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PUBLIC DOCUMENT No. 47

State Geological and Natural
History Survey

COMMISSIONERS
Henry Roberts, Governor of Connecticut (Chairtnan)
Arthur Twining Hadley, President of Yale University
Bradford Paul Raymond, President of Wesleyan University
Flavel Sweeten Luther, President of Trinity College (Secretary )
RUFUS WHITTAKER STIMSON, President of Connecticut Agricultural College

SUPERINTENDENT
William North Rice

Bulletin No. 5

Hartford Press

The Case, Lockwood & Brainard Company

1905

r

V3

THE USTILAGINE^, OR SMUTS,
OF CONNECTICUT

v..

By
GEORGE PERKINS CLINTON, S.D.

Botanist of Connecticut Agricultural Experiment Station

Hartford Press

The Case, Lockwood & Brainard Company

May, 1905

The Ustilagineae, or Smuts, of Connecticut.

GENERAL CHARACTERS OF THE SMUTS.

The Ustilagineae are an order of parasitic fungi commonly
known as the smuts, of which corn, oat, and onion smuts are
familiar examples. See Figs. 55, 43, 42. Thus the more evi-
dent characters of these fungi are the black, dusty masses that
break out on the surface of the infected plants (hosts). In
these macroscopic characters the smuts are often very similar
to the rusts, but with a little experience one can readily dis-
tinguish them from these even with the naked eye. The dusty
livedo stage of the rusts, for instance, is of a lighter reddish
color, and the darker teleuto stage is usually more firmly em-
bedded in the plant tissues, than are the smuts. Not all of the
smuts, however, produce dark dusty outbreaks, since the white
smuts are light colored and usually permanently embedded in
the tissues of the host. Generally they occur in the leaves,
and often are distinguished by light colored spots produced
without distortion of tissues. More experience is needed to
distinguish these forms. See Figs. 29-34.

The smuts occur on a great variety of flowering plants,
and at least 35 different families of plants in North America
and 17 in Connecticut are subject to their attack. The
grasses, however, are by far the most frequent hosts. Be-
tween 500 and 600 species of smuts have been described from
different parts of the world, and over 200 of these occur in
North America. The number of species known from this
state * is 50, and these represent 12 of the 19 genera reported
from North America. While time no doubt will reveal a few
more species and a greater number of hosts not reported here,
we may still consider the group rather thoroughly worked up,
since, as yet, only one other state has reported a greater num-
ber of species.

* The list of smuts reported in this paper is based chiefly on the collections
made by the writer during the past three seasons. Specimens of these are to be
found in the herbarium of the Conn. Agr. Exp. Station at New Haven and in the
writer's herbarium. Collections made by others are indicated in each case by the
name of the collector.

6 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

LIFE HISTORY OF THE SMUTS.

In the study of these fungi, there are three parts that we
need to consider, namely, the mycelium, the spores, and the
sori.

Mycelium is the technical name for the vegetative part of
the smut, by means of which it spreads through its host and
gathers nourishment. This consists of microscopic threads
confined to the interior of the host, so it is necessary to exam-
ine very thin cross sections of the tissues under the micro-
scope to see this stage. See Fig. 26. These threads vary but
little with the different species, and therefore are of no value
in classifying the different species. They are simple, hyaline,
more or less branched, and at first filled with protoplasmic
contents, but gradually lose most of their contents and be-
come septate. They push their way between the cells of
the host or sometimes through them. Usually, however, they
enter the cells only by short special branches called haustoria,
whose special function is to gather nourishment from the cell.
The mycelium may be localized or rather generally spread
through the host. In those cases where it gains entrance
through the germinating seed, it usually remains evident fi-
nally only at the nodes or where it breaks forth to the exterior
in its fruiting stage. In perennial plants the mycelium often
becomes established in the perennial parts, from which each
year it sends threads into the new growth.

Eventually the mycelium becomes prominent in certain
regions of the host, and there undergoes special modification
to give rise to the reproductive or spore stage. The walls of
the mycelial threads that form the spores are generally indis-
tinct through more or less complete gelatinization. The
spores are usually formed from contents in the interior of
these fertile threads. This gelatinization apparently serves
to nourish the developing spores, as often all signs of the
threads disappear on maturity of the spores. In some cases,
as in the genus Neovossia, Fig. 13, and often in Entyloma,
however, these threads remain as more or less evident envel-
opes to the spores.

Spores are the bodies by which the smuts reproduce them-
selves. See Figs. 1-20. They are the evident, usually ex-

No. 5.] USTILAGINE.E OF CONNECTICUT. 7

ternal part of the fungus, making up the smutty masses visible
to the naked eye, and are the characteristic and variable parts
upon which species are based. The spores are usually so
formed that at maturity they break out to the exterior of the
host in a dusty mass that is easily scattered, thus securing
their dispersal. In some species, however, they are embedded
in the host so that they are liberated only on the rotting of the
tissues, and sometimes they germinate in situ, and secure their
dispersal by the formation of secondary spores of a different
nature. While easily seen in mass, the spores are really
microscopic in size, varying according to species from 4ju to
35^.* They are usually subspherical or spherical, but vary
to ovoid, ellipsoidal, or even oblong. Pressure often makes
them more or less polyhedral, or irregularly so. They are
generally simple, consisting of single separate cells ; but are
sometimes bound more or less permanently into spore balls,
Figs, ya, 10a. These spore balls may consist entirely of fer-
tile cells (spores) or they may have an external coating (cor-
tex) of sterile cells, Fig. 19, or an internal matrix of sterile
cells or even of threads, Fig. 20. The spore walls may be
smooth, or marked with minute spines (echinulations if sharp,
verruculations if dull), or reticulated with a net-work of
ridges or wings. In color they are hyaline, yellowish, red-
dish or olive brown, violet, or purplish. Often the color is
so deep that the spores are opaque or nearly so. Upon these
variations the genera and species are chiefly classified, though
on the whole the variations are not so great as with many
of the other fungi. This is partly due to the fact that the
spore-producing threads are not diversified or permanent.

In some few species, most frequently in the genus Enty-
loma, the mycelium also gives rise to secondary spores called
conidia. These are usually hyaline, elongated, ephemeral,
and are produced originally on the exterior of the host, gener-
ally from mycelial tufts protruding through the stomates and
forming these conidia at their tips. Their object is to spread
the smut over the host or to new hosts immediately.

Sori are the evident masses of the spores that break but
singly or in clusters on the various parts of the hosts. See

* A ix, or micron, is one twenty-five thousandth of an inch.

8 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

Figs. 27-55. These are protected more or less permanently
by thin coverings of plant tissue, or in some genera, as Spha-
celotheca, and often in Cintractia, by false membranes com-
posed chiefly of 'fungous cells or threads, Fig. 5a. The sori
vary greatly in size, sometimes forming outbreaks smaller
than a pin-head, and sometimes, as often in corn smut, reach-
ing several inches in diameter. Usually each species has a
definite place on its host where the sori appear. This may be
on the leaves, Fig. 35, on the stems, Fig. 28, in the blossoms
as a whole, Fig. 54, or confined to the anthers, ovaries, Fig. 52,
or seeds, Fig. 48. Occasionally a smut may break out in any of
these places. In the grasses, when infecting the inflorescence,
the sorus may be confined to the ovary, infest the spikelet as a
whole, or even involve the entire inflorescence. Sometimes
all of the ovaries or spikelets are infested and sometimes only
part of them. The characters of the sori, of course, are modi-
fied somewhat by the part of the host in which they develop,
and also by the kind of host on which the smut occurs.
Usually the sori are dusty and easily dissipated. With a few
species they are hard, and the agglutinated spore mass is dis-
seminated by gradual disintegration, often through the ab-
sorption and loss of water. In these cases there is sometimes
a gradual ripening of the spores from the outside of the sorus
inward. With some genera {Entyloma, Doassansia, etc.) the
sorus is rather permanently embedded in the tissues of the
host, usually in the leaves, and often produces only a discolora-
tion of these. The species of one of the European genera
form gall-like growths in the roots.

Germination of Spores. There are two chief types of
germination of the Ustilaginese upon which the two families
are based. In the Ustilaginacecr, Fig. 21, the spores, when
placed in a drop of water, send out a single, hyaline thread,
several times their length, which usually divides into about
four cells by cross partitions, or septa. This is called the
promycelium. Usually at the apex of each cell one or more
elongated thin-walled spores, or sporidia, are budded out, the
apical cell bearing its sporidium terminally and the Others lat-
erally. The sporidia when full-sized are pinched off at the
base, and others are sent out until the protoplasm of the cell

NO. 5.] USTILAGINE^E OF CONNECTICUT. 9

is exhausted. These sporidia germinate by a short slender
thread (infection thread) that usually pushes out from near
one end, Fig. 24. In some species the cells of the promyce-
lium, instead of forming the sporidia, give rise directly to the
infection threads, Fig. 23. Sometimes the adjacent cells of the
promycelium become connected by closely applied short
threads, forming buckle or knee joints, and from these may
develop infection threads. Some species germinate directly
into elongated threads which scarcely partake of the nature of
a promycelium. When nutrient is added to the drop of
water, the spores germinate much more luxuriantly, the spor-
idia usually sprouting out other sporidia while still attached
to the promycelium. Especially in a solid cultural medium
these chains develop a more or less complicated system of
branching, Fig. 25. In a liquid solution, the sporidia usually
soon fall off from the promycelium, but continue to multiply
by the yeast fashions of budding new sporidia, which soon sep-
arate and develop others, until the nourishment is exhausted.
Then the sporidia may develop infection threads.

The second chief type of germination is shown by the spe-
cies of Tillctiacccc, Fig. 22. The simple or septate promyce-
lium in this case bears all of the elongated sporidia in a terminal
cluster. Sometimes these primary sporidia develop terminal
secondary sporidia. With some species the sporidia, before
or after falling off from the promycelium, become yoked in
pairs by short connecting threads. In nutrient solutions some
of the species develop finally a complicated mycelium that
bears numerous aerial sporidia or conidia quite different from
the normal type produced in water. The conidia develop in-
fection threads on germination.

