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THE BRITISH RUST FUNGI
(UREDINALES)

CAMBRIDGE UNIVERSITY PRESS °
Ponder: FETTER LANE, E.C.
C. F. CLAY, Manacrer *

Evinburgh: 100, PRINCES STREET
London: WILLIAM WESLEY AND SON, 28, ESSEX STREET, STRAND
Berlin: A. ASHER AND CO.
Deipsig: F. A. BROCKHAUS
Pew Work: G. P. PUTNAM’S SONS
Gombay and Calcutta: MACMILLAN AND CO., Ltn.

All rights reserved

THE BRITISH RUST FUNGI
(UREDINALES)

THEIR BIOLOGY AND CLASSIFICATION

BY

W. B. GROVE, M.A.

Cambridge :
at the University Press
1913

'
Cambridge :
PRINTED BY JOHN CLAY, M.A.
AT THE UNIVERSITY PRESS

PREFACE

te is now twenty-four years since the publication of Plowright’s

“Monograph of the Uredinez and Ustilaginee,” and during
that long period very great progress has been made in elucidating
the biology of the former group. To Plowright will ever belong
the honour of being one of the pioneers in this important work,
especially in that branch which is the most fascinating and
has the greatest number of secrets to unfold—viz. Hetercecism.
But since his time numerous investigators have followed in his
footsteps, while others have taken new ground and largely
increased our knowledge, so that the picture which he presented
of the biology and classification of the Rusts has now become,
in certain directions, very incomplete and misleading.

The descriptions in the present volume are naturally based
upon those in the “ Monographia Uredinearum ” of the brothers
Sydow, so far as that monumental work has been published.
Those of all the species of which British specimens could be
procured have been carefully revised, and there is hardly one
of them that has not been added to or amended. Fischer’s
“Uredineen der Schweiz,” and McAlpine’s “ Rusts of Australia”
have also been found extremely useful. No attempt has been
made to give a full synonymy, but merely so much as was
required to show the origin and authority of the name used,
and to include all the references to the various species contained
in the works consulted, especially those of Cooke and Plowright.
Dates have been added occasionally, but only for special reasons
or if the nomenclature adopted differs from that in the

a3

vi PREFACE

“ Monographia,” where a complete synonymy, with dates, lies
at the disposal of those who are interested in that matter.

A plan has been adopted, in a few cases, of grouping
some forms that are closely allied under a common name;
see Puccinia dispersa and P. sessilis, With more knowledge
(or more courage) this process might have been carried much
farther, and it is believed that in this device will be found the
first line of defence of sane systematists against the excessive
multiplication of “species” by “biological” nomenclators. It
cannot be justifiable to use the same word in the same branch
of science to denote two widely diverse grades of evolution.
Physiological, unaccompanied by morphological, distinctions
should never be allowed to constitute a difference of species,
unless it be as a temporary measure in cases which have not
been investigated.

In selecting the name for each species, the principle of
priority has been followed, subject to two conditions—(1) names
given to varieties need not be adopted (International Rules,
1905, Art. 49), and (2) names given to imperfect states are
not to be preferred, but the earliest name given to the perfect
(in this case, the teleutospore) stage (Brussels Congress, 1910);
sometimes, however, the name Uredo may have included the
perfect stage, as in some species of Uromyces (U. Scirpi, etc.).

Since ecidiospores are almost always “ rounded-polygonal,”
their shape is not mentioned unless it deviates from this form.
In the systematic part, all the spores are drawn to the same
magnification (600 times), except where indication to the
contrary is given. The drawings are all original and from
British specimens, unless a different source is stated. When
several similar spores are outlined, the surface-sculpture is not
always indicated upon every one of them.

My thanks are especially due to Professor G. S. W est, by
whose advice this work was undertaken and by whose assistance
it has been chiefly carried through. The ‘Plow right’ Herbarium
of Fungi, which is in the possession of the University, has been

PREFACE Vii

of considerable use, and the Principal (Sir Oliver Lodge) most
kindly obtained for me a grant towards the cost of preparing
the illustrations. By the courtesy of Dr A. B. Rendle, Keeper
of the Botanical Department of the British Museum, and that
of the Director of Kew Gardens, the Herbaria at those insti-
tutions have been consulted and much useful information has
been derived therefrom. Thanks are also due, and are hereby
gratefully tendered, to Herr H. Sydow (of Berlin) and Mr J.
Ramsbottom (of the British Museum) who have both given
unstinted help, to Mr T. B. Roe (a very successful collector
of specimens), Professor A. H. R. Buller, Mr A. D. Cotton
Mr H. J. Wheldon, Mr C. Crossland, Mr J. Adams, Sir Frederick
Moore, and many others in a smaller degree.

W. B. GROVE.

THE BotanicaL LAporatory,
Usiversity oF BirnmincHay.
July, 1913.

CONTENTS

PAGE
PREFACE . 4 2 ; : : F Vv
INTRODUCTION . : ; : . ; . Xi
GENERAL PART
CHAPTER I
THE Lire-History or Pvecinia Caricis, THE NETTLE AND SEDGE
Rost : ‘ j ‘ . : i j : 1
CHAPTER II
THE SEXUALITY OF THE UREDINALES—THE NATURE OF THE
SPERMATIA—NUCLEAR DIVISION IN THE UREDINALES—THE
ALTERNATION OF GENERATIONS “4 F ; : 17
CHAPTER III
SPORE-FORMS OF THE UREDINALES—-GROUPING ACCORDING TO SPORE-
FORMS ; , ‘ : 30
CHAPTER IV
Lire-Historizs OF OTHER UREDINALES—Pucernra Grawixis—P.
Poarum—P. MaLvackarumM—GYMNOSPORANGIUM CLAVARLE-
ForMmME—FEwpopuH¥LLum SEMPERVIVI—CRONARTIUM RIBICOLA—
MELAMPSORA PINITORQUA—CALYPTOSPORA GOEPPERTIANA. 41
CHAPTER V
SPECIALISATION—IMMUNITY ‘i ‘ : . . . 62
CHAPTER VI
CLASSIFICATION—-PHYLOGENY . 3 73

Note on MANIPULATION . ‘ { z 84

x CONTENTS

SYSTEMATIC PART

PAGE

Pucciniacece UrnoMyces. 85
PUCCINIA . 128

TRIPHRAGMIUM 287

PHRAGMIDIUM , ‘ 289

KUEHNEOLA . . 299

XENODOCHUS . : 302

GYMNOSPORANGIUM F 304

Cronartiacesze CHRYSOMYXA . . . 310
CRONARTIUM . 313

Coleosporiaceze CoLEosPoRIUM 319
OcHROPSORA . 329

ZAGHOUANIA . 331

Endophyllaceze ENnpoPHYLLUM 333
Melampsoraceze MELAMPSORA . , 336
MELAMPSORIDIUM . 3858

MELAMPSORELLA 360

PUcCcINIASTRUM 364

THECOPSORA . 368

CALYPTOSPORA : 372

HYALOPSORA . 373

MILESINA F 376

UREDINOPSIS . . 3879

Appendix HEMILEIA . 3881
CHRYSOMYXA . : 384

UREDO . P . 385

ARCIDIUM s ; . 386

‘CHOMA . ‘ ‘ 388

EXxcLUDED SPECIES . 889

GLOSSARY : ; : : . 391
BIBLIOGRAPHY : ‘ : : . 893

INDEXES ; ; . ‘ : . 898

INTRODUCTION

THE Uredinales form a group of Fungi which is also spoken
of as the Uredinee or the Rusts. An accurate acquaintance
with their nature is of great importance to the gardener, the
forester, or the agriculturist, on account of the enormous loss
which is caused by them every year and which can, at least in
part, be avoided by a fuller knowledge.

All the species are parasitic, growing upon or in a living
plant, which is called the host. The majority of the species of
Uredinales have more than one stage of growth, distinguished
by the form and arrangement of the spores which they produce ;
the number of distinct kinds of spores which a single species
can possess varies from one to five. If the various spore-forms
are all borne upon one host, the species is called autoecious.
But it is a remarkable fact that a large number of the
Uredinales pass their existence alternately upon two hosté,
certain of the spore-forms being always produced upon. the
one, and the remainder upon the other. Such species are
called heteroecious or metcecious. Many of those which grow
upon grasses or sedges are probably hetercecious, though this
has not been shown in every case, and there are a few proved
exceptions. In order to convey a notion of the complex nature
of the Uredinales, one of the hetercecious species will be taken
as the type, and its various stages will be described.

ADDITIONS AND CORRECTIONS

p. 24. Fig. 225 should be Fig. 226.

p. 28. Fig. 241 should be Fig. 242.

p. 139. Puccinia tinctoriicolu Magn. Caster. Bot. Zeitschr. p. 491 (1902)
should be added as a synonym to P. tinctoriae.

p. 150. P. Leontodontis. I have since found this species on L. hirtus
(Thrincia hirta).

p. 238. P. oblongata has also been found on Luzula silvatica.

pp. 296—7. It is stated that Phragmidium violacewm infests most of the
subspecies of Rubus fruticosus, except those belonging to the group ‘‘ Corylifolii,”
while Ph. Rubi is confined to that group and the allied Rubus caesius.

CHAPTER I

LIFE-HISTORY OF PUCCINIA CARICIS, THE NETTLE
AND SEDGE RUST

Puccinia Caricis has two of its stages, the spermogonial and
ecidial, on the Nettle (Urtica dioica and other species), and
two others, the uredo- and teleutospore-stages, on various species
of Carex, especially C. paludesa. The first appearance on the
nettle is in the spring, about the end of April or the beginning
of May, when small swollen yellowish spots can be seen on the
upper surface of the nettle-leaves. These spots are round and
convex above, sunken beneath, and about 3—4 mm. in diameter ;
soon they turn orange on the upper surface, owing to the
development thereon of the spermogones, small flagon-shaped
bodies walled in by a large number of slender orange hyphe
and filled with many hundreds of minute spore-like cells, the
spermatia, which are orange in mass, though singly they appear
colourless (Fig. 1).

Fig. 1. P. Caricis. Section of leaf of Nettle, showing the hypertrophy
produced by the mycelium of the ecidium-stage; a spermogone, on
the upper side of the leaf, and two ecidia, one closed, on the under
side. The upper face of the leaf is turned downwards. x 60.

G. U. 1

2 SPERMOGONES

The mycelium in the leaf is strictly localised, forming little
knotted masses (plectenchyma) just beneath the epidermal
cells of the affected spot; all the cells of this mycelium are
uninucleate. Some of these hyphe turn upwards, remaining
densely crowded and more or less parallel to one another, and
enclose the flask-shaped cavity, at length converging to a point
above it and piercing the epidermis at that place. Then the
upper ends of these hyphe diverge and form a brush-like bundle
surrounding a narrow canal, or ostiole, which connects the
cavity with the external air (Fig. 2). Meanwhile other hyphe

2%

Fig. 2. P. Caricis. Vertical section of spermo-
gone, on leaf of Nettle. x 200.

from the base have grown up within the flask, and made a
lining to its lower half; these hyphe are exceedingly delicate
and numerous, and each abstricts from its end, successively,
large numbers of the spermatia (Fig. 2), accompanied by a
quantity of sugary mucilaginous matter which binds the
spermatia into a coherent mass. The mucilage soon swells by
imbibition of moisture, and the spermatia are forced out of the
flask, through the ostiole, and form an orange globule between,
and on the top of, the diverging hairs. Ultimately the mucilage
dries up, or is washed away by rain, and the spermatia are
dispersed.

The spermatia are very small, thin-walled, oblong or roundish
cells, each containing a single relatively large nucleus, but
little cytoplasm and no reserve material. When placed in a
nutritive solution, they are capable of a kind of germination,

ACIDIA 3

but the mycelium produced is very scanty. Their function
will be discussed in a later paragraph, but it may be mentioned
here that all attempts to produce infection by means of them
have uniformly failed.

THE AXCIDIUM.

Before the spermogones have completed their development,
similar but larger conglomerations of hyphe arise on the lower
surface of the leaf, a little way below the epidermis, which they
raise up into a rounded dome. These masses enclose a number
of erect hyphe of two kinds—an
outer series of parallel closely-
joined colourless hyphe, forming
the peridiwm, consisting of more
or less hexagonal cells, which meet
above and roof over the dome-
shaped cavity; and an interior
series which remain shorter and
give off from their upper end
parallel chains of spores called
aecidiospores, which fill the whole
of the enclosed space. Each new
spore is produced beneath the older
ones, which are thereby pushed
gradually up. Finally, this struc-

: 5 cen Fig. 3. Leaf of Nettle (under
ture, which is called an aecidium, side) showing the scidia

of P. Caricis on the lamina

ruptures the epidermis, forces its andl petiole. 99%

way between the cells, the peridium

bursts at the summit, the edges become revolute, and there is
formed a white cup-shaped hollow, its floor covered with erect
chains of orange spores (see Fig. 1). These spores have a
rather thin, colourless, finely warted cell-wall, and are filled
with rich bright-orange granular and oily contents. The
mycelium which forms the ecidia is continuous with that
which bears the spermogones, and its cells are uninucleate, but
the spores themselves are binucleate. The origin and meaning
of this change will appear afterwards.

1—2

4 ECIDIA

The oldest spores being at the top, they separate as soon as
mature, and are blown away by the wind; fresh spores are
produced for a time from below. Since these cups are in dense
clusters over the whole of the affected spot, they are known as
“Cluster-cups.” The mycelium in the leaf continues to develop,
and the spot enlarges and ceases to be round. But the swelling
of the tissues within which the mycelium is living can be
carried to such an extent as to distort and curl the leaf, much
in the same way that Hxoascus deformans (the Peach Leaf-curl)
affects the Peach leaves. The epidermis above and around the
spot, also, often becomes coloured red or purple by an anthocyan-
derivative, both these effects being a response by the leaf-cells
to the stimulus of the parasitic growth, and perhaps part of an
effort to throw off or checkmate the invader. The mycelium,
moreover, is not confined ‘to the leaves; it may originate in
the petiole, in the stipules and in the stem. In the latter case
it causes notable curvature and distortion; an instance is
recorded where the curved gall-like mass, formed on the stem,
measured as much as 10 cm.‘in length, and similar but smaller
growths may frequently be met with.

Here should be noticed the close parallelism, up to a certain
point, in the formation of the spermogones and the ecidia—
they have similar shapes, they are both enclosed by a layer of
sterile hyphex, they arise in (usually) quick succession on the
same mycelium, they give off basipetal chains of spores frgm
their base. On the other hand, besides the difference any the
nuclei, the xcidiospores differ from the spermatia in their larger
size, In possessing a large store of reserve food, in their capacity
for germination, and for producing infection in another plant.

THE AICIDIOSPORES.

The mode in which the ecidiospores are produced is as
follows:—The upper cell of the sporiferous hypha usually
divides into two, an upper sterile cell, and a lower fertile or
basal cell, each with one nucleus. -The upper cell disintegrates
and perishes; the lower fertile cells conjugate with one another
in pairs, the process consisting in the formation of a small hole

ZCIDIOSPORES 5

in the dividing wall which afterwards enlarges until at length
hardly any trace of the wall is left. The two cytoplasms thus
form one mass, but the nuclei arrange themselves more or less
side by side without fusing or in most cases even touching.
This process must be regarded as an act of fertilisation or
rather as a substitute for such an act. The double cell is
called a fusion-cell.

The two adjacent nuclei are said to be “ paired,” and to-
gether constitute a synkaryon or dikaryon. Occasionally, in
some species of Puccinia and other genera, these’ fusion-cells
contain three nuclei, probably by a double fusion. Such cells
produce trinucleate spores, but the fate of these is not known.

The paired nuclei of the fusion-cell then divide side by side
and simultaneously—a process called conjugate division—and a
wall is formed between the two pairs. The wall is formed in
such a way that the two nuclei in the same cell are not sister-
nuclei (see Figs. 19, 20).

The fact that in conjugate division the paired nuclei divide
so that the two are usually in the same stage of mitosis at the

1 The stages connected with the sexual process in general are three in
number—(1) the association of two (almost always non-sister) nuclei in the
same cell, (2) the fusion of the two nuclei, preparatory to (3) the reduction in
the number of chromosomes, or at any rate in the amount of chromatin, to the
previous ordinary vegetative condition. These three stages may or may not
follow closely upon one another. The first and second, or the second and
third, or the third and first, may-be separated by an intervening series of cell-
divisions. Which is to be regarded as the actual fertilisation? In all proba-
bility the first. The nucleus is the director, the cytoplasmic mass is the work-
people of the cell-factory. The presence of the two directors is the essential
fact. The fusion is of less importance; it is often delayed for a long period,
and in certain cases it is known that, even when fusion has taken place, the
chromosomes still retain their individuality for a considerable time. There
is reason to believe that on the first introduction of fertilisation these three
stages followed immediately (i.e. without intervening cell-divisions) after one
another, the series of vegetative divisions being intercalated between (3) and (1),
as in Coleochaete (Allen, 1905) and Spirogyra (Tréndle, 1907; Karsten, 1909).
In the majority of the higher plants, stages (1) and (2) occur without much or
any interval, but a long series of divisions (the sporophyte) is intercalated
between (2) and (3). In most of the Uredinales, the chief series of vegetative
divisions is intercalated between (1) and (2), and (3) follows immediately after
(2). In certain Alge (Griffithsia, Dictyota) the intercalation takes place on an
extensive scale, both between (2) ‘and (3) and between (3) and (1).

6 ZECIDIOSPORES

same time, shows that they mutually influence each other, and
implies that all the cell-processes which go on are likewise
under their common control.

The upper cell, cut off from the fusion-cell, is the ezcidio-
spore-mother-cell; the lower grows a little longer and then
divides again in the same way, and thus a vertical series of
eecidiospore-mother-cells is formed, the
oldest at the top. Each of the zcidio-
spore-mother-cells, as soon as produced,
cuts off, by conjugate division, a small
cell below, called the intercalary cell;
this soon becomes disorganised and
disappears, while the other portion
becomes the aecidiospore. Thus the
chain at first consists of alternations
of «cidiospores and abortive spores,
both containing two nuclei (Fig. 4);
the function of the latter may be,
by their disintegration, to enable the
xcidiospores to separate more easily
from one another, and thus to aid dis-
persion by the wind.

The peridiwm is composed of cells
Fig. 4. P.Caricis. Chainof homologous with the spore-mother-

young weidiospores, x 500. elis) They arise from basal cells con-

a, fusion-tissue; b, basal rae —
(fusion) cell, with con- taining two nuclei in exactly the same

jugate nuclei; ¢, xcidio- . :
spore-mother-cell; d, in. WAY; Occasionally even an intercalary
tercalary cell; e, young ell is cut off, but it does not dis-
zcidiospore. : :
integrate. All of them thicken their
walls and remain in close contact with one another; ultimately
their contents disappear (Fig. 5). The central part of the
dome-shaped peridium is composed of the terminal cells of
the central spore-bearing hyphz ; they are probably less closely
connected with one another, and the peridium ultimately
bursts at that point.
The spores, if placed in a damp atmosphere, germinate
readily when mature and fresh. Only those spores which are
at the top of the chains and can be shaken out easily by

GERMINATION OF ACIDIOSPORES 7

tapping are mature enough to germinate, and even they, owing
to their thin walls, may lose this power in a few days according
to circumstances. Especially can they be killed by rapid

|

Fig. 5. P. Caricis. a, three cells of the peridium, on Nettle ; b, a cell in
optical section; c, the same in surface-view ; d, two xcidiospores. x 600.

drying. Instances are known, however, where some of them,
kept in a cool place, retained their capacity for germination
about seventy days, though most of them were dead after
eighty days.

Each spore has a number of germ-pores; in Puccinia Caricis
the number is about five or six; in other species of Uredinales
the number varies from two to eight.
These pores, which are scarcely visible
until germination begins, are thin places
in the inner layers of the outer wall, the
whole cell bearing a close resemblance to
many kinds of pollen-grains (microspores).
(Fig. 6.)

It is worthy of notice that in the
spores of Endophyllum, and others of the
less-advanced type,there are no real germ-
pores; the germ-tube merely forces its/ ; mo,
way out at the first place that gives way. gy ped Soak
From this state of things there isa gradual _in water. x 180.
transition from numerous to few germ-pores, in the highest
types of all (Uromyces and Puccinia teleutospores), each cell
has one and only one well-defined germ-pore.

8 GERMINATION OF ACIDIOSPORES

The germ-tube of the ecidiospore of P. Caricis can grow to
a length ten or twenty times greater than the diameter of the
spores, and often executes, at least in water, a number of
spiral turns (Fig. 6); it remains nearly of the same diameter
throughout, or may bear short stumpy branches here and there.
The granular contents of the spore travel along the tube with
its growth, remaining always towards the distal end. But
unless the germinating spore has fallen upon its proper habitat,
the leaf of a Carex, its attempt at growth comes to an end
when its reserve-material is exhausted.
If, however, the spore has fallen upon a Carew, its germ-
tube travels over the surface until it reaches a stoma, through
the pore of which it enters the respiratory chamber, forms a
swelling just inside as a kind of hold-fast, and then begins to
branch and traverse the intercellular spaces, occasionally sending
an haustorium into the mesophyll-cells
(Fig. 7). The cause of its entry is pro-
bably the search for water-vapour, since
the germ-tube of a Uredine is found
(De Bary, 1863; Gibson, 1904) to enter
the stomata as freely on another leaf as
on one of its proper host-plant, and also
to pass through a hole, comparable in
size to stomata, in a thin india-rubber
Fig. 7. Haustorium of Membrane which separated it from air
P. Caricis in chloro- saturated with water-vapour (Balls, 1905).
Lis anes But its further growth is influenced by
rium; m, nucleus of chemotaxis of a more complicated nature:
cell, x 500. : . : E
unless the right kind of stimulus is
furnished by its host, it cannot form effective haustoria, de-
velopment is poor and abnormal, and death soon ensues (Gibson,
1904). The resistance of the host to the parasite, shown
perhaps by the secretion of destructive enzymes, has also to be
considered. Once ipside the stomatal chamber, however, the
fungus is largely protected from outside influences, such as
desiccation: this preservative habit has no doubt contributed
much to the wide-spread prevalence of the Uredinales.

UREDOSPORES 9

THE UREDOSPORES.

The germ-tube soon forms a more or less extensive my-
celium, which may penetrate the greater part of the leaf’ of
the Carex, but in many species of Puccinia is strictly localised
to a small defined spot. All its cells are binucleate, like the
spore from which it originated. The cells of the mycelium, in
every stage, send haustoria into the cells of the host; when an
haustorium arises from a binucleate mycelium, it is itself

Fig. 8. Section of leaf of Carex paludosa, with a sorus of uredospores of
P. Caricis; a, upper epidermis; b, a vascular bundle. Most of the
pedicels have lost their spores. x 180.

likewise binucleate. After a few days this mycelium begins to
form the third kind of spore—the uredospore. A knot of
hyphz is formed just beneath the epidermis; some of the
branches turn upwards and form a regular
layer parallel to the surface—the spore-
bed (Fig. 8).

The upper rounded cell of each hypha
is divided into two daughter-cel]s, the
lower of which is developed into a
stalk, the upper becomes the uredospore
(Fig. 9). The spore is oval or roundish ;
when mature it is enclosed in a double
cell-wall, the outside being cutinised and wig.9. P. Caricis. De-
provided with spine-like projections, some- veloping uredospores

2 cones (1, 2, 3, 4, show stages
what like those of the scidiospore, only of growth ; 5 is a pe-
more pointed. In the inner layers of the _ dicel from which the

spore has vanished).
exospore there are usually three (rarely z 500. }

10 UREDOSPORES

four) germ-pores; in fact, the uredo- resembles the ecidiospore
in character, and must be considered as homologous with it—
the stalk-cell corresponding to the intercalary cell of the latter.
But they differ considerably in the fact that the uredospore is
always produced singly, not in chains.
(This is not true, however, of all
the Uredinales.) The membrane of
the uredospore is nearly colourless, but
it encloses a bright orange granular
and oily mass, with two nuclei. Every
cluster of uredospores produced on the
same spore-bed is called a sorus; it is
surrounded by the laciniz of the epi-
dermis, which is more or less torn or
Fig. 10. Leaf of Carex split by the enlarging mass. In many
pendula, with uredo- and .
teleuto-sori, slightly en- C488, several sori become confluent
larged. and form a larger pustule (Fig. 10).
In other species of Uredinales the uredospores have coloured
membranes or possess a larger or smaller number of germ-
pores. Moreover the distribution of these pores over the
surface is characteristic for each species: they may be placed
equatorially, as they are in P. Caricis (Fig. 11), or towards
the poles, or scattered over the surface with regularity or
without any order.
A uredospore may be very easily detached from its pedicel,
and a ie (chiefly by wind, though sometimes by insects)
ype to another leaf of Carew, on which it
germinates, the germ-tube enters a stoma,
produces a fresh crop of mycelium and
another sorus of uredospores; this process
can be repeated indefinitely. The my-
celium can also grow up and down the
Be map Hee leaf, producing fresh sori in its course ;
showing the three for this reason the sori are usually
germ-pores. x 600. arranged in linear series, owing to the
parallel venation of the Carex-leaf. The germ-tubes of the
uredospores are often curled or branched like those of the
secidiospores, and the germination is of the same character in

TELEUTOSPORES 11

both (Fig. 12). The uredospore retains its capacity for
germination for a longer time, even: .
for more than three months; in fact,
in certain foreign species, some of these
spores acquire a thicker wall which
enables them to act as a kind of
resting-spore—these are called amphi-
spores, but they are not formed by
P. Caricis.

It is found, generally, that if the
spores of the Uredinales are dried
gradually, they retain their power of
germination for a longer time and in
a better degree than if dried quickly
or not dried at all. Most likely a slow Fig. 12. Two uredospores

of P. Caricis, both germi-
drying enables them to mature more nating in distilled water.
perfectly. laa

THE TELEUTOSPORES.

After a time, probably in response to the weather or other
change of environment, the mycelium which has hitherto given
rise only to uredospores begins to produce, at first in the same,
afterwards in separate sori, the fourth kind of spore—the
teleutospore. In the genus Puccinia this is almost always a
compound body, formed of two superposed cells; each cell is
really a spore, and is capable of independent germination. In
many species of the genus the teleutospores readily break
apart at the septum, eg. in Puccinia fusca and P. Pruni-
spinosae, and the lower half may be, and has been, mistaken for
a uredospore. Those of P. Caricis do not easily break apart
until they are old and dead.

The teleutospores are formed on a pedicel, much in the
same way as the uredospores, except that the uppermost cell
is again divided, but apart from that they differ widely in
their character. They have a thick dark-brown exospore,
covered with a chitinous cuticle; in this species the exospore

12 TELEUTOSPORES

is much thicker at the apex than elsewhere (Fig. 13). While
the spores are standing in a densely crowded

\ sorus, the apex is the part most exposed to the
: weather, and therefore most needing protection.
There is a thin endospore to each cell: the

contents are granular and at first oily; there 1s
a large and conspicuous nucleus in the centre
of each. This nucleus, in its resting stage, 1s
almost homogeneous except for its nucleolus,
and was mistaken by the older observers for
a “vacuole.”

Since the mycelium from which the teleuto-
Fig.13. Teleuto- Spores, as well as the uredospores, were formed
spore of P. gontains paired nuclei, the cells of the teleuto-

Caricis. x 600. " . Lee
spore were at first in the same condition. When
its wall, however, begins to thicken, ie. when it is becoming
mature, the conjugate nuclei unite, and form one large fusion-
nucleus (Fig. 14). The two fusing nuclei, after the very

@
& o ae
©
4* 2
a b c d e

Fig. 14. Formation of teleutospores of P. Falcariae (after Dittschlag) ;
u, the spore-bearing hypha; b, the same divided into pedicel and
spore-cell; c, the spore divided into two cells; d, a young teleuto-
spore; ¢, the same after the fusion of the nuclei. x about 800.

numerous conjugate divisions during the long period of growth
from the formation of the fusion-cell of the secidium, would be
related, as it were, like very distant cousins, especially since
the nuclear divisions during this period, though indirect, appear
to show a very simplified form of mitosis, tending rather to be
of the nature of amitosis. The fusion, as already intimated, is
not to be considered as the act of fertilisation, but merely as a
necessary preliminary to chromatin-reduction.

MESOSPORES 13

It is in the germination of the teleutospore, presently to be
described, that its most distinctive feature is to be found. The
chief function of teleutospores is to act as resting-spores, and
in the majority of cases they will not germinate until they
have passed through a period of quiescence; in the present
instance this period is the winter, but it is not necessarily
always so. The resting-spore is primarily a device to tide over
an unfavourable period—whether of food-supply, moisture,
temperature, or resistance of host—without regard to season.
Some species, however, have teleutospores which can germinate
immediately, as in P. Malvacearum ; those teleutospores usually
have thin walls. P. Malvacearum is sometimes supposed to
hibernate by a perennial mycelium, but there is reason to
believe that in most cases infection each year proceeds from
over-wintered teleutospores. Most of the species which have
these thin-walled spores also produce some with thicker walls,
which act as resting spores in the ordinary way.

Besides the two-celled teleutospores, several species of
Puccinia also produce similar spores with only one cell—these
are called mesospores. A mesospore can occasionally be found
in many Puccinias, even in P. Caricis (Fig. 15),
but in others they are abundant, eg. in
P. Porri, where careful search is often required
before a two-celled spore can be detected.

Mesospores arise merely by the omission of the

last nuclear division; they are exactly of the

same nature as the two-celled teleutospores yi, 15 4 meso-
and germinate in the same way. By this spore of P. Car-
means they can be distinguished from the “” ae
amphispores previously mentioned, but not of course from the
teleutospores of Uromyces. In fact authors have described
some species which produce them as Uromyces, overlooking the
rarer two-celled spores that occur with them. See remarks .
under Puccinia Porri and Uromyces ambiguus.

14 GERMINATION OF TELEUTOSPORES

GERMINATION OF THE TELEUTOSPORE.

We now approach the consideration of a process which has
been in the past much discussed, and upon the right inter-
pretation of which the whole question of the systematic position
of the Uredinales depends. Each cell of the teleutospore of
P. Caricis has one germ-pore, though some genera allied to
Puccinia have teleutospores with more than one germ-pore to
each cell, e.g. Phragmidiwm, Uropyxis, Calliospora. The germ-
pore of the upper cell is in the thickening at the summit, that
of the lower cell is lateral and just beneath the septum. Each
of these pores is a canal passing through the cell-wall, and
covered only by the cuticle. Through these pores the germ-tube
passes, first appearing as a roundish swelling, the protoplasm
being surrounded by the thin endospore. This then elongates,
the nucleus squeezes through the relatively narrow pore and
enters the tube where it divides
twice, and forms four superim-
posed cells, separated by thin
cell-walls (Fig. 16). This row of
four cells was formerly known as
a promycelium, but is now called
a basidium. If kept in water
these cells can round off and
separate from each other!, and
germinate by sending out a tube,
se seve pe ets pati like the mycelial cells and spores

same, germinating and forming of many fungi. But if ina damp?
pteaeee cee a atmosphere, each cell without
separation produces a sterigma at
the end of which a basidiospore is formed, like the basidiospores
of Agaricini. These basidiospores can germinate at once, even
before they are detached from the sterigma, by sending out
a short tube which may produce a conidium resembling the
basidiospore at its end.

1 This method is said to take place normally in Barclayella.
* It is noted by many observers that, in a state of nature, it is a layer of
dew, not of rain, that is favourable to germination.

GERMINATION OF BASIDIOSPORES 15

If one of these easily detached basidiospores or conidia is
conveyed to the surface of a leaf or young stem of Nettle, its
germ-tube bores through the cuticle
and enters the tissues (Fig. 17),
where it ramifies and forms a my-
celium. The teleutospore is large
and heavy, and firmly attached to
its spore-bed on the leaf of Carex;
the basidiospores enable its con-
tents to be transferred easily to Fig 17. Endophyllum Sem-

pervivi. Germinating basi-
the surface on which alone they diospores (after Hoffmann) ;
are capable of further growth. But —%,,tbe spore: ® the germ.

. i ‘ ‘ vesicle, under the cuticle of
their wall is thin and they can live _ the epidermis; a, b, c, show

only for a short time; they contain bea care Sa oe Seecinley
but little food-supply and could not —-* about 200.

form a long germ-tube. That is the reason why their germ-
tubes do not, like those of the other spores, search for a stoma,
but enter by the quickest means. Nevertheless they can
abnormally enter by a stoma; De Bary records such a case in
his account of P. Dianthi (see Fig. 24).

The germination of the teleutospores of P. Caricis takes
place about the second week in April, and on the mycelium
produced by the basidiospores in the nettle there arise, in
about a fortnight, first spermogones and then ecidia like those
with which we started. But the mycelium arising from the
basidiospores does not always proceed immediately to spore-
production. In some species, e.g. Endophyllum Sempervint, it
hibernates in the growing point of the shoot, or in the leaves
if they are evergreen, as in Puccinia Bust, or in the stems or
branches in the case of some that live on shrubs or trees, as in
Cronartium ribicola.

Rather more than a twelfth of the species of Uredinales are
now known to be hetercscious. This mode of life may be
regarded partly as a device by which the parasite tides over
the time during which one of the host-plants is not available.
The leaves of the Nettle are delicate and soon perish in the
autumn; those of the Sedge persist throughout the winter.
The power of hetercecism increases the ability of the fungus to

16 HETERGCISM

adapt itself to new conditions and thus tends to perpetuate the
race, while the change of host, which is equivalent to a change
of diet, may very possibly tend to an increase of vigour in the
individual.

The former statement, however, must not be taken to refer
to all cases of hetercecism, since there are instances, e.g. in
species of Coleosporium, which cannot be explained on this
ground.

The reason why P. Caricis has been taken as the typical
Uredine, instead of the usual P. graminis, is that the ecidium
of the latter is now very rarely found in this country and is
therefore not available for demonstration, while that of P. Caricis
is common in all suitable localities. Even if not existing in
any place, it can be readily introduced if the three prerequisites
are at hand:—a pond bordered by Carex paludosa and by
Urtica dioica, and also a quantity of the leaves of Carex
infested by the parasite. The Nettle and the Sedge are not
injured appreciably by the disease, nor would it be of much
consequence if they were. To introduce the fungus into the
new locality, it is only necessary to obtain a bundle of the
required leaves (say about 500) from some place where the
Pucciniu exists, in January or February, and lay them on
the ground where a patch of nettles is known to occur. The
latter will be seen in spring to be beautified by the ecidium,
and in the summer the disease will spread to the surrounding
sedges (see Grove, Journ. Bot. 1913, p. 42).

4

CHAPTER II

THE SEXUALITY OF THE UREDINALES

De Bary suggested in 1884 that, if there was any sexual act
occurring in the life-cycle of the Uredinales, it would probably
be found in connection with the ecidium. At that time
nothing was known on this point, and De Bary anticipated that
something might take place analogous to what happens in the
formation of the asci of certain Ascomycetes.

The first discovery was made by Blackman (1904), who
found that, in laying the foundation of the spore-bed of the
ecidium of Phragmidium violuceum, cells became binucleate
by the passing into them of a
nucleus from an adjoining vege-
tative cell. He saw that this
passage took place through a nar-
row opening formed between the
cells (Fig. 18). The binucleate
cell then became the equivalent
of an oospore, and formed the be-
ginning of a sporophytic genera-
tion. In other words, he supposed
that the fertilised cell represented
a female gamete and the vegeta-
tive cell replaced a now vanished ae
or functionless male gamete. The Development of ecidium (after
fertilisation would then be of the Lapeer ERR Peed
nature of a semi-apogamy. He cells below, a nucleus is seen
considered that the spermatia ia iin Sides
were the functionless male gametes. From the ‘cell to be
fertilised he saw an upper sterile cell cut off, which soon

G. U. : 2

18 BASAL CELLS OF ACIDIA

degenerated ; this he considered to represent an abortive tri-
chogyne, in accordance with De Bary’s anticipation. The
acceptance of this interpretation implies the existence of a
close affinity between the Uredinales and the Red Seaweeds.

In 1905, Christman published the result of his researches
into Phragmidium speciosum, Caeoma nitens (= Gymnoconia
= Puccinia Peckiana), etc. According to him, the process that
took place was the fusion of the contents of two equal and
similar gametes, with the exception that the nuclei remained
side by side unfused. A considerable portion of the wall
between the two fusing cells was broken down, and the process
was of the nature of a conjugation, not a fertilisation.

Blackman and Fraser (1906) next examined a number of
other species, and in Melampsora Rostrupii they found the
same process which Christman had observed, though they still
considered that other species, e.g. Puccinia Poarum, showed
instanees of the migration of a nucleus as in the first subject
studied.

Christman, in 1907, showed that a similar act of conjugation
between two equal cells takes place in the formation of the
primary uredospores of Phragmidium Potentillae-canadensis
(= Kuehneola Tormentillae, Arthur, q.v.), the primary uredo-
spores in this species replacing the eecidium which is absent.

In 1908 Olive, in examining the primary uredospores of
Triphragmium Ulmariae, tried to reconcile the difference be-
tween these opposing views: he considered that conjugation
took place between two cells, one larger and one smaller, and
that either a large opening was formed so that the two proto-
plasts fused, or a narrow hole was produced through which the
nucleus of the smaller cell passed into the larger. He considered
the upper sterile cell as a degenerating tip-cell, not an abortive
trichogyne. The fusing cells might be placed in almost any
position with respect to each other. In Puccinia transformans,
a micro-form, possessing only teleutospores, he shows that the
basal cells which produce them arise equally by the fusion of
two uninucleate cells.

Kurssanow, in 1910, investigating Puccinia (Gymnoconia)
Peckiana, found both cases. that of Blackman and that of

STERILE CELLS 19

Christman, occurring side by side. All the conjugating cells
had an upper sterile cell which he calls a “buffer” cell; but
the passage of the nucleus only he put down, as others have
done, as a pathological phenomenon, caused perhaps by the
method of fixing.

In November of the same year Dittschlag, investigating
Puccinia Falcuriae, tried to settle the question and decide
definitely the function of the spermatia. This Puccinia is an
-opsis form, having spermogones and ecidia, followed later by
teleutospores, but without uredospores. He showed that the
cells of the spore-bed of the ecidium unite in pairs by the
disappearance of not quite all the separating wall. If a sterile
cell could be seen at all, it was seen equally on both (Figs.
19, 20).

But this does not militate against its being considered as
a degenerate trichogyne: it is certain that the two cells which

@ @ @@ }»
Ci fe : at
Oe \* 6
a b

Fig. 19. Puccinia Falcariae. Con- Fig. 20. P. Falcariae. Formation
jugation of two female cells to of cidiospores (after Ditt-

form the basa] cell of the wcidio-
spore-chain (after Dittschlag).
The uppermost cell on the left
in a does not belong to the others.
Each fertile cell has a sterile
cell above it. In b, the first
conjugate division is just com-
pleted (Diagrammatic).

schlag): u, the basal cell; b, an
zcidiospore-mother-cell; c, the
same in the act of conjugate
division (the nucleoli are seen
in the middle); d, the inter-
ealary cell cut off.

fuse are in most cases exactly alike, and therefore, if they
represent potential female cells, each of them would naturally
be provided with a trichogyne in equal degree, if at all.

2-9

20 BASAL CELLS OF ACIDIA

Again, in 1911, Hoffmann investigated a Uredine of a lower
type than most of those previously considered, viz. Endophyllum
Sempervivi. This genus differs from all the other Uredinales
in its mode of development. It has only spermatia and zxcidio-
spores, the latter functioning also as teleutospores in that their
conjugate nuclei fuse, and then on germination they produce a
basidium and basidiospores. These basidiospores reinfect the
host and produce both spermogones and ecidia. On the spore-
bed of the ecidium two ad-
jacent cells unite by the disso-
lution of the intervening walls
(Fig. 21); first a small hole
is formed, which grows larger
until at last almost no trace of
the wall is left. The disap-
pearing wall is often horizontal,
not vertical as in most of the
other cases, and the conjugating
cells are not situated in any

definite plane. In such cases
Fig. 21. Endophyllum Sempervivi.

Formation of scidio-teleutospores © sterile (trichogyne) cell was

(after Hoffmann). a, fusion-tissue; yot seen.

b, basal cell ; c, intercalary cell; d, :
spore; ¢, intercalary cell, degenera- Finally Fromme (1912)

ting. found that in Melampsora Lini
the spermogones and xcidia are produced simultaneously and
only from infection by basidiospores. The spermatiophores
ditfer from all others described in being many-celled, each cell
producing a single spermatium on a sterigma-like process; they
arise from a regular layer of large rectangular cells at the base
of the spermogone. The ecidia are stated to be undistinguish-
able from the spermogones externally, but produce female
gametes in the usual way, generally with one or two “ buffer”
cells which speedily disintegrate. The female gametes conju-
gate, in abundance, laterally in pairs, often in threes or fours;
the fusing cells are of equal rank, but need not be in the same
horizontal level. Aicidiospores were observed with several
nuclei, and one xcidiospore-mother-cell was seen with as many
as eleven nuclei. (See also note on p. 29.)

TRICHOGYNE HYPOTHESIS 21

From a comparison of all these observations it now becomes
certain that, in the Uredinales, the typical mode by which the
binucleate condition arises is by the conjugation of two similar
cells, each provided with a large nucleus and an abundant
supply of food. This fusion-cell can afterwards branch by the
formation of lateral buds, usually in basipetal succession, and
may thus produce several rows of spores at once ; by a similar
branching bunches of uredospores and teleutospores can arise in
the sori of those spore-forms (Blackman, ’06, Christman, ’07,
Dittschlag, 10, Hoffmann, ’11). See Figs. 37, 156.

At first, it may be supposed, the two conjugating cells .
belonged to a definite basal layer or spore-bed, as in Phrag-
midium, Puccinia spp., and Melampsora Rostrupii; but after-
wards they ceased to be arranged in a layer, and conjugation
took place between two purely vegetative cells in the mycelium
beneath. The beginnings of this change are seen in I.
Rostrupw, and its final product in such micro-forms as P.
Adoxae where the greater part of the mycelium has synkarya.

Whether the upper sterile cell which is so frequently met
with is to be considered as an abortive trichogyne is not so
certain. But it may be remarked that, since the sori generally
arise beneath the epidermis, no fertilisation could have taken
place by non-motile spermatia unless there were something of
the nature of a trichogyne to protrude through a stoma. In
this connection it is important to remember that in Uredinopsis,
one of the lowest of the Uredinales, the sori of primary uredo-
spores, i.e. ecidiospores, seem always to arise beneath a stoma ;
other sori can arise in many genera in the same way, and in
the genus Hemileia the pedicels of the uredospores protrude
into the air through the stomatal pore. Moreover we know
that, while ecidia of the enclosed higher types, as in P. Caricis,
arise at some depth in the host-tissues, and the basal layer and
its peridium are covered by a considerable thickness of dead
empty cells (which are afterwards pushed to the sides), the
eecidia of the more primitive form, the ceoma type, are shallow
and are not enclosed in a peridium, but are either quite naked
or surrounded only by a few paraphyses.

In the typical Uredinales, the conjugation of the two

ABSENCE OF ASCOGONIUM

male) cells takes place just before the formation of the
idium or its representative ; in the reduced mucro- and other
ms, such as P. Adowae and Uromyces Scillarum, it is probable
it the conjugation of the two (vegetative) cells takes place
some more or less indefinite period before the formation of
eutospores.

There is a general agreement among investigators that a
ucture resembling an ascogonium (from which the mass of
sal cells may be supposed to originate) does not exist in the
edinales, notwithstanding the suggestions to that effect by
wssee (1888) and Richards (1896). If it did exist, or had
sted, it would do something towards accounting for the
finiteness in form usually presented by an ecidium; if it is
ially unrepresented, the ecidium cannot be regarded as a
phological unit, but only as a collection of female cells. It
possible that traces of its existence are shown by the large
iultinucleate cells, containing 12—15 nuclei, which have been
scribed by Olive (1908) in the mycelium at the base of the
ung ecidium of Puccinia Cirsii-lanceolati and by others at
2 base of teleuto-sori, but this question must remain open till
‘ther investigations are made. The existence of an asco-
nium of that kind would, of course, be inconsistent with the
chogyne-interpretation of the sterile tip-cell in the ecidium.
fEcidiospore-mother-cells and excidiospores with three or
an more nuclei are frequently met with; these represent the
sult of a fusion of three or more cells of the spore-bed. But
ice uredospores and teleutospores with cells containing more
an two nuclei seem to be unknown, it is probable that these
normal zecidiospores undergo no further development.

THE NATURE OF THE SPERMATIA.

There are two and only two possible interpretations of the
srmatia—either they are male gametes, or they are conidia,
. merely additional multiplicative spores like the uredo-
ores. In favour of the former view the following arguments
a be adduced :

(1) The time of their appearance, just before the formation

NATURE OF THE SPERMATIA 23

of the ecidium or its representative, and on the same my-
celium. For instance, in those cases where there is no ecidium,
but primary uredospores which are formed, like ecidiospores,
from a fusion-cell, followed by secondary uredospores which are
not so formed although similar in all other respects—then the
primary spores alone are accompanied by the spermogones.
This argument is the most decisive.

(2) Their size and character. They are much smaller than
the other spore-forms, with thin walls, a large and not very
degenerate nucleus though often without a nucleolus, little
protoplasm and no reserve-stuff (oil, etc. with which the
ordinary spores of the Uredinales are so richly provided), thus
reminding one of the spermatia of the Floridez.

Sharp (1911) reports, in Puccinia Podophylli, spermatia
three times as long as the nucleus, and therefore containing
some appreciable amount of cytoplasm. But protoplasm is not
reserve-stuff.

(3) They will not reproduce the species. All the efforts
that have been made to cause them to do so have uniformly
failed. All the other reproductive cells of these Fungi can be
successfully used for that purpose, if applied to the proper host.
In some species, as Cronartium ribicola, the spermatia can be
collected in large quantities: Klebahn made numerous trials
with them, but entirely without result. Jaczewski and others
have confirmed his experience. It is a commonplace observa-
tion that highly specialised male cells cannot in themselves
reproduce the species, while female cells can, as in the cases of
parthenogenesis, both true and false. This difference in be-
haviour is partly correlated with the difference in the amount
of food-reserve available, with which the larger female gametes
are usually well supplied. In conjugation, where the two
(male and female) gametes are approximately of equal size (as
in certain Mucorini), each may form a functional azygospore.

(4) They will hardly germinate in water, probably because
they have no reserve-food. If food is supplied by cultivating
them in nutrient solutions, a little growth is obtained, but it
is very insignificant and soon perishes. The same thing is
true of male cells in other organisms. Conidia, under such

NATURE OF THE SPERMATIA

cumstances, would show quick and luxuriant growth ; they
inot be degenerate conidia, because the nucleus is large and
ll-formed though at times no nucleolus can be seen.

(5) They are sometimes accompanied by a sweet flurd,
ich gives off a pleasant, or more rarely an unpleasant, smell,
in P. suaveolens (obtegens), Uromyces Pisi, Cronartium Quer-|
sete. It is said that, in Japan, children lick the abundant
2rmogones of C. Quercus on account of the sweet juice that
zes from them. The presence of this can be readily under-
vod, if the aid of insects is invoked as well as wind, in order

carry the passive spermatia to the trichogyne projecting
rough a stoma, but otherwise is without explanation. The
‘va of a fly (Diplosis) or a similar organism, is to be found
iwling about the leaf and feeding on the spermatia and ecidio-
ores of many Uredinales; its body is quite orange in colour
rough being filled with them, and the spermatia would adhere
its outer surface. Though the spermogones of P. Caricis are
ually on the opposite leaf-surface to the ecidia, yet in very
my species they occur intermixed, and not infrequently
e ecidia grow habitually in circles round little groups of
ermogones ; a remarkable instance is seen in Phragmidium
sbi-idaei (= P. gracile); see Fig, 225.

(6) The most likely theory of the evolution of the Uredinales
that which places the majority of the micro- (including the
to-) forms as the most recent.

It is just in these, and in no others, that the spermogones
2 least often to be met with (see p. 39), as would be expected
they are furthest in descent from the primitive forms in
iich a true act of fertilisation occurred.

(7) If, on the other hand, we look upon the spermatia as
nidial forms, 7.e. as merely an additional means of vegetative
iltiplication, we are confronted by this difficulty (as well as
ose referred to above) that they appear just at that period of
velopment at which they are least wanted, whilst they are
ssing in many micro-forms where additional help would be
ost welcome. The xcidiospores have been shown in many
wys to possess an unusual amount of vigour and to be able to
oduce a stronger infection than the uredospores, which stand

NUCLEAR DIVISION 25

next in order. Even, therefore, if the spermatia could produce
an infection, their feeble aid would be wasted at such a time of
rejuvenation.

(8) There is also to be considered the fact that the
spermogones and spermatia of the Uredinales resemble those of
the Collemacez, which have been shown by Stahl (1877) and
Baur (1898) in all probability to fulfil the male function’. It
may be pointed out, in this connection, that the great similarity
of the spermogones to the pycnidia of the Ascomycetes has
been too much ignored, and that its significance is not yet fully
appreciated. In the Ascomycetes the pycnospores in most
cases undoubtedly act as conidial forms, and have lost all traces
of their primitive male function—in the Uredinales the sper-
mogones have equally lost their function, but have not taken
on the secondary réle of conidia: it may be suggested that the
latter are otherwise well provided for in that respect, and hence
feel no necessity for additional conidia. The spermatia of
Polystigma rubrum are, however, functionless either as male
cells or as conidia (Blackman and Welsford, 1912).

NucLearR DIVISION IN THE UREDINALES.

This is always of a simple type, not primitive, but reduced.
The number of chromosomes seems to be always somewhat
uncertain, and the chromatin forms masses which vary in
number from one to four. In the ordinary vegetative division,
which may be regarded as approaching rather to the nature of
amitosis, the nuclear membrane disappears, the nucleolus is
extruded, and the chromatin masses are drawn apart on a kind
of rudimentary spindle to form the daughter nuclei. In syn-
karya, the two paired nuclei in a cell are almost always in the
same stage of division at the same moment, and the four
resulting daughter nuclei move apart in such a way that the
two nuclei in each daughter cell are never sister-nuclei (Hoff-
mann, 1911).

1 A remarkable instance in Collema, though outwardly not at all resembling
the case of the Uredinales, is described by Bachmann (1912).

NUCLEAR DIVISION

In the teleutospore (ie. tetraspore-mother-cell), the first
ision is of a slightly higher type. The fusion-nucleus: is
ze, round and (when unstained) perfectly clear and homo-
1eous, but for its nucleolus, so that it looks like a vacuole:
vecupies almost invariably the middle of a cell. The dense
omatin mass is loosened out into a kind of spireme which
‘comes shorter and thicker; the nuclear membrane then
appears, and the spireme thread splits longitudinally, though
: splitting is often indistinct. It then divides transversely
o segments which become arranged or strung out on a
ndle (sometimes, but more rarely, in an equatorial plate):
mn the daughter nuclei are formed at the poles, and the next
ision, which is homotypic, follows immediately (Harper and
Iden, 1903; Blackman, 1904). Hoffmann considered that in
dophyllum Sempervivi he could count eight chromosomes
t before the reducing division.

Since each of these nuclear divisions of the teleutospore-
‘tents is usually followed at once by the formation of a cell-
ll, there are obtained four cells which are generally super-
ied inarow. But Weir(1912) records a case in Coleosporium
lsatullae where they were arranged in a “tetrad,” by which
ed he means presumably (for he gives no figures) in a
are or tetrahedral manner. :
According to Dittschlag (1910) the nucleus of the spermo-
ual hyphz is oblong and shows a slight chromatin network,
5 usually without a nucleolus. After abstriction (for which
Blackman, 1904), each nucleus enters again upon a resting
ge, and the chromatin network becomes looser. Each sper-
tium has a rather large nucleus, occupying about two-thirds
the cell, showing a decided chromatin network, but almost
ver a nucleolus. It has been frequently noticed that many
rmatia soon become binucleate, but the nuclei are sisters,
1 this condition is merely a beginning of vegetative growth
ich, however, usually aborts.

In the ecidia, the “fertile” (female) cells have a medium-
ed nucleus, with a fine chromatin network and a deeply
ouring nucleolus, as well as abundant finely granular proto-
sm. When the conjugate nuclei have arisen, they lose their

ALTERNATION OF GENERATIONS 27

nuclear membrane, the chromatin in each falls together in little
masses, the nucleolus lying by the side of the mass; these then
separate, a spindle is formed with an equatorial plate and the
chromatin masses pass at different rates towards the poles.
The nucleoli then disappear, and four chromatin balls are
formed which are the daughter nuclei. Only after this is com-
pleted, do they separate into two pairs, and the formation of a
cell-wall begins (Dittschlag).

THE ALTERNATION OF GENERATIONS.

Assuming, as we are now justitied in doing, the truth of the
foregoing ideas, we may represent the alternation of genera-
tions in a typical Uredine by the following diagram :

basidiospores

teleutospore basidium

SPOROPHYTE | GAMETOPHYTE

(2n generation)| (n generation)

. spermatium
uredospore ¢ gamete
ecidiosporé ? gametes

intercalary cell ea at

The n generation is that in which the nucleus has the
haploid number of chromosomes or, if there are no distinct
chromosomes visible, the single amount of chromatin ; in the
2n generation each cell has the double (diploid) number of
chromosomes or the double amount of chromatin, at first sur-
rounded by two nuclear membranes, afterwards by one.

28 TETRASPORE-MOTHER-CELL

The teleutospore-cell is a spore-mother-cell, exactly com-
parable with a tetraspore-mother-cell. The “ basidium” is merely
the same cell, removed outside the old outer cell-wall for the
convenience of the ensuing processes; in Coleosporium this
removal does not take place, but the “basidium” is formed
internally, the mucilaginous ‘nature of the cell-walls of the
teleutospores of that genus allowing the sterigmata to protrude
through them (see Fig. 241), which could not be done through
the hard cutinised cell-walls of the teleutospores of Puccinia
Caricis.

The first division of the nucleus of the teleutospore is
heterotypic, and really initiates the gametophyte; but, since
this is not a sufficiently definite point in the cycle, it is usual
to consider the gametophyte to begin after the next division,
Le. by the formation of the four basidium cells, which constitute
a tetrad. These four cells are the true tetraspores. In water,
deprived of air, they can each send out a germ-tube and, it is
presumable, could cause infection. The formation of basidio-
spores on sterigmata is a secondary process, viz. the production
of conidia suitable for wind-dispersion: this is shown by the
fact that, if the basidiospores are not on the proper host plant,
they can themselves germinate with the production of fresh
conidia of an exactly similar character.

The mycelium (thallus) of the gametophyte, issuing from
the basidiospores, bears male and female organs, the spermo-
gones and excidia. The spermatia disintegrate without any
result: the “fertile” cells of the zcidium (usually, perhaps
nearly always, after cutting off a sterile cell, as is the habit of
female gametes) are stimulated to further growth by con-
jugation with one another, the delay in the fusion of the nuclei
being of little or no importance from this point of view. This
so-formed fusion-cell is a zygote and is the beginning of the
sporophyte. The zcidiospores and uredospores which are borne
by this are conidial forms, devoted to increased multiplication,
and may continue indefinitely till the time arises for beginning
the cycle again. This is indicated by the dotted lines on the
left of the diagram.

It is known that a fusion of two nuclei, comparable with that

COMPARISON WITH BASIDIOMYCETES 29

which takes place in the maturing teleutospore, occurs also in
the basidium of the Agaricini and Polyporei, followed by a
division of the fusion-nucleus into four nuclei of which one
passes into each basidiospore, although it is not yet ascertained
how or where the cells of the hyphz of those Fungi become
binucleate. From this point of view it is evident that the
chief difference between the basidiospore-formation in the
Basidiomycetes and in the Uredinales lies in the fact that,
in the former, the four tetraspore-nuclei are not surrounded
by cell-walls previously to the production of conidia, as they
are in the latter.

The basidium of some of the Hemibasidiomycetes, e.g. of
the Auriculariez, is divided into a row of four super-imposed
cells of an exactly similar character to that of Puccinia, each
cell also giving rise to a basidiospore on a sterigma in the
same way. The similarity of this basidium to that of Coleo-
sportum is not diminished by the fact that it also is surrounded
by a gelatinous mass through which the sterigmata protrude.
Had not this primitive mode of forming the conidia been
modified into that typical of the Agaricini, there would have
been no opportunity for those wonderful and intricate contri-
vances for facilitating spore-dispersal which Buller has pointed
out (1909) and which find their highest and latest development
in Coprinus.

The Uredinales must be considered a highly organised
group of comparatively recent evolution, as is evident also from
their exceedingly complex parasitism. They are not a stage in
the evolution of the ordinary Basidiomycetes, but the end-group
of a different branch.

Nore. Werth und Ludwigs (1912) showed that the teleutospores of Puccinia
Malvacearum arise by the conjugation of two basal cells like those of an ecidium,
but usually of unequal size. The nucleus of the smaller cell passes into the
larger; the fertilised cell then forms, by conjugate division, a short chain of
binucleate cells, of which the two upper become the teleutospore.

CHAPTER ITI

SPORE FORMS OF THE UREDINALES

ANCIDIUM.

Acidia are usually of a cup-like shape, partly embedded in
the host, and with the free protruding edge more or less
recurved. This is the typical and presumably the most highly
evolved form. In it the spores are at first completely enclosed
by a firm structure, the peridiwm, the cells of which have the
membrane thickened on the inside wall or the outside or both,
and are arranged in very definite rows like the spores. But
there are a number of variations on this type, most, if not all,
of which belong to a lower stage of evolution. Sometimes the
peridium has the cells less definitely arranged in rows, and
therefore opening more irregularly ; at other times the peridium
is thin-walled and delicate, and in that case it usually opens by
a rounded pore and the edges do not roll back. In Hyalopsora
and its ales such a peridium is formed round the uredo-sori,
and there are reasons for believing that in these cases the pore
arises just beneath a stoma. A still simpler stage is represented
by those cases where there is no definite peridium at all, but
merely a surrounding circle of paraphyses which in a few cases
are almost or even totally non-existent. This is called a Ceoma
and indicates a more primitive form ; it is found in Phragmidium
and Melampsora. In the non-British Gymnoconia Peckiana
(= Puceinia Peckiana, on Rubus) peridium and paraphyses are
both entirely absent.

Again, if there is a definite peridium, it need not have the
shape of a cup. It may be elongated-cylindrical, straight or
curved like a horn: this is called a Restelia and is confined to

FORMS OF ACIDIA 31

the genus Gymnosporangium, on the Pomacee. Or it may be
oblong or globular, and more or less inflated: this is called a
Peridermium, and appears to be confined to the Conifers: (leaves
and stems) as hosts and to belong only to the genera allied to
Coleosporium and Cronartium. None of the Peridermia have
been found in Australia. It is a remarkable fact that ecidia
are never found upon Juncacee and Cyperacez, nor upon
Graminee with only two exceptions—the xcidium of Uromyces
Danthoniae, on Danthonia in Australia, and of Puccinia grami-
nella, on Stipa in North and South America. In all other
Uredines parasitic on these families, if ecidia, enter into the
life-cycle of the fungus at all, they are formed upon some broad-
and thin-leaved Monocotyledon, or upon a Dicotyledon, usually
though not invariably belonging to one of the more specialised
orders and above all to the Composite.

The essential characteristic of the «cidium is that its
spores are produced in chains from a fusion-cell, as described
in Chapter I. The spores themselves are always unicellular,
mostly with orange contents, and separated by intercalary
cells. Their customary polygonal shape arises entirely from
crowding and their verruculose sculpture presents a remark-
able similarity through all the group. The cells of the peridium
of the higher types are homologous with the ecidiospore-
mother-cells, and represent a division of labour for the sake of
protection: the paraphyses and the lower forms of peridium
are not of the same character, and may have had a somewhat
‘different origin. All ecidiospores, except those of Endophyllum,
germinate conidially as in P. Caricis and the germ-tubes enter
the host through a stoma: the germ-pores are numerous and
almost always indistinct.

When there are secondary ezcidiospores, i.e. such as arise
from the germination of a previous xcidiospore, they always
take the place of uredospores. In such cases, only the primary
eecidiospores arise from fusion-cells, and are accompanied by
spermogones. There is said to be one case where ecidiospores
are uninucleate, and thus comparable with azygospores, but
further evidence is required before this statement gan be
accepted (Moreau, 1911).

32 SPERMOGONES

SPERMOGONES.

While ecidia are always subepidermal, and in the higher
forms sunk rather deeply in the tissues of the host, the
spermogones are in certain genera subcuticular and in others
subepidermal, but always shallow. This doubtless corresponds
to the primitive form of the ecidium, when the trichogynes
protruded through a stoma in order to catch the spermatia.
When fertilisation was dropped, it became possible for the
wcidia to be surrounded for protective purposes by a continuous
and firm peridium and to be more deeply sunk within the host:
the spermogones, being outside the range of selective evolution
on account of their uselessness, have retained more or less of
their original character. The description of those of P. Caricis
applies essentially to nearly all, except that in certain genera
they are flat, not flask-shaped, and open by a wide pore, not
by an ostiole, and in these and others there are no protruding
filaments.

Spermogones never appear alone; they are always accom-
panied or closely followed by some other spore-form, either
ecidio-, uredo-, or teleutospores. In comparatively few cases,
as in P. Malvacearwm, have the spermogones disappeared en-
tirely.

UREDOSPORES.

Uredospores are usually distinguished from wcidiospores by
being produced singly at the apex of a short pedicel from
which they easily fall off: this pedicel is the homologue of the
intercalary cel] of the ecidiospores, as was shown by Christman.
In certain genera, however, e.g. Coleosporium and Chrysumyxa,
the uredospores are produced in short chains. Uredospores can
reproduce uredospores for an indefinite number of generations.
Occasionally there are two kinds, primary and secondary uredo-
spores; in such cases the primary ones arise from fusion-cells
and take the place of ecidiospores. So far as is known true
uredospores never arise from a uninucleate mycelium, though
they may spring from a mycelium which was at first uninu-
cleate, but became binucleate at some point in its development.

UREDOSPORES 33

This would be the case in a true Hemipuccinia, but none of
these have so far been cytologically investigated.

Uredospores are always unicellular, except in the case of
a few monstrosities. Primarily they must be considered a
device to aid in rapid propagation and hence may be called
summer-spores: for this reason they usually germinate with
great readiness when mature, always forming a tube which
enters a stoma of the host. The number of germ-pores varies
from 1 to 10 (usually 2 to 4): only one case is known in
Uromyces where they possess a single pore (U. uniporulus) and
one in Puccinia (P. monopora). They present a wonderful
sameness in shape throughout the whole group, and in colour
vary from yellow and orange to brown. It may be taken as
a general rule that if the wall is colourless, the contents are
yellow or orange from an abundance of that yellow oily
substance which occurs in ecidiospores ; if the wall is distinctly
brown, the contents are often colourless when mature, though
at first they frequently contain the usual yellow oil. It is only
in a few. instances, in the lowest genera, that uredospores are
quite without colour. The uredospores of Puccinia dispersa
and in a smaller degree of P. graminis are noticeable for a
curiously dull appearance which is very characteristic, because
they combine orange contents with a brownish membrane.
The outer wall of uredospores is almost always covered with
spines (echinulate), needles (aculeate) or warts (verruculose) ;
it is very rarely smooth; these projections enable them to cling
more readily to the surface of the host. They are often inter-
mixed with paraphyses, which are usually clavate or capitate
in shape; these are found in a few species of Puccinia, but
more especially in Melampsora and Phragmidiwm. These
paraphyses are homologous with the spores, being binucleate
at first, but the nuclei soon disintegrate.

There is very frequently found, in the uredo-sori of many
species a parasite belonging to the Deuteromycetes, called
Darluca Filum. It consists of a black pyecnidium, enclosing
numerous uniseptate pycnospores which are faintly apiculate:
at each end. This has been sometimes mistaken for another
spore form of the Uredine.

G. U. 3

34 AMPHISPORES

AMPHISPORES.

In countries which are arid or semi-arid, there is found in
certain species a form of uredospore which has led to several
mistakes owing to its misleading appearance. The spores are
provided with a thick cell-wall or have the summit thickened
conically, after the style of many a teleutospore of Uromyces,
and are supported on a persistent pedicel, so that one would
not take them for uredospores; nevertheless they will be found
to have more than one germ-pore and to germinate by a germ-
tube, although only after a period of rest. These were named
by Carleton amphispores ; they were first discovered in Puccinia
vexans. They are evidently a provision to enable the spore to
pass through an unfavourable period unharmed, and reinfect a
host of the same species when occasion arises. The amphi-
spores of Puccinia atrofusca Holway, though echinulate and
possessing two equatorial germ-pores, were first described by
the discoverers, Dudley and Thompson, as the teleutospores of
a Uromyces, and the same thing has happened in other cases,
eg. Puccinia convolvulr, P. tosta, and P. cryptandri. The
nearest approach to amphispores found in
British species is in P. Pruni-spinosae (Fig.
22), in which they have been mistaken for
paraphyses and were so figured in a well-
known text-book. There is another kind of

spore, presently to be described, called a

Hig, ee es mesospore, which bears a superficial resem-

spinosae. x600. blance to an amphispore. It isa mistake to

call amphispores a transition-form between

uredo- and teleutospores, since they are of later evolution than
the two latter.

TELEUTOSPORES.

The meaning of the word teleutospore is end-spore ; it was
considered to represent the stage when growth was ceasing for
the season. This is not the case, however, in all species, and
the word must now be used with another connotation, viz. a
teleutospore is one which germinates by the production of a
basidium and basidiospores.

TELEUTOSPORES 35

The teleutospores are generally produced in sori like those
of the uredospores; they frequently arise on the same mycelium,
and very often on the same spore-bed, mingled with the uredo-
spores. If both are found in any species, the teleutospores are
always formed at least not earlier, and usually later than the
uredospores. Their primary function now is to tide over an
unfavourable period; for this reason they are sometimes called,
in England, winter-spores. They may be one-celled as in

Fig. 23. Figures of various Teleutospores of Pucciniaces (after Sydow). From
left to right, they are (in the top row) Puccinia roesteliiformis, P. conspersa,
P. globosipes, P. Megatherium, Uromyces globosus, Hapalophragmium
Derridis; (in the second row) P. appendiculata, P. Euphorbiae var, intu-
mescens, P. deformata, U. achrous, U. giganteus, U. Ipomoeae, Triphragmium
Cedrelae. All x 480. (Non-British.)

Uromyces, two-celled as in Puccinia, radiately three-celled as
in Triphragmium, cruciately four-celled as in Pucciniastrum,
linearly many-celled as in Phragmidium and especially in
Xenodochus (see also Fig. 23). But it is misleading to speak
of a many-celled spore: each cell, taken separately, is a spore
and can germinate by itself without reference to the others.

3—2

36 TELEUTOSPORES

In the formation of teleutospores in the higher Uredinales,
the spore-mother-cell first divides into an upper fertile cell and
a lower sterile cell, which elongates more or less to form the
pedicel, The upper cell may remain undivided, or may divide
again: the lower of these two may then continue to divide and
so on, to form a many-celled chain. When the chain is long, as
in Xenodochus, it is seen very clearly that the spores are formed
like eecidiospores to this extent that the uppermost is always
the most mature. This may be taken as a sign that they are
modifications of zcidiospores to form resting-spores. In Hndo-
phyllum the sxcidiospores previously mentioned germinate as
soon as mature with a basidium, and are therefore teleutospores
also: this is the primitive state of things from which the
present wide division of labour into rejuvenating (ecidio-),
multiplying (uredo-), and resting (teleuto-) spores has been
evolved.

In some of the lower Uredinales, the teleutospores are
formed beneath the cuticle or in the epidermal cells, but the
usual position is directly beneath the epidermis. Throughout
the whole group the colour of teleutospores is almost uniformly
brown, varying in shade from a pale yellowish-brown up to
nearly black. Their contents are, like those of uredospores, at
first often oily and yellow, afterwards colourless. In the lowest
genera, those found on Ferns, the teleutospores are quite
hyaline.

Their surface is most often smooth externally, but occasion-
ally marked with superficial unevennesses, such as warts,
tubercles, lines, striw, reticulations, and pits; a few have spiny,
papillose, or finger-like processes, either at the summit or all
round. The majority of them have one pore to each cell, as in
Puccinia and Uromyces, covered at times by a distinct, often
hyaline, pore-cap; this is the highest type, being furthest
removed from the many-pored ecidiospores. Other genera
have 2 to 4 germ-pores to each cell, as in Phragmidium and
Gynmosporangium. In some cases, as in Uromycladium and
Ravenelia, the teleutospores are borne in bunches at the top of
» common stalk, either with or without accompanying hyaline
systs, i.e. abortive spores. An approach to this is found in the

“BASIDIOSPORES 37

British Puccinia Pruni-spinosae and other species, where the
short pedicels are all closely bound together in bunches at the
base. Paraphyses are naturally not so common in teleuto-sori
as with uredospores, since the former do not need such protection,
but they are found in P. Sonchi, P. dispersa, P. persistens, etc.,
although in these cases the so-called paraphyses are not at all
of the same character as those found in uredo-sori, e.g. of Puc-
cinia Baryi and others. When the teleutospore of a normally
two-celled species becomes one-celled (by the omission of the
last cell-division), it is called a mesospore: the mesospores of
Puccinia are practically identical with the teleutospores of
Uromyces and germinate like them.

BASIDIOSPORES.

All normal teleutospores develop under natural conditions
in the same way; the cell-contents divide themselves into four
parts, by a heterotype followed immediately by a homotype
mitosis.

This formation of what are really (and might with ad-
vantage be called) tetraspores can take place in two ways :—the
“basidium” can arise within the teleutospore-cell or outside it.
The first method is the more primitive, the second is an
adaptation to the tough cutinised or chitinous exospore of the
more advanced types. In the Coleosporiacez the teleutospore,
ie. the tetraspore-mother-cell, divides into four superposed cells
(like the tetraspores of Corallina) while still in the sorus, during
the autumn; each cell (spore) germinates, in late autumn, by
protruding a sterigma through the thin gelatinous wall of the
teleutospore and forming a basidiospore (conidium) at its apex.
Zaghouania shows an intermediate form of germination. But
in all the other families the cell-contents of the teleutospore,
clothed only in a thin endospore, pass out through a germ-pore
in the form of a longer or shorter tube (“basidium”); the
contents pass to the distal end of this, and are there divided
into four oblong cells. The median septum is sometimes formed
first, and the two lateral ones after. In water the “basidium ”
is usually long, in air it is short. In the absence of sufficient

BASIDIOSPORES -

, moreover, the four cells may not be formed ; the “ basidium ”
wy resemble, more or less, an ordinary germ-tube and possibly
iy function as such. Even if the four cells are formed, they
ay germinate by the protrusion of a germ-tube, which pre-
mably can cause infection by penetrating the cuticle (see
vons, 1912, p. 225). But, with access of air, each cell forms a
srigma and a basidiospore as previously described. These
nidia are obviously adapted for wind-dispersion. If they
ght on a suitable surface, they send forth a short tube which
variably bores straight through the cuticle into the under-
ing epidermal cell of the plant, and there begins to form
mycelium. The only instance in Puccinia known to the con-
ary?, out of the many observations that have been made and
rured of this process, is De Bary’s record of the case (Ann.
wi. Nat. Bot. 4. xx, 1863, pp. 88-9) where the germ-tubes of the
basidiospores of Puccinia Dianthi
(q.v.) penetrated through the stom-
atal openings of Dianthus barbatus
(Fig. 24). In grasses and sedges, it
is easy to see that the siliceous
we cuticle would present a great ob-
Pads oe peat ia stacle to the entry of such a tube,
ee Play trian while not impeding germ-tubes
ing straight for the stomata which enter through a stoma, and
(after De Bary’s figure). 390. this is probably the reason why
cidia are so rare on the order Glumiflore. In the hetercecious
yecies no one has yet brought forward ipdisputable evidence
» show that basidiospores can infect the host which bore the
‘leutospores, although statements to that effect are made.
The shapes of basidiospores are not irregular; they are
iore or less constant in each group. In Puccinia and Uromyces
1ey are ovate, somewhat flattened on one side, or kidney-
raped. In the Phragmidiee they are almost spherical; in
1e Melampsoracee small and roundish. In Endophyllum they °
re ovate; in Coleosporium they are large and ovate and a

1 Statements have been made of other instances, but most of them on
sufficient authority, and some have been specifically disproved. The genus
oleosporium seems, however, to be an exception.

GROUPS OF SPORE-FORMS 39

little flattened on one side, while in Ochropsora they are
spindle-shaped. This is one of the reasons for suspecting that
it is probably incorrect to classify these two latter genera in
the same family, on the basis of the “internal” basidium
merely. The genus Chrysopsora, which has the same kind of
basidium, belongs to the Pucciniacee.

GROUPING ACCORDING TO SPORE-FORMS.

For certain purposes it is convenient to have names for the
groups into which the Uredinales may be divided according to
the number of spore-forms possessed by each, though it must
never be forgotten that such a grouping does not in any way
indicate affinity. The method usually employed is that put
forward by Schréter, which (with a little modification according
to present ideas) may be presented in the following scheme.

Denoting spermogones by O, ecidia by I, uredospores by
II, and teleutospores (with the ensuing basidiospores) by III,
we call a fungus possessing

[ee if with all four on one
lant.

O TALTE |: Be eer, if O, I on one species,

and II, III on another.

OI III | an -opsis-form.

O ILIII | a Brachy-form.
JI III | a Hemi-form (in many cases perhaps the half of a

Hetereu-form).

[O] III | a Micro-form (spermogones sometimes absent).

Thus Melampsora Rostrupii is a Eumelampsora, Gymno-
sporangium Sabinae is a Gymnosporangiopsis, Uromyces Anthyl-
lidis is a Hemiuromyces, and Puccinia Campanulae is a Micro-
puccinia. A Leptoform is one, of whatever kind, in which the
teleutospores germinate as soon as mature, without any resting
period; thus the spores of P. Malvacearum belong chiefly to
the lepto-form, those of Endophyllum Sempervivi entirely so.

Maire (Progress. Rei Botan. 1911, iv, 115) has proposed a
much more complex arrangement on the same lines, which is
quite needless and, it is to be hoped, will be quietly ignored.

40 GROUPS OF SPORE-FORMS

The relations of the various groups to one another ar
represented in the following diagram. The circle represent
the Eu-forms; the substitution of any one chord in the plac
of the arc which it subtends shows how the life-history i
shortened in the other cases. Only the abnormal Endophyllun
cannot be included in such a scheme; its spore-grouping coulc
I)

only be represented by the symbol O 1" |

basidium

teleutospore

-opsis forms

uredospore

uredospore

mycelium
eecidiospore

fusion-cell

CHAPTER IV

LIFE-HISTORIES OF OTHER UREDINALES

Puccinia graminis.
THE Biack Rust or “MILDEW” oF CORN.

Another common Puccinia, whose life-history is of greater
economic importance than that of P. Caricis, is the well known
P. graminis, the Rust or Mildew of Corn. This species has its
spermogones and ecidia on Barberry (Berberis vulgaris and
Mahonia Aquifolium) and its uredo- and teleutospores on many
species of grasses, especially on cultivated wheat. It must not,
however, be assumed that any rust found on wheat is P. gra-
minis; there are at least two other species common on the
same host which, unless carefully examined, may be confounded
with it, viz. P. triticina and P. glumarum, not to mention a form
of P. coronata which sometimes also oécurs on cereals. For
this reason these four species are now distinguished as the
Black Rust, Brown or Orange Rust, Yellow Rust, and Crown
Rust of corn, respectively. The uredo-stage of P. graminis is
known as Red Rust.

If one merely substitutes Barberry for Nettle and Wheat
for Carex, what has been said about P. Caricis is true in all
essentials of P. graminis. The differences are not in the life-
history, but in ‘certain minor details of occurrence: e.g. the
spots caused on the leaves of Barberry are small, round and
red, while uredospores of P. graminis are most common on the
leaves, and the teleutospores form long black striz on the

42 PUCCINIA GRAMINIS

culms; for these differences the systematic part can be con-
sulted (Fig. 25).

There is, however, one point

of difference connected with P.

graminis which possesses great

biological interest—its virtual in-

dependence of the ecidial stage.

LJ For a long time it had been

known that Barberry bushes in

the hedges caused “mildew” on

the corn in the neighbouring

c fields, and when, in 1864-5, De

\
Bary proved the hetercecism by

i | experimental cultures, it was too

Big. 25. Puccinia graminis. a, hastily assumed that the ecidium
ecidia on Berberis; b, uredo- on the Berberis was just as es-
spore ; c, teleutospores. :

sential to the rust on the corn as
that on the Nettle is to the rust on the Sedge. Many facts
now tend to show that this is not the case.

In Australia and the plains of India the Barberry is un-
known except as an introduced plant, yet the Puccinia occurs
everywhere and does enormous damage. McAlpine records, in
his Rusts of Australia that he made numerous attempts to
infect imported species of Berberis with the rust of Australian
wheat which is morphologically undistinguishable from the
P. graminis of Europe, but all his efforts were in vain. The
inevitable inference is that P. graminis, as it occurs in those
countries, is a “ biological” race which maintains itself by other
than the primitive means. A similar thing is true, according
to Lagerheim, in Ecuador, where also rust flourishes and does
great harm. ;

The facts now known concerning the specialisation of the
Black Rust are treated of in a separate chapter, but there is
one point which must be mentioned here. This concerns the
mode by which fresh epidemics are produced each year. Even
if the Barberry is present, it is by no means certain that it
plays any important part in these annual attacks. Apart from
that, there are several possibilities: (1) the fungus may winter

ore ‘bd
3. a

PUCCINIA GRAMINIS 43

by its uredospores, (2) by a perennial mycelium, (3) by Eriks-
son’s mycoplasm.

The first possibility is entirely a matter of climate: it may
take place in one country and not in another, or in the same
country it may take place in one season and not in others.
McAlpine and Cobb find viable uredospores all the year round
in Australia, and Lagerheim says the same for Ecuador. But
in northern climates it has been shown that the uredospores of
P. graminis frequently lose all capacity for germination during
the winter; this is proved true of Sweden, North Germany,
North Dakota, etc., but in the United States, south of Ohio,
Bolley found germinable uredospores all through the year.
Similarly in Bohemia, uredospores of P. dispersa, P. glumarum
and P. Loli can survive mild winters or in sheltered places
(Baudys). Even though uredospores capable of germination
may sometimes be found on wild grasses during the winter, it
does not follow that those could start an epidemic next spring,
owing to the specialisation which has been proved to exist, by
which a form of P. graminis on one host is often incapable .of
infecting another host.

In regard to the second possibility, we find again two
opposing views. De Bary and others have searched in vain for
mycelium in the growing wheat plants, before infection becomes
visible, but Pritchard (1911) found mycelium resembling that
of P. graminis both in the pericarp of wheat grains and in
various parts of wheat seedlings. He showed that large num-
bers of wheat grains contained pustules of teleutospores, even
visible in the neighbourhood of the hilum, but also hidden
within the pericarp. He proved that the mycelium from the
pericarp penetrates through the intercellular spaces, as well as
through the cells, and “soon passes into the spaces between
the leaf-sheaths where it grows rapidly and attacks the tissues
at various points.” W.G. Smith figures teleutospores within
the seed of Oat (Gard. Chron. 1885, xxiv, 245, f. 58) and ecidia
in the pericarp and seed of Barberry (ibid. 1886, xxv, 309,
f. 58).

It is evident that, if this state of things prevailed on a
large scale, nothing more would be required to explain the

44, PUCCINIA GRAMINIS

origin of outbreaks of rust. It is not inconsistent with this
that Ward was able to prove that the mycelium of a uredo-
sorus extends only a little way round the margin of the sorus;
that may be and is true in certain cases, especially with regard
to secondary uredo-sori, but in P. Caricis the mycelium extends
up and down the leaf between the parallel vascular bundles,
producing uredo-sori all along its course. The practical bearing
of Pritchard’s discovery is to show that seed from an infected
crop should never be used for planting.

About the third possibility, it is difficult to come to any
definite conclusion. Eriksson’s hypothesis is that the proto-
plasm of the fungus is present in the grain, mixed with the
protoplasm of the host, in such a way that’ the two are
indistinguishable. As the plant grows up, he supposes that
the two grow together until, at a certain time, the protoplasm
of the fungus separates itself from that of its host in the form
of “Nucleoli,” passes into the intercellular spaces through
“invisible pores,’ then or earlier surrounds itself with a cell-
wall, forms a mycelium, and begins its ordinary life by producing
uredo-pustules. An intermediate stage, where the fungus-
protoplasm has surrounded itself by a cell-wall but is still
enclosed within the cells of its host, he named “special cor-
puscles.”

The difficulty in dealing with this theory lies in its in-
definiteness ; its author changes it from time to time to meet
objections, and supports it by hazy microscopical observations,
many of which are demonstrably the result of incorrect vision.
His “special corpuscles” have been shown by Ward and Klebahn
to be ordinary haustoria, Eriksson having completely overlooked
the intercellular hyphe to which those haustoria were attached.
It is incredible that the protoplasm of so highly evolved a fungus
could live outside its cell-walls, as he supposes. Such a state
of things is, of course, common in the lower fungi, Chytridinge
and allied groups. In Synchytrium Solani the fungus-protoplasm
and the host-protoplasm may be seen in the same cell, before
the latter has been completely devoured by the former, and in
that state they are even distinguishable by their microscopic
appearance. But it will need a great deal more “ proof” before

PUCCINIA POARUM 45

Eriksson’s startling hypothesis can be accepted in regard to
such ~ fungi as the Uredinales.

The futility of Eriksson’s mode of argument is seen in his
suggestion (1908) that “other diseases such as the American
Gooseberry Mildew can live within the infected shoots in a
form scarcely visible to our eyes.” But direct evidence against
the Mycoplasm Theory is accumulating. Jaczewski (1910)
grew seeds obtained from many much-rusted plants, but he
found that, when they were sown under glass and protected
with adequate care from all outside infection, they all produced
rust-free plants. Bolley, Linhart, Zukal and Klebahn had
similar experiences. Zach (1910) on investigating leaves and
culms of Rye, infected with P. graminis and P. glumarum,
found on the outskirts of the infection-patches all the states
described by Eriksson, but he proved that in all of them fungal
hyphe were present. In fact, Eriksson himself saw and repre-
sented these hyphe, but calls them “radialen Strange ” of his
supposititious “ Nucleoli,” the said “ Nucleoli” being merely the
deformed remains of the nucleus of the attacked cell. As
Marshall Ward (1905) remarked, Eriksson merely inverts all
the stages of a fungus attack on a cell, and supposes the last
state to be the first. This error and a misinterpretation of
the microscopic appearances account for the whole wearisome
persistence in an inherently improbable hypothesis.

Puccinia Poarum.
THE CoLTsFooT AND MEapow Grass Rwst.

This species is economically of no importance; its spermo-
gones and ecidia occur on the common Coltsfoot (T'ussilago
Furfara), and its uredo- and teleutospore stages on species of
Poa, to which however they do little harm. Here, again, the

1 This does not accord with Eriksson’s experience; but then on some of his
“protected” plants aphides also made their appearance, yet this does not seem
to have suggested to him that the zooplasm of the aphides must also have been
latent in the seed! If the aphides got in, so would fungus spores, since it has
been proved (Butler, 1905) that uredospores are carried by them and other.
insects.

46 PUCCINIA POARUM

life-history is in all essentials identical with that of P. Caricis,
but it differs in one striking particular—there are two genera-
tions of each stage during the year.

The spermogones and ecidia first appear on the leaves of
Coltsfoot in May and June, and are followed by the Puccinia
on neighbouring leaves of Poa in July and August. Then a
second crop of ecidia, also accompanied by spermogones, appears
from end of July to September, followed again by uredo- and
teleutospores in September to November. The latter rest
during the winter and infect the young Coltsfoot leaves again
in the following spring. In countries that have a climate
favourable for the growth of Poa, the uredospores may be found
the whole year round, and the fungus can maintain itself by
them alone. This is certainly the case in Australia, according
to McAlpine, where the Coltsfoot does not exist, and the uredo-
stage is most common in the winter months, 1e. June to
September. In this country, the teleuto-sori are rather incon-
spicuous, but can be found by searching carefully on the lower
leaves of species of Poa round the spot where the Coltsfoot has
been found affected by the ecidium, especially in July and
August.

The zcidium of this species has been examined cytologically
by Blackman and Fraser, and according to them the binucleate
condition of the fertile cells is produced by the migration of the
nucleus from one fertile cell to an adjoining one in the hymenial
layer, and also occasionally by a migration from one vegetative
cell to another at a point where the conjugating cells were
below the level of the hymenium, although only a little lower.
Cells with three or four nuclei were met with by them, and true
conjugate division was observed in such cases.

In sections which I have examined of this species, I have
seen evidence which seemed to indicate (although not with
perfect, certainty) that conjugation also took place by the
removal of a large part of the intervening cell-wall, and a con-
sequent fusion of the cells as described by Christman and
Dittschlag (Fig. 26).

As will be seen from the figure, the cells of the peridium of
this species differ slightly from those of P. Caricis in their shape,

PUCCINIA POARUM 47

as shown in radial section. The outer edge of each cell is more
prolonged downwards so as to overlap a portion of the cell
below in an imbricated manner. The xcidium, in both, as is the

{

Fig. 27. P. Poarum. An excidio-
spore germinating on leaf of
Poa annua. x 180.

Fig. 26. P. Poarum. Vertical section
through edge of excidium; showing a,
the crushed cells of the upper myce-
lium, pushed on one side ; b, the chain
of peridium-cells; c, the spore bed,
giving rise to d, the chains of scidio-
spores. x 600. The black dots are
the nucleoli; one #cidiospore-mother-
cell has three nuclei.

case in most of the higher forms
of the Uredinales, arises deep Fig. 28. P. Poarwm. u, an weidio-
- spore germinating in water,
in the leaf, and the densely x 250; b, the same, showing
packed knot of hyphe which — he germ-pores, x 500.
forms the starting-point of the hymenium, where the cells are
full of protoplasm, is covered over by a number of nearly empty
cells which are ultimately squeezed to the side by the develop-
ing eecidium and are shown in the figure at a.

It is easy to produce the teleutospores in a garden on a tuft

of (say) Poa annua or P. pratensis, by planting quite near to it

3 PUCCINIA MALVACEARUM

id overhanging it some Coltsfoot affected by the xcidium:
iother tuft about ten yards off can be used as a control. The
‘edo- and teleutospores will appear on the former tuft in about
k to 21 days. If this is done late in the year (September)
have found that only teleutospores are formed on the leaves
‘the Poa. For the germinating ecidiospores see Figs. 27, 28,

Puccinia Malvacearum.

THe HoLiyvock Rust.

This Rust differs from all the others that will be mentioned
the simplicity of its life-history, and also in the fact that it
not confined (as almost all the others are) to one kind of
plant or even to a few, but ap-
pears, so far as is known at
present, to range over the greater
part of a sub-family. It has
been found on over 50 species

of the Malvez, and it seems to be
identically the same in every case.
The mycelium develops in
spring in the intercellular spaces
of the young leaves and stems
and produces little knots under
the epidermis, on which a thick,
round, hard, pale-reddish cushion
of teleutospores is formed. These
spores have short or very long
pedicels according to their posi-
tion; they are mostly typical and
two-celled, but mesospores with
only one cell are not uncommon,
and occasionally a few may be
met with having three or even
fig. 29. P. Malvacearum, Ger- four cells, Most of them germi-
minating spore; a, basidium nate at once, in the sorus, pro-

breaking up into separate cells; 7. oes ‘
b, a basidiospore, x 600. ducing basidiospores in the usual

belonging to nearly all the genera |

4
@
5

PUCCINIA MALVACEARUM 49

way (Fig. 29). The accumulated basidia and spores give a
greyish tinge to the red-brown sorus. These spores can cause
fresh infection and so the disease spreads rapidly. It is most
active about the end of summer, and has often been the cause
of a serious epidemic on the more susceptible kinds of Holly-
hock. The sori are found on every green part of the plant,
stems, leaves, petioles, bracts, sepals, carpels and fruits, and are
even reported on the petals. There are no uredospores.

The chief biological interest of this fungus concerns the
way in which it passes the winter, a point about which there
has been much dispute. There are two possibilities, (1) by
perennial mycelium, (2) by over-wintering teleutospores. The
first has been strongly advocated, and it is very likely (though
one can hardly say it has been proved) that the mycelium does
winter in the young leaf-rudiments that are formed on shoots
at the base of last year’s stems. Freshly formed sori have also
been found on the cotyledons of seedlings which grow up in
late autumn round the parent plants and which in certain
cases can survive the dead season. But there seems to be
absolutely’ no justification for the claim that the mycelium
winters in the embryo of the seed. The disease can undoubt-
edly be carried with the seed, in sori either on the bracts
(portions of which are often mixed with the “seeds,” ie. fruits)
or on the outside of the carpels themselves.

Eriksson has lately (Ueber den Malvenrost, 1911) published
a theory, similar to his well-known theory about the Rust of-
Corn, and standing or falling with it: he says that P. Malvace-
arum perennates in the form of “mycoplasm” in the cells of
the autumn buds at the base of the shoots, as well as in the
embryos of the seeds of the infected plants. With these he
says it grows up in an imperceptible form, mingled with the
protoplasm of the host, permeating the newly-formed leaves
and at last suddenly breaking out in the form of pustules of.
primary teleutospores, which afterwards spread in the acknow-
ledged way. He explains the presence of this mycoplasm. by
stating that certain teleutospores of the previous autumn
germinated by sending out “ germ-tubes” which cut otf “ end-
conidia.” (This mode of germination of the late-formed spores

G. U. 4

50 PUCCINIA MALVACEARUM

of P. Malvacearum is well known, though not usually interpreted
in that way, see Fig. 29.) These “ end-conidia” do not form a
short tube, to penetrate the cuticle of the host, but “ pour forth
their protoplasm, as it seems, without the formation of an
opening, through the plasma-connections of the outer wall of
the epidermis of the host into an epidermal cell,” and so into
the tissues where it vegetates till required. It also exists in
the same state in the seeds of the infected plants. The fungus,
he says, “passes from the plasmatic into the filamentous state
just before the outbreak of the primary pustules.” It is clear,
however, that the figures he gives do not prove what he asserts.

Putting aside this purely supposititious and intangible
method, the chief means of perennation probably lies in the
fact that certain teleutospores produced at the end of the
growing season have the power of lasting through the winter
and germinating in the spring. Plowright, Massee and Tau-
benhaus all agree in this: the latter (1911) kept infected
leaves, gathered at Cornell University in the United States
from the living plant on November 26th, both indoors at a low
temperature and outdoors, and by testing spores taken from
them at intervals from December to April found that they still
remained germinable, though more and more slowly as time
went on.

Dandeno, however (9th Report Mich. Acad. Sci. 1907, p. 68),
states that the fungus does not winter in the seeds; he tried
seeds of diseased plants, carefully excluding infection from
outside, and found that they all produced healthy plants. His
experience also was that no teleutospores remained viable till
the next spring, but that the fungus maintained itself the
whole winter through on mallow plants in sheltered spots.

These differences may be partly a matter of climate, and as
regards the “seeds,” unless there were sori on them, they could
hardly be supposed to carry the infection, even if they came
from infected plants, except by the presence of “mycoplasm”
or mycelium, neither of which has been proved.

For this reason the chief means of preventing the disease
(apart from using “seed” from uninfected plants) must be to
gather and burn all dead leaves from the infested bed. When

peer erer

GYMNOSPORANGIUM CLAVARIZFORME 51

the disease does appear, spraying with Bordeaux mixture is the
best remedy against its spreading. It is the very young shoots
and the upper side of the leaves that most require spraying ;
although the pustules appear chiefly on the lower surface of
the leaves, there can be little doubt that infection by the
basidiospores takes place mainly through the upper surface.

Gymnosporangium clavarieforme.

THE HAWTHORN AND JUNIPER RwsT.

This fungus produces its zcidia on the branches, leaves and
fruit of the Hawthorn and on the leaves of the Pear, and its
teleutospores (there are no uredospores) on branches of the
common Juniper, where it causes fusiform swellings. There
are three other allied British species, of which G. Sabinae also
attacks the Pear, but its teleutospores are formed only on
Juniperus Sabina (the Savin Juniper).

On the swollen branches of the Juniper, the parasite pro-
duces in April and May numerous orange masses, which ooze
out and sometimes reach more than
1 cm. in height. They vary in
shape, but are usually more or less
conical or tongue-shaped (Fig. 30).
They consist of a mucilaginous
mass in which large numbers of
teleutospores are embedded. The
mucilage is produced by the gela-
tinisation of the cell-walls of the
fungus, especially of the very long
pedicels; it naturally swells and
becomes more conspicuous in wet
weather. The teleutospores ger-
minate at once, while still in the His Sh. Byanasnatoupin Bie
mass; the basidiospores are pro- variaeforme. Masses of teleuto-

: te spores on branch of Juniperus
duced in the usual way; it has communis (slightly reduced) ;
been shown lately in another species _‘*w° teleutospores, x 600.
(Coons, 1912) that they do not fall off, but are jerked off the

4—2

r~—

GYMNOSPORANGIUM CLAVARIZFORME

wigmata, much in the same way in which the basidiospores
> thrown off in the Agaricini. They accumulate in large
mbers on the outside of the mucilaginous mass, and present
> appearance of a golden-yellow powder. The mycelium of
2 fungus is perennial in the Juniper, spreading from branch
branch and producing a fresh crop of teleutospores each spring.
If one of the basidiospores is blown by the wind or carried
insects to a moist leaf or young fruit or stem of Hawthorn,
germinates and bores through the cuticle in the ordinary
y, and forms there a limited patch of mycelium. It is said
at the infection has been known to be conveyed for half

~Ss.

31. Gymnosporangium clavariaeforme. Aicidia on leaf, fruit, and branch
of Hawthorn (reduced); wu, peridium, x16. The fruit and gall on branch
are shown as they appear when the peridia are old, aud the mass looks
somewhat like » honeycomb.

mile. This mycelium produces the ecidium-stage, which is
t usually seen until the end of June, in about 18—20 days.
1e xcidia are not of the ordinary cup-like shape, but of the
m called Reestelia: they are cylindrical, brown at the base
d ochreous above where the peridium is torn into numerous
mder filaments (Fig. 31). On the leaves they appear in
undish patches a few mm. in diameter, but on the twigs
ey form large spongy masses and the fruits are often so
vered with them as to look like a cluster of little spikes.

* IT am indebted to Professor Buller for calling my attention to this fact.

ENDOPHYLLUM SEMPERVIVI 53

Within these ecidia the ecidiospores are produced; these
will only infect the Juniper, on which they begin the
cycle again.

For all these species of Gymnosporangium the only remedy
is to remove and burn the diseased Juniper, if it can be found ;
if it may not be destroyed, at least the affected branches should
be cut off, and the wounds dressed with Stockholm tar. It is
of no use to spray or otherwise treat the Hawthorn or Pear.
In them the disease is purely local; it comes to an end when
the summer ends, and will not recur next year unless fresh
infection is conveyed from the Juniper. The harm done to
them is confined to the loss of the foliage which naturally
weakens the tree to some extent.

Endophyllum Sempervivi.

THE HovuseLeEK Rust.

This parasite attacks the common Houseleek and numerous
other species of Sempervivum. It differs from nearly all the
other Uredinales in having only spermatia and ecidiospores,
the latter functioning also as teleutospores and producing basi-
diospores. This fungus has been thoroughly investigated by
Hoffmann (1911) from whom the following account has been
derived.

The ecidiospores mature on the leaves in April and May;
they have no visible germ-pores. They germinate at once,
while still in the ecidium; the germ-tube forces its way out
at some point of the circumference of the spore and elongates
to form the four-celled basidium. Each basidium produces four
basidiospores on long sterigmata; occasionally more are pro-
duced—Hoffmann observed as many as eight on one basidium.
The basidiospores may be blown on to the leaf of a Houseleek
where they germinate at once and bore through the cuticle ;
they form a holdfast (somewhat as a uredospore does) below
the outer wall and penetrate into the epidermal cell (see Fig. 17).
The mycelium then branches, passes through the intercellular

54 ENDOPHYLLUM SEMPERVIVI

spaces (sending haustoria into the cells) until it reaches the
base of the leaf; thence it penetrates into the axis and so up
to the growing point, where it hibernates till the following
year. In the spring it grows on into the freshly formed leaves
which become yellow and longer and more erect: on these,
on both sides, spermogones appear in March and April, followed

Fig. 82. Aicidia of En-
dophyllum on leaf of
Sempervivum monta-
num (reduced).

by excidia (Fig. 32) which repeat the
cycle. The affected plants are easily
recognised by the different attitude of
the leaves, which imparts an unusual

irregularity to the rosette (Fig. 33).

The most interesting point about
this species is that established by Hoff-
mann, that the ecidiospore-chain arises
in the way already described for Puccinia
Caricis from a cell produced by the
fusion of two adjacent cells of the spore-

bed, after the manner described by Christman except that the
conjugating cells were not situated in any definite plane. The
binucleate secidiospores then became uninucleate by the fusion

Fig. 33. Two plants of Sempervivum, one (left) affected by
Endophyllum Sempervivi, the other not.

of the conjugate nuclei. On germination, when the fusion-
nucleus divides into four, the first division shows slight
differences from the others so as to make it certain that it is

the reducing division,

CRONARTIUM RIBICOLA 55

The life-cycle may thus be represented as follows:
Basidiospore

Gametophyte Mycelium

(m generation) Acidium Spermogone

\ Female cells Spermatia
Fusion-cell

|
Spore-mother-cell

h
eee 4 Aicidio-teleuto-spore Intercalary cell
ome)
aa a
0 00 oOo Basidiospores °

This life-history 1s especially worthy of consideration because
it probably represents that which obtained at the first evolution
of the higher Uredinales. The various types of development,
seen in the genera Puccinia, Uromyces, etc., and described
under the names Eupuccinia, Micropuccinia and so on, may all
be derived from this original form. See Grove, New Phytologist,
1918, p. 89.

Cronartium ribicola.

Tue Rust OF CURRANTS AND FIVE-LEAVED PINES.

This disease, called the Currant Rust in one stage, and the
Weymouth Pine Blister Rust in the alternate stage, can do
enormous damage in the second phase ;"it threatens in places
to put a stop entirely to the cultivation of the Weymouth Pine.
It has been imported into England and the United States with
young trees of the latter from the continent of Europe. The
remedy is to inspect Weymouth Pines, in nurseries and plan-
tations, annually, cutting down and burning all those that
show infection, and to remove all currant bushes from their
neighbourhood. A currant-free belt, 300—500 ft. wide, is
considered sufficient for security.

The teleutospores are produced on leaves of various kinds

56 CRONARTIUM RIBICOLA

of Ribes (Fig. 34); they have been recorded on 26 out of about
50 known species. The spermogones and ecidia are formed on
stems and branches of the five-leaved species of Pinus: they
have been found on five out of the eighteen Pines of that group,
but do not attack species having 2 or 3 leaves in a fascicle.
The following account is founded on that of Spaulding (1911).
The basidiospores are formed about the beginning of August,
and if they are blown by the wind, and adhere to moist young
branches of the Pine, the germ-tube enters and produces a
mycelium which. lives in the branch for several years, ultimately
causing it to become considerably swollen in a fusiform or
irregular manner. In about half the cases it is the main trunk

Fig, 34. Cronartium ribicola. a, a spore of Peridermium Strobi, x 600;
6, the teleutospore-columns on leaf of Red Currant (reduced); c, a uredo-
spore; d, top of a column of teleutospores, x 600.

that is infested. On this swollen portion spermogones appear
at almost any period of the year, followed in spring by the
ecidia, which break through fissures in the bark; these may
be even as much as 1 cm. high, yellowish-white in colour, with
orange spores. When the peridium bursts open, in an irregular
manner, the spores may be carried by the wind to any plants
of Rabes that may be near and can at once infect them. The
distance to which the zcidiospores can be effectively carried is
estimated to be less than 500 ft. These spores cannot infect
the Pine; but if they fall upon a moist leaf of Ribes, the uredo-
pustules usually appear, on the underside, in from 10 to 20 days.

These uredospores can, as usual, reproduce themselves on

MELAMPSORA PINITORQUA 57

any leaves of Ribes to which they may be carried, thereby
forming a means of rapidly spreading an infection which is
once started. After a few weeks, brownish thread-like growths
appear in the uredo-sori: these are the filaments on which the
teleutospores are borne, and the latter may be found from July
until the fall of the leaf, and even upon the fallen leaves. The
teleutospores may germinate at once, perhaps also after a con-
siderable time, but the basidiospores which they produce can
only infect young branches of Pine as described above. They
are distributed by the wind, but probably cannot be carried to
any great distance. Owing to the long incubation period of
the mycelium which will produce the ecidiospores, the fungus
cannot be seen on the Pine until it is at least three years old,
although infection may have taken place in the seedling: by
that time the leaves have naturally fallen off the part which
received the infection.

Though this disease does little harm to the currant, it is
necessary to destroy the infested bushes, since they form a focus
of infection for the Pines. Weymouth Pines that are more
than 20—25 years old are rarely liable to attack.

Melampsora pinitorqua.

THE PINE AND AspEN Rust.

The ecidial stage of this Melampsora lives on young shoots
of Scots Pine (Pinus silvestris) and its uredo- and teleuto-stages
on leaves of Aspen (Populus tremula).

The teleutospores germinate after a winter's rest, and the
basidiospores infect the young pine-buds, just beginning to
elongate in May and June. The mycelium produced penetrates
the cortex, and reaches also the bast and medullary rays. On
these shoots the spermogones appear about the end of May,
and are followed by the ecidia. The cortex of the affected part
becomes orange and dead, while the unaffected part still con-
tinues to grow. Thus thin shoots may be killed altogether,
but in the thicker ones curvature takes place owing to the
one-sided growth: the negative geotropism of the growing

58 MELAMPSORA PINITORQUA

point, combined with this lateral curvature, causes S-shaped
distortions which have given rise to the name “ Pine Branch
Twist” for the disease (Fig. 35).

It is suggested by Massee that the zecidiospores can repro-
duce the excidia and thus propagate the parasite on the Pine
without reference to the alternate host, but no proof is given of
this statement. The mycelium is almost certainly perennial
in the affected branch, and thus fresh outbreaks arise year by
year more or less, according to the weather in the spring.

