CHP 3A10.1007 2F978 1 4020 4585 1 - 3

Chapter 3
Plant Diseases
and Their Pathogens
Because this is a reference book and not one to be read for pleasure or con-
tinuity, most of you will come to the material you need in this section by
way of the index or the lists of diseases given under the different hosts in
Chap. 4. At the beginning of
Chap. 4 you will nd a list of headings under

which diseases are grouped and described, from Anthracnose to Witchweed.
In the Host section,
Chap. 4, the key word, for example, rot or blight, is giv-

en in capital and small capitals, followed by the name of the pathogen (agent
causing disease) in boldface. In this Diseases section, Chap. 3, the pathogens
are listed in boldface in alphabetical order under each heading such as ROTS
or BLIGHTS and so on, followed by the common name of the disease. This
system was adopted for quick and easy reference because trying to alpha-
betize hundreds of similar common names would lead to endless confusion.
Also, it allows a very brief summary of the classication and diagnostic char-
acters of each genus before going on to a consideration of diseases caused
by the various species. This brief summary is in small type, so that it can be
readily skipped by readers uninterested in the technical details. Perhaps I am
the only one who feels the need for this quick review, to be used in conjunc-
tion with the classication given in

Chap. 2; perhaps others who have to
answer questions over a broad eld instead of their own specialty can make
use of these capsules sandwiched in between nontechnical descriptions.
An alphabetical arrangement has the great disadvantage of being thrown out
of alignment every time the name of a fungus is changed, as it so frequently
is. In some such cases the old name is retained to avoid change in order,
but the present accepted name is also given. Sometimes names have been
changed under several hosts and the old name inadvertently retained under
others. And sometimes the old name is purposely retained because it is so
82 Plant Diseases and Their Pathogens
familiar to everyone. This is particularly true of a few fungi far better known
by their anamorph states than by the correct name of the teleomorph state.
A fungus not only can have several names; it also can cause more than one
type of disease. For instance, Pellicularia lamentosa is the present name of
the fungus formerly known as Corticium vagum when causing Rhizoctonia
rot of potatoes and Corticium microsclerotia when causing web blight of
beans. As Rhizoctonia solani, the name given to the sclerotial stage, the same
fungus causes damping-off of seedlings, root rots of many plants, and brown
patch of lawn grasses. There are lots of plant diseases, and there are lots
of fungi causing them, but there are not nearly as many separate pathogenic
organisms as all the names would indicate. Thus, a Linkage Reference guides
the user to two or more common disease sites, i. e. Canker or Blight,
where the user searches for the pathogen alphabetically or the link may guide
the user directly to a pathogen in another chapter or section.
I cannot think of anything more deadly than ploughing straight through this
section from Anthracnose to Wilts. By doctors orders, take it in small dos-
es, as needed. But do read the few introductory remarks as you look up
each group, and please, please, before starting any control measures, read
the opening remarks in

Chap. 1 on Garden Chemicals, and look up, in the
list of chemicals, any material you propose to use, noting precautions to be
taken along the lines of compatibility, weather relations, and phytotoxicity.
Although the disease descriptions, fungus life cycles, and general principles
of control given here will remain fairly valid, it must be stressed that chem-
icals suggested for control are constantly changing. Todays discovery may
be obsolete tomorrow. This Plant Disease Handbook should, therefore, be
used in conjunction with the latest advice from your own county agent or
experiment station. Addresses of the state agricultural experiment stations
are given following

Chap. 4.
ANTHRACNOSE
The term anthracnose has been used for two distinct types of disease, one
characterized by a typical necrotic spot, a lesion of dead tissue, and the other
by some hyperplastic symptom, such as a raised border around a more or less
depressed center. The word was coined in France for the latter type, to dif-
ferentiate a grape disease from a smut of cereals, both of which were called
charbon. The new word was taken from the Greek Anthrax (carbuncle) and
nosos (disease), and was rst used for the grape disease, caused by Sphacelo-
ma ampelina, the chief symptom of which was a birds-eye spot with a raised
border.
A disease of brambles, raspberry and blackberry, was then named anthrac-
nose because it looked like the grape disease. The fungus, however, instead
of being correctly placed in the genus Sphaceloma, was mistakenly named
Gloeosporium venetum. The next disease entering the picture was a bean
trouble, and, because the fungus was identied as Gloeosporium (though lat-
er transferred to the genus Colletotrichum), this common bean disease with
typical necrotic symptoms was also called anthracnose and came to typify
diseases so designated.
The term spot anthracnose has been given to those diseases similar to the
original hyperplastic grape disease. Those with slight hyperplastic symptoms
are still commonly called anthracnose, and those with pronounced over-
growth of tissue are commonly called scab. Both types are caused by the
genus Elsino, anamorph state Sphaceloma, and are treated, in this revised
text, as a separate group.

Spot Anthracnose.
Anthracnose in the modern sense is a disease characterized by distinctive
limited lesions on stem, leaf, or fruit, often accompanied by dieback and
usually caused by a Gloeosporium or a Colletotrichum, anamorph fungi pro-
ducing slime spores oozing out of fruiting bodies (acervuli) in wet, pink-
ish pustules. These spores (conidia) on germinating form an appressorium
(organ of attachment) before entering the host plant. The teleomorph state of
the fungus, when known, is Gnomonia or Glomerella (see Fig. 3.1).
Figure 3.1 Bean Anthracnose. Pod and seeds with dark, sunken areas; section through bean seed showing
spores formed in an acervulus marked with prominent black setae
Apiognomonia
Apiognomonia errabunda (Anamorph, Gloeosporium quercinum). Oak
Anthracnose. See Discula umbrinella and Fig. 3.2.
Apiognomonia errabunda (formerly Gnomonia quercina). Oak Anthrac-
nose. The fungus is closely related to Gnomonia platani, usually reported as
G. veneta, but is now considered a separate species. The anthracnose appears
as brown areas adjacent to midribs and lateral veins.
Apiognomonia tiliae (formerly Gnomonia tiliae). Linden Anthracnose,
Leaf Spot, Leaf Blotch, Scorch on American and European linden.
Small, circular to irregular brown spots with dark margins form blotches
along main veins in leaves, leaf stalks, and young twigs, with rose-colored
pustules. In wet seasons, defoliation in early summer may be followed by
wilting and death of branches. Cut out and burn such branches.
Apiognomonia veneta (formerly Gnomonia platani) (G. veneta). Syca-
more Anthracnose, Twig Blight, general on American and Oriental planes
(London plane is rather resistant) and on California and Arizona sycamores.
The fungus winters as mycelium in fallen leaves, producing perithecia that
discharge ascospores when young foliage is breaking out. Mycelium also
winters in twig cankers. Young sycamore leaves turn brown and die, look-
ing as if hit by late frost. Leaves infected later in the season have irregu-
lar brown areas along the veins. Conidia ooze out from acervuli on under-
side of veins in esh-colored masses, in rainy weather, and are splashed to
other leaves. Twigs and branches have sunken cankers with more acervuli.
Native sycamores may be nearly defoliated, with smaller twigs killed. Larger
branches die with several successive wet springs. The trees usually put out
a second crop of leaves after defoliation, but this is a devitalizing process.
Dead twigs and branches give a witches broom effect to the trees.
ANTHRACNOSE 85
Figure 3.2 Oak Anthracnose
On white oaks anthracnose appears as brown areas adjacent to midribs and
lateral veins.
Control. Although raking up and burning all fallen leaves has been stressed
for years, the overwintering of the fungus on twigs makes this measure rather
ineffective. The spray schedule has called for three applications of bordeaux
mixture; a dormant spray, one when the buds swell, and another 7 days later.
Trees should be fertilized to stimulate vigorous growth.
Colletotrichum
Deuteromycetes, Coleomycetes
Spores are formed in acervuIi, erumpent, cushionlike masses of hyphae bearing conid-
iophores and one-celled, hyaline, oblong to fusoid conidia. Acervuli have stiff marginal
bristles (setae), which are sometimes hard to see. Conidia (slime-spores), held together
by a gelantinous coating, appear pinkish in mass. They are not wind-borne but can be
disseminated by wind-splashed rain. On landing on a suitable host, the conidium sends
out a short germ tube, which, on contact with the epidermis, enlarges at the tip into
a brown thick-walled appressorium. From this, a peglike infection hypha penetrates the
cuticle.
Colletotrichum acutatum. Anthracnose on almond, strawberry and black
gum.
Colletotrichum antirrhini. (
Glomerella cingulata). Snapdragon An-

thracnose, on snapdragon, chiey in greenhouses, sometimes outdoors in
late summer.
Colletotrichum atramentarium (or C. coccodes). Potato Anthracnose,
Black Dot Disease on potato stems and stolons following wilt and other
stem diseases, occasionally on tomato, eggplant, and pepper; general distri-
bution but minor importance. Starting below the soil surface, brown dead
areas extend up and down the stem. The partial girdling causes vines to lose
their fresh color and lower leaves to fall. Infection may extend to stolons and
roots. The black dots embedded in epidermal cells, inside hollow stems and
on tubers, are sclerotia to carry the fungus over winter and to produce conidia
the following spring.
The fungus is a wound parasite ordinarily not serious enough to call for
control measures other than cleaning up old refuse and using healthy seed
potatoes.
ANTHRACNOSE 87
Colletotrichum bletiae (
Glomerella cingulata) and other species. Orchid

Anthracnose, Leaf Spot on orchids coming in from the tropics.
Colletotrichum capsici. Ripe Fruit Rot of pepper.
Colletotrichum coccodes. Anthracnose on soybean.
Colletotrichum dematium (formerly Colletotrichum omnivorum). An-
thracnose on aspidistra and hosta. Large, whitish spots with brown margins
are formed on leaves and stalks. Remove and burn infected plant parts.
Colletotrichum dematium. Anthracnose on spinach.
Colletotrichum dematium f. sp. spinaciae. Spinach Anthracnose.
Known on spinach since 1880 but unimportant in most years. Leaves have
few to many circular spots, water-soaked, turning gray or brown, with setae
prominent in spore pustules. The fungus is seed-borne.
Colletotrichum dematium f. sp. truncata. Anthracnose on tomato.
Found in Georgia on Dolichos.
Colletotrichum erumpens (
Glomerella cingulata). Rhubarb Anthrac-

nose, Stalk Rot.
Colletotrichumfragariae (
Glomerella cingulata). Strawberry Anthrac-

nose found in Florida and Louisiana.
Colletotrichum fuscum. Foxglove Anthracnose small spots to 1/8 inch,
circular to angular, brown to purple brown, on leaves; sunken, fusiform
lesions on petioles and veins; minute black acervuli, with bristles, in center
of spots. Seedlings damp-off, older plants are killed or stunted in warm moist
weather. Use clean seed or treat with hot water (131F for 15 minutes).
Colletotrichum gloeosporioides (
Glomerella cingulata). Lime Anthrac-

nose, Withertip, only on lime in southern Florida.
Colletotrichum graminicola (formerly Colletotrichum sublineola). An-
thracnose on wild rice (Zizania).
Colletotrichum graminicola. Cereal Anthracnose widely distributed on
barley, oats, rye, wheat, sorghum, wild rice (Zizamia) and also on cultivated
lawn grasses, causing a root decay and stem rot. Leaf spots are small, circu-
lar to elliptical, reddish purple, enlarging and fading with age; centers have
black acervuli. The fungus winters on seed and plant refuse in or on soil.
Improved soil fertility reduces damage from this disease. This pathogen also
causes fruit anthracnose of tomato.
Colletotrichum higginsianum. Turnip Anthracnose, also on rutabaga,
mustard greens, radish, and Chinese cabbage in southeastern states. Very
small, circular gray spots on leaves, and elongate brown or gray spots on
midrib, petiole, and stem, show pink pustules in centers of dead tissue. Heav-
ily infected leaves turn yellow and die; young seeds in diseased pods may be
killed. Mustard variety Southern Curled Giant is highly resistant.
Colletotrichum lagenarium (see Colletotrichum orbiculare). Melon An-
thracnose on muskmelon, watermelon, cucumber, and other cucurbits.
Colletotrichum liliacearum (see Colletotrichum lilii). Found on dead stems
of daylilies and many other plants and perhaps weakly parasitic.
Colletotrichum lilii (formerly Colletotrichum liliacearum). Found on dead
stems of daylilies and many other plants and perhaps weakly parasitic.
Colletotrichum lindemuthianum (
Glomerella lindemuthianum). Bean

Anthracnose, a major bean disease, sometimes mistakenly called rust,
generally present in eastern and central states, rare from the Rocky Moun-
tains to the Pacic Coast.
Colletotrichum malvarum. Hollyhock Anthracnose, Seedling Blight
on hollyhock, mallow, and abutilon, particularly destructive to greenhouse
seedlings. Black blotches are formed on veins, leaf blades, petioles, and
stems. Remove and burn all old plant parts in autumn.
Colletotrichum omnivorum (see Colletotrichum dematium). Anthrac-
nose on aspidistra and hosta. Large, whitish spots with brown margins are
formed on leaves and stalks. Remove and burn infected plant parts.
Colletotrichum orbiculare. Anthracnose on watermelon.
Colletotrichum orbiculare (formerly Colletotrichum lagenarium). Melon
Anthracnose on muskmelon, watermelon, cucumber, and other cucurbits.
This is our most destructive disease of watermelons, found everywhere that
melons are grown and particularly destructive in the South. There are at least
three races of the fungus differing in ability to infect different cucurbits.
One race is virulent on cucumber, slight on watermelon, moderate on Butter-
nut squash; another is virulent on both watermelon and cucumber; Butternut
squash is immune to a third.
Leaf symptoms are small yellow or water-soaked areas, which enlarge and
turn black on watermelon, brown on muskmelon and cucumber. The dead tis-
sue shatters; leaves shrivel and die. Elongated, narrow, sunken lesions appear
on stems and petioles; vines may die. Young fruit darkens, shrivels and dies
if pedicels are infected; older fruit shows circular, black, sunken cankers or
depressions, from 1/4 to 2 inches across and 1/3 inch deep on watermelon. In
moist weather the centers of such spots are covered with gelatinous masses
of salmon-colored spores. Infected fruit has a bitter taste or the esh is tough
and insipid. Soft rots often follow the anthracnose. Epiphytotics occur only
in periods of high rainfall and temperature, near 75F.
ANTHRACNOSE 89
Control. Treating seed before planting is essential. Use a three year crop
rotation with non-cucurbits; destroy plant refuse. Watermelon varieties
Charleston Gray, Congo, Fairfax, and Black Kleckly are resistant but not
to all races of the fungus.
Colletotrichum phomoides (
Glomerella cingulata).Tomato Anthrac-

nose, common rot of ripe tomatoes, most frequent in Northeast and North
Central districts. Symptoms appear late in the season, causing more loss to
canning crops. Small, circular sunken spots, increasing to an inch in diame-
ter, penetrate deeply into the esh. At rst water-soaked, the spots turn dark,
with pinkish, cream, or brown spore masses in the depressed centers, often
arranged in concentric rings. The disease is worse in warm, moist weather.
The fungus winters in tomato refuse, sometimes in cucumber and melon
debris.
Control. Clean up trash and rotting fruit.
Colletotrichum pisi (
Glomerella cingulata). Pea Anthracnose, Leaf

and Pod Spot commonly associated with Ascochyta blight and often a sec-
ondary parasite.
Colletotrichum schizanthi. Anthracnose on buttery-ower. Symptoms
are small brown spots on leaves and water-soaked areas on young stems.
Cankers on stems and branches of older plants may cause leaves to turn
yellow, branches to die back from the tip, and nally death of all parts above
the canker.
Colletotrichum sublineola (see Colletotrichum graminicola). Anthrac-
nose on wild rice (Zizania).
Colletotrichum trichellum. Fruit Anthracnose of tomato and Hedera.
Colletotrichum truncatum. Stem Anthracnose prevalent in the South on
bean, lima bean, and soybean, also on clovers and on lentil in ND. Brick-
red spots appear on veins on underside of leaves and on pods. Plants are
chlorotic, stunted, may die prematurely; blossoms or pods may drop. Use
healthy seed grown in arid states; clean up plant refuse; rotate with non-
legumes.
Colletotrichum violae-tricoloris (
Glomerella cingulata). Anthracnose

of violet, pansy.
Colletotrichum sp. Azalea Anthracnose. New disease serious on Indian
and Kurume azaleas in Louisiana since 1954. Very small rusty brown spots
appear on both surfaces of young leaves, followed by defoliation. Spores
appear on fallen leaves, which serve as source of inoculum for the next sea-
son. Copper and organic fungicides are effective in control.
Discula
Blights.
Discula campestris. Anthracnose on maple.
Discula destructive. Anthracnose on dogwood.
Discula fraxinea. (Teleomorph, Gnomoniella fraxini). Anthracnose on
ash.
Gloeosporium
Genus characters are about the same as for Colletotrichum except that there are no setae
around the acervuli. Conidia are hyaline, one-celled, appearing in masses or pustules on
leaves or fruit. Leaf spots are usually light brown, with foliage appearing scorched.
Gloeosporium allantosporum (
Phlyctema vagabunda). Anthracnose,

Dieback on raspberry in Oregon, Washington.
Gloeosporium apocryptum (
Glomerella cingulata). Maple Anthrac-

nose, Leaf Blight, an important leaf disease of silver maple, common also
on other maples and boxelder, appearing from late May to August.
Gloeosporium aridum (Discula fraxinea). Anthracnose on ash.
Gloeosporium limetticolum (
Glomerella cingulata). Lime Anthrac-

nose, Withertip, only on lime in southern Florida.
Gloeosporium melongenae (
Glomerella cingulata). (possibly identical

with G. piperatum). Eggplant Anthracnose, Ripe Rot, an occasional
trouble.
Gloeosporium piperatum (
Glomerella cingulata). Pepper Anthrac-

nose, Fruit Spot, sometimes a leaf and stem spot but more often a disease
of green or ripe fruit.
Gloeosporium quercinum (Telemorph,

Apiognomonia errabunda). Oak
Anthracnose. See Discula umbrinella and Fig. 3.2.
Gloeosporium thuemenii f. sp. tulipi. Tulip Anthracnose found in Cal-
ifornia in 1939. Lesions on peduncles and leaf blades of Darwin tulips are
small to large, elliptical, rst water-soaked then dry with black margins and
numerous black acervuli in center of spots.
Gloeosporium sp.Peony Anthracnose on stems, leaves, owers, petals of
peony. Stem lesions are sunken, with pink spore pustules, and may complete-
ANTHRACNOSE 91
ly girdle the stalks, causing death of plants. Also a destructive anthracnose
on strawberry.
Glomerella
Ascomycetes, Phyllachorales
Perithecia are dark, hard, carbonaceous, usually beaked, immersed in substratum so only
the neck protudes. Ascospores are hyaline, one-celled; asci are thickened at tips, inoper-
culate but spores sometimes discharged with force; paraphyses present.
Glomerella cingulata (formerly Colletotrichum violae-tricoloris). An-
thracnose of violet, pansy. Circular dead spots with black margins, some-
times zonate, appear on leaves; owers have petals spotted or not fully devel-
oped and producing no seed; entire plants are sometimes killed. Remove and
burn infected plants or parts; clean up old leaves in fall. Copper sprays may
be injurious.
Glomerella cingulata (formerly Colletotrichum gloeosporioides). Anthrac-
nose, Canker, Dieback, Withertip, Fruit Rot of a great many plants, gen-
erally distributed except on the Pacic Coast, more common in the South.
Infection is often secondary, in tissues weakened from other causes. See also
under Canker and under Rots.
On citrus, orange, lemon, grapefruit there is a dying back or withertip of
twigs. Leaf spots are light green turning brown, with pinkish spore pustules
prominent in wet weather. Decayed spots are produced on ripening fruits
in storage. Similar withertip symptoms may also appear on avocado, aucu-
ba, cherimoya, g, loquat, roselle, rosemallow, royal palm, dieffenbachia,
rubber-plant, strawberry and other ornamentals and fruits. The disease has
also been reported on European white birch in Virginia. Lack of water and
nutrient deciency predispose plants to infection by this weak parasite.
The fungus attacks blue lupine and statice or sea-lavender; peach anthrac-
nose became important in Georgia when lupine was used as a ground cover
in orchards. Sweet pea anthracnose is often more severe near apple orchards
where the fungus winters on cankered apple limbs and in bitter rot mummies.
Whitish lesions disgure sweet pea leaves, shoots, and ower stalks. Leaves
wither and fall; stalks dry up before blossoming; seed pods shrivel. There
may be general wilting and shoot dieback.
Anthracnose and twig blight are widespread on privet. Leaves dry and cling
to the stem; cankers at the base of stems are dotted with pink pustules. Bark
turns brown and splits; death follows complete girdling of stems. European
privet is highly susceptible; California, Amur, Ibota, and Regal privets are
fairly resistant. Also found on black locust in GA and SC and pecan in
GA.
Control. Remove infected twigs and branches from trees and shrubs, taking
care to make smooth cuts at base of limbs and painting surfaces with a wound
dressing. Plant sweet peas, from healthy pods, at a distance from apple and
privet, in clean soil; rake up and burn plant refuse at the end of the season.
Glomerella cingulata (formerly Gloeosporium melongenae). (possibly
identical with G. piperatum). Eggplant Anthracnose Ripe Rot, an
occasional trouble. Yellow to brown spots on leaves and small to medi-
um depressed spots on fruit show pink spore masses following rain or heavy
dew. Spores are splashed by rain and spread by tools, insects, and workmen.
Rotation of crops and sanitary measures may be sufcient control.
Glomerella cingulata (formerly Gloeosporium limetticolum). Lime An-
thracnose Withertip, only on lime in southern Florida. Shoots, leaves, and
fruits are infected when young; mature tissues are immune. Twigs wither
and shrivel from one inch to several inches back from the tip; young leaves
have dead areas or are distorted; buds fail to open and may drop; fruits drop,
or are misshapen, or have shallow spots or depressed cankers.
Control. Spray with bordeaux-oil emulsion as fruit is setting, with two or
three applications of 1 to 40 lime sulfur at 714 day intervals.
Glomerella cingulata (formerly Gloeosporium apocryptum). Maple An-
thracnose Leaf Blight, an important leaf disease of silver maple, common
also on other maples and boxelder, appearing from late May to August. The
leaf spots are light brown, often merging with the leaves, appearing scorched.
The effect may be confused with the physiological scorch caused by hot
weather. On Norway maples the leaf lesions are conned to purple to brown
lines along the veins. In rainy seasons there may be severe defoliation.
Control. If trees have been affected more than a year or so, feed to stimulate
vigorous growth. Spray with a copper fungicide two or three times at 14-day
intervals, starting when buds break open.
Glomerella cingulata (formerly Colletotrichum bletiae) and other species.
Orchid Anthracnose Leaf Spot on orchids coming in from the tropics.
Lemon-colored acervuli are formed in soft, blackish spots in ragged leaves.
Burn diseased plants or parts. Spray with a copper fungicide.
Glomerella cingulata (formerly Colletotrichum pisi). Pea Anthracnose
Leaf and Pod Spot commonly associated with Ascochyta blight and often
ANTHRACNOSE 93
a secondary parasite. Spots on pods, stems, and leaves are sunken, gray, cir-
cular, with dark borders. Crop rotation is the best control.
Glomerella cingulata (formerly Gloeosporium piperatum). Pepper An-
thracnose, Fruit Spot, sometimes a leaf and stem spot but more often
a disease of green or ripe fruit. Spots are dark, sunken, with concentric rings
of acervuli and pink masses of spores, which are washed to other fruit. Seed
is infected internally and contaminated externally. Harvest seed only from
healthy fruit.
Glomerella cingulata (formerly Colletotrichum erumpens). Rhubarb
Anthracnose, Stalk Rot. Oval, soft watery spots on petioles increase
until whole stalks are included; leaves wilt and die. Small dark fruiting bod-
ies with setae survive winter in stems, produce conidia in spring. Clean up
all rhubarb remains in fall.
Glomerella cingulata (formerly Colletotrichum antirrhini). Snapdragon
Anthracnose on snapdragon, chiey in greenhouses, sometimes outdoors
in late summer. Stems have oval, sunken spots, grayish white with narrow
brown or reddish borders, fruiting bodies showing as minute black dots
in center. Spots on leaves are circular, yellow green turning dirty white,
with narrow brown borders. Stem cankers may coalesce to girdle plant at
base, causing collapse of upper portions, with leaves hanging limp along the
stem.
Control. Take cuttings from healthy plants; provide air circulation; keep
foliage dry; destroy infected outdoor plants in autumn. Spray, every 7 to 10
days.
Glomerella cingulata (formerly Colletotrichum phomoides). Tomato
Anthracnose, common rot of ripe tomatoes, most frequent in Northeast
and North Central districts. Symptoms appear late in the season, causing
more loss to canning crops. Small, circular sunken spots, increasing to an
inch in diameter, penetrate deeply into the esh. At rst water-soaked, the
spots turn dark, with pinkish, cream, or brown spore masses in the depressed
centers, often arranged in concentric rings. The disease is worse in warm,
moist weather. The fungus winters in tomato refuse, sometimes in cucumber
and melon debris.
Control. Clean up trash and rotting fruit.
Glomerella glycines, Fruit Anthracnose of tomato. Also, anthracnose on
soybean.
Glomerella gossypii, Fruit Anthracnose of tomato. Also, anthracnose on
cotton.
Glomerella lindemuthianum (formerly Colletotrichum lindemuthianum).
Bean Anthracnose, a major bean disease, sometimes mistakenly called
rust, generally present in eastern and central states, rare from the Rocky
Mountains to the Pacic Coast. It may also affect lima bean, Scarlet run-
ner, tepary, mung, kudzu, and broad beans, and cowpea. It is worldwide in
distribution, known in the United States since 1880. There are at least 34
strains of the fungus, in three different groups, but the disease has decreased
in importance with the use of western-grown, anthracnose-free seed.
The most conspicuous symptoms are on the pods, small, brown specks
enlarging to black, circular, sunken spots, in moist weather showing the
typical pinkish ooze of the slime-spores. Older spots often have narrow
reddish borders. After the spores are washed away, the acervuli look like
dark pimples. If pods are infected when young, the disease extends through
to the seed, which turns yellow, then rusty brown or black under the pod
lesion. The infection may extend deep enough to reach the cotyledons. Leaf
lesions are dark areas along veins on underside of the blade and on petioles.
Seedlings may show stem spotting below diseased cotyledons. The fungus
is spread by splashing rain, tools, and gardeners working with beans when
they are wet. Optimum temperature is between 63 and 75F, with maximum
around 85F.
Control. Use western-grown seed. Saving home-grown seed is dangerous
unless you can be sure of selecting from healthy plants and pods. Clean up,
or spade under, old bean tops; rotate crops. Never pick or cultivate beans
when vines are wet. There are some resistant varieties, but more reliance
should be placed on obtaining seed grown where the disease is not present.
Glomerella nephrolepidis. Fern Anthracnose, Tip Blight of Boston and
sword ferns. The soft growing tips of fronds turn brown and dry. Keep foliage
dry; remove and burn diseased leaves.
Gnomonia
Ascomycetes, Diaporthales
Perithecia innate, beaked, separate; paraphyses absent; ascospores two-celled, hyaline;
anamorph state Gloeosporium or Marssonina. Diseases caused by Gnomonia are classi-
ed as anthracnose, scorch, or leaf spot.
Gnomonia caryae. Hickory Anthracnose, Leaf Spot, widespread. The
disease is common in eastern states, causing defoliation in wet seasons.
ANTHRACNOSE 95
Large, roundish spots are reddish brown on upper leaf surface, dull brown
underneath. The fruiting bodies are minute brown specks, and the fungus
winters in dead leaves on the ground.
Gnomonia leptostyla (Marssonina juglandis). Walnut Anthracnose,
Leaf Spot, general on butternut, hickory, and walnut. Spring infection
comes from ascospores shot from dead leaves on the ground, secondary
infection from conidia. Irregular dark brown spots appear on leaets in early
summer; if these are numerous, there is defoliation. An unthrifty condition
of black walnuts and butternuts is often due to anthracnose.
Microdochium
Deuteromycetes, Coelomycetes
Hyaline, two-celled spores are formed in acervuli without setae. Spores are rounded at
ends and are formed in pale to black masses on leaves.
Microdochium panattonianum (formerly Marssonina panattoniana). Let-
tuce Anthracnose. Small, dead, brown spots appear on blades and petioles,
centers often falling out leaving black margined shot holes. Spots progress
from older to young inner leaves; outer leaves are broken off and blown
around by wind. The disease is important only during prolonged periods of
wet weather, when it may cause heavy losses. Sanitary measures and treating
seed before planting sufce for control.
Monographella
Ascomycetes, Dothideales
Perithecia immersed in substratum, not beaked, not setose, paraphyses lacking; spores
hyaline, two-celled. The genus contains more than 1000 species, many destructive to
plants, with conidial stages in many genera.
Monographella opuntiae (formerly Mycosphaerella opuntiae). Cactus
Anthracnose on Cereus, Echinocactus, Mammillaria, and Opuntia. The
curved spores of the anamorph state (Microdochium lunatum) form light
pink pustules on the surface of moist, light brown rotten areas. Cut out and
destroy diseased segments.
Pezicula
Ascomycetes, Helotiales
This is one of the discomycetes, cup fungi. The apothecia, formed on living plants, are
eshy, bright-colored with a peridium of dark cells forming a pseudoparenchyma. Spores
are hyaline, fusoid.
Pezicula malicorticis (formerly Neofabraea malicorticis). Northwestern
Apple Anthracnose on apple, crabapple, pear, quince, chiey in the Pacic
Northwest, where it is a native disease, serious in regions with heavy rainfall.
Cankers are formed on younger branches elliptical, dark, sunken, up to 3
or 4 inches wide and 10 to 12 inches long, delimited when mature by a crack
in the bark. Conidia of the anamorph state (Gloeosporium malicorticis) are
formed in cream-colored cushions, which turn black with age, in slits in the
bark. Young cankers, reddish brown, circular spots appear on the bark in
late fall. Fruit is infected, usually through lenticels from either ascospores or
conidia in pustules on bark, but the disease may not show up until the apples
are in storage.
Control. Cut out diseased limbs or excise cankers, burning all prunings and
dead bark. Spray with bordeaux mixture before fruit is picked and fall rains
start; repeat after harvest, and again about 2 weeks later.
Phlyctema
Pycnidia dark, separate or sometimes couent, developing in or under the epidermis
or bark. Conidiophores simle or forked; conidia hyaline, 1-celled, cylindrical or log,
spindle-shapped, mostly bent.
Phlyctema vagabunda (formerly Gloeosporium allantosporum). Anthrac-
nose, Dieback on raspberry in Oregon, Washington. See Elsino veneta
under Spot Anthracnose for the common raspberry disease called anthrac-
nose.
Pseudopeziza
ANTHRACNOSE 97
Apothecia brown, cup-shaped, arising from leaves on short stalks, not setose, paraphyses
present; spores one-celled, hyaline, ovoid.
Drepanopeziza ribis (formerly Pseudopeziza ribis). Current Anthrac-
nose, Leaf, Stem and Fruit Spot generally distributed on currant, ow-
ering currant, and gooseberry, rst reported on black currants in Connecticut
in 1873. Very small, brown, circular spots appear rst on lower, older leaves,
which turn yellow if spots are numerous. Hyaline, crescent-shaped coni-
dia are formed in moist, esh-colored masses in center of spots. In severe
infections there is progressive defoliation from below upward.
Other occasional symptoms are black, sunken spots on leaf stalks, light
brown to pale yellow lesions on canes, and black yspeck spots on green
berries, with considerable reduction in yield. Apothecia are formed on fallen
leaves; ascospores are forcibly discharged in spring and carried by wind to
young leaves.
Control. Clean up and burn old leaves under the bushes. Spray with bordeaux
mixture (preferred to the newer organics) shortly after leaves appear (about
3 weeks after blossoming) and immediately after picking. Include a good
spreader and cover both leaf surfaces thoroughly.
Pseudopeziza ribis (see Drepanopeziza ribis). Current Anthracnose,
Leaf, Stem and Fruit Spot generally distributed on currant, owering
currant, and gooseberry, rst reported on black currants in Connecticut in
1873.
BACTERIAL DISEASES
Rhizobiaceae
Agrobacterium
Small, motile, short rods, with two to six peritrichous agella or a polar or subpolar ag-
ellum, ordinarily Gram-negative, not producing visible gas or detectable acid in ordinary
culture media; growth on carbohydrate media usually accompanied by copious entracel-
lular, polysaccharide slime; gelatin liqueed slowly or not at all; optimum temperatures
25 to 30C. Found in soil, or plant roots in soil, or in hypertrophies or galls on roots or
stems of plants.
Agrobacterium rhizogenes. Hairy Root of apple, also recorded on coton-
easter, hollyhock, honey locust, honeysuckle, mulberry, peavine, peach,
quince, Russian olive, rose, and spirea. Woolly root and woolly knot
are other names given to this disease, which was long considered merely
a form of crown gall. Both diseases may appear on the same plant and in
early stages be confused. In hairy root a great number of small roots pro-
trude either directly from stems or roots or from localized hard swellings
that frequently occur at the graft union. The disease is common on grafted
nursery apple trees 1, 2, or 3 years old, and the root development may be
as profuse as witches brooms. Control measures are the same as for crown
gall.
Agrobacterium rubi. Cane Gall of brambles, on blackberry, black and pur-
ple raspberries, and, very rarely, red raspberry. Symptoms appear on fruiting
canes in late May or June as small, spherical protuberances or elongated
ridges of white gall tissue, turning brown after several weeks. Canes often
split open and dry out; produce small seedy berries. Cane gall is not as impor-
tant as crown gall, but one should use the same preventive measures. Avoid
runner plants from infected mother plants.
Agrobacterium tumefaciens. Crown Gall on a great variety of plants in
more than 40 families, general on blackberry, raspberry, and other brambles,
BACTERIAL DISEASES 99
Figure 3.3 Crown Gall on Rose
on grapes and on rose (see Fig. 3.3); on fruit trees apple, apricot, cherry,
g, peach and nectarine, pear (rarely), plum; on nuts almond very suscep-
tible, walnut fairly susceptible, pecan occasionally; on shade trees, willow
and other hard woods; rare on conifers but reported on incense cedar and
juniper; on many shrubs and vines, particularly honeysuckle and euonymus;
on perennials such as asters, daisies, and chrysanthemums; and on beets,
turnips, and a few other vegetables, with tomato widely used in experiments.
Crown gall was rst noticed on grape in Europe in 1853, and the organism
was rst isolated in 1904 in the United States from galls on Paris daisy. It
is of rst importance as a disease of nursery stock, but may cause losses of
large productive trees in neglected orchards, especially almonds and peaches
in California and other warm climates. It is very important to rose growers
and to the amateur gardeners who sometimes receive infected bushes.
Symptoms. The galls are usually rounded, with an irregular rough surface,
ranging up to several inches, usually occurring near the soil line, commonly
at the graft union, but sometimes on roots or aerial parts. On euonymus,
galls are formed anywhere along the vine. This is primarily a disease of the
parenchyma, starting with a rapid proliferation of cells in the meristematic
tissue and the formation of more or less convoluted soft or hard overgrowths
or tumors. The close analogy of the unorganized cell growth of plant galls
to wild cell proliferation in human cancer has intrigued scientists for many
years. In some fashion bacteria provide stimulus for this overdevelopment,
but similar galls have been produced on plants experimentally by injecting
a virus or growth-promoting substances.
Entrance of bacteria into plants for natural infection is through wounds.
In nurseries and orchards nematodes, the plow, the disc, or the hoe may
be responsible; on the propagating bench grafting tools are indicted. Many
claims have been made for the longevity of crown gall bacteria in soil, but it
now seems to be established that they do not live in the absence of host plants
more than a couple of years, and that sudden outbreaks of crown gall on land
not previously growing susceptible crops are due to irrigation water bringing
in viable bacteria from other infected orchards. The addition of lime to the
soil may encourage crown gall, for bacteria do not live in an acid medium.
The period of greatest activity is during the warm months.
Control. For home gardens rigid exclusion of all suspected planting stock
is the very best control. Do not accept from your nurseryman blackberries,
raspberries, roses, or fruit trees showing suspicious bumps. If you have had
previous trouble, choose a different location for new, healthy plants. Be care-
ful not to wound stems in cultivating.
For nurserymen, sanitary propagating practices are a must. Stock should be
healthy. Grafting knives should be sterilized by frequent dipping in 10%
Chlorox solution, 1 ounce in 2 gallons of water, or in denatured alcohol.
If nursery soil is infested, 2 years growth of cowpeas, oats, or crotalaria
between crops will minimize crown gall.
Fruit and nut growers can perhaps plant less susceptible varieties, although
fruit that is resistant in one locality may be diseased in another. American
grape varieties are considered more resistant than European. Apples may be
better on mahaleb root-stock, nut trees on black walnut understock. Budding
rather than grafting reduces the chance of infection.
Painting galls with a solution of Elgetol-methanol has given control of crown
gall on peaches and almonds in California. One part Elgetol (sodium dini-
trocresol) is shaken with 4 parts synthetic wood alcohol and applied with
a brush, covering the surface of the gall and extending 1/2 inch to 1 inch
beyond the margin into healthy bark.
Coryneform Group Mycobacteriaceae
Clavibacter
Slender, straight to slightly curved rods, with irregularly stained segments or granules,
often with pointed or club-shaped swellings at ends; nonmotile with a few exceptions
(C. accumfaciens and C. poinsettiae). Gram-positive.
Clavibacter agcopyri (see Corynebacterium agcopyei). Yellow Gum Dis-
ease on western wheat grass.
Clavibacter fascians (see Rhodococcus fascians). Fasciation, widespread
on sweet pea, also on carnation, chrysanthemum, gypsophila, geranium,
petunia,impatiens, Hebe sp. and pyrethrum.
Clavibacter accumfaciens (see Curtobacterium accumfaciens pv. ac-
cumfaciens). Bacterial Wilt of bean, widespread on kidney and lima beans
and soybean, causing considerable loss.
Clavibacter humiferum (see Corynebacterium humiferum). Reported from
wetwood of poplar, in Colorado.
Clavibacter michiganense (see Clavibacter michiganense subsp. michiga-
nense). Bacterial Canker of Tomato, widespread, formerly causing serious
losses of tomato canning crops.
Clavibacter michiganense subsp. michiganense (formerly Clavibacter
michiganens). Bacterial Canker of Tomato, widespread, formerly causing
serious losses of tomato canning crops. The disease has now been reported
on browallia, brunfelsia, cestrum, Datura sp., eggplant, Jerusalem-cherry,
bittersweet, pepper, painted-tongue, potato, ground-cherry, and buttery-
ower in Wyoming. This is a vascular wilt disease, seedlings remaining
stunted. Symptoms on older plants start with wilting of margins of lower
leaets, often only on one side of a leaf. Leaets curl upward, brown, and
wither, but remain attached to stem. One-sided infection may extend up
through the plant and open cankers from pith to outer surface of stem. Fruit
infection starts with small, raised, snow-white spots, centers later browned
and roughened but the white color persisting as a halo to give a birds-eye
spot. Fruits can be distorted, stunted, yellow inside. In the eld, bacteria are
spread by splashed rain and can persist in soil 2 or more years. Seeds carry
the bacteria internally as well as externally.
Control. Use certied seed, a 2- or 3-year rotation; clean up tomato refuse
at end of season and diseased plants throughout season. Fermenting tomato
pulp for 4 days at a temperature near 70F will destroy bacteria on surface
of seed; hot-water treatment, 25 minutes at 122F will kill some, perhaps not
all, of internal bacteria. Start seedlings in soil that has not previously grown
tomato.
Clavibacter poinsettiae (see Curtobacterium accumfaciens pv. poinsettia).
Stem Canker and Leaf Spot of Poinsettia, a relatively new disease, rst
noted in greenhouses in 1941.
Clavibacter sepedonicum (see Clavibacter michiganense subsp. sepe-
donicum). Bacterial Ring Rot of potato, widespread since 1931, when it
probably was introduced from Europe.
Clavibacter michiganense subsp. sepedonicum (formerly Clavibacter
sepedonicum). Bacterial Ring Rot of potato, widespread since 1931, when
it probably was introduced from Europe. All commercial varieties are sus-
ceptible, with losses formerly in millions of dollars in decay of tubers in
eld and storage. Now a single infected plant in a potato eld disqualies
the whole eld for certication. Symptoms appear when plants are nearly
full grown, with one or more stems in a hill wilted and stunted while the rest
seem healthy. Lower leaves have pale yellow areas between veins; these turn
deeper yellow, and margins roll upward. A creamy exudate is expelled when
the stem is cut across. This bacterium may also occur in sugar beet which
are symptomless.
Tuber infection takes place at the stem end, and the most prominent symp-
toms appear some time after storage. The vascular ring turns creamy yellow
to light brown, with a crumbly or cheesy odorless decay followed by decay
from secondary organisms. Bacteria are not spread from plant to plant in the
eld, but by cutting knife and ngers at planting. A knife used to cut one
infected tuber may contaminate the next 20 seed pieces.
Control. Use certied seed potatoes. Use several knives and rotate them
in disinfestant. Commercial growers use a rotating knife passed through
a chemical or hot-water bath between cuts. Disinfest tools, grader, digger,
and bags; sweep storage house clean and spray with copper sulfate, 1 pound
to 5 gallons of water.
Clavibacter xyli subsp. cynodontis. Stunting Disease of bermudagrass.
Clavibacter xyli subsp. xyli. Ratoon Stunt of sugarcane.
Corynebacterium agropyri (formerly Clavibacter agropyri). Yellow Gum
Disease on western wheat grass. Enormous masses of surface bacteria form
yellow slime between stem and upper sheath and glumes of ower head;
plants dwarfed or bent; normal seeds rare.
Corynebacterium humiferum (formerly Clavibacter humiferum). Report-
ed from wetwood of poplar, in Colorado.
Curtobacterium accumfaciens pv. accumfaciens (formerly Clavibacter
accumfaciens). Bacterial Wilt of Bean, widespread on kidney and lima
beans and soybean, causing considerable loss. Plants wilt at any stage from
seedling to pod-production, with leaves turning dry, brown, and ragged after
rains. Plants are often stunted. Bacteria winter on or in seed, which appear
yellow or wrinkled and varnished. When infected seed is planted, bacteria
pass from cotyledons into stems and xylem vessels. Other plants are infected
by mechanical injury and perhaps by insects, but there is not much danger
from splashed rain. Plants girdled at nodes may break over.
Control. Use seed grown in Idaho or California.
Curtobacterium accumfaciens pv. poinsettia (formerly Clavibacter
poinsettiae). Stem Canker and Leaf Spot of Poinsettia, a relatively
new disease, rst noted in greenhouses in 1941. Longitudinal water-soaked
streaks appear on one side of green stems, sometimes continuing through
leaf petioles to cause spotting or blotching of leaves and complete defoli-
ation. The cortex of stems turns yellow, the vascular system brown. Stems
may crack open and bend down, with glistening, golden brown masses of
bacteria oozing from stem ruptures and leaf lesions.
Control. Discard diseased stock plants; place cuttings from healthy moth-
er plants in sterilized media; avoid overhead watering and syringing; rogue
suspicious plants promptly.
Rhodococcus fascians (formerly Clavibacter fascians). Fasciation, wide-
spread on sweet pea, also on carnation, chrysanthemum, gypsophila, gera-
nium, petunia,impatiens, Hebe sp. and pyrethrum. Sweet pea symptoms are
masses of short, thick, and aborted stems with misshapen leaves developing
near the soil line at rst or second stem nodes. The fasciated growth on old
plants may have a diameter of 3 inches but does not extend more than an inch
or two above ground. The portion exposed to light develops normal green
color. Plants are not killed, but stems are dwarfed and blossom production is
curtailed.
Control. Sterilize soil or use fresh.
Rickettsialike bacteria. Bacterial Wilt on Toronto creeping bentgrass; bac-
teria found in xylem of roots, crown, and leaves. Initially, leaf blades wilt
from tip down and within several days entire leaf wilts, becomes dark green,
shriveled, and twisted; also leaf scorch of mulberry.
Enterobacteriaceae
Erwinia
Motile rods (usually) with peritrichous agella; Gram-negative; producing acid with or
without visible gas from a variety of sugars; invading tissues of living plants producing
dry necroses, galls, wilts, and soft rots. The genus is named for Erwin F. Smith, pioneer
in plant diseases caused by bacteria.
Enterobacter cloacae. Bulb Decay on onion.
Erwinia amylovora. Fire Blight, general on many species in several tribes
of the Rosaceae, particularly serious on apple, pear, and quince. Other hosts
include almond, amelanchier, apricot, aronia, blackberry, cherry, chokecher-
ry, cotoneaster, crabapple, exochorda, geum, hawthorn, holodiscus, India
hawthorn, kerria, Japanese quince, loquat, medlar, mountain-ash, plum, pho-
tinia, pyracantha, raspberry, rose, spirea, and strawberry.
Apparently a native disease, rst noticed near the Hudson River in 1780, re
blight spread south and west with increased cultivation of pears and apples.
By 1880 it had practically wrecked pear orchards in Illinois, Iowa, and other
states in the Northern Mississippi Valley. Then it devastated pears on the
Texas Gulf. Reaching California by 1910 it played havoc up the coast to
Washington.
Symptoms. Blossoms and leaves of infected twigs suddenly wilt, turn dark
brown to black, shrivel and die, but remain attached to twigs (see Fig. 3.4).
The bark is shrunken, dark brown to purplish, sometimes blistered with gum
oozing out. Brown or black blighted branches with dead persistent leaves
look as if scorched by re. The bacteria survive the winter in living tissue
at the edge of holdover cankers on limbs. These are dead, slightly sunken
areas with a denite margin or slight crack where dead tissue has shrunk
away from living. In moist weather bacteria appear on the surface of cankers
in pearly viscid drops of ooze, which is carried by wind-blown rain or insects
to blossoms. Infection spreads from the blighted bloom to the young fruit,
then down the pedicel to adjacent leaves, which turn brown, remaining hang-
Figure 3.4 Fire Blight. A hold over canker developed on apple limb at base of blighted twig; B bacteria swarming
through tissue
ing around the blighted blossom cluster. Leaf and fruit blight is also possible
by direct invasion, a secondary infection via bacteria carried from prima-
ry blossom blight by ants, aphids, ies, wasps, fruit-tree bark beetles, and
honeybees, sometimes tarnished plant bugs, and pear psyllids.
The tissue rst appears water-soaked, then reddish, then brown to black
as the bacteria swarm between the dying parenchyma cells. Division may
take place every half hour; so they multiply rapidly and are usually well in
advance of discolored external tissue. A collar rot may develop when cankers
are formed near the base of a tree. Water sprouts are common sources of
infection.
As spring changes to summer, the bacteria gradually become less active and
remain dormant at the edge of a woody canker until the next spring at sap
ow. Ordinarily they do not winter on branches smaller than 1/2 inch in
diameter.
Control. Spraying during bloom is now a standard means of preventing blos-
som blight. Use bordeaux mixture or a xed copper or streptomycin at 60 to
100 ppm. The latter is very effective at relatively high temperatures; at 65F
and below, copper is more satisfactory. Start spraying when about 10% of
the blossoms are open and repeat at 5- to 7-day intervals until late bloom is
over. A dormant spray for aphid control helps in preventing re blight. One
or more sprays may be needed for leafhoppers, starting at petal fall.
Inspect trees through the season and cut or break out infected twigs 12 inches
below the portion visibly blighted. If lesions appear on large limbs they may
be painted with one of the following mixtures:
I. 1 quart denatured alcohol, 1/4 pint distilled water, 3/4 ounce muriatic
acid, 1 1/2 pounds zinc chloride.
II. 100 grams cobalt nitrate, 50 cc glycerine, 100 cc oil of wintergreen, 50
cc acetic acid, 80 cc denatured alcohol.
III. 5 parts cadmium sulfate stock solution (1 pound stirred into 2 pints warm
water), 2 parts glycerine, 2 parts muriatic acid, 5 parts denatured alcohol.
Formulas I and II were developed for use on the West Coast, III for New
York. The paint is brushed on the unbroken bark over the lesions and for
several inches above and below the canker; it may injure if there are wounds
or cuts.
In cutting out cankered limbs during the dormant season, take the branch off
at least 4 inches back from edge of the canker, and disinfect the cut. The
home gardener may want to use 10% Chlorox for tools and bordeaux paint
for cut surfaces. Dry bordeaux powder is stirred into raw linseed oil until
a workable paste is formed.
Almost all desirable pear varieties are susceptible to re blight, particularly
Bartlett, Flemish Beauty, Howell, Clapps Favorite. Varieties Old Home, Ori-
ent, and the common Kieffer are more or less resistant. Jonathon apples are
very susceptible. Less apt to be severely blighted are Baldwin, Ben Davis,
Delicious, Duchess, McIntosh, Northern Spy, Stayman, and Winter Banana.
At the University of California some work has been done on susceptibility of
ornamentals to re blight. Pyracantha angustifolia is quite susceptible, but
P. coccinea and P. crenulata are rather resistant. Cotoneaster salicifolia is
susceptible; C. dammeri, C. pannosa, and C. horizontalis are more resistant;
and C. adpressa and C. microphylla show marked resistance.
Cultural methods inuence the degree of re blight, which is worse on fast-
growing succulent tissue. Avoid heavy applications of nitrogen fertilizers;
apply such nitrogen as is required in autumn or in spring in foliar sprays
after danger of blossom blight is over.
Erwinia carnegieana. Bacterial Necrosis of giant cactus in the entire
habitat of Carnegia gigantea. Long present in southern Arizona, this disease
was not described until 1942, after it had encroached on cactus parks and
private estates. Many giant cacti in the Saguaro National Monument have
been killed, with heaviest mortality in trees 150 to 200 years old.
Symptoms start with a small, circular, light spot, usually with a water-soaked
margin. The tissues underneath turn nearly black; the spot enlarges and has
a purplish hue with the center cracking and bleeding a brown liquid. The
rotten tissues dry, break up into granular or lumpy pieces, and fall to the
ground. Rotting on one side means leaning to that side; when the trunk is
girdled near the base, the giant is likely to fall in a wind-storm. If it does
not break, it stands as a bare, woody skeleton, with all parenchyma tissue
disintegrated. An insect, Cactobrosis fernaldialis, is largely responsible for
the rapid spread of the disease. The larvae tunnel inside the stems most of
the year, emerging from May to August to pupate for a month or so before
the adult, a tan and brown nocturnal moth, lays eggs.
Control. A phosphate dust, applied monthly from April to September, has
effectively controlled the insect vector. Incipient infections can be cut out
and the cavity allowed to dry out and cork over. Before the insect vector
was known, fallen trees were cut into short lengths, dragged to a burial pit,
covered with a disinfectant, and then with soil.
Erwinia carotovora subsp. carotovora (formerly Erwinia carotovora).
Soft Rot of calla, originally described from common calla, found on golden
calla, and also on beet, cactus, cabbage, cauliower, celery, cucumber, car-
rot, eggplant, geranium (Pelargonium), hyacinth, iris, onion, parsnip, pepper,
potato, salsify, sansevieria, tobacco, tomato, and turnip.
On calla lily the soft rot starts in upper portion of the corm and progresses
upward into leaf and ower stalks or down into roots, with the corm becom-
ing soft, brown, and watery. Sometimes infection starts at edge of petiole,
which turns slimy. Leaves with brown spots and margins die or rot off at the
base before losing color. Flowers turn brown; stalks fall over; roots are soft
and slimy inside the epidermis. Corms may rot so fast the plant falls over
without other symptoms, or the diseased portion may dry down to sunken
dark spots, in which the bacteria stay dormant to the next season.
On tomatoes, infection takes place through growth cracks, insect wounds, or
sunscald areas. The tissue is at rst water-soaked, then opaque, and in 3 to
10 days the whole fruit is soft, watery, colorless, with an offensive odor.
Control. Scrub calla corms, cut out rotted spots, and let cork over for a day or
two. Plant in fresh or sterilized soil in sterilized containers and keep pots on
clean gravel or wood racks, never on beds where diseased callas have grown
previously. Grow at cool temperatures and avoid overwatering.
Erwinia carotovora subsp. atroseptica (formerly Erwinia carotovora).
Potato Blackleg, Basal Stem Rot, Tuber Rot, general on potato. This
is a systemic disease perpetuated by naturally infected tubers. Lower leaves
turn yellow; upper leaves curl upward; stems and leaves tend to grow up
rather than spread out; stem is black-spotted, more or less softened at base
and up to 3 or 4 inches from ground, and may be covered with bacterial
slime; shoots may wilt and fall over. Tubers are infected through the stem
end. The disease is most rapid in warm, moist weather, and may continue in
storage. The bacteria are spread on the cutting knife, as with ring rot, and by
seed-corn maggots, and may persist for a time in soil.
Control. Use certied seed potatoes and plant whole tubers; if cut seed must
be used, allow to cork over to prevent infection from soil. Practice long rota-
tion; disinfest cutting knife. Late varieties seem to be more resistant.
Erwinia carotovora subsp. atroseptica (formerly Erwinia carotovora).
Delphinium Blackleg, Foot Rot, Bacterial Crown Rot of perennial
Delphinium; Stem and Bud Rot of Rocket Larkspur. In delphinium there
is a soft black discoloration at the base of the stem, with bacteria oozing out
from cracks. In larkspur there is a black rot of buds as well as yellowing
of leaves, blackening of stem, stunting of plants. The bacteria are appar-
ently carried in seed; hot-water treatment is helpful. Drenching delphinium
crowns with bordeaux mixture has been recommended in the past. Insect
larval control is helpful with potato. Avoid excessive watering or irrigation.
Erwinia carotovora subsp. carotovora (formerly Erwinia carotovora var.
carotovora). Wilt of sunower, Kalancho; zucchini squash, and draceana.
Soft Rot, general on many vegetables, in eld, storage, and transit, and
many ornamentals, especially iris. The bacteria were rst isolated from rot-
ten carrots, whence the name, but they are equally at home in asparagus, cab-
bage, turnips and other crucifers, celery, cucumber, eggplant, endive, garlic,
horseradish, melon, parsnip, pepper, spinach, sunower (stalk rot), sweet-
potato, and tomato. Besides wide distribution on iris, soft rot has been report-
ed, among ornamentals, on chrysanthemum, dahlia, Easter lily, geranium,
orchid, sansevieria, poinsettia, and yellow calla.
The bacteria enter through wounds, causing a rapid, wet rot with a most
offensive odor. The middle lamella is dissolved, and roots become soft and
pulpy. Soft rot in iris often follows borer infestation. Tips of leaves are with-
ered, the basal portions wet and practically shredded. The entire interior
of a rhizome may disintegrate into a vile yellow mess while the epidermis
remains rm. The rot is more serious in shaded locations, when iris is too
crowded or planted too deeply.
Control. Borer control, starting when fans are 6 inches high, has greatly
reduced the incidence of rot. If it appears, dig up the clumps, cut away all
rotted portions, cut leaves back to short fans. Allow to dry in the sun for
a day or two, then replant in well-drained soil, in full sun with upper portion
of the rhizome slightly exposed. Many good iris growers do not agree with
this sitting duck method, preferring to cover with an inch of soil; but the
sun is an excellent bactericide, and shallow planting is one method of disease
control. Clean off all old leaves in late fall after frost.
Prevent rot on stored vegetables by saving only sound, dry tubers, in straw or
sand, in a well-ventilated room with temperature not too much above freez-
ing. In the garden, rotate vegetables with eshy roots with leafy varieties.
Avoid bruising at harvest time.
Erwinia chrysanthemi. Bacterial Blight of Chrysanthemum, a orists
disease, rst noted in 1950. First evidence of blight is a gray water-soaked
area mid-point on the stem, followed by rot and falling over. The diseased
tissue is brown or reddish brown; the rot progresses downward to the base of
the stem or, under unfavorable conditions, may be checked with axillary buds
below the diseased area producing normal shoots. Cuttings rot at the base.
Sometimes affected plants do not show external symptoms, and cuttings tak-
en from them spread the disease. Bacteria can be spread via cutting knife,
or ngernails in pinching, and can live several months in soil. A form of
this species causes a leaf blight of philodendron and may also infect banana,
carnation, corn, and sorghum and pith/stem rot of tomato.
Control. Snap off cuttings; sterilize soil and tools.
Corn rot. Corn leaves show light or dark brown rotting at base; husks and leaf
blades have dark brown spots; lower portion of stalk is rotten, soft, brown,
with strong odor of decay; plants may break over and die, with little left but
a mass of shredded remnants of brovascular bundles. Bacteria enter through
hydathodes (water pores), stomata, and wounds.
Erwinia cypripedii. Reported from California, causing brown rot of Cypri-
pedium orchids. Small, circular to oval, water-soaked, greasy light brown
spots become sunken, dark brown to chestnut. Affected crowns shrivel;
leaves drop.
Erwinia herbicola (see Pantoea herbicola). Leaf Spot of dracaena.
On Dracaena sanderana, gypsophila and related plants.
Erwinia nimipressuralis. Wetwood of elm, slime ux, due to bacteria
pathogenic in elm trunk wood, especially Asiatic elms, but possibly occur-
ring in many other trees, including maple, oak, mulberry, poplar, and wil-
low. A water-soaked dark discoloration of the heartwood is correlated with
chronic bleeding at crotches and wounds and abnormally high sap pressure
in trunk, with wilting a secondary symptom. The pressure in diseased trees
increases from April to August or September, reaching 5 to 30 pounds per
square inch (as much as 60 pounds in one record). The bacteria inhabit ray
cells mostly and do not cause a general clogging of water-conducting tissues.
This pressure, caused by fermentation of tissues by bacteria, causes uxing,
a forcing of sap out of trunks through cracks, branch crotches, and wounds.
The ux ows down the trunk, wetting large areas of bark and drying to
a grayish white incrustation. Bacteria and yeasts working in the ux cause
an offensive odor that attracts insects.
Control. Bore drain holes through the wood below the uxing wound, slight-
ly slanted to facilitate drainage. Install 1/2-inch copper pipe to carry the drip-
ping sap away from the trunk and buttress roots. Screw the pipe in only far
enough to be rm; if it penetrates the water-soaked wood, it interferes with
drainage.
Erwinia rhapontica. Rhubarb Crown Rot, similar to soft rot.
Erwinia stewartii (see Pantoea stewartii). Bacterial Wilt of corn, Stew-
arts Disease on sweet corn, sometimes eld corn, in the middle regions of
the United States, from New York to California.
Erwinia tracheiphila. Bacterial Wilt of cucurbits, Cucumber Wilt on
cucumber, pumpkin, squash, and muskmelon but not watermelon. The dis-
ease is generally east of the Rocky Mountains and is also present in parts of
the West; is most serious north of Tennessee. Total loss of vines is rare, but
a 10 to 20% loss is common.
This is a vascular wound disease transmitted by striped and 12-spotted
cucumber beetles. Dull green abby patches on leaves are followed by sud-
den wilting and shriveling of foliage, and drying of stems. Bacteria ooze
from cut stems in viscid masses. Partially resistant plants may be dwarfed,
with excessive blooming and branching, wilting during the day but partial-
ly recovering at night. The bacteria winter solely in the digestive tract of
the insects and are deposited on leaves in spring with excrement, entering
through wounds or stomata.
Control is directed chiey at the insects. Start vines under Hotkaps and spray
or dust with rotenone or other insecticide when the mechanical protection is
removed. Experimental spraying with antibiotics streptomycin, terramycin,
and neomycin has reduced wilt and increased yield.
Pantoea ananatis, Leaf Blight and Bulb Decay of onion.
Pantoea herbicola (formerly Erwinia herbicola). Leaf Spot of dracaena.
On Dracaena sanderana, gypsophila and related plants. Galls are formed at
crown and roots of grafted plants from 1/4 to 1 inch in diameter, but with
a at nodular growth rather than the usual globose crown gall.
Pantoea stewartii (formerly Erwinia stewartii). Bacterial Wilt of corn,
Stewarts Disease on sweet corn, sometimes eld corn, in the middle
regions of the United States, from New York to California. This is a vascular
disease with yellow slime formed in the water-conducting system, resulting
in browning of nodes, and dwarng of plants; or long pale green streaks on
leaf blades, followed by wilting and death of whole plant. Tassels may be
formed prematurely and die before the rest of the plant. The bacteria are
chiey disseminated by corn ea beetles, and winter either in the beetles or
in seed. Primary infections come from ea beetles feeding in spring, from
infected seed, and occasionally from soil; but secondary spread is mostly by
insects.
Corn grown in rich soil is more susceptible to wilt, and so are early vari-
eties, especially Golden Bantam. Winter temperatures inuence the amount
of wilt the following summer. If the winter index, which is the sum of mean
temperatures for December, January, and February, is above 100, bacterial
wilt will be present in destructive amounts on susceptible varieties. If the
index is below 90, the disease will be very sparse in northeastern states; if
the index is between 90 and 100, there will be a moderate amount of wilt.
Disease surveys over a period of years testify to the reliability of such fore-
casts (based on the amount of cold the ea beetle vectors can survive); but
with the increasing use of hybrid sweet corn resistant to wilt, the importance
of winter temperatures is reduced.
Control. Use insecticides to control ea beetles; substitute commercial fertil-
izer for manure; destroy infected refuse; try treating seed with terracmycin or
streptomycin. Use resistant varieties such as Golden Cross Bantam, Carmel-
cross, Ioana, Marcross, and Iochief.
Pseudomonadaceae
Pseudomonas
Motile with polar agella; straight or curved rods; Gram-negative. Many species produce
a greenish, water-soluble pigment. Many species are found in soil and water; many are
plant pathogens causing leaf spots or blights.
Acidovorax avenae (formerly Pseudomonas albopreciptans). Bacterial
Spot of cereals, grasses, and corn. Light or dark brown spots or streaks on
grass blades. Bacteria enter through stomata or water pores.
Acidovorax avenae (formerly Pseudomonas avenae). Bacterial Leaf Spot
of sweet corn. Bacterial Leaf Blight of johnsongrass.
Acidovorax avenae subsp. citrulli (formerly Pseudomonas pseudoalcali-
genes). Angular Leaf Spot of muskmelon and watermelon. Fruit blotch;
on melon, squash, pumpkin, and watermelon.
Acidovorax cattleyae (formerly Pseudomonas cattleyae). Brown Spot of
orchids, Phalaenopsis and Cattleya, common in greenhouses. Infection is
through stomata of young plants, wounds of older plants. Dark green, circu-
lar water-soaked spots change to brown and nally black. On Cattleya the
disease is limited to older leaves.
Burkholderia andropogonis (formerly Pseudomonas andropogonis). Bac-
terial Stripe of sorghum and corn. Bacterial Leaf Spot on bougainvillea.
Black Spot on clovers and statice. Also causes blight of chickpea, and bac-
terial leaf spot on white clover. Red streaks and blotches appear on leaves and
sheaths, with abundant exudate drying down to red crusts or scales, readily
washed off in rains. Bacteria enter through stomata.
Bacterial Leaf Spot of velvet bean, clovers. Translucent angular brown
leaf spots have lighter centers and chlorotic surrounding tissue; there is
no exudate. Bacteria enter through stomata and ll intercellular spaces of
parenchyma.
Burkholderia caryophylli (formerly Pseudomonas caryophylli). Bacteri-
al Wilt of carnation, usually under glass. Plants wilt, turn dry, colorless
with roots disintegrating. Grayish-green foliage is the rst symptom, but
leaves rapidly turn yellow and die. Yellow streaks of frayed tissue in vas-
cular areas extend a foot or two up the stem. It takes a month for disease to
show up after inoculation, but it can be transmitted on cuttings taken from
plants before appearance of symptoms. The sticky character of diseased tis-
sue distinguishes this wilt from Fusarium wilt. Varieties Cardinal Sim, Lad-
die, Mamie, Portrait, and others may have severe cankers at base of stems,
orange-yellow when young, very sticky. Bacteria are spread by hands, tools,
splashing water. Also causes crown and leaf rot of statice.
Control. Remove and burn diseased plants and all within 1 1/2 -foot radius.
After handling wash with hot water and soap, sterilize tools (10% Clorox for
5 minutes). Obtain rooted cuttings from propagators of cultured, disease-free
material; keep in shipping bags until ready for benching and then place in
raised, steam-pasteurized benches. Never place cuttings in water or a liquid
fungicide (use dust if a fungicide is required for other diseases); never place
temporarily on an unsterilized table; never cut or trim with hands or knives;
never plant in outdoor nurse beds; never use overhead watering.
Burkholderia cepacia (formerly Pseudomonas cepacia). Sour Skin Rot of
onion. Slimy yellow rot of outer eshy scales, with a vinegar odor. Let crop
mature well before harvesting, tops dry before topping; cure bulbs thorough-
ly before storage.
Burkholderia gladioli (formerly Pseudomonas gladioli). Leaf Spot and
Blight on birds nest fern.
Onion Bulb Rot, a storage disease, inner scales of bulb water-soaked and
soft, sometimes entire bulb rotting.
Orchid Brown Rot and Leaf Spot.
Gladiolus Scab, Stem Rot, Neck Rot, widespread on gladiolus, also
on iris, bell peppers and tigridia. Lesions on corms are pale yellow, water-
soaked circular spots deepening to brown or nearly black, eventually sunken
with raised, horny, or brittle margins that are scablike and exude a gummy
substance. Bacteria overwinter on corms. First symptoms after planting are
tiny reddish raised specks on leaves, mostly near the base, enlarging to dark
sunken spots, which grow together into large areas with a rm or soft rot.
Sometimes plants fall over, but the disease is not ordinarily very damaging
in the garden. The chief loss is to the grower in disgured, unsalable corms.
Brown streaks in husks sometimes disintegrate, leaving holes.
Gladiolus scab is increased by bulb mites, may be related to grub and wire-
worm injury.
Pseudomonas aceris (see Pseudomonas syringae pv. aceris). Maple Leaf
Spot found in California on big leaf maple.
Pseudomonas adzukicola. Stem Rot of adzuki bean.
Pseudomonas albopreciptans (see Acidovorax avenae). Bacterial Spot of
cereals, grasses, and corn.
Pseudomonas andropogonis (see Burkholderia andropogonis). Bacterial
Stripe of sorghum and corn.
Bacterial Leaf Spot of velvet bean, clovers. Translucent angular brown
leaf spots have lighter centers and chlorotic surrounding tissue; there is
no exudate. Bacteria enter through stomata and ll intercellular spaces of
parenchyma.
Pseudomonas angulata (see Pseudomonas syringae pv. angulata). Black-
re of tobacco.
Pseudomonas asplenii. Bacterial Leaf Blight of birds-next fern, rst
reported from greenhouses in California. Small translucent spots enlarge to
cover whole frond; bacteria may invade crown and kill whole plant. Control
depends on strict sanitation sterilizing ats, pots, media, and foreceps used
in transplanting. Avoid excessive watering and too high humidity.
Pseudomonas avenae (see Acidovorax avenae). Bacterial Leaf Spot of
sweet corn. Bacterial Leaf Blight of johnsongrass.
Pseudomonas berberidis. Bacterial Leaf Spot of barberry. Small, irreg-
ular, dark green water-soaked areas on leaves turn purple-brown with age;
occasional spotting occurs on leaf stalks and young shoots. If twigs are
infected, buds do not develop in the next season; if they are girdled, the entire
twig is blighted. Cut out infected twigs and spray with bordeaux mixture or
an antibiotic.
Pseudomonas caryophylli (see Burkholderia caryophylli). Bacterial Wilt
of carnation, usually under glass.
Pseudomonas cattleyae (see Acidovorax cattleyae). Brown Spot of
orchids, Phalaenopsis and Cattleya, common in greenhouses.
Pseudomonas cepacia (see Burkholderia cepacia). Sour Skin Rot of
onion. Slimy yellow rot of outer eshy scales, with a vinegar odor.
Pseudomonas cichorii. Bacterial Leaf Spot on basil. Bacterial Blight
on Lobelia.
Pseudomonas cichorii. Bacterial Rot of chicory, Belgium endive, French
endive, iris, Soft Rot of potato, and Bacterial Leaf Spot of hibiscus, gera-
nium, magnolia and rhododendron. May also cause a Leaf Spot and Stem
Necrosis on chrysanthemum (see Fig. 3.5) and Bacterial Leaf Blight on
dwarf Schefera. A yellowish olive center rot, affecting young inner leaves.
Pseudomonas corrugata. Stem Rot of tomato, also Pith Necrosis.
Pseudomonas uorescens (marginalis). Marginal Blight of lettuce, Kan-
sas Lettuce Disease, also on witloof chicory, Soft Rot of potato tubers.
Leaf margins are dark brown to almost black, rst soft, then like parchment.
Yellowish red spots, turning dark, are scattered over leaves. Infected tissue
disintegrates into an odorous mass. Bacteria live in the soil, which should
not be splashed on plants by careless watering.
Pseudomonas gladioli (see Burkholderia gladioli). Leaf Spot and Blight
on birds nest fern.
Pseudomonas melophthora. Apple Rot, probably widespread. This is
a decay of ripe apples following after apple maggots and eventually rotting
whole fruit.
Figure 3.5 Bacterial Black Spot on Chrysanthemum
Pseudomonas pseudoalcaligenes (see Acetovorax avenae subsp. citrulli).
Angular Leaf Spot of muskmelon and watermelon. Fruit Blotch on mel-
on, squash, pumpkin, and watermelon.
Pseudomonas ribicola. On golden currant in Wyoming.
Pseudomonas sesami. Bacterial Leaf Spot of sesame. Brown spots on
leaves and stems. Can be controlled by treating seed with streptomycin.
Pseudomonas solanacearum (see Rolstonia solanacearum). Southern
Bacterial Wilt, also called Brown Rot, Bacterial Ring Disease, Slime
Disease, Granville Wilt (of tobacco), present in many states but particular-
ly prevalent in the South, from Maryland around the coast to Texas.
Pseudomonas syringae. Canker on kiwifruit; also Blight on impatiens and
mock orange. Also Leaf Spot on English and American elm, mountain-
laurel, arugula and coriander. Stem Dieback of Centaurea and fennel.
Pseudomonas syringae pv. aceris (formerly Pseudomonas aceris). Maple
Leaf Spot found in California on big leaf maple. Small, water-soaked spots,
surrounded by yellow zones, turn brown or black; cankers develop on peti-
oles and bracts in serious cases; leaves may drop; disease present in cool,
damp weather of early spring.
Pseudomonas syringae pv. angulata (formerly Pseudomonas angulata).
Blackre of tobacco.
Pseudomonas syringae pv. apii. Bacterial Blight of celery. Small, irregu-
larly circular rusty leaf spots, with a yellow halo, are occasionally numerous
enough to cause death of foliage, but commonly are only disguring. Spray
plants in seedbed with bordeaux mixture, or dust with copper lime dust; clean
up old refuse.
Pseudomonas syringae pv. apii. Bacterial Leaf Spot of celery.
Pseudomonas syringae pv. aptata. Bacterial Spot on beets, Swiss chard,
and nasturtium. Spots on nasturtium leaves are water-soaked, brownish, 1/8
to 1/4 inch across. On beets they are dark brown or black, irregular, and
in addition there are narrow streaks on petioles, midribs, and larger veins.
Petiole tissue may be softened as with soft rot. Infection is only through
wounds.
Pseudomonas syringae pv. coronafaciens. Halo Blight on grasses, such as
Poa spp. and Calamagrostis spp.
Pseudomonas syringae pv. delphinii. Delphinium Black Spot on del-
phinium and aconite (monkshood). Irregular tarry black spots on leaves,
ower buds, petioles, and stems may coalesce in late stages to form large
black areas. The bacteria enter through stomata or water pores. Occasionally
this bacterial leaf spot results in some distortion, but most abnormal growth
and blackening of buds is due to the cyclamen mite, a much more important
problem than black spot.
Control. Remove diseased leaves as noticed; cut and burn all old stalks at end
of season; avoid overhead watering. In a wet season spraying with bordeaux
mixture may have some value.
Pseudomonas syringae pv. glycinea. Bacterial Blight of soybean. Per-
haps the most common and conspicuous disease of soybean, appearing in
elds when plants are half-grown and remaining active until maturity, with
defoliation during periods of high humidity or heavy dews. Small, angular,
translucent leaf spots, yellow to light brown, turn dark reddish brown to near-
ly black with age. There is often a white exudate drying to a glistening lm
on under leaf surfaces. Black lesions appear on stems and petioles, and on
pods water-soaked spots enlarge to cover a wide area, darken, and produce
an exudate drying to brownish scales; seeds are often infected. Seedlings
from infected seed have brown spots on cotyledons and often die. Flambeau
and Hawkeye varieties are somewhat less susceptible. Use seed taken from
disease-free pods.
Pseudomonas syringae pv. helianthi. Bacterial Leaf Spot of sunower.
Leaves show brown, necrotic spots, rst water-soaked, then dark and oily.
Pseudomonas syringae pv. hibisci. Bacterial Leaf Spot on Hibiscus.
Pseudomonas syringae pv. lachrymans. Angular Leaf Spot of cucur-
bits, general on cucumber, muskmelon, summer squash, occasional on other
cucurbits. Leaves or stems have irregular, angular, water-soaked spots with
bacteria oozing out in tearlike droplets that dry down to a white residue.
Eventually the spots turn gray, die, and shrink, leaving holes in foliage. Fruit
spots are small, nearly round, with the tissue turning white, sometimes crack-
ing. The bacteria overwinter in diseased plant tissue and in the seed coat.
They are spread from soil to stems and later to fruit in rainy weather, also
transferred from plant to plant on hands and clothing. Infection is most severe
in plants gone over by pickers early in the morning before dew has dried off.
Control. Plow under or remove vines immediately after harvest.
Pseudomonas syringae pv. mori. Bacterial Blight of mulberry, gener-
al on black and white mulberry. Numerous water-soaked leaf spots join to
form brown or black areas with surrounding yellow tissue. Young leaves
may be distorted, with dark sunken spots on midribs and veins. Dark stripes
with translucent borders on young shoots exude white or yellow ooze from
lenticels. Dead twigs and brown leaves resemble re blight; trees are stunted
but seldom killed. Remove and burn blighted branches; do not plant young
mulberry trees near infected specimens.
Pseudomonas syringae pv. mors-prunorum. Bacterial Canker of stone
fruits, Citrus Blast, Lilac Blight on many unreleated plants, including
apple, plum, peach, cherry, pear, almond, avocado, citrus fruits, lilacs, ow-
ering stock, rose, beans, cowpeas, oleander, and leaf spot on peas.
On stone fruits all plant parts are subject to attack, but most destructive are
elongated water-soaked lesions or gummy cankers on trunks and branches,
usually sour-smelling. Dormant buds of cherry and apricot are likely to be
blighted, pear blossoms blasted. Small purple spots appear on leaves of plum
and apricot, black lesions on fruit of cherry and apricot. All varieties of apri-
cot are very susceptible to the disease. Plums on Myrobalan rootstock are
more resistant, and varieties California, Duarte, and President are tolerant.
On citrus, and particularly lemons, dark sunken spots, called black pit, are
formed on fruit rind, but there is no decay. The blast form of the disease is
most often on oranges and grapefruit water-soaked areas in leaves, which
may drop or hang on, twigs blackened and shriveled. The disease is most
serious in seasons with cold, driving rainstorms.
On lilac, brown water-soaked spots on leaves and internodes on young shoots
blacken and rapidly enlarge. Young leaves are killed; older leaves have large
portions of the blade affected. Infection starts in early spring in rainy weather.
The bacteria are primarily in the parenchyma, spreading through intercellular
spaces, blackening and killing cells, forming cavities. The vascular system
may also be affected, followed by wilting of upper leaves.
Control. Prune out infected twigs and branches. In California spray fruits in
fall with bordeaux mixture, at the time rst leaves are dropping. Grow bushy,
compact citrus trees less liable to wind injury; use windbreaks for orchards.
Pseudomonas syringae pv. papulans. Blister Spot of apple. Small, dark
brown blisters on fruit and rough bark cankers on limbs start at lenticels.
Bark may have rough scaly patches from a few inches to a yard long, bor-
dered with a pimpled edge, and with outer bark sloughing off in spring.
Pseudomonas syringae pv. phaseolicola. Bean Halo Blight, halo spot on
common, lima, and scarlet runner beans. The symptoms are those of other
bean blights except that there are wide green or yellowish green halos around
water-soaked leaf spots, such spots later turning brown and dry. Leaves wilt
and turn brown; young pods wither and produce no seed; sometimes plants
are dwarfed with top leaves crinkled and mottled. In hot weather, spots are
often angular, reddish brown, and without halo. Stem streaks are reddish,
with gray ooze; pod spots are red to brown with silver crusts; seeds are
small, wrinkled, with cream-colored spots. All snap beans are susceptible;
many dry beans Pinto, Great Northern, Red Mexican, Michelite are rather
resistant.
Control. Use seed from blight-free areas. Blight is rare in California, occa-
sional in Idaho. Plan a 3-year rotation. Do not pick beans when foliage is
wet.
Pseudomonas syringae pv. pisi. Bacterial Blight of pea, general on eld
and garden peas, especially in East and South, and causing a leaf spot of
sweet peas. Dark green water-soaked dots on leaves enlarge and dry to rus-
set brown; stems have dark green to brown streaks. Flowers are killed or
young pods shriveled, with seed covered with bacterial slime. Bacteria enter
through stomata or wounds, and if they reach the vascular system, either
leaets or whole plants wilt. Vines infected when young usually die. Alaska
and Telephone varieties are particularly susceptible.
Control. Avoid wounding vines during cultivation. Sow peas in early spring
in well-drained soil. Use disease-free seed and plan a 4-year rotation.
Pseudomonas syringae pv. porri. Bacterial Blight of shallot.
Pseudomonas syringae pv. primulae. Bacterial Leaf Spot of primrose in
ornamental and commercial plantings in California. Infection is conned to
older leaves irregularly circular brown lesions surrounded by conspicuous
yellow halos. Spots may coalesce to kill all or part of leaf. Spraying with
bordeaux mixture has prevented infection.
Pseudomonas syringae pv. savastanoi. Olive Knot, Bacterial Knot of
olive. Irregular, spongy, more or less hard, knotty galls on roots, trunk,
branches, leaf, or fruit pedicels start as small swellings and increase to sev-
eral inches with irregular ssures. Terminal shoots are dwarfed or killed;
whole trees may die. Bacteria enter through wounds, often leaf scars or
frost cracks. Variety Manzanilla is most susceptible of the olives commonly
grown in California. Another form of this species causes similar galls on
ash.
Control. Cut out galls carefully, disinfesting tools; paint larger cuts with
bordeaux paste and spray trees with bordeaux mixture in early November,
repeating in December and March if infection has been abundant. Do not
plant infected nursery trees or bring equipment from an infected orchard into
a healthy one.
Pseudomonas syringae pv. syringae. Brown Spot, Foliar on wild rice
(Zizania); leaf spot and stem collapse on urd bean; leaf spot and stem canker
on Ginkgo.
Oleander Bacterial Gall. Galls or tumors are formed on branches, herba-
ceous shoots, leaves, and owers but not on underground parts. Small
swellings develop on leaf veins, surrounded by yellow tissue, with bacterial
ooze coming from veins in large quantity. Young shoots have longitudinal
swellings with small secondary tubercles; young leaves and seedpods may
be distorted and curled. On older branches tumors are soft or spongy and
roughened with projecting tubercles; they slowly turn dark. Prune out infect-
ed portions, sterilizing shears between cuts; propagate only from healthy
plants.
Pseudomonas syringae pv. tabaci (see Pseudomonas tabaci). Blackre of
tobacco.
Pseudomonas syringae pv. tabaci. Tobacco Wildre on tobacco, tomato,
eggplant, soybean, cowpea, pokeberry, and ground-cherry, in all tobacco dis-
tricts sporadically. Leaf spots have tan to brown dead centers with chlorotic
halos. The disease appears rst on lower leaves and spreads rapidly in wet
weather. The bacteria persist a few months in crop refuse and on seed and
enter through stomatal cavities. In buried soybean leaves the bacteria have
lived less than 4 months; so fall plowing may be benecial. Seed stored for
18 months produces plants free from wildre.
Pseudomonas syringae pv. tagetis. Bacterial Leaf Spot. Circular necrotic
lesions on leaves and petioles. The lesions have dark purple margins. This
disease occurs on marigold, sunower, Jerusalem artichoke, and common
ragweed. Apical chlorosis is also caused by this pathogen on sunower and
sunower seed may be a source of inoculum.
Pseudomonas syringae pv. tagetis. Bacterial Leaf Spot on compass plant
and sunower.
Pseudomonas syringae pv. tomato. Bacterial Leaf Spot of crucifers,
Pepper Spot of cabbage, cauliower, chinese cabbage, and turnip, mostly
in northeastern and Middle Atlantic states. Numerous brown or purple spots
range from pinpoint to 1/8 inch in diameter. If spots are very numerous,
leaves yellow and drop off. Cauliower is more commonly affected than
cabbage. Bacteria, disseminated on seed or in diseased plant parts, enter
through stomata, and visible symptoms appear in 3 to 6 days. Disease is
most severe in seedbeds.
Control. Change location of hotbed starting seedlings; use 2-year rotation in
eld; have seed hot-water treated.
Pseudomonas syringae pv. tomato. Bacterial Speck of tomato. Numer-
ous, dark brown raised spots on fruit are very small, less than 1/16 inch; they
do not extend into esh and are more disguring than harmful.
Pseudomonas syringae pv. zizaniae. Leaf Spot and Stem Spot of wild
rice.
Pseudomonas tabaci (see Pseudomonas syringae pv. tabaci). Blackre of
tobacco.
Pseudomonas viburni. Bacterial Leaf Spot of viburnum, widespread.
Circular water-soaked spots appear on leaves, and irregular sunken brown
cankers on young stems, and the bacteria overwinter in leaves, stems or buds.
Remove and burn infected leaves. Spray with bordeaux mixture or an antibi-
otic such as Agrimycin two or three times at weekly intervals.
Pseudomonas viridiava. Bacterial Leaf Spot on basil. Bacterial
Canker on poinsettia.
Pseudomonas viridilivida. Louisiana Lettuce Disease on lettuce, bell
pepper, and tomatoes. Numerous water-soaked leaf spots fuse to infect large
areas, rst with a soft rot, then a dry shriveling. Sometimes outer leaves
are rotted and the heart sound. This bacterium also causes greasy canker of
poinsettia.
Pseudomonas washingtoniae. This bacterium causes spots on leaves of
Washington palm.
Pseudomonas woodsii. Bacterial Spot and Blight of carnation. Leaf
lesions are small, elongated, brown with water-soaked borders, wither-
ing to brown sunken areas, with masses of bacteria oozing out of stom-
ata. They are spread in greenhouses by syringing, and outdoors by rain.
Follow cultural practices suggested under P. caryophylli for carnation
wilt.
Pseudomonas sp. Blueberry Canker reported from Oregon. Reddish
brown to black cankers appear on canes of the previous season; all buds in
the cankered areas are killed; stems are sometimes girdled. Varieties Wey-
mouth, June, and Rancocas are resistant, but Jersey, Atlantic, Scammel,
Coville, and Evelyn are highly susceptible.
Rhizomonas suberifaciens. Corky Rot on lettuce.
Rolstonia solanacearum (formerly Pseudomonas solanacearum). South-
ern Bacterial Wilt, also called Brown Rot, Bacterial Ring Disease,
Slime Disease, Granville Wilt (of tobacco), present in many states but
particularly prevalent in the South, from Maryland around the coast to
Texas. Southern wilt is common on potatoes in Florida but also appears
on many other vegetables bean, lima bean, castor bean, soybean, velvet
bean, beet, carrot, cowpea, peanut, sweetpotato, tomato, eggplant, pepper,
and rhubarb. Ornamentals sometimes infected include ageratum, anthurium,
dwarf banana, garden balsam, geranium, canna, cosmos, croton, chrysan-
themum, dahlia, hollyhock, lead-tree, marigold, nasturtium, Spanish needle,
sunower, and zinnia. The symptoms are those of a vascular disease, with
dwarng or sudden wilting, a brown stain in vascular bundles, and dark
patches or streaks in stems. Often the rst symptom is a slight wilting of
leaves at end of branches in the heat of the day, followed by recovery at
night, but each day the wilting is more pronounced and recovery less until
the plant dies. Young plants are more susceptible than older ones. In potatoes
and tomatoes there may be a brown mushy decay of stems, with bacterial
ooze present. Potato tubers often have a browning of vascular ring, followed
by general decay.
Bacteria live in fallow soil 6 years or more and may persist indenitely in the
presence of susceptible plants. They are spread by irrigation water, in crop
debris, or soil fragments on tools and tractors, or by farm animals. Optimum
temperatures are high, ranging from 77 to 97F, with inhibition of disease
below 55F.
Control. Use northern-grown seed potatoes and Sebago and Katahdin vari-
eties, more resistant than Triumph and Cobbler. Use a long rotation for toma-
toes. Soil can be acidied with sulfur to kill bacteria, followed by liming in
the fall before planting.
Xanthomonas
Small rods, motile with a single polar agellum; form abundant slimy yellow growth.
Most species are plant pathogens causing necroses.
Ralstonia solanacearum, Race 3, Biovar 2. Bacterial wilting of geranium;
also yellowing and stunting.
Xanthomonas albilineans. Leaf Scald of sugarcane (FL, TX).
Xanthomonas axonopodis. Leaf Streak (water soaking) of African lily and
Leaf Blight of onion.
Xanthomonas begoniae (see Xanthomonas campestris pv. begoniae).
Begonia Bacteriosis, leaf spot of brous and tuberous begonias.
Xanthomonas campestris. Black Rot of cruciers, Bacterial Blight, Wilt,
Stump Rot of alder, arabidopsis, asparagus tree fern, avocado, cabbage,
cauliower, broccoli, brussels sprouts, kale, lavender, mustard, radish,
rutabaga, sunower, stock, turnip, and leaf blight of onion. Black rot was
rst observed in Kentucky and Wisconsin about 1890 and is generally dis-
tributed in the country, with losses often 40 to 50% of the total crop. It is
one of the most serious crucifer diseases, present each season but epidemic
in warm, wet seasons.
The bacteria invade leaves through water pores or wounds and progress to
the vascular system. Veins are blackened, with leaf tissue browning in a V-
shape. With early infection plants either die or are dwarfed, with a one-sided
growth. Late infection results in defoliation, long bare stalks with a tuft of
leaves on top. When stems are cut across, they show a black ring, result of the
vascular invasion, and sometimes yellow bacterial ooze. Black rot is a hard
odorless rot, but it may be followed by soft, odorous decays. Primary infec-
tion comes from bacteria carried on seed, or in refuse in soil, but drainage
water, rain, farm implements, and animals aid in secondary infection.
Control. Use seed grown in disease-free areas in the West or treat with hot
water, 122F, 25 minutes for cabbage, 18 minutes for broccoli, cauliower,
and collards. Plan a 3-year rotation with plants other than crucifers, and clean
up all crop refuse.
Xanthomonas campestris. Horse-Radish Leaf Spot. Leaves are spotted
but there is no vascular infection. Also causes leaf spot of Pilea sp., Pellionia
sp. and leaf spot and blight of bird of paradise, white buttery. Also, bacterial
leaf and stem lesions.
Xanthomonas campestris. Bacterial Leaf Spot on cabbage and radish.
Xanthomonas campestris pv. asclepiadis. Bacterial Blight on buttery
weed.
Xanthomonas campestris pv. barbareae. Black Rot of winter-cress (Bar-
barea vulgaris), similar to black rot of cabbage; small greenish spots turn
black.
Xanthomonas campestris pv. begoniae (formerly Xanthomonas begoniae).
Begonia Bacteriosis, leaf spot of brous and tuberous begonias. Blister-
like, roundish dead spots are scattered over surface of leaves. Spots are brown
with yellow translucent margins. Leaves fall prematurely, and in severe cases
the main stem is invaded, with gradual softening of all tissues and death of
plants. Bacteria remain viable at least 3 months in yellow ooze on surface
of dried leaves. Leaves are infected through upper surfaces during watering,
with rapid spread of disease when plants are crowded together under condi-
tions of high humidity.
Control. Keep top of leaves dry, avoiding syringing or overhead watering;
keep pots widely spaced; spray with bordeaux mixture and dip cuttings in
it.
Xanthomonas campestris pv. carotae (formerly Xanthomonas carotae).
Bacterial Blight of carrot. The chief damage is to ower heads grown
for seed, which may be entirely killed. Symptoms include irregular dead
spots on leaves, dark brown lines on petioles and stems, blighting of oral
parts, which may be one-sided. Use clean seed, or treat with hot water; rotate
crops.
Xanthomonas campestris pv. citri. Citrus Canker on all citrus fruits, but
not apparently eradicated from the United States. It came from the Orient
and appeared in Texas in 1910, becoming of major importance in Florida
and the Gulf States by 1914, ranking with chestnut blight and white pine
blister rust as a national calamity. But here is one of the few cases on record
where man has won the ght, where a disease has been nearly eradicated
by spending enough money and having enough cooperation early in the
game. Several million dollars, together with concerted intelligent effort by
growers, quarantine measures, destruction of every infected tree, sanitary
precautions so rigid they included walking the mules through disinfestant,
sterilization of clothes worn by workers ill saved us from untold later
losses.
Symptoms of citrus canker are rough, brown corky eruptions on both sides
of leaves and fruit. On foliage the lesions are surrounded by oily or yellow
halos. Old lesions become brown and corky.
Xanthomonas campestris pv. corylina (formerly Xanthomonas corylina).
Filbert Blight, Bacteriosis, the most serious disease of lberts in the Pacif-
ic Northwest, known since 1913 from the Cascade Mountains west in Oregon
and Washington. The disease is similar to walnut blight (see X. juglandis)
with infection on buds, leaves, and stems of current growth; on branches;
and on trunks 1 to 4 years old. The bacteria are weakly pathogenic to the
nuts. Copper-lime dusts are effective, with four to six weekly applications,
starting at the early prebloom stage.
Xanthomonas campestris pv. cucurbitae (formerly Xanthomonas cucur-
bitae). Bacterial Spot on winter squash and pumpkin. Leaf spots are rst
small and round, then angular between veins, with bright yellow halos; some-
times translucent and thin but not dropping out; often coalescing to involve
whole leaf. Bacterial exudate is present.
Xanthomonas campestris pv. cyamopsidis. Rot of Lithops spp.
Xanthomonas campestris pv. dieffenbachiae. Blight of Anthurium; also
Leaf Spot of cocoyam.
Xanthomonas campestris pv. dieffenbachiae (formerly Xanthomonas dief-
fenbachiae). Dieffenbachia Leaf Spot. Spots are formed on all parts of
leaf blade except midrib, but not on petioles and stems. They range from
minute, translucent specks to lesions 3/8 inch in diameter, circular to elon-
gated, yellow to orange-yellow with a dull green center. Spots may grow
together to cover large areas, which turn yellow, wilt, and dry. Dead leaves
are dull tan to light brown, thin and tough but not brittle. The exudate on
lower surface of spots dries to a waxy, silver-white layer.
Control. Separate infected from healthy plants; keep temperature low; avoid
syringing; try protective spraying with streptomycin.
Xanthomonas campestris pv. fragariae. Angular Leaf Spot on strawber-
ry; also Blossom Blight on strawberry.
Xanthomonas campestris pv. glycines (formerly Xanthomonas glycines
(phaseoli var. sojense). Bacterial Pustule of soybean, similar to regular
bean blight but chiey a foliage disease, present in most soybean areas, more
severe in the South. Small, yellow-green spots with reddish brown centers
appear on upper surface of leaves with a small raised pustule at the center
of the spot on the under leaf surface. Spots run together to large irregu-
lar brown areas, portions of which drop out, giving a ragged appearance.
Bacteria overwinter in diseased leaves and on seed. Variety CNS is highly
resistant; Ogden has some resistance.
Xanthomonas campestris pv. gummisudans (formerly Xanthomonas gum-
misudans). Bacterial Blight of Gladiolus. Narrow, horizontal, water-
soaked, dark green spots turn into brown squares or rectangles between
veins, covering entire leaf, particularly a young leaf, or middle section of
the blade. Bacteria ooze out in slender, twisted, white columns or in a gum-
my lm, in which soil and insects get stuck. Disease is spread by planting
infected corms or by bacteria splashed in rain from infected to healthy
leaves. The small dark brown corm lesions are almost unnoticeable. Soak
corms unhusked for 2 hours before planting.
Xanthomonas campestris pv. hederae (formerly Xanthomonas hederae).
Bacterial Leaf Spot of English ivy. Small water-soaked area on leaves
develop dark brown to black centers as they increase in size, sometimes
cracking, with reddish purple margins. Spots are sometimes formed on peti-
oles and stems, with plants dwarfed and foliage yellow-green. Spray with
bordeaux mixture or an antibiotic. Keep plants well spaced; avoid overhead
watering and high humidity.
Xanthomonas campestris pv. hyacinthi (formerly Xanthomonas hyacinthi).
Hyacinth Yellows, yellow rot of Dutch hyacinth, occasionally entering the
country in imported bulbs. The disease was rst noted in Holland in 1881
and named for the yellow slime or bacterial ooze seen when a bulb is cut.
The bulbs rot either before or after planting, producing no plants above
ground or badly infected specimens, which do not ower and have yellow to
brown stripes on leaves or ower stalks. Bacteria are transmitted by wind,
rain, tools, and clothes, with rapid infection in wet or humid weather, par-
ticularly among luxuriantly growing plants. The disease is usually minor in
our Pacic Northwest but worse in warm, wet weather on rapidly growing
plants. Innocence is more susceptible than King of the Blues.
Control. Cover infected plants with a jar or can until the end of the season;
then dig after the others. Never work or walk in elds when plants are wet;
avoid bruising; discard rotten bulbs; rotate plantings; avoid fertilizer high in
nitrogen.
Xanthomonas campestris pv. incanae (formerly Xanthomonas incanae).
Bacterial Blight of garden stocks causing, since 1933, serious losses on
ower-seed ranches in California; also present in home gardens. This is
a vascular disease of main stem and lateral branches, often extending into
leaf petioles and seed peduncles. Seedlings suddenly wilt when 2 to 4 inches
high, with stem tissues yellowish, soft and mushy, and sometimes a yellow
exudate along stem. On older plants, dark water-soaked areas appear around
leaf scars near ground, stem is girdled, and lower leaves turn yellow and
drop; or entire plants wilt or are broken by wind at ground level. Bacteria
persist in soil and on or in seed;they are also spread in irrigation water.
Control. Use a 2 to 3-year rotation. Treat seed with hot water, 127.5 to
131F for 10 minutes, followed by rapid cooling.
Xanthomonas campestris pv. juglandis (formerly Xanthomonas juglan-
dis). Walnut Blight on English or Persian walnut, black walnut, butternut,
Siebold walnut. Black, dead spots appear on young nuts, green shoots, and
leaves. Many nuts fall prematurely, but others reach full size with husk, shell,
and kernel more or less blackened and destroyed. Bacteria winter in old nuts
or in buds, and may be carried by the walnut erinose mite.
Control. Spray with a xed copper, as copper oxalate, or with streptomycin.
Apply when 10% of the blossoms are open, repeat when 20% are open, and
again after bloom.
Xanthomonas campestris pv. malvacearum. Leaf Spot on Hibiscus.
Xanthomonas campestris pv. oryzae (formerly Xanthomonas oryzae).
Carnation Pimple reported from Colorado as caused by a new form of the
rice blight organism. Very small, 1 mm, pimples are formed near base and
tips of leaves, which may shrivel.
Xanthomonas campestris pv. papavericola (formerly Xanthomonas papa-
vericola). Bacterial Blight of poppy on corn poppy and on Oriental, opi-
um, and California poppies. Minute, water-soaked areas darken to intense
black spots bounded by a colorless ring. Spots are scattered, circular, small,
often zonate, with tissue between yellow and then brown. There is a notice-
able, slimy exudate. Infection is through stomata and often into veins. Stem
lesions are long, very black, sometimes girdling and causing young plants
to fall over. Flower sepals are blackened, petals stop developing; pods show
conspicuous black spots.
Control. Remove and destroy infected plants; do not replant poppies in the
same location. Try Agrimycin as a preventive spray.
Xanthomonas campestris pv. pelargonii (formerly Xanthomonas pelar-
gonii). Bacterial Leaf Spot of geranium (Pelargonium). Irregular to cir-
cular brown leaf spots start as water-soaked dots on undersurface, becoming
sunken as they enlarge and with tissue collapsing. If spots are numerous, the
entire leaf turns yellow, brown, and shriveled, then drops. The leaves some-
times wilt and droop but hang on the plant for a week or so. Exterior of
stem is gray and dull, the pith and cortex black, later disintegrating into a dry
rot. The roots are blackened but not decayed. Cuttings fail to root, and rot
from the base upward. Bacteria can live 3 months in moist soil; are spread
by handling, splashing water, cutting knives, and whiteies.
Control. Remove diseased plants. Take cuttings from plants known to be
healthy; place in sterilized media and pots. Commercial growers should pur-
chase culture-indexed cuttings. Be sure to sterilize cutting knives. Use 1-year
rotation. Try Agrimycin as a preventive spray, or copper.
Xanthomonas campestris pv. pelargonii (formerly Xanthomonas pelar-
gonii) Geranium Leaf Spot on Pelargonium spp. Leaf spots are small,
brown, necrotic, sometimes with reddish tinge on upper surface and a slightly
water-soaked condition on underside. Young leaves may die and drop. Peti-
oles are occasionally spotted. Bacteria winter in old leaves or under mulch.
Xanthomonas campestris pv. phaseoli (formerly Xanthomonas phaseoli).
Bacterial Bean Blight, general and serious on beans but rare in some west-
ern states. Leaf spots are at rst very small, water-soaked or light green wilt-
ed areas, which enlarge, turn brown, are dry and brittle, and have a yellow
border around edge of lesions and often a narrow, pale green zone outside
that. Leaves become ragged in wind and rainstorms. Reddish brown horizon-
tal streaks appear in stem, which may be girdled and break over at cotyledons
or rst leaf node.
Pod lesions are rst dark green and water-soaked, then dry, sunken and brick
red, sometimes with a yellowish encrustation of bacterial ooze. White seeds
turn yellow, are wrinkled with a varnished look.
Control. Use disease-free western-grown seed. Keep away from beans when
plants are wet.
Xanthomonas campestris pv. pruni (formerly Xanthomonas pruni). Bac-
terial Spot of stone fruits, also called canker, shot hole, black spot; general
on plum, Japanese plum, prune, peach, and nectarine east of the Rocky
Mountains; one of the more destructive stone fruit diseases, causing heavy
losses in some states.
Symptoms on leaves are numerous, round or angular, small reddish spots with
centers turning brown and dead, dropping out to leave shot holes. Spots may
run together to give a burned, blighted, or ragged appearance, followed by
defoliation, with losses running high in devitalized trees. On twigs dark blis-
ters dry out to sunken cankers. Fruit spots turn into brown to black, saucer-
shaped depressions with small masses of gummy, yellow exudate, often with
cracking through the spot.
Control. Plant new orchards from nurseries free from the disease. Prune to
allow air in the interior of trees. Feed properly; trees with sufcient nitrogen
do not defoliate so readily. Zinc sulfate-lime sprays have been somewhat
effective.
Xanthomonas campestris pv. raphani (formerly Xanthomonas vesicatoria
var. raphani). Leaf Spot of radish, turnip, and other crucifers, similar to
bacterial spot on tomato.
Xanthomonas campestris pv. vesicatoria (formerly Xanthomonas vesi-
catoria). Bacterial Spot of tomato and pepper, common in wet seasons.
Small, black, scabby fruit spots, sometimes with a translucent border, pro-
vide entrance points for secondary decay organisms. Small, dark greasy
spots appear on leaets and elongated black spots on stems and petioles.
Bacteria are carried on seed.
Control. Rotate crops; destroy diseased vines. Spraying or dusting with cop-
per may reduce infection. These may be combined with streptomycin.
Xanthomonas campestris pv. vignicola (formerly Xanthomonas vignico-
la). Cowpea Canker on cowpeas and red kidney beans, a destructive dis-
ease, rst described in 1944. Beans are blighted; cowpea stems have swollen,
cankerlike lesions,with the cortex cracked open and a white bacterial exu-
date. The plants tend to break over. Leaves, stems, pods, and seeds are liable
to infection. Chinese Red cowpeas seem particularly susceptible, but the dis-
ease appears on other varieties.
Xanthomonas campestris pv. vitians (formerly Xanthomonas vitians).
Bacterial Wilt and Leaf Spot of lettuce, South Carolina Lettuce Dis-
ease, wilting and rotting of lettuce leaves and stems. In early stages plants
are lighter green than normal. Leaves may have denite brown spots coa-
lescing to large areas or may wilt following stem infection. Use windbreaks
to prevent injuries affording entrance to bacteria; also causes leaf spot of
pepper and tomato.
Xanthomonas campestris pv. zinniae. Leaf and Flower Spot of zinnia.
Xanthomonas campestris pv. zinniae (formerly Xanthomonas nigromacu-
lans). Leaf Spot on zinnia.
Xanthomonas carotae (see Xanthomonas campestris pv. carotae). Bacte-
rial Blight of carrot.
Xanthomonas corylina (see Xanthomonas campestris pv. corylina). Filbert
Blight, Bacteriosis, the most serious disease of lberts in the Pacic North-
west, known since 1913 from the Cascade Mountains west in Oregon and
Washington.
Xanthomonas cucurbitae (see Xanthomonas campestriis pv. cucurbitae).
Bacterial Spot on winter squash and pumpkin
Xanthomonas dieffenbachiae (see Xanthomonas campestriis pv. dieffen-
bachiae). Dieffenbachia Leaf Spot. Spots are formed on all parts of leaf
blade except midrib, but not on petioles and stems.
Xanthomonas glycines (phaseoli var. sojense) (see Xanthomonas campestris
pv. glycines). Bacterial Pustule of soybean, similar to regular bean blight
but chiey a foliage disease, present in most soybean areas, more severe in
the South
Xanthomonas gummisudans (see Xanthomonas campestris pv. gummisu-
dans). Bacterial Blight of Gladiolus.
Xanthomonas hederae (see Xanthomonas campestris pv. hederae). Bacte-
rial Leaf Spot of English ivy.
Xanthomonas hyacinthi (see Xanthomonas campestris pv. hyacinthi). Hya-
cinth Yellows, yellow rot of Dutch hyacinth, occasionally entering the
country in imported bulbs.
Xanthomonas incanae (see Xanthomonas campestris pv. incanae). Bacte-
rial Blight of garden stocks causing, since 1933, serious losses on ower-
seed ranches in California; also present in home gardens.
Xanthomonas juglandis (see Xanthomonas campestris pv. juglandis). Wal-
nut Blight on English or Persian walnut, black walnut, butternut, Siebold
walnut.
Xanthomonas oryzae (see Xanthomonas campestris pv. oryzae). Carna-
tion Pimple reported from Colorado as caused by a new form of the rice
blight organism. Xanthomonas papavericola (see Xanthomonas campestris
pv. papavericola). Bacterial Blight of poppy on corn poppy and on Orien-
tal, opium, and California poppies.
Xanthomonas pelargonii (see Xanthomonas campestris pv. pelargonii).
Bacterial Leaf Spot of geranium (Pelargonium).
Xanthomonas pelargonii (see Xanthomonas campestris pv. pelargonii).
Geranium Leaf Spot on Pelargonium spp.
Xanthomonas phaseoli (see Xanthomonas campestris pv. phaseoli). Bac-
terial Bean Blight, general and serious on beans but rare in some western
states.
Xanthomonas pruni (see Xanthomonas campestris pv. pruni. Bacterial
Spot of stone fruit, also called canker, shot hole, black spot; general on
plum, Japanese plum prune, peach, and nectarine east of the Rocky Moun-
tains.
Xanthomonas vesicatoria (see Xanthomonas campestris pv. vesicatoria.
Bacterial Spot of tomato and pepper, common in wet seasons.
Xanthomonas vesicatoria var. raphani (see Xanthomonas campestris pv.
raphani). Leaf Spot of radish, turnip, and other crucifers, similar to bacte-
rial spot on tomato.
Xanthomonas vignicola (see Xanthomonas campestris pv. vignicola).
Cowpea Canker on cowpeas and red kidney beans.
Xanthomonas vitians (see Xanthomonas campestris pv. vitians). Bacterial
Wilt and Leaf Spot of lettuce, South Carolina Lettuce Disease, wilting
and rotting of lettuce leaves and stems.
Xanthomonas nigromaculans (see Xanthomonas campestris pv. zinniae).
Leaf Spot on zinnia.
Xylella fastidiosa. Bacterial Leaf Scorch on maple, pecan, mulberry,
northern red oak and sweet gum.
Pierces Grape Disease. First described as California vine disease by Pierce
in 1892, now known as cause of grape degeneration in Gulf states; report-
ed from Rhode Island. First symptoms are scalding and browning of leaf
tissues, often with veins remaining green; canes die back from tips in late
summer; growth is dwarfed, fruit shriveled; roots die. The bacterium invades
the xylem and turns it brown. Alfalfa plants are stunted with short stems and
small leaves. Many species of sharpshooter leafhoppers transmit the bac-
terium to grape from alfalfa, clovers, grasses, also from ivy, acacia, fuch-
sia, rosemary, zinnia, and other ornamentals that are symptomless carriers.
There is no adequate control; roguing of diseased vines and spraying for
leafhoppers has proved ineffective. Propagate by cuttings from disease-free
vineyards.
Mycoplasmataceae
Phytoplasma
Ash Yellows and Witches Broom. On ash in Michigan, Montana, Nebraska,
North Carolina, and South Dakota and peanut in Oklahoma.
Aster Yellows. Throughout the United States, also called Lettuce Rio Grande
Disease, Lettuce White Heart, Potato Purple Top.
Bean Phyllody. Perhaps caused by a strain of aster-yellows MLO.
California Aster Yellows. In the West, also known as Celery Yellows, West-
ern.
Aster Yellows, Potato Late Break, Strawberry Green Petal. Aster yellows
may appear in more than 170 species of 38 families of dicotyledons. It is
serious on China aster, may also affect anemone, calendula, coreopsis, cos-
mos, purple coneower (Echinacea), delphinium, daisies, golden-glow, hy-
drangea, marigold, petunia, phlox, scabiosa, strawower, and other owers.
It is serious on lettuce, alfalfa, endive, carrot, parsley, New Zealand spinach,
radish, and some other vegetables, but not on peas, beans, or other legumes.
This disease is now known to be caused by a phytoplasma organism.
In most plants vein clearing is followed by chlorosis of newly formed tis-
sues, adventitious growth, erect habit, virescence of owers. Asters have
a stiff yellow growth with many secondary shoots; are stunted, with short
internodes; owers are greenish, dwarfed, or none. The chief vector is the
six-spotted leafhopper (Macrosteles fascifrons). The virus multiplies in the
insect, and there is a delay of 10 days or more after the insect feeds on a dis-
eased plant before it can infect a healthy specimen. There is no transmission
through insect eggs or aster seeds.
Celery petioles are upright, somewhat elongated, with inner petioles short,
chlorotic, twisted, brittle, often cracked, yellow. The celery strain of the virus
causes yellowing and stunting of cucumber, squash, pumpkin; infects gladi-
olus and zinnia.
Control of aster yellows is directed against the leafhoppers. Asters are grown
commercially under frames of cheesecloth, 22 threads to the inch, or wire
screening, 18 threads to the inch. In home gardens all diseased plants should
be rogued immediately and overwintering weeds, which harbor leafhop-
per eggs, destroyed. Spraying or dusting ornamentals and vegetables with
pyrethrum will reduce the number of vectors but will not entirely eliminate
the disease.
Recent work raises the probability that the etiological agent of aster yellows
is a mycoplasma rather than a virus. Therefore, treatment with antibiotics,
such as chlortetracycline, has suppressed the development of yellows symp-
toms. Mycoplasma-like bodies have been seen in microscopic study of dis-
eased plants and in transmitting leafhopper vectors, but not in healthy plants
or nontransmitting vectors.
Clover Proliferation. On strawberry and onion.
Corn Stunt. A dwarng disease present primarily in the South; transmitted
by leafhoppers. Mycoplasma-like bodies present; See Spiroplasma citri.
Elm Phloem Necrosis. On American elm from West Virginia and Georgia
to northern Mississippi, eastern Oklahoma, Kansas, and Nebraska. Origin
unknown but apparently present since 1882; the disease reached epidemic
proportions in Ohio in 1944, killing 20,000 trees that year near Dayton and
10,000 at Columbus. The most reliable diagnostic character is a buttercup
yellow discoloration of the phloem, often ecked with brown or black and
an odor of wintergreen. Destruction of phloem causes the bark to loosen and
fall away. Roots die rst, then the phloem in lower portions of tree, followed
by wilting and defoliation. American elms may be attacked at any age; they
wilt and die suddenly within 3 or 4 weeks or gradually decline for 12 to 18
months. This is now thought to be caused by a mycoplasma-like agent.
Transmission is by the white-banded elm leafhopper (Scaphoideus luteolus)
and possibly other species. Nymphs hatch about May 1 from eggs wintered
on elm bark and feed on leaf veins. Adults move from diseased to healthy
trees.
There is hope of propagating elms resistant to phloem necrosis. Communities
should interplant existing elms with Asiatic or European varieties or with
some other type of tree to provide shade if and when present elms die.
Peach Western X-Disease. Perhaps same as X-disease but usually treated
separately; also known as cherry buckskin and western-X little cherry. The
pathogen is transmitted by leafhoppers (Colladonus germinatus, Fieberella
orii, Osbornellus borealis, and others) to peach, nectarine and cherry in
western states. Symptoms vary according to rootstock, but cherry fruit is
smaller than normal. Sour cherries are puttylike, pinkish; sweet cherries are
small, conical, hang on trees late, fail to develop normal color. Symptoms on
peach are similar to those of X-disease.
Peach X-Disease. On peach and chokecherry, sometimes cherry in the north-
ern United States and of major importance in Connecticut, Massachusetts,
and New York. Peach trees appear normal in spring for 6 or 7 weeks after
growth starts, then foliage shows a diffused yellow and red discoloration
with a longitudinal upward curling of leaf edges; spots may drop out, leav-
ing a tattered effect. Defoliation starts by mid-summer. Fruits shrivel and
drop or ripen prematurely. Seed do not develop. Weakened trees are killed
by low temperatures or remain unproductive.
Chokecherry has conspicuous premature reddening of foliage, dead embryos
in fruit. The second and third seasons after infection foliage colors are duller,
there are rosettes of small leaves on terminals, and death may follow. Natural
infection is apparently from chokecherry to peach (not peach to peach or
peach to chokecherry) by a leafhopper (Colladonus clitellarius). Elimination
of chokecherries within 500 feet of peach trees provides the best control.
Peach Yellow Leaf Roll; a form of Western X-Disease; perhaps caused by
a more severe strain of the MLO.
Peach Yellows; Little Peach. First noted near Philadelphia in 1791 and so
serious that in 1796 the American Philosophical Society offered a $60 prize
for the best method of preventing premature decay of peach trees. Present in
eastern states on peach, almond, nectarine, apricot and plum. Not found west
of the Mississippi or in the South. In peach, clearing of veins, production
of thin erect shoots with small chlorotic leaves, premature ripening of fruit
(with reddish streaks in esh and insipid taste) is followed by death of the
tree in a year or so. The little peach strain of the MLO causes distortion
of young leaves at tips of branches, small fruit, delayed ripening. Plum is
systemically infected, with few obvious symptoms. Transmission is by the
plum leafhopper or budding.
Control. Budsticks and dormant nursery trees can be safely treated with heat
sufcient to kill the MLO (122F for 5 to 10 minutes), but cured trees are
susceptible to reinfection. Most effective control is removal of wild plum
trees around peach orchard and spraying to control leafhoppers.
Potato Apical Leaf Roll and Arizona Purple Top Wilt. Caused by aster
yellows.
Strawberry Green Petal. Perhaps due to a strain of aster yellows MLO, as
is chlorotic phyllody reported from Louisiana. Flowers have enlarged sepals,
small green petals.
Bud Proliferation and Delayed Maturity, on soybean.
Decline of ash.
Lethal Yellowing on palms.
Phloem Necrosis of chrysanthemum.
Spiroplasma citri. Corn Stunt. Has been reported on corn, onions,
horseradish, shepherds purse, yellow rocket, and wild mustard.
Stunt of blueberry.
Virescence on horseradish.
Witches Broom on pigeon pea (Cajanus cajan), and black raspberry.
Witches Broom on Japanese persimmon, and lilac.
Witches Broom and Yellowing on annual statice.
Yellows of elm.
BLACK KNOT
The term black knot is used to designate a disease with black knotty excres-
cences.
Apiosporina
Ascomycetes, Pleosporales
Asci are in locules, without well-marked perithecial walls, immersed in a massive, car-
bonaceous stroma, erumpent and supercial at maturity. Spores are hyaline, unequally
two-celled.
Apiosporina morbosa (formerly Dibotryon morbosum). Black Knot of
plum and cherry, Prunus Black Knot, Plum Wart, widespread and serious
on garden plums, also present on sweet and sour cherries, chokecherry, and
apricot. Apparently a native disease, destructive in Massachusetts by 1811
and the pathogen described from Pennsylvania in 1821, black knot has been
reported on peach, long thought to be immune.
The chief symptoms are black, rough, cylindrical or spindle-shaped enlarge-
ments of twigs into knots two to four times their thickness and several inches
long (see Fig. 3.6). Infection takes place in spring, but swelling is not evi-
dent until growth starts the following spring, at which time the bark ruptures,
and a light yellowish growth lls the crevices. In late spring this is covered
with an olive green, velvety layer made up of brownish conidiophores and
one-celled hyaline conidia of the anamorph Hormodendron state. Conidia
are spread by wind.
In late summer black stromata cover the affected tissues, and the galls
become hard. Asci are formed during the winter in cavities in the stroma;
ascospores are discharged and germinate in early spring, completing the
2-year cycle. Knots are produced from primary infection by ascospores or
from secondary infection from mycelium formed in old knots and growing
out to invade new tissue. Limbs may be girdled and killed; trees are stunted
BLACK KNOT 135
Figure 3.6 Black Knot on Prunus sp
and dwarfed, nearly worthless after a few years. Old knots may be riddled
with insects or covered with a pink fungus growing on the Apiosporina
mycelium.
Control. Cut out infected twigs and branches, 3 or 4 inches beyond the knot,
to include advancing perennial mycelium. Do this in winter or before April
1. Eradicate or thoroughly clean up wild plums and cherries in the vicinity.
Spray at delayed dormant stage in spring (just as buds break) with bordeaux
mixture or with liquid lime sulfur. The latter is preferable unless oil is com-
bined in the spray as an insecticide. Spray with lime sulfur at full bloom.
Dibotryon morbosum (see Apiosporina morbosa). Black Knot of plum
and cherry, Prunus Black Knot, Plum Wart, widespread and serious on
garden plums, also present on sweet and sour cherries, chokecherry, and apri-
cot.
Leptosphaeria
Perithecia in clusters on wood; spores dark, with several cells.
Gibberidea heliopsidis (see Leptosphaeria heliopsidis). Black Knot,
Black Patch on goldenrod and sunower.
Leptosphaeria heliopsidis (formerly Gibberidea heliopsidis). Black Knot,
Black Patch on goldenrod and sunower.
BLACKLEG
The term blackleg is used to describe darkening at the base of a stem or plant.
Blackleg of potatoes and delphinium are described under Bacterial Diseases;
blackleg of geraniums is under Rots.
Cylindrocarpon
Rots.
Cylindrocarpon obtusisporum. Blackleg; on grape.
Phoma
Pycnidia dark, ostiolate, lenticular to globose, immersed in host tissue, erumpent or with
short beak piercing the epidermis; conidiophores short or obsolete, conidia small, one-
celled, hyaline, ovate to elongate; parasitic on seed plants, chiey on stems and fruits,
rarely on leaves.
Phoma lingam. Blackleg of crucifers, Foot Rot, Phoma Wilt of plants of
the mustard family, including cabbage, cauliower, Chinese cabbage, brus-
sels sprouts, charlock, garden cress, pepper grass, kale, kohlrabi, mustard,
rape, radish, rutabaga, turnip, stock, and sweet alyssum. The teleomorph
state, Lystosphaeria maculans has been found on cabbage.The fungus was
rst noticed in Germany in 1791; the disease was reported in France in 1849,
and in the United States in 1910. It is generally distributed east of the Rocky
Mountains and formerly caused from 50 to 90% loss. With improved seed
and seed treatment it has become less important.
The rst symptom is a sunken area in the stem near the ground, which
extends until the stem is girdled and the area turns black. Leaves, seed stalks,
and seed pods have circular, light brown spots. Small black pycnidia appear-
ing on the lesions distinguish blackleg from other cabbage diseases. The
leaves sometimes turn purple and wilt, but there is no defoliation, as in black
rot.
The fungus reaches the soil via infected plant debris, remaining alive 2 or
more years. Spores are spread by splashing rain, or manure, on tools, and
perhaps by insects, with new lesions resulting in 10 to 14 days. But the chief
spread is by mycelium wintering in infected seed. When such seed is planted,
fruiting bodies are formed on cotyledons as they are pushed above ground,
and these serve as a source of inoculum for nearby plants. A few diseased
seed can start an epiphytotic in wet weather.
Control. Use seed grown on the Pacic Coast, which is usually, although
not always, disease-free. If the seed is infected, tie loosely in cheesecloth
bags and immerse in hot water, held at 122F for 30 minutes. It is some-
times possible to buy seed already treated. Sterilize soil for the seedbed; use
a 3-year rotation; do not splash seedlings when watering; do not transplant
any seedlings if the disease shows up in the seedbed; do not feed cabbage
refuse to cattle; do not transfer cultivators and other tools from a diseased to
a healthy eld without using a disinfestant.
Leptosphaeria
Leptosphaeria maculans. Blackleg on canola.
BLACK MILDEW
The terms black mildew, sooty mold, and black spot have been used to some
extent interchangeably. In this text the term sooty mold is restricted to those
fungi living on insect exudate and hence not true parasites. Included here
under Black Mildew are parasitic fungi that have a supercial dark myceli-
um. They are members of the Erysiphales (Meliolales according to some
classications) and hence similar to powdery mildews except for the dark
color, or they belong to the Hemisphaeriales, characterized by a dark stroma
simulating the upper portion of a perithecium. In a few cases the diseases are
called black spot rather than mildew.
Apiosporina
Ascomycetes, Pleosporales
Perithecia and mycelium supercial; mycelium with setae and perithecia usually hairy;
paraphysoids present; spores two-celled; dark.
Apiosporina collinsii. Witches Broom of serviceberry (Amelanchier)
widespread. Perennial mycelium stimulates the development of numerous
stout branches into a broom. A sooty growth on underside of leaves is
rst olive brown, then black. Numerous globose, beadlike, black perithecia
appear in late summer. The damage to the host is not serious.
Asterina
Ascomycetes, Asterinales
Asterina species are parasites on the surface of leaves and are usually found in warm
climates. In some cases the disease is called black mildew, in others, black spot. The
perithecia are dimidiate, having the top half covered with a shield, a small, round stro-
ma composed of radially arranged dark hyphae. Underneath this stromatic cover, called
scutellum, there is a single layer of fruiting cells; paraphyses are lacking; spores are
dark, two-celled. The mycelium, which is free over the surface, has lobed appendages,
hyphopodia, which act as haustoria in penetrating the cuticle and obtaining nourishment
from the host.
Asteridium lepidigenum (formerly Asterina lepidigena). Black Mildew
on lyonia, Florida.
Asterina anomala (see Limacinula anomala). Black Mildewon California-
laurel, California.
Asterina delitescens. Black Spot on redbay.
Asterina diplopoides. Black Spot on leucotho.
Asterina gaultheriae (see Schizothyrium pomi). Black Mildewon bearber-
ry, Wisconsin.
Asterina lepidigena (see Asteridium lepidigenum). Black Mildew on
lyonia, Florida.
Asterina orbicularis. Black Spot on American holly and Ilex spp.
Limacinula anomala (formerly Asterina anomala). Black Mildew on
California-laurel, California.
Schizothyrium pomi (formerly Asterina gaultheriae). Black Mildew on
bearberry, Wisconsin.
Asterinella
Ascomycetes, Asterinales
Like Asterina but lacking hyphopodia; with or without paraphyses; spores dark, two-
celled.
Asterinella puiggarii. Black Spot on eugenia.
Dimerosporium
According to some authorities this is the same as Asterina but the name
Dimerosporium is in common use.
Dimerosporium abietis (see Rasutoria abietiis). Black Mildew on Pacic
silver and lowland white rs.
Dimerosporium hispidulum. Black Mildew on boxelder.
Dimerosporium pulchrum. Black Mildew on ash.
Dimerosporium robiniae. Black Mildew on ailanthus.
Dimerosporium tropicale. Black Mildew on bignonia, Mississippi.
BLACK MILDEW 141
Rasutoria abietiis (formerly Dimerosporium abietis). Black Mildew on
Pacic silver and lowland white rs. Black patches are formed on older nee-
dles, usually on under surface. There is no apparent injury to trees.
(Irene) Asteridiella
Ascomycetes, Meliolales
Mycelium with capitate hyphopodia but no bristles; perithecia with larviform append-
ages; spores dark, with several cells.
Appendiculella araliae (formerly Irene araliae). Black Mildew on mag-
nolia, Mississippi.
Appendiculella calostroma (formerly Irene calostroma). Black Mildewon
wax-myrtle, Gulf States.
Appendiculella perseae (formerly Irene perseae). Black Mildew on avo-
cado, Florida.
Like Irene except that perithecia have no appendages.
Asteridiella manca (formerly Irenina manca). Black Mildew on wax-
myrtle, Mississippi.
Irene araliae (see Appendiculella araliae). Black Mildew on magnolia,
Mississippi.
Irene calostroma (see Appendiculella calostroma). Black Mildewon wax-
myrtle, Gulf States.
Irene perseae (see Appendiculella perseae). Black Mildew on avocado,
Florida.
Irenina manca (see Asteridiella manca). Black Mildew on wax-myrtle,
Mississippi.
Lembosia (Morenoella)
Brown vegetative mycelium with hyphopodia on surface of host; linear stroma, scutel-
lum, over single layer of fruiting cells; paraphyses present; spores dark, two-celled.
Echidnodella angustiformis (formerly Morenoella angustiformis). Black
Mildew on holly (Ilex spp.), Mississippi.
Echidnodella rugispora (formerly Lembosia rugispora). Black Mildewon
redbay, swampbay, Mississippi, North Carolina.
Lembosia cactorum. Black Mildew on cactus, Florida.
Lembosia coccolobae. Black Mildew on sea-grape, Florida; also L. por-
toricensis and L. tenella.
Lembosia illiciicola. Black Mildew on anise-trees, Alabama, Mississippi.
Lembosia rugispora (see Echidnodella rugispora). Black Mildew on red-
bay, swampbay, Mississippi, North Carolina.
Morenoella angustiformis (see Echidrodella angustiformis). Black Mil-
dew on holly (Ilex spp.), Mississippi.
Schiffnerula pulchra. On dogwood.
Meliola
Ascomycetes, Erysiphales (or Meliolales), Meliolaceae
Most abundant in tropics. Supercial dark mycelium with hyphopodia and setae;
perithecia globose, coal black without ostiole or appendages but often with setae;
spores several-celled, dark; paraphyses lacking. Conidia are lacking in most species, of
Helminthosporium type in others.
Diplotheca tunae (formerly Meliola wrightii). Black Mildew on chinaber-
ry.
Irenopsis cryptocarpa (formerly Meliola cryptocarpa). Black Mildew on
gordonia.
Irenopsis martiniana (see Meliola martiana). Black Mildew on redbay,
swampbay, Alabama, Florida, Mississippi.
Meliola amphitricha. Black Mildew on boxelder, magnolia, redbay,
swampbay.
Meliola bidentata. Black Mildew on bignonia.
Meliola camelliae. Black Mildewof camellia. Abundant black growth may
cover camellia leaves and twigs. Spraying with a light summer oil is some-
times effective.
Meliola cookeana. Black Mildew on callicarpa, lantana.
Meliola cryptocarpa (see Irenopsis cryptocarpa). Black Mildew on gor-
donia.
Meliola lippiae. Black Mildew on lippia.
Meliola magnoliae. Black Mildew on magnolia.
Like Irene except that mycelium has setae (stiff bristles) and perithecia lack
larviform appendages.
BLACK MILDEW 143
Meliola martiana (formerly Irenopsis martiniana). Black Mildew on red-
bay, swampbay, Alabama, Florida, Mississippi.
Meliola nidulans. Black Mildew on blueberry, wintergreen.
Meliola palmicola. Black Mildew on palmetto.
Meliola tenuis. Black Mildew on bamboo.
Meliola wrightii (see Diplotheca tunaei). Black Mildew on chinaberry.
Sthughesia
Perithecia smooth; spores two-celled, dark; paraphyses lacking.
Dimerium juniperi (see Sthughesia juniperi). Black Mildew on Rocky
Mountain juniper, California.
Sthughesia juniperi (formerly Dimerium juniperi). Black Mildew on
Rocky Mountain juniper, California.
BLACKSPOT
In common usage the term black spot without qualifying adjectives has come
to mean but one disease, rose black spot, with the two words currently written
as one, blackspot. This section is limited to the rose disease. Delphinium
black spot will be found under Bacterial Diseases, elm black spot under Leaf
Spots, other black spots under Black Mildew.
Diplocarpon
Ascomycetes, Helotiales, Dermateaceae (Mollisiaceae)
Apothecia innate, formed in dead leaves, but at maturity rupturing overlying tissues;
horny to leathery with a thick margin or outer wall (excipulum) of dark, thick-walled
cells; spores two-celled, hyaline; paraphyses present. Anamorph state is a Marssonina
with two-celled hyaline spores in an acervulus.
Diplocarpon rosae. Rose Blackspot, general on rose but less serious in
the semi-arid Southwest; reported from all states except Arizona, Nevada,
and Wyoming.
For nearly 100 years the fungus was known only by its anamorph state,
which has had about 25 different names. The rst denite record is by Fries
in Sweden in 1815, under the name Erysiphe radiosum, but the rst valid
description was by Libert in 1827 as Asteroma rosae. Later Fries called it
Actinonema rosae, and that term was widely used until Actinonema species
were transferred to Marssonina. The blackspot fungus was rst reported in
the United States in 1831, from Philadelphia, and in 1912 Wolf made the
connection with the teleomorph state, so that the correct name became Diplo-
carpon rosae.
Blackspot is probably the most widely distributed and best known rose dis-
ease. It is conned to roses, garden and greenhouse, and may affect practi-
cally all varieties, although not all are equally susceptible. There has been
some progress made in breeding resistant varieties, but recent investigation
BLACKSPOT 145
Figure 3.7 Rose Blackspot. Note mbriate margin to spot
disclosing many physiological races of the fungus explains why roses that
are almost immune to blackspot in one location may succumb in another.
Rosa bracteata is the only species thus far shown to be reasonably resistant
to all the different races tested. Roses with the Pernetiana parentage, which
has given us the lovely yellows, coppers, and blends, are especially prone
to blackspot. Some roses, like Radiance, are tolerant of blackspot, usually
holding their leaves, even though they cannot be considered resistant.
Symptoms are primarily more or less circular black spots, up to 1/2 inch
in diameter, with radiating mbriate or fringed margins (see Fig. 3.7). This
mbriate margin is a special diagnostic character, differentiating blackspot
from other leaf spots and from discolorations due to cold or chemicals. The
spots vary from one or two to a dozen or more on a leaf, usually on the upper
surface. With close examination you can see small black dots or pimples
in the center of the spots. These are the acervuli, bearing conidia, and they
glisten when wet (see Fig. 3.8).
In susceptible varieties the appearance of black spots is soon followed by
yellowing of a portion or all of leaets and then by defoliation. The leaf
Figure 3.8 Rose Blackspot. Two-celled conidia formed in Acervulus under cuticle
fall is apparently correlated with increased production of ethylene gas in
diseased tissue and perhaps by a difference in auxin gradient between leaf
and stem. Some roses lose almost all their leaves, put out another set and
lose those, and often are trying to leaf out for the third time by late sum-
mer. The process is so devitalizing that some bushes may die during the
following winter. On tolerant varieties leaf spots are present, though usually
in smaller numbers, but there is much less yellowing and defoliation. Cane
lesions are small indistinct black areas, slightly blistered, without mbriate
margins.
Infection occurs through either leaf surface, the fungus sending its germ tube
directly through the cuticle by mechanical pressure. The hyphae form a net-
work under the cuticle, joining together into several parallel laments radiat-
ing from the point of infection. The hyphae are actually colorless, the black
color of the spot coming from the death and disorganization of host cells.
The mycelial growth is between cells, with haustoria (suckers) invading epi-
dermal and palisade cells for nourishment.
Acervuli, summer fruiting bodies, formed just under the cuticle, bear two-
celled hyaline conidia on short conidiophores on a thin, basal stroma.
Splashed by rain or overhead watering, or spread by gardeners working
among wet plants, the conidia germinate and enter a leaf if there is contin-
ued moisture for at least 6 hours. Rain, heavy dew, fog, and sprinklers used
late in the day so foliage does not dry off before night provide the requisite
moisture. New spots show up within a week and new spores within 10 days.
Secondary cycles are repeated all summer from late May to late October
around New York City.
In my personal experience, the spread of disease is most rapid where large
numbers of susceptible varieties are massed together. If all the yellows, for
BLACKSPOT 147
instance, are planted together, the disease gets such a head start, and builds
up so much inoculum to spread to the more tolerant red and pink varieties
nearby, that these varieties also are more heavily infected than usual. When
roses are mixed in beds so that one or two particularly susceptible bushes are
surrounded by more resistant types, the infective material cannot increase
so rapidly, and the net result is less disease in the garden as a whole. Pro-
tected corners in the garden where air circulation is poor also increase the
disease potentiality. Spores are apt to be splashed farther when water hits
hard-packed soil without a mulch.
When old leaves drop to the ground, the mycelium continues a saprophyt-
ic existence, growing through dead tissue with hyphae that are now dark
in color. In spring three types of fruiting bodies may be formed: microac-
ervuli or spermagonia containing very small cells that perhaps act as male
cells; apothecia, the sexual fruiting bodies formed on a stroma between the
epidermis and palisade cells and covered with a circular shield of radiating
strands; and winter acervuli, formed internally and producing new conidia
in spring. The Diplocarpon or apothecial stage is apparently not essential;
it is known only in northeastern United States and south-central Canada.
The shield over the apothecium ruptures, and the two-celled ascospores are
forcibly discharged into the air to infect lowest leaves.
Where the sexual stage is not formed, primary spring infection comes from
conidia splashed by rain to foliage overhead, from acervuli either in over-
wintered leaves on the ground or in cane lesions. New roses from a nursery
sometimes bring blackspot via these cane lesions to a garden previously free
of disease.
Control. The importance of sanitation may have been somewhat over-
stressed; it cannot replace routine spraying or dusting. It is certainly a good
idea to pick off for burning the rst spotted leaves, if this is done when
bushes are dry so that the act of removal does not further spread the fun-
gus. Raking up old leaves from the ground at the end of the season makes
the garden neater and may reduce the amount of inoculum in spring, but,
because the fungus winters also on canes in most sections of the country,
removal of leaves cannot be expected to provide a disease-free garden the
next season. Comparative tests have shown that fall cleanup is ineffectu-
al. A good mulch, applied after uncovering and the rst feeding in spring,
serves as a mechanical barrier between inoculum from overwintered leaves
on the ground and developing leaves overhead. A mulch also reduces disease
by reducing the distance spores can be splashed from one bush to another
during the season. Drastic spring pruning, far lower than normal, reduces the
amount of inoculum from infected canes.
The importance of a dormant spray is debatable. Experiments have shown
that as a true eradicant, applied in winter, it has little value in reducing the
amount of blackspot the next summer. Use liquid lime sulfur after pruning,
provided the buds have not broken far enough to show the leaets.
Summer spraying or dusting, weekly throughout the season (from late April
to early November in New Jersey) is essential if you want to keep enough
foliage on bushes for continuous production of ne owers (it takes food
manufactured in several leaves to produce one bloom) and for winter sur-
vival. Some strong varieties will, however, live for years without chemical
treatment; they are usually scraggly bushes with erratic bloom. The idea that
oribunda varieties do not require as much spraying as hybrid teas is a mis-
conception. Some oribundas are quite resistant; others are very susceptible.
The same holds true for old-fashioned shrub roses. All too often blackspot
gets a head start in a garden from shrub roses we thought it unnecessary to
spray.
Roses can be defoliated as readily by chemicals as by the blackspot fungus;
so the fungicide chosen must be safe under the conditions of applications
as well as effective. There are many chemicals that will control blackspot
if they are applied regularly and thoroughly. Choice depends somewhat on
climate. Some copper sprays and dusts cause red spotting and defoliation
in cool, cloudy weather. Bordeaux mixture is both unsightly and harmful,
unless used in very weak dilution. At strengths recommended for vegetables
it will quickly turn rose leaves yellow and make them drop off. Dusts con-
taining more than 3 to 4% metallic copper are injurious under some weather
conditions. Dusting sulfur ne enough to pass through a 325-mesh screen has
been successfully used for years for blackspot control, but in hot weather it
burns margins of leaves. Copper and sulfur have a synergistic effect; a mix-
ture of the two is more effective than either used alone, but such a mixture
also combines injurious effects.
There are literally hundreds of combination rose sprays and dusts on the
market under brand names, and it seems to me easier, and even cheaper, con-
sidering the time saved, for home gardeners to make use of them to control
blackspot and other rose diseases as well as insects in one operation. You will
have to determine by trial and error the best combination for your area, and
you may not nd one that combines remedies for all the pests you may have
to ght through the season. Choose one that contains ingredients required
BLACKSPOT 149
every week all summer, and then add other chemicals if and when necessary.
Whatever mixture is chosen, coverage should be complete on upper and low-
er leaf surfaces, and applications must be repeated at approximately weekly
intervals. This may mean every 5 or 6 days when plants are growing rapidly
in a rainy spring and perhaps every 7 to 9 days in dry weather, when growth
is slow. Intervals of 10 to 14 days between sprays seldom give adequate con-
trol. Most directions call for application ahead of rain so that the foliage will
be protected when spores germinate during the rain; but if sprays are applied
every 7 days, there will always be enough residue left on the foliage to give
protection during the next rain. It is not necessary to make an additional
application immediately after a rain if your spraying is on a regular basis.
BLIGHTS
According to Webster, blight is any disease or injury of plants resulting in
withering, cessation of growth and death of parts, as in leaves, without rot-
ting. The term is somewhat loosely used by pathologists and gardeners to
cover a wide variety of diseases, some of which may have rotting as a sec-
ondary symptom. In general, the chief characteristic of a blight is sudden
and conspicuous leaf and fruit damage, in contradistinction to leaf spotting,
where dead areas are denitely delimited, or to wilt due to a toxin or oth-
er disturbance in the vascular system. Fire blight, discussed under Bacterial
Diseases, is a typical blight, with twigs and branches dying back but holding
withered, dead foliage.
Alternaria
Deuteromycetes, Hyphomycetes
Dark, muriform conidia formed in chains, simple or branches, or sometimes singly, on
dark, simple conidiophores growing from dark hyphae (see Fig. 3.9). The apical portion
of each conidium is narrowed and often elongated, bearing at its tip the next ovoid,
tapering conidium. Species with this characteristic formerly placed in Macrosporium
are now in Alternaria; those with spores rounded at both ends have been transferred to
Stemphylium.
There are many saprophytic species in Alternaria, the spores of which are
wind-borne for many miles and are a common cause of hayfever. There are
also parasitic forms causing blights and leaf spots. Sometimes the disease
starts as a leaf spot, but the lesions, typically formed in concentric circles,
run together to form a blight, the dark conidia making the surface appear
dark and velvety.
Alternaria alternata. Blight, Foliage and Pod of pea.
Alternaria cassiae. Seedling Blight of Cassia (sicklepod, and coffee sen-
na), and showy crotalaria.
BLIGHTS 151
Figure 3.9 Conidial Production Among Some Fungi Causing Blights. Alternaria, dark muriform spores in chains;
Botrytis, hyaline spores in clusters; Cercospora, pale to dark septate spores on dark conidia protruding from
stomata; Entomosporium, peculiarly appendaged spores in acervulus; Ovulina, hyaline spore with basal disjunc-
tor cell, borne free on mycelium; Pestalotia, in acervulus, median cells colored, end cells hyaline, apical cell
with appendages; Phomopsis, oval and liform hyaline spores in pycnidium; Septoria, septate hyaline spores in
Pycnidium; Volutella, hyaline spores formed on a hairy sporodochium
Alternaria cucumerina. Alternaria Blight of Cucurbits, Cucumber
Blight, Black Mold, general on cucumbers, muskmelon, watermelon, and
winter and summer squash. Symptoms appear in the middle of the season,
rst nearest the center of the hill. Circular brown spots with concentric rings
are visible only on upper surface of leaves, but a black, moldy growth, made
up of conidiophores and large brown spores, can be seen on both leaf sur-
faces. Leaves curl and dry up, cantaloupe foliage being more sensitive than
that of other cucurbits. The disease spreads rapidly in warm, humid weather,
and, with the vines drying, the fruit is exposed to sunburn. Sunken spots
develop on the fruit, covered with an olive green mass of conidia. Other
species of Alternaria cause a decay of melons in transit and storage.
Control. Purdue 44 and some other varieties of muskmelon are rather resis-
tant.
Alternaria dauci. Alternaria Blight of carrot, Carrot Leaf Blight, general
on carrot and parsley. Affected leaves and petioles are spotted, then turn
yellow and brown; entire tops are killed in severe infections. In California
the disease is known as late blight, with the peak coming in November. The
fungus apparently winters in discarded tops and on seed.
Control. Clean up refuse. Spray with a xed copper spray or dust, starting
soon after seedlings emerge and repeating at 7- to 10-day intervals.
Alternaria dianthicola. Carnation Collar Blight, Leaf Spot, Stem and
Branch Rot, general on carnation, widespread on garden pinks and sweet
william. The chief symptom is a blight or rot at leaf bases and around nodes,
which are girdled. Spots on leaves are ashy white but centers of old spots are
covered with dark brown to black fungus growth. Leaves may be constricted
and twisted, the tip killed. Branches die back to the girdled area, and black
crusts of spores are formed on the cankers. Conidia are spread during water-
ing in the greenhouse or in rains, outdoors. Entrance is through wounds,
stomata, or directly through the cuticle. The spores are carried on cuttings.
Control. Commercial growers can often avoid Alternaria blight by keeping
plants growing continuously in the greenhouse. Cuttings should be disease-
free, taken from midway up the stem, broken at the joint rather than cut, and
started in sterilized soil. Ordinarily the foliage should be kept dry, but under
mist propagation chemicals introduced into the mist system have reduced
blight.
Alternaria helianthi. Blight and Stem Lesion of sunower.
Alternaria panax. Alternaria Blight, Root Rot, Leaf Spot of ginseng,
ming aralia, and goldenseal, generally distributed. In Ohio the disease
appears each year in semiepidemic form and has been controlled with
bordeaux mixture or a xed copper spray plus a wetting agent, starting when
plants emerge in early May and repeating every 2 weeks until 3 weeks after
bloom.
Alternaria solani. Early Blight of potato and tomato, general on these
hosts, occasional on eggplant and pepper. The pathogen was rst described
from New Jersey, in 1882.
Leaf symptoms are dark brown, circular to oval spots, marked with concen-
tric rings in a target effect, appearing rst on lower, shaded foliage, with
the spots growing together to blight large portions or all of leaves, exposing
fruits. There may be a collar rot of young tomato seedlings, sunken spots
or cankers on older stems, blossom-drop with loss of young fruits, or dark
leathery spots near the stem end of older fruits. Alternaria blight is the most
common leaf spot disease of tomatoes in the Central and Atlantic States but
is somewhat less important elsewhere.
Foliage symptoms on potato are similar to those on tomato. Small round
spots on tubers afford entrance to secondary rot organisms. Each leaf spot
may produce three or four crops of dark spores, which remain viable more
BLIGHTS 153
than a year. They are blown by wind, splashed by rain, sometimes transmitted
by ea beetles. The fungus is a weak parasite, entering through wounds and
thriving in warm, moist weather, with 85F as optimum temperature. It can
survive in soil as long as the host refuse is not completely rotted; it also
winters on seed and on weed hosts.
Control. Plan, if possible, a 3-year rotation with crops not in the potato
family; dig under diseased refuse immediately after harvest. Use seed from
healthy tomatoes, or purchase plants free from collar rot.
Alternaria tagetica. Blight of marigold.
Alternaria tenuissima. Alternaria Blight, Leaf Spot of violet and pansy.
Spots vary from greenish yellow to light buff with burnt amber margins.
Brown patches run together to form large, blighted areas. Clean up and burn
old leaves in fall.
Alternaria zinniae. Zinnia Blight, Alternariosis on zinnia. Small reddish
brown spots with grayish white centers increase to irregular, large, brown,
dry areas. Similar spots on stem internodes or at nodes may girdle the stem,
with dying back of upper portions. Dark brown to black basal cankers with
sunken lesions are common. Roots may turn dark gray, rot, and slough off.
Small brown ower spots enlarge to include whole petals, causing conspic-
uous blighting. The fungus apparently winters on seed and in soil.
Control. Clean up refuse; use a long rotation if growing plants commercially.
Ascochyta
Pycnidia dark, globose, separate, immersed in host tissue, ostiolate; spores two-celled,
hyaline ovoid to oblong.
Ascochyta asparagina. Stem Blight of asparagus fern. Small branchlets
dry and drop prematurely; small branches are killed if attacked at crown.
Ascochyta chrysanthemi (Mycosphaerella ligulicola). (see Phoma chrysan-
themi (Didymella ligulica)). Ascochyta Ray Blight of chrysanthemum,
a conspicuous and rapid disease of ray owers.
Ascochyta fabae f. sp. spiricia. Leaf Blight of vetch.
Ascochyta piniperda. Spruce Twig Blight on young shoots of red, Nor-
way, and blue spruce; apparently a minor disease.
Ascochyta pisi, A. pinodes, A. pinodella. Ascochyta Blight or Myco-
sphaerella Blight of peas. All three fungi may be connected with the dis-
ease complex known as Ascochyta blight, are carried in infected seed and
overwinter in plant debris. A. pinodes has Mycosphaerella pinodes as its
ascospore stage so that the life cycle can start from either pycnidia or perithe-
cia produced on plants or stubble.
Lesions begin as small purplish specks on leaves and pods. When infection
is caused by M. pinodes or A. pinodella, the specks enlarge to round, tar-
getlike spots, which join together to form irregular, brownish purple blotch-
es. M. pinodes often withers and distorts young pods; A. pinodella causes
a severe foot rot, a dark region at the soil line. Elongated, purplish black
stem lesions are common. A. pisi causes leaf spots with dark brown margins,
stem and pod spots, but no foot rot.
Control. Use western-grown seed, usually free from the disease; clean up all
pea refuse and use a 3- or 4-year rotation.
The host range now includes many plants such as carrot, banana, and foliage
plants.
Phoma chrysanthemi (Telemorph, Didymella ligulica) (formerly Ascochy-
ta chrysanthemi (telemorph, Mycosphaerella ligulicola)). Ascochyta Ray
Blight of chrysanthemum, a conspicuous and rapid disease of ray owers. If
young buds are infected, the head does not open; if the attack is later, there
may be one-sided development of owers. A tan or brown discoloration pro-
ceeds from the base toward the tip of each individual ower, followed by
withering. Upper portions of stems and receptacles may turn black. Keep
plants well spaced; avoid overhead watering and excessive humidity.
Balansia
Ascomycetes, Hypocreales, Clavicipitaccae.
Balansia cyperi. Diseased Inorescence, Blight of purple nutsedge;
fungus is systemic and transmitted through tubers.
Beniowskia
Hyphae are coiled at the periphery of mature sporodochia; spherical spores are borne on
short denticles.
Beniowskia sphaeroidea. Blight of knotroot bristlegrass.
BLIGHTS 155
Botryodiplodia
Pycnidia black, ostiolate, erumpent, stromatic, conuent; conidiosphores simple, short;
conidia dark and 2- celled, ovoid to elongate
Botryosphaeria
Asci in locules in a stroma; spores one-celled, hyaline, eight in an ascus. There is a good
deal of variation in the genus. The locules may be scattered throughout the stromatic
tissue, or seated on the surface, or like perithecia, as in Botryosphaeria ribis. In B. ribis
there are two pycnidial forms, a Dothiorella stage containing very small spores that may
function as male cells and a Macrophoma stage containing larger spores, one-celled,
hyaline, functioning as other conidia.
Botryosphaeria ribis var. chromogena. Current Cane Blight, Canker,
Dieback of currant, owering currant, gooseberry, apple, rose, and many
other plants (also
Cankers). There are two forms of this species, one being

