,
389
217 Scirrhiella, 217

Soirrhia,
Sclcrodermatalcs, 398
Scleroderris, 155
606
328,
3G,
42,
52,
82,
301,
Sclerodiscus, 655 Sclerophoma, 482
617
Sclerospora, 82, 83, 89

185, 220,
Sambucus,
256
^ Graminicola,
90, 90, 101
Sanguisorba, 3G1 Santiella, 515

Sapindacea?, 127
Macrospora, 89
Sclerotinia, 135, 136, 138,
568
Alni, 143
Saponaria, 620
Saprolegniacea;, 75
Aucuparia", 143
Betula;, 143
741
Sclorotinia,
INDEX
Bulborum, 143 569 Crata^gi, 143, 569 Fructigcna, 137, 139, 569
("increa, 137, 139,
13!),
Seiridium, 558
Selenotila,
Selinia,
566
198
Sempervivum, 353
Senecio, 168, 333, 339
Fuckcliana, 581
140, 141, 579,
Sepedoniuin, 200, 577
Galanthi, 141, 581
Laxa, 137, 139, 569 Ledi, 137, 329

Libertiana, 140, 141, I4I,
Septobasidium, 405, 411 Pedicellata, 412

Septocylindrium, 588, 589
H2,
Areola, 589, 589
581
Linhartiana, 140, 569
Mespili, 140 Nicotianse, 142
Radicicolum, 590 Rufomaculans, 589

Septodothideopsis, 518
Septogloeum, 243, 556, 557

Arachidis, 557
Oxycocci, 140, 569

Padi, 140, 569
Cydoniae, 557
Fraxini, 557
Rhododendri, 141 Seaveri, 140, 569

Trifoliorum, 143
Tuberosa, 143 Urnula, 137
482 Sclerotium, 659, 660 Bulborum, 661 Cepivorum, 661

Sclerotiopsis,
Hartigianum, 557 Manihotis, 557 Mori, 249, 557 Frofusum, 557 Ulmi, 557 Septomyxa, 555 Septonema, 609 Septorella, 517
Oryzae, 661
Septoria, 243, 257, 265, 478, 517, 518
Rhizodes, 661
Rolfsii, 660, 661,
Aciculosa, 519
6S2
^sculi, 524
Tulipa;, 661
Tuliparum, 143, 661 ScolccopeUis Êruginea, 195 Scolecosporiurn, 557 'Scolccotrichum, 602, 607 Avenaj, 608 Fraxini, 608 Craminis, 608, 608 Iridis, 608 Melophthorum, 608 Musaj, 608 Scarias, 190 Scoriomyces, 641 Scorzonera, 305 Secale, 180, 379 Sedge, 89, 220, 329 Sedum, 221, 497, 522 Seiridiella, 558
Ampclina, 520 Antirrhini, 522 Armoracia?, 622 Avenaj, 520 Azalea;, 523 Beta;, 520 Canabina, 521 Caragana?, 524 Castanese, 524 Castanicola, 524 Cerasina, 520 Cercidis, 524
Chrysanthemella, 522 CitruUi, 520
Consimilis, 522
Cornicola, 524
Cucurbitacearum, 521 Curvata, 524
INDEX
Septoria, Curvula, 520
C'yclaniinis,
745
Tilia-,
522
Dianthi, 522
Divaricata, 523
Dolichi, 521
Exotica, 523
Fairmanii, 623
Fragaria;,
524 520 Ulmaria;, 624 Ulnii, 221 Varians, E22 Veronica', 524 Septosporiclla, 518
Septoria,
Tritici,
519
Septosporium,
(ilO,
620
Fraxani, 524
noterosporiuin, 620
Glaucescens, 520
Glumarum, 520
Graniinum, 520 Iledcra;, 522 Ilelianthi, 523 Ilippocastani, C24 Hydrangea), 522 Iridis, 522 Lactucse, 522 Limonum, 520 La?fgreni, 520 Longispora, 520 Lycopersici, 621, 522 Majalis, 523 Mcdicaginis, 521 Narcissi, 523 Nicotians;, 521 Nigro-maculans, C24 Nodorum, 620 Ochroleuca, 524 Oleandrina, 524 Parasitica, 523, 524 Pctrosclini, 521
Apii, 621, 521 Phlogis, 258, 619
Pini, 162
Sequoia, 243, U;}2 Service Berrj^, 191
Sesame, 34, 47
Setaria, CO, 209, 213,
305
Shad Bush, 39
Sida, 488
Sigmoidcomyccs, 570 SilUa, 283 Simblum, 464 Sircoccus, 482 Sirodesmium, G15 Siropatella, 53G Sirothecium, 500 Sirozythia, 526 Sisil, 552 Sistotrema, 413 Skeppcria, 40G SUme Fhix, 120 Slime Fungi, 3 Slime Molds, 1 Smut Fungi, 298
Smuts, G4 Snapdragon, 101, 492, 522, 553 Snowdrops, 141, 581 Sobralia, 270
Soft Rot, 105
Piricola, 24G,
Pisi,
519
Solanaceous, 86
250, 519
Populi, 249, 519
Solanum, 322, Solenia, 406
323-,
408
Pruni, 520
Pseudoplatani, 624
Ribis, 245, 619,
Sohdago, 179, 338 Solomon's Seal, 488