If the ecidiospores are blown on to a leaf of Aspen, they
germinate there and the mycelium produces uredospores during

Fig. 35. Melampsora pinitorqua (from a German specimen, ex herb.
Sydow). «, a young shoot of Pine, in June, with newly-formed
leaves, showing three ceomata (@), shaded; the leaves have been
removed from the affected portion, which is beginning to be curved;
b, w ceoma, x10; cv, ecidiospores; d, old leaf of Aspen, showing
numerous teleuto-sori on the lower surface ; e, teleutospores.

the summer: these are usually so abundant as to cover the
underside of the leaves, and the parts on which they occur show
plainly as yellow spots on the upper side. For a time, the
uredospores spread the disease rapidly during the growing
season, until sometimes all the leaves of a tree are more or less
attacked and are covered with a bright orange powder. As
the leaves begin to die, teleuto-sori are formed; at first these
are brown, and show as little angular crusts on the under side.
Afterwards, when the leaves are fallen off, the crusts begin to
turn almost black as the teleutospores mature. These spores
are produced beneath the epidermis, standing erect and side by

CALYPTOSPORA GOEPPERTIANA 59

side in crowded patches; they are brown and unicellular. They
are not perfectly ripe till the following April; then, if the leaves
are gathered from the forest-floor where they have lain all the
winter, brought into a room and kept moist, the teleutospores
will germinate in great numbers by sending out basidia in the
usual way, though it is not possible to discern a germ-pore in
the ungerminated spore. The basidiospores in turn infect the
Pine, and the cycle begins again.

In the Aspen the mycelium affects only the leaves and
causes little harm, since the leaves do not fall off prematurely :
in the allied species, Melampsora Rostrupwi, I have found the
root-suckers round the parent tree to be most infested at first.
Besides P. tremula, the Abele Poplar (P. alba) is also able to
propagate the disease, as well as the hybrid between them,
P. alba x tremula. The chief harm is done to the Pines:
Hartig showed that seedling pines are often killed by an attack,
but if the tree manages to survive over thirteen years it may
recover. In any case the tree is more or less spoilt by the
distorted and dead branches, and, if it is true that the zcidio-
spores can spread the disease on the pines, it is evident that
young seedlings, when seen to be infested, should be pulled up
and burnt at once. Another precaution suggested by the life-
history is not to allow plantations of the two species of poplar
near to a seed-bed of Scots Pine.

It must be remembered that there are several other
Uredinales on Pinus silvestris, and also others on the Aspen
which have no connection with Melampsora pinitorqua: the
latter species can be easily recognised by the curvature of the
young pine-branches, which is not produced by any of the
others. Fortunately the disease is rare in this country, as in
many other countries.

Calyptospora Goeppertiana.

THE COWBERRY AND SILVER Fir Rust.

This fungus produces its teleutospores on the Cowberry
(Vaccinium Vitis-idaea), and its ecidiospores chiefly on the
Silver Fir (Abies pectinata): it has no uredospores.

60 CALYPTOSPORA GOEPPERTIANA

It can live at any rate for a number of years in the Cowberry,
in which the mycelium is perennial, but in the Fir the mycelium
is short-lived and perishes when the leaves prematurely fall off.
In Europe only the Cowberry has been noticed as its teleuto-
spore-host, but in the United States it is recorded on eight é
other species of Vaccinium (including V. Myrtillus A. Gray);
strange to say, the fungus has not yet been observed on the Fir
in America. Besides Abies pectinata, it is recorded in this
country on A. Nordmanniana (from Wales, etc.), and the zcidia
have occurred or been produced
artificially elsewhere on at least 10
other species of the genus. A good
account is given in the Kew Bulletin
(1907) from which and other sources
the following is drawn.

The most noticeable effect is
produced upon the Cowberry. The
ecidiospores ripen in July and
August, and if one of them is
carried to a young branch of the
Cowberry, its germ-tube penetrates
through a stoma (or, it is said, bores
its way through the outer epidermis
wall), and penetrates into the cortex,
where it grows and next spring ex-
tends itself into the new shoots.
These present a remarkable appear-
ance : the internodes are lengthened,

they become spongy and _ strongly
a Ps Calyprospora Goepper- swollen and coloured red or pink,
ee ere afterwards turning brown (Fig. 36).
Abies pectinata, b, leaf of A. Lhe infested plants are taller than

Nordmanniana, with wscidia, ‘
moraniy (le luther bedlucea. uninfested ones and have smaller

leaves. The mycelium perennates
in the affected shoots, and passes each spring into the newly
formed ones; thus the diseased branches usually occur in
clusters. Finally the mycelium penetrates into the epidermis,
and the teleutospores are formed within the epidermal cells

CALYPTOSPORA GOEPPERTIANA 61

which they completely fill, They are mostly divided by two
crossed walls into groups of four cells, each provided with a
germ-pore at its upper and inner corner.

In the spring following their formation, they germinate in
situ about May, sending out their basidia through the dead
epidermis, and producing their basidiospores in the air. These
are blown by the wind on to the just-starting shoots of Fir, and
infect the young leaves, on which they produce the ecidia on
yellow spots in two rows, one on each side of the midrib. The
ecidia are cylindrical, white, with torn margin, }—1 mm. high,
filled with orange spores, and when empty look like the remains
of insects’ eggs. Their spores are soon ripe and can infect the
young Cowberry shoots, but not the Fir. The diseased leaves
soon turn yellow and begin to fall off during July; it is this
early defoliation of the Fir that does the harm.

There is obviously no cure except to remove and burn the
infested Cowberry plants, and for the sake of prevention these
should be searched for in the neighbourhood, when a plantation
of Silver Fir or its allies is going to be made. There is no
difficulty in detecting them on account of the peculiar appeatr-
ance which they present. The disease is rare in this country,
and is confined mainly to the moorland districts.

These life-histories have been selected in such a way as to
show, so far as could be done from British species, the remarkable
variations that exist in the cycle of development and in the
occurrence of the different spore-forms of the Uredinales.

CHAPTER V
SPECIALISATION

The Uredinales are strictly parasitic (obligate parasites).
Many parasitic Fungi can live for a time saprophytically, but
those belonging to the present group are quite incapable of
such an existence. They are wholly dependent upon their
host. Moreover, a study of the evolution of the Uredinales
shows us that they have sprung from some simple beginning
(resembling perhaps remotely the Ustilaginales) in such a way
that new forms ever appeared as new hosts were evolved, and
advanced part passu with them. The lowest forms are those
parasitic upon the Ferns, the highest are among those on the
Compositee and others of the more specialised orders.

Each form is more or less closely adapted to its particular
host, but there is a wide range among them in this respect. A
species which can find sustenance upon hosts of more than one
kind is called plurivorous. One of the widest is Puccinia
Malvacearum, confined, indeed, to the Mallow family, but
appearing to spread to nearly every genus of the group Malvex
of that family. It has been experimentally shown that it can
be transferred from Malva to Althaea and vice versd, and obser-
vations on its occurrence in nature imply that it can pass
equally to other genera (see under that species). Or a species
may be found only on part of a subfamily, as P. Arenariae on
many genera of the Alsinew. Others are confined to a single
genus, but appear to be equally at home on almost any species
belonging to it, as P. Violae. Still others are restricted so far
as we know to a single species, as P. Buai, and various species

ARTIFICIAL CULTURES 63

of a genus may have totally distinct rusts upon them, as in the
genus Galiwm. So far there is nothing that would contravene
one’s expectations, but it is impossible to avoid a little in-
credulity when one is told that on Hieraciwm there is a
Puccinia which is confined strictly to a single form of a variety
of a subspecies of a species of that genus (Probst, 1909).

These specialisations can be proved only by artificial cultures.
Certain experimenters have developed very successful methods
of infecting given plants with the spores. The ecidio- or uredo-
spores are the handiest for this purpose. A sorus of mature
uredospores is removed from the leaf, placed in a drop of water
and broken up with a needle; the spores are then thoroughly
shaken up with a suitable quantity of distilled water. The soil,
in which the plant to be used is growing, should have been
well watered before the experiment begins. The leaves to be
inoculated are first sprayed with distilled water from an atomiser,
and then the liquid containing the spores is similarly sprayed
upon them, naturally upon a surface which possesses stomata.
The plant is then placed under a bell-glass for 24—48 hours or
longer, and afterwards kept in a greenhouse at a suitable tem-
perature, protected if required by a larger glass shade with
sufficient ventilation. A similar uninoculated plant should be
kept near it asa control. In spraying, too great a quantity of
moisture should be avoided; in nature it is observed that the
germination of spores succeeds best in a layer of dew, not of
rain. The keeping under a closed bell-glass is for the purpose
of allowing the germ-tubes to enter the stomata; after that
nothing more is required but to grow the plant in the ordinary
way. The result of the infection will begin to appear in about
10—20 days or more, according to the species.

The inoculation with teleutospores which are ready to
germinate may be effected in the same way; or a leaf with
mature teleutospores may be tied, spores downwards, on the
plant at the selected spot, and left to itself or protected for a
time with a layer of wet cotton wool. If it be required to
discriminate, under special circumstances, between the artificial
infection and any accidental one that might occur, the leaf to
be used may be marked with lines in waterproof ink, and the

64 CORN-RUST

spores carefully placed between the lines with a camel-hair
brush.

A few instances of specialisation will now be given, in
addition to the less complicated cases which are treated of
under the several species in the systematic part. It will be seen
that greater economic importance attaches to this specialisation
than might at first be imagined. The first example taken will
be that of Puccinia graminis, which is found upon various
grasses, especially upon cultivated cereals. In the early days of
this study almost any rust upon corn was called P. graminis;
afterwards it was found that there are several kinds, which can
be easily separated by their form or colour, and the real
P. graminis is distinguished as Black Rust, on account of the
conspicuous black striae which its teleuto-sori form upon the
culms in autumn. Its uredospores also can easily be dis-
tinguished from the uredospores of the other species which live
upon the corn. But even after restricting the application of
the name by these morphological distinctions, the species is
still recorded on more than 180 kinds of grass, although of
course some few of these records may be erroneous.

When discussion took place in the past upon the mode by
which epidemics of Corn-rust were caused, apart from the
Barberry, year after year, it was considered sufficient to point
to this wide prevalence of the species, and to assert that it lived
through the winter upon the wild grasses and passed from them
to the corn when the time arrived. Eriksson is the experimenter
who has done most to refute this idea; by making artificial
inoculations he has proved that in certain cases the rust which
is found on wild grasses will not infect the wheat and vice versd.
In spite of this biological difference, however, in most cases no
morphological distinctions can be detected, or, if so, they are
very slight and somewhat variable. Nevertheless the difference
exists, though in varying degrees of definiteness; exactly the
same kind of specialisation has been proved to exist in the
Erysiphacee. The natural explanation is that the species,
P. graminis, was originally parasitic on numerous grasses, quite
indifferently; but as time went on, certain reasons, perhaps
geographical or ecological, caused some sets of individuals to

BLACK RUST 65

restrict themselves to a particular species of grass; in course of
years they became more and more closely adapted to this host,
and in so doing grew less and less able to infect other species.
But possibly they have seldom completely lost this power,
as is shown by the existence of “bridging” species of which
Ward produced the best evidence in P. bromina; these will be
referred to later.

It -is assumed that all the forms of P. graminis will infect
the Barberry; the restrictions are confined to the alternate
host. As a result of his experiments, Eriksson divided the
species into the following “special forms,” which are here called
“biological” races :—

f. Secalis—on Rye
f. Avenae—on Oat
f. Tritici—on Wheat
f. Atrae—on Aira

f. Agrostidis—on Agrostis
6. f. Poae—on Poa.

Sot re Oates

Race 1 grows not only on Rye, but also on Hordeum vulgare,
H. murinum, Agropyron repens, A. caninum, Elymus arenarius,
Bromus secalinus etc. (In all these enumerations non-British
species will be omitted.)

Race 2, on Oat, and on Arrhenatherum elatius, Dactylis
glomerata, Alopecurus pratensis, Miliwm effusum, Bromus
arvensis, B, madritensis, Festuca Myurus, F. sciuroides, F. ovina
(tenurfolia).

Race 3, on Wheat, but also though more rarely on Barley,
Oat, and Rye.

Race 4, on Aira caespitosa.

Race 5, on Agrostis canina, A. stolonifera, A. vulgaris.

Race 6, on Poa compressa, P. caesia, P. pratensis.

A seventh Race, f. Horde, is sometimes added, though
Eriksson included it under his f. Tritici or f. Secalis,

Jaczewski (1910) from numerous inoculation-experiments
arrived at somewhat different results: he found that he
could infect Rye only from Agropyron repens, A. caninwin,
Bromus secalinus, and Dactylis glomerata; Wheat only from A.
repens, Festuca gigantea, and Lolium perenne; Oat only from

GU. 5

66 BLACK RUST

Arrhenatherum elatius, Alopecurus pratensis, Avena pubescens,
and Festuca ovina; and Barley only from Zriticwm and Lolium
perenne. According to him it seems that only Barley and
Wheat could infect each other directly, although it is known
from other sources that Wheat can also infect Rye; this
could, however, be done even according to Jaczewski’s state-
ments, if Agropyron repens were employed as a “ bridging”
species.
Carleton, in North America, experimented with much the
same forms, but reached a still different result. According to
him (1899) there are only two biological races :
1. f. Lritici—on Wheat, Barley, Hordeum murinum, Koeleria cristata,
Festuca gigantea, Dactylis glomerata, Agrostis alba.

2. f. Avenae—on Oat, Avena pratensis, A. fatua, Hordeum murinum,
Dactylis ylomerata, Koeleria cristata, Arrhenatherum elatius, Holcus
mollis, Ammophila arenaria, Alopecurus pratensis.

In a further publication (1904) he adds to form (1) that
Holcus lanatus should probably be included, and furthermore
that ‘there is a form of P. graminis on Agrostis alba vulgaris
which could not be transferred to Wheat or Oat.

Freeman and Johnson (1911) in the U.S. cultivated P.
gramims by its uredospores alone for two years without any
loss of vigour. They found that the uredospores of f. Tritica
would infect Barley easily, rarely Rye, and never Oat, but by
using Barley as a “bridging” species, they could infect, with
the uredospores produced on that, Rye easily, and Oat in a less
degree. The uredospores of f. Hordei would infect Wheat and
Barley easily, and in a less degree Oat and Rye. The uredo-
spores of f. Secalis would infect Barley, and by using that as a
“bridging” species would infect Oat at the second step, but in
a less degree. The most specialised form was f. Avenae; besides
the Oat its uredospores would infect only Barley, and not always
that.

From this it is evident that either (1) the specialisation of
these races is less sharp than Eriksson would have us believe,
or (2) the specialisation is taking place along two distinct lines
in the United States and in Europe respectively. Probably
both of these statements are true, but in support of the former

YELLOW RUST 67

view we may adduce the fact that Eriksson could infect Berberis
vulgaris with teleutospores obtained from many grasses (Wheat,
Oat, Barley, Rye, Arrhenatherum elatius, Agropyron repens,
A. caninum, Dactylis glomerata, Agrostis stolonifera, Elymus
arenarius, Poa compressa, P. pratensis, Aira caespitosa, Bromus
secalinus, and many others, non-British) while Bolley was able
to infect a large number of the grasses with spores taken from
a single Barberry hedge.

The economic importance of the matter lies in the fact that,
if the specialisation is as strict as Eriksson maintains, the corn-
crops cannot often (in the absence of Berberis) be infected by
rust on other cereals or on the wild grasses in the neighbouring
hedges. This excessive strictness, however, no one else is
prepared to admit: in any case there are obviously plenty of
“bridging” species which would enable the rust to get at the
corn at the second step, if not at the first. The reason why we
cannot, so far, attribute any very great accuracy to the state-
ments regarding specialisation is that the conditions required
for infection are demonstrably very complex and at present ill-
understood, so that a negative result, even when repeatedly
occurring, often proves nothing whatever. This is manifest
from Eriksson’s own complaints about the “ capriciousness ” of
the germination of the spores, and from the frequent recurrence
of such remarks as this—“ Uredospores from Aira caespitosa
would not always infect Aira caespitosa.” There is another
very important conclusion that can be drawn from this survey,
viz. that the life-histories of hetercecious rusts must always be
worked out separately for each country in the world.

A second example of specialisation is given by the Yellow
or Golden Rust (P. glumarum): it is divided by Eriksson into
five biological races :

f. Triticc—on Wheat alone

f. Hordei—on Barley alone

f. Secalis—on Rye (perhaps also on Wheat)

f. Llymi—on Elymus arenarius alone

f. Agropyri—on Agropyron repens,

but of the last Eriksson remarks that he could not infect this

host with its own uredospores. With the exception mentioned,
5—2

SP wb

68 CROWN RUST

each of these forms would not infect the other hosts, so far as
they were tried. It is doubtful if the two last forms are dis-
tinct, as they were not tested sufficiently.

A more complicated case is seen in P. coronata Corda, the
Crown Rust. Not only is this divided by Eriksson and Klebahn
into two sub-species, P. coronata and P. Lolii (=coronsfera),
but each of these is still further sub-divided by them into
biological ‘races.

In P. coronata (ecidium on Rhamnus Frangula) they are:

f. Calamagrostidis—on C. lanceolata etc. (also on Phalaris)
f. Phalaridis—on P. arundinacea (also on Calamagrostis)
f, Agrostidis—on A. vulgaris, A. alba, A. stoloniferc

f. Holct—on H. lanatus, H. mollis

f. Agropyri—on A. repens.

She oe Poo

The state of P. coronata which occurs on Dactylis has not
yet been assigned to any of these forms. The last two of them
seem doubtful and may not belong to P. coronata, and the first
two of them may be identical, since each seems able to extend
at times to the other host.

In P. Loliz, the Crown Rust of Oat (ecidium on RA. cathar-
ticus), they are:

. Avenae—on A. sativa, A. fatua

. Lolii—on L. perenne (perhaps also on Festuca elatior)
. Festucae—on F. elatior, F. gigantea

. Holci—on HA. lunatus, H. mollis

. Alopecuri—on A. pratensis etc.

. Glyceriae—on G. aguatica.

To which of these the form of P. Lolii on Arrhena-
therum should be assigned seems to be undecided. There are
also Crown Rusts, of which little is known, on Melica nutans
and Sesleria coerulea.

Later (Arkw for Botanik, 1908, vol. viii) Eriksson revises
his previous conclusions in regard to P. Lolti as follows, making
eight races :

Oo Pp wee
re rh rh bh ph Ph

1. ff Avenae—on Avena

2. £. Alopecuri—on Al. pratensis, etc. (sometimes on Av. sativa)
3. f. Festucae—on F. elatior, F. gigantea
4.

f Lolii—on Lolium perenne and other species (also on F. elatior)

UROMYCES ON GRASSES 69

or

f. Glyceriae—on G. aquatica

6. f. Agropyri—on A. repens

f. Epigaei—on Calamagrostis epigeios (also, but rarely, on Avena
sativa)

8. f. Holei—on Holcus lanatus.

a

To these Miihlenthaler (1910) adds a ninth form, on several
species of Bromus. These results agree pretty well with those
of Klebahn, but not with those of Carleton. According to the
latter, the only host of P. Loli in nature, in the United States,
is Avena sativa: but in artificial cultures it can be foisted on
other species because of the unnatural conditions, especially
on account of the employment of very young and non-resistant
plants. In any case, however, there is a general agreement
that the form of P. Lolit on Avena sativa cannot be transferred
to Wheat, Barley, or Rye. In fact, with the exceptions
mentioned, it was found by all experimenters more or less in
all these cases, that attempts made to transfer the fungus from
the host of one “special form” to those of the others were
unsuccessful from some unknown cause.

The state of things in regard to the two common Uromyces
species, found upon Grasses, is more perplexing. There is no
agreement whatsoever between the various authors who have
experimented upon them. The latest results are, perhaps, the
following :

Krieg (1909) divides Uromyces Dactylidis into two biological
races or “ formee speciales ” :

l. f£ sp. with ecidium on Ranunculus bulbosus, R. repens
2. f. sp. with ecidium on several non-British species of Ranunculus.

Juel (Svensk Bot. Tidskr. ii. 169, 1908) divides Uromyces
Poae into nine biological races, of which the following seven
may be British:

. sp. Ficariae-nemoralis

. sp. Ficariae-trivialis

. sp. Ficariae-pratensis

. Sp. repentis-nemoralis (also on R. bulbosus)
. Sp. repentis-trivialis (also on Poa annua)

. Sp. repentis-pratensis

. Sp. auricomi-pratensis.

BT Se Ore ge NS
bor oP rh rh ph fh

70 BROWN RUST

The hosts of these are indicated by their names, but there
is a very high probability that their distinction depends entirely
on accidents of weather or manipulation at the time when the
inoculation was made.

Finally, in regard to another case, P. dispersa (sens. lat.)
the Brown Rust of Corn, it will be seen by referring to the
systematic part that it is divided into a number of subordinate
forms or sub-species, which are for the most part only dis-
tinguishable biologically ; though here the amount of difference
is much greater than in P. graminis, and there is more to be
said in favour of calling these forms distinct species, as is often
done. For, as will be seen by the descriptions, in some of them
an ecidium stage is known, in others not, though Klebahn
remarks that in the latter cases we might possibly find the
zecidium, if we could trace each form to its ancestral home.
Moreover, in some of these cases, the teleutospores germinate
in the spring, in others in the autumn.

One of the most remarkable of these forms is P. bromina,
on species of Bromus, from which Ward (1902) obtained such
important results. For instance, he showed that uredospores
taken from B. mollis always infected B. mollis and B. secalinus
and their close allies, but not B. sterilis and its allies; while
on the other hand those on B. steril¢s would infect B. sterilis
and its ally B. madritensis, but rarely the other Bromes. We
can reason, as Ward says, that uredospores from B. mollis infect
that species readily “because their food-supplies and previous
environment have affected their protoplasm in some way which
makes it easier for their germ-tubes and mycelium to grow in
tissues which afford them the same nutriment and present the
same obstacles, as they have hitherto enjoyed or been con-
fronted with” (p. 299). They can flourish in B. secalinus
because here also the food-supplies etc. offered are nearly the
same. But in B. sterilis the resistance of the plant to infection
is sufficiently great to present a barrier which is incapable of
being overcome except by an odd spore, here and there, varying
from the normal. In 4, out of 148, trials, uredospores from B.
mollis infected B, sterilis and these might then produce spores
which could pass on to B. madritensis, although in no single

BRIDGING SPECIES 71

case out of many experiments could B. madritensis be infected
directly from B. mollis. B. sterilis therefore acted as a
“bridging” species, and enabled the parasite to pass from
B. mollis to B. madritensis, though it could not do so without
this intervention. The same existence of “bridging” species
has been demonstrated in Erysiphe graminis, and no doubt
will be found in numerous other instances.

Ward further mentions (1903) that he found B. ardwennensis
var. villosus to be infectible by the spores from B. sterilis, B.
mollis, and B. patulus, as well as by those from B. arduennensis,
and therefore easily able to serve as a bridging species between
these others. Nevertheless, that such intermediary species
do not exist in all cases is proved by the fact that when he
grew “more than 200 species and varieties of Bromus side by
side or intermingled in contiguous beds, certain species invari-
ably caught the disease and became rusted, while others close
by showed no sign of infection.”

The occurrence of these abnormal spores, i.e. mutations,
which is proved in the case of Bromus, is of great significance
and gives us the clue by which we can understand how a
gradual or sudden passage can take place, and has taken place,
from one host to another, so that now an appreciable percentage
of the modern vegetable world have parasites more or less
specialised to themselves.

Another important consequence follows from this fact of
specialisation. If the parasite is so narrowly adapted to its
particular host, it may be expected that varieties of the host
can be found or bred which will be able to resist attack, that
is, will be immune. A great deal of research has been devoted
of late years, especially by Professor Biffen of the Agricultural
School at Cambridge, to this subject of breeding a race of
wheat which will be immune to Rust, and a certain amount of
progress has been made. Immunity depends chiefly (perhaps
entirely) upon the ability of the cytoplasm of the host-cells
to resist infection by secreting anti-toxins which will kill the
mycelium of the fungus. Immunity and susceptibility (which,
however, seem to be always relative only and not absolute) have
been proved to be inherited, and in fact to be Mendelian

72 IMMUNITY

characters, the latter being dominant (Biffen, 1907, 1912; Pole-
Evans, 1911). But owing to the minute specialisation which is
characteristic of many Rusts, a variety may be immune to one
Rust while susceptible to another, or may even be immune in
one country but susceptible to the same Rust in a different
climate. The latter change would depend upon a slight
disturbance (by climatic factors) of the delicate balance which
existed between the attacking and resisting powers of the two
organisms.

It may be pointed out here that this affords an opportunity
for dealing a final blow at the moribund “ mycoplasm” hypo-
thesis. For when a susceptible and an immune variety of
wheat were crossed (Biffen, 1905) both the reciprocal crosses
were susceptible. Yet in that cross in which the pollen used
was taken from the susceptible parent, while the other was
immune, the only means by which the “mycoplasm” could be
conveyed would be in the generative nuclei of the pollen-tube,
which is inconceivable. An attempted criticism of this con-
clusion (Butler, 1905) misses altogether the point of the
argument: the maternal parent could not be classed as
immune, if it usually contained the “ mycoplasm ” already in
its tissues.

CHAPTER VI

CLASSIFICATION AND PHYLOGENY

The Uredinales form a group of Fungi so closely allied that
they must be regarded as monophyletic. The number of species
considered in the present work is about 250. They may be
divided into five families.

A. IJMPEDICELLATA.

I. Melampsoracese—Teleutospores not pedicellate, but
seated on a dilated hyphal cell, produced singly in the
tissues of the host, or compacted side by side into flat crusts,
1—4-celled. Germination by an external basidium, with
minute round basidiospores (about 10). Uredospores ab-
stricted singly. Aicidia with or without a peridium.

II. Cronartiaceee—Teleutospores not pedicellate, produced
in chains, which are either separate or united into columnar,
wart-like, or lens-shaped bodies. Germination asin I. Basidio-
spores round, small (about 10 yu or less),

III. Coleosporiacee—Teleutospores in one (or rarely two)
layers, forming waxy, bright-coloured crusts, not pedicellate,
but seated on a dilated hyphal cell, at first one-celled. Ger-
mination by the formation of an internal basidium of four
superimposed cells, each of which protrudes a sterigma and a
large basidiospore (about 20 «).

IV. Endophyllaceze—AKcidio-teleutospores surrounded by
a hemispherical peridium and produced from a fusion-cell, in
chains with intercalary cells, but germinating with an external
basidium.

74 CLASSIFICATION

B. PEDICELLATZ.

V. Pucciniaceee — Teleutospores distinctly pedicellate
though the pedicel is often very short or caducous, united
into pulvinate sori, easily separable or immersed in gelatine,
each spore consisting of one or more cells arranged in rows or
groups. Germination by an external basidium ; basidiospores
more or less ovate. A¥cidia with or without peridium. Uredo-
spores always abstricted singly on distinct pedicels.

The following genera belonging to these families are British,
excluding Uredinopsis :

MELAMPSORACEA.

A. Teleutospores composed of 2—4 laterally adherent cells, the septa
in the latter case cruciately arranged.
a. Teleutospores colourless, scattered singly in the host-tissues or
formed in the epidermal cells. On Ferns.
[1. Teleutospores extracellular, scattered. Uredospores surrounded
by a peridium, colourless, without germ-pores. Uredtnopsis.]
2. Teleutospores intracellular. Uredospores surrounded by a
peridium, colourless, without evident germ-pores. Jfilesina.
3. Teleutospores intracellular. Uredospores without (or with a
very rudimentary) peridium, yellow, with germ-pores.
Hyalopsora.
b. Teleutospores brownish, in the epidermal cells or forming
subepidermal crusts. On Seed-plants.
1. Teleutospores intracellular. cidiospores without smooth

spot. Calyptospora.

2. Teleutospores intracellular. AEcidiospores smovcth on one side,
Thecopsora,

3. Teleutospores extracellular, subepidermal. Kcidiospores
smooth on one side. Pucciniastrum.

B. Teleutospores one-celled, united into crusts.

a. Teleutospores in the epidermal cells, usually one-celled, faintly
coloured. Uredospores with peridium, but without para-
physes, and without evident germ-pores. Acidium with a
peridium. Melampsorella,

b, Teleutospores not intracellular, one-celled, brown.

1. Teleutospores subepidermal. Uredo- and secidiospores as
above. Melampsoridium.,
2. Teleutospores subepidermal or subcuticular. - Uredospores
without peridium, but with paraphyses. cidium of the

cxoma type. Melampsora.

PHYLOGENY 75

CRONARTIACEA.
A. Teleutospores in pulvinate sori. Chrysomyxa.
B. Teleutospores in columnar sori. Cronartium.
COLEOSPORIACE.

A. Basidiospores fusiform. Uredospores formed singly. Ochropsora.
B. Basidiospores ellipsoid or lemon-shaped. Uredospores in chains.

cidia with an inflated peridium. Coleosporium.

C. Basidiospores globose. Basidium ultimately protruded. Teleuto-

spores with lateral pedicel. Zaghouania.
ENDOPHYLLACEZA.

Kcidio-teleutospores in a cup-shaped peridium, germinating with an
external basidium. Endophylium.
PUCCINIACEA.

A. Teleutospores embedded in gelatine, on Gymnosperms. No uredo-
spores, Gymnosporangium.

B. Teleutospores free, on Angiosperms.
a. Teleutospores of more than two cells.
1. Teleutospores of many cells. No uredospores. Xenodochus.
2. Teleutospores of several cells, dark-coloured. Czeomata en-
circled by paraphyses; uredospores the same and with
paraphyses intermixed. Phragmidium.
3. Teleutospores of several cells, faint-coloured. Czeomata with-
out paraphyses ; uredospores often with them. Kuehneola.

4, Teleutospores of three radiating cells. Triphragmium.
b. Teleutospores of one or two cells.
1. Teleutospores of two cells, usually. Puccinia.
2. Teleutospores of one cell. Uromyces.
PHYLOGENY.

In trying to comprehend the phylogenetic evolution of this
group of Fungi, there are several landmarks which can be
borne in mind. In the first place, their strict parasitism
implies a very close adaptation between them and their hosts:
this is not only a priori probable, but is confirmed by culture-
experiments and the existence of biological races. Therefore
those which are parasitic on the lowest hosts must be, on the
whole, most similar to the primitive forms, and those parasitic

76 PHYLOGENY

on the higher families would be expected to show the greatest
advance. This consideration alone is sufficient to determine
that Uredinopsis is like one of the primitive Uredinales and
that the genera Puccinia and Uromyces contain the highest
types. For Uredinopsis grows upon Ferns, and more than a
quarter of the Pucciniae live on the Composite.

Secondly a comparison of the spores of these two genera
and their respective allies suggests that the possession of a
single definite and well-formed germ-pore is a characteristic of
the latest forms, while the primitive ones had no germ-pores
at all, but protruded the germ-tube, as a conidium usually
does, at any convenient point or where the wall first gives way.
There is reason, from another point of view, to conclude that
germ-pores, when first existing, were numerous and indefinitely
scattered. A gradual reduction in their number and their
restriction to definite parts of the spore-wall occurred during
the course of evolution. The ecidio-teleutospore of Hndo-
phylam has no germ-pore; in the Pucciniacee the ecidio-
spores have usually several indistinct ones, the uredospores
have them fewer and more easily visible, and the teleutospores
have one or a small number, oftentimes very plainly marked.

Amongst the other Fungi, the group which presents the
nearest approach to the Uredinales is that of the Ustilaginales,
which are also parasites; their teleutospores (brandspores), in
the family Ustilaginacez, germinate in a similar way, but with
less definiteness, by the formation of a basidium and basidio-
spores. It may be inferred that this particular feature is one
of the most deeply seated characters of both groups, and is
therefore inherited from their ancestors.

Moreover, this feature is exhibited in the Uredinales by
cells which belong to the sporophytic generation, and after a
certain amount of growth the mycelium produced by the
basidiospores bears the two kinds of gametes. An exactly
similar course of events takes place in certain Algae, e.g.
Grifithsia, where the sporophyte bears tetraspores which on
germination produce a thallus which bears gametes. It is true
that in the Red Alge the tetraspores are more usually arranged
in tetrahedral fashion, but other modes also obtain, among them

yaa

ar

PHYLOGENY 77

(in Corallina) an arrangement of four superposed cells as in the
“basidium ” of the Uredinales. Moreover in one of the groups
of the latter, the Coleosporiaces, as well as in Chrysopsora, this
division into four cells takes place within the spore-mother-cell,
not outside it. It is a reasonable tentative hypothesis that
this “internal” formation of the four cells is the primitive
mode, inherited from the predecessors of the groups, and that
the formation of an external “basidium” is a later adaptation
of their successors to their environment. This would lead one
to look for the ancestors of the Uredinales among the Red
Algee.

The obvious implication is that the Coleosporiacee retain
much of the Uredinal primitive character, and this is borne out
by the fact that the ecidial host of Coleosporium is in every
case, so far as known, a species of Pinus. The teleutospore-
host may belong to the Composite or varivus other families, it
is true, but it is now generally admitted that the ecidial host
is the primitive, and that the others have been adopted by
successive mutations. It has an important bearing on this
argument that, in Gallowaya Pini, the teleutospores which are
of exactly the same nature as in Coleosporiwm are borne on
a Pine (Pinus inops Ait.).

Again, it has been shown that‘it is possible, without violence,
to interpret the female-cells of the ecidium as furnished with
a trichogyne, such as the carpogonia of the Floridez possess,
though in the Uredinales (possibly as a consequence of their
terrestrial habit) it has become abortive. Trichogynes are not,
uncommon in other groups of Fungi—in certain Ascomycetes,
in the Laboulbeniacez, and among the Lichens, in Collema and
other genera. Moreover in Collema the trichogyne and corre-
lative spermatia are almost certainly functional (Bachmann,
1912); the same is true in the Laboulbeniacer, but in most
Ascomycetes the trichogyne has either been lost altogether, or
if it survives has lost its function.

In order that the trichogynes in the ancestral Uredinales
should be effective, the female gametes must have been situated
beneath a stoma. It is a suggestive fact that in certain of the
group, belonging to the lowest forms, the sori of various kinds

78 PHYLOGENY

are always, or usually, so placed. For instance, in Melam-
psoridium betulinum the teleuto-sori almost invariably originate
directly below the stomatal pore. The cause of this cannot be
merely the need of oxygen for respiration, since it has been
shown that the intercellular spaces of a leaf are all well supplied
in that respect. Fig. 37 is drawn from the lower epidermis of
a leaf of Betula alba in which teleuto-sori were just beginning
to be produced. The same thing is true of the teleuto-sori of
Melampsora Larici-epitea and other Melampsorae,and apparently
even of Phragmidium. In others of the lower groups, Uredin-
opsis, Milesina, etc., uredo-sori are equally so placed, both

Fig. 37. Melampsoridium betulinum. a, young sori of teleutospores,
viewed through the epidermis, showing how they originate beneath
a stoma ; at the lower right-hand is a sorus with only two teleuto-
spores, x 300; b, three young teleutospores, forming a similar
sorus, seated on a common base, x 600.
primary uredo-sori (which represent zecidia) and secondary. It
can be justifiably inferred that this was the primitive position
in which the female gametes and afterwards the other kinds of
spore-sori were formed at the beginning: in Puccinia and its
allies this position is no longer maintained.

Since in the Ustilaginales (a comparatively non-progressive,
if not degraded, group) there is only one kind of spore besides
the basidiospore and the ensuing conidia, and this is produced
irregularly and not in definite sori, it may be inferred that the
same was true of the primitive Uredinales. This one kind of
spore must have been the equivalent of the teleutospore. It is

PHYLOGENY 79

found that, in Uredinopsis, the teleutospores are irregularly
scattered throughout the spongy mesophyll of the host. But,
in this position, their germination, or at least the liberation of
their basidiospores, could not take place easily until the leaf
had decayed, The transference to a place either (1) beneath the
cuticle, (2) in the epidermal cells, or (3) just beneath the
epidermis, and their aggregation into definite sori which by
their upward pressure would burst through the overlying layers,
would both be an advance in adaptation; so these various
positions are found to be occupied in successive genera, and the
most effective of all (the subepidermal sorus) is alone to be met
with in the highest groups.

In regard to a peridium, this can be supposed non-existent
at first, (1) because there is no peridium in the Ustilaginales,
(2) because a peridium could not exist so long as the trichogyne
was functional.

Here, it is true, there is a little deviation from what might
be expected; a peridium is found in Uredinopsis, Milesina,
Melampsorella and Melampsoridium (round the uredo-sori), but
in Hyalopsora this is very rudimentary or completely absent,
and it can scarcely be compared, in any case, with the peridium
of the ecidial stage of Puccinia, being of a very different
character. It must be. considered as a special development,
separately originated for the protection of the uredo-sori in
these lower groups. When one considers the cidiospores, one
finds them in the intermediate types either without a peridium,
or with encircling paraphyses, or with an irregularly shaped
peridium, and it is only in the higher forms, such as Puccinia
and Uromyces, that the beautifully outlined “Cluster-cup” arises.

In respect to these higher groups it has been shown else-
where (Grove, 1913) that the Endophyllaces constitute the
starting-point from which the varied forms of the Pucciniacese
have been evolved. A certain amount of advance went on, of
course, simultaneously among the Impedicellate, though to
nothing like the same extent.

In Endophyllum the ecidiospore which is the product of the
fusion-cell is also the teleutospore which germinates with a
basidium: in accordance with theory it is accompanied by

80 PHYLOGENY

spermogones. The first stage of evolution was the separation
of this spore-form into two, one (the ecidiospore) germinating
conidially, the other (the teleutospore) following it and germi-
nating basidially: types approximating to this stage are seen
in the section Pucciniopsis. It is quite certain that uredospores
are only modified xcidiospores, formed as a mere multiplying
device without the intervention of another fusion-cell. The
peridium which is found in these later stages of evolution round
the ecidium was at first represented (doubtless even in the
primitive Endophyllum) by a mere circle of paraphyses or
not.at all.

From a cytological point of view, the fusion of the two
nuclei in the teleutospore may be taken as paralleled by the
similar fusion in the basidium of the Basidiomycetes; the
division into four basidiospores follows in both cases, although
the mechanism is different. If the view propounded in a
previous chapter is adopted, that the four cells of the “basidium”
of the Uredinales are the real tetraspores' and the basidiospores
are merely conidia whose function is to facilitate dispersion by
wind, it will be seen that the difference in the Basidiomycetes
consists in the fact that cell-walls are not formed round the
tetraspores previously to the production of conidia. This may
recall the fact that in the Red Algee the four spores in a tetra-
sporangium are also not surrounded by cell-walls before their
discharge into the water. Of course, in the subaérial Uredinales
and Basidiomycetes such naked masses of protoplasm would be
comparatively ineffective for propagation, and are here replaced
by methods more suitable to a land environment. The throw-
ing off of the basidiospores with a jerk appears to be the same
in both these groups.

A similar comparison with the Ascomycetes cannot be made
with equal advantage, until the students of that group of Fungi
have come to some semblance of agreement as to the actual
course of its cytological history. But it is impossible to over-
look the remarkable parallelism between the cytology of the

1 In the Himalayan Barclayella, which is placed among the Melampsor-
aces (?), these tetraspores are said to round themselves off and separate,
apparently as the normal mode, without forming basidiospores.

PHYLOGENY 81

Uredinales, as now known, and that attributed by Claussen
(1912) to Pyronema confluens. In his paper, which is a con-
clusive reiteration and confirmation of his earlier work, he shows
that the numerous male nuclei of the antheridium enter the
ascogonium, and in it pair with the numerous female nuclei,
but without fusing with them. These synkarya then pass out
into the ascogenous hyphe, and there multiply by numerous
conjugate divisions. Finally a pair of descendants of these
nuclei are seen in the young ascogenic-cell, one being male
and the other female: here they divide conjugately into two
pairs, one pair being the ascus-nuclei, and the other pair
reserve-nuclet which may repeat the process in several ways:
The two non-sister ascus-nuclei fuse; then the fusion-nucleus
divides, the first division being heterotypic (meiotic, reducing,
possessing synapsis and diakinesis stages) and the two following
ones, by which eight spores are formed, being homotypic.
There is thus in the life-cycle a single fusion, followed by a
single reduction. The ascus is a spore-mother-cell, comparable
to the teleutospore of the Uredinales, but forming an octad, not
a tetrad of spores. The two “reserve-nuclei,” left after the
formation of the ascus, answer to the two nuclei Jeft in the
“basal” cell of the zcidium. Compare in this respect especially
the process as it takes place in Hndophyllum. The sporophyte
generation consists then in Pyronema only of the ascogenous
hyphe, whose cells contain the diploid number of chromosomes
though arranged within two nuclear membranes.

In certain species of Laboulbenia (Faull, 1912) there is a
similar cytological history. The ascogenic hyphe contain two
nuclei which divide homotypically by conjugate division, and
two non-sister nuclei pass into each ascus where they fuse; the
two left in the ascogenic cell may repeat the process. The
fusion-nucleus of the ascus divides to form eight nuclei of which
four soon degenerate: the first division is meiotic and the others
homotypic. There is no double fusion in this group and the
same statement may justifiably be inferred to be true of other
Ascomycetes.

On the other hand, Harper (1900), Blackman, Welsford,
Fraser, Brooks, Carruthers (1911) and others, maintain that in

a U. 6

82 PHYLOGENY

Pyronema confluens and in many other members of the Asco-
mycetes there is a double fusion (one in the ascogonium and
one in the young ascus), followed by a double reduction in the
ascus during the formation of the eight spores, the first division
being meiotic, the second homotypic, and the third brachymevotze.
In view of the established relationship between the Ascomycetes
on the one hand and the Uredinales and Basidiomycetes on
the other, this idea seems to be very unlikely. If correct, the
double process is a special development, peculiar to some only
of the Ascomycetes. The matter can only be decided by fresh
investigations, but it seems in all probability that the hypo-
thesis of a second fusion and subsequent brachymeiosis is the
result merely of a misinterpretation of the observed. phenomena.

According to Lutman (1910), in the Ustilaginales most of
the cells of the mycelium are binucleate, but the perfect resting
spores are always uninucleate, as are the cells of the basidia.

Rawitscher (1912) says the same, and adds that the con-
jugate condition arises (according to the species) by the
anastomosis in pairs either of the basidium-cells, or of the
basidiospores, or of the cells of the mycelium produced by
them, and the passage of the nucleus of the one cell into the
other to form a synkaryon.

Finally, it may be pointed out that the ideas embodied in
the foregoing discussion are in harmony with the now generally
accepted doctrine of the polyphyletic origin of the Fungi, by
which it is assumed that their various groups are not derived
from one or two ancestors, but originated separately from
distinct sub-divisions of the Alge, much in the same way in
which (on a smaller scale) the non-chlorophyllose Phanerogams
have arisen from various orders or families of Flowering Plants.
From this point of view, according to which the vast majority
of the Fungi originated from the Red Alge, it is not without
significance that already some of that group are known which
(though still rightly classed as Algee) have assumed a true holo-
parasitic habit—a statement which cannot be made to the same
extent, if at all, about other algal groups. Examples are found
in the well-known Harveyella mirabilis (Sturch, 1899) and in
Choreocolaz Polysiphoniae (Richards, 1891).

PHYLOGENY 83

From these considerations the probable phylogeny of the
Uredinales may be represented as in the following schemes.

Ascomycetes Basidiomycetes

Uredinales

- Ustilaginales
Rhodophycese
Pucciniacese
Coleosporiacese
Endophyll
ndop ie Cronartiacex
(Endophyllum-like) Melampsoracese

Primitive Uredine |

84 MANIPULATION

Nore oN MANIPULATION.

Dried specimens of Uredinales keep most of their characters
unchanged for an unlimited time, but the colours fade except
those of the teleutospores. The only two difficulties found in
examining them are in regard to the markings on the outer
surface of the spores, and the number of germ-pores. For the
first, different methods succeed in different cases, but the finer
markings can usually be seen by examining the spores under a
one-sixth inch in air, or in water after squeezing out their
granular contents by tapping or pressing hard upon the cover-
glass. For the second, boiling for about a minute in a drop of
lactic acid, on a glass slide over a spirit-lamp, is the best course,
although expulsion of the contents under pressure frequently
brings the germ-pores into view; in fact so plain do they often
become that they can be photo-micrographed with ease. Boiling
in lactic acid also restores old collapsed spores to their former
size and plumpness. j

The preparation from which Fig. 37 was drawn was
obtained in the following way: lay the side of the leaf opposite
to the sori in a thin layer of 5°/, KOH solution for an hour or
so, then reverse and brush or scrape away the softened tissue
as far as possible ; on mounting the remaining surface, epidermis
upwards, in glycerine and water, the arrangement of the parts
can be clearly seen.

If it is wished to observe the germination of the spores in a
hanging drop, almost any uredo will serve; for teleutospores
Puccinia Malvacearum and for ecidiospores Zeidium Ficariae
are usually the most handy. One of the best double stains
to use is Diamant Fuchsin and Light Green; the former stains
the nuclei red and the latter the cell-walls green. Stain heavily
with the former and wash out with alcohol till the desired tint
is arrived at; then use the Light Green dissolved in clove oil.

UREDINALES

A group of Fungi which are obligate parasites on ferns
and the higher plants. Mycelium filamentous, branched, septate,
developed within the tissues of the host, producing teleuto-
spores (resting spores, chlamydospores) which on germination
give rise to a generally four-celled “basidium,” each cell of
which may in turn produce, on a sterigma, a single basidio-
spore (conidium). In addition, there are often produced
spermatia (in spermogones), ecidiospores (in ecidia), and uredo-
spores (in sori).

PUCCINIACE

Pucciyiz&. Teleutospores of one or two cells,
Uromyces.

scarcely gelatinous (except the pedicels in some
= ( : Puccinia.

foreign species).
PHRAGMIDIES. Teleutospores of more than two
‘cells, the walls of the pedicels subgelatinous.

Triphragmium.
Teleutospores more or less verrucose. Phragmidium
; Kuehneola.
Teleutospores nearly or quite smooth. Savona
GYMNOSPORANGIES. Walls of pedicels of teleuto- ‘
spores becoming highly gelatinous. Gymmosporangium.

UROMYCES Link.

Autcecious or hetercecious.

Spermogones deeply embedded in the tissues of the host,
flask-shaped with conical mouth and ostiolar filaments. Aicidia
with an evident, usually cup-shaped peridium; «cidiospores
with indistinct germ-pores. Uredospores formed singly on
their ‘pedicels, with several usually rather distinct germ-pores
which are often surrounded by a- thickened border, rarely
accompanied by paraphyses. Teleutospores one-celled, on dis-
tinct pedicels, almost always with an apical germ-pore. Basi-
diospores flattened on one side or kidney-shaped.

4

86 \UROMYCES

The species are i Se agree to the families to which
the hosts belong: see Puccini This genus is often considered
the most highly (at least the latest) evolved of the Uredinales ;
but rather it forms a heterogeneous group, the species of which
have arisen at different times from various species of Puccima.

1. Uromyces Valeriane Fckl.

Uredo Valerianae Schum. Pl. Sal. ii. 233.

Beidium Valerianearum Duby, Bot. Gall. ii. 908. Cooke, Handb.
p. 540; Micr. Fung. p. 196.

Lecythea Valerianae Berk. ; Cooke, Handb. p. 532; Micr. Fung. p. 222.

Uromyces Valerianae Fckl, Symb. Mye. p. 63. Plowr. Ured. p. 128.
Sace, Syll. vii. 536. Sydow, Monogr. ii. 19. Fischer, Ured.
‘Schweiz, p. 54, f. 41.

Spermogones. Epiphyllous, in small clusters, honey-
coloured, turning black.

4icidiospores. Kcidia hypophyllous, and often on the
nerves, petioles and stalks, seated on pale thickened spots,
densely aggregated or circinate, cup-shaped, whitish-yellow ;
margin revolute and torn; spores covered with minute crowded
warts, yellow, 18 —25 x 16—20 mu.

Uredospores. Sori amphigenous, usually on indefinite
yellow spots, scattered or aggregated here and there, minute,
punctiform, pulverulent, brown; spores globose to broadly
ellipsoid, verrucose-echinulate, yellowish-brown or brown, 21—
28 w; epispore 24—3 wu thick, with two or three germ-pores.

Teleutospores. Sori similar, but longer covered by the
epidermis, dark-brown; spores ellip-
soid or ovate, with a flat subhyaline
papilla at the summit, smooth, pale
clear-brown 20—30 x 18—21 y; epi-
spore thin, scarcely thickened above ;
pedicels short, thin, hyaline, rather

Fig. 38. U.Valerianae. Teleu- deciduous.
tospores and uredospore (the . . eee a
latter viewed dry) on V. offici- On Valeriana dioica, V. offict-

sae nalis, Adcidia in May and June;
uredospores from June, teleutospores from July to October.
Common. (Fig. 38.)

in) 87
i

4
The uredospores seem to be vari bid in their markings; some are
distinctly verrucose with pointed warts ;-éthers are as distinctly echinulate.

DISTRIBUTION : Europe and South Africa,

2. Uromyces Scrophularie Fckl.

Aecidium Scrophulariae DC.; Cooke, Handb. p. 544; Micr. Fung.
p. 199.

Uromyces Scrophulariae Fckl. Symb. Myc. p. 63. Plowr. Ured. p. 139.
Sacce, Syll. vii. 559. Sydow, Monogr. ii. 27. Fischer, Ured.
Schweiz, p. 75, f. 56.

U. concomitans B. et Br. ; Cooke, Micr. Fung. p. 213.

Spermogones. Few, singly or in little groups, simul-
taneously with the ecidia.

4;cidiospores. Acidia hypophyllous or on the stems, on
yellowish spots, in rounded clusters or in more or less elongated
patches on the nerves and stems, cup-shaped, yellowish ; margin
involute, entire; spores verruculose, smooth below, yellowish,
18—21 x 1418 p,

Teleutospores. Sori small and roundish, arranged like
the ecidia except that they form

more elongated groups (as much as.
10 cm. long) on the stems, long
covered by the lead-coloured epider-
mis,at length naked and pulverulent,
dark-brown; spores very irregular,

obovate, fusiform, or ellipsoid, angu-
lar, rarely sub-globose, apex rounded, Fig. 39. U. Serophulariae.

- - Teleutospores on. 8. aquatica,
truncate or slightly pointed, some-
what thickened (up to 6 w), with a dark-coloured cap, at-
tenuated below, smooth, brown, 18—385 x 11—18,; pedicels
persistent, hyaline or yellowish, nearly as long as the spore.

On leaves, petioles and stems of Scrophularia aquatica,
S. nodosa. July—September. Not common. (Fig. 39.)

The spots on the leaves are pallid, edged with violet-brown. The
teleutospores especially cause considerable distortion of the leaves and
stems. The two kinds of spores may be produced on the same mycelium,
and the ecidia and teleuto-sori can occur simultaneously and intermixed,
or the latter surrounding the former (Grevillea, iii. 181, pl. 36). For this

88 \eduyces

is one of the species in which it is stated (Dietel, Flora, 1895, Ixxxi.
396), that the zcidiospores can reproduce the xcidia. Spermogones are
found sparingly only with the first generation of the wcidia and at the
same time. The secondary ecidiospores, in fact, take the place of
uredospores.

DisTRIBUTION : Europe generally.

3. Uromyces Limonii Lév.

Acidium Statices Desm. ; Cooke, Micr. Fung. p. 197; Grevillea, i. 7.

Uromyces Limonit Lév. Dict. Hist. Art. Uréd. p. 19. Cooke, Handb.
p- 518; Micr. Fung. p. 212. Plowr. Ured. p. 122, p.p. Sace. Syll.
vii. 582 p.p. Sydow, Monogr. ii. 41.

Aicidiospores. Aicidia amphigenous, often on red or brownish
spots, in roundish clusters or elongated along the nerves, usually
shortly cylindrical, whitish, with a torn margin; spores densely
and minutely verruculose, yellowish, 21—82 x 18—26 yp.

Uredospores. Sori amphigenous, scattered, generally round-
ish or, on the stem, oblong, long covered by the epidermis, at
length naked, pulverulent, cinnamon; spores varying from
globose to oblong, densely verruculose with minute papille,
yellowish-brown, 22—382 x 20—28 yw; epispore 14—24 uw thick,
with two or three germ-pores.

Teleutospores. Sori amphigenous or caulicolous, scattered
or circinate, roundish or oblong,
long covered by the epidermis,
pulvinate, black; spores subglobose
or more frequently oblong or clavate,
sometimes rounded, sometimes at-
tenuated at the apex, where the wall
is up to 10 w thick, attenuated
below, smooth, brown, 24—50 x 14—
25 w; pedicels as much as 80 u long,
thick, pale-brownish, persistent.

ig. 40. U. Limonii. Teleuto- :
a he Sn os ne ea On leaves and stems of Stutice

Timonium. Not common. Atcidia

in June and July; uredo- and teleutospores from July to
October. (Fig. 40.)

ON ELOMBAGINAEES! 89

This species was formerly united with U. Armertae (q.v.), but the
teleutospores are distinctly different.

DISTRIBUTION: Europe, North Africa, Siberia and North
America.

4. Uromyces Armerie Lév.

Caeoma Armeriae Schlechtd. Fl. Berol. ii. 126.

Uromyces Armeriae Lév. Ann. Sci. Nat. sér. 3, viii. 375. Sydow,
Monogr. ii. 40. Fischer, Ured. Schweiz, p. 52, f. 39.

U. Limonit Plowr. Ured. p. 122 p.p. Sace. Syll. vii. 532 p.p.

Spermogones. Scattered among the ecidia, honey-coloured.

4icidiospores. Aiicidia amphigenous, scattered or in small
clusters, at first hemispherical, then cup-shaped, with a whitish
incised margin ; spores densely and minutely verruculose, yellow,
17—28 x 16—22 p.

Uredospores. Sori amphigenous, sometimes on purplish
spots, rounded or elongated, surrounded or half-covered by the
cleft epidermis, pulverulent, cinnamon; spores globose to
oval, very densely and minutely verruculose, yellowish-brown,
24—32 x 21—28 w; epispore 24—3 w thick, with two or three
germ-pores.

Teleutospores. Sori similar, dark-brown; spores globose to
ovate, rounded and thickened (7 «) at the
apex, with a broad flat cap, usually
rounded below, smooth, brown, 24—36
x 21—32 w: pedicels hyaline, nearly as
long as the spore, seldom persistent.

On leaves and peduncles of Armeria
maritima. Not uncommon. Aicidia in
May and ;

y June; uredospores from J Be at annette
onwards; a few teleutospores begin to Teleutospore and
appear in the uredo-sori towards the end HEEB ORBAES:
of July. (Fig. 41.)

This species was united by Plowright with U. Limonii, but is distin-
guished by the more readily pulverulent sori, the shorter and broader
teleutospores, and the shorter hyaline pedicel which is easily detached.

The distinctness of the two species does not seem, however, to have been
tested by experimental cultures. Though the uredo- and teleutospores

90 UROMYCES

have occurred for many years consecutively on Thrift in my garden, I
have never noticed the ecidia; the uredospores last through the winter
on the evergreen leaves, and reproduce the fungus about June ; teleuto-
spores are rather scarce.

DistTRIBUTION : Central and North-Western Europe.

5. Uromyces Trifolii Lév.

Puceinia Trifolii Hedw. f. in DC. Flor. fr, ii, 225.

Oromyces Trifolii Lév. Ann. Sci. Nat. ser. 3, viii. 871. Plowr. Ured.
p. 124 pp. Sacc. Syll. vii. 534 p.p. Sydow, Monogr. ii. 132.
Fischer, Ured. Schweiz, p. 23 p.p.

Trichobasis fallens Cooke, Micr. Fung. p. 226 p.p.

Puccinia fallens Cooke, Handb. p. 508 p.p.

Nigredo fallens Arthur, N. Amer. FI. vii. 254.

Uredospores. Sori hypophyllous and on the petioles,
and like those of U. Trifoli-répentis ;
spores differing in having 5—7 germ-

pores.
RK Teleutospores. Indistinguishable from
7 those of U. Trifoli-repentis.
ig. 42. U. Trifolii. . ; : j
Se ee a v. On Trifolium hybridum, T. incarna-
pratense. tum, T. medium, T. pratense. Not com-

mon. (Fig. 42.)

Liro proved by culture-experiments that the U, romyces on Trifolium
repens could not be transferred to T. hybridum or T. pratense. Since this
difference is accompanied by the absence of the ecidium in the latter
species and by a difference in the number of the germ-pores, they are
considered distinct by Sydow. But on Trifolium pratense I have found
uredospores with not more than four gerra-pores, each covered with «a low
flat hyaline cap. In that case only the absence of the excidium would
separate the two forms, though the average number of germ-pores is no
doubt different in the two cases.

An ecidium has been found elsewhere on 7. pratense, but this has
been experimentally proved (Dietel, Flora, 1895, Ixxxi. 398) to belong to
another European and North American species, U. ménor Schrét. = VU. ob-
longus Vize (Grevillea, v. 110), which has no uredospores, but only ecidio-
and teleutospores.

Cooke’s species, U. apiculatus (Grevillea, vii. 136), is indefinite ; the
form on clover may belong here, that on Luthyrus pratensis to U. Pist.

DisTRIsurion: Europe, Asia Minor, Persia, North America,
ete.

ON LEGUMINOSA 91

6. Uromyces Trifolii-repentis Liro.

U. Trifolii-repentis Livro, Act. Soc. Faun. Flor. Fenn. xxix. 15. Sydow,
Monogr. ii. 131. Fischer, Ured. Schweiz, p. 23, f. 19.

U. Trifolii Plowr. Ured. p. 124 p.p. Sace. Syll. vii. 534 p.p.
McAlpine, Rusts of Australia, p. 97, f. 142, & pl. G, f. 32.

Trichobasis fallens Cooke, Micr. Fung. p. 226 p.p.

Puccinia fallens Cooke, Handb. p. 508 p.p.

Wigredo Trifoliz Arthur, N. Amer. FI. vii. 255.

Spermogones. Epiphyllous, honey-coloured, forming minute
clusters.

Aicidiospores. Aicidia hypophyllous, in clusters, roundish
on the leaves and as much as 5 mm. long on the nerves and
petioles, shortly cylindrical, whitish-yellow; margin white, torn,
hardly revolute; spores minutely verruculose, yellowish, 17—
21 x 14—18 p.

Uredospores. Sori hypophyllous and on the petioles,
scattered over the leaves or gregarious, small or rarely confluent
and larger, soon naked, pulverulent, pale-brown ; spores globose,
ovate or ellipsoid, echinulate, yellow-brown, 19—26 x 17—24 » ;
epispore about 14 thick, with two to four (generally two)
equatorial germ-pores.

Teleutospores. Sori surrounded by the cleft epidermis,
similar, but elongated on the
petioles, and darker brown;
spores globose to ovate, rounded
at the apex, with a very small
hyaline papilla, smooth or at
times bearing a few minute
warts arranged more or less in Fig. 43. U. Trifolii-repentis. a,
lines, brown, 18—30 x 16—25y;, —ecidia on young leaf; b, uredo-

j : : and teleuto-sori, on later-formed
epispore about 2u thick; pedi- — jear,
cels short, thin, hyaline, decidu-
ous.

On. leaves and petioles of Trifolium repens. Aicidia (rare)
from April and uredospores from May onwards. (Fig. 43.)
This species is distinguished from U, Trifolix Lév. by the smaller

number of germ-pores of the uredo, and also by the presence of the ecidia,
which cause long crooked swellings on the petioles and nerves, but not on

92 UROMYCES

the leaves. The mycelium of the ecidial stage is said to be perennial in
the host; Dietel says that in some localities the scidiospores can re-
produce themselves, and that then the uredo is suppressed.

Both this species and the preceding are distinguished from U. flectens
in the fact that the sori are smaller, distributed more uniformly over the
leaf, and do not cause distortions.

The zcidium is rare in Britain (I have seen specimens only from Perth) ;
most of our records of Uromyces on T. repens belong to the following
common species, U. flectens. Pseudopeziza, Trifolii (a Discomycete) is
common on leaves of white clover and is not infrequently mistaken for
the uredo-stage of U. Trifolii-repentis, but is distinguishable by its being
confined to the wpper surface of the leaves. No practical means of
prevention are known for either the Clover Rust (Uromyces) or the Clover
Leaf-spot (Pseudopeziza).

DISTRIBUTION: Europe, Asia Minor, Persia, North and
South America, Australia.

7, Uromyces flectens Lagerh.

Uromyces flectens Lagerh, Svensk Bot. Tidskrift, iii. 36. Sydow,
Monogr. ii. 360. Grove, Journ. Bot. 1911, p. 366.
Puecinia neurophila De Toni, Sacc. Syll. vii. 698.

Leleutospores. Sori hypophyllous, or more often on the
nerves and petioles where they cause swell-
ings and distortion, scattered, rather large,
4—2 mm. long or even confluent and larger,
long covered by the epidermis, then pul-

= verulent, dark-brown; spores as in U.

Fig. 44. U. flectens. TOON TEDENDS.

Teleutospores on 1’. On Trifolium repens. May—October.

repens. Common. (Fig. 44.)

It has been frequently noticed that the Uromyces on Trifolium repens
behaves differently in different localities ; sometimes forming teleutospores
only, from May to October; at others forming both ecidia and uredospores
during the same time. Plowright records an interesting experiment which
he performed (Ured. p. 125) ; in October he brought a plant of 7. repens,
with the Uromyces upon it, indoors and kept it there till the following
summer. During all this time it produced only teleutospores. Lagerheim,
in 1909, noticing that the form which produced only teleutospores had
sori which were larger, more predominant upon nerves and petioles, and

remained longer covered by the epidermis, described this as a distinct
%

ON LEGUMINOSA 93

species, to which evidently Plowright’s specimen may be ascribed. Cooke’s
figure of his U. apiculosa, on Trifolium repens (Micr. Fung. pl. vii. f. 154),
is probably the same species. ,

DIsTRIBUTION : Middle Europe and Persia.

8. Uromyces striatus Schrit.

Uredo apiculata vay. Trifolit Strauss, Ann. Wett. ii. 97 p.p.
Uromyces striatus Schrét. Abhandl. Schles. Ges. 1872, p. 11. Sace. Syll.
vil. 542 p.p. Sydow, Monogr. ii, 115. Fischer, Ured. Schweiz,
p. 31, f. 24. Cf. Plowr. Ured. p. 134.
[ Spermogones
Ajceudvospores
Uredospores. Sori amphigenous, scarcely ever on the nerves,
without spots, scattered, occasionally aggregated and confluent,
minute, pulverulent, cinnamon; spores globose to ellipsoid,
faintly and sparsely echinulate, yellowish-brown, 15—22 yw;
epispore 14—2,y thick, with 4—6 or even more germ-pores,
each with a small hyaline cap.

E As in Uromyces Pisi.]

Fig. 45. U. striatus. a, two teleutospores on T. minus (ex herb. Broome) ;
b, a teleutospore on 7. arvense (foreign, ex herb. De Thiimen); c, uredo-
spore from the same leaf as a.

Teleutospores. Sori similar, but darker; spores globose to
ovate, with a minute and narrow papilla, striated from apex to
base by longer or shorter lines of warts, brown, 18—24 x 15—
20 w; epispore 14—2 pw thick ; pedicels short, hyaline, deciduous.

[Acidia on Euphorbia Cyparissias, not known in Britain ;]
uredo- and teleutospores on leaves and stems of Trifolium
minus. Bath (Herb. Broome); King’s Norton (Worcestershire).
Very uncommon. July—August. (Fig. 45.)

See remarks made about the acidial stage under U. Pist. Schréter
proved the connection of an ecidium on Euphorbia Cyparissias with an

exactly similar Uromyces on Trifolium agrarium. U. striatus is found
elsewhere on Trifolium procumbens and also on many species of Medicago,

94, UROMYCES

including all the British species, but I have seen no specimens on these
from this country. The teleutospores on 7. minus which I have observed
are more distinctly verrucose and less striated than in the figures given by
Fischer, and may possibly not belong to the same species.

DistRIBUTION: Europe, North and South America, East
Indies.

9. Uromyces Loti Blytt.

Uromyces Loti Blytt, Christ. Vidensk.-Selskabs Forhandl. 1896, p. 37.
Sydow, Monogr. ii. 110. Grove, Journ. Bot. 1911, p. 367.

U. Euphorbiae-Corniculati Jordi, Centralbl. f. Bakt. 1904, 2. xi. 791.
Fischer, Ured. Schweiz, p. 34, f. 26.

[Spermogones. Hypophyllous, numerous, scattered amongst
the wcidia.

Mediospores. Aicidia distributed uniformly over the lower
surface of the leaf, cup-shaped, with a torn white revolute

Fig. 46. U. Loti. Four teleutospores, all on L. corniculatus ; a shows
how the spores look when wet, the others are viewed dry.

margin; spores densely and minutely verruculose, orange,
18—23 p.}

Uredospores. Sori amphigenous, but mostly hypophyllous,
scattered, minute, round, sometimes confluent, surrounded by
the cleft epidermis, soon naked, pulverulent, cinnamon ; spores
globose to ellipsoid, with short blunt and rather distant spines,
brownish, 17—25 x 16—23,; epispore 24—3,y thick, with
2—5 germ-pores.

Teleutospores. Sori similar, but darker in colour; spores
globose to obovate, often with a low flat pore-cap at the apex,
which is not thickened, beset with minute warts and ridges
which are often arranged in undulating longitudinal lines,
brown, 17—25 x 14—21 »; pedicels short, hyaline, deciduous,

ON LEGUMINOSA 95

[Mecidia on Euphorbia Cyparissias;] uredo- and teleuto-
spores on Lotus angustissimus, L. corniculatus; July, August.
(Fig. 46.)

Plowright refers to this species (but not as British) in a note on
p. 134. The markings on the teleutospore are very delicate and can
scarcely be seen except when the material is fresh and the spores are
viewed dry. I found that the longitudinal lines of warts were more
strongly marked and auastomosed more frequently on spores from L.
corniculatus than from L. angustissimus (Newquay, Cornwall), on which
they were fainter and more irregular, but this difference may have been
partly due to the fact that the latter had been gathered (by Dr Vigurs)
many years before they were examined. Jordi proved that acidiospores
from £. Cyparissias would freely infect L. corniculatus. But see U. Pisi.

DistrisuTion: Western, Central and Southern Europe,
and Japan.

10. Uromyces Anthyllidis Schrot.

Uredo Anthyliidis Grev. in Sm. Eng. FL. v. 383.

Uromyces Anthyllidis Schrét. Hedwig. xiv. 162. Plowr. Ured. p. 135.
Sacc. Syll. vii. 551. Sydow, Monogr. ii. 64. Fischer, Ured.
Schweiz, p. 36, 543, f. 28.

Uredospores. Sori amphigenous, widely and irregularly
scattered, or sometimes with a circle of small ones
round a larger one, minute, roundish, black and

shining, soon naked, then pulverulent, cinnamon ; fl
spores globose or subglobose, sparsely and finely
echinulate, yellowish-brown, 18—25u; epispore 3—

Big. 47. 0.

34 thick, with 4—6 germ-pores (4—5, Bubak; “snenyiiidis.
5—8, Fischer). . Uredospore.

Teleutospores. Sori similar, but darker in colour. Spores
globose to ovate, with a minute papilla at the rounded apex,
verrucose, brown, 16—22 x 15—20 m4; epispore rather thick ;
pedicels short, hyaline, deciduous.

On leaves of Anthyllis Vulneraria. Notcommon. June—
October. (Fig. 47.)

It is probable that this species occurs only on A. Vulnerarta (and on
the continent, A. maritima), but it has many close allies on other
Leguminose. Teleutospures are rarely formed ; in specimens gathered in
mid-September I have found only one or two, in the midst of abundant

96 UROMYCES

uredospores. The warts on the teleutospores are not numerous and are
rather easy to see.

DistriBuTION: North-western and Middle Europe.

11. Uromyces Ervi Westendorp.

Heidium Ervi Wally, Fl. Crypt. Germ. ii. 247.

Uromyces Ervi Westd. Bull. Acad. Roy, Sci. Belg. xxi. pt. 2, p. 246, f. 3.
Plowr. Ured. p. 140, Sydow, Monogr. ii. 96. Fischer, Ured.
Schweiz, p. 69, f. 53.

Acidiospores. Aicidia amphigenous, or on the petioles,
solitary or 2—8 together in little scattered groups, cup-shaped,
whitish ; margin faintly revolute, scarcely torn; spores densely
and minutely verruculose, pale-yellowish, 16—25 x 14—18 y.

Uredospores. Sori rarely formed, amphigenous or on the
petioles and stems, scattered, minute, ob-
long,surrounded by theruptured epidermis,
cinnamon ; spores ovate or ellipsoid, dis-
tantly echinulate, brownish-yellow, 20—
30 x 18—22 w, with two (rarely three)
germ-pores.

; Leleutospores. Sori amphigenous, or
Fig. 48. U. Ervi, Leaf -

of E. hirsutwn, with ™ore frequently on the petioles and stems,

pall slightly en- scattered, minute, oblong, surrounded by

arged; two teleuto- : a 3

spores. the ruptured epidermis, blackish-brown ;
= spores subglobose to obovate, usually
darker and rounded above (where the wall is up to 8 w or more
thick), rounded or attenuated at the base, smooth, brown,
20—28 x 14—20 4; pedicels brownish, persistent, as long or
twice as long as the spore.

On leaves, petioles, and stems of Hrvum hirsutum (Vicia
hirsuta). Acidia, May—October; teleutospores from July
onwards, lasting through the winter on the dead stems. (Fig. 48.)

It has been proved by many culture experiments that Plowright was
correct in his belief that this species is strictly confined to the one host.
The aecidiospores are capable of reproducing the ecidium and are found
throughout the season ; the uredospores are, perhaps in consequence, not
abundant, only a few being occasionally found and usually intermixed
with teleutospores.

DisTRIBUTION: Europe, Japan.

ON LEGUMINOSA: 97

12. Uromyces Fabe De Bary.

Uredo Fabae Pers. in Rém. Neu. Magazin, i. 93.

Uromyces Fabae De Bary, Ann. Sci. Nat. ser. 4, xx. 72. Plowr. Ured.
p. 119. Sace. Syll. vii. 531 p.p. Sydow, Monogr. ii. 103.
Fischer, Ured. Schweiz, p. 65, f. 49—51. McAlpine, Rusts of
Australia, p. 93, f. 307.

Trichobusis Fabae Cooke, Handb. p. 508; Micr. Fung. p. 225.

Uromyces appendiculatus Lév.; Cooke, Micr. Fung. p. 212, pl. vii.
f. 149—150 p.p.

Puccinia Fabae Link, referred by Cooke to this species, has no
existence in nature (Handb. p. 508; Micr. Fung. p. 211).

Spermogones. Hypophyllous, growing among the ecidia.
Aicidiospores. Aicidia hypophyllous, seated on pale-yellow
spots, solitary or in small round or elongated clusters, shortly
cup-shaped, with a whitish, torn, revolute margin; spores
densely and minutely verruculose, yellow, 14—22 yu.
Uredospores. Sori amphigenous, scattered or circinate, girt
by the ruptured epidermis, minute, pulverulent, pale-brown ;
spores globose to ovate, distantly echinulate, at length pale-
brown, 20—30 x 18—26 «4; epispore 14—24 yw thick, with three
or four germ-pores.
Teleutospores. Sori similar, but per-
sistent and darker or blackish-brown;
spores subglobose to obovate, rounded or
truncate and thickened above, where the
wall is dark and 7—11, thick, sometimes
with a colourless papilla, smooth, brown,
25—38 x 18—27 »; pedicels brownish,
persistent, thick and as much as 40—70 w
long.

On leaves and stems of Fuba vulgaris,
Lathyrus pratensis (?), Pisum sativum,
Views Cracca, V. sativa, V. septum. Alcidia
in April, May; uredospores from May,
teleutospores from July onwards, lasting : :
Fig. 49. U. Fabae.

through the winter on the dead stems. eleutospores on stem
(Figs. 49—52.) of Broad Bean.

One of the most widely spread of the Uredinales, occurring in every
G. U. 7

98

UROMYCES

part of the world; reported on many Leguminose, but doubtless some of
these are distinct species. Jordi has distinguished under U. Fabae three

Fig, U. Fabae.

yo YP

Teleutospores

a ae oe on Vicia Cracca.

“biological” races-—(

Fig. 51. U. Fabae. Telento-
spores and uredospore on
Vicia sepium.

1) on Faba vulgaris and Pisum sativum, (2) on

Lathyrus vernus and probably also on Pisum sativum, (3) on Vieta Cracca,
Pisum sativum, and possibly also Vicia hirsuta.
The ecidial generation is frequent on some hosts, such as eva sepium

Fig. 52. U. Fabae.
Ecidia on Pea, from
Plowright’s culture,
iu which he produced
them on both Pea
and Bean from the
same teleutospores.

and species of Lathyrus. On Faba vulgaris aud
Pisum sativum it is, on the contrary, very rare,
being recorded by Sydow, on the Pea, only from
Norway, East Indies and Japan (once from each).
It has been seen on the Bean in the East Indies, but
seems to be not uncommon in artificial cultures,
in which Plowright produced it both on Pea and
Bean from the same teleutospores (Plowr. Ured.
p. 121).

On the leaves of the common field Bean only
the uredospores are generally to be found, even as
late as mid-October, but on the stems the teleuto-
spores form large black sori. On Vicia sepium
the uredo-sori are often darker and covered by the
epidermis for a shorter time than on the Bean,

while the teleuto-sori occur in great abundance on the leaves and even

on the tendrils.
If all the infected

haulm, etc., were burnt instead of being put on the

manure heap or left to rot on the ground, the disease would become less
prevalent, especially if Jordi’s idea is true, that the Rust on the wild
Vetches is a distinct biological race. In Ecuador, at Quito, which has a

very equable climate

of “perpetual spring,” U. Fabae has, according to

Lagerheim, become almost an isolated uredo. The same thing is true of
it in other tropival climes.

DISTRIBUTION :

world-wide.

ON LEGUMINOSZ : 99

13. Uromyces Orobi Lév.

Aicidium Orobi Pers. in Rém. Neu. Magazin, i. 92. Cooke, Handb.
p. 542; Mier. Fung. p. 197.

Uromyces Orobi Lév. Ann. Sci. Nat. ser. 3, vill. 371, 376. Plowr. Ured.
p. 121. Sydow, Monogr. ii. 106. Fischer, Ured. Schweiz, p. 69,
f. 52.

Spermogones. Hypophyllous, mixed with the ecidia.
Aicidiospores. Aicidia hypophyllous, on yellowish spots, in
dense clusters 1—5 mm. long, rarely soli-
tary, shortly cup-shaped, with whitish revo-
lute margin; spores densely and minutely
verruculose, yellowish, 14—21 yp.
Uredospores. Sori amphigenous, scat-
tered, minute, punctiform, pulverulent,
brown; spores globose to ovate, distantly
echinulate, pale-brown then darker, 20—
28 x 18—25 pw; epispore 3—4 w thick, with
three or four germ-pores. ; ;
ea Sori similar, but darker; i ee
spores subglobose to ovate, rounded or sub- a
conical above, and much thickened (7—1]1 y) and darker, smooth,
brown, 25—35 x 18—28 yu; pedicels persistent, yellowish, thick,
as much as 100 » long.

On Orobus tuberosus (Lathyrus montanus = L. macrorrhizus).
Acidia in May and June; uredo- and teleutospores from June
onwards. (Fig. 53.)

Cooke says that the ecidia also occur on the stems. The thicker
membrane of the uredospores, which is also less strongly echinulate,
distinguishes them from those of U. Fubae. Jordi made attempts to
infect various other species of Lathyrus, and also species of Vicia and
Pisum, from U. Orobi, but in every case without success.

DistRiBuTion : North-western and Central Europe.

14. Uromyces Pisi Wint.

Uredo appendiculata var. Pist Pers. Obs. Myce. i. 17.
Ascidium Cyparissiae DC. Flor. fr. ii, 240.
Uromyces Lathyri Fckl. Symb. Myc. p. 62.

100 UROMYCES

U. Pist Winter, Krypt. Flor. i. 163. Cooke, Grevillea, vii. 135. Plowr.
Ured. p. 188. Sacc, Syll. vii. 542 p.p. Sydow, Monogr. ii. 124.
Fischer, Ured. Schweiz, p. 28, f. 22.

Spermogones. Hypophyllous, numerous, scattered amongst

the secidia.

Aicidiospores. Aicidia distributed uniformly over the lower
surface of the leaf, cup-shaped, with a white, torn, broadly
revolute margin; spores densely and minutely verruculose,
orange, 18—23 py.

Uredospores. Sori generally hypophyllous, scattered, minute,

soon naked, pulverulent, cinnamon; spores
, : globose or subglobose, minutely verruculose,
fi ! yellow-brown, 21—25 yw diam.; epispore 14—
8 E 24 thick, with 3—5 germ-pores.
ne Teleutospores. Sori similar, but sometimes
confluent and larger, dark-brown; spores sub-
ae he as globose to ovate, with a small hyaline papilla
spermogoneson (as much as 3m high), everywhere minutely
leaf of Euphor- FT her a ] l ti 20
bia Cyparissias 2d rather densely verruculose, brown, 20—
ee 28 x 14—22 #3 epispore 1} thick; pedicels
hyaline, short, deciduous.

Acidia on Euphorbia Cyparissias, May, June; uredo- and
teleutospores on Piswm sativum and Lathyrus pratensis, July—
September. Rare. (Fig. 54.)

Although both will equally infect 2. Cyparissias, it is probable that
the Uromyces on Pisum is biologically distinct from that on Lathyrus. It
is not certain that the latter has been found in this country, but the
former is recorded from various places. It must be remembered that
U. Fabae occurs also on the same two genera, though all the spore-forms
of the two can be easily distinguished.

An ecidium on £. Cyparissias, and attributed to U. Pis?, was found
at Dover, May, 1909 (Rev. T. Taylor); the specimen is in the British
Museum, but there is no proof that this belonged to U. Pisz, because it
has been shown that U. Loti, U. striatus (both of which are British), as
well as two other (non-British) species, equally produce on EZ. Cyparissias
wcidia which are norphologically indistinguishable. This acidium pos-
sesses a perennial mycelium, which permeates the whole host and deforms
and. bleaches it. The connection of one form of it with Uromyces Pist
has been experimentally demonstrated by Schroter, Rostrup, Fischer and
others. The Uromyces on Vicia Cracca which was formerly considered to

ON LEGUMINOSA 101

belong to U. Pisi has been proved by Jordi to be confined to that species
and not to be transmissible to Pisum sativum or Lathyrus. It has been
named by Magnus U. Fischeri-Eduardi, but is not known as British.

DIsTRIBUTION: Europe generally; North America less
commonly.

15. Uromyces Phaseolorum De Bary.

Ateidium Phaseolorum Wall. Fl. Crypt. Germ. ii. 256.
Uredo appendiculata var. Phaseoli Pers. Syn. p. 222.

Uromyces Phaseolorum De Bary, Ann. Sci. Nat. ser. 4, xx. 80 (1863).
Cooke, Grevillea, vii. 135.

U. Phaseoli Wint. Pilze, p. 157 (1884). Plowr. Ured. p. 122. York-
shire Fung. FI. p. 186.

U. appendiculatus Link, Obs. ii. 28. Sacc. Syll. vii. 535. Sydow,
Monogr. ii. 120, Fischer, Ured. Schweiz, p. 19, f. 16. McAlpine,
Rusts of Australia, p. 92, f. 306 (all pro parte).

Nigredo appendiculata Arthur, N. Amer. FI. vii. 257 p.p.

[Spermogones. In little clustars, whitish, then yellowish.
Aeidiospores. Aicidia hypophyllous, clustered in little
roundish groups 2—3 mm. wide on yellowish or brownish spots,
cup-shaped, whitish, with a torn revolute margin; spores poly-
gonal or oblong, densely and minutely verruculose, colourless,
18—36 x 16—24 y.]
Uredospores. Sori generally hypophyllous, on indistinct
spots, scattered or in little clusters here
and there, minute, soon naked, surrounded
by the cleft epidermis cinnamon; spores
subglobose to ovate, distantly but sharply
echinulate, brownish.-yellow, 18—-28 x 18
—22; epispore brownish-yellow, about
1} » thick, with two germ-pores; contents pig 55. 1, Phaseolorum.
colourless. Teleutospore and ure-
Teleutospores. Sori similar, but con- BOERS:
fluent, larger, amphigenous and blackish-brown; spores sub-
globose to ovate, rounded above, with a wide germ-pore and a
hemispherical hyaline papilla, smooth or rarely provided, espe-
cially neav the apex, with a few hyaline warts, chestnut-brown,
24—-35 x 18—25 «; epispore up to 34 yu thick; pedicels hyaline,
rather thin, about as long as the spore.

102 UROMYCES

On leaves of Phaseolus vulgaris. May, July—October ;
even earlier on forced plants. Uncommon. (Fig. 55.)

De Bary (J.c.) proved the genetic connection of the xcidia with the
uredo- and teleutospores. The xcidia are rarely met with; they may
occur either before or in company with the other spore-forms. I have
seen no proof that they have been found in this country. The description
is founded upon that of Sydow.| Fischer says that this species is very
common in Switzerland on Piast it may become a dangerous parasite
on forced Beans. All affected plants (leaves and stems) should be burnt.
U. appendiculatus of Sydow, which occurs on many Leguminosz, is
probably a collective species, though no experiments bearing on this point
are available. \

DISTRIBUTION : as a collective species (U. appendiculatus)
world-wide. a

16. Uromyces tuberculatus Fckl.

Ateidium Euphorbiae Gmel, in Linn. Syst. Nat. ii, 1473 p.p. Purton,
Mid, Flor. iii. 293. Cooke, Handb. p. 537 ; Micr. Fung. p. 195 p.p.
Plowr. Ured. p. 270.

Cromyces excavatus DC. ; Cooke, Grevillea, ii. 161; Micr. Fung. p. 213.

U. tuberculatus Fckl. Symb. Myc. p. 64. Sydow, Monogr. ii. 165.
Fischer, Ured, Schweiz, p. 43, f; 33.

U. proeminens Lév. ; Sace. Syll. vii. 553 p.p.

Spermogones |

Aicidiospores \ *
whole leaf; secidia immersed, cup-shaped, with a short denticu-
late margin ; spores orange, densely ver1.uculose, 17—25 x 14—
20 p.

Uredospores. Sori hypophyllous, scatte.-ed, at length naked,
cinnamon; spores more or less
globose, yellowish-brown, aculeo-
late, 20—25 uw; epispore 14—2) 4
thick, with 5—7 swollen germ-
pores (45, Fischer).

Fig. 56. U. tuberculatus. Two ma- Peleutosp Ores: Sori amphi-
ture teleutospores; «, a teleuto- genous and on the stems, round,
ice the tubercles are scattered or sometimes arranged

in little groups, pulverulent,
blackish-brown or black; spores globose to ellipsoid, occasionally

Hypophyllous, svread uniformly over the

ON GERANIACEAE 103

with a flat, broad, hyaline papilla, at first smooth, then covered
with more or less distant, broadly conical, obtuse, subhyaline
warts, chestnut-brown, 20—30 x 18—24 w; epispore 2—24 uw
thick; pedicels hyaline, deciduous.

On Euphorbia exigue. Rare; Hampshire, Mr Hill (Plowr.
l.c.); King’s Cliffe, Norths. (Grevillea, l.c.). Midlands (Purton,
ic). (Fig. 56.)

For a long time this species was considered to have only uredo- and
teleutospores, but the connection of these with the zcidium occurring on
the same species of Huphorbia was established by Tranzschel. Berkeley,
at King’s Cliffe, found them all together. The description given above is
partly founded upon those of Tranzschel, Sydow, and Fischer. The
mycelium of the ecidial stage infests the whole plant, that of the teleuto-
spores is more or less localised.

DISTRIBUTION: France, Germany, Switzerland.

17. Uromyces Geranii Otth et Wart.

Adium Geranit DC.; Cooke, Handb. p. 543; Micr. Fung. p. 199 p.p,

Trichobasis Geranti Cooke, Handb. p. 530.

Uromyces Geranit Otth et Wartm. Schweiz. Krypt. no. 401. Cooke,
Micr. Fung. p. 213. Plowr. Ured. p. 126. Sacc. Syll. vii. 535.
Sydow, Monogr. ii. 190. Fischer, Ured. Schweiz, p. 16, f. 14.

Spermogones. Mixed with the ecidia, orange.

4Aicidiospores. Aicidia hypophyllous or on the petioles, on
the leaves chiefly in the vicinity of the nerves and there
forming large dense clusters on thickened spots, on the petioles
forming elongated clusters and often causing great distortion,
at first hemispherical and closed, then opening by a round pore,
at length with a very slightly revolute incised margin, orange ;
spores somewhat ovate, densely and minutely verruculose, yellow,
2228 x 18—24 wu; epispore rather thick.

Uredospores. Sori hypophyllous, generally on brownish or
reddish-yellow spots, scattered or gregarious, minute, rounded,
pulverulent, cinnamon, surrounded by the cleft epidermis;
spores globose to obovate, sparsely echinulate, brown, 20—80 x
18—24 4; epispore about 2, thick, with one (rarely two)
germ-pores.

104 UROMYCES

Teleutospores. Sori similar, but less pulverulent, and
blackish-brown; spores subglobose to ovate,
not thickened above, but with a hyaline
papilla as much as 6 uw high, smooth, brown,
2235 x 1825 w; pedicels short, hyaline,
deciduous.

tie, 57. i Gerani, On Geranium dissectum, G. molle, G.
Teleutospores on G. pratense, G. pyrenaicum, G. silvaticum.
Cane Not common. Atcidia, March to June;

teleutospores, June—October. (Fig. 57.)

Liro proved that the ecidium of this parasite from G@. silvaticum
produced uredo- and teleutospores on the same plant, and Bock showed
that the uredospores from the same species reproduced themselves on
other hosts of the same genus. But there is another ecidium occurring
on G@. pratense and G. silvaticum, which belongs to a quite different life-
cycle. This is cidium sanguinolentum Lindr., and is the ecidial stage
of the hetercecious Puccinia Polygont-amphibii Pers. (q.v.). It differs
from the ecidium of U. Geranit in being seated on conspicuous blood-red
or deep-purplish spots which are not distinctly thickened ; moreover the
shape of the spores is that usual in ecidiospores, viz. rounded-polygonal,
while those of U. Geranii are always more or less ovate, and have a
thicker wall.

Again, there is an ecidium on G. pusillum which, according to Sydow,
is probably also found on G. molle and @. rotundifol/um, and which belongs
to Puceinia Polygoni-Oonvolvuli (q.v.)—a form of P. Polygoni-amphibii
which is often separated as a distinct species. The uredo- and teleuto-
sori would, of course, not follow the wzcidium on the same plant in either
of these two cases.

On G. pyrenaicum there is another Uromyces (U. Kabatianus) which
differs in the arrangement of its sori ; see below.

DIsTRIBUTION: Europe, except in the extreme South.

18. Uromyces Kabatianus Bubdk.

Uromyces Kabatianus Bubdk, Sitz. kin. béhm. Gesell. Wissen. 1902,
p. 1, f 1-5. Sace. Syll. xvii, 249. Sydow, Monogr. ii. 194.
Fischer, Ured. Schweiz, p. 18, f. 15.

Spermogones. Amphigenous, few, large, honey-coloured,
then darker, on the same spots as the ecidia.

cidiospores. ABcidia hypophyllous, on round yellowish
spots, in little clusters 2—4 mm. wide, hemispherical, opening

ON GERANIACE 105

by a pore; spores roundish-polygonal to oblong-ovate, densely
verruculose, yellow, 24—33 x 18—26 pu.

Uredospores. Sori hypophyllous, on yellow spots, in circi-
nate groups, seldom scattered, rather large, pulverulent, choco-
late-brown ; spores roundish, brown, distantly echinulate, 22—
26 «; epispore about 2 w thick.

Leleutospores. Sori hypophyllous, on yellowish or reddish

QVC

Fig. 58. U. Kabatianus. Teleutospores and uredospore, on G. pyrenaicum.

spots, rather large, covered with the thin silvery-shining epi-
dermis, generally in circinate groups, soon confluent, pulverulent,
brown; spores ovate, often oblong, scarcely thickened above, .
but with the germ-pore provided with a prominent subhyaline
papilla (up to 7 » high), smooth, pale-brown, 22—42 x 183—20 w;
pedicels short, hyaline, deciduous.

On Geranium pyrenaicum, and possibly on G. molle and
G. pusillum. (Fig. 58.)

This has been separated by Bub4k from U. Geranii on the ground of
the circinate arrangement of the uredo- and teleuto-sori, and the more
oblong and longer teleutospores : the description of the ewcidia and teleu-
tospores is taken from that of Bubdék. According to Sydow, Lind has
succeeded in transferring this species from G. pyrenaicum to G. molle and
G. pusiilum. Bub&k thought that only this form occurred on G. pyrenat-
cum, but Bock (Centralbl. f. Bakt. 2, xx. 584) showed that typical
Uromyces Geranii could also be produced on that host.

The two Uromyces are very closely allied, but several distinctions are
alleged: the uredospores themselves are identical, but not the sori ;
according to Sydow. the teleutospores of U. Aabatianus do not appear till
towards the end of October, are paler, longer and more oblong in shape
with a higher papilla, while their sori are generally circinate, paler and
less compact. The teleutospores of U. Geranit appear at the beginning of
summer, the sori are nearly black, rather compact, and more scattered.
I have specimens collected at Cambridge in August, on G. pyrenazcum,
having the uredo-sori in circinate groups on conspicuous yellow spots, and

106 UROMYCES

containing no teleutospores; it appears probable that these belong to
U. Kabatianus, which will no doubt be found in many places, if looked for.

DISTRIBUTION: a few places in Europe and Asia Minor.

19. Uromyces Alchemille Lév.

Uredo Alchemillae Pers. Obs. Mye. i. 98.

Uromyces Alchemillae Lévy. Ann. Sci. Nat. 3, viii. 371 (1847). Plowr.
Ured. p. 187, Sacc. Syll. vii. 558. Sydow, Monogr. ii. 196.
Fischer, Ured. Schweiz, p. 44, f. 34.

U. intrusa Cooke, Handb. p. 519; Micr. Fung. p. 213.

Trachyspora Alchemillae Fckl. Bot. Zeit. xix. 250. Arthur, N. Amer.
Fl. vii. 178.

Uredospores. Sori hypophyllous, radially arranged, occupy-
ing néarly the whole leaf-surface, rounded or elongated, often
confluent and covered by large fragments of the torn epidermis,
then pulverulent, orange, yellowish or even whitish; spores

ellipsoid to oblong, faintly echinulate, orange or yellowish,
16—25 x 14—21 p.

Fig. 59. U. Alchemillae. Teleutospores on A. vulyarix.

Leleutospures. Sori hypophyllous, scattered, rarely con-
fluent, minute, round, pulverulent, brown; spores globose to
obovoid or oblong, not thickened above, coarsely warted, brown,
26—40 x 20—80 uw; epispore 2—24 w thick; pedicels hyaline,
very deciduous, short or rather long; teleutospores are also
formed in the uredo-sori.

On.Alchemilla vulgaris. Common. Uredospores, April—
June; teleutospores, July—October. (Fig. 59.)
The mycelium perennates in the rhizome and grows up with the young

leaves, causing them to stand more erect, making them paler and con-
spicuous, but smaller and often deformed. The separate teleuto-sori are

ON RANUNCULUS 107

formed on other leaves on a localised mycelium, cause no deformation and
are not conspicuous; in them are a few secondary uredospores. The
teleutospores have unusually coarse warts, mostly towards the apex, or are
sometimes nearly or partially smooth. Bub4&k records (Centralbl. f.
Bakter. 2, xvi. 158) that in many trials in three years he could never get
the teleutospores to germinate, and could not artificially produce infection
in Alchemilla, though Klebahn (Zeitschr. f. Pflanzenkr. 1907) did so
readily with the uredospores. This species can be gathered at considerable
altitudes in Wales and Scotland (and as high as 7200 ft. in Switzerland).

DISTRIBUTION : Europe, Asia Minor, Greenland.

20. Uromyces Ficarie Lév.

Uredo Ficariae Schum. Pl. Sall. ii. 232.

Uromyces Ficariae Lév. Ann. Sci. Nat. 3. viii. 390. Cooke, Handb.
p. 518; Micr. Fung. p. 212, pl. 7, f. 156—7. Plowr. Ured. p. 140.
Sacc. Syll. vii. 568. Sydow, Monogr. ii. 208. Fischer, Ured.
Schweiz, p. 13, f. 12.

Teleutospores. Sori amphigenous or on the petioles, about
4 mm. diam., rounded, frequently col-
lected into dense orbicular or elongated
clusters, on pale-yellow spots, especially
on the petioles where they cause not-
able distortion, soon naked, pulverulent,
chocolate-brown; spores more or less
obovate,often irregular, not thickened
above, but with a conical hyaline papilla, Tata re sre
smooth, pale-brown, 22—38 x 18—26 py; Ficaria. é :
pedicels hyaline, deciduous; a few sub-
globose, pale-brownish, faintly echinulate uredospores, each
with three germ-pores, are occasionally found intermixed, but
are usually abortive.

On Ranunculus Ficaria. March to early June. Very
common. (Fig. 60; see also Fig. 79.)

The ecidium on the same host belongs to the life-cycle of Uromyces
Poae, and is considered to have no connection with the Uromyces on
A. Ficaria, though it may be found on the same leaf. Klebahn proved
that the teleutospores reproduce themselves. But there is a curious
conclusion arrived at by Tranzschel, as the result of his experiments

108 UROMYCES

(see Bot. Zeit. lxiii. 75), that an swcidium which he finds on &. Ficaria is
connected with U. Rumicis (q.v.). The spores of U. Ficariae and U.
Rumicis are very similar.

DistriBuTion : Europe generally, except the extreme South.

21. Uromyces caryophyllinus Wint.

Lycoperdon caryophyllinum Schrank, Baier. Flor. ii. 668.

Uromyces caryophyllinus Winter, Pilze, p. 149. Sacc. Syll. vii. 545.
Sydow, Monogr. ii. 210, 362. Fischer, Ured. Schweiz, p. 11, f. 10.
Trans. Brit. Myc. Soc, iii. 122. McAlpine, Rusts of Australia,
p- 102, f. 152—4 and pl. G, f. 30—1.

Nigredo caryophyllina Arthur, N. Amer. FI. vii. 246.

Uredospores. Sori amphigenous or on the stems, sometimes
on pallid spots, scattered, minute, round or oblong, soon
naked, pulverulent, cinnamon; spores globose to ellipsoid,
sparsely echinulate, yellowish-brown, 20—35 x 18—25 pw; epi-
spore 24—8 » thick, with three to five germ-pores.

x

nl

Fig. 61. U. caryophyllinus. Teleutospores and uredospore from the same
sorus, on carnation ; leaf of carnation with two groups of sori.

Teleutospores. Sori confluent and large, mostly oblong,
_surrounded and often covered by the cleft epidermis, sub-
pulverulent, brownish-black ; spores globose to ellipsoid, with

’

ON CARYOPHYLLACEH 109

a flat hyaline papilla, densely and minutely punctate, chestnut-
brown, 20—31 x 18—24 uw; epispore 2—3 uw thick, not thickened
at the summit; pedicels short, hyaline, deciduous.

On Dianthus barbatus, D. Caryophyllus, D. chinensis. On
cultivated carnations practically all the year round. (Fig. 61.)

The “Carnation Rust” was introduced into England on imported
plants about the year 1890; it sometimes occurs as an epidemic, causing
much injury. The. teleutospore-containing sori are often clustered on the
leaves and stems in circinate or elongated swollen patches ; uredospores
are mixed with them. The punctation of the teleutospores is perceptible
only when they are viewed dry, and at the best is very indistinct.

It is stated by Tranzschel and Fischer that this species is hetercecious,
and has its ecidium on Luphorbia Gerardiana, but as this Euphorbia
does not occur in Britain, the parasite probably maintains itself here
without hetercecism. It is remarked by Sydow that the same is true in
Switzerland, at least in certain cases; but see Fischer (p. 530) who
produced the fungus, from an wcidium on E, Gerardiana, on Saponaria
ocymoides, but not on Dianthus. The fungus has now spread round the
world in greenhouses, but only in the sporophytic stage ; the cidium has
not been recognised anywhere except in Europe. The best means of
prevention are (1) the selection of resistant varieties, (2) good and careful
cultivation, especially sufficient ventilation. If spraying is resorted to,
potassium sulphide solution ($ oz. to 1 gallon) ‘is perhaps the best, but
dilute Bordeaux mixture or copper sulphate solution (1 1b, to 50 gallons),
or sponging with a rose-red solution of permanganate of potash have also
been tried. The latter can be used even when the plants are in active
growth. Besides spraying, every infected leaf should be plucked off and
burnt as soon as discovered. This disease must not be confounded with
the outwardly similar “Fairy Ring of Carnations,” caused by Heterosporium,
but the same remedies apply to both. :

DISTRIBUTION : Europe, Western Asia, Japan, South Africa,
North America, Australia.

22. Uromyces Behenis Unger.
4eidium Behenis DC. Encyel. viii. 289. Cooke, Handb. p. 541;
Micr. Fung. p. 197.
Uromyces Behenis Unger, Einfluss d. Bod. p. 216. Cooke, Micr. Fung.
p. 218. Plowr. Ured. p. 138. Sace, Syll. vil. 559. Sydow,
Monogr. ii. 218. Fischer, Ured. Schweiz, p. 64, f. 48.
Acidiospores. Aicidia usually hypophyllous, seated on
spots that vary both in size and colour (yellow or purple)
and are generally very conspicuous, solitary or collected into

110 UROMYCES

clusters, cup-shaped, whitish-yellow, with a torn revolute
margin; spores densely and minutely verruculose, yellowish,

Fig. 62. U. Behenis. a, ecidia on early leaf, b, ecidia on later leaf, of
S. inflata ; two televitospores.

Teleutospores. Sori hypophyllous and on the stems, often
surrounding the secondary ecidia, irregularly scattered, gre-
garious or circinate, rounded or oblong, covered for a considerable
time by the lead-coloured epidermis, rather small and compact,
brownish-black or black ; spores subglobose or obovate, rounded
above and thickened (as much as 11), smooth, pale brown,
25—35 x 20—27 w; pedicels persistent, faintly yellow, thick,
as much as 75 p long.

On Silene inflata (latifolia), S. maritima. Not common.
Acidia and teleutospores, July—October. (Fig. 62.)

The spots occupied by the ecidia vary in colour, but the primary ones
are often tinged or margined with purple. This is one of the species
whose ecidiospores are capable of reproducing the ecidia, as Dietel has
shown (Flora, lxxxi. 395, 1895). The primary ecidia, on the earlier
leaves, are in roundish groups or concentric circles, only a few being
scattered. The secondary ecidia, on the younger leaves, stand more often
singly and are spread over a larger area; the teleuto-sori spring from
the same secondary mycelium or are formed separately. The secondary
wcidia are not confined to the beginning of the season, but continue to be
produced till the end of autumn, being in fact the’ representatives of
the uredo-sori.

On this accouut this species is very interesting biologically. The
primary scidia arise from infection by comparatively few basidiospores ;
the secondary arise from the more widely dispersed acidiospores of the

ON CARYOPHYLLACEEX 111

first generation, and their mycelium can produce either xcidiospores or
teleutospores or both. No spermogones seem to be known.

The ecidium requires to be carefully distinguished from that of
Puccinia Behenis (P. Silenes) (q.v.), which is much rarer and does not
extend throughout the season. ,

DistTRIBUTION : Europe generally.

23. Uromyces sparsus Lév.

Uromyces sparsus Lév. Ann. Sci. Nat. ser. 3, viii. 369. Cooke, Handb.
p. 519; Micr. Fung. p. 214. Plowr. Ured. p. 136. Sacc. Syll.
vil. 545. Sydow, Monogr. ii. 221.

Uredospores. Sori amphigenous and on the stems, on pallid
spots, scattered, roundish, $—1 mm.
diam., convex, covered for a considerable
time by the epidermis, which at length
splits and surrounds them, then pulveru-
lent, pallid-cinnamon; spores globose to
oblong, faintly cchinulate, brownish,
18—28 x 15—22 p.

Teleutospores. Sori similar, but
darker; spores subglobose to oblong,
rounded above where they are slightly
thickened (up to 4) and darker, gene-
rally tapering downwards, smooth, brown, Fig. 63. U. sparsus.

‘ ‘ Teleutospores on

22—32 x 14—21 yw; pedicels persistent, Spergularia salina

aes : 4 (foreign, J. Kunze,
thick, as much as 60 » long, brownish at Exsice., no. 216)
the apex.

On Spergularia rubra (Alsine rubra). May—July. Very
rare; I have seen no British specimens. (Fig. 63.)

DistrisuTion : Mid-western Europe.

24, Uromyces Chenopodii Schrit.

Uredo Chenopodii Duby, Bot. Gall. ii. 899.

Aedium Suaedae Thiim. Fung. egypt. iii. no. 53.

Aecidium Chenopodit, in Gard. Chron. (1895), xviii, 135.

Uromyces Chenopodit Schrot. in Kunz. Fung. Sel. no, 214. Plowright
in Trans. Brit. Myc. Soc. i. 56. Sace. Syll. vii. 548. Sydow,
Monogr. ii. 233.

112 UROMYCES

Aicidiospores. ABcidia amphigenous, clustered in circles 5—

10 mm. diam., cylindrical, whitish, margin deeply torn; spores
delicately verruculose, yellow, 18—22 pw diam.

Uredospores. Sori amphigenous, scattered or gregarious,

round or more frequently elongate, small,

surrounded by the conspicuous torn epi-

dermis, cinnamon; spores ‘globose. to oval
or oblong, delicately and sparingly echinu-
late, yellowish-brown, 18—25 x 16—21 yp;
epispore about 14 u thick.

UT hleisbaiores, Sori amphigenous but
mostly cauline, on the leaves rounded and
1—3 mm. diam., on the stems lanceolate
and even as much as 3 em. long (Sydow),
thick, compact, dark-brown; spores very
variable, oval to subpyriform, rounded
or subconical at the apex, thickened or
not, smooth, brown, 24—385 x 18—20u;

Bis. 80s ethenepale pedicels pale-brown, persistent, up to 80%

Teleutospores on long or more.
Suaeda fruticosa

(foreign, ex herb. De On stems, branches and leaves of
Thiimen). qe :
Suaeda maritima. Rare. Terrington
Marsh (Mr H. G. Ward); North Wootton Marsh (C. B. P.);
August. The ecidia are recorded from Worthing (Miss A. L.
Smith, Journ. of Bot. May, 1898), as well as by Plowright from
North Wootton. (Fig. 64.)