a saprophyte developing on already dying tissue. The parasitic form chro-
mogena is so named from its developing a purple-pink color when grown
on starch paste. There are also a number of pathogenic strains, varying from
high to low in virulence. Some currant varieties are quite resistant, but the
widely grown Wilder and Red Lake are rather susceptible.
Dieback and death of fruiting branches occur as the berries are coloring, with
leaves wilting and fruit shriveling. Later in the season small, dark, wartlike
fruiting bodies appear in rather denite parallel rows on the diseased canes.
Rose canes show a similar dying back and wilting above a canker. The fungus
winters in the canes; ascospores infect new shoots; secondary infection is by
spores oozing from pycnidia. The mycelium grows downward through bark
and wood to the main stem, which it encircles and kills.
Control. Cut out and burn diseased canes as soon as noticed. Take cuttings
from healthy bushes.
Botryotinia
Ascomycetes, Helotiales, Sclerotiniaceae
Stroma a typical black sclerotium, loaf-shaped or hemispherical, just on or beneath cuti-
cle or epidermis of plant and rmly attached to it; apothecia cupulate, stalked, brown;
ascospores hyaline, one-celled; conidiophores and conidia of the Botrytis cinerea type.
Botryotinia fuckeliana. The apothecial stage of Botrytis cinerea, the con-
nection having been made with isolates from grape, apple, celery, and potato.
The name of the conidial stage is still widely used for the pathogen causing
gray mold blights.
Botryotinia ricini. Gray Mold Blight of castor bean, Soft Rot of caladium.
A pale to olive gray mold develops on castor-bean inorescence, and when
fading owers drop onto stem and leaves, they are infected in turn.
Botrytis
Egglike conidia hyaline, one-celled, are formed on branched conidiophores over the
surface, not in special fruiting bodies (see Fig. 3.9). The arrangement of the spores gives
the genus its name, from the Greek botrys, meaning a cluster of grapes. Flattened, loaf-
shaped, or hemispherical black sclerotia are formed on or just underneath cuticle or
epidermis of the host and are rmly attached to it. These sclerotia, with a dark rind and
light interior made up of rmly interwoven hyphae, serve as resting bodies to carry the
fungus over winter. Microconidia, very minute spores that are spermatia or male cells,
function in the formation of apothecia in the few cases where a denite connection has
been made between the Botrytis stage and the ascospore form, Botryotinia.
Botrytis species are the common gray molds, only too familiar to every gar-
dener. Some are saprophytic or weakIy parasitic on senescent plant parts on
a wide variety of hosts; others are true parasites and cause such important
diseases as peony blight, lily blight, tulip re.
Botrytis cinerea. Gray Mold Blight, Bud and Flower Blight (see
Fig. 3.10), Blossom Blight, Gray Mold Rot, Botrytis Blight of gen-
eral distribution on a great many owers, fruits and vegetables. There are
undoubtedly many strains of this fungus and perhaps more than one species
involved, but they have not been denitely separated.
This gray-mold disease is common on soft ripe fruits after picking, as any
cook knows after throwing out half a box of strawberries or raspberries.
But in continued humid weather the blight appears on fruits before harvest.
BLIGHTS 157
Figure 3.10 Botrytis Petal Spot on Magnolia
Blackberries in the Northwest are subject to gray mold. The fungus winters
in blighted blueberry twigs, and spores infect blossom clusters.
Vegetables are commonly aficted as seedlings grown in greenhouses and in
storage after harvest. If lettuce plants are set in the garden too close together,
they may blight at the base in moist weather, as will endive and escarole.
Gray mold is common on lima beans, is sometimes found on snap and kid-
ney beans. In rainy or foggy periods globe artichoke may be covered with
a brownish gray, dusty mold, with bud scales rotten. Asparagus shoots are
sometimes blighted, tomato stems rotted.
Some of the ornamentals on which Botrytis cinerea is troublesome are given
in the following annotated list:
African violet leaf and stem rot, cosmopolitan in greenhouses.
Amaryllis gray mold, mostly in the South, on outdoor plants after chilling.
Anemone occasional severe rotting of crowns.
Arborvitae twig blight.
Aster brown patches in ower heads of perennial aster; gray mold on ow-
ers of China aster grown for seed in California.
Begonia dead areas in leaves and owers rapidly enlarging and turning
black in a moist atmosphere; profuse brownish gray mold.
Calendula gray-mold blight.
Camellia ower and bud blight, common after frost.
Carnation ower rot or brown spotting, worse in a cool greenhouse.
Century plant gray mold after overwatering and chilling.
Chrysanthemum cosmopolitan on owers, buds, leaf tips, and cuttings. Ray
blight on owers starts as small, water-soaked spots, which rapidly enlarge
with characteristic gray mold.
Dahlia bud and ower blight.
Dogwood ower and leaf blight. In wet springs anthers and bracts of aging
owers are covered with gray mold, and when these rot down on top of young
leaves, there is a striking leaf blight.
Eupatorium stem blight, common in crowded plantings. A tan area girdles
stem near ground with tops wilting or drying to that point.
Geranium (Pelargonium) blossom blight and leaf spot, most common in
cool, moist greenhouses where plants are syringed frequently. Petals are dis-
colored, owers drop, gray mold forms on leaves.
Lily Botrytis cinerea is common on lilies, but see also B. elliptica.
Marigold gray mold prevalent on fading owers.
Peony late blight, distinguished from early blight (see B. paeoniae) by
the sparse mold, usually standing far out from affected tissues, rather than
a thick, short velvety mold, and by much larger, atter sclerotia formed near
base of the stalk. Late owers are infected, and when they drop down onto
wet foliage, irregular brown areas are formed in leaves.
Pine seedling blight.
Pistachio shoot blight.
Poinsettia tip blight and stem canker.
Primrose crown rot and decay of basal leaves, with prominent gray mold,
very common in greenhouses where plants are heavily watered.
Rhododendron ower, twig, and seedling blight.
Rose bud or ower blight, cane canker. When half-open buds ball, the
cause is often an infestation of thrips; but if gray mold is present, Botrytis is
indicated. Canes kept too wet by a manure mulch, or wet leaves, or injured
in some way, are often moldy.
BLIGHTS 159
Snapdragon ower spikes wilt; tan cankers girdle stems.
Sunower bud rot and mold.
Sweet pea blossom blight.
Viola spp. gray mold and basal rot of violet and pansy.
Zinnia petal blight, head blight, moldy seed.
Botrytis cinerea may also infect arabis, cineraria, eucharis, euphorbia, fuch-
sia, gerbera, gypsophila, heliotrope, hydrangea, iris, lilac, lupine, May-apple,
pyrethrum, periwinkle, rose-of-Sharon, stokesia, viburnum, and wallower.
Control. Sanitation is more important than anything else. Carry around
a paper bag as you inspect the garden; put into it all fading owers and
blighted foliage; if infection is near the base, take the whole plant up for
burning. Keep greenhouse plants widely spaced, with good ventilation;
avoid syringing, overhead watering, and too cool temperature. Propagate
cuttings from healthy plants in a sterilized medium.
Botrytis douglasii. Seedling Blight of giant sequoia and redwood, perhaps
a form of B. cinerea.
Botrytis elliptica. Lily Botrytis Blight, general on lilies, also reported on
tuberose and stephanotis in California. Lily species vary in susceptibility
to the disease, but there are several strains of the fungus, and few lilies are
resistant to all strains. Madonna lily, L. candidum, is particularly susceptible,
with infection starting in autumn on the rosette of leaves developed at that
time.
If the blight strikes early, the entire apical growth may be killed with no
further development. More often the disease starts as a leaf spot when stems
are a good height. Spots are orange to reddish brown, usually oval. In some
species there is a denite red to purple margin around a light center; in others
the dark margin is replaced by an indenite water-soaked zone. If spots are
numerous, they grow together to blight the whole leaf. Infection often starts
with the lowest leaves and works up the stem until all leaves are blackened
and hanging limp. This is the result of many spot infections and not from an
invasion of the vascular system.
Buds rot or open to distorted owers with irregular brown ecks. There are
sometimes severe stem lesions, but the rot rarely progresses into the bulbs.
Spores formed in the usual gray-mold masses in blighted portions are spread
by rain, air currents, and gardeners. Optimum spore germination is in cool
weather, around 60F, but once infection has started 70F promotes most
rapid blighting. With sufcient moisture the cycle may repeat every few days
through the season. The fungus winters as very small black sclerotia, irregu-
lar or elliptical in shape, in fallen owers or blighted dead stems and leaves,
or as mycelium in the basal rosette of Madonna lilies.
Control. Avoid too dense planting, and shady or low spots with little air
circulation and subject to heavy dews. Clean up infected plant parts before
sclerotia can be formed. Copper sprays are more effective for the lily Botrytis
than the newer organics. Spray with bordeaux mixture; start when lilies are
5 or 6 inches high and continue at 10- to 14-day intervals until owering.
Botrytis galanthina. Botrytis Blight of snowdrop, sometimes found in the
sclerotial state on imported bulbs. If the black dots of sclerotia are present
only on outer scales, remove scales before planting; otherwise discard bulbs.
Botrytis gladiolorum. Gladiolus Botrytis Blight, Corm Rot, rst report-
ed in Oregon in 1939 and now serious in all important gladiolus-growing
areas the Pacic Coast, the Midwest, Florida in cool, rainy weather. In
northern areas the disease is a corm-rotting problem, in the South a ower
blight, damaging in transit, and in all areas it is a leaf spot or blight.
In dry weather and in more resistant varieties the leaf spots are very small,
rusty brown, appearing only on the exposed side of the leaf. In more humid
weather the spots are large, brown, round to oval or smaller, pale brown with
reddish margins. Flower stems have pale brown spots that turn dark. There
may be a soft rotting at the base of orets. The disease starts on petals as pin-
point, water-soaked spots, but in moist weather the whole ower turns brown
and slimy. Flowers with no visible spotting when packed often arrive ruined.
After the owers are cut, infection spreads down the stalk and into the corm,
producing dark brown spots, irregular in shape and size, most numerous on
the upper surface. Corms may become soft and spongy with a whitish mold.
Oval, at, black sclerotia, 1/8 to 1/4 inch long, are formed on corms in stor-
age and in rotting tissue in the eld or in refuse piles. They may persist in
the soil several years.
Control. Cure corms rapidly after digging; bury or burn all plant refuse.
Botrytis hyacinthi. Hyacinth Botrytis Blight recently found in Washing-
ton on plants grown from imported bulbs. Leaves have brown tips with gray
mold or brown spots on lower surface. Leaves may be killed, with small
black sclerotia formed in rotting tissue. Flowers rot and are covered with
powdery gray spores. Do not work with plants when they are wet; remove
infected parts or whole plants.
Botrytis narcissicola.

Sclerotinia narcissicola, under Rots.
Botrytis paeoniae. Peony Botrytis Blight, Early Blight, Bud Blast,
Gray Mold, probably present wherever peonies are grown. It is also record-
BLIGHTS 161
ed on lily-of-the-valley, but that may be a form of Botrytis cinerea. Peony
blight was rst noticed in epiphytotic form in this country in 1897 and has
been important in wet springs ever since.
Young shoots may rot off at the base as they come through the ground or
when a few inches high, with a dense velvety gray mold on the rotting por-
tions. This early shoot blight is far more common when the young stems are
kept moist by having to emerge through a mulch of manure or wet leaves.
Flowers are attacked at any stage. Buds turn black when they are very tiny,
never developing, or they may be blasted when they are half open. If it is
dry in early spring, infection may be delayed until owers are in full bloom,
at which time they turn brown. Infection proceeds from the ower down the
stem for a few inches, giving it a brown and tan zoned appearance. Leaf
spots develop when infected petals fall on foliage. Continued blighting of
leaves through the summer and late blasting of owers may be due to Botrytis
cinerea, which produces a sparser mold and conidiophores projecting farther
from the petal or leaf surface.
Conidia are blown by wind, splashed by rain, carried on gardeners tools, and
sometimes transported by ants. Secondary infection is abundant in cool moist
weather. In late summer small, shiny black, slightly loaf-shaped sclerotia
are formed near the base of stalks, just under the epidermis. They are quite
different from the large, at, black sclerotia often formed by B. cinerea on
the same stalks.
Control. Sanitation is the most important step. Cut down all tops in autumn
at ground level, or just below, to get rid of sclerotia wintering near base
of stems. Burn this debris; never use it for a mulch. Avoid any moisture-
retentive covering. If you insist on manure, apply it in a wide ring around the
plant, well outside the area of emerging shoots. Go around with a paper
bag periodically, cutting off for burning all blighted parts; never carry
these parts loose through the garden for fear of shedding spores to healthy
plants.
Botrytis polyblastis.

Sclerotinia polyblastis.
Botrytis streptothrix (see Streptobotrys arisaemae) (teleomorph state
Streptotinia arisaemae). Leaf and Stalk Blight of Jack-in-the-pulpit and
golden club.
Botrytis tulipae. Tulip Fire, Botrytis Blight of tulips, general wherever
tulips are grown, causing much damage in rainy springs. The rst indication
of disease is the appearance of a few malformed leaves and shoots among
healthy tulips or large light patches resembling frost injury on leaves. Gray
Figure 3.11 Botrytis Blight on Tulip
mold forming on such blighted areas of plants grown from infected bulbs
provides an enormous number of conidia to be splashed by rain to near-
by tulips. Secondary infection appears as minute, slightly sunken, yellowish
leaf spots, surrounded with a water-soaked area, and gray to brown spots on
stems, often zonate, and resulting in collapse. Small white spots appear on
colored owers, brown spots on white petals (see Fig. 3.11); but with contin-
ued moisture the spots grow together, and in a day or so the fuzzy gray mold
has covered rotten blooms and large portions of blighted leaves.
Very small, shiny black sclerotia are formed in leaves and petals rotting into
the ground, or on old ower stems or bulbs. Sometimes the latter have yellow
to brown, slightly sunken, circular lesions on outermost eshy scales without
the formation of sclerotia. Spring infection comes from spores produced on
such bulbs or from sclerotia on bulbs or sclerotia left loose in the soil after
infected tissues have rotted.
Control. Inspect all bulbs carefully before planting; discard those harboring
sclerotia or suspicious brown lesions. It is wise, though seldom possible in
a small garden, to plant new bulbs where tulips have not grown for 3 years.
Plant where there is good air circulation. Make periodic inspections, starting
early, removing into a paper bag plants with serious primary infection and
blighted leaves. Cut off all fading owers before petals fall; cut off all foliage
at ground level when it turns yellow. Burn all debris.
BLIGHTS 163
Streptobotrys arisaemae (formerly Botrytis streptothrix) (teleomorph state
Streptotinia arisaemae). Leaf and Stalk Blight of Jack-in-the-pulpit and
golden club. This species has conidiophores with strikingly twisted branches,
producing a reddish brown mat of conidia. Sclerotia are very small, seldom
over 1/32 inch, black, shiny, and somewhat hemispherical.
Briosia
Conidia on synnemata or coremia, erect fascicles of hyphae ending in a small head;
spores globose, dark, one-celled, catenulate (formed in chains).
Briosia azaleae (see Pycnosystanus azalea) (Pycnostysanus azaleae). Bud
and Twig Blight of azalea and rhododendron, widespread but occasional.
Pycnosystanus azalea (formerly Briosia azaleae) (Pycnostysanus azaleae).
Bud and Twig Blight of azalea and rhododendron, widespread but occa-
sional. The disease was reported from New York in 1874 and, as a rhodo-
dendron bud rot, from California in 1920. It was particularly serious on Mas-
sachusetts azaleas in 1931 and 1939. Flower buds are dwarfed, turn brown
and dry; scales are silvery gray. Twigs die when lateral leaf buds are infect-
ed. Successive crops of coremia are produced on old dead buds for as long as
three years, the rst crop appearing the spring after summer infection. The
coremia heads are dark, and the buds look as if stuck with tiny, round-headed
pins. Prune out and burn infected buds and twigs in late autumn and early
spring. Spraying with bordeaux mixture before blossoming and at monthly
intervals after bloom may be wise in severe cases.
Calonectria
Cylindrocladium under Blights.

Calonectria colhounii. Blight on Leucospermum.
Cenangium
Apothecia small, brown to black, sessile or substipitate on bark; spores hyaline, ellipti-
cal, one-celled; paraphyses liform.
Cenangiumferruginosum. Pine Twig Blight, Pruning Disease; Cenan-
gium Dieback of r and pine. The fungus is ordinarily saprophytic on
native pines but may become parasitic when their vigor is reduced by
drought. The disease is considered benecial to ponderosa pine in the South-
west because it prunes off the lower branches; on exotic pines it can be
damaging.
Infection starts near a terminal bud in late summer and progresses down
a twig into a node, sometimes beyond into 2-year wood. The needles redden
and die; they are conspicuous in spring but drop in late summer. Then brown
to black apothecia with a greenish surface to the cup appear on twigs. Cut
off and destroy infected twigs.
Cercospora
Conidia hyaline to pale to medium green or brown; long, usually with more than three
cross walls; straight or curved, with the base obconate or truncate, tip acute to obtuse;
thin-walled; not formed in a fruiting body but successively on slender conidiophores,
which emerge in fascicles or groups from stomata and usually show joints or scars where
conidia have fallen off successively. The conidiophores are always colored, olivaceous
to brown, pale to very dark (Fig. 3.9).
This is the largest group of the Dematiaceae, with about 400 species, all
parasitic, causing leaf spots or blights. The teleomorph state, when known,
is Mycosphaerella.
Cercospora apii. Early Blight of celery, general on celery and celeriac, rst
noted in Missouri in 1884 and since found in varying abundance wherever
celery is grown. The disease is most severe from New Jersey southward. The
name is somewhat misleading; in Florida early blight rarely appears before
the Septoria disease known as late blight. Foliage spots appear when plants
are about 6 weeks old. Minute yellow areas change to large, irregular, ash
gray lesions, covered in moist weather with velvety groups of conidiophores
and spores on both sides of leaves. Sunken, tan, elongated spots appear on
stalks just before harvest. The disease spreads rapidly in warm, moist weath-
er, the spores being splashed by rain, carried with manure or cultivators, or
blown by wind. The life cycle is completed in 2 weeks.
Control. Seed more than 2 years old is probably free from viable spores;
other seed should be treated with hot water, 30 minutes at 118 to 120F.
BLIGHTS 165
Bordeaux mixture and other copper sprays have been recommended. Spray
applications should start soon after plants are set and be repeated weekly, or
more often. Emerson Pascal is blight-resistant.
Cercospora carotae. Early Blight of carrot. Lesions on leaves and stems
are subcircular to elliptic, pale tan to gray or brown or almost black; lobes
or entire leaets are killed. The disease is more severe on young leaves and
builds up as the plant grows. Spores, produced on both leaf surfaces, are
spread by wind.
Control. Rotate crops and clean up refuse.
Cercospora microsora. Linden Leaf Blight, general on American and
European linden. Small circular brown spots with darker borders coalesce
to form large, blighted areas, often followed by defoliation; most serious on
young trees.
Cercospora sequoiae. Arborvitae Blight, Fire Blight, on oriental arborvi-
tae and Italian cypress in the South; destructive in ornamental plants. First
reported from Louisiana in 1943, the fungus was named as a new species
of Cercospora in 1945, but it is nearer Heterosporium in spore character.
Affected leaves and branchlets are killed, turn brown, and gradually fall off,
leaving shrubs thin and ragged. The lower two-thirds of the bush is affected
most severely, with a tuft of healthy growth at the top. When close to a house,
the side away from the wall shows most symptoms. Plants crowded in nurs-
eries are killed in 1 to 3 years, but in home gardens they may persist for years
in an unsightly condition. Conidiophores in fascicles produce conidia after
girdling cankers have killed the twigs. There is often a swelling above the
girdle that resembles an insect gall.
Cercospora sordida (Mycosphaerella tecomae) (see Pseudocercospora sor-
dida) Trumpetvine Leaf Blight from New Jersey to Iowa and southward.
Pseudocercospora sordida (formerly Cercospora sordida) (Mycosphaerel-
la tecomae). Trumpetvine Leaf Blight from New Jersey to Iowa and south-
ward. Small, angular, sordid brown patches run together; edge of leaets may
be purplish; the fungus fruits on underside of leaves. The blight is seldom
important enough to warrant control measures.
Cercosporidium
Cercosporidium punctum. Stem and Foliage Blight of fennel.
Choanephora
Zygomycetes, Mucorales
Mycelium profuse; sporangia and conidia present; sporangiola lacking. Sporangium
pendent on recurved end of an erect, unbranched sporangiophore with a columella, con-
taining spores provided at both ends and sometimes at the side with a cluster of ne,
radiating appendages. Conidia formed in heads on a few short branches or an erect coni-
diophore enlarged at the tip; conidia longitudinally striate, without appendanges.
Choanephora cucurbitarum. Blossom Blight, Fruit Rot, common on
summer squash and pumpkin, occasional on amaranth, cowpea, cucumber,
okra, and pepper; on sweetpotato foliage, on fading hibiscus, on vinca, and
other owers. This blight is often found in home gardens in seasons of high
humidity and rainfall. Flowers and young fruits are covered with a luxuriant
fungus growth, rst white, then brown to purple with a denite metallic lus-
ter. The fruiting bodies look like little pins stuck through this growth. Both
staminate and pistillate owers are infected, and from the latter the fungus
advances into young fruits, producing a soft wet rot at the blossom end. In
severe cases all owers are blighted or fruits rotted.
Control. Grow plants on well-drained land; rotate crops. Remove infected
owers and fruits as noticed.
Choanephora infundibulifera. Blossom Blight on hibiscus and jasmine.
Leaf Blight on soybean.
Ciboria
Stroma a dark brown to black sclerotium in catkins or seed, simulating in shape the
stromatized organ and not resembling a sclerotium externally. Apothecia cupulate to
shallow saucer-shaped; brown.
Ciboria acerina. Maple Inorescence Blight on red and silver maple.
Apothecia, developed in great numbers from stromatized inorescences on
ground beneath trees, start discharging spores when maple owers appear
overhead. Mycelium spreads through stamens, calyx, and bud scales until
ower cluster drops.
Ciboria carunculoides. Popcorn Disease of mulberry, a southern disease,
not very important. Sclerotia are formed in carpels of fruit, which swells to
resemble popcorn but remains green.
BLIGHTS 167
Ciborinia
Stroma a thin, at, black sclerotium of discoid type in leaves; one to several stalked
apothecia arise from sclerotia; apothecia small, brown, cupulate to at when expanded.
Ciborinia erythronii and C. gracilis. Leaf Blight of erythronium. Flat
black sclerotia are prominent in leaves.
Cladosporium
Blotch.
Cladosporium cladosporioides. Blossom Blight on strawberry.
Colletotrichum
Anthracnose.
Colletotrichum acutatum. Twig Blight and Fruit Spot on dogwood.
Colletotrichum gloeosporioides. Seedling Blight of papaya.
Colletotrichum dematium. Twig Blight on vinca.
Corticium
Basidiomycetes, Aphyllophorales
Hymenium or fruiting surface of basidia consisting of a single resupinate or horizontal
layer. This genus has contained a rather heterogeneous collection of species; some of the
more important have been transferred to the genus Pellicularia.
Corticium koleroga. Thread Blight.
Pellicularia koleroga.
Corticium microsclerotia. Web Blight.
Pellicularia lamentosa.
Corticium salmonicolor (see Erythricium salmonicolor). Limb Blight of
g, pear, apple in Gulf States.
Corticium stevensii. Thread Blight.
Pellicularia koleroga.
Corticium vagum, now Pellicularia lamentosa, teleomorph state of Rhi-
zoctonia solani, causing black scurf of potatoes and damping-off and root
rot of many plants. See both Pellicularia and Rhizoctonia under Rots.
Erythricium salmonicolor (formerly Corticium salmonicolor). Limb
Blight of g, pear, apple in Gulf States. The spore surface is pinkish.
Coryneum
Acervuli subcutaneous or subcortical, black, cushion-shaped or disc-shaped; conidio-
phores slender, simple; spores dark with several cross walls, oblong to fusoid; parasitic
or saprophytic (see Fig. 3.15).
Coryneum berckmansii (see Seimatosporium berckmansii). Coryneum
Blight of Oriental arborvitae, also on Italian cypress, causing serious losses
in nurseries and home gardens in the Pacic Northwest.
Coryneum microstictum (see Seimatosperium lichehnicola). Twig Blight
of American bladdernut.
Coryneum carpophilum (Cladosporium beijerinckii) (see Stigmina car-
pophila, Anamorph, Cladosporium beijerinckii). Peach Shoot Blight,
Coryneum Blight of stone fruits, Shot Hole, Fruit Spot, Winter Blight,
Pustular Spot, general on peach in the West, also on almond, apricot,
nectarine, and cherry.
Seimatosporiumberckmansii (formerly Coryneum berckmansii). Coryneum
Blight of Oriental arborvitae, also on Italian cypress, causing serious loss-
es in nurseries and home gardens in the Pacic Northwest. Small twigs
or branches are blighted, turn gray-green then reddish brown; many small
branchlets drop, leaving a tangle of dead gray stems; larger limbs may be
girdled. Twigs are dotted with black pustules bearing ve-septate spores.
As new growth develops in blighted areas, the spores spread the disease
to young contiguous foliage. Reinfection continues until the plant is so
devitalized it dies. The fungus fruits only on scale leaves or young stems.
Control. Remove and destroy blighted twigs. Apply a copper spray in
September to healthy bushes as a preventive spray; apply in September
and repeat in late October to infected bushes.
Seimatosporium lichenicola (formerly Coryneum microstictum). Twig
Blight of American bladdernut. Young twigs are killed; the fungus winters
in acervuli on this dead tissue, and spores are disseminated in spring. Prune
out and burn diseased twigs during the winter.
BLIGHTS 169
Stigmina carpophila (formerly Coryneum carpophilum (Cladosporium bei-
jerinckii)). Peach Shoot Blight, Coryneum Blight of stone fruits, Shot
Hole, Fruit Spot, Winter Blight, Pustular Spot, general on peach in the
West, also on almond, apricot, nectarine, and cherry. Twig lesions are formed
on 1 -year shoots, reddish spots developing into sunken cankers; fruit buds
are invaded, and there is copious gum formation. Small spots are formed on
foliage, dropping out to leave typical shot holes, followed by considerable
defoliation.
Apricot buds are blackened and killed during winter; fruiting wood in peach-
es is killed before growth starts. In late rains leaves and fruit are peppered
with small, round, dead spots. Fruit lesions are raised, roughened, scabby.
The fungus winters in twigs, diseased buds and spurs.
Control. In California, the standard spray for peach is bordeaux mixture
applied in autumn immediately after leaf fall and before the rainy season. On
apricots additional sprays are suggested for late January and at early bloom.
On almonds at least two spring sprays are recommended, one at the popcorn
stage of bloom, the other at petal fall.
Cryptocline
Cryptocline cinerescens. Twig Blight of oaks.
Cryptospora
Scomycetes, Amphisphaeriales
Perithecia immersed in a stroma, with long necks converging into a disc; ascospores
long, liform, hyaline; conidia on a stroma.
Cryptospora longispora (see Servazziella longispora). Araucaria Branch
Blight.
Servazziella longispora (formerly Cryptospora longispora). Araucaria
Branch Blight. Lower branches are attacked rst, with disease spreading
upward; tip ends are bent and then broken off; plants several years old may
be killed. Prune off and burn infected branches.
Cryptostictis
Deutermycetes, Coelomycetes
Spores dark, with several cross walls, formed in acervuli.
Cryptostictis sp. Twig Blight of dogwood.
Curvularia
Conidiophores brown, simple or sometimes branched, bearing conidia successively on
new growing tips; conidia dark, three- to ve-celled, with end cells lighter, more or less
fusiform, typically bent or curved with central cells enlarged; parasitic or saprophytic.
Curvularia cymbopogonis. Blight and Leaf Spot of itchgrass. Leafspots
coalesce after 3 or 4 days to form larger lesions and nal blighting symptoms.
Curvularia lunata (C. trifolii f. sp. gladioli). Gladiolus Flower Blight
and Leaf Spot, Curvularia Disease. Suddenly, in 1947, a blight showed
up in Florida as a serious threat to the gladiolus cut-ower industry, ruin-
ing hundreds of acres there and in Alabama in the next few months. The
disease is now recorded as far north as New York and Wisconsin and on
the Pacic Coast. The pathogen is usually identied as Curvularia luna-
ta, known as a crop pest for many years, especially in the tropics, but
studies indicate it is a special form of C. trifolii, cause of a leaf spot of
clover.
Curvularia spots on leaf or stem are oval, tan to dark brown, showing on
both sides of the leaf, bordered with a brown ring, slightly depressed and
with a narrow yellowish region between the spot and normal green of the
leaf. Tan centers of spots are covered with black spores resembling powder.
Premature death comes when stems of young plants are girdled; orets fail
to open when petioles are girdled.
Under favorable weather conditions tan spots on petals turn into a smudgy
ower blight. Brown to black irregular lesions appear on corms of blooming
stock and develop further in storage; the fungus survives in corms from one
season to the next. This is a high temperature fungus, with optimum for
growth 75 to 85F and no infection under 55F. A 13-hour dew period is
sufcient moisture. Leaf spots show up in 4 to 5 days, spots on orets and
stems in only 2 to 3 days. The complete life cycle is as short as a week in
BLIGHTS 171
warm rainy weather, and the fungus can survive in the soil for 3 years. Many
gladiolus varieties are more or less resistant; Picardy and some others are
very susceptible.
Cylindrocladium
Conidiophores dichotomously branched; spores hyaline, two- or several-celled.
Cylindrocladium clavatum. Blight on bird-of-paradise.
Cylindrocladium scoparium. Cylindrocladium Blight. Damping-off of
seedlings and cuttings conifers, azalea, magnolia, hydrangea, holly, pyra-
cantha, bottle brush, and poinsettia in greenhouses under very moist con-
ditions. Infected azalea leaves turn black, with petiole bases softened, and
drop in a few days; the bark turns brown. Leaves and stems are covered with
brownish mycelial strands and white powdery masses of conidia. Control by
proper humidity and aeration.
Cylindrocladium avesiculatum. Blight and Leaf Spot of Leucotho axil-
laris.
Cylindrosporium
Acervuli subepidermal, white or pale, discoid or spread out; conidiophores short, simple;
conidia hyaline, liform, straight or curved, one-celled or becoming septate; parasitic on
leaves. Many species have Higginsia or Coccomyces as a teleomorph state.
Cylindrosporium defoliatum. Leaf Blight of Hackberry. May cause defo-
liation but usually unimportant.
Cylindrosporium griseum. On western soapberry.
Cylindrosporium juglandis. On walnut.
Delphinella
Delphinella balsameae. Tip Blight of r.
Dendrophoma
Pycnidia dark or light brown, superfcial or submerged and erumpent; globose or elon-
gate, ostiolate; conidiophores elongated, branched; conidia hyaline, one-celled, elongate
to ellipsoid; parasitic or saprophytic.
Dendrophoma obscurans (see Phomopsis obscurans). Strawberry Leaf
Blight, Angular Leaf Spot. The lesions are large, circular to angular, red-
dish purple, zonate with age, having a dark brown center, a light brown zone,
and a purple border. Spots may extend in a V-shaped area from a large vein
to edge of the leaf, with black fruiting bodies appearing in the central por-
tion. Not serious before midsummer, the disease may be destructive late in
the season. The fungus winters on old leaves.
Phomopsis obscurans (formerly Dendrophoma obscurans). Strawberry
Leaf Blight, Angular Leaf Spot.
Diaporthe
Perithecia in a hard black stroma made up of host and fungal elements, rst immersed,
then erumpent; ascospores fusoid or ellipsoid, two-celled, hyaline. Anamorph state
a Phomopsis with two types of spores; alpha conidia, hyaline, one-celled ovate to
fusoid, and beta conidia, curved or bent stylospores.
Diaporthe arctii. Diaporthe Blight of Larkspur, Stem Canker, on annu-
al larkspur and delphinium. Lower leaves turn brown and dry but remain
attached; brown lesions at base of stems extend several inches upward and
down into roots. Scattered dark pycnidia are present in stems, petioles, leaf
blades, and seed capsules, the latter probably spreading the blight. Crowns
are sometimes developed in a cottony weft of mycelium; perithecia devel-
op on decaying stems. Remove and destroy diseased plants; use seed from
healthy plants.
Diaporthe phaseolorum. Lima Bean Pod Blight, Leaf Spot, apparent-
ly native in New Jersey, where it was rst noticed in 1891, more abundant
on pole than on bush beans. Leaf spots are large, irregular, brown, often
with discolored borders and large black pycnidia formed in concentric cir-
cles in dead tissue. Necrotic portions may drop out, making leaves ragged.
BLIGHTS 173
Pod lesions spread; pods turn black and wilted, with prominent black pyc-
nidia. Seeds are shriveled or lacking. Spores are produced in great numbers,
are disseminated by wind and pickers, and enter through stomata or wounds.
The disease is most severe along the coast; optimum temperature is around
80F. The fungus is seed-borne, but most lima bean seed is produced where
the disease does not occur. Use healthy seed; clean up refuse; rotate crops.
Diaporthe phaseolorum var. sojae. Soybean Pod and Stem Blight,
widespread. This disease was formerly confused with the more acute stem
canker caused by D. phaseolorum var. caulivora (
Rots). The pod blight

is a slower disease, killing plants in later stages of development. It can be
identied by the numerous small black pycnidia scattered over the pods and
arranged in rows on stems. The blight is more serious in wet seasons. The
fungus winters on the seed and on diseased stems in the eld. Use clean
seed; clean up plant refuse; rotate crops.
Diaporthe phaseolorum var. caulivora. Stem Blight of soybean; also
causes pod and seedling blight, stem canker, and seed decay of soybean.
Diaporthe vaccinii. Blueberry Twig Blight. The same fungus that causes
cranberry rot blights new shoots of cultivated blueberries, entering at tips,
progressing toward the base, and ultimately girdling old branches. Pycnidia
develop on leaves and dead twigs. The disease is seldom serious enough for
control measures.
Diaporthe vexans. Phomopsis Blight of Eggplant, Fruit Rot, general in
eld and market, especially in the South. Destruction is often complete, with
every above-ground part affected. Seedlings rot at ground level. The rst leaf
spots are near the ground, denite, circular, gray to brown areas with light
centers and numerous black pycnidia. The leaves turn yellow and die. Stem
cankers are constrictions or light gray lesions. Fruit lesions are pale brown,
sunken, marked by many black pycnidia arranged more or less concentrical-
ly. Eventually the whole fruit is involved in a soft rot or shriveling. Spores
winter on seed and in contaminated soil. There is no fungicidal control. Use
resistant varieties Florida Market and Florida Beauty.
Dichotomophora
Cankers.
Dichotomophora lutea. Stem Blight, of common parsley.
Didymascella (Keithia)
Ascomycetes, Phacidiales, Stictidiaceae
Apothecia brown, erumpent on leaves of conifers; spores dark, two-celled, ovoid; para-
physes liform; asci two- to four-spored.
Didymascella thujina. Arborvitae Leaf Blight, Seedling Blight of
arborvitae in eastern states and of giant arborvitae, sometimes called western
red cedar. The fungus is a native of North America and occurs abundantly
in the West, damaging seedlings and saplings, often killing trees up to 4
years old, if they are in dense stands in humid regions. Older trees do not
die, but foliage appears scorched, particularly on lower branches, and young
leaf twigs may drop. Cushionlike, olive brown apothecia embedded in leaf
tissue, usually upper, are exposed by rupture of the epidermis. After summer
discharge of spores (round, brown, unequally two-celled) the apothecia drop
out of the needles, leaving deep pits.
Control. Spray small trees and nursery stock several times during summer
and fall with bordeaux mixture.
Didymascella tsugae (see Fobrella tsugae). Hemlock Needle Blight. Nee-
dles of Canada hemlock turn brown and drop in late summer. Spores are
matured in apothecia on fallen needles with new infection in spring. The
damage is not heavy.
Fobrella tsugae (see Didymascella tsugae). Hemlock Needle Blight. Nee-
dles of Canada hemlock turn brown and drop in late summer.
Didymella
Ascomycetes, Sphaeriales, Mycosphaerellaceae
Perithecia (or perithecia-like stromata) membranous, not carbonaceous; innate; not
beaked; paraphyses present; spores two-celled, hyaline.
Didymella applanata. Raspberry Spur Blight, Purple Cane Spot,
Gray Bark, general on raspberries, also on dewberry, blackberry. Named
because it partially or completely destroys spurs or laterals on canes. The
disease, known in North America since 1891, may cause losses up to 75% of
the crop of individual plants of red raspberries. Dark reddish or purple spots
on canes at point of attachment of leaves enlarge to surround leaf and bud
and may darken lower portion of cane. Affected areas turn brown, then gray.
BLIGHTS 175
If buds are not killed outright during the winter, they are so weakened that
the next seasons spurs are weak, chlorotic, seldom blossoming. Pycnidia of
the anamorph Phoma state and perithecia are numerous on the gray bark;
ascospores are discharged during spring and early summer; on germination
they can penetrate unwounded tissue.
Control. Keep plants well-spaced, allowing plenty of sunlight for quick dry-
ing of foliage and canes. Remove infected canes and old fruiting canes after
harvest. A delayed dormant spray of lime sulfur or Elgetol may be advisable,
followed by two sprays of ferbam or bordeaux mixture, applied when new
shoots are 6 to 10 inches high and 2 weeks later.
Didymella bryoniae. Gummy stem blight and fruit spot; of watermelon.
Didymosphaeria
Perithecia innate or nally erumpent; not beaked; smooth; paraphyses present; spores
dark, two-celled.
Didymosphaeria populina (Venturia populina, V. tremulae, V. macularis
also cause this disease). Shoot Blight of polar, Leaf and Twig Blight.
Young shoots are blackened and wilted. In moist weather dark olive green
masses of spores are formed on leaves.
Diplodia
Pycnidia innate or nally erumpent; black, single, globose, smooth; ostiole present; coni-
diophores slender, simple; conidia dark, two-celled, ellipsoid or ovoid. Parasitic or sapro-
phytic.
Some species cause twig blights which are not too important: Diplodia
coluteae on bladder senna; D. longispora on white oak; Sphaeropsis
sapinea (formerly D. pinea) on pine; D. sarmentorum on pyracantha.
Lasiodiplodia theobromae (formerly Diplodia natalensis) (anamorph state
of Physalospora rhodina) causes blight, stem gumming, or stem-end rot of
melons, as well as twig blight of peach and citrus. See further under Rots.
Diplodia gossypina (see Lasiodiplodia theobromae). Blight of slash pine
and loblolly pine seedlings.
Lasiodiplodia theobromae (formerly Diplodia gossypina). Blight of slash
pine and loblolly pine seedlings
Discula
Discula quercina, Twig Blight of oaks.
Dothistroma
Stroma dark, elongate, innate, becoming erumpent and swollen, with a stalk extending
into the substratum, composed internally of dense, vertical hyphae; locules separate,
one to several in the upper part of the stroma; conidiophores simple, slender; conidia
several-celled, hyaline, long-cylindrical to liform.
Dothistroma pini. Needle Blight on Austrian pine and red pine.
Cryphonectria (Endothia)
Perithecia deeply embedded in a reddish to yellow stroma, with long necks opening
to the surface but not beaked; paraphyses lacking; spores two-celled, hyaline. Conidia
borne in hollow chambers or pycnidia in a stroma and expelled in cirrhi.
Cryphonectria parasitica. Chestnut Blight, Endothia Canker, general
on chestnut. To most gardeners this disease is of only historical importance,
for practically all of our native chestnuts are gone. The disease, however,
persists in sprouts starting from old stumps and in the chinquapin. One of the
most destructive tree diseases ever known, chestnut blight at least served to
awaken people to the importance of plant disease and to the need for research
in this eld.
First noticed in the New York Zoological Park in 1904, the blight rapidly
wiped out the chestnut stands in New England and along the Allegheny and
Blue Ridge Mountains, leaving not a single undamaged tree. In 1925 the
disease eliminated chestnuts in Illinois and by 1929 had reached the Pacic
Northwest.
BLIGHTS 177
Conspicuous reddish bark cankers are formed on trunk and limbs, often
swollen and splitting longitudinally. As the limbs are girdled, the foliage
blights, so that brown, dried leaves are seen from a distance. The fungus
fruits abundantly in crevices of broken bark, rst producing conidia extruded
in yellow tendrils from reddish pycnidia and later ascospores from perithe-
cia embedded in orange stromata. Fans of buff-colored mycelium are found
under affected bark.
Ascospores can be spread many miles by the wind, landing in open wounds,
but the sticky conidia are carried by birds and insects. The fungus can live
indenitely as a saprophyte, and new sprouts developing from old stumps
may grow for several years before they are killed.
Control. All eradication and protective measures have proved futile. Hope for
the future lies in cross-breeding resistant Asiatic species with the American
chestnut (and there has been some success in this line) or in substituting
Chinese and Japanese chestnuts for our own.
Diplocarpon (Fabraea)
Ascomycetes, Helotiales, Dermateaceae
Apothecia develop on fallen leaves; small, disclike, leathery when dry, gelatinous when
wet; asci extend above the surface of the disc; ascospores two-celled, hyaline. The
anamorph stage an Entomosporium with distinctive cruciate four-celled conidia, each
cell with an appendage, formed in acervuli (Fig. 3.9).
Didymascella tsugae (see Fabrella tsugae). Hemlock Needle Blight. Nee-
dles of Canada hemlock turn brown and drop in late summer.
Diplocarpon mespili (Entomosporium mespili) (formerly Fabraea mac-
ulate, Entomosporium maculatum). Pear Leaf Blight, Entomospo-
rium Leaf Spot, Fruit Spot, generally distributed on pear and quince,
widespread on amelanchier, sometimes found on apple, Japanese quince,
medler, mountain-ash, Siberian crab, cotoneaster, loquat, photinia.
Pears may be affected as seedlings in nurseries or in bearing orchards. Very
small purple spots appear on leaves, later extending to a brownish circular
lesion, 1/4 inch or less in diameter, with the raised black dot of a fruiting
body in the center of each spot. If spots are numerous, there is extensive
defoliation. Fruit spots are red at rst, then black and slightly sunken; the
skin is roughened, sometimes cracked. Quince has similar symptoms.
Twig lesions appear on the current seasons growth about midsummer, indef-
inite purple or black areas coalescing to form a canker. Primary spring infec-
tion comes more from conidia produced in these twig lesions than from
ascospores shot from fallen leaves on the ground. Most commercial varieties
of pear and quince are susceptible, although some are moderately resistant.
Fabraea maculate, Entomosporium maculatum (
Diplocarpon mespili,
Entomosporium mespili). Pear Leaf Blight, Entomosporium Leaf Spot,
Fruit Spot, generally distributed on pear and quince, widespread on ame-
lanchier, sometimes found on apple, Japanese quince, medler, mountain-ash,
Siberian crab, cotoneaster, loquat, photinia.
Fabraea thuemenii (Entomosporium thuemenii). Hawthorn Leaf Blight,
wide-spread on Crataegus species. Symptoms are similar to those of pear
leaf blight and for a long time the pathogen was considered identical. Small
dark brown or reddish brown spots, with raised black dots, are numerous
over leaves, which drop prematurely in August. In wet seasons trees may be
naked by late August.
Control. Because the fungus winters in twig cankers as well as in fallen
leaves, sanitation has little effect. Standard recommendation has been to
spray three times with bordeaux mixture, starting when leaves are half out
and repeating at 2-week intervals. The copper may be somewhat phytotoxic,
causing small reddish spots similar to those of blight, but it does prevent
defoliation.
Fabrella tsugae (formerly Didymascella tsugae). Hemlock Needle Blight.
Needles of Canada hemlock turn brown and drop in late summer. Spores are
matured in apothecia on fallen needles with new infection in spring. The
damage is not heavy.
Furcaspora
Starlike botryoblastospores; acervuli become erumpent at maturity and grade into
sporodochia and pycnidia.
Furcaspora pinicola. Needle Cast of pine.
BLIGHTS 179
Fusarium
Rots.
Fusarium graminearum. Head Blight of wild rice.
Fusarium moniliforme var. subglutinans. Blight of slash pine and loblolly
pine seedlings.
Fusarium solani. Stem and Leaf Blight on Spanish moss.
Fusarium subglutinans. Foliar Blight and Collar Rot on Chinese ever-
green. Seedling Blight on pine.
Fusarium tabacinum. Stem Blight of squash and pumpkin.
Gibberella
Ascomycetes, Hypocreales, Nectriaceae
Perithecia supercial, blue, violet, or greenish; spores hyaline with several cells. Coni-
dial stage in genus Fusarium with fusoid curved spores, several-septate. The species
causing stalk rots of corn and producing gibberellic acid are more important than those
causing blights.
Gibberella baccata (Fusarium lateritium). Twig Blight of ailanthus, cit-
rus, cotoneaster, g, hibiscus, hornbeam, peach, and other plants in warm
climates, sometimes associated with other diseases.
Glomerella
Anthracnose.
Glomerella cingulata. Cyclamen Leaf and Bud Blight Leaf and Shoot
Blight of poplar.
Gnomonia
Anthracnose.
Gnomonia rubi. Cane Blight of blackberry, dewberry, raspberry.
Hadrotrichum
Sporodochia cushion-shaped, dark; conidiophores dark, simple, forming a palisade and
arising from a stroma-like layer; conidia dark, nearly spherical, one-celled, borne singly;
parasitic on leaves.
Hadrotrichum globiferum. Leaf Blight of lupine.
Helminthosporium
Mycelium light to dark; conidiophores short or long; septate, simple or branched, often
protruding from stomata of host; more or less irregular or bent, bearing conidia succes-
sively on new growing tips; conidia dark typically with more than three cells, cylindrical
or ellipsoid, sometimes slightly curved or bent, ends rounded. Parasitic, often causing
leaf spots or blights of cereals and grasses.
Helminthosporium catenarium (Drechslera catenaria). Leaf Blight or
Crown Rot on creeping bentgrass; red leaf lesions and leaf tip dieback;
eventually entire plant becomes blighted to crown.
Helminthosporium gigantea (Drechslera gigantea). Blight or Zonate
Leaf Spot on wild rice and grasses.
Helminthosporium maydis (Cochliobolus heterostrophus). Southern
Corn Leaf Blight, easily confused with southern corn leaf spot due to
H. carbonum. The leaf blight occurs throughout the corn areas of the South
and north to Illinois, more important on eld than on sweet corn. Grayish
tan to straw-colored spots with parallel sides unite to blight most of the leaf
tissue. The elds appear burned by re. Resistant varieties offer the only
control.
Helminthosporium turcicum (see Setosphaeria turcica). Northern Corn
Leaf Blight on eld and sweet corn and on grasses; found from Wis-
consin and Minnesota to Florida but more severe in states with heavy
dews, abundant rainfall, and warm summers, losses running from a trace
to 50%.
Setosphaeria turcica (formerly Helminthosporium turcicum). Northern
Corn Leaf Blight on eld and sweet corn and on grasses; found from Wis-
consin and Minnesota to Florida but more severe in states with heavy dews,
BLIGHTS 181
abundant rainfall, and warm summers, losses running from a trace to 50%.
The disease starts on the lower leaves and progresses upward. Small, ellip-
tical, dark grayish green, water-soaked spots turn greenish tan and enlarge
to spindle-shape, 1/2 to 2 inches wide, 2 to 6 inches long. Spores develop-
ing on both leaf surfaces after rain or heavy dew give a velvety dark green
appearance to the center of the lesions. Whole leaves may be killed; entire
elds turn dry. The fungus winters in corn residue in the eld and produces
spores the next spring; these are spread by wind.
Control. Use a 3-year or longer rotation.
Herpotrichia
Ascomycetes, Dothidiales
Mycelium dark, perithecia supercial; spores with several crosswalls, olivaceous when
mature.
Herpotrichia juniperi. Brown Felt Blight of conifers at high eleva-
tions; on r, juniper, incense cedar, spruce, pine, yew when under snow.
When the snow melts, lower branches are seen covered with a dense felty
growth of brown to nearly black mycelium, which kills foliage by exclud-
ing light and air as well as by invading hyphae. Small, black perithecia
are scattered over the felt. This pathogen also found on dwarf mistle-
toe.
Heterosporium
Conidiophores dark, simple, bearing conidia successively on new growing tips; conidia
dark, with three or more cells, cylindrical, with rough walls (echinulate to verrucose);
parasitic, causing leaf spots, or saprophytic.
Heterosporium syringae. Lilac Leaf Blight. A velvety, olive green bloom
of spores if formed in blighted, gray-brown leaf areas, which may crack and
fall away. Infection is on mature leaves and the fungus is often associated
with Cladosporium. If necessary, spray after mid-June with bordeaux mix-
ture.
Higginisia
See Coccomyces under Leaf Spots.
Higginisia hiemalis. Cherry Leaf Blight. See Blumeriella jaapi and Coc-
comyces biemalisi, Cherry Leaf Spot.
Higginisia kerriae. Kerria Leaf and Twig Blight. See Blumeriella kerriae
and Coccomyces kerriae under Leaf Spots.
Hypoderma
Ascomycetes, Rhytismatales
Ascospores formed in hysterothecia (elongated perithecia or apothecia) extending along
evergreen needles; asci long-stalked; ascospores one-celled hyaline, fusiform, surround-
ed by a gelantinous sheath (see Fig. 3.29).
Hypoderma lethale (see Ploidoderma lethale). Gray Leaf Blight of hard
pines, from New England to the Gulf States.
Ploioderma lethale (formerly Hypoderma lethale). Gray Leaf Blight of
hard pines, from New England to the Gulf States. Hysterothecia are short,
narrow, black; often seen on pitch pine.
Hypodermella
Like Hypoderma but one-celled spores are club-shaped at upper end, tapering toward
base (see Fig. 3.29).
Hypodermella abietis-concoloris (see Lirula abietis-concoloris). Fir Nee-
dle Blight on rs and southern balsam.
Hypodermella laricis. Larch Needle and Shoot Blight. Yellow spots
are formed on needles, which turn reddish brown but stay attached, giving
a scorched appearance to trees. Hysterothecia are very small, oblong to ellip-
tical, dull black, on upper surface of needles.
Lirula abietis-concoloris (formerly Hypodermella abietis-concoloris). Fir
Needle Blight on rs and southern balsam.
BLIGHTS 183
Nectria (Hypomyces)
Ascomycetes, Hypocreales
Perithecia bright colored with a subicle (crustlike mycelial growth underneath); spores
two-celled, light, with a short projection at one end.
Hypomyces ipomoeae (see Nectria ipomoeae). Twig Blight of bladdernut.
Nectria ipomoeae (formerly Hypomyces ipomoeae). Twig Blight of blad-
dernut.
Hyponectria
Perithecia bright colored, soft; innate or nally erumpent; paraphyses lacking; spores
one-celled, light-colored, oblong.
Hyponectria buxi. Leaf Blight, Leaf Cast of boxwood.
Itersonilia
Cells reproduciug by budding and germinating by repetition; clamp connections as in
Basidiomycetes and probably anamorph species of Tremellales. The genus is not well
understood.
Itersonilia perplexans. Petal Blight of chrysanthemum and China aster.
The fungus was isolated from greenhouse chrysanthemums in Minnesota
in 1951 but apparently has been present, as a parasite or saprophyte, on
many other plants. On pompom chrysanthemums the tip half of outer petals
turns brown and dries; the diseased tissue is lled with broad hyphae and
clamp connections. Inoculated snapdragons show similar symptoms. Ade-
quate greenhouse ventilation seems to prevent trouble. This fungus has also
been reported on dill.
Itersonilia sp. Leaf Blight, Canker of parsnip, seasonal in New York and
neighboring states. Plants are defoliated in cool, moist weather. Spores from
leaves produce a chocolate brown dry rot on shoulder or crown of the root.
Good drainage and long rotation aid in control.
Kellermannia
Pycnidia black, globose, separate; immersed in host tissue; ostiolate; conidiophores
short, simple; conidia hyaline, mostly two-celled, cylindrical with an awl-shaped
appendage at the tip; parasitic or saprophytic.
Kellermania anomala (K. yuccaegena.) Yucca Leaf Blight, general on
nonarborescent forms of yucca; in Florida and California on arborescent
forms.
Kellermania sisyrinchii (see Scolecosporiella sisyrinchii.) Leaf Blight of
blue-eyed grass.
Scolecosporiella sisyrinchii (formerly Kellermannia sisyrinchii.) Leaf
Blight of blue-eyed grass.
Labrella
Pycnidia with a radiate shield, rounded; innate or erumpent; spores hyaline, one-celled.
Labrella aspidistrae. Leaf Blight of aspidistra.
Leptosphaeria
Perithecia membranous, not beaked, opening with an ostiole; innate or nally erumpent;
paraphyses present; spores dark, with several cells. Anamorph state a Coniothryium with
black, globose pycnidia and very small, dark, one-celled conidia, extruded in a black
cirrhus.
Diapleella coniothyrium (formerly Leptosphaeria (Melanomma) conithyri-
um; (Coniothyrium fuckelii)). Raspberry Cane Blight, general on rasp-
berry, dewberry, blackberry. The same fungus causes cankers of apple and
rose (
Cankers). On raspberry, brown dead areas extend into wood; whole

canes or single branches wilt and die; often between blossoming and fruit-
ing. The fungus enters the bark at any time during the season, through an
insect wound or mechanical injury. Smutty patches on the bark come from
BLIGHTS 185
small olive conidia of the Coniothyrium stage and larger, dark, four-celled
ascospores. Ascospores are spread by rain; conidia by rain and insects.
Control. Sanitation is very important; cut out and burn all diseased canes.
A control program for spur blight should sufce for cane blight.
Leptosphaeria (Melanomma) conithyrium (Coniothyrium fuckelii) (see
Diapleella coniothyrium) Raspberry Cane Blight, general on raspberry,
dewberry, blackberry.
Leptosphaeria korrae. Blight on turfgrass (associated with Fusarium blight
syndrome); disease is also called spring dead spot.
Leptosphaeria thomasiana. Cane Blight of dewberry, raspberry, in Pacic
Northwest.
Leptosphaeria sp. Blight; of Miscanthus.
Linospora
Leaf Spots.
Linospora tetraspora. Leaf Blight of poplar.
Lophodermella
Hymenium on a eshy-gelatinous stroma under the bark of woody plants; ascospores
aseptate.
Lophodermella sp. Needle Cast of pine.
Macrophomina
Rots.
Macrophomina phaseolina. Ashy Stem Blight, Charcoal Rot of soy-
beans, sweetpotatoes, many other plants.

Rots.
Micropeltis
A single hymenium, fruiting layer, covered with an open, reticulate scutellum; paraphy-
ses present; spores hyaline, with several cells.
Micropeltis viburni. Leaf Blight of viburnum.
Monilinia (Sclerotinia)
Stroma is a sclerotium formed in fruit by the fungus digesting eshy tissues and replac-
ing them with a layer of broad, thick-walled, interwoven hyphae forming a hollow
sphere enclosing core or seed of fruit, which has become a dark, wrinkled, hard mum-
my. Apothecia funnel-form or cupulate, rarely at-expanded, some shade of brown; asci
eight-spored; ascospores one-celled, ellipsoidal, often slightly attened on one side, hya-
line. Conidia hyaline, one-celled, formed in chains in grayish masses called sporodochia.
Monilinia azaleae. Shoot Blight of native or pinxter azalea (Rhododendron
roseum). Apothecia are formed on overwintered mummied fruits (capsules)
in leaf mold under shrubs in moist places. Ascospores infect leaves and suc-
culent shoots when the azalea is in full bloom. The conidial stage is common
on young developing fruits in late June and July (New York).
Monilinia fructicola. Leaf Blight and Shoot Blight of peach.
Monilinia johnsonii. Leaf Blight, Fruit Rot of hawthorn.
Monilinia laxa. Blossom Blight, Brown Rot of apricot, almond, cherry,
plum, and prune on Pacic Coast. Blossoms and twigs are blighted with
a good deal of gum formation. Monilinia laxa is sometimes coincident with,
and confused with, M. fructicola, which causes a more general rot of stone
fruits. Both are discussed more fully under Rots.
Monilinia rhododendri (Sclerotinia seaveri). Twig Blight, Seedling
Blight of sweet cherry.
Mycosphaerella
Anthracnose.
Mycosphaerella citrullina (M. melonis) conidial stage Didymella bryoni-
ae. Gummy Stem Blight, Stem End Rot, Leaf Spot of watermelon,
BLIGHTS 187
muskmelon, summer squash, pumpkin, and cucumber. Gray to brown dead
areas in leaves are marked with black pycnidia; leaves may turn yellow and
shrivel. Stem infection starts with a water-soaked oily green area at nodes.
The stem is girdled, covered with a dark exuded gum, and the vine wilts back
to that point. Fruit rot starts gray, darkens to nearly jet black, with gummy
exudate.
Control. Clean up crop refuse; practice rotation. Some varieties are more
resistant than others.
Mycosphaerella jiensis. Black Sigatoka on dwarf banana in FL.
Mycosphaerella melonis. Gummy Stem Blight of cucumbers.
Mycosphaerella pinodes. Pea Blight.
Ascochyta pinodes.
Mycosphaerella rabiei (Anamorph, Phoma rabiei). Blight of chickpea
Mycosphaerella sequoiae. Needle Blight of redwood.
Myriogenospora
Ascomata supercial or in a stroma, eshy, bright-colored; ascus with a thick cap tra-
versed by a slender pore; ascospores lifrom, multiseptate, often fragmenting.
Myriogenospora atramentosa. Blight on turf grass, centipedegrass.
Mystrosporium
Conidia dark, muriform; hyphae long.
Bipolaris iridis (formerly Mystrosporium adustum). Leaf Blight, Ink Spot
of bulbous iris; also on montbretia and lachenalia. Irregular black patches or
blotches appear soon after leaves push through the ground; under moist con-
ditions the foliage withers and dies prematurely. Inky black stains appear on
husks of bulbs (usually Iris reticulata), and yellow dots or elongated sunken
black craters show on eshy scales. The bulbs may rot,leaving only the husk
and a mass of black powder. The fungus spreads through the soil, invading
adjacent healthy bulbs.
Control. Dig bulbs every year; discard all diseased bulbs and debris; plant in
a new location. Spray with bordeaux mixture.
Mystrosporium adustum (see Bipolaris iridis). Leaf Blight, Ink Spot of
bulbous iris; also on montbretia and lachenalia.
Myxosporium
Conidia hyaline, one-celled, in discoid to pulvinate acervuli on branches.
Myxosporium diedickei. Twig Blight of mulberry.
Myxosporium everhartii. Twig Blight of dogwood. M. nitidum. Twig
blight and dieback of native dogwood. Prune twigs back to sound wood;
feed and water trees.
Neopeckia
Perithecia hairy, not beaked, formed on a mycelial mass; paraphyses present; spores
two-celled, dark.
Neopeckia coulteri. Brown Felt Blight on pines only, otherwise similar
to brown felt blight caused by Herpotrichia, a disease of high altitudes on
foliage under snow.
Ovulinia
Stroma a sclerotium, thin, circular to oval, shallowly cupulate, formed in petal tissue
but falling away; minute globose spermatia; apothecia of Sclerotinia type, small; asci
eight-spored; paraphyses septate with swollen tips; conidia large, obovoid, one-celled
except for basal appendage or disjunctor cell; borne singly at tips of short branches of
mycelium forming a mat over surface of petal tissue (see Fig. 3.9).
Ovulinia azaleae. Azalea Flower Spot, Petal Blight, very destructive
to southern azaleas in humid coastal regions, occasional on mountain-laurel
and rhododendron. Starting as a sudden outbreak near Charleston, South Car-
olina, 1931, the disease spread rapidly north of Wilmington, North Carolina,
down the coast to Florida, and around the Gulf. It reached Texas by 1938 and
BLIGHTS 189
was in California by 1940; it was reported in Maryland in 1945, in Virginia
in 1947, and in Philadelphia in 1959. Petal blight was reported from a Long
Island, New York, greenhouse in 1956, apparently present there since 1952,
and in 1959 infected all the azaleas in one New Jersey greenhouse. In both
cases the blight started on plants purchased from the South. This is the most
spectacular disease that I have ever witnessed, with most of the bloom on
all the azaleas in a town blighting simultaneously and seemingly overnight
under special weather conditions. The blight does not injure stem or foliage;
it is conned to the owers. The loss is aesthetic and economic from the
standpoint of tourist trade. For many years, before a control program was
worked out, the great azalea gardens of the South had to close their gates to
visitors far too early in the season.
Primary infection comes from very small apothecia produced from sclero-
tia on the ground under shrubs, usually in January or February, occasionally
as early as December. Spores shot into the air are carried by wind drift to
owers near the ground of early varieties, initial spots being whitish. If you
put your nger on such a spot, the tissue melts away. With continued high
humidity, heavy fog, dew, or rain, conidia are produced over the inner sur-
faces of petals and are widely disseminated to other petals by wind, insects,
and splashed rain. Within a few hours colored petals are peppered with small
white spots, and white owers have numerous brown spots. By the next day
owers have collapsed into a slimy mush, bushes looking as if they had had
scalding water poured over them. If the weather stays wet, small black scle-
rotia are formed in the petals in another 2 or 3 days. Infected blooms seldom
drop normally but remain hanging on the bushes in an unsightly condition
for weeks and months, some even to the next season. Many of the sclerotia,
however, drop out and remain in the litter on the ground ready to send up
apothecia the next winter.
Both Indian and Kurume varieties are attacked, the peak of infection com-
ing with mid-season varieties such as Pride of Mobile or Formosa. In some
seasons dry weather during early spring allows a good showing of azaleas;
in other years blight starts early and there is little color unless azaleas are
sprayed. On Belgian azaleas in greenhouses blight may start in December.
Control. Some mulches and soil treatments will inhibit apothecial produc-
tion. Secondary infection is bound to come from some untreated azalea in
the neighborhood. Spraying gives very effective, even spectacular, control
if started on time, when early varieties are in bloom and midseason azaleas
are showing color. Sprays must be repeated three times a week as long as
petal surface is expanding, about 3 or 4 weeks. After that, weekly spraying
is sufcient. Spraying is mandatory now for the big azalea gardens, and the
admission fees from the lengthened season pay for the program many times
over.
The original successful formula was: Dithane D-14 (nabam) 1 1/3 quarts to
100 gallons water, plus 1 pound 25% zinc sulfate, 1/2 pound hydrated lime,
and 1 ounce of spreader Triton B 1956. Later work showed that the lime
could be omitted, Dithane reduced to 1 quart, and zinc sulfate to 2/3 pound
to prevent injury in periods of drought. The spray should be a ne mist,
applied from several directions to get adequate coverage.
Commercial growers should beware of ordering azaleas from the South
unless they are bare-rooted and all ower buds showing color removed. As
a matter of fact, any potted or balled and burlapped plant grown in a nursery
near azaleas could very easily bring along some of the tiny sclerotia in the
soil, and they might remain viable more than 1 year. All traces of soil should
be washed off roots, and the plants wrapped in polyethylene for shipping.
Pellicularia
Includes some species formerly assigned to Corticium, Hypochnus, and Peniophora.
Hyphae stout, very short-celled; mycelium branching at right angles; basidia very stout,
formed on a resupinate, cottony or membranous layer of mycelium. Anamorph state
a Rhizoctonia, with sclerotia made up of brown, thin-walled, rather angular cells, or
Sclerotium, with sclerotia having a denite brown rind and light interior.
Athelia rolfsii (formerly Pellicularia rolfsii (Anamorph, Sclerotium rolfsii).
Southern Blight, Crown Rot. The disease has been known, in its scle-
rotium stage, for many years on hundreds of plants. The connection with
Pellicularia is recent, and the name does not have universal agreement. One
strain of the fungus has been called Sclerotium delphinii in the North, where
the disease is usually designated crown rot. This is, however, a variable fun-
gus with single spore cultures from the Pellicularia stage producing sclero-
tia typical of Sclerotium delphinii and of S. rolfsii, with intermediate forms.
Sclerotia of the southern blight strain are very small, round, tan, about the
size, shape, and color of mustard seed, the pathogen being frequently called
the mustard-seed fungus.
BLIGHTS 191
Southern blight affects almost all plants except eld crops like wheat, oats,
corn, and sorghum. Fruits and vegetables include Jerusalem artichoke, avo-
cado, bean, beet, carrot, cabbage, cucumber, eggplant, endive, lettuce, mel-
on, okra, onion, garlic and shallot, pea, peanut, pepper, potato, rhubarb,
strawberry, sweetpotato, tomato, turnip, and watermelon. Ornamentals, too
numerous to list in entirety, include ajuga, ageratum, amaryllis, azalea,
caladium, calendula, campanula, canna, carnation, cosmos, China aster,
chrysanthemum, dahlia, delphinium, daphne, duranta, gladiolus, hollyhock,
hydrangea, iris, jasmine, lemon verbena, lily, lupine, marigold, morning-
glory, myrtle, narcissus, orchids, phlox, pittosporum, rose, rose-mallow,
rudbeckia, scabiosa, sedum, sweet pea, star-of-bethlehem, tulip, violet, and
zinnia.
The rst sign of blight is the formation of white wefts of mycelium at the
base of the stem, spreading up in somewhat fan-shaped fashion and some-
times spreading out over the ground in wet weather. The sclerotia formed in
the wefts are rst white, later reddish tan or light brown. They may be numer-
ous enough to form a crust over the soil for several inches around a stem, or
they may be somewhat sparse and scattered.
In the white stage, droplets of liquid often form on the sclerotia, and the
oxalic acid in this liquid is assumed to kill plant cells in advance of the fungus
hyphae. This means that the pathogen never has to penetrate living tissue
and explains why so many different kinds of plants succumb so readily to
southern blight. Fruits touching the ground, as well as vegetables with eshy
roots, like carrots and beets, or plants with bulbs or rhizomes, like onions,
narcissus, and iris, seem particularly subject to this disease. Low ornamentals
such as ajuga blight quickly, the whole plant turning black; tall plants like
delphinium rot at the crown and then die back or topple over; bulbs have
a cheesy interior, with sclerotia forming on or between the scales.
Control. Remove diseased plants as soon as they are noticed. Take out sur-
rounding soil, for 6 inches beyond the diseased area, wrapping it carefully
so that none of the sclerotia drop back. Increasing the organic content of the
soil reduces southern blight, as does the addition of nitrogenous fertilizers,
such as ammonium nitrate. Treating narcissus bulbs in hot water for 3 hours,
as for nematodes, kills the fungus in all except the very largest bulbs.
Pellicularia lamentosa (see Thanatephorus cucumeris), teleomorph state
of Rhizoctonia solani. This is a variable fungus with some strains or forms
causing leaf blights but best known as cause of Rhizoctonia rot of potatoes
and damping-off of many plants.

Rots.
Pellicularia lamentosa f. sp. microsclerotia (Corticium microsclerotia)
see Thanatephorus cucumeris. Web Blight of snap bean, lima bean, also
reported on g, elder, hibiscus, hollyhock, tung oil, and phoenix tree, from
Florida to Texas. Many small brown sclerotia and abundant weblike myceli-
um are found on bean stems, pods, and foliage. Infection starts with small
circular spots that appear water-soaked or scalded. They enlarge to an inch or
more, become tan with a darker border, are sometimes zonate. The whitish
mycelium grows rapidly over the leaf blade, killing it, and spreads a web
from leaf to leaf, over petioles, owers, and fruit, in wet weather and at tem-
peratures 70 to 90F; in dry weather growth is inconspicuous except on
fallen leaves. The fungus is spread by wind, rain, irrigation water, cultivating
tools, and bean pickers; it survives in sclerotial form from season to season.
Control. Destroy infected plants; clean up refuse. In Florida, do not plant
beans between June and September if web blight has been present. Use a cop-
per spray or dust.
Pellicularia lamentosa f. sp. sasakii. See Thanatephonus cucumeris. Leaf
Blight of grasses, clover, etc.
Pellicularia lamentosa f. sp. timsii. See Thanatephonus cucumeris. Leaf
Blight of g.
Pellicularia rolfsii. See Athelia rolfsii, Anamorph,

Sclerotium rolfsii.
Southern Blight, Crown Rot. The disease has been known, in its scle-
rotium stage, for many years on hundreds of plants. The connection with
Pellicularia is recent, and the name does not have universal agreement.
One strain of the fungus has been called Sclerotium delphinii in the North,
where the disease is usually designated crown rot. This is, however, a vari-
able fungus with single spore cultures from the Pellicularia stage producing
sclerotia typical of Sclerotium delphinii and of S. rolfsii, with intermediate
forms. Sclerotia of the southern blight strain are very small, round, tan, about
the size, shape, and color of mustard seed, the pathogen being frequently
called the mustard-seed fungus.
Southern blight affects almost all plants except eld crops like wheat, oats,
corn, and sorghum. Fruits and vegetables include Jerusalem artichoke, avo-
cado, bean, beet, carrot, cabbage, cucumber, eggplant, endive, lettuce, mel-
on, okra, onion, garlic and shallot, pea, peanut, pepper, potato, rhubarb,
strawberry, sweetpotato, tomato, turnip, and watermelon. Ornamentals, too
numerous to list in entirety, include ajuga, ageratum, amaryllis, azalea,
caladium, calendula, campanula, canna, carnation, cosmos, China aster,
chrysanthemum, dahlia, delphinium, daphne, duranta, gladiolus, hollyhock,
BLIGHTS 193
hydrangea, iris, jasmine, lemon verbena, lily, lupine, marigold, morning-
glory, myrtle, narcissus, orchids, phlox, pittosporum, rose, rose-mallow,
rudbeckia, scabiosa, sedum, sweet pea, star-of-bethlehem, tulip, violet, and
zinnia.
The rst sign of blight is the formation of white wefts of mycelium at the
base of the stem, spreading up in somewhat fan-shaped fashion and some-
times spreading out over the ground in wet weather. The sclerotia formed in
the wefts are rst white, later reddish tan or light brown. They may be numer-
ous enough to form a crust over the soil for several inches around a stem, or
they may be somewhat sparse and scattered.
In the white stage, droplets of liquid often form on the sclerotia, and the
oxalic acid in this liquid is assumed to kill plant cells in advance of the fungus
hyphae. This means that the pathogen never has to penetrate living tissue
and explains why so many different kinds of plants succumb so readily to
southern blight. Fruits touching the ground, as well as vegetables with eshy
roots, like carrots and beets, or plants with bulbs or rhizomes, like onions,
narcissus, and iris, seem particularly subject to this disease. Low ornamentals
such as ajuga blight quickly, the whole plant turning black; tall plants like
delphinium rot at the crown and then die back or topple over; bulbs have
a cheesy interior, with sclerotia forming on or between the scales.
Control. Remove diseased plants as soon as they are noticed. Take out sur-
rounding soil, for 6 inches beyond the diseased area, wrapping it carefully
so that none of the sclerotia drop back. Increasing the organic content of the
soil reduces southern blight, as does the addition of nitrogenous fertilizers,
such as ammonium nitrate. Treating narcissus bulbs in hot water for 3 hours,
as for nematodes, kills the fungus in all except the very largest bulbs.
Pellicularia koleroga (Corticium stevensii). Thread Blight, a southern dis-
ease, from North Carolina to Texas, important on g and tung, sometimes
defoliating pittosporum, crape myrtle, roses, and other ornamentals, and
some fruits. The disease is recorded on apple, azalea, banana shrub, black-
berry, boxwood, camphor, cherry laurel, chinaberry, columbine, crabapple,
crape myrtle, casuarina, currant, dewberry, dogwood, elderberry, elm, eryth-
rina, euonymus, g, owering almond, owering quince, goldenrod, goose-
berry, guava, honeysuckle, hibiscus, morning glory, pear, pecan, pepper vine,
persimmon, pittosporum, plum, pomegranate, quince, rose, satsuma orange,
soapberry, silver maple, sweetpotato, tievine (Jacquemontia), tung, Virginia
creeper, and viburnum.
The fungus winters as sclerotia on twigs and leaf petioles, and in May and
June produces threadlike mycelium that grows over lower surface of leaves,
killing them and causing premature defoliation, although often dead leaves
hang on the tree in groups, matted together by thread-like spider webs.
Fruiting patches on leaves are rst white, then buff. The fungus ourishes in
moist weather, temperatures 75 to 90F.
Control. On gs, one or two applications of tribasic copper sulfate, or bor-
deaux mixture, are satisfactory until the fruit ripens in July. Pruning out
infected branches may be sufcient on tung and pecan, but at least one spray
of bordeaux mixture may be required.
Thanatephonus cucumeris (formerly Pellicularia lamentosa), teleomorph
state of Rhizoctonia solani. This is a variable fungus with some strains or
forms causing leaf blights but best known as cause of Rhizoctonia rot of
potatoes and damping-off of many plants.

Rots.
Thanatephonus cucumeris (formerly Pellicularia lamentosa f. sp. micro-
sclerotia (Corticium microsclerotia)). Web Blight of snap bean, lima bean,
also reported on g, elder, hibiscus, hollyhock, tung oil, and phoenix tree,
from Florida to Texas. Many small brown sclerotia and abundant weblike
mycelium are found on bean stems, pods, and foliage. Infection starts with
small circular spots that appear water-soaked or scalded. They enlarge to an
inch or more, become tan with a darker border, are sometimes zonate. The
whitish mycelium grows rapidly over the leaf blade, killing it, and spreads
a web from leaf to leaf, over petioles, owers, and fruit, in wet weather
and at temperatures 70 to 90F; in dry weather growth is inconspicuous
except on fallen leaves. The fungus is spread by wind, rain, irrigation water,
cultivating tools, and bean pickers; it survives in sclerotial form from season
to season.
Control. Destroy infected plants; clean up refuse. In Florida, do not plant
beans between June and September if web blight has been present. Use
a copper spray or dust.
Thanatephonus cucumeris (formerly Pellicularia lamentosa f. sp. sasakii).
Leaf Blight of grasses, clover, etc.
Thanatephorus cucumeris (formerly Pellicularia lamentosa f. sp. timsii).
Leaf Blight of g.
BLIGHTS 195
Penicillium
Cankers.
Penicillium oxalicum. Leaf Blight of grass.
Pestalotia
Acervuli dark, discoid or cushion-shaped, subcutaneous; conidiophores short, simple;
conidia fusiform, several-celled with median cells colored, end cells hyaline, a short
stalk at the basal cells and a crest of two or more hyaline appendages, setae, from the
apical cell (Fig. 3.9). Weak parasites or saprophytes; some are treated under Leaf Spots.
Pestalotia funerea (see Pestalotiopsis funerea). Tip Blight of conifers,
Needle Blight, Twig Blight of chamaecyparis, retinospora, cypress, bald
cypress, arborvitae, juniper, yew, and giant sequoia.
Pestalotia hartigii. Associated with a basal stem girdle of young conifers
but parasitism not proven. The stem has a swelling above the girdling lesions,
and the tree gradually turns yellow and dies. The effect may be more from
high temperature than the fungus; shading transplants is helpful.
Pestalotia sp. and Penicillium sp. Flower Blight on camellia.
Pestalotiopsis funerea (formerly Pestalotia funerea). Tip Blight of conifers,
Needle Blight, Twig Blight of chamaecyparis, retinospora, cypress, bald
cypress, arborvitae, juniper, yew, and giant sequoia. The fungus is sapro-
phytic on dead and dying tissue and also weakly parasitic, infecting living
tissue through wounds under moist conditions. It appears in sooty pustules
on leaves, bark, and cones.
Phacidium
Apothecia innate, concrete above with the epidermis and slitting with it into lobes; spores
one-celled, hyaline.
Phacidium abietinellum (see Nothophacidium abietinellum). Needle
Blight of balsam r.
Phacidium balsameae (see Sarcotrochilia balsameae). Needle Blight of
balsam r in New England, of white and alpine r in the Northwest.
Phacidium infestans. Snow Blight of conifer seedlings on r and young
pines in the Northeast, also on arborvitae and spruce; on white and alpine r
in the Northwest. This native fungus is most damaging in nurseries, attack-
ing foliage under the snow. The needles turn brown, with a covering of white
mycelium, just as the snow melts. In late summer and fall brown to nearly
black apothecia appear on underside of browned needles. Ascospores are
spread by wind, primary infection being in autumn. Additional infection
occurs in late winter, when mycelium grows out under the snow from dis-
eased to dormant, healthy needles.
Control. Spray nursery beds with dormant-strength lime sulfur in late fall;
remove infected seedlings; dip new stock in lime sulfur before planting.
Nothophacidium abietinellum (formerly Phacidium abietinellum). Nee-
dle Blight of balsam r.
Sarcotrochilia balsameae (formerly Phacidium balsameae). Needle
Blight of balsam r in New England, of white and alpine r in the North-
west.
Phaeoacremonium
Phaeoacremonium chlamydosporum. Black Goo on grape.
Phialophora
Rots.
Phialophora graminicola. Blight on turfgrasses (associated with Fusarium
blight syndrome).
Phloeospora
Pycnidia dark,imperfectly formed, globose, innate in tissue, not in distinct spots; conidia
hyaline or subhyaline, several-celled, elongate fusoid to liform; parasitic or saprophyt-
ic. One of the conidial forms linked with Mycosphaerella as a teleomorph state.
Phloeospora adusta. Leaf Blight of clematis.
BLIGHTS 197
Phoma
Blackleg.
Phoma conidiogena (see Phoma glomerata). Boxwood Tip Blight. Ashy
gray necrotic areas at leaf tips, with pycnidia on both leaf surfaces.
Phoma glomerata (formerly Phoma conidiogena). Boxwood Tip Blight.
Ashy gray necrotic areas at leaf tips, with pycnidia on both leaf surfaces.
Phoma fumosa. Twig Blight, occasional on maple.
Phoma macdonaldii. Blight, Premature Ripening of sunower.
Phoma mariae. Twig Blight on Japanese honeysuckle.
Phoma piceina. Twig and Needle Blight of Norway spruce. May cause
defoliation and sometimes death of forest trees.
Phoma sclerotioides. Brown Root Rot of alfalfa.
Phoma strobiligena (see Sclerophoma pythiophila), on cone scales of Nor-
way spruce.
Sclerophoma pythiophila (formerly Phoma strobiligena), on cone scales of
Norway spruce.
Phomopsis
Pycnidia dark, ostiolate, immersed, erumpent, nearly globose; conidiophores simple;
conidia hyaline, one-celled, of two types-ovate or ellipsoidal and long, lamentous,
sickle-shaped or hooked at upper end (Fig. 3.9). Anamorph state of Diaporthe; para-
sitic causing spots on various plant parts.
Dendrophoma obscurans (see Phomopsis obscurans). Strawberry Leaf
Blight, Angular Leaf Spot.
Phomopsis ambigua (teleomorph, Diaporthe eres). Twig Blight of pear,
widespread.
Phomopsis diospyri. Twig Blight of native persimmon.
Phomopsis japonica. Twig Blight of kerria.
Phomopsis juniperovora. Nursery Blight, Juniper Blight, Cedar
Blight, Canker on red-cedar and other junipers, cypress, chamaecyparis,
Japanese yew (Cephalotaxus), arborvitae, giant sequoia, and redwood. This
disease occurs in virulent form from New England to Florida and through
the Middle West; it may also occur on the Pacic Coast.
Tips of branches turn brown with progressive dying back until a whole
branch or even a young tree is killed. Trees over 5 years old are less seriously
injured. Spores produced in quantity in pycnidia on diseased twigs ooze out
in little tendrils in moist weather, to be spread by splashing water, insects,
and workers. Entrance is through unbroken tissue as well as wounds; the
stem is killed above and below the point of entrance. Small, sunken lesions
give a attened appearance to some seedlings. Overhead irrigation in a nurs-
ery is a predisposing factor, and a large amount of stock can be blighted in
a very short time. Older trees in home plantings suffer from twig blight. The
fungus winters on infected plant parts and remains viable at least 2 years.
Control. Have seedbeds well drained; water by ditch irrigation; remove and
burn diseased seedlings early in the season; keep seedbeds away from old-
er cedar trees; do not use cedar branches or needles for mulching. Spray
with xed copper or bordeaux mixture plus a wetting agent, starting when
growth begins and repeating to keep new foliage covered. Spiny Greek and
Hill junipers and Keteller red-cedars are somewhat resistant.
Phomopsis kalmiae. Mountain-Laurel Leaf Blight, Blotch. Circular,
brown, often zonate areas on leaves, frequently starting near margin or tip,
gradually enlarge and coalesce until most of the blade is involved. The fun-
gus often works down the petiole to cause a twig blight. The disease is more
prominent on bushes in the shade or under drip of trees. Remove blighted
leaves or clean up fallen leaves.
Phomopsis longicolla. Black Pod Spot and Seed on cowpea.
Phomopsis oblonga. Twig Blight on Chinese elm.
Phomopsis obscurans (formerly Dendrophoma obscurans). Strawberry
Leaf Blight, Angular Leaf Spot. The lesions are large, circular to angular,
reddish purple, zonate with age, having a dark brown center, a light brown
zone, and a purple border. Spots may extend in a V-shaped area from a large
vein to edge of the leaf, with black fruiting bodies appearing in the central
portion. Not serious before midsummer, the disease may be destructive late
in the season. The fungus winters on old leaves.
Phomopsis occulta. Shoot Blight of Colorado blue spruce.
Phomopsis vexans. Phomopsis Blight of eggplant.

Diaporthe vexans.
Phomopsis vaccinii. Twig Blight of blueberry.
BLIGHTS 199
Phyllosticta
Pycnidia dark, with ostiole, in spots in leaves; spores one-celled, hyaline. The charac-
teristics are the same as Phoma except that leaves rather than stems are infected. Other
species are listed under Leaf Spots.
Phyllosticta batatas. Sweet Potato Leaf Blight, occasional from New
Jersey to Florida, more prevalent in the South but seldom important enough
for control measures. Numerous white spots on leaves are bordered with
narrow reddish zones; pycnidia are numerous; spores are extruded in tendrils.
Phyllosticta cryptomeriae. Needle Blight found on Cryptomeria.
Phyllosticta lagerstroemiae. Tip Blight of crape-myrtle.
Phyllosticta multicorniculata. Needle Blight of r.
Phyllosticta pteridis. Tip Blight of fern. Leaves lose green color; spots are
ash gray with purple brown margins and numerous black pycnidia in center.
A very weak bordeaux mixture has been suggested for control; if overhead
watering is avoided, spraying may not be necessary.
Physalospora
Ascomycetes, Sphaeriales
Perithecia with papillate mouths, immerse in substratum but without well-dened stro-
mata; paraphyses present; spores one-celled, hyaline. A few species cause blights; many
cause rots.
Botryosphaeria obtusa (formerly Physalospora obtusa). Cane Blight of
rose, also Black Rot of apple, Canker and Dieback of many plants.
Cankers and also
Rots.
Glomerella cingulata (formerly Physalospora dracaenae). Dracaena Tip
Blight, Leaf Spot. Disease starts at the tips of lower leaves and spreads
down toward the base. Infected areas are sunken and straw-colored, dotted
with black specks of pycnidia. All leaves on the plant may die except a few
at the top. Remove infected leaves as soon as noticed. Spray with a copper
fungicide.
Physalospora dracaenae (
Glomerella cingulata). Dracaena Tip Blight,

Leaf Spot.
Physalospora gregaria. Twig Blight of yew.
Physalospora obtusa (see Botryosphaeria obtusa). Cane Blight of rose,
also Black Rot of apple, Canker and Dieback of many plants.
Phytophthora
Oomycetes, Peronosporales
This most important genus contains many species causing destructive blights, cankers,
and rots. The name, which means plant destroyer, was given in 1876 for the pota-
to blight fungus. Sporangia, formed successively on sporangiophores, slender, sparse-
ly branched hyphae emerging from stomata, germinate either by a germ tube or by
zoospores. The sexual spore is an oospore.
Phytophthora cactorum. Lilac Shoot Blight. Blossoms and succulent
growing tips are blighted and turn brown; suckers are killed back 4 or 5 feet.
Blight is most severe in wet springs when shrubs are crowded, shaded, and
improperly pruned. The same fungus causes a canker, foot rot, and dieback
of rhododendron and other plants and is considered again under Cankers.
Avoid planting lilacs and rhododendrons close together. Prune each year for
air circulation and to remove dead twigs.
Phytophthora capsici. Phytophthora Blight of Pepper, Leaf and Stem
Blight of Squash, Fruit Rot of pepper, eggplant, tomato, cucumber, and mel-
on. The disease was rst found in New Mexico in 1918 injuring chili pep-
pers; it occurs chiey in southwestern and Gulf states. In 1953, however, it
was reported that for some years it had been causing a leaf blight of squash
in North Carolina.
Pepper plants are girdled at the soil line with a dark green water-soaked
band, which dries and turns brown, followed by wilting and death of the
entire plant. Leaf spots are dark green and small at rst, later large bleached
or scalded areas. Dark, watersoaked patches on fruits are covered with white
mycelium. The fruit withers but remains attached; 60% of green fruit may
be infected in southwestern commercial plantings. Seed are infected from the
fruit. Symptoms on squash are somewhat similar; green leaf lesions spread-
ing over the blade, a basal stem rot, and wilting. Wet soil and high tempera-
tures encourage blight.
Control. Place seedbeds on land that has not previously grown peppers;
rotate crops. Avoid over irrigation.
Phytophthora citrophthora (also P. citricola and P. nicotianae var. para-
sitica). Shoot and Stem Blight on azalea. Needle Blight and Branch
Dieback of sequoia.
BLIGHTS 201
Phytophthora erythroseptica. Leaf Blight of pink and golden calla.
Leaves are wilted and distorted; petioles are black and soft.
Phytophthora ilicis. Holly Blight, Phytophthora Leaf and Twig Blight,
the most serious disease of English holly, particularly serious in the North-
west. For many years the trouble was ascribed to Boydia insculpta and called
Boydia canker, but this fungus merely invades tissue killed by Phytophtho-
ra. Leaf spots are dark, developing on lower leaves in cool rainy weather
and progressing upward in late fall and winter. Young twigs die back; black
stem cankers kill older twigs. Young plants in nurseries are defoliated and
sometimes killed.
Control. Choose a planting site with moderate air movement; space trees well
apart. Prune out all cankered and blighted twigs; prune also for air movement
through trees. Spray with tribasic copper sulfate, starting the middle of Octo-
ber.
Phytophthora infestans. Late Blight of potato and tomato, general on pota-
to in the Northeast, in Middle Atlantic and North Central states, sometimes
in Gulf and western states; on tomato in humid regions and seasons.
Here is a pathogen that has not lost its destructive virulence with passage of
time. In 1946, a whole century after potato blight caused the famous Irish
famine, tomato blight devastated tomatoes along the eastern seaboard, both
in home gardens and canning elds.
The potato went to Europe from South America shortly before 1600, seem-
ingly leaving its pathogens at home. For 200 years potatoes thrived in Europe
as the main source of carbohydrate food, but in August 1845, the Garden-
ers Chronicle reported: A fatal malady has broken out amongst the potato
crop. On all sides we hear of destruction. In Belgium the elds are said to
have been completely desolated. There is hardly a sound sample in Covent
Garden Market. The editor went on to describe the decay and to say: As
to cure for this distemper there is none. One of our correspondents is today
angry with us for not telling the public how to stop it; but he ought to con-
sider that Man has no power to arrest the dispensations of Providence. We
are visited by a great calamity which we must bear. And in 1946 American
gardeners were again blaming the editor, for lack of information on tomato
blight.
In 1845 the weather was continued gloom and fog, with below-average tem-
peratures. The Gardeners Chronicle editor was sure blight was due to pota-
toes being overladen with water. The Rev. M. J. Berkeley disagreed. He
insisted blight was due to a fungus, with the weather contributing to spread
of a moisture-loving parasite. The argument raged, for this was long before
Pasteur and his germ theory, and the rst time anyone believed a fungus
could be the cause and not the consequence of plant disease. A French scien-
tist, Montagne, named the fungus Botrytis infestans, but the rst really good
description of it was published by Berkeley, and it remained for the German
de Bary, in 1876, actually to prove the pathogenic nature of the fungus and
to erect the new genus Phytophthora to include it.
Meanwhile the disease was making history. The loss of the potato crop in
1845 and 1846 killed off a million people and caused another million and
a half to emigrate; the rst Government Relief program was instigated; and
the English Corn Laws were repealed with a change to a policy of free trade
and unbounded expansion of commerce.
Late Blight of Potato
Symptoms. After blossoming, large, dark green, water-soaked spots appear
on leaves in wet weather, rst on lower leaves. As a spot enlarges the center
is shriveled, dry, dark brown to black, and a downy, whitish growth appears
on the underside of leaves. Similar lesions are formed on stems and petioles,
and there is a characteristic strong odor as tops are blighted. On tubers, rst
symptoms are small brown to purple discolorations of skin on upper side,
changing to depressed pits when tubers are removed from soil and put in
storage (see Fig. 3.12). On cutting through the potato, a reddish brown dry
rot is seen.
Life History. The primary cycle starts with infected tubers, which have har-
bored mycelium in the dry rot patches over winter. If infected seed pieces
are planted, the fungus grows systemically into the shoots and nally fruits
by sending sporangiophores out through the stomata on lower leaf surfaces
(see Fig. 3.13). These swell at the tips into ovoid bodies, sporangia, then
branch and produce successively more sporangia. The latter may function
as conidia, putting out a germ tube, but more often are differentiated into
a number of swarmspores (zoospores), which have cilia enabling them to
swim about after they are splashed by rain to another leaf. Eventually they
stop swimming and send a germ tube in through the leaf cuticle or enter
through a stoma. Initial infection in the eld also comes from conidia blown
over from sprouts produced on infected tubers in cull piles. Blighting fol-
lows rapidly, with rst symptoms 5 days or less from the time of infection
BLIGHTS 203
Figure 3.12 Late Blight on Potato
Figure 3.13 Late Blight of Potatoes. Sporangiophores of Phytophthora infestans emerging from leaf, bearing
sporangia, sometimes called conidia, which germinate by zoospores
and with the fungus fruiting again in a whitish layer on the underside of
leaves.
Tubers with only a thin covering of soil may be infected by swarmspores
washing down onto them from blighted leaves overhead; they are also infect-
ed during digging if it is done in moist weather while tops are still green.
Swarmspores remain viable in the soil several weeks while awaiting favor-
able conditions. Oospores, the sexual spores, are apparently not required in
the life cycle for they are not found with potatoes grown in the eld. They
have been produced in culture.
Weather Relations. This is a disease entirely dependent on weather condi-
tions. Temperature and moisture conditions are right for an epiphytotic about
2 years out of 5. Zoospores are produced only in cool weather, 60F and
under, but they invade leaves most rapidly at higher temperatures. Because
they are swimming spores, rain is required. A cool, wet July is usually fol-
lowed by blight in August and September.
Control. Some varieties, such as Kennebec, Essex, Pungo, and Cherokee,
are resistant to the common strain of the fungus but not to some of the new-
er strains. Treat potato dumps and cull piles with a weed spray to control
sprouts. Delay digging crop until 2 weeks after tops die, or else kill the tops
with a weed killer to prevent infection at early digging.
Late Blight of Tomato
Although there are potato and tomato strains of Phytophthora infestans, each
is capable of infecting the other host. Ordinarily blight starts with potatoes in
midsummer; when the fungus moves over to tomatoes, it has to go through
several cycles to build up a strain virulent enough to produce general blight-
ing, and by that time the tomato season is nearly over. Now we know that
it is possible for the tomato strain to winter in potato tubers and be ready
to inict damage on tomatoes with the rst crop of zoospores produced on
potato sprouts. Conversely, tomato seedlings brought up from the South and
planted near potato elds can start an epiphytotic of late blight on pota-
toes.
The 1946 tomato blight saga the one that awakened eastern gardeners to
the fact that plant disease could be as important to home gardeners as to
farmers started in Florida late in November 1945. By January the disease
was extremely destructive in tomato seedbeds, and it continued so intermit-
tently whenever temperatures ranged from 60 to 70F and relative humid-
ity was nearly 100% for more than 15 hours. Evidence indicated spores
could be wind-borne for as far as 30 miles. The wave of late blight went
west to Alabama, taking 75% of the early crop, and rolled up the Atlantic
BLIGHTS 205
Figure 3.14 Late Blight on Tomato
Coast, reaching the Carolinas in May and Virginia and Maryland in June,
again taking 75% of the early crop. It rolled into Delaware and New Jer-
sey in July, but did not reach peak epidemic form until after an extended
rainy period in August, and ended in Massachusetts in August and Septem-
ber.
In 1947 a blight-forecasting service was started, based on weekly graphs pre-
pared by plotting daily the cumulative rainfall and mean temperatures and
aided by reports from key pathologists in various states. If conditions are
unfavorable for blight, we can save time and money by eliminating useless
spraying.
Symptoms. On seedlings small, dark spots on stems or leaves are followed
by death within 2 or 3 days. On mature plants blight starts with dark, water-
soaked leaf spots and large, dark brown spots on fruit, with most of the leaves
soon hanging lifeless and fruit rotting on the ground (see Fig. 3.14).
Control. Bordeaux mixture applied to young tomato plants will either pre-
vent fruit setting or cause stunting. It can be used after blossoming, or a xed
copper can be substituted.
Phytophthora meadii. Blight and Leaf Spots on West Indian holly.
Phytophthora medicaginis. Rot Root on Medicago spp.
Phytophthora nicotianae (formerly Phytophthora parasitica var, parasit-
ica. Leaf, Stem and Bub Blight on bougainvillea, dogwood, hibiscus,
artillery plant, and aluminum plant. Leaf blight; on jojoba.
Phytophthora parasitica var. parasitica (see Phytophthora nicotianae).
Leaf, Stem and Bub Blight on bougainvillea, dogwood, hibiscus, artillery
plant, and aluminum plant. Leaf blight; on jojoba.
Phytophthora syringae. Citrus Blight, also on lilac, but the more common
lilac blight is due to P. cactorum. On citrus trees leaves have semitransparent
spots similar to frost damage. Other Phytophthora species may be present
with P. syringae to cause brown rot of fruits.

Rots. On lilacs large irregu-
lar leaf patches have a lighter zone at margin. There may be some defolia-
tion.
Plectosporium
Plectosporium abacinum. Blight on Hydrilla.
Pyrenochaeta
Deuteromycetes, Sphaeropsidales, Sphaerioidaceae
Pycnidia dark, ostiolate, nearly globose, erumpent with a few bristles near ostiole; coni-
diophores simple or branched; conidia small, one-celled, hyaline, ovate to elongate; par-
asitic or saprophytic. See also under Rots.
Pyrenochaeta phlogis. Stem Blight of Phlox.
Pyricularia
Conidiophores long, slender, simple or rarely branched, septate, single or in tufts; coni-
dia pyriform to nearly ellipsoid, borne singly and attached at broader end; spores hyaline,
two- to three-celled; parasitic, chiey on grasses.
Pyricularia grisea. Leaf Blight on creeping bent grass and buffelgrass.
BLIGHTS 207
Pythium
Rots.
Pythium myriotylum. Blight of tomato.
Delphinella (Rehmiellopsis)
Perithecia single, globose, rupturing irregularly; asci in fascicles, no paraphyses; spores
hyaline, two-celled.
Delphinella balsameae (formerly Rehmiellopsis balsameae). Tip Blight,
Needle Blight of balsam r, on native balsam r in northern New England
and on ornamental rs in southern New England and New York. Infection is
in spring with needles of current season shriveled, curled, and killed, often
with a dieback of terminal or lateral shoots and sometimes cankers at base of
infected needles. Satisfactory control on ornamental rs has been obtained
by three sprays, at 10-day intervals, of bordeaux mixture, the rst application
made as new growth starts.
Rehmiellopsis balsameae (see Delphinella balsameae). Tip Blight, Nee-
dle Blight of balsam r, on native balsam r in northern New England and
on ornamental rs in southern New England and New York.
Rhizoctonia
Deuteromycetes, Mycelia Sterilia (Fungi Imperfecti)
Sclerotial form of some species of Pellicularia, Corticium, Macrophomina, and Heli-
cobasidium. Young mycelium colorless, with branches constricted at points of origin
from main axis, but soon colored, a weft of brownish yellow to brown strands, organized
into dense groups, sclerotia made up of short, irregular, angular or somewhat barrel-
shaped cells.
Rhizoctonia ramicola. Silky Thread Blight a southern disease similar to
web blight caused by Pellicularia koleroga. Perennial ornamental hosts in
Florida include elaeagnus, erythrina, crape-myrtle, holly, guava, pittospo-
rum, pyracantha, Carolina jessamine, feijoa, and rhododendron. Tan spots
with purple-brown margins appear on leaf blades, dead lesions on petioles
and young twigs. When leaves are abscissed, they are often held dangling and
matted together by brown fungus threads. Infection recurs annually in moist
weather with high daytime temperatures. The fungus winters as mycelium
in leaf lesions and diseased twigs. Sclerotia are apparently lacking in this
species.
Rhizoctonia sp. (teleomorph, Aquathanatephorus pendulus). Blight on
water hyacinth.
Rhizoctonia sp. (teleomorph, Thanatephorus cucumeris). Blight on beet.
Needle blight on pine.
Rhizoctonia solani. Blight of pistachio and Cynodon spp., and Foliar
Blight of soybean.
Rhizopus
Rots.
Rhizopus stolonifer. Seedling Blight on lupine; also caused by Pleio-
chaeta setosa, Alternaria sp., Aspergillus avus, Aspergillus niger, and
Curvularia sp.
Rosellinia
Ascomycetes, Xylariales
Perithecia separate, supercial from the rst, carbonaceous, not beaked, ostioles papil-
late; spores dark, one-celled with a small groove.
Rosellinia herpotrichioides. Hemlock Needle Blight. Needle-bearing
portions of twigs become covered on underside with a grayish brown
mycelial mat; black perithecia are produced in this mat in great abundance.
Ovoid, hyaline conidia are formed on Botrytis-like conidiophores.
Schirrhia
Ascomycetes, Dothideales, Dothideaceae
Asci usually short, cylindrical, and relatively numerous in spherical, ostiolate locules.
BLIGHTS 209
Scleropycnium
Pycnidia open out to a deep cupulate or discoid structure, tough, dark or black, subepi-
dermal or subcortical, then erumpent; spores hyaline, one-celled. Largely saprophytic
on twigs, sometimes parasitic on leaves.
Scleropycnium aureum. Leaf Blight of mesquite.
Sclerotinia (Whetzelinia)
Apothecia arising from a tuberoid sclerotium which, though formed free on aerial
mycelium, is sometimes enclosed in natural cavities of suscept or host, as in hollow
stem of perennials. Interior (medulla) of sclerotium white, completely enveloped by
a dark rind; gelatinous matrix lacking. Conidia wanting but spermatia (very small
microconidia) formed on sporodochia borne free or enclosed in cavities. Apothecia
some shade of brown; cupulate to funnel-form; usually at maturity saucer-shaped to
at expanded; ascospores hyaline, one-celled, ovoid. Species formerly included in
Sclerotinia but possessing monilioid conidia are now in Monilinia.
Botryotinia polyblastis (formerly Sclerotinia polyblastis). Narcissus Fire.
A serious ower blight in England, known here on the Pacic Coast. In
England overwintering sclerotia produce apothecia when Narcissus tazetta
comes into ower, the ascospores infecting the perianth and causing ower
spotting. From withered owers numerous large conidia, germinating with
several germ tubes, infect foliage, on which large sclerotia are formed late
in the season. Remove infected parts immediately; spray early in the sea-
son.
Ciberinia camelliae (formerly Sclerotinia camelliae). Camellia Flower
Blight, long known in Japan, rst noted in California in 1938, conrmed in
Georgia in 1948, although probably there several years previously, reported
in Oregon in 1949, Louisiana and North Carolina in 1950, South Carolina in
1954. The blight is now widespread in Virginia, conned to certain counties
in other states. It was not ofcially recorded from Texas until 1957 but must
have been there earlier. The 1950 outbreak at Shreveport, Louisiana, is said
to have started on plants brought in from Texas that probably originated in
California.
Floral parts only are affected, infection taking place any time after tips
of petals are visible in opening buds. Few to many brownish specks on
expanding petals enlarge until the whole ower turns brown and drops. In
early stages darkened veins are prominent diagnostic symptoms. When the
owers rest on moist earth, spermatia are produced on petals in shiny black
masses. Hard, dark brown to black sclerotia formed at the base of petals
frequently unite into a compound structure simulating petal arrangement.
This compound sclerotium may be an inch or more in diameter. Although
the petals do not melt when touched as do azaleas with petal blight, there is
a distinctive moist feeling that helps to differentiate ower blight from frost
injury. Rarely, a ower blight of camellias is caused by another Sclerotinia
(S. sclerotiorum).
Sclerotia lie dormant on ground or in mulching materials until the next winter
when, from January on (possibly earlier), after wet periods with rising tem-
perature, they produce one to several apothecia on long or short stipes with
brown, saucerlike discs 1/4 to 3/4 inch across, rarely up to 1 inch. Spores,
discharged forcibly, are carried by wind currents to owers, thus completing
the cycle. Spores may be wind-borne at least 1/3 mile, but presumably a large
proportion of them land on opening petals of the bush overhead. The scle-
rotia remain viable in the soil at least 2 or 3 years, sending up more apothe-
cia each season. No conidia are known; so there is no secondary infection
from ower to ower as with azalea blight. The amount of primary inocu-
lum is very large, however. One afternoon in New Orleans I collected nearly
1000 sclerotia that were producing apothecia from under a single camel-
lia.
Control. The rst line of defense is exclusion. Most southern states have
quarantines against known infected areas; they require that plants be shipped
bare-rooted, with all ower buds showing color removed. Northern garden-
ers ordering plants for greenhouses should insist on the same precautions
even without specic quarantines. Practically all outbreaks of camellia ow-
er blight have been traced to plants shipped in cans, presumably carrying
sclerotia in the soil. The disease has also appeared on owers shipped in by
air for camellia shows. Schedules should state that all specimens become the
property of the show committee, to be destroyed at the end of the show; no
blooms should be taken home for propagation.
Theoretically, because there is no conidial stage to spread the fungus, this
should be an easy disease to eradicate, but it has not proved so in prac-
tice. Camellias have thousands of owers produced over a period of months.
BLIGHTS 211
They drop into various ground covers, and it is almost impossible to nd and
destroy all infected blooms before rotting tissues release sclerotia into the
litter. Some cities have quarantined infected properties and provided a host-
free period of 2 years, during which all ower buds are removed from all
camellias in the area, but this approach has been only partially successful.
Various chemicals have been tried as ground treatment to inhibit formation
of apothecia.
Sclerotinia camelliae (see Ciberinia camelliae). Camellia Flower Blight,
long known in Japan, rst noted in California in 1938, conrmed in Georgia
in 1948, although probably there several years previously, reported in Oregon
in 1949, Louisiana and North Carolina in 1950, South Carolina in 1954.
Sclerotinia minor. Blight of soybean, peanut, and Eclipta.
Sclerotinia (Botryotinia) polyblastis (see Botryotinia polyblastis). Narcis-
sus Fire. A serious ower blight in England, known here on the Pacic
Coast. Sclerotinia rolfsii. Southern Blight on St. Johnswort.
Sclerotinia sclerotiorum. Shoot and Twig Blight of lilac, grape, pista-
chio, soybean, peanut, and malaviscus; ower blight of camellia resembling
that caused by S. camelliae but far less serious. Tuber blight and storage rot;
of Trillium. This ubiquitous fungus more often causes stem rots on its many
different hosts.

Rots.
Sclerotium
Deuteromycetes, Mycelia Sterilia (Fungi Imperfecti)
AsexuaI fruit bodies and spores lacking; there is merely a resting body, sclerotium, made
up of a compact, rounded mass of light-colored hyphae with a brown to black rind;
parasitic, often on underground plant parts. Pellicularia has proved to be the teleomorph
state for some forms.
Sclerotium bataticola. Ashy Stem Blight. See Macrophomina phaseoli
under Rots.
Sclerotium hydrophilum Blight of wild rice.
Sclerotium oryzae. Blight of wild rice.
Sclerotium rhizodes. White Tip Blight of grass.

Snowmold.
Sclerotium rolfsii. Southern Blight.
Pellicularia rolfsii.
Septoria
Pycnidia dark, separate, globose, ostiolate; produce in spots, erumpent; conidiophores
short, conidia hyaline, narrowly elongate to liform, several septate; parasitic, typically
causing leaf spots, but also blights and blotches (see Fig. 3.9). There are about 1000
species.
Septoria apiicola (Syn. Septoria apii and S. apii-graveolentis). Celery
Late Blight, general on celery, also on celeriac. The two species, singly or
together, produce the disease known as late blight, rst reported in Delaware
in 1891 and since causing much crop destruction, one California county
reporting half a million dollars loss from celery blight in 1908 and Michigan
a million in 1915. It was not known until 1932 that two distinct species were
involved.
Early symptoms are similar. Large leaf spot, due to S. apii, starts as a light
yellow area, which soon turns brown and dies. Spots are up to 1/4 inch
in diameter, with small black pycnidia. In small leaf spot, due to S. apii-
graveolentis, the more common and destructive pathogen, pycnidia appear at
the rst sign of chlorotic spotting and are often outside of the indenite mar-
gins of the spots, which are not over 2 mm. If infection is severe, the spots
fuse, and the leaves turn brownish black and rot. Leaf stalks may also be
infected. Pycnidia winter on seed and in plant refuse in garden and compost.
A single pycnidium of the small-spot fungus has an average of 3675 spores,
extruded in gelatinous tendrils. A single leaf spot may average 56 pycnidia,
and a single plant may have 2000 spots. Thus there are enormous amounts of
inoculum to be spread by rain, insects, people, and tools. Some years ago on
Long Island, when celery was inter-cropped with spinach, it was found that
workers spread blight spores on their sleeves as they cut the spinach in early
morning dew. And there is a case on record where a man walked through
his own blighted celery before taking a diagonal path across his neighbors
healthy eld. In a few days blight showed up all along that diagonal path.
Control. The fungus usually dies in the seed coat while the seed is still
viable. Using celery seed more than 2 years old obviates the necessity for
treatment. Fresh seed can be soaked in hot water for 30 minutes at 118
to 120F. Use crop rotation; do not plant near where celery was grown the
year before. Spray with bordeaux mixture or a xed copper, starting in the
seedbed when plants are just out of the ground.
BLIGHTS 213
Septoria leucanthemi*. Leaf Blight, Blotch on chrysanthemum, shasta
daisy, and oxeye daisy. The generally destructive Septoria on chrysanthe-
mum is S. chrysanthemi.

