Skip to main content

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

See other formats












Government Vegetable Pathologist. 



l!To~6 . 


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 introduced 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 the 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 Klebah'n's Die wirtswechselnden Rostpihe [The Heteroecious 
Rusts], 1904. This will give a good idea of what has already been done 
in the investigation of the rust's, but no one can fail to appreciate the 
work accomplished by that veteran mycologist, Dr. Cooke, in his 



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. 
Rod way, 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 supplying 
me with any specimens required for verification or illustration. 

Xo 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. 




I. Introduction ... ... ... ... ... ... ... 1 

II. Vegetative Organs Mycelium ... ... ... ... ... 3 

III. Reproductive Organs Spores ... ... ... ... ... 7 

IV. Spermogonia and Spermatia ... ... ... ... ... 13 

V. Aecidia and Aecidiospores ... ... ... ... ... 16 

VI. Uredospores ... ... ... 19 

VII. Teleutospores ... ... ... ... ... ... ... 23 

VIII. Mesospores and Amphispores ... ... ... ... ... 25 

IX. Sporidiola or Promycelial Spores ... ... ... ... 27 

X. Paraphyses and their Function ... ... ... ... ... 29 

XI. Origin of the Principal Spore-forms ... ... ... ... 31 

XII. Rusts in their relation to other Fungi ... ... ... ... 40 

XIII. Indigenous and Introduced Species ... ... ... ... 42 

XIV. Indigenous Species with their Hosts ... ... ... ... 45 

XV. Australian Distribution ... ... ... ... ... ... 50 

XVI. Origin and Specialisation of Parasitism ... ... ... ... 52 

XVII. Heteroecism and its Origin ... ... ... ... ... 55 

XVIII. Predisposition ... ... ... ... ... ... ... 60 

XIX. The present position of the Rust in Wheat question in Australia ... 64 


XX. Classification with special reference to Biologic forms ... ... 79 

XXI. Systematic Arrangement and Technical Descriptions 

Uromyces ... ... ... ... ... ... ... 84 

Uromycladium ... ... ... ... ... .. 104 

Puccinia ... ... ... ... ... ... ... 112 

Phragmidium ... .. ... ... ... ... 185 

Cronartium ... ... ... ... ... ... 189 

Melampsora ... ... ... ... .. 191 

Caeoma ... ... ... ... ... ... .. 193 

Aecidium ... ... ... ... ... ... ... 194 

Uredo 202 

Excluded or Doubtful Species ... ... ... ... 207 

Glossary of the principal scienti lie terms used ... ... ... 211 

Literature consulted ... ... ... ... ... ... 213 

Explanation of Plates ... ... ... ... ... 222 

Host Index with Rusts ... ... ... ... ... 329 

Fungus Index with Sy 110113-1118 and Hosts .. ... .. 337 

General Index . ... 343 





. The Uredineae, or rusts, constitute one of the most important groups 
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: 

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

tissues of the host-plant, and may either produce from its 
myoelium 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 aecidio spores, uredospores, and teleuto spores in 

succession, the aecidio spores being generally preceded or 
accompanied by a peculiar form of spore, known as a 

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. 

2 Introduction. 

This succession of spore-forms may be represented graphically by the 
following diagrams: 



Teleutonport L 4- Aecidlospore 


Fio. 1. 


Teleutospore "V 7* Aecidiospore 


Teleutospore -*^ f* Uredospore 

Fio. a FIG. 4. 

The first shows the complete cycle of development in which all the 
spore-forms follow each other in invariable order, and this is the most 
common form. In the second the uredospore is suppressed, and the 
number of species undergoing this contracted cycle is much reduced. In 
the third the aecidiospore is wanting, and the number of forms is still 
further reduced. And in the 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, probably ranks next to 
those with a complete development. 

The spermatia succeed the sporidiola, and generally present in the 
life-cycle, but they do not enter into the general development. 

Besides these regular forms, there are others which are generally re- 
garded as representing either stages in the life-history of imperfectly known 
species or degraded forms of which only the uredospores or aecidiospores 
are known. 

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

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

Vegetative Organs. 



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, it' is worthy of the most careful 

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 
*tand out on account of their completeness, owing to the use of the most 
modern histological methods the one by Professor Marshall Ward 9 on the 
Histology of Uredo dispersa Eriks, and the Mycoplasm hypothesis, and the 
other by Professor Eriksson 14> 15 on Das vegetative Leben der Getretderost- 
pihe [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 young 
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 appressorium, 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 inmer 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 liaiistoria 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. 

Vegetative Organs. 

with an a^ed mternal germ of disease, which he considers in 
certain cases to be a source of rust, in addition to the ordinary infection 
by spores. It is often stated that this rust passes the winter as mycelium 
n 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 panted, then, that there is no mycelium to start with, 
and it will be interesting to follow Eriksson's theory as to the 
manner in which the mycelium arises afresh in the tissues IB certain cells 
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 mycoplasm is stated to occur only in certain cells 
which favours the assumption that it is not a necessary constituent of 
the cell. 

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 protomyctlium 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 fungus is a sign of advancing 
maturity. By continued division a true pseudo-parenchyma 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. 5 

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 evem 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, an3 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 
over winter in the seed, as in the case of Uromyces euphorbiae, according to 
Carleton 3 . 

Witches'-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 a/re 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 thalt 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 PL 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 

6 Vegetative Organs. 

considered as a morbid enlargement of the affected part of the plant, due 
to ; agency (Connold 1 ), then there need be no hesitation in calling 

those structures galls. 

Perhaps the most striking illustration of a gall is seen in Uromy- 
dadium 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 Myrniong on Acacia implexa of an irregular leg-of- 
mutton shape and weighing about 3 Ib. (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 
tlv-ir 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. Magnus 3 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 is cz.' 

The i>eculiar gall-like swellings caused by Gymno sporangium may be 
mentioned, the mycelium of which is perennial in the various species of 
Juniper, and from their appearance are popularly known as " cedar 
[ in America. It may be an annual gall only bearing the teleuto- 
spores for one season, or a perennial gall, producing- successive crops of 
teleutospore:? year after year, and not requiring the transfer of the spores 
each M ycdhtm. 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 
Jimalayan species of nettle attacked by this fungus that the natives eat 
the overgrown and hypertrophied stems for food. 

In other cases the affected tissues may be so stimulated bv the localized 

mycelium as to cause their death Thus almond leaves have been found 

, riddled "shot-hole." due to the mvcelium of Pucdnia 

prum Pers just as P. malvacearum may also destroy a circumscribed 

. and falls Out, 

e mycelium, whether localized or perennial, is always beneath the 
surface of the plant and formed within the living tissues. It 's delicate 
in texture. l,ke all mternal mvcelia, and branches to form a regular Vet 
work ultimately forming compact cushions or spore-beds. It can often be 
traced from a ungle point of infection, whence it radiates all round and 
spreads, gathering matem! for the fresh production of spores 

Re-productive Organs. 



A detailed account of the different spore-forms will be given in the sys- 
tematic part in connexion with the vairious 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. 


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 1 . The asparagus 
rust (Puccinia asparagi, 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 rusted! 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. 
Lindroth 1 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 (Salmon 1 ). 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 any decrease from this cause. 


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 easily bored in them, and glass rods placed parallel 
!>ehveen them, so that the microscopic slides may be laid across the rods. 

g Reproductive Organs. 

The influence of various chemical substances on germination has been in - 
vesLate^ and t has been found that some substances, such as solutions 
of su g gar and nitrate solutions, hasten germination ; but for general purposes 
water alone is all that is necessary. Sometimes, however the spores will 
germinate and infect the leaves of the host-plant, although they may nol 

d Freema^'found this to be the case in dealing with the uredo-spores 
of Puccinia 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 many 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. 


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. gramims, 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 

'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, ajs 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 Carleton 1 , 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 
(June to August). The uredospores can germinate at once, and directly 
infect 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 
lusts, only after undergoing a period of rest in the old world 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. 

Eriksson 10 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 one or two winters, and then on exposing them for another 
winter, he found that they germinated, but this was exceptional. 

Reproductive Organs. 9 

In other species there is considerable variation. According to Woronin 1 , 
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. 


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 Gibson 1 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 to 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 his 1 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 Pucdnia 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 of 
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 u 
favorable to the others. The indirect relation of water or the effect of soil 
moisture, which acts upon the parasite through ifts host, has also been 
studied and shown to be of great importance. An abundance or sojl 
moisture at the growing season, in the case of asparagus rust m 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 
by 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 by 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 4 
gives illustrations to show that the germ-tube of the uredospores of P. 
triticina may enter direct through the epidermal cells as well as by the 


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 

The aecidio, uredo, and teleutospore forms are represented respectively 
for convenience by the Roman numerals I. II., III., and if the sporidioia 
arising from the teleutospore are designated by IV., and spermogonia 
by a cypher, 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 
in the cycle, although .there are various rusts in which it has not yet been 
found, so that I. or II., or both, may be omitted or suppressed, as well as O, 
\\hirh, however, is rarely absent. The germinating sporidiolum produces a 
myo'lium which may in turn give rise to spermogonia and aecidia. These may 
either occur on the same species of host-plant as that which Dears 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. distincta). 

This variation in the alternation of spore-farms 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 
\\hirh luuv similar associated stages. Taking the genus Puccinia as an 

Reproductive Organs. n 

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- 
puccinia, 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 Puccima malv ace arum, 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- 

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


In connexion with the suppression or omission of spore- forms, we may 
also consider 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. 


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

12 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 1 lind 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. Por 
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 a 
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 photooraphed 

ier, and the general shape can be determined by comparison while 

size is deduced from the exact measurement of individuals in large 

In this way the average size has been obtained for the teleu/to- 

res chiefly, of all the species accessible for observation. At the same 

ge number were measured direct by an eye-piece micrometer 

and I the results compared. All the microscopic investigations were irade 

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 honey dew 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 (the 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 Arthur 3 on the Taxo- 
notnic Importance of the Spermogonium, he shows that, in the first pla^t, 
ihe 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. 

I n 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 spermogonioim is the only one which does not repeat itself, 
as stated by Arthur 3 : "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 malvaceamm 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-Trouffy 2 , for instance, has observed in Uredineae having the 
various stages the following nuclear cycle : 


Mature teleutospores. 


Mvn-lium. producing spermatia and 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 

s nuclei may be interpreted otherwise. Blackman 2 has confirmed the 

ove nuclear cycle, and concludes that the spermatia are male cells which 

have become functionless, the nuclear characters being those of male cells 

and not of spores. 

The fertile cell of the aecidium or primary aecidiospore becomes bi- 

leate by the nucleus of a neighbouring vegetative cell migrating through 

>e wall, and this association of the two nuclei Blackman considers to be 

Hindoo 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.) 

Spermogonia and S-permatia. 15 

Massee also considers the aecidium to be a sexual product, and both 
authors ag,ree that the spermatia are not concerned in it. If the spermogonia 
and aecidia represented male and female reproductive 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 gametophyte starting with the uni-nucleate teleutospore. 
But the most probable view is that the spermatium is a functionless organ, 
and its presence is an indication that the ancestors of the rusts may have 
possessed an alternation of generations similar to that referred to above. 

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, with an orange-yellow or 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 post. 

Aecidia and Aecidiospores. 17 

The development of the aecidia has been traced by De Bary, Neumann 1 , 
and others. The hyphae derived from the promycelial spores form at 
certain points,, 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, Fischer 1 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. Mayus 1 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-peridium 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. 
Massee 1 , 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 
Fungus 1 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.'' 

1 8 Aecidia and Aecidiospores. 

Hut 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 Helicmthus annum, 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, only 
one instance was known, viz., Puccinia graminella, in which the aecidia 
and teleutospores occur together, the latter being often very rare, or wanting 

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. 




_ _ Aecidiospores 



t This repeated formation of aecidia was principally investigated bv 

.el, who named the aecidia arising directly from the germination of 

primary aecidia," and -those arising from the germination 

of aecidiospores "secondary aecidia;" and it was noticeable that spermo- 

nia were usually absent from the latter. Thus the germinating spori- 

um may produce a mycelium which bears only aecidia; but the aecidio- 

s, instead of giving rise directly to teleutospores, may repeat them- 

everal generations, and then give rise to teleutospores. How- 

x>mplete cvcle the aecidia originate only from sporidiola, while 


Uredospores. 19 



Uredospores may originate from the hyphae developed from the germ- 
tube of an aecidiospore, a promycelial spore, or another uredospore. They 
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 Chrysomyxa, 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 ibeen described and 
drawn by Roze l and Jacky 2 , in Puccinia chrysanthemi, 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 Uromvces on the 
other, in having two or more germ-pores, and this character also dis- 
tinguishes them from the .mesospores. Only in exceptional cases 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 only 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 5 mm. 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 Uredospares. 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 
either by infection from the promycelial spores, aecidiospores, or, it may 




FIG. 6. 

he, from a perennial mycelium. The spore-layer is usually distinguished 
by its larger size, and the corresponding injury it causes ^ to the host- 
plant; while 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 Triphragimum 
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 
leaves, while those of the latter are small 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, epiteosporiferotis 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 Lagerheim 1 , 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 
aufuimn months of April and May, or even earlier, and the harvest is 

Uredospores. 2 1 

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 (teleutospores, 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 Kmits, 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 Avas 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 

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 Henning 1 ) 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 Jacky 2 found that the uredospores of chrysanthemum 
rust still retained their germinating power, after exposure to the weather 
for 66 days, from ist December to 5th February. And Miss Gibson kept 
spores of the same rust in a dry test-tube in a cool room for 7 1 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 

22 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, ;unl <\irrv on the life of the species, independent of the teleuto- 
spore. This is well seen in Pucdnia 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. vcxans, 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 Pucdnia. 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 are 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-layers, 
and is recorded on Uromyces (9), Uromycladium (i), Pucdnia (22), Phrag- 
midium (i), and Aecidittm (2). 

Paraphyses most commonly occur in connexion with the uredosori, and 
are found in Australian species of Puccmia, Melampsora, and Phragn:idi::m 
as well as in Uredo. 

Occasionally they arise in both uredo and teleuto-sori, as in Pucdnia 
magnusiana and P. purpurea, and sometimes they are variable in their 
presence as in Pucdnia poarum, where Plowright found none in Britain, 
although they are common in Australian specimens. In Phragmidiuin 
subcortidum 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 paraphyses in 
tneir uredosori : Pucdnia lotii, P. magnusiana, P. poarum, P. ' pruni, 
P. pnrpurca; Melampsora hypericorum, M. lint; Phragmidium barnardi, 
P. subcortidum; Uredo kuehnii, and U. spyridit. 

Teleutos-pores. 23 



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 symphyti, 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 Bubak 2 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, hut 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. Sydow 1 states that in all Leptopuccinieae, or those only 
possessing iteleutospores 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, and 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 they germinate as 
soon as they are ripe, and the fungus winters by means of its mycelium 
in the host-plant. 




This genus is so anomalous, that it is now coming to be regarded, not 
an independent form, but as a biologic genus in connexion with Puccima 
or I'romyces. 

Teleutospores may be simple, as in Uromyces, or compound as i 
Puccinia. In the newly-constituted Australian genus, Uromycladtum, the 
teleutospores are 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 may be warted or striated in Urpmycladuim. 

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

'uccinia 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 1 has stated that there is only one 
germ-pore in each cell of P. barnardi, but three were invariably found by 
me in examining a large amount of material. 

Para-physes 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-known ex- 
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. -pnrpurea. and Uromyces -phyllodiorum. 

The germination of the teleutospores of Puccima graminis in Australia 
was tested under different conditions, and the most important condition 
seems to be the season of the year, for they were only known to germinate 
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 opem, 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 might 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. 



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 characteristics. The conclusion one would naturally draw 
is that they are teleutospores in the process of making, with t'Jie 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 gent-rally 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. 


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. Farlow 1 
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 

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. 

Besides the amphispores, mesospores are very numerous in this species, 
so that there is a Puccinia 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 Arthur 5 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, just like a teleutospore. 


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- 
spc-ra 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. malvaceantm, 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 
the other. But it is so difficult to draw the line sometimes, and since it is 
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- 
nrnation, 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 

Sporidiola or Promycehal Spores. 27 



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 my 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 * 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 ithe 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. 

with 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 aje 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 that 
P. gramhris 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, Plowright 2 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 germinaiting 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. 29 



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 spermaitia 
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 Melampsora and some species of Puccinia and Uredo, e.g., 
the uredo of P. poartim, 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 Dietel 9 . 

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 thatt 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 fuingus, 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 1 so many spores, and barren protective filamesnits 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, tEere 
would be little use for them ; but towards the summer, when the air is 

30 Paraphyses. 

ilry, 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 Ot-th. 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 pomum 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 Koern. 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 

Another function has been suggested by Magnus 6 in addition to \that 
of protection. In several species of Coleosporium he found that the 
paraphyses served both for protection and for raising and bursting the 
epidermis so as to make room for the growing spores. 

To a certain extent paraphyses may assist in raising the cuticle and 
hastening its rupture; but there are plenty of species which rupture the 
cuticle in the absence of paraphyses, and in the case of paraphyses asso- 
ciated with teleutospores, they rather appear to prevent the cuticle break- 
ing away until it decays. 

Origin of Spore-forms. 31 



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 
iormed. 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 
first 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.) 

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 1 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. 

32 Origin of Spore-forms. 


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. !). 

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 1 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 Coleosforium is evidently 
the representative of the septate basidium in the Auriculadaceae, although it 
s generally considered to be an exceptional form of teleutospore, which 
occurs in other genera of Rusts as well, such as Ockrojtsora, Trichofsora, 
and Chrysopsora. If the basidia in one of the saprophytic Auriculariaceae, 
s Saccoblastia ovtspora Moell., are compared 'with those of the 
parasitic Coltosponum senecionis, there is seen to be complete agreement in 
the structure. 

Origin of Spore-forms. 33 

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 f ungus'. 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. 


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 
growth 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 1 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). 

FIG. 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 Arthur 6 , to which he has given the 
name of Baeodromus. It occurs on Senecio, and resembles Coleos-porium 

34 Origin of Spore-forms. 

senecioms so much in general appearance, that it was at first regarded as 
that species, but the teleutospore produces a regular promycelium, with 
sterigmata and sporidiola, so that it well illustrates the transition from the 
basitlia bearing sterigmata in Coleosporium to the typical teleutospore 
with a promvoelium. The teleutospores are united into a solid mass and 
are arranged 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 regards 
as a multicellular 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 Chrysomyxa, in 

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 cells. 

A teleutospore is thusi a unicellular or multicellular spore, producing on 
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. 


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 first 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. Magnus 2 , 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- 

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 Avhich the wall still retains its thickness, but there 
are several germ-pores. They may resemble the uredospores in shape, 

Origin of Spore-forms. 35 

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 Gymno sporangium 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 as an equatorial band. But 
here again Magnus shows in species such as U. proeminens (DC.) Pass, 
and U. tuber culatus, Fckl., every transition from thq 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 

In Puccinia podolepidis there is often a germ-pore on either side of the 
upper cell of the teleutospore, as shown in PI. 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 asf 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-Gerloff 1 has shown that in Gym no sporangium clavariaeforme, 
Jacq., 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. Fischer 1 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 ordinary course, rounded 
themselves off and separated. Fischer observed in Gymno sporangium 
confusum that the cells of the promycelium separated before forming sterig- 
mata and promycelial spores, but Blackman, in a MS. note to his paper 1 , 
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 finally the promycelium bearing promycelial 

-6 Origin of Spore-forms. 

It is worthy of mention that Magnus 2 has repeatedly obsewed 

that when the teleutospores of Puccinia graminis were germinated in water 

te- produced an undivided germ-tube just like that of a uredospore, but; 

he could not satisfactorily settle the question, if this germ-tube could directly 

a thick to a thin wall, from one to a number ol 
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 e'ach promy- 
celium breaks up into three or four detached cells', apparently representing 
sporidiola, had been observed by Barclay' as early as 1891, in -P^cirna 
firainiana, Bard, 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 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 ^n 
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 
\vell -developed spore-layer they always precede the associated teleutospores, 
but in Uromyces scutellatus', as Magnus 2 has repeatedly observed, in 
different years they may appear simultaneously. 

The uredospores 1 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 aecidios pores. 


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 Phragmidium 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 Traverso l speak of them as epiteospores, to 
distinguish them from caeomospores, which are arranged in chains. There 

Origin of Spore-forms. 37 

is no essential point of distinction between aecidiosporcs 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-celled pTomycelium, 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 earl} origin 
from a teleutospore in its mode of germination, (Note 4, p. 75.) 


If the aecidiospores were the ^result 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 uhuariae, Uromy- 
cladium maritimum, and Puccinia obtegens (Lk.) Tul., then they accompany 
it, and if a teleutospore as in P. liliacearum, Duby. or Uromycladimn tep~ 
perianum, the spermogonia are associated with it. But Arthur 3 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 it'o 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 

38 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 (the 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 
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- 
pendent body capable of giving rise directly and without any further 
ood supplv 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. 39 

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 
tteady 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 prom'ycelial 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 


When the teleutospores in the different genera of rusts are carefully 
examined, it is often found that there is not only considerable variation in 
(he 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 (PI. 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. dichondrae^ 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 type; (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 and transversely divided, as in Sphaerophragmium. 
It is worthy of note (that there are no uredospores in P. dichondrae, but, 
even in P. hidwigii, 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 

40 Relation of Rusts 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 : 

( Ascomycetes 

Phycomycetes (Alga-like Fungi) J B (Hemibasidii (Smuts) 

' j Basidiomycetes 4 Protobasidii (Rusts) 

[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 srnuts 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. 41 

plants for their sustenance, they must accommodate themselves to their 
surroundings, and so they may pass through different stages, sometimes on 
the same plant, or on different plants. These various stages represented 
by different reproductive bodies, enable the rust to produce spores which 
can germinate at once if food supplies are available, or others which can 
rest if need be. The change of host is an evident advantage, not only 
from the point of view of a change of diet, but it may thus pass from one 
plant that dies down to another that is perennial. It may even become 
perennial itself in the underground parts of some plants, and then it 
vegetates, only producing rarely the reproductive bodies which would 
be formed under normal conditions. 

42 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 always 
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 fro stratum) which was 
possibly introduced with cultivated celery (Apium gravtolens), P. menthae 
on native minit introduced upon cultivated mint, and Melampsora 
lint, on. native and cultivated flax. On the other hand, species 
of rust occur on well-known imparted 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 raire 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 majority of the rusts here recorded are, of course, 
native, 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 Anthoxanrthum odoratum (1896). 
P. arenariae (Schum.), Schroet. on Stellaria media (1896). 
P. beckmanniae n. sp. on Beckmannia erucaefarmis (1904). 
P. chrysanthemi Roze, on Chrysanthemum indicum (1904). 
P. cichorii (DC.) Bell., on Cichorium intybus (1885). 
P. cvani (Schleich.) Pass, on Centaurea cyanus (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 catiano- (100=;) 
U. betae (Pers.), Kuehn, on Beta vulgaris (1878). 

The year in brackets indicates wten first recorded for or observed in Australia"! 

Indigenous and Introduced Species. 43 

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

U. fab'ae (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. Holway 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, 
and the rust (Puccinia im-patientis) appeared upon the plants in December 
of the same year. 

The latest addition to, our imported rust-fungi is that of Puccinia 
chry'santhemi, 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 likely 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 malvacearmri) 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 over 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), Italy (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 
Industry, U.S.A. Bolley, in a letter dated 29-th 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 Phragmidiwn 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 Phragrnidium, 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 ait 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 ^yild 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 



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- 
plants, 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 by 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 naitive 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. 



S | 

















* X 2 




























004 ~ 









8M > 




if! " 








.Jl l8| 


-i. If 111 







cs s ,0 o'w S a S -3 



-3-52, 5 






















Indigenous Species with their Hosts. 





3 HI 














1 s 







c >> 



































. carissae 


. mussoni 

. tetragoniae 

. loranthicola 





- - 











































Indigenous Species with their Hosts. 









A. nymphoi 

A. calthae 
A. ranuncul 

A. plantagi 








r J 

i i 





















3J. i 




|f|l s 


J2* O S Hji^f 





P;^; p; 

























. hypoxidis 















S I | i i i 1 


II lilt 4i i 8 H i H 
fill!!!] I 1 i j 1 I I 

Indigenous Species with their Hosts. 49 

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 -pi ants, but such rusts may either be confined exclusively to 
native planits 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 (Senccio vulgaris), but one stage of it is also found 
on native species of the same genus, and, therefore, it is regarded as in- 
digenous. 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. 
distincta. As regards introduced rusts, they may be found on both 
native and imported plants, the presumption being ithat 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 
Puccima 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, siuch as 
Puccinia chrysanthemi and Phragmidium subcorticium. 

Australian Distribution. 



It would be premature to attempt to arrange the Australian rusts in 
geographical districts, since they are as yet too imperfectly known, and 
large areas have not been explored sufficiently to give any exact idea as 
to the number of species or the distribution of those already known^ So 
for the present I will content myself with indicating their distribution m 
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 b 
lound 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. 

Uromyces ... 27 13 

Uromycladium ... 7 

Puccinia ... 90 ... 24 

Phragmidium ... 4 ... 4 

Cronartium ... i ... i 

Melampsora ... 2 .., 3 

Roestelia ... ... i 

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 Uromyces 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. 51 

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 



Now South 



Austral! ii. 

























Phragmidium .. 







Cronartium . . . 






Melampsora . . . 






















i 3 
















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 every 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 
preparatory 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 

It is assumed that the saprophytic habit was the normal one among 
fungi, and that parasitism is an acquired habit. Massee 9 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 positively 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 
flue 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, Massee 9 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 action 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 

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. 53 

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 Puccima 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 alnus, while that 
on Lolium perenne required for its aecidial host Rhamnus catliartica. Hence 
the old species was split up into two, which can also be separated by mor- 
phological characters. Eriksson 1 also proved in 1894 that the well- 
known and much-investigated species of Puccima 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. agro-pyrina, &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 l 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, 
Massee 9 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 

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. 'Fischer 3 and 
Klebahn 1 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, 
it may be referred to the " internal developmental tendencies 
Klebahn l until a better explanation is forthcoming. 

54 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. 

Neger 1 , in 1902, proved by numerous experiments that there were 
biologic forms of Oidium for several species of Erysiphe, and Marchal 1 
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. 



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, ialthough it is always 
desirable, where possible, to have experimental proof of it. In Uromyces 
fiolygoni, 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 plant 
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, Pucdnia 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 ithat 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 

But there is no evidence to prove that such a gradual separation took 
place, for even although the species of Pucdnia 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, (the view that heteroecism originated 
suddenly and without the different generations slowly and gradually pass- 
ing from plant to plant, 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. c 

5 6 Heteroecism. 

proceeded from the formation of teleutospores being succeeded by aecidio- 
spores, presumably at first both arising from the same mycelium. Next a 
division of labour took place, and the, mycelium of the aecidia was pro- 
duced by the sporidiola, while the mycelium of the teleutospore proceeded 
from the aecidiospore. The advance to heteroecism took place when the 
aecidiospores produced their mycelium in one host-plant and the : 
spores, through the sporidiola, in another, and the kerne 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-forms 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 
so But this latter view is hardly borne out by some experiments conducted 

liss Gibson 1 , in which the aecidia from different host-plants were use 
to infect Ranunculus ficaria, and while the germ-tube as a rule entered 
the stoma freely, 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 incapacity to penetrate the cells is due to lack ot attractive 
substance or to the presence of anything actively repellent is not clear 
though, as before stated, certain facts seem to suggest the presence ot 
something harmful to the hyphae." 

A few concrete examples may 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. rlw- 
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 
abietimim, Alb. and Schw. C, ledi (Alb. and Schw.) De Bary, forms uredo 
and teleutospores on Ledum palustre, and its aecidia also on Picea 
excelsa, there being little or no distinction between them and those of 
C. rhododendri. The third, C. ab'ietis (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. 

Barclay 2 , 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: 

1. P. roronata, Corda, I. Franguli 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. 

Hcteroechm. 57 

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


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

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

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

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 

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 regular 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 

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 aecicliospores 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 


This course of development may be represented graphically as follows, 
taking P. gr ambus as the type: 

, in Europe. &, in Australia. 

Spennatui Teleutospores 

Teleutos pores 



FIG. 14. 

Fio. 15. 

The aecidial stage of the spring rusts of wheat (P. tritidna) 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 plants for their continued existence, but while this 
is usually accomplished by means of the teleutospores, other and additional 
measures may be taken .to secure the same result. Sometimes the uredo- 
mycelium persists during the winter, and in the case of Pucdnia arrhenatheri 
the aecidium-mycelium becomes perennial, and reproduces the aecid'ia year 
after year, while the teleutospore is also produced. When the aecidiai 
stage is dropped, as in the case of Pucdnia graminis in Australia, then there 
is a profuse development of uredospores in comparison with teleutospores, 
and there is abundance forf present needs, as well as for future germination 
in the spring. In other cases where the aecidial host is absent, the same 
thing has been observed, as in Caleosporium senedonis, when occurring in a 
district destitute of fir trees, or Chrysomyxa rhododendri when the silver fir 
is absent. 

Heteroecism ds said to increase the vigour of the fungus, and a striking 
illustration is given by Pucdnia graminis on wheat. It is sometimes said 
that the aecidiospores from barberry are much more virulent than the 
uredosppres derived from the wheat itself, since the teleutospores produced 
are earlier, more copious, and more injurious to the wheat. With reference 
to this, Plowright says : " There is a wonderful difference in the amount 
of injury done by mildew, when derived directly from the barberry, and 
when derived from uredo that has reproduced itself through several gener- 
ations. . . . The fungus grows with such energy that it so injures the 
\vheat plant as to prevent it producing more than a few starved kernels." 
As against this view, it may* be well to bear in mind that in no country 
in the \yorld probably does P. graminis cause as great injury to wheat as 
it does in Australia, a country with barberries practically non-existent, amd 
in which the aecidial stage has never been found. 

Of course further observations on a number of species are necessary 
:abhsh the fact, but Klebahn considers that the utilization of the 
vegetative periods and other peculiarities of the host-plants, rendered pos- 
sible by the change of hosts, gives the fungus a decided advantage. 

Heteroecism. rp 

Whatever view we take as to the cause of heteroecism, it is a well- 
established fact that the promycelial spores, no matter how freely applied 
produce no effect on the grasses which bear them, and the most probable 
theory as to its origin assumes that the uredo and teleuto spore generation 
were at first associated with the aecidia, but ultimately passed over to 
other hosts. 

60 Predisposition. 


Wherever epidemic diseases caused by parasitic fungi occur, the ques- 
tion is raised as to the relative susceptibility of different varieties or indi- 
viduals. The fungus has the power of causing disease in the host plant 
attacked, but the latter in its turn may either be favorably or unfavor- 
ably disposed towards its development. This predisposition of the host 
for the attacks of the parasite is very variable, and is influenced by vari- 
ous factors. It is generally considered that a sickly plant is more liable 
to rust than a sound one ; but, on the contrary, strong and sound individuals 
are more easily and more virulently attacked, so that for artificial infection 
strong-growing plants are selected. Ward 7 has shown that when the host- 
plant is starved by withholding/ certain mineral salts, and thus stunted in 
growth and generally enfeebled, it is not affected in its susceptibility or 
otherwise. A starved plant certainly develops smaller pustules and fewer 
spores on account of the diminished supplies of food available for the 
mycelium, ibut the power of infection is just as great as in normal plants. 
As far as brome rust is concerned and we have no reason to doubt that 
it holds good for others as well predisposition and immunity on the part 
of the host, and impotence and virulence on the part of the parasite are 
alike independent of mere nutrition. But, as we shall see afterwards, 
certain substances not of the nature of food-material, introduced into the 
plant may affect its liability to disease. There are various factors, how- 
ever, which may either dispose the plant towards disease or tend to render 
it immune, and some of these may be given here. 

The age of the part attacked has an important influence on infection, 
particularly where the sporidiola are concerned. Young leaves and shoots 
are most easily infected, and when they get older little or no effect is pro- 
duced. This is owing to the germ-tube of the sporidiolum penetrating the 
epidermis direct, and it is well known that this laver becomes firmer and 
tougher and less easily penetrable as it gets older. The uredospores 
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 variously affected. Some- 
times it is the leaves, sometimes leaf and stem, and it may be on one or 
both sides of the leaf. The different species of rust on the same host- 
plant are apt to choose different portions. Thus Puccinia triticina, from 
its earlier attack, is found most commonly on the lower leaves, and extends 
on to the sheath, mostly near its junction with the leaf, while P. graminis 
5 worse on the upper leaves, and often particularly bad on sheath and 

^Different varieties or sorts of the same species vary considerablv in 

;ity, and, as will be shown later, it is by the selection 'and 

such rust-resistant sorts that solution of the rust-in-wheat ques- 

m Australia is being attempted. The same is the case with other 

have seen one kind of flax (Linum usitatissimum) badly 

l.y Mclampsora /*/, and another kind growing alongside quite 

Jennings 5 has recently made observations which tend to show that 

lants previously susceptible to the attacks of a parasitic fungus mav gradu- 

come immune when they are changed to rich ground where they are 

better nourished and more vigorous. 

Predisposition. 61 

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 importation. 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 plaints only showed the fungus very slightly 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. Salmon 3 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 

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 Bary 2 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. 

Hennings 2 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 
leaf may affect the result. 

Marshall Ward 1 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. ' ' 

Salmon 3 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 

6 , 


typical characters associated with the quality of rust ^ s ^ a " ce ' 
would appear therefore to have a share in bringing albout this, re suit. 

rust-resistant wheat, the leaves have a tough cuticle, and the straw 
is well glared often with a glaucous bloom. The flag is narrow 
not broad, erect not drooping, stiff and firm, not soft and flabby. 
Still all these characters may be present, and yet the climatic conditions 
may overrule them and weaken the power of rust-resistance. It will thus 
be 'seen that a variety of causes may contribute to rendering a plant im- 
mune, and that not only the chemical and other properties of protoplasm 
come into play, but there is also the mutual reaction of the living 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 another 
rust will appear. We express this by saying that there are rust-resistant 
wheats, but not " rust-proof." 

Animals are sometimes rendered immune, or, at least less susceptible 
to disease, by repeated infections, so that they become gradually accus- 
tomed to the effects of the parasite, and the important question arises, Can 
plants also be submitted to " protective inoculation " ? Inoculation witl 
attenuated bacteria is the simplest method with animals, and this treatment 
retards their vegetative development, and so lessens their injurious effects. 
This artificially-induced immunity only lasts a certain time, varying; in 
different cases. Ehrlich and Huebener 1 have shown, from numerous 
experiments upon animals, that an immunized mother can impart immunity 
to her offspring, but it is not lasting. 

In the case of plants, as in animals, there seem to be two opposing 
forces at work. On the one hand, the protoplasm of the fungus is 
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" or "partially immune" by inoculation with the parasitic fungus 
to protect it against further attack, but advantage has been taken of the 
great sensitiveness of certain spores to copper salts to protect the organism 
against them. (Note 6, p. 75.) 

Chemotaxis is the name given by Pfeffer 1 to a form of sensitiveness 
which certain organisms possess towards certain chemical substances. This 
power, which certain nutritive and other substances have of attracting 
bacteria and other organisms towards them, is known as -positive 
chemotaxis, and of organic substances with a high nutritive 
value which are positively chemofactic, may be mentioned asparagin an< 
peptone, while sugar, which is one of the best food stuffs and rich( 
sources of energy, has but little attractive power. The power of othe 
substances, on the contrary, to repel bacteria, is know as negative chei 
taxis, and free acids and alkalies, as well as alcohol, have this effe 
Glycerine, as far as known, is an inactive substance. Massee 9 has stal 
that immunity is owing to the absence or small proportion of the substance 
chemotactic to the parasite in the plant not attacked, and if a plant can 
be impregnated with some substance which is negatively chemotactic, and 
at the same time does not affect the utility of the plant, then immunity 
against parasitic fungi may be obtained in this way. 

Laurent 1 , acting on this principle, conducted a series of experiments 
to test the possibility of producing potatoes which would be proof against 
Phyto-phthora infestans. He grew very susceptible varieties in pots, to 
the soil of which sulphate of copper was added, and when the tubers were 
harvested, some were cut in two and their cut surface placed in contact with 

Predisposition. 63 

the mildew of potato leaves. After four days, the tubers grown in the 
soil containing copper did not show any infection, while those girown 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 pro- 
tection against rusts, but without success, 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. 

Massee 7 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 
coppesr '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. 

64 Wheat Rust in Australia. 


It will tend to clearness if we con-fine our attention to the rust in wheat, 
for the general considerations which hold in this case will apply to the rusts 
on the other cereals chiefly cultivated, viz., oats and barley. And the 
rust which is most important from the farmer's point of view, because* it 
does the most damage, will be chosen viz., Puccinia graminis, which is so 
distinct in its characters that there is no difficulty in recognising, it. There 
are only two kinds of rust in wheat in Australia, the positively injurious 
Puccinia graminis and the comparatively harmless P. triticina, because it 
does not pinch and shrivel the grain like the other. In the early days wheat 
and other cereals had to be imported into Sydney, but now it is grown to 
such an extent in the Australian Commonwealth that in the season 1903-4 
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 industry, whatever increases the yield or tends to diminish 
the losses from disease, will have a corresponding far-reaching effect. Ever 
since attention has been given to the subject, it has been found that not a 
year passes without its being present on wheat to a greater or less extent, 
and in some years, which are commonly spoken of as rusty years, it seriously 
injures the grain and considerably diminishes the yield. To give some con 
crete idea of the extent of the loss, it may be stated that in a particularly 
bad season like that of 1889, the loss was estimated to be for the whole o 
Australia between ^2, 000,000 and ^3, 000,000. In such favorable years 
for the rust, there is always an evident connexion between the weather and 
its epidemic nature. In 1889 it was a wet spring, and about the blooming 
season the weather was what was known as " muggy," consisting of showers 
with heat between, and heavy dews, so that the wheat-plant, at the time of 
coming into flower, was particularly susceptible, and the rust spores found 
a ready entrance into the tissues of the plant, with their accumulated stores 
of food, and thus it spread rapidly. 

Atmospheric conditions a,re often regarded as the prime factors in the 
production of rust, but they are only concerned in the matter in so far as 
they favour or hinder the development of the parasite which causes it, and 
this is strikingly shown in the existence of rust-resisting wheats. Owing to 
the importance of the subject, and the widespread losses, delegates from the 
various States were appointed to meet and confer periodically, and these 
various conferences extended from the first in 1890 to the last in 1896. 
The results of their labours may be seen in their voluminous reports, which 
practically cover the different phases of the question. But there were some 
important points with which they were not then in a position to deal, such 
as the effects of rust on the straw and grain, and experiments in the inocu- 
lation of the bartberry. These will be considered preliminary to the main 


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

-IV., where the straw of the rust-resistant Rerraf is bright, glossy deep 
yellow, and well ripened, while that of the badly rusted Queen's jubilee has 
a dirty, streaky, patchy appearance. The ears of the one are full-sized and 
well formed, with plump, well-filled grain, and the other has poor ears 
with light and shrivelled grain. 

Wheat Rust in Australia. 65 

But the effect upon the composition of the plant, and particularly its 
feeding value, is not known to the farmer, who cuts his crop for hay when 
rust threatens to ruin it. 

This has recently been determined by F. T. Shutt l , chemist, Dominion 
Experimental Farms, Canada, who analyzed two samples of wheat grown 
at Manitoba in the same field and of the same age, only the one was rusted 
and the other rust free, so sthat the results are strictly comparable. The 
analysis is as follows: 
























Straw from rust-free wheat 

7' 9 2 






Straw from rusted wheat 

.. . 






Grain from rust-free wheat 







Grain from rusted wheat 

I -4944 







The Straw. It is pointed out that in crude protein the rusted straw is 
much richer, and since this includes all the nitrogenous compounds of a food 
that go to repair waste, form blood and build up muscle, it may safely be 
concluded that the rusted straw is much superior in feeding value. There 
is also in the rusted straw slightly more fat and somewhat less fibre, so 
that all this affords additional evidence of its more highly nutritious nature. 

The handling and feeding of rusty straw in Australia from the farmer's 
point of view has received attention. The experience of one who has done 
a deal of threshing is to the effect that when very bad it caused an itchy 
sensation, and made the men about the thresher rub their skin until it was 
broken. As regards feeding rusty hay, another with large experience 
informs me that horses and cattle relish it far before ordinary hay. Of 
course, it was fed as chaff. 

The Grain. This from the rusted wheat is only about one-half the 
weight of that from the rust- free wheat, but as the protein content shows, 
it has, weight for weight, a considerably higher nutritive value. He 
accounts for the higher protein content in the smaller grain in its larger pro- 
portion of bran, but chiefly in the partial and incomplete transference and 
accumulation 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. 

begins to form the grain, and if that season is favorable for the develop- 
ment of rust, then the fungus has to draw upon the stored-up material, and 
consequently the grain is not fully formed, if at all. Incidentally this 
investigation also throws light upon the reason why the earlier rust (P. 
iriticind) 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. 

On account of the comparatively late appearance of P. gramtnis, it is 
by some considered to do the least damage to the grain, but, as a^ matter 
of observation, and on physiological grounds, it is known, at least in Aus- 
tralia, to be the most injurious. 



In Europe and America the identity of P. graminis is determined, not 
merely from its morphological characters, but from its ability to infect the 
barberry and produce aecidia. But in Australia infection of the barberry- 
has not been successful, although several attempts have been made, and 
some have doubted whether we have got the true P. graminis, and not a 
distinct biological form of it. From a comparison with European speci- 
mens and a critical examination of the sori, 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 this rust, therefore, to the barberry in Australia 
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, 
too, 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 zraminis of 

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. 67 

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 i6th 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 Gardeners' 
Chronicle for i5th January, 1898, he says 6 : "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 
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 Rave been made in the English autumn. As 
far as Australia is concerned, the rust which does the principal damage 
has apparently no intermediate, stage. 

gg Wheat Rust in Australia. 

The Australian rust has been determined as Puccinia gramas by 
Eriksson and other authorities; but if the ability to produce the aecidmm 
>n the barberrv is taken as the final criterion, then judgment must be 
suspended in 'numerous other cases. Thus Massee * has found P. 
wmiis on Alopecurus pratensis, and Avena elatior in the Royal Gardens 
ut he significantly adds-- Notwithstanding the great quantity of 
Jfcrfefii and Mahonia present in the grounds, the most careful and con- 
tim, , has failed to reveal the presence of the aecidiospore stage. 

" 7, P- 7 


In dealing with the rust question from a practical point of view, there 
were two main issues to be determined: 

i How is the rust spread and continued from season to season? 
2*. How may its injurious effects be mitigated or counteracted or 
prevented ? 

The first question is a most important one, for if we could find out 
where and under what conditions, the rust is lying dormant during the 
time from reaping the crop to sowing it again, then we might be able 
destrov it at this stage, and prevent its reappearance. Although the 
question may thus be simply stated, it is by no means easy to answer. 

The second subject of prevention or mitigation will evidently depend 
on our knowledge of the life-history of the rust fungus, as well as of the 
wheat plant itself, and how far the conditions can be controlled which 
render it susceptible to the fungus. 

How the Rust is Spread and Continued from Year to Year. We 
know exactly now, thanks to the labours of Eriksson, Marshall 
Ward, and others, how the, rust-spores enter the plant by means of their 
germ-tulbes, how they grow and ramify among the tissues, and drain them 
of their contents, until they again form a spore-layer, and reproduce the 
spores on the surface in great abundance. We thus know how fresh 
spores orginate once they have got a start; but it is the starting-point 
which is the difficulty. 