Teleutospores very variable, short and broad or long and narrow in
the same sorus ; thickening of apex also varying from 3 to 7»; pedicels
often very long and flexuous.

The name of this species is misleading; it has been found on S.
Fruticosa, but not on the present-day Chenopodium. Plowright mentions, as
showing the distinctness of this species from U. Salicorniae, that at North
Wvotton Marsh it did not spread to Salicornia herbacea, which was
growing near.

DISTRIBUTION: Germany, and most of the countries in South
Europe and North Africa.

ON CHENOPODIACEE 113
25. Uromyces Bete Lév.

Uredo Betae Pers. Syn. p. 220.

Trichobasis Betae Cooke, Handb. p. 530; Micr. Fung. p. 225,

Uromyces Betae Lévy. Ann. Sci. Nat. ser, 3, viii. 375. Cooke, Mier.
Fung. p. 213. Plowr. Ured. p. 127. Sacc. Syll. vii. 536. Sydow,
Monogr. ii. 224. Fischer, Ured: Schweiz, p. 10, f. 9. McAlpine,
Rusts of Australia, p. 100, f. 148—9, 316, and pl. H.

Nigredo Betae Arthur, N. Amer. FI. vii. 245.

Spermogones. In little clusters, honey-coloured.

ediospores. AKcidia amphigenous, often on rounded or
irregular yellowish spots, collected into rather large clusters
which are round or sometimes irregular and confluent, cup-
shaped, yellowish, with a reflexed incised margin; spores
delicately verruculose, pale-yellowish, 16—24 x 16—20 yu.

Uredospores. Sori amphigenous, scattered, sometimes con-
centrically arranged, thick, pulvinate, Jo
circular, up to 2 mm. diam., covered by aa
the epidermis which at length splits,
then pulverulent, cinnamon; spores
globose to obovate-oblong, sparsely and
minutely echinulate, yellowish, 21—382 x Fig. 65. U. Betae. Te-
16—26 1; epispore 243 thick, with amen tnt a
two equatorial germ-pores.

Teleutospores. Sori similar, but somewhat compact, dark-
brown; spores globose to obovate, rounded and slightly thick-
ened above, with a minute hyaline hemispherical papilla,
smooth, pale-brown, 22—34 x 18—25 w; pedicels short, hyaline.

On leaves of Beta maritima, B. vulgaris; also doing great
harm to cultivated mangels. Acidia rather rare, April—June ;

uredo- and teleutospores, rather common, May—October.
(Fig. 65.)

In May all four spore-forms can sometimes be seen on the same leaf.
Kiihn says that the mycelium of the ecidia is perennial, and that its
spores can reproduce the ecidia. If possible, the first leaves seen bearing
the ecidia should be collected and burnt; this will check the disease at,
the outset... If this is not possible, the plants may be sprayed with dilute
Bordeaux mixture or with potassium sulphide solution. Since in mangels.
the disease would chiefly be reproduced by teleutospores from old leaves

G. U. 8

114 UROMYCES

of the preceding crop, all affected mangel “tops” should be burnt :
rotation of crops is of course a sure preventive, as in all such cases,

DistRiBuTION : Europe, California, South Africa, Australia,
New Zealand.

26. Uromyces Salicornie De Bary.

Beidium Salicorniae DC. Flor. fr. vi. 92.
Uromyces Salicorniae De Bary, in Rab. Fung. Eur. nos. 1385—6.
Cooke, Grevillea, vii. 137. Plowr. Ured. p. 129. Sace. Syll. vii.
538, Sydow, Monogr. ii. 230.
4icidiospores. Aicidia on the cotyledons chiefly, scattered
or in smal] clusters, at first hemispherical, then cup-shaped,
with erect, torn, white margin; spores finely verruculose,
orange-yellow, 17—35 p.

Uredospores. Sori scattered or aggregated, minute, rounded,
long covered by the epidermis, pulverulent,
cinnamon ; spores ovate to pyriform, very finely
echinulate, yellow-brown, 24—35 x 18—25 w;_
epispore about 144 thick.

Teleutospores. Sori similar, but larger and
rather compact, dark-brown; spores subglobose
to obovate, rounded above and often thickened
(up to 4), and surmounted by a thin, broad,
dark cap, rounded below, smooth, brown, 25—
35 x 18—28 4; pedicels hyaline, thick, per-

Fig. 66. U. Sali- sistent, as much as 80 w long.
corniae. Teleu-

tospores on S. On leaves and stems of Salicornia europaea
Gemnarees (herbacea). | October — November. Rare.
(Fig. 66.)

The teleuto-sori are chiefly on the stems, as much as 3 mm. long, and
very pulvinate.

DisTRIBUTION: France, Germany.

27. Uromyces Rumicis Wint.

Uredo Rumicis Schum. Pl. Sail. ii. 231.

Trichobasis Rumicum DC. ; Cooke, Micr. Fung. p. 225 p.p.

Uromyces apiculosa Lév.; Cooke, Handb. p. 518; Micr. Fung,
p. 212 p.p. (excl. figs.).

ON POLYGONACE 115

0. Rumicis Wint. Krypt. Fl. i. 145. Plowr. Ured. p. 135 p.p. Sace.
Syll. vii. 544, Sydow, Monogr. ii. 238. Fischer, Ured. Schweiz,
p. 9, f. 8.

Uredospores. Sori amphigenous, on coloured spots, round,
minute, scattered, soon naked, —
pulverulent, cinnamon; spores

subglobose to ellipsoid, sparsely
echinulate, pale-brown, 20—28 x
18—24 yw, with two (more often

three) germ-pores. eo Fig. 67. U. Rumicis. Teleuto-
Teleutospores. Sori similar, spores and uredospore, on R.

but darker; spores subglobose to oe
pyriform, with a hemispherical hyaline papilla, often narrowed
below, smooth or nearly so, brown, 2435 x 18—24 4; epispore
rather thick ; pedicels thin, hyaline, deciduous.

On Rumex conglomeratus, R. crispus, R. Hydrolapathum,
R. nemorosus, R. obtusifolius, and perhaps others. May—
September. Common. (Fig. 67.)

The spots on the leaves are small, round, and of various colours ; often
the chlorenchyma in the immediate neighbourhood retains its green colour
long after the rest of the leaf has become faded and yellow.

It will be noticed that the spores of U. Rumicis are exactly like those
of U. Ficariae, and for this reason Tranzschel was led to suspect some
connection between the two, such as he demonstrated to exist between
P. fusca and P. Pruni-spinosae, whose teleutospores are equally alike. In
1905 he reported that he had produced an ecidium on Ranunculus Ficaria
from the spores of U. Rumicis; still later, he repeated this statement
(1909), and added that he had infected Rumex obtusifolius with xcidio-
spores from &. Ficaria. Other experimenters (Bubék, Krieg) have been
unable to repeat the former of these infections ; they could only produce
the ecidium on &. Ficaria with the spores of Uromyces Poae. It has
been suggested that there are two ecidia on &. Picaria, one belonging to
U, Poae and the other to U. Rumicis; I have tried to infect 2. obtusefolius
with ecidiospores from &. Ficaria, brought from a place where the ecidium
on it and the Uromyces on &. obtusifolius were both very abundant, but
the attempt, failed, Krieg (Centralbl. f. Bakt. 1906) obtained uredo-
spores on &, Acetosa with ecidiospores from &. Ficaria, but the same
material infected species of Poa (especially P. trivialis), and the possibility

8—2

116 UROMYCES

of contamination by foreign spores was not entirely excluded in his
experiments. Judgment on this point must be suspended.

DistRIBUTION: Europe, Algeria, Asia Minor, Africa, Cali-
fornia, Chili.

28. Uromyces Acetose Schrot.

Uredo bifrons DC. ; Cooke, Handb. p. 528 ; Micr. Fung. p. 217, pl. vii.
figs. 137—9.

Uromyces Rumicis Wint.; Plowr. Ured. p. 135 p.p. Fischer, Ured.
Schweiz, p. 9 (not f. 8) p.p.

U. Acetosae Schrét. in Rab. Fung. Europ. no. 2080 (1876). Sace. Syll.
vii. 587. Sydow, Monogr. ii. 241.

[Spernogones. Honey-coloured, clustered.
Aicidiospores. AKcidia amphigenous or on the petioles, in
dense clusters (up to 1 cm. broad), cup-

ff shaped, whitish-yellow, with a cut and
revolute margin; spores nearly smooth
or very minutely punctate, clear-yellowish,

Fig. 68. U. Acetosae. 18—21 x 12—18 p.]
Teleutospores, on Uredospores. Sori amphigenous, often

ceca seated on red or purple spots, scattered or
circinate, minute, pulverulent, cinnamon; spores subglobose to
ellipsoid, finely and densely verruculose, yellowish or pale
brownish-yellow, 18—25 x 17—22 w; epispore about 24, thick,
with three germ-pores.

Teleutospores. Sori similar, but dark-brown; spores sub-
globose to ellipsoid, not thickened above, or rarely with a paler
and very minute papilla, rounded below, beset with very minute
warts arranged in lines, brown, 21—26 x 20—24 py; Bpinne
rather thick ; pedicels thin, hyaline, deciduous.

On leaves and petioles of Rumex Acetosa, R. Acetosella.
May—September. Not uncommon. (Fig. 68.)

Allied to U. Rumieis, but U. Acetosae has shorter spores (of both kinds)
and the hyaline papilla of the teleutospores is almost always wanting.
The «cidium has not been found in Britain, but the other stages are
rather common : the uredo- and teleutospores are unusually alike, but can
be distinguished by the germ-pores and the fewer warts of the latter.

ON POLYGONACEE 117

On the same host-plants is a Puccinia, which (in the absence of the
teleutospores) can be distinguished only by the fact that the uredospores
have two (rarely three) germ-pores and are adorned with few and distant
spines. There is little doubt that many of the specimens recorded as
U. Acetosae are really the uredospores of Puccinia Acetosae.

DISTRIBUTION : Germany, France, Sweden, Norway, Finland.

29. Uromyces Polygoni Fckl.

Aicidium aviculariae Kze.; Cooke, Handb. p. 545; Micr. Fung. p. 199.

Puceinia vaginalium Link; Cooke, Handb. p. 495; Micr. Fung.
p. 204.

Trichobasis Polygonorum Berk.; Cooke, Micr. Fung. p. 226 p.p.

Uromyces Polygont Eckl. Symb. Myce. p. 64. Cooke, Handb. p. 519 ;
Micr. Fung. p. 213, Plowr. Ured. p. 123. Sacc. Syll. vil. 533.
Sydow, Monogr. ii. 236. Fischer, Ured. Schweiz, p. 61, f. 46.
McAlpine, Rusts of Australia, p. 99, f. 150—1.

Spermogones. Honey-coloured, conical, only a few together.
Aicidiospores. Mostly hypophyllous, on yellow or violet
spots, irregularly aggregated or in circular groups, cup-shaped,
whitish, with a cleft and revolute margin; spores verruculose,
yellowish, 15—21 x 14—18 yp.
Uredospores. Sori amphigenous or on the stems, scattered
or in small clusters, small, round, soon
naked, pulverulent,.cinnamon ; spores glo-
bose to ellipsoid, densely and minutely
verruculose, pale-brown, 18—26 x 17—24u;
epispore 1424 » thick, with three or four
germ-pores.
Teleutospores. Sori like the uredo-
sori, but larger and more confluent upon fig 69, Uv. Polygoni.
the stems, compact, dark-brown; spores Teleutospores, on P.
aviculare.
globose or obovate, rounded above and
thickened (up to 6), smooth, chestnut-brown, 22—38 x 14—
22 w; pedicels coloured, persistent, thick, as much as 90 wu long.

On Polygonum aviculare. Aicidia, rare, May—June, Man-
chester (T. Brittain), 1875; uredo- and teleutospores, very
common, July—November. (Fig. 69.)

118 UROMYCES

The connection of the ecidium with the later stages seems not yet to
have been experimentally demonstrated : but McAlpine found the ecidia
on young plants of P. aviculare along with the other spore-forms. In
Europe, ete. it is recorded on many other species of Polygonum.

This species is said to occur also on Rumesx Acetosella on the continent,
and should be looked for here on that host. If so, there would be three
species on R, Acetosella to be considered : U. Polygoni which has verrucu-
lose uredospores and smooth teleutospores ; U. Acetosae which is distin-
guished by having teleutospores beset with a few minute warts arranged in
lines ; while Puceinda Acetosae has aculeolate uredospores and teleutospores
with many delicate warts.

DISTRIBUTION: World-wide.

30. Uromyces Lilii Fckl.

Caeoma Lili Link, Sp. PL, ii. 8.

Uromyces Lilti Fckl, Symb. Myc. Nachtr. iii. 16. Sydow, Monogr. ii,
277. Fischer, Ured. Schweiz, p. 6, f. 5. Grove, Journ. Bot. 1911,
p. 368.

U. Brythronii DC. ; Sacc. Syll. vii. 564 p.p. Wild Fauna and Flora
of Kew, p. 163 (non U. Erythronti Pass. %).

Nigredo Litit Arthur, N. Amer. Fl. vii. 242.

Spermogones. Dispersed among the ecidia, brownish-
yellow.

Aicidiospores. Aicidia amphigenous, usually hypophyllous,
or even on the petioles and stems, seated on lanceolate
yellowish spots, in rounded or elongated groups of very different
sizes, and often extending widely, more or less crowded, for
a long time hemispherical and closed, at length opening by a
central pore, but scarcely cup-shaped, margin not revolute,
about 4mm. diam. yellow; spores densely and minutely
verruculose, yellowish, 22-35 x 18-26 w; epispore 3 thick
or less. ;

Teleutospores. Sori amphigenous, usually hypophyllous, on
large yellowish spots, scattered or collected in groups, rounded
or oblong, elongated on the petioles, at first covered by the
epidermis, which at length splits, large, pulverulent, dark-
brown; spores globose to oblong or pyriform, rounded above,
with a prominent hyaline papilla, covered when mature with
short ridges which are often disposed in lines more or less

ON LILIACEE 119

interrupted and anastomosing, brown, 283—44 x 22—30 p; epl-
spore 2—3} w thick; pedicels hyaline, slender, deciduous.

Fig. 70. U. Lilii. a, leaf of Lilium candidum, with ecidia; b, another, with
teleuto-sori, nat. size; c, cells of peridium, in section and inner face-view ;
d, teleutospores ; ¢, teleutospore seen from above ; f, ecidiospore, all x 600,

On Liliwn candidum. Kew Gardens; also at Birmingham,
1911-3 (C. W. Lowe). Aicidia in April, May; mature teleuto-
spores from June. (Fig. 70.)

The part of the leaf occupied by the ecidia is somewhat thickened,
and the ecidia scarcely protrude above the epidermis. The peridia are
slow in opening. The streaked teleutospores are very distinctive. The
lilies on which the parasite appeared at Birmingham had been in the
garden for some years, but it was not noticed till 1911. Fischer records
it on Fritillaria Meleagris, and Sydow and Arthur on other species of the
two genera. The true U. Hrythronii differs from this species in possessing
a truly cup-shaped ecidium with a distinctly revolute margin. But the
teleutospores of the Birmingham specimens were more like those figured
by Fischer under U. Erythronii than those which he figures on Fritillaria
Meleagris (f. 5), though devoid of the “ Queranastomosen.” Possibly the
species on Lilium is quite distinct from that on Fritillaria. The longitu-
dinal strize are so plainly marked as to be visible under a comparatively
low power. As Fischer remarks, the cells of the lower part of the peridium
are much thinner-walled than those in the upper part.

DISTRIBUTION : Central Europe, North America.

120 UROMYCES

31. Uromyces Gagee Beck.

Uromyces Gageae Beck, Verh, k.-k. zool. hak Gesell. Wien, xxx. 26.
Sace. Syll. vii. 568. Sydow, Monogr. ii. 273. Fischer, Ured.
Schweiz, p. 4, f. 3.

U. Ornithogali Plowr. Ured. p. 142.

Teleutospores. Sori amphigenous, scattered, roundish or
elliptical, 1—3 mm. long, covered by the
lead-coloured epidermis which at length
splits longitudinally, then naked, pul-
verulent, dark-brown; spores subglobose
to obovoid, not or scarcely thickened
above, but usually with a hyaline apiculus,

Fig. 71. U. Gageae. smooth, brown, 26—40 x 18—28 w; epi-

Teleutospores. i i o
spore 2 » thick; pedicels hyaline, shorter

than the spores.

On leaves of Gagea lutea. Rare. April, May. (Fig. 71.)

The teleutospores mature in spring, according to Fischer. Plowright
says that the mycelium causes variously shaped pale spots on the affected
leaves ; but I find no spots and in Sydow it is said that there are none.

DisTRIBUTION: Western and Central Europe.

32. Uromyces Scillarum Wint.

Uredo Scillarum Grev. in Smith, Engl. Fl. v. 376.

Uromyces concentricus Lév. ; Cooke, Handb. p. 519; Grevillea, vii. 138 ;
Micr. Fung. p. 213.

U. Scillarum Winter, Pilze Deutsch]. p. 142; Plowr. Ured. p. 141.
Sacc. Syll. vii. 567. Sydow, Monogr. ii. 278. Fischer, Ured.
Schweiz, p. 2, f. 1.

Teleutospores. Sori amphigenous, usually seated on pallid

or yellowish spots, small, round or

oblong, up to +mm. diam., collected
into round or oblong clusters, often
concentrically arranged, sometimes
confluent, long covered by the epi-

dermis which at length splits and

a ee ies ke on surrounds them, pulverulent, dark-

teleutospore, on S. campanu- brown; spores subglobose to oblong,
sao usually rounded and not thickened
above, smooth, occasionally marked with a few very faint lines

a

ON LILIACEA 121

running from apex to base, evenly coloured, brown, 18—82 x
14—22 w; epispore uniformly thin, about 14 thick; pedicels
hyaline, often deciduous, as long as or longer than the spore.

On leaves of Endymion non-scriptum (Scilla nutans), and

also of Scilla bifolia, S. campanulata. Common. April—June.
(Fig. 72.)

The yellow spots and the concentric arrangement of the sori are often
very marked. The mycelium is purely local. A few finely echinulate
uredospores, 27x 20 p, were found by Juel in the young sori on Seilla
obtusifolia (Bull. Soc. Myce. Fr. xvii, 259).

DISTRIBUTION: Central and Southern Europe and Morocco.

33. Uromyces ambiguus Lév.

Oredo ambigua DC. Flor. fr. vi. 64.

U. Alliorum Cooke, Handb. p. 528; Micr. Fung. p. 217 p.p.

Puccinia nuxta Fckl., forma simplicior Korn.

Uromyces Alliorum DC. ; Cooke, Handb. p. 518 ; Micr. Fung. p. 212.
Plowr. Ured. p. 137 p.p.

U. ambiguus Lévy. Ann. Sci. Nat. 3, viii. 375. Sacc. Syll. vii. 543.
Sydow, Monogr. ii. 262. Grove, Journ. Bot. 1911, p. 367.

Uredospores, Sori amphigenous, without spots, scattered,
roundish or oblong, long
covered by the epidermis,
yellowish; spores globose to
obovate, delicately verruculose,
pale-yellowish, 20—28 x 17— Roy
22 4; epispore 3—34 w thick,
with six or seven germ-pores. Fig. 73. U. ambiguus. Teleutospores
Teleutospores. Sori am- and uredospore on Allium Scorodo-
phigenousand onthestems,on = ?”™*"""
the leaves scattered, small and roundish, on the stems confluent
and larger, up to 15 mm. long, always covered by the blue-grey
epidermis; spores subglobose to pyriform, rounded above,
without a papilla and scarcely thickened, rounded or narrowed
below, smooth, brown, 20—35 x 17—24 w; pedicels thin, hyaline,
fragile, as much as 30 pw long.

122 UROMYCES

On Allium Schoenoprasum, A. Scorodoprasum, and (ac-
cording to Sydow) A. sphaerocephalum. Not common. (Fig. 73.)

It cannot be considered as certain that this is a species distinct from
Puccinia Porrt Wint., but as it presents slight differences, it is better, in
the total absence of culture experiments, to keep it separate for the time.
The difficulty lies in the fact that the teleutospores of this Uromyces agree
perfectly with the mesospores of the Puccznia (except perhaps in the greater
variability of the latter) ; but the teleuto-sori of the Uromyces are generally
larger and more persistently covered by the epidermis, and one searches
in vain in them for two-celled spores, such as are found freely in the sori
of the Puccinia. Fischer says that the uredospores of P. Porri have only
three germ-pores. According to Sydow, the Uromyces occurs only on the
three species of Allium mentioned above and A. rotundum, while the
Puccinia is found on them as well as on many other species of the genus.
Nevertheless the two forms are closely allied and from the evolutionary
point of view the Uromyces must be regarded as a specialised state or
mutation of the Puccinia. Neither of them has an ecidial stage.

DisTRIBUTION: Central and North-Western Europe.

34. Uromyces Colchici Massee.

U. Colchici Mass. Grevillea, xxi. 6, pl. 182, f. 16—18; Diseases of
Cult. Plants, p. 292, f. 85; Text-book of Plant Diseases, p. 227,
f. 56. Sydow, Monogr. ii. 268.

Teleutospores. Sori amphigenous, scattered, rather large,
elliptical, sometimes circinating, up to 2mm. long,
covered for some time by the epidermis which at
length splits, then sub-pulverulent, brown ; spores
subglobose to ovate, rounded above, with a broad
flat hyaline papilla, smooth, pale-brown, 28—40 x

Fig. 74. U. 20—28y; epispore 3—34 u thick; pedicels hya-

Colchici. — line, rather long, but very deciduous.
Teleutospore, . ’
from the ori- On leaves of Colchicum speciosum. Kew
ea specl- Gardens; unknown in the world elsewhere.
(Fig. 74.)

The teleutospores remain on the dead leaves and germinate in the
following spring, so that if Colchicum is again planted in the same ground
or allowed to remain there, it is liable to contract the disease year after
year. As in all similar cases, the best preventive against future attacks

ON JUNCACEA 123

is to remove carefully and burn all diseased leaves before they mature
their spores.

The fungus is stated to have attacked the foliage of the host for three
successive seasons, completely destroying it, and although for the first two
seasons it did not attack other species of Colchicum growing near, during
the third season it spread to C. autumnale and C. bavaricum.

35. Uromyces Junci Tul.

icidium zonale Duby, Bot. Gall. ii. 906. Cooke, Grevillea, xiv. 39.

Uromyces Junci Tul. Ann. Sci. Nat. ser. 4, ii. 146. Cooke, Grevillea,
vii. 1389; Micr. Fung. p. 213. Plowr. Ured. p. 132 ; Grevillea, xi.
52. Sacc. Syll. vii. 541. Sydow, Monogr. ii. 287. Fischer, Ured.
Schweiz, p. 57, f. 43,

Nigredo Junct Arthur, N. Amer. FI. vii. 238.

Spermogones. Usually epiphyllous.

Aicidiospores. Aicidia hypophyllous, seated on spots which
are zoned with yellow and purple, in
dense circinate clusters 2—5 mm.
wide, cup-shaped, yellowish-white,
with a torn revolute margin; spores
densely and minutely verruculose,
transparent-yellowish, 17—21 p.

Uredospores. Sori scattered,
roundish or oblong, up to 1 mm.
long, surrounded by the cleft epi- _
dermis, pulverulent, brown; spores Poe oo ieee
globose to ellipsoid, faintly echinu-
late, yellowish-brown, 20—28 x 16—22 yw, with two equatorial
germ-pores.

Teleutospores. Sori amphigenous or on the culms, scattered
or occasionally aggregated, similar to the uredo-sori, but darker ;
spores oblong-ovate to clavate, rounded or conical above and
much thickened (up to 14), attenuated below, smooth, dark-
brown, 24—42 x 12-18 w; pedicels thick, persistent, brownish,
as much as 60 yu long.

Aieidia on Pulicaria dysenterica, May—July; uredo- and
teleutospores on Juncus obtusiflorus, from July onwards, lasting
through the winter on the dead culms. Not common. (Fig. 75.)

124 UROMYCES

The connection of the two forms, first proved by Fuckel and Plowright,
has been confirmed by Fischer. On the continent and elsewhere, there
are forms of Uromyces on other species of Juncus, some of which (and
possibly most of them) have their ecidia on other hosts than P. dysenterica,

DiIsTRIBUTION: Central and Western Europe, Algeria, North
and South America.

36. Uromyces Scirpi Burr.

Aeidium Glaucis D. et M.; Cooke, Grevillea, xv. 29, Plowr. Ured.
p. 268 ; Gard. Chron, ser. 3, vii. 682, 746.

Oredo Scirpi Cast. Catal. Pl. Marseille, p. 214,

Uromyces Scirpi Burr. Parasit. Fung. Ilinois, p. 168. Sacc. Syll. vii.
558. Sydow, Monogr. ii. 302.

U. lineolatus Desm.; Plowr. Grevillea, xxi. 111.

U. maritimae Plowr. Gard. Chron. ser. 3, vii. 682, 746; Jour. Roy.
Hort. Soc. (1890), p. cix. Klebahn, Wirtswechs. Rostpilze, p. 328.

Nigredo Scirpi Arthur, N. Amer. Fl. vii. 233.

Spermogones. Usually epiphyllous.

Aicidiospores. Aicidia hypophyllous or on the petioles, in
rather small clusters, cup-shaped, with an incised revolute
margin; spores densely and minutely vérruculose, transparent-
yellowish, 16—24 x 1420 wu.

Uredospores. Sori hypophyllous, scattered or arranged in
lines, rounded or oblong, up to 1 mm. long,
surrounded by the cleft epidermis, pulverulent,
cinnamon ;. spores globose to ovate, distantly
and minutely echinulate, yellowish-brown,
22—35 x 16—25 p; epispore 14—2 pw thick,
with three equatorial germ-pores.

Teleutospores. Sori amphigenous, on in-

definite discoloured spots, scattered or confluent

Fig.76. U.Scirpi. 1n lines, long.covered by the epidermis, brownish-

ne black; spores oblong to clavate, tapering usually

Illinois, U.S.A. and thickened (up to 12) above, attenuated

below, smooth, pale-brown, 26—45 x 15—24 w;

pedicels brownish, persistent, as long as or longer than the
spore.

ON GRAMINE 125

fEcidia on leaves and petioles of Glauz maritima, May ;
uredo- and teleutospores on Scirpus maritimus, June to August.
Banks of the Humber, Hull. (Fig. 76.)

The researches by which Plowright proved the connection of these two
forms are given in Grevillea, xxi. 111, and in Journ. R. Hort. Soc. xii.
p. cx.; other observers have found a similar Uromyces on Scirpus mart-
timus and therefrom have produced ecidia on other host plants such as
Pastinaca sativa (Rostrup), Berula angustifolia, Daucus Carota (Bubak),
Ginanthe aquatica (Klebahn), Hippuris vulgaris and Sium latifolium
@ietel), etc. In North America, a morphologically indistinguishable
Uromyces on Scirpus fluviatilis, etc. has produced an xcidium on Cicuta
maculata (Arthur), and similar ecidia on allied Umbelliferze are suspected
to belong to the same life-cycle. It is evident that U. Seirpi, like Puceinia
Isiacae, is in its ecidial stage a plurivorous species, though possibly some
of these forms may be separated in the future as “biological” races. In
any case, they are not so sharply distinguished as in other instances, but
Klebahn isolates our British species as U. maritimae Plowr. See the full
account in Sydow, Monogr. ii. pp. 304—7.

DisTRIBUTION: Europe and North America.

37. Uromyces Dactylidis Otth.

Aicidium Ranunculi-acris Pers. Obs. Mye. ii. 22.

i. Ranunculacearum DC. FI. fr. vi. 97 p.p. Cooke, Handb. p. 539;
Micr. Fung. p. 196 p.p.

Uromyces Dactylidis Otth, Mittheil. Nat. Gesell. Bern, 1861, p. 85.
Plowr. Ured. p. 130. Sace. Syll. vii. 540 p.p. Sydow, Monogr. ii.
309. Fischer, Ured. Schweiz, p. 71, f. 54.

U. graminum Cooke, Handb. p. 520; Micr. Fung. p. 214,

Spermogones. Epiphyllous, honey-coloured, but also a few
scattered among the zcidia on the lower surface.

Aicidiospores. Aicidia hypophyllous or on the petioles,
seated on yellow spots, in roundish or, on the petioles, elongated
clusters, cup-shaped, yellow, with slightly torn, recurved margin ;
spores delicately verruculose, pale-yellowish, 17—25 p.

Uredospores. Sori amphigenous, scattered or in rows, small,
elliptic or oblong, long covered by the epidermis, pulverulent,
yellow-brown; spores globose to ovate, delicately echinulate,
yellow or yellow-brown, 21—32 x 18—25y; epispore 14—2 p
thick, with 7—9 germ-pores ; paraphyses generally wanting.

126 UROMYCES

Teleutospores, Sori generally hypophyllous, similar to the
uredo-sori but more often confluent,
always covered by the epidermis,

! compact, shining, black; spores ovate-

\ oblong, occasionally ellipsoid or pyri-

form, rounded above, rarely truncate,

often slightly thickened (up to 4,),

Fig. 77. U.Dactylidis. Te. ‘Smooth, yellow-brown, darker only

leutospores and the accom- glong the summit, 18—30 x 14—

Eee eee 20; epispore 14m thick; pedicels

brownish, persistent, nearly as long as the spore; paraphyses

numerous, brown, agglutinated, dividing the sori into compart-
ments.

AKcidia on Ranunculus acris, R. bulbosus, R. repens, March—
May; uredo- and teleutospores on Dactylis glomerata, from
July onwards, often covering the leaves, less often the sheaths

and culms, and persisting through the winter, especially on the
latter. (Fig. 77.)

This species and the following (U. Poae) are very closely allied, and
should possibly be united. Perhaps more experimental cultures have been
carried out with these two species than with most other Uredines; but the
result is only a wild confusion of contradictory statements, from which
one can infer, either that an immense number of intricately connected, but
morphologically indistinguishable forms, inhabit the species of Ranun-
cwlus and of Poa and Dactylis—or, preferably, that the factors which
govern the success of an attempted infection are so numerous and so little
known, that failure does not afford any ground for arriving at a definite
conclusion. Those who wish to learn further should consult the long
account of these results given in Sydow, Monographia, ii. pp. 312—16.

In the British specimens of U. Dactylidis which I have examined, the
paraphyses in the teleuto-sori, though often overlooked, are a conspicuous
feature. But upon the question of paraphyses in the uredo-sori no agree-
ment has been arrived at; the various authorities flatly contradict one
another. Hither, therefore, the paraphyses occur differently in different
countries, as Plowright suggests, or more than one species is included
under this title, or their presence or absence is a matter of no importance.
Against the latter suggestion, however, we must set the fact that in other
cases, e.g. in Puccinia Sonchi, the paraphyses form a constant and dis-
tinctive character. ;

An ecidium occurring on Ranunculus acris belongs to Puceinia
perplevans Plowr., but cannot be distinguished from the present one,

ON GRAMINEE 127

except by cultures ; see also under U. Poae. The teleuto-sori of the present
species are more numerous and much more conspicuous than those of U.
Poae, They resemble the spots of Phyllachora graminis on the same host.

DIstRIBUTION : Europe only, not yet found in America.

38. Uromyces Poe Raben.

Aeidium crassum var. 8 Ficariae Pers. ; Sow. pl. 397, f. 4.

42. Ranunculacearum DC.; Cooke, Handb. p. 539; Micr. Fung.
p. 196, pl. ii, figs. 12—14, p.p.

Uromyces Poae Rab. Unio itin. 1866, no. 38. Plowr. Ured. p. 131.
Sydow, Monogr. ii. 310. Fischer, Ured. Schweiz, p. 72, f. 55.

Spermogones. Epiphyllous, honey-coloured, a few also on
the lower surface among the ecidia.

Afcidiospores, AXcidia hypophyllous or on the petioles, in
roundish clusters on yellow spots, which on the petioles are
often elongated, cup-shaped, yellow, with torn, recurved margin ;
spores delicately verruculose, clear-yellow, 17—25 x 12—20 p.

Uredospores. Sori amphigenous, scattered or in rows, small,
elliptic or oblong, at first covered

by the epidermis, pulverulent,
yellowish-brown; spores globose
to ovate, faintly echinulate, yellow,
14—25 x 14—20 yw; epispore 14—
2 thick, with 4—9 germ-pores ;
a few paraphyses occasionally

intermixed. ; Fig. 78. U. Poae. Teleutospores
Teleutospores. Sori generally | and the accompanying para-

hypophyllous, similar to the uredo- PR ean

sori, but always covered by the epidermis, compact and black ;
spores oblong-ellipsoid to pyriform, rounded or truncate above,
not much thickened (up to 4 or 5 y), smooth, yellowish-brown,
apex alone darker, 17—28 x 14—20,y; epispore lip thick;
pedicels brownish, as long as or shorter than the spore; para-
physes numerous, brown, agglutinated, as in U. Dactylidis.

Acidia on Ranunculus auricomus, R. bulbosus, R. Ficaria,
R. repens, March—May; uredo- and teleutospores on Poa
annua, P. nemoralis, P. pratensis, P. trivialis, May—September.
(Fig. 78; see also Fig. 211.)

128

PUCCINIA

Concerning the difference between this species and U. Dactylidis

i/

Fig. 79. Acidia of U.
Poae (b) and teleuto-
sori of U. Ficariae
(a), on a leaf of R.
Ficaria, nat. size.

to this latter species.

(apart from the habitat) little that is definite can
be said. Some authors unite them, but I find the
teleutospores of U. Poae to be usually more oblong
and often provided with shorter pedicels and the
sori to be less conspicuous. The leaves of Poa are
smaller than those of Dactylis, and the teleuto-sori
do not cover them in such enormous numbers and
never extend to the culms.

Juel divides this species into 9 or 10 biological
races, but there is the usual conflict between
different experimenters as to their limits. Some
would even deny, what has been proved several
times, that an wcidium on kk. LYcarta belongs
here: see U. Rumic’s (p. 115). This ecidium
and U. Ficariwe may occasionally be found on the
sane leaf of R. Miearia. (Fig. 79.)

It must not be forgotten th t an eei 7ium which
occurs on £&. bulbosus and Lk. repens be
Puccinia Magnusiana : this is morphologically in-
distinguishable but is said to be later in its
appearance. The eecidia which can be seen on

pngs to

RL. repens in July and August possibly all belong
The teleutospores of (. Pode can most easily be

found by looking on the lower leaves of Poa in June or July along a damp
road-side where #. repens is abundant.

DISTRIBUTION: Europe, Asia Minor, Nova 8 zotia

PUCCINIA Pers.

Autcecious or hetercecious.
Spermogones asin Uromyces. Meidia witha peridium, which
is occasionally evanescent ; spores as in Uromyces. Uredospores

as in Uromyces. Teleutospores two-celled, rarely one- or several-
celled, the upper cell usually with an apical pore, less often the
pore is displaced to the side; the lower cells with a pore just
beneath the septum or rarely at a lower level. Basidiospores as

in Uronyces.

ON COMPOSIT 129

This genus occupies an intermediate position between
Uromyces and the less advanced genera, Phragmidium, etc., as
is shown by the fact that many species tend to form one-celled
teleutospores (mesospores) indistinguishable from those of
Uromyces, while others have spores with three or more cells,
arranged as in Phragmidium, Triphragmium, Sphaerophrag-
mium, ete.

The number of species is enormous, more than 1300 are
already known. The genus must therefore be subdivided, but
no quite satisfactory classification has as yet been discovered.
Schréter’s and Fischer’s separate nearly allied species, while
Arthur’s is a pathless chaos. As a temporary measure, instead
of adding a new imperfect scheme to those already existing,
the species are here arranged on the plan adopted in Sydows’
Monographia, in the order of the families and genera on which
they are parasitic; this has the advantage that it does, to a
great extent, bring nearly allied species
close together, while it is at the same time
very convenient for ccnsultation. In each
family the genera are arranged in the order
familiar to British botanists, but the fami-
lies themselves are in the order usually:
adopted on the continent, because that
will before long be accepted here also.

1. Puccinia Tripolii Wallr.

Puceinia Tripoli Wallr. Fl. Crypt. Germ. ii.
223. Cooke, Micr. Fung. p. 207; Gre-
villea, iii. 180.

P. Asteris Duby, Bot. Gall. ii. 888. Plowr.
Grevillea, ii. 48; Ured. p. 215. Sace.
Syll. vii. 687’ p.p. Sydow, Monogr. i.
15 p.p.

Teleutospores. Sori amphigenous, con-
fluent into rather large, pulvinate masses, hard, compact, very
dark-brown ; spores oblong-clavate or clavate, rounded above or
somewhat narrowed and much thickened (as much as 8p),
slightly constricted, generally attenuated downwards, smooth,

Fig. 80. P. Tripolii.
Teleutospores (one
abnormal).

G. U. 9

“130 PUCCINIA

pale clear-brown, 45—60 x 20—25 4; pedicels brownish, persis-
tent, thick, about as long as the spore or longer; no mesospores
were seen, but some irregular spores.

On Aster Tripolium. New Pitsligo, 1870 (Herb. Berk.);
Wolferton Beach, King’s Lynn, July—November, 1873 (Plow-
right). (Fig. 80.)

The greyish tinge mentioned by Plowright seems to be due to germ- ‘
tubes issuing from the spores of this Leptopuccinia. This species is
decidedly different from the American forms with which it is united by
Sydow ; there are no spots, the colour of the spore is paler, the apex not
so much thickened, and the sori are not confined to the lower leaf-surface.
Many spores were observed bifid at the summit, but not owing to germi-
nation, which had not taken place in them.

DistrizuTion: Northern and Central Europe, Siberia.

2. Puccinia Virgauree Lib.

Xyloma Virgaureae DC. Flor, fr. vi. 158.

Puccinia Virgaureae Lib. Crypt. Arduen. iv. No. 393. Cooke, Handb.
p. 500; Micr. Fung. p. 206. Plowr. Ured. p. 203. Sace. Syll.
vii. 679, Sydow, Monogr. i. 151. Fischer, Ured. Schweiz, p. 363,
f. 264.

Teleutospores. Sori hypophyllous, minute, crowded in
stellate or dendritic clusters,on round,
yellowish, purple-centred spots, com-

f= pact, shining, black ; spores oblong,

clavate or fusoid, above rounded,

attenuated or truncate, very much

thickened (as much as 12) and

darker or with a paler hood-like cap,

hardly constricted, tapering below,

| smooth, yellow-brown, paler down-

wards, 30—56 x 12—20 uw; pedicels

Fig. 81, P. Virgaureae, somewhat hyaline, half as long as the

Teleutospores. spore.

On Solidago Virgaurea. August and September. Un-
common; Surrey, etc. (Fig. 81.)

The sori are arranged in a radiating fashion; they are surrounded
each by a thick fence of dark-brown paraphyses, remain long covered by

ON COMPOSITAE 131

the epidermis and resemble rather a Dothidea or Asteroma than a Puccinia.
The spores bear a slight resemblance to those of P. Poarum, but are more
fusiform, and much more thickened at the apex. Mixed with them are
sometimes a number of mesospores.

DISTRIBUTION: Europe, except the southern parts.

3. Puccinia Millefolii Fckl.

Puceinia Millefolii Fckl. Symb. Myc. p.55. Cooke, Micr. Fung. p. 207.
Plowr. Ured. p. 215. Sydow, Monogr. i, 2. Fischer, Ured.
Schweiz, p. 296, f. 216.

Teleutospores. Sori amphigenous, on indistinct spots,
minute, roundish or irregular, gene-
rally scattered, compact, dark-
brown ; spores oblong or clavate,
rounded or gently attenuated at the
apex and thickened (4—9,), con-
stricted, more or less tapering below,
smooth, pale-brownish, 35—50 x
13—19 w; pedicels yellowish above,
thick, persistent, about 404 long;
a few mesospores are sometimes
intermixed,

On Achillea Millefolium. For-
den; St Leonards. August—October. (Fig. 82.)

a8)

Fig. 82. P. Millefolii. Teleuto-
spores.

Plowright demonstrated (/.c. p. 216) that this species is quite distinct
from P. Tripoli, with which it had been previously united, and his con-
clusion has been confirmed by Magnus and others. The species is rather
uncommon ; there is a closely allied one on A. Ptarmica (P. Ptarmicae
Karst.) which has not yet been seen in Britain. The spores of P. Mille-
Jfolii germinate as soon as mature.

DIsTRIBUTION : Central and Western Europe.

4, Puccinia Chrysanthemi Roze.

Uredo Chrysanthemi Roze ; Plowright in Trans. Brit. Myc. Soc. i. 98.

Puceinia Chrysanthemi Roze, Bull. Soc. Myc. Fr. 1900, p. 92. Sace.
Syll. xvi. 296. Sydow, Monogr. i. 46, 854. McAlpine, Rusts of
Australia, p. 153, f. 251—5, and pl. E, f.21. Fischer, Ured.
Schweiz, p. 190, f. 150.

P. Chrysanthemi-chinensis Henn. in Hedwig. xl. 26 (1901).

9—2

132

PUCCINIA

Uredospores. Sori generally hypophyllous, on irregular

pallid-yellow or brownish spots, scattered or in
clusters, about 1—14 mm. diam. often circi-
nate, pulverulent, snuff-brown ; spores globose
to ellipsoid, delicately echinulate, brown, 24—
52 x 17—27 w, mostly with three germ-pores.
[Teleutospores. Mixed with the uredospores,

Fig.83. P. Chry- oblong or ellipsoid, rounded and_ slightly

santhemi. Ure-
dospore (Brit-
ish).

thickened above, usually rounded or some-

h). what tapering at base, scarcely constricted,

delicately verruculose,

chestnut-brown, 35-—57 x 20—25 yz;

pedicels thick, hyaline, persistent, 35—60 long; mesospores

Fig. 84. P. Chrysanthemi.
Teleutospores and uredo-
spores, one abnormal (after
Fischer).

subglobose or pyriform, — slightly
thickened at the summit, 32—387 x
20-—21 p.]

On leaves of Chrysanthemum in-
dicum and C. sinense (not on other
species of the genus, much less on
other genera of Composite), in
greenhouses, all the year round. The
leaves that are attacked soon flag and
die. (Figs. 88, 84.)

This species is said to be very common
in Japan. It was first observed in England
in 1895, and has been found in other Euro-
pean countries and in North America ; in
1904 it reached Australia and New Zealand.

In Japan it produces teleutospores in
separate sori, which are hypophyllous,
roundish, dark-brown and naked, but in
Europe the teleutospores have been rarely
seen, though mesospores occasionally occur.
Abnormal and 2-celled uredospores (as well
as 3- or 4-celled teleutospores) have been

described and figured by Roze, Jacky and Fischer ; but these I have not

seen in British specimens,

Since, under the conditions in which the plants are grown here, the
young shoots appear above ground before the old ones die away, it is
probable that the parasite maintains itself by the uredospores alone; the
alternative would be the possession of a perennial mycelium, which has not

ON COMPOSITA 133

been found (Gibson, 1904, p. 188). If that is so, the disease can easily
be kept in check by rigid cleanliness and by spraying at intervals with
very dilute Bordeaux mixture or, better still, potassium sulphide solution.
Remove and burn all attacked leaves as soon as seen, water carefully with-
out wetting the leaves, choose resistant varieties (e.g. “‘ October Sun” and
“ William Tricker” are said to be immune), and there will be little fear
of an epidemic of the disease.

DistRiBuTION: Europe, Japan, North America, Australia.

5. Puccinia Leucanthemi Pass.

Puccinia Leucanthemi Pass. in Hedw. 1874, p.47. Sacc. Syll. vii. 705,
Sydow, Monogr. 1. 116, f. 95.

P. Asteris var. Chrysanthemi-Leucanthemi Massal. in Bull. Soc. Bot.
Ital. 1900, p. 258. Sacc. Syll. xvi. 297. Trans. Brit. Myc. Soc.
iii, 224.

Teleutospores. Sori amphigenous, generally hypophyllous,
often also on the petioles, scattered
or often circinate on indistinct spots,
or confluent into compact cushions
2—5 mm. wide, reddish-brown; spores
oblong or subclavate, somewhat
rounded or more often narrowed at
the apex, much thickened above (up
to 14), constricted, tapering down-
wards, smooth, yellowish, 40—70 x
1424; pedicels hyaline, thick, \

about as long as the spore.
Fig. 85. P. Leucanthemi.

On Chrysanthemum Leucanthe- Teleutospores, from an ori-
ginal specimen issued by
mum. Very rare. Lamorna. Cove, Passerini.

Cornwall, September, 1906 (F. J.
Chittenden). (Fig. 85.)

Only recorded for Britain and Italy. The similarity of the spores to
the teleutospores of Puceinia Afcidii-Leucanthemi Fisch., which has its
eecidiospores on C. Leutanthemum and its teleutospores on Carex montana,
seems to indicate that this is one of those instances, like P. fusca and P.
Pruni-spinosae, which give us a glimpse into the mode of evolution of
the Uredinales.

134 PUCCINIA
6. Puccinia Absinthii DC.

Trichobasis Artemisiae Berk. ; Cooke, Micr. Fung. p. 223.

Puceinia Discoidearum Link ; Cooke, Micr. Fung. p. 206.

P. Tanaceti Plowr. Ured. p. 189 p.p. Sace. Syll. vii. 637 p.p.

P. Absinthis DC. Flor. fr. vi. 56. Sydow, Monogr. i. 11. Fischer,
Ured. Schweiz, p. 188, f. 148.

Uredospores. Sori generally hypophyllous, on yellowish-
brown or indeterminate spots, scattered or aggregated, not
confluent, minute, roundish, pulverulent, pale-brown; spores
globose to ovoid, echinulate, pale yellowish-brown, 20—35 x
15—26 pw, mostly with three subequatorial germ-pores.

Teleutospores. Sori amphigenous, but generally on the
lower surface or sometimes on the
stems, similar to the uredo-sori but
occasionally confluent, soon naked,
dark-brown or blackish; spores oblong
to oblong-clavate, rounded and thick-
ened (8—7 p) above, constricted,
slightly attenuated below, the upper
cell punctate or verruculose, the
lower frequently smooth, especially
at the base, brown, 38—62 x 20—
27 w; pedicels hyaline, thick, persis-
tent, as much as 80 w long.

On Artemisia <Absinthium, A.
Fig. 86. P. Absinthii, Teleu- maritima, A. vulgaris. July—Sep-
tospores, on A. Absinthium. + ber, Rather uncommon, Also
recorded in Switzerland on A. campestris. (Fig. 86.)

The germ-pores of both kinds of spores are covered with paler, swollen
caps. P. Tanaceti, which was formerly united with this species, possesses
on the average narrower and shorter teleutospores, though in each these
spores are said to be marked in a similar way, chiefly in the upper half.
But the markings of P. Absinthii are stronger and less likely to escape
observation, especially on the pore-caps of both cells.

DISTRIBUTION: Europe, Siberia, Japan, North America

ON COMPOSITA 135

7. Puccinia Tanaceti DC.

Puccinia .Tanacett DC. Flor. fr. ii. 222. Cooke, Micr. Fung. p. 207, .
Plowr. Ured. p. 189 p.p. Sacc. Syll. vii. 657. Sydow, Monogr. i.
161. Fischer, Ured. Schweiz, p. 185, f. 146.

P. Discoidearum Link ; Cooke, Handb. p. 499 p.p.

Uredospores. Sori amphigenous, scattered, minute, not
confluent, orbicular, pallid-brown ; spores subglobose or broadly
ellipsoid, echinulate, yellow-brown, 25—82 x 16—25 w, with
three germ-pores, each covered by a convex colourless cap.

Teleutospores. Sori mostly hypophyllous, similar, but more
compact and blackish, 4—%mm, diam.; spores ellipsoid or

Fig. 87. P. Tanaceti, Teleutospores.

oblong, more or less rounded at both ends, thickened above (up
to 7), delicately verruculose (?) especially in the upper part,
rich-brown, 32—44 x 16—24; pedicels hyaline, thick, per-
sistent, as much as 120 long.

On Tanacetum vulgare. Not common. August, September.

(Fig. 87.)

136 PUCCINIA

Formerly united with Puccinia Absinthii, from which it is distinguished
by its (on the average) narrower and shorter teleutospores, with longer
pedicels. I could not see in any case that the teleutospores were verru-
culose, as they are said to be.

DistTRIBuTION: North-western and Central Europe.

8. Puccinia expansa Link.
Puccinia expansa Link, Sp. Pl. vi. 2, p. 75. Sydow, Monogr, i. 146.
Fischer, Ured. Schweiz, p. 182, f. 141—2.
P. Senecionis Cooke, Micr. Fung. p. 207. Plowr. Ured. p. 209,
(non Lib.).

Teleutospores. Sori amphigenous, but generally hypophyllous,
on round yellowish or brownish
spots, densely crowded and con-
fluent in roundish clusters 5—8
mm. broad, blackish-brown; spores
ovate or broadly ellipsoid, rounded
at both ends, with a minute paler

oe ae eae re apical papilla, hardly constricted,
spores and mesospore, on S. smooth, brown, 30—40 x 19—30x,
TRUE but sometimes smaller; pedicels
hyaline, very short; an occasional mesospore is seen.

On Senecio aquaticus, 8. Jacobaea. Not common. July—
September. (Fig. 88.)

The sori of this species are chiefly on the underside of the leaf; they
are collected into roundish or elongated groups and open by a small round
pore, after the manner of some ecidia, so that, when the spores have
fallen out, the group looks like a honeycomb. P. glomerata is said to be
distinguished by its paler, narrower and longer spores with a smaller
papilla, and by having the sori on both leaf-surfaces. I think the two
species are the same so far as our specimens are concerned, but have
followed Sydows’ Monographia pro tem. Plowright has confused the
characters of the two, partly because he thought his species on 8. aguaticus
was P. Senecionis Lib., which is incorrect. The true P. Senecion’s Lib.
has not been found in Britain.

DISTRIBUTION: Central Europe, Holland, California.

9, Puccinia glomerata Grev.

Puccinia glomerata Grev. in Berk. Eng. Fl. y. 365. Cooke, Handb.
p. 500; Mier, Fung. p. 206. Plowr. Ured. p. 209. Sydow,
Monogr. i. 148, t. 8, f. 125.

ON COMPOSIT 187

Teleutospores. Sori amphigenous, on brown orbicular spots,
densely gregarious and confluent in roundish clusters, 3—6 mm.
broad, occupying the whole spot, often circinate, long covered
by the epidermis, at length rather pulverulent, deep-brown ;
spores ovate to oblong, with a very minute paler apical papilla,
hardly constricted, rounded or slightly attenuated below,
smooth, pale-brown, 30—45 x 16—24; pedicels hyaline, very
short.

On Senecio Jacobaea. August—November. Not common.

Closely allied to the preceding species, but said to be distinguished by
its paler, narrower and longer spores which are provided with a smaller *
apical papilla. Moreover, the sori are said to be less dark in colour (not
darker as Plowright states), and are distributed more uniformly on both
leaf-surfaces, Nevertheless, I suspect they are identical.

DISTRIBUTION: Belgium.

10. Puccinia Carline Jacky.

Puceinia Hieracti Mart. ; Plowr. Ured. p. 184 p.p.
P. Carlinae Jacky, Compos.-Puce. p. 59, f. 14. Sacc. Syll. xvi. 297.
Sydow, Monogr. i. 35. Fischer, Ured. Schweiz, p. 216, f. 167.

Spermogones. Epiphyllous and on the petioles, singly or in
groups.

Uredospores. Sori amphigenous, without evident spots,
scattered, not confluent, mi-
nute, punctiform, pulverulent,
brown; spores globose or
broadly ellipsoid, very deli-
cately echinulate, _pallid-
brown, 24—380 x 20—25 p;
epispore uniformly thick, with
three germ-pores.

Teleutospores. Sori simi- Fig. 89. P. Carlinae. Teleutospores.
lar, but blackish-brown; spores
ellipsoid to obovate-oblong, rounded at both ends or more often
attenuated downwards, not thickened above, hardly constricted,
delicately verruculose, brown, 25—385 x 16—20p; pedicels
hyaline, short.

138 PUCCINIA

On Carlinavulgaris. July—October. Not common. (Fig. 89.)

Distinguished from P. Cirsti, which it much resembles, by the larger
uredospores, with very delicate hardly perceptible spines, and by the
slightly smaller teleutospores which are more frequently narrowed below.
But no cultures seem to haye been made to prove their distinctness.

DisTRIBUTION : Central and Northern Europe.

11. Puccinia Bardane Corda.
Puceinia Mieracit Plowr. Ured. p. 184 p.p.
P. Bardanae Corda, Icones, iv. 17. Sydow, Monogr. i. 113. Fischer,
Ured. Schweiz, p. 221, f. 173.
‘Spermogones. Similar to those of P. Cirsit,

Uredospores. Sori of two kinds; primary epiphyllous, on
discoloured rounded or irregular spots,
in concentric rings round the spermo-
gones, sometimes confluent, 1—5 mm.

broad, flat, surrounded by the cleft.
we epidermis, pulverulent, cinnamon;.
secondary amphigenous, without spots,

about 4—1 mm. diam., scattered or

Fig. 90. P. Bardanae. Te- gregarious; spores globose to ellipsoid,
aaa pe ae enn echinulate, pale-brown, 26—30 x 22—
27 w, with three (rarely four) germ-

pores.

Teleutospores. Sori amphigenous, generally hypophyllous,
without spots, scattered or in places densely gregarious,
3—1 mm. diam., rounded, pulverulent, black-brown; spores
ellipsoid, rounded at both ends, not thickened, gently constricted,
delicately verruculose, dark-brown, 30—42 x 22—27 uw; pedicels
hyaline, short.

On Arctium Lappa. September, October. A doubtful
native; I have seen no British specimens. Description after
Sydow. (Fig. 90.)

Distinguished from P. Cirsit by its larger teleutospores and by its two
kinds of uredo-sori. Jacky demonstrated the distinctness of the two
species by experimental cultures (Centralbl. f. Bakt. 2. ix. 796) ; also that
P. Bardanae could not be transferred to Taraxacum.

DisrRrisution: Europe only.

ON COMPOSITE 139

12, Puccinia tinctorie Magn.

Puceinia Compositarum var. Serratulae Cooke, Exsice. no. 33.
P. tinctoriae Magn. in Abhand. Nat. Gesell. Niirnberg, 1900, xiii. 37 ;

Ber. deutsch. Bot. Gesell. 1893, pl. 21, f. 27—8. Sydow, Monogr.
i. 150, f. 127, ,

Uredospores. Sori amphigenous, on inconspicuous pallid
spots, scattered, minute, roundish, pulverulent, dark-cinnamon ;

spores globose to ellipsoid, echinulate, brown, 24—40 x 19—26 u;
epispore with two germ-pores towards the summit.

Fig. 91. P. tinctoriae. Teleutospores and uredospore (from
Cooke’s specimens).

Teleutospores. Sori similar, but blackish; spores ellipsoid
to oblong, rounded at both ends, not thickened above, hardly
constricted, delicately verruculose, brown, 27—60 x 19—30 p;
pedicels hyaline, up to 14 long; epispore thin.

On leaves of Serratula tinctoria. Apparently very uncom-

mon; Belton Wood, Highgate (M. C. Cooke). Teleutospores,
July—September. (Fig. 91.)

DistrisuTion: France, Sweden, Germany, Austria, Italy,
Siberia.

13. Puccinia Centauree DC.

Puceinia Compositarum Sch.; Cooke, Handb. p. 499; Micr. Fung.
p. 206 p.p.

P. Centaureae DC. Flor. fr. vi. 595. Cooke, Micr. Fung. p. 207.
Plowr. Ured. p. 186. Sydow, Monogr. i. 39. Fischer, Ured.
Schweiz, p. 222, f. 1745.

140 PUCCINIA

Spermogones. Chiefly epiphyllous, in little clusters, orange.
Uredospores. Sori generally hypophyllous, on yellowish or
brownish spots (spots sometimes
none), primary rather large,
secondary very minute, much
scattered, pulverulent, brown;
spores globose to ellipsoid, acu-
leolate, brown, 22—30 x 16—
Fig. 92. P. Centaureae. Teleuto- 28 yw, with three (or two) germ-
spores and uredospore.
pores.

Teleutospores. Sori similar, but blackish-brown; spores
ellipsoid or somewhat obovate, rounded at both ends, not thick-
ened above, not or hardly constricted, delicately verruculose,
chestnut-brown, 24—40 x 16—25 4; epispore thin; pedicels
hyaline, thin, generally very short.

On Centaurea nigra. Teleutospores, August—November,
Common. (Fig. 92.)

As Plowright says, there are two generations of uredospores, The
primary uredospores, which are the equivalent of the zcidiospores, accom-
panied by spermogones, appear in May, and are followed by the secondary
uredospores, which form much smaller sori. The former, however, do not
occur on every infested plant, being only produced from direct infection
by the basidiospores. The primary uredospores are widely scattered by
the wind; Plowright proved that they produce the secondary spores in
about fourteen days. Fischer and Jacky record this also on C. Scabiosa,
but the Puccinie on other species of Centuurea are considered to be
distinct species or else biological races.

DIstTRIBUTION: Europe, Asia Minor, Siberia, North America.

14. Puccinia Cyani Pass.

Uredo Cyani Schleich. Plant. Hely. 95.

Puceinia suaveolens var. Cyant Winter, Pilze Deutschl. p. 190. Plowr.
Ured. p. 183. Sace. Syll. vii. 633.

P. Cyant Pass. in Rab. Fung. Eur. 1767. Sydow, Monogr. i. 38.

Spermogones. A few mixed with the primary uredo-sori.
Uredospores. Sori generally hypophyllous, without. spots,
scattered or crowded, minute, roundish, pulverulent, cinnamon ;

ON COMPOSITE 141

spores globose to ellipsoid, delicately echinulate, yellow-brown,
22—30 x 19—24 y, with two germ-pores.
Teleutospores. Sori amphigenous and on the stems, scattered,
minute, punctiform, pulverulent,
blackish-brown ; spores broadly ellip-
soid, rounded at both ends, not
thickened above, not constricted,
very delicately verruculose, chestnut-
brown, 830—35 x 22—27 w, sometimes

longer ; pedicels hyaline, short.
eon y , Fig. 93. P. Cyani.
On Centaurea Cyanus. Not un- ii cae

common in gardens. June—September. (Fig. 93.)

There is a similar species, P. montana, which occurs on the continent
on Centaurea montana ; it is distinguished by the much coarser warts of
the teleutospores, which are also relatively longer. Both these species,
like P. obtegens, permeate the whole plant with the mycelium of the
primary uredospores, but such infested plants of Centaurea can flower
freely, while those of Cirsiwm arvense never do. The mycelium of the
secondary uredospores, mixed with teleutospores, is more strictly localised.

DisTRIBUTION: Western and Central Europe.

15. Puccinia Carduorum Jacky.

Puceinia Hieracit Plowr. Ured. p. 185 p.p., see also p. 216.
P. Carduorum Jacky, Composit.-Puccin. p. 58, Sacc. Syll. xvi. 297.
Sydow, Monogr. i. 33.

Uredospores. Sori generally hypophyllous, on very indis-
tinct spots, scattered, minute, pulverulent, cinnamon; spores
globose or subglobose, densely echinulate, pale-brown, 22—28 p,
with three germ-pores.

Teleutospores. Sori similar, but dark-brown; spores variable,
oblong to ovate, rounded above, not thickened, hardly at all
constricted, rounded below, verruculose, brown, 25—88 x 17—
24 4; epispore thin; pedicels hyaline, short.

On Carduus crispus, C. nutans. Not common; Yorkshire,
Clare Island, etc.

142 PUCCINIA

This species was formerly undistinguished from the numerous forms
on allied species of Composite, until Jacky proved experimentally (in
1899) that it is confined to the sub-genus Carduus, and could not be
transferred to Cnicus or Cirsium. The uredo form described above (after
Sydow) is the secondary uredo ; the primary form, which does not always
occur, is described as amphigenous and larger, often on the mid-ribs
2—4 mm. long, and remaining long covered by the epidermis.

DisTRiBuTION : Europe and Siberia.

16. Puccinia Cardui-pycnocephali Syd.

Puccinia Cardut-pycnocephali Sydow, Monogr. i. 34, f. 35. Massee,
Journ. Bot. xlvi. 152; Trans. Brit. Myce. Soc. (1909), iii. 123.

Uredospores. Sori hypophyllous, without spots, scattered,
minute, pale-brown; spores globose or subglobose, very deli-
cately echinulate, pale-brown, 22—26 p.

Teleutospores. Sori similar, hidden in the tomentum of the
host, brown; spores oblong, rounded above and more or less so
below, not thickened, hardly constricted, smooth, pale-brown,
38—50 x 16—23 ; epispore thin; pedicels thin, hyaline, up
to 40 long, but deciduous.

On leaves and stems of Carduus pycnocephalus. Very rare;
Sidmouth (Dr Mayor), and between Eastbourne and Hastings
(G. Massee). Only recorded for Britain and Italy. July,
August.

Distinguished from P. Carduorum by its longer teleutospores, which
are smooth, and not plainly verruculose as in that species. But sometimes
the spores are said to approach those of P. galatica Sydow, which occurs
on the same host in Asia Minor, in being smaller, darker-coloured, thick-
walled, and delicately punctate. According to Massee (i.c.) these are
rather two poles of one species than two distinct species.

17. Puccinia Cirsii Lasch.

Trichobasis Cirsti Lasch ; Cooke, Micr. Fung. p. 224.

Puccinia Cirsii Lasch, in Rab. Fung. Eur., no. 89. Cooke, Grevillea,
iv. 109; Micr. Fung. p. 206. Sydow, Monogr. i. 55. Fischer,
Ured. Schweiz, p. 217, f 168—171.

Uredospores. Sori epiphyllous or hypophyllous, on paler
spots, scattered, minute, girt by the epidermis, pulverulent, cin-
namon ; spores globose to ovate, echinulate, pale chestnut-brown,

ee ON COMPOSITE 143

22—28 x 19—24 yu, with three more or less equatorial germ-
pores each provided with a thickening.

Fig. 94. P. Cirsii. Teleuto- Fig. 95. P. Cirsti. Teleuto-
spores, on C. palustre, from spore and uredospore, on
Hereford. C. lanceolatum.

Teleutospores. Sori mostly hypophyllous only, similar, but
blackish-brown or black ; spores ellipsoid or somewhat obovate,
rounded at both ends, not thickened above, hardly constricted,
verruculose or merely punctate, chestnut-brown, 25—88 x
17—25 «; epispore thin ; pedicels hyaline, very short:

On Cirsium, Dupplin Castle, Perth (M. C. Cooke). On
C. pratense, Ballyquirke Lake, Co. Galway (communicated by
J. Adams); Earlswood Lakes, near Birmingham. On C. palustre,
Hereford, Seckley Wood, Barnt Green, etc. Uredospores from
March; teleutospores, June—November. (Figs. 94, 95.)

There is no mention of this in Plowright’s Uredines, or in the Trans.
Brit. Myc. Soc. (Plowright’s list), but it is probably not uncommon. It
occurs frequently on the radical leaves, and can be easily distinguished
from P. Cnict-oleracex by the presence of the uredospores and the non-
circinate teleuto-sori, as well as by the absence of the apical thickening.
Fischer records it from Switzerland on many species of Cirstwm (but not
on those mentioned here), and also assigns to it spermogones on the upper
leaf-surface and petiole ; I have not been able to find any trace of these in
our British specimens. The uredospores, seen in water, sometimes
appear quite smooth, as Cooke describes them. The teleutospores have
the upper pore at the summit or at the side, the lower pore just beneath
the septum or lower down; they are at times faintly granulated, at others
distinctly verruculose. Mesospores are rare.

T have also a number of specimens on C. lanceolatum from Droitwich,
Wyre Forest, etc., bearing a great similarity to P. Cirsit-lanceolati Schrot.,
but the differences from P. Cirsit are so slight and elusive that, as the eecidial
stage by which the former is distinguished has not yet been found in
Britain, it is better for the present to place them under P. Cirsiv.

DisTRiBuTION: Europe, Siberia, Japan, North America.

144, PUCCINIA

18. Puccinia Cnici-oleracei Pers.

Puccinia Cnici-oleracei Pers. in Desm. Cat. des Pl. omis., p. 24,
Sydow, Monogr. i. 58, pl. 3, f. 47. Fischer, Ured. Schweiz,
p. 292, 594, f. 213.

P. Cardui Plowr. Ured. p. 216.

Teleutospores. Sori hypophyllous, on discoloured roundish
spots, minute, circinate, but mostly confluent in large patches
as much as 5 mm. diam. which remain for a long time covered
by the epidermis, compact, blackish-brown; spores clavate or
subfusiform, rounded or rarely somewhat conical above, very

c

Fig. 96. P. Cnici-oleracei. Teleutospores, on C. lanceolatum.,

much thickened (5—10 »), constricted, attenuated below,
smooth, yellowish-brown, 838—56 x 14—21 y; pedicels yellowish-
hyaline, thick, persistent, as much as 50 u long.

On Cirsium lanceolatum, Carduus crispus (?). August—
October. Not uncommon on the former. (Fig. 96.)

This is one of the species in which the upper cell separates from the
lower one. It is doubtful if Pucetnia Syngenesiarum (Cooke, Handbook,
p. 499; Micr. Fung. p. 206) belongs entirely here, as the figure in the
latter work (pl. 4, f. 64) does not give the true form of the teleutospore ;
but the majority of the specimens issued by him under that name are this
species. It is very questionable if this species occurs on Carduus crispus,
on which it is recorded by Plowright.

DISTRIBUTION : in a few places in Europe.

ON COMPOSITE 145

19. Puccinia obtegens Tul.

Caeoma obtegens Link, Obs. ii. 27.

Trichobasis suaveolens Lév.; Cooke, Handb. p. 530; Micr. Fung.
p. 226, pl. vii. figs. 151—3,

Puccinia obtegens Tul. Ann. Sci. Nat. ser. 4, ii. 87 (1854). Sydow,
Monogr, i. 53,

P. suaveolens Rost. Bot. Zeit. 1874, p. 556. Plowr. Ured. p. 182 p.p.
Sacc. Syl. vii. 633 p.p. Fischer, Ured. Schweiz, p. 219, f. 172.

Spermogones. Chiefly hypophyllous, a few epiphyllous,
crowded, covering the whole surface of the leaf, of a bright
honey-yellow colour and a pleasant smell.

Uredospores. Primary sori hypophyllous, occupying the
whole surface of the leaf, minute, crowded, often confluent,
pulverulent, reddish-brown, then darker; secondary, more scat-
tered; spores globose to broadly ellipsoid, echinulate, pale-
brown, 21—28 », with three irregularly placed germ-pores.

Teleutospores. Sori similar, always dark-brown; spores

ovate to ellipsoid, rounded at both ends, or

somewhat tapering below, not thickened
above but with a low flat pore-cap, hardly
constricted, delicately verruculose, brown,
26—42 x 17—25 py; epispore thin; pedicels
hyaline, thin, short.

Fig. 97. P. obtegens.
On Cirsium urvense (Carduus arvensis). Teleutospores,
Very common. (Fig. 97.)

The life-history of this species is peculiar. In spring the mycelium
permeates the host in every part. The affected plants can be recognised
immediately by their pale-green colour and spindly appearance; they
never flower. The spermogones are first seen towards the end of April,
and are easily detected by their bright colour, and their strong perfume,
resembling that of privet-flowers; the uredospores follow on the same
leaves during May. From these primary uredospores, a second generation
arises on other plants about July, and forms secondary uredospores and
tcleutospores in sori which are more scattered, never confluent, and
darker brown. This generation is not accompanied by spermogones,
The mycelium of these sori is localised to the infected spots and the host
does not suffer so severely. The sori of the primary uredospores rarely
bear a few teleutospores intermixed, but the secondary sori abound with
them from September to November, and it is from the germination of

G. U. 10

146 PUCCINIA

these latter that new infections arise in the spring as soon as the shoots
appear. The hibernation of the mycelium in the rhizome, which is stated
by Plowright, has not been proved.

The germ-pores are very easy to see in the uredospores of this species
and its allies. Each is often surrounded by a border like a bordered pit,
an appearance caused by a thickening of the membrane around and over
the pit. As usual the appearance of the spore changes according as it is
wet or dry ; if wet, it may appear merely punctate; if dry, it is seen to be
densely and coarsely echinulate.

DisTRIBUTION: Northern and Central Europe, Siberia,
North America.

20. Puccinia Andersoni B. et Br.

Puceinia Andersoni B. et Br. Ann. Nat. Hist. ser. 4, xv. 35. Cooke,
Micr. Fung. p. 206; Grevillea, iii, 179. Plowr. Ured. p. 204.
Sace. Syll. vii. 710. Sydow, Monogr. i. 58.

Teleutospores. Sori hypophyllous, seated on round yellow
spots 1—14 cm. diam. with a brown

C) border, almost concealed by the
pubescence of the leaf, minute, but
densely crowded in flat circular

clusters which are few on each leaf,

compact, blackish-brown or purplish-

black; spores oblong to clavate,

\ rounded or conically thickened (8—

10%) above, slightly constricted,

smooth, pale-brown, 40—54 x 16—

22; pedicels brownish, stout, per-

af ; sistent, as long as the spore or

Be recall ie longer; a number of mesospores

are found intermixed.

On Cirsium heterophyllum. June—October. A striking
and rare species. Glen Ogle (Rev. M. Anderson), Den of
Airlie (Mr Gardiner), Ingleton and Grassington, Yorkshire
(H. T. Soppitt), Alston, Cumberland (J. G. Baker). (Fig. 98.)

DISTRIBUTION : in a few places in Europe.

ON COMPOSITE 147

21. Puccinia Lapsane Fckl.

Heidium Lapsanae Schultz, Prod. Flor. Starg. p. 454. Cooke, Journ.
Bot. ii. 38, pl. 14, f. 2; Handb. p. 543; Micr, Fung. p. 198.

Trichobasis Lapsanae Cooke, Micr. Fung. p. 224.

Puceinia Lapsanae Fckl. Symb. Mye. p. 53. Cooke, Micr. Fung.
p. 207. Plowr. Ured. p. 149. Sacc. Syll. vii. 607. Sydow,
Monogr. i. 112. Fischer, Ured. Schweiz, p. 203, f. 159.

Spermogones. Crowded in little clusters, epiphyllous, honey-
coloured.

4Aicidiospores. Aicidia amphigenous, somewhat crowded on
large, roundish, purple spots, flattish, with torn white reflexed
margin; spores subglobose or ovate, nearly smooth, orange,
16—21 x 13—17 p.

Uredospores. Sori amphigenous, very minute, round, very
numerous, often confluent, pulverulent, chestnut-
brown ; spores globose to ovate, delicately echinu- fo
late, pale-brown, 17—22 x 15—18 p, with two
germ-pores.

Teleutospores. Sori amphigenous, minute,
scattered, numerous, pulverulent, blackish-brown; fig 99, p,
spores ellipsoid or ovate, rounded at both ends, Perc lea

A ‘ eleutospore.

not thickened above, scarcely constricted, very
delicately punctate, chestnut-brown, 22—33 x 17—26 yu; pedi-
cels hyaline, slender, short, often oblique.

On leaves and stems of Lapsana communis. Very common.
Aicidia, March—May ; uredospores, April—June; teleutospores,
June—September. (Fig. 99.)

The ecidium of this species shares with those on Ranunculus Ficaria
and Potentilla Fragariastrum the distinction of being the earliest Uredine
to make its appearance in the spring. It may be found on the young
leaves of the seedlings almost as soon as they are formed. Plowright
demonstrated (Expt. 499, May, 1885) that all three spore-forms belong to
the same life-cycle. The mycelium of the ecidia, when occurring on the
petioles, causes them to become pale and swollen ; on the leaves it often
produces conspicuous purple spots which bear the spermogones on the
upper surface.

DistRisuTion: Europe, Syria, Japan. |
10—2

148 PUCCINIA

22. Puccinia Cichorii Bell.

Uredo Cichorit DC. Flor. fr. vi. 74.

Puccinia Hieracié Mart. ; Plowr. Ured. p. 184 p.p.

P. Cichorti Bell..in Kickx, Fl. Fland. ii. 65. Sydow, Monogr. i. 49,
Fischer, Ured. Schweiz, p. 227, f. 179. McAlpine, Rusts of
Australia, p. 154, f. 61 and pl. D, f. 18.

Uredospores. Sori amphigenous or on the stems, numerous,
scattered, sometimes confluent, minute, pulverulent, surrounded
by the torn epidermis, cinnamon; spores globose to ellipsoid,
echinulate, yellowish-brown, 21—27 4; epispore moderately
thick, with two germ-pores.

Teleutospores. Sori similar, but blackish-brown; spores
ellipsoid, rarely obovate, rounded above and not thickened,
hardly constricted, generally rounded below, smooth or faintly
verruculose, brown, 27—38 x 19—25 uw; epispore thin; pedicels
hyaline, short, deciduous.

On Cichoriwm Intybus. Rare. August—October.

This species closely resembles Puccinia Hieracii, and there seem to
have been no cultures which give any information about its distinctness.
It is therefore separated merely on the ground of the difference of the
genus of the host; see under P. Hieraciz.

DIsTRIBUTION : Europe; introduced into Australia.

23. Puccinia Hypocheridis Oud.

Puceinia Hypochoeridis Oud. in Nederl. Kruidk. Archief. ser. 2, i. 175.
Sydow, Monogr. i. 100. Fischer, Ured. Schweiz, p. 232. McAlpine,
Rusts of Australia, p. 159, f. 62—38.

Uredospores. Sori amphigenous or on the stems, generally
on minute spots, scattered, pulverulent, cinnamon, primary
sori rather large, secondary minute ; spores globose to ellipsoid,
echinulate, brown, 22—28 yw, with two germ-pores.

Teleutospores. Sori similar, but punctiform (chiefly and
larger on the stems), black; spores ellipsoid to ellipsoid-obovate,
rounded at both ends or rarely attenuated downwards, not
thickened above, hardly constricted, delicately verruculose-
punctate (?), brown, 30—46 x 18—24 yw; epispore thin; pedicels
hyaline, short.

ON COMPOSITA 149

On Hypochoeris radicata. Not uncommon. July—Sep-
tember. The records from other countries include all the three
British species of Hypochoeris. Only uredospores were seen
in British specimens. (Figs. 100, 101.)

e / =

i

\

Fig. 100. P. Hypochoeridis. Fig. 101. P. Hypochoeridis. Teleuto-
Uredospore (British) on spores and uredospore, on H. glabra
H. radicata. (Berlin, ex herb. Sydow).

This species, which has been often stated to be a Brachypuccinia, differs
from most others of the type of P. Mveracii in having two kinds of
uredo-sori—the primary ones 1—2 mm. wide, and only on the leaves, the
secondary ones conspicuously smaller, almost punctiform. In this respect
it approaches P. Cyani and P. obtegens, but Jacky showed that in his
cultures it did not produce spermogones (Centralbl. f. Bakter. 1907, xviii.
83), as they do. The alleged punctations of the teleutospores were
invisible in all the specimens I have seen.

DIsTRIBUTION: Europe, Siberia, North America, Chili,
Australia.

24. Puccinia Leontodontis Jacky.

Puccinia Hieracti Plowr. Ured. p. 184 p.p.
P. Leontodontis Jacky, Composit.-Puccin. p. 75. Sydow, Monogr. i.
114. Fischer, Ured. Schweiz, p. 281, f. 182.

Uredospores. Sori amphigenous, but chiefly hypophyllous,
scattered, not confluent, minute, punctiform, surrounded by the
cleft epidermis, cinnamon; spores globose to ellipsoid, echinu-
late, brown, 25—32 p diam. or 27—35 x 24—27 p, always with
two opposite germ-pores above the equator.

Teleutospores. Sori similar, but darker-brown; spores
variable, ellipsoid to oblong or obovate, rounded at both ends, ©
not thickened above, not constricted, delicately verruculose or

150 PUCCINIA |

faintly granulated, chestnut-brown, 830—42 x 21—27 ; epispore
thin; pedicels hyaline, short or nearly as long as the spore.

Fig. 102. P. Leontodontis. Teleutospores and uredospore, on L. autumnalis,

On Leontodon autumnalis, L. hispidus. August, September.
Not uncommon. (Fig. 102.)

The teleutospores of this species are said to be more than usually
variable ; sometimes the sori, according to Sydow, are seated on coloured
spots, but more often the spots are wanting. Up to the present, no
experimental cultures appear to have been carried out with this Puccinia,
and it is separated from P. Hieracit mainly because it is parasitic upon a
different genus. I have found it mostly upon old yellowing leaves.

DISTRIBUTION : Europe generally.

25. Puccinia Tragopogi Corda.

Aeidium Tragopogi Pers. Syn. p. 211.

4%. Tragopogonis Cooke, Handb. p. 537; Micr. Fung. p. 195, pl. 1,
f. 1—3.

Puceinia Tragopogt Corda, Icon. v.50. Plowr. Ured. p. 197. Sace.

Syll. vii. 668 p.p. Sydow, Monogr. i. 167. Fischer, Uved.
Schweiz, p. 215, f. 166.

P. sparsa Cooke, Handb. p. 498; Micr. Fung, p. 205.

Spermogones. Epiphyllous, honey-coloured, sometimes
absent.

icidiospores. Aicidia hypophyllous, without spots, scat-
tered uniformly over the whole surface, and on other green
parts, cup-shaped, with a white torn revolute margin; spores
globose to ellipsoid, densely verrucose, pale-orange, 20—30 x
18—24 4, with three germ-pores.

ON COMPOSITA 151

Teleutospores. Sori amphigenous, without manifest spots,
scattered, occasionally aggregated, mi-
nute, punctiform, long covered by the
epidermis, at length pulverulent, dark-
brown; spores ellipsoid, rounded at
each end, not thickened above, gently
constricted, tuberculate, chestnut-
brown, 26—45 x 18—32; pedicels ae oa
hyaline, short.

On Tragopogon pratensis. Not uncommon. April—Sep-
tember. (Fig. 103.)

The mycelium arising from the infection of young plants by the basi-
diospores permeates the whole of the host, so that ecidia are produced on
every part—stems, leaves, bracts, and receptacles—and the infected plants
are noticeable for their paler colour and distorted form. It is believed to
hibernate also in the upper part of the root-stock. The mycelium of the
teleuto-sori is, on the contrary, strictly localised. Uredospores are not
produced in separate sori, but a few can be found intermixed with the
teleutospores, as well as a few mesospores ; the former can be distinguished
from the latter by being delicately aculeate, not tuberculate. The
teleutospores of this species are grossly and distinctly warted, reminding
one of those of Uromyces tuberculatus; they can be found at the same time
as the wcidia, but are much rarer, or at least less frequently observed.

DIsTRIBUTION: Europe, Asia Minor.

26. Puccinia Chondrille Corda.
Ascidium Prenanthis Pers. Syn. p. 208 (non Cooke, Handb. p. 542 ;
Micr. Fung. p. 198).
Puceinia Chondrillae Corda, Icones, iv. 15. Fischer, Ured. Schweiz,
p. 200, f. 156.
P. Prenanthis Auct. p.p.; Plowr. Ured. p. 148. Sydow, Monogr. i.
106. McAlpine, Rusts of Australia, p. 162 (?).

Ajcidiospores. Aucidia hypophyllous or on the petioles,
rarely a few on the upper surface, seated on large yellow and
purple spots, in clusters as much as 6—8 mm. broad, at first
hemispherical, opening by a pore, then flattened, whitish or
yellow, sometimes with a purplish tinge; spores globose to
ellipsoid, delicately verruculose, pallid-orange, 183—24y; peri-
dium poorly developed.

152 PUCCINIA

Uredospores. Sori hypophyllous, on pallid irregular spots,
scattered, minute, punctiform, pul-
verulent, pallid-brown; spores more-
or less globose, echinulate, yellow-

ED brown, 16—24 y, with three (rarely

ww four) germ-pores.

Teleutospores. Sori similar,
surrounded by the cleft epidermis,
blackish-brown; spores ellipsoid,
rounded and not thickened above,
not constricted, rounded below, very delicately verruculose,
brown, 26—86 x 16—24 uw; pedicels hyaline, very short.

Fig. 104. P. Chondrillae. Te-
leutospore and uredospores.

On Lactuca (Prenanthes) muralis only. Not common.
April—September. All three spore-forms may be found on the
same leaf. (Fig. 104.)

The ecidium of this species is not provided with a proper peridium
and opens with a pore, never assuming the form of a cup; moreover the
zecidiospores are more round than angular. The germ-pores of the uredo-
spores are covered with a broad convex colourless cap, which swells up
somewhat in water. The genetic connection of the three spore-forms has
been proved experimentally by Jacky (Centralbl. f. Bakt. ix. 1902, p. 842),
as also the fact that the parasite is not transmissible to other species of
Lactuca. There is another ecidium also found on Lactuca muralis on the
continent which possesses a well-developed peridium ; this belongs to
Puccinia Opizit Bubdék, and has its teleutospores on Corer muricata.

DISTRIBUTION: Europe generally.

27. Puccinia variabilis Grev.

Meidium Grevillei Grove, Journ. Bot. 1886, p. 129.

Puccinia variabilis Grev. Scot. Crypt. Fl. pl. 75. Cooke, Handb.
p. 500; Micr. Fung. p. 207, pl. 4, f. 82—3. Plowr. Ured. p. 150.
Sydow, Monogr. i. 163. Fischer, Ured. Schweiz, p. 202, f. 158.

Acidiospores. AXcidia amphigenous, on minute indetermi-
nate yellow or purplish spots, solitary or a few loosely aggre-
gated, cup-shaped, with whitish torn margin; spores subglobose
or ovate, with orange contents, verruculose, 20—25 x 15—20 p

Uredospores. Sori amphigenous, on very minute yellow or

ON COMPOSITE 153

purplish spots, scattered, minute, punctiform, soon naked,
brown; spores few, globose to ovate, echinulate, brown, 22—
32 x 19—26 w, with two germ-pores.

Teleutospores. Sori similar, but darker; spores ellipsoid to
oblong, rounded at both ends, not thickened above, not con-

se ON
Fig. 105. P. variabilis, Hicidia, on leaf of Taraxacum, and teleutospore.

stricted, delicately verruculose, brown, 28—40 x 18—25 pw;
epispore thin; pedicels hyaline, about as long as the spore, but
deciduous.

On Turaxacum officinale and its variety palustre. July—
October. Not common. (Fig. 105.)

Plowright and Soppitt both proved, vy laying leaves affected with the
ecidium of this species on healthy plants of Taraxacum, that the uredo-
and teleutospores were produced in about fourteen days. In July the
three spore-forms may be found on the same leaf.

There are two forms of ecidium found upon Taraxacum ; one, Aeidium
Grevillei Grove (=4. Taraxaci Grev. non K. et S.), spreads pretty
uniformly over the whole leaf in “numerous little clusters with single ones
scattered between them,” as Greville describes it (Flor. Edin. p. 444)—the
other, @. Tarazaci K. et &., forms large round clusters, and belongs to
P. silvatica, Fischer points out that the two ecidia differ in the form of
their peridium cells, those of P. variabilis having the membrane thickened
on the znner side, while those of P. silvatica have the outer wall most
strongly thickened. He states, furthermore, that it will be found that
this difference is characteristic in general of autcecious and hetercecious
species respectively. It is not, however, universally so, e.g. the ecidium
of P. albescens has the outer wall much more strongly thickened, although
it is autcecious.

Greville figures the teleutospores of his species (Scot. Crypt. Flor. pl]. 75)
as having either one or both of the cells sometimes divided by a vertical
septum.

DistTRIBUTION: Switzerland, Sweden.

154 PUCCINIA

28. Puccinia Taraxaci Plowr.

Puceinia Phaseoli var. Taraxaci Rebent. Fl. Neomarch. p. 356.

P. variabilis Grev. ; Cooke, Handb. p. 500; Micr. Fung. p. 207 p.p.

P. Taraxact Plowr. Ured. p. 186. Sacc. Syll. ix. 305. Sydow,
Monogr. i. 164. Fischer, Ured. Schweiz, p. 226, f. 178,

Spermogones. In little yellow roundish clusters.
Uredospores. Sori amphigenous, with or without spots,
scattered, minute, sometimes confluent and larger, roundish or
oblong, pulverulent, brown; spores glo-
bose to ovate, echinulate, pale-brown,
22—27 x 16—24 py, with two germ-pores.
Teleutospores. Sori similar, but
blackish, 4—3 mm. diam.; spores ellip-
Fig, 106, Fi Doravact, soid to ovate, rounded at both ends, not
Teleutospore anduredo- thickened above, not constricted, very
oe delicately verruculose, brown, 25—388 x
16—24; epispore thin; pedicels hyaline, short.

On Turaxacum officinale, Rather common. Spermogones
and primary uredospores in April; the teleutospores may be
found till November. The distinctions which Plowright attempts
to draw between the primary and secondary uredospores are
not so marked as is the case in P. Centaureae, and break down
in practice. (Fig. 106.)

This species differs from P. variabil’s, with which it was formerly
confused, chiefly in the absence of the ecidium. But, in addition to that,
the uredospores of P. Taraxact are far more abundant and the sori more
especially found on the upper leaf-surface ; the uredospores of P. variabilis
are scanty and are usually intermixed in the teleuto-sori. P. Taraxaci is
morphologically indistinguishable from P. Hieracii, but culture experi-
ments have proved that it cannot be transferred from Taraxacum to the
allied genera of the Composite.

Plowright’s remark (/. ¢. p. 187) that I considered this species to have
“a true eecidium” is a mistake, arising probably from a confusion between
it and P. variabilis.

DisTRIBUTION : Europe, North America, Japan, East Indies.

ON COMPOSITA 155

29. Puccinia Sonchi Rob.

Headium Sonchi Johnst.; Plowr. Ured. p. 266.

Puccinia Sonchi Rob. in Desm. Ann. Sci. Nat. ser. 3, xi. 274. Grove,
Sci. Gossip, 1885, p. 9, f. 6—9. Plowr. Ured. p. 196. Sauce. Syll.
vii. 638. Sydow, Monogy, i. 154,

[Spermogones. Obtained artificially on S. arvensis by Tranz-
schel. |

Uredospores. Sori generally hypophyllous, on the stem
collected in small oblong patches 4—5 mm. long, on yellowish
spots, scattered or gregarious, minute, at first covered by the
epidermis which is raised over them in a hemispherical vesicle,
then pierced at the top but never widely open, yellowish,
surrounded by a dark-brown line of paraphyses; spores more or
less oblong or obovate, densely verrucose, with a very thick
epispore, contents oily, yellowish, 2438 x 15—21 pu.

Fig. 107. P. Sonchi. a, the line of dark-brown paraphyses surrounding the
uredo-sori; b, a pale-brown paraphysis; c, a uredospore; all on S.
oleraceus. - x 600.

[Teleutospores. Sori hypophyllous, rarely on the stems, on
irregular brownish spots, more compact, rounded-pulvinate,
scattered or aggregated, confluent when on the stem, covered
by the epidermis, black; spores ovate or oblong, rounded or
truncate (rarely attenuated) above, thickened (8—8,), con-
stricted, rounded below, smooth, pale-brown, 30—60 x 20—30 4p ;
pedicels somewhat long, brownish, persistent; mesospores ovate,

156 PUCCINIA

clavate, or oblong, thickened at the apex, brownish, 45—
60 x 20—25 4; paraphyses numerous, reddish-brown, clavate
and somewhat thickened at the apex.]

On Sonchus arvensis, 8S. asper, S. oleraceus. Uredospores
only seen, July—September. Rather rare. (Fig. 107.)

In Sydows’ Monographia this species is said to be confined to the
neighbourhood of the coast or nearly so. It has been recorded by Prof.
Trail at Aberdeen, by Mr Johnston at Berwick-on-Tweed, by Mr D. A. Boyd
from Ayrshire; also from Sutton, near Askern, and Mulgrave Woods in
Yorkshire ; I have received specimens from Mr Hawkes collected near
Birmingham, from Mr Phillips near Hull, from Mr T. B. Roe near
Scarborough, and from Mr J. Adams at Howth, Co. Dublin, and Westport,
Co. Mayo.

It is a remarkable species, and worthy of close investigation. Though
the uredo is sometimes confounded with Coleosporium Sonchi, it is readily
distinguished by its brown paraphyses which form only a single row
round the sori and are easily seen with a lens through the epidermis as a
dark line surrounding the yellow spore-layer. It resembles at first sight
an ecidium, and has been more than once described as such : but it opens
by a pore the edges of which do not curl back. The so-called paraphyses
are really the upper part of a delicate imperfect peridium, composed of
hyaline pseudo-parenchyma (cells 5—10, diam.) ; at the top these cells
become elongated, linear, parallel, at first colourless, then brownish and
more or less clavate, and finally very dark brown, subopaque and irregular.
This colour is retained for many years in the dried specimens, though the
spores are bleached. The peridium resembles in some respects that which
surrounds the uredo-sori of Melampsorella Caryophyllacearum. The spores
themselves are at first sight like ecidiospores, with thick colourless walls,
and yellowish contents, the sculpture resembling that of the zcidiospores
of Endophyltium. Ultimately the wall becomes thinner and brownish; the
spores are borne singly on pedicels, like ordinary uredospores.

Tranzschel (Ann. Mycol. 1909, vii. 182) sowed the teleutospores on
S. arvensis and obtained spermogones, followed by the uredospores. This
species is widely different from a typical Puccinia.

DisTRIBUTION: Western Europe, Algeria, Canaries, J apan.

30. Puccinia Crepidis Schrit.

Puccinia Crepidis Schrot. Pilze Schles. p. 319. Sace. Syll. vii. 607.
Sydow, Monogr. i. 64. Fischer, Ured. Schweiz, p. 207, f. 163.

[Spermogones. Scattered amongst the ecidia, nearly always
present, in little clusters of 6—10, brownish.

ON COMPOSITE 157

4icidiospores. Akcidia hypophyllous, evenly spread over the
whole leaf-surface, flat, with white margin; spores delicately
verruculose, yellowish, 15—25 x 14—20 mu]

Uredospores. Sori amphigenous, minute, roundish, scat-
tered, surrounded by the epidermis, cin-
namon.; spores globose to ovoid, delicately
aculeate, brown, 20—24 x 16—20 uy,
with two or three germ-pores.

Teleutospores. Sori hypophyllous,
rarely also epiphyllous, minute, long (
covered by the epidermis, blackish-brown; Fig. 108. P. Crepidis.
spores ellipsoid to ovoid, rounded at both HRN Nona
ends, not thickened above, scarcely or not at all constricted,
very delicately punctate (7), chestnut-brown, 20—30 x 17—22y;
epispore thin ; pedicels hyaline, very slender.

On Crepis virens. Rare. August. (Fig. 108.)

This species seems to be very rare. It is recorded for Scotland
(Moray, Rev. Jas. Keith) in Scot. Nat. 1884, p. 270, and I have a specimen
showing teleutospores from Ireland (J. Adams), but I have not seen the
zcidia on British specimens. The punctations were not discernible on
the teleutospores.

DISTRIBUTION: Central Europe.

31. Puccinia major Dietel.

eidium Compositarum var. Prenanthis Cooke, Handb. p. 542; Micr.
Fung. p. 198.

Puccinia Lapsanae Plowr. Ured. p. 149 p.p.

P. major Dietel, Mitth. Thiir. Bot. Ver. 1894, heft vi. Sacc. Syll.
xiv. 310. Sydow, Monogr. i. 66. Fischer, Ured. Schweiz, p. 214.

Spermogones. Generally hypophyllous, on reddish or yellow-
ish spots.

4icidiospores.. A&cidia hypophyllous, often surrounding the
spermogones, on the same thickened spots, in roundish clusters
or more often forming oblong patches on the nerves and petioles;
shortly cylindrical, with white torn erect margin; spores ovoid
or rarely subglobose, delicately verruculose, orange, 20—30 x
16—24 pu.

158 PUCCINIA

Uredospores. Sori amphigenous, solitary, minute, cinnamon;
spores subglobose to ovoid, distinctly
echinulate, brownish, 24—30 x 21—
26 pu.

Teleutospores. Sori chiefly hypo-
phyllous, similar, but blackish-brown,
standing singly, scattered over nearly
the whole leaf-surface ; spores ellipsoid
to ovoid, rounded at both ends, not
Hig, 10h. 2: amaiens. Te thickened above, hardly constricted,

leutospore, on C. palu. very delicately verruculose, chestnut-
ne ie aa on leaf of brown, 33—48 x 22—30; epispore
thin; pedicels short, deciduous.

On Crepis paludosa. Aicidia in June; uredospores, mixed
with teleutospores, August and September. Yorkshire (Sop-
pitt), Braemar (Trail), Dolgelly, Wales, and in Ireland.
(Fig. 109.)

This species was formerly considered a variety (major) of P. Lapsanae
on account of the larger spores, but is now kept separate. Dietel proved
by cultures that the uredo- and teleutospores are connected with the
eecidial generation. From P. Crepidis, on Crepis virens, it is said to be
distinguished, not only by the size of the spores, but also by the difference
in the way in which the ecidia occur on the leaves; the minute, black,
solitary teleuto-sori seem also different.

DisTRIBUTION: Northern and Central Europe.

32. Puccinia Hieracii Mart.

Uredo Hieractt Schum. Enum. Pl. Sal. ii, 232.

Trichobasis Hieracti Cooke, Micr. Fung. p. 224 p.p.

Puccinia Hieracti Mart. Fl. Mosquen. p. 226. Cooke, Micr. Fung.
p- 207 p.p. Plowr. Ured. p. 184, p. min. p. Sace. Syll. vii.
633 p.p. Sydow, Monogr. i. 95. Fischer, Ured. Schweiz, p. 230,
f. 181.

Spermogones. In little yellowish clusters on leaves and
stems.

Uredospores. Sori amphigenous, usually epiphyllous, on
pale spots which are sometimes hardly perceptible, scattered,
scarcely confluent, minute, punctiform, soon naked, pulverulent,

ON CAMPANULACE 159

cinnamon ; spores globose to ellipsoid, echinulate, yellow-brown,
24—29 x 16—25 yw, with two germ-pores.

Teleutospores. Sori similar and often on the stems, but
dark-brown; spores ellipsoid or somewhat
ovate, rounded and not thickened above,
scarcely constricted, usually rounded
below, very delicately verruculose, brown,
25—40 x 16—24 w; epispore thin ; pedi-
cels hyaline, usually very short.

Fig. 110. P. Hieracii.

On leaves and stalks of Hieracium, Enon

H. boreale, H. murorum, H. Pilosella, H. umbellatum and

various subspecies. Wery common. May—November. (Fig.
110.)

It was proved by Jacky that this species, which occurs so abundantly
on Hieracium, cannot be transferred to other genera of Composite. Asa
similar fact has been demonstrated for many other species of Uredinales,
there is sufficient ground for the assumiption, now generally made, that
most species of Puccinia, etc., which are parasitic on different genera
should be regarded as distinct, even when no experimental evidence exists
in favour of that course. Jacky was also inclined to suspect that
P. Hieracii might hereafter be divisible into a number of biologic races, of
which, however, he only indicated one, that on ZH. villosum belonging to
the section Pilosella. René Probst (Centralbl. f. Bakt. 1909, 2. xxii. 676)
not only divided P. Hieraciz into 13 biologic races, arranged under two
subspecies, P. Piloselloidarum on the section containing H. Pilosella and
its allies, and P. Hieracii (sens. strict.) on the other species, forming the
section Euhieractwm,—but he goes on to reduce the question of such
races to an absurdity by “proving” that one of them was restricted to a
mere form of a variety of a subspecies (Hieracium Pilosella, subsp.
vulgare, var. genuinum, forma subpilosum).

The two subspecies may perhaps be defensible, if they are distinguished
morphologically, as Probst states, by the fact that in P. Piloselloidarum
the germ-pores of the uredospore are strictly equatorial, but in P.
Hieractt they are removed towards the upper pole.

DistrisuTion: Europe, Asia Minor, North America, Chili.

33. Puccinia Campanule Carm.

Puccinia Campanulae Carmich. in Berk. Engl. Fl. v. 365. Cooke,
Handb. p. 498; Micr. Fung. p. 205. Plowr. Ured. p. 200. Sace.
Syll. vii. 677. Sydow, Monogr. i. 196, f. 182. Fischer, Ured.
Schweiz, p. 175, f. 136.

160 PUCCINIA

Teleutospores. Sori hypophyllous, rarely epiphyllous, often
on the petioles and stems, scattered or circi-
nate, minute, sometimes (especially on the
stems) confiuent and larger, long covered by
the epidermis, then surrounded by it, round-
ish or irregular, ferruginous-brown ; spores
ellipsoid or oblong, rounded and not thick-

Fig. 111, P. Cam- ened above, but with a pale papilla to each
panulae. Teleuto- :
spores, on C. ro- germ-pore, constricted, rounded or somewhat
ey attenuated below, smooth, pale-brown, 26—
45 x 12—22 w; pedicels hyaline, thin, deciduous, as long as the

spore or shorter.

On Campanula Rapunculus, C. rotundifolia. Not common.
June—August. (Fig. 111.)

According to Sydow, the statement in Cooke’s Handbook that Jausione
montana is one of the hosts of this species is entirely without corroboration
and is probably due to an error in the determination of the plant: the
statement is copied from the notice in Berk. Ann. Nat. Hist. ser. 2, v.
462—“ On Jasione montana, Lampeter, J. Ralfs.” Fischer mentions that
he has occasionally met with one-celled teleutospores, which I have not
seen.

DISTRIBUTION : Central Eurupe.

34. Puccinia Adoxe Hedw. f.

Aeidium Adoxae Opiz, in Kl. Herb. Myce. i. no. 780.

Puccinia Adoxae Hedw. in DC. Flor. fr. ii. 220. Cooke, Micr. Fung.
p- 209 p.p. Plowr. Ured. p. 207. Sace. Syll. vii. 612. Sydow,
Monogr. i. 203 p.p. Fischer, Ured. Schweiz, p. 146, f. 111.

P. Saxifragarum Schl. ; Cooke, Handb. p. 506 p.p. W. G. Smith, in
Gard, Chron. xxiv. (1885), p. 21, f. 7.

Teleutospores. Sori roundish, united in large clusters, often
confluent, on discoloured spots on the leaves, or on the petioles
forming elongated swollen patches, long covered by the silver-
grey epidermis, at length naked, pulverulent, dark-brown;
spores ellipsoid to broadly fusiform, rounded or attenuated
above, with a conspicuous colourless papilla, mostly rounded
below, scarcely ‘constricted, smooth, chestnut-brown, 28—42 x
14—21 »; pedicels hyaline, delicate, short, deciduous.

ON ADOXA 161

On Adoxa Moschatellina. March—May. Not uncommon.
All parts of the plant are affected, rhizomes,
petioles, leaves, peduncles and flowers.
(Fig. 112.)

Puceinia Adoxae and the ecidium of P. albescens
(q.v.) are about equally common, but are rarely
found together; they occur not only on different
plants, but usually also in widely separated locali-
ties. In fact, it is agreed by all observers that there
are three cases, (1) where the zcidium alone occurs, (2) where the teleuto-
spores alone occur, and (3) where they both occur together ; in the latter
case uredospores are found with the teleutospores in the same sorus. The
first is considered to be the ecidium of P. argentata (q.v.), the second is
P. Adoxae, and the third is P. albescens.

Soppitt first proved that the teleutospores of P. Adowxae, laid upon
healthy plants when in active germination after passing the winter,
reproduce the teleutospores in about ten days without the intervention of
other spore-forms. It is therefore a Micropuccinia. Whether the wide-
spreading mycelium is perennial or not is uncertain. Worth. G. Smith
raised seedlings of Adowva from berries of an infected plant ; the seedlings
exhibited the Puccinia from the earliest stages of growth, but we are not
told what precautions were taken to prevent infection from outside. He
also found (é.¢.) teleuto-sori, in a state of nature, on fusiform swellings
of the underground parts of the plant (peduncles and petioles), as also on
rhizomes and scales, in March ; the spores were irregular, one-, two-, or
three-celled. In April the leaves, and in May the flowers and young
fruits were infected. No mycelium could be found in the rhizome. If
Plowright’s ascription of a perennial mycelium is incorrect, the infection
must have first taken place, in this instance, underground on the young
growth, and the mycelium gradually spread upwards. That this is
probably the case is shown by Fischer’s experiment; he kept plants which
had borne teleutospores in pots—if he removed all the leaves, they
produced healthy growth next spring, whereas, if the leaves were left on
and allowed to fall upon the soil, one plant at least (out of four) showed
teleutospores on the new shoots. Bubdék inclines to the same opinion
(Centralbl. f. Bakt. 2. xvi. 150).

As will be seen from the synonymy, P. Adowae and P. albescens are
united (under the former name) in Sydows’ Monographia, while Soppitt,
Plowright, Bubék and Fischer consider them distinct. In any case they
are very closely allied ; the probability is that P. Adoxae is a mutation
from P. albescens which has acquired the habit of reproducing teleutospores.
from teleutospores directly, while the original species from which it was.
evolved still maintains all the three spore-forms, though two of them are

Fig. 112. P. Adoxae.
Teleutospores.

GU. 1l

162 PUCCINIA

rarely observed. I have pointed out elsewhere that it is on spring
flowering plants of this kind that Micropucciniz would naturally arise (as
well as on alpine plants), if they are the result of an adaptation to a
shortened vegetative period. Uromyces Ficariae may be taken as another
instance of a similar character.

85. Puccinia albescens Plowr.

Aicidium albescens Grev. Fl. Edin. p. 444. Cooke, Handb. p. 536 ;
Micr. Fung. p. 194.

Puccinia albescens Plowr. Ured. p. 153. Fischer, Ured. Schweiz,
p. 144, f. 110.

P. Adoxae Hedw.; Cooke, Micr. Fung. p. 209 p.p. Sace. Syll. vii.
612 p.p. Sydow, Monogr. i. 203 p.p.

Spermogones. Scattered among the ecidia, yellowish.