Leaf Spots.
Septoria petrosellini. Leaf Blight of parsley, similar to late blight of celery
but conned to parsley.
*Recent study indicates these are one species and that the name should be
S. apiicola.
Septotinia
Stroma a denite, small, thin, elongate to angular black sclerotium maturing in host
tissue after it has fallen to ground. Apothecia shallow cup-shaped, stipitate; spores hya-
line, ovoid, one-celled. Conidial stage a Septotis, with hyaline spores, two or more cells,
formed on sporodochia.
Septotinia podophyllina. Leaf Blight of may-apple, found on leaves and
stalks of this plant only.
Servazziella
Ascomycetes, Amphisphaeriales
Perithecia immersed in a stroma, with long necks converging into a disc; ascospores
long, liform, hyaline; conidia on a stroma.
Cryptospora longispora (see Servazziella longispora). Araucaria Branch
Blight.
Servazziella longispora (formerly Cryptospora longispora). Araucaria
Branch Blight. Lower branches are attacked rst, with disease spreading
upward; tip ends are bent and then broken off; plants several years old may
be killed. Prune off and burn infected branches.
Sirococcus
Deuteromycetes, Sphaeropsidales, Sphaerioidaceae.
Small, rounded, black, semi-immersed pycnidia with wide ostioles; conidia hyaline,
fusiform, slightly constricted, 1-septate.
Sirrococcus elavigignenti-juglandacearum. Canker of black walnut and
butternut.
Sirococcus strobilinus. Shoot Blight of Picea, Abies, Pinus, and Tsuga
spp.
Sphaeropsis
Cankers.
Sphaeropsis sapinea. Shoot Blight of pine.
Dothiora (Sphaerulina)
Perithecia innate or nally erumpent, not beaked; paraphyses and paraphysoids lacking;
spores hyaline, several-celled.
Dothiora woli (formerly Sphaerulina polyspora). Twig Blight of sour-
wood, and oxydendron.
Dothiora taxicola (formerly Sphaerulina taxi). Needle Blight of yew.
Sphaerulina polyspora (see Dothiora woli). Twig Blight of sourwood,
and oxydendron.
Sphaerulina taxi (see Dothiora taxicola). Needle Blight of yew.
Sporidesmium
Conidiophores clustered, dark, short, simple, each bearing a terminal conidium; coni-
dia dark, quite large, muriform with many cells, oblong to ovoid; usually saprophytic,
sometimes parasitic.
Alternaria scorzonerae (formerly Sporodesmium scorzonerae). Salsify
Leaf Blight. Leaves have many circular ispots, varying from pin point to
1/4 inch, brown with red borders. Leaves or whole tops die; roots are small
BLIGHTS 215
and unsalable. The fungus winters as mycelium and spores in plant refuse.
May be the same as Alternaria tenuis.
Sporidesmium maclurae. Leaf Blight of osage-orange.
Sporodesmium scorzonerae (see Alternaria scorzonerae). Salsify Leaf
Blight. Leaves have many circular ispots, varying from pin point to 1/4
inch, brown with red borders.
Stemphylium
Leaf Spots.
Stemphylium vesicarium. Stemphylium Blight of onions. Lesions are
nondelineated, light yellow to brown, water-soaked and range in length from
one centimeter to the entire leaf.
Systremma
Asci in locules in an elongated stroma, which is erumpent and supercial at maturity;
spores light brown, two-celled. Conidial state Lecanosticta with brown conidia, two to
four cells, formed on a conidial stroma resembling an acervulus.
Mycosphaerella dearnessii (formerly Systremma acicola). Pine Brown
Spot Needle Blight, on southern pines, most serious on longleaf. The
name and classication of the fungus has been in dispute. The conidi-
al stage, known since 1876, was rst listed as Septoria, later placed in
Lecanosticta. The teleomorph state was named Scirrhia acicola in 1939 but
later transferred to Systremma because of its colored spores.
Most injurious on seedlings, needle blight may also injure large trees. Small,
gray-green spots on needles turn brown and form a narrow brown band,
the needle tips dying. Three successive seasons of brown spot kill longleaf
seedlings. The fungus is more severe on trees in unburned areas because
of accumulation of inoculum. Spray seedlings in plantations with bordeaux
mixture every 2 weeks from May to October or November.
Systremma acicola (see Mycosphaerella dearnessii). Pine Brown Spot
Needle Blight, on southern pines, most serious on longleaf.
Thelephora
Fruiting body leathery, upright, stalked; pileate or fan-shaped or much lobed, or in
an overlapping series; hymenium on the underside, smooth or slightly warty; spores
one-celled.
Thelephora spiculosa. Stem Blight found on azalea, fern, and other orna-
mentals in a Maryland garden. The fungus formed a dense weft of mycelium
on surface of the soil and on plants.
Thelephora terrestris. Seedling Blight, Smother. The mycelium rami-
es in the soil, and the leathery fruiting body grows up around the stem of
a seedling conifer or deciduous tree, smothering it or strangling it without
being actually parasitic on living tissue. The disease occurs most often in
crowded stands in nurseries. The damage is seldom important.
Tryblidiella
Ascomycetes, Patellariales
Apothecia opening by a wide cleft; spores dark, cylindrical, with several cells.
Rhytidhysteron rufulum(formerly Tryblidiella rufula). Twig Blight on cit-
rus.
Tryblidiella rufula (see Rhytidhysteron rufulum). Twig Blight on citrus.
Volutella
Sporodochia discoid, with marginal dark setae; conidiophores usually simple, in a com-
pact palisade; conidia hyaline, one-celled, ovoid to oblong; parasitic or saprophytic (see
Fig. 3.9).
Pseudonectria pachysandricola (see Volutella pachysandrae, Telemorph).
Pachysandra Leaf and Stem Blight. Large areas of leaves turn brown to
black, along with portions of stems, and in wet weather numerous pinkish
spore pustules appear along stems.
BLIGHTS 217
Volutella buxi. Boxwood Leaf Blight, Nectria Canker. Pinkish spore
occur as pustules on leaves and twigs. Leaves often turn straw-colored. See
further under Cankers.
Volutella pachysandrae (formerly, Pseudonectria pachysandricola). Pa-
chysandra Leaf and Stem Blight. Large areas of leaves turn brown to
black, along with portions of stems, and in wet weather numerous pink-
ish spore pustules appear along stems. The blight is most serious when
pachysandra has been injured or is too crowded or is kept too moist by tree
leaves falling into the bed. Spraying once or twice with bordeaux mixture
gives excellent control if severely blighted plants have been removed before
treatment. Keep pachysandra thinned and sheared back periodically.
BLOTCH DISEASES
Diseases designated as blotch have symptoms that are intermediate between
blights, where the entire leaf or shoot dies, and leaf spots, where the necrotic
lesions are denitely delimited. Blotches are irregular or indenite large or
small necrotic areas on leaves or fruit.
Alternaria
Blights.
Alternaria porri. Purple Blotch of onion, also on garlic, and shallot,
a problem in southern and irrigated areas. Small, white, circular to irregular
spots increase to large purplish blotches, sometimes surrounded by orange
and yellow bands, on leaves and ower stalks. Leaves often turn yellow
and die beyond the spots; girdled stalks die before seeds mature. Brown
muriform spores form a dusky layer on the blotches. Varieties with a waxy
foliage are more resistant than those with glossy leaves. The fungus winters
as mycelium and spores in crop refuse. Rotation, cleaning up plant debris,
and seed treatment are recommended.
Two other species of Alternaria, A. alternata and A. tenuissima, may cause
purple or brown blotches on onion, and there are physiological races as well.
Cercospora
Blights.
Cercospora concors (see Myrovellosiella concors). Potato Leaf Blotch.
An unimportant disease; leaets turn yellow with small blackened dead areas
or larger, irregular brown areas.
Cercospora purpurea (see Pseudocercospora purpurea). Avocado
Blotch, Cercospora Spot, considered the most important avocado disease
in Florida with no commercial variety entirely resistant.
BLOTCH DISEASES 219
Myrovellosiella concors (formerly Cercospora concors). Potato Leaf
Blotch. An unimportant disease; leaets turn yellow with small blackened
dead areas or larger, irregular brown areas.
Pseudocercospora purpurea (formerly Cercospora purpurea). Avocado
Blotch, Cercospora Spot, considered the most important avocado disease
in Florida with no commercial variety entirely resistant. Leaf spots are angu-
lar, brown to chocolate brown, scattered and distinct, less than 1/16 inch or
coalescing to larger patches. With a hand lens, grayish spore groups can be
seen on both sides of the leaf. Successive crops of spores are produced in
moist periods throughout the year. Fruit spots are 1/4 inch or less in diameter,
brown to dark brown, irregular, sunken, with cracked surfaces and grayish
spore tufts. Lesions are conned to the rind so that the esh is not affected,
but the cracks furnish entrance to anthracnose and other decay organisms.
The fungus winters in leaves, and appears to be progressively more abundant.
Cladosporium
Conidiophores dark, branched variously near upper or middle portion, clustered or sin-
gle; conidia dark, one- or two-celled, variable in size and shape, ovoid to cylindrical,
borne singly or in chains of two or three; parasitic or saprophytic.
Cladosporium herbarum. Leaf Blotch of lilac. The fungus is usually sec-
ondary, saprophytic, following blights.
Cladosporium paeoniae. Peony Leaf Blotch, Red Stem Spot, Mea-
sles. Leaf and stem spots are purplish or brownish red. On stems the spots
are raised, up to 4 mm long; on leaves the lesions are small specks. Small red-
dish spots are also present on oral bracts and petals. The disease is widely
distributed in commercial plantings and may sometimes destroy the value
of owers for cutting. Cut down tops in fall as for Botrytis blight. Spraying
the ground with Elgetol in spring before new growth starts has given good
control in some elds.
Geastrumia
Geastrumia polystigmatis. Sooty Blotch of Fruit on apple and blackber-
ry.
Gloeodes
Deuteromycetes, Sphaeropsidales, Leptostromataceae
Pycnidia dimidiate, having a radiate cover over the top half only, on a dark subicle or
mycelial crust; pseudoparaphyses present; conidia hyaline, one-celled.
Gloeodes pomigena. Sooty Blotch of Fruit on apple, crabapple, blackber-
ry, pear, and citrus, in eastern and central states down to the Gulf, rare in
the West. Fruit may be infected by heavy spore dissemination from pycni-
dia on twigs of various wild trees, including persimmon, prickly-ash, white
ash, bladdernut, hawthorn, red elm, sassafras, maple, sycamore, and willow.
On apples, clusters of short dark hyphae make a supercial thallus on the
cuticle, which appears as a sooty brown or black blotch, 1/4 inch in diame-
ter. Numerous spots may coalesce to cover the apple, a condition known as
cloudy fruit. Because the lesion is supercial the fruit esh is little affected,
but the grade and market value are reduced. On citrus the fungus does not
penetrate the rind, and spots can be removed by gentle hand rubbing. The
disease develops in cool rainy weather during the summer. To control open
up the trees in the orchards to facilitate quick drying.
Guignardia
Perithecia immersed in substratum, stroma lacking, mouths papillate; spores hyaline
unequally two-celled, with lower cell cut off just before maturity.
Guignardia aesculi. Horse-Chestnut Leaf Blotch, Buckeye Leaf
Blotch, general on horse-chestnut and Ohio buckeye, sometimes on red
and yellow buckeye. Large, reddish brown blotches in foliage are, usually,
surrounded by a yellowish area. Numerous pin-point black dots, pycnidia,
distinguish blotch from scorch due to drought. Petioles often have reddish
oval spots. In a rainy season there is a good deal of secondary infection from
spores spread by wind and rain. Blotches appear on nearly every leaet with
extensive defoliation. Primary infection in spring comes from ascospores
developed in fallen overwinter leaves.
Control. Rake up and burn leaves in fall. Feed trees that have been defoliated
for successive years.
BLOTCH DISEASES 221
Mycosphaerella
Blights.
Mycosphaerella dendroides (Cercospora halstedii, Anamorph). Pecan
Leaf Blotch, on pecan in the South, on hickory in East and South, a foliage
disease of nursery and orchard trees. Olive green velvety tufts of conid-
iophores and spores appear on undersurface of mature leaves in June and
July (in Florida), and yellow spots appear in corresponding areas on upper
leaf surfaces. Black pimplelike perithecia are produced in the tufts about
midsummer, united in groups to give the leaf a shiny black, blotched appear-
ance after the spores are washed away. In nursery trees, defoliation, starting
with basal leaves and progressing upward, may be serious. The disease is
of little consequence to orchard trees unless they have been weakened by
overcrowding, borer attack, or other cause. The fungus winters in fallen
leaves. To control clean up fallen leaves.
Mycosphaerella diospyri. Leaf Blotch of Japanese persimmon.
Mycosphaerella lythracearum (Cercospora punicae, Anamorph). Leaf
Blotch, Fruit Spot of pomegranate. The anamorph state has been thought
the same as that on crape-myrtle (Cercospora lythracearum), but is now
considered distinct. Leaf spots are circular, small, dark reddish brown to
almost black, sometimes grayish brown.
Phoma
Blackleg.
Phoma arachidicola. Web Blotch of peanut.
Phyllosticta
Blights.
Phyllosticta congesta. Leaf Blotch of garden plum.
Phyllosticta solitaria. Apple Blotch, widespread on apple and crabapple
in eastern states, serious in the South and in the Ozark section of Missouri,
Arkansas, Oklahoma, and Texas. The disease is also called fruit blotch, dry
rot, black scab, late scab, cancer, and tar blotch. From Kansas eastward it is
second in importance to apple scab. Leaf spots are very small, round, white,
with a single black pycnidium in the center of each. Larger elongate lesions
are formed on veins, midribs, and petioles. Leaves do not turn yellow, but
they drop prematurely if spots are numerous. Cankers on twigs and branches
are located at leaf nodes or base of spurs. The rst season they are small,
purple to olive in color; the next season this portion is tan and the new area
dark purple, often slightly raised. Pycnidia formed in twig lesions wash to
leaves, fruit, and new shoots, discharged only after heavy rains and in warm
weather. Heavily fertilized trees are more susceptible.
Fruit blotches are brown, irregular, feathery at the margin, studded with
numerous pycnidia. They afford entrance to secondary decay organisms and
may develop deep cracks, but the blotch fungus itself is supercial. It winters
in infected twigs and bark cankers.
Control. Secure healthy nursery stock. Some varieties, including Grimes
Golden, Jonathan, Stayman Winesap, and Winesap, are rather resistant.
Septoria
Blights.
Septoria agropyrina. Brown Leaf Blotch on wheat grasses.
Septoria elymi. Speckled Leaf Blotch on wheat grasses. A salt and pep-
per effect with numerous pycnidia in pale gray, tan, or fuscous lesions.
Septoria macropoda. Purple Leaf Blotch, general on blue grasses. Irreg-
ular blotches on blades are mottled greenish, then gray, tan or brown, nally
bleached nearly white. Pycnidia are round, attened, and light brown.
Zygophiala
A genus described from banana leaves in Jamaica.
Zygophiala jamaicensis. Greasy Blotch of carnation. A tropical fungus
found causing serious losses in California greenhouses in 1953 and reported
from Pennsylvania in 1957. Small, radiate patterns, resembling spider webs,
appear as if dipped in oil. Leaves become brittle, turn yellow, and die prema-
turely. The same fungus is present as a yspeck on apple.
BROOMRAPES
Broomrapes are parasitic seed plants like dodder and mistletoe. They are
leaess herbs, of the family Orobanchaceae, living on roots of other plants
and arising from them in clumps of whitish, yellowish, brownish, or purplish
stems. There are 130 or more species, mostly from North Temperate regions,
but few have any garden importance. The seed germinates in soil and pro-
duces a liform plant body that grows into the ground penetrating crown or
root of the host plant and forming a more or less tuberous enlargement, from
which the owering shoots arise. Such shoots may be nearly naked, clothed
only with a few scattered rudimentary leaves, or they may be covered with
conspicuous, overlapping scalelike leaves. The seed may remain viable in
the soil several years but probably not as long as has been believed, for they
can live on some weeds between crops.
Orobanche ludoviciana. Louisiana Broomrape on tomato and other
plants, including Spanish needle and coldenia, becoming a problem in Cali-
fornia. Tomatoes are stunted and do not produce a full crop of fruit.
Orobanche ramosa. Branched Broomrapeh, Hemp Broomrape, most
serious on hemp but parasitizing tomatoes, lettuce, tobacco, eggplant, Gan-
ra, Melitlotus, Silene, poppy mallow, cranesbil, Chaerophyllum, Verbena,
Coreopsis, eabank, engelmann daisy, and other hosts in California. In small
infections destroy the aerial stems before they set seed; practice crop rota-
tion. Deep plowing gives some control.
CANKERS AND DIEBACKS
A canker is a localized lesion or diseased area often resulting in an open
wound and usually on a woody structure. Starting as a denite necrotic spot,
it may girdle cane, stem, or tree trunk, killing the water-conducting tissues
so that the most prominent symptom becomes a dieback. When twigs and
branches die back from the tip, the condition may be a blight, with the
pathogen directly invading the dying area, or it may be a secondary effect
from a canker some distance below.
Aleurodiscus
Hymenium resupinate, of one layer, with projecting spinose or short-branching cystidia
(swollen sterile cells); spores hyaline. Facultative parasite on trees.
Aleurodiscus acerina (see Dendrothele acerina). Bark Patch, widespread
on maple.
Aleurodiscus amorphus. Balsam Fir Canker. Cankers are formed on
main stems of saplings, which are sometimes killed, but the fungus is
also widespread as a saprophyte on dead bark of rs and other conifers.
Cankers center around a dead branch, are narrowly elliptical with a raised
border; the dead bark is covered with a light-colored layer of the fun-
gus.
Aleurodiscus oakesii. Oak Bark Patch, Smooth Patch of white oak.
Irregularly circular, smooth, light gray sunken areas in bark vary from sever-
al inches to a foot across. The fungus is conned to dead bark; trees are not
injured.
Dendrothele acerina (formerly Aleurodiscus acerina). Bark Patch, wide-
spread on maple.
CANKERS AND DIEBACKS 225
Amphobotrys
Conidiophores are long, slender, pigmented, and highly branched; clusters of conidia at
apex of each branch; conidia ovoid, one-celled, hyaline.
Amphobotrys ricini. Stem Canker on texasweed and castorbean. Girdling
stem canker; of prostrate spurge.
Apioporthe
Perithecia in a black, carbonaceous stroma; spores two-celled, hyaline; conidia in cavi-
ties in a stroma.
Anisogramma anomala (formerly Apioporthe anomala). Canker, Twig
Blight of hazelnut.
Apioporthe anomala (see Anisogramma anomala). Canker, Twig Blight
of hazelnut.
Apioporthe apiospora. Twig Canker, Dieback of elm.
Ascospora
Ascomycetes, Sphaeriales, Sphaeriaceae
Perithecia with a subicle; paraphyses lacking; spores two-celled, hyaline.
Ascospora ruborum (Hendersonia rubi, Anamorph). Cane Spot, Die-
back of red and black raspberry, dewberry.
Atropellis
Apothecia black, sessile or with short stalk; asci clavate, with longer, hairlike paraphy-
ses; spores needlelike to slightly club-shaped, hyaline, one-celled.
Atropellis apiculata. Twig Canker. On southern pines.
Atropellis arizonica. Branch and Truck Canker. On western yellow pine.
Atropellis pinicola (syn. A. piniphila). Pine Branch and Trunk Canker,
on western white, sugar, and lodgepole pines in Pacic Northwest and Cal-
ifornia. Branches are girdled and killed, but not the trees. Perennial cankers
are smooth, elongated, attened depressions covered with bark, in which
appear very small black apothecia, 2 to 4 mm in diameter.
Atropellis piniphila (Cenangium piniphilum, Anamorph). Branch and
Truck Canker on lodgepole and ponderosa pines on Pacic Coast, on
cultivated pines in the South. Trees 5 to 25 years old are damaged by defor-
mation of main stem and branches. Infection is at branch whorls. Cankers
are elongated, attened depressions covered with bark and copious resin.
Apothecia have short stalks, are black with brownish discs, 2 to 5 mm
across.
Atropellis tingens. Branch and Truck Canker of native and exotic hard
pines from New England and Lake states to Gulf states. Slash pine saplings
are most susceptible. Smaller branches are girdled; perennial target cankers
are formed on larger branches and main stems. Cankers persist for many
years, but extension stops after about 10 years.
Botryodiplodia
Blights.
Botryodiplodia gallae (see Lasiodiplodia theobromae). Canker of oak.
Botryodiplodia theobromae. Canker of rose, and citrus.
Lasiodiplodia theobromae (formerly Botryodiplodia gallae). Canker of
oak.
Botryosphaeria
Blights.
Botryosphaeria dothidea. Canker, Gummosis, and Dieback on peach,
Bradford pear, thornless blackberry, sequoiadendron and sequoia.
Botryosphaeria obtusa. Canker, on thornless blackberry.
Botryosphaeria ribis. Saprophytic on dying tissue, and var. chromogena,
parasitic. Canker, Dieback of at least 50 woody plants, including apple,
avocado, eucalyptus, g, forsythia, hickory, pecan, pyracantha, quince,
rhododendron, sequoia, sequoiadendron, sweet gum, and willow. See under
Blights for the disease caused on currant and rose, under Rots for apple and
avocado diseases.
On redbud, sunken oval cankers nearly girdle branches, the fungus entering
through wounds, and dead and dying twigs. On rhododendron there is a leaf
spot and dieback similar to that caused by Phytophthora except that the
surface is roughened by protruding fruit bodies. Cankers on twigs, larger
branches, and trunks of willow may kill trees in a few years. Trunk lesions
are very small, 1/4 to 1/2 inch, and numerous or else large, from the union
of several small cankers, with ssured bark. Apples have watery blisters on
bark and decline in vigor. Forsythia has affected canes girdled and killed
with conspicuous brown dead leaves above the canker.
Control. Prune and burn dead twigs and heavily infected branches; paint
wounds with a disinfectant followed by tree paint; avoid injuries. Copper
sprays may help.
Botryosphaeria stevensii. Canker, on juniper.
Botrytis
Blights.
Botrytis cinerea. Canker of rose.
Caliciopsis
Ascomycetes, Coryneliales
Stroma lobed, each lobe containing a single locule, which is nally wide open; perithecia
stalked; asci on long slender stalks; spores dark, one-celled.
Caliciopsis pinea. Pine Canker on eastern white pine and other species,
also on Douglas r. Cankers are sharply depressed areas in bark, reddish
brown and smoother than rest of bark, up to several inches in diameter.
Small, globose, clustered black pycnidia, and stalked perithecia looking like
slender black bristles, arise from stroma in cankered bark. The disease is
most serious on suppressed saplings.
Encoelia (Cenangium)
Blights.
Cenangium singulare (see Encoelia pruinosa). Sooty-Bark Canker of
aspen, on Populus tremuloides in Rocky Mountain area.
Encoelia pruinosa (formerly Cenangium singulare). Sooty-Bark Canker
of aspen, on Populus tremuloides in Rocky Mountain area. Cankers on older
trees, at any point on trunk up to 60 to 70 feet may extend 10 to 15 feet
before they girdle the tree. The bark is sooty black with a thin white outer
layer.
Ceratocystis (Ceratostomella)
Ascomycetes, Micrascales
Perithecia with very long beaks, carbonaceous or leathery; ascospores hyaline, one-
celled; brown, ovoid conidia and one-celled rodlike endospores formed inside tubelike
conidiophores and extruded endwise. Some species are important tree pathogens; see
Oak Wilt and Dutch Elm Disease under Wilts.
Ceratocystis mbriata f. sp. platani (Endoconidiophora mbriata f. sp.
platani). Canker Stain of London Plane, Plane Blight, on London plane
and also on American plane or sycamore. This serious disease started as
a killing epidemic in the Philadelphia area about 1935, destroying city shade
trees by the thousands there and in Baltimore during the next few years. The
disease now extends from New Jersey to North Carolina and Mississippi.
Trees show sparse foliage, smaller leaves, and elongated sunken cankers on
trunks and larger branches. Cross sections through cankers reveal blue black
or reddish brown discoloration of wood, usually in wedge-shaped sectors.
First year cankers may not be more than 2 inches wide and a yard or so long,
but they widen annually, girdling and killing trees in 3 to 5 years. Several
cankers coalescing around the trunk kill more quickly. Once infection starts,
the tree is doomed.
Ascospores and the two types of conidia are produced in moist spring weath-
er (see Fig. 3.15). They may be spread by rain a short distance, but most
dissemination is by man in pruning operations, and ordinary tree paint car-
Figure 3.15 Spore Formation of Some Canker Fungi. Coniothyrium, small dark spores on short conidiophores in
pycnidium; Seiridium, formerly Coryneum), dark, septate spores in acervulus; Cytospora, sausage-shaped spores
in valsoid pycnidia expelled in cirrhi; Endoconidiophora, spores formed on inside of conidiophores; Nectria, two-
celled bright ascospores in reddish perithecia clustered on bark; Phoma, hyaline spores in pyncnidia formed in
spots on bark
ry viable spores. Some beetles may be vectors. Infection is solely through
wounds.
Control. Do not try to save trees where trunk has been invaded; diseased
branches may sometimes be removed, cutting at least 3 feet from infected
area. Do not prune unless absolutely necessary and then only in winter when
trees are less susceptible. Use tree wound dressing fortied with a disinfec-
tant.
Ceratocystis sp. Canker and Dieback on poplar.
Chondropodium
Pycnidia stromatic, stalked, columnar, externally black, hard, internally gelatinous; coni-
diophores simple; conidia hyaline, with several cells, crescent- or sickle-shaped; weakly
parasitic or saprophytic.
Chondropodium pseudotsugae. Bark Canker of Douglas-r. This is
a supercial canker with outer layers of bark killed over small, circular to
elliptical areas, in which pycnidia project as short, blunt, black spines. Trees
are not noticeably injured.
Colletotrichum
Anthracnose.
Colletotrichum acutatum. Canker and Dieback on Japanese maple.
Coniothyrium
Pycnidia black, globose, separate, erumpent, ostiolate; conidiophores short, sim-
ple; conidia small, dark, one-celled, ovoid or ellipsoid; parasitic or saprophytic (see
Fig. 3.15).
Coniothyrium fuckelii (Anamorph, Diapleela coniothyrium). Rose Com-
mom Canker, Stem Canker, widespread on rose, also causing raspberry
cane blight (see Leptosphaeria under Blights), sometimes associated with
apple rots, peach cankers, and stem canker of Virginia creeper. Of the three
species of Coniothyrium that cause rose cankers, C. fuckelii is by far the
most common. Any plant part may be affected. Pycnidia have even been
found within blackspot lesions on leaves, but this is primarily a cane disease,
starting as a red or yellow spot on bark, drying out and turning brown as it
increases in size, with the epidermis somewhat wrinkled and perhaps rup-
turing irregularly over sooty masses of very small, olive brown spores. The
stem may be girdled with dieback to that point.
Stem cankers are found around insect punctures, thorn pricks, leaf or thorn
scars, or abrasions caused by tying, but the majority of cankers are formed
at the cut end of a cane when a stub has been left in pruning above a leaf
axil or bud. Roses cut properly close to a bud seldom develop this canker.
A rose stub usually dies back to the rst node, and since this fungus is a weak
parasite, it starts most readily in such dead or dying tissue. When a cut is
made close to the node, it is quickly callused over, and the callus is a good
defense against wound fungi.
Control. Prune out cankered and dying stems as soon as noticed. Make all
cuts just above a bud or leaf axil, not only at spring pruning but in cutting
owers for the house or cutting off dead blooms during the season.
Coniothyrium rosarum. Rose Graft Canker. This is a disease of roses
under glass, starting at the union of stock and scion in the warm moist prop-
agating frame and continuing in a large amount of dead wood when plants
are removed to the greenhouse bench. Some consider the pathogen a form
of C. fuckelii. Having measured spores of the type specimen, in the Kew
Herbarium, I think they are distinct species, but that some cases of graft
canker are due to the common canker fungus.
Coniothyrium wernsdorfae. Rose Brand Canker, a rather rare but very
serious disease. The pathogen was named in Germany in 1905 and was not
reported in this country until 1925, although it was subsequently shown to
have been collected in Canada in 1912 and in Pennsylvania and Minnesota in
1914 and 1916. In 1926 a severe epiphytotic appeared at Ithaca, New York,
in the Cornell rose garden, infecting about 90% of the climbers so seriously
that the canes had to be cut to the ground. Since then it has been reported
from a few other states, but in several instances it has been confused with
common canker.
Small, dark reddish spots on canes enlarge and acquire a more or less denite
reddish brown or purple margin, contrasting sharply with the green of the
cane. The center of the spot turns light brown as the cells die, and little lon-
gitudinal slits appear over the developing pycnidia. Spores are olive brown,
nearly twice the size of C. fuckelii, and released through epidermal slits
instead of being spread in a sooty mass under the epidermis. Cankers formed
under the winter protection of soil are black when roses are rst uncovered
in spring, which explains the name Brandeckenkrankheit, meaning re-spot
disease.
C. wernsdorfae is a cold temperature fungus, infecting rose canes under
the winter covering, entering through insect wounds, thorn scars, scratch-
es, and occasionally through dormant buds. During a 4-year investiga-
tion at Ithaca, I found no infection on canes not hilled with earth or
other moist cover over winter and no natural infection during the sum-
mer.
Control. Omit the usual winter protection of soil or other materials that keep
canes moist. If brand canker is a problem, just fasten canes of climbers
down near the ground, uncovered, and hope for the best. Loss from win-
ter injury will be less than from the canker. Cut out diseased canes careful-
ly.
Seiridium (Coryneum)
Blights.
Coryneum cardinale (see Seiridium cardinale (Leptosphaeria sp., Tele-
morph)). Coryneum Canker of Cypress, Bark Canker of cypress, incense
ceder, common juniper and oriental arborvitae.
Coryneum foliicola. Twig Canker, Fruit Rot, widespread on apple, affect-
ing twigs, foliage and fruit.
Seiridium cardinale (formerly Coryneum cardinale (Leptosphaeria sp.,
Telemorph)). Coryneum Canker of Cypress, Bark Canker of cypress,
incense ceder, common juniper and oriental arborvitae. This disease, since
its discovery in 1927, has been gradually exterminating Monterey Cypress
in most parts of California and is also serious on Italian cypress. Twigs,
branches, and whole trees turn sickly, lose their leaves, and nally die.
The fungus attacks living bark and cambium, girdling twig and branch.
Cankers appear rst at base of lateral twigs; they are slightly sunken, dark,
resinous, rough, with black spore pustules. Conidia have dark median cells,
ve cross-walls (see Fig. 3.15). They are spread by tools, in nursery stock,
by wind and rain, and perhaps by birds and insects. Infection appears rst
in upper parts of trees, usually in spring during moist weather. Yellowing
and browning of foliage together with gummy ooze at the cankers form
conspicuous symptoms.
Control. Drastic surgery, removing wood well below infected parts, and
spraying foliage heavily with bordeaux mixture help some, but with heavy
infection the price of saving healthy trees is the removal and destruction of
all diseased specimens. California citizens, threatened with the loss of the
famous native stands of Monterey cypress at Point Lobos and Cypress Point,
voluntarily destroyed their own plantings by the thousands.
Cryphonectria
Production of perithecial ascocarps produced in a stroma of fungal and substrate tissues
or directly from somatic hyphae on the substrate. Ascospores are hyaline to brown and
one-to-several-septate.
Cryphonectria parasitica. Canker on oak.
Cryptodiaporthe
Ascomycetes, Sphaeriales, Valsaceae
Like Diaporthe but without blackened zones in substratum; spores hyaline, two-celled.
Amphiporthe aculeans (formerly Cryptodiaporthe aculeans; Sporocybe
rhois, Anamorph). Dieback, Canker of sumac.
Amphiporthe castanae (formerly Cryptodiaporthe castanea). Dieback,
Canker of Asiatic Chestnut, widespread, chiey on seedlings or on larger
trees in poor sites. Canker starts as a brown discoloration of bark of the
trunk, limb, or twig, often girdling twig and then invading larger branch.
Leaves on girdled branches wilt without yellowing, turn brown, and die.
Bark splitting over callus formation at edge of diseased area forms pro-
nounced canker. Conidia, two-celled, fusoid, are formed in pustules in bark;
beaked perithecia are formed in groups by midsummer.
Control. Maintain vigor; plant on well-drained, fertile soil. Prune out dis-
eased portions several inches below affected area.
Crytodiaporthe aculeans, Sporocybe rhois, Anamorph (see Amphiporthe
aculeans). Dieback, Canker of sumac.
Cryptodiaporthe castanea (see Amphiporthe castanae). Dieback, Canker
of Asiatic Chestnut, widespread, chiey on seedlings or on larger trees in
poor sites.
Cryptodiaporthe salicella. Twig and Branch Canker of willow.
Cryptomyces
Apothecia effuse, splitting irregularly; paraphyses present; spores hyaline, one-celled.
Cryptomyces maximus. Blister Canker on common and purple osier.
Cryptosporella
Perithecia in a circle in a stroma, with long necks converging in a common canal; spores
one-celled, hyaline; conidia borne on surface of stroma.
Cryptosporella umbrina. Rose Brown Canker, a widespread and serious
rose disease, rst reported in Virginia in 1917 but known from herbarium
specimens to have been present since 1903. The fungus was rst placed in
Diaporthe because of occasional two-celled spores.
Symptoms are most noticeable on canes, starting with very small purplish
spots, the center soon turning white with a reddish purple margin (see
Fig. 3.16). Many small spots may be grouped on a single cane. During the
winter, and especially on portions of canes covered with earth, cankers or
girdling lesions are formed, often several inches long, with tan centers and
purplish borders. In moist weather the surface of these large cankers is cov-
ered with yellow spore tendrils from pycnidia just under the bark; asci are
also extruded in tendrils from perithecia.
Leaf spots are small purplish specks or larger dead areas, cinnamon buff to
white, bordered with purple and with black pycnidia in the center. Marginal
spots are subcircular. Buds are sometimes blighted; exposed petals of ow-
ers have cinnamon-buff spots without the purple border. Infection is through
wounds and also uninjured tissue.
Control. The best time to take care of brown canker is at spring pruning. Cut
out every diseased cane possible. A dormant lime sulfur spray, immediately
after pruning, kills spores that may have been spread in the process and may
inhibit the fungus in initial lesions. Copper or sulfur sprays largely prevent
summer infections. Brown canker is more likely to be serious where roses
are overprotected for winter with salt, hay, leaves, or other material added to
the mound of soil. I have no trouble with brown canker when roses are left
unhilled over winter.
Cryptosporella viticola. Dead-Arm Disease of grapes, Branch Necro-
sis, widespread, especially in the Northeast, serious in Illinois, important
in California. Small, angular spots with yellowish margins and dark centers
are formed on leaves, stems of ower clusters and canes. The latter may
split to diamond-shaped cankers, and by the next season the arm is dead or
producing yellowed, dwarfed and crimped foliage. Lesions on cluster stems
advance into fruit late in the season causing rotting. Pycnidia are developed
on old wood; infection is often through pruning wounds.
Control. Make pruning cuts at least 6 inches below the lower margin of the
infected part. Spray with bordeaux mixture when spores are extruded.
Figure 3.16 Brown Canker on Rose
Cryptosporium
Acervuli erumpent, becoming cup-shaped or disclike; stroma brownish; conidiophores
simple or branched; conidia hyaline or subhyaline, one-celled liform.
Cryptosporium minimum. Canker on rose, not common.
Cryptosporium pinicola (see Gelatirosporium piricola). Canker, Branch
Mortality of Abies spp.
Gelatirosporium piricola (formerly Cryptosporium pinicola). Canker,
Branch Mortality of Abies spp.
Cylindrocarpon
Rots.
Cylindrocarpon didymium. Bole Canker on apple.
Cylindrocarpon cylindroides. Canker, Branch Mortality of Abies spp.
Cylindrocladium
Blights.
Cylindrocladium scoparium. Crown Canker of rose. The cane is attacked
at or just below the union of stock and scion, the bark darkening into a black,
water-soaked punky region. The cankers girdle but do not kill the canes; there
are fewer and more inferior blooms. The disease was long thought conned
to greenhouse roses but has appeared once or twice in outdoors elds. The
fungus lives in the soil and enters through wounds in the presence of suf-
cient moisture. Before planting of fresh stock, greenhouse benches should
be washed with boiling water and soil sterilized or changed.
The same fungus injures seedling conifers in nursery rows, causing damping-
off, root rot, stem canker and needle blight to white pine and Douglas-r.
See under Blights for a discussion of the pathogen on cuttings of azaleas and
other ornamentals.
Cytospora
Cosmopolitan species, anamorph state of Valsa. Pycnidia in a valsoid stroma with irreg-
ular cavities, incompletely separated; conidia hyaline, one-celled, allantoid, expelled in
cirrhi (see Fig. 3.15).
Cytospora abietis. Canker, Branch Mortality of Abies spp.
Cytospora pruinosa. Canker, Dieback of ash, on twigs and branches.
Cytospora chrysosperma (Teleomorph, Valsa sordida). Cytospora Can-
ker of poplar, aspen, cottonwood, willow, occasional on mountain-ash,
maple, cherry, and elder. Cankers form on trunks and large branches, most
often on trees of low vigor. Bark is discolored in more or less circular areas;
sapwood is reddish brown. In old cankers exposed wood is surrounded by
layers of callus tissue. In moist weather spring spore tendrils are extrud-
ed from pycnidia in dead bark. Perithecia are found infrequently in aspen,
arranged circularly around a grayish disc; they are ask-shaped with long
necks pushing through the bark. Twigs and small branches may die back
without a denite canker. The fungus is often present on healthy trees, not
becoming pathogenic until the trees are weakened by neglect, drought, pol-
larding or other causes. Entrance is through wounds. Lombardy and Simon
poplars are frequently killed.
Control. Remove dead and dying branches and trees with extensive cankers.
Avoid wounds; feed and water as necessary. Plant poplars that are less sus-
ceptible than Lombardy. Rio Grande cottonwood is resistant to twig blight.
Cytospora kunzei (Teleomorph, Leucostoma kunzei). Cutospora Canker
of spruce. Twig Blight, common and serious New England to the Mid-
west. Cankers start around bases of small twigs or on trunks. Browning and
death of Colorado blue spruce branches starts near the ground and progress-
es upward, a large ow of resin on affected limbs. Needles drop immediately
or persist for a time. Cankers are formed near resin spots and yellow tendrils
extruded. Spores are splashed by rain and wind to other branches; infection
is mostly through wounds.
Another form of the pathogen, Valsa kunzei var. supercialis, occurs on pine
and variety kunzei on balsam r, Douglas-r, larch and hemlock.
Control. There are no satisfactory control measures except removal of dis-
eased branches and perhaps carefully excising cankered bark. Spraying
with bordeaux mixture has been recommended but is seldom very effective.
Avoid wounding ornamental trees with lawn mowers; sterilize pruning tools
between cuts; feed to renew vigor.
Cytospora leucostoma. Canker of black cherry.
Cytospora nivea. Canker, Dieback of poplar and willow, similar to that
caused by C. chrysosperma; occasional.
Cytospora leucosperma. Branch Canker of elder.
Cytospora sp. Canker on alder and pecan.
Cytospora spp. Cytospora Canker of Italian Prunce, causing severe injury
to prune and apricot in Idaho orchards since 1951, also present on cherries,
peach, apple and willows. Some orchards have been lost, others hard hit.
Symptoms are yellow to brown ags of dead leaves and erumpent, gummy
cankers or elongated necrotic streaks in the bark. All suspicious wood should
be cut out, hauled out of the orchard, and burned.
Valsa cincta for further discussion of cankers on stone fruits.

Dasyscyphus
Ascomycetes, Helotiales, Helotiaceae
Apothecia stalked, white and hairy on the outside with a bright disc; paraphyses liform;
asci inoperculate; spores elliptical to fusoid.
Dasyscyphus agassizi (see Lachnellula agassizii). Common on blister-rust
lesions of white pine; saprophytic on dead branches.
Dasyscyphus calycina (Trichoscyphella hahniana) (see Lachnellula sub-
tilissima). On larch and r, ordinarily a saprophyte but can be a weak para-
site; occasional on blister-rust cankers.
Dasyscyphus ellisiana. Canker of Douglas-r and pine in eastern United
States. This is a native fungus on twigs and branches of native and introduced
pines and on basal trunk and branches of Douglas-r. Bark on trunk may be
infected for 10 to 15 feet, with copious resin ow and numerous swellings,
but trees are not killed. Apothecia are short-stalked, covered with white hairs,
with an orange to yellow disc, 2 to 4 mm across. Remove trees with trunk
cankers.
Dasyscyphus pseudotsugae (see Lachnellula pseudotsuga). Canker on
Douglas-r. Swollen open cankers, 2 to 3 inches long, are formed on sup-
pressed saplings.
Dasyscyphus resinaria (see Lachnellula resinaria). Canker on balsam r.
Swollen cankers at base of branches; younger stems girdled and killed.
Dasyscyphus willkommii (see Lachnellula willkommii (Trichoscyphel-
la willkommii syn. Lachnellula wilkommii)). European Larch Canker.
Found in Massachusetts in 1927 on nursery stock from Great Britain.
Lachnellula agassizi (formerly Dasyscyphus agassizi). Common on blister-
rust lesions of white pine; saprophytic on dead branches.
Lachnellula pseudotsuga (formerly Dasyscyphus pseudotsugae). Canker
on Douglas-r. Swollen open cankers, 2 to 3 inches long, are formed on
suppressed saplings.
Lachnellula resinaria (formerly Dasyscyphus resinaria). Canker on bal-
sam r. Swollen cankers at base of branches; younger stems girdled and
killed.
Lachnellula subtilissima (formerly Dasyscyphus calycina (Trichoscyphel-
la hahniana)). On larch and r, ordinarily a saprophyte but can be a weak
parasite; occasional on blister-rust cankers.
Lachnellula willkommii (formerly Dasyscyphus willkommii (Trichoscy-
phella willkommii syn. Lachnellula wilkommii)). European Larch Canker.
Found in Massachusetts in 1927 on nursery stock from Great Britain.
Infected trees were removed and the fungus not seen again until 1935,
near the original location. Perennial branch or trunk cankers are attened
depressions, swollen on the anks and on the opposite side of the stem.
Neighboring bark is somewhat cracked and dark with heavy exudation of
resin. Cup-shaped apothecia are 3 to 6 mm across with white hairs and
orange to buff discs, very short stalks. Young trees may be killed; older
trees usually survive. Frost wounds are a contributing but not an essen-
tial factor. Promptly remove all trees showing cankers; continue periodic
inspection.
Dermea (Dermatea)
Apothecia small, brownish to black with a circular opening; innate at rst, on a stromoid
base, rupturing host at maturity; spores one-celled, hyaline, globose to oblong. Cup fungi
(ascocarp cup-shaped); excipulum of subglobose cells; sclerotia absent.
Dermatea acerina (see Dermea acerina). Bark Canker of maple, occa-
sional.
Dermatea balsamea (see Dermea balsamea). Twig Canker of hemlock.
Dermatea livida (see Pezicula livida). Bark Canker of redwood.
Dermea acerina (formerly Dermatea acerina). Bark Canker of maple,
occasional.
Dermea balsamea (formerly Dermatea balsamea). Twig Canker of hem-
lock.
Dermea pseudotsugae. Branch Canker on r.
Pezicola livida (formerly Dermatea livida). Bark Canker of redwood.
Diaporthe
Blights.
Cryphonectria cubensis (formerly Diaporthe cubensis). Canker of Euca-
lyptus spp.
Diaporthe eres. Canker, Dieback of English holly in the Northwest. The
fungus name is a species complex that may include a Diaporthe on rose
petals and one causing a peach constriction disease.
Diaporthe cubensis (see Cryphonectria cubensis). Canker of Eucalyptus
spp.
Diaporthe eres. Canker, Dieback of English holly in the Northwest.
Diaporthe helianthi. Canker of sunower; also leaf spot of sunower.
Diaporthe oncostoma. Canker, Dieback of black locust.
Diaporthe phaseolorum var. caulivora. Canker of painted spurge, prickly
sida, redweed, morning-glory, black nightshade, jacquemontia, hemp sesba-
nia, indigo, spiny amaranth, vetch and soybean.
Diaporthe pruni. Twig Canker on black cherry; D. prunicola on American
plum.
Dichotomophthora
Conidiophores brown, branching dichotomous to subdichotomous, elongated, terminal
branches 48 lobed each lobe bearing single conidium; conidia dark, ovoid to elon-
gateovoid, 1 to 6 celled.
Dichotomophthora portulacae. Stem Canker and Root Rot on common
purslane.
Pseudomassaria (Didymella)
Blights.
Didymella sepincoliformis (see Pseudomassaria sepincolaeformis). Die-
back of rose.
Pseudomassaria sepincolaeformis (formerly Didymella sepincoliformis).
Dieback of rose.
Diplodia
Blights.
Diplodia sp. Rose Dieback, sometimes after drought and other contribut-
ing factors. In Texas the disease is most evident in autumn, progressing on
roses in storage or overwintering in the ground. Canes die from tip down-
ward, often starting in the ower stem. Diseased wood turns brown or black,
and is somewhat shriveled. Pycnidia are produced in dead canes. Improve
general rose vigor; use fungicides as for blackspot. May also cause canker
of Russian olive.
Diplodia camphorae. Canker, Dieback of camphor-tree.
Diplodia infuscans (see Sphaeropsis hyalina). Ash Canker and Dieback,
northeastern states.
Diplodia juglandis. Dieback, widespread on branches of walnut.
Diplodia mutila. Stem Canker on Laburnum. Branch Dieback on
juniper.
Diplodia natalensis (see Lasiodiplodia theobromae). Stem Canker of
prickly-ash; Dieback of citrus twigs, also causing citrus stem-end rot.
Diplodia quercina. Canker and Blight of oaks.
Diplodia sophorae. Dieback of pagoda tree.
Diplodia sycina. Canker, Dieback of g.
Lasiodiplodia theobromae (formerly Diplodia natalensis). Stem Canker
of prickly-ash; Dieback of citrus twigs, also causing citrus stem-end rot.
Rots.
Sphaeropsis hyalina (formerly Diplodia infuscans). Ash Canker and
Dieback, northeastern states.
Discella
Pycnidia cupulate or discoid; spores 2-celled, hyaline.
Discella carbonacea (see Discella microsperma). Twig Canker of willow.
Discella microsperma (formerly Discella carbonacea). Twig Canker of
willow.
Dothichiza
Pycnidia innate, nally erumpent; conidiophores lacking; conidia hyaline, one-celled.
Discosporium populeum (formerly Dothichiza populea). Dothichiza
Canker of poplar; European Poplar Canker, widespread but sporadic
as a branch and trunk canker. Lombardy poplars are most susceptible, but
hosts include black and eastern cottonwoods, balsam, black and Norway
poplars. Japanese poplars are rather resistant. Young trees in nurseries are
most injured, cankers often starting around wounds. They start as slightly
darker, sunken areas, often at base of twigs and limbs, and become elon-
gated. The bark is killed to the cambium; sapwood is brown. If a stem is
completely girdled, it dies; otherwise, callus formation goes on through the
summer, over the canker. In time diseased bark turns brown and cracks.
Spores are extruded in amber tendrils, drying to brown, and are washed to
wounds in the wood.
Control. Destroy infected stock in nurseries and plantations; do not move
stock from a nursery where the disease is known. Avoid pruning and other
wounds so far as possible; sterilize tools between cuts. Spraying nursery
trees with bordeaux mixture in spring may be helpful.
Dothichiza populea (see Discosporium populeum). Dothichiza Canker
of poplar; European Poplar Canker, widespread but sporadic as a branch
and trunk canker.
Dothiora
Ascomycetes, Pseudosphaeriales
Ascocarps hairy and phragonosporous or muriform ascospores are colored.
Dothiora polyspora (see Sydowia dothideoides). Canker of aspen.
Sydowia dothideoides (formerly Dothiora polyspora). Canker of aspen.
Dothiorella
Pycnidia dark, globose, grouped in a subcortical stroma; conidiophores simple, short;
conidia hyaline, one-celled, ovoid to ellipsoid; parasitic or saprophytic on wood.
Botryodiplodia gallae (formerly Dothiorella quercina). Dothiorella Can-
ker of oak, very destructive to red and white oaks in Illinois, affecting
twigs, branches, and occasionally trunks. Cankers are dark brown, elongat-
ed, sunken, often with cracks at the margin. Pustules of pycnidia develop in
bark and erupt through cracks, spores oozing on the surface. Sapwood has
dark streaks.
Dothiorella fraxinicola. Branch Canker of ash.
Dothiorella quercina (see Botryodiplodia gallae). Dothiorella Canker
of oak, very destructive to red and white oaks in Illinois, affecting twigs,
branches, and occasionally trunks.
Dothiorella sp. London Plane Canker, rst noted in New York City in
1947. Infected trees have sparse, undersized foliage and narrow, longitudinal
cankers on trunk and branches, varying from 1 to 4 inches wide and often
extending from ground level to branch top. The bark is rough, deeply s-
sured; inner bark is brown, dry; sapwood is only supercially discolored.
Branches wilt and die back.
Dothiorella ulmi. Dieback, Wilt of elm.
Wilts.
Endothia
Blights.
Endothia gyrosa. Branch Canker on oak.
Epicoccum
Leaf Spots.
Epicoccum nigrum. Canker on thornless blackberry.
Eutypa
Ascomycetes, Xylariales, Diatrypaceae
Stroma effuse; perithecia with necks at right angles to surface.
Eutypa armeniacae syn. E. lata. Cytosporina Dieback of apricot and
of grape; Twig Canker on cherry and chokecherry. Anamorph state report-
ed from California in 1962, perithecia in 1965. Bark cankers with gum are
formed at pruning wounds.
Fusarium
Rots.
Fusarium moniliforme var. subglutinans (see Fusarium subglutinans).
Pitch (Branch) Cankers and Shoot Dieback on southern pine species,
loblolly and pond pines.
Fusarium oxysporum. Stem Canker on peanut.
Fusarium solani. Stem Canker of sweetpotato, black walnut, oak, and
poinsettia.
Fusarium subglutinans (formerly Fusarium moniliforme var. subgluti-
nans). Pitch (Branch) Cankers and Shoot Dieback on southern pine
species, loblolly and pond pines.
Fusicoccum
Pycnidia one to several in a stroma, spherical or attened, subepidermal, erumpent;
opening separately or with a common pore; conidiophores simple, short; conidia hya-
line, one-celled, fusoid; parasitic or saprophytic.
Fusicoccum amygdali. Twig Canker of peach, increasingly important on
peaches in North Atlantic coastal area. Leaf spots are large, irregular or cir-
cular, often zonate, brown with scattered pycnidia near center. Cankers at
buds and bases of young twigs result in death of the distal portions; trunks
of young trees may be girdled. Infections occur throughout the season at bud
scales, stipules, fruit and leaf scars. Prune only in winter.
Fusicoccum elaeagni. Canker on Russian-olive.
Gibberella
Blights.
Gibberella baccata. Twig Canker of acacia, ailanthus, apple, boxwood,
mimosa, mulberry, and also on other plants where twig blight is the most
important symptom.

Blights.
Gloeosporium
Anthracnose.
Gloeosporium sp. Canker on holly.
Gloeosporium sp. (Gnomonia rubi, Teleomorph). Canker on thornless
blackberry.
Glomerella
Anthracnose.
Glomerella cingulata. Camellia Dieback, Canker, widespread; some-
times on azalea, blackberry, bittersweet, rose, raspberry, soapberry, moun-
tain-ash, and English ivy; also causing bitter rot of apple (
Rots) and
anthracnose of various hosts (
Anthracnose). Camellia tips die back; leaves

wilt, turn dull green and nally brown. The stem dries out, turns brown, and
there is a girdle of dead bark. Elliptical cankers are present on older wood.
Infection is solely through wounds, principally leaf scars in early spring
but also through bark wounded by cultivating tools or lawn mowers, frost
cracks, or the graft union.
Governor Moulton, Professor Sargent, and some other varieties are rather
resistant; Flora Plena, Prince Eugene Napoleon, and many others are highly
susceptible. Spraying with bordeaux mixture to prevent infection through
leaf and bud scars gives fair control.
Cryptosporiopsis (Glutinium)
Pycnidia innate, without a stroma; spores borne at tip and sides of conidiophores, hya-
line, one-celled.
Cryptosporiopsis pruinosa (formerly Glutinium macrosporum). Canker,
Fruit Rot of apple.
Glutinium macrosporum (see Cryptosporiopsis pruinosa). Canker, Fruit
Rot of apple.
Griphosphaeria
Perithecial wall carbonaceous, mouths papillate; spores dark, with several cells.
Discostroma corticola (formerly Griphosphaeria corticola (Anamorph,
Seimatosporium lichenicola). Rose Canker, Dieback. Cankers are formed
near base of canes, often showing dark glistening pustules of conidia. Occa-
sionally when the canker has girdled the cane, a large gall forms above
the lesion (see Fig. 3.17). It resembles crown gall but is apparently due to
interference with downward transfer of food. Cut out infected canes.
Griphosphaeria corticola (see Discostroma corticola (Anamorph, Seimato-
sporium lichenicola)). Rose Canker, Dieback. Cankers are formed near
base of canes, often showing dark glistening pustules of conidia.
Hendersonula
Pycnidia black, stromata, one to several per stroma, locules occurring at different levels
in stroma; conidophores long, exuous; conidia often extruded in cirrhi; at rst one-
celled, hyaline to yellowish, later becoming three- to four-celled and dark.
Hendersonula toruloidea. Canker on Arbutus menziesii.
Hymenochaete
Pileus, fruiting structure, resupinate, of several layers, with long, stiff, usually brown
setae (cystidia).
Hymenochaete agglutinans. Hymenochaete Canker on apple, birch,
hazelnut, sweetgum, mistletoe, and various young hardwoods. When an
infected dead stem comes in contact with a live one, the mycelium forms
a thin leathery fruiting body around the living stem, holding it to the dead
stem. This resupinate structure is deep brown in the center, with a yellow
margin. The stem is constricted at the point of encirclement, and the sapling
usually dies in 2 or 3 years. If the dead stem is removed before girdling,
Figure 3.17 Discostroma Canker on Rose
a sunken canker appears on one side, but this may be overgrown with callus
and disappear. Do not leave severed stems in contact with living seedlings
or saplings in nursery stands.
Hypoxylon
Perithecia in a pulvinate stroma, often conuent and crustose; ascospores with one cell,
rarely two, blackish brown; conidia in supercial layer on surface of young stroma.
Hypoxylon mammatum. Hypoxylon Canker of poplar. Aspen and large-
tooth aspen are most commonly attacked, balsam poplar less frequently. This
is usually a forest, rather than a home garden, disease. Trees less than 30
years old, growing on poor sites, are most susceptible. Trunk cankers start
as small, yellow to reddish brown, slightly sunken areas, centering around
a wound, there grow together to form a canker marked off by vertical cracks.
The bark is mottled, gray, with black patches where the blackened cortex
is exposed. Conidia appear in blisterlike stromata on rst- and second-year
cankers, whereas perithecia are formed on third-year cankers in hard, black
stromata covered with a white pruinose coat. Ascospores are ejected in win-
ter. Eliminate infected trees when thinning stands.
Kabatina
Kabatina juniperi. Blight on eastern red cedar; conidia produced in black
acervuli on discolored foliage.
Lachnellula
Apothecia mostly cup-shaped.
Lachnellula willkommii (Syn. Trichoscyphello willkommii). Canker of
European larch (
Dasyscypha).
Leptosphaeria
Blights.
Diapleella coniothyrium (formerly Leptosphaeria coniothyrium). Canker
on thornless blackberry.
Leptosphaeria coniothyrium (see Diapleella coniothyrium). Canker on
thornless blackberry.
Leucostoma
Leucostoma cincta. Canker on apple.
Macrophoma
Like Phoma, with discrete pycnidia arising innately, but with much larger spores; conidia
hyaline; one-celled.
Diplodia tumefaciens (formerly Macrophoma tumefaciens). Branch Gall
Canker of poplar. Nearly spherical round galls, not over 1 1/2 inches in
diameter, at base of twigs, which usually die; not serious.
Macrophoma candollei. Associated with Dieback of boxwood but appar-
ently saprophytic only. The large black pycnidia are, however, quite striking
on straw-colored leaves.
Macrophoma cupressi. Dieback of Italian cypress.
Macrophoma phoradendron. Defoliates mistletoe, but it grows back.
Macrophoma tumefaciens (see Diplodia tumefaciens). Branch Gall
Canker of poplar.
Massaria
Ascomycetes, Pyrenulales
Spores dark, with several cells, oblong-fusiform, with mucous sheath.
Massaria platani (see Splanchrorema platani). Canker, widespread on
branches of American, London, and California plane trees.
Splanchrorema platani (formerly Massaria platani). Canker, widespread
on branches of American, London, and California plane trees.
Melanconis
Perithecia in an immersed black stroma; paraphyses present; spores two-celled, light;
conidia supercial on a stroma.
Melanconis juglandis. Walnut Canker, Butternut Dieback, widespread
on butternut, also on black, Japanese, and English walnut. The disease was
rst described from Connecticut in 1923, but evidently was responsible for
slow dying of butternuts long before that. If trees have been previously weak-
ened, the fungus proceeds rapidly; otherwise there is the slow advance of
a weak parasite. Dead limbs are sprinkled with small, black acervuli, looking
like drops of ink and occasionally, in wet weather, developing spore horns
of olive gray conidia. In the teleomorph state, which is rare, perithecia are
embedded in the bark singly or in groups. Mycelium invades bark and wood,
with a dark discoloration, and grows slowly down a branch to the trunk.
When the latter is reached, the tree is doomed. In nal stages trees have
a stag-headed effect from loss of leaves.
Control. Remove diseased branches promptly, cutting some distance below
infection; remove trees developing trunk cankers; keep the rest growing well
with food and water.
Meria
Hyaline mycelium, branched; conidiophores simple, septate; conidia hyaline, one-
celled, produced singly or in clusters.
Meria laricis, Dieback and Blight, on western larch seedlings.
Monochaetia
Acervuli dark, discoid or cushion-shaped, subcutaneous; conidia several-celled, dark
median cells, hyaline end cells, and a single apical appendage; parasitic.
Monochaetia mali (see Seiridium unicorne). Canker, Leaf Spot of apple.
Seiridium unicorne (formerly Monochaetia mali). Canker, Leaf Spot of
apple. Fungus enters through deep wounds and grows into wood, then attacks
resulting wound callus and produces numerous fruiting bodies on exposed
wood and callus layer. Killing of successive callus layers results in a canker
similar to European apple canker. The disease is not common enough to be
serious.
Nectria
Perithecia bright, more or less soft and eshy, in groups, basal portion seated on a stroma;
spores two-celled, hyaline or subhyaline (see Fig. 3.15).
Nectria cinnabarina. Dieback, Twig Canker, Coral Spot, cosmopoli-
tan on hardwoods, most common on maples but also found on ailanthus,
amelanchier, apple, crabapple, apricot, ash, blackberry, chokecherry, beech,
birch, elm, hickory, horsechestnut, mimosa, linden, paper mulberry, pear,
peach, sophora, locust, and honey locust. It may also appear in stem cankers
on vines and shrubsampelopsis, barberry, boxwood, callicarpa, cotoneast-
er, currant, gooseberry, g, honeysuckle, kerria, California laurel, rose, and
syringa. The fungus is widespread as a saprophyte. On ornamental trees and
shrubs it is weakly parasitic, producing cankers around wounds and at base
of dead branches or causing a dieback of twigs and branches.
On maple, the fungus is more pathogenic, killing twigs, small branches,
young trees, and girdling larger branches. It is more frequent on Norway
maple and boxelder; it may also invade red, sycamore, Japanese, and other
maples. First symptoms are small, depressed, dead areas in bark near wounds
or branch stubs. Conspicuous esh-colored or coral pink sporodochia,
formed in dead bark, bear conidia. Later the pustules turn chocolate brown
and form pockets, in which perithecia are produced. The canker is most
common in severely wounded or recently pruned trees. Sapwood has a
greenish discoloration. Open cankers are eventually formed with successive
rolls of callus. Remove diseased wood and bark, cutting beyond the greenish
discoloration.
Nectria coccinea var. faginata. Nectria Beech Bark Canker on beech in
the Northeast. The disease occurs solely in connection with the woolly beech
scale insects (Cryptococcus fagi and C. fagisuga), but it has caused high
mortality in Canada, killing 50% of beech stands; it is epidemic in Maine on
American beech and is now present in much of New England and New York.
The scale nymphs, covered with a woolly white down, cluster thickly around
cracks and wounds in bark, often making trunk and branches appear to be
coated with snow. The small yellow larvae establish themselves on the bark
in autumn, each inserting its sucking organ, stylet, into the living bark, which
shrinks and cracks. Nectria enters through these cracks and kills surrounding
tissue in bark and cambium. When the cells are dead, the insects can no
longer obtain food; therefore, they disappear.
White pustules of sporodochia are pushed out through dead bark, bearing
elongate, three- to nine-celled, slightly curved macroconidia. Red perithe-
cia, slightly lemon-shaped, appear in clusters on the bark, often so abundant
that the bark appears red. After ascospores are discharged, the upper half of
the perithecium collapses and sinks into the lower. The eventual canker is
a deeply depressed cavity surrounded by callus. After the cambium dies, the
leaves wilt; the twigs, branches, and roots nally die.
Control. Ornamental trees can be sprayed or scrubbed to kill the insects.
A dormant lime sulfur spray is very effective. Oil sprays will kill the scale
but may injure beech. Late summer spraying for crawlers can supplement the
dormant spray.
Nectria desmazierii (Fusarium buxicola, Anamorph). Canker and Die-
back of boxwood (see Fig. 3.18).
Nectria ditissima. Sometimes reported but not conrmed in the United
States; reports probably refer to Nectria galligena.
Nectria fuckeliana. Canker on r.
Nectria galligena (Cylindrosporium mali, Anamorph). European Nectria
Canker, Trunk Canker, widespread on apple, pear, quince, aspen, beech,
birch, maple, hickory, Pacic dogwood, and various other hardwoods. This
is one of the more important diseases of apple and pear in Europe but is
less serious in this country. In eastern United States it is primarily an apple
disease; on the Pacic Coast it is more common on pear.
Young cankers are small, depressed or attened areas of bark near small
wounds or at base of dead twigs or branches, darker than the rest of the
bark and water-soaked. Older cankers are conspicuous and somewhat like
a target, with bark sloughed off to expose concentric rings of callus. Cankers
on elm, sugar maple and birch are usually circular; those on oak irregular;
on basswood elongate, pointed at ends. If the canker is nearly covered with
a callus roll, it indicates that the infection is being overcome.
Small red perithecia are formed singly or in clusters on bark or on wood
at margin of cankers. Ascospores discharged during moist weather are dis-
Figure 3.18 Volutella Blight or Nectria Canker on Boxwood
seminated by wind and rain. Creamy-white sporodochia protruding through
recently killed bark of young cankers produce cylindrical macroconidia and
ellipsoidal microconidia. Invasion is through bark cracks or other wounds
in living or dying, but not dead, wood. Infection is slow, with annual cal-
lus formation; only the smallest branches are likely to be girdled. Younger,
more vigorous apple trees receiving nitrogenous fertilizer appear to be more
susceptible.
Control. Remove and destroy small branches with cankers. Clean out trunk
cankers and cut back to sound bark; treat with bordeaux paste. On the West
Coast spray pome fruits immediately after leaf fall in autumn with bordeaux
mixture to prevent infection through leaf scars.
Nectria magnoliae. Nectria Canker, similar to the preceding but found on
magnolia and tuliptree.
Neofabraea
Anthracnose.
Neofabraea perennans (Gloeosporium perennans) (see Pezicula malicorti-
cis). Perennial Canker of apple, also bulls-eye rot of fruit.
Pezicula malicorticis (formerly Neofabraea perennans (Gloeosporium
perennans)). Perennial Canker of apple, also bulls-eye rot of fruit. The
disease is much like northwestern anthracnose. It often follows after winter
injury or starts at pruning cuts where aphids congregate, or may appear after
an application of wound dressing.
Nummularia
Stroma supercial, composed entirely of fungus elements, covered with a conidial layer
when young. Perithecia ask-shaped, embedded in stroma; spores one-celled, dark.
Biscogniauxia marginata (formerly Nummularia discreta). Blister Canker
of apple, crabapple, pear, mountain ash; also reported on serviceberry, birch,
elm, magnolia, and honey locust. This is a major apple disease east of the
Rocky Mountains, especially in Upper Mississippi and Lower Missouri Riv-
er valleys, where millions of apple trees have been killed. Large and small
limbs are affected. Cankers are dead areas, up to 3 feet long, mottled with
living wood and dotted with numerous round cushions of stromata, looking
like nailheads. Perithecia, with dark ascospores, are buried in the stroma-
ta; hyphae bearing small, light-colored conidia grow over the surface. The
fungus enters through branch stubs, bark injuries, and other wounds.
Control. Avoid especially susceptible varieties like Ben Davis. Shape trees
early to prevent large pruning wounds on older trees; the canker seldom
appears on trees less than 10 years old. Shellac pruning cuts immediately;
sterilize tools between cuts.
Nummularia discreeta (see Biscogniauxia marginata). Blister Canker of
apple, crabapple, pear, mountain ash; also reported on serviceberry, birch,
elm, magnolia, and honey locust.
Ophionectria (Scoleconectria)
Perithecia red to white, globoid, with a round ostiole, supercial, paraphyses lacking;
spores needle-shaped to liform, light colored.
Ophionectria balsamea (see Thyronectria balsamea). Bark Canker of bal-
sam r.
Ophionectria scolecospora (see Scoleconectria cucurbitula). Bark Canker
of balsam and alpine rs.
Scoleconectria cucurbitula (formerly Ophionectria scolecospora). Bark
Canker of balsam and alpine rs.
Thyronectria balsamea (formerly Ophionectria balsamea). Bark Canker
of balsam r.
Penicillium
Conidia in heads; conidiophores unequally verticillate at tip in whorls; globose conidia
formed in chains, one-celled, hyaline or brightly colored in mass; parasitic or saprophyt-
ic.
Penicillium vermoeseni. Penicillium Disease of Ornamental palms, seri-
ous in southern California with symptoms varying according to type of palm.
On queen palm (Arecastrum or Cocos plumosa) the disease is a trunk canker,
which may remain inconspicuous for several years but leads to weakening
and breaking of trunk. Infected trees should be removed at an early stage.
On Canary date palm the disease is a leafbase rot, and on Washington a bud
rot.

Rots.
Pezicula
Apothecia similar to Dermatea but lighter.
Pezicula carpinea. Bark Canker of hornbeam.
Pezicula corticola. Supercial Bark Canker and Fruit Rot, rather com-
mon on apple and pears. Hyaline, one-celled conidia of the Myxosporium
stage are formed in acervuli.
Pezicula pruinosa. Canker on branches of amelanchier.
Phacidiella
Asci borne in hymenial layers, covered with a membrane until mature, then splitting;
apothecia remain embedded in a stroma; paraphyses present; asci clavate.
Phacidiella coniferarum (Anamorph, Phacidium coniferarum). Phomop-
sis Disease of conifers. The fungus is usually saprophytic, but it is parasitic
on Douglas-r and larch in Europe and on living pine in Maine.
Phomopsis
Blights.
Phacidiopycnis boycei (formerly Phomopsis boycei). Phomopsis Canker
of lowland white r. Branches or main stem of saplings may be girdled and
killed; there is often swelling at base of canker where dead tissues join living.
The reddish brown needles of dead branches are prominent against living
foliage.
Phacidiopycnis piri (Teleomorph, Potabiamyces pyri, formerly Phomopsis
discolor). Pear branch canker and fruit rot.
Phomopsis alnea. Canker of European black alder.
Phomopsis amygdali. Branch Dieback on almond.
Phomopsis boycei (see Phacidiopycnis boycei). Phomopsis Canker of
lowland white r.
Phomopsis discolor (see Phacidiopycnis piri, see Teleomorph, see Potabi-
amyces pyri). Pear branch canker and fruit rot.
Phomopsis elaeagni (Syn. Phomopsis arnoldia). Canker of Russian-olive.
Phomopsis gardeniae. (Teleomorph, Diaporthe gardeniae). Gardenia
Canker, Stem Gall, widespread in greenhouses. Although not reported
until about 1933, this seems to be the most common gardenia disease.
Symptoms start with brown dead areas on stem, usually near the soil line.
The canker is rst sunken, then, as the stem enlarges, swollen with a rough,
cracked outer cork. The stem is bright yellow for a short distance above
the canker, a contrast to its normal greenish white. When stems are com-
pletely girdled, the foliage wilts and dies; the plant may live a few weeks
in a stunted condition. Flower buds fall before opening. When humidity is
high, black pycnidia on cankers exude yellowish spore masses. Entrance
is through wounds; spores may be spread on propagating knives. Infection
often starts at leaf joints at the base of cuttings after they have been placed
in a rooting medium. Because the cankers may be only slightly visible on
rooted cuttings, the disease may be widely distributed by the sale of such
cuttings.
Control. Use sterilized rooting medium. Use steam for a sand and peat mix-
ture. Destroy infected plants; sometimes it is possible to wait until blooms
are marketed.
Phomopsis lirella (Teleomorph, Diaporthe vincae). Canker, Dieback of
vinca, and periwinkle.
Phomopsis lokoyae. Phomopsis Canker of Douglas-r mostly on sap-
lings in poor sites in California and Oregon. Long, narrow cankers, some-
what pointed at ends, develop during the dormant season after young shoots
are infected. If the tree is not girdled during the rst season, the canker heals
over.
Phomopsis mali. Bark Canker of pear, and apple. The bark is rough.
Phomopsis padina (Telomorph, Diaporthe decorticans). Canker, Twig
Blight of sour cherry.
Phomopsis sp. Shoot Dieback on peach.
Phragmodothella
Asci in locules immersed in groups in a cushionlike stroma; spores hyaline, many-celled.
Dothiora ribesia (formerly Phragmodothella). Dieback, Black Pustule
on currant, owering currant, and gooseberry.
Phragmodothella ribesia (see Dothiora ribesia). Dieback, Black Pustule
on currant, owering currant, and gooseberry.
Physalospora
Blights.
Botryosphaeria corticis (formerly Physalospora corticis). Blueberry
Cane Canker, in Southeast on cultivated blueberries. The fungus enters
through unbroken bark, probably through lenticels, with cankers starting as
reddish, broadly conical swellings, enlarging the next year to rough, black,
deeply ssured cankers that girdle the shoots. The portions above cankers
are unfruitful and nally die. Avoid very susceptible varieties like Cabot and
Pioneer.
Botryosphaeria obtusa (formerly Physalospora obtusa (Sphaeropsis mal-
orum)). Dieback, Canker of hardwoods, New York Apple-Tree Canker,
Black Rot of Apple. The fungus attacks leaves, twigs, and fruits, is more
important east of the Rocky Mountains, and is found on many plants, includ-
ing alder, ampelopsis, birch, bignonia, bittersweet, callicarpa, catalpa, cean-
othus, chestnut, currant, cotoneaster, hawthorn, Japanese quince, maple,
peach, pear, and persimmon. On hardwoods the canker is similar to that
caused by P. glandicola on oaks. Limbs are girdled with large areas of rough
bark with numerous protruding black pycnidia. For the fruit rot phase of this
disease

Rots.
Botryosphaeria quercuum (formerly Physalospora glandicola (Sphaerop-
sis quercina, Anamorph)). Sphaeropsis Canker, Dieback of red, chest-
nut, and other oaks. Shade and ornamental trees of all ages may be killed.
Infection may start anywhere through wounds but more often on small twigs
and branches, passing to larger branches and trunk. Twigs and branches die;
leaves wither and turn brown; infected bark is sunken, and wrinkled, with
small black pycnidia breaking through. On larger stems the bark has a ridge
of callus around the canker, the sapwood in this area turning dark with black
streaks extending longitudinally for several inches. Numerous water-sprouts
grow from below the dead crown. The fungus winters on dead twigs, produc-
ing a new crop of conidia in spring, readily infecting most trees weakened
by unfavorable environmental conditions.
Control. Prune out diseased portions at least 6 inches below cankers. Fertilize
and water to improve vigor. Remove seriously diseased trees.
Botryosphaeria rhodina (formerly Physalospora rhodina). Black Rot
Canker of tung in Mississippi and Louisiana. Black, sunken cankers on
trunks, limbs, twigs, and shoots, may girdle and kill trees. Rogue and burn
diseased specimens.
Glomerella cingulata (formerly Physalospora miyabeana). Willow Black
Canker, accompanying scab to form the disease complex known as willow
blight in New England and New York. Starting in leaf blades, the fungus
proceeds through petioles into twigs; it also causes cankers on larger stems,
followed by defoliation. Pinkish spore masses of the anamorph Gloeospori-
um state are formed on dead twigs and branch cankers and then short-necked
perithecia, which overwinter. Remove and destroy dead twigs and branches
during the dormant period. Spray 3 times with bordeaux mixture, starting
just after leaves emerge in spring.
Physalospora cortices (see Botryosphaeria corticis). Blueberry Cane
Canker, in Southeast on cultivated blueberries.
Physalospora glandicola (Sphaeropsis quercina, Anamorph) (see Botryo-
sphaeria quercuum). Sphaeropsis Canker, Dieback of red, chestnut, and
other oaks.
Physalospora miyabeana (see Glomerella cingulata). Willow Black Can-
ker, accompanying scab to form the disease complex known as willow blight
in New England and New York.
Physalospora obtusa (Sphaeropsis malorum) (see Botryosphaeria obtusa).
Dieback, Canker of hardwoods, New York Apple-Tree Canker, Black Rot
of Apple.
Physalospora rhodina (see Botryosphaeria rhodina). Black Rot Canker
of tung in Mississippi and Louisiana.
Phytophthora
Blights.
Phytophthora cactorum. Bleeding Canker of maple, beech, birch, elm,
horsechestnut, linden, oak, sweetgum, and willow; Crown Canker of dog-
wood; Dieback of rhododendron; Trunk Canker of apple, almond, apricot,
cherry, and peach.
Bleeding Canker, rst noticed in Rhode Island on maple about 1939 and
found in New Jersey the next year, is now present on many trees in the
Northeast. The most characteristic symptom is the oozing of a watery light
brown or thick reddish brown liquid from ssures in bark at the root collar
and at intervals in trunk and branches. When dry, this sap resembles dried
blood, hence the name, bleeding canker. Sunken, furrowed cankers are more
denite on young trees than on older trees with rough bark. Symptoms are
most prominent in late spring and early fall, with trees in moist situations
most often affected. The fungus lives in the soil and advances upward from
a primary root infection. Wilting of leaves and blighting of branches is evi-
dently from a toxin. Mature trees have fewer, smaller, yellow-green leaves,
and there is an acute dieback of branches. Reddish-brown areas with intense
olive-green margins are found in wood extending vertically from roots to
dying branches, marked at irregular intervals with cavities containing the
watery uid.
Control. Although there is no real cure, injecting trees with Carosel, a mix-
ture of helione orange dye and malachite green, has inhibited the fungus and
neutralized the toxin. In some cases trees recover without treatment. Avoid
heavy feeding; this seems to encourage the spread of disease and causes
chronic cases to become acute.
Crown Canker, collar rot, is the most serious disease of dogwood reported
in New York, New Jersey, and Massachusetts. The rst symptom is a gener-
al unhealthy appearance, with leaves smaller and lighter green than normal,
turning prematurely red in late summer. Leaves may shrivel and curl dur-
ing dry spells (normal leaves often do likewise). Twigs and large branches
die, frequently on one side of the tree. The canker develops slowly on the
lower trunk near the soil level. Inner bark, cambium and sapwood are dis-
colored; the cankered area is sunken; the bark dries and falls away, leaving
wood exposed. Trees die when the canker extends completely around the
trunk base or root collar. The fungus lives in the soil in partially decayed
organic matter, and spores are washed to nearby trees. Entrance is through
wounds. The disease affects transplanted dogwoods, seldom natives growing
in woods.
Control. Transplant carefully, avoiding all unnecessary wounds; avoid hitting
base with lawnmower, by using a wire guard around the tree. It is difcult
to save trees already infected, but cutting out small cankers and painting the
wound with bordeaux paste is worth trying. If trees have died from crown
canker, do not replant with dogwoods in the same location for several years.
Rhododendron Dieback, is a disease in which terminal buds and leaves turn
brown, roll up, and droop as in winter cold. A canker encircles the twigs,
which shrivel with the terminal portion wilting and dying. In shady locations
leaves have water-soaked areas, changing to brown, zonate spots. Do not
plant rhododendrons near lilacs, for they are blighted by the same fungus.
Prune diseased tips well below the shriveled part, and spray after blooming
with bordeaux mixture, two applications 14 days apart.
Trunk Canker of Apple, is an irregular canker often involving the entire
trunk and base of scaffold branches, the rst outward symptom a wet area
on bark. Trees must be 5-years old or older for infection. Grimes Golden
and Tomkins King are especially susceptible, often being completely gir-
dled.
Phytophthora cinnamomi. Basal Canker of Maple, particularly Norway
maple. Trees have a thin crown, fewer and smaller leaves, and die a year or
two after cankers are formed at the base of the trunk. Sapwood is reddish
brown; the roots decay. Remove diseased trees. Plant new Norway maples
in good soil, well drained, rich in organic matter; treat injuries at base of
trunk promptly. See under Rots and Wilts for other manifestations of this
pathogen.
Phytophthora syringae. Pruning Wound Canker of almond.
Plenodomus
Pycnidia dark, immersed, irregular in shape and opening irregularly; conidia hyaline,
one-celled, oblong; parasitic.
Aposphaeria fuscomaculans (formerly Plenodomus fuscomaculans). Can-
ker on apple.
Plenodomus fuscomaculans (see Aposphaeria fuscomaculans). Canker on
apple.
Pseudonectria
Perithecia supercial, blight-colored, smooth; spores one-celled, hyaline.
Pseudonectria rouselliana. Nectria Canker of boxwood, Leaf Cast,
Twig Blight. The perithecia are formed on dead leaves, but the fungus is
thought to be the teleomorph state of Volutella buxi, which see.
Pseudovalsa
Perithecia in a stroma; spores dark, with several cells.
Pseudovalsa longipes. Twig Canker on coast live oak and white oak.
Rhabdospora
Pycnidia separate, not produced in spots, erumpent, ostiolate; conidiophores short, sim-
ple conidia hyaline, liform to needle-shaped, with several cells; parasitic or saprophyt-
ic.
Rhabdospora rubi (see Septocyta ruberum). Cane Spot, Canker of rasp-
berry.
Septocyta ruberum (formerly Rhabdospora rubi). Cane Spot, Canker of
raspberry.
Scleroderris
Apothecia black, opening with lobes, crowded together or with a stroma, short-stalked;
spores hyaline, elongate, with several cells.
Ascocalyx abietina (formerly Scleroderris lagerbergii =Gremmeniella abi-
etina). Canker on pine.
Grovesiella abieticola (formerly Scleroderris abieticola). Canker of bal-
sam r, on Pacic Coast. An annual canker, starting in autumn and ceasing
when cambium is active in spring, is formed on twigs, branches, and trunks
of saplings. Only twigs and small branches are girdled, and if this does not
happen before spring, the wound heals over. Small black apothecia with short
stalks appear on dead bark. Ascospore infection is through uninjured bark or
leaf scars.
Scleroderris abieticola (see Grovesiella abieticola). Canker of balsam r,
on Pacic Coast.
Scleroderris lagerbergii Syn. Gremmeniella abietina (see Ascocalyx abieti-
na). Canker on pine.
Scleroderris lateritium. Canker on pine.
Sclerotinia
Blights.
Sclerotinia (Syn. Whetzelinia) sclerotiorum. Basal Canker on Euonymus.
Stem Canker and Wilt on sage.
Septobasidium
Basidiomycetes, Septobasidiales
All species are on living plants in association with scale insects; the com-
bination causes damage to trees. Fungus body variable, usually resupinate,
dry, crustaceous or spongy, in most species composed of subiculum grow-
ing over bark; a middle region of upright slender or thick pillars of hyphae
supports the top layer, in which hymenium is formed. Basidium transversely
septate into two, three, or four cells, rarely one-celled; basidiospores ellipti-
cal, colorless, divided into two to many cells soon after formation, budding
with numerous sporidia if kept moist. Some species with conidia.
The fungus lives by parasitizing scales, obtaining food via haustoria. The
insects pierce the bark to the cambium, sometimes killing young trees.
The fungus kills a few scales but protects many more in its enveloping
felty or leathery covering, a symbiotic relationship. Spores are spread
by scale crawlers and by birds. Most felt fungi are found in the South,
abundant on neglected fruit, nut, or ornamental trees, rare on those well
kept.
Septobasidium burtii. Felt Fungus on southern hackberry, beech, pear,
apple, and peach. This is a perennial growth, with a new ring added to the
patch each summer. Probasidia are formed during the winter, and four-celled
basidia in spring.
Septobasidium castaneum. Felt Fungus abundant on willow and water
oaks, and holly; may injure azaleas. The surface is smooth, shiny, chocolate
brown to nearly black.
Septobasidium curtisii. Felt Fungus, widespread on many trees in the
Southeast, commonly on sour gum (tupelo) and American ash, also on hicko-
ry, hawthorn, Japanese quince, and others. The felt, purple-black throughout,
is mounded over the insects.
Septobasidium pseudopedicellatum. Felt Fungus, on citrus twigs, some-
times on main stem or branches of hornbeam. Surface is smooth, buff-
colored over dark brown pillars.
Solenia (Henningsomyces)
Fruiting layers erect, cylindrical, formed in groups, membranous.
Cyphellopsis anomala (formerly Solenia (Henningsomyces) anomala).
Bark Patch, Canker, widespread on alder.
Merismodes ochracea (formerly Solenia ochracea). Bark Patch of birch,
hornbeam, hickory, and alder.
Solenia (Henningsomyces) anomala (see Cyphellopsis anomala). Bark
Patch, Canker, widespread on alder.
Solenia ochracea (see Merismodes ochracea). Bark Patch of birch, horn-
beam, hickory, and alder.
Sphaeropsis
Pycnidia black, separate or grouped, globose, erumpent, ostiolate; conidiophores short;
conidia large, dark, one-celled, ovate to elongate, on liform conidiophores. Some
species have Physalospora as the teleomorph state.
Sphaeropsis sapinea. Bleeding Canker on pine.
Sphaeropsis tumefaciens. Canker and Gall on Carissa.
Sphaeropsis ulmicola. Sphaeropsis Canker of American elm. The dis-
ease spreads downward from small twigs to larger branches with a brown
discoloration of wood just under the bark. Secondary shoots sometimes
develop below the cankers. Trees weakened by drought or poor growing
conditions are particularly susceptible. Prune out infected wood, cutting
well below cankers.
Stegonsporium
Stegonsporium sp. Maple Canker, Dieback. Reported from New Jersey.
Large branches die back with conspicuous agging. Black tarlike fruiting
bodies are formed in cankers.
Strumella
Sporodochia wartlike, gray to black, of interwoven hyphae; conidiophores dark, branch-
es; conidia dark, one-celled, ovoid to irregular.
Conoplea globosa (formerly Strumella coryneoidea). Strumella Canker
of oak, especially the red oak group, also on American beech and chestnut,
occasional on pignut and hickories, red maple, and tupelo. Primarily a forest
disease, this canker may become important on red and scarlet ornamental
oaks. Starting as a yellowish discoloration of bark around a dead branch
or other point of infection, the canker develops into a diffuse lesion or into
a target canker with concentric rings of callus. Whitish mycelium is present
near outer corky bark, and the infected portion of the trunk may be attened
or distorted. Target cankers may be up to 2 feet wide and 5 feet long. The
small black nodules bear no spores while trees are living, but after death dark
brown spore pustules are formed, which blacken with age. New pustules are
formed yearly. Canker eradication has been unsuccessful in forest stands.
The diseased trees should be removed and utilized before spores can spread
infection.
Strumella coryneoidea (see Conoplea globosa). Strumella Canker of oak,
especially the red oak group, also on American beech and chestnut, occasion-
al on pignut and hickories, red maple, and tupelo.
Sydowia
Ascomycetes, Dothidiales
Asci usually short, cylindrical, and relatively numerous, in spherical, ostiolate locules.
Sydowia polyspora. Twig Dieback on r.
Thyronectria
Stroma valsoid with several perithecia, bright-colored; spores muriform, hyaline to sub-
hyaline.
Thyronectria austro-americana. Canker, Wilt of honeylocust. Slightly
depressed cankers ranging from pinhead size to 1/2 inch grow together and
enlarge to girdle a branch. Underlying wood is streaked reddish brown for
several inches from the canker, and there is often a gummy exudate. Some
trees die, but many survive.
Thyronectria balsamea. Canker on r.
Thyronectria berolinensis. Cane Knot Canker of fruiting and owering
currants.
Trichothecium
Rots.
Trichothecium roseum. Canker of rose.
Tubercularia
Forms bright colored cushions, mostly on wood or bark; ne branching conidiophores
bearing small, elipsoidal hyaline conidia.
Tubercularia ulmea. Canker on Russian olive and honeylocust.
Tympanis
Ascomycetes, Helotiales, Helotiaceae
Ascocarp cup-shaped; sclerotia absent; expiculum usually, if parallel hyphae.
Tympanis confusa. Canker on pine.
Valsa
Many perithecia in a circle in a stroma in bark; ask-shaped with long necks opening to
the surface; spores hyaline, one-celled, curved, slender.
Leucostoma cincta (formerly Valsa cincta). Perennial Canker of peach,
Dieback, also on nectarine. The fungus is apparently infective during the
dormant season, entering through wounds, dead buds, leaf scars, and fruit
spurs. It forms a canker complex with V. leucostoma and sometimes the
brown-rot fungus. It is more common in northern latitudes than in southern,
but is not important in well-cared-for orchards.
Leucostoma kunzei (formerly Valsa kunzei).

Cytospora kunzei.
Leucostoma persoonii (formerly Valsa leucostoma). Apple Canker,
Dieback, Twig Blight on apple, apricot, peach, pear, quince, plum, cherry,
willow, and mountain-ash. The fungus is a weak parasite entering through
wounds or twigs killed by frost.
Valsa cincta (see Leucostoma cincta). Perennial Canker of peach, Die-
back, also on nectarine. Valsa kunzei (see Leucostoma kunzei).
Cytospora
kunzei.
Valsa leucostoma (see Leucostoma persoonii). Apple Canker, Dieback,
Twig Blight on apple, apricot, peach, pear, quince, plum, cherry, willow,
and mountain-ash. Valsa salicina (Cytospora salicis). Twig and Branch
Canker of willow.
Valsa sordida.
Cytospora chrysosperma.
Vermicularia
Like Colletotrichum but setae are scattered throughout the acervuli, not just marginal;
spores hyaline, globose to fusoid.
Vermicularia ipomoearum. Stem Canker of morning glory.
Volutella
Blights.
Volutella buxi. Boxwood Nectria Canker, Volutella Blight. The
teleomorph state of the fungus is supposed to be Pseudonectria rouselliana,
which see. As a canker the disease often follows after winter injury, with
salmon-pink spore pustules on dying twigs, branches, and main stems. As
a blight, the fungus spreads rapidly in moist weather in summer, attacking
healthy twigs when humidity is high and often discernible at a distance by
a straw yellow ag. On such yellowing branches the backs of leaves and
the bark of twigs are both covered with the pinkish spore pustules.
Control. Cut out branches where the bark has been loosened by winter ice
and snow. Have a yearly housecleaning, brushing out accumulated leaves
and other debris from interior of bushes and cutting out all twigs with pink
pustules. If there are signs of disease, follow cleaning with thorough spray-
ing, from ground up through interior of bushes, with lime sulfur.
CLUB ROOT
Plasmodiophora
Plasmodiophoromycetes, Plasmodiophorales
This genus, founded on the club root organism, has a somewhat doubtful taxonomic
position. Formerly considered a slime-mold, one of the Myxomycetes, then placed in
the Chytridiales, lowest order of true fungi, it is now placed in a separate order, Plas-
modiophorales.
Thallus amoeboid, multinucleate in host cell; spores lying free in host cell at
maturity; frequently causing hypertrophy; parasitic on vascular plants.
Plasmodiophora brassicae. Club Root of cabbage and other crucifers;
nger-and-toe-disease, on alyssum, brussels sprouts, cabbage, Chinese cab-
bage, candytuft, cauliower, hesperis, honesty, peppergrass, garden cress,
mustard, radish, rutabaga, stock, turnip, and western wallower.
Club root was present in western Europe as early as the thirteenth century, but
the true cause was not known until the classic paper of the Russian Woronin
in 1878. The disease was important in the United States by the middle of the
nineteenth century, and is now present in at least 37 states. Losses come from
death of the plants and also from soil infestation, for susceptible crucifers
cannot be grown again on the same land for several years, unless it is treated.
The rst symptom is wilting of tops on hot days, followed by partial recov-
ery at night; affected plants may be stunted and not dead; outer leaves turn
yellow and drop. The root system becomes a distorted mass of large and
small swellings, sometimes several roots swollen like sweet potatoes, and
sometimes joined in one massive gall. Lateral and tap roots are scabby and
ssured, with rot starting from secondary fungi.
When diseased roots decompose, small spherical spores are liberated in the
soil; they are capable of surviving there many years between crops. In spring,
with suitable temperature and moisture, the resting spores germinate, each
CLUB ROOT 269
becoming a motile swarm spore with a agellum. This whiplike appendage
is soon lost, and the organism becomes amoebalike, moving by protoplas-
mic streaming until it reaches a root hair or other root tissue. The plasmodi-
um continues to grow and divide until it reaches the cambial cells, in which
it develops up and down the root. The swelling is produced by division of
plasmodia and of the infected cells. Eventually the multinucleate plasmodi-
um breaks up into many small resting spores, each rounded up around a sin-
gle nucleus. They are set free by the millions when the root rots, and are
spread in soil clinging to shoes or tools and in drainage water, manure, and
plant refuse. Spores are not seed-borne. Long-distance spread is probably by
infected seedlings. Infection takes place chiey in a neutral to acid soil, pH
5.0 to 7.0, at temperatures below 80F, and when moisture of soil is above
50% of its water-holding capacity.
Control. Inspect seedlings carefully before planting. Dispose of infested
crops with caution; resting spores passed through animals are still viable.
A long rotation of crops has been recommended, combined with adding lime
to soil, which must be applied in large amounts, about 6 weeks before the
cabbage crop is set. This brings the pH too high to use potatoes as a follow-
ing crop. Most turnip and rutabaga varieties are relatively resistant to strains
of the club root organism present in the United States.
DAMPING-OFF
Damping-off is the destruction of young seedlings by soil organisms. There
are two types. Pre-emergence damping-off rots the sprouting seed before it
breaks through the soil; it is recognized by bare spaces in what should be uni-
form rows. Such a poor stand may be due to poor viability of seed, but more
often it is due to soil fungi functioning in cold, wet soils when germination
is slow. Post-emergence damping-off is the rotting or wilting of seedlings
soon after they emerge from the soil. Succulent stems have a water-soaked,
then necrotic and sunken, zone at ground level; the little herbaceous plants
fall over on the ground or, in woody seedlings, wilt and remain upright. Root
decay follows. This type of damping-off is most common in greenhouses or
outdoors in warm humid weather and where seedlings are too crowded. Tree
seedlings in nursery rows are subject to this type of damping-off, and so are
perennial owers started in late summer for the next year.
Many fungi living saprophytically in the upper layers of soil can cause
damping-off. Pythium debaryanum, P. mastophorum and Rhizoctonia solani
are probably most common, but other species of these two genera and
Aphanomyces, Botrytis, Cylindrocladium, Diplodia, Fusarium, Macrophom-
ina, Helminthosporium, Sclerotium rolfsii, Fusarium equiseti, and Phytoph-
thora may be important on occasion. A synergistic interaction of Pythium
myriotylum, Fusarium solani, and Meloidogyne arenaria causes damping-
off of peanut which has been reported in Florida. See under Rots for
details. Also, Caloscypha fulgens (anamorph state, Geniculodendron pyri-
forme) causes damping-off of spruce seed, Colletotrichum gloeosporioides
of papaya, Colletotrichum acutatum of owering dogwood and Fusarium
moniliforme var. intermedia of pine.
Damping-off is prevented by starting seed in a sterile medium, such as ver-
miculite, perlite, or sphagnum moss, or by treating the soil or the seed before
planting. Commercial operators treat soil with steam or electricity.
Seed treatment, the coating of seed with a protectant dust, is crop insurance.
In some seasons, good stands can be obtained without it, but it scarcely pays
DAMPING-OFF 271
to take a chance. Seed disinfection is used to kill organisms of anthracnose
and other specic diseases carried on seed. The damping-off organisms are
in the soil, not on the seed, and coating the seed with a chemical is intended
to kill or inhibit fungi in the soil immediately surrounding the seed and so
provide temporary protection during germination.
DODDER
Dodders are seed plants parasitic on stems and other parts of cultivated
or wild plants. They are leaess, orange to yellow twining vines, without
chlorophyll and hence incapable of manufacturing their own food. They are
called love vine, strangle weed, gold thread, hairweed, devils hair, devils
ringlet, pull down, clover silk, and hell-bind, the last being most appropriate.
There are about 40 species in the United States, causing serious agricultural
losses in clovers, alfalfa, and ax, and becoming more and more important
in gardens on ornamentals and sometimes vegetables. Dodders belong to the
single genus Cuscuta, family Cuscutaceae, close to the morning-glory fami-
ly.
Dodder seed is grayish to reddish brown, resembling small legume seed but
roughened with three attened sides. It germinates as ordinary seed but is
synchronized to start a little later than its host seedlings. The parasite is
a slender, yellowish, unbranched thread with the growing tip circling around
in search of support. When it touches the host it twines like a morning-
glory and puts out little suckers, haustoria, into the stem of the victim, after
which its original connection with the soil dries up (see Fig. 3.19). Although
seedlings can live for a few weeks without a susceptible host, they nally die
if a connection is not established. Successful parasites continue to twine and
to spread orange tendrils from one plant to the next, often making a tangle of
matted orange hairs many feet across, with a black region in the center where
plants have died. Such tangles are conspicuous in weeds along roadsides.
In ornamental plantings host plants are not often killed but exhibit stunt-
ing and pallor, symptoms of starvation. Minute scales or rudimentary leaves
form on the dodder tendrils followed by dense clusters of beautiful white
blossoms (sometimes pale pink or yellow), which ripen seed in late summer,
with as many as 3000 seed being produced on a single plant.
Cuscuta spp. Much of the dodder infesting ornamentals is not readily iden-
tied as to species, but it is widespread on a great many shrubs, perennials
and annuals. It is found very commonly on chrysanthemum, also strangling
DODDER 273
Figure 3.19 Dodder on Oleander
any other plant in the vicinity. Many hours may be spent cleaning up ivy
and trumpet-vine, petunias and asters. Dodder is reported on camellias in the
South. It is even a pest of house plants, if eld soil has been used for the
potting mixture. Dodder has, however, one virtue for plant pathologists. It is
used as a bridge between plants to carry viruses and MLOs in testing their
host range.
Cuscuta americana on citrus. C. californica on beet.
Cuscuta coryli. Hazel Dodder. C. epithymum. Clover Dodder on
legumes.
Cuscuta exaltata on redbud, ilex, and sumac.
Cuscuta gronovii. Common Dodder on buttonbush, cucumber, raspber-
ry, members of the potato family, and many garden ornamentals, including
hedge plants.
Cuscuta indecora. Bigseed Alfalfa Dodder on alfalfa from Colorado
westward, also on sweet pea and tomato.
Cuscuta paradoxa on rose, Texas and Florida.
Cuscuta pentagona (C. arvensis). Field Dodder, widely distributed, most
common and serious east of Mississippi on many cultivated and wild herba-
ceous plants.
Cuscuta planifera. Littleseed Alfalfa Dodder, on some legumes in the
West.
Control. Avoid dodder-infested seed. Commercial seed containing one or
more dodder seed per 5-gram sample is prohibited entry into the United
States. Many states have laws regulating sale of infested seed, but it may
still be included inadvertently in a seed packet. If any contamination with
rough, at-sided seed is found, do not use any of the lot. Commercial dealers
sometimes clean infested seed by screening or treating with an iron powder,
which sticks to the rough dodder seed so it can be drawn out by magnets.
Before breaking new ground for a garden on native sod, examine it carefully.
If dodder is found, burn over the area, then hoe lightly but repeatedly for sev-
eral weeks to allow buried seed to germinate and die. When dodder is present
on cultivated plants, the only thing to do is to remove and burn infested parts
before seed is formed. Pulling off the orange tendrils is not sufcient. All
parts of the plant attacked must be cut off and burned, for even a small frac-
tion of a tendril left twined around a stem will start growing again.
A fungus, Colletotrichum destructivum, has been found to parasitize dodder
and offers a slight possibility of biological control.
DOWNY MILDEWS
Downy mildews, sometimes called false mildews, are Oomycetes, in the
order Peronosporales and all in the family Peronosporaceae except Phytoph-
thora in the Pythiaceae. They form mycelium in higher plants and produce
sporangiophores that protrude through stomata in great numbers, their spo-
rangia making white, gray, or violet patches on the leaves. The downy effect
distinguishes these mildews from the true or powdery mildews that form
white felty or powdery patches.
The sporangiophores are often branched; they bear a single sporangium at
the tip of each branch simultaneously, or successively in Phytophthora. Spo-
rangia germinate by swarm spores or with a germ tube as a conidium. An
oospore, resting spores with external ridges or knobs, is formed in an oogo-
nium, large globular multinucleate female cell, after it is fertilized by the
antheridium, a smaller male cell. The oospores are set free by weathering
and decay of host parts.
Basidiophora
Sporangiophore a single trunk with a swollen apex from which short branches grow out,
each bearing a nearly globose sporangium; germination by swarm spores; oospore wall
not conuent with that of oogonium. Mycelium is intercellular, haustoria small, knoblike
(Fig. 3.20).
Basidiophora entospora. Downy Mildewof aster, China aster, goldenrod,
and erigeron. Aster losses are reported by commercial growers in the South,
but apparently this is not an important garden problem.
Bremia
Figure 3.20 Downy Mildews Fruiting from Stomata on Underside of Leaves. Basidiophora, sporangiophoore with
swollen apex; Bremia, sporangiophore tip enlarged to a disc, dichotomous branching; Peronospora, sporangia on
sharply pointed terminal branches; Plasmopara, on obtuse tips; a and a
1
, sporangium; b
1
, zoospore
Dichotomous branching of sporangiophores; tips enlarged into discs bordered with
sterigmata bearing sporangia; swarm spores rare; germination usually by a germ tube
protruded through an apical papilla (Fig. 3.20).
Bremia lactucae. Downy Mildew of lettuce and other composites, endive,
cornower, centaurea, celtuce, escarole, romaine, and various weeds. First
noticed around Boston in 1875, the disease is serious in greenhouses and
in states where outdoor winter crops are grown. Light green or yellowish
areas on upper surface of leaves are matched by downy patches on the under
surface. Affected portions turn brown, and leaves die, the older ones rst.
Entrance is through stomata. The disease is worse in damp, foggy, cool
weather (43 to 53F).
Control. The pathogen has numerous physiological races so that lettuce vari-
eties like Imperial 44 and Great Lakes that are resistant in some localities
may not be so in others. Avoid excessive irrigation; eliminate crop residue
and weeds.
Peronospora
Mycelium intercellular; haustoria in a few species short and knoblike, but in most la-
mentous and more or less branched. Sporangiophore with erect trunk two to ten times
dichotomously branched, with branches somewhat reexed and terminal branches sharp-
pointed; sporangia colored, lacking an apical papilla, germinating from an indeterminate
DOWNY MILDEWS 277
point on the side. Oospores smooth or variously marked, germinating by germ tubes (see
Fig. 3.20).
Peronosclerospora sorghi (formerly Peronospora sorghi). Downy mildew,
on sweet corn.
Peronospora antirrhini. Snapdragon Downy Mildew, reported from
California, Oregon, Oklahoma, Pennsylvania, and Maryland.
Peronospora arborescens. Downy Mildew of prickly-poppy on leaves,
buds, and capsules. Yellow or light brown blotches on upper leaf surface
turn dark, with light gray mold on the underside. The fungus winters in old
plant debris in soil. Remove and burn infected plants. Use clean seed.
Peronospora arthuri. Downy Mildew of godetia, clarkia, gaura, and
evening primrose.
Peronospora destructor. Onion Dowy Mildew, Blight, general on onion,
shallot, Vidalia sweet onion, and garlic. One of the more serious diseases of
onion, reported in the United States in 1884. All varieties are susceptible, but
red onions have some resistance. Reduction in yield may be as high as 75%.
The rst sign of onion mildew is the production of conidiophores with a pur-
plish tinge a short distance back from tips of older leaves. Leaves turn yellow,
wither, and break over; seedstalks may be infected. Onion mildew is spo-
radic, abundant in years of heavy rainfall. Spores, produced in great numbers
in rain or when plants are wet with dew, lose vitality quickly when exposed to
sun. Low temperature, optimum 50F, favors infection. The fungus winters
as mycelium in bulbs, in overwintering plants in mild areas, or as oospores
in soil. Perennial onions in home gardens are considered an important source
of primary inoculum, but oospores have been known to survive 25 years in
soil.
Control. Calred is a resistant variety adapted to California. More onion seed
is being produced in Idaho, where dry summers preclude mildew.
Peronospora dianthicola. Carnation Downy Mildew, common in Cal-
ifornia on seedlings. Leaves turn pale, curl downward; terminal growth is
checked, and plants may die. There is a white growth on lower leaf sur-
faces.
Peronospora effusa. Spinach Downy Mildew, Chard Blue Mold, found
wherever spinach and swiss chard are grown, absent some seasons, nearly
destroying the crop in others. Large pale yellow spots grow together to cover
all or part of the leaf; lower leaves are infected rst, and then the blight is
scattered through the plant. Gray to violet mold forms on underside of leaves;
sometimes the whole plant decays and dries. Initial infection comes from
oospores in the soil; it requires humidity above 85% and a mean temperature
between 45 and 65F for a week. Secondary infection is from conidia. The
fungus is an obligate parasite and does not live over on hosts other than
spinach.
Control. Plant on well-drained, fertile ground; do not crowd; if overhead
irrigation is used, water early on sunny days; practice a 2- to 3-year crop
rotation. Resistant varieties such as Caliay and Texas Early Hybrid 7 are
being introduced.
Peronospora farinosa (formerly Peronospora schactii). Beet Downy
Mildew, on beet, sugar beet, and swiss chard. Inner leaves and seedstalks
are stunted and killed, covered with violet down. The disease appears on the
Pacic Coast during the fall rainy season. Oospores can survive in the soil
several years.
Peronospora fragariae. Strawberry Downy Mildew.
Peronospora grisea, on veronica, a grayish mildew on underside of leaves.
Peronospora manshurica. Soybean Downy Mildew, general. Yellow-
green foliage spots turn brown, with a grayish mold underneath; there may
be premature defoliation. The pathogen winters as mycelium in seed and
oospores in soil. There are at least three races.
Peronospora myosotidis. Forget-Me-Not Downy Mildew, also on lap-
pula. Pale spots on upper surface of leaves, with downy growth under-
neath.
Peronospora oxybaphi, on sand verbena and four-oclock.
Peronospora parasitica, on garden cress; P. leptosperma, on artemisia;
P. linariae, on linaria; P. lophanthi, on agastache.
Peronospora parasitica. Downy Mildewof crucifers, general on cabbage,
Chinese cabbage, broccoli, cauliower, horseradish, radish, turnip, cress,
peppergrass, also on sweet alyssum, arabis, arugula, stock, and hesperis.
Chief damage is to cabbage seedlings or plants grown for seed. Leaf lesions
are light green, then yellow, with downy mold on both sides of the leaf in the
widening yellow zone but not in the dead, shrunken, gray or tan central por-
tion. Secondary fungi often cover dead parts with a black sooty mold. Fleshy
roots of turnips and radishes may be discolored internally. Warm days and
cool nights favor the disease. The pathogen lives between crops in perennial
plants or winter annuals. There are several strains of P. parasitica; one, often
reported as P. matthiolae, blights stock in greenhouse and nursery. Leaves
wilt; tender stems and ower parts are stunted and dwarfed.
DOWNY MILDEWS 279
Control. Avoid crowding plants; keep foliage dry. Spray cabbage seedlings;
repeat two or three times a week until plants are set in eld. Treat heading
cabbage every 6 or 7 days beginning 1 to 3 weeks before harvest.
Peronospora pisi. Pea Downy Mildew. Water-soaked tissue and white
growth appear on any aerial plant part. The mycelium winters in vetch stems,
fruiting there in spring, and spores are disseminated back to peas. The dis-
ease is not important enough for control measures.
Peronospora potentillae. Downy Mildewof agrimony and mock strawber-
ry.
Peronospora radii. Downy Mildew of Marguerite daisy, Argyranthemum
frutescens (formerly Chrysanthemum frutescens).
Peronospora rubi. Downy Mildew of blackberry, dewberry, and black
raspberry.
Peronospora rumicis. Rhubard Downy Mildew. A European disease
reported from California on garden rhubarb. Fungus winters in rootstalks
and grows up into new leaves.
Peronospora schactii (see Peronospora farinosa). Beet Downy Mildew,
on beet, sugar beet, and swiss chard.
Peronospora sorghi (see Peronosclerospora sorghi). Downy mildew, on
sweet corn.
Peronospora sparsa. Rose Downy Mildew, chiey on roses under glass,
rarely outdoors. Young foliage is spotted, leaves drop; owers are delayed or
unmarketable. Abundant spores are produced on undersurface of leaves. To
control, keep humidity below 85% and daytime temperature relatively high.
Peronospora statices. Downy Mildew on statice.
Peronospora tabacina. Blue Mold of tobacco, Downy Mildew; also on
eggplant, pepper, and tomato. This is a seedling disease that can be controlled
by sprays on eggplant and pepper; it is unimportant on tomato.
Peronospora trifoliorum. Downy Mildew of lupine, and alfalfa.
Phytophthora
Blights.
Phytophthora phaseoli. Downy Mildew of lima bean, most important in
Middle and North Atlantic states, in periods of cool nights, heavy dews, and
fairly warm days. Some seasons it takes 50 to 90% of the crop; in other
years it is of little consequence. The white downy mold is conspicuous on the
pod, either in patches or covering it completely. The fungus grows through
the pod wall into the bean, then the pod dries, turns black. On leaves the
white mycelial weft appears sparingly, but veins are often twisted, purplish,
or otherwise distorted. Young shoots and owers are also attacked, bees and
other insects carrying spores from diseased to healthy blossoms. The fungus
fruits abundantly on pods, stems, and leaves; spores are splashed by rain.
Control. Use seed grown in the West where mildew is not present; plan a 2-
to 3-year rotation. Copper dusts are satisfactory.
Plasmopara
Sporangiophores with monopodial branches, with obtuse tips, arising more or less at
right angles; haustoria unbranched and knoblike; sporangia (conidia) small, hyaline,
papillate, germinating sometimes by germ tubes but usually by swarm spores; oospores
yellowish brown, outer wall wrinkled, sometimes reticulate, oogonial wall persistent but
not fused with oospore wall (see Fig. 3.20).
Plasmopara acalyphae. Acalypha Downy Mildew.
Plasmopara geranii on geranium. P. gonolobi on gonolobus.
Plasmopara halstedii. Downy Mildew of bur-marigold, centaurea, eriger-
on, eupatorium, gnaphalium, goldenrod, hymenopappus, Jerusalem arti-
choke, ratibida, rudbeckia, senecio, silphium, verbesina, and vernonia.
Zoospores germinate in soil moisture and invade seedlings via root hairs;
mycelium moving up into stem and leaves causes early wilting and death.
Older plants may not die but exhibit a light yellow mottling. Sporangio-
phores project through stomata on underside of leaves. The fungus winters
in seed and as oospores in soil.
Plasmopara crustosa (formerly Plasmopara nivea). Downy Mildew of
carrot, parsley, parsnip, and chervil. Yellow spots on upper surface of foliage
and white mycelial wefts on under surface turn dark brown with age. The
disease is relatively infrequent, important when plants are so crowded they
cannot dry off quickly after rain or heavy dew. Control by spacing rows prop-
erly.
Plasmopara nivea (see Plasmopara crustosa). Downy Mildew of carrot,
parsley, parsnip, and chervil.
DOWNY MILDEWS 281
Plasmopara pygmaea, on anemone, and hepatica. Fine white mildew covers
underside of leaves; plants are distorted, stems aborted.
Plasmopara viburni. Viburnum Downy Mildew.
Plasmopara viticola. Grape Downy Mildew, general on grape, also on
Virginia Creeper and Boston ivy. This is a native disease, endemic in eastern
United States, rst observed in 1834 on wild grapes. It appeared in France
after 1870, imported with American stock resistant to the Phylloxera aphid,
and in a few years had become as ruinous to the wine industry of Europe
as the potato blight had been to Ireland. The efcacy of bordeaux was rst
discovered in connection with this mildew.
In this country downy mildew is most destructive on European varieties of
grape. Pale yellow spots, varying in form but often nearly circular and some-
what transparent, appear on upper leaf surfaces, and a conspicuous white
coating appears on lower surfaces. The spots turn brown with age; in dry
weather the downy growth is scanty. Young canes, leafstocks, and tendrils
may be infected; owers may blight or rot; young fruits stop growing, turn
dark, and dry with a copious grayish growth. Older fruits have a brown rot
but lack the mildew effect. Fruits from diseased vines have less juice; bunch-
es are very poorly lled.
Initial infection comes from a swarm spore stopping on the lower side of
a leaf, putting out a germ tube and entering through a stoma. In 5 to 20
days the mycelium has spread through the leaf between cells, obtaining
food through thin-walled, globular haustoria. The hyphae mass in compact
cushions just beneath the stomata; under humid conditions a few grow out
through the openings and develop into branched conidiophores (sporangio-
phores). Each has three to six main branches, and they branch again. The
terminal branches end in two to four short, slender sterigmata, each of which
produces a single multinucleate spore. With moisture, each nucleus with
adjacent protoplasm is organized into a swarm spore, motile with two cil-
ia. They swim around for a while, then settle down, absorb their cilia, and
put out a germ tube. If they happen to be on the upper side of a leaf, nothing
happens; if on the lower surface, the germ tube may reach a stoma and start
an infection.
Toward the end of the growing season thick-walled resting spores, oospores,
are produced in intercellular spaces of the infected leaves. These are set free
in spring by disintegration of host tissue, are rain-splashed to other vines,
and germinate by production of a short, unbranched hypha bearing a single
large sporangium, to start the cycle anew.
Control. Copper sprays are effective. Apply bordeaux mixture immediately
before and just after blooming; repeat 7 to 10 days later and possibly when
fruit is half grown. Destroy fallen leaves by burning.
Pseudoperonospora
Like Plasmopara but with branches of sporangiophores forming more or less acute
angles; tips more acute.
Pseudoperonospora celtidis. Downy Mildew of hackberry.
Pseudoperonospora cubensis. Downy Mildew of cucurbits, destructive
to cucumber, muskmelon, and watermelon, particularly along the Atlantic
seaboard and the Gulf Coast, occasional on gourd, pumpkin, and squash. The
disease was rst noted in 1889 in New Jersey, and in 1896 destroyed most of
the cucumbers on Long Island. Irregular yellow spots appear on upper leaf
surfaces, often on leaves nearest the center of the hill. The lesion is brown
on the opposite side, covered with a purple growth in rain or dew. The whole
leaf may wither and die, with the fruit dwarfed to nubbins and of poor avor.
The fungus does not live in the soil and is not prevalent in the North until
July or August. It winters in greenhouses or comes up from the South by
degrees. Sporangia are spread by wind and cucumber beetles. The disease is
favored by high humidity, but temperatures need not be as cool as for other
downy mildews.
Control. Resistant cucumbers are of rather poor quality. Cantaloupe varieties
Texas Resistant No. 1 and Georgia 47 combine resistance to aphids with
resistance to downy mildew.
Sclerospora
Oospore wall conuent with that of oogonium; sporangiophore typically stout with
heavy branches clustered at apex; mycelium intercellular, with small, knoblike, un-
branched haustoria; germination by germ tube or swarmspores. Common in moist tropic
regions on corn, millet, sorghum, and sugar cane.
DOWNY MILDEWS 283
Sclerospora farlowii. Downy Mildewof Bermuda grass, in the Southwest.
Short, black, dead areas prune off tips of leaves without serious damage to
grass. Tissues are lled with thick-walled, hard oospores.
Sclerospora graminicola on cereals.
Sclerophthora macrospora (formerly Sclerospora macrospora). Downy
Mildew of oats, crazy top of corn, wheat, barley, St. Augustinegsrass, Ken-
tucky bluegrass and wild grasses. Plants bunch owing to shortening of intern-
odes.
FAIRY RINGS
Several species of mushrooms growing in circles in lawns and golf greens
cause a condition known as fairy ring, rather common when the soil is quite
moist and contains a superabundance of organic matter. Less commonly,
some of these mushrooms are responsible for a poor condition of other herba-
ceous plants and of roses. The chief symptom in turf is the appearance of
continuous or interrupted bands of darker green, due to the fungus myceli-
um breaking down organic matter into products easily assimilated by grass
roots. Following the zone of stimulated growth there may be a zone of dying
grass due to temporary exhaustion of nutrients, or to toxic substances from
the mushroom mycelium, or because a layer has developed that is rather
impervious to water. The green rings are more conspicuous on underfertil-
ized lawns, and their presence can sometimes be masked by adequate fer-
tilization. Breaking off the mushrooms, possibly spiking the sod, is all the
control ordinarily recommended.
The following species are merely representative of the Basidiomycetes found
in fairy rings. They are in the order Agaricales, family Agaricaceae.
Cyathus stercoreus (Birds Nest Fungus). Fairy Ring on turf.
Lepiota morgani. On turf and also in rose greenhouses, causing poor
growth. The caps are 2 to 12 inches across, white with scattered brown
scales; esh white; gills green when mature, spores green turning yellow,
stem bulbous at base with a large ring (annulus). Poisonous, though other
members of this genus, also causing fairy rings, are edible.
Marasmius oreades. Cap 2 inches or less, convex to plane, thin, tough, with-
ering but not decaying; gills free from stem; spores white. Edible.
Psalliota (Agaricus) campestris. Cap 1 1/2 to 3 inches; white, silky, nearly
at; esh white to pinkish; gills pink, then brown; spores brownish purple;
stem white, with a ring when young. Edible.
Other Basidiomycetes found on lawns in moist weather include puffballs,
which are very good eating when white and rm inside, and birds nest fungi,
which are tiny cups lled with seed, resembling a nest of eggs.
FAIRY RINGS 285
Trechispora
Trechispora alnicola. Blight, Fairy Ring of Kentucky bluegrass.
FRUIT SPOTS
Many fruit blemishes are symptoms of rot diseases and are treated under
Rots; others are due to physiological disturbances; a few others, limited to
fruits and known primarily as fruit spots or specks, are included here.
Aureobasidium
Deuteromycetes
Yeast-like growth characteristics.
Aureobasidium pullulans. Fruit Russet on apple.
Cribropeltis
Brown mycellium, branches profusely; black, irregularly circular pycnidia; simple,
hyaline, clavate conidiophores; pale, oblong, straight or slightly curved conidia.
Cribropeltis citrullina. Fly Speck of watermelon fruits.
Zygophiala
Blotches.
Zygophiala jamaicensis (Schizothyrium pomi). Fly Speck on apple.
Helminthosporium
Blights.
FRUIT SPOTS 287
Helminthosporium papulosum. Black Pox on apples and pears in eastern
states. Fruit spots are small, sunken, dark, scattered in profusion over the
surface. Blackish papules on bark are followed by a pitted or scaly condition.
Spray with sulfur (except at high temperatures).
Microthyriella
Ascomycetes, Hemisphaeriales
Vegetative mycelium lacking; stromata with radial structure appearing as black super-
cial dots on leaves or stems.
Microthyriella rubi (see Schizothyrium pevexiguum). Fly Speck of pome
fruits, general on apple, also on pear, quince, citrus fruits, banana, Japanese
persimmon, plum, blackberry, raspberry, and grape.
Schizothyrium pevexiguum (formerly Microthyriella rubi). Fly Speck of
pome fruits, general on apple, also on pear, quince, citrus fruits, banana,
Japanese persimmon, plum, blackberry, raspberry, and grape. The pathogen
has long been recorded as Leptothyrium pomi, but this is apparently a mis-
conception. The anamorph state is Zygophiala jamaicensis, originally iso-
lated from banana and recently reported as causing a greasy blotch of car-
nations. Flyspeck is often associated with sooty blotch on apples, but the
two diseases are distinct. Flyspeck looks like its name, groups of 6 to 50
very small, slightly elevated, supercial black dots connected with very ne
threads. Spots may extend entirely around blackberry canes and shoots.
Mycosphaerella
Anthracnose.
Mycosphaerella pomi. Brooks Fruit Spot, Phoma Fruit Spot. Quince
Blotch, of apple and quince, most prevalent in northeastern states. Spots
appear on fruits in July or early August, deeper red on the colored face
of apples, darker green on the lighter surface. They are irregular, slightly
sunken, more abundant near the calyx end of the fruit, usually with centers
ecked with black. The symptoms on quince are more of a blotch than a def-
inite spot.
Rhodotorula
Rodotorula glutinis. Fruit Russet on apple.
GALLS
Galls are local swellings, hyperplastic enlargements of plant tissue due to
stimulation from insects, bacteria, fungi, viruses, and occasionally physio-
logical factors. Crown gall, a common and serious problem, is discussed
under Bacterial Diseases. Cedar galls are treated under Rusts. See Black
Knot for hypertrophy of plum branches.
Exobasidium
Basidiomycetes, Exobasidiales
Mycelium intercellular with branched haustoria entering host cells; basidia extend above
the layer of epidermal cells much like the layer of asci in Taphrina; each basidium bears
two to eight basidiospores. Species cause marked hypertrophy in the Ericaceae.
Exobasidium vaccinii. Leaf Gall, widespread on ame azalea.
Exobasidium burtii. Leaf Gall, Yellow Leaf Spot on azalea and rhodo-
dendron.
Exobasidium camelliae. Camellia Leaf Gall on camellia in the South-
east, more common on sasanqua than on japonica. Symptoms are a strik-
ing enlargement and thickening of leaves and a thickening of stems of new
shoots. Diseased leaves are four or more times as wide and long as normal
leaves, very thick and succulent. Color of the upper surface is nearly normal,
but the underside is white with a thin membrane that cracks and peels back
in strips or patches exposing the spore-bearing layer. There is seldom more
than one diseased shoot on a stem, and not many on the whole bush; so the
disease does not cause serious damage.
Control. Handpicking of affected parts, searching carefully for diseased
leaves at base of new growth, removing them before spores are formed,
keeps sporadic infection at a minimum. Spraying with a low-lime bordeaux
may be effective but is seldom necessary.
Exobasidium oxycocci. Cranberry Rose Bloom, Shoot Hypertro-
phy on cranberry, and manzanita. The disease appears in cranberry bogs
soon after water is removed in spring. Bud infection results in abnormal
lateral shoots with enlarged, swollen, pink or light rose distorted leaves
that somewhat resemble owers. Excessive water supply promotes the dis-
ease. Remove water early in spring. If necessary, spray with bordeaux
mixture.
Exobasidium rhododendri. Rhododendron Leaf Gall. Large vesicular
galls, especially on Rhododendron catawbiense and R. maximum.
Exobasidium symploci. Bud Gall on sweetleaf.
Exobasidium uvae-ursi. Shoot Hypertrophy of bearberry.
Exobasidium vaccinii. Azalea Leaf Gall, Red Leaf Spot, Shoot Hyper-
trophy of andromeda, arbutus (A. menziesii), bearberry, blueberry (fruit
green spot), box sandmyrtle, chamaedaphne, cranberry, farkleberry, huck-
leberry, ledum, leucotho, manzanita, and rhododendron. On azaleas and
other ornamentals the galls are bladder-shaped enlargements of all or part
of a leaf, sometimes a ower bud (see Fig. 3.21). They are white or pink,
soft and succulent when young, brown and hard with age. This is seldom
a serious disease but in wet seasons, particularly in the South, and in shaded
gardens, the number of galls may become rather alarming. On cranber-
ries and blueberries the gall is a small, round, red blister in the leaf, with
spores packed in a dense layer on the underside. The fungus is systemic in
blueberries, fruiting on the leaves in June and July.
Control. Handpick and destroy galls as they appear. Spraying is seldom
required for cranberries and other fruits.
Exobasidium vaccinii-uliginosi. Shoot and Leaf Gall, Witches Broom
of rhododendron, manzanita, and mountain heath. An excessive number of
twigs is formed on infected branches. Leaves are yellowish white covered
with a dense mealy fungus growth. The mycelium penetrates the whole
plant so that it is wiser to remove the shrub than to attempt remedial mea-
sures.
Fusarium
Rots.
Fusarium decemcellulare (Teleomorph, Nectria rigidiuscula). Gall on
midge.
GALLS 291
Figure 3.21 Azalea Leaf Gall
Kutilakesa
Sporodochia erumpent, pale olive-green, cushion-shaped; similar to Kutilakesopsis but
differs by having larger two-celled conidia; teleomorph state is Nectriella.
Kutilakesa pironii. Stem and Leaf Gall, Cankers on croton, zebra plant,
and Clorodendron.
Nocardia
Actinomycetales
Related to bacteria with mycelial laments breaking up into rod forms.
Nocardia vaccinii. Blueberry Bud-Proliferating Gall, rst observed in
Maryland in 1944, described as a new species in 1952. Galls, similar to
crown gall, are formed at the soil line. Abnormal buds abort at an early stage
or grow into weak shoots, 1 to 6 inches high, forming a witches broom
effect.
Phoma, Phomopsis
Blights.
Phoma sp. or Phomopsis sp. StemGall on winter jasmine, privet, forsythia,
and rose, at scattered locations. Both pathogens have been reported causing
roundish, rather rough stem enlargements on ornamentals. It has not been
determined whether more than one fungus is involved.
Plasmopara
Downy Mildew.
Plasmopara halstedii. Basal Gall on sunower.
Protomyces
Archiascomycetes, Taphrinales
Protomyces gravidus. Stem gall on ragweed.
Protomyces macrosporus. Leaf gall on hedge parsley (Torilis sp.).
GALLS 293
Sphaeropsis
Cankers.
Sphaeropsis tumefaciens. Canker and Gall on Carissa.
Synchytrium
Chytridiomycetes, Chytridiales
Mycelium lacking; thallus converted into a soros with a membrane, at maturity function-
ing in entirety as a resting sporangium or divided to form many sporangia in a common
membrane; zoospores with one cillum at posterior end. Various species cause excres-
cences on leaves and fruit; potato wart.
Synchytrium anemones. Leaf Gall, Flower Spot of anemone and thalic-
trum. Flowers are spotted, distorted, dwarfed, and may fall. Red spots are
formed on leaves and stems.
Synchytrium aureum. Red Leaf Gall, False Rust on many plants, 130
species in widely separated genera, including calypha, artemisia, clintonia,
delphinium, geum, golden-glow, marsh-marigold, and viola. Pick off and
burn affected parts.
Synchytrium endobioticum. Potato Wart, Black Wart of potatoes,
a warty hypertrophy of tubers. A European disease wart was found in
1918 in backyard gardens in mining towns of Pennsylvania, Maryland, and
West Virginia. Diseased tubers had apparently been brought in by immi-
grants. A strict quarantine was placed on infested districts, and there has
been no spread to commercial potato elds. The disease shows as prominent
outgrowths or warts originating in the eyes, varying from the size of a pea
to that of the tuber itself. Numerous yellow sporangia are released into the
soil by decay of the malformed tissue. The disease, which may affect other
species of Solanum, is spread by contaminated soil or infected tubers. Buds
and adventitious shoots of tomato are infected below the soil line.
Control. By 1953 potato wart had been eradicated from more than half of the
1112 infested gardens in Pennsylvania. The plan called for applying copper
sulfate the rst year, keeping the land clean and cultivated, applying lime the
next year, growing vegetables the third year, and going back to potatoes the
fourth year to test results.
Synchytriumvaccinii. Red Leaf Gall on cranberry, azalea, chamaedaphne,
gaultheria, and ledum, from New Jersey northward. On cranberry the disease
appears just before blossoms open. Buds, owers and young leaves are cov-
ered with small, red, somewhat globular galls about the size of birdshot;
affected shoots bear no fruit. The disease is erratic in appearance but is most
frequent in bogs that have excessive or uneven water supply.
Synchytrium sp. Stem Gall on castor bean, in Texas. Small red galls on
stems, petioles, and leaves of seedlings.
LEAF BLISTER
AND LEAF CURL DISEASES
A single genus, Taphrina, is responsible for most of the hyperplastic (over-
growth) deformities known as leaf blister, leaf curl, or, occasionally, as pock-
ets.
Taphrina
Archiascomycetes, Taphrinales
Parasitic on vascular plants, causing hypertrophy. Asci in a single palisade layer, not
formed in a fruiting body; hyphal cells become thin-walled chlamydospores; on germi-
nation the inner spore protrudes from the host and is cut off by a septum to form an
eight-spored ascus, which may become many-spored by budding or the ascospores.
Taphrina spp. Maple Leaf Blister. Leaves after expanding in spring show
dark spots, shrivel, and fall. The disease may be locally epidemic; it is more
common in shaded locations.
Taphrina aceris. Western Maple Leaf Blister.
Taphrina aesculi. Leaf Blister of California buckeye; yellow turning to dull
red; witches brooms formed.
Taphrina australis. American Hornbean Leaf Curl.
Taphrina caerulescens. Oak Leaf Blister on various oak species, with
red oak particularly susceptible but often defoliating and sometimes killing
water, willow, laurel, and live oaks in the South. Blisters start on young par-
tially grown leaves as gray depressed areas on the undersurface, convex and
yellow on the upper surface. Individual blisters are 1/4 to 1/2 inch across but
often become conuent, causing the leaf to curl. Ascospores are borne on the
surface of the blistered area. The disease is most serious in a cool wet spring.
Control. A single dormant eradicant spray, before the buds swell, controls
the disease; later sprays are ineffective.
Taphrina carnea. Birch Red Leaf Blister.
Taphrina castanopsidis. California Chinquapin Leaf Blister.
Figure 3.22 Peach Leaf Curl; deformed leaf; palisade layer of asci formed on curled portion; germinating spore
Taphrina communis. Plum Pockets, common on American plums;
T. pruni, on European species, not in United States; T. prunisubcorda-
ta, in western United States. Leaves, shoots and fruits become puffy and
enlarged into reddish or white swollen bladders. Fruits are sometimes ten
times the size of normal plums. Most garden plums are of foreign origin
and not susceptible to the American species of Taphrina. Bordeaux mixture
applied in spring before ower buds open gives satisfactory control.
Taphrina coryli. Hazelnut Leaf Blister.
Taphrina deformans. Peach Leaf Curl, general on peach, also on nec-
tarine and almond but not on apricot. This is an old disease, known in the
United States for well over a century but not quite so important since 1900,
when a control was worked out. Young leaves are arched and reddened, or
paler than normal as they emerge from the bud, then much curled, puckered,
and distorted, greatly increased in thickness (Fig. 3.22). Any portion or the
entire leaf may be curled, and one or all leaves from a bud. The leaves often
look as if a gathering string had been run along the midvein and pulled tight.
Leaves may drop, lowering vitality of tree, with partial or total failure to set
fruit, and increasing chances of winter injury. Young fruits may be distorted
or cracked. Defoliation for several seasons kills tree outright.
The fungus has no summer stage, and the asci are formed not in a fruiting
body but in a layer over infected surfaces, giving them a silvery sheen. Before
leaves fall, ascospores are discharged from this layer, and land on bark or
twigs and bud scales, there to germinate by budding into yeastlike spores,
which remain viable over winter, sometimes for 2 years. In spring they are
washed by rain to opening leaf buds.
Control. One spray during the dormant season gives effective control. This
is best applied just before the buds swell, but can be done any time after leaf
fall in autumn when the temperature is above freezing. Applications after the
buds swell have little effect.
LEAF BLISTER AND LEAF CURL DISEASES 297
Taphrina faulliana. Leaf Blister of Christmas fern; T. licina, on sensitive
fern; T. struthiopteridis, on ostrich fern.
Taphrina ava. Yellow Leaf Blister of gray and paper birches in north-
eastern states.
Taphrina japonica (T. macrophylla). Leaf Curl on red alder. Young leaves
are enlarged to several times normal size and curled. They dry up after
ascospore discharge, and a new crop of healthy leaves is formed.
Taphrina populina. Leaf Blister, Yellow on poplar.
Taphrina populina. Poplar Yellow Leaf Blister. Conspicuous blisters,
small to large, an inch or more in diameter, are brilliant yellow on the con-
cave side when the asci are fully developed; later the color changes to brown.
Taphrina robinsoniana, T. occidentalis, T. alni. Catkin Hypertrophy of
alder. Scales of catkins enlarge and project as reddish curled tongues covered
with a white glistening layer. Infection can be reduced with a lime sulfur
spray.
Taphrina sacchari. Maple Brown Leaf Blister.
Taphrina ulmi. Elm Leaf Blister. Very small blisters on elm leaves. Dust-
ing nursery trees with sulfur has helped.
Taphrina weisneri. Cherry Witches Broom, Leaf Curl on wild and cul-
tivated cherries, T. avorubra, on sand cherry; T. ectans, on western wild
cherry; T. farlowii, leaf curl and fruit pockets on eastern wild cherry; T. con-
fusa, on chokecherry; T. thomasii, witches broom of cherry-laurel in Cali-
fornia.
LEAF SCORCH
According to the dictionary scorching means to heat so as to change color
and texture without consuming. Sometimes leaves are literally scorched in
summer heat, and sometimes symptoms caused by fungi resemble those of
a heat scorch. This section includes some of the latter.
Ceratocystis
Cankers.
Ceratocystis paradoxa. Black Scorch, Bud Scorch, Heart Rot of
coconut, Canary, Washington, and Guadaloupe palms, also causing a pineap-
ple disease in the tropics. The most striking symptom is a black, irregular,
necrotic condition of the leaf stalk. The tissues look as if they had been
burned, whence the name black scorch. Furled pinnae of leaf fronds show
pale yellow spots with broad margins that later converge and turn black;
infection spreads rapidly, and in severe cases the heart leaves dry up. The
heart rot discolors trunk tissues and rots the pithy material between cells.
Infection is through wounds during periods of relatively high humidity, or
through roots, or sometimes through uninjured fruit strands, petioles, or
pinnae. Palms with vitality lowered, as when the normal crown of leaves
has been reduced but the water supply to the leaves is not reduced, are most
susceptible.
Control. Destruction of infected parts seems to be the chief control measure.
It is easier to bury than to burn palm trunks.
Curvularia
Blights.
Curvularia sp. Leaf Scorch on pecan.
LEAF SCORCH 299
Diplocarpon
Blackspot.
Diplocarpon earlianum. Strawberry Leaf Scorch, general where straw-
berries are grown but more prevalent in the South. Dark purplish spots about
1/4 inch in diameter are scattered profusely over upper surface of leaves in
all stages of development. Later the spots enlarge to scorch wide areas of
the leaf, and black fruiting bodies give a tar spot appearance. Scorch spots
always lack the white centers so characteristic of Mycosphaerella leaf spot
on strawberry. Lesions are found on petioles, stolons, and fruit stalks as well
as leaves. If the fruit stems are girdled, owers or young fruits die. Rarely
the disease appears on green berries as a supercial red or brown discol-
oration and ecking. Spores, produced in quantity in acervuli on lesions, are
distributed by birds, insects, and pickers on tools and clothing. The fungus
winters in old leaves. Teleomorph and anamorph states are both produced in
spring, and repeated infections occur throughout the summer in moist weath-
er.
Control. Remove all old leaves when setting plants in spring. Spray with
bordeaux mixture at 10-day intervals, starting in January in Louisiana, late
February in North Carolina. Fairly resistant varieties include Catskill, Mid-
land, Fairfax, Howard 17, Blakemore, Southland.
Epicoccum
Leaf Spots.
Epicoccum sp. Leaf Scorch on pecan.
Fusarium
Rots.
Fusarium sp. Leaf Scorch on pecan.
Hendersonia
Deuteromycetes; Coelomycetes
Pycnidia dark, separate, globose, ostiolate, immersed then usually erumpent;
conidia dark, several-celled, elongate to fusoid; saprophytic or parasitic.
Hendersonia opuntiae. Scorch, Sunscald, common and serious on prick-
ly pear cactus (Opuntia). Segments turn reddish brown and die; centers are
grayish brown and cracked.
Pestalotia
Blights.
Pestalotia sp. Leaf Scorch on pecan.
Pseudopezicula
Hyaline, gelatinous apothecia containing paraphyses and 20 - 80 asci; asci contain 4
reniform, binucleate ascospores; ve-spored asci rarely observed.
Pseudopezicula tetraspora. Leaf Scorch of grapevines.
Septoria
Blights.
Septoria azaleae. Azalea Leaf Scorch, Leaf Spot. Small, yellowish,
round spots enlarge irregularly, turn reddish brown, with dark brown cen-
ters. Leaves fall prematurely; black fruiting bodies are produced in fallen
leaves. The disease is most severe in greenhouses in fall and winter and
under high humidity.
LEAF SCORCH 301
Stagonospora
Pycnidia dark, separate, supercial, or erumpent, globose, ostiolate; conidiophores short;
conidia hyaline, typically with three or more cells, cylindrical to elliptical; parasitic or
saprophytic.
Stagonospora curtisii. Narcissus Leaf Scorch, Red Blotch of Amaryl-
lis, Red Leaf Spot, Red Fire Disease, also on crinum, eucharis, hymeno-
callis, leucojum, nerine, sternbergia, vallota, and zephyranthes.
Leaf tips of narcissus are blighted for 2 or 3 inches as in frost injury and
separated off from healthy basal portions of leaves by a denite margin or
yellow area. Spores formed in pycnidia in the dead area furnish inoculum for
secondary infection, which consists of lesions in lower portions of leaves,
minute water-soaked or yellowish spots becoming raised, scabby, and red-
dish brown. Flower stalks may be spotted; brown spots appear on petals.
Bulbs suffer loss in weight due to killing of foliage a month or two before
normal dying down. All types may be infected but the most susceptible vari-
eties are in the Leedsii and Polyanthus groups. The fungus was described on
narcissus in 1878 but was not considered a threat to it, nor was it known to
be connected with amaryllis red blotch before 1929.
On amaryllis or hippeastrum red spots are formed on leaves, ower stems,
and petals. On foliage the spots are bright red to purplish, small at rst but
often increasing to 2 inches. Leaf or ower stalks are bent or deformed at
the point of attack. This disease should not be confused with red disease
caused by mites. The spores are variable in size and number of cells, one to
six. They are embedded in a gelatinous matrix and are disseminated in rain.
The fungus apparently winters in or on bulbs, infecting new leaves as they
grow out in spring.
Control. Treat suspected narcissus bulbs before planting. Control secondary
infection in the eld with bordeaux mixture. Discard seriously diseased
amaryllis bulbs; remove infected leaves and bulb scales; avoid syringing and
heavy watering.
LEAF SPOTS
Leaf spots are the most prevalent of plant diseases, so common we seldom
notice them, and rightly so, for if we should attempt to control all the miscel-
laneous leaf spots that appear in a small suburban garden in a single season,
we would quickly go mad. A typical leaf spot is a rather denitely delimited
necrotic lesion, often with a brown, sometimes white, center and a darker
margin. When the spots are so numerous they grow together to form large
dead areas, the disease becomes a blight, or perhaps a blotch, or scorch. Cer-
tain types of lesions are called anthracnose, spot anthracnose, blackspot. All
of these have been segregated out in their different sections. What is left is
a very large collection of names.
The genus Septoria, for instance, has about 1000 species, Mycosphaerella
500, Cercospora 400, chiey identied by the hosts on which they appear.
Cercospora beticola is so named because it causes a leaf spot of beet, C. apii
for its celery host. Species recorded in this country as causing a denite
disease are listed under their respective hosts. They are not repeated here
unless the leaf spot is of some importance or there is some useful information
that can be added to the name.
Most leaf spot diseases ourish in wet seasons. A comparative few may be
important enough to call for control measures other than general sanitation.
Adequate protection usually means several applications of fungicides, and
the cost of spraying trees and shrubs must be balanced against the expected
damage. Calling in a tree expert with high-pressure apparatus is often an
expensive proposition. If the budget is limited, it is more important to have
an elm sprayed for elm leaf beetles, which cause defoliation every season,
than for elm black spot, which may be serious in only one year out of three or
four. When it comes to rose blackspot (no relation to elm black spot), weekly
protection with a fungicide is necessary, but to save labor it can be combined
with insecticides.
LEAF SPOTS 303
Actinothyrium
Pycnidia supercial, globose, with a more or less mbriate shield; spores liform, hya-
line.
Actinopelte dryina (see Tubakia dryina). On oak.
Actinothyrium gloeosporioides (see Tubakia dryina). On oak. Leaf Spot
on sassafras.
Tubakia dryina (formerly Actinopelte dryina). On oak. Very small dark
spots between veins. Conspicuous in midsummer but not serious.
Tubakia dryina (formerly Actinothyrium gloeosporioides). Leaf Spot on
sassafras.
Alternaria
Blights.
Alternaria alternata. Leaf Spot of Calathea spp.
Alternaria alternata (formerly Alternaria fasciculata). Leaf Spot on rose-
acacia and asclepiodora.
Alternaria alternata (formerly Alternaria tenuis). Leaf Spot of magnolia,
hibiscus, clarkia, and many ornamental and other hosts. The fungus is a gen-
eral saprophyte and an occasional weak parasite. It discolors beet, chard, and
spinach seed.
Alternaria angustiovoidea. Leaf Spot and Blight of leafy spurge.
Alternaria brassicae (with large spores) and A. brassicicola (with small
spores). Black Leaf Spot of crucifers, cabbage, Chinese cabbage, col-
lards, turnip, garden cress, mustard greens, radish, and horseradish; Head
Browning leaf and pod spot of cauliower; Damping-off, Wire-stem of
seedlings.
Seedlings are subject to pre- or post-emergence damping-off, with dark
brown to black sunken spots on cotyledons, narrow dark spots on stems,
followed by wire-stem, a blackening toward the base. Leaf spots are small,
circular, yellowish, enlarging in concentric circles with a sooty black color
from the spores. In storage the spots unite to form a moldy growth over the
entire leaf. On seed pods, spots are purplish at rst, later brown; in moist
weather entire pods may be infected. Cauliower infection is a browning
of the head, starting at the margin of an individual ower or cluster. Spores
are blown, splashed by tools, spread on feet of men and animals. Seed
bears spores externally, mycelium internally. Wounds are not necessary for
infection.
Control. Hot water treatment of seed, 122F for 30 minutes, is fairly effec-
tive. Use long rotation for cauliower, avoiding all other crucifers in inter-
mediate years.
Alternaria brassicicola. Leaf Spot on Thlaspi.
Alternaria brassicicola (formerly Alternaria oleracea). Cabbage Leaf
Spot, occasional on crucifers. Has been confused with A. brassicicola.
Alternaria catalpae. Catalpa Leaf Spot, widespread in rainy seasons.
Small, water-soaked spots, up to 1/4 inch, appear over the leaf; they turn
brown and sometimes drop out leaving shot holes; there is more or less defo-
liation. The fungus is sometimes secondary following bacterial infection or
midge infestation. Rake up and burn fallen leaves.
Alternaria chrysanthemi (see Alternaria leucanthemi). Leaf Spot on shas-
ta daisy, and Canada thistle.
Alternaria citri. Cherry Leaf Spot, occasional, more often a rot of citrus
fruits.

