Skip to main content

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

See other formats

i i 

f * 

* * 





* 9 


* i 





♦B fiOfl MM5 



i : 




» ," ' f * 



■ ■ 



i * 



*< i 





| Ki'll 












> ♦ - 

wjj! y 


jljlijj [J 

kj ■■ . ' i ■ ( 


4 I 

1 ft \ 





€*m wv 

- 9 

























Government Vegetable Pathologist. 






1 906. 


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

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

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

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

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

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

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

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

in drawing from microscopic preparations. 

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



Handbook of Australian Fungi. When one considers that the material 

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

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

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

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


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

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

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

Melbourne, March, 1906. 









I. Introduction 

• • 

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

IV. Spermogonia and Spermatia 
V. Aecidia and Aecidiospores 

VI. Uredospores 

VII. Teleutospores ... 

VIII. Mesospores and Amphispores 

IX. Sporidiola or Promycelial Spores 

X. Paraphyses and their Function 

• - 


• • 

• • 

■ - • 

• * « 

• • • 

• • 

XI. Origin of the Principal Spore-forms 

- 9 

Rusts in their relation to other Fungi 

XIII. Indigenous and Introduced Species 

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

XVI. Origin and Specialisation of Parasitism 

XVII. Heteroecism and its Origin 

XVIII. Predisposition ... 

t * 

• * 

• • 

• • 

• • • 

• • 

- * - 

• • 

• • • 

■ • ■ 

• 9 

• * 

• • 

• • • 

• * 

• • 

The present position of the Rust in Wheat question in Austr 












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

Uromyces ... 

• • • 


• • 





• # 

Aecidium . . . 


• • • 

• • • 

• • 

• • 

# • 

• t • 

• • 

• • • 

t * 

« • 

• • • 

• • • 

* • 

• • 

• • • 

• • 

• • • 

• • 

Excluded or Doubtful Species 

Glossary of the principal scientific terms used 

Literature consulted 

Explanation of Plates 

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

• • 

- - - 

• • 

• • 

» - 

- - • 

• • 

• • 

9 • 

• • 













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

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

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

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

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

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

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

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

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

teleutospore ; or 

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

succession, the aecidiospores being generally preceded or 

accompanied by a peculiar form of spore, known as a 

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. 



This succession of spore-for 


be represented 

lly by the 

following diagrams 






Fig. 1. 


Fig. 1. 




I 'redospore 

Fig. 3. 

The first show 


plete cycle of develop 

spore- forms follow each other in 

common form 


invariable ( 

the second the uredosp 


Fig. 4. 

lent in which all the 
and this is the most 



d the 

number of species unde 
the third the aeeidiosix 

this contracted cycle is much reduced 

further reduced 



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

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




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

fe -cycle, but 
Besides th 


general development. 

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

species o 
are know 

degraded forms of which onlv the uredo 


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

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


sketched ; 

Vegetative Organs. 



Vegetative Organs — Mycelium. 

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

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

Among the recent investigations on the mycelium, there are two which 
stand out on account of their completeness, owing to the use of the most 
modern histological methods — the one by Professor Marshall Ward 9 on the 
Histology of Uredo dispersa Eriks, and the Mycoplasm hypothesis, and the 
other by Professor Eriksson u > 15 on Das vegetative Lcben der Getreiderost- 
pilze [The Vegetative Life of the Cereal Rusts]. The study of an indivi- 
dual case will prepare us for the more general examination of the 
mycelium throughout the rusts, and we will begin with that of Puccinia 
dispersa Eriks., or, strictly speaking, P. bromina Eriks., which Ward has 
so thoroughly dealt with and illustrated with such admirable clearness. 

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

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

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

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


Vegetative Organs. 

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

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

h the mycelium arises afresh 


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


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

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

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

Vegetative Organs. 


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

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

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

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


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


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

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

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

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


Vegetative Organs. 

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

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

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

& v-^"- "^f) 


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

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

In other cases the affected tissues mav be so stimulated bv 
mycelium as to cause their death. Thus almond Ip 9vm Kni, 

been found 

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



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

Reproductive Organs 



Reproductive Organs — Spores. 

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


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

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

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

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

Insects as well as other animals serve the same purpose. I have often 
observed the larvae of a species of Cecidomyia feeding on uredo and 
aecidio-spores, and at the same time spreading them while crawling along. 
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 anv 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 easilv bored in them, and glass rods placed parallel 
ietween them, so that the microscopic slides may be laid across the rods. 


Reproductive Organs. 

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

that is necessan 

Sometimes, however, the spores 

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

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

Duration of Germinating Power. 

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

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

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

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

drv. % . 

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

wheat if the uredospores can retain their germinating power during the 
winter. The results vary, as might be expected, according to the condi- 
tions prevailing at the time. In contrast to De Bary's results, Eriksson 
found that the uredospores of P. graminis lost their germinating power 
during the winter if kept in the open, but retained it if kept inside. 
Hitchcock and 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 
une to August). The uredospores can germinate at once, and directly 
feet the host-plant, or they can act as resting spores for a time, and 
freely infect the next season's growth, under the climatic conditions which 

prevail here. 

The germination of teleutospores takes place ait different periods, ac- 
cording to the nature of the species. They may either germinate imme- 
diately on reaching maturity, or, as is the case in the majority of heteroecious 
iusts, only after undergoing a period of rest — in the old w r orld 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 

and then on exposing them for another 

he found that they germinated, but this was exceptional 

Reproductive Organs 


In other species there is considerable variation. According to 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 itfo the formation of fresh spores. The first visible trace of 
this does not generally appear before eight days, but, according to the 
nature of the fungus, it may take much longer. Schimper, in 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 Puccinia on asparagus, and probably the same prin- 
ciple applies in a general way to other rusts. From a study of the direct 


Reproductive Organs. 

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

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


Suppression or Omission of S 

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


The aecidio, uredo, and teleutospore forms are represented respectively 

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

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

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

the cycle, although the 


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

hich. however, is rarelv absent 

diolum produces 

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

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

Reproductive Organs. 


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

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

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

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

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

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

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

Repetition of Spore-forms. 

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

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

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

Investigation of Spores. . 

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


Reproductive Organs. 

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

sometimes been overlooked by otherwise good observers. As a general 

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

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

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

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

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

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

guibiiea wirn a 

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


and the results compared. All the microscopic investigations were made 

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

Spermogonia and Spermatid. 13 


Spermogonia and Spermatia. 

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

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

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

the regular development of the fungus. 

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

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

14 Spermogonia and Spermatia. 

continue their further development. In a paper by Arthur 3 on the Taxo- 
nomic Importance of the Spermogonium, he shows that, in the first pla^t, 
• he presence of the spermogonium, along with its associated spore-forms, 
gives important information regarding the length of the life-cycle. Thus 
he concludes that if spermatia and uredospores arise from the same 
mycelium, aecidia do not occur, and if spermatia and teleutospores thus 
arise, neither uredo nor aecidio spores will occur. There are cases, how- 
ever, in which the spermogonia are found in association with both uredo 
and teleuto spores, as in Uromycladium maritimum, U. notabile, and U. 
robinsoni. When the spermatia are associated with aecidia, it is only where 
teleutospores and aecidia arise from the same mycelium that it can be 
definitely stated that there are no uredospores. 

In the second place, the characters of the spermogonium, such as posi- 
tion, size, form, and colour, and its relative position to the accompanying 
spore-forms, furnish characters for positive identification, although of 
minor value. There is an interesting relation of the spermogonium to 
other spore-forms to be noted in the cases afterwards referred to, where 
there is a repeated formation of aecidiospores, uredospores, or teleuto- 
spores, as the case may be, and these spore-forms do not directly proceed 
to their normal development. In such cases, as far as our present know- 
ledge goes, the spermogonium is the only one which does not repeat itself, 
as stated by 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 malvacearum and similar Lepto-Uredineae, the spermogonium 

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

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

Sappin-Trouffy 2 , for instance, has observed in Uredineae having the 
various stages the following nuclear cycle: — 

Uni-nucleate. — 

Mature teleutospores. 

Mycelium, 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 
the nuclei may be interpreted otherwise. Blackman 2 has confirmed the 
above nuclear cvcle. and concludes that the spermatia are male cells which 

have become functionless, the nuclear characters being those 

d not of 

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


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

Sfermogonia and Sfermatia. 

J 5 

Massee also considers the 

dium to be a sexual product, and both 

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

d aecidia represented 


and femal 

productive organs, a distinct 

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

bearing stage 

or gametophvte starting 

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


d its presence is an indication that the 

of th 


possessed an 

of generations similar to that referred to abo\ 


!6 Aecidia and Aecidiospores 


Aecidia and Aecidiospores. 

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

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

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

coloured, as a rule, 

orange- red oil, although this 

also occurs in many uredo and teleutospores. 

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

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

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

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

Aecidia and Aecidiospores. 17 

The development of the aecidia has been traced by De Bary, Neumann 1 , 
and others. The hyphae derived from the promycelial spores form at 
certain points, v deep down in the parenchyma of the leaf, little compact 
bodies which have been called primordia by De Bary, because they are 
the beginnings of the aecidia. These bodies gradually increase in size 
by the rapid multiplication of the mycelium until they assume a spherical 
form, only being slightly flattened at the part immediately beneath the 
epidermis, and considerably thickened at the base. If sections aie made at 
this stage the differentiation of the cells is seen to have begun. From the 
dense basal layer of hyphae arise numerous, closely-crowded, short, erect 
hyphae, somewhat club-shaped, and generally known as basidia. From 
each of these erect hyphae an apical cell is separated off, and beneath that 
another, and so on until a linear series is formed. At the same time, as the 
basal cells give rise to the so-called basidia, the surrounding cells develop 
into the external envelope or peridium. At. first they are absolutely indis- 
tinguishable from the others, but after the third or fourth division they are 
seen to be much broader. Then, simultaneously with the formation of the 
first spores, they acquire their characteristic markings and polygonal form. 
At first the peridial cells are filled with the coloured protoplasm or endo- 
chrome, but the colour gradually disappears. 

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

The origin of the aecidium has given rise to a good deal of discussion, 
and it is not universally accepted that it originates from non-sexual cells. 
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." 


Aecidia and Aecidiospores. 

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

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 






f iQ r f Aecidiospores 

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

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

give rise to teleutospores. H 

complete cycle, the aecidia originate only from sporidioi 


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





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

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

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

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

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

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

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



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






Fig. 6. 

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

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

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

hile those of 

and scattered on the under sur- 

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

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

A very striking case is recorded by 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 
autumn months of April and May, or even earlier, and the harvest is 



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

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

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

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

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

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

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 1st December to 5th February. And Miss Gibson kept 
spores of the same rust in a dry test-tube in a cool room for 71 days, from 
March to May, and at the end of that time about one quarter germinated, 
while a week after none germinated. 

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



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

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

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

Paraphyses most commonly occur in connexion with the uredosori, and 

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

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

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




Teleutospore s. 

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

As an example, Uredo sym-phyii, D'C. was considered by De Bary to 
be an independent species, and having lost its other spore-forms to be 
capable of existing without them, but 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, but generally 
it is darker than the uredo layer, being dark-brown to blackish, and only 
,rarely reddish. 

In the simplest cases teleutospores are unicellular, and originate in a 
similar manner to the uredospores. The spore-bed, consisting of inter- 
lacing and crowded hyphae, gives off erect branches, which become 
swollen at their free ends, and the finely granular protoplasmic contents 
are invested by an inner membrane, or endospore. in addition to the outer 
or epispore, which becomes relatively thick and dark in colour. The 
teleutospores are generally formed towards the end of the active vegetative 
period of the host-plant, and are often called winter spores in contrast 
to the uredo or summer spores. They are specially adapted and equipped 
for continuing the species over periods of drought, or damp, or cold, or 
seasons of scarcity. This is seen in the firm outer wall, which is often 
sculptured in various ways, as well as in the reserve material stored up in 
the contents. Sydow 1 states that in all Leptopuccinieae, or those only 
possessing (teleutospores which germinate at once, the epispore is perfectly 
smooth ; but there is one exception in Australia — P. plagianthi. In other 
groups the epispore may be smooth, warted, striated, &c. 

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

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






that it is now coming to be regarded 

or I 

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



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

In the newly -constituted Australian genus, Uromycladium, the 

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

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

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

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

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

conditions that they do not require such protection. The best 

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

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

be the season of the year, for they 

ger mini ate 

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

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

of P. malvacearum 

Mesospores and Amphispores. 25 


Mesospores and Amphispores. 

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

possess the same char 

The conclusion one would naturally d 

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

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

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

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

Ampiii spore. 

This peculiar kind of spore was first investigated in connexion with 
Puccinia vexans Farl. This species has a true uredo and teleuto-stage, 
in addition to a third form of unicellular spore, and the latter was the first 
to be recognised and recorded. In 1879 it was described as Uromyces 
brandegei by Peck, the unicellular spores being rough with minute warts, 
and therefore suggestive of Uromyces. Then, in 1883, Dr. 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 

b 2 

2 6 Mesospores and Amphispores. 

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

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

sorts of teleutospores. 

Up till recently this was the only instance known of the occurrence of 
amphispores, but 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, ius£ 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- 
spcra and P. simplex, in which the one-celled spores far outnumber the 
regular teleutospores, so much so that the latter have been frequently over- 
looked, and there is every probability that they undergo germination, and 
are therefore unicellular teleutospores in the fullest sense of the term. In 
P. heterospora there are no uredospores, and the unicellular spores are 
smooth, and otherwise resemble the teleutospores, while in P. simplex the 
uredospores are spinulose and yellow, and quite distinct from the smooth 
unicellular spores associated with the teleutospores. Owing to the teleuto- 
spores not having been obtained at first, the former has been variously 
named Uromyces pulcherrimus, B. and C, U. thwaitesii, B. and Br., and 
U. malvacearum, Speg., and the latter as Uromyces hordei, Rost. In fact, 
these spores have been regarded as a transition stage from the unicellular 
Uromyces to the bicelTular Puccinia spore. In such cases the one-celled 
spore functions as a teleutospore, and ithere would be a certain convenience 
in distinguishing between the undeveloped and immature spores and those 
which are fully formed, and in all probability capable of germination, 
reserving the term mesospore for the one, and unicellular teleutospore for 

other. But it is so difficult to d 


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

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


Sporidiola or Promycelial Spores. 27 


Sporidiola 1 or Promycelial Spores. 

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

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

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

28 Sporidiola or Promycelial Spores 

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

In the heteroecious rusts there is no evidence to show that the sporidiola 
can infect the plant bearing the teleutospores, hence it would appear ithat 
P. graminis in Australia can only be propagated, as far as spores are con- 
cerned, by means of the uredospores. It has not been definitely proved 
how the sporidiola are distributed, but no doubt the wind is an important 
factor, and probably also animals assist in the distribution. In connexion 
with the teleutospores of Gymno sporangium, 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 germinating teleutospores often lie upon the ground, it 
might seem at first sight as if they were not favorably situated for the wind 
to act upon them, but as they are usually produced in large numbers, suffi- 
cient of the sporidiola are likely to be suspended in the air to infect fresh 
plants when the conditions are suitable. 

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







Paraphyses and their function. 

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

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

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

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

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

structure and position. The wall is much thickened, so as to leave a very 
small cavity in the head, and in the stalk it may become so thick as 
completely to obliterate the cavity and thus render it solid. The 
thickened head has a great capacity for retaining moisture, and since they 
are crowded together and over-lap the spores, they will prevent them from 
getting soaked and at the same time protect them against excessive evapora- 
tion, which would have a drying effect according to 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 that their presence greatly 
depends upon some special condition of the fungus. " I find them con- 
stantly present with the uredospores of Puccinia perplexans, Plow., when 
these have arisen, not directly, but rather at a considerable distance from 
the aecidiospores. On the other hand, when the uredo arises directly 
from the aecidiospore, they are hardly present at all ; this looks very much 
as if they were an indication of exhaustion of vital energy on the part of 
the fungus, which was combated by protective efforts on the part of the 
parasite in conserving those spores which it does produce, but when full 
of vigour and fresh from the aecidiospore it is less careful of its spores. 
When it begins to feel the, effect of exhaustion, and is unable to develop 
such energetic spores, it takes more care of those which are produced." 

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



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


Another funct 

of protecti 

has been suggested 


in add 


In several species of Coleosp 
ed both for protection and for 

epidermis so as to make room for the growing spores 

m he found that the 
ng and bursting the 

To a certain extent paraph 

hastening its rupt 

may assist in raising the cuticle 


the absence of paraph v 

but there are plenty of species which rupt 


ted with teleutosp 

mg awa\ 

until it decay 

thev rather appea 

and in the case of parapl 


prevent the cuticle break 

Origin of Sf ore-forms 



Origin of the Principal Spore-forms. 

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

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

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

a. b 

Fig. 7. 

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

Fig. 8. 

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


Origin of Spore-forms. 


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

Fig. i). 


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

Fig. 10, 

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

Fig. 11. 

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


Origin of Spore-forms. 


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


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

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

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

Fm. 12. 

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

The next advance in the development of the teleutospore may be seen 
in an interesting genus, just described by Arthur 6 , to which he has given the 
name of Baeodromus. It occurs on Senecio, and resembles Coleosporium 


Origin of Spore-forms 

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

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


as a mu 

Fig. 13. 

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



The view that the uredospore is probably derived from the teleutospore 
is favoured by the variability of the latter in many species, and the grada- 
tions which are found to occur. At lirst sight the differences between the 
uredo and teleutospore seem so great as to be insurmountable, but there are 
distinct transitional forms from the one to the other. The membrane of the 
typical uredospore is covered with spines, and this is an evident adaptation 
for the spore which germinates immediately and is short-lived, and the 
characteristic spines serve to attach it to the surface of the host-plant in 
order that germination may be successfully accomplished. 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 which the wall still retains its thickness, but there 
are several germ-pores. They may resemble the uredospores in shape, 

Origin of Spore-forms. 


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

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

Kienitz-Gerloff 1 has shown that in Gymnosporangium clavariae forme, 
J acq., the thin- walled teleutospores do not produce a promycelium and 
promycelial spores, but simply a germ-tube like that of a uredospore. And 
Dietel 3 has further shown that both thick and thin- walled spores may 
sometimes germinate in this way, and thus serve the purpose of the uredo- 
spore occurring in other genera. 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 ordina.ry course, rounded 
themselves off and separated. Fischer observed in Gymnosporangium 
confusum that the cells of the promycelium separated before forming sterig- 
mata and promycelial spores, but Blackman, in a MS. note to his 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 finallv the promycelium bearing promycelial 

-,<5 Origin of Spore-forms. 


,pu«« It is worthy of mention that Magnus 2 has repeatedly obseived 
that when the teleutospores of Puccinia graminis were germinated in water 
thev produced an undivided germ-tube just like that of a uredospore, but 
be could not satisfactorily settle the question, if this germ-tube could directly 

penetrate the host-plant. 

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

This peculiar germination of the teleutospore, in which each promy- 
celium breaks up into three or four detached cells', apparently representing 
sporidiola, had been observed by Barclay 3 as early as 1891, in Pucctma 
lainiana, BarcL, and Uromyces solidaginis, Niessl. He considered this 
abnormal mode of germination to be due to the restricted supply of air 
obtained bv the spores in a hanging drop of water, for when allowed to 

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


tion was normal. 

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

In another respect the uredospores show a distinct^ transition. In a 
well -developed spore-layer they always precede the associated teleutospores, 
but in Uromyces scutellatus, as Magnus 2 has repeatedly observed, in 
different years they may appear simultaneously. 

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


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

aecidiospore always precedes the uredospoie in point of time. 

In Phragmidium subcorticium, for instance, the aecidiospores were not 
recognised at first as distinct from the uredospores, but although the indi- 
vidual spores resemble each other closely, the fact that one is produced in 
chains and the other not, distinguishes them. When aecidiospores are pro- 
duced without any special envelope or pseudo-peridium, as it is called, it 
is distinguished from the Accidium proper as a Caeoma, and there is every 
gradation from naked to covered aecidia. In Chrysomyxa the uredo is re- 
garded as a caeoma-form by Raciborski but in PJtragmidium the aecidio- 
spores are protected by a dense layer of paraphyses, which surround them, 
and thus take the place of a peridium. Triphragmium ulmariae (Schum.) 
Link, has what are called primary and secondary uredospores, although 
the former are described by De Toni as aecidiospores, but they are not pro- 
duced in chains. Winter regards them as biological representatives of the 
aecidium, and Drs. Milesi and Traverso 1 speak of them as epiteospores, to 
distinguish them from caeomospores, which are arranged in chains. There 

Origin of Spore-forms. 


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

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

Spermogonia and Spermatia. 

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

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

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

It is generally stated that the spermogonia either precede or accompany 
the aecidia, but they may occur with all the spore forms, according to the one 
which is first produced. Aecidia usually follow the germination of the 
sporidiola, and therefore spermogonia accompany them most frequently : but 
if the first formed spore is the uredo, as in Tripliragmium ulmariae, Uromy- 
cladium maritimum, and Puccinia obtegens (Lk.) Tub, then they accompany 
it, and if a teleutospore as in P. liliacearum, Duby. or Uromycladium tep- 
perianum, the spermogonia are associated with it. But 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 ifo follow." There are even instances where the spermogonium 
has entirely disappeared, as in Puccinia malvacearum, Mont. It is ac- 
knowledged by those who have given special attention to the subject that 
the spermogonium is an isolated organ, of uncertain origin and function, 
and that the balance of evidence is against its being a sexual organ. The 
spermogonium is seldom absent from the life-cycle, and vet it takes no direct 


Origin of Spore-forms. 

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

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

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

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

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

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

Origin of Spore-forms 


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

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

Variability of Teleutospores. 

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

40 Relation of Rusts to other Fungi. 


Rusts in their Relation to Other Fungi. 

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

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

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


f Ascomycetes 

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

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

I [Holobasidii (Mushrooms, &c.) 

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

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

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

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

Relation of Rusts to other Fungi. 


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


the same plant, or on different 

These various stag 


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

can germinate 

if food suppl 


ail able 

others which 

rest if need be. The change of host is 

dent advant 


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


dies down to another that is perennial. It mav 

perennial itself in 


dergiround parts of some 

gn become 
d then it 

vegetates, only producing 
be formed under normal ( 

arely the reproductive bodies which would 

42 Indigenous and Introduced Species. 

Indigenous and Introduced Species. 

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

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

indigenous. The great 

ded are, of course 

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

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

Puccinia anthoxanthi Fckl. on Anthoxanithum odoratum (1896). 

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

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

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

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

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

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

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

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

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

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

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

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

P. maydis Bereng. on Maize (1880). 

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

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

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

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

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

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

tratum (1892). 

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

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

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

The year in brackets indicates when first recorded for or observed in Australia 1 ? 

Indigenous and Introduced Species. 43 

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

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

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

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

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

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

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

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

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

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

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

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

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

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


44 Indigenous and. Introduced Species. 

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

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

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

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

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

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

Indigenous Species with their Hosts. 45 


Indigenous Species with their Hosts. 

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

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

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

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

4 6 

Indigenous Species with their Hosts. 















O eg 



a) C a. 

*0 © ce 

3 5'^ 










= := - 

= •§•2 

P >/E 

• • ; 


o * 

d > 

-a o 



3 5 S 

















5 xx *•- °*^ 








o ft© — 

d WT3 71 





.2 ©.: 




- — • 










• P^ 















- — - 












£' © 

d O ^ wji 

~^ — ©.d .i; ©.3 

- — ^ "3 ^ ft © ft*3 d 



— . ^— © 
ofi eC t- 
© © © 



to fc 

r2 5P © O 
O ft 











m ft a 


ft CO 

o o 

tt be 




s B £ 3 g^ ft 


P»i3 a X 5 

Oflp a ft od 

^ftO, anOiClHpHft^^ft^^ 

c3 $8 






c5 ce 



























■ — 




Indigenous Species with their Hosts 


co cd i 

a "So 

JS O co 

3 3 ,d £> 

d © o d 

• • • 

































co d 
d >> 






























CD Oi 




.2 CD 

is ca 

u cd 







CD ©.3 

2 rt 2 
2 d co 

P s ° 

^ 03 ^ 

p. bfi cd 

0^3 ft 

5 o o 

• • • 

ft ft ft 

d 2 

08 .2 

jo ,d 


on d 


X! ft 



• I— 


























_Z v- 





03 O 


• • 











ft ft 



■ — 



























CD ' 
























































































A s 

Indigenous Species with their Hosts. 

































£ a* 









S rr 


r J 




1 5 


& © 













o 2 




fafa fa 

















8 ! 







• — ^ 













c3 c3 

s * 

D © 










— . en 
S £ 

u o 

© s 





























>> '-= 3 

































3 e 

















Indigenous Species with their Hosts. 


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

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

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

therefore, it is regarded as in- 



Australian Distribution. 


Australian Distribution. 

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

geographical districts, since they are as yet too imperfectly known 



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

iound in some of the others. 

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

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

1905. 1892. 

Uromvces ... 27 ... 13 

Uromycladium ... 7 

Puccinia ... 90 ... 24 

Phragmidium •.. 4 ... 4 

Cronartium ... 1 ... 1 


• • 



Caeoma ... 2 ... o 

Aecidium ... 15 ... 16 

Uredo ... 13 ... 10 

161 72 

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

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

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

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

Australian Distribution. 



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


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

such as 

Puccinia tepperi and Uromycladium tepperianum. 


No. of 


Now South 








t • • 
















Phragmidium .. 
Cronartium . . . 




» • • 



m • • 

• • ■ 

• • • 



Melampsora . . . 




• • • 

• • • 



• • • 


• • • 


• • • 


" *5 




# • 









• • • 










5 2 



The Origin and Specialisation of Parasitism. 

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

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

eparatory stage to passing over to the living substance. 

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

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

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 positivelv chemotactic to the 
fungus into the tissues of the host. By similar means a parasitic fungus 
ran be induced to become parasitic on a new host." Parasitism, then, is 
due to chemotaxis, which is a form of sensitiveness in the plant whereby it 
has an affinity for certain substances, and is opposed to others. Thus there 
are various substances which are capable of attracting or repelling the germ- 
tubes of fungi, and the name of positive or negative chemotaxis has been 
given to this property. 

In an extensive series of experiments conducted with both parasitic and 
saprophytic fungi, 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 aation on the germ-tubes of obligate parasites is very 
slight. Experiments showed that " it was not sufficiently powerful in any 

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

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 



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

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

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


be separated bv mor 

phological characters. Eriksson 1 also proved in 1894 that the well- 
known and much-investigated species of Puccinia graminis could be split 
up into a series of forms, all of which agreed in producing aecidia on the 
barberry, but differed in the uredo and teleutospore generations, only being 
able to infect special host-plants. Puccinia dispersa, Eriks., was proved 

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

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

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

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

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, 
xt may be referred to the " internal developmental tendencies ot 
Klebahn x until a better explanation is forthcoming. 



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 Erysipke, 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. 




Heteroecism and its Origin. 

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

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

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

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

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

It has thus been assumed that heteroecious species originated from 

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

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

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

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




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

Next a 

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


| • • • 

dSSSV thJT^idSi, VhYle the mycelium of the teleutospore proceeded 
from the aecidiospore. The advance to heteroecism took place when the 
aecidlospores produced their mycelium in one host-plant and the teleuto- 
spores, through the sporidiola, in another, and the kernel of the matter lies 
in the answer to the question, How did this come about ? It may either 
have taken place bv a long series of slow and gradual changes, whereby 
the different spore-fonms gradually accustomed themselves to the new mode 
of life or it may have developed suddenly by one of the spore-forms 
terminating and growing on a different host-plant, and continuing to do 
*>. But this latter view is hardly borne out by some experiments conducted 
by Miss Gibson 1 , in which the aecidia from different host-plants were used 
to infect Ranunculus ficaria, and while, the germ-tube as a rule entered 
the stoma freelv, it was generally dead and shrivelled by the third day. 
This result was not supposed to be due to starvation, for she says 
" Whether the incapacitv to penetrate the cells is due to lack of attractive 
substance or to the presence of anything actively repellent is not clear 
though, as before stated, certain facts seem to suggest the presence of 
something harmful to the hyphae." 

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

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

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 : — 

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

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

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




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. staddo. 

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

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

Acacia notabilis, &c. 

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

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

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

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

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

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

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

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

c 2 



This course of development may be represented graphically as follows, 

taking P. graminis as the type:- 

a, in Europe. 


by in Australia. 







Fig. 14. 

Fio. 15. 

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

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

depend upon two host 

for their continued existence, but wh 


accomplished by means of the teleutospores, other and addit 

measures mav be taken to secure the same 

Sometimes the uredo 

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

the aecidium-mycelium becomes perennial, and reprod 





hile the teleutospore is also produced 

tage is dropped, as in the case of Puccinia 


When the aecidial 
Australia, then there 

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

dial host is absent, the same 

hen occurring 


In oth 

where the 

thing has been observed, as in Caleosf 

is absent 

f fir trees, or Chrysomyxa rhododendri when the silver fi 

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

that the 

given by Puccinia graminis 



sometimes said 

dios pores from barbe 

are much more virulent th 


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



There is a wonderful difference in the amount 

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

from the barberry 


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

ations. . . . 

wheat plant as to prevent it producing mon 

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

few i 
in mind th 

the world probably does P. graminis cause as gre 



hich the aecidial stage has never been found 

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


further obser 

to establish the fact, but Klebah 

vegetative periods and other peculiarities of th 7 

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

on a number of species are necessary 

that the utilization of the 

rendered pos- 




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

established fact that the promycelial spores 

produce no effect on the grasses 

matter how freely applied 

theory as t 
were at fii 
other hosts 

hich bear them, and the most probabl 


assumes that the uredo and teleuto spore 

th the 

dia. but 

imately passed over to 






Wherever epidemic diseases caused by parasitic fungi occur, the ques 

as to the relative susceptibility of different varieties or indi 

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


of causing di 

the host 

may either be favorably 




ards its de\ 

This predisposition of the host 

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

ous factors. It is generally 

dered that a sickly plant is more liabl 

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

more easilv and more virulently attacked 

for artificial infect 

selected. Ward 7 has shown that when the ho 

plant is starved by withholding' certain m 

salts, and thus stunted in 

growth and generally enfeebled, it is not affected in its 

A. starved plant certainly develops smaller pu 


f food 

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

sceptibility or 
les and fewer 
lable for the 

il plants. 


j far as brom< 
holds good fo 




to doubt that 

predisposition and immunity on the part 

of the host, and impotence and virulence 


of the parasite are 



of mere nutrition. But, as we shall see afterwards 

certain substances 

of the nature of food 

introduced into th 

plant may affect its liability to d 

There are various factors, ho 

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

it immune, 


The age of 

of these may be given here 

attacked has 

important infl 


particularly where the sporidiola are concerned. Young leaves and shoots 



epidermis d 

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

and it is well known that this laver becomes 


d less easilv penetrable as it 






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

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

times it is the 

both sides of the leaf 

are apt 

sometimes leaf and stem, a 
The different species of 


choose diffe 

may be on one or 
on the same host- 

portions. Thus Puccinia triticina, from 

attack, is found most commonly on the 

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

is worse on the upper leaves, 

Different varieties or sorts 
their susceptibilitv, and 

and extends 


d often parficularlv bad on sheath and 

f the same species vary 

be show 



it is 



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

being attempted 

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



d anoth 

same is the case with other 

) badly 

Hennings 5 has recentlv made observ 





hich tend to show that 

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

Predisposition . 




In 1894 several rhizomes of Peltandra virginica were sent to him from 
North America, and planted in pots. An aecidium developed on the sterns 
and midribs of the leaves, which was found to be new and named Aecid- 
ium import atum. One plant was left in the pot, which was placed in 
water, and the others were planted out in a soil composed of damp humus. 
The pot plant has annually produced the aecidia up till the time of writing 
(1902), while the other plants only showed the fungus very slightlv in 
1895 and 1896, after which they grew exceedingly strong, and since then 
have remained perfectly sound. It must be remembered that this is a 
hardy marsh-loving plant, and there is probably more than mere nutrition 
concerned in its freedom from disease, since it would be more reasonable to 
regard the result mentioned as due to the change of situation than to change 
of soil. 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 
]eaf 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 


) > 



typical characters associated \ 
would appear therefore to fm 

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

d they 
It. In 

a rust-resistant 


the leaves have a tough cuticle, 


glazed often with a gl 

not broad, erect not droopin 


id f 


d the straw 

The flag is narrow 
not soft and flabby. 


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

mav overrule them and weaken the power of 

It will thu 

be seen 


of causes may contribute 





mune, and that not only the chemical 

d other properties of protoplasm 


play, but the 

the mutual reaction of the 



plasm of host and parasite to be considered. 

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


We express this by sa> 
' proof " 


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


to disease, by repeated 


so that they become 

dually accus- 

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

be submitted to " protect i\ 

inoculation " ? 

Inoculation with 

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

different cases 

ncially-induced immunity 

__ . their 

lasts a certain time, vary 



Ehrlich and Huebener 



shown, from numerous 

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

forces at work 


the one hand, the protoplasm of the fung 


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


partially immune" by inoculation with the parasitic fun 

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

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

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


which certain nutritive and oth 

bacteria and other organisms 













. . . This 

as -p sitiv e 

value which are positively chemotactic, may be mentioned asparagin 

peptone, while sugar, which is 

of the best food stuffs and richest 

of energy, has but little attractive pow 

The power of 


the contrary 

pel bacter 

taxis, and free acids and alkal 

as well as alcohol, have this' effect 


far as kno 


Massee 9 has stated 

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

the parasite in the plant not attacked 

d if 



be impregnated with some substance which is negatively chemotactic, and 

at the same time does not affect the 

lty of the 

gainst parasitic fungi may be obtained in this way 

then immunity 

Laurent 1 , acting on this principl 

ducted a series of experiments 


the possibility of producing potatoes which would be proof against 


He grew very 

ptible varieties in pots, to 

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


know as negative chemo- • 






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

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

tection against rusts, but 

and the probability is that he did 

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

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 
coppar not sensibly greater than that found in the fruits obtained from un- 
treated plants. 

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




Wheat Rust in Australia. 


The Present Position of the Rust in Wheat Question in Au 

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

ated. viz. , oats 

d barlev 

And the 

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

does the most d 

be chosen 

Puccinia grami 

distinct in its characters that there is no difficulty in recognisin 

hich is so 





heat in Australia, the positively injurious 

Puccinia era 

d the comparatively harmless P. triticina, beca 



the early d*ays whe 

it is 

grown to 


does not pinch and shrivel the grain like the other. 

and other cereals had to be imported into Sydney, 

such an extent in the Australian Commonwealth that in the 

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

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

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

the losses from d 

have a corresponding far-reaching effect 


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


vear passes without its being present 

heat to a greater 

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

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

idea of the extent of the 

bad season like that of 

be stated that in a part 

the loss was estimated to be for the whole of 





such favorable y 

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


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

weather was what was known as " muggy," 

lg of show 

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

oming into fl 

rticularly susceptible, and the rust spores found 

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

: conditions are often regarded as the prime factors 


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


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


ferences extended from the first in 1890 to the last 

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

But there 


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


Effects of Rust on the Straw and Grai 


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

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

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

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

well formed, with plump 
with light and shrivelled 1 


grain, and the other has poor ears 

Wheat Rust in Australia. 





But the effect upon the com 

of the 

d particularly its 


is not known 

the farmer, who cuts his cron for h 

: threatens to ruin it. 

This has recently been determined by F. T. Sh 

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





analysis is as follow 

ithat the results are strictly comparable 


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



u- X 




iff c5 
















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


• • » 


• 9 • 




1 -4944 







1 -97 

2 *35 










The Sir a 

pointed out that in crude protein the rusted 

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

that go 

form blood and build up muscle, it may safely be 

concluded that the rusted 

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


is also in the rusted straw slightly 

that all this affords additional evidence of its more highly 


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

point of view has received 

The experience of one who has done 

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


d made the men about the thresher rub their sk 



egards feeding rusty hay, another with larg 


nforms me that horses and cattle relish it far before ord 


course, it 

fed as chaff. 

— This from the rusted wheat is onlv about 



The Grain — 

srht of that from the rust- free wheat, but 

half the 

it has 

protein content show 

ght for weight, a considerably higher nutritive value. 


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

portion of b 

but chiefly in the partial and incomplete transference and 

of starch 

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

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

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


Wheat Rust in Australia. 

beeins to form 

d if that 

favorable for the develop 


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

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

consequently the 
investigation als( 

On account of the comparatively 

appearance of P. graminis, it js 

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




be the 

Puccinia Graminis and the Barberry in Australia. 

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

But in Australia infection of the barbe 

has not been successful 


several attempts have been made, and 

have doubted whether we have got the 

P. graminis 

d not a 

distinct biological form of 

From a comparison with European speci- 


examination of the 

the uredo and teleuto spores 

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

proved. The relation of 

therefore, to the barberry in Austral 

became a pressing subject for experiment. 

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

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

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





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

Wheat Rust in Australia 



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

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

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

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

Cuttings from the English barberries have also been planted at Port 
Fairy in a rusty spot where they have thriven, and Queen's Jubilee 
wheat planted around them produced abundance of P. graminis. The 
rusted straw was allowed to die down on the spot, and every 
facility given for the inoculation of the barberry, but without 
result so far. Some P. graminis on wheat was sent by Dr. 
Plowright in March, 1903, which was gathered In September, 
1902, and kept in his garden till March. On arrival here 
some of it was kept inside, and a portion placed outside exposed to the 
weather, just as was done with Australian rusted straw. It was thus 
exposed during our winter months of June, July, and August, and in 
September it was tested, being then exactly twelve months old. While the 
teleutospores about six months old from the Australian wheat germinated 
freely, there was no change in the spores from the English wheat, and 
although attempts were made at different times, there was never any 
sign of germination. Probably they were kept too long, as they might 
have germinated on or about the English spring. Prior to this, I had sent 
rusty Australian straw for trial to Dr. Plowright, but none of the teleuto- 
spores showed the slightest trace of germination. Writing in the Gardeners 1 
Chronicle for 15th 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 

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

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

1 has apparently no intermediate, stage. 


Wheat Rust in Australia. 

The Aust 

rust has been determined as Puccinia graminis by 

Eriksson and other authorities; but if the ab 

produce the aecidium 

on the barberr 

is taken as 

the final criterion, then 


must be 

pended in numerous other cases. 

Thus Massee 4 has found P 

Kew ; 


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

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

but he significantly adds 

Notwithstanding the great quantity of 

and Mahonia present in the grounds 

most careful and con- 

h has failed 

the presence of the aecidiospore stag 

(Note 7, p. 75 

The Problem St 

In dealing with the rust question from a practical point of 


were two main 

issues to be determined 



How is the rust spread and continued f 



may its injurious effects be 


or counteracted or 


The first question is a most 

portant one, for if we could find out 

where, and under 

time from reapim 

ditions, the rust is 









destroy it at this stage, and prevent its reappearance 

might be able to 
Although the 

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

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




ledge of the life-history of 





d how far the conditions can be controlled which 

mt itself, ar 

susceptible to the fungus. 

the Rust is Spread and Continued / 

actly now, thanks to the labou 


to Y 

of Eriksson 



Ward, and 

how the, 

germ-tiibes, how the) 

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

of thei 

until thev 


form a spore 

and reproduce the 

pores on the surfac 




We thus know how fresh 

s pores 


once thev have 


a start: but it is the starting-point 


Although the rust was know 

d the effects produced by it were 

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



attacks of small insects 

i t 

Horse-hoeing Husbandry, attributes 




some think, by the East wind, which 

the wheat, leaving their excreta as black spots upon the 

shown by the microscope 

767, its true nature as a fungus, and 

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


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

The rust then is a fungus growing 

de the wheat-pl 



g at 

its expense 




or spores, which 1 
at certain seasons. 

itself by means of minute seed-like bod 

conspicuous on the leaves and stem of the wheat 

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

farmers that the barberrv bush had 

something to do with its spread 


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

compelling the 
Barv, in 1864, 


to extirpate barberry bushes. And when De 

fied the farmer 


d scientifically th 


was a connexion bet 


d th 

the fungus which appears on the barberrv 

hich appears on the wheat, then it was 

thought by 

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


the barberry bush in order to get rid of the 


it is 


Wheat Rust in Australia. 