Aicidiospores. Scattered uniformly over the whole surface
of the leaves, also on the petioles, shortly cylindrical, whitish-
yellow, with a deeply cut revolute margin; spores finely warted,
pale-yellowish, 15—22 pw.

Uredospores. Sori minute, scattered singly or in little
clusters, soon naked, brown; spores globose to ellipsoid, echinu-
late, pale-brown, 21—28 x 18—25 yw, with two germ-pores.

Teleutospores. Sori similar, or teleutospores at first in the

same sori as the uredospores ; spores

= ellipsoid to subfusiform, rounded or
[\ attenuated above, with a conspicuous
colourless papilla, usually more or
less rounded below, hardly con-

i

stricted, smooth, chestnut- brown,
Fig. 113. P. albescens. 32—45 x 1425; pedicels hyaline,
Teleutosperss. delicate, short, deciduous.

On Adoxa Moschatellina. The xcidia appear in April or
even in March, uredo- and teleutospores in May and June.
Apparently not uncommon in the ecidium stage, which makes
the affected leaves paler and dwarfed; but see the following
paragraphs. (Fig. 113.)

The uredo- and teleutospores seem to be rare in a natural state,
although Schréter, Nielsen, Soppitt and Fischer have all produced them
in small quantity artificially from the wcidiospores; the latter laid affected

ON RUBIACEE 163

leaves on healthy plants on the 26th of April, removing them after a time;
nevertheless the experimental plants showed both uredo- and teleutospores
on May 16th. Their mycelium is localised, but that of the ecidium
permeates the whole plant; it is a disputed point whether it perennates
in the rhizome or not; Plowright affirms, Fischer denies this, and Bubék
thinks it probably not perennial (Centralbl. fiir Bakt. 2. xvi. 150).
Fischer records and figures abnormal three- or four-celled teleutospores.

The distinctions between this species and P. Adoxae lie not only in
the presence of the ecidium and uredo, but also in the appearance
and character of the teleuto-sori. In P. albescens these are widely
scattered and mostly single, and only follow the ecidium towards the end
of May—in P. Adowae they are crowded in larger groups, on more or less
deformed parts of the plant, and can be found as early as April or even
March; moreover there are no uredospores in them.

An ecidium on Adoxa is found in North America, but it has been
said (Dietel, 1895) that teleutospores have only once been seen there on
that host. On that account Dietel considers that the xcidium is able to
reproduce its own spore-form. This is contradicted by Bubdk’s experi-
ments and, since P. argentata is common in North America, it is more
probable that the ecidium on Adoxa found there is not Medium
albescens, but belongs to P. argentata. Bubdk, in fact, considers that this
is the case with most of the records of P. albescens. He says that the
zcidium of P. argentata has golden-yellow spores, and that he has met
with the true Meidium albescens only from Yorkshire and Baden. I do
not quite share this view: I have found teleutospores, apparently not
mixed with uredospores, in mid-April, on the same plant on which were
abundant ecidia with golden-yellow spores. There are three possible
explanations of this occurrence: either (1) it was P. albescens with deep-
coloured spores, or (2) there were on the same plant P. Adoxae and the
eecidium of P. argentata (which is a very rare species in Britain), or (3) all
the distinctions usually given must be upset. Which is the true explana-
tion, future experiments alone can decide.

36. Puccinia Asperule-odorate Wurth.

Puceinia Asperulae Fckl. Symb. Mye. p. 56 p.p.

P. punctata Lk. ; Sydow, Monogr. i. 213 p.p.

P. Galit Plowr. Ured. p. 143 p.p. Fischer, Ured. Schweiz, p. 332,
f. 246 (see ibid. p. 555).

P. Asperulae-odoratae Wurth, Centralbl. f. Bakter. 2. xiv. 314,
f. 9—10.

Acidiospores. AXcidia hypophyllous, in little clusters, sur-
rounded by a paler zone, cup-shaped, with a slightly projecting
11—2

164 PUCCINIA

white margin; spores with a thin, colourless, faintly warted
membrane, 14—21 pu.

Uredospores. Sori mostly hypophyllous, roundish, scattered
or in minute clusters, on the
stems linear, soon naked, clear
chocolate-brown; spores globose
to pyriform, brown, finely echinu-
late, 18—30 pn, with two mostly
equatorial germ-pores.

Teleutospores. Sori similar, but

long covered by the epidermis;

eid, desea spores ellipsoid to clavate, rounded

Teleutospores and uredospore. or conical above and thickened (up

to 7 or more), distinctly con-

stricted, tapering below, smooth, clear brown, 30—52 x 17—21 p;
pedicels short.

On Asperula odorata. Rare. (Fig. 114.)

Distinguished by Wurth from P. punctata on the ground of cultures:
the teleutospores have, he says, a less thick cap, which is pale at the apex,
and the uredospores have only two germ-pores. Description mainly after
Fischer and Wurth, and not agreeing exactly with my specimens; the
species is hardly distinguishable, however, from P. punctuta. Fischer
figures also three-celled teleutospores.

37. Puccinia punctata Link.

eidium Gaulii A. et S. (non Pers.) ; Cooke, Handb. p. 540; Mier.
Fung. p. 196, pl. 2, f. 15—17.

Trichobasis Galit Lév. ; Cooke, Micr. Fung. p. 226 p.p.

Puccinia punctata Link, Obs. Mye. ii. 30 (1816). Sydow, Monogr. i.
213 p.p.

P. Galti Schw. Syn. Fung. Carol. p. 73 (1822). Plowr. Ured. p. 143
p-p. Sacc. Syll. vii. 600. Fischer, Ured. Schweiz, pp. 332, 554,
f. 2438—5.

P. Galiorum Link, Sp. Pl. ii. 76 (1825). Cooke, Handb. p. 501; Mier.
Fung. p. 208, pl. 8, f. 172—3 p.p.

Spermogones. Amphigenous, in little clusters, honey-
coloured,

ON RUBIACEE 165

4icidiospores. AKcidia hypophyllous, scattered or collected
in orbicular clusters on roundish or oblong paler spots, shortly
cylindrical, with a short white recurved margin; spores
globose or broadly ellipsoid, orange-yellow, somewhat smooth,
16—23 p.

Uredospores. Sori hypophyllous, minute, roundish, on the
stems linear, reddish-brown, often confluent; spores globose to
ovate, aculeolate, pale-brown, 22—30 x 17—23 u, with two or
three germ-pores.

Fig. 115. P. punctata. Teleutospores; a, on G. saxatile; b, on G. palustre ;
ce, on G. verum.

Teleutospores. Sori similar, but black, compact, and per-
sistent; spores ellipsoid to clavate, truncate, rounded or coni-
cally attenuate above, where they are much thickened (up to
14), and often darker, slightly constricted, tapering below,
brown, smooth, 30—56 x 14-244; pedicels brownish above,
persistent, thick, about as long as the spore; a few mesospores
occasionally.

On leaves and stems of Galiuwm Cruciata (?), G. Mollugo,
G. palustre, G. saxatile, G. uliginosum, G. verum. Not uncom-
mon. Atcidia, June; teleutospores, August and September.
(Fig. 115.)

The forms of P. punctata on these various species of Galium differ
considerably in the shape and size of the teleutospores, and have been
divided into several biological races. On G. Aparine there is a distinct
species, P. difformis (q.v.); but it is possible that P. punctata also occurs

166 PUCCINIA

on that plant. I am not certain that P. punctata lives on G. Cruciuta,
though I think I have seen it on that host ; the two usual species on that
Galium are P. Celukovskyana and P. Valantiae (q.v.).

The zcidium seems to be less common than the other spore-forms :
I have seen it on @. Mollugo, G. verum and G. uliginosum. Mesospores
have been observed on G. palustre, G. saxatile and G. verum. On
G. saxatedle the teleutospores were slightly larger than in the normal form,
but otherwise identical; P. Valantiae, which occurs on that same host, is
easily distinguishable.

DistRiBuTion : Europe, Siberia, North America, Chili.

38. Puccinia Celakovskyana Bubdak.

Puccinia Celakovskyana Buba&k, Ber. Béhm. Ges. Wissen. Prag, 1898,
p. 11. Wurth, Centralbl. f. Bakt. pt. 2, xiv. 212, 310, f. 2—8.
Sacc. Syll. xvi. 287. Sydow, i. 216. Fischer, Ured. Schweiz,
p. 335, 555, f 247.

Spermogones. Amphigenous, dark honey-coloured.
Uredospores. Primary sori surrounding the spermogones
and on the same mycelium,
more or less circinate, sur-
rounded by a yellowish zone,
long covered by the epidermis,
dark-brown; secondary, am-
phigenous, minute, scattered,
soon naked, pale chestnut-
brown or pale cinnamon; spores
globose to obovate, distinctly
echinulate, brown, 21—33 x 18
Fig. 116. P. Celakovskyana. ae a nearly a al
Teleutospores and uredospore. torial germ-pores.
Teleutospores. Sori hypo-
phyllous, on brownish spots, thickly scattered, minute, roundish,
on the stems linear and up to 1 mm. long, soon naked, pulvinate,
black-brown ; spores oblong to clavate, rounded (very seldom
narrowed) above where they are much thickened (up to 11)
and darker, faintly constricted, tapering below, smooth, chestnut-
brown, 85—56 x 17—25 w; pedicels hyaline, thick, persistent,
not as long as the spore; an occasional mesospore is found.

ON RUBIACEE 167

On Galium Cruciata. Not uncommon. Uredospores, May
—July; teleutospores, August, September. (Fig. 116.)

Distinguished at once from P. Valantiae on the same host by the dark
colour of its teleutospores, and their very great and dark apical thickening.
The presence of uredospores also distinguishes them ; in P. Celakovskyana
the two kinds of sori are often present together on the same leaf, the
uredo-sori pale brown, and the teleuto-sori almost black.

According to Bubak, almost its only distinction from P. punctata lies
in the absence of the zcidium whose place is taken by the primary uredo-
sori. Wurth reports it also (J.c.) on G. pedemontanum on the continent,
but demonstrated by culture-experiments that it could not be transferred
to other species of Galiwm. Most of the records of P. punctata on
G. Cruciata probably belong to this species, which certainly shows few
morphological distinctions from the former; [ find, however, that the
teleuto-sori are larger, more numerous, and more compact in this species
than in that found on G@. palustre.

Distrigutrion: Central Europe.

39. Puccinia Valantiz Pers.

Puccinia Valantiae Pers. Obs. Myc. ii. 25. Cooke, Handb. p. 500;
Micr. Fung. p. 207. Plowr. Ured. p. 212. Sace. Syll. vil. 685,
Sydow, Monogr. i. 217. Fischer, Ured. Schweiz, p. 336, f. 248,
P. acuminata Fckl. ; Cooke, Micr. Fung. p. 208.
Teleutospores. Sori hypophyllous, rather thick, pulvinate,
compact, orbicular, scattered, circinate
or confluent, up to 2 mm. diam., at first La
yellowish-,then chestnut-brown,atlength
greyish-brown (from the spores germi- ~
nating in situ), often elongated and
causing distortion on the stems; spores \°]
fusoid or slightly oblong, attenuated at
both ends, thickened (up to 9 «) above, a b
somewhat constricted, smooth, pallid-

‘brown, 35—65 x 10—17 pw; pedicels hya- pi, 117, p. valantiae.

line, persistent, up to 80 u long. Teleutospores; a, on
. : G. Cruciata; b, on
On Galium Cruciata, G. sazatile. G. saxatile.

June—September. Common. (Fig. 117.)

This Leptopuccinia is very distinct from P. Celakovskyana, which also
lives on G. Cruciata, not only by the absence of uredospores, but also by

168 PUCCINIA

the fusiform shape, thin walls, and pale colour of the teleutospores, which
often become totally devoid of thickening at the apex, by the dropping off
of the pale thickening cap, on germination. P. punctata is additionally
distinguished by the presence of the ecidium.

Fischer explains the fact, that distortions more usually accompany
P. Valantiae, by the consideration that infection takes place from the
basidiospores mainly through the cuticle of young and still growing parts,
while teleuto-sori of P. punctata and P. Celakovskyana are produced by
infection from spores whose germ-tubes can penetrate the stomata of
parts of the plant which are already fully developed. The relations
between the three species are very like those which subsist between
Oromyces Trifolit-repentis, U. Trifolii, and U. flectens.

DisTRIBUTION: Europe, North America.

40. Puccinia difformis K. et S.

Puceinia difformis K. et S. Myk. Heft. i. 71 (1817). Cooke, Handb,
p- 501; Micr. Fung. p. 208.

P. Galit Plowr. Ured. p. 144 p.p.

P. ambigua Lagh. in Sydow, Uredineen, no. 1056 (1897). Sace. Syll.
xvi. 288. Sydow, Monogr. i. 216.

4icidiospores. Aicidia hypophyllous, on yellow spots, soli-
tary or irregularly disposed over the
whole leaf, whitish-yellow, with torn
reflexed margin; spores verruculose,
orange, 13—25 p.

Teleutospores. Sori hypophyllous
or on the stems, small, elliptic, soli-
tary or clustered, on the stems often
elongated and confluent, long covered
by the ash-coloured epidermis, then
naked, firm, black ; spores ellipsoid to

Fig. 118. P. difformis, clavate, much thickened above, hardly

Teleutospores. constricted, tapering below, smooth,

brown above, paler downwards, 35—

55 x 15—25 4; pedicels brownish, persistent, as long as the
spore or longer.

On Galium Aparine. July—August. Surrey, Kent. (F ig.
118.)

ON VERONICA 169

The ecidia may be found together with the teleutospores right up to
September, often on the same spot. This species is very different from
the others on Galiwm; the teleuto-sori, as Cooke says, are “firm and
compact like little spots of pitch,” and may be accompanied by swellings
and distortion. There are no uredospores.

DISTRIBUTION: Central and Northern Europe, India, North
America.

41. Puccinia Veronice Schrit.

P. Veronicae Schrét. Pilz. Schles. p. 347. Plowr. Ured. p. 211 p.p.
Sydow, Monog. i. 256. Fischer, Ured. Schweiz, p. 323, f. 235.

Teleutospores. Sori hypophyllous, on orbicular brown spots,
minute, scattered or circinate,
roundish, at first yellowish-
brown, then brown; spores
fusoid, generally rounded above
and thickened (up to 5 p),
hardly constricted, tapering
below, smooth, yellowish or
very pale brown, 28—52 x
10—16 », pedicels hyaline,
persistent, as long as the spore Fig. reece aes
or shorter.

On Veronica montana. June (or earlier) to October. Not
common. (Fig. 119.)

Distinguished from P. Veronicarum by the very clear-coloured and
narrower spores, which are less strongly thickened at the apex. It has
also only one form of spore, viz. that with persistent pedicel which
germinates as soon as mature. It does not occur on V. alpona.

DisTRIBUTION : Western and Central Europe.

42. Puccinia Veronicarum DC.

P. Veronicarum DC. Flor. fr. ii. 594. Cooke, Handb. p. 496; Micr.
Fung. p. 204. Plowr. Ured. p. 214. Sydow, Monog. i. 257.
Fischer, Ured. Schweiz, p. 323, f. 236. See Sacc. Syll. vii. 685.

Teleutospores. Sori hypophyllous, on irregular or roundish

brown spots, circinate, minute, roundish, often confluent, some

170 PUCCINIA

compact, pulvinate and greyish, others brown and pulverulent ;
a spores oblong or obovate, attenu-
ated into a horny much thickened

(up to 8) conical point at the

apex, constricted, slightly tapering

C) below, smooth, pale or dark chest-
nut-brown, 28—40 x 1419 p;
pedicels hyaline or somewhat yel-

ee ee een lowish, rather long, deciduous or

leutospores on V. alpina (ex persistent.
herb. Berkeley).

On Veronica alpina, V. offici-
nalis and perhaps other species. Not common. July—October.
(Fig. 120.)

There are two forms of spores in this species; one, fragzlipes, with
deciduous pedicels ; the other, persistens, with persistent pedicels. In the
former, the sori are soon naked and pulverulent, the spore-wall is thicker
and darker-coloured, and germination takes place only after a winter's
rest. In the latter, the sori are compact, the spores are thin-walled and
paler and germinate as soon as mature, upon the living plant. But
occasionally both forms may be seen in the same sorus. It is the latter
form that spreads the parasite during the summer; the former causes
new infections in the following spring.

Distinguished from P. Veronicae by the relatively broader spores, and
the cone-shaped apical thickening. It is possible that the Pucezniae on
other species of Veronica, included by the Sydows with this, are really
distinct. But in Herb. Berkeley there is a Puccinia on Veronica alpina,
from Ben Aulder, Invernessshire (and also from Perthshire) which is
undoubtedly this species. The sori are hypophyllous only, but cover the
larger part of each leaf. The spores are quite smooth, by which it is
distinguished from the continental form on V. alpina, which has the upper
spore-cell occasionally warted on the upper part ; this form (P. albulensis
Magn.) has the sori densely gregarious on stems as well as leaves, some-
times covering a whole internode.

DISTRIBUTION: Europe generally.

43. Puccinia Menthe Pers.

Micidium Menthae DC. Flor. fr. vi. 95. Cooke, Handb. p. 544; Mier.
Fung. p. 199.

Trichobasis Labiatarum Lévy. ; Cooke, Micr. Fung. p. 224 p.p.

T. Clinopodti DO. ; Cooke, Micr. Fung. p. 224 (?).

ON LABIATA 171

Puccinia Menthae Pers. Syn. p. 227. Cooke, Handb. p. 496; Micr.
Fung. p. 204 p.p., pl. 4, f. 69, 70. Plowr. Ured. p. 157. Sace.
Syll. vii. 617. Sydow, Monogr. i. 282. Fischer, Ured. Schweiz,
p. 168, f. 131. McAlpine, Rusts of Australia, p. 140, f. 250.

P. Clinopodti DC. ; Cooke, Micr. Fung. p. 205 (?).

Spermogones. Scattered or arranged in little groups, honey-
coloured.

4icidiospores. Aicidia hypophyllous and often on the
stems, arranged on the leaves in clusters on orange or purplish
spots, or forming elongated patches on the stems and petioles
which are much thickened and deformed, opening irregularly,
margin scarcely torn, erect or even incurved ; spores verruculose,
pallid-yellow, 24—40 x 17—28 pu.

Uredospores. Sori hypophyllous, on yellowish or brownish
spots (or without spots), minute, roundish, scattered or aggre-
gated, soon naked, surrounded by the ruptured epidermis,
sometimes confluent, cinnamon; spores globose
to obovate, echinulate, pallid-brown, 17—28 x
14—19 p, with three equatorial germ-pores.

Teleutospores. Sori similar, but dark-brown;
spores subglobose to obovate, rounded at both
ends, with a broad pale-coloured apical papilla,
not or scarcely constricted, more or less in-
distinctly verruculose, sometimes smooth, dark-
brown, 26—35 x 19—23; pedicels hyaline, Fig. 121.

slender, longer than the spore. P. Menthae.

. : , Teleutospore
On Mentha aquatica, M. arvensis, M. citrata, on M. aquatica.

M. rotundifolia, M. silvestris, M. viridis, Ori-

ganum vulgare, Calamintha Clinopodium, C. officinalis. May—
October, teleutospores from August. Very common on garden
Mint, rather common on some of the other species. (Fig. 121.)

There can be little doubt that this is a collective species. Points of
difference are found in the finer or coarser warts of the teleutospores and
in the length of the pedicel ; but hitherto no certainty has been arrived at
in delimiting the various forms. I have seen the warts quite distinctly on
some of the teleutospores on V. aquatica and M. arvensis, when they are
viewed dry, but other spores in the same sorus seemed perfectly smooth.
The xcidium seems not.to occur on all species, though it is common on

172 PUCCINIA

garden mint; perhaps those forms which are without it may hereafter be
separated. But it has occurred on all the hosts mentioned above except
Origanum and WM. rotundifolia ; it may, however, according to Sydow, be
merely facultative. The form on C. Clinopodium really shows less difference
from that on Mentha aquatica than those on species of Mentha do from one
another. But Cruchet was unable to infect any one of the four Jf. arvensis,
M. aquatica, WV. silvestris, C. Clinopodium, except by spores from the same
species. As the result of his experiments, he divides P. Menthae into
eight biological races, as it occurs on Mentha and Calamintha ; and the
form on Origanwm is also biologically distinct. The Australian form of
P. Menthae, which is an introduced species on UM. Pulegium and M. lax-
flora, has no known ecidiospores, but occasional mesospores. Nothing
seems to be known about the form on Ajuga reptans mentioned by
Plowright, from Johnston’s Flor. Berwick.

In garden mint (J/. viridis) the mycelium of the ecidial stage is spread
throughout the whole plant, even in the rhizome; Klebahn was able to
trace the hyphe in some cases nearly up to the growing point. It lasts
for several years at least ; a bed of mint infested with it should be rooted
up and burnt ; there is no cure for the disease, although I have found that
cuttings taken from some of the more distant healthy-looking shoots and
planted elsewhere grow up without the parasite. The mycelium of the two
other stages is purely local. I have known the ecidia to occur for several
years in a garden without being followed by uredo- or teleutospores so far
as could be seen, and vice-versd, in another case, these spores occurred but
no ecidium was ever noticed.

DisTRIBuTION: Europe, Asia, Africa; the American and
Australian teleutospores are-more strongly warted.

44. Puccinia caulincola Schneid.

Puceinia caulincola Schneid. in Jahresb. Schles. Gesell. 1870, p. 120.
Sydow, Monogr. i. 301. Fischer, Ured. Schweiz, p. 172, f. 133.

P. Schneideri Schrét. in Herb. Schles. Pilze, no. 448. Plowr. Ured.
p. 201. Sacc. Syll. vii. 677.

Teleutospores. Sori on the stems and petioles, rarely on
the leaves, scattered, occasionally confluent, minute, roundish
or elongated, long covered by the bullate epidermis, at length
pulverulent, black, then cinnamon-brown; spores ellipsoid,
rounded at both ends, apex sometimes thickened in a papilli-
form fashion, rather constricted, smooth, pale-brown, 24—33
x 15—24; pedicels hyaline, thin, rather long, not very

ON LABIATA 173

deciduous ; a few mesospores intermixed, each with a low broad
pore-cap.

On Thymus Serpyllum. June— :
October. Very rare; links, Aber- () C)
deen (Prof. J. W. H. Trail). (Fig.
122.) wo) C)

The presence of the mycelium causes
the stems to stand more upright; the
internodes are considerably lengthened, and
the leaves fewer, so that the affected plants
can be readily distinguished, as in P.
Betonicae. The mycelium appears to be Fig. 122. P. caulincola.
perennial, and the sori to be confined Teleutospores and mesospore.
almost entirely to the stems, where they

cause a slight thickening and are more frequent at the nodes than
elsewhere.

DISTRIBUTION : Central and North-western Europe.

45. Puccinia Glechomatis DC.

P. Glechomatis DC. Encycel. viii. 245. Cooke, Handb. p. 496; Micr.
Fung. p. 204, pl. 4, f 73—4. Plowr. Ured. p. 214. Sacc. Syll. vii.
688. Sydow, Monogr. i. 277. Fischer, Ured. Schweiz, p. 327,
f. 239.

Teleutospores. Sori hypophyllous or on the petioles, on
brownish spots or sometimes none,
3—1 mm. diam., roundish, solitary
and scattered, or more often subcon-
fluent into rounded clusters as much
as 4 mm. diam., on the stem and
petioles often elongated, pulvinate,
at first yellowish, then chestnut, and
at last blackish; spores ellipsoid or
oblong, with an acute or rounded
horn-like process (8-12 high) which
is often obliquely placed and falls off on germination, faintly
constricted, rounded below, smooth, pale and clear-brown,
30—48 x 15—24 u; pedicels hyaline, persistent, as much as
75 long; an occasional mesospore is found.

Cj >

Fig. 123. P. Glechomatis.
Teleutospores and mesospore.

174 PUCCINIA

On leaves, petioles and stems of Glechoma hederacea
(Nepeta Glechoma). June—October. Not uncommon. (Fig.
123.)

The sori are especially large, round and compact late in the season,
when they produce spores which are darker and will not germinate
immediately (as the others do), but only after the winter’s rest. I havea
specimen, resembling this species, on Prunella vulgaris from Sutton Park,
Warwicks. ; Plowright mentions a similar one from Ben Lawers (i.c. p. 215).

DISTRIBUTION : Europe, Siberia, Japan.

46. Puccinia Betonice DC.

Puccinia Anemones var. Betonicae A. et S. Consp. p. 131.

P. Betonicae DC. Flor. fr. vi. 57. Cooke, Handb. p. 497; Micr. Fung.
p. 205. Grove, in Gard. Chron. xxiv. 180, f. 38 (1885). Plowr.
Ured. p. 199. Sacc. Syll. vii. 677 p.p. Sydow, Monogr. i. 275.
Fischer, Ured. Schweiz, p. 173, f. 134.

Teleutospores. Sori hypophyllous, on pallid irregular spots,
numerous, aggregated in patches, or more generally spreading
over nearly the whole of a leaf, more or less crowded on the
nerves, minute, perfectly round, surrounded by the torn erect
epidermis, pulverulent, reddish-brown; spores ellipsoid to

9ESQ0

Fig. 124. P. Betonicae. Normal and abnormal teleutospores.

ovate, rounded above with a small paler hemispherical pore-
cap, slightly constricted, rounded below, smooth, yellow-brown,
27—45 x 15—24 w; pedicels thin, hyaline, deciduous, about as
long as the spore; a few oval mesospores intermixed.

On Betonica officinalis (Stachys Betonica). Not common.
May—September. (Fig. 124.)

The affected leaves are paler, narrower, and stand more erect than the
healthy ones. Besides the mesospores, other anomalies are occasionally

ON LABIATAE 175

met with, viz. spores with three or more cells variously arranged. See
Grove, Gardener’s Chronicle, xxiv (1885), p. 180, f. 38. The mycelium is
probably perennial.

DIsTRIBUTION: Central and Western Europe.

47, Puccinia annularis Schlecht.

Uredo annularis Strauss in Wetter. Ann. ii. 106.
Puceinia annularis Schlecht. Flor. Berol. ii. 132 (1824). Plowr. Ured.

p. 217. Sace. Syll. vii. 689. Sydow, Monogr. i. 300. Fischer,
Ured. Schweiz, p. 329, f. 240.

P. Scorodoniae Link. Spec. ii. 72 (1825). Cooke, Handb. p. 497;
Micr. Fung. p. 205.

Teleutospores. Sori hypophyllous, on indefinite yellowish or
brownish concave spots, at first minute,
roundish, covered by the epidermis, in
orbicular clusters, then naked, con- bee
fluent, and forming a thick pulvinate
mass, ferruginous-brown; spores ob- \)
long, rounded or attenuated at the
apex and much thickened (up to 8 1),
slightly constricted, rounded or attenu-
ated at the base, smooth, very pale
pellpeel Rrowe, G0 ot <i at Ms Fig. 125. P. annularis.
pedicels hyaline, persistent, as much Teleutospores.
as 80 long.

On Teucrium Scorodonia. July—October. Not common.
(Fig. 125.)

Here, as in other Lepto-species, there seem to be two kinds of spores,
those which germinate at once and those which will not germinate till the
following year. The form on 7. Chamaedrys (not yet found in Britain)
has been proved to be biologically distinct.

DISTRIBUTION : Central and Southern Europe.

48. Puccinia Convolvuli Cast.

Uredo Betae 8. Convolvuli Pers. Syn. p. 221.

Puccinia Convolvuli Cast. Obs. 1.16. Plowr. Ured. p. 146. Sace. Syll.
vii. 610. Sydow, Monogr. i. 319. Fischer, Ured. Schweiz, p. 322,
f, 234.

176 PUCCINIA

[4cidiospores. Acidia hypophyllous, on brownish or pur-
plish spots, more or less circinate, often on the petioles and
then in elongated patches, cup-shaped, minute, with broad
recurved torn white margin; spores delicately verruculose,
pallid-yellow, 17—28 y.]

Uredospores. Sori scattered or circinate, minute, often
confluent, soon naked, brown; spores more or less ellipsoid,
rarely ovate, faintly echinulate, pale-brown, 22—30 x 18—26 p,
with two or three germ-pores just above the middle.

Teleutospores. Sori similar, but long
covered by the grey epidermis, black-
brown; spores ellipsoid to oblong, ob-
tuse or rounded above, more or less
thickened (up to 9 y), gently constricted,
rounded below, smooth, chestnut-brown,
38—66 x 18—30 4; with them are
intermixed (according to Sydow) ovoid
mesospores, much thickened at the

Fig. 126. P. Convolwuli. apex, brown, 25—35 x 20—26 uw; pedi-
Teleutospores on Convol- 681. brownish, thick, persistent, up to

vulus sepium, Poughkeep-
sie, U.S.A. 35 Be long.

On Convolvulus sepium (Miss Jelly). June—October. Very
rare. (Fig. 126.)

The connection of the ecidiospores with the teleutospores was experi-
mentally demonstrated by Arthur. According to Fischer, the uredospores
often have a smooth median equatorial zone, of which I could see no trace.
I have not seen the ecidia.

DistRipuTIoN: Europe, Africa, Japan, North America.

49. Puccinia Vince Berk.

Uredo Vincae DC. Flor. fr. vi. 70.

Trichobasis Vincae Berk. ; Cooke, Micr. Fung. p. 226, pl. 6, f. 130—1.

Puccinia Vineae Berk. Engl. Fl. v. 364. Cooke, Handb. p. 497 ; Mier.
Fung. p. 205, pl. 6, f. 182. Plowr. Ured. p. 161, pl. 2, f£. 11—14;
Gard. Chron. 1885, xxiv. 108, f. 22—3. Sacc. Syll. ix. 310.
Sydow, Monogr. i. 338. Fischer, Ured. Schweiz, p. 167, f. 130.

Spermogones. Hypophyllous or amphigenous, minute,

ON VINCA 177

punctiform, brownish, often very numerous, scattered over the
whole leaf-surface, sweet-scented, spherical, about 175 » diam.

Uredospores. Sori hypophyllous, pallid-brown, of two kinds ;
primary irregular, often elongated and curved, crowded and
confluent, naked; secondary scattered, on roundish dirty-brown
spots, long covered by the epidermis; spores globose to pyriform,
aculeate, pallid-brown, 20—32m diam. or 20—46 x 16—24 p,
with three germ-pores.

Fig. 127. P. Vineae. wu, teleutospore, seen dry ; b, the same, seen wet;
c, uredospore; d, the so-called ecidia. On Vinca major, all x 600.

Teleutospores. Sori hypophyllous, on scarcely perceptible
or conspicuous spots, minute, scattered or in groups, roundish
or irregular, surrounded or half-covered by the torn epidermis,.
pulverulent, dark-brown ; spores ellipsoid to oblong, rounded at
both ends or attenuated downwards, hardly thickened at apex
but with a pale papilla, not at all or faintly constricted, scrobi-
culate, ochreous-brown, 35—54 x 18—27 4; epispore 3—4 4
thick ; pedicels hyaline, deciduous, rather long.

On Vinca major, V. minor. Not common. Spermogones in
April; uredospores, May—June; teleutospores, July—October.
(Fig. 127.)

This is one of the most remarkable species of Puccinia found in Britain.
There is considerable difference of opinion about its structure. The bodies
referred to in the description given by Plowright (/.c. p. 161) as “secidia” are
of a puzzling nature: they are not described by Sydow or Fischer, but are
mistakenly considered by them as identical with the primary uredo-sori.
They accompany the spermogones on the under side of the leaves, and
are flatly pulvinate subepidermal erumpent sori, surrounded by the erect

G. U. 12

178 PUCCINIA

epidermis, and consisting of a floor of crowded erect narrow brownish
hyphe, each of which abstricts from its apex a small, smooth, round,
thick-walled, nearly colourless spore, 8—10y diam. Plowright describes
and figures these spores as forming basipetal chains, but this I could not
see. The sori are numerous, about 1mm, diam. and dark-brown with a
greyish bloom, due apparently to the overlying spores. Can they be a
parasitic fungus like Darluca Filum, which has itself occasionally been
considered as an additional spore-form of the Uredine on which it preyed ?

The plants affected by the mycelium of the spermogones are permeated
by it; they grow taller and more erect, the internodes are longer, the
leaves paler, shorter, and thicker. Plowright considered this mycelium to
be perennial, which is very probable. The spermogones have a distinct,
but faint odour.

The sculpture of the teleutospores is very striking and almost unique.
According to Sydow, it consists of a network of warts, but, in the British
specimens which I have seen, it would be more correctly described as a
series of pits arranged more or less in longitudinal lines (about 12 across
_ thespore). Fischer represents the network as much finer. As is frequently

the case, these markings can only be seen properly on the dry spore ; they
vanish almost completely when it is placed in water, unless it is emptied
of its contents. The germ-pore of the lower cell is placed near the insertion
of the pedicel, a very unusual position ; it is covered with a pale papilla
like the upper one.

DisTRIBUTION: Central and South-western Europe.

50. Puccinia Gentiane Link.

Medium Gentianae Jacz. Champ. Mont. p. 163 (1892), but ?

Oredo Gentianae Strauss in Wetter, Annal.'ii, 102 (1811).

Puceinia Gentianae Link, Spec. ii. 73. W. G. Smith, Gard. Chron.
xxlv. (1885) p. 372, f. 82. Cooke, Grevillea, xiv. 39. Plowr. Ured.
p. 147. Sace. Syll. vii. 604. Sydow, Monogr. i. 340. Fischer,
Ured. Schweiz, p. 164, f. 126.

[Spermogones. Honey-coloured, in small clusters.

dicidiospores. Aicidia hypophyllous or on the stems, on
circular brown spots, in irregular clusters, cup-shaped, with torn
white margin; spores delicately verruculose, orange, 16—23 x
1417 yp]

Uredospores. Sori amphigenous, but oftener on the upper
surface, scattered or circinate, minute, roundish, at first covered
by the epidermis, pale-chestnut; spores globose to ovoid,

ON GENTIANA 179

aculeolate, brownish-yellow, 20—80 x 18—24 py, with two,
rarely three, germ-pores.

Teleutospores. Sori similar and also on the stems, but
pulverulent and black-brown ;
spores ellipsoid to ovoid,
rounded at both ends, not
thickened above, but some-
times with a low broad papilla,
not constricted, smooth, dark-
chestnut, 28—38 x 24—30 w; Fig. 128. P. Gentianae. Teleutospore,
pedicels hyaline, thin, rather feawih ee pag
long, very deciduous; occa-
sionally there are a few mesospores intermixed.

On Gentiuna acaulis. Uredo- and teleutospores, Kew
Gardens, August, 1905 (M. C. Cooke), and Horsham (W. G.
Smith). (Fig. 128.)

I have a specimen on G. Andrewsii, from the United States, with
exactly similar teleutospores. The ecidia are said to appear in April and
June, and more frequently on the stems and peduncles than on the leaves.
Jaczewski’s AZicidium was on calyx, stem and leaf of G. angustifolia, on
pale orange spots.

DisTRIBUTION: Europe, Siberia, East Indies, North
America.

51. Puccinia Primule Duby.

Aicidium Primulae DC. Flor. fr. vi. 90. Cooke, Handb. p. 544;
Mier. Fung. p. 199.

Oredo Primulae DC. Flor. fr. vi. 68.

Trichobasis Primulae Cooke, Micr. Fung. p. 227.

Puccinia Primulae Duby, Bot. Gall. ii. 891. Cooke, Handb. p. 495 ;
Micr. Fung. p. 204. Plowr. Ured. p. 159. Sace. Syll. vii.
612. Sydow, Monogr. i. 348. Fischer, Ured. Schweiz, p. 161,
f. 124.

Aicidiospores. Aicidia hypophyllous, on yellowish spots,
densely but irregularly clustered in roundish groups, shortly
cylindrical, with a broad much cut revolute white margin ;
spores verruculose, orange, 17—23 x 12—18 p.

12—2

180 PUCCINIA

Uredospores. Sori hypophyllous, minute, scattered or
circinate, roundish, soon naked, brown; spores subglobose to
ovoid, echinulate, pallid-brown,

20—23 x 16—19 », with three

~ germ-pores.
Teleutospores. Sori similar,
y but long covered by the grey
epidermis, often confluent, or

; . standing in circles round the

Fig. 129. P. Primulae. a 2 ‘
Teleutospores and mesospore. ecidia or uredo-sori, blackish-
brown; spores ovoid or oblong,
rounded at both ends, hardly thickened above but with a
broad colourless papilla on each germ-pore, gently constricted,
smooth, pallid-brown, 22—30 x 15—18 uw; pedicels hyaline,

short, deciduous; an occasional mesospore was seen.

On Primula acaulis (vulgaris). Not common. Aicidia in
May; teleutospores, June—October. (Fig. 129.)

All three spore-forms may be found on the same leaf. The teleuto-
spores are rather irregular in shape ; mesospores are not infrequent, and

Fischer describes and figures three-celled spores. This parasite is recorded
on the continent also on Primula elatior and P. veris.

DISTRIBUTION: Central Europe.

52. Puccinia Soldanelle Fckl.

Aicidium Soldanellae Hornsch. in Rab, Krypt. Fl. p. 18 (1844).
Cooke, Handb. p. 537 ; Micr. Fung. p. 195.

Uredo Soldanellae DC. Flor. fr. vi. 85 (1815).

Puccinia Soldaneliae Fckl. Symb. Myc. Nacht. iii. 14. Plowr. Ured.
p. 159. Sace. Syll. vii. 618. Sydow, Monogr. i. 349. Fischer,
Ured. Schweiz, p. 159, f. 123.

Spermogones. Hypophyllous, numerous, punctiform, spheri-
cal.

eidiospores. ABcidia hypophyllous, scattered uniformly
over nearly the whole leaf-surface, shortly cylindrical or urceo-
late, with a white denticulate revolute margin ; spores delicately
verruculose, yellowish, 18—26 p.

[Uredospores. Sori generally epiphyllous, without. spots,

ON PRIMULACE 181

scattered or circinate, minute, surrounded by the torn epidermis,
brown; spores globose to ellipsoid, echinulate, pale-brown,
20—-32 x 18—28 w; epispore thick (2—3 yw), with three germ-
pores.

Teleutospores. Sori similar, but black-brown; spores ellipsoid
to ovate-oblong, with a broad paler apical papilla (5—8 pw high),
gently constricted, usually rounded below, smooth, brown, 35—
55 x 20—84 4; pedicels hyaline, deciduous, up to 50 » long.]

On Soldanella alpina (Botanic Garden, Glasgow). All three
spore-forms are said to occur together on the same plant in
July and August.

The zcidium stage is most common ; its mycelium is perennial and
diffused through the plant, and causes a conspicuous change in the leaves ;
they become smaller, paler, and longer-stalked. Only this stage seems
to have been met with in Britain ; no doubt on imported plants, as has
happened with P. Gentianae, P. Pazschket, etc. Description of the uredo-
and teleuto-stages after Sydow.

DIsTRIBUTION : Central Europe.

53. Puccinia Hydrocotyles Cooke.

Caeoma Hydrocotyles Link, Sp. Plant. ii. 22.

Trichobasis Hydrocotyles Cooke, Journ. Bot. ii. 844; Handb. p. 530;
Micr. Fung. p. 225, pl. 8, f. 168—9.

Puccinia Hydrocotyles Cooke, Grevillea, ix. 14. Ph. et Pl. Grevillea,
xiii. 53. Plowr. Ured. p. 195. Sacc. Syll. vii. 641. Sydow,
Monogr. i. 388.

[Hicidiospores. Acidia amphigenous, distributed pretty
uniformly over the whole leaf, rarely single, cup-shaped, with
a yellowish deeply cut revolute margin; spores punctate, pale-
yellowish, 19—26 y.]

Uredospores. Sori amphigenous, scattered, now and then
confluent, often circinate round a central larger one, very
minute, long covered by the epidermis, at length naked, pul-
verulent, cinnamon; spores subglobose or ellipsoid, echinulate,
brownish, 24—34 x 20—27 w, with two conspicuous germ-
pores.

182 PUCCINIA

Teleutospores. A very few are found in the uredo-sori,
ellipsoid to oblong, rounded at both
ends, hardly thickened above, gently
constricted, smooth, brown, 30—44 x 18
—28 yu; pedicels hyaline, thin, deciduous.

On Hydrocotyle vulgaris. Rare:
Kew Gardens; Epping Forest; Ireland,
Co. Dublin. Uredospores, July—Sep-
tember; teleutospores, October. (Fig.
130.)

This species is very imperfectly known.

The ecidium is recorded only from South

Fig. 180. P. Hydrocotyles. America ; in Europe the uredo-form alone has

Teleutospore and

uredospore. been observed, except for a few teleutospores

in the uredo-sori. Lindroth describes the

teleutospores (which are rare everywhere) as furnished with large isolated

depressed and rounded warts, while those I have seen are perfectly

smooth and with long and persistent pedicels. Cooke describes, both

uredo- and smooth teleutospores, intermixed, on Hydrocotyle, from Natal

(Grevillea Z.c.). Sydow states that the uredospores from all localities

agree perfectly ; those that I have examined from the Hawaian Islands

agree exactly with ours, having the same peculiar colour resembling a
strong wash of “raw sienna.”

This species illustrates, in the Epping Forest locality, what can be
frequently observed :—that, so long as the surroundings are undisturbed
by man, many species of Fungi occur year after year continuously in the
same spot. It is recorded as found there in 1863, 1864, 1871, 1882, and
1906, etc., and no doubt could have been found, or was found, there equally
in all the intervening years.

DIsTRIBUTION: France, Holland, Italy, North and South
America, Natal, Pacific Islands.

54. Puccinia Sanicule Grev.

Aeidium Saniculae Carm. in Cooke’s Brit. Aicid., Journ. Bot. ii. 39,
pl. 14, f. 1. Cooke, Handb. p. 543; Micr, Fung. p. 198.

Puceinia Saniculae Grev. Fl. Edin. p. 431. Cooke, Handb. p. 502 ;
Micr. Fung. p. 208, Plowr. Ured. p. 160. Sacc, Syll. vii. 618.
Sydow, Monogr. i. 413. Fischer, Ured. Schweiz, p. 122, f. 93.

Spermogones. Amphigenous, in little groups..
4Aicidiospores. AXcidia hypophyllous or on the petioles, on

ON UMBELLIFER 183

brown or purple spots, in small round clusters 2—4 mm. diam.,
elongated on the nerves and petioles, cup-shaped, with a whitish
lobed revolute margin; spores tending to become ellipsoid,
delicately verruculose, hyaline, 18—26 x 15—22 u.
Uredospores. Sori hypophyllous, on pale minute spots

2—3 mm. diam., scattered, rarely aggre- a

gated, minute, punctiform, pale-cinna-

mon; spore globose to ellipsoid, echinu- ( \)
late, not thickened above, brown, 25—- \ H
38 x 18—27 4; epispore thick (up to YD

34 4), with two (rarely three) germ-pores,

with slightly swollen caps. Fig. 181. P. Saniculae.
Teleutospores. Sori similar, but dar- eit cca iil

ker; spores ellipsoid or oblong, rounded

at both ends, sometimes slightly thickened above and provided

with a small papilla, hardly constricted, smooth, brown, 26—

45 x 18—26 w; pedicels hyaline, thin, deciduous.

On Sanicula europaea. Common. Aicidia, May and June;
teleutospores, August—October; but I have found all three
kinds of spores, in sheltered places, as early as April. (Fig. 181.)

DistRIBUTION: Central and North-western Europe.

55. Puccinia Cicute Lasch.

Puceinia Crcutae Lasch in Klotzsch, Herb. Myc. no. 787. Sydow,
Monogr. i. 372.

icidiospores. Aicidia on the nerves of the leaves, on
petioles and stems, in roundish or oblong groups as much as
ld em. long, pustular, with a feebly developed peridium, golden-
yellow; spores globose to ellipsoid, delicately punctate, sub-
hyaline, 17—26 x 10-—20 p.

Uredospores. Sori generally hypophyllous, scattered, mi-
nute, punctiform, pulverulent, cinnamon; spores subglobose to
ovate, echinulate, yellow-brown, 18—28 x 1422 yw, with three
germ-pores.

Telewtospores. Sori similar, but blackish-brown; spores
ellipsoid or oblong, rounded at both ends or rarely attenuated
downwards, not thickened above, gently constricted, somewhat

184 PUCCINIA

verruculose or distinctly verrucose-reticulated, even sometimes
nearly smooth, brown,, 28—46 x 18—30 p; pedicels hyaline,
thin, short, deciduous.

On Cicuta virosa. Rare. Mentioned by Plowright in his
arrangement of the British species, Trans. Brit. Myc. Soe. 11. 26;
he assigns spermogones to it as well. I have seen no specimens.

The fungus is similar to P. Smyrnii in its ecidia and teleutospores,
but differs in the presence of uredospores. Description after Sydow.

DISTRIBUTION: Central and Northern Europe, Siberia,
Japan, North America.

56. Puccinia Apii Desm.

Trichobasis Api Wallr. ; Cooke, Micr. Fung. p. 224.

Puccinia Apii Desm. Cat. des Pl. omis. p. 25. Cooke, Handb. p. 502;
Micr. Fung. p. 208. Plowr. Ured. p. 156. Sydow, Monogr. i. 359.
Fischer, Ured. Schweiz, p. 119.

Spermogones. Hypophyllous, mostly surrounded by the
eecidia, often circinate, shining reddish-brown.

Aicidiospores. Aicidia hypophyllous or on the petioles, on
minute, irregular, conspicuous, yellowish spots, in roundish or on
the petioles elongated clusters, shortly cylindrical, with white
torn margin; spores delicately verruculose, orange, 17—24 p.

PQY

Fig. 132. P. Apii. Teleutospores and uredospore.

Uredospores. Sori hypophyllous, scattered, here and there
confluent, surrounded by the epidermis, pulverulent, cinnamon-
brown; spores ellipsoid to obovate or even subclavate, shortly
echinulate, slightly thickened above (8—5 yu), brownish-yellow,
24—35 x 20—26 pw, with three equatorial germ-pores.

. Leleutospores. Sori hypophyllous, rarely epiphyllous, if on

ON UMBELLIFERA 185

the petioles sometimes very large, scattered or confluent,
roundish, pulverulent, blackish-brown; spores ellipsoid to ob-
long, rounded above, not thickened, hardly constricted, rounded
or gently attenuated below, smooth, brown, 30—50 x 15—23 p;
pedicels hyaline, thin, deciduous, about as long as the spore.

On Apium graveolens. Not common. Aicidia in May and
June; teleutospores September—November. Distinguished

from many of its close allies by the possession of an ecidium.
(Fig. 132.)

DistRisution: Central and Northern Europe, East Indies,
Japan, Tasmania.

57. Puccinia ASgopodii Mart.

Uredo Aigopodii Schum. Plant. Sal. ii, 233.

Puccinia Aigopodii Mart. Fl. Mosquen. p. 226. Cooke, Handb. p. 502;
Micr. Fung. p. 208. Plowr. Ured. p. 201. Sacc. Syll. vii. 678 p.p.
Sydow, Monogr. i. 353. Fischer, Ured. Schweiz, p. 105, f. 79.

Teleutospores. Sori amphigenous, but chiefly on the petioles
and nerves, on thickened yellowish spots,
small, but collected into dense irregular
clusters and confluent, at first black, {
covered by the shining epidermis which
splits in places longitudinally, soon naked,
pulverulent, blackish-brown; spores ob-
long to ovoid, often irregularly angled and
oblique, usually rounded above and with
a pale wart-like apiculus 2—3 yw high,
hardly or not at all constricted (often broadest at the septum),
more or less rounded below, smooth, clear chocolate-brown,
granular, 28—48 x 15—22 w; pedicels hyaline, short, deciduous.

Fig. 133. P. A gopodii.
Teleutospores.

On Aigopodium Podagraria. April—August. Rather com-
mon. (Fig. 133.)

According to Tranzschel, a few isolated uredospores are to be found
in the young sori; they are almost colourless, aculeate, 20—22 x18 p.
Semadeni proved that the spores of this fungus would infect only
Afgopodium, and not any of the allied Umbellifers. In this species,

186 PUCCINIA

J find the septum of the teleutospores almost always comparatively broad
and dark, far more so than in the majority of Puccinias.

DistRIBUTION: Europe generally.

58. Puccinia Bulbocastani Fckl.

Aicidium Bulbocastant Cumino, Fung, Vall. Pis. 1804—5.

4, Bunii DC. Syn. p. 51. Cooke, Handb. p. 540 p.p.; Mier. Fung.
p. 196. Plowr. Ured. p. 270 (?).

Puccinia Bulbocastant Fckl. Symb. Myc. p. 52. Cooke, Micr. Fung.

p. 209. Sydow, Monogr. i. 363. Fischer, Ured. Schweiz, p. 133,
f. 100.

Spermogones. Few, scattered amongst the ecidia, pale-
yellowish.

Aicidiospores. Aicidia rarely on the
leaves, hypophyllous, more often on the
petioles and stems, densely crowded, caus-
ing considerable hypertrophy and curva-
ture, between cup-shaped and pustulate,
whitish, with a white irregularly torn
margin; spores delicately verruculose,
yellowish, 15—22 p.

Teleutospores. Sori amphigenous,
scattered, minute, roundish, sometimes
on the petioles confluent and elongated,
long covered by the epidermis, black;
spores ellipsoid to obovate-oblong, gener-
ally rounded at both ends, not thickened
above, hardly constricted, minutely reti-
culate, brown, 25—42 x 14—24; pedicels

Fig. 184. P. Bulbocas- hyaline, thin, deciduous.
tani. Whole plant of

C. Bulbocastanum, with On Carwm (Bunium) Bulbocastanum.
ecidia, nat. size. (Dun-

stable, April, 1896.) Very rare. Dunstable (W. G. Smith).
(Fig. 134.)

This species has no uredospores. Plowright confused together this
and the Puceinia tumida on Conopodium denudatum (see his synonymy
on pp. 206, 270). The latter species has no excidia; this partly explains
his remarks that he was unable to obtain any evidence of the connection
between the ecidium and the Puccinia. Nevertheless, it appears not yet

ON UMBELLIFERE 187

to have been shown experimentally that the ecidia and the teleutospores
described above are genetically connected. The markings on the teleuto-
spore are really little, round, densely crowded pits, not actual reticulations
as in P. Chaerophylli. — Plowright’s statement (Jc. p. 270) that this
zcidium was found by him on Conopodiuin denudatum is a mere slip of the
memory, as am informed by Mr W. G. Smith, in whose company it was
found at Leagrave, near Dunstable, on the date mentioned.

DIsTRIBUTION: Western and Central Europe, Algeria.

59. Puccinia tumida Grev.
Puccinia tumida Grev. Flor. Edin. p. 430 (1824). Sydow, Monogr.
i. 376.
P. Bunii Winter ; Plowr. Ured. p. 206.
P. Umbelliferarum DC. ; Cooke, Micr. Fung. p. 208 p.p., pl. 4, f.'71—2.

Uredospores. Very few, oval, pale yellow, sparsely verrucu-
lose, 20—25 x 15—18, mingled with the teleutospores.

Teleutospores. Sori on the leaves, more often on the petioles
and nerves, minute, but many crowded
together and confluent in thickened
elongated masses (up to 1 cm. long),
covered by the ash-coloured epider-
mis, for a considerable time, black-
brown; spores ellipsoid to ovate,
rounded at both ends, not thickened Fig. 135. P. tumida., Teleu-
above, hardly constricted, smooth, pepe aaa creand use:
brownish, 26—36 x 1426 w; pedicels
hyaline, short, deciduous; an occasional mesospore is found.

Cl)

On Carum majus (= Conopodium denudatum = Bunium flexu-
oswm). Not uncommon. April, May. (Fig. 135.)

Formerly confounded with P. Bulbocastani, from which it is distin-
guished by its very different habit. In that species the teleuto-sori
are usually isolated on the leaves, and cause no swelling of the affected
part as P. tumida always does: the latter moreover is without scidia.

Plowright states that the mycelium is perennial, but this is doubtful.
The sori appear to be confined to the radical leaves ; I have never seen them
on the cauline leaves or (what practically means the same thing) attacked
plants do not flower. The fungus should therefore be looked for early,
before the radical leaves begin to fade ; the affected plants are then easily
distinguished by their yellowish appearance.

188 PUCCINIA

The ecidium of P. Conopodti-Bistortae (q.v.) is sometimes to be found
on the same plant as the teleuto-sori of P. twmida, though it is much rarer.
Plowright mentions that, though the Puccinia is very common round
King’s Lynn, he could never find the ecidium which he at that time
wrongly supposed to be connected with it. See also Puccinia Bulbocastant.

DisTRIBUTION: France, Germany, Norway.

60. Puccinia Pimpinelle Mart.

AGcidium Pimpinellae Kirchn. (1856) ; Cooke, Micr. Fung. p. 196.

4. Bunti DC. ; Cooke, Handb. p. 540 p.p.

Uredo Pimpinellae Strauss in Wetter, Annal. ii. 102 (1811).

Trichobasis Pimpinellae Cooke, Micr. Fung. p. 224.

Puceinia Pimpinellae Mart. Fl. Mosquen. ed. ii. p. 226. Cooke,
Micr. Fung. p. 209. Plowr. Ured. p. 155 p.p. Sace. Syll. vii.
616 p.p. Sydow, Monogr. i. 408. Fischer, Ured. Schweiz, p. 127,
f. 97.

Spermogones. Amphigenous, mostly scattered amongst the
xcidia, pale-yellowish.

Aicidiospores. AKcidia hypophyllous, in smaller or larger
groups, often along the nerves and causing slight hypertrophy,
between cup-shaped and pustulate, with a whitish irregularly
cut margin; spores verruculose, subhyaline, 20—28 yu.

Uredospores. Sori hypophyllous, scattered, minute, pul-
verulent, cinnamon; spores globose to
ellipsoid, echinulate, brown, 22—32 x
20—26 p, with two (rarely three) germ-
pores.

Teleutospores. Sori similar, but
blackish-brown; spores ellipsoid, rounded
Fig. 136. P. Pimpinellae. at both ends, not thickened above, hardly

Teleutospore,on P.Sazi- constricted, reticulate, brown, 283—387 x

fraga (British); a, the < : jl
same, on P. magna (after 19—25 po; pedicels hyaline, deciduous,

Fischer): rather short.

On Pimpinella magna, P. Suafraga. Not common.
Acidia, May and June; teleutospores, July—October. (Fig.
136.)

Very similar to P. Chaerophylli ; distinguished from it especially by

the uredospores, which have for the most part a thicker and darker
membrane with only two germ-pores. The peridium of the ecidia is

ON UMBELLIFERE ° 189

better developed, and the teleutuspores are plainly but not so densely
reticulate. Klebahn proved by cultures that P. Pimpinellae is distinct
from P. Chaerophylli, and Semadeni similarly proved its difference from
that species and from P. Heraclei; the latter showed (Centralbl. £ Bakt.
2. xill. 215) that it could be transferred to other species of the genus
Pimpinetia, but not to other genera of the Umbelliferae.

DisrRIBUTION : Europe, Asia Minor, East Indies, Algeria.

61. Puccinia Bupleuri Rud.

Heidium Falcariae B. Bupleuri-falcati DC. Flor. fr. vi. 91.

Puccinia Bupleurt Rud. in Linnea, iv. 514 (1829). Cooke, Grevillea,
vi. 47. Plowr. Ured. p. 154.

P. Bupleuri-falcati Wint. Pilze, p. 212 (1884). Sydow, Monogr. i. 364.
Fischer, Ured. Schweiz, p. 123, f. 94,

[Spermogones. Hypophyllous, numerous, generally scattered
over the whole surface among the ecidia.

Aicidiospores. Bcidia hypophyllous, or a few epiphyllous,
uniformly scattered, cup-shaped,
with a torn white revolute mar-
gin ; spores globose or ellipsoid,
punctate, yellow, 16—24 u.]

Uredospores. Sori amphi-
genous, scattered or occasion- ;
ally circinate, on minute paler: gees (Wellton he Manele
spots, small, rounded, cinna-
mon; spores globose to ellipsoid, echinulate, yellow-brown,
19—24 x 17—22 y, with three, four, or even five germ-pores.

Teleutospores. Sori amphigenous, minute, scattered, round-
ish, on the stems often larger and oblong, occasionally confluent,
covered by the epidermis, at length naked, blackish-brown ;
spores oblong to clavate, rounded at both ends, not thickened
above, hardly constricted, smooth, brown, 25—44 x 16—30 pu;
pedicels hyaline, thin, short, deciduous.

On Bupleurum tenuissimum. Very rare. Uredo- and
teleutospores, Walton-on-the-Naze, August, 1887. (Fig. 187.)

The uredospores were found in small quantities among the teleuto-
spores. Fischer says that the affected plants usually bear ecidia on every
leaf, the leaves are narrower and paler, and the plants do not flower; the

190 PUCCINIA

ecidium-stage appears in May and June, often abundantly. This applies
especially to the parasite on Bupleurum falcatum, which is probably
identical with that on B. tenuissimum, but the Walton plants bore flowers.

Distripution: Europe, Asia Minor, East Indies, Yunnan.

62. Puccinia ASthuse Mart.

Uredo Petroselini DC. Flor. fr. ii. 597.

Trichobasis Petroselini Berk. ; Cooke, Micr. Fung. p. 223 (?).

T. Cynapit DC. ; Cooke, Micr. Fung. p. 224.

Puccinia Afthusae Mart. Flor. Mosq. ed. ii. p. 225 (1817). Cooke,
Micr. Fung. p. 209.

P. bullata Wint. ; Plowr. Ured. p. 185 p.p.

P. Petroselint Lindr. Faun. et Flor. Fenn. xxii., no. 1, p. 84 (1902).
Sydow, Monogr. i. 399, 889. Fischer, Ured. Schweiz, p. 112,
f. 86,

Spermogones. Hypophyllous, yellow-brown, or almost hya-
line.

Uredospores. Sori generally hypophyllous, scattered or in
small clusters, very small, occasionally confluent and larger,
pulverulent, cinnamon; spores globose to ellipsoid, echinulate
either all over or only in the upper part, thickened above
(5—6 p), yellowish or brownish-yellow, 22—29 x 21—25 p, with
three (rarely two) equatorial germ-pores with conspicuous caps.

Teleutospores. Sori similar, but dark-brown, on the petioles
and stems often larger, confluent and
elongated; spores ellipsoid or ovate,
rounded at both ends or slightly at-
tenuated below, not thickened above,
hardly constricted, smooth or nearly
so, brown, 28—48 x 18—25 pu ; pedicels
hyaline, thin, short, deciduous.

Fig. 188, P. Athusae. On Athusa Cynapium, Petroselinum
Teleutospore anduredospore, ¢,4,°
iG hia. sativum. Not common. June—Oc-
tober. (Fig. 138.)

It is possible that the forms on these two hosts are distinct species, or
at least biological races. Semadeni showed (Centralbl. f. Bakter, 2. xiii.
443) that, while he could infect several (non-British) species of Umbelliferee
with uredospores from Hithusa Cynapium, he could not infect Petroselinum
sativum ; at the same time he could find no morphological difference

ON UMBELLIFERA 191

between the two forms. Conium maculatum took the infection very
weakly.

Lindroth and Fischer both describe the teleutopores as furnished with
numerous minute embedded granules, otherwise even or with low rounded
undulations, but Fischer figures them as perfectly smooth, as they
certainly are in the cases I have seen. The uredospores are spiny in the
upper part, nearly smooth below. When the few spines on the basal part
are not to be discerned (as sometimes happens), they closely resemble
those of P. Conzi, except in being relatively broader ; these two species are
closely allied.

DISTRIBUTION : Central and Northern Europe.

63. Puccinia Silai Fckl.

Puceinia bullata Wint. Pilze, p. 191 p.p. Plowr. Ured. p. 183 p.p.
Sydow, Monogr. i. 403 p.p. Fischer, Ured. Schweiz, p. 119, f. 91 8.
P. Silat Fckl. Symb. Myc. p. 58. Cooke, Grevillea, xiv. 39.

Spermogones. Scattered, pale yellowish, accompanying the
primary uredo-sori.

Uredospores. Primary sori generally on the nerves and
petioles, elongated and con-
fluent up to 3 cm. long, dark
cinnamon; secondary hypo- (\
phyllous or occasionally epi-
phyllous, scattered, minute,
punctiform, brown; spores glo- ce )
bose to ovate, more or less
thickened above(4—5 ),echin- Fig. 189. P. Silai. Teleutospores.
ulate, brown, 25—40 x 18—

28 «, with three (rarely four) germ-pores.

Teleutospores. Sori minute, similar, but sometimes con-
fluent on the stems, blackish; spores obovate or oblong,
rounded at both ends or gently attenuated below, not thick-
ened above, but often with a papilla, slightly constricted,
smooth, brown, 28—42 x 18—82,,; pedicels hyaline, rather
short, deciduous.

On Silaus pratensis. Rare; Pontrilas; Kew Gardens.
August, September. (Fig. 139.)

192 PUCCINIA

This species was included by Sydow under the old P. bullata of Winter ;
but since then it has been rendered probable by the experiments of
Semadeni that it is distinct from most of the forms still remaining under
that collective name (Centralbl. f. Bakter. 1904, 2. xiii. 530). Schroter
proved that the spore-forms described above are genetically connected.
The uredo- and teleutospores are generally found intermixed.

64, Puccinia Angelice Fckl.

Uredo Angelicae Schum, Pl. Sall. ii, 233.

Trichobasis Angelicae Cooke, Micr. Fung. p. 224.

Puecinia Angelicae Fckl. Symb. Myc. p. 52. Cooke, Micr. Fung.
p. 208. Sydow, Monogr. i. 356. Fischer, Ured. Schweiz, p. 117,
f. 90.

P. Pimpinellae Str. ; Plowr. Ured. p. 155 p.p.

Spermogones. Few, scattered, roundish, faintly coloured.

Uredospores. Primary sori chiefly along the nerves and
petioles, or on the underside of the leaves in minute clusters
on deep-yellow spots, at first deep-yellow, then darker, at
length cinnamon; secondary, hypophyllous, occasionally epi-
phyllous, on minute paler spots, scattered, minute, pulverulent,
yellow-cinnamon; spores obovate to oblong, echinulate, much
thickened (5—10 p) above, pale-brown, 25—40 x 22—28 yw, with
three equatorial germ-pores.

Fig. 140. P. Angelicae. Two normal teleutospores, also one abnormal and
& mesospore ; a, a uredospore; b, the primary uredo-sori (reduced). On
A, silvestris.

Teleutospores. Sori amphigenous, scattered, minute, rounded,
pulverulent, black-brown or black; spores oval to oblong, rounded
and scarcely thickened above, hardly constricted, rounded or
attenuated below, smooth, brown, 30—50 x 16—24 4; pedicels
hyaline, short, deciduous; an occasional mesospore is seen.

ON UMBELLIFER 1938

On Angelica silvestris. Not common. Uredospores, June
and July; teleutospores, July—November. (Fig. 140.)

Distinguished especially by the very bright yellow primary uredo-sori,
and by the slightly larger uredospores than in allied species. Semadeni
proved that it would grow also on Archangelica, but not on Athusa
Cynapium or Peucedanum palustre. The primary uredo-sori should be
compared with the ecidia of P..Smyrnii. P. Apii is also closely allied,
but differs in the possession of a true cup-shaped ecidium.

DIsrRIBUTION: Central and Northern Europe, Turkestan.

65. Puccinia bullata Wint.

Uredo bullata Pers. Obs. Mye. i. 98 p.p.

P. bullata Wint. Pilze, p. 191 p.p. Sace. Syll. vii. 634 p.p. Sydow,
Monogr. i. 408 p.p. Fischer, Ured. Schweiz, p. 119, f. 91 a.

P. Umbelizferarum and P. bullaria, p.p.

Spermogones. Pale yellowish.

Uredospores. Primary sori chiefly on the swollen nerves
and petioles, elongated, as much as
3 cm. long, dark-cinnamon, secondary
hypophyllous or rarely epiphyllous, scat-
tered, minute, punctiform, brown; spores
globose to obovate, echinulate, brown,
apex more or less thickened, 25—40 x
18—28 p, with three or four germ-pores
with swollen caps.

Teleutospores. Sori similar, often Fig. 141. P. bullata, Te-
confluent on the stems, long covered by ithe ee
the epidermis, blackish-brown; spores
oblong to obovate, rounded above, scarcely thickened, hardly
constricted, slightly narrowed below, smooth, uniformly brown,
30—45 x 18—24 w; pedicels hyaline, rather stout, deciduous.

On Peucedanum palustre. Shrapwick Bay, Somerset, 1883
(H. B. Waterfall). (Fig. 141.)

This species is placed temporarily in the collective group, P. bullata
Wint. All the forms which were included under that head by Plowright
are now separated on morphological or biological grounds, but nothing is
known as yet about the Puccinia on Peucedanum palustre, from the latter
point of view. It is a close ally of P. Stlad.

G. U. 13

194 PUCCINIA

66. Puccinia Heraclei Grev.

Trichobasis Heraclei Berk. ; Cooke, Micr. Fung. p. 225.

Puccinia Heraclei Grev. Scot. Cr. Flor. pl. 42. Cooke, Handb.
p. 502; Micr. Fung. p. 208. Sydow, Monogr. i. 387. Fischer,
Ured. Schweiz, p. 132.

P. Pimpinellae Strauss ; Plowr. Ured. p. 155 p.p.

Spermogones. Amphigenous, scattered amongst the ecidia,
pale-yellowish.

4Aicidiospores. icidia hypophyllous, frequently on the
petioles and especially on the nerves of the
leaves, on thickened yellowish spots, densely
crowded in irregular clusters, often causing
distortion, between cup-shaped and pustulate,
sometimes almost spherical and superficial ;
peridium feebly developed, opening by a
rounded pore; spores delicately verruculose,

: yellowish, 21—32 x 18—28 p.
ae gare ile Uredospores. Sori hi ttered
clei. Aicidia on sp amphigenous, scattered,
a (re- minute, chestnut-brown; spores globose to
ellipsoid, densely echinulate, pale-brown, 25—
32 x 19—27 yp, with three or four germ-pores.

Teleutospores. Sori similar or more or less confluent on the
nerves, surrounded by the ferruginous epidermis, pulverulent,
blackish ; spores ellipsoid, rounded at both ends, hardly con-
stricted, reticulated, brown, 26—387 x 18—27 w; pedicels hya-
line, short, deciduous.

On Heracleum Sphondyliwm. Aicidia, March—June; teleu-
tospores, August. Not common. (Fig. 142.)

This species closely resembles P. Chaerophylli, but is distinguished by
its less densely reticulated teleutospores. Semadeni proved by experi-
ment that they are distinct species, but no one has as yet reared all the
spore-forms of P. Heraclei from the basidiospores, as has been done
for P. Chaerophylli. The ecidia of this species are more conspicuous
than those of its allies; they occur in swollen patches, reminding one
of the acidia on Smyrnium Olusatrum, and being sometimes almost
spherical and superficial might be compared to a group of miniature Peziza
vesiculosa.

DISTRIBUTION: Central and North-western Europe.

ON UMBELLIFERZ 195

67. Puccinia Cherophylli Purt.

Pucceinia Chaerophylli Purton, Midl. Flor. iii. 303. Sydow, Monogr.
i. 367. Fischer, Ured. Schweiz, p. 129, f. 98.
P. Pimpinellae Str. ; Plowr. Ured. p. 155 p.p.

Spermogones. Pale-yellow, roundish.

4icidiospores. Aicidia on the leaves and petioles, on the
leaves scattered or circinate, on the petioles and nerves in
dense elongated clusters and causing a slight hypertrophy,
between cup-shaped and pustulate, yellowish; peridium poorly
developed ; spores verruculose, orange, 18—35 x 16—26 u.

Uredospores. Sori hypophyllous, scattered, minute, roundish,
pulverulent, cinnamon ; spores globose to obovate, echinulate,
pale brownish-yellow, 20—30 x 18—25; with three usually
equatorial germ-pores.

Teleutospores. Sori similar, but black-brown, on the petioles
more elongated ; spores ovate to oblong,
rounded at both ends or gently at-
tenuated below, not thickened above,
slightly constricted, reticulated, yellow-
ish-brown or brown, 24—36 x 16—25 p;
pedicels hyaline, thin, as long as the
spore or shorter.

On leaves, petioles, and stems of
Anthriscus silvestris, Chaerophyllum pig. 143. P Chuerophylli.

temulum, Myrrhis odorata. -Not com- Teleutospores, a, on
a Anthriscus, b, on Myr-
mon. Aicidia, May and June; teleuto- rhis.

spores, July—October. (Fig. 143.)

It was proved experimentally by’ Klebahn that this parasite is not
identical with P. Pimpinellae, and by Semadeni that it is not identical
with that or with P. Heraclei. The latter also reared spermogones and
zecidia from the basidiospores, and uredo- and teleutospores from the
zcidiospores, of P. Chaerophylli; thus proving that all the spore-forms
described above belong to the same species. The feeble development of
the peridium and the pustule-like, not cup-shaped, xcidia are paralleled
by those of P. Heracler. ‘The markings on the teleutospore are formed by
a network of low ridges, with small polygonal or roundish meshes. The
spore-reticulation is exactly of the same character as that of P. Pimpinellae
except that the meshes are a little smaller and not quite so easily seen.

13—2

196 PUCCINIA

Semadeni showed that the spores from A. s¢lvestris infected M. odorata
readily (Centralbl. f. Bakt. pt. 2, xiii. 217—9), but whether the form on
C. temulum belongs to the same species (or is a biological race of it) seems
at present to be undetermined.

DisTRIBUTION: Central and Northern Europe, Siberia.

68. Puccinia Conii Fckl.

Credo Cont Strauss in Wetter. Ann. ii. 96.

Trichobasis Conti Cooke, Micr. Fung. p. 225.

T. Umbellatarum Lévy. ; Cooke, Micr. Fung. p. 225 p.p.

Puccinia Conti Fckl. Symb. Myc. p. 538. Cooke, Handb. p. 209 p.p.
Sacc. Syll. xiv. 302. Sydow, Monogr. i. 375. Fischer, Ured.
Schweiz, p. 114, fi 87.

P. bullaria Link ; Cooke, Handb. p. 503 p.p.

P. bullata Wint. ; Plowr. Ured. p. 183 p.p.

Uredospores. Sori hypophyllous, occasionally on the petioles,
scattered, minute, rarely confluent, pul-
verulent, cinnamon ; spores ellipsoid to
obovate, thickened (up to 7) above
and echinulate in the upper part only,
pale-brownish, 2436 x 17—26 w, with
three germ-pores.

Teleutospores. Sori similar, but
blackish-brown, on the stems and petioles
ae eee often larger and long covered by the
leutospore and uredospore. gYey “epidermis ; spores ovate or ovate-

oblong or even clavate, rounded at both
ends or attenuated below, not thickened at the apex but with a
small papilla or pore-cap, hardly constricted, nearly or quite
smooth, pale-brown, 30—48 x 20—28 yw; pedicels hyaline, short,
deciduous.

On Conium maculatum. Not common; England, Wales,
Ireland (Clare Island). August, September. (Fig. 144.)

Distinguished by the uredospores, which are echinulate only in the
upper half ; the spines gradually diminish in size downwards and the lower
half is quite smooth. The teleutospores which I have examined are quite
smooth when empty, even under the highest power, but the protoplasm is
very granular and presents a misleading effect at first sight.

ON UMBELLIFERAE 197

69. Puccinia Smyrnii Corda.

Heidium Bunii var. Smyrnti-Olusatri DC. Flor. fr. vi. 96.
Trichobasis Petroselini Berk. ; Cooke, Handb. p. 529 p.p.
Puceinia Smyrnii Corda, Icon. iv. 18, f. 67 (1840). Cooke, Handb.
p. 563; Micr. Fung. p. 209, pl. 3, f. 55—6. Plowr. Ured. p. 199.
Sace. Syll. vii. 670.
P. Smyrnii-Olusatri Lindr. Faun. et Flor. fenn. xxii. no. 1, p. 9
(1902). Sydow, Monogr. i. 416.
Spermogones. Epiphyllous, on sunken spots.
Ajcidiospores. Aicidia hypophyllous or occasionally epi-
phyllous, in rather large irregular clusters, or on the petioles
and stems in elongated groups, on yellow spots, hemispherical,
yellow, opening by an irregular pore with a nearly entire
margin; spores globose or ovate to fusiform or pyriform,
delicately verruculose, yellowish, 16—40 x 16—20 p.
Teleutospores. Sori hypophyllous, on small yellow spots, scat-
tered or a few together, minute,
pulverulent, dark-brown; spores
ellipsoid to oblong, rounded at
both ends, not thickened above,
hardly constricted, coarsely and
remotely reticulate and tuber-
culate, brown, 30—48 x 17—
26 w; pedicels hyaline, thin, de-
ciduous, up to 604 long; epi-
spore rather thick. Fig. 145.

P. Smyrnii. Teleuto-
x spores; a, as really sculptured;
On Smyrnium Olusatrum. i sig oes iar aaa

Rather common near the coast.
Aicidia, A pril—June; teleutospores, June—August. (Fig. 145.)

The ecidia and teleuto-sori may occur on separate plants or on the
same. The markings on the teleutospore form a wide-meshed network,
which bears wart-like tubercles at the angles of the meshes. The ecidio-
spores bear more resemblance in form to uredospores than to what they
really are; but they are produced in chains with intercalary cells in the
usual way: they are not echinulate, but delicately verruculose ; the
markings can easily be seen on an empty spore. The peridium-cells are
grossly verrucose on the inner surface, and are not arranged in regular
rows. It is this irregularity which causes the peridium (as is usual in
such cases) not to split into lacinia, but to open by a pore.

198 PUCCINIA

DisTRIBUTION: Central and Eastern Europe, Crete, Cyprus,
Asia Minor, Algeria.

70. Puccinia Circee Pers.

Puccinia Circaeae Pers. Disp. Meth. p. 39, pl. 3, f. 4. Cooke, Handb.
p. 507; Micr. Fung. p. 211. Plowr. Ured. p. 213. Sace, Syll.
vii. 686. Sydow, Monogr, i. 422. Fischer, Ured. Schweiz, p. 319,
f, 232, ;

Teleutospores. Sori hypophyllous, on sunken yellowish or
purplish round spots, minute, pulvi-
nate, brown, then with a greyish
bloom, scattered or circinate and at
length confluent in a thick crust;
spores generally fusoid, rounded or
conically attenuated above and much
thickened (up to 12 4), gently constric-
Peale Laue ted, attenuated downwards, smooth,
Teleutospores. yellowish-brown or brown, 25—40 x
9—13,; pedicels hyaline, persistent,

about as long as the spore.

On Circaea alpina, C. lutetiana. Rather common. August—
October. (Fig. 146.)

The sori of this species present two different forms: the first-formed
are roundish, clear-brown, solitary or circinate and confluent ; the later-
formed, which appear round the others or on the stem and on the nerves
of the leaves, are darker-brown and never greyish. All the spores are of
the same shape, but the paler ones can germinate at once, in the sorus,
while the darker ones rest until the following spring, as in P. Veronicarum.
There is another Uredine on the same host, even occurring on the same
leaf: see Pucciniastrum Circaeae.

DIsTRIBUTION: Central and Northern Europe, North
America, East Indies.

71. Puccinia pulverulenta Grev.

Heidium Epilobi DC. Flor. fr. ii, 288. Cooke, Handb. p. 586; Mier.
Fung. p. 195.

Uredo vagans var. Kpilobii-tetragoni DO. Flor. fr. ii. 228,

Trichobasis Epilobit Berk. ; Cooke, Micr, Fung. p. 226.

ON EPILOBIUM 199

Pucecinia pulverulenta Grev. Fl. Edin. p. 432 (1824). Cooke, Handb.
p. 507; Micr. Fung. p. 211, pl. 4, f. 78—9. Plowr. Ured. p. 151.

P. Epilobit DC. ; Sace. Syll. vii. 608 p.p.

P. Epilobii-tetragoni Wint. Pilze, p. 214 (1884). Sydow, Monogr.
i. 424, Fischer, Ured. Schweiz, p. 152, f. 118. McAlpine, Rusts
of Australia, p. 170, f. 79—81.

Spermogones. Scattered among the ecidia, honey-coloured.

Aicidiospores. Aicidia hypophyllous or, when very abundant,
also epiphyllous, scattered rather closely over nearly the whole
surface of the leaf, cup-shaped, with a white torn revolute
margin; spores very delicately verruculose, orange, 16—26 yu.

Uredospores. Sori hypophyllous, scattered or circinate,
sometimes confluent, pulverulent,

chestnut-brown; spores globose to
ovoid, remotely echinulate, brown, ks

20—28 x 1525 p, with two germ-
pores. (0) ey,

Teleutospores. Sori hypophyl-
lous, often circinate, soon naked, _.

Fig. 147. P. pulverulenta. Te-
pulverulent, dark-brown > spores leutospores and uredospore.
ellipsoid or ovoid, rounded at both
ends, somewhat thickened above (up to 5) with a broad low
cap-like addition, gently constricted, smooth, brownish, 24—35
x 14—20 w; pedicels hyaline, slender, deciduous.

On LEpilobium hirsutum, E. montanum, HE. tetragonum.
fEcidia, May and June; teleutospores, June—November.
Common. (Fig. 147.)

The aecidium-forming mycelium appears (but perhaps falsely) to be
perennial, for the same plants are attacked year after year. The acidia
appear in May and cover leaf after leaf, as they are developed. The
affected plants are easily recognisable by their much paler and yellowish
colour. Soon the sori of uredo- and teleutospores begin to appear, at first
on the same leaves, but afterwards on the later-formed leaves higher up
the plant. In September and October the small last-formed leaves are
thickly covered by the teleutospores ; it is probably from the germination
of these in spring that the next attack proceeds. The mycelium of the
uredo- and teleuto-sori is strictly localised.

Plowright states (/.c.) that the ecidiospores sown on young seedlings of
E. hirsutum gave rise to ecidiospores in seventeen days, but very possibly

200 PUCCINIA

there is some oversight here, for Dietel obtained the wredo by sowing the
zcidiospores from £. tetragonum on that plant ; he also suggests that the
form on £. tetragonum is biologically distinct from that on E. hirsutum,
since on the latter he obtained no result.

DistRIBUTION: Central and Northern Europe, Nova Zembla,
Siberia, North America.

72. Puccinia Epilobii DC.

Puccinia Epilobit DC. Flor. fr. vi. 61. Cooke, Handb. p. 506; Micr.
Fung. p. 211. Plowr. Ured. p. 202. Sydow, Monogr. i. 427.
Fischer, Ured. Schweiz, p. 155, f. 120.

Teleutospores. Sori hypophyllous, scattered or rather crowded,
often uniformly distributed over the whole
surface of the leaf, rarely confluent, sur-
rounded by the torn epidermis, pulverulent,
reddish-brown; spores ellipsoid, oblong or
pyriform, rounded at both ends, hardly
thickened above, but with a minute papilla,
F .,.. much constricted, minutely verruculose,
ee clear brown, 27—48 x 1625 ps; pedicels
hyaline, 10—16 w long.

On Epilobium palustre. Rare. May—August. A sub-
alpine and arctic species. (Fig. 148.)

Distinguished from P. pulverulenta not only by its spores and the
absence of the zcidium- and uredo-stages, but also by the smaller teleuto-
sori which are scattered pretty uniformly over the leaf-surface. The
mycelium seems to be perennial; it permeates the whole plant, and
somewhat deforms the shoots, making the leaves smaller and thicker,
The warts of the epispore are sometimes hardly perceptible, but can
usually be seen if the spore is squeezed in the way recommended, see
p. 83.

DistTRIBUTION : Central and Northern Europe.

73. Puccinia Viole DC.

Aicidiwm Violae Schum. Pl. Sill. ii, 224. Cooke, Handb, p. 548;
Micr. Fung. p. 198.
Trichobasis Violarum Lév. ; Cooke, Micr, Fung. p. 226.

ON VIOLA 201 .

Puccinia Violae DC. Flor. fr. vi. 62 (1815). Plowr. Ured. p. 152.
Sydow, Monogr. i. 439. Sacc. Syll. vii. 609. Fischer, Ured.
Schweiz, p. 139, f. 106.

P. Violarum Link, Sp. Plant. ii. 80 (1824). Cooke, Handb. p. 504 ;
Micr. Fung. p. 210; Grevillea, iii. pl. 49, f. 5, 10 a, 6. ;

Spermogones. Crowded in little clusters, yellowish.

4icidiospores. Aicidia on all the green parts of the host,
on the leaves often forming swollen yellowish spots, generally
in roundish, or irregularly expanded, groups, on the stem some-
times scattered, flat, with a white irregularly torn revolute
margin; spores delicately verruculose, orange, 16—24 x 10—
18 py.

Fig. 149. P. Violae. u, leaf of V. silvatica with xcidia; b, teleutospore,
seen wet; c, the same, seen dry; d, a mesospore.

Uredospores. Sori hypophyllous, scattered or circinate,
minute, soon naked, pulverulent, cinnamon-brown; spores glo-
bose to ellipsoid, echinulate, brownish, 20—26 x 17—28 yw, with
two germ-pores.

Teleutospores. Sori similar, but darker; spores ellipsoid to
oblong, rounded at both ends or gently attenuated below,
thickened and paler above, hardly constricted, faintly punctate,
chestnut-brown, 20—40 x 15—23,; pedicels hyaline, deciduous,
rather long; a few mesospores are found.

On all green parts of Viola canina, V. hirta, V. odorata,
V. Riviniana, V. silvestris. Very common. Aicidia, April—
June; teleutospores, August—November. (Fig. 149.)

The punctation of the teleutospores is very delicate, like little pin-
pricks, and can only be seen when they are dry ; these spores are generally
described as smooth and appear so except in the most favourable
circumstances, The germ-pore of each of the cells is covered with a paler
convex cap. The connection of all the spore-forms of this species was

202 PUCCINIA

experimentally proved by De Bary and Jacky. Bock (Centralbl. fiir Bakt.
2. xx. 586) showed that what appeared to be identical with P. Violae could
also be artificially produced on V. cornuta, V. lutea, V. tricolor; the
zcidia formed small or large crowded well-defined groups, and thereby
differed from those of P. aegra which spread widely over the stems and
leaves of their hosts, mostly standing alone and not collected in groups.
In the latter, moreover, ecidia and teleutospores can be found in July at
the same time (or even on the same leaf), which is not usually the case
with P. Violae. This species can attack cultivated violets, but rarely does
much harm ; burning the infected plants is a sufficient remedy.

DisTRIBUTION : World-wide, except Australia.

74. Puccinia egra Grove.

Gcidium depauperans Vize, Gard. Chron. (1876) pp. 175, 361, 437.
Cooke, Micr. Fung. p. 195 ; Grevillea, v. 57.

Puccinia aegra Grove, Journ. Bot. xxi. (1883) p. 274. Plowr. Ured.
p. 158. Sace. Syll. vii. 614.

P. depauperans Sydow, Monogr. i, 442 (1903).

4icidiospores. Aicidia on all green parts of the host,
particularly on the stems, not in clusters, but spread pretty
uniformly over the whole surface, especially of the stems which
are swollen and distorted, cup-shaped, with a white torn revolute
margin; spores globose to ellipsoid, smooth, orange, 17—21 x
14—16 p.

Uredospores. Sori amphigenous, without spots, irregularly
scattered or aggregated, occasionally
confluent, long covered by the lead-

| coloured epidermis, cinnamon ; spores
globose to ellipsoid, delicately echinu-
late, brownish 20—28 pw diam.

en ae we re Teleutospores. Sori similar, but
spores and mesospore. darker ; spores ellipsoid or ovoid,
rounded or gently attenuated above,
where it is slightly thickened or rather surmounted by a minute
subhyaline papilla, hardly constricted, usually rounded below,
very delicately punctate or quite smooth, brown, 22—34 x 16—

20; pedicels hyaline, short; a few mesospores are found.

ON VIOLA 208

On Viola cornuta, V. lutea, V. tricolor, and most of the
many hybrid Violas and Pansies now cultivated in gardens.
Fresh ecidia are formed right through the summer till August,
while those of P. Violae cannot usually be found after early
June. (Fig. 150.)

The mycelium of the ecidia is perennial in the underground parts ; all
the shoots which arise from the affected plant are deformed, the internodes
are lengthened, the leaves become smaller, paler and often twisted. In
P. Violae this is not the case; only a slight swelling arises at the part
where the localised mycelium is producing its wxcidia. Bock (Centralbl.
fiir Bakt. 2. xx. 586) found that he could produce the ecidia of P. Violae
on the three species of Viola named above, by artificial infection ; but like
others he still considered the two species as distinct on account of their
different habit. Liro, on the contrary, considers them as the same. In
my experience, the uredo- and teleuto-sori of P, aegra are larger, and
remain longer covered by the epidermis, but the spores are identical.

This species can do considerable harm if allowed to spread ; there is
no remedy, but all infected plants should be carefully uprooted and burnt.
A Viola, badly attacked by the ecidium, was once sent to the Gardener’s
Chronicle by a correspondent as “a hybrid between a fern and a violet.”

DISTRIBUTION : Germany, Denmark.

75. Puccinia Fergussoni B. et Br.

Puccinia Fergussoni Berk. et Br. Ann. Nat. Hist. 1875, p. 35. Cooke,
Grevillea, iii. 179, pl. 49, f. 10 ¢; Micr. Fung. p. 210. Plowr.
Ured. p. 207. Sacc. Syll. vii. 682. Sydow, Monogr. i. 444. Grove,
Journ. Bot. 1912, p. 10.

Teleutospores. Sori hypophyllous or on the petioles, on
large roundish or irregular yellow spots,
in suborbicular or (on the petioles) elon-
gated clusters up to 14 em. long, densely
crowded and confluent, long covered by \)
the epidermis, then pulverulent, choco-
late-brown; spores irregular, generally \
oblong, attenuated or rarely rounded at fig.151. P. Fergussoni.
both ends, thickened above in a conical poner all and me-
form (up to 6 ,), gently constricted,
smooth, pale-brown, 26—45 x 12—18 yw; pedicels hyaline, thin,
deciduous, up to 30 w long; an occasional mesospore is found.

204 PUCCINIA

On Viola palustris. Rare; Wales, Scotland, and near
Birmingham. The beginnings of the sori may be seen by the
middle of May. (Fig. 151.)

This species is easily recognised by its large and pulvinate groups of
sori. The mycelium spreads considerably beyond the part occupied by
the spores, and consequently causes large yellow patches, usually only one
or at most two on each leaf, each the result of a separate infection by the
basidiospores.

P. asarina Cooke, Handbook, p. 504, Plowright, Uredinez, p. 202 (non
K. et §.), is this species, a mistake having been made in identifying the
host-plant. In continental specimens of P. asarina Kunze, so far as I have
seen, the sori on the lamina are as frequent on the upper leaf-surface
as on the lower, whereas in P. Fergussoni they are entirely hypophyllous.

DistrisuTion : Northern Europe, North America (?).

76. Puccinia argentata Wint.
Aeidium argentatum Schultz, Prod. Flor. Starg. p. 454 (1806 ; teleuto-
spores, on Impatiens).
Trichobasis Impatientis Rab. ; Cooke, Micr. Fung. p. 225.
Puccinia Noli-tangeris Corda, Icon. iv. 16, pl. 5, f. 57 (1840). Cooke,
Handb. p. 504; Micr. Fung. p. 210.
P. argentata Winter, Pilze, p. 194 (1884). Plowr. Ured. p. 193. Sace.
Syll. vii. 639. Sydow, Monogr. i. 450. Fischer, Ured. Schweiz,
p. 143, £. 109.
[Spermogones. Hypophyllous, scattered among the ecidia,
honey-coloured.
Aicidiospores. AKcidia hypophyllous, pretty uniformly dis-
tributed on discoloured swollen
iB al a) spots, on the petioles and
stems more scattered, white,
with a deeply-cut revolute
- margin ; spores 18—22 x 13—
CD 20 2; contents golden-yellow. ]
Uredospores. Sori hypo-
Fig. 152. P. argentata. Teleutospores, phy lous, scattered we circinate,
from the original specimen (L. Jenyns). sometimes on minute yellowish
spots, often confluent, covered
by the silvery epidermis, then pulverulent, roundish, ochra-
ceous; spores globose to broadly ellipsoid, delicately echinulate,

ON IMPATIENS 205

pale-yellowish, 16—22 x 1420, with 3—5 (usually four)
germ-pores.

Teleutospores. Sori similar, but chestnut-brown; spores
ellipsoid to subclavate, with a colourless conical cap to each
germ-pore, rounded or slightly attenuated at both ends, hardly
constricted, smooth, pale-brownish, 25—38 x 12—22 w; pedicels
hyaline, slender, short.

{[AMcidia on Adoxa Moschatellina, April—June ;] uredo and
teleutospores on Impatiens fulva, I. Noli-tangere, May, August—
October. Very rare; Albury, Surrey, October, 1864 (Rev. L.
Jenyns), Guildford (Rev. W. A. Vize), Shere (M. C. Cooke),
Kew Gardens (G. Massee). (Fig. 152.)

The teleutospores are at first produced in the same sori as the uredo-
spores ; the ecidial stage is not recorded, probably because it has been
confounded with the ecidium of P. albescens, from which it is distinguished,
according to Bubék, by its gold-coloured spores. I do not think this
distinction will hold good ; many of the specimens found in this country,
which appear to be P. albescens, have golden-yellow spores. See under
that species (p. 163).

The description given above of the spermogones and ecidia is taken
from Bubak, who showed (Centralbl. fiir Bakt. 2. xii. 413) that they could
produce the other stages on Jmpatiens in about ten days. He also showed
that the ecidium could be produced on the Adowa by over-wintered
teleutospores ; the incubation period was as long as one month, probably
because the mycelium first permeated the whole plant, from the leaf to
the stem, before producing spores. Afterwards he proved, contrary to his
former opinion, that the mycelium does not perennate in the rhizome, but
fresh infection must take place each spring (cbed. xvi. 150).

DisrrisuTion: Central and Northern Europe, North
America, Japan.

77. Puccinia Buxi DC.

Puceinia Buxt DC. Flor. fr. vi. 60. Cooke, Handb. p. 508; Micr.
Fung. p. 212. Plowr. Ured. p. 217. Sacc. Syll. vii. 688. Sydow,
Monogr. i. 453. Fischer, Ured. Schweiz, p. 316, f. 228—30.

Teleutospores. Sori amphigenous, on indefinite spots, scat-
tered or confluent, hemispherical, pulvinate, hard, compact,
soon naked, dark chestnut-brown or purplish-brown; spores

206 PUCCINIA

’

oblong to clavate, rounded above and not thickened, evidently
constricted, usually attenuated below,
smooth, brown, 55—90 x 20—35 w (or
even 100 u long); pedicels hyaline, per-
sistent, very long, (reaching 160 4).

On Buaxus sempervirens. Rather
common. September, October, lasting
through the winter and following spring.
(Fig. 153.)

The spores of this species easily fall apart
into their component cells. Ed. Fischer proved
that it has only the one spore-form : he gives
(.c.) figures showing the effect upon the leaf
of an infection by the basidiospores. Accord-
ing to him, the teleutospores germinate in
spring, and infect the delicate young leaves.
The mycelium grows slowly. During the
summer and autumn the infected spot becomes
much thickened : the sori are produced in late

; autumn or during the following winter. This
z= Nee = Sia is exactly in accordance with the suggestion
: made by Plowright (/.c.), without any experi-

mental evidence being at that time available.

DISTRIBUTION: Europe and Persia.

78. Puccinia Malvacearum Mont.
Puceinia Malvacearum Mont. in Gay, Hist. Chile, viii. 43. Cooke,
Mier. Fung. p. 205; Grevillea, ii. 47, 1387 and iii. pl. 35, pl. 49,
f.1. Plowr. Ured. p. 212. Sacc. Syll. vii. 686. Sydow, Monogr.
i. 476. Fischer, Ured. Schweiz, p. 313, f. 227. McAlpine, Rusts
of Australia, p. 178, f. 99, 100, 123—130, and pl. F, f. 28.
Teleutospores. Sori hypophyllous or amphigenous, and on
the petioles and stems, on conspicuous yellow or orange spots,
scattered but close together, small, hemispherical or on the
stems elongated, pulvinate, compact, hard, at first pale-reddish,
then reddish-brown; spores oblong to subfusoid, attenuated
at both ends or rarely rounded above, thickened at the apex,
gently constricted, smooth, yellowish-brown, 35—75 x 12—26 p;
pedicels hyaline, persistent, short or as much as 150 long;
one-, three-, or even four-celled spores also occur.

ON MALVACEE 207.

On many species of Malvacee (of the subfamily Malveze),
especially on Malva moschata, M. silves-

tris and Althaea rosea. Very common. .

May—October (also in April and Novem-

ber). (Fig. 154.)

This is.one of the most noticeable of the |

Uredinales. It is truly plurivorous ; so far

from being confined to a species, it is not even

confined to a genus. In botanic gardens, where ;

species of the family Malvacez are grown, side

by side, in the same plot, the disease can be

seen to spread to plants of all the allied genera

—NMalva, Lavatera, Althaea, Kitaibelia, Malope,

Abutilon, Sida, Sidalecea, Anoda, Malvastrum, ;

etc., have been recorded. A list containing ;

many (nearly forty) species of these genera is

given by Sydow, to which more are added by

Fischer, McAlpine, and Dandeno. Onallthese _ .-

it appears to be identical ; artificial infections ny Las ares Yaa
a eis eleutospores and meso-

have proved that it can be transferred from spore.

Malva to Althaea, and vice-versd.

It was first made known in 1852 by Montagne from a specimen found
in Chili. It was observed in Australia in 1857 (McAlpine). In Europe it
appeared in 1869, in South Africa in 1875, and it is now spread all
over the world. It is believed that Chili was its native home; the
rapidity of its distribution to other countries has few or no parallels among
plant diseases.

It has been proved by many experimenters that it produces only the
one kind of spore, which is capable of germinating at once when mature,
though some can hibernate. It is a disputed point whether the mycelium
can pass the winter in the plant or in the seed: the balance of evidence
goes to show that fresh infections arise each year by the germination
of over-wintered teleutospores, which can be found on all green parts, even
on the fruits. See p. 48.

79. Puccinia Pruni-spinose Pers.

Heidium punctatum Pers. in Uster. Annal. Botanik, xx. 135. Plowr.
Ured. p. 268.

4, quadrifidum DC. Flor. fr. vi. 90. Cooke, Handb. p. 586; Micr.
‘Fung. p. 194.
Puccinia Pruni-spinosae Pers. Syn. p. 226 (1801). Sydow, Monogr. i.
484, Fischer, Ured. Schweiz, pp. 157, 547, ff. 21, 122, 340.
McAlpine, Rusts of Australia, p. 171, f. 83-—6 and pl. D, f. 19—20.

208 PUCCINIA

P. Pruni DC. Flor. fr. ii, 222 (1805). Plowr. Ured. p. 192. Sace.
Syll. vii. 648.

P. Prunorum Link, Sp. Pl. ii. 82 (1825). Cooke, Handb. p. 507;
Micr. Fung. p. 211; Grevillea, iii. pl. 49, f. 11.

Tranzschelia punctata Arthur, North Americ. Fl. vii. 151.

Spermogones. Amphigenous, scattered, brown or blackish,
very shallow, punctiform.

Ajeidiospores. AKcidia hypophyllous, scattered over the
whole surface, flat, with a

broad revolute margin which
a is torn into few (8—5) lobes ;
spores roundish, pale yellow-
ish-brown, finely verruculose,

16—24 p.
Uredospores. Sori hypo-
“ b phyllous, generally on minute
Fig, 168, 2. Pnatspinowe Zale coloured spots, seattered, but
an ecidium on A. nemorosa, York- often crowded and confluent,
Seni fe ee and }, twoless soon naked, pulverulent, cin-
namon-brown; spores ellip-
soid to fusiform, ovoid-oblong or sub-pyriform, smooth and
more or less thickened at the summit in a conical shape and
darker, paler and narrowed below, where they are sharply

Fig. 156. P. Pruni-spinosae. Two teleutospores ; a, lower half of a teleuto-
spore; b, uredospore; c, paraphysis; d, a cluster of teleutospores. On
Wild Plum. (All x 600, except d, which is x 360.) :

verrucose or echinulate, pale-brown, 20—85 x 10—18 yp (with
three or four equatorial germ-pores, Arthur), mixed with
yellowish-brown or pale capitate paraphyses more or less
thickened at the apex.

ON PRUNUS 209

Teleutospores. Sori similar but blackish-brown; spores
ellipsoid to oblong, composed of two globose cells which readily
separate (or lower cell often narrower, paler and imperfect), not
thickened above, densely and coarsely verrucose, brown, 30—
45 x 18—25 »; pedicels hyaline, very short, deciduous, spring-
ing in clusters of about 10—20 from a common base.

kcidia on Anemone coronaria, A. nemorosa, April and May;
uredo- and teleutospores on Prunus domestica, P. insititia,
P. spinosa, also on cultivated species and varieties of Prunus
_and its allies, August—October. Common in certain districts.
(Figs. 155, 156.)

The discovery of the hetercecism of this parasite is comparatively
recent. Tranzschel first showed (in 1904) that it was hetercecious, using
Anemone coronaria and Amygdalus communis as the alternate hosts.
F. T. Brooks, at Cambridge in 1911, laid fresh eecidiospores from the same
species of Anemone on both sides of certain leaves of a “ Victoria” Plum,
leaving others uninoculated. The plant was enclosed by a bell-jar,
and three weeks later twenty-three of the inoculated leaves were found to
bear on their under-surface uredo-sori of P. Pruni-spinosae, while the
control leaves showed no signs of the rust (New Phytologist, x. 207).
Arthur, in the United States, proved a similar fact, but in that case the
host of the alternate phase was Hepatica acutiloba, a very close ally
of Anemone. The xcidium is also reported on other species of Hepatica,
Anemone ranunculoides and other species of Anemone, Eranthis hiemalis,
and various species of Thalictrum.

Scribner (Report of the Dept. Agric. U.S.A. 1887) describes the Puccinia
as found on Cherry, Apricot, and Peach: it is recorded by McAlpine
on leaves, fruit and stems of Peach and Nectarine, and leaves and fruit of
Almond and Apricot.

Trichobasis Rhamni of Cooke (Seem. Journ. Bot. ii. 344, iv. 104) on
“ Rhamnus catharticus,” which was afterwards referred by him to this species
(Handb. p. 508), is probably an error due to a mistake in the identification
of the host. He states that the Puccinia on the same leaf was absolutely
identical with P. pruni-spinosae, and omits the reference in the fourth
edition of “ Microscopic Fungi.” No one else has found such a Puccinia
on Rhamnus. The leaf in “ Micr. Fung.” ed. i. 210, is no doubt a Prunus
leaf.

The mycelium of the ecidial stage is perennial ; it penetrates in spring
into the growing shoots which become deformed, the affected leaves are
narrower and paler, and the flowers are usually imperfect or altogether
wanting. These plants it is which cause fresh infections of the Plum-trees

G. U. 14

210 PUCCINIA

every year; infection by over-wintered uredospores has been proved by
Tranzschel to be possible; but as Brooks shows (J. ¢.) it is probably rare,
because the plum-leaves are generally not attected until summer is well
advanced. Fresh infections of the Anemone can, of course, be produced
by the basidiospores of the over-wintered teleutospores.

The distinction usually made, by describing the uredospores as
“ echinulate,” and the teleutospores as ‘“ verrucose,” does not convey the
exact truth; the markings on both are very similar, but the warts of
the uredospores are sharp-pointed and usually turned downwards, while
those of the teleutospores are often blunted, and always darker and more
crowded.

The brothers Sydow describe a second form of uredospore, which I
cannot find. The two cells of the teleutospore separate with the greatest
readiness, and the lower cell which is very often paler and imperfect,
could then be easily mistaken for a uredospore and has been so described
and figured. The true uredospores, mentioned in the description, are very
similar to amphispores, and have been mistaken at times for paraphyses.
The teleutospores are attached by short fragile pedicels in bunches to
a common basal cell. This is one of the characters of Arthur’s genus,
Tranzschelia.

Arthur describes (North Americ. Flora, p. 150) a second species of
Tranzschelia (P. cohaesa Long, from Texas), agreeing in almost every
minute detail with P. Pruni-spinosae, but having all its four spore-forms
upon Anemone decapetala. In its teleutospores P. fusca (q.v.) agrees
exactly with both of tHese, so that 7’. cohaesa may be regarded as a
primitive form, from which both the others have been evolved. See
Grove, New Phytologist, 1913, p. 89.

Jacky (Centralbl. f. Bakter. 2. viii. 658) divides P. Pruni-spinosae into
two forms : f. typica, in which the teleutospore has both cells alike, on the
three species of Prunus mentioned ; and f. discolor, in which the teleuto-
spore is thickened above, and the lower cell is paler, narrower and
imperfect, on Amygdalus communis and P. Persica, less often on the other
species. On P. Armeniaca both forms are found. This difference is by no
means constant, however, and is hardly worthy of mention.

DisTRIBUTION: Europe, North and South America, Africa
and Australia.

80. Puccinia Rhodiole B. et Br.

Puceinia Rhodiolae B. et Br. Ann. Nat. Hist. ser. 2, v. 462. Cooke,
Handb. p. 505 ; Mier. Fung. p. 211. Plowr. Ured. p. 207. Sacc.
Syll. vii. 701. Sydow, Monogr. i. 491, f. 401.

ON CRASSULACE 211

Teleutospores. Sori amphigenous or on the stems, but
generally hypophyllous, scattered

orcrowded and confluent, minute, e
roundish, surrounded by the torn
epidermis, pulverulent, dark- UC)

brown; spores broadly ellipsoid,
depressed, rounded at both ends,

: ; Fig. 157. P. Rhodiolae. Teleuto-
scarcely thickened above, not spores, from Glen Callater (ex
constricted, smooth, dark chest- _—_—herb. Berkeley).
nut-brown, 20—35 x 17—24 uw;
pedicels hyaline, about as long as the spore; spores occasionally
three-celled, like Triphragmium.

On Sedum Rhodiola (roseum). Very rare. Glen Callater,
July, 1844 (W. Gardiner). (Fig. 157.)

DISTRIBUTION: Norway.