Although the rust was known, and 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, writing about it in his Horse-hoeing Husbandry, attributes it to the 
attacks of small insects " brought, some think, by the East wind, which 
feed upon the wheat, leaving their excreta as black spots upon the straw, 
as is shown by the microscope." In 1767, its true nature as a fungus, and 
therefore as a plant, was determined by Felice Fontana, and in 1797, 
Persoon gave it the name by which it is still known, Puccinia graminis. 
The rust then is a fungus growing inside the wheat-plant, and living at 
its expense, and reproducing itself by means of minute seed-like bodies 
or spores, which are so conspicuous on the leaves and stem of the wheat 
at certain seasons. 

For a long time there was a suspicion in the minds of many practical 
farmers that the barberry bush- had something to do with its spread', and 
so firmly was this believed in, that the State of Massachusetts passed an Act 
compelling the inhabitants to extirpate barberry bushes. And when De 
Bary, in 1864, justified the farmer, and proved scientifically that there 
was a connexion between the fungus which appears on the barberry 
bush, and that which appears on the wheat, then it was thought by 
many that we had reached the root of the matter, and that we had simply 
to destroy the barberry bush in order to get rid of the rust But it is well 

Wheat Rust in Australia. 69 

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 uredospores 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 " 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. Cobb 11 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 ithat 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. Molineux 1 , 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 j 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 la/rgely made 

^ Wheat Rust in Australia. 

i 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 Carrying over the rust from season to season, it has been suggested that 
infi-tion may be communicated to the wheat from other grasses which, as 
ave elsewhere shown, may also be attacked by Puccima gramims 
Klebahn 1 (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. 

c. 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 sporidiola, 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 said to germinate upon 
barberry leaves, and produce the aecidial form of the rustand it may be 
that here they are simply dying out are becoming functionless, because 
the barberry 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 
although 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, firom 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 mycoplasm 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 (Uromyces 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. 71 


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 themost 
favorable conditions for its healthy development, and thereby render the 
rust less injurious. But, as a matter of fact, when it was experimentally 
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 

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 Department 
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 

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- 
mental 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- 

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 

72 Wheat Rust in Australia. 

the rust. At Port Fairy, in Victoria, where rotation is regularly prac- 
tised, and where I have had crops grown the same st^ason, sometimes 
after mangels, or potatoes, or onions, there was no perceptible difference 
in rustiness from those crops of wheat grown in succession. From the 
stand-point of good farming, it is to be recommended ; but cropping year 
after year does not seem to affect the liability to rust further than this, 
that self-sown wheat is likely to appear in the interval between the two 
crops. Such "volunteer" wheat, if not killed in cultivating, is much 
more forward than sown grain, and on account of its earliness may escape 
the rust, but on the other hand, there is a risk attending it, for it is some- 
times more rusty than the ordinary wheat, and appears earlier, so that 
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. Professor Lowrie, speaking from a large experience, in South Aus- 
tralia, considers that the apparent exemption which fallow-land sometimes 
enjoys from rust is due to the fact that it is customary to sow bare f allow - 
land first, and the crop ripens sufficiently early to escape the rust in some 

Burning Stubble. It is sometimes recommended to burn the stubble 
in order to destroy any spores of rust that may be about ; but as a matter 
of experience, this has not been found to produce any appreciable differ- 
ence. Theoretically, the burning of the surface of the soil should destroy 
a number of spores, and to that extenti is beneficial ; but, practically, the 
difference in result is not marked. As far as most wheat-soils are con- 
cerned here, the loss of vegetable matter is much more serious than any 
possible increase of rust. 

Soils. Soils are sometimes said to be rust-liable and rust-free, like the 
wheat itself; but on closer investigation, it will be found that the soil 
is only one of several factors that require to be taken into consideration. 
Rich soils are said to surfer most, and in rusty years the best crops have 
sometimes been raised from the poorest soils. On the rich soils the crop 
is naturally heavier and more luxuriant than on poor soils, and the softer 
and more susceptible tissues are easily invaded by the parasite. Hence 
it is that the rust may sometimes be worst in the best crops. The mallee, 
in Victoria, vis said to be particularly rust-free ; but in some seasons the 
crop has to be cut for hay, in order to save it from being completely 
destroyed by rust. The reason for general freedom from rust lies not 
so much in the soil as in the light rainfall and the early ripening of the 
crop. But if " muggy " weather should prevail when the wheat is flower- 
ing, then the crop is just as liable to rust here as elsewhere. At Port Fairy 
the low-lying black soils more readily produce a rusty crop than the sandy 
soils, and this may be partly due to the fact that the dark-coloured soils 
absorb more heat, and are more likely to preserve the spores in a fit state 
for germination, or because they grow a heavier crop, which often lodges 
and thus encourages the rust. 

Manures. Perhaps no means for the prevention of rust have been 
more thoroughly and continuously tested than the application of different 
manures. In all the States, as well as in New Zealand, experiments 
have been conducted to this end, and the general opinion has been expressed 
by a late Director of Lincoln College, New Zealand: " No manure has 
yet been discovered that is a preventative of rust in cereal crops." The 
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; 
till, it was generally found that nitrogenous manures favoured the rust, 
while phosphatic ones had a tendency to diminish it. Nitrogenous manures 

\\heat Rust in Australia. 73 

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, and' thus enables the crop to escape the rust to a certain 

Treatment of Seed. This was thoroughly gone into, since ft 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 wheat 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 rust, and Dr. Cobb 10 
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 been 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 cam succumb to another in a different country, and this has 
really 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 

74 Wheat Rust in Australia. 

of the world (Europe, Asia, Africa, and America), it has 
been proved that no wheat is absolutely rust-proof, that 
to say, there is no wheat known which will be proof against the rust 
when grown under conditions favorable to its development. But the same 
experiments have shown that among a number of varieties grown in a 
particular district, some will resist the rust more than others, are able by 
their constitution it may be by their vigour of growth or toughness of 
their cuticle or glaucousness, or all combined to resist the inroads of the 
rust sufficiently to prevent its seriously injuring the grain, and such wheats 
are known as 'rust-resisting wheats. At present we have at 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 the 
rust. Rerraf is the wheat referred to, and perhaps it, too, in course of 
time, will succumb to the rust. 

For this great and burning rust-question the only measures I can 
suggest are : 

1. To produce wheats suited to our Australian conditions by crossing, 
as Mr. Farrer, Wheat Experimentalist of New South Wales, is now so 
successfully doing. 

2. To select and carefully cultivate the most rust-resisting plants from 
these and other wheats having the necessary hardy, prolific, grain-holding, 
and milling qualities, keeping up the strain and constantly renewing their 
constitutional vigour. 

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 

In judicious crossing and careful selecting lies the solution of the great 
rust problem, as far as our present knowledge goes, a)nd 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 80 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, says that " rust sometimes 
appears, but it is not very common." J. 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. 

NOTE i. 

P. 4. Eriksson 18 , in his latest work on the vegetative life of Puccima 
graminis, has described and figured a transition from the mycoplasm to the 
protomycelial stage, but it still remains to be seen how far his interpretation 
of the facts is borne out bv other observers. 

Wheat Rust in Australia. 75 

NOTE 2. 

P. 14. Christman 1 , 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 fusie. There is thus true sexual cell fusion with- 
out the intervention of spermatia. 

NOTE 3. 

P. 24. Arthur 8 considers the sorus in such species as Puccinia bromina 
and P. triticina to be compound and the modified hyphae which separate 
the individual sori to form a stroma, hence they are not paraphyses, strictly 

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. Beauverie 1 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 usedl 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 uredospores 
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. * 



Classification. 79 



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- 


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

The collective species known as Puccinia rubigo-vera (DC.) Wint. was 
first divided in 1894 by Eriksson ,and Kenning 1 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 forms were^:- 
i, secalis; 2, tritici ; 3, bromi ; 4, agropyri. Further investigation, 
however, led him in 1899 to raise the various biologic forms to the rank < 
independent species, and it will be interesting from our present stand- 
point to consider the reasons given by Eriksson for differentiating between 



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, on the following grounds: 

1. The uredo is only able to infect the host-plant, Secale. 

2. This form regularly appears several weeks earlier than that on 
wheat, even when the host-plants are growing alongside of each other. 

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

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 
slightly 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. hieradi (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 1 
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 (they 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 

It will be observed that only 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 formerly, where they were hitherto unsuspected 
and overlooked. 

The three types selected for illustration will show the spirit of our 

P. graminis is the type of a single species which is split up into a number 
of biologic forms, agreeing in the faot of producing aecidia on the bar- 
berry, and the aecidiospores in turn infect the grasses. 

P. dispersa, as originally constituted, is the type of a collective species 
at first divided into biologic forms, which were afterwards found *to be 
distinct species, both on morphologic and biologic grounds. And the 
original P. hieradi is the type of a collective species, the members of which 
Strictly limited in their infective power to single genera, and their 
main title to distinction as species is this biological peculiarity. 

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

the future not only to include under that name such forms as possess 

:t morphological characters, but also those which, although morpho- 

similar, are yet confined to definite host-plants. It will thus be 

necessary to split up the old species of P. hieradi into several of equal rank, 

the one growing on Hieracium spp. will form a type round which the 

others may be grouped. Just as in the old species there were differences of 

opinion as to the amount of structural differentiation necessary to separate 

Classification. 81 

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 
biologic forms. lit 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 
species, the physiological differences at present existing becoming ultimately 
associated with morphological distinctions, which will mark them off as true 
species. It is not easy to explain how this has come about, but we 
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's 4 view, and Klebahn 1 , 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 direction 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. 

82 Systematic Arrangement. 



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 wihich 
mark out for us that particular object from all others." fT-he 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 described 1 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 Pflanzenfamilien: 

Order UREDINEAE, Tul. 

Fungi parasitic on higher plants and developing in the interior of their 
hosts, a filiform, branching, septate mycelium. Spores arising terminally 
or laterally from erect, transversely divided, crowded hyphae, and usually 
of more than one kind. Teleutospores germinating by a short promy- 
celium. The order may be grouped in the following four families, of which 
the third is not represented in Australia: 

Teleutospores stalked, single, in groups or rows, or several cells in a 
head. Fam i. Pucciniaceae. 

Teleutospores sessile, in columnar or filiform masses. Fam. 2. Cro- 

Teleutospores sessile or stalked, in one or two-layered waxy masses. 
Fam. 3. Coleosporiaceae. 

Teleutospores sessile, in flattened one-layered masses, or loose in the 
tissues of their host. Fam. 4. Melampsoraceae. 

On account of their economic importance, the Graminaceous rusts are 
always treated first and the others are likewise grouped together under 
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. 


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

be treated at greater length than any of the others, and it will be 

nterestmg to trace (the different forms which the teleutospore assumes in 

the different genera. There are a number of genera based upon very slight 

ifTerentiating characters, and it is not always easy to settle whether they 

etamed or rejected, but I have given all those which are clearly 

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

lould be retained, where the teleutospore is two-celled, as in Puccima 

only the septum is longitudinal instead of transverse. There are several 

j of Puccima in which there is a tendency to an oblique, and even a 

longitudinal septum, as well as transverse, but this genus is retained for 

those species in which the spore is distinctly divided longitudinally with a 

single germ-pore at the apex of each. 

Systematic Arrangement. 83 

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 P.ucciniaceae 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 1 therefore the Uromyces 
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 
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. Uromyces, Link. Teleutospore one-celled, with single germ-pore, 

and solitary on its stalk. 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 


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, FIG. IP. FIG 20. 

8. Gymno sporangium, Hedw. Teleutospore two-celled (rarely three 

to five celled), and walls which ultimately form a common gela- 
tinous mass, with usually several germ-pores in each cell. Fig. 21. 

9. Hapalophragmium, Syd. Teleutosp :>re three celled, and consisting 

of two basal cells alongside of each other, with a third on top. 

Fig. 22. 

10. Triphragmimm, Link. Teleutospore three-celled, and consisting of 
basal cell supporting two others alongside of each other. Fig. 23. 


, , Phragmopyxis, Diet. Teleutospore three-celled in a longitudinal 
series, with outer layer swelling when moistened. Pig. 24 ^ 
12 Phrazmidium* Link. Teleutospore three or more celled in a longi- 

O i _^11* ___ ___ 1-n-v *-*-*^-i r*4- r\-r\rf-\ 

tudinal series, 
Fig. 25. 

and outer laver not swelling when moistened. 


FIJ. 21. 

ri... -1-2. 

FIG. 23. 

FIG. 25. 

13. Sphaerophragmium, Magn. Teleutospore consisting of a spherical 
head of four to nine cells, and arising from a single cell by longi- 
tudinal and transverse fission. Fig. 26. 

Anthomyces, Diet. Teleutospores forming a head of ^ three to 
eight cells, barne on a single stalk, and arising from a simple cell 
by the formation of longitudinal septa. Several small sterile cells 
at base. Fig. 27. 

Ravenelia, Berk. Teleutospores forming a more or less hemispheri- 
cal, many-celled head, transversely and longitudinally septate, 
arising from a stalk compounded of several hyphae, and with 
several sterile cells or cvsts at the base. Fig. 28. 

FIG. 27. 

FIG. 28. 

Only four of these genera occur in Australia, and they will be dealt 
with in the following order: Uromyces, Uromycladium, Puccinia, Phrag- 


In this genus the teleutospores are unicellular as well as the uredospores, 
and this has sometimes caused the one to be mistaken for the other, but the 
single germ-pore in the former serves ito distinguish it. Besides, the uredo- 
spores are always echinulate or warted, and while the teleutospores are 
usually smooth, they may also be striated or warted, but the generally 
greater thickening of the wall, and the fact that the contents are nearly 
colourless, also serve to separate them. All the different spore-forms may 
be present in the same host-plant, or they may be reduced to the teleutospore 
alone. In clover rust (U . trifolii} and beet rust (U. betae) all the spore- 
forms occur, but in the latter, although the rust is very common, I have 
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. 

Uromyces Gramineae. 85 

In carnation rust (U. caryophyllinus) 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, puccinioides, the uredospores being unknown, and teleutospores alone are 
found in U. bulbinis and U. di-ploglottidis. 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 pseuclo- 
peiidium; aecidiospores without distinct germ-pores. 

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

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

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

Australian species, 27. 



1. Uromyces danthoniae Me Alp. 

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 p in diam. ; 
pseudoperidial cells persistent, elongated and polygonal, with 
striated margin. 

Aecidiospores subglobose to polygonal, orange, average 1 6 jj. 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 /i. 

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 p, average 33 x 22 p ; pedicels elongated, persistent, 
slightly tinted adjoining spore, up to 96 ju long. 

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

Victoria Bacchus Marsh, I. Near Melbourne, Ardmona, Portland, 
Killara, Leongatha, Rutherglen, N"agambie, Kergunyah, fcc., 

Tasmania Domain, Hobart, Nov., 1894, II., III. (Rod way). 

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

Puccinia gramindla (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. 

86 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 jw." 

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. Br., &c., 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. 

Darlucafilum Cast., is common on uredosori. 

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


2. Uromyces ehrhartae McAlp. 

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 
scattered germ-pores on one face, 21-25 x 18-20 u, average 
'24 x 19 fi. 

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

Teleutospores pale chocolate brown to chestnut brown, only 
occasional uredospores intermixed, hooded and thickened at apex 
(8-9 p), and either conical or truncate, very variable in shape 
and size, elongated or short and stout, subglobose to elliptic, 
fusiform or oblong, 19-32 x 13-19 p, average 26 X 15 /i ; 
pedicels persistent, tinted, elongated, up to 45 p long. 
On Ehrharta stipoide* Labill. = Microlaena stipoides R. Br. 

Victoria Near Melbourne, Killara, Myrniong, Kergunyah. fcc., 

Uredospores common but teleutospores rather sparse, although in the 

eighbourhood of Melbourne during December and January the teleuto- 

jometimes rather common, usually on the lower withered leaves, 

rmixed with uredosori on those still partially green. Only the 

uredo-stage was found at first by Mr. Robinson, but latterly at Killara, in 

Kergunyah, in November, he secured the teleutospores. 
Darlucafilum Cast., is very common on uredosori. 

(Plate XVI., Fig. 13i>.) 

Uromyces Gramineae, Liliaceae. 87 


3. Uromyces tenuicutis McAlp. 

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 p, average 32 x 22 p. 

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

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 p ; pedicel sometimes persistent, usually 
tinted, commonly 20-35 p 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. 

Darlucafilum Cast., is very commonly associated with the uredospores. 
(Plate XVI., Fig. 133.) 



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, 

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 p, ; pedicel 
deciduous, unequal, straight or slightly curved, hyaline to yellowish 
brown, 12 x 4ju. 

On living leaves of Bulbine bulbosa Haw. 
Victoria Omeo (Morrison). 
New South Wales Upper Macquarie River. 
Specimen not seen. 


5. Uromyces tricorynes Me Alp. 

McAlpine, Agr. Gaz. N.S.W., VL, 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 17/;. 

88 Uromyces Orcliidaceae. 

\\. 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 p diam. or 22-25 x 19-20 p. 

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 p), round or conical, 25-32 x 20-29 p, average 
30 x 25 p ; pedicels persistent, pale yellow, especially towards 
spore, up to 80 p long and 9 p 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 
at septum, which might be about the middle or towards the base. They 
were longer than the ordinary spore, and measured about 37 x 27 p. The 
pedicel was occasionally rather lateral. 

Darluca filum Cast., occurred on the uredosori. 

(PlateXVI, Figs. 134, 135.) 


6. Uromyces microtidis Cooke. 

Cooke, Grev. XI V., 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, puncti- 
form, brown, pulverulent, girt by the ruptured epidermis. 

II. Uredospores globose, subglobose, ovate or ellipsoid, minutely acule- 

ate to warty, yellowish-brown, 22-30 x 17-25 p. 

III. Teleutospores intermixed with Uredospores, ovate, with hyaline 
apiculus, rough with warte, brown, 25-35 x 17-25 p, but mav reach 

1 p t average 32 x 24 p pedicel very short, hyaline. 
On living leaves of Microtis porrifolia R. Br. 

New South Wales Bullahdelah, II., III. 

Uromyces Orchidaceae. 89 

Only the teleutospores were described by Cooke, but in a portion of the 
original material kindly supplied by Massee there were numerous uredospores 
intermixed with the teleutospores. Sydow 2 has also found the uredospores 
with teleutospores intermixed, on specimens of the same host-plant sent from 
Chatham Island, New Zealand. 

If we compare the three rusts found on the Orchid genera, Microtis, 
Chiloglottis, and Thelymitra, it is found that aecidia occur on Chiloglottisj 
but are absent, as far as known, from the other two. The teleutospores are 
all provided with a hyaline apiculus, but in U. microtidis the epispore is very 
rough and knobby, and the size of the spore not generally exceeding 35 p. 
long, while in U. thelymitrae the epispore io generally thicker, and the spore 
polygonal and irregularly warted and altogether larger. The rust on Chtf.o- 
ylottis generally resembles that on Microtis, but the teleutospore is larger. 

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





Uromyces microtidis 

25-35 x 17- 25 p 

Generally prominently 

3-4 p 


warted all over 


U. orchiclearum 

30-50 X 19-24 p 

Irregularly warted and 

3-4 ft 


warts not so pro- 



U, thelymitrae 

35-45 x 25-30 fi 

Less closely but more 

5-6 p 


prominently warted, 


hence polygonal 

(Plate XVI., Fig. 139.) 


7. Uromyces orchidearum Cke. and Mass. 

Cooke and Massee, Grev. XVI., p. 74 (1888). 
Cooke, Handb. Austr. Fung., p. 332 (1892). 
Morrison, Viet. 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 p. or 18-22 x 15-20 p. 

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

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

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

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 p long), usually with irregularly 
warted epispore, which is 3-4 p thick and brightly coloured, 
30-51 x 19-24 p, average 36 x 21 p ; pedicel hyaline, attenuated 
downwards, up to 77 p long. 

D 2 

9 o 

Uromyces Orchidaceac, Scroplndariaceae. 

On leaves of Chiloylottis diphylla 11. Br., II., III. 
New South Wales Mt. Victoria (Hamilton). 
Victoria Oakleigh, July, 1894 (Morrison 1 ). 
Tasmania Bellerive Swamp, May, 1897 (Rod way l ). 

On C. (junnii Lindl., I., II., III. 

Victoria Oakleigh, Aug., 1892 (Morrison). 

In the original description of Cooke and Massee only teleutospores are 
recorded, but uredospores were also obtained from the original material 
labelled in Cooke's handwriting. 

Sometimes there are abnormal elongated teleutospores up to 60 JJL long. 

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

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


8. Uromyces thelymitrae McAlp. 

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 p, average 36 x 26 p pedicel hyaline, short, 
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 Thelymitra 
javanica Blame, but it has probably no connexion with our native species. 

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



9. Uromyces limosellae Ludw. 

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

I. Aecidia amphigenous, scattered or gregarious ; pseudoperidia with 
whitish lacmiate margin, not deeply incised. 
^Aecidiospores round to angular, hyaline, smooth, about 15 p 

III. Teleutosori mixed with the aecidia, often crowded, long covered 
by epidermis, pulvinate, dark brown. 

Uromyces Goodeniaceae, Rubiaceae. 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 p 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 
a spore layer. 

Specimen not seen. 


Selliera, Scaevola. 
it). Uromyces puccinioides Berk, and F.v.M. 

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^u. 
III. Teleutosori bullate, intermixed with aecidia or surrounding them 
as a ring, at first covered with greyish glistening epidermis, finally 

Teleutospores brown, apiculate, sometimes with the apex oblique 
or dentate, also thickened and rounded or flattened, 40-44 x 
18-23 p, average 41 x 22 ^ ; pedicels hyaline, persistent, up to 
48 p 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 (R/obinson). 

South Australia Glenelg (Holdfast Bay), 1854 (Berkeley 2 ). 

Tasmania Bellerive Swamp, Dec., 1890, and May, 1897 (Rod- 
way a ). 
On Scaevola sp. 

Queensland St. George (Wedd) (Bailey 13 ). 

It is recorded in Cooke's Handbook as being found on Goodenia, but there 
are no certain indications of this. It occurred on Goodenia herpystica 
Schlecht., which, however, is a synonym of Selliera radicans. 

(Plate XVII., Fig. 140.) 



11. Uromyces asperulae Me Alp. 

McAlpine, Agr. Gaz., N.S.W., VI., p. 851 (1895). 
Sacc. Syll. XIV., p. 276 (1899). 

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 u. 

92 Uromyces Leguminosae. 

III. Teleutosori on stems towards their base, elongated oval, dark- 
brown, appearing almost black, confluent, erumpent. 

Teleutospores at first intermixed with uredospores, yellowish- 
brown to chestnut-brown, smooth, oval to ovate, or elliptical to 
broadly clavate, apex much thickened (11 p) and rounded, some- 
times tapering or flattened, 27-37 x 15-19ju, average 30 x IS p ; 
pedicels long, persistent, hyaline, up to 45 ^ long. 

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-plant is given as Asperula scoparia Hook, f., in the Index 
Kewcnsis, but the synonymy seems a little confused. A. oligantha F.v.M. 
is first given as Rubia syrticola Miq., then R. syrticola is given as a 
synonym of Asperula scoparia Hook. f. 

(Plate XVII., Fig. HI.) 



12. Uromyces appendiculatus (Pers.) Link. 

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

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

[I. 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 p.] 

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 p. ; 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 p, while in 
the other the thickening is not perceptible. 

(Plate XLII., Fig. 306.) 

Uromyces Leguminosae. 93 


13. Uromyces bicinctus Me Alp. 

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 //, occasionally reaching a length of 45 /z, and variable in 
breadth, average 34 x 15 p pedicel similarly coloured to spore or 

III. Teleutospores intermixed with uredospores, pallid but thickened 
considerably at apex and brown (8-11 //), smooth, ellipsoid, and rounded at 
both ends, occasionally prolonged into one or two processes at apex, 28-34 

x 17-23 /t; pedicel persistent, hyaline, short. 

On phyllodes and pods of Acacia f asciculif era F.v.M. 

Queensland Rockhampton, 1867 (from host-plant in National 
Herbarium, Melbourne). 

This species approaches somewhat to U. phyllodiorum (B. and Br.), 
Me Alp., 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 

(Plate XLIIL, Fig. 318.) 


14. Uromyces fabae (Pers.) De Bary. 

De Bary, Ann. Sci. Nat. Ser. 4, XX. (1863). 
Sacc. Syll. VII., 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, flat, with torn white edges. 

Aecidiospores subglobose, orange, finely echinulate, 16-26 /t 

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 yu. 

[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 p), and rounded, 
truncated or conical, sometimes with colorless papilla, 24-47 x 
17-30 ju; pedicels hyaline or pale brown towards spore, persistent, 
up to 110 /z long.] 

On stems, leaves, and pods of Broad Bean (Vicia faba L.). 
New South Wales. 1894 (Cobb 9 ). 
Queensland. Gladfield (Gwyther) (Bailey 19 ). 


Uromyces Legmninosae. 

This rust has not been found in Victoria, and although it is said to 
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 


(Plate XLIL, Fig. 307.) 

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 

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 p, average 62 x 23 p. 

III. Teleutospores intermixed with uredospores, ruddy brown, globose 
to depressed globose, epispore rather thin, scarcely 2 p thick,, 
somewhat polygonal seen from above and then germ-pore very 
prominent, 25-30 p diam. or 16-21 X 25-35 //, average 18 x 28 p ; 
pedicels deciduous, hyaline, elongated, up to 96 p long, with/ 
septum at a short distance beneath spore. 

On phyllodes of Acacia salicina Lindl., and A. neriifolia A. Cunn. = 
A. retinodes Schlecht. 

Victoria Dimboola, Dec., 1895, and May, 1897 (Reader). 
New South Wales 1902 (Maiden). 

On Acacia sp. 

Queensland Islands of Torres Straits, June, 1897 (Bailey 18 ' 19 ). 

Portion of the original material named in Cooke's handwriting is in the 
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 wil 
a colourless vesicle or cyst produced laterally immediately beneath septum. 

The nature of the teleutospore and the presence of a septum in the stall 
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. 

(Plate XIX., Figs. 158-160.) 

16. Uromyces hardenbergiae McAlp. 


Sori on under surface of leaf, brown, crowded, globose to ellipsoid, often 
confluent, bullate, with ruptured epidermis, which usually remains. 
in large patches. 

Uromyces Leguminosae. 95 

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 /*, 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 //, average 26 x 19 p ; pedicel persistent, hyaline, 
elongated, up to 35 // long. 

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 

This species persists all the year round. 
Darluca filum Cast., very prevalent on uredosori. 

(Plate XVII., Figs. 143-145; Plate G., Fig. 34.) 


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. XVII., p. 70 


O. Spermogonia brown at first, becoming shining black, numerous, 
small, blister like, seated in centre of swollen, brownish, scattered, 
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 /z, 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 /z, but may even reach a length of 86 p ; 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 p. 

Uromyces Legnminosae. 

On phyllodes of Acacia notabilis F.v.M. 

S. Australia Near Gawler, July, 1885 (Tepper) (Ludwig 2 ). 
On phyllodes of Acacia dallachiana F.v.M., and A. penninervis Sieber, 

Victoria 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 Benth. and A. 
neriifolia A. Cunn. in National Herbarium, Melbourne. 

New South Wales Twofold Bay. 

Queensland Brisbane River. 
On phyllodes of Acacia pruinosa A. Cunn. 

New South Wales Gosford, Jan., 1906 (Froggatt). 

On Acacia sp. 

Queensland Brookfield, Brisbane River (Bailey 1 ' 19 ). 
New South Wales (Cobb 10 ). 

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 
but there are occasional elongated forms which may reach a length of 54 
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 f roi 
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 phi/llodiorum 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 arising from delicate filaments, 
rather fusiform, 55-58 p long, granulated, mixed with others which are 
elongated, uniseptate, fusiform, even, 22 fj. long." The longest uredospore 
found was 54 p long, and the uniseptate spores of Darluca filum Cast., wei 
a No very common. 

Next, Winter, in 1886, described the same fungus from S. Australia as 
Uromyoeu digitatus, the teleutospores being recorded for the first time. 

Then Cooke and Massee, in 1889, described a Uromyces phyllodiae f rom 
Brisbane as follows : " Sori minute, orbicular, compact, brown, crowded on 
<-lli).tir,lnill;ite, brown spots, 3-5 mm. long, at length naked, not pulverulent. 
Teleutospores elliptic, obtuse, rarely apiculate, 

Uredosporea not seen. 

brown ; epispore minutely warted, rather thick, hyaline, thickened at th 
apex, 40-45 x 16- IS /.-'' A portion of the original material ^as supplied 
by Mr. Bailey, and both uredospores and teleutospores have been found. 

Uromyces Leguminosae. 97 

Cooke and Massee mistook the uredospores for teleutospores, but the 
existence of several gerin-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 Afelampsora 
phyllodiorum, and gives a drawing of the uredospores with an average size of 
41 x 17 fi. He also found the perithecia of Darluca filum 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, I., p. 159 (1884). 
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 p, 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 ^ diam., or 22-25 x 14-16 yu. 

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/i. 

III. Teleutosori smaller and rounded on the leaves, larger and elon- 
gated on the petioles, dark brown, almost black, bullate, long 
covered bv 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 p, average 24 X 20 yu; 
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., L, II., III. Murramurrangbong 
Ranges, Jan., L, II., III. 

New South Wales (Cobb 12 ). 
Tasmania (Rod way 1 ). 

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 z considers this a common 

(Plate XVIL, Fig. 142; Plate G., Fig. 32.) 

9 g Uromyces Polygonaceae. 



11). Uromyces politus (Berk.) Me Alp. 

Berkeley, Linn. Journ. XIII., p. 174 (1872). 

Berkeley and Broome, Linn. Trans. II., p. 67 (1883). 

Cooke, Handb. Austr. Fung., p. 342 (1892). 

Sacc. Syll. VII., 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 
cells oblong to angular, with striated margin, 32-35 p long. 

Aecidiospores ochraceous, variable in size and shape, generally 
ellipsoid to ovoid or oblong, finely echinulate, 24-30x19-22^. 

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 ^u), 
occasionally two-celled, variable in size, 28-40 x 20-25 p, average 
32 X 24 p ; pedicel hyaline, persistent, elongated up to 150 p. 

On stems and branches of Muehlenbeckia cunninghami, F.v.M. 

New South Wales Pamamero Lake, Nov., 1860 (Berkeley 2 ). 
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 tl 

Berkeley 2 first recorded the aecidial stage of this species on Muehlenbeckia 
cunninghami in the Journal of the Linnean Society XIII., 174 (1872), 
and at the same time Cronartium asdepiadeum was noted on Jacksonia 
scoparia from the Darling Downs, Queensland. 

Then just ten years later, in the transactions of the same society for 1882, 
1. 67, R. polita is recorded on the latter plant from Brisbane (Berkeley and 

I Iroome 2 ), and it was rather peculiar, to say the least of it, that the same 
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 

II 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 determination 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. 99 

which Cronartium \tas first determined, and Mr. Bailey has sent me a speci- 
men from his herbarium labelled " Roestelia polita Berk., " and which was so 
named by Berkeley. On comparing the Brisbane specimen with the Darling 
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 011 Muehlenbeckia and not on Jacksonia it 
will reconcile certain discrepancies in Berkeley's original descriptions and 
drawings. In the original description of JR. polita the spores are given as 
large, while in connexion with its occurrence on Jacksoniait, is stated "the 
spores are globose about *0003 inch in diam. (7-7|- /u)." If we turn to the 
drawings on PI. ] 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 polygon! (Pers.) Fckl. 

Fuckel, Symb. Myc., p. 64 (1869). 

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 p 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 

Uredospores ellipsoid to obovate, yellowish brown, finely 
echinulate, with generally two lateral germ-pores on one face, 20-29 
x 16-21 p, average 24 x 20 p. 

III. Teleutosori scattered, blackish, pulvinate, roundish on leaves, 
elongated on stems, bursting through browned epidermis, confluent 
in masses, surrounding and distorting stem. 


Uromyces Chenopodiafeae. 

Teleutospores globose, subglobose or elliptfcal, smooth, bright 
chestnut brown, apex thickened (8 /*), rounded or occasionally 
somewhat pointed, 22-34 x 16--20 p, average 28 x 18 /* ; pedicels 
pale yellowish, persistent, firm, long, up to 90 p. 
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 
tdeutosori in May, 1904. 

(Plate XVIII., Figs. 150, 151.) 


A triplex. 

21. Uromyces atriplicis Me Alp. 

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 JJL. 

III. Teleutospores intermixed with uredospores, dark brown, sub-globose 
to shortly ellipsoid, finely striated longitudinally, slightly thick- 
ened at apex, with prominent single apical germ-pore, 22-29 x 
24-27 p, average 26 x 25 /z. 

On leaves of Atriplex semibaccata R. Br. 

Victoria Royal Park, near Melbourne, April, 1899. (Brittle- 

The groups of spores shown in the Figures are rather smaller than the 
average, being about 22 x 25 p. Occasionally an abnormal teleutospore 
occurred reaching a length of 37 ju. 

(Plate XVII., Figs. 146, 147.) 


22. Uromyces betae (Pers.) Kuehn. 

Kuehn, Bot. Zeit., p. 540 (1869). 

Cooke, Grev. XT., p. 98 (1883). 

Cooke, Handb. Austr. Fung., p. 330 (1892). 

Sacc. Syll. VII., p. 536 (1888). 

0. Spermogonia yellowish to honey-coloured, in small clusters, on both 

surfaces of leaf. 

Spermatia hyaline, ellipsoid, 5-6 x 3 p. 

1. Aecidia disposed on orbicular or oblong yellow spots, on both 

surfaces of leaf, mostly about mid-rib and on leaf stalk ; pseudo- 
peridia white, cup-shaped, with reflexed fringed margin ; peridial 

Uromyces Sapindaceae. lot 

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 /*, average 28 x 21 //. 

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 jj, average 32 x 21^; 
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 Faiiy 
in August and measured 27-33 x 22-24 //. 

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 

(Plate XVII., Figs. 148, 149 ; Plate XLIIL, Fig. 316; Plate H.) 



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 p. 
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. 

I02 Uromyces Caryophyllaceae. 



i 1 . Uromyces caryophyllinus (Schrank) Schroet. 
Schroeter, Brand, p. 10 (1869). 
McAlpine, Agr. Gaz. N.S.W. VII., p. 300 (1896). 
Sacc. Syll. VII., p. 545 (1888). 

II. Uredosori amphigenous, dark brown when exposed, but long 

covered by epidermis, round or oblong, scattered, often confluent, 
up to 3 mm. long. 

Uredospores roundish to oblong, yellowish brown to golden 
brown, decidedly echinulate, generally 3-4 scattered or nearly 
equatorial germ-pores on one face, 21-34 X 17-28 ju, average 
32 x 24 ,1. 

III. Teleutosori amphigenous, blackish-brown, oblong, confluent in elon- 
gated lines, long covered by ashy epidermis, then margined by the 
ruptured and ragged cuticle. 

Teleutospores at first intermixed with uredospores, roundish, 
oval or ovate, cinnamon brown, generally thickened, with broad 
colourless papilla, average 28 x 20 /t or 22-32 x 17-23 p ; 
pedicels very deciduous, hyaline, up to 45 ^ long. 

On stems and both surfaces of leaves of Dianthus caryophyllus L. and 
D. chinensis L. Carnation and Dianthus. 

Victoria Near Melbourne, and at Ardmona. Very common in 

some seasons all the year round. 

New South Wales Ashfield, near Sydney. (Maiden.) 
South Australia Feb., 1902. 
Queensland Roma, 1901 (Tryon 8 ). 

There is not much difference between the naked-eye characters of the 
uredo and teleuto sori, only the latter are a little darker in colour, but 
the spores themselves are quite distinct, for the uredospores are covered 
with decided spines, while the teleutospores are relatively smooth and have 
a clear papilla at apex. 

The teleutospore is filled with finely granular protoplasm, in the centre 
of which is a large vacuole-like body. When the spore is examined in the 
dry condition, the surface appears to be covered with numerous very fine 
points, which Fischer 5 regards as warts. The wall of the uredospore is beset 
with loosely arranged spines, which project at the margin when seen in 
optical section, while that of the teleutospore has densely crowded fine 
points, which are not visible at the margin. These projecting points are so 
fine that the spore is generally described as smooth. 

Darluca filum Cast, is often parasitic on the uredosori and teleutosori. 

(Plate XVIIL, Figs. 152-154; Plate G., Figs. 30, 31.) 


Uromyces scleranthi Rostr. 

Rostrup, Bot. Tidsskr., p. 40 (1897). 
Sacc. Syll. XIV., p. 275 (1899). 

II. Uredosori scattered, minute, orbicular or oblong, slightly raised, 
pale brown, erumpent, and surrounded by the ruptured 

Uromyces Zygophyllaceae. 103 

Uredospores golden brown, finely echinulate, ellipsoid to ovoid or 
oblong, 3-4 scattered germ-pores on one face, epispore 3 u thick 
25-32 x 17-21 /*. 

On Sderanthus 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 Sderanthus 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 ^ diam. or oblong, 24-25 X 18-20 p. 

Teleutospores rare, intermixed with uredospores, pear-shaped or obliquely 
ellipsoid, reddish-brown, apical papilla transversly thickened, 23-24 x 
19-24 p. ; pedicel hyaline, deciduous." 

(Plate XXVIII., Fig. 248.) 



26. Uromyces vesiculosus Wint. 

Winter, Hedw. p. 22 (1885). 

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 //, average 28 x 20 p. 

III. Teleutospores globose, elliptic, ovate or pear-shaped, thickened at 
the apex, rounded or with a broad apiculus more or less shortly 
conical, 21 /z diam., or 23-31 x 17-22 /z, average 25 X 20 p 
epispore thick, smooth, dark bay brown when mature; pedicel 
long, thick, persistent, may be flexuous, pale olivaceous, up to 
150 p. 

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 Dirnboola, 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. 

IC , Uromycladutm. 

Dietel observes that the occurrence of bicellular teleutospores is rare 
amoiiL' r/v,wyr/-x, for only once has he observed it in U. pisi and U. junci, 
and two were found in U.'limosel/ae. Dr. Cooke has also found a few in 
U. trifolii, and I have now to add U. orchidearum, U. tricorynes, U. vesi- 
culosus and U. politus. 

Darlucafilum Cast., is generally very plentiful on uredosori. 

(Plate XVIII., Fig. 155-157 ; Plate G., Fig. 33.) 


This new genus of Busts 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 
which seriously disfigure and ultimately destroy many of our Wattle-trees 

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- 
r rianum (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 
l>eing gradually and completely destroyed by this fungus. Some of the 
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 ssize 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 
the cuticle, without paraphyses at mouth, preceding the formation of any 
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 Leguminosae. 105 

This genus may be distinguished from Uromyces 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 witk Uromyces on the one hand and Ravenelia on the other. 

Australian species, 7. 

I. One teleutospore and vesicle in head. 

A. Uredospores fusiform, with hyaline apiculus, and finely 


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. 

Ur. bisporum. 

III. Two teleutospores and vesicle in head. 

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. 

Ur. alpinum. 

IV. Three teleutospores in head. 

A. Teleutospores finely warted, warts arranged in lines ; uredo- 

spores known. 

Ur. Jiotabile. 

B. Teleutospores with converging striae ; Uredospores unknown. 

Ur. tepperianum. 



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 /u. 

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 p. 

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 p ; vesicle arising from 
stalk immediately beneath septum, globose or slightly ellipsoid 
about 25 z diam. 

I0 <5 Uromycladium Leguminosae. 

X. Mesospores associated with uredospores, not uncommon, 
ellipsoid to oblong or obovate, rounded at apex, smooth-walled 
and wall of about equal thickness throughout, with colorless 
stalk, 15-25 x 10-15 ft. 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 daUachiana 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, 

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 sporophore. 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.) 

-*. Uromycladium bisporum Me Alp. 

III. Teleutosori on the branchlets forming elongated slightly swollen 
chocolate-brown masses, and on the under surface of the leaflets 
appearing as powdery patches. 

Teleutospores two in a head, yellowish-brown, subglobose , 
depressed globose, occasionally with very short, colored, stalk-likt, 
basal projection, slightly thickened at apex with germ-pore im- 
mediately beneath, 18-22 x 22-30 p. 

On branches, leaves and pod's of Acacia dealbata Link. 

Victoria Murramurrangbong Ranges, January, 1905 (Robinson). 
Tasmania 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 
with two spores and a vesicle in each head. The occasional presence of two 
teleutospores in U. simplex is a further indication of the passage from one to 
the other. 

(Plate XXIV., Figs. 207, 208 ; Plate XXXIII.) 

Uromycladium Leguminosae. 107 

29. 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 ^ diam. 

Spermatia hyaline, shortly ellipsoid, 3 x 2 p. 

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 ju ; 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 ^ diam., or 22-25 x 24-30 ju ; colorless 
vesicle globose, with very thin wall, and arising from stalk 
immediately beneath septum, 30-35 ju 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 /*, occasionally up to 57 JJL long. 
On phyllodes and stems of Acacia longifolia Willd. 
Victoria Saudringham 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.) 

jo8 Uromycladium Leguminosae. 

30. Uromycladium notabile (Ludw.) Me Alp. 

Ludwig, Bot. Centrbl. XLIII., p. 6 (1890). 
Cooke, Handb. Austr. Fung., p. 343 (1892). 
Sacc. Syll. XI., p. 222 (1895). 
Uredo notabilis Ludw. 

O. Spermogonia minute, punctiform, black, in swollen tubercles often 
intermixed with uredospores and teleutospores. 

Spermatia hyaline, ovate or ellipsoid, on elongated basidia, 4 x 
2-3 /i. 

II. Uredosori on both surfaces of phyllodes, on branches and pods, 

ochraceous to yellowish-brown, seated on a distorted inflated gall. 
Uredospores ellipsoid to oblong, bright yellow when fresh, 
becoming yellowish-brown, on elongated hyaline pedicels, with 
3-5 equatorial germ-pores on one face; epispore thick (3 p), reticu- 
late, 30-45 x 18-28 //. 

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, 
yellowish-brown, densely covered with warts arranged in lines, 
slightly thickened at apex, with germ-pore beneath, 16-23 x 
21-26 p. 

On phyllodes of Acacia notabilis F. v. M. 

South Australia Roseworthy, Sept., 1889, II. (Tepper). 
On branches, leaves and pods of Acacia dealbata Link. 

Victoria Murramurrangbong Ranges, Jan., 1905, O., II., III. 
(Robinson). Bright, June, 1905, II., III. (Davey). Bairns- 
dale and Orbost, Dec., 1905, III. 
Tasmania Hobart, May, 1905, II. (Rod way). Cataract Gorge 

and Dulverton, Jan., 1906, III. (Robinson). 
On branches of Acacia decurrens Willd. 

Victoria Near Melbourne, Feb., 1905, III. Myrniong, July, 

1905, III. 

New South Wales Exeter, near Moss Vale, May, 1905, III. 

On branches of Acacia elata A. Cunn. 

New South Wales Lawson on the Blue Mountains, April 1905, 

O., III. (Baker). 
On branches of A. binervata DC. 

New South Wales Sydney, Sept., 1905, III. (Maiden.) 
On branches of Acacia pruinosa A. Cunn. 

New South Wales Gosford, Jan., 1906, II. (Froggatt.) 
The ochraceous uredosori are generally distinct from the chocolate-brown 
teleutosori, but sometimes the two kinds of spores are found intermixed. 

The arrangement of the teleutospores is generally the same as in 
U. tepperianum, but the dense covering of warty spines instead of distinct 
striae differentiates them at once and the height of the spore is greater. The 
spermogonia are found in association with both uredospores and teleuto- 

The uredospores closely resemble in size and shape those of Uromyces 
phyllodwrum, but the surface markings serve to distinguish them. In the 

TJromycladium Leguminosae. 1 09 

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 indiam. 
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 
flourished 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. Dieter 5 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 Raveuelia, 
and, considering the relationship of this genus with Uromydadium, the 
suggestion turns out to be not far from the truth. 