6 9 

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

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

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

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

untreated seed. 

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



Wheat Rust in Australi 

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

uredospores is produced. 

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



> vc have elsewhere shown, may also be attacked by Pucctma gratnims 
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. 

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

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

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

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


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

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

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

Wheat Rust in Australia. 




Prevention or Mitigation of the Rust. 

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

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

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

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

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

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

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

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




Wheat Rust in Austr 

the rust. At Port F 

in Victoria, where rotation is regularly prac 



here I have had crops 


the same season, sometimes 


fter mangels, or potatoes, or 
rustiness from 

those crops 

d-point of good farming, it 

onions, there was no 

ptible difference 


heat grown in succession. 

From the 

be recommended ; but cropp 

fter vear does not seem to affect the liab 

to rust further th 

that self 


heat is likely to appear in the interval bet 




heat, if not killed in 


ng year 
in this, 
the two 
is much 

d on account of its earliness may escape 

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



more forward than sown grain, ai 

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

times more rusty than the ordinary wheat 

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

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

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


Professor Lowrie, speaking from a large experience 

South Au 

ders that the apparent exempt 

hich fallow-land sometimes 

land fii 

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

Burning Stubble 

It is sometimes recommended to b 

the stubbl 

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


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


Theoretically, the burning of the surface of the 

should destroy 

umber of 

and to that 

beneficial ; but, practically, the 

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

are con- 


possible increase of rust. 

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

d rust-free, like th 

wheat itself ; but on closer 

is only 

of several factors th 

gat ion, 

it will be found that the soi 

be taken 


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

sometimes been raised from the, poorest soils. 


the rich soils the crop 

naturally heavier and more luxuriant than on poor 

d the softer 

and more 



aded by the parasite 


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




said to be particul 

free : but in some seasons th 

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

destroyed by rust. The reason for g 

freedom from rust lies not 

much in the 


the light rainfall and the early ripening of the 


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

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

At Port Fairy 

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

soils, and this may be 
absorb more heat, and 

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

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

and thus encourages the rust. 

Perhaps no means for the prevention of rust h 

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



of different 

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

by a late D 

been discovered that 

of Lincoln Colleg 



No manure has 



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

it. Nitrogenous manures 

hile phosphatic ones had a tendency to diminish 

Wheat Rust in Australia. 


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

early maturity, and 1 thus enables the crop to escape the rust to a certain 

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

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

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

It was adopted as a conclusion at the last Rust-in-Wheat Conference 
that the treatment of the seed is valueless for rustt, and Dr. 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 ibeen found successful in treating peach and plum rust, 
still the mechanical difficulties to be overcome in spraying a wheat-fielH 
are so great, that it is no longer regarded as of practical importance. 

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

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

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


Wheat Rust in Australia. 



world (Europe 




there is no wheat k 









hich will be proof 


when grown under conditions favorable to its^ development 

experiments have show 
particular d 



the rust 
But the same 

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

constitution— it may be by their vigour of growth or 


their cuticle or glaucousness 



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

less of 
the inroads of the 

d such wheats 

least one such 

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


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


1 succumb to the rust, 
this great and burning 

question the only measures I can 


i. To produ 

heats suited 

Australian conditions by crossing 

Wheat Experimentalist of N 

buth Wal 

is now so 

as Mr. Farrer, 
successfully doing. 

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


d milling qu 

keeping up the 


g thei 


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, and to increase and 
extend this knowledge it would be desirable to follow the advice of 
Professor Eriksson, who has done more than any other single individual 
to enlighten us on the subject : — " The question of the rust of cereals being 
of the greatest practical importance for every country, means should be 
furnished to those in whose hands the direction of these investigations are 

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

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

appears, but it is not very common." 

Montague Smith, in 1828, notes 

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

P. 4 

Eriksson 18 

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

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


Wheat Rust in Australia 


Note 2. 

P. 14. — 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 fuse. 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 hyph'ae 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 used! in New Zealand 
for hedges, and, up to the present, I have never seen any sign of aecidia 
of Puccinia graminis on them, although I have examined hundreds. It 
would appear as if Puccinia graminis in Australia has lost the power of 
forming aecidia on barberries.' ' 

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

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






















Classification, with Special Reference to Biologic Forms. 


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

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

Gradations of Specific Variation. 

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

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

follows: — 1, secalis; 2, avenae; 

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



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

i. The uredo 
2. This form 


on the following grounds 

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



growing alongside of each other 

wheat, even when the host-plants are 

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

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

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 



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


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

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

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

It will be observed 

that onlv biological characters 

are here taken into 

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

striking character than 

and overlooked. 

The three t 


where they were hitherto unsuspected 

pes 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 

biologic fo 

d the aecid 

agreeing in the faot of producin 

dia on the bar 


feet the 

at first divided into biol 

tuted, is the type of 

distinct species, both on morphol 

forms, which were afterwards found to h 

and biol 

grounds. And the 

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

are strict lv 

ted in their infective 




d their 

n title to distinction as species is this biological p 

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

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

in the fut 

nclude under 

distinct morphological characters, but 

logical lv simil 

are yet 

fined to definite host 

It will thus be 

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

opinion as to the amount of 











biologic forms. 

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

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

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

Biologic forms 

in fact, may be regarded as incipient morphological 

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

species, the physiological differences at present existing becoming ultimately 

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


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 


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 direation of the evolution." 

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

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


Systematic Arrangement. 

Systematic Arrangement and Technical Descriptions. 

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

mark out for us that particular object from all others. 

> > 

[The descriptions 

are necessarily technical, and give those characters which enable the species 
to be discriminated from others, with the help of the illustrations. When 
the 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 Pfianzenfamilien: — 


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

a filiform, branching, septate mycelium 



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

of more than one kind 

ium. The order 



a short promy 


the third is not represented in Australia 

be grouped in the following four families, of which 




Fam i. P 

in groups 

rows, or several cells in a 

Teleutospores sessile, in columnar or filiform m 





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



Teleutospores sessile, in flattened one-lavered masses or loose 




of their host. Fam. 4. Melampsoraceae 

account of their economic 

alwavs treated fi 

portance, the Graminaceous rusts are 

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



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

be treated at greater length than any of the others 


the different genera 

(the different forms 

d it 


hich the teleutospore assumes in 

There are a number of genera based upon very 

and it is not always easy to settle whether they 

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

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


should be retained, where the teleutospo., 

only the septum is longitudinal instead of 

celled, as in Puccinia, 

There are several 

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

longitudinal sept 
those species 

as well 


but th 

hich the spore is distinctly divided longitudinally 

retained for 

germ-pore at the apex of 

Systematic Arrangement. 


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

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


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 sitalk. Uredospores echinulate or warty all 
over. Fig 16. 

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

but uredospore smooth on one face. 

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

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

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

and with only one germ-pore in each cell. 

Fig. 18. 

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


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. 16. 

Fig 17. 

Fig. 18 

Fro. 1 P. 

Fig 20 

8. Gymno sporangium, Hed 




ely three 

d), and walls which ultimately form a common gel 

mass, with usually several germ-pores in each 


9. Hapaloph 


Teleutoso^re three celled, and 


of two basal cells 



of each othe 

th a third on top 



Link. — Teleutospore three-celled, and consisting of 

Fig. 23. 

basal cell supporting two others alongside of each other 



! I. 




Phragmopyxis, Diet. 

Teleutospore three-celled in a longitudinal 


with outer laver swelling when moistened. Fig. 24. 

Phragmidium, Link. 

Teleutospore three or more celled in a longi- 

tudinal series, 
Fig. 25. 

and outer layer not swelling when moistened. 

Fi<;. 21. 

Fig. 22. 




Fig. *24. 

Fig. 25. 

Sphaerophragmium, M~ 

Teleutospore consisting of a spherical 

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


and transverse fission. Fig 

14. Authority 


— Teleuitospores forming a head of th 

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

Several sm 


Fig. 27. 

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

manv-celled head 

erselv and 

dinally septate 

arising from a stalk compounded of several hyphae 

d with 





Fig. 26. 

Fig. 27. 

Fig. 28. 

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


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


gle germ-pore in the former serves ito distinguish 

spores are 


echinulate or 

ted. and wh 


sides, the uredo- 
:eleutospores are 

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

greater thickening of the wall, and the fact that the 




All the different spore-forms mav 

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



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

alone. In 

forms occur, but 

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

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




Uromvces — Gramineae. 


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

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

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

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

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

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

Australian species, 27 


l. Uromyces danthoniae McAlp. 


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

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

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

or 16 x 12 


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

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

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

long covered by epidermis. 

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

slightly tinted adjoining spore, up to 96 fx long. 

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

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

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

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

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

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


Uromyces — Gramineae. 

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



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

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

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

Darluca filum Cast., is common on uredosori. 

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


. Uromyces ehrhartae McAlp. 


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

Uredo ehrhartae McAlp. 

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

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

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

24 x 


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



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

brown to black. 


pale chocolate b 

to chest 

brown, only 

9 /*), 


edospores intermixed, hooded and thickened at apex 

ther conical or truncate, 

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



stout, subglobose to 




26 x 1 



On Ehrharta stipoides Labill 

Microlaena stipoides R. Br 


Near Melbourne, Killara, My 




Uredospores common but teleutospores rather 

. e 'hbourhood of 

son are sometimes rathe 
and intermixed with ur 

Melbourne during December 


gh in 


edosori on those 

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

still partially green. 

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

Darluca filum Cast., is very common on uredosori. 

Only the 



(Plate XVI, Fig. 132.) 


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 11, average 32 x 22 /w. 

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 fx ; pedicel sometimes persistent 
tinted, commonly 20-35 ll long, but often much shorter. 

On leaves and stems of Sporobolus indicus R. Br. 

Victoria — Near Melbourne, Aug. — March. 

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

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

(Plate XVI., Fig. 133.) 



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

On living leaves of Bulbine bulbosa Haw. 


Victoria — Omeo (Morrison). 

New South Wales — Upper Macquarie II 

5. Uromyces tricorynes McAlp. 


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

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

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

Aecidiospores bright orange, subglobose to somewhat polygonal, 

smooth, average 20 x 1 7 li 




Uromvces — Orchidaceae. 

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

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

by ruptured epidermis. 

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

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

III. Teleutosori blackish on ruddy brown spots, 



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

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

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

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

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

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

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

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

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

Darluca filnm Cast., occurred on the uredosori. 

(PlateXVI, Figs. 134, 13 5.) 



6. Uromyces microtidis Cooke. 


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

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

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

II. Uredospores globose, subglob 

ate to warty, yellowish-bro 

III. Teleutosp 

termixed with 

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

-30 x 1 



psoid, minutely acule- 
ovate, with hyaline 

On living leaves of Microtis porrifolia R. B 

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

New South W 

Bullahdelah, II., Ill 

Uromyces — Orchidaceae 

8 9 

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

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

termixed with the teleutosp 

Sydow 2 has also found the uredospores 

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

Chatham Island, New Zealand. 

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

sts found on the Orchid 
is found that aecidia occi 



a occur on ChiloglotttSj 

The teleutospores are 

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

gh and knobby, and 

size of the spore not generallv exceeding 35 

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

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


Uromyces microtidis 

U. orchidearum 

• • 

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

U, thelymitrae 

1 - * 

35-45 x 25-30 /x 


Generally prominently 
warted all over 

Irregularly warted and 
warts not so pro- 

Less closely but more 
prominently warted, 
hence polygonal 



3-4 fi 

5-6 ju 


co iioid 



(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, Vict. Nat. XL, p. 90 (1894). 
Sacc. Syll. VII., p. 580 (1888). 

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

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

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

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 /x. 

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

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

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


warted epispore ; 

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

downwards, up to 77 ^ long. 

D 2 

£ Uromyces — Orchidaceac, Scrophalariaceae. 

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

New South Wales — Mt. Victoria (Hamilton 




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

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

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


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

labelled in Cool 


Sometimes there are abnormal elongated teleutospores up to 60 yu long 

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

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


8. Uromyces thelymitrae McAlp. 

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

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

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

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


9. Uromyces limosellae Ludw. 

Limo sella. 

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

I. Aecidia amphigenous, scattered or gregarious ; pseudoperidia with 

whitish laciniate margin, not deeply incised. 

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

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

by epidermis, pulvinate, dark brown. 

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

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

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

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

18-21 p. 

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

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

Uromyces — Goodeniaceae, Rubiaceaz. 91 

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

On leaves of Limosella aquatica L. 

S. Australia — Kangaroo Island (Tepper). 

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

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

Specimen not seen. 


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

Selliera, Scaevola 

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

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

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

III. Teleutosori bullate, intermixed with aecidia or surrounding them 

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

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

On leaves and flower stalks of Selliera radicans Cav. 

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

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


ham, Nov., 1905 (Robinson). 

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

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

On Scaevola sp. 

Queensland — St. George (Wedd) (Bailey 


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

(Plate XVIL, Fig. 140.) 


11. Uromyces asperulae McAlp. 

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

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

A sperula. 

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

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

9 2 

Uromyces — Leguminosae. 

III. Teleutosori on stems 

ds their base, elongated oval, dark 

brown, appearing almost black, confluent, erumpent 


first intermixed with 


brown to chestnut-b 



'es, yellowish- 
>r elliptical to 

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

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

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

The host 

is given 

Asperula scoparia Hook, f., in the Index 

Keivcnsis, but the synonymy seems 

confused. A. oligantha F.v.M 

is first gi\ 

as Rub 





synonym of Asperula scoparia Hook, f 

ticola is given as a 

(Plate XVII., Fig. 141.) 


12. Uromyces appendiculatus (Pers.) Link. 

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

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


Aecidia densely crowded in small 

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

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

14-23 /x.] 

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

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

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

III. Teleutosori dark-brown, almost black, easily detached, otherwise 

like uredosori. 

Teleutospores dark brown to chestnut brown, subglobose to ellip- 
soid or oblong, smooth, scarcely thickened at apex, with broad pale 
flattened papilla, 27-37 x 19-25 /u ; pedicel hyaline, persistent 
elongated, up to 70 p long. 

On Cowpea (Vigna catjang Walp.). 

New South Wales. 

Richmond, May, 1905 (Musson) 

The distinction between the teleutospores of U. fabae and U. appendicu- 
latus is very marked. In the one the apex is thickened up to 9 /*, while in 
the other the thickening is not perceptible. 


(Plate XLIL, Fig. 306.) 



Uromyces — Leguminosae 



13. Uromyces bicinctus McAlp. 

II., III. Sori ruddy-brown, gregarious often confluent, raised, compact, 

rupturing epidermis irregularly. 

II. Uredospores yellowish-brown, clavate to oval or ellipsoid, rounded 

at apex and slightly thickened, densely warted and warts arranged 
in regular longitudinal lines, with distinct germ-pores, generally 
arranged in two bands about one-third length of spore from either 
end, and usually three to four in each band on one face, 30-40 x 
13-18 /j,, occasionally reaching a length of 45 ^t, and variable in 
breadth, average 34 x 15 fi ; pedicel similarly coloured to spore or 

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

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

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

Queensland — Rockhampton, 1867 (from host-plant in National 

Herbarium, Melbourne). 

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

McAlp., in the uredospores, but they are shorter and much narrower, and 

distinguished at once by the two bands of germ-pores, on account of which 

the specific name is given. The teleutospores are also much thickened at 

apex and destitute of processes as a rule, although occasionally two may 

(Plate XLIIL, Fig. 318.) 


14. Uromyces fabae (Pers.) De Bary. 

De Bary, Ann. Sci. Nat. Ser. 4, XX. (1863). 
Sacc. Syll. VIL, p. 531 (1888). 

[O. Spermogonia yellowish, in small groups on leaves and stem.] 

[I. Aecidia scattered, in rings or orbicular patches ; pseudoperidia 

short, slightly prominent, fiat, with torn white edges. 

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

II. Uredosori amphigenous, roundish, pale brown, powdered, scattered, 

often confluent, soon naked. 

Uredospores subglobose or ovate, ochraceous, echinulate, with 
three equatorial germ-pores on one face, 20-30 x 17-20 fx. 

[III. Teleutosori rounded on the leaves, more abundant and elongated 

on the stems, often confluent, blackish-brown. 

Teleutospores variable in form, obovate or broadly clavate, 
smooth, dark-brown, apex darker, thickened (8-10 ^u), and rounded, 
truncated or conical, sometimes with colorless papilla, 24-47 x 
17-30 /i; pedicels hyaline or pale brown towards spore, persistent, 
up to 110 fi long.] 

On stems, leaves, and pods of Broad Bean (Vicia faba L.). 

New South Wales.— 1894 (Cobb 9 ). 
Queensland.— Gladfield (Gwyther) (Bailey 19 ). 


Vromyces — L eguminosae . 

This rust has not been found in Victoria, and although it is said to be- 
very common in New South Wales, on applying to Dr. Cobb for specimens, 
he informed me that they had all been destroyed by insects. Only the 
uredo-stage was found in New South Wales, and the exact species is still 


(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 u, average 62 X 23//. 

pedicels deciduous, hyaline, elongated, up to 96 fi long, with 
septum at a short distance beneath spore. 

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

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

On Acacia sp. 

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

Portion of the original material named in Cooke's handw 


National Herbarium, and our description is based upon that. In the original 
description by Cooke and Massee the uredospores were unfortunately mis- 
taken for teleutospores and vice versa. The uredospores are easily known 
from being obtusely warted and with equatorial germ-pores, apart altogether 
from size and shape. 

The teleutospore is solitary at the apex of the pedicel, but the sep- 
tum at a short distance from the spore foreshadows the Uromycladium with 
a colourless vesicle or cyst produced laterally immediately beneath septum. 

The nature of the teleutospore and the presence of a septum in the stalk 
was so suggestive of Uromycladium that the material was specially ex- 
amined to see if more than one spore was borne on a stalk, but the most 
careful search failed to reveal any indications that this was the case. 

III. Teleutospores intermixed with uredospores, ruddy brown, globose 

to depressed globose, epispore rather thin, scarcely 2 /x thick,, 
somewhat polygonal seen from above and then germ-pore veiy 
prominent, 25-30 \i diam. or 16-21 x 25-35 \x, average 18 x 28 fi ; 


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

16. Uromyces hardenbergiae McAlp. 


Sori on under surface of leaf, brown, crowded, globose to ellipsoid, often 

confluent, bullate, with ruptured epidermis, which usually remains 
in large patches. 

Uromyces — L eguminosae 


II. Uredospores golden-brown, elliptic to ovoid, echinulate, relatively 

thick-walled, with three distinct equatorial germ-pores on one face, 
25-35 x 20-22 n, average 27 x 20 p. 

III. Teleutospores intermixed with uredospores, very sparse, ruddy- 
brown, thick-walled, smooth, ellipsoid to obovoid, slightly 
thickened at apex, with hyaline apiculus and showing germ-pore, 
25-28 x 18-21 fi, average 26 x 19 ft; pedicel persistent, hyaline, 

ated, up to 35 

On leaves of Hardenbergia monophylla Benth. 

Victoria — Kergunyah, Dec, 1903, II., III. (Robinson). Frankston, 

January and August, 1904, II. (Robinson). 

The teleutospores were exceedingly rare, and somewhat resembled the 
uredospores, but slightly smaller, with ruddy smooth epispore and hyaline 

This species persists all the year round. 

Darluca filum Cast., very prevalent on uredosori. 

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


A cacia. 

17. Uromyces phyllodiorum (B. and Br.) McAlp. 

Berkeley and Broome, Linn. Trans. II., p. 67 (1883). 
Cooke, Handb. Austr. Fung., p. 331 (1892). 
Sacc. Syll. VII., p. 556 (1888). 

Melampsora phyllodiorum Berkeley and Broome, Linn. Trans., 

II., p. 67 (1883). 
Uromyces digitatus Winter, Rev. Myc, p. 209 (1886). 
Uromyces phyllodiae Cooke and Mass. Grev. XVIL, p. 70 


| O. Spermogonia brown at first, becoming shining black, numerous, 

I small, blister like, seated in centre of swollen, brownish, scattered, 

I discoid, amphigenous tubercles, varying in size from 1 to 4 mm. 


II., III. Sori surrounding spermogonia or alone, ruddy brown to dark- 
brown, round or elongated, crowded, often confluent, compact, 
raised, girt by the ruptured epidermis, which bursts irregularly. 

II. Uredospores golden yellow to golden brown, oval to ellipsoid, 

rounded at apex or bluntly pointed and slightly thickened, densely 
warted, and warts arranged in regular longitudinal lines, with 
distinct equatorial germ pores, 3-4 on one face, and even reaching 
to 6, 35-54 x 16-25 //, average 38 x 22 /u, with elongated 
and hyaline pedicel. 

III. Teleutospores at first intermixed with uredospores, lemon yellow, 

and sometimes almost colorless towards base, fusiform to wedge- 
shaped, thickened at apex, and bearing finger-like erect or divari- 
cate processes, sometimes a single process, or bifid, or several, 
60-70 x 14-20 /i, but may even reach a length of 86 ^ ; pedicels 
long, persistent, hyaline ; paraphyses intermixed with teleuto- 
spores or with uredospores accompanied by teleutospores, elongated, 
variously shaped, but generally somewhat cylindrical, thickened 
and rounded at apex, and tapering sometimes into a slender 
filament at base. 70-93 X 6-10 u. 

Vromyces — Legaminosae. 

On phyllodes of Acacia notabilis F.v.M 

S. Austra 

On phyllodes of 


Near Gawler, July, 1885 (Tepper) (Ludwig 2 ). 
daUachiana F.v.M., and A. penninerms Sieber 
Alps, near Bright, Dec, 1904 (C. French, jun.) 

On phyllodes of Acacia dealbata Link. 

Victoria— Orbost, Dec, 1905, II. III. 

On phyllodes of Acacia penninervis Sieber, A. microbotrya 
neriifolia A. Cunn. in National Herbarium, Melbourne. 

New South Wales— Twofold Bay. 
Queensland — Brisbane River. 

On phyllodes of Acacia pruinosa A. Cunn. 

Benth. and A, 

New South 
On Acacia sp. 

— Gosford, Jan., 1906 (Frogg 

Queensland — Brookfield, Brisbane River (Bailey 1 ' 19 ). 

New South Wales 



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 


by Winter as 32-35 x 20-25 p y 

reach a length of 54 


but there are occasional elongated forms which may 
They somewhat resemble those of Uromycladium notabile, but the markings 
on the epispore are much closer together. 

The teleutospores are very characteristic, but very variable both in shape 
and size. The apex may be prolonged into a single process, or there may be 

quite a number, at least up to 6. 

As regards size 

they are generally 

elongated, and the measurements given are the mean of a number taken from 

spores ending in a single process. They germinate at once without a period 
of rest. The specific name of digitatus is so appropriate that it is unfortu- 
nate it has to be set aside in obedience to the law of priority. 

This species was first described by Berkeley and Broome in a list of 
fungi from Brisbane, Queensland, as Melampsora phyllodiorum in 1883, and 
drawings accompany the description. Specimens of the original material 
have been kindly supplied to me by Mr. F. M. Bailey, Queensland State 
Botanist, and uredospores and teleutospores from these are shown in 
PI. XXV., Figs. 218-220. There is no doubt as to its identity. 


The following is* the original description of Berkeley and Broome 

Sori in amphigenous tubercles ; spores 

rather fusiform, 55-58 f. 

elongated, uniseptate, fusiform, even, 22 



from delicate filaments, 

granulated, mixed with others which are 

The longest uredospore I 


i long. 

found was 54 ^ long, and the uniseptate spores of Darluca filum Cast., were 
also very common. 

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

Then Cooke and Massee, in 1889, described a Uromyces phyllodiae from 
Brisbane as follows : — " Sori minute, orbicular, compact, brown, crowded on 
elliptic, bullate, brown spots, 3-5 mm. long, at length naked, not pulverulent. 
Uredospores not seen. Teleutospores elliptic, obtuse, rarely apiculate, 
brown ; epispore minutely warted, rather thick, hyaline, thickened at the 
apex, 40-45 x 16-18 ^/." A portion of the original material *ras supplie 
by Mr. Bailey, and both uredospores and teleutospores have been found. 

Uromyces — L eguminosae. 


Cooke and Massee mistook the uredospores for teleutospores, but the 
existence of several germ-pores, together with their general appearance, 
indicate their true nature. Spermogonia were present in this material. 

Finally Dr. Cobb, 10 in 1897, described the uredo stage of Melampsora 
phyllodioruniy and gives a drawing of the uredospores with an average size of 
41 x 17 fi. He also found the perithecia of Darluca filura Cast., among 
the sori, but their true nature was not recognised. 

(Plate XXV., Figs. 218-228.) 

18. Uromyces trifolii (Alb. and Schw.) Winter. 

Winter, Die Pilze, L, p. 159 (1884). 


Cooke, Handb. Austr. Fung., p. 330 (1892). 
Sacc. Syll. VII., p. 534 (1888). 

0. Spermogonia honey-coloured, disposed in small clusters. 

Spermatia hyaline, globose, about 3 jj, diam. 

1. Aecidia in circular clusters, on pallid spots ; pseudoperidia cup- 

shaped, with a white laciniate margin. 

Aecidiospores subglobose, ellipsoid or irregular, very finely 
verrucose, pale orange, 14-23 /j. diam., or 22-25 x 14-16 //. 

II. Uredosori rounded or elliptic, scattered, surrounded by the 

torn epidermis, ruddy brown. 

Uredospores irregularly globose or shortly elliptic, echinulate, 
golden-brown, with two equatorial germ-pores on one face, 22-26 
X 18-20//. 

III. Teleutosori smaller and rounded on the leaves, larger and elon- 
gated on the petioles, dark brown, almost black, bullate, long 
covered by the leaden epidermis. 

Teleutospores generally intermixed with uredospores, ellipsoid, 
globose or pear-shaped, thickened at the apex with a small pale 
wart, smooth, dark-brown, 22-30 X 19-22 ^u, average 24 X 20 \x\ 

pedicels long, hyaline, deciduous. 

On leaves, petioles and stems of Trifolium repens L. 

Victoria — Near Melbourne, Apr., I., II., III. Sep., I., III. Bunyip, 

Nov., I., II. Nyora, Apr., I., II., III. Murramurrangbong 
Ranges, Jan., I., II., III. 

New South Wales— (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* considers this a common 

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


Uromyces — Polygoriaceae. 



1<). Uromyces politus (Berk.) McAlp 

Berkeley, Linn. Journ. XIII., p. 174 (1872). 
Berkeley and Broome, Linn. Trans. II., p. 67 (1883). 
Cooke, Handb. Austr. Fung., p. 342 (1892). 
Sacc. Svll. VIL, p. 833 (1888). 

Roestelia polita Berk. 

I. Aecidia on dark- purplish, elongated patches, generally arranged in 

lines which may be parallel, bursting through epidermis, bright 
orange, becoming ivory- coloured with age, cylindrical to slightly 
compressed, straight, averaging 1| mm. high; pseudoperidia 

polished, smooth, with white, narrow, toothed margin ; peridial 

oblong to angular, with striated margin, 32-35 jj. Ion 

Aecidiospores ochraceous, variable in size and shape, generally 
ellipsoid to ovoid or oblong, finely echinulate, 24-30 x 19-22 /i. 

III. Teleutosori on the same discoloured area as the aecidia and 

associated with or distinct from them, solitary or confluent, 
brownish to blackish, bullate, elongated to oval, surrounded by the 
raised and ruptured epidermis, compact, 1-2 mm. long 

Teleutospores yellowish-brown, smooth, ellipsoid to oblong, 

rounded or pointed and apiculate at apex, thickened (6-8 
occasionally two-celled, variable in size, 28-40 x 20-25 /j, a 
32 x 24 u ; pedicel hyaline, persistent, elongated up to 150 

and at the same time Cronartium asclepiadeum was noted on Jacksonia 
scoparia from the Darling Downs, Queensland. 

Then just ten years later, in the transactions of the same society for 1882, 
p. 67, R. polita is recorded on the latter plant from Brisbane (Berkeley and 
Broome 2 ), and it was rather peculiar, to say the least of it, that the same 

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

New South Wales — Pamamero Lake, Nov., 1860 (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 the J 
Uromyces. ™ 

Berkeley 2 first recorded the aecidial stage of this species on Muehlenbeckia 
cunninghami in the Journal of the Linnean Society XIII., 174 (1872), 

species of Roestelia, or even a Roestelia at all, should occur on plants so 
widely separated in a botanical sense as Muehlenbeckia (Polygonaceae) and 
Jacksonia (Leguminosae). At the end of his description Berkeley remarks 

" The plant is identical with a specimen in the Kew Herbarium, and is 
growing on the same plant, Jacksonia scoparia R. Br. 

However, on examining the specimens on which this determi nation was 
based, it became evident how the error had arisen. Fortunately I have the 
specimen of Jacksonia scoparia from the National Herbarium, Melbourne, on 



Uromyces — Polygonaceae. 




r hich Cronartium was first determined, and Mr, Bailey has sent me a spec 
men from his herbarium labelled " Roestelia polita Berk., " and which was s 
named by Berkeley, On comparing the Brisbane specimen with the Darlin 
Downs specimen, they are seen to be the same, and Berkeley's original deter- 
mination of Cronartium is correct. When it is remembered that Roestelia 
polita Berk., has only been found on Muehlenbeckia and not on Jacksonia it 
will reconcile certain discrepancies in Berkeley's original descriptions and 
drawings. In the original description of R. polita the spores are given as 

5, while in connexion with its occurrence on Jacksonia it is stated — " the 
spores are globose about '0003 inch in diam. (7— 7J /j)" If we turn to the 
drawings on PL ] 5, the otherwise puzzling figures become clear when they 
are taken to represent a Cronartium, and it will be seen from the description 
of this fungus on Jacksonia that the so-called aecidiospores are really the 
promycelial spores of that fungus. 

If the identical specimen in the Kew Herbarium, to which Berkeley referred, 
is examined I have no doubt it will turn out to be a Cronartium. At the 
time Berkeley wrote our knowledge of the distinctions between these forms 
was not so clear as now, but still the projecting pseudoperidium of the 
so-called Roestelia, with its characteristic peridial cells and contained spores, 
is quite distinct microscopically from the outwardly similar column of 
teleutospores of Cronartium which are wedged together into a solid mass. 

In the original description the locality for New South Wales is given as 
" Bambamero " Lake, but Dr. Howitt informs me that the name is as 
above, and that the lake is situated 20 miles from Menindie, close to the 
Darling River. 

(Plate XXXIX., Figs. 297, 298; Plate XLIIL, Fig. 317.) 

20. Uromyces poiygoni (Pers.) Fckl. 

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


McAlpine, Agr. Gaz., N.S.W., VII., p. 301 (1896). 
Sacc. Syll, VII., p. 533 (1888). 

0. Spermogonia yellow to honey-coloured, conoid, in small groups. 

Spermatia hyaline, minute, 3 X 2 p. 

1. Aecidia crowded in roundish patches, bright orange, mostly hypo- 

phyllous, but often opposite ; pseudoperidia rather flat, with broad 
whitish torn edges. 

Aecidiospores subglobose, finely verrucose, pale yellow to orange 
yellow, 16-28 fx diam. 

II. Uredosori pale cinnamon to rusty brown, scattered, or arranged in 

a circinate manner and confluent, mostly amphigenous, pulveru- 
lent, round to oval, surrounded or partially covered by ruptured 

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

III. Teleutosori scattered, blackish, pulvinate, roundish on leaves, 

elongated on stems, bursting through browned epidermis, confluent 
in masses, surrounding and distorting stem. 


Uromyces — Chenopodiaceae. 

Teleutospores globose, subglobose or elliptical, smooth, bright 

chestnut brown, apex thickened 

/jl), rounded or occasionally 

somewhat pointed, 22-34 x 16-20 fx, average 28 x 18 fi ; pedicels 
pale yellowish, persistent, firm, long, up to 90 /*. 

On leaves and stems of Polygonum aviculare L. 

Victoria — Near Melbourne, Rutherglen, Myrniong, Goornong, 

Ardmona, Bunyip, Murramurrangbong Ranges, &c. Common 

generally throughout the year. 
Tasmania — Sandy Bay, January, 1906, II., III. (Rodway). 

It is only recorded for Victoria and Tasmania, but is probably to be 
found wherever the host plant occurs. Aecidia were first found on young 
plants along with the other stages in November, 1904. 

The uredospores are plentiful, more particularly in the spring and 
summer months, but the teleutospores are formed in the autumn months. 

Darluca filum Cast., commonly occurring on uredosori, was found on 
teleutosori in May, 1904. 

(Plate XVIIL, Figs. 150, 151.) 


21. Uromyces atriplicis McAlp. 

A triplex. 

Sori epiphyllous, orbicular, scattered, bullate, compact, ruddy brown, 

erumpent, girt by the ruptured epidermis, about \ mm. diam. 

II Uredospores pale brown, ellipsoid, finely echinulate, with 3-5 

scattered germ-pores on one face, 25-29 x 22 ju. 

III. Teleutospores intermixed with uredospores, dark brown, sub-globose 

to shortly ellipsoid, finely striated longitudinally, slightly thick- 

ened at apex 

24-27 h 


with prominent single 

apical germ-pore, 22-29 x 

average 26 

x 25 


On leaves of Atriplex semibaccata R. Br. 

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

The groups of spores shown in the Figures are rather smaller than the 


being about 22 x 25 /u 
occurred reaching a lenjrth of 37 

Occasionally an abnormal teleutospore 


(Plate XVIL, Figs. 146, 147.) 

22. Uromyces betae (Pers.) Kuehn. 

Kuehn, Bot. Zeit., p. 540 (1869) 
Cooke, Grev. XT., p. 98 (1883V 

Beta. . 


Cooke, Handb. Austr. Fung., p. 330 (1892). 
Sacc. Syll. VII., p. 536 (1888). 

ermogonia yellowish to honey-coloured, in small clusters, on both 

faces of leaf 

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

I. Aecidia disposed on orbicular or 


surfaces of leaf, mostly about mid-rib and on leaf 

yellow spots, on both 


peridia white, cup-shaped, with reflexed fringed margin ; peridial 


Uromyc es — Sapinda c ea 


cells firmly united, thickened all round but thicker on one side, 
striated, individually somewhat lozenge-shaped, collectively elon- 
gated polygonal. 

Aecidiospores angular, globose or oblong, orange-yellow, 
smooth, 18-28 x 16-22 //. 

II. Uredosori cinnamon or chestnut-brown, scattered or circinate, 

small, surrounded by the ruptured epidermis. 

Uredospores sub-globose, elliptic or ovoid, yellowish-brown, 
echinulate, with two distinct equatorial germ-pores on one face, 

25-32 x 16-25 /j, average 28 x 21 /j. 

III. Teleutosori dark-brown, margined by the ruptured cuticle, 

scattered or orbicularly arranged. 

Teleutospores ovate or ellipsoid with a colourless papilla at 
apex, dark-brown, smooth, 26-35 X 19-25 fj, average 32 x 21 p; 
pedicels thin, rather long, soon deciduous. 

On leaves of Beta vulgaris L. — Beet and Mangel. 

Victoria — This rust generally occurs wherever beet or mangels are 

cultivated, and has been found at Port Fairy, Maffra, Murtoa, 
Minyip, Ballarat, etc. It also occurs throughout the year 
from Jan. and Feb. to Nov. and Dec. As early as 1878 this 
rust from Ballarat was determined by Thuemen 2 . 

S. Australia — Aug., 1897 (Quinn). 

Tasmania — (Rodway ] ). Devenport, Jan., 1906 (Robinson) 

New South Wales — Hawkesbury Agric. College (Musson). 

The teleutospores are rather scarce, but they were found at Port Fairy 
in August and measured 27-33 x 22-24 /i. 

The aecidiospores were found towards the latter end of August and up 
to December on young leaves of beet growing from roots left in the ground 
during the winter. According to Plowright they very rarely occur in 

Britain in a state of nature. They were found to germinate very freely in 

(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 H . 

On fading leaves of Diploglottis cunninghamii Hook. f. 

Queensland — Woolston Scrub, Brisbane River (Bailey 6 > 19 ). 

This species was very destructive to the foliage of the Queensland 
Tamarind Tree. 

Specimen kindly sent by Mr. Bailey, but spores not obtainable. 



Uromyces — Caryofhyllaceae. 



24. Uromyces caryophyllinus (Schrank) Sch 

Schroeter, Brand, p. 10 (1869) 

Mc Gaz. N.S.W. VII., p. 300 (1896) 

Sacc. Syll. VII., p. 545 (1888) 

II. Uredo 

amphigenous, dark brown when exposed, b 

ed by epidermis, round or oblon 

it long 
ed, often confluent. 


to 3 mm 
U redosp 

ndish to oblong, yellowish b 


decidedly echinulate, 


• » 

x 24 


on one 

generally 3-4 scatl 
face. 21-34 x 17 









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

ed and rasced 

oval or ova 

s at first inte 
cinnamon bro 

mixed with uredospores, roundish, 
ra, generally thickened, with broad 

colourless papilla, average 
pedicels very deciduous, hyaline, up 

28 x 20 ix or 22-32 x 17-23 h i 

On stems 

d both surfaces of 

of Dianthus caryophyllus L. and 

D. chinensis L. — Carnation and Dianthus 

Victoria — Near Melbourne, and at Ardmona. Very common in 

year round 

New South Wa 
South Australia 
Queensland — Roma, 1901 (Try 

— Ashfield 
Feb.. 1902 

Sydney. (Maiden.) 



difference bet 

uredo and teleuto sori 

the naked-eye char 



spores the nisei 

the latter are a little darker 


quite d 

for the uredosp 



decided spines, while the teleutosp 

relatively smooth and 

pilla at ape 

The teleutosp 

filled with finely 



of which is a large vacuole-like body. When 

dry condition, the surface appears to be covered with numerous very fine 

spore is ex 

the centre 



points, which Fischer 5 regards 

ged spines, which project at the 

with loosely arrai 

optical section, while that of the 

points, which are not visible at the marg 

The wall of the uredospore is beset 

margin when seen in 

tospore has densely 

ded fine 




generally described as smooth 

L arluca filum Cast, is often parasitic on the uredosori and 

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

23. Uromyces scleranthi Rostr. 

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

II. Uredosori 




scattered, minute, orbicular or oblong, 
brown, erumpent, and surrounded by the 

slightly raised, 


Uromyces — Zygophyllaceae. 103 

Uredospores golden brown, finely echinulate, ellipsoid to ovoid or 
oblong, 3-4 scattered germ-pores on one face, epispore 3 a thick, 
25-32 x 17-21 11. 

On Scleranthus diander R. Br. 

Victoria — June, 1898 (Reader). 

Only the uredo-stage was found although numerous sori were examined 
and while the uredospores generally agree with those of the above species 
they are sometimes a little longer. 

It was originally found in Denmark on the stems and calyces as well as 
the leaves of Scleranthus perennis L. 