Rots.
Alternaria fasciculata (see Alternaria alternata). Leaf Spot on rose-acacia
and asclepiodora.
Alternaria leucanthemi (formerly Alternaria chrysanthemi). Leaf Spot on
shasta daisy, and Canada thistle.
Alternaria longipes. Brown Spot of tobacco, including ornamental ow-
ering tobacco. Small spots on lower leaves rapidly enlarge and turn brown.
The fungus winters on old stalks, which should be removed and burned.
Alternaria oleracea (see Alternaria brassicicola). Cabbage Leaf Spot,
occasional on crucifers.
Alternaria panax. Leaf Spot of schefera, Dizygotheca, and Tupidanthurs.
Alternaria passiorae. Brown Spot of passion ower. Minute brown leaf
spots, enlarging to an inch across are concentrically zoned with various
shades of brown. Dark green water-soaked spots on fruit turn brown; the
fruit shrivels, but the spots stay rm.
Alternaria polypodii. Fern Leaf Spot. Brown, circular to ovate, concen-
trically zonate spots are formed along margins of fronds. Chains of spores
are spread by syringing or air currents. Keep foliage dry; remove and burn
diseased leaves.
LEAF SPOTS 305
Alternaria raphani. Radish Leaf Spot. Yellow spots with black sporula-
tion, often with centers dropping out. Also occurs on turnip.
Alternaria sonchi. Leaf Spot of lettuce, escarole, endive, and chicory.
Alternaria tagetica. Leaf Spot of marigold.
Alternaria tenuis (see Alternaria alternata). Leaf Spot of magnolia, hibis-
cus, clarkia, and many ornamental and other hosts.
Alternaria tenuissima. Leaf Spot on blueberry.
Alternaria tenuissima (formerly Alternaria tomato). Nailhead Spot of
tomato, a leaf, stem, and fruit spot. On leaves and stems the disease is much
like early blight (see A. solani under Blights) with small dark brown spots
with yellow margins. But on fruit the disease is quite different. Very small
tan spots, 1/16 to 1/8 inch in diameter, become slightly sunken, with gray-
ish brown centers and darker margins. Spores produced abundantly on fruit
and foliage are spread by winds and splashing rain. Treat seed and spray as
for early blight. Varieties Marglobe, Pritchard, Glovel, and Break ODay are
quite resistant to nailhead spot. The same fungus causes ghost spot of apple.
Alternaria tomato (see Alternaria tenuissima). Nailhead Spot of tomato,
a leaf, stem, and fruit spot.
Alternaria sp. Leaf Spot of schefera, and umbrella tree.
Amerosporium
Pycnidia supercial, discoid to cupulate, hairy; spores one-celled, hyaline.
Amerosporium trichellum(see Colletotrichum trichellum). Leaf Spot and
Stem Spot on English ivy.
Colletotrichum trichellum (formerly Amerosporium trichellum). Leaf
Spot and Stem Spot on English ivy. In some cases stems are girdled,
causing collapse and death.
Annellophora
Conidiophores brown, simple, slender, elongating by successive proliferations through
conidial scars; conidia brown, multiseptate, obclavate to fusoid.
Annellophora phoenicis. Leaf Spot of date palm.
Aristastoma
Pycnidia brown, globose, erumpent, separate, with dark brown setae near ostiole;
conodiophores short, simple; conida hyaline, several-celled.
Aristastoma oeconomicum. Zonate Leaf Spot of cowpea, kidney bean.
Aristastoma sp. Leaf Spot on desert-rose.
Ascochyta
Blights.
Ascochyta abelmoschi (possibly identical with A. phaseolorum). Leaf
Spot, Pod Spot, Stem Spot of okra. Dark, small, water-soaked spots
slowly enlarge, turn brown, with many large black pycnidia in concentric
rings in dead tissue. Young okra pods are severely infected, and the myceli-
um grows into the seed.
Ascochyta althaeina (see Phoma exigna). Leaf Spot of hollyhock, rose-
mallow.
Ascochyta armoraciae. Leaf Spot of horse-radish.
Ascochyta aspidistrae. Aspidistra Leaf Spot. Large, irregular pale spots
on leaves.
Ascochyta asteris (see Phoma exigna). Leaf Spot of China aster.
Ascochyta boltshauseri. (see Stagonosporopsis hortensis). Leaf Spot,
Pod Spot of beans, on snap, kidney, lima, and scarlet runner beans, report-
ed in Oregon.
Aschochyta. Leaf Spot on big bluestem, little bluestem (both species of
Andropogon and on indiangrass.
Ascochyta cheiranthi. Leaf and Stem Spot of wallower. Grayish spots
up to 1/2 inch long, may girdle stems. Leaf spots are circular to elongate,
brown with darker brown margins. Dark pycnidia contain hyaline, two-celled
spores. Leaves wilt and fall; potted plants may be infected. Keep greenhouse
on the dry side.
Ascochyta clematidina. Clematis Leaf and Stem Spot, widespread. On
out-door plants stems are infected near the ground and are often girdled,
upper portions dying back. Spores for initial infection probably come from
LEAF SPOTS 307
pycnidia on stumps of old stems. Leaf spots are more common in greenhous-
es, small, water-soaked, then buff with reddish margins. Remove and destroy
infected leaves and stems.
Ascochyta compositarum. Leaf Spot on aster, eupatorium, silphium, and
sunower.
Ascochyta cornicola. Dogwood Leaf Spot.
Ascochyta cypripedii. Cypripedium Leaf Spot, reported on orchid from
Wisconsin. Leaf lesions are narrow, brownish, with a dark brown border.
Ascochyta juglandis. Walnut Ring Spot. Very small, round, brown leaf
spots between veins, ringed with targetlike ridges. The disease is unimpor-
tant in trees sprayed for walnut blight.
Ascochyta lycopersici (Didymella lycopersici) (see Phoma lycopersici)
Leaf Spot, Ascochyta Blight of tomato, eggplant, and potato.
Ascochyta phaseolorum (see Phoma exigua). Leaf Spot of snap beans.
Ascochyta pisi. Leaf Spot, Pod Spot of pea. General, but rare in the
Northwest. One of three species causing the disease complex known as
Ascochyta blight (also see Blights). Foliage spots are circular to irregular,
pinhead size to 1/2 inch. Stem lesions, at nodes or base, are brown to purplish
black. Brown pycnidia exude spore tendrils in wet weather.
Phoma exigua (formerly Ascochyta althaeina). Leaf Spot of hollyhock,
rose-mallow.
Phoma exigua (formerly Ascochyta asteris). Leaf Spot of China aster.
Spray foliage with bordeaux mixture.
Phoma exigua (formerly Ascochyta phaseolorum). Leaf Spot of snap
beans. Recent isolation and inoculation studies indicate that the Ascochyta
leaf blights of hollyhock, okra, pepper, eggplant, and tomato are all caused
by strains of the bean pathogen.
Phoma lycopersici (formerly Ascochyta lycopersici (Didymella lycopersi-
ci)). Leaf Spot, Ascochyta Blight of tomato, eggplant, and potato. Brown
spots with concentric rings are formed on leaves and stems, sometimes
cankers at base of young stems. Black pustules in center of spots discharge
spore tendrils in wet weather. The fungus winters in old plant refuse, is
a weak parasite, and is ordinarily too unimportant for control measures.
Stagonosporopsis hortensis (formerly Ascochyta boltschauseri). Leaf
Spot, Pod Spot of beans, on snap, kidney, lima, and scarlet runner beans,
reported in Oregon. Spots on leaves and pods are dark to drab, zonate; light
to dark brown pycnidia are numerous.
Asteroma
Pycnidia globose with a radiate subicle, a compact, crustlike growth of mycelium under-
neath; without an ostiole; spores hyaline, one-celled.
Asteroma garretianum. Black Spot on primrose.
Asteroma solidaginis. Black Spot, Black Scurf on goldenrod.
Asteroma tenerrimum. Black Spot on erythronium.
Asteromella (Stictochlorella)
Pycnidia smooth, with ostiole, densely gregarious in asteroma-like spots; spores hyaline,
one-celled.
Asteromella lupini. Leaf Spot on lupine.
Botrytis
Blights.
Botrytis fabae. Chocolate Leaf Spot on vetch.
Calonectria
See Cylindrocladium under Blights.
Calonectria colhounii. Leaf Spot on sentry palm.
Calonectria crotalariae. Leaf Spot on sentry palm.
Calonectria theae. Leaf Spot on sentry palm.
Cephaleuros
One of the green algae, possessing chlorophyll but not differentiated into root, stem, and
leaves; forming motile spores in sporangia.
Cephaleuros virescens. Algal Spot, Red Leaf Spot, Green Scurf in
the far South or in greenhouses on acacia, albizzia, ardisia, avocado, bixa,
LEAF SPOTS 309
bischoa, camellia, camphor-tree, cinnamon-tree, citrus, grevillea, guava,
jasmine, jujube, loquat, magnolia, mango, pecan, Japanese persimmon, priv-
et, rhododendron, viburnum.
On some hosts this is a disease of twigs and branches, which may be girdled
and stunted, covered with reddish brown hairlike fruiting bodies. On magno-
lia leaves velvety, reddish brown to orange, cushiony patches are formed, but
in the absence of sporangia (tiny globular heads on ne, dense reddish hairs)
the leaf spots remain greenish brown. Occasionally citrus fruits as well as
leaves are attacked.
The sporangia formed on the ne hairs germinate in moist weather, pro-
ducing zoospores that enter through stomata and form mycelium-like chains
of algal cells in host tissue. On twigs the alga invades outer cortical tissue,
which may swell abnormally, crack, and afford entrance to injurious fungi.
Weakened trees are most susceptible, and disease spread is most rapid in
periods of frequent and abundant rains. Twigs may die, and there may be
reduced yield of citrus fruit.
Control. Improve draining and other growing conditions; citrus trees sprayed
regularly with copper seldom have algal trouble. If it gets started, follow
cleanup pruning with a bordeaux mixture spray in December or January.
Repeat with bordeaux at start of rainy season or when red stage of the alga
is rst seen, and spray again 1 month later. A neutral copper may substitute
for bordeaux for the last two applications. The copper kills benecial insects
parasitic on scales, but the oil controls the scale insects.
Cephalosporium
Conidiophores slender or swollen, simple; conidia hyaline, one-celled, produced succes-
sively at the tip and collecting in a slime drop, produced endogenously in some species;
saprophytic or parasitic, some species causing vascular wilts of trees.
Cephalosporium apii. Celery Brown Spot, a new disease rst reported
from Colorado in 1943, later from New York and Ohio. Irregular light tan or
reddish brown shallow lesions are formed on celery leaf stalks, petioles, and
leaets. They may unite to make a scurfy brown streak up the inside of the
stalk and may develop transverse cracks. Utah and Pascal varieties are most
susceptible.
Cephalosporium cinnamomeum. Leaf Spot of nephthytis and syngoni-
um. Small circular to irregular spots, reddish brown with pale yellow bor-
ders enlarge, with centers becoming gray and papery. In severe cases leaves
turn yellow and die. Pick off infected leaves. Maintain low temperature and
humidity.
Cephalosporium dieffenbachiae. Dieffenbachia Leaf Spot. Small red
lesions with dark borders appear on young leaves. Spots sometimes run
together, and the whole leaf turns yellow and dies. Infection is often through
mealybug wounds. Avoid promiscuous syringing; keep temperature and
humidity low; control mealybugs, and ants that transport them.
Cercospora
Blights.
Cercospora abeliae. Abelia Leaf Spot, reported from Louisiana. Irregular
purple to brown spots; defoliation.
Cercospora abelmoschi (see Pseudocercospora abelmoschi). Leaf Spot on
okra, hibiscus.
Cercospora albo-maculans (Syn. Cercosporella brassicae) (see Pseudocer-
cospora capsellae). White Spot of turnip, Chinese cabbage, mustard, and
other crucifers, common in the Southeast.
Cercospora althaeina. Leaf Spot of hollyhock and abutilon. Spots circu-
lar, angular or irregular, 1.5 mm, olivaceous to grayish brown, with the dead
tissue falling out. The fungus winters in old plant parts.
Cercospora angulata. Leaf Spot on philadelphus, currant, owering cur-
rant, and gooseberry. Circular to angular spots, dingy gray centers, dark pur-
ple to nearly black margins.
Cercospora aquilegiae. Columbine Leaf Spot, reported from Kansas,
Wisconsin, Oregon. Spots circular to elliptical, reddish brown to nearly
black; fruiting is on both sides of the leaf.
Cercospora arachidicola (Mycosphaerella arachidicola, Teleomorph).
Peanut Early Leaf Spot. Spots light tan aging to reddish or dark brown
with a yellow halo, often conuent. Conidiophores on both sides of the leaf,
emerging from stomata or breaking through epidermal cells. Conidia color-
less to pale yellow or olive, with 5 to 12 cells. Control with sulfur-copper
dust.
LEAF SPOTS 311
Cercospora armoraciae. Horse-Radish Leaf Spot. Tan to dingy gray
lesions with yellow-brown margin; often slightly zonate.
Cercospora beticola. Cercospora Leaf Spot of beet, general on garden
and sugar beets, also on swiss chard, spinach. Brown ecks with reddish
purple borders become conspicuous spots with ash-gray centers and purple
margins. The brittle central tissue often drops out, leaving ragged holes. The
spots usually remain small but are often so numerous that foliage is killed.
If successive crops of leaves are lost, the crown of the beet root is elongated
and roughened. Leaf spotting is of little direct importance except in chard,
where foliage is used for greens. The beet root yield is reduced.
The grayish color of the spots is due to long, thin, septate conidia produced
on conidiophores protruded through stomata in fascicles or groups, com-
ing from a knotted mass of mycelium resembling a sclerotium. Conidia are
spread by rain, wind, tools, and insects. Infection is through stomata; disease
spread is most rapid under conditions of high humidity that keep stomata
open. Hot weather favors the disease.
Control. Crop rotation is highly important. In a small garden pick off the rst
spotted leaves.
Cercospora bougainvilleae (see Cercosporidium bougainvilleae). Leaf
Spot rst seen in Florida in 1962 and now the most important pathogen of
this host.
Cercospora brunkii. Geranium Leaf Spot, mostly in the South. Spots are
circular, light reddish brown with dark brown borders, sometimes coalescing
to kill entire leaf.
Cercospora calendulae. Calendula Leaf Spot. Spots run together to
blight and kill leaves; plants may be destroyed early in the season. Spores
enter through stomata of plants more than a month old.
Cercospora cannabina (see Pseudocercospora cannabina). Leaf Curl and
Wilt on hemp.
Cercospora cannabis. Leaf Spot on hemp.
Cercospora capsici. Pepper Leaf Spot, Stem-end Rot, common in the
Southeast, serious in rainy seasons. Spots 1/7 to 1 inch in diameter are rst
water-soaked then white with dark brown margins. Leaves turn yellow and
drop. The fungus grows through the pedicel into fruit, causing a rot of the
stem end. Loss of foliage exposes the fruit to sunscald. Spray or dust with
copper.
Cercospora circumscissa (Mycosphaerella cerasella, Teleomorph). Leaf
Spot, Shot Hole of apricot, plum, cherry, cherry-laurel, oriental cherry,
and chokecherry. Dead spots are somewhat larger than those caused by other
shot-hole fungi, but the damage is not serious.
Cercospora citrullina. Leaf Spot of watermelon, muskmelon, and other
cucurbits. Spots are small, circular, black with grayish centers, occurring
rst on leaves in center of watermelon hills. On cucumber, muskmelon, and
squash the spots are large and ochre-gray. Defoliation of vines causes reduc-
tion in fruit size, but the disease is not considered important. Clean up dis-
eased vines; use a 2- or 3-year rotation; spray or dust as for bacterial wilt.
Cercospora concors (see Mycovellosiella concors). Potato Leaf Spot,
Leaf Blotch.
Cercospora cornicola. Dogwood Leaf Spot, in the Gulf states, often with
Septoria orida. Spots irregular without denite borders.
Cercospora fusca (see Sirosporium diffusum). Pecan Brown Leaf Spot,
prevalent throughout the pecan belt but minor, serious only with high rainfall
and in neglected orchards where trees lack vigor.
Cercospora lathyrina. Leaf Spot on pea and sweet pea, in southern states
and north to New Jersey and Missouri. Angular to elongate spots have dirty
gray centers with a black line border.
Cercospora lythracearum. Leaf Spot on crape-myrtle, in Texas. Spots cir-
cular, pale brown to gray with a greenish fringe or yellow halo.
Cercospora magnoliae (see Cercosporidium magnoliae). (Mycosphaerella
milleri, Telleomorph). On magnolia in South.
Cercospora melongenae. Eggplant Leaf Spot, more common in tropical
areas. Yellow lesions change to large brown areas with concentric rings.
Cercospora nandinae. Nandina Leaf Spot, one of the few diseases of
this usually healthy shrub. Red blotches appear on upper leaf surface with
centers of older spots almost black. There is a scant sooty fruiting layer on
the undersurface. Reported from Alabama and North Carolina.
Cercospora personata (Mycosphaerella berkeleyii, Teleomorph) (see Phae-
oisariopsis personata). Peanut Leaf Spot, general on peanut.
Cercospora piaropi. Leaf Spot on water-hyacinths.
Cercospora pittospori. Pittosporum Leaf Spot, reported from Mississip-
pi, Florida, Louisiana, and Texas. Spots small, angular, yellow to dull brown,
fruiting in fawn-colored effuse patches on lower surface.
Cercospora puderi (see Pseudocercospora puderi). Leaf Spot on rose,
reported from Georgia and Texas.
Cercospora resedae. Leaf Spot, Blight of mignonette, a rapid disease
killing much of the foliage. Numerous small circular spots, pale yellow with
LEAF SPOTS 313
reddish brown borders, run together, discoloring the entire leaf. Spores are
spread by wind and rain; lower leaves are most affected.
Cercospora rhododendri (see Pseudocercospora handelii). Rhododen-
dron Leaf Spot.
Cercospora richardiaecola. Leaf Spot on calla lily, sometimes injurious.
Spots circular, brown, tan, or gray. Avoid syringing; keep plants well spaced;
ventilate greenhouse.
Cercospora rosicola (Mycosphaerella rosicola, Teleomorph). Cercospora
Spot of rose, wherever roses are grown but more important in the South.
Spots are circular, 1 to 4 mm, but coalescing to irregular areas, purplish or
reddish brown with pale brown, tan, or gray centers. Perithecia are formed
in fallen leaves.
Cercospora smilacis. Smilax Leaf Spot. Spots are more or less circular up
to 1/4 inch, dark purplish red, centers fading with age but margins remaining
denite and dark.
Cercospora sojina. Frog-Eye Disease of soybean. Typical frog-eye spots
are formed on leaves and elongated reddish lesions on stems, changing to
brown, gray, or nearly black with age. Pods of late varieties may be infected.
The fungus winters on diseased leaves and stems. Seed treatment is not effec-
tive; crop rotation is necessary. Early varieties often escape injury. There is
a wide difference in varietal susceptibility.
Cercospora symplocarpi. Leaf Spot on snowberry, coralberry, and wolf-
berry. Very small circular to angular spots, uniformly brown or with tan cen-
ters and brown margins.
Cercospora sp. Leaf Spot on kalancho.
Cercospora zebrina. Leaf and Stem Spot on bean, cowpea, groundnut,
peanut, birdsfoot trefoil and Lespedeza. Leaf Spot on clovers
Cercosporidium bougainvilleae (formerly Cercospora bougainvilleae).
Leaf Spot rst seen in Florida in 1962 and now the most important pathogen
of this host. Lesions are 1 to 5 mm, circular, depressed, with brown or tan
centers, reddish brown margins and a diffuse chlorotic area.
Cercosporidium magnoliae (formerly Cercospora magnoliae). (Myco-
sphaerella milleri, Telleomorph). On magnolia in South. Leaf spots are
small, angular, dark, with narrow yellow halo.
Mycovellosiella concors (formerly Cercospora concors). Potato Leaf
Spot, Leaf Blotch. Spots none to large irregular brown areas. Fruiting on
undersurface; conidiophores very pale; conidia almost hyaline.
Phaeoisariopsis personata (formerly Cercospora personata (Mycosphaerel-
la berkeleyii, Teleomorph)). Peanut Leaf Spot, general on peanut. Spots
are circular, 1 to 7 mm, but may coalesce; dark brown to black, often with
a yellow halo. Conidiophores on both sides of the leaf, more numerous on
the lower, are arranged concentrically in tufts; the epidermis is ruptured.
Spores are pale brown to olivaceous, one- to eight-septate. In wet seasons
vines may be nearly defoliated. Primary infections come from ascospores on
overwintered peanut leaves. Sulfur dust with 3.5% copper is recommended;
apply every 10 to 14 days.
Pseudocercospora abelmoschi (formerly Cercospora abelmoschi). Leaf
Spot on okra, hibiscus. Spots indistinct, but a sooty fruiting of spores on
under leaf surface.
Pseudocercospora cannabina (formerly Cercospora cannabina). Leaf
Curl and Wilt on hemp.
Pseudocercospora capsellae (formerly Cercospora albo-maculans (Syn.
Cercosporella brassicae)). White Spot of turnip, Chinese cabbage, mus-
tard, and other crucifers, common in the Southeast. Small, pale, circular
slightly sunken spots; may coalesce.
Pseudocercospora handelii (formerly Cercospora rhododendri). Rhodo-
dendron Leaf Spot. Angular dark brown spots with grayish down in cen-
ter. Control seldom necessary.
Pseudocercospora puderi (formerly Cercospora puderi). Leaf Spot on
rose, reported from Georgia and Texas. Spots are circular, to 5 mm, with
dingy gray centers, brown or reddish brown margins. Fruiting is chiey on
the upper surface in dense fascicles of short conidia.
Sirosporium diffusum (formerly Cercospora fusca). Pecan Brown Leaf
Spot, prevalent throughout the pecan belt but minor, serious only with high
rainfall and in neglected orchards where trees lack vigor. Spots are circu-
lar to irregular, reddish brown, often with grayish concentric zones. The
fungus winters in old spots on leaves. In Florida the disease appears rst
in June or July on mature leaves and may cause premature defoliation in
October. Stuart variety is particularly susceptible; others are more resistant.
Control with one application of bordeaux mixture between May 15 and
June 15.
LEAF SPOTS 315
Figure 3.23 Shot-Hole on Prunus sp
Cercosporella
Conidiophores hyaline, bearing conidia apically or on short branches; conidia hyaline,
cylindrical to liform with several cells (see Figs. 3.23 and 3.24); like Cercospora except
for light conidiophores; parasitic.
Cercosporella brassicae (see Pseudocercosporella capsellae). Leaf Spot
of cabbage, turnip, mustard, on West Coast.
Pseudocercosporella capsellae (formerly Cercosporella brassicae). Leaf
Spot of cabbage, turnip, mustard, on West Coast. Lesions on cabbage are
black, those on turnip and mustard gray with tan margins.
Cercosporidum
Cercosporidium personata (see Phaeoisariopsis personata). Leaf Spot on
peanut.
Phaeoisariopsis personata (formerly Cercosporidium personata). Leaf
Spot on peanut.
Ciborinia
Blights.
Ciborinia whetzelii (Syn. Sclerotinia whetzelii). Black Leaf Spot of
poplar, Ink Spot, from New England States to the Rocky Mountains on
aspen, black poplar, and other species. Saucerlike, thin black sclerotia are
formed in leaves, fall to the ground, and produce apothecia in spring. There
is often considerable defoliation, and small trees may be killed.
Ciborinia seaveri (Sclerotinia bifrons). Ink Spot, in western states, produc-
ing apothecia on ground under cottonwoods and poplars but pathogenic state
confused.
Cladosporium
Blotch Diseases.
Cladosporium colocasiae. Leaf Spot on elephants ear.
Cladosporium echinulatum. Leaf Spot on carnation.
Cladosporium epiphyllum. Leaf Spot on locust.
Cladosporium oxysporum. Leaf Spot on tomato
Blumeriella (Coccomyces)
Ascomycetes, Discomycetes
Blumeriella jaapii (formerly Coccomyces hiemalis and Higginisia hie-
malis). Cherry Leaf Spot, Blight, Shoot Hole, general on sweet and sour
cherries, the most common and destructive leaf disease of cherries. Leaf
spots are circular, rst purplish, then brown, falling out to give the shot-hole
effect (see Fig. 3.24). If lesions are numerous, the leaves turn yellow and fall
by midsummer, this premature defoliation reducing next seasons harvest.
The fungus winters in fallen leaves, producing disc-shaped apothecia for pri-
mary infection. Secondary infection comes from conidia, formed in whitish
masses on the spots in moist weather, more numerous on the undersurface.
New infection continues through the summer after harvest. Defoliation prior
to ripening reduces size and quality of fruit and exposes it to sunscald. Some
seasons shoots, spurs, and branches are killed, followed by a light crop the
next year. Thousands of sour cherry trees have been killed.
Control. An eradicant spray of a dinitro compound, such as Elgetol, applied
to the ground in early spring, reduces the amount of primary inoculum, but
summer sprays are also necessary. On sour cherry this may mean a spray at
petal fall, another 10 days later, two sprays in June, and another just after
LEAF SPOTS 317
Figure 3.24 Some Leaf-Spot Fungi. Ascochyta, hyaline, two-celled conidia in pycnidium; Cercosporella, hyaline,
septate spores on condiophores emerging from a stoma; Cladosporium (formerly Heterosporium), spiny, dark,
septate spores; Helmonthosporium, smooth, dark, septate spores; Mycosphaerella, two-celled hyaline ascospores
in a perithecium; Phyllosticta, hyaline, one-celled conidia in pycnidia formed in spots on leaves; Ramularia, hyaline
spores, becoming septate, formed successively on conidiophores; Stemphylium, colored muriform spores borne
free on mycelium
fruit is picked, with more applications, especially on nursery trees, needed
in some seasons. Consult your state experiment station for suitable materials
and schedule for your area.
Blumeriella jaapii (formerly Coccomyces lutescens). Leaf Spot, Shot
Hole on cherry-laurel, black cherry, and chokecherry. Similar to the disease
caused by C. hiemalis.
Blumeriella jaapii (formerly Coccomyces prunophorae). Leaf Spot, Shot
Hole on garden plum and apricot. Reddish to brown spots, dark blue initial-
ly, produce pinkish spore masses on underside of leaves in wet weather. The
shot-hole effect from dropping out of dead tissue may be very prominent and
accompanied by heavy fruit drop. Spray when shucks are off young fruit, 2
or 3 weeks later, and before fruit ripens, with lime sulfur, or with wettable
sulfur.
Blumeriella kerriae (formerly Coccomyces kerriae and Higginisia kerriae).
Kerria Leaf Spot, Twig Blight, widespread on kerris from eastern states
to Texas. Leaves have small, round to angular, light brown or reddish brown
spots with darker borders. When spots are numerous, leaves turn yellow and
die. Similar lesions on young stems may run together into extended cankers,
the bark splitting to show black pycnidia, from which ooze out masses of
long, white, curved spores. The fungus winters in old dead leaves. Spraying
with bordeaux mixture may help.
Coccomyces hiemalis and Higginisia hiemalis (see Blumeriella jaapii).
Cherry Leaf Spot, Blight, Shoot Hole, general on sweet and sour cher-
ries, the most common and destructive leaf disease of cherries.
Coccomyces kerriae and Higginisia kerriae (see Blumeriella kerriae) Ker-
ria Leaf Spot, Twig Blight, widespread on kerria from eastern states to
Texas.
Coccomyces lutescens (see Blumeriella jaapii). Leaf Spot, Shot Hole on
cherry-laurel, black cherry, and chokecherry.
Coccomyces prunophorae (see Blumeriella jaapii). Leaf Spot, Shot Hole
on garden plum and apricot.
Colletotrichum
Anthracnose.
Colletotrichum acutatum. Fruit Spot, Crown and Petiole Spot on
strawberry.
Colletotrichum coccodes. Leaf Spot and Slight Blight of velvetleaf.
Colletotrichum dematium f. sp. truncata. Leaf Spot and Stem Canker
of Stylosanthes spp.
Colletotrichum elastica (see Colletotrichum gloeosporioides). Leaf Spot
on g (Fiscus carica). Leaf Spot of basil, owering dogwood, cyclamen,
jasmine, passion ower, leaf and stem spot of calendula and dwarf mistletoe;
on many other hosts as anthracnose.
Colletotrichum gloeosporioides (formerly Colletotrichum elastica). Leaf
Spot on g (Fiscus carica). Leaf Spot of basil, owering dogwood, cycla-
men, jasmine, passion ower, leaf and stem spot of calendula and dwarf
mistletoe; on many other hosts as anthracnose.
Coniothyrium
Cankers.
LEAF SPOTS 319
Coniothyrium concentricum (see Microsphaeropsis concentrica). Leaf
Spot of century plant and yucca.
Coniothyrium hellebori. Black Spot of Christmas rose. Large, irregular,
dark brown to black spots on both sides of leaves, often running together with
concentric zonation; many leaves turn yellow prematurely and die; plants are
weakened and fail to mature the normal number of leaves. Stems may be
cankered, shrivel, and fall over, with wilting of unopened ower buds. Open
petals sometimes have black spots. In wet weather in spring and fall the
disease can spread through an entire planting in 2 or 3 days, but continuous
moisture is necessary for infection. Spray with bordeaux mixture.
Coniothyrium pyrina. Leaf Spot, Fruit Spot of apple, pear.
Microsphaeropsis concentrica (formerly Coniothyrium concentricum).
Leaf Spot of century plant and yucca. Spots are zoned, light grayish brown,
an inch or more in diameter, with concentric rings of tiny black pycni-
dia. Large portions of leaves may be destroyed. Remove and burn diseased
leaves.
Corynespora
Hyphae and conidia both dark.
Corynespora cassiicola (Syn. Helminthosporium vignicola). Soy Bean
Target Spot, also on cowpea, tomato, poinsettia, vinca, and privet; general
in South. Circular to irregular, reddish brown leaf spots, pin point to 1/4 inch,
often zonate and surrounded by yellow-green halos. Fruit necrotic pitting
and freckles are also found on infected fruit. Dark brown spots on petioles,
pods, and seed. Variety Ogden is moderately resistant. The same fungus
causes reddish purple spots on azalea, hydrangea and leaf spots on lipstick
vine, and on weeping g and leaf spot on thyme.
Cristulariella
Deuteromycetes, Moniliales, Moniliaceae
Sterile hyphae decumbent; fertile hyphae hyaline; ascending in a branched head with
conidia at tips of intermediate branches; spores globose, hyaline, one-celled.
Cristulariella depraedans. Leaf Spot on sugar and other maples. Spots
gray, denite or conuent.
Cristulariella moricola. Zonate Leaf Spot on Halesia. Leaf Spot on
hibiscus and tomato.
Cristulariella moricola (Telemorph, Grovesinia pyramidalis). Leaf Spot
on maple, tree-of-heaven, apple, bean, blueberry, cherry, dogwood, hibiscus,
sycamore, tung tree, viburnum, walnut, black walnut, beggar-ticks, trum-
pet vine, Mexican tea, dayower, blue waxweed, tick clover, mistower,
white snakeroot, morning glory, Indian tobacco, blue cardinal-ower, beef-
steak plant, poke, smart weed, false buckwheat, yellow dock, prickly mallow,
goldenrod, catbird grape, nectarine, grape, maple, serviceberry and boxelder.
Spots yellow-gray with denite margins.
Cryptomycina
Apothecium splitting irregularly into lobes, hyphal layer thin; spores hyaline, one-celled.
Cryptomycina pteridis. Tar Spot of fern, bracken. Spots are usually on
lower surface and between veins; leaves may roll.
Cryptostictis
Blights.
Cryptostictis arbuti (see Seimatosporium arbuti). Leaf Spot on Arbutus
menziesii, Manzanita, ledum.
Seimatosporium arbuti (formerly Cryptostictis arbuti). Leaf Spot on
Arbutus menziesii, Manzanita, ledum.
Cycloconium
Mycelium coiled, spores small, dark, two-celled; scarcely different from short hyphae.
Cycloconium oleaginum. Olive Leaf Spot, Peacock Spot, Ring Spot.
Blackish, more or less concentric rings on leaves, especially those weakened
or old.
LEAF SPOTS 321
Cylindrocladium
Conidiophores repeatedly dichotomously or trichotomously branched, each terminating
in two or three phialides (cells developing spores); conidia hyaline, with two or more
cells, cylindrical, borne singly; parasitic or saprophytic.
Cylindrocladium avesiculatum. Leaf Spot and Twig Dieback on holly,
and Leucotho sp.
Cylindrocladium colhounii. Leaf Spot on bottle-brush (Callistemon).
Cylindrocladium clavatum. Leaf Spot on bottle-brush (Callistemon).
Cylindrocladium pteridis. Leaf Spot, Leaf Blight of Washington palm.
Numerous small dark brown spots with light margins are somewhat disgur-
ing.
Cylindrocladium pteridis. Fern Leaf Spot, Leaf Blotch. Reddish brown
lesions run together to cover large areas. Pick off and burn infected fronds.
Cylindrosporium
Acervuli subepidermal, white or pale; conidiophores short, simple; conidia hyaline, li-
form, straight or curved, one-celled or septate; parasitic on leaves.
Cylindrosporium betulae. Brown Leaf Spot of Birch. Sometimes serious
enough to defoliate but not often present on ornamental trees.
Cylindrosporium chrysanthemi. Chrysanthemum Leaf Spot. Spots are
dark brown with yellowish margins, increasing to take in the whole leaf,
which hangs down. Similar to more common Septoria leaf spot.
Cylindrosporium clematidinis. Clematis Leaf Spot. Reddish brown
spots on lower leaves, which may drop. Dusting with sulfur has been sug-
gested.
Cylindrosporium salicinum. Willow Leaf Spot. Sometimes causing defo-
liation; can be controlled with bordeaux mixture if necessary.
Cylindrosporium sp. Leaf Spot on spirea, recorded from a Kansas nurs-
ery. Light yellow lesions turn dark brown, with masses of yellow conidia on
underside.
Cytospora
Cankers.
Cytospora sp. Leaf Spot on mulberry.
Dactylaria
Dactylaria higginsii. Leaf Spot on nutsedge.
Dichotomophthoropsis
Dichotomophthoropsis nymphaearum. Leaf Spot on water-lily, and
water shield.
Didymaria
Conidiophores simple, arising from leaf surface in loose groups; conidia hyaline, two-
celled, ovate-oblong, borne singly; parasitic on leaves.
Didymaria didyma (see Ramularia didyma). Leaf Spot on anemone.
Angular brown spots.
Ramularia didyma (formerly Didymaria didyma). Leaf Spot on anemone.
Angular brown spots.
Didymellina
Acomycetes, Sphaeriales, Mycosphaerellaceae
Perithecia separate, innate or nally erumpent, not beaked; spores two-celled, hyaline.
Didymellina macrospora (Heterosporium iridis, H. gracilis) (see Myco-
sphaerella macrospora, Anamorph). Irish Leaf Spot, Blotch, Fire on both
bulbous and rhizomatous iris.
Didymellina ornithogali (Heterosporium ornithogali) (see Mycosphaerella
ornithogali). Leaf Spot on star-of-bethlehem.
LEAF SPOTS 323
Didymellina poecilospora. A weak parasite sometimes causing black dis-
coloration of iris foliage.
Mycosphaerella macrospora (formerly Didymellina macrospora; Het-
erosporium iridis, H. gracilis, Anamorph). Irish Leaf Spot, Blotch, Fire
on both bulbous and rhizomatous iris. The spotting is conspicuous toward
the end of the season but is not too serious in a normally dry season. Usually
the spots are conned to the upper half of leaves, but if plants are crowded
and shaded and the summer is wet, the spotting appears earlier, covers more
of the leaf, and is more damaging.
Spots are dark brown at rst, surrounded by a water-soaked and then yel-
lowing region; they enlarge into rather oval lesions, up to 1/2 inch long, with
a red-brown border (Fig. 3.25). Flower buds and stems of bulbous iris may
be attacked. Tufts of olive conidia turn the centers grayish, the spores being
produced in abundance and splashed by rain to neighboring leaves. Infection
is through stomata or directly through the epidermis. The fungus winters as
mycelium in old leaves, and in spring produces a fresh crop of conidia or
perithecia of the Didymellina stage. Soils decient in lime apparently favor
Figure 3.25 Iris Leaf Spot
the disease. Repeated spotting reduces bloom and, after a number of years,
may kill plants.
Control. It is often sufcient to remove and burn all old leaves at the end
of the season; shearing back spotted leaves in midsummer is helpful. If the
disease is regularly a problem, spray with bordeaux mixture, starting when
fans are 6 to 8 inches high and repeating at 10- to 14-day intervals.
Mycosphaerella ornithogali (formerly Didymellina ornithogali; Heterospo-
rium ornithogali, Anamorph). Leaf Spot on star-of-bethlehem. Occasional
sooty spots on leaves, with foliage blackened and killed in severe infections.
Didymosporium
Conidia are slime-spores in acervuli; dark, two-celled.
Didymosporium arbuticola. Leaf Spot on Arbutus menziesii.
Dilophospora
Pycnidia distinct in a stroma; conidia very long, liform, with bristlelike hairs at each
end. Usually found on cereals and sometimes with the wheat nematode, causing a disease
called twist.
Dilophospora geranii (see Pestalozziella subsessilis). Leaf Spot on native
geranium.
Pestalozziella subsessilis (formerly Dilophospora geranii). Leaf Spot on
native geranium.
Diplodina
Pycnidia black, separate, immersed or erumpent, globose or attened, ostiolate; coni-
diophores simple, slender; conidia hyaline, two-celled, ovoid or ellipsoid; parasitic or
saprophytic. Similar to Ascochyta but not produced in spots.
Diplodia rhododendri (see Encoeliopsis rhododendron). Leaf Spot on
rhododendron.
LEAF SPOTS 325
Encoeliopsis rhododendron (formerly Diplodia rhododendri). Leaf Spot
on rhododendron.
Diplotheca (Stevensea)
Ascomycetes, Myriangiales
Asci born singly in locules at various levels in a massive stroma; spores dark, several-
celled.
Diplotheca wrightii. Black Spot, Charcoal Spot of Opuntia cacti in
Florida and Texas uncommon in the North. Dark spots, 1/4 inch or more in
diameter, are surrounded by a ring of fruiting bodies.
Dothichiza
Cankers.
Dothichiza caroliniana. Leaf Spot, Double Spot of blueberry, found only
on Vaccinium australis in North Carolina, but there causing extensive defo-
liation. Leaf spots are small, circular, with brown centers and a dark brown
ring, but in late summer infection spreads to a secondary necrotic area around
the original spot, giving the common name of double spot. Black pycnidia
are formed sparsely in the spots. All varieties of high bush blueberries are
somewhat susceptible, but Cabot, Dixie, Pioneer, and Rancocas are most
damaged.
Ectostroma
Black stromata formed in leaves and stems.
Ectostroma liriodendri. Tar Spot, widespread in tulip-trees but perhaps
secondary after insect injury.
Epicoccum
Sporodochia dark, rather cushion-shaped; conidiophores compact or loose, rather short;
conidia dark, with one or more cells, globose; mostly saprophytic.
Epicoccumasterinum(see Epicoccum nigrum). Leaf Spot of yucca; E. ne-
glectum, on royal palm; E. nigrum, on Magnolia grandiora; E. purpuras-
cens, on amaryllis.
Epicoccum nigrum (formerly Epicoccum asterinum). Leaf Spot of yucca;
E. neglectum, on royal palm; E. nigrum, on Magnolia grandiora; E. pur-
purascens, on amaryllis. All of these may be secondary infections. E. neglec-
tum and E. purpurascens are also synonyms of E. nigrum.
Exosporium
Conidia on subglobose to convex sporodochia; spores dark, with two to several cells,
somewhat club-shaped.
Discogloeum concentricum (formerly Exosporium concentricum). Leaf
Spot on euonymus and ligustrum (privet) in the South.
Exosporium concentricum (see Discogloeum concentricum). Leaf Spot
on euonymus and ligustrum (privet) in the South.
Fusicladium
Mycelium forming a stroma under cuticle of host; conidiophores dark, short; conidia
dark, two-celled, produced successively as pushed-out ends of new growing tips. Para-
sitic on higher plants, causing scab as well as leaf spots.
Fusicladium pisicola. Black Leaf of peas, rst reported in Utah in 1921,
causing trouble with canning peas. Spots start as small, irregular whitish
areas on undersurface of leaets and stipules, but they darken to gray or
black from the closely packed layer of dark conidia. The disease is not very
important.
Fusicladium robiniae (see Phaeoisariopsis robiniae). Leaf Spot, Seed-
ling Leaf Blight of black locust.
Phaeoisariopsis robiniae (formerly Fusicladium robiniae). Leaf Spot,
Seedling Leaf Blight of black locust. Spots are small, with light centers
and dark margins. There is frequently defoliation of seedlings, sometimes
stunting and death.
LEAF SPOTS 327
Gibbago
Gibbago trianthemae. Leaf Spot of horse purslane; a new genus and
species, recently described (1986), with potential for bioherbicide activity.
Gloeocercospora
Sporodochia formed on surface of host above stomata from hyphae emerging through
openings; conidiophores hyaline, simple or branched; conidia hyaline, elongate to li-
form, one- to many-septate, straight or curved, in a slimy matrix.
Gloeocercospora inconspicua. Leaf Spot of highbush and rabbit-eye blue-
berry. Circular to angular brownish spots on leaves, with sporodochia more
frequent on upper surface. These are at discs when dry, glistening globules
when wet, containing curved, septate conidia.
Gloeocercospora sorghi. Copper Spot of turf. (
Ramulispora sorghi).
Gloeosporium
Anthracnose.
Asteroma inconspicuum (formerly Gloeosporium inconspicuum). Elm
Leaf Spot, Twig Blight, Anthracnose on American and English elms.
Subcircular brown spots with darker margins and centers are visible on upper
and lower leaf surfaces.
Cryptocline betularum (formerly Gloeosporium betularum). Leaf Spot,
Anthracnose of river birch. Spots are more or less circular, 1/8 inch across,
brownish with pale centers and yellow margins.
Gloeosporium betularum (see Cryptocline betularum). Leaf Spot, An-
thracnose of river birch.
Gloeosporium inconspicuum (see Asteroma inconspicuum). Elm Leaf
Spot, Twig Blight, Anthracnose on American and English elms.
Gloeosporium mezerei (see Marssonina daphnes). Leaf Spot on daphne.
Gloeosporium rhododendri. Leaf Spot on rhododendron, tulip-tree.
Gloeosporium ulmicola. Elm Leaf Spot. Elongated spots on midribs,
veins, and margins, visible on both leaf surfaces.
Marssonina daphnes (formerly Gloeosporium mezerei). Leaf Spot on
daphne. Small brown spots on both sides of leaves.
Glomerella
Anthracnose.
Glomerella cingulata. Leaf Spot, widespread on queen palm, dracaena,
and maranta. Sobralia blight of orchids. Dark discoloration starts at tip of
leaves and advances toward base.
Glomerella cingulata. Leaf Spot on apple, aucuba, wampi, and croton. See
under Anthracnose for this fungus on many other hosts.
Glomerella sp. Black Spot of Vanda orchids.
Gnomonia
Anthracnose.
Gnomonia comari. Leaf Spot/Blotch and Fruit Rot of strawberry.
Gnomonia fragariae. Leaf Spot, Leaf Blotch of strawberry. Often asso-
ciated with Dendrophoma causing leaf blight, but not connected.
Gnomonia nerviseda (formerly Gnomonia caryae var. pecanae). Pecan
Liver Spot. Dark brown circular spots, mostly along midribs on underside
of leaves, appear in May and June. In autumn the color changes to cinnamon
brown, and dark fruiting bodies appear; there may be premature defoliation.
Spray in May with bordeaux mixture.
Pecan Vein Spot. Lesions resemble pecan scab on veins or leaf stems;
sometimes a narrow brown lesion extends nearly the length of a midrib.
Defoliation may be moderate or severe. Stuart variety is especially suscepti-
ble. Spray with bordeaux mixture just before and just after pollination; repeat
3 to 4 weeks later.
Gnomonia ulmea, Anamorph, Gloeosporium ulmeum (see Stegophora
ulmea). Elm Black Spot, Black Leaf Spot of Elm, general on Amer-
ican, English, and Chinese elms.
Gnomonia caryae var. pecanae (see Gnomonia nerviseda). Pecan Liver
Spot.
LEAF SPOTS 329
Stegophora ulmea (formerly Gnomonia ulmea, Anamorph, Gloeosporium
ulmeum). Elm Black Spot, Black Leaf Spot of Elm, general on Ameri-
can, English, and Chinese elms. Spots on leaves are small but conspicuous,
shining coal black, and slightly raised. Leaves may turn yellow and drop,
with severe defoliation in a wet season, especially on Siberian elm. Defoli-
ation in spring means death of twigs, but the disease is more common and
less important toward fall. Ascospores are formed in spring in perithecia on
fallen dead leaves; conidia are produced as a creamy exudate of spores in
summer. The fungus also winters as mycelium in dormant buds.
Control. Rake and burn fallen leaves. Chemical control is required only in
a wet spring, difcult to determine in advance.
Gnomoniella
Perithecia in substratum, beaked, membranous, separate; spores hyaline, one-celled.
Gnomoniella coryli (see Mamianiella coryli). Leaf Spot on hazel, frequent
in northern states.
Gnomoniella mbriata (see Mamianiella mbriata). Leaf Spot of horn-
beam.
Mamianiella coryli (formerly Gnomoniella coryli). Leaf Spot on hazel,
frequent in northern states. Controlled with bordeaux mixture aided by clean-
ing up fallen leaves.
Mamianiella mbriata (formerly Gnomoniella mbriata). Leaf Spot of
hornbeam.
Gonatobotryum
Conidiophores dark, with spiny inations at intervals, around which are borne ovoid,
dark, one celled conidia.
Gonatobotryum apiculatum. Leaf Spot on witchhazel.
Graphium
Synnema or coremium tall, dark, with a rounded terminal mass of conidia
embedded in mucus; simple, hyaline conidiophores; oblong conidia repro-
ducing by budding; parasitic.
Graphium sorbi. Leaf Spot of mountain-ash.
Guignardia
Blotch Diseases.
Guignardia bidwellii f. sp. parthenocissi. Leaf Spot on Boston ivy, pep-
per-vine, and Virginia creeper. Spots are numerous, angular, reddish brown,
usually dark brown at margins, with black dots in center, minute pycnidia of
the anamorph Phyllosticta state. Leaves are quite unsightly and there may be
defoliation. Bordeaux mixture applied two or three times, starting as leaves
are expanding, gives some control, but the cure looks about as bad as the
disease. This fungus is a form of the species causing black rot of grapes.
Helminthosporium
Blights.
Bipolaris cynodontis (formerly Helminthosporium cynodontis). Bermuda
Grass Leaf Blotch, general in South. Olive brown indenite lesions on dry
leaves.
Bipolaris setariae (formerly Helminthosporium setariae (Drechslera setari-
ae)). Leaf and Petal or Greasy Spot on geranium, areca palm, shtail
palm, rhapis palm, Calathea spp. Maranta spp., and Chamaedorea spp.
Bipolaris sorokiniana (formerly Helminthosporium sativum). Melting-
out, prevalent on bent grass in warm weather. Leaf Spot on Russian
wildrye (Elymus); Spot Blotch on switchgrass (Panicum).
Bipolaris sorokiniana (formerly Helminthosporium sorokiniana). Leaf
Spot and Stem Spot of wild rice. Leaf and Pod Spot on bean.
Drechslera catenaria (formerly Helminthosporium catenarium). Leaf
Spot on ribbon-grass.
Drechslera dictyoides (formerly Helminthosporium dictyoides). Fescue
Netblotch, general on fescue. Dark streaks across green leaves with darker
lengthwise streaks form a net pattern. Leaves turn yellow and die back from
tips.
LEAF SPOTS 331
Drechslera erythrospilum (formerly Helminthosporium erythrospilum).
Red Leaf Spot on redtop and bent grasses, widespread in eastern and
midwestern states. Under wet conditions lesions have small, pale centers
with russet borders; in dry weather leaves wither as in drought but with less
evident spotting. Conidia are typically cylindrical, rounded at both ends,
yellowish, and germinate from any or all cells (see Fig. 3.23).
Drechslera giganteum (formerly Helminthosporium giganteum). Zonate
Leaf Spot, Eye Spot on bent grasses, Canada and Kentucky bluegrass, and
Bermuda grass. The disease is present in turf and in nursery rows. Spots are
small, 1/16 to 1/8 inch, bleached-straw color in centers. In presence of mois-
ture (dew is sufcient) the fungus grows periodically into new areas, giving
the zoned appearance. In continued wet weather leaves are killed and grass
turns brown. Metropolitan and velvet bent grasses are less susceptible. Most
injury is in July and August. The fungus overwinters as dormant mycelium
in old leaves.
Drechslera poae (formerly Helminthosporium vagans). Bluegrass Leaf
Spot, Going-Out, Melting-out, Foot Rot, general but most injurious in
northeastern states, on bluegrass only. Scattered circular to enlongate leaf
spots, 0.5 to 3 by 1 to 8 mm, have prominent reddish brown to black borders;
centers are brown changing to straw-colored or white with age. The disease,
favored by cool rainy weather, usually appears in early spring, sometimes in
late fall, and is most severe on close-clippped turf. Grass thins out in large
areas; roots rot; weeds invade exposed soil.
Control. Merion bluegrass is quite resistant to leaf spot and will stand close-
clipping. For other bluegrasses cut high and fertilize well to help turf with-
stand the disease.
Drechslera siccans (formerly Helminthosporium siccans; Teleomorph,
Pyrenophora lolii). Brown Blight on fescue, and ryegrass. Leaves die
back with numerous dark chocolate-brown spots, oval to elongate and often
coalescing. The disease appears in early spring in cool, moist weather.
Drechslera stenacra (formerly Helminthosporium stenacrum). Leaf Mold
on redtop and bent grasses. Indenite spots; leaves dry, withered, in fall.
Drechslera triseptata (formerly Helminthosporium triseptatum). Leaf
Spot, Gray Leaf Mold on redtop, spike and bentgrasses in Oregon, Wash-
ington, and New York. Leaf tips are killed with vague lesions; gray mold
appears on dying tissue.
Drechslera tritici-repentis (formerly Helminthosporium tritici-repentis).
Leaf Spot on Russian wildrye (Elymus).
Exserohilum rostratum (formerly Helminthosporium rostratum). Leaf
Spot on bromelia, areca palm, shtail palm, rhapis palm, sweet sorghum,
and Chamaedorea spp.
Helminthosporium catenarium (see Drechslera catenaria). Leaf Spot on
ribbon-grass.
Helminthosporium cynodontis (see Bipolaris cynodontis). Bermuda
Grass Leaf Blotch, general in South.
Helminthosporium dictyoides (see Drechslera dictyoides). Fescue Net-
blotch, general on fescue.
Helminthosporium erythrospilum (see Drechslera erythrospilum). Red
Leaf Spot on redtop and bent grasses, widespread in eastern and midwest-
ern states.
Helminthosporium giganteum (see Drechslera giganteum). Zonate Leaf
Spot, Eye Spot on bent grasses, Canada and Kentucky bluegrass, and
Bermuda grass.
Helminthosporium rostratum (see Exserohilum rostratum). Leaf Spot
on bromelia, areca palm, shtail palm, rhapis palm, sweet sorghum, and
Chamaedorea spp.
Helminthosporium sativum (see Bipolaris sorokiniana). Melting-out,
prevalent on bent grass in warm weather. Leaf Spot on Russian wildrye
(Elymus); Spot Blotch on switchgrass (Panicum).
Helminthosporium setariae (Drechslera setariae) (see Bipolaris setariae).
Leaf and Petal or Greasy Spot on geranium, areca palm, shtail palm,
rhapis palm, Calathea spp. Maranta spp., and Chamaedorea spp.
Helminthosporium siccans; Teleomorph, Pyrenophora lolii (see Drech-
slera siccans). Brown Blight on fescue, and ryegrass.
Helminthosporium sorokiniana (see Bipolaris sorokiniana). Leaf Spot
and Stem Spot of wild rice. Leaf and Pod Spot on bean.
Helminthosporium stenacrum (see Drechslera stenacra). Leaf Mold on
redtop and bent grasses.
Helminthosporium triseptatum (see Drechslera triseptata). Leaf Spot,
Gray Leaf Mold on redtop, spike and bentgrasses in Oregon, Washington,
and New York.
Helminthosporium tritici-repentis (see Drechslera tritici-repentis). Leaf
Spot on Russian wildrye (Elymus).
Helminthosporium vagans (see Drechslera poae). Bluegrass Leaf Spot,
Going-Out, Melting-out, Foot Rot, general but most injurious in north-
eastern states, on bluegrass only.
LEAF SPOTS 333
Hendersonia
Pycnidia smooth, innate or nally erumpent, ostiolate; conidia dark, several-celled, elon-
gate to fusoid; saprophytic or parasitic.
Hendersonia concentrica. Leaf Spot on rhododendron.
Hendersonia crataegicola. Leaf Spot on hawthorn. Spots irregular, dark
brown.
Cladosporium (Heterosporium)
Conidiophores dark, simple; conidia dark, spiny, cylindrical, with three or more cells;
parasitic, causing leaf spots, or saprophytic.
Acroconidiella escholtziae (formerly Heterosporium escholtziae). Capsule
Spot, Leaf Spot, Stem Spot of California poppy. Lesions faint purplish
brown; seed capsules may shrivel. Treat seed with hot water, 125F, for 30
minutes.
Cladosporium allii (formerly Heterosporium allii). Leaf Spot on onion,
leek, shallot, chive, and garlic; rare in North America. Leaves have elliptical,
depressed, pale brown spots, and yellow and wither from tip downward.
Cladosporium echinulatum (formerly Heterosporium echinulatum). Fairy
Ring Spot, Leaf Mold on carnation, occasional in greenhouses. Bleached
spots on leaves have black spore groups in ring formation. Syringe as little
as possible and on bright days; control ventilation.
Cladosporium iridis (formerly Heterosporium gracile). Leaf Spot on
chlorogalum, daylily, same as H. iridis on iris (conidial state of Didymellina
macrospora). Leaf Spot on iris, blackberry, lily, freezia, and gladiolus.
Cladosporium variabile (formerly Heterosporium variabile). Leaf Spot,
pinhead rust of spinach, cabbage mold, sometimes severe in cold, wet
weather. Circular, chlorotic spots with brown or purple margins enlarge and
multiply until they cover most of the leaf, which turns yellow, withers, dies.
There is a greenish black mold on both leaf surfaces, made up of large olive
conidia, one- to six-celled, covered with warts. Keep plants growing vigor-
ously in wall-drained soil.
Heterosporium allii (see Cladosporium allii). Leaf Spot on onion, leek,
shallot, chive, and garlic; rare in North America.
Heterosporium echinulatum(see Cladosporium echinulatum). Fairy Ring
Spot, Leaf Mold on carnation, occasional in greenhouses.
Heterosporium escholtziae (see Acroconidiella escholtziae). Capsule
Spot, Leaf Spot, Stem Spot of California poppy.
Heterosporium gracile (see Cladosporium iridis). Leaf Spot on chloro-
galum, daylily, same as H. iridis on iris (conidial state of Didymellina
macrospora (Fig. 3.25)). Leaf Spot on iris, blackberry, lily, freezia, and
gladiolus.
Heterosporium variabile (see Cladosporium variabile). Leaf Spot, pin-
head rust of spinach, cabbage mold, sometimes severe in cold, wet weath-
er.
Illosporium
Sporodochia cushionlike, light-colored; conidiophores hyaline, branched with phialides
bearing conidia apically; spores hyaline, one-celled; parasitic or saprophytic, often sec-
ondary.
Illosporium malifoliorum. Leaf Spot of apple and crabapple.
Isariopsis
Dark, synnemata composed of loose conidiophores with spores at or near tips; conidia
dark or pale, with two or more cells, cylindrical to obclavate, often curved; parasitic.
Isariopsis griseola (see Phaeoisariopsis griseda). Angular Leaf Spot,
Pod Spot of beans, also sweet pea.
Phaeoisariopsis griseda (formerly Isariopsis griseola). Angular Leaf
Spot, Pod Spot of beans, also sweet pea. Small, angular brown spots are
so numerous they give a checkerboard appearance to leaves. The fungus
forms a gray moldy covering over dead areas on underside of leaves. Pod
spots are conspicuous when present, black with red or brown centers, varying
from a speck to the width of the pod. Small, dark synnemata scattered over
LEAF SPOTS 335
the surface bear large conidia, with two to four cells, at top of stalks. They
are probably wind-disseminated. Control measures are seldom practical.
Also, Leaf Spot on kidney bean.
Kabatia
Pycnidia with a radiate shield or scutellum, with an ostiole; spores two-celled, hyaline,
like a tooth at the apex.
Kabatia lonicerae. Leaf Spot on honeysuckle.
Lasiobotrys
Perithecia in a ring around a sclerotial stroma; spores dark, two-celled.
Lasiobotrys lonicerae. Leaf Spot on honeysuckle. Spot is well-marked
with small, dark, wartlike stromas.
Leptostromella
Pycnidia elongate, with a cleft; separate; spores liform, with rounded ends, hyaline,
continuous to septate on simple conidiophores.
Leptostromella elastica. Leaf Spot of rubber-plant. The symptoms appear
in spots and streaks, but infection spreads until the entire leaf is involved.
Black lines outline spots in which small black pycnidia produce long, color-
less spores. Remove and burn infected leaves.
Leptothyrella
Pycnidia with a radiate shield, separate; spores 2-celled, hyaline.
Leptothyrella liquidambaris (see Tubakia dryina). Leaf Spot red on
sweetgum.
Tubakia dryina (formerly Leptothyrella liquidambaris). Leaf Spot red on
sweetgum.
Leptothyrium
Pycnidium attened with a more or less radiate shield, opening with a ostiole; spores
one-celled, hyaline, on simple conidiophores.
Kabatia periclymeni (formerly Leptothyrium periclymeni). Leaf Spot on
honeysuckle, widespread.
Leptothyrium californicum. Leaf Spot on coast live oak.
Leptothyrium dryinum (see Tubakia dryina). Leaf Spot on white oak.
Leptothyrium periclymeni (see Kabatia periclymeni). Leaf Spot on hon-
eysuckle, widespread.
Tubakia dryina (formerly Leptothyrium dryinum). Leaf Spot on white oak.
Linospora
Perithecia innate, beak often lateral, with a shield; paraphyses lacking; spores spindle-
shaped to liform, hyaline.
Linospora gleditschiae. Leaf Spot, Tar Spot on honey locust in the South.
Numerous black fruiting bodies are formed on undersurface of leaves.
Lophodermium
Fruiting body a hysterothecium, midway between an elongated perithecium and a com-
pressed apothecium, hard, black, opening with a long narrow slit; paraphyses present;
hooked at tip; spores liform, septate or continuous. Most species cause needle casts.
Lophodermium schweinitzii. Rhododendron Leaf Spot. Large silvery
white spots with red, raised margins have very prominent oval, black fruiting
bodies on the upper surface. Lower side of spots is a light chocolate brown.
LEAF SPOTS 337
Infected portions may fall out, leaving irregular holes. The disease is more
common on native than on hybrid varieties.
Macrophoma
Cankers.
Macrophoma candollei. Leaf Spot of boxwood. Conspicuous black pyc-
nidia on dead leaves, usually straw-colored, sometimes brown or tan. The
fungus is a weak parasite coming in secondarily after winter injury or other
predisposing factors.
Marssonina
Anthracnose.
Cylindrosporium populinum (formerly Marssonina rhabdospora; Teleo-
morph, Pleuroceras populi). Leaf Spot of poplar. Brown spots on living
leaves; beaked pyriform perithecia formed in fallen leaves over winter.
Didymosporina aceris (formerly Marssonina truncatula). Leaf Spot and
Leaf Blight of Norway maple.
Diplosporonema delastrei (formerly Marssonina delastrei). Leaf Spot on
corncockle and campion.
Discella ochroleuca (formerly Marssonina ochroleuca). Leaf Spot on oak,
American chestnut. Spots are circular, yellow to brown with concentric
markings, small on chestnut, up to an inch on oak.
Marssonina daphnes. Daphne Leaf Spot. Small, thick brown spots on
both sides of leaf, which turns yellow, dies.
Marssonina delastrei (see Diplosporonema delastrei). Leaf Spot on corn-
cockle and campion.
Marssonina fraxini (see Piggotia fraxini). Ash Leaf Spot, sometimes seri-
ous in nursery stock, controlled by spraying with bordeaux mixture.
Marssonina juglandis. See Gnomonia leptostyla under Anthracnose.
Marssonina ochroleuca (see Discella ochroleuca). Leaf Spot on oak,
American chestnut.
Marssonina populi. Poplar Leaf Spot. Brown spots with darker margins.
There may be premature defoliation and killing of twigs.
Marssonina rhabdospora (Teleomorph, Pleuroceras populi) (see Cylin-
drosporium populinum). Leaf Spot of poplar.
Marssonina rosae. Anamorph state of the rose blackspot fungus, Diplocar-
pon rosae.
Marssonina tremulae. Leaf Spot on poplar.
Marssonina truncatula (see Didymosporina aceris). Leaf Spot and Leaf
Blight of Norway maple.
Piggotia fraxini (formerly Marssonina fraxini). Ash Leaf Spot, sometimes
serious in nursery stock, controlled by spraying with bordeaux mixture.
Mastigosporium
Conidiophores hyaline, very short, simple; conidia with four or more cells, with or with-
out apical appendages: broadly cylindrical with rounded or pointed ends; parasitic on
grasses.
Mastigosporium rubricosum. Leaf Fleck on redtop and bent grasses.
Spores with rounded ends, without appendages.
Melanconium
Acervuli subcutaneous or subcortical, conic or discoid, black; with setae; conidiophores
simple; conidia dark, one-celled, ovoid to ellipsoid; parasitic or saprophytic.
Melanconium pandani. Leaf Spot on pandanus.
Melasmia
Pycnidia in a broad, black, attened stroma that is supercial or nearly so, dimidiate;
conidiophores simple or branched; spores hyaline or subhyaline, one-celled, allantoid or
fusoid; parasitic on leaves; anamorph states of Rhytisma.
Melasmia falcata. Tar Spot of persimmon.
Melasmia menziesiae. Leaf Spot tar spot of azalea.
LEAF SPOTS 339
Micropeltis
Blights.
Micropeltis alabamensis. Black Leaf Spot on magnolia.
Microstroma
Basidiomycetes, Exobasidiales
Sporodochia small, white, breaking through epidermis; conidiophores hyaline, one-
celled, somewhat clavate, bearing conidia on short sterigmata; spores hyaline, one-
celled, small, oblong; parasitic. Conidia are now known to be basidiospores.
Microstroma juglandis. Leaf Spot, White Mold, Downy Spot, Witch-
es Broom of pecan, walnut, and hickory. Yellow blotching of upper side of
leaves and a glistening white coating on underside, due to pustules with enor-
mous numbers of spores, may be accompanied by premature defoliation. On
shagbark hickory the fungus also invades the stems, causing witches brooms
up to 3 feet across. Leaves formed on them in spring are yellow-green, with
white powder on underside. Leaets are small, curled, and soon drop. Prune
out witches brooms; spray with bordeaux mixture.
Microthyriella
Fruit Spots.
Microthyriella cuticulosa. Black Spot of holly. Dark spots on leaves of
American holly, Georgia.
Monochaetia
Cankers.
Monochaetia monochaeta. Leaf Spot on chestnut, white, red, and coast
live oaks, winged elm, hickories, especially destructive in the Southeast.
Spots are large, 1 to 2 inches in diameter, with pale green or yellow centers
with a red and brown border or concentric zones of gray, yellow, and brown.
Symptoms appear most often in late summer when loss of green tissue is not
so important.
Lembosina (Morenoella)
See Lembosia under Black Mildew.
Lembosina quercina (formerly Morenoella quercina). Leaf Spot, Black
Mildew of red and black oaks; twig blight of white oak, common in South-
east. Spots are purplish black, roughly circular, up to 1/3 inch across, on
upper surface and irregular brown areas on underside. Mycelium is super-
cial in early summer, but by late summer there are subcuticular hyphae and
a black shield formed over a at cushion of fertile cells. Asci are mature and
shield is ssured by spring.
Morenoella quercina (see Lembosina quercina). Leaf Spot, Black Mil-
dew of red and black oaks; twig blight of white oak, common in Southeast.
Mycosphaerella
Blights.
Mycosphaerella angulata. Angular Leaf Spot of muscadine grapes. Many
small, angular black spots, more conspicuous on lower surface of leaves,
which may turn yellow and die.
Mycosphaerella arachidis. Peanut Leaf Spot.

Cercospora arachidico-
la.
Mycosphaerella berkeleyi. Peanut Leaf Spot.
Cercospora personata.
Mycosphaerella (Anamorph, Pseudcercospora) bolleana. Leaf Spot of g,
and rubber-tree.
Mycosphaerella (Anamorph, Asteromella brassicae) brassicicola. Ring
Spot of crucifers, chiey cabbage and cauliower, sometimes brussels
sprouts, broccoli, and turnip. Dead spots in leaves, small to 1/2 inch, are
surrounded by a green zone that keeps its color even if the rest of the leaf
turns yellow. Small black pycnidia are deeply embedded in the dead tissue,
often in concentric rings. In moist weather conidia ooze from pycnidia in
pink tendrils. The fungus winters in old plant refuse, and ascospores are
forcibly ejected from perithecia in spring. The disease is conned to the
Pacic Coast and, as black blight, is serious on the seed crop in the Puget
Sound area. Sanitary measures and crop rotation keep it in check.
LEAF SPOTS 341
Mycosphaerella caroliniana. Leaf Spot, Purple Blotch on oxydendron
(sourwood). Reddish or purple spots on foliage in midsummer have dry,
brown centers. Pycnidia embedded in tissue break through lower surface,
spores being formed in great numbers.
Mycosphaerella caryigena. Pecan Downy Spot. Conidial stage has
been listed as a Pseudocercosporella caryigena. Leaf spots are pale yel-
low when young, turning yellow-brown, brown, or black. Conidia pro-
duced in minute acervuli on underside of leaves form a white downy or
frosty coating; leaves may drop early. Spores are spread in rain, fog, and
dew. The fungus overwinters in leaves, liberating ascospores in spring to
infect new foliage. Moneymaker and Stuart varieties are especially suscepti-
ble.
Control. Turn under old leaves before spring (plowing under winter cover in
spring takes care of this). Spray as for scab, bordeaux mixture when leaves
are half-grown and bordeaux plus 4 pounds of zinc sulfate when tips of small
nuts have turned brown.
Mycosphaerella cerasella.

Cercospora circumscissa.
Mycosphaerella (Anamorph, Cercospora) cercidicola. Redbud Leaf
Spot, general. Spots are circular to angular or irregular with raised dark
brown borders. With age, lesions become grayish above and rusty brown
on the undersurface, with the leaf tissue yellow-green outside the borders.
Spores are formed on fascicles of conidiophores projecting through stomata.
The fungus winters on fallen leaves, producing perithecia in spring. Twigs
may be attacked as well as foliage.
Mycosphaerella citri. Leaf Spot or Greasy Spot on citrus.
Mycosphaerella colorata. Mountain-Laurel Leaf Spot.

Phyllosticta
kalmicola.
Mycosphaerella (Anamorph, Pseudocercospora cruenta) cruenta. Leaf
Spot, Leaf Blotch of soybean, and kidney bean. Leaf spots distinct to
indistinct, circular to irregular, greenish to yellowish to rusty brown to
almost red, sometimes with gray centers.
Mycosphaerella efgurata (Anamorph, Piggotia fraxini). Ash Leaf Spot,
general east of the Plains. Spots small, purple to brown with yellow borders.
Mycosphaerella fragariae. Strawberry Leaf Spot, Black-Seed Dis-
ease, general on strawberries. Leaf spots are rst purple then reddish with
light brown or white centers, 1/8 to 1/4 inch across. Spots are also present
on petioles and fruit stems, and occasionally there are black spots on fruit,
with blackened achenes prominent against the white of unripe berries. Fruit
is poor; total yield is reduced; runner plants are weakened. Conidia of the
Ramularia stage are produced in clusters of short conidiophores on under-
side of diseased areas; perithecia are formed in autumn at the edge of the
leaf spots where the fungus winters. New conidia are produced in spring
with most infection taking place through stomata. There is a difference in
varietal susceptibility.
Control. Set healthy plants in well-drained soil; remove diseased leaves
before planting; spray with bordeaux mixture before planting and follow
with two or three more applications. The conidia are very sensitive to cop-
per, which prevents sporulation and kills nongerminated spores.
Mycosphaerella fraxinicola (Anamorph, Phyllosticta viridis). Ash Leaf
Spot, east of the Rocky Mountains.
Mycosphaerella juglandis. Leaf Spot of black walnut.
Mycosphaerella liriodendri (Phyllosticta liriodendrica). Tulip-Tree Leaf
Spot.
Mycosphaerella louisianae. Purple Leaf Spot of strawberry, in the South.
Large, irregular, reddish purple areas.
Mycosphaerella mori. Mulberry Leaf Spot, widespread, with the conidi-
al stage reported variously as Cercosporella, Cylindrosporium, Phloeospora,
Septogloeum, and Septoria. Yellow areas on upper leaf surface are matched
by whitish patches underneath, the fungus forming a white downy or pow-
dery coating. The disease is most serious in shady locations.
Mycosphaerella nigromaculans. Black Stem Spot of cranberry, report-
ed from all cranberry areas, often associated with red leaf spot. The fungus
enters through leaves, grows down the petioles, and forms elongated black
spots on the stems, which may be completely girdled, followed by defolia-
tion. Fruiting bodies are produced in autumn on dead stems with ascospores
discharged in rainy periods in spring. The anamorph state of the fungus is
a Ramularia nigromaculans.
Mycosphaerella nyssicola (Anamorph, Phyllosticta nyssae). Tupelo Leaf
Spot, on sour gum and water tupelo. Purplish irregular blotches, an inch
or more across, are scattered on upper leaf surface with lower surface dark
brown. There may be heavy defoliation. Perithecia mature in spring on fallen
leaves.
Mycosphaerella personata (Anamorph, Isariopsis clavispora). Leaf Spot,
widespread on muscadine and other grapes after midseason. Spots are dark
brown, 1/4 to 1/2 inch, surrounded by a yellow circle but with a narrow band
of normal green between spot and circle.
LEAF SPOTS 343
Mycosphaerella populicola (Anamorph, Septoria populicola); M. populo-
rum(S. musiva). Leaf Spot of native poplar; Canker on twigs and branches
of hybrid poplars.
Mycosphaerella psilospora (Anamorph, Septoria querceti). Oak Leaf
Spot on red and other oaks, common in Iowa. Spots very small, circular,
with strawcolor centers and dark margins.
Mycosphaerella ribis. Leaf Spot of owering currant.
Mycosphaerella ribis (M. grossulariae, Anamorph, Septoria ribis). Leaf
Spot of gooseberry, current. Numerous small brown spots with grayish cen-
ters are formed on both sides of leaves; there may be premature defolia-
tion. The fungus winters in leaves, producing ascospores in late spring. Two
sprays of bordeaux mixture plus 1 pint of self-emulsifying cottonseed oil per
100 gallons have given good control of leaf spot on gooseberries in New
York. The rst application is about June 1, the second in July right after fruit
is picked.
Mycosphaerella rosicola.

Cercospora rosicola.
Mycosphaerella rubi.
Septoria rubi.
Mycosphaerella pyri (Anamorph, Septoria pyricola). Pear Leaf Spot,
also on quince, occasional on apple. Spots are small, 1/8 to 1/4 inch, gray-
ish in center, dotted with black fruiting bodies, with a well-dened dark
brown margin. There are marked differences in susceptibility in pear vari-
eties. Flemish Beauty, Duchess, and Winter Nellis are moderately, and
Kieffer very, resistant. Sprays applied for leaf blight or scab control leaf
spot.
Mycocentrospora
Mycocentrospora verrucosa. Leaf Spot on euonymus.
Myrothecium
Sporodochia cushionlike, light or dark; conidiophores subhyaline to colored, repeatedly
branched, bearing conidia terminally; conidia subhyaline to dark, one-celled, ovoid to
elongate; weakly parasitic or saprophytic.
Myrothecium roridum. Leaf Spot on snapdragon, stock, eremurus, garde-
nia, hollyhock, aeschynanthus, aglaonema, aphelandra, dieffenbachia, epis-
cia, ttonia, nematanthus, hoya, peperomia, pilea, and sphathiphyllum. Tis-
sues are dry, brittle, with black sporodochia. Snapdragon leaves and ow-
ering stems wilt, with sunken cracked cankers. Avoid excessive moisture;
sterilize soil.
Nematostoma
Nematostoma occidentalis. Leaf Hair Discoloration on Artemisa.
Neottiospora
Deuteromycetes; Coelomycetes
Pycnidia dark, smooth, innate; spores hyaline, one-celled with two to several appendages
at the apex.
Alpakesa yuccifolia (formerly Neottiospora yuccifolia). Yucca Leaf Spot.
Neottiospora yuccifolia (see Alpakesa yuccifolia). Yucca Leaf Spot.
Ophiodothella
Asci in locules immersed in groups in a stroma, covered by host tissue at maturity;
paraphyses lacking; spores liform.
Ophiodothella vaccinii. Leaf Spot on huckleberry, and farkleberry.
Ovularia
Conidiophores emerging from leaves in clusters, simple or branched; conidia hyaline,
one-celled, ovoid or globose, apical or lateral, single or sometimes catenulate; parasitic.
Ovularia aristolochiae. Leaf Spot on Dutchmans-pipe.
Ovularia pulchella (see Ramularia pusilla). Tan Leaf Spot on creeping
bent grass.
LEAF SPOTS 345
Ramularia pusilla (formerly Ovularia pulchella). Tan Leaf Spot on creep-
ing bent grass.
Pestalotia
Blights.
Pestalotia aquatica (see Pestalotiopsis aquatica). Leaf Spot of arrow-
arum.
Pestalotia aucubae. Aucuba Leaf Spot. The fungus appears as a weak
parasite in sunscald spots or after other fungi.
Pestalotia cliftoniae. Leaf Spot on buckwheat-tree. Ashy or pale brown
spots. Spores usually curved, constricted at septa, three setae at crest.
Pestalotia funerea (see Pestalotiopsis funerea). Leaf Spot, Bark and
Cone Spot on conifers.
Pestalotia guepini (see Pestalotiopsis maculans). Camellia Leaf Spot,
widespread.
Pestalotia leucothos (see Pestalotiopsis leucothos). Leucotho Leaf
Spot, apparently following winter injury or other disease.
Pestalotia macrotricha (see Pestalotiopsis quepini var. macrotricha). Rho-
dodendron Leaf Spot, gray blight, twig blight, widespread on azalea and
rhododendron after winter injury.
Pestalotia palmarum (see Pestalotiopsis palmarum). Palm Leaf Spot,
Gray Leaf. Pestalotia rhododendri (see Pestalotiopsis sydowiana). Rho-
dodendron Leaf Spot.
Pestalotiopsis aquatica (formerly Pestalotia aquatica). Leaf Spot of
arrow-arum. Irregular, chestnut-brown spots, up to an inch in diameter,
have purplish or dark borders and are wrinkled concentrically. Acervuli are
sparse, black, erumpent on upper side of leaf. Spores are ve-celled with
three widely divergent setae.
Pestalotiopsis funerea (formerly Pestalotia funerea). Leaf Spot, Bark and
Cone Spot on conifers. Pathogenicity of the fungus is questionable. Medi-
an spore cells are dark brown; apical hyaline cell has four or ve erect
setae.
Pestalotiopsis leucothos (formerly Pestalotia leucothos). Leucotho
Leaf Spot, apparently following winter injury or other disease.
Pestalotiopsis maculans (formerly Pestalotia guepini). Camellia Leaf
Spot, widespread. Numerous, punctiform black fruiting bodies are scat-
tered over papery gray spots. The spores are ve-celled, bright olivaceous,
with one to four divergent, sometimes branched, setae, and a straight, short
pedicel. This species seems to be a true parasite.
Pestalotiopsis palmarum (formerly Pestalotia palmarum). Palm Leaf
Spot, Gray Leaf. Black pustules are sparsely produced on both surfaces
of pale, dead areas with denite, reddish brown borders. Spores are ve-
celled, with two or three setae, usually knobbed. The fungus is a wound
parasite.
Pestalotiopsis quepini var. macrotricha (formerly Pestalotia macrotricha).
Rhododendron Leaf Spot, gray blight, twig blight, widespread on azalea
and rhododendron after winter injury. Dark or pale spots with black raised
pustules are scattered over stems and leaves. Spots are often silvery gray on
upper surface and dark brown underneath, with densely gregarious acervuli
sooty from dark spores.
Pestalotiopsis sydowiana (formerly Pestalotia rhododendri). Rhododen-
dron Leaf Spot. Black pustules are scattered without order over dried
brown areas of living leaves. Spores are broader than those of P. macrotricha
and have shorter setae.
Pestalozziella
Conidia hyaline, one-celled, with a branched appendage at apex; acervuli subcutaneous;
conidophores slender, simple or branched.
Pestalozziella subsessilis. Leaf Spot on geranium.
Pezizella (Allophylaria)
Apothecia sessile, bright-colored, smooth; paraphyses liform, blunt; spores elliptical to
fusoid, hyaline, one-celled.
Discohainesia oenotherae (formerly Pezizella (Discohainesia) oenotherae).
Leaf Spot, Fruit Rot of blackberry, raspberry, and strawberry; leaf spot
of evening primrose, eugenia, galax, loosestrife, ludwigia, mock-strawberry,
May-apple, peony, and sumac. Spots are irregular, gray in center with a dark
LEAF SPOTS 347
brown border. Fruiting bodies are light amber discs; spores are amber in
masse.
Pezizella (Discohainesia) oenotherae (see Discohainesia oenotherae).
Leaf Spot, Fruit Rot of blackberry, raspberry, and strawberry; leaf spot
of evening primrose, eugenia, galax, loosestrife, ludwigia, mock-strawberry,
May-apple, peony, and sumac.
Phacidium
Blights.
Phacidium curtisii. Tar Spot, Leaf Spot of American holly, more seri-
ous in southern commercial plantings. Small yellow spots appearing in early
summer age to reddish brown with narrow yellow borders. At end of season
at, black, cushion-shaped stromata develop beneath the epidermis. Leaves
seldom drop prematurely, but infected areas may fall out leaving holes. In
years of heavy rainfall berries as well as leaves are spotted. Remove lower
branches; clean up and burn or turn under fallen leaves. Spray with bordeaux
mixture.
Phaeosphaeria
Ascomycetes; Dothideales
Phaeosphaeria maydis. Leaf Spot on sweet corn.
Phaeotrichoconis
Phaeotrichoconis crotalariae. Leaf Spot on areca palm; leaf spots on
palms which are similar in appearance are caused more often by Bipolaris,
Helminthosporium setariae and Helminthosporium (Exserohilum) rostra-
tum.
Phloeospora
Blights.
Phloeospora aceris. Leaf Spot of maple, including vine and dwarf maples.
The spot is small, rather angular, common but not important.
Phoma
Blackleg.
Phoma sp. Leaf Spot on ragweed.
Phomopsis
Blights.
Phomopsis viticola. Leaf Spot on grape.
Phlyctema
Pycnidia dark, separate or sometimes conuent, in or under epidermis or bark; closed
or ostiolate; conidiophores simple or forked; conidia hyaline, one-celled, cylindrical or
long spindle-shaped, mostly bent, sickle-shaped; saprophytic usually.
Phlyctema cuum. Leaf Spot on strangler g.
Phyllachora
Asci in locules, immersed in groups in a dark stroma covered by host tissue at maturity;
spores one-celled, hyaline; paraphyses present; asci cylindrical with short pedicels.
Phyllachora graminis. Tar Spot, Black Leaf Spot, general on wheat-
grass, ryegrass, fescues, redtop, and bent grass. Elongated grayish violet to
dark olive green spots, on both leaf surfaces, turn glossy black. The disease
is seldom serious.
Phyllachora sylvatica. Tar Spot on fescues in Northwest.
LEAF SPOTS 349
Phyllosticta
Blights.
Asterostomella saccardoi (formerly Phyllosticta saccardoi). Rhododen-
dron Leaf Spot, similar to that caused by P. maxima.
Discochora philoprina (formerly Phyllosticta ilicis (Teleomorph, Physa-
lospora ilicis)). Holly Leaf Spot on American and English holly and on
winterberry.
Phoma exigua (formerly Phyllosticta althaeina). Leaf Spot, StemCanker
on abutilon and hollyhock. Ashy spots have black dots of pycnidia. The tis-
sue sometimes becomes brittle and falls away, leaving jagged holes.
Phoma exigua (formerly Phyllosticta decidua). Leaf Spot of agrimony,
aralia, basil weed, betony, cynoglossum, eupatorium, germander, hierachia,
hoarhound, motherwort, lycopus, mint, and monarda.
Phyllosticta althaeina (see Phoma exigua). Leaf Spot, Stem Canker on
abutilon and hollyhock.
Phyllosticta andropogonivora. Leaf Spot on bluestem (Andropogon).
Phyllosticta antirrhini. Snapdragon Leaf Spot, Stem Rot, Blight.
Large circular, dark brown or black spots, with concentric ridges, are located
most often near tips and margins of leaves; centers may be cream to pale
brown, dotted with dark pycnidia. Young leaves may be curled, older leaves
shrivel and hang down along the stem. Petioles are girdled with brown elon-
gated lesions. Stems have rm brown rot with shoots or branches wilting or
have ashy white spots with dark brown or purplish margins and stems crack-
ing in area of spots. Young seedlings may damp off. Spray with bordeaux
mixture; keep greenhouse cool; avoid wetting foliage in watering; clean up
diseased plants.
Phyllosticta aucubae. Aucuba Leaf Spot. Brown or black zonate spots are
mostly along margins of leaves, sometimes with much defoliation. Spores
are exuded from leaves in yellow tendrils, then spread by rain, or syringing
in the greenhouse.
Phyllosticta camelliae (syn. P. camelliaecola). Camellia Leaf Spot.
Lesions are irregular brown spots.
Phyllosticta catalpae. Catalpa Leaf Spot. Dark brown or black spots 1/8
to 1/4 inch in diameter, may run together to give a blotched appearance.
Minute black fruiting bodies pepper the spots, which are often associated
with injury by the catalpa midge. Heavy infection may mean defoliation.
Phyllosticta circumscissa. Leaf Spot, widespread on apricot, peach, sour
cherry, chokecherry, and garden plum.
Phyllosticta concentrica. English Ivy Leaf Spot, also a twig blight,
widespread. Plants look ragged. Fruiting bodies are arranged in spots in
concentric circles.
Phyllosticta cookei. Magnolia Leaf Spot. Spots are grayish without de-
nite margins.
Phyllosticta decidua (see Phoma exigua). Leaf Spot of agrimony, aralia,
basil weed, betony, cynoglossum, eupatorium, germander, hierachia, hoar-
hound, motherwort, lycopus, mint, and monarda.
Phyllosticta ilicis (Teleomorph, Physalospora ilicis) (see Discochora philo-
prina). Holly Leaf Spot on American and English holly and on winterberry.
Phyllosticta hamamelidis. Witch-Hazel Leaf Spot. Small spots enlarge
to reddish brown blotches, causing some defoliation.
Phyllosticta hydrangeae. Hydrangea Leaf Spot, widespread. Brown
spots usually near leaf margins; in severe cases both leaves and blossoms are
killed. Spray with bordeaux mixture.
Phyllosticta kalmicola (Teleomorph, Mycosphaerella colorata). Moun-
tain-Laurel Leaf Spot, Kalmia Leaf Spot. Circular, grayish white to
silvery spots with red or purple borders, up to 1/4 inch across, are sparse-
ly or thickly covered with black pycnidia (see Fig. 3.26). Heavy infection
means disgured foliage and some defoliation. The disease is worse in shady
locations where shrubs are under drip of trees.
Phyllosticta maculicola. Dracaena Leaf Spot. Irregular small brown
spots have yellowish margins and long coils of spores from black pycnidia.
Phyllosticta maxima. Rhododendron Leaf Spot, widespread. Spots are
marginal or terminal, large, dark brown, and zonate.
Phyllosticta minima. Maple Leaf Spot, Gray Spot, also on boxelder,
widespread. Spots are irregular, 1/4 inch or more across, with brownish cen-
ters, containing black pycnidia, and purple-brown margins. The disease is
seldom serious enough for control measures.
Phyllosticta penicillariae. Leaf spot; also stunt chlorosis; on Pennisetum.
Phyllosticta richardiae. Calla Leaf Spot. Small, round, ash-gray spots run
together, producing irregular decayed areas.
Phyllosticta saccardoi (see Asterostomella saccardoi). Rhododendron
Leaf Spot, similar to that caused by P. maxima.
Phyllosticta sanguinariae. Bloodroot Leaf Spot. Spots reddish brown
with a darker border, then a zone of Indian red.
LEAF SPOTS 351
Figure 3.26 Phyllosticta Leaf Spot on Mountain-Laurel
Phyllosticta sojicola. Leaf Spot and Pod Spot of soybean; lesions have
purplish red borders surrounding lighter brownish centers which contain
numerous dark pycnidia (Fig. 3.26).
Phyllosticta vaccinii. Leaf Spot of farkleberry and highbush blueberry.
Also Blueberry; Fruit Rot. Early rot, scald, blast; on cranberry. Small, cir-
cular gray spots, with one to six pycnidia in center, have brown margins. The
disease is unimportant as a leaf spot; fruits have a hard, dry rot.
Phyllosticta wistariae. Wisteria Leaf Spot, more important in the South.
Physoderma
Chytridiomycetes, Blastocladiales
Denite mycelium with terminal and intercalary enlargements which are transformed
wholly or in part into sporangia and resting spores; sporangia rare, oospores abundant,
globose or ellipsoidal. Affected plant parts are discolored or slightly thickened.
Physoderma maydis. Brown Spot of corn, Corn Measles, Corn Pox,
Dropsy, most prevalent in the South. Very small, bleached or yellowish
spots darken to brown or reddish brown with a light margin. Adjacent spots
may coalesce to give the whole blade a rusty appearance. Spots on midrib
and leaf sheath are larger, up to 1/4 inch, irregular to square, darker than leaf
lesions. The entire sheath may turn brown on death of host cells; the epi-
dermis ruptures, exposing brown spore dust. In severe infections low nodes
are girdled so stalks break over. The resting spores remain in soil or plant
refuse over winter, germinating by swarm spores the next spring. A fairly
high temperature and low, wet land favor the disease. Remove plant refuse
early; rotate crops.
Phytophthora
Phytophthora ramorum. Leaf Spot (sudden oak death), California buck-
eye.
Tubackia (Pirostoma)
Pycnidia supercial, with a shield; spores one-celled, dark.
Pirostoma nyssae (see Tubackia dryira). Tupelo Leaf Spot.
Tubackia dryina (formerly Pirostoma nyssae). Tupelo Leaf Spot.
Placosphaeria
Pycnidia globose, dark, in a discoid stroma; spores hyaline, one-celled; teleo-
morph state in Dothideales.
LEAF SPOTS 353
Cheilaria agrostis (formerly Placosphaeria graminis). Tar Spot on redtop
grass.
Placosphaeria graminis (see Cheilaria agrostis). Tar Spot on redtop grass.
Placosphaeria haydeni. Black Spot, Tar Spot on goldenrod and aster,
stems and leaves.
Laestadia (Plagiostoma)
Spores two-celled, hyaline.
Laestadia asarifolia (formerly Plagiostoma asarifolia). Fruit Rot, Early
Rot, Scald, Blast on cranberry, also blueberry. Small, circular gray spots,
with one to six pycnidia in center, have brown margins. The disease is unim-
portant as a leaf spot; fruits have a hard, dry rot.). Leaf Spot on wild ginger.
Mycosphaerella prenanthis (formerly Plagiostoma prenanthis). Leaf
Spot on prenanthis.
Plagiostoma asarifolia (see Laestadia asarifolia). Fruit Rot, Early Rot,
Scald, Blast on cranberry, also blueberry.
Plagiostoma prenanthis (see Mycosphaerella prenanthis). Leaf Spot on
prenanthis.
Pleiochaeta
Pleiochaeta setosa. Leaf Spot on Genista
Pleospora
Perithecia membranous, paraphyses present; spores muriform, dark; some species have
Alternaria, some Stemphylium as anamorph state; wide saprophytic and pathogenic rela-
tionships.
Pleospora herbarum (Stemphylium botryosum; S. sarcinaeforme). Leaf
Spot of clovers, Leaf Blight of lilac, Seed Mold of China aster and
other plants. Spots on legumes are small, irregular, dark brown, sunken,
changing to concentric zonated light and dark brown areas. In nal stages
leaves are wrinkled, dark brown, and sooty. Conidia, like ascospores, are
muriform, olivaceous. Annual phlox has tan lesions. Asparagus has purple
spots.
Pleosphaerulina (Pringsheimia)
Ascomycetes, Sphaeriales, Dothioraceae
Perithecia innate, not beaked, paraphyses and paraphysoids lacking; spores muriform,
hyaline.
Pleosphaerulina sojicola (formerly Pringsheimia sojicola). Leaf Spot of
soybean.
Pseudocercosporella
Pseudocercosporella capsellae. White Leaf Spot on mustard greens.
Pyricularia
Blights.
Pyricularia grisea. Leaf Spot on grass
Ramularia
Conidiophores growing out from host through stoma, clustered, short, dark to hyaline;
conidia hyaline, cylindrical, mostly two-celled, often in chains; found on living leaves
causing leaf spots or white mold.
Cercosporella pastinaceae (formerly Ramularia pastinaceae). Leaf Spot
of parsnip. Lesions are circular, very small, at rst brown, then with a white
center and brown border. Long, slender, septate, hyaline conidia are pro-
duced on exposed conidiophores. No control is necessary.
Entylomella armoraciae (formerly Ramularia armoraciae). Pale Leaf
Spot of horseradish. Few to numerous light green to yellowish spots appear
on leaves in early summer, the invaded areas quickly turning thin and papery
with dead portions dropping out, leaving ragged holes late in the season.
LEAF SPOTS 355
Innumerable small sclerotium-like bodies in the dead tissue carry the fungus
over winter, producing short knobby conidiophores in spring, which either
push out through stomata or break through either epidermis. There is no
special control.
Ramularia armoraciae (see Entylomella armoraciae). Pale Leaf Spot of
horseradish
Ramularia pastinacae (see Cercosporella pastinaceae). Leaf Spot of
parsnip.
Ramularia primulae. Primrose Leaf Spot. Yellow blotches have ash-
colored centers.
Ramularia vallisumbrosae. Narcissus White Mold, sometimes destruc-
tive on Pacic Coast. Small, sunken, grayish or yellow spots appear on
leaves, especially near tips, increasing to dark green to yellow-brown patch-
es, on which, in moist weather, spores are formed in white powdery masses.
The disease may become epidemic with the foliage killed several weeks
before normal ripening. Flower stalks of late varieties may be attacked.
Black sclerotia winter in leaf fragments on ground, producing spores in
spring to infect young shoots.
Control. Spray with bordeaux mixture, starting when leaves are 4 to 6 inches
high. Clean bulbs thoroughly after digging and replant in a new location.
Ramularia variabilis. Foxglove Leaf Spot. Irregular spots, up to 1/4 inch
in diameter, brown with a reddish border, are formed most often on lower
leaves. Spores in tufts give a white, moldy appearance.
Ramulispora
Conidia on sporodochia, two- to many-septate, hyaline to subhyaline, oblong to fusoid,
irregularly united or branched at base; produced in gelatinous masses.
Ramulispora sorghi. Copper Spot of turf grasses, sooty stripe of sorghum,
Sudan grass, and Johnson grass. Black supercial sclerotia are formed on
both leaf surfaces, with conidia in pinkish gelatinous masses. Spots on leaves
are straw-colored with purple borders. Dead areas in turf are small, 1 to 3
inches, copper-red to orange. Velvet bent grass in acid soil is very suscepti-
ble. Liming the soil may help.
Rhizoctonia
Rots.
Rhizoctonia solani. Leaf Spot of tobacco, California pepper-tree (Schi-
nus).
Rhytisma
Apothecia concrete with epidermis and in black, stroma-like spots, tar spots, on leaves;
spores liform, typically hyaline.
Pseudorhytisma bistortae (formerly Rhytisma bistorti). Tar Spot on poly-
gonum. Black tarry spots similar to those on maple.
Rhytisma acerinum. Tar Spot of maple, especially on cut-leaf varieties.
Black, thickened, raised, tarlike spots, up to 1/2 inch in diameter, are formed
on upper leaf surface. They may be numerous enough to cause some defo-
liation but ordinarily are more disguring than destructive. Red and silver
maples are commonly affected in the East. The lesions are light yellow-
green at rst, forming black stomata in summer along with the conidial stage
(Melasmia acerina) (see Fig. 3.27). Ascospores are developed in spring in
tar spots on fallen overwintered leaves and are forcibly ejected, to be carried
by air currents to young leaves overhead.
Control. Collect and burn fallen leaves. Spray in early May with copper,
repeating in 3 weeks in an unusually wet season.
Rhytisma andromedae. Tar Spot on bog rosemary and lyonia.
Rhytisma bistorti (see Pseudorhytisma bistortae). Tar Spot on polygonum.
Rhytisma liriodendri. Leaf Spot on tulip-tree.
Rhytisma punctatum. Speckeled Tar Spot of maple, a black speckled
leaf spot on all species but especially on silver, striped, and bigleaf maple in
Pacic Coast states, rare in the East. Black, raised specks, pinhead size, are
formed in groups on upper leaf surface, in yellow-green areas about 1/2 inch
in diameter. Such areas retain their color even after leaves have faded in the
fall.
Rhytisma salicinum. Tar Spot of willow, on pussy willow and other vari-
eties. Spots are very thick, jet black, denitely bounded, 1/4 inch in diameter.
The fungus winters in old leaves which should be raked and burned.
LEAF SPOTS 357
Figure 3.27 Tar Spot of Maple; black tarry spot on leaf; section through spot; ascus, paraphyses, and liform
ascospores
Schizothyrium
Brown scutellum or shield, radiate at margin, with a single hymenium underneath;
apothecia round to linear, opening with a cleft or lobes; spores hyaline, two-celled.
Schizothyrium gaultheriae (see Schizothyrium pomi). Leaf Spot on win-
tergreen.
Schizothyrium pomi (formerly Schizothyrium gaultheriae). Leaf Spot on
wintergreen.
Sclerotinia
Blights.
Sclerotinia homoeocarpa. Leaf Spot on peanut.
Scolecotrichum
Conidiophores in loose clusters, simple, bearing conidia on pushed-out ends of suc-
cessive new growing points; spores dark, two-celled, ovoid or oblong, often pointed;
parasitic.
Cercosporidium graminis (formerly Scolecotrichum graminis). Brown
Stripe of lawn grasses, Streak of bluegrass and redtop. Grayish brown to
dark linear streaks on leaf blade may extend into leaf sheath and cause defo-
liation. Dark gray masses of conidiophores emerge in rows through stomata
of upper leaf surface.
Scolecotrichum graminis (see Cercosporidium graminis). Brown Stripe
of lawn grasses, Streak of bluegrass and redtop.
Selenophoma
Pycnidia brown, globose, immersed, erumpent, ostiolate; conidia hyaline, one-celled,
bent or curved, typically crescent-shaped, parasitic.
Pseudoseptoria everhartii (formerly Selenophoma everhartii). Speckle,
Leaf Blotch on bluegrass and other grasses. Brown ecks and frog-eye
spots on blades in early spring enlarge to straw-colored blotches scattered
with minute pycnidia. Spots may drop out, leaving holes.
Pseudoseptoria obtusa (formerly Selenophorma obtusa). Speckle, Leaf
Blotch on bluegrass and other grasses. Brown ecks and frog-eye spots on
blades in early spring enlarge to straw-colored blotches scattered with minute
pycnidia. Spots may drop out, leaving holes.
Selenophoma donacis; Syn. Pseudoseptoria everhartii (formerly S. ever-
hartii). Pseudoseptoria obtusa (formerly S. obtusa). Speckle, Leaf Blotch
on bluegrass and other grasses. Brown ecks and frog-eye spots on blades
in early spring enlarge to straw-colored blotches scattered with minute pyc-
nidia. Spots may drop out, leaving holes.
Selenophoma everhartii (see Pseudoseptoria everhartii). Speckle, Leaf
Blotch on bluegrass and other grasses.
Selenophoma obtusa (see Pseudoseptoria obtusa). Speckle, Leaf Blotch
on bluegrass and other grasses.
Septocylindrium
Conidiophores hyaline, short and simple or longer and branched, with irregular some-
what inated cells; conidia hyaline, two- to several-celled, in chains that are sometimes
branched; parasitic or saprophytic.
Septocylindrium hydrophylli. Hydrophyllum Leaf Spot.
LEAF SPOTS 359
Septogloeum
Acervuli subepidermal, erumpent, pale; conidiophores short, simple; conidia hyaline,
several-celled, oblong to fusoid; parasitic.
Diplodia acerina (formerly Septogloeum acerinum). Maple Leaf Spot.
A small leaf spot occasionally defoliating Norway and Schwedler maples in
the Middle West.
Cheilaria agrostis (formerly Septogloeum oxysporum). Char Spot of lawn
grasses. Lesions are tawny with yellow margins, circular becoming ellip-
tical, pointed at each end, covered with dull black to brown stromatic tis-
sue.
Septogloeum acerinum (see Diplodia acerina). Maple Leaf Spot.
Septogloeum oxysporum (see Cheilaria agrostis). Char Spot of lawn
grasses.
Septogloeum parasiticum. Elm Leaf Spot, Twig Blight.
Septogloeum rhopaloideum (Guignardia populi). Grayish brown, circular
or irregular spots on poplar.
Septoria
Blights.
Septoria agropyrina. Brown Leaf Blotch on wheatgrasses.
Septoria bataticola. Sweetpotato Leaf Spot, occasional, most common
in northern tier of sweetpotato states. Minute white spots on leaves are bor-
dered with a narrow reddish zone. Older lesions have one or more pycnidia
barely visible to the naked eye. The spores, oozing out in tendrils when water
is present on the leaf, are spread by rain and insects. No control is needed
except cleaning up crop refuse.
Septoria calamagrostidis. Leaf Spot on bent grasses. Scattered gray to
straw-colored lesions at tip of blades, appearing in Northwest in late winter
and early spring. Seaside creeping bent is especially susceptible.
Septoria callistephi. Leaf Spot, Damping-off, Stem Rot of China aster.
Septoria chrysanthemella and S. obesa. Chrysanthemum Leaf Spot,
also on oxeye daisy, general through eastern and central states to Florida;
also reported in the West. This disease is sometimes confused with nema-
tode injury, but the leaf nematode browns the leaves in wedge-shaped areas
between veins, and the fungi cause denite spots. These are rst small and
yellowish, then dark brown to nearly black. Sometimes the spots coalesce
into blotches; minute black fruiting bodies are faintly visible. Leaves may
turn yellow and drop prematurely or dry and hang down along the stems.
Spores are splashed from plant to plant in watering or rain, and are spread
on cultivating tools.
Control. Avoid syringing greenhouse plants; do not cultivate outdoor plants,
when they are wet.
Septoria citri. Citrus Septoria Spot on leaves but more serious on fruits.
Small, shallow, light brown depressions on green immature fruit retain
a green marginal ring as the fruit colors. Usually a minor trouble, sometimes
important in California.
Septoria citrulli. Watermelon Leaf Spot. The pathogen is like S. cucur-
bitacearum except that spores are shorter. Dogwood Leaf Spot.
Septoria cornicola. Angular lesions between veins are grayish with dark
purple margins.
Septoria cucurbitacearum. Septoria Leaf Spot of cucurbits, on cucum-
ber, winter squash, muskmelon, and watermelon. Foliage spots are small,
gray, circular, rather conspicuous, often bordered with a zone of yellow tis-
sue. The fungus fruits abundantly on upper side of leaves, with long thin
septate spores in black pycnidia. It winters in old plant parts; clean up all
refuse at end of the season.
Septoria cyclaminis. Leaf Spot on cyclamen.
Septoria dianthi. Septoria Leaf Spot of Dianthus, on carnation and sweet
william. Spots are more or less circular, light brown with purplish brown
borders, scattered over leaves and stems, particularly on lower leaves. The
spots may enlarge, and the leaves die. Take cuttings from disease-free plants;
avoid syringing, or do it early in the day.
Septoria divaricatae (see Septoria phlogis). Septoria Leaf Spot of phlox.
Septoria gladioli. Leaf Spot. More important as a hard rot of gladiolus
corms.
Septoria glycines. Brown Spot of soybean. Primarily a foliage disease,
this may also appear on stems and pods. It starts with irregular brown patch-
es on cotyledons, then reddish brown zones on both sides of leaves, often
with pale green or chlorotic zones surrounding the lesions. Spots may cover
the whole leaf, defoliation starting from lowest leaves. Brown discolorations
with indistinct margins extend an inch or more along stems. The pathogen
LEAF SPOTS 361
winters in diseased leaves and in seed. Some varieties are quite resistant. Use
healthy seed; treatment is unsatisfactory; rotate crops.
Septoria lactucae. Septoria Leaf Spot of lettuce, occasionally destructive
to some varieties. Lesions are more common on lower leaves irregular red-
dish marks, dotted sparsely with black pycnidia. The fungus is disseminated
with seed.
Septoria loligena. Leaf Spot on ryegrass, in California. Chocolate brown
spots, paler in the center, surrounded by lighter areas.
Septoria lycopersici. Septoria Leaf Spot of tomato, Leaf Blight, quite
destructive in Atlantic and central states, less important in the South and
West. In seasons with moderate temperature and abundant rainfall enough
foliage is destroyed so that fruits do not mature properly and are subject to
sunscald. The disease appears at any age but more often after fruit is set.
Infection starts on older leaves near the ground, with small, thickly scat-
tered, water-soaked spots, which become roughly circular with gray centers
and prominent dark margins. The spots are smaller, 1/16 to 1/8 inch, and
more numerous than those of early blight. Leaets may die with progres-
sive loss of foliage from the bottom up. The pathogen winters on tomato
refuse and solanaceous weed hosts; spores are washed from pycnidia by rain
or spread by brushing against moist leaves. Optimum temperature is 60 to
80F.
Control. Bury plant remains deep in soil or burn; control weeds; use long
rotations.
Septoria oudemansii. Leaf Spot of bluegrass, in northern states. Dark
brown, purple spots turning straw-colored appear on leaf sheaths and spread
to blades, with turf turning yellowish brown. Plants may be defoliated in cold
wet seasons, but they are rarely killed.
Septoria paeoniae. Septoria Leaf Spot of peony, Stem Canker. Round
gray spots with reddish borders are found on stems and leaves. Control with
sanitary measures.
Septoria phlogis (formerly Septoria divaricatae). Septoria Leaf Spot of
phlox. Dark brown circular spots, up to 1/4 inch in diameter, have light gray
to white centers and often run together in blotches.
Septoria pistaciarum. Leaf Spot on pistachio.
Septoria populicola. Leaf Spot of poplar.
Septoria rubi. (Teleomorph, Mycosphaerella rubi). Blackberry Leaf
Spot on blackberry, and dewberry, perhaps with more than one strain.
See Sphaerulina rubi for forms reported on red raspberry. Leaf spots are
light brown, sometimes with a purple border. Infection is usually so late in
the season that it is of minor importance, but it may cause some defolia-
tion.
Septoria secalis var. stipae. Leaf Spot on bent grass. White spots turn
straw-colored, with scattered pycnidia.
Septoria spraguei. Leaf Spot on Russian wildrye (Elysum).
Septoria tageticola. Marigold Leaf Spot, reported in 1958 from Flori-
da. Spots are oval to irregular, smoky gray to black, speckled with minute
black pycnidia. The disease advances upward from the lower leaves and also
infects younger branches, peduncles, bracts, and seed. African marigolds are
very susceptible, French almost immune.
Septoria tenella. Leaf Spot on fescue grasses. Small, vague, greasy brown
spots.
Septoria tritici var. lolicola. Leaf Spot on ryegrass. Indenite green to yel-
low mottled or blotched spots becoming fuscous to deep brown.
Sphaerulina
Perithecia separate, innate to erumpent, not beaked, lacking paraphyses and para-
physoids; hyaline, with several cells; clavate-cylindrical.
Sphaerulina rubi (Anamorph, Cylindrosporium rubi). Raspberry Leaf
Spot on red and black raspberry only, common east of the Rocky Mountains.
This disease and a similar one on blackberry and dewberry were for many
years considered due to Septoria rubi and then attributed to Mycosphaerella
as the teleomorph state. Later it was shown that two species were involved,
with Sphaerulina the ascomycete on raspberry, Septoria rubi the pathogen
commonly found on blackberry and dewberry.
Spots are small, circular to angular, rst greenish black, then grayish; pyc-
nidia produce elongate, three- to nine-septate spores. Perithecia, formed in
fallen leaves, are black, subepidermal, later erumpent; ascospores are cylin-
drical, curved, pointed at both ends, usually four septate.
Sporonema
LEAF SPOTS 363
Pycnidia dark, membranous or carbonaceous, innate, opening with torn lobes; spores
hyaline, one-celled.
Sporonema camelliae. Camellia Leaf Spot.
Stemphylium
Conidiophores dark, mostly simple, bearing a single terminal conidium or successive
conidia on new growing tips; conidia dark, muriform, smooth or spiny; parasitic or
saprophytic (see Fig. 3.24).
Stemphylium sp. sp. (Teleomorph, Pleospora herbarum). Red Leaf Spot
of gladiolus, widely distributed, causing an annual loss in Florida since 1938.
Spots are small, round, translucent, pale yellow with reddish brown cen-
ters. Leaves may be killed before owering or after spikes are cut, result-
ing in smaller corms. Infection takes place with 10 hours of dew or fog;
rain is unnecessary; optimum temperature is 75F. Leaves may be killed
within 2 weeks of inoculation. Picardy variety is moderately susceptible;
it is damaged more severely when grown near very susceptible Stoplight
and Casablanca. The disease, starting on particularly susceptible varieties,
spreads radially to less susceptible plants, decreasing in severity with dis-
tance from focal point. The leaf spot disappears in summer and autumn,
reappears in winter 3 weeks after a cold period.
Control. Use resistant varieties to separate very susceptible types from those
partly susceptible.
Stemphylium bolickii. Leaf Spot of echeveria, kalancho, and sedum. On
some species lesions are small, raised, irregular to circular, brown to purplish
black. On other species spots are larger, with tan centers, purplish margins.
Stemphylium botryosum (Teleomorph, Pleospora herbarum). Leaf Spot,
Black Seed Rot, Seed Mold on kidney beans, pea, onion, garlic, shallot,
salsify, asparagus, pepper, and tomato.
Stemphyliumcallistephi. Leaf Spot of China aster. Brown, nearly circular,
concentrically zonate spots with dark margins on leaves, bracts, petals, and
stems.
Stemphylium cucurbitacearum. Leaf Spot of cucurbits, on cucumbers,
muskmelon, and winter and summer squash. The pathogen is possibly sec-
ondary, perhaps confused with S. botryosum. Small brown spots with lighter
centers have mycelium growing over the lesion, producing globose, multi-
septate spores.
Stemphyliumlycopersici. Tomato Leaf Spot. Similar to gray leaf spot but
the conidia and conidiophores longer.
Stemphylium solani. Gray Leaf Spot, Stemphylium Leaf Spot in pep-
per, tomato, groundcherry, eggplant, and other Solanum species, mostly in
the South, occasionally a problem elsewhere. In warm, humid weather, plants
are defoliated in seedbed or eld. First infection is on older leaves, which
exhibit numerous small, dark brown spots extending through to the under-
surface. Centers are often a glazed gray-brown with cracking and tearing.
Leaves turn yellow and wither; all leaves may be killed except those at the
top; seedbeds are often destroyed.
Control. Use clean soil for seedbed; spray seedlings at weekly intervals.
Stemphylium vesicarium, Purple Spot of asparagus.
Stigmatea (Stigmea)
Fruiting structure subcuticular, hymenium a single disclike layer covered with a scutel-
lum; spores dark, two-celled; mycelium scanty.
Hormotheca rubicola (formerly Stigmea rubicola). Black Spot of raspber-
ry. Spot formed in late summer with a membranous layer under the cuticle;
fruiting bodies produced in spring.
Stigmea geranii. Black Leaf Speck of Geranium (cranesbill).
Stigmea rubicola (see Hormotheca rubicola). Black Spot of raspberry.
Stigmina (Stigmella)
Conidiophores short, dark, with a single terminal spore; conidia dark, muriform but with
few cells, ovoid to oblong to nearly spherical; parasitic on leaves.
Exosporium liquidambaris (see Stigmina liquidambaris). Leaf Spot on
sweet gum.
Exosporium palmivorum (see Stigmina palmivorum). Leaf Spot of palms,
in greenhouses and in the South.
LEAF SPOTS 365
Stigmina liquidambaris (formerly Exosporium liquidambaris). Leaf Spot
on sweet gum.
Stigmina palmivorum (formerly Exosporium palmivorum). Leaf Spot of
palms, in greenhouses and in the South. Small, round, yellowish transparent
spots run together to form large, irregular, gray-brown blotches; leaves may
die. The disease is more serious with insufcient light. Spores are long, club-
shaped, brown, with many cells. Remove and burn infected leaves. Spray
with bordeaux mixture.
Stigmella platani-racemosae (see Stigmina platani-racemosae). Leaf
Spot of California Sycamore, sometimes causing premature defoliation.
Stigmina platani-racemosae (formerly Stigmella platani-racemosae). Leaf
Spot of California Sycamore, sometimes causing premature defoliation.
Ulocladium
Ulocladium cucurbitae. Leaf Spot on cucumber
LICHENS
A lichen is a fungus body, usually one of the Ascomycetes with apothecia,
enclosing a green or blue-green alga. The fungus receives some food from
the alga and the alga some food and protection from the fungus, a relation-
ship termed symbiotic. Lichens frequently grow on living trees and shrubs,
but their injury is indirect, an interference with light or gas exchange to
stems or foliage, rather than from penetration of living cells of the suscept
plant. There are three types associated with plants: crustose, a crust closely
appressed to bark of main trunk or larger limbs; foliose, leaike, prostrate
but not so rmly attached to the substratum; and fructicose, bushlike, erect
or hanging.
Lichens are more abundant on garden shrubs boxwood, camellias, azaleas,
and so on and on citrus in the South. They ourish in neglected gardens
and orchards, and in shady damp locations, and may sometimes kill twigs
and branches of weak trees growing on poor sites.
In most gardens control is unnecessary. If lichens become too disguring or
too abundant for plant health, they may be killed by spraying affected parts
with bordeaux mixture or other copper spray; spray when the lichens are
dry. They may be removed from main trunks by rubbing the bark with a steel
brush after they are softened by rain.
MISTLETOE
Mistletoes are seed plants belonging to the family Viscaceae. They are semi-
parasites, manufacturing food but depending on a host plant for water and
mineral salts. There are three genera in North America: Phoradendron and
Viscum which are true mistletoes, and Arceuthobium, dwarf mistletoe.
The mistletoe seed is naked embryo and endosperm invested with a brous
coat and borne in white, straw-colored, pink, or red fruits berries
embedded in a sticky gelatinous pulp enabling them to cling to bark of trees
or stick to feet and beaks of birds, which disseminate them.
The seeds can germinate almost anywhere but penetrate only young thin
bark, by means of a haustorium sent out from a attened disc. Branches of
the haustorium extend up and down and around the tree and occasionally
produce secondary haustoria. The number of annual rings on a tree between
the tip of the primary haustorium and the bark tells the age of the mistletoe.
Many are 60 to 70 years old, and one has been reported as living 419 years.
The aerial portions of mistletoes are leafy, evergreen tufts of shoots on the
stems of host plants, most conspicuous on hardwoods after leaf fall (see
Fig. 3.28). The stems and leaves contain chlorophyll and are green but often
with a yellowish, brown, or olive cast, depending on the season. All species
have opposite leaves and round, jointed stems, and are dioecious with incon-
spicuous petal-less owers. They occasionally become so large or numerous
that the weight of the parasite breaks branches of the host. Growth is slow
at rst, but in 6 to 8 years the tufts may be 3 feet across. The aerial part
does not live much longer than that, but the haustoria live as long as the tree,
producing new bunches from adventitious buds.
Because they manufacture their own food, mistletoes require a lot of sun,
which may be one reason why they ourish so in the Southwest. Leafy
mistletoes are relatively harmless in some situations; in others they handi-
cap shade and forest trees, and occasionally kill hackberries and oaks. There
are a few leaf spots and other fungus diseases that keep mistletoes from get-
ting too abundant. They are harvested for Christmas greens with a curved
Figure 3.28 Mistletoe, Common in Southern Trees
mistletoe hook, which can be used to keep aerial portions cut off valuable
trees. Breaking off or cutting off the bunches, however, may lead to more
shoots in an ever-widening area.
Dwarf mistletoes are far more injurious, especially to forest trees, and much
less conspicuous. In western coniferous forests they rank next to heart rots in
importance, reducing the quality and quantity of timber and paving the way
for bark beetle infestations. Infected branches should be pruned out; if the
trunk is infected, the tree should be felled and removed.
MISTLETOE 369
Phoradendron (True Mistletoe)
Phoradendron means tree thief. The genus is restricted to the Americas, ranging from
southern New Jersey and Oregon southward. Most are on hardwoods.
Phoradendron californicum. California Mistletoe, ranging from southern
California to Arizona, chiey on Leguminosae mesquite, carob, squaw-
bush, creosote bush, parkinsonia. This is a leaess species, generally pen-
dent, with long stems and reddish pink berries.
Phoradendron juniperinum. Juniper Mistletoe, a leaess species with
straw- or wine-colored berries, ranging from Colorado and Utah through
New Mexico and Arizona.
Phoradendron libocedri. Incense Cedar Mistletoe, conned to incense
cedar and occurring throughout its range in Oregon, California, and Neva-
da. The pendent plants are leaess with straw-colored berries. It may injure
plants severely, causing spindle-shaped swellings in limbs at point of attack
and living in the trunk as a parasite for hundreds of years after external por-
tions have disappeared.
Phoradendron serotinum (avescens). Eastern Mistletoe, from south-
ern New Jersey west to Ohio and Missouri and south to the Gulf, on many
hardwoods oaks, elm, maple, sycamore, gums, hickory, pecan, hackber-
ry, hawthorn, persimmon, black locust, western soapberry, sassafras, and
trumpet-vine. This species has white berries and is the common Christmas
mistletoe.
Phoradendron tomentosum. Texas Mistletoe, abundant in Texas on elms,
oaks, mesquite, osage-orange, and sugarberry; has white berries.
Phoradendron villosum. Hairy Mistletoe, ranging from Oregon through
California, usually on oaks, also on Oregon myrtle, California buckeye,
chestnut, and manzanitas. It has pinkish white berries and may cause large
hypertrophies on oaks.
Viscum (True Mistletoe)
The genus is restricted to California. It is now known that Luther Burbank introduced
the parasite into the state in about 1900. Burbanks notes indicate that seed was supplied
to him by J. C. Vaughan of Chicago, Illinois. This mistletoe has spread about 3.5 miles
in 75 years.
Viscum album. European Mistletoe, on alder, ash, birch, hawthorn, hick-
ory, buckeye, maple, mountain ash, pear, persimmon, plum, poplar, pyracan-
tha, willow, crabapple, and elm.
Arceuthobium (Dwarf Mistletoe)
The genus is restricted to conifers, and most species are found in the Northwest. Trees
of any age may be deformed or killed, but the greatest mortality is among seedlings and
saplings, with lodgepole and ponderosa pines most susceptible. The most striking symp-
tom is the formation of witches brooms, with sometimes the whole crown transformed
into a huge broom. In other cases fusiform swellings in trunks turn into cankers. Foliage
of affected trees is reduced.
The mistletoes themselves are small, rarely attaining a maximum of 8 inches,
sometimes less than an inch. They are pereninal shoots, simple or branched,
jointed, with leaves reduced to opposite pairs of scales at the top of each
segment. Stems range in color from yellow to brown to olive green. Berries
are olive green to dark blue; each contains a single seed, rarely two. The seed
is ejected with force and is spread horizontally for some feet. Animals and
birds account for infection at a distance.
Arceuthobium americanum. Lodgepole Pine Dwarf Mistletoe, com-
mon on the Rocky Mountain form but not the Pacic lodgepole pine, found
also on r; rare on other pines. The owers bloom in spring, accessory
branches forming a whorl.
Arceuthobium campylopodum. Western Dwarf Mistletoe. It forms
witches brooms and owers late in summer. Widespread in Northwest
principally on coastal ponderosa pine; species that were formerly called
A. campylopodum are A. abietinum on white and grand rs, A. divaricatum
on pinon pines, A. laricis on western larch, A. microcarpum on blue and
Englemann spruce, A. tsugense on western hemlock, A. cyanocarpum on
limber pine. Found also on exotic pines in California.
Arceuthobium cyanocarpum. Dwarf Mistletoe on pine, timber pine, and
hemlock.
Arceuthobium douglasii. Douglas-Fir Dwarf Mistletoe, conned to this
host. Plants are small, only 1 1/2 inches high, greenish, slender.
Arceuthobium occidentale. Dwarf Mistletoe, on exotic pines in Califor-
nia.
Arceuthobium laricis. Dwarf Mistletoe on r and hemlock.
MISTLETOE 371
Arceuthobium pusillum. Eastern Dwarf Mistletoe, the only species in
the East, from Minnesota to New Jersey and north to Canada, common on
spruce, also on tamarack, and pines. The fruit matures in autumn; shoots are
very short, less than an inch.
Arceuthobium tsugense. Hemlock Dwarf Mistletoe on western and
mountain hemlock. Colletotrichum gleosporioides a hyperparasite.
Arceuthobium vaginatum subsp. cryptopodum. Southwestern Pon-
derosa Pine Dwarf Mistletoe. Plant yellowish, robust.
MOLDS
The word mold, or mould, has many meanings. The rst one given in Web-
ster is a growth, often woolly, produced on various forms of organic matter,
especially when damp and decaying, by saprophytic fungi. Leaf mold is
organic matter reduced to friable earth by these saprophytic fungi. When
rhododendrons are fed with a fertilizer having a cottonseed meal base, one
can often see a moldy growth, showing that benecial organisms are at work
breaking down the material for plant use.
Some of these saprophytic fungi have a harmful, parasitic phase. The com-
mon black bread mold, Rhizopus nigricans, causes soft rot of sweetpotatoes
and leak of strawberries and grapes. Penicillium spp., the common blue
molds on jellies, cause a decay of citrus and other fruits. Such diseases are
discussed under Rots.
The word mold is used loosely to cover any profuse fungus growth on the
surface of plant tissue. See Blights for a discussion of Botrytis gray mold,
so common on many plants; see Leaf Spots for Alternaria brown molds and
Ramularia white molds, and for moldy leaf spots due to Heterosporium and
Pleospora; see Sooty Molds for the black growths on insect exudate; and see
Snowmold for turf diseases.
Botryosporium
Conidiophores, tall, slender, hyaline producing numerous lateral branches of nearly
equal length, each producing two or more secondary branches that are enlarged at the tip
and bear heads of conidia; spores one-celled, hyaline; saprophytic.
Botryosporium pulchrum. Leaf Mold on tomato, also geranium (pelargo-
nium), occasional in greenhouses.
MOLDS 373
Chalara (Chalaropsis)
Mycelium at rst hyaline, then greenish; two types of conidia-macroconidia or chlamy-
dospores, olive green, thick-walled when mature, sessile or borne in short conidiophores
in compact groups; endoconidia, hyaline, formed inside end cells of a dark endoconid-
iophore and extruded in chains.
Chalara thielavioides (formerly Chalaropsis thielavioides). Black Mold
of rose grafts. Manetti mold, usually on grafted roses, sometimes on budded
roses in nursery elds. The fungus grows over and blackens cut surfaces of
stock and scion, preventing union and resulting in death of scions. When out-
door roses are budded on Manetti understock, the bud often turns black and
dies. Infection is only through wounds. Rosa odorata and R. chinensis var.
Manetti are both very susceptible understocks; R. multiora is moderately
susceptible; Ragged Robin is immune.
Control. Use healthy understock. Spray greenhouse benches, tools, etc., with
copper sulfate; prevent spread of spores by workmen on hands, clothing, and
budding knife.
Chalaropsis thielavioides (formerly Chalara thielavioides). Black Mold
of rose grafts.
Cladosporium
Blotch Diseases.
Cladosporium fulvum (see Fulvia fulva). Leaf Mold of tomatoes, general
on greenhouse crops, occasionally serious in gardens in wet seasons in the
Southeast and sometimes present in other states.
Cladosporium herbarum. Leaf Mold, Pod and Seed Spot. The fungus
is a weak parasite causing black mold of peanut, pod spot and seed mold
of lima and kidney beans, glume spot of bluegrass, leaf mold of pepper and
tomato, sometimes a fruit mold.
Cladosporium macrocarpum. Black Mold of spinach, on old leaves or
secondary after other leaf spots.
Fulvia fulva (formerly Cladosporium fulvum). Leaf Mold of tomatoes, gen-
eral on greenhouse crops, occasionally serious in gardens in wet seasons in
the Southeast and sometimes present in other states. Diffuse, whitish spots
on upper surface of older leaves enlarge, turn yellow; the undersurface of
the patches has a velvety olive brown coating of spores that are spread by
air currents and in watering. Spores remain viable about the greenhouse for
several months after plants are removed, and are sometimes carried on seed.
Infection occurs only when humidity is high.
Control. Resistant varieties such as Globelle, Bay State, and Vetomold have
been developed, but the fungus has mutated to more virulent forms. Regu-
lating ventilators in greenhouses to reduce humidity seems to be the most
practical control, sometimes providing heat on cool nights, even in summer.
Melanospora (Erostrotheca)
Ascomycetes, Melanosporales
Perithecia bright, more or less soft, without beak, paraphyses lacking; spores ellipsoid,
yellow to olivaceous. Conidial stage has many spore forms.
Erostrotheca multiformis (see Melanospora multiformis (Anamorph, Cla-
dosporium album)). White Mold of sweet pea, White Blight, also on peren-
nial pea, observed on greenhouse crops.
Melanospora multiformis (formerly Erostrotheca multiformis (Anamorph,
Cladosporium album)). White Mold of sweet pea, White Blight, also on
perennial pea, observed on greenhouse crops. Leaets are covered with tan
or buff, circular to irregular, small to large spots with cinnamon brown pus-
tules giving a granular appearance. White tufts of mold represent the Cla-
dosporium stage. Pseudosclerotia are also formed in the leaves, which may
die and drop. The fungus enters through stomata under conditions of high
humidity. Dusting with sulfur has been suggested.
Torula
Conidiophores lacking; entire branches of mycelium develop into simple or branched
chains of dark conidia, which separate readily; saprophytic.
Periconia maculans (formerly Torula maculans). Leaf Mold on yucca.
Torula maculans (see Periconia maculans). Leaf Mold on yucca.
NEEDLE CASTS
Certain diseases of conifers that result in conspicuous shedding of needles
are termed needle casts, sometimes needle blights. Most of the fungi causing
such symptoms are members of the Phacidiales.
Phaeocryptopus (Adelopus)
One of the black mildews, with supercial, dark mycelium; perithecia innate with a cen-
tral foot, without ostiole; spores two-celled, hyaline.
Adelopus gumannii (see Phaeocryptopus gaeumannii). Adelopus Nee-
dle Cast of Douglas-r, Swiss Needle Cast.
Phaeocryptopus gaeumannii (formerly Adelopus gumannii). Adelopus
Needle Cast of Douglas-r, Swiss Needle Cast. Although rst noted in
Switzerland in 1925, this seems to be a native American disease occurring
in relatively harmless fashion on the Pacic Coast, somewhat injurious to
native Douglas-r in the Southwest and to trees in New England and New
York. Needles fall prematurely, leaving only the current seasons growth. If
this happens for several consecutive years, trees have thin foliage, appear
yellow or brown, and nally die. Needles are yellow-green to brown, often
mottled, and on undersurface tiny black perithecia, issuing from stomata,
appear as sooty streaks, one on each side of the middle nerve.
Bifusella
Apothecia elongate, slitting with a cleft; paraphyses lacking; spores hyaline, one-celled,
club-shaped at both ends with halves joined by a narrow neck (Fig. 3.29).
Figure 3.29 Needle Cast Fungi, which form ascospores in hysterothecia, elongate apothecia opening with
a cleft. Bifusella, spores constricted in middle; Elytroderma, fusiform spores; Hypoderma, short fusiform spores;
Hypodermella, spores tapering at base; Lophodermium, ascus with liform spores, and paraphyses
Bifusella abietis (see Isthmiella abietis). Needle Cast of r on alpine and
corkbark r from New Mexico to Idaho.
Bifusella faullii (see Isthmiella faulii). Needle Cast of Balsam r, the most
common and destructive of the needle casts of this host.
Bifusella linearis. Needle Cast of pine, Tar Spot, on various pine species.
Hysterothecia are variable in length, shining black, on two-year needles.
Bifusella saccata. Needle Cast on pine.
Isthmiella abietis (formerly Bifusella abietis). Needle Cast of r on alpine
and corkbark r from NewMexico to Idaho. Dark brown to black hysterothe-
cia extend the entire length of the middle nerve on undersurface of needle.
Pycnidia are in two rows on upper surface.
Isthmiella faulii (formerly Bifusella faulli). Needle Cast of Balsam r, the
most common and destructive of the needle casts of this host. Ascospores
are discharged in July, but infected young needles do not change color until
spring, then turn light brown to buff. Effused pycnidia in the same color
appear in the groove on upper surface of the needle, followed by dusky brown
hysterothecia (apothecia with a covering), with maturing ascospores the sec-
ond summer.
Canavirgella
Canavirgella baneldii. Needle Cast of pine.
Elytroderma
NEEDLE CASTS 377
Ascospores two-celled, broadly fusiform (Fig. 3.29).
Elytroderma deformans. Needle Cast, Witches Broom on Coulter,
ponderosa, lodgepole, Jeffrey, pinon, and Jack pines. Elongated dull, dark
hysterothecia are on both leaf surfaces. The tissues of ponderosa and Jeffrey
pines may be penetrated and loose witches brooms formed. Saplings may
have entire crown converted; they die or make little growth.
Fusarium
Blights.
Fusarium lateritium. Needle Cast on Torreya taxifolia.
Hypoderma
Hysterothecia elliptical to oblong, opening by a cleft; asci long-stalked,
spores hyaline, fusiform, surrounded by a gelatinous sheath (see Fig. 3.29).
Hypoderma desmazierii (see Meloderma desmazierii). Needle Cast, Tar
Spot of pines, most frequent on eastern white pine.
Hypoderma hedgecockii (see Ploioderma hedgecockii). Needle Cast of
hard pines, in southeastern states.
Hypoderma lethali (see Ploioderma lethale). Gray Blight, Needle Cast
of hard pines, from New England to Gulf states.
Hypoderma robustum (see Virgella robusta). Needle Cast of Firs, in
West, usually white r.
Meloderma desmazierii (formerly Hypoderma desmazierii). Needle Cast,
Tar Spot of pines, most frequent on eastern white pine. Infected needles are
at rst yellow, then reddish brown, and nally deep brown with a grayish
cast. The tips are infected rst, the fungus being a weak parasite, completing
its cycle in a year. Hysterothecia are shining black, elliptical.
Ploioderma hedgecockii (formerly Hypoderma hedgecockii). Needle Cast
of hard pines, in southeastern states. Elliptical shining black hysterothecia
are present in discolored areas on green needles. Each ascus contains four
normal and four aborted spores.
Ploioderma lethale (formerly Hypoderma lethaei). Gray Leaf Blight,
Needle Cast of hard pines, from New England to Gulf states. Hysterothe-
cia are short, narrow, black, often found on pitch pine.
Virgella robusta (formerly Hypoderma robustum). Needle Cast of Firs,
in West, usually white r. Concolorous pycnidia, which form two rows, one
in each needle wing, often turn black after spore discharge.
Hypodermella
Like Bifusella with elongate apothecia, with a cleft, but paraphyses present;
spores hyaline, one-celled, club-shaped at upper end, tapering toward base
(see Fig. 3.29).
Davisomycella ampla (formerly Hypodermella ampla). Needle Cast of
jack pine. All needles may drop except those of the current season. Short,
elliptical, dull black hysterothecia are scattered over light buff-colored areas.
Hypodermella abietis-concoloris (see Lirula abietis-concoris). On rs and
southern balsam.
Hypodermella ampla (see Davisomycella ampla). Needle Cast of jack
pine.
Hypodermella concolor (see Lophodermella concolor). Needle Cast of
jack pine and lodgepole pines.
Hypodermella laricis. Larch Needle and Shoot Blight on eastern and
western larches. Yellow spots are formed on needles, which turn reddish
brown but stay attached, giving a scorched appearance to trees that are
normally deciduous. Hysterothecia are very small, oblong to elliptical, dull
black, on upper surface of needles.
Hypodermella nervata (see Lirula nervata). Needle Cast of Balsam.
Lirula abietis-concoloris (formerly Hypodermella abietis-concoloris). On
rs and southern balsam.
Lirula nervata (formerly Hypodermella nervata). Needle Cast of Balsam.
Pycnidia are in a groove along upper surface of needle in continuous or occa-
sionally interrupted row, turning nearly black after spores are discharged.
Lophodermella concolor (formerly Hypodermella concolor). Needle
Cast of jack pine and lodgepole pines. Virulent fungus infects young nee-
dles, in summer, which turn brown the next season. Short hysterothecia are
concolorous with the leaf and appear as shallow depressions.
NEEDLE CASTS 379
Lirula
Lirula macrospora. Needle Cast or Blight on spruce.
Lophodermium
Leaf Spots.
Lirula macrospora (formerly Lophodermium liforme). Spruce Nee-
dle Cast, sometimes causing serious defoliation of red and black spruce.
Hysterothecia are long or short, shining black (see Fig. 3.29).
Lophodermium durilabrum. Needle Cast on pine.
Lophodermium liforme (see Lirula macrospora). Spruce Needle Cast,
sometimes causing serious defoliation of red and black spruce.
Lophodermium juniperinum. Widespread and abundant on common
juniper and red-cedar but apparently not parasitic. Hysterothecia are ellipti-
cal, shining black, on both leaf surfaces.
Lophodermium nitens. Frequent but apparently saprophytic on ve-needle
pines. Hysterothecia short, black, shining.
Lophodermium piceae. Needle Cast, Needle Blight of r, Tar Spot
on r and spruce, most severe on young specimens. Needles turn yellow,
reddish, or brown, and drop. Short, shining black hysterothecia are formed
on all needle surfaces.
Lophodermiumpinastri. Pine Needle Cast, widespread. Pycnidia appear
in spring or early summer as tiny black spots on browned needles, followed
by dull, occasionally shining, black, short, elliptical hysterothecia. The fun-
gus is a weak parasite but can be epidemic in nurseries. Bordeaux mixture
will control it.
Lophodermium seditiosum. Needle Cast of scotch pine.
Mycosphaerella
Blights.
Mycosphaerella laricina. Needle Cast of European larch and western
larch.
Cyclaneusma (Naemacyclus)
Apothecia bright-colored, soft, opening with a cleft; paraphyses much
branched; spores worm-shaped.
Cyclaneusma niveum(formerly Naemacyclus niveus). Needle Cast, occa-
sional on various pines. Fruiting bodies tiny, elliptical, rst waxy, dark
brown, later concolorous with leaf surface.
Naemacyclus niveus (see Cyclaneusma niveum). Needle Cast, occasional
on various pines.
Pestalotia
Blights.
Pestalotia microspora. Needle Spot, Cast of Torreya.
Phoma
Blackleg.
Phoma eupyrena. Needle Cast and Blight of red r and Douglas-r.
Rhabdocline
Ascomycetes
Apothecia innate, brown, exposed by irregular rupture of epidermis; paraphyses present;
spores one-celled, becoming septate after discharge from ascus, rounded at ends and
constricted in the middle.
Rhabdocline pseudotsugae. Needle Cast of Douglas-Fir, Needle Blight,
common on Pacic Coast and in Rocky Mountain States on native Douglas-
r and in northeastern states on ornamental forms. The disease has reached
Europe on trees from western North America and is causing much concern
there.
Needles are infected in spring or early summer, with rst symptoms showing
as slightly yellow spots, usually at ends of needles, in autumn or winter. By
NEEDLE CASTS 381
the next spring the color is reddish brown, and leaves have a mottled appear-
ance. In severe infection needles turn a more uniform brown, and the entire
tree appears scorched. Apothecia are usually on underside of needles, some-
times on upper. They are at rst round cushions; then the epidermis ruptures
to expose a brown, elongated disc. Infected needles drop after ascospore dis-
charge, thereby living only 1 year instead of the normal 8 or so.
Control. Spraying with bordeaux mixture when new needles develop, repeat-
ing twice at 10- to 14-day intervals has been suggested; also, spraying with
lime sulfur at time of ascospore discharge in early summer. In forests, control
will probably depend on early elimination of susceptible trees.
Rhabdocline weirii. Needle Cast of Douglas-r.
Rhizosphaera
Deuteromycetes, Sphaeropsidales, Sphaerioidaceae
Pycnidia brown, on a stalk; spores ovoid, one-celled, hyaline.
Rhizosphaera kalkhofi. Needle Cast of blue spruce. Lowest needles are
affected rst, becoming mottled yellow, and the disease progresses up the
tree. It has been controlled in ornamentals with three sprays of bordeaux
mixture.
NEMATODES
In the six decades since the rst edition of this book was prepared, nematodes
have become of major importance in plant pathology. It used to be stated that
plant pests, insects, and diseases, took a toll of one-tenth of all our crops.
Now we believe that nematodes alone may cause a 10% crop loss, and some
place the gure as high as 25%. The monetary loss is not easy to gure.
Guesses range from $500,000 to $8 billion a year in the U.S. Nematodes
may be as damaging in home gardens as on farms.
Nematodes used to be considered primarily a southern problem, with the
root-knot nematode the major culprit. Now we know that nematodes can be
as serious in Maine or Minnesota as in Florida or Texas, and that root-knot
species are responsible for only a fraction of total losses.
A 1957 report from Maryland states that samples were taken from around the
roots of crop plants on 1210 different farms and gardens, and that every sam-
ple included at least one species of nematode known to be a plant parasite,
with root-knot nematodes making only 3.2% of the total. A 1959 report from
New Jersey states that, on the basis of 2500 soil and root samples taken since
1954, a very conservative estimate of annual loss in the state is $15 million.
The root-knot nematodes which are reduced by cold winters, were in third
place because of their importance as pests of greenhouse crops, including
African-violets, roses, and other ornamentals, as well as vegetable seedlings.
Nematodes (eelworms or roundworms) are threadlike animals in the phy-
lum Nematoda (or Nemata). The following two references were used in the
nematode taxonomic descriptions in this section:
Nickle, W. R. 1991. Manual of Agricultural Nematology. Marcel Dekker,
Inc., New York, NY. 1035 pp.
Blaxter, M. L., DeLey, P., Garey, J. R., Liu, L. X., Scheldeman, P., Vier-
straete, A., Vaneteren, J. R., Mackey, L. Y., Dorris, M., Frisse, L. M., Vida,
J. T., and Thomas, W. K. 1998. A molecular evolutionary framework for the
phylum Nematoda. Nature 392 (6671):7175.
NEMATODES 383
Nematodes live in moist soil, water, decaying organic matter, and tissues of
other living organisms. Some cause diseases of man or animals; others cause
plant diseases. The animal parasites include hookworms, pinworms, and the
worms in pork causing trichinosis, and they range in length from less than
an inch to nearly a yard. Most plant parasites are practically microscopic in
size, sometimes just barely visible to the naked eye. They mostly range from
0.5 to 2 mm long, or from 1/50 to 1/10 inch.
Nematode diseases of plants are not new. The wheat eelworm was recorded
more than two centuries ago (in 1743); root knot has been a recognized prob-
lem since 1855. Our systematic investigation of plant parasitic nematodes is
very new. Only in the past few years have we made surveys to nd out how
widespread nematodes are and how many cases of decline in plants are
due to them. Nematodes injure plants directly by their feeding, causing cell
death or gross modications and general stunting, and indirectly by afford-
ing entrance to bacteria and fungi causing rots and wilts. Some nematodes
also are vectors of ring spots and other virus diseases.
Many nematodes may merely live in the soil close to the plant and cause no
damage, and a few are actually benecial, feeding on such harmful pests as
Japanese beetle grubs. Only an expert nematologist can determine species
and decide which are responsible for a plants ill health. In submitting sam-
ples to your experiment station for diagnosis, dig up roots and some sur-
rounding soil, place immediately in a plastic bag to prevent drying out, and
mail as soon as possible.
Plant parasitic nematodes may be sedentary or migratory. They do not move
through soil to any great distance. Major dispersal is by shipment of infest-
ed nursery stock and soil; locally nematodes are spread on tools, and feet,
in irrigation water, in plant parts, and sometimes as dry cysts by the wind.
Plant nematodes are facultative or obligate parasites. They may be endopar-
asitic, living inside roots or other tissues, or ectoparasitic, living outside the
plant, inserting only the head for feeding; and some forms are intermediate
between the two types. Most plant nematodes are root parasites, but some
live in stems, bulbs, leaves, or buds. Some cause galls or other distinctive
symptoms; others produce a general yellowing, stunting, or dieback that is
often ascribed to other causes.
Nematodes are usually long and cylindrical, tapering at both ends, round
in cross section. In some genera the female is pear-shaped or saclike, but
the male is always vermiform. Nematodes in general lack coloration, being
transparent or with a whitish or yellowish tint. They are covered with a cuti-
cle, made up of three main layers,largely protein, under which is a cellular
layer called the hypodermis. The body cavity, pseudocoel, is lled with u-
id. The body wall musculature, directly beneath the hypodermis, consists of
longitudinal bers only. This means that nematodes cannot contract trans-
versely. They move through moist soil with a threshing motion, or a series
of undulations.
Nematodes have a complete digestive tract with a mouth at the anterior end.
This is surrounded by lips bearing the sensory organs, but there is no true
head, and nematodes lack eyes and nose. Basically there are six lips, but
they may be fused in pairs. The sense organs, amphids, are important diag-
nostic characters, one class of nematodes having amphids with conspicuous
openings, the other having amphids with minute pores. Most plant parasitic
nematodes belong to the latter group.
Behind the mouth there is a cavity (stoma), then the esophagus, the intestine,
and the rectum. The latter terminates in a ventral terminal or subterminal
anus in females, in a cloacal opening in males. The sexes are usually sepa-
rate, but sometimes males are missing and females are hermaphroditic. The
body region behind the anus or cloacal opening is called the tail.
Near the posterior end of many nematodes there is a pair of cuticular pouches
called phasmids, believed to be sense organs like the amphids. They are used
to divide nematodes into two main groups, the Secernentea, or Phasmidia,
with phasmids, and the Adenophorea, without phasmids.
All of the plant parasitic nematodes feed by means of a stylet, which works
something like a hypodermic needle. It is a conspicuous protrusible spear
used to puncture tissue. In most families this is a stomatostylet, a hollow
spear derived from the sclerotized walls of the buccal cavity or stoma.
Commonly the nematode punctures plant tissue with its stylet, then injects
a secretion from its salivary gland that predigests the food before it is sucked
in through the stylet. In the family Dorylaimidae the spear is an enlarged
tooth, odontostylet, originating in the esophagus wall. It is usually hollow,
but in the genus Trichodorus the tooth (onchiostyle) is solid but grooved.
The structure of the esophagus varies in different groups and is an important
diagnostic character. The esophagus commonly has one or two swellings,
known as bulbs. Those provided with a glandular apparatus are true bulbs;
those lacking such apparatus are pseudobulbs. True bulbs are the chief pump-
ing and sucking structures. They may be median, situated at midlength, or
posterior, at the end of the esophagus.
NEMATODES 385
Control measures for nematodes include crop rotation and other cultural
practices and soil treatment with chemicals. Most chemicals are meant for
fallow soil; a few are safe around living plants. Details of nematicides and
their application are given in