81. Puccinia Umbilici Guep.

Puceinia Umbilici Guep. in Duby, Bot. Gall. ii. 890. Cooke, Handb.
p. 505 ; Mier. Fung. p. 211, pl. 4, f 80—1. Plowr. Ured. p. 204.
Sacc. Syll. vii. 700. Sydow, Monogr. i. 492, f. 403.
Teleutospores. Sori amphigenous or on the petioles, on
yellowish spots, minute, roundish,
usually circinate, at length confluent
and forming large orbicular clusters up
to lem. diam., at first compact, then
pulverulent, dark reddish-brown; spores
broadly ellipsoid or subglobose, rounded
at both ends, not thickened above but
surmounted by a minute subhyaline
apiculus or pore-cap, not constricted, smooth, bright chestnut-
brown, 28—-32 x 18—26 y, the cells often.depressed (2.e. broader
than long) and frequently oblique; pedicels short, hyaline.

Fig. 158. P. Umbilici.
Teleutospores.

On Cotyledon Umbilicus. Locally common. May and June ;
in mild localities 1t can be found even as early as January.
(Fig. 158.)

DisTRIBUTION: France, Belgium, Portugal.

14—2

212 PUCCINIA

82. Puccinia Ribis DC.

Puceinia Ribis DC. Flor. fr, ii. 221. Gard. Chron. 1894, xvi. 135.
Trans. Brit. Myc. Soc. i. 57. Sace, Syll vii. 679. Sydow,
Monogr. i. 496.

Teleutospores. Sori epiphyllous, orbicular, surrounded by a
discoloured yellow zone, circinate

(-S and often confluent, pulverulent,
rich chestnut-brown ; spores oval

Gs oe or oblong, rounded above and be-
low, apex thickened slightly and

hooded, hardly constricted, verru-

culose or rather punctate, chest-

nut-brown, 20—30 x 15—20 yp;

Bigs 108. ries or eis pedicels hyaline, thin, deciduous,

about as long as the spore; a few
mesospores intermixed.

On leaves of Ribes rubrum. Very rare. Dallas Manse
Garden, Elginshire, July 16, 1894 (Rev. Dr Keith). (Fig. 159.)

The pore of the lower cell is always towards the base, near the insertion
of the pedicel. Eriksson showed that the teleutospores do not germinate
until they have passed through the winter. He considers the form on
Ribes rubrum as biologically distinct from that on 2. nigrum or R.
Grossularia.

DisrripuTion: Central and Northern Europe, North
America.

83. Puccinia Saxifrage Schlecht.

Puccinia Saaifragae Schlecht. Flor. Berol. ii. 134. Plowr. Ured.
p. 208. Sace. Syll. vii. 678. Sydow, Monogr. i. 500. Fischer,
Ured. Schweiz, p. 151, f. 117.

P. Saxifragarum Cooke, Micr. Fung. p. 209 (non Handb. p. 506).

Teleutospores. Sori generally hypophyllous, on discoloured
spots, round, scattered or aggregated and confluent and then
irregular, soon naked, pulverulent, dark-brown ; spores ellipsoid
or oblong, rounded at both ends or slightly attenuated below,
slightly constricted, often surmounted by a rather large pale

ON SAXIFRAGACEE 2138

conical papilla, marked with faint, sometimes curved, longitu-
dinal strie, pale-brown, 26—45 x
14—20 yw; pedicels hyaline, slender,
deciduous, not as long as the spore.

On leaves and petioles of Sast-
fraga granulata, S. stellaris, S. um-
brosa. Rare. August. (Fig. 160.)

The markings on the teleutospores are Fig. 160. P. Saxifva .
perfectly invisible when wet. This species inagaanaien: ud, on Pa an a
has no connection with Caeoma Sazxifragae (Ireland), b, on S. stellaris
which is also found on S. granulata. I (Lochnagar).
have specimens from both Scotland and Ireland, the latter on 8. wmbrosa
from Clare Island, and the former on S. stellaris from Lochnagar.

Distrisution: Central and Western Europe.

84. Puccinia Pazschkei Dietel.

Puceinia Pazschkei Diet. in Hedwig. 1891, p. 108; Ber. deutsch. Bot.
Gesell. ix. 44, pl. 3, £15. Sace. Syll. xi. 185. Sydow, Monogr.
i. 503, f. 411. Fischer, Ured. Schweiz, p. 148, f. 113. Trans.
Brit. Myc. Soe. iii. 123.

Teleutospores. Sori epiphyllous, about 4—1 mm. wide,
scattered or more often in orbicular
groups 2—3 mm. diam., a few occasion- : CoN
ally hypophyllous, surrounded by the }

swollen and torn epidermis, pulverulent, (0 }

dark reddish-brown ; spores ellipsoid or

oblong, rounded at both ends, very _. ;

‘ _ : Fig. 161. P. Pazschkei. Te-
slightly thickened above or with a jeutospores and mesospore,
minute flat papilla, gently constricted, (Sutton Coldfield).
faintly and irregularly verruculose, pale clear-brown, 25—35 x
13—18 »; pedicels hyaline, short, deciduous; a few mesospores
intermixed.

On leaves of Saaifraga longifolia, Kew Gardens (G. Massee),

Journ. Bot. xlvi. 152. On a hybrid between S. Cotyledon and
S. aizoon, Sutton Coldfield, April and May, 1911—12. (Fig. 161.)

A parasite doubtless introduced into this country with the plants. In
the Sutton example, the sori form two or three perfectly round groups

214 PUCCINIA

towards the tip of each affected leaf, on the upper side, Fischer records it
on S. aizoon and 8. elatior.

DISTRIBUTION: Switzerland, Austria.

85. Puccinia Chrysosplenii Grev.

Puceinia Chrysosplenti Grev. in Engl. Flora, v. 367. Cooke, Handb.
p. 506; Micr. Fung. p. 210. Plowr. Ured. p. 211. Sace. Syll.
vii, 685. Sydow, Monogr. i. 498. Fischer, Ured. Schweiz, p. 318,

f, 231,
Teleutospores. Sori amphigenous, but generally hypo-
phyllous, small, scattered or con-

fluent, often circinate, roundish,
fo) (O} pulvinate, pale-brown; spores of two

kinds—(forma persistens) broadly

y ( 4 fusoid, strongly thickened and

more or less conical at the apex,

a rounded or slightly attenuated

below, gently constricted, smooth,

; _ very pale-brown, 82—46 x 10—

w ET ob iigoiee eae 15 p; pedicels hyaline, rather long,

Fragilipes. persistent ; (forma fragilipes) ob-

long-ellipsoid, with a conical papilla, distinctly constricted,

yellowish-brown, with faint longitudinal ridges(?), 8542 x 14—
19 w; pedicels very deciduous.

On Chrysosplenium aliernifolium, C. oppositifolium. Not
common. End of March to August or September. (Fig. 162.)

The two kinds of spores are similar in form and function to those of
P. Veronicarum ; forma persistens consists of spores which germinate
as soon as mature, forma fragzlipes of spores which rest during the winter.
It is said that the latter are scarce, but when present they form smaller
sori which are less confluent, often solitary, and are frequently found

on the upper leaf-surface. Compare the two similar kinds of spores in
P, Circaeae.

DISTRIBUTION: Central and Northern Europe, Eastern Asia.

86. Puccinia Thalictri Chev.

Puceinia Thalictri Chevallier, Flor. Paris, i. 417. Plowr. Ured. p. 206.

Sace, Syll. vii. 680. Sydow, Monogr. i. 550. Fischer, Ured.
Schweiz, p. 94, f. 72.

ON RANUNCULACEX 215

Teleutospores. Sori hypophyllous, scattered or gregarious,
often occupying the whole leaf, roundish,
soon naked, pulverulent, dark-brown ;
spores much constricted, not thickened
at the apex, the upper cell nearly
globose, the lower globose, obovoid or
clavate, generally narrower; the spores
separate readily into their component
cells, are covered with large pointed
warts, dark-brown (the lower cell paler), _, Fed

Fig. 163. P. Thalictri. Te-

26—52 x 18—30 p; pedicels hy aline, leutospores, from Prof.
deciduous. Trail’s specimens.

On Thalictrum flavwn, T. minus. Very rare; Kinloch
Rannoch, Perthshire (Prof. Trail); Kew Gardens. Autumn.
(Fig. 163.)

This species has all the marks of a perennial mycelium. The same
plants are attacked by it year after year; they are somewhat deformed
and taller, with longer internodes, smaller and paler leaves and narrower
segments. There is in Fischer a figure of a teleutospore with three cells,
looking much like that of a Phragmidium. Cf. Puccinia fusca.

DistRisuTION: Northern and Central Europe, Siberia,
North America.

87. Puccinia fusca Wint.

Aicidium fuscum Pers, in Linn. Syst. Veg. p. 1472 (teleutospores).

Puceiniz Anemones Pers, Obs. ii. 24. Cooke, Handb. p. 503; Mier.
Fung. p. 209, pl. 4, f. 65—6.

P. fusca Wint. Pilze, p. 199. Plowr. Ured’p. 205. Sace. Syll. vii. 669.
Sydow, Monogr. i. 530. Fischer, Ured. Schweiz, p. 95, f. 73.

Spermogones. Hypophyllous, mixed with the teleuto-sori,
blackish.

Teleutospores. Sori hypophyllous, rarely on the upper
side, generally spread uniformly over the whole surface of
the leaves, here and there confluent, small, round, pulverulent,
dark-brown; spores very much constricted, composed of : two
almost globose or oblong cells which easily separate, densely

216

PUCCINIA

covered with large warts, brown, 30—55 x 15—26; pedicels

Fig. 164. P. fusca.
Teleutospores,

hyaline, up to 40 long; occasionally
a few one- or three-celled spores are
intermixed.

On Anemone nemorosa. Common.
March—June. (Fig. 164.)

It has been shown by De Bary and Ed.
Fischer that the mycelium is perennial in the
rhizome. The attacked plants are deformed
and never flower; they bear paler and narrower
leaves which are much thickened. The ecidia

on the same host are not connected with this species (see Ochropsora Sorbi
and P. Pruni-spinosae) ;, in fact they do not appear until some time later
than the teleutospores of P. fusca begin to show,

DisTRIBUTION: Europe, Siberia, North America.

88. Puccinia Calthe Link.

Aicidium Calthae Grev. Flor. Edin. p. 446. Cooke, Handb. p. 539 ;
Micr. Fung. p. 196.

Puccinia Calthae Link, Sp. Plant. ii. 79. Cooke, Handb. p. 504;
Micr. Fung. p. 210. Plowr. Ured. p. 145. Sacc. Syll. vii. 602.

Sydow, Monogr. i.

540. Fischer, Ured. Schweiz, p. 310, f. 225.

Spermogones. In little clusters, honey-coloured.
4Aicidiospores. Aicidia hypophyllous, in little clusters on

GD

O
L

Fig. 165. P. Calthae.
Teleutospores,

roundish yellowish spots, or on the
stems in elongated swellings, cup-
shaped, with a torn whitish recurved
margin; spores delicately verrucu-
lose, orange, 21—28 p.

Uredospores. Sori generally hy-
pophyllous, minute, scattered, round-
ish, pulverulent, chestnut; spores
globose to ellipsoid, echinulate, pale-
chestnut, 22—30 x 20—25 w, with

two germ-pores in the upper half.

Teleutospores. Sori amphigenous, small, irregularly scat-
tered or often circinate, pulverulent, but persistent, black-
brown; spores oblong-clavate or fusoid, generally with a paler

ON RANUNCULACEE 217

conical papilla at the apex, hardly constricted, perfectly smooth,
clear chestnut-brown, 30—44 x 13—22 %; pedicels hyaline,
thick, persistent, up to 75 long.

On Caltha palustris. Rather rare. icidia, May and June ;
teleutospores, July—October. (Fig. 165.)

There is another British species on Caltha palustris, P. Zopfii, which
has been usually confounded with the present one in herbaria. It differs
in having its teleutospores broader, darker, more oblong, and covered here
and there with minute warts, but is otherwise similar in appearance. There
are three others found on species of Caltha in North America, all so nearly
allied that they are difficult of discrimination; but of these one has
no uredospores, and the others have, so far as is known, teleutospores only.
Some of these may be found in Britain.

DISTRIBUTION : Europe, Siberia, North America:

89. Puccinia Zopfii Winter.

Puccinia Zopfii Wint. in Hedw. 1880, p. 39, 107. Sydow, Monogr.
i. 542. Fischer, Ured. Schweiz, p. 91, f. 70.

Spermogones. Amphigenous or on the petioles, in little
clusters of 6—10, brownish when old. ;

4icidiospores. Aicidia hypophyllous or on the petioles,
usually surrounding the groups of spermogones in scattered

Fig. 166. P. Zopfii. Leaf of Caltha with ecidia ; two teleutospores.

roundish clusters 1—2 mm. diam. (elongated on the petioles),
at first hemispherical, then shortly cup-shaped, flattish, yellow,
with a short torn scarcely reflexed margin; spores delicately
verruculose, yellow, 20—28 pu.

218 PUCCINIA

Uredospores. Sori generally hypophyllous, minute, scattered,
punctiform, round, soon naked, surrounded by the erect epidermis,
chestnut; spores ellipsoid, echinulate, brownish-yellow, 22—30 x
20—25 yu, with two or three germ-pores.

Teleutospores. Sori similar, but darker; spores oblong to
obovate, rounded at both ends, sometimes truncate above or
slightly narrowed below, scarcely thickened but with a broad
flat papilla at the apex, gently constricted, delicately verruculose,
dark chestnut-brown, 35—60 x 24—35 yu ; pedicels nearly hyaline,
short, deciduous.

On Caltha palustris. Rather rare; Shropshire, Scarborough,
Rhydd-y-fen, Gullan Loch, Ireland (co. Dublin). Aicidia in
May; teleutospores, August—December (June—September,
Fischer). (Fig. 166.)

Distinguished from P. Calthae essentially by its teleutospores which
are provided with a few, rather distant, minute warts, mostly towards the
upper end ; these are difficult to see except when viewed dry. The spores
are also relatively much broader and not narrowed towards the summit,
and are therefore easily seen to be distinct in shape; they are darker in
colour and have shorter pedicels. The ecidia are not known to be different
from those of P. Calthae; those described above probably belong to
P. Zopfii, because they were found in the same neighbourhood as the
teleutospores in Ireland: the cells of the peridium agreed with those
figured by Fischer (J.c.). Krieg showed (Centralbl. f. Bakt. 2. xv. 259)
that P. Zopfii is autecious, like P. Calthae.

The two species have been frequently confounded in herbaria ; but, if
I may judge by the specimens I have seen, the teleuto-sori of P. Calthae
are larger, more crowded, more often circinate, more compact, and remain
longer covered by the epidermis than those of P. Zop/iz, though this is not
always so well marked.

DISTRIBUTION: Central Europe.

90. Puccinia Lychnidearum Link.

Puceinia Lychnidearum Link, Sp. Pl. ii. 80 (1825), Cooke, Handb.
p. 505 ; Micr. Fung. p. 210 p.p. Plowr. Ured. p. 196.

P. Arenariae Winter, Pilze, p. 169 (1884), Plowr. Ured. p. 210. Sace.
Syll. vii. 683. Sydow, Monogr. i. 553. Fischer, Ured. Schweiz,
p. 307, f, 224. McAlpine, Rusts of Australia, p. 177, f. 97
(introduced on Stel/aria media).

ON CARYOPHYLLACEZ 219

P. Sperguliae DC. ; Cooke, Micr. Fung. p. 210. Sydow, Monogr. i. 560.
P. Moehringiae Fckl. ; Cooke, Micr. Fung. p. 210.

Teleutospores. Sori hypophyllous or rarely on the stems,
scattered or circinate, on pale spots, sometimes confluent,
pulvinate, pallid-brown, then darker, greyish-pulverulent from
the numerous basidiospores; spores oblong-fusoid or clavate,

/

Fig. 167. P. Lychnidearum. Fig. 168. P. Lychnidearum.
Teleutospores, on Lychnis Teleutospores, on Arenaria
diurna. trinervis.

rounded or somewhat pointed above and more or less thickened
(up to 10 4), gently constricted, rounded or attenuated below,
smooth, yellowish-brown, 30—50 x 10—20 4; pedicels hyaline,
persistent, 60—85 w long.

On various Caryophyllacee, such as Dianthus barbatus,
Lychnis diurna, L. vespertina, Arenaria trinervis, Gypsophila
elegans, Sagina nodosa (?), Spergula arvensis (?), Stellaria Holo-
stea, S. media, S. uliginosa. May—November. It is most
common on Lychnis diurna, on which as also on Dianthus the
sori are remarkably circinate. (Figs. 167, 168.)

The two species, P. Lychnidearum and P. Arenariae, are united in
Sydows’ Monographia, on the ground that our present knowledge, derived
from cultures, is insufficient to separate them, and that any apparent
morphological distinctions break down completely when 4 long series of

220 PUCCINIA

specimens is examined. The spores are, however, variable in form and
colour, and the sori differ in appearance and arrangement ; no doubt the
future will find this Puccinia divided into several biological races, if not
into distinct species. Meanwhile, on morphological grounds alone, our
British forms may be arranged under three heads :

1. forma Lychnidearum (Link), on Lychnis; sori medium-brown in
colour, often greyish, remarkably circinate, on conspicuous yellow
and purple spots.

2. forma Dianthe (DC.), on Dianthus; sori larger, darker, and more
pulvinate, usually somewhat circinate; =P. Dianthi DC.

3. forma Arenariae (Schum.), on Arenaria and Stellaria ; sori paler,
not so circinate, spores paler ; =P. Moehringiae Fckl.

The form on Sagina procumbens is so different that it is here reckoned
as a separate species, P. Saginae K. et 8. (q.v.). It will be noticed that
Plowright separated P. Lychnidearum as a distinct species from P.
Arenariae, chiefly on the ground that he considered the former to possess
uredospores. It has been observed by all, however, that the common
form on Lychnis diurna has no uredospores in its very abundant sori ; it
is therefore satisfactory to find that in Plowright’s herbarium there are
several leaves of Lychnis diurna, gathered by W. Phillips at Aberystwyth
apparently on one occasion only (July 1873), on which are uredospores
mixed with very few teleutospores. These he named P. Lychnidearum,
but examination shows that they have no connection with the common
fungus met with everywhere on the same host but belong to P. Behenis
Otth. See under that species.

The Puccinia recorded by Plowright (/.c. p. 210) on Spergula arvensis
may be a distinct species. See Cooke, Micr. Fung. p. 210. I have seen
no specimens.

When mature, the cells of the teleutospores of P. Lychnidearum
separate with great ease ; they germinate readily while still in the sori,
and the numerous basidiospores produced give them a greyish look, as
happens also in other Lepto-species. I have specimens on Arenaria
trinervis gathered in full germination at the end of May. During the
process the spores become denticulate at the summit; in the case
mentioned 75 °/, of the spores were in this state. Such spores have some-
times been wrongly described as having digitate processes like those of
P. coronata, — It was stated by De Bary that he had seen the germ-tubes
of the basidiospores of P. Dianthi enter the host-plant through the
stomata: no similar case has been detected by any other observer (see
p. 38).

DISTRIBUTION : Europe, Siberia, East Indies, North and
South America.

ON CARYOPHYLLACEX 221

91. P. Sagine K. et S.

Puccinia Saginae K. et S. Exsice. no, 221. Fuckel, Symb. Myce. p. 51.

P. Arenariae Wint.; Plowr. Ured. p. 210 p.p. Sydow, Monogr.
1. 553 p.p.

P. Lychnidearum Link; Cooke, Handb, p. 505; Micr. Fung. p. 210 p.p.

Teleutospores. Sori chiefly on the stems, compact, confluent,
forming blackish-brown masses, about 4 cm. long, encrusting at
intervals the flowering stems which are slightly thickened at
that spot, rarely on the leaves; spores oblong to oblong-clavate,

a

Fig. 169. P. Saginae. Teleutospores, from the Ercal specimen.

rounded and not thickened above, slightly constricted, usually
somewhat tapering below, smooth, very dilute brown, almost
hyaline, 36—41 x 15—19 ; pedicels hyaline, rather stout,
persistent, about as long as the spore.

On Sagina procumbens. Rare; specimen in Herb. Phillips
(Brit. Mus.) from Ercal, near Wellington, under the name
Puce. Lychnidearum. (Fig. 169.)

The spores of this species, as well as the character of the sori, render
it widely distinct from the various forms included under the name
P. Lychnidearum ; the most obvious difference ig in the total want of
thickening at the apex, unless this was due merely to the fact that all the
spores had germinated, which did not seem to be the case.

222 PUCCINIA

92, Puccinia Behenis Otth.
Aeeidium Behenis DC. Flor. fr. vi. 94 p.p.
Puceinia Behenis Otth, Mitth. Naturforsch. Gesell. Bern (1870), p. 89.
Fischer, Ured. Schweiz, p. 136, f. 103.
P. Silenes Schrét. in Winter, Pilze, p. 215 (1884). Cooke, Micr. Fung.
p. 211. Plowr. Ured. p. 147. Sacc. Syll. vii. 605. Sydow,
Monogr. i. 559.

Spermogones. In little clusters, honey-coloured.
Micidiospores. Aicidia hypophyllous, on pallid-yellow spots,

a

Fig. 170. P. Behenis. a, Teleutospore and mesospore, on Silene inflata;
b, uredospore and teleutospore, on Lychnis diurna.

in orbicular clusters, minute, shortly cylindrical, with whitish
torn margin; spores delicately verruculose, orange, 17—26 x
14—20 pw, or 15 diam.

Uredospores. Sori amphigenous, scattered or circinate,
sometimes confluent, on paler spots, minute, cinnamon-brown ;
spores subglobose to ellipsoid, echinulate, pale-brown, 20—26 x
17—22 yw, with three or four germ-pores.

Teleutospores. Sori similar, but black-brown ; spores oblong
to ellipsoid, rounded at both ends, faintly constricted, surmounted
by asmall pale apical papilla, smooth, chestnut-brown, 25—40 x
16—26 yw; pedicels hyaline, short, deciduous.

On Silene inflata (=8. latifolia, 8. Cucubalus) and Lychnis
diurna (dioica). Not common. Uredo- and _teleutospores,
July—October. (Fig. 170.)

The description of the spermogones and acidia is partly after Schroter,
Sydow, and Fischer. The distinction of the ecidium of this species from
that belonging to Uromyces Behenis is as follows :—The former has sper-
mogones and uredospores, which accompany and follow it, but it is never
accompanied by teleutospores. The early xcidia of the Uromyces are
situated on purplish spots which show above, the later ones are usually

ON POLYGONACEX 223

scattered singly : those of the Puccinia are in largish orbicular clusters
and are rarely found singly ; if not clustered, they spread over the whole
leaf. In Plowright’s herbarium are some leaves of Lychnis diurna, covered
with uredo-sori, which he mistakenly assigned to P. Lychnidearum : there
are no teleutospores of the latter, however, but a very few of P. Behenis
were found in the same sori, See under P. Lychnidearum. The circinate
arrangement of the sori, on paler spots, is very similar in both species.

DistRiBuTIoN: Central and Western Europe.

93. Puccinia Acetose Korn.

Uredo Acetosae Schum. Plant. Sail. ii. 231.

Puceinia Acetosae Korn. in Hedwig. 1876, p. 184. Plowr. in Trans.
Brit. Myc. Soc. i. 57. Sace. Syll. vii. 688. Sydow, Monogr.
i. 581. Fischer, Ured. Schweiz, p. 134, f. 101.

Uredospores. Sori amphigenous, scattered, minute, roundish
on the leaves, elongated on the
petioles and stems, soon naked,
ferruginous-brown; spores globose
to obovate, sparsely aculeate,
brownish, 24—30 x 20—28 yw, with
two (rarely three) germ-pores.

Teleutospores. Sori similar, but
dark-brown; spores ellipsoid, ob-
long, or subclavate, rounded at both ends or slightly attenuated
below, not thickened at the apex, but furnished with a broad
pore-cap, slightly constricted, delicately verruculose, chestnut-
brown, 28—42 x 19—24 4; pedicels hyaline, slender, deciduous,
as much as 35 wu long. ,

Fig. 171. P. Acetosae.
Teleutospores and uredospore.

On Rumex Acetosa. Malden, Yorkshire, July 16, 1894
(A. W. Saunders), Ireland, co. Antrim (J. Adams), August,
1909. Bewdley, Worcestershire, August, 1907, ete. (Fig. 171.)

There is no doubt that this species is often mistaken for Uromyces
Acetosae. In the absence of the two-celled teleutospores, which are rare,
it could be distinguished mainly by its smaller sori, and more spiny
uredospores (the spines are quite colourless). The teleutospores show the
delicate warts more clearly on the upper cell ; in the Bewdley specimens
they were perceptible only with great care. Magnus proved that this
species can pass the winter by its uredospores. It may be heterccious.

224, PUCCINIA

Disrrisution: Central and Northern Europe, Siberia,
North America.

94. Puccinia Oxyrie Fckl.

Puccinia Oxyriae Fckl. Symb. Nacht. iii. 14. Cooke, Grevillea, xi. 15.
Plowr. Ured. p. 194. Sace. Syll. vii. 642. Sydow, Monogr. i. 567.
Fischer, Ured. Schweiz, p. 135, f. 102.

Uredospores. Sori amphigenous, generally hypophyllous,
on minute purplish spots, scattered or aggregated, rounded,
sometimes confluent, surrounded by the cleft epidermis, cinna-
mon; spores globose to ovate, delicately echinulate, yellow-
brown, 23—30 x 20—26 p.

m

Fig. 172. P. Oxyriae. Teleutospores (from a Swiss specimen).

Teleutospores. Sori on the leaves similar, but also on the
petioles and peduncles and then elongated, pulverulent, black-
brown; spores oblong to obovate, rounded and slightly thick-
ened or hooded above, constricted, generally rounded below,
marked in the upper part with little unevennesses, almost as if
corroded, brown, 30—46 x 15—25 uw; pedicels nearly hyaline,
deciduous, rather long.

On Oxyria digyna (reniformis). Rare; Skye (Buchanan
White), Braemar (Prof. Trail). August. (Fig. 172.)
The teleutospores were described by the older authors as smooth ; but,

as Lagerheim first pointed out, they are really provided with faintly
perceptible markings, especially visible towards the summit.

DisTRIBUTION : Switzerland, Norway, Iceland, Colorado.

ON POLYGONACE 225

95. Puccinia Conopodii-Bistorte Kleb.

Puccinia Bistortae DC. Flor. fr. vi. 61. Cooke, Micr. Fung. p. 204;
Grevillea, ii. 161. Plowr. Ured. p. 192 (7). Sydow, Monogr.
i. 571 (all pro parte). Soppitt, Grevillea, xxii. 45 ; Gard. Chron.
1895, xviii. 773.

P. Conopodii- Bistortae Kleb. Wirtsw. Rostp. p. 318.

Acidiospores. AKcidia on large swellings of a bright orange
colour, immersed, spherical or flat, not at all cup-shaped, margin
not projecting; spores delicately verruculose, orange, 15—20 y.

Uredospores. Sori hypophyllous, minute, roundish, yellowish-
red, soon naked; spores globose to shortly ellipsoid, finely
echinulate, pale yellowish-brown, 21—24 w.

Teleutospores. Sori hypophyllous, scattered or united in
roundish groups, soon naked and pulverulent, dark-
brown; spores oblong or subclavate, rounded at ae
both ends or slightly oblique at the apex, faintly (
constricted, clear yellowish-brown, 28—42 x 16— ey)
25; epispore equally thick, smooth, but some-
times marked with a very few longitudinal or 5, 178.
oblique rows of delicate warts; no papilla. on P. Conopodii-
the germ-pores; pedicels hyaline, short, very tee
deciduous.

ficidia on Conopodiuwm denudatum, Yorkshire, May; teleuto-
spores on Polygonum Bistorta, June—August, not common.

(Fig. 173.)

The Puccinias on Pol. Bistorta are not yet well known. There appear to:
be at least four (or five) distinct forms, which are divisible morphologically
into two groups—(1) those whose teleutospores have no apical papilla,
(2) those which have a small hemispherical papilla at the summit; the
latter are called by Sydow in the Monographia P. mammillata Schrét.
(Pilz. Schles. p. 340). They are all hetercecious.

Of the former group, one form has an ecidium on Conopodium, and its
other spore-stages on P. Bistorta. Itis the one mentioned above ; the life:
history was first demonstrated by Soppitt (2.¢.). It is remarkable as
being the first instance known of a hetercecious Puccinia that had its
teleutospores on a Dicotyledon. The second form has its ecidium on
Angelica silvestris and Carum Caruz and its teleutospores on Pol. Bistorta
and Pol. viviparum. It would perhaps be better divided into P. Angelicae-
Bistortae Web. (= P. Cari-Bistortae) and P. Polygoni-vivipari Karst. The
connection in the former has been demonstrated by Klebahn and Fischer ;

G.U. 15

226 PUCCINIA

as regards Pol. viviparum Klebahn expresses doubts, but Semadeni was able
successfully to infect that species with uredospores from Pol. Bistorta, The
form on Pol. viviparum is here kept provisionally distinct, since the
eecidium on Angelica has not been found in Britain.

‘Of.the second group, P. mammillata, there are two biological races—
(1) P. Mei-mammillata Sernadeni, on Meum, and (2) P. Angelicae-mam-
millata Klebahn, on Angelica. Neither of these has been found in
Britain. All the four of these are closely allied ; in Sydows’ Monographia
it is suggested that possibly in all these cases the acidium on the
Umbellifer is merely facultative and the Puccinia can maintain itself
without that aid.

96. Puccinia Polygoni-vivipari Karst.
Puccinia Bistortae DC. ; Cooke, Grevillea, ii, 161 ; Mier. Fung. p. 204
pp. Plowr. Ured. p. 192. Sace, Syll. vii. 688. Sydow, Monogr.
i. 571 p.p.
P. Polygoni-vivipart Karst. Enum. Fung. Lapp. p. 221. Fischer,
Ured. Schweiz, p. 100, f. 76 (%).

Uredospores. Sori hypophyllous, scattered, small, roundish,
cinnamon-brown, girt by the erect epidermis; spores roundish
to ellipsoid, finely echinulate, pale-brown, 20—23 x.16—17 p.

Teleutospores. Sori similar, but blackish-brown; spores
elliptical to obovate-oblong, rounded and not thickened above,

A506

\V

Fig. 174. P. Polygoni-vivipari. a, two teleutospores from Mar Lodge;

b, uredo- and teleutospore from a specimen issued by Fuckel, Symb. Myc.

57; both on Polygonum viviparum.
hardly or not at all constricted, rounded or slightly tapering
below, smooth, brown, 20—30 x 15—20; epispore very thin and
translucent; no papilla on the germ-pores; pedicels deciduous,
short.

On Polygonum viparum. Very rare. Near Mar Lodge,
August, 1822 (Dr Greville). Braemar, August, 1882 (Prof.
Trail). (Fig. 174.)

This species is closely allied to P. Conopodii-Bistortae (q. v.), but in
the absence of all biological information is best kept distinct. I have —

ON POLYGONACEE 227

seen no British specimens but those collected at Mar Lodge and Braemar,
which are described above ; the spores of these agree with those of a
specimen on Pol. viviparum issued by Fuckel (see Symb. Myc. p. 57). It is
a purely Alpine species.

97. Puccinia Polygoni-amphibii Pers.
Medium Geranii DC. ; Cooke, Handb. p. 543; Micr. Fung. p. 199

p.p.(2).
4. sanguinolentum Lindr. Myk. Notiz. in Bot. Notis. 1900, p. 241.
Uredo Polygonorum DC. ; Grev. Sc. Cr. Fl. pl. 80.
Trichobasis Polygonorum Berk. ; Cooke, Micr. Fung. p. 226 p.p.
Puccinia Polygonorum Link ; Cooke, Handb. p. 495; Micr. Fung.

p. 203 p.p.
P. Polygoni Pers. ; Plowr. Ured. p. 188 p.p. Sacc. Syll. vii. 636 p.p.
P. Polygoni-amphibit Pers. Syn. p. 227; Sydow, Monogr.'i. 569 p.p.
Fischer, Ured. Schweiz, p. 301, f. 220.

[Spermogones. Few, amphigenous.

4icidiospores. Aicidia hypophyllous, mostly in concentric
groups, on well-marked spots which
are deep-red or purplish and often
surrounded by a conspicuous green- I
ish-yellow zone, sometimes occupying (0)
the greater part of a leaf, cup-shaped,
with a much cut recurved margin;
spores finely punctate-verruculose,
yellowish, 18—28 p.]

Uredospores. Sori amphigenous,
more often hypophyllous, scattered, i
roundish, soon naked, pulverulent,
brownish; spores ellipsoid to obovate, __ ; ;
faintly echinulate, yellowish-brown, Be a ree oes
25—28 x 18—21 pw, with two germ- — mphibium.
pores in the upper half.

Teleutospores. Sori chiefly hypophyllous, not projecting,
long covered by the epidermis, surrounding the uredo-sori in a
circular fashion, dark-brown; spores oblong to clavate, rounded
and thickened above (5—12 y), often obliquely conical at the
apex, more or less tapering below, gently constricted, smooth,
yellow-brown, 85—52 x 16—22,; pedicels nearly hyaline,
persistent.

15—2

228 PUCCINIA

[Acidia on Geranium pratense, G. silvaticum ;] uredo- and
teleutospores on Polygonum amphibium, P. lapathifolium, July—
October; not uncommon. Klebahn and Bubdk report the
ecidium on G. molle, G. phaeum and other species. (Fig. 175.)

Teleutospores are found not only in the separate sori described above,
but also in the old uredo-sori. These latter spores are rounded at the
apex ; those which grow in distinct sori often have the thickening cap
forced to one side, presumably by the pressure of the persistent epidermis.
For the distinctions of the ecidium of this Puccinia from that of Uromyces
Geranii which grows upon the same hosts, see under that species (p. 104).

The proof, of the connection of the zcidium with the Puccinia was first
given by Tranzschel in Russia, and has since been confirmed by Bubak. I
have not seen any British specimens of Meidium sanguinolentum ; the
description given above is taken from Lindroth. Similarly in Switzerland,
Fischer records only the uredo- and teleutospores. On Polygonum
amphibium they seem to be confined to the terrestrial form : I have never
seen them on the floating leaves.

DISTRIBUTION : World-wide.

98. Puccinia Polygoni-Convolvuli DC.
Puceinia Polygont-amphibii Pers. ; Sydow, Monogr. i. 569 p.p.
P. Polygoni-Convolvult DC. Flor. fr. vi. 61.
P. Polygont Plowr. Ured. p. 188 p.p. Sace. Syll. vii. 636 p.p.
Fischer, Ured. Schweiz, p. 303, f. 221.
Spermogones and dicidiu. Presumably similar to those of
the preceding species.
Uredospores. Sori hypophyllous, roundish, soon naked,
brown; spores ellipsoid to obovate,

echinulate, clear yellowish-brown, 24—
Co 30 x 15-20 w, with two opposite

germ-pores.
Teleutospores. Sori hypophyllous,

scattered, roundish, compact, pulvinate,

about 4mm. diam., soon naked and

pulverulent, black; spores oblong to

F eae, iaeaee clavate, rounded or occasionally conical

herb. Cooke). vbove and thickened (up to 8), faintly

constricted, gently attenuated below,

smooth, brown, 32—45 x 18—21 w; pedicels clear yellow-brown,
persistent, nearly as long as the spore.

ON THESIUM 229

fKeidia on Geranium pusillum, and possibly also on G. molle,
G. rotundifolium; uredo- and teleutospores on Polygonum Con-
volvulus, August and September. Uncommon. (Fig. 176.)

The connection of the «cidium on the first-named host and the
Puceinia on Pol. Convolwulus has been experimentally demonstrated by
Tranzschel. It is possible that the same parasite also attacks P. dume-
torum, P. Persicaria and others. The ecidium is not known for certain to
have occurred in Britain. The teleuto-sori of P. Polygoni-Convolvuli are
distinguished from those of P. Polygoni-amphibii by their compact
pulvinate form, and by being soon uncovered by the epidermis, while the
spores (perhaps in consequence of that) are much darker at the summit,
and the apex, if conical, is less often oblique. According to Sydow these
distinctions, however true they may be of the European forms of the
species, do not avail when the extra-European forms are considered. In
the Monographia, therefore, the two species are united, and only culture
experiments will be able to decide the question.

99. Puccinia Thesii Chaill.

Geidium Thesii Desv. in Journ. de Bot. ii. 311 p.p. Cooke, Handb.
p. 587; Micr. Fung. p. 195, pl. 3, f. 50—1.

Puceinia Thesii Chaill. in Duby, Bot. Gall. ii. 889. Cooke, Handb.
p. 495 ; Micr. Fung. p. 204. Plowr. Ured. p. 145. Sacc. Syll.
vii. 602. Sydow, Monogr. i. 585. Fischer, Ured. Schweiz, p. 300,
f. 219.

Spermogones. Amphigenous, numerous, amongst the ecidia.

Fig. 177. P. Thesii. Teleutospores, from a Surrey specimen.

Aicidiospores. Aicidia amphigenous, scattered uniformly
and rather thickly over the whole leaf-surface, seldom in roundish

230 PUCCINIA

or oblong groups, between cylindrical and cup-shaped, with a
white torn recurved margin; spores delicately verruculose,
orange, 16—24 p.

Uredospores. Sori amphigenous or on the stems, distributed
irregularly, minute, roundish, long covered by the epidermis,
brown ; spores globose to broadly ellipsoid, verruculose, yellow-
brown, 20-—28 p, with four or five germ-pores.

Teleutospores. Sori similar, but more compact, brown-black ;
spores oblong to clavate, generally rounded above and slightly
thickened, hardly or not at all constricted, rounded or attenuated
below, smooth, uniformly brown, 35—54 x 16—24,; pedicels
brownish, thick, not very persistent, short or as much as 95 w
long; a few mesospores intermixed.

On Thesium humifusum. Rare; Surrey, Dorset, Wilts.,
Hants., Cambs., ete. Aicidia, May—August (also recorded for
October); teleutospores, August—October. (Fig. 177.)

It was maintained by Vuillemin (Bull, Soc. Myc. France, 1894, p.
107 ff.) that this species has no xcidium-stage, the Heidiwm Thesit being
referred by him to another species, P. Desvauwit (= P. Passerinii),
distinguished by its broader and warted teleutospores ; but both Sydow
and Fischer have disproved this contention. The ecidia are rather more
common than the teleutospores, and are sometimes found among the
teleuto-sori on the same leaf, in October.

DISTRIBUTION: Europe, Eastern Siberia.

100. Puccinia Iridis Wallr.

"redo Iridis DC. Encyel. viii. 224. Plowr. Ured. p. 257.

Trichobasis Iridis Cooke, Micr. Fung. p. 227.

Puccinia Iridis Wally. in Rab, Krypt. Fl. ed. i. p. 23 (1844). Plowr.
Ured. p. 189. Sace. Syll. vii. 657. Sydow, Monogr. i. 598.
Fischer, Ured. Schweiz, p. 236, f. 186.

P. truncata B. and Br. Ann. Nat, Hist. ser. 2, xiii, 461 (1854). Cooke,
Handb. p. 494 ; Micr. Fung. p. 208.

Uredospores. Sori amphigenous, solitary or somewhat
aggregated, roundish or elongated, minute, long covered by the
epidermis, not pulverulent, reddish-brown; spores globose to

ON IRIS ° 231

ovate, ochraceous-brown, echinulate, 20—35 x 16—26 w; epi-
spore thick, with 2—4 (or more) germ-pores.

Teleutospores. Sori hypophyllous, few and irregularly scat-
tered, sometimes confluent, oblong,
compact, persistent, soon naked,
black; spores clavate or oblong,
much thickened (up to 14) at the
summit and rounded, less often
conical or truncate, gently constric-
ted, usually attenuated below,
smooth, fuscous-brown, darker above,

30—52 x 14-224; pedicels brown-
ish, thick-walled, persistent, about \
as long as the spore.

On Iris foetidissima, I. Pseuda-
corus, and on many cultivated Fig. 178. P. Iridis. Teleuto-
species of Iris. Not common. SEGRE) 00 Ae aeteaeae Tas
May—October; the teleutospores
may be found on the old leaves until the following spring.
(Fig. 178.)

The uredo-stage is most abundant and assumes various characters,
especially as regards the formation of spots; when present these are
conspicuous, oblong or oval, and yellowish, often with a greenish-brown
circumference. In that case the leaf looks remarkably variegated.

Plowright considered that the form which occurs on our cultivated
Irises is different from that on our native species, because he could not
find any teleutospores in the former ; other authors consider them as the
same, because the teleutospores on many species are difficult to find, and
appear only on dying leaves, especially towards the base, at the end of the
season. One can easily recognise them by their being naked ; for in this
species, contrary to the usual state of things, the uredo-sori remain long
concealed by the epidermis and the teleuto-sori soon become uncovered.
The uredospores are very thick-walled ; they can survive the winter and
reproduce the fungus in the spring.

This species might be hetercecious : no experimental cultures appear
to have been made. It will probably turn out to possess several biological
races, for it has been recorded on more than thirty-five species of Iris.

DISTRIBUTION: Europe, Asia, North America.

232 PUCCINIA
101. Puccinia Schroeteri Pass.

Puccinia Schroeteri Pass. Nuov. Giorn. Bot. Ital. vii. 255. W. G.
Smith, Gard. Chron. 1889, v. 725, f. 118. Wolley-Dod, Journ.
Roy. Hort. Soc. xii. p. liti, Plowr. Trans. Brit. Myc. Soc. 1. 57.
Sacc. Syll. vii. 732. Sydow, Monogr. i. 608. Fischer, Ured.
Schweiz, p. 78, f. 59.

Teleutospores. Sori amphigenous, chiefly epiphyllous, large,
oblong or elliptic, surrounded by a brownish-violet discoloration,
1—3 mm. long, solitary or in small clusters, long covered or
half uncovered and surrounded by the lead-coloured epidermis,

Fig. 179. P. Schroeteri. a, meso- and teleutospore, from the original specimen
of W. G. Smith; b, teleutospore and sori on leaf of daffodil, from Cooke’s
specimen. : «

blackish-brown; spores ellipsoid or oblong, rounded at both
ends, not thickened above, hardly constricted, obscurely reticu-
lated, golden-brown, then chestnut, 40—60 x 25—29 w; pedicels
hyaline, short, thick, deciduous ; mesospores also occur.

On Jonquil and Narcissus poeticus. Very rare; Malpas,
May, 1889 (Rev. C. Wolley-Dod); in Gard. Chron. and Journ.
Roy. Hort. Soc. (/.c.) it is stated to have been found also on the
“common double Narcissus” (? N. telamonius plenus). There
are some specimens in Herb. Brit. Mus. on daffodil leaves, sent
to the Gard. Chron. by a correspondent, May, 1894, and labelled
P. Lnliacearwm by Cooke, which on examination prove to be
this species. (Fig. 179.)

ON LILIACEA 233

é

Plowright observed that the spores would not germinate at once, but,
by securing the affected leaves during the winter near some plants of
NV. poeticus, he found the Puccinia reproduced next year and for eight
or nine years afterwards, though only on the tips of the leaves. The
reticulation of the spores varies in character, sometimes resolving itself
into longitudinal ridges or rows of warts. Mesospores and other abnormal
spores are recorded by Fischer.

DistRiBuTION: Belgium, Italy, Garniola.

102. Puccinia Asparagi DC.

Puccinia Asparagi DC. Flor. fr. ii. 595. Cooke, Handb. p. 494 ; Mier.
Fung. p. 203. Plowr. Ured. p. 144. Sace. Syll. vii. 601. Sydow,
Monogr. i. 615. Fischer, Ured. Schweiz, p. 235, f. 185.
Spermogones. In little clusters, honey-yellow.
Aicidiospores. ALcidia in oblong groups on the stems, for a
long time closed, then shortly cup-shaped, with a whitish, erect,
torn margin; spore delicately verruculose, orange, 15—28 p.

é
Fig. 180. 2. Asparagi. Teleutospores and mesospore.

Uredospores. Sori oblong, narrow, flat, long covered by the
epidermis, cinnamon-brown; spores globose to ovate, delicately
and densely echinulate, pale-brown, 20—30 x 17—25 pw, with
four germ-pores. °

234 PUCCINIA

Teleutospores. Sori oblong or linear, often confluent,
generally on the stems, rarely on the leaves (phylloclades),
blackish-brown ; spores ellipsoid to clavate, thickened (up to
8 w) and rounded above, very gently constricted, rounded below,
smooth, brown, 35—52 x 17—26 yw; pedicels hyaline or brown-
ish, persistent, as long as or longer than the spore; a few
obovate mesospores intermixed.

On Asparagus officinales. Aicidia, not common, May;
uredo- and teleutospores, September—December, rather fre-
quent. (Fig. 180.) ‘

Fischer points out that the connection between the ecidium and the
other spore-forms has not yet been demonstrated. This disease is often
very destructive to asparagus beds ; all diseased shoots should be gathered

and burnt. The best means of prevention is by the selection of resistant
varieties, and the avoidance of overcrowding.

DIsTRIBUTION: Europe, Abyssinia, North America.

103, Puccinia Liliacearum Duby.

Puccinia Liliacearum Duby, Bot. Gall. ii. 891. Plowr. Ured. p. 197.
Sace, Syll. vii. 668. Sydow, Monogr. i. 627. Fischer, Ured.
Schweiz, p. 76, 545, f. 57.

Spermogones. Numerous, especially at the apex of the

affected leaves, yellowish, conical.

[dicidiospores. Aicidia few, scattered, alae deeply im-
mersed, whitish, the narrow opening only pro-
jecting; spores minutely verruculose, orange,
about 15—20 p.]

Leleutospores. Sori amphigenous, embedded
in the dried yellowish parts of the leaf, hemi-
spherical, densely crowded, often confluent, long
covered by the ashy-grey epidermis which at
length opens by a cleft, then naked, pulverulent,
reddish-brown; spores oblong-fusiform or cla-
vate, not thickened above and rounded or fre-
quently somewhat narrowed, not constricted,
Fig. 181. P. indeed broadest at the septum, attenuated below,
Toate smooth, pallid-brown, 40—75 x 22—35 w; pedi-

(Lytham). cels hyaline, thick, rather long.

ON LILIACEA 235

On Ornithogalum umbellatum. Rare; Lytham and near
Carlisle (Rev. Hilderic Friend). March—May. (Fig. 181.)

As usual, only spermogones and teleutospores are present on these
specimens. Fischer records that he found the fungus on O. umbellatum in
March and April in great plenty, with spermogones and teleutospores, but
no ecidia ; the infested parts of the leaves were swollen, compact, and
harder than the healthy portions. He suggests (Centralbl. f. Bakt. 2. xv.
230) with great probability that the zxcidia with spermogones which are
also found on Ornithogalum (Afcidium ornithogaleum Bubdk, Annal. Myc.
ili, 222) belong to some heterecious species. The fungus on 0. umbellatum
is a biologic race; it does not attack O. nutans, much less other allied
species of Liliacez, such as Muscart and Hyacinthus, although P. Lilia-
cearum is recorded on them. Fischer adds that the mycelium is not
perennial, but infection takes place afresh each spring by the basidiospores
of the overwintered teleutospores, from leaves lying on the ground.

DIsTRIBUTION : Central Europe.

104. Puccinia Porri Wint.

Credo Porrt Sow. Engl. Fl. pl. 411.

U. Alliorum DC. Flor. fr. vi. 82 p.p. Cooke, Handb. p. 528; Mier.
Fung. p. 217 p.p.

Uromyces Alliorum DC. ; Cooke, Handb. p. 518; Mier. Fung. p. 212.
Plowr. Ured. p. 137 p.p.

Puccinia Porrt Wint. Pilze, p. 200. Plowr. Ured. p. 148. Sacc. Syll.
vii. 605. Sydow, Monogr. i. 610. Fischer, Ured. Schweiz,
p. 80, f. 61.

Uredospores. Sori amphigenous, on indeterminate pallid
spots, scattered or more or less in
rows, minute, at first covered by ()
the swollen epidermis, yellowish or
reddish-yellow ; spores globose to
ellipsoid, very delicately echinulate,
yellowish, 20—30 p.
Leleutospores. Sorlamphigenous
or caulicolous, generally without
spots, scattered, minute, oblong or fig 182, P. Porri. Telento-
roundish, about 1 mm. wide, but spores and mesospore, on 4.
: ‘ Schoenoprasum.
sometimes confluent into larger

patches, long covered by the lead-coloured epidermis, black-
brown; spores oblong or clavate, rounded or rather truncate

236 PUCCINIA

above, scarcely or slightly thickened, gently constricted, rounded
or attenuated below, smooth, brown, 28-—52 x 20—26 uw; pedicels
hyaline, short, deciduous; mesospores numerous, obovate or
pyriform, very irregular, 22—36 x 15—25 p.

On leaves, sheaths and stems of various species of Allium,
A. Cepa, A. Schoenoprasum, A. Scorodoprasum, etc. Rather
uncommon. June—August. (Fig. 182.)

Fischer describes the uredospores as pale-brown, distantly warted,
28—82 x 21—28 p, and provided with three germ-pores. The ecidium on
Allium which is usually placed with this species does not belong here.
The ecidium on Allium ursinum is known to belong to one of the forms of
P. sessilis : some, if not all, of the secidia on other species of Allium may
be jn the same class. It was Tranzschel (Ann. Mycol. 1910, viii. 415) who
proved that P. Porri is a Hemipuccinia ; he sowed the basidiospores on
Allium and obtained the uredospores direct. The supposed ecidium
is almost always found separate from the uredo- and teleutospores. There
is a very close alliance between P. Porri and Uromyces ambiguus, if indeed
they are not the two extremes of the same species.

If this disease attacks cultivated onions, as it sometimes does, remove
and burn all diseased plants and do not sow onions on the same ground
again for several years.

DistTRiBuTION: Europe, Syria.

105. Puccinia obscura Schrot.

Aeidium Compositarum var. Bellidis DC. ; Cooke, Handb. p. 543.

Puccinia obscura Schrot. in Nuov. Giorn. Bot. Ital. ix. 256. Plowr.
Ured. p. 174; Grevillea, xii. 86. Sacc. Syll. vii. 629. Sydow,
Monogr. i. 645, 898. Fischer, Ured. Schweiz, p. 237, f. 187.

Spermogones. Amphigenous, minute, in dense roundish
clusters, honey-coloured.

Aicidiospores. Aicidia amphigenous, on roundish or irregular
yellow spots, in loose clusters or scattered, between cup-shaped
and cylindrical, with whitish torn margin; spores delicately
verruculose, yellowish, 16—22 p.

Uredospores. Sori generally hypophyllous, on irregular
confluent purplish-brown spots, scattered, elliptical or linear,
long covered by the epidermis, pulverulent, rusty-yellow;

ON JUNCACE 237

spores ellipsoid to ovate, echinulate, pale-brown, 18—26 x ee
22 4; epispore rather thick, with two germ-pores.

Teleutospores. Sori similar, but compact, pulvinate, covered
or surrounded by the cleft epidermis, blackish-brown ; spores

Fig. 183. P. obscura. Teleutospores; two mesospores, one germinated in
the sorus; and two uredospores; all on L. campestris,

oblong, rounded, rarely truncate or conical above and thickened
(5—9 ), gently constricted, usually attenuated below, smooth,
brown, 30—48 x 14—20 #4; pedicels subhyaline, persistent, up.
to 80 « long; mesospores frequently intermixed with the teleu-
tospores.

Aicidia on Bellis perennis, September—December; uredo-
and teleutospores on Luzula campestris, L. silvatica. June—
November. Not common, except locally. (Fig. 183.)

Teleutospores are rarely produced, and I have seen them only on dead
leaves ; the fungus can winter by its uredospores, and in such cases, of
course, the «ecidium will not be formed. It is probably in consequence
of this that the ecidia and the uredospores are frequently not found near
one another ; there is a great difference in their mode of occurrence in
different districts. The uredospores sometimes show a small smooth
spineless area just below the germ-pores. Fischer records them in
Switzerland on Luzula masima and L. pilosa in September, and Sydow
includes all the species (except Z. arcuata) which grow in Britain.

The hetercecism of this parasite was first demonstrated by Plowright ;
the ecidia differ from nearly all others in being produced in late autumn
and winter. According to him, the teleutospores, when they occur, are not
formed till August and September, and germinate after a short resting
period ; thus the succeeding phase arises on the Daisy usually about
October. I have found that the spermogones (which are not mentioned by

238 PUCCINIA

other authors) appear first on yellow spots, and are slowly followed by the
zcidia towards November or earlier; at that time of the year the Daisy
plants are generally in a condition of vigorous, though slow, growth.

There is a species of Pucctnia (P. distincta McAlp.) found on Bellis
perennis in Australia, but this bears acidia and teleutospores on the same
leaves ; there are no uredospores. Since the Daisy is not a native of that
country and is grown there from seed imported from England and
Germany, it would seem that the fungus is imported with the seed ; yet in
our country and in Germany, the Daisy never bears teleutospores. The
teleutospores of the Australian species closely resemble those of P. obscura
on Luzula, and like them are often intermixed with numerous mesospores ;
moreover, they often grow amongst and round the ecidia, apparently
always from the same mycelium. Can it be that the European fungus has
been introduced with the seed and has adapted itself to an autcecious life ?
This, however, seems hardly probable, since Luzula campestris is found in
Australia, but the Puceinia which attacks it there (P. tenuispora McAlp.)
is different from either of our European species.

DIsTRIBUTION: Central and Northern Europe, North
America.

106. Puccinia oblongata Wint.

Caeoma oblongatum Link, Obs. Mye. ii. 27.

Trichobasis oblongata Berk. ; Cooke, Handb. p. 529; Micr. Fung.
p. 223, pl. 7, £. 158—9.

Puceinia Luzulae Lib. ; Cooke, Grevillea, iv. 109; Micr. Fung. p. 203.

P. oblongata Wint. Pilze, p. 183. Plowr. Ured. p. 190. Sace. Syll.
vil. 658. Sydow, Monogr. i, 646. Fischer, Ured. Schweiz, p. 239,
f. 188.

Uredospores. Sori amphigenous, on irregular and confluent
reddish-brown or blackish-brown spots, scattered, oblong, long
covered by the epidermis, ferruginous; spores oblong-ovate to
pyriform or clavate, irregular, smooth, rarely aculeolate at the
summit, rusty-yellow, 830—44 x 12—15 4; epispore colourless,
thick, without germ-pores.

Teleutospores. Sori similar, but soon naked, compact,
blackish-brown; spores clavate, much thickened (10—25 )
above where they are rounded or rarely more or less obliquely
tapering, gently constricted, tapering gradually downwards,
smooth, brown, 44—80 x 16—24 w; pedicels hyaline, persistent,
about as long as the spore or shorter.

ON JUNCACEA 239

On Luzula campestris, L. maxima, L. pilosa. Uredospores,
May—July ; teleutospores, September—November. (Fig. 184.)

PipRIgY

Fig. 184. P. oblongata. Teleutospores (one abnormal) and a mesospore;
b, uredospores; all on same leaf of L. pilosa.

The uredospores are said to be always smooth, more or less obovate,
and often irregular. It is stated by Sydow that they can survive the
winter. Fischer figures anomalous spores, of both kinds, including three-
celled and one-celled teleutospores.

Plowright’s suggestion that this is probably a hetercecious species has,
so far, received not the slightest confirmation. The greatly thickened
summit of the teleutospores is very striking ; they are produced in the
old uredo-sori, especially towards the tip of the leaves, so that in a
sorus which is producing uredospores a-few young teleutospores may be
found, and the fully-formed ones are surrounded by the numerous persistent
uredospore-pedicels.

It is, no doubt, very heterodox, but I cannot help expressing the
opinion that P. oblongata is merely an abnormal development of P. obscura,
On the same leaf of Luzula pilosa, if not in the same sorus; I have
found almost all the various kinds of spores figured by Fischer under both
species. Uredo oblongata Grev. Scot. Crypt. Flor. pl. 12, doubtless
includes this form, but his figure is P. Carvcis.

DisTRIBUTION: Central and Northern Europe.

107. Puccinia Scirpi DC.

Heidium Nymphoidis DC. Flor. fr. ii. 597 and vi. 93. Plowr. Gard.
Chron. 1895, xviii. 96, 135.

Puccinia Scirpt DC. Flor. fr. ii, 228, Plowr. Ured. p. 191. Sace.
Syll. vii. 659. Sydow, Monogr. i. 688. Trans. Brit. Myc. Soc.
i. 58. Fischer, Ured. Schweiz, p. 298, f. 218.

240 PUCCINIA

Spermogones. Epiphyllous, in roundish clusters.

Aeidiospores. Kcidia epiphyllous, on roundish yellow spots,
in orbicular clusters as much as 1 cm. diam. surrounding a
group of eae ine scutelliform, yellow, with a slightly and
irregularly torn narrow margin; spores delicately verruculose,
orange, 12—20 pw.

Uredospores. Sori scattered or in rows, often confluent,
oblong, elliptical or linear, long covered by the swollen epidermis
which is at length longitudinally split, ferruginous; spores
subglobose to ovoid, often flattened on one side, echinulate,
pale-brown, 19—32 x 12—24 y, with two germ-pores.

Fig. 185. P. Scirpi. Teleutospores and mesospore, from the original
specimens, King’s Lynn (ex herb. Plowright).

Teleutospores. Sori similar, generally numerous and con-
fluent, black-brown; spores oblong or subclavate, thickened
above (5—9) and rounded, truncate, or subconical, hardly
constricted, attenuated downwards, smooth, brown, 30-—60 x
12—24 1; pedicels yellowish, persistent, 25—45 uw long; meso-
spores more or less abundant, 24—40 yw long.

Ascidia on leaves of Limnanthemum (Villarsia) nymphueoides
(Nymphoides peltatum), July; uredo- and teleutospores on culms
of Scirpus lacustris, July—November. Rare; King’s Lynn;
near Earith (Huntingdonshire). (Fig. 185.)

Plowright suggested in 1889 (Monograph. p. 191) that this Puceinia
was most likely a heterwcious species. Chodat was led, in 1891, to suspect

the Limnanthemum as the alternate host, by finding the two stages both
together in the same pond in the Botanic Gardens at Geneya. Klebahn

ON CYPERACEX 241

had the same experience at Bremen. Bubdk experimentally demonstrated
the truth of the suspicion (Gisterr. Bot. Zeitschr. 1898, xlviii. 14).

The teleutospores were first found, in this country, on Scirpus lacustris
floating down the river Ouse at King’s Lynn, Nov. 17, 1877. The plants
had evidently been cut on that river or one of its tributaries, but it
was not till 1894 that the teleutospores were found in situ near Earith,
Huntingdonshire ; and a visit to the Old Bedford Level at that place,
in July, 1895, revealed the zecidia in abundance. See Gard. Chron. (/. ¢.).

DistrisuTion: Europe generally; a similar ecidium on
Limnanthemum indicum has been found in Queensland, but no
teleutospores have yet been found there.

108. Puccinia Caricis Reb.

Aeidium Urticae Schuin. Enum. Pl. Sall. ii, 222, Cooke, Handb.
p. 541; Micr. Fung. p. 197, pl. 1, f. 10, 11.

Uredo Caricis Schum. l. ¢. p. 231.

Trichobasis caricina Berk. ; Cooke, Micr. Fung. p. 223, pl. 8, f. 170—1.

Puceinia Caricis Rebent. Fl. Neom. p. 356. Plowr. Ured. p. 169.
Sacc. Syll. vii. 626. Sydow, Monogr. i. 648. Fischer, Ured.
Schweiz, p. 265, f. 201. McAlpine, Rusts of Australia, p. 133,
f. 29, 30:

P. striola Link, Sp. Pl. p. 67. Cooke, Handb. p. 493; Micr. Fung.

p. 2038 p.p.

Spermogones. Epiphyllous, in small clusters, honey-coloured.

Aicidiospores. Aicidia hypophyllous or occasionally amphi-
genous, often on the petioles and
stems, on reddish, yellow or pur-
plish spots, in dense clusters of
various sizes which are often very
large and cause -great swelling
and distortion on the stems, cup-
shaped, with torn white recurved
margin; spores verrtculose, orange,
16—26 x 12—20 p.

Uredospores. Sori amphige-
nous, generally hypophyllous, seat- Fig. 186. P. Caricis. a, teleuto-
tered, oblong, about } mm, long, ‘4%, PPE ell erhnating,
pale-brown ; spores subglobose to
oval, echinulate, yellow-brown, 21—30 x 15—22 p, with three

(rarely four) germ-pores.

GU. 16

242 PUCCINIA

Teleutospores. Sori generally hypophyllous, scattered or
arranged in lines, oblong or linear and confluent into long striz,
pulvinate, compact, black; spores clavate, usually rounded above
and much thickened (5—10 y), constricted, tapering downwards,
smooth, brown, darker at the apex, 35—66 x 1423 yu; pedicels
yellowish, persistent, about half as long as the spore or less.

fEcidia on Urtica dioica, April—June; uredo- and teleuto-
spores on Carex acutiformis (paludosa), C. hirta, C. pendula,
C. Pseudocyperus, C. riparia, June or July—April. Very
common, (Fig. 186.)

It is recorded also, in other countries, upon a long series of other
Carices (over forty-three), including C. acuta, C. Goodenovii, C. stricta, and
C. vesicarta among British species. It must not be assumed, however,
without trial that these are all identical ; the forms on @. hirta, C. vesicaria
seem to be biologically distinct (Klebahn, Fischer), and others may belong
to P. Pringsheimiana, ete. C. binervis is included by Plowright, but the
fungus on this host has not yet been experimentally shown to belong
to 4G, Urticae.

This parasite has been the subject of numerous investigations, since
Magnus in 1872 first showed that the wcidium on the Nettle belonged
to the life-cycle of P. Caricis on Curex hirta. Plowright, Schroter,
Klebahn, and many others have followed in his steps. It may be
remarked here that few species of Puceinia on Carex can be determined
with certainty until the ecidium-stage belonging thereto is known,

The teleutospores of this species germinate after the winter’s rest ;
they succeed most easily during April. They may be found on new or old
leaves of Carex all the year round. It is almost in vain to look for
the ecidium on the Nettle except in the vicinity of water where Sedges are
growing ; but where such a conjunction occurs, the parasite may often be
found upon both hosts in abundance every year. Klebahn records the
ecidium on Urtica wrens; Magnus states that the fungus can winter on
C. hirta by means of its uredospores.

DisTRIBUTION : Europe, Siberia, Japan, America, Australia.

109. Puccinia Pringsheimiana Kleb.

Aicidium Grossulariae DC, Flor. fr. vi. 92. Cooke, Handb. p. 541;
Micr. Fung. p. 197. Plowr. Ured. p. 263.

Puccinia Pringsheimiana Kleb. in Zeitschr. f. Pflanzenkr. v. 76.
Soppitt in Gard. Chron. 1898, xxiv. 145, f. 38. Sydow, Monogr.
i, 652. Fischer, Ured. Schweiz, p. 268. Massee, Diseases of
cultivated Pl. p. 300, f. 88.

ON CYPERACE 243

4Aicidiospores. cidia hypophyllous, crowded on red and
yellow spots, roundish, also in elon-
gatedclusters on the young branches,
petioles, and nerves, sometimes en-
tirely covering the young fruits,
shortly cylindrical, with broad, re-
curved, white, much torn margin;
spores orange, verruculose, 15—21 x
14—18 p.

Uredospores. Sori hypophyllous,
punctiform, about 4mm. long, on
yellowish spots; spores more or less
globose, pale-brown, echinulate, 18— Fig. 197, P. Pringsheimiana.
22 x 17—21y, with three, rarely a Ritespr ie ia
four, germ-pores.

Teleutospores. Sori amphigenous, linear or punctiform, up
to 1 mm, long, pulvinate, brownish-black; spores resembling
those of P. Caricis, 40—58 x 15—22 yp.

Adcidia on Ribes Grossularia, R. nigrum (?), May and June,
common; uredo- and teleutospores on C. acuta, C. caespitosa,
C. Goodenovit, C. stricta. (Fig. 187.)

The teleutospore-hosts are those given by Klebahn and Fischer.
Klebahn first suggested the connection of the wcidium with a Puccinia on
Carex, and has since demonstrated the truth of this idea by many culture
experiments. Soppitt also showed the same for Carex acuta and C. Goode-
novit. The ecidium is said to attack A. alpinum, R. aureum, R. rubrum,
R. sanguineum, but less frequently. Plowright records the ecidium on
leaves of Ribes nigrum (Norfolk, June, 1890), but there is no proof that it
belonged to this species.

This species is one of those forms originally named by Klebahn
P. Ribesti-Caricis ; he has since divided them under five heads which can
scarcely be reckoned anything but biological races :—P. Pringsheimiana,
P. Ribis-nigri-Acutae, P. Ribis-nigri-Paniculatae, P. Pseudo-cyperi, and
P. Magnusit (the latter on C. acutiformis and C. riparia). The same
species of Aibes serve as alternate hosts in each case, in varying degrees
of susceptibility, except that P. Magnusi is not recorded for R. rubrum
and &. Grossularia. The morphological differences between these forms
are slight and elusive.

P. Pringsheimiana can be distinguished from P. Caricis by its nearly

16—2

ee

244 PUCCINIA

round uredospores, but most of the other biological races of P. Aibesit-
Caricis have them oval or oblong. There is no remedy for this disease on
the Gooseberry but to gather and burn all diseased leaves and fruit, etc.,
and even this will be of no avail so long as the affected Carices continue
tu exist. Luckily the disease rarely does much harm,

110. Puccinia dioice Magn.

cidium Cirsti DC. Flor. fr. vi. 94.

Puccinia dioicae Magn. Tag. Nat. Vers. Miinchen, 1877, p. 200. Plowr.
Ured. p. 173. Sace. Syll. vii. 629. Sydow, Monogr. i. 653.
Fischer, Ured. Schweiz, p. 283, f. 208.

Spermogones. In little clusters, honey-coloured.
Acidiospores. Bcidia hypophyllous, on roundish yellow or
brownish spots, in clusters 2—5 mm. diam.,
cup-shaped, with torn white margin;
spores delicately verruculose, orange, 18
—25 mw
[Uredospores. Sori scattered, minute
punctiform, brown; spores globose to
ellipsoid, echinulate, pale-brown, 18—
ic Q5 pe.
Teleutospores. Sori scattered, roundish
Fig, 188. P. dioicae, oblong, 1 mm. long, soon naked, sur-
Hcidia on C. pratense, rounded by the cleft epidermis, pulvinate,
pen tia black; spores clavate, rounded or conical
and much thickened (up to 14y) above,
gently constricted, tapering below, smooth, brown, darker at
the apex, 85—56 x 14—20 u, occasionally 70 long; pedicels
brownish, persistent, as much as 50 pw long.]

Aeidia on Cirsium palustre, C. pratense, and (on the conti-
nent) on other species of Cirsium, June, July; uredo- and
teleutospores on Carex dioica, C. Davalliana(?). Very rare;

Scotland, Ireland. I have not seen the teleutospores. (Fig.
188.)

DistRiBuTION: Northern parts of Europe.

ON CYPERACEE 245

111. Puccinia silvatica Schrot.

Heidium Taraxaci K. et 8. Myk. Heft. i. 85.

Puccinia silvatica Schrot. in Cohn, Beitr. iii. 68. Plowr. Ured. p. 172.
Sace. Syll. vii. 627. Sydow, Monogr. i, 656. Fischer, Ured.
Schweiz, p. 289, f. 211.

Spermogones. In little clusters, yellowish.

Acidiospores. /Ecidia hypophyllous or amphigenous, on
roundish yellow or brown spots, in crowded
clusters 2—5 mm. wide, rarely solitary, occa-
sionally on the peduncle which they distort,
cup-shaped, with torn whitish revolute mar-
gin; spores nearly smooth, orange, 14—21 y.

[Uredospores. Sori hypophyllous, scat-
tered, minute, oblong, brown; spores globose
to ovate, echinulate, brown, 20—27 x 15—
22 pw.

Teleutospores. Sori hypophyllous, scat-
tered, minute, roundish or oblong, reaching
1 mm. in length, pulvinate, black; spores cla-
vate, rounded and much thickened (up to 11 p)
above, rarely conically attenuated, gently con-
stricted, tapering below, smooth, pale-brown, pig, 199. P. sil-

darker at the apex, 85—55x12—18y; pedicels *atica._— Mcidia
on leaf of Tarax-

brownish, persistent, as much as 40 yw long.] acum, from one
; of Soppitt’s spec-
Acidia on Taraxacum officinale, June, — imens, Saltaire.

July. I have seen specimens from both
England and Ireland. Teleutospores on species of Cures.
(Fig. 189.)

The question whether P. silvatica occurs in Britain is still in the same
state as in Soppitt’s time. The ecidium agrees with the one assigned
to that species, but the teleutospores have not been found; there is no
evidence that those found on Carex remota at Kew belonged to this species.
The distinction of this ecidium from that belonging to P. variabilis lies
chiefly in the clustered peridia, which are situated on a thickened part of
the leaf, the peridium-cells are arranged in evident rows and the spores
form, according to Juel, longer chains. Fischer insists that the peridium
cells are thickest on the outer side, while those of P. variabilis are thickest
on the inner side: he maintains that this difference is characteristic of

246 ‘ PUCCINIA

hetercecious and autecious species respectively, though this is certainly
not always true.

The only British species of Carex which have been proved by cultures
to be connected with this ecidium are C. caryophyllea (pruecox) and
C. urenaria, but it is supposed to grow also on a large number of other
species. The matter is, however, complicated by the fact that there are
two other Puceiniae (P. arenarticola and P. Schoeleriana) which are very
closely allied and differ chiefly in having their ecidia on other hosts.

DiIsTRIBUTION: Europe and Siberia.

112. Puccinia Schoeleriana Plowr. et Magn.

Heidium Jacobaeae Grev. Flor. Edin. p. 445.

44. Compositarum var. Jacobaeae; Cooke, Handb. p. 542; Micr.
Fung. p. 198 p.p.

i. Senecionis Fischer, Ured. Schweiz, p. 534, f. 335 (7).

Puccinia Schoeleriana P. et M. Quart. J. Micr. Sci. xxv. (new ser.)
pp. 167, 170. Phill. et Plowr. Grevillea, xiii. 54. Plowr. Journ.
Linn. Soc. xxiv. 91; Ured. p. 171. Sace. Syll. vii. 627. Sydow,
Monogr. i. 659. .

Spermogones. Epiphyllous, orange.

Leidiospores. AKcidia hypophy!l-
lous, clustered on roundish yellow or
brownish spots as much as ] cm. diam.,
cup-shaped, with a torn white reflexed
margin; spores delicately echinulate,
orange, 15—21 p.

Uredospores. Sori generally hypo-
phyllous, on yellowish spots, scattered,
minute, roundish or oblong, pulveru-
lent, surrounded by the cleft epidermis,
pale-brown; spores globose to ovate,
echinulate, yellow-brown, 24—30 x 16
—25 p.

Telentospores. Sori hypophyllous,
scattered or aggregated, 4—14 mm.
long, oblong, surrounded by the torn
epidermis, pulvinate, black; spores

Fig. 190. P. Schoeleriuna. Clavate or fusoid, rounded or conically
Teleutospores, from one of attenuated above and much thickened
Plowright’s cultures-on C.

arenaria, (up to 14), gently constricted, tapering

ON CYPERACE 247

downwards, smooth, brown, darker at the apex, 45—80 x 18—
22 4; pedicels brownish-yellow, persistent, 25—40 w long.

4Ecidia on Senecio Jacobaea, May and June; uredo- and
teleutospores on Carex arenaria, July—May. Rare; Norfolk,
Lincolnshire, Aberdeen, Burntisland. (Fig. 190.)

The life-history of this parasite was investigated by Plowright, who at
the same time demonstrated by his experimental cultures its distinctness
from Puceinia Caricis and P. arenarticola. Fischer records (l.c.) an
ecidium on Senecio Jacobaea, S. aquaticus and S. erucifolius closely
resembling this, but points out at the same time that C. arenaria does
not grow in Switzerland ; if, therefore, it is the same fungus, as it seems
without doubt to be, its alternate stage must occur there on some allied
species of Carex, such as C. disticha.

DistRiBuTion: Germany, Holland, Switzerland (?), Russia.

113. Puccinia arenariicola Plowv.

Puccinia arenariicola Plowr. Journ. Linn. Soc. Bot. xxiv. 90; Ured.
p. 170. Sace. Syll. ix. 311. Sydow, Monogr. i. 661.

4icidiospores. Aicidia generally hypophyllous, in round
clusters on circular yellow spots which
are as much as 1 cm. diam. and margined
with purple, cup-shaped, yellowish, with
torn revolute margin; spores nearly
smooth, yellow, 15—20 u.

Uredospores. Sori on yellowish spots,
linear or oblong, surrounded by the torn
epidermis, brown; spores globose to
ovate, delicately echinulate, pale-brown,
18—22 p.

Teleutospores. Sori generally hypo-
phyllous, scattered or sometimes aggre-
gated, oblong, as much as 1 mm. long, eT ae
pulvinate, black; spores clavate or ob- — Aicidia on leaf of ‘Cen-
long-clavate, rounded above where they pa a
are darker and much thickened (14,), __right’scultures(reduced).
gently constricted, tapering downwards,
smooth, brown, 40—65 x 14—22 w; pedicels brownish, persis-
tent, as much as 40 w long.

248 PUCCINIA

AKcidia on Centaurea nigra, May and June; uredo- and
teleutospores on Carex arenaria, July—April. Very rare; on
the sea-shore, Hemsby, Norfolk. (Fig. 191.)

" It has not been found anywhere else: but in all probability it could
fairly be regarded as merely a well-marked biological race or mutation of
P. Schoeleriana (q.v.). Plowright’s suggestion that it is identical with
P. tenuistipes Rost. seems less likely, but it is closely allied to P. Caricis-
montanae Fisch.

114. Puccinia extensicola Plowr.

Puccinia eaxtensicola Plowr. Ured. p. 181. Sacc. Syll. ix. 311. Sydow,
Monogr. i. 667.

4icidiospores. Aicidia amphigenous or on the stems, seated
on paler spots, scattered or in clusters, cup-shaped, whitish-
yellow, with torn margin; spores delicately verruculose, pale-
orange, 16—22 m.

Uredospores. Sori on extensive pale spots, scattered, mi-
nute, oblong or linear, reddish-brown; spores subglobose or
ovate, irregular, very delicately echintlate, yellowish-brown,
22—30 x 16—22 wu. “

Teleutospores. Sori oblong, $—1 mm. long, covered for a
long time by the epidermis which at length splits, pulvinate,
black; spores subclavate, rounded or truncate above, rarely
hooded, thickened (up to 8,), gently constricted, tapering
downwards, smooth, brown, 40—60 x 18—24 4; pedicels short,
hyaline ; a few mesospores occasionally intermixed.

Kcidia on Aster Tripolium, June, J uly; uredo- and teleu-

tospores on Cares eatensa, August—June. Rare; Wells,
Norfolk. f

The life-history of this species was worked out by Plowright in 1888 ;
it has hardly been found elsewhere except in Istria.

115. Puccinia paludosa Plowr.

Meidium Pedicularis Libosch, Mém. Moscou, v. 76, pl. 5, f. 1. Cooke,
Handb. p. 544; Micr. Fung. p. 199.

Puccinia paludosa Plowr. Ured. p. 174, Sace. Syll. ix. 311. Sydow,
Monogr. p. 671. Fischer, Ured. Schweiz, p. 273, f. 208.

ON CYPERACE 249

Spermogones. In little clusters, honey-coloured.

Aicidiospores. Aicidia hypophyllous or on the more or less
swollen petioles and stems, clustered in round elongated or
irregular groups, cup-shaped, with torn white revolute margin ;
spores delicately verruculose, pallid-orange, 15—25 pu.

Uredospores. Sori hypophyllous, on yellowish spots, scat-
tered or in little groups, very minute, roundish or oblong, soon
naked, pulverulent, yellowish-brown; spores more or less globose,
delicately echinulate, brownish, 20—26 u.

Li

|

Fig. 192. P. paludosa, Teleutospores on C. vulgaris, Norfolk.

Teleutospores. Sori hypophyllous, minute, scattered or
arranged in long lines, pulvinate, quite black; spores clavate,
rounded above and strongly thickened (up to 11 ,), constricted,
tapering below, smooth, brown (upper cell very dark), 45—60
x 16—22 4; pedicels brownish, persistent, rather long.

Aicidia on Pedicularis palustris, June, July; uredo- and
teleutospores on Carex fulva, C. panicea(?), C. stricta, C.
vulgaris (Goodenovit), from July onwards. As usual on Carex,
the teleutospores can be found throughout the winter. Rare;
Norfolk, Orkney. (Fig. 192.)

DIsTRIBUTION: Central and Northern Europe.

116. Puccinia uliginosa Juel.

Aicidium Parnassiae Gravis, in Bot. Gall. ii. 904. Cooke, Mier. Fung.

p. 198; Grevillea, i. 8 and ii, 161. Plowr. Ured. p. 129. Irish
Naturalist, Oct. 1907, p. 321.

250 PUCCINIA

Puccini uliginosa Juel, Ofvers. k. Vetensk.-Akad. Férh., 1894, no. 8,
p. 410. Sacec. Syll. xi. 198. Sydow, Monogr. p. 673. Fischer,
Ured. Schweiz, p. 267, f. 202.

icidiospores. Aicidia hypophyllous, clustered on yellow,
then brown circular spots 2—5 mm. diam., or sometimes
occupying the whole leaf-surface, cup-shaped, with torn recurved
yellowish margin; spores delicately verruculose, orange, 14—
18 yw.

[Uredospores. Sori amphigenous, scattered, very minute,
rounded or oblong, pulverulent, yellow-brown ; spores globose
to ovate, echinulate, brownish, 21—25 yu, with three germ-pores.

Teleutospores. Sori amphigenous or in little groups, minute,
punctiform, roundish or oblong, pulvinate, black ; spores oblong
or somewhat clavate, rounded above and thickened (up to 8 y),
gently constricted, rounded or attenuated below, smooth, brown,
darker at the apex, 30—38 x 12—18,; pedicels subhyaline,
persistent, 15—32 4 long.]

AKcidia on Parnassia vulgaris, June; uredo- and teleuto-
spores on Carex vulgaris (Goodenovit) and its var. juncella.
Only the xcidium recorded for Britain; Glasgow, Aberdeen,
Ireland.

117. Puccinia graminis Pers.

Aeidium Berberidis Gmel.; Cooke, Handb. p. 5388; Micr. Fung.
p. 195.

Trichobasis linearis Lév. ; Cooke, Micr. Fung. p. 223, pl. 7, f. 144.

P. gramints Pers. Disp. Meth. p. 39, Cooke, Handb, p. 493; Mier.
Fung. p. 202. Plowr. Ured. p. 162. Sace. Syll. vil. 622. Sydow,
Monogr. i. 692. Fischer, Ured. Schweiz, p. 243,.f190. McAlpine,
Rusts of Australia, p. 120, and many figures.

Spermogones. In little clusters, honey-coloured.

eidiospores. Aicidia hypophyllous, often also on the fruit,

on roundish, often thickened spots, 2—5 mm. diam., which are
margined with reddish-purple or yellow, clustered or scattered,
cylindrical, white, with a cut and somewhat erect margin;
spores appearing smooth, verging on orange, 14—26 yu diam.

Uredospores. Sori amphigenous, often also on the sheaths

and culms, scattered or arranged in rows, linear, 2—3 mm. long,

ON GRAMINEA: 251

often confluent and reaching a length of 1 cm. or more,
surrounded by the cleft epidermis, pulverulent, yellow-brown ;
spores ellipsoid or ovate-oblong, echinulate, yellow-brown, then
yellowish, 22—42 x 16—22 w, generally with four equatorial
germ-pores.

Cc
Fig. 193. P. graminis. a, ecidia Fig. 194. P. graminis.
on Berberis; b, uredo- and c, Teleuto- and uredo-
teleutospores on wheat. spore on Aira caespi-

tosa..

Yeleutospores. Sori similar, but forming long lines, soon
naked, pulvinate, black ; spores oblong-clavate, rounded or atten-
uated at the summit, much thickened (6—13 ,), slightly con-
stricted, attenuated’ below, smooth, chestnut-brown, 35—60 x
12—22 »; pedicels brownish, persistent, thick, as much as 60 »
long; paraphyses always absent.

AKcidia on leaves and fruits of Berberts vulgaris and Mahonia
Aquifolium ; uredo- and teleutospores on Agropyron caninum,
a. repens, Agrostis alba, A.canina, A. vulgaris, Aira cauespitosa,
A. fleauosa, Alopecurus pratensis, Arrhenatherum elatius, Avena
fatua, A. sativa, Brachypodium pinnatum, Bronus arvensis,
B. asper, B. erectus, B. giganteus, B. mollis, B. secalinus, Dac-
tylis glomerata, Festuca Myurus, PF. ovina, F. rubra, Holcus
lunatus, H. mollis, Hordeum murinum, H. pratense, H. vulgare,
Lolium perenne, Milium effusum, Molinia caerulea, Phalaris
cunariensis, Poa compressa, P. nemoralis, P. pratensis, P.

252 PUCCINIA

trivialis, Secale Cereale, Trisetum flavescens, Triticum vulgare.
These are the British grasses among those recorded by Sydow ;
the teleutospores have not been found, and probably do not
occur, on all these in this country. (Figs. 193, 194.)

This is the famous “rust” (uredo-) or “ mildew” (teleutospores) of corn
about which so much has been written. But all the earlier observers
confused together the various rusts of Cereals of which a number are now
distinguished. P. graminis is known as the “Black Rust,” on account of
the dark colour of the teleuto-sori; these are very distinctive, forming
narrow black lines, 4—1 cm. long, chiefly on the sheaths and culms.
However abundant this species may have been in the past, it is much less
common in mavy parts of England now than some of the following species.
Whether this is due to the general extirpation of wild Barberry bushes or
not, is not certain ; at any rate they are very uncommon, and the ecidium
on the cultivated species of Berberts and Mahonia is rarely met with in
England.

The test by which alone the Black Rust can be absolutely distinguished
is the power possessed by its basidiospores of producing the characteristic
eecidia on the Barberry. There is a European but possibly non-British
species of Puccinia (P. Arrhenatheri) which has also the Barberry for
its alternate host, on which it produces peculiar “ witches’-brooms,” the
mycelium living perennially in the twigs. This is Metdiwm graveolens
Shutt]., which was formerly wrongly identified with <. magelhaenicum
Berk. from Tierra del Fuego.

The uredo-stage of P. graminis can generally be recognised in the field
by its sori, which may reach a length of 10--15 mm. and are of a rusty-
orange or brownish-ochre colour; they do not become general till the
beginning of June. Forms of P. dispersa are often mistaken for it.

Microscopically, the uredospores are seen to be longer compared with
their breadth (more ellipsoid) than is the case with the other cereal
species ;- the teleutospores, which germinate only after a winter’s rest, are
longer and have longer pedicels ; their sori form much more conspicuous
lines and do not remain for long covered by the epidermis. It is the
uredo-stage which does the greatest harm to the crops; it is reported
to cause much loss in the United States, South Africa, Australia and
Tasmania, but not much in India.

This species has been divided by Eriksson into six biological races,
but they are of a very indefinite character and later researches (see
Carleton, ’99, p. 52) throw grave doubt upon their reality. At any rate,
they are not the same in America as in Europe, though this may be
explained by supposing that, since these forms are undergoing evolution
at the present moment, the course of this evolution is different in America
from what it is in Europe. The existence of these races is, however,
‘important ; they show that the wheat cannot necessarily be infected by

ON GRAMINEHX 2538

the forms which grow upon wild grasses ; according to Eriksson this is
absolutely true of his “f. sp. Zriticz,” but Carleton found that the uredo
from wheat would infect certain wild grasses and that the uredo from
some of them, in turn, would infect the wheat. Most of these races can
equally infect the Barberry ; yet Wheat-Rust abounds in South Africa,
Australia and in parts of India, where no species of Berberis are indigenous.
McAlpine in particular finds P. graminis on 27 species of Graminez, yet
he was unable to infect the Barberry-plants imported from England, even
though wheat planted closely around them was covered with the Puceinia.
He comes to the conclusion that the Wheat-Rust of Australia may be a
biological race which has lost the power of producing ecidia owing to the
absence of its zcidial host. According to the evidence at present available,
this seems also to be the case with other species, ¢.g. the xcidial host is
not known in Australia for P. Agrostidis, P. bromina, P. Festucae, P. Loli,
and especially P. Pourum. The latter case is the most striking, since the
Coltsfoot does not exist in Australia, and the uredospores on Poa have
been found there all the year round.

The only practicable remedy for this disease is to plant seeds of
varieties which have been shown to be immune: a certain progress has
already been made by Professor Biffen and others in the production of
these, and McAlpine mentions a variety, “ Rerraf,” which has been found
to be rust-resistant in many of the Australian States, though it lost that
power when transferred to other countries. In the year 1889, which had
a wet and “muggy” spring, the loss due to rust for the whole of Australia
was estimated to be between two and three million pounds sterling.

DIsTRIBUTION: In every country of the world.

118. Puccinia coronata Corda.
Acidium crassum Pers. Syn. Fung. p. 208 p.p. Cooke, Handb.
p. 5388; Mier. Fung. p. 196 p.p.
4G. Frangulae Schim. Pl. Sall. ii, 225.
-Puccinia coronata Corda, Ic. Fung. 4. 6, pl. 2, f. 96. Cooke, Handb.
p. 494; Micr. Fung. p. 2038, pl 4, f 60—2. Plowr. Ured.
p. 163 pp. Sace. Syll. vii. 623 p.p. Sydow, Monogr. i. 699.
Fiséher, Ured. Schweiz, p. 373, f. 270.
Spermogones. Epiphyllous or amongst the ecidia.
cidiospores. icidia hypophyllous or on the petioles, in
roundish groups or irregularly scattered, on yellow or purplish
spots, producing distortion especially of the petioles, cylindrical,
with a white torn revolute margin; spores very delicately
verruculose, orange, 16—25 x 15—20 p.
Uredospores. Sori amphigenous, but mostly epiphyllous,

254 PUCCINIA

scattered or arranged in rows, rarely confluent, minute, more
or less obloug, pulverulent, orange; spores globose to ovate,
shortly echinulate, yellow, 16—25 x 14—20y, with three or
four germ-pores (about ten, Fischer, but ?), and mingled with
a few paraphyses.

Teleutospores. Sori hypophyllous, irregularly scattered,
rarely confluent, oblong or linear, covered by the epidermis,
soon naked, black; spores cuneate, flat at the summit and
crowned with about 5—7 obtuse (digitaliform) darker-coloured
teeth, hardly or not at all constricted, gradually tapering
towards the base, smooth, brown, 35—60 x 12—22 w; pedicels
short, rather thick.

Ecidia on Rhamnus Frangula, May and June; uredo- and
teleutospores on Agropyron repens, Agrostis alba, A. stolonifera,
A. vulgaris, Calamagrostis lanceolata, Dactylis glomerata,
Festuca sylvatica, Holcus lanatus, H. mollis, Phalaris arun-
dinacea (but not yet recorded on all these grasses in Britain),
August—October. Common.

On account of the processes at the summit of the teleutospore this
species is called Crown Rust. It was surmised by Plowright that there '
are two Crown Rusts; these have since been called P. coronatu and
P. Lolii (=P. coronifera). They are equally widely distributed, but are
said to occur on different grasses, with the exception that they are both
found on the two species of Holcus.

In accordance with custom, they are here kept separate, but aside
from the distinction of the hosts they can be separated only by minute
differences. When they occur on Aolcus, therefore, the only test that
could absolutely decide the matter would be to await the maturation of
the teleutospores, and then try if they would infect R. Frangula. The
lighter-orange colour of tne uredo-pustules, and the character of the germ-
pores will, however, distinguish either of them from P. graminis, when
occurring on the same hosts; in the Crown Rusts, moreover, both kinds
of sori are confined almost entirely to the leaves.

The teleuto-sori of P. coronata have a less decided tendency to group
themselves round the uredo-sori than in P. Loli, and do not remain so
long covered by the epidermis, becoming naked early in the autumn. In
the uredo-sori, Eriksson says that paraphyses occur in P. coronata and
hardly at all in P. Loli ; but the evidence seems to favour the conclusion,
in general, that the presence or absence of paraphyses is a note of little
importance. Plowright mentions an interesting fact, confirmed by Pole-
Evans, that the Crown Rust, when it occurs on Dactylis, is au early summer

ON GRAMINEX 255

species and is accompanied by few uredospores, while that on Loliwm
perenne (P. Lolit) occurs only in the autumn with a profuse simultaneous
development of uredospores. This agrees with my experience. Never-
theless, I consider that P. coronata and P. Lolii ave merely biological
forms of one species. The experiments of Carleton, in the United States,
have shown that the distinctions of hosts are quite insufficient to dis-
criminate the two forms ; among others he succeeded in infecting a host,
stated hy Eriksson to be confined to P. coronata, with spores from a host
belonging to 2. Loliz.

P. Festucae Plowr. belongs to the same group, but has its acidium on
Honeysuckle. Crown Rusts have been found in Europe on many other
species and genera of grasses (Sydow, i. 705), but they cannot, in the
absence of cultures, be even temporarily arranged under the two heads.
Barclay described a form from Simla (P. coronata var. himalensis) on
Brachypodium silvaticum with its ecidium on Rhamnus dahurica ; this
has since been raised by Dietel to a species (P. himalensis). ,

It is a remarkable fact that the only other species of Puccinia known,
provided with the same processes, are two (P. Mesnieriana=digitata, and
P. Schweinfurthit) whose teleutospores occur on Rhamnus. A somewhat
similar form, but with much longer apical processes, is found on Lonicera
in Turkestan (P. longirostris Kom.). These instances are similar to those
of P. fusca and P. Pruni-spinosae already mentioned and can be explained
in the same way ; see Grove, New Phytologist, 1913, p. 89.

119. Puccinia Lolii Nielsen.

eidium erasswm Pers. ; Cuoke, Handb. p. 538 ; Micr. Fung. p. 196 PP,

44, Cathartici Schum. Pl. Sal. ii, 225.

Puccinia coronata Corda ; Cooke, Handb. p. 494 ; Micr. Fung.
D- 203 p.p. Plowr. Ured. p. 163 p.p. Sacc. Syll. vii. 623 p.p.

P. Lolti Niels. Ug. for Landmaend. i.549 (1875). Sydow, Monogr. i.
704, McAlpine, Rusts of Australia, p. 123 and many figures.

P. coronifera Kleb, Zeitsch. f. Pflanzenkr. iv. 132 (1894).

Fischer,
Ured. Schweiz, p. 375, f. 271.

Ajcidiospores. Kcidia hypophyllous or on the petioles,
seated on yellow or purplish spots, scattered or arranged in

groups, producing distortion of the parts, cylindrical, with a-

whitish torn revolute margin; spores very delicately verrucu-
lose, orange, 16—25 x 12—20 wu.

Uredospores. Sori amphigenous, scattered or in large
patches, forming blister-like swellings, minute, sometimes con-
fluent, lanceolate or more or less oblong, pulverulent, orange ;
spores globose to obovate, echinulate, yellow, 18—27 x 16—24,y,

Ca

256 PUCCINIA

with three or four inconspicuous germ-pores; paraphyses very
few or wanting.

Teleutospores. Sori hypophyllous, sometimes arranged in
circles round the uredo-sori, rarely scattered, occasionally con-

Fig. 195. P. Lolii. Acidia on leaves of R. catharticus ; Teleutospores,
a, on Lolium, b, on drrhenatherum.

fluent, oblong or linear, 3—1 mm. long, nearly always covered
by the epidermis, black; spores as in P. coronata, but very
irregular; mesospores also occur.

AKcidia on Rhamnus catharticus, May and June; uredo- and
teleutospores on Alopecurus pratensis, Arrhenatherum elatius,
Avena fatua, A. pratensis, A. sativa, Festuca elatior, Glyceria
aquatica, Holcus lanatus, H. mollis, Lolium perenne; not yet
recorded on all these species of grass in Britain. (Fig. 195.)

The Crown Rust of the Oat is most commonly found on Rye-grass,
frequently also on Arrhenatherum and Holecus, and also on cultivated
Oat which alone of the cereals it attacks, doing considerable damage.
The teleutospores can be found on .lrrhenatherum from the middle of
August onwards and, as Plowright remarked, are “accompanied by a
profuse development of uredospores”—so profuse, indeed, as to attract
the notice of even non-botanical eyes. The uredo-sori form more blister-
like swellings and the teleuto-sori remain longer covered by the epidermis
than is the case in P. coronata. The uredospores are much brighter in
colour than those of P. graminis.

This species has been divided by Eriksson into a varying number of
biological races, of which P. Lolii .lvenae is the most important; see p. 68.
It is found in Australia on Oat and Rye-grass (introduced with seed 2)
although no species of RAamnus is indigenous there (McAlpine).

DisTRIBUTION: Europe, Asia, North America, Australia.

ON GRAMINEEX 257

120. Puccinia Festuce Plowr.

Ateidium Periclymeni Schum. Enum. Pl. Sill. ii. 225. Cooke, Mier.
Fung. p. 196. Plowr. Ured. p. 264.
.E. crassum var. Periclyment Cooke, Handb. p. 539.
Puccinia Festucae Plowr. Gard. Chron. 1890, ii. 42, 139, and 1891,
i. 460; Grevillea, xxi. 109. Sacc. Syll. xi. 194. Sydow, Monogr.
i. 752. Fischer, Ured. Schweiz, p. 377, f. 272. McAlpine, Rusts
of Australia, p. 119, f. 13.
Spermogones. In‘small clusters, honey-coloured.
Ajcidiospores. Aicidia hypophyllous, on round yellow or
brownish spots, 1n roundish clusters 2—5 mm. diam., shortly
cylindrical, whitish-yellow, with recurved irregularly torn
margin; spores delicately verruculose, orange, 16—27 y.
Uredospores. Sori epiphyllous, scattered, minute, oblong,
yellow; spores globose to ellipsoid, echinulate, yellow-brown,
2230 w, without paraphyses.

ile

v
Fig. 196. P. Festucae. Teleutospores and mesospore.

Teleutospores. Sori hypophyllous, minute, scattered, oblong
or sublinear, black-brown; spores clavate-oblong, crowned at the
summit, with four to six curved and sometimes bifid processes,
gently constricted, attenuated downwards, smooth, pale-brown,
40—60 x 15—23 w; pedicels brown, persistent, 15—25 pu long;
a few mesospores intermixed.

Aicidia on leaves of Lonicera Periclymenum, June—August,
not uncommon; uredo- and teleutospores on Festuca duriuscula,
F. ovina, August—October, not common or at least rarely
observed. (Fig. 196.)

It was Plowright who first, in 1890 (after twenty-eight unsuccessful
trials), proved that the well-known xcidium on Londcera was connected

G. U. 17

258 PUCCINIA

with a Crown Rust on Festuca ; Klebahn and Fischer have since repeatedly
confirmed his results. Fischer records P. Festucae for Festuca rubra and
its var. fallax, and he connects it with an ecidium on Lonicera coerulea
and LZ. nigra. He states that the uredospores have about six germ-pores ;
that the teleuto-sori are situated in the groove of the upper side of the
leaf, at first covered by the epidermis which at length opens by a slit ; also
that the teleutospores have a thick-walled pedicel and are much thickened
and occasionally conical at the vertex ; mesospores were sometimes found
intermixed.

McAlpine found the uredospores with five scattered germ-pores on one
face (z.e. probably six or more altogether). His specimens were on Festuca
ovna and F. rig¢da, but in Australia no ecidium on Lonicera is known.
He found very commonly, mixed with the teleutospores, mesospores
provided with similar processes at the apex.

Distrisution: Central and Northern Europe, North
America, Australia.

121. Puccinia glumarum Er. et Henn.

Uredo glumarum Schmidt, Allg. dkon. Fl. i. 27 (1827).

Trichobasis glumarum Lév.; Cooke, Handb. p. 529; Micr. Fung.
p. 223.

Puccinia Rubigo-vera Plowr. Ured. p. 167 p.p.

P. straminis Cooke, Mier. Fung. p. 202 p.p.

P. glumarum Erikss. et Henning, Getreideroste, p. 141. Trans.
Brit. Myc. Soc. i, 59. Sydow, Monogr. i. 706. Fischer, Ured.
Schweiz, p. 366, f. 265.

Uredospores. Sori amphigenous and on the inflorescence,
minute, oblong, as much as 1 mm. long, arranged in long lines,
on yellow discoloured spots, not often confluent, lemon-yellow ;
spores globose to broadly ellipsoid, echinulate, yellow, 25—30 x
18—26 u; membrane always distinctly colourless.

Teleutospores. Sori hypophyllous or culmicolous, arranged
in long fine lines, a few scattered on the inflorescence, oblong,
dark-brown or black, covered by the epidermis ; spores clavate,
rounded, truncate or obliquely conical above, where the exo-
spore may be as much as 4—6 (or even 10) « thick, slightly
constricted, attenuated below, smooth, brown.30—70 x 12— 244 :
pedicels very short or almost none; paraphyses brown, numerous,
curved, surrounding each little group of teleutospores.

ON GRAMINEA 259

On leaves, culms and glumes of Agropyron caninum, A.
repens, Brachypodium sylvaticum, Bromus mollis and other
species, Elymus arenarius, Hordeum vulgare, Secale Cereale;

.
’
a
‘
a
‘

MW [
a b a

Fig. 197. P. glumarum. a, uredo-sori and b, teleuto-sori, on leaves, nat.
size; c, teleuto-sori, on glume; d, uredospore; e, teleutospore.

5
c
’
‘
"J
5
4
.
‘
4
‘
t
t
‘

Triticum vulgare. It is one of the few species that attack the
ears, to which it does serious damage. (Fig. 197°)

This is one of two forms originally included under the collective name
P. Rubigo-vera DC. ; they can best be distinguished in the uredo-stage.
In the one, P. glwmarum, the uredo-sori are abundant, clear lemon- or
sometimes orange-yellow, and stand in long lines, often occupying half the
leaf-blade ; in the other, P. dispersa and its subordinate forms, they tend
towards brownish-orange or even chocolate-brown, and are scattered,
rather thinly and without order, over the whole leaf-surface. Eriksson
and Henning first proved in 1896 what had been long surmised, that they
are quite distinct. P. glumarum has no known ecidial form, and has
been divided into five biological races (see p. 67). It is common in certain
districts of England and is called the Spring Rust on account of its
early appearance, and Yellow Rust on account of its bright colour, which
varies from sulphur- to pale cadmium-yellow. The lines of the uredo-sori
may be as much as 7 cm. long, chiefly on the upper face of the leaf; they
can be found all the year round on suitable leaves, and are frequently
abundant on Wheat as early as the beginning of May. The teleutospores
germinate as soon as they are mature; the basidium is yellow until
the basidiospores are formed, not colourless as in P. dispersa.

On Hordeum murinum there is recorded a form, P. Hordet Fckl.,
which has smaller yellow sori, arranged less evidently in lines; this may
belong to P. gluwmazum but has not yet been found in Britain. It is
probably not identical with P. simplex (q. v.).

DisTRIBUTION : Europe, Egypt, North America, Japan.
17—2

260 PUCCINIA

122. Puccinia dispersa (sens. lat.) Er. et Henn.

Trichobasis Rubigo-vera Lév.; Cooke, Micr. Fung. p. 222, pl. 7,
f. 140—2 (?).

Puccinia straminis Cooke, Mier. Fung. p. 202 p.p.

P. Rubigo-vera Plowr. Ured. p. 167 p.p. Sace. Syll. vii. 624 p.p.

P. dispersa Erikss. et Henning, Getreideroste, p. 210 (1896). Trans.
Brit. Myc. Soe. i. 58.

Uredospores. Sori generally epiphyllous or a few fhypo-
phyllous, 1—2 mm. long, scattered without order, rarely con-
fluent, oblong or punctiform, rust-coloured or dirty-ochre, be-
coming paler; spores more or less globose, shortly echinulate,
dirty-yellow or dull-orange, 16—28; membrane distinctly
brownish (pale chocolate-umber) when mature; germ-pores
7—10, scattered over the whole surface.

Teleutospores. Sori hypophyllous or less often on the
sheaths, scattered or slightly and irregularly aggregated, rarely
in distinct lines, small, oblong, covered by the epidermis, black ;
spores oblong to clavate, truncate, rounded, or obtusely and
obliquely pointed above, slightly thickened, gently constricted,
narrowed downwards, smooth, brown, 35—56 x 12—23 uw; pedi-
cels short ; paraphyses numerous, brownish, more or less curved,
surrounding the spores.

This is a general description of the forms included under the name
Brown Rust, to which the title P. dispersa was originally given. The
dirty-orange colour of the uredospores, which distinguishes them at a
glance from P. glumarum, is due to the fact that the membrane of the
spores is brownish, not hyaline ; the spore contents are orange in colour.
The germ-pores are scarcely perceptible in the immature or untreated
spore, but they can be seen easily if a spore is squeezed strongly between
the cover-glass and the slide, or by choosing a mature and empty spore.
In the teleutospores only the upper slightly thickened wall is dark-
chestnut, the rest being thin-walled and pale; there is usually also a
chestnut-brown band at the apex of the pedicel. The structures called
paraphyses here in the teleuto-sori are quite different from those called by
the samehame in Melampsora, etc. ; they are erect, coherent, thick-walled,
prismatic cells, which surround the teleuto-sori, or in the case of the
larger ones divide them into loculi. There are also paraphyses of the
ordinary shape, with a brownish membrane, mingled with the uredospores
in certain cases, but the occurrence of these seems, so far as is known at
present, to be somewhat fortuitous,

ON GRAMINEA 261

Eriksson has divided this species into a number of forms which show
certain differences, chiefly biological : they are given in what follows, but
it must be understood that they are distinguished almost entirely by their
host-plants. They all show the same scattered, brownish-orange uredo-
sori. Some, it is true, have ewcidia, others are not known to have them,
but this is a difference which time may remove; also Pole-Evans (1907)
has shown that they present minute differences in the mode of germination
of their uredospores. Many authors prefer to consider the biological races
which follow as distinct species, but if that is done it is a mistake, which
entails continual confusion, to retain the name P. dispersa for one of them.

DiIsTRIBUTION: Europe, Asia Minor, North America, and in
Australia (probably only introduced).

(1) PUCCINIA SECALINA nov. nom.

Puccinia dispersa (sens. strict.) Erikss. Ann. Sci. Nat. ser. 8, ix. 268,
pl. xi, f. 1—6 (1899). Sydow, Monogr. i. 709. Fischer, Ured.
Schweiz, p. 357, f. 261. Klebahn, Wirtswechs. Rostp. p. 237.