The description is as follows : — " Sori scattered, minute, orbicular or 
oblong, pale brown, long covered by epidermis. Uredospores yellowish- 
brown, globose, echinulate, 15-22 /x diam. or oblong, 24-25 x 18-20 fi 

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

(Plate XXVI1L, Fig. 248.) 


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 /u, average 28 x 20 u. 

III. Teleutospores globose, elliptic, ovate or pear-shaped, thickened at 

the apex, rounded or with a broad apiculus more or less shortly 

21 /i diam., or 23-31 x 17-22 yu, average 25 X 20 a 

epispore thick, smooth, dark bay brown when mature; pedicel 
long, thick, persistent, may be flexuous, pale olivaceous, up to 




On living leaves and stems of Zygophyllum billardieri DC. 

S. Australia — Spencer's Gulf (Tepper). (Winter 


On leaves, stems, and fruit capsules of Z. glaucescens F.v.M. 

Victoria — Near Dimboola, November, 1894, and Warracknabeal, 

March, 1904 (Reader). 
S. Australia — Yorke's Peninsula, June, 1902 (Molineux). 

The confluent sori may form large conspicuous elongated patches up to 
10 mm. long or more, and the greyish bladder-like epidermis remains 
unbroken for some time. Several two-celled teleutospores were met with 
and not constricted at the septum, which was decidedly thick and rather 
below the middle, sometimes approaching the base. They were, in other 
respects similar to the ordinary teleutospores, even to the size, and 
measured about 27 x 22 /u. 



Dietel observes that the occurrence of b 


is rare 

among Uromyces, for only once has he observed it in U. pisi and U. 

d two were found in U 


Cooke has also found 

U. trifolii, and I hav 
culosu8 and U. 

now to add U. orchidearum, U, 


w in 


Darlttca filum Cast., is generally very plentiful on uredosor 

(Plate XV1IL, Fi 

155-157 : Plate G., Fig. 33 


s new genus 


of Rusts is noteworthy, not only on account of the 
teleutospores forming a cluster at the top of the stalk, and generally accom- 
panied by a colorless cyst, but from the large galls produced by some species 
1 • ' • '-- JI - JS and ultimately destroy many of our Wattle-trees 

vhich seriously disfigure 


The seven species at present known show every gradation from one teleu- 

tospore accompanied by a cyst (U. simplex) through two teleutospores with 
or without a cyst ( U. maritimum), ( U. bisporum), and reaching to three teleu- 
tospores in a head always without a cyst (U. tepperianum) . The presence 
of more than one teleutospore at the end of a stalk is a novel feature in 
rusts, and the addition of a colorless vesicle or cyst at the base reminds one 
of Ravenelia to which this genus is allied. The unicellular teleutospore also 

links it on to Uromyces, and I consider that here we have the connecting 
link between Uromyces on the one hand and Ravenelia on the other. The 
genus Anthomyces with only a single species is composed of a head of three 

or more cells united together with sterile cells at the base, and may prove a 
bridging species from Uromycladium to Ravenelia. Spermogonia, uredo 
and teleuto sori occur, but no aecidia. The two gall-producing species 
hitherto found are U. notabile and U. tepperianum, both of which produce 
three teleutospores at the top of the stalk, and the former has uredospores 
in addition. The only species known outside of Australia is that of U. tep- 
perianum (formerly called Uromyces tepperianus, Sacc), and it is noted for 
causing deformation of the shoots, but with us it is ruining whole plantations 
of Wattles. Near Altona Bay, Victoria, the branches of the beautiful 
Golden Wattle (Acacia pycnantha) are deformed and destroyed by it when 
it forms numerous galls sometimes as large as a potato. In the neighbour- 
hood of Melbourne, too, hedges of the Kangaroo thorn (A. armata) are 

Some of the 

being gradually and completely destroyed by this fungus. 
shrubs have most of their branches infested with the chocolate colored 
galls, the color being due to the spores, and they may be in the form of a 
succession of small ones as large as peas, or large ones the size of walnuts. 
The phyllodes as well as the branches in some species are deformed. On 
cutting across these galls they are seen to be solid to the core, and 
not the product of insects but of the fungus, although in many cases bored 
and tunnelled by insects after being produced. When our Acacias are 
more closely examined for rusts no doubt the number of species will be con- 
siderably increased. 

General characters — Spermogonia somewhat hemispherical, produced under 

mouth, preceding the formation of 


the cuticle, without paraphyses at 
other spore. 

Uredospores solitary at apex of basidia, and generally much larger than 
teleutospores with several distinct germ-pores. 

Teleutospores in clusters, composed of one spore and cyst or two or more 
spores with or without a cyst, depressed globose. 

Uromycladium — L eguminosae. 1 05 

This genus may be distinguished from Urornyces by the arrangement of 
the teleutospores, by their shape, which is not elliptical, but depressed globose, 
and by the presence, in most cases, of a colorless vesicle or cyst. 

Allied with Urornyces on the one hand and Ravenelia on the other. 

Australian species, 7. 

Analytical Key to the Species. 

I. One teleutospore and vesicle in head. 

A. Uredospores fusiform, with hyaline apiculus, and finely 


Ur. simplex. 

B. Uredospores sub-elliptical, without hyaline apiculus, thickened 

at apex, and coarsely warted. 

Ur. robinsoni. 

II. Two teleutospores in head. 

A. Uredospores unknown. 

III. Two teleutospores and vesicle in head. 

Ur. bisporum. 

A. Uredospores sub-elliptical, warted all over, much thickened 

and dentate at apex. 

Ur. maritimum. 

B. Uredospores sub-clavate, evenly warted all over, and scarcely 

thickened at apex. 

IV. Three teleutospores in head. 

Ur. alpinum. 

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

spores known. 

Ur. notabile 

B. Teleutospores with converging striae ; uredospores unknown 

Ur. tepperianum 



27. Uromycladium alpinum McAlp. 

O. Spermogonia minute, black, shining, punctiform, crowded,- on both 

surfaces of phyllodes on discoloured patches, appearing before sori 
and ultimately surrounded or accompanied by them. 
Spermatia hyaline, ellipsoid, 3 X 2 fi. 

Sori amphigenous, rusty -brown, scattered or in groups, bullate, soon 

rupturing epidermis and becoming naked. 

II. Uredospores yellowish-brown to golden-brown, shortly or elon- 

gated clavate, occasionally oval or oblong, warted equally all 
over, scarcely thickened at apex, with 3-5 equatorial germ-pores on 
one face, 35-51 x 21-26 //, occasionally reaching a length of 58 \x. 

III. Teleutospores at first intermixed with uredospores, two in head 

with colorless vesicle, depressed globose to subglobose, yellowish- 
brown to dark-brown, smooth, very slightly thickened at apex 
with distinct germ-pore, 19-22 X 25-30 /i ; vesicle arising from 
stalk immediately beneath septum, globose or slightly ellipsoid 
about 25 ju diam. 



Uromycladium — Leguminosae. 


ellipsoid to 







or obovate, rounded at apex, smooth-walled 
and wall of about equal thickness throughout, with colorless 
stalk, 15-25 x 10-15 p. They differ from uredospores in being 
smooth and much smaller, and from the teleutospore in shape, in 
not being thickened at apex and without apical germ-pore. 

On phyllodes of Acacia dallachiana F.v.M. 

Victoria — Alps, near Bright, Dec, 1904 (C French, jr.). 


On phyllodes of Acacia buxifolia A. Cunn 

New South Wales 

New England 

(From type of host in National 

Herbarium, Melbourne.) 
On leaves and pods of A. dealbata Link. 

Victoria — Murramurrangbong Ranges, Jan., 1905 (Robinson). 

Tasmania— Risdon, Dec, 1905, and Mt. Wellington, Jan., 1906 
(Rod way). 

On phyllodes of A. implexa Benth. 

Victoria— Myrniong, May, 1905 (Brittlebank). 

On phyllodes and pods of Acacia linifolia Willd., in National Herbarium, 

New South Wales— Blue Mts. 

Queensland — Rockhampton, Nerbool Creek. 

This species very much resembles U. simplex in the appearance of the 
sori, but it is allied to U. maritimum in bearing two teleutospores and a 
vesicle on one sporophoi'e. It differs from the latter, however, in the 
uredospores which are generally clavate and warted equally all over. 

The vesicles vary in size, and are sometimes large and swollen when 
they are ready to burst. In old material they may have disappeared 
altogether. The teleutospores were much more numerous than the 
uredospores in December. 

(Plate XXIV., Figs. 209-215.) 

28. Uromycladium bisporum McAlp 


III. Teleutosori on the branchlets forming elongated slightly swollen 

chocolate-brown masses 



appearing as powdery patches 

der surface of the leaflets 


depressed globose, occasionally 

two in a head, yellowish-brow 

bsrlobose to 


basal projection, 
mediately beneath, 1 

htly thickened 

-99 x 22-30 H 

y short, colored, stalk 
* apex with germ-por 

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


Murramurrangbong Ranges, January, 1905 (Robinson). 
Risdon, Dec, 1905 (Rodway). 

No vesicle occurs below the septum, so that this species is an intermediate 
form between U. simplex with a single spore and vesicle and U. maritimum 

th two spores and a vesicle in each head 

The occasional presence of 

teleutospores in U. simplex is a further indication of the passage from 


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

Uromycladium — Legnminosae. 107 

9. Uromycladium maritimum McAlp 


O. Spermogonia at first ruddy-brown, ultimately black, dotted over the 

surface of prominent discoid tubercles, formed at corresponding 
points on both surfaces of the phyllodes, somewhat hemispherical, 
but broader than deep, and produced beneath the cuticle, 
averaging 120 jx diam. 

Spermatia Iryaline, shortly ellipsoid, 3 x 2 ji. 

II., III. Sori dark-brown, elongated, compact, confluent, rupturing 

epidermis, 2-3 mm. long, partially surrounding the black discoid 
spermogonial tubercles. 

II. Uredospores oval to elliptical or elongated elliptical, pale-brown to 

dark-brown, warted all over and thickened at apex, where spikes 
are specially prominent, forming a tuft, with very distinct 
equatorial pores, 3-7 on one face, very variable in length and 
breadth, 45-60 X 24-28 \i ; pedicel deciduous, hyaline, elongated, 
up to 106 x 5 ju. 

III. Teleutospores at first intermixed with uredospores, two on each 

sporophore, with a lower and lateral colorless vesicle, very rarely 
three spores in cluster without a vesicle, subglobose to de- 
pressed globose, dark-brown, thick walled, smooth, with finely 
granular contents, slightly thicker at apex, with very prominent 
apical germ-pore, 30-32 \x diam., or 22-25 X 24-30 \x\ colorless 
vesicle globose, with very thin wall, and arising from stalk 
immediately beneath septum, 30-35 \x diam. 

X. Mesospores intermixed with the uredospores, or even with uredospores 

and teleutospores, unicellular or very rarely bicellular, pale, 
smooth, with wall of equal thickness and finely granular contents, 
without any visible germ-pores, fusiform or ellipsoid, or even 
occasionally clavate, solitary at the end of a stalk like the 
uredospores, 22-45 X 11-19 /u, occasionally up to 57 \x long. 
On phyllodes and stems of Acacia longi folia Willd. 

Victoria — Sandringham and Beaumaris. 
Tasmania — Mersey Bluff, Jan., 1906 (Robinson). 

II. April to August, occurring alone, or sparingly mixed with teleuto- 
spores. III. September to latter part of spring and during summer, 
intermixed with a few uredospores. 

In one form or another it occurs all the year round. 

This rust was first found on the coast at Beaumaris in 1895 on Acacia 
longifolia, although not investigated at the time, and it was in this species 
that the peculiar grouping of the teleutospores and the presence of a 
colourless vesicle or cyst was first observed. 

The teleutospores germinate freely in water or moist air and without a 
period of rest. They germinate all round the sorus on the surface of the 
phyllodes, forming a flaky mass of spores and sporidiola, which easily peels off. 

The uredospores were only found to germinate in water in the spring. 

The spermogonia are associated with uredo and teleutospores occurring 
in the same sorus. 

Only in rare instances were sori found without being accompanied by 

spermogonial tubercles. 

(Plates XX., XXL, Figs. 166-184; XXIV., Figs. 216, 217; XXXII 

Fig. 273; XLIIL, Figs. 313, 314, 315.) 



Uromycladhim — Leguminosae. 


30. Uromycladium notabile (Ludw.) McAlp. 

Ludwig, Bot. Centrbl. XLIII., p. 6 (1890). 
Cooke, Handb. Austr. Fung., p. 343 (1892) 

O. Sp 

Sacc. Syll. XI., p. 222 (1895). 
Uredo notabilis Ludw. 
lOgonia minute, punctiform, black 

swollen tubercles often 

intermixed with uredospores and teleutosp 

Spermatia hyaline, ovate or ellipsoid, on elongated basidia, 4 x 

2-3 fx. 
II. Uredos 




surfaces of phyllodes, on branches and pod 
ochraceous to yellowish-brown, seated on a distorted inflated g 


ipsoid to oblon 



yellow when fresh 

3-5 ( 


on elongated hyaline pedicels, with 

quatorial germ -pores on one face; epispore thick (3 p) 
late, 30-45 x 18-28 jz. 

III. Teleutosori on branches, phyllodes and legumes, forming large, 

swollen, distorted galls, chocolate-brown, at length very powdery. 

Teleutospores intermixed with uredospores at first, in clusters 

of three (rarely two or four), sub-globose to depressed globose, 


arranged in 

yellowish-brown, densely covered with warts 

slightly thickened at apex, with germ-pore beneath, 16-23 x 

21-26 /i. 

On phyllodes of Acacia notabilis F. v. M. 

South Australia 

Roseworthy, Sept., 1889, II. (Tepper) 

On branches, leaves and pods of Acacia dealbata Link 






905. O.. II.. Ill 

(Robinson). Bright, June, 1905, II., III. (Davey). Bairns 
dale and Orbost, Dec, 1905, III. 
Tasmania— Hobart, May, 1905, II. (Rod way). Cataract Gorgt 

and Dulverton, Jan., 1906, III. (Robinson) 

On branches of Acacia decurrens Willd 

Victoria — Nea 

1905. Ill 



1905. Ill 

Myrniong, July 

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


On branches of Acacia elata A. Cunn 

New South 


0.> III. (Bake 

the Blue Mountains, April 1905 

On branches of A. binervata DC. 

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

— Gosford, Jan., 1906, II. (Froggatt.) 

New South Wales 

The ochraceous uredosori are generally distinct from the chocolate-bro 

iri, but sometimes the two kinds of spores are found intermixed. 


.me arrangement oi tne teleutospores is generally the same as in 
U. tepperianum, but the dense covering of warty spines instead of distinct 

ally the same as in 

differentiates them at once and the height of the spore 


spermogonia are found in association with both uredospores and teleuto 


The uredospores closely resemble in size and shape those of Uromu 

phijllodiorum, but the surface markings serve to distinguish them 

In the 

Vromycladium — Leguminosae. 109 

latter the warts are arranged in longitudinal lines, while in the former the 
surface markings are net-like. See Plate XXIII., Figs. 201, 202, and 
Plate XXV., Fig. 226. 

Some very large galls were found either surrounding or terminating 
branches of the Black Wattle (A. decurrens). A size of 3-4 inches in diam. 
was not uncommon, and one large clump resembling a big artichoke measured 
5 x 14 inches, and weighed 15 ounces. In some cases the branches still 
Nourished beyond the gall, but it was evidently an expiring effort of the tree to 
put forth leaves. The particular tree on which the largest galls occurred was 
about 30 feet high and 13 years old, but many of the branches were decay- 
ing, and it looked altogether rather dilapidated on account of the numerous 
galls which were often tunnelled by insects. 

The mycelium was evidently perennial, as some of these galls were 

several years old. 

Only the uredospores of this species were found at first, and were 

described as Uredo notabilis by Ludwig. Dietel 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 Ravenelia, 

and, considering the relationship of this genus with Vromycladium, the 

suggestion turns out to be not far from the truth. 

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


31. Uromycladium robinsoni McAlp. 


O. Spermogonia on discoid tubercles, minute, punctiform, ruddy -brown, 

partially or entirely surrounded by uredosori or teleutosori. 

Spermatia hyaline, minute, sub-globose, about 3-4 ll diam. 

II., III. Sori amphigenous, numerous, crowded, up to f mm. diam., 

light rust color, soon erumpent, powdery, and often surrounding 
spermogonial tubercles. 

II. Uredospores pale yellowish, oval to ellipsoid or elongated elliptical, 

thickened at apex (up to 6 /.i), warted all over, particularly at 
apex, with 2-3 equatorial germ -pores on one face, 38-45 x 
19-22 Li. 

III. Teleutospores at first intermixed with uredospores, solitary at end 

of sporophore, with lateral vesicle beneath, golden yellow to golden 
brown, depressed globose to sub-globose, smooth, slightly thickened 
at apex, 19-26 x 25-34 li ; vesicle hyaline, globose to shortly 
ellipsoid, often on distinct stalk with septum at base, 20-25 li diam. 
or 22-27 x 16-23 ll. 

X. Mesospores not uncommon, ellipsoid to elongated ellipsoid or 

elongated oblong, with thin and smooth walls, rounded at apex, 
without germ -pores, 18-22 x 9—12 yu. 

On phyllodes of Acacia melanoxylon It. Br. 

Victoria — Murramurrangbong Ranges, Nov. 1902, Dec. 1903, 

Jan. 1905 (Robinson). Myrniong, May, 1905 (Brittlebank). 
Tasmania — Hobart, Dec, 1905 (Rodway). 

This species was found near Kergunyah by my assistant, Mr. G. H. 

Robinson, in whose honor it is named. The teleutospores give a d 
appearance to the phyllodes on both surfaces, and sometimes the 



ded as to convey the impression of a continuous mass of rust, or even 

! j Uromycladium — L eguminosae. 

red dust. It resembles U. simplex in having a single teleutospore at the 
apex of the sporophore, with a lateral vesicle immediately below, but it 
differs chiefly in the shape and size of the uredospores. The powdery 
masses of teleutospores soon spread over the leaf and germinate in situ, 
forming flakes which are easily detached. 

The spermogonia are seated on tubercles along with uredo and teleuto- 
spores, but there may be powdery patches of both kinds of spores even on 
the same phyllode, without spermogonia or the associated tubercles (Plate 
XXXII., Fi<*. 274). This species represents one of the simplest forms of 
the genus, in which the sporophore bears a single teleutospore with a lateral 
colorless vesicle immediately beneath it. 

(Plate XXII., Figs. 185-189 ; Plate XXXII., Fig. 274.) 


32. Uromycladium simplex McAlp 

O. Spermogonia ruddy-brown, crowded, minute, punctiform, arranged 

in a circinate manner on both surfaces of phyllodes and very 
occasionally on young branches. 

Spermatia hyaline, shortly ellipsoid, 4-5 x 3-4 fi. 

Sori amphigenous, ruddy brown to dark brown, numerous, sometimes 

arranged in groups, bullate, soon rupturing epidermis and be- 
coming naked. 

II. Uredospores yellowish brown to golden brown, fusiform or oval, 

with hyaline apiculus, finely warted, with as many as six equatorial 
germ-pores on one face, three being very common, 48-58 x 21-25 fi 


III. Teleutospores at first intermixed with uredospores, solitary at end 

of sporophore and lateral vesicle immediately beneath, yellowish 
brown, depressed globose to sub-globose, smooth, slightly thickened 
at apex and germ-pore directly beneath, 22-25 X 25-32 ^u ; sporo- 
phore fitting into a sort of socket on base of spore, hyaline, elon- 
gated, 80 f.i or longer; vesicle hyaline, globose 19-22 fx diam. 

On phyllodes and branches of Acacia pycnantha Benth. 

Victoria— Grampians, Nov., 1900 (C. French, jun.), Dec, 1900 

(Robinson). Little River, Nov., 1902 and Jan., 1905 

(C. French, jun.), Feb., 1905. Werribee Gorge, Dec, 1902, 

and Nov., 1904. Malvern Gardens, near Melbourne, Sept., 

Very occasionally the colorless vesicle is replaced by an ordinary spore, 
thus showing that the sporophore bears at its apex two spores, or a spore and 
its substitute. 

The sori form numerous hemispherical pustules which may run together, 
and during the latter part of spring and early summer, while the teleuto- 
spores are being produced, the exuded spores are observed imbedded in gum 
freely germinating and readily detachable in flakes. 

The uredospores somewhat resemble those of Uromyces fusisporus, but in 
the latter there are only 3-4 germ-pores on one face. 

The spermogonia were first found in September on a young tree about 
four years old, and they usually occurred on distinct ruddy spots produced 
by Coniothyrium pycnanthae McAlp. and other fungi. 

Darlucafilum Cast, not uncommon on sori containing both uredo and 

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

Uromycladium — Leguminosae. in 

33. Uromycladium tepperianum (Sacc.) McAlp. 

Saccardo, Hedw. XXVIIL, 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 fx diam. 
Sperrnatia hyaline, ellipsoid, 3-3| x 2-2J li. 

III. Teleutosori on the leaves or phyllodes forming swollen distorted 

gall-like masses along their whole length, and on the branches 

long and broadly effused, or large somewhat spherical galls coated 
with cinnamon to chocolate- brown powdery spores. 

Teleutospores in clusters of three, sphaeroid to depressed 
globose, cinnamon brown, thickly channelled and striate, striae 
converging towards apex, slightly thickened in upper portion of 
wall, 14-17 X 18-25 fx ; sporophore hyaline, elongated, soon 
On branches of Acacia salicina Lindl. ; A. hakeoides A. Cunn. ; A. 
myrtifolia Willd.; and A. spinescens Benth. 

S. Australia — Blackhills, Sandy Creek, Murray Bridge, etc., 

Dec, 1889 and 1892 (Tepper). Dec, 1901 (Molineux). 

On phyllodes and branches of A. armata R. Br.; A. implexa Benth.; 
A. juniperina Willd. ; A. melanoxylon R. Br.; A. pycnantha Benth.; A. 
rigens A. Cunn.; A. siculiformis A. Cunn.; A. vomeriformis A. Cunn. 

Victoria — Mallee near Hopetoun, Oct., 1903 (C. French, jun.). 

Ringwood, Aug., 1904, (C. French, jun.). Werribee Gorge, 
Jan., 1905, (Brittlebank). Oakleigh, Jan., 1905. Little 
River, Jan., 1905, (C. French, jun.), and Feb., 1905. 
Cheltenham, May, 1905, (Robinson). Myrniong, July, 1905. 
Mt. Macedon, 1882, and Murray River, 1874, from types of 
A. siculiformis and A. vomeriformis in National Herbarium, 
Melbourne. Common around Melbourne. 
On A. diffusa Jjindl. ; A. vemiciflua A. Cunn. ; A. verticillata Willd. 

Tasmania. — Hobart, March- April, 1905 (Rodway and Lea). 

On Acacia longifolia Willd. 

New South Wales — Rose Bay, near Sydney, July, 1905 (Froggatt). 

On branches and phyllodes of A. erioclada Benth., and A. glaucoptera 

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 U. notabile of producing 
galls. This is particularly noticeable in the Golden W T attle (A. pycnantha), 
where the galls are as large as potatoes, and in some of the Wattle 
plantations where the trees are cultivated for their bark they hang in 

1 12 


large numbers from the branches like so many fruits, and the trees are 

dving or dead. (See Frontispiece.) 

The dark chocolate-brown spore-masses are quite powdery, and each spore 
nrominent ribs running from base to apex, so that it is easily known 


fluted. In fact, the appearance closely resembles the marki 
on the e£L's of some butterflies, and no doubt the purpose is the same 


then the memb 



the spore collap 

when dry conditions pr 

ly thick, and prever 
Occasionally a spor 

has been found germinating in situ, and they germinated freely in water 

in twenty hours. 

What is said to be the same fungus has been found on Albizzia t 
Benth., in Java, and it would be interesting to know if it occurred 
indigenous species of this genus in New South Wales, Queensland c 

1 any 


On A. implexa at Myrniong there 


and I found 

at the end of a branch in July somewhat of a leg-of-mutton shape and 
weighing about 3 lbs. (PI. XLI. 

Witches' brooms of 



rred on A. implexa, caused by this rust, and one of the 
iired 45 inches in circumference (PL XLIL). 


(Plates XXII., Figs. 190-195; XXIIL, Fig. 206; XXXIV.; XXXY 

XLI.; 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, sucli as chrysanthemum and corn- 
flower, hollyhock, marigold and daisy ; even a parasite, such as the native 
mistletoe, is subject to it. The two-celled teleutospore is easily recognized 
and distinguished from the unicellular uredospore. All the spore-forms may 
be present on the one plant, as in P. hederaceae on the native violets (Viola 
hederacea and V. betonicifolia) , or reduced to the teleutospore alone, as in 
P. malvacearum. There may also be heteroecious forms, such as P. caricis, 
with the aecidial stage on the nettle (Urtica). The teleutospore, although 
normally two celled, and with a horizontal septum, is sometimes very vari- 
able in these respects. It may not only, occasionally, be one-celled, but in 
P. dichondrae, for instance, it may be 3-4 celled, and it may be vertically, 
obliquely, or even muriformly divided by the septa. The most celebrated of 
all the species is P. graminis, or wheat rust, which seems to have lost the 
power here of infecting the barberry, for although germinating promycelial 
spores have been used upon specially imported barberries and rusty wheat 
grown around the latter, still no aecidia have been produced. 

Mesospores are common and paraphyses may be present in both the uredo 
and teleuto-layer. 

General characters. — Spermogonia when present, mostly epiphyllous, 
minute, sub-globose or flask-shaped, honey-coloured. 

Spermatia very minute, globose or ellipsoid, hyaline. 

Aecidia when present at first globose and closed, then cup-shaped and 

open, or elongated and cylindrical, with margins generally everted. 

4*1* • • j • • • i i - 

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


Puccinia — Gramineae* 


Uredosori when present, generally minute and flattened, sometimes 

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. 



34. Puccinia agropyri Ell. & Ev. 

Agropyron, Clematis. 

Ellis and Everhart, Journ. Myc. VII., p. 131 (1892). 
Sydow, Mon. Ured. L, p. 823 (1904). 
Sacc. Syll. XI., p. 201 (1895). . 

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

0. Spermogonia amphigenous, honey-coloured, in clusters on leaves, 

accompanying aecidia. 

Spermatia hyaline, globose, minute, 3 li diam. 

1. Aecidia hypophyllous, on definite spots, bright orange, in irregular 

clusters ; pseudoperidia cup-shaped, flattened, with reflexed lobed 
margin ; peridial cells piriform to quadrate, striated at margin 
and punctate all over, 28-32 x 16-22 li 

Aecidiospores orange-yellow, ellipsoid to sub-globose, finely 
echinulate, 21-29 x 16-19^, or 20-22 li diam. 

II. Uredosori epiphyllous, rarely on under surface, minute, elliptic or 

linear, sometimes confluent, yellowish to orange. 

Uredospores orange yellow, elliptical to ovoid, finely echinulate, 
4-5 scattered germ-pores on one face, 25-32 x 19-22 li, 
epispore up to 2 \ li thick. 

III. Teleutosori hypophyllous, minute, covered by the leaden-coloured 

epidermis, at length occasionally erumpent, oblong or linear, 
sometimes forming lines on sheaths. 

Teleutospores yellowish brown, cylindric clavate to elongated 
oblong, smooth, slightly constricted at septum, variable in size, 
• 40-70 x 12-25 /j, average 60 x 18 li ; upper cell rounded or 
squarely truncate at apex and decidedly thickened (6-9 li), some- 
times broader than long, darker in colour than lower, 19-29 x 
16-25 fi ; lower cell tapering towards pedicel or oblong, usually 
longer and narrower than upper, 25-41 x 12-20 li ; pedicel short, 
generally tinted. Occasionally three-celled teleutospores occur. 
X. Mesospores occasional, pale brown, elongated, rounded, or slightly 

pointed and thickened at apex, slightly tapering towards base, 
40-48 x 13-16 li, with short coloured pedicels. 
Aecidiospores on Clematis aristata R. Br. 

Victoria. — Murraruurrangbong Ranges, Dec, 1903, and Jan., 

1905 (Robinson). 

^^v^^ & 

Uredo and teleutospores on Agropyron scabrum Beau v. 

Victoria — Near Melbourne, Dec, 1892 (Robinson). Myrniong, 

March, Nov., Dec, Jan. Murramurrangbong Ranges, 
Dec, 1903 (Robinson). 


Puccinia — Gramineae. 

The teleutospores vary considerably in size and 


are some 

Limes elongated cylindric but usually cylindric clavate, the lower cell being 
comparatively narrow and tapering slightly towards base, while the upper 
cell is expanded and considerably flattened out at apex. The length may 


from 40-70 li and the breadth fro 

It is noticeable 



of elongated and 

thers of medium-sized spores 

In the Agricultural Gazette for New South Wales, Vol. VI., p 

1 895, the rust on this native grass was given as 
Henn., but on further examination I find that it a 
P. aqropyri Ell. and Ev. and is named accordingly 

P. d 

Eriks. and 

pproaches most closely 

Specimens of P. agropyrina Eriks., were examined from Eriks. Exs. 419 


evens Beauv., and the uredospores measured 21-24 jx diam 

or 24-27 x 16 



The teleutosp 


were considerably 

smaller than our own material, measuring only 33-42 x 1 

39 x 15 u. 


r 1 


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

Uredospore . 


25-32 x 19-22 n 40-70 x 12-25 /* 

P. agropyrina Eriks. on Agropyron repens, Beauv. 16-25 fi diam. 36-41 x 13-16 ft 

/'. agropyri Ell. & Ev. on A. glaucum, Roem & Schult. 20-25 x 18-22 ft 60-75 x 20-25 /* 
P. agropyri Ell & Ev. on A. scabrum, Beauv. 

Although the uredospores are described as smooth by Ellis and Everhart 
yet in specimens from Sydow's Ured. Exs. 1362, they are seen to be 

is of the P. 

decidedly echinulate and it is a question whether such 

a thing as a 

uredospore exists among Puccinias. 




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 

It is suggestive 

Klebahn, 1 p. 292, Puccinia (clematidi) agropyri Ell. and Ev. 

that both stages have been found in the Murramurrangbong Ranges growing 
near to each other. 

(Plate III., Fig. 25.) 

35. Puccinia agrostidis Plow. 


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

McAlpine, Agr. Gaz. N.S.W. VII., p. 149 (1896). 
Sydow, Mon. Ured. L, p. 717 (1903). 
Sacc. Syll. XL, p. 202 (1895). 

II. Uredosori elliptical to elongated, soon naked, linear and confluent. 

Uredospores orange yellow, elliptical, finely echinulate, as many as 
9 germ-pores seen on one face, forming a circle inside epispore, 

22-24 x 17-18 


III. Teleutosori 

minute, black to dark-brown, long covered by the 

epidermis, sometimes elongate, sometimes in groups. 
Teleutospores dark-brown, 

smooth, oblong or subclavate, 


thickened (up to 5 /;), truncate or rounded, markedly constricted, 
attenuated below, almost sessile, very shortly stalked, 40-54 X 

14-21 ^ 

average 44 

x 18 /i 



Puccinia — Gr amine ae. 


X. Mesospores fairly numerous, similarly coloured to teleutosp 


elongated elliptical, thickened at apex, 24-30 x 


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 orierinal 

material, and there is a general agreement in the spores. 

Dr. Cobb has described and drawn an unnamed species of Puccinia on 
Deyeuxia forsteri, which undoubtedly belongs to the same species. The 
teleutospores are given as 44-58 x 16-22 /j,, but the uredospores are rather 
larger, being 25-30 X 21-25 /x. 

The life-history of this species was made out by Dr. Plowright, who found 
after several experimental cultures, that the teleutospores produced Aecidium 
aquilegiae Pers. on Aquilegia vulgaris, and that the spores of A. aquilegiae 
when applied to Agrostis alba and Poa pratensis produced the rust. 

(Plate III., Fig. 27.) 


36. Puccinia anthoxanthi Fckl 

Fuckel, Symb. Myc. II., p. 15 (1873). 
McAlpine, Agr. Gaz. N.S.W. VII., p. 301 (1896). 
Sydow, Mon. Ured. I., p. 727 (1903). 
Sacc. Syll. VII., p. 665 (1888). 

II. Uredosori on both surfaces of leaves, solitary or in elongated 

groups, elliptic or linear, confluent in lines, soon naked, pulver- 
ulent, dusky orange. 

Uredospores yellowish-orange, elliptic to obovate, finely echinu- 
late, with two to four very distinct equatorial germ-pores on one 
face, 25-32 x 15-20 p. 


III. Teleutosori scattered, minute, dark-brown to black, elliptic, naked, 

surrounded by ruptured epidermis. 

Teleutospores at first intermixed with uredospores, chestnut- 
brown, elliptic to obovate or oblong-clavate, smooth, slightly 
constricted, rounded and thickened (6 fj) at apex, occasionally 
tricellular, 28-48 x 15—21 fi, average 35 x 20; upper cell 
usually darker than lower; pedicel persistent, tinted, 
20-25 x 6-7 /j,, sometimes reaching a breadth of 10 fi. 

X. Mesospores occasional, similarly coloured to teleutospores, obovate, 

thickened at apex, average 35 X 17 fx. 

On sheath, flag and inflorescence of Anthoxanthum odoratum L. 

Victoria — Near Melbourne, Dec, 1896, II. Rutherglen, Dec, 

1903, II., III. Leongatha, Feb., 1904., II. 

The teleutospores were not very common. In my own garden, near 
Melbourne, the uredo-stage was plentiful, but no teleutospores were found, 
while at Rutherglen the teleuto- stage occurred on several specimens. 

(Plate III., Figs. 20, 21.) 



Puccinia — Gramineae . 


7. Puccinia beckmanniae McAlp. 

II. Uredosori on both surfaces of leaf, but most common on under, 

orange, elliptic, often confluent in lines, erumpent and surrounded 
by ruptured epidermis. 

Uredospores orange, elliptic, echinulate, with 3 to 4 equatorial 

germ pores on one face, 25-29 x 16-19 /.i 

long covered by 

III. Teleutosori minute, black, linear, hypophyllous, 

epidermis, ultimately naked, about \ mm. long. 

Teleutospores yellowish-brown, elongated clavate, not or 



slightly constricted at septum, upper cell generally darker than 
lower and with numerous (up to 

45-60 x 16-25 p, 





56 x 18 ; pedicel very short or 

X. Mesospores brown, oblong to elongated elliptical, thickened at apex, 

and either bare or with short stumpy processes, 32-35 x 13-16 //. 

On Beckmannia erucaeformis Host. 

Victoria — Leongatha, February, March, 1904, II., III. 

Only uredospores were met with in February, but by March the teleu- 
tospores had developed. There is a general agreement with P. lolii, but the 
upper cell of teleutospore is generally broader. 

from seed sent by 

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. 


The grass on which this rust was found was 
United States Department of Agriculture for trial, and the spore 


Darlucafilum Cast., was very plentiful, both by itself and 

Probably in the former 


the hidden 

mycelium, and may have largely prevented spore formation 

(Plate II., Fig. 12.) 

38. Puccinia bromina Eriks. 


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

Puccinia dispersa, f. sp. bromi, Eriks. 

II. Uredosori 1-1 0mm. long, 1 

broad, ferr 


on upper surface 


leaf blade 

ded, sometimes on sheath and 


bright orange, 

with numerous scattered 

germ-pores, as many as 11 on one face, echinulate, 27-31 x 



III. Tele 

ated elliptical to oblong, black to dark-brown 

_ ^„ v „ v,xv,.i 6 o,^u euipuiciu w oDiong, DiacK to dark- brown, 

hypophyllous, scattered or subgregarious, sometimes on sheath and 


ed by epidermis, divided into 

each compartment separated by elongated, cylind._ 
clavate, yellowish-brown, barren filaments or paraphy 

partments, and 
5, or slightly 

Puccinia — Gr amine ae. 


Teleutospores oblong to clavate, dark chestnut-brown, con- 
stricted at septum, rounded or truncate at apex and slightly 
thickened, 40-56 x 18-24 li, average 48 x 21 li ; lower cell 
generally narrower than and almost as dark as upper ; pedicel 
short, hyaline to pale yellowish, up to 36 11 long. 

X. Mesospores comparatively rare, dark chestnut-brown like teleuto- 
spores, clavate, oval or oblong, rounded or truncate and slightly 
< . thickened at apex, base sometimes attenuated, 28-36 x 16-21 u. 

On Bromus mollis L. 

Victoria — Nalinga, Nov., 1898 (Robinson). Kergunyah, Nov., 

1902, Dec, 1903, and Jan., 1905 (Robinson). Domain, 
Melbourne, Dec, 1904, II., III., the latter very plentiful. 
Nagambie, Nov., 1904. 

New South Wales- 1890 (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 /u, thus being rather smaller 
than the above. The teleutospores were much paler in colour, and measured 
36-48 x 15—22 yu, the largest being about the average of the Victorian 

Fritz-Muller 1 observed aecidia on Symphytum officinale L. and Pulmonaria 
montana Lej., and by infection with the aecidiospores produced the rust on 
Bromus. He therefore named it Puccinia symphyti bromorum. 

(Plate III., Fig. 28 ; Plate C, Figs. 11-13.) 


39. Puccinia cacao McAlp 

* _ 

TJredo rottboelliae Dietel, Engler's Bot. Jahrb. XXXII., p. 5 


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 T 
32-40 x 24-32 h average 35 x 28 li. 

III. Teleutospores intermixed with uredospores, sparse, somewhat 

paler, oblong, smooth, rounded or flattened and unthickened at 
apex, slightly constricted at septum, 32-35 x 21-22 p ; lower 
cell tapering slightly towards base and generally about equal in 
size to upper ; pedicel hyaline, deciduous. 


Puccinia — Gramineae. 

On leaves and sheaths of Rottboellia comp-essa L 

Victoria— Creswick, Jan., 1893, II. (Robinson), WerribeeGo 


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 17 ). 

The Queensland specimen is labelled Puccinia straminis DeBary. 

The chocolate-brown colour of the uredospores is very characteristic. The 
uredo-stage of this species was first described by Dietel on a specimen from 
Japan ; on sending him some of our material he agreed that it was the 
same. Sydow in his Monograph (p. 800) has described a Puccinia on 
R. arundinacea with teleutospores alone, which however do not agree with 
these, being thickened at the apex up to 10 n and 34-56 x 22-27 n in size. 

The uredosori frequently look quite black owing to the presence of 

Darluca JUum Ca 

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

40. Puccinia cynodontis Desm 


Desmazieres Exsicc. III., No. 655. 

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

Sydow, Mon. Ured. I., p. 748 (1903). 

Sacc. Syll. VII., p. 661 (1888). 

P. altera McAlp., Agr. Gaz. N.S.W. VII., p. 151 (1896). 

II. Uredo-sori on both surfaces, minute, ellipsoid to lenticular, scat- 
tered or confluent, yellowish-brown to rusty -brown, soon naked, 


Uredospores globose to shortly elliptical, yellowish-brown to 

brownish-yellow, delicately verrucosa, 20-25 /i diam., or 20-25 x I 



III. Teleutosori roundish to ellipsoid or oblong, scattered or confluent, 

pulvinate, black. 

Teleutospores intermixed with uredospores, variously shaped, 
ellipsoid or oblong, smooth, chestnut-brown, slightly constricted at 
septum, occasionally tricellular, 30-50 x 15-21, average 35 x 
21 ^i ; upper cell thickened at apex and generally elongated 
pointed, sometimes rounded ; lower cell rounded at base or 
attenuated; pedicel firm, pale yellow, persistent, elongated up 

to 73 


X. Mesospores plentiful, dark-brown, ellipsoid, rounded and thickened 

at apex, 2t x 19 /<. 