Infection of Host. The host is entered by the fine infec-
tion threads of the promycelia or the sporidia boring through
its tissues into the interior. Once inside, these develop the
mycelium of which we have already spoken. Many of the
smuts infect their host only through the young hypocotyl or
epicotyl of the seedling. In these cases the smut usually can
not gain successful entrance after the plants appear above
ground. Once inside, the mycelium rapidly penetrates the
young tissues, seeking to gain access to the growing tip, where

IO CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

it then follows the upward growth of the plant, often not giv-
ing evidence of its presence until it breaks out in its spore
stage in the fruiting organs of the host two or three months
later. This is the case with the oat smuts, wheat and barley
smuts, and the grain smut of broom-corn. In this latter case
the mycelium has traveled upward through six to ten feet of
the cane. In other species infection may take place, through
any very young tissue of the host upon which the germs may
be blown or washed into contact. This is the case with corn,
and the corn smut appears soon after infection in its smutty
outbreaks, the mycelium usually remaining localized. With
the Entylomas the infection is largely confined to the leaves,
and apparently the mycelium is limited to the vicinity of the
sorus.

ECONOMIC IMPORTANCE.

Injury. While most of the smuts occur on plants of no
economic importance, there are at least twenty-five species in
America that do more or less injury to cultivated plants. Some
of these cause such serious injury that they are counted among
the worst species of parasitic fungi. It is with the cereals that
the smuts do most damage. Wheat, oats, barley, rye, and corn
are all subject to attacks from one or more species. With the
first three hosts the smuts break out in the inflorescence, en-
tirely preventing the formation of the seed. It thus becomes
an easy matter to determine rather accurately the per cent, of
loss in fields of these grains by counting the number of smutted
and of free heads in a given area. Oat smut, for instance, in
our central states generally claims from one to fifteen per
cent, of the grain, and in some few fields as high as thirty or
forty per cent, has been destroyed. In Illinois there is perhaps
an average loss per year of one million dollars worth of oats,
caused by the two smuts of this plant. In Indiana Arthur
found one field of wheat where fifty per cent, of the grain was
destroyed by the stinking smut. In the northwestern states,
in the wheat districts, the loss caused by this latter smut is
very great, as it is one of their worst fungous pests. When
abundant it renders the grain inferior for milling purposes, as
the spores may be so numerous in the flour that they darken

No. 5.] USTILAGINE^E OF CONNECTICUT. I I

it. In South Carolina during recent years a smut of rice has
been introduced from Japan, and causes some loss to this crop.
In the eastern states where onions are raised extensively, the
onion smut often does considerable injury. In this case the
smut becomes established in the soil and often prevents profit-
able onion culture on this land. Corn smut is common every-
where, and especially on sweet corn causes considerable injury.
In Connecticut the most injurious smuts, from an economic
standpoint, are those of onion, corn, barley, and oats.

Prevention. With those plants where infection takes place
through the germinating seed, the danger usually comes from
the spores that mechanically adhere to the seed. It has been
found that, if these are killed, the crop from this seed will be
free from smut. Investigations have shown that certain treat-
ment of the grain with chemical solutions or with hot water
will kill the spores with little or no injury to the seed. For
instance, it has long been known that soaking the seed of wheat
in a solution of copper sulphate of a certain strength for a cer-
tain time serves to prevent or lessen the amount of stinking
smut in the crop. Later it was found that soaking the seed
for ten or fifteen minutes in hot water at a temperature of
1320 to 1 350 F. was a more efficient remedy, since there was
less likelihood of injury to the seed. More recently still came
the less cumbersome practice of sprinkling piles of the grain
with formalin, one pint to fifty gallons of water, stirring the
grain to thoroughly wet it, and leaving it in piles or in bags
over night for the fumes to act on the spores. The smuts of
oats, the covered smut of barley, the stinking smut of wheat,
the grain smut of sorghum and broom-corn, and the grain smut
of millet, have all yielded to seed treatment. The loose smuts
of wheat and barley can be prevented or lessened by a severer
method of the hot water treatment (in which a preliminary
soaking of several hours in cold water precedes the hot water
treatment), but this also injures the seed more or less, so that
a greater quantity per acre must be sown. Corn smut can not
be prevented by seed treatment, as the smut gains entrance
through any young tissue of the host. Some experimenters
have advocated the removal of the smut-balls as soon as they
appear. It is known that the fungus develops aerial conidia

12 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

in manure, and so fresh manure applied to corn land is apt to
increase the amount of smut. With onion smut we have a
case where the seedlings while coming through the soil are in-
fected by the smut established there through a previous
smutted crop. Onions raised from sets in this land are little
infected. Some success has been had in preventing this smut
by treating the soil with a mixture of lime and sulphur, drilled
into it at time of seeding.

SYSTEMATIC TREATMENT OF CONNECTICUT

SPECIES.

The order Ustilagincce is divided into two families, based
chiefly on the method of spore germination. They are the
Ustilaginacecu (represented in this state by the genera Cintrac-
tia, Schhonella, Sorosporium, Sphacelotheca, Tolyposporium,
Ustilago), and the Tilletiacece (represented by Doassansia,
Entyloma, Neovossia, Tilletia, Tracya, Urocystis).

Key to Genera.

I. Spores simple.

A. Usually forming a dusty sorus at maturity (see also

IB and IIIAia).
i. Large, usually 16-35//.

a. With an elongated hyaline appendage

Neovossia.

b. Without a conspicuous appendage. Tilletia.
2. Small to medium, usually 5-18^.

a. Sorus covered with a false membrane of

definite fungous cells Sphacelotheca.

b. Protecting membrane of plant tissue only

(sometimes with gelatinized hyphae)

Ustilago.

B. More or less firmly agglutinated at maturity

Cintractia.

C. Permanently imbedded in leaves, producing dis-

colored areas Entyloma,

II. Spores chiefly in pairs, forming an agglutinated sorus. . .

Schizonella.

No. 5.] USTILAGINE^ OF CONNECTICUT. 13

III. Spores in more or less permanent balls.

A. Forming a dusty or granular sorus at maturity.

1. Spore balls consisting only of spores.

a. Often evanescent (Ustilago-like spores) . . .

Sorosporium.

b. Quite permanent, adhering by folds of spore

coats Tolyposporiiim.

2. Spore balls with a cortex of sterile cells

Urocystis.

B. Permanently embedded in the plant tissues.

1. Spore balls with a definite cortex of sterile cells

Doassansia.

2. Spore balls without cortex but with sterile

threads in center Tracya.

USTILAGINACEiE Schrot.

The sori usually form exposed, dusty, or agglutinated
spore masses. Germination is by means of a septate pro-
mycelium, which usually gives rise to terminal and lateral
sporidia (capable of yeast-like multiplication in nutrient solu-
tions), or else to infection threads. Figs. 21, 23-25.

Ustilago Rouss.

The sori occur on various parts of the host, according to
the species,' and at maturity form dusty, usually dark colored
spore masses. The spores are single, separate, of small to
medium size (5-18^), and are produced irregularly in fertile
threads that entirely disappear through gelatinization at ma-
turity. Figs. 1-4, 43-55-

This is the most common and the typical genus of the
smuts. Saccardo, in his Sylloge Fungorum, describes about
250 species for the world: 72 species occur in North America
and 18 in Connecticut. A number of the species possess eco-
nomic importance as parasites of the cereals.

Key to Species of Ustilago.
I. Spores olive or reddish brown.
A. Spores smooth.

1. Sori in leaves forming linear striae

U. longissima.

14 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

2. Sori in spikelets, ovoid.

a. Spores usually lighter colored on one side,

5-9/"-

* Spikelets entirely destroyed. . . U. Hordei.
** Spikelets smutted only at base. . U. levis.

b. Spores uniformly colored, 8-ii/i

U. Crameri.
B. Spores echinulate or verruculose.

1. Sorus in each spikelet, distinct.

a. Spores usually lighter colored on one side,

* Host Arrhenatherum avenaceum (Oat
Grass) U. perennans.

** Host Avena sativa (Oats) . . .U. Avence.

*** Host Hordewm vulgar e (Barley)

U. nuda.

b. Spores uniformly colored, 10-14^

U. Panici-glauci.

2. Sorus involving the entire panicle

U. Rabenhorstiana.

3. Sori in the ovaries.

a. Smooth and inconspicuous, 1-2 mm.

* Spores echinulate U. spermophora.

** Spores verruculose U. Eriocauli.

b. Hispid and conspicuous, 4-10 mm

U. sphcerogena.

4. Sori in leaves (rarely on stem).

a. Forming nodular swellings, often at nodes

U. Crus-?alli.

is1

b. Forming linear striae in the blades,

U. striccformis.

5. Sori breaking out on any part of host, often

large . . . .' U. Zea?.

II. Spores golden brown, verrucose; sori in seeds

U. Oxalidis.

III. Spores violet or purplish, reticulate, sori in flowers.

1. Reticulations rather fine, 1-3/i. . . . U. anomala.

2. Reticulations rather coarse, 2-4/t. U. utriculosa.

No. 5.] USTILAGINEtE OF CONNECTICUT. 1 5

Ustilago longissima (Sow.) Tul. Fig. I. The sori form
more or less distinct, linear striae from a few mm. to length of
leaf; the epidermal covering soon ruptures, and the dark
reddish brown spore mass becomes scattered from the more or
less shredded tissues. The spores are light brown, oblong, or
more commonly ellipsoidal to spherical, smooth (or sometimes
scarcely granular under an immersion lens), 4-8/z in length.

Host and Distr. : Glyceria grandis, Shaker Station, June
29, 1903.

This species undoubtedly has a wider distribution than in-
dicated above.

Ustilago Hordei (Pers.) Kell. & Sw. Fig. 46. The
sori occur in every spikelet, forming an ovate, adhering, purple-
black spore mass, 6-10 mm. in length, and rather permanently
covered by the thin, usually transparent plant tissue. The
spores are reddish brown, slightly lighter colored on one side,
chiefly subspherical or spherical, smooth, and 5-9^, rarely
(most elongated) 9-1 i/x, in length.

Host and Distr.: Hordeum vulgare, Whitneyville, Sept.
24, 1902.

This covered smut of barley is not nearly so common in
this state as is the loose smut of the same host.

Ustilago levis (Kell. & Sw.) Magn. Fig. 45. The sori
usually occur in all of the spikelets, are more or less hidden by
the enveloping glumes (which may be infected only at their
base or in the interior), and form rather permanent, black-
brown, semi-agglutinated, ovate spore masses 6-10 mm. in
length. The spores are reddish brown, slightly lighter col-
ored on one side, chiefly subspherical or spherical, smooth,
5-9 n, rarely (most elongated) lip., in length.

Host and Distr. : Avena sativa, New Haven, July 10,
July 28, 1902.

This hidden smut of oats is not so common in our oat fields
as is the loose smut, U. Avence.