(Plate XXIII., Figs. 196-205 ; Plate XXXVI.) 


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 p 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 p. 

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 p; vesicle hyaline, globose to shortly 
ellipsoid, often on distinct stalk with septum at base, 20-25 p diam. 
or 22-27 x 16-23 p. 

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 p. 
On phyllodes of Acacia melanoxylon R. 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 dingy 
appearance to the phyllodes on both surfaces, and sometimes the sori are so 
crowded as to convey the impression of a continuous mass of rust, or even 


Uromycladium Leguminosae. 

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., Fig. 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 p. 

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 p. 

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 n ; sporo- 
phore fitting into a sort of socket on base of spore, hyaline, elon- 
gated, 80 p. or longer; vesicle hyaline, globose 19-22 ^ 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., 

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. 

Darluca filum Cast, not uncommon on sori containing both uredo and 

(Plate XIX., Figs. 161-165; Plate XXXII., Fig. 275.) 

Uromycladium Leguminosae. 1 1 1 


33. Uromycladium tepperianum (Sacc.) McAlp. 

Saccardo, Hedw. XXVIII., p. 126 (1889). 
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 JJL diam. 

Spermatia hyaline, ellipsoid, 3-3 J x 2-2 J ju. 

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 p ; sporophore hyaline, elongated, soon 

On branches of Acacia salicina Lindl. ; A. hakeoides A. Cunn. ; A. 
myrtijolia 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 Lindl. ; A. verniciflua A. Cunn. ; A. verticillata Willd. 

Tasmania. Hobart, March- April, 1905 (Rodway and Lea). 
On Acacia lonyifolia Willd. 

New South Wales Rose Bay, near Sydney, July, 1905 (Froggatt). 
On branches and phyllodes of A. erioclada Benth., and A. glaucoptera 

West Australia From types of host in National Herbarium, 

On Acacia stricta Willd. 

Tasmania Gordon, D'Entrecasteaux Channel, Nov., 1905 (Rod- 

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 V. notabile of producing 
galls. This is particularly noticeable in the Golden Wattle (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 

, j 2 Puccini a. 

large numbers from the branches like so many fruits, and the trees are 
either dying or dead. (See Frontispiece.) 

The dark chocolate-brown spore-masses are quite powdery, and each spore 
has prominent ribs running from base to apex, so that it is easily known 
from being fluted. In fact, the appearance closely resembles trie markings 

the eggs of some butterflies, and no doubt the purpose is the same, to 
strengthen the membrane which is already relatively thick, and prevent 
the spore collapsing when dry conditions prevail. Occasionally a spore 
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 montana 
Benth., in Java, and it would be interesting to know if it occurred on any 
indigenous species of this genus in New South Wales, Queensland or West 

On A. implexa at Myrniong there were numerous galls, and I found one 
at the end of a branch in July somewhat of a leg-of-mutton shape and 
weighing about 3 Ibs. (PI. XLL). Witches' brooms of various sizes also 
occurred on A. implexa, caused by this rust, and one of the largest 
measured 45 inches in circumference (PL XLIL). 

(Plates XXIL, Figs. 190-195; XXIIL, Fig. 206; XXXIY.; XXXV.; 
XLL; XLIL, Fig. 305.) 


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, such 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. 

Aecidiospores originating in serial order and soon free, globose, sub- 
globose or angular, hyaline, yellowish or orange. 

Puccinia Gramineae. 113 

Uredosori when present, generally minute and flattened, sometimevS 

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- 

Teleutospores separate, variously shaped, pedicellate, 1 -septate, with 
one germ- pore in each cell. 

Sporidiola ovoid or reniform, generally hyaline. 

Australian species, 90. 


Agropyron, Clematis. 

34. Puccinia agropyri Ell. & Ev. 

Ellis and Everhart, Journ. Myc. VII., p. 131 (1892). 

Sydbw, 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 ^ 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 p. 

Aecidiospores orange-yellow, ellipsoid to sub-globose, finely 
echinulate, 21-29 X 16-19 ju, or 20-22 p 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 ju, 
epispore up to 2|- ^ 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 ^u, average 60 x 18 ^ ; upper cell rounded or 
squarely truncate at apex and decidedly thickened (6-9 /u), some- 
times broader than long, darker in colour than lower, 19-29 x 
16-25^; lower cell tapering towards pedicel or oblong, usually 
longer and narrower than upper, 25-41 x 12-20 /j, ; 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 ju, with short coloured pedicels. 
Aecidiospores on Clematis aristata R. Br. 

Victoria. Murramurrangbong Ranges, Dec., 1903, and Jan., 

1905 (Robinson). 
Uredo and teleutospores on Agropyron scci/irum Beauv. 

Victoria Near Melbourne, Dec., 1892 (Robinson). Myrniong, 
March, Nov., Dec., Jan. Murramurrangbong Ranges, 
Dec., 1903 (Robinson). 

1I4 Puccinia Gramineae. 

The teleutospores vary considerably in size and shape. They are some- 
times 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 from 40-70 /LI and the breadth from 12 p (in lower cell) to 25 /j, (in 
upper cell). It is noticeable that certain sori may consist of elongated and 
others of medium-sized spores. 

In the Agricultural Gazette for New South Wales, Vol. VI., p. 852, 
1895, the rust on this native grass was given as P. dispersa Eriks. and 
Henn., but on further examination I find that it approaches most closely to 
P. agropyri Ell. and Ev. and is named accordingly. 

Specimens of P. agropyrina Eriks., were examined from Eriks. Exs. 419 
on Agropyron repens Beauv., and the uredospores measured 21-24 /u diam. 
or 24^27 x 16-21 p. The teleutospores, however, were considerably 
smaller than our own material, measuring only 33-42 x 12-18 /u, average 
;J9 x 15 /LI. 

The following table will show at a glance the different sizes of the spores 
on different hosts of the two species of rust : 

Uredospore. Teleutospore. 

P. ajropyrina Eriks. on Agropyron repens, Beauv. 1 6-25 /Lt diam. 36-41 x 13-16 \i 

/'. agropyri Ell. & Ev. on A. glaucum, Roem & Schult. 20-25 x 18-22 p 60-75 x 20-25 /t 
P. ajropyriEll & Ev. on A. scabrum, Beauv. 25-32 x 19-22 p 40-70 x 12-25 fi 

Although the uredospores are described as smooth by Ellis and Everhart 
yet in specimens from Sydow's Tired. Exs. 1362, they are seen to be 
decidedly echinulate and it is a question whether such a thing as a smooth 
uredospore exists among Puccinias. The species altogether is of the P. 
dispersa type. Dietel 4 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 
Klebahn, 1 p. 292, Puccinia (clematidi) agropyri Ell. and Ev. It is suggestive 
that both stages have been found in the Murramurrangbong Ranges growing 
near to each other. 

(Plate III., Fig. 25.) 

35. Puccinia agrostidis Plow. 

Plowright, Grev. XXI., p. 110 (1893) and Gard. Chron., p. 


McAlpine, Agr. Gaz. N.S.W. VII., p. 149 (1896). 
Sydow, Mon. Ured. I., 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 /u. 

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 /j), truncate or rounded, markedly constricted, 
attenuated below, almost sessile, very shortly stalked, 40-54 x 
14-21 p, average 44 x 18 /u. 

Puccinia Gramineae. 115 

X. Mesospores fairly numerous, similarly coloured to teleutospores or 
lighter, oval to elongated elliptical, thickened at apex, 24-30 x 
12-14 p . 

On Deyeuxia forsteri Kunth. = Agrostis solandri, F.v.M. 

Victoria Near Melbourne, 1892 (Robinson). Ardmona, Oct.- 
Dec., 1894 (Robinson). Arthur's Creek, Aug., 1902 

New South Wales (Cobb 2 ). 

Through the kindness of Dr. Plowright, I received some of the original 
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 p, but the uredospores are rather 
larger, being 25-30 x 21-25 p. 

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.) 


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. VIL, 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 /*. 

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 p) at apex, occasionally 
tricellular, 28-48 X 15-21 p, average 35 X 20; upper cell 
usually darker than lower; pedicel persistent, tinted, 
20-25 x 6-7 p, sometimes reaching a breadth of 10 p. 
X. Mesospores occasional, similarly coloured to teleutospores, obovate, 

thickened at apex, average 35 x 17/7. 
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.) 

T 1 6 Puccinia Gramineae . 

37. Puccinia beckmanniae Me Alp. 

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 f.i. 

III. Teleutosori minute, black, linear, hypophyllous, long covered by 
epidermis, ultimately naked, about J 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 7) finger like processes, 
45-60 X 16-25 p, 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. 

The grass on which this rust was found was grown from seed sent 
the United States Department of Agriculture for trial, and the spoi 
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. 

Darluca filum Cast., was very plentiful, both by itself and in conjunction 
with the rust. Probably in the former case it was parasitic on the hidden 
mycelium, and may have largely prevented spore formation. 

(Plate IL, Fig. 12.) 

38. Puccinia bromina Eriks. 

Eriksson, Ann. Sci. Nat. IX., p. 271 (1899). 
Sydow, Mon. Ured. I., p. 712 (1903). 
Sacc. Syll. XYIL, p. 382 (1905). 

Puccinia dispersa, f. sp. bromi, Eriks. 

II. Uredosori 1-1 Omm. long, 1mm. broad, ferruginous, on leaf blade 

chiefly on upper surface, crowded, sometimes on sheath and 

Uredospores bright orange, ellipsoid, with numerous scattered 
gerni-pores, as many as 11 on one face, echinulate, 27-31 x 
18-24 p. 

III. Teleutosori elongated elliptical to oblong, black to dark-brown, 
hypophyllous, scattered or subgregarious, sometimes on sheath and 
panicle, covered by epidermis, divided into compartments, and 

Lch compartment separated by elongated, cylindric, or slightly 
clavate, yellowish-brown, barren filaments or paraphyses. 

Puccinia Gramineae. 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 /i, average 48 x 21 p lower cell 
generally narrower than and almost as dark as upper; pedicel 
short, hyaline to pale yellowish, up to 36 p 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 jl. 
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 (Cobb 2 ). 

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 p, thus being rather smaller 
than the above. The teleutospores were much paler in colour, and measured 
36-48 x 15-22/z, the largest being about the average of the Victorian 

Fritz-Muller 1 observed aecidia on Symphytum qfficinale 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.) 


39. Puccinia cacao Me Alp. 

Uredo rottboelliae Dietel, Engler's Bot. Jahrb. XXXII., p. 52 


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 echinulate f 
32-40 x 24-32 p, average 35 x 28 yu. 

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. 

! j g Puccinia Gramineae. 

On leaves and sheaths of Kottboellia compressa L. 

Victoria Creswick, Jan., 1893, II. (Robinson), Werribee Gorge, 
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. (Bailey 1 ?). 

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 p and 34-56 x 22-27 /z in size. 
The uredosori frequently look quite black owing to the presence of 
Darluca filum Cast. 

(Plate XXX., Figs. 259, 260.) 

40. Puccinia cynodontis Desm. 

Desmazieres Exsicc. III., No. 655. 

McAlpine, Agr. Gaz. N.S.W. VII., p. 150 (1896). 

Sydow, Mon. Ured. L, 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, 

Uredospores globose to shortly elliptical, yellowish-brown to 
brownish-yellow, delicately verrucose, 20-25 p diam., or 20-25 x 
17-22 p. 

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 p. 

X. Mesospores plentiful, dark-brown, ellipsoid, rounded and thickened 
at apex, 24 x 19 /t . 

On Cynodon dactylon Pers. 

Victoria Burnley, Caulfield, Armadale, Pakenham, Killara, 
Somerville, &c., 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. 

Puccinia Gramineae. 119-. 

It was likewise found by Magnus 5 that two kinds of uredospores 
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.) 


41. Puccinia festucae Plow. 

Plowright in Gard. Chron., p. 42 (1890) and Grev. XXL, p. 

109 (1893). 

Sydow, Mon. Ured. I., p. 752 (1903). 
Sacc. Syll. XL, p. 194 (1895). 

II. Uredosori 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- 

Uredospores sub-globose to ellipsoid, echinulate, yellowish to 
orange-yellow, up to 5 scattered germ-spores on one face, 20-25 x 
16-18 fi. 

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 ^u, often 60 X 16//; 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 ^ 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 11-14 p. 

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 IL, Fig. 13.) 


42. Puccinia flavescentis McAlp. 

McAlpine, Proc. Linn. Soc. N.S.W., XXVIIL, p. 558 (1903). 
Sacc. Syll. XVIL, p. 380 (1905). 

II. Uredosori on upper surface of leaf, minute, linear, often confluent, 
soon naked, p ulverulent, rusty brown, arranged along furrows of leaf. 
438. E 

I2 o Puccinia Gramincac. 

Uredospores globose to shortly elliptical, finely echinulate, 
golden-brown, with at least ."> uvnn-pores irregularly distributed, 
21 -24 p diam. or 25-32 x 21-24/i. 

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 ju, average 
44 x 24 /u ; 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 /i 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 fiavescens Labill. 

Victoria Near Melbourne, Dec.- April, II., III. 

On Stipa semibarbata R. 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. 

In specimens of P stipae (Op.) Hora, taken from Syd. Ured. Exs. No. 
23, on Stipa capillata L., the teleutospores are decidedly different. The 
apex is generally bluntly pointed, and the size 48-54 x 18-21 p. In speci- 
mens of P. stipae Arth., from Arthur and Holway's Ured. Exs. No. 27, on 
Stipa spartea Trin., the teleutospores are more pointed at the apex and 
rather thicker. 

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. 


Persoon, Disp. Meth., p. 39 (1797). 
Cooke, Handb. Austr. Fung., p. 335 (1892). 
Sydow, Mon. Ured. L, p. 692 (1903). 
Sacc. Syll. VIL, p. 622 (1888). 

II. Uredosori amphigenous, yellowish-brown, linear, 2-3 mm. ~ 
longer, either scattered or confluent in long streaks, especially on 
sheaths, pulverulent, soon naked, surrounded by ruptured epi- 

Puccinia Gramineae, 121 

Uredospores elongated ellipsoid to ovate oblong, brownish yellow, 
echinulate, generally with 3-4 equatorial germ-pores on one face, 
20-36 x 14-18 p. 

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 p, average 52 X 18 p ; upper cell rounded or pointed at 
apex, rarely truncate, considerably thickened (up to 12 p.), some- 
times as broad as long, 21-29 p 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 //. 

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 p ; 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 p, with elongated 

On Wheat (Triticum vulyare Yill.), Polish Wheat (T. polonicum L.), 
Oats (A vena sativa L.), Barley (HordeumvulgareL.),'Rje (Secale cerealelj.), 
and various species of the following genera of Grasses: Agropyron y 
Alopecurus, Amphibromus, Avena, Beckmannia, Briza, romus, Dactylis, 
Deyeuxia, Echinopogon, Elymus, Festuca, Gtyceria, Nordeum, Phalaris, Poa. 

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 p. in the oblong, dark-brown, perfectly mature 
spore to 63 ju, long, and sometimes only 14 p 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. Ptowngbt in Austra- 
lian specimens ; or the upper cell only may be pale in colour as shown in 
Plate L, 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 Oab (Avena fatua) and Glyceria 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 


Puccinia Gramineae. 

other authors, and a double asterisk indicates those which 
by Eriksson in Europe to produce aecidia on the Barberry 
were sown upon it : 

Agropyron divergens Nees 
*A. scabrum Beauv. 

Alopecurus geniculatus L. 
Amphibromus neesii Stead. 
*Avena t'atua L. 
**A. saliva L. 

Beckraannia 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 broraoides L. 

Glyceria dives F.v.M. ... 

G. stricta Hook.f. 
**Hordeum murinum L. 

H. secalinum Schreb. ... 
**H. vulgare L. 
**Phalaris canariensis L. 

P. minor Retz. 
**Secale cereale L. 

Triticum polonicum L. 
**T. vulgare Vill. 

have been found 
when their spores 

II., III. 
II., III. 

II., III. 
II., III. 
II., III. 
II., III. 

II., III. 
II., III. 
II., III. 
II., III. 

II.. III. 
II., III. 
II., III. 

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 various grasses 
are capable of infecting each other. 

(Plates I., Figs. 2, 5, 7, 8 ; XIV., Figs. 113-122 ; XL., Fig. 301 ; 
XLIIL, Fig.' 311 ; Plate A., Figs. 3, 4.) 

Gramineae. 123 


44. Puccinia impatientis (Schw.) Arthur. 

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, 1-1 J 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 

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 p, average 
40 X 16 IJL 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 p.. 

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 ^ and the teleutospores 45-100 ju long. 

P. montanensis Ellis also differs in the teleutospores being stouter and 
broader, sometimes reaching a breadth of 26 p. 

(Plate III., Fig. 26.) 

45. Puccinia lolii Niels. 

Nielsen in Ugeskrift for Landmaend. I., p. 549 (1875). 

Sydow, 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. 

j 24 Puccinia 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 p. ; 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, J-l mm. long. 
Teleutospores yellowish-brown to brown, elongated clavate, 
tapering towards base, smooth, constriction very slight or absent, 
sometimes unicellular, 35-60 x 12-20 p, average 53 X 16 /z; 
upper cell brown, truncate, thickened at apex, giving off irregular, 
blunt, generally curved processes variable in size and shape, 
averaging 6 to 7 or more ; lower cell generally paler in colour, 
longer than upper, and attenuated towards base ; pedicel persis- 
tent, coloured, short, stout, up to 1 1 /z 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. 

In 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 A vena saliva L., and Avenafatua 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 > 2 ). 

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 /u in length, and from 18-22 p. in 
breadth at the apex, and 7-8 p at the base. 

By infection experiments Klebahn 2 proved the connexion between this 
rust, and the aecidiurn on Rhamnns frangula L., but this genus does not 
occur naturally in Australia. 

P. coronata Corda, was the name originally given to the species occurring 
on Lolium percnne, Avena sativa, &c., with teleutospores surmounted by a 
crown of finger-like processes. It was found by culture experiments that 
the aecidial stage was produced on species of Rhamnus, and Klebahn deter- 
mined that the spores from some species of grasses produced aecidia 
exclusively on jRhamnus cathartica, while others did so exclusively on 
JR. frangula. This showed a biological distinction between the two, and 
Klebahn proposed the name of P. coronifera for the rust producing aecidia. 

Puccinia Gramineae. 125 

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. 

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/z; intermixed with 
large numbers of clavate paraphyses, club thickened at apex, of a 
dark smoky-brown and stalk hyaline, commonly 80-90 p 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 1 2 p}, sometimes bluntly 
pointed, hardly constricted at septum, 35-55 X 14-21 ^w, average 
38 X 18 IJL ; lower cell attenuated towards base; pedicels firm, 
persistent, coloured yellowish, about length of spore or longer, 
up to 70 ju. 

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 (Rodway ] ). 

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 otlier 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. 


Puccinia Gramineae. 

Subsequent research showed that there were still other species on the 
same host-plant. Plowright separated a third species in 1888, mainly based 
upon the fact that the aeeidiospores were only produced upon Rumex acetosa 
and not upon other plants. Next, Ludwig in 1892 distinguished another 
new rust on material sent from South Australia, viz., P. tepperi. Finally, 
Arthur in 1902 published another new species which he named P. simillima 
because it was very similar to P. magnusiana. 

There are thus at least five different species of Puccinia recorded upon 
Phragmites communis, and it is not always easy to differentiate them, at 
least by morphological characters. In three of them the aeeidiospores were 
proved to occur on different host-plants by a series of cultures, and this is 
considered by many a sufficient specific distinction. On these grounds they 
would 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 to 
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 
in Victoria and South Australia. 

The life-history of this fungus was first determined by 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 1 and Klebahn. 1 

The five species may be arranged according to their aecidial hosts whei 
known, as in the following table with their special characteristics shown :- 

Rumex sp. and 

Rumex acetosa. 

osori without paraph 



Ranunculus repens 
and R. bulbosa. 
Uredosori wil 

Anemone dichotoma. 
,h paraphyses. 

P. phragmitis. 

25-35 x 15-23/u 
45-65 x 16-25/t 
markedly con- 
150-200 [i long 

P. trailii. 

25-35 x 20-25 /* 

50-60 x 20-23 p, 
markedly con- 

75-100 p. long 

P. tepperi. 

27-30 x 20-23 ju 

45-68 x 18-26/i 
markedly con- 

1 80-250 n long 

P. magnusiana. 

24-35 x 16-19 fi 

35-55 x 14-21 /i 
hardly con- 

About length of 

P. simillima. 

26-44 x 18-22 /< 

42-56 x 15-20 / 
hardly con- 

About length of 

It will be seen from the above that there are two types clearly 
distinguishable morphologically, paraphysate and aparaphysate, and withi: 
these limits there are only " biological species " based upon a difference - 
aecidial host. 

If we compare P. trailii with P. phragmitis, tho teleutospores differ * 
having a granular spore-membrane, and shorter and stouter pedicels. If 
similar comparison is made of P. tepperi, the length of the pedicel is the 
striking feature, and it is, perhaps, rather premature to separate it from 
P. phragmitis until a distinct aecidial host has been proved. 

According to Arthur, P. simillima is distinguished from P. magnusiana 
in the slender pointed teleutospores, the more clavate and less roughened 
uredospores and the probably different aecidial host, viz., Anem ( 
dichotoma L 

(Plate II., Fig. 18.) 

Puccinia Gramineae. 127 


47. Puccinia maydis Bereng. 

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, 

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 p. 

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 p), sometimes truncate, roundated, 
base and rarely somewhat attenuated, occasionally 3 or 4 celled, 
32-52 x 16-24 ^, average 36 x 20 p. 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 p. 

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. 

New South Wales. Richmond (Cobb). 

Queensland. Toowoomba, 1887 (Tryon 1 ), (Bailey 1 , 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 
to be distinct Sydow in his Monograph has reserved the above name for 
this one. 

Arthur 4 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. 

Darlucafilum Cast., occurs on the uredosori. 

(Plate II., Figs. 15, 16; Plate C., Figs. 14, 15.) 


48. Puccinia perplexans Plow. 

Plowright, Quart. Journ. Micro. Sc. XXV., p. 164 (1885). 
Sydow, Mon. Ured. L, p. 719 (1903). 
Sacc. Syll. VIL, p. 632 (1888). 

II. Uredosori amphigenous and on sheath, orange, elliptic, rupturing 
epidermis and often confluent in lines. 

I2 8 Puccinia Gramineae. 

Uredospores orange-yellow, subglobose to elliptic, finely echini 
late, with 4-6 scattered germ-pores on one face, 25-29 x 19-22 

III. Teleutosori amphigenous and on sheath, covered by leaden- 
coloured epidermis, at first minute, linear, then confluent in ve 
long lines and even forming patches. 

Teleutospores variable in shape, clavate to oblong, constricted at 
septum, brown, upper cell darker than lower, rounded or truncate 
at apex and slightly thickened, lower cell attenuated towards base 
or rounded, 35-52 x 16-24 p, average 37 X 17 p ; pedicel short, 
hyaline, may reach a length of 20 p. 

X. Mesospores common, similarly coloured to teleutospores, slighth 
thickened at apex, elliptic to ovate or pear-shaped, 25-32 x 
16-21 p. 

Oil Alopecurus geniculatus L. 

Victoria Minyip, Nov., 1903 (Eckert). 

Among these plants one was badly rusted with the uredospores 
Puccinia graminis but no trace of the teleutospores of that fungus w 
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. 

Nielsen, Bot. Tids. II., p. 26 (1876). 
Cooke, Handb. Austr. Fung., p. 336 (1892). 
Sydow, Mon. Ured. L, p. 795 (1904). 
Sacc. Syll. VII., p. 625 (1888). 

II. Uredosori small, round or elliptical, scattered or aggregated togeth( 

orange-yellow or foxy- red. 

Uredospores shortly elliptical, finely echinulate, orange-yellow 
21-24 x 15-18 p, seated on hyaline pedicels up to 30 p long, am 
intermixed with numerous, stiff, capitate paraphyses, reaching 
length of 68 p. 

III. Teleutosori small or large, scattered or orbicularly dispos 
dark-brown to black, covered by the epidermis. 

Teleutospores elliptical or subclavate, very variable, aj 
truncate, rounded or conical, thickened (4-8 p), dark-brown, 
smooth, not or hardly constricted at septum, 35-45 x 15-21 
or may reach a length of 53 p when 3-celled, average 38 x 19 
pedicels persistent, short, brown. 

On leaves and stems of Poa annua L. 

Victoria Near Melbourne, Jan.-Dec., 1886, &c., II., III. 
New South Wales Sydney, 1890 (Cobb). Botanic Gardei 
Sydney, May, 1898. 

On Poa caespitosa Forst. 

Victoria Botanic Gardens, Melbourne, Dec., 1904. 

On Poa pratensis L. 

Victoria Port Fairy, Sept., 1904. Near Melbourne, Sept., 1904. 

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, 
the upper one being usually oblique, or the upper third may be divided 
longitudinally. In these abnormal forms the length may reach 53 jj. and 
the breadth 34 /j. 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 
Plowright in 1882, who found that the aecidiospores produced the uredo- 
spores on POOL 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. Lagerheim 1 , has found the spores on the 
leaves after the melting of the snow. 

Darlucafilum Cast., commonly occurs. 

(Plate III., Fig. 22.) 


50. Puccinia purpurea Cooke. 

Cooke, Grev. V., p. 15 (1876). 
Sydow, Mon. Ured. T., p. 803 (1888). 
Sacc. Syll. VIL, p. 657 (1888). 
Uredo sorghi Fckl. 

II. Uredosori amphigenous, seated on indeterminate, elongated and con- 
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 ^/, average 31 x 21 /j. ; 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 p. ; pedicel hyaline, persistent, 
thick, up to 100 n long ; paraphyses present.] 

On leaves of Johnson Grass (Sorghum halepense Pers.) and Sugar-cane 
{S. vulgar e Pers.). 

Queensland Gladfield (Gwyther), Nerang (Shirley), (Bailey H , 15 ). 

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. 

Darlucafilum Cast., is very common on uredosori. 

Puccinia Gramineae. 


430 and 

51. Puccinia simplex (Koern.) Eriks. and Henn. 

Eriksson and Henning, Getreider. p. 238 (1896). 
Me Alpine, Journ. Dep. Agr. Victoria I, pp. 

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 01 

sometimes numerous on both surfaces of leaves, but mostly on 
upper, citron yellow. 

Uredospores subglobose or ellipsoid, echinulabe, yellow, relatively 
thick walled, with numerous scattered germ-pores, 4-7 on one 
face, 21-24 p diam, or 24-30 X 17-20 u. 

III. Teleutosori covered by the epidermis, amphigenous, numerous, 
minute, punctiform, mostly oblong and confluent, black, somewhat 
longer on leaf sheath than on leaf, divided into compartments 
brown paraphyses, often strongly enlarged and thickened towards, 
the apex and spread out horizontally. 

Teleutospores oblong clavate to clavate, dark chestnut brown, 
smooth, usually rounded or truncate at apex, or obliquely conical, 
and slightly thickened at apex, but may sometimes reach 6 p, an<~ 
slightly constricted at septum, 40-54 X 17-24 p ; upper eel 
oblong, generally as broad as long, 15-21 X 17-22 p ; lower cell 
tapering towards pedicel, sometimes quite narrow at base, long* 
than upper, 21-33 x 15-18 p ; pedicel short, slightly tinted. 

X. Mesospores very numerous, similarly coloured and similarly 
thickened at apex to teleutospores, asymmetrical, sometim< 
elongated, saccate or inversely triangular, very variable, 24-45 x 

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 unicellub 
spores. A sorus may consist entirely of these or there may be a few norm* 
teleutospores intermixed. They are produced alongside the two-celled forms 
and are quite similar to them except that they are one-celled, and thei 
evidently show the transition from the one-celled to the two-celled teleul 
spore. There is usually a slight thickening at the apex of both, but it rareb 
exceeds 4-6 p. It was early recognised as an exceptional form, and 
named Uromyces hordei by Nielsen in 1875, and Puccinia anomala by Rostru] 
in 1876. 

The teleutospores only germinate after a period of rest, according 
Eriksson and Henning 1 . 

(Plate I, Figs. 1, 4, 9 ; Plate B., Figs. 9, 10.) 

Pucdnia Gramineae. 13! 


52. Puccinia subnitens Diet. 

Dietel, Erythea, p. 81 (1895). 

II. Uredosori' hypophyllous, orange, powdery, soon naked and sur- 

rounded by ruptured epidermis, often in long lines and confluent 

Uredospores yellowish-brown, globose to slightly ellipsoid, 
densely echinulate, with 3-4 scattered germ-pores on one face 
thick-walled, 21-23 p. diam. or 21-28 x 18-20 /u. 

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 at septum and 
thickened at apex (4-7 ju), smooth, 28-46 x 19-23 jj. ; pedicel 
persistent, yellow, up to 56 /z long. 

X. Mesospores not uncommon, similarly coloured to teleutospores, 
obovate, rounded and thickened at apex (up to 7 /u), smooth, 
28-37 x 19-22 ^ ; pedicel similar to that of teleutospore. 
Darlucafilum 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 Trin., the host-plant 
being mistaken for a Distichlis. 

Arthur 7 has obtained some remarkable results by sowing the- teleuto- 
spores of this species derived from Distichlis spicata 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 Kewensis, D. spicata is a synonym of D. maritima. 

(Plate XXX., Figs. 262-264.) 


53. Puccinia tepperi Ludw. 

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 p 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 p. 

III. Teleutospores intermixed with uredospores, subcylindrical to 
occasionally oblong, with rounded or acute slightly thickened 
apex (4-5 /*), 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 /i, 
average 50 x 19 n; pedicel hyaline, persistent, up to 250 // long. 

j-^2 Puccinia Gramineae. 

X. Mesospores very rare, similarly coloured to teleutospores, elongated 
ellipsoid, rounded and slightly thickened at apex, tapering very 
slightly towards base, 58 X 21 ju. 

On Phragmites communis Trin. 

S. Australia The -Grange, near Adelaide, April, 1891 (Tepper) 
(Ludwig 3 ). 

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.) 


54. Puccinia triticina Eriks. 

Eriksson, Ann. Sci. Nat. Series VIII., p. 270 (1899). 
Sydow, Mon. Ured. I. p. 716 (1903). 
Sacc. Syll. XVII., p. 376 (1905). 

P. dispersa f. sp. tritici Eriks. and Henn., Zeitschr. f. 
Pflanzenk. IV., p. 257 (1894). 

II. Uredosori 1-2 mm. long, reddish-brown, amphigenous, but mostly 
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 18-21 p. 

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 /u, average 48 x 16 ^; upper cell deeply 
coloured, generally rounded or flattened at apex and thickened, 
17-31 x 15-18 n ; lower cell paler and attenuated towards pedicel, 


longer and narrower than 
short, coloured. 

upper, 22-36 x 12-14 p ; pedicel 

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 p. 

On wheat, Triticum vulgar e Vill. and T. polonicum L. Common. 

Victoria, New South Wales, Queensland, South Australia, West 
Australia, and Tasmania. 

The uredospores are distinctly different from those of P. graminis in being 
subglobose instead of elongated, and the more numerous germ-pores are 
scattered instead of forming an equatorial band. 

Uredospares 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. 1, 2.) 

Puccinia Cy-peraceae. 133 


Carex, Urtica. 
55. Puccinia caricis (Schum.) Rebent. 

Rebentisch, Fl. Neom., p. 356 (1804). 

Sydow, Mon. Ured. L, p. 648 (1903). 

Sacc. Syll. VII., p. 626 (1888). 

Aecidium urticae DC. Fl. franc. II., p. 243 (1805). 

0. Spermogonia in small or large groups, honey coloured. 

Spermatia hyaline, ellipsoid, 4J x 2 p. 

1. 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 p. 

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 p. 

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 p) and 
rounded or bluntly pointed, lower cell generally attenuated 
towards base, 35-64 x 16-22/1, average 52 x 22/z; pedicels 
tinted, persistent, generally short, up to 30-40 /t long. 

X. Mesospores not common, elongated elliptical, coloured like telea- 
tospores, thickened and bluntly pointed at apex, 35-44 x 
12-16 p. 

Aecidia on leaves, leaf stalks and stems of Urtica dioica L. 

Victoria. Bacchus Marsh, Myrniong, Fern Tree Gully, etc. 
TJredo and teleutospores on leaves of Carex breviculmis R. Br. 

Victoria Killara, Sept., 1902 (Robinson.) 
On Carex pedunculata Muhl. 

Tasmania Cascades, May, 1892, II., III. (Rodway). 
On Carex yunniana 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 paniculata L. 

Victoria Gembrook Ranges, March, 1904, II., III. (C. French, 


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. 


Puccini a Cyperaceae. 

Magnus first experimentally proved, in 1872, that Aecidium urticae 
DC., was genetically connected with Puccinia caricis occuring on Car ex hirta. 
Dr ' Plowright also, in many cultures, has shown that the 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. 

Magnus 1 has also shown that this rust winters on Carex hirta by 
means of the uredo-generation. 

(Plate IV., Figs. 29, 30.) 

56. Puccinia cyperi Arthur. 

Arthur, Bot. Gaz., p. 226 (1891). 
Sydow, Mon. Ured. L, p. 681 (1903). 
Sacc. Syll. XL, p. 199 (1895). 

II. Uredosori hypophyllous, on brown effused spots, scattered irregu- 

larly or aggregated, ellipsoid to oblong, variable in size, 1-2 mm. 
long, long covered by the thick epidermis, ochraceous or brownish. 
Uredospores yellowish-brown, ellipsoid, echinulate, 22-25 x 
19-21 ju. 

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 
elongated fusiform, brown and paler beneath, smooth, constricted 
or hardly constricted at septum, 35-57 X 15-20 p, average 48 X 
15 fj. ; upper cell rounded or often obtusely and obliquely acuminate 
at apex, and thickened (up to 13 p) ; lower cell paler, oblong, but 
usually attenuated towards base ; pedicel coloured like lower cell, 
persistent, 20-30 ^ long, but may reach a length of 40 p. 

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 p), 30-32 X 13-17 /t ; pedicels 
similarly coloured, persistent, up to 22 f.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 p. 

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 steems to be that of paraphyses, and 
it may be in some other cases as well. 

Darluca filnm Cast., was common on uredosori. 

(Plate IV., Fig. 32.) 

Pticcinia Cyperaceae. ^ 

57. Puccinia longispora McAlp. 

IT. 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 /z diam. 

III. Teleutosori black, compact, elongated, buliate, crowded, confluent 
in long black lines, covered for some time, then girt by ruptured 

Teleutospores clear yellowish-brown to smoky-brown, elongated, 
narrow, smooth, constricted at septum, 50-80 X 14-19 ^, average 
63 X 19 fj. ; upper cell darker than lower, generally rounded and 
slightly expanded at apex, sometimes bluntly pointed, much 
thickened (9-12 p) ; 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. vulgar is 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 ^u. 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 intb 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 

(Plate IV., Fig. 31.) 


Puccini a Juncaceae. 


58. Puccinia juncophila Cooke and Mass. 

Cooke and Massee, Grev. XXII., p. 37 (1893). 

Sydow, Mon. Ured. I., p. 644 (1903). 

Sacc. Syll. XL, p. 199(1895). 

Uredo armillata Ludwig, Bot. Centrbl. XLIII. p. 



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 spiiiulose, 
relatively thick-walled (3-4 p) yellowish-brown, 24-32 x 16-22 /*. 

III. Telutosori dark-brown, elliptic to oblong, surrounded by ruptured 
epidermis, confluent. 

Teleutospores intermixed with uredospores, elliptical to oblong, 
golden-brown, with finely granular contents, slightly constricted 
at septum, rounied and a little thicker at apex (5-6 p), lower cell 
rounded at base and generally resembling upper, but often some- 
what narrower, 30-40 x 20-28 p, average 36 X 25 yn ; epispore 
thick, smooth ; pedicel hyaline, generally deciduous, often 
attached obliquely, up to 54 /u long. 

X. Mesospores similarly coloured to teleutospores, shortly elliptical, 
occasionally globose, smooth, thickened at apex, 23-28 x 19-21^. 

On Juncus effusus L., J. mwitimus Lam., J. pallidus R. Br., J. 
pauciflora R. Br., and other species. 

Victoria Coromby, Oct., 1889, II. (Tepper), the original of 
Uredo armillata Ludw. Dimboola, Nov., 1891, II., III. 
(Reader). 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 the uredospores, since 
they are given as 16-18 x 12-14 p. 

Through the kindness of Dr. Morrison, I have also been able to examine 
some of the original material from Oakleigh. 

Uredo armillata was the name given at first by Ludwig in 1889, from the 
uredospores only being found on Juncus pallidus, but the subsequent dis- 
covery of teleutospores intermixed with uredospores showed that it was a 
Puccinia. Material sent by Reader to Kew Herbarium was originally 
referred to Uromyces junci by the authorities there, but the identical 
specimens being later forwarded to us the two-celled toleutospores weiv 
found mixed with the uredospores. Uromyces junci, therefore, has not been 
shown to exist in Victoria. 

Darluca filum Cast., was very common, in some cases rendering the 
patches quite black. It occurred both on uredo and teleutosori. 

(Plate IV., Figs. 35, 36.) 

Pticcinia Juncaceae. 137 

59. Puccinia tenuispora McAlp. 

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 

Uredospores yellowish to pale yellowish -brown, elliptical to 
obovate, echinulate, with 1-2 germ-pores on one face 20-25 x 
15-22 p. 

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 
ll/i, attenuated towards base or rounded, constricted at septum, 
30-50 x 14-20 p, average, 42 x 17 ^ ; pedicel hyaline, per- 
sistent, generally about 30 p long. 

X. Mesospores abundant, clavate to obovate or oblong, smooth, 
thickened at apex and similarly coloured to teleutospore, 22-32 x 
13-16 p. 

On leaves and stems of Luzula campestris L. 

Victoria Murramurrangbong Ranges, Nov. and Dec., 1902-3, 
Jan., 1905 (Robinson). Wandin, Nov., 1903 (C. French, 

On Luzula oldfieldii Hook. f. 

Tasmania Mt. Wellington, Jan., 1892, II. (Rod way). 

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 belong to 
the Old World species, i 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 Luzula 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 p, and the size is 40-80 x 16-24 /i. 

The teleutospores are frequently found germinating, showing that they 
do not require to undergo a period of rest. 

Dariuca filum Cast, occurs frequently on the uredosori. 
(Plate IV., Figs. 33, 34.) 


Puc cinia Liliaceae . 



60. Puccinia burchardiae Sacc. 

Saccardo, Hedw. XXXII., p. 57 (1893). 

Ludwig, Zeitsch. f. Pflanzenkr. III., p. 137 (1893). 

McAlpine, Proc. Roy. Soc. Vic. VII., N.S., p. 215 (1894). 

Sydow, Mon. Ured. I., p. 620 (1903). 

Sacc. Syll. XI., 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.. 

III. Teleutosori amphigenous, numerous, small or large, erumpent, 
surrounded at base by cuticle of epidermis, circular cr elliptical, 

Teleutospores clavate, constricted at septum, thickened at apex 
(up to 14 or 15 p) and rounded, sometimes truncate or acute, 
40-60 x 14-23 yu, average 48 x 20 p., but sometimes attaining a 
length of 70 p; lowejr cell tapering towards pedicel, yellowish- 
brown; upper cell elliptical or rarely spherical, chestnut-brown, 
21-23 p broad; epispore smooth; pedicel hyaline, 30-60 x 5-6 p. 
X. Mesospores fairly common, similarly coloured to teleutospores or 
paler, clavate to ovate, rounded or pointed at apex and thickened, 
37-43 x- 12-15 p. 

On steins 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 differing from the type 
P. metanarthecii Pat. 

(Plate V., Figs. 38, 39.) 


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 p. 

III. Teleutospores clavate, slightly constricted at septum, smooth, 
upper cell convex or truncate, at apex darker, lower cell tri- 
angular, attenuated downwards into the short, hyaline pedicels, 
48-60 x 21-28 /z, average 50 x 24 p (according to Cooke and 
Massee, 60-70 X 20-25 p) ; pedicels persistent, generally short, 
but attaining a length of 52 p and a breadth of 7-8 p, being 
broadest at the junction with the spore. 

Puccinia Haemodoraceae, Amaryllidaceae. 139 

X. Mesospores numerous, similarly coloured to teleutospores, elongated, 
elliptical to oblong, thickened at apex and acute, rounded or trun- 
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.) 



62. 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 p, average 46 x 19^i; pedicel 
persistent, slightly brownish towards apex, 40-55 x 5-7 //. 

X. Mesospores very common, subfusoid or ovoid to subclavate, apex 
gibbous to apiculate, somewhat obtuse or two-horned, thickened at 
apex (up to 9 ju), brown, 25-40 x 12-18 p, pedicel hyaline to 
brownish. Occasionally a more deeply coloured spore occurs, 
with somewhat truncated apex, 30-46 x 5-7 p.. 

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 

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.) 



63. Puccinia hypoxidis McAlp. 

McAlpine, Agr. Gaz. N.S.W. VL, 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. 


Puccini a Labiatae. 

Uredospores shortly elliptical, yellowish, finely echinulate, 20-23 
x 16-18 jji, average 21 x 16 p. 

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 /*, 
average 42 X 18 /u ; upper cell squarish or oblong or club-shaped, 
apex very much thickened (up to 15 p), 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 p long. 

X. Mesospores chestnut to pallid, rounded or beaked at apex and 
thickened, somewhat fusiform to elongated elliptical, 25-36 x 
11-18 p. 

On leaves of Hypoxis glabella B. Br. 

Victoria Ardmona, July, 1893 (Bobinson). Burnley, Buther- 
glen, &c., July-Oct. 

It differs from P. burchardiae Sacc. in the much smaller uredospores, 
and decidedly in the teleutospores. which in the latter are generally rounded 
at the apex or beaked, and distinctly constricted at the septum. 

(Plate Y.,. Fig. 41.) 


64. Puccinia menthae Pers. 


Persoon, Syn., p. 227 (1801). 
Sydow, Mon. Ured. I., p. 282 (1902). 
Sacc. Syll. VIL, p. 617 (1888). 

[O. Spermogonia either arranged in small groups or scattered, honey- 

[I. Aecidia hypophyllous, or frequently on stems which are much'swollen, 
more rarely on purplish-red spots on leaves, seldom scattered ; 
pseudoperidia immersed, flat, opening irregularly and margin 
scarcely or irregularly torn. 

Aecidiospores subglobose, ellipsoid or polygonal, coarsely granular, 
pale yellowish, 24-40 x 17-28 p.] 

II. Uredosori hypophyllous, small, seated on pale spots, roundish or 

elliptical, scattered or aggregated, soon pulverulent and confluent, 
girt by the ruptured epidermis, cinnamon-brown. 

Uredospores subglobose, ellipsoid or obovate, echinulate, pale- 
brown, generally three equatorial pores, 20-30 X 16-19 p.. 

III. Teleutosori similar, but blackish-brown in colour. 

Teleutospores at first intermixed with uredospores, ellipsoid to 
ovate, rounded at both ends, apical papilla pale or hyaline, broadly 
expanded, hardly constricted at septum, finely warted, golden to 
chestnut-brown, 24-32 x 19-22 p, but commonly 24-25 x 
19-20 p ; pedicel hyaline, slender, generally surpassing length of 
spore, up to 40 p. 