On Cynodon dactyl on Pers. 

Victoria — Burnley, Caulfield, Armadale, Pakenham, Killara, 

Somerville, etc., Oct. -March. 

On some plants the teleutospores were of a very regular ellipsoid shape, 
and rounded and thickened at apex, so that it was at first thought there 
were two species on the same plant. But on examining a number of 
specimens every gradation was found from the teleutospore, rather constant 
in size and shape with rounded apex, to those necessarily longer on which 
the thickened apex was somewhat conical. 

Puccini a — Gramineae 


It was likewise found by Magnus 5 that two kinds of uredosp 

occur, the one thin-walled with numerous (up to 9) germ-pores and 

echinulate, while the other is thick walled, with few (1-3) germ-pores and 

either smooth or only with a few scattered spines. There were transition 
forms between the two, and this shows how variable a description might be 
according to the nature of the spores regarded as typical. 

(Plate III., Fig. 24.) 


41. Puccinia festucae Plow. 

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

109 (1893). 
Sydow, Mon. Ured. I., p. 752 (1903). 
Sacc. Syll. XL, p. 194 (1895). 

II. TJredosori mostly on under surface of leaf, but generally present 

on upper as well, causing conspicuous yellow spots on upper 

surface, minute, oblong to elliptic, scattered or confluent, orange- 

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

III. Teleutosori mostly on under surface of leaf, minute, scattered or 

often in groups and confluent, oblong to linear, brownish-black. 

Teleutospores at first intermixed with uredospores, clavate to 
cylindrical, brown, smooth, slightly constricted at septum, generally 
tapering towards base, 40-60 X 15-20 jjl, often 60 x 16 \x.\ upper 
cell more or less truncate and thickened at apex, surmounted 
• by a crown of 4-6 obtuse, straight or curved, sometimes bifid 
processes ; lower cell generally elongated wedge-shaped ; pedicels 
persistent, brown, stout, 15-25 \i long. 

X. Mesospores very common, similarly coloured to teleutospores, clavate 

to cylindrical to somewhat oblong, thickened at apex and sur- 
mounted by processes, 31-46 x 1 1—14 yu. 

On Festuca ovina L. 

Victoria — Leongatha, July, 1903, II. III. 
On F. rigida Kunth. 

Victoria — Rutherglen, Nov., 1895, II. 

In 1890 Plowright experimentally proved that the aecidium on Honey- 
suckle (Lonicera) was genetically connected with this species, and it has 
since been repeatedly proved by Fischer and Klebahn. 

(Plate II., Fig. 13.) 

42. Puccinia flavescentis McAlp 


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

II. TJredosori on upper surface of leaf, minute, linear, often confluent 

soon naked, p ulverulent, rusty brown, arranged along furrows of leaf 




Puccinia — Gramineae 



golden-brown, with at least 5 germ-pores 
21-24 p. diam. or 25-32 x 21-24 p. 

to shortly elliptical, finely echinulate, 

irregularly distributed, 

III. Teleutosori minute, elliptical, numerous, black, often confluent 

lengthwise, soon naked. 

Teleutospores intermixed with uredospores, dark chestnut brown, 
oblong, constricted at septum, with rounded and thickened apex (up 

to 9 p), smooth, occasionally tricellular, 33-48 x 18-26 p, average 
44 x 24 p ; upper cell generally hemispherical, and about equal 
in length to lower ; lower cell generally rounded at base, some- 
times narrowed and elongated like upper portion of pedicel ; pedicel 
persistent, tinted, elongated, up to 72 /j. long. 

X. Mesospores common, similarly coloured to teleutospores, elongated 

ellipsoid, rounded or truncate and thickened at apex, smooth, 
34-43 x 12-15 p. 

On Stipa ftavescens Labill. 

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

On Stipa semibarbata It. Br. 

Victoria — Nagambie, Nov., 1904, II., III. 

The pulverulent uredosori, and the numerous minute, black teleutosori 
are characteristic of this species. The uredospores form a rusty powder over 
entire upper surface of leaf. The pedicel of the teleutospore is sometimes 
lateral and the septum erect as in Diorchidium. It differs from P. stipae 
Arthur, in the uredosori being soon naked and decidedly ruddy brown, not 
yellowish, while the uredospores are broader. 


specimens of P 

2S, on Stipa 

(Op.) Hora, taken from Syd. Ured. Exs. N 

illata L., the teleutosp 

generally bluntly pointed, and the size 48-54 x 

are decidedly different 





In speci 

mens of P. stipae Arth., from Arthur and Holway's Ured. Exs. No. 27, on 

Stipa spar tea T 
rather thicker 

tel eu tosp 


pointed at the ape 


I have submitted specimens to Prof. J. C. Arthur, and he remarks that 
it is clearly distinct from his species, although there is very much similarity 
between the two, as one might expect, from the hosts being essentially alike. 

(Plate II., Fig. 17.) 

43. Puccinia graminis Pers. 


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

II. Uredosori amphigenous, yellowish-brown, linear, 2-3 mm. or 

longer, either scattered or confluent in long streaks, especially on 

sheaths, pulverulent, soon naked, surrounded by ruptured epi- 

Puccini a — Gr amine ae, 



Uredospores elongated ellipsoid to ovate oblong, brownish yellow, 
echinulate, generally with 3-4 equatorial germ-pores on one face, 
20-36 X H-18/i. 

III. Teleutosori sparingly on leaf blades, more commonly on sheaths 

stalks and inflorescence, linear, elongated, pulvinate, often con- 
fluent, up to 10 mm. or more, dark brown to dense black, soon 
rupturing epidermis which is prominent. 

Teleutospores clavate to oblong clavate, chestnut brown, smooth, 
somewhat constricted at septum, very rarely three-celled, 35-63 
X 14-25 fx $ average 52 X 18ju; upper cell rounded or pointed at 
apex, rarely truncate, considerably thickened (up to 12 //), some- 
times as broad as long, 21-29 fi long ; lower cell attenuated to- 
wards base, equal to or longer than upper, 18-35 p long; pedicel 
persistent, elongated, tinted, and sometimes as deeply coloured as 
spore, of equal thickness throughout, up to 73 x 8 fj. 

X. Mesospores very common, intermixed in the same sorus with uredo 

and teleutospores, similarly coloured, dark chestnut or paler, oblong 
to elongated ellipsoid, generally slender, rounded, pointed or trun- 
cated apex and thickened like teleutospore, smooth, 34-46 x 10- 
15 /j, ; pedicel generally short or of moderate length. Occasionally 
an obovate form occurs, deeply coloured, rounded and thickened at 
apex and much broader than usual, up to 22 \x, with elongated 
pedicel . 

On Wheat (Triticu?7i vulgar e VilL), Polish Wheat (T. polonicam L.), 
Oats {Avena sativa L.), Barley {Hordeum vulgar e L.), Rye {Secale cereale L.), 
and various species of the following genera of Grasses: — Agropyrvti, 
Alopecurus, Amphibromus, Avena, Beckrnannia, Briza, Bromus, Dactylis y 
Deyeuxia, Echinopogon, Elymus 9 Festuca, Glyceric^ Ilordeum, Phalaris, Boa. 

Common in all the States. 

Occasionally I have seen a uredospore with a slight indentation on either 
side, just where the circle of germ-pores occurs, but never any indications of 
a septum. The range of variation in the size of the teleutospores is great. 
It may vary from 35 X 25 fi in the oblong, dark-brown, perfectly mature 

spore to 63 ju, long, and sometimes only 14 ft broad in elongated clavate spores 
in the same sorus. 

The spore itself may be colourless, while the brown colouration is in the 
pedicel, and this has previously been observed by Dr. Plowright in Austra- 
lian specimens ; or the upper cell only may be pale in colour as shown in 
] Plate I., Fig. 8, which is from abnormal material met with on one occasion. 

There are no paraphyses, but there are numerous mesospores closely re- 
sembling teleutospores, but only one-celled. 

Darluca filum Cast., sometimes common on the uredosori, particularly 
on Wild Oat (Avena fatua) and Glyceric stricta. It has also been found on 
Agropyron scabrum, and Dr. Cobb 5 evidently mistook its true nature, for 
he says : — " Among its red-rust spots there are certain black bodies which 
may constitute a fourth spore of the rust." 

The following is a complete list of all the grasses and cereals on which I 
have found Puccinia graminis Pers. This determination of the fungus has 
been based on morphological characters alone, for I have not yet succeeded 
in infecting the Barberry. Those marked with an * are also recorded by 

E 2 



Puccinia — Gramineae 

have been found 

other authors, and a double asterisk indicates those 

by Eriksson in Europe to produce aecidia on the Barberry when their spores 

were sown upon it : 

Agropyron divergens Nees 
*A. scabrum Beauv. 

Alopecurus geniculatus L. 

Amphibromus neesii Steud. 
*Avena fatua L. 

• • • 

# • • 

• • • 

• • 

• • 

• • • 

• • • 

• • • 

- • • 


• * • 

• • 

. - • 

Beckmannia erucaeformis Host 

*Briza minor L. 
Bromus racemosus L. ... 
**B. secalinus L. 

• • • 

• • 

• - • 

• • • 

• • • 

• • • 

B. sterilis L. ... 
Dactylis glomerata L. 
Deyeuxia quadriseta Benth 
Echinopogon ovatus Beauv, 
Elymus striatus Willd. 
E. virginicus L. 
Festuca bromoides L. 

• - • 

• • 

• • • 

• • • 

• # 

• • 

• • t 

• • • 

# • 

• • 

• • 


• • 

• • 

• • • 

G. stricta Hook, f . 

• • • 

• • • 

• # • 


• • 

# • t 

H. secalinura Schreb. 

**H. vulgare L. 
**Phalaris canariensis L. 

P. minor Retz. 
**Secale cereale L. 

Triticum polonicum L. 

•*T. vulgare Vill. 

• • 

• • • 

• • 

• • 

* • 

* ♦ • 

• • • 

• • t 

• • • 

* • 

• • • 

• • • 

• • • 

• ■ • 

• • • 

• ■ • 

• - • 

• • 

• • 

• • 

• • 

• • 

• • 

• ■ 

• • • 

• • • 

• • • 

• • 

• • • 

• * • 

• • « 

• • • 

• • • 

• • • 

9 • • 

• • 

• • • 

• • • 

• • • 

II. , HI. 

II. , HI. 

II., 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 
are capable of infecting each other. 

various grasses 

(Plates I., Figs. 2, 5, 7, 8 ; XIV., Figs. 113-122 ; XL., Fig. 301 j 

XLIIL, Fig.' 311 ; Plate A., Figs. 3, 4.) 

Puccinia — Gramineae. 


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, l-l£ mm. long, pulvinate, 
erumpent, powdery, reddish-brown. 

Uredospores bright orange, subglobose or ellipsoid, finely 
echinulate, up to six scattered germ-pores on one face, 25-33 x 

III. Teleutosori hypophyllous, greyish -black, pulvinate, long covered by 

epidermis, densely crowded, linear, often confluent and arranged in 
lines between the veins, with dark-brown paraphyses in clusters. 

Teleutospores brownish, oblong to oblong-clavate, smooth, 
slightly constricted at septum, 35-55 X 16-22 ^, average 
40 x 16 fi ; upper cell rounded or truncate, and thickened at 
apex (3-5 p) ; lower cell generally paler in colour, and rounded or 
attenuated at base ; pedicel very short, hyaline or tinted, but 
dark-brown at apex. 

X. Mesospores not uncommon, pale brown, slightly thickened at apex, 

elongated, oblong, or obovoid, 29-35 x 12-17 /x. 

On living leaves of Elymus condensatus Presl. 

Victoria — Rutherglen, Dec, 1903. 

The seed of this grass was obtained from the United States, and the 
rust was doubtless imported with it. 

Arthur sowed germinating teleutospores from Elymus virginicus L. on 
Impatiens aurea Muhl., and in fifteen days aecidia were produced. This 
result was confirmed in the succeeding year. 

Specimens on E. virginicus, from Iowa, in Sydow's Ured. Exs. 1380, are 
labelled Puccinia rubigovera (DC.) Wint., and agree with this species. 

There is another Puccinia on the same host plant (P. procera Diet, and 
Holw.), and according to the description it is quite distinct. The 
uredospores are 32-45 x 30-38 /j. and the teleutospores 45-100 \x long. 

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

(Plate III., Fig. 26.) 


45. Puccinia lolii Niels. 

Nielsen in Ugeskrift for Landmaend. I., p. 549 (1 

I SydoM, Mon. Ured. I., p. 704 (1903; 

Sacc. Syll. XL, p. 203 (1895) 

P. coronifera Klebahn, Zeitschr. f. Pflanzenk. II., p. 340 


II Uredosori on upper and under surfaces of leaves, forming blister 

like swellings, oval or linear, orange, soon bursting through epider 
mis, pulverulent, often confluent, sometimes up to 2 mm. long. 



Puccini a — Gramineae . 

Uredospores orange- yellow, subglobose, ovate or broadly elliptic, 

strongly echinulate, with 2-3 equatorial germ-pores on one face, 

20-30 x 16-24 \x' y occasionally a few colourless capitate para- 

physes intermixed. 

III. Teleutosori on both surfaces of leaves, minute, black, linear or 

' oblong, often confluent, long covered by epidermis, \-\ mm. long. 

Teleutospores yellowish-brown to brown, elongated clavate, 


towards base, smooth, constriction very slight or abse 


sometimes unicellular, 35-60 x 12-20 h average 53 x 16 p 

upper cell brown, truncate, thickened at apex, giving off irregular 

blunt, generally curved processes variable in size and shape 

avera'^in" 6 to 7 or more ; lower cell generally paler in colour 


longer than upper, and attenuated towards base ; pedicel pei 
tent, coloured, short, stout, up to 11 /u long by 8 p broad. 

X. Mesospores exactly resembling teleutospores only one-celled. 
On stem, leaf, and inflorescence of Lolium perenne L. Common. 


Victoria— Near Melbourne, Myrniong, Port Fairy, &c, Aug. — May. 
South Australia — Mount Gambier, April, 1903, II., III. 

The teleutospores may be found as early as September, and through the 
summer and autumn. 

Tn some districts this rust is very severe and kills the grass down to the 
roots, especially if there are early autumn rains and warm weather. 

Darlucafilum Cast., on uredosori. 


Puccinia lolii avenae. 

On Avena saliva L., and Avena fatua L. 

Victoria— Port Fairy, Dec. 1903, and Sept.— Nov. 1904, II., III. 

Brighton, Jan., 1904, II., III. Myrniong, Nov., 1904, II. 
Near Melbourne, Jan., 1905, II., III. 

New South Wales— (Cobb 12 

Eriksson has shown that the form occurring: on Lolium will not infect 

Avena, nor the reverse, so that there are two biologic forms, and that found 
on the Oat may be distinguished as P. lolii avenae. 

This species may occur alongside of P. graminis, and the teleutosori 
were found together on the sheath of the cultivated oat, A. sativa, and! 

the wild oat, A. fatua. 

The teleutospores on the wild oat are sometimes very much longer than 
the normal. They range from 70 to 86 /i in length, and from 18-22 \i in 
breadth at the apex, and 7-8 fi at the base. 

By infection experiments Klebahn 2 proved the connexion between this* 
rust, and the aecidium on Phamnus frangula L., but this genus does no 
occur naturally in Australia. 

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

Puccinia — Gramineae. 12 


on Rhamnus cathartica, while the original name was retained for the rust 
producing its aecidia on Rhamnus frangula. Nielsen had previously used 
the name of P. lolii for Klebahn's species, and so it is retained on the score 
of priority. 

(Plate II., Figs. 11, 14 ; Plate XXX., Fig. 261 ; Plate B., Figs. 5-8.) 

46. Puccinia magnusiana Koern. 


Koernicke, Hedw. XV., p. 179 (1876). 
Sydow, Mon. Ured. I., p. 785 (1904). 
Sacc Syll. VII., p. 631 (1880). 

II. Uredosori on both surfaces of leaf, but mostly on upper, snuffy 

brown, erumpent, surrounded by ruptured epidermis, elliptic or 
linear, confluent lengthwise and forming long streaks, with clavate 

bright yellow paraphyses, 

Uredospores elliptic or obovate, golden yellow, echinulate, with 
four equatorial germ-pores, 24-35 X 16-19 /x; intermixed with 
large numbers of clavate paraphyses, club thickened at apex, of a 
dark smoky -brown and stalk hyaline, commonly 80-90 /x long. 

III. Teleutosori minute, black, very numerous, scattered, elliptic or 

linear, confluent into long black streaks on both surfaces of leaf, 
but mostly on upper, occasionally paraphysate as in uredosori. 

Teleutospores clavate to oblong, dark chestnut-brown, generally 
rounded and thickened at apex (up to 12 /x), sometimes bluntly 
pointed, hardly constricted at septum, 35-55 x 14-21 ^u, average 
38 X 18 yu ; lower cell attenuated towards base ; pedicels firm, 
persistent, coloured yellowish, about length of spore or longer, 
up to 70 jjl. 

X. Mesospores similarly coloured to teleutospores, variable in shape, 

somewhat ovoid to elongated, thickened at apex, 28-38 x 13-19 p. 

On Phragmites communis Trin. 

Victoria — Orbost, Aug., 1901 (Pescott). Flinders, Jan., 1902. 

Port Fairy, June-Aug., 1902 and 1905. Killara, March, 

1903. Bunyip, May, 1904. 
South Australia — The Grange, April, 1891 (Tepper). River 

Torrens, Adelaide, Apr., 1903 (Tepper). 
Tasmania — (Rod way ] ). 

The clavate paraphyses are usually described as hyaline, but Dr. Dietel 
compared the Australian specimens with intense dark-brown paraphyses 
with material from Europe, America, and the Cape of Good Hope, and he 
found that the latter were sometimes scarcely coloured, at other times light 
or dark brown. 

The rust on Phragmites communis Trin., was considered as one species up 
to 1876, viz., Puccinia phragmitis Schum. Then Koernicke separated it 
into two, viz., P. phragmitis and P. magnusiana. The former was 
characterized by the absence of paraphyses from the uredosori, the large 
bullate teleutosori and the very long stalked markedly constricted teleuto- 
spores, while the latter had numerous clavate paraphyses in the uredosori, 
minute, punctiform or linear teleutosori, and shortly stalked, hardly 
constricted teleutospores. The aecidia in the two cases likewise occurred 
on different hosts. 


Puccinia — Gramineae. , 


ch showed that there were still other species on the 

host-plant. Plowright separated a third species in 1888, mainly based 
upon the fact that the aeeidiospores were only produced upon Rumex 

and not upon other pi 

Next, Lud 


1892 distinguished another 

material sent from South Austr 

P. tepp 


Arthur in 1902 published another new species which he named P. sim 
because it was very similar to P. magnusiana. 

There are thus at least five different species of Puccinia recorded upon 
Phragmite8 communis, and it is not always easy to differentiate them, at 

by morphological characters 

In three of them the 


proved to occur on different host-plants by 

dered by many a sufficient specific distinction 

of cultures, and this is 

On these 

ounds they 


w«uld be considered biological species as in the case of Puccinias on various 
cereals and grasses, but at the same time an attempt will be made here 
show any points in which they differ morphologically. Only two of the five 
have hitherto been found in Australia, viz., P. magnusiana and P. tepperi y 
for although Dr. Cooke gives P. phragmitis in his Handbook of Australian 
Fungi, it was probably intended for P. magnusiana, which has been found 

Victoria and South Australia. 
The life-history of this fungus was first determined 

Dr. Plowright, 5 

who found that the promycelial spores produced aecidia on Ranunculus, 
repens and R. bulbosa, and conversely the aeeidiospores produced uredo- 
spores and teleutospores on Phragmites. This was afterwards confirmed by 
Fischer' and Klebahn.' 

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

Rumex sp. and 

Rumex acetosa. 


Uredosori without paraphyses. 

Ranunculus repens 
and R. bulbosa. 

Anemone dichotoma. 

Uredosori with paraphyses. 

P. phragmitis. 

25-35 x 15-23/i 

45-65 x 16-25 /i 
markedly con- 

150-200 n long 

P. trailii. 

25-35 x 20-25 \x 

50-60 x 20-23 \i 
markedly con- 

P. tepperi 


75-100 fi long 

27-30 x 20-23 \i 

45-68 x 18-26 fi 
markedly con- 

P. majrnusiana. 

180-250 fi long 

24-35 x 16-19^ 

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

About length of 

P. simillima. 

26-44 x 18-22 fi 

42-56 x 15-20 fi 
hardly con- 

About length of 

It will be seen from the above that the 

are two typ 


distinguishable morphologically, paraphysate and aparaphysate^ and with 
these limits there are only "biological species " based upon a differ 
aecidial host. 

If we compare P 


having a granular spore-membrane, and shorter 

th P. phragmitis, the teleutospores differ 




similar comparison is made of P. tepperi, the length of the pedicel is the 

triking feature, and 


ther premature to 


P. phragmitis until a distinct aecidial host has been proved. 
According to Arthur, P. simillima is distinguished from P 

it from 

the slender pointed 


more clavate and less 

redospores and the probably different aecidial host 

dichotoma L 





(Plate II., Fig. 1 

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 jx. 

III. Teleutosori amphigenous, scattered or subgregarious, variable in 

form, mostly linear or oblong, long covered by epidermis, pro- 
minent, very black, 1-2 mm. long. 

Teleutospores obovate to oblong or subclavate, bright chestnut 
brown, smooth, constricted at septum, rounded or conoid and 
thickened at apex (up to 8 /x), sometimes truncate, roundated, 
base and rarely somewhat attenuated, occasionally 3 or 4 celled, 
32-52 x 16-24 ^u, average 36 x 20 ji ; upper cell rarely 

vertically divided ; epispore thick ; pedicels persistent, elongated, 
yellowish, thickened particularly towards apex, up to 90 p. long. 

X. Mesospores occasional, similarly coloured to teleutospores, ovoid, 

rounded at both ends, thickened at apex, average 35 x 17 /*. 

On leaves of Zea mays L. 

Victoria.— Seville, April, 1897, II. (Hill). Tally Ho, April, 1902, 

II. (Cronin). Leongatha, March, 1904, II. Near Mel- 
bourne, April, 1904, II., III. 
I New South Wales. — Richmond (Cobb). 

Queensland. — Toowoomba, 1887 (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 
I to be distinct Sydow in his Monograph has reserved the above name for 
this one. 

Arthur* inoculated maize plants with uredospores from an aecidium on 
Oxalis cymosa Small, and in five days uredosori appeared in abundance. 
The Oxalis is supposed to have been inoculated from the teleutospores of 
P. maydis, but this infection has still to be verified. 

Darluca Jilum Cast., occurs on the uredosori. 

! , 

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

48. Puccinia perplexans Plow. 

Alopeciwus . 

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

II. Uredosori amphigenous and on sheath, orange, elliptic, rupturing 

epidermis and often confluent in lines. 



Puccinia — Gramineae. 

Uredospores orange-yellow, subglobose to elliptic, finely echin 

late, with 4-6 scattered germ-pores on one face 

9 x 19-22 n 

II [. Teleut 

ed epider 

phigenous and on sheath, covered by leaden- 

l confluent in very 

first minute 


oblong, constricted 

ng lines and even forming patches. 

Teleutospores variable in shape, clavate to 
ptum, brown, upper cell darker than lower, rounded or truncate 

apex and slightly thickened, lower cell attenuated towards base 

j 37 x 17 u; pedicel short, 

ded. 3 






hyaline, may reach a length of 20 

X. Mesospor 

16-21 yu 

On Alop 

at apex. 

ilarly coloured to 

teleutospores, slightly 
to ovate or pear-shaped, 25-32 x 



Victoria— Minyip, Nov., 1903 (Eckert) 

Among these plants one was badl) 
nnia graminis but no trace of the 


usted with the uredosp 



of that fu 


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. I., p. 795 (1904). 

Sacc. Syll. VII., p. 625 (1888). 

IT. Uredosori small, round or elliptical, scattered or aggregated togethe 

orange-yellow or foxy- red. 

Uredospores shortly elliptical, finely echinulate, orange-yellow, 

21-24 x 15-18 fi, seated on hyaline pedicels up to 30 n long, and 

intermixed with numerous, stiff, capitate paraphyses, reaching a 
length of 68 fi. I 

III. Teleutosori small or large, scattered or orbicularly disposed, 

dark-brown to black, covered by the epidermis. flj 

Teleutospores elliptical or subclavate, very variable, apex 


truncate, rounded or conical, thickened 

8 fi), dark -brown, 



smooth, not or hardly constricted at septum, 35-45 X 15-21 jj 

or may reach a length of 53 y. when 3-celled, average 38 x 19 /i ; 
pedicels persistent, short, brown. 

ives and stems of Poa annua L. 

Victoria— Near Melbourne, Jan.-Dec, 1886. &c. II.. III. 


South Wales 

Sydney, May, 1898 

Sydney, 1890 (Cobb). Botanic Gard 

On Poa 


Botanic Gardens, Melbourne, Dec, 1904 

On Poa vratensis L 


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

Puccinia — Gramineae 



Uredospores have been found all the year round and teleutospores from 
August to December. Teleutospores are fairly plentiful in October, and 
they are often three — or even four — celled. The three-celled forms have 
either transverse septa, or the upper cell may be divided vertically or 
slightly oblique. The four-celled is either produced by transverse septa, 
tlie upper one being usually oblique, or the upper third may be divided 
longitudinally. In these abnormal forms the length may reach 53 u and 
the breadth 34 li at the apex. 

The genetic connexion between the aecidium on Coltsfoot (Tussilago 
farfara) and this rust was first shown by Nielsen in 1876 and repeated by 
Plo wright in 1882, who found that the aecidiospores produced the uredo- 
spores on Poa annua in ten to twelve days. But the Coltsfoot does not 
exist in Australia, and, therefore, this rust can reproduce itself without the 
intervention of aecidia. As might be anticipated the uredo stage carries 
it over the winter, and, indeed, the fungus is most plentiful in our winter and 
early spring months — June, July, August, and September. Poa annua 
as a rule dies away early in October, as soon as the warm weather comes on, 
except in moist shady places. Lagerheim 1 , has found the spores on the 
leaves after the melting of the snow. 

Darluca filum Cast., commonly 

(Plate III., Fig. 22 


50. Puccinia purpurea Cooke 


Cooke, Grev. V., p. 15 (1876). 
Sydow, Mon. Ured. T., p. 803 (1888). 
Sacc. Syll. VII., p. 657 (1888). 

Uredo sorghi Fckl. 

II. Uredosori amphigenous, seated on indeterminate, elongated and cod- 

fluent, bright red to purple spots, scattered or in small irregular 
groups, covered by epidermis, and then splitting, yellowish-brown, 
1-1 J mm. long. 

Uredospores ellipsoid, obovate to piriform, yellowish-brown, 
roughly aculeate, with 3-4 germ-pores on one face, 28-34 X 
20-25 ix, average 31 x 21 li ; paraphyses intermixed. 

[III. Teleutosori on similar spots, mostly hypophyllous, oblong, elliptic, 

or linear, always or long covered by epidermis, reddish-brown. 

Teleutospores ellipsoid, oblong or ovate-oblong, rounded at apex, 
not or scarcely thickened, not or very slightly constricted at 
septum, mostly rounded at base, smooth, bright brown, with very 
thick epispore, 35-56 X 22-32 li ; pedicel hyaline, persistent, 
thick, up to 100 fi long ; paraphyses present.] 

On leaves of Johnson Grass (Sorghum halepense Pers.) and Sugar-cane 

{S. vulgare Pers.). 

h 15 

Queensland — Gladfield (Gwyther), Nerang (Shirley), (Bailey 

Only uredospores have been met with here. 

In P. maydis the teleutospores are thickened at apex. Originally P. 
sorghi Schwein. included rust on Zea and Sorghum, but it is now found 
that the same rust does not occur on both, and Sydow, in his Monograph, 
has wisely determined to reserve P. purpurea Cooke, for Sorghum and 
P. maydis Bereng., for Zea. 

Darluca filum Cast., is very common on uredosori. 


T ^o Puccini a — Gramineae. 


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

Eriksson and Henning, Getreider. p. 238 (1896). 
McAlpine, Journ. Dep. Agr. Victoria L, pp. 430 and 

529 (1902). 
Sydow, Mon. Ured. I., p. 756 (1903). 
Sacc. Syll. XVII., p. 377 (1905). 

Puccinia straminis Fckl. var. simplex Koern. in Land und 
- Forstwirtsch. Zeit. (1865). 

II. Uredosori very minute or up to \ mm. long, sparingly scattered or 

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

Uredospores subglobose or ellipsoid, echinulace, yellow, relatively 
thick walled, with numerous scattered germ-pores, 4-7 on one 

face, 21-24 ft diam, or 24-30 X 17-20 u. 
III. Teleutosori covered by the epidermis, amphigenous 


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

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

X. Mesospores very numerous, similarly coloured and similarly 
thickened at apex to teleutospores, asymmetrical, sometimes 
elongated, saccate or inversely triangular, very variable, 24-45 x 
15-24 fi. 

On Hordeum vulgare L. 

Victoria — Port Fairy, Warrnambool, Nov.-Jan. 

This form was first described in 1865 by Koernicke as a variety, but now 
Eriksson and Henning have raised it to the rank of a species. 

A very characteristic feature of it is the great predominance of unicellular 
spores. A sorus may consist entirely of these or there may be a few normal 
teleutospores intermixed- They are produced alongside the two-celled forms, 
and are quite similar to them except that they are one-celled, and they 
evidently show the transition from the one-celled to the two-celled teleuto- 
spore. There is usually a slight thickening at the apex of both, but it rarely 
exceeds 4-6 \x. It was early recognised as an exceptional form, and was 
named Uromyces hordei by Nielsen in 1875, and Puccinia anomala by Rostrup 
in 1876. 

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

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


Puccinia — Gramineae. j?j 

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 li diam. or 21-28 x 18-20 li. * 

III. Teleutosori similar to uredosori, but dark-brown. 

Teleutospores intermixed with uredospores, dark chestnut- 
brown, oblong to clavate, rounded at both ends or slightly 

tapering towards base, Slightly constricted ;it septum and 


thickened at apex (4-7 ^u), smooth, 28-46 x 19-23 ll ; pedicel 
persistent, yellow, up to 56 ll long. 

X. Mesospores not uncommon, similarly coloured to teleutospores, 

obovate, rounded and thickened at apex (up to 7 li), smooth, 
28-37 X 19-22 li ; pedicel similar to that of teleutospore. 

Darluca jilum Cast., common on uredosori. 

On living leaves of Distichlis maritima Rafin. 

Victoria — Flinders, Jan., 1902. Near Melbourne, April and 

Nov., 1905, II., III. (Robinson). 

This species is quite distinct from P. distichlidis E. and E., which, 
however, was originally found on Spartina gracilis Trim, the host-plant 
being mistaken for a Distichlis. 

Arthur 7 has obtained some remarkable results by sowing the teleuto- 
spores of this species derived from Distichlis sficata Greene, on various plants. 
The germinating teleutospores produced aecidia on three widely-separated 
families of plants, viz. — Chenopodiaceae, Capparidaceae, and Cruciferae. 

According to the Index Keivensis, D. spicata is a synonym of D. maritima. 

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

53. Puccinia tepperi Ludw. 


Ludwig, Zeitschr. f. Pflanzenk. II., p. 132 (1892). 
Sydow, Mon. Ured., I., p. 792(1904). 
Sacc. Syll. XL, p. 203 (1895). 

Sori epiphyllous, large, scattered or gregarious and confluent, up to 

3 cm. long, firm, thick, pulvinate, erumpent, dark-brown ; 
paraphyses in clusters, hyaline or pale yellow, capitate, thickened 
at apex, up to 75 ll long. 

II. Uredospores ellipsoid or pear-shaped, echinulate, pale yellowish, 

with as many as 9 scattered germ-pores on one face, 27-30 x 
20-23 ll. 

III. Teleutospores intermixed with uredospores, subcylindrical to 

lly oblong, with rounded or acute slightly thickened 

apex (4-5 ^u), markedly constricted at septum and upper cell often 

becoming detached, rounded or rarely attenuated at base, smooth 
yellowish-brown, occasionally 3-4 celled, 45-68 x 18-26 li 


50 x 19 ll; pedicel hyaline, persistent, up to 250 



Puc cinia — Gr amine ae . 

X. Mesospores very rare, similarly coloured to teleutospores, elongated 

slightly thickened 

ellipsoid, rounded and 

slightly towards base, 58 x 21 ii 

On Phragmites communis Trin. 

at apex, tapering very 

S. Australia — The xG range, near Adelaide, April, 1891 (Tepper) 


& )' 

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). 
Svdow, Mon. Ured. I. p. 716 (1903). 
Sacc. Syll. XVII., p. 376 (1905). 

P. dispersa i. sp. tritici Eriks. and Henn., Zeitschr 


Pflanzenk. IV., p. 257 (1894). 
II. Uredosori 1-2 mm. long, reddish-brow] 




on upper surface of leaf -blade, elliptic, sometimes confluent, scat- 
tered or somewhat gregarious, occasionally on sheath and stem. 
Uredospores subglobose to shortly ellipsoid, echinulate, orange- 

yellow, 4-6 scattered germ-pores on one face, 20-28 x 18 



average 48 x 16 fx ; upper cell deeply 

III Teleutosori oblong, black to dark-brown, hypophyllous, scattered, 

often arranged lengthwise in lines, sometimes on sheath and stem, 
covered by epidermis, divided into compartments surrounded by 
brown paraphyses. 

Teleutospores clavate to oblong, smooth, yellowish-brown to 
dark-brown, slightly constricted at septum, very occasionally 

3-celled, 39-57 x 15-18 yw, 
coloured, generally rounded or flattened at apex and thickened, 
17-31 x 15-18 jx ; lower cell paler and attenuated towards pedicel, 
longer and narrower than upper, 22-36 X 12-14 li ; pedicel 
short, coloured. 

X. Mesospores occasional, similarly coloured to teleutospores, ellipsoid 

to clavate or sub-clavate, shortly stalked, smooth, slightly thickened 
at apex, 25-38 x 13-16 li. 

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

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

Australia, and Tasmania. 

The uredospores 
subglobose instead 

distinctly different from those of P. graminis in being 

of elongated 

d the more numer 

scattered instead of forming an equatorial band 


Uredospores taken from young wheat plants in the winter months (June 

August) readily germinated in a moist chamber. 

(Plate I., Figs. 3, 6, 10; Plate XL., Fig. 302 ; Plate A., Figs. 


Puccinia — Cyperaceae 

l 33 


55. Puccinia Caricis (Schum.) Rebent. 

Rebentisch, Fl. Neom., p. 356 (1804). 
Sydow, Mon. Ured. L, p. 648 (1903). 
Sacc. Syll. VII., p. 626 (1888). 

Carex, Urlica, 

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

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

Spermatia hyaline, ellipsoid, 4J x 2jw 

I. Aecidia in rows or groups on yellowish or reddish spots, closely 

crowded, reaching 1 mm. in diam.; pseudoperidia cup-shaped, 
becoming wide at the mouth and campanulate, flattened or 
outstanding, with whitish margin which is toothed and reflexed. 

Aecidiospores orange, polygonal to subglobose, finely echinulate, 
16-26 x 12-20 fi. 

II. Uredosori elliptical or oblong, minute, hypophyllous, scattered or 

subgregarious, often confluent in long lines, erumpent, girt by the 
ruptured epidermis, ferruginous. 

Uredospores ellipsoid or ovoid, yellowish brown, echinulate 
1-2 germ-pores on one face, 25-30 x 19-24 li. 

III. Teleutosori minute, dark-brown or black, round to elliptic, 

scattered or gregarious, sometimes confluent, compact. 

Teleutospores clavate to elongated oblong, dark-brown, smooth, 
constricted at septum, apex much thickened (up to 12 /j) and 
rounded or bluntly pointed, lower cell generally attenuated 
towards base, 35-64 x 16-22^, average 52 x 22 li ; pedicels 
tinted, persistent, generally short, up to 30-40 n long. 

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

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

Victoria. — Bacchus Marsh, Myrniong, Fern Tree Gully, etc. 
Uredo and teleutospores on leaves of Car ex breviculmis 1Z. Br. 

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

Tasmania — Cascades, May, 1892, II., III. (Rodway). 
On Carex gunniana Boott., and C. inversa, R. Br. 

Tasmania — Longley, Dec, 1894, II. (Rodway). 
On Carex alsophila F.v.M. 

Victoria — Murramurrangbong Ranges, "Dec., 1903, II. (Robinson) 
On Carex paniadata L. 

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


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 — Cy-peraceae. 



experimentally proved, in 


that Aecidium urticae 

DC, was genetically connected with Puccinia caricis occuring on Car ex hirta 

Dr. Plowright also, in many cultures, has shown that 

aecidiospores pro 

duce the uredospores on species of Carex, and the teleutospores produce the 
aecidiospores on Urtica. 

On the under surface of the leaves the aecidia occur in small groups, but 
on the leaf -stalks and stems they are in large, bright-orange patches, 
thickening and distorting the parts on which they occur. 

Magnus 1 has also shown that this rust winters on Carex hirta by 

of the uredo 


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


56. Puccinia cyperi Arthur. 

Arthur, Bot. Gaz., p. 226 (1891). 


-2 mm 

Sydow, Mon. Ured. I., p. 681 (1903). 
Sacc. Syll. XL, p. 199 (1895). 

II. Uredosori hypophyllous, on brown effused spots, scattered 

larly or aggregated, ellipsoid to oblong, variable in size, 1 

lon< r , long covered by the thick epidermis, ochraceous or brownish. 

Uredospores yellowish-brown, ellipsoid, echinulate, 22-25 x 

19-21 ». 

III. Teleutosori hypophyllous, on similar spots, 1-3 mm. long, almost 

always covered by the epidermis, blackish brown. 

Teleutospores intermixed with uredospores, elongated oblong to 

gated fusiform, brown and paler beneath, smooth, constricted 

dly constricted 

57 X 




ge 48 x 

15 \x ; upper cell rounded or often obtusely and obliquely 

at apex, and thickened (up to 13 

cell paler, oblong, but 

usually attenuated towards base ; pedicel coloured like lo 
persistent, 20-30 /x long, but may reach a length of 40 \x 


X. Mesospores generally paler in colour than the teleutospores, ellip- 
soid to fusiform or even curved, obtusely and sometimes obliquely 
pointed at apex, thickened (up to 5 ju), 30-32 X 13-17 /t ; pedicels 
similarly coloured, persistent, up to 22 /.i long. 

On leaves of Nut grass (Cyperus rotundus L.). 

Queensland— Gatton, Aug., 1903 (Shelton) (Bailey, 18 , 20 ). 

The teleutospores are very variable in shape, and sometimes the cells 
may be placed almost at right angles to each other. They are sometimes 
excessively elongated, and may reach a length of 67 /i. 

Intermixed with the teleutospores there are numerous stalked bodies 
similarly coloured, and representing what are usually called mesospores, 
although Sydow describes them as paraphyses. While mesospores generally 
have an undoubted resemblance to the teleutospores with which they are 
associated, their function in this case seems to be that of paraphyses, and 
it may be in some other cases as well. 

Darhica filum Cast., was common on uredosori. 

(Plate IV., Fig. 32.) 




Puccini a — Cyp erac eae 


57. Puccinia longispora McAlp. 

Car ex. 

II. Uredo-sori hypophyllous, minute, elliptic to elongated, confluent in 

long lines, long covered by epidermis, rusty- brown. 