Ustilago Crameri Korn. Fig. 44. The sori occur in all
of the spikelets of the spike, destroying especially the inner
and basal parts, and form ovate dusty brown spore masses
2-4 mm. in length. The spores are reddish brown, ovoid to

1 6 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

subspherical, or occasionally more elongated or irregular,
smooth, often with pitted contents, and 8-in* in length.

Host and Distr. : Setaria Italica, New Haven, Sept. n,
1903.

So far this species has been found in this state only once,
in a small plot of Hungarian grass grown at the Experiment
Station. In some of the central and western states it some-
times causes considerable injury to its host. It can be pre-
vented by seed treatment with hot water, formalin, etc.

Ustilago perennans Rostr. The sori occupy all of the
spikelets, usually destroying the basal and inner parts (some-
times even running down slightly on the pedicles) ; they are
oblong, 3-8 mm. in length, and have a somewhat protected
semi-dusty olive brown spore mass. The spores are chiefly
subspherical or spherical, occasionally ellipsoidal to ovate, .
usually lighter colored on one side, minutely echinulate, espe-
cially on lighter side, and 5-8/x in length.

Host and Distr. : Arrhenathcrum avenaceum, South Man-
chester (Thaxter).

This is closely related to the oat smuts. Its mycelium be-
comes perennial in the underground parts of the host, and so
smuts the infested plants year after year.

Ustilago Avenae (Pers.) Jens. Figs. 23, 43. The sori
appear in all of the spikelets, completely destroying all the
floral parts, except a very temporary transparent covering
membrane; eventually the olive brown dusty spore mass be-
comes dissipated, leaving behind only the naked pedicles. The
spores are reddish brown, lighter colored on one side, sub-
spherical to spherical, or occasionally more elongated, mi-
nutely echinulate, especially on the lighter side, and chiefly
5-9^ in length.

Host and Distr. : Avena sativa, New Haven, July 8,
July 28, Aug. 17, 1902, Aug. 9, 1903; West Cornwall, July 18,
1902 ; Westville, July 7, 1903 ; West Haven, July 22, 1903.

This is a common pest in the oat fields of the state, though
the per cent, of infected plants is smaller than in the central
and western states. It yields to seed treatment.

Ustilago nuda (Jens.) Kell. & Sw. Fig. 47. The sori
infest all of the spikelets, changing each into an olive brown,

No. 5.] USTILAGINE.E OF CONNECTICUT. \"J

oblong, dusty spore mass about 6-10 mm. in length ; this is very
temporarily protected by a thin transparent membrane, and
upon dispersal of the spores nothing remains but the naked
rhachis. The spores are lighter colored on one side, sub-
spherical to spherical, or occasionally more elongated, minutely
echinulate, especially on the lighter side, and chiefly 5-9^ in
length.

Host and Distr. : Hordeum vidgare, Storrs, 1901 ; New
Haven, July 8, 1902, June 15, 1904; Whitneyville, Sept. 24,
1902 ; Westville, July 7, 1903.

This is the loose smut of barley, and is rather common in
the fields of this crop. The smuts of barley, oats, and oat
grass, as will be seen by the descriptions, are very similar, in
fact were not long ago considered one species. It requires
the modified form of the hot-water treatment to prevent the
loose smut of barley.

Ustilago Panici-glauci (Wallr.) Wint. Fig. 49. The
sori occur in all of the spikelets, are ovate, 2-3 mm. in length,
and at first protected by thin transparent glumes, but soon rup-
ture these, and scatter the dusty black-brown spore mass. The
spores are dark reddish brown, ovoid, spherical or sometimes
more elongated, prominently and abundantly echinulate, and
10-14,(4 in length.

Host and Distr. : Setaria glauca, Southington, Aug. 8,
1902; Bridgeport, Sept. 15, 1902; Berlin, Oct. 3, 1902; Glas-
tonbury, Oct. 23. 1902; Andover, Sept. 15, 1903; New Canaan,
Sept. 29, 1903 ; Manchester, Oct. 2, 1903.

Throughout the United States this is a common smut on
the yellow fox-tail grass, though it does not occur on the
green fox-tail.

Ustilago Rabenhorstiana Kuhn. Fig. 50. The sorus
involves the entire inflorescence, changing it into a linear or
oblong body, 3-5 cm. in length, which is usually hidden by
the enveloping leaf sheath ; it is covered for a short time by a
very fragile transparent plant membrane, and within the dusty
brown-black spore mass are often elongated remains of the
plant tissues. The spores are reddish or olive brown, ovoid
to spherical, or occasionally slightly angled, echinulate to ver-
ruculose, and 10-14^ in length.
2

1 8 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

Host and Distr. : Panicum sanguinale, Centreville, Aug.
10, Sept. i, 1902; Unionville, Aug. 26, 1902; New Haven,
Sept., 1902, Oct. 5, 1904; Westville, Aug. 25, 1903; Monto-
wese, Sept. 14, 1903 ; New Canaan, Sept. 29, 1903.

This is one of the common grass smuts here as elsewhere,
and is to be looked for in the fall of the year.

Ustilago spermophora B. & C. Fig. 52. The brown-
black dusty sori occur in the ovaries, infesting one here and
there ; they show as small ovate bodies, about 2 mm. in length,
extending between the spreading glumes, and are at first pro-
tected by a thin ovary membrane at the apex of which are the
remains of the styles. The spores are light brown, ovoid to
subspherical, usually prominently echinulate, and 8-1 i[i, or
occasionally 13^ in length.

Host and Distr.: Eragrostis major, New Haven, Oct. 31,
1902 ; Westville, Oct. 17, 1903.

The sori usually occur in only a few of the ovaries, and
are so inconspicuous that the smut is easily overlooked. Fig.
52 shows isolated spikelets of the grass with a single sorus in
each.

Ustilago Eriocauli (Mass.) Clint. The sori occur in the
ovaries, scarcely showing between the glumes as slightly
swollen ovoid bodies about 1 mm. in length; a thin membrane
protects the rather firm, at first semi-agglutinated, but finally
dusty, olive-black spore mass. The spores are polyhedral, sub-
spherical, or occasionally more elongated, rather prominently
verruculose, and 9-15^ in length.

Host and Distr. : Eriocaulon septangnlare, Whitneyville,
Sept. 21, 1902.

It requires very close examination to detect the flower heads
containing this inconspicuous smut, since they resemble the
normal ones. The smut should be searched for in the fall.

Ustilago sphaerogena Burr. Fig. 53. The sori occupy
the ovaries, forming ovate bodies, 4-10 mm. in length, which
are covered by a tough hispid plant membrane that ruptures
irregularly from the apex, disclosing at first an agglutinated,
but finally a dusty, olive brown spore mass. The spores are
ovoid to subspherical, prominently and sharply echinulate, and
9- 1 2/1 in length.

No. 5.] USTILAGINE.E OF CONNECTICUT. 19

Host and Distr. : Panicum Crus-galli, Conn. (Setchell) ;
Savin Rock, Sept. 14, 1902; Westville, Sept. 12, 1902.

This species and the next are very closely related, and oc-
cur on the same host, but are rarely, if ever, found together.
Fig. 53 shows a portion of the flower panicle with a single un-
usually large sorus.

Ustilago Crus-galli Tr. & Earle. The sori form nodules
often encircling the leaf node, or more rarely occur in place
of inflorescence, infecting both leaves and stem; they are one
to several cm. in diameter and protected by a tough hispid
plant membrane, which on rupture discloses an olive brown
spore mass. The spores are reddish brown, ovoid to spherical,
occasionally more elongated, rather bluntly echinulate or even
verruculose, and 10-14/i in length.

Host and Distr. : Panicum Crus-galli, New Haven, Sept.
11, 1903.

Only a single specimen of this smut was found on a culti-
vated variety of this barn-yard grass grown at the Experiment
Station.

Ustilago striaeformis (West.) Niessl. Fig. 51. The
sori occur in the leaves, rarely in the inflorescence, forming
short linear stria?, or by terminal fusion reaching several cm.
in length, and laterally are often so crowded as to cover most
of the leaf ; at first they are covered by the epidermis, but this
soon ruptures, and the dusty brown-black spore masses be-
come scattered from the shredded tissues. The spores are red-
dish brown, vary from ellipsoidal to spherical, or occasionally
irregular, are prominently echinulate, and 9-14^ in length.

Hosts and Distr. : Agrostis alba var. vulgaris, Whitney-
ville, July 20, 1902, May 9, 1903; Centreville, June 12, 1904;
Phlcitm pratense, Whitneyville, May 9, 1903; New Haven,
May 16, 1903.

Both of these hosts are economic plants grown for pasture
or hay, but so far the smut has been found on them in this state
only in door yards. Fig. 51 shows the leaves of Agrostis alba
var. vulgaris shredded by this fungus.

Ustilago Zeae (Beckm.) Ung. Figs. 2, 55. The sori
break out on any part of the host, often forming prominent
smut balls, though these vary from a few mm. to over a dcm.

20 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

in diameter, and also vary in shape according to the part at-
tacked ; the brownish black spore mass is at first covered with
a whitish membrane, composed largely of semi-gelatinized fun-
gous threads. The spores are ellipsoidal to spherical, occa-
sionally irregular, prominently echinulate, and 8-ii/x, rarely
even 15/x, in length.

Hosts and Distr. : Eiichlama hixurians, New Haven, Sept.
11, 1903; Zea Mays, New Haven (Sturgis), Aug. 18, 1901
(Rorer) ; Southington, July 17, 1902; Westville, Aug. 14, Sept.
2, 1902, June 28, 1904; Hartford, Oct. 20, 1902; New Canaan,
Sept. 29, 1903.

Corn smut is common on both the sweet and field varieties,
though in this state the former is more subject to its attacks.
Seed treatment will not prevent the 'smut, as it can gain en-
trance to its host through any exposed young tissue. The
first host given is teosinte, a plant that is closely related to
corn. Fig. 55 shows smutted staminate blossoms of corn re-
duced to one-half size.

Ustilago Oxalidis Ell. & Tr. Figs. 3, 48. The incon-
spicuous sori are found in the seeds, all or part of these being
changed into reddish brown dusty spore masses, that show to
the exterior only on the dehiscence of the otherwise little modi-
fied ovaries. The spores are golden yellow, ovoid to spherical,
or rarely more elongated or irregular, coarsely verrucose, and
13-20/Li in length.

