Full text of "The rusts of Australia: their structure, nature and classification"

See other formats

i i

f *

* *

NRLF

111

■H

* 9

* i

■

♦B fiOfl MM5

i :

■

» ," ' f *

■ ■

i *

*< i

itlllii!

•'

| Ki'll

■■■1

til?]

:.■•

> ♦ -

wjj! y

jljlijj [J

kj ■■ . ' i ■ (

4 I

1 ft \

€*m wv

- 9

THE

RUSTS

AUSTRALIA

(UREDINEAE).

+H>

438.

GALL-FUNGUS

DESTROYING GOLDEN WATTLE.

DEPARTMENT OF AGRICULTURE, VICTORIA

RUSTS

AUSTRALIA

THEIR

STRUCTURE, NATURE,

AND

CLASSIFICATION.

r>.

McALPINE,

Government Vegetable Pathologist.

WITH 55 PLATES (INCLUDING 366 FIGURES)

•

ittrlfcourne:

BY AUTHORITY : ROBT. S. BRAIN, GOVERNMENT PRINTER

438

1 906.

PREFACE.

The Rusts are among the most widespread and destructive of our fungus
parasites, and in order to mitigate the injury caused by them as much as
possible, it is necessary to know their nature and mode of life.

The rust of wheat has naturally received a large share of attention on
account of its influence upon one of our staple industries ; but it is still
only one of a number that require to be studied. Hence, at the several

Rust in Wheat Conferences held in the different States, investigations were
invariably recommended to be made " regarding all plants that are affected
by rust in the different colonies," because it was felt that such a wide
outlook was necessary even for understanding properly the history of a
single species. The present work, then, which has been in hand for
a number of years, aims at recording all rusts, as far as known in Australia
at present, and this will prepare the way for a consideration of the best
methods of preventing their appearance, or limiting their spread in the
numerous commercial crops subject to their ravages.

The familiar saying that to know any subject well we must know the
details of it, is very applicable in this case. The necessary details are
given here to enable one to recognise the different forms the rusts assume,
and the different stages through which they generally pass in order to
complete their life-history. Besides, there is a special object in view
in thus recording and describing the Rust-fungi of Australia, for this
can afterwards be used as a basis in working out the life-history of those
particular forms which attack our cultivated and economic plants, and
often do considerable damage.

All the species known to occur in Australia are included, and when
proved to be aliens, they are noted as introduce*} in the index.
Every species of which specimens are available is also figured in its
essential parts, so that there may be no doubt as to the form intended.

In this I have been ably aided by my assistant, Mr. G. H. Robinson,

who has supplied the numerous photomicrographs reproduced here,
which give such a vivid representation of trie peculiar and distinguishing
characters of the spores. A number of drawings have likewise been

executed by Mr. C. C. Brittlebank, who has had considerable experience

in drawing from microscopic preparations.

The principal works consulted, or referred to, are given in the
Bibliography at the end, and those wishing for fuller references, will find
them in Klebahln's Die wirtswechselnden Rostpilze [The Heteroecious
Rusts], 1904. This will give a good idea of what has already been done
in the investigation of the rusts, but no one can fail to appreciate the
work accomplished by that veteran mycologist, Dr. Cooke, in his

/5-J23

Handbook of Australian Fungi. When one considers that the material

had to be sent such long distances, and often limited in quantity, as well
as imperfectly preserved, it is surprising the number of rusts recorded,
and the general accuracy of the descriptions. It is a matter within my
personal experience, that in order to do justice to the rusts, it is necessary
to have plenty of material and to have it fresh, and there is always a
decided advantage in collecting your own specimens. The Australian
rusts recorded in the Handbook published in 1892 were 72, and the
number now has reached 161.

To all those who have contributed specimens, my best thanks are due.
The late Mr. Luehmann, F.L.S., Government Botanist of Victoria, allowed
me free access to the specimens in the National Herbarium and Mr. Bailey,
F.L.S., the Government Botanist of Queensland, who has done so much
in every division of botany, always willingly aided me with specimens or
information; the Government Botanist of New South Wales, Mr. Maiden,
F.L.S., as well as R. T. Baker, F.L.S., of the Technological Museum,.

and A. G. Hamilton also supplied me with any material required from the
sister State, as well as Dr. Morrison, of Western Australia, and Mr.
Rodway, F.L.S., of Tasmania. Messrs. Molineux and Quinn, of South

Australia, have also contributed, and Mr. J. G. O. Tepper, F.L.S., has

generously supplied me with numerous, and often type specimens from his
extensive collection.

My colleague, Mr. French, F.L.S., Government Entomologist, and his
assistant, Mr. C. French, junior, never lost an opportunity of securing
specimens in their frequent collecting trips; and Messrs. Reader and
Musson have added new species to the list. I have also to acknowledge
the courtesy of the Director of the Royal Gardens, Kew, in supplyii
me with any specimens required for verification or illustration.

No one is more conscious than myself of how much yet remains to
be done before the Rusts of Australia are thoroughly understood, but the
present work will at least lighten the labours of those who desire to
increase that knowledge, and by the combined efforts of various workers
in this promising field, their true nature and life-history may be so
revealed that the ravages due to them, in a congenial climate such as ours,
may be reduced to a minimum.

Melbourne, March, 1906.

•

CONTENTS

-r>

PART FIrv

CHAPTER.

PAGE

I. Introduction

• •

II. Vegetative Organs — Mycelium
III. Reproductive Organs — Spores

IV. Spermogonia and Spermatia
V. Aecidia and Aecidiospores

VI. Uredospores

VII. Teleutospores ...

VIII. Mesospores and Amphispores

IX. Sporidiola or Promycelial Spores

X. Paraphyses and their Function

• -

XII

• •

■ - •

• * «

• • •

• •

XI. Origin of the Principal Spore-forms

- 9

Rusts in their relation to other Fungi

XIII. Indigenous and Introduced Species

XIV. Indigenous Species with their Hosts
XV. Australian Distribution ...

XVI. Origin and Specialisation of Parasitism

XVII. Heteroecism and its Origin

XVIII. Predisposition ...

t *

• *

• •

• • •

• •

- * -

• •

• • •

■ • ■

• 9

• *

• •

• • •

• *

• •

The present position of the Rust in Wheat question in Austr

aim

1
3

7
13

1G
19

27
29
31

40
42
45
50
52

60
64

PART SECOND.

Classification with special reference to Biologic forms
XXI. Systematic Arrangement and Technical Descriptions-

Uromyces ...
Uromycladium

• • •

Puccinia

• •

Phragmidium

Cronartium

Melampsora

Caeoma

• #

Aecidium . . .

Uredo

• • •

• •

# •

• t •

• •

• • •

t *

« •

• • •

* •

• •

• • •

• •

• • •

• •

Excluded or Doubtful Species

Glossary of the principal scientific terms used

Literature consulted

Explanation of Plates

Host Index with Rusts ...
Fungus Index with Synonyms and
General Index ...

• •

- - -

• •

» -

- - •

• •

9 •

• •

84
104
112
185
189
191
193
194
202

207
211
213

OO0|

°29

337
343

PART FIRST.

GENERAL CHARACTERS AND MODE OF LIFE

CHAPTER 1

Introduction.

The Uredineae, or rusts, constitute one of the most important group

of parasitic fungi, and their ravages are known wherever plants are

cultivated. The cereals and grasses of our fields, the fruit trees of our
orchards, even the ornamental plants of our gardens, and many of our
forest trees are attacked by members of this family, and their study is not
only interesting from the point of view of the scientist, but from that
of every grower of plants for pleasure or for profit. Rusts are usually
so conspicuous that they attract the attention of even the ordinary observer,
and they have been known and recognised even from the earliest times,
particularly from their blighting effects on the wheat and corn crops.

Although so long known as regards their naked-eye characters, and
the effects they produce, their structure and life-history have only been
understood within comparatively recent times, and even now there are
many points concerning them which await investigation.

Their structure essentially consists of an inconspicuous mycelium
bearing the usually conspicuous spores, and while this vegetative mycelium
is generally similar throughout the group, the spores produced by it are
very dissimilar. The general study of this group will therefore mainly
resolve itself into a knowledge of the different spore forms, and their
relation to each other, either on the same plant or on different plants. The
finishing spore or teleutospore may be regarded as the ultimate stage of
the Uredineae, and which, after usually resting for a period, long or short,
germinates by putting forth a germ-tube, which bears, in turn, another kind
of spore. The germ-tube is known as the -pro- mycelium, and the spore as
the promycelial spore, or sporidiolum, so that if the latter is regarded as the
starting point, the teleutospore will constitute the finish.

Between these two forms there may be various intermediate stages, and
the series may consist of the following: —

i. The sporidiolum, when it produces its germ-tube, enters the

tissues of the host-plant, and may either produce from its
mycelium teleutospores similar ,to those from which it
originated ; or

2. It may give rise to uredospores at first, and subsequently

teleutospore ; or

3. It may produce aecidiospores, uredospores, and teleutospores in

succession, the aecidiospores being generally preceded or

accompanied by a peculiar form of spore, known as a
spermatium.

Hence the complete series of spore-forms will be spermatium, aecidiospore.
uredospore, teleutospore, and sporidiolum, although between the initial
sporidiolum and the final teleutospore, one or more of the above may be
suppressed or omitted in the life-cycle.

Introduction.

This succession of spore-for

be represented

lly by the

following diagrams

Sporidiolum

Teleutoapore

Sporidiolum

Aecidloepore

Teleutospore

Uredoepore
Fig. 1.

Aecidiospore

Fig. 1.

Sporidiolum

Teleutospore

I 'redospore

Fig. 3.

The first show

the

plete cycle of develop

spore- forms follow each other in

common form

invariable (

the second the uredosp

der

Teleutospore
Fig. 4.

lent in which all the
and this is the most

ppressed

d the

number of species unde
the third the aeeidiosix

this contracted cycle is much reduced

further reduced

And

is wanting, and the number of forms is still
fourth the cycle of development is reduced

to its lowest limits, a direct succession of teleutospores occurs, and the
number of species, instead of reaching a minimum, probablv ranks next to

those

The spermatia succeed the sporidiola, and are generally present in the

fe -cycle, but
Besides th

general development.

jse regular forms, there are others which are generally re-
garded as representing either stages in the life-history of imperfectlv known

species o
are know

degraded forms of which onlv the uredo

The subject, therefore, naturally divides itself into a consideration
the vegetative organs or mycelium and the various reproductive bodies
spores enumerated above, together with the structures accompanying thei

The life-history of each form, as far as known, will ibe briefly
but this has still to be investigated in most of the recorded snecie«

sketched ;

Vegetative Organs.

CHAPTER II.

Vegetative Organs — Mycelium.

The vegetative portion of many fungi is very inconspicuous as com-
pared with the reproductive, but its importance is not to be measured by
its size or extent, rather by the part it plays in the life of the organism ;
and since it is the foundation of the whole, if is worthy of the most careful
study.

Of late years, however, this part has come into special prominence,
particularly in the case of the cereal rusts, for it has been asserted that
it is not always by external infection that the rust begins its career in the
growing plant, but that in some cases it originates from within, and this
theory will engage our attention later on. Meanwhile this is referred to
to show that the key to the propagation of the rusts from year to year may
be not only on the surface, among the special reproductive bodies which
spread it throughout the growing season, but also in the interior among the
cells where the first beginnings of its life may appear.

Among the recent investigations on the mycelium, there are two which
stand out on account of their completeness, owing to the use of the most
modern histological methods — the one by Professor Marshall Ward9 on the
Histology of Uredo dispersa Eriks, and the Mycoplasm hypothesis, and the
other by Professor Eriksson u> 15 on Das vegetative Lcben der Getreiderost-
pilze [The Vegetative Life of the Cereal Rusts]. The study of an indivi-
dual case will prepare us for the more general examination of the
mycelium throughout the rusts, and we will begin with that of Puccinia
dispersa Eriks., or, strictly speaking, P. bromina Eriks., which Ward has
so thoroughly dealt with and illustrated with such admirable clearness.

Starting with the germination of the uredospore on the surface of the
leaf, which usually occurs within twenty-four hours, we find that the voung
germ-tube grows rapidly, and that the nucleus of the spore passes into it ;
sometimes, however, two or more nuclei may appear in it. The tip of the
tube begins to swell over a breathing pore or stoma into a thin vesicle, and
the contents derived from the spore accumulate here.

This external vesicle or appressoritim, as it is called, is the first stage
in inoculation from the outside, for a thin process is passed through the
opening of the stoma, and swells inside into another vesicle. The proto-
plasmic contents are transferred from the external to the internal vesicle,
and so the future growth takes place among the tissues of the leaf.

At one or more points this inner swelling forms a delicate tube, into
which the protoplasm is again transferred, and its nucleus soon divides.
This is the first-formed hypha, and the foundation of the vegetative system.
It soon branches and develops cross partitions or septa, and extends rapidly
among the cells of the host-plant to form the mycelium. Even at an early
stage, when the primary hypha is still unbranched and unseptate, suckers
or haustoria may be formed to provide a large imbibing surface for the
fungus. The haustorium begins as a small delicate process or projection
from the hypha, and this pierces the cell-wall and swells up into a minute
spherical head, which is provided with a nucleus. Shortly after entering
the cell this head takes on an irregular growth, and may assume a variety
of shapes.

The mycelium now becomes denser towards the surface, and prepares
for the production of the reproductive bodies or spores. This constitutes
the history of the mycelium from the time it starts as a germ-tube until
it reaches its full development.

•I

Vegetative Organs.

Eriksson has investigated on similar lines, and fully illustrated the
mycdium^f Puccima glumarum, Eriks. and Henn., but with this important
Xence °hat instead of starting with external infection from a spore, he
begins with an assumed internal germ of disease, which he considers in
certain cases to be a source of rust, in addition to the ordinary infection

spores. It is often stated that this rust passes the winter as mycelium
„i such leaves as are attacked in late autumn, and which persists till the
following spring; but the examination of hundreds of sections of leaves
taken from rusty plants, although not rusty at the place chosen for section,
failed to reveal the presence of such a mycelium dormant in the tissues.
It may be taken for granted, then, that there is no mycelium to start with,
and it will be interesting to follow Eriksson's theory as to the

h the mycelium arises afresh

of the autumn and spring leaves a peculiar thick plasma is found, containing
a distinct nucleus, and this Eriksson considers to be, not the ordinary pro-
toplasm of the cell, but a mixture of it, with the earliest vegetative form
of the fungus. This intimate mixture or symbiosis, or living together ot
the ordinary protoplasm of the host and that of the fungus he distinguishes
as mycoplasm. This mvcoplasm is stated to occur only in certain cells.
which favours the assumption that it is not a necessary constituent of

the

The next step and the youngest stage of mycelial formation, according
to Eriksson, is the presence of a plasma in the intercellular spaces, which
is partly in the form of growing filaments, partly as irregular masses.
There are no septa, and no distinctly recognizable nucleus, and even a dis-
tinct wall is not formed. The primary stage is quickly followed by a
secondary stage, in which the only visible' advance is a very distinct nucleus.
These two stages are very sharply marked off from the normal mycelium,
both by their plasmodia-like nature and the absence of transverse septa,
and for distinction the special name of protomycelium is given. Eriksson
has no doubt that the intracellular mycoplasm and the inter-
cellular protomycelium are genetically connected, but this, which is a
necessary link in the chain of evidence, requires to be further investigated.
(Note i, p. 74.) The formation of haustoria is the next* process, and consists
in a small straight prolongation of the protomycelium passing into the
interior of the cell, and at the apex forming a globular swelling, probably
containing a nucleus. Soon the whole forms a sac-like irregular organ.
which may become detached from the protomycelium. These detached
bodies in the cell were mistaken by Eriksson for a preliminary stage in

the formation of hyphae, and called "special corpuscles," but Ward pointed
out their true nature, and that they really had been formed by, instead
of giving rise to the hyphae, a correction which Eriksson himself has
acknowledged. The haustoria are often found closely adjoining the
nucleus, which thereby degenerates, and simultaneously with the shrinking
of the nucleus, and soon after the first entrance of the haustoria, trans-
verse septa begin to be formed in the protomycelium. In most of the cells
thus formed several nuclei are contained, and the stage is now reached
where a true mycelium is present, composed of hyphae.

This multiplication of the cells of the fung,us is a sign of advancing
maturity. By continued division a true pseudo-far enchyma is formed,
and at certain spots, where the cells appear to be particularly rich in food-
material, a kind of hymenium arises, from which ultimately the spores
are detached. Where spores are being formed, there the complete destruc-
tion of the cells of the host-plant occurs, and now the vegetative life of the
fungus is ended, and the reproductive phase is entered upon.

Vegetative Organs.

Perennial Mycelium. — In contrast to the localized mycelium, there may-
be a mycelium with unlimited growth which does not confine itself to par-
ticular spots, but may permeate entire shoots, or even the whole plant.
This is known as a perennial mycelium, and wherever it occurs the fungus
may reappear on the same plant year after year without the necessity for
reinfection by means of spores.

This vegetative reproduction through a perennial mycelium is not always
easy to prove, but its importance cannot be overrated, for hidden in the
tissues of the plant it cannot be reached by the ordinary means for con-
trolling the growth and spread of fungi, but involves the destruction of
the plant, or at least of those parts which harbour it. As Australian ex-
amples, we may note Uromyces trifolii, which attacks the white clover
(Trifolium repens) and Phragmidium subcorticium or rose-rust, in which
the mycelium of the aecidial stage penetrates all the tissues, and in each
succeeding year forms a new layer beneath the old.

Uromycladium notabile and U. tepperianum occurring on species of
Acacia are further examples, for the mycelium gives rise to large galls,
which persist from year to year and produce spores.

Even although the host-plant is an annual, and dies down every year,
it is still possible for the mycelium to be perennial, for it may be carried

Carleton

the seed, as in the case of Uromyces euphorbiae, according

W itc he sy -brooms. — It is not unusual among the forest trees and shrubs of
Europe to find shoots very much deformed and distorted, and looking
at a distance like large birds' nests or brooms, and to these the popular
name of witches' -brooms has been given. These peculiar and diseased
conditions were difficult to account for, and so the idea may have originated
with superstitious people that the trees were bewitched, in order that the
witches might be provided with brooms for their midnight rides, hence
the name.

But the true cause is seen when the matter is investigated in the light
of our present knowledge, and parasitic fungi are often found to be respon-
sible for the strange transformations of the normal shoots into the dense
twiggy, irregular tufts met with. This may also be produced by other
means, such as gall-mites, but a very striking case and the first recorded
instance in Australia is that of the rust-fungus, Cronartium jacksoniae,
which deforms the shoots of various leguminous plants as shown in
Pis. XXXVII., XXXVIII. Uromycladium tepperianuni also produces
this peculiar appearance on Acacias as shown in PI. XLII.

The perennial mycelium in the shoots stimulates a number of buds to
abnormal growth, quite different from the ordinary, and the result is seen
in the numerous densely crowded and considerably altered shoots as com-
pared with the normal. They are also thickly studded with the ruddy-
brown columns of teleutospores, in the case of Cronartium, which stand out
like so many curved or straight, stiff bristles, towards the ends of the
shoots, which are gradually being destroyed. Next year the mycelium will
grow into the young shoots and produce the same result.

Formation of Galls. — It is well known that the mycelium of fungi exer-
cises a stimulating effect upon growth, and not only causes the cells to grow
larger and divide more frequently than usual, and the chlorophyll to dis-
appear, but it may alter the character of the tissues. When an insect pierces
the young and living tissue of a plant with its proboscis or ovipositor,
it often causes the cells immediately surrounding it to grow and divide
more rapidly than elsewhere, so that a swelling of the tissue occurs, which
is known as a gall. So among the rusts there are instances where the stimu-
lation of growth occurs in a marked degree, and if a vegetable gall be

Vegetative Organs.

considered as a morbid enlargement of the affected part of the plant, due
to parasitic agency (Connold1), then there need be no hesitation in calling
these structures galls.

Perhaps the most striking illustration of a gall is seen in Uromy-
cladium tepperianum. In the neighbourhood of Melbourne hedges of Kan-
garoo Acacias (A. armata) are being gradually and completely destroyed by
the ravages of this fungus, which resemble on a superficial view large galls
caused by insects. Most of the branches, including the phyllodes, are
infested with the chocolate- brown swellings, which may be in the form of a
succession of small excrescences about the size of peas, or collected into
large clumps about the size of walnuts, and measuring 4 cm. across. In
some cases they are solid round knobs, and the external appearance is due
to the dense covering of the chocolate-brown teleutospores. One of the lar-
gest was met with at Mymiong on Acacia imflexa of an irregular leg-of-
mutton shape and weighing about 3 lb, (PI. XLI).

On A. pycnantha, or Golden Wattle, the galls are as large as potatoes,
and in some of the wattle plantations, where the trees are cultivated for
their bark, they hang in large numbers from the branches like so many
fruits, and numbers of the trees are either dying or dead. The swellings
are primarily caused by the fungus, and then various insect larvae may
ultimately invade them, boring and tunnelling through them. In A. implexa
the swellings may run along the whole length of the elongated phyllodes,
and in A. salicina there is an all-round swelling of the branches, and the
periderm is ultimately ruptured. Magnus3 found the galls to be per-
meated by an intercellular mycelium, which was multiseptate with numerous
and somewhat branched haustoria.

Some very large galls were also found on the Black Wattle (A. decur-
rens) and Silver Wattle (A. dealbata) either surrounding or terminating the
branches, and caused by U. notabile. Some measured 4-5 inches across,
and 3-4 inches was not uncommon, while one of the largest weighed

&v-^"- "^f)

The j>eculiar gall-like swellings caused by Gymno sporangium may be
mentioned, the mycelium of which is perennial in the various specie's of
Juniper, and from their appearance are popularly known as "cedar
apples " in America. It may be an annual gall only bearing the teleuto-
spores for one season, or a perennial gall, producing successive crops of
teleutospores year after year, and not requiring the transfer of the spores
each season. v

Localized Mycelium.— The localized mycelium may likewise produce
conspicuous swellings, particularly on the stem and midrib of the leaves
Thus, that of Aecidinm urticae causes hard curved thick swellings of con-
siderable extent, and such a development of starch takes place in some
Himalayan species of nettle attacked by this fungus that the natives eat
the overgrown and hypertrophied stems for food.

In other cases the affected tissues mav be so stimulated bv
mycelium as to cause their death. Thus almond Ip9vm Kni,

localized
been found

here badly riddled with "shot-hole," due to the mycelium of Puccim»
prum Pers just as P. malvacearum may also destroy a circumscribed
area which becomes separated around its circumference, and fal

out,

The mycelium, whether localized or perennial, is always beneath the
surface of the plant, and formed within the living tissue } I ^delicate
in texture, like all internal mycelia, and branch* to form a e.ular net
work ultimately forming compact cushions or spore-beds I^an often be
traced from a single point of infection, whence it radiates all round i and
spreads, gathering material for the fresh production of snorP,

Reproductive Organs

CHAPTER III.

Reproductive Organs — Spores.

A detailed account of the different spore-forms will be given in the sys-
tematic part in connexion with the various species ; but it will be convenient
here to take a preliminary view of some of the more essential general
features, such as distribution, germination, and infection, suppression or
omission, and repetition of spore-forms.

Distribution.

The most important means of distribution of the rust-spores, as for
fungus spores generally, is the wind. They are usually exposed on
longer or shorter stalks, often powdery, and their immense numbers and
lightness all render them easily detached, and spread by the slightest
breath of wind.

That the. wind is an important factor in the distribution of rust-spores
is strikingly shown in an example given by Halsted *. The asparagus
rust (Puccinia as-paragi, DC.) was very bad, but in one field the plants had
been cut over, and the rusty brush removed in order that the new growth
might escape. On examining this field about five weeks afterwards, he
found that the rust showed only on one side of the green plants, and that
was the side exposed to an old and very badly rustedl asparagus bed. That
this was the source of infection was obvious from the fact that a house with
a few trees around it intercepted a portion of this field, and there was less
rust upon that portion of the new bed in line with the house.

The dust-storms which occur will also be a fruitful source of spreading
them, and in our northern areas where the dust is sometimes carried in
such quantities as to obliterate fences, it can easily be understood how even
in virgin soil the spores of wheat rust may be found.

The rain will also help to scatter them, particularly over individual
plants, and on the surface of the soil, for it is a common observation how
rusty plants are cleared after a heavy shower.

Insects as well as other animals serve the same purpose. I have often
observed the larvae of a species of Cecidomyia feeding on uredo and
aecidio-spores, and at the same time spreading them while crawling along.
Lindroth1 has observed in Finland the occurrence of these larvae on no
less than sixty-one species of rusts, and similar larvae have been found feed-
ing on the conidia of various species of Oidium (Salmon1). In some pot
experiments with wheat, I found the larvae associated with the Oidium of
E. graminis, and feeding on the conidia. At first sight it might seem as if
this devouring of the spores would tend to reduce the fungus, but the
wholesale way in which the spores are carried about more than counter-
balances anv decrease from this cause.

Germination.

As a rule, uredo and aecidio-spores germinate easily in a damp chamber,
and I have found the method recommended by Plowright to be very con-
venient. A gardener's propagating glass is used, placed on a plate of
water, and a simple stand is made with two flat oblong pieces of gutta-
percha. Holes are easilv bored in them, and glass rods placed parallel
ietween them, so that the microscopic slides may be laid across the rods.

Reproductive Organs.

The influence of various chemical substances on germination has been in-
vesLated, and it has been found that some substances, such as solutions
ITSS*. nitrate solutions, hasten germination; but for ^ purp^s

that is necessan

Sometimes, however, the spores

germinate and infect the leaves of the host-plant, although they may not

d° Freem^'found this to be the case in dealing with the uredo-spores
of Puecinia bromina, and concludes that the negative results in distilled
water tests are not always an indication that the spores are incapable of
germination. The temperature seems to exercise an important influence.
Eriksson found that in manv cases the spores germinated more freely if
previously exposed to a temperature of o deg. C. or under and Marshall
Ward that the uredospores of P. bromina were not injured by being ex-
posed to a temperature of - 5 deg. C. for ten minutes, while the same tem-
perature continued for four to five hours killed them. Heat or cold, drought
or damp, age and ripeness, are all factors of importance in germination.

Duration of Germinating Power.

How long do the spores retain their germinating power is an important
question to settle; but not many definite determinations have been made.

De Bary states that the uredospores of P. graminis, kept dry, lose their
germinating power in one to two months.

Marshall Ward found that the uredospores of P. bromina preserved dry
for sixty-one days retained their germinating power ; but it was feeble.

Barclay found certain uredospores still capable of germination after
from two to eight months, the leaves on which they occurred being kept

drv. % .

It is a question of great scientific interest in connexion with rust in

wheat if the uredospores can retain their germinating power during the
winter. The results vary, as might be expected, according to the condi-
tions prevailing at the time. In contrast to De Bary's results, Eriksson
found that the uredospores of P. graminis lost their germinating power
during the winter if kept in the open, but retained it if kept inside.
Hitchcock and Carleton1, however, collected fresh uredospores from
growing plants of P. rubigo-vera at various times during the winter, and
found them capable of germination. In our comparatively mild winters
the uredospores retain their power of germination, and this seems to be the
means whereby the fungus is continued from season to season. I have
had freshly gathered uredospores of P. graminis from growing wheat and
oats, and P. triticina from wheat germinating freely in water in winter
une to August). The uredospores can germinate at once, and directly
feet the host-plant, or they can act as resting spores for a time, and
freely infect the next season's growth, under the climatic conditions which

prevail here.

The germination of teleutospores takes place ait different periods, ac-
cording to the nature of the species. They may either germinate imme-
diately on reaching maturity, or, as is the case in the majority of heteroecious
iusts, only after undergoing a period of rest — in the old wrorld usually in
the winter, but here, as no doubt sometimes elsewhere, the period of rest
is often partly in the summer, when drought checks growth as effectively
as the cold of a European winter.

Eriksson10 has shown that the teleutospores of P. graminis, with few

exceptions, only germinate in the spring following their formation, and
only then if kept in the open during the winter. He kept spores in the

herbarium for

and then on exposing them for another

he found that they germinated, but this was exceptional

Reproductive Organs

In other species there is considerable variation. According to Woronin1,
the teleutospores of P. helianthi germinated equally well when kept dry in a
room, or when taken from the leaves of a plant which had been under the
snow all winter; and Carleton 3 found them to germinate even without a
resting period.

After wintering, Eriksson found that the teleutospores of P. graminis
could retain their germinating power even to the beginning of autumn —
September in Sweden.

In Victoria the germination of the teleutospores of P. graminis has been
tested for several seasons, and it is found that after a period of rest ex-
tending over about eight months, and including the usually hot and dry
summer, they begin to germinate in our spring (September), and if the
weather conditions are favourable, may retain their germinating power
until November. Rusted straw has been kept outside exposed to the
weather, and in a room, but no germination took place outside of the above
months. When spores have once begun to germinate it is a mistake to
suppose that they will all germinate at the same time, for there is a certain
irregularity about germination in the open which causes the period of ger-
mination to be considerably extended.

Infection.

The mode of infection usually varies in different classes of spores. The
germ tube of the uredospore or aecidiospore generally enters by the stomata
into the tissues of the host-plant, while that of the sporidiolum bores through
the epidermis direct, and this difference in (the mode of infection influences
the stage at which it occurs in the development of the host-plant. The
sporidiola are, as a rule, only able to infect young and tender portions of
the plant; but the uredospores and aecidiospores can evidently attack old
and fully developed organs.

Further, since the germ-tube of the sporidiolum in piercing the epider-
mal cells must dissolve the wall at the spot where it enters, there is evidently
a reciprocal influence exerted between the parasite and its host; but even in

the case of the uredospores and aecidiospores, where an entrance is effected
through the stomata, there also exists some mutual relation, for it is only
in certain hosts that the parasite can grow and develop.

This relation will be more fully dealt with when the origin of para-
sitism is considered, but meanwhile the fact may be emphasized, that the
mere entrance of the germ-tube through the stoma does not constitute
infection. Miss Gibson1 carried out infection experiments with uredo-
spores and aecidiospores of various rusts on a number of plants other than
the original hosts, and she found that while the germ-tube may enter the
stoma freely, yet, once inside, death ensues sooner or later, and in no single
instance was a haustorium formed. The fungus was unable to penetrate
the cells of the foreign host-plant, and so died of starvation.

The period which elapses between the entrance of the germ-tube into
the host-plant and the appearance of the result is known as the incubation
period. During this time the germ-tube grows and ramifies among the
tissues, abstracting nourishment from the cells by means of haustoria, and
finally proceeds itfo the formation of fresh spores. The first visible trace of
this does not generally appear before eight days, but, according to the
nature of the fungus, it may take much longer. Schimper, in his1 masterly
work on Plant Geography, begins with the statement — " No factor affecting
plant life is so thoroughly clear as the influence of water," and even^ for
parasitic fungi this factor is of prime importance. Smith l has determined
the water relation for Puccinia on asparagus, and probably the same prin-
ciple applies in a general way to other rusts. From a study of the direct

Reproductive Organs.

relation or the effect of atmospheric moisture upon the spores or mycelium
it has been shown that dew is absolutely necessary for infection, and ot
more importance than rain, and, in fact, that without dew there can be no
infection. Further, that the effects of atmospheric dryness not only
influence spore germination, but also spore production, and the aecidial and
uredo stages art checked thereby, but if the mycelium is not completely
destroyed, the teleuto stage may appear independently of conditions un-
favorable to the others. The indirect relation of water or the effect of soil
moisture, which acts upon the parasite through lits host, has also been
studied and shown to be of great importance. An abundance of soil
moisture at the growing season, in the case of asparagus rust in California,
is stated to increase the vigour and vitality of the host, and retard the
development of the fungus. That the growth of the fungus is retarded
„.■ increased vitality of the host scarcely agrees with Marshall Ward s
conclusions in regard to brome rust (Chap. XVIII.). That the weather
exercises a commanding influence in hastening the development of rust in
wheat is abundantly shown by its rapid spread in hot " muggy weather,
especially when accompanied by heavy dews. Irrigation before ploughing
gives the wheat a good start, but if applied in the spring it makes the wheat
soft and very liable to rust, a case in which great vigor of growth of the
host is followed bv a strong development of the rust.

While the ordinary mode of infection is as stated above, exceptions may
occur. Thus, according to De Bary, the germ-tube of the sporidiolum of
Puccinia dianthi {P. arenariae) may enter by the stomata, and Bolley
gives illustrations to show that the germ-tube of the uredospores of P.
triticina mav enter direct through the epidermal cells as well as by the

stomata.

Suppression or Omission of S

As already pointed out in the Introduction, the complete cycle of develop-
ment follows an invariable order. The germinating teleutospore gives rise to
sporidiola, from the mycelium of which arises the spermogonia and aecidia ;
later the uredospores appear, and finally the teleutospores complete the

cycle.

The aecidio, uredo, and teleutospore forms are represented respectively

for convenience by the Roman numerals I. II., III., and if the sporidiola
arising from the teleutospore are designated by IV., and spermogonia

a cvpher, then it will be comparatively easy to represent the different
stages diagramatically. A complete cycle of development will be repre-
sented thus— O., I., II., III., IV.

The teleutospore with its sporidiola is believed to be a constant factor

the cycle, although the

been

found, so that I. or II., or both, may be omitted or suppressed, as well as O

hich. however, is rarelv absent

diolum produces

mycelium which may in turn give rise to spermogonia and aecidia. These may
either occur on the same species of host-plant as that which bears the teleuto-
spore, and the rust is thus said to be autoecious, or they may occur on a
totally different plant when the rust is said to be heteroecious. The aecidio-
spore on germinating may skip the uredo stage, and directly produce the
teleutospore, and thus the cycle is shortened. There are numerous examples
of this in Australian forms, among which may be mentioned the rust on the
marigold {P. calendulae), and that on the daisy (P. distinctd).

This variation in the alternation of spore-forms has been used by
Schroeter as a basis of grouping, and although these biological sections by
no means indicate close affinity, yet they conveniently group together forms
>\hich have similar associated stages. Taking the genus Puccinia as an

Reproductive Organs.

example — and it applies to other genera as well — the complete cycle is
designated Eu-puccinia, and represented diagramatically as O., I. II.,
III., IV.

Then the stage in which the uredo does not appear is known as

Pucciniopsis, and represented by O., I., III., IV. Again, the germinat-
ing sporidiolum may produce a mycelium which bears spermogonia and ure-
dospores, thus skipping the aecidial stage. This stage is called Brachy-
pucrinia, and represented by O., II., III., IV., or the spermogonia may like-
wise be suppressed when practically only half of the cycle is retained, and
it is known as Hemi-puccinia, represented by II., III., IV. This is a stage
which is probably based upon imperfect observation, and the presumed
absence of the spermogonia. Further, the germinating sporidiolum may skip
the aecidial and uredo-stage and produce the teleutospore direct, with its
accompanying spermogonium. This is known as Lepto-puccinia when the
teleutospore germinates immediately, and Micro-puccinia when it undergoes
a period of rest. lit would be represented diagramatically as O. III., IV.

As might be anticipated, the greatest number of species have a complete
life cycle, and next to that come probably those species in which the life
cycle is reduced to its lowest limits, thus showing that the aecidial and
uredo generations may be dispensed with. In the former case (there is, as
it were, a succession of forms adapted to the varying seasons, well fitted to
meet any sudden changes, and living upon the same, or having a change of
host-plant as may be found most convenient, while in the latter the change
consists not in the variety of spore-forms, but in having a general purpose
spore, which can either germinate upon the living plant as soon as it reaches
maturity or undergo a period of rest in ithe decayed organic matter before
producing sporidiola. Hollyrock rust, or Puccinra malvacearum, is a good
illustration of a single-spored form adapting itself to the most varied con-
ditions, and making its way in the world.

The complete scheme for grouping Puccinias, according to the alterna-
tion of their spore- forms, is as follows: —

Eu-puccinia, O., I., II., III., IV.
Pucciniopsis, O., I., III., IV.
Brachy-puccinia, O., II., III., IV.
Hemi-puccinia, II., III., IV.

Lepto-puccinia, O., III., IV. (Teleutospores germinating imme-
diately).
Micro-puccinia, O., III., IV. (Teleutospores germinating after
a period of rest).

Repetition of Spore-forms.

In connexion with the suppression or omission of spore-forms, we ma

sider the repetition of spore-forms, because it would appear that

in many instances the suppression is brought about or initiated by one of
the spore-forms being able to repeat itself in the course of the cycle, and
thus making up for the loss of one member of it. It is the doctrine of
substitution in another form. This will be more conveniently dealt with
when treating specially of the aecidio and uredo spores, where it will be
seen that when the uredo generation is suppressed or omitted, the aecidio-
spores often repeat themselves, and when the aecidial generation is
omitted the uredospores repeat themselves.

Investigation of Spores. .

The microscopic investigation of rust spores is comparatively easy,
although it requires careful observation with regard to their minute details.

Reproductive Organs.

If the loose spores are examined as to their individual structure, whether
aecidio uredo or teleutospores, it is simply necessary to mount them
direct, and I find the most convenient mounting medium to be glycerine
and water in equal proportions. For permanent preparations glycerine
jelly may be preferable; but for the rapid examination of material the
above serves very well, and where desirable the mount can be preserved.
The surface markings of the spores are often obscured in mounting, and have

sometimes been overlooked by otherwise good observers. As a general

rule, they are best seen in the dry condition and without any special pre-
paration, but they are usually also seen in glycerine and water, especially
if examined when freshly mounted.

The number and position) of the germ-pores require to be carefully
studied, since they are often of specific value, and there are various re-
agents which show them up with great clearness. Among these, I have
successfully used lactic acid and Bismarck brown.

A 50 per cent, solution of lactic acid was used as a mounting medium,
then the spores were added, and the slide was heated over a spirit-lamp
just until the liquid reached the boiling point. Sometimes it is necessary
to boil for a little in order to get the best results, taking care not to
burst the spores, but as a rule bringing it to the boil is sufficient. After
the preparation has cooled, the cover-slip is placed over it. Some, use a
mixture of equal parts of glycerine and alcohol for mounting the spores
before adding the acid.

Boiling in lactic acid not only drives out the air and makes the spores
more transparent, but it swells them to their normal shape and size, and
not only brings out the germ-pores, but the surface markings as well. For
pale-coloured spores or spores which have partially lost their colour
through age, I have found nothing to surpass Bismarck brown. The
clear germ-pores show up well against the brown stain. The spores are
simply laid in the smallest possible amount of water or water and glycerine
on a slide, then thoroughly separated by stirring with a needle. Sufficient
of the stain is next applied, covered, and after gentle boiling the spores
are ready to be examined. It stains quickly, but does not overstain, and
is fairly permanent.

Soaking hard tissues for twenty-four hours in a solution of caustic
potash or caustic soda softens them wonderfully, and mounts made from
minute pieces of the material thus treated are often extraordinarily beauti-
ful, and the relation of the parts to one another is preserved in a surpris-
ing manner. Though soaking for several days nearly always destroys the
colour of the spores, yet details may sometimes be distinguished with
certainty not obtainable by any other method.

For determining the average size and shape of the spores I know
nothing better than photography. Numbers of spores are photographed
together, and the general shape can be determined by comparison while

guibiiea wirn a

the size is deduced from the exact measurement of individuals in large
groups. In this way the average size has been obtained for the teleuto-
spores chiefly, of all the species accessible for observation. At the aMU*
time a large number were measured direct by an eve-piece micrometer

same

and the results compared. All the microscopic investigations were made

IfvilllZ' aJ,0Phromatlc <*J«t»«, and the photomicrographs with lenses
or Watson s holoscopic series.

Spermogonia and Spermatid. 13

CHAPTER IV.

Spermogonia and Spermatia.

The spermogonium, sometimes called pycnidium, is a small punctiform
body, hemispherical to flask-shaped, produced beneath the epidermis or
cuticle, and ultimately the narrow neck bursts through in order to allow
the contents to escape into the. air. The narrow opening is generally sur-
rounded by a brush of sterile projecting stiff hyphae, the so-called para-
physes, and in the interior of the spermogonium there are numerous con-
verging hyphae surmounted by very minute oval or rounded bodies borne
in short chains — the so-called spermatia. These bodies were considered to
be of the nature of male sexual organs, hence the name; but whatever
may have been their original function, now lost in the course of develop-
ment, there is no reliable evidence to support the view of their being
sexual in function.

The spermatia are embedded in a sugary secretion, and though it is
not known whether this serves to cause the spermatia to germinate, insects
are probably attracted to the spermogonia sometimes by means of this
sweet bait, and also by their powerful and penetrating odour, as in Puccinia
suaveolens Pers., which is so named on account of its sweet scent. The
honey colour of the spots may also serve as an attraction, and the project-
ing hairs or paraphyses are believed to retain the spermatia and prevent
them being washed away.

Colour, scent, and honeydew will thus co-operate in alluring insects
to the spot, and the evident resemblance of these relations to those which
prevail in the fertilization of flowers by insects naturally led to the belief
of the spermogonia and spermatia being concerned in some, way with
the fertilizing process. This view was further strengthened by the fact
that in Lichens, which possess similar structures, there called pycnidia, a
true process of fertilization occurs. But actual experiments fail to prove
any such connexion, and the probabilities are that the original function
has disappeared owing to modifications consequent upon the fungus becom-
ing parasitic. Spermatia do not germinate in water, but do so when
placed in a suitable nutritive solution, such as white cane-sugar dissolved
in water. Germination consists in a minute prolongation at one end, which
ultimately becomes like the parent spore, and thus, as far as present know-
ledge goes, the spermatia are isolated structures, and do not enter into

the regular development of the fungus.

Spermatia, whatever may be, their function, do not occur as solitary

spore-forms, but always precede or accompany one of the others. They
are usually associated with aecidia, but this is not invariably the case,
for there may be aecidia without them, and in the absence of aecidia they
may be associated with other spore-forms, such as uredospores in the
case of Uromycladium robinsoni, Puccinia hieracii (Schum.) Mart. &c., and
teleutospores, as in P. liliacearum Duby and Uromycladium tepperianum
(Sacc.) McAlp., and either uredo or teleuto spores, as in U. maritimum.
The same mycelium which proceeds from fthe teleutospores, and produces
the spermogonium, also produces aecidio uredo or teleutospores, as the
case may be, although it may seem a needless waste of material and an
unnecessary act, since the one is left behind in the race, while the others

14 Spermogonia and Spermatia.

continue their further development. In a paper by Arthur3 on the Taxo-
nomic Importance of the Spermogonium, he shows that, in the first pla^t,
• he presence of the spermogonium, along with its associated spore-forms,
gives important information regarding the length of the life-cycle. Thus
he concludes that if spermatia and uredospores arise from the same
mycelium, aecidia do not occur, and if spermatia and teleutospores thus
arise, neither uredo nor aecidio spores will occur. There are cases, how-
ever, in which the spermogonia are found in association with both uredo
and teleuto spores, as in Uromycladium maritimum, U. notabile, and U.
robinsoni. When the spermatia are associated with aecidia, it is only where
teleutospores and aecidia arise from the same mycelium that it can be
definitely stated that there are no uredospores.

In the second place, the characters of the spermogonium, such as posi-
tion, size, form, and colour, and its relative position to the accompanying
spore-forms, furnish characters for positive identification, although of
minor value. There is an interesting relation of the spermogonium to
other spore-forms to be noted in the cases afterwards referred to, where
there is a repeated formation of aecidiospores, uredospores, or teleuto-
spores, as the case may be, and these spore-forms do not directly proceed
to their normal development. In such cases, as far as our present know-
ledge goes, the spermogonium is the only one which does not repeat itself,
as stated by Arthur3 : — "An observation more pertinent to our inquiry is that
the spermogonium occurs but once in the cycle, not being repeated with
each generation of repeating aecidia or uredo. In case the teleutospore
takes on the conidial function (repeating process), it is uncertain whether
the spermogonium is repeated with each summer generation or not. In
Puccinia malvacearum and similar Lepto-Uredineae, the spermogonium

seems to be wholly suppressed even with the first generation in spring.

The question of sexuality has been raised in connexion with the presence
of nuclei, but what constitutes an act of fertilization is interpreted differ-
ently by different authors.

Sappin-Trouffy2, for instance, has observed in Uredineae having the
various stages the following nuclear cycle: —

Uni-nucleate. —

Mature teleutospores.
Sporidiola.

Mycelium, producing spermatia and aecidiospores
Spermatia.

Bi-nucleate.

Aecidiospores.

Mycelium, producing uredo and teleutospores.
Uredospores and
Young teleutospores.

In the mature teleutospore the two nuclei fuse, and this fusion was
regarded by Sappin-Trouffy as an act of fertilization, but the fusion of
the nuclei may be interpreted otherwise. Blackman2 has confirmed the
above nuclear cvcle. and concludes that the spermatia are male cells which

have become functionless, the nuclear characters being those

d not of

The fertile cell of the aecidium or primary aecidiospore becomes
~ ~ by the nucleus of a neighbouring vegetative cell mizratin" th

the wall, and this association of the two nuclei Blackman considers to be
fertilization rather than .the act of fusion in the teleutospore, since the fertile
cell is stimulated to further development by the entrance of a nucleus from
without. — (Note 2, page 75.)

Sfermogonia and Sfermatia.

Massee also considers the

dium to be a sexual product, and both

authors agree that the spermatia are not concerned in it. If the spermogonia

d aecidia represented

and femal

productive organs, a distinct

alternation of generations would be present, the spore-bearing stage or
sporophyte commencing with the fertilized cell in the aecidium, and the egg-

bearing stage

or gametophvte starting

th the uni-nucleate teleutospc
But the most probable view is that the spermatium is a functionless org

d its presence is an indication that the

of th

may

possessed an

of generations similar to that referred to abo\

!6 Aecidia and Aecidiospores

CHAPTER V.

Aecidia and Aecidiospores.

The aecidia, or cluster-cups, as they are often called, are usually
brightly coloured, and attract attention not only from their colouration, but
from their elegant forms as well. The mycelium derived from the germ-
tube of a promycelial spore may first produce its spermogonia, and then
proceed to the development of aecidia, or aecidia may be succeeded by
aecidia. The spore layers are contained in a receptacle or pseudo-pendium,
which is formed from a simple layer of flattened sterile cells. It increases
in size by' the formation of new cells at its base, and is cup-shaped or
cylindrical, rupturing at the apex to allow the escape of the spores.

The aecidiospores are always unicellular, and are arranged in linear
series, arising from densely-crowded, erect hyphae or basidia in basipetal
order. The young spores are at first separated from each other, according
to De Bary, by sterile, intermediate cells, which are soon, however.
absorbed, and this makes it often difficult to prove their presence. From
the mode of their formation, from above downwards, the ripe spores are
at the top, and readily separate to be blown away, while the young spores
become polygonal from mutual pressure, lateral and lengthwise.

The. spores have usually a colourless membrane, with the exception of
those of Gymno sporangium, in which it is usually a deep brown, and in
the great majority of species are provided with points, spines, or warts, to
make them adhere in order to germinate. The granular contents are

coloured, as a rule,

orange- red oil, although this

also occurs in many uredo and teleutospores.

Germination takes place as in the uredospores by means of a simple
cylindrical germ-tube which enters through the stoma into the interior of
the host-plant. At certain weak spots in the membrane germ-pores are
formed, which are usually only distinctly visible at germination, owing to
the swelling of the membrane at these spots, and through these the endo-
spore is protruded in the form of a tube. As the aecidiospore germinates
very readily in water, the process of germination can be easily followed.
A germ-tube rarely arises from more than one pore, and the contents of the
spore gradually pass into it along with the orange-colouring matter, so that
the spore is ultimately emptied and the endochrome is towards the
extremity of the tube.

I found aecidiospores of Puccinia tasmanica, from the common ground-
sel (Senecio vulgaris), to germinate freely in the manner indicated. They
were very plentiful in November, and in a drop of water they began to
germinate within a few hours, and soon produced germ-tubes of consider-
able length.

As to the length of time that aecidiospores retain their germinating
power there is much difference of opinion. De Bary states that they may
retain it for some weeks, while Plowright considers it a matter of hours.
Eriksson, on the other hand, found that the spores of Aecidium berberidis,
for instance, were very capricious and uncertain in their germination, but
there is always the possibility that although the spores do not germinate in
water, they may infect a host-plant when brought into direct contact with
the living leaf.

According to Bolley, the aecidiospores on barberry and other hosts are
still capable of germination, even after lying in the herbarium for some
time, or being sent through the Dost.

Aecidia and Aecidiospores. 17

The development of the aecidia has been traced by De Bary, Neumann1,
and others. The hyphae derived from the promycelial spores form at
certain points,v deep down in the parenchyma of the leaf, little compact
bodies which have been called primordia by De Bary, because they are
the beginnings of the aecidia. These bodies gradually increase in size
by the rapid multiplication of the mycelium until they assume a spherical
form, only being slightly flattened at the part immediately beneath the
epidermis, and considerably thickened at the base. If sections aie made at
this stage the differentiation of the cells is seen to have begun. From the
dense basal layer of hyphae arise numerous, closely-crowded, short, erect
hyphae, somewhat club-shaped, and generally known as basidia. From
each of these erect hyphae an apical cell is separated off, and beneath that
another, and so on until a linear series is formed. At the same time, as the
basal cells give rise to the so-called basidia, the surrounding cells develop
into the external envelope or peridium. At. first they are absolutely indis-
tinguishable from the others, but after the third or fourth division they are
seen to be much broader. Then, simultaneously with the formation of the
first spores, they acquire their characteristic markings and polygonal form.
At first the peridial cells are filled with the coloured protoplasm or endo-
chrome, but the colour gradually disappears.

With regard to the peridial cells, Fischer1 has shown that in
Gymno sporangium the sculpturing of the walls may be used for the dis-
tinction of species, and while in the genera Puccinia and Uromyces these
cells are much more uniform in their markings, yet there is considerable
variation. Mayus1 has examined peridial cells from several species of
these two genera, and has found that within the limits of the same species
the nature of the peridium undergoes variation under the influence of
external conditions. These variations chiefly consist in the relations of the
lumen and the thickness of the wall, the lumen being relatively larger in
shady places and the converse in sunny places.

The typical aecidium possesses an external envelope of cells or peri-
dium, which surrounds the spores, and when this becomes ruptured at the
top to allow the escape of the spores it assumes the form of a cup. But
the envelope may assume different forms, or even be absent, and then
different names are given to it for convenience, although in its essential
character of spore -formation it is still the same structure. When the peri-
dium is elongated, and often horn-like, it is called a Roestelia, or it may
be rather irregular in shape and confined to coniferous plants, when it is
called a Peridermium (not represented in Australia), and when the peridium
is absent it is known as a Caeoma, as in C. apocyni.

Even in the genus Puccinia the pseudo-periclium is wanting in indi-
vidual species. In a number of species, as in Puccinia prenanthis, the
aecidium is sunken and formed by the more or less altered tissue of the
host-plant, and a special pseudo-peridium is either completely wanting or
very imperfect. Such aecidia are to be regarded as intermediate forms
between the true distinctly walled aecidia and the caeoma form.

The origin of the aecidium has given rise to a good deal of discussion,
and it is not universally accepted that it originates from non-sexual cells.
Massee1, in his paper On the -presence of Sexual Organs in Aecidium.
considers it to be a sexual product, and has even drawn the swollen ends
of two mycelial hyphae imbedded in the tissues of the host-plant, supposed
to be in the act of conjugation. Arthur, in his Problems in the Study of
Plant Rusts,2 and The Aecidium as a Device to restore Vigour to the
Fungus1 comes to the conclusion \that " the aecidium, with its accom-
panying spermogonia, represents the original sexual stage of the fungus,
and that it still retains much of its invigorating power."

Aecidia and Aecidiospores.

But it is quite possible ithat the time of appearance of the aecidia has
something to do with their invigorating power, if such exists. They
usually appear in the spring, when the first rush of vegetation commences,
and naturally the parasite shares in the strong growth of the host-plant.
The nature of the host will likewise affect the result. It may appear on
the leaves of an annual such as Helianthus annuus, which also bears
teleutospores, or on those of a deciduous shrub such as barberry, the
uredo and teleutospores of which occur on wheat and other grasses. But
it is a striking fact that the aecidium is so rare upon a grass that, until
the discovery of an aecidium on a species of Danthonia in Victoria
one instance was known, viz., Puccinia graminella, in which the aecidia
and teleutospores occur together, the latter being often very rare, or wanting
altogether.

Repeated Formation of Aecidia. — The normal development of heteroe-
cious fungi, as well as of autoecious forms, follows a regular cycle, as
already shown. In those forms possessing all stages in the life cycle, the
sporidiola produce only the aecidia, and the aecidiospores in turn give rise
only to uredo and teleutospores, as in Uromyces trifolii (Alb. & Schw.) Wint.
But among those autoecious species which produce aecidiospores and teleuto-
spores without uredospores, this regular succession of forms may be departed
from in some cases, and (the aecidiospores, provided the mycelium is not
perennial in the host, may repeatedly produce new aecidia before the
teleutospores are reached, as in P. senecionis

Lib

Spennatia

Sporidiola

Teleutosjx>res

Aecidiospores

fiQ rf Aecidiospores

This repeated formation of aecidia was principallv investigated bv
Dietel, who named the aecidia arising directlv from the germination of
teleutospores primary aecidia," and .those arising from the germination
of aecidiospores " secondary aecidia;" and it was noticeable that spermo-
gonia were usually absent from the latter. Thus the germinating spori-
diolum may produce a mycelium which bears onlv aecidia.: hut rt»* »JGu.

X^S,f'nStead °! givinS.rise dirffly t0 teleutospores, may repeat them
seUes for several generations, and then give rise to telentnsTvJ^ w™

give rise to teleutospores. H

complete cycle, the aecidia originate only from sporidioi

i"e™r asi the aecidia at first origWte from ^°ridioik' and

Uredospores

CHAPTER VI.

Uredospores.

Uredospores may originate from the hyphae developed from the germ-
tube of an aecidiospore, a promycelial spore, or another uredospore. Thev
are generally ovate or elliptic, and are developed singly on the ends
of separate short upright hyphae, known as basidia. In some
genera, however, such as Coleosporium and C/irysomyxa, they arise like
the aecidiospores, in short chains, and thus resemble Caeoma- forms — in
fact, they are sometimes regarded as such. The spore-bearing hyphae are
crowded together just beneath the cuticle, or epidermis, of the plant, and
such an aggregation is known as a spore-bed, or sorus.

The uredospores are always unicellular, and never smooth, the mem-
brane being beset with projections in the form of short prickles (echinulate)
or fine warts (verrucose). Two-celled uredospores have been described and
• drawn by Raze l and Jacky 2, in Puccinia c/irysantkemi, but they are
very probably monstrosities, as suggested by Sydow, two unicellular spores
becoming united when young, and growing up together. In the same rust
I found, in one instance, two uredospores produced on the same stalk, the
one slightly beneath the other ; but this was merely a freak. They differ
generally from the aecidiospores on the one hand in the mode of forma-
tion, and from the unicellular teleutospores of the genus Uromyces on the
other, in having two or more germ-pores, and this character also dis-
tinguishes them from t he. mesos pores. Only in exceptional rases is there
only one, as in Puccinia monopora. They vary in colour, generally being

some shade of orange or brown, and in the brown spores De Bary has
shown that, as in teleutospores generally, the colouring matter is in the
wall, and not in the contents.

Germination occurs similarly to that of the aecidiospores. When ripe,
and kept moist, a germ-tube is readily protruded through one or more
of the germ-pores, and this enters the host-plant by a stoma, and' in
the interior develops a mycelium like that from which it originated.

It is interesting to notice that in some cases the uredospores may be
produced, not onlv at the surface, but within the tissues. This happened
with Puccinia pruni, in a peach fruit, where spore-beds of rust freely
producing uredospores were imbedded in the tissue, in more or less
roundied cavities, up to 5mm. below the surface. The decaying fruit would
form a splendid matrix for preserving the spores till next season. Since
they are chiefly produced in the summer, and adapted, as a rule, for
rapid germination, they are often spoken of as summer spores, and as
soon as they arrive at maturity become detached from their stalks.

Repeated Formation of Uredospores. — Just as aecidiospores may- pro-

duce aecidiospores for several generations, so may uredospores produce
uredospores. This is well seen, for instance, in P. graminis. where uredo-
spores are produced direct from the uredospores without the intervention
of aecidia and teleutospores, as is the case in Australia, and this repeated
formation of uredospores may continue indefinitely. But there are several
cases where the first-formed uredospores are different from those produced
later, and, in order to distinguish such forms, the two kinds of generations
are known respectively as primary and secondary

Uredospores.

The primary generation appears in the early spring, and originates
ther by infection from the promycelial spores, aecidiospores, or, it may

Spennatia

Sporidiola

Teleutospores

Uredospores

Fig. 6.

he, from a perennial mycelium. The spore-layer is usually distinguished
' its larger size, and the corresponding injury it causes to the host

hile the secondary generation, produced by infection from the
preceding uredospores, has a smaller spore-layer.

A well-investigated and very striking case is seen in Triphragmium «
ulmariae, in which the primary and secondary uredospores generally re-
semble each other; but the sori of the former are much larger and pro-
duced in great abundance, occurring, on the stalks and midribs of the

hile those of

and scattered on the under sur-

face of the leaf. So striking is the difference that a special name has been
proposed for the sorus, epiteosporiferous and epiteospore for the primary
spore; but it is quite superfluous, since the larger and more prominent
sori of the primary generation may be accounted for from the strong
vigour of growth in 'the plant at the time when the first infection occurs.

The wintering of the rust-fungi, in the form of the uredo, depends
on the nature of the rust itself, and also on that of the host-plant. If
any portion of the host-plant remains green and succulent during the
winter, then the fungus has an opportunity of surviving, and it is thus
seen that climatic conditions have a deal to do with the persistence of
the fungus. When the winter is mild and green vegetation flourishes, the
mycelium of the rust fungus may continue to grow, and may even produce
spores; whereas, if the winter is severe, and the mycelium does not
remain in the perennial parts of the plant, then the continuance of the
fungus is likely to be by teleutospores, which can last through the winter
on dead stems or other decaying vegetable matter. This so-called winter-
ing of the uredo depends so much on the climate that in a mild climate
the fungus may perpetuate itself exclusively by uredospores ; whereas,
under severe conditions, it has to resort to teleutospores.

A very striking case is recorded by Lagerheim1, in Uromyces fabae
(Pers.), De Bary, which in Europe passes through the three stages of the
aecidio-, uredo-, and teleuto-spore, while in Ecuador it only produces the
uredo-form. Heteroecism, or alternation of generations, is an arrange-
ment suited to conditions where the seasons are variable, but in an equable
climate such as Ecuador there is no occasion to produce such a variety of
spore-forms, and so the fungus adheres to the one which serves its purpose
best. Australia is also a case in point as far as Puccinia graminis is con-
cerned, and it is necessary to remember that our seasons are the reverse of
those of the Old World. Our cereal crops are generally sown in the
autumn months of April and May, or even earlier, and the harvest is

Uredospores.

reaped at the end of spring, or beginning of summer in November and
December, so that it is the heat ajnd drought of summer, not the cold and
wet of winter, which the fungus has to provide against. In fact, the
wintering of the uredo is a misnomer here, for it is the excessive dryness
and heat which is most injurious.

Bearing this in mind, it is easy to understand that P. graminis, although
it still continues to produce a certain amount of iteleutospores, is per-
petuated from season to season by means of uredospores. Self-sown
wheat or oats, or even the aftermath of either of these crops cut for hay,
is always more or less rusty during the late summer and autumn, the
uredospores being freshly produced then through the depth of winter.

The teleutospores of P. graminis seem unable to infect the barberry in
Australia, and this heteroecious rust would appear ito be fast becoming
like Uromyces fabae reduced to its lowest Mmits, and reproducing itself

only by uredospores. Of course the absence of the barberry would tend

to weaken if not destroy the capacity to produce the aecidial stage.

Although the germination of uredospores during winter has already been
generally referred to, some definite instances may be given here, and
I will select those of Puccinia graminis, P. triticina, and P. chrysanthemi
from a number of tests made. The rust appeared on some self-sown
wheat, which was growing vigorously during winter ,(June), and on
placing the uredospores of P. graminis in a drop of water, they were
found to germinate sparsely in seventeen hours, and in twenty-one hours
they germinated freely and very generally. At the same time, and
from the same wheat plants, uredospores of P. triticina were placed under
similar conditions, and they also began to germinate within 21 hours,
but after several hours longer, only a few were germinating, and not too
luxuriously.

The uredospores of P. chrysanthemi were also taken from green leaves
in May, and they germinated freely, producing long curved germ-tubes.

Thus uredospores taken from growing plants during the winter are cap-
able of germinating, and this proves conclusively that self-sown, or
volunteer wheat, on the headlands or elsewhere in the neighbourhood of
growing crops is one of the means whereby rust may be continued from
season to season. In one case, which I have every reason to believe is
quite exceptional, the season's wheat, sown at Wellington, New South Wales,
in April, was badly rusted as early as May, but, as a general rule, it is
exceedingly difficult, even for the trained observer, to find more than an
odd speck of rust in a crop of wheat earlier than ,the end of September,
though there may be plenty on self-sown plants.

There is a conflict of evidence, however, as to the conditions under
which germination takes place when the spores are not taken direct from
the fresh and growing plant.

Eriksson (Eriksson and Henning1) found that the uredospores of P.
graminis lost their capacity for germination during the winter if exposed
to the weather, but retained it if kept inside, and even then it gradually
disappeared, while Jacky2 found that the uredospores of chrysanthemum
rust still retained their germinating power, after exposure to the weather
for 66 days, from 1st December to 5th February. And Miss Gibson kept
spores of the same rust in a dry test-tube in a cool room for 71 days, from
March to May, and at the end of that time about one quarter germinated,
while a week after none germinated.

The uredospore is primarily a spore for the rapid reproduction of the
species. As a rule, it is produced in immense numbers, it is provided
with a thin wall, having projections of some sort to act as a holdfast, and
it generally infests the leaf or sheath, so that nutrition is not directly

Uredospores.

interfered with, as in the case of ithe teleutospores on the stem. But
the uredo may become inured to unfavorable conditions, such as drought
or cold, and carry on the life of the species, independent of the teleuto-
spore. This is well seen in Puccinia poarum, for in both Europe and
America it has been found on the leaves of Poa pratensis even after the
melting of the snows, and in Australia it occurs on Poa annua throughout
the winter months, the rust disappearing with the withering of the host,
which generally happens early in October. An extreme case seems to
have been reached in P. vexans, Farl., where, in addition to the ordinary
uredo, there is a specialised form to which the special name of amphispore
has been given, which is thick- walled, strongly papillate, and only ger-
minates after a period of rest.

It is quite common for the uredo-layer to be attacked by the parasitic
fungus, Darluca filum, Cast., so much so that it has been found upon 24 per
cent, of the species of Puccinia. It is somewhat unfortunate that Dr. Cobb*
has confounded this parasite with spermogonia, producing spermatia, for
in referring to peach rust, he writes : — " I frequently find among the
uredospores of a pustule of this rust, small black pycnidia, producing a
multitude of two-celled spores, which, when placed in a moist chamber,
often bud and multiply after the manner of yeast plants, but which occa-
sionally produce a mycelium. These two-celled bodies have, as I have
on several occasions publicly remarked, no slight resemblance to the so-
called spermogonia of several species of Aecidium." Although spermatia
are well-known to be unicellular, yet Carleton 2 quotes this authority for the
statement that ordinary germ-tubes a're produced in the germination of
spermatia as well as in the other spore-forms. This rust parasite is very
commonly distributed, attacking the mycelium and probably checking the
development of spores. It occurs on aecidia, uredo, and teleuto-lavers,

and is recorded on Uramyces (9), Uromycladium (1), Puccinia (22), PJirag-
midium (1), and Aecidium (2).

Paraphyses most commonly occur in connexion with the uredosori, and

are found in Australian species of Puccinia, Melampsora, and Phragmidium
as well as in Uredo.

Occasionally they arise in both uredo and teleuto-sori, as in Puccinia
magnusiana and P. purpurea, and sometimes they are variable in their
presence as in Puccinia poarum, where Plowright found none in Britain,
although they are common in Australian specimens. In Phragmidium
subcorticium not only are the uredosori provided with paraphyses, but like-
wise the aecidial patches, since they are without a surrounding membrane.

The following are the known Australian species, with paraphvses in
tneir uredosori :— Puccinia lolii, P. magnusiana, P. poarum, P. ' pruniy
P. purpurea; Melampsora hypericorum, M. lint; Phragmidium bamardi,
P. subcorticium; Uredo kuehnii, and U. spyridii.

Teleutospores

CHAPTER VII.

Teleutospore s.

Teleutospores are very varied in their shape and size, and are on that
account often regarded as the characteristic form for distinguishing genera.
They may be produced directly from ithe mycelium of the aecidiospore or
uredospore, or indirectly from the teleutospore itself by means of the
sporidiola. As the name denotes, it is the last formed, or finishing spore,
in the life-history of rusts, although in many instances it is the only spore
formed, and there are cases where it has not yet been found. Where
it exists alone it may be that the other spore-forms originally existed, but
have now come to be dispensed with, and where it does not exist it may
be a degenerate type like the other, only it is the teleutospore form which
has been dropped. It is a question, however, whether it is not a necessary
stage in the life of every rust, and its apparent absence is simply owing to
our not having discovered it.

As an example, Uredo sym-phyii, D'C. was considered by De Bary to
be an independent species, and having lost its other spore-forms to be
capable of existing without them, but Bubak2 afterwards found the teleuto-
spore in Bohemia, and so it may turn out in other cases.

They arise like the uredospores in smaller or larger spore-beds, often
closely crowded together, and usually directly beneath the epidermis which
they often rupture. It is seldom that they originate directly beneath
the cuticle. The colour of the spore layer is very variable, but generally
it is darker than the uredo layer, being dark-brown to blackish, and only
,rarely reddish.

In the simplest cases teleutospores are unicellular, and originate in a
similar manner to the uredospores. The spore-bed, consisting of inter-
lacing and crowded hyphae, gives off erect branches, which become
swollen at their free ends, and the finely granular protoplasmic contents
are invested by an inner membrane, or endospore. in addition to the outer
or epispore, which becomes relatively thick and dark in colour. The
teleutospores are generally formed towards the end of the active vegetative
period of the host-plant, and are often called winter spores in contrast
to the uredo or summer spores. They are specially adapted and equipped
for continuing the species over periods of drought, or damp, or cold, or
seasons of scarcity. This is seen in the firm outer wall, which is often
sculptured in various ways, as well as in the reserve material stored up in
the contents. Sydow1 states that in all Leptopuccinieae, or those only
possessing (teleutospores which germinate at once, the epispore is perfectly
smooth ; but there is one exception in Australia — P. plagianthi. In other
groups the epispore may be smooth, warted, striated, &c.

The portion of the hypha supporting the spore becomes the stalk, or
pedicel, by means of which it (remains attached to the spore-bed for a
longer or shorter period.

In Endophyllum the teleutospores originate in chains, andl are produced
within a peridium similar to aecidia. They would be called aecidiospores,
only they produce a fouir-celled promycelium, which bears promycelial
spores. They serve the purpose of summer spores, since thev germinate as
soon as they are ripe, and the fungus winters by means of its mycelium
in the host-plant.

438.

Teleutosp

that it is now coming to be regarded

or I

independent form, but as a biologic genus in connexion with Puccinia

pores

Teleutospores may be simple, as in Uromyces, or compound as in

In the newly -constituted Australian genus, Uromycladium, the

of the Uromyces type, but they have the peculiarity of
being produced, not solitary at the end of a stalk, but in groups sometimes
accompanied by a colourless vesicle. In Uromyces and Uromycladium
there is only one germ-pore, situated: at the apex, and the membrane is
generally smooth, although it raav be warted or striated in Uromycladium.

As yet there are known only two Australian genera with compound
spores— /W/wa and Phragmidium, in the one case consisting of two spore-
cells, and in the other of three or more in a vertical row.

In Puccinia the germ-pore of the upper cell is at the apex, and that of
the lower at the side just beneath the transverse partition. •

In Phragmidium, the number of germ-pores varies in Australian species
from one to three in each cell. Dietel l has stated that there is only one
germ-pore in each cell of P. bamardi, but three were invariably found by
me in examining a large amount of material.

Paraphyses are not frequently found in teleutosori, probably because
teleutospores are generally so well constructed for withstanding variable

conditions that they do not require such protection. The best

ample is that of the old species, Puccinia rubigo-vera, now split up into
several, such as Puccinia bromina and P. triticina, in which the teleuto-
sori are divided into compartments by the clavate brown paraphyses. — (Note
3> P- 75-) It is worthy of note that the teleutospores are capable of ger-
mination in the autumn of the year in which they are produced. There are
no other Australian species in which paraphyses are confined to the teleuto-
sori, but they may occur in connexion with the latter as well as with uredo-
sori in Puccinia magnusiana, P. purpurea, and Uromyces phyllodiorum.

The germination of the teleutospores of Puccinia graminis in Australia
was tested under different conditions, and the most important condition

be the season of the year, for they

ger mini ate

during the spring months. Badly rusted straw was placed in the cool
stores for three months, one portion being kept at a temperature of
4 deg. C, and another at- 18 de*g. C. A third portion of the same straw
was simply kept in the ope»n, and when tested for germination in the spring
only the spores exposed to the weather germinated.

Another feature of germination worthy of mention is the way in which
it is spread over a period of time, and the spores in the different sori are
not all ready at once. There seems to be a succession of ripening, for
among a patch of sori only one out of every fifteen or twenty will be found
to contain spores capable of germination. This shows the necessity for
germinating spores in bulk when tests are being made, for you mig^it happen
to select spores which would not germinate, being taken from an unripe
sorus. Not only are the spores ready for their work of germination at dif-
ferent times, but the sporidiola are produced in succession, for you never
find the promycelium bearing its four spores all at once, at least in P. gra-
minis. This is well shown in Plate XV., with the germinating teleutospores

of P. malvacearum

Mesospores and Amphispores. 25

CHAPTER VIII.

Mesospores and Amphispores.

In the great majority of Australian Puccinias, 80 per cent, at least,
there occur associated with the teleutospores, and quite distinct from the
uredospores, unicellular spores which somewhat resemble the two-celled
spore in coloration, though generally much paler. It is generally notice-
able that wherever the teleutospores are thickened, or apiculate, or pro-
longed into processes at the apex, or warty on the surface, these spores

possess the same char

The conclusion one would naturally d

is that they are teleutospores in the process of making, with t|he lower
cell wanting, just a survival of what is normal in the Uromyces. From
their partaking of the nature of a middle spore-form between Uromyces and
Puccinia, they are generally called mesospores, and simply represent an
imperfectly developed or abortive teleutospore, which may, however, in

certain cases perform the functions of a fully-developed teleutospore,
although only one-celled.

In the newly-constituted genus Uromycladium there are found in con-
siderable numbers among the uredosori, smooth- walled spores smaller than
the uredospores, produced singly on basidia and entirely different from the
teleutospores, to which the term mesospores has also been applied.

Just as there are two kinds of teleutospores, so there may be two kinds
of uredospores, which are represented at present in a few species belonging
to Uromyces and Puccinia, but have not hitherto been found in Australia.
This modified uredospore, while agreeing with the normal uredospore in
the mode of germination, possesses a thickened epispore, and a more or
less persistent pedicel. From its partaking of the characters of both
spores, those of the uredospore in its possession of two or more germ-pores,
and those of the teleutospore in its germinating only after a period of rest,
it has been called an Amphispore, by Carleton.

Ampiii spore.

This peculiar kind of spore was first investigated in connexion with
Puccinia vexans Farl. This species has a true uredo and teleuto-stage,
in addition to a third form of unicellular spore, and the latter was the first
to be recognised and recorded. In 1879 it was described as Uromyces
brandegei by Peck, the unicellular spores being rough with minute warts,
and therefore suggestive of Uromyces. Then, in 1883, Dr. Farlow1
found true bilocular teleutospores associated with the supposed Uromyces,
and named the fungus Puccinia vexans, the specific name referring to the
perplexing nature of the unicellular spores. As Dr. Farlow writes: — "The
perplexing question arises, are the one-celled spores a unilocular form of
teleutospores similar to what is known as P. cesatii, Schr., or are they the
uredospores of this species?" Their true nature was finally settled in
1897, when Carleton succeeded in germinating them, and ultimately they
were found to give rise to two germ-tubes, as in the true uredospore which
had been discovered the previous year. Now that the three spore-forms
are known in this species, it becomes possible to answer the question, in
what respect does the amphispore differ from the teleutospore, on the
one hand, and the uredospore on the other. It agrees with the uredospore
in being unicellular, and having more than one germ-pore, but differs in

b 2

26 Mesospores and Amphispores.

being strongly papillate instead of echinulate, thick instead of thin-walled,
with persistent pedicel and only germinating after a period of rest. Its
essential difference from the teleutospore is its unicellular character and
the possession of more than one germ-pore. ^ 9

Besides the amphispores, mesospores are very numerous in this species,
so that there is a Puccini a provided with two kinds of uredospores, and two

sorts of teleutospores.

Up till recently this was the only instance known of the occurrence of
amphispores, but Arthur5 has given descriptions and illustrations of
nine different species, one of which belongs to the Uromyces, in which this
form of spore is met with. According to this author, amphispores are
mainly developed in arid or semi-arid regions, and represent a resting or
winter form of uredospores, being provided with thickened walls to enable
them to withstand unfavorable conditions, ius£ like a teleutospore.

Mesospores.

In examining the relatively large number of Australian species of Puc-
cinia possessing mesospores, one finds that as a rule they are comparatively
scarce in point of numbers, and that while a few may resemble the ordinary
teleutospore in colouration the majority are paler and altogether with an
immature appearance. But there are a few cases, such as Puccinia hetero-
spcra and P. simplex, in which the one-celled spores far outnumber the
regular teleutospores, so much so that the latter have been frequently over-
looked, and there is every probability that they undergo germination, and
are therefore unicellular teleutospores in the fullest sense of the term. In
P. heterospora there are no uredospores, and the unicellular spores are
smooth, and otherwise resemble the teleutospores, while in P. simplex the
uredospores are spinulose and yellow, and quite distinct from the smooth
unicellular spores associated with the teleutospores. Owing to the teleuto-
spores not having been obtained at first, the former has been variously
named Uromyces pulcherrimus, B. and C, U. thwaitesii, B. and Br., and
U. malvacearum, Speg., and the latter as Uromyces hordei, Rost. In fact,
these spores have been regarded as a transition stage from the unicellular
Uromyces to the bicelTular Puccinia spore. In such cases the one-celled
spore functions as a teleutospore, and ithere would be a certain convenience
in distinguishing between the undeveloped and immature spores and those
which are fully formed, and in all probability capable of germination,
reserving the term mesospore for the one, and unicellular teleutospore for

other. But it is so difficult to d

and

not desirable to multiply names unnecessarily, I will use >the term meso
spore to designate a unicellular teleutospore form in Puccinia and Uromy
cladium, which may either be imperfectly developed and incapable of ger
mi nation, or fully formed and germinable

The presence of mesospores in a species would seem to indicate its still
close relationship to Uromyces, and that its separation from the parent form
had not yet proceeded sufficiently far to obliterate every trace of its former

connexion.

Sporidiola or Promycelial Spores. 27

CHAPTER IX.

Sporidiola1 or Promycelial Spores.

When the teleutospore germinates, whether at once or afiter a period of
rest, the endospore is protruded through the germ-pore as a germ-tube,
and the contained protoplasm passes into it. This germ-tube does not
behave like that of the aecidiospore or uredospore, and elongate and branch
indefinitely, but it soon ceases to grow in length, and terminates blindly.
Hence it has received the special name of promycelium, because it directly
produces its spores. The promycelium is soon divided by septa, generally
stated as formed from above downwards, but mv own observations in
regard to Puccinia malvacearum show that the median septum is first
formed, dividing the promycelium into two, and then each of these
subdivides again into two, making in all four cells, from each of
which there is a short lateral protuberance which dilates at the end, and
becomes a promycelial spore or sporidiolum. The two upper cells, as a
rule, produce their sporidiola first, then the next, and lastly the lowest;
but occasionally .the two median cells start first. The name sporidium is
often applied to this spore, but since it is already used as equivalent to
ascospore, the present name has been proposed by Saccardo.

The sporidiola are easily detached, and, provided with moisture, they
can germinate at once, and on the surface of a living leaf the germ-tube
can pierce the epidermis, and, growing and branching in the interior, pro-
duces a mycelium similar to that derived from the aecidiospore or uredo-
spore. But in (the case of grasses the walls of the epidermis often contain
much silica, and this may be one of the reasons why aecidia are so scarce
in that family. Blackman 1 has shown that there is considerable variation
in the length of the promycelium, according to the conditions of growth.
Teleutospores, germinating in moist air, produce very short promycelia, and
form sporidiola almost immediately, while those germinating in drop cul-
tures, with their germ-tube submerged, grew to a length only limited by
the reserve material, but no sporidiola we're formed. On reaching the air,
however, their formation took place. The free air necessary to their for-
mation is correlated with their distribution by the wind.

If the teleutospore be regarded as the final stage of the cycle, then the
sporidiolum will be the starting point, and from that there may proceed in
regular succession the various spore-forms already enumerated, viz., sper-
matia, aecidio, uredo, and teleutospores, back to sporidiola again. All these
may occur in the same species, and the general course of development is
briefly as follows: — From the teleutospore in the spring, sporidiola are
formed which develop on a suitable host-plant a mycelium, from which
usually on the upper side of the leaf spermogonia are developed, and either
on the same side, but generally on the opposite side, aecidia are soon after-
wards produced. Infection by the aecidiospores produces the uredo-stage,
and these spore-forms often reproduce themselves. The uredo is specially
adapted for (the rapid spread of the fungus, since from the time of infection
up to the formation of new uredospores, only eight to ten days may
elapse. Finally, along with the uredo, or in special layers, teleutospores
arise which, on germination, produce again the promycelia and sporidiola,
and thus complete the course of development. This is the typical mode in
which the alternation of spore-forms occurs in many species belonging^ to
different genera, but there is often variation in the order, or even omission
of some of the stages. So constantly are aecidial or uredo stages associated

28 Sporidiola or Promycelial Spores

w;th at least another stage, that when found alone ot isolated they are
regarded as incomplete, and it is taken for granted that the associated form
has yet to be found. In the case of teleutospores, however, there are
numerous species which produce them alone, and in such cases, all the other
spore-forms are considered to have been suppressed or never formed. And
of these surviving teleutospores, in some species they are able to germinate
at once, and thus produce successive generations in the course of a year,
while in others they can only do so after a winter's rest, and are thus pro-
duced only once a year.

In the heteroecious rusts there is no evidence to show that the sporidiola
can infect the plant bearing the teleutospores, hence it would appear ithat
P. graminis in Australia can only be propagated, as far as spores are con-
cerned, by means of the uredospores. It has not been definitely proved
how the sporidiola are distributed, but no doubt the wind is an important
factor, and probably also animals assist in the distribution. In connexion
with the teleutospores of Gymno sporangium, Plowright2 says: — " It is prob-
able that the promycelial spores are implanted upon the ovary by insects
which had previously visited the Podisoma under the delusion that it was a
flower, and carried the minute spores with them to the hawthorn.' ' The
wind, however, is likely to be the common agency. Since the leaves and
stalks bearing the germinating teleutospores often lie upon the ground, it
might seem at first sight as if they were not favorably situated for the wind
to act upon them, but as they are usually produced in large numbers, suffi-
cient of the sporidiola are likely to be suspended in the air to infect fresh
plants when the conditions are suitable.

How long the sporidiola retain (their germinating power, and how far they
can stand drying up, are questions not yet satisfactorily determined.

•

Paraphyses.

CHAPTER X.

Paraphyses and their function.

Paraphyses are variously shaped — often hair-like, or capitate unicellular
bodies, sometimes accompanying the spores, and are just branches of the
hyphae not concerned in, but accessory to, reproduction. These sterile fila-
ments may occur in the spermogonia, where they assume the form of stiff
hairs projecting from the mouth, and possibly serve to retain the spermatia
until they are carried away and distributed by insects.

But it is in the uredo-layer that they are most commonly met with, and
there they are of various shapes. They are recorded in Australian species
of Puccinia, Phragmidium, Melampsora, and in Uromyces phyllodiorum,
and along with other characters may be useful in the discrimination of
species, as in Puccinia magnusiana, Koern., where they at once distinguish
it from the other species occurring on Phragmites communis. They usually
surround and arch over the spore-bed, and the apex is often swollen in a
globose or clavate manner.

Their principal function is probably the same as that of the hairs in
some grasses and other plants — to protect the spores when exposed by the
rupture of the cuticle against excessive evaporation and consequent drying up
during ,the day, and to moderate the temperature in the cold nights.

In the genus Melam-psora and some species oi Puccinia &nd Uredo, e.g.,
the uredo of P. poarum, P. magnusiana, and U. spyridii growing on hosts in
damp situations or along rivers and in moist valleys, the paraphyses have

always swollen capitate heads, and their function is suggested by their

structure and position. The wall is much thickened, so as to leave a very
small cavity in the head, and in the stalk it may become so thick as
completely to obliterate the cavity and thus render it solid. The
thickened head has a great capacity for retaining moisture, and since they
are crowded together and over-lap the spores, they will prevent them from
getting soaked and at the same time protect them against excessive evapora-
tion, which would have a drying effect according to Dietel9.

Their great function is as protective organs when the spores are exposed
by the rupture of the cuticle, and Plowright compares them with the
pseudoperidial cells of the aecidiospores. He has made observations on
the paraphyses of certain species, and found that their presence greatly
depends upon some special condition of the fungus. " I find them con-
stantly present with the uredospores of Puccinia perplexans, Plow., when
these have arisen, not directly, but rather at a considerable distance from
the aecidiospores. On the other hand, when the uredo arises directly
from the aecidiospore, they are hardly present at all ; this looks very much
as if they were an indication of exhaustion of vital energy on the part of
the fungus, which was combated by protective efforts on the part of the
parasite in conserving those spores which it does produce, but when full
of vigour and fresh from the aecidiospore it is less careful of its spores.
When it begins to feel the, effect of exhaustion, and is unable to develop
such energetic spores, it takes more care of those which are produced."

Or it may be that when the fungus begins to feel exhausted, it is
unable to develop so many spores, and barren protective filaments take the
place of those which would normally produce spores. If the paraphyses
serve to prevent excessive evaporation, then it would follow that in the
early spring, when there is little need for protection on this account, there
would be little use for them; but towards the summer, when the air is

3°

Paraphyses.

dry, they would be required in much larger numbers. The presence or
absence of paraphyses at different periods of the year seems to have been
only definitely observed in this one species, but there are other instances
which may possibly belong to the same category. Thus Winter describes
Uromyces dactylidis Otth. with capitate thickened paraphyses, and Plow-
right distinctly states that they are absent, and this discrepancy may arise
from the observations having been made at different seasons of the year.
Again, Puccinia poarum Niels, is without paraphyses, according to Plow-
right; but Schroeter found them in Germany, and I have found numerous
long ones in Australia. In P. magnusiana Koem. the clavate paraphyses
are of a dark smoky brown colour in the head and hyaline in the stalk.
They only occur at the margin of the uredo-layer, and there is a slight
indentation on the inner side, so that the head bends over. In Phragmi-
dium subcorticium the tubular, thin walled paraphyses are always mar-
ginal both in the uredo- and aecidio-spore generation, and are curved

inwards.

Another funct

of protecti
paraphyses

has been suggested

Mag

in add

that

In several species of Coleosp
ed both for protection and for

epidermis so as to make room for the growing spores

m he found that the
ng and bursting the

To a certain extent paraph

hastening its rupt

may assist in raising the cuticle

the absence of paraph v

but there are plenty of species which rupt

ted with teleutosp

mg awa\

until it decay

thev rather appea

and in the case of parapl

asso-

prevent the cuticle break

Origin of Sf ore-forms

CHAPTER XI.

Origin of the Principal Spore-forms.

The most characteristic feature of the Uredineae and that which renders
them specially interesting is the variety of spore-forms which they pro-
duce. But while on the one hand there is a regular succession of spore-
forms, on the other the number may be reduced even to a single kind, and
the question arises in such cases whether the missing spore-forms have
•dropped out of the course of development, or whether they have never been
formed. Since, as we shall see, there is every reason to believe that the
various spores originated from a primitive form, the natural conclusion
would be that where they do not occur they have not yet been developed,
yet there are cases where intermediate forms may have been suppressed,
judging by what obtains in closely allied species.

In all rusts whose complete development is known there is one kind of
spore which is invariably present, and which serves to distinguish the
various genera, and that is the teleutospore. But the teleutospore on
germination gives rise to another kind of spore, the sporidiolum, which
may be regarded as the starting-point of the life-cycle, just as the teleuto-
spore is the last-formed or finishing spore.

The simplest form of spore, using this term in its widest sense, was at
fust developed from any joint of the exposed hyphal filament, but ulti-
mately as the differentiation between the vegetative and reproductive por-
tions of the fungus became more marked, the point of origin was restricted
to a definite spot of an upright hypha — towards the apex. By a process
of abstriction, the end of the hypha was rounded off and detached by simple
contraction without the formation of any septum. These might either be
produced solitary or several formed in succession, constituting a chain.
(Fig. 7 a, b.)

a. b

Fig. 7.

The spore might also originate by budding, as in the case of the
Yeasts, where a small protuberance quickly grows to its full size, becomes
rounded off, and detaches itself. In this process of budding the pro-
tuberance often narrows' itself at the point of attachment into a slender
stalk, whereby the connexion with the parent cell is maintained until the
spore is fully formed. (Fig. 8.)

Fig. 8.

This short delicate stalk proceeding from the parent cell is known as a
sterigma, and either by abstriction or budding the origin of the various
spore-forms may be explained.

Origin of Spore-forms.

Sporidiolum.

The sporidiolum was probably the earliest form of rust spore, and
represents the transition from the saprophytic to the parasitic mode of life.
If we start from undoubted saprophytes, the passage from the one to the
other will be made clear. It was not only necessary for the spread of these
fungi that the spores should germinate rapidly, but that they should be
produced in sufficient numbers, and so the basidium, or parent cell, had
either to produce more than one spore or divide up into several cells. In
accordance with this, in one type, the one-celled basidium produces mostly
four spores (Fig. 9) ; in another type the basidium divides usually finto

Fig. i).

four cells, each cell producing a spore. The latter type is well seen in
the Auriculariaceae where in such a genus as Saccoblastia the basidia are
transversely septate, and each cell bears a sterigma with its spore (Fig. 10).

Fig. 10,

Turning now to a parasitic genus such as Coleosporium, there is a close
resemblance in the mode of formation of spores. The body called the
teleutospore is found to consist of four cells' placed one above the other,
and each cell gives rise to a sterigma, with a sporidiolum at the end of it
(Fig. n). This is something very different from the typical teleutospore,

Fig. 11.

in which each cell produces, not an ordinary undivided germ-tube, but a
promyceltum divided into four cells, each of which bears a sterigma with
a sporidiolum. The so-called teleutospore of Coleosporium is evidently
the representative of the septate basidium in the Auriculariaceae, although it
is generally considered to be an exceptional' form of teleutospore, which
occurs in other genera of Rusts as well, such as Ochropsora, Trichofsora,
and Chrysopsora. If the basidia in one of the saprophytic Auriculariaceae,
such as Saccoblastia ovtspora Moell., are compared with those of the
parasitic Coleosporium senecionis, there is seen to be complete agreement in
the structure. °

Origin of Spore-forms.

The sporidiolum was thus at first the product of a basidium arising
from a mycelium, but there was nothing specially characteristic in this, nor
any advance upon the mode of reproduction in a saprophytic fungusi. That
which constituted the Rusts a distinct class, and separated them from their
nearest allies, was the development of a new kind of spore, the teleutospore,
which produced directly on germination, without the intervention of any
mycelium, a basidium bearing sporidiola. In the one case the sporidiola
were derived from a basidium borne by a mycelium nourished on dead or
decaying matter ; in the other the basidium, or so-called promycelium, was
the direct product of a living spore. This spore is the special feature of
this group of parasitic fungi, and will now be considered.

Teleutospore.

In the sporidiolum the fungus is provided with a spore capable of
germinating at once and reproducing the original form, but a parasitic
fungus requires further to accommodate itself to the varying seasons of
grcwth of the host-plant, and so a spore that could persist during the
winter while vegetative activity was practically suspended, became a neces-
sity. Hence the teleutospore, or typical resting-spore, was introduced into
the cycle to provide a thick-walled form, which could withstand the vicis-
situdes of climate and be ready to germinate when spring, with its revival
of growth, returned. While some teleutospores are capable of immediate
germination, the great majority undergo a period of rest, and a thickening
of the wall is associated with this condition.

While the sporidiolum was thus' probably the earliest -formed rust spore,
it did not meet all the requirements of the new mode! of life, and thus the
teleutospore became a necessity for parasitic life, a condition dependent on
the changes of a living organism. The great feature of a teleutospore is
the mode of germination and its product, and perhaps the simplest form
is represented by that of Bar clay ella Diet, in which there are several cells
in a row, and each cell produces a promycelium or septate germ-tube. The
peculiarity and primitiveness of this promycelium lies in the fact that,
instead of dividing transversely and each cell giving rise to a sporidiolum,

the promycelium itself breaks up into four divisions, each of which becomes
a sporidiolum (Fig. 12).

Fm. 12.

The only species belonging to this genus occurs on a Conifer (Pice a
morinda Link) in the Himalayas, and neither uredospores nor aecidiospores
are known. It produces the most primitive form of teleutospore, and the
geim-tube produced from each cell breaks up directly into sporidiola, which
again reproduce the fungus. This is an evident contrivance for multiply-
ing the spore-form (teleutospore) and providing a fresh start with a
sporidiolum, minute, light, and easily transported by the wind.

The next advance in the development of the teleutospore may be seen
in an interesting genus, just described by Arthur6, to which he has given the
name of Baeodromus. It occurs on Senecio, and resembles Coleosporium

Origin of Spore-forms

unecionis so much in general appearance, that it was at first regarded as
that species, but the teleutospore produces a regular promycelium, with
sterigrnata and sporidiola, so that it well illustrates the transition from the
basidia bearing sterigrnata in Coleosporium to the ty pica teleutospore

^^celium. The teleutospores are united into a solid mass and
ate aw«w5ed in chains consisting of 5-8 cells in a series, and while Arthur
considers each cell as a teleutospore, the chain of cells might be regarded
« . „_^.lular teleutospore. The spores germinate immediately and
the promvcelia, together with the four globose sporidiola, have bright
orange contents (Fig. 13). Closely related to this form is Chrysotnyxa, m

prorm

as a mu

Fig. 13.

which the teleutospore consists of a series of cells, and on germination pro-
duces promycelia of several cells, each of which bears a sterigma with
sporidiolum. In Melampsora the unicellular teleutospores form a compact
mass, producing promycelia of the typical form, and thus a teleutospore
may either consist of a simple cell or a series of superposed <
A teleutospore is thusi a unicellular or multicellular spore,
germination a promycelium, which dither directly breaks up into usually
four sporidiola or divides into four cells, each of which produces a
sporidiolum at the apex of a sterigma. There is one exception to this in
the genus Endophyllum, in which the promycelium is the product of an
aecidiospore, but this may be regarded as a case where the function of a
teleutospore has been transferred to a derivative form, the aecidiospore.

producing

Uredospore.