Uredospores sub-globose to shortly elliptic or obovate, yellowish- 
' brown, thick-walled, echinulate, with three equatorial <*erm-pores 

on one face, 22-30 x 19-22 /*, or 23-27 jx diam. 

III. Teleutosori black, compact, elongated, bullate, crowded, confluent 

in long black lines, covered for some time, then girt by ruptured 

Teleutospores clear yellowish-brown to smoky-brown, elongated, 

narrow, smooth, constricted at septum, 50-80 x 14-19 yu, average 

63 x 19 fx ; upper cell darker than lower, generally rounded and 

m slightly expanded at apex, sometimes bluntly pointed, much 

thickened (9-12 ^u) ; lower cell longer than upper, almost cylin- 
drical or slightly attenuated towards base ; pedicel yellow, short 
and often stout. 

On Carex caespitosa L., and C. vulgaris Fr. 

Victoria — Werribee Gorge, January, 1903. Gembrook Ranges, 
April, 1904 (C. French, jun.) 

On Carex sp. 

Victoria — Killara, March, 1903 (Robinson). 

In immature forms of teleutospore the upper and lower cells are much 
alike, but ultimately the lower cell is much elongated and nearly equally 
broad throughout. Although 33 different Puccinias are recorded on species 
of Carex, the elongated narrow teleutospore with short pedicel is quite 
characteristic for this species. In P. schoeleriana Plow, and Mag., the teleu- 
tospores may be equally long, but the lower cell is more cuneiform than 
cylindrical, and the pedicel may reach a length of 57 \x. An examination of 
teleutospores from Carex arenaria L. in Syd. Ured. Exs., No. 282, showed 
these differences distinctly. 

Specimens of P. caricis (Schum.) Rebent. were examined from Exsicc. 
Sydow Ured. No. 1065 and Exsicc. Briosi and Cavara, No. 12$, and the 
teleutospores were decidedly different. 

The lower cell in P. longispora is much more elongated and narrower 
and gradually merges into the short, stout and coloured pedicel, while in 
P. caricis besides being generally considerably shorter, more wedge-shaped 
and somewhat inflated, the pedicel was much narrower. It is in the upper 
cell, however, that the difference is most striking. In P. caricis the thick- 
ened apex is one-half the length of the cell, while in P. longispora it is only 
one-third, taking the average of a number. The thickened apex, also, in 
P. caricis is much darker than in P. longispora. Of course, as in so many 
other cases where there are a number of species on the same genus of host- 
plant, there is a family resemblance too, between the rusts, and P. caricis 
and P. longispora have many points in common along with differences in 

(Plate IV., Fig. 31.) 





Puccini a — Juncaceae. 



58. Puccinia juncophila Cooke and Mass. 

Cooke and Massee, Grev. XXII., p. 37 (189 
Sydow, Mon. Ured. I., p. 64-1 (1903). 
Sacc. Syll. XL, p. 199 (1895). 

Uredo armillata Ludwig, Bot. Centrbl. XLIII. p. 8 (1890). 

II. TJredosori ruddy-brown, ellipsoid, crowded, confluent, often girdling 

stem, surrounded by the ruptured epidermis and sparingly covered 
by shreds of it. 

Uredospores sub-globose, elliptic or piriform, very spinulose, 
relatively thick-walled (3-4 fi) yellowish-brown, 24-32 x 16-22 p. 

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

epidermis, confluent. 

Teleutospores intermixed with 
olden-brown, with finely granular contents, slightly constricted 


ptical to oblon 



Jed and a little thicker at apex (5-6 /x) 

at septum, rour 

rounded at base and generally resembling upper, but often some 

what narrower, 30-40 X 


smooth : 




ge 36 



ally decid 

25 a 

, oftt 

ttached obliquely, up to 54 p long 

X. Mesospores similarly coloured to teleutospores, shortly elliptical, 

occasionally globose, smooth, thickened at apex, 23-28 x 19—21 yu. 

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

Victoria — Coromby, Oct., 1889, II. (Tepper), the original of 
Uredo armillata Ludw. Diraboola, Nov., 1891, II., III. 


Oakleigh, April, 1893, II., III. (Morrison). 

Ardmona, Christmas Hills, Killara, Millbrook, Minyip, 
Mordialloc, Myrniong, near Melbourne, Phillip Island, 
Stawell, Werribee Gorge, Murramurrangbong Ranges, &c. 

Tasmania — (Rod way 1 ). 
In the original description of this species by Cooke and Massee, some 

mistake must have been made in the measurement of 

they are given as 16 

18 x 12-14 u. 


Through the kindness of Dr. Morrison, I ha 
some of the original material from Oakleigh. 

been able to examine 

Uredo armillata was the name given at first by Ludwig in 1 889, f r 

edospores only being found on Juncus 

y of teleutospores intermixed with uredosp 

but the subsequent dis- 
ss showed that it was a 


referred to Uromyc 

Material sent by Reader to Kew Herba 





by the authorities there, but the identical 

later forwarded to us the 


found mixed with the uredospores 
shown to exist in Victoria. 




therefore, has not bee 

Darluca fit 



y common 

in some cases rendering the 

patches quite black. It occurred both on uredo and teleutosori 

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



Puccinia — Juncaceae\ 


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 u. 

III. Teleutosori dark-brown, ellipsoid, scattered or crowded, often con- 
fluent, pulvinate, compact, soon naked, on similar spots. 

Teleutospores oblong to clavate, ochrey-yellow, fragile, smooth, 
apex generally rounded, rarely truncate or conical, thickened up to 
1 1 /u, attenuated towards base or rounded, constricted at septum, 
30-50 x 14-20 fi, average, 42 x 17 /u ; pedicel hyaline, per- 
sistent, generally about 30 \x long. 

X. Mesospores abundant, clavate to obovate or oblong 


ened at apex and similarly coloured to teleutospore, 22-32 x 
13-16 /u. 

On leaves and stems of Luzula campestris L. 

Victoria — Murramurrangbong Ranges, Nov. and Dec, 1902-3, 

Jan., 1905 (Robinson). Wandin, Nov., 1903 (C. French, 

On Luzula oldfieldii Hook. f. 

• Tasmania— Mt. Wellington, Jan., 1892, II. (Rodway). 

Cooke in his Handbook of Australian Fungi gives Aecidium bellidis 

Thuem. which is supposed to represent the aecidial stage of Puccinia obscura 
Schroet., occurring on Luzula, but the aecidium found on Bellis perennis here 
is associated with its own teleutospores, and belongs to a distinct fungus, 
P. distincta McAlp, so that whatever may be the case in Britain, the rust 
on the daisy is autoecious with us. 

The discovery of a rust on Luzula campestris, which does not belon 


the Old World species, is a further proof against its genetic connexion with 
that of the daisy. The species differs from P. obscura and P. oblongata. 
The teleutospores are characteristically fragile, thin- walled, and easily 
collapsible, and while of the same general type as in P. oblongata, they are 
much smaller. In P. obscura the colour is much deeper, and the wall is de- 
cidedly thicker and firmer. This is well shown in Plate IV., Fig. 34, in 
which the spores are taken from a specimen of Luzida campestris in Sydow's 
Ured. Exs., 1076, collected in 1896, and the material from which our de- 
scriptions are made is as recent as 1905. In P. oblongata the apex of the 
teleutospore is much thicker, being 10-20 /u, and the size is 40-80 X 16-24 /u. 

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

Darluca filum Cast, occurs frequently on the uredosori. 

(Plate IV., Figs. 33, 34.) 


Puccinia — Liliaceae 


Bur char dia 

60. Puccinia burchardiae Sacc. 

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

Ludwig, Zeitsch. f. Pflanzenkr. III., p. 137 

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

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


Sacc. Syll. XL, p. 197 (1895). 

II. Uredosori amphigenous, bullate, elliptical or sometimes circular, 

crowded, light-brown, erumpent, surrounded at base by dry cuticle 
of epidermis. 

Uredospores elliptic to subglobose, yellowish-brown, epispore 
echinulate, 28-31 x 22-25 p. 

numerous, small or large, erumpent 

III. Teleutosori amphigenous, 

surrounded at base by cuticle of epidermis, circular cr elliptical, 


Teleutospores clavate, constricted at septum, thickened at apex 

rounded, sometimes truncate or acute. 


to 14 or 


40-60 x 




gth of 70 

48 x 20 u, but sometimes 


u; lower cell tap 

towards pedicel, yellowisl 

'own; upper cell elliptical or rarely spherical, chestnut-brown, 
21-23 n broad; epispore smooth; pedicel hyaline, 30-60 X 5-6 jj. 

X. Mesospores fairly common, similarly coloured to teleutospores or 

paler, clavate to ( 
37-43 x 12-15 u 

ded or pointed at apex and thickened 

On stems and leaves of Eurchardia umbellata R. Br. 

Victoria — Dandenong Creek, Oct., 1891. (C. French, jun.). 


Melbourne, Sept. — Feb. Myrniong, Sept., 1898, and Dec, 
1903. Murrumurrangbong Ranges, Dec, 1903 (Robinson). 

S. Australia — Teatree Gully, Oct., 1890 (Tepper). 

The first record of this species 

appeared in Hedwigia 

of March and 

April, 1893, where Saccardo described it as 

P. metanartheeii Pat 

differing from the type 

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


61. Puccinia wurmbeae Cooke and Mass. 

Cooke and Massee, Grev. XVI., p. 74 (1887). 
Cooke, Handb. Austr. Fung., p. 337 (1892). 
Sydow, Mon. Ured. I., p. 640 (1903). 
Sacc. Syll. VII., p. 664 (1888). 

Sori elongated, bullate, dark-brown. 

II. Uredospores elliptic, granulate, brown, 25-28 x 15-18 u 

III. Teleutosp 


htly constricted at septum, smooth 

upper cell convex or truncate, at apex darker, lower cell tri- 

5 into the short, hyaline pedicels 

angular, attenuated downward 

X 21-28 /z, average 

Massee, 60-70 x 20-25 A t) ; 



but attaining a length of 


broadest at the junction with the spore 

ii (according to Cooke and 
pedicels persistent, generally short, 
ix and a breadth of 


8 ft, bein 



Puccinia — Haemodoraceae, Amaryllidaceae. 139 

X. Mesospores numerous, similarly coloured to teleutospores, elon" 

elliptical to oblong, thickened at apex and acute, rounded or 
cate, 40-49 x 17-21 fi. 

On leaves of Wurmbea dioica F. v. M. 

South Australia — Beltana, 1887 (Richards). 

No uredospores were found on the specimen in National Herbarium 

(Plate V., Fig. 37.) 



02. Puccinia haemodori P. Henn. 

Hennings, Hedw. XL., p. (96) (1901). 
Sydow, Mon. Ured. I., p. 609 (1903). 
Sacc. Syll. XVII., p. 366 (1905). 

III. Teleutosori amphigenous, round or often oblong, scattered or 

aggregated and then confluent, pulvinate, compact, erumpent, black, 
surrounded by the ruptured epidermis, 1-2 mm. long. 

Teleutospores oblong to oblong clavate, obtusely rounded or 

apiculate at apex and strongly thickened (8-13 ju), slightly con- 
stricted at septum, attenuated or rounded at base, smooth, 
chestnut-brown, 40-57 x 14-20 /u, average 46 x 19 ji, pedicel 
persistent, slightly brownish towards apex, 40-55 x 5-7 jx. 

X. Mesospores very common, subfusoid or ovoid to subclavate, apex 

gibbous to apiculate, somewhat obtuse or two-horned, thickened at 
apex (up to 9 //), brown, 25-40 x 12-18 jjl ; pedicel hyaline to 
brownish. Occasionally a more deeply coloured spore occurs, 
with somewhat truncated apex, 30-46 x 5-7 fi. 

On leaves of Haemodorum sp. 

West Australia— Near Perth (1900). 

Hennings described uredospores in his original diagnosis, but Sydow did 
not find them afterwards in the original material. He remarks that the 
teleutospores readily separate into their two constituent cells, and since the 
supposed uredospores agree with the upper cell of the teleutospore, they are 
probably the same. 

No uredospores were found by me in the specimen kindly sent by 

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. VI., p. 853 (1895). 
Sydow, Mon. Ured. I., p. 607 (1903). 
Sacc. Syll. XIV., p. 341 (1899). 

II. Uredosori on both surfaces and margins of leaves, minute, rust- 
coloured, rounded or oval, bullate, gregarious or scattered, bursting 
through and surrounded by epidermis. 


Puccini a — Labiatae. 

Uredospores shortly elliptical, yellowish, finely echinulate, 20-23 
X 16-18 /i, average 21 x 16 yu. 

III. Teleutosori amphigenous, black, blister-like, largely confluent and 

distorting leaf, long covered by epidermis. 

Teleutospores with a few uredospores intermixed, elongated 
clavate, chestnut-brown, slightly or not at all constricted at septum, 
commonly truncate and much thickened at apex, 36-54 x 15-22 p 

average 42 x 18 


upper cell squarish or oblong or club-shaped, 

apex very much thickened (up to 15 /j), sometimes strongly beaked, 
generally shorter than lower cell, dark chestnut-brown; lower cell 
tapering towards base, lighter in colour than upper, of a yellowish 
or golden brown ; pedicel pale yellow, persistent, generally about 
18-20 fi long. 

X. Mesospores chestnut to pallid, rounded or beaked at apex and 


11-18 /Lt. 

hat fusiform to elongated elliptical, 25-36 x 

On leaves of Hypoxis glabella II . Br. 

Victoria — Ardmona, July, 1893 (Robinson). Burnley, Ruther- 

glen, &c, July-Oct. 

It differs from P. burchardiae 


the much 


and decidedly in the teleutospores, which in the latter are generally rounded 
at the apex or beaked, and distinctly constricted at the septum. 

(Plate V.,, Fig. 41.) 




64. Puccinia menthae Pers. 

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

Spermogonia either arranged 


groups or scattered, honey 

Aecidia hypophyllous, or frequently on stems which are much 


ely on purplish-red spots on leaves, seldom scattered 
pseudoperidia immersed, flat, opening irregularly and 
scarcely or irregularly torn. 

Aecidiospores subglobose, ellipsoid or polygonal, coarsely granular 


pale yellowish, 24-40 x 
II. Uredosori hypophv 


seated on pale spots 




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


subglobose, ellipsoid or obovate, echinulate, pale 

brown, generally three equatorial pores, 20-30 X 16-19/1 
III. Teleutosori similar, but blackish-brown in colour 


first intermixed with uredospores, ellipsoid to 

rounded at both ends, apical papilla pale or hyaline, broadly 

xpanded, hardly constricted at sept 

chestnut-brown, 24-32 x 
1 9-20 a ; pedicel hval 


ted, golden to 




19-22 fx, but commonly 24-25 X 
slender, generally surpassing length of 






Puccinia — Acanthaceae. 141 

X. Mesospores, occasional, similarly coloured, nearly globose, very 

slightly roughened, 20 p diam. 

On leaves of Mentha laxijiora Benth. 

Victoria — near Melton (Reader). 

On leaves of Mentha pulegium L. 

Victoria— Myrniong, Aug., 1904, II., III. (Brittlebank). 

The teleutospores are prominently warted. 

(Plate XXIX., Fig. 250.) 


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 /1 diam., or 28-34 X 18-23 p. 

III. Teleutospores intermixed with uredospores, dark chestnut-brown, 

oblong, with thickened wall and roughened surface, sometimes 
slightly constricted, rounded at both ends, not thickened at apex, 
36-46 X 28-36 /z; upper cell resembling lower, but usually a little 
larger ; pedicel generally lateral and even sometimes on a level 
with the septum, flexuous, hyaline, up to 60 /j, long and 7 ^ 

On living leaves of Ruellia australis Cav. 

New South Wales — Richmond River, June, 1896 (Musson). 

The teleutospores were not very plentiful in the specimens examined, but 
the sori containing them could be detected by their darker brown colour. 
The lateral pedicel to the teleutospore naturally suggests P. lateripes B. and 
Rav., but a closer inspection reveals important differences in the two kinds 
of spores. 

I have compared the Australian species with specimens on the leaves of 
R. strepens L. from N. America in Sydow's Ured. Exs., No. 1374, and 
Kellerman's Ohio Fungi, No. 130, and it is evident that we are dealing with 
similarity of type due to the affinity of the host-plants, with considerable 
divergence in the character of the spores. Unfortunately, P. lateripes B. 
and Rav. and P. ruelliae (JB. and Br.) Lagh. are confounded, for although 
Sydow labels his specimen as the former, in his Monograph he names it 
the latter. In this specimen the finely echinulate uredospores are ellipsoid 
to ovate, and measure 24-28 X 19-21 /i, while here they are larger. The 

teleutospores likewise only measure 29-34 x 18-22 p, and are very strongly 

This species differs from P. longiana Syd., in the larger uredospores and 
the teleutospores not being thickened at apex, and from P. lateripes and 
P. ruelliae in the larger size of uredo- and teleuto-spores. 

(Plate V., Figs. 43, 44.) 

!42 Piiccinia — Convolvulaceae. 



66. Puccinia dichondrae Mont. 

Montagne in Gray's Fl. Chil. VIII., p. 46 (1845). 
Cooke, Handb. Austr. Fung., p. 338 (1892). 
Sydow, Mon. Ured. L, p. 321 (1902), and p. 881 (1904). 
Sacc. Syll. VII., p. 717 (1888). 

P. dichondrae Berk. Linn. Journ. XIII., p. 173 (1872). 
P. berkeleyana De Toni, Sacc. Syll. VII., p. 717 (1888). 
P. munita Ludwig, Zeitsch. f. Pflanzenk. II., p. 133 (1892). 

III. Teleutosori hypophyllous, punctiform, very minute, erumpent to 

superficial, somewhat pulverulent, densely gregarious and often 
covering entire surface, deep cinnamon, nestling among hairs, 

70-100 fx diam. 

Teleutospores clavate to oblong, golden brown, constricted at 
septum, with comparatively thin epispore, thickened at apex and 
produced into a hyaline apiculus (occasionally two), with granular 
contents, and each cell 1-guttulate, occasionally 3 to 4 celled, 
30-40 x 12-18 p, average 32 x 14 ju ; pedicel hyaline, sometimes 
tinted, persistent, generally slender, up to 46 ^t long, occasionally 
6 fj. broad. 

X. Mesospores sub-ellipsoid to oblong, similarly coloured to teleutospore, 

thickened at apex and usually with hyaline apiculus, 21-30 X 
15-22 /u. 

On Dichondra repens Forst. 

Victoria — Near Melbourne, 1886 (Reader). Cheltenham, Nov. 

1887 (French). Goulburn River Flats, Oct., 1896 (Robin, 
son). Christmas Hills, May, 1900 (Robinson). Point Cook 
May, 1902 (French, jun.). Murramurrangbong Ranges, Nov. 
1902 (Robinson). Mt. Blackwood, Dec, 1902. Port Fairy, 
Aug., 1905. Mentone, throughout the year, 1905, etc. 

New South Wales— 1901 (Camfield). Richmond, April, 1905 

(Musson). Recorded by 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 /u) by 
Berkeley, which is evidently a misprint. Next, Dr. Cooke in his Handbook of 
Australian Fungi makes a correction by giving the size of the spores as 
12-13 fi long, which evidently errs on the other side. Then De Toni 
changed the name to P. berkeleyana, seeing that the size of the spores as 
given did not at all agree with those of P. dichondrae, Mont. 

Another unfortunate error has arisen in connexion with this species 
through the wrong naming of the host-plaut. Mr. Tepper sent a rust to 
Prof. Ludwig from S. Australia, said to be on the living leaves of Hydrocotyle 
hirta R. Br., who determined it as a new species, P. munita. But on Prof. 
Ludwig kindly sending me some of the original material it was found that 
the leaves belonged to Dichondra repens and that the rust was P. dichondrae, 
a conclusion with which Prof. Ludwig agreed after examining the specimens 
sent to him. 

Puccinia — A-pocynaceae. 143 

The teleutospores are frequently 3-4 celled, and there is great variety in 
the arrangement of the septa. The spore may be divided vertically as in 
D ior chidium, or it may have a vertical or oblique septum in its upper or 
lower cell. . The septa may even be arranged muriformly. There may be 
a lateral hyaline apiculus in the lower as well as in the upper cell, and the 
pedicel may stand out at right angles to the lower cell. 

There is thus every gradation from the unicellular spore and the bicell- 
ular, in which the upper cell is more or less atrophied, up to the multicellular, 
which is vertically, obliquely or muriformly divided. 

Aecidia have been found on this plant and are regarded as belonging to 
this species, but although numerous specimens have been examined from 
various localities no trace of aecidiospores has been found here. 

It is worthy of note that some of our native species of Viola closely 
resemble the Dichondra, and the one may easily be mistaken for the other 
when no flowers are present. 

(Plate V., Fig. 42 ; Plate XL., Fig. 299.) 


67. PuCCillia alyxiae Cooke and Mass. 

A lyxia 

Cooke and Massee, Grev. XVI., p. 2 (1887). 

Cooke, Handb. Austr. Fung., p. 338 (1892). 
Sydow, Mon. Ured. I., p. 336 (1902). 
Sacc. Syll. VII., p. 714 (1888). 

III. Teleutosori generally hypophyllous, occasionally epiphyllous, 

discoid, compact, dark-brown, girt by the ruptured epidermis, 
l-2mm. diam. 

Teleutospores almost pear-shaped or oblong, yellowish to 
brownish, constricted in the middle, thickened at apex and 
generally apiculate, sometimes rounded or truncate, occasionally 
3-celled, 40-52 x 20-25 /jl, average 45 x 20 /* ; epispore thick, 
smooth, coloured ; pedicels persistent, hyaline, broadish, elongated, 
up to 130 fA, occasionally at right angles to the spore. 

X. Mesospores not uncommon, similarly coloured, elongated oblong, 

thickened and sometimes apiculate at apex, 38-50 X 19-21 p. 

On leaves of Alyxia buxifolia R. Br. 

Victoria— Brighton and Broadford, Sept., 1887 (Miss Campbell 1 ). 

Beaumaris, March, 1895. Cheltenham, May, 1902 (C.French, 

Sandringham, April, 1905. (Robinson). 
South Australia — Gawler, Sept., 1893. (Tepper). 
Tasmania — Near George's Bay, Nov., 1892. (Rodway 1 ). 

The sori are generally isolated, much inflated and surrounded by a 
conspicuous, brown, elevated ridge. When on both surfaces they are 
opposite to each other. 

In the Cheltenham material, the spores had germinated on the leaves 
lying on the ground on 19th May, while the spores on the Gawler material 
found in September had not. 

This spore probably undergoes a period of rest during the summer 
before germination. Cooke and Massee have given the length of the spores 
as 50-70 n, but this is evidently a misprint. 

(Plate VI., Fig. 46.) 


Puccinia — Apocynaceae, Goodeniaceae. 


68. Puccinia carissae Cooke and Mass. 

Cooke and Massee, Grev. XXII., p. 37 (1893) 
Sydow, Mon. TJred. I., p. 336 (1902). 

Sacc. Syll. XL, p. 195 (1895). 
III. Teleutosori hypophyllous, gregarious, on 

orbicular spots, forming 

rings which are at length confluent, rather compact, dark-brown. 

Teleutospores elliptic, constricted at septum, rounded at ends or 
occasionally flattened, smooth, brown, 30-34 x 17-25 n, average 
32 x 22 


occasionally 3-celled, then reaching to a length of 

deciduous or persistent, sometimes origin- 

49 fx ; pedicel hyaline, 
ating laterally, rather long. 

X. Mesospores occasional, just resembling a teleutospore, but without 

the septum, elliptic, with hyaline pedicel, 29-31 x 22-25 jx. 

On living leaves of Carissa ovata R. Br. 

Queensland — Gladfield, (Gwyther) (Bailey 13 ). Dalby, (Bancroft) 



Mr. Bailey has kindly sent me some excellent material from his herbarium 

In the original 

and the teleutospores have been carefully measured, 
description the size is given as 20-22 x 16 ft, but probably 30-32 ll was in- 
tended for the length. 

Sometimes both upper and lower cell have an oblique division. 

It differs from P. aly 

Cooke and Mass., in which the spores are much 

and thickened at the apex 

(Plate VI., Fig. 45.) 



69. Puccinia brunoniae McAlp. 

McAlpine, Agr. Gaz. N.S.W. VI., p. 851 (1895). 
Sydow, Mon. Ured. I., p. 193 (1902). 
Sacc. Syll. XtV., p. 320 (1899). 

0. Spermogonia dark honey-coloured, disposed in clusters on yellowish 

spots, or intermixed with aecidia on the same surface of the leaf, 
usually the upper. 

1. Aecidia scattered on 

more often 

cularly arranged 



htly swollen indeterminate spots, mostly on upper surface of 

leaves and 

slightly recurved 

petioles ; pseudoperidia with wh 


d soon 


isolated peridial cells, generally short and 

mm. diam.; 
stout, less commonly 





punctate, with striated 

24-31 x 14-17 /i, or 

Aecidiospores subglobose 

smooth, 17-21 u diam. or 1 





when elongated 38 

45 x 21-24 ii 

or even 


1 x 




ionally as long 

Puccinia — Goodeniaceae 


III. Teleutosori amphigenous, but mostly on upper surface, very rarely 

mixed with aecidia, forming clusters of black, bullate pustules on 
indefinite, pale, ruddy, thin spots, oval or elongated, usually 
confluent, arranged circularly, or in lines, at first girt by 
ruptured epidermis, which finally falls away. 

Teleutospores clavate, golden-brown to chestnut, thickened at 
apex, constricted at septum, sometimes 3 or 4 celled, 45-60 x 
18-21 /j, average 50 x 20 fx ; upper cell generally pointed, but 
often rounded and occasionally truncate, deeper in colour than 
lower ; lower cell tapering towards base and generally longer than 
upper; pedicels persistent, pale-yellowish to hyaline, 40-50 x 



X. Mesospores occasional, similarly coloured to teleutospores, rather 

oblong, thickened at apex, rounded or truncate, slightly narrowed 
at base, 28-38 x 17-21 F . 

On leaves and petioles of Brunonia australis Sm. 

Victoria — Drysdale, Oct., 1895, III. Murramurrangbong Ranges. 

Nov., 1902, Dec, 1903, O., I., III., and Jan., 1905, III, 
(Robinson). Alps, near Bright, Dec, 1904, III. (C. French, 
jun.). Rutherglen, Dec, 1904, III. 

Three-celled teleutospores occasionally seen, varying in length from 
60-73 fi and in breadth from 21-24 ju, the lower cell generally about 
as long as the other two. 

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

At first no aecidia were known, but these were found along with the 

teleutospores by G. H. Robinson. 

The discovery of aecidia rendered it necessary to carefully compare the 
two fungi found upon the Goodeniaceous plants, Brunonia and Goodenia, 
and a large number of specimens and slides were accordingly examined and 

While the rusts are of the same general type, they differ in several 
important respects. The aecidiospores in P. saccardoi are considerably 
smaller, and the peridial cells are generally much more elongated, and more 
than twice as long as broad. 

In the original description by Dr. Ludwig the size of the aecidiospores 
is given as 13-15 jjl diam., and the peridial cells as 18-25 x 15-18 ju, but 

if the latter are measured when isolated they are found to be much longer. 
Again in P. saccardoi the teleutospores are generally intermixed with or 
surround the aecidia, but this very rarely occurs in P. brunoniae. In the 
former the teleutospores are generally rounded at apex, but in the latter 
generally pointed and altogether narrower. 

In the Murramurrangbong Ranges where the aecidia and teleutospores 
of P. brunoniae were found very plentifully in November, though 
Goodeniaceous plants which, in other districts are affected by P. saccardoi y 
were exceedingly common and were often growing alongside the Brunonia, 
no rusts were found on them, even although the country was subjected to a 
most exhaustive search. The rusts, therefore, on the two different genera 
are considered to be specifically distinct. 

(Plate VI., Fig. 50.) 

146 Puccinia — Goodeniaceae 

70. Puccinia dampierae Syd. 

Sydow, Mon. Ured. I., p. 193 (1902). 
Sacc. Syll. XVIL, p. 315 (1905). 

I. Aecidia on stems and both surfaces of leaf, gregarious, bright 

orange ; pseudoperidia with white reflexed and torn margins, i-^ mm. 
in diam. ; peridial cells lozenge-shaped to oblong, with thick striated 
margins, 30-40 jx long. 

Aecidiospores orange, subglobose, ellipsoid to oblong, finely 
echinulate, 15-17 /x diam. or 17-19 X 13-14 ^u. 

III. Teleutosori on stems, scattered or aggregated, round or oblong, 

about 1 mm. diam., compact, dark-brown, girt by the ruptured 

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

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

On stem and leaves of Dampiera stricta K. Br. 

Victoria— Monbulk, Dec, 1905, I. (C. French, jr.). 

On wings of the stems of Dampiera alata Lindl. 

West Australia, III. (L. Preiss). 

Though I have not seen the West Australian specimens I prefer to 
regard the aecidia found in Victoria as a stage of the same fungus. If they 
should prove to be unrelated they would be easily separable at any time. 

71. Puccinia gilgiana P. Henn. 


Hennings, Hedw. XL., p. (95) (1901). 
Sydow, Mon. Ured. I., p. 194 (1902). 
Sacc. Syll. XVIL, p. 314 (1905). 

Aecidium perkinsiae P. Henn. Hedw., XL., p. (96) (1901). 

I. Aecidia on thickened and slightly deformed parts, gregarious or 

scattered ; pseudoperidia hemispherical to cup-shaped, semi- 
immersed, yellowish to ruddy ; peridial cells round or oblong- 
polygonal, subhyaline, reticulate, 18-24 X 16-20 li. 

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

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

almost covered by the ruptured brown epidermis, somewhat 

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

III. Teleutosori similar, but black. 

Teleutospores clavate or oblong, constricted at septum, smooth, 
obtusely rounded or truncate at apex and slightly thickened (up 
to 5 li), obliquely papillate and occasionally crested, rounded at 
base or attenuated, brown, 30-45 X 16-22 li, average 42 X 20 ft; 
pedicel brownish, thick, persistent, 30-50 a long. 

Puccinia — Goodeni 



X. Mesospores very common, oblong or oblong-clavate, thickened at 

apex, rounded or acute, chestnut-brown, smooth, stalked 23-32 
X 15-20/1. 

On petioles, calyx and stems of Leschenaultia linarioides DC. 

West Australia— Near Perth, 1900 (Pritzel). 
Aecidia on petioles, flower-stalks, and calyx ; uredo and teleutosori on 


Specimens kindly supplied by Hennings. 

(Plate VI.,Fig. 47.)' 

/ 2. Puccinia saccardoi Ludw. 

Goodenia, Velleia 



Ludwig, Hedw. XXVIIL, p. 362 (1889). 
Cooke, Handb. Austr. Fung., p. 337 (1892). 
Sydow, Mon. Ured. I., p. 193 (1902). 
Sacc. Syll. IX., p. 309 (1891). 

P. nigricaulis McAlp., Agr. Gaz. N.S.W. VII., p. 151 (1896). 
Aecidium g oodeniacearum (in part) Berkeley, Linn. Journ. 

XIII., p. 173 (1872). 

I. Aecidia on yellowish or brownish spots, scattered or in groups 
I arranged in circular patches (up to 6mm. diam.), amphigenous 

causing distortion of stem ; pseudoperidia cup-shaped, with white 
torn, everted edges, 215-325 /jl, diam. ; isolated peridial cells 
punctate, with striated margin, generally elongated oblong, or 
somewhat oval or lozenge-shaped, more than twice as long as 
broad, 35-42 X 16-21 ^. 

Aecidiospores subglobose, polygonal to oval, orange, very finely 

verrucose, 14-16 /i, diam., or 15-17 x 13-14 ^u. 

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

Teleutospores clavate to oblong, dark-brown, constricted at 
septum, smooth, variable in size, rarely 3-celled, 40-66 x 17-25//, 
occasionally up to 70 jx long, average, 54 x 20 // ; upper cell 
subglobose or somewhat quadrate, thickened at apex (up to 9 /u), 

rally rounded or truncate, occasionally pointed, 21-30 x 

17-25 /i, sometimes reaching 32 fi in length; lower cell generally 
paler in colour than upper, elongated and tapering towards base, 
sometimes subglobose, longer and narrower than upper, 17-35 X 
16-21 fi, occasionally 38 /t long; pedicel usually persistent, 
hyaline, occasionally pale-yellow, 35-60 X 7-12 /<. 

X. Mesospores common, similarly coloured to teleutospores or paler, 

variously shaped, ellipsoid to oval, or oblong to clavate, smooth, 
with pointed and thickened apex (up to 9 /*), and generally 

slightly tapering towards base, 25-49 x 12-21 /*, or averagin 
36 x 17 fx 



Puccinia — Campanulaceae . 

On stems, leaves and calyx of Goodenia geniculates It. Br., G. glauea 
F. v. M., G. pinnatifida Schlecht., G. albiflora Schlecht., G. hederacea 
Sm., and G. ovata Sm. 

Victoria— Port Phillip, 1886, I. Dimboola, Sept., 1892, I., III. 

(Reader). Minyip, near Colac, Grampians, Warracknabeal, 
Nhill, Killara, Mt. St. Bernard, Borung, Gembrook, near 


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

Creek, near Quorn, Sept., 1892 (Molineux). 


Tasmania — Cascades, Jan., May, Nov., 1892 and 1893, 

(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 pinnatifida, received from 
F. M. Reader, Dimboola, the fungus is' named A. goodeniacearum Berk, in 
Massee's handwriting, while the teleutospores of P. saccardoi are also met 
with intermixed with the aecidia. 

In Velleia paradoxa, from New South Wales, while the teleutospores 
are of the general type, there are quite a number in which the apex is 
truncate and prolonged laterally into a thickened, more or less beak-like 




observed in the 


type on Goodenia 
the spores on the genera Goodenia and 

geniculata a similar variety, so that 
Velleia cannot be differentiated. He writes — " Mixed among the normal 
teleutospores there are three and one-celled spores, the latter sometimes of 
abnormal size, as I have observed under similar conditions in P. heterospora. 
Frequently very peculiar forms appeared, in which the upper cell bore 1-3 
horn-like projections as long as the cell itself, or finger-like cells, as long as 
the upper, proceeded from the lower cell." 

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


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

Lobelia, Pratia, 

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. I., 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 /jl. 

1. Aecidia amphigenous, rounded or ovate, scattered or densely 

gregarious and occupying the entire surface of the leaves or 
petioles, at first covered by the epidermis, then free and rupturing 
like a broken blister, vesicular, ochraceous ; pseudoperidial cells, 
oval to oblong, covered with spines, loosely connected, 30-40 x 
18-25 11. 

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

III. Teleutosori on under surface, numerous, bullate, then girt 

ruptured epidermis so that they resemble little Pezizae. 

Teleutospores cylindric to sub-clavate, elongated, yellow, 
thickened apex (9-10 /i), rounded or truncate, deeply constricted 
at septum 56-70 x 15-26 /u, average 60 X 24, occasionally bi- 
and tri- septate when they may reach a length of 80 \x ; upper cell 

rally broader th 

rally cylindrical, and 

may taper slightly towards pedicel ; pedicel hyaline, short, stout, 
7-13 fi broad. 

Aecidiospores on Lobelia anceps L., L. pratioides Benth., L . jjurpurascens 
R. Br., Pratia erecta Gaudich., P. pedunculata Benth., P. platycalyx Benth. 

Victoria — Colac (F. v. Mueller), (von Thuemen). Murtoa, Oct., 

1892, (Reader). Ringwood, Oct., 1892 (Robinson). Minyip, 
Oct., 1901 (Eckert). Werribee Plains, June, 1902 (C. 
French, jun.). 

New South Wales — Guntawang, Sept., 1886 (Hamilton). Kurra- 

jong Heights, Apr., 1894 (Musson). 

Tasmania — Bellerive Swamp, Dec, 1891 (Rod way). 
Teleutospores on leaves of Lobelia anceps L. 

South Australia— Port Lincoln, Nov., 1852 (C. Wilhelm 



The aecidium was first determined by Thuemen on a specimen of 

Pratia platycalyx sent by the late Baron von Mueller from Colac, and named 
A. lobeliae. Then Berkeley received a specimen from the same source on 
P. pedunculata which he named A, microstomum. 

I have examined portions of the original material of both these 
specimens, and as a rule it is difficult to detect a peredial wall even in fine 
sections, so much so that they were as first taken for caeomata. But there is 
occasionally a loosely fitting layer of surrounding cells, so that we may 
regard them as aecidia. The aecidial stage is given in connexion with the 
teleuto stage since, although not found actually together, they occur separately 
on the same species, Lobelia anceps. 

The teleutospore is occasionally tri-septate, and is then slightly curved, 

and the lower septum oblique. 

The description of the' teleuto stage is drawn up from original specimens 
the National Herbarium, named in Berkeley's handwriting. 


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


Puccinia — Comfo sitae 




74. Puccinia angustifoliae McAlp. 

I. Aecidia in dense clusters on the 

leaves, more sparingly c 
about J to nearly J mm. diam. ; pseudoperidia with finely 
everted margins. 


quadrate, 16 X 


yellow, globose to elliptical or sub 

3 u or 13-14 a diam 

III. Teleutosc 

thro win 

on leaf and stem, black, long covered by epiderm 





pidermis, d 

patches, partially ruptu] 
:t from or along with 


clavate to oblong, smoky-bro 




stricted at septum, smooth, generally rounded and thickened 



htly tapering towards base or often 
average 50 X 20 // ; pedicel hyaline, 

l (up to 9 /x), s] 
rounded, 40-53 x 16-S 
deciduous, generally short, up to 38 fx long and 9 ll broad 


X. Mesosp 

common, similarly colored to teleutosp 


generally somewhat oval, thickened at ape 
On Scorzonera anguslifolia L. 

25-30 x 16 

17 a 

Victoria— Dimboola, Nov., 1892, L, III. (Reader). 


species differs from P. podospermi 




(Schum.) Jacky and P. tragopogi (Pers.) Corda, in the absence of 
uredospores and the smooth teleutospores. 

(Plate VII., Fig. 53.) 


75. Puccinia brachycomes McAlp. 

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 
pseudoperidial cells firmly united, oblong to sub-quadrate, punctulate 
all over, with broad striated margin. 

Aecidiospores yellowish, sub~globose to ellipsoid, finely echin- 
ulate, 13-16 fi diam. or 13-16 x 10-13 /jl. 

II. Teleutosori elliptical, pustulate, black to blackish-brown, 


covered by epidermis, crowded, confluent, and ultimately forming 
elongated patches several mm. in length. 

Teleutospores clavate, dark chestnut-brown, smooth, very 

slightly constricted at septum, 48-64 


18-25 n 


50 x 20 n ; upper cell truncate or conoid, occasionally rounded at 
apex, thickened (up to 9 ju) ; lower cell generally attenuated towards 
base, sometimes rounded and generally longer than upper ; pedicels 
persistent, tinted or hyaline, up to 35 fx long. 

X. Mesospores not uncommon, similarly coloured to teleutospores, 

ellipsoid, either rounded or pointed, and thickened at apex, 
28-30 x 14-19 p. 

On Brachycome ciliaris Less., I., III., and B. pachyptera Turcz., I. 

Victoria— Dimboola, May-July, 1896 and 1898 (Reader). 

On Brachycome scapiformis DC. and B. diversifolia, Fischer and Mey. 

Victoria— Buffalo Ranges, Dec, 1904, I., III. (C. French, jun.). 
Darluca ftlurn Cast., is a very common parasite on the aecidia. 