Host and Distr. : Oxalis stricta, Yalesville, July, 1902 ;
West Cornwall, July 18, 1902; New Haven, July, 1902; Whit-
neyville, Oct. 18, 1902, Sept. 9, 1903 ; Manchester, Oct. 2, 1903.

An inconspicuous conidial stage is also produced on the
surface of the anthers of the infected flowers. These temporary
spores are thin-walled and ovoid to subspherical. They are so
placed that they are probably carried from the flowers by in-
sects, as are pollen grains. Fig. 48 shows two ovaries in which
all of the seeds have been changed into smutty bodies.

Ustilago anomala Kze. The sori occur in the essential
organs of all the flowers, the floral envelopes forming a cov-
ering to the dusty, purplish spore mass. The spores are light
violet, ovoid to spherical, occasionally somewhat irregular,
provided with rather fine winged reticulations (1-3^ wide by
ifi deep), and 10-15/x, rarely 17^, in length.

No. 5.] USTILAGINEiE OF CONNECTICUT. 21

Hosts and Distr. : Polygonum Convolvulus (?), Mt. Car-
mel, May 8, 1904 (last year's flowers) ; Polygonum dume-
torum var. scandens, Montowese, Sept. 14, 1903 ; New Canaan,
Sept. 29, 1903.

This species is closely related to the next, but differs in
having usually lighter colored and somewhat larger spores,
with finer reticulations.

Ustilago utriculosa (Nees.) Tul. Figs. 4, 21, 54. The
sori destroy the essential organs of all of the flowers, and form
ovate, dusty, purplish spore masses 3-4 mm. in length, pro-
tected at first by the floral envelopes. The spores are violet
or purplish, chiefly subspherical or spherical, provided with
rather coarse, winged reticulations (2-4/i wide by about 1.5/z
deep), and chiefly 9-14^ in diameter.

Hosts and Distr. : Polygonum Hydropiper, Montowese,
Sept. 20, 1902 ; Polygonum hydro piperoides, Whitneyville, July
24, 1902; Polygonum lapathifolium, Westville, Sept. 2, 1902;
Polygonum Pennsylvanicum, Westville, Sept. 2, 1902; Ham-
den, Sept. 11, 1902; Hartford, Oct. 20, 1902; Glastonbury,
Oct. 23, 1902; Yalesville, Sept. 11, 1903, Oct. 14, 1904; Monto-
wese, Sept. 14, 1903; New Canaan, Sept. 29, 1903; Manches-
ter, Oct. 2, 1903 ; Green's Farms, Sept. 30, 1904.

This is one of the most common smuts of the state, espe-
cially on the last host.

Sphacelotheca DeBy.

The sori are usually found in the inflorescence (often con-
fined to the ovaries), are provided with a false membrane of
fungous cells that soon ruptures, disclosing a dusty spore mass
and a central columella composed chiefly of plant tissues. The
false membrane is formed largely or entirely of definite sterile
fungous cells which are hyaline or slightly tinted, and vary
in shape from linear to subspherical or cuboidal, and in size
from less than to larger than the spores. The spores are like
those of Ustilago, simple, free, usually reddish brown, and of
small to medium size. Figs. 5, 6, 37, 38.

Very often groups of the sterile subspherical cells are scat-
tered through the spore mass. The columella is usually the
remains of the woody plant tissues, and often protrudes above

22 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

the spore mass as the latter wears away. See Fig. 37. This
genus has not been thoroughly worked up for the whole world,
and so a number of the species belonging under it are now in-
cluded under Ustilago. Only two of the 16 species occurring
in North America are of economic importance. So far only
two species have been found in Connecticut.

Key to Species of Sphacelotheca.

I. Spores olive brown, smooth 5". Sorghi.

II. Spores purplish, verruculose 5". Hydropiperis.

Sphacelotheca Sorghi (Lk.) Clint. Figs. 5, 24, 26, 38.
The sori occur in the ovaries, forming oblong or ovate bodies
usually 3-8 mm. in length, or rarely fusing the aborted spike-
lets into longer forms. The brownish false membrane wears
away from the apex, revealing the olive brown spore mass and
finally the evident slender columella. The sterile cells of the
membrane easily break up into groups, and are hyaline, oblong
to subspherical, and chiefly 7-18/* in length. The olive brown
spores are subspherical or spherical, smooth, often have pitted
contents, and are 5.5-8.5/x in diameter.

Hosts and Distr. : Sorghum ■vulgare var. sorghum, New
Haven, Sept. 30, 1903 ; Sorghum vulgare var. technicum, New
Haven, Sept. 20, 1901 (Rorer), Sept. 11, 1903.

This species was found on both sorghum and broom-corn
grown at the Experiment Station. So far as I can learn,
neither of these plants is grown commercially in the state, so
the smut is of no economic importance here, though further
west it often does considerable injury to these plants.

Sphacelotheca Hydropiperis (Schum.) DeBy. Figs. 6,
37. The sori are found in the ovaries, forming oblong or
ovate bodies 3-5 mm. in length ; with the false membrane de-
hiscing at the apex, and revealing the purple-black spore mass
and finally the slender columella. The sterile cells, besides
forming the false membrane, constitute part of the columella;
they easily separate, are hyaline or slightly violet tinted, chiefly
subspherical, and 6-17/x in length. The spores are purplish,
broadly oblong or ovate to (chiefly) subspherical, very mi-
nutely but abundantly verruculose and 10-17^ in length.

Hosts and Distr.: Polygonum acre, Green's Farms, Sept.

No. 5.] USTILAGINE.E OF CONNECTICUT. 23

30, 1904; Polygonum sagittatum, Westville, Sept. 2, 1902,
Sept. 14, 1903 (Britton), Oct. 4, 1904; Montowese, Sept. 20,
1902; Whitneyville, Sept. 2iy 1902; Centreville, Sept. 27, 1902;
New Canaan, Oct. 9, 1902 (Britton), Sept. 11, 1903; Yales-
ville, Sept. 11, 1903; Andover, Sept. 15, 1903; Cheshire, Oct.

25, 1903-

This is another of the common smuts of the state, especially

on the arrow-leaf Polygonum, and is found in the fall. It is
the only purple-spored species of the genus, of which it is the
type, however. Fig. $7 shows the smut on Polygonum acre,
several of the sori exposing the slender central columella.

Cintractia Cornu.

The sori occur on various parts of the host, but most com-
monly in the ovaries ; at maturity they form a firmly aggluti-
nated, or more rarely a dusty, spore mass, which usually is pro-
tected at first by a false membrane of sterile threads or indefi-
nite cells. The single spores develop centripetally around a
central axis, the outermost wearing off as they ripen ; they are
usually of medium to large size, and of a dark reddish black
color, often opaque. Figs. 8, 27, 28.

The species of this genus occur on the Cyperaceas, or occa-
sionally on related families. So far 17 species have been de-
scribed, 13 occurring in North America, and 4 in Connecticut.
None of the species are of economic importance.

Key to Species of Cintractia.

I. Sori dusty at maturity, concealed by the glumes.

A. Spores usually irregular polyhedral C. Cyperi.

B. Spores usually ovoid to spherical (often with hya-

line wings) C. Montagnei.

II. Sori firmly agglutinated at maturity.

A. Subspherical, in ovaries C. Caricis.

B. Oblong to linear, surrounding base of pedicles

C. Junci.

Cintractia Cyperi Clint. The sori occur in the interior
of the spikelets, infecting all of the head, and are hidden by
the enveloping glumes until finally the dusty spore mass sheds

24 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

out on their exterior. The spores are reddish brown, oblong
to polyhedral, chiefly irregular, smooth, but often showing
darker lines on surface due to pressure of spore mass, and 12-
18/u, or (most elongated) even 22^, in length.

Host and Distr. : Cy perns iiliculmis, North Haven, July
26, 1902 ; Montowese, Sept. 14, 1903.

This species was described originally from this state by the
writer, having been found not uncommon on the sand plains
near North Haven and Montowese. The affected plants look
very much like those free from the smut, but can usually be
detected by the darker aspect of the spikelets, which in this
sedge are clustered into heads.

Cintractia Montagnei (Tul.) Magn. The sori occur
hidden in the ovaries, forming inconspicuous, oblong to sub-
spherical, usually dusty spore masses. The spores are brown
or brownish black, often compressed laterally, and so appear-
ing ovoid to subspherical or occasionally more irregular and
angled, smooth but minutely pitted, and 12719/z, chiefly 13-16/i,
in length ; they are very often provided with conspicuous, hya-
line, wing-like bladders on either side.

Host and Distr. : Rhynchospora alba, Berlin, Sept. 3,
1902; Cheshire, Oct. 25, 1903.

This is another smut readily overlooked, because the in-
conspicuous sori are hidden by the floral bracts.

Cintractia Caricis (Pers.) Magn. Figs. 8, 27. The sori
form subspherical bodies about 3-4 mm. in diameter in the
ovaries ; at first the sorus is protected by a white membrane
of sterile fungous tissue, but this soon wears off, revealing the
black, firmly agglutinated spore mass, the spores of which
gradually ripen and wear off toward the interior. The spores
arc black-brown, subopaque, chiefly irregular polyhedral, or
occasionally ovoid to subspherical, smooth or pitted to gran-
ular or even papillate, and 16-27^, chiefly 18-22/i, in length.

Host and Distr. : Carex Pennsylvania, Rainbow, June 6,
1903; East Hartford, June 1, 1904 (Weatherby).

This variable, smut has been reported on a large number of
Carex species in North America, but the above is the one upon
which it commonly occurs. The illustration shows it in the
ovaries at the base of the staminate spikelet of this host.

No. 5.] USTILAGINE.E OF CONNECTICUT. 2$

Cintractia Junci (Schw.) Trel. Fig. 28. The sori are
linear, usually surrounding the pedicles or peduncles for half
or more of their lower length, occasionally developing in the
basal parts of the flowers ; they form a black, rather firmly ag-
glutinated spore mass. The spores are black-brown, sub-
opaque, oblong to irregular polyhedral or subspherical, very
minutely pitted, and 1 4-22/1 in length.

Host and Distr. : Juncns tenuis, Westville, June, 1891
(Thaxter) ; Milford, June 26, 1894 (Sturgis), July 27, 1902;
Whitneyville, June 20, 1902.

This was one of the first smuts reported from North Amer-
ica, having been described by Schweinitz from Carolina in
1834. The figure shows one of the peduncles smutted for half
its length.

Schizonella Schrot.