The view that the uredospore is probably derived from the teleutospore
is favoured by the variability of the latter in many species, and the grada-
tions which are found to occur. At lirst sight the differences between the
uredo and teleutospore seem so great as to be insurmountable, but there are
distinct transitional forms from the one to the other. The membrane of the
typical uredospore is covered with spines, and this is an evident adaptation
for the spore which germinates immediately and is short-lived, and the
characteristic spines serve to attach it to the surface of the host-plant in
order that germination may be successfully accomplished. Magnus2, who
inclines to the view that the uredospores have developed out of teleuto-
spores, shows that in Uromyces scutellatus (Schrank), Lev., a gradation
can be traced between the reticulate or tuberculate membrane of the
teleutospore and the finely tuberculate or echinulate membrane of the uredo-
spore.

As to the thinning of the wall, there is also every gradation to be met
with from the thick brown membrane to the thin, almost colourless one.
The passage from the one to the other possibly took place through such a
form as the amphispore, a modified uredospore still capable of undergoing
a period of rest, and in which the wall still retains its thickness, but there
are several germ-pores. They may resemble the uredospores in shape,

Origin of Spore-forms.

echinulation, and germ-pores, but still they are resting-spores with thick-
ened walls. Fischer 1 has also illustrated the transition in a series of two-
celled teleutospores of Gymnosporangium confusum, which have all ger-
minated, and which show a dense thick membrane at one end of the series
and an excessively fine one at the other. And Dietel 3 has pointed out
that the occurrence of thick and thin-walled spores in species of Gymno-
sporangium is quite common, and that it is a character of the genus that
the teleutospores formed in the interior of the gelatinous mass are thin-
walled, while the external ones are thick-walled. The gelatinous substance
is formed from the gelatinous walls of the stalks of the teleutospore, and
readily absorbs the rain-water and thus facilitates the germination of the
spore. Another important difference between the uredospore and teleuto-
spore of Uromyces is that while the latter only possesses one germ-pore,
the former has two or more often arranged a9 an equatorial band. But
here again Magnus shows in species such as U. proeminens (DC.) Pass,
and U. tub er culatn s , Fckl., every transition from the. normal teleutospore
with apical germ-pore, through one with the germ-pore becoming somewhat
lateral, then with apical and lateral germ-pores, until the typical uredospore
is reached with a band of germ-pores confined to the equator, or several
scattered.

In Puccinia podolepidis there is often a germ-pore on either side of the
upper cell of the teleutospore, as shown in PL XXIX., Fig. 257. A third
and very important difference between the uredospore and teleutospore lies
in the mode of germination. Not only does the uredospore germinate im-
mediately on maturity, but it puts forth a germ-tube which penetrates
directly into the host-plant, while the teleutospore often undergoes a period
of rest and does not directly reproduce the fungus, but gives rise to a
piomycelium bearing promycelial spores, which germinate and enter a
host-plant. It would appear at first sight as? if this mode of germination
created a distinct barrier between the two kinds of spore, but when closely
looked into there is considerable variation in the germination of the teleuto-
spore, and under certain conditions it may forego the production of inter-
mediate spores.

Kienitz-Gerloff1 has shown that in Gymnosporangium clavariae forme,
J acq., the thin- walled teleutospores do not produce a promycelium and
promycelial spores, but simply a germ-tube like that of a uredospore. And
Dietel 3 has further shown that both thick and thin- walled spores may
sometimes germinate in this way, and thus serve the purpose of the uredo-
spore occurring in other genera. Fischer1 has also pointed out the influence
of external conditions on the mode of germination, the sporidiola only being
formed in air, while in water or in the interior of the gelatinous mass
surrounding the spores, an elongated germ-tube is formed. This has been
corroborated by Blackman 1, who found that the germ-tube is incapable of
sporidiola formation when submerged, and that under these conditions it
continues to grow in length until its reserve material is exhausted. He
also found that when germ-tubes of Phragmidium rubi were produced in
water, they might become divided into four cells, as if about to bear
sporidiola, but these cells, instead of following the ordina.ry course, rounded
themselves off and separated. Fischer observed in Gymnosporangium
confusum that the cells of the promycelium separated before forming sterig-
mata and promycelial spores, but Blackman, in a MS. note to his paper1,
kindly sent to me, says: "These are the cells of the promycelium which
are rounding themselves off and becoming directly sporidia." So that there
appears to be every gradation in the germination of the teleutospore, from
the production of a simple germ-tube to the division of the germ-tube into
cells which germinate, and finallv the promycelium bearing promycelial

-,<5 Origin of Spore-forms.

spores

,pu«« It is worthy of mention that Magnus2 has repeatedly obseived
that when the teleutospores of Puccinia graminis were germinated in water
thev produced an undivided germ-tube just like that of a uredospore, but
be could not satisfactorily settle the question, if this germ-tube could directly

penetrate the host-plant.

The passage from a thick to a thin wall, fiom one to a number o\
germ-pores, and from a typical promycelium to an ordinary germ-tube, has
been shown, and it all tends to support the view that the uredospore may
have been derived from a teleutospore.

This peculiar germination of the teleutospore, in which each promy-
celium breaks up into three or four detached cells', apparently representing
sporidiola, had been observed by Barclay3 as early as 1891, in Pucctma
lainiana, BarcL, and Uromyces solidaginis, Niessl. He considered this
abnormal mode of germination to be due to the restricted supply of air
obtained bv the spores in a hanging drop of water, for when allowed to

germinate in a watch-glass of water, with freer access of air, the g

germina-

tion was normal.

But this rounding off of the promycelial cells may be due to the influ-
ence of micro-organisms in the water. So far as our own observations
go, certain organisms are always present in great abundance when this
rounding off of the cells occurs. A similar phenomenon is observed in
connexion with the hyphae of numerous fungi growing in water or fluids
invaded by bacteria.

In another respect the uredospores show a distinct^ transition. In a
well -developed spore-layer they always precede the associated teleutospores,
but in Uromyces scutellatus, as Magnus2 has repeatedly observed, in
different years they may appear simultaneously.

The uredospores) may thus be regarded as having been derived from
the teleutospores, or from a mycelium common to both, and the question
naturally arises as to the origin of the aecidiospores.

Aecidiospores.

Aecidiospores and uredospores are often so much alike that they are
mistaken for each other, and sometimes there is a difference of opinion as
to whether a spore-form should be regarded as one or the other, as in
Triphragmium ulmariae; but, generally speaking, aecidiospores are produced
in chains, and uredospores singly on evident pedicels. In Coleosporium and
Chrysomyxa, however, the uredospores are developed in chains as well as
the aecidiospores, and this renders necessary the further distinction that the

aecidiospore always precedes the uredospoie in point of time.

In Phragmidium subcorticium, for instance, the aecidiospores were not
recognised at first as distinct from the uredospores, but although the indi-
vidual spores resemble each other closely, the fact that one is produced in
chains and the other not, distinguishes them. When aecidiospores are pro-
duced without any special envelope or pseudo-peridium, as it is called, it
is distinguished from the Accidium proper as a Caeoma, and there is every
gradation from naked to covered aecidia. In Chrysomyxa the uredo is re-
garded as a caeoma-form by Raciborski but in PJtragmidium the aecidio-
spores are protected by a dense layer of paraphyses, which surround them,
and thus take the place of a peridium. Triphragmium ulmariae (Schum.)
Link, has what are called primary and secondary uredospores, although
the former are described by De Toni as aecidiospores, but they are not pro-
duced in chains. Winter regards them as biological representatives of the
aecidium, and Drs. Milesi and Traverso1 speak of them as epiteospores, to
distinguish them from caeomospores, which are arranged in chains. There

Origin of Spore-forms.

is no essential point of distinction between aecidiospores and uredospores,
for though the former are always formed in chains, yet undoubted uredo-
spores may also be thus produced. Hence there is no valid distinction
between the two, even to the matter of nuclei, for both are binucleate.

The aecidiospore is just like the uredospore, thin-walled, and adapted
for immediate germination; but since it is essentially a spring form, and
required to keep pace with the rapid growth which then takes place, it is
not formed solitary upon a stalk, but tier upon tier, to make abundant
provision for the coming season. It is usually uredospores and aecidio-
spores which are confounded together, but there is one case at least in
which the aecidiospore partakes of the character of a teleutospore. In
Endophyllum the spores are produced in chains, and within a pseudo-
peridium just like normal aecidia, but instead of germinating in the usual
way, each one produces a four -eel led promycelium, giving rise to promy-
celial spores just like a teleutospore. They may either be described as
aecidiospores which produce promycelia or as teleutospores resembling aeci-
diospores. This genus is not now generally regarded as independent, but
as related biologically to Uromyces or Puccinia, and it will probably turn
out to be a case where an aecidiospore still retains marks of its earh origin
from a teleutospore in its mode of germination. (Note 4, p. 75.)

Spermogonia and Spermatia.

If the aecidiospores were the iresult of a sexual process, as is some-
times maintained, then of course they could not be derived from teleuto-
spores, and the mere fact that I have attempted to account for their origin
from this source shows that some other explanation must be forthcoming
for the so-called male sexual organs — spermogonia and spermatia. In

lichen-fungi such as a Collema a. true process of conjugation occurs, and
the male organ or spermogonium, with its contained spermatia, resembles
very closely in structure the similarly named bodies in the rusts. It was
Tulasne who originally discovered these bodies in 1851, and who suggested
their sexual nature, which he based partly on the fact that the spermatia
were not known then to germinate, and partly that they usually preceded
or accompanied the bodies they were supposed to fertilize, viz., the aecidia.

Great attention has, therefore, been paid to the spermatia, in order to
discover if they were capable of germination, and it has been found that
in a nutritive solution they grow and bud after the manner of yeast, but
no definite mycelium has been produced.

It is generally stated that the spermogonia either precede or accompany
the aecidia, but they may occur with all the spore forms, according to the one
which is first produced. Aecidia usually follow the germination of the
sporidiola, and therefore spermogonia accompany them most frequently : but
if the first formed spore is the uredo, as in Tripliragmium ulmariae, Uromy-
cladium maritimum, and Puccinia obtegens (Lk.) Tub, then they accompany
it, and if a teleutospore as in P. liliacearum, Duby. or Uromycladium tep-
perianum, the spermogonia are associated with it. But Arthur3 states the
case more generally when he remarks that " every one who has made cultures
of the rusts knows that in about a week after sowing the germinating teleu-
tospores there will appear spermogonia, without any regard to the kind of
spore that is ifo follow." There are even instances where the spermogonium
has entirely disappeared, as in Puccinia malvacearum, Mont. It is ac-
knowledged by those who have given special attention to the subject that
the spermogonium is an isolated organ, of uncertain origin and function,
and that the balance of evidence is against its being a sexual organ. The
spermogonium is seldom absent from the life-cycle, and vet it takes no direct

3»

Origin of Spore-forms.

part in the reproduction of the fungus. Its meaning seems to have been
lost, but its origin from the sporidiolum seems to indicate that it may be
the survival of the conidial reproduction of the sporidiolum.

Unless in cases where the mycelium is perennial, the sporidiolum ger-
minates and produces a mycelium, which gives rise to the spermogonium.
The same mycelium may either produce aecidia, as is usually the case, or
in the absence of aecidia, uredospores ; or, if both are absent, the teleuto-
spores, as in Uromycladium tepperianum. The spermogonia are never pro-
duced alone, but always precede or accompany some other spore form, for
the very good reason that they are incapable of reproducing the fungus.
When the spermogonium is about to be formed, the hyphae become inter-
woven, and form a tangled mass in it'he sub-epidermal tissues. From this
numerous delicate branches are given off, which are directed towards the
epidermis. These branches converge towards a central point and form a
somewhat round or piriform body, which is only covered by the cuticle at
the top. The periphery of this body consists of a dense felted mass of
sterile hyphae, quite unlike the peridium of the aecidia, which are likewise
always deeper-seated than the spermogonia. The so-called spermatia arise
inside the spermogonia in short chains by a process of abstriction at the
end of fertile converging hyphae, and are held together by a gelatinous sub-
stance which contains a certain amount of saccharine matter. (PI. XXIII.,
Fig. 203.)

The probable origin of this puzzling body may thus be accounted for :
We may suppose that the sporidiola were spread principally by wind and
rain, and at first only produced teleutospores. But in order to secure the
intervention of insects which were now becoming adapted to the floral
world, a new kind of reproductive body was developed, and the sper-
matia abstricted in chains were produced in spermogonia with a sweet
bait to attract insects. But when the teleutospores gradually developed
uredospores and the same mycelium could produce the three spore-forms —
spermatia, teleuto, and uredo spores — then the former became less and less
necessary. When finally the aecidiospores were developed, produced in gieat
abundance, and also in chains, then the competition was too keen, and
the necessity for spermatia practically ceased to exist, since the germinat-
ing sporidiola could produce the aecidia, and the germinating aecidia both
the uredospores and ithe teleutospores. Besides, the bright colour of the uredo
and aecidio spores may serve to attract insects for purposes of distribution,
and so they were able completely to replace the- spermatia. Finally, the
spermatia lost their capacity for germination, and now the spermogonium is
an organ which has survived its function, and only remains as a land-
mark to show what once had been the prevailing type of reproductive body
adapted for distribution by insect life.

We can thus picture to ourselves the progenitors of the rusts leading
a saprophytic existence and gradually adapting themselves to the new
mode of life when the parasitic habit was developed and a modification in
the spore-forms occurred. The evidence of this great change is shown in
the development of a spore primarily adapted for undergoing a period of
rest, and from the stored-up material directly producing a basidium with
sporidiola, instead of being preceded by a more or less scattered mycelium.

The peculiarity of the teleutospore lies in the fact that it is an in-
dependent body capable of giving rise directly and without anv further

food supply to fresh spores by the production of a septate germ-tube,
which develops secondary spores unlike the original, being smaller, thin-
walled, and ready to germinate at once on the damp surface of a living leaf
In course of time the teleutospores became differentiated into forms
adapted for extremes of temperature and resting, as well as other forms

Origin of Spore-forms

adapted for favorable conditions and rapid dissemination of the species —
uredospores. The third form of spore would probably be developed later
in point of time. The aecidiospores were at first just like the uredospores,
only produced tier upon tier as required, and the great purpose served by
all these different kinds of spores would be to provide a continuous succes-
sion of spore- forms — aecidiospores in the early spring, when the first rush
of growth commences; then uredospores during spring and summer, when
steady growth is maintained ; and finally teleutospores towards the autumn,
capable of remaining dormant during the winter and starting the whole
series again in the spring by means of promycelial spores.

The so-called spermogonium does not enter functionally into the present
cycle, being a relic of the past and a survival of the time when insects
were being catered for by the rust-fungi in order to aid in the distribution
of their spores ; but the development of such numerous and varied bright-
coloured spore-forms has rendered unnecessary this special form of fructi
fication.

Variability of Teleutospores.

When the teleutospores in the different genera of rusts are carefully
examined, it is often found that there is not only considerable variation in
the size and shape, but the seemingly constant character of the number of
cells is departed from. It is not always convenient to record this in a
systematic description, and I have selected a few species to give an indi-
cation of what is not at all uncommon throughout the Uredineae (PL XL.).
This variability not only shows how the uredospore in each genus might
have been derived from the teleutospore, but also how the multicellular form
of spore might have originated from the unicellular. Starting with the
genus Uromyces, there is no difficulty in showing how the bicellular spore of
Puccinia may have originated as in Uromyces orchidearum, U. tricorynes,
and U . vesiculosus (Pis. XVI., XVIII., XL.) Then in the genus Puccinia
there is not only the unicellular teleutospore or mesospore to indicate its pro-
bable origin from the Uromyces, but in addition to the two-celled spore
there are three and four celled spores, in which the cells are arranged in a
manner suggestive of various genera. The different forms seen in a single
preparation of P. dicJiondrae Mont, are shown in Plate XL., and there are
at least seven forms met with — (i) the unicellular spore, or mesospore,
generally with thickened apex, and resembling the uppermost cell of the
next form; (2) the typical bicellular teleutospore; (3) the three or four-
celled spore, with the cells arranged in linear series, superposed more or
less regularly, and after the Phragmidium type; (4) a three-celled spore,
consisting of one basal cell supporting two longitudinally divided on top,
and resembling the Triphragmium tvpe; (5) a three-celled spore, consisting
of two basal cells longitudinally divided and a single cell on top, as^ in
Hapalophragmium', (6) a four-celled spore with the two lower superim-
posed, and the two upper longitudinally divided ; (7) a four-celled ellipsoid
spore longitudinally andi transversely divided, as in Sphaerophragmium.
It is worthy of note (that there are no uredospores in P. dichondrae, but,
even in P. ludwigii, where uredospores are present, there^ is also a con-
siderable amount of variation. A two-celled spore is met with here, having
the septum longitudinal instead of transverse, and the pedicel in a line
with it, as in Diorchidium. P. graminis and P. triticina also show a
considerable amount of variation, if large quantities of material are
examined. Other genera exhibit variation of a similar kind, and there
is a basis here provided for natural selection to work upon, and evolve
the various forms which are used to characterize the different genera of the
Uredines.

40 Relation of Rusts to other Fungi.

CHAPTER XII.

Rusts in their Relation to Other Fungi.

The Fungi, as a class, are generally regarded as having descended
from the Algae, but since the latter possess the green colouring matter
or chlorophyll which enables them in the presence of sunlight to abstract
carbon from the carbon dioxide of the air, they do not seem to be the
most primitive forms. But the fission-fungi, such as the nitrifying bac-
teria occurring in the soil, are able, in the. absence of light and chloro-
phyll, to split up carbon dioxide and obtain the necessary carbon likewise
from inorganic material, so that the first forms of life to appear upon the
earth could thus obtain their nourishment without organic compounds at
all. The development of the chlorophyll would thus occur at a later
period, and the fungi proper, as well as the algae, may have had a common
origin from these primitive bacteria, instead of the one being a degenerate
form of the other.

Following the fate of the fungi, with which we are more immediately
concerned, their course of development ran parallel with that of the algae,
so much so that they have been regarded as degenerate algae or algae
without chlorophyll. This primitive stock resembling the algae so closely
is known as Phycomycetes, and from this divergence has taken place
in two directions, the offshoots representing two main divisions of fungi.
In the one case the Ascomycetes or fungi producing spores in delicate
sacs or asci, and in the other, the Basidiomycetes or fungi producing naked
spores on large terminal cells known as basidia. To this latter division
belong the Uredines or rusts, since they produce basidia which are trans-
versely divided, and bear naked spores; but they occupy a low position
as compared with the higher Basidiomycetes, including the mushrooms
and toadstools. In the Ustilagines or smuts closely related to the rusts,
the basidia are not as yet definitely fixed, since the spores are produced
at any part and new ones are developed when the old ones fall away,
which is not the case in the rusts.

The following arrangement will show the position of the rusts in this
scheme of classification : —

Fungi.

f Ascomycetes

Phycomycetes (Alga-like Fungi) \ ^ .,. 4 (Hemibasidn (Smuts)

J J 6 b ' | Basidiomycetes ^ Protobasidii (Rusts)

I [Holobasidii (Mushrooms, &c.)

Starting from the Phycomycetes or alga-like fungi, there is one divi-
sion of them which bears both sporangia and naked spores, and another
in which the sporangia may be wanting. The former would give rise to
the Ascomycetes, and the latter to the Basidiomycetes, which exclusively
reproduce themselves by naked spores.

The smuts are generally regarded as stepping-stones from the Phycomy-
cetes towards the rusts, which have become more closely identified with
the true Basidiomycetes, where the basidia are entire and not divided.

The above gives a very general' idea of the position of the rusts among
the fungi, and indicates briefly, without entering into detail, how they
may have originated.

The parasitic habit of the rusts will account for several features in
their life-history as well as in their structure. Being dependent on other

Relation of Rusts to other Fungi.

plants for their sustenance, they must accommodate themselves to their
surroundings, and so they may pass through different stages,

sometimes

the same plant, or on different

These various stag

presented

bv different reproductive bodies, enable the rust to produce spores which

can germinate

if food suppl

are

ail able

others which

rest if need be. The change of host is

dent advant

*nlv

from the point of view of a change of diet, but it may thus pass from

dies down to another that is perennial. It mav

perennial itself in

the

dergiround parts of some

gn become
d then it

vegetates, only producing
be formed under normal (

arely the reproductive bodies which would
ditions.

42 Indigenous and Introduced Species.

CHAPTER XIII.
Indigenous and Introduced Species.

In a large continent like Australia, where so many plants have been

introduced, both for economic and ornamental purposes, it is not alway
easy to determine what rust-fungi are native to the country, and those that
have been introduced on imported plants. A plant may be indigenous,
and yet the rust upon it may have been derived from an allied species, as
in the case of P. thuemeni on native celery (Apium prostratum) which was
possibly introduced with cultivated celery {Apium graveolens), P. menthae
on native mint introduced upon cultivated mint, and Melampsora
lint, on native and cultivated flax. On the other hand, species
of rust occur on well-known imported plants, such as the daisy, groundsel,
and marigold, which are not recorded elsewhere, and the natural conclusion
is that they are indigenous, or have been overlooked elsewhere. ^ It is only
in rare cases that the first introduction of any species of fungus is observed
and accurately determined, so that we must fall back upon some well
recognised principle to settle whether a rust is indigenous or not. Where
a new species is found on a native plant, it may be taken for granted that
it is indigenous, and even where the species is already known, but the plant
has a wide distribution, such as Phragmites communis, then there is no
reason to doubt that a rust upon it, such as Puccinia magnusiana is also

indigenous. The great

ded are, of course

and it will only be necessary to single out those which have, in all
probability, been imported firom other countries.

The following species may be regarded as having been introduced on
the grounds indicated* : —

Puccinia anthoxanthi Fckl. on Anthoxanithum odoratum (1896).

P. arenariae (Schum.), Schroet. on Stellaria media (1896).

P. beckmanniae n. sp. on Beckmannia erucaeformis (1904).

P. chrysanthemi Roze, on Chrysanthemum indicum (1904).

P. cichorii (DC.) Bell., on Cichorium intybus (1885).

P. cvani (Schleich.) Pass, on Centaurea cvanus (1904).

P. festucae Plowr. on Festuca pratensis (1903).

P. graminis Pers. on Wheat, &c. (1825).

P. helianthi Schwein. on Helianthus annuus (1887).

P. hypochoeridis Oud. on Hypochoeris radicata (1889).

P. impatientis (Schw.^ Arthur, on Elymus condensatus (1903).

P. lolii Niels, on Lolium perenne (1896).

P. malvacearum Mont, on Malva, &c. (1857).

P. maydis Bereng. on Maize (1880).

P. menthae Pers. on Mentha laxiflora (1884).

P. poarum Niels, on Poa annua (1890).

P. prenanthis (Pers.), Lindr. on Lactuca sp. (1892).

P. pruni Pers. on Prunus sp. (1883).

P. purpurea Cooke, on Sorghum halepense and S. vulgare (1892).
P. simplex (Koern.), Eriks. and Henn. on Barley (1902).

P. thuemeni (Thuem.) McAlp. on Apium graveolens and A. pros-

tratum (1892).

P. triticina Eriks. on Wheat (probably 182^).

Uromyces appendiculatus (Pers.). Link; on Vigna catjang (1905).

U. betae (Pers.), Kuehn, on Beta vulgaris (1878).

The year in brackets indicates when first recorded for or observed in Australia1?

Indigenous and Introduced Species. 43

U. caryophyllinus (Schrank), Schroet. on Carnations (1896).

U. fabae (Pers.), De By. on Beans (1898).

U. polygoni (Pers.), Fckl. on Polygonum aviculare (1896).

U. trifolii Alb. and Schw. on Trifolium repens (1892).

Phragmidium subcorticium (Schrank), Wint. on Rosa sp. (1892). •

Melampsora lini (Pers.), Tul. on Linum usitatissimum and L.

marginale (1889).
Uredo kuehnii Krueg. on Sugar-cane (1893).

It would be interesting to trace from what quarter these species found
their way into Australia, but from 'the very nature of the case, it is
impossible to tell exactly, except in a few instances, and one can only
make shrewd guesses as to the rest. The inquiry would mainly resolve
itself into the importation of the host-plants, either by cuttings or seed, and
the seed of such weeds as chick-weed (Stellaria media) and knot-weed (Poly-
gonum aviculare) might easily be carried in straw packing or in hay.

Mr. Ellery, F.R.S., late Government Astronomer, has pointed out that
the scarlet pimpernel (Anagallis arvensis) was first observed as an intro-
duced weed in the Observatory grounds, and he informs me that as many
English and African plants appeared there at different times after unpack-
ing cases from the different countries, he came to the conclusion that seeds
from the packing were the source. Both uredo and teleutospores of
Puccinia graminis have been found on wheat straw envelopes on bottles of
wine imported from France. — (Note 5, p. 75.)

In the case of the cereals and grasses, the rust spores would probably
be brought with the seed, and this is certainly true as regairds Puccinia
beckmanniae. The seed of Beckmannia erucaeformis Host, was forwarded
to me in 1903 by the United States Department of Agriculture, and on
growing it at Leongatha, the rust was very copiously developed in
February and March, 1904. E. D. Hoi way informs me that it is known
in Minnesota, U.S.A., although it has not hitherto been published.

The seed of Elymus condensatus was also sent from America in 1903,
nd the rust {Puccinia imfatientis) appeared upon the plants in December
of the same year.

The latest addition to. our imported rust-fungi is that of Puccinia
chrysanthemi, and it was observed for the first time in New South Wales
in 1904. Cuttings had been imported from England, and thus the rust
was carried ; but it has not as yet spread very much. Only the uredospores
occur here as in England, and I was able to germinate them freely in
tap-water towards the end of May. It is by means of this trade in
cuttings that the rust is likelv to be spread, unless proper precautions are
taken. The history of its distribution is rather interesting. It is pro-
bably indigenous to Japan, and in 1895 it first appeared in England, then
in France in 1897, and about the same time in Denmark and Germany.
In 1900 it reached America and Switzerland, and now, in 1904, or earlier,
it has come to Australia, probably by way of England. It was also re-
corded for New Zealand in 1904.

The mallow rust (Puccinia malvacearum) has been known in Australia
since 1857, when it was found in the neighbourhood of Melbourne, and
now it is one of the commonest rusts we have. It is remarkable for the
rapidity of its spread, nearly oveT the whole world, since it was first de-
scribed in 1852 from Chili, where it is indigenous. After its appearance
in Australia, the next record of it is in Spain (1869), then France (1872).
England and Germany (1873), Italv (1874), Switzerland and Cape of Good
Hope (1875), Austria and Hungary (1876), Greece (1877), North

America (1886), Sweden (1887), and even reaching Finland (1890).

44 Indigenous and. Introduced Species.

The hollyhocks on which this rust occurs are ornamental plants,
and the disease may have been spread in the ordinary course
of trade or exchange. The first record of a rust is by no means a
guide to its first appearance, for it is generally only when it has become
established and has proved injurious that it attracts attention. The prune
rust, Puccinia pruni, which now occurs in all the States, was first observed in
Queensland in 1886, and was recorded for Victoria in 1883. Although con-
fined at first to certain districts, it has since then spread considerably, and
as settlement increases it becomes more widespread^

The flax rust, Melampsora lini, was first determined on some cultivated
flax from South Australia in 1889 by Galloway of the Bureau of Plant
Industrv, U.S.A. Bolley, in a letter dated 29th December, 1904, informs
me that' it is a very abundant rust upon all the wild varieties, and is always
more or less destructive in the flax crop. It is common enough here on
the native flax, and was probably introduced with flax seed.

There are four species of Phragmidium in Australia a genus confined
to the Rose family, and only one of them is supposed to have been intro-
duced. Phr. subcorticium only occurs on the imported genus Rosa, and
was probably introduced in rose cuttings, since the mycelium of the aecidium
is known to winter in the stem.

Phr. potentillae on species of Acaena was determined by Winter, and*
although referred by him to this widely distributed species, it is probably
new. Phr. longissimum was first discovered at the Cape of Good Hope,
and is now known to occur in other parts of Africa. Its appearance on a
native Rubus in Queensland would seem to support the generally accepted
opinion of a former land -connexion between Africa and Australia. But
Wallace in his Island Life offers an alternative view. " We should prefer
to consider the few genera [of plants] common to Australia and South
Africa as remnants of an ancient vegetation, once spread over the Northern
Hemisphere, driven southward by the pressure of more specialised types
and now finding refuge in these two widely separated southern lands."
From the shape and arrangement of the teleutospores and their germination
immediately on ripening, this species stands apart from ithe others belonging
to this genus, and Dietel 13 draws the conclusion that it separated at a very
early period from the common stem of the genus Phragmidium, a conclusion
which harmonizes with the views of Wallace.

But the most interesting case of distribution is that of Phr. barnardi,
which is not confined to Australia as was formerly believed. Quite the
same type has now been found in Japan on the same host-plant (Rubus
parvifolius), and, according to Dietel 13, it is simply a variety of the Aus-
tralian species, having fewer cells in the teleutospore, and therefore dis-
tinguished as variety pauciloculare.

A number of plants are common to Eastern Asia and Australia, and
R. parvifolius is included by the late Baron von Mueller in a list of plants
which extend from Eastern Australia to Japan. There are various ways
in which the species may have attained to its present wide distribution,
which is given as Malaya, China, Japan, and Australia. Birds may have
carried the seeds, and with it some attached spores of the fungus to the
Asiatic continent, or inversely from Japan to Australia. There is also the
possibility of a former land-connexion between Australia and Asia, which
is assumed by the zoologists, and at that time the two forms of Phr. bar-
nardi may have existed. The flora of Japan, like that of Australia, is
regarded as being of the same character as that of the Tertiary period, so
that the wild raspberry and allied plants had plenty of time to spread
from a point to the north of both Australia and Japan, carrying with them
to their new homes, the rusts already developed upon them.

Indigenous Species with their Hosts. 45

CHAPTER XIV.

Indigenous Species with their Hosts.

The great majority of Australian rusts complete their life-history on one
and the same plant, and are thus autoecious ; but, although heteroecism, or
the division of the life cycle into two generations, each on different host-
ants, has not yet been proved for any of them, still it has been so well
established for several in other countries, that we may accept it for the
present as likely to hold good here. As to the recognised indigenous species
which are also heteroecious, there are only four — Puccinia agropyri, P.
agrostidis, P. magnusiana, and P. caricis — the three former on Gramineae
having their aecidial stage on Ranunculaceae, and the latter on Cyperaceae
with its aecidial stage on Urticaceae.

If we arrange the indigenous species of rusts known in Australia under
the different families of their host-plants, which are further classified ac-
cording to their predominance, as determined by the late Baron von Mueller
(Table) some interesting deductions may be made ; but it must always be
remembered that the number of known species is probably far short of those
actually existing. Under these circumstances our conclusions can only be
partial, still, even with these limitations, it will be instructive to compare the
predominance of the native host-plants with that of the native rusts. It
is found that the greatest number of rust-species occurs on the families of
native plants which are large in point of numbers. Thus the Leguminosae
with the greatest number of species have eighteen different rusts, while the
Compositae, which only stand fourth in the list, have seventeen. At the
same time it ought to be noted that future discoveries may alter this rela-
tion, for no less than seven species of the new genus, Uromycladium, have
been added to the Leguminosae within /the last few years.

The 'Cyperaceae, which succeed the Compositae, have only four species,
while the Gramineae, which come next, have thirteen species. Then the
Liliaceae have seven species and the Rubiaceae five ; but on the remaining
families they vary from one to five. The Leguminosae and
Compositae have eighteen and seventeen species respectively, the Gramineae
coming third with thirteen species. The grasses and composites are gene-
rally herbaceous, quick-growing plants, with succulent leaves, and
the rust-fungi can most readily penetrate their tissues and secure during the
growing season sufficiency of food. They would also be guided in their
choice bv the chemotactic nature of the substances contained in the host-

plants ; but there are so many factors which enter into the choice of a host-
plant by a rust that we -can only mainly at present note their preferences.
Confining our attention now to the species of Puccinia alone and compar-
ing them with the numbers and distribution as given in Sydow's Mono-
graph, it is found that while one-fourth of all the species inhabit Com-
positae and one-eighth occur in Gramineae, so with the native Puccinias in
Australia more than one-seventh belong to the Compositae and one-
ni'nth to the Gramineae. In the Leguminosae only one species has been
met with, and only fifteen species are recorded altogether.

Indigenous Species with their Hosts.

GO
GO

O eg

a) C a.

*0 © ce

3 5'^

D I DP

•-

= := -

= •§•2

P >/E

• • ;

=8*

o *

d >

-a o

3 5 S

a*
©

•

5 xx *•- °*^

U>PDPPDP

S
©

a>.£

o ft© —

d WT3 71

SlJfa

DDDD

.2 ©.:

ill

&5=>P

- — •

.59

• P^

*-3

micrc

orchi

thely

•

- — -

£' ©

d O ^ wji

~^— ©.d .i; ©.3

- — ^ "3 ^ ft © ft*3 d

©+*.d

OOP

— . ^— ©
ofi eC t-
© © ©

to fc

r2 5P © O
O ft

Sis

COS

cS-T-

ftft^aH^ftft^ft^ft^-

m ft a

'Ct3

ft CO

o o

tt be

s B £ 3 g^ ft

P»i3 a X 5

Oflp a ft od

^ftO, anOiClHpHft^^ft^^

c3 $8

ft
S

c5 ce

Pl^PhPhPh

r
O

ce
©

■ —

Indigenous Species with their Hosts

co cd i

a "So

JS O co

3 3 ,d £>

d © o d

• • •

ppp

•73

ft
CO

2
3
o

(-4

cd
cd

'3b

CD
CD

<x>

<tj^

co d
d >>

■a

.71

•3

CD Oi

ftftOn

.2 CD

is ca

u cd

p£

ftft

CD ©.3

2 rt 2
2 d co

P s °

^ 03 ^

p. bfi cd

0^3 ft

5 o o

• • •

ft ft ft

d 2

08 .2

jo ,d

2^
ft^

on d

£.2

X! ft

ftft

• I—

CO
C3

d
o

—
o

_Z v-

h.s

uct

£3

03 O

• •

•

ftft

ft ft

-2

■ —

CD
CD

CO
CD

CD
03
CD
CD

o
d

CD '

CD
CD

-3

■H

»-

«}

03
CD

-
CD

ft
O

d
o

CD
CD

O
O

CD
03
—
CD

CD
CD
CO

-ZI

can

icoi

Indigenous Species with their Hosts.

as
o

fa
O

fr-l

2Sg

rfl

£ a*
fa>

S rr

75*0

1 5

fc»8

& ©

—

o 2

fafa fa

o
o

8 !

•

fafa

• — ^

-I

c3 c3

s *

D ©

fafa

E'S

— . en
S £

u o

fafa

fa°

.-5

C
e2

>> '-= 3

—

-5

a
a

3 e

a
O

c3
©

Indigenous Species with their Hosts.

It is worthy of remark that on some of our most predominant families,
such as Myrtaceae and Proteaceae, which are only exceeded by the Legu-
minosae, the rusts are practically absent. It is passing strange that upon
our numerous Eucalypts and kindred species not a single rust-fungus
should have developed, while in the Proteaceae, with their wonderful variety
of foliage, only a single species, and that a Uredo, is recorded. Melamp-
sora eucalypti Rabh. found in Calcutta on the leaves of Eucalyptus globulus
is merely a name, and the specimens show no indications of a rust. I have
carefully examined the original specimen in Rabenhorst's Fungi europaei
2592, and while the leaves have numerous blister-like swellings over them,
they are found to consist of discoloured cells, the epidermal cells parti-
cularly being brown and discoloured, and might superficially be mistaken
for spores.

In the preceding table only indigenous rusts are given which occur on
native host-plants, but such rusts may either be confined exclusively to
native plants or they may occur on other allied introduced plants growing
here as well. Thus Puccinia tasmanica is found on the introduced weed
the common groundsel {Senecio vulgaris), but one stage of it is also found
on native species of the same genus, and,

The rust may also be found on native plants, though in other
parts of the world on allied species, and yet be regarded as native, as
in the case of P. perplexans and P. agropyri. There are even cases
where native rusts, or at least rusts not known elsewhere, are confined
exclusively to imported plants, as P. calendulae , P. cinerariae, and P.
distincia. As regards introduced rusts, they may be found on both
native and imported plants, the presumption being \that the rust spread
from one to the other, though some may incline to the view that these are
as much natives of Australia as of any other country. Disregarding
Puccinia graminis, there are four such species, P. menthae, P. malvace-
arum, P. thuemeni, and Melampsora lini. There is still another group
consisting of introduced rusts found here on imported plants alone, such as
Puccinia chrysanthemi and Phragmidium subcorticium.

therefore, it is regarded as in-

digenous.

5°

Australian Distribution.

CHAPTER XV.

Australian Distribution.

It would be premature to attempt to arrange the Australian rusts in

geographical districts, since they are as yet too imperfectly known

large

Kg „_ have not been explored sufficiently to give any exact idea as
w the number of species or the distribution of those already known.^ So
for the present I will content myself with indicating their distribution in
the different States, and this may lead to a filling up of many of the gaps,
when it is seen what species may exist in one State, and are probably to be

iound in some of the others.

There is one evident way in which our knowledge of this group might
be extended and the area of distribution made better known. They depend
for their existence on the occurrence of suitable host-plants, and since
these are given for each species, as far as known, wherever the host-plants
are to be found, there the parasites might be looked for.

The total number of species at present recorded is 161, and they are
distributed among the different genera as follows :

1905. 1892.

Uromvces ... 27 ... 13

Uromycladium ... 7

Puccinia ... 90 ... 24

Phragmidium •.. 4 ... 4

Cronartium ... 1 ... 1

Melampsora
Roestelia

• •

Caeoma ... 2 ... o

Aecidium ... 15 ... 16

Uredo ... 13 ... 10

161 72

It will be seen that the Puccinias constitute more than one-half of the
whole, and the Uromvces come next.

In Cooke's Handbook of Australian Fungi, published in 1892, there
are only 72 recorded, or less than half the number, and even some of these
do not stand the test of further investigation. In Melampsora, for instance,
there are three species given, and two of these must be withdrawn, one
belonging to another species also recorded, and another not being a rust
at all. And in Sydow's Monograph, just completed for the Puccinias,
only 43 are given for Australia.

The following list shows the distribution of species in the different
States, and it is naturally very unequal. In Victoria, which heads the list,
there has been a zealous band of collectors stimulated into activity by the
late Baron von Mueller and encouraged by a progressive and active Field
Naturalists' Club. In Queensland the Government Botanist has always
been most enthusiastic in working up the Fungi generally, and in New
South Wales and Tasmania good progress is being made. Although South
Australia does not possess an official Government Botanist, that State is

fortunate in having such a zealous Botanist as J. G. O. Tepper, F.L.S

Australian Distribution.

•

who has given special attention to this group, and has not only <

described

some species himself, but has had several named in his honour,

such as

Puccinia tepperi and Uromycladium tepperianum.

•

No. of
Species.

Victoria.

Now South
Wale*.

Queensland.

South
Australia.

WCRt

Australia,

t • •

Tasmania.

Uromyces

Uromycladium

Puccinia

7
90

7
69

3
26

7
1

6
6

Phragmidium ..
Cronartium . . .

4
1

» • •

m • •

• • ■

• • •

Melampsora . . .
Caeoma

2
2

2
1

• • •

...

• • •

Aecidium

" *5

# •

Uredo

• • •

Totals

161

5°

Parasitism.

CHAPTER XVI.
The Origin and Specialisation of Parasitism.

In a group of parasitic fungi like the rusts the question naturally arises,
How did this parasitic habit originate? Parasitism in fungi is an adapta-
tion whereby the fungus can directly draw its nourishment from the living
material, and in order to do this it must have become accustomed to a
new mode of life, for it is assumed that originally the fungi obtained the
requisite substances for food from dead or decaying organic material.

The saprophytic mode of life seems to have led up to the parasitic, tor
there is everv sort of gradation between the two. Some parasites are able
to complete their development entirely on artificial nutritive media. Others
require to infect the living plant first, and then undergo their final develop-
ment on dead tissue, while others begin their life on dead material, as a

eparatory stage to passing over to the living substance.

The origin of parasitism and that of specialisation are so intimately
bound up that they may be conveniently considered together, for at the
critical moment, when the spore first put forth its germ-tube into the living
tissue of a particular host-plant and was able to grow there, then parasit-
ism was established, and if the fungus confined itself to that host then
specialisation had begun.

What induced the fungus to enter the living plant by means of its
germ-tube, and afterwards confine itself to one or a few closely-allied
species is the question to be answered. It does not seem difficult to account
for the entrance of the germ-tube into the stoma, for it follows the lines
of junction of the cells, and ultimately comes to a stoma, into which it
dips just as it would into any other opening. But to be able to penetrate
the cells and abstract nourishment from them is the point which requires
explanation.

It is assumed that the saprophytic habit was the normal one among
fungi, and that parasitism is an acquired habit. Massee9 claims to have
proved this assumption to be a fact, for he says — " A saprophytic fungus
can be gradually educated to become an active parasite to a given host-
plant, by means of introducing a substance positivelv chemotactic to the
fungus into the tissues of the host. By similar means a parasitic fungus
ran be induced to become parasitic on a new host." Parasitism, then, is
due to chemotaxis, which is a form of sensitiveness in the plant whereby it
has an affinity for certain substances, and is opposed to others. Thus there
are various substances which are capable of attracting or repelling the germ-
tubes of fungi, and the name of positive or negative chemotaxis has been
given to this property.

In an extensive series of experiments conducted with both parasitic and
saprophytic fungi, Massee9 has shown that certain substances in the plant
are positively chemotactic in their nature, and others negatively so. Thus
it was found that sugar is the most general of positive chemotactic sub-
stances, although its aation on the germ-tubes of obligate parasites is very
slight. Experiments showed that " it was not sufficiently powerful in any

instance to attract the germ-tubes through perforations in mica or through
stomata."

Specialisation of Parasitism.

^Recent investigations in connexion with heteroecious rust-fungi, or those
which change their hosts and produce a different kind of fungus on each

Parasitism

host, together with the results of infection experiments, have considerably
modified our views as to the limits of species in such fungi.

Eriksson l, in dealing, with cereal rusts particularly, found that they
were not liable to infect indiscriminately the different cereals, but were
confined to one, or, at most, a few closely -allied host-plants, and to this
phenomenon, so widespread among parasitic fungi, he applied the appro-
priate name of specialisation.

As examples of specialisation among heteroecious rust-fungi may be
given those of Puccinia coronata, Corda, and P. graminis, Pers. Klebahn *
proved by infection experiments that the crown rust on Dactylis glomerata
and other grasses only produced its aecidium on Frangula alnusy while that
on Lolium perenne required for its aecidial host Rhamnus cathartica. Hence

old

be separated bv mor

phological characters. Eriksson1 also proved in 1894 that the well-
known and much-investigated species of Puccinia graminis could be split
up into a series of forms, all of which agreed in producing aecidia on the
barberry, but differed in the uredo and teleutospore generations, only being
able to infect special host-plants. Puccinia dispersa, Eriks., was proved

to be an independent species, with uredo and teleutospores on rye, and its
aecidia on species of Anchusa. Included in this were a number of forms

which had no known aecidial stage, and they were afterwards separated
and raised to specific rank as P. triticina, P. bromina, P. agropyrina, &c.

• If a general view be taken of this phenomenon it is found that when
two closely-related species, say, A and B, are attacked by a rust-fungus,
the one on A will not infect B, and that on B will not infect A, even although
the two fungi are the same species, regarded from a morphological point of
view. There must, however, be some adaptation between the host and the
fungus, so that the latter is attracted towards the one host and repelled by
the other. But it has been shown by Ward 1 that occasionally a spore from
A may gain a footing on B, and once having done this it can continue to
infect B, since it has now become adapted to it.

According to the same observer, parasites may be educated to attack
fresh plants by means of what he calls bridging-species. Thus, while the
parasite on A may be unable to infect B, it mav be able easily to infect
a related species C, and after establishing itself on C it may then have
the power to infect B, so that C becomes the bridging species from A to B.

Massee9 has also shown that a parasitic fungus can be led to attack a
new host-plant by injecting a substance positively chemotactic to the fungus

into the tissues of the living leaf.

Parasitism is thus an acquired habit, and, generally speaking, it be-
comes specialised, because only in certain plants are the substances present
which attract the fungi, while in others there are also certain substances
which repel, and thus prevent their germ-tubes gaining a footing in the
tissues.

But in contrast to this specialisation, there occurs in a few species what
may be called general parasitism, where the parasitic fungus is able to in-
fect host-plants widely separated in their affinities. Fischer3 and
Klebahn1 have shown that Cronartium asclepiadeum can attack plants
belonging to such distantly related families as Ranunculaceae and Scrophu-
lariaceae, as well as Asclepiadeae, so that it has become necessary to unite
under this name, species which were formerly separated on account of the

difference of host-plant.

Chemotaxis can hardly be held accountable for such a widely-divergent
distribution of host-plants, and although it does not clear the matter^ up,
xt may be referred to the " internal developmental tendencies ot
Klebahn x until a better explanation is forthcoming.

Parasitism.

The specialisation of parasitism resulting in the evolution of biologic
forms is not confined to the Uredineae, but probably extends to parasitic
fungi generally, which frequent more than one host, and this has been
experimentally proved, particularly in the mildews or Erysiphaceae. Seve-
ral biologic forms may occur within a morphological species, so that it
will be necessary in the future, for the proper understanding of any such
species, not only to determine its limits by means of structural characters,
but also the special forms included in it with restricted powers of infec-
tion. Hitherto it has been generally assumed that the same parasitic fungus
occurring on two closely related host-plants would be mutually infective,
but this does not necessarily follow, even with different species of the
same genus.

Neger1, in 1902, proved by numerous experiments that there were
biologic forms of Oidium for several species of Erysipke, and Marchal1
in the same year divided the one species of E. graminis into seven distinct
forms, using only the conidia for purposes of infection. He showed that
the biological form on barley was unable to infect wheat, oats, and rye,
and Salmon 2 carried the experiments a stage further by using the asco-
spores, which had the same restricted powers of infection.

This difference in infective power is not due to any apparent structural
change in the fungus, for the form of E. graminis on the wheat is indis-
tinguishable, even under the microscope, from that on the barley, and yet

the form on barley cannot infect the wheat, nor can that on wheat infect
the barley.

The difference, therefore, lies in the physiological peculiarities of the
host-plant, and it has been suggested that the cells of the leaf contain

an enzyme which is fatal to the growth of the haustorium of any other
form. But probably the action is reciprocal, and the germ-tube of the
fungus finds something in the particular host-plant which attracts it, and
is conducive to its growth.

Heteroecism

CHAPTER XVII.

Heteroecism and its Origin.

In a great many species the various stages of the self-same fungus,
as already indicated, occur on the same host-plant, but the variety in
the mode of reproduction has also brought about a variation in the mode
of nutrition, for there are a number of species in which one part of their
life is passed upon one species of plant, and the remainder on a totally
different species. The host-plants are not even related to each other, but
stand far apart in their natural affinities. Those which passed their entire
existence on one plant have been called autoecious species, while those
which spread it over different plants are called heteroecious species.

As a general rule it is assumed that the different forms of rust occur-
ring on the same host-plant are genetically connected, although it is always
desirable, where possible, to have experimental proof of it. In Uromyces

folygoniy for instance, the three stages of aecidio, uredo, and teleuto-
spores may all occur together on the same leaf, or the aecidia may occur
on one portion of the plant, and the uredo and teleuto sori on another ;

and in both cases the species is regarded as having three stages, which are
different forms of the same fungus. It is but a step further to have, say,
the aecidia on one host-plant and the uredo and teleuto stages on another,
and this affords a greater variety of food supply. Just as in the separa-
tion of the sexes in flowering plants, we are justified in assuming that all
the different stages occurred at first on the same host-plant, but gradually,
in the struggle for existence, one reproductive body matured on one
and the others on a different plant, so that a change of food was secuied
and a succession of crops insured.

This heteroecism of the rusts was first discovered by De Bary in 1864,
when he proved that the rust in wheat, Puccinia graminis, produced its
uredo and teleutospores on the Gramineae, while its aecidial stage
developed on the barberry.

It has thus been assumed that heteroecious species originated from

autoecious species in the simple and seemingly natural way ,that the two
generations separated, just to occupy fresh ground, at first passing over
to nearly allied plants, and gradually to plants further and further re-

moved in the natural system, until the present position of affairs was
brought about that the two generations of the same fungus attack plants
widely removed from each other, as far as their natural affinities are
concerned.

But there is no evidence to prove that such a gradual separation took
place, for even although the species of Puccinia on Phalaris may have
their related aecidia on other Monocotyledons, and the Uromyces on the
pea, with its aecidia on a Euphorbia, yet they are always considerably
remote from each other. In fact, ithe view that heteroecism originated
suddenly and without the different generations slowly and gradually pass-
ing from plant to plaint, seems to have most in its favour. It is admittedly
a difficult problem, since from the very nature of the case no one has been
able to observe an autoecious fungus becoming heteroecious.

Heteroecism is only possible when more than one spore-form occurs in
the life-cycle, and how the variety of spore-forms originated is capable
of different interpretation. It may either have been a progressive develop-
ment from the simplest forms, or it may have been a retrogression from
the most highly developed forms. We may conceive heteroecism to have

438.

Heteroecism.

proceeded from the formation of teleutospores being succeeded by aecidio
spores, presumably at first both arising from tr - — u"™ N~* J

Next a

of labour took place, and the, mycelium of the aecidia was pro

C»

| • • •

dSSSV thJT^idSi, VhYle the mycelium of the teleutospore proceeded
from the aecidiospore. The advance to heteroecism took place when the
aecidlospores produced their mycelium in one host-plant and the teleuto-
spores, through the sporidiola, in another, and the kernel of the matter lies
in the answer to the question, How did this come about ? It may either
have taken place bv a long series of slow and gradual changes, whereby
the different spore-fonms gradually accustomed themselves to the new mode
of life or it may have developed suddenly by one of the spore-forms
terminating and growing on a different host-plant, and continuing to do
*>. But this latter view is hardly borne out by some experiments conducted
by Miss Gibson1, in which the aecidia from different host-plants were used
to infect Ranunculus ficaria, and while, the germ-tube as a rule entered
the stoma freelv, it was generally dead and shrivelled by the third day.
This result was not supposed to be due to starvation, for she says
" Whether the incapacitv to penetrate the cells is due to lack of attractive
substance or to the presence of anything actively repellent is not clear
though, as before stated, certain facts seem to suggest the presence of
something harmful to the hyphae."

A few concrete examples mav be given to show how far these views are
borne out by facts.

De Bary considered the probable origin of three species of Chrysomyxa
occurring in the Alps, and the relation existing between them : — C. rho-
dodendri (DC.) De Bary, forms its uredo and teleutospores on species
of Rhododendron, while its aecidiospores occur on Picea excelsa, the name
given ito this form before its connexion was discovered being Aecidium
abietinum, Alb. and Schw. C. ledi (Alb. and Schw.) De Bary, forms uredo
and teleutospores on Ledum falustre, and its aecidia also on Picea
excelsa, there being little or no distinction between them and those of
C. rhododendri. The third, C. abietis (Wallr.) Ung., forms the same kind
of teleutospore on Picea excelsa, but the sporidiola from the germ-tube
produce mycelia which only form teleutospores and no aecidia or uredo-
spores have been observed. In seeking to account for this, he assumes
a common origin of the three forms, and considers that either the original
form from which they were all derived had no aecidial fructification to
start with, or there was an aecidial fructification, and C. abietis has in
course of time dropped it. The latter view is the one he favours. We
can imagine these three forms competing for Picea as an aecidial host,
and while two succeeded in establishing themselves, the third, C. abietis,
was compelled to drop it altogether.

Barclay2, in tracing the developmental history of Uredineae,
attempted to show that in the struggle for existence, heteroecism was
beneficial, and that if two species compete against one another for a host,
that which makes for heteroecism will more probably succeed than that
which makes for autoecism.

There is another interesting series of forms worthy of consideration
known as " coronate " rusts, because the apex of the teleutospore is pro-
longed into and crowned by a number of finger-like processes. There are
both heteroecious and autoecious species as follows : —

i. P. coronata, Corda, I. Frangula alnus, II., III., Grasses.

2. P. lolii, Niels., I. Rhamnus cathartica, &c, II., III., Grasses.

3. P. himalayensis (Barcl.), Diet. I. R. dahurica, II., III., Grasses.

Heteroecism

4. P. festucae, Plow., I. Lonicera periclymenium, &c, II., III.,

Grasses.

5. P. mesneriana, Thuem. III., Rhamnus alaternus.

6. P. digitata, Ell. and Hark. III., R. croceus.

7. P. schweinfurthii (P. Henn.) Magn. III., R. staddo.

8. P. longirostris, Komarov III., Lonicera hispida.

9. Uromyces phyllodiorum (B. and Br.) McAlp. O., II., III.,

Acacia notabilis, &c.

Fischer considered that the original forms of the heteroecious species lived
both on the grasses and Rhamnus as autoecious fungi, and that they could
undergo their complete development on either of them. These original
forms were thus supposed to be able to live on a variety of hosts, and it is
assumed thait only in recent times had they become specialised. In support

of this view there is a rust — Puccinia graminella which produces both
aecidia and teleutospores on a grass, and I have also found an aecidium

on Danthonia, and although teleutospores were not observed on the same
plant, I still regard the two as belonging to the same species, viz., Uromyces
danthoniae.

Fischer presupposes that the original form was both autoecious and
plurivorous, and it is reasonable to suppose that a fungus which could live
upon two such distinct hosts as Rhamrtus and grasses would, at the same
time, select numerous other plants as hosts, so that this view hardly explains
the fact. lit is much simpler to suppose that since these primary forms had
begun to form aecidia and eventually uredospores, they no longer carried
out their complete development on their original hosts, since a change of
host was in a sense equivalent to a cross in flowering plants. They accord-
ingly changed their teleutospores (and uredospores) to new hosts, so that
the autoecious stage was dispensed with because it was not so advantageous
as the other.

This change of host is not a haphazard affair, but takes place according
to a definite plan. lit may be confined to a single or' a few distinct species,
and attempts to bring it about on plants which do not belong to the regular
cycle, as a rule, end in failure.

But, of course, the legular host plants may fail, owing ,to drought or
some other climatic conditions, and then the fungus often obeys the law of
self-preservation, by repeating the same generation again and again. This
may even become a fixed habit until the single generation is more or less
independent, and then it is all that remains of what was once a complete
cvcle.

Among the heteroecious fungi there is a regular course of development
which is usually followed. The one host-plant bears the aecidium genera-
tion, and the other host-plant the uredo and teleuto spore generations, but
there are slight differences in detail which may be noted here.

The complete cycle of development, as already stated in the introduc-
tion, is the most common, in which the teleutospores germinate in the
spring after a winter's rest, and produce sporidiola. The sporidiola infect
the young leaves of the proper host, and produce aecidia, usually accom-
panied or preceded by spermogonia. Then the aecidiospores infect the host
which bears the uredo and teleuto spores, but as a rule the regular course
of development is interrupted by the repeated production of uredospores
before the final stage is reached. The fungus is thus widely spread from
plant to plant by means of the uredospores and then the teleutospores are
formed in the autumn, either from the same mycelium or from a teleuto-
spore-bearing mycelium proceeding from the uredospores.

c 2

5»

Heteroecism.

This course of development may be represented graphically as follows,

taking P. graminis as the type:-

a, in Europe.

Spennatia

by in Australia.

Teleutospores

Sporidiola

Teleutospores

Uredospores
(repeated)

Uredospores

Aecidiospores

Fig. 14.

Uredospores
Fio. 15.

The aecidial stage of the spring rusts of wheat (P. triticina) or barley
(P. simplex) is not known, but Klebahn hazards the suggestion that the hosts
to which they respectively belong may only exist in their original home, and
thus not hitherto observed. With such widely and extensively cultivated
plants, the uredospores could easily be carried on the grain or by the wind,
and the fungus could thus be perpetuated without the intervention of an
intermediate host.

The tiding over of the winter is most important for those fungi which

depend upon two host

for their continued existence, but wh

this

accomplished by means of the teleutospores, other and addit

measures mav be taken to secure the same

Sometimes the uredo

mvcelium persists during the winter, and in the case of Puccinia arrhenathe

the aecidium-mycelium becomes perennial, and reprod

after

year

hile the teleutospore is also produced

tage is dropped, as in the case of Puccinia

When the aecidial
Australia, then there

is a profuse development of uredospores in comparison with teleutospores,
and there is abundance iot present needs, as well as for future germination

dial host is absent, the same

hen occurring

the

In oth

where the

thing has been observed, as in Caleosf

is absent

f fir trees, or Chrysomyxa rhododendri when the silver fi

Heteroecism <is said to increase the vigour of the fungus, and a striki

that the

given by Puccinia graminis

heat

sometimes said

dios pores from barbe

are much more virulent th

■th

dospores derived from the wheat itself, since the teleutospores produced
earlier, more copious, and more injurious to the wheat. With reference

Plowright

There is a wonderful difference in the amount

f injury done by mildew, when derived d
rhen derived from uredo that

from the barberry

that has reproduced itself through several gener-
The fungus grows with such enercv that it so imiures the

ations. . . .

wheat plant as to prevent it producing mon

As against this view, it mav* be well to be

few i
in mind th

the world probably does P. graminis cause as gre

does

barbe

hich the aecidial stage has never been found

ved kernels."
in no country
y to wheat as
l-existent. amd

further obser

to establish the fact, but Klebah

vegetative periods and other peculiarities of th 7

sible by the change of hosts, gives the fungus a decided advantag

on a number of species are necessary

that the utilization of the

rendered pos-

■d

Heteroecism.

Whatever view we take as to the cause of heteroecism, it is a well-

established fact that the promycelial spores

produce no effect on the grasses

matter how freely applied

theory as t
were at fii
other hosts

hich bear them, and the most probabl

associated

assumes that the uredo and teleuto spore

th the

dia. but

imately passed over to

Predisposition.

CHAPTER XVIII

Predisposition.

Wherever epidemic diseases caused by parasitic fungi occur, the ques

as to the relative susceptibility of different varieties or indi

tion is raised as to the relativ
viduals. The fungus has the
attacked, but the latter in its

pow

of causing di

the host

may either be favorably

abl

disposed

ards its de\

This predisposition of the host

for the attacks of the parasite is very variable, and is influenced by

ous factors. It is generally

dered that a sickly plant is more liabl

than a sound one : but, on the contrarv, strong and sound individual

more easilv and more virulently attacked

for artificial infect

selected. Ward7 has shown that when the ho

plant is starved by withholding' certain m

salts, and thus stunted in

growth and generally enfeebled, it is not affected in its

A. starved plant certainly develops smaller pu

other

f food

spores on account of the diminished supplies <
mvcelium, but the power of infection is just as great

sceptibility or
les and fewer
lable for the

il plants.

norm

j far as brom<
holds good fo

concerned

to doubt that

predisposition and immunity on the part

of the host, and impotence and virulence

the

of the parasite are

alike

dep

of mere nutrition. But, as we shall see afterwards

certain substances

of the nature of food

introduced into th

plant may affect its liability to d

There are various factors, ho

which may either dispose the plant towards disease or tend to render

it immune,

The age of

of these may be given here

attacked has

important infl

feet

particularly where the sporidiola are concerned. Young leaves and shoots

fected

duced

epidermis d

d when they get older little or no effect is pro
This is owing to the germ-tube of the sporidiolum penetrating the

and it is well known that this laver becomes

tough

d less easilv penetrable as it

older

The

rmer
edosp

and aecidiospores, on the other hand, infect the older leaves as well as the
younger, and this is easily explained from the fact that their germ-tubes
enter through the stomata.

Different parts of the same plant are also variouslv affected. Some-

times it is the

both sides of the leaf

are apt

sometimes leaf and stem, a
The different species of

choose diffe

may be on one or
on the same host-

portions. Thus Puccinia triticina, from

attack, is found most commonly on the

the sheath, mostly near its junction with the leaf while P

is worse on the upper leaves,
stem.

Different varieties or sorts
their susceptibilitv, and

and extends

graminis

d often parficularlv bad on sheath and

f the same species vary

be show

derably

it is

the

and

breeding of such rust-resistant sorts that solution of the 'rust-in-wheat ques
tion in Australia is being attempted. The samp i« th* MCfl «itk «**•

being attempted

rusts, and I have seen one kind of flax {Linum
attacked by Mclampsora " '

free

litii,

d anoth

same is the case with other

) badly

Hennings5 has recentlv made observ

kind

gsid

quite

hich tend to show that

plants previously susceptible to the attacks of a parasitic fungus may grad
ally become immune when they are changed to rich ground where they
better nourished and more vigorous.

Predisposition .

In 1894 several rhizomes of Peltandra virginica were sent to him from
North America, and planted in pots. An aecidium developed on the sterns
and midribs of the leaves, which was found to be new and named Aecid-
ium import atum. One plant was left in the pot, which was placed in
water, and the others were planted out in a soil composed of damp humus.
The pot plant has annually produced the aecidia up till the time of writing
(1902), while the other plants only showed the fungus very slightlv in
1895 and 1896, after which they grew exceedingly strong, and since then
have remained perfectly sound. It must be remembered that this is a
hardy marsh-loving plant, and there is probably more than mere nutrition
concerned in its freedom from disease, since it would be more reasonable to
regard the result mentioned as due to the change of situation than to change
of soil. Salmon3 considers that the evidence which is gradually accumulat-
ing on the subject of the relations between host-plants and parasitic fungi
leads us to the conclusion that immunity and susceptibility are due to con-
stitutional (physiological) peculiarities, and not to any structural ones. He
has also shown experimentally that while the uninjured leaf may be im-
mune, the same leaf when cut or injured may become liable to infection,
and the conidia produced on such leaves are then able to infect uninjured
leaves.' In this way the range of infection of a biologic form may be in-
creased. Different species of the same genus, when they are generally at-
tacked by a rust-fungus, may vary considerably in their susceptibility to
infection.

If we attempt to explain the varying susceptibility of different plants
or different kinds of plants, then the difficulty is apparent, and the sym-
biotic relation between the parasite on the one hand, and the host-plant on
the other, complicates the matter. Why is the fungus able to infect certain
host-plants, and not others? Why is the host-plant capable of resisting
certain fungi, and not others? How is the fungus able to accommodate
itself to certain plants, and not to others ? These and other questions may
be asked, but cannot be fully answered at present. De Bary2 says : — "The
physiological reason for these predispositions cannot, in most cases, be ex-
actly stated ; but it may be said in general terms to lie in the material
composition of the host, and therefore to be indirectly dependent on the
nature of its food."

The question has been asked, if there is any relation between liability
to infection or power of resistance and the visible structural features of the
leaf, and it has been answered differently by various investigators.

Hennings2 lays stress upon the physical characteristics of the parts of
the plant on which the parasitic fungi occur. He considers that the para-
site develops differently on a thin-skinned, delicate leaf, and a thick-
skinned, firm, leathery leaf. Also that the venation and hairiness of the
]eaf may affect the result.

Marshall Ward1 fully investigated the structural peculiarities of the
leaves of the various species of Bromus used in his infection experiments
with brown rust — such as thickness of cell-wall and cuticle, " bloom,
size, number, and distribution of hairs, distribution of chlorophyll-tissue
and vascular bundles — and he arrived at the conclusion that " the resistance
to infection of the immune or partially immune species and varieties is
not to be referred to observable anatomical or structural peculiarities, but
to internal, that is, intra-protoplasmic properties beyond the reach of the
microscope. ' '

Salmon3 comes to the same conclusion from his infection experiments
with the oidium of Erysiphe, but the physical characters of the wheat-plant
seem to have some some effect on its liability to rust, for there are certain

) >

Predisposition.

typical characters associated \
would appear therefore to fm

ith the quality of rust-resistance,
i a share in bringing about this

d they
It. In

a rust-resistant

heat

the leaves have a tough cuticle,

glazed often with a gl

not broad, erect not droopin

stiff

bloom
id f

d the straw

The flag is narrow
not soft and flabby.

all these characters may be present, and yet the climatic conditions

mav overrule them and weaken the power of

It will thu

be seen

of causes may contribute

renderin

im-

mune, and that not only the chemical

d other properties of protoplasm

come

play, but the

the mutual reaction of the

proto

plasm of host and parasite to be considered.

As far as the rust fungus is concerned, there is no such thing as a per
fectly immune wheat-plant, for, given the necessary conditions of situation,
heat moisture, and spores at the right season, and ati some time or anoth"

ppear

We express this by sa>
' proof "

rust-resistant

heats, but not
Animals are sometimes rendered immune, or, at least less

ptible

to disease, by repeated

feet

so that they become

dually accus-

tomed to the effects of the parasite, and the important question arises, C

be submitted to " protect i\

inoculation " ?

Inoculation with

ttenuated bacteria is the simplest method with animals, and this treatment
retards their vegetative development, and

This
different cases

ncially-induced immunity

__ . their

lasts a certain time, vary

effec

Ehrlich and Huebener

ha\

shown, from numerous

periments upon animals, that an immunized mother can impart immunity
her offspring, but it is not lasting.
In the case of plants, as in animals, there seem to be two opposing

forces at work

the one hand, the protoplasm of the fung

endeavouring to overcome the resistance to its entrance offered by the host-
plant, while on the other hand the host is more or less successfully resisting
the inroads of the fungus. There are no known cases of a plant becoming

"immune"

partially immune" by inoculation with the parasitic fun

to protect it against further attack, but advantage has been taken of the
gireat sensitiveness of certain spores to copper salts to protect the organism

against them.—
Chemotaxis
which certain organisms possess towards certain chemical substa

ore 6, p. 75.)
is the name given by Pfeffer 1 to a form of sensitiveness

pow

which certain nutritive and oth

bacteria and other organisms
chemotaxis.

sub
them

hav

and

organic

substances

with

ki
a

high

. . . This
attracting

as -p 0 sitiv e
nutritive

value which are positively chemotactic, may be mentioned asparagin

peptone, while sugar, which is

of the best food stuffs and richest

of energy, has but little attractive pow

The power of

substances

the contrary

pel bacter

taxis, and free acids and alkal

as well as alcohol, have this' effect

Gly

far as kno

subst

Massee9 has stated

immunity is owing to the absence or small proportion of the subst

the parasite in the plant not attacked

d if

can

be impregnated with some substance which is negatively chemotactic, and

at the same time does not affect the

lty of the

gainst parasitic fungi may be obtained in this way

then immunity

Laurent1, acting on this principl

ducted a series of experiments

Phytofhth

the possibility of producing potatoes which would be proof against

He grew very

ptible varieties in pots, to

the soil of which sulphate of copper was added, and when the tubers
harvested, some were cut in two and their cut surface placed in contact with

know as negative chemo- •

Predisposition.

the mildew of potato leaves. After four days, the tubers grown in the
soil containing copper did not show any infection, while those grown in
untreated soil were infected in a very decided manner from being brought
into contact with a mildewed leaf.

Marchal 2, following on the same lines, tried to secure immunity to
lettuce against Bremia lactucae, and he found that the plants treated showed
considerable resistance, and the immunity seemed to be in proportion to
the strength of the solution. He also experimented with cereals for p

tection against rusts, but

and the probability is that he did

not use the proper substance which repelled the germ-tubes of the irust-
spores when they attempted to enter the plant.

Massee7 experimented with cucumber and tomato plants, watering the
soil on which they were grown with a solution of sulphate of copper. The
result was that " not a single one of the treated tomato plants showed a trace
of disease " after being sprayed with water containing the spores of the
fungus, while the untreated check plants were badly diseased.

It is important to note that tomatoes produced from plants treated with
solutions of copper sulphate were found on analysis to show amounts of
coppar not sensibly greater than that found in the fruits obtained from un-
treated plants.

These experiments at least show that certain substances entering into the
constitution of the host-plant render it for the time being immune to the
attacks of certain fungi, even although it was naturally predisposed to that
particular form of disease.

•

Wheat Rust in Australia.

CHAPTER XIX.

The Present Position of the Rust in Wheat Question in Au

It will tend to clearness if we confine our attention to the rust i
for the general considerations which hold in this case will apply to the rusts
on the other cereals chiefly

ated. viz. , oats

d barlev

And the

hich is most important from the farmer's point of view, because

does the most d

be chosen

Puccinia grami

distinct in its characters that there is no difficulty in recognisin

hich is so

There

are

kinds

heat in Australia, the positively injurious

Puccinia era

d the comparatively harmless P. triticina, beca

but

the early d*ays whe

it is

grown to

9°3-4

does not pinch and shrivel the grain like the other.

and other cereals had to be imported into Sydney,

such an extent in the Australian Commonwealth that in the

there were 5,566,340 acres under wheat, yielding a total of 74,149,634

bushels. Wheat will always form one of our staple products, and from the

great extent of the industrv, whatever increases the yield or tends to diminish

the losses from d

have a corresponding far-reaching effect

has been given to the subject, it has been found that

vear passes without its being present

heat to a greater

d in some years, which are commonly spoken of as rusty years, it seriously

the grain and considerablv diminishes the yield. To give some con-

idea of the extent of the

bad season like that of

be stated that in a part

the loss was estimated to be for the whole of

bet

d£s

such favorable y

for the rust, there is always an evident connexion between the weather and

epidemic

In 1889 it was a wet spring, and about the blooming

weather was what was known as " muggy,"

lg of show

th heat between, and heavy dews, so that the wheat-plant, at the time of

oming into fl

rticularly susceptible, and the rust spores found

a ready entrance into the tissues of the plant, with their accumulated
of food, and thus it spread rapidly.

: conditions are often regarded as the prime factors

Atmospheric

production of rust, but they are only concerned in the matter i
they favour or hinder the development of the parasite which ca
this is strikingly shown in the existence of rust-resisting wheats

Owing

the importance of the subject, and the widespread losses, delegates from
various States were appointed to meet and confer periodically

various

ferences extended from the first in 1890 to the last

The results of their labours may be seen in their voluminous reports, which
practically cover the different phases of the question.

But there

some

x>rtant points with which they were not then in a position to deal, such
the effects of rust on the straw and grain, and experiments in the inocu-
ion of the barberry. These will be considered preliminary to the main

question

Effects of Rust on the Straw and Grai

XLIV

The visible effects of the rust on the wheat plant is well shown in Pll

where the straw of the rust-resistant Rerraf is bright, glossy, deep

and well ripened, while that of the badly rusted Queen's Jubilee has

dirty, streaky, patchv appearance. The ears of the one are full-sized and

well formed, with plump
with light and shrivelled 1

filled

grain, and the other has poor ears

Wheat Rust in Australia.

But the effect upon the com

of the

d particularly its

feedin

is not known

the farmer, who cuts his cron for h

: threatens to ruin it.

This has recently been determined by F. T. Sh

ltt l, chemist, Dominion
arms, Canada, who analyzed two samples of wheat grown
Manitoba in the same field and of the same age, only the one was rusted

Experim

and

othe

analysis is as follow

ithat the results are strictly comparable

The

Analysts of Rusted and Rust-free Wheat-straw and Grain,

u- X

O CJ

•

iff c5

.£f*

•**

fi-p

Straw from rust-free wheat
Straw from rusted wheat
Grain from rust-free wheat
Grain from rusted wheat

Grams.

• • »

7-92

• 9 •

7-92

3*o5°4

12-26

1 -4944

io*66

2-44

7-69

10*50

13-69

165

1 -97
2-56

2*35

39-00

38-44

70-55
68-03

39*95

3678

2*29

3*o3

9-04
7-20

1-84
2-24

The Sir a

pointed out that in crude protein the rusted

much richer, and since this includes all the nitrogenous compounds of a food

that go

form blood and build up muscle, it may safely be

concluded that the rusted

much superior in feeding value,
more fat and somewhat less fib

There

is also in the rusted straw slightly

that all this affords additional evidence of its more highly

The handling and feeding of rusty straw in Australia from the farmer

point of view has received

The experience of one who has done

a deal of threshing is to the effect that when very bad it caused an itch

sensation,
broken.

d made the men about the thresher rub their sk

egards feeding rusty hay, another with larg

experience

nforms me that horses and cattle relish it far before ord

course, it

fed as chaff.

— This from the rusted wheat is onlv about

The Grain —

srht of that from the rust- free wheat, but

half the

it has

protein content show

ght for weight, a considerably higher nutritive value.

for the higher protein content in the smaller grain in its larger pro

portion of b
accumulatior

but chiefly in the partial and incomplete transference and

of starch

These results likewise afford interesting evidence as to the physiological
effect of the rust on the wheat plant, and agree with what has been deduced
from other data.

In the actively growing and feeding period of the plant's life, it is
apparently able to provide for the wants of the fungus as well as its own,
and therefore its vitality is not seriously affected. But when the second
period of forming and ripening the seed arrives, when feeding is gradually
ceasing, and the accumulated materials are being transferred to the seed,
then the fungus draws upon the plant's capital, crippling its energies, and
checking the movement of the food materials to the seed. As Shutt sum-
marizes the whole process: — " The growth of the rust arrests development,
and indicates premature ripeness, which, as we have seen, means a straw
in which still remains the elaborated food, and a grain small, immature,
rich in protein and deficient in starch."

This emphasizes what we have frequently insisted on, that the critical
period, literally the turning point in the plant's life, is reached when it

Wheat Rust in Australia.

beeins to form

d if that

favorable for the develop

grain

ment of rust, then the fungus has to draw upon the stored-up material, and

is not fulTy formed, if at all. Incidentally this
throws light upon the reason why the earlier rust (P.
) is comparatively harmless, since its period of greatest activity^ is
when root and leaf are busy manufacturing material sufficient to provide
for the necessities of both.

consequently the
investigation als(

On account of the comparatively

appearance of P. graminis, it js

by some considered to do the least damage to the grain, but,^ as a^ matter
of observation, and on physiological g

rounds

known

Aus

be the

Puccinia Graminis and the Barberry in Australia.

In Europe and America the fdentity of P. graminis is determined, not
merely from its morphological characters, but from its ability to infect the
barberry and produce aecid

But in Australia infection of the barbe

has not been successful

although

several attempts have been made, and

have doubted whether we have got the

P. graminis

d not a

distinct biological form of

From a comparison with European speci-

mens

examination of the

the uredo and teleuto spores

there is no doubt that the rusts are very much alike, only if the infection of
the barberry is accepted as a diagnostic character, then the identity is not

proved. The relation of

therefore, to the barberry in Austral

became a pressing subject for experiment.

The germination of the teleutospores may be easily accomplished at the
proper season, either by placing them in a drop of water on a slide under
a bell-jar, or, better still, if copious germination is required, iby taking some
of the rusty straw and placing it on a drop or two of water in a petri dish,
the cover of which is lined with damp blotting-paper. It is only after a
considerable rest that they will germinate, and they start on warm days
about the end of September, which is the beginning of our spring. This
may continue through the warmer days of October, almost or entirely ceas-
ing in a cold spell, and even until November germination continues if the
conditions are favorable ; but although numerous trials have been made, no
germination has occurred outside these months. Generally speaking, the
middle of October is the height of the season for germination, and then,

the uredospores may be multiplying rapidly on the growing wheat-
plant; so that there is no "off " season in Australia, as far as wheat is con-
cerned, when the teleutospores are active and the uredospores dormant,
In any case, as our wheat crops are usually harvested in November and
December, even if the barberry were common, and developed rust freely,
it could hardly be of much importance as a factor in spreading the wheat
rust, since the aecidia would not be developed profusely before the wheat
crop had passed the danger point.

The barberry is not a native of Australia, and very few hedges exist,
so that the question of its- infection is not of great immediate practical im-
portance, but it is of high scientific value to establish the fact that the rust
may pass one portion of its life on one plant, say, wheat, and continue it
on a very different plant as an intermediate host, say, barberry, and thus
settle that the rust with which we have to deal is the P.

too,

graminis

Europe.

As early as October, 1892, I succeeded in germinating the spores freely
and ^ copiously in a watch-glass with water, and infected four different
species of barberry obtained from the Botanic Gar'dens, two of which were

Wheat Rust in Australia

•

known elsewhere to carry the aecidial stage of P. graminis. In one case
the infected branch was placed under a bell-jar to preserve moist conditions,
but there was no result with any of the species, since the weather was too
hot and dry.

In 1902, in order to give the experiment another trial under the most
favorable conditions, Dr. Plowright kindly forwarded several young bar-
berry bushes from England, which arrived here in December in good con-
dition. Rusted straw was specially kept exposed to the weather in order to
inoculate the barberries. About 16th September the barberries were
putting forth their young leaves, and looked very healthy. One was kept
as a check, and the others were infected, either by scattering rusted straw
around the plants and tying it on to them, or, in two cases, by applying

germinating spores direct to the leaves. Some plants were kept under
bell-jars, others exposed, and all were attended to and watered freely. The
result was that not the slightest trace of any fungus appeared on any of the
barberry leaves. It may be stated that the conditions for fungus growth
were most favorable, as at times that muggy heat prevailed, which so
quickly spreads the rust in a growing crop.

In 1904 the experiments were continued in the pots, partly on the same
lines with rusted straw, and partly by planting a rusty wheat beside the
barberry, so that it might be naturally infecfed next season.

Cuttings from the English barberries have also been planted at Port
Fairy in a rusty spot where they have thriven, and Queen's Jubilee
wheat planted around them produced abundance of P. graminis. The
rusted straw was allowed to die down on the spot, and every
facility given for the inoculation of the barberry, but without
result so far. Some P. graminis on wheat was sent by Dr.
Plowright in March, 1903, which was gathered In September,
1902, and kept in his garden till March. On arrival here
some of it was kept inside, and a portion placed outside exposed to the
weather, just as was done with Australian rusted straw. It was thus
exposed during our winter months of June, July, and August, and in
September it was tested, being then exactly twelve months old. While the
teleutospores about six months old from the Australian wheat germinated
freely, there was no change in the spores from the English wheat, and
although attempts were made at different times, there was never any
sign of germination. Probably they were kept too long, as they might
have germinated on or about the English spring. Prior to this, I had sent
rusty Australian straw for trial to Dr. Plowright, but none of the teleuto-
spores showed the slightest trace of germination. Writing in the Gardeners1
Chronicle for 15th January, 1898, he says6: — "I have tried on two
occasions to get the teleutospore of Puccinia graminis from Australia
to germinate in England, but I have not succeeded. The first attempt was
made nine years ago, when Mr. D. McAlpine was good enough to send
me material ; but I was quite unsuccessful. Last year he was kind enough
to send me a further supply ; but, although I kept the straw out of doors
during the latter part of last winter and the spring of 1897, I was equally
unsuccessful. Is it probably like the seeds of some of the higher vege-
tables, the teleutospores require not only a period of rest, but also an

I exposure to a certain degree of cold?" Seeing that the spores germinated

freely here, the "exposure to cold " theory does not hold ; and the most
probable explanation is that they had been kept too long, and attempts
to germinate them should have been made in the English autumn. As
far as Australia is concerned, the rust which does the principal damage

1 has apparently no intermediate, stage.

Wheat Rust in Australia.

The Aust

rust has been determined as Puccinia graminis by

Eriksson and other authorities; but if the ab

produce the aecidium

on the barberr

is taken as

the final criterion, then

must be

pended in numerous other cases.

Thus Massee4 has found P

Kew ;

Herberts

Alo-peeurus pratensis, and Avetta elatior in the Royal Gardens

.' ./. .1 jj_ n vr^^f-kotnr^inrr thf* crrpnfr nua.ntitv o

but he significantly adds

Notwithstanding the great quantity of

and Mahonia present in the grounds

most careful and con-

h has failed

the presence of the aecidiospore stag

(Note 7, p. 75

The Problem St

In dealing with the rust question from a practical point of

were two main

issues to be determined

How is the rust spread and continued f

may its injurious effects be

^ated

or counteracted or

prevented

The first question is a most

portant one, for if we could find out

where, and under

time from reapim

ditions, the rust is

lvino-

dormant

du—b

the

rop

ing

again

destroy it at this stage, and prevent its reappearance

might be able to
Although the

question may thus be simply stated
The second subject of pre\

by no means easy to answer,
or mitigation will evidently depend

wheat
render

know

ledge of the life-history of

fun

well

d how far the conditions can be controlled which

mt itself, ar

susceptible to the fungus.

the Rust is Spread and Continued /

actly now, thanks to the labou

to Y

of Eriksson

Marshall

Ward, and

how the,

germ-tiibes, how the)

enter the plant by means of their
grow and ramify among the tissues, and drain them

of thei

until thev

again

form a spore

and reproduce the

pores on the surfac

great

abund

We thus know how fresh

s pores

orginate

once thev have

got

a start: but it is the starting-point

ffic

Although the rust was know

d the effects produced by it were

familiar from remote antiquity, yet its true nature was not discovered
until the latter half of the eighteenth century. As late as 1733, Jethro

Tull

abo

attacks of small insects

i t

Horse-hoeing Husbandry, attributes

the

brought,

feed

some think, by the East wind, which

the wheat, leaving their excreta as black spots upon the

shown by the microscope

767, its true nature as a fungus, and

therefore as a plant, was determined by Felice Fontana, and in

797

Persoon gave it the name bv which it is still known, Puc

The rust then is a fungus growing

de the wheat-pl

and

graminis

g at

its expense

prod

or spores, which 1
at certain seasons.

itself by means of minute seed-like bod

conspicuous on the leaves and stem of the wheat

For a long time there was a suspicion in the minds of many p

farmers that the barberrv bush had

something to do with its spread

so firmly was this believed in, that the State of Massachusetts passed an Act

compelling the
Barv, in 1864,

abit

to extirpate barberry bushes. And when De

fied the farmer

d scientifically th

was a connexion bet

bush

d th

the fungus which appears on the barberrv

hich appears on the wheat, then it was

thought by

many that we had reached the root of the matter, and that we had simpl

dest

the barberry bush in order to get rid of the

it is

well

Wheat Rust in Australia.

known that here in Australia, where barberry bushes are not native, and
where they are comparatively scarce, the rust is particularly bad in certain
seasons, so that there must be other causes to account for the prevalence
of rust.

The connexion between wheat-rust and barberry has already been dis-
cussed, so need not be further referred to here. There are several possible
ways in which the presence of the rust year after year may be accounted for,
and it may be worth while to consider some of these.

1. The uredospores produced in such immense numbers may serve to
carry it on. They are very minute and light, easily distributed by the
wind, and it has been shown, that they exist in the air and on the ground.
They might thus be readily transported and even carried to localities
far removed from wheat-growing areas, in some cases by the duststorms
which are very prevalent in the northern parts of Victoria. But the mere
presence of spores is not sufficient to account for the rust being spread,
since they must be capable of germination. I have often tried to germinate
uredospores taken from straw that had been left on the ground, but without
success. My latest attempt was with viredospores still retaining their
colour from a sorus on a dead leaf of Queen's Jubilee wheat on March 25th.
The spores^ were kept moist under a bell-jar, but not a single one ger-
minated. But the result is different when spores are taken from self-sown
wheat growing in the interval between the two crops. — (Note 8, p. 75.)

2. These spores are not only in the air and on the ground, but they
are commonly to be found entangled in the bearded tip or u brush " of the
grain. In one variety, Queen's Jubilee, this was so common that not a
single grain could be found without the uredospores. Dr. Cobb11 has
likewise examined the brush of a number of varieties in New South Wales,
and found in about 57 per cent, of the grains examined 'that the spores
were in the brush. This is an evident starting-point for the rust, but not
the only one, since seed wheat treated with bluestone, formalin, corrosive
sublimate, and other fungicides, produced rusty plants, and in fact there
was little difference as regards rust between the plants from treated and

untreated seed.

3. It was commonly supposed until recently that the rust could readily
pass from one cereal crop to another, and thus it was passed on to the
wheat at the proper season. This view was put forward as late as
September, 1904, in the Journal of Agriculture of South Australia, by
A. Molineux1, who states: — "I have observed for many years that when
ever we have a mild autumn and summer, accompanied with occasional
showers, we have complaints of red rust in the succeeding crop ; and I have
been led to the belief that until the new wheat crops have started, the rust
is nursed by the wild oats and other cereals that may always be found
growing on the headlands and by the roadsides." Of course, this is a
very convenient way of accounting for the presence of rust throughout the
year, but Eriksson has shown that the spores from oats will neither infect
wheat nor barley, nor will the spores from the barley infect wheat or oats.
It follows from this that adjacent fields of these crops will not affect or
be affected iby each other, so far as this rust is concerned. Probably,
however, Molineux's view is correct, except that it is the self-sown or
volunteer wheat growing in our paddocks or on the headlands that carry
it over. The system of harvesting practised in Australia with the com-
bined harvester, which takes off the heads only and delivers the winnowed
grain into bags, necessarily implies the scattering of a certain amount of
seed on the ground, and this germinates with the first rain, and is almost
always partly rusted, and often badly so. Our hay being largely made

Wheat Rust in Australi

from wheat, when rains fall shortly after it has been cut, the second
growth or aftermath is almost always rusted, and so a second crop of

uredospores is produced.

4 Since " intermediate hosts," such as the barberry, are not concerned
in earning over the rust from season to season, it has been suggested that
infection may be communicated to the wheat from overgrasses which^as

>vc have elsewhere shown, may also be attacked by Pucctma gratnims
Klebahn1 (p 230) has shown that uredospores from various grasses will
infect wheat, and it remains to be determined how far the grasses occurring
in our wheat-fields, and attacked by this rust, are capable of infecting it.

5. Towards the end of the growing season, a second kind of spore is
produced, known as the teleutospore. It will not germinate immediately,
but onlv after a period of rest, and it may also aid in carrying over the
rust from season to season. But although it can germinate in the spring,
and produce other minute spores known as spondiola, still they have not
been proved to infect the wheat-plant, and so we do not know what pur-
pose thev serve, if any. In other countries they a.re said to germinate upon
barberry leaves, and produce the aecidial form of the rust— and it may be
that here thev are simply dying out— are becoming functionless, because
the barberrv bush which they normally infect is not now available for them.

6. There is still another way in which some rusts are propagated, and
that is by means of the threads of the fungus or mycelium remaining in-
side the seed and starting into life with the germination of the grain. But

0h hundreds of seeds have been carefully examined by the micro-
scope, °no trace of this has been found, and therefore for the present we
must decline to regard it as a probable cause.

7. There still remains another possible means of continuation from sea-
son to season, which has been prominently brought forward by one who has
devoted considerable attention to the study of the rusts in Sweden, Profes-
sor Eriksson. Although he has not yet succeeded in giving scientific proof
of his theory, he considers that while infection by spores does occur, yet
the primary infection is from within, from an internal germ of disease
inherited from the parent plant and latent in the seed. He grew wheat in
closed chambers, where it was believed to be secure against infection from

without, and still the rust appeared all the same, and he can only account
for this by supposing that in the cells of the seed the protoplasm is asso-
ciated with the plasma of the fungus — what he calls mycoplastn — and from
this there arises, if the conditions are favourable, the mycelium of the rust
fungus, quite independent of external infection^ He does not seem
to have considered the possibility of the spores of the fungus
being attached to the seed, and until the soil and the seed are thoroughly
sterilized and every precaution taken to exclude infection from without,
and the disease still produced, until then we must suspend our judgment
and accept the Scotch verdict of Not Proven. A very striking
case, however, that the seed may be the means of continuing

rusts from season to season is given by Carleton 3 in connexion

with Euphorbia rust (Urotnyces euphorbiae, Cooke and Peck). The pods
of Euphorbia dentata, and even the naked seeds, were found to be affected
with aecidia, and on growing the rusted seeds under a bell-jar, those that
were disinfected produced plants without rust, while those not disinfected
gave rise to rusted plants. Here the seeds actually bore the aecidia, and
propagated the rust through the germinating seed. A similar instance is
met with in Aecidium platylobii McAlp., where the aecidial cups are borne
on the pods, and on opening the diseased ones, the seeds are frequently
found covered with the mycelium, which, on microscopic examination, is
found to penetrate them.

Wheat Rust in Australia.

Prevention or Mitigation of the Rust.

The all-important question now remains to be answered, how to pre-
vent or mitigate (the effects of the rust. In the various reports of the
Rust-in-Wheat Intercolonial Conferences, special attention was paid to the
solution of this question, and the effect on the crop of different cultural
methods ireceived a large share of attention.

Drainage. — It is commonly affirmed that rust is worst in hollows, where
water lodges, and on general grounds it is considered that drainage by
removing the surplus moisture would tend to afford the wheat the most

favorable conditions for its healthy development, and thereby render the
rust less injurious. But, as a matter of fact, when it was experimental Iv
tested in Victoria, an increased yield was the result ; but as regards rust, it
did not seem to affect it, for out of six areas artificially drained, five of them
were rusty. In a number of our wheat-growing areas, it is not excess of
moisture in the soil, but deficiency of it, which is complained of, and in
such areas the rust is very severe in seasons marked by copious late spring
rains.

Irrigation. — At the present time irrigation is the great problem which
overshadows all others in connexion with the utilization of the land in Aus-
tralia, and its relation to the development of rust has not been overlooked.
In irrigated areas where wheat was grown, the effect was observed, and it
was seen that the judicious application of water was beneficial if done at
the right time, and with due iregard to atmospheric conditions. There is
only one rational method of irrigation for wheat in Victoria, and pro-
bably for all Australia, and that is a thorough soaking of the ground
prior to ploughing. Mr. Geo. Pagan, of Ardmona, who has successfully
irrigated in the Goulburn Valley, states in the Journal of the Defartment
of Agriculture for March, 1905, that his routine practice is to flood the
land in March, unless there has been heavy rain, and then plough as soon
as the horses can work without sinking.

This one watering, followed by proper working of the land, will usually
mature the grain. Even in 1902, when the year's rainfall was only six
inches, this was the case. The application of water later in the growing
season is always risky, and often does more harm than good. It spoils
the grain, making it soft, may bring on rust, and results in very uneven
ripening.

Seed-bed. — A relatively dry and firm seed-bed is generally considered
best for wheat, and especially if there is a prospect of rain after sowing
to ensure a good germination. Its effect on rust has not been definitely
determined, but farmers are generally of opinion that sowing on a dry
seed-bed gives the plant a better chance to escape the rust. If the seed-
bed is moist, the condition will be favorable for the rust-spores on the
soil, or on the grain, to infect the germinating wheat-plant, but probably
it is not so much the nature of the seed-bed as the aftergrowth which will
affect the result. This also applies to the mode of sowing, for it did
not appear to make any difference, as regards rust, whether the wheat
was sown broad-cast or drilled.

Ploughing and Harrowing. — The question of deep as opposed to shallow
ploughing does not seem to have much to do with the rust. In experi-
oiental tests, deep ploughing yielded the heavier crop, and thus indirectly
favoured the development of rust ; but both were about equally affected.
Harrowing when the crop was about two feet high was equally incon-
clusive.

Rotation. — Rotation is undoubtedly good for the crop, and is a factor
in clean cultivation ; but it is another question, whether it is good or bad for

Wheat Rust in Austr

the rust. At Port F

in Victoria, where rotation is regularly prac

tised

here I have had crops

grown

the same season, sometimes

fter mangels, or potatoes, or
rustiness from

those crops

d-point of good farming, it

onions, there was no

ptible difference

heat grown in succession.

From the

be recommended ; but cropp

fter vear does not seem to affect the liab

to rust further th

that self
crops.

Such

heat is likely to appear in the interval bet

unteer

heat, if not killed in

atmg

ng year
in this,
the two
is much

d on account of its earliness may escape

it, for it is some-
ppears earlier, so that

more forward than sown grain, ai

the rust, but on the other hand, there is a risk attend

times more rusty than the ordinary wheat

it is one of the means of carrying over the rust' from one season to another

All such wheat should therefore be kept down by means of sheep.