Pucdnia Acanthaceae. 141 

X. Mesospores, occasional, similarly coloured, nearly globose, very 
slightly roughened, 20 p diam. 

On leaves of Mentha laxiflora 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.) 



65. Puccinia mussoni McAlp. 

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 /* 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 p; 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 p long and 7 p 

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 difterences 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 (B. 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 /*, while here they are larger. The 
teleutospores likewise only measure 29-34 x 18-22 p, and are very strongly 

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.) 


Puccinia Convolvulaceae. 


66. Puccinia dichondrae Mont. 

Montagne in Gray's Fl. Chil. VIII., p. 46 (1845). 

Cooke, Handb. Austr. Fung., p. 338 (1892). 

Sydow, Mon. Ured. I., 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 fj. diain. 

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 ju, average 32 x 14 /z ; pedicel hyaline, sometimes 
tinted, persistent, generally slender, up to 46 p long, occasionally 
6 p broad. 

X. Mesospores sub-ellipsoid to oblong, similarly coloured to teleutospore, 
thickened at apex and usually with hyaline apiculus, 21-30 x 
15-22 p. 

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 Maiden 1 . 

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 ^i) 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 /LI 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-plant. 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 Apocynaceae. 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 
Diorchidium, 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.) 



67. Puccinia alyxiae Cooke and Mass. 

Cooke and Massee, Grev. XVI., p. 2 (1887). 
Cooke, Handb. Austr. Fung., p. 338 (1892). 
Sydow, Mon. Ured. L, p. 336 (1902). 
Sacc. Syll. VIL, 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 p, average 45 x 20 p. ; epispore thick, 
smooth, coloured ; pedicels persistent, hyaline, broadish, elongated, 
up to 130 //, 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 B. Br. 

Victoria Brighton and Broadford, Sept., 1887 (Miss Campbell 1 ). 
Beaumaris, March, 1895. Cheltenham, May, 1902 (C.French, 
jun.) Sandringham, April, 1905. (Bobinson). 

South Australia Gawler, Sept., 1893. (Tepper). 

Tasmania Near George's Bay, Nov., 1892. (Bodway '). 

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 p, but this is evidently a misprint. 

(Plate VI., Fig. 46.) 


Puccinia Apocynaceae, Goodeniaceae. 


68. Puccinia carissae Cooke and Mass. 

Cooke and Massee, Grev. XXII., p. 37 (1893). 
Sydow, Mon. Ured. 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 p, average 
32 X 22 p ; occasionally 3-celled, then reaching to a length of 
49 f-i ; pedicel hyaline, deciduous or persistent, sometimes origin- 
ating laterally, rather long. 

X. Mesospores occasional, just resembling a teleutospore, but without 
the septum, elliptic, with hyaline pedicel, 29-31 x 22-25 p. 

On living leaves of Carissa ovata R. Br. 

Queensland Gladfield, (Gwyther) (Bailey l3 ). Dalby, (Bancroft) 
(Bailey 1G ). 

Mr. Bailey has kindly sent me some excellent material from his herbarium 
and the teleutospores have been carefully measured. In the original 
description the size is given as 20-22 x 16 /*, but probably 30-32 p was in- 
tended for the length. 

Sometimes both upper and lower cell have an oblique division. 

It differs from P. alyxiae Cooke and Mass., in which the spores are much 
larger and thickened at the apex. 

(Plate VI., Fig. 45.) 



69. Puccinia brunoniae Me Alp. 

McAlpine, Agr. Gaz. N.S.W. VI., p. 851 (1895). 
Sydow, Mon. Ured. L, p. 193 (1902). 
Sacc. Syll. XIV., 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 01^ more often circularly arranged on reddish, 

slightly swollen indeterminate spots, mostly on upper surface of 
leaves and on petioles ; pseudoperidia with white margin, torn, 
slightly recurved and soon falling away, average ^rnm. diam.; 
isolated peridial cells, generally short and stout, less commonly 
elongated oblong, punctate, with striated margin, generally 
24-31 x 14-17 ,u, or when elongated 38-45 x 21-24 jz. 

Aecidiospores subglobose or angular, or even oval, deep orange, 
smooth, 17-21 p diam. or 18-21 x 15-16 p, occasionally as long 
as 24 . 

Puc curia Goodeniaceae. 145 

III. Teleutosori amphigeiious, 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 ju, average 50 x 20 p. ; 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 p. 

X. Mesospores occasional, similarly coloured to teleutospores, rather 
oblong, thickened at apex, rounded or truncate, slightly narrowed 
at base, 28-38 x 17-21 //. 

On leaves and petioles of Brunonia australis Sm. 

Victoria Drysdale, Oct., 1895, III. Murramurrangbong Ranges. 
Nov., 1902, Dec., 1903, O., L, 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 p and in breadth from 21-24 //, the lower cell generally about 
as long as the other two. 

Four-celled teleutospores are very rare, elongated clavate, 63 X 24 p y 
the two upper cells about 33 x 24 p, and the two lower 30 x 14 p. In 
the same group there were three 3-celled teleutospores with an average 
size of 59 x 23 //. 

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 

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 p diam., and the peridial cells as 18-25 X 15-18 //, 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 Brunoma, 
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.) 


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, ^ ^ mm. 
in diam. ; peridial cells lozenge-shaped to oblong, with thick striated 
margins, 30-40 p long. 

Aecidiospores orange, subglobose, ellipsoid to oblong, finely 
echinulate, 15-17 p diam. or 17-19 x 13-14 p. 

III. Teleutosori on stems, scattered or aggregated, round or oblong, 
about 1 mm. diam., compact, dark-brown, girt by the ruptured 

Teleutospores clavate, rounded or acute at apex and strongly 
thickened (8-11 /;), constricted at septum, attenuated down- 
wards, rarely round, smooth, brown, 48-66 x 16-26 p.; pedicel 
yellowish, persistent, up to 80 JJL long. 

X. Mesospores intermixed with teleutospores, clavate, 40-55 x 14-22 /*. 

On stem and leaves of Dampiera stricta R. 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. 


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 /u. 

Aecidiospores subglobose or ellipsoid, angular, finely echinulate, 
yellowish, 15-20 /u diam. 

II. Uredosori on stems, gregarious in streaks, oblong, surrounded or 

almost covered by the ruptured brown epidermis, somewhat 

Uredospores subglobose, ovoid or ellipsoid, yellowish to brown, 
echinulate, 14-21 x 12-18 p. 

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 p.), obliquely papillate and occasionally crested, rounded at 
base or attenuated, brown, 30-45 x 16-22 //, average 42 X 20 /u ; 
pedicel brownish, thick, persistent, 30-50 fj, long. 

Puccinia Goodeniaceae. 147 

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.)' 

Goodenia, Velleia. 

* 2. Puccinia saccardoi Ludw. 

Ludwig, Hedw. XXVIII., 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 goodeniacearum (in part) Berkeley, Linn. Journ. 
XIIL, p. 173 (1872). 

I. Aecidia on yellowish or brownish spots, scattered or in groups 
arranged in circular patches (up to 6mm. diam.), amphigenous 
causing distortion of stem ; pseudoperidia cup-shaped, with white 
torn, everted edges, 215-325 p, 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 p. 

Aecidiospores subglobose, polygonal to oval, orange, very finely 
verrucose, 14-16 p, diam., or 15-17 x 13-14 p. 

III. Teleutosori on both surfaces of leaves and on stems, black, com- 
pact, roundish or elongated, confluent in loag swollen patches, 
generally intermixed with or surrounding aecidia, bursting through 

Teleutospores clavate to oblong, dark-brown, constricted at 
septum, smooth, variable in size, rarely 3-celled, 40-66 x 17-25/7, 
occasionally up to 70 p long, average, 54 x 20 p ; upper cell 
subglobose or somewhat quadrate, thickened at apex (up to 9 /*), 
generally rounded or truncate, occasionally pointed, 21-30 x 
17-25 //, sometimes reaching 32 fj. 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 /i, 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 /u), and generally 
slightly tapering towards base, 25-49 x 12-21 p, or averaging 
36 x 17 . 

j 48 Puccinia Campanulaceae . 

On steins, leaves and calyx of Goodenia geniculata R. 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 

S. Australia Tanunda Scrub, Oct., 1887 (Tepper). Mt. Brown 

Creek, near Quorn, Sept., 1892 (Molineux). 
Tasmania Cascades, Jan., May, Nov., 1892 and 1893, I. 

(Rodway 1 ). 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 pinna tifida, 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 
geniculata a similar variety, so that the spores on the genera Goodenia and 
Velleia cannot be differentiated. He writes " Mixed among the nori 
teleutospores there are three and one-celled spores, the latter sometimes 
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 a& 
the upper, proceeded from the lower cell." 

(Plate VI., Figs. 48, 49.) 


Lobelia , Pratit 

73. Puccinia aucta Berk, and F. v. M. 

Berkeley and F. von Mueller, Linn. Journ. XIII., p. 173 


Cooke, Grev. XL, p. 98 (1883). 
Cooke, Handb. Austr. Fung., p. 338 (1892). 
Sydow, Mon. Ured. L, p. 196 (1902). 
Sacc. Syll. VII., p. 676 (1888). 

Aecidium microstomum Berk. Journ. Linn. Soc. XIII, p. 173 

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 p. 

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 p. 

Aecidiospores irregularly globose, sub-angular or ellipsoid, finely 
echinulate, with distinct wall, ochraceous, 18-22 p. or 22-28 x 
18-20 p. 

III. Teleutosori on under surface, numerous, bullate, then girt by the 
ruptured epidermis .so that they resemble little Pezizae. 

Teleutospores cylindric to sub-clavate, elongated, yellow, 
thickened apex (9-10 ju), rounded or truncate, deeply constricted 
at septum 56-70 x 15-26 /i, average 60 x 24, occasionally bi* 
and tri- septate when they may reach a length of 80 p ; upper cell 
generally broader than lower ; lower cell generally cylindrical, and 
may taper slightly towards pedicel ; pedicel hyaline, short, stout, 
7-13 jj. broad. 

Aecidiospores on Lobelia anceps L., L.pratioides Benth., L.purpurascens 
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. Wilhelmi), 
(Berkeley 2 ). 

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 mueh 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 
in the National Herbarium, named in Berkeley's handwriting. 

(Plate VI., Fig. 51 ; Plate XXXIX., Fig. 289.) 

jr O Puccinia Compositae. 



74. Puccinia angustifoliae Me Alp. 

I. Aecidia in dense clusters on the leaves, more sparingly on stems, 
about J to nearly mm. diam. ; pseudoperidia with finely laciniate 
everted margins. 

Aecidiospores orange-yellow, globose to elliptical or sub- 
quadrate, 16 x 11-13 p or 13-14 p diam. 

III. Teleutosori on leaf and stem, black, long covered by epidermis, 
confluent, forming elongated patches, partially rupturing and 
throwing off epidermis, distinct from or along with aecidia. 
Teleutospores clavate to oblong, smoky-brown, slightly con- 
stricted at septum, smooth, generally rounded and thickened at 
apex (up to 9 /x), slightly tapering towards base or often 
rounded, 40-53 x 16-21 //, average 50 x 20 ^ ; pedicel hyaline, 
deciduous, generally short, up to 38 p, long and 9 ^ broad. 

X. Mesospores common, similarly colored to teleutospores. or paler, 
generally somewhat oval, thickened at apex, 25-30 x 16-17 yu. 

On Scorzonera anguslifolia L. 

Victoria Dimboola, Nov., 1892, I., III. (Reader). 

This species differs from P. podospermi DC., P. scorzonerae 
(Schum.) Jacky and P. tragopogi (Pers.) Corda, in the absence of 
uredospores and the smooth teleutospores. 

(Plate VII., Fig. 53.) 


75. Puccinia brachycomes Me Alp. 

I. Aecidia crowded on swollen and distorted portions of leaves, also on 

stems and branches, about ^ mm. diam.; pseudoperidia round, 
out-standing, with white, slightly everted, laciniate margin ; 
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 p diam. or 13-16 x 10-13 //. 

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 x 18-25 JJL, average 
50 X 20 /JL ; upper cell truncate or conoid, occasionally rounded at 
apex, thickened (up to 9 /JL) ; lower cell generally attenuated towards 
base, sometimes rounded and generally longer than upper ; pedic 
persistent, tinted or hyaline, up to 35 ju 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., L, 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, L, III. (C. French, jun.). 
Darluca filum Cast., is a very common parasite on the aecidia. 
(Plate VIIL, Fig. 66.) 

Puccinia Compositae. 151 


76. Puccinia calendulae Me Alp. 

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 Me Alp. Agr. Gaz. N.S.W. VII., 
p. 152 (1896). 

I. Aecidia araphigenous 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 jj. or 12- 16 p 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 p), attenuated towards base, smooth, 36-52 x 19-23 //, 
average 48 x 20 p ; upper cell darker in colour and broader than 
lower, 21-31 /x 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 yi/, 
but may attain a breadth of 10 ju 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 p. 

On leaves and all green parts, including fruits of Calendula officinalis L. 
Victoria Near Melbourne, growing in gardens, 1892, <fcc. Killara, 
July-Oct., 1902. Geelong, Nov., 1904, I., III. (Pescot-t). 
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 Gazette 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 VIII., Fig. 65 ; XLIIL, Fig. 312 ; Plate E., Figs. 22, 23.) 


77. Puccinia calocephali Me Alp. 

I. Aecidia very sparse on upper surface of hairy leaves, about J mm. 
diam. ; pseudoperidia saucer-shaped, laciniate at margin ; peridial 
cells, oblong to lozenge-shaped, striated, 25-32 /u long. 
438. F 

152 Puccinia Com-positae. 

Aecidiospores sub-globose to angular, orange-yellow, smooth, 
showing up to 4 germ-pores on one face, 16-19 X 12-13ju or 
14-16 ^u diam. 

III. Teleutosori black to blackish-brown, long covered by leaden 
epidermis, pulvinate, compact, round to elongated, sometimes 
confluent, 1-1 ^ mm. long. 

Teleutospores clavate, dark brown, particularly in upper cell, 
slightly constricted at septum, smooth, 40-57 X 17-24 ju, average 
45 X 22 p ; 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 p long and sometimes 10 p broad. 

X. Mesospores common, elongated, elliptical to oblong, generally paler 
than teleutospore, thickened at apex, and generally scoop-shaped, 
28-35 x 13-16 p. 

On leaves and stem of Calocephalus drummondii Benth., and C. lacteus 

Victoria Dimboola, Sept., 1891, III. (Reader). Phillip Island, 
Nov., 1901, I. (C. French, jun.) 

The teleutosori and teleutospores generally resemble those of P. 
tasmanica Diet., but they are not intermixed with aecidia. They both be- 
long, however, to the same general type. 

(Plate VII., Fig. 58.) 


78. Puccinia calotidis Me Alp. 

I. Aecidia amphigenous, in irregular clusters, orange ; pseudoperidij 
cup-shaped, with whitish, reflexed, laciniate margins. 

Aecidiospores subglobose to polygonal, finely echinulate, 13-14/ 
diam., or 13-15 x 12 p. 

III. Teleutosori minute, black, erumpent, gregarious, comp* 

Teleutospores dark chestnut-brown, oblong to elongated oblong 

clavate, smooth, constricted at septum, rounded or bluntly poinl 

and thickened at apex (up to 9 p), 37-50 X 18-28 p, avera, 

38-40 x 18-20^u; pedicel hyaline, persistent, generally taperii 

towards base, about 30 p. long, or shorter than spore. 

X. Mesospores not uncommon, similarly coloured to teleutospores, 

stalked, obovoid to ellipsoid, rounded or bluntly pointed 

thickened at apex (6-8 p), 28-37 x 15-18 p. 

On Calotis cuneifolia R. Br. and Calotis sp. 

New South Wales Guntawang, Sept., 1886 (Hamilton) 
Condoblin, Aug., 1897 (Maiden). 

The teleutospores generally resemble those of P. vittadiniae, but they 
longer on the average. 

(Plate XLIIL, Fig, 310.) 

Puccinia Com-positae. 153 

79. Puccinia chrysanthemi Roze. 

Roze, Bull. Soc. Myc. France, p. 92 (1900). 
Sydow, Mon. Ured. L, p. 46 (1902), p. 854 (1904). 
Sacc. Syll. XVL, p. 296 (1902). 

P. chrysanthemi chinensis P. Heim. 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 J mm. diam., causing dirty -brown, indefinite 
spots on upper surface of leaf, which gradually extend and 
ultimately decay. 

Uredospores ellipsoid to obovoid, yellowish-brown to golden- 
brown, echinulate, with 3 equatorial germ-pores on one face, 
28-35 X 22-25 p (32 p long very common, occasionally reaching 
a length of 45 p) ; pedicels hyaline, rarely persistent, stout, long, 
up to 60 f.i. 

[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 p), mostly rounded at base 
or sometimes slightly attenuated, chestnut-brown, with strong, 
finely punctulate epispore, 35-60 x 19-28 p; pedicel hyaline, stout, 
persistent, 40-80 x 7-9 p.] 

[X. Mesospores rare, ellipsoid to clavate, similarly coloured to teletito- 
spores, rounded and thickened at apex, finely ptinctulate, 28-46 x 
22-25 p.] 

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 
', 2 , 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. chrysanthemi-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 
uredospores are quite similar, having three equatorial germ-pores on one 
face, and measuring 28-37 X 22-25 fj. Jacky 3 succeeded in infecting 
C. indicum L., with material from C. sinense Sabine, and thus showed the 
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 these 
young shoots are taken as cuttings, so that there is no time when there are 
no leaves upon which the rust can 'live. 

F 2 


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 plani 
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 
on hollyhock, sunflower, and other plants may pass to the chrysanthemui 
when it has suddenly bacome temporarily weak through drought or otht 
causes ; but it ought to reassure them to know that the specific rust, known 
ajs Puncinii 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 miy be 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 by 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. It is strange that th< 
rust has not yet found its way into Victoria, considering the number 
chrysanthemums imported from England, France, America, and Japan. 

It was first observed in England in 1895, although it probably exist 
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, 
and at length it has found its way to Australia. In 1904 it was 
recorded for New Zealand by Kirk. 

It is found that many of the Australian-raised varieties are less liable 
rust than others when grown in England or America. 

(Plate XXIX., Figs. 251-255 ; Plate E., Fig. 21.). 

80. Puccinia cichorii (DC.) Bell. 

Bell., in Kickx. Fl. Fland. II. p. 65 (1867). 

Sydow, Mon. Ured. I., p. 49 (1902). 

Sacc. Syll. YIL, p. 606 (1888) ; XVII., p. 311 (1905). 

II. Uredosori amphigenous or on. stem, minute, pulverulent, scattered 

or confluent, girt or partially covered by ruptured epidermis, 

Uredospores globose, subglobose or ellipsoid, echinulat 
yellowish-brown, with one or two germ-pores on one face, sometin 
three to four, 22-29 x 19-23 /i, or 22-25 /i diam. 

III. Teleutosori similar, dark-brown. 

Teleutospores intermixed sparingly with uredospores, ellij 
or ovate-ellipsoid, brown to golden-brown, smooth, not or hard! 
constricted at septum, apex rounded not thickened, base for 
most part rounded or very slightly attenuated, epispore thii 
germ-pore in each cell prominent, with finely granular contenl 
27-38 X 19-25 ^, occasionally reaching 50 /* long, but averagii 
35 x 20 p. ; pedicel hyaline, short. 

Pvccinia Compositae. 155 

On leaves and stems of Cichorium intybus L. 

Victoria 1885 (Mueller 3 ) II. Near Melbourne, Au% 1896 
Phillip Island, Jan., 1900; March, 1904. 

The uredospores were common chiefly on the leaves, but the teleutospores 

e rather scarce, and occurred mostly on the midribs. Fischer 5 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 fj. long, and the upper colourless one only 6 p. 

Darluca filum Cast., was plentiful on the uredosori and the sori 
containing uredo and teleutospores intermixed. 

(Plate VIII., Fig. 61 ; Plate D., Fig. 18.) 


1. Puccinia cinerariae Me Alp. 

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 ju. 

Aecidiospores bright orange, subglobose to polygonal, very 
finely verrucose, 15-19 p 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 ^w, average 40 X 23 p. ; 
upper cell usually darker in colour, hemispherical, apex thickened 
to a depth of 9 p, 17-24 ju long; lower cell gradually tapering 
towards base, somewhat triangular, 18-30 p. long; pedicel per- 
sistent, tinted, up to 45 p long. 

X. Mesospores chestnut-brown, obovate to elongated elliptical, much 
thickened at apex, with long pedicel, 27-36 X 13-23 p. 

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 

ifected owing to the care that is taken to cut out the spots as soon as 
noticed. The disease spreads rapidly if neglected, and completely spoils a 
plant by denuding it of its foliage. 

Caeoma cinerariae Rostr., was found on leaves and stems in Jutland. 

(Plate VIII., Fig. 60.) 


Puccinia Compositae. 

Centaur ec 
82. Puccinia cyani (Schleich.) Pass. 

Passerini in Rabh. Fung. Eur., No. 1767. 

Sydow, Mon. Ured. I., p. 38 (1902). 

Sacc. Syll. VII., p. 634 (1888) ; XVII., p. 286 (1905). 

II. Uredosori mostly hypophyllous, scattered or crowded, minul 

orbicular or elliptic, pulverulent, reddish brown. 

Uredospores yellowish brown, globose, subglobose to ellipsoi( 
finely cchinulate, with two germ-pores 011 one face, 25-27 
19-24 ^ 

III. Teleutosori amphigenous, and on stem, scattered, pulverulent 
dark-brown, orbicular to elliptic, soon naked, J-l mm. 

Teleutospores intermixed with uredospores, chestnut-browj 
ellipsoid, rounded at both ends, unthickened at apex, not con- 
stricted at septum, delicately warted, germ-pores conspicuoi 
30-40 x 22-30 /u, average 32 x 24 yu ; epispore about 3 p thick 
pedicel hyaline, short, deciduous. 

On leaves and stems of Centaurea, cyanus L. 

Victoria Near Melbourne, Jan. and Feb., 1904 (C. French, jr.). 

This is evidently of the Puccinia hieracii type. 
The teleutospores are very regular and well defined, and there is n< 
tendency to 'abnormality. 

(Plate VIII., Fig. 64.) 

83. Puccinia distincta Me Alp. 

McAlpine, Agr. Gaz. N.S.W. VL, 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 
circinate manner, or irregularly disposed, and often distribul 
over entire surface ; pseudoperidia cup-shaped, with reflexed toi 
margins, about J-J mm. diam. ; peridial cells elongated elliptical 
oblong, punctulate all over, 25-29 x 13-19 //. 

Aecidiospores subglobose to oval or ellipsoid, orange-yellow, 
very finely echinulate, 14-19 ^ diam , or 14-20 x 13-16 p. 

III. Teleutosori intermixed with and often surrounding aecidii 
brownish black, oval, |-1 mm. long, generally confluent, burstii 
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 
15-21 p, average 42 x 17 /* ; upper cell generally darker 
colour than lower, rounded and thickened at apex (up to 9 p) 
pedicel persistent, pale yellow or coloured similarly to lower cell, uj 
to 50 /z long. 

X. Mesospores numerous, elongated clavate to somewhat ovate, chest 

nut brown, usually thickened at apex, 34-37 x 13-16 /*. 
On leaves, scapes, involucre and petals of Bellis perennis L. 

Victoria Near Melbourne, Oct., 1892, July-Oct,, 1904, L, III. 
(Robinson). Korumburra, May, 1903, I. 

Puccinia Compositae. 157 

Tasmania Near Huon Road, Nov., 1890, I. (Rodway). 
South Australia Norwood, April, 19Q3, 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 
Tlmem., 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.) 


4. Puccinia erechtitis Me Alp. 

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. Sperinogonia on brownish patches on upper surface of leaf, in groups, 

brown to honey-colour, yellowish by transmitted light. 
Spermatia hyaline, oval, 3 /JL 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 12-17 /i. 

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 p, average 47 X 19 p. ; 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 ju long and 9 p broad. 
X. Mesospores common, coloured like teleutospores, elongated oblong 

or oval, thickened at apex, 32-43 X 13-17 //. 

On leaves and stems of ErechtitesquadridentataDC., and E, argutaT>Q. 
Victoria Ardmona, Dimboola, Myrniong, and Marysville, Alps 

near Bright, and Mt. St. Bernard, &c. 
On E. quadridentata, E. arguta and E. pren&nthoides DC. 

Tasmania Huon Road, Dec., 1891, March, 1893, I. (Rodway). 

On Erechtites sp. 

New South Wales Guntawang, I. (Hamilton). 

I. Very common all the year round, except during middle of summer. 
III. 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.) 


Puccinia Com$ositae. 


85. Puccinia gnaphalii (Speg.) P. Henn. 

Hennings, Hedw. XLL, p. (66) (1902). 
Morrison, Victorian Nat. XL, 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). 
Puccinia gnaphaliicola P. Henn. Hedw. XXXVIII., p. (68) 

II. Uredo-sori common on stem and leaves, on the latter amphigenous,. 

but mostly on under surface, bursting through and surrounded by 
epidermis, also piercing woolly tomentum, scattered, sometimes; 
aggregated, compact, cinnamon, circular or oval, lenticular on 
stem, ^-i | mm. diam. 

Uredospores globose, sub -globose or ellipsoid, brown, finel 
echinulate, 21-24 /* diam. or 21-26 x 17-21 p. 

III. Teleutosori similar, but chestnut-brown to black. 

Teleutospores at first intermixed with uredospores, oblong o: 
oblong-clavate, dark chestnut-brown, slightly constricted at septum,, 
rounded or obtusely pointed or even beaked at apex, and 
thickened (up to 8 /i), usually slightly attenuated towards base 
smooth, 35-53 x 15-24 p, average 48 X 20 p, ; pedicel hyaline,, 
persistent, up to 66 x 8 p. 

X. Mesospores similarly coloured, few, oval or elongated, ellipti 

thickened at apex and sometimes beaked, 24-38 x 15-18 /i. 
On stems and leaves of Gnaphalium purpureum L., and G. japonic 

Victoria Oakleigh, Jan., 1904 (Morrison). Drysdale, Jan., 189 
and Oct., 1903. Whittlesea Ranges, Nov., 1898 (C. Frenc 
jun.) Killara, Oct., 1902. Near Melbourne, Feb. and Nov 
190-1. Murramurrangbong Ranges, Nov., 1904 (Aitken). 
The teleutospores are rather variable in length and breadth, sometime 
being short and stout and sometimes rather elongated and slender. The 
pedicels average 40-50 p. in length, and may vary in breadth from 4 p. up 
10 p. at junction with spore. Sydow gives size of teleutospores as 32-56 
18-24/1, and Dietel as 30-35 x 19-22 p. 

I am indebted to Dr. Morrison for some of the original material f: 
which he determined this fungus as P. investita Schwein. He found sp 
mogones immersed among spores in the sori, and the spermatia were fu 
form. Plenty of Darluca filum Cast., was found, and no doubt these we 
mistaken for spermogonia. 

(Plate VII., Fig. 57.) 


86. Puccinia helianthi Schwein. 

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). 
[O. Spermogonia honey-colored, in small clusters.] 

[I. Aecidia crowded or orbicular, or arranged in broadly expandu^ 
oblong *spots ; pseudoperidia cylindrical, plane, with white laciniate 


Puccinia Compositae. 159 

Aecidiospores orange-red, ellipsoid to polygonal, finely echin- 
ulate, 21-28 x 18-21/i.J 

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 /i diam. or 24-29 x 15-22 p. 

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 uredospares, chestnut- 
brown, oblong-elliptical or pear-shaped, smooth, but occasionally a 
little rough at apex, slightly constricted at septum, thickened at 
apex (6-9 p), generally rounded at base, 36-50 X 21-27 p, 
average 44 x 26 ^ ; 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 p pedicel hyaline, persistent, 
generally much longer than spore, up to 90 and 110 /i 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, Bailey 5 ; 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 

i possesses no aecidial stage, since he has examined specimens from numerous 

j localities without result, but Carleton 3 has collected the three stages in 

) America and remarks : "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 Woronin 1 the teleutospores germinate equally well 
whether they have been kept dry in a room or taken from leaves which had 
i lain under the snow throughout the winter. Carleton 2 says that they also 
I germinate at once without a resting period. 

(Plate VII., Fig. 56.) 


,S7. Puccinia hypochoeridis Oud. 

Oudemans in Nederl. Kruidk. Arch. II., Ser. 1, p. 175 (1873). 
Sydow, Mon. Ured. L, 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). 

160 Pucdnia Compositae. 

II. Uredosori umphigenous or frequently on stem, generally seated on 

minute spots, scattered, pulverulent, cinnamon-brown. 

Uredospores globose, sub-globose or ellipsoid, echinulate, pale- 
brown, with one prominent germ-pore on one face, 21-25 /j. dianu 
or 20-28 x 18-24 ft. 

III. Teleutosori amphigenous, often on stem, scattered or crowded, 
punctiform, blackish to dark-brown, pulverulent, generally 
roundish on leaf and up to 1 mm. diam., but often smaller, 
elongated on stem and up to 2 mm. long. 

Teleutospores at first intermixed with uredospores, oblong, 
ellipsoid or obovate, apex generally rounded, and not thickened, 
routed at base or slightly attenuated, hardly constricted at 
septum, dai'k-bro\v n, very delicately punctate, occasionally tri- 
cellular, 30-49 x 17-24 /u, average 32 x 20 /.t ; pedicel hyaline, 
generally deciduous, sometimes persistent and may reach a length 
of 77 p. 

On Hypochoeris radicata L., very common. 

Victoria Dimboola, Nov., 1889 (Reader). Ardmona, 1894 
(Robinson). Myrniong, March, 1900 (Brittlebank). Near 
Melbourne, Apr., 1901. Dookie, Oct., 1903. Altona Bay, 
Oct., 1903 (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 "). 

The punctation on the teleutospores is so fine that it may easily be o 
looked, but when the spores are mounted in water it is clearly seen. 

A specimen from Syd. Ured. Exs. 673 on H. glabra agreed very cl 
the uredospores being on an average 25 x 21 ft, and the teleutos 
34 x 18 ft. 

This species closely resembles in morphological characters P. hieracii 
(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 p. 

P. microseris described as a new species in the Agricultural Gazette of 
New South Wales was based upon a wrong determination of the host-plant 
by the collector since it was afterwards found to be Hypochoeris radicata 

Darluca jilum Cast., is sometimes common on the mixed uredo 

(Plate VIII., Figs. 62, 63.) 

88. Puccinia kalchbrenneri De Toni. 

De Toni in Sacc. Syll. VII., p. 645 (1888). 

Cooke, Handb. Austr. Fung., p. 337 (1892). 

Sydow, Mon. Ured. L, p. 93 (1902). 
II. Uredosori on both surfaces, but mostly on under, seated on in- 
determinate spots which are often confluent and variously coloured, 
scattered or gregarious, minute, at first covered, convex, firm, 
ultimately free and discoid, ochraceous. 

Puccmia Compositae. 161 

Uredospores globose, subglobose or ellipsoid, varied, yellow to 
pale brown, 20-30 x 19-26 p. 

III. Teleutosori hypophyllous, seated on the same spots, minute, brown. 

Teleutospores oblong or subclavate to lanceolate, apex thickened, 

attenuated or rarely truncate, constricted at septum, smooth, 

dark-brown, 40-57 x 15-22 p, rarely up to 27 p broad; pedicel 

hyaline, short, deciduous. 

On living leaves of Helichrysum sp. 

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. XIII., p. 6 (1884). 

Cooke, Handb. Austr. Fung., p. 335 (1892). 

Sydow, Mon. Ured., I., p. Ill (1902), and p. 863 (1904). 

Sacc. Syll. YIL, p. 612 (1888). 

P. hypochoeridis Me Alp., Proc. Roy. Soc. Yic. VII., N.S., 

p. 217 (1894). 
P. macalpini Sydow, Mon. Ured., I., p. 100 (1902). 

I. Aecidia amphigenous, spots none ; pseudoperidia scattered or in 
groups, semi-immersed, margin lacerated, white, 200-240 p diam.; 
peridial cells finely warted, lozenge shaped or elongated, 24-31 p 

Aecidiospores subglobose or elliptical, orange-yellow, very 
finely echinulate, 12-14 p diam. or 17-19 x 14 p, 

[II. Uredosori scattered, small, pulverulent, brown or mixed with 

Uredospores globose, brown, epispore rough, 20 p diam.] 

III. Teleutosori amphigenous, scattered, or surrounding aecidia, 
rather compact, dark-brown to black, raising and rupturing 

Teleutospores clavate, constricted at septum, dark-brown only 
in upper cell, epispore thickened, smooth, 45-66 X 16-22 p, 
average 49 x 17 p ; 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 Lngenopliora 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). 


Puccinia Composite** 

On L. huegelii Benfch., and L. Ullardien Cass. 

Tasmania Mt. Dromedary, near Hobart, Dec., 1894, I. (Rod- 
way 1 ). Mt. Direction, Oct., 1895, I., III. (Rodway 1 ). 
Devonport, Jan., 1906 (Robinson). 
Although material was very plentiful, careful search failed to reveal tl 
presence of uredospores, and I have given them 011 the authority 
Cooke. Massee informs me that P. lagenophorae Cooke, is not repi 
sented in the Kew Herbarium, and he, therefore, cannot refer to the tyj 
specimens. I have examined scores of plants with this rust upon thei 
from different localities and at different seasons, and I have never founc 

Tfte leaves on which P. hypochoeridis Me Alp., was found proved to 
not Hypochoeris radicata but Lagenophora billardieri, and the fungus 
agreed with the above. 

(Plate VII., Figs. 54, 55 ; Plate F., Fig. 27.) 

90. Puccinia podolepidis Me Alp. 

0. Spermogonia on small, honey-coloured spots, forming minute, dark- 

coloured points, hemispherical, pale yellow by transmitted ligh 
with round mouth, without projecting paraphyses, about 1 20 diai 
Spermatia minute, hyaline, oval, 3 x 2 p. 

1. Aecidia bright orange, on both surfaces of the leaf, numeroi 

generally arranged in circular groups, -which, however, ofte 
coalesce into large patches ; pseudoperidia cup-shaped, embedded h 
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 p, 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 anc 
thickened at apex (9-12 p), rounded or attenuated towards 
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 oft 
paler, ellipsoid, obovate to wedge-shaped or subclavate, rounded 
thickened at apex, pedicellate, 32-44 x 16-22 p. 
On leaves of Podolepis hngipedata A. Cunn. 

Victoria Buffalo Mts., Nov., 1903, I. (C. French, jun.). Alj 
near Bright, Dec., 1904, I., III. (C. French, jun.). 

Several teleutospores had two germ pores in upper cell, one on eith< 
side, but only one in lower cell. 

The teleutosori were rather sparse. 

(Plate XXIX., Figs. 256, 257.) 

91. Puccinia prenanthis (Pers.) Lindr. 

Lindroth, Myk. Mittheil, p. 6 (1901). 
Cooke, Handb. Austr. Fung., p. 334 (1892). 
Sydow, Mon. Ured. L, p. 106 (1902). 
Sacc. Syll. XVII., p. 306 (1905). 


Puccinia Compositae* 163 

I. Aecidia hypophyllous, a few rarely epiphyllous, or on petioles, seated 

on orbicular or elongated yellow or purple spots, up to 1 cm. in 
diam., in round or irregular groups, at first hemispherical, then 
flat, yellow or whitish, sometimes yellowish purple. 

Aecidiospores globose, subglobose or ellipsoid, delicately warted, 
pale orange, 13-24 /t diam. 

II. Uredosori hypophyllous, on pale indeterminate irregular spots, 

scattered, minute, punctiform, pulverulent, pale brown. 

Uredospores globose or subglobose, echinulate, yellowish-brown, 
16-24 ji 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 p ; pedicels hyaline, very short. 
On living leaves of Lactuca. 

Victoria, New South Wales, S. Australia. 

No specimen seen. It is given on the authority of Cooke, but Mueller 3 
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 

92. Puccinia tasmanica Diet. 

Dietel, Ann. Myc. I., p. 535 (1903). 
Sydow, Mon. Ured. 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 J 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 p. 

Aecidiospores spherical or angular, orange colored, very delicately 
warted, 13-16 /z diam., or 14-17 x 11-15 /*. 

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- 
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 ju, average 54 x 20 ^/, 
occasionally tricellular when 48-73 p long; upper cell deep chest- 
nut-brown, rounded or somewhat oval, conoid, or truncate and 
thickened at apex (up to 13 fj,) 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 

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 yu. 


Puccinia Compositae. 

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. (ftodway '). 

On Senecio peciinafus DC. 

N"ew South Wales Mount Kosciusko, Jan., 1898,1. (Maiden). 

On Senecio brachyglossus F.v.M. 

Victoria August, 1900, I. (Reader). 

On Senecio veUeioides, 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 Me Alp. 

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 p. 

III. Teleutosori minute, black, sparsely developed, intermixed wit 

Teleutospores dark chestnut-brown, oblong to elongated oblong 
clavate, smooth, constricted at septum, generally rounded at base 
and apex, sometimes bluntly pointed or truncate, and thickened at 
apex (6-9 p), 31-46 x 18-25 p, average 34 X 21 p ; upper cell 
darker than lower and similar or sometimes considerably broader ; 
pedicel hyaline, persistent, 30-40 p, long, and up to 9 p. broad 
adjoining spore. 

X. Mesospores very common, similarly coloured to teleutospores and 
stalked, ovoid to elongated ellipsoid, rounded or beaked at apex and 
thicHned (up to 9 p), 25-34 x 14-17 p. 

Puccinia Rubtaceac. 165 

On leaves including leaf stalks of Vittadiuia australis A. Rich. 
Victoria Dimboola, June, 1900 (Header). 

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 
ttnxtralis, 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.) 


94. Puccinia coprosmae Cooke. 

Cooke, Grev. XIX., p. 2 (1890). 
Sydow, Mon. Ured. L, p. 209 (1902). 
Sacc. Syll. IX., p. 300 (1891). 

P. coprosmatis Morrison, Viet. Nat. XL, p. 90 (1894). 

III. Telcutosori hypophyllous, sometimes epiphyllous, rusty-brown r 
prominent, compact, round or elliptic, usually in groups, sometimes 
forming mammilated tubercles, rarely solitary, densely crowded 
and continent, 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 6-8 /j), rounded or 
slightly tapering at base, smooth, constricted at septum, 
35-51 x 16-22 jj, average 43 x 18/i; pedicel hyaline, persistent, 
elongated, up to 110 p long and 8 ^ 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 p. 

On leaves of Coprosma billardieri Hook. 

Victoria Kew and Dandenong, Oct. and Feb., 1893. 
Tasmania (Rod way 1 ). 

On Coprosma hirtella Labill. 

Victoria Murramurrangbong Ranges, Dec. 1903, Jan. 1905 


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. 

Darhica jttum Cast., Avas very plentiful on the teleutosori on Coprosma 
billardieri, and was described by Morrison as a sperrnogone with spermatia. 

(Plate X., Fig. 78.) 


95. Puccinia oliganthae Me Alp. 

II. Uredosori hypophyllous, cinnamon-brown, round to ellipsoid, soon 
naked, girt by the ruptured epidermis, confluent into elongated 

1 66 

Puccinia Rubiaceae. 

TJredospores globose to shortly elliptical or obovate, golden- 
brown, finely echinulate, one to two germ-pores on one face, 
25-29 x 22-25 /u, or 25-27 /* 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 ^), and rounded or bluntly pointed, 
or may be truncate, lower cell generally attenuated towards base 
and paler in colour, 32-54 x 16-21 fi, average 48-16 ^u; pedicel 
yellowish, persistent, up to 48 jj. long. 
X. Mesospores occasional, similarly coloured to teleutospores, somewhat 

fusiform, with pointed and thickened apex, 28-32 X 12-13 /^. 
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 p broad, in P. punctata they are about 20-22 //. 
In the latter too the apex is much thicker, reaching up to 16 jw. The 
species closely resembles P. asperulae odoratae Wurth, but aecidia are 
present on the latter, and the teleutospores are only thickened at the apex 
to the extent of 7 ^u. 

(Plate X., Fig. 77.) 

96. Puccinia operculariae (Morr.) Syd. 


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-biown, oblong to oblong clavate, constricted 
at septum, smooth, 35-54 x 14-20 /it; upper cell rounded or 
pointed and thickened at apex (up to 11 ^u), about equal in length 
to lower ; lower cell usually tapering towards base or sometimes 
rounded ; pedicel hyaline, persistent, 80 p or more long and 6-7 /i 

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 way 1 ). 

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. 

Darluca jilum Cast., is common on the teleutosori, although it is usually 
only found on uredosori. 

(Plate XXIX., Fig. 258.) 


97. Puccinia loranthicola Me Alp. 

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 ^u. 

Aecidiospores ellipsoid to oblong or sub-angular, bright orange 
yellow, decidedly echinulate, 37-56 x 22-31 //. 

II., III. Uredosori amphigenous, brownish, bullate, in scattered 
groups, often arranged in circles and confluent, surrounding 
central darker teleutosori, epidermis splitting lengthwise and 

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 p, occasionally 80 X 22 /z; epispore 3-4J p 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/z, occasionally 3-celled, 
when about 86 ^ long ; pedicel hyaline, persistent, elongated up to 
200 p. long, and swollen towards apex up to 14 p. 

X. Mesospores subhyaline and with elongated pedicel like that of 
teleutospore, smooth, fusoid, hardly thickened at apex, 71-77 x 
25-28 p. 

On living leaves of Loranthus celastroides Sieb., growing on Stringybark 
(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.) 


Puccini a Umbelli ferae. 


98. Puccinia thuemeni (Thuemen) McAlp. 

P. castagnei Thuemen, Rev. Myc. II., p. 86 (1880). 

O. Spermogonia on both surfaces of leaf, pale yellow, round, in groups 
on minute raised yellow spots, about 150 /j. diam. 
Spermatia hyaline, ellipsoid, 3*5-4 x 2-2-5 /z. 

II. Uredosori amphigenous, bullate, round or ellipsoid, scattered or 

crowded, and becoming confluent, surrounded or partially covered 
by the ruptured epidermis, pulverulent, cinnamon-brown, 1 mm. or 
more long. 

Uredospores obovate to ellipsoid, golden-brown, echinulate, with 
thickened hood-like apex (7-8 /*), and three or four, more or less 
equatorial germ-pores on one face, 25-38 x 19-22 p, average 30-32 
x 21 fj, ; the hyaline pedicel may reach a length of 56 ^. 

III. Teleutosori similar to uredosori, dark-brown, also on both surfaces 
of leaf, but more common on under. 

Teleutospores similarly coloured to and intermixed with 
uredospores, shortly oblong to ovate-oblong, slightly or not at all 
constricted at septum, finely warted, rounded at apex, and not per- 
ceptibly thickened, 29-40 x 16-22 /*, average 32 x 20 p ; 
lower cell rounded at base or slightly attenuated ; pedicel 
hyaline, short, deciduous. 
On stems and leaves of Apium prostratum Labill. 

Victoria Beaumaris, Dec.- July, II., III. (III. in April). 
Portland, Jan., 1901, II., III. Sandringham, Jan. and Feb., 

1904, II., III. (III., very sparse). Mentone, Jan.-Sept., 

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 ^t 
in length. There are occasionally elongated teleutospores which reach a 
length of 40 j.t. The finely warted epispore is a constant feature of the 
teleutospores on both hosts. 

The illustrations of spores in PI. 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. 75 
shows the very rough and knobby teleutospores of P. bullata (Pers.) Schroet. 

Puccinia Umbelli ferae. 169 

onAethusa cynapium from Sydow's Ured. Ex.sicc., 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 confix 
The reason for the name I have adopted may first be given. P. cast< 
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. 