The sori form black agglutinated spore masses in the
leaves. The spores are united in pairs (formed by internal
division of a mother cell), and often become laxly connected
by their bulging contiguous surfaces, or even entirely separ-
ated ; they are of reddish brown color and of small or medium
size. Figs. 9, 35.

Only one species, with a variety, is known for this genus,
but it has a wide distribution on various species of Carex.
In North America it has been found chiefly on Carex Pennsyl-
vania!, its host in this state.

Schizonella melanogramma (DC.) Schrot. Figs. 9, 35.
The sori form black, agglutinated, linear, elevated striae, 1 or
2 mm. long, or, by terminal fusion, of considerable length,
chiefly on the upper sides of the leaves. The spores are dark
reddish brown, often with the cells entirely or partially sep-
arated by the bulging out of their contiguous surfaces, chiefly
ellipsoidal to hemispherical, or, when separated entirely, poly-
hedral or subspherical, and 8-12/x in, length.

Host and Distr. : Carex Pennsylvanica, Westville, July
12, 1902 ; Whitneyville, May 4, 1903.

This is a species found chiefly in the spring; it probably
has a much wider distribution than indicated here.

26 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

Sorosporium Rud.

The sori occur in various parts of the host, forming dusty
dark colored spore masses like Ustilago. The medium-sized
spore balls are composed of numerous spores, often so loosely
held together that in time they separate entirely. The spores
are like those of Ustilago, simple, olive to reddish brown, and
of medium size. Figs. 7, 36.

When the spore balls become separated into the individual
spores, it is difficult to distinguish the species from Ustilago.
The numbe* of species described by Saccardo is over 30, but
some of these probably belong under other genera. For
North America 9 species are now known, and 2 or 3 of these
occur in Connecticut.

Key to Species of Sorosporium.

I. Sori in ovaries, 1-2 cm. in length ; spores 8-12^

S. Everhartii.
II. Sori involving the entire infloresence usually.

A. Sorus 1-5 cm. ; spores 12-19^ 5\ Ellisii.

B. Sorus usually yj cm. ; spores 9-13^

S. Syntherismce.

Sorosporium Everhartii Ell. & Gall. The sori develop
in the ovaries, forming linear bodies 1-2 cm. in length, and are
covered with a prominent whitish false membrane that de-
hisces at the apex into several lobes, disclosing the black-
brown, semi-agglutinated spore mass and the flattened colu-
mella of plant tissue. The spore balls are oblong to subspher-
ical, composed of many firmly agglutinated spores, and vary
from 55 to 125/i in length. The spores are reddish brown, or
the interior ones often nearly hyaline, ovoid to subspherical
or polyhedral, smooth (outermost rarely granular), and 8-12^
in length.

Host and Distr. : Andropogon scoparius, Westville, Oct.
22, 1903; [Southington, July 16, 1902; North Haven, July 26,
1902; Montowese, Sept. 20, 1902; New Haven, Oct. 18,

1903]-

The first specimen mentioned above is typical of the species

as described here. Those included in the brackets were origi-

No. 5.] USTILAGINE.E OF CONNECTICUT. 2j

nally considered by the writer as belonging under S. Ellisii,
since their sori involve the entire inflorescence, instead of
being limited to the ovaries. The spores, however, are
smaller than those of the typical members of that species, and
both the spores and spore balls are like 6". Everhartii, so
they may possibly be only vigorous specimens of this species,
in which the sorus has involved the whole inflorescence.

Sorosporium Syntherismas (Pk.) Farl. Figs. 7, 36. The
elongated sori usually involve the entire inflorescence, 3-7 cm.
in length, or more rarely are limited to the individual spikelets,
when they are shorter; they are provided with a prominent
false membrane which ruptures irregularly, disclosing the black-
brown spore mass, within which are often shredded filaments
of plant tissues. The sterile cells of the membrane are hya-
line, oblong to cubical or subspherical, and tend to adhere in
filaments when crushed apart. The spore balls, often evanes-
cent when old, are irregular oblong to subspherical, and 40-
100 fj in length. The reddish brown spores are minutely ver-
rucose (the inner often smooth and more lightly colored),
subspherical, polyhedral or occasionally more elongated,
chiefly 9-1 3n in length.

Host and Distr. : Cenchrus tribidoides, Savin Rock, Aug.
30, 1902 ; Montowese, Sept. 20, 1902.

This species should also be looked for on Panicum pro-
liferum, which is a common host for it elsewhere.

Tolyposporium Wor.

The sori are usually found in the ovaries, forming a gran-
ular spore mass at maturity. The spore balls are of medium
size, dark colored, and composed of numerous permanently
united spores. The spores are bound together by ridged folds
or thickenings of their outer walls, and are of small to me-
dium size. Figs. 10, 40.

The spore balls can be ruptured by pressure, when the
outer darker colored covering often breaks apart as ridges or
spine-like projections on the light colored or hyaline spores.
So far 13 species have been described, though it is doubtful if
all belong to this genus; 2 of these occur in North America,

28 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

and i in Connecticut. None of the species apparently are of
economic importance.

Tolyposporium bullatum (Schrot) Schrot. Figs. 10, 40.
The ovate sori are found in the ovaries, are about 3-5 mm. in
length, and are covered by a smooth greenish plant membrane,
which upon rupturing discloses the granular, black spore
mass. The spore balls are opaque, black, oblong to spherical
or polyhedral, contain numerous (over 100) firmly aggluti-
nated spores, and are 50-1 60/z in length. The spores are light
reddish brown, or the inner ones semi-hyaline, and are covered
with a thin, tinted outer coat thrown more or less into ridges
or folds that bind the spores together; they vary from ovoid
to spherical or polyhedral, and are 7-12^ rarely 14^ in length.

Host and Distr. : Panicum Crus-galli, Woodmont (Setch-
ell) ; Unionville, Aug. 26, 1902; New Haven, Sept. 11, 1903;
New Canaan, Sept. 29, 1903.

Sometimes Ustilago sphcrrogena occurs in the ovaries of
the same plant with this. It can easily be distinguished from
this by its larger sori, which are covered by a hispid mem-
brane. Figure 40 shows the uppermost ovary only infected by
the fungus.

TILLETIACE.SE Schrot.

The sori form dusty erumpent spore masses, or are per-
manently embedded in the plant tissues, often without evident
distortion of these. The germination is by means of a pro-
mycelium, which usually gives rise to a terminal cluster of
elongated sporidia, which sometimes bear whorls of similar
secondary sporidia; or the primary sporidia, with or without
fusing in pairs, may give rise to infection threads, or in a
nutrient medium to a mycelium bearing dissimilar secondary
sporidia (aerial conidia). Fig. 22.

Tilletia Tul.

The sori occur in various parts of the host, usually in the
ovaries, forming a dusty, dark spore mass. The spores are
simple, separate, and originate singly in the ends of special
mycelial threads that generally disappear rather completely

NO. 5.] USTILAGINE/E OF CONNECTICUT. 29

through gelatinization ; they are of large size, 16-35/i. Figs.
11, 12, 39.

This is the type of the family, and is one of the larger
genera. The sorus is quite like that of Ustilago, but the
spores are usually considerably larger. Saccardo records 53
species ; 1 5 have been found in North America, but only 2 in
Connecticut. The important stinking smuts of wheat, which
cause so much damage in the western wheat districts, belong
in this genus ; but, as wheat is little grown in this state, neither
species has been reported here. The smooth-spored species,
Tilletia f ceteris (B. & C.) Trek, however, has been found, on
microscopic examination, in ground cattle food offered for
sale in the state, apparently made from smutted western wheat.
The cattle did not relish this food.

Key to Species of Tilletia.

I. Spores reticulate T. Anthoxanthi.

II. Spores apparently verruculose T. Maclagaui.

Tilletia Anthoxanthi Blytt. Figs. 12, 39. The sori arc
ovate, about 3 mm. in length, and usually occur in all of the
ovaries of the spike ; they are somewhat hidden by the envel-
oping glumes, and at first are covered by a thin plant mem-
brane, which eventually ruptures, disclosing the dusty, reddish
black spore mass. Hyaline cells, or immature spores, are
mixed with the spores, and are smaller than those, and have
thin to medium-thick walls. The mature spores are reddish
brown, ovoid to spherical, reticulate (3-6^ wide and 1-3/z
deep), and 24-30/*, occasionally even 34/i, in length.

Host and Distr. : Anthoxanthum odoratum, Whitney-
ville, July 6, 1902.

The sori are so inconspicuous in the ovaries that the in-
fected spikes scarcely differ in appearance from the normal.
See Fig. 39, in which the spike is magnified two diameters.
This European species has been found in this country only at
the above station.

Tilletia Maclagani (Berk.) Clint. Fig. 11. The sori are
found in the ovaries, and rarely also in the anthers, and are in-
conspicuous, being concealed by the enveloping glumes ; upon

30 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

the rupture of the enclosing plant membrane, a dusty, red-
dish brown spore mass is disclosed. The spores usually show
various stages of development, vary from light to dark reddish
brown, are subspherical or spherical, or occasionally elon-
gated or irregular, have a thick wall (3-4/O apparently closely
covered with verruculations (really very minutely areolately
pitted), and are 18-27^ in length.

Host and Distr. : Panicum virgatum, Conn. (Herb. Far-
low) ; East Hartford, July 7, 1903 (Weatherby).

The wild grass which is host for this smut is often cut for
hay in the state, but it is doubtful if the smut does enough
damage to be of economic importance.

Neovossia Korn.

The sori occur in the ovaries, forming dusty spore masses
protected at first by a membrane of plant tissue. The large
spores are simple, separate, and produced singly in the swollen
ends of the special fertile hyphae, which permanently invest
them and taper into conspicuous elongated hyaline appen-
dages. Fig. 13.

This genus is closely related to Tilletia, but can be dis-
tinguished by the tail-like appendage to the membrane invest-
ing the spores. There are only two species now known that
properly come under this genus.

Neovossia lowensis Hume & Hods. Fig. 13. The sori
are found in the ovaries, are ovoid, about 2-3 mm. in length,
and show inconspicuously between the spreading glumes.
Sterile cells, or immature spores, which are hyaline, thick-
walled, and chiefly smaller than the spores, are found in the
spore mass. The mature spores are reddish brown, subo-
paque, ovoid, ellipsoidal, or rarely subspherical, provided with
a hyaline envelope tapering into an irregular tail once or twice1
their length. They have a minutely reticulately pitted cell wall,
and are 19-28/M by 13-19/u in size.

Host and Distr. : Phragmites communis, Montowese, fall,
1901 (Evans) ; Oct. 18, 1902.