Fallowing. — This practice is similar in its effects to rotation as regards

rust,
tralia,

Professor Lowrie, speaking from a large experience

South Au

ders that the apparent exempt

hich fallow-land sometimes

land fii
seasons

from rust is due to the fact that it is customary to sow bare fallow-
rst, and the crop ripens sufficiently early to escape the rust in some

Burning Stubble

It is sometimes recommended to b

the stubbl

order to destroy any spores of rust that may be about; but

of experience, this has not been found to produce any appreciable diffe

ence.

Theoretically, the burning of the surface of the

should destroy

umber of

and to that

beneficial ; but, practically, the

dffference in result is not marked. As far as most wheat-soils
cerned here, the loss of vegetable matter is much more serious th

are con-

any

possible increase of rust.

Soils. — Soils are sometimes said to be rust-liable

d rust-free, like th

wheat itself ; but on closer

is only

of several factors th

gat ion,
require

it will be found that the soi

be taken

consideration

Rich soils are said to suffer most, and in rusty years the best crops have

sometimes been raised from the, poorest soils.

the rich soils the crop

naturally heavier and more luxuriant than on poor

d the softer

and more

ptibl

easily

aded by the parasite

Hence

that the rust may sometimes be worst in the best crops. The mallee

Victoria

crop

said to be particul

free : but in some seasons th

to be cut for hay, in order to save it from being completely

destroyed by rust. The reason for g

freedom from rust lies not

much in the

the light rainfall and the early ripening of the

crop

But if " muggy " weather should prevail when the wheat is flow

then the crop is just as liable to rust here as elsewhe

At Port Fairy

the low-lying black soils more readily produce a rusty crop than the sandy

soils, and this may be
absorb more heat, and

due to the fact that the dark-coloured soils
more likely to preserve the spores

for germination, or because they grow a heavier crop, which often lodg

and thus encourages the rust.
Ma

Perhaps no means for the prevention of rust h

more thoroughly and continuously tested than the appl
manures. In all the States, as well as in N

Zealand

been

of different
periments

have been conducted to this end, and the general opinion has been expressed

by a late D

been discovered that

of Lincoln Colleg

Zealand

No manure has

The

is a preventative of rust in cereal crops
tests were carried out under varying conditions, and, as might be expected,
the results of one year were often contradicted by those of the next;
still, it was generally found that nitrogenous manures favoured the rust,

it. Nitrogenous manures

hile phosphatic ones had a tendency to diminish

Wheat Rust in Australia.

tend to increase the amount of flag, and retard ripening, by affording an
excess of nitrogenous food, whereas phosphate of lime tends to induce

early maturity, and1 thus enables the crop to escape the rust to a certain
extent.

Treatment of Seed. — This was thoroughly gone into, since ix is so often
stated that the disease is in the seed. No doubt appropriate treatment
destroys the spores entangled in the " brush," but inside the seed no
mycelium has been traced. A great variety of " steeps " have been used,
and I have myself experimented with over twenty, including the hot-water
treatment; but they were all of no practical benefit. Last season (1904)
two plots of Queen's Jubilee wheat were sown the same day and grown
alongside of each other, in one of which the seed was treated with
formalin, while the other was untreated. The rust was bad on both plots,
and although special attention was given to the matter in the field, I could
not say that treatment of the seed with formalin gave any advantage as
regards rust.

Both sulphate of copper and formalin destroy the rust spores on the
seed-grains, and Dr. Hollrung, as the result of a series of experi-
ments, has recommended the latter as the best for this purpose. But since
infection chiefly takes place when the whear plant is above ground, it is
evident that the formalin treatment does not prevent it, and the experience
of numerous farmers who have used formalin successfully for the treat-
ment of stinking smut (Tilletia tritici), bears this out.

The hot-water treatment of the seed is constantly being brought forward
as a remedy for rust, but in 1892 the seed for 118 plots, consisting of
different varieties of wheat, was treated with hot water at 55 deg. C, and
in some cases the rust was just as bad as if no treatment had been given.

It was adopted as a conclusion at the last Rust-in-Wheat Conference
that the treatment of the seed is valueless for rustt, and Dr. Cobb10
one of the representatives of New South Wales, went so far as to say :
" As for curing rust by treating the seed, the idea is ridiculous. It would
be just as reasonable to expect to prevent measles among mankind by
soaking babies in some sort of pickle. "

It has not been thought necessary to refer specially to spraying as a
means of combating the rust ; for although this method is practicable in an
orchard, and has ibeen found successful in treating peach and plum rust,
still the mechanical difficulties to be overcome in spraying a wheat-fielH
are so great, that it is no longer regarded as of practical importance.

So far it would seem as if the rust in wheat defied treatment, and the
only practical measures' to be recommended for mitigating its effects were
to sow early and to select early maturing varieties. In this way it is often
possible to escape the rust, or the crop is too far advanced to suffer
seriously. But in this, one is at the mercy of the weather, and the only
hopeful remedy i's to grow wheats which will be able successfully to
resist the rust, even when the weather favours its development.

The question has been raised as to whether a wheat which resists one
kind of rust caini succumb to another in a different country, and this has
reallv been found to be the case. Professor Eriksson sent me ten varieties
of Swedish wheats which had been grown in the experimental plots, and
found to resist the rust which is prevalent in that part of the world, viz.,
Golden Rust (Puccinia glumarum). When grown here these rust-resisting
Swedish' wheats became rotten with rust, although of a different kind ; and
this, along with other experiments, points to the possibility that a wheat may
resist a rust such as P. graminis in one country and succumb to it in another.

Selection and Cross-breeding. — As the result of numerous experi-
ments, and the trial of hundreds of varieties of wheat from all parts

Wheat Rust in Australia.

of
been

the

world (Europe

that

say

proved
there is no wheat k

Asia
heat

Africa,
ahsol

America)
ust-proof

it
that

has

hich will be proof

against

when grown under conditions favorable to its^ development

experiments have show
particular d

among

the rust
But the same

umber of varieties grown in a
more than others, are able bv

constitution— it may be by their vigour of growth or

theii

their cuticle or glaucousness

combined

sufficiently to prevent its seriously injuring the
known as rust-resisting wheats. At present we ha\

less of
the inroads of the

d such wheats

least one such

wheat which has been grown in various districts of different States along
side of other wheats badly rusted, and it has been able to withstand th-

rust,
time

Rerraf is the wheat referred to, and perhaps it, too, in course of

1 succumb to the rust,
this great and burning

question the only measures I can

sug

i. To produ

heats suited

Australian conditions by crossing

Wheat Experimentalist of N

buth Wal

is now so

as Mr. Farrer,
successfully doing.

To select amd carefully cultivate the most rust-resisting plants from
these and other wheats having the necessary hardy, prolific, grain-holding,

d milling qu

keeping up the

g thei

igour

3. To cultivate early maturing varieties and sow them early, combined
with the best agricultural methods, such as clean cultivation, judicious
rotation, suitable (phosphatic) manuring and fallowing, to insure a good

crop.

In judicious crossing and careful selecting lies the solution of the great
rust problem, as far as our present knowledge goes, and to increase and
extend this knowledge it would be desirable to follow the advice of
Professor Eriksson, who has done more than any other single individual
to enlighten us on the subject : — " The question of the rust of cereals being
of the greatest practical importance for every country, means should be
furnished to those in whose hands the direction of these investigations are

placed, to meet from time to time, that is to say, at least every five years,
in order to discuss, according as experience is gained, the value of any new
observations, and to gain for their work the advantage of being based on
a plan common in its essentials to all."

Rust in wheat has been known in Australia for at least 8o years.
H. C. L. Anderson, Principal Librarian of the Public Library of New
South Wales, informs me that the earliest record of it is given by
Atkinson,1 who, referring to 1825, or even earlier, savs that " rust sometimes

appears, but it is not very common."

Montague Smith, in 1828, notes

that the plains and forest lands of the Hunter district of New South Wales
suffered from rust in wheat. As the wheat-growing area extended, the
rust also spread with amazing rapidity, until now it is undoubtedly the most
widely distributed and the most destructive of all the diseases to which
cereals are subject.

P. 4

Eriksson18

Note i.
his latest work on the vegetative life of Puc

graminis, has described and figured a transition from the mycoplasm to the
protomyoelial stage, but it still remains to be seen how far his interpretation
of the facts is borne out by other observers.

Wheat Rust in Australia

Note 2.

P. 14. — Christman1, however, has clearly shown that two fertile cells or
swollen hyphal branches come into contact, and at the point of contact an
opening is formed by solution of the cell wall, and thus their contents mix
although the nuclei do not fuse. There is thus true sexual cell fusion with-
out the intervention of spermatia.

Note 3.

P. 24. — Arthur8 considers the sorus in such species as Puccinia bromina
and P. triticina to be compound and the modified hyph'ae which separate
the individual sori to form a stroma, hence they are not paraphyses, strictly
speaking.

Note 4.

P. 37. — The origin of the aecidiospores as shown by Christman,1 from the
fusion of sexual cells and the peridium from morphologically equivalent cells
disposes of the view that the aecidiospores may have been derived from
teleutospores . If, as Blackman suggests, the rusts originated from the red
algae, then the sexual product or aecidium would represent an early stage
in the history of the Rusts.

Note 5.

P. 43. — Hooker, in his classical essay On the Flora of Australia (1859),
has a chapter on some of the naturalized plants, showing that even nearly

half a century ago, the chick-weed, knot-weed, scarlet pimpernel, daisy,
mallow, sweetbrier, and various other common plants had been introduced.
The groundsel, however, is not mentioned, and no doubt as trade increased
with other countries and exchange of products took place, the importation
of the seeds of weeds became common.

Note 6.

P. 62. — Beauverie1 has experimented with Botrytis cinerea, or grey rot,
and obtained, in sterilised soil, an attenuated form of fungus. He
infected soil liberally with this form, and grew plants therein from seeds
and cuttings perfectly free from the fungus, while plants not rendered
immune in this way perished.

Note 7.

P. 68. — Kirk informs me by letter what he has already stated in his
Annual Reports that " Barberries are being largely used! in New Zealand
for hedges, and, up to the present, I have never seen any sign of aecidia
of Puccinia graminis on them, although I have examined hundreds. It
would appear as if Puccinia graminis in Australia has lost the power of
forming aecidia on barberries.' '

Note 8.
P. 69. — Bolley,5 however, has announced that he found the uredosp

of Puccinia graminis successfully surviving upon dead leaves and straw,
even retaining their vitality when exposed to the drying winds of autumn,
and the intense cold of winter. «

■

«■

PART SECOND

CLASSIFICATION

AND

TECHNICAL

DESCRIPTIONS.

■

-4

•

Classification

CHAPTER XX.

Classification, with Special Reference to Biologic Forms.

Our views as to the limits of species have undergone a change owing to
the results of infection experiments in the rusts as well as in other groups
of fungi. Species have hitherto been distinguished on morphological
grounds, those possessing the same structural characters being considered
identical, and separated from those which differ from them in essential
points. But in recent times, when infection experiments have been carried
out on an extensive scale, it has been found that parasitic fungi, completely
agreeing in structural characters, or at least differing so slightly as to be
incapable of separation, have very different infective powers. It has
therefore become necessary to recognise such forms, and since the dif-
ferences are based upon physiological or biological characters, they will be
distinguished as "biologic forms." Various names have been proposed
for these different varieties, such as "special forms," by Eriksson; "bio-
logical species," by Rostrup; "sister species," by Schroeter ; and
"adapted races," by Magnus.

The truly morphological species, such as Puccinia graminis, Pers. or
Erysiphe graminis, DC, have still to be recognised; but each one may be
split up into a number of different forms, with distinct powers of infec-
tion.

Gradations of Specific Variation.

There is every possible gradation, however, between species which are
morphologically distinct and those which can only be separated on biologic
grounds. Puccinia graminis Pers., for instance, is recognised as an inde-
pendent species, because all the forms of this rust produce aecidia on the
barberry (except where it has lost this power), and the structural charac-
ters are always practically the same. It is regarded by Eriksson as a
collective species, in which the different members are so closely related, both
morphologically and biologically, that they are only separable into biologic
forms, and not to be distinguished as species. The forms on the different
hosts are not identical, but they constitute a series, each member of which
runs its course on definite host-plants, and is more or less strictly confined
to them. Arranged according to the principal ^ host-plants they are as

: — i, secalis; 2, avenae; 3, airae; 4, agrostidis; 5, poae; 6, tntici.

follows: — 1, secalis; 2, avenae;

The collective species known as Puccinia rubigo-vera (DC.) Wint. was
first divided in 1894 by Eriksson .and Hennirig1 into the two distinct species
of P. glumarum (Schum.) Eriks., the Yellow Rust, and P. dispersa, Eriks.
and Henn. the Brown rust. P. glumarum has not been found in Australia,
and it is not necessary here to refer to the various biologic forms into
which it has been divided. No aecidial stage has been found in connexion
with it.

P. dispersa, when first separated from P. glumarum, was split up into
four biologic forms by Eriksson, since he did not at that time consider
them sufficiently distinct to be designated species. These form
1, secalis; 2, tritici; 3, bromi; 4, agropyri. Further investigation,
however, led him in 1899 to raise the various biologic forms to the rank ot
independent species, and it will be interesting from our present stand-
point to consider the reasons given by Eriksson for differentiating between

Classification.

the biologic forms of P. dispersa and those of P. graminis. The name of
P. dispersa Eriks., was reserved for the form on rye, and it is separated
from that on wheat, for example,

i. The uredo
2. This form

regularly

on the following grounds

; to infect the host-plant, Se
appears several weeks earlier tha

growing alongside of each other

wheat, even when the host-plants are

3. The aecidiospores on Anchusa spp. only infect Secale, and the teleuto-
spores of this form can in turn only produce the aecidia on Anchusa.

4. The teleutospores are only capable of germination immediately after
they are formed, while those of the others only germinate in the following
spring.

It will be observed that these are only biological or physiological dif-
ferences> but in the two species recognised by Eriksson which occur in Aus-
tralia, viz., P. triticina and P. bromina, there are also structural differences
sufficient to separate them apart from other considerations. In P. bromina
the uredo-sori are much larger, and the bright orange uredospores are also

slightl\

larger.

It is in the teleutospores, however, that the difference is

most striking, and those of P. triticina are much nai rower, only being
about three-fourths that of the other.

P. hieracii (Schum.) Mart., as at first constituted, was a collective species,
but. like so many others, when the test of infective power is applied to
them, it had to be broken up into several. It has been proved by Jacky
that the common Puccinia occurring on species of Hieracium can only infect
this, and not other genera of Composites, so that the original name is

restricted to the species found on this genus. Other species of Puccinia
occur on Hieracium, but ithey are readily distinguished by their different
morphological characters. Similar results have been obtained with Puccinias
occurring on other genera, so that they also must be regarded as distinct
species.

It will be observed

that onlv biological characters

are here taken into

account, depending on the nature of the host-plant, but it is believed that
the closer and more careful investigation now rendered necessary will result
in structural differences being found, possibly of a more minute and less

striking character than

and overlooked.

The three t
classification.

formerly,

where they were hitherto unsuspected

pes selected for illustration will show the spirit of our

of
be

P. graminis is the type of a single species which is split up into a number

biologic fo

d the aecid

agreeing in the faot of producin

dia on the bar

turn

feet the

at first divided into biol

tuted, is the type of

distinct species, both on morphol

forms, which were afterwards found to h

and biol

grounds. And the

ginal P. hieracii is the type of a collective specie's, the members of which

are strict lv

ted in their infective

pow

genera

d their

n title to distinction as species is this biological p

The phenomenon of specialisation, first discovered bv Eriksson, has
thus caused us to revise our conceptions of species, and it will be necessary

at name such forms as possess
those which, although morpho-

in the fut

nclude under

distinct morphological characters, but

logical lv simil

are yet

fined to definite host

It will thus be

necessary to split up the old species of P. hieracii into several of equal rank,
and the one growing on Hieracium spp. will form a tvpe round which the
others may be grouped. Just as in the old species there were differences of

opinion as to the amount of

different

i-'

«
?

Classification.

■

biologic forms.

two allied forms, so in the new there is the same difficulty with the biological
characters, but it may be laid down, as a general rule, that the sharper the
distinction between two biologically different forms, the greater the reason
for calling them species, while the less distinctly marked would be called

Itt is all a matter of degree, and it must be left to the
good tact of the investigator, as Klebahn says, where to draw the line
between species and biologic forms.

Among the heteroecious rusts, there may be structural resemblances
between one generation and differences in the other. In that case, as a
matter of convenience, if the differences in the one generation are sufficiently
distinct, such should be designated species.

Biologic forms

in fact, may be regarded as incipient morphological

It is . not easy to explain how this has come about, but we

species, the physiological differences at present existing becoming ultimately

associated with morphological distinctions, which will mark them off as true

species.

may be sure that the explanation is to be sought, not along one line, but

along various converging lines.

The biologic forms of one and the same morphological species, such as
Puccinia graminis, may be supposed to have had a common origin, and the

most probable view is that the

original

form inhabited all the hosts on

which its descendants now live, and some of these descendants specialised
on one or other of these common host-plants. What caused them thus to
specialise? It may have been either through gradually becoming adapted
to certain of these host-plants in preference to others, or due to a spontaneous
change arising from internal causes, as expressed in the mutation theory
of De Vries.

This is Fischer's4 view, and Klebahn1, after examining the evidence.

sums up as follows: — "The manifold characters of the existing biological
species and races appear to have come about owing to the alternating exten-
sions and restrictions of the area of nutritive plants.. These changes, and
especially the restrictions of area, have been influenced by adaptation and
selection, but many observations indicate that internal developmental tenden-
cies have also played a part in determining the direation of the evolution."

The whole trend of this modern investigation is towards the recognition
of more deep-seated characters in the discrimination of species. One para-
site has become so thoroughly adapted to the physiological characters of a
host-plant that it cannot infect another, and so the plant on which the para-
site lives becomes a diagnostic feature of it. It is so much easier, and has
been so long the custom to be guided by morphological characters alone,
that physiological distinctions are not readily accepted, but now that they
are known, they must be recognised in some way, and the most convenient
is to incorporate them in the specific characters.

Again, the different stages of a fungus require to be known, in order
to classify it properly, and the mere knowledge of the uredo or aecidial
stage will not suffice for this purpose. Hence the life-history and infective
power, as well as structural characters, must all be taken into account in
fixing and determining species.

Systematic Arrangement.

CHAPTER XXI.
Systematic Arrangement and Technical Descriptions.

The different Rusts found in Australia are here named, described, and
systematically arranged. The names are necessary to distinguish one from
another, for, as George Eliot happily puts it — " The mere fact of naming
an object tends to give definiteness to our conception of it. We have then
a sign which at once calls up in our minds the distinctive qualities which

mark out for us that particular object from all others.

> >

[The descriptions

are necessarily technical, and give those characters which enable the species
to be discriminated from others, with the help of the illustrations. When
the described1 stage of any rust is enclosed in square brackets, this indicates
that it has not been found in Australia. The systematic arrangement
deals with the nine genera at present known, and arranges them according
to their natural affinities. Taking a general view of the entire order,
the following scheme of classification will be adopted, mainlv based upon
that of Engler and Prantl, in their Die natiirlichen Pfianzenfamilien: —

host

Order — Uredineae, Tul.
Fungi parasitic on higher plants and developing in the interior of th

a filiform, branching, septate mycelium

Spo

terminallv

terally from erect, transversely divided, crowded hyphae, and usually

of more than one kind

ium. The order

Teleutosp

germinating

a short promy

the third is not represented in Australia

be grouped in the following four families, of which

Teleutosp

stalked

Fam i. P

in groups

rows, or several cells in a

Teleutospores sessile, in columnar or filiform m
nartiaceae.

Fam

Cro

Teleutospores sessile or stalked, in one or two-layered waxy masses

Coleosporiaceae

Teleutospores sessile, in flattened one-lavered masses or loose

the

of their host. Fam. 4. Melampsoraceae

account of their economic

alwavs treated fi

portance, the Graminaceous rusts are

•st and the others are likewise grouped together undar
their respective families of host-plants, the order generally followed being
that of Baron von Mueller's Systematic Census of Australian Plants, start-
ing from the Grasses, and ascending to the higher forms.

Fam

Pucciniaceae

Since the great majority of our Australian Rusts belong to this family

be treated at greater length than any of the others

interesting

the different genera
differentiating

(the different forms

d it

hich the teleutospore assumes in

There are a number of genera based upon very

and it is not always easy to settle whether they

should be retained or rejected, but I have given all those which are clearly

It is a moot point whether such a genus as Diorchidium Kalch.,

distinct

should be retained, where the teleutospo.,

only the septum is longitudinal instead of

celled, as in Puccinia,

There are several

of Puccinia in which there is a tendency to an oblique, and

longitudinal sept
those species

as well

but th

hich the spore is distinctly divided longitudinally

retained for

germ-pore at the apex of

Systematic Arrangement.

From a consideration of the distribution of the Uredineae on their host-
plants, Dietel 12 has come to the conclusion that the genus Uredinopsis,
belonging to the Melampsoraceae, and occurring on ferns, represents the
oldest type of the Rusts, and that the Pucciniaceae may have developed
from the genus Melampsora. However that may be, we may assume that
the primitive form of this family was one-celled, and therefore the Uromvces
type forms the starting-point ; also that the Transition from the one-celled
to the two-celled spore as in Puccinia, was the next step in advance.

The actual evidence of this transition is seen not only in Puccinia hetero-
spora, B. and C, where the two-celled spore has not yet become completely
established, being mixed with a large number of unicellular spores, but in
such forms as Uromyces vesicul&sus, where there are occasional two-celled

Fisr.

spores, Fig. 157, U. tricorynes, Fig. 134, and U. politus, .Fig. 317.

Both Uromyces and Puccinia species occur on plants of the most diverse
character, and this would seem to indicate that many of the species came
into existence before that specialisation had begun, which resulted in the
genus Phragmidium, for instance, confining itself to the Rosaceae.

The principal genera are here arranged according to the nature of the
teleutospore, and the diagrammatic representation of the spores will show
at a glance the peculiarities of each : —

■
■

1
■

1. Uromyces, Link. — Teleutospore one-celled, with single germ-pore,

and solitary on its sitalk. Uredospores echinulate or warty all
over. Fig 16.

2. Hemileia, Berk and Br. — Teleutospore one-celled as in Uromyces,

but uredospore smooth on one face.

3. Uromycladium, McAlp. — Teleutospore one-celled as in Uromyces.

but the sporophore branching toward the apex produces either one
spore with a colourless sterile spore or cyst, or two or more spores
with or without a cyst. Fig. 17.

4. Puccinia, Pers. — Teleutospore two-celled, with transverse septum

and with only one germ-pore in each cell.

Fig. 18.

5. Gymnoconia, Lag. — Teleutospore as in Puccinia, but aecidia without

pseudoperidia.

6. Uropyxis, Schroet. — Teleutospore bicellular as in Puccinia, but each

cell with two or more germ-pores, and the teleutospore membrane
consisting of more than two layers. Fig. 19.

7. Diorchidium, Kalch. — Teleutospore two-celled, with longitudinal

septum and germ-pore at apex of each cell.

Fig.

20.

Fig. 16.

Fig 17.

Fig. 18

Fro. 1 P.

Fig 20

8. Gymno sporangium, Hed

Teleutosp

celled

ely three

d), and walls which ultimately form a common gel

mass, with usually several germ-pores in each

Fig

9. Hapaloph

Syd

Teleutoso^re three celled, and

of two basal cells

Fig.

alongside

of each othe

th a third on top

22.

Triph

Link. — Teleutospore three-celled, and consisting of

Fig. 23.

basal cell supporting two others alongside of each other

•

8.1

! I.

12.

Uromyccs.

Phragmopyxis, Diet.

Teleutospore three-celled in a longitudinal

series,

with outer laver swelling when moistened. Fig. 24.

Phragmidium, Link.

Teleutospore three or more celled in a longi-

tudinal series,
Fig. 25.

and outer layer not swelling when moistened.

Fi<;. 21.

Fig. 22.

»•

23.

Fig. *24.

Fig. 25.

Sphaerophragmium, M~0

Teleutospore consisting of a spherical

head of four to nine cells, and arising from a single cell by

tud

and transverse fission. Fig

14. Authority

Diet

— Teleuitospores forming a head of th

ght cells, borne on a single stalk, and arising from a simple
the formation of longitudinal septa.

Several sm

Fig. 27.

15. Ravenelia, Berk. — Teleutospores forming a more or less hemispher

manv-celled head

erselv and

dinally septate

arising from a stalk compounded of several hyphae

d with

sev

28.

Fig. 26.

Fig. 27.

Fig. 28.

Only four of these genera occur in Australia, and they will be dealt
with in the following order: — Uromyces, Uromycladium, Puccini a, Phrag-
midium.

UROMYCES Link.

mus the teleutospores are unicellular as well as the uredospores,
d this has sometimes caused the one to be mistaken for the other, but the

gle germ-pore in the former serves ito distinguish

spores are

way

echinulate or

ted. and wh

the

sides, the uredo-
:eleutospores are

usually smooth, they may also be striated or warted, but the generally

greater thickening of the wall, and the fact that the

earlv

colourless,

separate

All the different spore-forms mav

be present in the same host-plant, or they may be reduced to the teleutosp

the

folii) and beet rust (U. betae) all the spore
r, although the rust is very common, I hav<

alone. In

forms occur, but

seldom found the aecidia, and Plowright states that the aecidia are very
rarely found even in Britain.

Pea rust {Uromyccs pisi), with its aecidia on Euphorbia and its uredo-
spores and teleutospores on the pea, has not been found here, nor any other
heteroecious species of this genus.

•

Uromvces — Gramineae.

In carnation rust (U. caryofhyllinus) only uredo and teleutospores are
known, and this grouping of spore-forms is the most common with us.
Aecidia and teleutospores occur on the same host-plant in U. limosellae and
U. fuccinioides, the uredospores being unknown, and teleutospores alone are
found in U. bulbinis and V . diploglottidis. The rusts on clover, beet, and
carnation are the best known, and since the carnation is attacked at all
stages of its growth, it has suffered considerably from this disease. _

General Characters. — Spermogonia mostly globose, immersed, with
conical projecting neck.

Aecidia immersed, finally cup-shaped, with well-developed pseudo-
peiidium ; aecidiospores without distinct germ-pores.

Uredospores solitary on their stalks, with several usually distinct germ-
pores.

Teleutospores unicellular, pedicellate, only one produced from each
sporophore, with a single germ-pore at apex.

Sporidiola hyaline, ovoid, ellipsoid, or almost kidnev-shaped.

Australian species, 27

GRAMINEAE.

l. Uromyces danthoniae McAlp.

Danthonia.

I. Aecidia amphigenous, densely crowded in relatively large clusters,

bright orange ; pseudoperidia cup-shaped, ivory white, margin
regular, upright, very finely toothed, up to 320 u in diam. ;
pseudoperidial cells persistent, elongated and polygonal, with
striated margin.

Aecidiospores subglobose to polygonal, orange, average 16 u diam.

or 16 x 12

II. Uredosori minute, erumpent, somewhat scattered, ruddy brown.

Uredospores pale yellow to orange yellow, broadly elliptical to
ovate, finely echinulate, with as many as six scattered germ-pores
on one face, 27-32 x 23-26 li.

III. Teleutosori chocolate brown to black, elongated, running in lines,

long covered by epidermis.

Teleutospores at first intermixed with uredospores, pear-shaped to
ovate or broadly fusiform, smooth, with thickened apex, 30-41 x
20-26 li, average 33 X 22 fx ; pedicels elongated, persistent,

slightly tinted adjoining spore, up to 96 fx long.

I. on leaves of Danthonia sp. II., TIL on leaves of Danthonia
semiannularis R. Br.

Victoria — Bacchus Marsh, I. Near Melbourne, Ardmona, Portland,

Killara, Leongatha, Rutherglen, Nagambie, Kergunyah, ifcc,
II., III.

Tasmania— Domain, Hobart, Nov., 1894, II., III. (Rodway).

The aecidium found on a species of Danthonia at Bacchus Marsh is
described in connexion with this species as a matter of convenience.

Puccinia graminella (Speg.) Diet, and Holw., occurring on a species of
Stipa in Argentine, Chili, and California, is the only instance hitherto known
where the aecidium-stage is found on a grass. Teleutospores are developed
on the same mycelium alongside of the aecidia, and compressing them on
either side.

Uromyces — Gramineae.

The aecidia are described as follows : — " Epiphyllous, and

arranged

loosely in interrupted lines ; pseudoperidia persistent, composed of oblong
cells, with margin irregular and lacerated. Aecidiospores elliptic or ovoid,
21-29 x 18-21 fx

The aecidia on Danthonia are quite distinct. They occur on both sur-
faces of the leaf, are arranged in dense clusters, the margin of the pseudo-
peridium is regular and very finely toothed, and the aecidiospores are con-
siderably smaller.

There is a good deal of confusion over the species of Danthonia in Aus-
tralia, and they probably require to be dealt with by a specialist. The late
Baron von Mueller, in his Census, included D. pallida R. Br., D. semiannu-
laris R. Br., D. pilosa R. BrM <fcc, under D. penicillata F.v.M. Following
the Index Kewensis, these three species will be retained, but D. penicillata
F.v.M., will be regarded as a synonym.

Darluca filum Cast., is common on uredosori.

(Plate XVI., Fig. 131 ; Plate J.)

. Uromyces ehrhartae McAlp.

Ehrharta

McAlpine, Agr. Gaz. N.S.W, VI., p. 855 (1895).
Sacc. Syll. XIV., p. 405 (1899).

Uredo ehrhartae McAlp.

II. Uredosori on both surfaces of leaves and on sheath, minute, oval to

elongated linear, sometimes confluent, at first covered then naked,
yellowish-brown to orange.

Uredospores subglobose to oval, orange, finely echinulate, 3-4

24 x

ed
19

-pores on one face, 21-25 x 18-20 //,

average

III. Teleutosori similar, up to 1 mm. long, confluent in lines, chocolate

brown to black.

Teleutosp

pale chocolate b

to chest

brown, only

9 /*),

edospores intermixed, hooded and thickened at apex

ther conical or truncate,

and size, elongated or short and
fusiform or oblong, 19-32 x 13-19
pedicels persistent, tinted, elongated, up

iable

stout, subglobose to

shape
liptic,

average

26 x 1

On Ehrharta stipoides Labill

Microlaena stipoides R. Br

Victo

Near Melbourne, Killara, My

June-March

Kerguny

ifcc

Uredospores common but teleutospores rather

.e'hbourhood of

son are sometimes rathe
and intermixed with ur

Melbourne during December

sparse

gh in

the

edosori on those

and January the teleuto-
ally on the lower withered leaves,

still partially green.

uredo-stage was found at first by Mr. Robinson, but latterly at
March, and Kergunyah, in November, he secured the teleutospc

Darluca filum Cast., is very common on uredosori.

Only the

Killa

(Plate XVI, Fig. 132.)

Uromyces — Gramineae, Liliaceae. 87

3. Uromyces tenuicutis McAlp

Sporobolu

II. Uredosori on both surfaces of leaves, but mostly on upper, and on

stems, elliptic to oblong, orange-yellow, pulvinate, generally
minute, but may reach a length of 1 mm., at first covered by
epidermis, then erumpent, pulverulent.

Uredospores orange, ellipsoid to ovoid, spinulose, rather large,
with two distinct germ-pores on one face, equatorial or nearly so,
28-40 x 19-24 11, average 32 x 22 /w.

III. Teleutosori on lower surface of leaf, minute, pulvinate, covered by

epidermis.

Teleutospores yellowish-brown, smooth, obovate to oblong or piri-
form, irregularly rounded or truncate, or even pointed at apex,
and slightly thickened, sometimes broader than long, 22-35 x

16-25 ju, average 28 x 20 fx ; pedicel sometimes persistent
tinted, commonly 20-35 ll long, but often much shorter.

On leaves and stems of Sporobolus indicus R. Br.

Victoria — Near Melbourne, Aug. — March.

U. sporoboli E. and E., found on leaves of Sporobolus asper Kunth, in the
United States, has no uredo-stage, and the teleutosori are soon naked, while
the teleutospores may be subglobose and provided with long pedicels.

D arluca jilum Cast., is very commonly associated with the uredospores.

(Plate XVI., Fig. 133.)

LILIACEAE.

Bulbine.

4. Uromyces bulbinis Thuem.

Thuemen in Flora, p. 410 (1877).

Cooke, Handb. Austr. Fung., p. 409 (1892).

Sacc. Syll. VII., p. 572 (1888).

III. Sori amphigenous, small, densely gregarious, concentrically disposed

in large circles, covered by the epidermis, firm, rather concave,
brown.

Teleutospores clavate or oblong clavate, mostly rather acute at
the apex, and narrowed at the base, pedicellate, epispore smooth,
rather thick, especially at the apex, 30-36 x 20-22 li ; pedicel
deciduous, unequal, straight or slightly curved, hyaline to yellowish
brown, 12 x 4li.

On living leaves of Bulbine bulbosa Haw.

Victoria — Omeo (Morrison).

New South Wales — Upper Macquarie II

5. Uromyces tricorynes McAlp.

Tricoryne.

McAlpine, Agr. Gaz. N.S.W., VI., p. 756 (1895).
Sacc. Syll. XIV., p. 283 (1899).

I. Aecidia on pale spots in small clusters, amphigenous ; pseudoperidia

cup-shaped, with white edges irregularly torn and reflexed.

Aecidiospores bright orange, subglobose to somewhat polygonal,

smooth, average 20 x 1 7 li

438.

Uromvces — Orchidaceae.

II. Uredosori on pale green patches, brick coloured to orange,

amphigenous, but more common on under surface of leaves,
elongated oval to elliptical, bullate, solitary or in groups, not
confluent, at first papillate, then bursting through and surrounded

by ruptured epidermis.

Uredospores orange, subspherical to elliptic or ovate, finely
echinulate, relatively thick-walled, with two to three germ-pores

on one face, average 20-23 /i diam. or 22-25 x 19-20 fx

III. Teleutosori blackish on ruddy brown spots,

elongated

and

confluent, sometimes completely enveloping stem, at first bullate,
finally with greyish epidermis around or over them in shreds and
patches, generally 2 mm. long, sometimes up to 3 mm.

Teleutospores variable in form, oval, ovate, or somewhat
globular, yellowish - brown to chestnut - brown, smooth, apex
thickened (up to 8 fi), round or conical, 25-32 x 20-29 yu, average
30 x 25 /u ; pedicels persistent, pale yellow, especially towards
spore, up to 80 /x long and 9 \i broad.
On leaves and stems of Tricoryne elatior R. Br.

Victoria — Near Melbourne, Oct., 1892, II., III. (Robinson).

Rutherglen, July, Oct., 1893, 1., II., III. Murramurrangbong
Ranges, Nov. 1902 (Robinson). Nagambie, Nov., 1904, II., III.

New South Wales — Richmond, Oct. (Musson).

I. on both surfaces of leaves, July. II., on stems and leaves, not very
common in July, but prevalent in October and November. III. on stems
and branches and both surfaces of faded leaves, very common in July,
forming dark swollen patches.

Several two-celled teleutospores occurred, coloured similarly to the
ordinary teleutospore and thickened at apex. The spores were constricted

t septum, which might be about the middle or towards the base.

were longer than the ordinary spore, and measured about 37 X 27 p
pedicel was occasionally rather lateral.

Darluca filnm Cast., occurred on the uredosori.

(PlateXVI, Figs. 134, 13 5.)

They
The

ORCHIDACEAE.

6. Uromyces microtidis Cooke.

Microtis.

Cooke, Grev. XIV., p. 12 (1885).
Cooke, Handb. Austr. Fung., p. 332 (1892).
Sydow, Ann. Myc. I., p. 324 (1903).
Sacc. Syll. VII., p. 579 (1888).

Sori amphigenous, loosely arranged in irregular groups, minute, p

form, brown, pulverulent, girt by the ruptured epidermis.

II. Uredospores globose, subglob

ate to warty, yellowish-bro

III. Teleutosp

termixed with

piculus, rough with warts, bro
. length of 41 ut average 32 x

-30 x 1

redosp

psoid, minutely acule-
ovate, with hyaline

On living leaves of Microtis porrifolia R. B

35 x 17-25 fi, but may reach
•edicel very short, hyaline

New South W

Bullahdelah, II., Ill

Uromyces — Orchidaceae

Only the teleutospores were described by Cooke, but in a po

•tion of the
material kindly supplied by Massee there were numerous uredospores

termixed with the teleutosp

Sydow 2 has also found the uredospores

th teleutospores intermixed, on specimens of the same host-plant sent from

Chatham Island, New Zealand.

If we compare the three n
C hiloglottis , and Thelymitra, it
but are absent, as far as known, from the oth

sts found on the Orchid
is found that aecidia occi

genera,

otis.

a occur on ChiloglotttSj

The teleutospores are

all provided with a hyaline apiculus, but in U. microtidis the epispore is very

gh and knobby, and

size of the spore not generallv exceeding 35

long, while in U. thdymitrae the epispore is generally thicker, and the spore
polygonal and irregularly warted and altogether larger. The rust on Chil.o-
glottis generally resembles that on Microtis, but the teleutospore is larger.

The following table will show the points of resemblance and difference in
the teleutospores : —

Size.

Uromyces microtidis

U. orchidearum

• •

25-35 x 17-25 ft
30-50 x 1 9-24 fi

U, thelymitrae

1 - *

35-45 x 25-30 /x

Epispore.

Generally prominently
warted all over

Irregularly warted and
warts not so pro-
minent

Less closely but more
prominently warted,
hence polygonal

Thickness.

3-4//

3-4 fi

5-6 ju

Apiculus.

Generally
co iioid
Generally

conoid

Generally
obtuse

(Plate XVI., Fig. 139.)

Chiloglottis,

7. Uromyces orchidearum Cke. and Mass.

Cooke and Massee, Grev. XVI., p. 74 (1888).
Cooke, Handb. Austr. Fung., p. 332 (1892).
Morrison, Vict. Nat. XL, p. 90 (1894).
Sacc. Syll. VII., p. 580 (1888).

I. Aecidia hypophyllous, scattered, cup-shaped ; pseud operidial cells

polygonal to wedge or pear-shaped, finely notched, 32-40 p.

Aecidiospores pale yellow, subglobose to shortly elliptical, verru-
culose, 16-20 hi or 18-22 x 15-20 /u.

II., III. Sori mostly epiphyllous, bullate, at length erumpent, golden-
brown.

II. Uredospores elliptic to ovoid, honey-yellow to golden-yellow, echinu

late, with several germ-pores, 26-30 x 16-20 /x.

III. Teleutospores at first intermixed with uredospores, old gold colour

variable in shape and size, elliptic to ovoid or oblong, with pro-
minent hyaline apiculus (up to 9 jjl long), usually with irregularly

which is 3-4 /u thick and brightly coloured,

•

warted epispore;

30-51 x 19-24 /i, average 36 x 21 p ; pedicel hyaline, attenuated

downwards, up to 77 ^ long.

D 2

£0 Uromyces — Orchidaceac, Scrophalariaceae.

On leaves of Chiloylottis diphylla 11. Br., II., III.

New South Wales — Mt. Victoria (Hamilton

■

Victoria— Oakleigh, July, 1894 (Morrison
Tasmania— Bellerive Swamp, May, 1897 (Rod way

On C. yunnii Lindl., I., II., III.

Victoria— Oakleigh, Aug., 1892 (Morrison).

■

In the original description of Cooke and Massee only teleutospores are
>rded. but uredospores were also obtained from the original material

labelled in Cool

Sometimes there are abnormal elongated teleutospores up to 60 yu long

Several bicellular spores were met with similarly coloured to the normal
teleutospores, constricted at septum, irregularly warted, with prominent .1
hyaline apiculus, 59 x 28 /*.

(Plate XVI., Fig. 138 ; Plate XL., Fig. 303.)

Thelymitra

8. Uromyces thelymitrae McAlp.

deciduous, up to 1 1 p broad adjoining spore
On Thelymitra antennifera Hook. f. and T. flexuosa Endl.

Victoria— Near Melbourne, Sep. and Oct. (C. French, jun.).

An aecidial stage has been found in Java, by Raciborski, on Thelymitr
javanica Blume, but it has probably no connexion with our native species.

(Plate XVI., Figs. 136, 137.)

SCROPHULARIACEAE.

9. Uromyces limosellae Ludw.

Limo sella.

Ludwig in Dietel, Hedw., XXVIII., p. 182 (1889).
Sacc. Syll. IX., p. 293 (1891).

I. Aecidia amphigenous, scattered or gregarious ; pseudoperidia with

whitish laciniate margin, not deeply incised.

Aecidiospores round to angular, hyaline, smooth, about 15 u
diam. r

III. Teleutosori mixed with the aecidia, often crowded, long covered

by epidermis, pulvinate, dark brown.

Sori on leaf and sheath, scattered or gregarious, bullate, ellipsoid, |

yellowish to brownish, compact, surrounded or almost covered by
ruptured epidermis, except for a narrow slit, up to 1 J mm. long.

II. Uredospores ellipsoid to obovate or pear-shaped, golden-yellow,

echinulate, with as many as four equatorial germ-pores, 25-32 X

18-21 p.

III. Teleutospores ellipsoid to oblong, or polygonal, chestnut-brown,

relatively thick-walled, irregularly knobby, with hyaline apiculus
35-45 x 25-30 /u, average 36 X 26 fi ; pedicel hyaline, short,

Uromyces — Goodeniaceae, Rubiaceaz. 91

Teleutospores obovate, oblong or clavate, rarely round, with
yellowish-brown, thick, smooth wall, strongly thickened at apex,
and for the most part paler, 32-40 x 18-22 11 ; pedicel as long as
or shorter than teleutospore.

On leaves of Limosella aquatica L.

S. Australia — Kangaroo Island (Tepper).

It differs from U. scrophulariae (DC), to which it is otherwise allied,
in the size and colour of the spores.

Dietel observed two bicellular teleutospores among the ordinary ones in
& spore layer.

Specimen not seen.

GOODENIACEAE

10. Uromyces puccinioides Berk, and F.v.M

Selliera, Scaevola

Berkeley and Mueller, Linn. Journ. XIII., p. 173 (1872).
Cooke, Handb. Austr. Fung., p. 332 (1892).
Sacc. Syll. VII., p. 585 (1888).

I. Aecidia aggregate, on brown orbicular spots, arranged in a circinate

manner, opposite ; pseudoperidia with abbreviated margin.
Aecidiospores orange, subglobose, 16-19 X' 13-16 fx.

III. Teleutosori bullate, intermixed with aecidia or surrounding them

as a ring, at first covered with greyish glistening epidermis, finally
naked.

Teleutospores brown, apiculate, sometimes with the apex oblique
or dentate, also thickened and rounded or flattened, 40-44 x
18-23 fx, average 41 x 22 11 ; pedicels hyaline, persistent, up to
48 ix long.

On leaves and flower stalks of Selliera radicans Cav.

Victoria — Near Melbourne, Nov. — June. Wimmera Flats, May,

1897 (Reader). Phillip Island, Jan., 1900. Portland, Jan.,
1901. Point Cook, May, 1902 (C. French, jun.). Sandring-

ham, Nov., 1905 (Robinson).

South Australia — Glenelg (Holdfast Bay), 1854 (Berkeley

Tasmania — Bellerive Swamp, Dec, 1890, and May, 1897 (Rod-
way 1).

On Scaevola sp.

Queensland — St. George (Wedd) (Bailey

It is recorded in Cooke's Handbook as being found on Goodenia, but t
are no certain indications of this. It occurred on Goodenia herpy
Schlecht., which, however, is a synonym of Selliera radicans.

(Plate XVIL, Fig. 140.)

RUBIACEAE.

11. Uromyces asperulae McAlp.

I McAlpine, Agr. Gaz., N.S.W., VI., p. 851 (1895).

Sacc. Syll. XIV., p. 276 (1899).

A sperula.

II. Uredosori hypophyllous, minute, bullate, bursting through epidermis.

Uredospores globose, subglobose or ovate, finely echinulate,
orange-yellow, single germ -pore visible on one face, 21-25 X
20-21 a.

Uromyces — Leguminosae.

III. Teleutosori on stems

ds their base, elongated oval, dark

brown, appearing almost black, confluent, erumpent

Teleutosp

first intermixed with

dosp

brown to chestnut-b

smooth

'es, yellowish-
>r elliptical to

broadly clavate, apex much thickened (11 /*) and rounded, some-
times tapering or flattened, 27-37 x 15-19 /i, average 30 X 18 At

pedicels long, persistent, hyaline, up to 45
On stems and leaves of Asperula oligantha F.v.M

Victoria— Ardmona, May to Oct., and in moist places all the
year round (Robinson).

The host

is given

Asperula scoparia Hook, f., in the Index

Keivcnsis, but the synonymy seems

confused. A. oligantha F.v.M

is first gi\

as Rub

Miq

synonym of Asperula scoparia Hook, f

ticola is given as a

(Plate XVII., Fig. 141.)

LEGUMINOSAE

12. Uromyces appendiculatus (Pers.) Link.

Link, Obs. II., p. 28 (1825).
Sacc. Syll. VIL, p. 535 (1888).

Uromyces phaseoli (Pers.) Wint.
[O. Spermogonia on minute spots, whitish.]

Vigna-

Aecidia densely crowded in small

annular groups ; pseudoperidia
sliortly cylindrical, whitish, with deeply cut reflexed margins.

Aecidiospores polygonal, finely warted, colourless, 17-32 x

14-23 /x.]

II. Uredosori on both surfaces of leaf, round, pale cinnamon brown,

pulverulent, solitary or crowded and confluent, soon naked and
surrounded by ruptured epidermis.

Uredospores yellowish brown, ellipsoid to ovate, finely echin-
ulate, 23-32 x 17-22 p.

III. Teleutosori dark-brown, almost black, easily detached, otherwise

like uredosori.

Teleutospores dark brown to chestnut brown, subglobose to ellip-
soid or oblong, smooth, scarcely thickened at apex, with broad pale
flattened papilla, 27-37 x 19-25 /u ; pedicel hyaline, persistent
elongated, up to 70 p long.

On Cowpea (Vigna catjang Walp.).

New South Wales.

Richmond, May, 1905 (Musson)

The distinction between the teleutospores of U. fabae and U. appendicu-
latus is very marked. In the one the apex is thickened up to 9 /*, while in
the other the thickening is not perceptible.

(Plate XLIL, Fig. 306.)

■

Uromyces — Leguminosae

Acacia

13. Uromyces bicinctus McAlp.

II., III. Sori ruddy-brown, gregarious often confluent, raised, compact,

rupturing epidermis irregularly.

II. Uredospores yellowish-brown, clavate to oval or ellipsoid, rounded

at apex and slightly thickened, densely warted and warts arranged
in regular longitudinal lines, with distinct germ-pores, generally
arranged in two bands about one-third length of spore from either
end, and usually three to four in each band on one face, 30-40 x
13-18 /j,, occasionally reaching a length of 45 ^t, and variable in
breadth, average 34 x 15 fi ; pedicel similarly coloured to spore or
paler.

III. Teleutospores intermixed with uredospores, pallid but thickened
considerably at apex and brown (8-11 f.i), smooth, ellipsoid, and rounded at
both ends, occasionally prolonged into one or two processes at apex, 28-34

x 17-23 /j. ; pedicel persistent, hyaline, short.

On phyllodes and pods of Acacia fasciculi f era F.v.M.

Queensland — Rockhampton, 1867 (from host-plant in National

Herbarium, Melbourne).

This species approaches somewhat to U. phyllodiorum (B. and Br.),

McAlp., in the uredospores, but they are shorter and much narrower, and

distinguished at once by the two bands of germ-pores, on account of which

the specific name is given. The teleutospores are also much thickened at

apex and destitute of processes as a rule, although occasionally two may
appear.

(Plate XLIIL, Fig. 318.)

Vicia

14. Uromyces fabae (Pers.) De Bary.

De Bary, Ann. Sci. Nat. Ser. 4, XX. (1863).
Sacc. Syll. VIL, p. 531 (1888).

[O. Spermogonia yellowish, in small groups on leaves and stem.]

[I. Aecidia scattered, in rings or orbicular patches ; pseudoperidia

short, slightly prominent, fiat, with torn white edges.

Aecidiospores subglobose, orange, finely echinulate, 16-26 /t
diam.]

II. Uredosori amphigenous, roundish, pale brown, powdered, scattered,

often confluent, soon naked.

Uredospores subglobose or ovate, ochraceous, echinulate, with
three equatorial germ-pores on one face, 20-30 x 17-20 fx.

[III. Teleutosori rounded on the leaves, more abundant and elongated

on the stems, often confluent, blackish-brown.

Teleutospores variable in form, obovate or broadly clavate,
smooth, dark-brown, apex darker, thickened (8-10 ^u), and rounded,
truncated or conical, sometimes with colorless papilla, 24-47 x
17-30 /i; pedicels hyaline or pale brown towards spore, persistent,
up to 110 fi long.]

On stems, leaves, and pods of Broad Bean (Vicia faba L.).

New South Wales.— 1894 (Cobb9).
Queensland.— Gladfield (Gwyther) (Bailey19).

Vromyces — L eguminosae .

This rust has not been found in Victoria, and although it is said to be-
very common in New South Wales, on applying to Dr. Cobb for specimens,
he informed me that they had all been destroyed by insects. Only the
uredo-stage was found in New South Wales, and the exact species is still

doubtful.

(Plate XLIL, Fig. 307.)

Acacia*

15. Uromyces fusisporus Cke. and Mass.

Cooke and Massee, Grev. XVI., p. 2 (1887).
Cooke, Handb. Austr. Fung., p. 331 (1892).
Sacc. Syll. VII., p. 555 (1888).

Sori amphigenous, solitary or in groups and then confluent, elliptic to>

discoid, dark-brown or black, erumpent and girt by the ruptured
epidermis.

II. Uredospores golden-brown, fusiform, obtusely warted, with acute

and more or less hyaline apiculus, 3-4 equatorial germ-pores on
one face, 57-77 x 17-25 u, average 62 X 23//.

pedicels deciduous, hyaline, elongated, up to 96 fi long, with
septum at a short distance beneath spore.

On phvllodes of Acacia salicina Lindl., and A. neriifolia A. Cunn
A. retinodes Schlecht.

Victoria— Dim boola, Dec, 1895, and May, 1897 (Reader).
New South Wales— 1902 (Maiden).

On Acacia sp.

Queensland — Islands of Torres Straits, June, 1897 (Bailey18'19

Portion of the original material named in Cooke's handw

National Herbarium, and our description is based upon that. In the original
description by Cooke and Massee the uredospores were unfortunately mis-
taken for teleutospores and vice versa. The uredospores are easily known
from being obtusely warted and with equatorial germ-pores, apart altogether
from size and shape.

The teleutospore is solitary at the apex of the pedicel, but the sep-
tum at a short distance from the spore foreshadows the Uromycladium with
a colourless vesicle or cyst produced laterally immediately beneath septum.

The nature of the teleutospore and the presence of a septum in the stalk
was so suggestive of Uromycladium that the material was specially ex-
amined to see if more than one spore was borne on a stalk, but the most
careful search failed to reveal any indications that this was the case.

III. Teleutospores intermixed with uredospores, ruddy brown, globose

to depressed globose, epispore rather thin, scarcely 2 /x thick,,
somewhat polygonal seen from above and then germ-pore veiy
prominent, 25-30 \i diam. or 16-21 x 25-35 \x, average 18 x 28 fi ;

(Plate XIX., Figs. 158-160.)

16. Uromyces hardenbergiae McAlp.

Hardenbergia*

Sori on under surface of leaf, brown, crowded, globose to ellipsoid, often

confluent, bullate, with ruptured epidermis, which usually remains
in large patches.

Uromyces — L eguminosae

II. Uredospores golden-brown, elliptic to ovoid, echinulate, relatively

thick-walled, with three distinct equatorial germ-pores on one face,
25-35 x 20-22 n, average 27 x 20 p.

III. Teleutospores intermixed with uredospores, very sparse, ruddy-
brown, thick-walled, smooth, ellipsoid to obovoid, slightly
thickened at apex, with hyaline apiculus and showing germ-pore,
25-28 x 18-21 fi, average 26 x 19 ft; pedicel persistent, hyaline,

ated, up to 35

On leaves of Hardenbergia monophylla Benth.

Victoria — Kergunyah, Dec, 1903, II., III. (Robinson). Frankston,

January and August, 1904, II. (Robinson).

The teleutospores were exceedingly rare, and somewhat resembled the
uredospores, but slightly smaller, with ruddy smooth epispore and hyaline
apiculus.

This species persists all the year round.

Darluca filum Cast., very prevalent on uredosori.

(Plate XVII., Figs. 143-145; Plate G., Fig. 34.)

A cacia.

17. Uromyces phyllodiorum (B. and Br.) McAlp.

Berkeley and Broome, Linn. Trans. II., p. 67 (1883).
Cooke, Handb. Austr. Fung., p. 331 (1892).
Sacc. Syll. VII., p. 556 (1888).

Melampsora phyllodiorum Berkeley and Broome, Linn. Trans.,

II., p. 67 (1883).
Uromyces digitatus Winter, Rev. Myc, p. 209 (1886).
Uromyces phyllodiae Cooke and Mass. Grev. XVIL, p. 70

(1889).

| O. Spermogonia brown at first, becoming shining black, numerous,

I small, blister like, seated in centre of swollen, brownish, scattered,

I discoid, amphigenous tubercles, varying in size from 1 to 4 mm.

II., III. Sori surrounding spermogonia or alone, ruddy brown to dark-
brown, round or elongated, crowded, often confluent, compact,
raised, girt by the ruptured epidermis, which bursts irregularly.

II. Uredospores golden yellow to golden brown, oval to ellipsoid,

rounded at apex or bluntly pointed and slightly thickened, densely
warted, and warts arranged in regular longitudinal lines, with
distinct equatorial germ pores, 3-4 on one face, and even reaching
to 6, 35-54 x 16-25 //, average 38 x 22 /u, with elongated
and hyaline pedicel.

III. Teleutospores at first intermixed with uredospores, lemon yellow,

and sometimes almost colorless towards base, fusiform to wedge-
shaped, thickened at apex, and bearing finger-like erect or divari-
cate processes, sometimes a single process, or bifid, or several,
60-70 x 14-20 /i, but may even reach a length of 86 ^ ; pedicels
long, persistent, hyaline ; paraphyses intermixed with teleuto-
spores or with uredospores accompanied by teleutospores, elongated,
variously shaped, but generally somewhat cylindrical, thickened
and rounded at apex, and tapering sometimes into a slender
filament at base. 70-93 X 6-10 u.

Vromyces — Legaminosae.

On phyllodes of Acacia notabilis F.v.M

S. Austra

On phyllodes of

Victoria-

Near Gawler, July, 1885 (Tepper) (Ludwig2).
daUachiana F.v.M., and A. penninerms Sieber
Alps, near Bright, Dec, 1904 (C. French, jun.)

On phyllodes of Acacia dealbata Link.

Victoria— Orbost, Dec, 1905, II. III.

On phyllodes of Acacia penninervis Sieber, A. microbotrya
neriifolia A. Cunn. in National Herbarium, Melbourne.

New South Wales— Twofold Bay.
Queensland — Brisbane River.

On phyllodes of Acacia pruinosa A. Cunn.

Benth. and A,

New South
On Acacia sp.

— Gosford, Jan., 1906 (Frogg

Queensland — Brookfield, Brisbane River (Bailey1'19).

New South Wales

(Cobb

The appearance presented by this rust varies according to the presence or
absence of spermogonia. In the Queensland specimens there are black
discoid tubercles with spermogonia in the centre and surrounded by the sori,
while in the Alpine specimens the numerous sori are scattered over the green
surface of the phyllode.

The size of the uredospores is

given

by Winter as 32-35 x 20-25 py

reach a length of 54

but there are occasional elongated forms which may
They somewhat resemble those of Uromycladium notabile, but the markings
on the epispore are much closer together.

The teleutospores are very characteristic, but very variable both in shape
and size. The apex may be prolonged into a single process, or there may be

quite a number, at least up to 6.

As regards size

they are generally

elongated, and the measurements given are the mean of a number taken from

spores ending in a single process. They germinate at once without a period
of rest. The specific name of digitatus is so appropriate that it is unfortu-
nate it has to be set aside in obedience to the law of priority.

This species was first described by Berkeley and Broome in a list of
fungi from Brisbane, Queensland, as Melampsora phyllodiorum in 1883, and
drawings accompany the description. Specimens of the original material
have been kindly supplied to me by Mr. F. M. Bailey, Queensland State
Botanist, and uredospores and teleutospores from these are shown in
PI. XXV., Figs. 218-220. There is no doubt as to its identity.

The following is* the original description of Berkeley and Broome

Sori in amphigenous tubercles ; spores

rather fusiform, 55-58 f.

elongated, uniseptate, fusiform, even, 22

long,

arising

from delicate filaments,

granulated, mixed with others which are

The longest uredospore I

i long.

found was 54 ^ long, and the uniseptate spores of Darluca filum Cast., were
also very common.

Next, Winter, in 1886, described the same fungus from S. Australia as
Uromyces digitatus, the teleutospores being recorded for the first time.

Then Cooke and Massee, in 1889, described a Uromyces phyllodiae from
Brisbane as follows : — " Sori minute, orbicular, compact, brown, crowded on
elliptic, bullate, brown spots, 3-5 mm. long, at length naked, not pulverulent.
Uredospores not seen. Teleutospores elliptic, obtuse, rarely apiculate,
brown ; epispore minutely warted, rather thick, hyaline, thickened at the
apex, 40-45 x 16-18 ^/." A portion of the original material *ras supplie
by Mr. Bailey, and both uredospores and teleutospores have been found.

Uromyces — L eguminosae.

Cooke and Massee mistook the uredospores for teleutospores, but the
existence of several germ-pores, together with their general appearance,
indicate their true nature. Spermogonia were present in this material.

Finally Dr. Cobb,10 in 1897, described the uredo stage of Melampsora
phyllodioruniy and gives a drawing of the uredospores with an average size of
41 x 17 fi. He also found the perithecia of Darluca filura Cast., among
the sori, but their true nature was not recognised.

(Plate XXV., Figs. 218-228.)

18. Uromyces trifolii (Alb. and Schw.) Winter.

Winter, Die Pilze, L, p. 159 (1884).

Trifolium

Cooke, Handb. Austr. Fung., p. 330 (1892).
Sacc. Syll. VII., p. 534 (1888).

0. Spermogonia honey-coloured, disposed in small clusters.

Spermatia hyaline, globose, about 3 jj, diam.

1. Aecidia in circular clusters, on pallid spots ; pseudoperidia cup-

shaped, with a white laciniate margin.

Aecidiospores subglobose, ellipsoid or irregular, very finely
verrucose, pale orange, 14-23 /j. diam., or 22-25 x 14-16 //.

II. Uredosori rounded or elliptic, scattered, surrounded by the

torn epidermis, ruddy brown.

Uredospores irregularly globose or shortly elliptic, echinulate,
golden-brown, with two equatorial germ-pores on one face, 22-26
X 18-20//.

III. Teleutosori smaller and rounded on the leaves, larger and elon-
gated on the petioles, dark brown, almost black, bullate, long
covered by the leaden epidermis.

Teleutospores generally intermixed with uredospores, ellipsoid,
globose or pear-shaped, thickened at the apex with a small pale
wart, smooth, dark-brown, 22-30 X 19-22 ^u, average 24 X 20 \x\

pedicels long, hyaline, deciduous.

On leaves, petioles and stems of Trifolium repens L.

Victoria — Near Melbourne, Apr., I., II., III. Sep., I., III. Bunyip,

Nov., I., II. Nyora, Apr., I., II., III. Murramurrangbong
Ranges, Jan., I., II., III.

New South Wales— (Cobb12).
Tasmania — (Rod way1).

All the stages were found together in January and April, and occurred
on petiole, leaf, and stalk of inflorescence. The uredospores and teleutospores
were very common, but the aecidiospores were not so common, and they were
found also on the calyces. The leaf stalk was often much swollen and dis-
torted. The spermogonia usually precede the aecidia. The teleutospores
are considered to germinate only after a period of rest, but they were ob-
served in April, to germinate freely while on the leaf.

The mycelium may be perennial, and Dietel* considers this a common
occurrence.

(Plate XVII., Fig. 142 ; Plate G., Fig. 32.)

Uromyces — Polygoriaceae.

POLYGONACEAE

Muehlenbeckia.

1<). Uromyces politus (Berk.) McAlp

Berkeley, Linn. Journ. XIII., p. 174 (1872).
Berkeley and Broome, Linn. Trans. II., p. 67 (1883).
Cooke, Handb. Austr. Fung., p. 342 (1892).
Sacc. Svll. VIL, p. 833 (1888).

Roestelia polita Berk.

I. Aecidia on dark- purplish, elongated patches, generally arranged in

lines which may be parallel, bursting through epidermis, bright
orange, becoming ivory- coloured with age, cylindrical to slightly
compressed, straight, averaging 1| mm. high; pseudoperidia

polished, smooth, with white, narrow, toothed margin ; peridial

oblong to angular, with striated margin, 32-35 jj. Ion

Aecidiospores ochraceous, variable in size and shape, generally
ellipsoid to ovoid or oblong, finely echinulate, 24-30 x 19-22 /i.

III. Teleutosori on the same discoloured area as the aecidia and

associated with or distinct from them, solitary or confluent,
brownish to blackish, bullate, elongated to oval, surrounded by the
raised and ruptured epidermis, compact, 1-2 mm. long

Teleutospores yellowish-brown, smooth, ellipsoid to oblong,

rounded or pointed and apiculate at apex, thickened (6-8
occasionally two-celled, variable in size, 28-40 x 20-25 /j, a
32 x 24 u ; pedicel hyaline, persistent, elongated up to 150

and at the same time Cronartium asclepiadeum was noted on Jacksonia
scoparia from the Darling Downs, Queensland.

Then just ten years later, in the transactions of the same society for 1882,
p. 67, R. polita is recorded on the latter plant from Brisbane (Berkeley and
Broome 2), and it was rather peculiar, to say the least of it, that the same

On stems and branches of Muehlenbeckia cunninghami, F.v.M.

New South Wales — Pamamero Lake, Nov., 1860 (Berkeley2).
Victoria — Murray River, Koondrook, Nov., 1905 (C. French, jr.)

This species differs from U. polygoni in the projecting Roestelia-like
aecidia, and in the larger teleutospores with elongated pedicels. Several two-
celled teleutospores occurred, oblong, constricted at septum, upper cell with
apiculate apex, thickened, and more deeply coloured than lower, which is
sometimes colourless, size that of the largest ordinary teleutospores. The
somewhat elongated aecidia, in the absence of any other stage, led Berkeley
to regard this as a species of Roestelia, but the finding of specimens by Mr.
C. French, jr., with teleutospores in addition, showed it to be one of the J
Uromyces. ™

Berkeley 2 first recorded the aecidial stage of this species on Muehlenbeckia
cunninghami in the Journal of the Linnean Society XIII., 174 (1872),

species of Roestelia, or even a Roestelia at all, should occur on plants so
widely separated in a botanical sense as Muehlenbeckia (Polygonaceae) and
Jacksonia (Leguminosae). At the end of his description Berkeley remarks

" The plant is identical with a specimen in the Kew Herbarium, and is
growing on the same plant, Jacksonia scoparia R. Br.

However, on examining the specimens on which this determi nation was
based, it became evident how the error had arisen. Fortunately I have the
specimen of Jacksonia scoparia from the National Herbarium, Melbourne, on

Uromyces — Polygonaceae.

rhich Cronartium was first determined, and Mr, Bailey has sent me a spec
men from his herbarium labelled " Roestelia polita Berk., " and which was s
named by Berkeley, On comparing the Brisbane specimen with the Darlin
Downs specimen, they are seen to be the same, and Berkeley's original deter-
mination of Cronartium is correct. When it is remembered that Roestelia
polita Berk., has only been found on Muehlenbeckia and not on Jacksonia it
will reconcile certain discrepancies in Berkeley's original descriptions and
drawings. In the original description of R. polita the spores are given as

5, while in connexion with its occurrence on Jacksonia it is stated — " the
spores are globose about '0003 inch in diam. (7— 7J /j)" If we turn to the
drawings on PL ] 5, the otherwise puzzling figures become clear when they
are taken to represent a Cronartium, and it will be seen from the description
of this fungus on Jacksonia that the so-called aecidiospores are really the
promycelial spores of that fungus.

If the identical specimen in the Kew Herbarium, to which Berkeley referred,
is examined I have no doubt it will turn out to be a Cronartium. At the
time Berkeley wrote our knowledge of the distinctions between these forms
was not so clear as now, but still the projecting pseudoperidium of the
so-called Roestelia, with its characteristic peridial cells and contained spores,
is quite distinct microscopically from the outwardly similar column of
teleutospores of Cronartium which are wedged together into a solid mass.

In the original description the locality for New South Wales is given as
" Bambamero " Lake, but Dr. Howitt informs me that the name is as
above, and that the lake is situated 20 miles from Menindie, close to the
Darling River.

(Plate XXXIX., Figs. 297, 298; Plate XLIIL, Fig. 317.)

20. Uromyces poiygoni (Pers.) Fckl.

Fuckel, Symb. Myc, p. 64 (1869).

Polygonum

McAlpine, Agr. Gaz., N.S.W., VII., p. 301 (1896).
Sacc. Syll, VII., p. 533 (1888).

0. Spermogonia yellow to honey-coloured, conoid, in small groups.

Spermatia hyaline, minute, 3 X 2 p.

1. Aecidia crowded in roundish patches, bright orange, mostly hypo-

phyllous, but often opposite ; pseudoperidia rather flat, with broad
whitish torn edges.

Aecidiospores subglobose, finely verrucose, pale yellow to orange
yellow, 16-28 fx diam.

II. Uredosori pale cinnamon to rusty brown, scattered, or arranged in

a circinate manner and confluent, mostly amphigenous, pulveru-
lent, round to oval, surrounded or partially covered by ruptured
epidermis.

Uredospores ellipsoid to obovate, yellowish brown, finely
echinulate, with generally two lateral germ-pores on one face, 20-29
x 16-21 fi, average 24 x 20 ^.

III. Teleutosori scattered, blackish, pulvinate, roundish on leaves,

elongated on stems, bursting through browned epidermis, confluent
in masses, surrounding and distorting stem.

IOO

Uromyces — Chenopodiaceae.

Teleutospores globose, subglobose or elliptical, smooth, bright

chestnut brown, apex thickened

/jl), rounded or occasionally

somewhat pointed, 22-34 x 16-20 fx, average 28 x 18 fi ; pedicels
pale yellowish, persistent, firm, long, up to 90 /*.

On leaves and stems of Polygonum aviculare L.

Victoria — Near Melbourne, Rutherglen, Myrniong, Goornong,

Ardmona, Bunyip, Murramurrangbong Ranges, &c. Common

generally throughout the year.
Tasmania — Sandy Bay, January, 1906, II., III. (Rodway).

It is only recorded for Victoria and Tasmania, but is probably to be
found wherever the host plant occurs. Aecidia were first found on young
plants along with the other stages in November, 1904.

The uredospores are plentiful, more particularly in the spring and
summer months, but the teleutospores are formed in the autumn months.

Darluca filum Cast., commonly occurring on uredosori, was found on
teleutosori in May, 1904.

(Plate XVIIL, Figs. 150, 151.)

CHENOPODIACEAE.

21. Uromyces atriplicis McAlp.

A triplex.

Sori epiphyllous, orbicular, scattered, bullate, compact, ruddy brown,

erumpent, girt by the ruptured epidermis, about \ mm. diam.

II Uredospores pale brown, ellipsoid, finely echinulate, with 3-5

scattered germ-pores on one face, 25-29 x 22 ju.

III. Teleutospores intermixed with uredospores, dark brown, sub-globose

to shortly ellipsoid, finely striated longitudinally, slightly thick-

ened at apex

24-27 h

with prominent single

apical germ-pore, 22-29 x

average 26

x 25

On leaves of Atriplex semibaccata R. Br.

Victoria — Royal Park, near Melbourne, April, 1899. (Brittle-
. bank).

The groups of spores shown in the Figures are rather smaller than the

average,

being about 22 x 25 /u
occurred reaching a lenjrth of 37

Occasionally an abnormal teleutospore

(Plate XVIL, Figs. 146, 147.)

22. Uromyces betae (Pers.) Kuehn.

Kuehn, Bot. Zeit., p. 540 (1869)
Cooke, Grev. XT., p. 98 (1883V

Beta. .

Cooke, Handb. Austr. Fung., p. 330 (1892).
Sacc. Syll. VII., p. 536 (1888).

ermogonia yellowish to honey-coloured, in small clusters, on both

faces of leaf

Spermatia hyaline, ellipsoid, 5-6 x 3 u.

I. Aecidia disposed on orbicular or

oblong

surfaces of leaf, mostly about mid-rib and on leaf

yellow spots, on both

pseudo

peridia white, cup-shaped, with reflexed fringed margin ; peridial

Uromyc es — Sapinda c ea

iot

cells firmly united, thickened all round but thicker on one side,
striated, individually somewhat lozenge-shaped, collectively elon-
gated polygonal.

Aecidiospores angular, globose or oblong, orange-yellow,
smooth, 18-28 x 16-22 //.

II. Uredosori cinnamon or chestnut-brown, scattered or circinate,

small, surrounded by the ruptured epidermis.

Uredospores sub-globose, elliptic or ovoid, yellowish-brown,
echinulate, with two distinct equatorial germ-pores on one face,

25-32 x 16-25 /j, average 28 x 21 /j.

III. Teleutosori dark-brown, margined by the ruptured cuticle,

scattered or orbicularly arranged.

Teleutospores ovate or ellipsoid with a colourless papilla at
apex, dark-brown, smooth, 26-35 X 19-25 fj, average 32 x 21 p;
pedicels thin, rather long, soon deciduous.

On leaves of Beta vulgaris L. — Beet and Mangel.

Victoria — This rust generally occurs wherever beet or mangels are

cultivated, and has been found at Port Fairy, Maffra, Murtoa,
Minyip, Ballarat, etc. It also occurs throughout the year
from Jan. and Feb. to Nov. and Dec. As early as 1878 this
rust from Ballarat was determined by Thuemen 2.

S. Australia — Aug., 1897 (Quinn).

Tasmania — (Rodway ]). Devenport, Jan., 1906 (Robinson)

New South Wales — Hawkesbury Agric. College (Musson).

The teleutospores are rather scarce, but they were found at Port Fairy
in August and measured 27-33 x 22-24 /i.

The aecidiospores were found towards the latter end of August and up
to December on young leaves of beet growing from roots left in the ground
during the winter. According to Plowright they very rarely occur in

Britain in a state of nature. They were found to germinate very freely in
water.

(Plate XVII., Figs. 148, 149; Plate XLIIL, Fig. 316; Plate H.)

SAPINDACEAE.

Diploglotlis.

23. Uromyces diploglottidis Cooke and Mass.

Cooke and Massee, Grev. XVII., p. 55 (1889).
Cooke, Handb. Austr. Fung., p 331 (1892).
Sacc. Syll. IX., p. 294 (1891).

III. Sori epiphyllous, scattered, convex, minute, for a long time

covered, at length splitting, pale brown, seated on orbicular
greenish spots.

Teleutospores elliptic, apex obtusely acuminate, base attenuated
into a short pedicel, epispore hyaline, thick, contents granular,

pallid, 50-60 x 20-30 H.

On fading leaves of Diploglottis cunninghamii Hook. f.

Queensland — Woolston Scrub, Brisbane River (Bailey 6> 19).

This species was very destructive to the foliage of the Queensland
Tamarind Tree.

Specimen kindly sent by Mr. Bailey, but spores not obtainable.

102

Uromyces — Caryofhyllaceae.

CARYOPHYLLACEAE

Dianthu

24. Uromyces caryophyllinus (Schrank) Sch

Schroeter, Brand, p. 10 (1869)

A.gr. Gaz. N.S.W. VII., p. 300 (1896)

Sacc. Syll. VII., p. 545 (1888)

II. Uredo

amphigenous, dark brown when exposed, b

ed by epidermis, round or oblon

it long
ed, often confluent.

to 3 mm
U redosp

ndish to oblong, yellowish b

decidedly echinulate,

• »

x 24

m-por

on one

generally 3-4 scatl
face. 21-34 x 17

golden

nearly

average

III. Teleutosori amphigenous, blackish-brown, oblong, confluent in elon-
gated lines, long covered by ashy epidermis, then margined by the

ed and rasced

Teleutosp
oval or ova

s at first inte
cinnamon bro

mixed with uredospores, roundish,
ra, generally thickened, with broad

colourless papilla, average
pedicels very deciduous, hyaline, up

28 x 20 ix or 22-32 x 17-23 hi

On stems

d both surfaces of

of Dianthus caryophyllus L. and

D. chinensis L. — Carnation and Dianthus

Victoria — Near Melbourne, and at Ardmona. Very common in

year round

New South Wa
South Australia
Queensland — Roma, 1901 (Try

— Ashfield
Feb.. 1902

Sydney. (Maiden.)

The

difference bet

uredo and teleuto sori

the naked-eye char

spores the nisei

the latter are a little darker

are

quite d

for the uredosp

are

but
>red

decided spines, while the teleutosp

relatively smooth and

pilla at ape

The teleutosp

filled with finely

the

of which is a large vacuole-like body. When

dry condition, the surface appears to be covered with numerous very fine

protoplasm
spore is ex

the centre

the

points, which Fischer5 regards

ged spines, which project at the

with loosely arrai

optical section, while that of the

points, which are not visible at the marg

The wall of the uredospore is beset

margin when seen in

tospore has densely

ded fine

The

pro

generally described as smooth

L arluca filum Cast, is often parasitic on the uredosori and

(Plate XVIIL, Figs. 152-154; Plate G., Figs. 30, 31.)

23. Uromyces scleranthi Rostr.

Rostrup, Bot. Tidsskr., p. 40 (1897)
Sacc. Syll. XIV., p. 275 (1899).

II. Uredosori

pale

epidermis.

Scleranthus

scattered, minute, orbicular or oblong,
brown, erumpent, and surrounded by the

slightly raised,

ruptured

Uromyces — Zygophyllaceae. 103

Uredospores golden brown, finely echinulate, ellipsoid to ovoid or
oblong, 3-4 scattered germ-pores on one face, epispore 3 a thick,
25-32 x 17-21 11.

On Scleranthus diander R. Br.

Victoria — June, 1898 (Reader).

Only the uredo-stage was found although numerous sori were examined
and while the uredospores generally agree with those of the above species
they are sometimes a little longer.

It was originally found in Denmark on the stems and calyces as well as
the leaves of Scleranthus perennis L.

The description is as follows : — " Sori scattered, minute, orbicular or
oblong, pale brown, long covered by epidermis. Uredospores yellowish-
brown, globose, echinulate, 15-22 /x diam. or oblong, 24-25 x 18-20 fi

Teleutospores rare, intermixed with uredospores, pear-shaped or obliquely
ellipsoid, reddish-brown, apical papilla transversly thickened, 23-24 x
19-24 \x ; pedicel hyaline, deciduous."

(Plate XXVI1L, Fig. 248.)

ZYGOPHYLLACEAE.

26. Uromyces vesiculosus Wint.

Winter, Hedw. p. 22 (1885).

Zygophyttum

Cooke, Handb. Austr. Fung., p. 330 (1892).
Sacc. Syll. VII., p. 547 (1888).

Sori amphigenous, scattered or gregarious, often confluent, rounded

or irregular, ruddy brown, covered by the cinereous vesicular
epidermis, which ultimately ruptures and surrounds the sorus.

II. Uredospores sub-globose, elliptic or ovate, yellow to brownish,

densely warted, even prickly, epispore thin, with 3-5 equatorial
germ-pores on one face, 26-35 x 19-24 /u, average 28 x 20 u.

III. Teleutospores globose, elliptic, ovate or pear-shaped, thickened at

the apex, rounded or with a broad apiculus more or less shortly

21 /i diam., or 23-31 x 17-22 yu, average 25 X 20 a

epispore thick, smooth, dark bay brown when mature; pedicel
long, thick, persistent, may be flexuous, pale olivaceous, up to

On living leaves and stems of Zygophyllum billardieri DC.

S. Australia — Spencer's Gulf (Tepper). (Winter

On leaves, stems, and fruit capsules of Z. glaucescens F.v.M.

Victoria — Near Dimboola, November, 1894, and Warracknabeal,

March, 1904 (Reader).
S. Australia — Yorke's Peninsula, June, 1902 (Molineux).

The confluent sori may form large conspicuous elongated patches up to
10 mm. long or more, and the greyish bladder-like epidermis remains
unbroken for some time. Several two-celled teleutospores were met with
and not constricted at the septum, which was decidedly thick and rather
below the middle, sometimes approaching the base. They were, in other
respects similar to the ordinary teleutospores, even to the size, and
measured about 27 x 22 /u.

io4

Uromycladium.

Dietel observes that the occurrence of b

tosp

is rare

among Uromyces, for only once has he observed it in U. pisi and U.

d two were found in U

Cooke has also found

U. trifolii, and I hav
culosu8 and U.

now to add U. orchidearum, U,

fe
U.

w in

Darlttca filum Cast., is generally very plentiful on uredosor

(Plate XV1IL, Fi

155-157 : Plate G., Fig. 33

Tin

s new genus

UROMYCLADIUM McAlp.

of Rusts is noteworthy, not only on account of the
teleutospores forming a cluster at the top of the stalk, and generally accom-
panied by a colorless cyst, but from the large galls produced by some species
1 • ' • '-- JI-JS and ultimately destroy many of our Wattle-trees

vhich seriously disfigure

(Acacias).

The seven species at present known show every gradation from one teleu-

tospore accompanied by a cyst (U. simplex) through two teleutospores with
or without a cyst ( U. maritimum), ( U. bisporum), and reaching to three teleu-
tospores in a head always without a cyst (U. tepperianum) . The presence
of more than one teleutospore at the end of a stalk is a novel feature in
rusts, and the addition of a colorless vesicle or cyst at the base reminds one
of Ravenelia to which this genus is allied. The unicellular teleutospore also

links it on to Uromyces, and I consider that here we have the connecting
link between Uromyces on the one hand and Ravenelia on the other. The
genus Anthomyces with only a single species is composed of a head of three

or more cells united together with sterile cells at the base, and may prove a
bridging species from Uromycladium to Ravenelia. Spermogonia, uredo
and teleuto sori occur, but no aecidia. The two gall-producing species
hitherto found are U. notabile and U. tepperianum, both of which produce
three teleutospores at the top of the stalk, and the former has uredospores
in addition. The only species known outside of Australia is that of U. tep-
perianum (formerly called Uromyces tepperianus, Sacc), and it is noted for
causing deformation of the shoots, but with us it is ruining whole plantations
of Wattles. Near Altona Bay, Victoria, the branches of the beautiful
Golden Wattle (Acacia pycnantha) are deformed and destroyed by it when
it forms numerous galls sometimes as large as a potato. In the neighbour-
hood of Melbourne, too, hedges of the Kangaroo thorn (A. armata) are

Some of the

being gradually and completely destroyed by this fungus.
shrubs have most of their branches infested with the chocolate colored
galls, the color being due to the spores, and they may be in the form of a
succession of small ones as large as peas, or large ones the size of walnuts.
The phyllodes as well as the branches in some species are deformed. On
cutting across these galls they are seen to be solid to the core, and
not the product of insects but of the fungus, although in many cases bored
and tunnelled by insects after being produced. When our Acacias are
more closely examined for rusts no doubt the number of species will be con-
siderably increased.

General characters — Spermogonia somewhat hemispherical, produced under

mouth, preceding the formation of

any

the cuticle, without paraphyses at
other spore.

Uredospores solitary at apex of basidia, and generally much larger than
teleutospores with several distinct germ-pores.

Teleutospores in clusters, composed of one spore and cyst or two or more
spores with or without a cyst, depressed globose.

Uromycladium — L eguminosae. 1 05

This genus may be distinguished from Urornyces by the arrangement of
the teleutospores, by their shape, which is not elliptical, but depressed globose,
and by the presence, in most cases, of a colorless vesicle or cyst.

Allied with Urornyces on the one hand and Ravenelia on the other.

Australian species, 7.

Analytical Key to the Species.

I. One teleutospore and vesicle in head.

A. Uredospores fusiform, with hyaline apiculus, and finely

warted.

Ur. simplex.

B. Uredospores sub-elliptical, without hyaline apiculus, thickened

at apex, and coarsely warted.

Ur. robinsoni.

II. Two teleutospores in head.

A. Uredospores unknown.

III. Two teleutospores and vesicle in head.

Ur. bisporum.

A. Uredospores sub-elliptical, warted all over, much thickened

and dentate at apex.

Ur. maritimum.

B. Uredospores sub-clavate, evenly warted all over, and scarcely

thickened at apex.

IV. Three teleutospores in head.

Ur. alpinum.

A. Teleutospores finely warted, warts arranged in lines ; uredo-

spores known.

Ur. notabile

B. Teleutospores with converging striae ; uredospores unknown

Ur. tepperianum

LEGUMINOSAE.

Acacia

27. Uromycladium alpinum McAlp.

O. Spermogonia minute, black, shining, punctiform, crowded,- on both

surfaces of phyllodes on discoloured patches, appearing before sori
and ultimately surrounded or accompanied by them.
Spermatia hyaline, ellipsoid, 3 X 2 fi.

Sori amphigenous, rusty -brown, scattered or in groups, bullate, soon

rupturing epidermis and becoming naked.

II. Uredospores yellowish-brown to golden-brown, shortly or elon-

gated clavate, occasionally oval or oblong, warted equally all
over, scarcely thickened at apex, with 3-5 equatorial germ-pores on
one face, 35-51 x 21-26 //, occasionally reaching a length of 58 \x.

III. Teleutospores at first intermixed with uredospores, two in head

with colorless vesicle, depressed globose to subglobose, yellowish-
brown to dark-brown, smooth, very slightly thickened at apex
with distinct germ-pore, 19-22 X 25-30 /i ; vesicle arising from
stalk immediately beneath septum, globose or slightly ellipsoid
about 25 ju diam.

io6

Uromycladium — Leguminosae.

Mesospores
ellipsoid to

associated

with

uredospores,

not

uncommon

oblong

or obovate, rounded at apex, smooth-walled
and wall of about equal thickness throughout, with colorless
stalk, 15-25 x 10-15 p. They differ from uredospores in being
smooth and much smaller, and from the teleutospore in shape, in
not being thickened at apex and without apical germ-pore.

On phyllodes of Acacia dallachiana F.v.M.

Victoria — Alps, near Bright, Dec, 1904 (C French, jr.).

On phyllodes of Acacia buxifolia A. Cunn

New South Wales

New England

(From type of host in National

Herbarium, Melbourne.)
On leaves and pods of A. dealbata Link.

Victoria — Murramurrangbong Ranges, Jan., 1905 (Robinson).

Tasmania— Risdon, Dec, 1905, and Mt. Wellington, Jan., 1906
(Rod way).

On phyllodes of A. implexa Benth.

Victoria— Myrniong, May, 1905 (Brittlebank).

On phyllodes and pods of Acacia linifolia Willd., in National Herbarium,
Melbourne.

New South Wales— Blue Mts.

Queensland — Rockhampton, Nerbool Creek.

This species very much resembles U. simplex in the appearance of the
sori, but it is allied to U. maritimum in bearing two teleutospores and a
vesicle on one sporophoi'e. It differs from the latter, however, in the
uredospores which are generally clavate and warted equally all over.

The vesicles vary in size, and are sometimes large and swollen when
they are ready to burst. In old material they may have disappeared
altogether. The teleutospores were much more numerous than the
uredospores in December.

(Plate XXIV., Figs. 209-215.)

28. Uromycladium bisporum McAlp

III. Teleutosori on the branchlets forming elongated slightly swollen

chocolate-brown masses

appearing as powdery patches

der surface of the leaflets

Teleutosp

depressed globose, occasionally

two in a head, yellowish-brow

bsrlobose to

basal projection,
mediately beneath, 1

htly thickened

-99 x 22-30 H

y short, colored, stalk
* apex with germ-por

On branches, leaves and pod's of Acacia dealbata Link

Victor
Tasm

Murramurrangbong Ranges, January, 1905 (Robinson).
Risdon, Dec, 1905 (Rodway).

No vesicle occurs below the septum, so that this species is an intermediate
form between U. simplex with a single spore and vesicle and U. maritimum

th two spores and a vesicle in each head

The occasional presence of

teleutospores in U. simplex is a further indication of the passage from

the

(Plate XXIV., Figs. 207, 208 ; Plate XXXIII

Uromycladium — Legnminosae. 107

9. Uromycladium maritimum McAlp

O. Spermogonia at first ruddy-brown, ultimately black, dotted over the

surface of prominent discoid tubercles, formed at corresponding
points on both surfaces of the phyllodes, somewhat hemispherical,
but broader than deep, and produced beneath the cuticle,
averaging 120 jx diam.

Spermatia Iryaline, shortly ellipsoid, 3 x 2 ji.

II., III. Sori dark-brown, elongated, compact, confluent, rupturing

epidermis, 2-3 mm. long, partially surrounding the black discoid
spermogonial tubercles.

II. Uredospores oval to elliptical or elongated elliptical, pale-brown to

dark-brown, warted all over and thickened at apex, where spikes
are specially prominent, forming a tuft, with very distinct
equatorial pores, 3-7 on one face, very variable in length and
breadth, 45-60 X 24-28 \i ; pedicel deciduous, hyaline, elongated,
up to 106 x 5 ju.

III. Teleutospores at first intermixed with uredospores, two on each

sporophore, with a lower and lateral colorless vesicle, very rarely
three spores in cluster without a vesicle, subglobose to de-
pressed globose, dark-brown, thick walled, smooth, with finely
granular contents, slightly thicker at apex, with very prominent
apical germ-pore, 30-32 \x diam., or 22-25 X 24-30 \x\ colorless
vesicle globose, with very thin wall, and arising from stalk
immediately beneath septum, 30-35 \x diam.

X. Mesospores intermixed with the uredospores, or even with uredospores

and teleutospores, unicellular or very rarely bicellular, pale,
smooth, with wall of equal thickness and finely granular contents,
without any visible germ-pores, fusiform or ellipsoid, or even
occasionally clavate, solitary at the end of a stalk like the
uredospores, 22-45 X 11-19 /u, occasionally up to 57 \x long.
On phyllodes and stems of Acacia longi folia Willd.

Victoria — Sandringham and Beaumaris.
Tasmania — Mersey Bluff, Jan., 1906 (Robinson).

II. April to August, occurring alone, or sparingly mixed with teleuto-
spores. III. September to latter part of spring and during summer,
intermixed with a few uredospores.

In one form or another it occurs all the year round.

This rust was first found on the coast at Beaumaris in 1895 on Acacia
longifolia, although not investigated at the time, and it was in this species
that the peculiar grouping of the teleutospores and the presence of a
colourless vesicle or cyst was first observed.

The teleutospores germinate freely in water or moist air and without a
period of rest. They germinate all round the sorus on the surface of the
phyllodes, forming a flaky mass of spores and sporidiola, which easily peels off.

The uredospores were only found to germinate in water in the spring.

The spermogonia are associated with uredo and teleutospores occurring
in the same sorus.

Only in rare instances were sori found without being accompanied by

spermogonial tubercles.

(Plates XX., XXL, Figs. 166-184; XXIV., Figs. 216, 217; XXXII

Fig. 273; XLIIL, Figs. 313, 314, 315.)

•>

io8

Uromycladhim — Leguminosae.

Acacia.

30. Uromycladium notabile (Ludw.) McAlp.

Ludwig, Bot. Centrbl. XLIII., p. 6 (1890).
Cooke, Handb. Austr. Fung., p. 343 (1892)

O. Sp

Sacc. Syll. XI., p. 222 (1895).
Uredo notabilis Ludw.
lOgonia minute, punctiform, black

swollen tubercles often

intermixed with uredospores and teleutosp

Spermatia hyaline, ovate or ellipsoid, on elongated basidia, 4 x

2-3 fx.
II. Uredos

both

surfaces of phyllodes, on branches and pod
ochraceous to yellowish-brown, seated on a distorted inflated g

Uredosp

ipsoid to oblon

bright

yellow when fresh

becom
3-5 (

yellowish-br

on elongated hyaline pedicels, with

quatorial germ -pores on one face; epispore thick (3 p)
late, 30-45 x 18-28 jz.

III. Teleutosori on branches, phyllodes and legumes, forming large,

swollen, distorted galls, chocolate-brown, at length very powdery.

Teleutospores intermixed with uredospores at first, in clusters

of three (rarely two or four), sub-globose to depressed globose,

lines,

arranged in

yellowish-brown, densely covered with warts

slightly thickened at apex, with germ-pore beneath, 16-23 x

21-26 /i.

On phyllodes of Acacia notabilis F. v. M.

South Australia

Roseworthy, Sept., 1889, II. (Tepper)

On branches, leaves and pods of Acacia dealbata Link

Yict

Murramurrangb

Jan

905. O.. II.. Ill

(Robinson). Bright, June, 1905, II., III. (Davey). Bairns
dale and Orbost, Dec, 1905, III.
Tasmania— Hobart, May, 1905, II. (Rod way). Cataract Gorgt

and Dulverton, Jan., 1906, III. (Robinson)

On branches of Acacia decurrens Willd

Victoria — Nea

1905. Ill

Melbour

Feb

1905. Ill

Myrniong, July

New South Wales— Exeter, near Moss Vale, May, 1905, III

(Baker).

On branches of Acacia elata A. Cunn

New South

Lawson

0.> III. (Bake

the Blue Mountains, April 1905

On branches of A. binervata DC.

New South Wales —Sydney, Sept., 1905, III. (Maiden.)
On branches of A'.acia pruinosa A. Cunn.

— Gosford, Jan., 1906, II. (Froggatt.)

New South Wales

The ochraceous uredosori are generally distinct from the chocolate-bro

iri, but sometimes the two kinds of spores are found intermixed.

The

.me arrangement oi tne teleutospores is generally the same as in
U. tepperianum, but the dense covering of warty spines instead of distinct

ally the same as in

differentiates them at once and the height of the spore

The

spermogonia are found in association with both uredospores and teleuto

spores

The uredospores closely resemble in size and shape those of Uromu

phijllodiorum, but the surface markings serve to distinguish them

In the

Vromycladium — Leguminosae. 109

latter the warts are arranged in longitudinal lines, while in the former the
surface markings are net-like. See Plate XXIII., Figs. 201, 202, and
Plate XXV., Fig. 226.

Some very large galls were found either surrounding or terminating
branches of the Black Wattle (A. decurrens). A size of 3-4 inches in diam.
was not uncommon, and one large clump resembling a big artichoke measured
5 x 14 inches, and weighed 15 ounces. In some cases the branches still
Nourished beyond the gall, but it was evidently an expiring effort of the tree to
put forth leaves. The particular tree on which the largest galls occurred was
about 30 feet high and 13 years old, but many of the branches were decay-
ing, and it looked altogether rather dilapidated on account of the numerous
galls which were often tunnelled by insects.

The mycelium was evidently perennial, as some of these galls were

several years old.

Only the uredospores of this species were found at first, and were

described as Uredo notabilis by Ludwig. Dietel5 has thrown out the suggestion

in his paper on "The Genus Ravenelia," that from the nature of the

uredospores in Ludwig's species they may be found to belong to Ravenelia,

and, considering the relationship of this genus with Vromycladium, the

suggestion turns out to be not far from the truth.