Sorbus, 235, 255, 33G, 368, 371, C07
Sordaria, 224
Sordariacca;, 222, 224
519
Rosa;, 522
Rostrupii, 522
Secalina, 520
Sedi, 522
Key
Sorghum,
to,
224
49, 121, 221, 305, 310, 311,
Spadicca, 524
312, 314, 384, 613
746
Sorokina, 151 Sorolpidium, 8 Beta?, 8
Sorosphaera,
6,
INDEX
Sphaerioidacese,
to,
Phaeodidymae,
Key
510
Phaeophragmise, 480, 514
Key Key
to,
514
Graminis, 8
Phaeospora;, 480, 500

to,
Sorosporium, 302, 312, 312 Consanguineum, 312

Dianthi, 312
Ellisii,
500
Phragmosporse, 480 Scolecosporae, 480, 517
312
Key
to,
517
Everhartii, 312
Sorothelia, 227
Sparassis,
Staurosporae, 480
Sphajrita, 68,
238
412
Sphserocolla, 640
Spathularia, 131
Speira, 615
Spelt, 206
Spermatia, 325
Sphaerographium, 517 Sphaeromyces, 656 Sphaeronema, 482, 484 Adiposura, 495
Spermodermia, 655 Spermogonia, 334, 325

Sphacelia, 196, 211, 212, 640, 643
Fimbriatum, 494, 495

Oryzse, 495
Phacidioides, 148, 494
Segetum, 213, 643 Typhina, 643

Sphacelotheca, 302, 303, 310
Reiliana, 312, 312
Pomarum, 495
Spurium, 152, 495 Sphaeronemella, 527
Sphaeropeziza, 156
Sorghi, 311, 311

Sphajrella,
244
225
to,
Sphajriaceffi, 222,
Key
225 475
Spha^riales, 124, 195, 221,
222 Spha?ridium, 641
Key
to,
Sphajrophragmium, 454 Spha:ropsdidales, 479, 564 Key to, 479 Sphaeropsis, 284, 501 Japonicum, 503 Magnolia?, 503
Malorum,
480
IMori,
284, 502, 502, 546
Spha?rioidacese, 479,
503
Key
to,
480
Pseudodiplodia, 603
Amerosporae, 480
Dictyosporse, 480
Ulmi, 503
Vinca?, 503
Viticola, 284 Sphaerosoma, 132 Sphaerosporium, 639
Didymospora;, 480 Helicospora?, 480 Hyalodictya?, 480
Hvalodidyma^, 480, 505 Key to, 505 Hyalophragmia?, 480, 513 Key to, 513 Hyalosporoe, 480 Key to, 480 Phaôdictyae, 480, 516 Key to, 516 Phajodidymae, 480, 503
Spha;rostilbe, 195, 196, 198, 207
Flavida, 207
Repens, 207
Sphaerotheca, 172, 175
Castagnei,
-
115
Humuli, 175, 569
Lanestris,
Mali, 184
Var. Fuliginea, 176 177
INDEX
Sphserotheca, Mors-uvse, 176, 176
.Si)oioniiia,
717
224
Pannosa, 176, 569 Spha^rulina, 236 Sphinetrina, 153