This rare species, which has been reported elsewhere only
from Iowa, was first found in this state by Professor Evans
of Yale.

No. 5.] USTILAGINE^E OF CONNECTICUT. 3 1

Urocystis Rab.

The sori usually occur in the leaves or stems, where they
often cause considerable distortion, or more rarely in the
floral parts, and form dark colored, dusty spore masses. The
spore balls are permanent, composed of an enveloping cortex
of tinted sterile cells, enclosing one to several spores, and are
of small to medium size. The spores are reddish brown, and
of variable shape and size. Figs. 14, 15, 41, 42.

The sori of this genus look very much like those of Usti-
lago. Saccardo in his Sylloge Fungorum describes 34 spe-
cies; 12 of these occur in North America, and 4 have been
listed from this state. A few of the species are of some
economic importance.

Key to Species of Urocystis.

I. Spore balls usually with 1 fertile cell, rarely with 2 ;
sori forming extended outbreaks on leaves and

bulbs U. Cepulce.

II. Spore balls usually with 1 or 2 fertile cells, rarely
with 3 or 4.

A. Sori forming pustular or irregular swellings

on leaves and stem U. Anemones.

B. Sori in striae usually on under side of leaf

sheath U. occulta.

III. Spore balls usually with 3 to 5 fertile cells, rarely

with more ; sori oblong in base of flowers

U. Hy poxy is.

Urocystis Cepulee Frost. Figs. 14, 42. The sori occur
in the leaves and bulbs as isolated pustules or often as more
extended areas, and are at first covered by a thin plant mem-
brane, but eventually rupture this, disclosing a black, dusty
spore mass. The spore balls are ovoid to spherical, small,
17-25^ in length, and contain one or rarely two spores. The
sterile cells are tinted, ovoid to spherical, small, 4-8^, and
rather completely cover the spores. The spores are reddish
brown, ovoid to spherical, and chiefly \2-i6fi in length.

Host and Distr. : Allium Cepa, Green's Farms, June,

32 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

1890 (Thaxter), Oct. 22, 1902, June 24, 1904; Bridgeport,
Sept., 1902 ; New Haven, June 23, 1904.

This is one of the most injurious smuts in this state. It
occurs early in the season on the young seedlings, many of
which may be killed outright, thus thinning out the stand ir-
regularly. It is also found throughout the season, eventually
doing considerable damage to the bulbs. The smut often be-
comes established in the soil, which is then rendered unfit for
raising onions from seed. Sets, however, transplanted in the
infected soil, suffer little from this trouble. Soil once infected
remains so for years, and for this reason there is considerable
land in the onion districts of the state that cannot be used to
advantage for this crop. Experiments with treating the in-
fected soil, at time of planting the seed, with various fungi-
cides, such as formalin, and sulphur mixed with lime, have
given some favorable results in keeping down the smut. As
yet, however, such treatment is little practiced by the growers.

Urocystis Anemones (Pers.) Wint. The sori develop
in the leaf blades, petioles, and stems, forming conspicuous
pustules of varying shape and size, and soon disclose dusty,
black spore masses. The spore balls are irregular, contain
1-5, usually 1-2 spores, and are 22-35/x, rarely 45/x, in length.
The sterile cells usually incompletely cover the spores, and
sometimes become separated from them ; they are smoky
brown or yellowish tinted, ovoid to spherical, and about 8-14/i
in length. The spores are reddish brown, irregular, oblong
or ovoid to polyhedral or subspherical, smooth, and chiefly
\2-1Jn, rarely 20/i, in length.

Host and Distr. : Anemone nemorosa, Milford, May,

1 89 1 (Thaxter).

This smut is found early in the spring, and no doubt has a
wider distribution and other hosts in the family Ranunculaceae
than that reported here.

Urocystis occulta (Wallr.) Rab. Figs. 15, 41. The
sori of this smut are usually confined to the leaves, especially
the inner side of the leaf sheaths, but occasionally occur in
the culm or inflorescence ; they form linear striae of consider-
able length, and often are so closely placed that they merge
into a dusty, reddish black stratum. The spore balls are ob-

No. 5.] USTILAGINE^ OF CONNECTICUT. 33

long to subspherical, contain I or 2, rarely 3 or 4, spores, and
are 16-32/z in length. The sterile cells usually incompletely
cover the spores, are hyaline or yellowish tinted, and sub-
spherical to oblong. The spores are reddish brown, oblong
to subspherical, often irregularly flattened, smooth, and 11-18^
in length.

Host and Distr. : Secale cereal e, Milford, June, 1890
(Thaxter) ; Rainbow, June 6, 1903.

The smut possesses some economic importance, since it
occurs on cultivated rye. However, it apparently rarely be-
comes so abundant in the rye fields of this state as to cause
serious injury. So far the writer has been able to find only
a single smutted plant, though a number of fields have been
examined.

Urocystis Hypoxyis Thaxt. The sori occur in the inner
and basal parts of the flowers, distorting and destroying them
more or less, and often run down somewhat on the pedicles ;
the infected parts have an irregular oblong shape, and finally
disclose the dusty, purple-black spore mass. The spore balls,
vary from ovoid to spherical, contain 1-8, chiefly 3-5, spores,
and are 25-60/x in length. The sterile cells completely cover
the spores, are reddish yellow, ovoid, and 9-14/u in length.
The spores are reddish brown, polyhedral, subspherical, or oc-
casionally ovoid to oblong, and usually 13-16/x in length.

Host and Distr. : Hypoxys ere eta, Westville, July, 1889
(Thaxter), July 12, 1902.

This smut was originally described from the specimens
found by Professor Thaxter of Harvard in West Rock Park,
Westville. Since then it has been reported from Massa-
chusetts and South America.

Entyloma DeBy.

The sori occur chiefly in the leaves, being permanently em-
bedded in their tissues, and forming discolored but usually
scarcely distorted areas. The spores are hyaline or yellowish
tinted, rarely darker colored, simple, separate, or occasionally
slightly adhering together in rows or irregular masses, and of
medium size. Elongated conidia are often produced from the

34 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

mycelium, protruding in tufts through the stomates. Figs.
16-18, 30-34.

About 70 species have been described, of which 25 occur
in North America. Of the latter, four occur on cultivated
hosts, but are not of especial economic importance. So far 8
species have been reported in Connecticut.

Key to Species of Entyloma.

I. Spores dark tinted, reddish brown, often, adhering to-
gether.
A. Sori forming minute, black, oblong or linear striae.

1. Spores golden brown, 7-1 ifx E. lineatum.

2. Spores darker brown, 8-14^... E. crastophilum.
II. Spores hyaline or yellowish tinted; sori forming discol-
ored spots.

A. Spores not apiculate, often adhering somewhat.

1. With hypophyllous conidia or sporidia.

a. Sori yellowish or reddish brown, 1-2 mm.

E. Thalictri.

b. Sori whitish or yellowish, 1-10 mm

E. Lobelice.

c. Sori yellowish or reddish, often bordered,

y2-6 mm E. Physalidis.

2. Usually without conidia.

a. Sori light yellowish, y2-2 mm. . .E. Linaria.

b. Sori yellowish or reddish brown, 2-5 mm. . .

E. polysporum.

B. Spores apiculate and pedicellate, never adhering ; sori

yellowish, later reddish brown E. Nymphaa.

Entyloma lineatum (Cke.) Davis. Figs. 16, 32. The
sori occur in the leaves, leaf sheaths, or occasionally in the
culm, and are small, ^-3 mm. in length, subcircular to
(chiefly) linear, scattered or fusing somewhat, lead-colored
or black, and rather permanently covered by the epidermis.
The spores are light golden brown, firmly agglutinated, usually
ovoid to subspherical or somewhat polyhedral, smooth, and
7-1 ifi in length.

No. 5.] USTILAGINE/E OF CONNECTICUT. 35

Host and Distr. : Zizania aquatic a, New Haven, 1892
(Setchell) ; Whitneyville, July 24, 1902; Montowese, Oct. 18,
1902; Westville, Aug. 21, 1903.

Entyloma crastophilum Sacc. This species is closely re-
lated to the preceding, but differs in its slightly smaller sori
and somewhat darker colored spores which are 8-14/i in length.

Host and Distr. : Holcus lanatus, West Haven, July 2^,
1903; Undetermined grass, West Haven, Aug. 12, 1903.

Both this and the preceding species differ from the ordinary
Entylomas in their darker colored spores.

Entyloma Thalictri Schrot. The sori are found in the
leaves, forming small, angular, yellowish or reddish spots
about 1-2 mm. in diameter, or by confluence becoming more
extended and indefinite. The spores are hyaline or yellowish,
ovoid to subspherical or occasionally somewhat angled and
more irregular, thin-walled to medium thick-walled, smooth,
and 8-13/i in length. The conidia are apparently produced on
the under surface of the sori.

Host and Distr. : Thalictrum polygamum, Montowese,
Sept. 14, 1903.

This species is not very common, and has been reported on
the above host only from this state.

Entyloma Lobeliae Farl. Figs. 17, 31. The sori form
conspicuous whitish or yellowish areas in the leaves, and are
1-10 mm. in diameter. The spores are hyaline or yellowish
tinted, oblong to subspherical or somewhat angled, rather
thick-walled, and 11-15/* in length. The conidia form a whit-
ish growth on the under surface of the sori, and are fusiform,
and 10-25^ by 2-3^ in size.

Host and Distr.: Lobelia inflata, Hamden, Sept. 11, 1902;
Hartford, Oct. 20, 1902 ; Cheshire, Aug. 3, 1903 ; Yalesville,
Oct. 14, 1904.

This is probably the most common Entyloma occurring in
the New England states, but so far it has been found only on
the above species of Lobelia.

Entyloma Physalidis (K. & C.) Wint. Fig. 34. The
sori form at first yellowish, but later darker colored, roundish
or angular spots from 3/2-6 mm. in diameter. The spores vary
from slightly tinted to light or reddish yellow, are ovoid, spher-

36 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

ical or slightly angled, have a thickish wall, and are 10-16^ in
length. The conidia may occur on either surface of the sorus
as a whitish growth, and are linear, somewhat curved, and
30-55/i by 1-2/i in size.

Host and Distr. : Physalis pitbescens, South Manchester
(Thaxter) ; New Haven, Sept. 16, 1902, Sept. 30, 1903 ;
Physalis Virginiana, Hamden, Oct. 14, 1903.