(Plate XXIII., Figs. 196-205 ; Plate XXXVI.)

Acacia

31. Uromycladium robinsoni McAlp.

■

O. Spermogonia on discoid tubercles, minute, punctiform, ruddy -brown,

partially or entirely surrounded by uredosori or teleutosori.

Spermatia hyaline, minute, sub-globose, about 3-4 ll diam.

II., III. Sori amphigenous, numerous, crowded, up to f mm. diam.,

light rust color, soon erumpent, powdery, and often surrounding
spermogonial tubercles.

II. Uredospores pale yellowish, oval to ellipsoid or elongated elliptical,

thickened at apex (up to 6 /.i), warted all over, particularly at
apex, with 2-3 equatorial germ -pores on one face, 38-45 x
19-22 Li.

III. Teleutospores at first intermixed with uredospores, solitary at end

of sporophore, with lateral vesicle beneath, golden yellow to golden
brown, depressed globose to sub-globose, smooth, slightly thickened
at apex, 19-26 x 25-34 li ; vesicle hyaline, globose to shortly
ellipsoid, often on distinct stalk with septum at base, 20-25 li diam.
or 22-27 x 16-23 ll.

X. Mesospores not uncommon, ellipsoid to elongated ellipsoid or

elongated oblong, with thin and smooth walls, rounded at apex,
without germ -pores, 18-22 x 9—12 yu.

On phyllodes of Acacia melanoxylon It. Br.

Victoria — Murramurrangbong Ranges, Nov. 1902, Dec. 1903,

Jan. 1905 (Robinson). Myrniong, May, 1905 (Brittlebank).
Tasmania — Hobart, Dec, 1905 (Rodway).

This species was found near Kergunyah by my assistant, Mr. G. H.

Robinson, in whose honor it is named. The teleutospores give a d
appearance to the phyllodes on both surfaces, and sometimes the

•

ded as to convey the impression of a continuous mass of rust, or even

! j 0 Uromycladium — L eguminosae.

red dust. It resembles U. simplex in having a single teleutospore at the
apex of the sporophore, with a lateral vesicle immediately below, but it
differs chiefly in the shape and size of the uredospores. The powdery
masses of teleutospores soon spread over the leaf and germinate in situ,
forming flakes which are easily detached.

The spermogonia are seated on tubercles along with uredo and teleuto-
spores, but there may be powdery patches of both kinds of spores even on
the same phyllode, without spermogonia or the associated tubercles (Plate
XXXII., Fi<*. 274). This species represents one of the simplest forms of
the genus, in which the sporophore bears a single teleutospore with a lateral
colorless vesicle immediately beneath it.

(Plate XXII., Figs. 185-189 ; Plate XXXII., Fig. 274.)

32. Uromycladium simplex McAlp

O. Spermogonia ruddy-brown, crowded, minute, punctiform, arranged

in a circinate manner on both surfaces of phyllodes and very
occasionally on young branches.

Spermatia hyaline, shortly ellipsoid, 4-5 x 3-4 fi.

Sori amphigenous, ruddy brown to dark brown, numerous, sometimes

arranged in groups, bullate, soon rupturing epidermis and be-
coming naked.

II. Uredospores yellowish brown to golden brown, fusiform or oval,

with hyaline apiculus, finely warted, with as many as six equatorial
germ-pores on one face, three being very common, 48-58 x 21-25 fi

III. Teleutospores at first intermixed with uredospores, solitary at end

of sporophore and lateral vesicle immediately beneath, yellowish
brown, depressed globose to sub-globose, smooth, slightly thickened
at apex and germ-pore directly beneath, 22-25 X 25-32 ^u ; sporo-
phore fitting into a sort of socket on base of spore, hyaline, elon-
gated, 80 f.i or longer; vesicle hyaline, globose 19-22 fx diam.

On phyllodes and branches of Acacia pycnantha Benth.

Victoria— Grampians, Nov., 1900 (C. French, jun.), Dec, 1900

(Robinson). Little River, Nov., 1902 and Jan., 1905

(C. French, jun.), Feb., 1905. Werribee Gorge, Dec, 1902,

and Nov., 1904. Malvern Gardens, near Melbourne, Sept.,
1905.

Very occasionally the colorless vesicle is replaced by an ordinary spore,
thus showing that the sporophore bears at its apex two spores, or a spore and
its substitute.

The sori form numerous hemispherical pustules which may run together,
and during the latter part of spring and early summer, while the teleuto-
spores are being produced, the exuded spores are observed imbedded in gum
freely germinating and readily detachable in flakes.

The uredospores somewhat resemble those of Uromyces fusisporus, but in
the latter there are only 3-4 germ-pores on one face.

The spermogonia were first found in September on a young tree about
four years old, and they usually occurred on distinct ruddy spots produced
by Coniothyrium pycnanthae McAlp. and other fungi.

Darlucafilum Cast, not uncommon on sori containing both uredo and
teleutospores.

(Plate XIX., Figs. 161-165; Plate XXXIL, Fig. 275.)

Uromycladium — Leguminosae. in

33. Uromycladium tepperianum (Sacc.) McAlp.

Saccardo, Hedw. XXVIIL, p. 126 (1889).

Acacia.

Cooke, Handb. Austr. Fung., p. 331 (1892
Sacc. Syll. IX., p. 291 (1891).
Uromyces tepperianus Sacc.

O. Spermogonia minute, ruddy at first, then black, brownish by trans-
mitted light, depressed globose, 150 fx diam.
Sperrnatia hyaline, ellipsoid, 3-3| x 2-2J li.

III. Teleutosori on the leaves or phyllodes forming swollen distorted

gall-like masses along their whole length, and on the branches

long and broadly effused, or large somewhat spherical galls coated
with cinnamon to chocolate- brown powdery spores.

Teleutospores in clusters of three, sphaeroid to depressed
globose, cinnamon brown, thickly channelled and striate, striae
converging towards apex, slightly thickened in upper portion of
wall, 14-17 X 18-25 fx ; sporophore hyaline, elongated, soon
deciduous.
On branches of Acacia salicina Lindl. ; A. hakeoides A. Cunn. ; A.
myrtifolia Willd.; and A. spinescens Benth.

S. Australia — Blackhills, Sandy Creek, Murray Bridge, etc.,

Dec, 1889 and 1892 (Tepper). Dec, 1901 (Molineux).

On phyllodes and branches of A. armata R. Br.; A. implexa Benth.;
A. juniperina Willd. ; A. melanoxylon R. Br.; A. pycnantha Benth.; A.
rigens A. Cunn.; A. siculiformis A. Cunn.; A. vomeriformis A. Cunn.

Victoria — Mallee near Hopetoun, Oct., 1903 (C. French, jun.).

Ringwood, Aug., 1904, (C. French, jun.). Werribee Gorge,
Jan., 1905, (Brittlebank). Oakleigh, Jan., 1905. Little
River, Jan., 1905, (C. French, jun.), and Feb., 1905.
Cheltenham, May, 1905, (Robinson). Myrniong, July, 1905.
Mt. Macedon, 1882, and Murray River, 1874, from types of
A. siculiformis and A. vomeriformis in National Herbarium,
Melbourne. Common around Melbourne.
On A. diffusa Jjindl. ; A. vemiciflua A. Cunn. ; A. verticillata Willd.

Tasmania. — Hobart, March- April, 1905 (Rodway and Lea).

On Acacia longifolia Willd.

New South Wales — Rose Bay, near Sydney, July, 1905 (Froggatt).

On branches and phyllodes of A. erioclada Benth., and A. glaucoptera
Benth.

West Australia — From types of host in National Herbarium,

Melbourne.
On Acacia stricta Willd.

Tasmania — Gordon, D'Entrecasteaux Channel, Nov., 1905 (Rod-
way).

This species was first described and illustrated by Saccardo in 1889,
who considered that the unicellular teleutospores were borne singly on long
stalks, hence he placed it in the genus Uromyces. He failed to observe,
however, that each sporophore bore a cluster of three in a head. It is the
most widespread of all the known forms being found on nineteen different
species of Acacia, and it shares the property with U. notabile of producing
galls. This is particularly noticeable in the Golden WTattle (A. pycnantha),
where the galls are as large as potatoes, and in some of the Wattle
plantations where the trees are cultivated for their bark they hang in

1 12

Pttccinia.

large numbers from the branches like so many fruits, and the trees are

dving or dead. (See Frontispiece.)

The dark chocolate-brown spore-masses are quite powdery, and each spore
nrominent ribs running from base to apex, so that it is easily known

fluted. In fact, the appearance closely resembles the marki
on the e£L's of some butterflies, and no doubt the purpose is the same

then the memb

hich

eady

the spore collap

when dry conditions pr

ly thick, and prever
Occasionally a spor

has been found germinating in situ, and they germinated freely in water

in twenty hours.

What is said to be the same fungus has been found on Albizzia t
Benth., in Java, and it would be interesting to know if it occurred
indigenous species of this genus in New South Wales, Queensland c
Au

1 any

West

On A. implexa at Myrniong there

and I found

at the end of a branch in July somewhat of a leg-of-mutton shape and
weighing about 3 lbs. (PI. XLI.

Witches' brooms of

sizes

also

rred on A. implexa, caused by this rust, and one of the
iired 45 inches in circumference (PL XLIL).

largest

(Plates XXII., Figs. 190-195; XXIIL, Fig. 206; XXXIV.; XXXY

XLI.; XLIL, Fig. 305.)

•y

PUCOINIA Pers.

This genus includes more than half of all the Australian Rusts, and is
important, not only on account of its numbers, but from its appearing on so
many of our cultivated crops. It occurs on all the cereals and many of the
grasses, on celery and chicory, on fruit trees, such as peach and plum, and
many garden favorites are attacked by it, sucli as chrysanthemum and corn-
flower, hollyhock, marigold and daisy ; even a parasite, such as the native
mistletoe, is subject to it. The two-celled teleutospore is easily recognized
and distinguished from the unicellular uredospore. All the spore-forms may
be present on the one plant, as in P. hederaceae on the native violets (Viola
hederacea and V. betonicifolia) , or reduced to the teleutospore alone, as in
P. malvacearum. There may also be heteroecious forms, such as P. caricis,
with the aecidial stage on the nettle (Urtica). The teleutospore, although
normally two celled, and with a horizontal septum, is sometimes very vari-
able in these respects. It may not only, occasionally, be one-celled, but in
P. dichondrae, for instance, it may be 3-4 celled, and it may be vertically,
obliquely, or even muriformly divided by the septa. The most celebrated of
all the species is P. graminis, or wheat rust, which seems to have lost the
power here of infecting the barberry, for although germinating promycelial
spores have been used upon specially imported barberries and rusty wheat
grown around the latter, still no aecidia have been produced.

Mesospores are common and paraphyses may be present in both the uredo
and teleuto-layer.

General characters. — Spermogonia when present, mostly epiphyllous,
minute, sub-globose or flask-shaped, honey-coloured.

Spermatia very minute, globose or ellipsoid, hyaline.

Aecidia when present at first globose and closed, then cup-shaped and

open, or elongated and cylindrical, with margins generally everted.

4*1* • • j • • • i i -

Aecidiospores originating in serial order and soon free, globose,
globose or angular, hyaline, yellowish or orange.

sub-

Puccinia — Gramineae*

Uredosori when present, generally minute and flattened, sometimes
paraphysate.

Uredospores globose, sub-globose, ellipsoid or ovate, originating singly
on the terminal ends of the hyphae, with germ-pores mostly in pairs or
several, rarely one, never smooth.

Teleutosori variable in size, flattened or pulvinate, sometimes para-
physate.

Teleutospores separate, variously shaped, pedicellate, 1-septate, with
one germ- pore in each cell.

Sporidiola ovoid or reniform, generally hyaline.
Australian species, 90.

GRAM1NEAE.

■

34. Puccinia agropyri Ell. & Ev.

Agropyron, Clematis.

Ellis and Everhart, Journ. Myc. VII., p. 131 (1892).
Sydow, Mon. Ured. L, p. 823 (1904).
Sacc. Syll. XI., p. 201 (1895). .

Aecidium clematidis DC. Fl. franc. II., p. 243 (1805).

0. Spermogonia amphigenous, honey-coloured, in clusters on leaves,

accompanying aecidia.

Spermatia hyaline, globose, minute, 3 li diam.

1. Aecidia hypophyllous, on definite spots, bright orange, in irregular

clusters ; pseudoperidia cup-shaped, flattened, with reflexed lobed
margin ; peridial cells piriform to quadrate, striated at margin
and punctate all over, 28-32 x 16-22 li

Aecidiospores orange-yellow, ellipsoid to sub-globose, finely
echinulate, 21-29 x 16-19^, or 20-22 li diam.

II. Uredosori epiphyllous, rarely on under surface, minute, elliptic or

linear, sometimes confluent, yellowish to orange.

Uredospores orange yellow, elliptical to ovoid, finely echinulate,
4-5 scattered germ-pores on one face, 25-32 x 19-22 li,
epispore up to 2 \ li thick.

III. Teleutosori hypophyllous, minute, covered by the leaden-coloured

epidermis, at length occasionally erumpent, oblong or linear,
sometimes forming lines on sheaths.

Teleutospores yellowish brown, cylindric clavate to elongated
oblong, smooth, slightly constricted at septum, variable in size,
• 40-70 x 12-25 /j, average 60 x 18 li ; upper cell rounded or
squarely truncate at apex and decidedly thickened (6-9 li), some-
times broader than long, darker in colour than lower, 19-29 x
16-25 fi ; lower cell tapering towards pedicel or oblong, usually
longer and narrower than upper, 25-41 x 12-20 li ; pedicel short,
generally tinted. Occasionally three-celled teleutospores occur.
X. Mesospores occasional, pale brown, elongated, rounded, or slightly

pointed and thickened at apex, slightly tapering towards base,
40-48 x 13-16 li, with short coloured pedicels.
Aecidiospores on Clematis aristata R. Br.

Victoria. — Murraruurrangbong Ranges, Dec, 1903, and Jan.,

1905 (Robinson).

^^v^^ &

Uredo and teleutospores on Agropyron scabrum Beau v.

Victoria — Near Melbourne, Dec, 1892 (Robinson). Myrniong,

March, Nov., Dec, Jan. Murramurrangbong Ranges,
Dec, 1903 (Robinson).

ii4

Puccinia — Gramineae.

The teleutospores vary considerably in size and

They

are some

Limes elongated cylindric but usually cylindric clavate, the lower cell being
comparatively narrow and tapering slightly towards base, while the upper
cell is expanded and considerably flattened out at apex. The length may

vary
upper

from 40-70 li and the breadth fro

It is noticeable

may

of elongated and

thers of medium-sized spores

In the Agricultural Gazette for New South Wales, Vol. VI., p

1 895, the rust on this native grass was given as
Henn., but on further examination I find that it a
P. aqropyri Ell. and Ev. and is named accordingly

P. d

Eriks. and

pproaches most closely

Specimens of P. agropyrina Eriks., were examined from Eriks. Exs. 419

Ail

evens Beauv., and the uredospores measured 21-24 jx diam

or 24-27 x 16

The teleutosp

were considerably

smaller than our own material, measuring only 33-42 x 1

39 x 15 u.

•g

The following table will show at a glance the different sizes of the spores
different hosts of the two species of rust :

Uredospore .

Teleutospore.

25-32 x 19-22 n 40-70 x 12-25 /*

P. agropyrina Eriks. on Agropyron repens, Beauv. 16-25 fi diam. 36-41 x 13-16 ft

/'. agropyri Ell. & Ev. on A. glaucum, Roem & Schult. 20-25 x 18-22 ft 60-75 x 20-25 /*
P. agropyri Ell & Ev. on A. scabrum, Beauv.

Although the uredospores are described as smooth by Ellis and Everhart
yet in specimens from Sydow's Ured. Exs. 1362, they are seen to be

smooth
is of the P.

decidedly echinulate and it is a question whether such

a thing as a

uredospore exists among Puccinias.

The

species

altogether

dispersa type. Dietel4 infected Clematis vitalba with the germinating teleu-
tospores of this species and produced spermogonia and aecidia (A clematidis
DC.) so that it is considered to be a heteroecious species and called by

It is suggestive

Klebahn,1 p. 292, Puccinia (clematidi) agropyri Ell. and Ev.

that both stages have been found in the Murramurrangbong Ranges growing
near to each other.

(Plate III., Fig. 25.)

35. Puccinia agrostidis Plow.

Deyeuxia.

Plowright, Grev. XXI., p. 110 (1893) and Gard. Chron., p.
139(1890).

McAlpine, Agr. Gaz. N.S.W. VII., p. 149 (1896).
Sydow, Mon. Ured. L, p. 717 (1903).
Sacc. Syll. XL, p. 202 (1895).

II. Uredosori elliptical to elongated, soon naked, linear and confluent.

Uredospores orange yellow, elliptical, finely echinulate, as many as
9 germ-pores seen on one face, forming a circle inside epispore,

22-24 x 17-18

III. Teleutosori

minute, black to dark-brown, long covered by the

epidermis, sometimes elongate, sometimes in groups.
Teleutospores dark-brown,

smooth, oblong or subclavate,

apex

thickened (up to 5 /;), truncate or rounded, markedly constricted,
attenuated below, almost sessile, very shortly stalked, 40-54 X

14-21 ^

average 44

x 18 /i

•

Puccinia — Gr amine ae.

X. Mesospores fairly numerous, similarly coloured to teleutosp

2-1

elongated elliptical, thickened at apex, 24-30 x

H-

On Deyeuxia forsteri Kunth. = Agrostis solandri, F.v.M.

Victoria — Near Melbourne, 1892 (Robinson). Ardmona, Oct.-

Dec, 1894 (Robinson). Arthur's Creek, Aug., 1902
(Robinson).

New South Wales— (Cobb 2).

Through the kindness of Dr. Plowright, I received some of the orierinal

material, and there is a general agreement in the spores.

Dr. Cobb has described and drawn an unnamed species of Puccinia on
Deyeuxia forsteri, which undoubtedly belongs to the same species. The
teleutospores are given as 44-58 x 16-22 /j,, but the uredospores are rather
larger, being 25-30 X 21-25 /x.

The life-history of this species was made out by Dr. Plowright, who found
after several experimental cultures, that the teleutospores produced Aecidium
aquilegiae Pers. on Aquilegia vulgaris, and that the spores of A. aquilegiae
when applied to Agrostis alba and Poa pratensis produced the rust.

(Plate III., Fig. 27.)

Anthoxanthum

36. Puccinia anthoxanthi Fckl

Fuckel, Symb. Myc. II., p. 15 (1873).
McAlpine, Agr. Gaz. N.S.W. VII., p. 301 (1896).
Sydow, Mon. Ured. I., p. 727 (1903).
Sacc. Syll. VII., p. 665 (1888).

II. Uredosori on both surfaces of leaves, solitary or in elongated

groups, elliptic or linear, confluent in lines, soon naked, pulver-
ulent, dusky orange.

Uredospores yellowish-orange, elliptic to obovate, finely echinu-
late, with two to four very distinct equatorial germ-pores on one
face, 25-32 x 15-20 p.

III. Teleutosori scattered, minute, dark-brown to black, elliptic, naked,

surrounded by ruptured epidermis.

Teleutospores at first intermixed with uredospores, chestnut-
brown, elliptic to obovate or oblong-clavate, smooth, slightly
constricted, rounded and thickened (6 fj) at apex, occasionally
tricellular, 28-48 x 15—21 fi, average 35 x 20; upper cell
usually darker than lower; pedicel persistent, tinted,
20-25 x 6-7 /j,, sometimes reaching a breadth of 10 fi.

X. Mesospores occasional, similarly coloured to teleutospores, obovate,

thickened at apex, average 35 X 17 fx.

On sheath, flag and inflorescence of Anthoxanthum odoratum L.

Victoria — Near Melbourne, Dec, 1896, II. Rutherglen, Dec,

1903, II., III. Leongatha, Feb., 1904., II.

The teleutospores were not very common. In my own garden, near
Melbourne, the uredo-stage was plentiful, but no teleutospores were found,
while at Rutherglen the teleuto- stage occurred on several specimens.

(Plate III., Figs. 20, 21.)

Puccinia — Gramineae .

Beckmannia

7. Puccinia beckmanniae McAlp.

II. Uredosori on both surfaces of leaf, but most common on under,

orange, elliptic, often confluent in lines, erumpent and surrounded
by ruptured epidermis.

Uredospores orange, elliptic, echinulate, with 3 to 4 equatorial

germ pores on one face, 25-29 x 16-19 /.i

long covered by

III. Teleutosori minute, black, linear, hypophyllous,

epidermis, ultimately naked, about \ mm. long.

Teleutospores yellowish-brown, elongated clavate, not or

only

slightly constricted at septum, upper cell generally darker than
lower and with numerous (up to

45-60 x 16-25 p,

absent.

finger

processes,

average

56 x 18 ; pedicel very short or

X. Mesospores brown, oblong to elongated elliptical, thickened at apex,

and either bare or with short stumpy processes, 32-35 x 13-16 //.

On Beckmannia erucaeformis Host.

Victoria — Leongatha, February, March, 1904, II., III.

Only uredospores were met with in February, but by March the teleu-
tospores had developed. There is a general agreement with P. lolii, but the
upper cell of teleutospore is generally broader.

from seed sent by
the

must have been imported with the seed. Holway informs me that he has
collected it in the State of Minnesota, and kindly forwarded me a specimen.

grown

The grass on which this rust was found was
United States Department of Agriculture for trial, and the spore

with

Darlucafilum Cast., was very plentiful, both by itself and

Probably in the former

parasitic

the hidden

mycelium, and may have largely prevented spore formation

(Plate II., Fig. 12.)

38. Puccinia bromina Eriks.

Bromus.

Eriksson, Ann. Sci. Nat. IX., p. 271 (1899).
Sydow, Mon. Ured. L, p. 712 (1903).
Sacc. Syll. XVIL, p. 382 (1905).

Puccinia dispersa, f. sp. bromi, Eriks.

II. Uredosori 1-1 0mm. long, 1

broad, ferr

chiefly
panicle

on upper surface

leaf blade

ded, sometimes on sheath and

Uredosp

bright orange,

with numerous scattered

germ-pores, as many as 11 on one face, echinulate, 27-31 x

8-24

III. Tele

ated elliptical to oblong, black to dark-brown

_ ^„v„ v,xv,.i6o,^u euipuiciu w oDiong, DiacK to dark- brown,

hypophyllous, scattered or subgregarious, sometimes on sheath and

panicle

ed by epidermis, divided into

each compartment separated by elongated, cylind._
clavate, yellowish-brown, barren filaments or paraphy

partments, and
5, or slightly

Puccinia — Gr amine ae.

117

Teleutospores oblong to clavate, dark chestnut-brown, con-
stricted at septum, rounded or truncate at apex and slightly
thickened, 40-56 x 18-24 li, average 48 x 21 li ; lower cell
generally narrower than and almost as dark as upper ; pedicel
short, hyaline to pale yellowish, up to 36 11 long.

X. Mesospores comparatively rare, dark chestnut-brown like teleuto-
spores, clavate, oval or oblong, rounded or truncate and slightly
< . thickened at apex, base sometimes attenuated, 28-36 x 16-21 u.

On Bromus mollis L.

Victoria — Nalinga, Nov., 1898 (Robinson). Kergunyah, Nov.,

1902, Dec, 1903, and Jan., 1905 (Robinson). Domain,
Melbourne, Dec, 1904, II., III., the latter very plentiful.
Nagambie, Nov., 1904.

New South Wales- 1890 (Cobb2).

On Bromus arenarius Labill.

Victoria — Murray River.

The specimen of B. arenarius in the National Herbarium attacked by an
Ustilago and numbered Berkeley 202, also shows this rust, which, however,

was overlooked. It is interesting as probably indicating that this is a
native rust, although it also occurs on the imported B. mollis.

Three-celled teleutospores are occasionally met with, sometimes owing to
the upper cell being divided vertically, but more frequently there are
two transverse septa instead of one.

Specimens were examined from Eriksson's Fung. Paras. Scand., Exs. 420,
and the uredospores measured 20-25 X 17-20 /u, thus being rather smaller
than the above. The teleutospores were much paler in colour, and measured
36-48 x 15—22 yu, the largest being about the average of the Victorian
specimens.

Fritz-Muller1 observed aecidia on Symphytum officinale L. and Pulmonaria
montana Lej., and by infection with the aecidiospores produced the rust on
Bromus. He therefore named it Puccinia symphyti bromorum.

(Plate III., Fig. 28 ; Plate C, Figs. 11-13.)

Rottboellia

39. Puccinia cacao McAlp

* _

TJredo rottboelliae Dietel, Engler's Bot. Jahrb. XXXII., p. 5

(1902).

Sori on both surfaces of leaves but mostly on under, scattered or in

small groups, minute, elliptical, bullate, long covered by epidermis,
about J mm. long.

II. Uredospores brownish or chocolate brown, elliptic to ovoid or oblong,

with distinct germ -pores, generally three equatorial on one face or
may be scattered, epispore thin, dark-coloured, finely echinulateT
32-40 x 24-32 h average 35 x 28 li.

III. Teleutospores intermixed with uredospores, sparse, somewhat

paler, oblong, smooth, rounded or flattened and unthickened at
apex, slightly constricted at septum, 32-35 x 21-22 p ; lower
cell tapering slightly towards base and generally about equal in
size to upper ; pedicel hyaline, deciduous.

Puccinia — Gramineae.

On leaves and sheaths of Rottboellia comp-essa L

Victoria— Creswick, Jan., 1893, II. (Robinson), WerribeeGo

■w^

Dec., 1902, II. Killara, March, 1903, II., III. (Robinson).
Kergunyah, Dec, 1903, II. (Robinson). Near Melbourne,
Nov. and Jan., 1904. Various other localities.
Queensland— II. (Bailey17).

The Queensland specimen is labelled Puccinia straminis DeBary.

The chocolate-brown colour of the uredospores is very characteristic. The
uredo-stage of this species was first described by Dietel on a specimen from
Japan ; on sending him some of our material he agreed that it was the
same. Sydow in his Monograph (p. 800) has described a Puccinia on
R. arundinacea with teleutospores alone, which however do not agree with
these, being thickened at the apex up to 10 n and 34-56 x 22-27 n in size.

The uredosori frequently look quite black owing to the presence of

Darluca JUum Ca

(Plate XXX., Figs. 259, 260.)

40. Puccinia cynodontis Desm

Cynodo

Desmazieres Exsicc. III., No. 655.

McAlpine, Agr. Gaz. N.S.W. VII., p. 150 (1896).

Sydow, Mon. Ured. I., p. 748 (1903).

Sacc. Syll. VII., p. 661 (1888).

P. altera McAlp., Agr. Gaz. N.S.W. VII., p. 151 (1896).

II. Uredo-sori on both surfaces, minute, ellipsoid to lenticular, scat-
tered or confluent, yellowish-brown to rusty -brown, soon naked,

compact.

Uredospores globose to shortly elliptical, yellowish-brown to

brownish-yellow, delicately verrucosa, 20-25 /i diam., or 20-25 x I

7-2

III. Teleutosori roundish to ellipsoid or oblong, scattered or confluent,

pulvinate, black.

Teleutospores intermixed with uredospores, variously shaped,
ellipsoid or oblong, smooth, chestnut-brown, slightly constricted at
septum, occasionally tricellular, 30-50 x 15-21, average 35 x
21 ^i ; upper cell thickened at apex and generally elongated
pointed, sometimes rounded ; lower cell rounded at base or
attenuated; pedicel firm, pale yellow, persistent, elongated up

to 73

X. Mesospores plentiful, dark-brown, ellipsoid, rounded and thickened

at apex, 2t x 19 /<.

On Cynodon dactyl on Pers.

Victoria — Burnley, Caulfield, Armadale, Pakenham, Killara,

Somerville, etc., Oct. -March.

On some plants the teleutospores were of a very regular ellipsoid shape,
and rounded and thickened at apex, so that it was at first thought there
were two species on the same plant. But on examining a number of
specimens every gradation was found from the teleutospore, rather constant
in size and shape with rounded apex, to those necessarily longer on which
the thickened apex was somewhat conical.

Puccini a — Gramineae

119-.

It was likewise found by Magnus 5 that two kinds of uredosp

occur, the one thin-walled with numerous (up to 9) germ-pores and

echinulate, while the other is thick walled, with few (1-3) germ-pores and

either smooth or only with a few scattered spines. There were transition
forms between the two, and this shows how variable a description might be
according to the nature of the spores regarded as typical.

(Plate III., Fig. 24.)

Festuca

41. Puccinia festucae Plow.

Plowright in Gard. Chron., p. 42 (1890) and Grev. XXI., p.

109 (1893).
Sydow, Mon. Ured. I., p. 752 (1903).
Sacc. Syll. XL, p. 194 (1895).

II. TJredosori mostly on under surface of leaf, but generally present

on upper as well, causing conspicuous yellow spots on upper

surface, minute, oblong to elliptic, scattered or confluent, orange-
yellow.

Uredospores sub-globose to ellipsoid, echinulate, yellowish to
orange-yellow, up to 5 scattered germ-spores on one face, 20-25 x
16-18 /u.

III. Teleutosori mostly on under surface of leaf, minute, scattered or

often in groups and confluent, oblong to linear, brownish-black.

Teleutospores at first intermixed with uredospores, clavate to
cylindrical, brown, smooth, slightly constricted at septum, generally
tapering towards base, 40-60 X 15-20 jjl, often 60 x 16 \x.\ upper
cell more or less truncate and thickened at apex, surmounted
• by a crown of 4-6 obtuse, straight or curved, sometimes bifid
processes ; lower cell generally elongated wedge-shaped ; pedicels
persistent, brown, stout, 15-25 \i long.

X. Mesospores very common, similarly coloured to teleutospores, clavate

to cylindrical to somewhat oblong, thickened at apex and sur-
mounted by processes, 31-46 x 1 1—14 yu.

On Festuca ovina L.

Victoria — Leongatha, July, 1903, II. III.
On F. rigida Kunth.

Victoria — Rutherglen, Nov., 1895, II.

In 1890 Plowright experimentally proved that the aecidium on Honey-
suckle (Lonicera) was genetically connected with this species, and it has
since been repeatedly proved by Fischer and Klebahn.

(Plate II., Fig. 13.)

42. Puccinia flavescentis McAlp

Stip

McAlpine, Proc. Linn. Soc. N.S.W., XXVIIL, p. 558 (1903)
Sacc. Syll. XVIL, p. 380 (1905).

II. TJredosori on upper surface of leaf, minute, linear, often confluent

soon naked, p ulverulent, rusty brown, arranged along furrows of leaf

438.

IZO

Puccinia — Gramineae

Uredospores

globose

golden-brown, with at least 5 germ-pores
21-24 p. diam. or 25-32 x 21-24 p.

to shortly elliptical, finely echinulate,

irregularly distributed,

III. Teleutosori minute, elliptical, numerous, black, often confluent

lengthwise, soon naked.

Teleutospores intermixed with uredospores, dark chestnut brown,
oblong, constricted at septum, with rounded and thickened apex (up

to 9 p), smooth, occasionally tricellular, 33-48 x 18-26 p, average
44 x 24 p ; upper cell generally hemispherical, and about equal
in length to lower ; lower cell generally rounded at base, some-
times narrowed and elongated like upper portion of pedicel ; pedicel
persistent, tinted, elongated, up to 72 /j. long.

X. Mesospores common, similarly coloured to teleutospores, elongated

ellipsoid, rounded or truncate and thickened at apex, smooth,
34-43 x 12-15 p.

On Stipa ftavescens Labill.

Victoria — Near Melbourne, Dec-April, II., III.

On Stipa semibarbata It. Br.

Victoria — Nagambie, Nov., 1904, II., III.

The pulverulent uredosori, and the numerous minute, black teleutosori
are characteristic of this species. The uredospores form a rusty powder over
entire upper surface of leaf. The pedicel of the teleutospore is sometimes
lateral and the septum erect as in Diorchidium. It differs from P. stipae
Arthur, in the uredosori being soon naked and decidedly ruddy brown, not
yellowish, while the uredospores are broader.

specimens of P

2S, on Stipa

(Op.) Hora, taken from Syd. Ured. Exs. N

illata L., the teleutosp

generally bluntly pointed, and the size 48-54 x

are decidedly different

The

In speci

mens of P. stipae Arth., from Arthur and Holway's Ured. Exs. No. 27, on

Stipa spar tea T
rather thicker

tel eu tosp

pointed at the ape

I have submitted specimens to Prof. J. C. Arthur, and he remarks that
it is clearly distinct from his species, although there is very much similarity
between the two, as one might expect, from the hosts being essentially alike.

(Plate II., Fig. 17.)

43. Puccinia graminis Pers.

Gramineae.

Persoon, Disp. Meth., p. 39 (1797).
Cooke, Handb. Austr. Fung., p. 335 (1892)
Sydow, Mon. Ured. I., p. 692 (1903).
Sacc. Syll. VII., p. 622 (1888).

II. Uredosori amphigenous, yellowish-brown, linear, 2-3 mm. or

longer, either scattered or confluent in long streaks, especially on

sheaths, pulverulent, soon naked, surrounded by ruptured epi-
dermis.

Puccini a — Gr amine ae,

121

Uredospores elongated ellipsoid to ovate oblong, brownish yellow,
echinulate, generally with 3-4 equatorial germ-pores on one face,
20-36 X H-18/i.

III. Teleutosori sparingly on leaf blades, more commonly on sheaths

stalks and inflorescence, linear, elongated, pulvinate, often con-
fluent, up to 10 mm. or more, dark brown to dense black, soon
rupturing epidermis which is prominent.

Teleutospores clavate to oblong clavate, chestnut brown, smooth,
somewhat constricted at septum, very rarely three-celled, 35-63
X 14-25 fx$ average 52 X 18ju; upper cell rounded or pointed at
apex, rarely truncate, considerably thickened (up to 12 //), some-
times as broad as long, 21-29 fi long ; lower cell attenuated to-
wards base, equal to or longer than upper, 18-35 p long; pedicel
persistent, elongated, tinted, and sometimes as deeply coloured as
spore, of equal thickness throughout, up to 73 x 8 fj.

X. Mesospores very common, intermixed in the same sorus with uredo

and teleutospores, similarly coloured, dark chestnut or paler, oblong
to elongated ellipsoid, generally slender, rounded, pointed or trun-
cated apex and thickened like teleutospore, smooth, 34-46 x 10-
15 /j, ; pedicel generally short or of moderate length. Occasionally
an obovate form occurs, deeply coloured, rounded and thickened at
apex and much broader than usual, up to 22 \x, with elongated
pedicel .

On Wheat (Triticu?7i vulgar e VilL), Polish Wheat (T. polonicam L.),
Oats {Avena sativa L.), Barley {Hordeum vulgar e L.), Rye {Secale cereale L.),
and various species of the following genera of Grasses: — Agropyrvti,
Alopecurus, Amphibromus, Avena, Beckrnannia, Briza, Bromus, Dactylisy
Deyeuxia, Echinopogon, Elymus9 Festuca, Glyceric^ Ilordeum, Phalaris, Boa.

Common in all the States.

Occasionally I have seen a uredospore with a slight indentation on either
side, just where the circle of germ-pores occurs, but never any indications of
a septum. The range of variation in the size of the teleutospores is great.
It may vary from 35 X 25 fi in the oblong, dark-brown, perfectly mature

spore to 63 ju, long, and sometimes only 14 ft broad in elongated clavate spores
in the same sorus.

The spore itself may be colourless, while the brown colouration is in the
pedicel, and this has previously been observed by Dr. Plowright in Austra-
lian specimens ; or the upper cell only may be pale in colour as shown in
] Plate I., Fig. 8, which is from abnormal material met with on one occasion.

There are no paraphyses, but there are numerous mesospores closely re-
sembling teleutospores, but only one-celled.

Darluca filum Cast., sometimes common on the uredosori, particularly
on Wild Oat (Avena fatua) and Glyceric stricta. It has also been found on
Agropyron scabrum, and Dr. Cobb 5 evidently mistook its true nature, for
he says : — " Among its red-rust spots there are certain black bodies which
may constitute a fourth spore of the rust."

The following is a complete list of all the grasses and cereals on which I
have found Puccinia graminis Pers. This determination of the fungus has
been based on morphological characters alone, for I have not yet succeeded
in infecting the Barberry. Those marked with an * are also recorded by

E 2

122

Puccinia — Gramineae

have been found

other authors, and a double asterisk indicates those

by Eriksson in Europe to produce aecidia on the Barberry when their spores

were sown upon it :

Agropyron divergens Nees
*A. scabrum Beauv.

Alopecurus geniculatus L.

Amphibromus neesii Steud.
*Avena fatua L.

• • •

# • •

• • •

• •

• • •

- • •

sat

• * •

• •

. - •

Beckmannia erucaeformis Host

*Briza minor L.
Bromus racemosus L. ...
**B. secalinus L.

• • •

• •

• - •

• • •

B. sterilis L. ...
Dactylis glomerata L.
Deyeuxia quadriseta Benth
Echinopogon ovatus Beauv,
Elymus striatus Willd.
E. virginicus L.
Festuca bromoides L.

• - •

• •

• • •

• #

• •

• • t

• • •

# •

• •

• • •

G. stricta Hook, f .

• • •

• # •

mur

• •

# • t

H. secalinura Schreb.

**H. vulgare L.
**Phalaris canariensis L.

P. minor Retz.
**Secale cereale L.

Triticum polonicum L.

•*T. vulgare Vill.

• •

• • •

• •

* •

* ♦ •

• • •

• • t

• • •

* •

• • •

• ■ •

• • •

• ■ •

• - •

• •

• ■

• • •

• •

• • •

• * •

• • «

• • •

9 • •

• •

• • •

II. , HI.

II. , HI.
II.

II., III.

II. , III.

II., III.
II., III.

II.

II., III.

II.

II.. III.

II., III.

II.

II., III.

II., III.
II., III.

II., III.
II., III.
II., III.

II. , III.
II., III.

•

While P. graminis has been found on the above grasses, it must not be
-assumed that the one will affect the other if growing alongside of each other.
It used to be considered that the same species would attack indiscriminately
any of its hosts, but it has now been proved that a selection or specialisation
goes on, and that the same morphological species living on different hosts is
not identical in the sense of affecting all alike. By means of infection ex-
periments with uredospores and aecidiospores obtained from definite teleuto-
spores, Eriksson arrived at this result, that P. graminis resolved itself into
a series of specialised forms, each of which was confined to a definite and
circumscribed series of host-plants.

As the result of experiments so far, six biologically distinct forms are
constituted :

1. Secalis — On Rye.

2. Avenae — On Oat.

3. Tritici— On Wheat.

4. Airae — On Aira.

5. Agrostidis — On Agrostis

6. Poae — On Poa.

When time and opportunity offer, I hope to carry out experiments to

determine how far the rusts of the P. graminis type on the
are capable of infecting each other.

various grasses

(Plates I., Figs. 2, 5, 7, 8 ; XIV., Figs. 113-122 ; XL., Fig. 301 j

XLIIL, Fig.' 311 ; Plate A., Figs. 3, 4.)

Puccinia — Gramineae.

123

44. Puccinia impatientis (Schw.) Arthur.

Elymus

Arthur, Bot. Gaz. XXXV., p. 19 (1903).
Sydow, Mon. Ured. L, p. 751 (1903).

Aecidium impatientis Schw.

II. Uredosori epiphyllous and occasionally hypophyllous, scattered or

arranged in lines and confluent, oblong, l-l£ mm. long, pulvinate,
erumpent, powdery, reddish-brown.

Uredospores bright orange, subglobose or ellipsoid, finely
echinulate, up to six scattered germ-pores on one face, 25-33 x
18-23^..

III. Teleutosori hypophyllous, greyish -black, pulvinate, long covered by

epidermis, densely crowded, linear, often confluent and arranged in
lines between the veins, with dark-brown paraphyses in clusters.

Teleutospores brownish, oblong to oblong-clavate, smooth,
slightly constricted at septum, 35-55 X 16-22 ^, average
40 x 16 fi ; upper cell rounded or truncate, and thickened at
apex (3-5 p) ; lower cell generally paler in colour, and rounded or
attenuated at base ; pedicel very short, hyaline or tinted, but
dark-brown at apex.

X. Mesospores not uncommon, pale brown, slightly thickened at apex,

elongated, oblong, or obovoid, 29-35 x 12-17 /x.

On living leaves of Elymus condensatus Presl.

Victoria — Rutherglen, Dec, 1903.

The seed of this grass was obtained from the United States, and the
rust was doubtless imported with it.

Arthur sowed germinating teleutospores from Elymus virginicus L. on
Impatiens aurea Muhl., and in fifteen days aecidia were produced. This
result was confirmed in the succeeding year.

Specimens on E. virginicus, from Iowa, in Sydow's Ured. Exs. 1380, are
labelled Puccinia rubigovera (DC.) Wint., and agree with this species.

There is another Puccinia on the same host plant (P. procera Diet, and
Holw.), and according to the description it is quite distinct. The
uredospores are 32-45 x 30-38 /j. and the teleutospores 45-100 \x long.

P. montanensis Ellis also differs in the teleutospores being stouter and
broader, sometimes reaching a breadth of 26 /*.

(Plate III., Fig. 26.)

Lolium

45. Puccinia lolii Niels.

Nielsen in Ugeskrift for Landmaend. I., p. 549 (1

I SydoM, Mon. Ured. I., p. 704 (1903;

Sacc. Syll. XL, p. 203 (1895)

P. coronifera Klebahn, Zeitschr. f. Pflanzenk. II., p. 340

II Uredosori on upper and under surfaces of leaves, forming blister

like swellings, oval or linear, orange, soon bursting through epider
mis, pulverulent, often confluent, sometimes up to 2 mm. long.

124

Puccini a — Gramineae .

Uredospores orange- yellow, subglobose, ovate or broadly elliptic,

strongly echinulate, with 2-3 equatorial germ-pores on one face,

20-30 x 16-24 \x'y occasionally a few colourless capitate para-

physes intermixed.

III. Teleutosori on both surfaces of leaves, minute, black, linear or

' oblong, often confluent, long covered by epidermis, \-\ mm. long.

Teleutospores yellowish-brown to brown, elongated clavate,

tape

towards base, smooth, constriction very slight or abse

sometimes unicellular, 35-60 x 12-20 h average 53 x 16 p

upper cell brown, truncate, thickened at apex, giving off irregular

blunt, generally curved processes variable in size and shape

avera'^in" 6 to 7 or more ; lower cell generally paler in colour

'©*"©

longer than upper, and attenuated towards base ; pedicel pei
tent, coloured, short, stout, up to 11 /u long by 8 p broad.

X. Mesospores exactly resembling teleutospores only one-celled.
On stem, leaf, and inflorescence of Lolium perenne L. Common.

Victoria— Near Melbourne, Myrniong, Port Fairy, &c, Aug. — May.
South Australia — Mount Gambier, April, 1903, II., III.

The teleutospores may be found as early as September, and through the
summer and autumn.

Tn some districts this rust is very severe and kills the grass down to the
roots, especially if there are early autumn rains and warm weather.

Darlucafilum Cast., on uredosori.

Puccinia lolii avenae.

On Avena saliva L., and Avena fatua L.

Victoria— Port Fairy, Dec. 1903, and Sept.— Nov. 1904, II., III.

Brighton, Jan., 1904, II., III. Myrniong, Nov., 1904, II.
Near Melbourne, Jan., 1905, II., III.

New South Wales— (Cobb 12

Eriksson has shown that the form occurring: on Lolium will not infect

Avena, nor the reverse, so that there are two biologic forms, and that found
on the Oat may be distinguished as P. lolii avenae.

This species may occur alongside of P. graminis, and the teleutosori
were found together on the sheath of the cultivated oat, A. sativa, and!

the wild oat, A. fatua.

The teleutospores on the wild oat are sometimes very much longer than
the normal. They range from 70 to 86 /i in length, and from 18-22 \i in
breadth at the apex, and 7-8 fi at the base.

By infection experiments Klebahn2 proved the connexion between this*
rust, and the aecidium on Phamnus frangula L., but this genus does no
occur naturally in Australia.

P. coronata Corda, was the name originally given to the species occurring
on Lolium ferenne, Avena sativa, &c, with teleutospores surmounted by
crown of finger-like processes. It was found by culture experiments that
the aecidial stage was produced on species of Phamnus, and Klebahn deter
mined that the spores from some species of grasses produced aecidia
exclusively on Phamnus cathartica, while others did so exclusively
P. frangula. This showed a biological distinction between the two, an
Klebahn proposed the name of P. coronifera for the rust producing aecidia

Puccinia — Gramineae. 12

on Rhamnus cathartica, while the original name was retained for the rust
producing its aecidia on Rhamnus frangula. Nielsen had previously used
the name of P. lolii for Klebahn's species, and so it is retained on the score
of priority.

(Plate II., Figs. 11, 14 ; Plate XXX., Fig. 261 ; Plate B., Figs. 5-8.)

46. Puccinia magnusiana Koern.

Phragmites

Koernicke, Hedw. XV., p. 179 (1876).
Sydow, Mon. Ured. I., p. 785 (1904).
Sacc Syll. VII., p. 631 (1880).

II. Uredosori on both surfaces of leaf, but mostly on upper, snuffy

brown, erumpent, surrounded by ruptured epidermis, elliptic or
linear, confluent lengthwise and forming long streaks, with clavate

bright yellow paraphyses,

Uredospores elliptic or obovate, golden yellow, echinulate, with
four equatorial germ-pores, 24-35 X 16-19 /x; intermixed with
large numbers of clavate paraphyses, club thickened at apex, of a
dark smoky -brown and stalk hyaline, commonly 80-90 /x long.

III. Teleutosori minute, black, very numerous, scattered, elliptic or

linear, confluent into long black streaks on both surfaces of leaf,
but mostly on upper, occasionally paraphysate as in uredosori.

Teleutospores clavate to oblong, dark chestnut-brown, generally
rounded and thickened at apex (up to 12 /x), sometimes bluntly
pointed, hardly constricted at septum, 35-55 x 14-21 ^u, average
38 X 18 yu ; lower cell attenuated towards base ; pedicels firm,
persistent, coloured yellowish, about length of spore or longer,
up to 70 jjl.

X. Mesospores similarly coloured to teleutospores, variable in shape,

somewhat ovoid to elongated, thickened at apex, 28-38 x 13-19 p.

On Phragmites communis Trin.

Victoria — Orbost, Aug., 1901 (Pescott). Flinders, Jan., 1902.

Port Fairy, June-Aug., 1902 and 1905. Killara, March,

1903. Bunyip, May, 1904.
South Australia — The Grange, April, 1891 (Tepper). River

Torrens, Adelaide, Apr., 1903 (Tepper).
Tasmania — (Rod way ]).

The clavate paraphyses are usually described as hyaline, but Dr. Dietel
compared the Australian specimens with intense dark-brown paraphyses
with material from Europe, America, and the Cape of Good Hope, and he
found that the latter were sometimes scarcely coloured, at other times light
or dark brown.

The rust on Phragmites communis Trin., was considered as one species up
to 1876, viz., Puccinia phragmitis Schum. Then Koernicke separated it
into two, viz., P. phragmitis and P. magnusiana. The former was
characterized by the absence of paraphyses from the uredosori, the large
bullate teleutosori and the very long stalked markedly constricted teleuto-
spores, while the latter had numerous clavate paraphyses in the uredosori,
minute, punctiform or linear teleutosori, and shortly stalked, hardly
constricted teleutospores. The aecidia in the two cases likewise occurred
on different hosts.

126

Puccinia — Gramineae. ,

Subsequent

ch showed that there were still other species on the

host-plant. Plowright separated a third species in 1888, mainly based
upon the fact that the aeeidiospores were only produced upon Rumex

and not upon other pi

Next, Lud

1892 distinguished another

material sent from South Austr

P. tepp

Finally

Arthur in 1902 published another new species which he named P. sim
because it was very similar to P. magnusiana.

There are thus at least five different species of Puccinia recorded upon
Phragmite8 communis, and it is not always easy to differentiate them, at

by morphological characters

In three of them the

diosp

proved to occur on different host-plants by

dered by many a sufficient specific distinction

of cultures, and this is

On these

ounds they

\\-

w«uld be considered biological species as in the case of Puccinias on various
cereals and grasses, but at the same time an attempt will be made here
show any points in which they differ morphologically. Only two of the five
have hitherto been found in Australia, viz., P. magnusiana and P. tepperiy
for although Dr. Cooke gives P. phragmitis in his Handbook of Australian
Fungi, it was probably intended for P. magnusiana, which has been found

Victoria and South Australia.
The life-history of this fungus was first determined

Dr. Plowright,5

who found that the promycelial spores produced aecidia on Ranunculus,
repens and R. bulbosa, and conversely the aeeidiospores produced uredo-
spores and teleutospores on Phragmites. This was afterwards confirmed by
Fischer' and Klebahn.'

The five species may be arranged according to their aecidial hosts where-
known, as in the following table with their special characteristics shown : —

Rumex sp. and
Rheum.

Rumex acetosa.

Uredosori without paraphyses.

Ranunculus repens
and R. bulbosa.

Anemone dichotoma.

Uredosori with paraphyses.

P. phragmitis.

Uredospores,
25-35 x 15-23/i

Teleutospores,
45-65 x 16-25 /i
markedly con-
stricted

Pedicels,
150-200 n long

P. trailii.

25-35 x 20-25 \x

50-60 x 20-23 \i
markedly con-
stricted

P. tepperi

75-100 fi long

27-30 x 20-23 \i

45-68 x 18-26 fi
markedly con-
stricted

P. majrnusiana.

180-250 fi long

24-35 x 16-19^

35-55 x 14-21 /i
hardly con-
stricted

About length of
spore

P. simillima.

26-44 x 18-22 fi

42-56 x 15-20 fi
hardly con-
stricted

About length of
spore

It will be seen from the above that the

are two typ

clearly

distinguishable morphologically, paraphysate and aparaphysate^ and with
these limits there are only "biological species " based upon a differ
aecidial host.

If we compare P

having a granular spore-membrane, and shorter

th P. phragmitis, the teleutospores differ

ped

similar comparison is made of P. tepperi, the length of the pedicel is the

triking feature, and

perhap

ther premature to

sepa

P. phragmitis until a distinct aecidial host has been proved.
According to Arthur, P. simillima is distinguished from P

it from

the slender pointed

the

more clavate and less

redospores and the probably different aecidial host

dichotoma L

viz.

ghened

Anemone

(Plate II., Fig. 1

Puccinia — Gramineae. 127

47. Puccinia maydis Bereng.

Zea.

Bereng., Atti VI., Riun. sc. ital., Milano, p. 475 (1844).
Cooke, Handb. Austr. Fung., p. 337 (1892).
Sydow, Mon. Ured. I., p. 830 (1904).
Sacc. Syll. VII., p. 659 (1888).

Puccinia sorghi (in part) Schweinitz, N. Amer. Fung., p. 295,

(1831).

II. Uredosori amphigenous, elliptic or oblong, scattered or united

into larger or smaller groups, here and there confluent, rather
convex, soon erumpent from the longitudinal fissure of the
cuticle, reddish-brown.

Uredospores ellipsoid or obovate, slightly warted, at first
yellowish then reddish brown, with two to three equatorial germ-
pores on one face, 23-38 x 20-26 jx.

III. Teleutosori amphigenous, scattered or subgregarious, variable in

form, mostly linear or oblong, long covered by epidermis, pro-
minent, very black, 1-2 mm. long.

Teleutospores obovate to oblong or subclavate, bright chestnut
brown, smooth, constricted at septum, rounded or conoid and
thickened at apex (up to 8 /x), sometimes truncate, roundated,
base and rarely somewhat attenuated, occasionally 3 or 4 celled,
32-52 x 16-24 ^u, average 36 x 20 ji ; upper cell rarely

vertically divided ; epispore thick ; pedicels persistent, elongated,
yellowish, thickened particularly towards apex, up to 90 p. long.

X. Mesospores occasional, similarly coloured to teleutospores, ovoid,

rounded at both ends, thickened at apex, average 35 x 17 /*.

On leaves of Zea mays L.

Victoria.— Seville, April, 1897, II. (Hill). Tally Ho, April, 1902,

II. (Cronin). Leongatha, March, 1904, II. Near Mel-
bourne, April, 1904, II., III.
I New South Wales. — Richmond (Cobb).

Queensland. — Toowoomba, 1887 (Tryon1), (Bailey1,5).

Schweinitz considered that the same rust occurred on sorghum and
maize to which he gave the name of P. sorghi, but since they are now found
I to be distinct Sydow in his Monograph has reserved the above name for
this one.

Arthur* inoculated maize plants with uredospores from an aecidium on
Oxalis cymosa Small, and in five days uredosori appeared in abundance.
The Oxalis is supposed to have been inoculated from the teleutospores of
P. maydis, but this infection has still to be verified.

Darluca Jilum Cast., occurs on the uredosori.

(Plate II., Figs. 15, 16; Plate C, Figs. 14, 15.)

48. Puccinia perplexans Plow.

Alopeciwus .

Plowright, Quart. Journ. Micro. Sc. XXV, p. 164 (1885).
Sydow, Mon. Ured. I., p. 719 (1903).
Sacc. Syll. VII., p. 632 (1888).

II. Uredosori amphigenous and on sheath, orange, elliptic, rupturing

epidermis and often confluent in lines.

128

Puccinia — Gramineae.

Uredospores orange-yellow, subglobose to elliptic, finely echin

late, with 4-6 scattered germ-pores on one face

9 x 19-22 n

II [. Teleut

ed epider

phigenous and on sheath, covered by leaden-

l confluent in very

first minute

the

oblong, constricted

ng lines and even forming patches.

Teleutospores variable in shape, clavate to
ptum, brown, upper cell darker than lower, rounded or truncate

apex and slightly thickened, lower cell attenuated towards base

j 37 x 17 u; pedicel short,

ded. 3

6-24

hyaline, may reach a length of 20

X. Mesospor

thickene
16-21 yu

On Alop

common
at apex.

ilarly coloured to

teleutospores, slightly
to ovate or pear-shaped, 25-32 x

icul

Victoria— Minyip, Nov., 1903 (Eckert)

Among these plants one was badl)
nnia graminis but no trace of the

usted with the uredosp

tosp

of that fu

Puccinia

found, although all the specimens were growing together.

Plowright in a series of more than thirty cultures worked out the life-
history of this fungus and showed the genetic connexion between the
aecidium on Ranunculus acris and this species, which was afterwards con-
firmed by Dietel and Klebahn.

(Plate III., Fig. 23.)

49. Puccinia poarum Niels.

Poa

Nielsen, Bot. Tids. II., p. 26 (1876).
Cooke, Handb. Austr. Fung., p. 336 (1892).
Sydow, Mon. Ured. I., p. 795 (1904).

Sacc. Syll. VII., p. 625 (1888).

IT. Uredosori small, round or elliptical, scattered or aggregated togethe

orange-yellow or foxy- red.

Uredospores shortly elliptical, finely echinulate, orange-yellow,

21-24 x 15-18 fi, seated on hyaline pedicels up to 30 n long, and

intermixed with numerous, stiff, capitate paraphyses, reaching a
length of 68 fi. I

III. Teleutosori small or large, scattered or orbicularly disposed,

dark-brown to black, covered by the epidermis. flj

Teleutospores elliptical or subclavate, very variable, apex

truncate, rounded or conical, thickened

8 fi), dark -brown,

smooth, not or hardly constricted at septum, 35-45 X 15-21 jj

or may reach a length of 53 y. when 3-celled, average 38 x 19 /i ;
pedicels persistent, short, brown.

ives and stems of Poa annua L.

Victoria— Near Melbourne, Jan.-Dec, 1886. &c. II.. III.

South Wales

Sydney, May, 1898

Sydney, 1890 (Cobb). Botanic Gard

On Poa

Victo

Botanic Gardens, Melbourne, Dec, 1904

On Poa vratensis L

Victo

Port Fairy, Sept., 1904. Near Melbourne, Sept

Puccinia — Gramineae

129

Uredospores have been found all the year round and teleutospores from
August to December. Teleutospores are fairly plentiful in October, and
they are often three — or even four — celled. The three-celled forms have
either transverse septa, or the upper cell may be divided vertically or
slightly oblique. The four-celled is either produced by transverse septa,
tlie upper one being usually oblique, or the upper third may be divided
longitudinally. In these abnormal forms the length may reach 53 u and
the breadth 34 li at the apex.

The genetic connexion between the aecidium on Coltsfoot (Tussilago
farfara) and this rust was first shown by Nielsen in 1876 and repeated by
Plo wright in 1882, who found that the aecidiospores produced the uredo-
spores on Poa annua in ten to twelve days. But the Coltsfoot does not
exist in Australia, and, therefore, this rust can reproduce itself without the
intervention of aecidia. As might be anticipated the uredo stage carries
it over the winter, and, indeed, the fungus is most plentiful in our winter and
early spring months — June, July, August, and September. Poa annua
as a rule dies away early in October, as soon as the warm weather comes on,
except in moist shady places. Lagerheim1, has found the spores on the
leaves after the melting of the snow.

Darluca filum Cast., commonly

(Plate III., Fig. 22

50. Puccinia purpurea Cooke

Sorghum

Cooke, Grev. V., p. 15 (1876).
Sydow, Mon. Ured. T., p. 803 (1888).
Sacc. Syll. VII., p. 657 (1888).

Uredo sorghi Fckl.

II. Uredosori amphigenous, seated on indeterminate, elongated and cod-

fluent, bright red to purple spots, scattered or in small irregular
groups, covered by epidermis, and then splitting, yellowish-brown,
1-1 J mm. long.

Uredospores ellipsoid, obovate to piriform, yellowish-brown,
roughly aculeate, with 3-4 germ-pores on one face, 28-34 X
20-25 ix, average 31 x 21 li ; paraphyses intermixed.

[III. Teleutosori on similar spots, mostly hypophyllous, oblong, elliptic,

or linear, always or long covered by epidermis, reddish-brown.

Teleutospores ellipsoid, oblong or ovate-oblong, rounded at apex,
not or scarcely thickened, not or very slightly constricted at
septum, mostly rounded at base, smooth, bright brown, with very
thick epispore, 35-56 X 22-32 li ; pedicel hyaline, persistent,
thick, up to 100 fi long ; paraphyses present.]

On leaves of Johnson Grass (Sorghum halepense Pers.) and Sugar-cane

{S. vulgare Pers.).

h 15

Queensland — Gladfield (Gwyther), Nerang (Shirley), (Bailey

Only uredospores have been met with here.

In P. maydis the teleutospores are thickened at apex. Originally P.
sorghi Schwein. included rust on Zea and Sorghum, but it is now found
that the same rust does not occur on both, and Sydow, in his Monograph,
has wisely determined to reserve P. purpurea Cooke, for Sorghum and
P. maydis Bereng., for Zea.

Darluca filum Cast., is very common on uredosori.

T^o Puccini a — Gramineae.

Hordeum*

51. Puccinia simplex (Koern.) Eriks. and Henn.

Eriksson and Henning, Getreider. p. 238 (1896).
McAlpine, Journ. Dep. Agr. Victoria L, pp. 430 and

529 (1902).
Sydow, Mon. Ured. I., p. 756 (1903).
Sacc. Syll. XVII., p. 377 (1905).

Puccinia straminis Fckl. var. simplex Koern. in Land und
- Forstwirtsch. Zeit. (1865).

II. Uredosori very minute or up to \ mm. long, sparingly scattered or

sometimes numerous on both surfaces of leaves, but mostly on
upper, citron yellow.

Uredospores subglobose or ellipsoid, echinulace, yellow, relatively
thick walled, with numerous scattered germ-pores, 4-7 on one

face, 21-24 ft diam, or 24-30 X 17-20 u.
III. Teleutosori covered by the epidermis, amphigenous

numer

minute, punctiform, mostly oblong and confluent, black, somewhat
longer on leaf sheath than on leaf, divided into compartments by
brown paraphyses, often strongly enlarged and thickened towards,
the apex and spread out horizontally.

Teleutospores oblong clavate to clavate, dark chestnut browm
smooth, usually rounded or truncate at apex, or obliquely conical,,
and slightly thickened at apex, but may sometimes reach 6 /u, and
slightly constricted at septum, 40-54 x 17-24 fi; upper cell
oblong, generally as broad as long, 15-21 X 17-22 \x\ lower cell
tapering towards pedicel, sometimes quite narrow at base, longer
than upper, 21-33 x 15-18 n ; pedicel short, slightly tinted.

X. Mesospores very numerous, similarly coloured and similarly
thickened at apex to teleutospores, asymmetrical, sometimes
elongated, saccate or inversely triangular, very variable, 24-45 x
15-24 fi.

On Hordeum vulgare L.

Victoria — Port Fairy, Warrnambool, Nov.-Jan.

This form was first described in 1865 by Koernicke as a variety, but now
Eriksson and Henning have raised it to the rank of a species.

A very characteristic feature of it is the great predominance of unicellular
spores. A sorus may consist entirely of these or there may be a few normal
teleutospores intermixed- They are produced alongside the two-celled forms,
and are quite similar to them except that they are one-celled, and they
evidently show the transition from the one-celled to the two-celled teleuto-
spore. There is usually a slight thickening at the apex of both, but it rarely
exceeds 4-6 \x. It was early recognised as an exceptional form, and was
named Uromyces hordei by Nielsen in 1875, and Puccinia anomala by Rostrup
in 1876.

The teleutospores only germinate after a period of rest, according to
Eriksson and Henning1.

(Plate I., Figs. 1, 4, 9 ; Plate B., Figs. 9, 10.)

Puccinia — Gramineae. j?j

52. Puccinia subnitens Diet.

Distichlis

Dietel, Erythea, p. 81 (1895)

II. Uredosori hypophyllous, orange, powdery, soon naked and sur-

rounded by ruptured epidermis, often in long lines and confluent,
bullate.

Uredospores yellowish-brown, globose to slightly ellipsoid,
densely echinulate, with 3-4 scattered germ-pores on one face
thick-walled, 21-23 li diam. or 21-28 x 18-20 li. *

III. Teleutosori similar to uredosori, but dark-brown.

Teleutospores intermixed with uredospores, dark chestnut-
brown, oblong to clavate, rounded at both ends or slightly

tapering towards base, Slightly constricted ;it septum and

•

thickened at apex (4-7 ^u), smooth, 28-46 x 19-23 ll ; pedicel
persistent, yellow, up to 56 ll long.

X. Mesospores not uncommon, similarly coloured to teleutospores,

obovate, rounded and thickened at apex (up to 7 li), smooth,
28-37 X 19-22 li ; pedicel similar to that of teleutospore.

Darluca jilum Cast., common on uredosori.

On living leaves of Distichlis maritima Rafin.

Victoria — Flinders, Jan., 1902. Near Melbourne, April and

Nov., 1905, II., III. (Robinson).

This species is quite distinct from P. distichlidis E. and E., which,
however, was originally found on Spartina gracilis Trim, the host-plant
being mistaken for a Distichlis.

Arthur7 has obtained some remarkable results by sowing the teleuto-
spores of this species derived from Distichlis sficata Greene, on various plants.
The germinating teleutospores produced aecidia on three widely-separated
families of plants, viz. — Chenopodiaceae, Capparidaceae, and Cruciferae.

According to the Index Keivensis, D. spicata is a synonym of D. maritima.

(Plate XXX., Figs. 262-264.)

53. Puccinia tepperi Ludw.

Phragmites

Ludwig, Zeitschr. f. Pflanzenk. II., p. 132 (1892).
Sydow, Mon. Ured., I., p. 792(1904).
Sacc. Syll. XL, p. 203 (1895).

Sori epiphyllous, large, scattered or gregarious and confluent, up to

3 cm. long, firm, thick, pulvinate, erumpent, dark-brown ;
paraphyses in clusters, hyaline or pale yellow, capitate, thickened
at apex, up to 75 ll long.

II. Uredospores ellipsoid or pear-shaped, echinulate, pale yellowish,

with as many as 9 scattered germ-pores on one face, 27-30 x
20-23 ll.

III. Teleutospores intermixed with uredospores, subcylindrical to

lly oblong, with rounded or acute slightly thickened

apex (4-5 ^u), markedly constricted at septum and upper cell often

becoming detached, rounded or rarely attenuated at base, smooth
yellowish-brown, occasionally 3-4 celled, 45-68 x 18-26 li

50 x 19 ll; pedicel hyaline, persistent, up to 250

132

Puc cinia — Gr amine ae .

X. Mesospores very rare, similarly coloured to teleutospores, elongated

slightly thickened

ellipsoid, rounded and

slightly towards base, 58 x 21 ii

On Phragmites communis Trin.

at apex, tapering very

S. Australia — The xG range, near Adelaide, April, 1891 (Tepper)

(Ludwi

& )'

The mesospores had all the appearance of teleutospores without the
septum. The rounded apex and stalk at base showed that this was not
the basal cell of a normal teleutospore.

(Plate II., Fig. 19.)

Triticum

54. Puccinia triticina Eriks.

Eriksson, Ann. Sci. Nat. Series VIII., p. 270 (1899).
Svdow, Mon. Ured. I. p. 716 (1903).
Sacc. Syll. XVII., p. 376 (1905).

P. dispersa i. sp. tritici Eriks. and Henn., Zeitschr

Pflanzenk. IV., p. 257 (1894).
II. Uredosori 1-2 mm. long, reddish-brow]

but

ostly

on upper surface of leaf -blade, elliptic, sometimes confluent, scat-
tered or somewhat gregarious, occasionally on sheath and stem.
Uredospores subglobose to shortly ellipsoid, echinulate, orange-

yellow, 4-6 scattered germ-pores on one face, 20-28 x 18

average 48 x 16 fx ; upper cell deeply

III Teleutosori oblong, black to dark-brown, hypophyllous, scattered,

often arranged lengthwise in lines, sometimes on sheath and stem,
covered by epidermis, divided into compartments surrounded by
brown paraphyses.

Teleutospores clavate to oblong, smooth, yellowish-brown to
dark-brown, slightly constricted at septum, very occasionally

3-celled, 39-57 x 15-18 yw,
coloured, generally rounded or flattened at apex and thickened,
17-31 x 15-18 jx ; lower cell paler and attenuated towards pedicel,
longer and narrower than upper, 22-36 X 12-14 li ; pedicel
short, coloured.

X. Mesospores occasional, similarly coloured to teleutospores, ellipsoid

to clavate or sub-clavate, shortly stalked, smooth, slightly thickened
at apex, 25-38 x 13-16 li.

On wheat, Triticum vulgar e Vill. and T. polonicum L. Common.

Victoria, New South. Wales, Queensland, South Australia, West

Australia, and Tasmania.

The uredospores
subglobose instead

distinctly different from those of P. graminis in being

of elongated

d the more numer

scattered instead of forming an equatorial band

germ-por

Uredospores taken from young wheat plants in the winter months (June

August) readily germinated in a moist chamber.

(Plate I., Figs. 3, 6, 10; Plate XL., Fig. 302 ; Plate A., Figs.

Puccinia — Cyperaceae

l33

CYPERACEAE.

55. Puccinia Caricis (Schum.) Rebent.

Rebentisch, Fl. Neom., p. 356 (1804).
Sydow, Mon. Ured. L, p. 648 (1903).
Sacc. Syll. VII., p. 626 (1888).

Carex, Urlica,

Aecidium urticae DC. Fl. franc. II., p. 243 (1805).

O. Spermogonia in small or large groups, honey coloured.

Spermatia hyaline, ellipsoid, 4J x 2jw

I. Aecidia in rows or groups on yellowish or reddish spots, closely

crowded, reaching 1 mm. in diam.; pseudoperidia cup-shaped,
becoming wide at the mouth and campanulate, flattened or
outstanding, with whitish margin which is toothed and reflexed.

Aecidiospores orange, polygonal to subglobose, finely echinulate,
16-26 x 12-20 fi.

II. Uredosori elliptical or oblong, minute, hypophyllous, scattered or

subgregarious, often confluent in long lines, erumpent, girt by the
ruptured epidermis, ferruginous.

Uredospores ellipsoid or ovoid, yellowish brown, echinulate
1-2 germ-pores on one face, 25-30 x 19-24 li.

III. Teleutosori minute, dark-brown or black, round to elliptic,

scattered or gregarious, sometimes confluent, compact.

Teleutospores clavate to elongated oblong, dark-brown, smooth,
constricted at septum, apex much thickened (up to 12 /j) and
rounded or bluntly pointed, lower cell generally attenuated
towards base, 35-64 x 16-22^, average 52 x 22 li ; pedicels
tinted, persistent, generally short, up to 30-40 n long.

X. Mesospores not common, elongated elliptical, coloured like teleu-
tospores, thickened and bluntly pointed at apex, 35-44 x
12-16 fx.

Aecidia on leaves, leaf stalks and stems of Urtica dioica L.

Victoria. — Bacchus Marsh, Myrniong, Fern Tree Gully, etc.
Uredo and teleutospores on leaves of Car ex breviculmis 1Z. Br.

Victoria — Killara, Sept., 1902 (Robinson.)
On Carex pedunculata Muhl.

Tasmania — Cascades, May, 1892, II., III. (Rodway).
On Carex gunniana Boott., and C. inversa, R. Br.

Tasmania — Longley, Dec, 1894, II. (Rodway).
On Carex alsophila F.v.M.

Victoria — Murramurrangbong Ranges, "Dec., 1903, II. (Robinson)
On Carex paniadata L.

Victoria — Gembrook Ranges, March, 1904, II., III. (C. French,

jun.)

Dr. P. Hennings has recorded a new species, Uredo breviculmis, on
Carex breviculmis var. leucochlora Bunge, from Japan, and seeing that the
host plant is a native of Japan as well as of Australia, I forwarded some of
this material to him, and he replied that he scarcely thinks the two species
are identical.

134

Puccini a — Cy-peraceae.

Mag

experimentally proved, in

187

that Aecidium urticae

DC, was genetically connected with Puccinia caricis occuring on Car ex hirta

Dr. Plowright also, in many cultures, has shown that

aecidiospores pro

duce the uredospores on species of Carex, and the teleutospores produce the
aecidiospores on Urtica.

On the under surface of the leaves the aecidia occur in small groups, but
on the leaf -stalks and stems they are in large, bright-orange patches,
thickening and distorting the parts on which they occur.

Magnus1 has also shown that this rust winters on Carex hirta by

of the uredo

(Plate IV., Figs. 29, 30.)

Cyperus.

56. Puccinia cyperi Arthur.

Arthur, Bot. Gaz., p. 226 (1891).

irregu

-2 mm

Sydow, Mon. Ured. I., p. 681 (1903).
Sacc. Syll. XL, p. 199 (1895).

II. Uredosori hypophyllous, on brown effused spots, scattered

larly or aggregated, ellipsoid to oblong, variable in size, 1

lon<r, long covered by the thick epidermis, ochraceous or brownish.

Uredospores yellowish-brown, ellipsoid, echinulate, 22-25 x

19-21 ».

III. Teleutosori hypophyllous, on similar spots, 1-3 mm. long, almost

always covered by the epidermis, blackish brown.

Teleutospores intermixed with uredospores, elongated oblong to

gated fusiform, brown and paler beneath, smooth, constricted

dly constricted

57 X

ge 48 x

15 \x ; upper cell rounded or often obtusely and obliquely

at apex, and thickened (up to 13

cell paler, oblong, but

usually attenuated towards base ; pedicel coloured like lo
persistent, 20-30 /x long, but may reach a length of 40 \x

cell,

X. Mesospores generally paler in colour than the teleutospores, ellip-
soid to fusiform or even curved, obtusely and sometimes obliquely
pointed at apex, thickened (up to 5 ju), 30-32 X 13-17 /t ; pedicels
similarly coloured, persistent, up to 22 /.i long.

On leaves of Nut grass (Cyperus rotundus L.).

Queensland— Gatton, Aug., 1903 (Shelton) (Bailey, 18, 20).

The teleutospores are very variable in shape, and sometimes the cells
may be placed almost at right angles to each other. They are sometimes
excessively elongated, and may reach a length of 67 /i.

Intermixed with the teleutospores there are numerous stalked bodies
similarly coloured, and representing what are usually called mesospores,
although Sydow describes them as paraphyses. While mesospores generally
have an undoubted resemblance to the teleutospores with which they are
associated, their function in this case seems to be that of paraphyses, and
it may be in some other cases as well.

Darhica filum Cast., was common on uredosori.

(Plate IV., Fig. 32.)

Puccini a — Cyp erac eae

*35

57. Puccinia longispora McAlp.

Car ex.

II. Uredo-sori hypophyllous, minute, elliptic to elongated, confluent in

long lines, long covered by epidermis, rusty- brown.

Uredospores sub-globose to shortly elliptic or obovate, yellowish-
' brown, thick-walled, echinulate, with three equatorial <*erm-pores

on one face, 22-30 x 19-22 /*, or 23-27 jx diam.

III. Teleutosori black, compact, elongated, bullate, crowded, confluent

in long black lines, covered for some time, then girt by ruptured
epidermis.

Teleutospores clear yellowish-brown to smoky-brown, elongated,

narrow, smooth, constricted at septum, 50-80 x 14-19 yu, average

63 x 19 fx ; upper cell darker than lower, generally rounded and

m slightly expanded at apex, sometimes bluntly pointed, much

thickened (9-12 ^u) ; lower cell longer than upper, almost cylin-
drical or slightly attenuated towards base ; pedicel yellow, short
and often stout.

On Carex caespitosa L., and C. vulgaris Fr.

Victoria — Werribee Gorge, January, 1903. Gembrook Ranges,
April, 1904 (C. French, jun.)

On Carex sp.

Victoria — Killara, March, 1903 (Robinson).

In immature forms of teleutospore the upper and lower cells are much
alike, but ultimately the lower cell is much elongated and nearly equally
broad throughout. Although 33 different Puccinias are recorded on species
of Carex, the elongated narrow teleutospore with short pedicel is quite
characteristic for this species. In P. schoeleriana Plow, and Mag., the teleu-
tospores may be equally long, but the lower cell is more cuneiform than
cylindrical, and the pedicel may reach a length of 57 \x. An examination of
teleutospores from Carex arenaria L. in Syd. Ured. Exs., No. 282, showed
these differences distinctly.

Specimens of P. caricis (Schum.) Rebent. were examined from Exsicc.
Sydow Ured. No. 1065 and Exsicc. Briosi and Cavara, No. 12$, and the
teleutospores were decidedly different.

The lower cell in P. longispora is much more elongated and narrower
and gradually merges into the short, stout and coloured pedicel, while in
P. caricis besides being generally considerably shorter, more wedge-shaped
and somewhat inflated, the pedicel was much narrower. It is in the upper
cell, however, that the difference is most striking. In P. caricis the thick-
ened apex is one-half the length of the cell, while in P. longispora it is only
one-third, taking the average of a number. The thickened apex, also, in
P. caricis is much darker than in P. longispora. Of course, as in so many
other cases where there are a number of species on the same genus of host-
plant, there is a family resemblance too, between the rusts, and P. caricis
and P. longispora have many points in common along with differences in
detail.

(Plate IV., Fig. 31.)

•

Puccini a — Juncaceae.

JUNCACEAE.

Juncus

58. Puccinia juncophila Cooke and Mass.

Cooke and Massee, Grev. XXII., p. 37 (189
Sydow, Mon. Ured. I., p. 64-1 (1903).
Sacc. Syll. XL, p. 199 (1895).

Uredo armillata Ludwig, Bot. Centrbl. XLIII. p. 8 (1890).

II. TJredosori ruddy-brown, ellipsoid, crowded, confluent, often girdling

stem, surrounded by the ruptured epidermis and sparingly covered
by shreds of it.

Uredospores sub-globose, elliptic or piriform, very spinulose,
relatively thick-walled (3-4 fi) yellowish-brown, 24-32 x 16-22 p.

III. Telutosori dark-brown, elliptic to oblong, surrounded by ruptured

epidermis, confluent.

Teleutospores intermixed with
olden-brown, with finely granular contents, slightly constricted

uredosp

ptical to oblon

Jed and a little thicker at apex (5-6 /x)

at septum, rour

rounded at base and generally resembling upper, but often some

what narrower, 30-40 X

thick,

smooth :

ped

10-28
hyali

ge 36

ally decid

25 a

epispo
, oftt

ttached obliquely, up to 54 p long

X. Mesospores similarly coloured to teleutospores, shortly elliptical,

occasionally globose, smooth, thickened at apex, 23-28 x 19—21 yu.

On Juncus effusus L., J. mxritimus Lam., J. pallidus R. Br., J.
pauciflora R. Br., and other species.

Victoria — Coromby, Oct., 1889, II. (Tepper), the original of
Uredo armillata Ludw. Diraboola, Nov., 1891, II., III.

(Header).

Oakleigh, April, 1893, II., III. (Morrison).

Ardmona, Christmas Hills, Killara, Millbrook, Minyip,
Mordialloc, Myrniong, near Melbourne, Phillip Island,
Stawell, Werribee Gorge, Murramurrangbong Ranges, &c.

Tasmania — (Rod way 1).
In the original description of this species by Cooke and Massee, some

mistake must have been made in the measurement of

they are given as 16

18 x 12-14 u.

edosp

Through the kindness of Dr. Morrison, I ha
some of the original material from Oakleigh.

been able to examine

Uredo armillata was the name given at first by Ludwig in 1 889, f r

edospores only being found on Juncus

y of teleutospores intermixed with uredosp

but the subsequent dis-
ss showed that it was a

Puccinia.

referred to Uromyc

Material sent by Reader to Kew Herba

was

«y

specimens

beinq

by the authorities there, but the identical

later forwarded to us the

celled

found mixed with the uredospores
shown to exist in Victoria.

Uromy

por

were

therefore, has not bee

Darluca fit

Cast

was

y common

in some cases rendering the

patches quite black. It occurred both on uredo and teleutosori

(Plate IV., Figs. 35, 36.)

Puccinia — Juncaceae\

137

59. Puccinia tenuispora McAlp.

Luzula

II. Uredosori hypophyllous, round to ellipsoid, scattered or often con-
fluent in lines, yellowish-brown, soon naked and girt by the
ruptured epidermis, compact, on irregular, confluent, purplish
spots.

Uredospores yellowish to pale yellowish-brown, elliptical to
obovate, echinulate, with 1-2 germ-pores on one face, 20-25 x
15-22 u.

III. Teleutosori dark-brown, ellipsoid, scattered or crowded, often con-
fluent, pulvinate, compact, soon naked, on similar spots.

Teleutospores oblong to clavate, ochrey-yellow, fragile, smooth,
apex generally rounded, rarely truncate or conical, thickened up to
1 1 /u, attenuated towards base or rounded, constricted at septum,
30-50 x 14-20 fi, average, 42 x 17 /u ; pedicel hyaline, per-
sistent, generally about 30 \x long.

X. Mesospores abundant, clavate to obovate or oblong

smoo

ened at apex and similarly coloured to teleutospore, 22-32 x
13-16 /u.

On leaves and stems of Luzula campestris L.

Victoria — Murramurrangbong Ranges, Nov. and Dec, 1902-3,

Jan., 1905 (Robinson). Wandin, Nov., 1903 (C. French,
jun.'

On Luzula oldfieldii Hook. f.

• Tasmania— Mt. Wellington, Jan., 1892, II. (Rodway).

Cooke in his Handbook of Australian Fungi gives Aecidium bellidis

Thuem. which is supposed to represent the aecidial stage of Puccinia obscura
Schroet., occurring on Luzula, but the aecidium found on Bellis perennis here
is associated with its own teleutospores, and belongs to a distinct fungus,
P. distincta McAlp, so that whatever may be the case in Britain, the rust
on the daisy is autoecious with us.

The discovery of a rust on Luzula campestris, which does not belon

the Old World species, is a further proof against its genetic connexion with
that of the daisy. The species differs from P. obscura and P. oblongata.
The teleutospores are characteristically fragile, thin- walled, and easily
collapsible, and while of the same general type as in P. oblongata, they are
much smaller. In P. obscura the colour is much deeper, and the wall is de-
cidedly thicker and firmer. This is well shown in Plate IV., Fig. 34, in
which the spores are taken from a specimen of Luzida campestris in Sydow's
Ured. Exs., 1076, collected in 1896, and the material from which our de-
scriptions are made is as recent as 1905. In P. oblongata the apex of the
teleutospore is much thicker, being 10-20 /u, and the size is 40-80 X 16-24 /u.

The teleutospores are frequently found germinating, showing that they
do not require to undergo a period of rest.

Darluca filum Cast, occurs frequently on the uredosori.

(Plate IV., Figs. 33, 34.)

138

Puccinia — Liliaceae

LILIACEAE.

Bur char dia

60. Puccinia burchardiae Sacc.

Saccardo, Hedw. XXXII., p. 57 (1893).

Ludwig, Zeitsch. f. Pflanzenkr. III., p. 137

McAlpine, Proc. Roy. Soc. Vic. VII., N.S., p. 215 (1894).

Sydow, Mon. Ured. I., p. 620 (1903).

(1893).

Sacc. Syll. XL, p. 197 (1895).

II. Uredosori amphigenous, bullate, elliptical or sometimes circular,

crowded, light-brown, erumpent, surrounded at base by dry cuticle
of epidermis.

Uredospores elliptic to subglobose, yellowish-brown, epispore
echinulate, 28-31 x 22-25 p.

numerous, small or large, erumpent

III. Teleutosori amphigenous,

surrounded at base by cuticle of epidermis, circular cr elliptical,

black.

Teleutospores clavate, constricted at septum, thickened at apex

rounded, sometimes truncate or acute.

to 14 or

40-60 x

gth of 70

48 x 20 u, but sometimes

u; lower cell tap

towards pedicel, yellowisl

'own; upper cell elliptical or rarely spherical, chestnut-brown,
21-23 n broad; epispore smooth; pedicel hyaline, 30-60 X 5-6 jj.

X. Mesospores fairly common, similarly coloured to teleutospores or

paler, clavate to (
37-43 x 12-15 u

ded or pointed at apex and thickened

On stems and leaves of Eurchardia umbellata R. Br.

Victoria — Dandenong Creek, Oct., 1891. (C. French, jun.).

Near

Melbourne, Sept. — Feb. Myrniong, Sept., 1898, and Dec,
1903. Murrumurrangbong Ranges, Dec, 1903 (Robinson).

S. Australia — Teatree Gully, Oct., 1890 (Tepper).

The first record of this species

appeared in Hedwigia

of March and

April, 1893, where Saccardo described it as

P. metanartheeii Pat

differing from the type

(Plate V., Figs. 38, 39.)

Wurmbea.

61. Puccinia wurmbeae Cooke and Mass.

Cooke and Massee, Grev. XVI., p. 74 (1887).
Cooke, Handb. Austr. Fung., p. 337 (1892).
Sydow, Mon. Ured. I., p. 640 (1903).
Sacc. Syll. VII., p. 664 (1888).

Sori elongated, bullate, dark-brown.

II. Uredospores elliptic, granulate, brown, 25-28 x 15-18 u

III. Teleutosp

clavate,

htly constricted at septum, smooth

upper cell convex or truncate, at apex darker, lower cell tri-

5 into the short, hyaline pedicels

angular, attenuated downward
48-60

X 21-28 /z, average

Massee, 60-70 x 20-25 At) ;

0 x

but attaining a length of

broadest at the junction with the spore

ii (according to Cooke and
pedicels persistent, generally short,
ix and a breadth of

8 ft, bein

Puccinia — Haemodoraceae, Amaryllidaceae. 139

X. Mesospores numerous, similarly coloured to teleutospores, elon"

elliptical to oblong, thickened at apex and acute, rounded or
cate, 40-49 x 17-21 fi.

On leaves of Wurmbea dioica F. v. M.

South Australia — Beltana, 1887 (Richards).

No uredospores were found on the specimen in National Herbarium

(Plate V., Fig. 37.)

HAEMODORACEAE.

Haemodorum

02. Puccinia haemodori P. Henn.

Hennings, Hedw. XL., p. (96) (1901).
Sydow, Mon. Ured. I., p. 609 (1903).
Sacc. Syll. XVII., p. 366 (1905).

III. Teleutosori amphigenous, round or often oblong, scattered or

aggregated and then confluent, pulvinate, compact, erumpent, black,
surrounded by the ruptured epidermis, 1-2 mm. long.

Teleutospores oblong to oblong clavate, obtusely rounded or

apiculate at apex and strongly thickened (8-13 ju), slightly con-
stricted at septum, attenuated or rounded at base, smooth,
chestnut-brown, 40-57 x 14-20 /u, average 46 x 19 ji, pedicel
persistent, slightly brownish towards apex, 40-55 x 5-7 jx.

X. Mesospores very common, subfusoid or ovoid to subclavate, apex

gibbous to apiculate, somewhat obtuse or two-horned, thickened at
apex (up to 9 //), brown, 25-40 x 12-18 jjl ; pedicel hyaline to
brownish. Occasionally a more deeply coloured spore occurs,
with somewhat truncated apex, 30-46 x 5-7 fi.

On leaves of Haemodorum sp.

West Australia— Near Perth (1900).

Hennings described uredospores in his original diagnosis, but Sydow did
not find them afterwards in the original material. He remarks that the
teleutospores readily separate into their two constituent cells, and since the
supposed uredospores agree with the upper cell of the teleutospore, they are
probably the same.

No uredospores were found by me in the specimen kindly sent by
Hennings.

The teleutospores are somewhat variable in shape, and occasionally the
upper cell may be ellipsoid and deep chestnut, while the lower is cylindrical,
much paler in colour, and two-thirds the entire length.

(Plate V., Fig. 40.)

AMARYLLIDACEAE.

63. Puccinia hypoxidis McAlp.

Hypocois.

McAlpine, Agr. Gaz. N.S.W. VI., p. 853 (1895).
Sydow, Mon. Ured. I., p. 607 (1903).
Sacc. Syll. XIV., p. 341 (1899).

II. Uredosori on both surfaces and margins of leaves, minute, rust-
coloured, rounded or oval, bullate, gregarious or scattered, bursting
through and surrounded by epidermis.

140

Puccini a — Labiatae.

Uredospores shortly elliptical, yellowish, finely echinulate, 20-23
X 16-18 /i, average 21 x 16 yu.

III. Teleutosori amphigenous, black, blister-like, largely confluent and

distorting leaf, long covered by epidermis.

Teleutospores with a few uredospores intermixed, elongated
clavate, chestnut-brown, slightly or not at all constricted at septum,
commonly truncate and much thickened at apex, 36-54 x 15-22 p

average 42 x 18

upper cell squarish or oblong or club-shaped,

apex very much thickened (up to 15 /j), sometimes strongly beaked,
generally shorter than lower cell, dark chestnut-brown; lower cell
tapering towards base, lighter in colour than upper, of a yellowish
or golden brown ; pedicel pale yellow, persistent, generally about
18-20 fi long.

X. Mesospores chestnut to pallid, rounded or beaked at apex and

thickened,

11-18 /Lt.

hat fusiform to elongated elliptical, 25-36 x

On leaves of Hypoxis glabella II . Br.

Victoria — Ardmona, July, 1893 (Robinson). Burnley, Ruther-

glen, &c, July-Oct.

It differs from P. burchardiae

Sacc

the much

redosp

and decidedly in the teleutospores, which in the latter are generally rounded
at the apex or beaked, and distinctly constricted at the septum.

(Plate V.,, Fig. 41.)

LABIATAE

Mentha

64. Puccinia menthae Pers.

Persoon, Syn., p. 227 (1801).
Sydow, Mon. Ured. I., p. 282 (1902).
Sacc. Syll. VII., p. 617 (1888).

Spermogonia either arranged

coloured.]

groups or scattered, honey

Aecidia hypophyllous, or frequently on stems which are much

mor

ely on purplish-red spots on leaves, seldom scattered
pseudoperidia immersed, flat, opening irregularly and
scarcely or irregularly torn.

Aecidiospores subglobose, ellipsoid or polygonal, coarsely granular

margin

pale yellowish, 24-40 x
II. Uredosori hypophv

7-2

seated on pale spots

ptical, scattered or aggregated, soon pulverulent and confluent
by the ruptured epidermis, cinnamon-brown.

Uredosp

subglobose, ellipsoid or obovate, echinulate, pale

brown, generally three equatorial pores, 20-30 X 16-19/1
III. Teleutosori similar, but blackish-brown in colour

Teleutosp

first intermixed with uredospores, ellipsoid to

rounded at both ends, apical papilla pale or hyaline, broadly

xpanded, hardly constricted at sept

chestnut-brown, 24-32 x
1 9-20 a ; pedicel hval

finely

ted, golden to

spore

19-22 fx, but commonly 24-25 X
slender, generally surpassing length of

Puccinia — Acanthaceae. 141

X. Mesospores, occasional, similarly coloured, nearly globose, very

slightly roughened, 20 p diam.

On leaves of Mentha laxijiora Benth.

Victoria — near Melton (Reader).

On leaves of Mentha pulegium L.

Victoria— Myrniong, Aug., 1904, II., III. (Brittlebank).

The teleutospores are prominently warted.

(Plate XXIX., Fig. 250.)

ACANTHACEAE

65. Puccinia mussoni McAlp.

Ruellia

Sori amphigenous, minute, usually rounded, bullate, scattered or some-
times in groups.

II. Uredospores yellow with thick, chestnut wall, globose to shortly

ellipsoid, strongly echinulate, with two germ-pores on one face,
30-38 /1 diam., or 28-34 X 18-23 p.

III. Teleutospores intermixed with uredospores, dark chestnut-brown,

oblong, with thickened wall and roughened surface, sometimes
slightly constricted, rounded at both ends, not thickened at apex,
36-46 X 28-36 /z; upper cell resembling lower, but usually a little
larger ; pedicel generally lateral and even sometimes on a level
with the septum, flexuous, hyaline, up to 60 /j, long and 7 ^
broad.

On living leaves of Ruellia australis Cav.

New South Wales — Richmond River, June, 1896 (Musson).

The teleutospores were not very plentiful in the specimens examined, but
the sori containing them could be detected by their darker brown colour.
The lateral pedicel to the teleutospore naturally suggests P. lateripes B. and
Rav., but a closer inspection reveals important differences in the two kinds
of spores.

I have compared the Australian species with specimens on the leaves of
R. strepens L. from N. America in Sydow's Ured. Exs., No. 1374, and
Kellerman's Ohio Fungi, No. 130, and it is evident that we are dealing with
similarity of type due to the affinity of the host-plants, with considerable
divergence in the character of the spores. Unfortunately, P. lateripes B.
and Rav. and P. ruelliae (JB. and Br.) Lagh. are confounded, for although
Sydow labels his specimen as the former, in his Monograph he names it
the latter. In this specimen the finely echinulate uredospores are ellipsoid
to ovate, and measure 24-28 X 19-21 /i, while here they are larger. The

teleutospores likewise only measure 29-34 x 18-22 p, and are very strongly
warted.

This species differs from P. longiana Syd., in the larger uredospores and
the teleutospores not being thickened at apex, and from P. lateripes and
P. ruelliae in the larger size of uredo- and teleuto-spores.

(Plate V., Figs. 43, 44.)

!42 Piiccinia — Convolvulaceae.

CONVOLVULACEAE

Dichondra.

66. Puccinia dichondrae Mont.

Montagne in Gray's Fl. Chil. VIII., p. 46 (1845).
Cooke, Handb. Austr. Fung., p. 338 (1892).
Sydow, Mon. Ured. L, p. 321 (1902), and p. 881 (1904).
Sacc. Syll. VII., p. 717 (1888).