(Plate VIII., Fig. 66.) 

Puccinia — Compositae. 

I 5 I 

76. Puccinia calendulae McAlp 



McAlpine, Proc. Linn. Soc. N.S.W. XXVIII., p. 558 (1903). 
Sydow, Mon. Ured. I., p. 852 (1904). 
Sacc. Syll. XVII., p. 280 (1905). 

Aecidium calendulae McAlp. Agr. Gaz. N.S.W. VII., 

p. 152 (1896). 

I. Aecidia amphigenous or on stems, orange-yellow, in clusters up to 

6 mm. diam, sometimes circinate, or may be scattered irregularly ; 
pseudoperidia with white, torn and reflexed margin ; peridial cells 
quadrate or polygonal, striated at margin, 21-24 p long. 

Aecidiospores very irregular, sub-globose to polygonal, very finely 
echinulate, pale orange, 14-17 X 11-12 ll or 12-16 ll diam. 

III. Teleutosori intermixed with aecidia, black, erumpent, soon naked, 

girt by the ruptured epidermis, circular to elliptical, minute, 

compact, often confluent. 

Teleutospores yellowish-brown, clavate to oblong clavate, con- 
stricted at septum, rounded or acute at apex and thickened (up to 
12 fi), attenuated towards base, smooth, 36-52 x 19-23 /<, 
average 48 x 20 ll ; upper cell darker in colour and broader than 
lower, 21-31 li long; lower cell slightly or not at all tapering 
towards pedicel and averaging same length as upper ; pedicel 
persistent, hyaline, but coloured towards apex, 28-37 x 7-8 ^/, 
but may attain a breadth of 10 ll at junction with spore. 

X. Mesospores not uncommon, similarly coloured to teleutospores, 

ovate to elliptical or pear-shaped, thickened at apex, 30-42 x 
21-23 ll. 

On leaves and all green parts, including fruits of Calendula officinalis L. 

Victoria — Near Melbourne, growing in gardens, 1892, etc. Killara, 

July-Oct., 1902. Geelong, Nov., 1904, J., 111. (Pescott). 
Frankston, August, 1904, I. 

Aecidial stage all the year round, but less common in mid-summer, 
persisting only in shady gardens. Teleutostage from March to November. 

The aecidial stage was the only one found at first, and was described in 
the Agricultural Gazetti of New South Wales for 1896 at p. 152. Then 

in March, 1902, the teleutostage was found by Mr. G. H. Robinson, and 
was very plentiful. Both stages very common in University Gardens, 
Melbourne, in autumn and winter of 1905. Mesospores germinating at apex 
like teleutospores, and producing sporidiola. 

It causes swelling, discoloration and distortion of the flower-stems and 
leaves, and the bright orange colour of the aecidia on the leaves at once 
attracts attention from its harmonising with the flower-head. 

(Plates VIIL, Fig. 65 ; XLIIL, Fig. 312 ; Plate E., Figs. 22, 23.) 


77. Puccinia calocephali McAlp. 


I. Aecidia very sparse on upper surface of hairy leaves, about J mm. 

diam.; pseudoperidia saucer-shaped, leciniate at margin; peridial 

cells, oblong to lozenge-shaped, striated, 25-32 ll long. 



J 5 2 

Puccinia — Com-positae. 


to angular, orange-yellow, smooth, 
up to 4 germ-pores on one face, 16—19 X 12-13 jx or 

Aecidiospores sub-globose 

14-16 p diani. 
III. Teleutosori black to 

blackish-brown, long covered 

by leaden 

epidermis, pulvinate, compact, round to elongated, sometimes 

confluent, 1-1 h mm. long. 


Teleutospores clavate, dark brown, particularly in upper cell, 

slightly constricted at septum, smooth, 40-57 X 17-24 
45 X 22 

ju, average 

fi ; upper cell rounded, scoop-shaped, sometimes truncate 
and thickened at apex (up to 1 1 p) : lower cell yellowish brown, 
attenuated towards base ; pedicels persistent, tinted to hyaline, 
up to 35 fx long and sometimes 10 fi broad. 

X. Mesospores common, elongated, elliptical to oblong, generally paler 

than teleutospore, thickened at apex, and generally scoop-shaped, 
28-35 x 13-16 /x. 


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


Victoria — Dimboola, Sept., 1891, III. (Reader). Phillip Island, 

Nov., 1901, I. (C. French, jun.) 

The teleutosori and teleutospores generally resemble those of 1\ 
tasmanica Diet., but they are not intermixed with aecidia. They both be- 
long, however, to the same general type. 

(Plate VII., Fig. 58.) 


78. Puccinia calotidis McAlp. 



Aecidia amphigenous, in irregular clusters, orange ; pseudoperidia 

cup-shaped, with whitish, reiiexed 


Aecidiospores subglobose to polygonal, finely echinulate, 13-14/i 

III. Tele 

15 x 12 p. 

minute, black 

umpent, gregarious, compact 

Teleutospores dark chestnut-brown, oblong to elongated obi 
ivate, smooth, constricted at septum, rounded or bluntly pointed 

to 9 ft), 37-50 X 18-28 H , average 

and thickened at 



38-40 x 18-20 fi; pedicel hy 

generally tap 

X. Me 

wards base, about 30 fi long, or shorter than spore 

ly coloured to teleutosp 


ed, obovoid to ellipsoid, rounded or bluntly pointed and 

thickened at apex (6-8 


x 1 






On Calotis cuneifolia R. Br. and Calotis sp. 

New South Wales — Guntawang, 

Condoblin, Aug., 1897 (Maiden). 

The teleutospores generally resemble those of P. vittadiniae, but they 


(Plate XLIIL, Fig, 310.) 

Puccinia — Compo sitae 

J 53 

79 # Puccinia chrysanthemi Roze. 


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

. P. chrysanthemi chinensis P. Henn. Hedw. XL., p. (26) (1901). 

II. Uredosori on both surfaces of leaf, but mostly hypophyllous, 

sparingly on stem, generally round, soon naked, powdery, scattered 
or crowded, often confluent, sometimes arranged circularly, snuff 
brown, average 1-1 £ nim. diam., causing dirty-brown, indefinite 
spots on upper surface of leaf, which gradually extend and 
ultimately decay. 

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

[III. Teleutosori on both surfaces of leaf, but chiefly on under surface, 

prominent, roundish, pulvinate, solitary or confluent in groups, often 
concentrically arranged, dark-brown to black, soon naked, compact. 
Teleutospores at first sparingly intermixed with uredospores, 
ellipsoid to pear- shaped or clavate, rounded or occasionally 
truncate at apex, and thickened (up to 9 /u), mostly rounded at base 
or sometimes slightly attenuated, chestnut-brown, with strong, 
finely punctulate epispore, 35-60 x 19-28 n; pedicel hyaline, stout, 
persistent, 40-80 x 7-9 ^u.] 

[X. Mesospores rare, ellipsoid to clavate, similarly coloured to teleuto- 
spores, rounded and thickened at apex, finely punctulate, 28-46 X 
22-25 ft.] 

On cultivated Chrysanthemums (Chrysanthemum indicum L.). 

New South Wales — Sydney Botanic Gardens, April, 1904 

(Maiden). Sydney, May, 1904. 

This species has been assigned to P. hieracii by Massee , but Jacky 

12 3 

» > 

has shown, by means of infection experiments, that it confines itself 
to the one genus, and does not infect other Compositae. He has also experi- 
mented with the Japanese chrysanthemum rust, P. (hrysanthemi-chinensis P. 
Henn., and proved it to be the same species as the other. 

Through the kindness of Professor Ideta, of the Agricultural College, 
Osaka, Japan, I have received abundant material of the Japanese rust, both 
in its uredo and teleuto stages, and am thereby enabled to compare it with 
the rust occurring in Australia, as far as the uredo stage is concerned. The 
uredospore3 are quite similar, having three equatorial germ-pores on one 
face, and measuring 28-37 X 22-25 /w. Jacky 3 succeeded in infecting 

C. indicum L., with material from C. sinense Sabine, and thus showed 
identity of the two rusts, although apparently in Europe and Australia it is 
gradually losing the capacity to form teleutospores. Indeed, Miss Gibson ' 
goes so far as to say that there is no necessity for resting-spores, for the 
young shoots are above ground long before the old ones die away, and th 
young shoots are taken as cuttings, 

time when there are 

pon which 

F 2 


Puccinia — Compositae. 

The sams author also records the fact that there are certain varieties of 
chrysanthemum which do not take the rust, although growing among plants 
badly rusted, but Jacky 3 tested a number of different varieties and found 
them equally predisposed to the rust. 

Our chrysanthemum growers sometimes express a fear that the rust seen 
on hollyhock, sunflower, and other plants may pass to the chrysanthemum 
when it has suddenly bscome temporarily weak through drought or other 
causes ; but it ought to reassure them to know that the specific rust, known 
as Puccmii chrysanthemi, is alone capable of producing infection on that 
particular host. 

In the European material Jacky detected what he considers to be two- 
celled uredospores, but Sydow regards them as two unicellular spores stuck 
together. It may b3 mentioned here to show the wonderful variety, that in 
the Japanese material I found two uredospores borne on the same stalk, the 
one slightly beneath the other. 

The teleutospore is sometimss described as smooth, but 

careful focus- 

sing it may be seen to be finely punctulated all over in the young stage, 
although in the mature and deeply-coloured spore this is difficult to see, only 
being visible on the thickened apex. 

The mesospores were obtained from the Japanese material, and are 
rather rare. 

This rust was probably introduced into Europe from Japan, and it may 
have reached Australia either from Europe or Japan, 
rust has not yet found its way into Victoria, considering the number of 
chrysanthemums imported from England, France, America, and Japan. 

It was first observed in England in 1895, although it probably existed 
there before that time ; in America in 1896 ; in France and Italy in 1897 ; 
and about 1898 it spread to Germany, then to Denmark and Switzerland, 

to Australia. In 1904 it was also 

It is strange that the 


at length it has found its 


recorded for New Zealand by Kirk. 

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

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

80. Puccinia cichorii (DC.) Bell. 


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

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

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

If. Uredosori amphigenous or on stem, minute, pul 



fluent, girt 

or partially covered by ruptured epidermis 


Uredospores globose, 



ellipsoid, echinulate, 

yellowish-brown, with one or two germ-pores on one face, sometimes 

three to four, 22-29 x 19-23 fi 
III. Teleutosori similar, dark-brown. 


or 22-25 f± diam. 

Teleutospores intermixed sparingly with uredospores, ellipsoid 
or ovate-ellipsoid, brown to golden-brown, smooth, not or hardly 
constricted at septum, apex rounded not thickened, base for the 


27-38 X 19-25 /t, occasionally reaching 50 \x long, but averaging 
35 x 20 p. ; pedicel hyaline, short. 

most part rounded or very slightly attenuated, epispore 
germ-pore in each cell prominent, with finely 


Pvccinia — Compositae. 155 

On leaves and stems of Cichorium intybus L. 

Victoria — 1885 (Mueller 3 ) II. Near Melbourne, Aug., 1896. 

Phillip Island, Jan., 1900; March, 1904. 

The uredospores were common chiefly on the leaves, but the teleutospores 
vyere rather scarce, and occurred mostly on the midribs. 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 fx long, and the upper colourless one only 6 li. 

Darluca filum Cast., was plentiful on the uredosori and the sori 


uredo and teleutospores intermixed 

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



81. Puccinia cinerariae McAlp. 

I. Aecidia on both surfaces of the leaf, running along veins and gre- 

garious, causing depressions on upper surface and swelling of 
the veins on under surface, ^ mm. diam. ; pseudoperidia very 
slightly projecting, with margins white, everted, fimbriate ; peri- 
dial cells roughly quadrangular, often rounded at one end, longer 
than broad, with radiating marginal lines, 22-26 X 19-21 fi. 

Aecidiospores bright orange, subglobose to polygonal, very 
finely verrucose, 15-19 li diam. or 15-17 x 13 /*. 

III. Teleutosori intermixed with aecidia, black, comparatively rare. 

Teleutospores clavate, chestnut-brown, generally rounded and 
much thickened at apex or bluntly pointed, occasionally truncate, 
constricted at septum, 36-54 x 17-24 la, average 40 x 23 fx ; 
upper cell usually darker in colour, hemispherical, apex thickened 
to a depth of 9 li, 17-24 li long; lower cell gradually tapering 
towards base, somewhat triangular, 1 8-30 li long ; pedicel per- 
sistent, tinted, up to 45 li long. 

X. Mesospores chestnut-brown, obovate to elongated elliptical, much 

thickened at apex, with long pedicel, 27-36 x 13-23 fx. 

On leaves of cultivated Cineraria. 

Victoria — In nursery near Melbourne, April, 1899 (Cronin). 

The following note accompanied the specimen : — The plants were grown 

under glass, and were generally very healthy. There are not many leaves 

••so affected owing to the care that is taken to cut out the spots as soon as 

I noticed. The disease spreads rapidly if neglected, and completely spoils a 

j plant by denuding it of its foliage. 

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

(Plate VIII., Fig. 60.) 


i 5 6 

Puccini a — Compo sitae. 


82. Puccinia cyani (Sehleich.) P 

Passerini in Rabh. Fung. Eur., N 



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

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

II. Uredosori mostly hypophyll 

ed or 

ded, minute 

orbicular or elliptic, pulverulent, reddish br 


yellowish b 


lobose, subglobose to 


finely cchinulate, with two 


on one face 


7 x 




III. Teleut 


and on stem, scattered, pulverulent 

dark-brown, orbicular to elliptic, soon naked 


1 mm 







ellipsoid, rounded at both end 
stricted at septum, delicately 
30-40 x 22-30 /i, average 32 x 
pedicel hyaline, short, 

thickened at apex, not 


erm-pores conspicuou 


24 fx ; epispore about 3 /x thick 

On leaves and stems of Centaur ecu cyanus L. 

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

This is evidently of the Puccinia hieracii type. 

ular and well defined, and there is no 


teleutospores are very reg 

tendency to 'abnormality. 

(Plate VIII., Fig. 64.) 


83. Puccinia distincta McAlp. 

McAlpine, Agr. Gaz. N.S.W. VI., p. 853 (1895). 
Sydow, Mon. Ured. I., p. 29 (1902), and p. 850 (1904) 

Sacc. Syll. XIV., p. 311 (1899). 
I. Aecidia seated on discoloured spots, amphigenous, 

arranged in a 

circinate manner, or irregularly disposed, and often distributed 
over entire surface ; pseudoperidia cup-shaped, with reflexed torn 

mm. diam. ; peridial cells elongated elliptical to 



l i 


oblong, punctulate all over, 25-29 x 13-19 \i 



to oval or ellipsoid, orange-yellow, 

very finely echinulate, 14-19 p diam , or 14-20 x 13-16 \x 

III. Teleutosori intermixed with and often 

brownish black, oval, ^-1 



mm. long, generally confluent, bursting 

through and surrounded by the leaden-coloured epidermis. 

Teleutospores chestnut-brown, clavate or 

oblong clavate, at- 

tenuated at base, smooth, constricted at septum, 34-50 X 

15-21 H 


42 x 17 fi ; upper cell generally darker in 

colour than lower, rounded and thickened at apex (up to 9 ft) ; 
pedicel persistent, pale yellow or coloured similarly to lower cell, up 

to 50 /x long. 
X. Mesospores numerous, elongated 

to somewhat ovate, chest 

brown, usually thickened at apex, 34-37 X 




On leaves, scapes, involucre and petals of Bellis perennis L 


oria— Near Melbourne, Oct., 1892, July-Oct., 1904, I., Ill 
(Robinson). Korumburra, May, 1903, I. 

Puccinia — Compositae 


Tasmania — Near Huon Road, Nov., 1890, I. (Rodway). 
South Australia — Norwood, April, 1903, I., III. (Tepper). Port 

Pirie, Oct., 1903, J., III. (Dickens). 

Aecidiospores abundant from April to September, and even to January. 
The aecidia may even occur on the petals, and they have been described 
before the Microscopical Society of Victoria as early as 1880. 

Teleutospores in October or November, although they occur sparingly as 
early as April, The spots are generally pale-yellowish to pale greenish in 
colour. This aecidium from Victoria was determined by Cooke as A. bellidis 
Thuem., which belongs to P. obscura Schroet., but now that the teleutospores 
belonging to it have' been found on the same host-plant the Puccinia is seen 
to be a distinct species. 

(Plate VIIL, Fig. 67 ; Plate E., Fig. 26.) 



S4. Puccinia erechtitis McAlp. 

McAlpine, Proc. Roy. Soc. Vic. VII., N.S., p. 216 (1894). 
Sydow, Mon. Ured. I., p. 78 (1902). 
Sacc. Syll. XIV., p. 309 (1899). 

0. Spermogonia on brownish patches on upper surface of leaf, in groups, 

brown to honey-colour, yellowish by transmitted light. 

Spermatia hyaline, oval, 3 \x long. 

1. Aecidia on stem and leaves, causing distortion and swelling, pale- 
yellow at first, becoming orange-yellow, arranged close together in 
lines or irregularly ; pseudoperidia cup-shaped, with white, torn, 
revolute edges ; peridial cells firmly united, warted all over, 
lozenge-shaped, and nearly iso-diametric, or elongated and oblong, 
with striated margins. 

Aecidiospores variable in shape, irregularly globose or elliptic 

orange-yellow, smooth, 16-19 x 12-1/ jx. 

III. Teleutosori long covered by epidermis, crowded together, 

pulvinate, black. 

Teleutospores clavate to oblong, yellowish-brown, constricted 
at septum, smooth, 41-57 x 17-25 li, average 47 x 19 li ; upper 
cell dark-brown, rounded or pointed, occasionally truncate and 
thickened at apex ; lower cell usually yellowish-brown and tapering 
towards base, elongated wedge-shaped ; pedicel persistent, pale- 
yellow or hyaline, up to 40 li long and 9 li broad. 

X. Mesospores common, coloured like teleutospores, elongated oblong 

or oval, thickened at apex, 32-43 X 13-17 yu. 
On leaves and stems of Erechtites quadridentata DC, and E. arguta DC. 

Victoria — Ardmona, Dimboola, Myrniong, and Marysville, Alps 

near Bright, and Mt. St. Bernard, etc. 

On E. quadridentata, E. arguta and E. prendnthoides DC. 

Tasmania— Huon Road, Dec, 1891, March, 1893, I. (Rodway). 


On Erechtites sp. 

New South Wales — Guntawang, I. (Hamilton). 

I I. Very common all the year round, except during middle of summer. 

| Til. From January to June, but not so common. 

The New South Wales specimen was named Aecidium senecionis Desm. 

by Cooke. 

(Plates VII., Fig. 59 ; XXVII., Fig. 240.) 


Puccinia — Compo sitae. 

85. Puccinia gnaphalii (Speg.) P. Henn 

Hennings, Hedw. XLI., p. (66) (1902). 
Morrison, Victorian Nat. XI., p. 120 (1894). . 
Sydow, Mon. Ured. I., p. 88 (1902). 
Sacc. Syll. XVI., p. 295 (1902). 

Uredo gnaphalii Speg. Fung. Arg. IV., p. 28 (1882) 


Puccinia gnaphaliicola P. Henn. Hedw. XXXVIII., p. (68) 


II. Uredo-sori common on stem and leaves, on the latt< 

but mostly on under surface, bursting through and 

epidermis, also pier 

lly torn en turn 

l* amphigenous,. 
surrounded by 
ed, sometimes 

gated, compact, cinnamon, circular or oval, lenticular on 


-\ mm. diam. 
Uredospores globose 

sub globose 


echinulate, 21-24 a diam. or 21-26 x 17 





III. Teleutosori similar, but chestnut-brown to black 


at first 

ed with uredospores, oblon 



oblong-clavate, dark chestnut-brown, slightly constricted at septum 

rounded or obtusely pointed or even beaked at apex 


thickened (up to 8 /*), 
smooth, 35-53 x 15-24 

lly slightly attenuated towards base 

persistent, up to 66 




ge 48 

x 20 


pedicel hyal 



X. Mesospores similarly coloured, few, oval or elongated, elliptical^ 

thickened at apex and sometimes beaked, 24-38 x 15—18 yu. 
On stems and leaves of Gnaphalium purpureum L., and G. japonicum 


Oakleigh, Jan., 1904 (Morrison). Drysdale, Jan., 1896,. 
and Oct., 1903. Whittlesea Ranges, Nov., 1898 (C. French,, 
jun.) Killara, Oct., 1902. Near Melbourne, Feb. and Nov.,. 
1904. Murramurrangbong Ranges, Nov., 1904 (Aitken). 

The teleutospores are rather variable in length and breadth, sometimes 

being short and stout and sometimes rather elongated and slender. 



pedicels average 40-50 /j, in length, and may vary in breadth from 4 /j. up to 
10 /* at junction with spore. Sydow gives size of teleutospores as 32-56 x 
18-24^, and Dietel as 30-35 x 19-22 ju. 


I am indebted to Dr. Morrison for some of the original material 
which he determined this fungus as P. investita Schwein. He found sper- 
mogones immersed among spores in the sori, and the spermatia were fusi- 
form. Plenty of Darluca filum Cast., was found, and no doubt these were 

mistaken for spermogonia. 

(Plate VII., Fig. 57.) 

86. Puccinia helianthi Schwein. 


Schweinitz, Syn. Carol., p. 73 (1821). 

Cooke, Handb. Austr. Fung., p. 333 (1892). 
Sydow, Mon. Ured. I., p. 92 (1902). 
Sacc. Syll. VII., p. 603 (1888). 

Spermogonia honey- colored, in small clusters.] I 

[I. Aecidia crowded or orbicular, or arranged in broadly expanded,, 

oblong \spots ; pseudoperidia cylindrical, plane, with white laciniatft 


Puccinia — Compositae. 150 

Aecidiospores orange-red, ellipsoid to polygonal, finely echin- 

ulate, 21-28 x 18-21/*.] 

II. Uredosori roundish, chestnut-brown, scattered or confluent, pul- 

verulent, often on yellow or pale green spots on upper surface of 
leaf but generally forming brown mass on under surface. 

Uredospores sub-globose, elliptic or obovate, golden-yellow, echin- 
ulate, epispore sub-hyaline, showing one germ-pore on one face. 
21-24 yu diam. or 24-29 x 15-22 F . 

III. Teleutosori roundish, dark -brown or black, prominent, scattered at 

first but ultimately in clusters, confluent, dotting both surfaces of 
leaf, but most prominent on under. 

Teleutospores at first intermixed with uredospores, chestnut- 
brown, oblong-elliptical or pear-shaped, smooth, but occasionally a 
little rough at apex, slightly constricted at septum, thickened at 

apex (6-9 /*), generally rounded at base, 36-50 x 21-27 yu, 
average 44 x 26 jx ; upper cell similar in colour or just a 
shade darker than lower, and rather larger, 22-29 x 21-27 /u; 

lower cell either the counterfeit of upper or slightly tapering 
towards base, 17-23 X 20-22 fi; pedicel hyaline, persistent, 
generally much longer than spore, up to 90 and 110 yu long. 

On leaves, branches, involucral bracts and corolla-leaves of Helianthus 
<innuus L., and H. tuber osus L. 

Victoria — Very common, December- April. 

New South Wales — Mudgee and Guntawang, Feb., 1887 (National 

Herb.) (Hamilton). Sydney Botanic Gardens, Feb., 1901. 
Queensland — Ipswich, 1888, 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 
possesses no aecidial stage, since he has examined specimens from numerous 
localities without result, but Carleton 3 has collected the three stages in 
America and remarks : — u The aecidium occurs rarely in comparison with the 
occurence of other stages, but it is to be found on a number of hosts and 
occasionally in considerable abundance. This rarity of its occurrence, 
together with the occurrence of spermogonia so often with the uredo, may 
be accounted for by the fact that the uredo is often produced by direct 
teleutosporic infection." 

According to Woronin 1 the teleutospores germinate equally well 
whether they have been kept dry in a room or taken from leaves which had 
lain under the snow throughout the winter. Carleton 2 says that they also 
I germinate at once without a resting period. 

(Plate VII., Fig. 56.) 


37. Puccinia hypochoeridis Oud. 

Oudemans in Nederl. Kruidk. Arch. II., Ser. 1, p. 175 (1873) 

Sydow, Mon. Ured. I., p. 100 (1902). 

Sacc. Syll. VII., p 634 (1888); XVII., p. 302 (1905). 

Puccinia microseris McAlp., Agr. Gaz. N.S.W. VI., p 

757 (1895). 


Puccinia — Com-po sitae. 


Uredosori amphigenous or frequently on stem, genera ^ 

llv seated 


minute spots, scattered, pulverulent 


Uredospores globose, sub-globose or ellipsoid, echinul 

brown, with one prominent germ-por 




0-28 x 1 



III. Teleutosor 


blackish to 

often on stem, scattered or crowded 









to 1 mm. diam., but often smaller 

d on stem and up to 

mm. long. 


at fir 

ed with uredospor 



ellipsoid or obovate, apex 

Uy rounded, and not thickened 

rounded at base or 
septum, dark-brow n, 
cellular, 30-49 x 17 


uated, hardly 

stricted at 

ery delicately punct 

occasionally tri- 


h l 


,ge 3 

x 20 fx ; pedicel hyaline, 

lly deciduous, sometimes persistent and may 

reach a length 

of 77 ft. 

On Hypochoeris radicata L 




(Robinson). My 

L889 (Reader). Ardmona 
March, 1900 (Brittlebank). 



Melbourne, Apr., 1901. Dookie, Oct., 1903. Altona Bay 
Oct.. ] 

X. w 

903 (C. French, jun.). Murramurrangbong Ran 
Dec, 1903 (Robinson). , . - 

New South Wales — Wagga (Maiden). 

On H. glabra L. 

Victoria— Dimboola, Nov., 1889, and July, 1892 (Reader). 

Queensland — Brisbane, 1886 (Bailey 5 ). 

The punctation on the teleutospores is so fine that it may easily be over- 
looked, but when the spores are mounted in water it is clearly seen. 

A specimen from Syd. Ured. Exs. 673 on II. glabra agreed very closely, 

25 x 21 ju, and the teleutospores 

the uredospores being 

on an 


34 x 18 ft. 

This species closely resembles in morphological characters P. hieracd 
(Schum.) Mart., but Jacky has shown by means of cultures that the latter 
can only grow on Hieracium species, and cannot be transferred to other 
genera of Composites. Occasionally there may be a three-celled teleutospore 
reaching a length of 56 /x. \ 

P. microseris described as a new species in the Agricultural Gazette 
New South Wales was based upon a wrong determination of the host-plan 
by the collector since it was afterwards found to be Hypochoeris radicata L. 

Darluca filum Cast., is sometimes common on the mixed uredo and 

(Plate VIII., Figs. 62, 63.) 

88. Puccinia kalchbrenneri I)e Toni. 


De Toni in Sacc. Syll. VII., p. 645 (1888) 
Cooke, Handb. Austr. Fung., p. 337 (189 
Sydow, Mon. Ured. I., p. 93 (190 

II. Uredosori on both surfaces, but 


under, seated on in 

determinate spots which are often confluent and variously coloured 
scattered or gregarious, 


at first covered, convex, 


imately free and discoid 

Puccinia — Compositae. 


Uredospores globose, subglobose or ellipsoid, warted, yellow to 
pale brown, 20-30 x 19-26 ju. 

III. Teleutosori hypophyllous, seated on the same spots, minute, brown. 

Teleutospores oblong or subclavate to lanceolate, apex thickened 


attenuated or rarely truncate, constricted at sept 
dark-brown, 40-57 x 15-22 /n, rarely up to 27 /x broad; pedicel 
hyaline, short, deciduous. 

On living leaves of Helichrysum sp. 


There is no specimen of this species from Australia in the Kew Her- 
barium, and I have not found it on any species of Helichrysum, though this 
genus is exceedingly common near Melbourne. 

89. Puccinia lagenophorae Cooke. 

Cooke, Grev. XIIL, p. 6 (1884). 

Cooke, Handb. Austr. Fung., p. 335 (1892). 



Sydow, Mon. Ured., L, p. Ill (1902), and p. 863 (1904). 
. Sacc. Syll. VII., p. 612 (1888). 

P. hypochoeridis McAlp., Proe. Roy. Soc. Yie. VII., N.S., 

p. 217 (1894). 

P. macalpini Sydow, Mon. Ured., L, p. 100 (1902). 

I. Aecidia amphigenous, spots none ; pseudoperidia scattered or in 

groups, semi-immersed, margin lacerated, white, 200-240 fx diam.; 
peridial cells finely warted, lozenge shaped or elongated, 24-31 p 

Aecidiospores subglobose or elliptical, orange-yellow, very 
finely echinulate, 12-14 p diam. or 17-19 x 14//. 

[II. Uredosori scattered, small, pulverulent, brown or mixed with 


Uredospores globose, brown, epispore rough, 20 ft 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, 

J average 49 x 1 7 jx ; upper cell subglobose or oblong or somewhat 

conical, rounded, flattened, or bluntly pointed at apex, which is 
considerably thickened, generally equal in length to lower cell ; 
lower cell pale in colour, attenuated into pedicel and narrower 
than upper cell ; pedicel persistent, generally slightly tinted, 
attaining a size of 42 x 7 p. 

X. Mesospores relatively numerous, pale coloured, elongated elliptical, 

thickened at apex, with persistent hyaline pedicel, 38-42 x 

16-17 p. 

On stems and leaves of Lagenophora billardieri Cass. 

Victoria — Omeo, 1884 (Stirling). Ardmona, Oct., 1894 (Robin- 
son). Near Melbourne, Sept., 1900, I. (C. French, jun.) 
Murramurrangbong Ranges, Nov., 1902, Dec., 1903, Jan., 

1905 I., III. (Robinson). 


Puccinia — Com-po sitae. , 

On L. huegelii Benth., and L. billardieri Cass 

Tasmania — Mt. Dromedary 


Mt. Direction 






89-1, I. (Rod 

(Rod way 1 ) 

Devonport, Jan., 1906 (Robinson) 

Although material was very plentiful, careful search failed 


have given 

presence of uredospores, and I 

Cooke. Massee informs me that P 

sented in the Kew Herbarium, and he, therefor 






from diffe 

I have 


of plants with th 

localities and at different 

authority of 
is not repre- 
refer to the type 
5 rust upon them 
and I have never found 

Trie leaves on which P. hypochoeridis McAlp 

found proved to be- 

not Hypochoeris 
agreed with the above 


but Lagenophora billardieri, and the fung 

(Plate VII., Figs. 54, 55 

Plate F., Fi<?. 27.) 

90. Puccinia podolepidis McAlp. 


0. Spermogonia on small, honey-coloured spots, forming minute, dark- 

coloured points, hemispherical, pale yellow by transmitted light,, 
with round mouth, without projecting paraphyses, about 1 20 p diam. 
Spermatia minute, hyaline, oval, 3 X 2 ju. 

1. Aecidia bright orange, on both surfaces of the leaf, numerous 



in circular groups, which, however, often 
coalesce into large patches ; pseudoperidia cup-shaped, embedded in 
tissue, with white reflexed, laciniate margins ; peridial cells oblong 
to lozenge shaped, with striated margins, 28-32 x 16-18 p. 

Aecidiospores deep orange, ellipsoid to subglobose, smooth, 
24-32 x 16-22 it, or 22-24 p diam. 

III. Teleutosori black, surrounding the aecidia. 

Teleutospores chestnut-brown, oblong to clavate, constricted at 
septum, smooth, rounded or truncate or occasionally pointed and 
thickened at apex (9-12 u), rounded or attenuated towards base,. 

upper cell darker than lower, 38-54 X 22-32 p. ; pedicel per- 
sistent, hyaline, relatively short. 
X. Mesospores common, similarly coloured to teleutospores or often 

paler, ellipsoid, obovate to wedge-shaped or subclavate, rounded and 
thickened at apex, pedicellate, 32-44 X 16-22 p. 
On leaves of Podolepis longipedata A. Cunn. 

Victoria— Buffalo Mts., Nov., 1903, I. (C. French, jun.). 

near Bright, Dec, 1904, I., III. (C. French, jun.). 

Several teleutospores had 


side, but only one in lower cell. 

two germ 

pores in upper cell, one on either 

The teleutosori were rather sparse 

(Plate XXIX., Figs. 256, 257.) 


91. Puccinia prenanthis (Pers.) Lindr. 

Lindroth, Myk. Mittheil, p. 6 (1901). 
Cooke, Handb. Austr. Fung., p. 334 (1892) 
Sydow, Mon. Ured. I., p. 106 (1902). 
Sacc. Syll. XVII., p. 306 (1905). 

Puccinia — Comfo sitae* 


I. Aecidia hypophyllous, a few rarely epiphyllous, or on petioles, seatec 

on orbicular or elongated yellow or purple spots, up to 1 cm. ir 
diam., in round or irregular groups, at first hemispherical, thei 
flat, yellow or whitish, sometimes yellowish purple. 

Aecidiospores globose, subglobose or ellipsoid, delicately warted 
pale orange, 13-24 f.i diam. 

II. TIredosori hypophyllous, on pale indeterminate irregular spots 

scattered, minute, punctiform, pulverulent, pale brown. 

Uredospores globose or subglobose, echinulate, yellowish-brown 
16-24 u diam. 

III. Teleutosori similar, girt by the ruptured epidermis, dark-brown. 
' Teleutospores ellipsoid, rounded but not thickened at apex, 

mostly rounded at base, not constricted at septum, delicately 
warted, brown, 26-36 x 16-24 fx ; pedicels hyaline, very short. 

On living leaves of Lactuca. 

Victoria. New South Wales. S. Aust 

No specimen seen. It is given on the authority of Cooke, but 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. Tired. L, p. 867 (1904). 
Sacc. Syll. XVIL, p. 277 (1905). 

I. Aecidia on blister-like swellings on stem and branches, on upper and 

under surface of leaves, on flower-head stalks and involucre, causing 
discoloration and distortion, and usually surrounded by paler green 
tissue, about ^ mm. in diam., disposed in large circular or oval 
clusters or irregularly ; pseudoperidia cup shaped, with white, 
irregularly laciniated everted edges, tubercular before opening ; 
peridial cells firmly united, overlapping each other, with striated 
margins and usually broader at one end, individually slipper- 
shaped, 25-35 x 13-17 fi. 

Aecidiospores spherical or angular, orange colored, very delicately 
warted, 13-16 /u diam., or 14-17 x 11-15 p. 

III. Teleutosori for a long time covered by epidermis, then erumpent 

and epidermis usually thrown off or remaining in shreds and 
patches, intermixed or running parallel with aecidia, black, pulvi- 
I nate, oval, up to 1 mm. long, often confluent in elongated lines. 

Teleutospores oblong to clavate, chestnut-brown, slightly con- 
stricted at septum, smooth, 36-63 X 15-25 y, average 54 x 20 //, 
occasionally tricellular when 48-73 fj. long ; upper cell deep chest- 
nut-brown, rounded or somewhat oval, conoid, or truncate and 
thickened at apex (up to 13 yu) ; lower cell usually paler in colour, 
rounded at base or tapering, often elongated relatively to upper ; 
pedicels persistent, pale yellow to hyaline, sometimes longer than 

X. Mesospores very common, elongated oval, oblong, or somewhat 

elliptical, apex rounded or pointed and usually thickened, golden- 
yellow to chestnut-brown, 29-44 x 13-17 /j. 


Puccini a — Compo sitae. 

On leaves and stems of Senecio vulgaris L. 

Victoria (Ralph), (Robinson), (French, junr.), &c, I. III. 
New South Wales (Hamilton), I. 
* Tasmania— 1891, 1893, 1895, 1897, I. III. (Rodway >). 

On Senecio pectinatus DC. 

New South Wales — Mount Kosciusko, Jan., 1898, I. (Maiden). 

On Senecio brachyglossas F.v.M. 

Victoria — August, 1900, I. (Reader). 

On Senecio velleioides, A. Cunn. 

Victoria— Sealers' Cove, 1854, I. Port Phillip, 1886, I. 


It differs from P. senecionis Lib., in the aecidia being amphigenous and 
not hypophyllous ; also in the teleutospores being considerably larger, 
thickened at apex, and not provided with a hyaline papilla, while the pedicel 
is decidedly persistent and elongated. Groundsel, or S. vulgaris, being an 
imported plant, and so cosmopolitan in its character, it was considered 
strange that a new species of rust should be found upon it, and although I 

had named it in MS. P. australiensis, yet I delayed publishing it under 

that name, hoping bo find the teleutospores on a native Senecio. 

Teleutospores are very common in some localities during the winter and 

spring months, and the Groundsel growing on the coast at Port Fairy was 

so overrun with both aecidia and teleutosori that the plants were stunted 

in growth. 

S. pectinatus DC, occurs in the three States from which Puccinia 

tasmanica has been recorded, but only the aecidia! stage has been met with 

in New South Wales. 

(Plate VII., Fig. 52.) 

93. Puccinia vittadiniae McAlp. 



I. Aecidia on both surfaces of leaf, but mostly on upper, scattered or in 

groups and confluent, minute ; pseudoperidia white becoming 
yellowish, immersed, with lobed margin ; peridial cells with striated 
margin in section, striae projecting and appearing as points in sur- 
face view, oblong, 25-32 x 13-16 p. 

\ ^ ] Aecidiospores yellowish, ellipsoid, very finely verrucose, 14-17 x 

11-13 /Li. 

III. Teleutosori minute, black, sparsely developed, intermixed with 


Teleutospores dark chestnut-brown, oblong to elongated oblong or 
clavate, smooth, constricted at septum, generally rounded at base 
and apex, sometimes bluntly pointed or truncate, and thickened at 


34 X 21 fx ; upper 


apex (6-9 /u), 31-46 x 18-25 /u, 
darker than lower and similar or sometimes considerably broader ; 
pedicel hyaline, persistent, 30-40 /x long, and up to 9 fx broad 
adjoining spore. 

X, Mesospores very common, similarly coloured to teleutospores and 

stalked, ovoid to elongated ellipsoid, rounded or beaked at apex and 
thickened (up to 9 /u), 25-34 x 14-17 /u. 

Puccinia — Rubiaceae. 


On leaves including leaf stalks of Vittadinia australis A. Rich. 

Victoria — Dimboola, June, 1900 (Reader). 

In Agr. Gaz. N.S.W. VI., p. 757 (1895), I described a new species under 
the name of Aecidium vittadiniae upon a plant forwarded to me as Vittadinia 
australis, but which was afterwards found to be an Erechtites, so that the 
present description replaces the previous one. 

(Plates XXXIX., Fig. 292 ; XLIIL, Figs. 308, 309.) 


94. Puccinia coprosmae Cooke. 


Cooke, Grev. XIX., p. 2 (1890). 



Sydow, Mon. Ured. I., p. 209 (1902). 
Sacc. Syll. IX., p. 300 (1891). 

P. coprosmatis Morrison, Vict. Nat. XL, p. 90 (1894). 

III. Telcutosori hypophyllous, sometimes epiphyllous, rusty-brown, 

prominent, compact, round or elliptic, usually in groups, sometimes 
forming mammilated tubercles, rarely solitary, densely crowded 
and confluent, sometimes surrounded by the ruptured epidermis 
or naked and deforming the leaves. 

Teleutospores elongated oblong to clavate, pale yellowish to 
brownish, the two cells about equal and more or less ovate, apex 
bluntly pointed or rounded, thickened, (from Q-8 //), rounded or 
slightly tapering at base, smooth, constricted at septum, 
35-51 X 16-22 fj, average 43 x 18^/; pedicel hyaline, persistent, 
elongated, up to 110 fx long and 8 li broad. 