Chap. 1. Greenhouse soils are often steam-
sterilized, and plants are sometimes dipped in hot water, the duration of the
soak and the temperature depending on the tolerance of the plant and the
kind of nematode to be eradicated. Some plants are antagonistic to nema-
todes. Asparagus roots produce a chemical that is toxic to many species, and
marigolds grown with or in advance of some ower crops reduce the num-
bers of Pratylenchus, lesion nematodes. Some soil fungi trap nematodes but
do not provide a practical control. The endospore-forming bacterium Pas-
teuria penetrans is known to effectively suppress certain root-knot nema-
todes.
Anguina
Anguinidae. Endoparasitic nematodes feeding in above ground plant tissue and trans-
forming seeds or leaves into galls. Males and females both elongate (wormlike), but
females are obese. Cuticle nely striated; stylet short with well-developed basal knobs;
tail coneshaped; single ovary.
Anguina agrostis. Grass Nematode, serious on bent grass and chewings
fescue in the Pacic Northwest. Second-stage larvae remain in sheaths near
growing tips most of the year, entering embryonic owers in late spring.
There the larvae mature, and the females lay large quantities of eggs. The
quickly hatching young larvae transform developing seed into elongated dark
purple galls. When the galls fall to the ground, nematodes are released to
reinfect grass in the vicinity. There is only one generation a year, and larvae
cannot exist in moist soil more than a year without access to a host plant with
developing inorescence. The disease is important only on grass grown for
seed; it is not a problem on clipped turf. When seed is threshed, galls can be
carried 300 feet or more from the machines by air currents, and still further
in heavy winds.
Control. Rotate with a crop other than bentgrass or fescue or plow under and
prevent inorescence for 1 year. Soak seed for 2 hours in tepid water with
a wetting agent; then hold for 15 minutes at 126F.
Anguina balsamophila. On balsam-root; galls on underside of leaves.
Anguina graminis. Galls on leaves of fescue grasses.
Anguina tritici. Wheat Nematode on wheat and rye, a eld crop pest
forming galls in place of grain. The disease was recognized in 1745, the rst
to be attributed to nematodes. The species is long-lived, viable nematodes
having been found in seed stored 28 years. Brine otation was the old method
of eliminating galled seed.
Aphelenchoides
Aphelenchoididae. Bud and leaf nematodes, foliar nematodes. Ecto- and endoparasites;
males and females wormlike, very slender; cuticle nely annulated; stylet with small
basal knobs; tail with acute tip.
Aphelenchoides besseyi (including A. oryzae). Summer Dwarf Nema-
tode of strawberry, present from Maryland to Louisiana, also reported from
Oklahoma, Missouri, southern Illinois, California, and Washington. The
nematodes live in the soil and are washed into buds by rains and irriga-
tion water, affecting young leaves as they develop. Leaets are crimped or
crinkled, cupped, narrow, with a reddish cast to veins and petioles. Older
leaves are darker green, more brittle than normal. This is a major disease
in Florida, commonly noted from July to October. Cold weather checks its
progress often masking symptoms, but plants do not recover; runner plants
from infested mother plants are diseased. In spring the nematode population
may be low, allowing nearly normal formation of early leaves, but in summer
a single bud may harbor up to 1300 individuals, causing center leaves to be
deformed and dwarfed. The same species causes a serious disease of rice in
Arkansas and Louisiana.
Control. Buy certied plants; rogue and burn diseased plants as soon as
noticed. Treat dormant infested plants with hot water, 2 minutes at 127F.
Aphelenchoides fragariae (including A. olesistus). Spring Dwarf Nema-
tode of strawberry; Fern Nematode, a leaf nematode. A bud parasite of
strawberry from Cape Cod to Maryland and found in scattered localities
along the Pacic Coast. This is a cold-weather species, persisting through
the winter with several thousand nematodes present in a single bud as leaves
unfold in spring. The foliage is small twisted, thickened, glossy, with swollen
petioles; blossom buds are killed or poor, and no fruit is set. Some plants are
killed; others recover.
As the fern nematode, or begonia leaf blight nematode, this species is record-
ed on anemone, aquatic plants (Cabomba sp., Limnophila sp., Peplis sp., and
NEMATODES 387
Potamogeton sp.), begonia, bouvardia, calceolaria, chrysanthemum, clema-
tis, coleus, crassula, dianthus, doronicum, fern, geranium, hosta, hydrangea,
Lamium, lily, peony, primrose, saintpaulia, scabiosa, zinnia, and other orna-
mentals. Fern leaves have a patchy or blotched appearance with dark brown
to black areas on the fronds. In some species these are rather narrow dark
bands from midrib to border, limited by parallel side veins; in birds-nest
fern there is a profuse brown discoloration from the base halfway up the
leaf.
On begonias the disease is most serious on semituberous varieties grown in
greenhouses. Small brown spots with water-soaked margins, on underside of
leaves, enlarge, coalesce, turn dark brown, and become visible on the upper
surface. Whole leaves may turn dark; plants may be stunted. On brous-
rooted begonias spots stay small, and leaves become shiny with a tendency
to curl, lose color, and drop. Nematodes are spattered from plant to plant
by syringing or careless watering; there is no disease spread when foliage is
kept dry.
Dieback of Easter lilies grown in the Northwest is also attributed to this bud
and leaf nematode. Leaves are rst blotched with yellow, then turn brownish,
drooping and curling against the stem (see Fig. 3.30). The nematodes live
over in the bulbs and are splashed from leaves of one plant to another in
the eld. Lilies from diseased bulbs develop bunchy-top symptoms, with
thick, twisted foliage and dieback.
Control. Strawberry plants in nurseries should be inspected and certied in
spring. Mother plants, near the end of the dormant period, can be treated with
hot water, 2 minutes at 127F. Crop rotation helps.
Bulbs may be treated with hot water, for 1 hour at 111F. Potted begonias
can be submerged, pot and all, for 1 minute at 120F, or for 3 minutes at
116F. African violets may be treated for 30 minutes at 110F, ferns for 10
to 15 minutes at the same temperature.
Aphelenchoides parietinus. Causing root-plate and scale necrosis of bul-
bous iris.
Aphelenchoides ritzemabosi. Chrysanthemum Foliar Nematode, com-
mon and serious on this host in home gardens and greenhouses, rst reported
in New Jersey in 1890. It is also recorded on dahlia, zinnia, and some oth-
er ornamentals but possibly confused with A. fragariae. A morphological-
ly similar species produces a yellow bud blight of Vanda orchids. The rst
symptoms are dark spots on areas on underside of leaves, but by the fth day
after infestation discolored veins stand out sharply on upper leaf surface, and
Figure 3.30 Foliar Nematode on Lily
diseased leaves turn brown or black, starting in distinctive wedge-shaped
areas between veins (see Fig. 3.31). Later the leaves dry, wither, and hang
down along the stems. The nematodes swim from the soil up the stem in
a lm of water, the disease going from lowest leaves progressively upward.
Almost any variety may be attacked, but Koreans are particularly suscepti-
ble. The nematodes may not survive the winter in old dead leaves but they
do survive in living leaves in old crowns.
Control. Keep foliage dry; avoid overhead watering. Use a mulch to avoid
splashing. Avoid crown divisions; make tip cuttings which are usually free
from nematodes. Dormant plants can be treated with hot water, 5 minutes at
122F or 30 minutes at 112F.
NEMATODES 389
Figure 3.31 Leaf Nematode of Chrysanthemum. Wormlike male and female nematodes cause wedge-shaped
browning between veins, followed by general blighting of leaf
Aphelenchoides ritzemabosi. Current Nematode, a bud parasite on black
currants and gooseberries in England; reported from California on gooseber-
ries. Treat cuttings for 30 minutes in hot water, 110F.
Aphelenchoides subtenuis. Bud and Leaf Nematode on narcissus, caus-
ing scale necrosis. Reported from the Southeast and Pacic Coast states.
Belonolaimus
Belonolaimidae. Sting nematodes, migratory obligate ectoparasites, usually found free
in soil near growing tips; both sexes long, slender, with blunt ends; body strongly annu-
late; about 2 mm long, stylet long, with well-developed knobs; two ovaries.
Belonolaimus gracilis. Sting Nematode on a wide variety of hosts from
Virginia southward, also reported from New Jersey and from a rose green-
house in Connecticut. This is a major pest of strawberries, celery, and sweet
corn in Florida. It injures Bermuda, centipede, and St. Augustine grasses and
seedlings of slash and long-leaf pines, being rst recorded from pine. Other
plants damaged by Belonolaimus species include peanut, pea, lupine, Austri-
an winter pea, cowpea, bean, lima bean, soybean, beets, cabbage, cauliow-
er, lettuce, endive, onion, potato, and sweetpotato. The slender worms feed
at root tips and along the sides. Soil fungi enter roots through feeding punc-
tures. Roots develop short stubby branches with necrotic lesions; plants are
stunted. On woody plants decline symptoms include chlorosis, twig dieback,
premature dropping of fruit (such as grapefruit), and rapid wilting under
moisture stress. The nematodes seem to be limited to light, sandy soils.
Control. Rotate crops; cultivate to remove weed hosts.
Belonolaimus longicaudatus. This species may be responsible for some of
the injury ascribed to B. gracilis. It occurs in the same southeastern states
and may injure roots of celery, peanut, grasses, cabbage, bean, and other
vegetables. Potato and soybean are considered especially susceptible. It has
also been reported on magnolia.
Bursaphelenchus
Aphelenchoididae. Ecto - and endoparasites; females (adult) have a vulval ap.
Bursaphelenchus lignicolus. Causes wilt of pine and the nematode is vec-
tored by cerambycid beetle (pine sawyer beetle).
Bursaphelenchus xylophilus. Pinewood Nematode on pine.
Cacopaurus
Tylenchulidae. Cuticle nely annulate; female small but very obese; eggs large; male
lacks stylet.
Cacopaurus pestis. Reported from roots of Persian (English) walnut in Cal-
ifornia, causing typical decline with reduction in size and number of leaves,
fewer nuts, eventually complete defoliation and death.
Criconema
Criconematidae. Ring nematodes, short, thick, sedentary ectoparasites; cuticle thick with
spines or scales; usually found in woodlands, in damp areas, seldom in cultivated soil.
Criconema civellae. Reported on citrus roots in a Maryland greenhouse.
Criconema decalineatum. Fig Spine Nematode on gs.
Criconema spinalineatum. Zoysia Spine Nematode on Zoysia.
Mesocriconema
Criconella xenoplax (see Mesocriconema xenoplax). Ring Nematode on
peach cover crops including curly dock, perennial ryegrass, vetch, crimson
clover, hairy vetch, and cowpea; also tall fescue, and white clover.
NEMATODES 391
Mesocriconema xenoplax (formerly Criconella xenoplax). Ring Nema-
tode on peach cover crops including curly dock, perennial ryegrass, vetch,
crimson clover, hairy vetch, and cowpea; also tall fescue, and white clover.
Criconemoides (Genus dubium)
Criconematidae. Ring nematodes; short, thick-bodied; cuticle thick with retrose (inclin-
ing backward) annules; ectoparasites with a wide host range
Criconemoides annulatum. On holly oak, Montana; beans and citrus,
Louisiana.
Criconemoides citri. Citrus Ring Nematode on citrus in Florida. The
broadly annulated head is often buried deep in root tissue, which dies near
the feeding puncture.
Criconemoides crotaloides. On Douglas-r and poplar, Utah.
Criconemoides curvatum. Reported in large numbers on carnations but
apparently not very injurious; also on grasses, Ohio.
Criconemoides cylindricum. On peanut, in Georgia.
Criconemoides komabaensis. On camellia, in Florida.
Criconemoides lobatum. On pines, Florida; potato, New York; also grasses.
Criconemoides mutabile. On marigold, D.C.
Criconemoides ornatum. On grasses, Ohio.
Criconemoides parvum. On grasses, Ohio.
Criconemoides rusticum. On grasses, Ohio.
Criconemoides similis. Cobbs Ring Nematode. Apparently an impor-
tant factor in decline of peaches in Maryland and North Carolina, reported
on pine in Florida and North Carolina.
Criconemoides teres. On oak, California.
Criconemoides xenoplax. On carnation, causing reduced root system, stunt-
ing, reduced ower yield; also reported on grape, peach and grasses.
Crossonema
Crossonema sp. Decline of Alaska cedar.
Ditylenchus
Anguinidae. Bulb and stem nematodes, slender, of moderate length, conelike tail, nely
striated cuticle, mostly endoparasites.
Ditylenchus destructor. Potato Rot Nematode. Feeding on underground
stem structures of a large number of plants but important on potato, especial-
ly in Idaho and Wisconsin. Discolored spots on tubers progress to a gray or
brown decay. The tissues have a granular appearance; they dry and shrink
and the skin may crack. Invasions continue in storage, sometimes with com-
plete destruction of tubers.
Ditylenchus dipsaci. Stem and Bulb Nematode. An internal parasite
of bulbs, stems, leaves, rarely roots, causing Eelworm Disease of Narcis-
sus, Ring Disease of Hyacinth, Onion Bloat, Stem Disease of Phlox. The
name dipsaci covers many strains and probably more than one species. The
type was found in 1857 on Fullers teasel. The nematodes are thought to
release a pectinase during feeding, which results in a dissolution of the mid-
dle lamella and the production of large intercellular spaces. They injure,
besides hyacinth and narcissus, grape-hyacinth, tulip, galtonia, garlic, shal-
lot, and onion, and cause a stem disease of alfalfa and many owers besides
phlox.
The strains of hyacinth and narcissus are not reciprocally infective, although
the hyacinth strain does infect onions. Hyacinths have yellow ecks or
blotches on the leaves, which are often twisted, short, and split. In narcissus
there are pustules or blisters, called spikkels, in leaves, which can be felt
when the leaf is drawn through the ngers. Nematodes in such pustules
probably enter leaves as they push up through the soil. Bulbs badly diseased
at planting produce no foliage, or a few leaves that are premature, twisted,
and bent.
When leaves are dry, nematodes are inactive; but when the foliage is moist
and decayed, they revive and pass down into the soil or the neck of the bulb.
They enter bulb scales, move down to the basal plate, and then enter the base
of other scales. Infected scales are brown, and, since there is little lateral
movement of nematodes, the cut surface of a bulb shows one or more brown
rings contrasting with healthy tissue. Eggs, larvae, and adults are all present
in the brown areas. Male and female adults are wormlike, up to 1.9 mm long.
Infective larvae issue in large numbers in whitish tufts in a break between
basal plate and scales, and work through the soil to invade adjacent plants.
NEMATODES 393
They are also spread in irrigation water, on tools, and by animals. Some
winter in weed hosts, some in seed of composites. In moist soil they die in
a year or so, but they have been recovered from plants after 5 or 6 years.
The strain on phlox attacks campanula, sweet william, evening primrose,
goldenrod, schizanthus, anemone, foxglove, and orchids. The leaves are very
narrow, crinkled, and waved, often brittle, with a tendency to lengthen peti-
oles. Stems may be swollen near the top or bent sidewise; plants are stunted,
often fail to bloom, may die prematurely. The nematodes enter through stom-
ata of young shoots and work upward as the stems develop. They infest seed
of phlox and other composites, and may be so disseminated.
In onions the inner bulb scales are enlarged, causing a split onion that sel-
dom owers and sometimes rots at the base. Seedlings are twisted, stunted,
covered with yellow spots. On plants grown from sets, a slight stunting and
accid condition of outer leaves is followed by leaf-tip necrosis and contin-
ued stunting. The larvae may live long in infested soil and may be carried in
set onions.
Control. Commercial growers routinely treat narcissus bulbs in hot water,
4 hours at 110 to 112F. All infected plants, parts, and debris should be
removed from elds and destroyed; a 2- to 4-year rotation may be tried.
Take up and burn infested phlox or similar plants. Put new plants in a new
location or in fumigated soil.
Ditylenchus (Sychnotylenchus) gallicus. On elm.
Ditylenchus iridis. Probably a strain of D. dipsaci, on bulbous iris. Mild-
ly infected plants dry up prematurely and have poor root systems. Heavi-
ly infected plants are stunted, having few if any roots, and the bulbs decay
before harvest. Treat bulbs with hot water as for narcissus, but soak only 3
hours and as soon after curing as possible.
Dolichodorus
Dolichodoriadae. Awl nematodes similar to sting nematodes with long stylet with well-
developed knobs; coarsely annulated cuticle; both sexes wormlike; male tail has a bursa
(lateral extension); female has two ovaries; ectoparasites.
Dolichodorus heterocephalus. Awl Nematode, causing decline of cel-
ery, bean, tomato, corn, pepper, and water chestnut in the Southeast, also
recorded on pecan. It feeds largely on root tips and sometimes along the side
of roots, causing necrotic lesions. It also feeds on germinating seeds and
hypocotyls, sometimes penetrating the seedcoat to reach the embryo. Poor
seedling emergence may be due to this nematode.
Dolichodorus (Neodolichlodorus) obtusus. On arctostaphylus and pecan,
California.
Dorylaimus
Dorylaimidae. Spear nematodes, with an odontostylet (hollow tooth), bottle-shaped
esophagus; cuticle with longitudinal ridges; both sexes wormlike, tails rounded to
cone-shaped; not proven plant parasites.
Dorylaimus spp. Found in soil near soybean, sweetpotato, and other plants
but not known as a pathogen.
Helicotylenchus
Hoplolaimidae. Spiral nematodes, ectoparasites or semiendoparasites; long strong stylet
with basal knobs; cuticle annulated. The head is inserted in a root, but the body remains
outside in a ventrically curved spiral with one or more turns.
Helicotylenchus dihystera. On gardenia, corn, and bluegrass.
Helicotylenchus erythrinae. Zimmermans Spiral Nematode. Rather
common in Florida around roots of grasses. Present in other states on blue-
berry, boxwood, cauliower, cedar, clovers, corn, cranberry, turf grasses,
oak, oat, pachysandra, pepper, pieris, pine, rhubarb, soybean, strawberry,
wheat, and yew.
Helicotylenchus multicinctus. Cobbs Spiral Nematode. Associated
with roots of many plants, including azalea, cherry, cranberry, marsh grass,
hibiscus, peach, pine, spruce, and yew.
Helicotylenchus nannus. Steiners Spiral Nematode, a small species
common in the Southeast. Found damaging roots of apple, azalea, boxwood,
asparagus fern, calathea, camellia, centipede grass, civet bean, gardenia,
peperomia, philodendron, rubber-plant, royal palm, laurel oak, soybean,
peanut, and tomato. There is a gradual decline, stunting, and failure to form
ower buds.
Helicotylenchus pseudorobustus. On corn, grape, tomato, and soybean.
NEMATODES 395
Hemicriconemoides
Criconematidae. Ectoparasites; female with cuticular sheath, anchor-shaped stylet with
anteriorly concave knobs; males without sheath or stylet. Commonly associated with
turf and woody plants in warm climates, but pathogenicity not yet demonstrated.
Hemicriconemoides biformis. Oak Sheathoid Nematode. On roots of
oak, Florida.
Hemicriconemoides chitwoodi. Associated with stunting of camellias.
Hemicriconemoides oridensis. Pine Sheathoid Nematode. On pine.
Hemicriconemoides gaddi. On camellias.
Hemicriconemoides wessoni. On myrica, Florida.
Hemicycliophora
Criconematidae. Sheath nematodes; ectoparasites with sedentary habits; female retains
last molt as an extra cuticle; knobs of stylet spheroid; males rare, without stylet.
Hemicycliophora arenaria. Causing root galls on rough lemon, also repro-
ducing in tomato, pepper, celery, squash, and bean. Celery has large, multi-
branched galls.
Hemicycliophora brevis. On California-laurel.
Hemicycliophora obtusa. On beet, Utah.
Hemicycliophora parvana. Tarjans Sneath Nematode, damaging cel-
ery in Florida, also recorded on corn, beans, and dracaena.
Hemicycliophora similis. Grass Sneath Nematode. Also causes small
galls on roots of blueberry and cranberry.
Heterodera and Globodera
Heteroderidae. Cyst nematodes, highly specic, attacking members of but few genera
in a given plant family, partially endoparasitic, quite sedentary, attached to root by neck
only. The female is lemon-shaped to globoid, white, yellow, or brown, 0.5 to 0.75 mm.
Eggs are deposited or retained in body of mother, whose leathery wall forms a true cyst.
Eggs remain alive for years in cysts, which are spread by wind or in soil around nonhost
plants. Males are slender worms, up to 1.75 mm. Root-knot nematodes, formerly all
classed as Heterodera marioni, have been reclassied as various species of Meloidogyne.
The stylet Heterodera is twice as long as that in Meloidogyne, and the latter does not
form true cysts.
Globodera rostochiensis (formerly Heterodera rostochiensis). Golden
Nematode on white potatoes, also eggplant, tomato, and other members
of the Solanaceae, but not on tobacco. It was rst discovered in the United
States on Long Island in 1941, and it was kept there, by a rigorous quar-
antine, until 1967, when it was found at a single location in upstate New
York. In 1968, it was found on a potato farm in Delaware. Known as potato
sickness, the disease has been serious in the British Isles for many years.
Crops do not show much damage until heavy populations have built up in
the soil; then there is midday wilting, stunting, poor root development, early
death, with up to 85% reduction in potato yield. The eggs live in the soil
inside cysts barely visible to the naked eye. Each may contain up to 500
eggs, and some hatch one year, some another. Cysts have remained viable
17 years. In spring, when soil temperature is around 60F, a chemical given
off by potato or tomato roots stimulates hatching, and the larvae (which have
had a rst molt inside the egg) leave the cysts and migrate to host plants,
entering the roots. The females become stationary, swell to pear shape, and
break through the roots, remaining attached by a thin neck. The cylindrical
males work out of the roots and cluster around to mate with the females.
Eggs are formed, and the dead female becomes the cyst, rst white, then
gold, orange, nally brown. Cysts detached from roots remain in the soil
or may be spread in potato bags, crates, machinery, even in trouser cuffs of
farm workers. Lily-of-the-valley pips, cacti, and other plants intercepted at
quarantine have had golden nematode cysts in fragments of soil around the
roots.
Control. A quarantine restricts movement of potatoes, nursery stock, root
crops and top soil from infested land. Healthy potatoes are sold in paper bags
to prevent reinfestation from secondhand burlap bags. The Peconic strain of
potato is said to be resistant; Rosa, Elba, and NY 71 are also resistant.
Heterodera avenae. Oat Cyst Nematode on pea.
Heterodera cacti. Cactus Cyst Nematode. Obtained from various local-
ities in Mexico, where it is probably indigenous, and likely to occur on cacti
wherever grown. The cyst is lemon-shaped.
Heterodera carotae. Carrot Cyst Nematode.
Heterodera cruciferae. Cabbage Cyst Nematode, closely related to
the sugarbeet nematode. On crucifers in California. Hosts include broc-
coli, Brussels sprouts, cabbage, cauliower, kale, kohlrabi, mustard, radish,
rutabaga, seakale, lobularia, sweet alyssum, wallower, and garden cress.
NEMATODES 397
Heterodera ci. Fig Cyst Nematode on g in Florida and California.
Heterodera glycines. Soybean Cyst Nematode causing Yellow Dwarf
Disease. An immigrant from Japan and Korea, rst noted in North Carolina
in 1954, thence spread to Arkansas, Florida, Illinois, Kentucky, Louisiana,
Mississippi, Missouri, Tennessee, and Virginia. Plants are yellow, stunted;
roots are small and dark with few or no bacterial nodules but with lemon-
shaped brown cysts clearly visible. This nematode reproduces only in roots
of lespedeza, vetch, tomato and bean, besides soybean, but the cysts occur as
contaminants of narcissus bulbs and gladiolus corms grown in infested soil
and may be so disseminated.
Infested areas are under federal and state quarantines. Soil fumigation tem-
porarily reduces nematode populations and increases plant growth and yield.
Heterodera gottingiana. Pea Cyst Nematode on pea.
Heterodera humuli. Hop Cyst Nematode on bean, pea, and cucumber.
Heterodera iri. On grasses.
Heterodera mothi. Cyst Nematode on nutsedge.
Heterodera punctata (Punctodera punctata). Grass Cyst Nematode
found on wheat and small grains, also associated with bentgrasses in North
Dakota, Michigan, and Minnesota, and turfgrass in New Jersey.
Heterodera rostochiensis (see Globodera rostochiensis). Golden Nema-
tode on white potatoes, also eggplant, tomato, and other members of the
Solanaceae, but not on tobacco.
Heterodera schachtii. Sugar Beet Nematode, occurring in sugar-beet
areas from California to Michigan, also infesting table beets and crucifers
cabbage, broccoli, rape, turnip, rutabaga, and radish. The females, numerous
white specks clinging to roots, contain 100 to 600 eggs. Slender larvae punc-
ture root cells with their strong stylets and pass through three molts inside
the roots. The wormlike males then leave the roots to search for the ask-
shaped females, which are attached to the roots only by their heads. Eggs are
deposited in a gelatinous mass. These soon hatch to start other generations,
but the females die with more eggs inside their bodies, which turn brown
and become cysts. Eggs inside cysts may remain viable 5 or 6 years. Control
depends on a very long crop rotation or soil fumigation.
Heterodera tabacum ((Globodera tabacum tabacum). Tobacco Cyst
Nematode. Reported from Connecticut on tobacco, tomato, and other
solanaceous plants, but not potato; also reported on Jerusalem-cherry, egg-
plant, and pepper in Virginia. Stunting is also caused on tobacco.
Heterodera trifolii. Clover Cyst Nematode. On clover and other legumes
except peas. Spinach, beet, soybean, and carnation are minor hosts. Cysts are
brown, lemon-shaped.
Heterodera zeae. Corn Cyst Nematode. On sweet corn, eld corn, and
barley.
Hoplolaimus
Hoplolamidae. Lance nematodes, somewhat migratory, some species tropical or sub-
tropical, of moderate length; strong stylet with basal knobs; often in a spiral or C-shape
position.
Hoplolaimus coronatus. On Nerine, grasses, carnation, oak, citrus, pine,
sweetpotato, and tomato.
Hoplolaimus galeatus (H. coronatus). Crown-Headed Lance Nema-
tode, wide-spread. On turf grasses, zoysia, nursery crops, corn, sugarcane,
citrus, tomato, sweetpotato, pine seedlings, and carnation. This species may
feed from the outside, burying the head only, or it may enter the root com-
pletely, destroying the cortex, which is sloughed off, and feeding on the
phloem.
Hoplolaimus magnistylus. Stunt on hibiscus.
Hoplolaimus uniformis. On various ornamentals, reported from Rhode
Island.
Hypsoperine
Heteroderidae. Similar to Meloidogyne, the root-knot nematode, but female body oval
rather than pear-shaped.
Hypsoperine graminis. Described in 1964 from roots of grass and forming
inconspicuous galls, primarily on members of the Gramineae. St. Augus-
tine grass may become chlorotic and die. Bermuda grass may decline. Also
present on zoysia.
Longidorus and Paralongidorus
Longidoridae. Needle nematodes; relatively large ectoparasites with long, slender stylet;
similar to Xiphinema but not causing galls.
NEMATODES 399
Longidorus elongatus. On grape, causing necrosis and excessive root-
branching.
Longidorus maximus. Reported associated with celery, leek, lettuce, and
parsley.
Paralongidorus sylphus. Thorns Needle Nematode, fairly common in
the Pacic Northwest, causing severe stunting of peppermint.
Meloidodera
Heteroderidae. A new genus, a link between Heterodera and Meloidogyne; eggs are
retained in the female, but there is no distinct cyst stage; second stage larvae invade
roots but no galls are formed.
Meloidodera oridensis. In roots of slash pine in Florida.
Meloidoderita
Tylenchulidae sp. On grapes. Males developed in soil and have a degenerate
esophagus that lacks a stylet.
Meloidogyne
Heteroderidae. Root-knot nematodes, formerly considered one species, now known to
be several, distinguished by slight morphological differences such as striations, per-
ineal pattern of the tail, type of galls formed, host preferences, and somewhat by local-
ity. Females are white, pear-shaped to sphaeroid with elongated necks, slender stylets
with well-developed basal knobs; males are slender, wormlike. Females deposit eggs in
a gelatinous mass, and the body is not turned into a cyst as in Heterodera (see Fig. 3.32).
Root knot is the best known nematode disease, with over 2000 plant species
susceptible to one or more forms of Meloidogyne. Root knot was rst
reported in England, in 1865 on cucumbers; in 1876 it was recorded in the
United States on violet. Infected plants are stunted; they often wilt, turn
yellow, and die. The chief diagnostic symptom is the presence of small or
large swellings or galls in the roots (see Fig. 3.33). They are nearly round
or long and irregular, but they are an integral part of the root and cannot
be broken off. This differentiates them from benecial nodules, formed
Figure 3.32 Root-Knot Nematode Galls on Potato
on legume roots by nitrogen-xing bacteria, which can be readily broken
off.
Root-knot nematodes occur in practically every state. We used to think they
were killed by northern winters, but some species can survive extreme cold.
They do have fewer generations in the North and do not build up such large
populations as in southern sandy or peat soils. Grasses and grains are about
the only plants immune or resistant to root knot.
The long, thin young larva takes form inside the egg, breaks out, and migrates
through the soil to a root. It moves in to the axial cylinder and there becomes
sedentary. It injects a secretion of its esophageal glands into the tissue by
means of its short buccal stylet, and this stimulates the formation of 3 to 5
giant cells around the injection point. The nematode absorbs its food from
these nectarial cells the rest of its life. As it feeds, the larva swells rapidly
into a sausage-shaped body, which, in the female, becomes whitish and pear-
shaped, large enough to be just visible to the naked eye. The male changes
into a threadlike cylindrical form, folded up inside the larval molt, from
which it nally escapes.
The female deposits its eggs in an extruded yellow-brown jelly. There may
be up to 3000; the average is nearer 300 to 500. The larvae develop inside
the eggs and become free in the soil when the host root cracks or decays.
They may attack the same root in a new place or another root. At 80F a gen-
eration takes only 25 days; at 67F the cycle averages 87 days, and below
55F activity ceases. Root-knot nematodes may be injurious by their feeding
punctures even if typical swellings are not formed. Some have been shown
to increase Fusarium and bacterial wilts, and they almost surely complicate
the crown-gall problem.
NEMATODES 401
Figure 3.33 Root-Knot Nematode; pear-shaped female with egg sac; encysted young larva; and wormlike adult
male
Control. Rotation of crops may be practical only for species with a narrow
host range, and a few varieties of vegetables, fruits, and ornamentals have
been developed resistant to particular species. Soil fumigation before plant-
ing is a common control; see

Chap. 1 for satisfactory chemicals. These
usually kill larvae free in the soil but not all of those inside root knots.
Meloidogyne arenaria. Root-Knob Nematode. Causes stunting and root
gall on dwarf gardenia, compacta holly, and japanese boxwood. Also report-
ed on watermelon.
Meloidogyne arenaria (formerly Meloidogyne arenaria thamesi). Thames
Root-Knob Nematode. Occurring naturally in Florida, on Chinese silk-
plant (Boehmeria); found elsewhere in greenhouses. Also reported on tomato
and scindapsus.
Meloidogyne arenaria thamesi (see Meloidogyne arenaria). Thames
Root-Knob Nematode.
Meloidogyne chitwoodi. Columbia Root-Knob Nematode. Has been
associated with alfalfa, potato, tomato, sugar beet, wheat, and corn. This is
signicant because wheat and corn are commonly grown in rotation with
potato and sugar beets to reduce M. hapla. Both monocotyledonous and
dicotyledonous plant species are good hosts, indicating a wide host range
for this nematode.
Meloidogyne graminicola. Rice Root-Knob Nematode. On purple
nutsedge and yellow nutsedge. Root-Knob Nematode on clovers.
Meloidogyne hapla. Northern Root-Knob Nematode. Common on
many outdoor crops in the North and in orist and nursery stock. Hosts
include abelia, anoda, barberry, bean, blueberry, boxwood, California-laurel,
cantaloupe, carrot, cherry, clematis, clovers, cocklebur, corn, cress, cucum-
ber, dog fennel, eggplant, escarole, forsythia, geranium, germander, glad-
iolus, grape-hyacinth, goldenchain, jimsonweed, kale, lettuce, marigold,
mock-orange, morning-glory, mulberry, myrtle, mustard, parsnip, pachysan-
dra, pansy, peanut, peony, pepper, periwinkle, potato, privet, rose, sainfoin,
sequoia, soybean, spirea, spurge, strawberry, sugar beet, tomato, velvetleaf,
viburnum, wheat, and weigela.
This species is a particular pest of peanut and is probably the most impor-
tant nematode on strawberries. It causes galls, reduces growth of main roots,
resulting in excessive branch roots; plants are stunted and may die. Injury is
more serious in sandy soils. Yields have been increased by using granular
Nemagon, mixed with fertilizer, as a side-dressing or by planting in fumigat-
ed beds. Rotation with corn and some grains may be practical.
NEMATODES 403
Meloidogyne incognita. Root-Knob Nematode. On kiwi, sequoia, soci-
ety garlic, sweet potato, and jacquemontia.
Meloidogyne incognita. Cotton Root-Knob Nematode. A southern
native associated with many plants forage crops, bean, cabbage, can-
taloupe, carrot, celery, chard, corn, cucumber, grape, lettuce, pepper, potato,
radish, rhubarb, soybean, New Zealand spinach, squash, tobacco, tomato,
turnip, watermelon; also on azalea, boxwood, camellia, calthea, coleus,
collinsia, daylily, gardenia, hibiscus, hollyhock, iris, India love grass, neph-
thytis, roystonea, schefera, and scindapsus. It was reported on iris in 1955,
from New York and Texas, the rst instance of rhizomatous iris credited as
host to a root-knot nematode. Tips of leaves turn yellow, then brown, with
whole leaf gradually dying. There are some resistant soybean varieties, and
asparagus, strawberry, and peanut can be used in a rotation.
Meloidogyne incognita. Southern Root-Knob Nematode. Native to the
South and common there, but overwintering as far north as New Jersey. This
is the most important root-knot species on peach; it is also recorded on abelia,
banana, bean, carrot, coleus, corn, cucumber, daylily, eggplant, gardenia,
geranium, hibiscus, onion, okra, sweetpotato, pepper, tomato, watermelon,
and willow. It causes stunting and chlorosis of gardenia, but does not occur
on peanuts or strawberries, and these may be used in a rotation. Resistant
crotalaria and oats can be used as cover crops in peach orchards, and some
peach understocks are highly resistant.
Meloidogyne javanica. Javanese Root-Knob Nematode. Common in
southern peach orchards and nurseries, widespread in Georgia on peaches
such as Yunnan and Shali that are otherwise resistant to root knot. Found
in northern greenhouses. May be associated with azalea, bean, beet, cab-
bage, calendula, carrot, carnation, corn, Cocos plumosa, cucurbits, eggplant,
impatiens, radish, sequoia, snapdragon, soybean, tomato, and watermelon.
Resistant peanut, strawberry, cotton, and pepper can be used in the rotation.
Meloidogyne ovalis. On maple.
Naccobus
Pratylenchidae. Males wormlike; females swollen in the middle, saclike,
with a short, narrow tail; eggs extruded in a gelatinous matrix or held within
the body; stylet with small basal knobs; endoparasites.
Naccobus aberrans. False Root-Knob Nematode. Important in western
sugar-beet elds, also present on garden beets, cacti, carrot, crucifers, gaillar-
dia, lettuce, and salsify. Root galls are similar to those caused by Meloidog-
yne, and may be fairly large.
Naccobus batatiformis. On cabbage, cactus, carrot, and stock
Naccobus dorsalis. Reported on heronsbill (erodium), probably on other
hosts.
Nacobbodera
Heteroderidae
Nacobbodera chitwoodi. Reported on Douglas r, and spruce.
Nothanguina
Anguinidae
Nothanguina (orrina) phyllobia. Foliar Nematode. On nightshade.
Paratylenchus
Tylenchulidae. Pin nematodes, related to ring nematodes but thinner, primarily ectopar-
asites; minute; cuticle nely annulated; female with long stylet, body ventrally curved
but too short for a spiral.
Paratylenchus (Gracilacus) anceps. On California-laurel.
Paratylenchus dianthus. Carnation Pin Nematode. First reported on car-
nation in 1955 in Maryland, now well distributed through the Northeast.
Paratylenchus elachistus. On Boehmeria.
Paratylenchus (Gracilacus) epacris. California Sessile Nematode.
Associated with a decline of black-walnut trees in California.
Paratylenchus hamatus. Celery Pin Nematode, Fig Pin Nematode. On
azalea, bean, boxwood, celery, chrysanthemum, clover, corn, g, geranium,
gladiolus, turf grasses, hemlock, holly, horse-radish, iris, oak, onion, pars-
ley, peach, pieris, pine, mountain-pink, prune, rose, soybean, strawberry, and
tomato. This species is responsible for celery losses in New England, plants
being stunted and chlorotic, and with decline of g in California, symptoms
being chlorosis and leaf drop and undersized fruit. On mint it has caused
NEMATODES 405
one-third reduction in growth. The nematodes can be starved out of celery
elds by a 2-year rotation with lettuce and spinach.
Paratylenchus micoletzkyi. On marigold.
Paratylenchus macrophallus. On mint.
Paratylenchus projectus. Reported from Maryland in 1955 on pasture
grasses, also found on roots of alfalfa, bean, clover, corn, spruce, and soy-
bean. A serious decline of celery and parsley in New Jersey was attributed to
this species. Preplanting fumigation has produced a striking growth response.
Paratylenchus penetrans. On snapdragon.
Pratylenchus
Pratylenchidae. Lesion nematodes, sometimes called root-lesion or meadow nematodes,
widely distributed migratory endoparasites; males and females wormlike, small, 0.3 to
0.9 mm, with short stylet. Conspicuous necrotic spots are formed on roots, and eggs
are deposited in root tissues or in soil. Feeding punctures afford entrance to pathogenic
microorganisms.
Pratylenchus brachyurus (P. leiocephalus). Godfreys Meadow Nema-
tode, Smooth-Headed Meadow Nematode. On corn, grasses, cereals,
asparagus, avocado, citrus, collinsia, dogwood, peanut, pieris, pine, pineap-
ple, potato, soybean, strawberry, and tomato. Unsightly lesions are formed
on peanut shells, and the nematode survives through curing. Preplanting soil
fumigation has increased yield.
Pratylenchus coffeae (P. musicola). Associated with strawberry black root
and decline, in Arkansas.
Pratylenchus crenatus. Associated with many kinds of nursery plants.
Pratylenchus fallax. Lesion nematode on grass.
Pratylenchus hexincisus. Described from corn roots, Maryland.
Pratylenchus minyus. On pear and grape, in California. Also, on laree.
Pratylenchus musicola. On banana, g, olive, and walnut.
Pratylenchus nannus. Lesions nematode; On zinnia.
Pratylenchus negelectus. Lesion nematode on potato.
Pratylenchus penetrans. Cobbs Meadow Nematode. Associated with
decline in alfalfa, amaranth, apple, arborvitae, azalea, bean, blackberry,
blueberry, boxelder, cabbage, carrot, cedar, celeriac, celery, cherry, chrysan-
themum, clover, corn, cucumber, eggplant, fern, garden balsam, gayfeather,
gladiolus, grass, hemlock, holly, horseradish, lettuce, lily, maple, mock-
orange, onion, parsnip, peach, pear, peony, pepper, pieris, pine, mountain-
pink, phlox, plum, peach, pear, potato, raspberry, rose, safower, sequoia,
soybean, spinach, spirea, strawberry, sweetpotato, tobacco, tomato, turnip,
zinnia, and yarrow. This species is distributed throughout the United States.
Apples have necrotic black or amber spots on white rootlets; roots may be
stunted and distorted; tree vigor is reduced; leaves are small. The disease has
been called little leaf and rosette. The nematodes invade cortex only;
secondary fungi may play a part in symptoms. Control measures include root
dips and soil fumigation, hot-water treatment for strawberry stock plants, and
removal of all old roots on lilies before forcing. Marigolds produce a chem-
ical toxic to nematodes and can be used in rotations.
Pratylenchus pratensis. DeMans Meadow Nematode. Important on
grasses, strawberry, lily, and narcissus; reported on a great many other hosts,
but there may have been some confusion with other species.
Pratylenchus safaenis. On soybean, corn, cotton, millet, rice, and sorghum.
Pratylenchus scribneri. Scribners Meadow Nematode. First reported
on potatoes in 1889 in Tennessee. Associated with amaryllis, hibiscus, straw-
berry in Florida, roses in California, and in New Jersey, clover, corn, dahlia,
orchids, parsnip, peach, potato, raspberry, rose, soybean, and tomato.
Pratylenchus subpenetrans. Described from pasture grasses, Maryland.
Pratylenchus thornei. Thorns Meadow Nematode. On wheat, other
grains, and grasses. It has also been reported on maple and nectarine.
Pratylenchus vulnus. Walnut Meadow Nematode. Described in 1951
from California as an important parasite of walnut and rose on the West
Coast, also present elsewhere. It may affect avocado, boxwood, almond,
g, forsythia, gayfeather, apricot, citrus, peach, plum, raspberry, loganber-
ry, rose, sequoia, strawberry, Japanese boxwood, spiny Greek juniper, blue
rug juniper, walnut, and yew. Soil fumigation has increased growth of roses
by 400%.
Pratylenchus zeae. Corn MeadowNematode. Associated with corn, also
alfalfa, bean, chrysanthemum, cucumber, grasses, pea, phlox,.potato, soy-
bean, tobacco, and tomato.
Pratylenchus spp. Lesion Nematodes. Probably as widespread as a group
as root-knot nematodes and even more serious, though less readily recog-
nized. The brown or black root condition usually comes from secondary fun-
gi entering and rotting the roots after cells are pierced and torn by the nema-
todes. In boxwood and other ornamentals there is often a brush or witches
broom of new surface roots to compensate for old roots sloughed off. First
NEMATODES 407
symptoms are usually yellow, black, or brown lesions on ne feeder roots.
Boxwood becomes sickly, stunted; foliage is dark brown to orange, some-
times drops; some branches may be killed. Tuberous begonias may be heav-
ily infested in roots and tubers, with poor growth. Where possible, fumigate
soil before planting. Help plants to recover from root injury by mulching,
adequate watering, and feeding.
Radopholus
Pratylenchidae. Burrowing nematodes; endoparasites with entire life cycle inside plants,
including copulation and egg deposition. Male and female wormlike, with short stylet.
Female with at lip region, two ovaries; 0.6 mm long; male with rounded lip region.
Radopholus similis. Burrowing Nematode, Associated with SPREAD-
ING DECLINE OF CITRUS. The most important citrus disease in Flori-
da. This is a subtropical species, rst reported in 1893 from banana roots
in the Fiji Islands. Citrus decline was known for many years before the
nematode connection was made in 1953. This species is also responsi-
ble for AVOCADO DECLINE and in 1963 was reported as infesting 237
plants in many families. Possible hosts include acanthus, allamanda, alu-
minum plant, calathea, Barbados cherry, banana, castor-bean, cocculus,
hibiscus, Japanese boxwood, Japanese persimmon, ixora, jacobinia, ginger-
lily, loquat, Momordica, pandanus, peperomia, philodendron, periwinkle,
pothos, podocarpus, palms, guava, as well as corn, pepper, tomato, and other
vegetables, and various trees. Asparagus, marigold, and crotalaria are among
the few nonhosts. The burrowing nematode has been found in Louisiana as
well as central Florida.
The nematodes enter the cortical parenchyma of young succulent roots just
back of the tip and form burrows, leaving behind avenues of infection for
soil fungi and bacteria. Infected trees seldom die outright, but have poor
growth and cease to produce a protable crop. The disease spreads in all
directions from an infected specimen, but somewhat unevenly, the distance
ranging from 25 to 200 feet in a year, averaging about 50 feet. Long-distance
spread is by transplants from nurseries.
Control. Living trees, once infected, cannot be restored to vigor. Diseased
trees in quarantined areas are pulled and burned, including two trees beyond
those known to be infested in an orchard, and the soil is treated with D-
D. Bare-rooted nursery stock can be treated with hot water, 10 minutes at
122F. After the pull and treat, nonhosts are grown for 2 years before citrus
is replanted. There is some hope of resistant varieties.
Rotylenchulus
Hyplolaimidae. Reniform nematodes, partially endoparasitic root parasites. Female
swollen, kidney-shaped; two ovaries; male wormlike, unable to feed.
Rotylenchulus reniformis. Reniform Nematode. First described from
pineapple roots in Hawaii, now found in Florida and other warm states on
turf, cotton, peanut, sweetpotato, tomato, gardenia, jacquemontia, and other
ornamentals. The head of the female, with elongated neck, goes in the corti-
cal parenchyma of the rootlet, and her kidney-shaped body projects outside.
It is covered with a gelatinous material containing eggs and larvae, so that
soil particles adhere.
Rotylenchus
Hoplolaimidae. Spiral nematodes, worldwide in temperate and tropical climates; mostly
ectoparasitic but partially endoparasitic, somewhat migratory; body wormlike but held
in shape of a spiral; long stylet; female with two ovaries; 0.5 to 1 mm long.
Rotylenchus blaberus. Spiral Nematode on spider-lily.
Rotylenchus buxophilus. Boxwood Spiral Nematode. Associated with
boxwood decline in Maryland and nearby states; also found with barberry,
privet, and peony. The roots have minute brown spots, and the root system is
much reduced.
Rotylenchus cristiei. On grasses.
Rotylenchus robustus. Reported on azalea.
Rotylenchus uniformis. Reported on many ornamental trees and shrubs in
New Jersey nurseries.
Scutellonema
Hoplolaimidae. Spiral nematodes, similar to Rotylenchus.
Scutellonema blaberum (Rotylenchus blaberus). West African Spiral
Nematode. On banana, yam, red spider lily, and African-violet.
NEMATODES 409
Scutellonema brachyurum. Carolina Spiral Nematode. Working at
crown and roots of African-violet, destroying root cells, depositing eggs in
cortical tissues. Also on amaryllis.
Scutellonema bradys. Yam Nematode.
Scutellonema christiei. Christies Spiral Nematode. Common on lawn
grasses in Florida, also reported on apple and grasses in Maryland and West
Virginia.
Sphaeronema
Tylenchulidae
Sphaeronema sp. sp. Decline of Alaska-cedar.
Tetylenchus
Belonolaimidae. Male and female wormlike, stylet short.
Merlinius joctus. On blueberry.
Trichodorus and Paratrichodorus
Trichodoridae. Stubby-root nematodes; migratory ectoparasites with wide host ranges;
thick-bodied, cylindrical; 0.5 to 1.5 mm long; smooth cuticle; tail short, bluntly rounded;
long, slender stylet is a grooved tooth.
Paratrichodorus allius. Reported reducing onion yield in Oregon.
Paratrichodorus christiei. Christies Stubby Root Nematode. Wide-
spread in southern states but also present elsewhere feeding on many plants
in many different plant families. These include azalea, avocado, blueberry,
bean, beet, cabbage, citrus, corn, cranberry, chayote, onion, potato, squash,
strawberry, tomato, and turf grasses St. Augustine, Bermuda, and zoysia.
On tomato there is general stunting and formation of short lateral roots. The
stubby effect is apparently caused by a secretion and not just mechanical
piercing by the stylet; there is reduced cell multiplication. The host list is too
long for crop rotation to be practical, and soil fumigation is not as effective as
with some other species. Asparagus and poinsettia are nonhosts, and aspara-
gus has a nematicidal effect. This nematode and some other Trichodorus
species are vectors of tobacco rattle virus, cause of potato corky ringspot.
Trichodorus obtusus. Cobbs Stubby Root Nematode. On Bermuda
grass.
Paratrichodorus pachydermus. Seinhorst Stubby Root Nematode.
On turf and dahlia.
Trichodorus primitivus. On azalea.
Tylenchorhynchus
Belonolaidae. Stylet nematodes, sometimes called stunt nematodes, primarily ectopar-
asities, somewhat migratory, common in roots of nursery stock and cultivated plants.
Male and female wormlike, 0.6 to 1.7 mm long; stylet variable in length with well devel-
oped knobs; female has rounded tail, two ovaries; male tail is pointed; cuticle coarsely
annulated.
Tylenchorhynchus brevidus. On grasses.
Tylenchorhynchus capitatus. Causes stunting and chlorosis of pepper,
bean, tomato, and sweetpotato.
Tylenchorhynchus claytoni. Tesselate Stylet Nematode. Common
and widespread through southeastern and eastern states. Associated with
andromeda, apple, arborvitae, azalea, bean, blueberry, boxwood, broccoli,
cherry, cereals, clovers, corn, cranberry, dogwood, forsythia, grape, grasses,
hemlock, holly, lettuce, lilac, maple, peach, peanut, pepper, pine, potato,
raspberry, rhododendron, soybean, strawberry, sweetpotato, tomato, tulip-
tree, veronica, willow, and yew. Azaleas may be severely injured, with
reduced root system, short twigs, leaf chlorosis and increased susceptibil-
ity to winter injury. Soil treatment with the standard fumigants and also
with systemics gives adequate control. Nonhosts include peanut, pepper,
cucumber, and crotalaria.
Tylenchorhynchus dubius. Reported on cereals, grasses, clovers, also aza-
lea and carnation.
Tylenchorhynchus annulatus (formerly Tylenchorhynchus martini). Su-
garcane Stylet Nematode. On sugarcane, rice, soybean, and sweetpotato.
Tylenchorhynchus martini (see Tylenchorhynchus annulatus). Sugarcane
Stylet Nematode. On sugarcane, rice, soybean, and sweetpotato.
Tylenchorhynchus maximus. On turf.
NEMATODES 411
Tylenchulus
Tylenchulidae. Female sedentary, with elongated anterior portion entering the root and
swollen, ask-shaped posterior outside the root; well-developed stylet with large basal
knobs; male remains small, cylindrical; does not feed.
Tylenchulus semipenetrans. Citrus Nematode. First noted in California
in 1912, now widespread in citrus regions; important in California and Flori-
da, present also in Arizona and Texas. Hosts other than citrus include olive,
persimmon, grape, and lilac. Citrus trees exhibit a slow decline resulting
from reduced root activity. Symptoms also include twig dieback, chlorosis
and dying of foliage, wilting under moisture stress, and reduced fruit produc-
tion. Control measures include resistant rootstock, and hot-water treatment
of nursery stock, 25 minutes at 113F or 10 minutes at 116F.
Tylenchus
Tylenchidae. This genus, described in 1865, originally contained most species with
stomato-stylets, but many of these have been transferred to other genera. Those left are
common in soil around plants but apparently not important parasites.
Xiphinema
Longidoridae. Dagger nematodes; very common migratory ectoparasites; very long,
males and females both wormlike; long, slender stylet from a bottle-shaped esophagus.
Xiphinema americanum. American Dagger Nematode. A native, rst
described in 1913 from specimens taken around roots of corn, grasses, and
citrus trees. Found all over the United States associated with many kinds of
plants, including ash, azalea, bean, boxwood, clover, camellia, citrus, dog-
wood, elm, geranium, melon, oak, palm, pea, pecan, peach, pepper, pine,
poplar, rose, soybean, strawberry, sweetpotato, tomato, viburnum, vinca, and
walnut. In addition to its causing decline and sometimes winterkill by its
feeding on roots, this species is believed to transmit tomato ringspot, peach
yellow bud mosaic, and grape yellow vein viruses and to increase the inci-
dence of Cytospora canker on spruce. Dagger nematodes may be introduced
into greenhouses with virgin soil from the woods and may destroy almost all
the feeder roots of plants. There may be very high soil populations.
Xiphinema bakeri. Dagger Nematode. On sequoia.
Xiphinema chambersi. Chambers Dagger Nematode. Causing a de-
cline in strawberries, with stunting and sunken, reddish brown root lesions.
Xiphinema diversicaudatum. European Dagger Nematode. A proven
pathogen of rose, strawberry, peanut, g, tomato, soybean, garden balsam,
and other plants. This species is very common in commercial rose green-
houses, reducing vigor, causing chlorosis. Galls are formed on rose roots;
they are similar to root-knot galls but more elongate and nearer the tip of the
root, causing it to curl. Cleaning up a greenhouse infestation means disposal
of all plants in a bed, careful sterilization of soil, and replanting with clean
stock.
Xiphinema index. California Dagger Nematode. Reported on Boston
ivy, grape, g monkshood vine, pistachio, and rose. Feeding in root tips caus-
es a terminal swelling with angling of main roots, death of lateral roots.
Xiphinema radicicola. Pacic Dagger Nematode. Reported on oak, in
Florida.
NONPARASITIC DISEASES
Plants in poor health from one or more environmental conditions far out-
number those aficted with diseases caused by parasites bacteria, fungi,
and nematodes. When foliage turns yellow from lack of nitrogen, or from
unavailability of iron in an alkaline soil, or from lack of oxygen in a water-
logged soil, we call it a physiological or physiogenic or nonparasitic disease.
The adverse condition may be continuing, as it is with a nutrient decien-
cy, or it may be transitory, an ice storm, perhaps, lasting but a day but with
resultant dieback continuing for the next two years. It may be chemical injury
from injudicious spraying or fertilizing or from toxic substances in the atmo-
sphere. It may be due to a toxin injected by an insect.
Trees and crops can be insured against hail, hurricanes, lightning, and other
acts of God, but not the misguided zeal of gardeners. Years of working in
gardens in my own state and visiting gardens in other states from coast to
coast have convinced me that plants often suffer more from their owners than
from pests and diseases. Azaleas die from an overdose of aluminum sulfate
applied to correct acidity, when the original cause of ill health was a too-wet
soil. Rhododendron die when a deep, soggy mass of maple or other soft
leaves is kept around the trunks. Roses die when the beds are edged with
a spade and soil is mounded up in the center, burying some plants too deeply
and exposing roots of others. Seedlings die from an overdose of fertilizer in
hot weather. Trees die from grading operations.
Spray injury is exceedingly common, with the gardener thinking the red or
brown spots are fungus leaf spots and increasing the chemical dosage until all
foliage is lost. Weed killers take their unexpected toll of nearby ornamentals.
Either a deciency or an excess of plant nutrients can cause a physiologi-
cal disease. Greenhouse operators and commercial growers in the eld must
watch nutrition very carefully. The backyard farmer gets along pretty well
by using a complete fertilizer containing nitrogen, phosphorus, and potas-
sium in large amounts and minor elements in trace amounts. There are kits
available for amateur diagnosticians who wish to check soil deciencies and
acidity, but you may prefer to send a soil sample to your state experiment
station for a correct interpretation of nutrients and soil acidity. Take a slice
through the soil to spade or trowel depth from several places in the garden,
mix those samples together, and send a small sample of the mixture.
Acidity, Excess
Soil acidity or alkalinity is measured on a pH scale that runs from 0 to 14.
When the number of acid or hydrogen ions balances the number of alkaline
or hydroxyl ions, we have pH 7.0 or neutral. Above pH 7.0 the soil is alkaline
and may contain free lime; below it, the soil is acid. Few crop plants will
grow below pH 3.5 or above pH 9.0. If the soil becomes very acid, roots are
poorly developed and may decay, growth is slow, and foliage is mottled or
chlorotic. This result is due either to actual excess of hydrogen ions or to
physical structure of the soil and solubility of nutrients.
Most owering plants, fruits, and vegetables do well in a soil just slightly
acid, in a pH range of 6 to 7 or 6 to 8. Plants ourishing in a very acid soil, pH
4 to 5, are few: alpines, azalea, arbutus, andromeda, bunchberry, wild calla,
camellia, Chamaecyparis (white cedar), a few ferns, wild orchids, pitcher-
plants, galax, and mountain-ash. In the pH 5 to 6 list are: arbutus-tree, azalea,
bleeding-heart, birch, blueberry, bent grasses, bracken, camellia, Carolina
jessamine, Clarkia, cranberry, cypress, Daphne odora (but not D. mezerium,
which is in the 6 to 8 group), hemlock, juniper, mountain-laurel, some ferns,
some orchids, some oaks, pine, rhododendron, sour gum, spruce, silver-bell
tree, Styrax, strawberry, sweetpotato, and yew.
The small kits for home testing of soils include a booklet giving the pH
preferences of a long list of plants and the amount of lime required to correct
the acidity. This varies with the type of soil and the original pH. To bring
a sandy soil from pH 4 to above 6 takes only 1/2 pound of hydrated lime; it
takes 2 pounds of lime to effect the same change in a clay soil.
Air Pollution
Polluted air is not conned to cities. Even in the country crops suffer when
sunlight plus automobile exhaust produce ozone and other gases. Air pollu-
tants come from smelters, pulp mills, factories, power plants, incinerators,
and other sources. Ozone injury is common in pine, resulting in chlorotic
and needle mottling, tipburn, blight, needle ecking, and stunting; in tobac-
co, causing weather eck; in spinach, with oily areas followed by white
NONPARASITIC DISEASES 415
necrotic spots on upper leaf surface; in grape, with a dark stippling. Other
sensitive plants include bean, celery, corn, tomato, carnation, orchid, radish,
marigold, and petunias. Some varieties are more susceptible than others.
Smog occurs from a chemical reaction of unburned hydrocarbons, as from
automobiles, ozone, sunlight, and, usually, thermal inversion. Tremendous
losses in California orchid houses come when smog appears when plants are
in the budding stage.
Chrysanthemums may be prevented from owering by ethylene in the atmo-
sphere; tomatoes are also very sensitive. Injury from sulfur dioxide, a product
of fuel combustion, is at a high level in the colder months. Foliage has white
spots, tips, or margins. Soot particles entering houses from smokestacks
cause necrotic spots.
Control. For orchids and other high-priced greenhouse crops, air can be
passed through a lter of activated charcoal. Taller smokestacks reduce
injury from gases and soot. Increasing the vitamin C content of plants by
treating them with a substance such as potassium ascorbate may reduce
injury from ozone. Installation of purication devices in automobiles and
industrial plants may provide some future relief.
Alkali Injury
Some semiarid soils are nearly barren from excess of chemicals with a basic
reaction. Composition varies, but three common salts are sodium chloride,
sulfate of soda, and carbonate of soda; these salts become concentrated at the
soil surface with a whitish incrustation. Other soils are black alkali, where
the organic matter has been dissolved. Applications of gypsum or sulfur,
cultivation, and mulching are correctives.
Alkalinity
Either aluminum sulfate or sulfur, or both mixed together, can be used to
reduce the pH for plants doing best in a somewhat acid soil.
Aluminum Toxicity
Occasional, if aluminum is used in excess. Browning, dieback, sometimes
death of azaleas and other plants may occur.
Arsenical Injury
Leaves of peaches, apricots, and other stone fruits are readily spotted or
burned with lead arsenate unless lime or zinc sulfate is added as a correc-
tive. There may be similar leaf spotting and defoliation when these tender
fruits are grown in old apple land that has accumulated a residue of lead
arsenate over a period of years. Even apple trees can be severely injured by
arsenical sprays under some conditions.
Baldhead
In beans this is loss of the growing point, due to mechanical injury in thresh-
ing seed.
Bitter Pit
On apples this is called stippen or Baldwin spot and is characterized by
small, circular, slightly sunken spots on fruit, increasing in storage, especial-
ly at warm temperatures, most frequent on varieties Jonathan, Baldwin, Spy,
Rhode Island Greening. It seems to be related to uctuation of the moisture
supply in soil and increased by abundant rainfall shortly before harvest. On
pear, bitter pit is sometimes associated with moisture deciency; in olives,
with overnutrition.
Black End
In pear, the whole blossom end of the fruit may turn black and dry; the dis-
ease appears when oriental pear rootstocks are used in poor soil. In walnut,
black end of nuts is probably drought injury.
Black Heart
In beets, this is generally boron deciency (see below); occasionally it is
potassium or phosphorus deciency. In apple wood it may be freezing injury;
in potatoes, lack of oxygen; in celery, uctuating soil moisture.
Black Root
Defective soil drainage and accumulation of toxins are associated with black
roots, but so too are soil fungi and root nematodes.
Blasting
Blasting of inuorescence and failure to produce seeds. These symptoms
seem associated with extremes of soil moisture, too wet or too dry, at blos-
som time. Onion Blast, prevalent in the Connecticut Valley, appears within
a few hours after bright sunshine follows cloudy, wet weather. Leaf tips are
rst white, then brown.
Blindness
Blindness of tulips and other bulbs. Failure to ower may be due to Botry-
tis blight or other disease, but it may come from root failure in dry soil or
from heating of bulbs in storage or transit. Too early forcing may result in
blindness.
Blossom-End Rot
Very common on tomatoes, also on pepper, squash, watermelon. The tissues
at the blossom end of the fruit shrink, causing a dark, attened or sunken,
leathery spot, which may include nearly half the fruit (see Fig. 3.34). The
disease is most common on plants that have had an excess of rainfall in the
early part of the season, followed by a period of drought. There are, however,
various contributing factors, the most important being a deciency of calci-
um, which is needed for synthesis of rigid cell walls of the tomato. Adding
calcium oxide to the soil or spraying with 1% calcium chloride has reduced
the disease. For home gardens, deep soil preparation, use of a complete bal-
anced fertilizer, and mulching to conserve moisture should help.
Bordeaux Injury
Both the copper and the lime in bordeaux mixture can be injurious to some
crops. Cucurbits are stunted, and blossoming and fruit-setting are delayed in
Figure 3.34 Blossom-End Rot on Tomato
tomatoes. Red-spotting of foliage of roses and apples is followed by yellow-
ing and defoliation. See Copper Spray Injury; Lime-Induced Chlorosis.
Boron Deciency
A small quantity of boron is required for normal growth of most plants. For
some there is not much leeway between necessary and toxic amounts; other
plants require or tolerate large amounts. Deciency symptoms vary with the
crop.
Fruit trees. Internal and external cork of apples, dieback, rosette; dieback,
blossom blight of pear; stunting, excessive branches, internal necrosis of
peaches. Apple leaves on terminal shoots turn yellow, are convex with red
veins; twigs die back from tip; dwarfed, thickened, brittle leaves are in tufts
at nodes; internodes are abnormally shortened. Fruit has dry corky lesions
throughout the esh or diffuse brown lesions and bitter taste. McIntosh,
Baldwin, Rome, Northwestern Greening, and Jonathan exhibit external cork
with severe russeting of surface. Control by applying borax, 1 ounce per each
inch of diameter of tree trunk, in a 1-foot band outside the drip of the branch-
es. Apply only once in 3 years, and reduce the amount by half for peaches
and other stone fruits and for very sandy soils.
Beets, turnips, other root crops. Black Heart, Brown Heart. Roots have dark
spots; plants are gradually stunted and dwarfed; leaves are small, variegated,
twisted. The interior of the beet or turnip has a dark brown to nearly black
water-soaked area, sometimes with a hollow center. The amount of borax that
can be added without injury depends on type of soil and moisture content.
Celery. Cracked Stem. Leaves have a brownish mottling; stems are brittle,
cracked with brown stripes.
Lettuce. There is malformation of young leaves, death of growing point.
Ornamentals. Terminal ower bud dies; top leaves are thick and brittle.
Application of boron in fritted form has prevented splitting in carnations,
and has increased ower production in greenhouse roses.
Boron Toxicity
Retardation or prevention of germination, death or stunting of plants, bleach-
ing or yellowing of tops, disappearance of color along midrib and veins, all
are indications of excess boron. Beans are extremely sensitive to boron, with
injury from as little as 4 pounds borax broadcast per acre. If borax has been
used for root crops, boron-tolerant cabbage should follow before beans in
the rotation.
Brown Bark Spot
Brown Bark Spot of fruit trees. Perhaps this is arsenical injury from residue
in the soil.
Brown Heart
Brown Heart of turnip, cabbage, cauliower.

Boron Deciency.
Bud Drop
In sweet pea very young ower buds turn yellow and drop off when there is
a deciency of phosphorus and potassium during periods of low light inten-
sity. Water sparingly at such periods; avoid excess of nitrogen. Gardenias
often drop their buds when taken from greenhouses to dry homes, but there
is also bud drop in greenhouses with high soil moisture, high temperature,
and lack of sunlight in winter.
Calcium Chloride Injury
Trees may be damaged when this dust-laying chemical is washed off country
roads or driveways down to roots.
Calcium Deciency
All plants require calcium, which is built into walls of cells, neutralizes
harmful by-products, and maintains a balance with magnesium and potas-
sium. Calcium is leached out of the soil as calcium carbonate and should be
replaced by adding ground limestone, or dolomite (calcium magnesium car-
bonate), or gypsum (calcium sulfate), which does not increase the pH of the
soil.
In fruits, calcium deciency shows rst in the roots, which are short and stub-
by with a profuse growth behind the tips that have died back. Basal immature
peach leaves sometimes have reddish discolorations, and twigs may die back.
Corn and legumes require large amounts of calcium, which may become
unavailable under conditions of high soil acidity.
Catface
Fruit deformity, due to insects or growth disturbances.
Chlorine Injury
A tank of chlorine gas for the swimming pool carelessly opened too close to
trees and shrubs causes foliage browning and sometimes death. Leaf margins
are sometimes killed by chlorine gas from manufacturing processes.
Chlorosis
Yellowing or loss of normal green color may be due to deciency of nitro-
gen, magnesium, or manganese. Occasionally boron deciency or toxicity,
insufcient oxygen to the roots in a waterlogged soil, or alkali injury may
cause chlorosis but in the majority of cases, and particularly with broad-
leaved evergreens, it occurs because iron is unavailable in an alkaline soil.
Iron Deciency.
Chlorosis
Hydroponically grown basil with interveinal chlorosis associated with CO
2
enrichment.
Copper Deciency
Exanthema or dieback of fruits apple, apricot, citrus, olive, pear, prune;
failure of vegetables on muck soils. Copper deciency in fruits is widespread
in Florida and occurs frequently in California. Leaves are unusually large
and dark green, or very small and quickly shed, on twigs that die back, with
a reddish brown gummy discharge. Citrus fruits are bumpy and drop, or have
insipid avor and dry pulp. Application of copper sulfate to the soil corrects
the deciency, but often spraying trees once or twice in the spring with bor-
deaux mixture provides sufcient copper indirectly. Spraying almonds with
a copper chelate has prevented shriveling of kernels. Muck or peat soils in
New York, formerly unproductive, now grow normal crops of onions and
lettuce with the addition of copper sulfate. On copper-decient Florida soils,
many truck crops fail to grow or are stunted, bleached, and chlorotic.
Copper Spray Injury
Some xed copper sprays are less injurious than bordeaux mixture, but all
coppers may be harmful to some plants under some conditions. Foliage spots
are small, numerous, reddish, sometimes brown. In peach leaves the centers
of the spots may fall out, leaving shot holes. Rosaceous plants follow spotting
with yellowing and dropping of leaves. Even mild coppers may be injurious
if the temperature is below 55F, or the weather continues rainy or cloudy.
Treated leaves are often harsher than normal and more subject to frost injury.
Dwarng and stunting are important symptoms on many crops, especially
cucurbits. Tomato owering is injured or delayed; apple and tomato fruits are
russeted. Tree roots are injured by overow from pools treated with copper
for algae.
Cork
Boron deciency, in apple.
Cracked Stem
Boron deciency, in rhubarb, celery.
DDT Injury
Foliage of some plants cucurbits particularly, roses occasionally turns
yellow or orange, often with stunting. Certain camellia varieties have been
injured when shrubs are under trees sprayed with DDT. Continued spraying
with DDT builds up a residue in the soil which may eventually have a toxic
effect on the root system, the effect varying with the type of soil and plant.
Dieback
This is due to deciency or excess of moisture, nutrients; winter injury; also
cankers, nematodes, borers.
Drought
The effects of a prolonged dry period may be evident in trees and shrubs for
two or three years thereafter.
End Spot
End Spot of avocado. Unequal maturity in both ends of the fruit seems to
be a factor in withering, spotting, and cracking at lower end. Pick promptly,
instead of leaving on trees.
Exanthema
Copper deciency, in fruits.
Frost Injury
This injury is caused by low temperature after plants have started growth
in spring or before they are dormant in fall (see Winter Injury for freezing
Figure 3.35 Frost Injury on Holly
during the dormant period). Yellow color of some leaves in early spring is
due to temperatures unfavorable for chlorophyll formation. Some leaves,
including those of rose, are reddened or crinkled with frost (Fig. 3.35). Blos-
som buds of fruit trees are critically injured by frost late in spring. In the
South, where plants come out of dormancy early, orchard heaters, smudge
res, power fans, and airplanes ying low to stir up the air are all used to
help save the crop. Many ornamentals are injured when a long, warm autumn
ends in a sudden very cold snap, or warm weather in February or March is
followed by heavy frosts. Cracks in tree trunks come from such temperature
uctuations.
Gas Toxicity
Illuminating gas escaping from aging gas mains causes slow decline or
sudden death, depending on the plant. Tomatoes are extremely sensitive
and indicate the slightest trace of gas by leaves and stems bending sharply
downward. Plane trees develop rosy canker long, narrow cankers near
the trunk base with inner bark watermelon-pink and swollen. With large
amounts of gas escaping, foliage wilts and browns suddenly, followed by
death of twigs and branches; with slow leaks, the symptoms appear gradu-
ally over a year or two. After the leak is repaired, it is sometimes possible to
save trees by digging a trench to aerate the roots, applying large quantities
of water, burning out severely injured roots, then replacing soil and feeding
to stimulate new growth.
Natural gas is, apparently, not as injurious.
Girdling Roots
Unfavorable conditions sometimes deect roots from their normal course,
and one or two may grow so closely appressed to a tree as to almost strangle
it. If one side of a tree shows lighter green leaves with tendency to early defo-
liation, dig down on that side to see if a root is choking the trunk under the
soil surface. The root should be severed and removed, then all cut surfaces
painted.
Grading Injuries
Many shrubs die when they are planted much deeper than the level at which
they were grown in the nursery. Similarly, many trees die when they are
covered over with ll from house excavations. Roots require oxygen for sur-
vival, and a sudden excess of soil cuts off most of the supply. A tree expert
should be on hand to give advice before any digging starts. Afterward is too
late. And if grading means lling in soil around trees, a little well around the
trunk is not enough. There must be radial and circular trenches laid with tile,
and then crushed stone and gravel, before the top soil goes in place. Con-
sult Tree Maintenance by P. P. Pirone for clear descriptions and diagrams for
protecting trees from contractors.
Graft Incompatibility
Lilacs are sometimes blighted from incompatibility of the lilac scion on priv-
et stock. Walnut girdle is due to incompatibility of scions on black walnut
roots.
Gummosis
Formation of gum on bark of fruit trees is commonly formed in cases of
bacterial canker, brown rot, crown rot, and root rots from soil fungi and in
connection with the peach tree borer, but other cases of gummosis seem con-
nected with adverse sites and soil moisture conditions irrespective of para-
sitic organisms.
Heart Rot
Boron deciency, in root crops.
Heat Injury
There are many ways in which excessive high temperatures can injure
plants, ranging from death to retarded growth or failure to mature owers
and fruit. Sunstroke, outright killing of plants, is a limiting factor in ower
and vegetable production in summer in the South. Seedlings, especially tree
seedlings and beans, may have heat cankers with stem tissues killed at the
soil line. See also Sunscald, Leaf Scorch, Tipburn.
Hollow Heart
This is sometimes due to excessive soil moisture.
Hopperburn
Marginal chlorosis, burning and curling of leaves of potatoes and dahlias is
due to leafhoppers.
Internal Browning or Cork
Internal Browning or Cork of apple. Boron deciency.
Iron Deciency
Iron is seldom, or never, actually decient in the soil, but it is often in such
an insoluble form in neutral or alkaline soils that plants cannot absorb it, or
it may be precipitated as insoluble iron phosphate where excessive amounts
of phosphates are added to the soil. Chlorosis is an indication of the lack of
iron, for it is necessary for the formation of chlorophyll, the green pigment
(see Fig. 3.36). In acid soils iron is usually available; in alkaline soils leaves
turn yellowish green, often remaining green along the veins but yellowing in
Figure 3.36 Iron Deciency in Chrysanthemum
interveinal areas. Terminal growth of twigs is small, and the shrub or tree is
generally stunted.
To obtain a quick response it is possible to spray leaves with a solution of
ferrous sulfate. More lasting is a soil treatment of a 50-50 mixture of ferrous
sulfate and sulfur.
Rather recent is the use of chelated iron, sold as Sequestrene and under other
trade names. In this form the iron cannot be combined with soil elements and
remains available to the plant even under alkaline conditions. The solution,
prepared according to directions on the package, is poured on the soil around
the unthrifty bush, and often the green color returns in a matter of days. Iron
chelates are now extensively used for citrus and for ornamentals.
Leaf Scorch
Leaf Scorch, of maple, horse-chestnut, beech, walnut, and other trees. Scat-
tered areas in the leaf, between the veins or along the margins, turn light or
dark brown, with all the leaves on a branch affected more or less uniform-
ly. The canopy of the tree looks dry and scorched; leaves may dry and fall,
with new leaves formed in summer. Lack of fruiting bodies distinguishes
scorch from a fungus leaf blotch. It appears during periods of high tempera-
ture and drying winds and often after a rainy period has produced succulent
growth.
Leaf scorch of Easter lilies has been a problem for years but can be prevent-
ed by keeping the pH of soil near 7.0 with lime, adequate nitrogen, but low
phosphorus. It may have some connection with root rots.
Leaf scorch of iris has puzzled amateur growers in the past few years; it is
more serious in the Southwest but has appeared in gardens elsewhere. Leaves
turn bright reddish brown at the tips in spring before owering, and in a few
days the whole fan is scorched and withered, and the roots have rotted with
a reddish discoloration (see Fig. 3.37). Many theories, including nutrition
and nematodes, have been advanced, but there is no general agreement as to
cause.
Lightning Injury
Trees may be completely shattered or a narrow strip of bark and a shallow
layer of wood torn down the trunk. Tall trees or those growing in the open
Figure 3.37 Scorch in Iris
are most likely to be struck. Valuable trees can be protected with lightning
conductors, installed by a competent tree expert.
Lime-Induced Chlorosis
Plants are sickly, with yellow foliage, in calcareous soils or near cement
foundations.

Iron Deciency.
Little Leaf
Little Leaf, on almond, apricot, avocado and other fruits.

Zinc Deciency.
Magnesium Deciency
Large areas in the Atlantic and Gulf Coast truck crop regions are low in
magnesium because of natural lack of magnesium rock, extensive leaching
from heavy rainfall, removal of large quantities in crops, and use of fertiliz-
ers lacking this element. In tomatoes, veins remain dark green while rest of
leaf is yellow or chlorotic. Cabbages have lower leaves puckered, chlorotic,
mottled, turning white at the margin and in center. In strawberries, leaves are
thin, bright green, then with necrotic blotches. On fruit trees, fawn-colored
patches are formed on mature, large leaves, with affected leaves dropping
progressively toward the tip. In owering plants there are a greatly reduced
rate of growth, yellowing between veins of lower leaves, sometimes dead
areas between veins, sometimes puckering.
Control by using dolomitic limestone, or with fertilizers containing mag-
nesium, or with Epsom salts (magnesium sulfate) around azaleas and other
shrubs in home gardens.
Manganese Deciency
Top leaves become yellow between veins, but even smallest veins retain
green color, giving a netted appearance. Lower the pH below 7 and add man-
ganese sulfate to the soil.
Marginal Browning
Potassium deciency or hopperburn.
Mercury Toxicity
Roses are extremely sensitive to mercury vapor and have been gravely
injured when paints containing mercury were used to paint sash bars in
greenhouses. Covering the paint with a paste of dry lime sulfur mixed with
lime, our, and water reduced the amount of toxic vapor.
Molybdenum Toxicity
Cause of whiptail in broccoli and cabbage, chlorosis of citrus in Florida, of
grapes in Michigan. Citrus leaves have large interveinal yellow spots with
gum on undersurface and may fall. Injecting the trunk with sodium molyb-
denate has corrected the condition quickly. On grapes chlorosis of terminal
leaves was attributed to molybdenum deciency correlated with nitrogen
toxicity and was corrected by adding 0.01 ppm molybdic acid to nutrient
solutions.
Mottle Leaf
Zinc deciency.
Nitrogen Deciency
Symptoms are paleness or uniform yellowing of leaves, and stems, ring
or burning of lower leaves, sometimes red pigments along veins, stunted
growth, reduced yield with small fruit. Immediate results can be obtained by
side-dressing with a quickly available nitrogenous fertilizer, but long-range
planning includes use of legumes in the rotation, green-manure crops, and
balanced fertilizers. Urea is recommended for turf, one application providing
a slow release through the season.
Nitrogen Excess
Too much nitrogen leads to overdevelopment of vegetative growth at the
expense of owers and fruit; to bud drop of roses, sweet peas, and toma-
toes; and, in high concentrations, to stunting, chlorosis, and death. Excessive
nitrogen decreases resistance to winter injury and to such diseases as re
blight, powdery mildew, and apple scab.
Oedema
Small, wartlike, sometimes corky, excrescences are formed on underside of
leaves of many plants cabbage, tomatoes, geraniums, begonia, camellias,
etc. When roots take up more water than is given off by leaves, the pres-
sure built up may cause enlarged mesophyll cells to push outward through
the epidermis. This condition is rare outdoors but is found in greenhouses
and sometimes on house plants where they have been overwatered. Copper
sprays sometimes produce similar intumescences. Camellias frequently have
corky swellings on bottom surface of leaves, often due to water relations,
sometimes to a spot anthracnose fungus.
Oxygen Deciency, Asphyxiation
Overwatered house plants and crops in poorly drained low situations
often show the same symptoms as those caused by lack of water, for the
roots cannot respire properly and cannot take up enough water. Improve
drainage; lighten soil with compost and sand; avoid too much articial
watering.
Phosphorus Deciency
Young leaves are dark green; mature leaves are bronzed; old leaves are mot-
tled light and dark green. In some plants there is yellowing around leaf mar-
gins. Stems and leafstalks develop reddish or purplish pigments; plants are
stunted, with short internodes; growth is slow, with delayed maturity. Most
complete commercial fertilizers have adequate phosphorus, but it can be
added separately in the form of superphosphate. In preparing rose beds apply
a liberal amount at the second spade depth as well as in the upper soil.
Potassium Deciency
Marginal browning, bronzing, or scorching appears rst on lower leaves
and advances up the plant, which is stunted. Leaves are often crinkled, curl
inward, develop necrotic areas; the whole plant may look rusty. The lack of
potassium can be made up with a complete fertilizer containing 5 to 10%
potash. Wood ashes also help to supply potassium.
Ring Spot
Yellow rings on African-violet foliage come from breaking down of the
chloroplasts when the leaf temperature is suddenly lowered, as in watering
with water considerably colder than room temperature.
Rosette
Zinc deciency in pecan and walnut, boron deciency in apple.
Rust
This term is used by amateur gardeners for any rust discolorations for a leaf
blight of phlox of unknown origin (probably a water relation), a spot necrosis
of gladiolus, red-spider injury, and many other troubles that have nothing to
do with true fungus rusts.
Salt Injury
Trees and shrubs along the seacoast are injured by ocean spray, and after
hurricanes and high winds traces of injury can be found 35 to 40 miles
inland. Conifers are usually affected most; they appear damaged by re, with
needles bright yellow, or orange-red. Eastern white pine is very susceptible;
Austrian and Japanese black pines, blue spruce, and live oak are highly resis-
tant. Roses have often survived submersion in salt water during hurricanes.
Roadside trees, and e specially maples, may be injured by salt used on high-
ways during the winter. Either sodium chloride or calcium chloride may be
harmful.
Scald
Scald, of apple. Asphyxiation injury to fruit in storage from accumulation of
harmful gases; most important when immature fruit is stored without ade-
quate ventilation at too high temperature and humidity. Wrapping fruit in
oiled paper or packing with shredded oiled paper, and storage near 32F,
with a high concentration of carbon dioxide at the start, control scald.
Scorch
Leaf Scorch.
Shot Berry
Shot Berry, of grape. Defective pollination.
Smog Injury
Unsaturated hydrocarbons and ozone in the atmosphere are the cause, with
many kinds of plants injured in the Los Angeles area. Tan lesions appear
on fern leaves in 24 hours with necrosis in 24 more (Fig. 3.38). Many orna-
mentals and vegetables are injured, with annual loss $3 million. Spraying
carnations in greenhouses with Vitamin C prevents sleepiness from smog.
Some greenhouses have installed activated-carbon lters for polluted air.
Figure 3.38 Ozone Injury on Tobacco
Smoke Injury
The most important agent in smoke injury is sulfur dioxide, a colorless gas
with a suffocating odor released from smelters and many industrial pro-
cesses. Acute smoke injury shows in rapid discoloration of foliage, defo-
liation, sometimes death. Conifer needles turn wine red, in whole or part,
then brown. Leaves of deciduous trees have yellow to dark brown dead
areas between veins, with tissue next to larger veins remaining green. Chron-
ic injury results in unhealthy, stunted trees, but less apparent discoloration
and defoliation. Roses, grapes, and legumes are seriously injured. Gladiolus
leaves appear burned from the tips down.
Control of injurious smoke must be at the source by lters, tall smoke-
stacks, neutralizing the acid gases, or using them in the manufacture of sulfur
and sulfuric acid.
Soot Injury
City trees and shrubs acquire an accumulation of soot, the solid residue of
smoke, which screens out the sunlight. Evergreens can be sprayed with a
soapy solution of Calgon (sodium hexametaphosphate), followed by syring-
ing with clear water.
Stigmonose
Dimpling of fruit by insect punctures.
Sulfur Injury
Sulfur sprays and dusts are likely to burn foliage in hot weather, when tem-
perature is much over 85F. There is often a browning of tip or margin of
leaves. Lime sulfur is injurious to some plants in any weather, russeting
peach foliage, causing apple drop, etc. When roses or other plants are con-
tinuously dusted with sulfur over a period of years, the soil may become too
acid and require lime as a corrective.
Sunscald
Trees with smooth bark are subject to sunscald when trunks or branches are
suddenly exposed to the sun, as when the next tree is removed. Young trees
are subject to sunscald the rst year or two after planting and should have
trunks wrapped in burlap or sprayed with a protective wax to prevent the
cambium under the thin bark from drying out.
Boxwood foliage is subject to sunscald in spring after winter covering is
removed, particularly if this is done on a sunny day with drying winds.
Sunscald is common on green tomatoes when fruits are exposed to sun in hot
dry weather (Fig. 3.39). This happens when foliage is lost through disease
or excessive irrigation, or when too much is removed in training tomatoes
to a single stem. A yellow or white patch appears on the side of the tomato
nearest the sun, often developing into a blister, then into a large, attened
spot with a papery white surface darkened by the growth of secondary fungi
and internal decay.
Figure 3.39 Tomato Sunscald
Sunstroke
Outright killing in excessive heat.
Tipburn
Potassium deciency may produce a tipburn, but more often this is a reac-
tion to heat, common in potatoes and particularly in lettuce, which shows
marginal browning of leaves and small brown or black spots in tissues near
larger veins. A regular supply of moisture and avoidance of excessive fertil-
ization in warm weather reduce tipburn, but more reliance should be placed
on growing varieties resistant to summer heat.
Topple
Topple, of gladiolus. Toppling over is apparently due to calcium deciency;
reduced by a spray of 2% calcium nitrate.
Variegation
Chlorophyll deciency, genetic factors, and virus diseases can produce var-
iegated plants.
Water Deciency
Practically all of the injury laid to excessive heat or cold is basically due to
lack of water. Winter winds and summer sun evaporate it from cells faster
than it can be replaced from roots, so that the cells collapse and die.
Weed-Killer Injury
There has always been some unintentional injury to neighboring plants in
the use of weed killers of the kill-all variety on driveways; but since we
have had 2,4-D as a selective weed killer for lawns, the damage to innocent
bystanders has been enormous, not only from spray drift and volatile material
in the atmosphere but from using for other spraying purposes equipment that
Figure 3.40 Weed-Killer Injury; Tomato and Oak
has applied 2,4D. It is impossible adequately to clean out such a sprayer;
mark it with red paint and keep it for weeds only. Symptoms of injury are
curling, twisting, and other distortions; there is often a fern-leaf effect instead
of normal-size foliage (Fig. 3.40). I have seen roses seriously malformed
when a factory several hundred feet away mixed up some 2,4-D. I have seen
tall oaks with all leaves unrecognizable after powdered 2,4-D was applied
to the lawn. I have seen chrysanthemum in a greenhouse utterly deformed
when 2,4-D was used on a lawn outside. Fortunately, unless the dose is too
heavy, the plants gradually grow back to normal.
Winter Injury
Most winter browning of evergreens is due to rapid evaporation of water in
sudden warm or windy spells. Copious watering late in the fall, a mulch, and
windbreaks are helpful for broad leaf evergreens, as is spraying them with
a waxy material, Wilt-Pruf, which prevents evaporation.
Sudden icestorms cause obvious breaking in trees; in boxwood and similar
shrubs they result in bark sloughing off and gradual dieback for months,
even years afterwards. I have seen symptoms on azaleas long after the ice
was forgotten.
Yellows
This term is used for some deciency disease but also for various virus dis-
eases and Fusarium wilts.
Zinc Deciency
Little Leaf of almond, apricot, apples, grape, peach, plum. Foliage is small,
narrow, more or less crinkled, chlorotic at tips of new growth, with short
internodes producing rosettes of leaves. Defoliation progresses from base to
top of twigs. The method of supplying zinc depends somewhat on the fruit.
Spray apples, peaches, plums, pears during dormant period with zinc sulfate.
Swab grape vines immediately after winter pruning.
Mottle Leaf of citrus. Leaves are small, pointed, with a sharply contrasting
pattern of green along midrib and main laterals and light green or yellow
between veins.
Rosette of pecans and walnuts. Narrow, crinkled leaets with dead or perfo-
rated areas have a rosette appearance; trees often bear no nuts. Pecan growers
in southeastern states broadcast zinc sulfate on soil under each tree in winter.
Variety Money-maker is resistant to zinc deciency.
Vegetable crops corn, beans, tomato, soybean have been protected by
amending the soil with 23 pounds zinc sulfate per acre.
POWDERY MILDEWS
Mildew is a disease in which the pathogen is seen as a growth on the surface
of plants. The same word is used for the fungus causing the disease. Mildews
are Ascomycetes. Black mildews are parasites in the order Meliolales with
a dark mycelium to give a sooty effect. They are common in the South or
on tropical plants in greenhouses (
Black Mildew). Powdery mildews are

plant parasites in the order Erysiphales. They have white mycelium, in a del-
icate weft or thick felt, made up of a criss-cross tangle of hyphae. Colorless
spores borne in chains on upright conidophores give the white powdery effect
(see Fig. 3.41). False or downy mildews are oomycetes, and the conspicuous
growth is not vegetative mycelium but fruiting structures and conidia pro-
truding through stomata or epidermis to give a white frosty appearance in
moist weather (
Downy Mildews).
True powdery mildews and in speaking of them we usually eliminate the
word powdery are widely distributed but sometimes more abundant in
semiarid regions than in areas of high rainfall, where other diseases ourish.
Unlike those of most other fungi, powdery mildew spores do not require free
water for germination. Some species require high humidity, but it is usually
provided at the leaf surface when cold nights change to warm days or when
plants are grown in crowded, low, or shady locations without sufcient air
circulation. Spores of other species can germinate with very low humidity.
When a mildew spore lands on a leaf and puts out its germ tube, it does
not make its nearest way inside the leaf but produces a tangle of septate
threads, hyphae, on the surface. Special sucking organs, haustoria, pene-
trate the epidermal cells, occasionally the subepidermal cells, in search of
food. The penetrating tube is slender, but, once inside the cells, the haustori-
um becomes a round or pear-shaped enlargement or a branched affair, with
greatly increased absorbing surface.
Condiophores, growing at right angles from the mycelium, produce one-
celled conidia in rows or chains of somewhat barrel-shaped hyaline cells,
which become oval as they are dislodged from the top of the chain and dis-
Figure 3.41 Powdery Mildew: mycelium and conidia formed on surface of a leaf and two types of the haustoria
in host cells
seminated by wind. Mildews known only in this anamorph state are called by
the form genus name Oidium. It requires the sexual fruiting bodies, perithe-
cia, to place mildews in their proper genera.
Perithecia are round with a dark membranous wall, technically cleistothecia
because they have no beak or ostiole, and rupture irregularly to free the asci.
They are held in place in the mycelium by appendages. The form of these
appendages and the number of asci in the perithecium are the chief charac-
ters differentiating the six genera important in this country (see Fig. 3.42).
Sphaerotheca and Erysiphe both have simple appendages; but the former
has only one ascus, the latter several. Podosphaera has appendage tips
dichotomously branched and one ascus; Microsphaera has the same type of
appendage but several asci. Phyllactinia has lancelike appendages swollen
at the base; those of Uncinula are coiled at the tip. Both have more than one
ascus.
Powdery mildews are obligate parasites, having no saprophytic growth peri-
ods in dead plant parts, although the perithecia carry the fungus through the
winter on either living or dead tissue. Mycelium sometimes winters in buds.
Symptoms of mildew are dwarng and stunting, often with a slight redden-
ing and curling of leaves before the white mycelium is noticeable. There may
be deformation of ower buds. Such symptoms are due to the withdrawal of
plant foods by the fungus and to excessive respiration.
Sulfur dust and lime sulfur sprays have long been considered specic reme-
dies for powdery mildews; some copper sprays are effective. Many of the
newer organics are ineffective. However, inorganics, in a formulated form,
i. e. potassium biocarboante are very effective and safe.
POWDERY MILDEWS 441
Figure 3.42 Powdery Mildews. Perithecia (cleistothecia) of the six genera: Erysiphe, simple appendages and
several asci; Sphaerotheca, same with one ascus; Mycrosphaera, dichotomously branched appendages and sev-
eral asci; Podosphaera, same with one ascus; Phyllactinia, appendages bulbous at base; Uncinula, appendages
coiled at tip
Erysiphe
Cleistothecia globose, or globose-depressed, sometimes concave; asci several, two- to
eight-spored; appendages occose (cottony), simple or irregularly branched; sometimes
obsolete, usually similar to mycelium and interwoven with it; mycelium brown in rare
cases.
Blumeria graminis (see Erysiphe graminis). Powdery Mildew of cereals
and grasses, economically important on bromegrass, wheat, oats, barley, and
rye; aesthetically important on lawn grasses, wheatgrass, fescue, and blue-
grass.
Brasiliomyces trina (formerly Erysiphe trina). Oak Powdery Mildew,
on tanbark oak and coast live oak, in California, causing witches brooms
(but

Sphaerotheca lanestris for the common live-oak mildew). Perithecia
are small, yellow-brown, with appendages lacking or rudimentary; asci have
two, rarely three, spores.
Erysiphe aggregata. Alder Powdery Mildew. Perithecia large, asci with
eight spores, rarely six or seven.
Golovinomyces cichoracearum (formerly Erysiphe cichoracearum). Pow-
dery Mildew of cucurbits and many ornamentals, mostly composites, per-
haps best known to gardeners as the Phlox Mildew. Asci are two-spored,
perithecia rather small, haustoria not lobed. There are nearly 300 hosts
including: cucumber, squash, pumpkin, gourds, cantaloupe, watermelon, let-
tuce, endive, Jerusalem artichoke, pepper, potato, salsify, Echinacea, Euca-
lyptus, Achillea, Anchusa, Artemisia, aster, begonia, Boltonia, calendula,
campanula, chrysanthemum, clematis, coreopsis, cosmos, dahlia delphinium
Eupatorium, gaillardia golden-glow, goldenrod, Helenium, hollyhock, Inula,
mallow Mertensia phlox, rudbeckia, Salpiglossis, salvia (sage), sunower,
stokesia and zinnia.
There are various strains of the fungus, the form on cucurbits not affecting
ornamentals, the strain of phlox (see Fig. 3.43) is reportedly limited to that
host, the strain on zinnia with a wide range of host plants. The lettuce strain,
perhaps a mutation of the form on wild lettuce, was not reported on cultivated
lettuce before 1951 and is important only in California and Arizona.
Powdery mildew was reported on cucurbits in North America in 1890, but
did not gain much prominence until 1926, when it suddenly reduced the
melon crop in the Imperial Valley of California by 5000 carloads. By 1939
mildew-resistant Cantaloupe 45 had been developed to meet the situation,
but in another decade the fungus had produced a different strain to which
Cantaloupe 45 was susceptible. Plant breeders can never rest on their laurels
because fungi that are obligate parasites seldom stay long outwitted. Other
varieties, Cantaloupes 5, 6, and 7, were bred resistant to both strains of the
fungus.
Powdery mildew is the principal disease of cucumbers in greenhouse culture,
with tiny white supercial spots on leaves and stems enlarging and becoming
powdery. Young watermelon fruits in greenhouses have small pimples or
warts under the area covered by mildew mycelium.
Phlox mildew is only too familiar to gardeners. The white coating often
appears on variety Miss Lingard in June, but on other varieties (in New Jer-
sey) more prominently in July and August. The mycelium is present on both
leaf surfaces and forms a thick felt on stems. In late summer black perithecia
are formed in great abundance. Powdery mildew on zinnias and chrysanthe-
mums usually starts so late in the season that it is more conspicuous than
harmful.
POWDERY MILDEWS 443
Figure 3.43 Erysiphe cichoracearum on Phlox
Control. Sulfur dust gives excellent control but is phytotoxic to some melons
and other cucurbits. Sulfur-tolerant cantaloupes have been produced. The
variety Homegarden is supposedly resistant to mildew and other pests. Keep
phlox and other ornamentals well spaced, and dust with sulfur at the rst
sign of white growth. Because the perithecia winter on old stems and leaves,
plants should be cut at ground level in autumn and burned.
Erysiphe cruciferarum. Powdery Mildew on cabbage and turnip.
Erysiphe graminis (syn. Blumeria graminis). Powdery Mildew of cere-
als and grasses, economically important on bromegrass, wheat, oats, bar-
ley, and rye; aesthetically important on lawn grasses, wheatgrass, fescue,
and bluegrass. The foliage is yellow or chlorotic with a white weft or mealy
coating on upperside of leaf, which later turns yellow and is often studded
with tiny brown perithecia. Seed from infected plants is small and shriveled.
Apply sulfur dust or wettable sulfur sprays.
Erysiphe heraclei. Powdery Mildew on carrot, parsley, chervil (culinary
herb), and celery.
Erysiphe orontii. Powdery Mildew on mint.
Erysiphe panax. Powdery Mildew on ginseng.
Erysiphe polygoni. Powdery Mildew of legumes, and many other veg-
etables and ornamentals, about 200 species in 90 plant genera. Appendages
are long or short, interwoven with the mycelium, but the perithecia are not
immersed in it. Asci have three to six spores. Peas exhibit a white powdery
coating over leaves and pods, with the latter often discolored. Leaves are
sometimes yellowish and deformed. The disease may be severe on peas in
arid sections of western states, particularly on late homegarden varieties.
On beans, the mildew is grayish. It is prevalent in California in cloudy weath-
er or in autumn when humidity is increased, but it is more important in the
Southeast. Other vegetables infected by this species include lima bean, soy-
bean, cabbage, turnip, radish, horse-radish, and carrot. Spores germinate at
quite low humidity.
The legume mildew is widespread on lupine, occasional on sweet pea. Other
ornamental hosts include acacia, anemone, arrowleaf clover, candytuft, cal-
endula, California poppy, iceland poppy, China aster, clematis, columbine,
dahlia, delphinium, Erigeron, gardenia, geranium, hydrangea, honeysuckle,
locust, matrimony-vine, peony, sugar beet, and tulip-tree. Teleomorph state
found on sugar beet in CO, MT, NB, WA, and WY.
Control. Choose resistant vegetable varieties or dust with sulfur. Spray or
dust ornamentals with Karathane or sulfur.
Erysiphe taurica (
Leveillula taurica). On leaves of mesquite.

Erysiphe trina (see Brasiliomyces trina). Oak Powdery Mildew, on tan-
bark oak and coast live oak, in California, causing witches brooms (but
Sphaerotheca lanestris for the common live-oak mildew).

Erysiphe sp. Powdery Mildew on tomato.
POWDERY MILDEWS 445
Leveillula
Cleistothecia globose, or globose-depressed, sometimes concave; asci two- to eight
spored; appendages occose (cottony), simple or irregularly branched; usually similar
to mycelium and interwoven with it.
Leveillula taurica (formerly Erysiphe taurica). On leaves of mesquite.
Commonly causes powdery mildew on tomatoes in Eastern Mediterranean
region and reported on fresh market tomato in California and Utah; also
on cucumber, hibiscus, onion, guar, wild tobacco, cotton and desert bird of
paradise.
Microsphaera
Cleistothecia globose to globose-depressed; appendages branched dichotomously at
apex, often ornate; asci several, with two to eight spores.
Microsphaera penicillata. Named for the alder (Alnus), on which it is
widespread, but best known to gardeners as the Lilac Mildew. It also
infects many other trees, shrubs, and vines, including azalea, beech, bitter-
sweet, birch, catalpa, dogwood, elder, elm, euonymus, forestiera, hazelnut,
magnolia, mountain-holly, plane, New Jersey tea, privet, trumpetvine, and
viburnum. According to some taxonomists the proper name of this species is
Microsphaera penicillata, but M. alni is more familiar and still widely used.
Mildew is prevalent on lilac in late summer and fall, sometimes in dry sea-
sons, almost completely covering foliage with a thick white coating; but,
because it comes so late in the season, it is not very injurious. It is also
common on deciduous azaleas in late summer, forming a very thin gray-
ish white coating with numerous prominent dark perithecia. This species
is more prevalent than the legume mildew on sweet peas, but it is chiey
a greenhouse problem in spring, when temperatures and humidity are less
uniform. The foliage may be malformed, dropping prematurely or drying
out and shriveling.
Among tree hosts oaks are probably most susceptible, but it would seldom
pay to attempt control measures except in nursery rows. On pecans the white
coating starts forming on leaves and nuts in July with occasional defoliation,
shuck splitting, and shriveled kernels. Most commercial pecan varieties are
mildew-resistant.
Blueberry Mildew is caused by a special strain of lilac mildew, reported as
Microsphaera vaccinii, and M. penicillata var. vaccinii. Varieties Pioneer,
Cabot, and Wareham are said to be particularly susceptible; Concord, Jersey,
and Rubel are intermediate; and Stanley, Rancocas, Harding, and Katherine,
highly resistant. On some blueberries the mycelium is conspicuous on upper
leaf surfaces, on others barely visible on underside. Midsummer defoliation
weakens the bushes. Cranberries, farkleberries, trailing arbutus and lyonia
are possible hosts to this strain.
Control. Bordeaux mixture is recommended for pecans two applications,
June and July. Dust blueberries with sulfur. Use sulfur on lilacs and other
ornamentals.
Microsphaera diffusa. General on snowberry, widespread on wolfberry,
coralberry, occasional on black locust, lima bean, kidney bean and soybean.
Appendages are two to four times the diameter of the perithecia, with ulti-
mate branches long, forming a narrow fork.
Microsphaera euphorbiae. On lima bean, euphorbia, roselle.
Microsphaera grossulariae. European Powdery Mildew, occasional on
currant, gooseberry. There is a light weft of mycelium mainly on upper sur-
face of leaves. For the more important American mildew

Sphaerotheca
mors-uvae.
Microsphaera penicillata. Powdery Mildew on Leucotho axillaris.
Microsphaera pulchra. Powdery Mildew on owering dogwood.
Oidium
This term is used for mildews known solely from the conidial stage. In some cases the
type of conidial fructitication may suggest correct genera, but until perithecia are found,
Oidium is preferred.
Oidium araliacearum. Powdery Mildew on English Ivy.
Oidium begoniae. Begonia Mildew, especially important on tuberous
begonias on the West Coast, though it may also occur on brous-rooted
begonias.
Oidium dianthi. Carnation Powdery Mildew.
Oidium euonymus japonici. Euonymus Mildew, general throughout the
South and on the Pacic Coast on Euonymus japonicus. The mycelium forms
a thick felt on the leaf surface, causing some yellowing and defoliation (see
Fig. 3.44). I have seen this disease rampant in foggy coast towns like Beau-
POWDERY MILDEWS 447
Figure 3.44 Powdery Mildew on Euonymus; prevalent in the South
fort, South Carolina, or Mobile, Alabama, and equally severe in semiarid El
Paso, Texas. The washing effect of a water spray applied with pressure, either
by adjusting the hose nozzle or putting the thumb over a portion of the orice,
is a deterrent to this mildew. Sulfur dust can be used, probably Karathane.
Oidium obductum. On oriental plane.
Oidium pyrinum. On crabapple.
Oidium tingitaninum. Citrus Mildew, common in Java, Ceylon, India, but
in this country causing only limited injury to tangerine trees in California.
White patches are formed on upper surface of leaves, the tissue underneath
rst a darker, watery green, then losing color, turning yellowish.
Oidium sp. On greenhouse snapdragons, a white powdery growth on both
leaf surfaces, sometimes on young stems. Control with Karathane or sulfur.
Oidium sp. On avocado, occasionally in Florida, in nurseries or on young
trees in shaded locations. Tips of shoots are killed back; dark green spots
appear on upper leaf surfaces with white mycelium on the underside. The
mildew can be controlled with lime sulfur. On alfalfa in Wisconsin.
Oidium sp. On shrub banana in LA.
Oidium sp. On black medic in CA.
Oidium sp. On Melilotus alba in FL.
Oidium sp. On poinsettia.
Oidium sp. On tomato.
Oidium sp. On Torena.
Phyllactinia
Perithecia are large; appendages are lancelike with a bulbous base. Mycelium does not
send haustoria into epidermal cells of host but forms special branches that pass through
stomata into intercellular spaces; each of these intercellular branches or hyphae sends
a single haustorium into the adjacent cell.
Phyllactinia angulata. Powdery Mildew on elm.
Phyllactinia corylea (syn. P angulata). Powdery Mildew of trees, named
for the hazelnut or lbert but prevalent on many other trees and shrubs,
such as amelanchier, ash, barberry, beach, birch, boxwood, catalpa, chin-
aberry, crabapple, currant, blackberry, raspberry, gooseberry, crape-myrtle,
dogwood, buttonbush, chestnut, elm, elder, fringe-tree, hawthorn, hickory,
hornbeam, holly, linden, oak, plane trees, quince, rose, sassafras, tulip-tree,
walnut and willow. Mildew is seldom serious enough on shade trees to war-
rant control measures, but in the nursery dusting with sulfur may be advis-
able. It is common on lberts in Oregon, but comes so late in the season it
does not affect yield.
Podosphaera
Perithecia globose; one ascus, with eight spores; appendages dark brown or colorless,
dichotomously branched at tip; rarely an extra set of basal appendages present.
Podosphaera leucotricha. Powdery Mildew of apple, also crabapple,
pear, quince, photinia. First noted in Iowa on seedling apples in 1871, this
mildew became more important in orchards when organic fungicides, inef-
fective for mildew, were substituted for sulfur and copper in the apple-scab
schedule. Twigs, foliage, blossoms and fruits may be disgured, stunted,
deformed or killed. Gray to white felty patches are formed on leaves, usually
on underside. Leaves are crinkled, curled, sometimes folded longitudinally
and covered with masses of powdery spores. They soon turn brittle and
POWDERY MILDEWS 449
die, resulting in decreased yield. The same powdery growth starts on 1-year
twigs, but in midsummer it is transformed into a brown, felty covering, in
which minute, dark perithecia are embedded in dense aggregations. Infect-
ed twigs are stunted or killed. The fungus winters as dormant mycelium
on twigs or in buds. Such buds produce shriveled blossoms and no fruit.
Fruit produced on infected twigs is stunted or russeted. Jonathan variety is
especially susceptible.
Podosphaera clandestina (P. clandestina var. tridactyla). Powdery Mil-
dew of cherry, occasional on plum, peach, apricot, apple, pear, quince,
hawthorn, serviceberry, spirea. Budded sour cherry is most severely attacked,
but the disease is seldom serious except on nursery stock. Young leaves and
twigs are covered with a white mycelium and powdery spores. Leaves are
curled upward; terminal leaves are smaller; twig growth is stunted. Sulfur
sprays or dusts will control.
Podosphaera pannosa (formerly Sphaerotheca pannosa var. rosae). Rose
Mildew, general on rose; distinct from peach mildew but apparently not
conned to rose, since apricots growing near roses have been infected. More
than one strain may be involved. Rose mildew is found wherever roses grow.
Always a problem with greenhouse roses, it was enhanced when aerosol
treatments for red spiders and other pests were substituted for old-fashioned
syringing. Mildew increased in garden roses when ferbam and other new
organics replaced the old sulfur and copper in the blackspot sprays. Rose
mildew is omnipresent along the Pacic Coast and is serious in the semi-
arid Southwest. In the East, it appears on small-owered ramblers such as
Dorothy Perkins and Crimson Rambler in May, and may be quite serious
on hybrid teas and some oribundas in late summer, with the advent of cool
nights.
The rst symptom may be a slight curling of leaves, with the mycelial growth
such a light and evanescent weft as to be almost unnoticed. Later the white
coating is conspicuous from the chains of conidia produced lavishly over
the surface. The coating may cover buds, resulting in no bloom or distorted
owers. Leaves often have a reddish or purplish cast under the white myceli-
um and sometimes turn black. They may be slightly blistered. On canes,
the growth is heavier and more felty, especially near thorns. Toward the end
of the season perithecia may be found on canes, but they are not common,
and I have not seen them on leaves except on a Rugosa rose at Ithaca, New
York. Mildew is prevalent on soft succulent shoots, fostered by an excess of
nitrogen.
Control. Sulfur dusts have been standard treatment for garden roses for many
years; to be effective dusting must be started at the rst sign of mildew,
before the mycelium gets too thick. Sulfur may be injurious to roses in very
hot weather. Choice of variety of rose is important. Shiny-leaved climbers
like Dr. Van Fleet seldom have mildew, and the shrub polyantha, The Fairy,
is very resistant. Many red roses, hybrid teas and uoribundas, are especially
susceptible, but the orange-red oribunda Spartan remains free from it (in
my own experience). Garden planning avoids a lot of mildew trouble. Keep
the plants well spaced, in beds away from buildings, and not surrounded by
tall hedges or walls.
Podosphaera tridactyla. Recently reported on almond in California and
the most common mildew on apricot, causing large nonnecrotic lesions on
leaves.
Sphaerotheca
Appendages simple, exuous, resembling hyphae; only 1 ascus in a perithecium.
Cystotheca lanestris (formerly Sphaerotheca lanestris). Powdery Mildew
of coast live oak on Quercus agrifolia in California, reported also on white,
southern red, bur and post oaks. The disease is most destructive in the nar-
row coastal plain. The most conspicuous symptom is a powdery white, stunt-
ed growth developing from certain terminal or lateral buds. The shoots are
swollen, eshy, with much shortened internodes. Foliage on such shoots is
often reduced to pale yellow, bractlike leaves, which turn brown, dry, and
shrivel; these shoots resembles witches brooms. On leaves developing from
normal buds and shoots, the fungus forms a dense layer on both surfaces,
more abundant on the lower side. This species is sometimes called the brown
mildew because the grayish-white mycelium changes to tan and then brown
with age. Perithecia are formed in the brown felt, abundantly in some years,
rarely in others. In southern California the fungus may winter in the conidi-
al state, with widespread leaf and shoot infections coming from wind-borne
spores.
Control is not easy. Spraying with lime sulfur in March and October is fairly
effective but may be phytotoxic at high temperatures and low humidity. Wet-
table sulfur has not been consistently effective. Removal of witches brooms
by pruning back to normal lateral branches is effective only if the tree is
slightly susceptible and conditions for reinfection are unfavorable. Heavy
POWDERY MILDEWS 451
pruning stimulates new growth and increases the amount of mildew. The
Holm or holly oak is apparently resistant to mildew and well adapted to the
coastal region.
Sphaerotheca castagnei (see Sphaerotheca fuligena). On buffaloberry,
spirea.
Sphaerotheca fuliginea (formerly Sphaerotheca castagnei). On buffalober-
ry, spirea.
Sphaerotheca fuliginea. Powdery Mildew of summer squash and cucur-
bits.
Sphaerotheca fusca. Powdery mildew; on ground-cherry.
Sphaerotheca lanestris (see Cystotheca lanestris). Powdery Mildew of
coast live oak on Quercus agrifolia in California, reported also on white,
southern red, bur and post oaks.
Sphaerotheca macularis (S. humuli). Hop Mildew, also on fruits, black-
berry, dewberry, gooseberry, raspberry, strawberry, rose (probably rarely
in this country), and other ornamentals, including Agastache, betony, buf-
faloberry, delphinium, Epilobium, Erigeron, gaillardia, geranium, geum,
gilia, hawksbeard, hawkweed, Hydrophyllum, kalancho, matricaria, mead-
owsweet, ninebark, Polemonium, phlox, sumac, spirea, tamarisk and Ver-
nonia.
This mildew may be important on Latham variety of raspberry, appearing
on new canes when they are 2 to 3 feet high. The tip leaves are dwarfed,
mottled, and distorted, almost as if they had mosaic. The undersurface of
leaves is water-soaked or has the familiar white coating. There is no specic
control except to space plants for free air circulation.
The powdery mildew sometimes serious on strawberries in northeastern and
Pacic Coast States is probably a special strain. The edges of affected leaves
curl upward, exposing the lower surface, where the powdery frosty growth
is evident. Fruit, stems, and berries may be affected, with fruit often failing
to color. Resistant varieties include Sparkle, Puget Beauty, Siletz and India.
Sphaerotheca mors-uvae. American Gooseberry Mildew, also on cur-
rant; sometimes the limiting factor in gooseberry production. Fruits dry up
with a brown, felty covering. Leaves and canes are stunted with the usual
white coating. Perithecia are formed on canes, and ascospores are discharged
in early May as fruit is set. Conidia for secondary infection are produced
within 10 days. Spray with lime sulfur immediately after bloom.
Sphaerotheca pannosa var. persicae. Peach Mildew, general on peach,
also on almond, apricot, nectarine, matrimony-vine, and Photinia. The
mycelium is pannose (ragged) or in dense patches, persistent, usually satiny,
shining white, or sometimes grayish or brown. Immature fruits are highly
susceptible. They have brown blotches and are scabby and malformed. The
fungus winters in shoots. Nonglandular varieties Peak and Paloro are more
affected than glandular Walton, Johnson, Halford, and Stuart, at least in
California. Sulfur in the spray schedule for brown rot should control mildew
without additional treatments. Karathane is effective but very slightly phyto-
toxic. Lime sulfur is recommended.
Sphaerotheca pannosa var. rosae (
Podosphaera pannosa). Rose Mil-

dew, general on rose; distinct from peach mildew but apparently not con-
ned to rose, since apricots growing near roses have been infected.
Sphaerotheca phytoptophila. Associated with gall mites causing witches
brooms on hackberry. The mycelium is evanescent; perithecia are formed
inside loose scales of enlarged buds.
Sphaerotheca sp. On Tolmiea, pick-a-back plant, in greenhouse.
Uncinula
Perithecia globose; appendages uncinate, slightly coiled at tips; several asci, with two to
eight spores.
Pleochaeta prosopidis (formerly Uncinula prosopidis).
Uncinula circinata. On maples, Virginia creeper, western soapberry.
Uncinula clintonii. General on American linden.
Uncinula exuosa (see Uncinuliella exuosa). Horse-Chestnut Pow-
dery Mildew, on Aesculus spp., including red, yellow and Ohio buckeye,
widespread in central and eastern states.
Uncinula macrospora. General on American and winged elms.
Uncinula necator. Grape Powdery Mildew, general on grapes, also on
Ampelopsis; common in late summer on eastern grapes but not serious;
a major problem in California. Leaves, canes and young fruits are cov-
ered with white patches; growth is often distorted. Late in the season the
white mycelium disappears and the spots appear brown or black; berries are
russeted or scurfy, failing to mature.
Control. Keep California grapes covered with a light coating of sulfur dust.
Apply when new shoots are 6 to 8 inches long; when they are 12 to 16 inches,
14 days later; when shoots are 2 to 3 feet; when fruit is half.grown; when
POWDERY MILDEWS 453
fruit begins to ripen. If some of the applications are omitted, and mildew
gets a head start, wettable sulfur is used as an eradicant spray. Karathane is
also effective. Copper sprays are often used in the East, if any are necessary.
Bicarbonate sprays are excellent as protectant and eradicant.
Uncinula parvula; U. polychaeta. Widespread on hackberry and southern
hackberry.
Uncinula prosopidis (see Pleochaeta prosopidis). On mesquite.
Uncinula salicis (see Uncinula adunca). Willow Powdery Mildew, also
on pussy willow and poplar, sometimes causing defoliation but not often
serious.
Uncinuliella australiana. (not Erysiphe lagerstroemiae as reported earli-
er in U.S.). Crape-Myrtle Powdery Mildew on crape-myrtle only, from
Maryland to Florida and Texas, the most serious disease of this shrub. The
perithecia have been found only in Florida, but presumably it is the same
species throughout the host range. The disease appears on young shoots in
early spring, later infecting leaves and different parts of the inuorescence.
Affected parts are covered heavily with a white mealy to dusty growth;
young leaves are stunted, often less than one-third normal size but abnor-
mally thickened. Internodes are short, ower stems stunted; buds often fail
to develop owers. Infected portions often have a reddish discoloration under
the white coating. Diseased leaves and buds drop in a week or two, but
stems may sprout again and sometimes produce normal growth in hot weath-
er.
The fungus winters as mycelium in dormant buds and in spring covers such
buds with a dense white coating of conidia, the source of primary infection,
which starts as small, circular white patches on young leaves. Spores pro-
duced in abundance on these patches account for rapid spread of the disease
until midsummer heat.
Control. Spray with lime sulfur when buds burst in spring and repeat 2 weeks
later. If the initial infection is not checked, spray wettable sulfur or dust with
sulfur.
Uncinuliella exuosa (formerly Uncinula exuosa). Horse-Chestnut
Powdery Mildew, on Aesculus spp., including red, yellow and Ohio buck-
eye, widespread in central and eastern states. This mildew gives a very thin
coating on the leaf surface, supposedly mostly on the underside although
I have seen it on the upper. Perithecia are numerous, small, barely dis-
cernible with the naked eye. Control is usually unnecessary except in nurs-
eries. A copper spray used for blotch will also control mildew.
Uncinula adunca (formerly Uncinula salicis). Willow Powdery Mildew,
also on pussy willow and poplar, sometimes causing defoliation but not often
serious. The growth is in diffused or circumscribed patches on both leaf sur-
faces.
ROTS
A rot is a decay, a decomposition or disintegration of plant tissue. It may be
a hard dry decay or a soft and squashy one. It may affect root or rhizome,
stem, tree trunk, blossom or fruit. Some rots also affect leaves, but diseases
that are primarily of foliage are more often designated leaf spots or blights.
Rots caused by bacteria are discussed under Bacterial Diseases.
There are a great many wood rots of trees, recognized by the sporophores
or conks of the various species of Fomes, Polyporus, and other shelving or
bracket fungi. By the time these signs appear, it is usually too late to do any-
thing about the disease. The tree-rot fungi enter through unprotected wounds
either pruning cuts or breaks due to wind and icestorms. For proper prun-
ing methods and treatment of wounds, see U.S. Department of Agriculture
Farmers Bulletin 1896, Care of Damaged Shade Trees, Tree Maintenance
by P. P. Pirone or Tree Experts Manual by Richard R. Fenska. The fact that
tree wound dressings are now available in convenient aerosol bombs should
make it easier for home gardeners to protect pruning cuts from wood-rotting
fungi.
Physalospora (Acanthorhynchus)
Perithecia separate, innate, beaked; spores one-celled, dark.
Acanthorhynchus vaccinii (see Physalospora vaccini). Cranberry Blotch
Rot, a common fruit rot thriving in warmer sections, more important in New
Jersey than in Massachusetts.
Physalospora vaccini (formerly Acanthorhynchus vaccinii). Cranberry
Blotch Rot, a common fruit rot thriving in warmer sections, more important
in New Jersey than in Massachusetts. The rot starts as a small, light-colored
spot on the berry, spreading to destroy the whole fruit, with dark blotches
on the skin. The fungus may invade leaves, but it seldom fruits on them
until they have fallen. Cranberry bogs in New Jersey may need three or four
sprays of Bordeaux mixtures starting at midbloom, but in Massachusetts two
are sufcient.
Acremonium
Acremonium sp. Root Rot on melon and watermelon.
Alternaria
Blights.
Alternaria alternata. Fruit Rot on tomato and black pit disease on potato
tubers (stored).
Alternaria citri. Alternaria Rot of citrus fruits, navel-end rot, black rot,
widespread, prevalent in warm dry sections, but not too serious. In oranges
the rot is most common in the Washington Navel variety a rm, dry, black
rot at the navel end, often in only one segment, with fruit coloring prema-
turely, appearing sound on the outside. In lemons the disease is a soft, dark
internal rot of old or weak fruit in storage. Firm dark brown spots are formed
on the rind. Grapefruit sometimes has a dark internal storage rot, not readily
discernible externally.
Control. Chemical treatment after picking is not very satisfactory. Produce
sound fruit in the orchard; avoid holding too long on the tree; avoid holding
weak or old fruit too long in storage; store at low temperatures.
Alternaria mali. Fruit Rot, widespread storage rot of apple, sometimes
quince. Also a weak parasite enlarging injured spots on foliage. Try captan
at 6- to 14-day intervals.
Alternaria radicina (Syn. Stemphylium radicinum). Black Rot of carrots,
a soft storage rot of roots held over winter. Rot may start at the crown or
from some wound on the side of the root. Initial infection may be in eld or
in storage house; a black mycelial weft with large, brown muriform spores
develops over the rotted tissue. There is no control except to choose rm,
healthy roots for storage and to store at low temperatures.
Alternaria solani. Collar Rot of tomato, also fruit rot and early blight, gen-
eral on tomato with the collar rot stage most frequent in the South.
Blights.
Alternaria zinniae. Stem Rot on Ageratum.
ROTS 457
Alternaria sp. Flower Rot of Vanda orchids, causing infection in transit
along with Botrytis.
Alternaria sp. On Schefera in Florida.
Alternaria sp. Calyx-End Rot on apple.
Amphobotrys
Cankers and Dieback.