‘ Adium Anchusae Erikss. et Henning, Getreideroste, p. 210.
4. Asperifolia Pers. ; Cooke, Handb. p. 541; Micr. Fung. p. 197 p.p.

Aicidiospores. Aicidia hypophyllous or often on the calyx
and fruit, seated on rounded yel-
lowish or reddish-yellow spots,
cup-shaped, with an incised revo-
lute margin; spores verruculose,
orange, 20—26 p.

fAKcidia on Anchusa (Lycopsis)
arvensis, very rare, Shere, Folkes-
tone, Eltham, etc, August; uredo-

and teleutospores on Secale Cereale, /
May—October. (Fig. 198.)

The Brown Rust of Rye in the Fig. 198. P. secalina. Aicidia
uredo- and teleuto-stages seems to be ate ace
confined to that cereal. Eriksson and of Rye.

Klebahn have proved that it can be :

transferred from species of Anchusa to the Rye and vice-versdé, but not,
they say, to other plauts. Plowright’s observation (Ured. p. 168) that
the zcidium was produced on Anchusa by infection from a rusted bundle
of wheat straw is discredited by them, but possibly without sufficient
reason. The teleutospores are capable of germination as soon as they

mature ; hence the ecidium is usually met with in August and September.

262 PUCCINIA

(2) Pucctnra BRoMINA Erikss.

Puceinia bromina Erikss. l.c, p. 271, pl. xii, f.12—17., Sydow, Monogr.
i. 712. McAlpine, Rusts of Australia, p. 116, pl. C, and fig. 28.

P. Symphyti-Bromornm ¥, Miill. Beihefte Bot. Centralbl. 1901, x. 201.
Klebahn, /.c. p. 239. Fischer, Ured. Schweiz, p. 359, f. 262.

P. dispersa Erikss.; Marshall Ward, Ann. Bot. 1902, xvi. 233;
Annal. Mycol. 1903, p. 132. Freeman, Ann. Bot. zbed. p. 487.

Aeidium Symphyti and Al. Pulmonariae Thiim.

Spermogones. Honey-coloured.

Aicidiospores. Aicidia hypophyllous or on the petioles,
sometimes even on the calyx, seated on large round or irregular
spots which are purplish-brown and surrounded by a yellow
zone, cup-shaped, with an incised revolute margin; spores
verruculose, orange, 18—27 p.

Fig. 199. P. bromina. a, teleutospores; b, a mesospore; c¢, paraphyses
surrounding the teleuto-sori ; d, uredospores (empty); e, paraphysis in the
uredo-sori. On Bromus sterilis. .

Aicidia on Symphytum officinale; uredo- and teleutospores
on many species of Bronwus, sometimes even on the inflorescence,
teleutospores from June onwards. (Fig. 199.)

The teleutospores germinate only after the winter’s rest ; Ward found
germinable uredospores even in February and March; it has been proved
that an ecidium on Pulmonaria montuna belongs to the same life-cycle.
F. Miiller, E. 8. Freeman, and especially Marshall Ward, have investigated
the Brown Rusts of the Bromes, and have discovered a wonderful series of
inter-relations among them.

(8) PuccInia TRITICINA Erikss.

Puceinia triticina Evikss. l.c. p. 270, pl. xi, f. 7-11. Sydow, Monogr.
i. 716. Klebahn, Zc. p. 245. Fischer, Ured. Schweiz, p. 366.
McAlpine, Rusts of Australia, p. 182, pl. i, f. 3, 6, 10; pl. xl,
f. 302; pl A, f. 1, 2

ON GRAMINEA 263

On Triticum vulgare. June—August. No ecidium is known
to belong to it. (Fig. 200.)

The Brown Rust of Wheat has been frequently so abundant in this
country in its uredo-stage as to cause great loss. The uredospores can be
distinguished from those of P. graminis, when both occur upon wheat, by
being subglobose, not elongate-ellipsoid, and by the more numerous germ-
pores which are scattered instead of forming an equatorial band ; also
they appear early in spring, before those of P. graminis. Sometimes the
teleuto-sori occur on the culms, and are then arranged more or less in
lines, but they are most common on the underside of the “flag”; their
spores germinate only after a winter’s rest.
Mesospores are not frequent in this species.
Klebahn tested the basidiospores of this
Rust on forty-two likely species of plants
in the hope of discovering an ecidium in
its life-cycle, but without any result. The
uredospores were found to be capable of

surviving the winter by McAlpine in |
Australia (where it is an introduced 7 4
species), and by Carleton in the United

Fig. 200. P. triticina.
States south of lat. 40° N. “ Telepisesarn

(4) Puccinia Houcina Erikss.

Puceinia holeina Erikss. é.c. p. 274, pl. xiii, f. 22—5. Sydow, Monogr.
i. 715. Klebahn, Zc. p. 249. Fischer, Ured. Schweiz, p. 365.

On Holcus lanatus, H. mollis. June—October. Common.

The uredo-stage must be carefully distinguished from that of P. coronata,
which occurs on the same hosts ; the number of germ-pores at once decides
the question. The uredo-sori are of a brighter colour than in the other
forms of P. dispersa and stand upon conspicuous pale spots. The teleuto-
spores are more rarely produced and require to be looked for closely ; they
resemble those of P. triticina, but are mingled with a few mesospores.

(5) PuccInia AGROPYRINA Erikss.

Puceinia agropyrina Erikss., l.c. p. 273, pl. xii, f. 18—21. Klebahn,
Zc. p. 249. Sydow, Monogr. i. 712. Fischer, Ured. Schweiz, p. 365.

On <dgropyron caninum, A. repens. August—October.
(Fig. 201.)

This is one of the commonest of the Rusts on wild grasses in the
autumn, and is easily recognised by the following points: The small
scattered dull-orange uredo-sori on the upper leaf-surface; the round
faintly echinulate uredospores, which when empty show a pale-chocolate

264 PUCCINIA

membrane marked with about nine (7—10) germ-pores, which are each
surrounded by a little thickening of the cell-wall, so that they look some-
what like a “ bordered pit” ; the teleuto-sori mostly on the lower leaf-surface
or sheath, black, covered by the epidermis ; the teleutospores obconical or
with nearly parallel sides, truncate, rounded, or pointed (obtusely and
often obliquely) at the apex; the slightly thickened apical wall and a
broad band at the base chestnut-brown, but the remainder rather pale ;
each little group of teleutospores surrounded by a dense wall of brown
closely coherent “paraphyses.” Oftentimes there are numerous meso-
spores, especially on the margin of the sorus. The clavate paraphyses
which are frequently present in the uredo-sori of other forms of P. dispersa

Fig. 201. P. agropyrina. a, teleutospores; b, paraphyses with the same;
c, plan of paraphyses; d, paraphysis with uredospores ; ¢, uredospore.

are mostly absent in this; when present they have a thin brownish
membrane exactly of the same colour as that of the uredospores. This
fungus is as distinctively an autumn parasite as P. glumarum is a spring
one on the same hosts.

(6) Puccinra TrisEtr Erikss.

Puceinia Triseti Evikss. Ann, Sci. Nat. ser. 8, ix. 277, pl. xiii, f. 26—9
(1899). Sydow, Monogr. i. 716. Fischer, Ured. Schweiz, p. 364.
Klebahn, Zc. p. 249.

.On TLrisetum flavescens. Uncommon ; Alvechurch, Hereford,
etc. June—October. P. graminis occurs on the same host.

(7) Puccinta simpLex Er. et Henn.

Puccinia simples E. et H. Getreideroste, p. 258 (1896). Sydow,
Monogr. i, 756. Klebahn, Zc. p. 248. Fischer, Ured. Schweiz,
p. 368, f. 266. McAlpine, Rusts of Australia, p. 130, pl. i, f. 1,
4, 9 and pl. B, f. 9, 10 (introduced in 1902).

P. Rubigo-vera var. simplex Korn. ; Plowr. Ured. p. 168.

ON GRAMINEZ 265

On leaves and culms of Hordewm vulgare and other species
of Hordeum. Teleutospores, August, September. (Fig. 202.)

The Dwarf Brown Rust of Barley, distinguished by the fact that it
bears few two-celled teleutospores, but very numerous mesospores, which
are variable and asymmetrical, slightly thickened at the apex (46),
measuring 25—45x16—24p. It is to be found in the uredo-stage all the
year round. The teleutospores germinate in spring; Klebahn tried to
infect, with their basidiospores, the same forty-two species which he
tested with P. triticina, but equally in vain.

009

Fig. 202. P. simplex. Teleutospores.

P. simplex may be worthy of being regarded as a distinct species ; it
presents a little more difference from the other forms of P. dispersa than
they do from one another. The sori of both kinds are amphigenous and
more minute and punctiform (except on the culms), and the uredospores
are of a brighter yellow. On Hordeum distichum I have found sori of
P. simplex on the leaves, and with them those of P. graminis on the culms,

It will: be noticed that all the last five races are without
any known ecidium; it follows, apparently, that they must
maintain themselves by their uredospores, but one might
venture to suggest that future, unexpected, discoveries will
throw light upon this obscure matter. Eriksson and Klebahn
have both proved, by numerous infection experiments, that
these races or subspecies of P. dispersa are all biologically
distinct ; with few, and doubtful, exceptions none of them can
be transferred from its own to the other hosts.

There are other forms of Brown Rust of which little is
known. The ecidium on Lchium vulgare mentioned by Plow-
right (Ured. p. 168) may belong to one of these; this plant was
one of the forty-two previously mentioned, tested by Klebahn.

I have also some specimens of uredospores on Aira flexuosa

-and A. caespitosa sent by Mr T. B. Roe from Scarborough; those

266 PUCCINIA

on A. flecuosa have no paraphyses, as those on A. caespitosa
have, but otherwise they are almost identical and are undoubt-
edly P. dispersa (sens. lat.). I find similar spores on A. caes-
pitosa round Birmingham, and have a specimen on the same
host sent by the late H. T. Soppitt from Saltaire; this latter
is the plant referred to in a note to P. Bary: (Plowr. Ured.
p. 192)—both these have abundant paraphyses with the uredo-
spores. Herr H. Sydow informs me that he considers the
presence or absence of these paraphyses to be a character of
little importance in P. dispersa. It must be remembered that
P. graminis also grows on A. caespitosa, but the uredospores
can be easily distinguished by their elliptical shape and three
or four subequatorial yerm-pores ; their membrane is brownish,
but without the chocolate tinge of P. dispersa.

123. Puccinia sessilis Schneid,

Puccinia sessilis Schneid. in Schrot. Brandpilze Schles. p. 19.
P. linearis Rob. ; Cooke, Mier. Fung. p. 203.

cidiospores. See the descriptions given for the four

specialised biological races.
Uredospores. Sori amphigenous, scattered, minute, puncti-
form or shortly linear, yellow; spores globose to ellipsoid,
echinulate, brownish-yellow, 20

: —28 x 18—24 y; epispore thin,

( ) with about seven germ-pores.
Teleutospores. Sori similar,
(_) sometimes confluent, long cov-
ered by the epidermis, pul-

vinate, black; spores oblong

Fig. 2038. P._ sessilis. or oblong-clavate, rounded or
Teleutospores and mesospore.

truncate above where they are

darker and slightly thickened (2—5), hardly constricted,

somewhat narrowed below, smooth, brown, 35—52 x 15—22 py;

pedicels very short or absent ; an occasional mesospore is found,

Aicidia on various species of Monocotyledons; uredo- and

teleutospores on Phalaris (Digraphis) arundinacea, uncommon,
July—May. (Fig. 203.)

ON GRAMINEE 267

The four following biological races agree exactly in the teleutospores,
and these can only be distinguished by their successful use to infect
the alternate host; though sometimes the question may presumably be
decided by finding one or more of those hosts, in the immediate neigh-
bourhood, affected by the ecidium.

It is evident from the disagreement between various authors that it
is impossible to decide to which of the four biological races the name
P. sessilis Schneid. should be applied: it will be better, therefore, to use it
as a collective title, which can be employed in cases where the xcidial
host cannot be determined. A fifth race, P. Schmidtiana Dietel, having
its ecidia on Leucojum, has not yet been found in Britain.

DISTRIBUTION: Europe and North America.

(1) Puccinia DicRapHipis Soppitt.

Aicidium Convallariae Schum. Enum. Pl. Sill. ii. 224. Plowr. Ured.
p. 264. Soppitt in Gard. Chron. ser. 3, vii. 648.

Puceinia Digraphidis Soppitt in Journ. Bot. 1890, p. 213.

P. intermixta Friend in Gard. Chron. ser. 3, viii. 270 p.p.

P. Paridis Plowr. in Gard. Chron. 1892, p. 137; Journ. Linn. Soc.
1893, p. 43.

P. Smilacearum-Digraphidis Kleb. Zeitschr. f. Pflanzenk. 1896, p. 261.
Fischer, Ured. Schweiz, p. 340, f. 251.

P. sessilis Sydow, Monogr. i. 781.

Spermogones. Epiphyllous or in the midst of the ecidia.

Aicidiospores. /Ecidia hypophyllous, loosely clustered on
roundish or irregular yellow spots, cup-shaped, with a cut white
revolute margin; spores verruculose, yellowish, 19—27 p.

fEcidia on Convallaria majalis, Paris quadrifolia. Not
common. May and June.

Attempts have been made to subdivide still further the fungi included
under this head. P. Digraphidis Soppitt, on Convallaria, and P. Paridis
Plowr., on Paris, are two of these forms which to their authors appeared
under cultivation to be confined to their respective ecidial hosts. But,
on the other hand, Klebahn has been able to infect, from one and the
same Puceinia, both Convallaria, Matianthemum, Paris, and Polygonatum ;
nevertheless his attempts to induce specialisation, by cultivating the
fungus year after year on Polygonatum alone, had the result that towards
the end (while it still grew freely on that genus) it could be transferred
only with difficulty or not at all to the other genera. Evidently we have
here a case where specialisation is naturally in progress, but has not yet
proceeded far enough to effect complete separation.

268 PUCCINIA

(2) PuccINIA ORCHIDEARUM-PHALARIDIS Kleb.

eidium Orchidearum Desm. Cat. Plant. omis. p. 26. Cooke, Handb.
p. 545; Mier. Fung. p. 200. Plowr. Ured. p. 179.

Puceinia Orchidearum-Phalaridis Web. Zeitschy. f. Pflanzenkr. 1899,
p. 155. Sydow, Monogr. 1. 782. Fischer, Ured. Schweiz, p. 343.

Spermogones. Epiphyllous or in the midst of the groups of
weidia.

cidiospores. ASeidia hypophyllous, usually in circular
clusters on yellow spots, cup-shaped, with a cut white reflexed
margin; spores verruculose, yellow, 17—26 yw, sometimes slightly
ellipsoid.

Aicidia on Orchis latifolia. Not common. May—July.

It is recorded, on the continent, on several other species of Orchidacee.
This ecidium must not be confounded with
the Caeoma Orchidis, which belongs to a Me-
lampsora.

(3) Puccinta WINTERIANA Magn.

Beidium Alii Grev. Flor. Edin. p. 447.
Cooke, Handb. p. 545; Mier. Fung.
p- 200.

Puccinia sessilis Plowr. Ured. p. 165. Save.
Syll. vii. 624.

P. Winteriana Magn. in Hedwig. 1894,
p. 78. Sydow, Monogr. i. 783.

P. Allvi-Phalaridis Web. in Pringsh. Jahrb.
1899, p. 399. Fischer, Ured. Schweiz.
p. 343.

Spermogones. Epiphyllous or in the
midst of the weidia.

Beidiospores. Meidia hypophyllous,
in circinate clusters on large yellow
spots, cup-shaped, with a cut white
recurved margin; spores very delicately

verruculose, yelowish, 17—26 u.

Fis. 204. Ps Winweertana, Meidia on Allium ursinum. Not
Spermogones (sp) and eommon. End of May—July. (Fig.
weidia (aec) on Allivn ’ : ;

wrsinum. xX 4. 204.)

ON GRAMINEZ 269

I have found that in many cases only one cluster is formed (or at
most two small ones) on a leaf; this probably indicates a scarcity of
active basidiospores. In other localities, however, eight or ten clusters
may be found on a single leaf. This ecidium must not be confounded
with Caeoma Alliorum, which belongs to the Melampsore.

(4) Puccinia PHauaripis Plowr.

eidium Ari Desm, Cat. Plant. omis. p. 26. Cooke, Handb. p. 545 ;
Micr. Fung. p. 199.

Puccinia Phalaridis Plowr. Journ. Linn. Soc. 1888, xxxiv. 88; Ured.
p. 166. Sydow, Monogr. i. 783.

P: Art-Phalaridis Kleb. in Pringsh. Jahrb, 1899, p. 399. Fischer,
Ured. Schweiz, p. 344, f. 252.

Spermogones. Epiphyllous or a few hypophyllous, dark
honey-coloured, in roundish groups.

Ajeidiospores. Aicidia hypophyllous, in roundish clusters
(often surrounding a little group of
spermogones), on conspicuous pale-
yellow spots, cup-shaped, with a broad
cut white revolute margin ; spores deli-
cately verruculose, yellow, 15—26 p.

Aicidia on Arum maculatum, Not
common. May—July. (Fig. 205.)

The same remark may be made about the
occurrence of this as about the preceding form.
The three latter races are biologically quite dis-
tinct in so far that, in experimental cultures,
each of them will produce the ecidium only
on that particular genus to which it has Fig. 205. P. Phalaridis.

i Aicidia on Arum
become accustomed. This has been abun- maculatum. x}.
dantly proved by Plowright, Klebahn, Fischer,
Dictel and others. But the groups of zcidia are exactly of the same type
in each case, and must have had a common origin in the past.

124. Puccinia Anthoxanthi Fckl.

Puceinia Anthoxanthi Fckl. Symb. Myc. Nachtr. ii. 15. Plowr. Ured.
p. 194. Sace. Syll. vii. 665. Sydow, Monogr. i. 727. Fischer,

Ured. Schweiz, p. 261. McAlpine, Rusts of Australia, p.°115,
f. 20—1. :

270 PUCCINIA

Uredospores. Sori amphigenous, on indefinite yellowish
spots, scattered or in groups, elliptical or linear, soon naked,
minute, rusty-yellow; spores ellipsoid to ovate, delicately echinu-
late, yellowish, 20—30 x 15—20p (with 2—4 very distinct
equatorial germ-pores on one face, McAlpine),

Teleutospores. Sori very inconspicuous, amphigenous, scat-
tered, minute, soon naked, elliptical or linear, blackish-brown ;
spores elliptical or subpyriform or oblong, usually rounded
above and thickened (up to 8), gently constricted, rounded
or rarely attenuated below, smooth, chestnut-brown, 28—48 x
16—22 4; pedicels persistent, brownish, up to 45 long.

On leaves (living or fading) of Anthoxanthum odoratum.
Very rare; King’s Lynn, 1884.

The description is from Sydow, where it is stated that the uredo is
frequent in mid-Germany, but the teleutospores are exceedingly rare.
Plowright mentions that, mixed with the uredospores, he found a large
number of hyaline capitate paraphyses; but Fischer states that the
specimen in Sydows’ Uredineen (no. 458) has no paraphyses and none are
mentioned in the description. In a doubtful specimen on Anthoxanthum
from Switzerland, however, Fischer found paraphyses. McAlpine also
found no paraphyses, but occasional mesospores, and states that the teleu-
tospores are much more uncommon than the uredospores. He found his
specimens on sheath and inflorescence of A. odoratum, as well as on the
leaves. The only British specimens I have seen are those collected at
King’s Lynn by Plowright in 1884, but it is also recorded for Yorkshire.
The suggestion that this is a hetercecious species has so far received no
confirmation.

DistrisuTion: Belgium, Germany, Australia.

125. Puccinia perplexans Plowr.

P. perplexans Plowr. Quart. Journ, Mier. Sci. xxv. 164; Ured. p. 179;
Hedw. 1886, p. 38; Grevillea, xiii. 53. Sacc. Syll. vii. 632.
Sydow, Monogr. i. 719. McAlpine, Rusts of Australia, p. 127,
f, 23.

cidiospores. Aicidia hypophyllous and on the petioles,
clustered on roundish yellow spots, somewhat cylindrical, with
a white incised margin; spores faintly verruculose, orange,
18—27 pw diam.

ON GRAMINEZ 271

Uredospores. Sori amphigenous, scattered, roundish, ob-
long or linear, occasionally con-
fluent, minute, yellow-brown ;
spores globose to ovate, faintly
echinulate, yellow, 20—28 w
diam. (with 4—6 scattered
germ-pores on one face, McAl-
pine).

Teleutospores. Sori amphi-
genous, scattered, occasionally
confluent, minute, generally
oblong or linear, about 1—1} Fig. snr tla
mm. long, always covered by :
the epidermis, black; spores variable, generally oblong to
clavate, rounded, truncate or obliquely attenuated at the apex,
slightly thickened (3 ) and darker, gently constricted, narrowed
below, smooth, brown, 36—57 x 18—24 yu, with a very short
pedicel.

fEcidia on leaves and ‘petioles of Ranunculus acris; uredo-
and teleutospores on Alopecurus pratensis. Very uncommon,

(Fig. 206.)

The connection of the ecidium with the Puccinia was first demon-
strated by Plowright, and has since been confirmed by Dietel and by
Klebahn. All the teleutospores I have seen were full of a very coarsely
granular protoplasm. It must not be forgotten that an ecidium on
Ranunculus acris belongs also to Uromyces Dactylidis,

DISTRIBUTION: Holland, Germany and Australia.

126. Puccinia Magnusiana Korn.

eidium Ranunculacearum DC. ; Cooke, Handb. p. 539: Mier, Fung.
p. 196 p.p.

Puccinia graminis var. Arundinis Cooke, Handb, p. 493; Micr. Fung.
p. 202.

P. Magnusiana Korn. in Hedwig. 1876, p. 179. Phill. and Plowr. in
Grevillea, xiii. 53. Plowr. Ured. p. 177; Proc. Roy. Soc, Lond.
xxxvi. 47; Quart. Journ. Mier, Sci. xxv. 156. Sace. Syll. vii. 631.
Sydow, Monogr. i. 785. Fischer, Ured. Schweiz, p. 241, f. 189.
McAlpine, Rusts of Australia, p. 125, f. 18.

272 PUCCINIA

Acidiospores. Aicidia hypophyllous in small clusters on
yellowish spots, or on the petioles or stems forming elongated
groups, cup-shaped, with a cut white margin; spores densely
and finely verruculose, yellowish, 15—25 p.

Uredospores. Sori amphigenous, scattered, rarely confluent,
elliptical or oblong, 1—2 mm. long, pulverulent, pale yellowish-
brown ; spores mostly ovate or ellipsoid, delicately echinulate,
pale brownish-yellow, 20—85 x 12—20; germ-pores indis-
tinct ; paraphyses numerous, clavate, hyaline or pale brownish.

a

5

Fig, 207, P. Magnusiana. a, Teleutospores (one abnormal); b, paraphysis
with uredospores; ¢, sori on leaf, and d, sori on stem of Phragmites.

Teleutospores. Sori amphigenous, very numerous, usually
scattered over the whole leaf-surface, oblong or sublinear,
minute, 1—2 mm. long, or on the culms and leaf-sheaths forming
narrow striz several centimetres long, flat, compact, persistent,
blackish: spores oblong or clavate, rounded above or rarely
conically attenuate or truncate, with .a distinct thickened cap
(5—10 p), hardly constricted, attenuated downwards, smooth,
brown, darker above, 32—55 x 16—26 4; pedicels thick,
brownish, persistent, as long as or shorter than the spores.

Aeidia on Ranunculus bulbosus, R. repens, April—June or
even July and August; uredo- and teleutospores on Phragmites
communis, June—April. Not uncommon in suitable localities.

(Fig. 207.)

ON GRAMINEE 273

It was Plowright who first proved, by a long series of cultures, that
the ecidium of this species is produced only on the two species of
Ravunculus given above; Klebahn and Fischer have since abundantly
confirmed his results. The ecidia belonging to Uromyces Poae and
U. Dactylidis occur on the same hosts and are morphologically quite
indistinguishable, but begin to appear earlier in the spring.

P. Mugnusiana is distinguished from the two following species by its
numerous small teleuto-sori, and the abundant paraphyses mixed with
its uredospores; the teleutospores also show hardly any constriction.
McAlpine describes mesospores 28—38 x 13—19 yu, and says that the apex
of the paraphyses in the Australian specimens is of a dark smoky-brown.
This species and P. Phragmitis may occur together upon the same leat.

DistRisuTion: Europe, South Africa, Japan, Australia.

127. Puccinia Phragmitis Korn.

Evidium rubellum Gmel. Syst. Nat. ii. 1473. Cooke, Handb. p, 544 ;
Micr. Fung. p. 199 p.p. Purton, Midl. Flor. iii. 294, pl. 26.

Uredo Phragmitis Schum. FI. Sall. ii. 231.

Pucetnia arundinacea Hedw. in DC. Flor. fr. v. 59. Cooke, Handb.
p. 493; Micr. Fung, p. 202.

P. Phragmitis Korn. in Hedwig. 1876, p. 179. Plowr. Ured. p. 175.
Sacc. Syll. vii. 630. Sydow, Monogr. i. 787. Fischer, Ured.
Schweiz, p. 250, f. 192.

Spermogones. Whitish.

Aeidivspores. Aicidia hypo-
phyllous, on circular red or deep J
purple spots 4—I}cm. diam., in
dense clusters, shortly cylindrical or
cup-shaped, with a cut white re-
curved margin; spores verruculose,
nearly hyaline, 16—26»; spore-
mass white.

Uredospores. Sori amphige-
nous, scattered or subgregarious,
elliptical, lanceolate or linear, some-
times confluent, rather large, con-
vex, pulverulent, brown, without _

Fig. 208. P. Phragmitis. Teleu-

paraphyses ; spores subglobose OF tospores and uredospore; u,
obovate, verruculose rather than %ori on leaf of Phragmites.

G U. 18

274 PUCCINIA

echinulate, brownish, 25—35 x 16—26 ; contents colourless ;
epispore rather thick, with four equatorial germ-pores.
Teleutospores. Sori numerous, similar, but larger and
thicker, very convex and compact, black ; spores oblong, rounded
at both ends, thickened (4—9 yu) above, constricted, smooth,
deep yellowish-brown, 45—65 x 16-—25 u, sometimes 75 uw long;
pedicels hyaline or yellowish, thick, persistent, 100—200 p long.

AKcidia on Rumex acutus, R. crispus, R. conglomeratus,
R. Hydrolapathum, R. obtusifolius, Rheum officinale, May and
June; uredo- and teleutospores on Phragmites communis,
July—May, often in the same sori. Not common, except
locally. (Fig. 208.)

It was Plowright who first showed that P. Phragmitis has its ecidium,
not on Ranunculus, but on species of Rumex and Rheum (not, however,
on Rumex Acetosa). Klebahn and Fischer have confirmed his results,
and Arthur has done the same for the North-American forms. It is a
remarkable fact, however, that the scidium had not been previously
found in North America until Arthur obtained it artificially by infection
of Rumex crispus and R. obtustfolius with the teleutospores of P. Phrag-
mitis, Afterwards it was found in Nebraska on various species of Rumezx
and Rheum. It is suggested by Sydow that this species is dispensing
with the ecidium, in which case it must winter by means of its uredo-
spores. Even in England the e«cidium seems relatively scarce, but it is
very conspicuous, and can be found on Rumex growing amidst Reeds.

DISTRIBUTION: Europe, South Africa, Japan, North America,
Chili.

128. Puccinia Trailii Plowr.

Aicidium rubellum Gmel.; Cooke, Handb. p. 544; Micr. Fung.

p. 199 p.p.
Puceinia Traalii Plowr. Ured. p. 176. Sace. Syll. ix. 312. Sydow,
Monogr. i. 790. Fischer, Ured. Schweiz, p. 252, f. 193.

Ajcidiospores. Similar to those of the last species; but the
spots are purple, surrounded by a yellow margin, the ecidia are
wider and flatter, and the spores are on the average somewhat
larger.

Uredospores. Sori amphigenous, scattered, rather large,

ON GRAMINEA 975

elliptical or linear, reddish-brown, pulverulent, without para-
physes; spores subglobose or
ovate, échinulate, brownish, 25—
35 x 20—25 pu.

Teleutospores. Sori similar,
but larger (2—4 mm.), compact,
pulvinate, black’; spores oblong,
rounded at both ends, with a cap-
like thickening (5—10 yz) above,
plainly constricted, brown, 50—
60 x 20—23 w; pedicels brownish,
thick, persistent, 75—100 yw long.

fcidia on Rumex Acetosa,
May and June; uredo- and teleu-
tospores on Phragmites communis,
from July. Rare. (Fig. 209.)

The results of Plowright’s cultures
have been confirmed by Klebahn, but
at any rate this species is very closely
allied to the previous one, and should

; : : Fig. 209. P. Trailii. Teleuto-
rather be consider ed as merely a spores, and sori on leaf of Phrag-
biological race of it. mites.

XE

DisTRIBUTION: Holland, Germany.

129. Puccinia Agrostidis Plowr.

Hieidium Aquilegiae Pers. ; Plowr. Ured. p. 263.
42, Ranunculacearum var. Aquilegiae DC. ; Cooke, Handb. p. 539.
P. Agrostidis Plowr. Gard. Chronicle, 1890, ii. 139; Grevillea, xxi.
110. Sacc. Syll. xi. 202. Sydow, Monogr. i. 717. McAlpine,
Rusts of Australia, p. 114, f. 27(2). Fischer, Ured. Schweiz,
p. 353, f. 257.
Spermogones. Honey-coloured, on round spots.
Ajcidiospores. Aicidia on rather large roundish yellow spots
which are often thickened and margined with brown, crowded,
hypophyllous, shortly cylindrical, with a torn white margin ;
spores faintly verruculose, orange, 16—30 x 14—20 pu.
Uredospores. Sori amphigenous, on yellow spots, elongated
or linear, about 1mm. long, bright-orange; spores globose to

18—2

276 PUCCINIA

ovate, yellow, faintly echinulate, 20—25 x 16—22 ~; membrane
colourless.

Teleutospores. Sori hypophyllous, minute, covered by the
epidermis, oblong or linear, rarely confluent, black; spores
somewhat clavate, rounded, truncate or gently narrowed above,
slightly thickened (about 5), faintly constricted, narrowed
below, smooth, brown, darker upwards, 38—48 x 12—20,;
pedicels very short.

Acidia on Aquilegia vulgaris, Lewes, Sussex ; Lake Win-
dermere; Wyre Forest, etce., May, June; uredo- and teleuto-
spores on leaves and sheaths of Agrostis alba, A. vulgaris,
August. Uncommon. The Puccinia should be looked for on
the grass near the place where the eecidium was seen.

The connection between the ecidium on Aguilegia and the Puccinia
on Agrostis was first demonstrated by Plowright (see Gard. Chron. 1890,
ii, 139, and 1891, i. 683); the fact has since been confirmed and extended
to Aqualegia alpinu. McAlpine’s species seems to be rather different ; he
records numerous mesospores, and uredospores with as many as nine
germ-pores, circularly arranged, on one face.

DistRIBUTION : Central and Western Europe, Siberia, India,
and (?) Australia.

130. Puccinia Molinie Tul.

Aieidium Melampyri i. et 8. exsicc. no. 165.

Puceinia Moliniae Tul. Ann. Sci. Nat. ser. 4, ii, 141, pl. ix, f. 9—11.
Cooke, Grevillea, v. 57; Micr. Fung. p. 203. Plowr. Ured. p. 179.
Sacc. Syll. vii. 631. Sydow, Monogr. i. 762. Fischer, Ured.
Schweiz, p. 256, f. 195.

P. nemoralis Juel, Ofvers. K. Vet.-Akad. Forh. 1894, p. 506, f. a—f.

P. Heidti-Melampyri Livro, Act. Soc. Faun. et Flor. Fenn. xxix. 55.

[4icidiospores. Aicidia hypophyllous, clustered on roundish
red or purple spots 3—5 mm. diam., cup-shaped, with a cut
white revolute margin; spores very minutely verruculose,
yellowish (?), 15—18 p.]

Uredospores. Sori amphigenous, generally hypophyllous,
often on brownish or purplish spots, scattered or arranged in
lines and confluent, oblong or linear, brown; spores more or less

ON GRAMINE 277

globose, aculeate, yellow-brown, 20—28 x 20—24 4; epispore
3—6 wu thick, with three germ-pores.
Teleutospores. Sori similar, often confluent and as much as
8 mm. long, conspicuous, pulvi-
nate, black; spores ellipsoid,
rounded at both ends, slightly
thickened (up to 5) above,
hardly constricted,smooth, brown, )
32—46 x 20—30 yw; pedicels hya-
line or yellowish, curved, per-
sistent, rather thin, very long (as
much as 120); a few mesospores
sometimes intermixed.

{[Aicidia on Melampyrum
spp.]; uredo- and teleutospores
on Molinia coerulea, July—Oc-
tober, Perthshire (Dr B. White).
This zcidium is not recorded for
Britain, and appears to be very
rare everywhere. (Fig. 210.)

Plowright, relying upon the experi-
ments of Rostrup, connected the
ecidiuin on Orehis latifolia with this
Puccinia, though he himself could not
succeed in the infection. Others have
similarly failed, and there seems to be a ee ene re
little doubt that Rostrup’s conclusions tiers, Raniioeh (Dr Bashanal
were inaccurate. Juel has since then White).
succeeded in showing that an ecidium
on Melumpyrum pratense is part of the life-cycle of a Puceinia on Molinia,
which he named P. nemoralis, but of which there is no proof that it is
different from P. Moliniae Tul. (Juel, Z.c.). Liro confirms this result and
names his. species P. Meidii-Melampyri (l.c.). The ecidiospores are
described by Sydow as “yellowish,” but Juel describes them as colourless,
like those of the allied species P. Phragmitis and P. Trailit.

Since the ecidium on Melampyrum has not been found in this country,
the British species may turn out, on investigation, to be different from
these. For there is a closely allied species or biological race, named by
Cruchet (Centralbl. f. Bakter. 2. xiii. 96) P. Brunellarum-Moliniae, which
has teleutospores very like those of P. JJoliniae Tul. but its ecidium on

278 PUCCINIA

Brunella (= Heidiuin Prunellae Wint.). This was found by Fischer in
Switzerland, and the connection of the two hosts was proved by Cruchet.
To this, doubtless, belongs the ecidium found on Prunella vulgaris by
Dr Keith at Forres (Plowr. Ured. p. 264), and Qr Buchanan White's
Puccinia on Molinia may belong there likewise. The apex of the teleuto-
spores in his specimens is, however, much less strongly thickened than in
the figures given by Fischer.

DIsTRIBUTION: Throughout Europe, except the south.

131. Puccinia Poarum Niels.

Aeidium Tussilaginis Gmel. in Linn. Syst. Nat. ii. 1473.

AG, Compositarum var. Tussilaginis Cooke, Handb. p. 542; Mier.
Fung. p. 198.

Puccinia Poarum Niels. in Botan. Tidsskr. ii. 26. Phill. et Plow.
Grevillea, xiii. 54, Plowr. Ured. p. 168; Grevillea, xi. 52. Sace.
Syll. vii. 625. Sydow, Monogr. i. 795. Fischer, Ured. Schweiz,
pp. 361, 556, f. 268. McAlpine, Rusts of Australia, p. 128, f. 22.

Spermogones. Epiphyllous, pale-yellow, often very numer-
ous.

Micidiospores. Aicidia hypophyllous, usually in dense
clusters on circular yellowish or reddish thickened spots
1—2 om. diam., seldom scattered, cup-shaped, with a dentate
white revolute margin; spores verruculose, orange, 18—25 x
16—20 p.

Uredospores. Sori on the leaves, sometimes on the culms,
minute, roundish or elliptic, soon naked, yellow ; spores globose
to ellipsoid, densely and minutely verruculose, yellow, 17—28 x
17—25 y, with about five scarcely perceptible scattered germ-
pores, and intermixed with numerous, hyaline, capitate para-
physes.

Teleutospores. Sori similar, oblong or linear, more or. less
in short rows, long covered by the epidermis, surrounded by
a small pale area, black; spores oblong-clavate, cylindrical, or
obconical, variable, rounded, truncate, or rarely conically at-
tenuated above where they are slightly thickened (4—8 p),
hardly or not at all constricted. more or less tapering below,
smooth, chestnut-brown, becoming gradually paler downwards

ON GRAMINEE 279

30—45 x 16—22 4; pedicels short, brownish, persistent; an
occasional mesospore is found.

ns.

N0

Fig. 211. P. Poarum. Teleutospores ; a, uredospore on P. nemoralis; b, para-
physes with same; c, mesospore; d, teleuto-sori on P. pratensis; e, typical
teleuto-sori of Uromyces Poae, on the same.

fEcidia on Tussilago Farfara, about May, June, and August,
September, very common; uredo- and teleutospores on Poa
annua, P. nemoralis, P. pratensis, P. trivialis, about July,
August and October—December, common but easily overlooked
unless searched for. (Fig. 211.)

First stated by Nielsen, the connection of the two hosts was demon-
strated by Plowright and Klebahn. This hetereecious Puccinia differs
from all others in having two generations in one year. The earlier crop
of wcidia begins to appear in May, and is followed by the uredo- and
teleutospores on the surrounding leaves of Poa; these germinate quickly
and the second crop of ecidia is produced about August, and the second
generation of teleutospores may be found on Poa from October. The
latter germinate in the following spring, but according to Lagerheim the
uredospores algo, in a favourable climate, can survive the winter. This is
certainly true in Australia, where (though the Puccinia is an introduced
one) the uredospores have been found the whole year round. In that
country the Coltsfoot does not exist, and the fungus is carried through
the winter by the uredo-stage; in fact, according to McAlpine, it is most
common in the winter there. Arthur and Carleton say that the fungus
does the same as far north as Nebraska in North America, where the
Coltsfoot is only a naturalised plant. Uredospores were found alive in
every month of the year at Washington, D.C.

In the Scottish Naturalist (’84, p. 270) this species is recorded for Poa
Jluitans, but there may possibly be some error in this statement. Plow-
right says that the uredospores are not accompanied by paraphyses,

280 PUCCINIA

though others find them habitually. I myself have always found para-
physes (capitate, but not ‘“ stiff”) in the uredo-sori. McAlpine records
three- and even four-celled teleutospores in Australia, and I have found a
very few mesospores in the sori.

Perhaps the easiest way to obtain the teleutospores is to search the
lower leaves of species of Poa, growing round leaves of Coltsfoot, as soon
as ecidia of the second crop are perceived upon the latter towards the
end of July or beginning of August.

DistTRIBUTION: Europe, Japan, North America, Australia.

132. Puccinia Baryi Wint.

Epitea Baryi Berk. et Br. Ann. Mag. Nat. Hist. 1854, no. 755.

Lecythea Baryi Berk. ; Cooke, Handb. p. 532 ; Mier. Fung. p. 222.

Puceinia Baryi Wint. Pilz. Deutsch. p. 178. Plowr. Ured. p. 191.
Sace. Syll. vii. 660. Sydow, Monogr. i. 737. Fischer, Ured.
Schweiz, p. 369, f. 267.

Uredospores. Sori mostly epiphyllous, on hnear brown spots,
scattered or in groups, often disposed in long linear series,
minute, elongated, reddish-brown; spores globose to obovate,
delicately verruculose, yellow, 18—25 ~; paraphyses numerous,
clavate to capitate.

Fig. 212. P. Baryi. Teleutospores; a, abnormal teleutospore ;
b, paraphysis; c, uredospore; all on B. silvaticum.

Teleutospores. Sori similar, but long covered by the epi-
dermis, blackish-brown ; spéres very irregular, ellipsoid or sub-
clavate or pyriform, obtuse or truncate and slightly thickened
and undulated above, hardly constricted, somewhat attenuated
below, smooth, clear-brown, darker above, 25—40 x 15—25 p;
pedicels none or very short, brownish, darker at base of spore
where the transverse wall is much thickened.

ON GRAMINEA 281

On leaves (living or fading) of Brachypodium pinnatum,
B. silvaticum. Not uncommon. July—November, teleuto-
spores not before September. (Fig. 212.)

Both Plowright and Fischer mention that, mixed with the uredospores,

are numerous hyaline capitate paraphyses ; Sydows’ Monographia omits
all mention of these. The specimens which I have seen show them always

in great numbers. ’

Often the pedicel of the teleutospores is almost non-existent, and the
basal cell-wall is strongly thickened. The upper margin of the teleuto-
spore is often undulated ; occasionally one is met with having three
cells. An ecidium, though often suggested, has not yet been discovered

for this species.

DiIsTRIBUTION : Central and North-Western Europe.

133. Puccinia Agropyri Ell. et Ev.

Aeidium Clematidis DC. Flor. fr. ii. 243, Plowr. Ured. p. 265.

Zi. Ranunculacearum var. Clematidis Cooke, Handb. p. 539.

Puceinia Agropyri Ell. et Ev. in Journ. Mycol. vii. 131. Sace. Syll. xi.

201. Sydow, Monogr. i. 823. Fischer, Ured. Schweiz, pp. 350, 555,
f. 255. McAlpine, Rusts of Australia, p. 113.

Spermogones. Amphigenous.

Aicidiospores. AMcidia hypophyllous and on the petioles
and stems, usually on brownish spots, causing considerable
distortion, scattered or in clusters of very varied size, shortly
cylindrical, with white torn
broadly revolute margin; spores

verruculose, orange, 18—27 p.
[Uredospores. Sori amphi-
genous but generally hypophyl-
lous, on irregular yellow spots,
scattered, oblong or linear, 1—
14 mm. long, cinnamon; spores -

more or less globose, delicately
echinulate, pallid-yellow, 19— Fig. 218. P. dyropyri. Teleuto-
e : spores and mesospure, on d. repens,

27 4; epispore rather thin, with — Pomerania (ex herb. Sydow).
three or four germ-pores.

Teleutospores. Sori epiphyllous, scattered, sometimes con-
fluent, oblong or linear, as much as 3 mm. long, covered always
by the lead-coloured epidermis, black; spores cylindric-clavate;

282 PUCCINIA

usually truncate above and thickened (up to 6), gently con-
stricted, attenuated downwards, smooth; pale-brown, darker at
the apex, 40—80 x 11—22; pedicels very short, persistent,
somewhat hyaline, with a dark-brown band at the apex.]

cidia on Clematis Vitalba, May—July ; [uredo- and teleuto-
spores on species of Triticum (Agropyron), but not recorded as
yet for Britain]. (Fig. 213.)

Dictel first showed, by culture experiments, that the ecidium on
Clematis Vitalba is connected with a Puccinia on Triticum glaucum.
In Europe the ecidium is frequently observed (probably because of the
striking distortions it produces), but the Puccinia has been far more
rarely noted, being by no means conspicuous in its appearance. According
to Sydow, it occurs also on Triticum junceum and T. repens, on which it
may presumably be found in this country, if looked for.

The statement of Rathay, that Herdium Clematidis belongs to a
Melampsora on Poplar, is now universally discredited. It has been
observed on many (nineteen) species of Clematis, but it must be noted, as
remarked by Sydow, that the genus Z7yiticum is also very widely distributed,

According to Fischer, the teleutospores are often accompanied by
brown paraphyses, which subdivide the sori into smaller compartments,
something like what obtains in P. dispersa and P. persistens. Other
authors have not found the paraphyses, but they are easily overlooked ;
abnormal three-celled teleutospores are recorded. Probably more than
one species has been included under the name. P. agropyrina Eriks.
is distinctly different.

DistRisuTion: Europe, North and South America, Turk-
estan, Japan, Australia.

134. Puccinia persistens Plowr.

Meidium Ranunculacearum var. Thalictri Cooke, Handb. p. 540.
Puceinia persistens Plowr. Ured. p. 180. Sacc. Syll. ix. 312. Sydow,
Monogr. i. 825. Fischer, Ured. Schweiz, pp. 347, 555, f. 254.

Spermogones. Epiphyllous, in little clusters, orange.

cidiospores. Aicidia hypophyllous, in clusters on thick-
ened spots which are purple-brown above and yellow with a
brownish border below, urn-shaped or subcylindrical, yellow,

with a torn white margin; spores minutely verruculose, orange,
14—28 uy.

ON GRAMINE 283

Uredospores. Sori minute, roundish or elongated, orange,
on yellowish spots; spores more or
less globose, minutely echinulate,

yellow, 25—80 p. ( 7
Teleutospores. Sori hypophyl-

lous, minute, ovate, oblong or
linear, black, long covered by the
epidermis; spores clavate-oblong
or irregular, rounded, truncate
or obliquely attenuate above,
slightly thickened (4—7 1), more
or less constricted, rounded or
often tapering below, smooth,
brown, 50—60 x 15—-20 uw; pedicels short, hyaline, persistent.

Fig. 214. P. persistens.
Teleutospores.

cidia on Thalictrum flavum, T. minus, May—July ; uredo-
and teleutospores on Agropyron repens, from July onwards.
(Fig. 214.)

The connection of these spore-forms was ascertained by Plowright and
confirmed by Rostrup. The secidia on various other species of Thalictrum
may belong to the same Puccinia or to Puccinias on Agropyron caninum,
Arrhenatherum elatius or even on species of Poa. There is abundant scope
here for further investigation. According to Fischer the teleuto-sori
are subdivided into compartments by groups of paraphyses, in the
same way as in P. Agropyrt. In fact the three species, P. persistens,
P. Agropyrt, and P. Actaeae-Agropyri Fischer, form a natural group
which should be regarded rather as biological races of one species, having '
their teleutospores on Triticum (Agropyron) and their ecidia on
Thulictrum, Clematis and Actaea respectively ; the latter (P. Actaeae-
Agropyrt) has not yet been found in Britain.

DistTRIBUTION: Europe, Siberia, Japan, Himalaya, North
America.

The evidence for considering the three following species as
British is not yet sufficient :—

135. Puccinia Phlei-pratensis Erikss. et Henn.

Puccinia Phlei-pratensis Erikss. et Henn., in Zeitschr. fiir Pflanzenkr.
1894, p. 140; Getreideroste, pl. 5, f. 55—6. Sacc. Syll. xi. 204,
Sydow, Monogr. i. 784. Fischer, Ured. Schweiz, p. 260.

284 PUCCINIA

Uredospores. Sori on the leaves or most often on the
culms, on the latter and on the sheaths confluent in long lines
(as much as 8 cm. long) and.splitting the epidermis, on the
leaves more often scattered, minute and oblong, 1—2 mm. long,
pulverulent, dark yellowish-brown; spores ellipsoid to oblong
or even somewhat pear-shaped, aculeate, dirty-yellow, 18—30 x
15—20 pu.

Teleutospores. Sori similar, but.chiefly on the culms, black-
brown; spores clavate, rounded or conically attenuate above,
thickened (up to 8), constricted, attenuated below, smooth,
brown, 38—52 x 1420 w; pedicels brownish, persistent, rather
thick, as much as 60 uw long.

On leaves and culms of Phleum pratense (and P. nodosum) ;
also on Festuca elatior.

This was separated by the authors from P. gramznis on the ground
that it will not infect Berberis; Eriksson has shown that it will feebly
infect Phleum Micheli, Secule cereale, and Avena sativa, but not Triticum
vulgare, Hordeum vulgare or Poa pratensis. It chiefly occurs in the uredo-
stage, and winters thereby or by its mycelium; in many localities the
teleutospores are not formed at all. This may be one of the cases where
the ecidium has been dropped completely from the life-cycle. I have not
seen a British specimen, but it is extremely likely to occur here.

DisTRiBuTION: Germany, Austria, Denmark, Sweden. \\ it

136. Puccinia Arrhenatheri Erikss.

Heidium graveolens Shuttl. apud Cooke, in Bull. Soc. Bot. Fr. 1877,
xxiv. 315, Saec. Syll. vil. 778.

Puceinia perplexans Plowr. f. Arrhenathert Kleb. in Abhandl. Natur.
Ver. Bremen, 1892, p. 366.

P. Arrhenathert Erikss. in Cohn’s Beitrage zur Biologie d. PAlanz. 1898,
vi. 1. Sydow, Monogr, i. 729. Fischer, Ured. Schweiz, p. 345.

[Spermogones. Very numerous, minute, covering a great
part of the leaf or the whole leaf uniformly.

Beidiospores. .Kcidia hypophyllous, sometimes even on
the flowers, deforming the affected branches, generally distri-
buted densely and evenly over the whole leaf, between cylin-
drical and cup-shaped, with a torn whitish revolute margin:

ON GRAMINEE 285

spores subglobose or ellipsoid, verruculose, yellowish, 19—82 x
16—24 u.]

Uredospores. Sori epiphyllous, very rarely hypophyllous,
on minute yellow spots, elliptical or oblong, minute, pale rust-
coloured ; spores globose to ovate, echinulate, yellowish, 19—30
x 19—26 yw, with numerous germ-pores, and mixed with nume-
rous capitate paraphyses which are 10—14, broad and as

much as 80 uw long.
| a | c y

Fig. 215. P. drrhenatheri. a, teleutospores on drrhenatherum, from Hampton-
in-Arden ; b, another, and ¢, two abnormal ones, from Lichfield; d, a
paraphysis and uredospore from the latter.

Np

Teleutospores. Sori hypophyllous, scattered, minute, puncti-
form or shortly linear, covered
by the epidermis, black; spores

ellipsoid-oblong or oblong-

clavate, rounded, truncate or

rarely gently attenuated a-
bove where they are thickened
(5—10 yw) and darker, hardly \°/

or not at all constricted, gen- : 7

erally attenuated downwards, _.
1h ~ Fig. 216. P. dArrhenatheri. Teleuto-
smooth, pale-brown, 380—45 spores, paraphysis and uredospore,,

x 18—24y, mixed with brown- oa Mama Silesia (ex herb.
‘ 2 ydow).

ish paraphyses; pedicels short,

brownish.

[Aicidia on leaves and flowers of Berberis vulgaris ;] uredo-
and teleutospores on Arrhenatherum elatius. (Figs. 215, 216.)
This species produces on the Berberis its characteristic witches’ -brooms,

composed of many upright branches ; it was at first mistakenly identified
with . Magelhaenicum Berk. The ecidium has not been seen in Britain,

286 PUCCINIA

but I have found on Arrhenatherum on many occasions teleutospores and
uredospores which seem to be identical with those of this species, though
the former are, in my specimens, often irregularly three- or four-celled.

DistrRisuTion : Central and Northern Europe, Turkestan.

137. Puccinia paliformis Fckl.

Puccinia paliformis Fckl. Symb. Myc. p. 59, pl. ii. f.17. Plowr. Ured.
p- 203, Sacc. Syll vii. 731. Sydow, Monogr. i. 759, f. 534.
Fischer, Ured. Schweiz, p. 264, f. 200. ;

Teleutospores. Sori on the leaves and culms, scattered,
minute, roundish or oblong, up to
1mm. long, pulvinate, surrounded
by the cleft epidermis, black-
brown; spores clavate, usually
truncate above, more rarely round-
ed or conically attenuate, much
thickened (10—16 yp), hardly
constricted, tapering downwards,

Fig.217. P.paliformis(?). Uredo- smooth, pale-brown, 40—56 x 10—
spore and telenionpores, from 99 4, pedicels hyaline, persistent,
about as long as the spore.

On Koeleria cristata. Very rare. September and October,
near Aberdeen (Prof. Trail). (Fig. 217.)

There is much doubt about this fungus ; it was suspected by Winter,
on account of the likeness of its teleutospores to those of P. Carteczs, that
the original specimens on which the species was founded grew not on
Koelerta, but on Carer. It seems to have been recorded only twice, once
by Morthier in the Jura, in spring on old leaves of the previous year, and
once as above. The three figures quoted in the synonymy agree fairly
well, but appear to have been all taken from the same material, viz. that
gathered by Morthier, which according to Sydow may well be Koeleria and
not Carex. I have examined the specimens, preserved in Herb. Kew,
gathered by Prof. Trail; they are undoubtedly Aoeleria, they have split
sheaths and, though not in flower, agree perfectly with specimens of
Koeleria cristata from the Highlands, in the same herbarium. But they
bear uredospores in slightly swollen epiphyllous yellow patches, sunken in
the leaf (these are mentioned by Prof. Trail in the Scottish Naturalist,
1883, p. 85) ; therefore the fungus cannot be P. paliformis as described by
Fuckel, but is probably P. langissima, Schrét. I found the uredospores oval
or obovate, pale brownish-yellow, sparsely echinulate, 23-—24 x 17—18 py.

TRIPHRAGMIUM 287

It should be noted that there is an xcidium belonging to P. longissima, on
species of Sedum, formerly called by mistake Endophyllum Sedi. This
has been found on S. acre, and according to Mayor (Annal. Mycol. 1911,
* ix. 841) on S. reflexum, but not, so far as I know, as yet in Britain.
Further search in the Highlands will no doubt readily decide which of the
two species occurs there. ,

TRIPHRAGMIUM Link.

Auteecious.

Spermogones subcuticular, flattish, without ostiolar fila-
ments. Czomata indefinite, large, without paraphyses; uredo-
sori small, definite, encircled by paraphyses; spores in both
borne singly on pedicels: pores not evident. Teleuto-sori more
or less definite; spores coloured, radiately 3-celled, more or less
verrucose, with one pore in each cell, equidistant from the septa,
Le. apical.

It is most likely that the foreign species, usually classed
with ours on account of the form of the teleutospores, are not
closely allied. Spores of this particular shape are met with,
abnormally, in other genera, as in several species of Puccinia,
even in Puccinia graminis. Neither is this genus closely allied
to Phragmidium; the teleutospores have no gelatinous outer
coat, and the germ-pores are differently placed.

Triphragmium Ulmarie Wint.

redo Ulmariae Schum. Enum. PI. Sail. ii. 227.

Uromyces Ulmariae Lév.; Cooke, Micr. Fung. p. 212, pl. 7, f. 147—8.

Triphragmium Ulmariae Wint. Pilze, p. 225. Cooke, Handb. p. 492;
Micr. Fung. p. 202, pl. 3, f. 47—9. Plowr. Ured. p. 218, pl. 4, f. 6.
Sace. Syll. vii. 768. Fischer, Ured. Schweiz, p. 423, f. 293.
Arthur, N. Amer. Fl. vii. 178. Sydow, Monogr. iii. 171.

T. Filipendulae Pass. Nuov. Giorn. Bot. Ital. vii. 255. Cooke,
Grevillea, xi. 15. Plowr. Ured. p, 219. Massee, in Grevillea,
xxi. 115, Fischer, Ured. Schweiz, p. 425, f. 294. Sacc. Syll. vii.
769. Sydow, Monogr, iii. 174.

288 TRIPHRAGMIUM

Spermogones. On the leaves and petioles, circinate, flat,
yellowish.

Uredospores. Sori of two kinds—primary, Le. ceomata,
amphigenous, large, expanded, bright-orange, mostly on the
veins and petioles where they cause distortion, without para-
physes; secondary, hypophyllous, small, round, scattered, orange,
surrounded by paraphyses; spores brilliant-orange, ellipsoid to
obovate, verrucose, 25—28 x 18—21 y, without evident germ-
pores.

Fig. 218. J. Ulmariae. Normal telentospore ; a and b, two abnormal
ones; on 8. Ulmaria.

Teleutospores. Sori hypophyllous, small, round, brownish-
black, persistent, but pulverulent, sometimes arising in the
primary uredo-sori; spores subglobose, flattened, chestnut-
brown, more or less rough with obtuse warts, 35—49 w; each
cell has, at a point opposite to the inner corner, a germ-pore
round which the warts are often crowded; pedicels colourless,
persistent, variable in length; abnormal spores may have two
or four to five cells.

On Spiraea Ulmaria, S. Filipendula. Very common on the
former host. Primary uredospores, May—July; teleutospores,
August—November. (Fig. 218.)

The primary uredo-sorus may be Jooked upon as a ceoma, ie. an
ecidium, and in any case it corresponds to that developmental stage ;
Klebahn proved (Zeitschr. f. Pflanzenkr. 1907) that it is produced by
infection by the basidiospores. Dietel observed that, in elevated situations,
the secondary uredo-spore generation on S. Ulmaria was almost absent,
and the teleutospores arose in connection with the primary uredo-sori ;
this is in agreement with the usual shortening of the life-history that

PHRAGMIDIUM 289

takes place in arctic and alpine Uredinales. As Plowright remarks, the
teleutospores can be found in spring on last year’s leaves, and germinate
readily when placed in water.

The form on §. Filipendula, which is very uncommon in this country,
differs from that on S. Ulmaria in having the teleutospores usually smooth,
and only occasionally provided with a few warts round the germ-pores ;
but similar spores may be found on both hosts. In both, some of the
spores have three super-imposed cells, as in Phragmidium, also two cells
placed’as in Puccinia or laterally as in Diorchidium. The uredospores on
S. Filipendula are often pyriform and as much as 35 p long; it may bea
distinct species.

Arthur, who records 7. Ulmariae in the North American Flora on
Filipendula rubra, says that it is confined to one locality “ of less than
a half hectare extent,” at Lafayette, Indiana.

DISTRIBUTION : Europe, Siberia, Japan, Indiana.

PHRAGMIDIUM Link.

Autecious.

Spermogones subcuticular, conical or flattened, without
ostiolar filaments. Czomata indefinite, usually encircled by
incurved paraphyses; spores in chains, each with numerous
germ-pores. Uredo-sori definite, usually encircled by para-
physes; uredospores borne singly on pedicels, often with
paraphyses intermixed, pores numerous, scattered, indistinct.
Teleutospores two- to several-celled by transverse septa; wall
thick, laminate, usually coarsely verrucose, the middle layer dark
and rigid; pores two or more in each cell, placed laterally ;
pedicels often swollen below; basidiospores globose.

This genus is confined entirely to the family Rosacee.
There are many species in North America, but with the
exception of P. Potentiliae and those introduced on cultivated
roses they are all different from those of Europe. The warts
often found on the outer surface of the teleutospores are due to
the contraction of the external gelatinous layer, which can be
caused to swell up enormously by boiling in lactic acid. This

GU. 19

290 PHRAGMIDIUM

character, as well as the rigid middle lamina and the position
of the germ-pores, distinguishes the genus from all the neigh-
bouring ones.

l. Phragmidium Fragariastri Schrot.

Puccinia Fragariastri DC. Flor. fr. vi. 55.

Phragmidium obtusum Link, Sp. Pl. ii. 84 p.p. Cooke, Handb.
p. 491; Micr. Fung. p. 201 (as obtusatum).

P. Fragariastri Schrét. Flor. Schles. iii. 351. Plowr. Ured. p. 220.
Fischer, Ured. Schweiz, p. 412, f. 287.. Sacc. Syll. vii. 742.
Sydow, Monogr. iii. 101, f. 45.

Spermogones. In little clusters, honey-coloured.

4icidiospores. Cxomata mostly hypophyllous or on the

veins and petioles, irregular,
scattered, often confluent and
large, bri ght-orange, surrounded
by clavate paraphyses; spores
densely verruculose, orange-
yellow, 17—28 x 14—21 ps.
Uredospores. Sori hypo-
phyllous, scattered, roundish,
soon naked, surrounded by and
mixed with hyaline, thin-walled
Fig. 219. Ph. Fragariastri. Uredo- ee itate palep hy BES “BpOIeS
spore, paraphysis, and teleutospore ; roundish, densely verruculose,
ae teleutospore boiled in lactic orange-y ellow, 18—24 be
Teleutospores. Sori hypo-
phyllous, scattered, minute, roundish, pulverulent, brown; spores
cylindrical or rarely somewhat clavate, of 2—5 (mostly four)
cells, rounded, at both ends, sometimes slightly thickened and
paler at the summit but never papillate, faintly constricted,
rather pale-brown, 45—70 x 2228 y, sometimes with a few
delicate warts which are more abundant towards the apex, but
generally quite smooth ; usually three germ-pores to each cell ;
pedicels colourless, 14—21 w long.

a

On Potentilla Fragariastrum (=P. sterilis), and possibly
on other species of the genus, but never on Fragaria vesca.
March—October. Very common. (Fig. 219.)

PHRAGMIDIUM 291

The uredospores of this species are distinguished from those of its
allies by being densely and rather coarsely verruculose and very similar to
the cz#oma-spores, from which, in fact, they differ almost solely in being
abstricted singly and not in chains. The ceoma-stage is one of the earliest
Uredines of spring, showing on the leaves as soon as they are well
developed, and extending even to the calyx. The teleutospores are
entirely devoid of papilla on the apical cell; the gelatinous outer coat
is sometimes almost non-existent, and the spores are but slightly changed
by boiling in lactic acid.

DisTRIBUTION: Europe.

2. Phragmidium Potentille Karst.

Puceinia Potentiliae Pers. Syn. p. 229.

Phragmidium Potentillae Karst. Fung. fenn. iv. 49. Plowr. Ured.
p. 221. Fischer, Ured. Schweiz, p. 410, f. 286. Sacc. Syll. vii.
743, McAlpine, Rusts of Australia, p. 188 (2). Arthur, N. Amer.
Fl. p. 174. Sydow, Monogr. iii. 97.

Spermogones. Few, amphigenous, surrounded by the ecidia.
Aicidiospores. Czomata as in P. Fragariastri.

Fig. 220. Ph. Potentillae. a, teleutospore x 360; b, the same x 600; c¢, the
same, boiled in lactic acid for one minute; d, a four-celled teleutospore,
boiled and distorted by pressure.

Uredospores. Sori hypophyllous, roundish, often confluent,
at first covered by the swollen epidermis, surrounded by abun-
* Ke, * . >
* dant, clavate, curved paraphyses; spores ellipsoid to obovate,
finely echinulate, yellow, 21—24 x 16—19 yp.
Teleutospores. Sori hypophyllous, orbicular, soon naked,

19—2

292 PHRAGMIDIUM

black; spores cylindrical or subclavate, of 3—6 cells (occasionally
one or two), rounded or bluntly papillate at the apex, hardly
constricted, smooth, brown, 42—80 x 20—28 w%; two or three
germ-pores to each cell; pedicels thick, hyaline, persistent, as
long as or much longer than the spores (100—150 y).

On Potentilla argentea, P. verna, and various cultivated
species. April—September. Not common. (Fig. 220.)

This species is more closely allied to P. Sanguisorbae than to
P. Fragariastri. The finely echinulate uredospores and the papillate
teleutospores distinguish it from the latter.

DistTRiBUTION: Europe, Asia Minor, Siberia, Japan, North
America, Australia (?).

3. Phragmidium Sanguisorbe Schrit.

Puceinia Sanguisorbae DC. Flor. fr. vi. 54.
Lecythea Poterti Lév. ; Cooke, Micr. Fung. p. 221, pl. 3, f. 31.
Phragmidium Sanguisorbae Schroét. Flor. Schles. p. 352. Plowr. Ured.

p. 221. Fischer, Ured. Schweiz, p. 408, f. 285. Sace. Syll. vii.

742. Sydow, Monogr. iii. 156. i
P. acuminatum Fr. ; Cooke, Handb. p. 490; Micr. Fung. p. 201, pl. 3,

f. 30, 32,

Spermogones. Amphigenous, flat, clustered.

Aicidiospores. Czomata amphigenous, oblong, circinate
round the spermogones, or irregular
and swollen on the nerves and petioles ;
spores verruculose, orange-yellow, 18—
214; paraphyses curved.

Uredospores. Sori small, scarcely
¢mm., soon naked, surrounded by a
circle of clavate, curved paraphyses ;
spores globose to ovate, echinulate,
orange-yellow, 18—24 p.

Teleutospores. Sori punctiform, +—
1mm., soon naked, black ; spores eylin-
drical-oblong, of 2—5 (mostly four) cells,
apex drawn out into a papillate beak,
Fig.221, Ph. Sanguisorbae. faintly constricted, base rounded, some-

a, teleutospore x 360 ; b,
teleutospore x 600, what verruculose, yellowish-brown, 56—

PHRAGMIDIUM 293

70 x 21—244; about three germ-pores, to each cell; pedicels
hyaline, rather short.

On Poteritwm Sanguisorba. Rather common. Cxomata,
April—June ; teleutospores, July—November. (Fig. 221.)

DIsTRIBUTION: Europe, Algeria, Asia Minor, Turkestan.

4. Phragmidium disciflorum James.

Ascophora disciflora Tode, Fung. Meck. sel. i. 16 (see, Sydow).

Coleosporium pingue Lév.; Cooke, Handb. p. 520; Micr. Fung. p. 217.

C. miniatum Pers. ; Cooke, Micr. Fung. p. 217.

Lecythea Rosae Lév. ; Cooke, Micr. Fung. p. 221, pl. 3, f. 37.

Phragmidium bullatum West. ; Cooke, Micr. Fung. p. 202. ,

P. mucronatum Fr.; Cooke, Handb. p. 490; Micr. Fung. p. 201, pl. 3,

f, 38.

P. subcorticium Wint. Pilze, p. 228. Plowr. Ured. p. 224. Fischer,
Ured. Schweiz, p. 400, f. 281. Sace. Syll. vii. 746 p.p. McAlpine,
Rusts of Australia, p. 188, f. 229—233, and pl. I, f. 37. Arthur,
N. Amer. Flor. vii. 172.

P. disciforum J ames, Contr. U.S. Nat. Herb. iii. 276. Sydow, Monogr.
iii, 115.

Spermogones. Flat, subcuticular, pale honey-yellow.

Aicidiospores. Czomata on branches, petioles, leaf-nerves
and fruits, often confluent on the branches.for long distances,
on the leaves mostly roundish, bright-orange; the smaller ones
are surrounded by a circle of clavate, thin-walled, colourless
paraphyses ; spores in short chains, verruculose, orange-yellow,
24—28 x 18—21 yp.

Uredospores. Sori hypophyllous, very small, scattered,
round, soon naked, pale-orange, surrounded by a circle of
clavate, curved, colourless paraphyses ; spores ellipsoid or ovate,
yellow, echinulate, 21-28 x 14—20 uw, with numerous germ-
pores.

Teleutospores. Sori similar, but black; spores ellipsoid to
subfusiform-cylindric, of 6—8, rarely five or nine cells, with a
pointed papilla which is pale above and darker and scabrous
below, not constricted, rounded at base, unevenly warted, red-
brown, then blackish-brown, 65—120 x 30—45 w; each cell
with two or three germ-pores; pedicels colourless, persistent,
about as long as the spore, swollen at the base.

294 PHRAGMIDIUM

On Rosa canina, R. spinosissima, and many kinds of culti-

Fig. 222. Ph. disciflorum. a,
secidiospore, on cultivated Rose ;
b, uredospore, and c, paraphyses,
on Dog Rose; d, a small teleuto-
spore, on the same; e, a teleuto-
spore, on Burnet Rose; all x 600.

vated roses. Very common. Ce-
oma-spores in May and June;
teleutospores, August—October.
(Fig. 222.)

The teleutospores can germinate in
spring, and produce the ceoma (Jacky),
but they germinate with difficulty
(Miiller); the parasite is propagated
chiefly by the mycelium of the czoma-
stage, which passes the winter in the
branches and in spring bursts out into
wide-spreading spore-beds. These at-
tack and destroy the buds ; afterwards
the uredo- and teleutospores appear on
the leaves, with localised mycelium. The
infested branches are often swollen and
deformed. To keep the disease under
control, all teleutospore-bearing leaves
and the branches which are permeated
by the mycelium should be collected
and burnt, and the plant may be
sprayed, during the winter, with copper
sulphate solution.

It is noteworthy that this fungus
occurs in North America only on cul-
tivated roses; the indigenous Rose-

Phragmidia are all different species. It has been introduced, probably by,
cuttings, also into Australia and other parts of the world.

DIstTRIBUTION: Europe, Asia Minor, Persia, Africa, North
America, South America, Australia.

5. Phragmidium fusiforme Schriter.

Uredo pinguis var. Rosae-alpinae DC. Flor. fr, ii. 235.

P. fusiforme Schrét. Rostpilze Schles. p. 24. Fischer, Ured. Schweiz,
p. 404, f. 283. Sacc. Syll. vii. 747. Sydow, Monogr. iii. 121.

Phragmidium Rosae-alpinae Wint. Pilze, p. 227. Plowr. Ured. p. 226.

Aicidiospores. Sori on the petioles, nerves and fruits,
similar to those of P. disciflorum, but not so extensive; on
the leaves, punctiform and surrounded by hyaline clavate para-
physes ; spores echinulate, 17—30 x 15—20 p.

PHRAGMIDIUM

295

Uredospores. Sori small, punctiform, yellow; parapbyses

like those of the last species; spores roundish,
echinulate, yellow, 18—21 p.

Teleutospores. Sori similar, with similar
paraphyses, in little clusters, black ; spores
cylindrical or fusiform, of 8—13 cells, atten-
uated upwards into a pale horny conical
process, not constricted, rounded below,
verrucose, dark-brown, 80—105 x 21—24 p ;
each cell with two or three germ-pores ;
pedicels colourless, persistent, sometimes
longer than the spore.

On Rosa alpina. Rare ; Scotland (intro-
duced). June—October. (Fig. 223.)

This species is very abundant on &. alpina in
Switzerland. It is distinguished by its numerous,
very short, and crowded cells, which. are separated
by thin partitions.

DISTRIBUTION: Europe.

6. Phragmidium violaceum Wint.

Puccinia violacea Schultz, Flor. Starg. p. 459.

Leeythea ruborum Lév.; Cooke, Micr. Fung.
p. 221 p.p.

Phragmidium violaceum Wint. Pilze, p. 231.
Plowr. Ured. p. 223. Fischer, Ured.

Fig. 223.

TUL

Ph. fusi-

Teleutospore.

Schweiz, p. 416, f. 289. Sacc. Syll. vii. 744. Sydow, Monogr.

iii. 139.

P. bulbosum Schlecht. ; Cooke, Handb. p. 491; Micr. Fung. p. 201,

f. 41, 45, 46.

Spermogones. Epiphyllous, in crowded clusters.

4Agcidiospores. Czomata hypophyllous, roundish or elon-
gated, often in circular clusters, on conspicuous spots which are
reddish above and surrounded by a violet-red margin, frequently
also on the stems; paraphyses few, clavate, straight; spores
roundish or ellipsoid, echinulate, orange-yellow, 19—30 x 17—

24: pe.

296 PHRAGMIDIUM

Uredospores. Sori yellow, roundish, often confluent, pul-
verulent; spores ellipsoid to ovate, distantly verruculose, yellow,
28-—32 x 21—24 yu.

Teleutospores. Sori hypophyllous, large, roundish, thick,
pulvinate, black, on conspicuous purple-bordered spots; spores

b

Fig. 224. Ph. violaceum. a, ecidiospore; b, uredospore; c, outlines of various
mature teleutospores from the same sorus; d, young teleutospore before
the middle coat is fully developed, showing the fusion-nucleus in each cell,
and the gelatinous outer coat; e, mature teleutospore, the pearl-like drops
are caused by the shrinking of the outer coat; ¢ x 200, the rest x 600.

a

cylindrical, of 1—5 (mostly four) cells, rounded at both ends,
with a short yellowish papilla at the apex, hardly constricted,
verrucose, brown, 65—100 x 80—385 «4; two germ-pores to each
cell; pedicels long, colourless, swollen at the base.

On Rubus fruticosus. Very common, especially near the
coast. August—November. (Fig. 224.)

This species is easily distinguished by its large conspicuous red and
purple spots, and on microscopical examination by the predominance of
four-celled teleutospores. These pass the winter on the leaves, which often
remain green on the plant; they germinate with the greatest readiness in
April. It is an interesting fact that on some portions of the coast, such
as in North Wales, this species predominates, but on other portions,
e.g. in parts of Yorkshire, as I was informed by the late Mr. R. H. Philip,
its place is largely taken by P. Rubi.

DisTRisuTIon : Europe, Egypt.

PHRAGMIDIUM 297

7. Phragmidium Rubi Wint.

Puccinia mucronata var. Rubi Pers. Disp. Meth. p. 38.
Phragmidium Rubi Wint. Pilze, p. 230. Plowr. Ured. p. 224. Fischer,

Ured. Schweiz, p. 418, f. 290.
Monogr. iii. 141.

Sace. Syll. vii. 745 p.p. Sydow,

Spermogones. Epiphyllous, in minute clusters.
Acidiospores. Caomata hypophyllous, roundish, or on the

nerves elongated, often in little
groups,surrounded byclavate para-
physes; spores resembling those
of P. molaceum.

Uredospores. Sori hypophyl-
lous, scattered; spores smaller
than those of P. violaceum.

Teleutospores. Sori small, on
brownish spots, round, scattered,
seldom more than 4mm. diam.,
black ; spores cylindrical, of 4—7
(mostly six) cells, rounded above
and mucronate with a colourless
acute papilla (5—10 long), not
constricted, rounded below, brown,
beset with numerous little warts,
70—115 x 2832, with three
(or two) germ-pores to each cell;
pedicels hyaline, as long as or
longer than the spore, swollen at
the base.

Fig. 225. Ph. Rubi. Teleuto-
spore x 360; uredospore x 600;
a, teleutospore, boiled in lactic
acid for one minute, x 600. On
R. fruticosus.

On Rubus fruticosus, R. caesius, R. saxatilis (?). Rather less
common than P. violaceum. July—September. (Fig. 225.)

Distinguished from P. violaceum by its smaller teleuto-sovi, and by the
predominance of six-celled spores; the spots on the leaves are usually
much less brilliant in colour. The form on Rubus saxatilis may be a
distinct species, P. Rubi-saxatilis Liro, Ured. Fenn. 1908, p. 421; Sydow,

Monogr. iii. 144,

DisTRiBUTION : Europe.

298 PHRAGMIDIUM

8. Phragmidium Rubi-Idzi Karst.

Puccinia Rubi-Idaet DC. Flor. fr. vi. 54.

Lecythea gyrosa Berk. ; Cooke, Micr. Fung. p. 222, pl. 8, f. 162—4.

Phragmidium gracile Cooke, Handb. p. 491; Micr. Fung. p. 201, pl. 3,
f. 42, 48; Grevillea, iii, 171, pl. 45, f. 9.

P. Rubi-Idaei Karst. Myc. Fenn. iv. 52. Plowr. Ured. p. 226. Fischer,
Ured. Schweiz, p. 420, f 291. Sacc. Syll. vii. 748. Sydow,
Monogr. iii. 146.

Spermogones. Epiphyllous, in little groups, yellowish.

ox

Fig. 226. Ph. Rubi-Idaei. a, part of leaf of Raspberry, showing the circular
groups of ceomata, nat. size; b, a single group, magnified, showing the
spermogones in the centre, surrounded by a pulvinate ring of cxomata,
still partly covered by the epidermis; c, scidiospore and d, paraphysis,
x 600; e, teleutospore x 300; f, teleutospore after boiling in lactic acid;
g, Single cell of same, showing the three coats, (1) hyaline and gelatinous,
(2) dark and rigid, (3) the thin endospore; hk, one of the middle coats,
detached by pressure, showing the three bordered germ-pores.

4icidiospores. Czeomata epiphyllous, small, round, yellow,
usually forming a circle round a group of spermogones, sur-
rounded by a dense border of hyaline, clavate, inward-curved,

KUEHNEOLA 299

thin-walled paraphyses; spores ellipsoid or ovate, sparsely
echinulate, yellow, 21—24 x 18 p.

Uredospores. Sori hypophyllous, small, scattered, pale-
orange, surrounded by thin-walled clavate paraphyses; spores
roundish or ellipsoid, sparsely echinulate, yellow, about 21 x
18 »; pores obscure.

Teleutospores. Sori similar, but black; spores cylindrical,
of 6—10 (mostly 7 or 8) cells, slightly tapering at the apex and
papillate, not constricted, rounded below, verrucose, brown,
80—135 x 28—35 «; three germ-pores to each cell; pedicels
very long, thick, colourless, persistent, swollen towards the base.

On Rubus Idaeus, and possibly on other species of the
genus. Not uncommon. June—October. (Fig. 226.)

Lecythea gyrosa of Berkeley is the ceeoma-stage, in which a pulvinate
ring of spores is formed round the little group of spermogones, leaving the
centre, at tirst sight, apparently unoccupied. This form is not confined
to the early part of the season, but may be found as late as August or
September. It is much rarer than the other spore-forms.

Although this Phragmidium is not found in America, there is a very
similar one, P. imitans Arthur, on allied species of Rubus, inhabiting
the United States and Canada, in which the comata are similarly formed.

On the cultivated Raspberry, when this disease is present, it can be
prevented from spreading by spraying with potassium sulphide solution or
dilute Bordeaux mixture. All diseased leaves should be burnt.

DISTRIBUTION : Europe, Siberia, Japan.

KUEHNEOLA Magnus.

Auteecious.

Spermogones subcuticular, somewhat flattened, without
ostiolar filaments. Uredo-sori of two kinds: primary, ie. the
equivalent of the ceomata, often surrounding the spermogones,
without paraphyses; secondary, similar, but scattered, some-
times with paraphyses; uredospores borne singly on pedicels,
with few and inconspicuous equatorial pores. Teleutospores of

o

300 KUEHNEOLA

several cells as in Phragmidium, but the wall is faintly coloured
or colourless, and smooth; pores one in each cell, apical.

This genus is not confined to Rosacee, being recorded on
Malvacee in America, where also both the British species are
found. It is not closely alhed to Phragmidium: the wall of
the teleutospores and the germ-pores are quite different. But
Tam also of the opinion that the two species included here are
not in reality congeneric.

1. Kuehneola albida Magnus.

Uredo Milleri Schrét. Flor. Schles. iii. 375. Plowr. Ured. p- 256.
Chrysomyxa albida Kiihn, Bot. Centralbl. xvi. 154 (1883). See also
: Bull. Soc. Myc. Fr. xvii. 31.
Phragmidium albidum Lagerh. Mitth. Bad. Bot. Ver. 1888, p. 44.
Fischer, Ured. Schweiz, p. 415.
Kuehneola albida Magn. Bot. Centralbl. Ixxiv. 169 (1898).
K. Uredinis Arthur, N. Amer. Fl. vii. 186 (1912).

Spermogones. Epiphyllous, clustered on small reddish spots.

Fig. 227. K. albida. Uredospore and teleutospores.

Uredospores. Sori, primary epiphyllous, yellow, often in
rings surrounding the spermogones, secondary hypophyllous, seat-
tered, occasionally on the calyx and stems, smaller, punctiform

KUEHNEOLA 301

and paler, when old whitish; spores globose, obovate or irregu-
larly polygonal, closely verruculose-echinulate, yellow, about
20—26 w; pores indistinct (? three or four).

Teleutospores. Sori hypophyllous, singly or in little round-
ish groups, but never confluent, }—4 mm., pulvinate, yellowish-
white; spores cylindrical-clavate, flattened and _ irregularly
coronate at the summit, of 2—13 (mostly 5 or 6) trapezoidal
cells which are smooth and colourless, each measuring 17—30
x 15—25 w; the wall of each cell becomes thicker from below
upwards, and the upper edge is irregularly undulated; the
germ-pore is situated in one of the finger-like projections at
the upper edge of each cell; pedicels very short, sometimes
wanting; each cell is really a perfectly distinct spore.

On Rubus fruticosus. Not rare; Taunton, Hereford,
Worcestershire, Woolmer, New Forest, North Wootton, Ayrshire,
Ireland, etc. Teleutospores, September—November. (Fig. 227.)

The life-history of this species is imperfectly known. The uredospores
may precede the teleutospores, but may also be found simultaneously with
them and (presumably the secondary uredospores) even in the same sorus.
The primary uredospores seem to occur in chains, represent zecidiospores,
and probably germinate at once. The teleutospores germinate im situ on
the leaves as early as the beginning of September. Juel, who experi-
mentally demonstrated the connection of the two spore-forms, suggested
that some uredospores survive the winter and germinate in the spring ;
J. Miiller and S. Strelin state the same of the secondary uredospores.

This species is said to be found on Rubus caesius ; it occurs on many
species of Hubus in North America. The name given by Arthur
(K. Uredinis) rests upon a. (probably true) idea that the teleutospores
constitute the fungus named by Link Ozdiwm Uredinis and placed among
the Hyphomycetes : see Sacc. Syll. iv. 16.

DISTRIBUTION: Europe, North America.

2, Kuehneola Tormentill# Arthur.

Phragmidium Tormentillae Fckl. Symb. Myc. p. 46. Plowr. Ured.
p. 222. Fischer, Ured. Schweiz, p. 414, f. 288. Sace. Syll. vii.
“744. Sydow, Monogr. iii. 105.

P. Potentiliae-canadensis Dietel in Hedwig. Beibl. xlii. 179. Sydow,
Monogr. iii. 106, f. 48.

Kuehneola Tormentiliae Arthur, Res. Sci. Congr. Vienn. p. 342 (1904).

&. obtusa Arthur, N. Amer. Fl. vii. 185 (1912).

302 XENODOCHUS

Spermogones. Epiphyllous, in little groups.

Uredospores. Sori of two kinds, primary epiphyllous, sur-
rounding the spermogones, secon-
dary hypophyllous, scattered, small,

(.) round, punctiform, orange, sur-

) rounded byafew clavate paraphyses;

spores spherical to obovate, finely

echinulate, reddish-orange, 20—
C | 23 x 14—20 p.
Teleutospores. Sori hypophyl-
( } lous, similar, but bright-brown;
spores cylindrical, fusiform or cla-
vate, of 2—7 (mostly five) cells,
often curved, thickened at the apex
Fig. 298. K. Tormentillae like many Puceiniae, slightly con-
Teleutospores. stricted, tapering below, smooth,
sienna-brown, 52—140 x 18—24 p;
epispore thin, with one germ-pore in each cell; contents orange;
pedicels varying in length, persistent, not much widened below.

On Potentilla Tormentilla (=P. erecta) and possibly on
other species of the genus. Very rare. September, October.
(Fig. 228.)

This species resembles a Puccinia in some respects, especially in the
thickening of the apex of the teleutospores, and the position of the solitary
germ-pore of each cell; the wall of each cell becomes darker upwards, the
lower cells being nearly colourless, and the uppermost a pale clear brown,
all quite free from any warts or projections. They can germinate in
autumn (September) like those of K. albida. Dietel says that the
uredospores and their mycelium can survive through the winter.

DIsTRIBUTION: Europe, North America.

XENODOCHUS Schlecht.

Auteecious.

Ceomata large, indefinite, without paraphyses or peridium.
Uredospores absent, represented by the secondary smaller
ceomata. Teleuto-sori similar, often on the same mycelium;
teleutospores of very long chains of cells, not verrucose.

XENODOCHUS 303

Arthur records and names another species, X. minor, on
Sanguisorba latifolia, from Alaska.

Xenodochus carbonarius Schlecht.
Yenodochus carbonarius Schlecht. in Linnea, i. 237, pl. 3, f. 3. Cooke,
Handb. p. 489; Micr. Fung. p. 201, pl. 3, f. 29. Plowr. Ured.
p. 227. Sace. Syll. vii. 751.
Phragmidium carbonarium Wint. Pilze, p. 227. Fischer, Ured.
Schweiz, p. 406, f. 284. Sydow, Monogr. iii. 157, f. 67.
Aicidiospores. Czomata hypophyllous, on coloured spots,
elongated, large, erumpent and pul-
verulent on the nerves, petioles and
stems, roundish and scattered on the
leaves, bright orange-red or vermilion;
spores roundish to oblong, verruculose,
orange, 18—24 y, in short chains.
Teleutospores. Sori amphigenous,
often confluent with the c#omata,
roundish, soon naked, thick, large,
pulvinate, black; spores elongated-
cylindrical, often curved, of 4—22
cells, rounded at both ends, strongly
constricted, smooth, dark-brown, 200—
800 x 24-28 «4; each cell with two
opposite germ-pores at the upper
margin, except the uppermost which
has one apical germ-pore; pedicels Fig. 229. X. carbonarius.
very short, persistent. The whole - Bare rihy *<600 pe
chain is surrounded by a distinct sub-
hyaline membrane, which swells up considerably in lactic acid.

On Sanguisorba officinalis. Not uncommon. June—Oc-
tober. (Fig. 229.) ~

The teleutospore-cells of the Burnet Chain Rust mature from above
downwards, the upper ones being darkest and the lower ones often
colourless, There are no uredospores ; perhaps they are represented by
secondary xcidiospores. The distinctions of this species from the other
Phragmidiew are quite sufficient to justify its generic separation.

Winter assigns to the ceeomata clavate paraphyses which I cannot find.

DISTRIBUTION: Europe, Asia.

304 GYMNOSPORANGIUM

GYMNOSPORANGIUM Hedvw. fil.

Hetereecious. All the sori subepidermal.

Spermogones spherical, with ostiolar filaments. Atcidia
more or less elongated or tubular; peridium membranous,
rupturing by lateral slits; spores brown, with numerous evident
germ-pores, Teleuto-sori erumpent, naked, variously shaped,
gelatinous, expanding when moist; spores with long pedicels,
usually two-celled, generally with two pores (1—4) in each cell,
mostly near the septa; the cell-wall of the pedicels becomes
gelatinised. Germination by an ordinary basidium and roundish
basidiospores: these are thrown off with a jerk, at maturity,
like the spores of the Hymenomycetes (Coons, 1912).

There are many species of Gymnosporangium in America
(Kern in N. American Flora gives 32), of which only three
occur in Europe, and one or two in Japan. Several of these
have three, four, or five-celled teleutospores. The ecidia of the
Gymnosporangia are on Rosacee (except one on Hydranges) ;
the teleutospores are all on Cupressinese (Juniperus, Cupressus,
Chamaecyparis, Libocedrus), on which they often form swellings,
Le. galls. But we get also one remarkable exception to this
rule in the autcecious species (the only one known) G. bermu-
dianum, which produces both its ecidia and its teleuto-sori on
Junipers (J. bermudiana and other species).

1. Gymnosporangium clavarieforme DC.

Tremella clavariaeformis Jacq. Collect. ii. 174.

Podisoma Junipert Fr. ; Cooke, Handb. p. 510; Mier. Fung, p. 214.

Roestelia lacerata Tul. ; Cooke, Handb. p. 534; Micr. Fung. p. 193,
pl. 2, £. 226.

R. carpophila Bagnis, Flora, 1xiii. 317.

Gymnosporangium clavariaeforme DC, Flor. fr. ii. 217. Plowr. Ured.
p. 233. Fischer, Ured. Schweiz, p. 383, f. 275. Sacc. Syll. vii.
737 p.p. Kern, in North Amer. Fl. vii. 202. Sydow, Monogr.
iii. 59, f. 29,

Spermogones. Numerous, amphigenous, but chiefly epi-
phyllous, in little clusters on red spots, yellow, then dark-brown.

GYMNOSPORANGIUM 305

Aicidiospores. ABcidia clustered on yellow or orange thick-
ened spots on the leaves, fruits, and stems, cylindrical, up to

Fig. 230. G.clavariaeforme, Aicidia (Reestelia) on leaf, fruit and branch of
Hawthorn (reduced) ; a, peridium x16, The fruit and gall on branch are
shown as they appear when the peridia are old and the mass looks some-
what like a honey-comb.

24 mm. high, fimbriate above, at length lacerate to base, pale-
brown; spores verruculose, brownish, about 28—30 4; pores
8—10, scattered.

Teleutospores. Spores collect-
ed in long, cylindrical, conical,
ribbon-like or tongue-shaped masses
about 1 cm. long; which are at first
firm, then gelatinous, finally horny
when dry, pale-orange, protruding
from fusiform swellings of the
branches; spores fusiform, varied
much in length and breadth, from
thick-walled, 50—60 x 15—21 p to
thin-walled, 100—120 x 10—12 y,
rounded or attenuated above,
hardly constricted, tapering gradu-
ally below, smooth, dark yellowish
or pale-brown ; germ-pores two in Fig. 231. G. clavariaeforme.

7 - : Masses of teleutospores on
each cell; pedicels hyaline, very branch of J. communis (re-

long. duced); teleutospores x 600.

a

G. U. 20

306 GYMNOSPORANGIUM

feidia on Crataegus Oxyacantha, C. monogyna, Pyrus
communis, July—August; teleutospores on Juniperus ecom-
munis, April and May. Rather common. (Figs. 280, 231.)

The teleutospore-mycelium is perennial in the branches of the Juniper ;
the spores are produced in spring and germinate immediately. The
zcidia can be produced on the Hawthorn in two to four weeks or more
after infection. Their spores in turn infect young shoots of Juniper,
in which the mycelium gives rise to teleutuspores in one year (von Tubeuf),
or two years (Plowright), and in each succeeding year so long as the branch
survives. The mycelium in Crataegus is purely local and annual.

The experimental culture of this species has been performed
unnumbered times by many mycologists ; the ecidia have been produced
on several species of Crataegus and possibly on the Apple and the Medlar ;
also (spermogones only in some cases) on Cydonia, Sorbus, and Amelanchier
—and the teleutospores on Juniperus nuna and J. oxycedrus, as well as
on J. communis. This culture is one of the easiest to perform of all that
are known among the Uredinales.

The only means of checking this disease on the pomaceous hosts is by
the removal of the affected Junipers, if they can be found.

Disrrisution: Europe, Algeria, North America.

2. Gymnosporangium confusum Plowr. /

Medium Mespili DC. Flor. Fr. vi. 98.

Roestelia Cydoniae Thiim. in Sacc. Syll. vil. 834.

Gymnosporangium confusum Plowr. Ured. p. 232, pl. 4, f 13, 14.

Fischer, Ured. Schweiz, p. 385, f 276; Zeitschr. f. Pflanzenkr.
1891, i. 193, 260. Sydow, Monogr. iii. 56, f. 27.

Spermogones. Orange-coloured.

Ajcidiospores. AKcidia’on thickened spots, obconical or
cylindrical, opening at the summit which is at length fimbriate,
orange above, often surrounded by a reddish or purple, line;
spores verruculose, pale-brown, 21—24 py.

Teleutospores. Spore-masses at first pulvinate, dark choco-
late-brown, almost black, then irregularly conical, 5—8 mm.
long, chestnut-brown, swelling when moist; spores oval to
fusiform, smooth, of two kinds, some with hyaline spore-walls
and orange contents, tapering above, others with dark-brown
thick walls, rounded above, about 30—50 x 20—25 fs germ-
pores 2—4; pedicels hyaline, rather long.

GYMNOSPORANGIUM 307

Aicidia on leaves, branches and fruits of Crataegus Oxy-

acantha, C. monogyna, Mespilus ger-
manica, Cydoma vulgaris, June—
August; teleutospores on Juniperus
Sabina, April and May; uncommon.
(Fig. 232.)

The course of the life history of this
species is identical with that of the pre-
ceding. It is distinguished by its shorter
and broader teleutospores, which resemble
those of G. Sabinae, but the ecidia are
quite different from those of that species
and resemble those of G. clavariaeforme.,
The ecidia of G. confusum are reported
also doubtfully on Pyrus communis and
Cotoneaster integerrimus. The cells of the
peridium have their side-walls marked
with elongated obliquely placed ridges,
while those of @. clavariaeforme have them
coarsely warted. This description is taken
from Fischer and Plowright, both of whom
give long accounts of its peculiarities. In
all Plowright’s specimens of G. confusum
(produced artificially on Hawthorn, Quince,
Medlar) the peridia were smaller, shorter,

au:

.

we
Fig. 232. G. confusum. u,

cluster of ecidia on Haw-
thorn twig, nat. size; b, two
peridia of same x 10; ¢, clus-
ter of acidia on Quince leaf,
x5; d, two teleutospores on
J. Sabina x 600. All from
cultures by Plowright.

less deeply torn, and rather more inflated than those of G. clavariae-

JSorme, i.e. of the typical £. lacerata.

DistriputTion: Europe, Northern Persia.

3. Gymnosporangium Juniperi Link.

Tremella juniperina L. Sp. Pl. p. 1625.

Roestelia cornuta Tul.; Cooke, Handb. p. 534, f. 218; Micr. Fung.
p. 193, pl. 2, f. 18, 19. :

Gymnosporangium Junipert Link, Obs. i. 7. Cooke, Handb. p. 509,
f. 203 ; Micr. Fung. p. 214. Sydow, Monogr. iii. 27, f. 11.

G. juniperinum Fr. Syst. Myc. iii. 506. Plowr. Ured. p. 235. Fischer,
Ured. Schweiz, p. 391, f. 278. Sauce. Syll. vii. 738 p.p.

G. cornutum Arthur, Mycologia, i. 240. Kern, N. Amer. Fl. vii. 198.

Spermogones. Epiphyllous, subepidermal, in roundish

crowded groups, yellowish, at length black.

308 GYMNOSPORANGIUM

Aeidiospores. Aicidia hypophyllous, in irregular or circular
groups, horn-shaped, conical, curved,
Ve 4mm. wide, 2mm. long, at length
a open and fimbriate above, yellowish-
ie brown, on round spots which are
brownish below, and bright-orange or
x red on the upper side; spores finely
verruculose, brown, 21—28 x 19—24u;

Mig 2c gig ais gern-pres 8—10, sntered
of Mountain Ash x14; Teleutospores. Spore-masses on
ve unopened peridium sung twigs and occasionally on leaves,
more or less globose, 1—3 mm. across,
at first chocolate-brown, then orange, soft, gelatinous; spores
obtusely fusiform, of two kinds, first thick-walled and brown,
second thin-walled and yellowish, 31—52 x 21—30 yw (Dietel),
66—75 x 17—27 w (Plowright); germ-pores one or two in each

cell; pedicels rather long.

Zcidia on Pyrus Aucuparia, July—October ; teleutospores
on Juniperus communis, May and June. Not common; Surrey,
etc. /Ecidia very abundant at Blair Athol, August, 1905
(D. Prain). (Fig. 233.)

The teleutospores differ from those of the other European species in
the possession of a broad colourless papilla over each germ-pore. Their
mycelium causes fusiform swellings of the smaller branches. Brebner
infected a leaf of Mountain Ash from the teleutospores and obtained
spermogones in eleven days.

DistRiBuTION: Europe, North America.

4. Gymnosporangium Sabine Wint.

Tremella Sabinae Dicks. Pl. Crypt. Brit. i. 14.

Podisoma Sabinae Fr.: Cooke, Handb. p. 510; Micr. Fung. p. 214.

Roestelia cancellata Reb. Fl. Neom. p. 350. Cooke, Handb. p. 533 ;
Micr. Fung. p. 193, pl. 2, f. 20, 21.

Gymnosporangium Sabinae Wint. Pilze, p. 232. Plowr. Ured. p. 230,
pl. 4, f. 11, 12. Fischer, Ured. Schweiz, p. 394, f. 279. Sace.
Syll. vii. 739. Sydow, Monogr. iii. 51, f. 26.

Spermogones. Epiphyllous, on large yellow or orange spots,
very crowded, at length black.

GYMNOSPORANGIUM 309

icidiospores. Aicidia hypophyllous, on the same spots, flask-
shaped, 1—2 mm. broad, pale-brown,
split to the base into lacinie which
remain united at the summit, and at
first are joined at intervals by short
transverse bands; spores finely verru-
culose, brown, 28—30 u (average).

Teleutospores. Spore-masses on
the branches, at first pulvinate, dark-
brown, then irregularly conical, 8—
10 mm. high, red-brown, gelatinous;
spores of two kinds, thick-walled and
thin-walled, broadly and obtusely bi-
conical, scarcely constricted, smooth,
brown, 40—50 x 25—30 w; germ-pores
four, two in each cell.

Fig.234. G. Sabinae. Groups

Zicidia on Pyrus communis, July— os ew leet eae

September; teleutospores on Juniperus
Sabina, April and May. Not uncommon. (Fig. 234.)

This is said to occur on other species of Pyrus and Juniperus. The
life-history is similar to that of the other Gymnosporangia. The spermo-
gones are said by Fischer to have occurred on the fruit of the Pear ;
other authors record the ecidia on both the young fruits and the petioles.
The ecidia are easily distinguishable from all the others, the upper part of
the peridium, after dehiscence, looking very like the calyptra of Polytrichum;
but the teleutospores are similar to those of G. confusum, the chief
difference being that the thick-walled spores of the latter are rounded
at the summit, not bluntly conical.

In this, as in all the similar cases, when the ecidium is found on
its host, search should be made in the neighbourhood for the alternate
host ; the Juniper is often found in a neighbouring garden. Since it
is. always the teleutospore-mycelium that is perennial, the only successful
remedy for this plant-disease is to destroy and burn the Juniper, or
at least the affected part ; it is useless to spray the ecidial host.

DISTRIBUTION: Europe.

310 CRONARTIACEE

CRONARTIACE.

Nearly all hetercecious.

Teleutospores one-celled, without pedicels, produced in
chains; the chains remaining united laterally into bundles
which may be either columnar, wart-like or discoid; germinating
when mature by typical basidia. All the sori subepidermal ;
spermogones are known in both genera. '

Teleutospores in long chains, united into pulvinate sori; cell-wall
smooth, colourless. Uredospores catenulate, surrounded by a very
delicate evanescent peridium. AXcidia not known in the British species.

Chrysomyxa.

Teleutospores in long chains, united into columnar sori; wall smooth,
slightly coloured. Uredospores borne singly on pedicels, echinulate, sur-
rounded by a peridium which ruptures at the summit. Acidia erumpent,
inflated, with a membranous peridium which ruptures at the sides;
wcidiospores partly smooth, not uniformly verrucose over the whole
surface, owing to partial fusion of the warts. Cronartium.

CHRYSOMYXA Unger.

Spermogones hemispherical. icidia with a well-developed
peridium; scidiospores with coarsely verrucose membrane,
without germ-pores. Uredospores produced in rows by basi-
petal abstriction, resembling ecidiospores, but without or with
a very delicate peridium. Teleutospores forming velvety
pulvinate sori, in simple or branched chains, one-celled, with
thin colourless membrane, germinating without a resting
period.

In addition to the two species mentioned below, it is stated
in Massee (Plant Diseases, p. 266) that the xcidium, named
Peridermium coruscans (Fr.) and assigned by Tranzschel to a
Chrysomyxa on Ledum, has been seen on Picea Pinsapo in
England, doubtless on newly imported plants. It is common on

CHRYSOMYXA 311

Picea eacelsa in the North of Europe and has its uredo- and
teleutospores on Ledum palustre, on which it produces witches’-
brooms. See Klebahn, Wirts. Rost. p. 391. It is named by
Tranzschel Chrysomyxa Woroniric.

Quite recently also Chrysomyxa Rhododendri has been
detected in Scotland by Mr D. A. Boyd. See Appendix.

]. Chrysomyxa Empetri Schrit.

Uredo Empetri Pers. ; Cooke, Handb. p. 527 ; Micr. Fung. p. 216.

Chrysomyxa Empetri Schrét. Krypt. Flor. Schles. iii. 372. Plowr.
Ured. p. 258. Sacc. Syll. vii. 762. Fischer, Ured. Schweiz,
p. 557.

Melampsoropsis Empetri Arthur, N. Amer. Fl. vii. 118.

Uredospores. Sori hypophyllous, occasionally (according to
Magnus) epiphyllous, small,
roundish or elongated, some-
times arranged in lines parallel
to the midrib, covered by the
raised epidermis, orange; spores
in short chains, ellipsoid or
polygonal, densely verrucose,
26—35 x 18-25 wu, 30—35 x
21—28 w (according to Fischer),
25—30 x 17—25 (Plowright), __ ; ;
"O6—OT 18-98 w (Artliutl; "ett wien, decile me
wall rather thick, colourless; “edo-sori, x10; b, uredospores

x 600.
contents orange.

On Empetrum nigrum. Uncommon; North Wales, ete.
May—October. Often in small quantity, and very inconspicuous
except when fresh. (Fig. 235.)

This fungus is considered to be a Chrysomywxa because the uredospores
are produced in chains. The teleutospores seem doubtfully to have been
observed by Rostrup and Lagerheim, and might possibly be discovered in
this country if looked for. This parasite also occurs on the same host in
the northern half of North America, but there also no teleutospores have
ever been seen.

DistTRIBUTION : Europe, North America:

312 CHRYSOMYXA

2. Chrysomyxa Pyrole Rostr.

icidium Pyrolae DC. Flor. Fr. vi. 99.

Trichobasis Pyrolae Berk.; Cooke, Handb. p. 529; Micr. Fung.
p. 223 p.p.

Chrysomyxa Pyrolae Rosty. Bot, Centr. iii. 126 (1881). Plowr. Ured.
p. 253, Fischer, Ured. Schweiz, p. 429. Sace. Syll. vii. 761.

Melampsoropsis Pyrolae Arthur, N. Amer. FI. vii. 118.

Uredospores. Sori hypophyllous, often covering the whole
surface uniformly, roundish, 3—1 mm.
diam., soon naked, surrounded by the
torn epidermis and a very delicate
evanescent peridium, yellow; spores
in chains, roundish or polygonal,
verrucose, orange, 21—28 x 18—21 pu.

Teleutospores. Sori hypophyllous,
covering the whole leaf-surface uni-
formly, up to $ mm. wide, roundish
or oblong, flat, waxy, yellowish- then
blood-red, when dry brown; spores
ellipsoid, about 8 « wide, in rows as
much as 100—120 wu long.

On Pyrola minor, P. rotundifolia
and its var. P. maritima. Uncom-
mon; Edinburgh, Kew Gardens,

Fig. 236. C.Pyrolae. a, uredo- Lancashire, ete. April— August.
sori, on underside of leaf, .
nat. size; b, chains of young (Fig. 236.)
, showing inter- . . ‘
ee : - oa This parasite may possibly be heterce-

dospore. On P. maritima cious; Fraser (Mycologia, 1911, iii. 67)

Grecides) ex herb. H. J. syooests that Peridermium conorum-Piceae

. is its ecidium ; Rostrup, Arthur, and Kern

had already expressed the same idea. The

teleutospores are rarely formed, and the fungus probably maintains

itself by its uredospores, which can be distinguished from those of

Pucciniasirum Pyrolae by their sori keing scattered (not in groups), and
by the absence of a distinct peridium.

DisTRIBUTION: Europe, North America.

CRONARTIUM 313

As the remarkable suggested ecidium-stage may possibly
be found in this country, it will be convenient to add -here
the description given of it, so as to aid in its identification :

PERIDERMIUM CONORUM-PICE%
(Hcidium conorum-Piceae Reess.)