The cultivated strawberry tomato, Physalis pitbescens, is
sometimes rather seriously injured by this smut. The figure
shows the sori on this host, the spots having a definite darker
border.

Entyloma Linariae Schrot. Fig. 30. The sori show as
small, faint, yellowish spots on the under surface of the leaves,
and are oval to circular in outline, and l/>-2 mm. in diameter.
The hyaline or yellowish spores are chiefly subspherical or
spherical, smooth, have evident double wall, and are ii-i$h,
in length. The conidia have not been observed.

Host and Distr. : Linaria vulgaris, Westville, Oct. 22,
1903.

This species has been reported on this host only twice in
North America. The variety is more common.

Entyloma Linariae var. Veronicas Wint. The sori are
somewhat more evident than in the typical form of the species,
showing on both sides of the leaf. The spores also are deeper
tinted and larger, 13-16/z, or rarely 19^, in length.

Host and Distr. : Veronica peregrina, New Haven, May
16, 1904.

Entyloma polysporum (Pk.) Farl. The sori form sub-
circular or more irregular, yellowish or (later) dark brown
spots, 2-5 mm., in the leaves; the surrounding tissue is often
killed, thereby merging the sori. The spores are usually hya-
line to yellowish, or more rarely even chestnut brown, ovoid
to spherical or somewhat polyhedral, smooth, provided with
evident thick double wall, and chiefly 12-17/u in length. Coni-
dia, apparently, are lacking.

Host and Distr. : Ambrosia artemisiccfolia, Cheshire, Aug.

3. I903-

The closely related species Entyloma Compositarum Farl.,

also on Compositas as hosts, probably also occurs in this state,

though not yet reported.

No. 5.] USTILAGINE^ OF CONNECTICUT. 37

Entyloma Nymphaeae (Cunn.) Setch. Figs. 18, 33. The
sori occur in the leaves, forming yellowish, or with age reddish
brown, variable, usually irregular areas most prominent on
the under surface. The hyaline spores are ovoid to subspheri-
cal, usually distinctly apiculate, and with remains of hypha as
an appendage at opposite end, smooth or, under an immersion
lens, very minutely verruculose, 10-14^ in length. Conidia
have not been found, but the spores germinate in situ.

Hosts and Distr. : Nuphar advena, New Haven (Setch-
ell) ; Nymphcea odorata, Ledyard (Setchell) ; Westville, Sept.
2, 1902, Oct. 3, 1904.

The appendaged and apiculate spores easily distinguish
this species from any other. Fig. 33 shows the sori on a por-
tion of the leaf of Nymphcea odorata.

Doassansia Cornu.

The sori occur on various parts of the host, usually in the
leaves, and are rather permanently embedded in the tissues.
The spores are united into large, permanent spore balls ; these
consist of a distinct cortical layer of sterile cells, with the spores
entirely filling the interior, or limited to one or two layers be-
neath the cortex, while the interior is filled with sterile cells
or threads. The spores are hyaline or yellowish tinted, smooth,
and of small to medium size. Figs. 19, 22, 29.

Our knowledge of this and the next genus is largely due
to the investigations of Professor Setchell of the University
of California, but at one time a resident of this state, where he
collected part of the material with which he worked. The
species occur chiefly on plants growing in moist situations, es-
pecially on the Alismacese. Of the 24 species described, 10
occur in North America, and 5 in Connecticut. They are of
no economic importance.

Key to Species of Doassansia.

I. Spore balls with spores entirely filling their interior;
sori forming lead-colored, slightly elevated pustules

in leaves D. opaca.

II. Spore balls with a single layer of spores within which
are sterile cells.

38 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

A. Sori in the slightly swollen ovaries D. occulta.

B. Sori forming discolored areas in leaves

D. MartianoMana.

C. Sori forming conspicuous distortions usually in

leaves D. deformans.

III. Spore balls with several irregular layers of spores, within
which are hyphal threads ; sori indefinite, hidden in
leaf petioles D. obscura.

Doassansia opaca Setch. Figs. 19, 22, 29. The sori
form in the leaves evident lead-colored pustules, about 2-5
mm., which show as slight elevations on both surfaces; they
are more or less scattered, usually in yellowish or reddish
brown widely discolored areas. The spore balls are closely
compacted in a single layer, occupying most of the space be-
tween the epidermal layers of the leaf, and consist of a distinct
cortex with a mass of fertile cells entirely filling the interior;
they are oblong to subspherical or cuboidal, and 200-300/i in
greatest length. The cortical cells are reddish brown, oblong
or subcubical, prominent, about 14-27/u in length. The spores
are rather loosely compacted, chiefly subspherical, 10-15^ in
diameter.

Host and Distr. : Sagittaria variabilis, Norwich, Aug. 14,
1889 (Setchell); New Haven (Setchell) ; Whitneyville, Aug.
10, 1902 ; Montowese, Sept. 20, 1902 ; Westville, Sept. 8, 1904.

A portion of one of the spore balls is shown in cross section
in Fig. 19.

Doassansia occulta (Hoffm.) Cornu. The sori occur in
the ovaries, forming swollen, ovate bodies. The spore balls
consist of a distinct cortex within which is a single layer of
fertile cells surrounding a central mass of parenchymatous tis-
sue; they are subspherical or more irregular, and 100-160^ in
length. The cortical cells are polyhedral or more elongated
tangentially, and 8-10^ in length. The spores adhere rather
firmly, and are about 10-12^ in length.

Host and Distr. : Potamogeton Pennsylvanicus, Norwich,
Aug. 26, 1889 (Setchell); Bridgeport (Morong).

Doassansia Martianoffiana (Thiim.) Schrot. In this
species the sori occur in the leaves, forming unthickened, yel-

No. 5.] USTILAGINE.E OF CONNECTICUT. 39

lowish or eventually reddish brown subcircular spots, which
sometimes merge into indefinite areas. The spore balls are
situated in the spongy parenchyma of the leaf, and consist of
a distinct cortex surrounding a single layer of fertile cells
within which is a central mass of sterile cells ; they are chiefly
subspherical or spherical, and 100-160^ in diameter. The
cortical cells are brown and small. The spores are slightly
yellowish tinted, chiefly polyhedral or slightly elongated ra-
dially, and S-12/j. in length.

Host and Distr. : Potamogeton sps., Norwich (Setchell) ;
New Haven (Setchell) ; Simsbury (Setchell) ; Whitney ville,
Aug. 18, Sept. 2i, 1902; Westville, Sept. 8, 1904.

Doassansia deformans Setch. The sori occur in various
parts of the host, usually in the leaves, where they form con-
spicuous distortions in the midribs and petioles. The spore
balls occur in the intercellular spaces, and consist of a cortical
layer surrounding a single layer of fertile cells within which
is a central mass of sterile cells ; they are chiefly subspherical,
and vary from 100-140/u in diameter. The cortical cells are
polyhedral, occasionally slightly elongated tangentially, small,
about 4-6^ by 8-12/z. The spores are ovoid to polyhedral,
rather firmly united, and chiefly 8-12/x, rarely 15/i, in length.

Host and Distr.: Sagittaria variabilis, Norwich, Aug. 17,
1889 (Setchell) ; New Haven (Setchell).

This species was originally described by Setchell from
Connecticut material.

Doassansia obscura Setch. The sori are obscured, the
spore balls being hidden in the interior of the basal parts of
the petioles without especial discoloration or distortion. The
spore balls are arranged in a single row in the air chambers
of the host, and consist of a distinct cortex surrounding several
irregular layers of spores and a central mass of indefinite fun-
gal hyphse; they are oblong to subspherical, and of very large
size, 150-300/i in length. The conspicuous cortical cells are
light brown, ovoid to obovate or subcordate, and 12-18/i by
8-i2^u. The spores are chiefly subspherical and small, 8-12/j.
in diameter.

Host and Distr. : Sagittaria variabilis, Norwich, Sept.
(Setchell).

40 CONNECTICUT GEOL. AND NAT. HIST. SURVEY. [Bull.

Setchell described this species originally from Connecticut
and Massachusetts. This is the only collection reported for
this state; the fungus is so hidden in the host that it is not
easily detected.

Tracya Syd.

The sori occur permanently embedded in the tissue of the
leaves (fronds). The spore balls are destitute of a cortical
layer, and consist of a single layer of spores enclosing a net-
work of septate filaments. The spores are hyaline or yellow-
ish, firmly united, and of small to medium size. Fig. 20.

This genus, first described by Setchell under the name
Cornuella, contains a single species, which has been reported
only from North America and on a single species of duckweed.
It is closely related to Doassansia, but lacks the sterile cortex.

Tracya Lemnae (Setch) Syd. Fig. 20. The sori occur
in the languishing fronds, showing the spore balls under a
hand lens as minute, clustered or scattered, opaque embedded
bodies. The spore balls are situated in the spongy paren-
chyma above the lower epidermis, are subspherical and rather
small, 50-100/i in diameter. The spores are yellowish, firmly
compacted, cuboidal, polyhedral or often more elongated ra-
dially, and chiefly 10-12/x in length; they arise from the ends
of the sterile network of brownish hyphse that fill the interior.

Host and Distr. : Spirodela polyrrhiza, New Haven
(Setchell) ; Whitneyville, Oct., 1902.

This is one of the most interesting species of the Ustila-
gineae. So far it has been reported only from four states. As
in Doassansia, sections of the infected tissue are necessary to
make out the structure of the fungus. Fig. 20 shows merely
a portion of the spore ball in cross section.

LIST OF HOSTS, ACCORDING TO FAMILIES.

MONOCOTYLS.

Naiadace,e. Alismace.e.

Potamogeton Pennsylvanicus. Sagittaria variabilis.

Doassansia occulta. Doassansia deformans.

Potamogeton sps. Doassansia obscura.

Doassansia Martianofhana. Doassansia opaca.

No. 5.]

USTILAGINE/E OF CONNECTICUT.

41

'GRAMINE.E.

Agrostis alba var. vulgaris.

Ustilago stritzformis.
Andropogon scoparius.

Sorosporium ? Ellisii.

Sorosporium Everhartii.
Anthoxanthum odoratum.

Tilletia Anthoxanthi.
Arrhenatherum avenaceum.

Ustilago perennans.
Avena sativa.

Ustilago Avence.

Ustilago levis.
Cenchrus tribuloides.

Sorosporium Syntherisma.
Eragrostis major.

Ustilago spermophora.
Euchlaena luxurians.

Ustilago Zees.
Glyceria grandis.

Ustilago longissima.
Holcus lanatus.