Amphobotrys ricini. Stem Rot and Wilt of poinsettia.
Aphanomyces
Phycomycetes, Saprolegniales.
Thallus composed of cylindricaI branching hyphae without denite constrictions; spo-
rangium cylindrical, threadlike, swarm spores arranged in a single row and encysting at
the mouth; saprophytic or parasitic, living in the soil and causing root rots or damping-
off.
Aphanomyces cladogamus. Causing rootlet necrosis of tomato, pepper,
spinach, and a severe root rot of pansy.
Aphaomyces cochlioides. A seedling disease of sugar and table beets, part
of the complex called black root; causing tip rot, a wilting of tops. Crop
rotation and proper fertilization are helpful.
Aphanomyces euteiches. Pea Root Rot, also on bean, sweet pea and
perennial pea. The fungus is also a weak parasite in roots of many non-
legumes. First described in 1925, the fungus probably existed earlier in
various root disease complexes and was responsible for giving up land for-
merly devoted to canning peas. Considered the most important of the pea
root rots, found in every district, it is particularly destructive in eastern and
central states.
The fungus is parasitic on subterranean parts, causing root and stem rot in
peas of all ages, symptoms and crop yield varying with the time of infection.
If the root system is invaded when only three or four nodes are formed, the
plant may wilt and die suddenly; later invasion results in dwarng and drying
out of foliage from the ground upward. When seedlings are pulled out of the
ground, the roots do not break off but come out as a brous string or vascular
cylinder freed from cortex. The fungus invades only the cortex or roots and
base of stem, causing softening and rapid decay of tissue. Large numbers of
thick-walled oospores are formed in the cortex; these may remain viable in
the soil more than one season.
It is a novel root pathogen on alfalfa in Canada.
Control. A well-drained soil with low moisture content decreases rot. When
soil moisture is at 45% of saturation, there is no disease; at 75% there may
be more than 70% infection. Nitrogenous fertilizers are helpful.
Aphanomyces raphani. Radish Black Root and Damping-Off, wide-
spread; more important on long-rooted icicle varieties. Also on Abyssinian
mustard, cabbage, Chinese cabbage, Chinese kale, honesty, mustard green,
rape, rocuet salad, sea-kale, Spanish mustard, wild radish and Brassica
robertiana. Small, steel-gray to black areas appear around point of emer-
gence of secondary roots. Enlarging roots are constricted and turn black.
Rotation is essential for control. Choose globe rather than long varieties.
Armillaria
Basidiomycetes, Agaricales.
One of the mushrooms, cap-shaped on a stalk with an annulus or ring but no volva (cup)
at the base; gills attached to the stem; spores white (see Fig. 3.45).
Armillaria mellea. Mushroom Root Rot of trees and shrubs, also known
as Armillaria root rot or toadstool disease, rst described in America in 1887,
known in Europe a hundred years earlier. The fungus is called honey mush-
room, honey agaric, oak fungus and shoestring fungus. Although the honey-
colored toadstools are often seen in the East around rotting tree stumps and
may occasionally cause death to weak ornamental trees, the chief damage is
west of the Rocky Mountains, especially in California, where most fruit and
nut crops and ornamental trees and shrubs are menaced.
The decay is of the roots and root crown. Sheets of tough, fan-shaped myceli-
um are found between bark and wood, the latter changing to light tan, becom-
ing soft and watery in texture. Clumps of toadstools are often found at the
base of dead or dying trees, especially in autumn, but do not always appear
in dry seasons. They are honey-colored or light tan, with a stalk 4 to 6 inches
or more high and a cap 2 to 4 inches across, often dotted with brown scales.
Basidiospores formed along the gills are wind-borne. They can establish
themselves in old stumps and dead trees but cannot infect healthy trees.
The latter are infected in the ground by means of black or brown cordlike
ROTS 459
Figure 3.45 Mushroom Root-Rot Fungus, Armillaria mellea
rhizomorphs, the shoestrings, which grow out from infected roots a short
distance through the soil. On meeting and penetrating a healthy root, the fun-
gus progresses along the cambium layer, working up to and girdling the root
crown. Leaves are dwarfed, turn yellow or fall prematurely; on small trees
all foliage may die simultaneously. On conifers, particularly pines, there is
an abnormal ow of resin from the root collar.
Trees subnormal in vigor and suffering from drought are most injured.
Orchards of citrus and other fruits on lands recently cleared of oaks are
liable to enormous damage unless resistant rootstocks are used. The rot is
found less often on dry hillsides than in valleys near streambeds, where ood
waters deposit soil and infected debris around root crowns, or in places kept
too wet by articial watering. Ornamental trees and shrubs are often injured
when extra soil is added in grading and terracing, and are then kept too wet
by watering the lawn frequently.
The list of susceptible plants is far too long to be given in entirety. A repre-
sentative selection includes almond, apple, apricot, avocado, cherry, citrus,
currant, grape, incense cedar, peach, pear, plum and raspberry, hickory, l-
bert and walnut, California pepper-tree; oaks, pines, spruce and sycamore;
azalea, rhododendron, boxwood and rose; (root, crown, and stem rot on)
African daisy and (stem rot on) sunower;and sometimes other herbaceous
plants such as begonia, carnation, dahlia, narcissus, peony, rhubarb, and
strawberry.
Control. Use resistant plants where possible. Of fruits, only French pear,
Northern California black walnut, g and persimmon are sufciently resis-
tant to grow safely on infested soil. Some plants can be grafted onto resistant
rootstock such as Myrobalan 29. The University of California has prepared
a list of resistant or moderately resistant ornamental shrubs. Some on the
list are Acacia decurrens var. mollis, A. verticillata, Buxus semipervirens,
Ilex aquifolium, Lonicera nitida, Prunus ilicifolia (hollyleaf cherry), P. lyoni
(Catalina cherry), Pyracantha coccinea and var. lalandii but not P. angusti-
folia, which is susceptible.
Moderately resistant shrubs include Abelia grandiora, Darwin, Japanese,
and Mrs. Wilson barberry, Mexican orange (Choisya), Elaeagnus argentea,
Euonymus japonica, Japanese privet, Myrtus communis, Pittosporum tobira
and Spiraea prunifolia.
Mechanical measures are often helpful. Excavate and expose the root crown;
remove diseased portions of bark and affected small roots. Paint wounds
with a pruning wound compound. Leave the treated roots exposed until cool
weather in autumn. Trenching or digging a ditch around a plot will restrict
the disease temporarily, but roots will grow through the ditch in time.
Carbon disulde is still recommended as a soil disinfectant, applied in stag-
gered rows, in holes 18 inches apart each way, which should be immediately
closed by tamping. Hand applicators are available for injecting the disulde
6 to 7 inches deep. This treatment is for land where valuable trees have been
removed; one cannot go closer to a healthy tree than the edge of the branch
spread. After treatment, the land should remain fallow for at least 60 days,
and then be ploughed before planting.
Ascochyta
Blights.
Ascochyta pinodes. Foot Rot of peas. Of the three species that make up the
Ascochyta blight complex, this one produces most denitely a foot rot, with
infection at the root crown or base of stem.
ROTS 461
Aspergillus
Conidiophores have a round head at the top, with radially arranged bottle-shaped sterig-
mata that bear conidia in chains; spores are one-celled, globose to ellipsoid, hyaline.
Bread molds are in this genus. When, rarely, a sexual fruiting body (cleistothecium) is
formed, the species is placed in the order Eurotiales.
Aspergillus alliaceus. Cladode Rot, Stem and Branch Rot on Cereus
and Opuntia cacti. This is a high temperature species. Spores are yellow in
mass.
Aspergillus fumigatus. Wound Rot (storage) on beet.
Aspergillus niger. Calyx-End Rot of dates, Fig Smut, Bunch Mold of
grapes, Pomegranate Rot, Black Mold of peach. Crown Rot of peanut;
also market and storage rot of shallot, onion, apple and potatoes. The fungus
is a weakly parasitic black mold invading ripe tissue through wounds. In
dates, the interior of the fruit is lled with a black dusty mass of spores,
spread to a large extent by the dried-fruit beetle. Practice orchard sanitation;
keep decaying fruits cleaned up so insects cannot carry spores.
Aspergillus niger var. oridanus. Wound parasite on Dracaena. Lower
stem black, rotted, with dark brown spore masses.
Aspergillus spp. Green and yellow molds causing secondary rots of many
fruits and some vegetables in storage.
Lasiodiplodia (Botryodiplodia)
Blights.
Botryodiplodia theobormoae (see Lasiodiplodia theobromae). Root Rot
on apple
Lasiodiplodia theobromae (formerly Botryodiplodia theobormoae). Root
Rot on apple.
Botryosphaeria
Blights.
Botryosphaeria dothidea. Fruit Rot of peach and grape.
Botryosphaeria obtusa. Fruit Rot of peach, Black Root Rot of apple.
Botryosphaeria rhodina. Fruit Rot of peach.
Botryosphaeria ribis (Anamorph, Dothiorella gregaria). Dothiorella Rot
of avocado and citrus, Black Fruit Rot of apple and pear, Nut Rot of
tung oil. On avocado this is a soft rapidly spreading surface rot, starting
from small spots when fruit begins to soften. The fruit may be covered with
decay spots by the time it is usable. The fungus winters in dead twigs, in tip-
burned leaves, and enters the avocado while it is still on the tree. Two sprays,
mid-September and early October, using bordeaux mixture, have given fairly
good control. Remove dead wood from trees, to reduce source of inoculum,
and pick fruit early.
On lemons and other citrus fruits the rot starts as a discoloration around the
button, becoming a brown, leathery but pliable decay. When fruit is entire-
ly involved it becomes olivaceous black. On tung, brown lesions appear on
green fruit, which drops prematurely. See further under Cankers.
Botryotinia
Blights.
Botryotinia convoluta. Botrytis Crown Rot of iris, Gray Mold Rot on
rhizomatous iris, rst recorded in Canada in 1928 and apothecia later pro-
duced in culture. The chief diagnostic character is the presence of many shin-
ing black sclerotia, much convoluted and agglomerated into large clusters on
rotting rhizomes. These are often found in spring on plants that started into
the winter apparently healthy, for the fungus is active in cool, wet weath-
er. Conidiophores are brown, formed in fascicles, and bear dense clusters of
light brown ovate or slightly pyriform conidia. They appear in spring grow-
ing from or near sclerotia. Affected plants do not start spring growth.
Botrytis
Blights.
Botrytis aclada. Gray Mold Neck Rot of onion, also shallot and garlic;
widespread. This is usually found on bulbs after harvest, infection taking
place through neck tissue and scales appearing sunken and cooked. Scle-
ROTS 463
rotia are rst white, then dark, 2 to 4 mm across. Conidiophores and conidia
forming the gray mold are produced directly from mycelium in tissue or
from sclerotia. Articially cure bulbs after harvest to cause rapid dessication
of neck tissue; store at low temperature. Colored varieties keep better than
white.
Botrytis byssoidea. Mycelial Neck Rot of Onion. The fungus is much like
B. aclada but produces more mycelium and less profuse gray mold.
Botrytis cinerea. Gray Mold Fruit Rot, Cosmopolitan on peach, cherry,
plum, pomegranate, quince, pear, grape, strawberry, pepper, tomato and egg-
plant. Also causing a leaf rot of hothouse rhubarb and a rot of carrot, lettuce,
celery and onion. See further under Blights.
Botrytis gladiolorum. Botrytis Neck Rot, CormRot, Blight of gladiolus.
Blights.
Botrytis porri. Seedborne causing natural infection.
Botrytis (Teleomorph, Botryotinia) squamosa. Small Sclerotial Neck
Rot of onion. Elliptical leaf lesions with withering of tips.
Botrytis tulipae. Bulb Rot of Vidalia sweet onion.
Brachysporium
Conidiophores brown, erect, usually solitary, septate; conidia dark, unequally two- or
more-septate; attached to apical cell of conidiophore by a short narrow cell; saprophytic.
Brachysporium tomato. Fruit Rot of tomato.
Calonectria
See Cylindrocladium under Blights.
Calonectria crotalariae. Basal Stem Rot of oleander.
Calonectria sp. (Anamorph, Cylindrocladium). Crown and Collar Rot on
papaya.
Catenularia
Hyphae dark; conidophores simple or sparingly branched, with terminal chains of coni-
dia; spores dark, one-celled.
Catenularia fuliginea. Fruit Rot of date.
Mycocentrospora
Spores hyaline, liform, with long, whiplike tapering beaks, several cross walls and
a swordlike appendage from basal cell; mycelium dark.
Centrospora acerina (see Mycocentrospora acerina). Black Crown Rot
of celery, Storage Rot of carrot.
Mycocentrospora acerina (formerly Centrospora acerina). Black Crown
Rot of celery, Storage Rot of carrot (see Fig. 3.46). On celery the dis-
ease appears 7 or 8 weeks after stock has been placed in cold storage with
pale, ochraceous lesions at the crown end, gradually turning black, some-
times reddish. The fungus lives in the soil; use infested elds for early celery
to be marketed without storage. The same species also causes leaf spot of
pansy.
Cephalosporium
Leaf Spots.
Cephalosporium carpogenum. Fruit Rot on apple in storage, reported
from Washington and Pennsylvania.
Cephalosporium gregatum (see Phialophora gregata). Brown Stem Rot
of soybean.
Phialophora gregata (formerly Cephalosporium gregatum). Brown Stem
Rot of soybean.
A vascular disease of major importance in the Midwest, also present in Flori-
da, North Carolina and Virginia. It has been controlled with a long rotation
5 years corn, 1 year soybeans.
Ceratocystis
Cankers.
ROTS 465
Figure 3.46 Storage Rot on Carrot
Ceratocystis mbriata (Endoconidiophora mbriata). Sweet Potato
Black Rot, found wherever sweetpotatoes are grown, most destructive
in storage but present also in seedbed and eld. Round, blackish spots
extend into vascular ring or deeper; sprouts are sickly with black cankers
below ground or are killed. The fungus winters in storage houses, on wild
morning-glory and other weeds near the eld and in soil, where it remains
viable for several years. Spores are spread by the sweetpotato weevil and in
wash water if potatoes are washed before storing. This fungus also infects
Jacquemontia.
Control. Standard treatment has been disinfection of planting stock in a solu-
tion of borax. Using pulled sprouts provides plants free from black rot. Plan
a 4-year rotation; sort carefully before storage; cure quickly at high temper-
ature and humidity. Yellow Jersey is highly susceptible; some varieties are
quite resistant.
A strain of this fungus is reported causing Black Cane Rot in propagating
bed of Syngonium auritum (Philodendron trifoliatum) in a California nurs-
ery. Brown to black water-soaked girdling cankers, often on parts in contact
with the soil, cause yellowing and death of foliage. The fungus can be erad-
icated by treating canes with hot water, 120F for 30 minutes.
Ceratocystis wageneri (see Ophiostoma wageneri). Root Rot of lodgepole
pine and ponderosa pine.
Ophiostoma wageneri (formerly Ceratocystis wageneri). Root Rot of
lodgepole pine and ponderosa pine.
Apostrasseria (Ceuthospora)
Pycnidia in a valsoid stroma; conidia oblong to bacillar, extruded in tendrils; conidio-
phores obsolete or none.
Apostrasseria lunata (formerly Ceuthospora lunata). Black Rot of cran-
berry, developing in berries after picking. The fruit turns dark and soft. The
disease is more important in Washington and Oregon. Spraying for other
cranberry diseases largely controls this rot. Pick berries when dry; avoid
bruises; keep them cool.
Ceuthospora lunata (see Apostrasseria lunata). Black Rot of cranberry,
developing in berries after picking.
Chalara (Chalaropsis)
Molds.
Chalara thielaviopsis (formerly Chalaropsis thielaviopsis). Root Rot on
poinsettia.
Chalaropsis thielaviopsis (see Chalara thielaviopsis). Root Rot on poin-
settia.
ROTS 467
Armillaria (Clitocybe)
Basidiomycetes, Agaricales
One of the mushrooms, with gills typically decurrent (running down the stem), cap
homogenous and conuent with eshy stripe, which has neither ring nor cup; spores
white or very lightly colored.
Armillaria tabescens (formerly Clitocybe monadelpha). On privet, apple.
Armillaria tabescens (formerly Clitocybe tabescens). Mushroom Root
Rot, Clitocybe Root Rot of citrus, pecan and other fruits and many orna-
mentals. This root rot is as devastating in Florida as Armillaria rot is in Cal-
ifornia and very similar (some say the pathogen is identical). It is important
in the decline of citrus groves, on orange, grapefruit, lemon, tangerine and
lime on rough lemon stock; is very destructive to Australian-pine (Casua-
rina); and has been reported on more than 200 species in 59 plant fami-
lies, including Acalypha, avocado, arborvitae, apricot, camellia, castor-bean,
cherry-laurel, crape-myrtle, cotoneaster, cypress, dogwood, Eugenia, euca-
lyptus, grape, guava, glorybush, Hamelia, holly, Ligustrum, juniper, jasmine,
loquat, oleander, poinciana, pomegranate, pear, Parkinsonia, rose, viburnum
and wax-myrtle. In recent years Clitocybe root rot has become economically
important on Georgia peach trees and has killed many lychee trees in Florida.
It is said to account for 75% of rose mortality in some sections.
Symptoms of decline do not ordinarily develop until the pathogen has been
working a number of years and has killed a large part of the root system.
Often mushrooms are present at the base of trees before the tops show more
than a slight yellowing or lack of vigor; but if soil is removed from the
root crown, many lateral roots are found dead, and often the taproot is also
gone. Infection starts at some point on the lateral roots, spreads to the base
of the tree, and then to other roots. Sometimes there is gumming at the
crown extending upward on the trunk. Mycelial fans or sheets are present
between bark and wood; the clusters of mushrooms developing at the base
are similar to those of Armillaria, but the black shoestring rhizomorphs
are lacking. Instead, there are sometimes black, hard stromatic outgrowths
from ssures in bark of infected roots. The fruiting clusters develop in fall,
from mid-September to December. The caps are light tan to honey-colored,
2 to 3 1/2 inches in diameter. The rot is most prevalent on land cleared
of oaks and other hardwoods, also on sandy, well-drained land subject to
drought.
Control. Citrus trees on sour orange stock are quite resistant. Surgical treat-
ment for fruits and ornamental trees is often quite successful. Remove the
soil at least 2 or 3 feet from the trunk, working carefully to avoid injuring
healthy roots. Cut off all dead roots, ush with the root crown and remove
any infected oak or foreign roots in the vicinity. Cut out dead and infected
bark at the root crown or the base of the trunk, being sure to collect all chips
(on heavy paper placed under exposed roots) for burning. Paint all exposed
surfaces with a pruning wound compound and ll in partially, disinfesting
the soil with bordeaux mixture. The root crown can be left exposed to aer-
ation and drying or, if too large a proportion of the root system has been
lost, new roots can be stimulated by mounding the soil around the base to
a height of several inches above the partial girdle. The new roots will come
from callus formed at the margin of living bark.
Trenches 2 or 3 feet deep will aid in preventing spread to healthy trees. Fal-
low soil can be treated with carbon disulde;

Armillaria mellea.
Clitocybe monadelpha (see Armillaria tabescens). On privet, apple.
Clitocybe tabescens (see Armillaria tabescens). Mushroom Root Rot,
Clitocybe Root Rot of citrus, pecan and other fruits and many ornamen-
tals.
Colletotrichum
Anthracnose.
Colletotrichum acutatum. Bitter Rot of apple fruit; fruit rot of grape.
Colletotrichum capsici. Ripe Rot of pepper, Boll Rot of Cotton.
Colletotrichum circinans. Onion Smudge, surface rot, also on shallot,
garlic and leek. Bulb or neck has a dark green or black smudge, often covered
with stiff bristles of the acervuli of the fungus. Smudge is more prominent in
white onions; it is conned to the neck of colored bulbs. The fungus winters
on mature onions, on sets or in soil. It develops in the eld at a fairly high
temperature and soil moisture with most of the damage just before harvest.
Cure rapidly after harvest; rotate crops; clean up debris; change to colored
onions if the rot is too serious on white.
Colletotrichum coccodes. Root Rot and Wilt of greenhouse tomato.
Colletotrichum gloeosporioides. Fruit Rot of grape and pepper.
Colletotrichum gloeosporioides (formerly Colletotrichum nigrum). Fruit
Rot of pepper, probably general on pepper in South and East. The fun-
ROTS 469
gus is a wound parasite on pepper pods. The spots are irregular, indenite,
depressed, blackish. Numerous acervuli with stout setae are scattered over
spots.
Colletotrichum lilii. Black Scale Rot of Easter lily, Brown Scale. First
noticed in Louisiana in 1937, the rot immediately threatened the lily industry
in that section. Bulbs are brown to nearly black when dug, with outer scales
most affected. Young lesions start as irregular light brown areas, then become
black and sunken owing to collapse of epidermal cells and subepidermal
layers. Oldest lesions are nearly black, with tissue dry and shriveled. Stems
and roots are not affected. The acervuli are small, gregarious, with many dark
brown setae and continuous hyaline conidia.
Colletotrichum nigrum(see Colletotrichum gloeosporioides). Fruit Rot of
pepper, probably general on pepper in South and East. The fungus is a wound
parasite on pepper pods. The spots are irregular, indenite, depressed, black-
ish. Numerous acervuli with stout setae are scattered over spots.
Frammulina (Collybia)
Margin of young cap turned in; gills not decurrent; stipe central; no annulus or volva;
spores white or light; causing wood rots.
Collybia velutipes (see Frammulina velutipes). Heart Rot, White Sap-
wood Rot of hardwoods.
Frammulina velutipes (formerly Collybia velutipes). Heart Rot, White
Sapwood Rot of hardwoods. The fungus is a small toadstool with central
stem, base covered with dark brown velvety hairs, cap yellowish or brown-
ish. The disease is a soft spongy white rot of sapwood of living hardwoods,
particularly basswood, horse-chestnut, American elm and on catalpa. The
toadstools are formed in clusters at wounds.
Coniophora
Pileus resupinate, effuse; hymenium with one layer, cystidia lacking; spores dark; wood-
destroying.
Coniophora puteana. Brown Cubical Rot of conifers and sometimes
hardwoods on slash, building timbers, and sometimes living trees. The
crustlike eshy fruiting bodies are a little over 2 inches in diameter, olive to
brown with whitish margins and smooth to slightly waxy surface.
Coniophora corrugis (see Cylindrobasidium corrugum). Sapwood Rot of
alpine r.
Cylindrobasidium corrugum (formerly Coniophora corrugis). Sapwood
Rot of alpine r.
Coniella (Coniothyrium)
Cankers.
Coniella diplodiella (formerly Coniothyrium diplodiella). White Rot of
grapes, appearing spasmodically on grapes but not one of the more impor-
tant diseases. Small pycnidia appear on outside of fruit cuticle as shiny, rosy
points, also on leaves. Infection is usually through wounds. Spots on ripe
grapes are grayish, with brown borders.
Coniothyrium diplodiella (see Coniella diplodiella). White Rot of grapes,
appearing spasmodically on grapes but not one of the more important dis-
eases.
Coprinus
Inky cap mushrooms; hymenium lining gills; gills deliquesce into a black, inky liquid.
Coprinus urticicola. Fruit Rot of pear.
Corticium
Pileus resupinate, effuse; hymenium with one layer, cystidia lacking; spores hyaline.
Corticium vagum and other species with a thin lm of mycelium with short, broad cells
on substratum have been transferred to Pellicularia. Species with cystidia have been
placed in Peniophora. See also Corticium under Blights.
Butlerela eustacei (formerly Corticium centrifugum). Fisheye Fruit Rot
of apple, generally distributed. A dry, spongy rot often following scab.
ROTS 471
Corticium centrifugum (see Butlerela eustacei). Fisheye Fruit Rot of
apple, generally distributed.
Corticium fuciforme (see Laetisaria fuciformis). Pink Patch of turf, red
thread.
Corticium galactinum (see Scytinostroma galactinum). White Root Rot
of apple, also recorded on blackberry, dewberry, wineberry, peach and many
ornamentals baptisia, dogwood, holly, owering almond, owering plum,
iris, winter jasmine, kalmia, pearl bush, peony, spirea, sumac, viburnum and
white campion.
Corticium radiosum(see Vesiculomyces citrinus). White Butt Rot on sub-
alpine r in Colorado.
Laetisaria fuciformis (formerly Corticium fuciforme). Pink Patch of turf;
red thread. Grass is rst water-soaked, then dead, in isolated patches, 2 to
15 inches in diameter, with pinkish red gelatinous strands of the fungus mat-
ting the blades together and growing into coral red horns, 1/8 to 2 inches
long. These turn brittle, break into pieces to spread the pathogen. Velvet
bent grasses are more susceptible than colonial and creeping bents. Cad-
mium compounds will control if applied as protectants before the disease
appears.
Scytinostroma galactinum (formerly Corticium galactinum). White Root
Rot of apple, also recorded on blackberry, dewberry, wineberry, peach and
many ornamentals baptisia, dogwood, holly, owering almond, owering
plum, iris, winter jasmine, kalmia, pearl bush, peony, spirea, sumac, vibur-
num and white campion. The fungus also causes a root rot of white pine and
a decay of rs, affecting also western white cedar and spruce. The disease
starts at the collar or on larger roots and advances rapidly outward on small-
er roots. The collar may be girdled and killed while distal portions are still
alive. A dense weft of white mycelium covers roots and penetrates to wood,
causing the white rot. The disease is prevalent on lands recently cleared of
oaks.
Vesiculomyces citrinus (formerly Corticium radiosum). White Butt Rot
on subalpine r in Colorado.
Corynespora
Leaf Spots.
Corynespora cassiicola. Root Rot on soybean.
Cryptochaete
Basidocarp cartilaginous or coriaceous, erumpent, at rst tuberculiform; gloecystidia
yellowish or hyaline; cystidia present or lacking; spores hyaline, curved-cylindrical to
allantoid, smooth.
Cryptochaete (Corticium) polygonia (see Peniophora polygonia). White
Rot on aspen in Colorado.
Peniophora polygonia (formerly Cryptochaete (Corticium) polygonia).
White Rot on aspen in Colorado.
Cylindrocarpon
Conidia on sporodochia; spores with several cells, like Fusarium but more nearly cylin-
drical with rounded ends; cosmopolitan in soil, occasionally pathogenic.
Cylindrocarpon liriodendri. Root Rot of tulip poplar.
Cylindrocarpon destructans. Sometimes listed as cause of Scale-Tip Rot
of Easter lily in Pacic Northwest, but probably secondary. True cause of rot
unknown.
Cylindrocladium
Blights.
Cylindrocladium clavatum. Root Rot on Norfolk-island-pine.
Cylindrocladium crotalariae (Teleomorph, Calonectria crotalariae). Cy-
lindrocladium Black Rot on peanut, and Root Rot on tulip tree and kiwi.
Red crown rot; on soybean.
Cylindrocladium scoparium. Root Rot on peach and tulip-tree.
Cylindrocladium heptaseptatum. Postharvest Decay on leatherleaf
fern.
Cylindrocladium pauciramosum. Root and Crown Rot on heath.
Cylindrocladium parasiticum. Black Rot of hoary-tick clover and on
peanut.
Cylindrocladium pteridis. Postharvest Decay on leatherleaf fern.
ROTS 473
Cylindrocladium scorparium. Root Rot on pine, sweet gum and tulip-
tree.
Daedalea
Pileus dimidiate to caplike and stipitate; pores waved, mazelike or somewhat resembling
gills; without cystidia; hymenium labyrinthine.
Cerrena unicolor (formerly Daedalea unicolor). Heart Rot, Canker of
maples and other living hardwoods, including alder, ailanthus, amelanchier,
birch, chestnut and hackberry. Decayed wood is yellow at rst, later white
and soft. Conks are small, corky, often occuring in clusters, varying from
brown to gray.
Daedalea confragosa (see Daedaleopsis confragosa). White Mottled
Wound Rot of hardwoods, also on r.
Daedalea quercina. Brown Cubical Rot of dead timber; Heart Rot of
living trees in immediate vicinity of butt wounds, usually on oak, chest-
nut, sometimes on maple, birch and hickory. In advanced stages the wood
is reduced to a yellow-brown friable mass, with a tendency to break into
small cubes. Conks are corky and shelf-shaped, up to 7 inches wide, grayish
to almost black with smooth upper surface and cream to brownish under-
surface. Mouths are large, elongated, irregular. The conks are more or less
perennial.
Daedalea unicolor (see Cerrena unicolor). Heart Rot, Canker of maples
and other living hardwoods, including alder, ailanthus, amelanchier, birch,
chestnut and hackberry.
Daedaleopsis confragosa (formerly Daedalea confragosa). White Mottled
Wound Rot of hardwoods, also on r. This is a white soft rot, a slash
destroyer in eastern hardwood forests but sometimes on living trees, espe-
cially willows, near wounds. Annual leathery to rigid conks (sporophores)
are shelf-shaped, up to 6 inches wide, and may occasionally encircle a small,
dead stem. The upper surface is gray to brown, smooth, concentrically zoned.
Mouths of tubes on undersurface are elongated, wavy in outline.
Daldinia
Ascomycetes, Xylariaceae
Perithecia in a globoid to pulvinate, concentrically zoned stroma, carbonaceous to leath-
ery, 3 to 5 cm across; spores one-celled; dark.
Daldinia concentrica. Wood Rot of ash, beech, various hardwoods and
occasionally citrus. There is a supercial white rot on dead parts of living
trees. On English ash the decay is called calico wood and is strikingly marked
with irregular brown to black bands. Stroma containing perithecia are hemi-
spherical, black, carbonaceous.
Diaporthe
Blights.
Diaporthe phaseolorum. Sweetpotato Dry Rot. If diseased potatoes are
planted, the sprouts are affected, but the disease shows little in the eld. The
roots, infected at the stem end, continue to rot in storage. They are shrunken,
often mummied, covered with papillae, which are pycnidia under the skin
massed in a coal-black stroma. Optimum temperature for the fungus is 75
to 90F. Use cool storage.
Diaporthe citri. Phomopsis Stem End Rot, Melanose, general on cit-
rus; Stem Rot of mango. The rot on fruits is a leathery, pliable, buff to brown
area at the button end. The melanose is a supercial marking of fruits with
yellow or brown, scabby, waxy dots or crusts, on leaves, twigs and fruit,
often in streaks. On lemon trees, especially variety Eureka, there is a condi-
tion known as decorticosis or shell bark. The outer bark dies, loosens, peels
off in longitudinal strips. New bark forms below this, and the tree may recov-
er only to develop the disease again in 4 or 5 years. Some leaves and twigs
die; the fungus winters in dead wood.
Control. A single copper spray, bordeaux or a neutral copper, applied within
1 to 3 weeks after fruit is set, controls melanose. Copper applied in summer
induces excessive cork formation in the melanose lesions, a condition known
as star melanose. Applied early, it is noninjurious.
Diaporthe phaseolorum. Fruit Rot of pepper and tomato, also pod blight
of lima bean.

Blights.
Dichotomophthora
Cankers.
Dichotomophthora portulacae. Black Stem Rot on common purslane.
ROTS 475
Diplodia
Blights.
Diplodia natalensis (see Lasiodiplodia theobromae). Diplodia Collar and
Root Rot, Fruit Rot, Gummosis, general on citrus, sometimes peach,
mango and avocado.
Diplodia opuntia. Cladode Rot of cactus.
Diplodia phoenicum. Leaf and Stalk Rot of date palms, Fruit Rot. The
disease is sometimes fatal to transplanted offshoots. Leaves decay and die
prematurely; spores are produced in great abundance. Infection is through
wounds. Remove diseased tissue as far as possible and apply copper-lime
dust.
Diplodia pinastri. Collar Rot of pine.
Diplodia theobromae (see Lasiodiplodia theobromae). Sometimes consid-
ered a synonym of D. natalensis.
Diplodia tubericola (see Lasiodiplodia theobromae). Java Black Rot,
general on sweetpotatoes, especially in the South.
Diplodia zeae (see Stenocarpella maydis). Diplodia Corn Ear Rot, Root
and Stalk Rot, seedling blight.
Lasiodiplodia theobromae (formerly Diplodia natalensis). Diplodia Col-
lar and Root Rot; Fruit Rot, Gummosis, general on citrus, sometimes
peach, mango and avocado. On fruit, the rot resembles Phomopsis rot in
being a leathery pliable decay of the stem end. It can be prevented by spray-
ing with bordeaux mixture, adding 1% oil to check the increase in scale
insects after the copper kills entomogenous fungi keeping them in check.
The collar rot may girdle young trees and produce some gumming. Trees
affected with root rot seldom recover and should be removed.
Lasiodiplodia theobromae (formerly Diplodia theobromae). Sometimes
considered a synonym of D. natalensis but differentiated by pycnidia devel-
oped in a stroma instead of on a subiculum and by darker spores. Causing
rots of tropical fruits, stem-end rot of avocado and collar rot of peanuts. The
peanut rot appears in Georgia, Florida and Alabama. Runners and central
stem are invaded; they are brown at rst, then black with pycnidia.
Lasiodiplodia theobromae (formerly Diplodia tubericola). Java Black
Rot, general on sweetpotatoes, especially in the South. So named because
the rst diseased specimens came from Java; this is strictly a storage rot. The
inner part of the tuber is black and brittle; innumerable pycnidia are pro-
duced under the skin, giving it a pimply appearance. The potato is nally
mummied. Use care in handling so skins are not broken or bruised; cure
properly after harvest; have suitable temperature in the storage house.
Stenocarpella maydis (formerly Diplodia zeae). Diplodia Corn Ear Rot,
Root and Stalk Rot, seedling blight. This is one of several fungi commonly
causing ear rot in corn. The rot is dry, varying from a slight discoloration of
kernels to complete rotting of the ear. Seedlings and inner stalks have a dry,
brown decay. Another species (D. macrospora) is similar but less common,
found in more humid, warmer regions. The rot is greater in smutted plants.
Treat seed before planting with Spergon.
Diplodina
Leaf Spots.
Diplodina persicae. Fruit Rot of peach, found in Louisiana in 1952, affect-
ing stem and leaves as well as fruit. All varieties are susceptible.
Epicoccum
Leaf Spots.
Epicoccum nigrum. Postharvest Decay on cantaloupe.
Echinodontium
Hymenium in the form of teeth with spiny serrate margins; pileus caplike to crustose.
Echinodontium tinctorium. The Indian paint fungus causes Brown Strin-
gy Rot, Heartwood Rot of living conifers balsam r, hemlock, Engel-
mann spruce, larch, and Douglas-r chiey in the West, often with large
losses in forest stands. Light brown to tan spots are produced in heartwood
accompanied by small radial burrows resembling insect galleries. Rusty
streaks follow the grain. In older trees rot can extend entire length of heart-
wood and into roots. External signs of decay are hard, woody, hoof-shaped
perennial conks, the upper surface dull black, cracked, the undersurface gray,
ROTS 477
covered with coarse teeth, the interior rust are brick red with a pigment used
by the Indians for paint. Even one fruiting body is indicative of extensive
decay.
Polyporus (Favolus)
Pileus usually stipitate; lamellae forking irregularly to form elongate, rhomboidal pores.
Favolus alveolaris (see Polyporus mori). Heart Rot of hickory.
Polyporus mori (formerly Favolus alveolaris). Heart Rot of hickory.
Fomes
Pileus woody, perennial, with tubes in layers; common cause of wood decay. Spores
hyaline to brown to nearly black.
Fomes annosus (see Heterobasidion annosum). Heart Rot, Root and Butt
Rot, Spongy Sap Rot of conifers, sometimes hardwoods; also Root Rot
on juniper and rhododendron.
Fomes applanatus.
Ganoderma applanatum.
Fomes connatus. White Spongy Rot of heartwood of living hardwoods,
most prevalent on maples, especially red and sugar maples. Entrance is
through wounds or branch stubs, but fruiting is usually on basal stems or
scars. Conks appear annually but are perennial, small, less than 6 inches
wide, hoof-shaped, corky to woody, white to yellowish, the upper surface
covered with moss or algal growth. There is usually a limited area of decay.
Fomes everhartii (see Phellinus everhartii). Yellow Flaky Heart Rot of
living hardwoods, including birch and beech and especially oaks.
Fomes fomentarius. White Mottled Rot of birch, beech, poplar, maple,
and other hardwoods. This fungus mostly decays dead timber; sometimes
it attacks living trees. The wood is brownish, rm in early stages of decay,
but in advanced stages is yellowish white, soft, spongy, with narrow dark
zone lines and small radial cracks lled with yellow mycelium, giving a mot-
tled effect. Decay starts in upper part of the bole and progresses downward.
Conks are profuse on dead trees. They are hard, perennial, hoof-shaped, up
to 8 inches wide, with a smooth concentrically zoned upper surface, gray to
brown undersurface. The interior is brown, punky, with tubes encrusted with
white.
Fomes fraxinophilus (see Perenniporia fraxinophila). White Mottled Rot
of ash, a heartwood rot most common on white ash, also on green ash and
willow.
Fomes igniarius (see Phellinus igniarius). White Spongy Rot, White
Trunk Rot, Heart Rot, on a wide variety of hardwoods but not on conifers.
Fomes ofcinalis (Fomitopsis ofcinalis) (see Fomitopsis ofcinalis).
Brown Trunk Rot of conifers infecting heartwood of living larch and
other trees.
Fomes pini (Trametes pini) (see Phellinus pini). Red Ring Rot, white
pocket rot, of conifers, especially Douglas-r, larch, pine and spruce, causing
heavy forest losses.
Fomes pinicola (see Fomitopsis pinicola). Brown Crumbly Rot of many
conifers and some hardwoods maple, birch, beech, hickory, peach sually
on dead trees, occasionally in heartwood of living trees.
Fomes rimosus (see Phellinus robiniae). Heart Rot on locust.
Fomes robustus (see Phellinus robustus). Heart Rot of cacti and other
desert plants; of oak, r, juniper, in different strains.
Fomes roseus (see Fomitopsis rosea). Brown Pocket Rot, cubical rot of
heartwood of living conifers, particularly Douglas-r.
Fomitopsis ofcinalis (formerly Fomes ofcinalis (Fomitopsis ofcinalis)).
Brown Trunk Rot of conifers infecting heartwood of living larch and other
trees. Intensely white spore surface; very bitter, known as the quinine fungus.
Fomitopsis pinicola (formerly Fomes pinicola). Brown Crumbly Rot of
many conifers and some hardwoods maple, birch, beech, hickory, peach
ually on dead trees, occasionally in heartwood of living trees. Sporophores
are shelf- to hoof-shaped, 2 to 10 inches across, sometimes up to 2 feet, upper
surface gray to black, often with a red margin, underside white to yellow
when fresh.
Fomitopsis rosea (formerly Fomes roseus). Brown Pocket Rot, cubical rot
of heartwood of living conifers, particularly Douglas-r. Decay originates in
upper part of bole. Wood is yellow to reddish brown, soft, breaking into
irregular cubes. Woody bracket conks, up to 6 inches wide, have black tops
and rose undersurface. Infection is through dead branch stubs and broken
tree tops.
Heterobasidion annosum (formerly Fomes annosus). Heart Rot, Root
and Butt Rot, Spongy Sap Rot of conifers, sometimes hardwoods; also
ROTS 479
Root Rot on juniper and rhododendron. Infection is through wounds. Tis-
sue thin, mycelial felts are formed between bark and wood, which is pinkish
to violet in incipient states. In advanced stages white pockets are formed
in wood. Perennial conks are bracket-shaped to at layers, upper surface
zonate, light to dark grayish brown, undersurface beige with small pores.
Infection is sometimes through dead roots from mycelium growing through
soil, sometimes by spores washed by rain or carried by rodents.
Perenniporia fraxinophila (formerly Fomes fraxinophilus). White Mot-
tled Rot of ash, a heartwood rot most common on white ash, also on green
ash and willow. Conks are up to a foot wide, with dark, rough upper surface,
brownish underneath, appearing rst when wood has decayed only a short
distance. Infection is usually through branch stubs.
Phellinus everhartii (formerly Fomes everhartii). YellowFlaky Heart Rot
of living hardwoods, including birch and beech and especially oaks. Infec-
tion is usually limited to the lower trunk, and the aky character is because
the decay is more rapid between rays. There are narrow, dark brown zone
lines. Gnarled swellings on the trunk indicate sapwood invasion. The conks
are perennial, hard, woody, shelf-shaped, up to a foot wide, with the yellow-
brown upper surface becoming black, charred, rough, concentrically grooved
with age. The undersurface is reddish brown.
Phellinus ingiarius (formerly Fomes igniarius (Phellinus igniarius)). White
Spongy Rot, white trunk rot, heart rot, on a wide variety of hardwoods but
not on conifers. Aspen and birch are particularly susceptible. Decay is mostly
conned to heartwood, but in yellow birch living sapwood is killed, causing
cankers on the trunk. In an advanced stage the decay is soft, whitish, with
ne black lines running through it. The conks are perennial, hard, woody,
thick, usually hoof-shaped, up to 8 inches wide, the upper surface gray to
black, becoming rough and cracked with age; undersurface is brown and
the interior rusty brown with many layers of tubes, the oldest stuffed with
white. Infection is through branch stubs and open wounds. A single conk
may indicate 15 linear feet of rot in heartwood.
Phellinus pini (formerly Fomes pini (Trametes pini)). Red Ring Rot, white
pocket rot, of conifers, especially Douglas-r, larch, pine and spruce, caus-
ing heavy forest losses. Decay starts as a purplish or red discoloration of
the heartwood, but in an advanced stage there are many soft, white brous
pockets separated by sound wood. Sporophores vary from shelf- to bracket-
to hoof-shaped, averaging 4 to 8 inches across, rough gray to brownish black
with light brown margin on upper surface and gray to brown underneath.
Tube mouths are circular to irregular. On living trees conks are formed at
knots or branch stubs.
Phellinus robiniae (formerly Fomes rimosus) Heart Rot on locust.
Phellinus robustus (formerly Fomes robustus). Heart Rot of cacti and other
desert plants; of oak, r, juniper, in different strains. Context of sporophores
bright yellow-brown; spores hyaline.
Fusarium
Mycelium and spores generally bright in color. Macroconidia fusoid-curved, septate, on
branched conidia in slimy masses, sporodochia; smaller microconidia with one or two
cells; resting spores, chlamydospores, common. Teleomorph state when known usually
in Hypocreales, Nectria or Gibberella. Cause of many important rots, wilts, and yellows
diseases. Classication difcult, with different systems and synonyms, many forms and
races. (see Fig. 3.47)
Fusarium acuminatum. Fruit Rot of squash and pumpkin.
Fusarium avenaceum. Crown Rot of Eustoma. Associated with cereal dis-
eases, fruit and storage rots, but now included in F. roseum by many pathol-
ogists. Fruit Rot of squash and pumpkins.
Fusarium culmorum. Also on cereals, included in F. roseum by many
pathologists. Fruit Rot of squash and pumpkins, also Root and Basal Rot
of leak.
Fusarium equiseti. Fruit Rot of squash and pumpkin.
Fusarium moniliforme (Teleomorph, Gibberella fujikuroi). Ripe Rot of
gs, carried by the pollinating g wasp; Root, Stalk, Pink Kernel Rot of
corn. The rotted kernels are pink to reddish brown; the stalks have brown
lesions, may break over or ripen prematurely.
Figure 3.47 Forms of Fusarium. A septate macroconidia; B micoconidia in chains or a head; C clamydospores;
D sclerotium
ROTS 481
Fusarium oxysporum. Reported as causing a new disease of soybean in
Missouri and Iowa. Root rot, with rapid wilting and drying of leaves; most
severe on seedlings.
Fusarium oxysporum. Root Rot on apple and sage; tomato hypocotyl rot
on sugar pine, red and white rs; stem rot on zygocactus; and rot of stone
plant. This pathogen may also be seedborne and pathogenic on Douglas-r.
Root and crown rot; of leafy spurge.
Fusarium oxysporum. Iris Basal Rot on bulbous varieties of iris. Plants
fail to emerge, or turn yellow, wilt and die. Roots are few or none. The bulb
is infected at the base, which shrinks; the husk adheres rmly, sometimes
with a white or reddish mass of mycelium. The rot is more serious in warm
climates and on yellow rather then blue varieties. De Wit is very susceptible;
Wedgewood is quite resistant.
Control. Avoid bruising bulbs in digging; sort and discard diseased bulbs
right after digging; dry bulbs rapidly.
Fusarium oxysporum. Tulip Basal Rot. Leaves growing from diseased
bulbs turn red, wilt and die; roots are few or none. Bases of bulbs have
a rather rm rot with white or pink felty masses of spores. The diseased
area usually turns chalky. This is primarily a storage disease in bulb sheds
and warehouses.
Fusarium oxysporum f. sp. batatas. Sweetpotato Stem Rot, a wide-
spread eld disease, destroying more than 50% of plants in some elds. This
fungus also infects Jacquemontia. The stem rot is conspicuous about 2 weeks
after sprouts are set. Sprouts are yellow or dead, and the vines brown or
black, often split near the ground. Some plants develop new roots above
the decayed section and so survive. Sweetpotatoes from infected plants are
small, decayed at the stem end, with vascular tissues brown. The fungi winter
in stored roots and can live indenitely in soil. Varieties Big Stem Jersey,
Little Stem Jersey, Maryland Golden and Nancy Hall are very susceptible;
Porto Rico is intermediate; Southern Queen, Triumph, and Yellow Strassburg
are quite resistant.
Fusarium oxysporum f. sp. cepae. Bulb Rot, Basal Rot of onion, shallot
and garlic. In the eld there is progressive yellowing and dying back from
tips, the roots commonly turning pink and gradually decaying. The rot is
often associated with wounds of maggots and other insects. In storage the
rot is most active at room temperature or above.
Fusarium oxysporum f. sp. chrysanthemi. Fusarium Wilt of chrysanthe-
mum.
Fusarium oxysporum f. sp. gladioli (F. orthoceras var. gladioli). Fusari-
um Brown Rot, Yellows of gladiolus, a major disease in some sections.
Most infection takes place in the eld, but subsequent decay appears in
storage. Corm lesions are rst small, reddish brown, more often on lower
half of corm. They enlarge in storage to irregular to circular, sometimes
zonate brown areas, which do not infrequently advance until the whole
corm is a hard, dry, brownish black mummy. Infection comes from old
corms, the fungus penetrating through the basal plate and the center of the
new corm. The latter may be entirely decayed in storage, with the fungus
advancing from the center to the outside, causing brown to black surface
lesions.
Symptoms of yellows, a vascular disease, include bending of young leaf
stalks, cupping of leaf stalks in older plants, crooked ower stems, often
greener than normal and a curving of growth away from the side of the corm
showing rot. There is gradual yellowing and dying of foliage, starting with
the oldest leaves. Picardy and Spotlight varieties are particularly susceptible.
Nitrogenous fertilizers and manures, especially where phosphorus is low,
increase corm rot.
Control. Cure immediately after digging at 95F to develop wound periderm
and cuticle resistant to the fungus; use resistant varieties where possible or
a 3- to 4-year rotation.
Fusarium oxysporum f. sp. lilii. Basal Rot of lily, on bulbs, roots, stems
of garden and native lilies; Corm Rot of crocus, also on freesia and cactus
(Cereus). A chocolate rot at base of scales next to the basal plate progresses
until the scales fall away. The disease is more destructive to Madonna and
some other garden lilies; it is seldom a problem with Easter lilies grown in
the Northwest. Keep bulbs cool in storage, and plant in cool soil. Infection
comes from contaminated soil as well as diseased bulbs.
Fusarium oxysporum f. sp. narcissi (F. bulbigenum). Narcissus Basal
Rot, general on hardy varieties, rare on polyanthus varieties. Rot begins at
the root plate at base of bulbs and spreads through central portions rst,
extension of the rot being more rapid in affected scales than across to adja-
cent healthy scales. Rotted tissue is chocolate or purplish brown, the myceli-
um a delicate weft of white or pink threads. The rot is dry, spongy, with little
external evidence; it is primarily a storage or transit disease, but it may occur
in the eld late in the season. When lightly infected bulbs are planted, there
is no root development, and plants are stunted. Basal rot is spread in hot-
water treatment for nematodes. It is more prevalent where soil temperatures
ROTS 483
are above 65F and on large trumpet varieties. Golden Harvest is much more
susceptible than King Alfred.
Control. Discard all bulbs showing rot, or that are soft when pressed; if dis-
ease has occurred previously, plant in a new location.
Fusarium oxysporum f. sp. radicis-lycopersici. Root and Crown Rot of
tomato.
Fusarium oxysporum var. redolens. Root Lesions of pine.
Fusarium poae. Carnation Bud Rot, Silver Spike Disease of bluegrass.
The interior of carnation buds is brown or pink, decayed, moldy and often
infested with grass mites, which have introduced the spores. The disease
is favored by excessive dampness. Pick and destroy diseased buds; control
mites.
On bent grasses, fescues and especially Kentucky bluegrass, seed heads with-
er before they are fully expanded, appearing silvery. Seeds are aborted, and
in moist weather copious mycelium grows from decayed areas in culms. The
pathogen is disseminated and grass inoculated by the grass mite (Siteroptes
graminum). Burning over dead grass is a practical means of control.
Fusarium proliferatum. Root Rot and Stem Wilt on asparagus in CT.
Root Rot and Crown Rot of clovers and leafy spurge, Bulb Rot of onions.
Fusarium roseum. Peppermint Root and Rhizome Rot. Reported from
Oregon as part of a complex with Rhizoctonia solani and Pythium sp. Necrot-
ic lesions girdle rhizomes; new shoots damp-off. Fall-plowed mint gave
stronger stands. This pathogen also causes seedling stem rot on Douglas-r.
Fusarium cerealis. Stem Rot of carnation and cereals. Roots and stems of
cuttings and young plants rot; in older stock the diseased tissue turns brown-
ish red or crimson. Infection is only through injured, weak or old tissue.
Fusarium semitectum. Corky Dry Rot on cantaloupe.
Fusarium solani. Tuber Rot on caladium and Stem Rot on chrysanthe-
mum, Fraser r, Douglas-r, dieffenbachia (cutting rot), and sweetpotato
(root rot); shefera is susceptible with no symptoms. Root rot; this pathogen
also causes root rot of apple. Root and Crown Rot of leafy spurge.
Fusariumsolani (Teleomorph, Nectria haematococca). Stem Rot and Wilt
of Exacum.
Fusariumsolani f. sp. cucurbitae (Teleomorph, Hypomyces solani). Fusar-
ium Root Rot of cucurbits, primarily pumpkin and squash, occasionally
muskmelon, watermelon and cucumber. The fungus usually girdles the plant
at ground level with a soft dark decay, resulting in a striking wilt of the entire
vine. Fruits on the ground may be rotted and the fungus carried on seed to
infest clean soil. Do not plant cucurbits in land known to be contaminated.
Fusarium solani f. sp.phaseoli. Dry Root Rot of bean and lima bean, com-
mon but most important in New York, Idaho and other areas intensively
cropped for many years. Indenite reddish lesions or streaks on taproot and
subterranean stem turn dark brown to black. Lateral roots are reduced and
plants stunted. This is a late season disease favored by warm soil. The fun-
gus winters in crop refuse and soil and may be carried in dust on seed. The
best control is a long rotation between crops.
Fusarium solani f. sp. pisi. Root Rot on chick-pea, spruce, pine, r, and
hemlock.
Fusarium subgutinans. Collar Rot and Foliar Blight on Chinese ever-
green.
Fusarium sp. Root and Seed Rot of bird-of-paradise (Strelitizia), part of
a fungus complex. Controlled by treating seed in hot water, 135F for 30
minutes, and immediately cooling in cold water and treating planting medi-
um with methyl bromide or steam.
Fusarium sp. Dill Root Rot, Wilt, discovered in Ohio in 1949. Symptoms
include browning of roots, necrosis of vascular system, yellowing, wilting
and death. Young plants are most susceptible. Seed treatment did not give
satisfactory control.
Fusarium sp. Root Rot of sweet peas, reported as prevalent in Montana.
Plants turn yellow when in bloom with necrosis of vascular system which
leads to drying up of plant.
Gaeumannomyces
Gaeumannomyces graminis var. graminis. Blight of centipede grass and
Bermudagrass. Root rot; of St. Augustinegrass. Leaf yellowing and root mass
reduction; of Zoysia.
Ganoderma
Differing from Fomes in having spores truncated at one end and two-layered, the spines
of the brown endospore projecting into hyaline exospore. Sporophore has a hard crust,
formed by a layer of thick-walled, elongated cells.
ROTS 485
Ganoderma applanatum (Syn. Fomes applanatus). White Mottle Rot,
widely distributed on hardwoods, maple, beech, alder, acacia, birch, horse-
chestnut, hawthorn and hickory, and sometimes on conifers. The rot is ordi-
narily on dead timber, but the fungus can attack living trees through wounds
and destroy heartwood for a few feet. In early stages the wood is some-
what bleached, surrounded by a dark brown band. This shelf fungus is called
artists conk because the white undersurface immediately turns brown when
bruised and can be used for writing or etching pictures. The upper surface is
smooth, zoned, gray or gray-black; up to 2 feet wide.
Ganoderma curtisii (see Ganoderma lucidum). Perennial, with several lay-
ers of pores.
Ganoderma lucidum (formerly Ganoderma curtisii). Perennial, with sever-
al layers of pores.
Ganoderma lucidum (Syn. Polyporus lucidus). The varnish or lacquer fun-
gus causes Heart Rot of eastern hardwoods and conifers, especially hem-
lock, reported also on boxwood, hackberry, sassafras, maples and citrus. This
fungus may be an important facultative parasite on city shade trees. The rot is
white, spongy, with black spots scattered throughout. The conks are annual,
with a reddish, shiny, lacquered upper surface and a short, thick lateral stalk;
common on logs, stumps, standing or fallen trees.
Ganoderma zonatum. Butt Rot of queen palms, Florida.
Ganoderma zonatum. On mesquite, in Texas.
Greeneria
Greeneria uvicola. Bitter Rot of grape.
Gibberella
Blights.
Gibberella zeae (Anamorph, Fusarium graminearum). Corn Root Rot,
Stalk Rot, Ear Rot, also Fusarium Head Blight or Scab of cereals and
grasses. Corn is attacked at all ages, with both roots and kernels rotted. Coni-
dia are pinkish in mass; black perithecia are numerous on overwintered corn
stalks and residues. Hybrid corn with loose husks exposing the ear tip or vari-
eties with upright ears retaining water are more apt to be infected. Rotation
and clean plowing aid in control.
Gilbertella
Gilbertella persicaria. Fruit Rot on peach.
Gloeosporium
Anthracnose.
Colletotrichum gloeosporioides (formerly Gloeosporium foliicolum). (Glo-
merella cingulata). Fruit Rot on citrus fruits.
Gloeosporium foliicolum (see Colletotrichum gloeosporioides). (Glomerel-
la cingulata). Fruit Rot on citrus fruits.
Gloeotinia
Gloeotinia granigera (formerly Gloeotinia temulenta). Blind Seed Rot on
grass.
Gloeotinia temulenta (see Gloeotinia granigera). Blind Seed Rot on
grass.
Glomerella
Anthracnose.
Glomerella cingulata. Bitter Rot of apple and pear, Fruit Rot of peach,
also Stem Rot, Canker, Dieback of many fruits and ornamentals, Ripe
Rot of grapes. Bitter rot is a late season disease of apple, often destructive in
central and southern states. The fruits have light brown circular spots, which
gradually enlarge; they cover rotting esh, which has a bitter taste. Lesions
become concave and have concentric rings of pink to dark spore pustules
in sticky masses. Spores are splashed by rain or carried by ies and other
ROTS 487
insects. Eventually apples turn into dry, shriveled mummies, in which the
fungus overwinters and where the ascospore stage is produced. Large limbs
have oval, roughened, sunken cankers. The disease is favored by hot muggy
weather.
Apple varieties vary greatly in resistance, and some, like Yellow Newtown,
are resistant to the canker but susceptible to fruit rotting. Varieties somewhat
resistant include Delicious, Rome Beauty, Stayman Winesap, Winesap and
York Imperial. Ripe rot starts on grapes as they mature and gives a bitter
taste to the pulp. To control disease remove mummies from trees and prune
out dead twigs and cankers.
Glomerella cingulata var. vaccinii. Cranberry Bitter Rot, a eld and stor-
age rot. A soft brownish yellow discoloration develops on fruit late in the
season, most serious in a hot July and August.
Godronia
Apothecia coriacious, pitcher-shaped; spores liform, hyaline.
Godronia cassandrae (Fusicoccum putrefaciens). Cranberry End Rot,
general on cranberry, with the ascospore stage also found on dead branches
of leatherleaf (Cassandra). The rot appears late, often after picking and pack-
ing, and is enhanced by injuries during harvesting and screening. It starts at
either blossom or stem end of the berry; the fruit becomes soft and light-
colored.
Godronia cassandrae f. sp. vaccinii. On blueberry.
Guignardia
Blotch.
Botryosphaeria vaccinii (formerly Guignardia vaccinii). Cranberry Early
Rot, Scald, Blast, general on cranberry and sometimes on huckleberry. All
aerial plant parts are attacked, but the disease is more destructive to the fruit.
Young fruit may blast and shrivel, but more often rot starts as a light-colored
soft spot when fruit is half grown. The berry mummies, turns black and is
covered with small pycnidia. Leaves have reddish brown spots, sometimes
drop prematurely.
Figure 3.48 Black Rot of Grapes
Guignardia bidwellii. Black Rot of grapes, widespread, principal cause
of failure of European grapes in eastern United States, causing more loss
than all other grape diseases combined. All parts of the vine are attacked.
On leaves, reddish brown dead spots are sprinkled with black pycnidia. Rot
starts on half-grown fruit as a pale spot, soon turning brown and involving
the entire berry, which shrivels into a black wrinkled mummy, dropping or
remaining in the cluster (see Fig. 3.48). Some berries shatter if attacked ear-
ly. Ovoid conidia and sometimes microconidia (spermatia) are formed on
leaves, berries and canes. Ascospores are produced in overwintered mum-
mied berries. Primary infection in spring comes from either spore form.
Thecaphora deformans. Seed Smut of lupine.
Control. If mildew is also a problem, use a xed copper. Cultivate in early
spring so as to cover old mummies with soil and so eliminate that source of
inoculum.
Guignardia vaccinii (see Botryosphaeria vaccinii). Cranberry Early Rot,
Scald, Blast, general on cranberry and sometimes on huckleberry.
Helicobasidium
Basidiomycetes, Septobasidiales
ROTS 489
An exposed cottony hymenium or fruiting layer; basidia transversely septate; spores
coiled like a watch spring.
Helicobasidium corticioides. Brown Pocket Rot on subalpine r, in Col-
orado.
Helicobasidium brebissonii (Anamorph, Rhizoctonia crocorum). Violet
Root Rot of potato, sweetpotato, asparagus, beet, carrot and some orna-
mentals ash, catalpa, chinaberry, crocus, elm, mulberry, parthenocissus
and western soapberry. The fungus invades roots from the soil, turning them
reddish or violet. The disease is conned to underground parts unless con-
tinuously wet weather allows the reddish-purple mycelium to grow up the
stem. Small, darker sclerotia are embedded in this purplish mat, which turns
brown with age.
Helminthosporium
Blights.
Bipolaris cactivora (formerly Helminthosporium cactivorum). Stem Rot
of cacti, Basal or Top Rot of seedling cacti, which turn into a shrunken
brown mummy covered with spores. Initial symptoms are yellow lesions;
rotting may be complete in 2 to 4 days.
Exserohilum turcicum (formerly Helminthosporium turcicum). Crown
Rot of sweet corn, Leaf Blight.
Blights.
Helminthosporium cactivorum (see Bipolaris cactivora). Stem Rot of
cacti, Basal or Top Rot of seedling cacti.
Helminthosporium sesami. Stem Rot on sesame in Texas.
Helminthosporium turcicum (see Exserohilum turcicum). Crown Rot of
sweet corn, Leaf Blight.
Hericium
Fleshy, branched or unbranched, with subulate spines long and pendant; spores spheri-
cal or subspherical, staining blue with iodine. Like Hydnum but sporophore formed on
wood, not on the ground.
Hericium erinaceus (Syn. Hydnum erinaceus), hedgehog fungus. White
Heart Rot, occasional on living oak, maple and other trees. The soft white
spongy rot may entirely decompose the tissue, leaving large hollows lined
with yellowish mycelium. Sporophores are annual; soft, white, browning
with age, globular with a hairy top and long slender teeth on the lower sur-
face.
Grandinia (Hyphodontia)
Grandinia granulosa (formerly Hyphodontia aspera). Root and Butt Rot
of spruce.
Hypholoma
Basidimycetes, Agaricales
Margin of cap with a curtainlike veil; stipe with incomplete or vanishing ring; spores
purple.
Hypholoma sublateritium. Root Rot of currant.
Idriella
Mycelium hyaline to brown; conidophores brown, simple, nonseptate, narrowed above,
with prominent spore scars; conidia (sympodulospores) lunate to falcate, with pointed
ends, produced in clusters near apex of the conidiophore; aleuriospores brown, several-
celled.
Idriella lunata. Root Rot on strawberry.
Inonotus
Inonotus tomentosus (Syn. Polyporus tomentosa). Root Rot of spruce.
ROTS 491
Irpex
Resupinate, effused-reexed, or shelf-like; younger parts of hymenophore are poroid;
with increasing age produce attened teeth.
Irpex lacteus. Wood Rot and Decline of apple.
Gliocladium (Isaria)
Conidiophores equally distributed on a synnema, erect fascicle of hyphae; conidia hya-
line, one-celled, ovoid; some species in insects.
Gliocladium roseum (formerly Isaria clonostachoides). Isaria Rot of
tomato. Fruits are partly covered with cottony mycelium, white turning pink
or orange and becoming granular, but rot remaining rm. Reported from
around Washington, D.C.
Isaria clonostachoides (see Gliocladium roseum). Isaria Rot of tomato.
Kluyveromyces
Ascomycetes, Saccharomycetaceae
Kluyveromyces marxianus var. marxianus. Soft Rot of onion caused by
a true yeast on bulbs.
Lasiodiplodia
Lasiodiplodia theobromae. Collar Rot on peanut.
Lentinus
Gills are notched or serrate at edge, decurrent, stipe often lateral or lacking cap, tough-
eshy to leathery; spores white.
Lentinus lepideus (see Neolentinus lepideus). Scaly Cap, causing a brown
cubial rot of coniferous wood and sometimes decaying heartwood of living
pines.
Lentinus tigrinus. Sapwood Rot, white mottled butt rot of living hard-
woods, commonly associated with re scars and one of the most impor-
tant decay fungi in the Mississippi Delta. Fruiting body is white with cap
depressed in center, more or less covered with blackish brown hairy scales,
rarely developing on living trees.
Neolentinus lepideus (formerly Lentinus lepideus). Scaly Cap, causing
a brown cubial rot of coniferous wood and sometimes decaying heartwood
of living pines.
Lenzites
Pores elongated radially to resemble gills; pileus shelike; woolly and zonate above.
Gloeophyllum sepiarium (formerly Lenzites saepiaria). Timber Rot,
Brown Pocket Rot, usually of dead sapwood, occasionally a heart rot,
rarely on living trees. This is the common destroyer of coniferous slash; it is
found on telephone poles and other timber. Fruiting bodies are long narrow
shelves coming from cracks, the upper surface a yellow red to dark reddish
brown.
Lenzites betulina. Heart Rot of birch and cypress.
Lenzites saepiaria (see Gloeophyllum sepiarum). Timber Rot, Brown
Pocket Rot, usually of dead sapwood, occasionally a heart rot, rarely on
living trees.
Leptosphaeria
Blights.
Leptosphaeria korrae. Root and Crown Rot of turf grasses (necrotic ring
spot).
ROTS 493
Macrophoma
Cankers.
Macrophoma sp. Fruit Rot of grape.
Macrophomina
Spores hyaline, one-celled, in pycnidia.
Macrophomina phaseolina (Anamorph, Sclerotium or Rhizoctonia batati-
cola). Charcoal Rot, Ashy Stem Blight, on many plants in warm climates
and sometimes in temperate zones. The name for the sterile stage comes
from sweetpotato, and the term charcoal rot is used because the interior of
the potato becomes jet black.
The fungus lives in the soil, is particularly prevalent in warm soils and attacks
roots and stems of a varied list of hosts, including bean, lima bean, soybean,
beet, corn, cowpea, cabbage, eggplant, garlic, gourds, pepper, strawberry and
watermelon; also chrysanthemum, dahlia, garden mallow, mountain-laurel,
marigold and zinnia. In most cases the pycnidial stage is not formed. The
mycelium spreads through the soil, and very small black sclerotia are formed
in great abundance on or in lower stems and roots. On beans, black sunken
cankers appear just below the cotylendonary node, and the lesion may extend
up the stem, ashy gray in the center. Stems may break over, or the growing
point may be killed. In sweetpotatoes the disease is a storage rot, the tissue
becoming a dark red-brown with the outer zone black from the formation of
myriads of sclerotia. The decay is spongy, then hard, mummied. The fungus
is spread in irrigation water, crop debris, imported soil and on seed.
Control. Use bean seed grown in western disease-free regions. Keep plants
growing vigorously with proper food and water; practice general sanitation.
Macrophomina phaseolina. Charcoal Rot on soybean, sunower, Ama-
ranthus, Euphorbia spp., Ipomea, Sonchus and Tidestrominia; root rot on
caper spurge.
Magnaporthe
This is characterized by the production of perithecial ascocarps produced in
a stroma of fungal and substrate tissues or directly from somatic hyphae on
the substrate.
Magnaporthe poae. Summer patch and large brown patch on grass.
Melanconium
Leaf Spots.
Greeneria uvicola (formerly Melanconium fuligineum). Bitter Rot of
grapes, widespread but especially serious on Muscadine grapes in Georgia.
Decayed berry pulp has a bitter taste; up to 30% of fruit is reduced to dry,
hollow shells. Spray with bordeaux mixture three times at 14-day intervals
beginning after fruit is set. The later sprays for black rot should control bitter
rot.
Melanconium fuligineum (see Greeneria uvicola). Bitter Rot of grapes,
widespread but especially serious on Muscadine grapes in Georgia.
Mycocentrospora
Mycocentrospora acerina. Dry Rot of carrot.
Monilinia
Blights.
Monilinia fructicola (Syn. Sclerotinia fructicola). Brown Rot of stone
fruits, blossom blight, general on peach, plum and cherry, also on apricot,
almond, beach, plum, Japanese quince, and, rarely, apple and pear. The fun-
gus is distinct from the species in Europe (Sclerotinia fructigena) causing
brown rot of stone fruits and also a serious apple rot. In the United States,
brown rot is our most destructive stone-fruit disease, causing an annual
peach loss of over $5 million. Monilina fructicola is the usual causative
agent east of the Rocky Mountains; M. laxa causes a similar rot and blossom
blight on the Pacic Coast. See also

Blights.
ROTS 495
Flowers turn brown prematurely, rot in moist weather; the calyx cup is black-
ened, and the discoloration may extend down into the pedicels. Infrequently
there is a leaf and twig blight; cankers are formed on the larger limbs, with
exudation of gum. The fruit rot is the familiar stage seen in any backyard with
a fruit tree and usually in baskets of peaches, plums or cherries purchased for
preserving and held over to the next day. The rot starts as a small, circular
brown spot but spreads rapidly to take in the entire fruit, with the rotted sur-
face covered with gray to light brown spore tufts or cusions (sporodochia),
sometimes in concentric rings (see Fig. 3.49). Conidia are formed in chains
on the sporodochia. The fruit nally shrinks and mummies and either falls
to the ground or remains clinging to the tree.
The fungus and decayed tissue together form a stroma that acts as a scle-
rotium; in spring, if the mummy has been kept moist and partially or wholly
covered with soil, cup-shaped brown apothecia are produced. Primary infec-
tion is from ascospores, forcibly ejected and carried up to blossoms by air
currents or from a new crop of conidia formed on mummies hanging on
trees. Secondary infection is from conidia wind-borne from blossom to blos-
som and later from fruit to fruit. Entrance is often through wounds made by
the plum curculio, oriental fruit moth and other insects. Rotting and conidial
production continue after picking.
The rot is favored by wet weather, conidia germinating only in a lm of wa-
ter. Acid soil is said to increase apothecial production from mummies on the
ground. In a normal season reduction from blossom blight is not important
because some thinning is advantageous; but if blossom blight is not prevent-
ed, inoculum is provided for the fruit rot that causes such enormous losses.
Control. Sanitary measures are important. In the small garden rake up and
burn or bury deeply the fallen mummies; pick mummies from trees; cut out
twigs showing gum; in summer remove infected fruit before conidia form.
Standard control has been wettable sulfur sprays or sulfur dust, applied:
every 3 or 4 days during bloom to control blossom blight; when shucks are
falling; 2 or 3 weeks after shuck fall; and 2 to 4 weeks before fruit ripens.
In some instances the newer organic fungicides are preferred to sulfur, and
sometimes they are used with it. Control of the plum curculio is very impor-
tant. For one or two trees in a home garden one of the all-purpose fruit sprays
or dusts now available under various trade names may be satisfactory. Con-
sult your county agent for the schedule right for your locality.
Monilinia laxa (Syn. Sclerotinia laxa). Brown Rot, green and ripe fruit rot,
blossom blight, on almond, apple, apricot, cherry, peach, plum, pear, nec-
Figure 3.49 Brown Rot of Plums
tarine, quince and Japanese quince in Washington, Oregon and California;
also reported from Wisconsin and Michigan. Although this disease is similar
to that caused by M. fructicola, the blossom and twig blight phase is more
important than the brown rot. Sulfur, which can be used in later sprays for
most stone fruits, may injure apricots.
Monilinia oxycocci (Syn. Sclerotinia oxycocci). Cranberry Hard Rot, Tip
Blight in Pacic Northwest and Wisconsin. Young growing tips wilt and
ROTS 497
dry just before blossoming; grayish spores tufts are formed on tips. Fruit is
attacked through blossoms or wounds. The berries are yellowish white, rm,
leathery, cottony inside, turning dark and mummifying late in the season.
The disease is too erratic to justify cost of regular spraying; clean harvest
will prevent overwintering.
Monilinia urnula (Syn. Sclerotinia vaccinii-corymbosi). Blueberry Brown
Rot, Mummy Berry, Twig Blight of high bush blueberry, similar to hard
rot of cranberry. Varieties differ in susceptibility, with June and Rancocas
often showing severe primary infection.
Monilochaetes
Conidiophores dark, erect, slender, usually simple; septate; conidia hyaline or becoming
pigmented in age, borne singly at apex or produced in chains under conditions of high
humidity.
Monilochaetes infuscans. Root Rot of weed species of genus Ipomoea.
Monosporascus
Ascomycetes, Sordariales
Monosporascus cannonballus. Root Rot on cantaloupe and watermelon.
Mucor
Mycelium profusely developed. Sporangiophores erect, simple or branched, all branches
terminated by sporangia which are globose to pyriform with a columella and thin wall;
gametangia essentially alike, suspensors without denite outgrowths; hyaline chlamy-
dospores sometimes formed.
Mucor mucedo. Postharvest Rot of tomato.
Mucor piriformis. Postharvest Rot of tomato. Fruit Rot of cherry.
Mucor racemosus. Storage Rot of sweetpotato, occasional after chilling;
Fruit Rot of citrus. Control with low temperatures and dry atmosphere in
the storage house.
Mycoleptodiscus
Mycoleptodiscus terrestris. Root and Stem Rot of soybean and birdsfoot
trefoil.
Myrothecium
Leaf Spots.
Myrothecium roridum. Ring Rot of tomato, Crown Rot of snapdragon
and pansy. Crowns of greenhouse snapdragons appear water-soaked, then
covered with a thin white mycelium and numerous black sporodochia. Irreg-
ular brown spots on tomato fruits are surrounded by slight depressions. Also
causes root rot of red clover and alfalfa.
Myrothecium sp. On Bells of Ireland, causing crown necrosis. Stems are
girdled at ground level; tops wilt; basal branches die.
Nematospora
Ascomycetes, Saccharomycetales
This is a yeast or budding fungus, following after insect injury; asci, with 8 to 16 spores,
derived directly from vegetative mycelial cells; spores elongate, fusiform to needle-
shaped, agellate.
Nematospora coryli. Yeast Spot of soybean, Dry Rot of pomegranate,
citrus, Pod Spot of pepper, bean and soybean, Cloudy Spot of toma-
to, Kernel Spot of pecan. The yeast is almost always associated with
plant bug injury. The western leaf-footed plant bug carries the fungus from
pomegranate to citrus. On pomegranates depressed light spots in esh around
seeds are followed by general browning and collapse. In citrus, the juice sacs
just inside the rind dry out with a brownish to reddish stain. Cloudy spot
on tomato fruit is associated with pumpkin bugs and leaf-footed plant bugs.
Brown areas are formed on pecan kernels.
Nematospora phaseoli. Yeast Spot of lima bean, a seed disease, destruc-
tive from Maryland southward. Infection follows puncture of pods by the
southern green stinkbug and possibly other insects. The seed lesions are dark
brown, sunken, wrinkled.
ROTS 499
Neurospora
Ascomycetes, Sordariales
Perithecia ask-shaped, membranous; ascospores dark, one-celled with gelatinous coat-
ing; conidial stage monilioid
Neurospora sitophila. Ripe Rot of pear. The fungus is the same one caus-
ing pink bakery mold on bread. There is a luxuriant pink growth over fruit;
conidia are formed in chains.
Nigrospora
All hyphae more or less creeping, hyaline; conidiophores short, dark, cells somewhat
inated; conidia black, one-celled, situated on a attened, hyaline vesicle at top of the
conidiophore.
Nigrospora oryzae. Ripe Fruit Rot of tomato, Nigrospora Cob Rot of
corn. Corn cobs are shredded, with the pith completely disintegrated; kernels
are lled with masses of black spores. Corn on poor soil is more susceptible;
stalks break over at any point. Rapid drying checks infection of seed corn.
Olpidium
Chytridiomycetes, Spizellomycetales
Endobiotic, living in host cells or tissues, living or dead.
Olpidium brassicae. Sometimes found in outer cells of rootlets of cabbage
and other crucifers, tomato, lettuce and other plants, producing zoosporangia
and resting spores in the cells. The effect on the host is usually merely a slight
unthriftiness. Olpidium has been found associated with a disease of lettuce,
Big Vein, now thought due to a virus.
Omphalia
Basidomycetes, Agaricales
Gills decurrent, cap sunken in center, somewhat funnel-shaped; central car-
tilaginous stem; spores white.
Marasmiellus pigmentatus (formerly Omphalia pigmentata). Omphalia
tralucida. Decline Disease of date palms. Growth is retarded; roots decay;
leaves die prematurely; fruit is worthless. Deglet Noor variety is most sus-
ceptible. Select thrifty offshoots from healthy plants for new date gardens.
Soil can be treated with carbon disulde, as for Armillaria rot.
Omphalia pigmentata (Omphalia tralucida) (see Marasmiellus pigmenta-
tus). Decline Disease of date palms.
Oospora (Geotrichum)
Slender branched or unbranched mycelium breaking up into ellipsoidal or spherical hya-
line or light-colored conidia called oidia.
Geotrichum citriaurantii (formerly Oospora citri-aurantii). Sour Rot of
citrus. This is a soft, putrid slimy rot of fruit, mostly of stored lemons, where
it is spread by contact. The mycelium forms a thin, compact, somewhat wrin-
kled layer over the surface. Fruities help to spread the spores. Fruit should
be stored as short a time as possible and frequent inspections made during
storage.
Geotrichum candidum (formerly Oospora lactis). Sour Rot, Watery
Fruit Rot of tomato, common in transit and market, especially on fruit from
the South. There is a velvety or granular coating over the surface or a uffy
growth along the margin of cracks, and a disagreeable odor and avor. The
rot is common on ripe fruit touching the ground, occasional on green fruit.
The fungus is a weak parasite, entering through wounds.
Oospora citri-aurantii (see Geotrichum citriaurantii). Sour Rot of citrus.
Oospora lacti (see Geotrichum candidum). Sour Rot, Watery Fruit Rot
of tomato, common in transit and market, especially on fruit from the South.
Ophiosphaerella
Ophiosphaerella sp. Large Brown Patch Rot on bermudagrass and creep-
ing bentgrass.
ROTS 501
Paecilomyces
Conidiophores and branches more divergent than in Penicilllium; conidia (phialospores)
in dry basipetal chains, one-celled, ovoid to fusoid, hyaline.
Paecilomyces buxi (see Sesquicillium buxi). Root Rot and Decline on box-
wood.
Sesquicillium buxi (formerly Paecilomyces buxi). Root Rot and Decline
on boxwood.
Pellicularia
Blights.
Thanatephorus cucumeris (Syn. Pellicularia lamentosa) (considered by
some Botryobasidium). Rhizoctoniose, Black Scurf of potatoes, stem
canker and soil rot of beans (see under Blights for Web Blight of beans and
other plants); Rhizoctonia Dry Rot Canker of Beets, Crown and Crater Rot
of Carrots; Rhizoctonia Disease of Celery, Crucifers, Cucurbits; Bottom Rot
of Lettuce; Damping-Off of Pepper and Eggplant; Root Rot of Onion; Root
and Basal Stem Rot of Pea; Crown Rot of Rhubarb. The sterile state of this
fungus, Rhizoctonia solani, was rst named in 1858 in a German textbook
and is still the most familiar term for a fungus with many pathogenic strains
causing many types of diseases.
Any cook has seen signs of the pathogen on potato tubers small brown to
black hard ecks, sclerotia, on the skin. They look like particles of dirt but
do not scrub off when potatoes are washed. There may be only one or two
sclerotia, or they may nearly cover the whole surface of the tuber. When such
potatoes are planted, the growing point may be killed. Some sprouts renew
growth after being girdled, which may be repeated until they die. Larger
plants have stems decayed just below the soil line, interrupting the downward
transfer of food and resulting in a cluster of green or reddish aerial tubers.
Roots may be killed back extensively. Most of the tubers are small, often
with a brown jelly rot at the stem end.
Under moist conditions a white cobwebby weft of mycelium is formed at the
base of potato stems, and the basidial stage is produced as a powdery crust on
this weft. The fungus winters as mycelium or sclerotia in soil or tubers. The
mycelium can grow saprophytically long distances in the soil independent
of any plant. Infection is favored by cool temperatures; the disease is most
serious in wet seasons on heavy soils. The average yearly loss for the country
is about 10 million bushels, 2 to 3%, but individual losses may be from 5 to
50%. For control use healthy tubers for seed.
Thanatephorus cucumeris (Syn. Pellicularia lamentosa (Anamorph, Rhi-
zoctonia solani)). Brown Patch of turf, Root and Leaf Rot of lawn
grasses, wheat grass, bentgrass, fescues, ryegrass, Kentucky bluegrass (infre-
quently on Canada bluegrass), St. Augustine grass, and Bermuda grass.
Brown or blackish patches on the turf resemble sunscald or chinch bug
injury. The areas are roughly circular, from an inch to 3 feet across, some-
times up to 20 feet. The fungus works outward with a smoke ring of
grayish black mycelium at the advancing margin. The leaves are rst water-
soaked, black, then collapsed, dry and light brown, but the roots are seldom
killed. The disease develops most rapidly during warm humid periods and
with an excess of nitrogen.
Thanatephorus cucumeris (Syn. Pellicularia lamentosa (Anamorph, Rhi-
zoctonia solani)). Root and Stem Rot, Damping-Off of ornamentals. In
wet weather cobwebby mycelium develops on lower portions of stems; the
lower leaves rot and upper portions of plants wilt and die. Seedlings and old-
er plants so rotted include Aconitum, abelia, Achillea, Ageratum, aster, arti-
choke, begonia, calendula, campanula, carnation, endive, dahlia, delphini-
um, geranium, iris, lettuce, lupine, orchids, platycodon, poinsettia, salsify,
sunower and tulip. For control avoid excessive use of manure.
Penicillium
Cankers.
Various species cause blue, green, occasionally pink molds, including the common blue-
green mold on jellies. Some produce antibiotics, Penicillium notatum being the one used
for production of penicillin.
Gliocladium roseum (formerly Penicillium roseum). Fruit Rot of citrus and
of dates. A pink mold, found on lemons but not oranges.
Gliocladium vermoeseni (formerly Penicillium vermoeseni). Bud Rot of
palms. The terminal bud is killed and base of leaf stalks rotted. Affected trees
of very susceptible Washingtonia lifera should be replaced with resistant
Washingtonia robusta, Mexican fan palm. Also
Cankers.
ROTS 503
Penicillium aurantiogriseum. Crown Rot of asparagus, a seedling dis-
ease recently prevalent in Washington, following freezing injury. Bright blue
spore masses appear on diseased crowns. Protect seedlings for winter by
slight hilling in fall; avoid mechanical injury in harvesting; prevent drying
out of crowns between digging and replanting.
Penicilliumdigitatum. Green Mold of citrus fruit, Clove Rot of garlic. On
lemons and other citrus, olive-green powdery spore masses, forming a dust
cloud when disturbed, cover fruit except for a band of white mycelium out-
side the green area. Garlic plants are yellow and stunted. Avoid injury in har-
vesting and packing. Commercial growers use chemicals in the wash water
to prevent decay.
Penicillium expansum. Blue Mold Rot of many fruits; Soft Rot of apple,
pear, avocado, pomegranate, Japanese persimmon, quince and feijoa. The
decay on avocados is slow, and often the affected portions can be trimmed
off. This fungus causes 80 to 90% of the decay of storage apples. The rot-
ted portions are light-colored, soft, watery, with a disagreeable moldy taste
and odor. A few rotted apples spoil all the others in a container. Use great
care in harvesting and grading to avoid wounds; keep temperature as low as
possible.
Penicillium gladioli. Blue Mold Rot, Penicillium Dry Rot of gladiolus,
also found in imported bulbs scilla, tritonia (montbretia). This is a storage
rot. Light to dark brown sunken lesions appear on any part of corms with
border of the decayed area water-soaked and greenish. Small grayish sclero-
tia are formed, and under moist conditions masses of blue mold. Dry rapidly
after harvest, 80F for 10 to 14 days, then store at low temperature; avoid
wounds and bruises; sort before planting.
Penicillium italicum. Blue Contact Mold of citrus, Fruit Rot. The mold
is blue in the older portion but powdery white at margins. It spreads readily
from fruit to fruit by contact, through uninjured skin.
Penicillium roseum (see Gliocladium roseum). Fruit Rot of citrus and of
dates. A pink mold, found on lemons but not oranges.
Penicillium vermoeseni (see Gliocladium vermoeseni). Bud Rot of palms.
Peniophora
Like Corticium but with cystidia.
Chaetoderma luna (formerly Peniophora luna). Brown Rot in lodgepole
pine, Rocky Mountain area.
Peniophora luna (see Chaetoderma luna). Brown Rot in lodgepole pine,
Rocky Mountain area.
Pestalotia
Blights.
Pestalotia longisetula. Root, Stolon, and Petiole Rot on strawberry.
Phaeoacromonium
Blights.
Phaeoacremonium aleophilum. Decline of grape.
Phaeoacremonium chlamydosporum. Decline of grape.
Phaeoacremonium inatipes. Decline of grape.
Phialophora
Conidiophores dark, short, single or clustered; phialides broader near middle, tapering
toward ends, producing conidia endogenously, spores subhyaline to dark, one-celled.
Phialophora malorum. Storage Rot of apples.
Phlebia
Basidiocarp effuse, typically monomitic; spores even in general outline, hyaline or pale
in color, typically nonamyloid.
Phlebia chrysocrea. Heart Rot on oak.
ROTS 505
Pholiota
Basidomycetes, Agaricales
Spores ochre yellow to rusty brown; gills attached to stipe, which has an
annulus but no cup at the base.
Pholiota adiposa. Brown Mottled Heart Rot of maple and other living
hardwoods basswood, birches, poplars and more rarely conifers. The wood
has brown mottled streaks. The sporophores are formed in clusters on trunks
and stumps m ushrooms with yellow central stems and caps, sticky yellow
slightly scaly upper surface, yellow to brown gills.
Phoma
Blackleg.
Phoma apiicola. Phoma Root Rot of celery, occasionally serious, espe-
cially in Golden Self Blanching, also on carrot, parsnip, parsley and caraway.
The disease appears rst in the seedbed, a black rot of the crown or base of
leafstalks. Plants are stunted, outer leaves or entire plant killed, falling over
as roots rot off. Spores are produced in tendrils from black pycnidia and
spread in rains and irrigation water. Use clean seed, grown in California,
where the disease is rare; sterilize seedbed soil or use a fresh location.
Phoma betae (Teleomorph, Pleospora betae). Phoma Rot of beets, causing
black root of seedlings, necrotic streaks on seedstalks, brown spots on old
leaves and rot of eshy roots. The fungus is seed-borne and winters in roots
carried over for seed production and in debris. Crop rotation is essential.
Phoma destructiva. Phoma Rot of tomato, pepper, nearly general, espe-
cially in the South, but not in North Central States. Small, irregular dark spots
appear on leaves in great numbers; zonate markings are similar to those of
early blight. Severely infected leaves turn yellow, wither. Fruit spots in eld
are small, 1/8 inch, slightly depressed, with numerous tiny black pycnidia.
After harvest, spots enlarge to 1/2 to 1 1/2 inches and become black, leathery,
with minute pustules. The fungus winters in decaying refuse in soil; seedbed
infection is common, and the disease reaches the eld via infected seedlings.
Masses of spores produced on leaves are washed to fruits by rain or spread
by workers and are distributed during harvesting and packing.
Control. Locate seedbeds away from land that has previously grown toma-
toes; spray as for early blight; do not harvest when wet.
Phoma macdonaldii (Teleomorph, Leptosphaeria lindquistii). Stem Rot
on sunower.
Phoma terrestris on sweet corn.
Phoma sp. Crown and Root Rot on bugleweed.
Phomopsis
Blights.
Phomopsis amygdali. Fruit Rot on almond.
Phomopsis mali. Fruit and Core Rot (Postharvest) on apple.
Phomopsis vaccinii. Fruit Rot on blueberry.
Phomopsis sp. Fruit Rot on peach.
Phymatotrichopsis (Phymatotrichum)
Deuteromycetes, Hypohomycetes
Conidiophores stout with inated tips bearing loose heads of conidia; spores hyaline;
one-celled, produced on surface of soil.
Phymatotrichopsis omnivors (formerly Phymatotrichum omnivorum).
Texas Root Rot, Phymatotrichum Root Rot, Cotton Root Rot. This
is the most destructive plant disease in Texas, a limiting factor in gardening
and crop production. It occurs in the Red River counties of Oklahoma, the
southwestern half of Arizona, the southeastern edge of Nevada and Califor-
nia, the southeastern corner of Arkansas and Utah, the northwestern corner
of Louisiana and in most of Texas except the Panhandle.
The list of susceptible plants owers, vegetables, fruits, eld crops and trees
is much longer than that of plants resistant to this omnivorous fungus,
so aptly named. At least 1700 plant species are attacked, more than by any
other known pathogen. Because of the wide host range and destructiveness,
the economic losses are enormous, $100 million a year in Texas alone, with
perhaps $50 million in adjacent states.
Crops that either are resistant or escape the disease are the cereals and grass-
es, annuals grown in winter only, and sweet alyssum, amaranth, sweet basil,
ROTS 507
beauty-berry, bee-balm, collinsia, diosma, calceolaria, calla lily, California-
poppy, candytuft, canna, chicory, cranberry, cucumber, currant, cyclamen,
daffodil, dahoon, deutzia, dill, fenner, fern, staghorn, foxglove, freesia, gold-
entuft, mustang grape, gypsophila, hackberry, hoarhound, hyacinth, iris, lily,
nigella, marsh-marigold, mignonette, mints, mimulus, muskmelon, mustard,
nasturtium, oak, osage-orange, oxalis, Indian paint-brush, palms, pansy,
petunia, phlox, Chinese pink, pitcher-plant, pomegranate, poppy, portulaca,
primrose, pumpkin, red-cedar, sage, scarlet-brush, snapdragon, snowdrop,
stock, strawberry, strawower, tuberose, valerian, verbena, violet, wallow-
er, wandering jew, water cress, watermelon, yaupon, yucca and zinnia.
Phymatotrichum root rot occurs from July until frost. It kills plants in more
or less circular spots, ranging from a few yards to an acre or more. Death
may come within a few days of rst wilt symptoms, and just preceding the
wilt plants actually run a fever, with a higher than normal temperature. If
plants next to the wilted ones are pulled out, these apparently healthy plants
will often be found to be covered with yellow to buff mats of mycelium,
and under moist conditions spore mats appear on the surface of the soil
around diseased plants. Such mats are 2 to 12 inches in diameter, rst snow
white and cottony, later tan and powdery from spores produced in quantities.
The fungus spreads through the soil by means of rhizomorphs, smooth, dark
brown strands. The rate of spread may be 2 to 8 feet a month in an alfalfa
eld, 5 to 30 feet a season in a cotton eld, or around fruit trees.
Sclerotia are formed along the mycelial strands. They are small, roundish,
light at rst, then dark and warty. The fungus winters either as sclerotia
in soil, persisting several years in the absence of live hosts, or as dormant
mycelium in living roots. The disease is most common and severe on heavy,
alkaline soils. Abundant organic material reduces rot by favoring antagonis-
tic soil saprophytes.
Control. In ornamental plantings replace diseased plants with some of those
given in the resistant list. Monocotyledons are generally resistant. In locat-
ing new orchards, make sure that root rot has not been present previously by
growing an indicator crop of cotton for a year. Grow immune crops in rota-
tion with susceptible crops, and grow susceptible annuals in winter rather
than summer. Try heavy manuring.
Ammonium sulfate can sometimes save a valuable ornamental tree or shrubs
already infected with root rot. Prune back the top, make a circular ridge about
the plant at the edge of the branch spread, and work ammonium sulfate into
the soil within the ridge then ll the basin with water to a depth of 4 inch-
es. The chemical treatment and watering is repeated in 5 to 10 days, then
no more chemical the same season. Follow through with frequent water-
ing.
Phymatotrichum omnivorum (see Phymatotrichopsis omnivors). Texas
Root Rot, Phymatotrichum Root Rot, Cotton Root Rot.
Physalospora
Cankers.
Botryosphaeria stevensii (formerly Physalospora mutila). Black Rot of
apple, in the West, similar to disease by P. obtusa in the East.
Botryosphaeria obtusa (formerly Physalospora obtusa). Black Rot of
apple, New York Apple Tree Canker, Frog-Eye Leaf Spot, general on
apple and crabapple, from Atlantic Coast to the Great Plains; also widespread
on pear, mountain-ash, peach, quince, currant and various woody species.
The fungus, in its anamorph state (Sphaeropsis malorum), was rst reported
as causing apple rot in 1879.
The lesions start as small brown spots, frequently at a wormhole, but they
darken and turn black as they expand. There is usually one lesion to an apple,
often at the calyx end, with concentric zones of black and brown, and minute
black pycnidia. The rot eventually takes in the whole fruit, which is shriveled
and wrinkled and nally mummies. The pycnidia are black, carbonaceous,
and may contrain three types of spores large one-celled brown spores, large
hyaline spores, and two-celled colored spores. Perithecia, sometimes formed
in cankers or on twigs, apparently play little part in the life history, the fungus
wintering as dormant mycelium or in the pycnidial state. Conidia, entering
through wounds, start primary infection in spring on leaves with the small
frog-eye leaf spots.
Control. Use the same spray schedule as for apple scab, starting with the
petal-fall application. Clean up mummied apples; avoid bruising; cut out
cankers.
Botryosphaeria rhodina (formerly Physalospora rhodina). Diplodia Rot
of citrus, g, rubber-tree and pear, possibly apple. The conidial stage is
a Diplodia, probably D. natalensis, with dark, two-celled spores.
Physalospora mutila (see Botryosphaeria stevensii). Black Rot of apple,
in the West, similar to disease by P. obtusa in the East.
ROTS 509
Physalospora obtusa (see Botryosphaeria obtusa). Black Rot of apple,
New York Apple Tree Canker, Frog-Eye Leaf Spot, general on apple
and crabapple, from Atlantic Coast to the Great Plains; also widespread on
pear, mountain-ash, peach, quince, currant and various woody species.
Physalospora rhodina (see Botryosphaeria rhodina). Diplodia Rot of cit-
rus, g, rubber-tree and pear, possibly apple.
Phytophthora
Blights.
Phytophthora cactorum. Stem Rot, Foot Rot of lily, Photinia, tulip,
Hydrastis, blue laceower, babys breath, Centaurea, peony, clarkia, rhubarb
and tomato; leather rot of strawberries; collar rot of dogwood, walnut, apple
and pear; crown rot of Euonymus and strawberry; root rot of boxwood and
vinca, and crown rot of peach and Cannan r; kernel and shuck rot of pecan.
With foot rot, lilies suddenly fall over, wilt and die; the lower part of the stem
is shrunken. Plant only healthy bulbs and where the disease has not occurred
previously.
Strawberry leather rot occurs when berries come in contact with soil, start-
ing with a brown rotted area on green fruit and a discoloration of vascular
bundles. Ripe fruit has a bitter taste. Crown rot of rhubarb starts with slightly
sunken lesions at base of petiole, which enlarge until the entire leaf collaps-
es. Spraying crowns with bordeaux mixture is helpful. Start new beds with
healthy plants. Collar rot on English walnut is a bark disease starting below
the ground with irregular dark brown or black cankers and soft, spongy areas
at the crown, a black uid in cambial cavities. Trees are stunted, with sparse
yellow-green top growth. There may be an unusually heavy crop of nuts, but
the tree dies the next season. Grow walnuts grafted on Persian or Paradox
rootstocks. See under Cankers for symptoms on apple and dogwood.
Stem rot and wilt of snapdragon starts with water-soaked lesions on the stem;
these turn yellow, brown, enlarge to girdle the stem; plant wilts. Sterilize soil
before planting.
Phytophthora capsici. One of the species causing buckeye rot of tomato.
See under Blights for pepper rot and blight.
Phytophthora cinnamomi. Avocado Root Rot, Pine Little Leaf, Collar
Rot of hardwoods and conifers, seedling root rot, on more than 100 hosts,
including rs, cedars, cypress, juniper, Japanese umbrella tree, larch, pine,
spruce, arborvitae, heaths, heather, azalea, Heuchera, cranberry, highbrush
blueberry, rhododendron, camellia, birch, western swordfern, manzanita,
walnut, oak, locust, yew, venus-ytrap and gold-dust plant. In conifers root
rot is dry with resin ow; needles gradually lose color. Infected tissue of
hardwoods turns reddish brown except in black walnut, where it is black;
seedlings die. The disease is aggravated in pine by poor aeration and low
fertility.
Root rot is the most serious avocado disease in California, present also in
Florida and Texas. It occurs on soils with poor drainage, excess moisture
being necessary. As the roots rot, leaves become light-colored and wilt even
if soil is moist; trees decline over a period of years. The fungus can be spread
with seed if fruit is allowed to lie on the ground. Treat suspected seed with
hot water, 120 to 125F for 30 minutes; use nursery stock grown in fumigat-
ed soil; prevent movement of soil water from infested areas; plant on well-
drained soil; water trees individually to avoid excess moisture.
Phytophthora citricola. Root Rot of pine, hemlock and Fraser r seedlings
in Christmas tree plantings; also fruit rot of avocado.
Phytophthora citrophthora. Root and Crown Rot of Penstemon.
Phytophthora citrophthora. Brown Rot, Gummosis, Foot Rot of citrus.
Masses of amber gum break out from the trunk near crown; the bark is killed
above and below ground; foliage turns yellow; trees may die. The disease is
prevalent where excess water stands around the tree after irrigation or where
there is poor drainage. Brown rot of fruit is a decay with no visible surface
mold, except in moist air, but a slightly rancid, penetrating odor. Lemons and
oranges may be affected on the tree, on branches near the ground, and there
is much loss in storage. The fungus lives in the soil; spores are splashed
up in rainy weather and are spread in the washing tank. Lemons are most
susceptible to gummosis, then lime, pumelo, grapefruit, sweet orange and
nally sour and trifoliate oranges. The latter two are used as fairly resistant
understocks.
Control. Plant susceptible trees high, with lateral roots barely covered;
expose the root crown of infected trees with a basin 6 inches deep and 4
feet across. Once a year cover crown and lower trunk with bordeaux paste.
To control fruit rot, spray ground and lower branches, up to 3 feet, with
bordeaux mixture just before rains begin. If fumigation is to be practiced,
substitute a copper-zinc-lime spray for the bordeaux.
Phytophthora colocasiae. Root Rot of ginseng.
ROTS 511
Phytophthora cryptogea. Collar Rot of rhododendron, China aster, mari-
gold, gloxinia and zinnia; root, crown, and stem rot on watercress, juniper,
African daisy, chicory, beet, globe thistle, lettuce, parsley, peach, sage,
spruce, and ice plant; stem rot on sunower, pink rot of potato. Stems and
roots appear water-soaked, then black from a soft rot. Sterilize soil.
Phytophthora cryptogea var. richardiae. Root Rot of calla. The feeder
roots rot from tips back to rhizomes, leaving the epidermis a hollow tuber.
New roots sent out from the rhizome rot in turn. Leaves turn yellow and drop,
starting with outer leaves; plants do not ower, or the tips of blossoms turn
brown. Rot in the rhizome is dry and spongy, not wet and slimy. Clean old
rhizomes thoroughly; cut out rotted spots. Grow in sterilized pots rather than
benches.
Phytophthora drechsleri. Root Rot on r, basil, blackberry, columbine,
juniper, sage, linden, pine, and spruce. Sometimes associated with tomato
buckeye rot, basal decay of sugar beets, tuber rot of potato, root rot of saf-
ower.
Phytophthora erythroseptica. Pink Watery Rot of potato, Rot of calla
lily and golden calla, Crown and Root Rot of wild rice. The rot starts
at stem end of potatoes; affected tissues exude water under pressure. When
tubers are cut, esh turns pink or red, then black. The fungus can exist in soil
4 years.
Phytophthora fragariae. Strawberry Red Stele Disease, Brown Core
Rot, a very serious strawberry disease, rst noticed in Illinois in 1930, now
widespread in northern strawberry sections and in California. A strain of
this pathogen causes root rot of loganberry. The fungus attacks roots only,
destroying ne feeding roots rst, then invading the central cylinder, stele,
which turns dark red. New spring leaves on badly affected plants are small,
bluish, have short petioles; large leaves from the previous season dry up;
little or no fruit is produced; plants die in the rst dry period or are stunt-
ed.
The fungus is most active in cold, wet soil, in rainy periods in late fall, win-
ter, and early spring except when ground is frozen. Zoospores produced on
roots are spread by water; resting spores formed in the red stele carry the
pathogen in a dormant state through the heat of summer. There are at least
three physiological races, and once the fungus infests a eld it is worthless
for strawberries for 10 years.
Control. Buy clean, certied plants. Aberdeen and Stelemaster varieties are
resistant; Temple, Sparkle, Fairland, Redcrop, and Pathnder, fairly so.
Phytophthora lateralis. Cypress Root Rot on Lawson cypress (Chamae-
cyparis lawsoniana) often called Port Orford cedar, and Hinoki cypress
(C. obtusa), killing thousands of trees in Oregon nurseries and landscape
plantings. It is found on juniper and azalea in North Carolina. It is also
reported on mountain-laurel and Photinia in NC and on cypress from Wash-
ington and apparently native to the Northwest. The fungus enters through
the roots and spreads to lower part of main trunk killing the tissues. Blue
cypress changes to purple, green, nally tan and dies. The color changes take
several months in cool, damp weather, only 2 or 3 weeks in hot, dry weather.
There is no practical chemical control, and Lawson cypress seems to be
incompatible with resistant rootstocks. Grow disease-free propagating stock
in new soil. Avoid large plantings of Lawson cypress such as windbreaks or
hedges. Remove and destroy infected plants, getting the entire root system.
Phytophthora megasperma. Root Rot, occasional on cabbage, cauliow-
er, brussel sprouts, carrot, artichoke, stock, citrus, soybean and wallower.
Diseased plants wilt suddenly; leaves turn red to purple; underground stems
and roots rot. The disease is more prevalent in winter plantings in California
and in low, poorly drained areas. Level ground properly before planting to
avoid waterlogged spots. Root and crown rot on peach.
Phytophthora megasperma f. sp. glycinea. Root and Stem Rot on soy-
bean.
Phytophthora nicotianae var. nicotianae. Crown Rot, Root Rot, and
Stem Canker on annel bush.
Phytophthora nicotianae var. parasitica Syn. Phytophthora parasitica
(P. terrestris). Brown Rot of citrus, in Florida; Buckeye Rot of tomato,
also on lily roselle, sempervivum, potato (tuber rot), zebra plant (stem rot),
sage (root rot), and Christmas cactus (root rot). The disease appears on the
lowest tomato fruits, where water stands after rains. The lesions have con-
centric narrow dark brown bands alternating with wide light brown bands.
The decay is rapid and the internal tissue semi-watery, though the exterior
is rm. Control by staking tomatoes; avoid poorly drained soil or plant on
ridges. This species is often present with P. citrophthora in cases of citrus
foot or collar rot.
Phytophthora nicotianae var. parasitica. Crown Rot on petunia and poin-
settia (stem rot).
Phytophthora palmivora. Palm Bud Rot, Leaf Drop, Wilt of coconut,
Washingtonia, and queen palm, root and crown rot of mango; also root rot
on English ivy. The fungus is an omnivorous tropical species, presumably the
ROTS 513
one causing stem rot of dieffenbachia and peperomia. It has been prevented
in nurseries by using cuttings from healthy plants in pasteurized soil.
Phytophthora parasitica var. nicotianae (Syn. P. nicotianae var. parasiti-
ca). Root Rot on pine.
Phytophthora porri. Head Rot on cabbage.
Phytophthora sojae (Syn. P. megasperma f. sp. glycinea). Root and Stem
Rot of soybean, a relatively new disease reported from Illinois, Indiana, Mis-
souri, North and South Carolina and Ohio. Serious in cool rainy weather,
causing pre- and post-emergence damping-off.
Phytophthora syringae. Root Rot on shore juniper and Photinia.
Phytophthora torulosum. Root Rot and Damping-off on soybean.
Plectospira
Oomycetes, Saprolegniales
Sporangium with much inated branching; swarm spores are formed in basal portion
and cut out into a single row in an elongate lamentous apical portion, which acts as an
exit tube. Swarm spores encyst at the mouth as in Aphanomyces. Oogonium terminal or
intercalary, accompanied by up to 65 antheridia.
Plectospira myriandra. Rootlet Necrosis on tomato. The fungus is weak-
ly parasitic on roots.
Plenodomus
Pycnidia coriaceous or carbonaceous, more or less sclerotoid. Condiophores obsolete or
none; conidia one-celled, hyaline.
Plenodomus destruens. Foot Rot of sweetpotato, one of the more impor-
tant eld diseases and sometimes a storage rot. The base of the stem turns
brown from just under the soil surface to 4 or 5 inches above; leaves turn yel-
low and drop off; vines wilt unless adventitious roots are put out; pycnidia
are numerous. The root has a rm brown rot, not affecting the whole potato
but enough to make it worthless for food. The fungus winters in old plant
refuse but not in soil. Use clean seed potatoes; rotate crops. This fungus also
infects Jacquemontia.
Pleospora
Leaf Spots.
Pleospora herbarum. Fruit Rot of tomato. A rm dark rot develops in fruit
after picking, starting from infections through cracks near stem end of fruit.
Progress is most rapid at 65 to 70F and is checked by storage at 45F.
Pleurotus
Stipe off center or lacking; cap sometimes inverted; gills more or less eshy and separa-
ble into two layers, edges acute; spores white.
Hypsizygus ulmarius (formerly Pleurotus ulmarius). Brown Heart Rot
sapwood wound rot of elm, maple, and other living hardwoods. Rot start-
ing in heartwood may extend into sapwood; infected wood separates along
annual rings. Annual sporophores have a long excentric stalk, and white to
yellow to brown smooth upper surface. They issue from crotches and pruning
wounds.
Pleurotus ostreatus, oyster cap. White Flaky Sapwood Rot of maple and
other hardwoods, sometimes on living trees. A light-colored decay is sur-
rounded by a narrow brown zone. Fleshy annual conks are shelving, sessile,
or with a short, stout excentric stalk. The upper surface is smooth, white or
grayish, gills extending onto the stalk, an edible fungus. Infection is through
open wounds. Wood Rot of grape.
Pleurotus ulmarius (see Hypsizygus ulmarius). Brown Heart Rot sap-
wood wound rot of elm, maple, and other living hardwoods.
Polyporus
Pileus tough, thick, with a stipe, or as a shelf; pores rounded, small, tubes crowded.
Dichomitus squalens (formerly Polyporus anceps). Red Ray Rot on west-
ern conifers, causing heart rot of living trees but benecial as a cause of rapid
decay of slash in forests. Fruiting bodies rarely develop on living trees.
Ganoderma lucidum (formerly Polyporus lucidus). Root Rot on redbud.
ROTS 515
Inonotus dryadeus (formerly Polyporus dryadeus). White Root Rot, occa-
sional in oaks and conifers in the West. Roots are killed; tree dies. Decayed
wood is white to cream; bark is loosened and shredded.
Inonotus hispidus (formerly Polyporus hispidus). White Spongy Heart
Rot of living trees of black ash, oak, maple and birch; does not decay dead
trees. Heartwood in upper portion of trunk is reduced to soft spongy yellow
or white mass. Shelf sporophores, up to 10 inches wide, have dark brown,
coarse, velvety to hairy upper surface and golden brown undersurface, turn-
ing dark brown with age. They are formed at branch stubs, frost cracks, or
trunk cankers.
Inonotus tomentosus (formerly Polyporus tomentosus var. circinatus Syn.
Inonotus circinatus). Root Rot of sand pine.
Laetiporus sulphureus (formerly Polyporus sulphureus Syn. Laetiporus
sulphureus), sulphur fungus. Red Brown Heart Rot, Brown Cubical Rot
in heartwood of maple and other living hardwoods and conifers, widespread
on oak, balsam, Douglas-r and spruce. The annual, shelike fruiting bodies
are most conspicuous soft, eshy, moist when fresh, with bright orange-red
upper surface and brilliant yellow underneath, formed in overlapping clus-
ters. When old they are hard, brittle, dirty white (see Fig. 3.50). Infection is
through dead branch stubs and wounds.
Phaeolus schweinitzii (formerly Polyporus schweinitzii) Root Rot on pine.
Phellinus gilvus (formerly Polyporus gilvus). White Sapwood Rot, preva-
lent on dead trees, occasional on living trees. Small, annual, yellow to red,
brown leathery to corky sporophores, developed in profusion.
Pitoporus betulinus (formerly Polyporus betulinus Syn. Piptoporus betuli-
nus). Brown Cubical Rot of dead or dying gray and paper birches. Conks
have smooth grayish upper surface with incurved margin.
Polyporus abietinus Syn. Hirshioporus abietinus (see Trichaptum abiet-
inum). Pitted Sap Rot, Hollow Pocket, White Pocket Rot on r.
Polyporus anceps (see Dichomitus squalens). Red Ray Rot on western
conifers.
Polyporus balsameus (see Postia balsamea). Balsam Butt Rot of living
balsam r, eastern hem lock, northern white-cedar, western red-cedar, also
prevalent on dead trees.
Polyporus betulinus Syn. Piptoporus betulinus (see Piptoporus betulinus).
Brown Cubical Rot of dead or dying gray and paper birches.
Polyporus dryadeus (see Inonotus dryadeus). White Root Rot, occasional
in oaks and conifers in the West.
Figure 3.50 Shelf Fungus (Laetiporus sulphureus) on Oak
Polyporus gilvus (see Phellinus gilvus). White Sapwood Rot, prevalent
on dead trees, occasional on living trees.
Polyporus hispidus (see Inonotus hispidus). White Spongy Heart Rot of
living trees of black ash, oak, maple and birch; does not decay dead trees.
Polyporus lucidus (see Ganoderma lucidum). Root Rot on redbud.
Polyporus pargamenus. White Pocket Rot of dead sapwood in eastern
United States but sometimes on living maple and other hardwoods.
Polyporus sulphureus Syn. Laetiporus sulphureus (see Laetiporus sul-
phureus), sulphur fungus. Red Brown Heart Rot, Brown Cubical Rot in
ROTS 517
heartwood of maple and other living hardwoods and conifers, widespread on
oak, balsam, Douglas-r and spruce.
Polyporus schweinitzii (see Phaeolus schweinitzii). Root Rot on pine.
Polyporus squamosus. White Mottled Heart Rot on maple, buckeye,
birch and occasional on living trees near wounds. Conks are annual, eshy,
white to dingy yellow with a short, thick lateral stalk, upper surface with
broad appressed scales, up to 18 inches wide.
Polyporus tomentosus var. circinatus Syn. Inonotus circinatus (see Inono-
tus tomentosus). Root Rot of sand pine.
Polyporus versicolor Syn. Coriolus versicolor (see Trametes versicolor),
rainbow conk. Sapwood Rot.
Postia balsamea (formerly Polyporus balsameus). Balsam Butt Rot of
living balsam r, eastern hemlock, northern white-cedar, western red-cedar,
also prevalent on dead trees. Advanced decay is brown, brittle, breaking into
large cubes, easily crushed to a clay-colored powder. In living trees the rot
column is usually only 3 or 4 feet from ground. Sporophores are shelving,
up to 2 inches wide, with pale brown upper surface with concentric zones,
white underneath.
Trametes versicolor (formerly Polyporus versicolor Syn. Coriolus versi-
color), rainbow conk. Sapwood Rot. This is the most common fungus on
hardwood slash in woods and sometimes on conifers. The rot is soft, white
spongy. Heartwood of living catalpa may be decayed, the fungus entering
through wounds and dead branches. The conks are thin, tough, leathery,
annual, up to 2 inches wide with a hairy or velvety surface multicolored
white, yellow, brown, gray, and black. The undersurface is yellow or white.
This pathogen also causes wood decay and decline of apple and has been
reported as Trametes versicolor.
Trichaptum abietinum (formerly Polyporus abietinus Syn. Hirshioporus
abietinus). Pitted Sap Rot, Hollow Pocket, White Pocket Rot on r.
May attack dead sapwood in wounds of living trees.
Poria
Pileus resupinate, thin, membranous; tubes wartlike, separate.
Junghuhnia luteoalba (formerly Poria luteoalba). Brown Rot of conifer-
ous wood.
Perenniporia subacida (formerly Poria prunicola). White Rot of cherry
and other Prunus spp.
Phellinus laevigatus (formerly Poria laevigata). Red Mottle Rot on
Prunus spp.
Phellinus weirii (formerly Poria weirii). Douglas-Fir Root Rot. The dis-
ease is most destructive to trees 70 to 150 years old, which are killed in
groups. The fungus can persist in dead roots for a century. Less susceptible
conifers should be planted with judicious cutting of infected stands.
Poria cocos (see Wolporia cocos). Root Rot on roots of various trees,
especially pine, in southeastern United States.
Poria luteoalba (see Junghuhnia luteoalba). Brown Rot of coniferous
wood.
Poria prunicola (see Perenniporia subacida). White Rot of cherry and oth-
er Prunus spp.
Poria laevigata (see Phellinus laevigatus). Red Mottle Rot, on Prunus spp.
Poria subacida. Feather Rot, Spongy Root Rot, String Butt Rot of
living conifers and dead hardwood. Decay rarely extends more than 6 to 10
feet in the trunk. Irregular pockets run together forming masses of white
bers; annual rings separate readily. Sporophores are white to straw-colored
to cinnamon-buff crusts forming sheets several feet long on underside of
fallen trunks or on underside of root crotches or exposed roots of living trees.
Poria weirii (see Phellinus weirii). Douglas-Fir Root Rot.
Wolporia cocos (formerly Poria cocos). Root Rot on roots of various
trees, especially pine, in southeastern United States. Huge sclerotia, weigh-
ing up to 2 pounds, are formed; this stage is known as Pachyma cocos.
Pyrenochaeta
Blights.
Phoma terrestris (formerly Pyrenochaeta terrestris). Pink Root of onions,
widespread on onions, garlic and shallot; also on grasses. Roots of affect-
ed plants shrivel and turn pink. New roots replacing the old are infected in
turn; plants are stunted, bulbs small. The fungus persists indenitely in the
soil and is distributed on onion sets and transplants. Yellow Bermuda is the
most resistant of commercial onion varieties. The green Beltsville Bunching
onion, Nebuka strain of Welsh onion, Evergreen variety of shallot, and leaks
ROTS 519
and chives are resistant. In Arizona, Granex gives a better yield than other
onions despite pink root.
Pyrenochaeta lycopersici. Root Rot on tomato.
Pyrenochaeta terrestris (see Phoma terrestris). Pink Root of onions, wide-
spread on onions, garlic and shallot; also on grasses.
Pythium
Wall of sporangium smooth; discharging swarm spores in imperfectly formed state into
thin-walled vesicle, which later ruptures to allow spores to escape. Sporangia terminal
or intercalary. Species live in moist soil causing damping-off and root rots.
Pythiumacanthicum; P. myriotylum; P. periplocum. Causing rot of water-
melon fruit.
Pythiumaphanidermatum. Leak, Root Rot. Damping-off of muskmelon,
cucumber, squash, also papaya, bean, radish, spinach, sugar beet, guayule,
caper spurge, and ice plant. There is a watery decay with a yellow brown
liquid leaking out when fruit is pressed. Lesions are brown and wrinkled.
The fungus lives in the soil; primary infection is in the eld, secondary from
contact in transit or storage. Sort carefully before packing. Refrigerate at 45
to 50F in transit.
Pythium aristosporum. Root Rot of bean.
Pythium arrhenomanes. Root Rot on tomato, broadleaf signalgrass, large
crabgrass, barnyardgrass, nutsedge, goosegrass, itchgrass and johnsongrass.
Pythium carolinianum. Root and Stem Rot of parrotfeather (Myrio-
phyllum).
Pythium catenulatum. Root Rot of bean.
Pythium debaryanum. Damping-Off of seedlings. Watery Leak of pota-
toes. Leak starts as a brown discoloration around a wound and soon spreads
to include the whole potato, which is soft, easily crushed, and drips a brown
liquid with the slightest pressure. Entrance to the tuber is usually through
harvest wounds. Pythium hyphae grow through the soil in great profusion
and can enter seedlings through either stomata or unbroken epidermis. See
Damping-Off, for rot of seedlings.
Pythium dissotocum. Root Rot of bean, and spinach.
Pythium irregulare. Associated with Melon Root Rot and Fruit Rots of
Figure 3.51 Pythium Blackleg on Geranium
other cucurbits in cool weather and Seed Decay of corn. Root Rot and
Crown Rots of clovers and basil.
Pythium mastophorum. Damping-off on celery.
Pythium myriotylum. Root Rot on tomato.
Pythium paroecandrum. Stem, Crown and Root Rot on lupine.
Pythium polymastum. Bottom Rot and Damping-off on cabbage.
Pythium splendens. On Chinese evergreen, peperomia, and philodendron.
Pythiumultimum. Fruit Rot of muskmelon, often with luxuriant white fun-
gus growth; Damping-Off, Root Rot of many seedlings in greenhouse and
ROTS 521
eld. Root Rot and Crown Rot of clovers. Crown Rot of impatiens. Root
Rot of kiwi.
Pythium uncinulatum. Stunt and Leaf Yellowing on lettuce.
Pythiumspp. Most soils contain several species of Pythium ready to perform
at optimum moisture and temperature. Exact determination is not always
practical. Nematode wounds often dispose plants to rot. Diseases include
African-violet rot, aloe root rot, black rot of orchids, begonia root rot, coleus
black leg, geranium cutting rot (see Fig. 3.51), bean and parsley root rot,
rhubarb crown rot, mottle necrosis of sweetpotato, and other rots. Some
plants can be treated with hot water, 115F for 30 minutes. Sterilize soil
before use; avoid excessive watering. See Damping-Off, for seedling rots.
Rhizina
Ascomycetes, Pezizales
Cup-shaped apothecia with rhizoids underneath; asci operculate, opening with a lid,
eight-spored; spores fusoid, spindle-shaped, paraphyses present.
Rhizina undulata. Seedling Root Rot, Damping-Off. Coniferous seed-
lings in the Pacic Northwest are sometimes killed in isolated circular patch-
es 2 to 4 feet in diameter, particularly in burned areas. Infected roots are
matted together with white mycelium. More or less resinous annual fructi-
cations are formed on the ground. They are irregular, an undulating brown
upper surface with narrow white margin, 2 to 3 inches across. There is no
control, but the disease is minor.
Rhizoctonia
Sclerotial form of Pellicularia, Corticium, Macrophomina, and Helicobasidium. Young
mycelium colorless; branches constricted at points of origin from main axis; older
mycelium colored, wefts of brownish yellow to brown strands, organized into dense
groups of hyphae, sclerotia, made up of short, irregular, angular or somewhat barrel-
shaped cells (see Fig. 3.52).
Rhizoctonia sp. Postharvest Decay on leatherleaf fern.
Rhizoctonia bataticola. Charcoal Rot.
Macrophomina phaseoli
Rhizoctonia crocorum. Violet Root Rot.
Helicobasidium purpureum.
Figure 3.52 Forms of Rhizoctonia solani . A young mycelium, constricted at branches; B loosely formed angular
to barrel-shaped colored cells; C section through sclerotium formed from aggregation of cells in B; D basidium
and spores of Pellicularia, teleomorph state of R. solani
Rhizoctonia solani. Black Scurf of potatoes, Brown Patch of turf.
Pellicularia lamentosa Root Rot and Crown Rot of clovers..

Rhizoctonia solani. Root and Stem Rot of poinsettia and other ornamen-
tal plants, including basil, begonia, camellia, calla, carnation, chrysanthe-
mum, coleus, cornower, geranium, gloxinia, impatiens, lily, pansy, pothos,
peperomia, primrose, ragweed, rosemary, caper spurge, and sainfoin (crown
rot). Although Pythium ourishes best in the low oxygen content of poor-
ly drained soils, Rhizoctonia, causing similar root rots, is serious in well-
drained soils. On poinsettia, dark brown lesions at or above soil level are
often covered with brown mycelium; the leaves turn yellow and drop, the
roots rot, and the plant dies. Web Blight and Stem Canker on alfalfa.
Damping-off and Bottom Rot on Cichorum (endive, escarole and witloof
chicory) and Nicotiana.
Rhizoctonia tuliparum. Gray Bulb Rot of tulips, in northeastern and
Pacic states. The most conspicuous sign of this disease is a bare patch
in spring where tulip shoots should be showing. Occasionally an infected
bulb will produce some above-ground growth, but the plants are slow and
often wither and die before owering. Bulbs rot from the top down; myceli-
um forms felty masses between scales; on bulbs and in surrounding soil are
masses of brown to black, attened sclerotia, composed of the yellow-brown,
thin-walled irregular cells typical of Rhizoctonia. These can survive in soil
for years, germinating to infect bulbs after planting or in very early spring.
Occasionally sclerotia are transported on bulbs, but the bulbs are usually so
noticeably diseased that they are not sold.
Control. Remove soil and plants from affected area and for at least 6 inches
beyond. Destroy all infected bulbs at harvest. Use a 4- to 5-year rotation.
ROTS 523
Rhizopus
Zygomyetes, Mucorales
Sporangium large, globose, multispored, with a columella and a thin wall; sporangio-
la and conidia lacking. Sporangiophores arise in fascicles from aerial arching stolons,
which develop rhizoids at point of contact with substratum (Fig. 2.2).
Rhizopus arrhizus. Soft Rot on gladiolus corms, light brown and on beet.
Rhizopus arrhizus (formerly Rhizopus oryzae). Head Rot on sunower.
Rhizopus nigricans (Syn. R. stolonifer), the common black bread mold.
Soft Rot of sweetpotato and other vegetables; Rhizopus Rot, Whiskers,
Leak of peach, strawberry, and other fruits. This is one of the more serious
storage rots of sweetpotato, soft, watery, progressing rapidly, with rotting
complete inside 5 days after visible infection. The tuber is brownish with-
in, covered with a coarse whiskery mycelial growth; there is a mild odor.
Cucurbits, crucifers, carrots, beans, lima beans, onions, peanuts, potatoes,
Jerusalem artichoke, and guava are susceptible to this black mold. Nancy
Hall and Southern Queen are among the more resistant varieties of sweet-
potato. To prevent rot, cure at 80 to 85F for 10 to 14 days, at high humidity,
to permit rapid corking over of wounds; then store at 55F.
The fungus is a weak parasite on ripe fruit peach, g, strawberry, citrus,
persimmon, pear, avocado and melons. A coarse cottony mold appearing in
wounds and over the surface is covered with sporangia, white when young,
black at maturity. A watery uid with an offensive odor leaks from the soft
fruit. Avoid wounding in harvesting; do not pack overripe fruit; keep at low
temperature in transit and market. Amaryllis, lily, and tulip bulbs may be
infected.
Rhizopus oryzae (see Rhizopus arrhizus). Head Rot on sunower.
Rhizopus stolonifer. Soft Rot of Euphorbia trigona.
Roesleria
Mycelium inconspicuous; apothecia cup-shaped, opening more or less completely; asci
disappearing early, leaving a persistent mass of spores and paraphyses. Spores hyaline,
one-celled, globose.
Roesleria hypogaea. Root-Rot of grape.
Rosellinia
Perithecia smooth, ostiole simple or with a low papilla; with a subicle under the fruiting
layer; paraphyses present; spores olive to brown, one-celled.
Rosellinia necatrix (Syn. Dematophora). White Root Rot of g, grape,
avocado, apricot, cherry, apple, pear, peach, walnut, holly osmanthus, privet
and poplar. This is like Armillaria root rot in that all trees in a certain area are
killed, but there are no rhizomorphs or toadstools formed. A white mycelial
growth on surface of affected roots turns black and cobwebby. During wet
weather a delicate mold forms on surface of bark and on soil around base
of tree. Foliage is sparse and wilting, growth slow or none. Crabapples are
quite resistant. Plums and apricots can be grown on resistant understock.
Schizophyllum
Pileus leathery; stipe lateral or none; edge of gills split; spores white.
Schizophyllum commune. Wound Rot, common on dead parts of liv-
ing trees maple, boxelder, almond, acacia, ailanthus, birch, catalpa, hick-
ory, peach, pecan and citrus, g. Fruiting bodies are small, thin, sometimes
lobed, up to 2 inches wide, fan-shaped with gray-white downy upper surface,
brownish forked gills on underside, common on fruit trees. This pathogen
also causes wood decay and decline of apple.
Sclerotinia
Blights.
Botryotinia narcissicola (formerly Sclerotinia narcissicola). Narcissus
Smoulder. Perhaps the fungus should be transferred to Botryotinia, since
there is a conidial stage. The disease is a decay of stored narcissus bulbs,
also known on snowdrop, and a rot of foliage and owers in the open, espe-
cially during cold wet seasons. Leaves are distorted, stuck together as they
ROTS 525
emerge from soil. Sclerotia are small, black, attened bodies, up to 1/2 inch
long when several grow together, just below outer papery bulb scales. In
prolonged storage there is a yellow-brown rot.
Control. Remove and destroy diseased plants as soon as noticed; destroy
weeds to provide air circulation; spray with bordeaux mixture; discard rot-
ting bulbs at harvest; change location every year.
Sclerotinia homoeocarpa. Dollar Spot. Small Brown Patch of turf on
bent grasses, fescues and bluegrass. Spots are brown at rst, then bleached
and straw-colored, about 2 inches in diameter but coalescing to large irregu-
lar patches. While leaves are being killed, a ne white cobwebby growth of
mycelium can be seen in early morning when dew is present.
Sclerotinia intermedia. Stem Rot, market disease of celery, carrot and sal-
sify.
Sclerotinia minor. Stem Rot of lettuce, celery, carrot, cocklebur, Austrian
winter pea, lana woolly pod vetch, phacelia and basil; also stem rot on fennel,
root and crown rot on cabbage, crown rot and wilt on Cichorium (radicchio).
Root and Pod Rot of peanut. Resembles rot due to S. sclerotiorum, but
sclerotia are much smaller. Crown Rot of pepper.
Sclerotinia narcissicola (see Botryotinia narcissicola). Narcissus Smoul-
der. Perhaps the fungus should be transferred to Botryotinia, since there is
a conidial stage.
Sclerotinia sclerotiorum. Black Rot of bulbous iris, hyacinth, narcissus
and tulip. Iris fails to start growth, or plants turn yellow, wilt, and die, often in
clumps. Bulbs are covered with thin gray masses of mold with black irregular
sclerotia between scales. Tulip leaves develop reddish color early in spring,
wilt and die; stems and bulbs are rotted into a crumbly mass of fragments
and black sclerotia. This is a cool-temperature fungus that stops action about
the time the heat-loving Sclerotium rolfsii starts in. Remove diseased plants
and surrounding soil as soon as noticed. Discard all small bulbs at harvest;
plant healthy bulbs in clean soil. The pathogen supposedly dies out after 2
years in soil without suitable host.
Sclerotinia sclerotiorum. Calyx-End Rot on apple and Root and Pod
Rot of peanut, Root and Stem Rot of alfalfa.
Sclerotinia sclerotiorum. Green Fruit Rot of almond, peach, apricot, g
and strawberry; Rhizome Rot of ginseng. In almond, young shoots and
fruits are killed and wither soon after petals fall. Infection takes place through
jackets from apothecia produced under trees where weeds or crop plants have
been previously infected with cottony rot. Losses are serious only when there
is continuous wet weather during and after blooming. Spraying for brown
rot helps to control green rot. Shaking or jarring trees after bloom to remove
jackets from young fruits is suggested.
Sclerotinia sclerotiorum. Leaf and Crown Necrosis of African daisy;
Root Rot and Wilt of peanut. White Mold Rot of soybean.
Sclerotinia sclerotiorum (S. libertiana). Lettuce Drop, Watery Soft Rot
of endive; Pink Rot of celery, Cottony Rot of bean, carrot, parsnip, cab-
bage, and other crucifers and cucurbits. In lettuce, older leaves wilt and fall
at on the ground, leaving center leaves erect, but these are soon invaded by
mycelium and reduced to a slimy wet mass. In continued moisture a thick,
white cottony mold is formed, bearing large black sclerotia up to the size
of peas (see Fig. 3.53). They winter in the soil, send up groups of apothe-
cia in spring. These are brown, cup- to saucer-shaped, up to an inch across,
on a stalk. Ascospores are ejected in a veritable cloud; there is no known
conidial stage.
Control. In commercial celery elds deep plowing or ooding is used to
inhibit apothecial production. Sterilize seedbed soil before planting.
Sclerotinia sclerotiorum. Stem Rot of pepper, cocklebur, tomato, and
many ornamentals aconite, basil, calendula, chervil, canola, chrysanthe-
mum, cynoglossum, dahlia, daisy, delphinium, Gaillardia, gazania, holly-
hock, peony, purple coneower, snapdragon, sunower, zinnia, and others.
The same sort of cottony mold is formed on ower stems as on vegeta-
bles, but here the sclerotia are usually inside the pith and so are rather
Figure 3.53 Sclerotinia sclerotiorum. A sclerotium formed in pith of stem and one falling out from broken stem;
B apothecia produced from sclerotium on ground; C ascus with paraphyses; D, section through sclerotiorum, with
colorless medulla and dark wall on rind cells
ROTS 527
long and thin. You can feel them by running thumb and nger along the
stem; sometimes the cottony mycelium, or cracks in the stem, or one or two
external sclerotia indicate their presence. They are common in peony stems.
When sclerotia are formed in owers, the shape corresponds to oral parts.
Sunowers have large compound sclerotia.
Control. Cut out and destroy affected parts, trying to keep sclerotia from
falling out onto soil. Dusting with sulfur sometimes checks rapid spread of
mycelium.
Sclerotinia trifoliorum. Root Rot, Stem Rot of cocklebur and alfalfa.
Sclerotium
Blights.
Sclerotium cepivorum. White Rot of onion, shallot and garlic. Affected
plants die from a rotting at the neck, at which point there is a surface crust
of small black sclerotia and a thin weft of white mycelium. The sclerotia are
smaller and rounder than those of Botrytis. Roots are often rotted off, and
sometimes spots in a eld covering several square yards are infested.
Sclerotium rolfsii (including Sclerotium delphinii). Crown Rot of delphini-
um, iris, ajuga, aconite, quinoa, sainfoin, kiwi, and many other ornamentals
and vegetables, Root Rot and Wilt of peanut, Wet Scale Rot of narcissus,
Southern Blight. For a full discussion see Pellicularia rolfsii under Blights.
Seaverinia
Apothecia shallow, cup- to disc-shaped; a stroma formed but no denite sclerotia; coni-
dia botryose.
Seaverinia geranii (Syn. Sclerotinia geranii). Rhizome Rot on geranium.
Steccherinum
This genus has been separated off from Hydnum. The pileus is sessile or substipitate and
laterally attached, on a woody substratum; spines are terete or attened; cystidia present;
spores white, smooth.
Climacoden septentrionalis (formerly Steccherinum septentrionale Syn.
Hydnum septentrionale) White Spongy Rot of heartwood of living maples,
beech, hickory, and other hardwoods. A zone of brown discolored wood is
around the white rot area, and there are ne black zone lines. The fruiting
bodies are large, soft, soggy, creamy white, in very large, bracket-shaped
clusters on trunks.
Hericium obietis (formerly Steccherinum abietis Syn. Hydnum abietis).
Brown Pocket Rot of heartwood of living rs and western hemlock in
Pacic Northwest. Elongated pockets, empty or with white bers, are sep-
arated by rm reddish brown wood. Sporophores are like coral, white to
cream, up to 10 or 12 inches high and wide, usually on dead trees, some-
times in wounds of living trees.
Steccherinum abietis Syn. Hydnum abietis (see Hericium obietis). Brown
Pocket Rot of heartwood of living rs and western hemlock in Pacic
Northwest.
Steccherinum septentrionale Syn. Hydnum septentrionale (see Climaco-
den septentrionalis) White Spongy Rot of heartwood of living maples,
beech, hickory, and other hardwoods.
Stereum
Effused-reexed to stipitate; spore-bearing surface smooth, pale brown, upper surface
with a velvety coating of hairs, formed in several distinct layers; gloeocystidia and cys-
tidia present or lacking; spores smooth, colorless.
Chondrostereum purpureum (formerly Stereum purpureum). Silver Leaf,
Sapwood Rot, common on plums and other fruit trees, sometimes impor-
tant on apples, occasional on shade and ornamental trees, widespread but
more serious in the Northwest. The fungus enters through wounds; grows
rst in heartwood, and then kills sapwood and bark; infected branches devel-
op foliage with dull leaden or metallic luster. If the disease is not checked,
the entire tree may be lost. The sporophores appear after death, resupinate to
somewhat shelf-shaped, with purple undersurface.
ROTS 529
Stereum fasciatum (Syn. S. ostrea). Brown Crumbly Rot, mostly on
slash, sometimes on maple and birch. Thin, leathery grayish sporophores
with undersurface light brown, smooth.
Stereum hirsutum. Wood Rot, Sapwood Wound Rot, occasionally near
wounds of living trees birch, maple, hickory, mountain-mahogany, euca-
lyptus, peach, and others. Thin, leathery crustlike sporophores have hairy,
buff to gray upper surface, smooth gray undersurface.
Control. Remove branches and burn at rst sign of silvering. Protect trees
from wounds; paint pruned surface with bordeaux paste or other disinfectant;
keep brush removed from orchard.
Stereum purpureum (see Chondrostereum purpureum). Silver Leaf, Sap-
wood Rot, common on plums and other fruit trees, sometimes important
on apples, occasional on shade and ornamental trees, widespread but more
serious in the Northwest.
Stereum sanguinolentum. Red Heart Rot of slash and living conifers
rs and eastern white pine. Fruiting bodies are small, not over 2 inches wide;
upper surface is a silky pale olive buff; lower surface bleeds readily when
wounded, dries to grayish brown. Sporophores are produced in profusion on
dead wood, occasionally on dead branches of living trees.
Streptomyces
Schizomycetes, Actinomycetales
Intermediate form between bacteria and fungi. Much-branched mycelium that does not
fragment in bacillary or coccoid forms; conidia in chains on sporophores; primarily soil
forms, some parasitic.
Streptomyces ipomoea (Syn. Actinomyces ipomoea) Soil Rot or Pox of
sweetpotatoes, general New Jersey to Florida and in the Southwest. This
pathogen also infects Jacquemontia. Leaves are small, pale green to yellow;
plants are dwarfed, make little or no vine growth, and may die before end of
the season; the root system is poorly developed with most roots rotted off,
or breaking off if plant is pulled from the soil. Small dark lesions are formed
on stems below the soil line. Pits with jagged or roughened margins, often
coalescing, are formed on mature roots. The rot is found in soils at pH 5.2
or above; and is worse in dry soils and seasons. Variety Porto Rico is very
susceptible.
Control. Apply sulfur to acidify soil to pH 5.0.
Stromatinia
Apothecia arising from a thin, black, subcuticular, effuse sclerotium or stroma; small
black sclerotia are borne free on mycelium, not giving rise to apothecia. There is no
conidial stage; apothecia resemble those of Sclerotinia.
Stromatinia gladioli (Syn. Sclerotinia gladioli). Dry Rot of gladiolus,
also found on crocus, freesia and tritonia. Lesions on corms start as red-
dish specks, with slightly elevated darker border; spots enlarge, and centers
become sunken, dark brown to black with lighter raised edges; they grow
together into irregular areas. On husks the lesions are tobacco brown. Very
small black sclerotia are formed on husks, in corm lesions, and on dead
stems. Plants in the eld turn yellow and die prematurely owing to decay
of leaf sheath. Corms may appear normal when dug, the rot developing in
storage. The disease is more prevalent in heavy soils, and the fungus can
survive several years in soil. Apothecia have been produced articially by
fertilizing receptive bodies on sclerotia with spermatia (microconidia). They
are densely crowded, 3 to 7 mm broad, on stipes 6 to l0 mm high.
Control. Use soil with good drainage and a 4-year rotation. Removing husks
before planting helps to reduce gladiolus rot diseases. Cure corms rapidly
after harvest.
Stromatinia narcissi. Large-scale speck fungus on narcissus and zephyran-
thes. Black, thin, round, at sclerotia 1/2 to 1 mm, adhere rmly to outer
scales. The fungus is mostly on bicolor varieties and seems to be saprophyt-
ic without causing a denite disease.
Thielaviopsis
Hyphae dark; two kinds of conidia-small, cylindrical, hyaline endogenous spores and
large, ovate, dark brown exogenous spores, both formed in chains.
Thielaviopsis basicola. Black Root Rot, seedling root rot of tobacco and
many vegetables bean, carrot, corn, chickpea, lentil, okra, onion, pea, toma-
to, corn-salad, vinca, and watermelon; and ornamentals begonia, cyclamen,
gerbera, elm, oxalis, lupine, pelargonium, peony, poinsettia, pansy, scindap-
sus, and others. There is blackening and decay of roots; young plants damp-
ROTS 531
off and die; older plants are stunted, with the decay proceeding until all roots
are destroyed. Stem discoloration extends 2 to 3 inches above the soil line.
The fungus lives in soil as a saprophyte, entering through nematode wounds.
Hyaline conidia produced inside conidioles are forced out through hyphal
tips. Chlamydospores are larger, dark, club-shaped, with several cells; they
break up so that each pillbox acts as a spore. This disease is especially serious
on poinsettia, dwarng plants, causing misshapen leaves and ower bracts.
The rot is most destructive in heavy, cold, slightly acid to alkaline soils well
supplied with humus. Long wet periods after transplanting increase rot. Soils
with pH lower than 5.6 or sandy soils low in organic matter are less con-
ducive to disease.
Control. Sterilize soil for seedbeds; use clean pots for poinsettias and other
greenhouse plants; reduce pH with sulfur or by using half peat moss and half
soil.
Trametes
Pileus without stipe, sessile to effuse-reexed, rm; hymenium white or pallid, punky to
corky, not friable when dry; tubes unequally sunken.
Trametes suaveolens. White Wood Rot of willow and poplar, after wound-
ing. A dry, corky decay with an anise odor begins in lower trunk and pro-
gresses upward. Leathery to corky sporophores 6 inches wide are white when
young, gray to yellow with age.
Trichoderma
Conidia in heads on conidiophores divided into two or three tips, a single head on each
tip; spores hyaline, one-celled.
Trichoderma viride. Green Mold Rot, Cosmopolitan on narcissus, also
on shallot, garlic, occasional on citrus, but saprophytic. This fungus has an
antibiotic or antagonistic effect on Rhizoctonia, Pythium, and other damping-
off fungi and is quite helpful in reducing Armillaria root rot and crown rot
due to Sclerotium rolfsii.
Trichoderma harzianum. Fruit Rot of apples in storage.
Trichothecium
Conidiophores long, unbranched; conidia two-celled, hyaline or bright, single, at apex
of conidiophore; upper cell usually larger than basal cell; mostly saprophytic.
Trichothecium roseum. Fruit, Storage Rot on tomato, g, celery, carrot,
occasional on quince and pear; a pink mold. Pink Mold Rot on plum, nec-
tarine and peach.
Ustulina
Stroma globoid, cupulate to pulvinate; carbonaceous, black, somewhat hollow; spores
dark, one-celled.
Ustulina deusta. White Heart Rot, a brittle white rot with prominent black
zones in butts of living hardwoods; prevalent on sugar maple sprouts. Black
crusts appear on stumps, logs, and on at cankered areas of American beech.
Valsa
Cankers and Diebacks.