Spicaria, 201, 584, 585
Sporormiella, 224
Sporoschismeic, 609
Sporotrichella, 576
Colorans, 205, 585

Solani, 585
Sporotrichum, 230, 576, 577 Poa;, 577, 578

Spruce, 145, 162, 230, 391, 418, 423,
424, 431, 432, 434, 436, 438, 440, 509, 524 Spumaria, 11
Spicularia, 571
Spilomium, 655
Spinach, 96, 321, 487, 551, 590, 605,
611, 628, 629
Spincllus, 104
Alba,
184,
1 1
Spirea,
175,
176,
336,
524,
Spumariacca', Spurge, 544
10,
11
637
Spirechnia, 354
Spirillacese,
Spirilli,
Squash, 95, 105, 179, 247, 540, 548 Stachybotryclla, 598

Stachybotrys, 598 Stachylidiea;, 595
19
13
Spirillum, Cholerse-asiatica?, 19 Volutans, 13

Spirodelia, 315
Stagonospora, 514, 514

Carpathica, 514
Iridis,
514
Spondylocladium, 609, 614 Atrovirens, 614, 614 Spongospora, 6, 8 Subterranea, 8 Sporidium, 326 Sporocybe, 630 Sporoderma, 640
Staurochaeta, 482
Staurospora?, 633
Steccherinum, 414, 416

Ballouii, 416 Stemmaria, 630
Stemonitacea', 10
Stemphyliopsis, 592
Sporodesmium, 257, 615, 616, 617 Brassicsp, 617

Dolichopus, 617
Exitiosum, 258, 616 Var. Solani, 616
Stemphylium, 616, 617, 617 Citri, 618 Ericoctonum, 617 Tritici, 618
Stenocybe, 153
Glomorulosum, 610 Ignobile, 617 Melongenae, 617 Mucosum, 617 Piriforme, 260, 616 Putrcfacicns, 617 Scorzonera?, 617 Solani Varians, 617 Sporoglena, 600 Sporonema, 274, 534, 535 Oxycocci, 535, 535 Phacidioides, 148, 535 Platani, 274, 535 Pulvinatum, 536
Stereum, 405, 409 Frustulosum, 409, 410 Hirsutuin, 409 Purpurcurn, 410 Quorcinum, 409 Rugosum, 410
Sterigma, 298
Sterigmatocystis, 167, 310, 572, 673
Ficuum, 573
Lutco-nigra, 573
Niger, 573
StictidaccJC, 154
Stictis, 154,
155
Panizzei, 156
748
Stigmatea, 150, 236, 243, 243, 244 Alni, 243
Juniperi, 243
INDEX
Sugar Cane, 37, 47, 206, 209, 227, 228,
248, 258, 305, 374, 392, 448, 463, 484, 495. 499, 503, 512, 554, 5G0, 598, 608, 620, 630, 658
Stigmatella, 641
Stigmella, 615
Stigmina, 608, 610, 670

Briosiana, 610
Stilbacese, 565,
Sulla, 277,
Maple, 415 630

Cedar, 416 37, 268
Sunflower, 179, 321, 523
Key
to,
632 632
Swamp
Sweet Pea,
Amerospora^, 632 Key to, 633
Pepper, 37
Potato, 82, 105, 204, 337, 408,
483, 492, 495, 513, 574,
597-,
Didymosporse, 632 Helicospora?, 632