As regards the synonyms, Cooke, in his Australian Handbook, gives P. 
apii Corda at first, and then in an addendum substitutes P. castagnei Thuem. 
for it. 

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 ] has shown that the 
host-plant of P. castagnei Schroet., is not Thalictrum angustifolium, but an 
TTmbellifer, 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.) 


Puccinia xanthosiae McAlp. 

II. Uredosori amphigenous, and on leaf-stalks, dark-brown, pulvinate, 

gregarious, elliptical, often confluent, rupturing epidermis, about 
^ rnm. long, often causing entire leaf to become brownish. 

Uredospores golden-brown, thick-walled, eehinulate, globose to 
elliptical, 30-32 ^ diam., or 30-37 x 24-28 p, average 34 x 26 /tx; 
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 p, 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 p. 

On Xanthosia pusilla Bunge. 

Victoria Frankston, Oct., 1899, II. : Oct., 1903, III. Sandring- 
ham, Sept., March, II, III. ; Feb., II. 

(Plate IX., Fig. 76.) 

T y O Puccinia Onagraceae, Rosaceae. 



100. Puccinia epilobii-tetragoni (DC.) Wint. 

Winter, Pilze, p. 214 (1884). 

Sydow, Mon. Ured. I., p. 424, (1902). 

Sacc. Syll. VIL, p. 608 (1888). 

I. Aecidia distributed over the entire surface of the leaf, scattered or 

crowded, hypophyllous ; pseudoperidia cup-shaped, with white, 
laciniate, revolute margins. 

Aecidiospores orange yellow, polygonal, finely warted, 16-26 p. 
diam., 18-21 p. being common. 

II. Uredosori chestnut-brown, scattered, or often orbicularly arranged, 

sometimes confluent, pulverulent, hypophyllous, soon naked. 

Uredospores ellipsoid or ovoid, clear brown, echinulate, with 
two germ-pores on one face, 21-27 x 16-21 p, occasionally 
reaching a length of 31 p. 

III. Teleutosori dark-brown, round, often arranged on ring-like spots, 
hypophyllous, soon naked, pulverulent. 

Teleutospores intermixed with uredospores, elliptic or oblong, 
yellowish-brown, slightly constricted at septum, smooth, 27-36 
x 15-21 p, average 30 x 18 p; upper cell with a cap-like 
thickening at apex (5-6 p) ; lower cell usually rounded at base 
or sometimes slightly tapering ; pedicel hyaline, slender, deciduous, 

X. Mesospores exceedingly rare, similarly coloured to teleutospores and 
thickened at apex (5 p), subclavate to obovoid, smooth, 21-27 x 
13-15 p, with hyaline pedicel. 

On leaves of Epttobium glabellum Forst. 

Victoria Dimboola, Dec., 1892, I., II., III. (Reader). Ardmona, 
Arthur's Creek, Kergunyah, Nyora, Pakenham, <kc., Aug. 
Dec., I., II., III. 
On E. billardierianum Ser. 

Tasmania Great Lake, Feb., 1894 (Rodway l ). 
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 epUobii-Jleischeri Fisch., with aecidia in addition to 
teleutospores, and the present species with the three stages. 

(Plate X., Figs. 79, 80, 81.) 


101. Puccinia gei Me Alp. 


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. 

Puccinia Rosaceae. j^i 

Teieufcospore brownish-yellow, smooth, fusiform to clavate, con- 
stricted at sepfcam, epispore chestnut-brown, 30-40 x 13-18 u, 
average 35 x 14 ^ ; upper cell elongated and tapering or 
rounded, thickened at apex, 18-22 /* long; lower cell elongated 
and tapering towards base, or bulging, 16-22 /z long; pedicel 
pellucid, sometimes 54 p 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 p. 

On living leaves of Geum renijolium F. v. M. 

Tasmania The Calf, Adamson's Peak, March, 1895 (Rod way 1 ). 
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. 

Persoon, Syn. Fung., p. 226 (1808). 
Cooke, Grev. XL, 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 p, or papillate up to 9 /*) 
with spines less prominent, bluntly conical or rounded, 25-38 x 
12-18 jit, occasionally reaching a length of 40 p or more, average 
30 x 15 /j, ; paraphyses intermixed, numerous, capitate, pale yel- 
low, long-stalked, sometimes attaining a length of 70 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 

Teleutospores at first intermixed with uredospores, dark-brown, 
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 p, average 
36 x 25 p ; upper cell usually darker in colour, and broader than 
lower, globose or depressed globose, often slightly thickened 
at apex, and thickly studded with short stout spikes, average 
16-22 p diam.; lower cell generally oblong to obovoid, and equal to 
or longer than upper, 14-24 x 14-19 p ; pedicel short, hyaline, 

On leaves, fruits and stems of Peach (Prunus persica Stokes). 

On leaves and fruits of Almond and Apricot (Prunus amygdalus Stokes, 
and P. armeniaca L.). 


Piiccinia Lcguminosac. 

On leaves of Plum (Primus domestica L.) and Nectarine. 

Victoria, New South Wales (Cobb 1 ), Queensland (Bailey 5 ) 
(Tryon 1 ), South Australia, West Australia, and Tasmania 
(Rod way 1 ). 

Common. November to June, the teleutospores being very common in 
March on Prunus 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 Maire 1 , but 
I 33 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 Prillieux 1 , 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. (McAlpine 34 ). 

On the leaves of the Peach the sori are situated on yellow spots, which 
are 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 z 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 punctatum Pers. 

Darluca filum Cast., is found on the uredosori. 

(Plate X., Figs. 83-86 ; Plate D., Figs. 19, 20.) 



103. Puccinia zorniae (Diet.) Me Alp. 

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^. 

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 p, average 36 x 21 p ; upper cell rounded and thickened 
at apex (up to 6 /u), lower cell rounded at base ; pedicel hyaline, 
persistent, about length of spore. 

Puccinia 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 
Zornia 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 n epispore brown, echinulate, with three germ-pores." 

Darluca Jilum Cast., occasionally on sori. 

(Plate X., Fig. 87.) 



104. Puccinia tetragoniae McAlp. 

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 /j. 

in diam. 

Spermatia hyaline, globose to oval, 2J-3-J- ft 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 ^ diam. 

Aecidiospores orange-yellow, minutely warted, variously shaped, 
globose to ovate or mussel-shaped, 27-39 x 22-25 /.t. 

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 /w, average 31 x 24 p. 

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 
septum, smooth, 40-60 x 25-36 jw, average 50 x 30 ju ; pedicel 
hyaline, stout, persistent, up to 80 ^ long. 

X. Mesospores similarly coloured to teleutospores, ellipsoid or elongated, 
thickened at apex, 35-54 x 25-29 ju. 

On leaves and stems of Tetragonia implexicoma Hook. f. 

Victoria. On the coast at Sandringham, Beaumaris, Flinders, 

Portland, &c., all the year round. 

Tasmania Hobart, about rocks, Aug., 1897, I., II., III. (Rod- 
way 1 ). 

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 


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.) 


105. Puccinia ludwigii Tepp. 

Tepper, Bot. Centralb., XLIIL, p. 6 (1890). 
Cooke, Handb. Austr. Fung., p. 336 (1892). 
Sydow, Mon. Ured. I., 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 J mm. diam. 

II. Uredospores sub-globose to ovate, very finely echinulate, pale 
yellowish-brown, 22-28 x 16-20 yu, 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 ju, 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 yu. 

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 4 ). 

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 
on 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. brownii, 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 
Phragmidium 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. 

Darlucafiium Cast., is common on the uredo and teleutosori. 

(Plates XI., Figs. 90, 91, 92 ; XL., Fig. 300.) 


106. Puccinia muehlenbeckiae (Cooke) Syd. 

Sydow, Mon. Ured. I., 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 p, 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 p), and rounded or obtuse, sometimes 3-4 celled, 32-45 x 
14-19 /i, occasionally reaching a length of 55 p, average 40 X 
1 8 /it ; lower cell generally somewhat triangular ; pedicel hyaline, 
generally persistent, up to 40 p long. 

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 p. 

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) (Bailey 15 ). 
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 
characters with the exception of the septum. 


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 p. Sydow himself had not seen 
any specimens of the rust on JIuehlenbeckia, 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 XI., Fig. 93.) 



107. Puccinia dielsiana P. Henn. 

Hennings, Hedw. XL., p. (95) (1901). 

Sydow, Mon. Ured. L, p. 566 (1902). 

Sacc. Syll. XVII., p. 361 (1905). 

III. Teleutosori on stems, large, compact, bullate, erumpent, up to J cm. 

Teleutospores ellipsoid or oblong ellipsoid, generally rounded at 
both ends or obtuse and slightly thickened at apex (3-5 ^u), smooth, 
very slightly constricted at septum, with granular contents, 
chestnut or dark-brown, 35-18 x 18-28, average 38 x 22 p ; 
pedicel sub-hyaline, persistent, 80-120 x 5-6 /.t. 

X. Mesospores occasional, ellipsoid to ovoid, coloured and thickened at 
apex like teleutospores, 34 x 23 p. 

On steins 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 p. 

(Plate XI.-, Fig. 94.) 

Kochia, Enchylaena. 

108. Puccinia kochiae Mass. 

Massee, Grev. XXII., 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 /z, 32 x 23 p. 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 ^ average 37 x 25 p ; pedicel tinted 
yellowish, persistent, 30-40 x 6 /*, but may reach a length of 96 ju. 

X. Mesospores few, similarly coloured to teleutospores, oval to flattened 
at apex, 28-35 x 22-25 p. 

Puccinia Caryophyllaceae, Malvaceae. 177 

On leaves and stems of Kochia sedifolia F. v. M.; Kochia villosa, Lindl.; 
and Etichylaena 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 teleutosporen 
have occasionally their septum lengthwise. 

(Plate XL, Pigs. 95, 96.) 



109. Puccinia arenariae (Schum.) Schroet. 

Schroeter, Pilz. Schles., p. 345 (187 2). 
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 p), base rounded or attenuated, slightly 
constricted at septum, smooth, pale yellowish brown, 
30-50 x 14-20 p, average 35 x 16 p, very rarely tricellular, 
48 x 16 p j 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 p, generally resembling 
the upper cell of the teleutospore, with elongated pedicel, 22-29 
X 16-17 p. 

On leaves and stems of Stellaria media Cyrill. 

Victoria Gellibrand, Feb., 1896 (Hill). Gembrook 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.) 


Abutilon, Hibiscus. 

110. Puccinia heterospora Berk, and Curt. 

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 jt ; pedicel hyaline, 
slender, elongated, three to four times as long as spore. 

178 Puccinia Malvaceae. 

X. Mesospores much more common than teleutospores, subglobose or 
elliptical, apex more thickened, pale-brown, 17-22 p diam., or 
25-32 x 17-22 p. 

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. 

111. Puccinia malvacearum Mont. 

Montagne in Gay's Hist. Chili VIII., p. 43 (1852). 
Cooke, Handb. Austr. Fung., p. 338 (1892). 
Syflow, Mon. Ured. I., p. 476 (1903). 
Sacc. Syll. VII., 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 
thickening slight, smooth, yellowish-brown, rarely tricellular, 
35-75 x 12-26 p, average 50 x 17 ^; pedicels firm, long, per- 
sistent, occasionally septate, sometimes measuring 170 p 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. sylvestris L., Althaea 
rosea Cav., Lavatera plebeia Sims, and Plagianthus spicatus Benth. Ex- 
ceedingly common. 

Victoria (Berkeley 2 ) . 

New South Wales (Sacc. and Berl. 1 ). 

Queensland (Bailey 7 ). 

South Australia. 

West Australia (Morrison). 

Tasmania (Rod way l ). 

The sori first appearing were carefully examined for any trace of 
accompanying spermogonia, bulb none were found. Spores occasionally 
3-celled, and reaching a length of 77 p. 

This fungus, which was originally described by Montagne, has now 
overspread the earth and occurs on both wild and cultivated species of 
Malvaceous plants. Dr. Plowright 5 states that he has a specimen in his 
herbarium from Melbourne, gathered as early as 1865 on Malva 

Puccinia Malvaceae, Geraniaceae. 179 

Puccinia malvacearum 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 Me Alp. 

McAlpine, Proc. Roy. Soc. Vic. VII, N.S., p. 218 (1894). 
Sydow, Mon. tired. 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 at 
septum, very rarely three celled, epispore with reticulated markings, 
38-50 X 17-23 p, average 41 X 20 /u ; pedicel hyaline, short, 
deciduous, but may attain a length of 60 p. 
On leaves and flowers of Plagianthus sidoides Hook. Very common. 

Tasmania Mount Wellington, southern slope, Aug.-April (Rod- 
way 1 ). 

The spore is described as smooth by Sydow, but it is seen to be covered 
with a fine irregular network. 

(Plate XII., Fig. 98.) 



113. Puccinia geranii-pilosi McAlp. 

II. Uredosori hypophyllous, chocolate-brown, scattered or sub-gregar- 

ious, orbicular, soon naked and surrounded by ruptured epidermis, 
compact, up to 1 mm. diam., often on purplish-red spots or portions 
of leaf becoming coloured. 

Uredospores yellowish-brown, elliptical to obovate, echinulate, 
epispore slightly thickened at base, 25-30 X 19-22 ^. 

III. Teleutosori elongated, dark-brown to black, sometimes confluent, 
erumpent, compact, 1-1 J /< long, commonly on petiole. 

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 //, 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 ^t long. 


Puccini a Geraniaceae. 

X. Mesospores occasional, similarly coloured, subglobose, thickened at 
apex, 25-26 fj. diam. 

On living leaves of Geranium pilosum Sol. 

Victoria Killara, Nov., 1902, II., March, 1903, II., EIL (Robin- 
New South Wales Richmond (Musson). 

Occasionally a teleutospore occurs in which the septum is vertical as in 

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. 

I. Aecidia amphigenous, delicate, minute, orange, circinate, either alone 

or accompanied by teleutospores ; pseud operidia with expanded and 
lobed margin ; peridial cells subquadrate to oblong with striated 
margin, and punctate all over, firmly united, 28-32 x 16-22 p. 

Aecidiospores pale orange, finely echinulate, elliptic to subquad- 
rate, 19-22 x 16 p, or 18-19 p 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 p diam., or 22-27 x 19-22 p. 

III. Teleutosori intermixed, solitary or arranged in small confluent 
groups, generally round, black, compact, surrounded by ruptured 

Teleutospores chestnut brown, clavate, constricted at septum, 
smooth, often obliquely pointed or rounded and thickened 
at apex (up to 9 p), 33-64 x 19-25 p, average 45 x 22 p ; 
lower cell usually paler in colour and attenuated towards base ; 
pedicels tinted, generally short, persistent, up to 3Sp long by 6-9 p 

X. Mesospores coloured like teleutospores, oblong to oval, bluntly 
pointed and thickened at apex, 33-38 X 16-20 p. 

Puccinia Rutaceae. 181 

On stems, leaf -stalks and leaves of Pelargonium australe Jacq. 

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 Murrarnurangbong 

It differs from P. geranii Corda, to which this species was referred by 
ooke in Grevillea, XXI., p. 39, in the uredospores being nearly twice a* 
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. 
granularis Kalch. and Cooke. 

In P. pelargonii the aecidia are hypophyllous, and not amphigenous, and 
tho teleutospores are rather shorter and narrower. In P. granularis there 
are no aecidia. 

(Plate XII., Fig. 102 ; Plate F., Fig. 29.) 



115. Puccinia boroniae P. Henn. 

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 ft), gen- 
erally rounded at base, 28-40 x 16-21 /u, average 35 x 20 /x, 
occasionally 3-celled and elongated, 52 x 25 /* ; pedicel hyaline to 
yellowish, persistent, up to 120 p 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 p. 
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 /x, by Sydow. 22-35 
x 16-19 ^u, and I have made the average 35 x 20 ^, my measurements 
being corroborated by those in the photographs, which are also about 
35 x 20 f.i. 

(Plate XIII., Fig. 107.) 


116. Puccinia correae Me Alp. 

McAlpine, Proc. Roy. Soc. Viet. 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. 


Puccinia Rutaccae, Tremandraceae. 

Teleutospores yellowish-grey, long-stalked, elongated fusoid, con- 
stricted at septum, 44-60 x 17-20 p, average 50 x 18 p; upper 
cell elongated, tapering and rounded at apex ; lower cell tapering 
towards base ; pedicel light grey, several times as long as 

X. Mesospores similarly coloured and stalked, ovoid, with truncated 
apex, 25-28 x 16-19 p. 

On leaves of Correct, lawrenciana Hook. 

Tasmania Mount Wellington, Dec. Coast near Trial Harbour, 
Jan., 1894 (Rodway 1 ). 

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. Although the plant is very common, 
the rust was very rare. 

(Plate XIII., Fig. 106.) 


117. Puccinia eriostemonis Me 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, 30-15 x 15-18 p, average 32 x 16 /*; 
pedicel persistent, hyaline, elongated, up to 125 p long by 3-5^ p 

X. Mesospores occasional, similarly coloured to teleutospores, ellipsoid, 
thickened and bluntly pointed at apex, 30-32 x 13-16 p. 

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., in the sori being orange 
instead of brown, the spores are smaller, with finely granular orange con- 
tents, and the.epispore rather thicker. 

(Plate XIII., Fig. 105.) 

118. Puccinia pritzeliana P. Henn. 



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 
purplish margin, scattered or in orbicular groups, pulvinate, 
cinnamon- brown, J-1J mm. diam. 

Puccinla Violaceae. 

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 p, 
average 48 X 20 p ; pedicels hyaline, persistent, up to 150 p long. 
X. Mesospores rare, similarly coloured to teleutospores, somewhat 
oblong and bluntly apiculate at apex, 43 x 28 p. 

On living leaves of Tremandra stelligera B. Br. 

West Australia Near Perth, Oct., 1900 (Pritzel). 

Quite distinct from P. tremandrae Berk, from Ceylon. 

(Plate XIII., Fig. 108.) 



119. Puccinia hederaceae McAlp. 

0. Spermogonia arranged in small or large groups, crowded, honey- 


Spermatia hyaline, globose, minute, 2J-3 p. diam. 

1. Aecidia 011 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 J mm. diam. with white, torn, 
re volute edges. 

Aecidiospores subglobose to shortly elliptical, finely warted, 
orange-yellow, about 16 p diam., or 16-19 x 12-14 p. 

II. Uredosori minute, brown, roundish, erumpent, scattered, soon 

naked, on both surfaces of leaf. 

Uredospores subglobose or shortly elliptical, golden-brown, 
strongly echinulate, 2-3 germ-pores on one face, 25-29 X 18-22//, 
average 25 x 21 p and 22-25 p. 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 p, average 35 
X 19 p - } pedicel hyaline, deciduous, short. 
On living leaves of Viola hederacea Labill. 

Victoria Near Melbourne, Oct., Nov., and Dec., 1885, 1886 
(Beader). Wandin, .Dec., 1895. Murramurrangbong Banges, 
Nov., 1902, Dec., 1903, Jan., 1905 (Bobinsony. Mt. Black- 
wood, Dec., 1902. Alps, near Bright, Dec., 1904 (C. 
French, jun.) 

Tasmania Cascade Bivulet, Dec., 1892 (Bodway '). Mt. Drome- 
dary, Dec., 1894 (Bodway 1 ). Devonport, Jan., 1906 
On V. betonicifolia Sm. 

Tasmania Ben Nevis, April, 1896 (Bodway). 

New South Wales Monga (Bauerlen), recorded as Aecidium 

violae Schum. by Baker 1 . Bichmond (Musson). 
Victoria Murramurrangbong Banges, Nov., 1902 (Bobinson). 

438. G 

1 84 

Pucdnia Crticiferae. 

A few one-celled teleutospores were observed among the ordinary 
teleutospores, which they resembled in every respect except size and shape, 
even having the germ-pore quite distinct beneath the apex, ellipsoid, with 
small hyaline apiculus, 25-26 X 20 /u. 

Aecidium violae Schum. is given by Dr. Cooke in his Handbook as 
occurring on Victorian native violets, but it is the aecidium belonging to 
P. hederaceae Me Alp. 

This species was at first determined as P. aegra Grove by Winter 2 , 
but on comparing the two species they are seen to be decidedly different. 

It is closely allied to P. violae (Schum.) DC., but differs in the 
teleutospores being larger, and warted all over. Specimens of P. violae 
were examined from Syd. Ured. Exs., No. 286, on Viola arenaria, and from 
Kellerman's Ohio Fungi, No. 96, on Viola blanda. The average size of the 
teleutospores was found to be 22-32 x 16-19 /u, and were generally 
smooth, although occasionally a little warted. 

P. aegra Grove, is recorded by Cooke for Australia; but all the speci- 
mens I have met with, including part of the same collection of material 
forwarded to Dr. Cooke, have echinulate aecidiospores, and warted 
teleutospores, and are therefore not this species. 

(Plate XIII., Figs. 109-112 ; Plate E., Figs. 24, 25.) 

120. Puccinia cruciferae McAlp. 

I. Aecidia anaphigenous and on stems, in irregular clusters or circinat 
large, cup-shaped, J-J mm. diam.; pseudoperidia with whitisl 
edges, crenulate, slightly revolute ; pseudoperidial cells ovoid 01 
oblong, punctulate all over, with striated margin, average 25 

Aecidiospores orange, subglobose, finely echinulate, 14-16 

III. Teleutosori on upper surface of basal leaves, associated wit 
aecidia, a few together, round, bullate, compact, covered at fii 
by raised grey epidermis, which splits irregularly, up to 1 mi 

Teleutospores black in mass, dark chestnut-brown individual!] 
oblong to clavate, smooth, slightly constricted at septum, bluni 
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 p. long and 
10 p broad. 

X. Mesospores common, similarly coloured to, or paler than, teleut 
spores, variously shaped, ellipsoid, oblong or ovate, point 
rounded, or truncate at apex, and thickened up to 7 p, 
25-34 x 14-20 p, with pedicels equally long and similar to those 
of teleutospores. 
On Cruciferous plant. 

New South Wales Guntawang (Hamilton). 

It differs from P. cruciferarum Rud., in having aecidia, and fi 
P. barbareae Cooke, in which the teleutospores are broadly lanceolate ai 
considerably narrower. 

Phragmidium. 185 

This is a portion of the original material which was named Aecidium 
bai'hareae in Cooke's Handbook of Australian Fungi 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.) 


121. Pnccinia hibbertiae Me Alp. 

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 ^ ; upper cell 
rounded at apex to somewhat conical, usually thickened (up to 
6 //) ; lower cell generally rounded at base, sometimes slightly 
attenuated ; pedicel hyaline, persistent, elongated, slender, 
120-150 p long. 

X. Mesospores occasional, similarly coloured to teleutospores, generally 
fusiform and thickened at apex, sometimes elongated oblong, 
33-35 x 12-16 //. 

On Hibbertia sericea Benth. 

Victoria Cheltenham, July, 1904 (Robinson). Oakleigh, Sept., 

The small size of the leaf together with the scanty sori render this 

species easily overlooked. The teleutospores are often noticed germinating 
in situ. 

(Plate XXX., Kg. 265.) 


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. barnardi) on the native raspberry (Rubus parvifolius) does considerable 
damage, and the imported species P. subcorticium is spreading extensively 
on the sweetbrier. 

><ral Characters. Spermogonia flattened or slightly raised, generally 
arranged in concentric circles. 

Aecidia after caeoma type, surrounded by a dense crown of inwardly- 
curved paraphyses. 

Caeomospores with numerous distinct germ-pores distributed over the 
entire surface. 

I ^redosori 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-pores, 
the others about three to four each, placed laterally. 

Sporidiola spherical. 

Australian species, 4. 

186 Phragmidium Rosaceae. 


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 uppe 
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 p. 

TI. 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 
in colour. 

Uredospores globose or ellipsoid, epispore thick, warty, dai 
yellow, 17-19 p diarn. or 21-26 X 17-19/u; paraphyses surroi 
ing them, at first colourless, finally yellowish-brown, variously ai 
peculiarly shaped, generally curved and pointed at apex, 30-40 
9-12 p. 

III. Teleutosori scattered or gregarious, minute, dark brown, n< 
rarely confluent, pulverulent. 

Teleutospores at first intermixed with uredospores, golden-browi 
cylindrical, equal, rounded at apex, sometimes with a shoi 
hyaline, conoid apiculus, 6-9 septate (8 very common), constrict 
at septa, smooth, usually two germ-pores on one face in each 
or three altogether, 60-115 x 23-28 p ; pedicels hyaline, cyl 
drical, or a little inflated, sometimes uniseptate towards base, vei 
long, up to 207 x 10-19 p. 

On living leaves of Native Raspberry (Rubus parvifolius L.). 

Victoria Alps near Bright, Arthur's Creek, Flinders, Kill 

Murramurrangbong Ranges, Myrniong, near Melbourn 

Phillip Island, Seymour, &c. 
South Australia Mount Lofty (Tepper 2 ). 
Tasmania Huonville, Jan., 1892 (Rodway 1 ). Mount Wellin 

Jan., 1892 (Rodway). 

As early as 1886 this species was determined by Plowright and Win 
from specimens sent by that veteran microscopist, Mr. F. Barnard, K 

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 anpl 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 n 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 

Phragmidium Rosaceae. 187 

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 p 

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 longlssimum Thuem. 

Thuemen, Flora, p. 379 (1875). 

Cooke, Handb. Austr. Fung., p. 340 (1892). 

Sacc. Syll. VII., p. 750 (1888). 

Hamaspora longissima Koern. Hedw. XVL, 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 ^ 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 fj.', 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 /.i. 

On leaves of Rubus moluccanus L. 

Queensland Eudlo Creek (Bailey 9 ). 

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 

Scattered over the leaves are groups of silky-looking twisted filaments, 
and OH 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 ft diam. 

The teleutospores were very commonly 140 /< long, and one 6 -celled in- 
dividual attained a length of 190 p. 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 /<, then Massee reduced it to 
70-130 n, and in the Queensland specimens I have found it to be 110-190 p. 
The stalks are excessively long, and they may reach a length of 500 f.i 
or more. 

The teleutospores agree with those of Phragmidium in being three or 
more transversely septate, but they are agglutinated together into a 
gelatinous cylindrical filamentous mass, and each cell only possesses one germ 

1 88 

Plirag midium Rosaceae . 

Massee 3 from an examination of fresh material comes to the conclusion 
that it is not a Phragmidium nor a ffamaspora, although he does not under- 
take the responsibility of forming a new genii?. 

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 Gymnosporangium in the arrangement and structui 
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). 
Cooke, Handb. Austr. Fung., p. 339 (1892). 
Sacc. Syll. VII., p. 743 (1888). 
[I. 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 p.] 

II. Uredosori hypophyllous, orange-red, roundish, scattered 01 
gregarious, often confluent, surrounded by a crown of club-shaj 

TJredospores spherical, elliptical or ovate, yellowish, echinulat 
17-24 x 14-20 p, average 18 x 15 p. 

[III. Teleutosori orbicular, black, pulvinate, minute. 

Teleutospores oblong, 2-6 septate (usually 3-5), slightly coi 
stricted, obtuse, or obtusely apiculate above, slightly attenuat 
below, smooth, yellowish-brown, 50-70 X 20-22 p ; pedicel 
colourless, firm, 100-150 p, long.] 

On living leaves of Acaena sanguisorba Yahl. 

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, and 
portion of the same material, kindly supplied by Reader, shows only 

Darlucafilum Cast, is plentiful on some of the sori. 

125. Phragmidium subcorticium (Schrank) Winter. 

Winter, Die Pilze L, p. 228 (1884). 
Cooke, Handb. Austr. Fung., p. 339 (1892). 
Sacc. Syll. VII., p. 746 (1888). 

I. 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 14-20 yu. 

Cronartium. 189 

II. Uredosori hypophyllous, on yellow spots, yellowish red, minute, 

orbicular, scattered or crowded, with tubular paraphyses slightly 
swollen towards apex, and not particularly club-shaped. 

Uredospores ellipsoid, ovoid or piriform, finely echinulate, 
reddish orange, 4-6 germ-pores on one face, 22-29 x 16-20 /u. 

III. Teleutosori hypophyllous, minute, scattered or crowded, black, 
readily detachable. 

Teleutospores oblong, dark-brown, warted, obtuse at the apex 
with a colourless apiculus, 7-8 celled, with 2-3 germ-pores on one 
face in each cell, 75-102 x 32-36 p. ; pedicel hyaline, except 
pale-brown towards apex, expanded in the lower half, 100-150 ju 
long and broadened up to 22 p. 

On Rosa rubiginosa L., R. canina L., and R. laxa Retz. 

Victoria Pascoe Vale, Oct., 1898 (Cronin). Ferntree Gully, 

April, 1899 (Hill). Brighton, Nov., 1901, Aug., 1903, June, 

Oct. and Dec., 1904, March, 1905, II. 
S. Australia Third Creek Garden, Mt. Lofty Range, Nov., 1899 

(Tepper). Nov., 1900 (Agricultural Bureau), II., III. 
Tasmania Launceston, Oct., 1902 (Littler). Hobart, Apr., 1903, 

II., 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 caeorna may winter in the stems, so that by means 
of cuttings the disease was probably introduced into Australia. It 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 L, Fig., 37.) 


Only one species of this genus is known here, and it occurs rather com- 
monly on certain leguminous plants. So far teleutospores only have been 
found. This genus produces uredo and teleutospores on one host plant, 
and the aecidial stage on another, but the latter has not yet been 
found in Australia. In Europe there is a bladder-rust on the bark of JPMMM 
sylvestris, and this has been proved to be the aecidial stage of Cronart 
asclepiadeum (Willd.) Fr. As in so many other similar cases, this stage wu> 
considered to be an independent form, and named Peridermium cornui 
Rostr. and Kleb. 

General characters. Sperm ogonia truncate to conoid. 

Aecidia with bladder-like, irregularly opening pseudoperidia (Pet^ider- 

Aecidiospores without germ-pores, separated by distinct intermediate 
cells, epispore always more or less regularly warty reticulated. 

Uredo-layer included in hemispherical pseudoperidum, which opens at apex 
by a narrow pore. 

i o,o Cronartium Leguminosae. 

Uredospores echinulate, without germ-pores. 

Teleutospores unicellular, united into a cylindrical, vertical, horny coluirn 
germinating in situ, surrounded at base by the uredospores. 
Sporidiola subglobose, hyaline. 
Australian species, 1. 


Jacksonia, &<. 

126. Cronartium jacksoniae P. Henn. 

P. Hennings, Hedw. XL., p. 127 (1901). 

O. Spermogonia honey- coloured, crowded or in lines, circular, flattene 
on branches and lower surface of leaves. 

Spermatia hyaline, ovoid to ellipsoid, 5-6 X 2J-3 p. 
III. Teleutosori forming ruddy-brown, straight or curved, simplt 
columns, crowded, 2-2 J 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 
end, with large vacuole, cylindrical to fusiform, contents granular, 
40-50 X 8-10 JJL ; teleutospores germinate at once and produce 
promycelia with promycelial spores, which are hyaline, globose, 
and average about 6-7 /z diam. 
On Jacksonia scoparia R. Br. 

Queensland Darling Downs (Law), (Bailey 3 ). Near Brisbai 
(Bailey 4 ). 

On Platylobium formosum Sm. 

Victoria Murramurrangbong Ranges, Nov. and Dec., 1902-{ 

On Gompholobium lati folium Sm. 

Victoria Murramurrangbong Ranges, Dec., 1903 (Robinson). 

On Bossiaea cinerea R. Br. 

Victoria Near Melbourne, Nov., 1904 (Robinson). 
Tasmania Bellerive, Dec., 1905 (Rodway). 

On Aotus villosa Sm. 

Victoria Near Melbourne, Sept., O. 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 is 
so common in some parts that the great majority of the shrubs 

The spermogonia were only found on Aotus and before the appearanc 
of the teleutospores, but the multiplication and deformation of the shoot 
had already been produced. On Bossiaea cinerea on one occasion Aecidiu 
eburneum McAlp. was found on the same plant near Melbourne. 

The horn-like aggregation of teleutospores varies in length from 1-2 mi 
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 plants, and the germinating 
teleutospores were clearly seen. 

Melampsora Hypericaceae. i 9I 

Considering its occurrence on indigenous Leguminous plants, together 
with the narrower teleutospores and colourless promycelial spores, as com- 
pared with C. asdepiadeum, I considered it a new species and had named it 
C. leguminum. But I find that Hennings in a footnote in Hedwigia XL., 
p. 127, states that the form occurring on Jacksonia is specifically distinct 
from C. asclepiadeum, and had better be named C. jacksoniae, the name now 



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. amygdcdinae 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 

General description. Spermogonia forming a minute, hemispherical, 
flattened stratum, often subcuticular, otherwise subepidermaL 

Aecidia after the eaeoma 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 

Australian species, 2. 



127. Melampsora hypericorum (DC.) Schroet. 

Schroeter, Brand und Rost., p. 26 (1872). 
Sacc. Syll. VII., p. 591 (1888). 

II. Uredosori mostly hypophyllous, scattered or subgregarious, at first 

bright orange, becoming pale, pulverulent, small, up to J 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 ^, 
average 17 X 1 4 p ; 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- 

Teleutospores yellowish-brown, intercellular, subcylindrical or 
prismatic, 25-37 + 6-9 p. 
On leaves and occasionally on stems of Hypericum japonicum Thunb. 

Victoria Murramurrangbong Ranges, Nov. and Dec., 1902-3, and 
Mt. Blackwood, Jan., 1903 (Robinson). Buffalo Mts. and 
Alps, near Bright, Nov. and Dec., 1903-4 (C. French, jr.). 
Near Melbourne, Nov., Jan. 


Melampsora Linaceae. 

The paraphyses are noted as absent by Dr. Plowright in his Monograph 
of the Uredineae, but in these specimens they are particularly abundant, and 
form a striking feature of the uredosori. Fischer 5 also found the paraphys 
wanting, and he came to the conclusion with others, that the caeonaa form 
had been mistaken for the uredo, since the spores were in chains. But 
Klebahn 3 has proved the existence of both the caeoma form without para- 
physes and the uredo form with paraphyses, so that the three stages may 
occur on species of Hypericum. Aecidium disseminatum Berk, is found hex 
on the same host-plant and in the same locality as this species. 



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). 

II. Uredosori scattered, rounded, orange, soon pulverulent, up to 1^ 

mm. diam. 

Uredospores round or ovate, bright orange-yellow, echinulate, 
pedicellate, 15-25 x 13-18 p, exceptionally reaching a length of 
28 fji paraphyses curved, markedly capitate, 17-20 ^t thick. 

III. Teleutsori flattened, often confluent, reddish brown, then black, 

Teleutospores densely crowded beneath epidermis, cylindrical] 
prismatic, intercellular, polygonal in section, 45-60 x 17-20 //, 
very occasionally two-celled. 

On leaves and stems of Linum, marginals A. Cunn. Widely distributed. 

Victoria Near Melbourne, Oct., 1885 (Reader). Ardmona, 

Kergunyah, Killara, near Melbourne, Rutherglen, &c., Oct., 


New South Wales Hume River, 1886 (Jephcott). Guntawan< 

Mudgee (Hamilton); Merilla, Oct., 1890 (Cobb 2 ). 
South Australia Murray River, 1890 (Tepper), (Ludwig 2 ). 
Tasmania Near Waterworks, Hobart, Jan., 1892 (Rodway 1 ). 
On Linum usitatissimum L. wherever flax is cultivated. 

Victoria Near Melbourne, 1902. Donald, Nov., 1903, II., III. 

Port Fairy, Jan., 1904, II., III., &c. 
South Australia Near Adelaide, 1889 (Crawford). 
New South Wales Brungle, 1891 (Cobb 4 ). 

In the Journal of Mycology for 1889, Galloway 1 records this rust 
being sent to him from South Australia by the late Frazer S. Crawford, w] 
wrote that it had destroyed a crop of flax near Adelaide, and was likely 
spread and prove a troublesome pest. Bolley informs me by letter thai 
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 flax-sick soil disease. 

Dr. Cobb 4 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. 193 


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, Aecidium, and Uredo, and of the former there are wnlv 
two known for Australia, one of which was called an Aecidium by Cooke 
and the other a Uredo by Berkeley. This is not surprising when one 
considers that the caeoma is not distinct from the aecidium, 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. Barclay 3 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 Aecidium and Caeoma. In 
the aecidial stage of Puccinia aucta (Aecidium lobeliae 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- 
^atenulate 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 

Caeoma-forms, 2. 


Tabernaemon 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 p. ; epispore hyaline, coarsely verrucose, 5 p. thick. 

On Tabernaemontana orientalis B. Br. 
Queensland Brisbane, (Bailey 1 ). 

Mr. Bailey kindly sent me a specimen of this from his herbarium, 
which had been named by Dr. Cooke Aecidium 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 (Aecidium) apocynatum 
Schwein., it is constituted a new species. 

i 9 4 

Aecidium Scrophulariaceae. 


130. Caeoma clematidis Thuem. 

Thuemen, Myc. Univ. No. 539. 
Berkeley, Hook. Journ. VI., p. 205 (1854). 
Cooke, Handb. Austr. Fung., p. 344 (1892). 
Sacc. Syll. VII., p. 867 (1888). 
Uredo clematidis Berk. 

I. Sori hypophyllous, generally on pale-green portions of leaf, solitary or 
arranged in circular groups, more or less rounded, pulvinate, ruddy- 
brown at first and bright orange when burst. 

Spores in rows, bright orange, variable in shape, ellipsoid, 
ovate, oblong or polygonal, epispore hyaline, scolloped, 22-32 x 
16-20 p. 

On living leaves of Clematis aristata R. Br., and C, microphylla DC. 

Victoria Near Colac, Sept., 1897 (Hill). Phillip Island, Jan., 
1900. Murramurrangbong Ranges, Dec., 1903 (Robinson). 
Near Melbourne, Feb. and July, 1904. Frankston, Jan., 1904, 
Aug., 1904 (Robinson). 

Queensland Bailey ' 5 . 

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. 


Several of the aecidia given in Dr. Cooke's Handbook oj 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 Me Alp., 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 revolute. 

Spores, globose or angular, commonly orange yellow, growing in chains. 

Aecidium-forms, 15. 



131. Aecidium disciforme Me Alp. 

O. Spermogonia on under surface of upper and younger leaves, honey- 
coloured, flat, numerous, minute. 

Spermatia hyaline, elliptical, 5-6 p. 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 J mm. ; 
peridial cells hyaline, loosely adherent, subglobose, elliptic or 
lozenge-shaped when united, with striated margin, 25-30 p. 

Aecidiospores orange-yellow, subglobose to elliptic, 16-20 u. 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. 

Berkeley in Herb., Grev. XI., p. 97 (1883). 

Sacc. Syll. VII., p. 814 (1888). 

I. Aecidia numerous, crowded, semi-immersed, on under-surface of leaf 
arid a few scattered on upper surface, J 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 2o p. 

Aecidiospores dark yellow, variously shaped, ellipsoid, oblong or 
angular, with finely -granular contents, 24-30 x 16-20 ju ; epispore 
hyaline, finely verrucose, about 2 p. broad. 

On Veronica sp. 

Victoria (F. v. Mueller). 
Tasmania ( Rod way) . 

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 

(Plate XXXIX., Fig. 287.) 


133. Aecidium plant aginis-variae McAlp. 

0. Spermogonia on both surfaces of leaf, but mostly on upper, honey- 

coloured, numerous, 160-180 p, diam. 
Spermatia hyaline, elliptical, 4-5 p. 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 p long. 

Aecidiospores bright orange-yellow, elliptic to oblong, finely 
echinulate, 22-25 x 16-23 p, sometimes reaching a length of 28 p. 

196 Aecidium Genticwaccae, Compositae. 

On leaves of Plnntagn vnria R. Br. 

Victoria Murramurrangbong Ranges, Nov., 1902 (Robinson). 

Mt. Black wood, March, 1905 (Cyril Brittlebank). 
New South Wales Guntawang, near Mudgee (Hamilton). 
Tasmania Knocklofty, Oct., 1895 (Rodway 1 ). 

This species is given as A. plantaginis 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 
Darlucafilum Cast. 

(Plate XXXIX., Fig. 288.) 



134. Aecidium nymphoidis DC. 

De Candolle, Fl. fr. II., p. 597 (1805). 
Cooke, Handb. Austr. Fung., p. 341 (1892). 
Sacc. Syll. VIL, 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 
ji diam. 

On Limnanthemum indicum Thw. 

Queensland Near Brisbane, 1884 (Bailey 4 ). 

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 Chodat 1 . Then Bubak 1 , by means of cultures, proved the 
connexion, but until the teleuto stage is found in Australia only the 
aecidium stage will be given. 

Specimen not seen. 


135. Aecidium cymbonoti Thuem. 


Thuemen in Muell. Supp. Phyt. Austr. XI., p. 96 (1880). 
Sacc. Syll. VII., p. 833 (1888). 

0. Spermogonia minute, honey-coloured, in groups, surrounded bj 

aecidia, 130-160 ju diam. 

Spermatia minute, hyaline, ellipsoid, 3 fj. long. 

1. Aecidia on upper surface of leaves, markedly circinate, about J mm. 

diam.; pseudoperidia yellowish, with fimbriate margin ; peridial cells 
considerably overlapping each other, elongated oblong to slipper- 
shaped, with striated margin and reticulate all over, 28-38 X 
16-17 p. 

Aecidiospores orange-yellow, ellipsoid to sub-globose, finely 
echinulate, 13-18 x 13-15 p. 

Aecidium Compositae. 197 

On Cymbonotus lawsonianus Gaudich. 

Victoria Dookie, Jan., 1892 (Pye). Dimboola, May, 1896. 
(Reader). Preston, April, 1900 (C. French, jun.). Myrniong 
(C. C. Brittlebank). 

Some of the specimens were overrun with Darluca filum Cast. 
There is no clue to the description of this species, but I have drawn out 
the above from abundant material. 

(Plate XXXIX., Fig. 290.) 


136. Aecidium monocystis Berk. 

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 ^. 

Aecidiospores pale orange-yellow, ellipsoid, finely verrucose, 
25-30 x 19-22 p. 

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 tine 
leaves. It was first recorded on Phyllachne (Helophyllum) 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. forsterioides, 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.) 

01 c 
137. Aecidium oleariae Me Alp. 

I. Aecidia crowded on steins 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 p. 

On stems and leaves of Olearia axillaris F.v.M. 

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. 


Aecidium Rubiaceae, Leguminosae. 



138. Aecidium plectroniae Cooke. 

Cooke, Grev. X., p. 124 (1882). 

Bailey, Queensland Agr. Journ. IV., p. 284 (1899). 

Sacc. Syll. VII., p. 795 (1888). 

0. Spermogonia on upper surface of same spots which bear the aecidif 

sometimes accompanied by one or a few aecidia. 

1. Aecidia hypophyllous, rotund, in small clusters on orbicular dis- 

coloured spots ; pseudoperidia somewhat prominent, whitish, mar- 
gins sub-entire ; peridial cells ovate to somewhat quadrate, 
punctate all over, and with striate margin. 

Aecidiospores yellow, finely echinulate, sub-globose to ellipsoid 
or polygonal, 25-26 x 19-20 /*, or 22-28 p diam. 

On leaves of Canthium coprosmoides F. v. M. 

Queensland Endeavour River (Roth) (Bailey, 31 - 22 ). 

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. 


139. Aecidium eburneum McAlp. 

McAlpine, Proc. Roy. 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 ii 
dense clusters, bright orange, average J mm. diam.; pseudoperidia 
cup-shaped, becoming ivory-coloured then brownish, with reflected 
finely toothed margin ; peridial cells polygonal, reticulated, 24-31 

Aecidiospores ellipsoid, bright orange, very finely echinulate, 
18-28 x 12-17 fi. 