AKcidia on the outer surface of the scales, large, 4 cm. or more
in diam., oblong or irregular in shape,
one or two (or in any case few) on
each scale, immersed, white; spores
ellipsoidal, orange; 28—35 x 18—
24 4; epispore with numerous, large,
crowded, prismatic warts, each about
3—4 yw wide.

Fig. 237. P. conorum-Piceae.

icidiospore, from scales of

On cones of Picea excelsa. Au- a a as
gust, September. In some years

rather common in the Alps and Jura Mountains. (Fig. 237.)

CRONARTIUM Fries.

Spermogones hemispherical. Atcidia with a broad, inflated,
irregularly torn peridium; ecidiospores with a coarsely verru-
cose membrane, smooth on one side, without germ-pores,
separated by well-marked intercalary cells. Uredo-sori enclosed
in a hemispherical peridium which opens at the summit by a
narrow pore; uredospores produced singly, on pedicels, echinu-
late, without germ-pores. Teleutospores abstricted in long
chains, and remaining united into cylindrical columns which
are horny when dry, germinating as soon as mature. Basidio-
spores round, very minute.

1. Cronartium asclepiadeum Fr.

Cronartium asclepiadeum Fr. Obs. Mye. i. 220 (1815). Fischer, Ured.
Schweiz, p. 481, f. 295. Sacc. Syll. vii. 597.

C. Paeoniae, Cast. Catal. Pl. Marseill. p. 217 (1845). Cooke, Mier.
Fung. p. 215.

314 CRONARTIUM

C. flaccidum Wint. Pilze, i. 236 (1884). Plowr. Ured. p. 254. Sace.
Syll. vii. 598.

Peridermium Cornut Keb. Zeitschr. f. Pllanzenkr. 1892, ii. 269, pl. 5,
f. 2.

Aieidiospores. Aicidia (P. Cornut) erumpent from the bark,
forming large reddish-yellow bladders, generally occupying a
portion of a branch in large numbers; spores ellipsoid, 22—26
x 16—20 4; epispore 3—4 yw thick, verrucose, thinner on part

‘of its surface and there smooth or somewhat reticulate.

Uredospores. Sori small, pustular, surrounded by a peridium
which opens at the summit with a pore; spores ellipsoid or
ovate, sparsely echinulate, 21—24 x 17—21 p.

Fig. 238. C. asclepiadeum. a, Peridermium Cornui on branch of Pine;
b, teleuto-sori on leaf of Peony (reduced); ev, uredospore x 600; d, part
of a column of teleutospores x 300.

Teleutospores. Sori cylindrical, often curved, arranged in
large clusters, over 1 mm. high, brown, at length horny,
compact; spores ellipsoid or cylindrical-oblong, reaching 56 u
long and 14 ~ broad; epispore thin, slightly thicker above.

fKcidia on the branches of Pinus silvestris, May, June;
uredo- and teleutospores on Paeonia officinalis in gardens,
July—October. Very uncommon. (Fig. 288.)

It has been proved by the researches of Cornu, Klebahn, Fischer and

many others, not only that the remarkable ecidia on Pine are genetically
connected with the other spore-forms on Peony, but also that they can

CRONARTIUM 315

produce the same on Vincetoxicum, Nemesia, Cynanchum, and Verbena, ass
well as on many species of Paeonia. There is reason for believing that
the parasitism on Memesia, at least, has arisen at a very recent date. This
species is therefore plurivorous in its teleuto-stage, but not in its ecidial
stage. The mycelium is perennial, according to Fischer, in the pine-
branches ; it produces ecidiospores in May and infects the alternate hosts,
on which uredo- and teleutospores are borne during the summer—the
latter can germinate at once.

2. Cronartium Quercuum Miyabe.

Uredo Quercus Brondeau in Duby, Bot. Gall. ii. 893 (1830). Cooke,
Handb. p. 526; Micr. Fung. p. 216. Plowr. Ured. p. 257.
Fischer, Ured. Schweiz, p. 539.

Peridermium deformans and P. giganteum Tubeuf, Pflanzenkr. p, 429.

Melampsora (2) Quercus Schrét.; see Sacc. Michel. ii. 308 ; Syll. vii.
594.

Cronartium asclepiadeum var. Quercuum Berk. Grevill. iii. 59 (from
the U.S.). Sace. Syll. vii. 597.

C. Quercuum Miyabe, Bot. Mag. Tokyo, xiii. 74 (1899).

C. Quercus Arthur, North Americ. Fl. vil. 122 (1907).

[Zcidiospores. Aicidia caulicolous, forming subglobose
swellings 5—25 cm. across, arranged in
tortuous lines or cerebroid, at first
orange-yellow, bladdery; peridium col-
ourless, circumscissile, soon falling away
in flakes or sheets, about 2 cells thick,
outer surface smooth, inner verrucose; Fig. 239. C. Querewum.

7 ; Uredospores, on Quercus
cells roundish or irregularly compressed, (ex herb. Thos. Brittain).
walls very thick, lumen small; spores
obovate, 25—32 x 17—23 w; wall colourless, uniformly thick
(24381 »), coarsely verrucose, usually with smooth spot at base
and extending up one side, tubercles somewhat deciduous. ]

Uredospores. ‘Sori hypophyllous, thickly scattered, round,
small (4 mm.), hemispherical, dehiscing by an apical pore, at
length surrounded by the torn epidermis, yellow; peridium
delicate or wanting; spores obovate to broadly ellipsoid, orange-
yellow, 15—25 x 10—17 4; wall colourless, 3 w thick, evenly
echinulate with short,strong points.

[Teleutospores. Sori hypopkyllous, columnar, filiform, 2—3
mm, long, 150—175 uw thick; spores fusiform to oblong, 30—40

316 CRONARTIUM

x 15—20,; wall nearly colourless, smooth, 2—3 thick;
basidiospores oval.]

[4écidia on branches of Pinus ;] uredo- and teleutospores on
leaves of Quercus pedunculata. Rare; Hastings, St Leonards,
Shere, Hurstmonceaux, Salisbury. October. (Fig. 289.)

Only the uredogpores have been found in this country, and usually so
also on the continent. The wcidia’ have been found in the United States
on five species of Pinus, as well as on others in Japan; and the other
spore-forms on at least 20 species of Quercus in various localities. The
description (except for the uredospores) is taken from Arthur (.c.). The
uredospores seem to be most common on the leaves of the shoots that
spring up from the stools of felled oaks. They occur also in France and
Switzerland in the same way. Their dimensions are smaller in this
country than those given by Arthur.

Distrisution: Europe, United States, Guatemala, Japan.

3. Cronartium ribicola F. de Waldh.

Cronartium ribicola Fisch. de Waldh. in Rab. Fung. Eur. no. 1595;
Hedwig. xi, 182. Klebahn, Wirtsw. Rostp. p. 382. Sacc. Syll.
vii. 598. Fischer, Ured. Schweiz, p. 433, f. 296. Gard. Chron.
ser, 3, xi. 736; xii. 44, 183, 187, 501; sili. 425; xxiii. 202, 244;
xxvi. 72—38, 94. Arthur, N. Amer. Fl. vii. 122.

Peridermium Strobi Kleb, Abhandl. Nat. Ver. Bremen, x. 153, pl. 1,
f, 5—10, 13, 14: Zeitschr. f. Pflanzenkr. ii. 269, pl. 5, f. 1.

Spermogones. Irregular elevations of the bark, 2—3 mun.
broad, yellow, with a minute opening through which exudes
a sweet fluid in which the spermatia are included; spermatia
ovoid to elliptical, 2—44 p.

Aicidiospores. Aicidia erumpent from the bark in the form
of a bladder, with an inflated peridium, about 1 cm. high,
yellowish-white ; spores roundish or polygonal, coarsely verru-
cose, except on an elongated smooth patch, orange, 22—29 x
18—20 pw; epispore 2—24 » thick, 3—33 y at the smooth spot.

Uredospores. Sori hypophyllous, small, pustular, forming
orbicular groups 1—5 mm. across, surrounded by a delicate
peridium which opens at the summit with a pore; spores
ellipsoid to obovoid, distantly and sharply echinulate, orange,
21—24 x 14—18 uw; epispore colourless, 2—3 w thick.

CRONARTIUM 317

Teleutospores. Columns hypophyllous, cylindrical, curved,
up to 2 mm. long, crowded especially along the veins of the
leaf, sometimes covering the whole leaf, orange to brownish-
yellow ; spores oblong, smooth, reaching 70 u long x 21 wu broad.

Aicidia on stems and branches of Pinus Cembra, P. monti-
cola, P. Strobus, March—June; uredo- and teleutospores on.
Ribes nigrum, R. rubrum, etc., ‘July—October. Uncommon
except when imported; Surrey, King’s Lynn, Exeter, West-
bury, Woburn, Windsor Forest, Perth, etc. (Fig. 240.)

Fig. 240. GC. ribicola. a, spore of Peridermium Strobi; b, the teleutospore-
columns on leaf of Red Currant (reduced); ¢, uredospore; d, top of a
column of teleutospores, x 600.

This dangerous parasite, sometimes called the Weymouth Pine Rust,
is confined in its ecidial stage to the five-leaved Pines ; it is reported on
the continent also on P. excelsa and P. Lambertiana. The ecidiospore is
distinguished from that of Peridermium Pini and P. Cornui by the fact
that a great part of the surface is smooth. The attacked trees are
stunted, the tops show a bushy growth that can easily be recognised ;
the part where the mycelium is growing is swollen. In the Currant, the
attacked leaves become thicker in texture, and different in colour.

It was first discovered, in this country, at King’s Lyun by Dr Plow-
right, who found the Cronartium on leaves of black, white, and red
currant on July 3, 1892. On August 13 he exhibited in London a branch
of P. Strobus affected by the mycelium of the alternate stage; the tree
grew about fifteen yards away. from the currant bushes. On March 19,
1893, he found the Peridermium in full perfection at the same place.

Though the ecidium occurs in Europe chiefly on the Weymouth Pine
(P. Strobus), yet that cannot be its original host, since neither Cronartium
ribicola nor Peridermium Strobi was found in America (the home of
P. Strobus) until the fungus was imported on it from Germany. P. Cembra,

318 COLEOSPORIACE

the Swiss Stone Pine, however, will equally serve as host, as has been
shown by Tranzschel, and it has been found on that tree in Russia and in
Switzerland. One of these countries was probably the original home of
the parasite, from which it is spreading wherever its hosts will grow.

This is one of the species with which attempts have been made to
produce infection by the use of the abundant spermatia (Klebahn,
Wirtsw. Rost. p. 387), but numerous trials on healthy Weymouth Pines
were entirely without result.

DistTRIBUTION: Europe (north of the Alps), Siberia, Japan,
North America.

‘

COLEOSPORIACE.

AKcidium furnished with a peridium. Spermogones sub-
epidermal, flattish, linear, without ostiolar filaments, dehiscing
by a slit. Teleutospores in one (more rarely two) subepidermal
layers, dividing as they mature into four superimposed cells,
each of which germinates by a sterigma bearing one basidio-
spore.

CoLEospoRIE&. Hetercecious.

fcidia (Peridermium) more or less cylindrical, with inflated
peridium, irregularly torn at summit. Uredospores abstricted
in chains. Teleutospores with a strongly thickened gelatinous
wall above. Basidiospores ovate. Coleosporium.

OcHRoPsOREH. Hetercecious.

Aicidia with cup-shaped peridium. Uredospores abstricted singly.
Teleutospores thin-walled. Basidiospores fusiform.

Ochropsora,

ZAGHOUANIEH. Autcecious.

For the present, the abnormal genus Zayhouania may be arranged
as a subfamily of the Coleosporiacew; it is distinguished
especially by the fact that the four-celled basidium is formed
internally, but emerges from the teleutospore before the
formation of the round basidiospores. See p. 331.

Zaghouania,

The internal “basidium” which has been considered as a
character of this family is not confined to it, being found also
in Chrysopsora, which belongs to the Pucciniacex. It is
remarkable that no species of the family has up till now been
discovered in Australia, while only one of the Eurasian species

COLEOSPORIUM 319

is indigenous to North America, This-family retains probably,
in the mode of germination of its teleutospore, a very primitive
character, but has nevertheless undergone a large amount of
recent evolution, and is no doubt worthy of subdivision.

There is a North American species, belonging to the
Coleosporiaceee, which is deserving of great attention. It is
Gallowaya Pini Arthur (formerly Coleosportum Pint Galloway),
which has teleutospores only, and on leaves of Pinus inops,
i.e. on trees of the same order on which Coleosportwm has its
ecidia. Similarly, among the Melampsoracez, there is a like
case in Necwum Farlow Arthur, which has its teleutospores on
Abies canadensis, while various hetercecious Melampsoracer,
with similar teleutospores on other (non-coniferous) plants,
have their ecidia on Conifers. Again in the Cronartiacez,
Chrysomyxa Ledi and C. Rhododendri are hetercecious species
having their ecidia on Picea excels; but there is also
C. Abietis having its teleutospores on the same host (P. excelsa)
and no other spore form. Gymmnosporangiwm bermudianum,
already mentioned (p. 304), furnishes a somewhat similar
instance. The evolutionary significance of these facts has: not
yet been elucidated.

COLEOSPORIUM Lévy.

fcidia with a more or less cylindrical inflated peridium,
which opens by a cleft and becomes irregularly torn; ecidiospores
with colourless membrane, without germ-pores, superficially.
tuberculate, the tubercles somewhat deciduous. Uredospores
not enclosed in a peridium, abstricted in short chains, resembling
the xcidiospores. Teleutospores in flat, waxy, indehiscent sori,
with a colourless gelatinous membrane, which is thin and wavy
at the sides but strongly thickened above, at first filled with
a rich orange-red oily mass; at length each spore divides into
four superimposed cells, which in autumn can germinate in
situ as soon as mature, with a long sterigma.

The species of Coleosporiwm are morphologically very much
alike, and are distinguished chiefly by their hosts. Moreover,

320 COLEOSPORIUM

since the scidia of all of them live on needles of two-leaved
Pines, it is impossible to say where any given ecidium of this
class belongs, without culture-experiments. If a Coleosporium
can be found in the immediate neighbourhood, the xcidium
may be conjecturally assigned to that. But though the ecidia
are very much alike, there are certain differences to be seen,
although nothing is yet known about their constancy as specific
characters. In certain cases nearly all the peridia will be found
on one only of the two leaves of Pine in each fascicle; this is,
according to my experience, almost always true of C. Senecionis.
In others, the peridia occur on both leaves of the fascicle; this
seems to be the case usually with C. Tussilaginis. Some
peridia are very bladdery and inflated, others are more flat
or cylindrical. It may be possible in the future to distinguish
them by these means, especially since the Coleosporia lend
themselves easily to artificial cultures; see under C. Rhin-
anthacearum and C. Tussilaginis.

1. Coleosporium Senecionis Fr.

Oredo farinosa var. Senecionis Pers. Syn. p. 218.
Peridermium Pini Chev.; Cooke, Handb. p. 535 p.p., f. 219; Mier.
Fung. pl. 2, f. 27, 28 (not the description p. 194).

P. acicolum Link ; Cooke, Mier. Fung. p. 194.

Puccinia glomerata (uredospores) Cooke, Handb. p. 500.

Coleosporium Senecionis Fr. Summ. Veg. Scand. p. 512. Cooke, Micr.

Fung. p. 218, pl. 7, f 145, 146. Plowr. Ured. p. 248 p.p. Sace.
Syll. vii. 751. Fischer, Ured. Schweiz, p. 451. Arthur, N. Amer.
Fi. vii. 94.

Spermogones. Amphigenous, scattered, conspicuous.

Micidwospores. Aicidia (Peridermium acicolum) oblong or
shortly cylindrical, fragile, rupturing irregularly, whitish; spores
almost all oblong, seldom roundish, densely verrucose, orange,
25—85 x 15—25 pm.

Uredospores. Sori hypophyllous, roundish or on the stems
elongated, soon naked and pulverulent, orange; spores mostly
oblong, verruculose, 26—31 x 14—17 j; epispore rather thick.

Teleutospores. Sori hypophyllous, forming little red crusts ;
spores prismatic, length up to 100u, breadth 18—244; epispore
at summit up to 22 w thick.

COLEOSPORIUM 321

ficidia on (?one of the two) leaves of Pinus austriaca,
P. silvestris, May, June; uredo- and teleutospores on Senecio
Jacobaea, S. palustris, S. silvaticus, S. viscosus, S. vulgaris, all
through the year; also recorded for S. pulcher, S. sarracenicus,
etc, in Botanic gardens, and on cultivated Cinerarias (ie.
Senecio) at Sydenham; see Journ. Roy. Hort. Soc. 1908, xxxiil.
511. Very common. (Figs. 241, 242.)

Fig. 241. C. Senecionis. a, chain Fig. 242, C. Senecionis. Teleuto-
of uredospores; b, two fascicles spore germinating.
of leaves of P. silvestris, bearing
peridia on one leaf only of each
fascicle (reduced).

This is the species of Coleosporiwm whose life-history has been longest
known; Wolff first experimentally demonstrated it in 1872, and he was
followed by Plowright in 1882, as well as by Cornu, Hartig, Rathay, Von
Thiimen, Rostrup, Klebahn and Fischer. The ecidium had previously
been called Peridermium. Since Senecio vulgaris continues to live through
the winter in our climate, and Magnus and others have found the uredo-
spores throughout the year, the intervention of the alternate host is
not in this case necessary. Yet it is generally quite easy to find the
Peridermium on the needles of P. silvestris if one searches in June any
trees that may be growing in the neighbourhood of Groundsel infected
with the parasite; since the Peridermium is not conspicuous, it may easily
be overlooked unless special search is made. It is probable that there are
several biological races of this fungus, on different species of Senecio, and
it has been proved by Fischer that it cannot be transferred to Cacalia or
Sonchus.

GU. 2]

322 COLEOSPORIUM

Since there are other so-called “species” of Peridermzum on the leaves
of P. silvestris, which are morphologically not distinguishable from that
belonging to this species, it is always advisable, when such a one is found,
to look on the possible hosts in the neighbourhood for the corresponding
Coleosporium. But the Percdermium found on the bark of Scots Pine is
totally distinct, both morphologically and biologically, although Wolff and
Plowright recorded them as identical. Plowright, however, failed to infect
S. vulgaris by spores from “a specimen of dc. Pint on the bark of a
young fir-branch” (i.e. pine-branch)—naturally enough ; and he also puts
on record (/.c. p. 250) his frequent failures to infect the Groundsel with
spores from ecidia (4c. Pind var. acicola) which seemed to him to be like
those with which he succeeded. His consequent suspicion, that ‘there
must be more than one species included under this name,” is now
abundantly confirmed. He was experimenting, in these latter cases, with
ecidia belonging to some of the species of Coleosporium mentioned in the
following pages.

In North America, C. Senecionis has been found on 8S. vulgaris,
apparently in one locality only (Rhode Island), probably introduced from
Europe ; the Peridermium was not observed. In that quarter of the globe
there afte many indigenous species, biologically resembling ours, but
mostly on different hosts, including one on a species of Orchidaceze.

This fungus does not do much harm to the Pine, but in any case the
removal of Senecio from the neighbourhood arrests the disease. The
teleutospores germinate in the autumn in which they are produced.

DIsTRIBUTION : Europe, North America (once).

2. Coleosporium Tussilaginis Tul.

Uredo Tussilaginis Schum. Pl, Sall. ii. 229.

Coleosporium Tussilaginis “ Lév.” in Tul. Mém. Uréd. 1854, p. 136.
Cooke, Handb. p. 520, f. 211; Micr. Fung. p. 217, pl. 8, f. 180—2.
Fischer, Ured. Schweiz, p. 449.

C. Sonchi Plowt. Ured. p. 250 p.p. Sace. Syll. vii. 752 p.p.

aoe Plowrightii Kleb, Zeitschr. f. Pflanzenkr. ii, 258, pl. 5,
. 6

Aicidiospores. Acidia (P. Plowrighti’) like those of the
allied species; spores oval or mostly round, delicately verrucose,
20—380 x 15—24 pw.

Uredospores. Sori hypophyllous, small, scattered or aggre-
gate, orange; spores roundish, very densely verruculose, 23—
28 x 17—21 yw; epispore rather thick.

COLEOSPORIUM 323

Teleutospores. Sori filling large intercellular spaces of the
mesophyll towards the lower sur-
face of the leaf; spores prismatic,
length up to 140y, breadth 18—
28 w; epispore 18—21 yw thick, or
more, at the summit.

Aicidia on (?both) leaves of
Pinus silvestris; uredo- and teleu-
tospores on Tussilago Farfara,
May—November, very common.
(Fig. 248.)

The connection of the spore-forms
on the alternate hosts has been demon- Frig.243. C. Tussilaginis. Mcidia
strated by Plowright, Klebahn, Fischer > eee
and Wagner. Klebahn and Fischer experiment), reduced; uredo-
showed moreover that the parasite can- and young teleutospore x 600.
not be transferred to Petasites or Sonchus.

Plowright produced the ecidia (P. Plowrightiz) on leaves of Scots Pine
from the Coleosporium on Tussilago (experiment 1248), 25th May, 1899.
His specimens show the peridia, in almost every case, on both leaves of
the fascicle. The same is true of the following experiment by Blackman :
In October, 1904, he tied leaves of Coltsfoot bearing germinating teleuto-
spores of C. Tussilaginis on four trees of P. silvestris at Crockham Hall,
Kent. On April 14th, 1905, the beginnings of spermogones and ecidia
were visible in three cases on neighbouring needles, but nowhere else. In
June, 1905, all four showed the Peridermiwm. Specimens in Herb. Kew.

The uredospores are often very densely and rather coarsely verruculose,
but also occasionally smooth either wholly or in parts ; in fact the warts
are deciduous and may disappear entirely.

DISTRIBUTION: Europe.

3. Coleosporium Petasitis Lév.

Coleosporium Petasitis Lév. Ann. Sci. Nat. 1847, p. 373. Cooke,
Handb. p. 521; Micr. Fung. p. 217. Fischer, Ured. Schweiz,
p. 450.

C. Sonchi Plowr. Ured. p. 250 p.p. Sace. Syll. vii. 752 p.p.

Peridermium Boudiert Fischer, Contrib. étude du genre Coleosporium,
in Bull. Soc. Bot. Fr. xli. 171.

Aicidiospores. ABcidia (P. Boudiert) like those of neigh-
bouring species.
21—2

324 COLEOSPORIUM

Uredospores. Sori scattered, orange, at first covered by the
epidermis, soon pulverulent; spores ovate or ellipsoid, densely
and evenly verruculose, 21—34 x 14—2] p.

Teleutospores. Sori forming little red crusts; spores pris-
matic, length up to 100 yw, breadth 18—24 1; epispore up to
14 thick at the summit.

Aicidia on leaves of Pinus silvestris; uredo- and teleutospores
on Petasites officinalis, August—November, not uncommon.

The life-cycle has been demonstrated by Fischer and Wagner for
P. officinalis: the parasite may also extend to other species of Petasites.

DistTRisurion : Europe.

4, Coleosporium Sonchi Lévy.

Uredo Sonchi Pers. Syn. Fung. p. 217.
Coleosporium Sonchi Lév. Ann. Sci. Nat. 1847, p. 373. Cooke, Handb.
p. 521; Micr. Fung. p. 218, pl. 8, f. 178, 179. Plowr. Ured.
p. 250 p.p. Sace, Syll. vii. 752. Fischer, Ured. Schweiz, p. 453.
Peridermium Fischeri Kleb. Zeitschr. f. Pflanzenkr. v. 13,
Spermogones
Aycidiospores
those of neighbouring species; spores ellipsoid or polygonal-
roundish, verrucose, 25—32 x 18—25 wu.

} fKcidia (P. Fischert) and spermogones like

Fig. 244. C. Sonchi. On S. arvensis. Two chains of young uredospores ;
teleutospore, showing fusion-nucleus ; and uredospore.

COLEOSPORIUM 325

Uredospores. Sori small, round or oblong, girt by the
epidermis, in irregular groups, especially along the veins,
yellowish-orange; spores roundish-oval, densely and _ finely
verruculose, 18—25 x 15—21 u.

Teleutospores. Sori forming small, flat, roundish, waxy-red
crusts; spores prismatic, length reaching 100 p, breadth 18—
30 4; epispore at summit reaching 18 « thick.

AKcidia on leaves of Pinus silvestris; uredo- and teleuto-
spores on Sonchus arvensis, S. asper, S. oleraceus, August—
November, not uncommon. (Fig. 244.)

Klebahn, Fischer, and Wagner have demonstrated the life-cycle of this
parasite; the two former have also proved that it cannot be transferred to
Cacalia, Campanuta, Senecio, or Tussilago. Fischer records this species
on Sonchus palustris as well as on the hosts given above.

DISTRIBUTION : Europe.

5. Coleosporium Cacalie Fckl.

Oredo Cacaliae DC. Flor. fr. vi. 65.

Coleosporium Cacaliae Fckl. Symb. Myc. p. 43. Cooke, Micr. Fung.
p. 218. Fischer, Ured. Schweiz, p. 446.

C. Soncht Plowr. Ured. p. 250 p.p.

Peridermium Magnusianum Fisch. Contrib. étude du genre Coleo-
sporium, in Bull. Soc. Bot. Fr, xl, 171.

P. Magnusti Wagner, Zeitschr. f. Pflanzenkr. vi. 9 and viii. 257.

[Spermogones

Aeidiospores

Uredospores. Sori numerous, roundish, at first covered by
the epidermis, at length pulverulent, orange; spores ellipsoid,
verruculose, deep-yellow, 24—35 x 21—24 p.

Teleutospores. Sori hypophyllous, forming flat, red, waxy
crusts; spores prismatic, up to 140 long, 18—25 4 broad,
thickened (up to 28 ) at the summit.

} Resembling those of other Coleosporva.]

[Mcidia on leaves of Pinus montana, perhaps also of P.
silvestris ;] uredo- and teleutospores on Cacalia hastata, C.
suaveolens. The spermogones and ecidia have not been observed
in Britain; the other spore-forms at Bath (Rev. J. E. Vize),

326 COLEOSPORIUM

Batheaston (C. E. Broome), Oxford Botanic Gardens (Herb.
Bloxam). October. This is, of course, an introduced species.

DISTRIBUTION : Europe.

6. Coleosporium Rhinanthacearum Lév.

Coleosporium Rhinanthacearum Lévy. Ann. Sci. Nat, 1847, p. 373.
Cooke, Handb. p. 521; Micr. Fung. p. 218, pl. 8, f 176, 177.
C. Euphrasiae Plowr. Ured. p. 252. Sace. Syll. vii. 754.

This species is now divided into two, but on purely bio-
logical grounds: no morphological distinctions worthy of the
name can be discerned, and since the hosts are all closely allied
it is perhaps best to retain the collective name, at any rate for
a time.

(1) CoLkrosporium EvupHRasiz Wint.

Uredo Huphrasiae Schum. Pl. Sal. ii. 230,

Coleosporium Euphrasiae Winter, Pilze, p.246. Fischer, Ured. Schweiz,
p. 442. °

Peridermium Stahlii, Kleb. Zeitschr. f. Pflanzenkr. ii. 269, pl. 5, f. 5.

Spermogones

Aicidia
spores oval or roundish, 20—30 x 15—24 p.

Uredospores. Sori small, scattered, roundish, flat, yellowish-
red; spores irregularly polygonal, densely verruculose, 20—24
x 14—17 «; epispore thin, colourless.

Teleutospores. Sori small, flat, roundish, red; spores pris-
matic, orange, up to 105m long, 18—24y wide; epispore at
summit about 14 thick.

Like those of the allied species. cidio-

AKcidia on leaves of Pinus silvestris; uredo- and teleutospores
on ELuphrasia officinalis, Burtsia Odontites, Rhinanthus Crista-
galli, July—September, very common.

It is not certain that Bartsia can be infected from Luphrasia or
Rhinanthus; no experiments ou that point are recorded. Klebahn proved
abundantly that the parasite can be transferred from Rhinanthus to
Euphrasia, but not to Senecio, Sonchus, or Tussilago. Wagner (Zeitschr.
f. Pflanzenkr. viii. 261) infected Huphrasia with ecidiospores from Pinus
montana.

COLEOSPORIUM 327

(2) CoLEosporium MeELampyri Karst.

Uredo Melampyri Rebentisch, Flor. Neomarch. p. 355.

Coleosporium Melampyri Karst. Myc. Fenn. iv. 62. Fischer, Ured.
Schweiz, p. 440, f. 269.

Peridermium Soraueri Kleb. Zeitschr. f. PAlanzenkr. iv. 194,

The only apparent differences from C. Huphrasiae are in
the size of the spores: uredospores
24—35 x 21-28 w; teleutospores as
much as 115y long, 21—28, wide;
epispore very thick (up to 28) at the
summit.

AKcidia on leaves of Pinus silvestris ;
uredo- and teleutospores on Melampy-
rum arvense, M. pratense and _ its
var. montanum, July—September, not
uncommon. (Fig. 245.)

Wagner records the ecidium also on Bt
P. montana. Klebahn has demonstrated that
the spores of this species will not infect

Euphrasia, Rhinanthus, or Campanula. Ihave Fig, 245. C. Melampyri.

not seen the thickening on the summit of the a, teleutospore on M.
pratense, Randan Woods;

teleutospores so pronounced in ours as in the i teloureapone on he
continental specimens, possibly because they same, gathered at Bonn,
were not so mature. Germany.

Sincethe uredo-hosts of C. Rhinanthacearum

are all annual and die at the approach of winter, it would seem probable
that fresh infections must occur each year from the ecidium, but as this
is, at any rate, not commonly found, the ecidiospores must be widely
distributed by the wind; it is very possible, however, that the fungus
winters in some other manner as yet unsuspected, or that the ecidia are
more abundant than is thought to be the case. They should be searched
for in Mayand June. There is here great scope for experimental research,
especially since young pot-plants of Pinus can be used for infection.
Klebahn placed such a Pine amongst a clump of Melampyrum, strongly
infested with Col. Melampyr?, and left it from July to September (the pot
sunk in the earth); in the latter month spermogones appeared and the
wecidium (Peridermium Sorauert) in the following spring.

DisTRiBuTION : Europe.

328 COLEOSPORIUM

7. Coleosporium Campanule Lév.

Uredo Campanulae Pers. Syn. p. 217.

Coleosporium Campanulae, Lév. Ann. Sci. Nat. ser. 3, vill, 373.
Cooke, Handb. p. 521; Mier. Fung. p. 218. Plowr. Ured. p. 251.
Sacc, Syll. vii. 753. Fischer, Ured. Schweiz, p. 448. Arthur,
N. Amer. Flor. vii. 88.

Peridermium oblongisporium Fckl. Symb. Myc. p. 42. Rostr. Bot.
Tidsskrift, xix. pp. 40, 41, 49.

P. Rostrupit Fischer in Bull. Soc. Bot. Fr. xli. p. clxxii.

Spermogones. Amphigenous, scattered, conspicuous.

Aicidiospores. Aicidia (Peridermium oblongisporium) like
those of allied species, but spores distinctly oblong, 834—40 x
18—22 (2).

c
Fig. 246. C. Campanulae. a, uredo- Fig. 247. C. Campanulae. Teleuto-
sori on C. rotundifolia, nat. size ; spores, beneath the epidermis,
b, uredospore and young and x 360; and uredospore x 600. On
mature teleutospores, on C. glome- C. rotundifolia.

rata, x 600.

Uredospores. Sori hypophyllous, scattered, often confluent,
covering the whole leaf-surface, also on the stems, roundish or
irregular, covered by the epidermis for some time, then naked,
yellow or yellowish-red, fading to whitish; spores ellipsoid to
oblong, subangular, strongly and rather densely verruculose,
orange, 21—35 x 14—2] p.

Teleutospores. Sori hypophyllous, subepidermal, scattered,
often confluent, forming small, flat, red, then pale-brownish
crusts; spores prismatic, up to 100 long, 21—28, wide;
epispore hyaline, reaching 35, at the summit, but thin
elsewhere.

OCHROPSORA 329

AKcidia on leaves of Pinus silvestris; uredo- and teleutospores
on Campanula glomerata, C. latifolia, C. rotundifolia, C. Tra-
chehium, C. turbinata, end of July—October, not uncommon ;
sometimes abundant on cultivated species of Campanula, in
both uredo- and teleuto-stages. (Figs. 246, 247.)

Some of the forms on the various species are separated by Klebahn as
biologically distinct, but the specialisation is not in any case sharply fixed
(Wirtswechs. Rostp. pp. 365—9). C. Campanulae is also recorded, in
other countries, on many other species of Campanula and on Phyteuma
and Specularia. In North America the xcidia are found on Pinus rigida
but, as in this country, are much less common, or at least less frequently
observed, than the spore-forms on Campanulaceae. On C. rotundifolia,
this species and Puccinia Campanulae may be found in company.

DisTRIBUTION: Europe, North America, China, Japan, East
Indies.

OCHROPSORA Dietel.

Aicidia with cup-shaped peridium. Uredospores solitary,
on pedicels. Teleutospores united loosely into waxy crusts,
club-shaped or cylindrical, not thickened above, dividing as
they mature into four superimposed cells.

This genus is not closely allied to Coleosporiwm. It is,
indeed, doubtful whether the character upon which the Coleo-
sporlacee are united into one group, viz. the internal basidium,
is really an indication of close affinity. There can be little
doubt that Chrysopsora, which also has an internal basidium,
belongs to the Pee and not to the Coleosporiacez.

Ochropsora Sorbi Diet.

Aeidium leucospermum DC. Flor. fr. ii. 239. Cooke, Handb. p. 536;
Micr. Fung. p. 194, pl. 1, f. 4—6. Plowr. Ured. p. 269.
’ Ochropsora Sorbt Dietel, Ber. Deutsch. Bot. Gesell. 1895, xiii. 401.
Fischer, Ured. Schweiz, p. 455.
Spermogones. On the foliage leaves (loosely spread over
the whole wpper surface) and even on the sepals, whitish, then
brownish.

330 OCHROPSORA

Aicidiospores. Aicidia scattered pretty regularly over the
lower surface of the leaves, not very
crowded, shortly cylindrical, white,
with torn revolute margin; spores
irregularly oblong, colourless, thin-
walled, very delicately verruculose,
18—30 x 15—21 p.
[Uredospores. Sori hypophyllous,
small, roundish, scattered, not more
than + mm. diam.; spore-mass grey-
Fig. 248. 0. Sordi. cid- ish or yellowish-white, surrounded
ium leucospermum. a, ecidia

on leaf of 4. nemorosa, nat. by a circle of paraphyses, which form
size; b, the same, x2; c,
ecidiospores, x 600.

a kind of peridium, but their upper
ends, when mature, are free and sub-
clavate; spores subglobose to ovate, pale-brownish, distantly
verrucose, 25—28 x 18—25 uw; epispore 1—14 yu thick, with no
perceptible germ-pores.

Fig. 249. 0. Sorbi. Section of teleuto-sorus, before the division of the spores
into four cells (one spore is shaded) ; u, basidiospores. (After Fischer.)

Teleutospores. Sori hypophyllous, 4—4 mm. diam., at first
covered by the epidermis, pustulate, pale flesh-colour, roundish
or oblong, clustered in groups; spores cylindrical, rounded
above, crowded, grey, granular, subopaque, 70 x 10—18 p, at
length divided into four cells; basidiospores fusiform, 22—
25 x 8 p.]

AKeidia on Anemone nemorosa, April—June, not common,
Oxford, Cambridge, Devon, North Wales, Yorkshire, Scotland,
ete. [Uredo- and teleutospores on Pyrus Aucuparia, August
and September, not yet observed in Britain.] (Figs. 248, 249.)

ZAGHOUANIA 331

The description of the uredo- and teleutospores is after Fischer; the
latter mature in autumn and germinate at once. According to him, the
mycelium of the excidial stage is perennial in the rhizome, but Klebahn
proved (Zeitschr. f. Pflanzenkr. 1907, p. 144) that the teleutospores infect
the growing points of the rhizome in autumn, and produce the ecidia in
the following spring. He could also infect other species of Pyrus (Aria,
torminalts, scandica, Malus) from the Anemone; Fischer did the same for
P. fennica and P. communis. In the Anemone nearly every leaf of the
affected plant will be attacked, as well as the flower-shoots. The leaves
become longer, narrower and of a paler green, and are borne on longer
petioles. They are often divided into more segments than the normal
leaves. Fischer remarks that, when the fungus appears on the sepals, the
cells in the neighbourhood develop chloroplasts.

The discovery of this hetercecism was due to Dietel and was confirmed
by Klebahn ; previously the ecidium has been mistakenly attributed, by
Soppitt, to Lndophyllum as £. leucospermum. The mode of germination
of the spores will, of course, easily distinguish them: about this there
seems to have been some misapprehension—Soppitt (Journ. Bot. 1893,
p. 274) distinctly stated that the spores did not germinate with “ promy-
celial” spores, but in the Trans. Brit. Myc. Soc. i. 84, 98, this is altered
into the statement that the spores ‘germinate as do those of the (other)
Endophylla.”

Puccinia fusca lives upon the same plant (Anemone nemorosa) but
affects its host in a different way (see p. 215). Meidium punctatum,
a stage of Puccinia Pruni-spinosae, is found on A. nemorosa as well as on
garden Anemones (A. coronaria), but differs in the character of the
broader peridium and in having faintly coloured spores. Moreover, its
spermogones are dark-coloured and are found on both sides of the leaf
(see p. 207).

ZAGHOUANIA Patouillard.

Sorl erumpent, subpulverulent. .Aicidia (Peridermium)
with a peridium which is irregularly lacerate above; margin
slightly involute ; spores in short chains, soon seceding. Sper-
mogones flask-shaped, with ostiolar filaments. Uredospores
pedicellate, solitary. Teleutospores one-celled, ovoid, pedicel-
late, with a slightly thickened hyaline and verruculose epispore,
germinating as soon as mature; basidium four-celled, semi-
internal; basidiospores nearly sessile.

The description is founded upon that given by Dumée and
Maire in Bull. Soc. Myce. Fr., 1902. The semi-internal basidium
is characteristic. Dumée and Maire make a separate family,

332 ZAGHOUANIA

Zaghouaniaceee, for its reception and compare it to Septobasi-
dium among the Auricularies; it seems more likely that
Zaghouania will be proved to belong to the Pucciniacee.

Zaghouania Phillyree Pat.

Acidium Phillyreae DC. Flor. fr. vi. 96. Sacc. Syll. vii. 807. Massee,
Journ. Bot. 1908, xlvi. 153. Trans. Brit. Myc. Soe. iii, 123.

4. crassum var. Phillyreae Cooke, Handb. p. 539.

Uredo Phillyreae Cooke, Exsicc. i. 592. Plowright, Ured. p. 258.
Sace. Syll. vii. 856.

Zaghouania Phillyreae Pat. Bull. Soc. Myc. Fr. 1901, xvii. 187.
Dumée et Maire, Bull. Soc. Myc. Fr. 1902, xviii. 23.

Spermogones. Flask-shaped, with well-developed ostiolar
filaments; spermatia ovoid,
hyaline, 4—5 x 2—3 p.

Aieidiospores. Aicidia nu-
merous, densely crowded, semi-
immersed, with a more or less
involute, nearly entire, whitish
margin; spores variable, glo-
bose, ovoid or oblong-pyriform,
alveolate-reticulate, deep or-
ange-yellow, 20—30 x 14—
18 p.

Uredospores. Sori hypo-

Fig. 250. Z. Phillyreae. a, uredo-sori

on leaf of P. media, Chichester, nat. phyllous, usually on. yellowish

size; b, uredospore from the same; spots, solitary or aggregated,

c, teleutospore beginning to germi- b 1 1

nate (after Dumée and Maire). subangular, at ength naked,
orange-yellow ; spores globose

to ovoid, echinulate, orange-yellow, 24—32 x 12—16 uw.

[Teleutospores. Mixed with the uredospores, oblong-ovoid,
45—65 x 15—18 yw, with a rather thick, hyaline, verruculose epi-
spore ; basidiospores sessile, smooth, subglobose, 12—14 wu diam. ]

Aicidia on leaves and young shoots of Phillyrea latifolia,
Pevensey Churchyard, August, 1907 (G. Massee); on Phillyrea
media, near Chichester, ecidia, 1869, uredo, April, 1874
(F. Paxton). The uredospores have been gathered in Italy as
early as February. (Fig. 250.)

ENDOPHYLLACE 333

The fungus in the ecidium-stage forms rounded swollen pustules on
the leaves or extensive patches on the stems. Every shoot of the year is
usually attacked and contorted, and in August is covered and made
conspicuous by a copious development of the orange spores. The teleuto-
spores have not been detected in this country. They are remarkable for
their mode of germination—the basidium is formed internally as in
Coleosporium; this then emerges through a rupture at the base of the
spore and produces its large nearly sessile basidiospores externally, one
from each cell.

DIsTRIBUTION: France, Germany, Italy, Corsica, Tunisia,
Algeria.

ENDOPHYLLACEA

Teleutospores in long chains, surrounded by a peridium,
which is formed like that of a typical ecidium of Puccinia from
the peripheral cell-rows, but is sometimes less strongly devel-
oped; spores separated by intercalary cells, produced from a
fusion-cell, but germinating as soon as mature by a typical
basidium and basidiospores ; germ-pores not perceptible; spore-
wall coloured, verruculose. Spermogones present; both kinds
of sori subepidermal.

ENDOPHYLLUM Léveillé.

Characters of the genus the same as of the family.

1. Endophyllum Euphorbie-silvatice Wint.

Aicidium Euphorbiae-silvaticae DC. Flor. fr. ii. 241.

4, Euphorbiae Pers. ; Cooke, Handb. p. 537; Micr. Fung. p. 195 p.p.

Endophyllum Euphorbiae-silvaticae Winter, Pilze, i. 251. Sacc. Syll.
vii. 767. Fischer, Ured. Schweiz, p. 437, f. 298.

E. Euphorbiae Plowr. Ured. p. 228.

Spermogones. Epiphyllous, or a few amongst the ecidia,
yellowish, then brown.

334 ENDOPHYLLUM

Teleutospores. Sori hypophyllous, or occasionally epiphyllous,
more or less covering the whole surface, crowded, zcidium-like,
sunk in the leaf-tissue which is slightly swollen, surrounded by
a thin peridium in the shape of a shallow cup, with a short,

Fig. 251. EH. Euphorbiae-silvaticae. wu, affected leaf of E. amygdaloides,
nat. size; b, secidio-teleutospore; c, germinating spore; d, another, not
so advanced, x 540; ¢, inner, and f, outer wall of peridium-cell, x 600.

torn, slightly revolute margin; spores in evident chains, bluntly
polygonal or subglobose, densely but finely verruculose, orange,
18—23 w; epispore about 1 thick; basidiospores obovate, yellow.

On Euphorbia amygdaloides (= E£. silvatica). Rather un-
common. April—June. (Fig. 251.)

The mycelium usually perennates in the plant. The affected shoots
are taller than the normal ones, with shorter, wider and paler leaves;
they seldom flower. This species externally closely resembles an acidium,
and can be distinguished from one only by the different mode of germina-
tion of the spores. The ecidia which occur on other species of Zuphorbia
belong to different life-cycles, Uromyces Pis? etc. ;

The peridium of this species is more strongly developed than that of
the following one; its cells are densely warted, and arranged in distinct
rows. A long account of the behaviour of the parasite is given by
Plowright (/.c.). The spores germinate readily in the summer as soon
as produced.

DISTRIBUTION: Europe.

ENDOPHYLLUM
2. Endophyllum Sempervivi De Bary.

Uredo Sempervivi A. et 5. Consp. p. 126.

335

Endophytlum Sempervivi De Bary, Morphol. p. 304 (see. Saccardo).
Cooke, Handb. p. 546; Micr. Fung. p. 200.

Plowr. Ured. p. 229. Sacc. Syll. vii. 767.
Fischer, Ured. Schweiz, p. 436, f. 297.
Hoffmann, Centralbl. f. Bakter. xxxii. 137,

f. 1—14 and pl. i, ii.

Spermogones. Scattered amongst the

ecidia, roundish.

Teleutospores. Sori rather large, amphi-
genous, sunken in the leaf, ecidium-like, sur-

Fig. 252. E. Sem-

rounded by many layers of hyphae and by a _ pervivi. Sori on

peridium, which opens by a pore at the summit

Sempervivum mon-
tanum (reduced).

and finally becomes cup-shaped; spores bluntly
polygonal or roundish, yellowish-brown, densely reticulate-ver-
rucose, 2435 x 21—28 »; epispore 3—4 4 thick.

On Sempervivum tectorum; also found
on 8. calcareum, S. globiferum, S. montanum
(Plowright), S. arachnoideum (Fischer) and
others (Saecardo). Not common, Warwick-
shire, Forden, Kew Gardens, etc. April—
August. (Figs, 252, 253.)

It has been proved by De Bary, Hoffinann
and others that the basidiospores produced by the
teleutospores infect the leaves, and from them arises
a mycelium which perennates in the stem. It
produces spermogones and teleutospores in the
following spring. The affected leaves are more
erect than normal ones, twice as long, narrower
and yellowish at the base: infested plants should
be burnt, so that they may not infect others. See
the fuller account given on pp. 53-5.

DISTRIBUTION : Europe.

Fig. 253. E. Semper-
vivi. Aicidio-teleu-
tospore germinating
(after Hoffmann).

336 MELAMPSORACEH

MELAMPSORACE/E

Teleutospores not pedicellate, but seated on a dilated
hyphal cell, produced singly in the tissues of the host or com-
pacted side by side into flat crusts, one-celled or divided
longitudinally into 2—4 cells. Germination by an external
basidium, with minute round basidiospores (about 10 «). Uredo-
spores abstricted singly. Uredo-sori and ecidia with or without
a peridium.

Melampsora.
Melampsoridium.
Melampsorella.
Pucciniustrum.
Thecopsora,
Calyptospora.

{ Hyalopsora.
Hyanopsore&. Teleutospores hyaline; on Ferns. { Milesina.

| Uredinopsis.

MELAMPSOREH, Teleutospores brown or brownish,

on Seed-plants. |

MELAMPSORA Cast.

Hetercecious, or in a few species autcecious. The sori, of all
kinds, may be subcuticular or subepidermal. Teleutospores
one-celled, rarely septate, compacted laterally into flat, irregular,
dark-coloured crusts; wall coloured, smooth. Uredospores not
enclosed in a peridium, abstricted singly, without evident germ-
pores, intermixed with capitate paraphyses. Acidia of the
ceeoma-type, pulvinate, without peridium and generally without
paraphyses. Spermogones, shallow, hemispherical, without
ostiolar filaments.

There are seven species of Melampsora recorded for North
America, but only one of these, A/. Lint, is found in Britain,
and one other, M. alpina on arctic and alpine species of
Saaifraga and Salix, in Europe. The Melampsoras on Salix
and Populus form a very complex group; this is no doubt
correlated with the fact that the genera Salia and (to a
smaller extent) Populus are themselves in a state of flux,
evolving many closely related species and possessing many

MELAMPSORACEA 337

hybrids. Klebahn is the author who has chiefly studied the
group, and the following account of the known or supposed
British species on Salix is mainly founded upon his work.

The artificial key to the species, here given, is based upon
that of Fischer, but must be taken-with a little reservation
since, as is now known, the life-histories of such hetercecious
forms require to be worked out for each country separately.

HETERGCIOUS.

A. Teleutospores on Salix.

a. Uredospores oblong, smooth at upper end. Czeoma on Allium.
1. Teleutospores subepidermal. M. Allit-Salicis-albae.
2. Teleutospores subcuticular. M. Allii-fragilis.

b. Uredospores roundish, without smooth spot.
1. Teleutospores subcuticular, strongly thickened above, with a
conspicuous germ-pore. Coma on Larix.
M. Larici-Caprearum.
2. Teleutospores not strongly thickened above, without an evident

germ-pore.
(1) Teleutospores subepidermal.

Ceoma on Larix. M. Larici-epitea.

» on Orchidacee. M. Orchidi-repentis.

» on Huonymus. M. Euonymi-Caprearum.

» on R&ebes. M. Ribesti-purpureae.

(2) Teleutospores subcuticular. M. Ribesti-viminalis.

B. Teleutospores on Populus.
a. Uredospores elongated, smooth above. Teleutospores on P.
nigra, P. balsamifera.
1. Cexoma on Larix. Teleutospores epiphyllous.
M. Lariei-populina.
2. Coma on Allium. Teleutospores hypophyllous.
, M. Allit-populina.
b. Uredospores roundish, not smooth above. Teleutospores on
P. alba, P. tremula.

1. Ceoma on Larix. M. Lariet-trenvutae.

2. » on Pinus. AM. pinitorqua.

3. » on Mercurialis. M. Rostrupit.
AUTCECIOUS.

Teleutospores not on Salicacee.
M. Hypericorum, M. Lint, M. Euphorbiae, M. vernalis.

G. Ue 22

338 MELAMPSORA

1. Melampsora Larici-Caprearum Kleb.

Caeoma Laricis Plowr. Ured. p. 262 p.p.

Melampsora salicina Lév. ; Cooke, Handb. p. 522 ; Micr. Fung, p. 219.

MM. farinosa Schrot. Flor. Schles. iv. 360. Plowr. Ured. p. 238 p.p.
(see note).

M. Larici-Caprearum Keb. in Forstl.-naturw. Zeitschr. 1897, p. 469.
Fischer, Ured. Schweiz, p. 483, f. 312.

Acidiospores. Ceomata minute, pale-orange; spores round-
ish, oblong, or polygonal, 15—25 x 12—17 w; epispore up to
2 thick, finely verruculose, with many thin places (? germ-

pores).
ASS
(ii

:

Fig. 254. M. Larici-Caprearum. Para- Fig. 255. M. Larici-Caprearum.

physis and uredospores (one showing Teleuto-sori, on upper face of leaf
the thin places in the epispore); b, of S. Caprea, nat. size; teleuto-
teleutospores. On S. Caprea. spore, germinating, x 360.

Uredospores. Sori hypophyllous, showing as yellow spots
on the upper side, variable in size and arrangement, 1—3 mm.
wide; spores roundish, oval, or polygonal, 14—21 x 18—15n;
epispore 2—24y thick, firm, distantly echinulate, with thin
places (? germ-pores); paraphyses capitate, 50—60 x 18—26 pn,
thickened (up to 5 w) above.

Teleutospores. Sori epiphyllous, 1 mm. or more wide, dark
reddish-brown, frequently confluent in extensive crusts, lying
between the cuticle and the epidermis; spores prismatic,
rounded below, 830—45 x 7—14y, rather unequal in length;
epispore clear-brown, thin (1), but thickened (up to 10)

ON WILLOW 339

above where it is pierced by an evident laterally placed germ-
pore.

Ceomata on Larix europaea; uredo- and teleutospores on
Salia Caprea, more rarely on S. awrita and other Salices. The
commonest species. (Figs. 254, 255.)

The teleutospores germinate the following spring. They are distin-
guished from those of all the allied species by being thickened above.
Plowright remarks that the czeoma is not uncommon early in the year on
Larch foliage, but is very inconspicuous and easily overlooked ; he found
the zcidiospores, in company with the uredospores on 8. Caprea, at West
Malvern, June, 1900. But it is almost impossible to say, without experi-
ment, to which form of Melampsora any given coma on Larch is to be
assigned. .

Plowright’s If. farinosa seems to be chiefly this species, but several of
the allied forms are continually recorded under the same name. The
reddish teleuto-sori on the upper side of the leaf, are distinctive and are
easily found by looking for them from September onwards. It may be
mentioned here that both uredo- and teleutospores of the Melampsoras on
Willow and Poplar germinate readily : if the germinating uredospores are
placed upon healthy leaves and kept in a damp chamber, infection usually
follows in 7—10 days.

2. Melampsora Euonymi-Caprearum Kleb.

Uredo confluens var. Euonymi Mart. Flor. Mosq. p. 280. Cooke,
Handb. p. 527.

U. Euonymi Cooke, Micr. Fung. p. 216.

Cacoma Euonymi Plowr. Ured. p. 260.

Melampsora Euonymi-Caprearum Kleb. in Pringsh. Jahrb. f. Wissensch,
Bot. 1900, xxxiv. 358. Fischer, Ured. Schweiz, p. 489.

Spermogones. Flatly pulvinate.

4cidiospores. Cxeomata mostly hypophyllous, bright-orange,
in elongated clusters on orange spots, }—1 mm. diam.; spores
oval, rarely oblong, 18-23 x 1419 4; epispore thick (up to
5 w), finely and densely verrucose.

Uredospores. Sori hypophyllous, on discoloured spots which
show distinctly on the upper side, small, } mm. wide, pulvinate,
single or in groups; spores mostly roundish, orange, 14—19 x
14—17 p, distantly echinulate without smooth spots ; epispore
thin, or at times thickened (up to 4), with several germ- .

22—2

340 MELAMPSORA

pores; paraphyses capitate with a slender pedicel, thickened
(up to 8 yz) above, 50—70 x 18—25 pu.
Teleutospores. Sori hypophyllous,
covered by the epidermis, small, about
4 mm. diam., but united mto groups
bounded by the veins, brown with a
tinge of bluish-grey; spots brown on the
upper surface; spores irregularly pris-
matic, rounded at both ends, 25—40
x 7—13 4; epispore thin, clear-brown,
scarcely thickened above, with a barely
perceptible apical germ-pore.

Fig. 256. M. Euonymi- - :
Cuprearum, OA dead Ceomata on HLuonymus europaeus,

leaf of S. Caprea,show- August, September, rather rare; uredo-

ing teleuto-sorion lower . zn

face (slightly reduced). and teleutospores on ; Saliz aurita, 8.
Caprea, S. cinerea. (Fig. 256.)

Fischer records the coma in Switzerland in May and June. The
distinction of this species from the preceding one (apart from the ecidial
host) lies in the teleuto-sori ; these are hypophyllous and subepidermal,
while those of Jf, Larici-Caprearum are epiphyllous and subcuticular. It
is not certain, however, that this distinction is absolute; Fischer in a
culture on S. cinerea obtained a few teleuto-sori showing also the latter
characters. There is a further difference in the form of the teleutospores.

3. Melampsora Larici-epitea Kleb. (emend. Fischer).

Uredo epitea K. et 8. Mycol. Heft. i. 68.

Lecythea epitea Lév. ; Cooke, Micr. Fung. p. 221.

Melampsora epitea Thiim. ; Plowr. Ured. p. 239.

M. Larici-epitea Fischer, Ured. Schweiz, p. 485, f. 313. Kleb. in
Zeitschr. f. Pflanzenkr. 1899, ix. 88.

Aicidiospores. Ceeomata hypophyllous, scattered or in rows,
with yellow spots on the upper surface, roundish or oblong,.
3—14 mm. long, pale orange-yellow ; spores roundish or some-
what polygonal, finely warted, 15—25 x 10—21,; epispore
14—3 w thick, with no recognisable germ-pores.

Uredospores. Sori amphigenous, seated on yellow spots,
orange-yellow, t—14 mm. diam.; spores mostly oval, sometimes
oblong, roundish, or angular, echinulate, crange, 12—25 x 9—

ON WILLOW 341

19 w; epispore 14—3} w thick, without perceptible germ-pores ;
paraphyses capitate, with a thin pedicel, occasionally clavate,
thickened (up to 10 yz) above, hyaline, 35—80 x 15—24 u.

Fig. 257. M. Larici-epitea. Fig. 258. Mf. Iarici-epitea.
Cxeoma on Larch leaf, x2 Uredospores and _ para-
(one of Plowright’s cultures physes, on S. viminalis.

from S. cinerea).

Teleutospores. Sori amphigenous, covered by the epidermis,
at length dark-brown, sometimes tinged round with greyish-
blue or violet, }—1 mm. diam., densely clustered or confluent ;
spores prismatic or subclavate, rounded at both ends, occasion-
ally narrowed above, 20—50 x 7—14 ; epispore clear-brown,
uniformly thin, without an evident germ-pore.

Cxeomata on Larix europaea; uredo- and teleutospores on
many species of Saha The teleutospores in October and
November. (Figs. 257, 258.)

This species is one of the most complex of all the Melampsoracez ; it
has been divided into about eight biological races, all of which have their
ceoma on Larch, while the other spore-forms are specialised to certain
species of Willow. It is recorded on the following British Salices for
other countries, but it is not possible as yet to say on which of these it is
found in this country: S. aurita, Caprea, cinerea, fragilis, Smithiana,
viminalis for the typical form ; nigricans for the form M. Lariei-nigricantis
Schneider ; purpurea for the form M. Larici-purpureae Schneider ; and
reticulata for M. Larici-reticulatae Schneider. Morphological differences
between these forms are not discoverable.

The teleutospores germinate in the spring following after their forma-
tion. The teleuto-sori are formed under the epidermis, and appear to
start usually just below a stoma; they often occupy entire areas bounded
by the veins. This description is taken mainly from Fischer. The
teleutospores are distinguished from those of M. Larici-Caprearum by the
total want of thickening at the apex. Plowright produced the ceoma on
Larch from the teleutospores on Salix cinerea, May 24th, 1900.

342 MELAMPSORA

4, Melampsora Ribesii-viminalis Keb.

Caeoma Ribestt Link, Sp. Pl. ii. 28 p.p.
Melampsora Ribesii-viminalis Kleb. in Pringsh. Jahrb. f, Wissensca.
Bot. 1900, xxxiv. 363. Fischer, Ured. Schweiz, p. 494.

Spermogones. Pulvinate, with flat hymenium.

Aicidiospores. Czeomata hypophyllous, on discoloured spots
which show on both sides, arranged in groups, 1°5 mm. wide,
bright-orange ; spores roundish, rarely oval, 18—23 x 14—17 p;
epispore 2—4 4 thick, with many thinner places (? germ-pores),
finely and densely verruculose.

Uredospores. Sori hypophyllous, minute, about +mm.,
scattered or in groups, pale orange-yellow ; spores more or less
round, 15—19 x14—16 yp; epispore 2 w thick, uniformly echinu-
late; paraphyses capitate or more often clavate, hardly thick-
ened above, 50—70 x 18—25 p.

Teleutospores. Sori epiphyllous, developed between the
cuticle and the epidermis, }—4 mm., scattered or in groups
over the whole leaf, dark-brown, shining; spores prismatic,
rounded at both ends, more or less irregular, 25—40 x 7—14p;
epispore thin, clear-brown, not thickened above, with no evident
germ-pore,

Ceomata on Ribes Grossularia, R. mgrum, R. rubrum;
uredo- and teleutospores on Salix viminalis.

Klebahn showed, by many trials, the genetic connection of the parasites
on these species. The teleutospores can be distinguished from all others
except those of M. Larict-Caprearum and M. Allii-Galanthi-fragilis by
their being subcuticular. M. Larici-epitea is also recorded for Salix
viminalis, but that has its teleutospores subepidermal, and the paraphyses
of the uredo-sori strongly thickened at the apex. The ceoma on Ribes
does not seem to be recorded for this country and is apparently rare.

5. Melampsora Ribesii-purpuree Kleb.

Melampsora Ribesii-purpureae Keb. in Pringsh. Jahrb. f. Wissensch.
Bot. 1901, xxxv. 667, Fischer, Ured. Schweiz, p. 492.
Spermogones, Subconical, with flat hymenium.
Aicidiospores. Cxeomata mostly hypophyllous, on pale-yellow
spots which show on both sides, scattered or in subcircinate
groups, 4—14 mm., sometimes confluent, surrounded by the

ON WILLOW 343

torn epidermis, orange; spores roundish or subpolygonal, 18—
20x 15—18; epispore about 3, thick, finely and densely
verruculose, with a few thinner places (? germ-pores).

Uredospores. Sori mostly hypophyllous, on conspicuous
bright-yellow spots which show on both sides, 4—14 mm.,
pulvinate, surrounded by the torn epidermis, bright orange-red ;
spores roundish, uniformly but not densely echinulate, 15—23
x14—19.; epispore up to 24 thick, with a few thinner
places; paraphyses variable, capitate or clavate, 40—70 x 12—
21 w, not thickened above.

Teleutospores. Sori amphigenous, but mostly hypophyllous,
scattered or in little groups over the whole surface, subepider-
mal, small, }—4 mm. diam., brownish-black; spores irregular,
prismatic, rounded at both ends, 25—35 x 7—10p; epispore
thin, clear-brown, not thickened above, without evident germ-
pore.

Ceomata on Ribes alpinum, R. Grossularia (not on R.
nigrum, R. rubrum); uredo- and teleutospores on Salia pur-
purea.

Description according to Fischer ; Klebahn’s experiments have demon-
strated the connection of the parasites on these hosts. It is probable that
MM. Ribesit-auritae Kleb. is only a biological race of the same fungus, having
spermogones and czeoma-spores (according to Klebahn) on Lubes nigrum
as well as on the two mentioned above, and its other spore-forms on
Salix aurita and possibly 8. Caprea. They are scarcely distinguishable,
if at all, morphologically ; both may be British.

M. Larici-epitea is also recorded on S. purpurea, but can be distin-
guished by its densely clustered teleuto-sori on spots often bounded by
veins, and the paraphyses of the uredo-sori strongly thickened above,
M. mixta (Plowr. Ured. p. 289) may belong to either, but his species ex-
tended to the branches and inflorescence.

6. Melampsora Orchidi-repentis Kleb.

Uredo confluens var. Orchidis A. et 8. Consp. p. 122, Cooke, Handb.
p. 527.

U. Orchidis Mart. Flor. Mosq. p. 229. Cooke, Micr. Fung. p. 216.

Caeoma Orchidis Wint. Pilze, p. 256. Plowr. Ured. p. 261; Gard.
Chron. 1890, viii. 41. Sacc. Syll. vii. 868.

Melampsora Orchidi-repentis Kleb. in Pringsh. Jahrb. f. Wissensch.
Bot. 1900, xxxiv. 369. Fischer, Ured. Schweiz, p. 488.

344 MELAMPSORA

Spermogones. Hardly projecting, flat, mostly under the
stomata.

Aicidiospores. Ceeomata irregular in outline, clustered or
circinate on large pale-yellowish spots, often
confluent, 1—2 mm. diam.; bright orange-
yellow; spores roundish-polygonal, 11—20 x
11—15 yw; epispore very thin, delicately
warted.

Uredospores. Sori hypophyllous, minute,
deep-orange, causing yellow spots on the
upper side; spores more or less round, orange,
echinulate, 183—17 x 12—14; paraphyses
Fig.259. M.Orehiai- mostly capitate, with thin pedicels, 40—

reece, Gems 70 x16—20,

ovata, from one of Teleutospores. Sori hypophyllous or a

aegis ae w- few on the upper side, subepidermal, small,
dark-brown; spores prismatic, rounded at
both ends, 16—48 x‘7—14y; epispore clear-brown, uniformly
thin (about 1»), without any evident germ-pore.

Ceomata on Lnstera ovata, Orchis latifolia, O. maculata,
May, June; uredo- and teleutospores on Sahix aurita, 8. repens.

Not common. (Fig. 259.)

The ceoma is also recorded on Listera cordata, Goodyera repens, and
Gymnadenia conopsea, but no experimental proof of the connection seems
to be forthcoming in the case of these species, though both Plowright and
Klebahn have demonstrated it for the other species. The teleutospores
germinate, as usual, after the winter’s rest. Soppitt found the coma on
O. maculata and the teleutospores on S. repens growing together near
Southport.—The ecidium of Puccinia Orchidearum-Phalaridis can be
readily distinguished from the ‘ceoma of this species by its evident
peridium and the regularity of its form.

7. Melampsora Allii-fragilis Kleb.

Uredo Alltiorum DC. ; Cooke, Micr. Fung. p. 217 p.p.

Cacoma Alliorum Link ; Plowr. Ured. p. 261 p.p..

Melampsora Vitellinae Plowr. /.c. p. 240 p.p.

M. AUii-fragitis Web. in Pringsh. Jahrb. f. Wissensch. Bot. 1901,
xxxv. 671, Fischer, Ured. Schweiz, p. 481.

ON WILLOW 345

Spermogones. Scarcely projecting, with flat hymenium.

4icidiospores. Ceomata on the leaves and stems, also on
the bulbils, clustered on discoloured spots, usually oblong, up
to 2mm. long, surrounded by the torn epidermis, bright orange-
yellow; spores irregular, mostly polygonal, seldom round, 18—
25 x 12—19 w; epispore 1—1$ w thick, finely verruculose.

Uredospores. Sori hypophyllous or partly epiphyllous,
minute, $mm., circular, surrounded by the torn epidermis,
reddish-orange, causing reddish-yellow spots on the upper side ;
spores distinctly oblong or obovate, 22—83 x 13—15y; epispore
up to 3 thick, with thinner places (? germ-pores), distantly
echinulate, but smooth and somewhat thinner above; paraphyses
capitate or clavate, 50—70 x 15—20 yy, with thin pedicel and
uniformly thickened membrane (3—5 yp).

Teleutospores. Sori chiefly epiphyllous, between the cuticle
and the epidermis, scattered or in groups, pulvinate, 1—14 mm.
broad, dark-brown, shining; spores -prismatic, rounded at both
ends, 30—48 x 7-144; epispore clear-brown, uniformly thick
(about 1), without evident germ-pore; basidiospores orange.

Ceomata on Allium Cepa, A. ursinum and others; uredo-
and teleutospores on Salia fragilis, S. pentandra, and the
hybrid between them.

The teleutospores germinate after a winter’s rest. Klebahn’s J.
Galanthi-fragilis is morphologically identical, even occurring on the same
species of Salix, but has its ceomata on Galanthus nivalis ; it can only be
considered as a biological race. The same author’s M. Larici-pentandrae
infests the same species of Salix, but has its ceomata on Larix ; it is
distinguished, however, by its teleuto-sori, which occur on both sides of
the leaves, are more minute (though often confluent) and arise below the
epidermis. I have a Melampsora on Salix fragilis, which has minute
teleuto-sori abundantly on the upper surface and beneath the epidermis.
This might belong to M. Larici-pentandrae Kleb. ;

8. Melampsora Allii-Salicis-albe Kleb.

Melampsora Altii-Salicis-albae Kleb. in Zeitschr. f, Pflanzenkr. 1902,
xii. 19. Fischer, Ured. Schweiz, p. 480.

Spermogones. Rather flat.
Acidiospores. Ceomata on the leaves and stems, in groups

346 MELAMPSORA

on yellowish spots, about 1 mm. diam., surrounded by the
epidermis, bright orange-yellow ; spores mostly roundish, 17—
26 x 15—18 yw, densely verruculose.

Uredospores. Sori of two kinds: (1) in summer and autumn
on the leaves, hypophyllous, 4 mm. wide, on inconspicuous
discoloured spots, (2) in spring, erumpent from the bark of
young twigs and as much as 5 mm. long, afterwards on the
young leaves, as much as 2 mm. long and densely crowded ;
spores all similar, distinctly oblong, sometimes clavate or pyri-
form, 20—36 x 11—17 4; epispore 2m thick, smooth above,
distantly echinulate below ; paraphyses capitate, 50—70 x 15—
20 », not thickened above, absent from the cortical sori.

Leleutospores. Sori amphigenous, subepidermal, scattered
thinly over the leaf-surface singly or in groups, dark-brown ;
spores prismatic, rounded at both ends, clear-brown, 25—45 x
7—10 4; epispore scarcely 1 thick, not thickened above,
without evident germ-pore.

Keidia on Allium ursinwm and other species; uredo- and
teleutospores on Salix alba.

The ceoma on Allium is indistinguishable from that of MM. Alli-
fragilis or M. Altii-populina. This species can winter by its teleutospores
which produce the ceoma on Alliwm, or by the perennial mycelium in the
cortex of the branches on which the uredospores appear in spring before
the cxzoma is produced; these sori are without paraphyses. The whole
of this account is due to Klebahn: I have a specimen from Yorkshire
(C. Crossland) which is referred, doubtfully, to this species.

9. Melampsora arctica Rostr.

MM. arctica Rostr, Fung. Groenland. 1888, p. 535. Sacc, Syll. Fung.
vii. 595. Annals of Scott. Nat. Hist. 1911, p. 37.

Uredospores. Sori hypophyllous, gregarious, yellow ; spores
spheroid to ovoid, echinulate, 18—23 «; paraphyses clavate.

Teleutospores. Sori hypophyllous, scattered, very small,
dark-brown; spores prismatic, reddish-brown.

On leaves of Salta herbacea, Scotland, Ben-an-Dothaidh, at
3100 ft. (J. A. Wheldon and A. Wilson),

ON POPLAR 347

Plants of the Salix were brought from Scotland and cultivated at
Walton, near Liverpool; there were then uredospores only. When the
leaves fell, they were left on the ground, and two teleuto-sori were developed
on them, of a rufous black colour. I have not seen the specimens.

10. Melampsora populina Lévy.

Melampsora populina Lévy. Ann. Sci. Nat. 1847, p. 375. Cooke,
Handb. p. 523; Micr. Fung. p. 219. Plowr. Ured. p. 242. Sace.
Syll. vii. 590.

(1) MELAmpsorA ALLII-POPULINA Kleb,

Caeoma Alliorum Link ; Plowr. Ured. p. 261 p.p.

M. populina Cooke, Micr. Fung. pl. 9, f. 195, 196.

Melampsora Allii-populina Keb. in Zeitschr. f. Pflanzenkr. 1902, xii.

25. Fischer, Ured. Schweiz, p. 504.

Spermogones. Pulvinate, projecting.

Aicidiospores. Ceomata about 1 mm. wide, mostly in
groups on yellowish-white spots on the leaves, surrounded by
the epidermis, bright orange-red; spores roundish or oval or
polygonal, 17—23 x 1419»; epispore about 2, thick, but
sometimes thicker and then obviously thin at certain spots
(?germ-pores), verruculose.

Uredospores. Sori hypophyllous, or even rarely epiphyllous,
round, scarcely 1 mm. wide, surrounded by the epidermis,
causing yellowish spots, bright reddish-orange; spores distinctly

YL

Fig. 260. i. Allii-populina, a, teleutospores ; b, paraphysis ; c, uredospores.
On Populus nigra.

oblong or clavate, rarely oval, 24—38 x 11—18,; epispore
2—4 thick, with sunken places (? germ-pores), sometimes
thicker at one end, but without any equatorial thickening,
distantly echinulate, but smooth above; paraphyses mostly
capitate, 50—60 x 14—22 », with a wall uniformly 2—3 4
thick.

348 MELAMPSORA

Teleutospores. Sori hypophyllous, subepidermal, scattered
over the leaf singly and in groups, pulvinate, +—? mm,
blackish-brown, not shining; spores prismatic, rounded at both
ends, 35—60 x 6—10 w; epispore clear-brown, 1—1} pu thick,
scarcely thickened above, without evident germ-pore.

Ceomata on Allium Cepa, A. ursinum and other species,
May; uredo- and teleutospores on Populus nigra, P. balsama-
fera. (Fig. 260.)

The description is chiefly after Klebahn. The teleutospores mature in
February of the following year; I find them shorter than given above,
viz. 25—32 p, and somewhat truncate at the summit. The czoma of

species of Melampsora on Allium is easily distinguished from the xcidium
of Puccinia Winteriana on the same host by the absence of a peridium.

(2) Metampsora Larici-poputina Kleb.

Caeoma Laricis Plowr. Ured. p. 262 p.p.

Melampsora populina Lév.; Plowr. Ured. p. 242 p.p. (see note,
“ Biology,” p. 243).

M. Larici-populina Kleb. in Zeitschr. f. Pflanzenkr. 1902, xii. 43.
Fischer, Ured. Schweiz, p. 502, f. 316.

4Aicidiospores. Ceeomata on hardly perceptible spots, scarcely
1 mm. long, bright yellowish- orange; spores oval or roundish,

17—22 x 14—18 uw; epispore 14—2 4 thick, colourless, finely
verruculose.

ete xO eee
i

Fig. 261. Mf. Larici-populina. a, teleutospores; b, uredospores and
paraphysis, On P. ontariensis,

Uredospores. Sori mostly hypophyllous, in little groups,
causing yellowish angular spots on the upper side, rarely

ON POPLAR 349

solitary and epiphyllous, up to 1 mm. wide, at first covered by
the raised epidermis and a layer of hyphe, at length uncovered
and surrounded by the same, distributed over the whole leaf-
surface ; spores distinctly oblong, 30—40 x 13—17 pw; epispore
about 2 « thick, at the equator thickened up to 5—6 yu, covered
with rather distant spines except at the summit which is
smooth; paraphyses clavate-capitate, 40—70 x 14—18 wv, strongly
thickened (up to 10 u) above.

Teleutospores. Sori epiphyllous, minute but united in
groups and confluent, distributed over nearly the whole leaf,
covered by the epidermis, clear translucent-brown, then red-
brown, and at length black; spores prismatic, rounded above
and less so below, 40—50 or more x 7—10y; epispore pale,
scarcely 1 thick, but reaching 24—3y above where it is
faintly coloured and without evident germ-pore.

Cxomata on Laria europaea; uredo- and teleutospores on
Populus balsamifera (ontariensis), P. canadensis, P. nigra, P.
pyramidalis (atalica), and also on other species, in September
and October. Not uncommon. (Fig. 261.)

The description of the czeomata is after Klebahn. This species is
distinguished from M. Allii-populina by the following points: the elon-
gated uredospores have a distinct equatorial thickening or rather swelling ;
the teleuto-sori are epiphyllous, and the spores are very slightly thickened

at the summit. On P. ontariensis I have seen the teleuto-sori so thick
and crisp as to remind one of Phyllachora Ulmi on elm-leaves.

11. Melampsora Tremule Tul.

Melampsora Tremulae Tul. Ann. Sci. Nat. 1854, p. 95. Cooke, Handb.

p. 522; Micr. Fung. p. 219. Plowr. Ured. p. 240. Sacc. Syll.
vii. 589.

(1) Me tampesora LaRIcI-TREMULAE Kleb.

Caeoma Laricis Plowr. Ured. p. 262 p.p.
Melampsora Larici-tremulae Kleb. in Forstlich.-naturwiss. Zeitschr.
1897, p. 468. Fischer, Ured. Schweiz, p. 498, f. 315.
4cidiospores. Ceomata solitary or in little groups, on
yellowish spots, minute, scarcely 1 mm. diam., pale-orange or
flesh-colour ; spores roundish, oval or polygonal, 14—17 x 12—
16 «4; epispore about 1 » thick, verruculose,

350 MELAMPSORA

Uredospores. Sori hypophyllous, minute, 4 mm., somewhat
pulvinate, pulverulent, spots obsolete; spores oval, oblong or
obovate, rarely round, 15—22 x 10—15y; epispore about 2
thick, covered with stout spines; paraphyses distributed through-
out the sorus, elongated-clavate rather than capitate, 40—45 x
8—17 p, with a thick wall (3—9 ,).

Teleutospores. Sori hypophyllous, minute, $—1 mm. diam.,
covered by the epidermis, dark-brown; spores prismatic, rounded
at both ends, 40—60 x 7—12,y; epispore thin (1—2), not
thickened above, with a scarcely evident germ-pore.

Ceomata on Laria europaea; uredo- and teleutospores on
Populus alba, P. tremula, rarely P. balsamifera, September,
October.

The description is after Klebahn. The teleutospores germinate after
a winter’s rest. Though Plowright records that he could obtain no result
“by placing the germinating teleutospores of M. tremulae on young larch
trees” (2c. p. 241), the truth of the hetercecism has been abundantly
demonstrated by Hartig, Klebahn and Fischer. Plowright mentions that’
Prof. Trail had found these two forms growing in company near Aberdeen ;
and the meaning of his failure is simply that he was using one of the
other Melampsorae that grow on P. tremula. It has been suggested, but
not proved, that Jf pinitorqua is itself only a biological race, and there
are also M@. Magnusiana (Klebahnz), and M. Rostrupit which may come
under the same category; see p. 353.

(2) Metampsora PINITORQUA Rostr.

Cacoma pinitorquum A. Braun, in Monatsb. Berl. Akad. 1863, p. 624.
Melampsora pinitorgua Rostr. Tidsskr. f. Skovbrug, 1889, xii. 177.
Fischer, Ured. Schweiz, p. 499. Yorkshire Fungus Flora, p. 183.
Massee, Text-book of Fungi, f. 22, 98; Text-book of Plant
Diseases, p, 235, f. 60; Dis. Cultiv. Plants, p. 325, f. 98.
Aicidiospores. Czomata erumpent through the cortex of
young shoots, solitary, linear, reaching 20 x 3mm., reddish-
orange ; spores roundish or oval, pale reddish-yellow, 14—20 x
13—17 u, or oblong, 22 x 104; epispore about 2 thick, deli-
cately verruculose.
Uredospores. Sori hypophyllous, on spots which show yellow
chiefly on the upper side, solitary or in groups, often scattered
over the whole surface, scarcely $ mm. wide, pulvinate; spores

ON POPLAR 351

oval to ovate-oblong, 15—22 x 11—16 u; epispore uniformly
about 2 thick, with two germ-pores(?), echinulate all over;
paraphyses distributed throughout the sorus, clavate, not capi-
tate, 40 —50 x 12—17 y, with a uniformly thick wall (3—7 w).

Fig. 262. M. pinitorqua (from 4 German specimen, ex herb. Sydow). a, a
young shoot of Pine, in June, with newly-formed leaves, showing three
ceomata (@), shaded; the leaves have been removed from the affected
portion, which is beginning to be curved; b, a ceoma, x10; ¢, @ecidio-
spores; d, old leaf of Aspen, showing numerous teleuto-sori on the lower
surface ; e, teleutospores.

Teleutospores. ‘Sori hypophyllous, mostly in clusters, about
4mm. wide, covered by the epidermis, crust-like, brown, not
shining; spores prismatic, rounded at both ends, 20—35 x 7—
11 »; epispore pale-brownish, scarcely 1 » thick, not thickened
above, without evident germ-pore.

Ceomata on young shoots of Pinus silvestris which they
distort, May, June; uredo- and teleutospores on Populus alba,
P. tremula, and their hybrid (P. canescens), not on P. balsami-
fera. Rare in this country. (Fig. 262.)

The connection of the forms on the two hosts was first demonstrated
by Rostrup and confirmed by Hartig and Klebahn. See p. 57.

(3) MeELampsora Rostrupi Wagner.

Uredo confluens DC.; Cooke, Handb. p. 527; Micr. Fung. p. 216,
pl. 7, £. 133, 134.

Caeoma Mercurialis Link ; Plowr. Ured. p. 260. Sacc. Syll. vii. 868.

Melampsora aecidioides Plowr. Ured. p. 241. Sace. Syll. vii. 590.

M. Rostrupii Wagner, in Oester. Bot. Zeitschr. 1896, xlvi. 273
Fischer, Ured. Schweiz, p. 501.

352 MELAMPSORA

Spermogones. Epiphyllous, or a few hypophyllous, in large
round clusters, honey-coloured.

Aicidivspores. Ceeomata hypophyllous and on the petioles
and stems, in clusters on pale-yellow spots, often circinate
round the spermogones, about 1 mm. wide, often confluent in
patches 1—14 cm. wide, bright-orange; spores roundish-poly-
gonal or oval, 13—18x12—16y; epispore 1—l}¥ thick,
finely and densely verruculose.

(>)

WA

Fig. 263. M. Rostrupii. u, teleutospores on P. tremula; b, teleutospore on
P, alba; c, wcidiospores on Mercurialis perennis; d, paraphyses and
uredospores on P. tremula (one paraphysis contains the orange remains of
the protoplasm).

Uredospores. Sori hypophyllous, about 1 mm. wide, pulvi-
nate, compact, causing large yellow spots on both sides; spores
oval or sometimes roundish-polygonal, 21—25 x 14—18 1;
epispore up to 34 thick, covered with stout rather distant
spines; paraphyses distributed throughout the sorus, clavate or
somewhat capitate, 50 x 15—23 yw, with a thick wall (3—6 p).

Teleutospores. Sori hypophyllous, $—1 mm. wide, scattered
over the whole leaf, covered by the epidermis, dark-brown ;
spores prismatic, rounded at both ends, pale-brown, 40—55 x
8—10,; epispore thin (1—2 ), not thickened above, without
any evident germ-pore.

Ceomata on Mercurialis perennis, April—June ; uredo- and
teleutospores on Populus alba, P. tremula, and occasionally on
other species, September and October. (Fig. 263.)

The connection of the two forms has been shown by Rostrup, Nielsen,
Plowright, Klebahn, Wagner, and Jacky, The teleutospores may be found

MELAMPSORA 353

by looking in spring on fallen leaves of P. tremula, P. alba, at places where
Mercurialis is found to be affected. The ceeoma on the latter is very
capricious in its occurrence ; in some years it may be found almost every-
where, in other years hardly a specimen can be met with. The large
yellow spots show conspicuously on the upper surface of the leaves, so
that when it does occur it is impossible to overlook it.

There are two places near Birmingham where all the spore-forms occur
year after year in close proximity: it is from these specimens that the
descriptions are taken. In one case the teleutospores are on P. alba, in
the other on P. tremula.

The morphological differences of the four (or five) species of Melam-
psora which appear on P. ulba and P. tremula are very slight. It is better
to include them all for the present under Melampsora tremulae, as
biological races. The fourth species is J/. Magnusiana Wagner, which
has its ceomata on Chelidonium, and which, according to Klebahn, is
identical with WM. Alebahni Bubdk, on Corydalis ; this has not yet been
recorded for Britain, but Plowright mentions (Jc. p. 241) a Melampsora
on P. tremula from which he could not obtain ceomata on either Larix or
Pinus or Mercurialis—this might be M. Magnusiana.

12. Melampsora Euphorbie Cast.

Uredo Helioscopiue Pers. Disp. Meth. p. 13.

Leeythea Euphorbiae Lév. ; Cooke, Micr. Fung. p. 221.

Melampsora Euphorbiae Cast. Obs. ii..18, with plate (1843). Cooke,
Handb. p. 523; Micr. Fung. p. 219, pl. 9, f. 193, 194.

M. Helioscopiae Wint. Pilze, p. 240 (1884). Plowr. Ured. p. 236.
Sace, Syll. vii. 586. Fischer, Ured. Schweiz, p. 508, f. 318,

[Spermogones. Flat, hemispherical.
Aicidiospores. Cxomata minute, }—} mm. diam. on the
leaves, 1—4 mm. long on the stems, yellowish-red; spores in

Fig. 264. M. Euphorbiae. a, teleutospores on E, exigua; b, teleutospore,
and c, uredospore and paraphysis, on EL. Peplus.

GU. 23

354 MELAMPSORA

short chains, without paraphyses, roundish to ellipsoid, densely
verrucose, 21—28 x 19—24 y.]

Uredospores. Sori hypophyllous and on the stems, scattered
roundish or oblong, soon naked, but surrounded by the epider-
mis, golden-brown, mixed with numerous capitate paraphyses ;
spores roundish, orange, 15—17 x 11—14 y; epispore colourless,
echinulate, without perceptible germ-pores; paraphyses hyaline,
with strongly thickened heads, 16—23 w diam.

Leleutospores. Sori amphigenous, subepidermal, minute,
roundish or oblong, reddish-brown, then black, more or less
pulvinate, on the stems often confluent; spores prismatic, 50—
60 x 10—14 wz; epispore brown, thin, at length thickened above
(up to 5 w) and darker.

On Euphorbia exgua, E. Helioscopia, FE. Peplus. May—
October. Very common. (Fig. 264.)

Miiller, on the ground of cultures, considers the form on £. Helioscopia
as a separate species from those on Z. Peplus and E. exigua, the latter
two also being biologically distinct from each other (Centralbl. f. Bakter.
2, xix. 441). In his experiments he observed a czeoma-stage on £. exigua
which the others did not have. This stage seems to be very rarely met
with, and has probably not occurred in this country: the description of
the spermogones and czeomata is from Fischer. In the same species, as it
occurs on £. Cyparissias, Jacky states that he obtained the uredospores
by infection with the basidiospores without the intervention of the
ceoma-stage, so that M. Huphorbiae may be in a transition state between
a eu-form and a hemi-form. The ecidium on £. exigua recorded by
Plowright (Zc. p. 270) belongs to Uromyces tuberculatus ; sce p. 102.

DistRIBUTION: Europe, Siberia.

13. Melampsora Hypericorum Wint.

Uredo Hypericorum DC. Flor. fr. vi, 81. Cooke, Handb. p. 526;
Micr. Fung. p. 215, pl. 8, f. 174, 175.

Melampsora Hypericorum Winter, Pilze, p. 241, Plowr. Ured. p. 243.
Sace, Syll. vii. 591. Fischer, Ured. Schweiz, p. 506, f. 317.
McAlpine, Rusts of Australia, p. 191.

«Ecidiospores. Czomata hypophyllous, scattered, roundish
or oblong, flatly pulvinate, often very small, subepidermal, long
covered, at length erumpent, orange, showing as indefinite pale-
yellow or orange spots on the upper face; spores in short

MELAMPSORA 355

chains, ellipsoid to polygonal or subclavate, 18—28 x 10—18 w;
epispore colourless, about 2 yu thick, —

rather densely verruculose, with no aye
perceptible germ-pores; no para-

physes.

Teleutospores. Sori hypophyllous, i
subepidermal, small, roundish, red- ACRND a b
dish-brown, then dark-brown ; spores
prismatic, more or less rounded above, Fig. 265. M. Hypericorum.

i a, teleutospores, under the
pale-brown, 28—40 x 10—17 p; epi- epidermis; b, sxcidiospore,

. without paraphyses. On
spore thickened (up to 3 ») above. Pain agen

On Hypericum Androsaemum, H. humifusum, H. perfora-
tum, H. pulchrum. Not common. May—October. (Fig. 265.)

What was described by Plowright as the uredo-stage of this fungus is
stated by Fischer, Tranzschel, and others, to be the coma stage—the
spores “being produced in short chains, with sterile intercalary cells,
without paraphyses, but sometimes” (at least on Hypericum montanum)
“surrounded by a layer of swollen colourless cells which might almost be
considered as an undeveloped peridium.” Miiller considers the form on
Hf. montanum as a biological race, since he could not infect other species
of Hypericum with spores from it.

But Klebahn has proved (Zeitschr. f. Pflanzenkr. 1905, xv. 106) that a
species of Hypericum can bear both the caeoma-form without paraphyses
and the uredo-form with paraphyses. McAlpine (Rusts, p. 192) récords
that in Australia the uredo-sori have very abundant paraphyses, inter-
mixed; he describes them as “hyaline, capitate, overtopping the spores,
50—68 x 18—24 p.” His species was on leaves and stems of H. japonicum,
and differs slightly from the British ones. His description of the uredo-
sori is as follows: “Sori mostly hypophyllous, scattered or gregarious, at
first bright-orange, becoming pale, pulverulent, up to 4mm. diam.,
erumpent and surrounded by the ruptured epidermis. Spores subglobose
to ellipsoid, finely verrucose, orange-yellow, 14—21x 11—17 p, with two
germ-pores on one face.” On the British specimens which I have seen,
there are no paraphyses and the spores are decidedly in chains.

DistRIBUTION: Europe, Siberia, India.

14. Melampsora Lini Desm.

Uredo miniata var. Lini Pers, Syn. Fung. p. 216.
U. Taini Schum. Pl. Sail. it. 230. Arthur, N. Amer. Fl. vii. 101.

23—2

356 MELAMPSORA

Lecythea Lini Berk. ; Cooke, Handb. p. 582 ; Micr. Fung. p. 222,
pl 8, £ 165—%,

Melampsora Lini Desi. Pl. Crypt. no. 2049. Plowr. Ured. p. 237.
Sace. Syll. vii. 588. Fischer, Ured. Schweiz, p. 507. McAlpine,
Rusts of Australia, p. 192, f. 236 and pl. I, f. 36.

Spermogones

Aicidiospores

Uredospores. Sori amphigenous and on the stems, small,
scattered, roundish or oblong, flatly pulvinate, subepidermal,
(2at first covered by a parenchymatous peridium, Fischer),
orange; spores roundish to ellipsoid, echinulate, orange-yellow,
16—24 x 12—17 yu; paraphyses not numerous, hyaline, strongly
capitate, much thickened above, 20—25 pw diam.

} See below.

Fig. 266. M. Lini. a, teleutospores; 6, plan of same; ve, paraphysis
and two uredospores. On L. catharticum.

Teleutospores. Sori similar, but confluent, chiefly on the
stems, subepidermal, reddish-brown, at length black and shining;
spores prismatic, 35—60 x 7—10 yp; epispore thin, yellowish-
brown, rather thickened above and darker.

On Linum catharticum. June—October. Not uncommon.
(Fig. 266.)

Forms of Melampsora Lint occur widely on many species of Linwm and
_have usually been regarded as identical. That which often acts as a very
destructive parasite wherever the common Flax is cultivated has cousider-
ably wider teleutospores (17—20,p, McAlpine) and attempts to infect
L. usitatissemum from L. catharticum have uniformly failed ; it is therefore
considered by some as a biological race or even species=.JJelampsora
liniperda Kornicke (Centralbl. f. Bakter. 1911, 2. xxxii. 278). Teleuto-
spores of this have been described as much as 80 long. Fromme (1912)
has recently described spermogones and ecidia to this form on cultivated

MELAMPSORA 3857

Flax ; the spermogones are flask-shaped, subepidermal, without ostiolar
filaments, the ecidia are small and apparently difficult to distinguish.
No other author seems to have met with these, unless the structures to
which Fischer assigns a “ parenchymatous peridium” were such ecidia.

It would keep the disease in check if infected Flax plants were pulled
up and burnt as soon as seen, but such a remedy is impracticable on a
large scale. No really immune varieties of Flax are:known, but fortunately
the parasite seems not to occur in the Irish flax-fields.

DISTRIBUTION: Europe, North and South America, Australia.

15. Melampsora vernalis Niessl.

Uredo Saxifragarum DC. Flor. fr. vi. 87. Cooke, Handb. p. 525;
Micr. Fung. p. 215.

Caeoma Saxifragae Wint. Pilze, p. 258. Plowr. Ured. p. 259.

Melampsora vernalis Niessl, in Wint. Pilze, p. 237 (1881). Plowr.
Gard. Chron. 1890, viii. 41; Jour. Roy. Hort. Soc. xii. p. cxi;
Trans. Brit. Myc. Soc. i. 59. Sacc. Syll. vii. 592.

M. Saxifragarum Schrot. in Cohn’s Krypt. Flor. Schles, p. 375 (1887).
Fischer, Ured. Schweiz, p. 511.

Spermogones. Scattered, yellow.

4icidiospores. Cxomata small, elliptic, flat, solitary, golden-
yellow; spores in chains, roundish, finely verruculose, 17—30 u.

[Uredospores. Sori epiphyllous, very small, round; spores
ellipsoid, echinulate, golden-yellow, 16—20 x 15 y.]

Teleutospores. Sori densely clustered, subepidermal, small,
irregular, chestnut-brown ; spores oblong to clavate, yellowish-
brown, 40—50 x 14 w.

On Sazifraga granulata. Rare. June—September.

Plowright states that, on the specimens collected by Mr James Taylor
at Clark Farquhar, N.B., in June, 1890, the teleutospores were found on
the lower leaves and stems, and there were no uredospores. The descrip-
tion of the latter is after Voglino and Fischer, and may not belong to the
British species. The connection of the coma with the teleutospores has
been proved by Plowright and Dietel.

DisTRiBUTION: Germany, Switzerland, Italy.

358 MELAMPSORIDIUM

MELAMPSORIDIUM Klebahn.

aie 2

Hetercecious. id

Teleutospores one-celled, with brownish membrane, united
into flat waxy crusts, but each little group starts almost always
directly beneath a stoma. Uredo-sori surrounded by a hemi-
spherical peridium which opens by an apical pore, often begin-
ning beneath a stoma; uredospores abstricted singly, more or
less smooth at one end, with indistinct germ-pores, not mixed
with capitate paraphyses. Atcidia with a well-developed inflated
peridium. Spermogones subcuticular, other sori subepidermal.

This genus is, in some respects, as closely allied to Puccint-
astrum as to Melampsora, or rather more to the former than to
the latter. .

Melampsoridium betulinum Kleb.

Melampsora betulina Desm. ;XCooke, Handb. p. 522, f. 212 (misnamed
winter spores); Micr.”Fung. p. 219, pl. 9, f. 189, 190. ‘Plowr.
Ured. p. 248. Mace. Syll. vii. 592.

Melampsoridium betulinum Wleb. Zeitschr. f. Pflanzenkr. 1899, p. 21
(see also 1891, p. 1380); Wirtswechs. Rostp. p. 401. VPischer,”
Ured. Schweiz, p. 512, f. 320.

Uredo populina var. betulina P. ys p. 219.
fe

Acidiospores. Aicidia hypophyllous, single or in longitu-

Fig. 267. M. betulinum. w, peridium of uredo-sorus, viewed from above,
showing the point at which it will dehisce, x 300; b, uredospores, x 600.

dinal rows parallel to the mid-rib, small, oval or oblong, 1—14
mm. long, clear reddish-orange, with irregularly torn margin;

MELAMPSORIDIUM 359

spores roundish, 14—21 x 11—16 y, echinulate; epispore thinner
and smoother above.

Uredospores. Sori hypophyllous, with yellow spots showing
on the upper side, collected in groups and mostly limited by
the veins, each sorus scarcely ;4, mm. wide, surrounded by a
dome-shaped peridium which at length opens at the summit
(where its cells are drawn out into long sharp points, Fischer) ;
spores decidedly oblong or subclavate, orange, 22—40 x 8—12 uw;
epispore colourless, with distant spines, often smooth above.

Fig. 268. 2. betulinum. Sorus of teleutospores. The fusion-nucleus
is seen in four of them. x 600.

Teleutospores. Sori hypophyllous, always covered by the
epidermis, scarcely 4 mm. wide, in dense clusters limited by
the veins, often spread over the whole leaf, reddish, then brown ;
spores prismatic, rounded at both ends, somewhat oblique,
80—50 x 7—15 4; epispore thin, scarcely thickened above,
nearly colourless, without perceptible germ-pore.

AKcidia on leaves of Laria europaea, May ; uredo- and teleu-
tospores on Betula alba (both verrucosa and pubescens), August
—November, lasting through the winter on the decaying leaves.
(Figs. 267, 268; see also Fig. 37, p. 78.)

It was Plowright who first, in 1890 (after many unsuccessful trials)
discovered that the Melampsora on Birch was connected with an zcidium-
form on Larch (see Zeitschr. f. Pflanzenkr. i. 130; Gard. Chron. 1890, viii.
41). He performed the experiment in both directions, and his conclusions
were confirmed, eight years later, by Klebahn. The ecidium in this cage
does not belong to the czoma-type, but to that of Peridermium (P. Laricis
Kleb.), having a peridium and resembling in its spores also Peridermium
Strobi. The forms on B. verrucosa and B. pubescens are, to a small extent,

360 MELAMPSORELLA

biologically distinct. The ecidium stage of this parasite is very rare
everywhere, and seems not to be recorded in Britain, except by Plowright
at King’s Lynn; the other stages are exceedingly common all over the
country, but do little damage. The teleutospores germinate after a
winter’s rest, but since it is supposed they cannot infect the Birch and
the ecidium-stage appears to be so rare, it is doubtful by what means the
fungus propagates itself from year to year. Liro states that in Northern
Europe the ecidium on the Larch does not occur, though the fungus is
very common in the uredo-stage, especially on Birch seedlings ; but as he
also says that the uredospores do not survive the winter there, the question
of its perennation is left in a very unsatisfactory state.

On the Hornbeam (Carpinus Betulus) there is in Europe an allied
species (Melampsoridium Carpint) to which an ecidium has not yet been
discovered.

DISTRIBUTION: Europe, Asia, North America.

MELAMPSORELLA Schroter.

Hetercecious.

Teleutospores in the epidermal cells, with a thin, colourless
membrane, usually one-celled, germinating at once on maturity.
Uredo-sori furnished with a peridium, and without any para-
physes amidst the spores; uredospores yellow, sessile, produced
in short chains or singly, without evident germ-pores. icidia
with a peridium. Uredo-sori and ecidia subepidermal : sper-
mogones subcuticular, without ostiolar filaments.

This genus bears a certain resemblance to Milesina. but is
distinguished from it by the absence of vertical septa in the
teleutospores, as well as by its habitat on Phanerogams.

1. Melampsorella Caryophyllacearum Schrot.

Beidium elatinum A. et 8. Consp. p. 121. Plowr. Ured. p. 270.

Peridermium elatinum Link; Cooke, Handb. p. 535; Micr. Fung.
p. 194.

Uredo Caryophyllacearum Johnst.; Cooke, Handb. p. 526; Mier.
Fung. p. 216. 5

Melampsorella Curyophyllacearum Schrét. in Hedwig. 1874, xiii. 85.
Klebahn, Wirtswechs. Rostp. p. 396. Fischer, Ured. Schweiz,
p. 516, f. 522—6,

MELAMPSORELLA 361

Jelampsora Cerastit Wint. Pilze, p. 242 (1881). Plowr. Ured. p. 247.

Melumpsorella Cerastii Schrot. Flor. Schles, p. 366 (1887). Sace. Syll
vii. 596.

M. elatina Arthur, N. Amer. FI. vii. 111.

Spermogones. Epiphyllous, scattered, conical, honey-coloured.
Aicidiospores. Aicidia hypophyllous, arranged in an irregu-
lar row on each side of the mid-rib, erumpent,
shortly cylindrical, roundish or compressed,
pale orange-red, with torn white margin;
spores ellipsoid or polygonal, orange, 16—

30x 14—17 4; epispore thin, densely verru- i
cose. aa

Uredospores. Sori generally hypophyl- y
lous, usually arising beneath a stoma, sur-

rounded by a peridium which slowly opens

by an apical pore, small, crowded, pustular, Beene a
yellow; spores sometimes in short chains, — dermium elatinum,
ovoid-oblong or ellipsoid, yellowish, 20—30 alighely Leen
x 16—-21 4; epispore thin, beset with a 4, aleaf, x10.
small number of pointed warts which are

only visible when dry, with an occasional glabrous strip (?),
without perceptible germ-pores. .

Teleutuspores. Hypophyllous, often covering the whole leaf,
developed within the epidermal cells, whitish-yellow or pinkish,
in little groups in each cell, roundish or flattened, one-celled,
1421 yw; epispore smooth, thin; basidiospores globose, nearly

colourless, 7—9 p.

Aividia on leaves of Abies pectinata, June—September ;
uredo- and teleutospores on Cerastiwm arvense, C. triviale and
its var. alpestre, C. viscosum, Stellaria graminea, S. media
(more rarely); uredospores from May onwards. Not very
common. (Figs. 269, 270.)

In North America it occurs on other species of Adzes, and on Alsine
and other species of Cerastium ; also in Europe on numerous allied species
of the subfamily Alsinee. The teleutospores are developed on those
leaves of the second host which live through the winter ; they germinate
about May and can infect the Silver Fir. There are comparatively few
records of the ecidium-stage in this country; it causes small erect

362 MELAMPSORELLA

witches’-brooms on the Fir, the infected buds producing dense clusters of
small leaves which are not.pseudo-distichous (as in the normal shoots) but
spirally arranged, moreover they are not evergreen, but fall off in August,
On these leaves the spermogones and excidia are seen in June; and the
zcidiospores infect the second host and produce uredo-sori in 10—14 days.
It will be seen that the teleutospores found in spring do not belong to the
same generation as the uredospores found in the following summer.

The mycelium of both stages is perennial, so long as the host survives,
this being the only known instance of such a state of things ; for this
reason the parasite can maintain itself on either the Fir or the other host
quite independently. On the Fir, it produces in time large cankerous
growths. There is some evidence that a specialisation into biological
races on different species of Alsinee has begun. In the latter the

On 008
©, 28 ot So% B2

°
a
o
°
\g

Fig. 270. M. Caryophyllacearum. a, cells of the peridium of a uredo-sorus,
on Stellaria graminea, showing the opening pore being formed beneath a
stoma, seen through the epidermis, x 600; b, an epidermal cell of S.
nemorum, containing teleutospores (after Fischer) x about 400 ; c, attacked
leaf of S. graminea, slightly enlarged; d, one young and two mature
uredospores, x 600.

mycelium grows up with the young shoots and gradually spreads upwards
into every leaf, making the leaves in many cases smaller, but not much
different in colour. I have seen a few uredo-sori on the upper side of the
leaves, and even on the sepals of Cerastium.

The peridium of the uredospores is formed of a definite hemispherical
pseudo-parenchymatous layer, one cell thick, most of the cells being
polygonal by mutual pressure, nearly smooth, and containing at first a
little of the same yellow oil as the uredospores. It bears a very close
resemblance in texture to that round the uredo-sori of Pucedniastrum
Circaeae, Milesina Blechni, Uredinopsis filicina, and their allies.

The heterecism of this parasite was established by Fischer in 1901
(Zeitschr. f. Pflanzenkr. xi. 321) ; the result was attained only after a long

MELAMPSORELLA ‘ 363

and vain search extending over many years; it has been confirmed by
Tubeuf, Klebahn, and Bub4k. The ecidia of three other species, Melam-
psorella Symphyti, Pucciniastrum pustulatum and Calyptospora Goeppertiana,
which also live on the same host, do not cause any deformation of the
shoots, and can therefore be easily distinguished. ,

DISTRIBUTION : Europe, North America.

2. Melampsorella Symphyti Bubdak.

Oredo Symphytt DC, Encycl. viii. 232. Plowr. Ured. p. 255. Sace.
Syll. vii. 861.

Trichobasis Symphyti Lév. ; Cooke, Handb. p. 529.

Coleosporium Symphyti Fckl. Symb. Myce. p. 48. Cooke, Micr. Fung.
p. 218.

Melampsorella Symphyti Bubdk, Centralbl. f. Bakter, 2. xii. 423.
-Fischer, Ured. Schweiz, p. 523.

[Spermogones. Chiefly hypophyllous, crowded, often spread
over the whole leaf, orange-yellow.

Aicidiospores. Aicidia hypophyllous, in two rows parallel
to the mid-rib, not crowded, shortly
cylindrical 4—#% mm. high, opening at
the summit by a cleft, at length torn
to the base into 8—5 segments; spores.
orange, verrucose, 20—40 x 18—29 y.]

Uredospores. Sori hypophyllous,
small, rounded, closely crowded, rich
golden-yellow, often spread over the Fig. 271. -M. Symphyti.
whole leaf, at first covered by the eae a
epidermis and a thin peridium, then
pulverulent ; spores ovate or ellipsoid, finely verruculose, yellow,
28—35 x 21—28 p, without evident germ-pores.