X. Mesospores common, similarly coloured to teleutospores, ovoid to 

ellipsoid, or somewhat fusoid, usually pointed and thickened at 
apex, 32-41 x 19-22 /u. 

On leaves of Coprosma billardieri Hook. 

Victoria — Kew and Dandenong, Oct. and Feb., 1893. 
Tasmania — (Rod way ' ). 

On Coprosma hirtella Labill. 

Victoria — Murramurrangbong Ranges, Dec. 1903, Jan. 1905 


I am indebted to Dr. Morrison for some of the original material on 
which he founded his new species P. coprosmatis, but it is identical with 
the above species first determined by Dr. Cooke on Coprosma lucida from 
New Zealand. 

Darlucafilum Cast., was very plentiful on the teleutosori on Cojyrosma 
billardieri, and was described by Morrison as a spermogone with spermatia. 

(Plate X., Fig. 78.) 

95. Puccinia oliganthae McAlp. 

A sperula 

II. Uredosori hypophyllous, cinnamon-brown, round to ellipsoid, soon 

naked, girt by the ruptured epidermis, confluent into elongated 


Puccinia — Rubiaceae. 

V redospores globose to shortly elliptical or obovate, golden- 
brown, finely echinulate, one to two germ-pores on one face, 
25-29 X 22-25 li, or 25-27 li diam. 

III. Teleutosori on stems, dark-brown to black, elongated, bullate, 

soon naked, compact, 2 mm. long or more. 

Teleutospores narrowly clavate to oblong, brown, constricted at 
septum, thickened at apex (9-10 li), and rounded or bluntly pointed 

ay be 

lly attenuated towards base 

and paler in colour, 32-54 x 16-21 li, average 48-16 li ; pedicel 
yellowish, persistent, up to 48 li long. 

X. Mesospores occasional, similarly coloured to teleutospores, somewhat 

fusiform, with pointed and thickened apex, 28-32 x 12-13 li. 

On stem and leaves of Asperula oligantha F. v. M. 

Victoria — In shady gullies of Murramurrangbong Ranges, Nov., 

1902 (Robinson). 

The examination of specimens of P. punctata Link, on species of Asperula 
from Exsicc. Sydow Ured. 465, 466, showed that the two were distinct. In 
P. oligantha the uredospores are broader, and while the teleutospores 
average only about 16-18 li broad, in P. punctata they are about 20-22 /<. 
In the latter too the apex is much thicker, reaching up to 16 li. The 
species closely resembles P. asperulae odoratae Wurth, but aecidia are 
present on the latter, and the teleutospores are only thickened at the apex 

the extent of 7 u 

(Plate X., Fig. 7 

96. Puccinia operculariae (Morr.) Syd. 


Morrison, Victorian Nat. XI., p. 119 (1894). 

Sydow, Mon. Ured. L, p. 224 (1902). 

Sacc. Syll. XIV., p. 321 (1899) ; XVII., p. 317 (1905). 

Aecidium cystoseiroides Berkeley, Fl. Tasm., p. 270 (1860). 

I. Pustulate, deforming the leaves ; pseudoperidia immersed. 

Aecidiospores orange, subangular. 

III. Teleutosori hypophyllous, on orbicular brownish to yellowish spots, 

solitary or a few together, round or elliptic, compact, reddish- 
brown, girt or partially covered by the ruptured epidermis, up to 
2 mm. long. 

Teleutospores golden-bi own, oblong to oblong clavate, constricted 
at septum, smooth, 35-54 x 14-20 /u; upper cell rounded or 
pointed and thickened at apex (up to 11 li), about equal in length 
to lower ; lower cell usually tapering towards base or sometimes 
rounded ; pedicel hyaline, persistent, 80 li or more long and 6-7 ii 
broad . 

On living leaves of Opercularia aspera Gaertn. 

Victoria — Genoa River (F. V. Mueller). 

On leaves and petioles of Opercularia varia Hook. f. 

Victoria — Oakleigh, Nov., 1893 (Morrison). 

Tasmania— Near Hobart Rivulet, Nov., 1860, I. (Gunn), 1902 

(Rod way 1 ). 


Puccinia — Loranthaceae. 


Morrison states that this fungus deforms the leaf and produces a con- 
cavity on the opposite side. He made this a variety of P. coprosmae Cooke, 
and certainly there is a very close resemblance, but the sori in the latter are 
generally in groups, and the apex of the spore is not quite so thick. 

No specimens of the aecidial stage were seen, but being on the same 
host-plant as the teleutospores, it is included in this species. v 

Darluca filum Cast., is common on the teleutosori, although it is usually 

nly found on ured 

(Plate XXIX., Fig. 258 



97. Puccinia loranthicola McAlp 

I. Aecidia imbedded in raised brownish to dark brownish, often con- 
fluent cushions on one or both surfaces of leaf, orange ; pseudo- 
peridia tubular, with white, reflexed, much torn margins ; peridial 
cells oblong, with striated margins, 43-50 X 27-31 a. 


Aecidiospores ellipsoid to oblong or sub-angular, b 

-,"*«,*, »x*gwu viaug 

yellow, decidedly echinulate, 37-56 x 22-31 li 

II., III. Uredosori amphigenous, brownish, bullate, in scattered 

groups, often arranged in circles and confluent, surrounding 
central darker teleutosori, epidermis splitting lengthwise and 

II. Uredospores orange yellow, ellipsoid to oval, obovate or oblong, 

coarsely echinulate, with 3-5 equatorial germ-pores on one face, 
40-65 x 22-32/7, occasionally 80 x 22/*; epispore 3-4J li thick. 

III. Teleutospores intermixed with uredospores, subhyaline, cylindrical 

to clavate cylindrical, smooth, slightly constricted at septum, 
rounded and slightly or not at all thickened at apex, tapering 
slightly towards base, 65-94 x 15—24 yu, occasionally 3-celled, 
when about 86 fx long ; pedicel hyaline, persistent, elongated up to 
200 Li long, and swollen towards apex up to 14 fi. 

X. Mesospores subhyaline and with elongated pedicel like that of 

teleutospore, smooth, fusoid, hardly thickened at apex, 71-77 x 

25-28 ft. 

On living leaves of Loranthus celastroides Sieb., growing on Stringy bark 
{Eucalyptus sp.). 

Victoria — Murramurrangbong Ranges, Jan., 1905 (Robinson). 


All the stages were found on the same tree, but the aecidia on separate 
leaves. All the spore-forms are particularly large, a fact which probably has 
some relation to the peculiar nutrition of the fungus, and the height at which 
it occurs, some specimens being obtained fully 50 feet from the surface of 
the ground. 

The aecidia are very conspicuous on raised cushions, with corresponding 
depressions on the opposite side, where aecidia may also occur. 

The uredosori are of a ruddy brown, in isolated groups, generally 
arranged circularly and with teleutosori in the centre. Puccinia loranthi 
Speg., has only teleutospores, and it is doubtful if they are to be regarded 
as such. P. macrocarya Rac. on leaves of Loranthus in Java has only aecidia 

and teleutospores. 

(Plate XXXI., Figs. 268-271.) 


Puccinia — Umbelli ferae . 


98. Puccinia thuemeni (Thuemen) McAlp. 

P. castagnei Thuemen, Rev. Myc. II., p. 86 (1880). 


O. Spermogonia on both surface 


leaf, pale yellow, round, in groups 

ed yellow spots, about 150 /u diam 

Spermatia hyaline, ellipsoid, 3*5-4 x 




II. u 

amphigenous, bullate 






ded, and becoming confluent, surrounded or partially covered 

by the ruptured epidermis, pulverulent, cinnamon-brown, 1 mm 

more long 

Uredospores obovate to ellipsoid, golden-b 

echinulate* with 

ened hood 



d three or four, mor 

quatorial germ-pores on one face 




x 21 


hyaline pedicel may reach a length of 56 

III. Teleutosori similar to uredosori, dark-brown, also on both surfaces 

of leaf, but more 


coloured to 

dospores, shortly oblong to ovate-oblon 

d intermixed with 
ightly or not at all 

cted at septum, finely warted, rounded at apex, and not pe 

ceptibly thickened, 29-40 x 
lower cell rounded at bas 
hyaline, short, decid 





32 x 20 ft 

attenuated ; pedicel 

On stems and leaves of Avium prostratum Labill 

Victoria — Beaumaris, Dec.-Julv, II., III. (III. in 


Portland, Jan., 1901, II., III. Sandringham, Jan. and Feb., 

Mentone, Jan. -Sept., 


1904, II., III. (III., very sparse). 

1905, II. III. 

Tasmania — Hobart, Sept., 1905 (Rod way). Mersey River, January, 

1906 (Robinson). 
It may be found in sheltered spots all the year round. 

On Apium graveolens L. 

Victoria — Near Melbourne, April-Sept., II. III. 
Tasmania — Hobart, Sept., 1905 (Rod way). 

Very common in the early spring months. 

Spermogonia were only found on the native celery (Apium prostratum) 
associated with uredospores and teleutospores. 

The germ-pores of the uredospores may be in a transverse band, or one 
may be placed above the other. When fully developed the uredospores are 
similarly coloured to the teleutospores, otherwise they are yellowish. On 
Apium graveolens both uredospores and teleutospores agree almost exactly in 
average size, though of the latter there are usually very few more than 33 p 
in length. There are occasionally elongated teleutospores which reach a 
length of 40 p. The finely warted epispore is a constant feature of the 
teleutospores on both hosts. 

The illustrations of spores in PL IX., Figs. 68-72, show that the rust on 

the native celery (Apium prostratum), and on cultivated celery (A. 
graveolens) is the same in Australia. 

Figs. 73, 74 show the smooth teleu- 

tospores of P. apii Corda on Apium graveolens from Sydow's Ured. Exsicc, 
558, which is quite distinct from the finely warted P. thuemeni, and Fig. 7d 
shows the very rough and knobby teleutospores of P. bullata (Pers.) Schroet. 


Puccinia — Umbelli ferae. 169 

on Aethusa cynapium from Sydow's Ured. Exsicc, 1261, which is readily dis- 
tinguishable from either of the above. There is thus a clear distinction be- 
tween these three recognised species, but the synonymy is rather confusin 
The reason for the name I have adopted may first be given. P. castagnei 
Thuem., on cultivated celery agrees with our species, but P. castagnei 
Schroet., in Cohn's Beitrage, III., 62 (1879), was first applied to a Puccinia 
on Thalictrum angustifolium, and Thuemen's name being thus preoccupied, 
I have substituted P. thuemeni. 


As regards the synonyms, Cooke, in his Australian Handbook, gives P 
i Corda at first, and then in an addendum substitutes P. castaqnei Thuem 

Then Saccardo, in his Sylloge, gives P. bullata with P. apii as a 
synonym, and P. castagnei is regarded as distinct. Further, Sydow in his 
Monograph includes P. apii Corda, and P. castagnei Thuem., under P. apii 
Desm., and P. bullata is given separately, although he remarks that the 
latter is probably a collective species. The teleutospores of P. bullata are 
described as smooth, and therefore do not agree with the quoted specimen 
from Sydow. Finally, it may be noted that Tranzschel 1 has shown that the 
host-plant of P. castagnei Schroet., is not Thalictrum angusti folium, but an 
Umbellifer, and probably Silaus pratensis, so that this species is a synonym 
of P. bullata. 

Darluca filum Cast., occurs on the uredosori and teleutosori. 

(Plate IX., Figs. 68-75 ; Plate C, Figs. 16, 17.) 


99. Puccinia xanthosiae McAlp. 

II. Uredosori amphigenous, and on leaf -stalks, dark-brown, pulvinate, 

gregarious, elliptical, often confluent, rupturing epidermis, about 
j mm. long, often causing entire leaf to become brownish. 

Uredospores golden-brown, thick-walled, echinulate, globose to 
elliptical, 30-32 )< diam., or 30-37 x 24-28 p, average 34 x 26 /x; 
pedicel colourless, 4-6 p broad. 

III. Teleutosori rare, on leaves of previous year, minute, black. 

Teleutospores oblong to oblong clavate, dark-brown, smooth, 
constricted at septum, 40-60 x 20-25 jj, average 42 x 20 p ; 
upper cell generally darker than lower, rounded and slightly 
thickened at apex, occasionally truncate ; lower cell rounded at 
base or attenuated towards stalk ; pedicel persistent, tinted, about 
20 p long. 

X. Mesospores occasional, similarly coloured to teleutospores, obovate, 

thickened at apex, 32 x 20 ju. 

On Xanthosia pusilla Bunge. 

Victoria— Frankston, Oct., 1899, II. : Oct., 1903, III. Sandring- 

ham, Sept., March, II, III. ; Feb., II. 

(Plate IX., Fig. 76.) 


Puccinia — Onagraceac^ Rosaceae. 


100. Puccinia epilobii-tetragoni (DC.) Wint. 

Winter, Pilze, p. 214 (1884). 
Sydow, Mon. Ured. L, p. 424, (190 
Sacc. Syll. VII., p. 608 (1888). 


I. Aecidia distributed en 

crowded, hypophyll 
laciniate. revolute re 

surface of the leaf, scattered 

doperidia cup-shaped 

th white, 


Aecidiospores orange yellow, polygonal, finely warted, 16-26 
diam., 18-21 \i being common. 

II. Uredosori chestnut-brown, scattered, or often orbicularly arranged 


sometimes confluent, pulverulent, hypophyllous, soon naked 


psoid or 

cL clear bro 



two germ-pores on one face 
reaching a length of 31 p. 


7 x 16-21 


III. Teleutosori dark-brown, round, often arranged on 


hypophyllous, soon naked, pul\ 


like spot 

intermixed with 


htly constricted at 

elliptic or oblong, 
smooth, 27-36 


x 15 



erage 30 x 

18 /.i ; upper cell with a cap-like 
thickening at apex (5-6 fj) ; lower cell usually rounded at base 
or sometimes slightly tapering ; pedicel hyaline, slender, deciduous, 

X. Mesospores exceedingly rare, similarly coloured to teleutospores and 

thickened at apex (5 ft), subclavate to obovoid, smooth, 21-27 x 
13-15 /j, with hyaline pedicel. 

On leaves of Efilobmm glabellum Forst. 

Victoria — Dimboola, Dec, 1892, I., II., III. (Reader). Ardmona, 

Arthur's Creek, Kergunyah, Nyora, Pakenham, &c, Aug. 
Dec, I., II., III. 

On E. billardierianum Ser. 

Tasmania — Great Lake, Feb., 1894 (Rodway 

On Epilobium sp. 


Tasmania — Summit of Mt. Wellington, Jan. 1892, I. (Rodway). 
Sydow, in his Monograph, distinguishes between P. epilobii DC, with 
teleutospores alone, P epilobii-fleischeri Fisch., with aecidia in addition to 
teleutospores, and the present species with the three stages. 


(Plate X., Figs. 79, 80, 81.) 


101. Puccinia gei McAlp. 


McAlpine, Agr. Gaz. N.S.W. VI., p. 756 (1895). 
Sydow, Mon. Ured. I., p. 484 (1903). 
Sacc Syll. XIV, p. 297 (1899). 

III. Teleutosori hypophyllous, confluent, coffee-brown, on greenish- 
yellow spots, which also occur on upper surface of leaf. 

Puccini a — Rosac eae. 


Teleutospore brownish-yellow, smooth, fusiform to clavate, con- 
stricted at sepfcam, epispore chestnut-brown, 30-40 x 13-18 //, 
average 35 X 14 p ; upper cell elongated and tapering or 
rounded, thickened at apex, 18-22 /* long; lower cell elongated 
and tapering towards base, or bulging, 16-22 p long; pedicel 
pellucid, sometimes 54 /t long. 

X. Mesospores occasional, coloured like teleutospores, elongated 

ellipsoid to oblong, rounded and thickened at apex, occasionally 
somewhat pear-shaped, and the basal portion prolonged like a 
stalk, as sometimes happens in the bicellular spore, 28-35 x 
12-18 ft. 

On living leaves of Geum renifolium F. v. M. 

Tasmania — The Calf, Adamson's Peak, March, 1895 (Rod 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. XI., p. 97 (1883). 
Sydow, Mon. Ured. I., p. 484 (1903). 
Sacc. Syll. VII., p. 648 (1888). 

II. Uredosori hypophyllous, occasionally epiphyllous, minute, light 

brown to cinnamon brown, orbicular, scattered, but often grouped 
in patches and confluent, soon naked, pulverulent, usually seated 
on discoloured spots, which also show on upper surface. 

Uredospores variable in shape, oblong to ovoid oblong, ellipsoid 
to somewhat piriform, closely echinulate, yellowish, with two or 
occasionally three sub-equatorial germ-pores on one face, apex 
yellowish brown, thickened (average, 5-6 ^u, or papillate up to 9 yu) 
with spines less prominent, bluntly conical or rounded, 25-38 x 
* 12-18 ju, occasionally reaching a length of 40 fx or more, average 
30 X 15 fi ; paraphyses intermixed, numerous, capitate, pale yel- 

low, long-stalked, sometimes attaining a length of (0 p. 

III. Teleutosori hypophyllous, scattered or confluent, isolated or in 

groups, very pulverulent, seal-brown, paraphysate, known from the 
uredosori when fully developed by their dark, almost black 


Teleutospores at fi 

dospores, dark-b 

oblong to ellipsoid or obovoid, densely warted, often composed of 
two globose or depressed globose cells, readily separating from 

each other, very rarely 3-celled, 25-45 x 17-25 n> average 

36 x 25 jx ; upper cell usually darker in colour, and broader 
lower, globose or depressed globose, often slightly thickened 
at apex, and thickly studded with short stout spikes, average 
16-22 fi diam.; lower cell generally oblong to obovoid, and equal to 
or longer than upper, 14-24 X 14-19 n ; pedicel short, hyaline, 

On leaves, fruits and stems of Peach {Primus persica Stokes). 
On leaves and fruits of Almond and Apricot (Prunus amygdalus Stokes, 
and P. armeniaca L.). 




Puccini a — Leguminosae. 

On leaves of Plum [Primus domes tica 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 Primus domestica. 

This rust is usually most prevalent towards the autumn, but in some 
seasons it appears in the early summer. It not only destroys the foliage, 
but may even attack the fruit, and in some seasons it actually ruptures the 
bark, especially in rapidly growing nursery stock. When the rust occurs on 
the fruit, uredospores may be produced both at the surface and in the pulp. 

There are said to be two kinds of uredospores by Dumee and 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- 


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 
s,ve very prominent on the upper surface. There they often run together, 
and have the appearance of yellow ochre. In the Apricot the spot is hardly 
noticeable at first, but when held up to the light the indeterminate yellow 
spots are seen clearly. The leaf soon turns yellow, and then the spots are 
of a pale green. In the Plum the spots are very pale on the lower surface, 
but on the upper surface the yellowish-green is very prominent. 

Tranzschel 2 has proved experimentally that the aecidial stage of the rust 
on Almond occurs on Anemone coronaria L., and it has hitherto been known 
as Aecidium <punctatnm Pers. 

Darluca filum Cast., is found on the uredosori. 

(Plate X., Figs. 83-86 ; Plate D., Figs. 19, 20.) 



103. Puccinia zorniae (Diet.) McAlp. 

Uredozorniae Dietel, Hedwigia XXXVIII., p. 257 (1899). 

II. Uredosori on both surfaces of leaf, but most numerous on under, 

yellowish when young, then ruddy-brown, round to elliptic, 

bullate, at first covered, then erumpent and surrounded by rup- 
tured epidermis, gregarious and sometimes confluent. 

Uredospores old-gold colour, ellipsoid to occasionally obovoid, 
finely echinulate, with two germ-pores on one face, 25-32 x 19-22/1. 
III. Teleutosori as above, but dark-brown. 

Teleutospores intermixed with uredospores and similarly 
coloured, oblong, smooth, deeply constricted at septum, two cells 
about equal in size or lower sometimes a little longer, 32-44 x 
20-24 /i, average 36 x 21 /i ; upper cell rounded and thickened 
at apex (up to 6 /lc), lower cell rounded at base ; pedicel hyaline, 
persistent, about length of spore. 


Puccini a — Ficoideae. 


On living leaves of Zornia diphylla Pers. 

New South Wales — Richmond River, May, 1904 (Musson). 

Uredozorniae Berk., is mentioned by Cooke in Grev. XX., p. 110, as on 
Zcrnia in Mauritius, but no diagnosis is given. This is the first record of 
the teleuto stage, since Dietel in his Uredineae brasilienses only men- 
tions the uredo stage on the same plant from Rio Janiero, and thus describes 
it: — "Sori hypophyllous, scattered, surrounded by the ruptured ochraceous 
epidermis, minute, brown. Uredospores globose, obovate or ellipsoid, 
21-28 x 18-23 fx ; epispore brown, echinulate, with three germ-pores." 

Darluca Jilum Cast., occasionally on sori. 


(Plate X, Fig. 87.) 


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 f.i 

in diam. 

Spermatia hyaline, globose to oval, 2 J- 3 J /j. diam. 

1. Aecidia hypophyllous, crowded, often arranged in lines, greenish to 

yellowish-brown knob-like swellings before bursting ; pseudoperidia 
white, cylindrical, with torn edges, average 630 ji diam. 

Aecidiospores orange-yellow, minutely warted, variously shaped, 


lobose to ovate or mussel-shaped, 27-39 x 22-25 /u 

II. Uredosori amphigenous, soon naked, bullate, orange-red, generally 

orbicular, 1-1 ^ mm. diam. 

Uredospores orange-yellow, ellipsoid to ovoid, strongly echinu- 
late, 27-37 x 22-25 fj, average 31 x 24 ju. 

III. Teleutosori amphigenous, black, compact, bullate, orbicular, irre- 
gular or elongated, scattered, covered by epidermis for some time, 
then erumpent. 

Teleutospores golden brown to chestnut-brown, elliptical to 
oblong, usually rounded at both ends, slightly constricted at 
ptum, smooth, 40-60 x 25-36 ^, average 50 x 30 ju ; pedicel 

hyaline, stout, persistent, up to 80 

X. Mesospores similarly coloured to teleutospores, ellipsoid or elongated, 

thickened at apex, 35-54 x 25-29 ja. 

On leaves and stems of Tetragonia implexicoma Hook. f. 

Victoria. — On the coast at Sandringham, Beaumaris, Flinders, 

Portland, <fcc, all the year round. 
Tasmania — Hobart, about rocks, Aug., 1897, I., II., III. (Rod- 
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 

x74 Puccinia — Polygonaceae. 

large, orange, blister-like clusters quite conspicuous against the green of the 
leaf, and the rust-devouring Diplosis was ^frequently met with there and 
probably scatters the spermatia. 

The aecidia are usually swarming with the Diplosis, dipping into them 
as an insect inserts its proboscis into a flower, and sometimes it is entirely 
inside the cups. The larvae are invariably coated with the aecidiospores. 
Occur August to October and subsequently. 

Uredospores just beginning to appear in October, and the Diplosis was 

also found on the sori. 

Teleutospores are scanty in October, and they occur on the same leaves 

as uredospores. 

The uredo and teleutosori are often associated together in the same 
cluster, and the three spore-forms may all occur on one leaf. 

(Plate XL, Figs. 88, 89.) 



105. Puccinia ludwigii Tepp. 

Tepper, Bot. Centralb., XLIIL, p. 6 (1890). 
Cooke, Handb. Austr. Fung., p. 336 (1892) 

Sydow, Mon. Ured. L, p. 581 (1903). 

Sacc. Syll. XL, p. 196 (1895). 

II., III. Sori minute, circular, irregularly scattered or aggregated into 

small groups, soon naked and girt by the ruptured epidermis, on 
both surfaces of leaf, up to \ mm. diam. 

II. Uredospores sub-globose to ovate, very finely echinulate, pale 

yellowish-brown, 22-28 X 16-20 /x, or 19-22 /x diam. 

III. Teleutospores at first intermixed with uredospores, oblong to 

ellipsoid, rounded at both ends, very coarsely warted, slightly con- 
stricted at septum, not thickened at apex but generally with 
hyaline apiculus, sometimes three or four celled, dark brown, 30-40 

x 20-25 p, average 32 x 21 p; pedicel short, hyaline, deciduous. 
X. Mesospores similarly coloured to teleutospores, warted and with 

hyaline apiculus, oval, with short hyaline pedicel, 28-31 x 18-23 /u. 

On leaves of Rumex brownii Campd. 

"Victoria — Coromby, Oct., 1889 (Tepper). Shepparton, on river 

flats, Nov., 1895 (Robinson). Flinders, Jan. Murramurrang- 
bong Ranges, Dec. Killara, March. Myrniong, Aug. 

Queensland — Ennogera (Bancroft). Brisbane, Sept., 1886 



Tasmania — Devonport, January, 1906 (Robinson). 
On Rumex flexuosus Sol. 

Victoria — Warracknabeal, Oct., 1903 (Reader) 

I have had plenty of material of this species, both from Victoria and 
Queensland, as well as some of the original material from Ludwig, and ex- 
amination shows that it is the same fungus in each case. 

Mr. Bailey kindly supplied specimens from Queensland, and there is one 
in the National Herbarium, Melbourne, sent from Queensland to Dr. Cooke 
in 1886, and it is named in his own handwriting P. rumicis Lasch., 

P. acetosae (Schum.) Koern. It has the same characters as the others, and 

comparing it with P. acetosae from Syd. Ured. Exs., 1163, I find that it 


Puccinia — Polygonaceae. 175 

differs in the more finely echinulate uredospores, but more particularly in 
the teleutospores, which are coarsely knobbed and altogether characteristic. 

The introduced Rumex crispus and R. acetosella are exceedingly common 
weeds in Victoria, and may be seen growing alongside rusty R. broumii, but 
no rust has ever been detected on them. 

R. brownii is a native species, and its parasite is evidently a native also. 

The teleutospores are varied in shape, and three and four celled forms 
are met with. In the three-celled forms they may either be after the 
P hragmidium or Triphragmium type, and in the four-celled forms the upper 
third cell is vertically divided. Just as in P. dichondrae there is every 
gradation from the one to the four-celled spore. 

Darluca filum Cast., is common on the uredo and teleutosori. 

(Plates XI., Figs. 90, 91, 92 ; XL., Fig. 300.) 


106. Puccinia muehlenbeckiae (Cooke) Syd. 

Sydow, Mon. Ured. L, p. 566 (1903). 
Sacc. Syll. IX., p. 299 (1891). 

Puccinia rumicis- scutati (DC) Wint. var. muehlenbeckiae 
Cooke, Grev. XIX., p. 47 (1890), and Handb. Austr. Fung., p. 

336 (1892). 

II. Uredosori hypophyllous, scattered or in irregular groups, some- 
times circularly arranged, round or elliptic, girt by the torn 
epidermis, cinnamon-brown. 

Uredospores yellowish-brown, ellipsoid or ovoid, echinulate, 

24-32 x 16-21 /a, average 28 x 19 p. 

III. Teleutosori similarly arranged to uredosori, but dark-brown. 

Teleutospores at first intermixed with uredospores, clavate to 
oblong, dark-brown, constricted at septum, thickened at apex (up 
to 7 /u), and rounded or obtuse, sometimes 3-4 celled, 32-45 x 
14-19 ju, occasionally reaching a length of 55 /*, average 40 X 
1 8 a ; lower cell generally somewhat triangular ; pedicel hyaline, 


ally persistent, up to 40 f.i Ion"" 


X. Mesospores occasional, similarly coloured to teleutospores, ellipsoid to 

elongated oblong, smooth, rounded or conoid and thickened at apex 
(up to 5 p), 31-34 x 12-18 /w. 

On leaves of Muehlenbeckia adpressa Meissn. 

Victoria— Brighton, May, 1894 (Morrison). Near Melbourne, 

Jan.-July. Cape Schanck, March, 1903 (C. French, jun.). 
Frankston, Feb., 1904 (Robinson). Sandringham, Sept., 
1905, II., III. 

On Muehlenbeckia gracillima Meissn. 

Queensland— Gladfield (Gwyther) (Bailey 1 *). 
N.S. Wales — Kurrajong Heights (Musson). 

Some of the mesospores show their connexion with teleutospores very 
clearly. In one instance two germ-pores were formed in the usual positions, 
while in another there was a slight notch on one side as if the beginning of 
a septum. They are evidently one-celled teleutospores, and have all their 
characters with the exception of the septum. 


1 76 Puccinia — Chenopodiaceae. 

There were spots on the upper surface of the leaves with purplish-red 
margins, but these were due to gall-mites. The spores are of the general type 
of P. rumicis-scutati, but both kinds are somewhat smaller. I had a large 
amount of material to operate upon, and found the sori to be hypophyllous, 
although Cooke inadvertently describes them as epiphyllous. In a specimen 
of P. rumicis-scutati, from Syd. Ured. Exs. 25, the teleutospores are of a deep 
chestnut-brown, and attain a length of 57 /m. Sydow himself had not seen 
any specimens of the rust on Muehlenbeckia, but even from the brief 
description of Cooke he was right in making a species of it. 

Darluca filum Cast., sometimes common on uredosori. 

(Plate XL, Fig. 93.) 




10/. Puccinia dielsiana P. Henn. 

Hennings, Hedw. XL., p. (95) (1901). 
Sydow, Mon. Ured. L, p. 566 (1902). 
Sacc. Syll. XVIL, p. 361 (1905). 

III. Teleutosori on stems, large, compact, bullate, erumpent, up to f cm. 


Teleutospores ellipsoid or oblong ellipsoid, generally rounded at 
both ends or obtuse and slightly thickened at apex (3-5 ll), smooth, 
very slightly constricted at septum, with granular contents, 
chestnut or dark-brown, 35-18 x 18-28, average 38 x 22 fx ; 
pedicel sub-hyaline, persistent, 80-120 x 5-6 fx. 

X. Mesospores occasional, ellipsoid to ovoid, coloured and thickened at 

apex like teleutospores, 34 x 23 fi. 

On stems of Chenopodiaceae — Threlkeldia drupata Diels. 

W. Australia— Near Perth, 1900. 

The teleutospore has occasionally a vertical septum, and in elongated 
slender specimens the breadth is reduced to 18 u. 

XL, Fiff. 94 

108. Puccinia kochiae Mass. 

Kochia, Enchylaena. 

Massee, Grev. XXIL, p. 17 (1893). 
Sydow, Mon. Ured. I., p. 565 (1903). 
Sacc. Syll. XL, p. 196 (1895). 

II., III. Sori amphigenous, discoid, plane, very compact, blackish- 
brown to black, often aggregated together, girt by the ruptured 
epidermis, up to 1 mm. diam. 

II. Uredospores elliptical, golden-brown, decidedly echinulate, with 

numerous prominent germ-pores, as many as twelve on one face, 
29-34 x 18-25 li, 32 x 23 li being very common. 

III. Teleutospores densely packed, intermixed with uredospores, 

elliptic-oblong, rounded at both ends, or the apex with a slight 
indication of a papilla which is often oblique, perfectly smooth, 
chestnut-brown, hardly constricted at septum, epispore about 3 /t 
thick, 35-45 x 22-29 li, average 37 X 25 li ; pedicel tinted 
yellowish, persistent, 30-40 x 6 li, but may reach a length of 96 /*. 

X. Mesospores few, similarly coloured to teleutospores, oval to flattened 

at apex, 28-35 x 22-25 li. 


Puccinia — Caryophyllaceae, Malvaceae. 177 

On leaves and stems of Kochia sedifolia F. v. M.; Kochia villosa, Lindl.; 
and Enchylaena tomentosa R. Br. 

Victoria — Dimboola, Nov. and Dec. (Reader). 

I have some of the material from Dimboola, portion of which was sent 
to Massee, and the golden-brown uredospores are seen along with the 
teleutospores. No uredospores were mentioned in the original description, a 
striking feature of which is the numerous germ-pores. The teleutospores 
have occasionally their septum lengthwise. 

(Plate XI., Figs. 95, 96.) 




109. Puccinia arenariae (Schum.) Schroet. 

Schroeter, Pilz. Schles., p. 345 (1872). 
Sydow,Mon. Ured., I., p. 553 (1903). 
Sacc. Syll. VII., p. 683 (1888). 

III. Teleutosori compact, pulvinate, roundish, scattered, often circinate, 

pale-brown to dark -brown. 

Teleutospores fusoid or clavate, apex pointed or rounded, 

often thickened (6-8 /u), base rounded or attenuated, slightly 
constricted at septum, smooth, pale yellowish brown, 
30-50 X 14-20 /j, average 35 x 16 /u, very rarely tricellular, 
48 X 16 ft; pedicels hyaline, persistent, equal to or longer than 
the spores, 60-100 p. 

X. Mesosperes occasional, similarly coloured to teleutospores, obovoid 

or somewhat wedge-shaped, rounded or obtusely pointed or 
truncate at apex and thickened up to 6 /w, generally resembling 
the upper cell of the teleutospore, with elongated pedicel, 22-29 
X 16-17 /x. 

On leaves and stems of Stellaria media Cyrill. 

Victoria — Gellibrand, Feb., 1896 (Hill). Gem brook Ranges, April, 

1904 (C. French, jun.). 

The germ-tubes of the sporidiola of this species have been known to enter 
the stomata of Dianthus barbatus L., and it seems to be the only known case 
in the genus Puccinia of such germ-tubes entering stomata. 

(Plate XII., Fig. 97.) 


110. Puccinia heterospora Berk, and Curt. 

Abulilon, Hibiscus. 

Berkeley and Curtis, Journ. Linn. Soc. X., p. 356 (1868). 
Cooke, Handb. Austr. Fung., p. 338 (1892). 
Sydow, Mon. Ured. I., p. 472 (1903). 
Sacc. Syll. VII., p. 695 (1888). 

III. Teleutosori minute, hypophyllous, soon naked, crowded in orbicular, 

glomerules, dark-brown, on determinate purplish or yellow spots. 

Teleutospores very variable in size and shape, smooth, thick- 
walled and more deeply thickened at apex, dark chestnut-brown, 
elliptic or oblong, not or scarcely constricted at septum, 
occasionally divided vertically, 20-38 X 17-28 ya; pedicel hyaline, 
slender, elongated, three to four times as long as spore. 

i 7 8 

Puccinia — Malvaceae. 

X. Mesospores much more common than teleutospores, subglobose or 

elliptical, apex more thickened, pale-brown, \i 

25-32 x 17-22 //. 

2 2 

\i diam., or 

On leaves of Abutilon crispum Sweet, and Hibiscus sp. (Bailey 1G ). 

Queensland — Gulf of Carpentaria and St. George, Oct., 1885 

(F. v. Mueller in National Herbarium). 

On Abutilon avicennae Gaertn. 

New South Wales — North-west part, Oct., 1887 (Bauerlen). 

There are only relatively few bicellular spores present, and at first they 
were overlooked, the fungus being named Uromyces pulcherrimus B. and C, 
^ind U. thwaitesii B.^and Br. 

The so-called mesospores are probably of the nature of one-celled 
teleutospores, and this species may be regarded as a transition stage from the 
unicellular Uromyces to the bicellular Puccinia. 

(Plate XII., Fig. 101.). 

Althaea, Lavatera, Malva, Plagianthus. 

ill. Puccinia malvacearum Mont, 

Montagne in Gay's Hist. Chili VIII., p. 43 (1852). 
Cooke, Handb. Austr. Fung., p. 338 (1892 
Sydow, Mon. Ured. I., p. 476 (1903). 
Sacc. Syll. VIL, p. 686 (1888). 

III. Teleutosori generally hypophyllous, brown to reddish-brown or 

orange, but grayish when spores are germinating, compact, round, 
pulvinate, elongate on the stems, scattered or crowded, seldom 
confluent, at first covered by epidermis, soon naked. 

Teleutospores fusiform, attenuated at both ends, apex sometimes 
rounded, slightly constricted at septum or not at all, apical 

smooth, yellowish-brown, rarely tricellular, 

17 \i\ pedicels firm, long, per- 

thickening slight, 

35-75 x 12-26 u, 

average 50 x 

sistent, occasionally septate, sometimes measuring 170 fx long. 

X. Mesospores occasional, same colour as teleutospores, somewhat 

fusiform, slightly thickened at apex, 40-45 X 17-19 p. 

On all green parts of Malva rotundifolia L., M. sylrestris 

rosea Cav 

Lavatera plebeia Si 

Plagianthus spicatus Benth 

L., Althaea 


(Berkeley 2 ). 

Victoria — 

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, but none were found. Spores occasionally 
3-celled, and reaching a length of 77 fi. 

This fun 


which was 

ally described by Monta 

spread the earth and occurs on both wild and 




cultivated species 
Malvaceous plants. Dr. Plowright 5 states that he has a specimen in 
herbarium from Melbourne, gathered as early as 1865 on Malva sylvestris 






Puficinia — Malvaceae, Geraniaceae. 179 

Puccinia malvace arum is one of the commonest of rusts, and the 
different stages in the growth of the spore are well seen in this species. A 
group may be observed springing from a common hypha, and at first they 
are without septa, but soon a transverse septum is formed, and, although 
specially looked for, I have never seen a partial septum as recorded by 
Fischer 5 . 

Germination occurs as soon as the spore is ripe, but some may hibernate. 
Occasionally the pedicel has a septum. Mr. French, Government Entomolo- 
gist, has observed this rust on Hollyhocks in a nursery near Melbourne as 
early as 1857. 

(Plate XII., Figs. 99, 100 ; Plate XV., Figs. 123-130 ; Plate F., Fig. 28.) 

112. Puccinia plagianthi McAlp. 



McAlpine, Proc. Rov. Soc. Vic. VII, N.S., p. 218 (1894). 
Sydow, Mon. Ured. I., p. 480 (1902). 
Sacc. Syll. XIV., p. 295 (1899). 

III. Teleutosori hypophyllous, sparingly epiphyllous, scattered, pul 

verulent, soon naked, up to 1-1 J mm. 

Teleutospores golden-brown, oblong to clavate, rounded at apex 
slightly attenuated towards base or rounded, slightly constricted al 
septum, very rarely three celled, epispore with reticulated markings 
38-50 x 17-23 /j, average 41 x 20 jj. ; pedicel hyaline, short, 
deciduous, but may attain a length of 60 ft. 

On leaves and flowers of Plagianthus sidoides Hook. Very common. 

Tasmania — Mount Wellington, southern slope, Aug. -April (Rod 



The spore is described as smooth by Sydow, but 

be covered 

with a fine irregular network 

(Plate XII., Fig. 98.) 



113. Puccinia geranii-pilosi McAlp 


II. Uredosori hypophyllous, chocolate-brown, scattered or sub-gr 

ious, orbicular, soon naked and surrounded by ruptured epidermis 
compact, up to 1 mm. diam., often on purplish-red spots or portion: 

of leaf becoming coloured. 

Uredospores yellowish-brown, elliptical to obovate, echinulate 

epispore slightly thickened at base, 25-30 X 19-22 /i. 

III. Teleutosori elongated, dark-brown to black, sometimes confluent 

erumpent, compact 

ly on peti 

Teleutospores at first intermixed with uredospores, oblong to 
oblong-clavate, chestnut-brown, smooth, slightly constricted at 
septum, sometimes short and stout and about as broad as long, 
32-48 x 22-32 /i, average 36 X 25 /* ; upper cell rounded at apex, 
not or slightly thickened ; lower cell rounded at base or occasion- 
ally slightly tapering; germ pore generally indicated at apex by a 
paler groove ; pedicel hyaline, deciduous, elongated, up to 80 /i long. 



Puccini a — Geraniaceae. 

X. Mesospores occasional, similarly coloured, subglobose, thickened at 

apex, 25-26 \i diam. 