On Bossiaea cinerea R. Br., stems, leaves, flower-stalks, calyx, ant 

Victoria Near Melbourne, Sept.-Nov. (Barnard, Robinson, &c.). 
Tasmania Bellerive swamp (Rodway !), 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 of which 

Aecidium Leguminosae. 199 

he kindly sent me. The spores in both cases are very finely echinulate, and 
in the West Australian specimens they are 18-23 x 15-18 p, 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.) 


140. Aecidium platylobii Me Alp. 

0. Spermogonia honey-coloured, minute, scattered, either associated 

with or on opposite side to aecidia. 

Spermatia very numerous, hyaline, cylindrical, 3 x 1 p. 

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 11. 
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 soleniiforme 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, having 
hitherto only been found on the first or second pair of leaves of young 

The mycelium bearing the aecidia penetrated the fruit cover and entered 
the seed, so that the young plant'is probably infected from the start. 

(Plate XXVII., Figs. 237, 238.) 


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 
J mm., white, radiately laciniate at margin. 

Aecidiospores rather angular, orange, 25-26 fi diam. 
On leaves and fruits of Goodia lotifolia Salisb. 

Victoria Nov., 1895 (French). Gellibrand River, Dec., 1895 


Tasmania 1860 (Archer). (Berkeley 1 ) 

The aecidia, as the specific name denotes, are rather tubular at first. 
(Plate XXXIX., Fig. 293.) 

200 Aecidium Amarantaceae, Hyp eric aceae. 



142. Aecidium deeringiae Cooke and Mass. 

Cooke and Massee, Ann. Rep. Dept. Agr. Queensland, p. 40 

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 ^ 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 ^ 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.) 



143. Aecidium disseminatum Berk. 

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 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 p.. 

On Hypericum japonicum Thunb. 

Victoria Murramurrangbong Ranges, Nov. 1902, and Dec., 1903 

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 



144. Aecidium calthae Grev. 

Greville, Fl. Ed., p. 446 (1824). 

McAlpine, 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, about 

J mm. diam. ; pseudoperidia white, margin laciniate ; peridial 
cells pentagonal or hexagonal, margin finely streaked, 28-32 p. 

Aecidiospores bright orange, angular to sub-globose or oval, finely 
echinulate, average 22-23 x 17-20 p. 
On living leaves of Caltha introloba F. v. M. 

New South Wales Mt. Kosciusko, Jan., 1899 (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.) 


145. Aecidium ranunculacearum DC. 

De Candolle, Fl. fr. VI., p. 97 (1805). 
Cooke, Handb. Austr. Fung., p. 340 (1892). 
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 p. 

Aecidiospores orange-yellow, polygonal to oblong or ellipsoid, 
very finely echinulate, 17-30 x 14-20 p, average 22 x 17 p. 
On Ranunculus parvi/lorus 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), (Berkeley 1 ). 
This aecidium occurs on several species of Ranunculus, and varies some- 
what in general characters, although the aecidiospores all agree in being 
very finely echinulate. As the result of cultures, this has been found to be 

2O2 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 filum 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. 

General characters. Seri 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. 



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. 

Uredospores globose to ellipsoid or piriform, brown or orange, 
rather thin-walled, echinulate, 47-53 x 28-35 ^u ; pedicels hyaline, 
clavate, somewhat shorter than spore. 
Common on leaves of Snccharum offiicinarum L. 

New South Wales Clarence River (Cobb 7 ). 
Queensland Brisbane and Mackay, 1898 (Tryon 5 ). 

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 p. 
Specimen not seen. 



147. Uredo scirpi-nodosi McAlp. 

II. Sori bullate, elongated, splitting lengthwise, running in lines on stem, 
1J-2 mm. long. 

Uredospores brown, subglobose, ellipsoid to oblong, variable in size, 
25-32 n diam., or 27-36 X 19-25 /j. ; epispore echinulate, often 
3 thick. 

Uredo Liliaceae. 203 

On Scirpus nodosus Rottb. 

Victoria Mordialloc, Dec., 1885 (Reader) (Winter 2). 

This specimen is given as Puccinia rimosa (Link) Wint., by Winter, but 
on carefully examining some of the original material, there are coarsely 
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 
named it as above. 

(Plate XXVIII., Fig. 241.) 



148. Uredo anguillariae Cooke. 

Cooke, Grev. XIV., p. 11. (1885). 

Cooke, Handb. Austr. Fung., p. 343 (1892). 

Sacc. Syll. VII., p. 840 (1888). 

II. Sori on both surfaces, gregarious, elliptic, bullate, long covered by the 

Uredospores globose or globosely oval, smooth, brown, epispore 
thin, 20-22 x 20 /z, 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 not seen. 


149. Uredo geitonoplesii McAlp. 

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 ^. 

On Geitonoplesium cymosum' A. Cunn. 

Victoria Orbost, July, 1901 (Pescott). 


150. Uredo schelhammerae Me Alp. 

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 ^, average 24 x 18ju. 
On Schelhammera undulata R. Br. 

New South Wales Kurrajong Heights, 1892 and 1903 (Musson). 


Uredo RJiamnaceae, Stylidiaceae, Compositae 


Spyridium, Pomaderris. 

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, 

Uredospores yellowish to orange-yellow, ellipsoid to obovate, 
distinctly echinulate, with coarsely granular contents, 22-31 x 
17-22 nj 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- 
stage has been found. 

(Plate XXVIII., Fig. 245.) 



152. Puccinia stylidii Me Alp. 

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 p, or 22-24 p diam. 

III. Teleutosori, p. 210. 

On leaves of Stylidium graminifolium Sm. 

Tasmania New Waterworks, Hobart, Nov. 1892, II. (Rod way). 
Darluca filum, Cast., literally covered some of the sori. 

(Plate XXVIII., Fig. 242.) 

153. Uredo bidentis P. Henn. 


Henniiigs, 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 ^ thick, finely echinulate or almost 
smooth, 25-35 X 22-27 p t in exceptional cases 41 p long; para- 
physes numerous, hyaline, clavate, 50-60 ^ long. 

Urcdo Compositae. 205 

On leaflets and leaf stalks of Bidens pilosa L. 
Queensland Brisbane, 1886 (Bailey 5 ). 

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 
is 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. 

Lindroth, Acta Soc. Fauna et Flora fennica, XXII., p. 1 1 

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 ^ thick, 
14-18 fji diam. 
On leaves of Crepis japonica Benth. 

Specimen not seen. 


155. Puccinia oleariae McAlp. 

Me Alpine, Agr. Gaz. KS.W. VI., p. 756 (1895). 
Sacc. Syll. XIV., p. 278 (1899). 
Uromyces asteris Me Alp. 

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 ^; epispore 
longitudinally striate, average thickness 1 J // ; 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 /uL- } pedicel hyaline, persistent, short, up to 56 ^ 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. (Rod way 1 ). 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 Puccinia. 
(Plate XXVIII., Fig. 249.) 


TJredo Proteaceae, Crasstdaceae, Leguminosae. 



156. Uredo angiosperma Thuem. 

Thuemen, Symb. Myc. Austr. IV., No. 95 ("1880). 
Cooke, Handb. Austr. Fung., p. 343 (1892). 
Sacc. Syll. VII., p. 840 (1888). 

II. Sori on both surfaces, large, commonly disposed about a circle, 

covered by the splitting, torn, and elevated epidermis, powdery, 


Uredospores oval or ellipsoid, rounded at the apex, somewhat 

narrowed at the base, 45 x 30 n ; epispore smooth, pale brownish, 

5-7 ju thick. 
On living leaves of Hakea sp. 

West Australia Oct., 1877 (F. v. Mueller) (Thumen 2 ). 
Specimen not seen. 



157. Uredo tillaeae McAlp. 

McAlpine, Agr. Gaz. N.S.W. VL, 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 ^t diam. or 20 x 16 p. 
On leaves of Tillaea sieberiana Schult. 

Victoria Studley Park, near Melbourne (Barnard). 



158. Uredo bossiaeae McAlp. 

II. Uredosori on both surfaces of leaves but mostly on under, chocolate 
brown, erumpent, surrounded by ruptured epidermis, compact, 
scattered or aggregated, sub-globose, 1-1J mm. diam. 

Uredospores elliptic, finely echinulate, golden-brown, epispore 
about 3 p thick, 24-31 x 17-21 /x. 

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. 

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 ji. 

Excluded or Doubtful Species. 207 

On leaves, twigs, and legumes of Cassia sp. 

Queensland Gladfield (Gwyther) (Bailey 13 ). 

In the original description, the spores are given as smooth, and 12-14 x 
S-10 (.(, but on examining material kindly supplied by Mr. Bailey, of 
Brisbane, tkey were found to be as above. 

(Plate XXVIIL, Fig. 247.) 



160. Uredo rhagodiae Cke and Mass. 

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 p 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 p. 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 p. 

Darluca filum Cast, is often found on the sori. 

(Plate XXX., Fig. 267.) 


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's 
Handbook of Australian Fungi, which on further investigation must be 
removed from the list of Australian species. Sometimes, although rarely, it 
happened that the host-plant of the parasite was wrongly given, and then a 
new species has been created which afterwards turned out to be a known one. 
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 Puccinia 

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 
in the National Herbarium, Melbourne, named in his own handwriting, it 


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. depauperans (Vize) Syd. The rust 011 
the native violets ( Viola hederacea and V. betonicifolia) 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 
vvarted, 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. 

8. Puccinia investita Schw. was determined by Dr. Morrison on 
species of Gnaphalium, but on investigating the original material I found it 
to be P, gnaphalii (Speg.) P. Henc. 

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. maynusiana Koern and }'. tepperi Ludw. 

11. Puccinia rimosa (Link.) Wint. This species is given by Winter 2 
as occurring on Isolepis nodosa R. ~Br.Scirpus nodosus 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 h 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 oehraceous 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 p. 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 
Harbarea, and a portion of the original specimen shows teleutosori as well as 
aecidia. I have named the rust Puccinia cruciferae. 

17. Aecidium bellidis Thuem., was determined by Cooke on Bellis 
perennis from Victoria. This is considered to represent the aecidial stage 
of Puccinia 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 Puccinia distincta. 

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 (Puccinia 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 senecionis 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 have been 
new species of Purcinia discovered. As regards the grcrundsel, 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 Jtidi'-n* piloMi 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, 





161. Uromyces polycnemi McAlp. 

I. Aecidia bright orange, generally covering lower surface of leaves, 

crowded, cylindrical, surrounded at base by ruptured epidermis, up 
to H ram. long; pseudoperidia colourless, with toothed margin; 
peridial cells oblong to somewhat lozenge-shaped, with moderately 
thick striated margins, 37-40 p, long. 

Aecidiospores orange, ellipsoid to oblong or sub-globose, very 
finely echimilate, 22-23 X 17-20 p or 20-22 p 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 2mm. in diam. 

Uredospores ellipsoid to oval, orange coloured, finely echinulate, 
with three equatorial germ-pores on one face, 28-32 X 18-22 p.. 

III. Teleuto-sori similar to uredo-sori, but generally larger and darker in 

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 surface of 
every leaf, and as the host plants grew in dense masses, in the saline marshes 
near the coast, it was difficult to find one without the rust. 

A Tuberculina was frequently parasitic on the aecidia. 

(Plate XLIIL, Fig. 319.) 



152. Puccinia stylidii McAlp. 

II. Uredosori, p. 204. 

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 p), 
rounded or attenuated at base, not infrequently three-celled, very 
variable in shape and size, 40-62 x 17-28 ju; pedicel hyaline, 
persistent, up to 45 p long. 
X. Mesospores common, similarly coloured to teleutospores, thickened 

at apex, oval to ovoid or elongated ellipsoid, 30-40 x 15-23 p. 
On leaves of Stylidium graminifolium 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. 

Glossary 211 


Abstriction. The formation of a spore by pinching off the end of a spore-forming 
hypha, without the production of a septum. 

Aecidios-pore. A spore formed in an aecidium, serially and successively abstricted. 

Aecidium. A spore-body consisting of a cup-shaped envelope or peridium with a 
hymenium at the bottom of the cup, from which aecidiospores are successively 
detached; also the generic name originally applied to the aecidiospores. 

'Amphigenous. Growing on both surfaces of a leaf. 

Am^his-pore. A modified uredospore with thickened wall, and only germinating 
after a resting period. 

A-p^pressorium. 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-petal. 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. 

Caeomos-pore. A spore formed in a caeoma. 

Ca-pitate. 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. 

Echinulate. Covered with short spines. 

Elliptical. Shaped like an ellipse or oval. 

Endochrome. The peculiar colouring matter in the cells, and usually applied to the 
coloured cell-contents of spores. 

Endos-pore. The innermost coat of a spore. 

E-pi-phyllous. Growing on the .upper surface of a leaf. 

E-pileos'pore. A name suggested for a uredospore which functions as an aecidiospore, 
and the sorus contains or is surrounded by prominent paraphyses. 

Erum-pent. 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. 

Gatnetofhyte.lhe stage in the life-cycle of the plant which bears the sexual organs. 

Germ-fore. An opening on tho surface of a spore through which the germ-tube 

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. 

Heteroecious 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. 



Hymenium. A spore-bearing layer of hyphae. 

Hy-pertro-phy. An abnormal enlargement or excessive growth of particular parts, 
which may be caused by excess of nourishment or induced by parasites. 

Hy-pha. A tubular thread-like cellular structure, collectively constituting the vege- 
tative body or mycelium of the fungus. 

Hy-po-phyllous. Growing on the under surface of a leaf. 

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. 

Mesos-pore. A single-celled spcre-form related to the teleutospore, and probably 
representing a transition from the unicellular Uromyces. 

Micron or 'Micromillimetre. Indicated by the sign p, 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 variation produced by internal influences, in which the 
equilibrium of the organism is disturbed and 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 which a parasitic mode of life is indispensable for com- 
plete development. 

Orbicular. Applied to a flat body with a circular outline. 

Papillate. Covered with protuberances or terminating in a papilla. 

Paralyses. Barren filaments associated with the spores, either slender or variously 
swollen, hyaline or coloured. 

Parasite. An organism living on r,r 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. 

Firifcrm. Pear-shaped. 

Plasmodium. A mass of naked protoplasm, containing many nuclei and exhibiting 

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 aecidium. 

Promycelial S'pore. The spore produced by a promycelium, also known as a sporid- 

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. 

Protoplasm. The living substance in plants, consisting of albuminoid matter. 

Pseudo-parenchyma. False tissue or compact mycelium, so called because it is 
formed by the union of previously separate elements or hyphae into a dense 
mass, while true tissue is derived from a single cell, which divides and 
forms a coherent, more or less firm, mass of cells. 

Pseudo--peridium. The outer envelope of the aecidium, consisting of sterile cells. 

Pulverulent. Powdery 

Pul-vinate. Cushion-shaped. 

Reticulate. Covered with lines or ridges crossing each other so as to form a net- 

Saccate. Bag-shaped. 

Sa-pro-phyte. A fungus which preys upon dead organic matter only, in contrast to 
a parasite. 

Se-ptate. Partitioned off into distinct divisions. 

Sorus. A cluster of spores constituting a spore-bed, as uredosorus, teleutosorus. 

Literature. 213 

S-permatium. A minute detachable cell produced in a spermogonium, and formerly 

considered to have a male sexual function. 

Spermogonium. The receptacle in which spermatia are produced and detached. It 
is sometimes called a pycnidium, although that term is generally reserved for 
a spore-receptacle in another division of Fungi. 
S-pore. A detachable cell, capable of reproducing the fungus. 

S-poridiolum. 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. 

S-poro-phyte. 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 


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. 
Teleutosfiore. Generally 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. 
Uredos-pore.A spore detached from the apex of a pedicel and producing a 


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. 



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 

2. - - Problems in the Study of Plant Rusts. Bull. Torr. Club, Vol. 30, p. i 

3 . 1 -'Taxonomic Importance of the Spermogonium. Ibid. Vol. 31, p. 3- 

-'The Aecidium of Maize Rust. Bot. Gaz., Vol. 38, p. 64. 1904. 

- Amphisporea of the Grass and Sedge Rusts. Bull. Terr. Club, Vol. 32, 

f, P ' 35 A N 9 ew Uredineous Fungus from Mexico Baeodromous holwayi Arth. 
Ann. Myc., Vol. 3, p. 18. 1905. 

-. Cultures of Uredineae in 1904. Journ Myc. Vol. n, p. 54- X 93- 

8 Terminology of the Spore-Structures in the Uredmales. ., V 

i. ARTHUR, J. C., and HOLWAY, E. W. D.-Descriptions of American Uredineae. 

Bull. Lab. Nat. Hist., Univ. Iowa, Vol. 5, p. 3*9- I( > 2 - 
i. ATKINSON, J.-An account of Agriculture and Grazing in New South 

London. 1826. 


Vol. i, Part I., p. 18. 1884. 
' ^opsis'of Yhi ^ue 9 ensland 4 Flora, First Supplement, pp. 84, 85. Bris- 

Supplement) p p. 125-127. Brisbane . l888 ' 

__ e ) . . 

6. - Ibid, Third Supplement, p. 123. Brisbane. iboo. 

Additional Fungus Blights observed to have injured Plant , di 
year. Ann. Rep. De* P . Agric. of Queensland, p. 45- Brisbane. 1890-9 1. 
8. - - Contributions to the Queensland Flora. Botany Bull. 2, p. 34. B 

o. l89I ' A nn. Rep. Dep. Agric. of Queensland p. $1. Brisbane. 1891-2. 

214 Literature. 

io. 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. 1892. 

u. Contributions to the Queensland Flora. Botany Bull. 5, p. 36. 1892. 

12. - Ann. R'ep. Dep. Agric. of Queensland, p. 40. Brisbane. 1892-3. 

13. Contributions to the Queensland Flora. Botany Bull. 8, pp. 110-113. 

Brisbane. 1893. 

14. Ann. Rep. Dep. Agric. of Queensland, p. 31. Brisbane. 1893-4. 

15. Contributions to the Queensland Flora. Botany Bull. 9, pp. 16, 17. 

Brisbane. 1894. 

16. Ann Rep. Dep. Agric. of Queensland, p. 35. Brisbane. 1894-5. 

17. Queensland Blight Fungi. Proc. Conf. Austr. Fruit-growers, New 

Zealand, p. 208. Wellington. 1896. 

18. Ann. Rep. Dep. Agric. of Queensland, p. 39. Brisbane. 1897-8. 

19. Botany Contributions. Queensland Agric. Journ., Vol. 2, p. 38. Brisbane. 


20. Contributions to the Flora of Queensland. Ibid. Vol. 3, p. 205. Bris- 
bane. 1898. 

21. Ann. Rep. Dep. Agric.' of Queensland, p. 33. Brisbane, 1898-9. 

22. Contributions to the Flora of Queensland. Agric. Journ., Vol. 4, p. 284. 

Brisbane. 1899. 
i. BAKER, R. T. Contributions to a Knowledge of the Flora of Australia. Proc. 

Linn. Soc., New South Wales, Vol. 24, p. 447. 1899. 
i. BANDI, W. Beitrage zur Biologic der Uredineen (Phragmidium subcorticium 

(Schrank) Winter). Hedw., Vol. 42, p. 118. 1903. 

1. BARCLAY, A. Aecidium ttrticae Schum. var. himalayense. Sci. Mem. Med. 

Off. Army of India, p. 29. Calcutta. 1887. 

2. On the Life History of a Remarkable Uredine on Jasminum grandi- 

florum, L. (Uromyces cunning namianus, n. sp.). Trans. Linn. Soc., London, 
Vol. 3, p. 141. 1891. 

3. On Two Autoecious Caeomata in Simla. Scientific Memoirs by Med. 

Officers of Army of India, Part VI. 1891. 

4. - Rust and Mildew in India. Journ. Bot, Vol. 30, p. 349. 1892. 

1. BARY, A. DE. Neue 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. 

2. Comparative Morphology and Biology of the Fungi, Mycetozoa and Bac- 
teria. English Edition, Oxford. 1887. 

i. BEAUVERIE, M. J. Essais cl'immunisation des vegetaux centre les maladies crypto- 
gamiques. Compt. rend. Vol. 133, p. 107. 1901. 

1. BERKELEY, M. J. Flora of Tasmania Fungi. Hooker's Botany of the Ant- 

arctic Voyage, Part III., Vol. 2, p. 270. 1860. 

2. Australian Fungi. Journ. Linn. Soc., London, Vol. 13, pp. 173, i74> 


i. BERKELEY, M. J., and BROOME, C. E. List of Fungi from Brisbane, Queens- 
land. Trans. Linn. Soc., London, Vol. i, p. 407. 1878. 
2. Ibid, Part II., Vol. 2, p. 67. 1882. 

3. Ibid, Part III., Vol. 2, p. 221. 1886. 

i. BLACKMAN, V. H. On the conditions of Teleutospore germination and Spori- 
dia formation in the Uredineae. New Phytologist. Vol. 2, p. io. 1903. 

2. -On the Fertilisation, Alternation of Generations, and general Cytology 

of the Uredineae. Ibid. Vol. 3, p. 23. 1904. 

3. On the Fertilisation, Alternation of Generations, and general Cytology 

of the Uredineae. Ann. Bot., Vol. 18, p. 323. 1904. 

1. BOLLEY, H. L. The Heteroecismal Puccinieae. Am. Micro. Journ., Vol. io, p. 

169. 1889. 

2. Wheat Rust. Bull. 26. Agr. Exp. Station, Indiana. 1889. 

3. Wheat Rust : Is the infection local or general in origin? Agric. Science, 

Vol. 5. 1891. 

4. Einige Bemerkungen ueber die symbiotische Mykoplasmatheorie bei dem 

Getrefderost. Centralb. f. Bakt., Vol. 4, p. 855. 1898. 

5. New work upon wheat rust. Science N.S. Vol. 22, p. 50. 1905. 

1. BUBAK, 'F. Puccinia scir-pi DC. Oesterr. Bot. Zeitschr., Vol. 48, p. 14. 1898. 

2. Uredo sym-phyti DC., und die zugehorige Tele'utosporen und Aecidienform. 

Ber. d. Deutsch Bot. Gesellsch., Vol. 21, p. 356. 1903. 

i. BUTLER, E. J. The Indian Wheat Rust Problem. Bull. No. i, Dept. Agric., 
India. Calcutta. 1903. 




i. CAMPBELL, F. M. Victorian Fungi. Viet. 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. 

i. CHODAT, R. Identite du Puccinia scir-pi DC. avec Aecidiam nym-phoidis DC. 

Verh. Schweiz. Nat. Ges., p. 43. 1888-9. 
i CHRISTMAN, A. H. Sexual Reproduction in the Rusts. Hot. Gaz., Vol. 39, p. 

267. 1905. 

1. COBB, 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. 

- 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. 1892. 

- Fungus Diseases of the Sugar-cane. Ibid. Vol. 4, p. 799. 1893. 

S Host and Habitat Index of the Australian Fungi. Dep. Agric., N.S> 

Wales. 1893. 
q. - Notes on Diseases of Plants. Agric. Gaz., N.S. Wales Vol. 5, p. 384. 


tetters on the Diseases of Plants. Ibid. Vol. 8, pp. 216, 217, 232-234. 


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. 

i. 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. 
i. CONNOLD, E. T. British Vegetable Galls. London. 1901. 

1. COOKE, M. C. Australian Fungi. Grevillea, Vol. n, pp. 97, 98. 1883. 

2. - - Ibid, Vol. 12, p. 20. 1883. 

3. - Fungi australiani (reprinted from Grevillea, Vols. 9, 10, n, 12). 1883. 

4. - Australian Fungi. Grevillea, Vol. 14, pp. n, 12. 1885. 

- Ibid. Vol. 15, p. 99. 1887. 

6. - - Ibid. Vol. 16, p. 2. 1887. 

7. - -- Ibid. Vol. 16, p. 74. 1888. 

Ibid. Vol. 17, pp. 55, 70. 1889. 

9- Ibid. Vol. 19, p. 46. 1890. 

10. - - Handbook of Australian Fungi, pp. 330-344. London. 1892. 
i. CORDA, A. Icones Fungorum. 6 Fasc. 1838-56. 


i. DANGEARD, P. A., and SAPPIN TROUFFY, P. Recherches histologiques sur les 
Uredinees. Compt. rend. Vol. 116, p 211. 1893. 

Une pseudofecondation chez les Uredinees. Ibid. Vol. 116, p. 267. 

1. DIETEL, P. Beitrage zur Morphologic und Biologic 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 

Gymno sporangium. Hedwigia, Vol. 28, p. 99. 1889. 

4- - - Ueber den Generationswechsel von Puccinia agro-pyri 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. 81, p. 

394. 1895. 

7- - - Waren die Rostpilze in friiheren Zeiten plurivor? Bot. Centralb. Vol. 
79, p. 81 and 113. 1899. 

438. II 

2 1 6 Literature. 

8. DIETEL, P. Uredinaies in ENGLER and PRANTL, Die natiirlichen Pflan 

familien. 1900. 
9. 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. 
p. 3. 1903. 
-ii. Eine neue Puccinia auf Senecio. Ibid., p. 535. 1903. 

12. Betrachlungen ueber die Vertheilung der Uredineen auf ihrer Nahr- 

pflanzen. Centralb. f. Bakt., Vol. 12, p. 218. 1904. 

13. Ueber die Arten der Gattung Phragmidium. Hedwigia, Vol. 44, p. 122 

and p. 330. 1905. 

i. DUMEE, P., and MAIRE, R. Remarques sur les Uredospores de Puccinia firuni 
Pers. Bull. Soc. Myc., France. Vol. 17, p. 308. 1901. 


i. EHRLICH, and HUEBENER. Ueber die Vererbung der Immunitat bei Tetani 
Zeitschr. f. Hygiene. Vol. 18. 1894. 

1. ERIKSSON, J. Ueber die Spezialisierung des Parasitismus bei den Getreiden 

pilzen. Ber. Deutsch. Bot. Ges. Vol. 12, p. 292. 1894. 

2. Ueber die Forderung der Pilzsporenkeimung durch Kaite. Centralb. 

Bakt. Vol. i, p. 557. 1895. 

Ueber die verschiedene Rostempfanglichkeit verschiedener Getreidesorte 

Zeitschr. f. Pflanzenk. Vol. 5, pp. 80 and 156. 1895. 

1st die verschiedene Widerstands fahigkeit der Weizensorten konst 

oder nicht? Ibid., p. 198. 1895. 

Welche Rostarten zerstoren die australischen Weizenernten ? Ibid. Vc 

6, p. 141. 1896. 

6. Welche Grasarten kb'nnen die Berberitze mit Rost anstecken? Ibid., 

193. 1896. 

7. - Neue Untersuchungen ueber die Spezialisierung, Verbreitung und H 

kunft des Schwarzrostes (Puccinia graminis Pers.). Jahrb. f. Wiss. Bot. Vc 
29, p. 499. 1896. 

8. Der heutige Stand der Getreiderostfrage. Ber. Deutsch. Bot. G< 

Vol. 15, p. 183. 1897. 

9. Vie latente et plasmatique de certaines Uredinees. Compt. rend. Vol. 

124, p. 475. 1897. 

10. Ueber die Dauer der Keimkraft in der Wintersporen gewisser Rostpilze. 

Centralb. f. Bakt. Vol. 4, p. 376. 1898. 

Sur POrigine 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 Getreideschwarzrostes in Schweden und 

in anderen Landern. Centralb. f. Bact. Vol. 9, p. 590. 1902. 

The Researches of Professor H. Marshall W T ard on the Brown Rust of 

the Bromes and the Mycoplasm Hypothesis. Arkiv for Botanik udg. of K. 
Sv. Vet. Akad. Vol. i, p. 139. 1903. 

14. Ueber das vegetative Leben der Getreiderostpilze. K. Sv. Vet. Aka 

Vol. 37, No. 6, I. 1904. 

15. - - Ibid. Vol. 38, No., 3, II. -III. 1004 

16. - - On the Vegetative Life of some Uredineae. Ann. Bot. Vol. 19, p. 55. 

1 7- Zur Frage der Entstehung und Verbreitung der Rostkrankheiten der 

Pflanzen. Arkiv. f. Botanik. K. Sv. Vet. Akad. Vol. 5, No. 3. 1905. 
J ^- - Ueber das vegetative Leben der Getreiderostpilze. K. Sv. Vet. Akad. 

Vol. 39, No. 5, IV. 1905. 

i. ERIKSSON, J., and HENNING, E. Die Hauptresultate einer neuen Untersuchung 
ueber die Getreideroste. Zeitschr. f. Pflanzenk.* Vol. 4, pp. 66 and 140. 

2 - T)ie Getreideroste, ihre Geschichte und Natur, sowie Massregeln gegen 

dieselben. Stockholm. 1896. 


i. FARLOW, W. G. Notes on some Species in the Third and Eleventh Centuries of 
Ellis' North American Fungi. Proc. Am. Acad. Arts and Sci. Vol. 
1 8. Boston. 1883. 

2. The Conception of Species as affected bv recent Investigations on Fungi. 

American Naturalist. Vol. 32, p. 675. 1898. 



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 ; 
Swedish research into the rust of cereals. Agric. Gaz., N S W'des Vol o 
p. 2 3 q. 1898. 

2. - - 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 Gymnosporangium sabinae (Dicks.) Wint. and G. confusum 

Plowright. Zeitschr. f. Pflanzenk. Vol. i, p. 260. 1891. 

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 -penicillatum. Hedw. Vol. 34, p i. 


4- - - Die biologischen Arten der parasitischen Pilze und die Entstehung neuer 
Formen im Pflanzenreich. Verh. Schweiz. Nat. Ges. Vol. 86. Locarno, p. 49, 

Die Uredineen der Schweiz. Beitr. zur Krypt. Flor der Schweiz. Vol. 2,. 

part 2, 1904. 

i. FREEMAN, E. M. Experiments on the Brown Rust of Bromes (Puccinia dis- 
$ersa}. Ann. Bot., Vol. 16, p. 487. 1902. 


i. GALLOWAY, B. T. Rust of Flax. Journ. Myc., Vol. 5, p. 215. 1889. 
i. GIBSON, C. M. Notes on Infection Experiments with various Uredineae. New 
Phytologist. Vol. 3, p. 184. 1904. 

II . 

i. HALSTED, B. D. Mycological Notes Observation in Wind infection of a Rust. 

Bull. Torr. Bot. Club. Vol. 25, p. 159. 1898. 
i. 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 beziiglich der physikalischen 

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. 

1. HITCHCOCK, 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. 

i. 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. 1899. 

2. - - Der Chrysanthemum-Rost. Ibid.. Vol. 10, p. 132. 1900. 

3- - Der Chrysanthemum-Rost. Centralbl. f. Bakt., Vol. 10, p. 369. 1903. 


i. KIENITZ-GERLOFF, F. Die Gonidieu von Gvmnos-porangium clavariae forme. Bot. 
Zeit. Vol. 46, p. 389. 1888. 

i. KLEBAHN, H. Die Wirtwechselnden Rostpilze. 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. 

i. KUEHN, J. Der Rost der Runkelrubenblatter, Uromyces betae. Bot. Zeit, p. 
540. 1869. 

H 2 




i. LAGERHEIM, G. v. Ueber Uredineen mit variablem Pleomorphismus. Troms 

Mus. Vol. 16, p. 105. 1894. 
i. LAURENT, E. De 1'action interne du sulfate de cuivre dans la resistance de h 

pomme de terre au Phyto-phthora infestans. Compt. rend., Vol. 135, p. 

1040. 1902. 

i. LINDROTH, J. I. Cecidomyia-larver, som ata rostsporer. Medd. Soc. Faun*. 

Fl. Fenn. Vol. 26, p. 25. 1900. 
i. LUDWIG, F. Ueber einen neuen Goodeniaceenrost aus Siidaustralien. Hedw. 

Vol. 28, p. 362. 1889. 

Ueber einige neue Pilze aus Australien. Bot. Centralb. Vol. 43, p. 5. 


Ueber neue australische Rostkrankheiten. Zeitschr. f. Pflanzenkrank- 
heiten, Vol. 2, p. 130. 1892. 

Ueber einige Rost-und Brandpilze Australiens. Ibid., Vol. 3, p. 137. 


Lehrbuch der niederen Kryptogamen, p. 455. Stuttgart. 1892. 


Verb. Bot. Ver. Prov. 

MAGNUS, P. Ueberwinterung der Puccinia caricis. 
Brand, Vol. 18, p. 27. 1885. 

Ueber das Auftreten der Stylosporen bei den Uredineen. Ber. dei 

Deutsch. Bot. Gesellsch., Vol. 9, p. 85. 1891. 

Zur Kentniss des Verbreitung einiger Pilze. Ber. Deutsch. Bot. Ges., 

Vol. 10, p. 195. 1892. 

Die systematische Unterscheidung nachst verwandter parasitischer Pilz< 

auf Grund ihres verschiedenen biologischen Verhaltens. Hedwigia, Vol. 
33, p. 362. !8 9 4. 

Fungi Pars. II. in J. Bornmueller, Iter Persico-turcicum. 1892-9: 

Verhandl. Zool.-bot. Gesellsch. Wien, Vol. 49, p. 95. i8q9. 

Ueber eine Function der Paraphysen von Uredolagern. Ber. Deutsct 

Bot. Ges., Vol. 20, p. 334. 1902. 

Kurze Bemerkung zur Biologic des Chrysanthemum-rostes. Centralb. f. 

Bakt. Vol. 10, p. 575. 1903. 

MAIDEN, J. H. Report on Botanic Gardens and Domains, p. 10. N.S. Wales. 

MARCHAL, E. De la specialisation du parasitisme ohez /' Erysi-phe graminis. 

Compt. Rend., Vol. 135, p. 210. 1902. 

De F immunisation de la Laitue (Lettuce) contre le Meunier. Ibid 

1067. 1902. 

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. 

Mycological Flora of the Royal Gardens, Kew. Kew Bull., p. 151. 


Chrysanthemum Rust. Gardeners' Chronicle, 8th Oct., 

- 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. 

On the Origin of Parasitism in Fungi 

Vol. 197, p. 7. 1904. 

London. 1903.. 
Phil. Trans. Royal Soc., Ser. 

MAYUS, O. Die Peridienzellen rTer Uredineen in ihrer Abhangigkeit von Stan< 

ortsverhaltnissen. Centralb. f. Bakt., Vol. 10. p. 644. 1903. 

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. 

Beet-leaf Rust or Blighted Mangel leaves (Uromyces betae}. Ibid, pj 

45, 46. 1891. 

- Report on Peach and Plum Leaf-Rust (Puccini -pnini}. Ibid, pp. 
144, pis. 3. 1891. 

- Report on Rust in Wheat as Victorian Delegate. Proc. Conf., N.I 
Wales. 1801. 

Literature. 219 

6. McALPiXE, 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. i8q.2. 

- The Undescribed Uredospores of Pnccinia burchardiac. Viet. Nat., 
Vol. 10, p. 192. 1894. 

9. - Report on Rust in Wheat Experiments at School of Horticul- 
ture, Burnley. Illustrated with maps and drawings, p. 66. Dep. Agric., 
Victoria, 1894. 

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 -pruni Pers. 

Proc. Linn. Soc., N.S. Wales. 2nd series, Vol. 10, p. 440. 1895. 

14. - - Puccinia on Groundsel with Trimorphic Teleutospores. Ibid., p. 461. 


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, 1897. 

22. - - The Fungi on the Wheat Plant in Australia/ Agric. Gaz., N.S. Wales, 

Vol. q, p. 1009. 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 

I'ndnula in Australia. Proc. Linn. Soc., N.S. Wales. Vol. 24, p. 301. 1899. 

- Two Mallee Fungi. Viet. Nat., Vol. 16, p. 141. 1899. 

- Rose Rust. Journ. Dept. Agric., Vic., p. 81. 1902. 

- Fungus Diseases of Stone-fruit Trees in Australia, and their Treatment. 
Dept. of Agric., Victoria. 1902. 

Cereal Rusts. Jour.i. 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 -pruni 

Pers. Ann. Myc., Vol. 2, p. 344. 1904. 

34. Note on the Arrangement of Teleutospores in Puccinia 'pruni Pers. 

Ibid. p. 348. 1904. 

35. - - Bobs A Rust-resisting Wheat. Journ. Dept. Agric.. Vic., p. 166. 1905. 
36. A new genus of Uredineae Uromydadium. Ann. Myc. Vol. 3, p. 303. 


i. MILESI, M., and TRAVERSO, G. B. Saggio di una monografia del genere Tri- 
phragmium. Ann. Myc., Vol. 2, p. 143. 1904. 

i. MOLINEUX, A. Some Remarks about Red Rust. Journ. Agric., S. Australia. 
Vol. 8, p. 95. 1904. 

i. MORRISON, A. New Victorian Micro-fungi. Victorian Nat., Vol. u, pp. 90, 
91. 1894. 

- Notices of Victorian Fungi. Ibid. pp. 119, 120. 1894. 

i. MUELLER, F. v. Fragmenta Phytographiae Australiae Supplement, Vol. ir, 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. 

i. MUELLER, FRITZ. Beitnige zur Kentniss der Grasroste. Hot. Centralb. Beihefte 

10, p. 181. IQOI. 
i. MUELLER, J. Die Rostpilze der Rost-und Rubusarten und die auf ihnen vor- 

kommenden Parasiten. Landw. Jahrb., Vol. 15, p. 719- 1886. 




i. NEGER, F. W. Beitrage zur Biologic cler Erysipheen. Flora 90, p. 221. 1902. 
i. NEUMANN, R. Ueber die Entwickelungsgeschichte der Aecidien und Spermo- 
gonien der Uredineen. Hedwigia, Vol. 33, p. 346. 1894. 


i. PFEFFER, W. Ueber chemotaktische Bewegungen von Bakterien, Flagellaten 
und Volvocineen. Untersuch. Bot. Inst., Tubingen. Vol. 2, p. 582. 1888. 

i. PLOWRIGHT, C. B. The connexion of Wheat Mildew (Puccinia graminis Pers.) 
with the Barberry Aecidium (Aecidium berberidis Gmel.) Card. Chron. 
Ser. 2, Vol. 18, p, 231. 1882. 

Podisoma juni-peri and Roestelia lacerata. Ibid, p. 553. 

3. Reproduction of Heteroecious Uredines. Linn. Soc., London. Botany, 

Vol. 21, p. 368. 1884. 

4. On the Life-history of certain British Heteroecismal Uredines. Quart. 

Journ. Micro. Sci., Vol. 2$, p. 156. 1885. 

5. A Monograph of the British Uredineae and Ustilagineae. London, i! 

6. - Barberry and Wheat Mildew. Gar'd. Chron., Vol. 23, p. 45. 18 

i. PRAIN, D. Rust in Wheat in the Australian Colonies. Agric. Ledger No. i( 

Calcutta. 1897. 
i. PRILLIEUX, E. Maladies des Plantes Agricoles. Pt. i, p. 242. Paris, 1895. 


i. RALPH, T. S. On the Aecidium affecting the Senedo vulgaris or Groundsel. 

Victorian Nat., Vol. 7, pp. 18, 19. 1890. 
i. RATHAY, E. Untersuchungen ueber die Spermogonien der Rostpilze. Denkschrift 

d. K. Akad. d. Wiss, Wien, Vol. 46, p. 51. 1883. 
i. RICHARDS, H. M. On some points in the development of Aecidia. Proc. 

Amer. Acad. Sci., Vol. 31, p. 255. 1895. 
i. RODWAY, L. Tasmanian Fungi. Proc. Roy. Soc., Tasmania, pp. 142, 143. 

i. ROZE, M. E. Le Puccinia chrysanthemi. Bull. Soc. Myc. France, Vol. i6 r 

p. 88. 1900. 


1. SACCARDO, P. A. Sylloge Fungorum. Vols. 7, 9, n, 14, 16, 17 1888-1905. 

2. Mycetes aliquot australiensis. Hedw., Vol. 28, p. 126. 1889. 

3. Ibid. Bull. Soc. Myc. France, Vol. 5, p. 118. 1889. 

4. Ibid. Hedw., Vol. 29, p. 155. 1890. 

5. - Ibid. Hedw., Vol 31, p. 57. 1893. 

i. SACCARDO, P. A., and BERLESE, A. N. Fungi australiensis. Rev. Myc., Vol. 
7, p. 93. 1885 

1. SALMON, E. S. Mycological Notes. Journ. Bot., Vol. 42, p. 184. 1904. 

2. Cultural Experiments with " Biologic Forms 1 ' of the Erysiphaceae. 

Phil. Trans., B., Vol. 197, p. 107. 1904. 

3. Cultural Experiments with the Barley Mildew, Erysi-phe graminis 

DC. Ann. Myc., Vol. 2, p. 88. 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. 1893. 

2. Recherches histologiques sur la famille des Ur^dinees. Le Botaniste, 

Vol. 5, pp. 59-244. 1896-97. 

i. SHUTT, F. T. The effect of Rust on the Straw and Grain of Wheat. Not 

West Farmer, Winnipeg, Manitoba. 2oth Oct., 1904. 

i. SMITH, W. G. Diseases of Field and Garden Crops. London. 1884. 
i. SMITH, R. E. The Water relation of Puccinia as-paragi. Bot. Gaz., Vol. 

p. 19. 1904. 
i. SORAUER, P. Das. Verhalten des Getreiderostes in trockenen und nassen ]'< 

ren. Zeitschr. f. Pflanzenkr., Vol. 4, p. 121. 1894. 
2. Die Predisposition der Pflanzen fur parasitare Krankheiten. Deuts 

Landwirts. Ges., Berlin. 1903. 

1. SYDOW, P. and H. Monographia Uredinearum. Vol. i. 1902-04. 

2. Neue und kritische Uredineen. Ann. Myc., Vol. i, p. 324. 1903. 

Literature. 221 


i. TASSI, F. La^Ruggine dei Crisantemi (Pitccinia chrysanthemi Roze). Bull. 

Lab. Bot., Univ. di Siena. November, 1903. 
i. TATE, R.A list of the Charas, Mosses, Liverworts, Lichens, Fungi, and Al- 

gals of Extra-tropical South Australia. Proc. Roy. Soc. S. Australia, Vol. 

4, p. 12. 1881. 

1. TEPPER, J. G. O. Red Rust : Its nature, approximate cause and probable cure. 

Proc. Roy. Soc., S. Australia, Vol. 3, p. 13. 1879. 

2. - - Notes on Australian Fungi. Ibid., Vol. 12, p. 150. 1889. 

3. - - Contributions to the Fungal Flora of Australia. Proc. Roy. Soc. S. 

Australia, Vol. 13, p. 55. 1890. 
i. TENISON-WOODS, J. E., and BAILEY, F. M. On some of the Fungi of New 

South Wales and Queensland. Proc. Linn. Soc., N.S. Wales, Vol. 5, p. 83. 

i. THOMPSON, E. H. A Hand-book to the Insect Pests of Farm and Orchard 

(including Fungi). Dep. Agric. of Tasmania. Bull, i, pp. 29-34. 1892. 
i. THUEMEN, F. v. Symbolae ad floram mycologicam Australiae. I. Grevillea, 

Vol. 4. p. 75. 1875. 
. II. 

Ibid. II. Flora No. 28. 1878 

3. - - 1st der Berberitzenrost notwendig zur Erzeugung des Grasrostes? 
Oesterr. Landw. Wochenbl. 1883. 

1. TRANZSCHEL, W. Ueber einige auf Grund von irrtumlicher Bestimmung der 

Nahrpflanzen aufgestelle Puccinia Arten. Ann. Myc., Vol. 2, p. 158. 1904. 