Hyalostilbea;, 632, 633
Phajostilbeae, 632
608,
083
William, 508
Phragraospora;, 632
Siilbella, 633,
635
Sycamore, 275, 408, 524, 525, 541, 546, 560, 580, 603 Sydowia, 236
Symphoricarpus, 187 Symphytum, 178 Synchytriacese, 67, 69 Key to, 70 Synchytrium, 70, 70 Endobioticum, 70 Papillatum, 7l
Vaccini, 71, 71
Flavida, 635
Nanum, 635
Populi, 636 These, 635
Stilbonectria, 198
Stilbospora, 558 S;ilbothamnium, 630 Stilbum, 207 Stone Fruits, 139, 278, 569
Stoneworts, 3
Synsporium, 598 Synthetospora, 593

176,
Strawberry,
11,
52,
100,
486,
Syringa, 35, 186

Syzygites, 104
494, 507, 519, 529, 542, 555, 590,
591
Streptococcus, 18
Streptothrix, 599
Dassonvillei, 599
Stromatinia, 137
Stropharia, 448
Tamarind, 250, 546

Tapesia, 146 Taphrina, 126
Strumella, 655, 656
Sacchari, 656
StuarteUa, 223
Stypinella, 393
Aurea, 127 Bassei, 130

Bullata, 130
Mompa, 393
Stypinellese,
Coerulescens, 127
392
Communis, 130
Cratajgi, 130
Stysanus, 630, 636, 638
Stemonites, 637
Ulmari*, 637 Veronicae, 637

Sugar-beet, 22, 36, 37, 41, 408
Dccipiens, 130 Deformans, 127, 128, 129 Farlowii, 130

Insititiffi,
130
INDEX
Taphrina, Johonsonii, 127 Lotifîpcs, 129
Thielaviopsis, 595, 596
749
Ethaceticus, 228, o98
Maculans, 130 MirabilLs, 129 Pruni, 128 Rhizipcs, 129 Rostrupiana, 130 Tlieobroma>, 130 Ulmi, 127 Taphrinopsis, 126
Paradoxa, 59G Podocarpi, 507

Thiobactcrialcs, 19
Thistle, 328
Thoracella, 505
Thozetia, 641 Thyridella, 278
Taraxacum, 17G
Taxus, 192, 259, 493 Tea, 8, 231, 243, 2G3, 287, 403, 409,
411, 415, 418, 438, 448, 544, 553,
614, 630, 635 Teasel, 100
Thyridium, 278 Thyrococcum, 658 Sirakoffi, 658

Thyronectria, 198 Thyrsidium, 553
Tiarospora, 505
Tecoma, 631
Tcleutospore, 326, 327
Telia, 326
Tichothecium, 236 Tilachlidium, 633

Tilia,
193, 202, 489, 509, 524, 545,
631, 658
Tilletia, 301, 314,
Terfeziacese, 165, 166
315
Testicularia, 303
Foetens, 315, 316
Tcstudina, 167
Tetracium, 593 Tetracladium, 593

Tetracoccosporium, 616 Tetradia Salicicola, 252 Tetramyxa, 6, 8
Tetraploa, 615 Thalictrum, 321, 389 Thallophyta, 2
Glomerulata, 315 Hordei, 317

Horrida, 317
315 317 Texana, 316

Panlcii,
Secalis,
Tritici, 316,
Tilletiaceae, 302,
317 314
Thamnidiese, 105 Thaxteria, 227
Key to, 314 Tilmadoche, 12 Timber, 415

Timothy, 24, 310, 385, 550 Titania, 280
Titca), 593,
Thecaphora, 302, 313, SIS Deformans, 313 Thecospora, 641 Thelephora, 406, 410 Galactina, 411 Laciniata, 4^0, 411 Thelephoracea?, 402, 405, 433 Key to, 405 Theleporus, 440
Thelocarpon, 197 Theobroma, 205 Thorrya, 251 Thielavia, 166, 167
Basicola, 167, 168
593
Maxilliformis, 593
Toad
89,
Stool,
398
Tobacco, 27, 33, 44, 45, 48, 50, 52, 69,

143, 260, 486, 506, 521, 570, 573, 581, 619, 621, 624, 627, 654
Tolypomyria, 576 Tolyposporella, 303