P. dichondrae Berk. Linn. Journ. XIII., p. 173 (1872).
P. berkeleyana De Toni, Sacc. Syll. VII., p. 717 (1888).
P. munita Ludwig, Zeitsch. f. Pflanzenk. II., p. 133 (1892).

III. Teleutosori hypophyllous, punctiform, very minute, erumpent to

superficial, somewhat pulverulent, densely gregarious and often
covering entire surface, deep cinnamon, nestling among hairs,

70-100 fx diam.

Teleutospores clavate to oblong, golden brown, constricted at
septum, with comparatively thin epispore, thickened at apex and
produced into a hyaline apiculus (occasionally two), with granular
contents, and each cell 1-guttulate, occasionally 3 to 4 celled,
30-40 x 12-18 p, average 32 x 14 ju ; pedicel hyaline, sometimes
tinted, persistent, generally slender, up to 46 ^t long, occasionally
6 fj. broad.

X. Mesospores sub-ellipsoid to oblong, similarly coloured to teleutospore,

thickened at apex and usually with hyaline apiculus, 21-30 X
15-22 /u.

On Dichondra repens Forst.

Victoria — Near Melbourne, 1886 (Reader). Cheltenham, Nov.

1887 (French). Goulburn River Flats, Oct., 1896 (Robin,
son). Christmas Hills, May, 1900 (Robinson). Point Cook
May, 1902 (French, jun.). Murramurrangbong Ranges, Nov.
1902 (Robinson). Mt. Blackwood, Dec, 1902. Port Fairy,
Aug., 1905. Mentone, throughout the year, 1905, etc.

New South Wales— 1901 (Camfield). Richmond, April, 1905

(Musson). Recorded by Maiden1.

South Australia — Mount Lofty, near Adelaide, Oct., 1891 (Tepper).
Tasmania — North East (Mueller).

I have examined the original material from the National Herbarium,
Melbourne, and find that there must have been some mistake over the

measurements of the spores, which were given as *005 inch (130 /u) by
Berkeley, which is evidently a misprint. Next, Dr. Cooke in his Handbook of
Australian Fungi makes a correction by giving the size of the spores as
12-13 fi long, which evidently errs on the other side. Then De Toni
changed the name to P. berkeleyana, seeing that the size of the spores as
given did not at all agree with those of P. dichondrae, Mont.

Another unfortunate error has arisen in connexion with this species
through the wrong naming of the host-plaut. Mr. Tepper sent a rust to
Prof. Ludwig from S. Australia, said to be on the living leaves of Hydrocotyle
hirta R. Br., who determined it as a new species, P. munita. But on Prof.
Ludwig kindly sending me some of the original material it was found that
the leaves belonged to Dichondra repens and that the rust was P. dichondrae,
a conclusion with which Prof. Ludwig agreed after examining the specimens
sent to him.

Puccinia — A-pocynaceae. 143

The teleutospores are frequently 3-4 celled, and there is great variety in
the arrangement of the septa. The spore may be divided vertically as in
D ior chidium, or it may have a vertical or oblique septum in its upper or
lower cell. . The septa may even be arranged muriformly. There may be
a lateral hyaline apiculus in the lower as well as in the upper cell, and the
pedicel may stand out at right angles to the lower cell.

There is thus every gradation from the unicellular spore and the bicell-
ular, in which the upper cell is more or less atrophied, up to the multicellular,
which is vertically, obliquely or muriformly divided.

Aecidia have been found on this plant and are regarded as belonging to
this species, but although numerous specimens have been examined from
various localities no trace of aecidiospores has been found here.

It is worthy of note that some of our native species of Viola closely
resemble the Dichondra, and the one may easily be mistaken for the other
when no flowers are present.

(Plate V., Fig. 42 ; Plate XL., Fig. 299.)

APOCYNACEAE.

67. PuCCillia alyxiae Cooke and Mass.

A lyxia

Cooke and Massee, Grev. XVI., p. 2 (1887).

Cooke, Handb. Austr. Fung., p. 338 (1892).
Sydow, Mon. Ured. I., p. 336 (1902).
Sacc. Syll. VII., p. 714 (1888).

III. Teleutosori generally hypophyllous, occasionally epiphyllous,

discoid, compact, dark-brown, girt by the ruptured epidermis,
l-2mm. diam.

Teleutospores almost pear-shaped or oblong, yellowish to
brownish, constricted in the middle, thickened at apex and
generally apiculate, sometimes rounded or truncate, occasionally
3-celled, 40-52 x 20-25 /jl, average 45 x 20 /* ; epispore thick,
smooth, coloured ; pedicels persistent, hyaline, broadish, elongated,
up to 130 fA, occasionally at right angles to the spore.

X. Mesospores not uncommon, similarly coloured, elongated oblong,

thickened and sometimes apiculate at apex, 38-50 X 19-21 p.

On leaves of Alyxia buxifolia R. Br.

Victoria— Brighton and Broadford, Sept., 1887 (Miss Campbell1).

Beaumaris, March, 1895. Cheltenham, May, 1902 (C.French,

Sandringham, April, 1905. (Robinson).
South Australia — Gawler, Sept., 1893. (Tepper).
Tasmania — Near George's Bay, Nov., 1892. (Rodway 1).

The sori are generally isolated, much inflated and surrounded by a
conspicuous, brown, elevated ridge. When on both surfaces they are
opposite to each other.

In the Cheltenham material, the spores had germinated on the leaves
lying on the ground on 19th May, while the spores on the Gawler material
found in September had not.

This spore probably undergoes a period of rest during the summer
before germination. Cooke and Massee have given the length of the spores
as 50-70 n, but this is evidently a misprint.

(Plate VI., Fig. 46.)

144

Puccinia — Apocynaceae, Goodeniaceae.

Carissa

68. Puccinia carissae Cooke and Mass.

Cooke and Massee, Grev. XXII., p. 37 (1893)
Sydow, Mon. TJred. I., p. 336 (1902).

Sacc. Syll. XL, p. 195 (1895).
III. Teleutosori hypophyllous, gregarious, on

orbicular spots, forming

rings which are at length confluent, rather compact, dark-brown.

Teleutospores elliptic, constricted at septum, rounded at ends or
occasionally flattened, smooth, brown, 30-34 x 17-25 n, average
32 x 22

occasionally 3-celled, then reaching to a length of

deciduous or persistent, sometimes origin-

49 fx ; pedicel hyaline,
ating laterally, rather long.

X. Mesospores occasional, just resembling a teleutospore, but without

the septum, elliptic, with hyaline pedicel, 29-31 x 22-25 jx.

On living leaves of Carissa ovata R. Br.

Queensland — Gladfield, (Gwyther) (Bailey 13). Dalby, (Bancroft)

(Bailey

Mr. Bailey has kindly sent me some excellent material from his herbarium

In the original

and the teleutospores have been carefully measured,
description the size is given as 20-22 x 16 ft, but probably 30-32 ll was in-
tended for the length.

Sometimes both upper and lower cell have an oblique division.

It differs from P. aly

Cooke and Mass., in which the spores are much

and thickened at the apex

(Plate VI., Fig. 45.)

GOODENIACEAE

Brunonia

69. Puccinia brunoniae McAlp.

McAlpine, Agr. Gaz. N.S.W. VI., p. 851 (1895).
Sydow, Mon. Ured. I., p. 193 (1902).
Sacc. Syll. XtV., p. 320 (1899).

0. Spermogonia dark honey-coloured, disposed in clusters on yellowish

spots, or intermixed with aecidia on the same surface of the leaf,
usually the upper.

1. Aecidia scattered on

more often

cularly arranged

eddisl

htly swollen indeterminate spots, mostly on upper surface of

leaves and

slightly recurved

petioles ; pseudoperidia with wh

margin,

d soon

falling

away,
isolated peridial cells, generally short and

torn,
mm. diam.;
stout, less commonly

average

elongated

obi

punctate, with striated

24-31 x 14-17 /i, or

Aecidiospores subglobose

smooth, 17-21 u diam. or 1

margin

■)iy

when elongated 38

45 x 21-24 ii

or even

1 x

5-16

deep

ionally as long

Puccinia — Goodeniaceae

*45

III. Teleutosori amphigenous, but mostly on upper surface, very rarely

mixed with aecidia, forming clusters of black, bullate pustules on
indefinite, pale, ruddy, thin spots, oval or elongated, usually
confluent, arranged circularly, or in lines, at first girt by
ruptured epidermis, which finally falls away.

Teleutospores clavate, golden-brown to chestnut, thickened at
apex, constricted at septum, sometimes 3 or 4 celled, 45-60 x
18-21 /j, average 50 x 20 fx ; upper cell generally pointed, but
often rounded and occasionally truncate, deeper in colour than
lower ; lower cell tapering towards base and generally longer than
upper; pedicels persistent, pale-yellowish to hyaline, 40-50 x

9-10

X. Mesospores occasional, similarly coloured to teleutospores, rather

oblong, thickened at apex, rounded or truncate, slightly narrowed
at base, 28-38 x 17-21 F.

On leaves and petioles of Brunonia australis Sm.

Victoria — Drysdale, Oct., 1895, III. Murramurrangbong Ranges.

Nov., 1902, Dec, 1903, O., I., III., and Jan., 1905, III,
(Robinson). Alps, near Bright, Dec, 1904, III. (C. French,
jun.). Rutherglen, Dec, 1904, III.

Three-celled teleutospores occasionally seen, varying in length from
60-73 fi and in breadth from 21-24 ju, the lower cell generally about
as long as the other two.

Four-celled teleutospores are very rare, elongated clavate, 63 X 24 /u,
the two upper cells about 33 x 24 f.i, and the two lower 30 x 14 /u. In
the same group there were three 3-celled teleutospores with an average
size of 59 x 23 ji.

At first no aecidia were known, but these were found along with the

teleutospores by G. H. Robinson.

The discovery of aecidia rendered it necessary to carefully compare the
two fungi found upon the Goodeniaceous plants, Brunonia and Goodenia,
and a large number of specimens and slides were accordingly examined and
compared.

While the rusts are of the same general type, they differ in several
important respects. The aecidiospores in P. saccardoi are considerably
smaller, and the peridial cells are generally much more elongated, and more
than twice as long as broad.

In the original description by Dr. Ludwig the size of the aecidiospores
is given as 13-15 jjl diam., and the peridial cells as 18-25 x 15-18 ju, but

if the latter are measured when isolated they are found to be much longer.
Again in P. saccardoi the teleutospores are generally intermixed with or
surround the aecidia, but this very rarely occurs in P. brunoniae. In the
former the teleutospores are generally rounded at apex, but in the latter
generally pointed and altogether narrower.

In the Murramurrangbong Ranges where the aecidia and teleutospores
of P. brunoniae were found very plentifully in November, though
Goodeniaceous plants which, in other districts are affected by P. saccardoi y
were exceedingly common and were often growing alongside the Brunonia,
no rusts were found on them, even although the country was subjected to a
most exhaustive search. The rusts, therefore, on the two different genera
are considered to be specifically distinct.

(Plate VI., Fig. 50.)

146 Puccinia — Goodeniaceae

70. Puccinia dampierae Syd.

Sydow, Mon. Ured. I., p. 193 (1902).
Sacc. Syll. XVIL, p. 315 (1905).

I. Aecidia on stems and both surfaces of leaf, gregarious, bright

orange ; pseudoperidia with white reflexed and torn margins, i-^ mm.
in diam. ; peridial cells lozenge-shaped to oblong, with thick striated
margins, 30-40 jx long.

Aecidiospores orange, subglobose, ellipsoid to oblong, finely
echinulate, 15-17 /x diam. or 17-19 X 13-14 ^u.

III. Teleutosori on stems, scattered or aggregated, round or oblong,

about 1 mm. diam., compact, dark-brown, girt by the ruptured
epidermis.

Teleutospores clavate, rounded or acute at apex and strongly
thickened (8-11 lx), constricted at septum, attenuated down-
wards, rarely round, smooth, brown, 48-66 x 16-26 li ; pedicel
yellowish, persistent, up to 80 li long.

X. Mesospores intermixed with teleutospores, clavate, 40-55 x 14-22 li.

On stem and leaves of Dampiera stricta K. Br.

Victoria— Monbulk, Dec, 1905, I. (C. French, jr.).

On wings of the stems of Dampiera alata Lindl.

West Australia, III. (L. Preiss).

Though I have not seen the West Australian specimens I prefer to
regard the aecidia found in Victoria as a stage of the same fungus. If they
should prove to be unrelated they would be easily separable at any time.

71. Puccinia gilgiana P. Henn.

Leschenaultia.

Hennings, Hedw. XL., p. (95) (1901).
Sydow, Mon. Ured. I., p. 194 (1902).
Sacc. Syll. XVIL, p. 314 (1905).

Aecidium perkinsiae P. Henn. Hedw., XL., p. (96) (1901).

I. Aecidia on thickened and slightly deformed parts, gregarious or

scattered ; pseudoperidia hemispherical to cup-shaped, semi-
immersed, yellowish to ruddy ; peridial cells round or oblong-
polygonal, subhyaline, reticulate, 18-24 X 16-20 li.

Aecidiospores subglobose or ellipsoid, angular, finely echinulate,
yellowish, 15-20 li diam.

II. Uredosori on stems, gregarious in streaks, oblong, surrounded or

almost covered by the ruptured brown epidermis, somewhat
pulverulent.

Uredospores subglobose, ovoid or ellipsoid, yellowish to brown,
echinulate, 14-21 x 12-18 li.

III. Teleutosori similar, but black.

Teleutospores clavate or oblong, constricted at septum, smooth,
obtusely rounded or truncate at apex and slightly thickened (up
to 5 li), obliquely papillate and occasionally crested, rounded at
base or attenuated, brown, 30-45 X 16-22 li, average 42 X 20 ft;
pedicel brownish, thick, persistent, 30-50 a long.

Puccinia — Goodeni

stems

X. Mesospores very common, oblong or oblong-clavate, thickened at

apex, rounded or acute, chestnut-brown, smooth, stalked 23-32
X 15-20/1.

On petioles, calyx and stems of Leschenaultia linarioides DC.

West Australia— Near Perth, 1900 (Pritzel).
Aecidia on petioles, flower-stalks, and calyx ; uredo and teleutosori on

Specimens kindly supplied by Hennings.

(Plate VI.,Fig. 47.)'

/ 2. Puccinia saccardoi Ludw.

Goodenia, Velleia

Ludwig, Hedw. XXVIIL, p. 362 (1889).
Cooke, Handb. Austr. Fung., p. 337 (1892).
Sydow, Mon. Ured. I., p. 193 (1902).
Sacc. Syll. IX., p. 309 (1891).

P. nigricaulis McAlp., Agr. Gaz. N.S.W. VII., p. 151 (1896).
Aecidium g oodeniacearum (in part) Berkeley, Linn. Journ.

XIII., p. 173 (1872).

I. Aecidia on yellowish or brownish spots, scattered or in groups
I arranged in circular patches (up to 6mm. diam.), amphigenous

causing distortion of stem ; pseudoperidia cup-shaped, with white
torn, everted edges, 215-325 /jl, diam. ; isolated peridial cells
punctate, with striated margin, generally elongated oblong, or
somewhat oval or lozenge-shaped, more than twice as long as
broad, 35-42 X 16-21 ^.

Aecidiospores subglobose, polygonal to oval, orange, very finely

verrucose, 14-16 /i, diam., or 15-17 x 13-14 ^u.

III. Teleutosori on both surfaces of leaves and on stems, black, com-
pact, roundish or elongated, confluent in long swollen patches,
generally intermixed with or surrounding aecidia, bursting through
epidermis.

Teleutospores clavate to oblong, dark-brown, constricted at
septum, smooth, variable in size, rarely 3-celled, 40-66 x 17-25//,
occasionally up to 70 jx long, average, 54 x 20 // ; upper cell
subglobose or somewhat quadrate, thickened at apex (up to 9 /u),

rally rounded or truncate, occasionally pointed, 21-30 x

17-25 /i, sometimes reaching 32 fi in length; lower cell generally
paler in colour than upper, elongated and tapering towards base,
sometimes subglobose, longer and narrower than upper, 17-35 X
16-21 fi, occasionally 38 /t long; pedicel usually persistent,
hyaline, occasionally pale-yellow, 35-60 X 7-12 /<.

X. Mesospores common, similarly coloured to teleutospores or paler,

variously shaped, ellipsoid to oval, or oblong to clavate, smooth,
with pointed and thickened apex (up to 9 /*), and generally

slightly tapering towards base, 25-49 x 12-21 /*, or averagin
36 x 17 fx

148

Puccinia — Campanulaceae .

On stems, leaves and calyx of Goodenia geniculates It. Br., G. glauea
F. v. M., G. pinnatifida Schlecht., G. albiflora Schlecht., G. hederacea
Sm., and G. ovata Sm.

Victoria— Port Phillip, 1886, I. Dimboola, Sept., 1892, I., III.

(Reader). Minyip, near Colac, Grampians, Warracknabeal,
Nhill, Killara, Mt. St. Bernard, Borung, Gembrook, near

Melbourne.

S. Australia — Tanunda Scrub, Oct., 1887 (Tepper). Mt. Brown

Creek, near Quorn, Sept., 1892 (Molineux).

Tasmania — Cascades, Jan., May, Nov., 1892 and 1893,

(Rodway1). Devonport, Jan., 1906, I., III. (Robinson).

On Velleia macrocalyx De Yriese.

New South Wales — Guntawang, Sept., 1886 (Hamilton).

On Velleia paradoxa R. Br.

Victoria — Pine Plains, I. (Reader).

New South Wales — Guntawang, Sept., 1886, I. III. (Hamilton).

I have received from Prof. Saccardo some of the original material on
G. geniculata from S. Australia, and on comparing it with P. nigricaulis
McAlp., have no doubt but that they are the same. The Victorian
specimens, however, occur very plentifully on the stems as well as on the
leaves, and under favorable conditions the fungus obtains such profuse
development as to cause blackening and distortion of the stems.

Aecidium goodeniacearum Berk, belongs partly to P. saccardoi and to
Uromyces puccinioides. In specimens of Goodenia pinnatifida, received from
F. M. Reader, Dimboola, the fungus is' named A. goodeniacearum Berk, in
Massee's handwriting, while the teleutospores of P. saccardoi are also met
with intermixed with the aecidia.

In Velleia paradoxa, from New South Wales, while the teleutospores
are of the general type, there are quite a number in which the apex is
truncate and prolonged laterally into a thickened, more or less beak-like

projection.

But

Ludwig

observed in the

original

type on Goodenia
the spores on the genera Goodenia and

geniculata a similar variety, so that
Velleia cannot be differentiated. He writes — " Mixed among the normal
teleutospores there are three and one-celled spores, the latter sometimes of
abnormal size, as I have observed under similar conditions in P. heterospora.
Frequently very peculiar forms appeared, in which the upper cell bore 1-3
horn-like projections as long as the cell itself, or finger-like cells, as long as
the upper, proceeded from the lower cell."

(Plate VI., Figs. 48, 49.)

CAMPANULACEAE

73. Puccinia aucta Berk, and F. v. M.

Lobelia, Pratia,

Berkeley and F. von Mueller, Linn. Journ. XIII., p. 173

(1872

Cooke, Grev. XL, p. 98 (1883).
Cooke, Handb. Austr. Fung., p. 338 (1892).
Sydow, Mon. Ured. I., p. 196 (1902).
Sacc. Syll. VII., p. 676 (1888).

Aecidium microstomum Berk. Journ. Linn. Soc. XIII, p. 173

(1872).
A. lobeliae Thuem. Grev. IV., p. 75 (1875).

Puccinia — Campanulaceae. 149

0. Spermogonia slightly raised, with projecting paraphyses, honey

coloured, in groups.

Spermatia numerous, hyaline, ellipsoid, 5-6 X 3 /jl.

1. Aecidia amphigenous, rounded or ovate, scattered or densely

gregarious and occupying the entire surface of the leaves or
petioles, at first covered by the epidermis, then free and rupturing
like a broken blister, vesicular, ochraceous ; pseudoperidial cells,
oval to oblong, covered with spines, loosely connected, 30-40 x
18-25 11.

Aecidiospores irregularly globose, sub-angular or ellipsoid, finely
echinulate, with distinct wall, ochraceous, 18-22 n or 22-28 x
18-20 u.

III. Teleutosori on under surface, numerous, bullate, then girt

ruptured epidermis so that they resemble little Pezizae.

Teleutospores cylindric to sub-clavate, elongated, yellow,
thickened apex (9-10 /i), rounded or truncate, deeply constricted
at septum 56-70 x 15-26 /u, average 60 X 24, occasionally bi-
and tri- septate when they may reach a length of 80 \x ; upper cell

rally broader th

rally cylindrical, and

may taper slightly towards pedicel ; pedicel hyaline, short, stout,
7-13 fi broad.

Aecidiospores on Lobelia anceps L., L. pratioides Benth., L . jjurpurascens
R. Br., Pratia erecta Gaudich., P. pedunculata Benth., P. platycalyx Benth.

Victoria — Colac (F. v. Mueller), (von Thuemen). Murtoa, Oct.,

1892, (Reader). Ringwood, Oct., 1892 (Robinson). Minyip,
Oct., 1901 (Eckert). Werribee Plains, June, 1902 (C.
French, jun.).

New South Wales — Guntawang, Sept., 1886 (Hamilton). Kurra-

jong Heights, Apr., 1894 (Musson).

Tasmania — Bellerive Swamp, Dec, 1891 (Rod way).
Teleutospores on leaves of Lobelia anceps L.

South Australia— Port Lincoln, Nov., 1852 (C. Wilhelm

(Berkeley

The aecidium was first determined by Thuemen on a specimen of

Pratia platycalyx sent by the late Baron von Mueller from Colac, and named
A. lobeliae. Then Berkeley received a specimen from the same source on
P. pedunculata which he named A, microstomum.

I have examined portions of the original material of both these
specimens, and as a rule it is difficult to detect a peredial wall even in fine
sections, so much so that they were as first taken for caeomata. But there is
occasionally a loosely fitting layer of surrounding cells, so that we may
regard them as aecidia. The aecidial stage is given in connexion with the
teleuto stage since, although not found actually together, they occur separately
on the same species, Lobelia anceps.

The teleutospore is occasionally tri-septate, and is then slightly curved,

and the lower septum oblique.

The description of the' teleuto stage is drawn up from original specimens
the National Herbarium, named in Berkeley's handwriting.

(Plate VI., Fig. 51 ; Plate XXXIX., Fig. 289.)

i5°

Puccinia — Comfo sitae

COMPOSITAE.

Scorzonet

74. Puccinia angustifoliae McAlp.

I. Aecidia in dense clusters on the

leaves, more sparingly c
about J to nearly J mm. diam. ; pseudoperidia with finely
everted margins.

stems

Aecidiospores
quadrate, 16 X

yellow, globose to elliptical or sub

3 u or 13-14 a diam

III. Teleutosc

confluent
thro win

on leaf and stem, black, long covered by epiderm

forming

gated

off

pidermis, d

patches, partially ruptu]
:t from or along with

Teleutosp

clavate to oblong, smoky-bro

;htly

and
idia.

con-

stricted at septum, smooth, generally rounded and thickened

ape

htly tapering towards base or often
average 50 X 20 // ; pedicel hyaline,

l (up to 9 /x), s]
rounded, 40-53 x 16-S
deciduous, generally short, up to 38 fx long and 9 ll broad

X. Mesosp

common, similarly colored to teleutosp

generally somewhat oval, thickened at ape
On Scorzonera anguslifolia L.

25-30 x 16

paler
17 a

Victoria— Dimboola, Nov., 1892, L, III. (Reader).

This

species differs from P. podospermi

DC,

scorzonerae

(Schum.) Jacky and P. tragopogi (Pers.) Corda, in the absence of
uredospores and the smooth teleutospores.

(Plate VII., Fig. 53.)

Brachycome.

75. Puccinia brachycomes McAlp.

I. Aecidia crowded on swollen and distorted portions of leaves, also on

stems and branches, about ^ mm. diam.; pseudoperidia round,

margin

out-standing, with white, slightly everted, laciniate
pseudoperidial cells firmly united, oblong to sub-quadrate, punctulate
all over, with broad striated margin.

Aecidiospores yellowish, sub~globose to ellipsoid, finely echin-
ulate, 13-16 fi diam. or 13-16 x 10-13 /jl.

II. Teleutosori elliptical, pustulate, black to blackish-brown,

long

covered by epidermis, crowded, confluent, and ultimately forming
elongated patches several mm. in length.

Teleutospores clavate, dark chestnut-brown, smooth, very

slightly constricted at septum, 48-64

18-25 n

average

50 x 20 n ; upper cell truncate or conoid, occasionally rounded at
apex, thickened (up to 9 ju) ; lower cell generally attenuated towards
base, sometimes rounded and generally longer than upper ; pedicels
persistent, tinted or hyaline, up to 35 fx long.

X. Mesospores not uncommon, similarly coloured to teleutospores,

ellipsoid, either rounded or pointed, and thickened at apex,
28-30 x 14-19 p.

On Brachycome ciliaris Less., I., III., and B. pachyptera Turcz., I.

Victoria— Dimboola, May-July, 1896 and 1898 (Reader).

On Brachycome scapiformis DC. and B. diversifolia, Fischer and Mey.

Victoria— Buffalo Ranges, Dec, 1904, I., III. (C. French, jun.).
Darluca ftlurn Cast., is a very common parasite on the aecidia.

(Plate VIII., Fig. 66.)

Puccinia — Compositae.

I5I

76. Puccinia calendulae McAlp

Calendula

■

McAlpine, Proc. Linn. Soc. N.S.W. XXVIII., p. 558 (1903).
Sydow, Mon. Ured. I., p. 852 (1904).
Sacc. Syll. XVII., p. 280 (1905).

Aecidium calendulae McAlp. Agr. Gaz. N.S.W. VII.,

p. 152 (1896).

I. Aecidia amphigenous or on stems, orange-yellow, in clusters up to

6 mm. diam, sometimes circinate, or may be scattered irregularly ;
pseudoperidia with white, torn and reflexed margin ; peridial cells
quadrate or polygonal, striated at margin, 21-24 p long.

Aecidiospores very irregular, sub-globose to polygonal, very finely
echinulate, pale orange, 14-17 X 11-12 ll or 12-16 ll diam.

III. Teleutosori intermixed with aecidia, black, erumpent, soon naked,

girt by the ruptured epidermis, circular to elliptical, minute,

compact, often confluent.

Teleutospores yellowish-brown, clavate to oblong clavate, con-
stricted at septum, rounded or acute at apex and thickened (up to
12 fi), attenuated towards base, smooth, 36-52 x 19-23 /<,
average 48 x 20 ll ; upper cell darker in colour and broader than
lower, 21-31 li long; lower cell slightly or not at all tapering
towards pedicel and averaging same length as upper ; pedicel
persistent, hyaline, but coloured towards apex, 28-37 x 7-8 ^/,
but may attain a breadth of 10 ll at junction with spore.

X. Mesospores not uncommon, similarly coloured to teleutospores,

ovate to elliptical or pear-shaped, thickened at apex, 30-42 x
21-23 ll.

On leaves and all green parts, including fruits of Calendula officinalis L.

Victoria — Near Melbourne, growing in gardens, 1892, etc. Killara,

July-Oct., 1902. Geelong, Nov., 1904, J., 111. (Pescott).
Frankston, August, 1904, I.

Aecidial stage all the year round, but less common in mid-summer,
persisting only in shady gardens. Teleutostage from March to November.

The aecidial stage was the only one found at first, and was described in
the Agricultural Gazetti of New South Wales for 1896 at p. 152. Then

in March, 1902, the teleutostage was found by Mr. G. H. Robinson, and
was very plentiful. Both stages very common in University Gardens,
Melbourne, in autumn and winter of 1905. Mesospores germinating at apex
like teleutospores, and producing sporidiola.

It causes swelling, discoloration and distortion of the flower-stems and
leaves, and the bright orange colour of the aecidia on the leaves at once
attracts attention from its harmonising with the flower-head.

(Plates VIIL, Fig. 65 ; XLIIL, Fig. 312 ; Plate E., Figs. 22, 23.)

77. Puccinia calocephali McAlp.

Calocephahis.

I. Aecidia very sparse on upper surface of hairy leaves, about J mm.

diam.; pseudoperidia saucer-shaped, leciniate at margin; peridial

cells, oblong to lozenge-shaped, striated, 25-32 ll long.

438.

J52

Puccinia — Com-positae.

.showing

to angular, orange-yellow, smooth,
up to 4 germ-pores on one face, 16—19 X 12-13 jx or

Aecidiospores sub-globose

14-16 p diani.
III. Teleutosori black to

blackish-brown, long covered

by leaden

epidermis, pulvinate, compact, round to elongated, sometimes

confluent, 1-1 h mm. long.

Teleutospores clavate, dark brown, particularly in upper cell,

slightly constricted at septum, smooth, 40-57 X 17-24
45 X 22

ju, average

fi ; upper cell rounded, scoop-shaped, sometimes truncate
and thickened at apex (up to 1 1 p) : lower cell yellowish brown,
attenuated towards base ; pedicels persistent, tinted to hyaline,
up to 35 fx long and sometimes 10 fi broad.

X. Mesospores common, elongated, elliptical to oblong, generally paler

than teleutospore, thickened at apex, and generally scoop-shaped,
28-35 x 13-16 /x.

On leaves and stem of Calocephalus drummondii Benth., and C. lactens

Less.

Victoria — Dimboola, Sept., 1891, III. (Reader). Phillip Island,

Nov., 1901, I. (C. French, jun.)

The teleutosori and teleutospores generally resemble those of 1\
tasmanica Diet., but they are not intermixed with aecidia. They both be-
long, however, to the same general type.

(Plate VII., Fig. 58.)

Calotis

78. Puccinia calotidis McAlp.

Aecidia amphigenous, in irregular clusters, orange ; pseudoperidia

cup-shaped, with whitish, reiiexed

Aecidiospores subglobose to polygonal, finely echinulate, 13-14/i

III. Tele

15 x 12 p.

minute, black

umpent, gregarious, compact

Teleutospores dark chestnut-brown, oblong to elongated obi
ivate, smooth, constricted at septum, rounded or bluntly pointed

to 9 ft), 37-50 X 18-28 H, average

and thickened at

ape

38-40 x 18-20 fi; pedicel hy

generally tap

X. Me

wards base, about 30 fi long, or shorter than spore

ly coloured to teleutosp

and

ed, obovoid to ellipsoid, rounded or bluntly pointed and

thickened at apex (6-8

28-3

x 1

Sept.,

1886

(Hamilton)

Ion

On Calotis cuneifolia R. Br. and Calotis sp.

New South Wales — Guntawang,

Condoblin, Aug., 1897 (Maiden).

The teleutospores generally resemble those of P. vittadiniae, but they

(Plate XLIIL, Fig, 310.)

Puccinia — Compo sitae

J53

79# Puccinia chrysanthemi Roze.

Chrysanthemum.

Roze, Bull. Soc. Myc. France, p. 92 (1900).
Sydow, Mon. Ured. L, p. 46 (1902), p. 854 (1904).
Sacc. Syll. XVI., p. 296 (1902).

. P. chrysanthemi chinensis P. Henn. Hedw. XL., p. (26) (1901).

II. Uredosori on both surfaces of leaf, but mostly hypophyllous,

sparingly on stem, generally round, soon naked, powdery, scattered
or crowded, often confluent, sometimes arranged circularly, snuff
brown, average 1-1 £ nim. diam., causing dirty-brown, indefinite
spots on upper surface of leaf, which gradually extend and
ultimately decay.

TJredospores ellipsoid to obovoid, yellowish-brown to golden-
brown, echinulate, with 3 equatorial germ-pores on one face,
28-35 X 22-25 fi (32 n long very common, occasionally reaching
a length of 45 jj) ; pedicels hyaline, rarely persistent, stout, long,
> up to 60 fi.

[III. Teleutosori on both surfaces of leaf, but chiefly on under surface,

prominent, roundish, pulvinate, solitary or confluent in groups, often
concentrically arranged, dark-brown to black, soon naked, compact.
Teleutospores at first sparingly intermixed with uredospores,
ellipsoid to pear- shaped or clavate, rounded or occasionally
truncate at apex, and thickened (up to 9 /u), mostly rounded at base
or sometimes slightly attenuated, chestnut-brown, with strong,
finely punctulate epispore, 35-60 x 19-28 n; pedicel hyaline, stout,
persistent, 40-80 x 7-9 ^u.]

[X. Mesospores rare, ellipsoid to clavate, similarly coloured to teleuto-
spores, rounded and thickened at apex, finely punctulate, 28-46 X
22-25 ft.]

On cultivated Chrysanthemums (Chrysanthemum indicum L.).

New South Wales — Sydney Botanic Gardens, April, 1904

(Maiden). Sydney, May, 1904.

This species has been assigned to P. hieracii by Massee , but Jacky

12 3

» >

has shown, by means of infection experiments, that it confines itself
to the one genus, and does not infect other Compositae. He has also experi-
mented with the Japanese chrysanthemum rust, P. (hrysanthemi-chinensis P.
Henn., and proved it to be the same species as the other.

Through the kindness of Professor Ideta, of the Agricultural College,
Osaka, Japan, I have received abundant material of the Japanese rust, both
in its uredo and teleuto stages, and am thereby enabled to compare it with
the rust occurring in Australia, as far as the uredo stage is concerned. The
uredospore3 are quite similar, having three equatorial germ-pores on one
face, and measuring 28-37 X 22-25 /w. Jacky 3 succeeded in infecting

C. indicum L., with material from C. sinense Sabine, and thus showed
identity of the two rusts, although apparently in Europe and Australia it is
gradually losing the capacity to form teleutospores. Indeed, Miss Gibson '
goes so far as to say that there is no necessity for resting-spores, for the
young shoots are above ground long before the old ones die away, and th
young shoots are taken as cuttings,

time when there are

pon which

F 2

154

Puccinia — Compositae.

The sams author also records the fact that there are certain varieties of
chrysanthemum which do not take the rust, although growing among plants
badly rusted, but Jacky 3 tested a number of different varieties and found
them equally predisposed to the rust.

Our chrysanthemum growers sometimes express a fear that the rust seen
on hollyhock, sunflower, and other plants may pass to the chrysanthemum
when it has suddenly bscome temporarily weak through drought or other
causes ; but it ought to reassure them to know that the specific rust, known
as Puccmii chrysanthemi, is alone capable of producing infection on that
particular host.

In the European material Jacky detected what he considers to be two-
celled uredospores, but Sydow regards them as two unicellular spores stuck
together. It may b3 mentioned here to show the wonderful variety, that in
the Japanese material I found two uredospores borne on the same stalk, the
one slightly beneath the other.

The teleutospore is sometimss described as smooth, but

careful focus-

sing it may be seen to be finely punctulated all over in the young stage,
although in the mature and deeply-coloured spore this is difficult to see, only
being visible on the thickened apex.

The mesospores were obtained from the Japanese material, and are
rather rare.

This rust was probably introduced into Europe from Japan, and it may
have reached Australia either from Europe or Japan,
rust has not yet found its way into Victoria, considering the number of
chrysanthemums imported from England, France, America, and Japan.

It was first observed in England in 1895, although it probably existed
there before that time ; in America in 1896 ; in France and Italy in 1897 ;
and about 1898 it spread to Germany, then to Denmark and Switzerland,

to Australia. In 1904 it was also

It is strange that the

and

at length it has found its

way

recorded for New Zealand by Kirk.

It is found that many of the Australian-raised varieties are less liable to
rust than others when grown in England or America.

(Plate XXIX., Figs. 251-255 j Plate E., Fig. 21.).

80. Puccinia cichorii (DC.) Bell.

Cichorium

Bell., in Kickx. Fl. Fland. II. p. 65 (1867).

Sydow, Mon. Ured. I., p. 49 (1902).

Sacc. Syll. VII., p. 606 (1888) ; XVII., p. 311 (1905)

If. Uredosori amphigenous or on stem, minute, pul

scattered

fluent, girt

or partially covered by ruptured epidermis

cinnamon.

Uredospores globose,

subglobose

ellipsoid, echinulate,

yellowish-brown, with one or two germ-pores on one face, sometimes

three to four, 22-29 x 19-23 fi
III. Teleutosori similar, dark-brown.

or 22-25 f± diam.

Teleutospores intermixed sparingly with uredospores, ellipsoid
or ovate-ellipsoid, brown to golden-brown, smooth, not or hardly
constricted at septum, apex rounded not thickened, base for the

thin,
contents,

27-38 X 19-25 /t, occasionally reaching 50 \x long, but averaging
35 x 20 p. ; pedicel hyaline, short.

most part rounded or very slightly attenuated, epispore
germ-pore in each cell prominent, with finely

granular

Pvccinia — Compositae. 155

On leaves and stems of Cichorium intybus L.

Victoria — 1885 (Mueller 3 ) II. Near Melbourne, Aug., 1896.

Phillip Island, Jan., 1900; March, 1904.

The uredospores were common chiefly on the leaves, but the teleutospores
vyere rather scarce, and occurred mostly on the midribs. Fischer5 describes
the teleutospores as being finely warted, while Sydow considers them to be
smooth. When the uredo and teleutospores are examined together, as they
often occur in nature, the former are seen to be covered with decided
prickles, which project from the margin, while the latter have fine points on
the surface, which are only to be seen at the margin by careful focussing.

Although no mesospores were met with there were several cases where
the upper cell of the teleutospores was colourless, shrivelled, and much
reduced in size, while the lower cell was enlarged. The two cells are
generally equal in length, but in one of the cases referred to the lower cell
was 25 fx long, and the upper colourless one only 6 li.

Darluca filum Cast., was plentiful on the uredosori and the sori

£3

uredo and teleutospores intermixed

(Plate VIII., Fig. 61 ; Plate D., Fig. 18.)

•

Cineraria

81. Puccinia cinerariae McAlp.

I. Aecidia on both surfaces of the leaf, running along veins and gre-

garious, causing depressions on upper surface and swelling of
the veins on under surface, ^ mm. diam. ; pseudoperidia very
slightly projecting, with margins white, everted, fimbriate ; peri-
dial cells roughly quadrangular, often rounded at one end, longer
than broad, with radiating marginal lines, 22-26 X 19-21 fi.

Aecidiospores bright orange, subglobose to polygonal, very
finely verrucose, 15-19 li diam. or 15-17 x 13 /*.

III. Teleutosori intermixed with aecidia, black, comparatively rare.

Teleutospores clavate, chestnut-brown, generally rounded and
much thickened at apex or bluntly pointed, occasionally truncate,
constricted at septum, 36-54 x 17-24 la, average 40 x 23 fx ;
upper cell usually darker in colour, hemispherical, apex thickened
to a depth of 9 li, 17-24 li long; lower cell gradually tapering
towards base, somewhat triangular, 1 8-30 li long ; pedicel per-
sistent, tinted, up to 45 li long.

X. Mesospores chestnut-brown, obovate to elongated elliptical, much

thickened at apex, with long pedicel, 27-36 x 13-23 fx.

On leaves of cultivated Cineraria.

Victoria — In nursery near Melbourne, April, 1899 (Cronin).

The following note accompanied the specimen : — The plants were grown

under glass, and were generally very healthy. There are not many leaves

••so affected owing to the care that is taken to cut out the spots as soon as

I noticed. The disease spreads rapidly if neglected, and completely spoils a

j plant by denuding it of its foliage.

Caeoma cinerariae Rostr.. was found on leaves and stems in Jutland.

(Plate VIII., Fig. 60.)

i56

Puccini a — Compo sitae.

Centaurea*

82. Puccinia cyani (Sehleich.) P

Passerini in Rabh. Fung. Eur., N

P-n*

176

Sydow, Mon. Ured. I., p. 38 (1902).

Sacc. Syll. VIL, p. 634 (1888) ; XVII., p. 286 (1905)

II. Uredosori mostly hypophyll

ed or

ded, minute

orbicular or elliptic, pulverulent, reddish br

Uredosp

yellowish b

lobose, subglobose to

finely cchinulate, with two

por

on one face

7 x

III. Teleut

ampl

and on stem, scattered, pulverulent

dark-brown, orbicular to elliptic, soon naked

1 mm

Teleutospor

caixed

edospor

chestnut-b

ellipsoid, rounded at both end
stricted at septum, delicately
30-40 x 22-30 /i, average 32 x
pedicel hyaline, short,

thickened at apex, not

ted

erm-pores conspicuou

decid

24 fx ; epispore about 3 /x thick

On leaves and stems of Centaur ecu cyanus L.

Victoria — Near Melbourne, Jan. and Feb., 1904 (C. French, jr.).

This is evidently of the Puccinia hieracii type.

ular and well defined, and there is no

The

teleutospores are very reg

tendency to 'abnormality.

(Plate VIII., Fig. 64.)

Bellis.

83. Puccinia distincta McAlp.

McAlpine, Agr. Gaz. N.S.W. VI., p. 853 (1895).
Sydow, Mon. Ured. I., p. 29 (1902), and p. 850 (1904)

Sacc. Syll. XIV., p. 311 (1899).
I. Aecidia seated on discoloured spots, amphigenous,

arranged in a

circinate manner, or irregularly disposed, and often distributed
over entire surface ; pseudoperidia cup-shaped, with reflexed torn

mm. diam. ; peridial cells elongated elliptical to

margins,

about

l i

4"2

oblong, punctulate all over, 25-29 x 13-19 \i

Aecidiospores

subglobose

to oval or ellipsoid, orange-yellow,

very finely echinulate, 14-19 p diam , or 14-20 x 13-16 \x

III. Teleutosori intermixed with and often

brownish black, oval, ^-1

surrounding

aecidia,

mm. long, generally confluent, bursting

through and surrounded by the leaden-coloured epidermis.

Teleutospores chestnut-brown, clavate or

oblong clavate, at-

tenuated at base, smooth, constricted at septum, 34-50 X

15-21 H

average

42 x 17 fi ; upper cell generally darker in

colour than lower, rounded and thickened at apex (up to 9 ft) ;
pedicel persistent, pale yellow or coloured similarly to lower cell, up

to 50 /x long.
X. Mesospores numerous, elongated

to somewhat ovate, chest

brown, usually thickened at apex, 34-37 X

3-16

On leaves, scapes, involucre and petals of Bellis perennis L

Victoria—

oria— Near Melbourne, Oct., 1892, July-Oct., 1904, I., Ill
(Robinson). Korumburra, May, 1903, I.

Puccinia — Compositae

157

Tasmania — Near Huon Road, Nov., 1890, I. (Rodway).
South Australia — Norwood, April, 1903, I., III. (Tepper). Port

Pirie, Oct., 1903, J., III. (Dickens).

Aecidiospores abundant from April to September, and even to January.
The aecidia may even occur on the petals, and they have been described
before the Microscopical Society of Victoria as early as 1880.

Teleutospores in October or November, although they occur sparingly as
early as April, The spots are generally pale-yellowish to pale greenish in
colour. This aecidium from Victoria was determined by Cooke as A. bellidis
Thuem., which belongs to P. obscura Schroet., but now that the teleutospores
belonging to it have' been found on the same host-plant the Puccinia is seen
to be a distinct species.

(Plate VIIL, Fig. 67 ; Plate E., Fig. 26.)

Erechtites.

■

S4. Puccinia erechtitis McAlp.

McAlpine, Proc. Roy. Soc. Vic. VII., N.S., p. 216 (1894).
Sydow, Mon. Ured. I., p. 78 (1902).
Sacc. Syll. XIV., p. 309 (1899).

0. Spermogonia on brownish patches on upper surface of leaf, in groups,

brown to honey-colour, yellowish by transmitted light.

Spermatia hyaline, oval, 3 \x long.

1. Aecidia on stem and leaves, causing distortion and swelling, pale-
yellow at first, becoming orange-yellow, arranged close together in
lines or irregularly ; pseudoperidia cup-shaped, with white, torn,
revolute edges ; peridial cells firmly united, warted all over,
lozenge-shaped, and nearly iso-diametric, or elongated and oblong,
with striated margins.

Aecidiospores variable in shape, irregularly globose or elliptic

orange-yellow, smooth, 16-19 x 12-1/ jx.

III. Teleutosori long covered by epidermis, crowded together,

pulvinate, black.

Teleutospores clavate to oblong, yellowish-brown, constricted
at septum, smooth, 41-57 x 17-25 li, average 47 x 19 li ; upper
cell dark-brown, rounded or pointed, occasionally truncate and
thickened at apex ; lower cell usually yellowish-brown and tapering
towards base, elongated wedge-shaped ; pedicel persistent, pale-
yellow or hyaline, up to 40 li long and 9 li broad.

X. Mesospores common, coloured like teleutospores, elongated oblong

or oval, thickened at apex, 32-43 X 13-17 yu.
On leaves and stems of Erechtites quadridentata DC, and E. arguta DC.

Victoria — Ardmona, Dimboola, Myrniong, and Marysville, Alps

near Bright, and Mt. St. Bernard, etc.

On E. quadridentata, E. arguta and E. prendnthoides DC.

Tasmania— Huon Road, Dec, 1891, March, 1893, I. (Rodway).

■

On Erechtites sp.

New South Wales — Guntawang, I. (Hamilton).

I I. Very common all the year round, except during middle of summer.

| Til. From January to June, but not so common.

The New South Wales specimen was named Aecidium senecionis Desm.

by Cooke.

(Plates VII., Fig. 59 ; XXVII., Fig. 240.)

158

Puccinia — Compo sitae.

85. Puccinia gnaphalii (Speg.) P. Henn

Hennings, Hedw. XLI., p. (66) (1902).
Morrison, Victorian Nat. XI., p. 120 (1894). .
Sydow, Mon. Ured. I., p. 88 (1902).
Sacc. Syll. XVI., p. 295 (1902).

Uredo gnaphalii Speg. Fung. Arg. IV., p. 28 (1882)

Gnaphaliurru.

Puccinia gnaphaliicola P. Henn. Hedw. XXXVIII., p. (68)

(1899).

II. Uredo-sori common on stem and leaves, on the latt<

but mostly on under surface, bursting through and

epidermis, also pier

lly torn en turn

l* amphigenous,.
surrounded by
ed, sometimes

gated, compact, cinnamon, circular or oval, lenticular on

stem

-\ mm. diam.
Uredospores globose

sub globose

echinulate, 21-24 a diam. or 21-26 x 17

lipsoid
1«.

bro

finely

III. Teleutosori similar, but chestnut-brown to black

Teleutosp

at first

ed with uredospores, oblon

oblong-clavate, dark chestnut-brown, slightly constricted at septum

rounded or obtusely pointed or even beaked at apex

thickened (up to 8 /*),
smooth, 35-53 x 15-24

lly slightly attenuated towards base

persistent, up to 66

ge 48

x 20

pedicel hyal

Thunb.

X. Mesospores similarly coloured, few, oval or elongated, elliptical^

thickened at apex and sometimes beaked, 24-38 x 15—18 yu.
On stems and leaves of Gnaphalium purpureum L., and G. japonicum

Victoria

Oakleigh, Jan., 1904 (Morrison). Drysdale, Jan., 1896,.
and Oct., 1903. Whittlesea Ranges, Nov., 1898 (C. French,,
jun.) Killara, Oct., 1902. Near Melbourne, Feb. and Nov.,.
1904. Murramurrangbong Ranges, Nov., 1904 (Aitken).

The teleutospores are rather variable in length and breadth, sometimes

being short and stout and sometimes rather elongated and slender.

The

pedicels average 40-50 /j, in length, and may vary in breadth from 4 /j. up to
10 /* at junction with spore. Sydow gives size of teleutospores as 32-56 x
18-24^, and Dietel as 30-35 x 19-22 ju.

from

I am indebted to Dr. Morrison for some of the original material
which he determined this fungus as P. investita Schwein. He found sper-
mogones immersed among spores in the sori, and the spermatia were fusi-
form. Plenty of Darluca filum Cast., was found, and no doubt these were

mistaken for spermogonia.

(Plate VII., Fig. 57.)

86. Puccinia helianthi Schwein.

Jlelianthus^

Schweinitz, Syn. Carol., p. 73 (1821).

Cooke, Handb. Austr. Fung., p. 333 (1892).
Sydow, Mon. Ured. I., p. 92 (1902).
Sacc. Syll. VII., p. 603 (1888).

Spermogonia honey- colored, in small clusters.] I

[I. Aecidia crowded or orbicular, or arranged in broadly expanded,,

oblong \spots ; pseudoperidia cylindrical, plane, with white laciniatft

margins.

Puccinia — Compositae. 150

Aecidiospores orange-red, ellipsoid to polygonal, finely echin-

ulate, 21-28 x 18-21/*.]

II. Uredosori roundish, chestnut-brown, scattered or confluent, pul-

verulent, often on yellow or pale green spots on upper surface of
leaf but generally forming brown mass on under surface.

Uredospores sub-globose, elliptic or obovate, golden-yellow, echin-
ulate, epispore sub-hyaline, showing one germ-pore on one face.
21-24 yu diam. or 24-29 x 15-22 F.

III. Teleutosori roundish, dark -brown or black, prominent, scattered at

first but ultimately in clusters, confluent, dotting both surfaces of
leaf, but most prominent on under.

Teleutospores at first intermixed with uredospores, chestnut-
brown, oblong-elliptical or pear-shaped, smooth, but occasionally a
little rough at apex, slightly constricted at septum, thickened at

apex (6-9 /*), generally rounded at base, 36-50 x 21-27 yu,
average 44 x 26 jx ; upper cell similar in colour or just a
shade darker than lower, and rather larger, 22-29 x 21-27 /u;

lower cell either the counterfeit of upper or slightly tapering
towards base, 17-23 X 20-22 fi; pedicel hyaline, persistent,
generally much longer than spore, up to 90 and 110 yu long.

On leaves, branches, involucral bracts and corolla-leaves of Helianthus
<innuus L., and H. tuber osus L.

Victoria — Very common, December- April.

New South Wales — Mudgee and Guntawang, Feb., 1887 (National

Herb.) (Hamilton). Sydney Botanic Gardens, Feb., 1901.
Queensland — Ipswich, 1888, Bailey5; Brisbane, 1889.

This rust was first observed in South Carolina and Pennsylvania, then it
appeared in Russia where the sunflower is largely cultivated, and soon it
spread over Europe, extending to Australia. The teleutospores are very
regular and definite in shape.

Aecidia have not been found in Australia, although the rust is plentiful.

Sydow in his Monograph comes to the conclusion that this species
possesses no aecidial stage, since he has examined specimens from numerous
localities without result, but Carleton3 has collected the three stages in
America and remarks : — u The aecidium occurs rarely in comparison with the
occurence of other stages, but it is to be found on a number of hosts and
occasionally in considerable abundance. This rarity of its occurrence,
together with the occurrence of spermogonia so often with the uredo, may
be accounted for by the fact that the uredo is often produced by direct
teleutosporic infection."

According to Woronin1 the teleutospores germinate equally well
whether they have been kept dry in a room or taken from leaves which had
lain under the snow throughout the winter. Carleton2 says that they also
I germinate at once without a resting period.

(Plate VII., Fig. 56.)

Hyvoch

37. Puccinia hypochoeridis Oud.

Oudemans in Nederl. Kruidk. Arch. II., Ser. 1, p. 175 (1873)

Sydow, Mon. Ured. I., p. 100 (1902).

Sacc. Syll. VII., p 634 (1888); XVII., p. 302 (1905).

Puccinia microseris McAlp., Agr. Gaz. N.S.W. VI., p

757 (1895).

i6o

Puccinia — Com-po sitae.

II.

Uredosori amphigenous or frequently on stem, genera ^

llv seated

minute spots, scattered, pulverulent

Uredospores globose, sub-globose or ellipsoid, echinul

brown, with one prominent germ-por

face

diam

0-28 x 1

III. Teleutosor

punctiforn

mphigenous,
blackish to

often on stem, scattered or crowded

k-b

pul

dish

leaf

generally

to 1 mm. diam., but often smaller

d on stem and up to

mm. long.

Teleutosp

at fir

ed with uredospor

oblo

ellipsoid or obovate, apex

Uy rounded, and not thickened

rounded at base or
septum, dark-brow n,
cellular, 30-49 x 17

htly

uated, hardly

stricted at

ery delicately punct

occasionally tri-

,ge 3

x 20 fx ; pedicel hyaline,

lly deciduous, sometimes persistent and may

reach a length

of 77 ft.

On Hypochoeris radicata L

Victor

Dimbool

Nov

(Robinson). My

L889 (Reader). Ardmona
March, 1900 (Brittlebank).

1894

Nea

Melbourne, Apr., 1901. Dookie, Oct., 1903. Altona Bay
Oct.. ]

X. w

903 (C. French, jun.). Murramurrangbong Ran
Dec, 1903 (Robinson). , . -

New South Wales — Wagga (Maiden).

On H. glabra L.

Victoria— Dimboola, Nov., 1889, and July, 1892 (Reader).

Queensland — Brisbane, 1886 (Bailey 5).

The punctation on the teleutospores is so fine that it may easily be over-
looked, but when the spores are mounted in water it is clearly seen.

A specimen from Syd. Ured. Exs. 673 on II. glabra agreed very closely,

25 x 21 ju, and the teleutospores

the uredospores being

on an

average

34 x 18 ft.

This species closely resembles in morphological characters P. hieracd
(Schum.) Mart., but Jacky has shown by means of cultures that the latter
can only grow on Hieracium species, and cannot be transferred to other
genera of Composites. Occasionally there may be a three-celled teleutospore
reaching a length of 56 /x. \

P. microseris described as a new species in the Agricultural Gazette
New South Wales was based upon a wrong determination of the host-plan
by the collector since it was afterwards found to be Hypochoeris radicata L.

Darluca filum Cast., is sometimes common on the mixed uredo and
teleutosori.

(Plate VIII., Figs. 62, 63.)

88. Puccinia kalchbrenneri I)e Toni.

Ilelichrysum

De Toni in Sacc. Syll. VII., p. 645 (1888)
Cooke, Handb. Austr. Fung., p. 337 (189
Sydow, Mon. Ured. I., p. 93 (190

II. Uredosori on both surfaces, but

tly

under, seated on in

determinate spots which are often confluent and variously coloured
scattered or gregarious,

minu

at first covered, convex,

fir

imately free and discoid

Puccinia — Compositae.

161

Uredospores globose, subglobose or ellipsoid, warted, yellow to
pale brown, 20-30 x 19-26 ju.

III. Teleutosori hypophyllous, seated on the same spots, minute, brown.

Teleutospores oblong or subclavate to lanceolate, apex thickened

attenuated or rarely truncate, constricted at sept
dark-brown, 40-57 x 15-22 /n, rarely up to 27 /x broad; pedicel
hyaline, short, deciduous.

On living leaves of Helichrysum sp.

Victoria.

There is no specimen of this species from Australia in the Kew Her-
barium, and I have not found it on any species of Helichrysum, though this
genus is exceedingly common near Melbourne.

89. Puccinia lagenophorae Cooke.

Cooke, Grev. XIIL, p. 6 (1884).

Cooke, Handb. Austr. Fung., p. 335 (1892).

Lagenophora.

■

Sydow, Mon. Ured., L, p. Ill (1902), and p. 863 (1904).
. Sacc. Syll. VII., p. 612 (1888).

P. hypochoeridis McAlp., Proe. Roy. Soc. Yie. VII., N.S.,

p. 217 (1894).

P. macalpini Sydow, Mon. Ured., L, p. 100 (1902).

I. Aecidia amphigenous, spots none ; pseudoperidia scattered or in

groups, semi-immersed, margin lacerated, white, 200-240 fx diam.;
peridial cells finely warted, lozenge shaped or elongated, 24-31 p
long.

Aecidiospores subglobose or elliptical, orange-yellow, very
finely echinulate, 12-14 p diam. or 17-19 x 14//.

[II. Uredosori scattered, small, pulverulent, brown or mixed with

teleutosori.

Uredospores globose, brown, epispore rough, 20 ft diam.]

III. Teleutosori amphigenous, scattered, or surrounding aecidia,

rather compact, dark-brown to black, raising and rupturing
epidermis.

Teleutospores clavate, constricted at septum, dark-brown only

in upper cell, epispore thickened, smooth, 45-66 X 16-22 p,

J average 49 x 1 7 jx ; upper cell subglobose or oblong or somewhat

conical, rounded, flattened, or bluntly pointed at apex, which is
considerably thickened, generally equal in length to lower cell ;
lower cell pale in colour, attenuated into pedicel and narrower
than upper cell ; pedicel persistent, generally slightly tinted,
attaining a size of 42 x 7 p.

X. Mesospores relatively numerous, pale coloured, elongated elliptical,

thickened at apex, with persistent hyaline pedicel, 38-42 x

16-17 p.

On stems and leaves of Lagenophora billardieri Cass.

Victoria — Omeo, 1884 (Stirling). Ardmona, Oct., 1894 (Robin-
son). Near Melbourne, Sept., 1900, I. (C. French, jun.)
Murramurrangbong Ranges, Nov., 1902, Dec., 1903, Jan.,

1905 I., III. (Robinson).

162

Puccinia — Com-po sitae. ,

On L. huegelii Benth., and L. billardieri Cass

Tasmania — Mt. Dromedary

way

Mt. Direction

Oct

Hoba

189

Dec
I..

Ill

89-1, I. (Rod

(Rod way1)

Devonport, Jan., 1906 (Robinson)

Although material was very plentiful, careful search failed

eal

have given

presence of uredospores, and I

Cooke. Massee informs me that P

sented in the Kew Herbarium, and he, therefor

tit

the

>ph

Cool

specimens,
from diffe
uredosp

I have

ned

of plants with th

localities and at different

authority of
is not repre-
refer to the type
5 rust upon them
and I have never found

Trie leaves on which P. hypochoeridis McAlp

found proved to be-

not Hypochoeris
agreed with the above

radicata

but Lagenophora billardieri, and the fung

(Plate VII., Figs. 54, 55

Plate F., Fi<?. 27.)

90. Puccinia podolepidis McAlp.

Podolepis*

0. Spermogonia on small, honey-coloured spots, forming minute, dark-

coloured points, hemispherical, pale yellow by transmitted light,,
with round mouth, without projecting paraphyses, about 1 20 p diam.
Spermatia minute, hyaline, oval, 3 X 2 ju.

1. Aecidia bright orange, on both surfaces of the leaf, numerous

generally

arranged

in circular groups, which, however, often
coalesce into large patches ; pseudoperidia cup-shaped, embedded in
tissue, with white reflexed, laciniate margins ; peridial cells oblong
to lozenge shaped, with striated margins, 28-32 x 16-18 p.

Aecidiospores deep orange, ellipsoid to subglobose, smooth,
24-32 x 16-22 it, or 22-24 p diam.

III. Teleutosori black, surrounding the aecidia.

Teleutospores chestnut-brown, oblong to clavate, constricted at
septum, smooth, rounded or truncate or occasionally pointed and
thickened at apex (9-12 u), rounded or attenuated towards base,.

upper cell darker than lower, 38-54 X 22-32 p. ; pedicel per-
sistent, hyaline, relatively short.
X. Mesospores common, similarly coloured to teleutospores or often

paler, ellipsoid, obovate to wedge-shaped or subclavate, rounded and
thickened at apex, pedicellate, 32-44 X 16-22 p.
On leaves of Podolepis longipedata A. Cunn.

Victoria— Buffalo Mts., Nov., 1903, I. (C. French, jun.).

near Bright, Dec, 1904, I., III. (C. French, jun.).

Several teleutospores had

Alps,.

side, but only one in lower cell.

two germ

pores in upper cell, one on either

The teleutosori were rather sparse

(Plate XXIX., Figs. 256, 257.)

Lactuca

91. Puccinia prenanthis (Pers.) Lindr.

Lindroth, Myk. Mittheil, p. 6 (1901).
Cooke, Handb. Austr. Fung., p. 334 (1892)
Sydow, Mon. Ured. I., p. 106 (1902).
Sacc. Syll. XVII., p. 306 (1905).

Puccinia — Comfo sitae*

163

I. Aecidia hypophyllous, a few rarely epiphyllous, or on petioles, seatec

on orbicular or elongated yellow or purple spots, up to 1 cm. ir
diam., in round or irregular groups, at first hemispherical, thei
flat, yellow or whitish, sometimes yellowish purple.

Aecidiospores globose, subglobose or ellipsoid, delicately warted
pale orange, 13-24 f.i diam.

II. TIredosori hypophyllous, on pale indeterminate irregular spots

scattered, minute, punctiform, pulverulent, pale brown.

Uredospores globose or subglobose, echinulate, yellowish-brown
16-24 u diam.

III. Teleutosori similar, girt by the ruptured epidermis, dark-brown.
' Teleutospores ellipsoid, rounded but not thickened at apex,

mostly rounded at base, not constricted at septum, delicately
warted, brown, 26-36 x 16-24 fx ; pedicels hyaline, very short.

On living leaves of Lactuca.

Victoria. New South Wales. S. Aust

No specimen seen. It is given on the authority of Cooke, but Mueller3
only records the aecidial stage for Victoria. The aecidial wall is very im-
perfectly formed in this species, and there is a close approach to caeoma
forms.

92. Puccinia tasmanica Diet

Senecio

Dietel, Ann. Myc. I., p. 535 (1903).
Sydow, Mon. Tired. L, p. 867 (1904).
Sacc. Syll. XVIL, p. 277 (1905).

I. Aecidia on blister-like swellings on stem and branches, on upper and

under surface of leaves, on flower-head stalks and involucre, causing
discoloration and distortion, and usually surrounded by paler green
tissue, about ^ mm. in diam., disposed in large circular or oval
clusters or irregularly ; pseudoperidia cup shaped, with white,
irregularly laciniated everted edges, tubercular before opening ;
peridial cells firmly united, overlapping each other, with striated
margins and usually broader at one end, individually slipper-
shaped, 25-35 x 13-17 fi.

Aecidiospores spherical or angular, orange colored, very delicately
warted, 13-16 /u diam., or 14-17 x 11-15 p.

III. Teleutosori for a long time covered by epidermis, then erumpent

and epidermis usually thrown off or remaining in shreds and
patches, intermixed or running parallel with aecidia, black, pulvi-
I nate, oval, up to 1 mm. long, often confluent in elongated lines.

Teleutospores oblong to clavate, chestnut-brown, slightly con-
stricted at septum, smooth, 36-63 X 15-25 y, average 54 x 20 //,
occasionally tricellular when 48-73 fj. long ; upper cell deep chest-
nut-brown, rounded or somewhat oval, conoid, or truncate and
thickened at apex (up to 13 yu) ; lower cell usually paler in colour,
rounded at base or tapering, often elongated relatively to upper ;
pedicels persistent, pale yellow to hyaline, sometimes longer than
spore.

X. Mesospores very common, elongated oval, oblong, or somewhat

elliptical, apex rounded or pointed and usually thickened, golden-
yellow to chestnut-brown, 29-44 x 13-17 /j.

164

Puccini a — Compo sitae.

On leaves and stems of Senecio vulgaris L.

Victoria (Ralph), (Robinson), (French, junr.), &c, I. III.
New South Wales (Hamilton), I.
* Tasmania— 1891, 1893, 1895, 1897, I. III. (Rodway >).

On Senecio pectinatus DC.

New South Wales — Mount Kosciusko, Jan., 1898, I. (Maiden).

On Senecio brachyglossas F.v.M.

Victoria — August, 1900, I. (Reader).

On Senecio velleioides, A. Cunn.

Victoria— Sealers' Cove, 1854, I. Port Phillip, 1886, I.

It differs from P. senecionis Lib., in the aecidia being amphigenous and
not hypophyllous ; also in the teleutospores being considerably larger,
thickened at apex, and not provided with a hyaline papilla, while the pedicel
is decidedly persistent and elongated. Groundsel, or S. vulgaris, being an
imported plant, and so cosmopolitan in its character, it was considered
strange that a new species of rust should be found upon it, and although I

had named it in MS. P. australiensis, yet I delayed publishing it under

that name, hoping bo find the teleutospores on a native Senecio.

Teleutospores are very common in some localities during the winter and

spring months, and the Groundsel growing on the coast at Port Fairy was

so overrun with both aecidia and teleutosori that the plants were stunted

in growth.

S. pectinatus DC, occurs in the three States from which Puccinia

tasmanica has been recorded, but only the aecidia! stage has been met with

in New South Wales.

(Plate VII., Fig. 52.)

93. Puccinia vittadiniae McAlp.

Vittadinia

I. Aecidia on both surfaces of leaf, but mostly on upper, scattered or in

groups and confluent, minute ; pseudoperidia white becoming
yellowish, immersed, with lobed margin ; peridial cells with striated
margin in section, striae projecting and appearing as points in sur-
face view, oblong, 25-32 x 13-16 p.

\ ^]Aecidiospores yellowish, ellipsoid, very finely verrucose, 14-17 x

11-13 /Li.

III. Teleutosori minute, black, sparsely developed, intermixed with

aecidia.

Teleutospores dark chestnut-brown, oblong to elongated oblong or
clavate, smooth, constricted at septum, generally rounded at base
and apex, sometimes bluntly pointed or truncate, and thickened at

average

34 X 21 fx ; upper

cell

apex (6-9 /u), 31-46 x 18-25 /u,
darker than lower and similar or sometimes considerably broader ;
pedicel hyaline, persistent, 30-40 /x long, and up to 9 fx broad
adjoining spore.

X, Mesospores very common, similarly coloured to teleutospores and

stalked, ovoid to elongated ellipsoid, rounded or beaked at apex and
thickened (up to 9 /u), 25-34 x 14-17 /u.

Puccinia — Rubiaceae.

'«5

On leaves including leaf stalks of Vittadinia australis A. Rich.

Victoria — Dimboola, June, 1900 (Reader).

In Agr. Gaz. N.S.W. VI., p. 757 (1895), I described a new species under
the name of Aecidium vittadiniae upon a plant forwarded to me as Vittadinia
australis, but which was afterwards found to be an Erechtites, so that the
present description replaces the previous one.

(Plates XXXIX., Fig. 292 ; XLIIL, Figs. 308, 309.)

RUBIACEAE.

94. Puccinia coprosmae Cooke.

Coprosma

Cooke, Grev. XIX., p. 2 (1890).

Sydow, Mon. Ured. I., p. 209 (1902).
Sacc. Syll. IX., p. 300 (1891).

P. coprosmatis Morrison, Vict. Nat. XL, p. 90 (1894).

III. Telcutosori hypophyllous, sometimes epiphyllous, rusty-brown,

prominent, compact, round or elliptic, usually in groups, sometimes
forming mammilated tubercles, rarely solitary, densely crowded
and confluent, sometimes surrounded by the ruptured epidermis
or naked and deforming the leaves.

Teleutospores elongated oblong to clavate, pale yellowish to
brownish, the two cells about equal and more or less ovate, apex
bluntly pointed or rounded, thickened, (from Q-8 //), rounded or
slightly tapering at base, smooth, constricted at septum,
35-51 X 16-22 fj, average 43 x 18^/; pedicel hyaline, persistent,
elongated, up to 110 fx long and 8 li broad.

X. Mesospores common, similarly coloured to teleutospores, ovoid to

ellipsoid, or somewhat fusoid, usually pointed and thickened at
apex, 32-41 x 19-22 /u.

On leaves of Coprosma billardieri Hook.

Victoria — Kew and Dandenong, Oct. and Feb., 1893.
Tasmania — (Rod way ' ).

On Coprosma hirtella Labill.

Victoria — Murramurrangbong Ranges, Dec. 1903, Jan. 1905

(Robinson).

I am indebted to Dr. Morrison for some of the original material on
which he founded his new species P. coprosmatis, but it is identical with
the above species first determined by Dr. Cooke on Coprosma lucida from
New Zealand.

Darlucafilum Cast., was very plentiful on the teleutosori on Cojyrosma
billardieri, and was described by Morrison as a spermogone with spermatia.

(Plate X., Fig. 78.)

95. Puccinia oliganthae McAlp.

A sperula

II. Uredosori hypophyllous, cinnamon-brown, round to ellipsoid, soon

naked, girt by the ruptured epidermis, confluent into elongated
patches.

i66

Puccinia — Rubiaceae.

V redospores globose to shortly elliptical or obovate, golden-
brown, finely echinulate, one to two germ-pores on one face,
25-29 X 22-25 li, or 25-27 li diam.

III. Teleutosori on stems, dark-brown to black, elongated, bullate,

soon naked, compact, 2 mm. long or more.

Teleutospores narrowly clavate to oblong, brown, constricted at
septum, thickened at apex (9-10 li), and rounded or bluntly pointed

ay be

lly attenuated towards base

and paler in colour, 32-54 x 16-21 li, average 48-16 li ; pedicel
yellowish, persistent, up to 48 li long.

X. Mesospores occasional, similarly coloured to teleutospores, somewhat

fusiform, with pointed and thickened apex, 28-32 x 12-13 li.

On stem and leaves of Asperula oligantha F. v. M.

Victoria — In shady gullies of Murramurrangbong Ranges, Nov.,

1902 (Robinson).

The examination of specimens of P. punctata Link, on species of Asperula
from Exsicc. Sydow Ured. 465, 466, showed that the two were distinct. In
P. oligantha the uredospores are broader, and while the teleutospores
average only about 16-18 li broad, in P. punctata they are about 20-22 /<.
In the latter too the apex is much thicker, reaching up to 16 li. The
species closely resembles P. asperulae odoratae Wurth, but aecidia are
present on the latter, and the teleutospores are only thickened at the apex

the extent of 7 u

(Plate X., Fig. 7

96. Puccinia operculariae (Morr.) Syd.

Opercularia.

Morrison, Victorian Nat. XI., p. 119 (1894).

Sydow, Mon. Ured. L, p. 224 (1902).

Sacc. Syll. XIV., p. 321 (1899) ; XVII., p. 317 (1905).

Aecidium cystoseiroides Berkeley, Fl. Tasm., p. 270 (1860).

I. Pustulate, deforming the leaves ; pseudoperidia immersed.

Aecidiospores orange, subangular.

III. Teleutosori hypophyllous, on orbicular brownish to yellowish spots,

solitary or a few together, round or elliptic, compact, reddish-
brown, girt or partially covered by the ruptured epidermis, up to
2 mm. long.

Teleutospores golden-bi own, oblong to oblong clavate, constricted
at septum, smooth, 35-54 x 14-20 /u; upper cell rounded or
pointed and thickened at apex (up to 11 li), about equal in length
to lower ; lower cell usually tapering towards base or sometimes
rounded ; pedicel hyaline, persistent, 80 li or more long and 6-7 ii
broad .

On living leaves of Opercularia aspera Gaertn.

Victoria — Genoa River (F. V. Mueller).

On leaves and petioles of Opercularia varia Hook. f.

Victoria — Oakleigh, Nov., 1893 (Morrison).

Tasmania— Near Hobart Rivulet, Nov., 1860, I. (Gunn), 1902

(Rod way1).

Puccinia — Loranthaceae.

167

Morrison states that this fungus deforms the leaf and produces a con-
cavity on the opposite side. He made this a variety of P. coprosmae Cooke,
and certainly there is a very close resemblance, but the sori in the latter are
generally in groups, and the apex of the spore is not quite so thick.

No specimens of the aecidial stage were seen, but being on the same
host-plant as the teleutospores, it is included in this species. v

Darluca filum Cast., is common on the teleutosori, although it is usually

nly found on ured

(Plate XXIX., Fig. 258

LORANTHACEAE.

Loranth

97. Puccinia loranthicola McAlp

I. Aecidia imbedded in raised brownish to dark brownish, often con-
fluent cushions on one or both surfaces of leaf, orange ; pseudo-
peridia tubular, with white, reflexed, much torn margins ; peridial
cells oblong, with striated margins, 43-50 X 27-31 a.

Aecidiospores ellipsoid to oblong or sub-angular, b

-,"*«,*, »x*gwu viaug

yellow, decidedly echinulate, 37-56 x 22-31 li

II., III. Uredosori amphigenous, brownish, bullate, in scattered

groups, often arranged in circles and confluent, surrounding
central darker teleutosori, epidermis splitting lengthwise and
persistent.

II. Uredospores orange yellow, ellipsoid to oval, obovate or oblong,

coarsely echinulate, with 3-5 equatorial germ-pores on one face,
40-65 x 22-32/7, occasionally 80 x 22/*; epispore 3-4J li thick.

III. Teleutospores intermixed with uredospores, subhyaline, cylindrical

to clavate cylindrical, smooth, slightly constricted at septum,
rounded and slightly or not at all thickened at apex, tapering
slightly towards base, 65-94 x 15—24 yu, occasionally 3-celled,
when about 86 fx long ; pedicel hyaline, persistent, elongated up to
200 Li long, and swollen towards apex up to 14 fi.

X. Mesospores subhyaline and with elongated pedicel like that of

teleutospore, smooth, fusoid, hardly thickened at apex, 71-77 x

25-28 ft.

On living leaves of Loranthus celastroides Sieb., growing on Stringy bark
{Eucalyptus sp.).

Victoria — Murramurrangbong Ranges, Jan., 1905 (Robinson).

All the stages were found on the same tree, but the aecidia on separate
leaves. All the spore-forms are particularly large, a fact which probably has
some relation to the peculiar nutrition of the fungus, and the height at which
it occurs, some specimens being obtained fully 50 feet from the surface of
the ground.

The aecidia are very conspicuous on raised cushions, with corresponding
depressions on the opposite side, where aecidia may also occur.

The uredosori are of a ruddy brown, in isolated groups, generally
arranged circularly and with teleutosori in the centre. Puccinia loranthi
Speg., has only teleutospores, and it is doubtful if they are to be regarded
as such. P. macrocarya Rac. on leaves of Loranthus in Java has only aecidia

and teleutospores.

(Plate XXXI., Figs. 268-271.)

i68

Puccinia — Umbelli ferae .

UMBELLIFERAE

98. Puccinia thuemeni (Thuemen) McAlp.

P. castagnei Thuemen, Rev. Myc. II., p. 86 (1880).

Apium

O. Spermogonia on both surface

leaf, pale yellow, round, in groups

ed yellow spots, about 150 /u diam

Spermatia hyaline, ellipsoid, 3*5-4 x

II. u

amphigenous, bullate

psoid

ded, and becoming confluent, surrounded or partially covered

by the ruptured epidermis, pulverulent, cinnamon-brown, 1 mm

more long

Uredospores obovate to ellipsoid, golden-b

echinulate* with

ened hood

apex

d three or four, mor

quatorial germ-pores on one face

30-3

x 21

hyaline pedicel may reach a length of 56

III. Teleutosori similar to uredosori, dark-brown, also on both surfaces

of leaf, but more
Teleutospores

similarly

under,
coloured to

dospores, shortly oblong to ovate-oblon

d intermixed with
ightly or not at all

cted at septum, finely warted, rounded at apex, and not pe

ceptibly thickened, 29-40 x
lower cell rounded at bas
hyaline, short, decid

6-22

slightly

average

32 x 20 ft

attenuated ; pedicel

On stems and leaves of Avium prostratum Labill

Victoria — Beaumaris, Dec.-Julv, II., III. (III. in

A.pril).

Portland, Jan., 1901, II., III. Sandringham, Jan. and Feb.,

Mentone, Jan. -Sept.,

1904, II., III. (III., very sparse).

1905, II. III.

Tasmania — Hobart, Sept., 1905 (Rod way). Mersey River, January,

1906 (Robinson).
It may be found in sheltered spots all the year round.

On Apium graveolens L.

Victoria — Near Melbourne, April-Sept., II. III.
Tasmania — Hobart, Sept., 1905 (Rod way).

Very common in the early spring months.

Spermogonia were only found on the native celery (Apium prostratum)
associated with uredospores and teleutospores.

The germ-pores of the uredospores may be in a transverse band, or one
may be placed above the other. When fully developed the uredospores are
similarly coloured to the teleutospores, otherwise they are yellowish. On
Apium graveolens both uredospores and teleutospores agree almost exactly in
average size, though of the latter there are usually very few more than 33 p
in length. There are occasionally elongated teleutospores which reach a
length of 40 p. The finely warted epispore is a constant feature of the
teleutospores on both hosts.

The illustrations of spores in PL IX., Figs. 68-72, show that the rust on

the native celery (Apium prostratum), and on cultivated celery (A.
graveolens) is the same in Australia.

Figs. 73, 74 show the smooth teleu-

tospores of P. apii Corda on Apium graveolens from Sydow's Ured. Exsicc,
558, which is quite distinct from the finely warted P. thuemeni, and Fig. 7d
shows the very rough and knobby teleutospores of P. bullata (Pers.) Schroet.

Puccinia — Umbelli ferae. 169

on Aethusa cynapium from Sydow's Ured. Exsicc, 1261, which is readily dis-
tinguishable from either of the above. There is thus a clear distinction be-
tween these three recognised species, but the synonymy is rather confusin
The reason for the name I have adopted may first be given. P. castagnei
Thuem., on cultivated celery agrees with our species, but P. castagnei
Schroet., in Cohn's Beitrage, III., 62 (1879), was first applied to a Puccinia
on Thalictrum angustifolium, and Thuemen's name being thus preoccupied,
I have substituted P. thuemeni.

for

As regards the synonyms, Cooke, in his Australian Handbook, gives P
i Corda at first, and then in an addendum substitutes P. castaqnei Thuem

Then Saccardo, in his Sylloge, gives P. bullata with P. apii as a
synonym, and P. castagnei is regarded as distinct. Further, Sydow in his
Monograph includes P. apii Corda, and P. castagnei Thuem., under P. apii
Desm., and P. bullata is given separately, although he remarks that the
latter is probably a collective species. The teleutospores of P. bullata are
described as smooth, and therefore do not agree with the quoted specimen
from Sydow. Finally, it may be noted that Tranzschel 1 has shown that the
host-plant of P. castagnei Schroet., is not Thalictrum angusti folium, but an
Umbellifer, and probably Silaus pratensis, so that this species is a synonym
of P. bullata.

Darluca filum Cast., occurs on the uredosori and teleutosori.

(Plate IX., Figs. 68-75 ; Plate C, Figs. 16, 17.)

Xanthosia

99. Puccinia xanthosiae McAlp.

II. Uredosori amphigenous, and on leaf -stalks, dark-brown, pulvinate,

gregarious, elliptical, often confluent, rupturing epidermis, about
j mm. long, often causing entire leaf to become brownish.

Uredospores golden-brown, thick-walled, echinulate, globose to
elliptical, 30-32 )< diam., or 30-37 x 24-28 p, average 34 x 26 /x;
pedicel colourless, 4-6 p broad.

III. Teleutosori rare, on leaves of previous year, minute, black.

Teleutospores oblong to oblong clavate, dark-brown, smooth,
constricted at septum, 40-60 x 20-25 jj, average 42 x 20 p ;
upper cell generally darker than lower, rounded and slightly
thickened at apex, occasionally truncate ; lower cell rounded at
base or attenuated towards stalk ; pedicel persistent, tinted, about
20 p long.

X. Mesospores occasional, similarly coloured to teleutospores, obovate,

thickened at apex, 32 x 20 ju.

On Xanthosia pusilla Bunge.

Victoria— Frankston, Oct., 1899, II. : Oct., 1903, III. Sandring-

ham, Sept., March, II, III. ; Feb., II.

(Plate IX., Fig. 76.)

170

Puccinia — Onagraceac^ Rosaceae.

ONAGRACEAE

100. Puccinia epilobii-tetragoni (DC.) Wint.

Winter, Pilze, p. 214 (1884).
Sydow, Mon. Ured. L, p. 424, (190
Sacc. Syll. VII., p. 608 (1888).

Efilob

I. Aecidia distributed en

crowded, hypophyll
laciniate. revolute re

surface of the leaf, scattered

doperidia cup-shaped

th white,

Aecidiospores orange yellow, polygonal, finely warted, 16-26
diam., 18-21 \i being common.

II. Uredosori chestnut-brown, scattered, or often orbicularly arranged

H-

sometimes confluent, pulverulent, hypophyllous, soon naked

Uredosp

psoid or

cL clear bro

echinulate

two germ-pores on one face
reaching a length of 31 p.

7 x 16-21

nally

III. Teleutosori dark-brown, round, often arranged on

hypophyllous, soon naked, pul\

Teleutospores
yellowish-brown

like spot

intermixed with

dosp

htly constricted at

elliptic or oblong,
smooth, 27-36

x 15

erage 30 x

18 /.i ; upper cell with a cap-like
thickening at apex (5-6 fj) ; lower cell usually rounded at base
or sometimes slightly tapering ; pedicel hyaline, slender, deciduous,
short.

X. Mesospores exceedingly rare, similarly coloured to teleutospores and

thickened at apex (5 ft), subclavate to obovoid, smooth, 21-27 x
13-15 /j, with hyaline pedicel.

On leaves of Efilobmm glabellum Forst.

Victoria — Dimboola, Dec, 1892, I., II., III. (Reader). Ardmona,

Arthur's Creek, Kergunyah, Nyora, Pakenham, &c, Aug.
Dec, I., II., III.

On E. billardierianum Ser.

Tasmania — Great Lake, Feb., 1894 (Rodway

On Epilobium sp.

■

Tasmania — Summit of Mt. Wellington, Jan. 1892, I. (Rodway).
Sydow, in his Monograph, distinguishes between P. epilobii DC, with
teleutospores alone, P epilobii-fleischeri Fisch., with aecidia in addition to
teleutospores, and the present species with the three stages.

(Plate X., Figs. 79, 80, 81.)

ROSACEAE

101. Puccinia gei McAlp.

Geum

McAlpine, Agr. Gaz. N.S.W. VI., p. 756 (1895).
Sydow, Mon. Ured. I., p. 484 (1903).
Sacc Syll. XIV, p. 297 (1899).

III. Teleutosori hypophyllous, confluent, coffee-brown, on greenish-
yellow spots, which also occur on upper surface of leaf.

Puccini a — Rosac eae.

*7*

Teleutospore brownish-yellow, smooth, fusiform to clavate, con-
stricted at sepfcam, epispore chestnut-brown, 30-40 x 13-18 //,
average 35 X 14 p ; upper cell elongated and tapering or
rounded, thickened at apex, 18-22 /* long; lower cell elongated
and tapering towards base, or bulging, 16-22 p long; pedicel
pellucid, sometimes 54 /t long.

X. Mesospores occasional, coloured like teleutospores, elongated

ellipsoid to oblong, rounded and thickened at apex, occasionally
somewhat pear-shaped, and the basal portion prolonged like a
stalk, as sometimes happens in the bicellular spore, 28-35 x
12-18 ft.

On living leaves of Geum renifolium F. v. M.

Tasmania — The Calf, Adamson's Peak, March, 1895 (Rod way1).
The specimen was gathered on an out-of-the-way mountain, at an altitude

of nearly 4,000 ft.

(Plate X., Fig. 82.)

102. Puccinia pruni Pers.

Primus

Persoon, Syn. Fung., p. 226 (1808).

Cooke, Grev. XI., p. 97 (1883).
Sydow, Mon. Ured. I., p. 484 (1903).
Sacc. Syll. VII., p. 648 (1888).

II. Uredosori hypophyllous, occasionally epiphyllous, minute, light

brown to cinnamon brown, orbicular, scattered, but often grouped
in patches and confluent, soon naked, pulverulent, usually seated
on discoloured spots, which also show on upper surface.

Uredospores variable in shape, oblong to ovoid oblong, ellipsoid
to somewhat piriform, closely echinulate, yellowish, with two or
occasionally three sub-equatorial germ-pores on one face, apex
yellowish brown, thickened (average, 5-6 ^u, or papillate up to 9 yu)
with spines less prominent, bluntly conical or rounded, 25-38 x
* 12-18 ju, occasionally reaching a length of 40 fx or more, average
30 X 15 fi ; paraphyses intermixed, numerous, capitate, pale yel-

low, long-stalked, sometimes attaining a length of (0 p.

III. Teleutosori hypophyllous, scattered or confluent, isolated or in

groups, very pulverulent, seal-brown, paraphysate, known from the
uredosori when fully developed by their dark, almost black

appearance

Teleutospores at fi

dospores, dark-b

oblong to ellipsoid or obovoid, densely warted, often composed of
two globose or depressed globose cells, readily separating from

each other, very rarely 3-celled, 25-45 x 17-25 n> average

36 x 25 jx ; upper cell usually darker in colour, and broader
lower, globose or depressed globose, often slightly thickened
at apex, and thickly studded with short stout spikes, average
16-22 fi diam.; lower cell generally oblong to obovoid, and equal to
or longer than upper, 14-24 X 14-19 n ; pedicel short, hyaline,
deciduous.

On leaves, fruits and stems of Peach {Primus persica Stokes).
On leaves and fruits of Almond and Apricot (Prunus amygdalus Stokes,
and P. armeniaca L.).

;

172

Puccini a — Leguminosae.

On leaves of Plum [Primus domes tica L.) and Nectarine.

Victoria, New South Wales (Cobb1), Queensland (Bailey5)

(Tryon1), South Australia, West Australia, and Tasmania
(Rod way1).
Common. November to June, the teleutospores being very common in

March on Primus domestica.

This rust is usually most prevalent towards the autumn, but in some
seasons it appears in the early summer. It not only destroys the foliage,
but may even attack the fruit, and in some seasons it actually ruptures the
bark, especially in rapidly growing nursery stock. When the rust occurs on
the fruit, uredospores may be produced both at the surface and in the pulp.

There are said to be two kinds of uredospores by Dumee and Maire1, but
I33 have shown that this is simply due to an error of interpretation, and
that they are always thickened at the apex when seen in their natural
position, and not obliquely. Teleutospores are comparatively rare on the
Apricot and Peach, less so on the Almond, and very common on the Plum.
In April, 1904, both stages of the rust were found on leaves of all the host-

plants.

According to Prillieux1, teleutospores often appear alone, without

having being preceded by uredospores, but this has never been known to
occur in Australia. The arrangement of the teleutospores in the sorus is
worthy of note. With a magnifying glass the spores can be seen to be
arranged in minute clusters, and each cluster under the microscope is seen to
consist of a number of spores, the stalks of which have become agglutinated
together, and to the free end of each a spore is attached. (McAlpine34).

On the leaves of the Peach the sori are situated on yellow spots, which
s,ve very prominent on the upper surface. There they often run together,
and have the appearance of yellow ochre. In the Apricot the spot is hardly
noticeable at first, but when held up to the light the indeterminate yellow
spots are seen clearly. The leaf soon turns yellow, and then the spots are
of a pale green. In the Plum the spots are very pale on the lower surface,
but on the upper surface the yellowish-green is very prominent.

Tranzschel 2 has proved experimentally that the aecidial stage of the rust
on Almond occurs on Anemone coronaria L., and it has hitherto been known
as Aecidium <punctatnm Pers.

Darluca filum Cast., is found on the uredosori.

(Plate X., Figs. 83-86 ; Plate D., Figs. 19, 20.)

LEGUMINOSAE.

Zornia.

103. Puccinia zorniae (Diet.) McAlp.

Uredozorniae Dietel, Hedwigia XXXVIII., p. 257 (1899).

II. Uredosori on both surfaces of leaf, but most numerous on under,

yellowish when young, then ruddy-brown, round to elliptic,

bullate, at first covered, then erumpent and surrounded by rup-
tured epidermis, gregarious and sometimes confluent.

Uredospores old-gold colour, ellipsoid to occasionally obovoid,
finely echinulate, with two germ-pores on one face, 25-32 x 19-22/1.
III. Teleutosori as above, but dark-brown.

Teleutospores intermixed with uredospores and similarly
coloured, oblong, smooth, deeply constricted at septum, two cells
about equal in size or lower sometimes a little longer, 32-44 x
20-24 /i, average 36 x 21 /i ; upper cell rounded and thickened
at apex (up to 6 /lc), lower cell rounded at base ; pedicel hyaline,
persistent, about length of spore.

;

Puccini a — Ficoideae.

173

On living leaves of Zornia diphylla Pers.

New South Wales — Richmond River, May, 1904 (Musson).

Uredozorniae Berk., is mentioned by Cooke in Grev. XX., p. 110, as on
Zcrnia in Mauritius, but no diagnosis is given. This is the first record of
the teleuto stage, since Dietel in his Uredineae brasilienses only men-
tions the uredo stage on the same plant from Rio Janiero, and thus describes
it: — "Sori hypophyllous, scattered, surrounded by the ruptured ochraceous
epidermis, minute, brown. Uredospores globose, obovate or ellipsoid,
21-28 x 18-23 fx ; epispore brown, echinulate, with three germ-pores."

Darluca Jilum Cast., occasionally on sori.

(Plate X, Fig. 87.)

FICOIDKAE.

104. Puccinia tetragoniae McAlp.

Tetragonia

McAlpine, Agr. Gaz. N.S.W. VI., p. 854 (1895).
Sydow, Mon. Ured. I., p. 563 (1903), p. 895 (1904).
Sacc. Syll. XIV., p. 295 (1899).

0. Spermogonia orange, numerous, aggregated, amphigenous, 150-170 f.i

in diam.

Spermatia hyaline, globose to oval, 2 J- 3 J /j. diam.

1. Aecidia hypophyllous, crowded, often arranged in lines, greenish to

yellowish-brown knob-like swellings before bursting ; pseudoperidia
white, cylindrical, with torn edges, average 630 ji diam.

Aecidiospores orange-yellow, minutely warted, variously shaped,

lobose to ovate or mussel-shaped, 27-39 x 22-25 /u

II. Uredosori amphigenous, soon naked, bullate, orange-red, generally

orbicular, 1-1 ^ mm. diam.

Uredospores orange-yellow, ellipsoid to ovoid, strongly echinu-
late, 27-37 x 22-25 fj, average 31 x 24 ju.

III. Teleutosori amphigenous, black, compact, bullate, orbicular, irre-
gular or elongated, scattered, covered by epidermis for some time,
then erumpent.

Teleutospores golden brown to chestnut-brown, elliptical to
oblong, usually rounded at both ends, slightly constricted at
ptum, smooth, 40-60 x 25-36 ^, average 50 x 30 ju ; pedicel

hyaline, stout, persistent, up to 80

X. Mesospores similarly coloured to teleutospores, ellipsoid or elongated,

thickened at apex, 35-54 x 25-29 ja.

On leaves and stems of Tetragonia implexicoma Hook. f.

Victoria. — On the coast at Sandringham, Beaumaris, Flinders,

Portland, <fcc, all the year round.
Tasmania — Hobart, about rocks, Aug., 1897, I., II., III. (Rod-
way1).

All the stages are usually very plentiful in the month of August.

The spermogonia precede or accompany the aecidia on upper and under
surface of the leaves, are flask-shaped and the neck projects slightly between
the watery vesicles which stud the surface of the leaf. They often form

x74 Puccinia — Polygonaceae.

large, orange, blister-like clusters quite conspicuous against the green of the
leaf, and the rust-devouring Diplosis was ^frequently met with there and
probably scatters the spermatia.

The aecidia are usually swarming with the Diplosis, dipping into them
as an insect inserts its proboscis into a flower, and sometimes it is entirely
inside the cups. The larvae are invariably coated with the aecidiospores.
Occur August to October and subsequently.

Uredospores just beginning to appear in October, and the Diplosis was

also found on the sori.

Teleutospores are scanty in October, and they occur on the same leaves

as uredospores.

The uredo and teleutosori are often associated together in the same
cluster, and the three spore-forms may all occur on one leaf.

(Plate XL, Figs. 88, 89.)

POLYGONACEAE.

Rumex.