Valsa ambiens subsp. leucostomoides. Causing decay around holes of
tapped sugar maples.
Leptographium (Verticicladiella)
Deuteromycetes, Hyphomyceres
Conidiophores upright, tall, brown, branched only near apex, penicillate; conidia (sym-
podulospores) hyaline, one-celled, ovoid to clavate, often curved, apical on sympodially
formed new growing points, in slime droplets.
Leptographium abietinum (formerly Verticicladiella abietina). Root Rot
on white pine.
Leptographium penicillata (formerly Verticicladiella penicillata. Root
Rot on white pine.
ROTS 533
Leptographium procera (formerly Verticicladiella procera. Root Rot and
Decline of eastern white pine, sand pine and red pine.
Leptographium wagenerii (formerly Verticicladiella wagenerii. Root Rot
on r and pine.
Verticicladiella abietina (see Leptographium abietinum). Root Rot on
white pine.
Verticicladiella penicillata (see Leptographium penicillata). Root Rot on
white pine.
Leptographium procera (see Verticicladiella procera). Root Rot and
Decline of eastern white pine, sand pine and red pine.
Leptographium wagenerii (see Verticicladiella wagenerii). Root Rot on
r and pine.
Xylaria
Stroma is upright, simple or branched; perithecia, immersed laterally, are produced after
conidia; spores dark, one-celled.
Xylaria hypoxylon. Root Rot of hawthorn and gooseberry.
Xylaria mali. Black Root Rot of apple, also honey locust. Wood is soft,
spongy, dirty white, with narrow conspicuous black zones forming fantas-
tic patterns. Roots are covered with thin compact white mycelium, which
changes to black incrustations. Fruiting bodies are dark brown to black, club-
shaped, 1 to several inches high, united at the base, extending upward like
a fan. The disease is not common, and where it does occur, only a few trees
are killed.
Xylaria polymorpha. On decaying wood, identied by cylindrical thumb-
like fruiting bodies.
RUSTS
Rust fungi belong to the Uredinales, a highly specialized order of the Basid-
iomycetes. In common with mushrooms they have spores of the sexual stage
borne in fours on a club-shaped hypha known as a basidium, but apart from
this they differ very decidedly from woody and eshy Basidiomycetes. The
term rust is applied both to the pathogen and to the disease it inicts. There
are more than 4000 species of rusts, all obligate parasites on ferns or seed
plants. Many are heteroecious, completing their life cycle on two differ-
ent kinds of plants; but some are autoecious (monoecious), having all spore
forms on a single host species. There are only two families, Melampsoraceae
and Pucciniaceae.
Many rusts show physiological specialization, the existence within a species
of numerous strains or races that look alike but attack different varieties of
crop plants, thus greatly complicating the problem of breeding for rust resis-
tance. Rusts with a complete life cycle have ve different spore forms, num-
bered 0 to IV.
0. Pycniospores (spermatia) formed in pycnia (spermagonia). The pycnia
resemble pycnidia of Ascomycetes, are usually on upperside of leaves.
They discharge one-celled pycniospores with drops of nectar, and these,
usually distributed by insects attracted to the sweet secretion, function
in fertilization.
I. Aeciospores (aecidiospores), one-celled, orange or yellow, formed, often
in chains, in a cuplike sorus or aecium, which has a peridium (wall)
opening at or beyond the surface of the host.
II. Urediospores (uredospores, summer spores, red rust spores), one-celled,
walls spiny or warty, reddish brown, on stalks or in chains in a uredium
(uredinium or uredosorus), over which the epidermis of the host is bro-
ken to free the spores. Resting II spores, formed by some rusts, have
thicker and darker walls.
III. Teliospores (teleutospores, winter spores, black rust spores), one or more
cells, in telia (teleuto sori), either on stalks, as in the family Puccini-
aceae, or sessile, in crusts or cushions as in the Melampsoraceae.
RUSTS 535
IV. Basidiospores (sporidia) on a basidium or promycelium formed by the
germinating teliospore. Basidium is usually divided transversely into
four cells, with one sporidium formed from each cell at the tip of a sterig-
ma.
In heteroecious rusts spore stages 0 and I are formed on one host and II
and III on another, and are so indicated in the information given with each
species. Stage IV always follows III on germination. Although most autoe-
cious rusts have all spore forms, on one host, there are a few short-cycle
(microcylic) rusts with some spore stages dropped out. For a detailed life
history of a heteroecious rust,

Puccinia graminis.
Gardeners frequently mistake a reddish discoloration of a leaf, perhaps due
to spray injury or weather or a leaf-spot fungus, for rust. True rust is iden-
tied by the presence of rust-colored spores in powdery pustules or perhaps
gelatinous horns. With rusts, the discoloration of tissue is yellowish, not red,
and it is due to increased evaporation from the broken epidermis. Plants are
often stunted.
Losses in food crops due to rust have been enormous since the beginning of
history. The Romans had a festival to propitiate the rust gods. Now we try to
do it by removing the alternate host, barberry to save wheat, black currants
to save white pine; or by developing more and more resistant varieties for
the ever increasing rust strains; or by the use of fungicides, classically sulfur,
latterly some of the carbamates, and, in a few cases, antibiotics.
Achrotelium
Melampsoraceae. Telia on underside of leaves; spores one-celled at rst, four-celled on
germination, stalked.
Achrotelium lucumae. II, III on lucuma and egg fruit, Florida.
Aecidium
This is a form genus, a name applied to the aecial stage where the full cycle is unknown
and 0 and I are the only spores. Aecia have a peridium and catenulate spores. There are
many species.
Aecidium avocense. On poppy-mallow, probably aecial stage of Puccinia
avocensis.
Aecidium conspersum. On houstonia and galium, Wisconsin.
Aecidium rubromaculans. On viburnum, Florida.
Angiospora
Physopella.
Aplopsora
Melampsoraceae. Teliospores sessile, hyaline, one-celled, in a single layer;
aecia unknown.
Aplopsora nyssae. On tupelo, II, III.
Baeodromus
Pucciniaceae. Spores one-celled; telia pulvinate, erumpent; short chains of spores.
Baeodromus californicus. On senecio, III.
Baeodromus eupatorii (see Coleosporium steviae). On eupatorium.
Coleosporium steviae (formerly Baeodromus eupatorii). On eupatorium.
Bubakia (Phakopsora)
Melampsoraceae. Telia indehiscent, lenticular, spores formed in irregular succession,
one-celled. Uredia without peridium or paraphyses.
Bubakia erythroxylonis. On erythroxylon.
Caeoma
Form genus. Aecia with catenulate spores but no peridium.
Caeoma faulliana (see Melampsora medusae). Needle Rust on alpine r.
Aecia orange-yellow, on needles of current year.
Caeoma torreyae. On torreya, California.
Melampsora medusae (formerly Caeoma faulliana). Needle Rust on
alpine r. Aecia orange-yellow, on needles of current year.
RUSTS 537
Cerotelium
Pucciniaceae. Spores one-celled; teliospores in a many-layered mass; hyaline, not exsert-
ed through stomata; aecia with peridium; uredia with paraphyses; spores borne singly.
Cerotelium dicentrae. 0, I on bleeding heart; II, III on Urticastrum.
Cerotelium ci (Physopella ci). Fig Rust, II, III on common g, Flori-
da strangler g and osage-orange, Alabama, Florida, Louisiana, Minnesota,
South Carolina, Texas.
Chrysomyxa
Melampsoraceae. Teliospores in cylindrical or branching chains; promycelium exserted;
urediospores typically in short chains; uredia without peridium.
Chrysomyxa arctostaphyli. On bearberry, III.
Chrysomyxa chiogenis. II, III on creeping snowberry; 0, I on spruce.
Chrysomyxa empetri. II, III on crowberry; 0, I on red and white spruce.
Aecia on upper and lower surfaces of needles.
Chrysomyxa ilicina. II, III on American holly.
Chrysomyxa ledi. 0, I, on black, red and Norway spruce; II, III on underside
of leaves of Ledum spp.
Chrysomyxa ledi var. cassandrae. Spruce Needle Rust. 0, I on black,
red, blue and Engelmann spruce; II, III on bog rosemary (Chamaedaphne).
May become epidemic on spruce, causing considerable defoliation.
Chrysomyxa ledi var. groenlandici. On Labrador-tea, Michigan, New
Hampshire.
Chrysomyxa ledi var. rhododendri. II, III on rhododendron, Washington.
A European rust rst noted on Pacic Coast in 1954, apparently entering
despite quarantine on nursery stock. Yellow uredia on leaves.
Chrysomyxa ledicola. 0, I on white, black red, blue, Engelmann, and Sitka
spruce; II, III on upperside of leaves of Ledum spp. Spruce needles may be
so discolored that trees appear yellow.
Chrysomyxa moneses. On Sitka spruce and moneses.
Chrysomyxa piperiana. 0, I on Sitka spruce; II, III on underside leaves of
Rhododendron californicum, California, Oregon, Washington.
Chrysomyxa pirolata (C. pyrolae). 0, I on cones of black, blue, Engelmann,
Norway, red and white spruce; II, III on pyrola. Aecia are on upperside of
cone scales; infected cones turn yellow, produce no seed.
Chrysomyxa weirii. Spruce Needle Rust. III on Engelmann and red
spruce. Waxy orange to orange-brown elongate or elliptical telia occur on
1-year needles. This is the only spore stage known; teliospores can reinfect
spruce.
Coleosporium
Melampsoraceae. Pycnia and aecia are on pines; uredia and telia on dicotyledons. Pyc-
nia subepidermal or subcortical, attish, linear, dehiscent by a slit; aecia on needles,
erumpent, with prominent peridium, spores ellipsoid or globular; uredia erumpent, pow-
dery without peridia; urediospores globose or oblong, catenulate, with verrucose (warty)
walls; telia indehiscent, waxy, gelatinous on germination; spores sessile or obscurely
catenulate, one-celled, smooth but with thick and gelatinous walls.
Coleosporium apocyanaceum. 0, I on loblolly, longleaf, and slash pines; II,
III on Amsonia spp. in the Southeast.
Coleosporium asterum (C. solidaginis). Needle Blister Rust of pine. 0,
I on all two- and three-needle pines in eastern United States; II, III on aster
and goldenrod, on China aster (except far South), on golden aster (Chrysop-
sis), erigeron, grindelia, seriocarpus, and other composites. This blister rust
on pine needles has pustules higher than they are long, in clusters or short
rows. The rust is fairly common on ornamental pines in gardens, wintering
on aster and related composites. Older needles of young pines may be severe-
ly infected, with white aecia conspicuous in spring and early summer. Aster
leaves have bright orange-yellow spore pustules on undersurface. Destroy
goldenrod near pines.
Coleosporium crowellii. III only stage known; on needles of pinon and lim-
ber pines, Arizona, Colorado, New Mexico, Utah, Nevada, and California.
Coleosporium delicatulum. Pine Needle Rust. 0, I on two- and three-
needle pines; II, III on goldenrod and euthamia.
Coleosporium helianthi. 0, I on two- and three-needle pines, especially in
the Southeast; II, III on silphium and parthenium.
Coleosporium helianthi. Sunower Rust. 0, I on pitch and short-needle
pines; II, III on wild and cultivated sunower, Jerusalem artichoke and
heliopsis. Sunower leaves, with brown rust pustules, dry up and drop.
Control is not easy.
RUSTS 539
Coleosporium ipomoeae. 0, I on southern and Chihuahua pines; II, III
on moonower, morning-glory, sweetpotato, jacquemontia, and quamoclit;
most abundant in warmer regions. The uredia are orange-yellow, telia deep
reddish orange on sweetpotato.
Coleosporium jonesii. 0, I on pinon pine; II, III on owering currant and
gooseberry.
Coleosporium lacinariae. 0, I on loblolly, longleaf, and pitch pines; II, III
on liatris.
Coleosporium mentzeliae. On mentzelia.
Coleosporium minutum. 0, I on loblolly and spruce pines; II, III on
forestiera.
Coleosporium pacicum. 0, I on Monterey, Coulter, and Jeffrey pines; II,
III on marigold, sunower, tarweed, and other composites.
Coleosporium pinicola. III on Virginia or scrub pine.
Coleosporium tussilaginis. 0, I on pitch, red, and Virginia pines; II, III on
campanula, lysimachia, and specularia, Underside of bluebell leaves are cov-
ered with orange to reddish brown pustules. Leaves dry; plants are stunted.
Coleosporium tussilaginis. 0, I on Scotch pine; II, III on sow-thistle.
Coleosporium tussilaginis. 0, I, unknown; II, III on senecio.
Coleosporium vernoniae. 0, I on two- and three-needle pines in South; II,
III on elephantopus.
Coleosporium vernoniae. 0, I on various two- and three-needle pines; II, III
on ironweed.
Coleosporium viburni. 0, I, unknown; II, III on Viburnum spp.
Cronartium (Causing Blister Rusts)
Melampsoraceae. Heteroecious; pycnia and aecia on trunk and branches of pine; uredia,
telia on herbaceous or woody dicotyledons.
Pycnia on stems, caeomoid, forming blisters beneath host cortical layer; dehiscent by
longitudinal slits in bark; aecia on trunks, erumpent, with peridium sometimes dehiscent
at apex, more often spliting irregularly or circularly at side; aeciospores ellipsoid with
coarsely warted walls, sometimes with smooth spot on one side. Uredia on underside of
leaves or on stems of herbaceous hosts; delicate peridium, dehiscent at rst by a central
pore; urediospores borne singly on pedicels, ellipsoidal with spiny walls; telia erumpent,
often coming from uredia; catenulate, one-celled teliospores often forma extended cylin-
drical or liform column, horny when dry (see Fig. 3.54).
Figure 3.54 Pine Rusts. Coleosporium asterum, uredial (II) and telial (III) stages on aster, teliospores germinating
in situ. Cronartium ribicola, II and III stages on currant
Blister rusts are characterized by swellings that are globose, subglobose, or fusiform,
depending on species. A rust on a pine stem is invariably a Cronartium, although this
stage has often gone under the name of Peridermium.
Cronartium appalachianum (Peridermium appalachianum). I on Virginia
pine, North Carolina, Tennessee, Virginia, West Virginia. Girdling bark
lesions with columnar aecia.
Cronartium coleosporioides. Ponderosa Pine Rust, widespread in
Rocky Mountains; II, III on Indian paintbrush.
Cronartium coleosporioides (C. lamentosum). Western Gall Rust,
Paintbrush Blister Rust. 0, I on lodgepole, ponderosa and Jeffrey pines, in
West; II, III on Indian paintbrush, birds-beak, owls-clover and wood-betony.
Slight swellings are formed on twigs, trunks, and branches; many lodgepole
pine seedlings are killed.
Cronartium comandrae. Comandra Blister Rust. 0, I on ponderosa, Ari-
zona, and lodgepole pines in West and pitch, mountain,jack, loblolly, Austri-
an, Scotch, and maritime pines in the East; II, III on bastard toadax (Coman-
dra spp.). Destructive effect is limited to distribution of toadax, which is
widespread but locally restricted to small areas. Ponderosa pine suffers most
severely, with many seedlings and saplings destroyed; occasionally a large
tree is attacked.
Cronartium comptoniae. Sweet-Fern Blister Rust. 0, I on two- and
three-needle pines; II, III on sweet-fern and sweet gale in northern pine
regions and south to North Carolina, and on Pacic wax-myrtle on Pacif-
ic Coast. Young pines may be girdled and killed, but are fairly safe after
attaining a trunk diameter of 3 inches. Losses in nurseries and plantations are
high, especially among lodgepole and ponderosa pines. Affected stems swell
slightly near the base with long fusiform swellings or depressed streaks on
RUSTS 541
eastern hard pine; pitch oozes out from insect wounds in these areas. Killing
of main stem often results in multiple-stemmed shrublike trees. Orange aecia
appear on 3-year seedlings, preceded by pycnia the year before; spores are
wind-borne many miles to herbaceous hosts.
Control. Remove Myrica species for several hundred yards around nurseries
or pine plantations, and allow no large groups within a mile.
Cronartium conigenum. Pine Cone Rust. 0, I on cones of Chihuahua
pine; II, III on oaks in Southwest. Cones develop in large galls producing
aecia with distinct, erumpent peridium 2 or 3 years after infection.
Cronartium harknessii (see Endocronartium harknessii). Western Gall
Rust. 0, I on Jeffrey, ponderosa, lodgepole, and digger pines; II, III on Indian
paintbrush, lousewort, owls-clover, or omitted, with direct infection from
pine to pine.
Cronartium occidentale. Pinon Blister Rust. 0, I in pinon and Mexican
pinon; II, III on currant, gooseberry and owering currant. This rust cannot
be told from whitepine blister rust on Ribes hosts, but is differentiated by the
type of pine attacked. Aecia on Mexican or singleleaf pinon are distinct sori;
on pinon they form broad layers under bark.
Cronartium quercuum f. sp. fusiforme. Rust on pine.
Cronartium quercuum f. sp. fusiforme. Southern Fusiform Rust, 0, I
on hard pines in southern states, especially loblolly, slash, and pitch pine; II,
III on evergreen oaks on underside of leaves. Pine stems have pronounced
spindle-shaped swellings, sometimes with witches brooms. Branch infec-
tions that do not reach the main trunk are not serious, but those that go on to
the trunk may kill the tree. Longleaf pines are rather resistant, and shortleaf
P. echinata almost immune. Pines well spaced in good locations grow more
rapidly and may have more rust than those in poor sites. It has also been
reported on oaks.
Control. Prune branches yearly before swellings reach main stem.
Cronartium quercuum (C. cerebrum). Eastern Gall Rust. 0, I on pines,
especially scrub and shortleaf in the South; II, III on chestnut, tanbark and
oak. Globose to subglobose galls are formed on pine stems; in spring aecia
break through the bark in more or less cerebroid (brainlike) arrangement.
Cronartium ribicola. White Pine Blister Rust. 0, I on eastern white pine
from Maine to Virginia and Minnesota, on western white pine in the Pacic
Northwest, on sugar pine in California; II, III on currant, owering currant
and gooseberry. Occurs also on limber pine in Northcentral and Southeastern
Wyoming.
This dread disease is supposed to have originated in Asia, whence it spread
to Europe, where the eastern white pine introduced from America was very
susceptible. White pine blister rust was found in Russia in 1854, and by
1900 had spread over most of Europe. It was recorded on Ribes at Geneva,
New York, in 1906, but probably was there some years previously. In 1909
it was found on pine, at which time it was learned that infected pines from
a German nursery had been widely planted throughout the Northeast. The
next year the disease reached Vancouver, British Columbia, in a shipment
from a French nursery, whence it spread to Washington, Oregon, Northern
California, Idaho, and western Montana. Thus from cheap stock brought in
for forest planting has come one of our greatest forest hazards. Our present
quarantine laws are designed to prevent such introductions.
The western white sugar and whitebark pines are even more susceptible
to blister rust than eastern white pine; but in either case robust, dominant
trees are more severely attacked, with frail individuals lightly infected. This
however, is partly explained by more vigorous trees having more needles to
receive spores. Of the Ribes species, black currant is most susceptible and
dangerous. Cultivated red currants are somewhat resistant, causing a min-
imum of pine infection; Viking and Red Dutch varieties are practically
immune. Wild gooseberries and skunk currant are highly susceptible in the
Northeast, as are western black currant, stink currant, and red owering
currant. The greater the susceptibility of the Ribes species, the more spores
are produced to inoculate pines, with proportionate damage.
Symptoms and Life History. When a spore arrives on a pine needle from
a currant, the rst sign of infection is a small golden yellow to reddish brown
spot. The next season, or possibly in two years, the bark looks yellowish,
often with an orange tinge to the margin of the discolored area, and there
may be a spindle-shaped swelling. If such symptoms appear early in the
season, pycnia are formed in bark by July or August; but if discoloration
is delayed until midsummer, they appear the next year. The male fruiting
bodies are small, honey yellow to brown patches, swelling to shallow blisters
and rupturing to discharge drops of a yellowish, sweet liquid. After this is
eaten by insects or washed away by rain, the lesions turn dark. The next
spring or summer aecia push through the bark in the same region. These
are white blisters, rupturing to free orange-yellow aeciospores, which are
carried away by wind. The bark then dries out and cracks, with death of
cambium and underlying wood. The disease has taken 3 to 6 years to reach
this stage.
RUSTS 543
Production of aecia continues yearly until stem is killed beyond the lesion.
Dead foliage assumes a conspicuous red-brown color. This ag of brown
on a green background is the most conspicuous symptom of blister rust
before death of the pine. Infection progresses downward from small to larg-
er branches and into trunk. Swellings are not apparent on stems much over
2 inches in diameter on eastern white pine, but in the West they sometimes
show up in stems 5 inches through. Larger limbs and trunks sometimes show
constriction in the girdled area.
The aeciospores, large, ellipsoidal, with thick, warty walls, are carried by
wind great distances to Ribes species (they cannot reinfect pine). They send
their germ tubes into a currant or gooseberry leaf through stomata, and with-
in 1 to 3 weeks pinhead-size blisters appear in clusters on yellowed leaf
tissue. These uredia rupture to release large, ellipsoidal, yellow urediospores
with thick, colorless walls and short, sharp but sparse spines. The spores are
somewhat moist and sticky, and are windborne short distances to other Ribes
bushes nearby. There may be up to seven generations in a summer, or the
spores may remain viable over winter in uredia; this stage can infect only
currant.
In late summer telia follow uredia in the same or new leaf lesions, appear-
ing as short brown bristles on underside of leaves or looking like a coarse
felt. Each felty bristle is composed of vertical rows of broad, spindle-shaped
spores, which germinate in situ to a ve-celled promycelium with each of the
four upper cells bearing at the point of a sterigma a small, thin-walled, round
basidiospore. This cannot reinfect currant and soon dies from exposure to
the sun unless the wind blows it immediately to a pine needle. The effective
range is around 300 feet except for spores from black currants, which can be
carried a mile. The spores from pine to currant can be carried many miles,
up to 300. Blister rust is more important at elevations of 1000 feet or over,
where it is increased by lower temperatures and more rainfall.
Control. Eradication of the Ribes host is denitely effective in controlling
white pine blister rust. This means complete removal of black currants and
local removal of cultivated red and wild currants and gooseberries within 300
or 900 feet of pines, according to state regulations, taking care to get all the
root system capable of resprouting.
Blister rust is seldom found on ornamental pines in cities; the smoke and
fumes are unfavorable to the fungus. Elsewhere valuable ornamentals can
be saved by cutting off infected branches and cleaning out trunk infection,
stripping off diseased bark and a 2-inch side margin, 4-inch margin at top
and bottom, of healthy bark. If the cankers are nearer to the trunk than 6
inches, the bark should be excised around the branch stub. The red currant
Viking is immune to blister rust, and a couple of black currant hybrids are
resistant. Some white pines are exhibiting resistance.
Cronartium stalactiforme (see Peridermium stalactiforme). 0, I on lodge-
pole pines in Rocky Mountain regions; II, III on Indian paintbrush.
Cronartium strobilinum. Pine Cone Rust. 0, I on cones of longleaf and
slash pines; II, III on evergreen oak. Cones are swollen, reddish; 25 to 90%
drop.
Endocronartium harknessii (formerly Cronartium harknessii). Western
Gall Rust. 0, I on Jeffrey, ponderosa, lodgepole, and digger pines; II, III
on Indian paintbrush, lousewort, owls-clover, or omitted, with direct infec-
tion from pine to pine. Galls are globose, with large, conuent aecia; bark
sloughs off in large scales; witches brooms are formed. A variety of this
species, alternate stage unknown, occurs on Monterey and knobcone pines
in California.
Control. Remove trees with galls for a distance of 300 yards around nurs-
eries. Do not ship infected trees from nurseries.
Peridermium stalactiforme (formerly Cronartium stalactiforme). 0, I on
lodgepole pines in Rocky Mountain regions; II, III on Indian paintbrush.
The rust enters pine trunks through small twigs, producing diamond-shaped
lesions that elongate an average of 7 inches a year, but grow laterally less
than 1/2 inch. Removal of diseased trees is the only known control.
Cumminsiella
Pucciniaceae. Autoecious; teliospores two-celled; pycnia and other sori subepidermal;
aecia cupulate.
Cumminsiella mirabilissima. 0, I, II, III on barberry and mahonia in the
West, Arizona, California, Colorado, Idaho, Montana, Nebraska, New Mex-
ico.
Cumminsiella texana. On barberry, Texas.
RUSTS 545
Desmella
Pucciniaceae. Uredia and telia subepidermal, protruding in tufts; uredia without peridi-
um or paraphyses. Spores globoid, on pedicels, two-celled.
Desmella aneimiae. On Boston fern, Florida.
Endocronartium
Badisiomycete, Uredinales, Pucciniaceae.
Endocronartium harknessii. Western Gall Rust or Pine-Pine Gall
Rust on pine.
Endophyllum
Pucciniaceae. Teliospores in form of aeciospores; telia with cupulate peridium.
Endophyllum sempervivi. III on houseleek and hen-and-chickens. Succu-
lent leaves may be covered with reddish pustules. This is not common, but
may be serious. Clean out infected parts.
Endophyllum tuberculatum. III on hollyhock and checkermallow.
Frommeella
Pucciniaceae. Teliospores two-to many-septate; aecia and uredia erumpent.
Frommeella duchesneae. II, III on mock-strawberry, false strawberry, or
Aztec Indian berry.
Arthuriomyces (Gymnoconia)
Pucciniaceae. Uredia lacking; aecia present but without peridium; teliospores two-
celled, one pore in each cell.
Arthuriomyces peckianus (formerly Gymnoconia peckiana (G. intersti-
tialis). Orange Rust of blackberry. 0, I, III on blackberry, dewberry and
black raspberry, rst described from eastern United States in 1822, present
from Canada to Florida and from Alaska to southern California. Very bright
orange spores cover underside of leaves in spring. The mycelium is perennial
in the bush, living throughout the year between cells of the stem, crown, and
roots, each season invading new tissue as new growth begins. Shoots may be
bunched, often with a witches broom effect; plants are dwarfed. Spraying
is useless; infected plants never recover. Plant only healthy stock, obtained
from a nursery where the disease is unknown. Remove infected plants show-
ing upright habit of growth, yellow color, and glistening yellowish dots of
pycnia before the orange spore stage appears. Blackberry varieties Eldorado,
Orange Evergreen, Russell, Snyder, Ebony King, dewberry Leucretia, and
boysenberries are quite resistant.
Gymnoconia peckiana (G. interstitialis) (see Arthuriomyces peckianus).
Orange Rust of blackberry. 0, I, III on blackberry, dewberry and black
raspberry, rst described from eastern United States in 1822, present from
Canada to Florida and from Alaska to southern California.
Gymnosporangium
Pucciniaceae. All but one species heteroecious. Picnia and aecia usually on trees and
shrubs of the apple family; telia conned to cedars and junipers except for one species
on cypress; uredia wanting. Teliospores thick- or thin-walled, various in form but mostly
at, tongue-shaped, expanding greatly when moistened, usually with two cells; walls
smooth, one to several pores in each cell; pedicel colorless, usually with outer por-
tion swelling and becoming jellylike when moistened. Aecia are highly differentiated
and conspicuous, with catenulate aeciospores, deeply colored with verrucose walls (see
Fig. 3.55).
The life cycle is similar in all juniper leaf rusts. In early summer, small, slightly swollen
spots appear on leaves of the pomaceous host, then small raised specks in this area on the
upper surface, openings of ask-shaped pycnia embedded in leaf tissue. After exuding
an orange liquid containing pycniospores, the specks are black. Later, aecia push out on
the underside of the same spots as dingy white columns, rostelia, with the outer coat-
ing rupturing to release a powdery mass of yellow to brown aeciospores. The ruptured
segments sometimes make the open aecium look star-shaped, but in the common cedar-
apple rust aecia are cup-shaped. Aecia are also formed on fruit and tender green stems.
Aeciospores released during summer are wind-borne to junipers. Mycelium winters in
the juniper needle or stem, and in spring galls are started that take a year or more to
produce teliospores in cushions or horns.
RUSTS 547
Figure 3.55 Cedar-Apple Rust. Gymnosporangium juniperi-virginianae. A section through crabapple leaf with
pycnidia (0) on upper surface and aecium (I ) with prominent peridium and aeciospores in chains on under-
surface; B two-celled teliospores on gelatinous stalks, which help form the jellylike telial horns on cedar galls;
C, teliospores germinating with a promycelium and basidiospores; D teliospore of G. clavipes, the quince rust
Gymnosporangium bermudianum. 0, I, III on stems of eastern and south-
ern red-cedar in the Gulf states. No alternate host; aecia precede telia on
small galls.
Gymnosporangium bethelii. III on Rocky Mountain juniper; 0, I on fruits
of hawthorn.
Gymnosporangium bethelii. III on stems of prostrate and Rocky Moun-
tain junipers; 0, I on leaves, fruit of hawthorn. Telia are 3 to 4 mm high on
irregular galls on cedar twigs and branches.
Gymnosporangium biseptatum. III on stems of Chamaecyparis thyoides;
0, I on amelanchier. Spindle-shaped swelling in stem; trees may die.
Gymnosporangium clavariiforme. III on common and mountain juniper;
0, I on chokeberry, amelanchier, pear and quince. Slender telia 5 to 10 mm
high produced on long fusiform swellings on branches.
Gymnosporangium clavipes. Quince Rust. III on eastern red-cedar,
dwarf, mountain, and prostrate junipers; 0, I on fruits and young stems of
amelanchier, apple, chokeberry, crabapple, hawthorn, mountain-ash, quince,
Japanese quince and pear. Short slight swellings, somewhat spindle-shaped,
occur in cedar twigs and branches, many of which die. On the main trunk,
infected areas are black rough patches or rings around the bark. Mycelium
is perennial, conned to the outer layer of living bark; it can sometimes be
scraped out by scraping the bark. On pomaceous hosts, the disease is most
frequent on fruits, often causing distortion. Rust sometimes affects twig and
buds but seldom leaves. Aecia are particularly prominent on hips of English
hawthorn, with long whitish perithecium around orange spores.
Control. Some apple varieties susceptible to apple rust are rather resistant to
quince rust, including Jonathan, Rome, Ben, Davis, and Wealthy. Red Deli-
cious is quite susceptible. Destroy cedars in neighborhood of orchards; spray
as for apple rust.
Gymnosporangium confusum. III on Savin Juniper; 0, I on hawthorn.
Gymnosporangium cornutum (G. auriantiacum). Juniper Gall Rust. III
on leaves and stems of common juniper; 0, I on mountain-ash.
Gymnosporangium cunninghamianum. III on Arizona cypress; 0, I on
amelanchier.
Gymnosporangium davisii. III on mountain and common juniper; 0, I on
leaves of red and black chokecherry. Telia are usually on upper surface of
needles, sometimes at base of stems.
Gymnosporangium effusum. III on eastern red-cedar; 0, I on chokeberry.
Fusiform swellings on cedar trunk and branches.
Gymnosporangium ellisii. Witches Broom Rust. III on southern white
cedar (Chamaecyparis); 0, I on sweet-fern, gale, bayberry, wax-myrtle
leaves, fruits and young stems. Aecia are cluster cups; telia are cylindrical,
liform, 3 to 6 mm high, appearing on leaf blade or axil the rst season after
infection, thereafter only on stems, invading inner bark and wood. Witches
brooms are abundant; even large trees die if heavily broomed.
Gymnosporangium exiguum. III on leaves of alligator and Mexican
junipers, eastern red-cedar; 0, I on leaves, fruits of hawthorn.
Gymnosporangium exterum. III on stems of eastern red-cedar; 0, I on
gillenia. Flattened telia anastomose over short fusiform swellings with
roughened bark on cedars. Also galls on stems of juniper.
Gymnosporangium oriforme. III on red-cedar; 0, I on leaves of hawthorn.
Cedar galls are small.
Gymnosporangium fraternum (G. transformans). III gall on Chamaecy-
paris thyoides; 0, I on chokeberry.
Gymnosporangiumglobosum. Hawthorn Rust, III general on eastem red-
cedar, also on dwarf, prostrate, and Rocky Mountain junipers; 0, I mostly on
hawthorn, also on apple, crabapple, pear and mountain-ash. Leaf galls on
cedar are very similar to those of common cedar-apple rust, but are smaller,
seldom over 1/2 inch, nearer mahogany red in color, and not perennial, pro-
RUSTS 549
ducing telial horns one season only. Apple and pear foliage may be slightly
affected but not the fruit; aecia are common on hawthorn pips.
Gymnosporangium gracile. III Witches Broom on juniper; 0, I on
hawthorn, quince, and shadbush.
Gymnosporangium asiaticum. III on leaves of Chinese juniper; 0, I on Chi-
nese owering quince and pear.
Gymnosporangium harknessianum. III on western juniper; 0, I on ame-
lanchier, chiey on fruits, sometimes stems. Papery margins of aecia are
usually long.
Gymnosporangium hyalinum. III on southern white-cedar; 0, I on haw-
thorn and pear leaves. Slight swellings are formed on small twigs and branch-
es of white-cedar.
Gymnosporangium inconspicuum. III on Utah juniper; 0, I on fruits, most-
ly of amelanchier and squaw-apple. Juniper leaves turn yellow; rarely telia
appear on branches.
Gymnosporangium japonicum (G. photiniae). III gall on stems of Chinese
juniper; 0, I on photinia.
Gymnosporangium juniperi-virginianae. Cedar-Apple Rust. III gener-
al on red cedar, eastern and southern, on prostrate and Rocky Mountain
junipers; 0, I general on apple and crabapple east of Great Plains. The fun-
gus is a native of North America and does not occur elsewhere. It is more
important commercially in the apple-growing regions of the Virginias and
Carolinas and certain states in the Mississippi Valley. It is important in many
areas on ornamental crabapples in home plantings.
The cedar apples or galls vary from 1/16 inch to over 2 inches across.
Leaves are infected during the summer, and by the next June a small, green-
ish brown swelling appears on upper or inner leaf surface. This enlarges until
by autumn the leaf has turned into a chocolate brown, somewhat kidney-
shaped gall covered with small circular depressions. The next spring in moist
weather orange telial horns are put forth from the pocketlike depressions.
The teliospores are enveloped in a gelatinous material that swells vastly,
a gall covered with horns sometimes reaching the size of a small orange.
They germinate in place to produce the basidiospores, which are carried by
wind to infect apple or other deciduous host.
By midsummer, apple leaves show yellow areas with amber pustules on
upper surface; but after pycnia have exuded drops of sticky liquid,they appear
as black dots in a rather reddish circle. On the undersurface of these spots
small cups are formed, with recurved mbriate margins. These aecia may
also appear near stem end of apples and are common on swollen twigs of
crabapple. Spores from these cups are blown back to the cedar in late sum-
mer, the entire cycle thus taking 2 years, 18 to 20 months on the cedar, 4 to
6 on the apple host.
Chief injury is to the apple host, the rust causing premature defoliation,
dwarng and poor-quality fruit. On very susceptible crabapples, such as
Bechtels crab, repeated infection may cause death of the branches or of the
entire tree. All our native crabapples are susceptible; most Asiatic varieties
are resistant.
Control. Care in planning is most important. Dont let your landscape archi-
tect or gardener put cedars and native crabapples or hawthorns close together.
Keep them separated as far as possible with a windbreak in between of some
tall nonsusceptible host. Some states have laws prohibiting red-cedars within
a mile of commercial apple orchards, but for practical garden purposes a few
hundred yards is sufcient, the danger markedly decreasing with distance,
especially with a house or hedge as a windbreak.
If junipers are already planted, it is possible in late winter to go over small
specimens and remove galls before spore horns are formed. Spraying in
spring inhibits telial development and germination of teliospores. Spray red-
cedars in August to prevent infection from crabapples.
Fairly resistant apple varieties are Baldwin, Delicious, Rhode Island and
North-western Greening, Franklin, Melrose, Red Astrachan, Stayman, and
Transparent. Avoid susceptible Jonathan, Rome, Wealthy, and York Imperi-
al. Most junipers susceptible to apple rusts are cultivars of Juniperus virgini-
ana and J. scopulorum. Many cultivars of J. chinensis and J. horizontalis are
resistant, and there are even some resistant forms of J. virginiana.
Gymnosporangiumkernianum. III on alligator, Utah, and western junipers;
0, I on amelanchier and pear. Telia arise between leaves on green twigs, but
mycelium is perennial in stems, causing dense witches brooms 6 to 18
inches in diameter.
Gymnosporangium libocedri. III on incense cedar; 0, I on leaves, fruits,
of amelanchier and hawthorn, also apple, crabapple, pear, quince, Japanese
quince and mountain-ash. Aecium is a cluster cup on foliage; telia are always
on leaves; witches brooms and swellings are produced on branches, rarely
on trunks. The fungus is said to persist in the mycelial stage up to 200
years.
Gymnosporangium multiporum. III on stems of western, one -seed, and
Utah juniper between leaves; 0, I unknown.
RUSTS 551
Gymnosporangium nelsonii. III on juniper and red-cedar; 0, I on leaves of
amelanchier.
Gymnosporangium nelsonii. III on one-seed, prostrate, Rocky Mountain,
Utah, and western junipers; 0, I on hawthorn, quince, Oregon crab, pear,
squaw-apple and Pacic mountain-ash. Galls are rm, woody, round, up to
2 inches in diameter.
Gymnosporangium nidus-avis. Witches Broom Rust. III on eastern and
southern red-cedars, on prostrate and Rocky Mountain junipers; 0, I on fruit,
young stems, leaves of apple, hawthorn, mountain-ash, quince, Japanese
quince, amelanchier or serviceberry. Trunks and branches of large trees have
witches brooms and long spindle-shaped swellings. Aecia are on both leaf
surfaces.
Gymnosporangium nootkatense. Gall Rust. II, III on Alaska cedar; 0, I
on mountain-ash, and Oregon crabapple. This is the only Gymnosporangium
species with uredial stage. Uredia are bright orange fading to pale yellow;
teliospores appear later in the same pustules. Aecia are cluster cups.
Gymnosporangium speciosum. III on alligator, one-seed, and Utah juni-
pers; 0,I on leaves of syringa (Philadelphus) and fendlera. Telia are in lon-
gitudinal rows on long fusiform swellings on juniper branches, which are
girdled and die. In severe infections the whole tree dies.
Gymnosporangium trachysorum. III on stem of eastern red-cedar; 0, I on
hawthorn leaves. Swellings on cedar are abruptly fusiform to globoid with
prominent telia 6 to 10 mm high.
Gymnosporangium tremelloides (G. juniperinum). III, stem gall on moun-
tain juniper; 0, 1 on Pacic mountain-ash. On smaller branches swellings
are subglobose galls up to 3/4 inch in diameter; hemispherical swellings on
larger branches are covered with attened telia.
Gymnosporangiumvauqueliniae. Witches BroomRust. III on one-seed
juniper; 0, I on Vauquelinia californica. This rust is the only Gymnospo-
rangium causing witches brooms on the aecial host.
Hyalopsora
Melampsoraceae. Telia on ferns, teliospores several-celled, in epidemis; urediospores of
two kinds, with pores.
Hyalopsora aspidiotus. Fir-Fern Rust. 0, I on balsam r; II, III on oak fern
(Phegopteris dryopteris). Pycnia are slightly raised orange-yellow spots on
needles; aecia are yellow to white, columnar, on 2-year needles.
Hyalopsora cheilanthis. Fir-Fern Rust. 0, I on balsam r; II, III on rock
brake, parsley fern, and cliff brake.
Hyalopsora polypodii. Fir-Fern Rust. General in northern and western
states on polypody fern and woodsia.
Kuehneola
Puciniaceae. Teliospores two- to many-celled; wall faintly colored or colorless.
Kuehneola malvicola. II, III on hibiscus and malvaviscus.
Kuehneola uredinis. Yellow Rust, Cane Rust. 0, I, II, III on blackberry,
dewberry, and raspberry. The disease appears to be increasingly prevalent,
especially on leaves, but there is a great difference in varietal susceptibility.
Eldorado, Foster,Jumbo, Lawton blackberries are highly susceptible; Nanti-
choke, Austin Thornless, Boysen Brainerd, Burbank Thornless, Jersey Black
are resistant. European varieties are generally resistant.
Kunkelia
Pucciniaceae. Pycnia subcuticular; telia subepidemal, caeomoid; teliospores catenulate,
one-celled.
Gymnoconia nitens (formerly Kunkelia nitens). Short-Cycle Orange
Rust of blackberry. I, general on blackberry but more common in the South
and West, also on dewberry and black,but not red raspberry. This is a peren-
nial rust, a systemic disease with only the aecial stage present. Underside
of leaves may be covered with quantities of orange-yellow spores. Remove
infected bushes.
Kunkelia nitens (see Gymnoconia nitens). Short-Cycle Orange Rust of
blackberry.
Melampsora
Melampsoraceae. Telia more or less indenite; teliospores sessile, subcuticular or
subepidermal, forming crusts of a single layer; aecia when present with rudimentary
peridium; uredia erumpent, pulverulent; spores globoid or ellipsoid, single on pedicels.
RUSTS 553
Figure 3.56 Teliospores. Melampsora, sessile in crust under host epidermis; Phragmidium, stalked, with several
cells; Puccinia, stalked, two-celled; Uromyces, stalked, one-celled
Species heteroecious when telia are on woody plants; autoecious if telia are on herba-
ceous plants (Fig. 3.56).
Malampsora abieti-capraearum. Fir-Willow Rust. 0, I on balsam, white,
and alpine rs; II, III on willows, widespread. Yellow spots on willow leaves
in early summer are followed by dark pustules when the telial stage is pro-
duced. There may be some defoliation.
Melampsora abietis-canadensis. Hemlock-Poplar Rust. 0, I on eastern
hemlock; II, III on various poplars. Cones have golden powdery masses of
spores over the surface; later shrivel, turn black, and hang as mummies; no
viable seed produced. Uredia are golden powdery pustules on undersurface
of poplar leaves; in late summer telia are formed in orange-yellow crusts that
change to black; in spring basidiospores reinfect hemlock.
Melampsora arctica. 0, I on saxifrage; II, III on willow.
Melampsora farlowii. Needle And Cone Rust of hemlock. 0, I unknown;
III on hemlock. Reddish slightly raised telia are on undersurface of needles,
shoots of the current year, and on cones. Young shoots may be twisted and
killed. Injury may occur in nurseries and in ornamental hedges.
Melampsora hypericorum (Mesopsora hypericorum). On St. Johnswort,
Montana.
Melampsora larici-populina. Rust; 0, I, on pine and larch; II, III, on poplar.
Melampsora medusae. Douglas-Fir Needle Rust. 0, I on Douglas-r,
big-cone spruce; II, III on native poplars. Pycnia are on upper surface of
current-year needles; aecia, of the caeoma type, are orange-yellow on the
undersurface. The rust is often epidemic on young trees but with little per-
manent ill effect.
Melampsora medusae. Larch Needle Rust. 0, I on larch in northeastern
states; II, III on native and introduced poplars except in far South.
Melampsora medusae f. sp. deltoidae. Rust; 0, I on Douglas r, pine and
larch; II, III, on poplar.
Melampsora occidentalis. Poplar Rust. 0, I unknown; II, III on native
poplars in the West.
Melampsora paradoxa (M. bigelowii). Larch-Willow Rust. 0, I on larch;
II, III on many species of willow. The damage to larch is insignicant. The
fungus winters on willow as mycelium in catkins, terminal buds, and young
stems and can maintain itself on willow in the uredial stage without larches.
Melampsora ribesii-purpureae. 0, I on currant, owering currant and
gooseberry; II, III on willow species.
Melampsorella
Melampsoraceae. Heteroecious on r, spruce, and dicotyledons; pycnia subcuticular,
aecia and uredia subepidermal, telia in epidemal cells. Only one species in United States.
Melampsorella caryophyllacearum (M. cerastii). Yellow Witches
Broom Rust. 0, I on many rs; II, III on chickweed. Infected evergreen
branches develop numerous upright lateral shoots from one point, forming
a compact witches broom; twigs are dwarfed, and needles turn yellow and
drop, leaving brooms bare. The fungus is perennial in stems, and shoots
develop with yellow leaves. Pycnia appear in raised orange spots on both
surfaces of dwarfed leaves in spring; aecia form in summer on underside,
in two rows of orange blisters. The disease is seldom serious enough for
control measures. In forest practice remove trees with main stem infections
early in life of the stand.
Melampsoridium
Melampsoraceae. Heteroecious, on larch and dicotyledonous shrubs and trees; pycnia
subcuticular; other sori subepidermal; teliospores sessile, one-celled.
Melampsoridium betulinum. Birch Leaf Rust. 0, I on larch; II, III on
birches. Uredia on underside of birch leaves are small reddish yellow pow-
dery pustules, followed later in summer by telia, rst waxy yellow, then dark
brown to nearly black.
RUSTS 555
Milesina
Melampsoraceae. Heteroecious on rs and ferns. All spores are colorless;
urediospores obovate or laceolate; teliospores in epidemal cells.
Milesina fructuosa. 0, I on balsam r; II, III on Dryopteris spp. Aecia are
white on current needles, maturing by midsummer.
Milesina laeviuscula. Needle Rust. 0, I on grand r; II, III on licorice fern,
in West.
Milesina marginalis. 0, I on balsam r; II, III on Dryopteris marginalis.
Pycnia are on both sides of needles, aecia of needles of current year, maturing
by midsummer.
Milesina pycnograndis (M. polypodophila). 0, I on balsam r; II, III on
Polypodium virginianum. Hyphae are perennial in needles and small stems
of balsam r; aecia on needles 3 to 9 years old.
Nyssopsora
Puccinaceae. Autoecious; teliospore with three cells.
Nyssopsora clavellosa. III on Aralia hispida.
Peridermium
A form genus with 0, I, on Gymnosperms. Aecia have peridia and are cylindrical,
tonguelike or bullate.
Peridermium bethelii. On dwarf mistletoe.
Peridermium ornamentale. 0, I on white, alpine, and noble rs.
Peridermium rugosum. 0, I on Pacic silver and lowland white rs.
Phakopsora
Melampsoraceae. Telia indehiscent, lenticular; spores formed in irregular succession,
not in chains.
Phakopsora cherimoliae. On cherimoya.
Phakopsora jatrophicola. On cassava.
Phakopsora pachyrhizi. On soybean.
Phakopsora zizyphi-vulgaris. On Zizyphus jujuba, Florida.
Phragmidium
Pucciniaceae. Autoecious. Pycnia subcuticular, other sori subepidermal; aecia caeo-
moid; teliospores large, conspicuous, of one to ten or more cells, each with two or
three lateral pores; walls somewhat layered, inner layer colored, outer nearly colorless,
smooth or verrucose; pedicel colorless except near spore; often swelling in lower por-
tion (see Fig. 3.56). Aecia with catenulate globoid or ellipsoid verrucose spores; uredia
when present circled with paraphyses; urediospores single on pedicels, walls verrucose
or echinulate with indistinct scattered pores.
Phragmidium americanum. 0, I, II, III on leaves of native and cultivated
roses. Teliospores with eight to eleven cells.
Phragmidium fusiforme (P. rosae-acicularis). 0, I, II, III on several hosts
species. Teliospores with ve to eleven cells, walls chocolate brown, verru-
cose.
Phragmidium montivagum. 0, I, II, III on many species of roses. Telio-
spores with six to nine cells.
Phragmidium mucronatum (P. disciorum). Leaf Rust of Rose. 0, I on
leaves and stems; II, III on leaves of cultivated roses, eastern states to the
Rocky Mountains and on the Pacic Coast. This is the common rust of hybrid
teas and other roses with large, rm leaets. It is not much of a problem in
the East, although sometimes found in New York and New England gar-
dens, but it is a serious menace along the Pacic Coast. Aecia appear on
leaves as small, roughly circular spots, 1/25 inch across, bright orange on the
underside of leaf, from the spore masses, light yellow on the upper surface,
sometimes bordered with a narrow green zone. Leaf lesions may be slightly
cup-shaped viewed from the upper surface. Stem lesions are long and narrow.
The summer uredial stage has reddish orange spores in very small spots,
1/50-inch, over underside of leaves. This stage may repeat every 10 to 14
days in favorable weather, with wilting and defoliation. In mild climates
the uredial stage continues; in cooler areas the telial stage is formed toward
autumn black pustules of stalked dark spores, rough, with a point, ve to
nine cells.
The leaf surface must be continuously wet for 4 hours for rust spores to
germinate and enter the leaf; this means liquid water and not high humid-
ity as with mildews. High summer temperatures adversely affect infection,
RUSTS 557
summer spores retaining viability for only a week at 80F. In southern Cali-
fornia temperatures are uniformly favorable for rose rust, and from October
to April there is sufcient rainfall. In drier months fog may provide requisite
moisture.
Control. Removing infected leaves during the season and all old leaves left
at the time of winter or early spring pruning may be somewhat helpful.
Phragmidiumrosae-arkansanae. 0, I, II, III on Rosa arkansana and R. suf-
fulta. Teliospores with ve to eight cells.
Phragmidium rosae-californicae. 0, I, II, III on many rose species. Telio-
spores with eight to eleven cells.
Phragmidium rosicola. III on Rosa engelmanii and R. suffulta. Teliospores
one-celled, nearly round.
Phragmidium rubi-idaei. Leaf and Cane Rust of raspberry; Western
Yellow Rust, general but important only in the Pacic Northwest. 0, I, II,
III on red rasberries, sometimes black but not on blackberries. Small, light
yellow spore pustules appear in young leaves, with black teliospores follow-
ing in the same spots later in the season. Deep, cankerous lesions are formed
on canes in the fruiting year, Cuthbert variety being particularly suscepti-
ble. Spring infection probably comes from sporidia formed in telia on fallen
leaves. A dormant spray may be helpful, along with cleaning out infected
canes at winter pruning.
Phragmidiumspeciosum. 0, I on stems and leaves, III on stems of cultivated
and native roses, throughout United States except far South.
Phragmidium subcorticium. Obsolete name. Some specimens former-
ly recorded as this species belong to P. mucronatum, others to P. rosae-
pimpinellifoliae.
Phragmidium tuberculatum. On Rosa sp. Connecticut and Alaska.
Phragmopyxis
Pucciniaceae. Teliospores colored, two- to many-septate; wall three-layered, the middle
layer swelling in water; aecia, uredia, and telia with a border of paraphyses.
Phragmopyxis acuminata. 0, III on Coursetia.
Physopella (Angiopsora)
Pucciniaceae. Only uredia and telia known. Telia indehiscent, lenticular;
teliospores in chains.
Physopella ampelopsidis (Phakopsora vitis). On ampelopsis and grape,
Florida.
Physopella compressa. On paspalum, southern ornamental grass.
Pileolaria
Pucciniaceae. Autoecious, on members of family Anacardiaceae. Teliospores stipitate,
dark, with pores, one-celled; pycnia subcuticular; uredia present.
Pileolaria cotini-coggyriae. On smoke tree.
Pileolaria patzcuarensis. 0, I, II, III on sumac.
Prospodium
Pucciniaceae. Autoecious on Bignoniaceae and Verbenaceae in warm climates.
Prospodium appendiculatum. On tecoma, Florida, Texas.
Prospodium lippiae. On lippiae, Arizona.
Prospodium plagiopus. On tabebuia, Florida.
Prospodium transformans. On tecoma, Florida.
Puccinia
Pucciniaceae. A very large genus, comprising nearly half of all known rusts; autoecious
and heteroecious. Teliospores smooth, two-celled with apical pores, rm pedicels, col-
ored; aecia cluster cups with peridium (see Fig. 3.56). The species listed here are a small
selection of those on garden plants; others are listed in host section.
Puccinia acroptili. Rust on Centaurea.
Puccinia allii (P. porri). Autoecious on onion, garlic and shallot, but 0, I
stages rare. Occasional on cultivated onion, more common on garlic, wild
garlic, and wild onion. Uredia are yellowish, telia black.
Puccinia amphigena (Aecidium yuccae). On yucca.
Puccinia andropogonis, with various strains. 0, I on lupine, Indian paint-
brush and turtlehead; II, III on andropogon.
Puccinia antirrhini. Snapdragon Rust. II, III general on snapdragon, also
on linaria, corydylanthus; 0, I unknown. Pustules of spores on underside of
leaves are chocolate brown, often in concentric circles (see Fig. 3.57). The
RUSTS 559
Figure 3.57 Rust on Snapdragon
area over the pustule is pale or yellow on upper surface. Spores also appear
on stems; there is a drying and stunting of whole plant. The rust is spread by
wind-blown spores and on cuttings. For infection, plants need to be wet with
rain or dew 6 to 8 hours with day temperatures around 70 to 75F. Spores
are killed above 94F. There are at least two races.
Control. Purchase only rust-resistant variety. Bordeaux mixture controls sec-
ondary fungi following rust but not the rust itself. Sulfur dust is still useful,
or a spray made by adding 1 ounce rosin soap to a gallon of water and then
adding 1 ounce dry lime sulfur.
Puccinia arachidis. Peanut Rust, occasional in Alabama, Florida, Texas.
Puccinia aristidae and varieties. II, III on wild grasses, Aristides and Dis-
tichlis; 0, I on eriogonum, greasewood, beet, spinach, western wallower,
garden cress, radish, California bluebell, heliotrope, cleome, primrose, sand-
verbena, and others.
Puccinia asparagi. Asparagus Rust. II, III general on susceptible vari-
eties; 0, I not reported in natural infections. Also on onion. Asparagus rust
reached America in 1896 from Europe and spread with devastating sudden-
ness from Boston and New Jersey to California, reaching there by 1912, one
of the fastest cases of disease spread in our history. If tops are attacked sev-
eral years in succession, the root system is so weakened that shoots fail to
appear in spring or are culls.
The rst symptom is a browning or reddening of smaller twigs and needles,
with the discolored area spreading rapidly until the whole planting looks as
if it had ripened prematurely. The reddish color is due to numerous small
pustules of urediospores that give off a dusty cloud when touched. These
appear in successive generations until autumn, or a spell of drought, when
they are replaced by black teliospores, either in the same or a new fruiting
body. They remain on old stems until spring, germinating then to infect new
shoots as they emerge from the ground.
Control. For a long time resistant varieties Mary Washington and Martha
Washington were the answer to the rust problem, but the fungus has devel-
oped resistant strains. Waltham Washington, Seneca Washington, and Cal-
ifornia 500 have some resistance. Clean up volunteer or wild asparagus
around beds. A parasitic fungus, Darluca lum, helps keep rust in check.
Puccinia brachypodii var. poae-nemoralis (formerly Puccinia poae-
nemoralis (Syn. P. poae-sudeticae)). Bluegrass Leaf Rust, Yellow Leaf
Rust. II, III on turf grasses, mostly Canada and Kentucky bluegrass; 0,
I, unknown; general east of the Rocky Mountains. The uredia are orange-
yellow with numerous peripheral paraphyses. Telia are covered rather per-
manently with epidermis; spores are dark brown with short pedicels. The
wheat stem rust is more important on Merion bluegrass.
Puccinia calcitrapae var. centaureae (formerly Puccinia carthami). Widely
distributed on safower in Great Plains and California. Spores carried on
seed or persisting in soil infect seedlings, which often die.
Puccinia canaliculata. Rust on purple nutsedge and yellow nutsedge.
Puccinia carduorum. Rust on Carduss tenniorus and Carduss thoermeri.
Puccinia caricina (P. caricis var. grossulariata, P. pringsheimia). 0, I on
currant, owering currant, gooseberry; II, III on Carex spp. Common only on
RUSTS 561
wild species or in neglected gardens. Leaves are thickened, sometimes curled
in reddish cluster cup areas; there are enlargements on stems and petioles, red
spots on berries. Control by eliminating the sedge host.
Puccinia carthami (see Puccinia calcitrapae var. centaureae). Widely dis-
tributed on safower in Great Plains and California.
Puccinia claytoniicola. On claytonia, Wyoming.
Puccinia conoclinii. On ageratum, Ohio.
Puccinia coronata. Crown Rust of oats; Orange Leaf Rust of Oats. 0, I on
buckthorn and rattan vine; II, III on oats and grasses. There are several vari-
eties and many physiological races of this rust, which is as destructive to oats
as leaf rust is to wheat. Redtop, meadow fescue, ryegrass, and bluegrass are
among the lawn grasses that may show orange or black pustules on leaves.
Puccinia crandallii. 0, I on snowberry, wolfberry, coralberry; II, III on grass-
es, fescues, bluegrass.
Puccinia cynodontis. On Bermuda grass, New Mexico.
Puccinia cypripedii. On orchids.
Puccinia dioicae (P. extensicola) in many varieties. 0, I on aster, golden-
rod, erigeron, senecio, lettuce, oenothera, rudbeckia, and helenium; II, III on
Carex spp.
Puccinia dracunculi (see Puccinia tanaceti var. dracunculina). On artemi-
sia, Wisconsin to the Pacic Coast.
Puccinia averiae (see Puccinia melampodii). On Calendula.
Puccinia graminis. Stem Rust of grains and grasses. 0, I on barberry and
mahonia, especially in north central and northeastern states; II, III on wheat
and other cereals and wild and cultivated grasses.
This is the classic example of rust, the one used in school textbooks and
known through the ages as the major limiting factor of wheat production.
Proof of the connection between barberry and wheat in the life cycle was not
made until 1864, but long before that farmers had noticed that wheat suffered
when barberry plants were near. France in 1660, Connecticut in 1726, and
Massachusetts in 1755 enacted laws requiring the destruction of barberry
near grain elds.
There are six commonly recognized varieties of stem rust:
Puccinia graminis f. sp. avenae on oats, sweet vernal grass, brome grasses,
some fescues.
P. graminis f. sp. agrostidis on redtop and other Agrostis spp.
P. graminis f. sp. graminicolaon St. Augustine grass.
P. graminis f. sp. phlei-pratensis on timothy and some related grasses.
P. graminis f. sp. poae on Kentucky and other bluegrasses.
P. graminis f. sp. secalis on rye, some wheat, and barley grasses.
P. graminis f. sp. tritici, wheat rust on wheat, barley, rye, and many grasses.
Stem rust occurs wherever wheat is grown, but is most serious in northern
states. It is dependent on weather conditions, with epidemics and disastrous
losses in certain seasons. The amount depends on the maturity of the crop
when rust strikes, but losses may run 25% of expected yield for the nation
and much higher for individual states. There are a great many physiological
races.
On grains and grasses the rst rust appears as long, narrow streaks on stems,
leaf sheaths, leaf bases, and distal portions of blades. These streaks are ure-
dial sori, the epidermis being torn back to form a white collar around a dark
red powdery mass of one-celled urediospores. Later the same sori turn black
as dark, two-celled teliospores replace summer urediospores. Stems may be
broken at this stage.
The summer spores appear about 10 days after infection. This stage can be
repeated, the spores reinfecting wheat, and, since they are carried by wind
from one plant to another, one state to another, even to hundreds of miles,
they account for large outbreaks of disease. In Mexico and southern Texas
this II stage continues through the winter and causes spring infection with-
out the intervention of barberry. Waves of urediospores coming up from the
South may start northern infection.
Normally in the North, spring infection starts on barberry from sporidia
(basidiospores) produced on a promycelium put forth by a teliospore win-
tered on a wheat stem. Two sexes occur in this rust, designated + and rather
than male and female. A young teliospore contains two nuclei, one + and the
other ; as the spore matures, these fuse to a single nucleus, which divides
twice in the production of the four-celled basidium (promycelium). Each
cell produces a sporidium; two of these are + and two . A sporidium falling
on a barberry leaf germinates, sends in an infection thread, and develops
a mononucleate (haploid) feeding mycelium and nally a ask-shaped pyc-
nium containing pycniospores, which correspond to the sex of the sporidium
starting infection. The pycnia are in reddish lesions on the upper leaf surface.
Hyphal threads, receptive hyphae, extend through the mouth of the pycni-
um. Aided by insects, which are attracted by a sweet nectar, pycniospores
(spermatia) of one sex are brought into contact with receptive hyphae of the
opposite sex, and sexual union takes place, without which there is no further
development of the rust.
RUSTS 563
The dicaryotic or binucleate mycelium formed from the fertilized hypha
grows through the cells of the barberry leaf and masses together on the under-
side to produce aecia lled with a yellowish waxy layer of aeciospores in
cluster-cup formation. These spores, unable to reinfect barberry or mahonia
are wind-borne to the cereal or grass host, the subsequent mycelium contin-
uing binucleate until the fusion in the teliospore. New crops of urediospores
can be produced every 10 to 14 days.
Control. Resistant varieties are of primary importance, but they are difcult
to maintain because the sexual process in rusts allows the continuous devel-
opment of new strains. More than 200 strains are known, but only a dozen
or so are important in any one year. Race 15B is prevalent most years and
can attack all varieties of wheat grown in this country. Eradication of the
barberry eliminates the alternate host and also the breeding place of new rust
varieties. Most barberry and mahonia species are under quarantine, but some
have been designated rust-resistant by the U.S. Department of Agriculture
and may be shipped interstate under permit.
Puccinia helianthi. Sunower Rust. 0, I, II, III general on sunower,
Jerusalem artichoke, and heliopsis. Numerous brownish pustules in which
repeating spores are formed develop on underside of leaves, which may dry
and drop.
Puccinia heterospora. III on abutilon, hollyhock, mallow, and malvavis-
cus.
Puccinia heucherae. III on coral bells, woodland star, saxifrage, bishops-
cap, and foam-ower.
Puccinia hieracii. 0, I, II, III widespread on endive and hawksbeard. Endive
leaves are spotted and blighted with dusty spore pustules. The crop is occa-
sionally lost, but no control has seemed practical.
Puccinia horiana. White Rust. III, IV on chrysanthemum; no alternate host
known. First reported in England in 1964; became widespread there in 1976.
Found in amateur chrysanthemum plantings in New Jersey and Pennsylvania
in 1977.
Puccinia iridis. Iris Rust. 0, I, II, III on bulbous iris, serious in the South-
east, uncommon in Northwest. Small, oblong to oval, red or dark brown pow-
dery spots, often surrounded by a yellow margin, are present on leaves and
stems, which may die prematurely. In inoculation tests with Dutch iris, vari-
eties Early Blue, Gold and Silver, Golden West, Imperator, Lemon Queen,
and Texas Gold were resistant.
Puccinia jaceae var. diffusa. Rust on Centaurea.
Puccinia lagenophorae. On English daisy.
Puccinia malvacearum. Hollyhock Rust. III general on hollyhock, also on
mallow, and lavatera. This rust is so common and destructive it limits the use
of hollyhocks as ornamentals. Stems, leaves, bracts may be attacked. There
are yellow areas on the upper surface of leaves, orange-red spore pustules
on the underside, and elongated lesions on stems. Spore pustules are some-
times grayish from formation of sporidia, but the alternate host is unknown.
In severe infections leaves dry and hang down along the stem. The fungus
winters in pustules in basal leaves and in old stems.
Control. Cleaning up all infected plant parts in fall and again very early in
spring is most important; infection starts early in the season, and once it is
under way, it is very difcult to curb with a fungicide.
Puccinia melampodii (formerly Puccinia averiae). On Calendula.
Puccinia melampodii. On Baccharis, Texas.
Puccinia menthae. Spearmint Rust. 0, I, II, III on spearmint, pepper-
mint, oregano, also horse-mint, mountain-mint, dittany, bee-balm, yerba bue-
na, and germander; especially serious for mint farmers in Middle West and
Northwest. In spring and early summer the disease appears as light yel-
low to brown raised spots on deformed stems and leafstalks, sometimes on
main veins; golden to chocolate brown spots appear in late summer and fall.
Affected leaves dry, and the yield of oil is reduced. The pathogen has at least
10 races. Dusting with sulfur and early cutting are recommended.
Puccinia nakanishikii. Rust on lemon grass.
Puccinia pelargonii-zonalis. Pelargonium Rust. The uredinial stage of
a rust, presumably this species, was found on geranium in New York and
California in 1967. It has now been reported in Pennsylvania and Florida.
Brown spore pustules appear on leaves, petioles, and stems; leaves turn yel-
low and drop. Destroy infected plants.
Puccinia phragmitis. 0, I on rhubarb; II, III on reed grass, sometimes
present in California but not serious. Aecia are white, on underside of
rhubarb leaves, surrounded by pycnia.
Puccinia poae-nemoralis (Syn. P. poae-sudeticae) (see Puccinia brachy-
podii var. poae-nemoralis). Bluegrass Leaf Rust, Yellow Leaf Rust. II,
III on turf grasses, mostly Canada and Kentucky bluegrass; 0, I, unknown;
general east of the Rocky Mountains.
Puccinia polygoni-amphibii. Rust; II, III, on jointweed.
Puccinia polysora. Southern Corn Rust. 0, I, unknown; II, III on corn and
grasses. Present in the South, requiring higher temperatures than common
RUSTS 565
corn rust; not very important. Urediospores are yellow to golden, teliospores
chestnut brown, angular; often parasitized by Darluca lum.
Puccinia psidii. Rust on allspice (Pimenta dioica) Melaleuca quinquen-
ervia, and Syzygium jambos.
Puccinia pygmaea. Rust on grasses.
Puccinia recondita (P. rubigo-vera). Leaf Rust of cereals and grasses, with
several varieties:
P. recondita tritici (P. triticina). II, III on wheat (but not grasses); 0, I on
meadow rue. This rust is worldwide and more serious than stem rust in the
southern half of the American wheat belt, sometimes epiphytotic with loss-
es up to 30%. The leaf tissue is progressively destroyed through the sea-
son, resulting in a reduced number of kernels, shriveled grain, low weight
and protein content. Rust pustules breaking through the epidermis greatly
increase transpiration losses. Orange uredial pustules are followed later by
gray telial sori, but urediospores are the effective spore form and can survive
southern winters. There are many physiological races.
P. recondita agropyri. II, III on wheat grasses and wild ryegrasses; 0, I on
clematis, buttercup, columbine, larkspur, and other Ranuculaceae. Common
in Rocky Mountain area.
P. recondita agropyrina. Similar to the above but occurring outside moun-
tainous areas.
P. recondita apocrypta. II, III on wheat and wild grasses; 0, I on waterleaf
and mertensia.
P. recondita impatientis. II, III on redtop and related grasses; 0, I on touch-
me-not.
P. recondita secalis. II, III on rye; 0, I on bugloss (Lycopsis).
Puccinia solheimi. On dodocatheon, Wyoming.
Puccinia sorghi. Corn Rust. 0, I on oxalis; II, III on corn, sweetcorn, gen-
eral in northeastern and north central states. Cinnamon brown spore pustules
cover both leaf surfaces with black pustules toward autumn. The disease is
not often serious enough for control measures.
Puccinia sparganioides (P. peridermiospora). Ash Rust. 0, I, general on
ash east of the Great Plains; II, III on marsh and cord grasses (Spartina spp.).
Ash twigs and petioles are swollen and leaves distorted. Cluster cups lled
with yellow powdery aeciospores are formed in the swellings. In New Eng-
land, where rust is often severe, the most important infection period on ash
is May 15 to June 20, with 6 to 8 hours of damp air necessary. Marsh grasses
are infected and reinfected July 20 to August 20.
Puccinia stenotaphricola. On St. Augustine grass, Florida.
Puccinia striiformis (P. glumarum). Stripe Rust of wheat. II, III on wheat,
barley, rye, redtop, orchardgrass, and many other grasses. Uredial stage is
yellow, and pustules are formed in streaklike clusters on leaves; telia are in
black streaks.
Puccinia substriata. Rust on eggplant.
Puccinia taneceti. Chrysanthemum Rust. II general; III known only in
Japan; 0, I unknown. Small blisters of pinhead size appear on underside of
leaves and occasionally on upper surface. The spore mass is dark reddish
brown and powdery. The rust is more common in greenhouses than outdoors.
Optimum germination is at 60 to 70F; spores are killed at high tempera-
tures.
Puccinia tanaceti var. dracunculina (formerly Puccinia dracunculi). On
artemisia, Wisconsin to the Pacic Coast.
Puccinia thaliae (P. cannae). II, III on edible canna, garden canna,and
maranta.
Pucciniastrum
Melampsoraceae. Heteroecious with perennial mycelium, pycnia and aecia on conifers:
rs and spruces; pycnia subcuticular, other sori subepidermal; telia may be intraepider-
mal; aecia and urediospores yellow.
Pucciniastrum americanum. Late Leaf Rust of raspberry. 0, I on white
spruce; II, III on red raspberry, not black. This rust appears late in the season
on Cuthbert and other susceptible varieties, in northern half of the country,
most common east of the Mississippi. Fine light yellow powdery masses of
spores appear on basal leaves, leaf petioles, shoots, and even fruit.
Pucciniastrum epilobii. Fuchsia Rust, the alternate hosts are species of
Abies.
Pucciniastrum goeppertianum. Fir-Huckleberry Rust, Blueberry
Witches Broom. 0, I on rs; III on low and high bush blueberries. The fun-
gus is systemic and perennial in blueberries, producing short swollen twigs
in a witches broom effect, and telia forming a polished red layer around the
shoots. Destroy diseased bushes; keep blueberry plantations some distance
from rs.
Pucciniastrum hydrangeae. 0, I on eastern and Carolina hemlock; II, III on
hydrangea.
RUSTS 567
Pucciniastrum vaccinii (P. myrtilli). Hemlock Rust, Leaf Rust of blue-
berry; widespread. 0, I on eastern hemlock; II, III on azalea, blueberry, cran-
berry, lyonia, menziesia, and rhododendron. This is the most common hem-
lock rust, but often only a single leaf or twig is infected. Aecia are formed on
current-year needles. Blueberries have yellow pustules, on leaves only, with
defoliation in mid- or late summer.
Ravenelia
Pucciniaceae. Autoecious, tropical with only a few species in United States. Teliospores
more or less muriform, with compound stalks.
Ravenelia dysocarpae (see Ravenelia fragrans var. evernia). On Mimosa,
Arizona.
Ravenelia fragrans var. evernia (formerly Ravenelia dysocarpae). On
Mimosa, Arizona.
Ravenelia humphreyana. On Poinciana, Florida, Texas.
Ravenelia indigoferae. On Indigofera, Arizona.
Maravalia (Scopella)
Pucciniaceae. Tropical. Uredia and telia subepidermal. Teliospores one-celled, on pedi-
cel.
Maravalia sapotae (formerly Scopella sapotae, Syn. Uredo sapotae). On
sapodilla in Florida, infecting leaves in winter and early spring.
Scopella sapotae, Syn. Uredo sapotae (see Maravalia sapotae). On sapodil-
la in Florida, infecting leaves in winter and early spring.
Sphenospora
Pucciniaceae. Tropical. Telia and peridia subepidermal, then erumpent; teliospores
waxy, two-celled, on pedicel.
Sphenospora mera. On bletilla, Florida.
Sphaerophragmium
Pucciniaceae. Teliospores stalked, four- to several-celled, with transverse and horizontal
septa; on legumes.
Sphaerophragmium acaciae. On lebbek, Florida.
Tranzschelia
Pucciniaceae. Teliospores two-celled, stalked; uredia with pseudoparaphyses; on Ranun-
culaceae and Prunus.
Tranzschelia discolor (T. pruni-spinosae var. discolor). Rust of stone fruits.
Peach Rust. 0, I on Anemone coronaria; II, III on apricot, peach, plum,
prune, almond, and cherry, in late summer. Yellow angular spots appear on
leaves with powdery spore pustules on underside, reddish on peach, dark
brown on almonds; sometimes with late season defoliation. Peach fruit may
have round sunken green spots; twigs may have oval blisters in early spring.
Urediospores wintering on sucker shoots can start spring infection without
the alternate host. The Drake variety of almond is most susceptible.
Tranzschelia pruni-spinosae var. typica. 0, I on anemone, hepatica, thalic-
trum, and buttercup; II, III on wild species of Prunus.
Triphragmium
Pucciniaceae. Teliospores stalked, with three cells forming a triangle, each with a single
pore.
Triphragmium ulmariae. 0, I, II, III on meadowsweet.
Uredinopsis
Melampsoraceae. Telia on ferns; teliospores scattered irregularly in mesophyll, rarely in
subepidermal crust, typically several-celled; aecia white.
Uredinopsis osmundae. Fir-Fern Rust. 0, I on balsam r, widespread; II,
III on Osmunda spp.
RUSTS 569
Uredinopsis phegopteridis. Fir-Fern Rust. 0, I on balsam r; II, III on
Phegopteris dryopteris.
Uredinopsis pteridis (U. macrosperma). Fir-Fern Rust. 0, I on various rs;
II, III on Pteridium aquilinum. Aecia are on 1- to 5-year needles of Pacic
silver, white, lowland white, alpine, and noble rs.
Uredinopsis struthiopteridis. Fir-Fern Rust. 0, I on balsam, lowland
white, alpine, and noble rs; II, III on ostrich fern.
Uredo
Form genus; uredia with or without peridia.
Uredo artocarpi. Breadfruit in Hawaii.
Uredo coccolobae. On sea-grape, Florida.
Uredo ericae (Pucciniastrum ericae). On erica, California.
Uredo phoradendri. On mistletoe.
Uromyces
Pucciniaceae. Like Puccinia but teliospores with one cell, yellow to dark; aecia when
present with a persistent peridium (see Fig. 3.56).
Uromyces appendiculatus. Bean Rust. 0, I rare on bean; II, III general
on dry beans, widespread but infrequent on lima bean, scarlet runner bean.
This is the true bean rust, an old disease reported as far back as 1798 and
quite distinct from anthracnose that is sometimes called rust. It is particularly
serious and prevalent on Kentucky Wonder pole beans.
Small rust pustules are formed on leaves most frequently, sometimes on
stems and pods. The reddish brown sori are most numerous on underside
of leaves, with the upper surface yellowing in the same areas. There may be
nearly complete defoliation. In late summer in the North, dark telia replace
summer spores, but in the South, urediospores survive the winter and start
early spring infection. Rust spores are spread by wind and on tools and cloth-
ing. Some even cling to supporting poles and can start a fresh outbreak of rust
if poles are not disinfested before reuse.
Control. No bean variety is resistant to all of the more than 30 races so far
identied. Most snapbeans are highly tolerant of rust; and pole beans White
Kentucky Wonder, U.S. 4 Kentucky Wonder, Potomac, and Rialto are fairly
tolerant.
Uromyces appendiculatus var. appendiculatus (Syn. U. phaseoli). Rust
on bean.
Uromyces ari-triphylli. On jack-in-the-pulpit; Autoecious, O, I, II, III
stages (entire ife cycle) on one host.
Uromyces betae. Beet Rust. II, III on beets, and swiss chard, in Califor-
nia, Oregon, occasionally Arizona and New Mexico. Reddish brown pustules
may be numerous on foliage in late summer or in wet seasons. Control is sel-
dom attempted for table beets; some sugar beet varieties are resistant. The
seed-borne fungus also persists in volunteer plants and debris.
Uromyces ciceris-arietini. Rust on chickpea.
Uromyces costaricensis. Rust on wild bamboo.
Uromyces dianthi (U. caryophyllinus). Carnation Rust. 0, I on euphorbia
(but not in United States); II, III general on carnation and sweet william,
a serious disease under glass. Chocolate brown pustules, varying from 1/16
to 1/4 inch, break out on both sides of leaves and on buds and stems. Leaves
curl up, often die; infected plants are stunted.
Control. Use surface watering where possible, avoiding syringing; keep
greenhouses properly ventilated; use rust-free cuttings.
Uromyces fabae. Pea Rust. 0, I, II, III on pea, peavine, occasionally on
broad bean; not very serious.
Uromyces galii-californici. On galium, California.
Uromyces punctatus. Rust on Astragulus in ID and OR.
Uromyces trifolii, in several varieties. 0, I, II, III on clovers. Pale brown
pustules surrounded by torn epidermis, appear on underside of leaves and on
petioles and stems.
Uromyces sp. Rust on birdsfoot trefoil.
Uropyxis
Pucciniaceae. Autoecious. Teliospores two-celled, on pedicels; uredia with paraphyses.
Uropyxis daleae var. eysenhardtiae (formerly Uropyxis eysenhardtiae). On
Dalea and Eysenhardtia in Arizona.
Uropyxis eysenhardtiae (see Uropyxis daleae var. eysenhardtiae). On
Dalea and Eysenhardtia in Arizona.
SCAB
Diseases characterized by an overgrowth of tissue in a limited area are
commonly called scab. The hyperplastic scablike lesions correspond to the
necrotic or dead areas of leaf spots and cankers. Diseases called scab caused
by Elsino or its anamorph, Sphaceloma, are treated under Spot Anthrac-
nose.
Cladosporium
Blotch Diseases.
Cladosporium bruneo-atrum. Possible cause of russeting of citrus fruit
hitherto attributed solely to citrus mite.
Cladosporium carpophilum (Syn. Fusicladium carpophilum), apparently
a different strain from peach scab fungus. Almond Scab. Water-soaked
symptoms on young shoots turn brown; leaves turn black, drop prematurely;
circular, olivaceous spots coalesce on fruit.
Cladosporium carpophilum. Peach Scab, general on peach, widespread
on apricot, nectarine, cherry, and plum. The form on cherry and European
plum has been attributed to Venturia cerasae (Cladosporium cerasi). Small,
round, olive black spots appear on infected fruits about 6 weeks after petals
have fallen. These are usually on upperside of fruit, and cracking may follow.
Twigs show nearly circular yellow-brown blotches with gray or bluish bor-
ders; cambium may be killed and twig die. Leaf spots are brown, scattered,
with tissue drying and falling out, leaving circular holes.
Control. The brown-rot spray schedule should also control scab, a sulfur
spray 4 to 6 weeks after petal fall being especially important. A fungicide
can be combined with an insecticide spray for curculio.
Cladosporium caryigenum. Pecan Scab, Leaf Spot, general on pecan,
and hickory. Scab is perhaps the most important limiting factor in pecan
production in the Southeast. All varieties are somewhat susceptible, even
those, like Stuart, that have been quite resistant in the past. Crop losses may
reach 75 to 95%.
The fungus attacks rapidly growing tissue in leaves, shoots, and nuts; mature
growth seems to be immune. On Schley and other highly susceptible vari-
eties, primary infection shows in elongated olive brown lesions on veins and
underside of leaves. With secondary infection leaves appear almost black, as
a result of coalescing of spots; defoliation follows. On more resistant vari-
eties, such as Moore and Stuart, infection is often delayed until the leaves
are nearly mature, and so scab spots are conned to nuts. Nut lesions are
small, black, circular, slightly raised at rst, then sunken. Spots may be so
close together that the entire surface turns black; the nuts drop prematurely or
remain attached to shoots indenitely. Infection is correlated with spring and
early summer rainfall, continuous moisture for 6 to 8 hours being required
for the spores to germinate and enter the host. First lesions appear in 1 or 2
weeks.
Control. Knock off old shucks and leaf stems before trees leaf out in spring.
When they are wet after a rain, a slight jarring of branches will make such
diseased material drop. Prune off low limbs for better air circulation. Four
protectant sprays are required in Georgia, ve in Florida.
Cladosporiumcladosporioides f. sp. pisicola (formerly Cladosporium pisi-
cola). Pea Scab, Black Spot of pea. Dark spots covered with velvety mold
are formed in moist weather on leaves, stems, where black streaks may devel-
op into cankers and pods may be distorted. The fungus is seed-borne, and
lives in soil in plant refuse.
Cladosporium coreopsidis. Reported on coreopsis in Wisconsin, causing
stunting and suppression of owering.
Cladosporium cucumerinum. Cucumber Scab, general on cucumber in
greenhouses, an important transit and storage decay of muskmelon, some-
times serious on late-planted squash. The disease was rst noted in New
York in 1887. Leaves with water-soaked spots may wilt, stems have slight
cankers, but most injury is to the fruit. First symptoms, while cucumbers are
still small, are gray, slightly sunken spots, sometimes exuding a gummy sub-
stance. They darken with age, and the collapsed tissue forms a pronounced
cavity, lined with a dark green velvety layer of greenish mycelium, short
conidophores, and dark, one- to two-celled spores. On leaves, these fruit-
ing fascicles are extruded through stomata. The disease becomes epidemic
after mid-summer, when night temperatures are cold or with heavy dews and
fog.
SCAB 573
Control. Resistant cucumber varieties include Maine No. 2, Wisconsin SR
10, SR 6, and Highmoor. A long rotation is advised.
Cladosporium pisicola (see Cladosporium cladosporioides f. sp. pisicola).
Pea Scab, Black Spot of pea.
Fusarium
Rots.
Fusarium heterosporum. Head Scab of tall fescue.
Spilocaea (Fusicladium)
Leaf Spots.
Fusicladium dendriticum (see Spilocaea pomi). Conidial stage of the
apple-scab fungus.

Venturia inaequalis.
Fusicladium eriobotryae (see Spilocaea pyracanthae). Loquat Scab,
widespread on leaves, stems, fruit of loquat.
Fusicladium photinicola (see Spilocaea photinicola). Christmasberry
Scab on Photinia arbutifolia.
Fusicladium pyracanthae (see Spilocaea pyracanthae). Pyracantha
Scab, widespread on leaves and fruit.
Fusicladium saliciperdum (Syn. Venturia chlorospora) (see Pollaccia
saliciperda). Willow Scab, Blight, rst noticed on willow in Connecticut
in 1927, apparently introduced from Europe.
Pollaccia saliciperda (formerly Fusicladium saliciperdum (Syn. Venturia
chlorospora). Willow Scab, Blight, rst noticed on willow in Connecti-
cut in 1927, apparently introduced from Europe. Repeated defoliation has
killed thousands of trees in the Northeast. Young leaves are attacked and
often killed in spring, almost within a few hours, and from the leaf blades
the fungus enters twigs, kills back young shoots, and causes cankers. Olive
green felty spore masses are formed on the long veins on underside of leaves.
Overwintering is as dormant mycelium in twigs infected the previous spring.
Another fungus, Physalospora miyabeana, is found with the scab fungus,
and the two together form the disease complex known as willow blight.
Physalospora usually attacks later in the season than Fusicladium and causes
cankers on larger stems.
Control. Prune heavily to remove diseased parts. Spray with bordeaux with
excess lime.
Spilocaea photinicola (formerly Fusicladium photinicola). Christmasber-
ry Scab on Photinia arbutifolia. Brown velvety spots appear on leaves,
ower stalks, and green berries; the berries being disgured when mature.
Prune in winter to remove dead wood and foliage. Spray before blossoming
with bordeaux mixture.
Spilocaea pomi (formerly Fusicladium dendriticum). Conidial stage of the
apple-scab fungus.

Venturia inaequalis.
Spilocaea pyracanthae (formerly Fusicladium eriobotryae). Loquat Scab,
widespread on leaves, stems, fruit of loquat. This is similar to pear and apple
scab. Dark velvety spots cause more or less deformation of fruit, but the
disease is seldom important.
Spilocaea pyracanthae (formerly Fusicladium pyracantha). Pyracantha
Scab, widespread on leaves and fruit. The unsightly black scabs spoil the
appearance of bright berries. The fungus winters in the mycelial state in
attached leaves. Frequent spraying with bordeaux mixture controls scab but
causes some defoliation.
Spongospora
Plasmodiophoromycetes, Plasmodiophorales
Spores in a hollow sphere with several openings; zoosporangia formed; zoospores ante-
riorly bi-agellate; sexual fusion of myxamoebae.
Spongospora subterranea. Powdery Scab of potatoes, Canker, Spongy
Scab. Indigenous to South America and introduced into Europe more than
a century ago, potato scab was not noticed in North America before 1913, in
Maine. Ordinarily not important, it causes economic loss in some seasons.
Slightly raised pimples appear on tubers when they are less than an inch in
diameter; they are varying shades of brown on the surface, faintly purple
underneath. The epidermis, not growing as fast as the pimple, breaks and
curls back over the pustule, which, by this time, is a brown powdery mass
of spore balls and decomposed plant tissue. The lesions are often corked
off, but under favorable conditions large, depressed cankers may form. The
fungus winters on stored tubers or in soil, remaining viable for many years.
In the presence of a potato tuber and enough moisture, each spore in the ball
SCAB 575
germinates by swarmspores, which stay grouped together in a plasmodium,
dissolving cuticle and killing cells. When the food supply diminishes, the
plasmodium again breaks up into spore balls.
Control. Avoid low soggy ground; if such soil must be used, acidify it with
sulfur as for common scab.
Streptomyces
Rots.
Streptomyces acidiscabies. Acid Scab on beet, carrot, radish, parsley and
turnip.
Streptomyces scabies (Syn. Actinomyces scabies). Common Scab of
potatoes, Beet Scab, Corky Scab, Actinomycosis, general on potatoes,
widespread on beets, also reported on carrot, parsnip, radish, rutabaga, and
turnip. This disease may have been in America as long as potatoes have
been grown, but the causal organism was not described until 1890. Scab-
by potatoes, by lowering the market grade, mean an annual loss of several
million dollars. Chief symptoms are the tuber lesions, starting as minute
brown specks and progressing to scabs that are warty or with corky ridges,
or are pitted and depressed with the skin cracking open. Such potatoes can
be eaten, but have poor customer appeal and are wasteful because of the
deep peeling required. On beets, the scabs are similar but more bulging.
The pathogen can be found even in virgin soil. It invades young tubers and
may sometimes be seen as a grayish coating on freshly dug potatoes. It is
most destructive in soils with pH 5.7 and over, with its activity sharply lim-
ited in soils slightly more acid. Although its optimum temperature is 72 to
86F, the fungus can withstand great extremes of temperature and moisture
and can pass through the digestive tract of animals, returning to the eld in
manure.
Control. Seed tubers have been treated with formalin, but the organism is
so prevalent in potato soils that such treatment may have little result. Soils
already slightly acid may be further acidied with sulfur. Enough sulfur
to acidify highly alkaline soil would be too expensive and too injurious to
potatoes. Alkaline materials time, wood ashes, and manure should not
be applied to potato soil. Somewhat resistant varieties include Menonimee,
Ontario, Cayuga, and Seneca.
Venturia
Ascomycetes, Pleosporales, Venturiaceae
Perithecia setose, often only near apex, papillate; paraphyses absent; spores unequally
two-celled, olive.
Venturia cerasi. On cherry and European plum, perhaps a strain of the peach
scab fungus but not infecting peach.
Venturia inaequalis (Anamorph, Spilocaea pomi). Apple Scab, Scurf,
Black Spot, general on apple except in far South, widespread on crabapple;
reported also on mountain-ash and hawthorn, but probably other species of
Fusicladium infect these hosts. Scab is the worlds top-ranking apple disease
and is probably coextensive with the host. In this country it takes a fourth or
more of the crop in a favorable scab year, the average national loss running
around 8%, or over 10 million bushels. Scab is somewhat less important in
the South and in irrigated sections of Washington, but it is important in the
humid coastal areas (Fig. 3.58). The pathogen was rst described and named
by Fries in Sweden in 1819 and was recognized in New York and New Jersey
in 1834, apparently having come here with some European imports.
Symptoms. The rst symptom of scab on leaves is a dull smoky area that
changes to an olive-drab moldy spot, 1/4 inch or more in diameter, without
a sharp outline. Sometimes the leaf is raised or domed in the vicinity of the
Figure 3.58 Apple Scab on Leaf and Fruit
SCAB 577
Figure 3.59 Venturia inaequalis, the apple-scab fungus. A one-celled dark conidia of Fusicladium stage;
B perithecium with two-celled ascospores
spot; sometimes it turns brown and drops prematurely. Similar spots may be
formed on blossom pedicel, calyx, and petals, followed by dropping of young
fruit. Scabby lesions sometimes appear on twigs, but are less common.
On fruits, spots are small, more or less raised, rounded, dark olive areas (see
Fig. 3.58). As they increase in size, the cuticle ruptures to form a white rim
around a dark, velvety center, and still later the center may be raised, corky,
and tan in color, after dark mycelium and spores have disappeared. Lesions
are usually most abundant near calyx end of fruit; if they are too numerous,
the fruit splits.
Life History. The fungus winters in dead fallen leaves, producing small, dark,
ask-shaped perithecia and, toward spring, asci with eight brown ascospores,
unequally two- celled, with the upper cell wider than the lower (Fig. 3.59).
The ascospores mature about the time blossoms show pink, and are forcibly
expelled during warm spring rains. Each ascus elongates, protrudes its tip
through the mouth of the perithecium, and explodes its spore content. When
a spore, carried by wind, arrives on a young leaf or bud, it penetrates the cuti-
cle with a germ tube and develops a layer of branching mycelium just under
it. The scab spot is evident in about 10 days, when brown conidiopores bear-
ing olive brown, one-celled, somewhat pointed spores appear on the surface.
Secondary infection occurs when these conidia are carried to new infection
courts.
The expulsion of ascospores proceeds in a series of discharges over a rather
long period, up to 3 months, starting in February, on the West Coast, but
a shorter period, beginning in April, in New York. Germination and infection
take place from 41 to 79F. Length of wetting period necessary for primary
infection decreases as the temperature rises 13 to 18 hours of continuous
wetting at 43F and only 4 to 6 at 70F. Secondary infection from conidia
continues all season in rainy periods and even in storage scab may show up
on apples infected just before picking.
Control. No varieties are immune to scab. Resistance varies with the season
and the part of the country. McIntosh apples are very susceptible; Baldwins
are fairly resistant but may scab badly some years. There is more than one
strain of the fungus. Nitrogenous fertilizers increase yield of the fruit but also
susceptibility to scab.
Protective spraying, having a chemical lm on blossom, fruit, or foliage at all
times when weather makes infection probable, is the only real answer to scab.
This may mean more than a dozen applications in a wet year and a minimum
of ve any season, a program more suited to the commercial grower than to
the amateur. Timing is all-important, and most states have a spray warning
service that tells of imminent discharge of ascospores. Any spray schedule
must be tailored for the locality, the season, and apple varieties grown. The
apple grower gets this specic help from his county agents.
Venturia pyrina (Anamorph, Fusicladium pyrorum). Pear Scab, general
on pear, also on quince, similar to apple scab. The pear species of Venturia
overwinters in fallen leaves and also in affected twigs; the perithecia mature
somewhat later than those of apple scab. Conidia are formed on pear twigs
and washed to leaves and fruit.
Pear scab is not serious except on such varieties as Flemish Beauty, Winter
Nelis, Seckel, Anjou, Bosc, and Duchess. Bartlett pears are rather resistant.
SCURF
Two diseases, one of sweetpotatoes and one of potatoes, are commonly
called scurf.
Monilochaetes
Hyphae and conidiophores dark, spores hyaline, one-celled, oblong-cylindric, in chains.
Monilochaetes infuscans. Sweetpotato Scurf. Small, circular, brown or
black spots are formed on all underground parts, often forming a uniform
patch over the whole potato or a black patch on red-skinned varieties. The
skin cracks, and potatoes shrink in storage. The black conidiophores stick up
from the surface of the lesions like bristles. The fungus winters on the roots
and on decaying vines.
Control. Scurf, formerly present in 50% of New Jersey sweetpotatoes, is now
rare because of proper care. Set only healthy sprouts, grown from potatoes
bedded in sand that has not grown sweetpotatoes before.
Helminthosporium (Spondylocladium)
Conidiophores dark, straight, septate, the upper cells bearing whorls of conidia; conidia
dark with three or more cells.
Helminthosporium solani (formerly Spondylocladium atrovirens). Silver
Scurf of potatoes. Scab, Dry Rot, present in almost all potato districts but
not too important. Light brown lesions become somewhat blistered, giving
the skin a marked silvery appearance. The disease is only skin deep, and
control measures are seldom used.
Spondylocladium atrovirens (see Helminthosporium solani). Silver Scurf
of potatoes. Scab, Dry Rot, present in almost all potato districts but not too
important.
SLIME MOLDS
Slime molds belong to the Myxomycetes, a group intermediate between bac-
teria and fungi. Their assimilative phase is a plasmodium, which is trans-
formed into distinct fructications on a substratum. They are not parasitic
and are often found in rotting logs. Sometimes they are a nuisance in lawns,
for the plasmodium after ingesting decayed organic matter or microorgan-
isms for food moves up a grass blade for fruiting. Their spores are pro-
duced on or in aerial sporangia and are spread by wind. On absorbing water
the spore cracks open and the contents escape as a swarmspore, sometimes
two, with two agella. The swarmspore ingests food like an amoeba, divides
by ssion into a myxamoeba, unites with another to form a zygote, which
enlarges, with mitotic division, into a multinucleate plasmodium. There are
many species. Two only are listed here, as being common on turf.
Fuligo septica (formerly Mucilago spongiosa). Cream to yellow plasmodi-
um forms large grayish white structures, 2 to 6 cm long by 1 to 6 cm wide,
that are lobed and branched sporangia lled with a dark mass of purple, spiny
spores.
Mucilago spongiosa (see Fuligo septica).
Physarum polycephalum. Plasmodium colorless, watery-white or yellow.
Fruiting bodies small, gray, sessile, crowded on grass blades, and scattered
in groups or rings over an area of several feet. Spores are purple brown in
mass. The sporangia develop during humid weather in summer and autumn.
Use a stream of water to wash the spore masses off the grass.
SMUTS
Smuts, of the fungus order Ustilaginales, are named for their sooty black
spore masses. Like the rusts, they belong to the Basidiomycetes and are all
plant parasites, of most economic importance on cereals and grasses, but they
differ from rusts in having a less complicated life history and in being able
to live part of their lives saprophytically in rich organic matter or in culture
media. There are two spore forms. The teliospore, usually called a chlamy-
dospore, is formed by the rounding up of a hyphal cell. In addition to a thin
inner endospore wall, it has a thick outer exospore wall, usually dark, smooth
or ornamental. Teliospores are formed singly or united into balls. They can
be distributed long distances by wind, and spores of some species remain
viable for years. Some have to ripen several months before they can germi-
nate.
Occasionally the teliospore puts out a germ tube that penetrates host tissue
directly. More often it produces a promycelium that gives rise to sporidia,
which can bud to more sporidia. Classically true smuts have been divided
into two families on the type of sporidial formation: Ustilaginaceae, with
sporidia produced on the sides of a four-celled promycelium, and Tilleti-
aceae, with sporidia produced at the end of a one- or two-celled promyceli-
um. Fischer, however, points out that there are so many variations that it is
preferable to include all species in a single family, Ustilaginaceae, and to dif-
ferentiate the species on the basis of morphological characters and the host
family. This is logical, but we include here the families as they are given
in most textbooks and also the false smuts, Graphiolaceae (Ustilaginales),
which have an uncertain taxonomic position.
There are three types of infection with smuts, with control measures modied
according to type. The mycelium always penetrates the young host tissue
directly; it does not enter through stomata.
1. Infection of seedlings as the seed germinates, from spores adhering to
the outside of the seed or present in soil; controlled by dusting seed and
planting in noninfested soil.
SMUTS 583
2. Seedling infection by mycelium within the seed as a result of ovary infec-
tion from spores germinating on the stigma; controlled by treating seed
with hot water.
3. Infection of any actively growing meristematic tissue (roots, shoot, tas-
sels, or young ears) by spores transported by wind from decaying plant
material; controlled, partially, by spraying or dusting susceptible plants.
Burrillia
Tilletiaceae. Sori in various host parts, often in leaves, rather permanently embedded.
Spore balls with a central sterile mass surrounded by fertile teliospores but no sterile
cortex (surface layer). Teliospore hyaline to yellowish, rather rmly united. On water
plants.
Burrillia decipiens. Leaf Smut of oating heart (Nymphoides).
Cintractia
Ustilaginaceae. Sori usually in ovaries, black, more or less agglutinated spore mass-
es with a peridium. Teliospores single, olive to reddish brown, formed from a fer-
tile stroma surrounding a central columella of host tissues. On Cyperaceae and Jun-
caceae.
Doassansia
Tilletiaceae. Sori usually in leaves; spore balls large and conspicuous, with a sterile layer
around fertile cells. Teliospores pale yellowish brown to hyaline, thin walled. Germina-
tion often in situ. On water plants.
Doassansia epilobii. Leaf Smut on epilobium.
Entyloma
Tilletiaceae. Sori generally in leaves forming light spots, giving the name white smut, or
slightly raised darker blisters. Teliospores produced singly but often adhering in irreg-
ular groups hyaline to pale green, yellow, or brown. Sporidia formed on the surface
give the white powdery appearance.
Entyloma calendulae. Calendula White Smut. Spots are pale yellow,
turning brown to black, 1/4 inch in diameter. The smut is common but not
very serious in commercial calendula plantings around San Francisco. Plant
debris should be cleaned up, perhaps the location changed.
Entyloma compositarum, White Smut of composites, boltonia, calendu-
la, erigeron, eupatorium, gnaphalium, golden-glow, helenium, and prairie
coneower.
Entyloma dactylidis (E. crastophilum and E. irregulare). Bluegrass Blis-
ter Smut, on Poa spp., Oregon, Washington, Minnesota, North Dakota.
Gray-black, blister areas in leaves from subepidermal masses of chlamy-
dospores. A series of ne dotlike masses of sporidia (conidia), appear scat-
tered in rows along surface of the blisters.
Entyloma dahliae. Dahlia Leaf Smut, a European disease occasionally
reported here. It showed up in one location in California where overhead
watering was used, but disappeared when the practice was discontinued.
Leaves are marked by more or less circular spots, rst yellow-green, then
brownish and dry. Primary spores germinate in leaves and send projections
to the outside, where secondary spores are formed to spread the disease. Late
planting seems to increase disease incidence.
Entyloma ellisii. Spinach Smut. An occasional disease with infected
leaves pale and worthless. Spores are produced in irregular, marginal necrot-
ic lesions.
Entyloma lineatum. Smut of wild rice.
Entyloma polysporum. Leaf Smut of gaillardia, golden-glow, senecio, syl-
phium, andsunower.
Graphiola
Ustilaginales. This family and genus are sometimes included in the smuts, sometimes
not. The sori are erumpent, enclosed in a compact black peridium on leaves of palms.
The spores are formed in parallel chains, and bud laterally to form two or more sporidia,
which become somewhat colored with thickened walls.
Graphiola phoenicis. False Smut of palms, Leaf Spot on queen, canary
date, royal and Washington palms and on palmetto. Leaves are yellow-
spotted with small black scabs or warts having a dark, horny outer surface
and long, exuous sterile hyphae protruding from an inner membrane con-
taining powdery yellow or light brown spore masses. Seriously infected
SMUTS 585
leaves may die. The disease occurs on date palm where humidity is continu-
ously high, but is checked in desert areas best suited to date culture. Kustawy
variety is less susceptible than some others.
The disease also appears on small ornamental palms in greenhouses and con-
servatories. Cut out and burn infected leaves or parts.
Mycosyrinx
Ustilaginaceae. Spores united in pairs; sori with a double peridium in swollen pedicels
and peduncles. Mostly tropical.
Mycosyrinx osmundae. Inorescence Smut on osmunda fern.
Neovossia
Tilletiaceae. Sori in ovaries, semi-agglutinated to powdery. Teliospores borne singly,
each with a long pedicel appendage, and producing many sporidia.
Neovossia iowensis. On grains, affecting kernels in the dough stage.
Schizonella
Ustilaginaceae. Sori in leaves; short to long striae; black, agglulinated teliospores in
pairs, germinating with three- to four-celled promycellium with lateral sporidia. Two
species on Cyperaceae.
Sorosporium
Ustilaginaceae. Spores loosely united into balls, readily separable by pressure, in various
hosts, more often in reproductive parts. Germination by promycelium and sporidia or
germ tube. Mostly on grains.
Sorosporium saponariae. Flower Smut of silene.
Sporisorium
Ustilaginaceae. Sori in various host parts but mostly in inorescence; granular to pow-
dery, covered at rst by a peridium. Teliospores single, formed around a central columel-
la. Germination usually with sporidia. Most species on grains and grasses, sometimes
causing severe stunting.
Sphacelotheca cruenta (see Sporisorium cruentum). Loose Kernel Smut
on sorghum, causing smutting and excessive branching.
Sphacelotheca reiliana (see Sporisorium holci-sorghi). Head Smut of
corn, in Pacic states and scattered locations in South.
Sphacelotheca sorghi (see Sporisorium sorghi). Covered Kernel Smut.
Kernels replaced by smut galls
Sporisorium cruentum (formerly Sphacelotheca cruenta). Loose Kernel
Smut on sorghum, causing smutting and excessive branching. Controlled by
seed treatment and resistant varieties.
Sporisorium holci-sorghi (formerly Sphacelotheca reiliana). Head Smut
of corn, in Pacic states and scattered locations in South. Galls on tassels and
ears breaking into loose dark brown spore masses. Do not plant in a smutted
eld for 2 years; use certied seed, resistant hybrids.
Sporisorium sorghi (formerly Sphacelotheca sorghi). Covered Kernel
Smut. Kernels replaced by smut galls.
Thecaphora
Ustilaginaceae. Sori in various host parts, mostly inorescence; powdery or granular.
Spores rmly united into balls, with no sterile cells. Chiey on Leguminosae and Con-
volvulaceae.
Tilletia
Tilletiaceae. Sori mostly in ovaries, occasionally in vegetative parts of host forming
a powdery or semi-agglutinated spore mass, often foetid. On grains and grasses, called
bunt; interior of seed a solid mass of spore balls (see Fig. 3.60).
Tilletia buchloana. Bunt of buffalograss.
Tilletia caries. Dwarf Bunt of wheat. Plants a fourth or half size of healthy
plants.
Tilletia foetida. Stinking Smut, Common Bunt of Wheat, on wheat and
wheat grasses wherever grown, occasionally on rye. A major agricultural
disease, especially in Pacic Northwest, this is of historical importance as
the rst disease controlled by seed disinfection. In 1670 a ship was wrecked
SMUTS 587
Figure 3.60 Smut Spores. Ustilago (left), spiny chlamydospore germinating with promycelium and sporidia
formed at sides; Tilletia (right), reticulate chlamydospore with long H-shaped sporidia formed at end of promyceli-
um and sometimes forming small secondary sporidia
off the Coast of England, but the cargo of wheat was salvaged, free from
bunt because of its salt-water bath. Dark smut balls replace kernels, and
there is a shy odor. Plants are stunted, but not as much as with dwarf bunt.
Spore balls are broken in threshing and seed contaminated. Many materials
are offered for seed treatment. Seed dealers treat seed for farmers in special
machinery at low cost.
Tilletia pallida. Bunt on velvet and creeping bent grass, Oregon, Rhode
Island. Seeds are lled with black spores, plants stunted. The disease is seri-
ous where grass is grown for seed, with up to 80% nonviable seed.
Urocystis
Tilletiaceae. Sori mostly in leaves and stems, blackish; embedded in host tissues. Spore
balls permanent, without sterile cortex but sometimes with a layer of hyaline, hyphal
fragments. On Liliaceae, Primulaceae.
Tuburcinia trienthalus (see Urocystis trientalus). Leaf and Stem Smut
of starower.
Urocystis trientalus (formerly Tuburcinia trientalus). Leaf and Stem
Smut of starower.
Urocystis
Tilletiaceae. Sori usually in leaves, stem sheaths, occasionally in owers;
dark brown to black, powdery to granular. Spore balls with distinct sterile
spores on the surface, only a few fertile spores. Sori without peridium.
Urocystis agropyri. Flag Smut of wheat, also on wheat grass, red top, and
bluegrasses. Symptoms are similar to those of stripe smut.
Urocystis anemones (including U. hepaticae-trilobae). Leaf and Stem
Smut of anemone, hepatica, and trautveteria.
Urocystis carcinodes. Smut of aconite, baneberry, clematis, and cimicifu-
ga.
Urocystis colchici (Fischer includes U. cepulae in this species). Leaf Smut
of autumn crocus, camassia, Solomons-seal and false Solomons-seal.
Urocystis gladiolicola. Gladiolus Smut. This disease had been intercepted
several times at quarantine and appeared once in California elds, in 1950,
but apparently is eradicated there. Growers should be on the lookout for
corms with low blister swellings, with ridges paralleling veins, bluish black,
breaking open to expose dense black spore balls. Seedlings exhibit blister-
ing, shredding, and necrosis of stem and leaf tissues; they die if the disease
is severe.
Urocystis kmetiana. Floral Smut of eld pansy (Viola bicoler).
Urocystis magica (U. colchici). Onion Smut, general on onion, also on
leek, shallot, garlic, and chives. This is the most destructive onion disease,
found in the Connecticut Valley as early as 1861 and thence spread to
all northern onion-growing sections, but more important where onions are
grown from seed rather than sets as in most home gardens.
Black elongated blisters or pustules of spores break out on scales or leaves
of young plants. Many plants die; others survive and have black or brown
smut pustules on the cured bulbs. Plants are stunted but not rotten, although
smut may be followed by secondary rot organisms.
The spores can live in soil for years, but infection is possible only in young
plants from the second day after seed germination until the seedling is in
rst leaf, a period of 10 to 15 days. The spore is able to penetrate the onion
through root and cotyledon but cannot enter a true leaf. After entrance it
spreads through the seedling until it reaches the leaves to form fruiting pus-
tules just below the epidermis. When this ruptures, spores are dropped, to
be disseminated by running water and tools, on feet of persons and animals,
and on roots of transplanted vegetables. Onion smut is conned to states with
cool summers, optimum soil temperature for infection being 72F.
Urocystis tritici. Flag Smut of wheat. Plants are dwarfed with twisted leaf
blades; sheaths are marked with grayish-black stripes; diseased tissues dry
up and are shredded. Infected plants rarely produce heads.
SMUTS 589
Ustilago
Ustilaginaceae. Sori in various host parts; spore masses powdery to agglutinated; usually
dark brown to black, in some species yellow to purple without a peridium. Spores single,
not united in balls (see Fig. 3.60).
Ustilago avenae. Nigra Loose Smut, general on barley.
Ustilago avenae (including U. perennans). Black Loose Smut on oats and
some grasses. Individual owers in panicle are largely replaced by a spore
mass. The young seedling is diseased from the seed, and the fungus grows
systemically in the plant.
Ustilago buchloes. Stripe Smut on grass.
Ustilago bullata. Head Smut on many grasses.
Ustilago esculenta. Smut on wild rice.
Ustilago heueri. Erythronium Smut. Large dusty pustules lead to crack-
ing and dying of leaves of dogtooth violet.
Ustilago hordei. Covered Smut of barley. Heads are converted into hard,
black, smutted masses, enclosed within thin membranes.
Ustilago kolleri. Covered Smut of oats. Spore balls remain intact with-
in glumes until threshing, when spores are distributed over surface of seed,
ready to infect young seedlings.
Ustilago maydis (U. segetum). Corn Smut, Boil Smut, general on corn
but most destructive to sweet corn. The average annual loss is 3 to 5% but
it can be 100% in any one eld. The fungus was described in Europe in
1754 and not reported here before 1822, but it may be native along with its
host. There are many physiological races, and smut resistance is likely to
be correlated with lack of vigor, so that it has been hard to breed desirable
resistant varieties.
Any plant parts aboveground may be attacked stalks, prop roots, leaves, tas-
sels, husks, and ears (see Fig. 3.61). Large boils are formed, at rst covered
with a greenish white membrane, said to be good eating when boiled or fried.
Later the membrane breaks and releases myriads of dark chlamydospores.
The plant is often distorted. Infections are local; each boil is formed where
a spore lands, and there is no systemic growth through the plant. The fun-
gus is not seed-borne, and germinating seedlings are not affected. Chlamy-
dospores winter in soil, corn debris, and manure. They produce sporidia,
which may bud to form secondary sporidia, and these are carried by wind
and other agencies to corn plants, which are 1 to 3 feet high. Mycelium from
Figure 3.61 Corn Smut
SMUTS 591
spores of two sexes is needed for active development. Spores formed in the
rst boils provide inoculum for secondary infection of ears.
Corn smut thrives in warm weather, optimum temperature for spore germi-
nation being 80 to 92F. Heaviest infection occurs when scant rainfall in
early stages of growth is followed by moderate rainfall as corn approaches
maturity. Vigorous plants are most susceptible, but may escape the most seri-
ous effects because of their rapid growth. Spores retain viability 5 to 7 years.
They remain viable in passage through an animal into manure, but are killed
by the acids in silage.
Control. Seed treatment is not effective. Some hybrid varieties are rather
resistant. Most reliance in home gardens should be placed on cleanliness,
cutting off and burning all smutted parts before the boils break open to
release spores.
Ustilago mulfordiana. Fescue Smut on fescue grasses.
Ustilago tritici. Nuda Loose Smut. Normal heads replaced by black pow-
dery masses.
Ustilago striiformis. Stripe Smut, general on grasses wheatgrasses, red-
top, bentgrasses, fescues, ryegrass, and bluegrass; does not occur on cereals.
Long dark narrow striations develop in leaves; as the sori mature, spores
are freed, and the blade splits into ribbons. Plants are systemically infect-
ed, make poor growth, and inorescences are stunted or absent. Perennial
mycelium may overwinter in the plant.
Ustilago violacea. Anther Smut of carnation, dianthus, lychnis, and silene.
Infected plants grow slowly, produce many weak axillary shoots; stem
internodes are shortened; ower buds are short and squatty; calyxes tend to
split; owers are sprinkled with black sooty dust from the anthers, whose
pollen grains are replaced by smut spores. The fungus enters through owers
or injured surfaces and grows systemically. Spores are spread on cuttings.
Control by roguing diseased plants before owering. Do not take cuttings
from plants with grassy or bushy habit.
SNOWMOLD
Northern lawns and turf of golf greens often show round light patches as the
snow melts in early spring. Such a disease is called snow mold and may be
due to one of several fungi, sometimes to two appearing together.
Microdochium (Fusarium)
Rots.
Fusarium nivale, Teleomorph, Monographella nivales). (see Microdochi-
um nivale). Pink Snowmold, Fusarium Patch, most important on bent-
grass on golf courses but infecting other turf grasses and winter wheat and
winter rye.
Microdochium nivale (formerly Fusarium nivale, Teleomorph, Mono-
graphella nivales). Pink Snowmold, Fusarium Patch, most important on
bentgrass on golf courses but infecting other turf grasses and winter wheat
and winter rye. Irregularly circular patches, from 1 to 2 inches to a foot or
more, appear as snow is melting. They are whitish gray, often with a pinkish
tinge, and several patches may run together to cover large areas. Individual
plants have a bleached appearance, feel slimy when wet. Spores are formed
in salmon-pink sporodochia over stomata in leaves. They are sickle-shaped,
one- to three-septate. Perithecia are produced on the luxuriant white mycelial
mat.
Abundant moisture in the fall, snow falling on unfrozen ground, deep snow,
and a prolonged, cold wet spring are predisposing factors, but the presence
of snow is not a requisite for the disease. Severity is increased by applying
fertilizer in late autumn and an excess of organic matter in the soil. Reports
differ as to susceptibility, but Colonial, Washington and Metropolitan bent-
grasses appear to be more resistant then Seaside bent.
SNOWMOLD 593
Sclerotium
Blights.
Sclerotium rhizodes. Frost Scorch, String of Pearls, in northern states.
Not exactly a snowmold but appearing in early spring with bleached, with-
ered leaves covered with rows of tiny sclerotia. Collect clippings when mow-
ing diseased areas to remove sclerotia on leaf tips.
Typhula
Fruit body erect, simple, like a little club, on a long stipe from a sclerotium; basidia with
four sterigmata and simple, hyaline spores.
Typhula incarnata. Snowmold of turf and lawn grasses, Typhula Blight,
common in eastern United States. As the snow disappears in spring, a felty
white mycelial mat is seen over grass and adjacent soil. Plants wither and
turn light brown or tan in roughly circular patches, very conspicuous against
the green of the rest of the lawn. The chief diagnostic character is the pres-
ence of very small, tawny to hazel brown spherical sclerotia in large numbers
over affected parts. These can be made to fruit in the laboratory into rose-
colored sporophores up to 1 inch tall.
Control. The disease gradually disappears as moisture decreases and temper-
ature and sunlight increase; so control seldom seems necessary. Six weeks
after striking cases of snowmold, lawns are often uniformly green and show
little sign of having been affected. Phosphate fertilizers are said to decrease
injury from Typhula.
Typhula idahoensis. Snowmold on wheat and grasses in Idaho and Mon-
tana. Sclerotia are chestnut brown, sporophores fawn to wood brown, less
than 1/2 inch high.
SOOTY MOLD
Sooty mold is a black coating on surface of leaves or fruit composed of a weft
of dark mycelial threads. As here used, the term applies to saprophytic fungi
that live on insect honeydew and harm plants only indirectly. See

Black
Mildew for the true parasites with dark mycelium and spores giving a sooty
appearance to foliage.
Capnodium
Ascomycetes, Capnodiales
Mycelium supercial, dark; spores muriform, in perithecium-like conceptacles at tips of
branches of a carbonaceous stroma; associated with insect secretion on living plants.
Capnodium citri. Sooty Mold on citrus, on honeydew secreted by scale
insects, aphids, whiteies, especially abundant following whiteies in Flori-
da, black scale in California. A black velvety membranous coating is formed
over leaves, twigs, and fruit. If honeydew is slight, the coating appears in
spots; but if the insect secretion is abundant, the entire surface may be cov-
ered by a dense continuous membrane resembling black tissue paper. With
age, under dry conditions it may be blown off in fragments. The black mem-
brane is made up of hyphae that are individually olive green to deep brown,
with wide short cells. Branches may crisscross or be cemented together.
There are several spore forms: simple conidia that are cut off from upright
hyphae, others formed in small, black pycnidia, stylospores in very long
ask-shaped conceptacles, and muriform brown ascospores in perithecia.
Although sooty molds do not obtain food from the plant, the black membrane
interferes greatly with photosynthesis and food manufacture. Affected fruit
is smaller, with coloring retarded; it is more likely to decay than normal fruit.
Control is directed against the insects, either by spraying with insecticides or
by using entomogenous fungi and insect parasites. Oil sprays kill the insects
and help to clean the trees of the sooty covering.
SOOTY MOLD 595
Capnodium elongatum. Sooty Mold of tulip-tree, oleander, holly-osman-
thus, and others. Foliage of tulip-trees very frequently has a black coating,
often on honeydew secreted by the tulip-tree aphid, sometimes following
attacks of tulip-tree scale. A dormant oil spray controls the latter.
Capnodium spp. Sooty Mold on gardenia, g, crape-myrtle, azaleas, and
many other plants. Gardenias are especially subject to sooty mold following
whiteies, crape-myrtle after aphids, azaleas after mealybugs and magnolias
after scales. A summer oil spray helps to control the insects and loosens the
black coating so that it is more readily washed off.
Very often rhododendrons and other broad-leaved evergreens growing inside
the branch spread of tulip and other trees aficted with aphids and scales are
covered with sooty mold growing in the honeydew dropped down on foliage
from the tree overhead.
Fumago
Mycelium dark, creeping over surface of leaves; conidiophores dark, variable, bearing
conidia terminally or laterally; conidia variable, dark, muriform, frequently in chains;
saprophytic on honeydew; probably conidial stage of Capnodium.
Fumago vagans. A heavy black moldlike growth on leaves of linden and
many other ornamental trees and shrubs, also on house plants in honeydew
from aphids, mealybugs and scale insects.
Scorias
Ascomycetes, Capnodiales
Mycelium with parallel walls, forming a thick spongy mass; perithecium long-stalked,
round; spores four-celled.
Scorias spongiosa. Sooty Mold. Often on trees alder, beech, pine, etc.
SPOT ANTHRACNOSE
Diseases caused by species of Elsino anamorph state Sphaceloma, are
characterized by some overgrowth of tissue. When this hyperplasia is pro-
nounced, the disease is usually called scab; when the overgrowth is scarcely
noticeable (merely a slightly raised border around a necrotic center), the
disease has been commonly known as anthracnose. Recently, the term spot
anthracnose was introduced to differentiate a Sphaceloma malady from
anthracnoses caused by fungi with slime spores (Colletotrichum, Glom-
erella) and from the Venturia type of scab diseases. Consequently, all spot
anthracnose diseases are included in this section, but with the common
designation, scab or anthracnose, also listed.
Elsino
Ascomycetes, Myriangiales, Elsinoaceae
Asci are borne singly, at different levels, in an effused stroma, having a gelatinous inte-
rior and crustose rind, under or within the epidermis, which ruptures to expose the asci.
The anamorph state is a Sphaceloma.
Elsino ampelina. Grape Anthracnose, Birds Eye Rot, widespread on
grape. Small sunken spots with dark margins and light centers occur on fruit,
young shoots, tendrils, petioles, and leaf veins. Leaves may be distorted and
ragged from diseased portions dropping out. Outbreaks are sporadic in east-
ern states. Varieties Catawba, Campbell Early, Diamond, Norton, and Salem
are quite susceptible; Concord, Delaware Moore Early, and Niagara are resis-
tant. The fungus winters on canes.
Control. Apply a dormant lime-sulfur spray and four or ve sprays of bor-
deaux mixture: when new shoots are 7 to 8 inches long; just after bloom; 7
to 10 days later; and when berries are half grown.
Elsino cinnamomi. Camphor-Tree Scab. Inconspicuous brown leaf
spots, sometimes dropping out; elongated raised lesions on veins, petioles,
and stems. Reported from South Carolina.
SPOT ANTHRACNOSE 597
Elsino corni. Dogwood Spot Anthracnose, a serious threat to owering
dogwood from Delaware to Florida, also reported from Louisiana. Infected
buds do not open, or they produce stunted, malformed owers, marked
with numerous small, circular to elongated spots with light tan centers, pur-
ple to brown borders, up to 50 on a bract. Leaf spots are 1 to 2 mm, slightly
raised at the margin, purple paling to yellow-gray at centers, which may be
broken in a shot-hole effect. There may be 100 spots on a leaf, scattered or
concentrated at tip, margin, or midrib. Spots on petioles, fruit clusters, and
stems are similar to leaf spots.
Elsino diospyri. Spot Anthracnose on leaves of native persimmon,
reported from Florida, 1955.
Elsino euonymi-japonici. Spot Anthracnose on evergreen euonymus,
Florida. Small, roundish spots, mostly on upper surface of leaves, brown with
raised, orange-cinnamon margin; stem cankers circular to elliptical, wrinkled
or ssured, grayish white with raised orange margins.
Elsino fawcettii. Sour Orange Scab, Citrus Scab, Verrucosis on cit-
rus fruits, except rare on sweet orange. Lemons, sour orange, King orange,
bitter orange, and calamondin are very susceptible; Mandarin and Satsuma
oranges, tangerines, and all grapefruit except Royal and Triumph are mod-
erately susceptible. Climatic conditions play a part. Grapefruit and lemons
in the Rio Grande Valley are less susceptible than in Florida, but Satsumas
in Alabama are more susceptible than those in Florida. Known in the Orient
since ancient times, scab is believed to have come to Florida on Satsumas
from Japan. It was rst recorded there in 1885; the fungus was identied as
a Sphaceloma in 1925.
Tender growth is most readily infected, and the disease is most important
on young trees. On leaves, minute, semitranslucent spots change to raised
excrescences with corky crests, pale yellow to pinkish, then dull olive drab
with a conical depression opposite the crust. Foliage may be wrinkled or
stunted. Fruits have slightly raised scabs or are warty with corky crests,
which may grow together into large irregular patches. Scabs on grapefruit
may ake off as the fruit matures, with the area remaining green.
Spores are spread by wind, rain, dew-drip, possibly by insects. The young
fruit of grapefruit is very susceptible right after petal fall, but becomes pro-
gressively resistant and is practically immune when it reaches 3/4 inch in
diameter. Temperature range for severe infection is from 59 to 73F. Exces-
sive nitrogen increases scab. The pathogen winters on infected leaves, some-
times fruits.
Control. Apply a neutral copper spray or bordeaux mixture just before
growth starts in spring. A second copper spray, just after owers shed,
controls melanose as well as scab.
Elsino ilicis. Chinese Holly Spot Anthracnose. Numerous black spots,
1 to 2 mm, coalesce to large black patches on upperside of leaf mostly the
apical half, with distortion. Shoots and berries have brown to gray lesions
with slightly raised margins.
Elsino jasminae. Jasmine Scab. Reported from Florida. Spots numerous,
round or irregular, up to 2 mm.
Elsino ledi. Ledum Spot Anthracnose, widespread on ledum, Labrador
tea, and salal in Northwest, leucotho in Florida. Leaf spots are grayish white
with red-brown borders and purple margins. The disease is not serious.
Elsino lepagei. Scab on sapodilla and canistel in Florida (found on young
nursery stock in cans). Small, raised spots, gray at center.
Elsino leucospila. Camellia Scab, also recorded on ternstroemia in Flori-
da. Some corky excrescences on camellia foliage are due to this pathogen,
others to moisture relations.
Elsino magnoliae. Magnolia Scab on Magnolia grandiora from Geor-
gia to Louisiana. Spots are circular to angular, with black papillae in centers,
on upper leaf surface along midrib, margin, or tip. Infected leaves may drop.
Elsino mangiferae. Mango Scab. Spots usually originate on underside of
young mango leaves but become visible above. They are circular to angular,
dark brown to black with olive buff centers. Spots on mature leaves are larger,
slightly raised with narrow brown margins and dirty white centers. Stems
have irregular grayish blotches; fruit, gray to brown spots with dark margins.
Elsino mattirolianum. Spot Anthracnose on madrona and strawberry
tree (Arbutus spp.) in California.
Elsino parthenocissi. Virginia Creeper Soft Anthracnose. Leaf spots
are few to numerous, circular, scattered or along midribs and veins; they
have buff centers with narrow brown margins; fruit spots are grayish white;
lesions on petioles are somewhat raised. Also reported from pepper-vine.
Elsino phaseoli. Lima Bean Scab. First United States report from North
Carolina probably from imported seed. Lesions on pods, stems, and leaves.
Elsino piri. Pome Fruit Spot Anthracnose on pear, apple, and quince in
moist sections of western Washington and Oregon, more prevalent in home
gardens than commercial orchards. Fruit spots are small, up to 2 mm, red or
reddish purple with pale centers, upwards of 100 on an apple.
Elsino quercicola. Spot Anthracnose on water oak, Florida.
Elsino quercus-falcatae. Spot Anthracnose on southern red oak, Geor-
gia, South Carolina. Blackish brown leaf spots are few to abundant, scattered
over upper surface.
Elsino randii. Pecan Anthracnose, Nursery Blight on pecans in the
Southeast, an important nursery disease, limiting factor in production of
budded pecans in wet seasons. Small reddish lesions develop on both leaf
surfaces, those on the upper surface later turning ash gray. Diseased tissues
become brittle and fall out, leaving ragged margins and perforations. Spray
trees with bordeaux mixture when rst leaves are half grown; follow with
three sprays of bordeaux at 3- to 4-week intervals.
Elsino rosarum. Rose Anthracnose, widespread on rose, collected on
wild roses as early as 1898, in most areas more important on climbing roses
than on hybrid teas. Leaf spots are scattered or grouped, sometimes running
together, usually circular, up to 1/4 inch. Young spots are red, varying brown
or dark purple on upper leaf surface, showing up to 2 to 6 days after inoc-
ulation but not visible on under surface for 2 to 4 weeks, then dull reddish
brown to pale purple. On aging, the center of the spot turns ashen white, with
a dark red margin. Leaves may turn yellow or reddish in area of spots, may
have slits or perforations as the centers fall out.
Cane spots are circular to elongated, raised, brown or purple, with depressed
light centers and acervuli in barely visible dark masses. The fungus winters
in cane spots; spores are produced and spread only in rainy periods. A single
leaf lesion may produce 10,000 spores within an hour after wetting and will
continue production as long as the rain lasts.
Control. Where possible, prune out infected canes in spring. Keep foliage
protected as for blackspot. Sulfur and copper compounds are effective.
Elsino solidaginis. Goldenrod Scab in Florida, South Carolina, and
Georgia. New growth is affected as it develops. Lesions formed on midrib,
veins, petioles, and leaf blades are raised on one surface, sunken on the other,
with white to gray centers and brown borders.
Elsino tiliae. Linden Spot Anthracnose reported from Nova Scotia and
Virginia. Gray spots with black margins are numerous on leave blades and
petioles.
Elsino veneta. Bramble Anthracnose, general on blackberry, dewberry,
raspberry, being most common on black raspberry. Circular, reddish brown
sunken spots with purple margins and light gray centers, up to 3/8 inch in
diameter, appear on young shoots. On older canes these grow together into
large cankers. Similar spots, not always with purple margins, are formed on
fruit, leaf, and ower stalks. Leaf spots are rst yellowish, then with a red
margin around a light center, which may drop out. Leaves may drop pre-
maturely; fruit may dry up as a result of loss of water from infected canes.
Primary spring infection comes from ascospores produced in old lesions on
canes; secondary spread is by conidia.
Control. Cut old canes or handles from black raspberries after setting;
remove and burn old fruiting canes after harvest. In some cases the single
late dormant spray has controlled anthracnose without later sprays; in oth-
ers three foliage sprays have been effective without a dormant spray. Black
raspberry Quillen is quite resistant.
Sphaceloma
Acervuli disc- or cushion-shaped, waxy; conidiophores simple, closely grouped or com-
pacted, arising from a stromalike base; spores one-celled, hyaline, ovoid or oblong.
Teleomorph state where known is Elsino.
Sphaceloma araliae. Aralia Scabon Hercules club (Aralia spinosa), Mary-
land and Missouri.
Sphaceloma cercocarpi. Spot Anthracnose of birch-leaf mahogany, in
California. Leaf spots are nearly circular, up to 3 mm across, with pale cen-
ters and slightly elevated purple margins.
Sphaceloma hederae. English Ivy Scab. Leaf spots are raised with red-
brown margins, pale depressed centers, often numerous.
Sphaceloma lippiae. Lippia Spot Anthracnose on fog-fruit. Closely
resembling mint anthracnose and found in same elds in Indiana, also
reported from Florida. Numerous spots on leaves and stems are scattered
or grouped and nearly conuent; centers are depressed, buff-colored, with
purple margins.
Sphaceloma menthae. Mint Anthracnose, Leopard Spot Disease. Cir-
cular, oval or irregular spots on leaves, stems, and rootstocks are black with
white centers, up to 5 mm. Formerly serious, this disease is now controlled
in commercial mint elds by fall plowing, covering plants deeply.
Sphaceloma morindae. Morinda Scab. Buff-colored spots on leaves,
stems, and petioles, Florida.
Sphaceloma murrayi. Gray Scab of willow. Leaf spots are round, irregu-
lar, somewhat raised, grayish white with narrow, dark brown margins, often
conuent, sometimes fragmenting; long narrow patches along veins.
Sphaceloma oleandri. Oleander Scab. Leaf spots spherical to irregular,
densely grouped over entire surface, whitish with brownish black margin,
slightly elevated, 1 to 4 mm.
Sphaceloma perseae. Avocado Scab, one of the most important avocado
diseases in Florida, some years with nearly 100% infection; also occurring
in Texas. Leaf lesions are mostly on upper surface, very small red spots with
a dark olive conidial growth. Fruit lesions are corky, raised, brownish, oval,
but often coalescing giving a russeted appearance; sometimes cracking to
allow entrance of fruit-rotting fungi. Avoid highly susceptible varieties like
Lulu. Spray with bordeaux mixture as for blotch.
Sphaceloma poinsettiae. Poinsettia Scab. Light raised lesions on stems,
veins, and midribs, pale buff at center with purple to nearly black margins.
Sphaceloma psidii. Guava Scab, reported in feijoa in Florida.
Sphaceloma punicae. Pomegranate Spot Anthracnose. Very small pur-
ple spots with paler centers on both leaf surfaces.
Sphaceloma ribis. Gooseberry Scab, Washington. Leaf spots numerous,
very small, raised, and grayish.
Sphaceloma spondiadis. Mombin Scab. On purple mombin (Spondias)
Florida.
Sphaceloma symphoricarpi. Snowberry Anthracnose, widespread on
snowberry, impairing beauty of ornamental plants, rst described from New
York in 1910; also on coralberry. Leaf spots appear in early spring, minute,
dark purple to black, aging with dirty gray centers, coalescing into large
areas subject to cracking. Leaves may be misshapen from early marginal
infections. Spotting is inconspicuous in owers but pronounced on berries,
with purple areas becoming sunken and pinkish. Secondary infection by an
Alternaria shrivels fruit into brown mummies. A dormant lime-sulfur spray
followed by foliage sprays may help.
Sphaceloma viburni. Snowball Spot Anthracnose.
Sphaceloma violae. Violet Scab, Pansy Scab, widespread on violet and
pansy from Connecticut to Louisiana and Texas, a limiting factor in main-
taining violet collections. Reddish spots with white centers change to irregu-
lar to elongated raised scabs on leaves and stems, often with much distortion.
Remove and burn old leaves.
Sphaceloma spp. Undetermined species have been reported on Bignonia,
catalpa, camellia, and sambucus, in Louisiana, on buttonwood in Florida,
rhododendron in Washington.
VIRUS, VIROID, PHYTOPLASMA
PATHOGENS AND DISEASES
For many years the classication of plant viruses was in a state of chaos.
Fortunately recent biochemical and molecular investigations on organization
and structure of genome as well as on structural and nonstructural viral pro-
teins provided enough data to create a denition of species. A virus species
is a polythetical class of viruses consisting of replicating lineage and occu-
pying a particular ecological niche. This indicated that viruses and biological
entities that possess genes, replicate, interact with hosts and are exposed to
selection pressure, thus specialize and evolve. Guidelines provided in the
Sixth and Seventh Reports of the International Committee or the Taxono-
my of Viruses are partly followed, especially by use of a virus species name
or vernacular name for the not fully described viruses. Following are virus
species (including viroids and phytoplasmas) and virus diseases in alphabet-
ical order by common names.
Abelia Latent Tymovirus
Symptomless on Abelia; occurs in Eastern USA.
Abutilon Infectious Variegation; Abutilon Mosaic Bigeminivirus
A single variegated seedling found among green plants imported into Eng-
land from the West Indies in 1868 was propagated vegetatively as an orna-
mental variety. The bright yellow mottling on green leaves tends to disappear
in subdued light. Transmission is by grafting, occasionally by seed, and, in
native Brazil, by whitey Bemisia tabaci. Plants may recover if variegated
leaves are persistently removed but may be reinfected.
VIRUS, VIROID, PHYTOPLASMA PATHOGENS AND DISEASES 603
Albutilon Yellows Closterovirus
Transmitted by Triaulerodes abutilonea. Infected plants are chlorotic.
Alligatorweed Stunting Closterovirus
Occurs in North America region and Florida.
Alfalfa Mosaic Alfamovirus
Potato, Celery Calico; Bean Yellow Dot. Various strains of the alfalfa virus
are transmitted by cotton, pea, and other aphids to bean, clovers, pea,
cucumber, potato, tomato, zinnia, tumbleweed, poison hemlock, wild carrot,
Japanese pachysandra, and other hosts. Calico is a minor potato disease in
California and Idaho. Leaf spots are irregular, brilliant yellow to gray; yield
may be reduced. Celery has a conspicuous yellow-green mosaic; bean has
small, necrotic lesions.
Almond Bud Failure
Virus on Drake almond, in California, is transmissible by grafting. Limbs
have many branches, some dead at the end; leaves are darker green, more
upright, retained longer than normal; fruits few, often misshapen.
Almond Calico
On almonds in California, graft transmissible. Chlorotic blotches in leaves.
Alstroemeria Mosaic Potyvirus
and Alstromeria Streak Potyvirus
Both viruses are transmitted by aphids in a non-persistent manner.
Alternanthera Mosaic Potexvirus
Found in Florida, Maryland, and Pennsylvania on skullcap, recracker plant
and moss rose.
Apple Chlorotic Leaf Spot Trichovirus
Originally considered as a closterovirus but now is the type species of Tri-
chovirus genus.
Apple Green Mottle
On Duchess variety in New York. Fruit with discolored rings, of little value.
Apple Mosaic Ilarvirus
Occurring naturally only on apples; no insect vector known; transmitted by
budding. Small irregular cream to yellow leaf spots coalesce to large chlorot-
ic areas, with or without vein-banding. Three strains cause severe mosaic,
veinbanding mosaic and mild mosaic. There is no marked reduction in yield.
Hosts other than members of Rosaceae family include lbert, hop, birch, and
horse chestnut.
Apple Stem Grooving Capillovirus
First reported in Viriginia Crab. The main symptoms are stem grooves and
abnormal graft union.
Tulare Apple Mosaic Ilarvirus
Reported from California; has a wider host range than apple mosaic.
Apple, Dapple
Fruit with circular islands or patches remaining green; on trees with Virginia
crab or Robusta V bodystock; rst noted in New Hampshire.
Apple Stem-Pitting
Wood-pitting in Virginia Crab bodystock, sometimes followed by bark
cracks, dwarng, early fruit production.
Apricot Gummosis
First noted in Washington in 1947; transmitted by budding. Profuse gum-
ming on branches and trunks, necrosis and dieback of new shoots; dead,
punky areas in fruit.
Apricot Ring Pox
In California and Colorado. Irregular ring spots with marked vein clearing in
some varieties, chlorotic mottling in others; dead tissue may fall out leaving
shot holes. Fruit bumpy or with reddish brown necrotic spots extending into
esh.
Arabis Mosaic Nepovirus
It is one of the viruses that causes lilac yellow ring symptoms. Occurred on
many genera of ornamental, vegetable and orchard plants. Transmitted by
nematodes. This virus causes foliar blanching on hosta.
Artichoke (Globe) Curly Dwarf Potexvirus
In California on artichoke, cardoon, and zinnia, milk thistle. Leaves curled,
plants dwarfed, killed.
Artichoke Latent Potyvirus
Symptoms on Cynara spp. are none or stunting and yellow ecking of plants.
Ash Ring Spot =Arabis Mosaic Nepovirus
On white ash, New York. Chlorotic green and reddish spots, rings, line pat-
terns; stunting; dieback.
Ash Witches Broom
Reported from Louisiana on Arizona ash. Yellowish leaves, a fourth to
a third normal size; multiple, spindly shoots.
Asparagus 1 Potyvirus
Symptomless on Asparagus ofcinalis.
Asparagus 2 Ilarvirus
Caused stunting and decline on Asparagus ofcinalis plants. Virus is trans-
mitted by pollen to the seed and to the pollinated plant.
Aster Chlorotic Stunt Carlavirus
Reported in Aster novae-angliae in Ontario, Canada.
Aster Ring Spot
In Florida on China aster, pepper, pansy, and other plants. Yellow ring, line,
and oakleaf patterns. A strain of Tobacco Rattle Tobravirus.
Avocado Sun Blotch Viroid
There are long, narrow, shallow, longitudinal grooves, buff-colored on stems,
whitish on green fruit, reddish purple on purple fruit. Shoots may be twisted
and abnormal. Transmitted through seeds.
Avocado 3 Alphacryptovirus
Transmitted only by seeds.
Barley Yellow Dwarf Luteovirus
Occurs on tall fescue, and various Poa and Festuca spp. Occurs on cereal
crops in Alaska.
Barley Yellow Streak Mosaic
Occurs on barley in Alaska.
Bayberry Yellows
Wavy margins and tips on young apical leaves, distorted margins on older
leaves, which are pale, yellow, small. Plant is stunted, with shortened intern-
odes, few or no fruits.
Bean Common Mosaic Potyvirus
Distributed worldwide in common beans wherever they are grown. Many
strains of the virus were distinguished. They are transmitted by vectors, sap,
pollen and seed. Virus is serologically related to 17 other Potyviruses.
Bean Mosaic =Bean Common Mosaic Potyvirus
Found wherever beans are grown, transmitted by many species of aphids
pean, cotton, cowpea, cabbage, peach, spirea, turnip and in seed. First
leaves are crinkled, stiff, chlorotic; older leaves have chlorotic mottling,
often with leaf margins rolled down. Mosaic-resistant varieties include
Robust, Great Northern, U.S. No. 5 Refugee, Idaho Refugee, and Wisconsin
Refugee.
A strain known as bean greasy pod virus causes a greasy appearance of the
pods in some western states. The asparagus-bean mosaic is a light and dark
green mosaic with leaf rolling transmitted by seed and by the pea aphid. The
virus may be a strain of bean mosaic virus or a different virus.
Bean Leaf Roll Luteovirus
Legume virus transmitted by aphids.
Bean Pod Mottle Comovirus
Systemic mottling in some varieties; circular, light brown local lesions on
pods of other varieties. May also be seed transmitted in soybean.
Bean Southern Mosaic Sobemovirus
Chlorotic mottling or localized necrosis of foliage; pods with dark green
blotches or shiny areas, slightly malformed, short, curled at end. Virus
present in new seed but not in that stored 7 months.
Bean Yellow Mosaic Potyvirus
Mild Mosaic of Gladiolus. On beans, peas, sweet peas, clover, Tahitian bridal
veil, gladiolus, false lupine, and freesia. In beans there is a coarse yellow
mottling and distortion of leaves, which are pointed downward; proliferation
of stems; shortening of nodes and general stunting; reduced pod production;
delayed maturity. In pea and sweet pea there is veinal chlorosis, with slight
rufing. Gladiolus owers are striped or ecked, young leaves have an angu-
lar green mottling, but symptoms are mild compared with the disease on
beans and freesia, which should not be planted near gladiolus and clovers.
The virus is transmitted by bean and pea aphids but not through seed. Rogue
infected plants as soon as noticed.
Bean Yellow Stipple =Cowpea Chlorotic Mottle Bromovirus
Mild mottle and chlorotic spots on bean leaves, sometimes coalescing.
Beet 2 Alphacryptovirus
Transmitted only by pollen and seeds.
Beet Black Scorch
Necrovirus, Tombusriridae. Reported in Colorado.
Beet Curly Top Hybrigeminivirus
Conned to North America, curly top is especially important in the com-
mercial sugar beet industry west of the Continental Divide, but it is com-
mon on many plants. Vegetables include bean, beet, carrot, celery, cabbage
and other crucifers, cucumber, melon, squash, pumpkin, eggplant, spinach,
chard, New Zealand spinach, horseradish, and tomato. Ornamentals include
alyssum, blue ax, campanula, carnation, columbine, coreopsis, cosmos, del-
phinium, foxglove, geranium, larkspur, nasturtium, pansy, petunia, poppy,
portulaca, pyrethrum, scabiosa, Shasta daisy, stock, strawower, veronica,
and zinnia.
In beets there are clearing of veins, leaf curling, with sharp protuberances
from veins on underside of leaf, increase in number of rootlets. In tomato,
where the disease is called western yellow blight or tomato yellow, seedlings
turn yellow and die. Older plants show twisting and upward rolling of
leaets, stiff and leathery foliage, with leaf petioles curling downward;
branches and stems are abnormally erect; the whole plant is dull yellow,
often with purple veins; roots are killed, few fruits formed.
In cucurbits, tips of runners bend upward; old leaves are yellow, tip leaves
and stems abnormally deep green. In beans, there is a thickening and down-
ward curling of rst true leaf, which becomes brittle. The plant stops growing
and may die. Older plants survive until the end of the season, with puckering
and downward curling at the top of the plant, reduction in size of new leaves,
shortened internodes.
Ornamentals grown near diseased beets are usually infected. Zinnias have
shortened internodes, chlorotic secondary shoots arising from leaf axils. Cos-
mos leaets are twisted and curled, petioles bent down. Geranium leaves are
chlorotic between veins with protuberances on lower surface.
The virus is conned to phloem in plants and is transmitted by the beet
leafhopper (Circulifer tenellus). The insects winter on weed hosts, laying
eggs and maturing the rst generation there before migrating in swarms,
often hundreds of miles, to sugar beet elds. When the beets are plowed
out, the hoppers migrate to neighboring gardens.
Control. Destruction of weed hosts helps somewhat, as does early planting.
There are resistant varieties of sugar beets, none of table beets. Tomatoes are
sometimes protected with temporary muslin tents. Infected plants must be
destroyed.
Beet Distortion Mosaic Virus
Transmitted probably by a fungus Polymyxa betae; spreads in California.
Beet Latent Virus
A symptomless virus in beets.
Beet Mosaic Potyvirus
On sugar and table beets, spinach. Discrete yellowish lesions, then chlorotic
mottling, darkening of vascular tissue; leaves bend back near the tips, which
sometimes die. Transmission is by pea, peach, bean, and other aphids.
Beet Ring Mottle
On sugar beet and spinach; stunting, distortion, mottling; transmission by
aphids.
Beet Necrotic Yellow Vein Tobamovirus
On sugar beet and transmitted by soil-borne fungus Polymyxa betae.
Beet Pseudo-Yellows Closterovirus
Yellowing of sugar beet, carrot, spinach, cucumber, lettuce, and ornamentals;
transmission by greenhouse whitey.
Beet Savoy
Leaves are dwarfed, curled down, with small veins thickened; roots have
phloem necrosis. A plant bug (Piesma cinerea) is the vector.
Beet Western Yellows Luteovirus
In Europe known as Beet Mild Yellowing Luteovirus. More than 150 species
from 23 families are susceptible. Virus-transmitted by insects but principal
natural vector is Myzus persicae.
Beet Yellow Net Luteovirus
On beets and chard. Leaves have a yellow network of veins against a green
background. Transmission by the peach aphid.
Beet Yellows Closterovirus
On beets and spinach. Outer and middle leaves are yellowed, thickened, brit-
tle, with chlorotic areas waxy. Vectors are peach and bean aphids.
Bidens Mottle
On Rudbeckia, Zinnia, and Ageratum.
Bidens Mottle Potyvirus
On Rudbeckia, Zinnia, and Ageratum.
Blackberry Dwarf
Loganberry Dwarf.
Blackberry Dwarng
On brambles in California.
Blackberry Mosaic
Mottling, crinkling, vein clearing and distortion.
Blackberry Variegation
On raspberry and blackberry. Infected leaves are nearly white at maturity.
Blackeye Cowpea Mosaic
On urd bean.
Blueberry Leaf Mottle Nepovirus
Reported on cultivars Rubel and Jersey in Michigan. Virus is transmitted by
pollen and by honeybees which carry the pollen.
Blueberry Necrotic Ring Spot
A strain of tobacco ring spot virus, causing stunting and distortion; transmit-
ted by dagger nematodes.
Blueberry Necrotic Shock Ilarvirus
Leaf and ower necrosis symptoms occurred for 14 years, then plants
recover and remain symptomless.
Blueberry Red Ring Spot Caulimovirus
Red spots and rings, oak leaf patterns.
Blueberry Ring Spot
A minor disease chiey on Cabot with red rings and dots in leaves.
Blueberry Scorch Carlavirus
Transmitted by grafting, it caused marginal chlorosis and necrosis of leaves
but some cultivars remain symptomless.
Blueberry Shoestring Sobemovirus
Symptoms of shoestring disease included reddish streak on stem, narrow
strap-like leaves and ower breaking.
Blueberry Shoestring Sobemovirus
On highbush bluberry.
Blueberry Stunt Phytoplasma
Bushes are dwarfed with small leaves, yellowing in summer, brilliant red
in fall; berries are small, poor. Transmission by a leafhopper (Scaphytopius
magdalensis). Variety Rancocas is quite resistant; Harding is tolerant. Elim-
inate wild blueberries near cultivated.
Broad Bean Severe Chlorosis Closterovirus
Caused chlorosis, necrosis and leaf malformation on broad beans.
Broad Bean Wilt Fabavirus
Virus is the type species of genus. It is known also as a catalpa chlorotic
leaf spot, tropaeolum, nasturtium and petunia ringspots, pea streak virus and
parsley virus 3.
Broad Bean Wilt Fabavirus
Causes leaf mottle, ring spots, and poor growth of brous-rooted begonia;
also found on clockvine, bean, lettuce, spinach, lambsquarter, ajuga, and
dogwood.
Brome Grass Mosaic
Recently reported on Kentucky bluegrass, in Kansas.
Brome Mosaic Bromovirus
On cowpea.
Cabbage Black Ring Spot
Turnip Mosaic Potyvirus.