Tolyposporium, 302, 313, S14 Bullatum, 313 Filiferum, 314 Volkensii, 314
750
Tomato,
21, 27, 30, 36, 41, 42, 44, 47,
INDEX
Trichocladium, 602 Trichocollonema, 517 Trichocomaceaj, 165
52, 86, 268, 497, 522, 540, 551, 564,
605, 606, 623, 624, 643, 653
Tomentella, 403 Torsellia, 483

Torula, 191, 595, 596, 597
Exitiosa, 597
Sphserella, 597 Toruleee, 594, 595
Trichoderma, 571 Trichodytes, 562

Tricholoma, 123, 450, 460, 460 Rutilans, 460
Saponaceum, 460
Trichopeltulum, 528 Trichopezizese, 135
Trichophila, 529
Key
Toxins, 2
to,
595
Toxosporium, 557, 558 Abietinum, 558 Trabutia, 276 Trachyspora, 354 Tracy a, 315 Tracyella, 528
Ti-agopogon, 178, 310, 378
Trichopsora, 336 Trichoseptoria, 517, 518

Alpei, 518
Trichosphseria,
226,
228,
228,
654,
596
Sacchari, 228
Trichosporie.ne, 594,
598
Trametes, 417, 437 Pini, 401, 437

Radiciperda, 401, 431
Robinophila, 438 Suaveolens, 438
Thea, 438
Tranzschelia, 354, 356
Punctata, 356, 357

Trees, 193, 407, 409, 411, 426, 428,
598 Trichosporium, 599 Trichostroma, 655 Trichotheca, 639 Trichothecium, 586 Trichurus, 630 Tridentaria, 593 Trientahs, 315
Key
to,
432, 451, 452, 453, 454, 460, 568,
662
Coniferous, 418, 419, 424, 433,
440, 456
Deciduous, 414, 418, 419, 421, 425, 427, 430, 452, 454, 456
Forest, 132, 153
Fruit, 418, 421, 439
Nut, 420, 439 Orchard, 419 Ornamental, 421

Shade, 419 Timber, 419, 448 Treleasiella, 527 Tremellales, 323 Trichsegum, 616 Trichiacese, 9 Trichobelonium, 146 Trichobotrys, 598
Trifohum, 168, 178, 313, 373, 374 Trigonella Coerulea, 168 Trimmotostroma, 657, 657 Abietina, 657 Trinacrium, 593 Triphragmium, 354, 358 Ulmariae, 358, 358 Triplicaria, 655 Triticum, 180, 260, 262, 379 Trochila, 156, 157, loS, 539 Craterium, 157, 541 Popularum, 157, 555 Trogia, 443, 444 Faginea, 443 Tropaeolum, 37, 81, 362, 621 Trullula, 553, 554 Vanillae, 554 Tryblidiacese, 151, 154 Tryblidiella, 150 Tsuga, 229, 391, 416
INDEX
Tubaria, 449 Tuboralcs, 124
Tubercularia, 196, 201, 396, 639, 642,
Umbollifcrs,
6,
751
74,
91,
377,
592,
607
Uncigora, 583
642
Fici, 642, 642 Vulgaris, 202, 642
Uncinula, 175, 180 Aceris, 182

Circinata, 182
Tuberculariacea;, 565, 638
Key
to,
638
Clandcstina, 182 Flexuosa, 182
Dematiese, 638
Mori, 182
Necator, 181, 181 Prunastri, 182
Salicis,
Key
to,
655
182, 569
Amerospora;, 638, 654 Dictyosporaj, 639, 658
182
Didymospora;, 638 Hclicospora;, 639

Phragmospora^, 639, 657 Key to, 657
Scolecosporse, 639
Uredinales, 137, 323, 394, 475, 643
Key
to,
335
Biologic Specialization, 332
Cytology, 330 Form Genera, 334

Imperfecti, 335
StaurosporB, 639
Mucedincse, 638
Amerosporese, 638, 639
Key
to,
389
Infection Experiments, 334
Key to, 639, Dictyosporse, 638