105. Puccinia ludwigii Tepp.

Tepper, Bot. Centralb., XLIIL, p. 6 (1890).
Cooke, Handb. Austr. Fung., p. 336 (1892)

Sydow, Mon. Ured. L, p. 581 (1903).

Sacc. Syll. XL, p. 196 (1895).

II., III. Sori minute, circular, irregularly scattered or aggregated into

small groups, soon naked and girt by the ruptured epidermis, on
both surfaces of leaf, up to \ mm. diam.

II. Uredospores sub-globose to ovate, very finely echinulate, pale

yellowish-brown, 22-28 X 16-20 /x, or 19-22 /x diam.

III. Teleutospores at first intermixed with uredospores, oblong to

ellipsoid, rounded at both ends, very coarsely warted, slightly con-
stricted at septum, not thickened at apex but generally with
hyaline apiculus, sometimes three or four celled, dark brown, 30-40

x 20-25 p, average 32 x 21 p; pedicel short, hyaline, deciduous.
X. Mesospores similarly coloured to teleutospores, warted and with

hyaline apiculus, oval, with short hyaline pedicel, 28-31 x 18-23 /u.

On leaves of Rumex brownii Campd.

"Victoria — Coromby, Oct., 1889 (Tepper). Shepparton, on river

flats, Nov., 1895 (Robinson). Flinders, Jan. Murramurrang-
bong Ranges, Dec. Killara, March. Myrniong, Aug.

Queensland — Ennogera (Bancroft). Brisbane, Sept., 1886

(Bailey

Tasmania — Devonport, January, 1906 (Robinson).
On Rumex flexuosus Sol.

Victoria — Warracknabeal, Oct., 1903 (Reader)

I have had plenty of material of this species, both from Victoria and
Queensland, as well as some of the original material from Ludwig, and ex-
amination shows that it is the same fungus in each case.

Mr. Bailey kindly supplied specimens from Queensland, and there is one
in the National Herbarium, Melbourne, sent from Queensland to Dr. Cooke
in 1886, and it is named in his own handwriting P. rumicis Lasch.,

P. acetosae (Schum.) Koern. It has the same characters as the others, and

comparing it with P. acetosae from Syd. Ured. Exs., 1163, I find that it

■

Puccinia — Polygonaceae. 175

differs in the more finely echinulate uredospores, but more particularly in
the teleutospores, which are coarsely knobbed and altogether characteristic.

The introduced Rumex crispus and R. acetosella are exceedingly common
weeds in Victoria, and may be seen growing alongside rusty R. broumii, but
no rust has ever been detected on them.

R. brownii is a native species, and its parasite is evidently a native also.

The teleutospores are varied in shape, and three and four celled forms
are met with. In the three-celled forms they may either be after the
P hragmidium or Triphragmium type, and in the four-celled forms the upper
third cell is vertically divided. Just as in P. dichondrae there is every
gradation from the one to the four-celled spore.

Darluca filum Cast., is common on the uredo and teleutosori.

(Plates XI., Figs. 90, 91, 92 ; XL., Fig. 300.)

Muehlenbeckia

106. Puccinia muehlenbeckiae (Cooke) Syd.

Sydow, Mon. Ured. L, p. 566 (1903).
Sacc. Syll. IX., p. 299 (1891).

Puccinia rumicis- scutati (DC) Wint. var. muehlenbeckiae
Cooke, Grev. XIX., p. 47 (1890), and Handb. Austr. Fung., p.

336 (1892).

II. Uredosori hypophyllous, scattered or in irregular groups, some-
times circularly arranged, round or elliptic, girt by the torn
epidermis, cinnamon-brown.

Uredospores yellowish-brown, ellipsoid or ovoid, echinulate,

24-32 x 16-21 /a, average 28 x 19 p.

III. Teleutosori similarly arranged to uredosori, but dark-brown.

Teleutospores at first intermixed with uredospores, clavate to
oblong, dark-brown, constricted at septum, thickened at apex (up
to 7 /u), and rounded or obtuse, sometimes 3-4 celled, 32-45 x
14-19 ju, occasionally reaching a length of 55 /*, average 40 X
1 8 a ; lower cell generally somewhat triangular ; pedicel hyaline,

ally persistent, up to 40 f.i Ion""

&•

X. Mesospores occasional, similarly coloured to teleutospores, ellipsoid to

elongated oblong, smooth, rounded or conoid and thickened at apex
(up to 5 p), 31-34 x 12-18 /w.

On leaves of Muehlenbeckia adpressa Meissn.

Victoria— Brighton, May, 1894 (Morrison). Near Melbourne,

Jan.-July. Cape Schanck, March, 1903 (C. French, jun.).
Frankston, Feb., 1904 (Robinson). Sandringham, Sept.,
1905, II., III.

On Muehlenbeckia gracillima Meissn.

Queensland— Gladfield (Gwyther) (Bailey1*).
N.S. Wales — Kurrajong Heights (Musson).

Some of the mesospores show their connexion with teleutospores very
clearly. In one instance two germ-pores were formed in the usual positions,
while in another there was a slight notch on one side as if the beginning of
a septum. They are evidently one-celled teleutospores, and have all their
characters with the exception of the septum.

1 76 Puccinia — Chenopodiaceae.

There were spots on the upper surface of the leaves with purplish-red
margins, but these were due to gall-mites. The spores are of the general type
of P. rumicis-scutati, but both kinds are somewhat smaller. I had a large
amount of material to operate upon, and found the sori to be hypophyllous,
although Cooke inadvertently describes them as epiphyllous. In a specimen
of P. rumicis-scutati, from Syd. Ured. Exs. 25, the teleutospores are of a deep
chestnut-brown, and attain a length of 57 /m. Sydow himself had not seen
any specimens of the rust on Muehlenbeckia, but even from the brief
description of Cooke he was right in making a species of it.

Darluca filum Cast., sometimes common on uredosori.

(Plate XL, Fig. 93.)

■

CHENOPODIACEAE

Threlkeldia.

10/. Puccinia dielsiana P. Henn.

Hennings, Hedw. XL., p. (95) (1901).
Sydow, Mon. Ured. L, p. 566 (1902).
Sacc. Syll. XVIL, p. 361 (1905).

III. Teleutosori on stems, large, compact, bullate, erumpent, up to f cm.

long.

Teleutospores ellipsoid or oblong ellipsoid, generally rounded at
both ends or obtuse and slightly thickened at apex (3-5 ll), smooth,
very slightly constricted at septum, with granular contents,
chestnut or dark-brown, 35-18 x 18-28, average 38 x 22 fx ;
pedicel sub-hyaline, persistent, 80-120 x 5-6 fx.

X. Mesospores occasional, ellipsoid to ovoid, coloured and thickened at

apex like teleutospores, 34 x 23 fi.

On stems of Chenopodiaceae — Threlkeldia drupata Diels.

W. Australia— Near Perth, 1900.

The teleutospore has occasionally a vertical septum, and in elongated
slender specimens the breadth is reduced to 18 u.

XL, Fiff. 94

108. Puccinia kochiae Mass.

Kochia, Enchylaena.

Massee, Grev. XXIL, p. 17 (1893).
Sydow, Mon. Ured. I., p. 565 (1903).
Sacc. Syll. XL, p. 196 (1895).

II., III. Sori amphigenous, discoid, plane, very compact, blackish-
brown to black, often aggregated together, girt by the ruptured
epidermis, up to 1 mm. diam.

II. Uredospores elliptical, golden-brown, decidedly echinulate, with

numerous prominent germ-pores, as many as twelve on one face,
29-34 x 18-25 li, 32 x 23 li being very common.

III. Teleutospores densely packed, intermixed with uredospores,

elliptic-oblong, rounded at both ends, or the apex with a slight
indication of a papilla which is often oblique, perfectly smooth,
chestnut-brown, hardly constricted at septum, epispore about 3 /t
thick, 35-45 x 22-29 li, average 37 X 25 li ; pedicel tinted
yellowish, persistent, 30-40 x 6 li, but may reach a length of 96 /*.

X. Mesospores few, similarly coloured to teleutospores, oval to flattened

at apex, 28-35 x 22-25 li.

Puccinia — Caryophyllaceae, Malvaceae. 177

On leaves and stems of Kochia sedifolia F. v. M.; Kochia villosa, Lindl.;
and Enchylaena tomentosa R. Br.

Victoria — Dimboola, Nov. and Dec. (Reader).

I have some of the material from Dimboola, portion of which was sent
to Massee, and the golden-brown uredospores are seen along with the
teleutospores. No uredospores were mentioned in the original description, a
striking feature of which is the numerous germ-pores. The teleutospores
have occasionally their septum lengthwise.

(Plate XI., Figs. 95, 96.)

CARYOPHYLLACEAE.

Stellaria

109. Puccinia arenariae (Schum.) Schroet.

Schroeter, Pilz. Schles., p. 345 (1872).
Sydow,Mon. Ured., I., p. 553 (1903).
Sacc. Syll. VII., p. 683 (1888).

III. Teleutosori compact, pulvinate, roundish, scattered, often circinate,

pale-brown to dark -brown.

Teleutospores fusoid or clavate, apex pointed or rounded,

often thickened (6-8 /u), base rounded or attenuated, slightly
constricted at septum, smooth, pale yellowish brown,
30-50 X 14-20 /j, average 35 x 16 /u, very rarely tricellular,
48 X 16 ft; pedicels hyaline, persistent, equal to or longer than
the spores, 60-100 p.

X. Mesosperes occasional, similarly coloured to teleutospores, obovoid

or somewhat wedge-shaped, rounded or obtusely pointed or
truncate at apex and thickened up to 6 /w, generally resembling
the upper cell of the teleutospore, with elongated pedicel, 22-29
X 16-17 /x.

On leaves and stems of Stellaria media Cyrill.

Victoria — Gellibrand, Feb., 1896 (Hill). Gem brook Ranges, April,

1904 (C. French, jun.).

The germ-tubes of the sporidiola of this species have been known to enter
the stomata of Dianthus barbatus L., and it seems to be the only known case
in the genus Puccinia of such germ-tubes entering stomata.

(Plate XII., Fig. 97.)

MALVACEAE.

110. Puccinia heterospora Berk, and Curt.

Abulilon, Hibiscus.

Berkeley and Curtis, Journ. Linn. Soc. X., p. 356 (1868).
Cooke, Handb. Austr. Fung., p. 338 (1892).
Sydow, Mon. Ured. I., p. 472 (1903).
Sacc. Syll. VII., p. 695 (1888).

III. Teleutosori minute, hypophyllous, soon naked, crowded in orbicular,

glomerules, dark-brown, on determinate purplish or yellow spots.

Teleutospores very variable in size and shape, smooth, thick-
walled and more deeply thickened at apex, dark chestnut-brown,
elliptic or oblong, not or scarcely constricted at septum,
occasionally divided vertically, 20-38 X 17-28 ya; pedicel hyaline,
slender, elongated, three to four times as long as spore.

i78

Puccinia — Malvaceae.

X. Mesospores much more common than teleutospores, subglobose or

elliptical, apex more thickened, pale-brown, \i

25-32 x 17-22 //.

2 2

\i diam., or

On leaves of Abutilon crispum Sweet, and Hibiscus sp. (Bailey 1G).

Queensland — Gulf of Carpentaria and St. George, Oct., 1885

(F. v. Mueller in National Herbarium).

On Abutilon avicennae Gaertn.

New South Wales — North-west part, Oct., 1887 (Bauerlen).

There are only relatively few bicellular spores present, and at first they
were overlooked, the fungus being named Uromyces pulcherrimus B. and C,
^ind U. thwaitesii B.^and Br.

The so-called mesospores are probably of the nature of one-celled
teleutospores, and this species may be regarded as a transition stage from the
unicellular Uromyces to the bicellular Puccinia.

(Plate XII., Fig. 101.).

Althaea, Lavatera, Malva, Plagianthus.

ill. Puccinia malvacearum Mont,

Montagne in Gay's Hist. Chili VIII., p. 43 (1852).
Cooke, Handb. Austr. Fung., p. 338 (1892
Sydow, Mon. Ured. I., p. 476 (1903).
Sacc. Syll. VIL, p. 686 (1888).

III. Teleutosori generally hypophyllous, brown to reddish-brown or

orange, but grayish when spores are germinating, compact, round,
pulvinate, elongate on the stems, scattered or crowded, seldom
confluent, at first covered by epidermis, soon naked.

Teleutospores fusiform, attenuated at both ends, apex sometimes
rounded, slightly constricted at septum or not at all, apical

smooth, yellowish-brown, rarely tricellular,

17 \i\ pedicels firm, long, per-

thickening slight,

35-75 x 12-26 u,

average 50 x

sistent, occasionally septate, sometimes measuring 170 fx long.

X. Mesospores occasional, same colour as teleutospores, somewhat

fusiform, slightly thickened at apex, 40-45 X 17-19 p.

On all green parts of Malva rotundifolia L., M. sylrestris

rosea Cav
ceedingly

Lavatera plebeia Si

Plagianthus spicatus Benth

L., Althaea

Ex-

(Berkeley2).

Victoria —

New South Wales — (Sacc. and Berl.1).

Queensland — (Bailey7).

South Australia.

West Australia — (Morrison).

Tasmania — (Rod way l ).

The sori first appearing were carefully examined for any trace of
accompanying spermogonia, but none were found. Spores occasionally
3-celled, and reaching a length of 77 fi.

This fun

which was

ally described by Monta

spread the earth and occurs on both wild and

now

cultivated species
Malvaceous plants. Dr. Plowright5 states that he has a specimen in
herbarium from Melbourne, gathered as early as 1865 on Malva sylvestris

his

•

Puficinia — Malvaceae, Geraniaceae. 179

Puccinia malvace arum is one of the commonest of rusts, and the
different stages in the growth of the spore are well seen in this species. A
group may be observed springing from a common hypha, and at first they
are without septa, but soon a transverse septum is formed, and, although
specially looked for, I have never seen a partial septum as recorded by
Fischer 5.

Germination occurs as soon as the spore is ripe, but some may hibernate.
Occasionally the pedicel has a septum. Mr. French, Government Entomolo-
gist, has observed this rust on Hollyhocks in a nursery near Melbourne as
early as 1857.

(Plate XII., Figs. 99, 100 ; Plate XV., Figs. 123-130 ; Plate F., Fig. 28.)

112. Puccinia plagianthi McAlp.

Plagianthus.

•

McAlpine, Proc. Rov. Soc. Vic. VII, N.S., p. 218 (1894).
Sydow, Mon. Ured. I., p. 480 (1902).
Sacc. Syll. XIV., p. 295 (1899).

III. Teleutosori hypophyllous, sparingly epiphyllous, scattered, pul

verulent, soon naked, up to 1-1 J mm.

Teleutospores golden-brown, oblong to clavate, rounded at apex
slightly attenuated towards base or rounded, slightly constricted al
septum, very rarely three celled, epispore with reticulated markings
38-50 x 17-23 /j, average 41 x 20 jj. ; pedicel hyaline, short,
deciduous, but may attain a length of 60 ft.

On leaves and flowers of Plagianthus sidoides Hook. Very common.

Tasmania — Mount Wellington, southern slope, Aug. -April (Rod

The spore is described as smooth by Sydow, but

be covered

with a fine irregular network

(Plate XII., Fig. 98.)

GERANIACEAE

Geranium

113. Puccinia geranii-pilosi McAlp

£—

II. Uredosori hypophyllous, chocolate-brown, scattered or sub-gr

ious, orbicular, soon naked and surrounded by ruptured epidermis
compact, up to 1 mm. diam., often on purplish-red spots or portion:

of leaf becoming coloured.

Uredospores yellowish-brown, elliptical to obovate, echinulate

epispore slightly thickened at base, 25-30 X 19-22 /i.

III. Teleutosori elongated, dark-brown to black, sometimes confluent

erumpent, compact

ly on peti

Teleutospores at first intermixed with uredospores, oblong to
oblong-clavate, chestnut-brown, smooth, slightly constricted at
septum, sometimes short and stout and about as broad as long,
32-48 x 22-32 /i, average 36 X 25 /* ; upper cell rounded at apex,
not or slightly thickened ; lower cell rounded at base or occasion-
ally slightly tapering; germ pore generally indicated at apex by a
paler groove ; pedicel hyaline, deciduous, elongated, up to 80 /i long.

i8o

Puccini a — Geraniaceae.

X. Mesospores occasional, similarly coloured, subglobose, thickened at

apex, 25-26 \i diam.

On living leaves of Geranium pilosum Sol.

Victoria —

-Killara, Nov., 1902, II., March, 1903, II., £11. (Robin-
son).

New South Wales — Richmond (Musson).

Occasionally a teleutospore occurs in which the septum is vertical as in
Diorchidium.

In the recorded species of Puccinia on Geranium there are only teleu-
tospores with the exception of P. callaguensis Neger, on leaves of Geranium
berterianum Colla, in Chili.

It differs from this species however in the teleutospores being smooth
and not minutely verrucose, and in the elongated pedicel.

The host-plant is variously named. It is found in all the Australian
States, and was called G. pilosum by Solander.

In the Index Kewensis it is given as a synonym of G. dissectum, but
the late Baron von Mueller used the name of G. pilosum, considering it
"more exact than that of G. carolinianum and G. dissectum."

F. M. Bailey, in his Queensland Flora, divides G. dissectum into two
principal races, one of which is pilosum.

The species of Puccinia found upon it is new, and while it is not cus-
tomary to determine a host-plant from the species of rust found upon it,
still in this instance it lends probability to the view that we are here dealing
with an indigenous form of Geranium, differing from G. dissectum. For
this reason I have not followed the Index Kewensis in naming the host.

(Plate XII., Figs. 103, 104.)

114. Puccinia morrisoni McAlp.

Pelargonium

I. Aecidia amphigenous, delicate, minute, orange, circinate, either alone

or accompanied by teleutospores ; pseudoperidia with expanded and

lobed margin

peridial cells subquadrate to oblong with striated

margin, and punctate all over, firmly united, 28-32 x 16-22 ft.

Aecidiospores pale orange, finely echinulate, elliptic to subquad-
rate, 19-22 x 16 li, or 18-19 fx diam.

II. Uredosori mostly on under surface of leaf, round, small, brownish,

pulverulent, solitary or arranged in confluent groups.

Uredospores subglobose to ellipsoid, yellowish brown, echinulate,
22-24 li diam., or 22-27 x 19-22 u.

in small confluent

III. Teleutosori

i intermixed, solitary or arranged

groups, generally round, black, compact, surrounded by ruptured
epidermis.

Teleutospores chestnut brown, clavate, constricted at septum,

smooth, often obliquely pointed or rounded and thickened

at apex (up to 9 li), 33-64 x 19-25 li, average 45 x 22 ^ ;

lower cell usually paler in colour and attenuated towards base ;

pedicels tinted, generally short, persistent, up to 38/.* long by 6-9 li
broad.

X. Mesospores coloured

like teleutospores, oblong to oval, bluntly

pointed and thickened at apex, 33-38 X 16-20 fi

Puccini a — Rutaceae.

181

On stems, leaf -stalks and leaves of Pelargonium australe Jacq

Range

Victoria— 1892, II., III. (Morrison). Phillip Island, Jan., 1900

II., III. Murramurrangbong Ranges, Dec, 1903., I., II.
III. (Robinson).

Tasmania — Devonport, Jan., 1906 (Robinson).
The only locality where aecidia were found was the Murramurangbon

©~~"»

It differs from P. geranii Corda, to which this species was referred bv
Cooke in Grevillea, XXI., p. 39, in the uredospores being nearly twice as
large, and in the much longer and broader teleutospores.

There are two species of Puccinia already recorded upon the genus
Pelargonium from S. Africa, viz., P pelargonii (Thuem.) Syd., and P.
granular is Kalch. and Cooke.

i In P. pelargonii the aecidia are hypophyllous, and not amphigenous, and

'ores are rather shorter and narrower. In P. grauularis there

aecid

(Plate XII., Fig. 102 ; Plate F., Fig. 29.)

RUTACEAE.

115. Puccinia boroniae P. Henn.

Boronia

Hennings, Hedw. XLIL, p. (73) (1903).
Sydow, Mon. Ured. I., p. 891 (1904).
Sacc. Syll. XVII., p 351 (1905).
III. Teleutosori on branches, erumpent, pulvinate, tuberculate, dark

cinnamon, compact, confluent lengthwise up to 2 cm. long, sur-
rounded at base by the ruptured epidermis.

Teleutospores ellipsoid to ovate, oblong, brown, smooth, rounded or
bluntly pointed at apex and slightly thickened (up to 5 n), gen-
erally rounded at base, 28-40 x 16-21 /u, average 35 x 20 yu,
occasionally 3-celled and elongated, 52 x 25 li ; pedicel hyaline to
yellowish, persistent, up to 120 /u long.
X. Mesospores common, similarly coloured to teleutospores, ellipsoid to

obovoid, rounded or bluntly pointed at apex and slightly thickened,
30-34 x 17-18 fx.

On Boronia spinescens Benth.

West Australia — Avon, Oct., 1903 (Diels).

A very interesting Leptopuccinia which projects from the cortex of the
branches in thick, often confluent pustules, and partially surrounds them.

A specimen was kindly forwarded by Hennings, and the mesospores were
observed which are not given in the original description. The size of the

teleutospores, as given by Hennings, is 20-32 x 15-18 ^i, by Sydow. 22-35

X 16-19 fx, and I have made the average 35 x 20 /u, my measurements

being corroborated by those in the photographs, which are also about

35 x 20 it

(Plate XIII., Fig. 107.)

Correa.

16. Puccinia correae McAlp

Mc Alpine, Proc. Roy. Soc. Vict. VII., N.S., p. 215 (1894).
Sydow, Mon. Ured. I., p. 462 (1903).
Sacc. Syll. XIV., p. 297 (1899).
III. Teleutosori hypophyllous, cushion-shaped, circular or interruptedly

circular, dirty brown, scattered, soon naked.

l82

Puccinia — Rutaceae, Tremandraceae.

Teleutospores yellowish-grey, long-stalked, elongated fusoid, con-
stricted at septum, 44-60 X 17-20 ju, average 50 x 18 yu; upper
cell elongated, tapering and rounded at apex ; lower cell tapering

several times as long as

towards

base ; pedicel light

grey,

teleutospore.

X. Mesospores similarly coloured and stalked, ovoid, with truncated

apex, 25-28 x 16-19 p.

On leaves of Correa lawreitciana Hook.

Tasmania — Mount Wellington, Dec. Coast near Trial Harbour,

Jan., 1894 (Rodway').

The sori stand out very distinctly from the cinnamon-brown under
surface of the leaves, causing corresponding circular depressions on upper
surface, of a yellowish-green colour,
the rust was very rare.

Although the plant is

very

common

(Plate XIII., Fig. 106.)

Eriostemon,

117. Puccinia eriostemonis Mc Alp.

III. Teleutosori hyphophyllous, dark-orange, ellipsoid, somewhat cir-
cularly arranged, erumpent, surrounded by ruptured epidermis,
up to 2 mm. long.

Teleutospores clavate to ellipsoid, with finely granular orange
contents, compact, smooth, generally rounded at apex, sometimes
pointed, slightly thickened, usually tapering towards base, con-

stricted at septum, 30-45 x 15-18 /.i

average

32 x 16 y.

pedicel persistent, hyaline, elongated, up to 125 /i long by 3
broad.

°2

X. Mesospores occasional, similarly coloured to teleutospores, ellipsoid,

thickened and bluntly pointed at apex, 30-32 X 13-16 y.

On leaves of Eriostemon myoporoides DC.

Victoria — Mt. Bernard, Nov., 1903 (C. French, jr.). Alps, near

Bright, Dec, 1904 (C. French, jr.)

Occasionally a spore occurs in which the septum is longitudinal.

This species differs from P. correae McAlp
instead of brown, the spores are smaller, with
tents, and the.epispore rather thicker.

(Plate XIII., Fig. 105.)

, in the sori being orange
finely granular orange con-

TREMANDRACEAE.

118. Puccinia pritzeliana P. Henn.

Tremandra

Hennings, Hedw. XL., p. (95) (1901).
Sydow, Mon. Ured. I., p. 460 (1902).
Sacc. Syll. XVII., p. 350 (1905).

III. Teleutosori hypophyllous, on round brown spots, frequently with

scattered or in orbicular groups, pulvinate,

purplish margin,

cinnamon- brown, \-\\ mm. diam.

Puccinia — Violaceae.

1 83.

Teleutospores oblong or oblong clavate, obtusely rounded at
apex, which may or may not be thickened, sometimes two-papillate,
or occasionally somewhat crested, slightly constricted at septum,
generally rounded at base, yellowish, smooth, 40-60 x 15-25 /u,

'»

48 X 20 fj. ; pedicels hyaline, persistent, up to 150

X. Mesospores rare, similarly coloured to teleutospores, somewhat
x oblong and bluntly apiculate at apex, 43 x 28 /*.

On living leaves of Tremandra stelligera R. Br.

West Australia— Near Perth, Oct., 1900 (Pritzel).

Quite distinct from P. tremandrae Berk, from Ceylon.

(Plate XIII., Fig. 108.)

VIOLACEAE.

119. Puccinia hederaceae McAlp.

Viola.

O. Spermogonia arranged in small or large groups, crowded, honey-
coloured.

Spermatia hyaline, globose, minute, 2^-3 \x d

I. Aecidia on all green parts of the plant and flowers, on both surfaces of

leaf but mostly hypophyllous, disposed in circular or irregular
groups ; pseudoperidia flat, about A mm. diam. with white, torn,

re volute edges.

Aecidiospores subglobose to shortly elliptical, finely warted,
orange-yellow, about 16 /x diam., or 16-19 x 12—14 yu.

II. Uredosori minute, brown, roundish, erumpent, scattered, soon

naked, on both surfaces of leaf.

Uredospores subglobose or shortly elliptical, golden-brown,
ly echinulate, 2-3 germ-pores on one face, 25-29 X 18-22 u,

average 25 x 21 ji and 22-25 /u diam.

III. Teleutosori minute, black, roundish, erumpent, usually surrounded

by ruptured epidermis, pulverulent.

Teleutospores at first intermixed with uredospores, elliptic or
oblong, rounded at both ends or slightly attenuated towards base,,
hardly constricted at septum, distinctly warted all over but prom-
inently on upper cell, chestnut-brown, usually with hyaline
apiculus, occasionally unicellular, 29-49 x 17-22 //, average 35
X 19 fi ; pedicel hyaline, deciduous, short.

On living leaves of Viola hederacea Labill.

Victoria — Near Melbourne, Oct., Nov., and Dec, 1885, 1886

(Reader). Wandin, Dec, 1895. MurramurraDgbong Ranges,

Nov., 1902, Dec, 1903, Jan., 1905 (Robinson). Mt. Black-
wood, Dec, 1902. Alps, near Bright, Dec, 1904 (C.
French, jun.)
Tasmania — Cascade Rivulet, Dec, 1892 (Rodway 1). Mt. Drome-
dary, Dec, 1894 (Rodway1). Devonport, Jan., 1906
(Robinson).

On V. betonicifolia Sm.

Tasmania — Ben Nevis, April, 1896 (Rodway).

New South Wales — Monga (Bauerlen), recorded as Aecidium

violae Schum. by Baker1. Richmond (Musson).
Victoria — Murramurrangbong Ranges, Nov., 1902 (Robinson).

438.

184

Puccinia — Cruciferae.

A fe

celled teleutospores were ob

among

the

pores, which they resembled i

pore quite dist
25-26 X 20 hi

y respect

d shap

even having the germ
small hyaline apiculus,

Aecidium violae Schum. is ^
occurring on Victorian native violets, but it

P. hederaceae McAlp.

beneath the apex, ellipsoid

given

by D

Cooke in his Handbook as
is the aecidium belonging to

at first determined as P. aegra Grove by Winte
but on comparing the two species they are seen to be decidedly different

species

It is closely allied to P. violae (Schum

teleutospores being

'«

and

ted all over. Sp

but differs in the

of P

eel from Syd. Ured. Exs., No. 286

d from

were examn

Kellerman's Ohio Fungi, No. 96, on Viola blanda.

teleutospores was found to be 22-32 X 16-19 ju, and were generally

The average size of

smooth, although
P. aegra Grov«
mens I have met
forwarded to D

little

ted

ded by Cooke for Australia; but all the speci

th, including part of the same collection of

Cool

echinulate

tospores, and are therefore not this species

diospores

d warted

(Plate XIII., Figs. 109-112 ; Plate E., Figs. 24, 25.)

CRUCIFERAE.

120. Puccinia cruciferae Mc Alp.

I. Aecidia amnhi^enous and o

large,

cup-shaped, §-| mm. diam.; pseudoperidia w
edges, crenulate, slightly revolute ; pseudoperidial cells

th whitish

oblong, p
lonsr.

all over, with

ted margin, average

oid
25

Aecidiospores orange,

subglobose, finely echinulate, 14-16

III. Teleutosori on upper surface of basal leaves, associated with

aecidia, a few together, round, bullate, compact, covered at first

by raised grey ej>idermis, which
diam.

splits

irregularly,

to 1 mm.

Teleutospores black in mass, dark chestnut-brown individually,
oblong to clavate, smooth, slightly constricted at septum, bluntly
pointed, rounded or truncate at apex and thickened up to 9 />/,
tapering slightly or rounded at base, 37-62 X 18-25 p ; pedicels
usually slightly tinted towards apex, persistent, up to 50 /x long and
10 fi broad.

X. Mesospores common, similarly coloured to, or paler than, teleuto-

spores

variously shaped, ellipsoid, oblong

or ovate, pointed,

rounded, or truncate at apex, and thickened up

to 7

25-34 x 14-20 /./, with pedicels equally long and similar to those
of teleutospores.

On Cruciferous plant.

New South Wales

It differs from P. cru

Guntawang (H

P. barb

ifi

Rud

in having

and

Cooke, in which the

derabh

broadly lanceolate

Phragmidium.

185

This is a portion of the original material which was named Aecid

barbareae in Cooke's Handbook oj Australian Funyi but, on closely
examining it, teleutosori were found. The host-plant is not Barbarea
vulgaris R. Br., which is the only species of that genus in Australia.

(Plate XXX., Fig. 266.)

DILLENIACEAE

121. Puccinia hibbertiae McAlp.

Hibbertia

•

III. Teleutosori ruddy-brown to black, compact, soon naked, up to

1 mm. diam., on under surface of leaves.

Teleutospores pale yellowish-brown, oblong to somewhat clavate,
smooth, constricted at septum, 35-41 X 18-21 ft; upper cell
rounded at apex to somewhat conical, usually thickened (up to
6 fi) ; lower cell generally rounded at base, sometimes slightly
attenuated ; pedicel hyaline, persistent, elongated, slender,
120-150 //long.

X. Mesospores occasional, similarly coloured to teleutospores, generally

fusiform and thickened at apex, sometimes elongated oblong,

33-35 x 12-16 p.

On Hibbertia sericea Benth.

Victoria — Cheltenham, July, 1904 (Robinson). Oakleigh, Sept.,

1904.

The small size of the leaf together with the scanty sori render this
species easily overlooked. The teleutospores are often noticed germinating

situ.

(Plate XXX., Pig. 265.)

PHRAGMIDIUM Link.

(

This genus occurs exclusively on the Rose family, and of the 46 species
recorded by Dietel 13, only four are found in Australia. They are all autoe-
cious, and in some instances spermogonia accompany aecidia. The aecidia are
of the caeoma type, but paraphyses take the place of a peridium. The species,
(P. hamardx) on the native raspberry (Rubtie parvifolius) does considerable
iamage, and the imported species — P. subcorticium — is spreading extensively
m the sweetbrier.

General Characters. — Spermogonia flattened or slightly raised, generally
irranged in concentric circles.

Aecidia after caeoma type, surrounded by a dense crown of inwardly-
curved paraphyses.
I Caeomosporcs with numerous distinct germ-pores distributed over the

| entire surface.

Uredosori likewise surrounded by paraphyses.
Uredospores solitary on basidia, with numerous germ-pores.
Teleutospores separate, pedicellate, consisting of from three to ten,
superimposed cells, the uppermost of which has one to three germ-pore
the others about three to four each, placed laterally.

spherical.

G 2

186 Phragmidium — Rosaceae.

ROSACEAE.

Rubus

122. Phragmidium barnardi Plow, and Wint

Winter, Rev. Myc, VIII., p. 208 (1886).

Cooke, Handb. Austr. Fung., p. 339 (1892).
Sacc. Syll. VII., p. 746 (1888).

O. Spermogonia on numerous, minute, discoloured patches on upper

surface of leaf, appearing as ruby coloured, minute, blister-like
puckered swellings, with round central pore, and arranged in
small concentric circles.

Spermatia golden-yellow in mass, hyaline individually, minute,
narrowly ellipsoid, embedded in mucilage, 4-6 x 2J-3 ix.

II. Uredosori scattered on under surface of leaf, corresponding to

spots on upper surface, very irregular in outline, ochraceous,
also sparingly on upper surface of leaf, usually on veins, ellipsoid
to elongated ellipsoid, surrounded by ruptured epidermis, and pale
in colour.

Uredospores globose or ellipsoid, epispore • thick, warty, dark
yellow, 17-19 fi diam. or 21-26 x 17-19 fx ; paraphyses surround-
ing them, at first colourless, finally yellowish-brown, variously and
peculiarly shaped, generally curved and pointed at apex, 30-40 X

9-12

III. Teleutosori scattered or gregarious, minute, dark brown, not

rarely confluent, pulverulent.

Teleutospores at first intermixed with uredospores, golden-brown,
cylindrical, equal, rounded at apex, sometimes with a short,
hyaline, conoid apiculus, 6-9 septate (8 very common), constricted
at septa, smooth, usually two germ-pores on one face in each cell,
or three altogether, 60-115 x 23-28 /u ; pedicels hyaline, cylin-
drical, or a little inflated, sometimes uniseptate towards base, very
long, up to 207 x 10-19 li.

On living leaves of Native Raspberry (Rubus farvifolius L.).

Victoria — Alps near Bright, Arthur's Creek, Flinders, Killara,

Murramurrangbong Ranges, Myrniong, near Melbourne
Phillip Island, Seymour, &c.

South Australia — Mount Lofty (Tepper

Tasmania — Huonville, Jan., 1892 (Rodway1). Mount Wellington

Jan., 1892 (Rodway).

As early as 1886 this species was determined by Plowright and Winter
from specimens sent by that veteran microscopist, Mr. F. Barnard, Kew,
Victoria.

The teleutospores occur on the same leaves as the uredospores, and they
germinate on the living plant immediately on ripening. The germ-pores in
each cell may be seen very clear and distinct. There are two in each seen
on one face, situated laterally an(I immediately below each of the septa, and
by means of careful focussing it can be seen that there is one behind, thus
making three in each cell. In the apical cell there are also two just above
the septum, and a third may also be seen as in the other cells.

Dietel ] remarks that there is only one germ-pore in each cell of this
species, but, as indicated above, three can usually be seen with such dis-
tinctness that they are capable of being photographed, as shown in the
Figures.

Phragmidium — Rosaceae

i87

Teleutospores were found germinating on the living plant. The promy-
celium was stout and consisted of four cells with a basal cell, the contents
being greyish like the sporidiola. The sporidiola are subglobose and 7-8 u
diam.

This fungus does considerable damage to the native raspberry, causing
the leaves to fall prematurely.

(Plate XXVI., Figs. 234, 235 • Plate I., Fig. 38.)

123. Phragmidium longissimum Thuem.

Thuemen, Flora, p. 379 (1875).

Rubus

Cooke, Handb. Austr. Fung., p. 340 (1892).
Sacc. Syll. VII., p. 750 (1888).

Hamas'pora longissima Koern. Hedw. XVI., p. 23 (1877).

II. Uredosori hypophyllous, scattered or gregarious, or even confluent,

clear orange, surrounded by a dense layer of paraphyses.

Uredospores globose or obovate or shortly elliptical, pale yellow,
with thick epispore, 16 fi diam.

III. Teleutospores on both surfaces of leaf, agglutinated in long fila-
ments up to 6 mm. long, rather gelatinous when moist, but when
dry twisted up and silky-fibrillose, 4 to 6 celled, commonly 4-celled,
yellowish to cream-coloured, containing granular protoplasm,
cylindric acuminate, with apex sharply pointed and hyaline,
110-190 x 12-15 fx ; pedicel hyaline, tapering gradually towards
base to a fine point, hollow, just slightly narrower than spore,
10-12 a broad, and reaching a length of 500 u.

On leaves of Rubus molnccanus L.

Queensland — Eudlo Creek (Bailey

This species was first observed on Rubus in S. Africa, and destroys
the foliage of one of the blackberries indigenous to Queensland. Bailey
kindly sent me specimens in which the teleutospores are germinating in
situ.

Scattered over the leaves are groups of silky-looking twisted filaments,
and on being teased out are seen under the microscope to consist of innumer-
able spores with their long pedicels inextricably blended.

The germinating spores are seen to have one germ pore in each cell, and
the order of germination is basipetal. The promycelial branches are stout,
elongated, and transversely septate, and they may either all arise on one
side of the spore, or partly only on one side. The promycelial spores are
subglobose, finely warted and 9-10 fj. diam.

The teleutospores were very commonly 140 /< long, and one 6-celled in-
dividual attained a length of 190 ja. As might be expected in such a long
and variously septate spore there is considerable difference in the lengths
recorded. It was originally given as 200-240 /u, then Massee reduced

0-130 fi, and in the Queensland specimens I have found it to be 1 10—190 ^i
The stalks are excessivelv long, and they may reach a length of 500 /

mor

The teleutospores agree with those of Phragmidium in bei

more transversely septate, but they are agglutinated together into a

cylindrical filamentous mass, and each cell only

pore

i88

Phragmidium — Rosaceae.

Massee3 from an examination of fresh material comes to the conclusion
that it is not a Phragmidium nor a Hamaspora, although he does not under-
take the responsibility of forming a new genu?.

Dietel 8, 10, on the other hand, considers that there are no substantial
grounds for separating this species from Phragmidium, although he recog-
nises a resemblance to Gymnosporangimn in the arrangement and structure
of the teleutospores.

(Plate XXXI., Fig. 272.)

124. Phragmidium potentillae (Pers.) Karst.

Karsten, Fung. Fenn., No 94 (1868).
Winter, Rev. Myc. VIII., p. 208 (1886).

Acaena-

Cooke, Handb. Austr. Fung., p. 339 (1892).
Sacc. Syll. VII., p. 743 (1888).

, Caeomata arranged in circular groups, often confluent, orange yellow
to reddish.

Caeomospores crowded in a short series, globose, ovate or ellipsoid,
echinulate, 17-24 x 14-19 ft.]

II. Uredosori hypophyllous, orange-red, roundish, scattered or

gregarious, often confluent, surrounded by a crown of club-shaped
paraphyses.

Uredospores spherical, elliptical or ovate, yellowish, echinulate,,
17-24 X 14-20 A/, average 18 x 15 /i.

[III. Teleutosori orbicular, black, pulvinate, minute.

Teleutospores oblong, 2-6 septate (usually 3-5), slightly con-
stricted, obtuse, or obtusely apiculate above, slightly attenuated
below, smooth, yellowish-brown, 50-70 X 20-22 \i ; pedicels
colourless, firm, 100-150 \i long.

On living leaves of Acaena sanguisorba Vahl.

Victoria — Near Melbourne, 1886 (Reader). (Winter.2)
On Acaena ovina A. Cunn.

S. Australia — Sept., 1898 (Quinn).

The specimen from Victoria was originally determined by Winter, an
portion of the same material, kindly supplied by Reader, shows only
uredospores.

Darlucajilum Cast, is plentiful on some of the sori.

125. Phragmidium subcorticium (Schrank) Winter.

Winter, Die Pilze I., p. 228 (1884)

Cooke, Handb. Austr. Fung., p. 339 (1892)
"--- Syll. VII., p. 746 (1888).

Rosa

Caeomata forming large dense

swellings on stems, leaf -stalks, and
fruits, and lesser swellings on lower surface of leaves, effused,
generally confluent, bright orange, with club-shaped paraphyses.
Caeomospores ellipsoid to ovoid, echinulate, epispore hyaline, and

contents orange, 25-29 x 14

■I

Cronartium.

189

II. Ured

hypophyll

orbicular, scattered or crowded

yello

pots, yellowish red, minute

tubul

Phy

ds apex, and not particularly club-shaped

htly

Uredospores ellipsoid, ovoid or p
eddish orange. 4-6 germ-pores on one face

finely ech

29 x

III. Teleutosori hypophyll

readily detachable.

minute, scattered or

6-20 p.
ded. black

Teleutospores

th a colourless apiculus. 7-8 celled

oblong, dark-brown, warted, obtuse at the apex

face in each cell. 7

x 3

ped

3 germ-pores

hyaline

one
Pt

pale-brown towards apex, expanded in the lower half, 100-150 y
long and broadened up to 22 u.

On Rosa rubiginosa L., R. canina L., and R. laxa Ret

Victo

Pascoe Vale, Oct., 1898 (C

April, 1899 (H
Oct. and Dec.

Gully

il). Brighton, Nov., 1901, Aug., 1903, June
904. March. 1905. II.

S. Australia — Third Creek Garden, Mt. Lofty Range, Nov., 1899

(Tepper). Nov., 1900 (Agricultural Bureau), IL, III.

Tasmania — Launceston, Oct., 1902 (Littler). Hobart, Apr., 1903,

IL, III. (Rodway).

This species is now very common around Melbourne, particularly on the
sweetbrier (R. rubiginosa). It has also become a pest in the nurseries,
causing considerable damage to the dog-rose stocks (R. canina). Seedlings
are attacked very severely in a favorable season and killed outright. This
rust occurs in some of its stages, practically all the year round, but is less
common in midsummer. The uredospores resemble the caeomospores in their
size and shape, but may be distinguished by their long stalks (up to 56 /<)
and their germ-pores.

The mycelium of the caeoma may winter in the stems, so that by means

of cuttings the disease was probably introduced into Australia.

may

also have been introduced with the seeds of the sweetbrier, for that was
among the first European plants to be imported into Tasmania and New
South Wales, and used as hedges.

(Plate XXVI., Figs. 229-233 ; Plate I., Fig., 37.)

Only

CRONARTIUM Fries.
s senus is known here, and

monly on certain leguminous plants. So far

found

This genus produces uredo and teleutospores on

stage

other, but the latter h

have been
one host plant,
s not yet been

Europe there is a bladder-rust on the bark of

and the aecidial

found in Australia.

sylvestris, and this has been proved to be the aecidial stage of Ci

asclepiadeum (Willd.) Fr. As in so many other similar cases

considered to be an independent form, and named Peridermium cornui

Rostr. and Kleb.

stage was

General characters
Aecidia with bladder-like

— Spermogonia truncate to conoid

mium)

ly opening pseudoperidia (P

Aecidiosp

thout crerm-

parated by d

mediate

epispore always more or less regularly warty reticulated.
Uredo-layer included in hemispherical pseudoperidum, which opens at ape
by a narrow pore.

190

Cronartium — Legnminosae.

Uredospores echinulate, without germ-pores.

Teleutospores unicellular, united into a cylindrical, vertical, horny column,,
germinating in situ, surrounded at base by the uredospores.
Sporidiola subglobose, hyaline.
Australian species, 1.

LEGUMINOSAE.

126. Cronartium jacksoniae P. Henn.

Jacksonia, dec.

P. Hennings, Hedw. XL., p. 127 (1901).

O. Spermogonia honey-coloured, crowded or in lines, circular, flattened,.

on branches and lower surface of leaves.

Spermatia hyaline, avoid to ellipsoid, 5-6 X 2|-3 p.
III. Teleutosori forming ruddy-brown, straight or curved, simple

columns, crowded, 2-2 \ mm. high, with blunt apex, on young and
swollen branches.

Teleutospores ruddy-brown in mass, more or less colourless indi-
dividually, elongated, generally somewhat tapering towards each
end, with large vacuole, cylindrical to fusiform, contents granular,
40-50 X 8-10 fi ; teleutospores germinate at once and produce
promycelia with promycelial spores, which are hyaline, globose,
and average about 6-7 \i diam.

On Jacksonia scoparia R. Br.

Queensland — Darling Downs (Law), (Bailey2). Near Brisbane

(Bailey4).

On Platylobium formosum Sm.

Victoria

Murramurrangbong

Ranges, Nov.

and Dec, 1902-3

(Robinson).
On Gompholobmm latifolium Sm.

Victoria — Murramurrangbong Ranges Dec, 1903 (Robinson).

On Bossiaea cinerea R. Br.

Victoria — Near Melbourne, Nov., 1904 (Robinson)
Tasmania — Bellerive, Dec, 1905 (Rodway).

On Actus villosa Sm.

Victoria

Near

(Robinson).

Melbourne,

Sept.,

Nov.,

1904,

III.

It deforms the shoots of the

plant on which it grows,

causing mal-

formation of branches and multiplication of shoots (witches' brooms), and
so common in some parts that the great majority of the

shrubs are

ttacked

The spermogonia were only found on Aotus and before the appear

of the teleutosp

the multiplication and deformation of the shoots

had already been produced. On Bossiaea cinerea on one occasion Aeeid
eburneum McAlp. was found on the same plant near Melbourne.

The horn-like aggregation of teleutospores varies in length from 1-2 mm.
on Aotus and Bossiaea to 2-2^ mm. in Platylobium and Gompholobium.

The original specimen on Jacksonia scoparia in the National Herbarium,
Melbourne, is given as a variety of C. asclepiadeum (Willd.) Fr., in Berkeley's
handwriting, and is recorded in the Journal of the Linnean Society, p. 174
(1883), with the remark that the only specimens are in a young state. The
same species was subsequently found by G. H. Robinson very plentifully in
Victoria

on various species of Leguminous pi

and the germiuating

teleutospores were

rly

Melampsora — Hypericaceae. 191

Conside

•ing its occurrence on indigenous Leguminous plants, together
with the narrower teleutospores and colourless promycelial spores, as com-
pared with G. asclepiadeum, I considered it a new species and had named it
C. leguminum. But I find that Hennings in a footnote in Hedwi^ia XL.
p. 127, states that the form occurring on Jacksonia is specifically distinct
from G. asclepiadeum, and had better be named C. jacksoniae, the name now
adopted.

(Plates XXXVIL, XXXVIII.)

MELAMPSORA Castagne.

In this genus there are both heteroecious and autoecious species, but only
the latter occur in Australia. The two species known are on native plants,
but they also occur in Britain on the same genera. They only possess uredo
and teleutospores, but the three spore-forms may occur on the same host
{M. amygdalinae Kleb.), or only aecidia and teleutospores (M. saxi-
fragarum (DC.) Schroet.).

The teleutospores form flat irregular crusts, and are united to each other
like the cells of a honeycomb.

Rostrup first pointed out in 1883 the connection between this genus and
Caeoma species, and now the relation between a number has been proved by
cultures.

General description. — Spermogonia forming a minute, hemispherical,
flattened stratum, often subcuticular, otherwise subepidermal.

Aecidia after the caeoma type, destitute of a pseudoperidium, usually
without paraphyses, pustular.

Uredospores usually without distinct germ-pores, included in more or less
developed peridia.

Teleutospores unicellular, rarely transversely divided, coalescing in a
plane, firm, black or dark-brown stratum. Sporidiola globose, yellow, then
orange.

Australian species, 2

HYPERICACEAE.

127. Melampsora hypericorum (DC.) Schroet.

Schroeter, Brand und Rost., p. 26 (18
Sacc. Syll. VII., p. 591 (1888).

Hypericum

II. Uredosori mostly hypophyllous, scattered or subgregarious, at first

bright orange, becoming pale, pulverulent, small, up to £ mm.
diam, erumpent and surrounded by the ruptured epidermis.

Uredospores subglobose to ellipsoid, finely verrucose, orange
yellow, with two germ-pores on one face, 14-21 x 11-17 ji,
average 17 x 14 /* ; paraphyses intermixed, numerous, hyaline,
capitate, over-topping the spores, 50-68 p. long, with head 18-24 p.

III. Teleutosori minute, dark-brown, flat, irregular, solitary or gre-
garious.

Teleutospores yellowish-brown, intercellular, subcylindrical or

prismatic, 25-37 + 6-9 u.
On leaves and occasionally on stems of Hypericum j aponicum Thunb.

Victoria — Murramurrangbong Ranges, Nov. and Dec, 1902-3, and

M"t. Blackwood, Jan., 1903 (Robinson). Buffalo Mts. and
Alps, near Bright, Nov. and Dec, 1903-4 (C. French, jr.).

Near Melbourne, Nov., Jan

192

Melampsora — Linaceae.

The paraphyses are noted as absent by Dr. PIov

Mono

of the Uredineae, but in these specimens they are particularly abundant, and

form a striking feature of the uredosori

Fischer 5 also found the paraphy

had bee
Klebahn

and he came to the conclusion with others
l mistaken for the uredo, since the spor

that the caeoma form

But

proved

of both the caeoma form without

physes and the uredo form with paraphy

so that the th

stages

may

species of Hyp

Aecidium disseminatum Berk, is found he

same host-plant and in the same locality as this species

LINACEAE.

mum*

128. Melampsora lini (Pers.) Tul.

Tulasne, Ann. Sci. Nat,, p. 93 (1854).
Cooke, Grev. XL, p. 98 (1883).
Cooke, Handb. Austr. Fung., p. 332 (1892).
Sacc. Syll. VII., p. 588 (1888).

Uredosori scattered, rounded, orange,
mm. diam.

11.

soon pulverulent, up to 1|

•_>

Uredospores round or ovate, bright orange-yellow, echinulate,

a length of

pedicellate, 15-25 x 13-18 fxy exceptionally reaching

28 fi ; paraphyses curved, markedly capitate, 17-20 fi thick.

III. Teleutsori flattened, often confluent, reddish brown, then black,

shining.

Teleutospores densely crowded beneath epidermis, cylindrically
prismatic, intercellular, polygonal in section, 45-60 x 17-20 //,
very occasionally two-celled.

On leaves and stems of Linuni marginale A. Cunn. Widely distributed.

Vic to

Near Melbou

1885

(Reade

Ardmona

Kergunyah, Killara, near Melbourne, Rutherglen, &c, Oct

March.
South Wales — Hume Ri

1886 (Jeph

Mudgee (Hamilton) ; Merilla, Oct., 1890 (Cobb

Guntawang,

South Australia — Murray River, 1890 (Tepper), (Ludwig2)

Tas

Near Waterworks, Hobart, Jan., 1892 (Rodway1)

On Linum usitatissimum L. wherever flax is cultivated

Victo

Near Melbourne. 190

Donald. Nov.. 1903. II

III

Port Fairy, Jan., 1904, II., Ill

South Australia

New South Wales— Brungle, 1891 (Cobb4)

—Near Adelaide, 1889 (Crawford)

Journal of

1889. G

ecords this

t as

being sent to him from South Australia by the late Frazer S. Crawford, who

pread

that it had destroyed a crop of flax near Adelaide, and was likely

d prove a troublesome pest

forms me by letter that

Linum. and

in Dakota it is a very abundant rust on all the wild varieties of Linum,
is always more or less destructive in the flax crop. He further states that
it was especially destructive in 1904, in many cases practically destroying
crops which he had bred from seeds supposed to be immune to the wilt disease
or dax-sick soil disease.

Dr. Cobb4 in the Agricultural Gazette of New South Wales for 1891,
notes it as causing serious injury to cultivated flax, and in some parts of
Victoria, particularly at Port Fairy, I have found it injuring the crop.

(Plate XXVI., Fig. 236 ; Plate I., Fig. 36.)

Caeoma — Apocynaceae.

*93

CAEOMA Link.

There are several imperfect forms which only occur in one stage and
cannot with certainty be assigned to their proper genera. For such it is
convenient to have a form-genus, which simply serves as a resting-place until
their true affinities are determined. At first they were considered to be
independent fungi and had generic names assigned to them. These form-
genera are Caeoma, Aeeidium, and Uredo, and of the former there are *nlv
two known for Australia, one of which was called an Aeeidium by Cooke

and the other a Uredo by Berkeley. This is not surprising when one
considers that the caeoma is not distinct from the aeeidium, as in those
cases where the one merges into the other. Thus in Puccinia prenanthis
(Pers.) Lindr., the aecidial wall is very imperfectly formed, so that the
aecidia approach caeoma-forms. Barclay3 also found a variety of this
species in Simla, in which there was not a vestige of a peridium, and
he considers this, along with others, an interesting example of an inter-
mediate and mostly vanishing stage between Aeeidium and Caeoma. In
the aecidial stage of Puccinia aucta (Aeeidium lobelias Thuem., A. micro-
stomum Berk.), the peridial wall was sometimes absent, so that I described
it at first as a Caeoma.

Caeomata are generally considered to be stages in the life-history of the
Melampsoraceae, but since true caeoma-forms occur in connexion with
Puccinia, and species such as Melampsora tremulae Tul. have so-called
caeoma-forms as Caeoma laricis (Westd.) Hart, with an investment of
barren cells, the evidence for this connexion is weakened and the necessity
for retaining this as a form-genus much reduced. In Saccardo's Sylloge
Fungorum the isolated forms are treated as a sub-genus of Uredo with sub-
"Catenulate spores. They have no special significance here, but elsewhere
they form witches'-brooms.

General Characters. — They are simply aecidia without peridia, the spores
are produced in chains, with or without paraphyses, and accompanied by
spermogonia.

Caeoma-forms, 2.

APOCYNACEAE.

Tahernaem on tana

129. Caeoma apocyni McAlp.

I. Sori on both surfaces of leaf, minute, crowded, brownish, bullate.

Spores yellowish, very variable in shape, ellipsoid, ovoid, pear-
shaped, oblong or angular, with finely granular contents, 25-34
x 20-26 /j. ; epispore hyaline, coarsely verrucose, 5 /x thick.

On Tabernaemontana orientalis R. Br.

Queensland — Brisbane, (Bailey1).

Mr. Bailey kindly sent me a specimen of this from his herbarium,
which had been named by Dr. Cooke Aeeidium apocyni Schwein., as
given in his Handbook of Australian Fungi, p. 341 (1902). It has no
pseudoperidium, however, and is therefore a Caeoma, and since the spores do
not apparently agree even with those of Caeoma (Aeeidium) apocynatum
Schwein., it is constituted a new species.

194

Aecidium — Scrophulariaceae

RANUNCULACEAE.

Clematis

130. Caeoma clematidis Thuem.

Thuemen, Myc. Univ. No. 539.

Berkeley, Hook. Journ. VI., p. 205 (1854).

Cooke, Handb. Auatr. Fung., p. 344 (1892).
Sacc. Syll. VII., p. 867 (1888).

Uredo clematidis Berk.

I. Sori hypophyllous, generally on pale-green portions of leaf, solitary o

ged in circular groups, more or less rounded, pulvinate, ruddy

brown at first and bright orange when b

in rows,

bright

orange,

able

shape, ellipsoid

oblong or polygonal, epispore hyaline, scolloped, 2

of Clematis aristata R. Br., and C. microphylla DC

Victoria— Near Colac, Sept., 1897 (H

1900

Murramurrangbong

Ran

Dec

Phillip Island, Jan.
, 1903 (Robinson)

Near Melbourne, Feb. and July, 1904. Frankston, Jan., 1904, *
Aug., 1904 (Robinson).

Queensland

Bailey

This species was first described from S. Africa as a Uredo, but it has the
spores in chains and is a true Caeoma. Specimens were examined from
Thuemen's Myc. Univ. 539 and found to be the same.

AECIDIUM Pers.

Several of the aecidia

given

in Dr. Cooke's Handbook of Australian

Fungi are now referred to their proper Puccinias. A. urticae is accepted
as a stage in the life-history of Puccinia caricis, and described in connexion
with that form. A. goodeniacearum, A. senecionis and A. violae are placed
under their proper Puccinia or Uromyces, and the aecidium on the daisy
(A. bellidis) is now found to belong to P. distincta McAlp., and to have no
connexion, as in Britain, with P. obscura on Luzula. The most noteworthy
aceidium is that found on a grass (A. danthoniae), and constitutes the second
known example. They often cause deformation of the branches known as
witches' brooms, and various swellings, and in some instances compound
galls are formed up to 5 cm. across (A. englerianum). One species causes
thickening and distortion of the leaves in Veronica, and nettle stems are
often abnormally swollen from the same cause.

General Characters. — Pseudoperidia cup-shaped or urn-shaped, rarely
cylindrical, with margin often crenate or laciniate and re volute.

Spores, globose or angular, commonly orange yellow, growing in chains*

Aecidium-forms, 15.

SCROPHULARIACEAE

Veronica.

131. Aecidium disciforme McAlp.

O. Spermogonia on under surface of upper and younger leaves, honey-
coloured, flat, numerous, minute.

Spermatia hyaline, elliptical, 5-6 /x long.

Aecidium — Plantaginaceae 195

I. Aecidia on both surfaces of leaves, but mostly on under, bullate,

brownish, resembling dotted discs, often confluent, at first entirely
enclosed, ultimately opening irregularly by small pore, 1-1 A mm. ;
peridial cells hyaline, loosely adherent, subglobose, elliptic or
lozenge-shaped when united, with striated margin, 25-30 fj.

Aecidiospores orange-yellow, subglobose to elliptic, 16-20 a diam
or 19-22 x 14-16 /u.

On Veronica gracilis R. Br.

Tasmania — Sandy Bay, Hobart, Aug., 1896 (Rodway).

On Veronica calycina R. Br.

New South Wales— Kogarah, Oct., 1900 (Maiden).

This species causes thickening and distortion of the leaves. It is quite
distinct from A* veronicae Berk., of which there is a portion of the original
material in the National Herbarium, Melbourne. The latter forms a regular
cup with fimbriate margins, while the former only opens by a minute pore.
The spores are also quite different.

132. Aecidium veronicae Berk.

Veronica

Berkeley in Herb., Grev. XL, p. 97 (1883).
Sacc. Syll. VIL, p. 814 (1888).

I. Aecidia numerous, crowded, semi-immersed, on under-surface of leaf

and a few scattered on upper surface, \ mm. diam.; pseudo-
peridia cup-shaped, becoming discoloured and dark-brown with
age, with fimbriate margins ; peridial cells, oblong to angular, with
very broad striated margin, average 32 x 25 it.

Aecidiospores dark yellow, variously shaped, ellipsoid, oblong or
angular, with finely-granular contents, 24-30 x 16-20 fx ; epispore

hyaline, finely verrucose, about 2 it broad.

On Veronica sp.

Victoria — (F. v. Mueller).
Tasmania — (Rodway).

There is an old specimen in the National Herbarium which had been

submitted to Berkeley by the late Baron von Mueller. There are plenty of

■ spores still in the cups, but probably the thoroughly mature ones have fallen

away

(Plate XXXIX., Fig. 287.)

PLANTAGINACEAE

133. Aecidium plantaginis-variae McAlp

Plantag

0. Spermogonia on both surfaces of leaf, but mostly on upper, honey-

coloured, numerous, 160-180 li diam.
Spermatia hyaline, elliptical, 4-5 li long.

1. Aecidia on both surfaces, sub-gregarious or gregarious ; pseudoperidia

cup-shaped, outstanding, with recurved and lobed margins;
peridial cells oblong to lozenge-shaped, or polygonal, with broad

striated margin, 35-40 li long.

Aecidiospores bright orange-yellow, elliptic to oblong, finely
echinulate. 22-25 x 16-23 u, sometimes reaching a length of 28 u.

196

Aecidium — Gentianaceae, Comfo sitae.

On leaves of Plantago vnria R. Br.

Victoria — Murramurrangbong Ranges, Nov., 1902 (Robinson).

Mt. Blackwood, March, 1905 (Cyril Brittlebank).
New South Wales — Guntawang, near Mudgee (Hamilton).
Tasmania — Knocklofty, Oct., 1895 (Rodway 1).

This species is

given as A.

plant aginis Ces., in Cooke's Handbook of

Australian Fungi, but there are no spots on the leaves, and the spores are
echinulate not tuberculate, as well as considerably larger.

The Victorian specimens were rare, and the fungus was much overrun by

Darluca filum Cast.

(Plate XXXIX., Fig. 288.)

GENT1ANACEAE.

134. Aecidium nymphoidis DC.

Limnanthemum

De Candolle, Fl. fr. II., p. 597 (1805).
Cooke, Handb. Austr. Fung., p. 341 (1892)
Sacc. Syll. VII., p. 809 (1888).

I. Aecidia on upper surface of leaf, gregarious, disposed without order

on rounded spots, or in concentric zones ; pseudoperidia scutelliform,
margin scarcely prominent, entire, or irregularly laciniate.

Aecidiospores orange-yellow, angular, delicately warted, 12-20
ju diam.

On Limnanthemum indicum Thw.

Queensland — Near Brisbane, 1884 (Bailey

This aecidium was first found on the leaves of Limnanthemum nymphoides
Hoff. and Link and its connexion with Puccinia scirpi DC, was first sug-
gested by Chodat1. Then Bubak1, by means of cultures, proved the
connexion, but until the teleuto stage is found
aecidium stage will be given.

in Australia only the

Specimen not seen.

COMPOSITAE.

135. Aecidium cymbonoti Thuem.

Cymbonotus.

Thuemen in Muell. Supp. Phvt. Austr. XI., p. 96 (1880).
Sacc. Syll. VII., p. 833 (1888).

0. Spermogonia minute, honey-coloured, in groups, surrounded

aecidia, 130-160 /i diam.

Spermatia minute, hyaline, ellipsoid, 3 fx long.

1. Aecidia on upper surface of leaves, markedly circinate, about ^ mm.

diam.; pseudoperidia yellowish, with fimbriate margin ; peridial cells
considerably overlapping each other, elongated oblong to slipper-

ma rgm

and reticulate all over, 28-38 X

shaped, with striated
16-17 ^.

Aecidiospores orange-yellow, ellipsoid to sub-globose, finely
echinulate, 13-18 x 13-15 fi.

Aecidium — Compositae. 107

On Cymbonotus lawsonianus Gaudich.

Victoria— Dookie, Jan., 1892 (Pye). Dimboola, May, 1896

(Reader). Preston, April, 1900 (C. French, jun.). Myrniom
(C. C. Brittlebank). J *

Some of the specimens were overrun with Darluca filum Cast.
There is no clue to the description of this species, but I have drawn oul
the above from abundant material.

(Plate XXXIX., Fig. 290

136. Aecidium monocystis Berk.

Abrotanella.

Berkeley, Flor. N.Z. II., p. 196 (1855).

McAlp., Proc. Roy. Soc. Vic. VII., N.S., p. 218 (1894)

Sacc. Syll. IX., p. 319 (1891).

I. Aecidia solitary, relatively large (J mm. diam.) persistent on the upper

surface of the leaves towards their tips, surrounded with a tough
border ; pseudoperidia white, with margin toothed ; peridal cells
somewhat wedge-shaped to angular oblong, appearing hexagonal
when united, covered with projecting points, 50 x 22 p

Aecidiospores pale orange-yellow, ellipsoid, finely verrucose,

25-30 x 19-22 H.

On leaves of Abrotanella J or sterioides Hook. f.

Tasmania — Summit of Mt. Wellington, Feb. 1891 (Rodway).

This is a very characteristic species, occurring solitary at the tips of the
leaves. It was first recorded on Phyllachne (H el ophyllum) coleusoi Berggr.
from New Zealand, but L. Rodway, Government Botanist of Tasmania, has
grave doubts as to the correct determination of the host-plant. He is of
opinion that this rust is attributed to New Zealand by mistake, and that
it was really found in Tasmania. He informs me that this Aecidium is
abundant on every tuft of A. for sterioides, which grows in profusion near
Hobart, and is not likely to have been overlooked by the early botanists.
The coloured drawing of the plant in the New Zealand Flora certainly closely
resembles Abrotanella, and I have forwarded a specimen of the plant to the
Director of the Royal Gardens, Kew, for comparison with the original. It
is not likely that the same Aecidium occurs on a Composite and a

Stylidiaceous plant.

(Plate XXXIX., Fig. 291.)

Olearia

137. Aecidium oleariae McAlp

I. Aecidia crowded on stems and leaves, generally running in lines

lengthwise, about 1 mm. ; pseudoperidia round to elongated, with
lacerated margins ; peridial cells hexagonal, striated, 35-42 p long.
Aecidiospores orange, elliptic to oblong, with granular contents,
22-26 x 14-17 /a.

On stems and leaves of Olearia axillaris F.v.JVL

Victoria — Port Fairy, June, 1899 (Robinson).

The aecidia occur most commonly on the stem, and are deeply imbedded
in the tissues of this coast shrub.

198

Aecidium — Rubiaceae, Leguminosae.

RUBIACEAE.

Can thium

138. Aecidium plectroniae Cooke.

Cooke, Grev. X., p. 124 (1882)

Bailey, Queensland Agr. Journ. IV., p. 284 (1899).
Sacc. Syll. VII., p. 795 (1888).

O. Spermogonia on upper surface of same spots which bear the aecidia,

sometimes accompanied by one or a few aecidia.

I. Aecidia hypophyllous, rotund, in small clusters on orbicular dis

coloured spots ; pseudoperidia somewhat prominent, whitish, mar

ovate to somewhat quadrate

gins sub-entire ; peridial cells
punctate all over, and with striate mar

Aecidiospores yellow, finely
polygonal, 25-'

ub-globose to ellipsoid

6 x 19

2-28 u diam

On leaves of Canthium coprosmoides F. v. M.

Queensland — Endeavour River (Roth) (Bailey, 21, 25

The host-plant is sometimes placed under the genus Plectronia.

This species was first found on Canthium (Plectronia) guenzii Sond.,
from Natal, and the spores were undescribed, but I have been able to
redescribs it from specimens kindly sent from Queensland by Bailey.

LEGUMINOSAE

Bossiaea

139. Aecidium eburneum McAlp.

McAlpine, Proc. Rov. Soc. Vic. VII., N.S., p. 218 (1895).
P. Hennings, Hedw. XL., p. 352 (1901).
Sacc. Syll. XIV., p. 375 (1899).

A. bossiaeae P. Hennings, Hedw. XL., p. 352 (1901).

I. Aecidia mostly hypophyllous or on stems and legumes, scattered or in

dense clusters, bright orange, average £ mm. diam.; pseudoperidia

cup-shaped, becoming ivory-coloured then brownish, with reflected

finely toothed margin ; peridial cells polygonal, reticulated, 24-31 /x
long.

Aecidiospores ellipsoid, bright orange, very finely echinulate,
18-28 x 12-17 /i.

On Bossiaea drier ea R. Br., stems, leaves, flower-stalks, calyx, and

legumes.

Victoria — Near Melbourne, Sept.-Nov. (Barnard, Robinson, &c).
Tasmania — Bellerive swamp (Rod way !), and Dec, 1905.

On leaves of Bossiaea linophylla R. Br.

West Australia

— King George's Sound, July, 1901 (Pritzel).

On fruit of Bossiaea rhombifolia Sieber and B. microphylla Sm.

New South Wales — Richmond, Nov., 1896, and Falconbridge,

Oct., 1904 (Maiden).

On fruit of Bossiaea heterophylla Vent.

New South Wales, Sep., 1896 (Maiden).
I have revised the description of A. eburneum from fresh specimens, and

have no doubt but it is the same as Hennings' species, a specimen

ich

.•

Aecidium — Leguminosae. 190

he kindly sent me. The spores in both cases are very finely eehinulate, and
in the West Australian specimens they are 18-23 x 15—18 /u, while in the
other they are 18-28 x 12-17 p.

On the stems of B. cinerea there are oval to lenticular swellings, some-
times half an inch long, separate or close together, with dark purplish
border, and often becoming depressed in the centre. On the leaf-stalks the
swelling may completely surround them, and on the leaf there are brown
raised orbicular spots, with aecidia on both surfaces, but sparingly on upper.
On the flower-stalks the swellings are similar to those on the leaf-stalks, and
on the legumes large circular or irregular patches are formed on margins and
both valves.

(Plate XXVII., Fig. 239.)

Platylobium.

140. Aecidium platylobii McAlp.

0. Spermogonia honey-coloured, minute, scattered, either associated

with or on opposite side to aecidia.

Spermatia very numerous, hyaline, cylindrical, 3 X 1 jx.

1. Aecidia on pallid spots or on swellings on pods, amphigenous, in

small groups or in dense clusters, deep orange ; pseudoperidia
cylindrical, white, with small portion inserted in matrix, margin
finely toothed, up to 1 mm. long.

Aecidiospores orange, elliptic to sub-angular, finely warted,
21-25 x 14-18 /u.

On leaves and legumes of Platylobium formosum Sm.

Victoria — Murramurrangbong Ranges, Nov., 1902, Dec. 1903, and

Jan. 1905 (Robinson).

It is closely related to Aecidium solenii/orme Berk., but until further
stages are found it will be retained as a distinct species.

While the bright orange aecidia are very conspicuous and very plentiful
on both surfaces of the legumes, they are very rare upon leaves, ha\

hitherto only been found on the first or second pair of leaves of young

seedling

The mycelium bearing the aecidia penetrated the fruit cover and entered
seed, so that the young plant'is probably infected from the start.

(Plate XXVII., Figs. 237, 238.)

Goodia.

141. Aecidium soleniiforme Berk.

Berkeley, Fl. Tasm. II., p. 270 (1860).
Cooke, Handb. Austr. Fung., p. 340 (1892).
Sacc. Syll. VII., p. 788 (1888).

I. Aecidia on brown orbicular spots, sometimes densely crowded, par-
ticularly when on fruit ; pseudoperidia cylindrical, elongated up to
^ mm., white, radiately laciniate at margin.

Aecidiospores rather angular, orange, 25-26 /u diam.

On leaves and fruits of Goodia lotifolia Salisb.

Victoria— Nov., 1895 (French). Gellibrand River, Dec, 1895

(Hill).
Tasmania— 1 8 6 0 (Archer). (Berkeley1)

The aecidia, as the specific name denotes, are rather tubular at first.

(Plate XXXIX., Fig. 293.)

200

Aecidium — Amarantaceae, Hyfericaceae

AMARANTACEAE.

142. Aecidium deeringiae Cooke and Mass.

Deeringia.

Cooke and Massee, Ann. Rep. Dept. Agr. Queensland, p. 40

(1893).

0. Spermogonia on discoloured spots on the upper surface.

1. Aecidia crowded on orbicular pallid spots (2-3 mm.) on the under

surface, which are somewhat thickened, semi-immersed, margin
white, crenulate.

Aecidiospores sub-globose, very finely echinulate, 12 fi diam.

On living leaves of Deeringia relosioides R. Br.

Queensland — Gladfield (Gwyther), (Bailey.12)

The colour of the spores could not be determined in the dried specimens.

Only the name of this fungus was given in the Annual Report, and Dr.
Cooke has kindly furnished me with the description in MS.

Uromyces deeringiae Syd. occurs on Deeringia indica in Java and the
Philippine Islands, but the aecidiospores are 22-28 /x diam., so that they
are quite distinct from the present form unless there has been a mistake in
the transcription of the size of the spores. The host in both cases is the
same according to the Index Kewensis.

(Plate XXXIX., Fig. 294.)

HYPERICACEAE.

1 1:3. Aecidium disseminatum Berk.

Hypericum

Berkeley in Hook. Handb. Fl. N.Z., p. 756 (1867).
Sacc. Syll. IX., p. 322 (1891).

I. Aecidia on under surface of leaves, causing corresponding eleva-

tions on upper

surface, sub-gregarious
i

or crowded, sometimes

confluent, fully \ mm. diam. ; pseudoperidia raised, with white
margin, reflexed, fimbriate; peridial cells sub-quadrate, average
30 x 22 fi.

Aecidiospores orange-yellow, sub-ellipsoid to sub-angular,
finely verrucose, 16-17 x 11-13 fi.

On Hypericum japonicum Thunb.

Victoria — Murramurrangbong Ranges, Nov. 1902, and Dec, 1903

(Robinson).

The original description by Berkeley is very brief : " Spots none or effused.

Peridia scattered, short ;

margin lobed ;

spores white." Since it was

hardly possible from this description to be certain of the species, a specimen
was sent to the Royal Gardens, Kew, for comparison with the type specimen
there, and it was found to be identical.

This aecidium occurs on the same host-plant as Melampsora hypericorum
(DC.) Schroet. and is occasionally found on the same leaf with the uredosori.

Aecidium — Ranunculaceae. 201

RANUNCULACEAE.

144. Aecidium calthae Grev

Caltha

Greville, Fl. Ed., p. 446 (1824).

Mc Alpine, Proc. Linn. Soc. N.S.W., XXIV., p. 301 (1899).

Sacc. Syll. VII., p. 602 (1888).

0. Spermogonia arranged in small groups, honey-coloured.

1. Aecidia distributed on upper surface of leaf and often confluent, abou

mm. diam. ; pseudoperidia white, margin laciniate ; peridial

cells pentagonal or hexagonal, margin finely streaked, 28-32 ^i.

Aecidiospores bright orange, angular to sub-globose or oval, finely
echinulate, average 22-23 x 17-20 ^.

On living leaves of Caltha introloba F. v. M.

New South Wales — Mt. Kosciusko, Jan., 1S99 (Maiden).

I have provisionally named this as above, although Greville describes the
aecidium as hypophyllous and on the petioles, while here it was epiphyllous,
and only on the blade of the leaf. However, the spores and pseudoperidial
cells quite agree with British specimens. The host-plant is indigenous to
Australia, and was found in a rocky creek, on eastern side of Mueller's
Peak, Mt. Kosciusko, at a height of about 6,500 feet. This was the only
micro-fungus found there by Mr. Maiden. The Puccinia-stage of this
fungus occurs in Europe and America, and has hitherto only been found on
the marsh marigold (Caltha palustris L.) ; until that stage has been dis-
covered here, it cannot be definitely stated that the two fungi are the same.

(Plate XXXIX., Fig. 295.)

•

Ranunculus

145. Aecidium ranunculacearum DC.

De Candolle, Fl. fr. VI., p. 97 (1805).
Cooke, Handb. Austr. Fung., p. 340 (189
Sacc. Syll. VII., p. 776 (1886).

I. Aecidia hypophyllous or surrounding lobes of leaf and on stalk,

densely crowded, very bright orange, often causing a swelling ;
pseudo-peridia cup-shaped to cylindrical, whitish, with margin
brittle and lacerated, 1 mm. high ; peridial cells somewhat quadrate
and tapering to a point where they dovetail into each other,
with striated margin, 27-28 x 18-22 ^u.

Aecidiospores orange-yellow, polygonal to oblong or ellipsoid,
very finely echinulate, 17-30 x 14-20 /u, average 22 x 17 yu.

On Ranunculus parvijlorus L., R. rivularis Banks and Sol., R. lappaceus
Sm. and R. gunnianus Hook.

Victoria— Mount Emu Creek, 1854. Omeo, Nov., 1882 (Stirling).

Ardmona, 1896 (Robinson). Near Dimboola, Nov., 1897
(Reader). Murramurrangbong Ranges, Nov., 1902 (Robin-
son). Buffalo Ranges, Dec, 1904 (C. French, jun.).

On Ranunculus sp.

New South Wales — Guntawang and Mudgee (Hamilton). Rich-
mond (Musson).
Tasmania— St. Patrick's River, Nov., 1844 (Gunn), (Berkeley1).

varies some-

■

This aecidium occurs on several species of Ranunculus, and varies
what in general characters, although the aecidiospores all agree in be v
very finely echinulate. As the result of cultures, this has been found to be

02 Uredo — Gramineae, Cy-peraceae.

a composite species, the aecidiospores infecting various grasses on which
they produce uredo and teleutospores. Until similar experiments are
carried out here, the different aecidia, occurring on species of Ranunculus
must be grouped together.

Darluca Jilum Cast, was found associated with the aecidia.

(Plate XXXIX., Fig. 296.)

UREDO Pers.

This stage probably belongs to teleutospore-forms which have not been
observed, and which may only rarely be produced. But it is also quite
possible that some may possess a perennial mycelium, and dispense with the
necessity of forming teleustopores . Several in Dr. Cooke's Handbook of Aus-
tralian Fungi have been referred to their proper genera. U. notabilis Ludw.,
belongs to Uromycladium notabile, and U. armillata Ludw., to Puccinia
juncophila. U. clematidis Berk., has already been shown to be a Caeoma, and
U. cichoracearum DC, as determined by Cooke, belongs to U. bidentis P.
Henn.

General characters. — Sori without pseudoperidium, covered or erumpent,
for the most part orange-yellow, rather pulverulent, and spores produced
singly on the terminal ends of hyphae, never smooth. Germination by germ-
tube, which enters the host-plant through stomata.

Uredo-forms, 13.

GRAMINEAE

Saccharum

146. Uredo kuehnii Krueg

»•

Krueger, Bericht. Zuck. Java IX., p. 117(1891).
Cobb, Agr. Gaz. N.S.W. IV., p. 799 (1893).
Sacc. Syll. XL, p. 182 (1895).

Uromyces kuehnii Krueg.

II. Sori mostly hypophyllous, linear, at length bursting through epidermis

and forming elongated brownish or blackish narrow streaks, single
or confluent, with clavate, brown paraphyses.

TJredospores globose to ellipsoid or piriform, brown or orange,
rather thin-walled, echinulate, 47-53 x 28-35 /j, ; pedieels hyaline,
clavate, somewhat shorter than spore.

Common on leaves of Saccharum ojjfiicinarum L.

New South Wales — Clarence River (Cobb7).
Queensland — Brisbane and Mackay, 1898 (Tryon5).

Dr. Cobb found the spores to possess four equatorial germ-pores, and I

have given his measurements, although Krueger makes them out to be
28-57 x 18-34 u.

O" *"" "~&

Specimen

CYPERACEAE.

147. Uredo scirpi-nodosi McAlp

Scirp

II. Sori bullate, elongated, splitting lengthwise, running in lines on stem,

1^-2 mm. long.

TJredospores brown, subglobose, ellipsoid to oblong, variable in size,
25-32 jx diam., or 27-36 X 19-25 /u; epispore echinulate, often

3 a thick.

Uredo — Liliaceae.

203

On Scirpus nodosus Rottb.

j Victoria— Mordialloc, Dec, 1885 (Reader) (Winter 2).

[ This specimen is given as Puccinia rimosa (Link) Wint., by Winter, but

I on carefully examining some of the original material, there are coarsely

I echinulate uredospores and nothing else. It is evidently not that species

described by Winter in Hedwigia, p. 28 (1880), and since it does not agree

with the uredo stage of Puccinia scirpi DC. of which I have examined
specimens from Sydow's Ured. Exs. 685 on Scirpus lacustris L., I have

ed it as abo\

(Plate XXVIII., Fig. 241.)

LILIACEAE.

148. Uredo anguillariae Cooke.

Wurmbea.

Cooke, Grev. XIV., p. 11. (1885).

Cooke, Handb. Austr. Fung., p. 343 (1892).

Sacc. Syll. VIL, p. 840 (1888).

II. Sori on both surfaces, gregarious, elliptic, bullate, long covered by the

epidermis.

Uredospores globose or globosely oval, smooth, brown, epispore
thin, 20-22 x 20 /u, on short deciduous hyaline pedicels.

On leaves of Wurmbea (Anguillaria) dioica F. v. M.

New South Wales — Guntawang (Hamilton).

It is curious to note that Cooke and Massee have also described a
Puccinia with uredo and teleutospores on this plant, the uredospores being
25-28 x 15-18 p.

Specimen

149. Uredo geitonoplesii Mc Alp

G eitonoplesium

II. Sori on under surface of leaves on vinous patches, in small groups

minute, elliptical, ultimately rupturing epidermis.

Uredospores yellowish to yellowish brown, elliptical to ovoid,
very finely echinulate, 19-27 X 17-19 p.

On G eitonoplesium cymosum A. Cunn.

Victoria — Orbost, July, 1901 (Pescott).

Schelhammera

150. Uredo schelhammerae McAlp.

II. Uredosori epiphyllous, yellowish-brown, round to ellipsoid, compact,

scattered or in groups, occasionally confluent, soon naked and sur-
rounded by ruptured epidermis, ^ mm. diam. or up to 1 mm. long.
Uredospores golden yellow, ellipsoid to obovate, very finely
echinulate, usually with two germ-pores on one face, 22-26 x
15-20 ju, average 24 x 18 yu.

On Schelhammera undulata R. Br.

New South Wales— Kurrajong Heights, 1892 and 1903 (Musson).

204 Uredo — Rhamnaceae, Stylidiaceae, Compositae

RHAMNACEAE.

Spyridium, Pomaderi

151. Uredo spyridii Cooke and Mass.

Cooke and Massee, Grev. XV., p. 99 (1887).
Cooke, Handb. Austr. Fung., p. 343 (1892)
Sacc. Syll. VII., p. 861 (1888;.

II. Sori hypophyllous, punctiform, generally scattered, sometimes con-
fluent, ochraceous to snuffy brown, soon bursting through epidermis,
pulverulent.

Uredospores yellowish to orange-yellow, ellipsoid to obovate,
distinctly echinulate, with coarsely granular contents, 22-31 x
17-22 /u; paraphyses enveloping sori, long, curved, tufted, hyaline
to pallid, generally slightly swollen at apex.

On leaves of Spyridium parvifolium F.v.M.

Victoria— Oakleigh, June, 1886 (Watts). Sept., 1887 (Mrs.

Martin). Murramurrangbong Ranges, Dec, 1903, and Jan.,
1905 (Robinson).

On Pomaderris apetala Labill.

Victoria — Murramurrangbong Ranges, Jan., 1905 (Robinson).
Tasmania — Devonport, Jan., 1906 (Robinson).

On both genera of host plants the sori are surrounded by paraphyses.
The rust is very plentiful on the leaves, and, although a large quantity of
material has been examined at different seasons of the year, only the uredo-

ge has been found

(Plate XXVIII., Fig. 245.)

STYLIDIACEAE.

152. Puccinia stylidii McAlp.

Stylidium.

II. Sori on both surfaces of leaf, solitary or sub-gregarious, elliptic,

bullate, and splitting raised epidermis, sometimes confluent later-
ally, reaching 1 mm. or more in length.

Uredospores orange-yellow, elliptic to subglobose, echinulate, with
solitary germ-pore on one face, 22-24 X 16-18 ju, or 22-24 jx diam.

III. Teleutosori, p. 210.

On leaves of Stylidium gramini folium Sm.

Tasmania — New Waterworks, Hobart, Nov. 1892, II. (Rod way).
Darluca filum, Cast., literally covered some of the sori.

(Plate XXVIII., Fig. 242.)

COMPOSITAE

Bidens.

153. Uredo bidentis P. Henn.

Hennings, Hedw. XXXV., p. 251 (1896)

Sacc. Syll. XIV., p. 395 (1899)

II. Sori scattered or densely gregarious, amphigenous, most numerous

on under surface, minute, pulvinate, then flattened or somewhat
patelliform, ochraceous, surrounded by the epidermis.

Uredospores ovoid, ellipsoid or subglobose, yellow or pale brown,
epispore pale chestnut-brown, 2-3 li thick, finely echinulate or almost
smooth, 25-35 X 22-27 fx, in exceptional cases 41 il long; para-
physes numerous, hyaline, clavate, 50-60 u long.

Urcdo — Co?npositae

205

On leaflets and leaf stalks of Bidens pilosa L

: Queensland — Brisbane, 1886 (Bailey5).

Part of the original specimen named by Dr. Cooke Uredo cichoracearum

DC, was kindly forwarded to me by Mr. Bailey, and on comparing it with

|- the above species in Exsicc. Syd. Ured., 1647, it was found to be the same.

Even the paraphyses, which are not recorded by Hennings, occurred in both.
Puccinia bidentis Diet, and Holw. was described in the Botanical
Gazette, XXIV., 32 (1897), on a species of Bidens in Mexico, and probably
the complete stage of this uredo. The original species was founded on

Bidens pilosa from Brazil

(Plate XXVIII., Figs. 243-244.)

154. Uredo crepidis-japonicae Lindr.

Crepis

Lindroth, Acta Soc. Fauna et Flora fennica, XXII., p. 1 1

(1902).
Sacc. Syll. XVII., p. 438 (1905).

II. Sori amphigenous, but often hypophyllous, cinnamon brown, round,

minute, at first surrounded by epidermis.

Uredospores sub-globose or broadly ellipsoid, yellowish, finely
echinulate, with three scattered germ-pores and epispore 1 /t thick,
14-18 n diam.

On leaves of Crepis japonica Benth.

Queensland.

Specimen not seen.

Olearia

155. Puccinia oleariae McAlp.

Mc Alpine, Agr. Gaz. N.S.W. VI., p. 756 (1895).
Sacc. Syll. XIV., p. 278 (1899).

Uromyces aster is McAlp .

II. Uredosori hypophyllous, pulvinate, scattered, bright orange.

Uredospores oval or irregularly pear-shaped to spathulate,
stalked, bright orange, apiculate at apex and somewhat thickened,
with three equatorial pores on one face, 39-51 x 19-24 fx; epispore
longitudinally striate, average thickness 1 J f ; pedicels relatively
short, hyaline, and continuous with spore.

III. Teleutosori similar but brownish.

Teleutospores intermixed with uredospores, bright orange,
oblong to oblong-clavate, smooth, fragile, constricted at septum,
rounded at apex, generally tapering slightly towards base, 62-90
x 18-28 fx; pedicel hyaline, persistent, short, up to 56 it long.

X. Mesospores occasional, similarly coloured, elongated ellipsoid,

rounded at both ends, 60-68 x" 18-22 p.

On leaves of Olearia aryophylla F. v. M. (Aster argophyllus Labill.).

Tasmania — Near Hobart, November, II. (Rodway1). Cataract

Gorge, Jan., 1906, II., III. (Robinson).

This species was originally described as Uromyces asteris, but on extended
critical examination of the type material I placed it in the genus Uredo on
account of the number of the equatorial germ-pores. Since then additional
material has been found, showing it to be a Fuccinia.

(Plate XXVIII., Fig. 249.)

2o6 Uredo — Proteaceae, Crassulaceae, Leguminosae.

PROTEACEAE.

Hakea

156. Uredo angiosperma Thuem.

Thuemen, Symb. Myc. Austr. IV., No. 95 (1880).
Cooke, Handb. Austr. Fung., p. 343 (1892).
Sacc. Syll. VIL, p. 840 (1888).

II. Sori on both surfaces, large, commonly disposed about a circle,

covered by the splitting, torn, and elevated epidermis, powdery,

brown.

Uredospores oval or ellipsoid, rounded at the apex, somewhat
narrowed at the base, 45 x 30 fi ; epispore smooth, pale brownish,
5-7 f.i thick.

On living leaves of Hakea sp.

West Australia— Oct., 1877 (F. v. Mueller) (Thumen2).

Specimen not seen.

CRASSULACEAE

Tillaea

157. Uredo tillaeae McAlp,

LEGUMINOSAE

Bossiaea

158. Uredo bossiaeae McAlp.

II. TJredosori on both surfaces of leaves but mostly on under, chocolate

brown, erumpent, surrounded by ruptured epidermis, compact,
scattered or aggregated, sub-globose, 1-1^ mm. diam.

Uredospores elliptic, finely echinulate, golden-brown, epispore
about 3 fA thick, 24-31 x 17-21 /u.

On leaves of Bossiaea prostrata R. Br.

Tasmania — Hobart, April, 1893 (Rodway).

An Aecidium has been found on species of Bossiaea in West Australia,
Victoria, New South Wales, and Tasmania, but there is no evidence to con-
nect it with this form. It is also noteworthy that a Cronartium is found on
species of Bossiaea.

(Plate XXVIII., Fig. 246.)

159. Uredo pallidula Cke. and Mass.

Cassia

Cooke and Massee, Grev. XXII, p. 37 (1893).
Sacc. Syll. XI., p. 222 (1895).

II. Sori pallid, convex, gregarious, splitting irregularly and then girt by

the ruptured epidermis, on both surfaces.

Uredospores pulverulent, tawny in the mass, pale-yellow by
transmitted light, elliptical to ovoid, finely echinulate, 16-19 X
12-13 u.

McAlpine, Agr. Gaz. N.S.W. VI., p. 757 (1895).
Sacc. Syll. XIV., p. 390 (1899).

II. Sori hypophyllous, minute, round to oval, sometimes confluent.

Uredospores variable in size and shape, usually globose or some- \

what oval, smooth, orange-yellow, about 19 li diam. or 20 X 16 p..

On leaves of Tillaea sieberiana Schult.

Victoria — Studley Park, near Melbourne (Barnard).

;•

Excluded or Doubtful Sp

On leaves, twigs, and legumes of Cassia sp

In the original description, the spores are given as smooth, and 12-14

Queensland — Gladfield (Gwyther) (Bailey

8-10 ;/, but on examining material kindly supplied by Mr. Bailey, of
Brisbane, they were found to be as above.

(Plate XXVIII., Fig. 247.)

CHENOPODIACEAE

160. Uredo rhagodiae Cke and Mass.

Rhayodia

Cooke and Massee, Grev. XV., p. 99 (1887).
Cooke, Handb. Austr. Fung., p. 343 (1892).
Sacc. Syll. VII., p. 859 (1888).

II. Sori hypophyllous, scattered, globose, ruddy-brown, a long time
| covered, at length torn and girt by the remains of the epidermis,

up to 1 mm. diam.

Uredospores yellowish, ellipsoid, echinulate, with four or more
scattered germ-pores on one face, 24-30 x 17-22 p.

On leaves of Rhagodia billardieri R. Br.

Victoria — Botanic Gardens, Melbourne, June, 1886 (Watts).

Sandringham, April, 1905 (Robinson).

Cooke and Massee gave measurements of spores as 20 X 15 /j from
Watts' specimen from Botanical Gardens, Melbourne. I have been able to
examine part of that same material returned by Cooke and Massee to the
National Herbarium and find the spores to measure 24-30 x 17-22 /u. The
rust is fairly abundant along the eastern shores of Port Phillip, and in fresh

material examined the spores have the same measurements, roughly about
27 x 20 H.

Darluca filum Cast, is often found on the sori.

(Plate XXX., Fig. 267.)

EXCLUDED OR DOUBTFUL SPECIES.

When one considers that specimens of the various Rusts have hitherto
been mostly sent to Britain or Germany for determination, it can readily be
understood that sometimes the material was in an imperfect condition or not
in the best possible state for proper examination. Hence in some cases the
species were wrongly determined, and there are several recorded in Cooke

Handbook of Australian Fungi, which on further investigation must be

removed from the list of Australian species. Sometimes, although rarely

happened that the host-plant of the

ly given, and

new species has been created which afterwards turned out to be a k
On these and other grounds the following are excluded

1. Uromyces amygdali Pass. — Cooke determined this on Peach and
Almond leaves from Victoria, New South Wales, and Queensland, but when
complete specimens were examined it was found to be merely the uredo-
stage of Puccinia pruni Pers.

2. Uromyces junci (Desm.) Wint. — This species has not been found in
Australia, as it was wrongly determined at Kew Herbarium for Pziccinia
juncophila Cooke and Mass.

3. Puccinia acetosae (Schum.) Koern.— This rust was determined by
Cooke on a native species of Rumex, but on the examination of a specimen

the National Herbarium, Melbourne, named in his own handwritin

2o8 Excluded or Doubtful Species.

was found to agree with P. ludwigii Tepper. P. acetosae and its synonym,
P. rumicis Lasch. are therefore excluded.

4. Puccinia aegra Grove. — P. depauperaiis (Vize) Syd. — The rust on
the native violets ( Viola hederacea and V. bet onicif olio) is comparatively
common, and has been collected at various localities in Victoria, New South
Wales, and Tasmania. It was at first determined by Winter as this species,
and Cooke has also recorded it, but the examination of numerous specimens,
including part of the same material forwarded to Dr. Cooke, showed the
teleutospores to be warted, not smooth, and aecidiospores likewise finely
warted, and therefore not this species.