Cabbage Ring Necrosis =Turnip Mosaic Potyvirus
On cabbage and other crucifers, also infecting petunia, zinnia, calendula,
cucumber, beet, chard, and spinach. Concentric necrotic rings on leaves and
irregular dark, slightly sunken lesions on stems.
Cactus X. Potexvirus
On night-blooming cactus (Hylocereus).
Calibrachoa Mottle Carmovirus
Chlorotic blotch, leaf mottling and interveinal yellowing on Calibrachoa
species.
Camellia Yellow Mottle Varicosavirus
Infectious Variegation; Color-Breaking. A graft-transmissible disease with
at least four strains; CV1 large white spots on petals; CV2 small white
ecks on petals, leaf variegation; CV3 feathery mottling of owers; CV4
slower variegation and leaf mottle.
Canna Mosaic
Leaves with irregular pale yellow stripes from midrib to margin, parallel with
veins, may be wrinkled and curled with chlorotic areas often dusty brown.
Stems, sepals, and petals have yellow bands. Transmitted by peach and other
aphids.
Canna Yellow Mottle Badnavirus
Bacilliform virions, not enveloped, 120130 nm length and 28 nm wide.
Virus caused systemic veinal yellowing, mottling and chlorosis of Canna
spp.
Carnation Etched Ring Caulimovirus
Causes an etched-ring pattern on carnation leaves.
Carnation Italian Ringspot Tombusvirus
It is synonym for carnation strain of Tomato Bushy Stunt Tombusvirus.
Carnation Latent Carlavirus
This virus is serolocicaly related to potato viruses S and M, chrysanthemum
virus B and other carlaviruses.
Carnation Mosaic
Widespread on carnation and sweet william. Light and dark green mottling
in young leaves is followed by yellow or necrotic spots or streaks. Flowers
may be spotted or striped. The vector is the peach, not the carnation, aphid.
Greenhouse fumigation helps in control.
Carnation Mottle Carmovirus
Common in commercial carnations but producing only faint leaf mottling
and ower streaks; transmitted by root contact or cutting knife.
Carnation Necrotic Fleck Closterovirus
Severe necrotic ecking and streaking to mild yellow ecks on carnation.
Carnation Ring Spot Dianthovirus
Concentric rings on sweet william with vein clearing, then general mosaic;
necrotic rings on carnation, often combined with reddening and curling of
older leaves. Sap-transmissible on cutting knife.
Carnation Streak
A strain of Carnation Necrotic Fleck Closterovirus. Yellow or reddish
spots and streaks parallel to veins; lower leaves turn yellow and die. Graft-
transmissible.
Carnation Vein Mottle Potyvirus
The best test for diagnosis of this virus is immunosorbent electron microsco-
py technique.
Carnation Yellows
Foliage mottling, ower streaking due to combination of streak and mosaic
viruses.
Carrot Motley Dwarf =Carrot Mottle Umbrarvirus
An Australian disease now present in California. Leaets are small, chlorot-
ic, with twisted petioles; plants are stunted, roots unmarketable. Vectors are
aphids.
Carrot Red Leaf Luteovirus
Transmitted by aphids in persistent manner, but does not multiply in the vec-
tor. No transmission by sap, seeds, or pollen.
Carrot Thin Leaf Potyvirus
Caused vein clearing and malformation of leaves (narrowing).
Cassava Common Mosaic Potexvirus
Reported from Florida, virus is transmitted by mechanical inoculation and
by grafting.
Cauliower Mosaic Caulimovirus
Widespread on crucifers, broccoli, brussels sprouts, cauliower, chinese cab-
bage, collard, kale, annual stock, and honesty. Clearing of veins in cauliow-
er is followed by mild chlorotic mottling, with veins usually banded by dark
green necrotic ecks. Midribs are curled, leaves distorted, plants stunted,
terminal heads dwarfed. Stock is rosetted, with shortened internodes. Trans-
mission is by cabbage, false cabbage, peach, and other aphids.
Celery Calico
On cucumber, crookneck squash, tomato, celery, and larkspur. In celery
there are vein clearing, puckering, and downward cupping of younger leaves,
green islands in yellows areas of outer leaves, yellow and green zigzag bands
on leaets. Basal and middle leaves of delphinium have orange-amber or
lemon-yellow areas, chlorotic ring and line patterns; younger leaves are nor-
mal green. Transmission is by many species of aphids celery, celery-leaf,
rusty-banded, cotton, erigeron-root, foxglove, greenpeach, and honeysuckle.
Celery Mosaic Potyvirus
On celery, celeriac, and carrot in California. Young leaets are mottled green
and yellow, in advanced stages narrow, twisted, cupped; plants are stunted,
with central leafstalks shortened, outer in a horizontal position with rusty
necrotic spots. Transmission is by many species of aphids. A crinkle leaf
strain of this virus causes a yellow mottle with raised blister areas and crin-
kling on celery leaves.
Celery Yellow Dwarf Poleroviridae
Occurs on cereal crops in Alaska.
Celery Yellow Spot Luteovirus
On celery and parsnip in California. Yellow spots and stripes, mostly along
veins; circular white spots on petioles; transmission by honeysuckle aphid.
Cherry Albino
On sweet cherry in Oregon. Branches die back in spring; leaves golden
brown with up-rolled margins; late summer, new growth small and rosetted;
fruit small and white; trees soon killed. Transmission by tissue union.
Cherry Bark Splitting
On sour cherry and apricot.
Cherry Black Canker
On sweet cherry, Oregon. Black, rough cankers on twigs and branches.
Cherry Buckskin
Peach Western X-Disease (under Bacterial Diseases, Phytoplasma).

Cherry Bud Abortion
On sweet and sour cherry.
Cherry Chlorosis
On Malaheb and chokecherry.
Cherry Freckle Fruit Disease
On sweet cherry.
Cherry Green Ring Mottle
On sour cherry.
Cherry Gummosis
On sour cherry. Dieback of terminal shoots, excessive gumming of branches.
Cherry Little Cherry
On sweet and sour cherry; fruits are half-size. Flowering cherry acts as
a reservoir for the virus.
Cherry Midleaf Necrosis
Dark brown necrosis starting in midvein; heavy leaf fall; trees small and less
vigorous but fruit normal. On sour cherry in Oregon.
Cherry Mora
Abnormal delay of a month or more in ripening fruit; leaves yellowish, small,
rosetted, twisted on fruit spurs. On sweet cherry, Oregon.
Cherry Mottle Leaf Trichovirus
On sweet cherry in Northwest. Leaves show chlorotic mottling, are puckered,
wrinkled, distorted but not perforated. Fruit is small, hard, insipid, uneven or
delayed in ripening; crop is reduced. Branches are shortened, trees stunted.
Transmission by grafting or budding with no insect vector known, but the
disease spreads from wild bitter cherry to sweet cherry. Remove diseased
trees. Propagate with scions from virus-free trees.
Cherry Necrotic Rusty Mottle
On sweet cherry. Foliage and blossoms delayed in spring; brown necrotic or
rusty chlorotic spots, often with shot holes and defoliation.
Cherry (Sour) Pink Fruit
Fruit bitter, pink; tree stunted; foliage pale.
Cherry Pinto Leaf
On sweet cherry. Chlorotic patterns of varying size with disease tissue chang-
ing to bright yellow or white. Transmission by budding.
Cherry Rasp Leaf Nepovirus
Transmitted by nematode Xiphinema amerciana. Virus differs from other
cherry diseases that cause leaf enations.
Cherry Rasp Leaf Nepovirus
On sweet cherry, in Northwest. Enations, elongated protuberances from
underside of leaves with depressed lighter areas on upper surface.
Cherry Ring Spot =Prunus Necrotic Ringspot Ilarvirus
On sour cherry, sweet cherry (tatter leaf), peach, plum, prune, widespread in
Northeast. Chlorotic or necrotic rings and spots on leaves, with lacerations
to give the tatter-leaf effect. Transmission by grafting or budding, and, to
a small extent, seed. Control by testing budwood sources on a differential
host, such as Shirfugen variety of Prunus serrulata, sensitive to all strains.
Cherry (Flowering) Rough Bark
On Kwanzan owering cherry. Internodes shortened; leaves in clusters and
arched downward from necrosis and cracking of midribs; bark deep brown,
rough with longitudinal splitting; trees dwarfed.
Cherry Rugose Mosaic =Strain of Prunus Necrotic Ringspot
Ilarvirus
On sweet cherry. General chlorosis of leaf between midvein and margin,
with distortion; fruit yield reduced; fruits attened, angular. Transmission by
grafting; incubation 9 months.
Cherry Rusty Mottle
On sweet cherry. Many leaves turn bright yellow to red with islands of green,
and drop before harvest; remaining leaves have yellow-brown spots, rusty
appearance; fruit is small, late, insipid. Remove diseased trees. Select graft-
ing material from virus-free trees.
Cherry Twisted Leaf
On Bing cherry, severe stunting, leaves small, distorted, distal portion bent
abruptly downward; sometimes defoliation.
Cherry Vein Clearing
Sweet Cherry Crinkle. A viruslike disease but not transmissible; probably
genetic. Clearing of veins throughout leaves or in localized areas; margins
irregular; some leaves with elongated, slotlike perforations; small blisters
on lower side of veins, upper silvery. Leaves may fold along midrib, wilt
and drop in midsummer. Rosetting of some branches. Blossoms abnormally
abundant, but fruit reduced, pointed, attened on one side with swollen ridge.
Cherry Yellows =Prune Dwarf Ilarvirus
Widespread on sour cherry. Yellow areas enlarge to cover whole leaves;
defoliation. Diseased leaves and fruit larger than on healthy trees, but yield
reduced by half. Transmission by budding and through seed.
Chickpea Filiform Potyvirus
On chickpea. Found in Washington State, but there is no evidence of spread.
Chrysanthemum Aspermy
Tomato Aspermy Cucumovirus.

Chrysanthemum Chlorotic Mottle Viroid
Widespread in greenhouses and gardens. Bonnie Jean, Ridge, and Delaware
varieties are used as indicators.
Chrysanthemum Flower Distortion
Apparently not widespread in United States. Virus is carried without symp-
toms in leaves of White Wonder, but if this is grafted to Friendly Rival,
owers are extremely dwarfed and distorted, with ray orets short, narrow,
incurved, or irregularly curved.
Chrysanthemum Mosaic, Chrysanthemum B Carlavirus
Noordams B, Kellers Q, and other virus strains are widespread in chrysan-
themum with mild to severe leaf mottling and sometimes a brown streaking
of owers. Transmission is by grafting and by aphids. Control by indexing
tips from heat-treated plants on reliable test varieties to make sure they are
virus-free.
Chrysanthemum Ring Spot
Reported from Alabama in plants also aficted with Stunt. Large yellowish
chlorotic ring patterns, severe leaf dwarng and distortion.
Chrysanthemum Rosette
A strain from symptomless Ivory Seagull produces veinbanding, crinkle, dis-
tortion, rosetting on Blazing Gold.
Chrysanthemum Stunt Viroid
Widespread in greenhouses and gardens. Symptoms vary with variety, but
plants are dwarfed, with small owers and leaves and bloom earlier than
normal or later in some varieties. Blazing Gold, Blanche, Mistletoe, Daunt-
less, and Bonnie Jean often used as indicator varieties. Leaves of Blanche
are crinkled, and Mistletoe has a measles pattern. Transmitted by dod-
der, sap-inoculation, grafting with incubation period 6 weeks or longer; no
insect vector is known. Many plant species have been infected experimental-
ly. Commercial growers go to great lengths to select and reselect a virus-free
stock, and great care is taken to prevent recontamination. Garden varieties
are now indexed and available.
Cineraria Mosaic
Mottling, dwarng, and distortion of leaves, transmitted by seed, mechani-
cally, and by Aphis marutae
Citrus Enation Woody Gall Luteovirus
Found in different species of Citrus, Ulmus, Morus, Rosa, Prunus, Viburnum
and Ecualyptus.
Citrus Exocortis Viroid
Probably same as Rangpur lime disease; in Florida and Texas on red grape-
fruit and sweet orange trees, on Rangpur lime and trifoliate orange root-
stocks. Trees are stunted with bark shelling.
Citrus Leaf Rugose Ilarvirus and Citrus Variegation Ilarvirus
Disease known as citrus psorosis virus complex. Viruses spread in California
and caused psorosis of young leaves and malformation of the old ones.
Citrus Leprosis Rhabdovirus
Transmitted by mite-vector Brevipalpus phoenicis.
Citrus Psorosis Complex: Citrus Leaf Rugose Ilarvirus,
Citrus Ringspot Virus, Citrus Variegation Ilarvirus
Found wherever citrus is grown. Leaf symptoms are small elongated white
or yellow areas near veins. Bark symptoms are scales or small pustules with
irregular growth and gum deposits. With B strain of the virus, leaves have
dots, rings, or large translucent areas and small corky pustules; fruit has sur-
face rings. In the concave gum strain, cavities develop on trunks and larg-
er limbs. The blind-pocket strain usually produces troughlike depressions in
bark, sometimes bark scaling. The crinkly leaf strain, usually on lemon, caus-
es warping and pocketing of mature leaves, and rough, bumpy fruit. Trans-
mission is by budding or through natural root grafts.
Remove trees with advanced infection; use budwood from trees known to
be free from psorosis. Sometimes bark can be scraped, going several inch-
es beyond the margin of affected areas and painting the scraped areas with
bordeaux paste.
Citrus Ringspot Virus
Caused epinasty, chlorotic ecks, leaf mottling Ringspot and large irregular
chlorotic pattern on mature leaves of Citrus sp.
Citrus Stubborn Disease
Oranges have multiple buds, abnormal branching, acorn-shaped fruit, which
is sour and bitter at the navel end.
Citrus Tatter Leaf Cappillovirus
Reported from California and Texas on Meyer lemon and lime. Blotchy spot-
ting of younger leaves and ragged margins.
Citrus Tristeza Closterovirus
Quick Decline. In California, Florida, and Texas; usually in trees on sour
orange rootstock. First symptoms are partial or complete suppression of new
ushes of growth. Older leaves are dull or bronzed, later yellow. Defoliation
continues progressively from base of twigs to tip. Rootlets and then roots
die. Limbs die back, and weak shoots are produced from main limbs and
trunk. Transmission is by melon and other aphids. Make new plantings with
stock-scion combinations known to be resistant. Best rootstocks are sweet
orange, rough lemon, Rangpur lime, and sweet lime.
Citrus Vein Enation
In California on sour orange, Mexican lime, and other citrus. Veins swell and
enations develop on lower surface. Transmission by grafts and aphids.
Citrus Xyloporosis
Cachexia. In Florida, chiey on sweet lime rootstock, but also on mandarins
and some of the tangelo oranges. Symptoms include stunting of 2- or 3-year
trees, small yellow leaves, partial leaf drop, early blooming and fruiting; hor-
izontal growth of branches in middle section of trees; dieback, followed by
decay of entire trunk and roots. Fruits are more rounded, with a thicker rind.
Transmission is by budding, possibly through seed. Use resistant rootstocks,
as sour oranges of Israel and Bagdad, Valencia orange.
Citrus Yellow Vein
In California on limequat. Petioles and veins are bright yellow.
Clerodendron Zonate Ring Spot
In Florida on bleeding-heart vine. Chief symptoms are cleared veins.
Clover Club Leaf
On crimson clover. Young leaves are light-colored, have club leaf appearance
due to delayed opening. Yellow margins of leaves turn red or purple during
summer. Transmission by a leafhopper.
Clover (Alsike) Mosaic =Clover Yellows Vein Potyvirus
On pea, causing chlorotic spotting and dark green banding of veins, leaves
slightly cupped or distorted. Leaf puckering and plant stunting on bean. Pea
aphid is vector.
Clover (Red) Vein Mosaic Carlavirus
On garden pea, causing pea stunt, broad bean, sweet pea, and red clover.
Vein clearing and chlorosis are chief symptoms with, on peas, curling of
leaves and rosetting of younger shoots, wilting, and collapse. Vein clearing
is the only symptom on sweet pea. Broad beans may be stunted and killed.
Transmission is by the pea aphid without incubation period or long retention.
The Wisconsin pea stunt virus may be a strain of the red clover virus.
Clover Yellow Mosaic Potexvirus
Transmitted by mechanical inoculation; serologically related to White
Clover Mosaic, Cactus X and Hydrangea Ringspot Potexviruses.
Clover Yellow Vein Mosaic
On winged bean, wild carrot, poison hemlock, and red bean.
Clover Wound Tumor Phytoreovirus
Big vein disease, causing enlargement of veins, sometimes with enations,
woody tumors on roots, sometimes stems. The virus was discovered acci-
dentally in leafhopper nymphs and has been transmitted experimentally to
many plants besides clovers.
Cocksfoot Streak Potyvirus
Caused chlorotic streaks on leaves of Dactylis glomerata.
Coleus Mosaic =Cucumber Mosaic Cucumovirus
Reported from Illinois on coleus, symptoms varying with variety. Leaves
may be puckered, crinkled, asymmetrical, with oak-leaf markings or ring
spots or small necrotic spots.
Commelina Diffusa Potyvirus and Commelina Mosaic Potyvirus
Both viruses infected Commelina diffusa and Rhoeo spatecea on which
mosaic or systemic mottling symptoms occurred. Both viruses induced
pinwheel structures in the cytoplasm.
Coriander Feathery Red Vein Nucleorhabdovirus
Virus multiplies in vectors (Myzus perscicae and Hydaphis foeniculi) and is
transmitted congenitally to the progeny of the vector.
Corn Leaf Fleck
On eld and sweet corn in California. Small, circular pale spots on leaves
with tip and marginal burning, leaves dying 7 to 10 days after initial symp-
toms. Transmission by corn, peach, and apple grain aphid, which retain the
virus for their entire lives.
Corn (Sweet) Mosaic
Leaves have broken or continuous interveinal chlorosis.
Cotton Leaf Curl Bigeminivirus
Caused chlorosis, thickening and malformation of veins, petioles and leaves
on Gossypium barbadense and Hibiscus esculentus plants.
Cowpea Chlorotic Mottle Bromovirus
Reported in peanut, common bean, cowpea and soybean plants.
Cowpea Mosaic Comovirus
Clearing of veins is followed by chlorotic mottling, slight convex cupping
of leaets, shortened internodes, abortion of owers, twisting of petioles,
and delayed maturity. Yield is reduced. Vectors are potato, pea, and cotton
aphids. Another cowpea mosaic, known in Trinidad and probably the same
as one in the United States, is transmitted by bean leaf beetles. May infect
soybeans, hoary-tick clover (Desmodium canescens).
Cranberry False Blossom
The most serious cranberry disease in Massachusetts, New Jersey, and Wis-
consin; known also on the Pacic Coast. American and European cranber-
ries are the only natural hosts, but the virus has been dodder-transmitted
to other plants. Cranberry owers are erect, instead of pendent, with calyx
lobes enlarged, petals short, streaked with red or green, stamens and pis-
tils abnormal. Flowers may be replaced by leaves or short branches. Axil-
lary buds produce numerous erect shoots forming witches brooms; diseased
fruits are small and irregular. Transmission is by the blunt-nosed leafhopper.
Select strains resistant to the vector or ood the bogs after leafhoppers have
hatched. Spray with pyrethrum.
Crimson Clover Latent Nepovirus
Symptomless on Trifolium incarnatum.
Cucumber Mosaic Cucumovirus
General with many strains in cucumber, squash, melon, winged bean, peri-
winkle, wild violets, desert-rose, and a wide range of other plants, includ-
ing spinach, where the disease is called blight; tomato, causing shoestring
disease with liform leaets; pepper, petunia, grlic mustard, fuschia, and
tobacco. Wintering is on ground cherry, milkweed, pokeweed, catnip, Texas
bluebell and Peristroph sp. and other weed hosts. Transmission is by peach,
cotton, potato, and lily aphids and, in some cases, through seed.
In cucurbits there is a yellow-green systemic mottling, with leaves small,
distorted, curled, plants dwarfed with shortened internodes, few fruits set
and those mottled and misshaped, a condition called white pickle. The lily
mosaic strain produces a masked infection or chlorotic mottling and necrosis
when mixed with Lily Symptomless Virus. The lima-bean, southern celery
mosaic, and cowpea strains cause chlorotic mottling.
Geraniums are stunted and mottled; gladiolus owers are color-broken;
dahlia foliage has oakleaf patterns; periwinkle (myrtle) has a streaky mottle,
down-curved leaves, small owers with a white streak in the blue color. Petu-
nias have distorted leaf blades, few or no blossoms. In delphinium, which is
very susceptible, the disease is called ring spot, stunt, witches broom.
Control. Resistant varieties of spinach, cucumber, and squash are available.
Diseased lilies and other owers should be rogued immediately. Control
aphids by systemic ground treatments or sprays; repel aphid vectors by an
aluminum foil mulch. Lilies may possibly be freed of the virus by scale prop-
agation.
Cucurbit Leaf Curl Begomovirus
On pumpkin, cucumber, muskmelon, honeydew, zucchini, banana squash,
and squash.
Cucurbit Leaf Crumple Begomovirus
On cucumber.
Cucurbit Yellow Stunting Crinivirus
On melon.
Currant (Red) Mosaic =Tomato Ringspot Nepovirus
Irregular, light green circular spots along midrib and larger veins enlarge to
bands. Canes are stunted; plants decline.
Cymbidium Mosaic Potexvirus
Virus was isolated from many orchid species showing mosaic symptoms. It
is transmitted by contact between plants.
Dahlia Mosaic Caulimovirus; Stunt
General. Bands along midrib and veins remain yellow-green. In some vari-
eties leaves are distorted and blistered; in others, leaves are yellowed with
margins up-rolled; in others, plants are very short and bushy with short ow-
er stems. Transmission is by peach and other aphids.
Dahlia Oakleaf
May be a separate entity or a strain of Tomato Spotted Wilt Tospovirus.
A pale chlorotic line across the leaf suggests the outline of an oak leaf.
Dahlia Ring Spot
Caused by a strain of Tomato Spotted Wilt Virus Tospovirus. Leaves have
concentric rings or irregular zigzag markings. In Utah a yellow strain causes
bright yellow rings and zigzags.
Dasheen Mosaic Potyvirus
Widespread in plants from Araceae family.
Delphinium Ring Spot
Faint chlorotic rings around green and yellow centers appear on young
leaves, irregular necrotic spots or rings with yellow bands on mature leaves.
Desmodium Mosaic Potyvirus
Transmitted by aphids, sap and seeds. Leaf sap contains only a few lamen-
tous (775 nm length) virus particles.
Desmodium Yellow Mottle Tymovirus
Transmitted by mechanical inoculation. Leaf sap contains many isometric
(30 nm diameter) virions.
Diodia Vein Chlorosis Closterovirus
Transmitted by Trialeurodes abultilonea; caused chlorotic vein banding and
chlorosis of Diodia virgianiana plants.
Dodder Latent Mosaic
Three species of dodder transmit mosaic to cantaloupe, potato, tomato, and
celery.
Dogwood Mosaic Nepovirus
Considered earlier as a strain of Arabis Mosaic Nepovirus.
Elderberry Disease
=Elderberry Carlavirus, Elderberry Latent Carmovirus
A virus disease reported from golden elderberry can infect various stone
fruits and is considered a potential threat to the fruit industry.
Elm Mosaic
On American elm in Ohio and eastern states. Some leaves are larger than
normal, others small, distorted, with yellow and green mottling. There may
be some branch brooming, gradual decline in vigor. Transmission is by graft-
ing; no insect vector is known.
Elm Zonate Canker
On American elm, New Jersey, Ohio, Missouri. Zonate cankers appear in
bark as rings of dead and living tissue in cortex or phloem. Some leaves
develop brown necrotic spots. Transmission is by bark patch grafts; no insect
vector is known.
Euonymus Mosaic
Infectious Variegation. Persistent yellowing along veins; transmission by
grafting and budding.
Euphorbia Mosaic Begiminivirus
Spreads in Florida by nymphs and adults of Bemisia tabaci. Infected plants
showed mosaic and leaf malformation.
Fig Mosaic
Systemic chlorotic mottling is accompanied by severe leaf distortion; fruits
have light circular areas or rusty spots, may be deformed and drop prema-
turely. Transmission is by grafting and the g midge (Aceria cus).
Figwort Mosaic Caulimovirus
The viral DNAhas little homology with that of Cauliower Mosaic Caulimo-
virus.
Filaree Red Leaf Luteovirus
On Erodium in California. Early symptoms are mild vein clearing, outward
curvature of petioles, inward cupping of leaets. Later leaets cup outward,
with reddish discoloration, are brittle, with petioles stify upright; owers
are dwarfed or suppressed. Aphids are vectors. Caused vein clearing and
chlorosis of young leaves; older leaves are red and breakable. Infected plants
are stunted and do not develop owers.
Foxtail Mosaic Potexvirus
Virus related to Narcissus Mosaic and Viola Mottle Potexvirus.
Geranium Chlorotic Spot
General on geranium. It is caused by tomato and tobacco ring spot nepovirus-
es.
Geranium Crinkle
Pelargonium Leaf Curl Tombusvirus. General on geranium. Hyaline spots
are small, circular to irregular, sometimes star- or tree-shaped, with brown
centers. Young leaves are crinkled, small, sometimes puckered and split;
severely infected leaves turn yellow and drop. Petioles and stems have corky,
raised necrotic streaks; tops may die. The disease is most severe in spring,
inconspicuous in summer. Transmission is by grafting (not by knife prepar-
ing cuttings) and probably by whiteies.
Geranium (Pelargonium) Mosaic
Leaf Breaking. Leaves smaller, with purple spotting along veins, and sup-
pression of horseshoe pattern in foliage of some geranium varieties.
Gladiolus Mosaic
Bean Yellow Mosaic Potyvirus.

Grapevine Fanleaf Nepovirus
Infectious degeneration in California; Court-nou and Roncet in Europe.
New growth is severely stunted; leaves are dwarfed and puckered or with
deep indentations and folded like a half-closed fan; fruit set is poor. The
virus is present in soil and can be transmitted, apparently, by nematodes.
Grapevine Fleck Virus
Generally symptomless in most cultivars; transmitted only by grafting.
Grapevine Rupestris Vein Feathering Maravirus
On grape.
Grape Leaf Roll =Grapevine Leafroll-Associated Closterovirus
White Emperor Disease. In California, restricted to Emperor variety. Fruit
is greenish yellow or pink rather than normal red; leaves are darker than
normal, turning bronze or reddish along veins, yellow between veins.
Grapevine Stem Pitting Associated Chlosterovirus
Virions (800 nm length) found in phloem, but they are very difcult to extract
from host tissue. Virus related only to Citrus Tristeza Closterovirus.
Grape Yellow Mosaic F Strain of Grapevine Fanleaf Nepovirus
(Panachure)
Yellowing of leaves of young shoots in some varieties; various types of leaf
mottling; blossom shedding. Transmission by grafting and in soil.
Grape Yellow Vein =Tomato Ringspot Nepovirus
Can be transmitted by dagger nematodes.
Guar Symptomless Potyvirus
Symptomless or caused mild green mottle on Cyanopsis tetregonoloba.
Hellenium S. Carlavirus
Hellenium and Impatiens strains differ in host range and aphid transmissibil-
ity.
Henbane Mosaic
Clearing or yellowing of veins of youngest leaves, then a yellow mosaic and
dark green vein banding.
Hibiscus Chlorotic Ringspot Carmovirus
Caused very variable symptoms on the leaves of Hibiscus rosa-sinensis; from
mottling and chlorotic spots to vein banding and rings.
Hibiscus Latent Fort Pierce Tobamovirus
On Hibiscus.
Hippeastrum Mosaic Potyvirus
Virus known also as Amaryllis Mosaic Virus, caused chlorotic streaking on
leaves and ower stalks.
Hollyhock Mosaic
Pronounced yellow and green mottle on hollyhock and malva.
Holodiscus Witches Broom
On ocean spray. Diseased branches form clusters of thin, wiry shoots with
abnormally short internodes, crowded small leaves; foliage turns bronze red
early. Transmission by the spirea aphid and by grafting.
Hop American Latent Carlavirus, Hop Latent Carlavirus,
Hop Mosaic Carlavirus
All three carlaviruses are distantly serologically related. Usually they are
symptomless or show mild mosaics.
Hyacinth Mosaic Potyvirus
Spreads by aphids. Virus caused chlorotic spots on leaves, stunt and color-
breaking of owers.
Hydrangea Latent Carlavirus
Reported on Hydrangea macrophylla in U.S.A.
Hydrangea Phyllody Phytoplasma
Witches Broom, green owers.
Hydrangea Ring Spot Potexvirus
Chlorotic blotches and rings, brown rings and oak-leaf patterns are common
in orists hydrangea. A probable cause of hydrangea running out. Trans-
mission is by cutting knife. Virus can infect snapdragon, sweet william and
globe amaranth.
Iris Fulva Mosaic Potyvirus and Iris Severe Mosaic Potyvirus
Both potyviruses are spread generally by aphids. For diagnostic purposes
they can be easily separated by pinwheel inclusions morphology.
Iris Mosaic =Iris Mild Mosaic Potyvirus
Widespread on bulbous iris, especially serious on Pacic Coast. Plants are
stunted with yellowish streaks on leaves and dark, teardrop markings on
white, blue, or lavender owers, clear feathery markings on yellow owers.
Transmission is by peach and potato aphids. Establish disease-free founda-
tion stock; rogue all diseased plants; spray for aphids.
Iris Yellow Spot Tospovirus
On Allium sp. Including A. cepa.
Impatiens Latent Potexvirus
Symptomless virus transmitted only by mechanical inoculation.
Impatiens Necrotic Spot Tospovirus
Previously called strain of Tomato Spotted Wilt Tospovirus. It is the most
common and most damaging virus in the greenhouse industry. The virus
has an extremely broad host range and its vector, the western ower thrips,
Frankliniella occidentalis, are widespread and difcult to control. Symp-
toms caused by virus were variable but generally occurred as brown or black
necrotic spots, blotch, necrotic rings on leaves and ower distortion. Often
the center leaves collapse. Infected plants are stunted and die prematurely.
The control of vector is essential in greenhouses.
Ixia Mosaic
Perhaps iris mosaic.
Johnsongrass Mosaic Potyvirus
Known also as Maize Dwarf Mosaic Virus strains 0 and Kansas and as Sug-
arcane Mosaic Virus. On Johnsongrass virus caused systemic mosaics.
Kalanchoe Top-Spotting Badnavirus
Kalanchoe blosfeldiana plants showed sunken yellow spots and sometimes
leaf deformation.
Laburnum Mosaic
Infectious variegation. Bright mottling of foliage, often with veins picked out
in yellow.
Lettuce Big Vein Varicosavirus
Transmitted by a vector-fungus, Olpidium brassicae. Vein clearing followed
by enlarging and bleaching
Lettuce Infectious Yellows Closterovirus
Virus infected lettuce, sugarbeets, cantaloupe, watermelons, melons, squash,
and carrots. Main symptoms included chlorosis, reddening and brittle leaves;
plants stunted.
Lettuce Speckles Mottle Umbravirus
Virus in mixed infection is encapsidated in coat protein of Beet Western
Yellows Luteovirus.
Lettuce Mosaic Potyvirus
Widespread on lettuce. Leaves mottled, deformed, yellowed, browned; plants
stunted or dead. Transmission is by peach and root aphids and in seed. Con-
trol vectors, use virus-free seed; rogue seedbeds.
Lilac Mottle Carlavirus
Caused leaf chlorosis and mottling on Syringa spp.
Lilac Ring Spot Carlavirus
Pale green to yellow spots, lines, broad diffuse rings, and bands on lilac
leaves, often with distortion and holes in tissue.
Lilac Witches Broom Phytoplasma
On lilac, privet in Maryland. Brooming symptoms; lateral buds produce two
to six slender shoots, which branch freely, with very small leaves on Japanese
lilac. In common lilac and Regal privet there is yellow vein clearing with less
prominent brooming. Transmission by grafting, and by dodder; no insect
vector known.
Lily Color Adding
On lily.
Lily Color Removing
On lily.
Lily Fleck
Caused by Lily Symptomless Carlavirus and Cucumber Mosaic Cucu-
movirus. Yellow ecks on Easter lily leaves change to gray or brown,
elongating parallel to veins; surface is depressed but unbroken. Plants are
dwarfed with curled leaves, owers small with brown streaks.
Lily Latent Mosaic
In Easter lily and tulip, symptoms masked or systemic chlorotic mottling.
Lily Mottle Potyvirus
It is a lily strain of Tulip Breaking Potyvirus.
Lily Ring Spot =Cucumber Mosaic Cucumovirus
Possibly a form of cucumber mosaic. There is only a faint mottling on some
species, but on Lilium tigrinum and L. regale dark ring markings develop
into necrotic areas. The growing point is killed; no owers are formed; whole
plant is twisted, stunted, deformed. Peach aphid is the vector.
Lily Rosette
Yellow Flat. On Easter lily. Leaves curl downward; plants are dwarfed, yel-
lowed, mature early; bulbs are small. Transmission by the melon aphid, not
by seed. Rogue diseased plants; spray for aphids.
Lily Symptomless Carlavirus
Present in Easter lilies wherever grown commercially but producing no
symptoms alone; in combination with Cucumber Mosaic Cucumovirus
causing necrotic eck. Transmission by melon aphid.
Locust Witches Broom
Brooming disease on locust from Pennsylvania to Georgia, Ohio, and Ten-
nessee. Vein clearing is followed by reduction in size of new leaves, growth
of spindly shoots to witches brooms. Roots are more brittle, shorter, and
darker than normal; rootlets branch excessively to root brooms. Transmis-
sion is by budding and grafting; no insect vector is known.
Loganberry Dwarf
Blackberry Dwarf. On loganberry and phenomenal blackberry in Northwest.
Leaves are small, obovate, rigid, with new canes short and spindly. Young
plants have crinkled leaves with some chlorosis or necrosis along veins.
Flowers are small, drupelets ripen unevenly and tend to fall apart when fruit
is picked. Transmission is by aphids.
Lolium Latent Potexvirus
On ryegrass (Lolium perenne and L. multiorum).
Lonicera Infectious Variegation
Vein yellowing and variegation on honeysuckle; graft-transmitted.
Maize Dwarf Mosaic Potyvirus
First noted in Ohio in 1962 and since devastating to corn in many states. Red
to purple streaks in upper leaves, ears usually incomplete; plants dwarfed
with great reduction in yield. Transmission by corn leaf and peach aphids.
Occurs also on Sorghum sp. and Triticum sp.
Maize Chlorotic Mottle Machlovirus
Usually occurred together with many potyviruses on infected maize plants
showing severe systemic necrosis and stunting. Plant often dying.
Maize Rayado Fino Maravirus
Initially described as a strain of Corn Stunt Virus now in the type species
of Maravirus. Virus frequently occurred in mixed infection with Phytoplas-
mas. It is multiplied in the vector, Dalbulus maydis, Cocadellidae and is not
transmitted by sap, grafting or seeds.
Maize Stripe Tenuivirus
Caused yellow striping and chlorosis on natural hosts. Zea mays, Sorghum
vulgare, S. bicolor, Triticum aestivum and Hordeum vulgare.
Maize Stunt =Maize Chlorotic Dwarf Waikavirus
Corn Stunt.
Maize White Line Mosaic
On eld corn and weed hosts including Panicum, Setaria and Digitaria.
Malva Vein Clearing Potyvirus
Caused vein clearing and yellow mosaics on many Malva and Lavatera
species. Virus is known as Malva Mosaic, Malva Green Mosaic or Malva
Yellow Vein Mosaic Virus.
Melon Leaf Curl Bigeminivirus
Caused leaf mottling and chlorosis on Cucurbits.
Melon Necrotic Spot Carmovirus
Transmitted by fungus Olpidium radical, by sap, by contact between plants
and by seeds. It caused chlorotic and necrotic spots on Cucurbits.
Mimosa Striped Chlorosis Badnavirus
Infected Albizzia sp. plant showed chlorotic stripes along leaf vein and inter-
veinal chlorosis.
Mint Crinkle Closterovirus
On golden ginger mint.
Mirabilis Mosaic Caulimovirus
Genome consists of DNA, virus-transmitted in semi-persistent manner by
aphid, Myzus persicae and by sap.
Muskmelon Mosaic =Squash Mosaic Comovirus
Widespread on melon. First leaves have dark green bands parallel with main
leaf veins; later leaves are mottled, sometimes deformed. Transmitted by
seed and sap; insect vectors unknown
Muskmelon Vein Necrosis Carlavirus
Caused vein necrosis and leaf chlorosis on cucumber and pea plants.
Mustard Mosaic
On black mustard, California. Small, brown local lesions are followed by
a general mottling.
Nandina Mosaic Potexvirus
Occurs in California; transmitted by mechanical inoculation.
Narcissus Chocolate Spot
Often present with white streak in a decline complex.
Narcissus Flower Streak
Strong breaking of owers but normal foliage in Oregon bulb crops.
Narcissus Mosaic Potexvirus
Widespread on narcissus, but with mild symptoms, seldom apparent before
plants bloom; has been confused with yellow stripe.
Narcissus White Streak
Silver Leaf. Paper tips and white streaks in leaves are primary symptoms,
with wilting and falling over of foliage long before harvest so bulbs are small.
Causes decline combined with chocolate spot. Transmission by aphids.
Replant only the largest bulbs.
Narcissus Yellow Stripe Potyvirus
Strong yellow streaking and mottling of foliage, often roughened near veins
and with a peculiar twist. Flowers are streaked. Transmission by several
species of aphids. Select the best plants for a mother block, with nal selec-
tion during bloom; rogue plantings early before symptoms are masked by
hot weather.
Nasturtium Mosaic Potyvirus
Vein clearing, rufing and cupping of young leaves, dark green vein band-
ing in older leaves, sometimes chlorotic spots or white rings between veins.
Flower color may be broken, petals crinkled. Transmitted by several aphids.
Nothoscordum Mosaic Potyvirus
False garlic (wild amaryllis) mosaic transmitted through bulbs but not seed.
Typical mosaic mottling of foliage.
Oak Ringspot Virus
Caused mosaic, chlorotic ringspot and oak leaf pattern on the older leaves.
Oat Blue Dwarf Maravirus
Synonym for Flax Crinkle Virus transmitted by aster leafhopper. Infected
Avena sp. plants turned deep blue.
Oat Golden Stripe Furovirus
Transmitted by fungus, Polymyxa graminis; systemically infected Avena sp.
plants.
Oat Mosaic Bymovirus
Only Avena sp. are susceptible (developing mottle). Virus transmitted by fun-
gus, Polymyxa graminis.
Odontoglossum Ringspot Tobamovirus
It is synonym for orchid strain of Tobacco Mosaic Tobamovirus.
Onion Yellow Dwarf Potyvirus
Yellow streaks develop at base of leaves, with yellowing crinkling, and at-
tening of new leaves. Leaves may be prostrate, ower stalks bent, twist-
ed, and stunted; yield is reduced. Some species are relatively tolerant; tree
onions are symptomless. Bean, apple-grain, corn leaf, and other aphids are
vectors. Control is by indexing, growing sample lots of sets and mother bulbs
in greenhouse beds or production of virus-free stocks in areas where disease
is absent, and roguing of infected volunteer onions. Some varieties are resis-
tant to the onion strain of the virus but not to the strain from shallot or garlic.
Opuntia Sammons Tobamovirus
Symptomless on Opuntia spp., virus is transmitted only by mechanical inoc-
ulation.
Orchid (Cattleya) Blossom Brown Necrotic Streak
Brown spots, streaks of whole ower; leaves may have yellow streaks; trans-
mission by knife. In removing ower spikes use hot knife, with attached
propane torch.
Orchid Fleck Rhabdovirus
Caused chlorotic and necrotic eck on many orchid species. Virions are typ-
ical rhabdo or bullet-shaped.
Orchid (Cattleya) Mosaic Potexvirus
Flower-Break. On Cattleya and other orchids. There are apparently two dis-
eases: mild color break, with variegation in the ower but no distortion,
and severe color break, with owers distorted or twisted as well as varie-
gated. Leaves are mottled and sometimes twisted. The virus may be present
in apparently healthy plants but can be detected with antisera, and infected
plants removed. Transmission is by the green peach aphid.
Orchid (Cymbidium) Mosaic Potexvirus
Black Streak; Cattleya Leaf Necrosis. The most common virus disease on
many kinds of orchids. On Cymbidium there is initially a mosaic mottle,
then necrotic spots, streaks, and rings on leaves but no effect on owers.
In Cattleya there are sunken brown to black leaf patterns, sometimes rings,
more often elongated streaks on older leaves. If leaves are killed prematurely,
owers are fewer and smaller but normal in form and color. No insect vector
is known.
Orchid (Odontoglossum) Ring Spot Tobamovirus
On Odontoglossum only. Small, necrotic spots and rings on older leaves,
light green to yellow areas on young leaves. Leaves may turn yellow and
drop in 2 or 3 months or persist longer. There are no ower symptoms; no
insect vector is known.
Orchid (Oncidium) Ring Spot
On mature leaves of Oncidium; round to irregular, slightly sunken yellow
areas on both leaf surfaces; becoming necrotic with age.
Orchid (Vanda) Ring Spot
On 22 cultivated orchid species.
Ornithogalum Mosaic Potyvirus
On ornithogalum, galtonia, hyacinth, lachenalia, agapanthus, hebe, ne light
and dark green leaf mottling becomes gray or yellow as leaves mature. Flow-
er stalks are marked with light and dark green blotches; there are thin lon-
gitudinal streaks on perianth segments. Transmission is by melon, peach,
potato and lily aphids.
Palm Mosaic Potyvirus
Infected Washingtonia robusta plants showed mosaics, ringspots and line
pattern. Plants are often stunted.
Panicum Mosaic Sobemovirus
On St. Augustine grass.
Papaya Mosaic Potexvirus
Caused mosaic and stunting on Carica papaya.
Papaya Ringspot Potyvirus
Symptoms on infected Carcia papaya plants included mottling, malforma-
tion of leaves and streaking on owers and fruits.
Pea Enation Mosaic Enamovirus
On pea, sweet pea, broad bean, soybean and sweet clover. Symptoms are yel-
lowish spots on leaves, which are later white, with crinkling and savoying.
Very susceptible varieties like Alderman have necrotic spots and prolifera-
tions or enations from underside of leaves. Pods may be markedly distorted
and twisted with seeds small and yellow. Transmission is by pea, potato, and
peach aphids.
Pea Mosaic Potyvirus
On pea, sweet pea, red clover and broad beans. Sweet pea has leaf mottling,
chlorosis, breaking of ower color. Garden pea has vein clearing followed
by mottling or general chlorosis and stunting. Transmission is by pea, peach
and bean aphids. Perfection and Horal varieties are resistant to this virus but
not to pea enation mosaic.
Pea Mottle
Pea Mottle caused by Clover Yellow Mosaic Potexvirus and Clover Yellow
Vein Potyvirus. Fairly widespread on garden pea, snapbean, white clover
and broad bean. On pea a severe systemic mosaic may be fatal. Some plants
have chlorotic mottling of leaves and stipules, but stems, pods, and seeds are
normal. Bean and pea aphids are probably vectors.
Pea Streak Carlavirus
Light brown to purple, oblong, necrotic lesions are scattered along stems and
petioles with stems often girdled. Leaves and pods are roughened with light
brown necrotic areas.
Pea Wilt
Causing severe streak in pea if pea-mottle virus is also present.
Peach Asteroid Spot
Discrete, chlorotic lesions spread along veins forming starlike spots; some
chlorophyll is retained in lesions as leaves turn yellow.
Peach Calico
Leaves are rst mottled, then yellowed, then papery white. Creamy white
streaks develop on twigs. Fruit is shorter, rounder, with creamy white to red
patches. Transmission is by budding.
Peach Dwarf
Only on Muir peach. Profusion of large, at, dark green leaves, closely
appressed on short twigs; witches broom showing in dormant period; fruit
larger than normal, misshapen.
Peach Golden Net
Probably identical with line pattern.
Peach Little Peach
Related to peach yellows, and in same host range, eastern United States.
Peach Mosaic
In Southwest on peach, apricot, nectarine, plum and capable of infecting
almond. Spring growth of peach has short internodes, with sometimes ow-
er breaking, chlorotic mottling, and foliage distortion early in the season,
with masking of symptoms or dropping out of affected areas in midsum-
mer. Fruit is small, irregular in shape, unsalable. Apricot stones have white
rings and blotches. Transmission is by budding, grafting, a mite (Eriophyes
insidiosus), and perhaps the plum aphid. Removal of infected trees, nursery
inspection and quarantine reduce the incidence of mosaic.
Peach Mottle
Known only in Idaho.
Peach Necrotic Leaf Spot
On peach but with sweet cherry as a symptomless carrier. Light brown, dead,
membranous areas in unfolding leaves fall out, leaving holes. The disease is
recurrent on peach.
Peach Phony Disease
The most important peach disease in the Southeast. Trees are dwarfed;
foliage is abnormally green, fruit small; there are ecks in wood, especially
in roots. Phony trees have short terminal and lateral twigs; profuse lateral
branching. Growth starts in spring earlier than on normal trees. Production
gradually decreases, with trees worthless in a few years. Transmission is by
root grafting and sharpshooter leafhoppers. Control has been by eradication
and by quarantine to restrict movement of nursery stock.
Peach Red Suture
Probably a form of yellows. On peach and Japanese plum. Fruit ripens pre-
maturely with softening, swelling, and red blotching on the suture, esh
coarse and watery while rest of fruit is hard and green. Eradicate diseased
trees; propagate from healthy budwood.
Peach Ring Spot Ilarvirus
Cherry Ring Spot.

Peach Rosette
On peach and plum. Trees suddenly wilt and die, or there are abnormally
short stems bearing dwarfed leaves, with veins cleared and thickened; death
follows in a few months. The virus can be inactivated by heating at 122F
for 10 minutes.
Peach Rosette Mosaic Nepovirus
Of minor importance on peach, highbush blueberry, and plum. Delayed foli-
ation, chlorotic mottling, rosetting of shoots, dark green color; transmission
by grafting and through soil. Eradicate trees; do not replant in same soil
without fumigation.
Peach Stubby Twig
A new disease of peach and nectarine in California. Chlorotic leaves, stubby
twig growth, decreased fruit production; transmitted with infected budwood.
Peach Wart
Foliage is normal but fruits are blistered, welted, and have conspicuous
raised warty outgrowths. Tissues are light tan to red, rough, cracked, and
russeted or smooth, with severe gumming. Transmission by budding or
inarching.
Peach Yellow Bud =Tomato Ringspot Nepovirus
Winters Peach Mosaic. On peach, apricot and almond in California. Pale
yellow, feather-edged blotches along the midvein with leaf distortion, and
defoliation near base of shoots. Transmission is by grafting. In eld spread
is only to adjacent trees, perhaps through soil.
Peanut Mottle Potyvirus
On wild peanut (Arachis chacoense).
Peanut Stunt Cucumovirus
First noted on peanuts in Virginia in 1964 and also occurs in bean, red and
white clover. Severe dwarng and malformation of foliar parts and suppres-
sion of fruit development. Transmission by grafting and green peach aphid.
Peanut Top Paralysis Potyvirus
Found in Oklahoma but there is no evidence of spread.
Pear Decline Phytoplasma
A relatively new and devastating disease in California, Oregon, and Wash-
ington, trees showing a slow decline or rapid collapse. First thought due to
a toxin of the pear psylla (Psylla pyricola), now considered a virus disease
transmitted by the psylla.
Pear Stony Pit
On Bosc and other pears in Pacic Northwest. Dark green areas appear just
beneath epidermis of fruit, 10 to 20 days after petal fall, resulting in deeply
pitted or deformed fruit at maturity, with corky or necrotic hard tissue at base
of pits. The fruit is gnarled, hard to cut. Transmission is by grafting; no insect
vector is known. Bosc and Anjou pears can be top-worked with Bartlett to
reduce losses from stony pit.
Pelargonium Flower Break Carmovirus
and Pelargonium Line Pattern Carmovirus
Viruses are usually symptomless on Pelargonium or caused by ower streak-
ing, line pattern or chlorotic spotting. They are transmitted by grafting and
by contact between plants.
Peony Leaf Curl
Plants half normal height, with crooked ower stalks, curled leaves. Trans-
mission is by grafting but not contact; no insect vector is known.
Peony Ring Spot =Tobacco Rattle Tobravirus
Marked yellow mosaic, irregular or in rings, sometimes small necrotic spots.
Peperomia Ring Spot
Concentric brown, necrotic rings on leaves, which may be cupped, curled, or
twisted and may fall. Severely affected plants are stunted. Grower losses in
Florida may be 25%. Take cuttings from healthy, vigorous plants.
Pepper Mild Mottle Tobamovirus
Infectious virus particles were found in water for greenhouse irrigations.
Pepper Mild Tigre Bigeminivirus
and Pepper Texas Bigeminivirus
Both viruses are transmitted by Bemisia tabaci and spread in Texas and Mex-
ico.
Pepper Strain of Alfalfa Mosaic
On pepper.
Pepper Vein Banding Mosaic
Probably caused by Potato Y Potyvirus, a new disease in Florida. Plants are
stunted with up to 50% loss of marketable fruit. Vein clearing and banding
on leaves, fruit roughened with chlorotic spots or stripes. Transmission is by
green peach and melon aphids. Eradicate deadly nightshade as a weed host
for 150 feet from peppers, or use sunower as a barrier.
Pepper Mottle Potyvirus
On pepper.
Phlox Streak
Streaks evident in leaves and stems. Clearing of veins is followed by necrosis
in leaf veins and petioles. Graft-transmissible.
Physalis Mosaic Tymovirus
Known as Belladonna Mottle Virus caused by systemic mosaic or mottle.
Pigeon Pea Golden Mosaic Begomovirus
On pigeon pea.
Pineapple Wilt Associated Closterovirus
Occurs in the Hawaiian pineapples causing yellowing and ecks on leaves,
leaf tip necrosis and plant wilt.
Plantago Mottle Tymovirus
Related to Andean Potato Latent, Dulcamara Mottle, Belladonna Mottle,
Onion Yellow Mosaic and Turnip Yellow Mosaic Viruses.
Plum Line Pattern =Plum American Line Pattern Ilarvirus
On plum, oriental cherry, widespread. Some plum varieties have yellow vein
banding, brilliant green and yellow patterns of the oakleaf type, formed by
single or multiple irregular lines or bands; in early summer the yellow fades
to creamy white. In other varieties patterns are faint or absent. On owering
cherries discolored areas are bounded by a chlorotic band. Transmission is
by budding or grafting.
Plum White Spot
Small pale yellow to white spots, mostly aggregated near leaf tips on Santa
Rosa plum.
Poinsettia Mosaic Tymovirus
Caused systemic leaf mosaic and malformation and discoloration of bracts.
Poplar Decline Potyvirus
Pinwheel structures are induced in infected cells. Main disease symptoms
are chlorotic and necrotic leafspots, necrosis of leaves and death of branches
and even whole trees.
Poplar Mosaic Carlavirus
Many cultivars are symptomless or they develop mosaics or necrosis. Virus
usually occurs alone in infected poplar trees.
Potato A Potyvirus
Known as Potato Mild Mosaic, Potato Virus P and Solanum Virus 3. Trans-
mitted by aphids in non-persistent manner. Virus caused no symptoms.
Potato Acropetal Necrosis
Caused by potato viruses Y and X.
Potato Aucuba Mosaic Potexvirus
Bright yellow mottle in most varieties, sometimes necrosis of tubers.
Potato Bouquet Disease
Caused by Tobacco Ring Spot Nepovirus.
Potato Calico
Caused by strain of AlfalfaMosaic Alfamovirus.
Potato Crinkle
Mild Mosaic. Due to Potato Virus X plus A. Leaf mottling and crinkling are
often inconspicuous, but plants die prematurely. Plant healthy tubers; isolate
seed plots. Varieties Katahdin, Chippewa, Houma and Sebago are resistant.
Potato Green Dwarf
Caused by a strain of Beet Curly Top Hybrigeminivirus. Terminal growth is
dwarfed and deformed; leaets cupped upward.
Potato Leaf Roll Luteovirus
Important wherever potatoes are grown. Symptoms show about a month after
plants appear above ground. Leaves are thick, leathery, rolled, with excessive
starch; sometimes with a reddish or purple discoloration on the underside.
Plants are dwarfed; tubers are few, crisp, with net necrosis brown strands of
dead tissue in some varieties; sprouts are spindling; yield may be reduced
one-half. Transmission is by peach and other aphids.
Use certied seed potatoes. These come from a foundation stock obtained by
indexing. Seedpieces or tubers are planted consecutively in a row, and if any
show virus symptoms, the whole unit is destroyed.
Potato Leaf Rolling Mosaic =Potato M Carlavirus
Leaves are mottled, accid, with some upward rolling but without distinct
rolling, rigidity of leaf roll. Transmission by peach, potato and geranium
aphids.
Potato Mottle
Caused by potato X potexvirus.
Potato Rugose Mosaic
Caused by potato Y potyvirus, often with PVX. Leaves are crinkled, mottled;
lower leaves with black veins; plants are stunted, die prematurely. Control by
careful roguing.
Potato S Carlavirus
Generally symptomless in most potato cultivars.
Potato Spindle Tuber Viroid
General on all tested varieties of potatoes. Plants are more erect than normal
but spindly, lacking vigor. Stems are stiff, leaves small, dark green; tubers are
elongated, pointed at the end, the eyes staring. Symptoms are accentuated
by high soil moistures. Transmission is by contaminated knives in cutting,
by contact between freshly cut seed pieces. Control by using certied seed.
Potato Vein Banding =Potato Y Potyvirus
On potato and many other hosts, transmitted mechanically and by many
aphids. On some varieties there is leaf drop and necrotic streak or chlorotic
mottling; on others there is no sign of disease.
Potato Virus A
Present in nearly symptomless form in some varieties, causing crinkle with
virus X.
Potato X Potexvirus
Almost universally present in commercial potato stocks. Cause of latent
mosaic.
Potato Witches Broom
Apical leaves are slightly rolled, upright, light green with reddish or yellow-
ing margins. There is proliferation of axial buds with tendency to bloom
and set fruit; there are aerial tubers and numerous small subterranean
tubers. Such tubers put out spindle shoots without a rest period and pro-
duce dwarfed, very bushy plants with small, round, or heart-shaped leaves.
Use certied seed potatoes.
Potato Yellow Dwarf Nucleorhabdovirus
Formerly causing heavy losses in Northeast but now mostly controlled by
seed certication. Potato leaves are rolled and yellowed; the plant is dwarfed
with split stems showing rusty ecks. Transmission is by clover leafhoppers;
overwintering hosts are ground cherry, oxeye daisy, vinca and other plants.
Potato Yellow Spot
Reported from Maine, mostly on Katahdin variety. Spots are small, circular,
bright yellow, chiey on lower leaves.
Primrose Mosaic
Plants are chlorotic, stunted, rugose, with upward, sometimes downward,
cupping of leaves. Petioles and peduncles are shortened; owers are white-
streaked; leaves are coarsely mottled yellow-green, with green islands; tips
of leaves are narrowed. No insect vector is known.
Privet Ring Spot
Reported on privet in Texas. Leaves are smaller, lighter green, drop early.
Prune (Standard) Constricting Mosaic
Spots are concentrated in a band across tip of the leaf; this area is killed, and
all tissue except the midvein drops out.
Prune Diamond Canker
Symptoms expressed only on French prune diamond or oval excrescences
on secondary branches, often excess sprouts from body of tree.
Prune Dwarf Ilarvirus
On prune, plum, cherry and peach. Leaves are small, narrow, rugose, dis-
torted, glazed. Internodes are short, but some branches escape and appear
normal. Blossoms are numerous, but mature fruits few; pistils are aborted,
petals narrowed. Most injurious to Italian prune, symptomless in Bradshaw
and damson plums. Transmission by grafting and budding.
Prunus Necrotic Ringspot Ilarvirus
Very common in sweet cherry and plum orchards, worldwide. Its natural
hosts are species from Prunus, Rosa, Humulus and Cucumis. Earlier virus
was described as Plum Line Pattern, Peach Ringspot, Prunus Ringspot, Red
Currant Necrotic Ringspot, Rose Chlorotic Mottle, Rose Line Pattern, Rose
Vein Banding and Sour Cherry Necrotic Ringspot. Virus is transmitted by
grafting, by seeds and by pollen to seeds and to the pollinated plants.
Quail Pea Mosaic Comovirus
Caused mosaics on Streptostyles helvola as well as on soybean and common
bean plants.
Radish Mosaic Comovirus
Chlorotic spotting and mottling of foliage; plants not stunted.
Ranunculus Mottle Potyvirus
Caused foliar mottling and distortion on Ranunculus asiaticus plants.
Raspberry Alpha Leaf Curl Luteovirus
Common on red raspberry. Veins are retarded in growth, causing down-
ward curling and crinkling of leaves. Foliage is dark green, but bronzed in
late summer with glistening surface. Berries are small, poor; diseased canes
are readily winter-killed. Transmission is by small raspberry aphid (Aphis
rubiphila). Cuthbert variety is most susceptible. Rogue diseased plants.
Raspberry Beta Leaf Curl Luteovirus
Infecting blackcaps, especially Cumberland, but also causing severe curling
on Cuthbert and hybrid purple Columbian.
Raspberry Decline
On red raspberry.
Raspberry (Red) Mosaic
Green Mottle; Mild Mosaic; Yellows. Widespread on red and black raspber-
ries, dewberry and blackberry. Symptoms vary greatly, but usually mottled
areas are darker green than rest of leaf tissue; there may be blistering and
curling downward. On blackcaps, tips are stunted, fruiting laterals shortened,
fruit seedy or with poor avor. Foliation of diseased plants is delayed. Trans-
mission by aphids.
Raspberry (Black) Necrosis
On red and black raspberries and perhaps related to red raspberry mosaic.
Leaves are curled down, have necrotic spots.
Raspberry Ringspot Nepovirus
Virus known also as Red Currant Ringspot is transmitted by nematode,
Longidorus spp. Raspberry cultivars differ in susceptibility.
Raspberry Streak
Eastern Blue Streak, Rosette. On black raspberry. Plants are stunted, smaller
in successive seasons, leaves usually curled, close together on canes, dark
green, often twisted upside down. New canes have bluish dots or streaks near
the base and sometimes on branches of fruiting spurs. Fruit is small, poor;
plants are short-lived. Symptoms are less severe in the mild streak strain.
Roguing aids in control.
Raspberry Yellow Mosaic =Rubus Yellow Net Bednavirus
Black raspberries are severely dwarfed; fruiting laterals are more upright
than normal; foliage is yellow; leaets are long and narrow. Symptoms show
at high temperatures, while those of red raspberry mosaic are masked. Plants
are weakened, die in 2 or 3 years. Transmitted by the raspberry aphid.
Red Clover Vein Mosaic Carlavirus
Caused vein yellowing and mosaic on infected red clover plants.
Rhododendron Necrotic Ringspot Potexvirus
Transmitted by grafting; infected Rhododendron spp. showed necrotic rings
on leaves.
Rhubarb Chlorotic Ring Spot
Chlorotic spots and rings, necrotic stippling and rings on leaves of rhubarb,
reported from Oregon.
Rhubarb Ring Spot
Caused by Turnip Mosaic Potyvirus.
Rhynchosia Golden Mosaic Begomovirus
Reported on soybean in Mexico.
Ribgrass Mosaic Tobamovirus
It is ribgrass strain of Tobacco Mosaic Tobamovirus.
Robinia Brooming
Locust Witches Broom.

Rose Mosaic
Infectious Chlorosis. Common on garden roses on the Pacic Coast, some-
times on greenhouse roses in the East and on garden roses originating in
the West. Chlorotic areas feather away from midribs of leaets, often with
local distortion, sometimes with ring, oakleaf and watermark patterns. Plants
are dwarfed, with buds often bleached, imperfect, on short stems. The virus
is carried in understock and infects tops after budding or grafting; no insect
vector is known. More than one virus is probably involved. Much rose mosa-
ic seems to be due to the Prunus Necrotic Ring Spot and Apple Mosaic
Ilarviruses. Rootstocks can be heat-treated to provide a virus-free source.
Rose Rosette
On species roses, Wyoming, California. Leaets and ower parts are mis-
shapen, stems dwarfed, with precocious growth of lateral buds, indenite
chlorotic pattern in leaves, increase in thorniness of stems. The general effect
resembles 2, 4-D injury. Graft and mite transmissible, but the disease devel-
ops slowly.
Rose Streak
On rose in eastern United States. Leaves have brownish or reddish ring and
vein-banding patterns; stems have ring patterns and sometimes necrotic areas
near inserted buds, causing girdling, wilting of foliage. Transmission is by
grafting.
Rose Yellow Mosaic
Chlorotic areas are brighter and lighter yellow than in typical rose mosaic;
there is less puckering of leaves.
Saguaro Cactus Carmovirus
Symptomless on Saguaro cactus plants. Virus was found in Arizona but there
is no evidence of spread.
Schefera Ringspot Badnavirus
Bacciliform particles were found in Brassica actinophylla.
Shamrock Chlorotic Ringspot Potyvirus
Caused chlorotic Ringspot on leaves and decline of Oxalis regnellii plants.
Smithiana Potex Virus
Infected plants are usually symptomless.
Solanum Yellows Luteovirus
Occurs on potato plants.
Sonchus Yellow Net Nucleorhabdovirus
On lettuce.
Sorghum Chlorotic Spot Furovirus
Found only in Kansas; caused chlorotic spot on infected plants.
Sorghum Mosaic Potyvirus
Virus is one of the potyvirus complex infecting tropical grasses.
Sorghum Stunt Mosaic Nucleorhabdovirus
Transmitted by leafhopper, Graminella sonora, only in temperatures below
38C.
Sowbane Mosaic Sobemovirus
Occurred in Europe, South and Central American region; found also in USA.
Sowthistle Yellow Vein Nucleorhabdovirus
Susceptible hosts are Sonchus oleraceus and Lactuca sativa.
Soybean Dwarf Luteovirus
There are two different strains of this virus; dwarng strain and leaf yellow-
ing strain. Virions occurred in cytoplasm and cell vacuoles of phloem. Virus
transmitted only by aphid vectors in persistent manner.
Soybean Mosaic Potyvirus
Widespread on soybean. Leaves are distorted, narrow, with margins turning
down, some with rufing along main veins; plants are often stunted, pods
misshapen with fewer seeds. Transmission by peach, pea and other aphids
and in seed. Control by roguing.
Soybean Yellow Mosaic
Soybean Yellow Mosaic is caused by the Bean Yellow Mosaic Virus.
Younger leaves show chlorotic mottling, followed by necrotic spots. Soy-
bean bud blight, due to Tobacco Ring Spot Virus, is serious in the Midwest,
causing losses up to 100%. Tip buds turn brown, dry brittle; plant is dwarfed,
produces no seed.
Sparaxis Mosaic
Strong leaf mottling and crinkling.
Spinach Blight
Caused by Cucumber Mosaic Cucumovirus.
Spinach Latent Ilarvirus
Transmitted by means not involving a vector. Symptoms disappeared soon
after infection.
Spinach Yellow Dwarf
In California, conned to spinach, with vein clearing, curvature of midrib;
young leaves with mottling, puckering, curling, blisters; old leaves with
yellow blotches becoming necrotic. All varieties are equally susceptible.
Mechanical transmission and by aphids.
Spring Beauty Latent Bromovirus
Symptomless; best serological test for identication is double diffusion test.
Squash Leaf Curl Bigeminivirus
It is probably the same as Muskmelon Necrotic Mosaic Virus causing severe
stunting and leaf curl on plants from family Cucurbitaceae.
Squash Mosaic Comovirus
On squash and muskmelon, mostly in California. Foliage is severely mottled
and malformed with dark green blisters. Transmission by banded, western
striped and 12-spotted cucumber beetles, but not by pollen.
Squash (Southern) Mosaic
On squash in Florida, infecting also cucumber and watermelon.
Stock Mosaic
On stock, reported from California. Denite mottling with dark green islands
conspicuous against light green areas. Plants are stunted; seed pods small;
owers broken with petals undersized. Cut-ower elds are often a total loss,
but there are resistant varieties.
Strawberry Crinkle Cytorhabdovirus
Chlorotic and necrotic spotting with crinkled leaves and vein clearing. Trans-
mission by strawberry aphid.
Strawberry Latent Ringspot Nepovirus
Infects strawberry and rose; latent and seedborne in parsley.
Strawberry Latent Virus Rhabdovirus
Causing no distinct symptoms but intensifying those caused by other viruses.
Strawberry Leaf Curl
Caused by Strawberry Veinbanding Caulimovirus plus Strawberry Latent
Rhabdovirus.
Strawberry Leaf Roll
Leaets are rolled down, pale green, small, on spindly petioles.
Strawberry Mild Crinkle
Caused by Strawberry Vein Chlorosis Virus with or without Strawberry Mot-
tle Virus.
Strawberry Mild Yellow Edge Chlorosis Potexvirus
Slight chlorosis of leaf margin.
Strawberry Mild Yellow Edge Luteovirus
Transmitted by aphids in persistent manner. The relation to Soybean Dwarf
Luteovirus is so close that they are probably the same species.
Strawberry Mottle
Chlorotic spotting, leaf distortion.
Strawberry Multiplier Disease
Resembling witches broom and stunt; transmitted by leaf grafting.
Strawberry Necrotic Shock =Tobacco Streak Ilarvirus
Blackish spots on leaves and petioles; whole crown may be killed, but plants
recover, and virus becomes latent.
Strawberry Pallidosis =Cucurbit Yellows Crinovirus
and Beet Pseudo-Yellows Closterovirus
On strawberry.
Strawberry Pallidosis Virus
Caused no disease symptoms and is transmitted only by grafting; dsRNAwas
found in infected cells.
Strawberry Phyllody Strawb PhF Phytoplasma
On strawberry.
Strawberry Pseudo Mild Yellow Edge Carlavirus
Symptomless on infected plants; transmitted by aphids in semi-persistent
manner. Virus related to Carnation Latent Carlavirus.
Strawberry Severe Crinkle
Due to Strawberry Mottle Virus plus Strawberry Crinkle Cytorhabdovirus.
Strawberry Stunt
In the Pacic Northwest. Plants are erect but short; leaves at rst folded, later
open, dull with a papery rattle; leaets cupped or with margins turned down;
midveins tortuous; petioles short; fruits small, hard, seedy. Transmission by
the strawberry aphid.
Strawberry Veinbanding Caulimovirus
Diffuse banding along veins; leaets with epinasty, mild crinkling, wavy
margins. Transmission by several aphids, grafting, dodder.
Strawberry Witches Broom
Leaves are numerous, light in color with spindly petioles; margins of leaets
are bent down; runners are shortened, plants dwarfed; ower stalks spindly
and unfruitful. Transmission by the strawberry aphid.
Strawberry Yellow Edge
Central leaves dwarfed, with yellow edges.
Strawberry Yellows
A complex caused by mild yellow edge, crinkle, and mottle viruses. June
yellows is a genetic leaf variegation, not due to a virus.
For control of strawberry viruses buy certied plants. Nurseries on the Mary-
land eastern shore provide 37 varieties virus-free from a foundation stock of
indexed plants.
Streptanthera Mosaic
Mottled foliage. Caused by Bean Yellow Mosaic Virus.
Subterranean Clover Red Leaf Luteovirus
Caused mild yellowing, stunting and reddening of many plants from Legu-
minosae family.
Sugarcane Bacilliform Badnavirus
Occurs in Florida and Hawaii; serologically virus is related only to Banana
Streak Badnavirus.
Sunower Mosaic Potyvirus
Known also as Helianthus Mosaic Virus; caused mild systemic mosaic and
mottling, sometimes necrosis on leaves and stems.
Sweet Clover Sweet Latent Nucleorhabdovirus
Virus often together with Bean Common Mosaic Potyvirus caused leaf
chlorosis.
Sweet Potato Caulimovirus
There are no conspicuous symptoms on Ipomoea batata. There is no evi-
dence of it spreading in the United States.
Sweet Potato Feathery Mottle Potyvirus
First symptom is a yellowing along veins or small diffuse yellow spots. Some
leaves are abnormally dark green with feathery yellow areas along veins.
Leaves may be slightly rugose and dwarfed. Transmitted by aphids, white-
ies, and sprouts. Caused feathering, clearing and chlorotic spots on Ipo-
moea batata but symptoms usually disappeared soon after infection.
Sweet Potato Internal Cork
=Sweetpotato Feathery Mottle Potyvirus
First recognized in South Carolina in 1944, now in most sweetpotato areas,
most prevalent in Georgia and the Carolinas. Dark brown to blackish corky
spots in esh of roots, which appear normal outside. Some are present when
sweetpotatoes are dug, but cork spots increase in number and size during
storage, especially at temperatures higher than the recommended 55 to 60F.
Foliage symptoms are vein feathering and mottling followed by reddish to
purple blotching sometimes in ring form. Quality of Porto Rico variety is
severely affected, but not yield. Transmission is by peach and potato aphids
and by grafting, with morning-glories used as index plants. Control insects to
reduce disease; there is little spread to new plantings 100 yards or more from
diseased elds. Cure immediately after digging at 85F with 90% humidity;
then store at 55F, except seed stocks, which should be kept at 70F so that
lots with internal cork can be selected and discarded.
Sweet Potato Mosaic
Transmitted by eshy-core and sprout grafts and by sweetpotato whitey.
Sweet Potato Russet Crack Strain of Sweetpotato
Feathery Mottle Potyvirus
Dark lesions and ne cracks in skin of eshy roots.
Teasel Mosaic Potyvirus
On Fullers teasel and scabiosa, with vein-clearing, asymmetry, strong mosa-
ic pattern, malformation, death of plant. Transmission by peach and rose
aphids.
Tigridia Mosaic
Pale to yellow-green irregular streaks and blotches in leaves and ower
bracts, occasionally dark streaks in owers. Transmission by lily and melon
aphids.
Tobacco Broad Ring Spot
In tobacco, Wisconsin, experimentally to other plants. Chlorotic or necrotic
rings, sometimes concentric; young leaves puckered at rst.
Tobacco Etch Potyvirus
Mild and severe strains widespread on tobacco, tomato, pepper, petunia,
potato and other plants. Symptoms are vein clearing with ne necrotic etch-
ing, usually toward base of leaves. Plants are stunted with smaller, mottled
leaves. Transmission is by peach, lily, bean and other aphids.
Tobacco Mild Green Mosaic Tobamovirus
Transmitted on the surface of the affected seeds. Infectious virus particles
were found in water used for irrigation in greenhouses.
Tobacco Mosaic Tobamovirus
Tomato Mosaic; Pepper Mosaic. General in gardens, elds, greenhouses on
tobacco, tomato, pepper, eggplant, petunia, Moraine ash, Achimenes, Aeschy-
nanthus, Chirita, Codononthe, Episcia, gloxiana, Kohleria, Nematanthus,
Streptocarpus, Smithantha, Rhoeo, and nearly all solanaceous plants. Toma-
to foliage has a light and dark green mottling, accompanied by some curling
and malformation of leaets, often with a fernleaf effect. A yellow strain
of the virus causes striking yellow mottling of leaves, sometimes stems and
fruits. Yield is greatly reduced. In pepper, yellowish chlorotic lesions are fol-
lowed by systemic chlorosis. Spinach has some mottling, stunting, necrosis.
Eggplant is often killed.
Transmission is by mechanical means by handling, on tools, through soil,
by grafting, possibly but not probably by seed. The virus can be transmitted
by feeding of grasshoppers, but apparently there is little spread by the usual
aphid vectors. This is the most resistant and highly infectious of all viruses. It
withstands heat, even alcohol and various germicides, and retains infectivity
in a dried state for many years. The most common source of inoculum is
smoking tobacco. Gardeners contaminate their hands by smoking and then
infect plants as they transplant, tie, prune, etc., the virus entering through
scratches or broken hairs. The rst symptoms appear in 8 to 10 days. In
greenhouses, even doorknobs, faucets, and ats can be contaminated after
handling virus-infected plants and remain a source of infection.
There are many strains of the virus, causing cowpea mosaic, tomato aucuba
mosaic, tomato enation mosaic, tomato streak, orchid aucuba, etc.
Control. Remove and burn any suspicious plants in the seedbed along with
neighboring plants. Destroy weeds, especially ground cherry and other
solanaceous species. Never smoke while working with plants, and always
wash hands thoroughly with soap after handling tobacco in any form or
touching diseased plants, before handling young seedlings or healthy plants.
Tobacco Necrosis Necrovirus
On tobacco, tomato, aster, geranium and bean, conned to roots, or systemic
without symptoms, or systemic with symptoms. In Holland, the virus causes
a severe crippling of tulips called Augusta disease, often preventing ower-
ing, resulting in death.
Tobacco Rattle Tobravirus
On Romaine lettuce and transmitted by Paratrichodorus christiei.
Tobacco Ring Spot Nepovirus
General on tobacco, petunia, potato, cucumber, celery, Moraine ash and gera-
nium, causing pimple disease of watermelon, bud blight of soy bean, in glad-
iolus, iris, Astilbe, and Easter lily without symptoms. Causing large chlorot-
ic areas on spinach leaves; faint zigzag lines on beet; pin-point necrotic
spots with yellow haloes on cucurbits and fruits rst pitted, then with ele-
vated pimples; eggplant yellows, bouquet disease of potatoes, with stems
curved, shortened, sometimes with black lesions on underside of veins. Petu-
nia seedlings are stunted, rst leaves are mottled, and seed pods have few
seeds. Mint is stunted.
Transmission is through seed of petunia (but not of tobacco), by nematodes
and in some crops by grasshoppers.
Tobacco Streak Ilarvirus
On tobacco, soybean, sweet clover, tomato, common yellow mustard, wild
radish, milk thistle, and experimentally a wide range of hosts. Irregular spots,
lines, and rings. Distributed worldwide. Virus is known as Asparagus Stunt
Virus, Datura Quercina Virus and Strawberry Necrotic Shock Virus. It is
transmitted by thrips, by sap and by seeds. Virus is also transmitted by pollen
to the pollinated plants.
Tobacco Vein Mottling Potyvirus
Transmitted by aphids in non-persistent manner. Its natural host plants are
Rumex sp., Nicotiana tabacum and Solanum carolinense. Virus caused
chlorotic vein banding symptoms on infected plants.
Tobacco Yellow Net Luteovirus
Spreads by aphids Myzus persicae in California. Virus differs in host range
from Beet Yellow Net Luteovirus.
Tomato Aspermy Cucumovirus
Chrysanthemum Aspermy. On tomato the growing point of the main stem
is inhibited, axillary shoots giving the plants a bushy appearance; fruit pro-
duction is curtailed; there may be failure to set seed. The disease was intro-
duced into North America on European and Asiatic varieties of chrysanthe-
mums, which have mottled leaves. Transmission is by foxglove, green peach,
and green and black chrysanthemum aphids. Perennial chrysanthemums near
tomato elds are a source of infection.
Tomato Big Bud
Leaves curl and hang down; stems are shortened and calyxes greatly
enlarged.
Tomato Black Ring Nepovirus
Transmitted by nematodes, seeds and pollen. Virus caused necrotic ringspots,
systemic chlorotic ringspots, mottling, stunting and leaf malformation of the
following plant genera: Allium, Apium, Beta, Fraxinus, Lactuca, Lycopersi-
con, Narcissus, Phaseolus, Robinia, Rubus, Solanum, Tulipa and Vitis.
Tomato Bushy Stunt Tombusvirus
Strains of this virus are known as Carnation Italian Ringspot Tombusvirus,
Pelargonium Leaf Curl Tombusvirus and Petunia Asteroid Mosaic Tombus-
virus. Virus transmitted by means not involving vectors. Virions were found
in all parts of the host plant: in cytoplasm, in nuclei, in nucleoli, in mitochon-
dria and in cell vacuoles. It forms crystals in the cytoplasm.
Tomato Chino La Paz Begomovirus
Found on tomato in Baja California, Mexico.
Tomato Enation Mosaic
Caused by a strain of Tobacco Mosaic Tobamovirus.
Tomato Fernleaf
Shoestring. Caused by Cucumber Mosaic Cucumovirus sometimes with
Tobacco Mosaic tobamovirus.
Tomato Infectious Chlorosis Closterovirus
Transmitted by white y Trialeurodes vaporariorum; caused interveinal
yellowing and necrosis in infected tomato plants.
Tomato Mosaic Tobamovirus
Circular, water-soaked necrotic spots on leaves; black streaks on veins; con-
centric sunken rings on fruit.
Tomato Pseudo Curly Top Hybrigeminivirus
Spreads among tomato plants in Florida. Virus is transmitted by treehopper
Micrutalis malleifer. Virion genome consists of circular ssDNA.
Tomato Ring Spot Nepovirus
Curling and extensive necrosis of shoot terminals; brown rings and streaks
on leaets, stems, fruits, more pronounced at high temperature. Causes leaf
streaks in iris, crumbly fruit of red raspberry and chlorotic spot of geranium.
May infect chicory, healall, black medic, moth mullein, impatiens, apple,
common cinquefoil and dandelion. Transmission is by dagger nematodes.
Plants that may serve as reservoir plants for Tomato Ringspot Virus are:
chickweed, henbit, dandelion, woodsorrel, plantain, strawberry, sorrel and
red clover.
Tomato Spotted Wilt Tospovirus
More serious on the Pacic Coast, but also occurring in Florida, Texas and
some central and eastern states in greenhouses and sometimes outdoors on
plants such as ragwort, purslane, nightshade, desert-rose, and puncture vine.
Spotted wilt is common on tomato, potato, tobacco, lettuce, pea, pepper,
celery and other vegetables. Ornamentals include amaryllis, aster, begonia,
blackberry-lily, calendula, calla, chrysanthemum, dahlia, delphinium, fuch-
sia, gaillardia, gloxinia, nasturtium, geranium, primrose, petunia, Rieger
begonia, hydrangea, stephanotis, salvia, stock, verbena and zinnia.
In tomato there are bronze, ringlike secondary lesions; plants are stunted
with some necrosis; there may be a yellowish mosaic with leaf distortion.
Fruits are often marked with concentric rings of pale red, yellow, or white.
Potatoes have zonate necrotic spots on upper leaves, streaks on stems, which
collapse at the top; plants are stunted, with small yield. Lettuce is yellowed,
with retarded growth, brown blemishes on central leaves; affected spots are
like parchment but with brown margins. Peas have purplish necrotic spots
on stems and leaves following mottling, and circular spots and wavy lines
on pods. Spots on outer stalks of celery are rst yellow, then necrotic with
pockets of dead tissue inside petioles; plants are stunted and worthless.
China asters have dead tan areas in leaves, brown surface blotches on stems.
Calla lillies have whitish, then brown spots and streaks. On sweet pea, red-
dish brown to purple streaks may run full length of the stem. Circular to
oval leaf spots with diffuse margins are followed by yellowing and death of
leaves and stems. Blossoms sometimes develop a circular pattern in the pig-
ment. Delphiniums may have numerous distinct double rings. The viruses
causing oak leaf and ring spot in dahlia are probably strains of the spotted
wilt virus.
Transmission is by onion and ower thrips. Only the larvae can become vir-
uliferous by feeding on infected plants, but then there is an incubation period
of 5 to 9 days, during which the insect becomes adult, before the virus can
be transmitted to healthy plants.
Tomato Streak; Double Streak
Caused by Tobacco Mosaic Tobamovirus plus Potato X Potexvirus. Leaves
are mottled green with numerous small, grayish brown papery spots, may
wither and dry. Later growth is mottled green and yellow with small choco-
late brown spots and dark brown streaks on stems; fruit has brown greasy
spots. The disease is more important in greenhouses; workers should refrain
from handling tobacco or potatoes while working with tomatoes.
Tomato Top Necrosis Neopvirus
Occurs in Indiana, Missouri and Illinois. Virus is very rare on tomato plan-
tations. It is serologically related to many nepoviruses.
Tomato Western Yellow Blight
Beet Curly Top Hybrigeminivirus.

Tomato Yellow Leaf Curl Begomovirus
On tomato.
Tomato Yellow Net =Tobacco Yellow Net Luteovirus
Pronounced yellow necrosis of veins and veinlets. Transmission by the peach
aphid.
Tomato Yellow Top =Strain of Potato Leafroll Luteovirus
Leaets small, curled, yellow; or purplish in cool weather.
Tradescantia Zebrina Potyvirus
Transmitted by aphids to Tradescantia, Zebrina, Commelina and Rhoeo sp.
plants on which it caused mosaics, stuntings and malformation of leaves.
Tritonia Mosaic
Mottling at base of young leaves.
Tulare Apple Mosaic Ilarvirus
Caused leaf chlorosis on Corylus and Malus sp., it is transmitted only by
mechanical inoculation. Virus is related to Citrus Leaf Rugose Ilarvirus.
Tulip Breaking Potyvirus
Due to Lily Latent Mosaic virus often present with Tulip Color-adding virus.
Broken tulips appear wherever hybrids are grown. There is little or no obvi-
ous effect on foliage and little interference with growth, but there are marked
color patterns on the owers; differences in named broken varieties possibly
due to the proportion of color-breaking and color-adding viruses present.
Most pure white owers do not change; some turn pink or red. Pink and
bright red owers have strong color changes; very dark tulips turn even dark-
er. There may be dark stripes due to pigment intensication. Transmission is
by aphids, and roguing should take place early, before insects are active.
Broken varieties should not be grown near those with solid colors.
Turnip Mosaic Potyvirus
Cabbage Black Ring Spot; Watercress Mottle; Nasturtium Mosaic. On
turnip, rutabaga, rape, mustard, cabbage, collard, horse-radish, watercress,
garden balsam, nasturtium, stock, ladys slipper, impatien, safower and
sweet rocket. Turnip shows a systemic chlorotic mottling with crinkling, leaf
distortion, stunting of plants. Cabbage has numerous small, black necrotic
rings or spots; cauliower and broccoli have a diffuse systematic mot-
tling. Horse-radish has blotchy mottling, necrotic rings, ecks, and streaks
on petioles and leaf veins. Variegated owers appear on nasturtium stock,
wallower and sweet rocket; the last may be severely crippled or killed.
Transmission is by peach and cabbage aphids. Protect seedlings by spraying
or screening seedbeds.
Walnut Brooming Disease
Bunch Disease. Presumably virus.
Watermelon Curly Mottle Bigeminivirus
Occurs in Arizona. Infected Citrullus lanatus and Cucumis melo var. can-
talupensis plants showed leaf necrosis, vein banding, curling and stunting
symptoms.
Watermelon Mosaic Potyvirus
In Florida and probably other states. Symptoms include mild interveinal
chlorosis, stunt, distortion, mottle, consisting of green bands along veins or
raised green blisters. Leaf apices often form long narrow, sometimes twisted
projections, shoestrings. A yellow strain of the virus causes more yellow
spotting with less shoestring effect.
Watermelon Silver Mottle Tospovirus
On watermelon.
Wheat Streak Mosaic Rymovirus
On wild rice (Zizania); foliar streak symptoms with chlorotic areas becoming
necrotic with eventual leaf death.
White Clover Mosaic Potexvirus
Known also as Clover Mosaic Virus and Pea Wilt Virus, caused systemic
mosaics and chlorotic mottlings on clover plants. Virus is transmitted by
contact between plants and by seeds.
White Line Mosaic
A mosaic disease of sweet and dent corn with short chlorotic lines along leaf
veinal tissue.
Wild Cucumber Mosaic Tymovirus
Occurs in California and Oregon. Virus caused mild leaf chlorosis on natural
hosts: Marah sp. and Echinocistis sp.
Wineberry Latent Virus
Known as Loganberry Calico Virus induced calico disease-like symptoms on
blackberry. Virus is transmitted by mechanical inoculation and by grafting
but on many Rubus sp. remains symptomless.
Wisteria Mosaic
Diffuse yellowish blotches with scattered green islands; mature leaets twist-
ed.
Wisteria Vein Mosaic Potyvirus
On wisteria.
Yellow Mottle Begomovirus
On okra.
Zucchini Yellow Mosaic Potyvirus
On cucurbits, melon, squash, pumpkin and watermelon.
Zygocactus Montana X Potexvirus
Serologically related to many potexviruses, caused reddening of pads of
infected Zygocactus truncatus plants. It was found in the USA but there is
not evidence of spread.
WHITE RUSTS
White rusts are all members of a single genus, Albugo, in the Oomycetes and
are apparently obligate parasites like the true rusts. They form a white blister
just underneath the epidermis.
Albugo (Cystopus)
Sporangia are borne in chains at apex of a short, clavate, usually unbranched sporangio-
phore, forming a limited sorus beneath the host epidermis and exposed by its rupture.
The mycelium is intercellular except for small, knoblike haustoria. The sporangia dry to
a white powder and are disseminated by wind, germinating by swarmspores. Fertiliza-
tion of a globose oogonium and a clavate antheridium produces a single oospore, also
germinating by swarmspores.
Albugo bliti. White Rust or White Blister on beet, amaranth, globe ama-
ranth, seabeach amaranth, and smooth pigweed. Blisterlike white pustules
formed in leaves change to reddish brown when mature. Flowers and stems
are dwarfed, distorted. The fungus winters in seed coats. Destroy infected
plants and debris at end of season. Change location of plantings.
Albugo candida. White Rust of crucifers on arugula, cabbage, chinese cab-
bage, radish, horse-radish, turnip, watercress, garden cress, peppergrass, sal-
sify, mustard, arabis, sweet alyssum, boerhavia, draba, hesperis, candytuft,
stock, wallower and western wallower.
Blisters appear on any part of the plant except root. They vary in size and
shape and are often conuent in extended patches. There seem to be two
types of infection: general or systemic, resulting in stunting of entire plant
and formation of pustules on all parts; or local, with direct invasion of single
leaves, stems, or owers. Upper surface of leaves often has yellow areas with
white pustules on the underside. The latter are powdery when mature, and the
epidermis is ruptured to free chains of sporangia that are carried by wind to
moist surfaces. They germinate by 6 to 18 zoospores, swarmspores, which
settle down, produce germ tubes, and enter plants through stomata.
Stems have localized or extended swellings, sometimes sharp bends, pro-
liferation from lateral buds giving a bushy growth. Various ower parts are
deformed with pronounced distortion of ower pedicels. When these thick-
ened parts die, oospores are formed to survive the winter in crop refuse. The
disease ourishes in cool, wet weather; the spores germinate better when
slightly chilled.
Control. Remove infected parts of ornamentals as noticed. Clean up all veg-
etable refuse at end of season and all cruciferous weeds nearby. Spraying is
impractical.
Albugo ipomoeae-panduratae. White Rust, general on sweetpotato, also
on morning-glory, moonower, Jacquemontia and quamoclit. The disease is
usually late on sweetpotato, after vines have made their growth, but it is very
conspicuous with irregular yellow areas on upper surfaces and white cheesy
pustules on lower surface. Oospores wintering in host tissue are liberated by
decay in spring. There are no control measures.
Albugo occidentalis. White Rust of spinach. After a report from Virginia in
1910, the disease went unrecorded until 1937, when it appeared in epidemic
form in Texas; it has since been serious in Oklahoma and Arkansas and has
attacked all commercial varieties tested at the University of Wisconsin. The
white blisters are small, usually on underside of leaves, sometimes on upper.
Infected leaves become chlorotic, then brown; the entire crop may be lost.
Albugo platensis. White Rust, on trailing four oclock, common four
oclock, and boerhavia.
Albugo portulacae. White Rust of portulaca. Swollen and deformed
branches bear white pustules. Shoots tend to become more erect and spin-
dling.
Albugo tragopogonis. White Rust of salsify, also on African daisy, anten-
naria, artemisia, centaurea, feverfew, matricaria, senecio and sunower.
Light yellow areas appear on leaves. The epidermis, forced into domelike
swellings, bursts to show chalky sori of spores. Foliage may die; plants are
dwarfed. There is no control.
WILT DISEASES
To wilt means to lose freshness or to become accid. Wilting in plants may
be temporary, due to too rapid transpiration; or it may be permanent, due to
continued loss of water beyond the recovery point. Disease organisms, by
reducing or inhibiting water conduction, may cause permanent wilting.
Because wilt diseases are systemic, and tied up with the entire vascular sys-
tem of a plant, they are usually more important, and harder to control, than
localized spots or cankers. In many cases the fungus enters the plant from
the soil through wounds or root hairs and cannot be controlled by protec-
tive spraying. Often, although the fungus is present only near the base of
a plant, the rst symptom is a agging or wilting or yellowing of a branch
near the top. Many species of Fusarium are responsible for important wilts
and yellows. Verticillium is a common cause of wilt in maples, other trees,
and shrubs, but most important among the wilt pathogens are two species of
Ceratocystis, one causing oak wilt, the other Dutch elm disease.
Acremonium (Cephalosporium)
Leaf Spots.
Acremonium diospyri (formerly Cephalosporium diospyri). Persimmon
Wilt, a lethal disease of common persimmon. Wilt appears in scattered local-
ities from North Carolina to Florida and west to Oklahoma and Texas, but
most infection is in north central Florida and central Tennessee. Spread is
rapid and death quick. First notice of the disease was in Tennessee in 1933.
By 1938 only 5% of the persimmons in the infected stand were alive. Top-
most branches wilt suddenly, then the rest of the tree, with defoliation and
death. The fungus fruits in salmon-colored spore masses in cracks in dead
bark of dying trees or under bark of dead rings. Fine, blackish streaks are
present in ve or six outer rings of trunk, branches, and roots. No control is
known.
Cephalosporium diospyri (see Acremonium diospyri). Persimmon Wilt,
a lethal disease of common persimmon.
Cephalosporium sp. Sunower wilt.
Ophiostoma (Ceratocystis)
Ascomycetes, Ophiostomatales
Perithecia enlarged at base, with thin walls, and long slender neck, ascus wall evanes-
cent, ascospores hyaline. Conidial stage may be Chalara with endogeneous spores or
Graphium with external conidia or conidiophores united into a dark stalk (synnema).
Ceratocystis fagacearum (Chalara quercina, Endoconidiophora faga-
cearum). Oak Wilt, our most serious disease of oaks, now known in 20
states from Texas and Oklahoma east to Pennsylvania and South Carolina. It
has also been reported in Florida. Although apparently present in the Upper
Mississippi Valley for many years, the disease did not cause concern, and
the fungus was not described until 1943, since when it has become a major
threat to our forest economy and to trees in residential areas. All native oak
species are susceptible, also chinquapin, chestnut, lithocarpus (and apples in
experimental inoculation); but red oaks succumb most rapidly. Scouting for
the disease has been done largely by airplane, the discolored foliage being
visible up to a half mile.
First symptoms are a slight crinkling and paling of leaves, followed by pro-
gressive wilting, bronzing, and browning of leaf blades from margins toward
midribs and defoliation progressively downward and inward throughout the
tree. Red oaks almost never recover and may be killed within 4 to 8 weeks
after symptoms appear. White and burr oaks may persist for some years, with
affected branches dying in a staghead effect.
The rst internal symptoms are the formation of gums and tyloses in the
xylem. After wilting, mycelial mats are formed between the bark and wood,
and the bark cracks from the pressure exerted. Perithecia are formed in these
mats, which have a sour odor and attract insects. Nitulid beetles, fruit ies,
brentids, springtails, bark beetles, and possibly other insects get conidia and
ascospores on or in their bodies as they feed, and can inoculate other trees
through wounds. We know that ascospores remain viable several months on
insects and can be distributed through fecal pellets, but we do not yet know
how great a role they play in the spread of oak wilt. Birds have been suspect-
ed as carriers but are not yet indicted. Local spread is largely by root grafts,
WILT DISEASES 683
one tree infecting others within 50 feet and with grafts possible between red
and white oaks, not limited to the same species.
Control. In residential areas infected trees should be removed. In forests,
felling may wound other trees and spread the disease more than letting the
dead tree remain but treated so that it is not infective. Different states handle
the problem in different ways. In Pennsylvania, each infected tree is cut, with
all other oaks within 50 feet, and ammate crystals are placed on each stump.
In North Carolina stumps and felled trees are thoroughly sprayed. In West
Virginia the trees are left standing, but a deep girdle into the heartwood dries
out the tree so that mycelial mats and spores do not form.
Ceratocystis (Ceratostomella) ulmi (Graphium ulmi) (see Ophiostoma
ulmi and O. novo-ulmi). Dutch Elm Disease, on American, Sibirian, Slip-
pery and European elms in 31 states, Maine to North Carolina and west to
Oklahoma, and on cedar.
Ophiostoma ulmi and O. novo-ulmi (formerly Ceratocystis (Ceratostomel-
la) ulmi (Graphium ulmi)). Dutch Elm Disease, on American, Sibirian,
Slippery and European elms in 31 states, Maine to North Carolina and west
to Oklahoma, and on cedar. This fatal disease is not really of Dutch origin
but is so named because it was rst investigated in Holland. It was noticed in
Europe about 1918, rst in France, then in Belgium and Holland. It spread
throughout central and southern Europe, then into England and Wales. In
many places it virtually exterminated the elms, including those on the famous
avenues at Versailles. It is suspected that the fungus came to Europe from
Asia during World War I.
Dutch elm disease was discovered in Ohio in 1930 and in New Jersey in
1933. It has spread north through New England and has become very serious
in the Midwest. In 1948, the disease was found in Denver, Colorado, and in
1976 in California. It is now fairly widespread in reports of its occurrence in
the United States. The spread of the fungus is linked with the presence of the
large and small European bark beetles, Scolytus scolytus and S. multistriatus.
Only the latter is established in this country, having arrived in Boston about
1919. Patient detective work established the fact that the fungus came here
in elm burl logs imported for furniture veneer. After one such infected elm
burl was found in Baltimore in 1934, months of scouting went on in the
vicinity of ports of entry, railroad distributing yards, and veneer plants. Such
backtracking showed the infected material had come in at four ports of entry
and had been carried by 16 railroads over 13,000 miles in 21 states. From
this source the disease got its start in at least 13 areas in 7 states.
Elm nursery stock is, of course, quarantined, and elm burls are embargoed;
but who who would have believed that dishes could have anything to do
with killing our elms? Dishes have to be crated, however, and several times
since 1933 English dishes crated with elm wood carrying bark beetles and
Ceratocystis have been intercepted. All American and European elms are
susceptible. Asiatic elms, Ulmus parvifolia and U. pumila, are resistant.
A seedling elm, named Christine Buisman for its Netherlands discoverer, is
highly resistant, though not immune, and is now available. Other promising
seedlings have been tested by the U.S. Department of Agriculture.
Symptoms are apparent from the latter part of May until late fall. The acute
form of the disease is characterized by sudden and severe wilting. First the
young leaves, then all leaves wilt and wither, sometimes so rapidly that they
dry, curl, and fall while still green, before they can turn the usual brown of
dead leaves. Sometimes terminal twigs are curled into a shepherds crook.
Chronic disease symptoms are gradual, often taking all summer for com-
plete defoliation. In many cases individual branches or ags appear, the
yellowed leaves conspicuous against the rest of the tree; but sometimes all
leaves gradually turn yellow. In another type of chronic disease, trees leaf
out late in spring, with sparse chlorotic foliage and a staghead appearance.
When an affected twig is cut across, the vessels or water-conducting tubes
show dark brown or black, being clogged with bladderlike tyloses and
brown gummy substances (see Fig. 3.62). The production of these sub-
stances is thought to be stimulated by a toxin secreted by the fungus and
carried in the sapstream. Symptoms are not dependent on the physical
presence of fungal hyphae in all parts of the tree. The fungus lives in the
sapwood, fruiting in cracks between wood and loosened bark and in bark
beetle galleries under the bark. This fruiting is of the anamorph state, spores
being produced in structures called coremia. These are black stalks about
1 mm high with enlarged heads bearing vast numbers of minute, pear-shaped
spores embedded in a translucent drop of sticky liquid. Spores in the vessels
increase in a yeastlike manner. The perithecial stage, not found in nature,
has been produced in culture by crossing plus and minus strains of the
fungus.
Although the smaller European elm bark beetle is chiey responsible for
spread of the pathogen, at times the native elm bark beetle, Hylurgopinus
rupes, is the agent. When the adult beetles emerge from under the bark
of dead or dying trees, they bring along sticky spores on their bodies and
deposit them as they feed in the crotches of young twigs or leaf axils of near-
WILT DISEASES 685
Figure 3.62 Dutch Elm Disease. A branch cut to show discoloration of wood; B wound in twig crotch due to
beetle feeding; C bark-beetle carrier of the fungus; D egg and larval galleries of the beetle engraved on sapwood
by healthy trees. Although the beetles feed on healthy wood, usually within
200 feet of their original tree, they breed only on weakened or dying wood
and may y some distance for it. The European female tunnels out a brood
gallery 1 or 2 inches long in the wood, and when the larvae hatch, they tunnel
at right angles across the wood (Fig. 3.62). There is a second brood in August
and September, but the overwintering one, emerging in May, is most to be
feared. Because the disease often follows trafc routes, automobiles proba-
bly account for a good deal of long-distance spread. So far as we know, the
only other natural means of infection is by root grafts, made when trees are
planted so close together that their roots touch. This is another argument for
diversied planting, rather than streets closely lined with but one type of tree.
Control. In the rst few frantic years an enormous amount of money (more
than $26 million) was spent on trying to eradicate the disease by removing
and burning diseased trees; and while this was undoubtedly helpful, it did
not stop the spread of wilt. The Federal government has now left the control
of Dutch elm disease up to the communities and is restricting its efforts to
research. Many towns have taken a laissez-faire attitude, thinking that our
elms are doomed anyway, so why waste money? Other, more enlightened
communities have proved that a sustained control program keeps the disease
down to a negligible 1 or 2%, or less, and that the cost is far, far less than
that of continuous removal of dead trees.
The Midwestern Chapter of the National Shade Tree Conference, in its Guide
for Community-Wide Control of Dutch Elm Disease, suggests:
1. Survey of the total elm tree population to be protected.
2. Symptom scouting for detection of diseased trees and sanitation scouting
for badly weakened elms and wood piles containing elm wood.
3. Destruction of known sources of elm wood actually or potentially haz-
ardous for spread of disease. Elm wood piles should be destroyed com-
pletely, or each log stripped of bark and the bark destroyed. Diseased trees
should be burned, on site if possible, or thoroughly sprayed. Wood chips
from diseased elms may still carry the fungus; material should be burned,
not used for mulches.
4. Spraying of healthy trees to prevent infection.
5. Maintenance of elms in healthy condition to prevent invasions of bark
beetles. This includes proper watering and fertilizing, spraying to control
summer foliage pests if necessary.
A single annual DORMANT SPRAY is now considered sufcient to protect
healthy elms from bark beetles if enough material is used and complete cov-
erage is obtained. This spray was originally a very heavy dosage of DDT,
which caused some bird mortality and other environmental problems. Some
communities, of which Greenwich, Connecticut is a good example, gure
that they cannot afford not to spray, for it costs less to spray for control than
to remove a dead tree. Where dormant spraying and sanitation have been
combined consistently, the annual loss from Dutch elm disease has been kept
to 1% or less.
Chemotherapy, injection of chemicals that will inactivate the fungus, has
been a promising line of research for many years. A parasitic European wasp
is now being bred at several laboratories for release against the bark beetles.
To have elms in our future we must keep on planting them. Some forms, such
as the Christine Buisman and Groeneveld elms, are quite resistant although
not immune. Chinese and Siberian elms are resistant.
Dothiorella
Cankers.
Dothiorella ulmi. Dothiorella or Cephalosporium Wilt of elms. Die-
back, rather common on American elms, occasional on slippery and Siberi-
an elms in central and eastern states. The names are confusing. In culture
the fungus develops spores as in Cephalosporium, but in nature Dothiore-
lia-type pycnidia are developed on bark of killed twigs. The fungus has also
WILT DISEASES 687
been classied as Deuterophoma. Spores are extruded in a sticky mass and
are disseminated by wind, rain, possibly insects. Infection is through insect
or other wounds on foliage. The mycelium proceeds from leaf petioles into
wood, where it is conned to the vessels. The foliage wilts and yellows; there
are gradual dying back of the crown and a brownish discoloration in outer
rings of the wood. Without laboratory diagnosis the disease cannot be pos-
itively separated from Dutch elm disease, but the elliptical cankers on the
stems, with small black specks of pycnidia, provide one diagnostic symp-
tom. Older trees die 3 to several years after rst symptoms; nursery trees, in
1 or 2 years. Some trees recover, and some remain infected for many years
without showing much effect.
Control. Prune out infected branches a foot or more below the lowest point of
discoloration. Promote vigor by feeding, watering, aerating soil. The inclu-
sion of a fungicide in sprays for elm-leaf beetles or cankerworms might be
helpful.
Fusarium
Rots.
Fusarium annuum (F. solani). Fusarium Wilt of chili pepper. Under-
ground stems are dry, brown, but the roots soft and water-soaked; plants wilt
and die rapidly. Spores are spread in irrigation water and with wind-blown
particles of soil. Avoid heavy, poorly drained soils.
Fusarium foetens. Wilt on begonia.
Fusarium oxysporum. Wilt on pyracantha and basil. Blight and Wilt on
purple coneower (Echinacea).
Fusarium oxysporum f. sp. apii. Celery Wilt, Yellows, general in north-
ern celery districts. There are three strains of the fungus, all causing stunting,
vascular discoloration, crown and root rot, but one form causes the entire
plant to turn yellow at high temperatures, producing brittle stalks with a bit-
ter taste. Another strain causes downward curling of young heart leaves, and
the third produces no above-ground symptoms except stunting. The fungus
persists indenitely in soil. Golden, self-blanching varieties are more sus-
ceptible. Grow green petiole celery or somewhat resistant Michigan Golden,
Cornell 19, Tall Golden Plume, Golden Pascal or Emerson Pascal.
Fusariumoxysporumf. sp. asparagi. FusariumWilt of asparagus, a major
factor in asparagus decline in California, found in most plantings. The fungus
lives in soil and may be distributed on seed.
Fusariumoxysporum f. sp. barbati. FusariumWilt of sweet william. New
growth is yellowed; plants are stunted; leaves point downward and are tinged
with tan as they die. Roots and lower stem are discolored brown. Plant in new
or sterilized soil.
Fusarium oxysporum f. sp. batatas.

Rots.
Fusarium oxysporum f. sp. betae. Fusarium Yellows on sugar beet.
Fusarium oxysporum f. sp. callistephi. Aster Wilt, one of the most serious
diseases of China aster, unless resistant seed is used. Plants wilt, wither, and
die at any age from seedlings to full bloom. Older plants are often stunted,
with a one-sided development and a brown discoloration of the vascular sys-
tem. Sometimes all lower leaves are wilted, with blackening at base of stem,
often with a pink spore mass at ground level. Plants in full bloom may sud-
denly droop their heads. Such symptoms are in contrast to the mycoplasma-
like disease, aster yellows, where the plant remains upright, although stunted
and yellow. The fungus is seed-borne and persists in the soil many years.
Control. Sterilize soil for seedbeds. Some seedsmen provide seed of wilt-
resistant varieties, but maintaining resistance means continuous selection
from asters grown on heavily infested soil under conditions highly favorable
for infection, and this is an expensive process.
Fusarium oxysporum f. sp. cattleyae. Wilt of cattleya orchids. The fungus
was isolated from a private collection in Ohio. Leaves wilted, roots abscised
and decayed; owers fewer, smaller, short-lived.
Fusarium oxysporum f. sp. chrysanthemi. Fusarium Wilt on chrysanthe-
mums.
Fusarium oxysporum f. sp. conglutinans. Cabbage Yellows, Fusarium
Wilt, general on cabbage and other cruciers, probably the most destructive
disease of such hosts in the Midwest, perhaps other sections. It is serious
on cabbage, kohlrabi, and collards. Brussels sprouts, cauliower and broc-
coli are moderately susceptible in hot dry seasons. The fungus, which can
live many years in the soil, enters through the roots, usually right after trans-
planting or at the rst hot weather, with potassium deciency as well as heat
thought to favor infection. The fungus progresses upward in the xylem, not
invading other elements until the plant dies.
The most striking symptom is the dull yellow to greenish color of the foliage,
together with a warping or curling of basal leaves. Leaves are killed and
WILT DISEASES 689
shed from the base up; the woody tissue in the stem is brown, with a water-
soaked appearance. The fungus is spread by soil clinging to farm imple-
ments, drainage, water, wind, animals and infected seedlings. Once the dis-
ease is established, general sanitation and crop rotation are of little help
against a fungus that can survive so long without a susceptible host.
Control. Once soil is infested resistant varieties offer the only hope. Many
have been developed, including Jersey Queen, Marion Market, Wisconsin
Golden Acre, Resistant Detroit, resistant strains of Early Jersey Wakeeld,
Charleston Wakeeld, Globe, Wisconsin All Season and Wisconsin Hollan-
der.
Fusarium oxysporum f. sp. cucumerinum. Cucumber Wilt. A newly rec-
ognized form of Fusarium highly pathogenic to cucumber and muskmelon
in Florida, only slightly pathogenic to watermelon.
Fusarium oxysporum f. sp. cyclaminis. Fusarium Wilt on cyclamen.
Fusarium oxysporum f. sp. cubense. Wilt of banana.
Fusarium oxysporum f. sp. dianthi. Carnation Fusarium Wilt, Yellows,
Branch Rot, general. The rst symptom is a slow withering of shoots, often
accompanied by change of color from normal deep green to lighter green to
pale straw yellow. Plants appear wilted, especially during the warmer part of
the day. Only one side of the plant may be affected, resulting in distortion and
tendency to curl. If the stem is split, a brownish streak is seen in the vascular
system. There may be a dry, shreddy rot of affected wood and cortex. Plants
may be infected at any age, but succumb faster if attacked when young. This
species of Fusarium does not rot roots; see F. roseum under Rots for the form
causing stem and root rot on carnation.
Control. Sterilize greenhouse soil and benches; take cuttings from healthy
mother block; avoid overwatering. Drenching newly atted or benched
plants has reduced the number of wilted plants but does not replace steaming
or otherwise sterilizing soil.
Fusarium oxysporum f. sp. erythroxyli. Wilt of Erythroxylum.
Fusarium oxysporum f. sp. gladioli.

Rots.
Fusarium oxysporum f. sp. hebae. Fusarium Wilt of Hebe buxifolia, and
veronica. Reported as killing nursery plants in California.
Fusarium oxysporum f. sp. lactucum. Wilt of lettuce.
Fusariumoxysporum f. sp. lycopersici. Fusarium Wilt of tomato, general,
in many sections the most damaging tomato disease in eld and greenhouse.
Chief losses are in states where air temperatures are rather high during most
of the season, susceptible varieties dying or producing little fruit. Losses go
up to 30,000 tons of canning tomatoes, or 10 to 35% of the crop in many
states.
In seedlings there is downward curvature of the oldest leaves followed by
wilting and death. In older plants the disease is most evident as fruit begins
to mature, lower leaves turning yellow, rst on one side of the stem or leaets
on one side of the petiole. One shoot may be killed before the rest of the plant
shows symptoms.
The fungus enters through roots and grows into the stem, where it produces
the toxic substances causing wilting and eventual death. The vascular system
in the stem shows a dark brown discoloration. In severe infections the fungus
grows into fruit and seeds, but such fruits usually drop, and seed is not used.
Almost all original infection comes from the soil, the Fusarium operating
best in light sandy soils and at temperatures between 80 and 90 F, but the
disease is spread widely in transplants. It is encouraged by low potassium
and high nitrogen nutrients.
Control. Start seedlings in clean soil; do not grow in the same land more
than once in 4 years. The use of resistant varieties is the chief means of
control. Marglobe, Pritchard and Rutgers are moderately resistant, but infes-
tation by nematodes may predispose even these to wilt. Pan America, South-
land, Homestead and Jefferson are more highly resistant. Treating soil with
nematicides may reduce incidence of wilt even though the wilt pathogen is
not killed.
Fusarium oxysporum f. sp. melonis. Muskmelon Fusarium Wilt, similar
to that of watermelon, important in Minnesota, New York, New Jersey and
Maryland. Seeds rot in soil; seedlings damp-off; vines wilt. Fungus persists
in soil and is carried internally in seed. Varieties Golden Gopher and Iroquois
are quite resistant.
Fusarium oxysporum f. sp. niveum. Watermelon Wilt, general on water-
melon, also on citron. The fungus is transported in and on seed and persists in
soil 15 to 18 years. It rots seeds or seedlings, causes wilting of plant, some-
times with cottony mycelium on surface of dying vines. Resistant varieties
include Improved Kleckley Sweet and Klondike.
Fusarium oxysporum f. sp. perniciosum. Mimosa Wilt on mimosa from
New Jersey and Maryland to Florida. This extremely pernicious wilt start-
ed about 1930 at Tryon, North Carolina, and mimosas have wilted and died
at a rapid rate ever since. The wilt appeared in one city block at Morgan-
town, North Carolina in 1943, and by 1947 trees were dead and dying on
232 blocks.
WILT DISEASES 691
The rst external symptom is a wilting and yellowing of leaves on some of
the branches, causing foliage to hang down, then die and drop. Death of the
tree follows from a month to a year after rst infection. The trunk has a brown
ring of discolored sapwood, usually in the current annual ring, and the color
may extend out into the branches. The xylem is plugged with brown gum-
my substances. Small branches may have a one-sided wilting with the bark
attened over collapsing tissue. The disease has been spreading in Maryland
since 1947, in Florida since 1952.
As with other Fusaria, this is a soil fungus entering through the roots, and
eradication of diseased trees has no effect on spread of the wilt. Nema-
todes, by their wounds, may increase the incidence of wilt. Out of a great
many seedlings grown from seed collected from Maryland to Louisiana,
inoculated several times with the fungus and planted in infested soil, some
have remained mostly disease-free. These have been propagated by the
U.S. Department of Agriculture. Released for commercial sale are Charlotte
and Tryon.
Fusarium oxysporum f. sp. pisi. Pea Wilt, caused by race 1 of this
pathogen and Near Wilt, caused by race 2. Race 1, conned to pea, produces
stunted plants, pale yellow green, with leaves curled downward, stem thick-
ened and brittle near the ground. Plants wilt and die prematurely. The disease
may cause more or less circular bare spots in the eld, enlarging each year
if peas are planted continuously, encouraged by high soil temperature. Some
commercial pea varieties are resistant to race 1 but not to race 2. Delwiche
Commando was the rst variety introduced resistant to both races.
Fusarium oxysporum f. sp. raphani. Radish Wilt. Young plants turn yel-
low and die; others are stunted, with discoloration of roots.
Fusarium oxysporum f. sp. spinaciae. Fusarium Wilt of spinach. Plants
are pale; leaves roll inward, gradually die. The wilt is serious in Texas and
Virginia. One form of the mosaic-resistant Savoy spinach is also resistant to
wilt.
Fusarium oxysporum f. sp. tracheiphilum. Wilt of cowpea.
Fusarium solani f. sp. pisi. Wilt of chick-pea.
Hendersonula
Pycnidia dark, separate; spores dark with several cells.
Hendersonula toruloidea. Branch Wilt of walnut. Canker, destructive to
Persian walnuts but associated with sunburn of affected branches. The fun-
gus is a wound parasite.
Phialophora
Rots.
Phialophora gregata. Wilt of chick-pea.
Phomopsis
Blights.
Phomopsis sp. Wilt on ice plant.
Phytophthora
Blights.
Phytophthora cactorum. Wilt of blue laceower and babys breath.
Phytophthora cinnamomi. Rhododendron Wilt. A wilt of young stock,
grafted plants 2 to 3 years old, seldom on older shrubs, most severe on
Rhododendron ponticum. The foliage is rst dull yellow, then permanently
wilted, roots are decayed; stems are brown at soil level and below. Remove
infected stock from frames immediately; avoid excessive irrigation; keep soil
acidity at pH 4.0 to 4.5; provide shade and mulch for young plants. This
pathogen also causes wilt of Japanese umbrella tree.
See under Rots for this fungus at work on many other plants.
Pythium
Rots.
Pythium myriotylum. Wilt on peanut.
Pythium tracheiphilum. Wilt on lettuce and also leaf blight.
Pythium aphanidermatum. Wilt of Nicotiana.
WILT DISEASES 693
Rhizoctonia
Blights.
Rhizoctonia solani. Wilt of watermelon.
Sclerotinia
Blights.
Sclerotinia minor. Wilt of lupine and wild garlic.
Verticillium
Conidia one-celled, hyaline, globose to ellipsoid, formed at tips of whorled branches and
separating readily from tips.
Verticillium albo-atrum. VerticilliumWilt, Maple Wilt of many ornamen-
tal trees, shrubs, fruits, owers and vegetables. The fungus was rst isolated
from potatoes in Germany in 1870 but apparently was present in California
as early as 1850. It attacks nearly 300 cultivated plants of widely diverse
types and may persist as a saprophyte in the soil 15 years or more.
Of the ornamental tree hosts silver maples are most susceptible, then sugar
and red maples, elms, with occasional reports on ailanthus, alfalfa, aspen,
ash, boxelder, beech, black locust, camphor-tree, carob, catalpa, Chinaberry,
cucumber, deerbrush, dogwood, goldenrain, horse-chestnut, India hawthorn,
redbud, linden, magnolia, oak, osage-orange, olive, pistachio, persimmon,
periwinkle, Russian olive, sassafras, strawberry, smoke-tree, tulip-tree, wal-
nut, mango, sunower and hickory. Maples may wilt suddenly in midsum-
mer, often a large branch or one side of the tree drying and dying while the
other side stays fresh. The sapwood of the infected side has greenish streaks,
and sometimes slime ux develops on the bark. The disease can be chronic,
progressing slowly for several seasons, or acute, affecting the entire tree in
a few weeks. In elms the leaves may be smaller than normal, with a drooping
accidity in hot hours of the day. Later there is a slight yellowing, deepen-
ing until the foliage is a striking lemon yellow. Defoliation starts at time of
rst yellowing, and quite often branchlets drop as well as leaves. Sapwood
discoloration is brown, and the disease cannot be told positively from Dutch
elm disease without laboratory cultures. Tyloses and gums are formed in
the wood as with other toxin-producing fungi. The fungus always progress-
es upward through the xylem vessels so there is little danger of downward
infection of the main trunk from pruning operations. Progress is slowed by
adequate moisture and by high nitrogen fertilizers, ammonium sulfate being
particularly helpful.
Verticillium wilt is also a problem on rose understock. Ragged Robin, Odora-
ta, and Multiora are very susceptible, Dr. Huey less susceptible, and Manetti
resistant.
In fruit trees the wilt is often known as black heart or verticillosis. It is com-
mon in apricots, less so in almonds and peaches; branches may drop their
leaves and die. Also susceptible are sweet and sour cherry, avocado, plum
and prune.
On bush fruits raspberry, blackberry, dewberry and youngberry the dis-
ease is commonly known as blue stem. The symptoms appear late in the
season, leaves turning pale, cane tips bending downward, canes taking on
a bluish color, lower leaves wilting and drying. Death is often delayed until
the season after rst infection. Black raspberries are more susceptible than
red. The disease is sometime serious on strawberries, especially in Califor-
nia, but cannot always be separated from root rots. Plants may collapse in
large areas at the beginning of hot weather.
Verticillium wilt is very destructive to mint in Michigan and Indiana, also
reported, though not so serious in Oregon and Washington. Infected plants
Figure 3.63 Verticillium Wilt on Tomato
WILT DISEASES 695
are stunted, defoliated, and killed; yield of oil is greatly reduced. The fungus
attacks all species of mint, but peppermint is most susceptible. There are
some resistant hybrids. Deep plowing, inverting the soil, has reduced the
amount of wilt.
Verticillium is especially damaging to tomatoes in Utah and California. First
symptoms are yellowing of older leaves and wilting of tips during the day;
Figure 3.64 Verticillium Wilt on Potato
Figure 3.65 Verticillium Wilt on Snapdragon
WILT DISEASES 697
later, margins of all leaves curl upward, then leaves drop (see Fig. 3.63).
Plants are stunted; fruit is small. Moderately resistant varieties Riverside
and Essar have been developed for California. Symptoms on potatoes are
rather indenite, but often there is yellowing of lower leaves, shortening of
internodes, and rosetting of the top (see Fig. 3.64). Resistant varieties may
be symptomless hosts. Verticillium wilt is common on eggplant and okra,
rather rare on pepper. It occurs on Chinese yard-long bean, rhubarb and New
Zealand spinach.
On herbaceous perennials in eastern gardens I nd Verticillium wilt com-
mon on aconite and chrysanthemum, with leaves turning dark brown and
hanging down along the stem. When the stem is cut across near the base,
a circle of black dots indicates the fungus in the vessels. Such plants seldom
die immediately but ower poorly and gradually peter out. Wilt was serious
on greenhouse chrysanthemums until a wholesale commercial concern start-
ed to provide healthy propagating stock from cultured cuttings. Other orna-
mental hosts include abutilon, aralia, barberry, begonia, China aster, carna-
tion, dahlia, fremontia, geranium, marguerite, peony, poppy, snapdragon (see
Fig. 3.65), stock and viburnum.
Control. Sometimes it is possible to prune out an infected maple and still save
the tree, but often the dying tree must be taken out. Neither maple nor elm
should be replanted in the same spot. Do not transfer plants from areas where
wilt has appeared. Do not set raspberries following potatoes or tomatoes; do
not use tomatoes after eggplant or potatoes without a long rotation. Proper
fertilization and adequate watering may help trees to recover from wilt.
Verticillum dahliae is considered by some a synonym of V. albo-atrum and
by others as a distinct species; reported as causing wilt of dahlias, mint,
marigold, ice plant, barley, wheat, oat, potato, Leucospermum, impatiens,
giant hyssop, globe artichoke, ash, cabbage, Cineraria and Echinacea, and
other plants. This form has microsclerotia and grows on agar at slightly high-
er temperatures.
Verticillium fungicola. Dry Bubble of oyster mushroom. Infection of
sporophores at pin or button stage cause development of typical dry bubbles;
mature sporophores show cracking and curling of tissues, and depressed,
brown, necrotic areas.
WITCHWEED
A parasitic weed, Striga asiatica, new to the western hemisphere, was report-
ed from North Carolina in 1956 and later from South Carolina, although
apparently it was rst seen in the latter state in 1951 following construction
of a power line across a farm. The plant is an obligate root parasite of corn
and crabgrass, perhaps other plants. It is 2 to 15 inches high, foliage varying
from dark to light green, with linear leaves curving downward, tubular ow-
ers with two-lipped corolla, cardinal red on the upper surface with a yellow
eye, straw yellow on the lower surface. The numerous brown seeds are very
minute.
Witchweed is reported from other countries on 63 plant species, 56 of them
members of the Gramineae (grains and grasses). Tests in the United States
with 77 non-gramineous hosts found none parasitized by witchweed, but 45
species of our grasses and grains are susceptible to this new pest. To help
in eradication, report suspicious weeds immediately to your county agent or
extension pathologist.

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CHP 3A10.1007 2F978 1 4020 4585 1 - 3

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Chapter 3

Chap. 4. At the beginning of

Chap. 4 you will nd a list of headings under

Chap. 4, the key word, for example, rot or blight, is giv-

Glomerella cingulata). Snapdragon An-

Glomerella cingulata) and other species. Orchid

Glomerella cingulata). Rhubarb Anthrac-

Glomerella cingulata). Strawberry Anthrac-

Glomerella cingulata). Lime Anthrac-

Glomerella lindemuthianum). Bean

Glomerella cingulata).Tomato Anthrac-

Glomerella cingulata). Pea Anthracnose, Leaf

Glomerella cingulata). Anthracnose

Phlyctema vagabunda). Anthracnose,

Glomerella cingulata). Maple Anthrac-

Glomerella cingulata). Lime Anthrac-

Glomerella cingulata). (possibly identical

Glomerella cingulata). Pepper Anthrac-

Cankers). There are two forms of this species, one being

Cylindrocladium under Blights.

Rots). The pod blight

Cankers). On raspberry, brown dead areas extend into wood; whole

Cankers and also

Glomerella cingulata). Dracaena Tip Blight,

Valsa cincta for further discussion of cankers on stone fruits.

Anthracnose). Camellia tips die back; leaves

Black Mildew). Powdery mildews are

Leveillula taurica). On leaves of mesquite.

Sphaerotheca lanestris for the common live-oak mildew).

Podosphaera pannosa). Rose Mil-

Cankers and Dieback.

Pellicularia lamentosa Root Rot and Crown Rot of clovers..

Cankers and Diebacks.

Turnip Mosaic Potyvirus.

Peach Western X-Disease (under Bacterial Diseases, Phytoplasma).

Tomato Aspermy Cucumovirus.

Bean Yellow Mosaic Potyvirus.

Cherry Ring Spot.

Locust Witches Broom.

Beet Curly Top Hybrigeminivirus.

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