Entyloma crastophilum.
Hordeum vulgare.

Ustilago Hordei.

Ustilago nuda.
Panicum Crus-galli.

Tolyposporium bullatum.

Ustilago Crus-galli.

Ustilago sphcerogena.
Panicum sanguinale.

Ustilago Rabenhorstiaua.
Panicum virgatum.

Tilletia Maclagani.
Phleum pratense.

Ustilago striceformis.
Phragmites communis.

Neovossia Iowensis.
Secale cereale.

Urocystis occulta.
Setaria glauca.

Ustilago Panici-glauci.
Setaria Italica.

Ustilago Crameri.
Sorghum vulgare and vars.

Sphacelotheca Sorghi.

Zea Mays.

Ustilago Zea.
Zizania aquatica.
Entyloma lineatum.
Cyperace.e.

Carex Pennsylvanica.
Cintractia Caricis.
Schizonella melano gramma.
Cyperus filiculmis.
Cintractia Cyperi.
Rhynchospora alba.
Cintractia Montagnei.
Lemnace.e.

Spirodela polyrrhiza.
Tracya Lemnce.
Eriocaulace^:.
Eriocaulon septangulare.
Ustilago Eriocauli.
Jojcace.*:.

Juncus tenuis.
Cintractia J unci.
Liliace.*:.

Allium Cepa.
Urocystis Ce pules.

AM ARYLLIDACE.E .

Hypoxys erecta.

Urocystis Hypoxyis.

DICOTYLS.

POLYGONACE.E.

Polygonum acre.

Sphacelotheca Hydropiperis.
Polygonum Convolvulus (?)

Ustilago anomala.
Polygonum dumetorum
var. scandens.

Ustilago anomala.
Polygonum Hydropiper.

Ustilago utriculosa.
Polygonum hydropiperoides.

Ustilago utriculosa.
Polygonum lapathifolium.

Ustilago utriculosa.
Polygonum Pennsylvanicum.

Ustilago utriculosa.

^
^

42

CONNECTICUT GEOL. AND NAT. HIST. SURVEY.

[Bull.

Polygonum sagittatum.

Sphacelotheca Hydropiperis.
Nymph.eace.e.
Nuphar advena.

Entyloma Nymphcece.
Nymphaea odorata.
Entyloma Nymphcece.
Ranunculace^e.

Anemone nemorosa.

Urocystis Anemones.
Thalictrum polygamum.
Entyloma Thalictri.

OxALIDACE.E.

Oxalis stricta.

Ustilago Oxalidis.
Solanace^:.
Physalis pubescens.

Entyloma Physalidis.
Physalis Virginiana.
Entyloma Physalidis.
Scrophulariaceve.
Linaria vulgaris.

Entyloma Linarice.
Veronica peregrina.
Entyloma Linarice
var. Veronicce.
Campanulace^e.
Lobelia inflata.
Entyloma Lobelice.
Composite.

Ambrosia artemisiaefolia,
Entyloma polysporum.

EXPLANATION OF FIGURES.

Figs 1-20. Spores of various smuts, part of which are

shown in optical cross section. Magnified about 650 diameters

except where otherwise stated.

Fig. 1. Ustilago longissinia from Glyceria grandis.

Fig. 2. Ustilago Zece from Zea Mays.

Fig. 3. Ustilago Oxalidis from Oxalis stricta.

Fig. 4. Ustilago utriculosa from Polygonum Pennsylvanicum,

Fig. 5. Sphacelotheca Sorghi from Sorghum vulgare ; a,
sterile cells of false membrane.

Fig. 6. Sphacelotheca Hydropiperis from Polygonum sagit-
tatum; a, sterile cells of false membrane.

Fig. 7 Sorosporium Syntherismce from Cenchrus tribuloides;
a, spore ball magnified only about 100 diameters.

Fig. 8. Cintractia Caricis from Carex Pennsylvanica.

Fig. 9. Schiconella melanogramma from Carex Pennsyl-
vanica.

Fig. 10. Tolyposporium bullatum from Panicum Crus-galli;
a, spore ball magnified only about 100 diameters.

Fig. 11. Tilletia Maclagani from Panicum virgatum.

Fig. 12. Tilletia Anthoxanthi from Anthoxanthum odora-
tum; a, sterile cell.

Fig. 13. Neovossia Iozvensis from Phragmites communis.

1

No. 5.] USTILAGINE.E OF CONNECTICUT. 43

Fig. 14. Urocystis Cepuloe from Allium Cepa.

Fig. 15. Urocystis occulta from Secale cercale.

Fig. 16. Entyloma lineatum from Zizania aquatica.

Fig. 17. Entyloma Lobelia from Lobelia inflata.

Fig. 18. Entyloma Nymphcece from Nymphcca odorata.

Fig. 19. Doassansia opaca from Sagittaria variabilis. This

is a cross section of a spore ball merely showing one end ;

a, superficial cortex of sterile cells ; b, spores filling the

interior of ball.
Fig. 20. Tracya Lemnaz from Spirodela polyrrhiza. This is

a cross section of a spore ball merely showing one end ;

a, superficial layer of spores ;. b, sterile hyphae filling in-
terior of ball.

Figs. 21-25. Germination of spores of smuts. Magnified
about 650 diameters.

Fig. 21. Germination of spore of Ustilago utriculosa from
Polygonum Pcnnsylvanicum in water; a, promycelium;

b, sporidia.

Fig. 22. Germination of spore of Doassansia opaca from Sag-
ittaria variabilis in water ; a, promycelium ; b, sporidia ;

c, secondary sporidia fallen off.

Fig. 23. Germination of spore of Ustilago Avence from Ave-

na sativa in horse dung; a, promycelium; b, infection

threads.
Fig. 24. Germination of a sporidium of Ustilago Sorghi into

an infection thread.
Fig. 25. Small portion of a group of sporidia developed from

promycelium of Tolyposporium Eriocauli in potato agar.
Fig. 26. Cross section of epicotyl of broom corn infected
by Ustilago Sorghi; a, epidermal cells ; b, parenchyma cells of
cortex ; c, woody cells of central cylinder, and d, mycelium of
the fungus ramifying through the parenchyma cells of the
cortex. Magnified about 125 times.

Figs. 27-55. Sori of various smuts, showing general ap-
pearance and parts of different hosts infected. Natural size
except where otherwise indicated.

Index.

Page.

Cintractia Cornu 23

Caricis (Pers.) Magn.... 24

Cyperi Clint 23

Junci (Schw.) Trel 25

Montagnei (Tul.) Magn... 24

Doassansia Cornu 37

deformans Setch 39

Martianoffiana (Thiim.)

Schrot 38

obscura Setch 39

occulta (H off in.) Cornu. . 38

opaca Setch 38

Entyloma DeBy 33

crastophilum Sacc 35

Linariae Schrot 36

Linariae var. Veronicas

Wint 36

lineatum (Cke.) Davis.... 34

Lobelias Farl 35

Nymphaeae (Cunn.) Setch. 37

Physalidis (K. & C.) Wint. 35

polysporum (Pk.) Farl... 36

Thalictri Schrot 35

Neovossia Korn 30

Iowensis Hume & Hods. . 30

Schizonella Schrot 25

melanogramma (DC.)

Schrot 25

Sorosporium Rud 26

Everhartii Ell. & Gall 26

Syntherismaj (Pk.) Farl.. 27

Sphacelotheca DeBy 21

Hydropiperis (Schum.)

DeBy 22

Sorghi (Lk.) Clint 22

Page.

Tilletia Tul 28

Anthoxanthi Blytt 29

Maclagani (Berk.) Clint.. 29

Tilletiaceae Schrot 28

Tolyposporium Wor 27

bullatum (Schrot.) Schrot. 28

Tracya Syd 40

Lemnse (Setch.) Syd 40

Urocystis Rab 31

Anemones (Pers.) Wint.. 32

Cepulae Frost 31

Hypoxyis Thaxt 33

occulta (Wallr.) Rab 32

Ustilaginaceae Schrot 13

Ustilago Rouss 13

anomala Kze 20

A venae (Pers.) Jens 16

Craineri Korn 15

Crus-galli Tr. & Earle . . 19

Eriocauli (Mass.) Clint. . 18
Hordei (Pers.) Kell. &

Szu 15

levis (Kell. & Sw.) Magn. 15

longissima (Sow.) Tul... 15

nuda (Jens.) Kell. & Silk. 16

Oxalidis Ell. & Tr 20

Panici-glauci (Wallr.)

Wint 17

perennans Rostr 16

Rabenhorstiana Kiihn 17

spermophora B. & C 18

sphserogena Burr 18

striaeformis (West.) Niessl. 19

utriculosa (Nees.) Tul.... 21

Zese (Beckm.) Ung 19

Fig. 27, p. 24.

Pig. 28, p. 25.

Cinliai.ua Caricis ■ 2.

Cinlractia Junci

Fig. 29, p. 38.

Fig. 3..,, p

Duassansia opaea.

Enlyloma Linariae.

Fig. 31, p- 35-

Fig. 32, p. 34.

Kntvloma Lobelia-

Fig. 33, p

Entyloma lineatum.

Fig. 34, p. 35-

Entyloma Nytnphaeae.

Entvloma Physalidis.

Fig- 35, P- 25.

Fig. 36, p. 27.

Schizonella melanogramma
Fig. !7, p. 22

s sporiutn Syntherismse.

Fig. j8, p. :j.

Sphacelotheca Hydropiperis < 2.

Sphacelotheca Sorglii \ 2.

Fig- 39. P- 29-

Fig-. 40, p. 28.

Tilletia Anthoxanthi • 2.
Fig. 41, p

I

Tolyposporium bullatum

Fig. 42, p 31.

Urocystis occulta.

Urocystis Cepulse.

Pig. 43. P- '6.

Fig. 44, p. 1 -

Pig. 45, p. I;

Ustilagi 1 A vi 1 1
Fig. 46, p. 15.

Usl ilagi 1 Crameri.
Fig. 47. p. i6.

Ustilago Hordei.

Ustilago Oxalidis

Ustilago nuda.

Pig. w, p. ■?•

Fig. 50, p. 1?.

Pig. 51, p. tq.

U. Paniei-glauci.

Pig- 53. P- 18.

U. liabenhorstiana.

Fig. 52, p. 18.

Qstilago spermophora

Fig. 5.1, p. 2r.

U. striaeformis.

Fig. ,,, p. 19.

U. sphaerogena.

U. utriculosa.

U. Zea_- -h ,'a