Didymosporaj, 638 Helicospora;, 638
Phragmosporae, 638, 645 Key to, 645
Staurosporse, 638
Uredinium, 325
Uredinopsis, 341, 391
Uredo, 334, 335, 389, 390, 392 Arachidis, 392 Aurantiaca, 392 Autumnalis, 392
Kuhnii, 392 Muelleri, 361
Orchidis, 392
Satyrii,
Tuberculina, 335, 640, 643
Tuburcinia, 315
Tulip, 143, 310, 389, 564, 661
Tumeric, 130 Tupelo, 412

Turnip, 25, 26, 36, 41, 42, 44, 46, 81,
95, 177, 568, 592,
Tropa-'oli,
392 392
Uredospore, 327 Urobasidium, 403

Urocystis, 301, 314, 318
619
Tympanis, 151 Typha, 188 Typhula, 412
Agropyri, 320 Anemonis, 320

Cepula;, SIS, 318. 319
Colchici, 320
Graminum, 412
Variabilis, 412,
413
u
Uleomyces, 199 Ulmaria, 358
Ulmus, 188, 202, 610
320 320 Kmetiana, 320 Occulta, 319, 319 Ornithogali, 320 Primulicola, 320 Viola>, 319 Urohendersonia, 515
Gladioli,
Italica,
752
Uromyces, 355, 371, 375, 390
INDEX
Usti'.ago, Avensp, 303, 303,
306
Appendiculatus, 371, 372, 373
Bulgarica, 305
Betae, 374,
374
Caryophyllinus, 328, 375

Colchici, 375
Crameri, 304 Cruenta, 310 Crus-galli, 305

Esculenta, 310
Dactylidis, 374 Ervi, 375
Erythronii, 375
Fabse, 373
Ficuum, 310 Fischeri, 310

Hordei, 305, 306
374 Ficarise, 375 Jaffrini, 375 Kuhnei, 374

Fallens,
306 Macros pora, 306

La;vis, 306,
Medians, 305
Nuda, 306, 308

Panici Miliacei, 310
Medicaginis, 374
Minor, 374 Pallidus, 375

Pisi, 329, 330,
Poffi,
Perennans, 307
Phccnicus, 310
372, 374
Rabenhorstiana, 307
Sacchari, 305
Scorzonora?, 305
375 374
Scillarum, 375
Trifolii, 373, 373,
Secalis,
310
Uromycladium, 327
Urophlyctis, 73
Alfalfffi,
Shiriana, 310
Sphserogena, 310
Striffiformis,
74
309
Homispherica, 74 Kriegeriana, 74 Leproides, 73 Major, 74 Pluriannulata, 74

Pulposa, 74, 74
Trifolii,
Tragopogonis, 310 Pratensis, 305
Rubsaamcrl, 74 74 Uropyxis, 354 Urospora, 251 Urosporium, 608 Ustilaginaceae, 301, 302 Key to, 302
Ustilaginales, 214, 299, 326, 392
307 310 Vaillantii, 310 Violacea, 310 Vrieseana, 310 Zeae, 308, 308, 309 Ustulina, 285, 286 Zonata, 287
Tritici, 307,
Tulipse,
Key
to,
302
Ustilaginoidea, 199, 213, 214, 640, 643

Virens, 214, 214, 643
Ustilasinoidella, 199, 214,
Vaccinium, 184, 234, 242, 347, 397, 543, 569

Valerian, 101
650
Graminicola, 214
Valeriana, 178 Valerianella, 101

Valsa, 208, 277, 278
Musajperda, 214
(Edipigera, 214
Ustilago, 299, 300, 301 302, 303, 310,
Ambiens, 278
Caulivora, 278
311, 312, 313, 315
(Eutypa) Erumpens, 278
INDEX
Valsa(Eutypellii) Prunastri, 278
Viala-a,
753
277
Leucostoma, 278 Oxystoma, 278 Valsaceae, 223, 277 Key to, 277 Valsaria, 279
Valsonectria, 19S, 208, 208, 484
Parasitica, 208
Vibcrnuni, 101 Vibrio Rugiila, 15