2. - Ueber die Moglichkeit, die Biologic wirtswechselnder Rostpilze auf Grund 

morphologischer Merkmale vorauszusehen. Arbeit, der Kais. St. Petersburg. 
Naturf. Gesell. Vol. 35, p. i. 1904. 

1. TRYON, H. Report on Insect and Fungus Pests, pp. 97, 196. 1889. 

2. - - Ann. Rep. Dep. Agric. of Queensland Vegetable .Pathology, pp. 39, 40. 


3. -- Ibid. p. 39. 1895-6. 

4 . - Ibid. p. 37. 1896-7. 

5. -- Ibid. pp. 43, 44. 1897-8. 

6. -- Ibid. p. 35. 1898-9. 

7. - Ibid. p. 30. 1899-1900. 

8. -- Ibid. p. 26. 1900-1. 

i. TUBEUF, K. F. V., and SMITH, WM. G. Diseases of Plants induced by Crypto- 
gamic Parasites. London. 1897. 


i. WARD, H. Marshall. Illustrations of the Structure and Life-history of Puc- 

cinia graminis, Ann. Bot., Vol. 2, p. 217. 1888. 
i. - - The Bromes and their Brown Rust. Brit. Assoc. Rep., Glasgow, p. 

836. 1901. 
3. - The Bromes and their Rust-fungus (Puccinia dispersa). Ann Bot., Vol. 

15, p. 560. 1901. 
4< _ _ Q n th e question of " Predisposition " and " Immunity" in Plants. 

Camb. Phil. Soc., Vol. n, p. 326. 1902. 

5. - - On pure Cultures of a Uredine, Puccinia dis-persa. Proc. Roy. Soc., 

Vol. 69, p. 451. 1902. 

6. - - On the Relations between Host and Parasite in the Bromes and their 

Brown Rust, Puccinia dispersa. Ann. Bot., Vol. 16, p. 233. 1902. 

7. - - Experiments on the effect) of mineral starvation on the Parasitism of th 

Uredine-fungus, Puccinia disfersa, on species of Bromus. Proc. Roy. bac., 
Vol. 71, p. 138. 1902. 

Further Observations on the Brown Rust of the Bromes, Puccinia dis- 

pcrsa, and its adaptive Parasitism. Ann. Myc., Vol. i, p. 132. i93- 

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. 19, p. i. 


i. WEBBER, H. J. 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. 22. 1885. 

2. Fungi australienses. Rev. Myc., Vol 8, p. 208. 1886. 

i. WCROXIX, M. Untersuchungen ueber die Entwickelung des Rostpilzes (Puc- 
cinia helianthi), welcher die Krankheit der Sonnenblume verursacht. Bot. 
Zeit., p. 677. 1872. 


Explanation of Plates. 


1. Uredosori on upper surface of leaf. 

2. Teleutosori on under surface. 


(All Figures vat. size.} 


3. Uredosori on upper and under surface, also on sheath. 

4. Teleutosori oa stem. 


$ MPALpmc. .Dvezu,. 

22 4 

Explanation of Plates. 


(All Figures nat. size.) 

5. Uredosori on under surface of leaf. 

6. Teleutosori on under surface 



7. Uredosori on under surface of leaf. 

8. Teleutosori on upper surface. 


9. Teleutosori on sheath and blade. 
10. Uredosori on under surface of leaf. 




Explanation of Plates. 


(All Figures nat. size.} 



11. Uredosori on under surface of leaf. 

12. Teleutosori on under surface. 

13. Teleutosori on stem. 


14. Uredosori on under surface of leaf. 

15. Teleutosori on young stem. 


16. Uredosori on upper surface of leaf. 

17. Uredosori on under surface. 



Explanation of Plates. 


(All Figures nat. size.) 


18. Chiefly uredosori on lower surface of chicory leaf. 

P. armeniaca). 

19. Uredo and teleutosori on lower surface of peach leaf, 
ao. Uredo and teleutosori on lower surface of apricot leaf. 


Explanation of Plates. 


(All Figures nat, size.) 

ai. Uredosori on under surface of leaf. 


22. Aecidia and teleutosori on under surface of leaf. 

23. Uredo and teleutosori on stem. 


24. Uredo and teleutosori on under surface of leaf of Viola hederacca. 

25. Aecidia on under surface of leaf of V. betonicijoiia. 


26. Aecidia and teleutosori on rpper surface of leaf, showing the dark tel( 
surrounding the other. 




Explanation of Plates. 


(All Figures nat. size.) 


27. Aecidia and teleutosori on both surfaces of leaves. 


28. Teleutosori on under surface of leaf and on leaf-stalk. 

29. Aecidia, uredo, and teleutosori on under surface of leaf, and teleutosori on stem. 


RSJireux, Gov*Prtnter. 



Explanation of Plates. 


(All Figures nat. size.) 


30. Uredosori on upper surface of leaf. 

31. Teleutosori on both surfaces. 


32. Aecidia, uredo, and teleutosori on leaf. 


33. Uredo and teleutosori on both surfaces ot leaves and on stems. 


34. Uredosori on under surface of leaf. 



Explanation of Plates. 


(Natural Size.) 


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. 




Explanation of Plates. 



(All Figures nat. size.) 

36. Uredo and teleutosori on stem, 


37. Aecidia (Caeomata) showing large swollen patches on stems and small patches on 
lower surface of leaves. 


38. Ochraceous uredosori on lower surface of leaves and flower-stalks, powdery 
teleutosori accompanying uredosori. 


1C BnttLeknJi . Del. 

r.e, 0irut 




Explanation of Plates. 




39. Leaves with aecidia in dense clusters 

40. Cluster of aecidia on leaf ... 

41. Section of aecidium showing peridial wall and spores in chains 

42. Chain of aecidiospores 

43. Single spore detached and more or less rounded 

44. Pseudoperidial cells seen from the surface ... 

nat. size 

... X 15 

X 50 

... X 1000 
... X 1000 
... X 550 





Explanation of Plates. 


(All Figures X 250 unless otherwise stated.} 


1. Portion of sheath of barley (Hordeum vulgare) showing numerous crowded black 

dots, the teleutosori of Puccinia sim-plex, the spring or orange rust of 
barley ... ... ... ... ... ... ... 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. sim-plex 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, with 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- 


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. triticina 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. 


II. Robinson, I'hot. 

to arid x 260. 




Explanation of Plates. 


(All Figures X 250.) 



if. Section of teleutosorus of Puccinia lolii avenae on oat (Avena sativa] with several 
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 $ratensis. 

14. Section of teleutosorus of P. lolii on Lolium -perenne. 

15. 16. Groups of teleutospores of P. maydis on Zea mays, including one four- 

celled spore. 

17. Teleutospores of P. flavescentis on Sti-pa flavescens. 

18. Section of teleutosorus of P. magnusiana on Phragmites communis. 

19. Five teleutospores of P. te-p-peri on Phragmites communis, showing the extremel] 

Jong pedicels. 


G. H. Robinson, Phot. 

X 250. 




Explanation of Plates. 


(All Figures X 250.) 



20. Section of teleutosorus of Puccinia anthoxanthi on Anthoxanthum odoratutn, 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. -per-plexans on Alo-pecurus geniculatus. 

24. Teleutospores of P. cynodontis on Cynodon dactylon. 

25. Section of teleutosorus of P. agro-pyri on Agro-pyron scabrnm. 

26. Group of teleutospores of P. impatientis on Elymus condensatus. 

27. Group of teleutospores of P. agrostidis on Deyeuxia forsteri. 

28. Section of teleutosorus of P. bromina on Bromus mollis. 


G. H. Robinson, Phot. 

x 250. 





Explanation of Plates. 


(All Figures X 250.} 


29. Group of teleutospores of Puccinia caricis on Carex breviculmis, Killara, Vic- 


30. Group of teleutospores of Puccinia caricis on Carex stricta, Berlin, Germany 

(Sydow, Uredineen, 460). 

31. Teleutospores of P. longis-pora on Carex caes-pitosa. 

32. Teleutospores of P. cy-peri on Cyferus rotundus. 

33. Group of teleutospores of P. tenuis-pora on Luzula campeatris, Murramurrangbong 

Ranges, Victoria. 

34. Teleutospores of P. obscura on Luzula cam-pestris, Berlin, Germany (Svdow, Ure- 

dineen, 1076). The spores of this species are differently shaped, much thicker 
in the wall and much darker than those of P. tenuis'pora. 

35. Uredospores of P. junco-phila from sorus containing both uredo and teleutosporc* 

on lunciis maritimus. 

36. Teleutospores of the s.ame. 



<;. II. Robinson, Phot. 




1 2 


Explanation of Plates. 



(All Figures X 250.) 

37. Teleutospores and mesospores of Puccinia wurmbeae on Wurmbea dioica 

38. Uredospores of P. burchardiae on Burchardia 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. hy-poxidis on Hy-poxis glabella. 

42. Teleutospores, two three-celled, of P. dichondrae on DicJiondra re-pens. 

43. Teleutospore and uredospore of P. mussoni on Ruellia australis Richmond River, 

New South Wales. 

44. Teleutospores of P. ruelliae on Ruellia sire-pens, Columbus, Ohio, U.S.A. (W. 

A. Kellerman, Ohio Fungi, 130). 


1 m#- 



G. U. Robinson, Phot. 



252 Explanation of Plates. 


(All Figures X 250.) 


45. Group of teleutospores of Puccinia carissae on Carissa ovata. 

46. Section of teleutosorus of P. alyxiac on Alyxia buxifolia, with teleulospores and 


47. Teleutospores of P. gilgiana on Leschcnaultia linarioides. 

48. 49. Teleutospores and mesospores of P. saccardoi on Goodenia geniculaia. 

50. Teleutospores and mesospores of P, brunoniae on Brunotna australis, with one 

three-celled and one four-celled teleutospore. 

51. Teleutospores of P. aucta on Lobelia ance-ps. (From a drawing by C. C. Brittle- 

' bank from the original material from Berkeley in the National Herbarium, 

1'J.ATK VI, 

G. H. Robinson, Phot 




Explanation of Plates. 


(All Figures X 250.} 


52. Teleutospores and mesospores of Puccinia tasmanica 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. lageno-phorae on Lagenofhora billardieri, show- 

ing mesospores and teleutospores. 

56. Teleutospores and mesospoies of P. helianthi on Helianthus annuus. 

57. Teleutospores of P. gnaphalii on Gna-phalium ja-ponicum. 

58. Teleutospores and mesospores of P. caloce-phali on Caloce-phalus drummondii. 

59. Teleutospores and mesospores of P. erechtitis on Erechtites qnadridentata. 


G. II. Robinson, Phot. 

x 250. 



Explanation of Plates. 


(All Figures X 250.) 


60. Section of teleutosorus of Pucdnia cinerariae on Cineraria sp. cult, with teleuto- 

spores and mesospores. 

61. Three uredospores and several teleutospores of P. cickorii on Cichorium intybus. 

62. Uredospores and teleutospores of P. hy-pochoeridis on Hy-pochoeris radicata. 

63. Teleutospores of the same. 

64. Two uredospores and numerous teleutospores of P. cyani on Centaurea cyanus. 

65. One mesospore and several teleutospores of P. calendulae on Calendula ojficinalis. 

66. Mesospores and teleutospores of P. brachycomes on Brachycome ciliaris. 

67. Teleutospores and mesospores of P. distincta on Bellis perennis. 


<;. H. Robinson, Phot. 

x 250 



Explanation of Plates. 



(All Figures X 250 unless otherwise stated.) 

Three uredospores and four teleutospores of Puccinia thuemeni on A-pium -pros- 
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 

7 1 - 


Teleutospores of P. thuemeni on A-pium graveolens, from Brighton, Victoria. 

Uredospore and teleutospore from the same as 71, showing roughened epispore 
in both forms more clearly ... ... ... ... ... x 500 

Uredospores and teleutospores of P. a-pii on A-pium graveolens^ Berlin, Ger- 
many. (Sydow, Uredineen, 558.) 

Teleutospores as in Fig. 73, showing distinctly smooth epispore 

X 500 

Teleutospores of P. bullata on Aethusa cyna-pium, with coarsely warted epispore. 
(Sydow, Uredineen, 1261.) ... ... ... ... ... X 500 

76. Teleutospores of P. xanthosiae on Xanthosia -pusilla. 


V, ~-:j 
, '^~\ 





Explanation of Plates. 


(All Figures X 250.) 



77. Teleutospore of I'uccinia oliganthae on Asperula oligantha. 

78. Teleutospores, some having germinated, of P. coprosmae, on Coprcsma hirtella. 

79. Uredospores and teleutospores of P. epilobii-tetragoni on Ep>ilobium glabellntn, 

Murramurrangbong Ranges, Victoria. 

80. Uredospores and teleutospores of P. epilobii-tetragoni on Epilobitim montanum. 

(Sydow, Uredineen, 1369.) 

Si. Teleutospores of P. epilobii DC. on Epilobium roseum, the teleutospores having 
much thinner walls, and being more variable in shape than those of P. 
e-pilobii-tetragoni and also finely verrucose. (Sydow, Uredineen, 1418.) 

82. Teleutospores of P. gel on Gettm renifoliutn. 
8$. Uredospores of P. -pruni on Prunus -persica. 

84. Paraphysis and uredospores of P. pruni on Prunus persica. 

85. Teleutospores of P. pruni on Prunus persica, rather smaller than average. 

86. Teleutospores of P. -pruni on Prunus domestica. 

87. Uredospores and teleutospores of P. zorniae on Zornia diphylla. 


G. II. Robinson, Phoc. 




Explanation of Plates. 


(All Figures X 250.} 


88, 89. Teleutospores of Puccinia tetragoniae on Tetragonia im-plexicoma, 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 Rumex brownii, one spore being 


92. Uredospore and teleutospores of P. acetosae on Rumex arifohus, all the latter 

with smooth walls. (Sydow, Uredineen, 954.) 

93. Teleutospores of P. muehlenbeckiar on Muehlenbcckia adfressa. 

94. Teleutospores and mesospore of P. diclsiana on Threlkeldia sp. 

95. Uredospores of P. kochiae on Kochia sedifolia, showing the characteristic nume- 

rous germ pores. 

96. Teleutospores of P. kochiae on Kochia sedifolia. 



G. II. Ilobiuson, Phot. 




Explanation of Plates. 


(All Figures X 250.) 


97. Teleutospores of Puccinia arenariae on Stellaria media. 

98. Teleutospores of P. -plagianthi on Plagianthus sidoides, the epispore being 

slightly channelled. 

99. Teleutospores (stained) of P. malvacearum on Malva rotundijolia. 

100. Teleutospores and mesospore of P. malvacearum on Lavatera -plcbeia. 

101. Teleutospores, mostly one-celled, of P. keteros-pora on Abutilon cris-pum. 
702. Teleutospores and mesospores of P. morrisoni on Pelargonium australc. 

103. Uredospores of P. geranii~-pilosi on Geranium -pilosum. 

104. Teleutospores of the same, the single germ pore being noticeable as a pal 

channel through the apex of the cell wall. 


x 250. 




Explanation of Plates. 


(All Figures X ?jo unless otherwise stated.) 


105. Teleutospores of Puccinia enostemonis on Eriostemon myoporoidcs. 

106. Teleutospores of P. correae on Correa lawrenciana. 

107. Teleutospores and mesospores of P. boroniae on Boronia s-pinescens. 

108. Teleutospores and mesospores of P. 'pritzeliana 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, 3113) ... ... ... ... X 500 

H2. Teleutospores of P. i-iolae on Viola arenaria, wtih smooth epispore. (Sydow, 
Uredineen, 286) ... ... ... ... ... ... X 5 


G. II. Kobinson, Phot. * 250 and 500. 



Explanation of Plates. 


(All Figures X 250.} 


113. Teleutospores and mesospores on wild oat, Avena fatua. 

114. Teleutospores on barley, Hordeum vulgare. 

1115, 116. Teleutospores, mesospores, and uredospore, one of the first three-celled, or 
native barley, Echino-pogon ovatus. 

117. Teleutospores on cocksfoot, Dactylis glomerata. 

118. Teleutospores and uredospores on silver grass, Festuca bromoides. 

IK;. Teleutospores and uredospores on an imported barley grass, Hord.enm secalinnm. 

120. Teleutospores on small canary-seed grass, Phalaris minor. 

121. Teleutospores on native wheat grass, Agro-pyron scabrum. 

122. Teleutospores on Am-phibromus neesii. 

1Y.VTK X 1 V 

G. H. Robinson, Phot. 



Explanation of Plates. 


(All Figures X 250 unless otherwise stated.} 


123. Germinating teleutospore, from each cell of which a promycelium has been 

pushed forth, cell contents collected towards apex of tube, and segmentation 

124. Germinating teleutospore at a later stage, the promycelium having become divided 

into four segments, and comparatively stout tubes emitted from each seg- 

125. Two germinating teleutospores, in the one on the left only the two median seg- 

ments of the promycelium as yet bear sporidiola. 

126. 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 which the upper two promycelial cells have produced 

sporidiola, while in the third from the apex the formation of the sporidiolum 
is just commencing. 

128. Germinating teleutospore, the promycelium bearing three sporidiola on stout 

elongated sterigmata. The sporidiolum second from the base is already 
germinating and putting forth a germ tube extending across to the promy- 
celium. The segment second from the apex has not so far produced a 
sporidiolum, though a stout elongated filament has been emitted. 

129. Promycelial spores 

130. Germinating promycelial spore 

X 500 
X 500 

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 orlycerine and water. 


G. H. Robinson, Phot. x .->.- & 500 



Explanation of Plates. 


(All Figures X 250.} 


131. Group of teleutospores of Uromyces dant/ioniae on Danthonia scmianmdaris. 

132. Teleutospores of U '. ehrhartae on Microlaena (Ehrharta] sti-poides. 

133. Teleutospores of U . tenuicutis on S-porobolus indicus. 

1 34. Two one-celled and one two-celled teleutospore of U . trlcorynes on Tricoryne 


135. Teleutospores of U '. tricorynes on Tricoryne elatior. 

136. Uredospores of V '. thelymitrae on Thelymitra antennifera. 

137. Teleutospores of the same. 

138. Teleutospores of U. orchid earum on Chilo glottis di-phylla, from original type 

material in National Herbarium. 

130. Teleutospores of U. microtidis on Microtis -porrifolia from type material from 


G. H. Robinson, Phot 




Explanation of Plates. 


(All Figures X 250.} 


140. Group of teleutospores of Uromyces -puccinioides on Selliera radicans. 

141. Teleutospores of V '. as-fcerulae on As-perula oligantha. 

142. Uredospores and teleutospores of U. trifolii on Trifolium re-pens. 

143. Uredospores of U . hardenbergiae on Hardenbergia mono-phylla. 

144. Teleutospore of the same. 

145. Two Uredospores of same at top and one teleutospore below. 

146. Section of teleutosorus of V '. atri-plicis on Atri-plex semibaccata The teleuto- 

spores are faintly striate with one prominent germ pore at apex. 

147. Two teleutospores of same at left and four uredospores at right. 

148. Uredospores of U. betae on Beta vulgaris. 

149. Teleutospores of same with prominent hyaline apiculus. 


G. II. Robinson, Phot. 



2 7 6 

Explanation of Plates. 


(All Figures X 250 unless otherwise stated.} 


150. Uredospores of Uromyces polygoni on Polygomim aviculare. 

151. Teleutospores of same. 

152. Uredospores of U. caryo-phyllinus on Dianthus caryo-phyllus, with prominent 

scattered germ pores. 

153. Teleutospores and one uredospore of same. 

154. Section through leaf of Dianthus caryo-phyllus, showing uredosorus of U '. caryo- 

phyllimts with two pycnidia of Darluca filum growing upon it, and two 
young pycnidia on opposite surface of leaf ... ... X 5 

155. Uredospores of U. -uesiculosus on Zygo-phyllnm glaucesccns. 

156. Teleutospores of same. 

157. Two-celled teleutospore of same. 


X 50 & 250. 

G. H. Robinson, Phot. 



2 7 8 

Explanation of Plates. 


(All Figures X 250 unless otherwise stated.} 


158. Teleutospores, seen from above, of Uromyces fusis-porus on Acacia nerii folia. 

159. Side view of same. 

160. Uredospores of same, with pronounced apiculus and equatorial band of germ 



161. Teleutospores of Uromydadmm simplex on Acacia -pycnantha, 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). 

163. 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. 

[64. Germinating sporidiolum of same. (Stained) 

X 5< 

165. Uredospores of same, being very similar to those of Uromyces iusis-porus, fig. 
160, though somewhat shorter and broader and' possessing more germ pores. 

'l.ATK XIX. 

G* H. Robinson, Phot. X 250 & 6UO. 





Explanation of Plates. 




(All Figures X 250.) 

Section of teleutosorus of U. 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. 

175. One-celled spores (mesospores ?), occasionally found ' intermixed with uredo- 


176. Immature two-celled colourless spore, of same character, from .uredosorus. 



li !!. Kuliinson, Phot. 

X 250. 



K 2 

Explanation of Plates. 


(All Figures X 500.} 


177. Two young teleutospore heads, arising from common stalk, the one on the 

right being younger than that on the left, and both intermediate between 
T2 and T3 in next figure. (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 ; TI, similar filament, somewhat 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 two dark 
teleutospores ; U, basidium of a detached uredospore. (Stained.) 

179. Abnormal teleutospore head of three teleutospores, a septum beneath the lower- 


r8o. Normal teleutospore head, with two teleutospores above septum and a vesicle 
below, there being no septum below the vesicle. 

181. 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 dentat( 
at apex. 

'i. ATI: XXI. 

-f ClHJ. 



28 4 

Explanation of Plates. 


(All Figures X 250 unless otherwise stated.} 


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. 


1 88. Mature teleutospores of same, with vesicles. 

189. Mature teleutospore of same germinating, the sporidiola just about to be formed. 


190. 191, 192. Successive stages in the development of a. teleutospore head of U. 

te-p-perianum 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 ... ... ... ... ... X 500 

194. Mature teleutospore head of same. (The markings are indistinct on account 

of poor condition of material) ... ... ... ... X 500 

195. Detached teleutospores of same. 


-250 i: SCO. 




Explanation of Plates. 


(All Figures X 250 unless otherwise stated.} 


196. Uredospores of U. notabile on Acacia notabilis. Original type material of 

Uredo notabilis Ludw. (Somewhat swollen by lengthened treatment with 
caustic potash.) 

197. Uredospores of the same, showing the net-like surface markings. ... x 500 

198. Compound stem, with basidia and uredospores of the same. 

199. Compound structure from uredosorus of same, being probably commencement 

of teleutospore formation ... ... ... ... ... x 500 

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 teleutospore?, separated from each other, of U. te$- 

perianum^ on Acacia melanoxylon, from Cheltenham, showing striated mark- 
ings on the surface. These are considerably smaller than the average. 


G. H Robinson, Phot. 

x 250 & 600. 




Explanation of Plates. 


(All Figures X -50 unless otherwise stated.} 



207. Teleutospores of U. bis-porum, on Acacia dealbata, one cluster of two showing 

the common staik. 

208. Teleutospores of the same. 

209. Uredospores of U. al-pinum, on Acacia deaHata. 

210. 211. Teleutospores of the same. 

212. Five uredospores of U. alpinum on Acacia dallachiana^ mixed with teleutospores 

and one mesospore (M). 

213, 214. Successive stages in the development of teleutospore clusters of the same. 


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. 


(J. H. Robinson, Phot. 

x 30 & 250. 




Explanation of Plates. 


(All Figures X 250 unless otherwise slated.} 


218. Two uredospores of U. -phyllodiorum (B. aud Br.) McAlp., on Acacia sp. from 

Queensland. (Type material of Melam-psora -phyllodiorum 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.) 

^i, 222. Uredospores of V '. phyllodiorum, on Acacia sp. from Queensland. (Type 
material of Uromyces -phyllodiae Ccoke rnd Mass., these uredospores being 
described as teleutospores. From Herbarium of F. M. Bailey.) 

223, 224. Two of the few digitate teleutospores present in the same material. 
(Material of above old and much faded.) 

2215. Uredospores of U. -phyllodiorum on Acacia dallachiana, from Bright, Victoria. 

226. Two of the same, more highly magnified, to show surface markings ... X 

227, 228. Teleutospores from the same material, some beginning to germinate. 


NOTE. The markings of the uredospores are arranged in distinct lines, and are not 
net-like as in Uromycladium notabile. 

G. H. Robinson, Phot. 

X 250 & 500. 




Explanation of Plates. 



(All Figures X 250 unless otherwise stated.} 


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 Rubus parvifolius, two 

germ pores being often seen on one face of each cell. 

236. Teleutospores, closely attached to each other, of MeJam-psora lini on Linum 


G H. Robinson. Phot. Nat. size & X 250. 



2 9 4 

Explanation of Plates. 



237. Young seedling of Platylobium formosum, with Aecidium platylobii on leaves 
and leaf stalks ... ... ... ... ... nat. size 

238. 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- 

-phylla ... ... ... X 5 

240. Section of aecidial cup of Puccinia erechtitis on Erechtites quadridentata X 50 


G. H. Robinson, Phot 

Nat. size & x 50. 




Explanation of Plates. 



(All Figures X 250). 

241. Spore of Uredo scir-pi-nodosi or Scir-pus nodosus. 

242. Uredospores of Puccinia stylidii on Stylidium gramini folium. 

243. U. bidentis on Bidens -pilosa from Queensland. 

244. U. bidentis on Bidens -pilcsa from Brazil. (Sydow, Uredineen, 1647.) 

245. V '. s-pyridii on S-pyridium par-vi folium. 

246. U. bossiaeae on Bossiaea -prostrata. 

247. U. -pallidula on Cassia sp. 

248. Uredospores of Uromyces sderanthi on Sderanthus diander. 

249. Uredospores of Puccinia oleariae on Olearia argo-phylla, epispore finely striate. 


G. H. Robinson, Phot. 

X 250 


Explanation of Plates. 


(All Figures X 250.) 


250. Uredospores and teleutospores of Puccinia menthae on Mentha pulegium. 

251. Uredospores of P. chrysanthemi on Chrysanthemum indicum^ from New South 


252. Isolated teleutospore found associated with the same. 

253. Uredospores and teleutospores of P. chrysanthemi on Chrysanthemum cult. 3 

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. 'podolefodis on Podole-pis longi- 


257. Teleutospore from same, with hyaline germ pore at each side of apex of upper 


258. Teleutospores of P. o-perculariae on O-percularia varia. 


G. H. Robinson, Phot. 




Explanation of Plates. 


(All Figures X 250.} 

259. Uredospores of Puccinia cacao on Rottboellia compressa. 

260. Uredospores and one teleutospore of same. 


261. Uredospores of P. lolii avenae on Avena sativa, germinating in water after 

24 hours. (Stained.) 

262. Uredospores of P. subnitens on Distichlis maritima. 

263. 264. Teleutospores from same. 

265. Teleutospores of P. liibbertiae on Hibbertia sericea. 

266. Teleutospores of P. crucifcrae on an undetermined Crucifer. From type 

material referred to Aecidium barbareae DC., at Kew. 

267. Uredo rJiagodiae on Rhagodia billardieri. 


G. H. Robinson, Phot. 


3 02 

Explanation of Pla'es. 


(All Figures X 250 unless otherwise stated.} 


268. Leaf of Loranthus celastroides, showing uredosori and teleutosori of Puccinia 

loranthicola ... ... ... ... ... ... ... nat. size 

269. Teleutospores from same. 

270. Uredospores from s^me. 

271. Leaf of Loranthus celastroides, with aecidia of same ... ... nat. size 

272. Teleutospores of Phragmidium longissimitm. 

a. Four and five celled spores, before germination. 

b. Spores commencing to germinate, with one germ tube divided by a 


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. 


G. H. Robinson, Phot. x 250 & 500. 



Explanation of Plates. 



273. Leaves of Acacia longifoh'a, 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 *pvcnantha, with the minute powdery sori of U . simplex, nat. size 


Nat. size. 

U. H. .Robinson, Phot. 




Explanation of Plates. 



276. Sori of U. bis-porum on stems and leaf stalks of Acacia dealbata 

277. Sori on fruits 

. nat. size, 
nat. size 

G. H. Robinson, Phot. 



3 o8 

Explanation of Plates. 



278. Branch of Acacia armata showing the powdery galls caused by the growth of 
U. tefferianum nat . s i ze 


G. H. Robinson, Phot. 

Nat. sizi 



3 io 

Explanation of Plaits. 



279. Galls attached to branches of Acacia fycnantha, caused by U. let- 

nat. size 

280.. Galls detached 

G. H. Rot inson, Phot, 



Nat. size. 



Explanation of Plates. 



281. Numerous young galls on Acacia dealbata, bearing the uredospores of U. 
notabile ... ... ... ... ... ... ... nat. size 

(Being on the young growth of the current season, the galls are necessarily 
comparatively small, though many of several years' growth bearing teleuto- 
spores have been found as large as potatoes, and weighing nearly i Ib. 

PLATE \\\V[ 

G. H. Robinson, Phot. 

Nat. size. 



L 2 


Explanation of Plates. 



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 


G. H. Robinson, Phot. Nat. size. 


Explanation of Plates. 


285. Witches' broom, due to C. jacksoniae on Gom'pholobium latifolium ... 

286. Normal healthy shoot of Gom-pholobium latifolium 


nat. size 
nat. size 

PLATE .\.\\V] 1 1. 

G. H. Robinson, Phot. Nat. size. 


Explanation of Plates. 


(All aecidia X 50, and aecidio scores X 300.) 


287. A. veronicae on Veronica sp. 

288. A. $lantaginis-variae on Plan fa go varia. 

289. A. lobeliae Thuem., on Lobelia pratioides, described in connexion with Pttccinia 

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. calttiae on Caltha introloba. 

296. A. ranunculacearum on Ranunculus rivularis. 

297. Aecidium of Uromyces *politus on leaves of Muehlenbeckia cunningnami a. 

nat. size, b. aecidial tubes X 5. 

298. Aecidiospores of same(X 300) and pseudoperidial cells (X 150). 

PLATK \\.\l.\ 

\ I 



C. C. Brittlebank, Del. 


320 Explanation of Plates. 


(All Figures X 500.) 



299. 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 S^haero-phragmium. The variation even in the same sorus is sometimes 
greater than it is between some different species. 


300. Teleutospores may be divided generally as in the above, although the four- 

celled forms are not quite so frequent. 


301. Teleutospores are also one to four celled, but the septa are generally transverse, 

while the upper cell may occasionally be longitudinally divided. 


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 


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. 

C. C. Brittlebank. Del. 



X 5UU 

3 22 

Explanation of Plates. 


304. Large gall on Acacia im$lexa, weight 3 Ibs. 



G H. Robinson, Phot. 




3 2 4 

Explanation of Plates. 



305. Witches' broom on Acacia im-plexa, due to Uromycladium te-pperianum ... x i 


306. Teleutospores of Uromyces a'p-pendiculatus on Vigna catjang or Cow-pea, from 

Richmond, New South Wales ... .. ... ... ... X 250 

307. Teleutospores of Uromyces fabae on Lathyrus venosus. (Sydow, Uredineen, 

1353) X 2 5 


G. H. Robinson, i'hot. 



326 Explanation of Plates. 


(All Figures X 250.} 



308. Teleutospores of P. vittadiniae, mostly of a short broad type, on Vittadinia 


309. Teleutospores and mesospores of the same. 

310. Teleutospores and mesospores of P. calotidis on Calotis sp. 

311. Uredospore of P. graminis on Avena Jatua, germinating in water, showing 

branching growth at the end of twenCy-four hours. jStained with Congo 

312. Mesospore of P. calendulas on Calendula officinalis, germinating in water after 

three days, two sporidiola produced, both of which have already germinated. 
(Stained with Bismarck brown.) 


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 shown 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.) 


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. -politus, one being two-celled, on Muehlenbeckia cun- 


318. Uredospores with two bands of germ pores, and teleutospores of U. bicinctus 

on Acacia -fasciculi-fera. 

319. Uredospores and teleutospores of U, polycnemi on Polycnemum -pentandrum. 


G. H. Robinson, Phot. ,-.- 


Explanation of Plates. 




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. 

PLATE \\.\\. 

S-colour process. 

Fig 320. 


Straw perfectly clean. 



Sfraw b.idlv rustr<l. 

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.) 

Acacia armata R.Br. 

[Jromycladium tepperianum (Sacc.) 

Acacia buxifolia A. Cunn. 

Uromycladium alpinum, McAlp. 

Acacia binervata DC. 

Uromycladium notabile (Ludw. ) 

Acacia dallachiana F.v.M. 

Uromyces phyllodiorum (Berk, and 

Br.) McAlp. 
Uromycladium alpinum, McAlp. 

Acacia dealbata Link. 

Uromyces phyllodiorum (B. and Br.) 


Uromycladium alpinum, McAlp. 
U. bisporum, McAlp. 
U. notabile (Ludw.) McAlp. 

Acacia decurrens Willd. 

Uromycladium notabile ( Ludw.) 

Acacia diffusa Lindl. 

Uromycladium tepperianum (Sacc.) 

Acacia elata A. Cunn. 

Uromycladium notabile ( Ludw.) 

Acacia erioclada Benth. 

Uromycladium tepperianum (Sacc. ) 

Acacia fasciculifera F. v. M. 

Uromyces bicinctus, McAlp. 

Acacia glaucoptera Benth. 

Uromycladium tepperianum (Sacc.) 

Acacia hakeoides A. Cunn. 
Uromycladium tepperianum (Sacc.) 

Acacia implexa Benth. 

Uromycladium tepperianum (Sacc.) 

U. alpinum, McAlp. 

Acacia juniperina Willd. 

Uromycladium tepperianum (S*cc.) 

Acacia linifolia Willd. 

Uromycladium alpinum, McAlp. 
Acacia longifolia Willd. 

Uromycladium maritimum, McAlp. 

U. tepperianum (Sacc. ) McAlp. 

Acacia melanoxylon R.Br. 

Uromycladium robinsoni, McAlp, 
Uromycladium tepperianum (Sacc. 

Acacia microbotrya Benth. 

Uromyces phyllodiorum (B. and Br.) 

Acacia myrtifolia Willd. 

Uromycladium tepperianum (Sacc. ) 

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.) 

Uromycladium notabile (Ludw.) 


Acacia penninervis Sieber. 
Uromyces phyllodiorum (B. and Br. ) 

Acacia pruinosa A. Cunn. 

Uromyces phyllodiorum (B. and Br.) 



notabile (Ludw.) 


Acacia pycnantha Benth. 

Uromycladium simplex, McAlp. 
U. tepperianum (Sacc.) McAlp. 

Acacia retinodes Schlect.= Acacia 
neriifolia, A. Cunn. 

Acacia rigens A. Cunn. 

Uromycladium tepperianum (Sacc.) 

Acacia salicina Li mil. 

Uromyces fusisporus, Cke. ami 
Uromycladium tepperianum (Sacc.) 


Host Index. 

Acacia siculiformis A. Cunn. 

Uromycladium tepperianum (Sacc.) 

Acacia spinescens Benth. 

Uromycladium tepperianum (Sacc.) 

Acacia stricta Willd. 

Uromycladium tepperianum (Sacc. ) 

Acacia verniciflua A. Cunn. 

Uromycladium tepperianum (Sacc.) 

Acacia verticillata Willd. 

Uromycladium tepperianum (Sacc.) 

Acacia vomeriformis A. Cunn. 

Uromycladium tepperianum ( Sacc. ) 

Acaena ovina A. Cunn. 

Phragmidium potentillae (Pers.) 

Acaena sanguisorbae Vahl. 

Phragmidium potentillae (Pers. ) 

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 thuemeni, McAlp. 

Apium prostratum Labill. 
Puccinia thuemeni, McAlp. 

Asperula oligantha F.v.M. (A. 

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. 

Bidens pilosa L. 

Uredo bidentis, P. 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 R. Br. 

Aecidiiim 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 

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. 

Uromj/ces bulbinis, Thuem. 

Burchardia umbellata R. Br. 

Puccinia burchardiae, Sacc. 

Calendula omcinalis L. 

Puccinia calendulae, McAlp. 

Calocephalus drummondii Benth. 

Puccinia calocephali, McAlp. 

Calocephalus lacteus Less. 

Puccinia calocephali, McAlp. 


Puccinia calotidis, McAlp 

Calotis cuneifolia R. Br. 

Puccinia calotidis, McAlp. 

Caltha introloba F.v.M. 
Aecidiurn calthae, Grev. 

Candollea serrulata Labill. = Sty- 

lidium graminifolium, Sw. 

Canthium (Plectronia) coprosmoides 

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. 
Puccinia carissae, Cke. and Mass. 

Cassia Tourn. 
Uredo pallidula, Cke. and Mass. 

Centaurea cyanus L. 

Puccinia cyani (Schleich.) Pass. 

Chiloglottis diphylla R.Br 

Uromyces orchidearum, Cke. and 

Chiloglottis gunnii Lindl. 

Uromyces orchidearum, Cke. and 

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 chmatidis 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, Liiidr. 

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. 


Host Index. 

Dampiera stricta R. Br. 

Puccinia dampierae, Syd. 

Danthonia DC. 

Uromyces danthoniae, McAlp. I. 

Danthonia semiannularis R.Br. 
Urorayces danthoniae, Me Alp. II., III. 

Deeringia celosioides R.Br. 

Aecidium deeringiae, Cke. and Mass. 

Deyeuxia fqrsteri Kunth. (Agrostis 

solandri F. v. M.) 
Puccina agrostidis, Plow. 

Deyeuxia quadriseta Benth. 

Puccinia graminis, Pers. 

Dianthus caryophyllus L. 

Uromyces caryophyllinus (Schrank) 

Dianthus chinensis L. 

Uromyces caryophyllinus (Schrank) 

Dichondra repens Forst. 

Puccinia dichondrae, Mont. 

Diploglottis cunninghamii Hook. f. 
Uromyces diploglottidis, Cke. and 

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.) 

Epilobium billardierianum Ser. 

Puccinia epilobii-tetragoni (DC.) 

Epilobium glabellum Forst. 

Puccinia epilobii-tetragoni (DC.) 

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 renifolium F. 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. 


Hakea Schm.l. 

Uredo angiosperrna, Thuem. 

Hardenbergia monophylla Benth. 

(Kennedya monophylla Vent.) 
Uromyces hardenbergiae, McAlp. 
Helianthus annuus L. 

Puccina helianthi, Schw. 

Helianthus tuberosus L. 

Puccinia helianthi, Schw. 
Helichrysum Vaill. 

Pucciuia kalchbrejmeri, 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 (t)C.) 

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. Hena. 

Juncus effusus L. 

Puccinia juncophila, Cke and Mass. 

Juncus maritimus Lam. 

Puccinia juncophiia, 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, Ck\ 

Lagenophora huegelii Benth. 

Puccinia lagenophorae, Cke. 

Lavatera plebeia sim>. 

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 liui (Pers. ) Tul. 

Lobelia anceps L. 
Puccinia aucta, Berk, and F. v. M. 

Lobelia pratioides Benth. 

Puccinia aucta, Berk, and F. v. M. 

Lobelia purpurascens R. Br. 

Puccinia aucta, Berk, and F. v. M. 

Lolium perenne L. 

Puccinia lolii, Niels. 

Loranthus celastroides Sieber. 
Puccinia loranthicola, McAJp. 

Luzula campestris DC. 
Puccinia tenuispora, McAlp. 

Luzula oldfieldii Hook. f. 
Puccinia tenuispora, McAlp. 

Malva rotundifolia L. 

Puccinia malvacearum, Mont. 

Malva sylvestris L. 

Puccinia ^malvacearum, Mont. 

Mentha laxiflora Benth. 
Puccinia menthae, Pers. 

Mentha pulegium L. 
Puccinia menthae, Pers. 

Microtis porrifolia R.Br. 
Uromyces microtidis, Cke. 

Microlaena stipoides R. Br. (Ehrharta 

*ti]ioi<l>* Labill.) 
Uromyces ehrhartae, McAlp. 

Muehlenbeckia adpressa Meissn. 
Puccinia muehlenbeckiae (Cke.) Syd. 

Muehlenbeckia cunninghami 1 

Uromyces politus (B. and Br.) McAlp. 


Host Index. 

Muehlenbeckia gracillima Meissn. 
Puccinia muehlenbeckiae (Cke.) Syd. 

Olearia argophylla F. v. M. (Aster). 

Puccinia oleariae, McAlp. 

Olearia axillaris F.v.M. 

Aecidium oleariae, Me Alp. 

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 plagia'nthi, 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) 

Rosa laxa Retz. 

Phragmidium subcorticium (Schrank) 

Rosa rubiginosa L. 

Phragmidium subcorticium (Schrank ) 

Rottboellia compressa L. 
Puccinia cacao, McAlp. 

Rubus moluccanus L. 

Phragmidium longissimum, Thuem . 

Rubus parvifolius L. 

Phragmidium barnardi, Plow, and 

Ruellia australis Cav. 

Puccinia mussoni, McAlp. 

Rumex brownii Campd. 

Puccinia ludwigii, Tepp. 

Host Index. 


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 tasiiianica, 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. M. 

(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. 

Piaccinia 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 Emll. 
Uromyces thelymitrae, McAlp. 

Threlkeldia drupata Diels. 
Puccinia dielsiana, P. Henn. 

Tillaea sieberiana Schult. 
Uredo tillaeae, McAlp. 

Tremandra stelligera R. Br. 

Puccinia pritzeliana, P. Heiin. 

Tricoryne elatior R. Br. 
Uromyces tricorynes, McAlp. 

Trifolium repens L. 
Uromyces trifolii (Alb. and Schw.) 

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 discJorme, McAlp. 

Veronica gracilis R. Br. 
Aecidium disciforme, McAlp. 

Vicia fabaL. 
Uromyces fabae (Pers.) De Bary. 

Vigna catjang Walp. 

Uromyces appendiculatus (Pers.) 

Viola betonicifolia Sm. 
Puccinia hederaceae, McAlp. 


Host Index. 

Viola hederacea LabilL 
Puccinia hederaceae, McAlp. 

Vittadinia australis Rich. 
Puccinia vittadiniae, Me Alp. 

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. 

Pnccinia zorniae (Diet.) McAlp. 

Zygophyllum billardieri DC. 

Uromyces vesiculosus, Wint. 

Zygophyllum glaucescens F.v.M. 

Uromyces vesiculosus, Wint. 

Fungus Index. 337 


Synonyms in italics ; * Species excluded from Australia ; t Doubtful species. 


Aecidium Pers. ... ... ... ... _ 194 

*apoeyni, Schwein. (See Caeoma apocyni, Me Alp., p. 193) 209 

*barbareae, DC. (SeePuccinia cruciferae, McAlp., p. 184) ... 209 

*bellidis, Thuem. (See Puccinia distiucta, McAlp., p. 156) "' 209 

bossiaeae, P. Henn. = Aecidium eburneum, McAlp. ... ... jgg 

calendulae, Me Alp. = Puccinia calendulae, McAlp. ... ... "". 151 

calthae, Grev. Caltha introloba .. ... ... ... .201 

clematidis, DC. = Puccinia agropyri, Ell. & Ev. I. ... ... "' 113 

*ccmpositarum, Mart. (See Puccinia tasmaiiica, Diet., p. 163) ... ... 209 

cymbonoti, Thuem. Cymbonotus lawsonianus ... ... ... ... 196 

cystoseiroides, Berk. = Puccinia operculariae (Morr.) Syd. I. ... ... 166 

deeringiae, Cke. and Mass. Deeringia celosioides ... ... '200 

disciforme, McAlp.- Veronica calycina, V. gracilis ... ... 194 

disseminatum, Berk. Hypericum japonicum ... ... ... ... 200 

eburneum, McAlp. Bossiaea cinerea, B. heterophylla, B. linophylla, B. 

microphylla, B. rhombifolia ... ... ... ... ... 198 

goodejiiacearum, Berk. = Uroniyces puccinioides, Berk. & F.v.M. L, and 

Puccinia saccardoi, Ludw. I. ... ... ... ... 91, 147 

impatientis, Schw. = Puccinia impatientis (Schw. ) Arth. I. ... ... 123 

lobeliae, Thuem. = Puccinia aucta, Berk. & F.v.M. I. ... ... ... 148 

microstomum. Berk. Puccinia aucta, F.v.M. I. ... ... ... 148 

monocy stis, Berk. Abrotanella f orsterioides ... ... ... ... 1 97 

nymphoidis, DC. Limnanthemum indicum ... ... ... ... 196 

oleariae, McAlp. Olearia axillaris 

perkinsiae, P. Henn. = P. gilgiana, P. Henn. I. ... ... 146 

*plantaginis, Ces. (See A. plantaginis-variae, McAlp., p. 195) ... 
plantaginis-variae, McAlp. Plan tago varia 
platylobii, McAlp. Platylobium formosum ... ... ... 199 

plectroniae, Cke. Canthium (PUctronici) coprosmoides ... 198 

ranunculacearum, DC. Ranunculus sp., R. gunnianus, 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) 
mttadiniae, McAlp. = Puccinia vittadiniae, McAlp. 