5. Puccinia apii Desm. — The rust found on Celery, both native and
imported, in Australia, is found to be quite distinct from this species.

6. Puccinia caulincola Corda is given by Cooke on Hypochoeris glabra,
but the species on this host-plant is undoubtedly P. hypochoeridis Oud.

7. Puccinia geranii Corda was the name given by Cooke to the rust
occurring on Pelargonium australe, but it is now found to be a different
species, P. morrisoni McAlp.

s. Puccinia investita Schw. was determined bv Dr. Morrison on

species of Gnaphalium, but on investigating the original material I found it
to be Pn gnaphalii (Speg.) P. Henn.

9. Puccinia microseris McAlp. — Since the host-plant of this species
was really Hypochoeris radicata it is now excluded.

10. Puccinia phragmitis (Schum.) Koern. — This species has not yet
been found in Australia, only P. magnusiana Koern and 1 \ tepperi Ludw.

11- Puccinia rimosa (Link.) Wint. — This species is given by Winter2
as occurring on Isolepis nodosa R. T$r.=Scirpus 7iodosus Rottb. in Victoria,
but an examination of the original material only shows uredospores which
do not agree with those of this species, and Sydow * has arrived at the same
conclusion. The local specimen I have named Uredo scirpi-nodosi.

12. Puccinia violae (Schum.) DC. — This species is also given by Cooke
as occurring on violets in Victoria, but the history of the specimens shows
how necessary it is to guard against recording species as Australian which
may have been imported in the dry condition. The original material, with
the name in Cooke's handwriting, is accompanied by the following note
from the collector, Mr. Reader: — "On leaves of what appears to be Viola
sp. found in an old book." I was so convinced, however, that P. violae did ;
not occur on our native violets, and had not been found growing on culti-
vated species, that I communicated with the collector, who informed me
that the leaves on which this species was found had been brought from
Europe and forwarded with other specimens for determination to Dr.
Cooke. Unfortunately, on the strength of this determination, Mr. Tryon, in
Queensland, and others, have also credited Australia with this species. Mr.
Reader, who is a chemist, also sent myself some jaborandi leaves
(Pilocarpus pennatifolius, Lem.) with a rust upon them, and it was found
to be Puccinia pilocarpi Cooke, but of course it had been imported.

13. Cronartium asclepiadeum (Willd.) Fries. — This widely distributed
species is recorded on a leguminous plant from Queensland, and the name is
given in Berkeley's handwriting on the specimen in the National Herbarium.
Dietel, however, regards it as a new species, and has named it C. jacksoniae.

14. Melampsora nesodaphnes B. and Br. — This species was found on
the fruit of Beilschmiedia {Nesodaphne) obtusifolia Benth., sent by Bailey to
Berkeley from Queensland, and I also found it on the fruit of Cinnamomum
oliveri Bail., sent by Baker from New South Wales. There is a mass of
powdery spores on the surface of the fruit which is swollen and distorted.

Excluded or Doubtful Species. 209

These spores are exceedingly numerous, and very variable in size and shape.
They are hyaline individually, although ochraceous in the mass and finely
granular. They vary from elliptic to ovoid or pear-shaped, and are
occasionally slightly curved. There is a clear central axis to the spore, and
it is generally broken in the centre, or may be several times transversely
divided. Their average size is 24-32 x 9-10 ^. They are usually borne on
short hyaline filaments, and occasionally a long filament is adherent to a
detached spore.

There is nothing in the appearance or nature of this fungus to indi-
cate that it is a Uredine ; it rather suggests some form of Hyphomycete.

15. Aecidium apocyni Schwein. — The specimen referred by Cooke to
this species is Caeoma apocyni.

16. Aecidium barbareae DC— The Cruciferous host-plant is not a

Barbarea, and a portion of

specimen shows

aecidia. I have named the rust Puecinia cruciferae.

17. Aecidium bellidis Thuem., was determined by Cooke on Bellis
perennis from Victoria. This is considered to represent the aecidial stage
of Puecinia obscura Schroet. which occurs on Luzula, but since teleuto-
spores have been found from year to year on Victorian specimens along
with the aecidia, it is evidently quite a distinct species. I have, therefore,
named it Puecinia distinct a.

Mr. Cheeseman, one of our leading nurserymen, informs me that the
seeds of Bellis perennis are imported from England and Germany, mostly
from the latter country. He also states that some have had to give up
growing the plants on account of the rust (Puecinia distincta), since it
attacks the youngest seedlings sometimes, and at other times the plants
remain healthy till late in spring and then become badly attacked. One
would naturally conclude that the rust had been imported with the seed, but
it is strange that no teleutospores have been found accompanying the aecidia
in the countries named. Although I have provisionally recorded it as a
native rust, the probabilities are that it will be found to exist in the
countries from which the seed has been derived.

18. Aecidium COmpOSitarum Mart. — All the species occurring on
Compositae have been distributed among their respective host-plants.

19. Aecidium plantaginis Ces. — The species on the native Plantago
varia is distinct from this, the spores being echinulate instead of
tuberculate, and considerably larger. It has been named A. plantaginis variae.

20. Aecidium seuecionis Desm. — This has also been determined in the
absence of complete specimens, but since the teleutospores have been found
associated with the aecidia, it has been determined as a new species by
Dietel, and named P. tasmanica. It is a striking fact that on three such
common imported plants as the garden daisy, Bellis perennis, groundsel,
Senecio vulgaris, and marigold, Calendula officinalis, there should hav^e been
new species of Puecinia discovered. As regards the groundsel, aecidia have
been found on native species of Senecio, probably belonging to the same rust.

21. Aecidium Violae Schum., is also recorded, but this belongs to P.
violae, and is therefore excluded.

22. Uredo Cichoracearum DC. originally determined by Dr. Cooke
on Bidens pilosa is now found to be U. bidentis P. Henn.

23. Uredo leguminum Desm. — This species originally occurred on
Phaseolus, and is recorded by Cooke as being found on pods of Acacia in
Queensland, but since there is every reason to believe that the rust on a bean
is distinct from that on a wattle it is excluded from the list. Probably it is
a stage of a Uromycladium.

2IO

Addenda.

ADDENDA

AMARANTACEAE.

161. Uromyces polycnemi McAlp.

Polycnemum.

I. Aecidia bright orange, generally covering lower surface of leaves,

crowded, cylindrical, surrounded at base by ruptured epidermis, up
to \\ mm. long; pseudoperidia colourless, with toothed margin;
peridial cells oblong to somewhat lozeuge-shaped, with moderately
thick striated margins, 37-40 /x long.

Aecidiospores orange, ellipsoid to oblong or sub-globose, very
finely echinulate, 22-23 X 17-20 ^ or 20-22 /x diam.

II. Uredo-sori on both surfaces of leaf, bullate, at first covered by

leaden-coloured epidermis, then splitting, scattered or gregarious,
sometimes confluent, up to 2 mm. in diam.

Uredospores ellipsoid to oval, orange coloured, finely echinulate,
with three equatorial germ-pores on one face, 28-32 X 18-22 \x.

III. Teleuto-sori similar to uredo-sori, but generally larger and darker in

colour.

Teleutospores dark golden-brown in mass, tawny brown indivi-

dually, ellipsoid to

elongated ellipsoid, rounded

at both ends,

occasionally slightly thickened at apex and germ-pore prominent,
smooth, 34-46 X 17-25 /x; pedicel hyaline, persistent, compara-
tively short.

On Polycnemum pentandrum F.v.M.

Victoria — Port Fairy. Dec, 1905.

The aecidia were very conspicuous, often covering the under sin

every leaf, and as the host pla
near the coast, it was difficult

surface of
dense masses, in the saline marshes

A Tuberculina was frequently parasitic on the aecid

(Plate XLIIL, Fig. 319.)

STYLIDIACEAE.

152. Puccinia stylidii McAlp

II. Uredosori, p. 204.

Stylidium

III. Teleutosori on both surfaces of leaf, dark-brown to black, round to

elliptic, sometimes confluent in long lines, pulvinate, splitting and
surrounded by epidermis, up to 1^ mm. or longer. %

Teleutospores at first intermixed with uredospores, bright chest-
nut-brown, clavate to oblong, smooth, slightly constricted at sep-
tum, rounded or bluntly pointed and thickened at apex (9-11 ^),
rounded or attenuated at base, not infrequently three-celled, very
variable in shape and size, 40-62 x 17-28 /jl; pedicel hyaline,
persistent, up to 45 fx long.

X. Mesospores common, similarly coloured to teleutospores, thickened

at apex, oval to ovoid or elongated ellipsoid, 30-40 X 15-23 /x.

On leaves of Stylidium gramini folium Sm.

Tasmania — Devonport, Jan., 1906 (Robinson).

Only the uredospores were at first sent by Rodway in Nov., 1892, and

the discovery of the final stage shows that Uredo stylidii is a Puccinia, the
description of which is now completed.

i
i

Glossary

211

GLOSSARY

Abstriction. — The formation of a spore by pinching off the end of a spore-forming

hypha, without the production of a septum.

Aecidiosfore. — A spore formed in an aecidium, serially and successively abstricted.

Aecidium. — A spore-body consisting of a cup-shaped envelope or peridiuni with a

hymenium at the bottom of the cup, from which aecidiospores are successively
detached; also the generic name originally applied to the aecidiospores.

'Amfkigenous. — Growing on both surfaces of a leaf.

Amfhisfore. — A modified uredospore with thickened wall, and only germinating

after a resting period.

Affressorium. — The vesicle formed by the tip of a germ-tube over the stoma of a

leaf before entering it.

Autoecious. — Applied to a parasitic fungus of which all the spore-forms are capable of

being produced on the same host.

Basidium. — The mother-cell or hypha from which spores are given off.

Basi fetal. — Growing in the direction of the base from above downwards.

Bullate. — Raised like a blister.

Caeoma. — A spore-layer in which the spores are formed in chains but without an

envelope of peridial cells; an aecidium without a peridium.

Caeomosfore. — A spore formed in a caeoma.

Ca fit ate. — Ending in a head, as when the free end of an upright hypha enlarges in

a spherical manner.

Chemotaxis. — A form of sensitiveness which certain organisms possess towards cer-
tain chemical substances.

Circinate. — Arranged in a circular manner or nearly so.
Clavate. — Club-shaped, enlarging towards the apex.

Deciduous. — Falling away, applied to the pedicel of a spore when it does not re-
main attached to it.

E chinul at e . — Covered with short spines.
Elliftical. — Shaped like an ellipse or oval.

Endochrome. — The peculiar colouring matter in the cells, and usually applied to the

coloured cell-contents of spores.

Endosfore. — The innermost coat of a spore.

Efifhyllous. — Growing on the upper surface of a leaf.

Efiteosfore. — A name suggested for a uredospore which functions as an aecidiospore,

and the sorus contains or is surrounded by prominent paraphyses.

Erumfent. — Bursting through the surface of the host-plant.

Fimbriate. — Fringed, when the margin is bordered by slender processes.

Form-genus. — A genus in which the species are only represented by single stages of

what is believed to be an incomplete life-history, as Uredo, Aecidium.

Gall. — A morbid enlargement or hypertrophy of the cells of a plant, due to para-
sitic agency.
Gametofhyte. — The stage in the life-cycle of the plant which bears the sexual organs.

Germ-fore.— An opening on the surface of a spore through which the germ-tube

passes.
Germ-tube.— A tubular process developed from the spore in germination, and may

either become a mycelium or promycelium.

Gibbous. — Swollen at the base.

Globose. — Spherical.

Haustorium.—A short lateral branch of a hypha, penetrating a cell of the host-plant

and acting as a sucker as well as an organ of attachment.
Htteroccious.— Applied to a parasitic fungus of which certain spore-forms occur on

one host and others on plants widely remote.
Host. — A plant which nourishes a parasite.
Hyaline. — Colourless or translucent.

212

Glossary

Hymenium. — A spore-bearing layer of hyphae.

Hypertrophy. — An abnormal enlargement or excessive growth of particular parts,.

which may be caused by excess of nourishment or induced by parasites.

Hypha. — A tubular thread-like cellular structure, collectively constituting the vege-
tative body or mycelium of the fungus.

—Growing on the under surface of a leaf.

Hypophyllous.—

Indigenous. — Native to a country, and not introduced.

Intercellular. — Between the cells of the host-plant.

Intracellular. — Within the cells of the host-plant.

Laciniate. — Cut up into narrow lobes.

Lumen. — The cavity bounded by the walls of an organ.

Mesospore. — A single-celled spcre-form related to the teleutospore, and probably

representing a transition from the unicellular Uromyces.

Micron or "Micro millimetre. — Indicated by the

sign

jiy is the standard unit for

microscopical measurements, and equals i-ioooth of a millimetre, or i-25,oooth
of an inch.

Mutation. — That kind of

equilibrium of the organism is disturbed and

variation produced by internal influences, in which the

a new position of equilibrium

found strikingly different from the original one, sometim-es called spontaneous
variation. ,

Mycelium. — The vegetative portion of a fungus composed of one or more hyphae.

Nucleus. — An organized proteid constituent of the cell, necessary for its continued

growth and life.

Obligate parasite. — One in wThich a parasitic mode of life is indispensable for com-

plete development.

Orbicular. — Applied to a flat body with a circular outline. i

Papillate. — Covered with protuberances or terminating in a papilla.

Paraphyses. — Barren filajnents associated with the spoies, either slender or variously

swollen, hyaline or coloured.

Parasite. — An organism living on or in another living organism, and at its expense.

Pedicel. — Spore-bearing stalk.

Persistent. — Lasting, not soon falling away, applied to the pedicel of a spore.
Piriform. — Pear-shaped.

Plasmodium. — A mass of naked protoplasm, containing many nuclei and exhibiting

movement.

Plurivorous. — Applied to a parasite which can exist on a variety of host-plants.

Predisposition. — The tendency to disease exhibited by an organism when the con-
ditions are favorable to the development of the parasite.

Primordia. — The beginnings of any structure, such as the aeciiium.

Promycelial spore. — The spore produced by a promycelium, also known as a sporid-

iolum.

Promycelium. — The short germ-tube of a teleutospore, which produces a few spores

unlike the teleutospores, and then perishes.

Protomycelium. — A term used by Eriksson to denote the early plasmodia-like stage-*

of the mycelium in certain Rusts.

Protopl

■The living substance in plants, consisting of albuminoid matter.

False tissue or compact mycelium, so called because it is
formed by the union of previously separate elements or hyphae into a dense

and

mass, while true tissue is

forms a coherent, more or less firm, mass of cells.

derived from a single cell, which divides

■t

— The outer tnvelopc of the aecidium, consisting of sterile cells.

Pulverulent. — Powdery
Pulvinate. — Cushion-shaped.

Covered with lines or ridges crossing each other so as to form a net-

Reticulate.—

work.
Saccate. — Bag-shaped?

Saprophyte. — A fungus which preys upon dead organic matter only, in contrast to

a parasite.

Septate. — Partitioned off into distinct divisions.

Sorus. — A cluster of spores constituting a spore-bed, as uredosorus, teieutosorus.

Literature

213

Sfermatium. — A minute detachable cell produced in a spermogonium, and formerly

considered to have a male sexual function.

Sfermogonium. — The receptacle in which spermatia are produced and detached. It

is sometimes called a pyenidium, although that term is generally reserved for
a spore-receptacle in another division of Fungi.

Spore. — A detachable cell, capable of reproducing the fungus.

Sporidiolum. — Spore produced by the germ-tube of a teleutospore. Also called

sporidium, but since this term is already used as equivalent to an ascospore,
Saccardo has proposed the present name.

Sporophyte. — The stage in the life-cycle of a plant which bears the spores.

Sterigma. — A delicate stalk from which a spore is detached and arising from a

basidium.

Stoma. — A breathing pore or aperture in the epidermis forming a means of communi-
cation between the inside of the plant and the outer air.

Stroma. — A cushion or receptacle bearing reproductive bodies.
Symbiosis. — The living together of dissimilar organisms, both contributing to their

mutual welfare. It differs from parasitism' in which the benefit is all on the

side of the parasitic organism.
Teleutos<pore.—Genera\\y regarded as the final spore in the life cycle of the Rusts,

which germinates and produces a promycelium and promycelial spores.

Truncate. — Cut off abruptly.

Uredo. — The generic name originally applied to the uredospores.
Vredosfore.—A spore detached from the apex of a pedicel and producing a

mycelium.
Verrucose. — Covered with small warts.
Witches' broom.'— A diseased state of the shoots of a plant, when they are developed

in tangled masses or tufts, due to parasitic agencies and especially fungi.

LITERATURE.

1. Arthur, J. C— The Aecidium as a Device to Restore Vigour to the Fungus.

Proc. Soc. Prom. Agric. Sci., Vol. 23, p. 65. 1902.

Problems in the Study of Plant Rusts. Bull. Torr. Club, Vol. 30. P-

903

904

Taxonomic Importance of the Spermogonium. Ibid. Vol. 31, p. H3-

-. The Aecidium of Maize Rust. Bot. Gaz., Vol. 38, p. 64. 1904.

J. Amphisporesi of the Grass and Sedge Rusts. Bull. Terr. Club, \ol. 32,

6. P" 3A N9ew' Uredineous Fungus from Mexico— Baeodromous holwayi Arth.

Ann. Myc, Vol. 3, p. 18. 1905.

- - • J

i9°5

7. Cultures ot ureameac m iyu4. j«-u. MJ~. .—. --, r ^ ' ^ v ,

8. Terminology of the Spore-Structures in the Uredinales. Bot. Gaz., vol.

39, P- 2I9- i9°5

1. Arthur, J. C, and Holway, E. W. D.-Descriptions of American Ured.neae.

Bull. Lab. Nat. Hist., Univ. Iowa, Vol. 5, P- 3*9- If><>2-
1. Atkinson, J.-An account of Agriculture and Grazing in New South ttales.

London. 1826.

Brisbane. 1883.

1. Bailey, F. M.— Synopsis of the Queensland Mora pp. 774"Z7»- ""g

2. Contributions to the Queensland Flora. Proc. Roy. Soc, Q

Vol. 1, Part I., p. 18. 1884.

* =1 l^psU of *e SXs.al.dVra, Firs, Supplement, pp. 84, 85. Bri,

bane. 1886. -n • u .~ Tcoo

5. Ibid, Second Supplement, pp. 125-127-. . Bnsban|- lb88'

6- Ibid, Third Supplement, p. 123. Brisbane. 1890.

7- Additional Fungus Blights observed to have ^e.l^ 5w *

' - • -c. of Queensland, p. 45- Brisbane. 1890-01.

leensland Flora. Botany Bull. 2, p. 34. Brisbane.

. of Queensland, p. 51. Brisbane. 1891-2.

year

189I.

214

Literature.

10. Bailey, F. M. — A Review of the Fungus Blights which have been observed to in-
jure living vegetation in Queensland. Rep. Aust. Assoc. Adv. Sci., Hobart,
Vol. 4, pp. 392, 395, 396, 397, 398, and 400.

11.
12.

*3-

14.
15-

16.
17-

18.

19.

20.

21.

395.

Contributions to the Queensla]
Ann. Rep. Dep. Agric. of Q
Contributions to the Queensh

1892.

Botany Bull. 5, p

p. 40. Brisbane.

36.. 1892.
1892-3.

Flora. Botany Bull. 8, pp. 110-113

Brisbane. 1893.

Ann. Rep. Dep.
Contributions to

Brisbane. 1894.

Agric. of Queensland, p
the Queensland Flora.

Brisbane. 1893-4.

Ann Rep. Dep.
Queensland

Agric.

Blight

of Queensland, p.

Fungi. Proc. Conf.

31-

Botany Bull. 9, pp. 16, 17

35. Brisbane. 1894-5.
Austr. Fruit-growers, New

Zealand, p. 208. Well

1896.

Ann. Rep. Dep.

Agric.

of Q

p. 39. Brisbane. 1897-8.

1898.

Botany Contributions. Queensland Agric. J

38. Brisbane.

ban

1898.
Ann. Rep. Dep. Agric/ of Q

Ibid. Vol. 3, p. 205

Bris-

Brisbane. 1899.

p. 33. Brisbane, 1898-9.
. Agric. Journ. Vol. 4,

284.

1. Baker, R. T.

Contributions to a Knowledge of the Flora of Australia.

Proc.

Linn. Soc, New South Wales, Vol. 24, p. 447. 1899.

1. Bandi, W. — Beitrage zur

Biologie

der Uredineen (Phragmidium subcorticium

{Schrank) Winter). Hedw., Vol. 42, p. 118. 1903

1. Barclay, A. — Aecidiwn

w.,

urticae

Schum. var. himalayense. Sci. M

Off. Army of India, p. 29. Calcutta. 1887.

flort

Vol.

On the Life History of a Remarkable Uredine on J

grandi

(Uromyces cunninghamianus, n. sp.). Trans. Linn. Soc, London,

3, p. 141. 1891.
On Two Autoecious

Caeomata in Simla. Scientific M

by M

Officers of Army of India, Part VI. 1891.

Rust and Mildew in India. J

1. Bary, A. de. — Neue

Bot, Vol. 30, p. 349

1892.

Untersuchungen

ueber die Uredineen, insbesondere die
Entwickelung der Puccinia graminis und den Zusammenhang derselben mit
Aecidium berberidis. Monatsber. K. Acad. d. Wiss, p. 25.

1865.
Comparative Morphology and Biology of the Fungi, Mycetozoa

teria. English Edition, Oxford. 1887.

is d'immunisation des vegetaux contre les maladies crypto-

e, M. J.-

gamiques. Compt. rend. Vol. 133, p. 107

1 901

1. Berkeley, M. J. — Flora of Tasmania

Fungi.

Hooker's Botany of the Ant-

arctic Voyage, Part III., Vol. 2, p. 270. i860.

& 11 ^^^ a 4 — * 4 ^^^B

Australian

1872.

Fungi.

Linn. Soc, London, Vol. 13, pp. 173* ^A*

1. Berkeley, M. J., and Broome, C. E. — List of

Fungi

land. Trans. Linn. Soc, London, Vol. 1, p. 407

Ibid, Part II., Vol. 2, p. 67. 1882.

from Brisbane, Q
1878.

Ibid, Part III., Vol. 2, p. 221. 1886.

3- .

1. Blackman, V. H. — On the conditions of Teleutospore germination and Spori-

dia formation in the Uredineae. New Phytologist. Vol. 2, p. 10. 1903.

On the Fertilisation, Alternation of Generations, and general Cytology

2. -

of the Uredineae. Ibid. Vol. 3, p. 23. 1904.

On the Fertilisation, Alternation of Generations, and

— -- j

of the Uredineae. Ann. Bot., Vol. 18, p. 323. 1904

general Cytology

1. Bolley, H. L. — The Heteroecismal Puccinieae. Am. Micro. J

10, p

169. 1889.

2
3

Bull. 26. Agr. Exp. Station, Indiana. 1889.

Wheat Rust : Is the infection local or general in origin?

Vol. 5. 1891.

Einige Bemerkungen ueber die symbiotische Mykoplasm

Getrefderost. Centralb. f. Bakt., Vol. 4, p. 855. 1898.
New work upon wheat rust. Science N.S. Vol.

Agric.

Science,
bei dem

I. BUBAK, F.

22, p. 50

I905

fhyt

fi DC. Oesterr. Bot. Zeitschr., Vol. 48, p. 14. 1898.

Ber. d. Deutsch Bot. Gesellsch., Vol. 21, p. 356.
Butler, E. J.— The Indian Wheat Rust Problem.
India. Calcutta. 1903.

1903.
Bull.

No. 1, Dept. Agric,

Literature.
*

i. Campbell, F. M. — Victorian Fungi. Vict. Nat. Vol. 4, p. 96. 1887.

1. Carleton, M. A. — Cereal Rusts of the United States. U.S. Department of

Agriculture, Division of Vegetable Physiology and Pathology, Bull. No. 16.
Washington. 1899.

2. Culture Methods with Uredineae. Journ. Applied Microscopy and Lab.

Methods. Vol. 6, p. 2109. Rochester, N.Y. 1903.

3. Investigations of Rusts. U.S. Department of Agriculture, Bureau of

Plant Industry. Bull. 63. Washington. 1904.

1. Chodat, R. — Identite du Puccinia scirfi DC. avec Aecidium nymfhoidis DC.

Verh. Schweiz. Nat. Ges., p. 43. 1888-9.

1 Christman, A. H. — Sexual Reproduction in the Rusts. Bot. Gaz., Vol. 39, p.

267. 1905.

1. Cobb5 N. A. — Peach Rust in our Orchards. Agric. Gaz., N.S. Wales. Vol. i%

p. 93. 1890.

2. Contributions to an economic knowledge of Australian Rusts (Uredineae).

Ibid., pp. 185-214. 1890.

3. Maize Rust. Ibid. Vol. 2, p. 215. 1891.

4. Notes on Diseases of Plants. Ibid, pp. 157, 286. 1891.

5. Report on Rust in Wheat. Proc. Conf. N.S. Wales. 1891.

6. Contributions to an economic knowledge of Australian Rusts (Uredineae).

Agric. Gaz., N.S.W., Vol. 3, pp. 44-68. 18Q2.

7. Fungus Diseases of the Sugar-cane. Ibid. Vol. 4, p. 799. 1893.

% Host and Habitat Index of the Australian Fungi. Dep. Agric, N.S.

Wales. 1893.
9. Notes on Diseases of Plants. Agric. Gaz., N.S. Wales Vol. 5, p. 384.

1894.

10. Letters on the Diseases of Plants. Ibid. Vol. 8, pp. 216, 217, 232-234.

l897- . . .

11. Comparative Observations on the Brush of about fifty Varieties of

Wheat. Ibid. Vol. 13, p. 647. 1902.

12. Letters on the Diseases of Plants. Ibid. Vol. 15, pp. 8-14. 1904

1. Cobb, N. A., and Oliff, A. S. — Insect Larvae {Cecidomyia sp.) eating Rust

on Wheat and Flax. Agric. Gaz., N.S.W. Vol. 2, p. 67. 1891.

1. Connold, E. T. — British Vegetable Galls. London. 1901.

1. Cooke, M. C— Australian Fungi. Grevillea, Vol. 11, pp. 97, 98. 1883.

2. Ibid, Vol. 12, p. 20. 1883.

3. Fungi australiani (reprinted from Grevillea, Vols. 9, 10, 11, 12). 1883.

4. Australian Fungi. Grevillea, Vol. 14, pp. 11, 12. 1885.

5. Ibid. Vol. 15, p. 99. 1887.

6. Ibid. Vol. 16, p. 2. 1887.

7. Ibid. ' Vol. 16, p. 74. 1888.

8. Ibid. Vol. 17, pp. 55, 70. 1889.

9. Ibid. Vol. 19, p. 46. 1890.

xo. Handbook of Australian Fungi, pp. 330-344. London. 1892.

1. Corda, A. — Icones Fungorum. 6 Fasc. 1838-56.

1. Dangeard, P. A., and Sappin Trouffy, P. — Recherches histologiques sur les

Uredinees. Compt. rend. Vol. 116, p 211. 1893.

2. Une pseudofecondation chez les Uredinees. Ibid. Vol. 116, p. 267.

1893.

1. Dietel, P. — Beitrage zur Morphologie und Biologie der Uredineen. Bot.

Centralb. Vol. 32, p. 54, &c. 1887.

2. Verzeichniss der in der Umgebung von Leipzig beobachten Uredineen.

Ber. Natf. Ges. Leipzig. 1888-89.

, 3. Ueber das Vorkommen von zweierlei Teleutosporen bei der Gattung

Gymnosforangium. Hedwigia, Vol. 28, p. 99. 1889.

4. Ueber den Generationswechsel von Puccinia agrofyri Ell. and Ev.

Oesterr. Bot. Zeitschr. Vol. 42, p. 261, 1892.

5. Die Gattung Ravenelia. Hedw., Vol. 33, pp. 22 and 367. 1894.

6. Ueber Rostpilze mit wiederholter Aecidienbildung. Flora. Vol. 8i, p.

394. 1895.

7. Waren die Rostpilze in friiheren Zeiten plurivor? Bot. Centralb. Vol.

79, p. 81 and 113. 1899.

438.

:2l6

Literature.

8. Dietel, P. — Uredinaies in Engler and Prantl, Die natiirlichen Pflanzen-

familien. 1900

TO.

II.

12.

Ueber die biologische Bedeutung der Paraphysen in den Uredolagern

von Rostpilzen. Hedw., Vol. 41, p. (58). 1902.

Ueber die auf Leguminosen lebenden Rostpilze und die Verwandt-

schafts verhaltnisse der Gattungen der Pucciniaceen. Ann, Myc. Vol. 1
p. 3. 1903.

Eine neue Puccinia auf Senecio. Ibid., p. 535. 1903.

Betrachlungen ueber die Vertheilung der Uredineen auf ihrer Nahr-

pflanzen. Centraib. f. Bakt., Vol. 12, p. 218. 1904

and

Ueber die Arten der Gattung Phragmidium. Hedwigia, Vol. 44, p. 122

P- 33o

1905

1. Dumee, P., and Maire, R. — Remarques sur les Uredospores de Puccinia fruni

Pers. Bull. Soc. Myc, France. Vol. 17, p. 308. 1901

•

1. Ehrlich, and Huebener.

Zeitschr. f. Hygiene.

' - •

Ueber die Vererbung der Immunitat bei Tetanus.

Vol. 18. 1894.

J. — Ueber die Spezialisierung des Parasitismus bei den Getreiderost-

J5-
16.

*7-

18.

pilzen. Ber. Deutsch. Bot. Ges. Vol. 12, p. 292

1894.

Ueber die Forderung der Pilzsporenkeimung durch Kalte. Centraib. f.

Bakt. Vol. 1, p. 557. 1895.

Ueber die verschiedene Rostempfanglichkeit verschiedener Getreidesorten.
Zeitschr. f. Pflanzenk. Vol. 5, pp. 80 and 156. 1895.
1st die verschiedene Widerstands fahigkeit der Weizensorten konstant

oder nicht? Ibid., p. 198.

Welche Rostarten zerstoren

1896.

1895.

Ibid. Vol.

6, p. 141.

Welche Grasarten konnen die Berberitze mit Rost anstecken? Ibid., p.

193

1896.

Neue Untersuchungen ueber die Spezialisierung, Verbreitung und

kunft des Schwarzrostes (Puccinia graminis Pers.). Jahrb. f. Wiss. Bot.

Her-
Vol.

p. 499
Der

1896.

heutig

Vol. 15, p. 183. 1897.

Stand der Getreiderostfrage. Ber. Deutsch. Bot. Ges.

Vol.

124. P- 475 . ~ ..

Ueber die Dauer der Keimkraft in der Wintersporen gewisser Rostpilze.

Centraib. f. Bakt. Vol. 4, p. 376. 1898.

Vie latente et plasmatique de certaines Uredinees. Compt. rend.

1897.

Sur l'Origine et la Propagation de la Rouille des Cereales par la Semence.

Ann. Sci. Nat. Bot., Ser. 8. Vol. 14, p. 107. 1901.

Ueber die Spezialisierung des Getreidescliwarzrostes in Schweden und

in anderen Landern. Centraib. f. Bact. Vol. 9, p. 590. 1902.

The Researches of Professor H. Marshall Ward on the Brown Rust of

for Botanik udg. of K.

the Bromes and the Mycoplasm Hypothesis. Arkiv
Sv. Vet. Akad. Vol. 1, p. 139. 1903.

Ueber das vegetative Leben der Getreiderostpilze.

Vol. 77, No. 6, I. 1004.

Ibid. Vol. 38, No, 3, II. -III.

K. Sv. Vet. Akad.

1004

1905

On the Vegetative Life of some Uredineae. Ann. Bot. Vol. 19, p. 55

Zur Frage der Entstehung und

Pflanzen. Arkiv. f. Botanik. K. Sv. Vet. Akad. Vol. 5

Ueber das vegetative Leben der Getreiderostpilze.

Vol. 39, No. 5, IV. 1905
Eriksson, J., and Henxing, E.

ueber die Getreideroste.

Verbreitune der Rostkrankheiten der

No. 3. 1905.
K. Sv. Vet. Akad.

1894.

Die

Die Hauptresultate einer neuen Untersuchung
Zeitschr. f. Pflanzenk/ Vol. 4, pp. '66 and 140.

Getreideroste, ihre Geschichte und Natur, sowie Massregeln gegen

dieselben. Stockholm. 1896.

1. Farlow, W. G. — Notes on some Species in the Third and Eleventh Centuries of

Ellis' North American Fungi.
18. Boston. 1883.

Proc. Am. Acad. Arts

d Sci.

Vol.

The Conception of Species as affected bv recent Investigations on Fungi

American Naturalist.

Vol. 32, p. 675.

189S.

Literature.

217

1. Farrer, W.— The making and improvement of Wheats for Australian condi-

tions, together with Eriksson's General review of the principal results of s»
Swedish research into the rust of cereals. Agric. Gaz., N.S. Wales, Vol. q*
p. 239. 1898.

- Some notes on the Wheat " Bobs"; its Peculiarities, Economic Value, and

Origin. Agric. Gaz., N.S. Wales, Vol. 15, p. 739 and p. 849. 1904

1. Fischer, E. — Ueber Gymnosforangium sabinae (Dicks.) Wint. and G. confusum

Plowright Zeitschr. f. Pflanzenk. Vol. 1, p. 260. x8qi.

2. Experiences d'infection avec les Puccinia helvetica Schroet. et P. mag-

nusiana

Koern. Arch. Sci. Phys. and Nat. Vol. 28, p. 376. 1892.

3. Die Zugehoerigkeit von Aecidium fenicillatum. Hedw. Vol. 34 p 1.

. l895'
4- Die biologischen Arten der parasitischen Pilze und die Entstehung neuer

Formen im Pflanzenreich. Verh. Schweiz. Nat. Ges. Vol. 86. Locarno, p. 49,
1904.

5. Die Uredineen der Schweiz. Beitr. zur Krypt. Flor der Schweiz. Vol. 2y

part 2, 1904.

1. Freeman, E. M. — Experiments on the Brown Rust of Bromes {Puccinia dis-

fersa). Ann. Bot., Vol. 16, p. 487. 1902.

1. Galloway, B. T. — Rust of Flax. Journ. Myc, Vol. 5, p. 215. 1889.

1. Gibson, C. M. — Notes on Infection Experiments with various Uredineae. New

Phytologist. Vol. 3, p. 184. 1904.

■

1. Halsted, B. D. — Mycological Notes — Observation in Wind infection of a Rust-
Bull. Torr. Bot. Club. Vol. 25, p. 159. 1898.

1. Henning, E. — Ueber verschiedenartige Predisposition des G°.treides fur Rost.

Land. Akad. Handl. Tids. 1894.

1. Hennings, P. — Fungi Warburgiani. Hedw., Vol. 32, p. 216. 1893.

2. Anpassungsverhaltnisse bei Uredineen bezuglich der physikalischei>

Beschaffenheit des Substrats. Hedwigia, Vol. 40, p. 125. 1901.

3. Fungi Australiae occidentalis I. Ibid., pp. 95 and 352. 1901.

4. Fungi australienses. Ibid., Vol. 42 (p. 73). 1903.

5. Einige Beobachtungen ueber das Gesunden pilzkranker Pflanzen bei veran-

derten Kulturverhaltnissen. Zeitschr. f. Pflanzenk. Vol. 13, p. 41. 1903.

tchcock, A. S., and Carleton, M. A. — Preliminary Report on Rusts of Grain,
Kans. Agr. Expt. Sta. Bull. No. 38. 1893.

2. Rusts of Grain, II. Ibid. Bull. No. 46. 1894.

1. Hooker, J. D., and Jackson, B. D. — Index Kewensis. Oxford. 1895, &c-

1. Jacky, E. — Die Compositen-bewohnenden Puccinien vom Typus der Puccinia

hieracii und deren Spezialisierung. Zeitschr. f. Pflanzenk. Vol. 9, pp. 193,
263, and 330. 18Q9.

2. Der Chrysanthemum-Rost. Ibid.. Vol. 10, p. 132. 1900.

3. Der Chrysanthemum-Rost. Centralbl. f. Bakt., Vol. 10, p. 369. 1903.

1. Kienitz-Gerloff, F. — Die Gonidien von Gymnosforangium clavariaeforme. Bot.

Zeit. Vol. 46, p. 389. 1888. " ,

1. Klebahn, H. — Die Wirtwechselnden Roslpilze. Berlin. 1904. (This masterly

work includes references to Klebahn's numerous contributions to the litera-

ture of the Rusts.)

2. Kulturversuche mit heteroecischen Uredineen. Zeitschr. f. Pflanzenkr.

Vol. 2, p. 337. 1892. ^

3. Kulturversuche mit Rostpilzen xii. Bericht (1903-1904). Zeitschr. f. Pflan-
zenkr. Vol. 15, p. 106. 1905.

1. Kuehn, J.— Der Rost der Runkelriibenblatter, Uromyces betae. Bot. Zeit, p.

540. 1869.

H 2

2l8

Literature.

I. Lagerheim, G. v. — Ueber Uredineen mit variablem Pleomorphismus. Tromso

Mus. Vol. 16, p. 105. 1894.

1. Laurent, E. — De Taction interne du sulfate de cuivre dans la resistance de la

pomme de terre au Phytofhthora infestans. Compt. rend., Vol. 135, p.

1040

1902

Lindroth, J. I. — Cecidomyia-larver, som ata rostsporer. Medd. Soc. Fauna.
Fl. Fenn. Vol. 26, p. 25. 1900.

1. Ludwig, F. — Ueber einen neuen Goodeniaceenrost aus Siidaustralien. Hedw.

Vol. 28, p. 362.

1889.

Ueber einige neue Pilze aus Australien. Bot. Centralb. Vol.

1890.

43i P- 5

Ueber neue australische Rostkiankheiten. Zeitschr. f. Pflanzenkrank-

heiten, Vol. 2, p. 130.

1892.

1893.

Ueber einige Rost-und Brandpilze Australiens. Ibid., Vol.

3. p. 137-

Lehrbuch der niederen Kryptogamen, p. 455. Stuttgart. 1892.

1. Magnus, P. — Ueberwinterung der Puccinia caricis. Verh. Bot. Ver. Prov.

Brand, Vol. 18, p. 27. 1885.

Ueber das Auftreten der Stylosporen bei den Uredineen.

Deutsch. Bot. Gesellsch., Vol. 9, p. 85. 1891.

Ber. der

Vol.

Zur Kentniss des Verbreitung einiger Pilze. Ber. Deutsch. Bot. Ges.,

10, p. 195

1892.

Die systematische Unterscheidung nachst verwandter parasitischer Pilze

auf Grund ihres verschiedenen biologischen Verhaltens. Hedwigia, Vol.

33> P- 362.

1894.

5
6

Fungi Pars. II. in J. Bornmueller, Iter Persico-turcicum. 1892-93.

Verhandl. Zool.-bot. Gesellsch. Wien, Vol. 49, p. 95.

i8qq.

Ueber eine Function der Paraphysen von Uredolagern. Ber. Deutsch.

Bot. Ges., Vol. 20, p. 334

1Q02

Kurze Bemerkung zur Biologie des Chrysanthemum-rostes. Centralb. f.

Bakt. Vol. 10, p. 575. 1903

1. Maiden, J. H. — Report on Botanic Gardens and Domains, p. 10. N.S. Wales.

1902.

1. Marchal, E. — De la specialisation du parasitisme ohez V Erysifhe graminis. '•

Compt. Rend., Vol. 135, p. 210. 1902.

De 1' immunisation de la Laitue (Lettuce) contre le Meunier. Ibid p

1067.

1902

1. Massee, G. — On the Presence of Sexual Organs in Aecidium. Ann. Bot., Vol.

2, p. 47.

1888.

- Notes on Exotic Fungi in the Royal Herbarium, Kew. Grev., Vol. 21

p. 4

1892.
Australian Fungi. Grev., Vol. 22, p. 17

1893.

1897.

Mycological Flora of the Royal Gardens, Kew. Kew Bull., p. 151.

5
6

8
9

Chrysanthemum Rust. Gardeners' Chronicle, 8th Oct., 1898.

The Cereal Rust Problem. Does Eriksson's Mycoplasma exist in Nature?

Natural Science, Vol. 15, p. 337. 1899.

To protect Cucumbers and Tomatoes from fungus. Journ. Roy. Hort.

Soc, Vol. 28, p. 142. 1903.

A Text-book of Plant Diseases, 2nd. Ed. London. 1903..

On the Origin of Parasitism in Fungi. Phil. Trans. Royal Soc, Ser. B,

Vol. 197, p. 7.

1904

cTer Uredineen in ihrer Abhangigkeit von Stand-
ee o

1. Mayus, O. — Die Peridienzellen

ortsverhaltnissen. Centralb. f. Bakt., Vol. 10. p. 644. 1903
1. McAlpine, D.— The Life-history of the Rust of Wheat. Bull. 14, Dep. Agric,

Victoria, pp. 22-32, pis. 2. 1891.

Rust of Wheat. Ibid. pp. 33, 34

■

1891.

45, 46.

Beet-leaf Rust or Blighted Mangel leaves

1891.

[Uromyces betae). Ibid, pp

Report cm Peach and Plum Leaf-Rust {Puccini fruni). Ibid, pp. 138,

144, pis. 3.

i8qi.

Wales.

Report on Rust in Wheat as Victorian Delegate. Proc. Conf., N.S.

1 891.

Literature.

219

6. McAlpine, D. — Ueber die Verwendung geschrumpfter Koerner von rostigem

Weizen als Saatgut. Zeitschr. f. Pflanzenkrankh., Vol. 2, p. 193. 1892.

7. Report on Rust in Wheat as Victorian Delegate. Proc. Conf. S. Aus-
tralia. i8q2.

8. The Undescribed Uredospores of Puccinia bur char diac. Vict. Nat.,

Vol. 10, p. 192. 1894.

9. Report on Rust in Wheat Experiments at School of Horticul-
ture, Burnley. Illustrated with maps and drawings, pp. 66. Dep. Agric,
Victoria, 1894.

10. Report on Rust in Wheat as Victorian Delegate. Proc. Conf., Queens-
land. 1894.

11. Australian Fungi. Proc. Roy. Soc, Victoria. Vol. 7, p. 214. 1895.

12. Systematic Arrangement of Australian Fungi, together with Host Index

and List of Works on the subject. Dep. Agric, Victoria. 1895.

13. Notes on Uromyces amygdali — a Synonym of Puccinia fruni Pers.

Proc. Linn. Soc, N.S. Wales. 2nd series, Vol. 10, p. 440. 1895.

14. Puccinia on Groundsel with Trimorphic Teleutospores. Ibid., p. 461.

1895.

-15. Australian Fungi. Agric. Gaz., N.S. Wales, Vol. 7, p. 752. 1895.

16. Ibid, p. 850. 1895.

17. Ibid. Vol. 7, p. 147. 1896.

18. Ibid, p. 299. 1896

19. Two Additions to the Fungi of New South Wales. Proc. Linn. Soc,

N.S. Wales. Vol. 21, p. 722. 1896.

20. New South Wales Fungi. Ibid. Vol. 22, p. 36. 1897.

21. Rust in Wheat Experiments, 1894-1897. Dep. Agric, Victoria, 1S97.

22. The Fungi on the Wheat Plant in Australia. Agric Gaz., N.S. Wales,

Vol. q, p. iooq. 1898.

23. Rust in Wheat during the dry season of 1897. Ibid. p. 1421. 1898.

24. On a micro-fungus from Mount Kosciusko, and on the first record of

Uncinula in Australia. Proc Linn. Soc, N.S. Wales. Vol. 24, p. 301. 1899.

25. Two Mallee Fungi. Vict. Nat., Vol. 16, p. 141. 1899.

26. Rose Rust. Journ. Dept. Agric, Vic, p. 81. 1902.

.27. Fungus Diseases of Stone-fruit Trees in Australia, and their Treatment.

Dept. of Agric, Victoria. 1902.

28. Cereal Rusts. Joura. Dept. Agric, Vic, p. 425 1902.

29. Peach and Plum Rust. Ibid. p. 617. 1902.

30. Australian Fungi, new or unrecorded, Decades V.-VI. Proc. Linn.

Soc, N.S. Wales. Vol. 28, p. 558. 1903.

31. Rerraf — A Rust-resisting Wheat. Journ. Dept. Agric, Vic, Vol. 2,

p. 531. 1904.

32. Diseases of Cereals, Rust and Take-all in Wheat. Ibid. p. 700. 1904.

33. Some Misconceptions concerning the Uredospores of Puccinia fruni

Pers. Ann. Myc, Vol. 2, p. 344. 1904

34. Note on the Arrangement of Teleutospores in Puccinia fruni Pers.

Ibid. p. 348. 1904.

35. Bobs — A Rust-resisting Wheat. Journ. Dept. Agric. Vic, p. 166. 1905.

36. A new genus of Uredineae — Urotnycladium. Ann. Mvc Vol. 3, p. 303.

1905.

1. Milesi, M., and Tra verso, G. B. — Saggio di una monografia del genere Tri-

fhragmium. Ann. Myc, Vol. 2, p. 143. 1904.

1. Molixeux, A. — Some Remarks about Red Rust. Journ. Agric, S. Australia.
. Vol. 8, p. 95. 1904.

1. Morrison, A. — New Victorian Micro-fungi. Victorian Nat., Vol. 11, pp. 90,

91. 1894.

2. Notices of Victorian Fungi. Ibid. pp. 119, 120. 1894.

1. Mueller, F. v. — Fragmenta Phytographiae Australiae — Supplement, Vol. 11, p.

96. 1880. (Fungi enumerated by Dr. Cooke.)

2. Census of the genera of plants hitherto known as indigenous to Australia.

Proc. Roy. Soc N.S. Wales, Vol. 15, pp. 254, 255. 1881.

3. Notes on Victorian Fungi. Victorian Nat., Vol. 2, p. 80. 1885.

4. Lecture on Rust in Cereals. Sandhurst, Victoria. 1865.

1. Mueller, Fritz.— Beitrage zur Kentniss <ler Grasroste. Eot. Centralb. Beihefte

10, p. 181. 1901.

x. Mueller, J. — Die Rostpilze der Rost-und Rubusarten und die auf ihnen vor-

kommenden Parasiten. Landw. Jahrb., Vol. 15, p. 719- 1886.

•

220

Literature.

i. Neger, F. W. — Beitrage zur Biologie der Erysipheen. Flora 90, p. 221

1902

1. Neumann, R. — Ueber die

Entwickelungsgeschichte der

gonien der Uredineen. Hedwigia, Vol. 33, p. 346.

Aecidien und Spermo-
1894.

5*
6.

i. Pfeffer, W. — Ueber chemotaktische

Bewegungen

von Bakterien, Flagellaten

und Volvocineen. Untersuch. Bot. Inst., Tubingen. Vol. 2, p. 582. 1888.

1. Plowright, C. B.

with the Barberry Aecidium (Aecidium berberidis Gmel.) Gard. Chron.
Ser. 2, Vol. 18, p, 231. 1882.

Podisoma junif

Reproduction of Heteroecious Uredines. Linn.

Ibid, p. 553.

Soc, London.

Botany*

Vol. 21, p. 368.

1884.

On the Life-history of certain British Heteroecismal Uredines. Quart

Micro. Sci., Vol. 25, p. 156.

1885.

A Monograph of the British Uredineae and Ustilagineae. London. 1889.

Barberry and Wheat Mildew. Gard. Chron., Vol. 23, p. 45

1898.

Prain, D. — Rust in Wheat in the Australian Colonies.

Agric. Ledger No.

i6r

Calcutta.

1897.

1. Prillieux, E.

Maladies des Plantes Agricoles.

Pt. 1, p. 242. Paris, 1895.

1. Ralph, T. S. — On the Aecidium affecting the Senecio vulgaris or Groundsel.

Victorian Nat., Vol. 7, pp. 18, 19

1890.

1. Rathay, E. — Untersuchungen ueber die Spermogonien der Rostpilze. Denkschrift.

Wiss Wien

1883.

1. Richards, H. M. — On some points in the development of Aecidia. Proc.

Amer. Acad. Sci., Vol. 31, p. 255.

189

1. Rodway, L. — Tasmanian

1897.

Fungi.

Proc. Roy. Soc, Tasmania, pp. 142, 143

1. Roze, M. E. — Le Puccinia chrysanthemi. Bull. Soc. Myc. France, Vol. 16

88.

1900

1. Saccardo, P. A. — Sylloge Fungorum. Vols. 7, 9, n, 14, 16, 17 1888-1905.

Mycetes aliquot australiensis. Hedw., Vol. 28, p. 126. 1889.

Bull. Soc. Myc. France, Vol. 5, p. 118.

Ibid.

Ibid. Hedw., Vol. 29, p. 155

Ibid. Hedw., Vol 31, p. 57

1889.

1890.
1893.

1. Saccardo, P. A., and Berlese, A. N.

Fungi australiensis.

Rev. M)

Vol.

7> P. 93

188

.-)

Salmon, E. S. — Mycological Notes. J

Cultural Experiments with

Phil. Trans., B., Vol. 197, p. 107

t c

Biologic
1904.

184. 1904

Forms'' of the Erysiphaceae-

DC.

Cultural Experiments with the Barley Mildew,

'fhe gr a minis

1904

1. Sappin-Trouffy, P. — La Pseudo-Fecondation chez les Uredinees el les pheno-

menes qui s'y rattachent. Compt. rend., Vol. 116, p. 1304.
Recherches histologiques sur la famille des Uredinees.

1893.

Le Botaniste,

Vol. 5, pp. 59-244

1896-97.

Shutt. F. T— The effect of Rust on the Straw and Grain of Wheat. Nor'-

W est Farmer. W

20th Oct., 1904

1. Smith, W. G. — Diseases of Field and Garden Crops. London.

1884.

1. Smith, R. E. — The Water relation of Puccinia as far a gi. Bot. Gaz., Vol.

p. 19

1904

1. Sorauer, P. — Das. Verhalten des Getreiderostes in trockenen und nassen

ren. Zeitschr. f. Pflanzenkr., Vol. 4, p. 121. 1894.

Die Predisposition der Pflanzen fur parasitare Krankheiten. Deutsch.-

Landwirts. Ges., Berlin.

1903

1. Sydow, P. and H. — Monographia Uredinearum. Vol. 1. 1Q02-04

Neue und kritische Uredineen.

i9°3

Literature.

O ^> J

.2.

5
6

jo.

i. Tassi, F. — La

Ruggine

Lab. Bot., Univ. di Siena. November, 1903
1. Tate, R. — A list of the

dei Crisantemi {Puccinia chrysanthemi Roze). Bull.

gals
4, P

Charas, Mosses, Liverworts, Lichens, Fungi, and Al-
of Extra-tropical South Australia. Proc. Roy. Soc. S. Australia, Vol.

12.

1881.

ri"*

J. G. O.

Red Riist : Its nature, approximate cause and probable cure.
Proc. Roy. Soc, S. Australia, Vol. 3,

P- l3
Ibid., Vol.

Contributions to the Fungal Flora of

Notes on Australian Fungi.

1879.

12, p. 150
Australia.

Australia, Vol. 13, p

1. Tenison-Woods, J. E.,

South Wales and Qu<
1880.

55. 1890.
and Bailey,

1889.
Proc. Roy. Soc. S.

F. M. — On some of the

Fungi

of New

Proc. Linn. Soc, N.S. Wales, Vol. 5, p. S3.

1. Thompson, E. H. — A Hand-book to the Insect Pests of Farm and Orchard

(including Fungi).

Dep.

Agric

of Tasmania.

Bull.

*> PP- 29-34

1892.

i. Thuemen, F. v. — Symbolae ad floram mycologicam Australiae. I. Grevillea,

Vol.

4. P
Ibid.

75
II.

1875-

Flora No. 28. 1878

■ 1st der Berberitzenrost

Oesterr. Landw. Wochenbl

notwendig

1883.

zur Erzeugung des Grasrostes?

1. Tranzschel, W

— Ueber einige auf Grund von irrtlimlicher
Nahrpflanzen aufgestelle Puccinia Arten.

Ann. Myc, Vol. 2

Bestimmung

der

p. 155. 1904

Moglichkeit, die Biologie wirtswechselnder Rostpilze auf Grun<
' Merkmale vorauszusehen. Arbeit, der Kais. St.

Naturf. Gesell. Vol. 35, p. 1. 1904.
1. Tryon, H. — Report on Insect and Fungus Pests, pp. 97, 196. 1889.

Petersburg,

Ann.

1894.5.

Vegetable Pathology, pp. 39, 40

3- "

4- *

6. -

8. -

Ibid. p. 39. 1895-6.

Ibid. p. 37. 1890-7.

Ibid. pp. 43
Ibid. p. 35

Ibid.

1897-8.

P
Ibid, p

30.
26.

44.

1898-9.
1 899- 1900.
1 900- 1.

1. Tubeuf, K. F. V., and Smith, Wm. G. — Diseases of Plants induced by Crypto-

gamic Parasites. London. 1897.

Ward, H. Marshall.— Illustrations of the Structure and Life-history of Puc-

1888.
Brit. Assoc. Rep., Glasgow, p.

cinia graminis. Ann. Bot., Vol. 2, p. 217
The Bromes and their Brown Rust.

836.

1901.
The Bromes and their Rust-fungus {Puccinia dispersa). Ann Bot., Vol.

15. P- 56°-

1 go 1

On the question of " Predisposition " and " Immunity" in Plants. Proc

Proc. Roy. Soc,

Camb. Phil. Soc, Vol. n, p. 326. 1902

On pure Cultures of a Uredine, Puccinia disp

Vol. 69, p. 451. 1902. .

On the Relations between Host and Parasite in the Bromes and their

Brown Rust, Puccinia dispersa. Ann. Bot., Vol. 16, p. 233. 1902. ^

Experiments on the effect of mineral starvation on the Parasitism of the

Uredine-fungus, Puccinia dispersa, on species of Bromus. Proc Roy. Soc,
Vol. 71, p. 138. 1902.

Further Observations on the Brown Rust of the Bromes, Puccinia dis-
persa, and its adaptive Parasitism. Ann. Myc, Vol. 1, p. 132. i9°3-

On the Histology of Uredo dispersa Eriks., and the Mycoplasm Hypo-

thesis. Phil. Trans. B., Vol. 196, p. 29. 1903.

Recent Researches on the Parasitism of Fungi. Ann. Bot., Vol.

*9> P

1Q05

r. Webber,

Peridial Cell Characters in the Classification of the Uredineae.

Amer. Nat., Vol. 24, p. 177. 1890.

1. Winter, G. — Exotische Pilze II. Hedwigia, Vol. 24, p

Rev. Myc, Vol 8, p. 208.

Fungi australienses.

18S6.

1885.

x. Woroxix, M. — Untersuchungen ueber die Entwickelung des Rostpilzes (Puc*

cinia helianthi), welcher die Krankheit der Sonnenblume verursacht.

Bot,

Zeit., p. 677.

1872.

212

Explanation of Plates

PLATE A.

(All Figures mat. size.)

PUCCINIA TRITICINA on WHEAT

Fig.
i. Uredosori on upper surface of leaf.

2. Teleutosori on under surface.

PUCCINIA GRAMINIS ON WHKAT.

3. Uredosori on upper and under surface, also on sheath

4. Teleutosori on stem.

■■

• I

-*1

- ? '

•

: i

C C Brzt/JtbanJc.De/,.

-V--

.-J

D M:'Alpif?& Dvjc

• I

PLATE B

■

• *

A c>/i- :za ^i > ' Pnntr

OAT, RYE-GRASS, AND BARLEY RUSTS.

}

224

Explanation of Plates

PLATE B.

(All Figures nat. size.)

PUCCINTA LOLII AVENAE on OATS

Fig.

5. Uredosori on under surface of leaf.

6. Teleutosori on under surface

PUCCINIA LOLII on RYEGRASS.

7. Uredosori on under surface of leaf

8. Teleutosori on upper surface.

PUCCINIA SIMPLEX cn BARLEY

9. Teleutosori on sheath and blade.

10. Uredosori on under surface of leaf

•

PLATE A

CCBrtitLtbcwkBeL

JL 1

)

if'. *

D MPJl^ui^Jhrexit

RSBrauui. u<rvt>Pnn£er.

WHEAT RUST.

4 4

'■'•■

•

226

Explanation of Plates.

PLATE C.

{All Figures nat. size.)

PUCCINIA BROMINA on SOFT BROME (Bromus mollis)*
Fig-

ii. Uredosori on under surface of leaf.

. Teleutosori on under surface.

13. Teleutosori on stem.

PUCCINIA MAYDIS ON MAIZE (Zea mays)

14. Uredosori on under surface of leaf.

15. Teleutosori on young stem

PUCCINIA THUEMENI on CELERY (Apium

16. Uredosori on upper surface of leaf

17. Uredosori on under surface.

;

• l

PLATK ('

* »■

« » ■

JVr

C C £rUtlehank Del

I)McALpiKt Direxu

\ S Brow,, (kv4 * printer

BROME-GRASS, MAIZE, AND CELERY RUSTS.

•

228

Explanation of Plates

PLATE D.

{All Figures nat. size.)

PUCCINIA CICHORII on CHICORY (Cichorium intybus)
Fig.

18. Chiefly uredosori on lower surface of chicory leaf.

PUCCINIA PRUNI on PEACH AND APRICOT (Prunus persica ano

P. armeniaca).

19. Uredo and teleutosori on lower surface of peach leaf.

20. Uredo and teleutosori on lower surface of apricot leaf

PLATE I)

^M'(^k

CC.BruttUhank.Del.

DMcAlpuiei Direxiz

F S Bra j/ w T p -utter

CHICORY, PEACH, AND APRICOT RUSTS.

■

y -

;

230

Explanation of Plates.

PLATE E.

(All Figures nat. size.)

PUCCINIA CHRYSANTHEMI on CHRYSANTHEMUM
Fig.

ai. Uredosori on under surface of leaf.

PUCCINIA CALENDULAE ON MARIGOLD (Calendula officinalis)
22. Aecidia and teleutosori on under surface of leaf.

23. Uredo and teleutosori on stem.

PUCCINIA HEDERACEAE on NATIVE VIOLETS.

24. Uredo and teleutosori on under surface of leaf of Viola hederacea

25. Aecidia on under surface of leaf of V. betonicijolia.

PUCCINIA DISTINCTA on DAISY (Bellis perennis).

26. Aecidia and teleutosori on upper surface of leaf, showing the dark teleutofori

surrounding the other.

PLATE E

CCBrUdeiasAjtl

DWAlpuze,, Direxvt

R. S.Braui GwtPrviUr

CHRYSANTHEMUM, MARIGOLD, VIOLET, AND DAISY RUSTS.

■ ■

•

■

•

■

•

■

•

■

- f

•

■

•

232

Explanation of Plates

PLATE F.

{All Figures nat. size.)

PUCCINIA LAGENOPHORAE on LAGENOPHORA BILLARDIERL

Fig.

27. Aecidia and teieutosori on both surfaces of leaves.

PUCCINIA MALVACEARUM on LAVATERA PLEBEIA,

28. Teieutosori on under surface of leaf and on leaf -stalk.

PUCCINIA MORRISONI on PELARGONIUM AUSTRALE.

29. Aecidia, uredo, and teieutosori on under surface of leaf, and teieutosori on stem

PLATE F

CC£nttUi?a7ihfDd

DMC Alp 1716 Direjcio

. . "ma ■ . ' r> • v-

LAGENOPHORA, MALLOW, AND PELARGONIUM RUSTS.

•

234

Explanation of Plates

PLATE G.

(All Figures nat. size.)

UKOMYCES CARYOPHYLLINUS ON CARNATION (Dianthus caryophyllus).

Fig.

30. Uredosori on upper surface of leaf.

31. Teleutosori on both surfaces.

UROMYCES TRIFOLII on WHITE CLOVER (Trifolium repens)

32. Aecidia, uredo, and teleutosori on leaf.

UROMYCES VESICULOSUS on ZYGOPHYLLUM GLAUCESCENS.

33. Uredo and teleutosori on both surfaces ot leaves and on stems.

UROMYCES HARDENBERGIAE on HARDENBERGIA MONG'FHYLLA.

34. Uredosori on under surface of leaf.

PLATK G

CCBnMeiank.Vel.

D ACcllfiVU,, Dtrtxib

R S Bravi, 6ovtPnnler.

CARNATION, CLOVER, AND OTHER RUSTS.

■

•

236

Explanation of Plates.

PLATE H.

[Natural Size.)

UROMY

BETAE ON MANGEL (Beta vulgaris)

Fig.

35. Uredo and teleutosori on under surface of leaf.

This leaf was obtained from Mangels, the roots of which had been planted in
August for seed, and in January the plants were fully six feet high, with abundance
of foliage covered on both sides with rust.

PLATE II

* t • •• *

c I >

' '

• ' ••

•

* 9 ,-

MANGEL RUST

' .

' '

. -*

238

Explanation of Plates.

•

PLATE I.

(All Figures nat. size.)

Fig.

MELAMPSORA LINI on NATIVE FLAX (Linum marginale)

36. Uredo and teleutosori on stem.

PHRAGMIDIUM SUBCORTIC1UM on SWEET-BRIER (Rosa rubiginosa).

37. Aecidia (Caeomata) showing large swollen patches on stems and small patches on

lower surface of leaves.

PHRAGMIDIUM BARNARDI on NATIVE RASPBERRY (Rubus parvifolius).

38. Ochraceous uredosori on lower surface of leaves and flower-stalks, powdery

teleutosori accompanying uredosori.

PLATE I

Y ' BnttUharJi. Dd

'■' ft ■ ■■ DirexU

$. /train C*>\* P>

SWEET BRIAR, RASPBERRY, AND FLAX RUSTS.

■

•

■

•

■

240

Explanation of Plates

•

PLATE

AECIDIUM on DANTHONIA

Fig.

39. Leaves with aecidia in dense clusters

nat. size

40. Cluster of aecidia on leaf ...

X 15

41. Section of aecidium showing peridial wall and spores in chains

42. Chain of aecidiospores

X 50

.. X 1000

43. Single spore detached and more or less rounded

.. X 1000

44. Pseudoperidial cells seen from the surface

.• X

PLATE J

C. CJtrzttUbcuiJi. Dd

DIFAlputfi, Diructi,

RSJfrtusis.GovtPwtlar

WALLABY-GRASS RUST.

242

Explanation of Plates.

•

PLATE I.

(All Figures X 250 unless otherwise stated.}

PUCCINIA.

Fig.

1. Portion of sheath of barley (Hordeu?n vulgare) showing numerous crowded black

dots, the teleutosori of Puccinia simplex, the

barley

spring or

• • •

orange rust of
• .. nat. size

2. Portion of sheath of wheat [Triticum vulgare), the broad dark lines representing

the uredosori of P. graminis, summer rust of wheat

nat. size

3. Portion of flag of wheat, the small dark dots representing the uredo and teleu-

tosori of P. triticina, the spring or orange rust of wheat

nat. size

4. Uredospores of P. simplex from barley, the numerous germ pores appearing as

paler dots scattered promiscuously over the face of the spore.

5. Uredospores of P. graminis from wheat, w7ith their three equatorial germ pores.

6. Uredospores of P. triticina from wheat with germ pores promiscuously scattered

over the face.

7. Teleutospores of P. graminis from wheat, the thickened apex being very pro-
nounced.

8. Teleutospores of P. graminis from wheat, upper cells checked in their develop-

ment by some unknown cause as shown by their pale colour, not due to ger-
mination having already occurred.

9. Teleutospores of P. simplex from barley, only one being two-celled.

10. Section of teleutosorus of P. triticifia on wheat, with paraphyses at left.

Note. — The whole of the photomicrographs in this and the succeeding plates were
taken with the horizontal laboratory camera made by Messrs. Watson and Sons, of
London, using their Holoscopic lens of 12 mm. .65 N.A. to obtain the magnifica-
tion of 250 diams., and the 6 mm. .95 N.A. for 500 diams.

Platk I

{,. II. Robinson, Phot.

Sut. size and v iou

PUCCINIA.

GRAMINEAE.

•

••

■

• r

-
-

•

244

Explanation of Plates

PLATE II.

(All Figures X 250.)

PUCCINIA.

Fig.

ii. Section of teleutosorus of Puccinia lolii avenae on oat (Avena saliva) with severaE-

unicellular as well as bicellular teleutospores.

12. Group of teleutospores of P. beckmanniae on Beckmannia erucaeformis

13. Section of teleutosorus of P. festucae on Festuca fratensis.

14. Section of teleutosorus of P. lolii on Lolium -perenne.

15, 16. Groups of teleutospores of P. tnaydis on Zea mays, including one four

celled spore.

17. Teleutospores of P. flavescentis on Stifa flavescens.

18. Section of teleutosorus of P. magnusiana on Phragmites communis.

19. Five teleutospores of P. tefferi on Phragmites communis, showing the extremeljT

long pedicels.

Plate II.

G. II. Robinson, Phot.

x 250

PUCCINIA.

GRAMINEAE.

■

246

Explanation of Plates.

PLATE III.

{All Figures X 250.)

PUCCINIA.
Fig.

20. Section of teleutosorus of Puccinia anthoxanthi on Anthoxanthum odoraium, one

spore with a septate pedicel laterally inserted.

21. Group of teleutospores of the same, one with a much inflated pedicel.

22. Teleutospores of P. -poarum on Poa annua.

23* Teleutospores of P. ferflexans on Alofecurus geniculates

24. Teleutospores of P. cynodontis on Cynodon dactylon.

25. Section of teleutosorus of P. agrofyri on Agrofyron scab rum

26. Group of teleutospores of P. imfatientis on Elymus condensatus.

27. Group of teleutospores of P. agrostidis on Deyeuxia forsteri

28. Section of teleutosorus of P. bromina on Bromus mollis.

Platk III.

G. H. Robinson, Phot.

x 250

PUCCINIA.

GRAMINEAE.

• *

' »

■

t •

. ■

•J SiiS ,; l i- • v * . « •

438

248

Explanation of Plates

PLATE IV.

(All Figures X 250 .)

PUCCINIA.

Fig.

29. Group of teleutospores of Puccinia caricis on Carex breviculmis, Killara, Vic-
toria.

30. Group of teleutospores of Puccinia caricis on Carex stricta^ Berlin, Germany

(Sydow, Uredineen, 460).

31. Teleutospores of P. longisfora on Carex caesfitosa.

32. Teleutospores of P. cyferi on Cyferus rotundas.

33. Group of teleutospores of P. tenuisfora on Luzula eamfestris, Murramurrangbong

Ranges, Victoria.

34. Teleutospores of P. obscura on Luzula camfestris, Berlin, Germany (Sydow, Ure-
dineen, 1076). The spores of this species are differently shaped, much thicker
in the wall and much darker than those of P. tenuisfora.

35. Uredospores of P. juncofhila from sorus containing both uredo and teleutospores

on 1 uncus maritimus.

36. Teleutospores of the same.

•

Plati; I \ .

G^ H. Robinson, Phot.

x 250.

PUCCINIA.

CYPERACEAE and JUNCACEAE

1 2

250

Explanation of Plates.

PLATE V.

(All Figures X 250.)

PUCCINIA.

Fig.

37. Teleutospores and mesospores of Puccinia wurmbeae on Wurmbea dioica

38. Uredospores of P. burchardiae on Bur char dia umbellata.

39. Teleutospores, one three-celled, of the same.

■

40. Teleutospores and mesospores of P. haemodori on Haemodorum sp.

41. Teleutospores and mesospores of P. hyfoxidis on Hyfoxis glabella.

42. Teleutospores, two three-celled, of P. dichondrae on Dichondra re-pens.

43. Teleutospore and uredospore of P. tnussoni on Ruellia australis^ Richmond River,

New South Wales.

44. Teleutospores of P. ruelliae on Ruellia strefens, Columbus, Ohio, U.S.A. (W.

A. Kellerman, Ohio Fungi, 130).

Plate V.

(i. 1J. Robinson, Phot.

x 2."".

PUCCINIA.

LILIACEAE, HAEMODORACEAE, AMARYLLIDEAE, and ACANTHACEAE.

•

■

252

Explanation of Plates.

PLATE VI.

Fig.

{All Figures X 250.)

PUCCINIA.

45. Group of teleutospores of Puccinia carissae on Carissa ovata

46. Section of teleutosorus of P. alyxiae on Alyxia buxifolia, with teleutospores and

mesospores.

47. Teleutospores of P. gilgiana on Leschenaultia linarioides.

48, 49. Teleutospores and mesospores of P. saccardoi on Goodenia geniculate

50. Teleutospores and mesospores of P. brunoniae on Brunonia au straits, with one

three-celled and one four-celled teleutospore.

51. Teleutospores of P. aucta on Lobelia ancefs. (From a drawing by C. C. Brittle-

1 bank from the original material from Berkeley in the National Herbarium,
Melbourne).

Plate VI

G. H. Robinson, Phot

x 250

PUCCINIA.

APOCYNACEAE, GOODENIACEAE, and CAMPANULACEAE

•

254

Explanation of Plates.

PLATE VII.

(All Figures X 250.)

PUCCINIA.

Fig.

52. Teleutospores and mesospores of Puccinia t as manic a on Senecio vulgaris, one

teleutospore being three-celled.

53. Teleutospores and mesospores of P. angustifoliae on Scorzonera angustifolia.

54, 55. Sections of teleutosori of P. lagenofhorae on Lagenofhora billardieriy show-

ing mesospores and teleutospores.

56. Teleutospores and mesospoies of P. helianthi on Helianthus annuus.

57. Teleutospores of P. gnafhalii on Gnwphalium jafonicum.

58. Teleutospores and mesospores of /\ calocefhali on Calocefhalus drummondii.

59. Teleutospores and mesospores of P. erechtitis on Erechtites quadridentata.

Plate VII.

G. II. Robinson, Phot

x 2.")0

PUCCINIA.

COMPOSITAE

*5«

Explanation of Plates

Fiu.

PLATE VIII.

{All Figures X 250.)

PUCCINIA

60. Section of teleutosorus of Puccinia cinerariae on Cineraria sp. cult, with teleuto-
spores and mesospores.

61. Three uredospores and several teleutospores of P. cichorii on Cichorium intybus

62. Uredospores and teleutospores of P. hyfochoeridis on Hyfochoeris radicata.

63. Teleutospores of the

64, Two uredospores and numerous teleutospores of P. cyani on Centaur ea cyanus.

65. One mesospore and several teleutospores of P. calendulae on Calendula officinalis

66. Mesospores and teleutospores of P. brachycomes on Brachycome ciliaris.

67. Teleutospores and mesospores of P. distincta on Bellis ferennis.

Platk VII!

('•. H. Robinson, Phot

x 250

PUCCINIA.

COMPOSITAE

258

Explanation of Plates

PLATE IX.

(All Figures X 250 unless otherwise stated.)

* •

'lg-

PUCCINIA.

68. Three uredospores and four teleutospores of Puccinia thuemeni on Afium fros-

tratum^ showing thickened apex of uredospore and slight warting of teleuto-

spore

Beaumaris, Victoria.

69. Uredospores and teleutospores from same, but in greater variety.

70. One uredospore and two teleutospores from the same, but more highly magnified,

and showing more clearly the roughened epispore of both uredo and teleuto-

spores

• *

• • •

• •

• *

x 500

71. Teleutospores of P. thuemeni on Afium graveolens, from Brighton, Victoria.

72. Uredospore and teleutospore from the same as 71, showing roughened epispore

in both forms more clearly

• •

• • .

X 5°°

73. Uredospores and teleutospores of P. afii on Afium graveolens, Berlin, Ger-
many. (Sydow, Uredineen, 558.)

74. Teleutospores as in Fig. 73, showing distinctly smooth epispore

X 500

75. Teleutospores of P. btillata on Aethusa cynafium^ with coarsely warted epispore.

(Sydow, Uredineen, 1261.)

• •

c •

X 5°°

76. Teleutospores of P. xanthosiae on Xanthosia fusilla.

Platk IX

G. II. Robinson, Phot.

x 230 ami 5u«>.

PUCCINIA.

UMBELLIFERAE.

z6o

Explanation

Plates.

PLATE X.

(All Figures X 250.)

PUCCINIA.

Fig.

77. Teleutospore of Puccinia oliganthae on Asferula oligantha

78. Teleutospores, some having germinated, of P. cofrosmae, on Cofrosma hirtella

79. Uredospores and teleutospores of P. efilobii-tetragoni on Efilobium glabel/um,

Murramurrangbong Ranges, Victoria.

80. Uredospores and teleutospores of P. efilobii-tetragoni on Efilobium montanum.

(Sydow, Uredineen, 1369.)

Si. Teleutospores of P. efilobii DC. on Efilobium roseum, the teleutospores having

much thinner walls, and being more variable in shape than those of P.
efilobii-tetragoni and also finely verrucose. (Sydow, Uredineen, 1418.)

82. Teleutospores of P. gei on Geum reni folium.

83. Uredospores of P. fruni on Prunus fcrsica.

84. Paraphysis and uredospores of P. fruni on Prunus fcrsica.

85. Teleutospores of P. fruni on Prunus fersica^ rather smaller than average

86. Teleutospores of P. fruni on Prunus domestica.

87. Uredospores and teleutospores of P. zorniae on Zornia difhylla.

Platk X.

G. II. Robinson, Phot.

x 25u

PUCCINIA.

RUBIACEAE, ONAGRACEAE, ROSACEAE, and LEGUMINOSAE.

262

Explanation

Plates.

Fig.

PLATE XI.

(All Figures X 250.)

PUCCINIA.

88, 89. Teleutospores of Puccinia tetragoniae on Tetragonxa imflexicoma, one with

nearly vertical septum in upper cell.

90. Teleutospores, strongly warted, of P. ludwigii on Rumex flexuosus.

91. Teleutospores strongly warted, of P. ludwigii on Rum ex brownii, one spore being

Trifihragmium-Wke.

92. Uredospore and teleutospores of P. acetosae on Rumex arifohus^ all the latter

with smooth walls. (Sydow, Uredineen, 954.)

93. Teleutospores of P. mue/ilnibrcfciar on MuehlenbecTzia adfressa.

94. Teleutospores and mesospore of P. diclsiana on Threlkeldia sp.

95. Uredospores of P. kochiae on Koclria sedifolia, showing the characteristic nume-
rous germ pores.

96. Teleutospores of P. kochiae on Kochia sedifolia.

Plate XI

G. II. Robinson, Phot.

x UoO

PUCCINIA.

FICOIDEAE, POLYGONACEAE, and CHENOPODI ACEAE.

> '

264

Explanation of Plates

PLATE XII.

(All Figures X 250.)

PUCCINIA.

1 or

97. Teleutospores of Puccinia arenariae on Stellaria media

98. Teleutospores of /\ -plagianthi on Plagianthus sidoides, the epispore

slightly channelled.

being

99. Teleutospores (stained) of P. malvacearum on Malva rotundij alia.

100. Teleutospores and mesospore of P. malvacearum on Lavatcra flebeia.

101. Teleutospores, mostly one-celled, of P. heterosfora on Abutilon crisfum

T02. Teleutospores and mesospores of P. morrisoni on Pelargonium australe.

103. Uredospores of P. geranii-filosi on Geranium filosum.

104. Teleutospores of the

same, the single germ pore being

noticeable as a paler

channel through the apex of the cell wall.

1'I.ATK XII

G. H. Kobinson, Phot.

x 250

PUCCINIA.

CARYOPHYLLACEAE, MALVACEAE, and GERANIACEAE

-. — _ '

266

Explanation

Plates.

Fig.

PLATE XIII.

(All Figures X 250 unless otherwise stated.)

PUCCINIA.

05. Teleutospores of Puccinia eriostemonis on Eriostemon myo-poroidcs.

106. Teleutospores of P. correae on Correa lawrenciana.

107. Teleutospores and mesospores of P. boroniae on Boronia sfinescens.

108. Teleutospores and mesospores of P. fritzeliana on Tremandra stelligera, some

teleutospores having germinated.

109. Teleutospores of P. hederaceae on Viola hederacea.

no. Teleutospores of the same, more highly magnified, to show more clearly the

slightly warted epispore

- . -

• • •

• *

• •

X 500

in. Teleutospore of P. aegra on Viola tricolor with smooth epispore. (W. B. Grove

in Rabh. Fungi Europaei, 31 13)

• • •

• •

X 5°°

112. Teleutospores of P. violae on Viola arenaria, wtih smooth epispore. (Sydow,

Uredineen, 286)

- • •

• • •

X 5°°

•

L'late Mil

G. II. Robinson, Phot.

x 250 ami 5l0

PUCCINIA.

RUTACEAE, TREMANDREAE, and VIOLACEAE

■

268

Explanation of Plates

PLATE XIV.

(All Figures X 250.)

PUCCINIA GRAMTNIS on VARIOUS GRASSES

Fig.

113. Teleutospores and mesospores on wild oat, Avena fatua.

114. Teleutospores on barley, Hordeum vidgare.

11^, 116. Teleutospores, mesospores, and uredospore, one of the first three-celled, on

native barley, Echinofogon ovatus.

117. Teleutospores on cocksfoot, Dactylis glomerata.

118. Teleutospores and uredospores on silver grass, Festuca bromoides

ii(). Teleutospores and uredospores on an imported barley grass, Hordeum secalinum

120. Teleutospores on small canary-seed grass, Phaiaris minor.

121. Teleutospores on native wheat grass, Agrofyron scabrum

122. Teleutospores on Amfhibromus neesii.

I'j.ATK XI \

G. H. Robinson, Phot.

X 2'MI

PUCCINIA GRAMINIS ON VARIOUS GRASSES.

I ;

■

•

270

Explanation of Plates.

PLATE XV.

(All Figures X 250 unless otherwise stated.)

GERMINATION of TELKUTOSPORES of PUCCINIA MALVACEARUM.

Fig.

123. Germinating teleutospore, from each cell of which a promycelium has been

pushed forth, cell contents collected towards apex of tube, and segmentation
commenced.

124. Germinating teleutospore at a later stage, the promycelium having become divided

into four segments, and comparatively stout tubes emitted from each

seg-

ment.

125. Two germinating teleutospores, in the one on the left only the two median seg-

126.

128.

ments of the promycelium as yet bear sporidiola.
Germinating teleutospore with stout elongated and

contorted filaments

arising

from the promycelial cells, those from the two median cells ultimately giving
rise to abnormally small sporidiola.

127. Germinating teleutospore in wrhich the upper two promycelial cells have produced

sporidiola, while in the third from the apex the formation of the sporidiolum

is just commencing.

Germinating teleutospore, the promycelium bearing three

sporidiola on stunt

The sporidiolum second from the base is already
germinating and putting forth a germ tube extending across to the promy-

apex has not so far produced a

elongated sterigmata.

celium.

The segment second from the

sporidiolum, though a stout elongated filament has been emitted.

129. Promycelial spores

130. Germinating promycelial spore

• -

- - -

X 500

• •

• • *

• • •

X 5°°

Note. — All the above were grown in droplets of water on slides in a moist cham-
ber, and prepared for photographing by being first dried, then fixed with a saturated
solution of mercuric bichloride, washed, rinsed in dilute acetic acid, washed again,
and after drying stained with alcoholic Bismarck brown, again washed and mounted
in glycerine and water.

Plate

G. H. Robinson, Phot.

x 250 & 50 1

PUCCINIA.

TELEUTOSPORE GERMINATION OF P. MALVACEARUM.

•

272

Explanation of Plates

* •

tig.

PLATE XVI.

[All Figures X 250.)

UROMYCES.

131. Group of teleutospores of Uromyces danthoniae on Danthonia semi annularis

132. Teleutospores of U. ehrhartae on Micro Laena [Ehrharta) stifoides.

133. Teleutospores of U . tenuicutis on S-porobolus indicus.

1 ^4. Two one-celled and one two-celled teleutospore of U . tricorynes on Tricoryne

elatior.

135. Teleutospores of U . tricorynes on Tricoryne elatior.

136. Uredospores of U . thelymitrae on Thelymitra antennifera

137. Teleutospores of the same

138. Teleutospores of U. orchid earitm on Chilo glottis difihylla, from original type

material in National Herbarium.

130. Teleutospores of U . microtidis on Microtis forri folia from type material from

Massee.

Platk XVI

G. H. Robinson, Phot.

X -■"•"

UROMYCES

GRAMINEAE, LILIACEAE AND ORCHIDACEAE

274

Explanation of Plates

Fig.

PLATE XVII.

(All Figures X 250.)

UROMYCES.

140. Group of teleutospores of Uromyces fuccinioides on Selliera radicans.

141. Teleutospores of U . as ferulae on As ferula oligantha.

142. Uredospores and teleutospores of U. trifolii on Trifolium refens

143. Uredospores of U . hardenbergiae on Hardenbergia monofhylla.

14a. Teleutospore of the same.

145. Two uredospores of same at top and one teleutospore below

146. Section of teleutosorus of U . atriflicis on Atriflex semibaccata

spores are faintly striate with one prominent germ pore at apex.

The teleuto-

147. Two teleutospores of same at left and four uredospores at right.

148. Uredospores of U. brtae on Beta vulgaris

149. Teleutospores of same with prominent hyaline apiculus.

Platk XVII

G. II. Robinson, Phot.

UROMYCES.

GOODENIACEAE, RUBIACEAE, LEGUMINOSAE, AND CHENOPODIACEA

•

■

276

Explanation of Plates.

PLATE XVIII.

(All Figures X 250 unless otherwise stated.)

UROMYCES.

"Ficr

150. Uredospores of Uromyccs folygoni on Polygonum aviculare

151. Teleutospores of same.

152. Uredospores of U. caryofhyllinus on Dianthus caryofhyl/us, with prominent

scattered germ pores.

153. Teleutospores and one uredospore of same.

154. Section through leaf of Dianthus caryofhyllus, showing uredosorus of U. caryo-

-phyllinus with two pycnidia of Darluca filum growing upon it, and two

young pycnidia on opposite surface of leaf

• • •

155. Uredospores of U. vesiculosus on Zygofhyllum glaucescens.

156. Teleutospores of same.

157. Two-celled teleutospore of same

X 5°

Platk XVIII

G. H. Robinson, Phot.

X 50 & 250

UROMYCES.

POLYGONACEAE, CARYOPHYLLACEAE, AND ZYGOPHYLLACEAE

■

■ >

■•

, Si

278

Explanation

Plates.

Fig.

PLATE XIX.

{All Figures X 250 unless otherwise stated.)

UROMYCES.

158. Teleutospores, seen from above, of Uromyces fusisforus on Acacia nerii folia

159. Side view of same.

160. Uredospores of same, with pronounced apiculus and equatorial band of germ

pores.

UROMYCLADIUM.

161. Teleutospores of Uromycladium simplex on Acacia fycnantha, part of the

pedicel being still attached with the remains of the vesicle. (Stained).

162. Young teleutospores of the same showing the vesicles still attached. (Stained).

16}. Portion of flake of gummy material on leaf of Acacia fycnantha, in which nume-
rous teleutospores of U. simplex are embedded, nearly all germinating and

producing sporidiola.

164. Germinating sporidiolum of same. (Stained)

• •

x 5°°

165. Uredospores of same, being very similar to those of Uromyces fusisforus, fig.

160, though somewhat shorter and broader and' possessing more germ pores.

Pl.ATK XIX

G. H. Robinson, Phot.

X 250 & 500,

UROMYCES AND UROMYCLADI UM.

LEGU M I N OSAE-ACAC I A.

■

438

280

Explanation of Plates.

PLATE XX

{All Figures X 250.)

UROMYCLADIUM MARITIMUM.

Fie.

j66. Section of teleutosorus of V \ maritimum on Acacia longifolia^ the mature

teleutospores having the remains of the vesicles attached to the pedicels.

167. Group of mature teleutospores of the same.

168. Formation of teleutospore head ; a young cluster, showing the two young spores

at the apex, and the lateral vesicle below the septum. (Stained.)

169, 170. Similar heads, more advanced. (Stained.)

171. Mature teleutospore head, two teleutospores above and one vesicle below.

172.

Abnormal development of head of teleutospores, the lateral vesicle being re-
placed by a true spore, and a septum placed in the stalk beneath it. In the
normal head of two spores and a vesicle there is no septum below the vesicle.

173. Teleutospores germinating while still attached to each other : two sporidiola on

one promycelium, others so far undeveloped.

174. Uredospores with prominent equatorial germ pores and serrated epispore

thickened and dentate at apex.

75. One-celled spores (mesospores ?), occasionallv found " intermixed with uredo-
spores.

176. Immature two-celled colourless spore, of same character, from .uredosorus.

(Stained).

G. 11. Robinson, Phot.

X 2 50.

UROMYCLADIUM MARITIMUM

LEGUMINOSAE-ACACIA LONGIFOLIA

K 2

Explanation of Plates

PLATE XXI.

(All Figures X 500.)

Fig.
177. Two

UROMYCLADIUM MARITIMUM.

younnr

teleutospore heads,

right being younger than
T2 and T3 in next figure.

arising from common stalk, the one on the
that on the left, and both intermediate between

(Stained.)

178. Main stem,

bearing teleutospore

heads in various stages of development; T,

filament with two septa, the upper two segments destined to become teleuto-
spores, the lowermost a vesicle ; Tl, similar filament, 9omewhat more ad-

vanced, the vesicle beginning to push out from the lowermost segment; T2,
similar filament still more advanced, the uppermost cell distending, in the

next growth is taking place at the side, and in

the lowest the vesicle has

attained considerable development; T3, nearly mature head, with twro dark
teleutospores; U, basidium of a detached uredospore. (Stained.)

179. Abnormal teleutospore head of three teleutospores, a septum beneath the lower-

most.

r8o. Normal teleutospore head, with two teleutospores above septum and a vesicle

below, there being no septum below the vesicle.

1S1. Teleutospore head in which the vesicle is apparently at the apex, but this is

probably due to displacement in mounting.

182, 183. Normal teleutospore heads.

184. Uredospores with prominent germ pores, serrated epispore thickened and dentate

at apex.

I'LATI. XXI

(J. II. Robinson, Phot.

-I- ;uu

UROMYCLADIUM M4RITIMUM.

LEGUMINOSAE-ACACIA LONGI FOLIA.

2«4

Explanation of Plates.

PLATE XXII.

(All Figures X 250 unless otherwise stated.)

UROMYCLADIUM.

Ftp

1 i^-

185, 186. Uredospores of U. robinsoni on Acacia melanoxylon. The solitary large

smooth-walled spore (?) in Fig. 186 is difficult of interpretation.

187. Young and mature teleutospores of same, with and without attached vesicles.

(Stained.)

188. Mature teleutospores of same, with vesicles.

189. Mature teleutospore of same germinating, the sporidiola just about to be formed.

(Stained.)

190, 191, 192. Successive stages in the development of a teleutospore head of U.

tefferianum on Acacia salicina^ the striated epispore being clearly shown in
192. (Stained.) ... ... ... ... ... ... X 500

193. Single teleutospore of same, seen from above, and showing striated mark-

• • • ••• ••• • • • ••• ••• ••• ••• ^\ ^^^^^

ings

194. Mature teleutospore head of same.

of poor condition of material)

(The markings are indistinct on account

X 500

• • •

195. Detached teleutospores of same.

Platk XXII

4
1

G. II Robinson, Vt.vi

x 250 & 510.

UROMYCLADIUM.

LEGUMINOSAE-ACACIA.

■

•

286

Explanation of Plates.

PLATE XXIII.

(All Figures X 250 unless otherwise stated.)

UROMYCLADIUM.

196. Uredospores of U. notabile on Acacia notabilis.

Original

Uredo notabilis Ludw.
caustic potash.)

type material of

(Somewhat swollen by lengthened treatment with

197. Uredospores of the same, showing the net-like surface markings

X 500

iq8. Compound stem, with basidia and uredospores of the same.

199. Compound structure from uredosorus of same, being probably commencement

of teleutospore formation

• • •

• •

X 5°°

200. Uredospores of U. notabile on Acacia dealbata.

201, 202. Uredospores of the same, showing the net-like surface markings... X 500

203. Portion of spermogonium of the same, with basidia bearing spermatia in

chains. (Stained)

* • •

• • •

X 500

204. Cluster of three immature spores of the same, attached to their stalk. (Stained.)

205. Mass of mature teleutospores of the same, separated from each other, from

Acacia decurrens.

206. Mass of scarcely mature teleutospores, separated from each other, of V . tef-

ferianum, on Acacia melanoxylon^ from Cheltenham, showing striated mark-
ings on the surface. These are considerably smaller than the average.

Plate XX HI

G. H Robinson, Phot.

x 250 & 600.

UROMYCLADIUM.

LEGUMINOSAE-ACACIA.

288

Explanation of Plates

PLATE XXIV.

(All Figures X ~jo unless otherwise stated.)

UPOMYCLADIUM.

Fig.

207. Teleutospores of U. bisforum^ on Acacia dealbata, one cluster of t

the common staik.

o showing

208. Teleutospores of the same.

209. Uredospores of U. alfinum, on Acacia dealbata.

210, an. Teleutospores of the same.

212. Five uredospores of U. alpinum on Acacia dallachiana, mixed with teleutospores

and one mesospore (m).

13, 214. Successive stages in the development of teleutospore clusters of the same

(Stained.)

215. Group of teleutospores of the same.

216. Section of leaf of Acacia longifolia attacked by U. maritimum, the uredosori

being at the margin of the inflated tubercle, and the spermogonia in the

centre

• * •

• •

# • •

• • ■

X 30

217. Uredospore of U. maritimum attacked by some Hyphomycete, which has gained

access to the interior of the spore and grown within it preparatory to forming
the long septate threads which have pierced the wall near the germ pores.

Plate XXIV

(J. II. Robinson, Phot.

x 30 & L'6U

UROMYCLADIUM.

LEGUMINOSAE-ACACIA.

290

Explanation of Plates.

PLATE XXV.

{All Figures X 250 unless otherwise stated.)

UROMYCES PHYLLODIORUM.

Fig

218. Two uredospores of U. fhyllodiorum (B. aud Br.) McAlp., on Acacia sp. from

Queensland. (Type material of Melamfsora fhyllodiorum B. and Br.,
from Herbarium of F. M. Bailey, Government Botanist, Queensland.)

219. One of the very sparse digitate teleutospores present in the same material.

220. Uredospore from the same material

• • •

X 500

(All the above from old and much faded material.)

1, 2

22. Uredospores of U. fhyllodiorum, on Acacia sp. from Queensland. (Type

material of Uromyces fhyllodiae Ccoke rnd Mass., these uredospores
described as teleutospores. From Herbarium of F. M. Bailey.)

being

223, 224. Two of the few digitate teleutospores present in the same material

(Material of above old and much faded.)

225. Uredospores of U . fhyllodiorum on Acacia dallachiana, from Bright, Victoria.

226. Two of the same, more highly magnified, to show surface markings ...

X 500

227, 228. Teleutospores from the same material, some beginning to germinate

(Stained.)

Note. — The markings of the uredospores are arranged in distinct lines, and are not

net-like as in Uromycladium notabile.

Pl.ATK XXV

-
*

G. H. Robinson, Phot.

X 250 & 500

UROMYCES PHYLLODIORUM

LEGUMINOSAE-ACACIA.

292

Explanation of Plates

PLATE XXVI.

(All Figures X 250 unless otherwise stated.)

PHRAGMIDIUM, MELAMPSORA
Fig.