Vicia,
99,
178,
313, 372,
375,
408,
506, 582
Vigna Sinensis, 168

Vinca, 101, 488, 503
Violet, 72, 73, 96, 99,
KiS, 176, 320,
Vanda, 205, 544 Vanguicria, 356

Vanilla, 204, 205, 253, 280, 375, 510,
388, 416, 488, 507, 544, 550, 591, 599, 620, 630 Virgaria, 599
Vitis, 181, 238, 323, 620, 624 Volutella, 497, 564, 641, 644
553, 554, 607 Vegetables, 51, 105 Velutaria, 150
Venturia, 251, 253, 227

Ccrasi, 255, G06
Buxi, 204 Concentrira, 645 Dianthi, 645

Fructi, 644, 644
Chlorospora, 606 Crata^gi, 255

Ditricha, 255, 607
Fraxini, 255, 606
Inajqualis, 253, 254, 607, 611
Leucotricha, 644 Volutellaria, 641

Volutina, 641
Volvaria, 449, 452
Cinerascens, 255, 607
Bombycina, 453, 454
Pomi, 253
Pyrina, 253, 607
Tremula}, 255, 607 Verbena, 176, 178, 1S7, 570 Vermicularia, 482, 496, 564
Circinans, 497
W
Walnut,
28, 275, 419, 421, 428, 430, 524, 555, 606
Lilies,
Conccntrica, 497
Water
322
5-10,
Dematium,
496, 496
Denudata, 497 Melica?, 497 Miorocha^ta, 497

Polygoni-virginica, 497
Oak, 435 Watermelon, 247, 408, 4C0, 521, 598, 629, 651
Weinmannodora, 501 Wheat, 21, 73, 90, ISO, 200, 205, 206,
207, 213, 257, 258, 304, 307, 308, 316, 329, 333, 349, 379, 380, 382, 412, 491, 493, 520, 550, 571. 572,
587, 600, 613, 618
Subcffigurata, 497
Telephii, 497
Trichclla, 495
Varians, 497
Veronica,
8, 69, 523, 524,
637
Willia, 121
Verticillieai, 566,
583
Willow, 36, 44, 155, 157, 158, 159,

182, 284, 340, 342, 344, 421, 428, 433, 438, 454, 509, 530, 560, 682
Key
to,
583
Verticilliopsis, 583,
684
Infestans, 584 Vertioillium, 196, 200, 5R3, 584, 587

Albo-atrun:i, 584
Wisteria, 21
Witrhes Broom, 126,

Wojnowicia, 515
130,
191, 211,
215, 330, 348, 349, 369, 648
Vetch, 99, 373, 409, 506
754
Woronina, 70
Woroniniella, 70
INDEX
Yeasts, 120, 121
Yew, 249
Ypsilonia, 482
Wound
Parasites, 399
Yucca, 503
Z
Xanthoxylum, 188 Xenodochus, 355, 361 Carbonarium, 361 Xenopus, 575 Xenosporium, 615
Xerotus, 445
Xylariaceae, 224, 284 Zea, 384
Zignojlla,
227
Zingiber, 46, 52, 130
Zinnia, 141
Zizania, 310 Zopfia, 189
Zopfiella, 189
Key
to,
285
Zukalia, 190, 191
Xylariea;, 285
Stuhlmanniana, 191
Zj'gochytriaceae, 67
Xylocladium, 637 Xylostroma, 657, 663 Xylotroma, 659, 633
Zygodesmus, 599 Albidus, 599

Zygomycetes,
66, 101,
114
Key
to,
102
Yam, 543
Yeast, 301
Zygorhynchus, 104 Zygosaccharomyces, 121 Zythia, 527 Fragaria;, 527
I'Ovri
icwar
SB733
.58 gen Stevens, Frank Line/The Fungi which caus
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THE FUNGI WHICH CAUSE PLANT DISEASE