[Teleutospores. Hypophyllous, developed within the epi-
dermal cells, forming large whitish or pinkish patches, many
crowded in each cell, pale-yellowish, smooth, 11—18 x9—15 u.]

[Aicidia on leaves of Abies pectinata, May, June;] uredo-
[and teleutospores] on Symphytum officinale, rather uncommon,
May to September. (Fig. 271.)

Only the uredospores have so far been recorded for Britain, but the

teleutospores would in all probability occur on the same plants at a later
date, while the zcidia could doubtless be found, if carefully looked for, in

364 PUCCINIASTRUM

woods where Abies and Symphytum both grow together. This was the
experience of Bubdk (see Ber. Deutsch. Bot. Gesell. 1903, xxi. 356). The
teleutospores appear to arise (or at least to mature) about May on a
perennial mycelium from the previous year, and are capable of immediate
germination: their basidiospores infect the young Fir-needles, but the
mycelium thus produced is not perennial and causes no witches’-brooms,
The ecidiospores are distinguished from those of the other species which
grow upon the same host by their much larger size. The whole of the
foregoing particulars (except as regards the uredospores) are due to Bubak
and Fischer ; the former found his uredospores chiefly upon Symphytum
tuberosum.
It will be useful to tabulate here by what characters the various excidia
on Abies pectinata can be discriminated : that belonging to
Melampsorella Caryophyllacearum causes witches’-brooms.

Melampsorella Symphyti has large spores 20—40 x 18—29 p.
Puceiniastrum pustulatum has a smooth line down the spore.
Calyptospora Goeppertiana has spores 21—30 x 14—18 p.

DISTRIBUTION : Europe.

PUCCINIASTRUM Otth.

Hetercecious or eecidia unknown.

Teleutospores extracellular, in a single layer, subepidermal,
with a brownish membrane, divided by vertical septa into
2—4 cells. Uredo-sori surrounded by a delicate hemispherical
peridium, opening at the summit with a pore; uredospores
yellow in mass, with indistinct or no germ-pores.

Aicidia with a thin cylindrical peridium (so far as known) ;
eecidiospores verrucose except on one side which is thinner and
smooth (?always), not provided with germ-pores.

1. Pucciniastrum Agrimoniz Tranzschel.

Uredo Potentillarum var. Agrimoniae DC. Flor. fr. vi. 81.

U. Agrimoniae Plowr. Ured. p. 255. Sace. Syll. vii. 839.

Coleosporium ochraceum Fekl. ; Cooke, Micr. Fung. p. 218.

Puceiniastrum Agrimoniae Tranz. Script. Bot. Hort. Petrop. iv. 301.
Fischer, Ured. Schweiz, p. 465. Arthur, N. Amer, Fl. vii. 106.

Uredospores. Sori chiefly hypophyllous, pulvinate, small,
confluent, sometimes spread over the whole leaf, covered by the.

epidermis and surrounded by a thin peridium which opens at
the summit with a pore, orange-yellow, fading to ochraceous;

PUCCINIASTRUM 365

spores shortly ellipsoid or obovate, echinulate, orange, 18—21 x
14; epispore rather thick, with indistinct

germ-pores. y
[Teleutospores. Sori similar, but indefinite, \

clear-brown; spores subepidermal, extracellular,
cuneate, smooth, each divided into four cells . _ Fig. 272.

: é P. Agrimoniae.
by two longitudinal walls at right angles to Uredospores.

one another, 30 x 21—30 u.]

On Agrimonia Eupatoria. Uredospores common, July—
September ; teleutospores, very rare everywhere, not yet found
in Britain. (Fig. 272.)

We owe our knowledge of the teleutospores to Tranzschel and Dietel ;
see Engler u. Prantl, Natiiyl. Pflanzenfam. vol. i. pt. 1**, p. 24. Until
they were discovered, the position of the
fungus was quité uncertain. Klebuhn (see
Zeitschr. f. Pflanzenkr. 1907, xvii. 149)
proved that the parasite could maintain
itself by over-wintered uredospores.

DISTRIBUTION : Europe, Asia,
North and South America.

2, Pucciniastrum Circee Speg.

Uredo Circaeae Schum. Pl. Sall. it. 228.
Cooke, Micr. Fung. p. 217, pl. 7,

f. 135, 136.

Pucecinia Cireaeae Pers. ; Cooke, Handb.
p. 507 p.p.

Melampsora Circaeae Winter ; Plowr.
Ured. p. 245. .

Puceiniastrum CircaeaeSpeg. Dec.Myc. Fig. 273. P. Circaeae. a, half

g 7: ‘ of a leaf of C. lutetiana,
65. Sacc. Syl. vil. 763. Fischer, showing greipeont Gliehily
Ured. Schweiz, p. 461, f. 302. enlarged); , uredospore

x 600; ¢, part of peridium
Uredospores. Sori hypophyllous, x 180; d, teleutospores, be-

on paler patches bounded by veins, Br MMs ae a

minute, yellowish, crowded, slightly confluent, surrounded by
the epidermis and by a peridium which opens at the summit
with a pore; spores ovate, 21—24 x 12—14,4; epispore thin,
covered with minute distant warts, without evident germ-pores ;
paraphyses wanting; peridium usually opening beneath a stoma.

366 PUCCINIASTRUM

Teleutospores. Hypophyllous, in little subepidermal groups,
roundish or flattened at the sides, divided longitudinally into
about 2—4 cells, 17—24 x 21—28; epispore of uniform
thickness (about 2 4), clear-yellowish, smooth.

On Circaea intermedia, C. lutetiana. Rather common in the
uredo-stage. June—September. (Fig. 273.)

It is prubable that this, like Pucciniastrum pustulatum, has an ecidium-
stage on a conifer, but nothing has yet been discovered to confirm the
suspicion. Puceinia Circaeae has sometimes been confounded with this
species, though they have nothing in common but the host-plant. They
may both be found on the same leaf.

DISTRIBUTION : Central Europe.

3. Pucciniastrum pustulatum Dietel.

Uredo pustulata Pers. Syn. p. 219. Cooke, Handb. p. 526; Micr.
Fung. p. 215.

Melampsora pustulata Schrot. Pilz. Schles. p. 364. Plowr. Ured. p. 244.

Pucciniastrum Lpilobit Otth in Mittheil. Nat. Gesell. Bern, 1861, p. 72.
Sace. Syll. vii. 762. Fischer, Ured! Schweiz, p. 459.

P. Abieti-Chamaenerit Kleb. Wirtswechs. Rostpilze, p. 393.

P. pustulatum Dietel, in Engler u. Prantl, Nat. Pflanzen. i. 1**, 47.

Arthur, N. Amer. FI. vii. 107.
[Spermogones. Hypophyllous, abundant, subcuticular.
Acidiospores. cidia hypophyllous, whitish, mostly in two
rows corresponding to the two white lines,
4mm. wide, 1 mm. high, cylindrical, with
an evident peridium, opening by longitu-
dinal slits; spores mostly oval, 13—22 x
10—14 w (15—17u, Dietel); epispore finely
verrucose, but with a thinner and smooth
area which often forms a longitudinal strip. ]
Uredospores. Sori hypophyllous, caus-
ing reddish or yellowish spots on the upper
surface, roundish, | mm. wide, scattered or
Fig: 874 Pi) preee in little groups, often contiuent, orange,
latum. AScidia on surrounded by the peridium and the remains
leaf of A. pectinata, : :
slightlyenlarged;a, Of the epidermis; spores mostly oval, orange-
a peridium, x 10. yellow, distantly echinulate, 15—22 x 11—

From a foreign :
specimen. 14; paraphyses wanting.

PUCCINIASTRUM 367

Leleutospores. Sori hypophyllous, clustered or widely con-
fluent, {mm. wide, covered by the epidermis, reddish, then
blackish-brown; spores shortly cylindrical or prismatic, 17—
35 x 7—14 p, those in the middle of the sorus Melampsora-like,
but at the periphery roundish or oval, and composed of 1—3
cells, 1.e. divided by longitudinal walls; epispore clear-brown,
thickened (up to 3) above, but with a thinner spot (? germ-
pore).

[Aicidia on Abies pectinata, June, July ;] uredo- and teleuto-
spores on E’pilobium (Chamaenerion) angustifolium, E. palustre.
August—October. Shere, Abinger, Surrey. (Fig. 274.)

The ecidiospores are distinguished from those of Calyptospora Goep-
pertiana by their smooth area and generally shorter length. The ecidium
of Melampsorella Caryophyllacearum lives also on Abves pectinata, but
produces thereon witches’-brooms, and finally swellings and canker of the
branches, while P. pustulatum attacks the leaves only. Klebahn, Fischer,
and Tubeuf have all demonstrated experimentally the genetic connection
of the stages of this parasite, so far as concerns Z. angustzfolium ; but the
form on £. palustre and its allies is probably, at least biologically, distinct.
Klebahn’s name, quoted above, includes only the form on £. angustrfolium
and the allied £. Dodonaet. He restricts the name P. Abzeti-Chamaenerwt
to this, calling the form on £. palustre, E. montanum, etc. by the name
P. Epilobii, and assigns to them also certain small morphological differ-
ences. The collective species is conimon in North America, but the
zecidium has not yet been recognised there.

DISTRIBUTION: Europe, North America.

4, Pucciniastrum Pyrolz Dietel.

Eeidium Pyrolae Gmel. in Linn. Syst. Nat. ii. 1473.

Uredo Pyrolae Grev. Flor. Edin. p. 440 p.p. Fischer, Ured. Schweiz,
p. 539, f. 337.

Trichobasis Pyrolae Berk.; Cooke, Handb. p. 529; Micr. Fung.
p. 223 p.p.

Melampsora Pyrolae Schrét. ; Plowr. Ured. p. 247.

Pucciniustrum Pyrolae Dietel, in Engler u. Prantl, Nat. Pflanzen.
i. 1**, 47, Arthur, N. Amer. FI. vii. 108.

Thecopsora(?) Pyrotae Karst. ; Sace. Syll. vii. 766.

Uredospores. Sori hypophyllous, minute, hemispherical,
orange, on yellowish or purplish discoloured spots, surrounded
by the epidermis and by a peridium which both open at the

368 THECOPSORA

summit with a pore; spores elongated-ellipsoid or clavate,
provided with distant and pointed warts, yellowish, 26—35 x
1416; epispore rather thick, without perceptible germ-
pores.

[Leleutospores. Sori hypophyllous, adjoining the uredo-sori,
inconspicuous, flat, subepidermal, forming an
even layer of laterally united cells; spores
columnar or oblong, 24—28 x 10—12 py;
epispore colourless, uniformly thin (1 «)—
(description after Dietel).]

Fig. 275. P. Pyrolae. On Pyrola minor, P. rotundifolia. May
Uredospores, on P. October, uredospores only; Scotland,
rotundifolia. 5 ‘

Ludlow, ete. (Fig. 275.)

Teleutospores have been met with by few observers ; previous to their
discovery, it was uncertain in what genus the fungus should be placed.
It may possibly be hetercecious. The uredospores are often more coarsely
warted at one end, though this is not invariably the case. Fischer figures
the cells of the peridium round the pore as furnished above with pointed
warts, of which one is distinctly longer than the others: those cells are
enormously thickened on the lower wall.

DisTRIBUTION: Europe, North America.

THECOPSORA Magnus.

Hetercecious or ecidia unknown.

Teleutospores intracellular, occupying and filling the epi-
dermal cells of the leaves, united into a brown crust, other
characters as in Pucciniastrum. Uredo as in Pucciniastrum.
Aicidia hemispherical, with a thick brown peridium (so far as
known); scidiospores verrucose, with a narrow, thin, smooth
strip down one side (? always).

1. Thecopsora Padi nov. comb.

Licea strobitina A. et S. Consp. p. 109, pl. 6, f. 5.

Phelonitis strobilina Pers, ; Cooke, Handb. p. 409, f. 141.
Perichuena strobilina Fr. ; Greville, Scot. Cr. Fl. pl. 275.
eAvidium strobilinum Wint. Pilze, p. 260; Plowr. Ured. p. 266.

” ‘THECOPSORA 369

Uredo Padi K. et S. exsice. 187. Cooke, Handb. p. 527.
U. porphyrogenita Kze. ; Cooke, Micr. Fung. p. 216.
Melampsora Padi Cooke, Handb. p. 523 (1871). Plowr. Ured. p. 246.
Fung. Fl. Yorkshire, p. 184.
Pucciniastrum Padi Dietel in Eng. u. Prantl, Natiirl. Pflanz. i. 1**,
p. 47. Fischer, Ured. Schweiz, p. 463, f. 303; Centralbl. f.
_ Bakter. 2. xv. 227.
Thecopsora areolata Magn. in Hedwigia, 1875, p. 123. Sacc. Syll.
vii. 764,
Spermogones. Whitish, pustular, flat, open, exhaling a
strong smell.
4icidiospores. Aicidia crowded, covering on the upper
(sometimes the under) side the lower
part of the scales of the fallen cones,
hemispherical or polygonal; peridium
thick, brown, woody, opening by a
slit; spores oval, inequilateral, yellow,
21—28 x 1720; epispore very
thick (up to 6), echinulate, with a
narrow, thinner, smooth stripe.
Uredospures. Sori hypophyllous,
clustered on spots 1—5 mm. wide Fig. 276. Th. Padi. a, leaf
which are brownish above, reddish of P. Padus, showing uredo-

; sori; b, scale of cone of Picea
or purplish below, and more or less excelsa, showing wxcidia (both

bordered by the veins, covered by the ee
epidermis and by a peridium which

opens at the summit by a pore; spores oblong-oval or irregular,
echinulate, pale-yellowish, 15—21 x 10—14; epispore about
13 thick.

Teleutuspores. Developed in the epidermal cells, several in
each, epiphyllous or occasionally hypophyllous, forming dark-
brown shining crusts which are bounded by the veins; spores
oval-cylindrical or prismatic, 22—30 x 8—14 y, divided by thin
longitudinal walls into 2—4 cells; epispore thin, slightly
thickened above, clear-brown, smooth, with a germ-pore in the
upper and inner corner of each cell..

Acidia on cones of Picea excelsa, Scotland, Yorkshire,
August, November; uredo- and teleutospores on Prunus Padus,
August, September. Very rare. (Fig. 276.)

G. U. 24

370 THECOPSORA

The connection of the spore-forms on these two hosts has been experi-
mentally demonstrated by Klebahn, Tubeuf, and Fischer. The basidio-
spores in spring (about the time of pollination of the Fir) infect the female
flowers of the Spruce Fir, which are usually at the top of the high trees ;
occasionally also the young shoots are affected, but they do not produce
zcidia. The ecidia are developed in late summer, and mature on the
fallen cones; their spores germinate in the following May, and then infect
the leaves of the Bird Cherry, on which they produce uredospores in the
summer and teleutospores in the autumn (Fischer, J.c. and Centralbl. f.
Bakter. 2, xv. 1906, p. 227). The description of the teleutospores is
taken from Klebahn and Fischer. As will be seen from the synonymy,
the ecidium was originally classed among the Myxomycetes. The three
stages appear in Cooke’s Handbook (according to the knowledge then
prevailing) on three different pages, the ecidium from Appin, the uredo
from some place in Scotland, and the “Melampsora” from Swanscombe,
Kent (Cooke, 1865). It is also recorded on P. Padus in Yorkshire Fung.
FI. p. 184, while the eecidium is recorded on ‘pine-cone scales” on p. 369.
The uredo has also been found at Braemar, Aboyne, Perth, etc.; and the
zecidium in Dumfriesshire.

DISTRIBUTION: Europe.

2. Thecopsora Galii De Toni.

Caeoma Galii Link, Sp. Pl. ii. 21.
Melampsora Galiz Wint. Pilze, p. 244. Plowr. Trans. Brit. Myc. Soc.
i. 59.

Puceiniastrum Gali Fischer, Ured. Schweiz, p. 471, f. 307.

Thecopsora Galii De Toni in Sace, Syll. vii. 765.

Uredospores. Sori scattered or gregarious, small, round,
pulvinate, reddish, covered by the epidermis and by a peridium
which opens at the summit with a pore; spores shortly ellipsoid
or ovate, sparsely echinulate, orange-yellow, 17—20 x 14—16 yp;
epispore colourless, without evident germ-pores.

Teleutospores. Developed in the epidermal cells, forming
httle dark-brown crusts, crowded, roundish, longitudinally
septate into 2—4 cells, 21—24 x 21—32 yu; epispore rather
thick, yellowish-brown, smooth, with an evident germ-pore at
the upper and inner corner.

On Galium verum (H. T. Soppitt), June—September, 1889.
Very rare,

It is reported, in continental Europe, as occurring on other species
of Galium, also ou Sherardia and Asperula.

THECOPSORA 371

3. Thecopsora Vacciniorum Karst.

Uredo Vacciniorum Link, Sp. Pl. ii. 15. Wonks, Handb. p. 527;
Micr. Fung. p. 216.

Melampsora Vacciniorum Schrét.; Plowr. Ured. p. 246.

Puceiniastrum Vacciniorum Dietal, in Eng. u. Prantl], Nattirl. Pflanz.
i. 1**, 47. Fischer, Ured. Schweiz, p. 467, f. 305.

P. Myrtillé Arthur, North Amer. Fl. vii. 109.

Thecopsora Vaceiniorum Karst. Bidr. Finl. Nat. xxxi. 59. Sac. Syll.
vii. 765.

Uredospores. Sori hypophyllous, rather small, scattered or
in groups on (sometimes orange) dis-
coloured spots which are visible for /
some time before the sori appear, long
covered by the epidermis, brownish,
surrounded by a peridium which opens Fiz. 277. Th. Vaccini-
at the summit with a pore; spores T Myrtilles (Rontkma)
ovate to ellipsoid, sparsely echinulate-
verrucose, orange-yellow, 15—16 x 12—18p, (21—-28 x 14—
18 », Fischer).

[Teleutospores. Hypophyllous, developed in the epidermal
cells which are filled by them and form little brown indehiscent
crusts, pale-brown, divided by longitudinal walls, 14—17 x
7—10; epispore uniformly thin.]

On leaves of Vaccinium Myrtillus, V. Vitis-cdaea. Uredo-
spores only, May—October, Shropshire, Yorkshire, North Wales,
Scotland. (Fig. 277.)

The teleutospores are very rare, and can be found only on the dead or
fallen leaves. Fischer suggests that this may be a hetercecious species as
indeed is most likely the case, though no probable suggestion has yet been
made as to the nature of the alternate host. He records it on V. uligino-
sum also; and Arthur on several other species of Vacciniacesw, in the
United States.

DistRisuTIoN: Europe, North America.

bo
ree
.

372 CALYPTOSPORA

CALYPTOSPORA J. Kiihn.

Hetercecious. Teleutospores intracellular, occupying the
swollen epidermal cells all round the stem of the host for a
considerable distance,_ otherwise as in Pucciniastrum. No
uredo. Kcidia cylindrical, with a thin peridium; zcidiospores
verrucose, without germ-pores, and with no smooth spot.

Calyptospora Goeppertiana Kiihn.

Zicidium columnare A. et 8. Consp. p. 121, pl. 5, f. 4.

Fig. 278. C. Goeppertiana.
Affected branch of V. Vitis-
idaea, Scotland (slightly re-
duced); a, leaf of A. pecti-
nata, b, leaf of A. Nordman-
niana, with ecidia, Torquay
(slightly enlarged),

Peridermium columnareS. et K.; Cooke,
Handb. p. 535; Micr. Fung. p. 194,
pl. 2, f 27, 28.

? HMeidium pseudo-columnare Plowr.
Ured. p. 271 (not Kiihn 2).

Calyptospora Goeppertiana Kiihn in
Hedwigia, 1869, viii. 81. Sace.
Syll. vii. 766. Kew Bulletin, 1907,
p. 1, with plate.

C. columnaris Kiibn (1886); Arthur,
N. Amer. Flor. vii. 114.

Pucciniastrum Goeppertianum Keb.
Wirtswechs. Rostp. p. 391. Fischer,
Ured. Schweiz, p. 466, f. 304.

Afcidiospores. Aicidia hypophyl-
lous, arranged in two long rows
parallel to the mid-rib, cylindrical,
white, with torn or slit margin;
spores ellipsoid, uniformly verrucose,
orange-red, 21—30 x 14—18 yp.

Teleutospores. Caulicolous (the
fusiform attacked part of the stem
being swollen and at first clear-
pink, passing into brown), developed
in the epidermal cells, densely
crowded, prismatic, mostly divided
by crossed longitudinal walls into
four cells, up to 42 «4 high; epispore

yellowish-brown, smooth, thickened (up to 3) at the summit,
with a germ-pore at the upper and inner corner of each cell.

CALYPTOSPORA 373

ficidia on leaves of Abies pectinata, A. Nordmanniana,
June—September; teleutospores on stems of Vaccinium Vitis-
idaea, July—September. England, Wales, Scotland. Very
rare. (Fig. 278.)

The life-history has been experimentally demonstrated by Hartig,
Kiihn, and Bubak. It has been shown that the ecidia can be developed
in artificial cultures on other species of Abzes (but not on Tsuga canadensis
or Pseudotsuga Douglasiz), though it is not recorded on any of these in
natural conditions. Saccardo’s citation of “Abies canadensis” in the
Sylloge is probably an error. The teleutospores are recorded also on
several other species of Vaccinium, including V. Myrtillus A. Gray, in the
United States, but the «cidia have not been observed there.

The infested branches of the Cowberry stand perfectly erect; the
plant becomes taller and the leaves stunted. No uredo-stage occurs.
For the life-history see p. 59. The mycelium is perennial in the Cow-
berry, producing fresh crops of teleutospores year after year; this may be
the origin of the (presumably) erroneous statement in the Kew Bulletin
(2.c.) that the basidiospores are able to infect the Vaccinium again, as well
as the Silver Fir. This statement was originally made by Hartig, but is
unsupported by any experimental evidence.

When planting any of the species of Abzes liable to attack, it would be
well to look for and burn all infested bushes of Cowberry in the neighbour-
hood ; they are easily recognisable by their peculiar habit. The fungus
cannot attack the Firs unless the infested Cowberry is near enough to
convey the infection. ?

DisTRIBUTION: Europe, North America.

HYALOPSORA Magnus.

Teleutospores in one or two layers, produced in the
epidermal cells which are united into crusts; spores with
colourless membrane, each divided by vertical septa into 2—4
(or more) cells. Uredo-sori subepidermal, without a peridium
or with a very rudimentary one, but surrounded by paraphyses,
uredospores of two kinds, yellow, sessile, furnished with evident
germ-pores. On Ferns.

It has been suggested that the species of Hyalopsora are
hetercecious, Abies and Pinus being named by Bubak (1906) as

374 HYALOPSORA

possible alternate hosts. This is in itself improbable and is
inconsistent with Arthur’s suggestion that pf the two kinds of
uredospores the first kind represents zcidiospores.

1. Hyalopsora Aspidiotus Magn.

Uredo Polypodti Schrét. Krypt. Flor. Schles. iii. 374. Plowr. Ured.
p. 256 p.p. Sacce. Syll. vii. 857 p.p.

U. Aspidiotus Peck in Ann. Rep. N. Y. State Mus. xxiv. 88.

Melampsorella Aspidiotus Magn. Ber. Deutsch. Bot. Gesell. xiii. 288.

Puceiniastrum Aspidiotus Karsten, Bidr. Finl. Nat. Folk, 1879, xxxi.
143.

Hyalopsora Aspidiotus Magn. Ber. Deutsch. Bot. Gesell. 1901, xix.
582, Arthur, N. Amer. Flor. vii. 112.

H. Polypodii-Dryopteridis Magn. in Hedwig. 1902, p. 224. Fischer,
Ured. Schweiz, p. 472, f. 308.

Uredospores. Sori amphigenous, scattered, small, round,
up to $mm., golden-yellow, without a peridium, dehiscing
irregularly, often seated on yellowish spots; spores oblong or
ellipsoid, of two kinds, (1) thick-walled (24—34 4), with very

Fig. 279. H. Aspidiotus. Part of frond of P. Dryopteris, showing uredo-sori,
nat. size ; three spores, all from the same sorus (Scotland).

faint, hardly perceptible warts, 36—72 x 30—40 p, with 6—8
scattered germ-pores, (2) thin-walled (about 14,), covered
uniformly with very faint scattered warts, 283—40 x 16—26 xu,
with four indistinct equatorial germ-pores ; paraphyses few.
Teleutospores. In the epidermal cells, often filling them
completely, roundish or irregular, flattened where they are in
contact, sometimes arranged in two layers, about 25 u high,
21—35 w or more wide, divided by vertical septa into 3—5

HY ALOPSORA 375

(mostly four) cells; epispore thin, smooth, colourless; germ-
pore not perceptible.

On Polypodium Dryopteris. Uredospores, June—August ;
teleutospores, May and June, on young leaves (Magnus). Rare.
(Fig. 279.)

Fischer, who records it also on P. Robertianum, states that the teleuto-
spores germinate in June. _ Arthur prefers to call the first kind of
uredospore the ecidiospore; this is possibly correct but, until something
is known about their development, it is premature to decide. According
to him, the ecidia have no peridium, and the uredo-sori a very rudimentary
one, but I have repeatedly found both kinds of spores in the same sorus.
They appear perfectly smooth, when seen in water.

DISTRIBUTION : Europe, North America.

2. Hyalopsora Polypodii Magn.

Uredo linearis var. Polypodii Pers. Syn. p. 217.

U. Filicum Desm.; Cooke, Handb. p. 526; Micr. Fung. p. 215 p.p.
White, Scot. Nat. 1877, iv. 27, pl. 2, f. 7.

U. Polypodiz DC. Flor. fr. vi. 81. Plowr. Ured. p. 256 p.p. Fungus
Flor. Yorkshire, p. 204. Sacc. Syll. vii. 857 p.p.

Pueciniastrum Polypodii Dietel, in Hedwig. 1899, xxxviii. 260.

Hyalopsora Polypodii Magn. Ber. Deutsch. Bot. Gesell. 1901, xix. 582.
Fischer, Ured. Schweiz, p. 474, f. 309. Arthur, N. Amer, Flor,
vii. 112. Dietel, Annal. Mycol. 1911, ix. 530.

Uredinopsis Polypodii Liro, Uredin. Fennic. 1908, p. 496.

Uredospores. Sori hypophyllous, minute, scattered, bullate,
golden-yellow, without a peridium, rup-
turing irregularly; spores more or less
globose or ellipsoid, of two kinds, (1)
thick-walled (2—3 »), with very faint
warts, 26—38 x 18—29y, with 6—8
scattered germ-pores, (2) thin-walled _ .
(1—14 #), covered uniformly with faint ae eee a laces
distant warts, 22—35 x 13—20 yw, with pee fragilis (Shrews-
four equatorial germ-pores.

Teleutospores. In the epidermal cells, often filling them
completely, showing as yellowish-brown spots on the under side
of the leaf, densely crowded, divided into 2—4 cells, each about
14—18 in diam., single cells subglobose; epispore thin,
colourless; germ-pore perceptible at the upper end.

376 MILESINA ;

On Cystopteris fragilis (=Polypodium fragile Linn.) and its
var. dentata. June—September. Rather rare (2700 ft. on Ben
Lawers); occurring freely on cultivated Cystopteris in fern-cases,
(Fig. 280.)

The same remarks may be made about the uredospores of this as of
the previous species. In both cases it is not certain that the teleutospores
have been found in Britain, and the description is taken from Dietel and
Fischer. Dietel has shown (Ann. Mycol. ic.) that infection by the
uredospores can be performed easily in a room on C. fragilis; time of
incubation fourteen days. This would, therefore, be a very convenient
species for cytological investigation among the lowest types, but Dietel
obtained zo teleutospores.

DisTRIBUTION: Europe, North America.

MILESINA Magnus.

Teleutospores intracellular, hyaline, septate. Uredo-sori
furnished with a peridium which opens at the summit with a
pore; uredospores hyaline, pedicellate, with a thin epispore,
but without germ-pores. On Ferns.

1. Milesina Dieteliana Magn.

Milesia Polypodit B, White, Scot. Nat. 1877, iv. 162, pl. 2, f. 5. Sace.
Syll. vii. 768,

Melampsorelia Dieteliana Sydow, in Mycoth. Germ. no. 62 (1908).

Milesina Dieteliana Magn. Ber. Deutsch. Bot. Gesell. xxvii. 325 (1909).

Uredospores. Sori hypophyllous, small, round, pustular,
clustered loosely in small groups

on brown irregular spots, pale-
brown ; spores colourless, oblong
or obovate, strongly but sparsely
and irregularly echinulate above,

Fig. 281. M. Dieteliana. Uredo- smooth below, 22—30 x 15—16 4;

spores, on P. vulgare var. serra- wall moderately thick (14—2 p).
tum (C. H. Wright). They were Tel i
enclosed in a thin white peridium. [Leleutospores. Not found in

Britain. ]

MILESINA 377

On Polypodium vulgare var. serratum, Scotland, December,
1906 (C. H. Wright in Herb. Kew). On Polypodium vulgure,
Dolgelly, May, 1913 (A. D. Cotton). (Fig. 281.)

The genus Milesia is now dropped, because it was founded on an
imperfect state which might belong to any one of several genera.

2. Milesina Blechni Sydow.
Uredo Scolopendrii Fckl.; Plowr. Ured. p. 256 p.p. Sacc. Syll. vii.
860 p.p.
Melampsorella Blechni Syd. Annal. Mycol. 1903, p. 537.
Milesina Blechni Sydow in Mycoth. Germ. no. 877 (1910).

Uredospores. Sori hypophyllous, hemispherical, pustular,
4—3 mm. diam., yellowish, loosely scattered on green or brownish
leaf-segments, enclosed in a thin white peridium, opening at

Fig. 282. M. Blechni. «, cells of peridium, x 360; 5, uredospores, x 600;
c, portion of frond of B. Spicant, showing uredo-sori, nat. size.

the summit by a round pore which always begins to be formed
at a stoma; spores colourless, oblong, obovate or clavate, faintly
or irregularly echinulate, 32—45 x 12—18 uw; epispore 142 pu
thick.

[Teleutatpores. Unknown in Britain.]

On Blechnum Spicant. Very uncommon. July—Septem-
ber. (Fig. 282.)

This fungus closely resembles the Milesinu on Polypodium vulgare,
and was included under the name Milesia Polypodii B. and B. White.
The markings on the spores of this and the allied species are more often
of the nature of mere roughnesses than like the neat and regular echinula-
tion of the higher types (Puccinza, etc.).

378 MILESINA

3. Uredo Scolopendrii Schrét. (probably a Milesina).

Ascospora Scolopendrii Fckl. Symb. Myce. Nacht. ii, 19.

Uredo Scolopendrit Schrét. Krypt. Flor. Schles. p. 374. Plowr. Ured.
p. 256 p.p. Sace. Syll. vii. 860 p.p.

U. (2) Pteridum White, Scot. Nat. 1877, iv. 27, pl. 2, f. 6.

Milesia Polypodii White p.p.; see Gard. Chron. 1892, 3. xii. 776
(specimen on Scolopendrium in Herb. Cooke).

Uredospores. Sori similar to those of Milesina Blechni, but
larger, about 1 mm., brown, in loose irregular groups; spores

Fig. 283, U. Scolopendrii. Portion of frond of Scolopendrium, showing
sori, nat. size; three uredospores, x 600.

very numerous, oozing out, colourless, clavate, strongly echinu-
late all over, especially at the apex, 35—45 x 15—23 p; epispore
13—2 p thick.

On Scolopendrium vulgare. Rare; Scotland, England
(Cooke), Forden (Rev. J. E. Vize), Warton, N. Lancs., March,
1908 (J. W. Hartley in Herb. Kew). (Fig. 283.)

- Only known in the uredo-stage, but closely allied to Mf. Blechni. The
name, Uredo Scolopendrit, has been used to include many other species.

Besides these parasites on Ferns, Plowright also records under Uvedo
Polypodii (p. 256) a fungus on Adiantum Capillus-Veneris. If this is
correct, it was probably Hyalopsora A dianti-Capilli- Veneris Sydow (Annal.
Mycol. 1903, p. 248), which, I am informed by Herr H. Sydow, occurs in
Istria and North Italy.

It is quite possible that two others may be found in Britain, viz.
Milesina Kriegeriana Magn, on Aspidium spinulosum and Credinopsis
Jjiticina Magn. on Polypodium Phegopteris, but I have seen no British
specimens. The latter is so likely to occur here, that it is advisable to
give a description of it.

UREDINOPSIS 379

UREDINOPSIS Magnus.

Teleutospores solitary, extracellular, hyaline, septate. Uredo-
sori subepidermal, with a distinct peridium ; uredospores hyaline,
pedicellate, without germ-pores. On Ferns.

This genus is distinguished from all others by the fact that
the teleutospores are dispersed without order among the cells
of the mesophyll. There are three known forms of spores,
which seem to occur simultaneously, and not in a fixed order of
succession as in most pleomorphic Uredinales. It is one of the
lowest genera from the point of view of organisation; and its
cytological investigation would doubtless yield a rich harvest of
new ideas,

Not yet discovered in Britain. .

Uredinopsis filicina Magn.

Protomyces ? filicinus Niessl, in Rab. Fung. Europ. no. 1659, 1873.

Uredo Polypodii var. Phegopteris Winter, Pilze, p. 253. Sacc. Syll.
vil. 858, .

Uredinopsis filicina Magn. Atti Congress. Bot. Internaz. 1892, p. 167.
Fischer, Ured. Schweiz, p. 476, f. 310, 311.

Uredospores. Sori hypophyllous, small, roundish, thickly
scattered, yellowish-brown, of two kinds—(1) smaller, about

-

b, uredospores (no. 2) and paraphysis; c, teleutospores. From a German
specimen, issued by Sydow.

4 mm. diam., flattish, semi-transparent, always situated beneath

a stoma, surrounded by a rather tough peridium, rupturing

above; spores ;iore or less obovate-polygonal, on slender
“8

?
ra

380 UREDINOPSIS

pedicels as long as or longer than the spore, colourless, densely
verruculose, averaging 20—26 x 12—14 4; epispore about 1 p
thick ; (2) larger, about 4mm. diam., more elevated, pustular,
without (?) peridium, but with what looked like hyaline clavate
thin-walled paraphyses; spores ovate-fusiform, colourless, smooth,
thickened at the apex with a conical, acute, usually oblique
process, 85—45 x 10—12 yw; pedicels short; epispore thin.

Teleutospores. Scattered singly throughout the mesophyll,
Le. not in definite clusters, roundish-oval or oblong, 1—3
(mostly two) celled, 18—24~x 15—16 4; epispore very thin,
almost perfectly hyaline and smooth.

On Polypodium Phegopterts. Europe. Not uncommon;
appearing about the time when the fern-sori are being formed.
The uredo-sori are of about the same size as a fern-sporangium,
and are scattered among the fern-sori. (Fig. 284.)

The description and figures are taken from a German specimen. The
fungus will no doubt be found in this country, if looked for ; it is rather
conspicuous on account of the uredo-sori. The warts on the uredospores
(no. 1) are clearly perceptible even when wet; the peridium is pseudoparen-
chymatous, with cells isodiametric near the apex, becoming longer and
prosenchymatous towards the periphery ; texture rather tough. and con-
sistent, not friable. Fischer gives a beautiful and accurate drawing of the
nature of the peridium and its relation to the epidermis (/.c. fig. 310).

The uredospores (no. 2) are described by Arthur in his generic character
(N. Amer. Flora, vii. 115) as smooth, except for two longitudinal, thickened,
more or less warted ridges, which I could not discern.

As in Hyalopsora, Arthur prefers to call the two kinds of uredospores
respectively eecidio- and uredo-spores: the same remark may be made as
under that genus, for the supposed ecidiospores occur in company with
the teleutospores on the dying parts of the frend, and are said to germinate
only after having passed the winter. Uredospores (no. 2) can germinate
at once,

The teleutospores are to be found in large numbers in the tissues
immediately round both kinds of sori, and simultaneously with them.
They are borne on short lateral branches of the mycelium. Dietel, who
gave a long account of this fungus (Ber. Deutsch. Bot. Gesell. 1895, xiii.
326), showed that in the allied European species, U. Struthiopteridis, the
teleutospores germinated easily with a typical basidium (occasionally
branched) which bore round basidiospores.

DistrisuTion: Germany, Switzerland, Italy.

APPENDIX
HEMILEIA Berk. et Broome.

Ecidia, if any, unknown, Uredospores borne singly on
hyphe which protrude in fascicles through a stoma. Teleuto-
spores formed later on pedicels in the centre of the same
fascicles, one-celled, with apical germ-pore, germinating as in
Uromyces.

The character given for Hemileia, when only H. vastatrix
was known, of having one side of the uredospore smooth, is now
known to be not of universal application.

Hemileia americana Mass.

Hemileia americana Mass. in Gard. Chron. 1905, xxxviii. 153, f. 53;
Kew Bulletin, 1906, p. 40, with plate.

Uredospores. Sori hypophyllous, forming broadly effused
pulverulent deep-orange patches,
often several centimetres in ex-
tent; spores perfectly spherical,
shortly stipitate, 24—32, diam.;
epispore bearing small, rather
sparsely scattered, round warts,
and with two germ-pores.

Teleutospores. Occupying the
centres of the heads of uredo-
spores, shortly stipitate, colour-
less, broadly obovate or turbinate,
often with a small obtuse apl- Fig. 285. H. americana. Tip of
culus, densely verruculose, 30 x Cattleya leaf, showing sori (slightly

reduced); a, uredospore, and b,
22—25 p. teleutospore, x 600.

382 HEMILEIA

On leaves of Cattleya Dowiana Batem., imported from
Costa Rica, 1899. (Fig. 285.)

Only a small patch of Rust was present on the leaf when the plant
was received from Costa Rica, but this continued to increase in size and
the falling spores infected other leaves. The uredospores germinated
readily, and young Cattleya leaves, inoculated on the under surface,
produced mature uredospores in thirteen days. No success attended the
efforts to infect other orchids, not belonging to the genus Cattleya.

This description is founded on that given in the Kew Bulletin; on
referring to the Gardener's Chronicle (/.c.) it will be seen that the
particulars there given differ in several respects. The specimen is in the
Kew Herbarium. This fungus and the others of the same genus might
easily become dangerous parasites in orchid houses, if allowed to spread ;
though it seems probable, on the slender evidence at present known, that
each is confined, like most other Rusts, to a single genus, if not species.

Hemileia Phaji Sydow.

Oredo Phojr Raciborski, Parasit. Alg. und Pilz. 1900, ii. 32.

U. Lynch Adams, Irish Naturalist, 1911, xx. 68.

Hemileia Phaji Sydow, Monogy. iii. (ined.). Grove, Journ. Bot. 1913,
p. 44.

Fig. 286. H. Phaji. a, epidermis, showing fascicle of uredospores emerging
from a stoma, x 180; b, the same in section; c, uredospores, x 600; d, a
portion of a leaf, with uredo-sori, in Herb. Kew, x}.

Uredospores. Sori hypophyllous, densely scattered, round,
distinct, formed of little dense fascicles of hyphe (20—25)
which issue through a stoma; pedicels divergent, clavate up-
wards, each surmounted by a spore; spores subglobose or rarely

HEMILEIA 383

obovate, strongly but sparsely echinulate-verrucose all over,
13—15 mw diam. or 18—25 x 12—16 uw.

On Phajus Wallichti. Botanic Gardens, Glasnevin, Dublin
(Sir Frederick Moore). April. (Fig. 286.) .

The description and figures a, b, and ¢ are derived from some slides
mounted by Sir Frederick; I have not seen the leaves, which were not
preserved, but there is an exactly similar form on Phajus sp. in Herb.

Kew, unlocalised but apparently sent up by some gardener for identifica-
tion. This also was named Uredo Lynchii at first.

Hemileia Oncidii Griff. et Maubl.
Hemileaa Oneidii Griff. et Maubl., Bull. Soc. Myc. Fr. 1909, p. 138,
pl. vi.
Uredospores. Sori hypophyllous, numerous, pulverulent,
orange, minute, 50—100 w diam.; fertile hyphe issuing from
stomata, fasciculate, branched, 25—30 yu long, clavate above;

ms

Fig. 287. H. Oncidii. Uredospores, a, wet, b, dry : c, part of leaf, showing
uredo-sori, x}. From the specimen in Herb. Kew.

spores globose or rarely obovate, echinulate-verrucose, filled
with orange drops, 16—18, diam., occasionally 20 long;
epispore hyaline.

[Teleutospores. Sori growing in the centre of the spots,
pallid-brownish ; spores subglobose or pyriform, at first hyaline,
aculeate, then pallid-brown and somewhat smooth, 20—23 x
15—20 p.]

On Oncidium varicosum, imported by Messrs Stuart Low
& Co. from San Paulo, Brazil, August, 1909. Specimen in
Herb. Kew. (Fig. 287.)

In the Kew specimen, the spots occupied by the crowded groups
of uredo-sori are more or less oval, $—1 cm. across, and covered with
orange-yellow dust, but as the sori are limited, each by the stoma
through which it issues, they never become confluent. The description is
founded upon that of Griffon and Maublanc; only uredospores were seen.
What is doubtless the same species has been found at the Botanic

384 CHRYSOMYXA

Gardens, Glasnevin, Dublin, on Oncidium varicosum and O. Forbesti.
Others of the same character were also seen there by Sir Frederick Moore
on Epidendrum vitellinum and Lycaste Skinneri. These tropical parasites
are imported with the plants, and occasionally spread to a small extent,
under glass, in this country. Persistent examination of such imports
would no doubt discover still others of the same kind,

Chrysomyxa Rhododendri De Bary.

Aeidium abietinum A, et S. Consp. p. 120 p.p.

Uredo Rhododendri DC. Flor. fr. vi. 86.

Chrysomyxa Rhododendri De Bary, in Bot. Zeit. 1879, p. 809, pl. 10,

f.1—6. Sacc. Syll. vii. 760. Fischer, Ured. Schweiz, p. 426.

[&cidiospores. Aicidia irregular, membranaceous, com-
pressed and elongated parallel to the mid-rib of the leaf (up
to 3 mm. long), erumpent on transverse yellowed zones, at length
irregularly torn, whitish; spores more or less ellipsoid, yellowish,
1745 x 12—22 w; epispore thin, thickly verruculose except
for a smooth longitudinal stripe.]

Uredospores. Sori almost always hypophytlous, on yellowish
or brownish spots, minute, roundish or oblong, scattered or in
small groups, at length pulverulent, orange; spores in chains
with intercalary cells, irregular or oval, verruculose, orange-
yellow, 17—28 x 15—22 yw, without perceptible germ-pores.

[Teleutospores. Sori similar, in densely clustered groups,
brownish-red ; spores single or in the centre of the sorus several
(46) superposed in a row, prismatic, 20—30 x 10—14yz;
epispore colourless, thin, but with an annular thickening at the
summit of the uppermost cell.]

[Acidia on leaves of Picea eacelsa, August—October ;]
uredospores on Rhododendron hirsutum, Douglas Castle, Lanark-
shire, June, 1913 (D. A. Boyd).

While this book was passing through the press, Mr D. A. Boyd kindly
communicated specimens of this most interesting find. It occurred in
small quantity, and only uredospores were observed. Description of the
other forms after De Bary. This parasite is very common in Switzerland,
in all parts where the Fir and the Alpine Roses occur together. The
teleutospores germinate in June or July and the spermogones and ecidia
appear on the Fir leaves from that time onwards, The xcidiospores can
at once infect the Rhododendron leaves, where the mycelium winters,

UREDO 385

producing its spores in the following spring. The fungus is recorded also

on R. ferruginewm and R. dahuricum, but does not attack the American
or Himalayan species.

DISTRIBUTION: Central Europe, Siberia.

Uredo Lynchii Plowr.

Trichobasis Lynchit Berk. in Gard. Chron. 1877, viii. 242. Berk. et
Br. Ann. Nat. Hist. 1878, ser. 5, i. 26. W. G. Smith in Gard.
Chron. 1885, xxiii. 693, f. 154.

Uredo Lynchii Plowr. Ured. p. 259. Sace. Syll. vii. 852.

Uredospores. Sori subepidermal, erumpent, on small pallid
spots, scattered, rarely confluent ; spores yellow, obovate, beau-
tifully echinulate, with short pedicels, 28-—35 x 20—80 u.

Fig. 288. U. Lynchii. Sori on leaf of Spiranthes, surrounded by the ruptured
epidermis, x 40; two uredospores. From the original specimens.

On a Spiranthes from Trinidad, Kew Gardens, August,
September; present on the plant when imported (R. Irwin
Lynch). On living leaves of Spiranthes in hothouse, Kelvinside,
Glasgow, September, 1890 (D. A. Boyd). Specimens in Herb.
Kew. (Fig. 288.)

Uredo Plantaginis B. et Br.
Uredo Plantaginis B. et Br. Ann. Nat. Hist. ser. 5, vii. 1380 (1881).
Plowr. Ured. p. 259. Sace. Syll. vii. 857.

Uredospores. Sori epiphyllous, on round yellowish spots
3—6 mm. wide, erumpent, surrounded by the epidermis; spores
roundish to obovate, faintly echinulate with sharp but distant
spines, yellowish, varying from 19 x 20 w up to 27 x 16 yw, with
(apparently) two or three equatorial germ-pores.

On Plantago lanceolata(?), P. major. Very rare. (Fig. 289.)

“@. vu. 25

386 UREDO

Plowright mentions this with doubt; Berkeley and Broome (/.¢.) record
it as found on Plantago at Wood Newton, and
on P. lanceolata at Dolgelly (Ralfs). In the
Trans. Worcester. Nat. Club, 1910, p. 291, it is
recorded at Pirton Pool. These I have not seen.
Plowright suggests that the Dolgelly specimen
may be a Synchyirium, and some of those I have

Fig. 289. U.Plantaginis. seen in herbaria under the name U. Plantaginis
Uredospores, on PP. are not Uredines. But in the British Museum
Wight a ale of there is one on P. major, collected in the Isle of

Wight by J. F. Rayner, October, 1907, which
appears to belong to this class; the foregoing description is taken from
this specimen, the sori lie in the centre of thickened yellow spots which

are sometimes confluent. .

Uredo Tropezoli Desm.
Uredo Tropacolt Desm. Ann. Sci. Nat. 1836, vi. 243. Cooke, Journ,
Bot. iv. 97; Handb. p. 528; Micr. Fung. p. 216. Plowr. Ured.
p. 258. Sacc. Syll. vii. 862. :

Uredospores. Sori hypophyllous, on pale-yellow _ spots,
minute, roundish, scattered or confluent; spores subglobose
to ovoid, orange-yellow, 20 w diam.

On leaves of Tropagolum aduncum (= T. canartense). Very
rare; Shere, near Guildford, October, 1865 (Dr Capron).

This species has been found only once in England. Desmaziéres who
records it on Tropacolum minus, in August, says that it causes, on the

upper face of the leaf, numerous irregular spots, barely 1 mm. across.
It has been recorded from France and Belgium.

4&cidium Hellebori Fischer.
4%, Hellebori Fischer, Ured. Schweiz, p. 526.

Spermogones. Epiphyllous, or hypophyllous amongst the
ecidia, immersed, with projecting paraphyses, about 185—150 wu
diam.

Aicidiospores. AKcidia hypophyllous, crowded, in roundish
groups, cup-shaped, with a torn spreading margin; spores
densely verruculose, 18—24 x 18—21 py.

On Helleborus viridis. May.

Included with all reserve. Winter mentions an ecidium on Helleborus
foetidus (Pilze, p. 269), which he places under .£e. Ranunculacearum.

ACIDIUM 387

Rev. C. Wolley-Dod records (Journ. Roy. Hort. Soc. xii. p. liii.) an wcidium
allied to 4c. Ficariae on his Hellebores at Malpas. I have not met
with any other reference to an ecidium on that genus. The whole of the

description is taken from Fischer, who adds that it belongs presumably to
a hetercecious species.

4écidium Ranunculacearum DC. var. Lingue.

4. Ranunculacearum DC. Flor. fr. vi. 97 p.p. Grev. Flor. Edin.
p. 446. Plowr. Ured. p. 266.

4écidiospores. Aicidia hypophyllous, in roundish or elongated
clusters of various sizes, cup-shaped, whitish, with a brittle
margin; spores polygonal, orange-yellow, 17—28 x 14—20 u.

On Ranunculus Lingua. Very rare; Duddingston Loch
(Dr Greville).

iT
It was at one time suggested by the brothers Sydow that this acidium
belonged to their Puccinta Calamagrostidis on Calamagrostis neglecta, but
they have since seen reason to doubt the truth of this idea.

7&cidium Poterii Cooke.

Aicidium Poterii Cooke, Journ. Bot. ii. 39, pl. 14, f. 3; Handb.
p- 540; Micr. Fung. p. 198. Plowr. Ured. p. 268. See also
Berkeley, Eng. Fl. v. 373.

Aicidiospores. AXcidia hypophyllous, in subrotund or elon-
‘gated clusters on the nerves of the leaves, also upon the
petioles, scattered or circinate, immersed, edges torn into minute
fugacious teeth ; spores oval, yellowish.

On Poterium Sanguisorba. May and June. A species of
which nothing else seems to be known.

ZEcidium Dracontii Schweinitz.

eidium Dracontii Schwein. Trans. Amer. Phil. Soc. 1834. Cooke,
Handb. p. 538; Micr. Fung. p. 200. Plowr. Ured. p. 266. Sace.
Syll. vii. 831.
4icidiospores. AKcidia on extensive pallid spots on the
leaves, sometimes almost covering them, arranged without
order, elongate; spores orange.

25—2

388 CHOMA

On Arum triphyllum. In gardens, Melbury, 1863 (Rev.
M. J. Berkeley), An introduced species, from North America.

Czeoma Ari-italici Rud.

Uredo Ari-ttatict Requien in Duby, Bot. Gall. ii. 899.

Caeoma Ari Winter Pilze, p. 256.

C. Ari-ttalicti Rud. in Linnea, iv. 512. Sacc. Syll. Fung. vii. 868.
Trans. Brit. Myc. Soc. i. 60.

Aicidiospores. Czomata hypophyllous, irregular, flattened,
scattered or concentric, often confluent, without a peridium,
orange-yellow ; spores round or elliptic, often somewhat angular,
verrucose, orange, 15—30 x 15—20 p.

On Arum maculatum. Very rare; near Salisbury, April,
1897 (Mr E. J. Tatum). Found also in France and Germany.

Requien’s description is as follows: Uredo hypophylla, maculis latis
lutescenti-albidis sparsis, acervulis rufis, annulatim digestis, orbicularibus
ovatisve, compactis, planis, epidermide rupta cinctis, sporidiis dilute rufis,
grossis, pellucidis, subglobosis.

Phragmidium tuberculatum J. Miiller.

This species is recorded for Britain by Sydow (Monographia,
ili. 114), on the ground that it was distributed by Baxter,
“Stirp. Crypt. Oxon.” no. 37. There are specimens of this
exsiccatum both at Kew and in the British Museum, but in
both these the fungus on Rose-leaves is typical Ph. disciflorum,
almost all the teleutospores having six septa, not 8—5 as in
Ph. tuberculatum. The latter species, being widely distributed
in Europe, is likely to be found here, but the evidence of its
occurrence is at present insufficient.

There is in Herb. Kew an ecidium on “Atriplex littoralis,
Maldon, Essex, M. A. Irvine,” June Ist, 1864, Alcidia hypo-
phyilous, covering' the whole leaf, densely crowded, shortly
cylindrical, with a slightly torn margin. Of this nothing else
seems to be known.

EXCLUDED SPECIES 389

EXCLUDED SPECIES.

Uromycres Parwassi@ Cooke, Grevillea, vii. 134. Plowr. Ured.
p. 128.

This species does not exist. The ecidium on Parnassia,
which was at first assigned to
it, is now known to belong to
Puceinia uliginosa (q.v.), and the
other stages are merely Uromyces
Valerianae on Valeriana dioica,
as has been pointed out by Sydow.
The leaves issued by Cooke under
this name (Exsicc. 1. 74) are
obviously the radical leaves of
the Valerian (Fig. 290) which Fig. 290. Radical leaves of Va-
grows in similar places to those —_—‘leriana dioica (left) and Parnassia
suitable for the Parnassia. In ee MENDES
Sutton Park (Warwicks.) the two plants grow side by side, and
when I first found the Uromyces there I put it down as JU.
Parnassiae. Rusty marks do frequently appear on the leaves of
the Parnassia which are placed in herbaria as U. Parnassiae,
but on examination no spores will be found in them.

Uromyces Urntica Cooke, Grevillea, vii. 137. Plowr. Ured. p. 142.

This also is a non-existent species. The origin of the
mistake can now never be unravelled.

URoOMYCES SCUTELLATUS Cooke, Grevillea, vii. 137. Plowr. Ured. p. 134,

This is probably nothing but an error in identification.

PuccINIA ASARINA Cooke, Handb. p. 504, Plowr. Ured. p. 202.

This is merely Puccinia Fergussont; the leaves on which
the specimens are in Cooke’s Exsice. i. 110 are obviously those
of Viola palustris. It is rather strange that the spores of
this species are exceedingly like the spores of the continental
P. asarina, and the sori are almost identical in appearance.

390 EXCLUDED SPECIES

PUCCINIA CLANDESTINA Carm. in Berk. Engl. Fl. v. 365, “on Scabiosa
succisa”; Plowr. Ured. p. 185.
Perhaps the host-plant was mistaken; no Puccinia on
Scabiosa is known.

XENODOCHUS cURTUS Cooke, Micr. Fung. p. 201. Plowr. Ured. p. 228.

Not a Uredine, but may be a species of Hyphomycetes.

Ectp1uM BaRBARE Cooke, Grevillea, x. 115. Plowr. Ured. p. 265.

This record depends, so far as this country is concerned, on
a single leaf which was sent to M. C. Cooke by a correspondent,
who informed him (on the authority of another person) that
the leaf belonged to Burbarea praecox, But on examination
the leaf is.seen to be one of the radical leaves of Lapsana
communis, of which it has the peculiar hairs, and the fungus is
obviously Acidium Lapsanae. The true 4. Barbareae is quite
different. The leaf is preserved in the Kew Herbarium.

/JECIDIUM INCARCERATUM B. et Br. Plowr. Ured. p. 267.

Plowright’s suspicion that this is merely Doassansia Sagit-
turiae (on Sagittaria sagittefulia) is perfectly correct.

ZECIDIUM PSEUDO-COLUMNARE J. Kiihn.

Recorded in Plowright, Ured. p. 271, on “Abies pectinata,
nordmanniana, amabilis, cephalonica. Lyme Regis, Mr Munro.”
Nothing else seems to be known of this as British ; most likely
the specimens belonged to Calyptospora Goeppertiana, or were
introduced on imported plants. 4. pseudo-columnare differs
from the ecidium of C. Goeppertiana in having oblong white
spores which are larger, more irregular in form, unevenly warted
or even smooth at the end (see Hedwigia, 1885, xxiv. 108).

UrEpo OXALIDIS.

There is a specimen bearing this name in the Herbarium of
the British Museum, on leaves of Oxalis Acetosella (W. Phillips,
July 31, 1880) from “Orton”; and a similar one, part of the
same gathering, in the Kew Herbarium, from Orton Longueville,
Hunts. (Rev. M. J. Berkeley, August, 1880). Both these are
not Uredines, but show marks probably of insect-bites.

GLOSSARY

aculeate. Covered with needle-like projections,

alveolate. Having depressions all over the surface, like a honeycomb.

amphigenous. Growing upon both sides of a leaf.

autcecious. Having all spore-forms upon the same species of host.

basipetal. Having each new part (e.g. spore) formed nearer to the base
than the similar preceding one.

brachymeiosis. A modification of meiosis in which the separation
of the chromosomes is not preceded by a- contraction of the nuclear
material,

capitate. Surmounted by a nearly globular head.

chlorenchyma. Parenchymatous tissue containing chloroplasts.

circinate. Arranged in a circle.

cuneate. Tapering downwards, with straight sides.

dendritic. Having a branched form, like a tree.

denticulate. Provided with small teeth-like projections,

digitaliform. Having the shape of a finger of a glove.

echinulate. Covered with spiny projections.

ellipsoid. Having an oval outline, rounded equally at both ends.

endokaryogamy. The intracellular fusion of nuclei after a series of
conjugate divisions.

epiphyllous. Growing on the upper side of a leaf.

erumpent. Bursting through the tissues of the host, and becoming
superficial.

fusoid. Having the shape of a shuttle, tapering at each end.

gamete. A cell specialised for reproducing the species, by fusing with
another gamete.

haustoria. Short mycelial branches which penetrate from the inter-
cellular spaces into the cells of the host, and absorb their nutritive
contents.

hetercecious. Having some spore-forms upon cne species of host and
the others upon another species of a different genus”

hypophyllous. Growing on the underside of a leaf.

infection. The successful attack of the mycelium upon the cells of the
host.

inoculation. The entry of a germ-tube into a host-plant.

392 GLOSSARY

intercalary. Occupying a position between other bodies in a row.

karyogamy. See endokaryogamy.

laciniate. Torn into a ragged form at the edge.

meiosis. A special type of nuclear division by which the number of
chromosomes in each daughter nucleus is reduced to half the number
present in the nucleus before meiosis.

metcecious. The same as hetercecious.

monophyletic. Descended from a single species or closely allied group
of species.

pallid. Of the colour of fresh chamois leather.

paraphysis. A more or less thread-like organ which grows by the side
of the spores.

plectenchyma. A kind of pseudo-parenchyma, formed by a mass of
intertwined hyphe.

polyphyletic. Descended, in distinct lines, from widely different
ancestors,

pulverulent. Having a powdery appearance from the loose spores.

pulvinate. Having the shape of a cushion.

punctate. Marked with little dots, like pin-pricks.

punctiform. Having the form of a small pin’s head.

reticulate. Covered with a network of liues.

scrobiculate. Having the surface hollowed out into little shallow pits.

semi-apogamy. A fusion of cells for reproduction, where one at least
of the fusing cells is still more or less sexually specialised, but the
cells are not of opposite sexes. If both cells represent female gametes,
it may be called parthenogamy.

trichogyne. A long hair-like projection from a female cell, suitable for
arresting a passing male cell.

truncate. As if cut off at the top, with rather square corners.

verrucose. Warted, ie. covered with rounded elevations.

verruculose. Covered with minute rounded elevations.

viable. Able to put forth a germ-tube.

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INDEX OF HOST-PLANTS MENTIONED

Abies alba=A. pectinata Anthyllis Vulneraria 95
amabilis 390 Apium graveolens 185
cephalonica 390 Aquilegia vulgaris 276
Nordmanniana 3873, 390 Arctium Lappa 138
pectivata 361, 363, 367, 373, 390 Arenaria trinervis 219

Achillea Armeria maritima 89
Millefolium 131 Arrhenatherum elatius 251, 256, 285
Ptarmica 131 Artemisia

Adiantum Capillus-Veneris 378 Absinthium 1384

Adoxa Muschatellina 161, 162, 205 campestris 134

Aigopodium Podagraria 185 maritima 134

AAthusa Cynapium 190 vulgaris 134

Agrimonia Eupatoria 365 Arum

Agropyron 282 maculatum 269, 388
caninum 251, 259, 263 triphyllum 388
repens 251, 254, 259, 263, 283 Asparagus officinalis 234

Agrostis Asperula odorata 164
alba 251, 254, 276 Aspidium spinulosum 378
canina 251 Aster Tripolium 130, 248
stolonifera 254 Atriplex littoralis 388
vulgaris 251, 254, 276 Avena

Aira elatior (see Arrhenatherum)
cespitosa 251, 265 fatua 251, 256
flexuosa 251, 265 pratensis 256

Ajuga reptans 172 sativa 251, 256

Alchemilla vulgaris 106

Allium Barbarea preecox 390
Cepa 236, 345, 348 Bartsia Odontites 326
Schenoprasum 122, 236 Bellis perennis 237
Scorodoprasum 122, 236 Berberis vulgaris 251, 285
‘spherocephalum 122 Beta
ursinum 268, 345, 346, 348 maritima 113

Alopecurus pratensis 251, 256, 271 vulgaris 113

Alsine rubra 111 Betonica officinalis 174

Althwa rosea 207 Betula alba. 359

Amygdalus communis 209 Blechnum Spicant 377

Anchusa arvensis 261 Brachypodium \

Anemone pinnatum 251, 281
coronaria 209 silvaticum 255, 259, 281
decapetala 210 Bromus 262
nemorosa, 209, 216, 330 arvensis 251
ranunculoides 209 asper 251

Angelica silvestris 193, 225 erectus 251

Anthoxanthum odoratum 270 giganteus. 251

Anthriscus silvestris 195 mollis 251, 259

INDEX OF HOST-PLANTS MENTIONED 399

Bromus (cont.)
secalinus 251
sterilis 262

Bunium
Bulbocastanum 186
flexuosum 187

Bupleurum
faleatum 190
tenuissimum 189

Buxus sempervirens 206

Cacalia
hastata 325
suaveolens 325

Calamagrostis lanceolata 254

Calamintha
Clinopéddium 171
officinalis 171

Caltha palustris 217, 218

Campanula

glomerata 329
latifolia 329
Rapunculus 160
rotundifolia 160, 329
Trachelium 329
turbinata 329
Carduus

arvensis 145
crispus 141, 144
nutans 141
pyenocephalus 142
Carex

acuta 243
acutiformis 242, 243

arenaria 246, 247, 248

binervis 242
exspitosa 243
earyophyllea 246
Davalliana 244
dioica 244
extensa 248

fulva 249
Goodenovii 243, 249
hirta 242
montana 133
muricata 152
paludosa 242, 248
panicea 249
pendula 242
precox 246
Pseudocyperus 242
remota 245
viparia 242, 243
stricta 243, 249
vulgaris 249, 250
Carlina vulgaris 138
Carpinus Betulus 360
Carum
Bulbocastanum 186
Carui 225

Carum (cont.)

majus 187
Cattleya Dowiana 381
Centaurea

Cyanus 141

montana 141

wnigra 140, 248
“Scabiosa 140
Cetastium
.,arvense 361
‘- triviale 361

viscosum 361
Cherophyllum temulum 195
‘Chelidonium majus 353
Chrysanthemum

indicim 132

Leucanthemum 133

sinense’ 132
Chrysosplenium

alternifolium 214

oppositifolium 214
Cichorium Intybus 148
Cicuta virosa 184
Cineraria 321

Circea

alpina 198

intermedia 366

lutetiana 198, 366
Cirsium

arvense 145

heterophyllum 146

lanceolatum 143, 144

palustre 1438, 244

pratense 143, 244
Clematis Vitalba 282
Cnicus (see Cirsium)
Colchicum speciosum 122
Conium maculatum 196
Conopodium denudatum 187, 225
Convallaria majalis 267
Convolvulus sepium 176
Corydalis 353
Cotoneaster integerrimus 307
Cotyledon Umbilicus) 211
Crateegus

monogyna 306, 307 j

Oxyacantha 306, 307

Crepis
paludosa 158
virens 157

Cydonia vulgaris 306, 307
Cystopteris fragilis 376

Dactylis glomerata 126, 251, 254
Deschampsia ‘(see Aira)
Dianthus : ;
barbatus 109, 219
Caryophyllus 109
chinensis 109
Digraphis arundinacea 266

400 INDEX OF HOST-PLANTS MENTIONED

Echium vulgare 265 Gymnadenia conopsea 344
Elymus arenarius 259 Gypsophila elegans 219
Empetrum nigrum 311
Endymion non-seriptum 121 Helleborus
Epidendrum vitellinum 383 foetidus 386
Epilobium viridis 386
angustifolium 367 Hepatica acutiloba 209
hirsutum 199 Heracleum Sphondylium 194
montanum 199 Hieracium
palustre 200, 367 boreale 159
tetragonum 199 murorum 159
Eranthis hiemalis 209 Pilosella 159
Ervum hirsutum 96 umbellatum 159
Euonymus europeus 340 Holcus
Euphorbia lanatus 251, 254, 256, 263
amygdaloides 334 mollis 251, 254, 256, 263
Cyparissias 93, 95, 100 Hordeum
exigua 108, 354 murinum 251, 259
Helioscopia 354 pratense 251
Peplus 354 vulgare 251, 259, 265
silvatica 334 Hydrocotyle vulgaris 182
Euphrasia officinalis 326 Hypericum
Androsemum 355
Faba vulgaris 97 humifusum 355
Festuca montanum 355
duriuscula 257 perforatum 355
elatior 256, 284 pulchrum 355
Myurus 251 Hypocheris radicata 149
ovina 251, 257
rubra 251 Impatiens
silvatica 254 fulva 205
Fragaria vesca 290 Noli-tangere 205
Fritillaria Meleagris 119 Iris Y
feetidissima 231
Gagea lutea 120 Pseudacorus 231
Galanthus nivalis 345
Galium Jasione montana 160
Aparine 168 Jonquil 232
Cruciata 165, 167 Juucus obtusiflorus 123
Mollugo 165 Juniperus
palustre 165 communis 306, 308
saxatile 165, 167 Sabina 307, 309
uliginosum 165
verum 165, 370 Koeleria cristata 286
Gentiana acaulis 179
Geranium Lactuca muralis 152
dissectum 104 Lapsana communis 147
molle 104, 105, 228, 229 Larix europrea 339, 341, 349, 350, 359
pheum 228 Lathyrus
pratense 104, 228 macrorrhizus 99
pusillum 104, 105, 229 montanus 99
pyrenaicum 104, 105 pratensis 90, 97, 100
rotundifolium 104, 229 vernus 98
silvaticum 104, 228 Leontodon
Glaux maritima 125 autumnalis 150
Glechoma hederacea 174 hirtus (see Additions, p. xii)
Glyceria hispidus 150
aquatica 256 Lilium candidum 119
fluitans 279 Limnanthemum nymphmoides 240

Goodyera repens 344 Limonium (sve Statice)

INDEX OF HOST-PLANTS MENTIONED 401

Linum
catharticum 356
usitatissimum 356

Listera
cordata 344
ovata 344

Lolium perenne 251, 256
Lonicera Periclymenum 257
Lotus

angustissimus 95

corniculatus 95
Luzula

campestris 237, 239

maxima 239

pilosa 239

silvatica 237 and p. xii
Lycaste Skinneri 383
Lychnis

diurna (dioica) 219, 222

vespertina 219
Lycopsis arvensis 261

Mahonia Aquifolium 251
Malva silvestris 207
Malvesr 207

Mangels 113

Medicago 93
Melampyrum 277
arvense 327

pratense 327

Mentha

aquatica 171

arvensis 171

citrata 171
rotundifolia 171
silvestris 171

viridis 171
Mercurialis perennis 352
Mespilus germanica 306, 307
Milium effusum 251
Molinia cerulea 251, 277
Myrrhis odorata 195

Narcissus

poeticus 232
Pseudonarcissus 232
Nepeta Glechoma 174
Nymphoides peltatum 240

Oncidium

Forbesii 383

varicosum 383

Orchis

latifolia 268, 344

maculata 344

Origanum vulgare 171
Ornithogalum umbellatum 235
Orobus tuberosus 99

Oxalis Acetosella 390

Oxyria digyna (reniformis) 224

G. U.

Peonia officinalis 314
Paris quadrifolia 267
Parnassia palustris 250, 389
Pedicularis palustris 249
Petasites officinalis 324
Petroselinum sativum 190
Peucedanum palustre 193
Phajus Wallichii 382
Phalaris

arundinacea 254, 266

canariensis 251
Phaseolus vulgaris 102

Phillyrea
latifolia 332
media 333

Phleum

nodosum 284

pratense 284
Phragmites communis 272, 274, 275
Picea excelsa 313, 369

Pimpinella
magna 188
Saxifraga 188

Pinus
austriaca 321
Cembra 317

montana 325
monticola 317
silvestris 314, 321, 323, 324, 325,

326, 327, 329, 351
Strobus 317

Pisum sativum 97, 100
Plantago

lanceolata 385

major 385

Poa

annua 127, 279
aquatica 256
compressa 251
fluitans 279
nemoralis 1427, 251, 279
pratensis 127, 251, 279
trivialis 127, 251, 279
Polygonum

amphibium 228
aviculare 117

Bistorta 225
Convolvulus 229
lapathifolium 228
viviparum 226
Polypodium

Dryopteris 375
Phegopteris 380
Robertianum 375
vulgare 877

Populus

alba 350, 351, 352
balsamifera var. 348, 349, 350
canadensis 349

canescens 351

: 26

402 INDEX OF HOST-PLANTS MENTIONED

Populus (cont.)
nigra (serotina) 348, 349
ontariensis (candicans) 349
pyramidalis (italica) 349
tremula 350, 351, 352
Potentilla
argentea 292
erecta 302
Fragariastrum 290
sterilis 290
Tormentilla 302
verna 292
Poterium Sanguisorba 293, 387
Primula
acaulis (vulgaris) 180
elatior 180
veris 180
Prunella vulgaris 174, 278
Prunus
domestica 209
insititia 209
Padus 369
spinosa 209
Pulicaria dysenterica 123
Pulmonaria montana 262
Pyrola
minor 312, 368
rotundifolia 312, 368
Pyrus
Aucuparia 308, 330
communis 306, 307, 308
torminalis, etc. 331

Quercus pedunculata 316

Ranuneulus
acris 126, 271
auricomus 127
bulbosus 126, 127, 272
Ficaria 107, 115, 127
Lingua 387
repens 126, 127, 272
Rhamnus
catharticus 209, 256
Frangula 254
Rheum officinale 274
Rhinanthus Crista-galli 326
Rhododendron
dahuricum 385
ferrugineum 3885
hirsutum 384
Ribes
alpinum 343
Grossularia 248, 342, 343
nigrum 243, 317, 342
rubrum 212, 317, 342
Rosa
alpina 295
canina 294, 388
spinosissima 294

Rubus
cesius 297 and p. xii
fruticosus 296, 297, 301 and p. xii
Ideus 299
saxatilis 297

Rumex
Acetosa 116, 223, 275
Acetosella 116, 118
acutus 274
conglomeratus 115, 274
crispus 115, 274 |
Hydrolapathum 115, 274
nemorosus 115
obtusifolius 115, 274

Sagina

nodosa 219
procumbens 221
Sagittaria sagittifolia 390
Salicornia

europea 114

herbacea 114
Salix

alba 346

aurita 339, 340, 341, 344
Caprea 339, 340, 341
cinerea, 340, 341
fragilis 341, 345
herbacea 346
nigricans 341
pentandra 345
purpurea 341, 343
repens 344

reticulata 341
Smithiana 341
viminalis 341, 342
Sanguisorba officinalis 303
Sanicula europea 183
Saponaria ocymoides 109
Saxifraga

Aizoon 213

Cotyledon 213

elatior 214

granulata 213, 357
longifolia 213
stellaris 213

umbrosa 213

Scabiosa 390

Scilla

bifolia 121
campanulata 121
nutans 121

Scirpus

lacustris 240
maritimus 125
Scolopendrium vulgare 378
Scrophularia

aquatica 87

nodosa 87

Secale cereale 252, 259, 261

INDEX OF HOST-PLANTS MENTIONED 403

Sedum

acre 287

reflexum 287
Bhodiola 211
roseum 211
Sempervivum
arachnoideum 3835
calcareum 835
globiferum 335
montanum 335
tectorum 335
Senecio

aquaticus 136
Jacobwa 136, 187, 247, 321
palustris 321
pulcher 321
sarracenicus 321
silvaticus 321
viscosus 321

vulgaris 321
Serratula tinctoria 139
Sherardia 370

Silaus pratensis 191
Silene

Cucubalus 110, 222
inflata 110, 222
latifolia 110, 222
maritima 110
Smyrnium Olusatrum 197
Soldanella alpina 181
Solidago Virgaurea 130
Sonchus

arvensis 156, 325
asper 156, 325
oleraceus 156, 325
Spergula arvensis 219, 220
Spergularia rubra 111
Spirea

Filipendula 288
Ulmaria 288
Spiranthes 385
Stachys Betonica 174
Statice Limonium 88
Stellaria

graminea 361
Holostea 219
media 219, 361
uliginosa 219
Sueeda

fruticosa 112
maritima 112°
Symphytum officinale 262, 363

Tanacetum vulgare 135
Taraxacum officinale 153, 154, 245
Teucrium

Chamedrys 175

Scorodonia 175

Thalictrum 209

flavum 215, 283

minus 215, 283

Thesium humifusum 230
Thrincia hirta (see Additions, p. xii)
Thymus Serpyllum 173
Tragopogon pratensis 151
Trifolium

agrarium 93

hybridum 90

incarnatum 90

medium 90

minus 93

pratense 90

procumbens 93

repens 91, 92

Trisetum fiavescens 252, 264
Triticum (see also Agropyron)
-glaucum 282

junceum 282

vulgare 252, 259, 263
Tropzolum

aduncum 386

canariense 3886

minus 3886 ;
Tussilago Farfara 279, 323

Urtica dioica 242, 389

Vaccinium
Myrtillus 371, 373
uliginosum 3871
Vitis-idea 371, 373
Valeriana
dioica 86, 389
officinalis 86
Veronica
alpina 170
montana 169
officinalis 170.

Vicia
Cracca 97
hirsuta 96
sativa 97
sepium 97
Vinca
major 177
minor 177
Viola

canina 201
cornuta 203

hirta 201

lutea 203

odorata 201
palustris 204, 389
Riviniana 201
silvestris 201
tricolor 203

GENERAL INDEX

Synonyms and species incidentally mentioned are in italics. When more than
one page is given, the one on which the description of the species occurs is

printed in heavy type..

eecidiospores 4

ecidium 38, 30

“ECIDIUM 386
abietinum A, et 8. 384
Adoxae Opiz 160
albescens Grev. 162
Allii Grev. 268
Anchusae EH, et H. 261
Aquilegiae Pers. 275
argentatum Schultz 204
Ari Desm. 269
Asperifolit Pers. 261
aviculariae Kze. 117
Barbareae Cooke 390
Behenis DC. 109, 222
Berberidis Gmel. 250
Bulbocastanit Cum. 186
Bunii DC. 186, 188
Bunii var. Smyrnii-Olusatri DC. 197
Calthae Grev. 216
Cathartici Schum. 255
Chenopodit Duby 111
Cirstt DO. 244
Clematidis DC. 281
columnare A. et 8. 372
Compositarum var, Bellidis DC. 236
Compositarum var. Jacobaeae Cooke

Compositarum var. Prenanthis
Cooke 157

Compositarum var. Tussilaginis
Cooke 278

conorum-Pices Reess 313

Convallariae Schum. 267

crassum Pers. 253, 255

crassum var, Ficariae Pers. 127

crassum var, Periclyment Cooke 257

crassum var. Phillyreae Cooke 332

Cyparissiae DC. 99

depauperans Vize 202

Dracontii Schw. 387

elatinum A. et 8S. 360

Fpilobii DC. 198

Eervi Wallr. 96

Euphorbiae Gmel, 102

Euphorbiae Pers. 333

ECIDIUM (cont.)

Euphorbiae-silvaticae DC. 333

Falcariae var. Bupleuri-faleati DC.
189

Frangulae Schum. 253

fuscum Pers. 215

Galii A. et S. 164

Gentianae Jacz. 178

Geranii DC. 103, 227

Glaucis D. et M. 124

graveolens Shut. 252, 284

Grevillei Grove 152

Grossulariae_DC. 242

Hellebori Fisch. 386

incarceratum B, et Br. 390

Jacobaeae Grev. 246

Lapsanae Schultz 147

leucospermum DC, 329

Aelampyri K. et 8. 276

Menthae DC. 170

Mespili DC. 306

Nymphoidis DC. 239

Orchidearum Desm. 268

ornithogaleum Bub, 235

Orobi Pers. 99

Parnassiae Grav. 249

Pedicularis Libosch 248

Periclyment Schum. 257

Phaseolorum Wallr. 101

Phillyreae DC. 332

Pimpinellae Kirech. 188

Poterii Cooke 387

Prenanthis Pers. 151

Primulae DC. 179

pseudo-columnare Plowr. 372, 390

Pulmonariae Thiim. 262

punctatum Pers. 207, 331

Pyrolae DC. 312

Pyrolae Gmel. 367

quadrifidum DC. 207

Ranunculacearum DC. 125, 127, 271

Ranunculacearum var. Aquilegiae
DO. 275

Ranunculacearum var. Clematidis
Cooke 281

Ranunculacearum var. Lingue 387

GENERAL INDEX

ABCIDIUM (cont.)

Ranunculacearum vay.
Cooke 282

Ranunculi-acris Pers. 125
rubellum Gmel. 273, 274
Salicorniae DC. 114
sanguinolentum Lindr.
Saniculae Carm. 182
Scrophulariae DC, 87
Senecionis Fisch. 246
Soldanellae Horn. 180
Sonchi Johnst. 155
Statices Desm. 88
strobilinum Wint. 368
Suaedae Thiim. 111
Symphyti Thiim, 262
Tarazaci K. et 8. 245
Thesit Desv. 229
Tragopogi Pers. 150
Tragopogonis Cooke 150
Tussilaginis Gmel. 278
Urticae Schum. 241
Valerianearum Duby 86
Violae Schum. 200
zonale Duby 123

alternation of generations 27

104, 227

amphispores 11, 34
ascogonium 22
Ascomycetes 77, 80

Ascophora disciflora Tode 293
Ascospora Scolopendrit Fekl. 378
azygospores 23, 31

Barclayella 14, 80

basal cells 18

Basidiomycetes 29

basidiospores 14, 37

basidiospores, germination of 15, 38

basidium 14, 37

basidium, internal 28, 39, 77, 318,
329, 332

Blackman 17

Blackman and Fraser 18

Blackman and Welsford 25

brachymeiosis 82

bridging-species 65, 66, 71

COMA 30, 388
Alliorum Link 269, 344, 347
Avi-italici Rud, 388
Armeriae Schl. 89
Euonymi Plowr. 339
Galiit Link 370
Hydrocotyles Link 181
Laricis Plowr. 838, 348, 349
Lilii Link 118
Mercurialis Link 351
dblongatum Link 238
obtegens Link 145
Orchidis Wint. 268, 343

Thalictri

405

CAZ0MA (cont.)
pinitorquum A. Br. 350
Ribesit Link 342
Saxifragae Wint. 213, 357
CALYPTOSPORA 74, 372
columnaris Kiithn 372
Goeppertiana Kiihn 59, 363, 364,
367, 372 |
Christman 18°
CHRYSOMYXA 75, 310
albida Kiihn 300
Empetri Pers. 311
Pyrola Rost. 3812
Rhododendri De By.
Woronini Tranz. 311
classification 73
Claussen 81
Coleosporiacen 73, 75, 318
COLEOSPORIUM 75, 319
Cacalie Fekl. 325
Campanule Lév. 328
‘Euphrasie Wint. 326
Melampyri Karst. 827
miniatum Pers. 293
ochraceum Fekl. 364
Petasitis Lév. 3823
pingue Lév. 293
Rhinanthacearum Lév. 326
Senecionis Fr. 320
Sonchi Lév. 156, 322, 323, 324, 325
Symphyti 363
Tussilaginis Tul. 322
Collemaces 25, 77
conjugate division 5
Cronartiaces 78, 75, 310
CRONARTIUM 175, 813
asclepiadeum Fr, 313
asclepiadeum var. Quercuum Berk.
315
flaccidum Wint, 314
Paeoniae Cast. _ 313
Quercus Arth. 315
Quercuum Miy. 315
tibicola F. de W. 55, 316

311, 384

Darluca Filum Cast.
dikaryon 5
Dittschlag 19
Doassansia Sagittariae Fisch. 390

33, 178

Endophyllacee 73, 75, 333
ENDOPHYLLUM 75, 333
Euphorbiae Plowr. 333
Euphorbiw-silvatice Wint. 333
leucospermum Sopp. 331
Sedi Lév. 287
Sempervivi De By. 53, 336
Epitea Baryi B. et Br, 280

Fairy Ring of Carnations 109
26—38

406

Fromme 20

fusion of nuclei 12, 28

gametophyte 27

germ-pores 7, 76

germ-tube 8

GYMNOSPORANGIUM 75, 304
bermudianum Earle 304
clavarieforme DC. 51, 304
confusum Plowr. 306
cornutum Arth. 307
Juniperi Link 307
juniperinum Fr. 307
Sabine Wint. 308

haustoria 8

HEMILEIA 381
americana Mass. 381
Oncidii G. et M. 383
Phaji Syd. 382

heterceecism 16

Heterosporium 109

heterotype mitosis 26, 28

Hoffmann 20

HYALOPSORA 74, 373
Adianti-Capilli-Veneris Syd. 378
Aspidiotus Magn. 374
Polypodii Magn. 375
Polypodii-Dryopteridis Magn. 374

immunity 71
inoculation 63
intercalary cell 6, 10
intercalation 5

KUEHNEOLA 75, 299
albida Magn. 300
obtusa Arth. 301
Tormentille Arth. 301
Uredinis Arth. 300

Kurssanow 18

Laboulbenia 77, 81

lactic acid, action of 84, 289, 291,
297, 298

LECYTHEA
Baryi Berk. 280
epitea Lév. 340
Euphorbiae Lév.
gyrosa Berk, 298
Lini Berk. 356
Poteriit Lév. 292
Rosae Lév. 293
Ruborum Lév. 295

Licea strobilina A, et S. 368

Lutman 82

Lycoperdon caryophyllinum Schr. 108

353

manipulation 84
Melampsoracene 73, 74, 336

GENERAL INDEX

MELAMPSORA 74, 336
aecidioides Plowr, 351
Allii-fragilis Kleb. 344, 346
Allii-populina Kleb. 346, 347
Allii-Salicis-albe Kleb. 345
arctica Rost, 346
betulina Desm. 358
Cerastit Wint. 361
Circaeae Schrét. 365
elatina Arth. 361
epitea Thiim, 340
HKuonymi-Caprearum Kleb. 339
Euphorbie Cast. 353
farinosa Schrét. 338
Galii Wint. 370
Helioscopiae Wint. 353
Hypericorum Wint. 354
Klebahni Bub. 350, 353
Larici-Caprearum Kleb. 338
Layrici-epitea Fisch. 340, 343
Larici-pentandrae Kleb. 345
Larici-populina -Kleb. 348
Larici-tremule Tul. 349
Lini Desm. 355
Tiniperda Korn. 356
Magnusiana Wagn. 350, 353
mixta Plowr. 343
Orchidi-repentis Kleb. 343
Padi Cooke 369
pinitorqua Rost.
populina Lév. 347, 348
pustulata Schrot. 366
Pyrolae Schrot. 367
Quercus Schrét. 315
Ribesii-auritae Kleb. 343
Ribesii-purpurer Kleb. 342

57, 350

Ribesii-viminalis Kleb. 342
Rostrupii Wagn. 351
salicina Lévy. 3388
Saxifragarum Schrét. 357
Tremulm Tul. 349
Vacciniorum Schrét. 371

vernalis Niessl 357
Vitellinae Plowr. 344
MELAMPSORELLA 74, 360
Aspidiotus Magn. 374

Blechni Syd. 377
Caryophyllacearum Schrit.
360, 364, 367
Cerastit Schrét. 361
Dieteliana Syd. 376
Symphyti Bub. 363
MELAMPSORIDIUM 74, 358
betulinum Kleb. 358
Carpini Fisch. 360
MELAMPSOROPSIS
Empetri Arth. 311
Pyrolae Arth. 312
mesospores 13
Milesia Polypodii White 376, 377, 378

156,

GENERAL INDEX 407

MILESINA 74, 376
Blechni Syd, 362, 377
Dieteliana Magn. 376
Kriegeriana Magn. 378

mycoplasm 43, 44, 49, 72

NIGREDO
appendiculata Arth. 101
Beta Arth. 113
caryophyllina Arth. 108
fallens Arth. 90
Junci Arth. 123
Liltt Arth. 118
Scirpi Arth. 124
Trifolii Arth. 91
nuclear division 25

OCHROPSORA
Sorbi Diet.
Olive 18

75, 329

Perichaena strobilina Fr. 368
PERIDERMIUM 31
acicolum Link 320
Boudiert Fisch. 323
columnare Cooke 372
conorum-Piceae 313
Cornui Kleb. 314
deformans Tub. 315
elatinum Link 360
Fischert Kleb. 324
giganteum Tub. 315
Laricis Kleb. 359
Magnusianum Fisch. 325
Magnusit Wagn,. 325
oblongisporium Fekl. 328
Pini Chev. 320
Plowrightii Kleb. 322
Rostrupii Fisch. 328
Soraueri Kleb. 327
Stahlii Kleb. 326
Strobi Kleb, 316, 359
peridium 3, 6, 30, 79
Phelonitis strobilina Pers. 368
PHRAGMIDIUM 75, 289
acuminatum Fr. 292
albidum Lagh. 300
bulbosum Schl. 295
bullatum West. 293
earbonarium Wint. 303
disciflorum James 293
Fragariastri Schrét. 290
fusiforme Schrét. 294
gracile Cooke 298
imitans Arth. 299
mucronatum Fr, 293
obtusum Link 290
Potentille Karst. 291
Potentillae-canadensis Diet. 301
Rosae-alpinae Wint. 294

PHRAGMIDIUM (cont.)
Rubi Wint. 297
Rubi-Idei Karst. 298
Rubi-saxatilis Liro 297
Sanguisorbe Schriét, 292
subcorticium Wint, 293
Tormentillae Fckl. 301
tuberculatum Mull. 388
violaceum Wint. 295
PHYLLACHORA
graminis 127
Ulmi 349
phylogeny 75
plectenchyma 2
PODISOMA
Juniperi Fr. 304
_ Sabinae Fr, 308
promycelium 14
Protomyces filicinus Niessl 379
Pseudopeziza Trifolit Fckl. 92
PUCCINIA 75, 128
Absinthii DC. 134
Acetose Kirn. 223
Actaeae-Agropyri Fisch. 283
acuminata Fekl. 167
Adoxe Hedw. f. 160, 162
Acidii-Leucanthemi Fisch. 133
Aicidii-Melampyri Liro 276
figopodii Mart. 185
egra Grove 202
Aithuse Mart. 190
Agropyri HE. et E. 281
agropyrina Eriks. 263
Agyrostidis Plowr. 275
albescens Plowr. 161, 162, 205
Allii-Phalaridis Kleb. 268
ambigua Lagh, 168
Andersoni B. et Br. 146
Anemones Pers. 215
Anemones var. Betonicae A. et S.
174
Angelic Fekl. 192
Angelicae-Bistortae Kleb. 225
annularis Schl. 175
Anthoxanthi Fekl. 269
Apii Desm. 184
Arenariae Wint. 218, 221
arenariicola Plowr. 247
argentata Wint. 161, 163, 204
Ari-Phalaridis Kleb. 269
Arrhenatheri Eriks, 284
arundinacea Hedw. 273
asarina Cooke 204, 389
Asparagi DC. 233
Asperulae Fekl. 163
Asperule-odorate Wurth 163
Asteris Duby 129
Asteris var. Chrysanthemi-Leucan-
themi Massal. 133
Bardane Cord. 138

408 GENERAL INDEX

PUCCINIA (cont.)

PUCCINIA (cont.)

Baryi Wint. 280

Behenis Otth 220, 222

Betonicw DC. 174

Bistortae DC. 225, 226

bromina Eriks. 262

bromina, specialisation of 70
Brunellarum-Moliniae Cruch. 277
Bulbocastani Fckl. 186

bullaria Link 193, 196

bullata Wint. 190, 191, 193, 196

Bunit Wint. 187

Bupleuri Rud. 189
Bupleuri-falcati Wint. 189
Buxi DC. 205

Calthe Link 216
Campanula Carm. 159, 329
Cardui Plowr. 144
Cardui-pyenocephali Syd. 142
Carduorum Jacky 141
Cari-Bistortae Kleb. 225
Caricis Reb. 241

Caricis (life-history) 1
Carline Jacky 137
caulincola Schn. 172
Celakovskyana Bub. 166
Centaurew DC. 139
Cherophylli Purt. 195
Chondrille Cord. 161
Chrysanthemi Roze 131
Chrysanthemi-chinensis Henn. 131
Chrysosplenii Grev. 214
Cichorii Bell. 148

Cicutss Lasch 183

Circeerr Pers. 198, 365, 366
Cirsii Lasch 142
Cirsii-lanceolati Schrét. 143
clandestina Carm. 390

a Clinopodit DC. 171

“ Gnici-oleracei Pers. 144
cohaesa Long 210
Compositarum Sch. 139
Conii Fekl. 196
Conopodii-Bistortee Kleb. 225
Convolvuli Cast. 175
-coronata Cord. 253, 255
coronata, specialisation of 68
coronifera Kleb. 255
Crepidis Schrét. 156

Cyani Pass. 140
depauperans Syd. 202
Diantht DC. 15, 38, 220
difformis K. et S. 168
Digraphidis Sopp. 267
dioice Magn. 244
Discoidearum Link 134, 135
dispersa E. et H. 260, 262
dispersa Hiriks. 261
distincta McAlp. 238
Epilobii DC. 199, 200

Epilobii-tetragoni Wint. 199
expansa Link 136
extensicola Plowr. 248
Fabae Link 97
fallens Cooke 90, 91
Fergussoni B. et Br. 203
Festuce Plowr. 257
Fragariastri DC. 290
fusca Wint. 215, 331
galatica Syd. 142
Galii Schw. 163, 164, 168
Galiorum Link 164
Gentiane Link 178
Glechomatis DC. . 173
glomerata Grev. 136, 320
glumarum FE. et H. 258
glumarum, specialisation of 67
graminis Pers. 41, 250
graminis, specialisation of 64
graminis var. Arundinis Cooke 271
Heraclei Grev. 194 —
Hieracii Mart. 137, 138, 141, 148,
149, 158
holeina Eriks. 263
Hordei Fekl. 259
Hydrocotyles Cooke 181
Hypocheridis Oud. 148
intermixta Friend 267
Tridis Wallr. 230
Lapsane Fekl. 147, 157
Leontodontis Jacky 149
Leucanthemi Pass. 133
Liliacearum Duby 234
linearis Rob. 266

-Lolii Niels. 255

Lolii, specialisation of 68
longissima, Schrét. 286
Luzulae Lib. 238
Lychnidearum Link 218, 221
Magnusiana Wirn. 128, 271
major Diet. 157
Malvacearum Mont. 48, 206
mamnillata Schrét. 225
Menthw Pers. 170

Millefolii Fekl. 131

mixta f. simplicior Korn, 121

. Moehringiae Fckl. 219

Molinie Tul. 276

mucronata var. Rubi Pers. 297

nemoralis Juel 276

neurophila De Toni 92

Noli-tangeris Cord. 204

oblongata Wint. 238

obscura Schrét. 236

obtegens Tul. 145

Opizti Bub. 152

Orchidearum-Phalaridis Kleb. 268,
344

Oxyrie Fekl. 224

GENERAL INDEX 409

PUCCINIA (cont.) PUCCINIA (cont.)

paliformis Fekl. 286 . Smyrnii-Olusatri Lindr. 197
paludosa Plowr. 248 Soldanelle Fekl. 180
Paridis Plowr. 267 Sonchi Rob. 155
Pazschkei Diet. 213 sparsa Cooke 150
perplexans Plowr. 126, 270 Spergulae DC. 219
perplexansf. drrhenatheri Kleb. 284 straminis Cooke 258, 260
persistens Plowr. 282 striola Link 241
Petroselini Lindr. 190 suaveolens Rost. 145
Phalaridis Plowr. 269 suaveolens var. Cyani Wint. 140
Phaseoli var. Taraxaci Reb. 154 Symphyti-Bromorum Mill. 262
Phlei-pratensis E. et H. 283 Syngenesiarum Cooke 144
Phragmitis Kirn. 273 Tanaceti DC. 134, 135
Piloselloidarum Probst 159 i Taraxaci Plowr. 154
Pimpinelle JJurt. 188, 192, 194, Thalictri Chev, 214

195 Thesii Chaill. 229
Poarum Niels, 45, 278 tinctorie Magn. 139
Polygont Pers. 227, 228 tinctorticola Magn.=P. tinctorie
Polygoni-amphibii Pers. 104, 227, Magn. 139

228 Tragopogi Cord. 150
Polygoni-Convolvuli DC. 104, 228 Trailii Plowr. 274
Polygoni-vivipari Karst. 226 Trifolit Hedw. 90
Polygonorum Link 227 Tripolii Wally, 129
Porri Wint. 122, 235 Triseti Eriks. 264
Potentillae Pers. 291 triticina Eriks. 262
Prenanthis Plowr. 151 truncata B. et Br. 230
Primule Duby 179 tumida Grev. °187
Pringsheimiana Kleb. 242 uliginosa Juel 249
Pruni DC. 208 Umbelliferarum DC. 187, 193
Pruni-spinose Pers. 207 Umbilici Guep. 211
Prunorum Link 208 vaginalium Link 117
pulverulenta Grev. 198 Valantize Pers. 167.
punctata Link 163, 164 variabilis Grev. 152, 154
Rhodiola B. et Br. 210 Veronice Schrét. 169
Ribesii-Caricis Kleb. 2438 Veronicarum DC. 169
Ribis DC. 212 Vinee Berk. 176
Rubigo-vera Plowr. 258, 260 violacea Schultz 295
Rubigo-vera var. simplex Korn. 264 Viole DC. 200
Rubi-Idaei DC. 298 Violarum Link 201
Sagine K. et S. 221 Virgauree Lib. 130
Sanquisorbae DC. 292 Winteriana Magn. 268, 348
Sanicule Grev. 182 Zopfii Wint. 217
Saxifrage Schl. 212 Pucciniacee 74, 75, 85
Sazifragarum Cooke 160, 212 PUCCINIASTRUM 74, 364
Schmidtiana Diet. 267 Abicti-Chamaenerii Kleb. 366
Schneideri Schrét. 172 Agrimonis Tranz. 364
Schoeleriana P. et J. 246 Aspidiotus Karst. 374
Schroeteri Pass. 232 Circewe Speg. 362, 365
Scirpi DC. 239 Epilobit Otth 366
Scorodoniae Link 175 Galii Fisch. 370
secalina Grove 261 Goeppertianum Kleb. 372
Senecionis Cooke 136 Myrtilli Arth. 371
sessilis Schn. 266, 267, 268 Padi Diet. 369
Silai Fekl. 191 Polypodii Diet. 375
Silenes Schrot. 222 pustulatum Diet. 363, 364, 366
silvatica Schrét. 158, 245 Pyrole Diet. 312, 367
simplex E. et H. 264 Vacciniorum Diet. 371
Smilacearum-Digraphidis Kleb. Pyronema conjluens Tul. 81

267

Smyrnii Cord. 197 Rawitscher 82

410 GENERAL INDEX

Red Alge 18, 23, 77, 83

RGSTELIA 30
cancellata Reb, 308
carpophila Bagnis 304
cornuta Tul. 307
Cydoniae Thiim. 306
lacerata Tul. 304

Schréter’s classification 39
sexuality 17
specialisation 62
spermatia 2, 22
spermogones 1, 32
sporophyte 27

syukaryon 5

teleutospores 11, 34

teleutospores, germination of 14, 49

tetraspore-mother-cell 26, 37
tetraspores 26, 28, 37, 80
THECOPSORA 74, 368
areolata Magn. 369
Galii De Toni 370
Padi Grove 368
Pyrolae Karst. 367
Vacciniorum Karst. 371

Trachyspora Alchemillae Fekl.

TRANZSCHELIA
cohaesa Arth. 210
punctata Arth. 208

TREMELLA
clavariaeformis Jacq. 304
juniperina Linn. 3807
Sabinae Dicks. 308

TRICHOBASIS
Angelicae Cooke 192
Apii Wallr. 184
Artemisiae Berk. 134
Betae Cooke 113
caricina Berk. 241
Cirsii Lasch 142
Clinopodit DC. 170
Conit Cooke 196
Cynapii DC. 190
Epilobii Berk. 198
Fabae Cooke 97
fallens Cooke 90, 91
Galiit Lév. 164
Geranii Cooke 103
glumarum Lévy. 258
Heraclei Berk. 194
Hieracti Cooke 158
Hydrocotyles Cooke 181
Impatientis Rab. 204
Iridis Cooke 230
Labiatarum Lév. 170
Lapsanae Cooke 147
linearis Lév. 250
Lynchit Berk. 385
oblongata Berk. 238

106

TRICHOBASIS (cont.)
Petroselint Berk. 190, 197
Pimpinellae Cooke 188
Polygonorum Berk. 117, 227
Primulae Cooke 179
Pyrolae Berk. 312, 367
Rhamni Cooke 209
Rubigo-vera Lév. 260
Rumicum DC, 114
suaveolens Lév. 145
Symphyti Lév. 363
Umbellatarum Lévy. 196
Vineae Berk. 176
Violarum Léy. 200

trichogyne 18, 19, 77

TRIPHRAGMIUM 75, 287
Filipendulae Pass. 287
Ulmarie Wint. 287

UREDINOPSIS 74, 379
filicina Magn. 362, 378, 379
Polypodii Liro 3875
Struthiopteridis Diet. 380
UREDO 378, 385
Acetosae Schum. 223
Egopodii Schum. 185
Agrimoniae Plowr. 364
Alchemillae Pers. 106
Alliorum Cooke 121, 235, 344
ambigua DC. 121
Angelicae Schum. 192
annularis Str. 175
Anthyllidis Grev. 95
apiculata var. Trifolit Str. 93
appendiculata var. Phaseoli Pers.
101
appendiculata var. Pisi Pers. 99
Aspidiotus Peck 3874
Betae Pers. 113 ,
Betae var. Convolvuli Pers. 175
bifrons DC. 116
bullata Pers. 193
Cacaliae DC. 325
Campanulae Pers. 328
Caricis Schum, 241
Caryophyllacearum Jobust. 360
Chenopodii Duby 111
Chrysanthemit Roze 131
Cichorii DC. 148
Circaeae Schum. 365
confluens DO. 351
confluens var. Euonymi Mart. 339
confluens var. Orchidis A. et 8. 343
Conit Str. 196
Cyani Schl. 140
Empetri Pers. 311
epitea K. et S. 340
Euonymi Cooke 339
Euphrasiae Schum, 326
Fabae Pers. 97

GENERAL INDEX 411

UREDO (cont.) uredospores 9, 32

farinosa var. Senecionis Pers. 320 UROMYCES 75, 85
Ficariae Schum. 107 Acetose Schrét. 116, 223
Filicum Desm. 375 Alchemille Lév. 106
Gentianae Str. 178 Alliorum DC, 121, 235
glumarum Schmidt 258 ambiguus Lévy. 121
Helioscopiae Pers. 353 Anthyllidis Schrét. 95
Hieracit Schum. 158 apiculatus Cooke 90
Hypericorum DC. 354 apiculosa Cooke 93, 114
Iridis DC. 230 appendiculatus Lév. 97, 101
linearis var. Polypodii Pers. 375 Armerize Lév. 89
Lint Schum, 355 . Behenis Ung. 109
Lynchii Plowr. 382, 385 Bete Lév. 118
Melampyri Reb. 827 caryophyllinus Wint. 108
miniata var. Lini Pers. 3855 Chenopodii Schrét. 111
Miilleri Schrét, 300 Colchici Mass. 122
Orchidis Mart. 343 concentricus Lév. 120
Oxalidis 390 concomitans B. et Br. 87
Padi K. et S. 369 Dactylidis Otth 125, 271, 273
Petroselini DC. 190 Dactylidis, specialisation of 69
Phaji Rac. 382 Ervi Westd. 96
Phillyreae Cooke 332 Erythronii DC. 118
Phragmitis Schum. 273 Euphorbiae-corniculati Jordi 94
Pimpinellae Str. 188 excavatus DC. 102
pinguis var. Rosae-alpinae DC, 294 Fabe De Bary 97
Plantaginis B. et Br. 385 Ficaria Lév. 107
Polygonorum DC. 227 Fischeri-Eduardi Magn. 101
Polypodii Schrét. 374, 375 flectens Lagh. 92
Polypodii var. Phegopteris Wint. Gageae Beck 120

379 Geranii O. et IF. 103
populina var. betulina Pers. 358 graminum Cooke 125
porphyrogenita Kze. 369 intrusa Cooke 106
Porri Sow. 235 Junci Tul. 123
Potentillarum DC. = chiefly the Kabatianus Bub. 104

ceoma-spores of Phragmidium Lathyri Fekl. 99

Fragariastri i Lilii Fekl. 118
Potentillarum var. Agrimoniae DC, Limonii Lév. 88, 89

364 lineolatus Desm. 124
Primulae DC. 179 Loti Blytt 94
Pteridum White 378 maritimae Plowr. 124
pustulata Pers. 366 minor Schrét. 90
Pyrolae Grev. 367 oblongus Vize 90
Quercus Brond, 315 Ornithogali Plowr. 120
Rhododendri DC. 384 Orobi Lév. 99
Rumicis Schum. 114 Parnassiae Cooke 389
Saxifragarum DC, 357 Phaseoli Wint. 101
Seillarum Grev. 120 Phaseolorum De Bary 101
Scirpi Cast. 124 Pisi Wint. 99
Scolopendrii Schrét. 377, 378 Pox Rab. 127, 273
Sempervivi A. et S. 335 Por, specialisation of 69
Soldanellae DC. 180 Polygoni Fekl. 117
Sonchi Pers. 324 proéminens Lév. 102
Symphyti DC. 363 Rumicis Wint. 114, 116
Tropeoli Desm. 386 Salicornie De Bary 114
Tussilaginis Schum, 322 Scillarum Wint. 120
Ulmariae Schum. 287 Scirpi Burr. 124
Vacciniorum Link 371 Scrophularie Fckl. 87
vagans var. Epilobii-tetragoni DC. scutellatus Plowr. 389

198 sparsus Lév. 111

Vineae DC. 176 striatus Schrét. 93

412 GENERAL INDEX

UROMYCES (cont.) “vacuole” 12
Trifolii Lév. 90, 91
Trifolii-repentis Liro 91 XENODOCHUS 75, 302
tuberculatus Fekl. 102, 354 carbonarius Schl. 303
Ulmariae Lév. 287 curtus Cooke 390
Urticae Cooke 389 Xyloma Virgaureae DC. 130
Valeriane Fekl. 86
Ustilaginacesee 76, 78, 82 ZAGHOUANIA 75, 331
Ustilaginales 82 Phillyrese Pat. 332
[2
FINIS

CAMBRIDGE: PRINTED BY JOHN CLAY, M.A. AT THE UNIVERSITY PRESS.