Caeoma Link 

apocyni, McAlp. Tabernaemon tana orientalis 
clematidis, Thuem. - Clematis aristata, C. microphylla ... 

Cronartium Fr. ... 

*asclepiadeum (Willd.) Fries (See C. jacksoniae, P. Henn., p. 190) 
jacksoniae, P. Henn. Jacksonia scoparia, Aotus villosa, Bossiaea cmerea, 
Gompholobium latifolium, Platylobium formosum . 

Hamaspora Koem. 

longissima, Koern.=Phragmidium lougissimum, Thuem. 

Melampsora Cast 

hypericorum (DC.) Schroet. Hypericum japonicum ... Jjj 

lini(Pers.)Tul.-Linummargmale, L. usitatissimum. (Introduced.) ... 192 
tnesodaphnes, Berk, and Br. Probably a Hyphomyccte ., r>r\ 

pkyllodiorum, Berk, and Br.-Uromyces phyllodiorum (B. and ^ 



Fungus Index. 

Phragmidium Link 

barnardi, Plow, and Wint. Rubus parvifolius 
longissimum, Thuem. Rubus moluccanus 

potentillae (Pers.) Karst. Acaena ovina, A. sanguisorba ... 

subcorticium (Schrank) Wint. Rosa canina, R laxa, R. rubiginosa. 




Puccinia Pers. ... ... ... 112 

*acetosae (Schum.) Koern. (See P. ludwigii, Tepp. p. 174) ... ... 207 

*aegra, Grove. (See P. hederaceae, McAlp., p. 183) ... ... ... 208 

agropyri, Ell and Ev. Ayropyron scabrum II., III. ; Clematis aristata I. 113 

agrostidis, Plow. Deyeuxia forsteri ... ... ... ... ... 114 

altera, McAlp. = P. cynodontis, Desm. ... ... ... ... ... 118 

alyxiae, Cke. and Mass. Alyxia buxifolia ... ... ... ... 143 

angustifoliae, McAlp. Scorzonera augustifolia ... ... ... 150 

anthoxanthi, Fckl. Anthoxanthum odoratum. (Introduced.) ... ... 115 

*apii, Desm. (See Puccinia thuemeni, McAlp., p. 168) ... ... ... 208 

arenariae (Schum.) Schroet. Stellaria media. (Introduced.) ... ... 177 

aucta, Berk, and F.v.M. Lobelia anceps, L. pratioides, L. purpurascens, 

Pratia erecta, P. pedunculata, P. platycalyx ... ... ... 148 

beckmanniae, McAlp. Beckmannia erucaeformis. (Introduce 1.) ... 116 

berkeleyana, De Toni = P. dichondrae, Mont. ... ... ... ... 142 

boroniae, P. Henn. Boronia spines cens ... ... ... ... 181 

brachycomes, McAlp. Brachycome ciliaris, B. diversifolia, B. pachyptera, 

B. scapiformis ... ... ... ... ... ... ... 150 

bromina, Eriks. Bromus arenarius, B. mollis ... ... ... ... 116 

brunoniae, McAlp. Brunonia australis ... ... ... ... 144 

burchardiae, Sacc. Burchardia umbellata ... ... ... ... 138 

cacao, McAlp. Rottboellia compressa ... ... ... ... 117 

calendulae, McAlp. Calendula officinalis ... ... ... ... 151 

calocephali, McAlp. Calocephalus drummondii, C. lacteus ... ... 151 

colotidis, McAlp. Calotis sp., C cuneifolia ... ... ... ... 152 

caricis (Schum.) Reb. Carex alsophila, C. breviculmis, C. gunniana, C. 

inversa, C. paniculata, C. pedunculata, II., III. ; Urtica dioica I. ... 133 

carissae, Cke. and Mass. Carissa ovata ... .. ... ... 144 

castagnei, Thuem. = P. thuemeni (Thuem.) McAlp. ... ... ... 168 

*caulincola, Cda. (See P. hypochoeridis, Oud., p. 159.)... ... ... 208 

chrysanthemi, Roze Chrysanthemum indicum. (Introduced.) ... ... 153 

cichorii (DC.) Bell. Cichorium intybus. (Introduced.)... ... ... 154 

cinerariae, McAlp. Cineraria sp. cult. ... ... ... ... 155 

coprosmae, Cke. Coprosma billardieri, C. hirtella ... ... ... 165 

coprosmatis, Morr. = P. coprosmae, Cke. ... ... ... -.. 165 

coronifera, Kleb. =P. lolii, Niels ... ... ... 123 

correae, McAlp. Correa lawrenciana , ... ... ... ... 181 

cruciferae, McdJp. Crucifer unknown ... ... ... ... 184 

cyani (Schleich.) Pass. Centaurea cyanus. (Introduced.) ... ... 156 

cynodontis, Desm. Cynodon dactylon 

cyperi, Arth. Cyperus rotundus ... ... ... ... ... 134 

dampierae, Syd. Dampiera alata, D. stricta ... ... ... ... 146 

dichondrae, Mont. Dichondra repens. ... ... ... ... 142 

dielsiana. P. Henn. Threlkeldia drupata, Diels. 

*dispersa Eriks. (See P. bromina, Kriks. and P. triticina, Eriks.)... 116, 132 

distincta, McAlp. Bellis perennis ... ... ... ... ... 156 

epilobii-tetragoni (DC.) Wint Epilobiurn sp., E. billardieranum, E. 

glabellum ... ... ... 170 

erechtitis, McAlp. Erechtites sp., E. arguta, E. prenanthoides, E. quad- 

ridentata ... ... ... ... ... ... ... 157 

eriostemonis, McAlp. Eriostemon myoporoides 

festucae, Plow. Festuca ovina, F. rigida. (Introduced.) ... ... 119 

flavesceritis, McAlp. Stipa navescens, S. semibarbata 

gei, McAlp. Geum renifolium ... ... ... ... 170 

*geranii, Corda (See P. morrisoni, McAlp., p. 180 )... ... ... 208 

geranii-pilosi, McAlp. Geranium pilosum ... ... ... ... 179 

gilgiana, P. Henn. Leschenaultia linarioides ... ... 146 

gnaphalii, (Speg.) P. Henn. Gnaphalium japonicum, G. purpureum ... 158 

gnaphaliicola, P. Henn. P. gnaphalii (Speg.) P. Henn. ... ... 158 

Fungus Index. 339 


Puccinia Pers. continued. 

graminis, Pers. Agropyron divergens, A. scabrum, Alopecurus geniculatus, 
Amphibromus neesii, Avena fatua, A. sativa, Beokmannia erucae- 
formis, Briza minor, Bromus racemosus, B. secalinus, B. sterilis, Dactylis 
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 
haemotlori, P. Henn. Haemodorum sp. ... ... ... ... 139 

hederaceae, McAlp. Viola betonicifolia, V. he deracea ... ... ... 183 

helianthi, Schwein. Helianthus annuus, H. tuberosus. (Introduced.) ... 158 
heterospora, Berk, and Curt. Abutilon avicennae, A. crispum, Hibis- 
cus sp. ... ... ... ... ... ... ... ... 177 

hibbertiae, McAlp. Hibbertia sericea ... ... ... ... 185 

hypochoeridis, Oud. Hypochoeris glabra, H. radicata. (Introduced.) ... 159 
hypochoeridis, McAlp. (P.macalpini, Syd. ) = P. lagenophorae, Cke. ... 161 

hypoxidis, McAlp. Hypoxis glabella ... ... ... ... ... 139 

impatientis, (Schw.) Arth. Elymus condensatus. (Introduced.) ... 123 

*investita, Schw. (See P.gnaphalii (Speg.) P. Henn. 

juncophila, Cke. and Mass. Juncus effusus, J. maritimus, J. pallidus ... 136 
kalchbrenneri, De Toni. Helichrysum sp. ... ... ... ... 160 

kochiae, Mass. Enchylaena tomentosa, Kochia sedifolia, K. villosa ... 176 

lagenophorae, Cke. Lagenophora billardieri, L. huegelii 

lolii, Niels. Avena fatua, A. sativa, Lolium perenne. (Introduced.) 

longispora, McAlp. Carex sp., C. caespitosa, C. vulgaris 

loranthicola, McAlp. Loran thus eel astroides 

ludwigii, Tepp. Rumex brownii, R. flexuosus... 

macalpini,Syd. P. lagenophorae, Cke. 

magnusiana, Koern. Phragmites communis ... ... ; 125 

malvacearum, Mont. Althaea rosea, Lavatera plebeja, Malva rotundifolia, 

Plaghnthusspicatus. (Introduced.) 
maydis, Bereng. Zea mays. (Introduced.) ... ... ... j 2 ? 

menthae, Pers. Mentha laxiflora, M. pulegium. (Introduced.)... ... HO 

microseris, Me Alp. P. hypochoeridis, Oud. 

morrisoni, McAlp. Pelargonium australe 

muehlenbeckiae (Cke.) Syd. Muehlenbeckia adpressa, M. gracillima ... 1/5 

munita, Ludw. = P. dichondrae, Mont. ... "** i2 

mussoni, McAlp. RuelJia australis ... - |*f 

nigricaulis. McAlp. = P. saccardoi, Ludw. **? 

oleariac, McAlp. Olearia argophylla ... 

oliganthae, McAlp. Asperula oligantha 

operculariae (Morr.) Syd. Opercularia aspera, 0. varia . ... 

perplexans, Plow. Alopecurus geniculatus ... '" ona 

*phragmitis (Schum.) Koern. (See P. magnusiana, Koern, p. 125) 
plagianthi, McAlp. Plagian thus sidoides ... *** 

poarum, Niels. Poa annua, P. caespitosa, P. pratensis. (Introduc 

podolepidis, McAlp. Podolepis longipedata 

prenanthis (Pers.) Lindr. Lactuca sp. (Introduced.) ... ... J?J 

pritzeliana, P. Henn. Tremandra stelligera ... ... 

pruni, Pers. Prunus amygdalus, P. armeniaca, P. domestica, f. p ^ 

(Introduced.) ... ... i<xi 

purpurea, Cke.-Sorghum halepense, S. vulgare. (Introduced.) ... l- 

nmosa, Link. (See Uredo scirpi-nodosi, McAlp., p. 203) 
*rumicis, Lasch. (See P. ludwigii, Tepp., p. 174) ... 

runncis-scutati var. muehlenbecktae, Cke. =P. muehlenbeckiae (Cke ) Syd. ... 

saccardoi, Ludw. - Goodenia albiflora, G. gemculaU G. glauca G. 

sorghi _. .. . , 

stylidii, McAlp. Stylidium graminifohum 

subniteris, Diet. Distich) is maritima ... 

tasmanica, Diet. Senecio brachyglossus, S. pectmatus, & 

teimisporaf McAlp. Luzula campestris, L. oldheldii ... 

tepperi, Ludw. Phragmites communis... ^3 

tetragoniae, McAlp. Tetragonia implexicoma Wntmdu'ced ) 168 

thuemeni, McAlp.-Apium graveolens A. prostratum. (Intrc 

triticina, Eriks. Triticum vulgare. (Introducea.) 


Fungus Index. 

Puccinia Pers. continued. 

*violae, (Schum.) DC. (See P. hederaceae, McAlp., p. 183) 
vittadiniae, McAlp. Vittadinia australis 
wurmbeae, Cke. and Mass. Wurmbea dioica ... 
xanthosiae, McAlp. Xanthosia pusilla 
zorniae (Diet.) McAlp. Zornia diphyl la 

Uredo Pers. 

angiosperma, Thuem. Hakea sp. 

anguillariae, Cke. Wurmbea dioica 

armillata, Ludw. = Puccinia juncophila, Cke. and Mass. 

bidentis, P. Henn. Bidens pilosa 

bossiaeae, McAlp. Bossiaea prostrata 
*cichoracearum, DC. (See Uredo bidentis, P. Henn., p. 204) 

dematidis, Berk.=Caeoma clematidis, Thuem. 

crepidis-japonicae, Lindr. Crepis japonica 

ehrhartae, McAlp. = Uromyces ehrhartae, Me Alp. 

geitonoplesii, McAlp Geitonoplesium cymosum 

gnaphalii Speg. = Puccinia gnaphalii (Speg. ) P. Henu. 

kuehnii, Krueg. Saccharum officinarum. (Introduced.) 
*leguminum, Desm. Acacia sp. 

notabilis, Ludw. = Uromycladium notabile (Ludw.) McAlp. 

pallidula, Cke. and Mass. Cassia sp. ... ... ... .., 

rhagodiae, Cke. and Mass. Rhagodia billardieri 

rottboelliae, Diet. = Puccinia cacao, McAlp. 

schelhammerae, McAlp. Schelhammera undulata 

scirpi-nodosi, McAlp. Scirpus nodosus 

sorghi, Fckl. = Puccinia purpurea, Cooke. 

spyridii, Cke. and Mass. Pomaderris apetala, Spyridium parvifolium 

tillaeae, McAlp. Tillaea sieberiana 

Uromyces Link ... 

*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 semibaccatuin 

betae (Pers. ) Kuehn. Beta vulgaris. (Introduced.) 

bicinctus, McAlp. Acacia fasciculif era ... ... 

bulbinis, Thuem. Bulbine bulbosa 

caryophyllinus (Schrank) Schroet. Dianthus caryophyllus, D chinensis. 

danthoniae, McAlp. Danthonia sp., D. semiannularis ... 

digitatiM, W int. = Uromyces phyllodiorum (B. and Br. ) McAlp. ... 

diploglottidis, Cke. and Mass. Diploglottis cunninghamii 

ehrhartae, McAlp Microlaena (Ehrharta) stipoides 

fabae (Pers. ) De Bary. Vicia faba. (Introduced.) 

fusisporus, Cke. atd 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 appendiculatus (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 ... 

polygoni (Pers.) Fckl. Polygonum aviculare. (Introduced.) 

puccinioides, Berk, and F. v. M. Selliera radicans, Scaevola sp. 

scleranthi, Rostr. Scleranthus diander 

tenuicutis, McAlp. Sporobolus indicus 

tepptrianus, Sacc. = Uromycladium tepperianum ( Sace. ) Me Alp. ... 

thelymitrae, McAlp Thelymitra antennifera, T. flexuosa 

tricorynes, McAlp. Tricoryne elatior ... 

trif olii (Alb. and Schw. ) Wint. Trifolium repens. (Introduced.) 

vesiculosus, Wint. Zygophyllum billardieri, Z. glaucescens 





















Fungus Index. 341 


Uromycladium McAlp. ... ... ... ... ... 104 

alpinum, Me Alp. Acacia buxifolia, A. dallachiana, A. dealbata, A. implexa, 

A. linifolia ... ... ... ... ... ... ... 105 

bisporum, McAlp. Acacia dealbata ... ... ... ... ... 100 

maritimum, McAlp. Acacia longifolia ... ... ... ... 107 

notabile (Ludw.) McAlp. Acacia binervata, A. dealbata, A. decurrens, A. 

elata, A. notabilis, A. pruinosa ... ... ... ... ... 108 

robinsoni, McAlp. Acacia melanoxylon ... ... ... ... 109 

simplex, McAlp. Acacia pycnantha ... ... ... ... ... llo 

tepperianum (Sacc.) McAlp. Acacia armata A. diffusa, A. erioclada, A 
glaucoptera, A. hakeoides, A. implexa, A. juniperina, A. lonpifolia, A. 
melanoxylon, A. myrtifolia, A. pycnantha, A. rigens, A. salicina. A. 
siculiformis, A. spinescens, A. stricta, A. verniciflua, A. verticillata, 
A. vomeriformis ... ... ... ... ... ... ... Ill 

General Index. 



Acacia, gall fungi ... ... 

Kangaroo ... 

Acacia armata ... 

dealbata ... 

decurrens ... 

implexa ... t>< 

pycnantha ... 

salicina .... ... 

Aecidiospores, binucleate ... 

coloured membrane in Gymno- 
sporangium ... 

colouring matter ... ... 

development ... ... 

distinction from uredospores 

germination in Puccinia 
tasmanica ... 

infection ... 



partaking of character of 

repeated formation... 
retention of germinating 

power ... 
Aecidium, colour 
invigorating power 

in heteroecious species 
on grass ... 
on seeds ... 
peridiurn ... 
primary and secondary 
pseudoperidial cells 
rep'eated formation 
sterile cells 

sexuality ... 14 

Aecidium abietinum 

bellidis ... ... ." 

berberidis, spore germination 

platylobii, mycelium in seed 

Alopecurus pratensis with Puc- 
cinia graminis ... 
Amphispore ... 

Anagallis arvensis, importation 
Apium, graveolens (Puccinia) ... 

prostrat um ( Puccinia) 
Asclepia'ieae, Oronartiam 
Ascomycetes :.. 
Asparagus rust (Puccinia aspar- 

agi), distributed by wind 
water relation 
Atmospheric influences on spore 

A uriculariaceae 
Australian rusts, number 







18, 57 

17, 75 
22, 36 





5 80 








A i PAOB - 

Autoecious species ... ... 10, 55 

most numerous ... ' j.-, 

repetition of aecidia ... ]s 

A vena elatior with Puccinic 

graminis ... gg 

Baeodromus ... 

Barberry, connexion with wheat 

rust ... KK KO 

. ... oo, OH 

in Australia ... gg 

infection experiments, local 60 

in Kew gardens ... ... gg 

Barberry rust (Aecidium ber 

beridis) ... ... jg 

absent in Australia " gg 

connexion with wheat rust .. 55 

increasing vigor of wheat rust 58 

local attempts at inoculation 66 

Barclayella, ... ... .. 33 

Basidia in aecidia ... ... ig 

Basidiomycetes ... 40 
Bean rust (Uromycesfabae) ... 20 
Beckmanniaerucaeformis, importa- 
tion of rust on seed ... 43 
Beet rust (Uromyces betae) ... 84 
Binucleate spores ... ... 14 

Biologic forms, and classification 79 

evolution of ... ... 54 

Bismarck brown .. ... 12 

Black wattle gall fungus (Uro- 

mycladium notabile) ... li 

Brachypuccinia ... ... 11 

Bremia lactuca, effect of copper 

salts ... ... ... 63 

Bridging species ... ... 53 

Brome rust (Puccinia brotnina) ... 3 

infection by uredospores ... 3 

starvation of host ... ... 60 

uninfluenced by structural 

peculiarities of leaf ... 61 

Brown Rust (Puccinia dispersa) ... 79 
Burning stubble, effect on wheat 

rust ... ... ... 72 

Caeoma ... ... ... 17, 36 

Caeomospores ... ... ... 36 

Carnation rust (Uromyces caryo- 

phyllinus) ... ... 85 

Caustic potash for softening tissues 12 

Cecidomyia, eating rust spores ... 7 

Cedar apples ... ... ... 6 

Celery rust (Puccinia thuemeni) ... 42 
Cereal rusts not liable to affect 
other cereals indiscrimin- 
ately ... ... ... 53 

Chemotaxis, parasitism due to 

positive and negative ... 52, 62 
Chick weed rust (Puccinia aren- 

ariae) ... ... ... *'-' 

Chicory rust (Pua-inia rirhorii) . . 42 
Chrysanthemum rust (Puccinia 

chrysanthemi) ... ... -1 

germination of uredospores 

importation ... ... 43 


General Index. 

Chrysomyxa, absence of fir trees 58 

germination of teleutospore 34 

origin of species in Alps ... 56 

uredospores in chains ... 19 

Chrysomyxa, abietis ... ... 66 

ledi ... ... ... 56 

rhododendri ... ... 56 

Chrysopsora, teleutospore of ex- 
ceptional form ... ... 32 

Classification and biologic forms 

of Uredines ... ... 79 

Clover rust (Uromyces trifolii) ... 18, 84 

Cluster cups ,.. ... ... 16 

Coleospcriaceae ... ... 82 

Coleosporium, function of paraphyses 30 

teleutospore in ... ... 32 

uredospores in cha ins ... 19 

Colecsporium senecionis, and fir trees 58 

Collema, fertilisation in .. 37 

Compositae, native rusts on .. 45 

Compound teleutospores .. 24 

Copper salts, influence on fungi.. 62 

Cronartiaceae ... ... .. .82 

Cronartium, asclepiadeum on varied 

hosts ... ... ... 53 

jacksoniae producing witches' 

brooms ... ... ... 5 

Crossbreeding wheats for rust- 
resistance ... ... 74 

Crown or coronate rusts ... 56 

Cucumber diseases and copper salts 63 
Cycle of development, in heteroe- 

cious species ... ... 58 

in rusts ... ... ... 2, 10 

Cyperaceae, number of native rusts 

on ... ... ... 45 

Dactylis glomerata ... ... 53 

Daisy rust (Puccinia distincta) ... 10 
introduced on seed ... ... 209 

Danthonia aecidium ... ... 18, 57 

Darluca ftlum, a common parasite 22 

mistaken f or spermogonia 22, 158, 165 

Development of fungi ... ... 40 

Dew necessary for infection ... 10 

Diorchidium ... ... ... 83 

doubtful genus ... ... 82 

Distribution of Australian species 50 

Distribution of spores... ... 7 

Doubtful species ... ... 207 

Drainage, effect on rust ... 71 
Dry atmosphere and spore ger- 
mination ... ... 10 

Early maturing wheat escaping rust 74 

Echinulate uredospores... ... 19 

Effect of rust on straw and grain 64 
Elymus condensatus, importation 

of rust on seed ... ... 43 

Endochrome ... ... ... 17 

Endophyllum, teleutospores ... 23 

Endospore, in aecidiospores ... 16 

in teleutospores ... ... 23 

Enzymes in leaf cells ... ... 54 

Epispore of teleutospore ... 23 

Epiteospores ... ... ... 20 

Erysiphaceae, specialisation of 

parasitism ... ... 54 

Erysiphe graminis, biologic forms 54 

Eucalyptus globulus, supposed rust 49 

Eu2)horbia, aecidium of Pea rust 55 

dentata ... ... ... 70 

rust ( Uromyces euphorbiae) ... 70 
Eupuccinia ... ... ... 11 

Excluded species ... ... 207 

Fallowing, effect on rust ... 72 

Fertilisation, in aecidium ... 17, 75 

in Uredineae ... ... 14, 75 

Fir trees and rust ... 58 

Flax rust (Melampsora lini) ... 43, 60 
first record in Australia ... 44 
Formalin treatment of seed wheat 73 
Frangula alnus :.. ... 53 

Fungi, origin and development ... 40 
Fusion of nuclei ... ... 14, 75 

Gall formation... 

Gall fungi, on wattles (Acacias) ... 

Genera, Australian, and number of 

species ... 
General parasites 
Germinating power, duration, in 

in teleutospores 
in uredospores 

Germinating spores, methods for 
aecidiospores and uredo- 
spores ... 
Germination, of aecidiospores of 

Puccinia tasmanica 
of amphispores 
of teleutospores, factors in- 

of Puccinia graminis 8, 24, 66 
of P. malvacearum ... 24, 27 

of Phragmidium rubi abnor- 
mal ... 
of uredospores, of Puccinia 


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 

solitary in Puccinia mono- 

pora ... 
Glycerine, and water ... 

jelly ... 

Golden rust (Puccinia glumarum) 

Golden wattle galls 

Gradations of specific variation ... 

Gramineae, native rusts on 

Grass aecidium 

Grasses, and heteroecious rusts ... 

and wheat rust 
Groundsel rust (Puccinia tas- 
manica) ... ... 16, 49 

Gymnoconia, characters ... 
Gymnosporangium, aecidiospores 
colored membrane 


insects conveying spondiola 

sculpturing of peri dial cells ... 






3, 8 








18, 57. 

' 57 






General Index. 


Gymnosporangium clavariae forme, 
direct infection by teleuto- 
spores ... 
confusum, promycelial cells 

teleutospores, thickness of 

cell wall 

Hard i issues, softening... 
Harvesting methods and rust 

Hetianthus annuus, aecidia 
J-J emibasidii ... 
Hemipuccinia ... 
Heteroecious species 
regular development 
Heteroecism ... 
beneficial ... 
causing increased vigor 
discovery ... 

when possible 
Hieracium, Puccinia ... 
Hollyhock rust (Puccinia malvace 

arum) ... 

wide distribution ... 

Honeydew of spermogonia 
Host and parasite, relation 
Host-plants, imported, and new 


indigenous, and rusts 

Hot- water treatment of seed 
Hypertrophy of nettle 
Immunity acquired 

and predisposition .. 
attempts to secure ... 
due to physiological peculiari- 

factors influencing ... 
Importation of rusts ... 
on cuttings 
on seed ... 
on straw ... 
Incubation period 
Indigenous species, table of 
and hosts... 
Infection, aided by injury 

and age of parts attacked 
by aecidiospores 
by internal germ of disease 
By sporidiola 

germ tubes entering stomata 
by teleutospores ... 
by uredospores 

germ tubes piercing epider- 
mal cells 

dew necessary for ... 
experiments, with barberry in 

with foreign host plants 
factors influencing 




















9, 60 








Injuries, influence on infection ... 
Inoculation, protective... 
Insects attracted, by colour of 
spores ... 

by honey dew 

by scent of spermogonia 

conveying sporidiola 

eating spores 
Intermediate hosts of wheat rust 

Internal developmental tendencies 
Introduced species 

how introduced 

on native hosts 
Investigation of spores... 
Irrigation and wheat rust 
Kangaroo thorn, gall fungus 
Knot- weed rust ( Urcmyctspolygoni) 
Lactic acid showing up germ pores 
Ledum palustre 
Leguminosae, native rusts on 
Leptopuccinia ... 

with warted 






42, 4!) 




Lettuce, attempt to confer immu- 
nity from disease 
Liability to disease 

influenced or not by structural 

Lichen-furgi ... 
Life -cycles of rusts 

Life history of rusts 
Liliaceae, native rusts on 
Linum marginale 

rust -resisting variety 
Lolium perenne, crown rust 
with introduced rust 

Mahonia in Kew gardens, rust free 

Maize rust (Puccinia maydii] 

Mallow rust (Puccinia ma/i-acearum) 

Manures and wheat rust 

Marigold rust (Puccinia calendulae) 10, 42 

Measuring spores by photography 

Melampsora, germination of teleu- 

Melampsora eucalypti, merely 



Mesospores, in Puccinia 

in Uromycladium ... 
Mint, native, rust (Puccinia 


Mounting spores 
Mycelium, development of 
in seed of annuals ... 

localized, causing hypertrophy 
perennial causing galls 

no evidence in wheat gram 
of Puccinia arrhenatheri ... 
of Uromyces euphorbiae ... 









... 22, 49 


... 25, 26 








General Index, 

Mycoplasm theory 

bearing on rust in wheat 
Native celery ... 


mint ... ... .. 

rusts, and native hosts 

on imported hosts 
Nettle rust (Aecidium urticae) ... 
Nuclear cycle ... 
Nuclei, fusion of 
Oat, wild, and wheat rust 
Obligate parasites 

Odour of spermogonia ... 
Omission of spore-forms 
Origin of heteroecism ... 

of parasitism 

of spore- forms 
Paraphyses, function ... 

in aecidia 

in spermogonia 

in teleutosori ... 22, 

in uredosori 

Australian species with . . . 
Parasitism, origin 

Pea rust ( Uromyces pisi) 
Peach or prune rust (Puccinia 

pruni) .... 

Pedicel of teleutospore ... 
Peltandra virginica 
Perennial mycelium 
Peridermium ... 

Peridial cells 

Peridium, = Pseudoperidium 

often wanting 

varied forms 
Phalaris, Puccinia 
Photography in measuring spores 
Phragmidium barnardi, germ pores 



rubi, teleutospore germination 


paraphyses, in aecidia 

in uredosori 
perennial mycelium 
Phragmitcs communis, Puccinia 
Phy corny cetes ... 
Phytopthora infestans, and copper 

Picea excelsa ... 

morinda . . 

Ploughing, influence on wheat rust 
Plum or prune rust (Puccinia 

pruni) ... 
Poa annua 

pratensis ... 

Polygonum avicu lare 
Potash, caustic, for softening 

tissues ... 

Promycelial spores (see Spori- 

4, 74 

42, 44 

14, 75 

22, 30 

13, 29 

24, 75 







5, 36 


29, 42 




Promycelium ... ... j 

abnormal ... ... ... 35 

cells of, separating ... 35 

elongated ... ... . . 27 

in various genera ... .. 33 

septa ... ... . . 27 

Protective inoculation ... .. 62 

Protobasidii ... ... . . 40 

Protomycelium ... . 4 

Pseudoparenchyma ... . . 4 

Pseudoperidial cells (see Peridial cells) 
Pseudoperidium (see Peridium) 

Puccinia, introduced species ... 42 

variation in teleutospores ... 39 

Puccinia agropyri, heteroecious ... 45 

agropyrina ... ... 53 

agrostidis ... ... ... 45 

anthoxanthi ... ... 42 

arenariae, introduced ... 42 
sporidiola germ-tubes enter- 
ing by stomata ... 10 
arrhenatheri, perennial myce- 
lium ... ... ... gg 

asparagi, influence of wind ... 7 

water relation ... ... 9 

beckmanniae, imported ... 42 
bromina contrasted with P. 

triticina... ... ... 80 

paraphyses in teleutosori ... 24, 75 
uredospore, duration of 

germinating power ... 8 

germination and infection 3, 8 

calendulae, marigold rust ... 10 

Australian species ... 49 

short cycle ... ... 10 

caricis, heteroecious ... 45 

cesatii ... ... ... 25 

chrysanthemi, germination of 

uredospores ... ... 21 

introduced ... ... 42 

two-celled uredospores ... 19 

cichorii, introduced ... 42 

cinerariae... ... ... 49 

coronata ... ... . 53 

cyani, introduced ... ... 42 

dianthi, sporidiola germ -tubes 

entering stomata ... 10 
ditkondrae, variation in 

teleutospores ... ... 39 

dispersa, biologic forms ... 79 

subdivision ... ... 53 

distincta, daisy rust, Aus- 
tralian species ... 49 
introduced on seed ... 209 
short cycle ... ... 10 

festucae, introduced ... 42 

glumarum, mycoplasm theory 4 
gramineOa, with aecidium on * 

grass ... ... ... 18 

graminis, biologic forms ... 79 
carried over from year to 


in Australia ... 21 , 

collective species 
cycle of development ... 
forms on special hosts ... 
heteroecism increasing 
vigor ... 




General Index. 


Pucdnia graminis, importation on 

straw ... ... 43 

injurious rust in Australia 64 

no aecidia in Australia ... 58 

on grasses, wheat, &c. ... 70 

repeated formation of 

uredospores ... 19 
reproduced by uredo- 
spores in Australia ... 19 
specialisation of forms ... 53 
teleutosporeSjfrom Australia 
not germinating in Eng- 
land, and vice v< rsd ... 67 
germinating power ... 8 
germination, exposure to 

cold not necessary ... 67 
in Australia... ... 24, 67 

in water ... ... 36 

time for ... ... 8 

variation ... ... 39 

uredospores, germinating 

power ... ... 8 

on old straw ... 69 

germination ... ... 21 

persist through winter ... 8 
graminis and barberry rust, 

connexion first discovered 55 
in Australia ... ... 58, 66 

in Kew Gardens ... ... 68 

helianthi, germination of 

teleutospore ... ... 9 

heterospora, one -celled teleuto- 

spores most common ... 26 

transition form ... ... 83 

hieracii, spermogonia with 

uredo... ... ... 13 

subdivision of ... ... 80 

impatientis, introduced ... 43 
liliacearum, spermogonia with 

teleuto ... ... 13, 37 

lolii, introduced ... ... 42 

paraphyses in uredosori ... 22 

magnusiana, heteroecious ... 45 
paraphyses, in teleutosori 

and uredosori ... 22 
malvacearum, causing "shot- 

hole" ... ... 6 

germination of teleutospores 24, 27 
importation and widespread 

distribution ... ... 43 

suppression of spermogonia 14 

maydis, introduced... ... 42 

menthae, introduced ... 42 
obtegens, spermogonia with 

uredo... ... ... 37 

perplexans, on native host ... 49 

paraphyses in uredosori ... 29 

plagianthi, epispore ... 23 

poarum, introduced ... 42 

paraphyses in uredosori ... 22, 29 

uredo withstanding cold ... 22 

podolepidis, two germ pores in 

upper cell of teleutospores 35 
prainiana, abnormal teleuto- 
spore germination ... 36 
prenanthis, introduced ... 42 
peridium often wanting ... 17 

Pucdnia pruni, causing "shot-hole" 6 
first record in Australia ... 44 
producing uredospores 

within fruits ... ... 19 

supposed spermogonia ... 22 
purpurea, introduced ... 42 

rufiifjovera, germinatingpower 

of uredospores 

paraphyses in teleutosori .. 24 
subdivision ... ... 79 

senecionis, repetition of aeeulia ! s 

xiiitfih-x, aecidia unknown ... 58 

introduced ... ... .jj 

mesospores ... ... _>; 

suavcolens, scent of sper- 
mogonia ... ... 13 

tasmanica, germination ol 

aecidiospores ... ... 16 

groundsel rust ... ... 42, 49 

thuemeni, introduced rust 

attacking native host .. 42, 49 

triticina, aecidia unknown 58 

comparatively harmless ... 64, 66 

contrasted with P. bromina 80 

germination of uredospores 21 

paraphyses in teleutosori ... 2*J, 7"- 

preference for certain parts 60 

vexans, amphispores ... 25 

Pucciniaceae ... ... ... 82 

Pucciniopsis ... ... ... 11 

Queen's Jubilee wheat, rust spores 

on grain ... ... ... 69 

Ranuculaceae, Cronartium ... 53 
Ranunculus Jicaria, infection ex- 
periments ... 56 
Ravenelia ... ... ... 84 

compared with Uromydadium 104 
Repetition, of aecidia ... ... 18 

of spore forms ... ... 11 

and spermogonia... ... 14 

of uredospores ... .. 19 

Rerraf, a rust-resisting wheat ... 64, 74 


Rhododendron, Chrysomyxa ... 56 
Roestelia, ... ... ... 17 

Rose rust (Phragmidium subcorti- 

cium) ... ... 5 

Rotation of crops and rust ... 71 
Rubiaceae, native rusts... 45 

Rust Conference 
Rust of wheat... 

connexion with rust on grasses 
early theories ... ... 68 

effect of, drainage ... ... 71 

early maturing wheats ... 
fallowing ... ... 7 '2 



muggy weather ... 67, 7'- 


rotation of crops 

seedbed Tl 

seed treatment 
stubble burning 
effect on straw and grain ... 
first appearance in season ... 
first record in Australia ... 74 


General Index. 

Rust of wheat, how carried over 

from year to year ... 21, 68 

how spread ... ... 69 

infection from within, my co- 
plasm theory ... ... 4, 70 

influence on yield ... ... 65 

losses from ... ... 64 

mycelium not in seed ... 70 

mycoplasm theory ... 4, 70 

not spread from other cereals 53 

prevention and mitigation ... 71 
question, present position in 

Australia ... ... 64 

spores, in soil ... ... 69 

on seed ... ... 69 

to destroy ... ... 73 

wintering ... ... 8, 20 

spraying impracticable ... 73 

Rust-liable soils ... ... 72 

Rust -proof wheat unknown ... 62 
Rust-resistance and structural 

characters ... ... 61 

Rust-resisting wheats ... ... 62, 74 

Rusts, Australian, and their hosts 45 

cycle of development ... 2, 10 

in heteroecious species ... 58 

indigenous and introduced ... 42 

influence of parasitic habit ... 40 

relation to other fungi ... 40 

sexuality ... 14, 17, 75 

Rusty straw and grain analyzed 65 

Rye, rust ... ... ... 53 

Saccoblastia ovispora ... ... 32 

Saprophytes becoming parasitic 52 

Scarlet pimpernel ... ... 43 

Scent of spermogonia ... ... 13 

Scrophiilariaceae, Cronartium ... 53 

Secale ... ... ... 80 

Seed, perennial mycelium ... 5 

not in wheat ... ... 70 

Seed-bed" and rust ... ... 71 

Seed-treatment for rust ... 73 

Seeds with aecidia ... ... 70 

Selection and crossbreeding wheat 74 
Self-sown wheat and rust 21, 69, 72 

Senecio vulgaris ... ... 16 


formation in promycelia ... 27 
Sexuality and nuclei ... ...14,75 

of aecidia ... ... 15, 17 

Shot-hole caused, by Puccinia mal- 

vacearum ... ... 6 

by P. pruni ... .... 6 

Silver wattle gall fungus ( Uromy- 

cladium notabile) ... ... 6 

Softening tissues for microscopical 

examination ... ... 12 

Soil moisture, influence on rust ... 10 

Soils, rust-liable and rust- free .., 72 
treatment with copper salts 

to prevent disease ... 62 

Sorus, uredo ... ... ... 19 

Special corpuscles ... ... 4 

Specialisation of parasitism ... 52 

in Erysiphaceae ... ... 54 

Specific variation, gradations ... 79 
Spermatia, embedded in sugary 

secretion ... ... 13 

formation... ... ... 13, >8 

germination ... ... 13, 37 

not male cells ... ... 13 

uninucleate ... ... 14 

Spermogonia, associated with cer- 
tain spore forms ... ... 1 3, 37 

attractive to insects ... 13, 38 

Darluca flum mistaken for... 22 
functionless ... 13, 37, 39 

importance ... ... 14 

occurrence ... ... 13 

origin ... ... ... 38 

paraphyses ... ... 13 

repetition of spore forms ... 14 
suppressed ... ...14,37 

Spore-bed ... ... ... 19 

Spoie-forms ... ... ... 7 

investigation .. ... 11 

origin of principal ... ... 31 

repetition... ... 11, 18, 19 

suppression ... ... 10 

Spores, distribution ... ... 7 

germination, of aecidiospores 7, 16 
of amphispores ... ... 25 

of spermatia ... ... 13, 37 

of sporidiola ... ... 27 

of teleutospores ... 8, 24, 27, 66 
abnormal ... ... 35 

of uredospores ... 7,19,21,27 
nuclei ... ... ... 14 

on seed ... ... ... 69 

parasites ... ... ... 7, 22 

wintering, of teleutospores ... 8 

of uredospores ... ... 8,20 

Sporidiola, air necessary for for- 
mation ... ... 27 

conveyed by insects ... 28 

distribution ... ... 28 

earliest spore form... ... 32 

formation... ... ... 27 

germination ... ... 27 

incapable of infecting grasses 

bearing them ... ... 59 

infection ... ... ... 9, 28 

origin ... ... ... 32 

uninucleate ... ... 14 

Spraying for rust of wheat im- 
practicable 73 

Staining spores ... ... 12 

Starvation of host and infective 

power of rust ... ... 60 

Stellaria media ... .. 43 

Sterigma ... ... ... 31 

Structural characters of host and 

susceptibility, to disease ... 61 
to wheat rust ... ... 61 

Stubble burning and rust ... 72 

Sugar, influence on germ tubes of 

fungi ... ... ... 52, 62 

Sulphate of copper, in seed treat- 
ment for rust ... ... 73 

treatment of soil with ... 62 

Suppression of spore forms ... 10 

Surface markings of spores ... 12 

General Index. 


Susceptibility of host, due to 

physiological peculiarities 
influence of structural charac- 
ters 61 

Teleutospores ... 
. direct infection by germ tube 
function ... 

germination, factors influenc- 
ing ... 

in Phragmidium rubi 
in Puccinia graminis 
in P. malvacearum 




8, 24, 66 
.. 24, 27 

.. 33, 35 
... 24, 75 



origin and development 


simple or compound 

two kinds... 


variability, of form 

of germination ... 
Teleutsori paraphysate... 
Tomato, copper salts and disease 
Trifolium repens 
Triphragmium ulmariae ... 36 

primary and secondary uredo 

spores ... ... 20, 36 

spermogonia with uredo ... 13, 37 
Uninucleate spores ... 14 

Uredineae in relation to other 



Uredo dispersa, histology 
kuehnii, paraphyses 
spyridii, paraphyses 
symphyti ... 
Uredosori, paraphyses ... 

parasites ... 
Uredospores, binucleate 

derived from teleutospores 
function ... 
germinating power 

duration of 
germination, in Puccinia chry- 


in P. rubigovera ... 
germ tubes, piercing cells ... 
infection by germ tubes 

primary and secondary 
produced within fruit in Puc- 
cinia pruni 
repeated formation 
two celled, in Puccinia chry- 

santhemi ... 19 

wintering... ... 20, 21 

Uromyces ... ... 24, 83 

distinction between uredo- 
spores and teleutospores ... 

betae 42,84 

bulbinis ... ... "J 

caryophyllinus ... ... 43, 85 





22, 29 

22, 29 














Uromyces dactylidit, paraphyses .'$< > 

danthoniae, with aecidium on 

grass ... ... ...18,57 


euphorbiae, perennial mycelium 5,70 . 

j'abae in Ecuador ... .... 20 

limosellae ... ... ... 85 

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 ... ... 85 

scutettatus, origin of uredo- 

uredo with teleuto spore ... 36 
solidaginis ... ... 36 

tricorynes, two-celled teleuto- 
spores ... ... ... 39, 83 

trifolii complete cycle 
introduced ... ... 43 

perennial mycelium 
tuberculatus ... 35 

vesiculosus, two-celled teleu- 
tospores ... ... 39, 83 

Uromydadium 24, 83, 104 

forming galls often 

mesospores ... 25, 26 



vesicles ... ... - 24 

Uromydadium maritimum, occur- 
rence of spermogonia ... 13, 37 
notabile ... - 1* 

perennial mycelium and 

galls ... -. 6 

robinsoni, spermogonia ... i> 
tepperianum, spermpgonia ... 13, 3/ 
perennial mycelium and 



Ustilagines ... 

Vegetative organs, mycelium ... 3 

Verrucose uredospores ... 

Vesicle in Uromydadium 

Violet rust, native (Puccima 

Water relation, for Puccinia on 


to infection, direct and in- 
direct ... 

Wattles, gall- fungi on ... ... 

Wheat rust. See Rust of Wheat. 

Wheat, rust-resisting ...' J 

self-sown and rust ... ^ b., /- 

Wild oats and wheat rust ... J 

Wind distribution of spores ... / 

Wintering of rust spores 

Witches' brooms 

By Authority: ROBT. S. BRAIN, Government Printer, Melbourne. 




MAR 1.6 
MAR 2 3 1536 

APR 1 1935T 

JAN 23 1963 

LD 21 3m-8,'3 

MoAlpine, D 
The ruats 


of Australia 

V . J 


8 3 1C