229. Aecidiospores of Phr. subcorticium on Rosa rubiqnosa.

230. Uredospores of same.

231, 232. Teleutospores of same.

233. Twig of Rosa rubiginosa, the uppermost shoot of which is swollen and dis-

torted by the aecidia of Phr. subcorticium.

nat. size

■

234, 235. Teleutospores and uredospore of P. barnardi on Rub us farvifolius, two

germ pores being often seen on one face of each cell.

236. Teleutospores, closely attached to each other, of Melamfsora lini on Linum

usitatissimum.

1'I.ATK XXV]

G H. Robinson, Phot.

Nat. size & X 250

PHRAGMIDIUM AND MELAMPSORA.

ROSACEAE AND LINACEAE.

■

294

Explanation of Plates

PLATE XXVII.

AECIDIUM.

Fior.

237. Young

seedling of Platylobium formosum, with Aecidium flatylobii on leaves

and leaf stalks

nat. size

2i:8. Fruiting branch of the same, with aecidial cups on the pods

nat. size

239. Section through cups of Aecidium eburneum on pod of Bossiaea hetero-

X 5°

fhylla

• • •

• • 1

240. Section of aecidial cup of Puccinia erechtitis on Erechtites quadridentata X 50

t'LA'VK XXV [I.

G. H. Robinson, Phot

Nat, size & x 50

AECIDIA.

LEGUMINOSAE AND COMPOSITAE

•

296

Explanation of Plates

•

PLATE XXVIII.

{All Figures X 250).

Fig.

UREDO.

241. Spore of Uredo scirfi-nodosi or Scirfus nodosus

242. Uredospores of Puccinia stylidii on Stylidium graminijolium.

243. U. bidentis on Bidens filosa from Queensland.

244. U. bidentis on Bidens filcsa from Brazil. (Sydow, Uredineen, 1647.)

245. U . sfyridii on Sfyridium farvifolium

246. U. bossiaeae on Bossiaea frostrata.

247. U. fallidula on Cassia sp.

24S. Uredospores of Uromyces scleranthi on Scleranthus diander.

249. Uredospores of Puccinia oleariae on Olearia argofhylla^ epispore finely striate.

Plate XXVI II

G. H. Robinson, Phot

X 250

UREDO

•

* t

■

-»

■m

■

298

Explanation of Plates

Fig.

PLATE XXIX.

{All Figures X 250.)

ADDENDA to PUCCINIA.

250. Uredospores and leleutospores of Puccinia menthae on Mentha fulegium

251. Uredospores of P. chrysanthemi on Chrysanthemum indicum^ from New South

Wales.

252. Isolated teleutospore found associated with the same.

253. Uredospores and teleutospores of P. chrysanthemi on Chrysanthemum cult.,

Japan. (Herbarium A. Ideta.)

254. Abnormal teleutospore associated with uredospores and teleutospores of same.

255. Mesospore from same.

256. Group of teleutospores and mesospores of P. -podolefidis on Podolefis longu

fedata.

257. Teleutospore from same, with hyaline germ pore at each side of apex of upper

cell.

258. Teleutospores of P. oferculariae on Ofercularia varia.

Platk XXIX

G. H. Robinson, Phot.

X 250.

PUCCINIA.

LABIATAE, COMPOSITAE AND RUBIACEAE.

3oo

Explanation of Plates

PLATE XXX.

(All Figures X 250.)

ADDENDA to PUCCINIA and UREDO

Fig.

259. Uredospores of Puccinia cacao on Rollboellia comfressa

260. Uredospores and one teleutospore of same

261. Uredospores of P. lolii avenae on Avena saliva, germinating in water after

24 hours. (Stained.)

262. Uredospores of P. subnitens on Distichlis maritima

263, 264. Teleutospores from same.

sertcea.

266. Teleutospores of P. crucifrrae on an undetermined Crucifer. From type

material referred to Aecidium barbareac DC, at Kew.

267. Uredo rhagodiae on Rhagodia billardieri.

Platk XXX

I *

G. H. Robinson, Phot.

x 2:>0,

PUCCINIA AND UREDO.

• «

* *

-A

302 Explanation of Plates.

PLATE XXXI.

[All Figures X 250 unless otherwise stated.)

ADDENDA to PUCCINIA and PHRAGMIDIUM.
Fig.

26S. Leaf of Loranthus celastroides^ showing uredosori and teleutosori of Puccinia

loranthicola

• • • • • • •

... nat. size

69. Teleutospores from same.

270. Uredospores from s^me.

271. Leaf of Loranthus celastroides, with aecidia of same ... ... nat. size

272. Teleutospores of Phragmidium longissimum.

a. Four and five celled spores, before germination.

b. Spores commencing to germinate, with one germ tube divided by a

septum.

c. Spore cell, giving rise to four-celled promycelium, with loose sporidiola

around it.

The material was fourteen years old, and although there were plenty of sporidiola,
none were found attached to the promycelia.

Note. — Figs. 272, a, b, c, from drawings.

Plate XXXI

G. H. Robinson, Phot

x 2f>0 & 5<m>.

PUCCINIA AND PHRAGMIDIUM.

LORANTHACEAE AND ROSACEAE.

> .

•

■

3°4

Explanation of Plates.

PLATE XXXII.

UROMYCLADIUM

Fig.

273. Leaves of Acacia longifolia, with the tuberculate spermogonial sori of Uromy-

cladium maritimum

• • •

nat. size

274. Leaves of Acacia melanoxylon^ the upper with tuberculate spermogonial sori, and

the lower with minute pulverulent sori, both of U. robinsoni. ...

nat. size

275. Leaf of Acacia fycnantha, with the minute powdery sori of U . simflex. nat. size

Plate XXXII

O. H. Kobinson, Phot.

UROMYCLADIUM.

Nat. size

U. MARITIMUM, U. ROBINSONI, AND U. SIMPLEX ON ACACIA.

.. »

*■ I

• (-

-< - -

4 »

*o6

Explanation of Plates.

PLATE XXXIII.

Fi(T

UROMYCLADIUM BISPORUM

276. Sori of U. bisforum on stems and leaf stalks of Acacia dealbata

nat. size.

277. Sori on fruits

• • •

• •

nat. size

Plate XXXIII

G. H. Robinson, Phot.

Nat. size

UROMYCLADIUM.

LEGUMINOSAE-ACACIA DEALBATA

3°8 Explanation of Plates

PLATE XXXIV.

UROMYCLADIUM TEPPERIANUM.
Fig.

278. Branch of Acacia armata showing the powdery galls caused by the growth of

U. tefferianum

• • •

nat. size

Plate XXXIV

■

■ ■

. ■ v» ■ ■ ■

■
■ ■

■

■ ■

■

■ -

•

■

. ■

■

* 4 I*

G. H. Robinson, Phot

Nat. size.

UROMYCLADIUM.

LEGUMINOSAE-ACACIA ARMATA

"10

Explanation of Plates

PLATE XXXV.

Fig.

UROMYCLADIUM TEPPERIANUM.

279. Galls attached to branches of Acacia fycnantha, caused

ferianum

• • •

• •

U. tef.

nat. size

280. Galls detached

• • •

• •

nat. size

Pi-ATK XX \\

G. H. Rot inson, Phot.

Nat, size

UROMYCLADIUM.

U. TEPPERIANUM ON ACACIA PYCNANTHA.

•

■

♦•..

■

» ■

•

•^

438

312

Explanation of Plates

PLATE XXXVI.

UROMYCLADIUM NOTABILE.
Fig.

281. Numerous young galls on Acacia dealbata, bearing the uredospores of U.

« « • • • • ••• • • • ••• llclL.ol L* \Z

notabile

• • •

(Being on the young growth of the current season, the galls are necessarily
comparatively small, though many of several years5 growth bearing teleuto-
spores have been found as large as potatoes, and weighing nearly 1 lb.
each.)

Plate XXXVI

G. H. Robinson, Phot.

Nat. size

UROMYCLADIUM.

U. NOTABILE ON ACACIA DEALBATA.

•

■

• J

• ■

•

■

• *

•

L 2

314

Explanation of Plates.

^^s

PLATE XXXVII.

Fig.

CRONARTIUM JACKSONIAE.

282. C. jacksoniae deforming shoot of Platylobium formosum

• • «

nat. size

283. Witches' brooms due to C. jacksoniae on branches of Aotus villosa .

nat, size

284. Normal healthy shoot of Aotus villosa

nat. size

Pf'ATK XXXVII.

■

G. H. Robinson, Phot.

Nat. size.

CRONARTIUM.

C. JACKSONIAE ON PLATYLOBIUM AND AOTUS

■

•

316

Explanation of Plates.

PLATE XXXVIII.

CRONARTIUM JACKSONIAE.

Fig.

285. Witches' broom, due to C. jacksoniae on Gomfholobiitm lati folium

nat. size

2S6. Normal healthy shoot of Gomfholobium latifolium

nat. size

G. H. Robinson, Phot.

Nat. size

CRONARTIUM.

C. JACKSONIAE ON GOMPHOLOBIUM LATIFOLIUM

1 -

• k

* - *

■

•

■

•

v -

* •

• .

* •

•

3'8

Explanation of Plates.

PLATE XXXIX.

(All aecidia X 50, and aecidiosfores X 300.)

AECIDIUM

Fig.

287. A. veronicae on Veronica sp

288. A. flantaginis-variae on Plantago varia.

289. A. lobeliae Thuem., on Lobelia fratioides, described in connexion with Puccinia

aucta. The free margin ruptures irregularly, and it is sometimes difficult to
detect a peridial wall, so that it partakes of the nature of a Caeoma.

290. A. cymbonoti on Cymbonotus lawsonianus.

291. A. monocystis on Abrotanella forsterioides.

292. A. vittadiniae on Vittadinia australis.

293. A. soleniiforme en Goodia loti folia.

294. A. deeringiae on Deeringia celosioides.

295. A. calthae on Caltha introloba.

206. A. ranunculacearum on Ranunculus rivularis.

297. Aecidium of Uromyces folitus on leaves of Muehlenbeckia cunninghami — a

nat. size, b. aecidial tubes X 5.

298. Aecidiospores of same(X 300) and pseudoperidial cells (X 150).

Platk XXXIX

C. C. Brittlebank, Del.

X 5, 50, 15U & 30U

AECIDIUM

I • . —

•

■

320

Explanation of Plates

PLATE XL.

(All Figures X 500.)

NORMAL and ABNORMAL SPORES of PUCCINIA and UROMYCES.

PUCCINIA DICHONDRAE.

Fig.

209. The teleutospores exhibit a great variety of shape and size, and range from one

to four celled forms. In the two-celled forms the septum may either be
transverse or longitudinal, and in the three-celled forms the septa may
either be transverse or oblique, and the upper or lower cell may be divided
longitudinally. The four-celled forms may, in addition, become somewhat
spherical, and be divided longitudinally and obliquely, so as to resemble
a Sfhaero-phragmium. The variation even in the same sorus is sometimes
greater than it is between some different species.

PUCCINIA LUDWIGIL

300. Teleutospores may be divided generally as in the above, although the four-
celled forms are not quite so frequent.

PUCCINIA GRAMINIS.

301. Teleutospores are also one to four celled, but the septa are generally transverse,

while the upper cell may occasionally be longitudinally divided.

PUCCINIA TRITICINA.

302. The teleutospores do not show such a wide variation as in P. graminis, but they

may be from one to three celled, and the upper cell may be longitudinally
divided.

UROMYCES ORCHIDEARUM.

303. This species shows very well the transition from the unicellular [Uromyces) to

the bicellular {Puccinia) teleutospores. Some of the two-celled spores are
equally divided by the transverse septum and constricted, so that they depart
entirely from the Uromyces type.

Plate XL

G. C. Brittlebank. Del.

X 6UU.

PUCCINIA AND UROMYCES

NORMAL AND ABNORMAL TELEUTOSPORES.

' ■

% *

">22

Explanation of Plates.

PLATE XLI.

UROMYCLADIUM TEPPERIANUM

Fig
3°4-

Large gall on Acacia imflcxa, weight 3

lbs.

• • •

X *

G IJ. Robinson, Phot.

X J

UROMYCLADIUM.

LEGUMINOSAE-ACACIA IMPLEXA.

324

Explanation of Plates.

PLATE XLII.

UROMYCLADIUM.

Fig.

305. Witches' broom on Acacia im-plexa, due to Uromycladium tefferianum

X i

UROMYCES.

306. Teleutospores of Uromyces affendiculatus on Vigna catjang or Cow-pea, from

Richmond, New South Wales
307. Teleutospores of Uromyces fabat

# •

x 25°

on Lathyrus venosus. (Sydow, Uredineen,

1353)

• •

• * •

X 210

Pl.XTK XIJI.

G. 11. Robinson, Phot.

x { &, 2*>u

UROMYCLADIUM, UROMYCES.

LEGUMINOSAE.

•

\
V

• . ~

•if o ■ ■• ;

•

326

Explanation of Plates.

PLATE XLIIL

(All Figures X 250.)

PUCCINIA.

Fig.

308. Teleutospores of P. vittadiniae, mostly of a short broad type, on Vittadinia

austral is.

309. Teleutospores and mesospores of the same.

310. Teleutospores and mesospores of P. calotidis on Calotis sp

311. Uredospore of P.

branching growth at

graminis

the

red.)

on Avena Jatua, germinating in water, showing
end of twenfy-four hours. ^Stained with Congo

312. Mesospore of P. calendulae on Calendula officinalis, germinating in water after

three days, two sporidiola produced, both of which have already germinated.
(Stained with Bismarck brown.)

UROMYCLADIUM.

313. Teleutospore of U . maritimum on Acacia longifolia, germinating after twenty-

four hours in water, and producing four sporidiola from the lengthy pro-
mycelium. (Stained with Congo red.) (In some cases the promycelium
may be quite three times the length of that showrn when germination takes
place in water.)

314. Sporidiola of the same germinating in water three hours after formation.

(Stained with Congo red.)

315. Uredospores of the same germinating after three days in water. The germ

tubes are very broad, with almost colourless contents, and extremely thin
wall. At first they are generally much contorted, and sometimes bifurcate
at the apex. (Stained with Congo red.)

UROMYCES.

316. Aecidiospore of U. betae on Beta vulgaris, germinating in water after twenty-
four hours, the contents being collected towards the end of the tube where
branching is commencing. (Stained with Congo red.)

317. Teleutospores of U. folitus, one

ninghami.

being two-celled,

on Muehlenbeckia cun-

318. Uredospores with two bands of germ pores, and teleutospores of U. bicinctus

on Acacia fasciculifera.

319. Uredospores and teleutospores of U. folycnemi on Polycnemum fentandrum

Pl-ATK XMIT

G. II. Robinson, Phot.

x 250

PUCCINIA, UROMYCLADIUM, AND UROMYCES.

• **ir. '

•

i#*

.■

3^8

Explanation of Plates

PLATE XLIV.

RUST-RESISTING and RUST-LIABLE WHEATS.
Fig.

320. Rerraf, a variety of wheat generally found to be rust-resisting, and the straw

is shown to be perfectly clean.

321. Queen's Jubilee, a rust-liable variety, grown alongside of the other, and badly

attacked by Puccinia graminis.

IVvn WAY.

3-colour process

I . « . .*

RERRAF.

Straw perfectly clean

R. S. Hkain, (iovt. l'r

QUEEN'S JUBILEE.

Straw ba«ll\ ni-' • 1.

•

■

r*--

■

J *

■

- *

■

•

.
*

■J

•

Host Index

329

HOST INDEX.

Abrotanella forsterioides Hook. f.

Aecidium monocystis, Berk.

Abutilon avicennae Gaertn.

Puccinia heterospora, Berk, and Curt.

Abutilon crispum Sweet.

Puccinia heterospora, Berk, and Curt.

Acacia L.

Uromyces fusisporus, Cke. and Mass.
Uromyces phyllodiorum (B. and Br.)
McAlp.

Acacia armata R.Br.

Uromycladium tepperianum (Sacc.)
McAlp.

Acacia buxifolia A. Cunn.

Uromycladium alpinum, McAlp.

Acacia binervata DC.

Uromycladium notabile (Ludw.)
McAlp.

Acacia dallachiana F.v.M.

Uromyces phyllodiorum (Berk, and
Br.) McAlp.

Uromycladium alpinum, McAlp.

Acacia dealbata Link.

Uromyces phyllodiorum (B. and Br.)

McAlp.
Uromycladium alpinum, McAlp.
U. bisporum, McAlp.
U. notabile (Ludw.) McAlp.

Acacia decurrens Willd.

Uromycladium notabile ( Ludw.)
McAlp.

Acacia diffusa Lindl.

Uromycladium tepperianum (Sacc.)
McAlp.

Acacia elata A. Cunn.

Uromycladium notabile ( Ludw.)
McAlp.

Acacia erioclada Benth.

Uromycladium tepperianum (Sacc. )
McAlp.

Acacia fasciculifera F. v. M.

Uromyces bicinctus, McAlp.

Acacia glaucoptera Benth.

Uromycladium tepperianum (Sacc.)
McAlp.

Acacia hakeoides A. Cunn.

Uromycladium tepperianum (Sacc.)
McAlp.

Acacia implexa Benth.

Uromycladium tepperianum (Sacc

McAlp.

)

McAlp

Acacia juniperina Willd.

Uromycladium tepperianum (Sa
McAlp.

Acacia linifolia Willd.

)

Acacia longifolia Willd

McAlp

Alp

Acacia melanoxylon R.Br.

Uromycladium robinsoni, McAlp.

Uromycladium tepperianum (Sacc.)
McAlp.

Acacia microbotrya Benth.

Uromyces phyllodiorum (B. and Br.)
McAlp.

Acacia myrtifolia Willd.

Uromycladium tepperianum (Sacc.)
McAlp.

Acacia neriifolia A. Cunn.

Uromyces fusisporus, Cke. and Mass.
U. phyllodiorum (B. and Br.) McAlp.

Acacia notabilis F.v.M.

Uromyces phyllodiorum (B. and Br.)

( Ludw. )

McAlp.
Uromycladium notabile
McAlp.

Acacia penninervis Sieber.

Uromyces phyllodiorum (B. and Br. )
McAlp.

Acacia pruinosa A. Cunn.

Uromyces phyllodiorum (B. and Br.)

McAlp.
Uromycladium notabile (Ludw.)

McAlp.

Acacia pycnantha Benth.

Uromycladium simplex, McAlp.
U. tepperianum (Sacc.) McAlp.

Acacia retinodes Schlect. = Acacia

neriifolia, A. Cunn.

Acacia rigens A. Cunn.

Uromycladium tepperianum (Sacc.)
McAlp.

Acacia salicina Lindl.

Uromyces fusisporus, Cke. and Mass.
Uromycladium tepperianum (Sacc)

McAlp.

33°

Host Index.

Acacia siculiformis A. Cunn.

Uromycladium tepperianum (Sacc. )
McAlp.

Acacia spinescens Benth.

Uromycladium tepperianum i Sacc. )
McAlp.

Acacia stricta Willd.

Uromycladium tepperianum (Sacc. )
McAlp.

Acacia verniciflua A. Cunn.

Uromycladium tepperianum (Sacc.)
McAlp.

Acacia verticillata Willd.

Uromycladium tepperianum (Sacc. )
McAlp.

Acacia vomeriformis A. Cunn.

Uromycladium tepperianum ( Sacc. )
McAlp.

Acaena ovina A. Cunn.

Phragmidium
Karst.

potentillae (Pers. )

Acaena sanguisorbae Vahl.

Phragmidium potentillae (Pers.)
Karst.

Agropyron divergens Nees.

• Puccinia graminis, Pers.

Agropyron scabrum Beauv.

Puccinia agropyri, Ell. and Ev.
P. graminis, Pers.

Agrostis solandri F.v.M.=Deyeuxia

forsteri, Kunth.

Alopecurus geniculatus L.

Puccinia graminis, Pers.
P. perplexans, Plow.

Althaea rosea Cav.

Puccinia malvacearum, Mont.

Alyxia buxifolia R. Br.

Puccinia alyxiae, Cke. and Mass.

Amphibromus neesii Steud.

Puccinia graminis, Pers.

Anthoxanthum odoratum L.

Puccinia anthoxanthi, Fckl.

Aotus villosa Sm.

Cronartium jacksoniae, P. Henn.

Apium graveolens L.

Puccinia tbuemeni, McAlp.

Apium prostratum Labill.

Puccinia thuemeni, McAlp.

Asperula oligantha F.v.M

scoparia, Hook. f. )

Uromyces asperulae, McAlp.
Puccinia oliganthae, McAlp.

Atriplex semibaccata R. Br.

Uromyces atriplicis, McAlp.

Avena fatua L.

Puccinia graminis, Pers.
P. lolii avenae.

Avena sativa L,

Puccinia graminis, Pers.
P. lolii avenae.

Beckmannia erucaeformis Host.

Puccinia beckmanniae, McAlp.
P. graminis, Pers.

Bellis perennis L.

Puccinia distincta, McAlp.

Beta vulgaris L.

Uromyces betae (Pers.) Kuehn.

[A.

Bidens

Henn

Boronia spinescens Benth.

Puccinia boroniae, P. Henn.

Bossiaea cinerea R. Br.

Aecidium eburneum, McAlp.
Cronartium jacksoniae, P. Henn.

Bossiaea heterophylla Vent.

Aecidium eburneum, McAlp.

Bossiaea linophylla K. Br.

Aecidium eburneum, McAlp.

Bossiaea microphylla Sm.

Aecidium eburneum, McAlp.

Bossiaea prostrata R. Br.

Uredo bossiaeae, McAlp.

Bossiaea rhombifolia Sieber.

Aecidium eburneum, McAlp.

Brachycome ciliaris Less.

Puccinia brachycomes, McAlp. \

Brachycome diversifolia Fisch. and

Mey.
Puccinia brachycomes, McAlp.

Brachycome pachyptera Turcz.

Puccinia brachycomes, McAlp.

Brachycome scapiformis DC.

Puccinia brachycomes, McAlp.

Briza minor L.

Puccinia graminis, Pers.

Bromus arenarius Labill.

Puccinia bromina, Eriks.

Host Index.

Bromus mollis L.

Puccinia bromina, Eriks.

Bromus racemosus L.

Puecinia graminis, Pers.

Bromus secalinus L.

Puccinia graminis, Pers.

Bromus sterilis L.

Puccinia graminis, Pers.

Brunonia australis Sm.

Puccinia brunoniae, McAlp.

Bulbine bulbosa Haw.

Uromyces bulbinis, Thuem.

Burchardia umbellata R. Br.

Puccinia burchardiae, Sacc.

Calendula officinalis L.

Puccinia calendulae, McAlp.

Calocephalus drummondii Benth.

Puccinia calocephali, McAlp.

Calocephalus lacteus Less.

Puccinia calocephali, McAlp.

Calotis R. Br.

Puccinia calotidis, McAlp

Calotis cuneifolia P. Br.

Puccinia calotidis, McAlp.

Caltha introloba F.v.M.

Aecidium calthae, Grev.

Candollea serrulata Labill. = Sty-

lidium graminifoliura, Sw.

Canthium (Plectronia) coprosmoides

F.v.M.
Aecidium plectroniae, Cooke.

Carex L.

Puccinia longispora, McAlp.

Carex alsophila F.v.M.

Puccinia caricis (Schum.) Reb.

Carex breviculmis, R. Br.

Puccinia caricis (Schum.) Reb.

Carex caespitosa L.

Puccinia longispora, McAlp.

Carex gunniana Boott.

Puccinia caricis (Schum.) Reb.

Carex in versa R.Br.

Puccinia caricis (Schum.) Reb.

Carex paniculata L.

Puccinia caricis (Schum.) Reb.

Carex pedunculata Muhl.

Puccinia caricis (Schum.) Reb.

Carex vulgaris Fr.

Puccinia longispora, McAlp.

Carissa ovata R.Br.

Mass

Cassia Tourn.

Uredo pallidula, Cke. and Mass.

Centaurea cyanus L.

Pass

Chiloglottis diphylla R.Br

Uromyces orchidearum, Cke. and
Mass.

Chiloglottis gunnii Lindl.

Uromyces orchidearum, Cke. and
Mass.

Chrysanthemum indicum L.

Puccinia chrysanthemi, Roze.

Cichorium intybus L.

Puccinia cichorii (DC. ) Bell.

Cineraria L.

Puccinia cinerariae, McAlp.

Clematis aristata R.Br.

Caeoma clematidis, Thuem.
Puccinia agropyri, Ell. and Ev. I.
(Aecidium clematidis DC).

Clematis microphylla DC.

Caeoma clematidis, Thuem.

Coprosma billardieri Hook.

Puccinia coprosmae, Cke.

Coprosma hirtella Labill.

Puccinia coprosmae, Cke.

Correa lawrenciana Hook.

Puccinia correae, McAlp.

Crepis japonica Benth.

Uredo crepidis-japonicae, Lindr.

Cruciferae.

Puccinia cruciferae, McAlp.

Cryptandra hookeri F.v.M.

= Spyridium parvifolium, F.v.M.

Cymbonotus lawsonianus Gaudich.

Aecidium cymbonoti, Thuem.

Cynodon dactylon Pers.

Puccinia cynodontis, Desm.

Cyperus rotundus L.

Puccinia cyperi, Arth.

Dactylis glomerata L.

Puccinia graminis, Pers.

Dampiera alata Lindl.

Puccinia dampierae, Syd.

S32

Host Index.

Dampiera stricta R. Br.

Puccinia dampierae, Syd.

Danthonia DC.

Uromyces danthoniae, McAlp. I.

Danthonia semiannularis R. Br.

Uromyces danthoniae, McAlp. II., III.

Deeringia celosioides KBr.

Aecidium deeringiae, Cke. and Mass.

Deyeuxia forsteri Kunth. [Agrostis

isolandri F. v. M.)
Puccina agrostidis, Plow.

Deyeuxia quadriseta Benth.

Puccinia graminis, Pers.

Dianthus caryophyllus L.

Uromyces caryophyllinus (Schrank)
Schroet.

Dianthus chinensis L.

Uromyces caryophyllinus (Schrank)
Schroet.

Dichondra repens Forst.

Puccinia dichondrae, Mont.

Diploglottis cunninghamii Hook. f.

Uromyces diploglottidis, Cke. and
Mass.

Distichlis maritima Rafin.

Puccinia subnitens, Diet.

Echinopogon ovatus Beauv.

Puccinia graminis, Pers.

Ehrharta stipoides Labill. = Micro-

laena stipoides, R.Br.

Elymus condensatus Presl.

Puccinia impatientis (Schw.) Arth.

Elymus striatus Willd.

Puccinia graminis, Pers.

Elymus virginicus L.

Puccinia graminis, Pers.

Enchylaena tomentosa R.Br.

Puccinia kochiae, Mass.

Epilobium Dill.

Puccinia epilobii-tetragoni (DC)
Wint.

Epilobium billardierianum Ser.

Puccinia epilobii-tetragoni (DC.)
Wint.

Epilobium glabellum Forst.

Puccinia epilobii-tetragoni (DC)
Wint.

Erechtites Rafin.

Puccinia erechtitis, McAlp.

Erechtites arguta DC.

Puccinia erechtitis, McAlp.

Erechtites prenanthoides DC

Puccinia erechtitis, McAlp.

Erechtites quadridentata DC

Puccinia erechtitis, McAlp.

Eriostemon myoporoides DC

Puccinia eriostemonis, McAlp.

Festuca bromoides L.

Puccinia graminis, Pers.

Festuca ovina L.

Puccinia festucae, Plow.

Festuca rigida Kunth.

Puccinia festucae, Plow.

Geitonoplesium cymosum A. Cunn.

Uredo geitonoplesii, McAlp.

Geranium pilosum Sol.

Puccinia geranii-pilosi, McAlp.

Geum renif olium P. v. M.

Puccinia gei, McAlp.

Glyceria dives F. v. M.

Puccinia graminis, Pers.

Glyceria stricta Hook. f.

Puccinia graminis, Pers.

Gnaphalium japonicum Thunb.

Puccinia gnaphalii (Speg.) P. Henn.

Gnaphalium purpureum L.

Puccinia gnaphalii (Speg.) P. Henn.

Gompholobium latifolium Sm.

Cronartium jacksoniae, P. Henn.

Goodenia albiflora Schlecht.

Puccinia saccardoi, Ludw.

Goodenia geniculata R. Br.

Puccinia saccardoi, Ludw.

Goodenia glauca F. v. M.

Puccinia saccardoi, Ludw.

Goodenia hederacea Sm.

Puccinia saccardoi, Ludw.

Goodenia ovata Sm.

Puccinia saccardoi, Ludw.

Goodenia pinnatifida Schlecht.

Puccinia saccardoi, Ludw.

Goodia lotifolia Salisb.

Aecidium soleniiforme, Berk.

Haemodorum Sm.

Puccinia haemodori, P. Henn

)

Host Index.

333

■•

Hakea Schrad.

Uredo angiosperma, Thuem.

Hardenbergia monophylla Benth.

(Kennedy a monophyUa Vent. )
Uromyces hardenbergiae, McAlp.

Helianthus annuus L.

Puccina helianthi, Schw.

Helianthus tuberosus L.

Puccinia helianthi, Schw.

Helichrysum Vaill.

Puccinia kalchbrenneri, De Toni.

Hibbertia sericea Benth.

Puccinia hibbertiae, McAlp.

Hibiscus L.

Puccinia heterospora, B. and C.

Hordeum inurinum L.

Puccinia graminis, Pers.

Hordeum secalinum Schreb.

Puccinia graminis, Pers.

Hordeum vulgare L.

Puccinia graminis, Pers.

P. simplex (Koern.) Eriks. and Henn.

Hypericum japonicum Thunb.

Melampsora hypericorum (DC.)

Schroet.
Aecidium disseminatum, Berk.

Hypochoeris glabra L.

Puccinia hypochoeridis, Oud.

Hypochoeris radicata L.

Puccinia hypochoeridis, Oud.

Hypoxis glabella R. Br.

Puccinia hypoxidis, McAlp.

Jacksonia scoparia R. Br.

Cronartium jacksoniae, P. Henn.

Juncus effusus L.

Puccinia juncophila, Cke and Mass.

Juncus maritimus Lam.

Puccinia juncophila, Cke and Mass.

Juncus pallidus R. Br.

Puccinia juncophila, Cke and Mass.

Juncus pauciflora R. Br.

Puccinia juncophila, Cke and Mass.

Kennedya monophylla, Vent. = Har-
denbergia monophylla, Benth.

Kochia sedifolia F. v. M.

Puccinia kochiae, Mass.

Kochia villosa Lindl.

Puccinia kochiae, Mass.

Lactuca L.

Puccinia prenanthis (Pers.) Lindr.

Lagenophora billardieri Cass.

Puccinia lagenophorae, Cke.

Lagenophora huegelii Benth.

Puccinia lagenophorae, Cke.

Lavatera plebeia Sims.

Puccinia malvacearum, Mont.

Leschenaultia linarioides DC.

Puccinia gilgiana, P. Henn.

Limnanthemum indicum Thw.

Aecidium nymphoidis, DC.

Limosella aquatica L.

Uromyces limosellae, Ludw.

Linum marginale A. Cunn.

Melampsora lini (Pers.) Tul.

Linum usitatissimum L.

Melampsora lini (Pers.) Tul.

Lobelia anceps L.

Puccinia aucta, Berk, and F. v. M.

Lobelia pratioides Benth.

Puccinia aucta, Berk, and F. v. M.

purpurascens

Lolium perenne L.

Puccinia lolii, Niels.

Loranthus celastroides Sieber.

Puccinia loranthicola, McAlp.

Luzula campestris DC.

Puccinia tenuispora, McAlp.

Luzula oldfieldii Hook. f.

Puccinia tenuispora, McAlp.

Malva rotundifolia L.

Puccinia malvacearum, Mont.

Malva sylvestris L.

Pucciniainalvacearum, Mont.

Mentha laxiflora Benth.

Puccinia menthae, Pers.

Mentha pulegium L.

Puccinia menthae, Pers.

Microtis porrifolia R.Br.

Uromyces microtidis, Cke.

Micr olaena s tipoides R. Br. ( Ekrharta

atipoides Labill . )
Uromyces ehrhartae, McAlp.

Muehlenbeckia adpressa Meissn.

Puccinia muehlenbeckiae (Cke.) Syd.

Muehlenbeckia cunninghami F.v. M.

Uromyces politus (B. and Br.) McAlp.

334

Host Index.

Muehlenbeckia gracillima Meissn.

Puccinia muehlenbeckiae (Cke.) Syd

Olearia argophylla F. v. M. (Aster).

Puccinia oleariae, McAlp.

Olearia axillaris F.v.M.

Aeciclium oleariae, McAlp.

Opercularia aspera Gaertn.

Puccinia operculariae (Morr. ) Syd.

Opercularia varia Hook f .

Puccinia operculariae (Morr.) Syd.

Pelargonium australe Jacq.

Puccinia morrisoni, McAlp.

Phalaris canariensis L.

Puccinia graminis, Pers.

Phalaris minor Retz.

Puccinia graminis, Pers.

Phragmites communis Trin.

Puccinia magnusiana, Koern.
P. tepperi, Ludw.

Plagianthus sidoides Hook.

Puccinia plagianthi, McAlp.

Plagianthus spicatus Benth.

Puccinia malvacearum, Mont.

Plantago varia R. Br.

Aecidium plantaginis-variae, McAlp.

Platylobium formosum Sm.

Aecidium platylobii, McAlp.
Cronartium jacksoniae, P. Henn.

Plectronia coprosmoides = Canthium

coprosmoides, F. v. M.

Poa annua L.

Puccinia poarum, Niels.

Poa caespitosa Forst.

Puccinia poarum, Niels.

Poa pratensis L.

Puccinia poarum, Niels.

Podolepis longipedata A. Cunn.

Puccinia podolepidis, McAlp.

Polygonum aviculare L.

Uromyces polygoni, Fckl.

Pomaderris apetala Labill.

Uredo spy rid ii, Cke. and Mass.

Pratia erecta Gaudich.

Puccinia aucta, Berk, and F. v. M.

Pratia pedunculata Benth.

Puccinia aucta, Berk, and F. v. M.

Pratia platycalyx Benth.

Puccinia aucta, Berk, and F. v. M.

Prunus amygdalus Stokes.

Puccinia pruni, Pers.

Prunus armeniaca L.

Puccinia pruni, Pers.

Prunus domestica L.

Puccinia pruni, Pers.

Prunus persica Stokes.

Puccinia pruni, Pers.

Ranunculus L.

Aecidium ranunculacearum, DC.

Ranunculus gunnianus Hook.

Aecidium ranunculacearum, DC.

Ranunculus lappaceus Sm.

Aecidium ranunculacearum, DC.

Ranunculus parviflorus L.

Aecidium ranunculacearum, DC.

Ranunculus rivularis Banks and Sol

Aecidium ranunculacearum, DC.

Rhagodia billardieri R.Br.

Uredo rhagodiae, Cke. and Mass.

Rosa canina L.

Phragmidium subcorticium (Schrank)
Wint.

Rosa laxa Retz.

Phragmidium subcorticium (Schrank)
Wint.

Rosa rubiginosa L.

Phragmidium subcorticium (Schrank )
Wint.

Rottboellia compressa L.

Puccinia cacao, McAlp.

Rubus moluccanus L.

Phragmidium longissimum, Thuem .

Rubus parvifolius L.

Phragmidium barnardi, Plow, and
Wint.

Ruellia australis Cav.

Puccinia mussoni, McAlp.

Rumex brownii Campd.

Puccinia ludwigii, Tepp.

Host Index.

335

Rumex flexuosus Sol.

Puccinia ludwigii, Tepp.

Saccharum officinarum L.

Uredo kuehnii, Krueg.

Scaevola L.

Uromyces puccinioides, Berk, and
F. v. M.

Schelhammera undulata R. Br.

Uredo schelhammerae, McAlp.

Scirpus nodosus Rottb.

Uredo scirpi-nodosi, McAlp.

Scleranthus diander R. Br.

Uromyces scleranthi, Rostr.

Scorzonera angustifolia L.

Puccinia angustifoliae, McAlp.

Secale cereale L.

Puccinia graminis, Pers.

Selliera radicans Cav.

Uromyces puccinioides, Berk, and
F. v. M.

Senecio brachyglossus F. v. M.

Puccinia tasmanica, Diet.

Senecio pectinatus DC.

Puccinia tasmanica, Diet.

Senecio velleioides A. Cunn.

Puccinia tasmanica, Diet.

Senecio vulgaris L.

Puccinia tasmanica, Diet.

Sorghum halepense Pers.

Puccinia purpurea, Cke.

Sorghum vulgare Pers.

Puccinia purpurea, Cke.

Sporobolus indicus R. Br.

Uromyces tenuicutis, McAlp.

Spyridium parvifolium F. v.

(Cryptandra hookeri F. v. M.)
Uredo spyridii, Cke and Mass.

Stellaria media Cyrill.

Puccinia arenariae (Schum.) Schroet.

Stipa flavescens Labill.

Puccinia flavescentis, McAlp.

Stipa semibarbata R. Br.

Puccinia flavescentis, McAlp.

Stylidium graminifolium Sm. (Can-

dollea serrulata Labill.)
Puccinia stylidii, McAlp.

Tabernaemontana orientalis R. Br.

Caeoma apocyni, McAlp.

Tetragonia implexicoma Hook. f.

Puccinia tetragoniae, McAlp.

Thelymitra antennifera Hook. f.

Uromyces thelymitrae, McAlp.

Thelymitra flexuosa Endl.

Uromyces thelymitrae, McAlp.

Threlkeldia drupata Diels.

Puccinia dielsiana, P. Henn.

Tillaea sieberiana Schult.

Uredo tillaeae, McAlp.

Tremandra stelligera R. Br.

Puccinia pritzeliana, P. Henn.

Tricoryne elatior R. Br.

Uromyces tricorynes, McAlp.

Trifolium repens L.

Uromyces trifolii (Alb. and Schw.)
Winter.

Triticum polonicum L.

Puccinia graminis, Pers.
P. triticina, Eriks.

Triticum vulgare Vill.

Puccinia graminis, Pers.
P. triticina, Eriks.

Urtica dioica L.

Puccinia caricis (Schum.) Reb. I
(Aecidium urticae Schum.)

Velleia macrocalyx De Vriese.

Puccinia saccardoi, Ludw.

Velleia paradoxa R. Br.

Puccinia saccardoi, Ludw.

Veronica L.

Aecidium veronicae, Berk.

Veronica calycina R. Br.

Aecidium disc\forme, McAlp.

Veronica gracilis R. Br.

Aecidium disciforme, McAlp.

Vicia faba L.

Uromyces fabae (Pers.) De Bary.

Vigna catjang Walp.

Uromyces appendiculatus (Pers.)
Link.

Viola betonicifolia Sm.

Puccinia hederaceae, McAlp.

336

Host Index.

Viola hederacea LabilL

Puccinia hederaceae, McAlp.

Vittadinia australis Rich.

Puccinia vittadiniae, McAlp.

Wurmbea dioicaF.v.M.

Puccinia wurmbeae, Cke. and Mass
Uredo anguillariae, Cooke.

Xanthosia pusilla Bunge.

Puccinia xanthosiae, McAlp.

Zea mays L.

Puccinia maydis, Bereng.

Zornia diphylla Pers.

Puccinia zorniae (Diet.) McAlp.

Zygophyllum billardieri DC.

Uromyces vesiculosus, Wint.

Zygophyllum glaucescens F.v.M.

Uromyces vesiculosus, Wint.

Fungus Index.

337

FUNGUS INDEX.

Synonyms in italics ;

•::•

Species excluded from Australia ; t Doubtful species

Aecidium Pers.

f • •

193)

apocyni, Schwein. (See Caeoma apocyni, McAlp., p
*barbareae, DC. (SeePuccinia cruciferae, McAlp., p. 184)
*bellidis, Thuem. (See Puccinia distincta, McAlp., p. 156)

bossiaeae, P. Henn. = Aecidium eburneum, McAlp.

cakndulae, McAlp. = Puccinia calendulae, McAlp.

calthae, Grev. — Caltha introloba

dematidis, DC. = Puccinia agropyri, Ell. & Ev. I.

• •

• • •

• •

ccmpositarum, Mart. (See Puccinia tasmanica, Diet., p. 163)
cymbonoti, Thuem. — Cy mbonotus lawsonianus ...
cystoveiroides, Berk. = Puccinia operculariae (Morr.) Syd. I.

• • •

• •

(

leeringiae, Cke. and Mass. — Deeringia celosioides
disciforme, McAlp. — Veronica calycina, V. gracilis
disseminatum, Berk. — Hypericum japonicum
eburneum, McAlp. — Bossiaea cinerea, B. heterophylla, B. linophylla, B.

microphylla, B. rhombifolia

• •

goodeniacearum, Berk. = Uromyces puccinioides, Berk. & F.v.M. I., and

PAOK.

194

209

200

209

198
151

201

113
209

196
166

200
194
200

198

Puccinia saccardoi, Ludw. I.

• • •

impatientis, Schw. = Puccinia impatientis (Schw. ) Arth. I.
lobeliae> Thuem. = Puccinia aucta, Berk. & F.v.M. I.

microstomum, Berk. = Puccinia aucta, F.v.M. I.
monocystis, Berk. — Abrotanella forsterioides
nymphoidis, DC. — Limnanthemum indicum
oleariae, McAlp. — Olearia axillaris
perkinsiae, P. Henn. = P. gilgiana, P. Henn. I.

• •

- - .

• •

• • •

• •

• • •

• •

plantaginis, Ces. (See A. plantaginis- variae, McAlp., p. 195)

• • •

plantaginis- variae, McAlp. — Plantago varia
platylobii, McAlp. — Platylobium formosum
plectroniae, Cke. — Canthium (Plectronia) coprosmoides
ranunculacearum, DC. — Ranunculus sp., R. gunnianus,

• ■ ■

• •

- • ■

91, 147

. . 123

. . 148

.. 148

.. 197

,.. 196

... 197

.. 146

.. 209

.. 195

.. 199

.. 198

R. lappaceus, R.

parviflorus, R. rivularis

- • ■

• •

• • •

• •

senecionis, Desm. (See Puccinia tasmanica, Diet., p. 163)

• • •

• - •

soleniiforme, Berk. — Goodia lotifolia
urticae, DC. = Puccinia caricis (Schum.) Reb. I.
veronicae, Berk. — Veronica sp.
*violae, Schum. (See Puccinia hederaceae, McAlp., p. 183)
vittadiniae, McAlp. = Puccinia vittadiniae, McAlp.

• •

• ■ •

• •

Caeoma Link

• • •

• •

apocyni, McAlp. — Tabernaemontana orientalis
clematidis, Thuem. - Clematis aristata, C. microphylla

• • •

• * •

• • •

Cronartium Fr.

• •

♦ • •

• • •

p. 190)

asclepiadeum (Willd.) Fries (See C. jacksoniae, P. Henn.,
jacksoniae, P. Henn.— Jacksonia scoparia, Aotus villosa, Bossiaea cinerea,

Gompholobium latifolium, Platylobium formosum ...

£01
209
199
133
195
209
165

193

193
194

189
208

190

Hamaspora Koern.

longissima, Koern

Melampsora Cast

Phragmidium longissimum, Thuem

- - •

• •

• • •

hypericorum(DC.)Schroet.— Hypericum japonicum ... •••

lini (Pers.) Tul.-Linum marginale, L. usitatissimum. (Introduced.)
tnesodaphnes, Berk, and Br. Probably a Hyphomycete ... - .••

phylMwrum, Berk, and Br. = Uromyces phyllodiorum (B. and Br.)

Mc Alp.

187

191

191
192

208

• •

Fungus Index.

Phragmidium Link

• • I

• • •

• t

• t •

• •

- - •

• • •

barnardi, Plow, and Wint. — Rubus parvifolius

longissimum, Thuem. — Rubus moluccanus

potentillae (Pers.) Karst. — Acaena ovina, A. sanguisorba ...

subcorticium (Sclirank) Wint. — Rosa canina, R laxa, R. rubiginosa.

(Introduced).

• • »

• • •

PAGE.

185

186
187
188

188

Puccinia Pers.

• • •

• •

» • •

• • •

*acetosae (Schum.) Koern. (See P. ludwigii, Tepp. p. 174)
*aegra, Grove. (See P. hederaceae, McAlp., p. 183)

agropyri, Ell. and Ev. — Ayropyron scabrum II., III.; Clematis aristatal.

agrostidis, Plow. — Deyeuxia forsteri ...

altera, McAlp. = P. cynodontis, Desm. ...

alyxiae, Cke. and Mass. — Alyxia buxifolia

angustifoliae, McAlp. — Scorzonera augustifolia

anthoxanthi, Fckl. — Anthoxanthum odoratum. (Introduced.)
*apii, Desm. (See Puccinia thuemeni, McAlp., p. 168) ...

arenariae (Schum.) Schroet. — Stellaria media. (Introduced.)

aucta, Berk, and F.v.M. — Lobelia anceps, L. pratioides, L. purpurascens,

• • t

• • •

» • •

• • •

- - .

• •

Pratia erecta, P. pedunculata, P. platycalyx

• • •

beckmanniae, McAlp. — Beckmannia erucaeformis. (Introduce!.)

berkekyana, De Toni = P. dichondrae, Mont. ...

boroniae, P. Henn. — Boronia spines cens

brachycomes, McAlp. — Brachycome ciliaris, B. diversifolia, B. pachyptera,

B. scapiformis

• •

• • •

• *

• •

• • •

9 •

• • •

# • •

• •

• ■ ■

bromina, Eriks. — Bromus arenarius, B. mollis

brunoniae, McAlp. — Brunonia australis

burchardiae, Sacc. — Burchardia umbel lata

cacao, McAlp. — Rottboellia compressa

calendulae, McAlp. — Calendula officinalis

calocephali, McAlp. — Calocephalus drummondii, C. lacteus

colotidis, McAlp. — Calotis sp., C cuneifolia

caricis (Schum.) Reb. — Carex alsophila, C. breviculmis, C. gunniana, C.

inversa, C. paniculata, C. pedunculata, II., III. ; Urtica dioica I.
carissae, Cke. and Mass. — Carissa ovata
castagnei, Thuem. = P. thuemeni (Thuem.) McAlp. ... .

*caulincola, Cda. (See P. hypochoeridis, Oud., p. 159.)...
chrysanthemi, Roze — Chrysanthemum indicum. (Introduced.)
cichorii (DC.) Bell. — Cichorium intybus. (Introduced.)...
cinerariae, McAlp. — Cineraria sp. cult,
coprosmae, Cke. — Coprosma billardieri, C. hirtella
coprosmatisy Morr. = P. coprosmae, Cke.
coronifera, Kleb. =P. lolii, Niels. ... ...

correae, McAlp. — Correa lawrenciana »

cruciferae, Mc^lp. — Crucifer unknown

cyani (Schleich.) Pass. — Centaurea cyanus. (Introduced.)

cynodontis, Desm. — Cynodon dactylon

cyperi, Arth. — Cyperus rotundus

dampierae, Syd. — Dampiera alata, D. stricta

dichondrae, Mont. — Dichondra repens.

dielsiana. P. Henn. — Threlkeldia drupata, Diels.

dispersa Eriks. (See P. bromina, Kriks. and P. triticina, Eriks.)...

distincta, McAlp.— Bellis perennis

epilobii-tetragoni (DC.) Wint — Epilobium sp., E. billardieranum, E.

• • •

• •

• • •

• *

« • •

• • •

• •

1 • •

• • •

• -

• • •

• ■

• • t

112

207

208
113
114
118
143
150
115
208

177

148
116
142
181

150
116
144
138

117
151
151
152

133

... 144

... 168

... 208

... 153

... 154

... 155

... 165

... 123

... 181

... 184

... 156

... 118

... 134

... 146

... 142

... 176

116, 132

... 156

glabellum

• * ■

• • •

170

erechtitis, McAlp. — Erechtites sp., E. arguta, E. prenanthoides, E. quad-

ridentata

• - -

• •

• • •

• * •

• • •

- ■ -

• • •

• •

eriostemonis, McAlp. — Eriostemon myoporoides

festucae, Plow. — Festuca ovina, F. rigida. (Introduced.)

flavescentis, McAlp. — Stipa flavescens, S. semibarbata

gei, McAlp. — Geum renifolium

geranii, Corda — (See P. morrisoni, McAlp., p. 180)...

geranii-pilosi, McAlp. — Geranium pilosum

gilgiana, P. Henn. — Leschenaultia linarioides

gnaphalii, (Speg.) P. Henn. — Gnaphalium japonicum, G. purpureum

gnaphaliicola, P. Henn. = P. gnaphalii (Speg.) P. Henn.

. . .

• •

• • •

• •

157

182
119
119
170
208
179
146
158
158

Fungus Index.

339

PAGE.

Puccinia Pers. — continued.

graminis, Pers.— Agropyron divergens, A. scabrum, Alopecurus geniculates,

Amphibromus neesii, Avena fatua, A. sativa, Beokmannia erucae-
formis, Briza minor, Bromus racemosus, B. secalinus, B. sterilis, Dactylia
glomerata, Deyeuxia quadriseta, Echinopogon ovatus, Elymus striatus,
E. virginicus, Festuca bromoides, Glyceria dives, G. stricta, Hordeum
murinum, H. secalinum, H. vulgare, Phalaris canariensis, P. minor,
Secale cereale, Triticum polonicum, T. vulgare. (Introduced.) ... 120

haemodori, P. Henn. — Haemodorum sp. ... ... ... ... 139

McAlp. — Viola betonicifolia, V. he deracea ... ... ... 183

helianthi, Schwein. — Helianth

(Introduced.) ... 158

heterospora, Berk, and Curt. — Abutilon avicennae, A. crispum. Hibis

cus sp

• • • ••• ••• •••

McAlp. — Hibbertia sericea

• . .

• - •

McAlp. — Hypoxis glabella

• • • • • • ••• - • « • • -

loranthicola, McAlp. — Loranthus celastroides
ludwigii, Tepp. — Rumex brownii, R. flexuosus...

• • •

• • • . ■ •

• •

• • • • • • •

macalpini, Syd. =P. lagenophorae, Cke.
magnusiana, Koern. — Phragmites communis

Mont.— Althaea rosea, Lavatera plebeja, Malva rotundifolia,

Plagia nthus spicatus . ( Introduced. )

• ■

••• ••• •••

• • •

• • • • • •

mussoni, McAlp. — RuelJia australis

• * •

McAlp. = P. saccardoi, Ludw.
McAlp. — Olearia argophylla

• • • • •

• •

• • • • •

perplexans, Plow.— Alopecurus geniculatus

• * •

177

185

hypochoeridis, Oud. — Hypochoeris glabra, H. radicata. (Introduced.) ... 159
hypochoeridis, McAlp. — (P.macalpini, Syd.) = P. lagenophorae, Cke. ... 161

139

impatientis, (Schw.) Arth. — Elymus condensatus. (Introduced.) ... 123

*investita, Schw. — (See P.gnaphalii (Speg.) P. Henn. ... ... ... 158

juncophila, Cke. and Mass. — Juncus effusus, J. maritimus, J. pallidus ... 136

mm art. ... ... ... ... 160

kalchbrenneri, De Toni — Helichry

kochiae, Mass. — Enchylaena tomentosa, Kochia sedifolia, K. villosa ... 176

lagenophorae, Cke. — Lagenophora billardieri, L. huegelii ... ... 161

lolii, Niels. — Avena fatua, A. sativa, Lolium perenne. (Introduced.) 123

longispora, McAlp. — Carex sp., C. caespitosa, C. vulgaris ... ... 135

167
174

161
125

178
127

maydis, Bereng. — Zea mays. (Introduced.)

menthae, Pers.— Mentha laxiflora, M. pulegium. (Introduced.)... ... 140

microseris, McAlp. = P. hypochoeridis, Oud.

159
180

morrisoni, McAlp. — Pelargonium australe

muehlenbeckiae (Cke. ) Syd.— Muehlenbeckia adpressa, M. gracillima ... Uo

munita, Liidw. = P. dichondrae, Mont — ••• 142

141

147
205
165

McAlp. — Asperula oligantha ... ...

Morr.) Syd.— Opercularia aspera, O. varia . ... ... loo

127

phragmitis (Schum.) Koern.— (See P. magnusiana, Koern, p. 1^5) ... **>

plagianthi, McAlp.— Plagian thus sidoides ... # ... ••• ••• {£?

poarum, Niels.— Poa annua, P. caespitosa, P. pratensis. (Introduced.) ... j^»

podolepidis, McAlp.— Podolepis longipedata ... •• — *" i«o

prenanthis(Pers.)Lindr.— Lactuca sp. (Introduced.) J?f

pritzeliana, P. Henn.— Tremandra stelligera ... ... «• »•

pruni, Pers.— Prunus amygdalus, P. armeniaca, P. domestica, P. persica. ^

(Introduced.) ... ... ••• ••• ••• , ,*V '*' i4>q

purpurea, Cke. -Sorghum halepense, S. vulgare. (Introduced.) ... i-»

*rimosa, Link.— (See Uredo scirpi-nodosi, McAlp., p. 203) £°

*rumicis, Lasch.— (See P. ludwigii, Tepp., p. 174) ... - , - n5

rwmcis-sciUati var. muehlmbeckiae, Cke. =P. muehlenbeckiae (Cke ) Syd. ... uo

saccardoi, Ludw. - Goodenia albiflora, G. geniculata, G. g»«ca. u- -

hederacea, G. ovata,G. pinnatifida, Velleia macrocalyx, \ . pa"£ox» •» J*0

simplex (Koern.) Eriks. and Henn. -Hordeum vulgare. (/ntr^ucwU •» ™
sorqhi Schw. = P. maydis, Bereng., and P. purpurea, Cke. (Introduced.) 127, 1£

•••

subnitens, Diet.— Distich) is maritima ... ... ••• vpllpinidea'

tasmanica, Diet. - Senecio brachyglossus, S. pectmatus, S. n elleioides,

131

... 163

S. vulgaris ... ••• ••• ••• . ••' 137

McAlp.— Luzula campestns, L. oldneidn ... ••• •••

tepperi, Ludw.— Phragmites communis... ••• •• 173

tetragoniae, McAlp.— Tetragonia implexicoma "/r.^nAn^A \ '" 168

thuemeni, McAlp.-Apium graveolens A. prostratum. (Introduced.) ... ig

triticina, Eriks.— Triticum vulgare. (Introduced.) ...

340

Fungus Index.

Puccinia Pers. — continued.

violae, (Schum. ) DC.

PAGE,

McAlp. — Vittadinia australis
Ike. and Mass. — Wurmbea dio]
McAlp. — Xanthosia pusilla

McAlp., p. 183)

• • •

• •

• • •

)McA

Zornia diphylla

■ • -

• • 4

» • •

Uredo

• • •

t • *

» • •

* •

• • 4)

• • •

• t •

angiosperma, Thuem. — Hakea sp.
anguillariae, Cke. — Wurmbea dioica
armillata, Ludw.

bidentis, P. Henn. — Bidens pilosa *
bossiaeae, McAlp. — Bossiaea prostrata
*cichoracearum, DC.
clematidis, Berk. = Caeoma clematidis, Thuem.
crepidis-japonicae, Lindr. — Crepis japonica

• •

• • •

Mass

• • •

« • •

• - .

Henn

204)

• • •

• - •

• •

chrhartae, McAlp.

McAlp

gnaphalii Speg.

McAlp — Geitonoplesium cymosum

- - •

• • •

• t

- t

(Speg. )

• •

kuehnii, Krueg. — Saccharum officinarum. (Introduced.)

leguminum, Desm. — Acacia sp.

• • •

• •

notabilis, Ludw.

Cassia sp.

McAlp

• •

. - •

• • t

• .

• •

rhagodiae, Cke. and Mass. — Rhagodia billardieri
rottboelliae, Diet. = Puccinia cacao, McAlp.
schelhammerae, McAlp. — Schelhammera undulata
scirpi-nodosi, McAlp. — Scirpus nodosus
sorghi, Pckl. = Puccinia purpurea, Cooke,
spyridii, Cke. and Mass. .....__

tillaeae, McAlp. — Tillaea sieberiana

♦ • *

• • •

• ^

• • •

• •

• • •

.mm

• •

• • •

Uromyces Linl

• • •

• •

• • •

• •

• • •

• • »

• • •

amygdali, Pass. (See Puccinia pruni, Pers., p. 171)
appendiculatus, (Pers.) Link — Vigna catjang. (Introduced.)
asperulae, McAlp — Asperula oligantha

asteris, McAlp. = Uredo dubia, McAlp

atriplicis, McAlp. — A triplex semibaccatum

betae (Pers.) Kuehn. — Beta vulgaris. (Introduced.)

bicinctus, McAlp. — Acacia fasciculifera

bulbinis, Thuem. — Bulbine bulbosa

caryophyllinus (Schrank) Schroet. — Dianthus caryophyllus, D chinensis

• •

• • •

• •

* •

• •

(Introduced.)

• •

• * •

• •

danthoniae, tylcAlp. — Danthonia sp., D. semiannularis ...
digit at us, W int. = Uromyces phyllodiorum (B. and Br.) McAlp.
diploglottidis, Cke. and Mass. — Diploglottis cunninghamii
ehrhartae, McAlp — Microlaena (Ehrharta) stipoides

• •

• ■

• •

» • *

• • •

• •

* • •

• • «

• 9

fabae (Pers.) De Bary. — Vicia faba. (Introduced.)

fusisporus, Cke. a&d Mass. — Acacia neriifolia, A. salicina

hardenbergiae, McAlp — Hardenbergia [Kennedya) monophylla

junci (Desm.) Tul. — (See Puccinia juncophila, Cke. and Mass. p. 136.)

kuehnii, Krueg. =Uredo kuehnii, Krueg.

limosellae, Ludw. — Limosella aquatica...

microtidis, Cke — Microtis porrifolia

orchidearum, Cke. and Mass — Chiloglottis diphylla, C. gunnii

phaseoli (Pers. ) Wint. = Uromyces appeadiculatus (Pers.) Link. ...

phyllodiae, Cke. and Mass. = Uromyces phyllodiorum (B. and Br.) McAlp.

phyllodiorum (B. and Br-) McAlp. — Acacia sp., A. dallachiana, A. micro-

botrya, A. neriifolia, A notabilis, A. penninervis, A. pruinosa
politus (Berk) McAlp. — Muehlenbeckia cunninghami
polycnemi, McAlp. — Polycnemum pentandrum

• • •

• f

polygoni (Pers.) Fckl. — Polygonum aviculare. (Introduced.)
puccinioides, Berk, and P. v. M. — Selliera radicans, Scaevola sp.
scleranthi, Rostr. — Scleranthus diander ... ...

tenuicutis, McAlp. — Sporobolus indicus

tepperianus, Sacc. = Uromycladium tepperianum (Sacc. ) McAlp.

thelymitrae, McAlp — Thelymitra antennifera, T. flexuosa

• •

• • •

• •

• • •

• • «

tricorynes, McAlp. — Tricoryne elatior ...

trifolii (Alb. and Schw.) Wint. — Trifolium repens. (Introduced.)

vesiculosus, Wint. — Zygophyllum billardieri, Z. glaucescens

• • •

< • •

• •

208
164
138
169
172

202

206
203
136
204

206
209
194
205
86
203
158
202
209

108
206
207

117

203
202
129
204
206

207
92

91
205
100
100

102
85
95

101
86
93
94
94

207
202
90
88
89
92
95

210

91
102

111

103

Fungus Index.

34i

Uromycladium McAlp.

■ - • • - ' • •

• - • ••• ••

Page.

104

alpinum, McAlp. — Acacia buxifolia, A, dalluchiana, A. dealbata, A. implexa,

A. linifolia
bisporum, McAlp. — Acacia dealbata
maritimum, McAlp. — Acacia longifolia
notabile (Ludw.) McAlp. — Acacia binervaba, A. dealbata, A. decurrens, A.

9 w » ••• ••• ••• • • •

• • • • . • • • • • ■ . •••

• • • • • • • ■ • • • •

elata, A. notabilis, A. pruinosa

• •• ••• . . .

robinsoni, McAlp. — Acacia melanoxylon

simplex, McAlp. —Acacia pycnantha

tepperianum (Sacc.) McAlp. — Acacia armata A. diffusa, A. erioclada, A

glaucoptera, A. hakeoides, A. implexa, A. juniperina, A. longifolia, A.

melanoxylon, A. myrtifolia, A. pycnantha, A. rigens, A. salicina, A.

siculiformis, A. spinescens, A. siricta, A. verniciflua, A. verticiliata,

105
IOC

107

10s

109
110

A. vomer if or mis . . .

• •

■

•

■

General Index.

343

GENERAL INDEX

• •

• • i

Acacia, gall fungi
Kangaroo

Acacia armata

dealbata

decurrens
implexa

pycnantha
salicina

Aecidiospores, binucleate

• • •

a •

• •

• « •

• • ■

• •

• • I

- • »

• • •

• I •

coloured membrane in Gymno-

sporangium
colouring matter
development
distinction from uredospores
germination in Puccinia

I'AGK.

5
6
6
6
6
6

• • •

16
16

17
36

tasmanira
infection
origin
partaking of

teleutospores
repeated formation...

• -

• • •

retention

16
16

... 17, 36

character of

• •

germinating

power . . .
Aecid ium, colour

development
invigorating power

in heteroecious species
on grass ...

on seeds ...
origin

t • •

• • i

• * •

paraphy ses
peridiurn ...
primary and secondary
pseudoperidial cells
repeated formation
sterile cells
sexuality
A ecidium abietinum

* * ■ ■ ■

bellidis

berberidis, spore germination
importatum

platylobii, mycelium in seed

Algae

Alopecurus pratensis with Puc-

16
16
17

18,57

... 17, 75
22, 36

17
18

14, 17, 75

209

70
40

Anaph

cinia gramims

- • *

• • •

germination

• •

• • •

• • t

Anagallis arvensis, importation
Anchusa

• • • - - ■

Anthomyces

Apium, graveolens (Puccinia)

proHtratum (Puccinia)
Appressorium
Asclepia'leae, Cronartium
Ascomycete8 ...

Asparagus rust (Puccinia axpar-

agi), distributed by wind

68
25
25
43
53, 80
84
42
42

3
53
40

water relation

• *

• •

• • •

• •

Atmospheric influences on spore

germination
A uric xdariaceae
Australian rusts, number

distribution

433.

10
32
50

Autoecious species

most numerous

repetition of aecidia ...

A vena elatior with Puccinio

graminis

Baeodromus ...

• • • •• •

Barberry, connexion with wheat

PAOR.

), 55
45

68
33

rust

• • t

• •

ft • ft

• ft ft

in Australia

infection experiments, local

in Kew gardens
Barberry rust (A ecidium ber-
beridis) ...

absent in Australia

connexion with wheat rust ...

increasing vigor of wheat rust

local attempts at inoculation
Barclay e lla

Basidia in aecidia

Basidiomycetes

55, 58

66
66

• • •

— *

fabae)
formis, impor

• • •

• •

16
66
55
5S
66
33
16
40
20

• •

tion of rust on seed
Beet rust ( Uromyces betae)
Binucleate spores

Biologic forms, and classification

evolution of
Bismarck brown
Black wattle gall fungus (Uro-

mydadium notabile)
Br achy puccinia
Bremia lactuca% effect of copper

Sell Lo ♦ • • ••• . . ,

Bridging species

Brome rust (Puccinia bromina) ...
infection by uredospores
starvation of host ...
uninfluenced by structural
peculiarities of leaf

Brown Rust (Puccinia dispersa) ...
Burning stubble, effect on wheat

43
84
14

79
54
12

(i
11

63
53

rust

ft •

■ • ft

t • •

• € •

• • •

Caeoma

Caeomospores ...

Carnation rust (Uromyces caryo-

17,36
36

phyllinus)

Caustic potash for softening tissues

Cecidomyia, eating rust spores ...

Cedar apples ...

Celery rust (Puccinia thwmeni) ...

Cereal rusts not liable to affect

other cereals indiscrimin-
ately ... ••• •••

Chemotaxis, parasitism due to

positive and negative

Chick weed rust (Puccinia aren-

ariae)
Chicory rust (Puccinia cichorii) . .

Chrysanthemum rust (Puccinia

chrysanthemi)
germination of uredospores
importation

85
12

52,62

42
42

1
21
43

I I

344

'II

General Index.

Chrysomyxa, absence of fir trees
germination of teleutospore
origin of species in Alps

» • •

• i •

• •

uredospores in chains
Chrysomyxa, abietis

ICCuw • • « •• •

rhododendri

Chrysopsora, teleutospore of ex-
ceptional form ...

Classification and biologic forms

of Uredines

PAGE.

58
34
56
19
66
56
56

• • •

trifolii) ... 18, 84

Cluster cups ...
Coleospcriaceae

Coleosporium, function of paraphyses 30

teleutospore in
uredospores in chains

Coleosporium senexionis, and fir trees 58

Collema, fertilisation in ... 37

• • •

Compositae, native rusts on
Compound teleutospores
Copper salts, influence on fungi...
Cronartiaceae ...
Cronartium>asclepiadeum on varied

.82

hosts

• •

jacksoniae producing witches'
brooms ...

Crossbreeding wheats for rust-
resistance

Crown or coronate rusts

Cucumber diseases and copper salts

Cycle of development, in heteroe-

74
56
63

cious species
in rusts

• •

Cuperaceae, number of native rusts

, 58
2, 10

• • •

Dactylis glo merata

Daisy rust (Puccinia distincta) ...

introduced on seed ...
Danthonia aec
Darluca Mum

45
53
10

209

... 18, 57
parasite 22
mistaken for spermogonia 22, 158, 165

Development of fungi ...
Dew necessary for infection
Diorchidium ...

doubtful genus

• •

• • •

• •

* • •

Distribution of Australian species
Distribution of spores
Doubtful species

• • •

• # •

Drainage, effect on rust

Dry atmosphere and spore ger-
mination

Early maturing wheat escaping rust

Echinulate uredospores...

Effect of rust on straw and grain

Elymus condensatus, importation

of rust on seed ...

Endochrome ...

Endophyllum, teleutospores

Endospore, in aecidiospores
in teleutospores

Enzymes in leaf cells ...

Epispore of teleutospore

Epiteospores ....

Erysiphaceae, specialisation

83
82

207

10
74

19
64

# •

• • •

• • «

17
23
16
23
54
23
20

Eru

parasitism

• • •

54
54

• • 1

• • •

Euphorbia, aecidium of Pea rust

Gu&fvLCLLCL ••• ••• •••

rust ( Uromyces euphorbiae) . . .
Eupuccinia
Excluded species
Fallowing, effect on rust
Fertilisation, in aecidium

in Uredineae ...

Fir trees and rust
Flax rust (Melampsora lini)

first record in Australia
Formalin treatment of seed wheat
Frangula alnus

Fungi, origin and development . . .
Fusion of nuclei
Gall formation...

t>kQti.

55
70
70
11

207

... 17, 75
... 14, 75

% • •

... 43, 60

73
53
40

... 14, 75

• - *

Gall fungi, on wattles (Acacias) ..•
Genera, Australian, and number of

OlJL/V-/lt>0 »•• ••• •••

General parasites

Germinating power, duration, in

5
6

50
53

aecidiospores
in teleutospores
in uredospores

• • •

• •

• * •

8
8, 21

Germinating spores, methods for

aecidiospores and uredo-

bUOI vS ••• ••• •••

teleutospores
Germination, of aecidiospores of

• • •

Puccinia tasmanica
of amphispores
of teleutospores, factors in-

16
25

fluencing
of Puccinia graminis

of P. malvacearum

8, 24, 66
... 24, 27

of Phragmidium rubi abnor-

mal

• • ♦

of uredospores, of Puccinia

bromina

• • •

of P. chrysanthemi

of P. dispersa

of P. graminis ...

of P. rubigovera . . .

of P triticina ...
Germ-pores, demonstrating

in aecidiospores
in teleutospores

of Puccinia podolepidis

in uredospores

• • •

. • •

• •

• • #

• •

■ ■ ■

3, 8
21

3
21

8
21
12
16
24

35
19

solitary in Puccinia mono-

pora

• • •

Glycerine, and water ...

I Li i y ••• ••• • • •

Golden rust (Puccinia glumarum)

Golden wattle galls

Gradations of specific variation ...

Gramineae, native rusts on

Grass aecidium

Grasses, and heteroecious rusts ...

19
12
12
73
6
79
45

and wheat rust

• • •

• •• lo, O/.

57
69

Groundsel rust (Puccinia tas-

manica)

• •

Gymnoconia, characters...
Gymnosporangium, aecidiospores

colored membrane

cTclllS . . • ••• •••

insects conveying sporidiola
sculpturing of peri dial cells ...

... 16, 49

16
6

General Index.

345

PAGE.

I'AOI.

mosporangium clavariaeforme9

direct infection by teleuto-

spores ...
confusum, promycelial

separating

teleutospores, thickness of

cells

cell wall

• •

• • •

« •

Hap a

Hard tissues, softening...

Harvesting methods and rust

Haustoria

Hawthorn

Helianthus anmtus, aecidi

// em ibasidii ...
Hemileia
Hemipuccinia ...
Heteroecious species

indigenous

regular development
Heteroecism

• - »

• #

• • •

• - *

• • •

• - •

• • •

• ■ •

• • •

• •

. . .

Hit

Hollyl

beneficial ...

causing increased vigor

discovery ...

origin

when possible

• • •

• •

♦ • •

• * •

- - -

35
83
12
69

28
18
40
83
11
53
45

57
55

arum )

• • •

wide distribution

- - •

• •

• • •

• • «

• ■ •

Holobctsidii

Honeydew of spermogonia
Host and parasite, relatior
Host-plants, imported, and new

* •

43
40
13

rusts
indigenous, and rusts

starved

• • •

Hot- water treatment of seed
Hypertrophy of nettle

Hyphae
Immunity acquired

and predisposition ..

attempts to secure ...

due to physiological peculiari-

42
45
60

6
3

60
60

62, 75

ties

• •

factors influencing .•.
Importation of rusts ...

on cuttings

on seed ...

on straw ...
Incubation period
Indigenous species, table of

• • •

* • •

• • •

and hosts ...
heteroecious

• • •

• •

Infection, aided by injury

and age of parts attacked ...

by aecidiospores ...

by internal germ of disease ...

15y sporidiola

germ tubes entering stomata

by teleutospores
by uredospores
germ tubes piercing epider-

43,44

9, 60

mal cells

• • I

dew necessary for ...
experiments, with barberry in

10
10

Australia

• • •

with foreign host plants

factors influencing

• t •

66
9
9

Injuries, influence on infection
Inoculation, protective...

• ■ •

61
62

Insects attracted

colour of

spores

• • ■

• • •

♦ • •

by honeydew

by scent of spermogonia

conveying sporidiola

eating spores
Intermediate hosts of wheat
Internal developmental tendencies
Introduced species

rust

• • .

• • #

• • •

how introduced

on native hosts
Investigation of spores ...
Irrigation and wheat rust

Kangaroo thorn, gall fungus

K not- wet d rust ( Uromyces polygoni)

Lactic acid showing up germ pores
Ledum palustre
Leguminosae, native rusts on
Leptopuccinia ...

species with war ted
epipspore
Lettuce, attempt to confer immu-

• • •

38
13

7
70

42,49

6
43

56
45
11

one

lsease

Liability to disease

influenced or not by structural

characters
Lichen-fungi ...

Life-cycles of rusts
heteroecious

Life history of rusts

Liliaceae, native rusts on

Linum marginale

63
60

usitatissimum

• •

rust -resisting variety
Lolium perenne, crown rust
with introduced rust

Mahonia in Kew gardens, rus
Maize rust (Puccinia maydis)
Mallow rust (Puccinia malva*
Manures and wheat rust

M ■

• • •

• - •

37
2.10

68
42
43

Marigold rust {Puccinia calendulae) 10, 42
Measuring spores by photography 12
Melampsora, germination of teleu-

tospore ... ••• •••

Melampsora eucalypti, merely a

name

• • •

icorurn

• •

• • •

lini

introduced
paraphyses
M elampsoraceae^

Mesospo

• • •

■ - •

in Uromycladium

22, 49
44
22

25, 26
25

Mint, native, rust (Pucctma

menthae)
Mounting spores

Mycelium, development of
in Refill of annuals

. • -

• • •

investigation
localized, causi w
perennial causing galls

hypertro

• • •

no evidence in wheat grain
of Puccinia arrhenatheri .••

of Uromyces euphortriae

t ♦ t

42
12

3
5,70
3
6
5
70

34^

General Index.

Mycoplasm theory

• • •

• ft

• • •

• . •

• • •

bearing on rust in wheat
Native celery ...

flax

mint

rusts, and native hosts
on imported hosts
Nettle rust (Aecidium urticae)
Nuclear cycle ...
Nuclei, fusion of
Oat, wild, and wheat rust
Obligate parasites
Ochropsora

Odour of spermogonia .
Omission of spore-forms
Origin of heteroecism

of parasitism

of spore-forms
Paraphyses, function

in aecidia

in spermogonia

in teleutosori

in uredosori

- ■ .

• •

• • •

• •

• • #

• •

• *

• •

• ♦ '

• • •

PAOK.

4, 74

42, 44

14, 75

13
10

55
52

• •

22,30

13, 29

• • •

Australian species with

• • #

Parasitism, origin

specialisation
Pea rust ( Uromyces pisi)
Peach or prune rust {Puccinia

22, 24, 75

22
22

52
52
55

pruni )

• • •

• •

Pedicel of teleutospore ...
Peltandra virginica
Perennial mycelium
Periderm ium ...
Peridial cells ...

• • •

* • •

Peridium, =Pseudoperidium

often wanting

varied forms
Phalaris, Puccinia
Photography in measuring spores
Phragmidium bamardi, germ pores

paraphyses

longissimum

potentillae...

rubi9 teleutospore germination
subcortkium

ft • 4

paraphyses, in aecidia

in uredosori

• . •

perennial mycelium

... 19, 44

23
61

• • • o

17
17

55
12
24

22
44
44

5, 36

22
5

* •

• •

• * •

• • g

Phragmites communis, Puccinia ... 29, 42
Phragmopyxis ... ... 84

Phy corny cetes^ ... ... ... 40

Phytopthora infestans, and copper

ScHuS ••• mm*

Picea excelsa ...

morinda

Ploughing, influence on wheat rust

Plum or prune rust (Puccinia

pruni) ...

Poa annua

62
56
33
71

pratensis . . .
Podisoma

Polygonum aviculare
Potash, caustic,

tissues

• • •

22
22
28
43

for softening

r • •

Predisposition
Primordia

• • *

• • •

Promycelial spores (see Spori-

diola)

12
60

Promycelium

abnormal

• . .

• •

cells of, separating

elongated

in various genera
septa

• •

• - .

• • •

• •

• *

• •

• «

Protective inoculation

Protobasidii

Protomycelium

Pseudoparenchyma

Pseudoperidial cells (see Peridial cells)

Pseudoperidium (see Peridium)

Puccinia, introduced species

variation in teleutospores ...
Puccinia agropyri, heteroecious

PAGE.

1
35
35

27
33

27
62

40
4
4

agropyrina

agrostidis ...
anthoxanthi
arenariae, introduced

* • •

• •

• • •

• • *

• •

45
53

sporidiola germ-tubes enter-
ing by stomata

arrhenatheri, perennial myce-
lium

• • •

asparagi, influence of wind ...

water relation ...
beckmanniae, imported
bromina contrasted with P.

triticina ... 80

paraphyses in teleutosori ...24, 75
uredospore, duration of

8
3, 8
10
49
10
45
25

• ft *

germinating power
germination and infection
calendidae, marigold rust
Australian species
short cycle

caricis, heteroecious
cesatii

chrytanthemi, germination of
uredospores
introduced

two-celled uredospores
cichorii, introduced
cinerariae . . .
coronata . . .

• • •

« * •

♦ • •

C ft ft

cyani, introduced ...
dianthi, sporidiola germ-tubes

21
42
19
42
49
53
42

entering stomat a

diihondrae ,

variation

teleutospores

• • I

ft • •

111

• • •

dispersa, biologic forms

subdivision

• •

39
79

distincta, daisy rust, Aus-

tralian species
introduced on seed
short cycle

♦ • •

• •

glumarum, mycoplasm theory
graminella> with aecidium on

££^ciSS ... ... • ■ .

graminis, biologic forms

carried over from year to

49
209

•

18
79

year
in Australia
collective species
cycle of development
forms on special hosts
heteroecism
vigor

• • •

• * #

69
21, 69

58
53

increasing

General Index.

347

PAGE.

Puccinia graminw, importation on

straw

• • •

• •

injurious rust in Australia
no aecidia in Australia ...
on grasses, wheat, &c. ..•
repeated formation of

uredospores
reproduced by uredo -

43
64

spores in Australia

• • *

specialisation of forms ...
teleutospores,from Australia
not germinating in Eng-
land, and vice virsd ...
germinating power
germination, exposure to
cold not necessary ...

19
53

in Australia...
in water
time for
variation

• • •

... 24, 67

. . . o

uredospores, germinating
power

on old straw
germination ...
persist through winter . . .
graminis and barberry rust,
connexion first discovered

8
69
21

in Australia
in Kew Gardens ...
helianthi, germination

teleutospore

• • ■

55
... 58, 66

68
of

• • • t/

heterospora, one celled teleuto-

spores most common

* • •

transition form

• •

hieracii, spermogonia with
uredo...
subdivision of

• • •

impatientis, introduced
liliacearurti) spermogonia with

teleuto
lolii, introduced

80
43

paraphyses in uredosori ...
magnusiana, heteroecious ...
paraphyses, in teleutosori
and uredosori
malvacearwn, causing "shot-
hole"

... 13, 37

distribution

germination of teleutospores 24, 27
importation and widespread

43
14
42

suppression of spermogonia

maydis, introduced...
menthae, introduced
obtegens, spermogonia with

uredo...

37
49
29

42
paraphyses in uredosori ... 22, 29
uredo withstanding cold ... 22

perplexans, on native host ...
paraphyses in uredosori ...

plagianthiy epispore
poarum, introduced

podolepidis, two germ pores in

upper cell of teleutospores

prainiana, abnormal teleuto-
spore germination
prenanlhis, introduced

peridium often wanting ...

36
42

cinia prunt, causing "shot-hole
first record in Australia ...
producing uredospores

within fruits ...
supposed spermogonia
purpurea f introduced
rubigovera, germinating power
of uredospores
paraphyses in teleutosori ..

PAOK.

6
44

19
22
42

subdivision

• • •

senecionisy repetition of aecidia

simplex, aecidia unknown ...

introduced

mesospores
suaveoleris, scent

• • •

S
24

79
18
58
42
28

sper-
mogonia

tasmanica, germination oi

• • *

16
... 42, 49

aecidiospores
groundsel rust
thuemeniy introduced

attacking native host .. 42
triticina, aecidia unknown

rust

58
comparatively harmless ... 64, 66

contrasted with P. bromina

ospores

80
21

■— ' — — ----- »— - •— - ■ -w ^ -*^ «^— — v v ~ w w« w v «-r -v a. m www

preference for certain parts
vexansy amphispores

• • •

Pucciniaceae
Pucciniopsis
Queen's Jubilee wheat, rus

on grain ...
fianuculaceae, Cronartium

• • •

60
25

82
11

• t

69
53

periments

infection ex-

Ravenelia

compared with Uromycladium

Repetition, of aecidia ...

of spore forms

" spermogonia...

of uredospores

• •

104

11
14

1?)

Rhamnua

Rerraf, a rust-resisting wheat ... 64, 74

• • • t)*9

56
17

at hart tea

Rhododendron, Chrysomyxa

Poestelia

Rose rust (Phragmidium subcorti-

cium)

• •

Rotation of crops and rust
Rubiaceae, native rusts...
Rust Conference
Rust of wheat ...

• •

•

* •

45
71

connexion with rust on grasses
early theories
effect of, drainage ...

early maturing wheats ...

* • ■

fallowing

irrigation

manures

muggy weather

ploughing

rotation of crops

seedbed

seed treatment

stubble burning
effect on straw and grain
first appearance in season
first record in Australia

70
68

71
74

71
72

... 67| 72

71
73

• • •

348

General Index.

PAGK.

PAGE.

Rust of wheat, how carried over

from year to year

• #

• • •

• •

how spread

infection from within, myco-

plasm theory
influence on yield ...
losses from
mycelium not in seed
mycoplasm theory
not spread from other cereals
prevention and mitigation ...

question, present position in

Australia
spores, in soil

21, 68
69

4,70
65
64
70

4, 70
53

on seed

to destroy
wintering

• *

• •

• • •

• •

spraying impracticable

• •

• • %

Rust-liable soils

Rust -proof wheat unknown

Rust-resistance and structural

• •

64
69
69
73

8, 20

73
72

characters

♦ • •

• •

Rust-resisting wheats ...

Rusts, Australian, and their hosts
cycle of development

in heteroecious species
indigenous and introduced ...
influence of parasitic habit ...
relation to other fungi
sexuality

62,74

45
2, 10
58
42
40
40

14, 17, 75

Rusty straw and grain analyzed
Rye, rust

• •

• ■ •

• •

Saccoblastia ovispora
Saprophytes becoming parasitic
Scarlet pimpernel
Scent of spermogonia ...
Scrophulariaceae, Cronartium . . .

kD tSC/Cvt'O ••• ••• •••

Seed, perennial mycelium

not in wheat

Seed-bed and rust

Seed-treatment for rust

Seeds with aecidia

Selection and crossbreeding wheat

Self-sown wheat and rust
Senecio vulgaris
Septa...

21, 69,

65
53
32
52
43
13
53
80
5
70

71
73

• • •

- ■ .

formation in promycelia
Sexuality and nuclei

of aecidia

• •

16
3

... 14, 75
... 15, 17

Shot-hole caused, by Puccinia mal-

vacearum
by P. pruni

Silver wattle gall fungus ( Uromy-

cladium notabile) ...

Softening tissues for microscopical

examination
Soil moisture, influence on rust ...

Soils, rust-liable and rust- free ...
treatment with copper salts

6
6

12
10

to prevent disease

Sorus, uredo ...

Special corpuscles

Specialisation of parasitism
in Erysiphaceae

9 • •

• • •

52
54

Specific variation , gradations
Spermatia, embedded in sugary

secretion
formation ...
germination
not male cells
uninucleate

■ • •

• •

• . . 1 0 , ' o
. . . 1 «"> , . > /

Spermogonia, associated with cer-

tain spore forms ...
attractive to insects
Darluca filura mistaken for...
functionless

... 13, 37

... ±0) oo

• • •

importance
occurrence

• • •

origin
paraphyses

repetition of spore forms
suppressed

* •

Spore -bed

Spoie-forms

investigation
origin of principal
repetition...
suppression

Spores, distribution

• •

V •

* •

• *

13, 37, 39

13
38
13
14

14,37
19

7
11

31
11, 18, 19

• •

• • •

• •

• • •

germination, of aecidiospores
of amphispores
of spermatia
of sporidiola
of teleutospores

abnormal
of uredospores

• •

• • •

« • •

• • •

7
7, 16

*.. 13, 3/

8, 24, 27, 66

nuclei
on seed
parasites ...

wintering, of teleutospores
of uredospores

7, 19, 21, 27

14
69

• •

• • •

7,22

8
8, 20

Sporidiola, air necessary for for-

• • •

mation
conveyed by insects
distribution
earliest spore form ...
formation ...
germination ...

incapable of infecting grasses

bearing them
infection ...

• • •

28
32

27
27

origin

• • •

uninucleate

• • •

9,28

Spraying for rust of wheat im-
practicable
Staining spores

73
12

Starvation of host and infective

power of rust

• • •

Stellar ia media
Sterigma

Structural characters of host and

susceptibility, to disease ...

to wheat rust
Stubble burning and rust
Sugar, influence on germ tubes of

43
31

61
61

fungi

• • •

... 52, 62

Sulphate of copper, in seed treat-

ment for rust

• •

treatment of soil with
Suppression of spore forms
Surface markings of spores

• • t

• • «

73
62

•

General Index.

349

PAGK.

I'AOB.

Susceptibility of host, due to

physiological peculiarities
influence of structural charac-

ters

Teleutospores ...

• • •

• •

direct infection by germ tube

function ...

germination, factors influenc-

61, 62
23

mg
in Phragmidiumrubi

in Puccinia graminis
in P. malvacearum

• *

• • •

germ-pores
occurrence
origin and development

peculiarity

- • -

• « •

• •

simple or compound

two kinds...
uninucleate
variability, of form

of germination

• ■ •

Teleutsori paraphysate...

Tomato, copper salts and disease

Trichopsora

Trifolium repens ...

Triphragmium ulmariae

35
35

8, 24, 66

.. 24, 27

33, 35

24,75

63
32

5
36

• • ♦

• • t

primary and secondary uredo

spores

4 *

• • •

spermogonia with uredo

Uninucleate spores

Uredineae in relation to other

fungi

Uredinopsis

Uredo dispersa, histology

kuehnii, paraphyses

spyridii, paraphyses

20, 36

13,37
14

# •-•

• • •

• •

symphyti

• •

Uredosori, paraphyses ...

parasites ...
Uredospores, binucleate

derived from teleutospores . . .

distribution

formation...

function ...
germinating power

duration of
germination, in Puccinia chry-

sait themi
in P. rubigovera ...
germ tubes, piercing cells ...
infection by germ tubes

origin

primary and secondary

produced within fruit in Puc-

3
22

22, 29
23

22, 29

7,22

8
10

3
34
19

cinia pruni

• ft •

repeated formation

two celled, in Puccinia chry-

santhemi

wintering ...

19
19

Uromyces

• •

19
20, 21
24, 83

distinction between uredo-
spores and teleutospores . .

betae
bidbinis

caryophyllinus

84
42,84

85
43,85

Uromyces dactylidis, paraphyses
danthoniae. with aecidium (

grass
diploglottidi*

... 18, 57

euphorbiae, perennial mycelium 5, 70

fabae in Ecuador
limosellae ...
orchidearum, two-celled teleu-
tospores ... ... 39

phyllodiorum , paraphyses ... 24, 29

pisi ... ... ... 84

politus, two-celled teleuto-

spores ...
polygoni ...

• • •

... 43, 55

proeminens transition forms
from teleutospore to uredo-

spore
puccinioides
scutellatus, origin of uredo-

spore

• ■

uredo with teleuto spore ...

solid aginis

tricoryiiesy two-celled teleuto-
spores ...

trifolii — complete cycle
introduced
perennial mycelium

34
36
36

tuberculatum

• • •

. . . 39, 83

43
5

idosus. two-celled teleu-

tospores
Uromycladium . . .

• 9 •

forming galls often

mesospores

spermogonia

teleutospores

vesicles

• • •

... 3*7, Ot>

24, 83, 104

... 25, 2li

14
24
24

Uromycladium maritimum, occur-
rence of spermogonia

notabile

perennial mycelium and

galls ...

... 13, 37

• • •

6
13

robiwoni, spermogonia
tapper ianum, spermogonia ... 13, 37
perennial mycelium and

galls ...

• • •

• •

• • •

• ■ •

Uropyxis

Ustilagines

Vegetative organs, mycelium
Verrucose uredospores . . .
Vesicle in Uromycladium
Violet rust, native

hederaceae)
Water relation, for Puccinia on

ucc

208

• • •

asparagus ...

to infection, direct and m-

direct

• • •

9
6

Wattles, gall- fungi on ... • • •

Wheat rust. See Rust of Wheat.

• • • D^j # ^

Wheat, rust -resisting ...'
self-sown and rust ...
Wild oats and wheat rust
Wind distribution of spores
Wintering of rust spores
Witches' brooms

21, 69, 72

7
... 8,20

Brain. Government Printer, Melbou

.♦

•

■

•

■

&BB

A.' J>

THIS BOOK IS DUE ON THE LAST DATE

STAMPED BELOW

INITIAL FINE OF 25 CENTS

WILL BE ASSESSED FOR FAILURE TO RETURN
THIS BOOK ON THE DATE DUE. THE PENALTY
WILL INCREASE TO 50 CENTS ON THE FOURTH

DAY AND TO $1.00 ON THE SEVENTH DAY
OVER DUE.

334

, I***

I ■*

trl

* .

APR

193*

JAN 2 3 1963

>4f

' V

<\\

■*.'

f U

h ^*» -

- ^^h^^^^jj»W& ^<

lr> - X

•

• <

IP*

•

i *

-xm

im-mZ

■

f
/

LD 21 3m-8,'3°

fr? ^

s .

^t.

• •

• ,

■J

- ri>

tf*

;«,^-

»'#!

' 5fc>

1
d

--"T

. »>

'■■

F>-

•c

• fc

/ -

*fc

.•

. ,&

1 i

.lifk*

«• »

•

■ *

: ---':-.

, —

* • *v

V V ^B

i *

1^ t ' ■ - -

• < « ~ « - *

• •-•.#•-'•4. . « i k * . . \ 1^. 114 t #1 ? , 1 91 1 • 1

... . ■

' , ■ :

!«4> ; " J •

; :. .... ; r 1 _ ; . - .! ; :r

■■.'•♦*f-'»'-

P ;"-it -OBI;;

Bt2 *; ;**£*}'•{ §t;; 'iliffJrl ,::.

i:?|M |j{f iSiii jmiiiflAMfe*

4 •

- j

- I

< '

■

• . f

«• »

I * 1 • ■ • * ? ♦ % 4f 14**' 'iflUC

tifUf —***ttfr vV

f 1 TYlf .««.»• t^»f>i| -i. j *4*f

-,;>;:,; •;:.;„. — •
J]

i :■•::■■ ■, ; I

• 1 ■ • - 4 ...

- *>— • • • •* 4

r «

• :?»i:Mi ij^iiitiSKiti

•*

-:,&

* * • til

• 1*M 1 t K *■ "• • » 1 ,•»*•-* » % *

' M^PJJJJJH

■ »

Pl^Hlii^^pj^HpHpfp^H - * - -

-* « t

^ «

ihllfS

*1 T

* * *

» —

4 v

• * » «

* fk

ID; Hi- »J- .:i - IHilll. ilxr, tJuHi-. «•;;«

- t

■

■ ■

- * ->

p»«?

pfrr

3: r

itf

« *• *

1f4

■ V

*■*•**

- ■Bl?9>4f •" "4%™- *

*. •• .--. . . m * . •

* *•-—

^ * f

. N , ,

* 4

•~ — » ■

• •

A I

* 7 --

• t

41 * *■

A • <■

■ * *

7 .: : i

}

*t t

• * 1

t# •

4 • •.-" «

* *

■• I *

m m v

1 I

* - 4

-*l *— *

•

1 .

■

i ' • .

tt ■

jjlljjl

■

ii fl

'HpJHWtt^n RfflUIBffl

m\V •::ft|Hif!- '-•'{'; i :

in iffl n

■

HfliUlil^

* * a X ^ w iVl r 4 J I l&l iff 1 tMlllf llll'ijtlf 15 12t IU

' • ill il 0 n 1 III 1 it il ! Oiliiiifttfl

■ •

•

' • rlf r ll#I ill 24 it !tI§j llnlii^pitp^Bi^HI i

if j

! I f

i :

- •

• *

. .

_, •

■

•

. ' •

■

•

I li'

, * #

mnuitiiiutt.

rt ■ •'" #»r*

Internet Archive Audio

Featured

Top

Images

Featured

Top

Software

Featured

Top

Books

Featured

Top

Video

Featured

Top

Mobile Apps

Browser Extensions

Archive-It Subscription

Save Page Now

Full text of "The rusts of Australia: their structure, nature and classification"

See other formats