On living leaves of Geranium pilosum Sol. 

Victoria — 

-Killara, Nov., 1902, II., March, 1903, II., £11. (Robin- 

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 ; pseudoperidia with expanded and 

lobed margin 

peridial cells subquadrate to oblong with striated 

margin, and punctate all over, firmly united, 28-32 x 16-22 ft. 

Aecidiospores pale orange, finely echinulate, elliptic to subquad- 
rate, 19-22 x 16 li, or 18-19 fx diam. 

II. Uredosori mostly on under surface of leaf, round, small, brownish, 

pulverulent, solitary or arranged in confluent groups. 

Uredospores subglobose to ellipsoid, yellowish brown, echinulate, 
22-24 li diam., or 22-27 x 19-22 u. 

in small confluent 

III. Teleutosori 

i intermixed, solitary or arranged 

groups, generally round, black, compact, surrounded by ruptured 

Teleutospores chestnut brown, clavate, constricted at septum, 

smooth, often obliquely pointed or rounded and thickened 

at apex (up to 9 li), 33-64 x 19-25 li, average 45 x 22 ^ ; 

lower cell usually paler in colour and attenuated towards base ; 

pedicels tinted, generally short, persistent, up to 38/.* long by 6-9 li 

X. Mesospores coloured 

like teleutospores, oblong to oval, bluntly 

pointed and thickened at apex, 33-38 X 16-20 fi 

Puccini a — Rutaceae. 


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 Murramurangbon 


It differs from P. geranii Corda, to which this species was referred bv 
Cooke in Grevillea, XXI., p. 39, in the uredospores being nearly twice as 
large, and in the much longer and broader teleutospores. 

There are two species of Puccinia already recorded upon the genus 
Pelargonium from S. Africa, viz., P pelargonii (Thuem.) Syd., and P. 
granular is Kalch. and Cooke. 

i In P. pelargonii the aecidia are hypophyllous, and not amphigenous, and 

'ores are rather shorter and narrower. In P. grauularis there 


(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 n), gen- 
erally rounded at base, 28-40 x 16-21 /u, average 35 x 20 yu, 
occasionally 3-celled and elongated, 52 x 25 li ; pedicel hyaline to 
yellowish, persistent, up to 120 /u long. 
X. Mesospores common, similarly coloured to teleutospores, ellipsoid to 

obovoid, rounded or bluntly pointed at apex and slightly thickened, 
30-34 x 17-18 fx. 

On Boronia spinescens Benth. 

West Australia — Avon, Oct., 1903 (Diels). 

A very interesting Leptopuccinia which projects from the cortex of the 
branches in thick, often confluent pustules, and partially surrounds them. 

A specimen was kindly forwarded by Hennings, and the mesospores were 
observed which are not given in the original description. The size of the 

teleutospores, as given by Hennings, is 20-32 x 15-18 ^i, by Sydow. 22-35 

X 16-19 fx, and I have made the average 35 x 20 /u, my measurements 

being corroborated by those in the photographs, which are also about 

35 x 20 it 

(Plate XIII., Fig. 107.) 


16. Puccinia correae McAlp 

Mc Alpine, Proc. Roy. Soc. Vict. VII., N.S., p. 215 (1894). 
Sydow, Mon. Ured. I., p. 462 (1903). 
Sacc. Syll. XIV., p. 297 (1899). 
III. Teleutosori hypophyllous, cushion-shaped, circular or interruptedly 

circular, dirty brown, scattered, soon naked. 



Puccinia — Rutaceae, Tremandraceae. 

Teleutospores yellowish-grey, long-stalked, elongated fusoid, con- 
stricted at septum, 44-60 X 17-20 ju, average 50 x 18 yu; upper 
cell elongated, tapering and rounded at apex ; lower cell tapering 

several times as long as 


base ; pedicel light 



X. Mesospores similarly coloured and stalked, ovoid, with truncated 

apex, 25-28 x 16-19 p. 

On leaves of Correa lawreitciana Hook. 

Tasmania — Mount Wellington, Dec. Coast near Trial Harbour, 

Jan., 1894 (Rodway'). 

The sori stand out very distinctly from the cinnamon-brown under 
surface of the leaves, causing corresponding circular depressions on upper 
surface, of a yellowish-green colour, 
the rust was very rare. 

Although the plant is 



(Plate XIII., Fig. 106.) 


117. Puccinia eriostemonis Mc Alp. 

III. Teleutosori hyphophyllous, dark-orange, ellipsoid, somewhat cir- 
cularly arranged, erumpent, surrounded by ruptured epidermis, 
up to 2 mm. long. 

Teleutospores clavate to ellipsoid, with finely granular orange 
contents, compact, smooth, generally rounded at apex, sometimes 
pointed, slightly thickened, usually tapering towards base, con- 

stricted at septum, 30-45 x 15-18 /.i 



32 x 16 y. 

pedicel persistent, hyaline, elongated, up to 125 /i long by 3 


A 1 

X. Mesospores occasional, similarly coloured to teleutospores, ellipsoid, 

thickened and bluntly pointed at apex, 30-32 X 13-16 y. 

On leaves of Eriostemon myoporoides DC. 

Victoria — Mt. Bernard, Nov., 1903 (C. French, jr.). Alps, near 

Bright, Dec, 1904 (C. French, jr.) 

Occasionally a spore occurs in which the septum is longitudinal. 

This species differs from P. correae McAlp 
instead of brown, the spores are smaller, with 
tents, and the.epispore rather thicker. 

(Plate XIII., Fig. 105.) 

, in the sori being orange 
finely granular orange con- 


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 

scattered or in orbicular groups, pulvinate, 

purplish margin, 

cinnamon- brown, \-\\ mm. diam. 

Puccinia — Violaceae. 

1 83. 

Teleutospores oblong or oblong clavate, obtusely rounded at 
apex, which may or may not be thickened, sometimes two-papillate, 
or occasionally somewhat crested, slightly constricted at septum, 
generally rounded at base, yellowish, smooth, 40-60 x 15-25 /u, 


48 X 20 fj. ; pedicels hyaline, persistent, up to 150 

X. Mesospores rare, similarly coloured to teleutospores, somewhat 
x oblong and bluntly apiculate at apex, 43 x 28 /*. 

On living leaves of Tremandra stelligera R. Br. 

West Australia— Near Perth, Oct., 1900 (Pritzel). 

Quite distinct from P. tremandrae Berk, from Ceylon. 

(Plate XIII., Fig. 108.) 


119. Puccinia hederaceae McAlp. 


O. Spermogonia arranged in small or large groups, crowded, honey- 

Spermatia hyaline, globose, minute, 2^-3 \x d 

I. Aecidia on all green parts of the plant and flowers, on both surfaces of 

leaf but mostly hypophyllous, disposed in circular or irregular 
groups ; pseudoperidia flat, about A mm. diam. with white, torn, 

re volute edges. 

Aecidiospores subglobose to shortly elliptical, finely warted, 
orange-yellow, about 16 /x diam., or 16-19 x 12—14 yu. 

II. Uredosori minute, brown, roundish, erumpent, scattered, soon 

naked, on both surfaces of leaf. 

Uredospores subglobose or shortly elliptical, golden-brown, 
ly echinulate, 2-3 germ-pores on one face, 25-29 X 18-22 u, 

average 25 x 21 ji and 22-25 /u diam. 

III. Teleutosori minute, black, roundish, erumpent, usually surrounded 

by ruptured epidermis, pulverulent. 

Teleutospores at first intermixed with uredospores, elliptic or 
oblong, rounded at both ends or slightly attenuated towards base,, 
hardly constricted at septum, distinctly warted all over but prom- 
inently on upper cell, chestnut-brown, usually with hyaline 
apiculus, occasionally unicellular, 29-49 x 17-22 //, average 35 
X 19 fi ; pedicel hyaline, deciduous, short. 

On living leaves of Viola hederacea Labill. 

Victoria — Near Melbourne, Oct., Nov., and Dec, 1885, 1886 

(Reader). Wandin, Dec, 1895. MurramurraDgbong Ranges, 

Nov., 1902, Dec, 1903, Jan., 1905 (Robinson). Mt. Black- 
wood, Dec, 1902. Alps, near Bright, Dec, 1904 (C. 
French, jun.) 
Tasmania — Cascade Rivulet, Dec, 1892 (Rodway 1 ). Mt. Drome- 
dary, Dec, 1894 (Rodway 1 ). Devonport, Jan., 1906 

On V. betonicifolia Sm. 

Tasmania — Ben Nevis, April, 1896 (Rodway). 

New South Wales — Monga (Bauerlen), recorded as Aecidium 

violae Schum. by Baker 1 . Richmond (Musson). 
Victoria — Murramurrangbong Ranges, Nov., 1902 (Robinson). 




Puccinia — Cruciferae. 

A fe 

celled teleutospores were ob 






pores, which they resembled i 

pore quite dist 
25-26 X 20 h i 

y respect 

d shap 


even having the germ 
small hyaline apiculus, 

Aecidium violae Schum. is ^ 
occurring on Victorian native violets, but it 

P. hederaceae McAlp. 

beneath the apex, ellipsoid 


by D 

Cooke in his Handbook as 
is the aecidium belonging to 


at first determined as P. aegra Grove by Winte 
but on comparing the two species they are seen to be decidedly different 


It is closely allied to P. violae (Schum 

teleutospores being 




ted all over. Sp 

but differs in the 

of P 

eel from Syd. Ured. Exs., No. 286 

d from 

were examn 

Kellerman's Ohio Fungi, No. 96, on Viola blanda. 

teleutospores was found to be 22-32 X 16-19 ju, and were generally 

The average size of 

smooth, although 
P. aegra Grov« 
mens I have met 
forwarded to D 






ded by Cooke for Australia; but all the speci 

th, including part of the same collection of 




tospores, and are therefore not this species 


d warted 

(Plate XIII., Figs. 109-112 ; Plate E., Figs. 24, 25.) 


120. Puccinia cruciferae Mc Alp. 

I. Aecidia amnhi^enous and o 


cup-shaped, §-| mm. diam.; pseudoperidia w 
edges, crenulate, slightly revolute ; pseudoperidial cells 

th whitish 


oblong, p 

all over, with 

ted margin, average 




Aecidiospores orange, 

subglobose, finely echinulate, 14-16 


III. Teleutosori on upper surface of basal leaves, associated with 

aecidia, a few together, round, bullate, compact, covered at first 

by raised grey ej>idermis, which 




to 1 mm. 

Teleutospores black in mass, dark chestnut-brown individually, 
oblong to clavate, smooth, slightly constricted at septum, bluntly 
pointed, rounded or truncate at apex and thickened up to 9 />/, 
tapering slightly or rounded at base, 37-62 X 18-25 p ; pedicels 
usually slightly tinted towards apex, persistent, up to 50 /x long and 
10 fi broad. 

X. Mesospores common, similarly coloured to, or paler than, teleuto- 



variously shaped, ellipsoid, oblong 

or ovate, pointed, 

rounded, or truncate at apex, and thickened up 

to 7 

25-34 x 14-20 /./, with pedicels equally long and similar to those 
of teleutospores. 

On Cruciferous plant. 

New South Wales 

It differs from P. cru 

Guntawang (H 

P. barb 



in having 



Cooke, in which the 


broadly lanceolate 



This is a portion of the original material which was named Aecid 

barbareae in Cooke's Handbook oj Australian Funyi but, on closely 
examining it, teleutosori were found. The host-plant is not Barbarea 
vulgaris R. Br., which is the only species of that genus in Australia. 

(Plate XXX., Fig. 266.) 


121. Puccinia hibbertiae McAlp. 



III. Teleutosori ruddy-brown to black, compact, soon naked, up to 

1 mm. diam., on under surface of leaves. 

Teleutospores pale yellowish-brown, oblong to somewhat clavate, 
smooth, constricted at septum, 35-41 X 18-21 ft; upper cell 
rounded at apex to somewhat conical, usually thickened (up to 
6 fi) ; lower cell generally rounded at base, sometimes slightly 
attenuated ; pedicel hyaline, persistent, elongated, slender, 
120-150 //long. 

X. Mesospores occasional, similarly coloured to teleutospores, generally 

fusiform and thickened at apex, sometimes elongated oblong, 

33-35 x 12-16 p. 

On Hibbertia sericea Benth. 

Victoria — Cheltenham, July, 1904 (Robinson). Oakleigh, Sept., 


The small size of the leaf together with the scanty sori render this 
species easily overlooked. The teleutospores are often noticed germinating 


(Plate XXX., Pig. 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. hamardx) on the native raspberry (Rubtie parvifolius) does considerable 
iamage, and the imported species — P. subcorticium — is spreading extensively 
m the sweetbrier. 

General Characters. — Spermogonia flattened or slightly raised, generally 
irranged in concentric circles. 

Aecidia after caeoma type, surrounded by a dense crown of inwardly- 
curved paraphyses. 
I Caeomosporcs with numerous distinct germ-pores distributed over the 

| entire surface. 

Uredosori likewise surrounded by paraphyses. 
Uredospores solitary on basidia, with numerous germ-pores. 
Teleutospores separate, pedicellate, consisting of from three to ten, 
superimposed cells, the uppermost of which has one to three germ-pore 
the others about three to four each, placed laterally. 




G 2 

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 upper 

surface of leaf, appearing as ruby coloured, minute, blister-like 
puckered swellings, with round central pore, and arranged in 
small concentric circles. 

Spermatia golden-yellow in mass, hyaline individually, minute, 
narrowly ellipsoid, embedded in mucilage, 4-6 x 2J-3 ix. 

II. Uredosori scattered on under surface of leaf, corresponding to 

spots on upper surface, very irregular in outline, ochraceous, 
also sparingly on upper surface of leaf, usually on veins, ellipsoid 
to elongated ellipsoid, surrounded by ruptured epidermis, and pale 
in colour. 

Uredospores globose or ellipsoid, epispore • thick, warty, dark 
yellow, 17-19 fi diam. or 21-26 x 17-19 fx ; paraphyses surround- 
ing them, at first colourless, finally yellowish-brown, variously and 
peculiarly shaped, generally curved and pointed at apex, 30-40 X 



III. Teleutosori scattered or gregarious, minute, dark brown, not 

rarely confluent, pulverulent. 

Teleutospores at first intermixed with uredospores, golden-brown, 
cylindrical, equal, rounded at apex, sometimes with a short, 
hyaline, conoid apiculus, 6-9 septate (8 very common), constricted 
at septa, smooth, usually two germ-pores on one face in each cell, 
or three altogether, 60-115 x 23-28 /u ; pedicels hyaline, cylin- 
drical, or a little inflated, sometimes uniseptate towards base, very 
long, up to 207 x 10-19 li. 

On living leaves of Native Raspberry (Rubus farvifolius L.). 

Victoria — Alps near Bright, Arthur's Creek, Flinders, Killara, 

Murramurrangbong Ranges, Myrniong, near Melbourne 
Phillip Island, Seymour, &c. 

South Australia — Mount Lofty (Tepper 

Tasmania — Huonville, Jan., 1892 (Rodway 1 ). Mount Wellington 

Jan., 1892 (Rodway). 


As early as 1886 this species was determined by Plowright and Winter 
from specimens sent by that veteran microscopist, Mr. F. Barnard, Kew, 

The teleutospores occur on the same leaves as the uredospores, and they 
germinate on the living plant immediately on ripening. The germ-pores in 
each cell may be seen very clear and distinct. There are two in each seen 
on one face, situated laterally an(I immediately below each of the septa, and 
by means of careful focussing it can be seen that there is one behind, thus 
making three in each cell. In the apical cell there are also two just above 
the septum, and a third may also be seen as in the other cells. 

Dietel ] remarks that there is only one germ-pore in each cell of this 
species, but, as indicated above, three can usually be seen with such dis- 
tinctness that they are capable of being photographed, as shown in the 


Phragmidium — Rosaceae 

i8 7 

Teleutospores were found germinating on the living plant. The promy- 
celium was stout and consisted of four cells with a basal cell, the contents 
being greyish like the sporidiola. The sporidiola are subglobose and 7-8 u 

This fungus does considerable damage to the native raspberry, causing 
the leaves to fall prematurely. 

(Plate XXVI., Figs. 234, 235 • Plate I., Fig. 38.) 

123. Phragmidium longissimum Thuem. 

Thuemen, Flora, p. 379 (1875). 



Cooke, Handb. Austr. Fung., p. 340 (1892). 
Sacc. Syll. VII., p. 750 (1888). 

Hamas'pora longissima Koern. Hedw. XVI., p. 23 (1877). 

II. Uredosori hypophyllous, scattered or gregarious, or even confluent, 

clear orange, surrounded by a dense layer of paraphyses. 

Uredospores globose or obovate or shortly elliptical, pale yellow, 
with thick epispore, 16 fi diam. 

III. Teleutospores on both surfaces of leaf, agglutinated in long fila- 
ments up to 6 mm. long, rather gelatinous when moist, but when 
dry twisted up and silky-fibrillose, 4 to 6 celled, commonly 4-celled, 
yellowish to cream-coloured, containing granular protoplasm, 
cylindric acuminate, with apex sharply pointed and hyaline, 
110-190 x 12-15 fx ; pedicel hyaline, tapering gradually towards 
base to a fine point, hollow, just slightly narrower than spore, 
10-12 a broad, and reaching a length of 500 u. 

On leaves of Rubus molnccanus L. 

Queensland — Eudlo Creek (Bailey 


This species was first observed on Rubus in S. Africa, and destroys 
the foliage of one of the blackberries indigenous to Queensland. Bailey 
kindly sent me specimens in which the teleutospores are germinating in 

Scattered over the leaves are groups of silky-looking twisted filaments, 
and on being teased out are seen under the microscope to consist of innumer- 
able spores with their long pedicels inextricably blended. 

The germinating spores are seen to have one germ pore in each cell, and 
the order of germination is basipetal. The promycelial branches are stout, 
elongated, and transversely septate, and they may either all arise on one 
side of the spore, or partly only on one side. The promycelial spores are 
subglobose, finely warted and 9-10 fj. diam. 

The teleutospores were very commonly 140 /< long, and one 6-celled in- 
dividual attained a length of 190 ja. As might be expected in such a long 
and variously septate spore there is considerable difference in the lengths 
recorded. It was originally given as 200-240 /u, then Massee reduced 


0-130 fi, and in the Queensland specimens I have found it to be 1 10—190 ^i 
The stalks are excessivelv long, and they may reach a length of 500 / 


The teleutospores agree with those of Phragmidium in bei 


more transversely septate, but they are agglutinated together into a 

cylindrical filamentous mass, and each cell only 



Phragmidium — Rosaceae. 

Massee 3 from an examination of fresh material comes to the conclusion 
that it is not a Phragmidium nor a Hamaspora, although he does not under- 
take the responsibility of forming a new genu?. 

Dietel 8 , 10 , on the other hand, considers that there are no substantial 
grounds for separating this species from Phragmidium, although he recog- 
nises a resemblance to Gymnosporangimn in the arrangement and structure 
of the teleutospores. 

(Plate XXXI., Fig. 272.) 

124. Phragmidium potentillae (Pers.) Karst. 

Karsten, Fung. Fenn., No 94 (1868). 
Winter, Rev. Myc. VIII., p. 208 (1886). 


Cooke, Handb. Austr. Fung., p. 339 (1892). 
Sacc. Syll. VII., p. 743 (1888). 

, Caeomata arranged in circular groups, often confluent, orange yellow 
to reddish. 

Caeomospores crowded in a short series, globose, ovate or ellipsoid, 
echinulate, 17-24 x 14-19 ft.] 

II. Uredosori hypophyllous, orange-red, roundish, scattered or 

gregarious, often confluent, surrounded by a crown of club-shaped 

Uredospores spherical, elliptical or ovate, yellowish, echinulate,, 
17-24 X 14-20 A /, average 18 x 15 /i. 

[III. Teleutosori orbicular, black, pulvinate, minute. 

Teleutospores oblong, 2-6 septate (usually 3-5), slightly con- 
stricted, obtuse, or obtusely apiculate above, slightly attenuated 
below, smooth, yellowish-brown, 50-70 X 20-22 \i ; pedicels 
colourless, firm, 100-150 \i long. 

On living leaves of Acaena sanguisorba Vahl. 

Victoria — Near Melbourne, 1886 (Reader). (Winter. 2 ) 
On Acaena ovina A. Cunn. 

S. Australia — Sept., 1898 (Quinn). 

The specimen from Victoria was originally determined by Winter, an 
portion of the same material, kindly supplied by Reader, shows only 

Darlucajilum Cast, is plentiful on some of the sori. 

125. Phragmidium subcorticium (Schrank) Winter. 

Winter, Die Pilze I., p. 228 (1884) 


Cooke, Handb. Austr. Fung., p. 339 (1892) 
"--- Syll. VII., p. 746 (1888). 


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 





II. Ured 



orbicular, scattered or crowded 


pots, yellowish red, minute 


P h y 

ds apex, and not particularly club-shaped 


Uredospores ellipsoid, ovoid or p 
eddish orange. 4-6 germ-pores on one face 

finely ech 

29 x 

III. Teleutosori hypophyll 

readily detachable. 

minute, scattered or 

6-20 p. 
ded. black 


th a colourless apiculus. 7-8 celled 

oblong, dark-brown, warted, obtuse at the apex 

face in each cell. 7 

x 3 


A 1 


3 germ-pores 


P t 

pale-brown towards apex, expanded in the lower half, 100-150 y 
long and broadened up to 22 u. 

On Rosa rubiginosa L., R. canina L., and R. laxa Ret 


Pascoe Vale, Oct., 1898 (C 


April, 1899 (H 
Oct. and Dec. 


il). Brighton, Nov., 1901, Aug., 1903, June 
904. March. 1905. II. 

S. Australia — Third Creek Garden, Mt. Lofty Range, Nov., 1899 

(Tepper). Nov., 1900 (Agricultural Bureau), IL, III. 

Tasmania — Launceston, Oct., 1902 (Littler). Hobart, Apr., 1903, 

IL, III. (Rodway). 

This species is now very common around Melbourne, particularly on the 
sweetbrier (R. rubiginosa). It has also become a pest in the nurseries, 
causing considerable damage to the dog-rose stocks (R. canina). Seedlings 
are attacked very severely in a favorable season and killed outright. This 
rust occurs in some of its stages, practically all the year round, but is less 
common in midsummer. The uredospores resemble the caeomospores in their 
size and shape, but may be distinguished by their long stalks (up to 56 /<) 
and their germ-pores. 

The mycelium of the caeoma may winter in the stems, so that by means 

of cuttings the disease was probably introduced into Australia. 



also have been introduced with the seeds of the sweetbrier, for that was 
among the first European plants to be imported into Tasmania and New 
South Wales, and used as hedges. 

(Plate XXVI., Figs. 229-233 ; Plate I., Fig., 37.) 



s senus is known here, and 

monly on certain leguminous plants. So far 


This genus produces uredo and teleutospores on 



other, but the latter h 

have been 
one host plant, 
s not yet been 

Europe there is a bladder-rust on the bark of 

and the aecidial 

found in Australia. 

sylvestris, and this has been proved to be the aecidial stage of Ci 

asclepiadeum (Willd.) Fr. As in so many other similar cases 

considered to be an independent form, and named Peridermium cornui 

Rostr. and Kleb. 

stage was 


General characters 
Aecidia with bladder-like 

— Spermogonia truncate to conoid 


ly opening pseudoperidia (P 


thout crerm- 

parated by d 


epispore always more or less regularly warty reticulated. 
Uredo-layer included in hemispherical pseudoperidum, which opens at ape 
by a narrow pore. 


Cronartium — Legnminosae. 

Uredospores echinulate, without germ-pores. 

Teleutospores unicellular, united into a cylindrical, vertical, horny column,, 
germinating in situ, surrounded at base by the uredospores. 
Sporidiola subglobose, hyaline. 
Australian species, 1. 


126. Cronartium jacksoniae P. Henn. 

Jacksonia, dec. 

P. Hennings, Hedw. XL., p. 127 (1901). 

O. Spermogonia honey-coloured, crowded or in lines, circular, flattened,. 

on branches and lower surface of leaves. 

Spermatia hyaline, avoid to ellipsoid, 5-6 X 2|-3 p. 
III. Teleutosori forming ruddy-brown, straight or curved, simple 

columns, crowded, 2-2 \ mm. high, with blunt apex, on young and 
swollen branches. 

Teleutospores ruddy-brown in mass, more or less colourless indi- 
dividually, elongated, generally somewhat tapering towards each 
end, with large vacuole, cylindrical to fusiform, contents granular, 
40-50 X 8-10 fi ; teleutospores germinate at once and produce 
promycelia with promycelial spores, which are hyaline, globose, 
and average about 6-7 \i diam. 

On Jacksonia scoparia R. Br. 

Queensland — Darling Downs (Law), (Bailey 2 ). Near Brisbane 

(Bailey 4 ). 

On Platylobium formosum Sm. 



Ranges, Nov. 

and Dec, 1902-3 

On Gompholobmm latifolium Sm. 

Victoria — Murramurrangbong Ranges Dec, 1903 (Robinson). 

On Bossiaea cinerea R. Br. 

Victoria — Near Melbourne, Nov., 1904 (Robinson) 
Tasmania — Bellerive, Dec, 1905 (Rodway). 

On Actus villosa Sm. 










It deforms the shoots of the 

plant on which it grows, 

causing mal- 

formation of branches and multiplication of shoots (witches' brooms), and 
so common in some parts that the great majority of the 

shrubs are 


The spermogonia were only found on Aotus and before the appear 

of the teleutosp 


the multiplication and deformation of the shoots 

had already been produced. On Bossiaea cinerea on one occasion Aeeid 
eburneum McAlp. was found on the same plant near Melbourne. 

The horn-like aggregation of teleutospores varies in length from 1-2 mm. 
on Aotus and Bossiaea to 2-2^ mm. in Platylobium and Gompholobium. 

The original specimen on Jacksonia scoparia in the National Herbarium, 
Melbourne, is given as a variety of C. asclepiadeum (Willd.) Fr., in Berkeley's 
handwriting, and is recorded in the Journal of the Linnean Society, p. 174 
(1883), with the remark that the only specimens are in a young state. The 
same species was subsequently found by G. H. Robinson very plentifully in 

on various species of Leguminous pi 

and the germiuating 

teleutospores were 


Melampsora — Hypericaceae. 191 


•ing its occurrence on indigenous Leguminous plants, together 
with the narrower teleutospores and colourless promycelial spores, as com- 
pared with G. asclepiadeum, I considered it a new species and had named it 
C. leguminum. But I find that Hennings in a footnote in Hedwi^ia XL. 
p. 127, states that the form occurring on Jacksonia is specifically distinct 
from G. asclepiadeum, and had better be named C. jacksoniae, the name now 



In this genus there are both heteroecious and autoecious species, but only 
the latter occur in Australia. The two species known are on native plants, 
but they also occur in Britain on the same genera. They only possess uredo 
and teleutospores, but the three spore-forms may occur on the same host 
{M. amygdalinae Kleb.), or only aecidia and teleutospores (M. saxi- 
fragarum (DC.) Schroet.). 

The teleutospores form flat irregular crusts, and are united to each other 
like the cells of a honeycomb. 

Rostrup first pointed out in 1883 the connection between this genus and 
Caeoma species, and now the relation between a number has been proved by 

General description. — Spermogonia forming a minute, hemispherical, 
flattened stratum, often subcuticular, otherwise subepidermal. 

Aecidia after the caeoma type, destitute of a pseudoperidium, usually 
without paraphyses, pustular. 

Uredospores usually without distinct germ-pores, included in more or less 
developed peridia. 

Teleutospores unicellular, rarely transversely divided, coalescing in a 
plane, firm, black or dark-brown stratum. Sporidiola globose, yellow, then 

Australian species, 2 


127. Melampsora hypericorum (DC.) Schroet. 

Schroeter, Brand und Rost., p. 26 (18 
Sacc. Syll. VII., p. 591 (1888). 


II. Uredosori mostly hypophyllous, scattered or subgregarious, at first 

bright orange, becoming pale, pulverulent, small, up to £ mm. 
diam, erumpent and surrounded by the ruptured epidermis. 

Uredospores subglobose to ellipsoid, finely verrucose, orange 
yellow, with two germ-pores on one face, 14-21 x 11-17 ji, 
average 17 x 14 /* ; paraphyses intermixed, numerous, hyaline, 
capitate, over-topping the spores, 50-68 p. long, with head 18-24 p. 

III. Teleutosori minute, dark-brown, flat, irregular, solitary or gre- 

Teleutospores yellowish-brown, intercellular, subcylindrical or 

prismatic, 25-37 + 6-9 u. 
On leaves and occasionally on stems of Hypericum j aponicum Thunb. 

Victoria — Murramurrangbong Ranges, Nov. and Dec, 1902-3, and 

M"t. Blackwood, Jan., 1903 (Robinson). Buffalo Mts. and 
Alps, near Bright, Nov. and Dec, 1903-4 (C. French, jr.). 

Near Melbourne, Nov., Jan 


Melampsora — Linaceae. 

The paraphyses are noted as absent by Dr. PIov 



of the Uredineae, but in these specimens they are particularly abundant, and 

form a striking feature of the uredosori 

Fischer 5 also found the paraphy 


had bee 

and he came to the conclusion with others 
l mistaken for the uredo, since the spor 

that the caeoma form 



of both the caeoma form without 

physes and the uredo form with paraphy 

so that the th 



species of Hyp 

Aecidium disseminatum Berk, is found he 

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). 

Uredosori scattered, rounded, orange, 
mm. diam. 


soon pulverulent, up to 1| 


Uredospores round or ovate, bright orange-yellow, echinulate, 

a length of 

pedicellate, 15-25 x 13-18 fx y exceptionally reaching 

28 fi ; paraphyses curved, markedly capitate, 17-20 fi thick. 

III. Teleutsori flattened, often confluent, reddish brown, then black, 


Teleutospores densely crowded beneath epidermis, cylindrically 
prismatic, intercellular, polygonal in section, 45-60 x 17-20 //, 
very occasionally two-celled. 

On leaves and stems of Linuni marginale A. Cunn. Widely distributed. 

Vic to 

Near Melbou 





Kergunyah, Killara, near Melbourne, Rutherglen, &c, Oct 


South Wales — Hume Ri 

1886 (Jeph 

Mudgee (Hamilton) ; Merilla, Oct., 1890 (Cobb 



South Australia — Murray River, 1890 (Tepper), (Ludwig 2 ) 


Near Waterworks, Hobart, Jan., 1892 (Rodway 1 ) 

On Linum usitatissimum L. wherever flax is cultivated 


Near Melbourne. 190 

Donald. Nov.. 1903. II 


Port Fairy, Jan., 1904, II., Ill 


South Australia 

New South Wales— Brungle, 1891 (Cobb 4 ) 

—Near Adelaide, 1889 (Crawford) 

Journal of 


1889. G 

ecords this 

t as 

being sent to him from South Australia by the late Frazer S. Crawford, who 


that it had destroyed a crop of flax near Adelaide, and was likely 

d prove a troublesome pest 

e y 

forms me by letter that 

Linum. and 

in Dakota it is a very abundant rust on all the wild varieties of Linum, 
is always more or less destructive in the flax crop. He further states that 
it was especially destructive in 1904, in many cases practically destroying 
crops which he had bred from seeds supposed to be immune to the wilt disease 
or dax-sick soil disease. 

Dr. 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. 



There are several imperfect forms which only occur in one stage and 
cannot with certainty be assigned to their proper genera. For such it is 
convenient to have a form-genus, which simply serves as a resting-place until 
their true affinities are determined. At first they were considered to be 
independent fungi and had generic names assigned to them. These form- 
genera are Caeoma, Aeeidium, and Uredo, and of the former there are *nlv 
two known for Australia, one of which was called an Aeeidium by Cooke 

and the other a Uredo by Berkeley. This is not surprising when one 
considers that the caeoma is not distinct from the aeeidium, as in those 
cases where the one merges into the other. Thus in Puccinia prenanthis 
(Pers.) Lindr., the aecidial wall is very imperfectly formed, so that the 
aecidia approach caeoma-forms. 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 Aeeidium and Caeoma. In 
the aecidial stage of Puccinia aucta (Aeeidium lobelias Thuem., A. micro- 
stomum Berk.), the peridial wall was sometimes absent, so that I described 
it at first as a Caeoma. 

Caeomata are generally considered to be stages in the life-history of the 
Melampsoraceae, but since true caeoma-forms occur in connexion with 
Puccinia, and species such as Melampsora tremulae Tul. have so-called 
caeoma-forms as Caeoma laricis (Westd.) Hart, with an investment of 
barren cells, the evidence for this connexion is weakened and the necessity 
for retaining this as a form-genus much reduced. In Saccardo's Sylloge 
Fungorum the isolated forms are treated as a sub-genus of Uredo with sub- 
"Catenulate spores. They have no special significance here, but elsewhere 
they form witches'-brooms. 

General Characters. — They are simply aecidia without peridia, the spores 
are produced in chains, with or without paraphyses, and accompanied by 

Caeoma-forms, 2. 


Tahernaem on tana 

129. Caeoma apocyni McAlp. 


I. Sori on both surfaces of leaf, minute, crowded, brownish, bullate. 

Spores yellowish, very variable in shape, ellipsoid, ovoid, pear- 
shaped, oblong or angular, with finely granular contents, 25-34 
x 20-26 /j. ; epispore hyaline, coarsely verrucose, 5 /x thick. 

On Tabernaemontana orientalis R. Br. 

Queensland — Brisbane, (Bailey 1 ). 

Mr. Bailey kindly sent me a specimen of this from his herbarium, 
which had been named by Dr. Cooke Aeeidium apocyni Schwein., as 
given in his Handbook of Australian Fungi, p. 341 (1902). It has no 
pseudoperidium, however, and is therefore a Caeoma, and since the spores do 
not apparently agree even with those of Caeoma (Aeeidium) apocynatum 
Schwein., it is constituted a new species. 



Aecidium — Scrophulariaceae 



130. Caeoma clematidis Thuem. 


Thuemen, Myc. Univ. No. 539. 

Berkeley, Hook. Journ. VI., p. 205 (1854). 

Cooke, Handb. Auatr. Fung., p. 344 (1892). 
Sacc. Syll. VII., p. 867 (1888). 

Uredo clematidis Berk. 

I. Sori hypophyllous, generally on pale-green portions of leaf, solitary o 

ged in circular groups, more or less rounded, pulvinate, ruddy 

brown at first and bright orange when b 


in rows, 




shape, ellipsoid 


oblong or polygonal, epispore hyaline, scolloped, 2 




of Clematis aristata R. Br., and C. microphylla DC 

Victoria— Near Colac, Sept., 1897 (H 






Phillip Island, Jan. 
, 1903 (Robinson) 

Near Melbourne, Feb. and July, 1904. Frankston, Jan., 1904, * 
Aug., 1904 (Robinson). 




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 


in Dr. Cooke's Handbook of Australian 

Fungi are now referred to their proper Puccinias. A. urticae is accepted 
as a stage in the life-history of Puccinia caricis, and described in connexion 
with that form. A. goodeniacearum, A. senecionis and A. violae are placed 
under their proper Puccinia or Uromyces, and the aecidium on the daisy 
(A. bellidis) is now found to belong to P. distincta McAlp., and to have no 
connexion, as in Britain, with P. obscura on Luzula. The most noteworthy 
aceidium is that found on a grass (A. danthoniae), and constitutes the second 
known example. They often cause deformation of the branches known as 
witches' brooms, and various swellings, and in some instances compound 
galls are formed up to 5 cm. across (A. englerianum). One species causes 
thickening and distortion of the leaves in Veronica, and nettle stems are 
often abnormally swollen from the same cause. 

General Characters. — Pseudoperidia cup-shaped or urn-shaped, rarely 
cylindrical, with margin often crenate or laciniate and re volute. 

Spores, globose or angular, commonly orange yellow, growing in chains* 

Aecidium-forms, 15. 



131. Aecidium disciforme McAlp. 

O. Spermogonia on under surface of upper and younger leaves, honey- 
coloured, flat, numerous, minute. 

Spermatia hyaline, elliptical, 5-6 /x long. 

Aecidium — Plantaginaceae 195 

I. Aecidia on both surfaces of leaves, but mostly on under, bullate, 

brownish, resembling dotted discs, often confluent, at first entirely 
enclosed, ultimately opening irregularly by small pore, 1-1 A mm. ; 
peridial cells hyaline, loosely adherent, subglobose, elliptic or 
lozenge-shaped when united, with striated margin, 25-30 fj. 

Aecidiospores orange-yellow, subglobose to elliptic, 16-20 a diam 
or 19-22 x 14-16 /u. 

On Veronica gracilis R. Br. 

Tasmania — Sandy Bay, Hobart, Aug., 1896 (Rodway). 

On Veronica calycina R. Br. 

New South Wales— Kogarah, Oct., 1900 (Maiden). 

This species causes thickening and distortion of the leaves. It is quite 
distinct from A* veronicae Berk., of which there is a portion of the original 
material in the National Herbarium, Melbourne. The latter forms a regular 
cup with fimbriate margins, while the former only opens by a minute pore. 
The spores are also quite different. 

132. Aecidium veronicae Berk. 


Berkeley in Herb., Grev. XL, p. 97 (1883). 
Sacc. Syll. VIL, p. 814 (1888). 

I. Aecidia numerous, crowded, semi-immersed, on under-surface of leaf 

and a few scattered on upper surface, \ mm. diam.; pseudo- 
peridia cup-shaped, becoming discoloured and dark-brown with 
age, with fimbriate margins ; peridial cells, oblong to angular, with 
very broad striated margin, average 32 x 25 it. 

Aecidiospores dark yellow, variously shaped, ellipsoid, oblong or 
angular, with finely-granular contents, 24-30 x 16-20 fx ; epispore 

hyaline, finely verrucose, about 2 it broad. 

On Veronica sp. 

Victoria — (F. v. Mueller). 
Tasmania — (Rodway). 

There is an old specimen in the National Herbarium which had been 

submitted to Berkeley by the late Baron von Mueller. There are plenty of 

■ spores still in the cups, but probably the thoroughly mature ones have fallen 


(Plate XXXIX., Fig. 287.) 



133. Aecidium plantaginis-variae McAlp 


0. Spermogonia on both surfaces of leaf, but mostly on upper, honey- 

coloured, numerous, 160-180 li diam. 
Spermatia hyaline, elliptical, 4-5 li long. 

1. Aecidia on both surfaces, sub-gregarious or gregarious ; pseudoperidia 

cup-shaped, outstanding, with recurved and lobed margins; 
peridial cells oblong to lozenge-shaped, or polygonal, with broad 

striated margin, 35-40 li long. 

Aecidiospores bright orange-yellow, elliptic to oblong, finely 
echinulate. 22-25 x 16-23 u, sometimes reaching a length of 28 u. 


Aecidium — Gentianaceae, Comfo sitae. 

On leaves of Plantago vnria R. Br. 

Victoria — Murramurrangbong Ranges, Nov., 1902 (Robinson). 

Mt. Blackwood, March, 1905 (Cyril Brittlebank). 
New South Wales — Guntawang, near Mudgee (Hamilton). 
Tasmania — Knocklofty, Oct., 1895 (Rodway 1 ). 

This species is 

given as A. 

plant aginis Ces., in Cooke's Handbook of 

Australian Fungi, but there are no spots on the leaves, and the spores are 
echinulate not tuberculate, as well as considerably larger. 

The Victorian specimens were rare, and the fungus was much overrun by 

Darluca filum Cast. 

(Plate XXXIX., Fig. 288.) 


134. Aecidium nymphoidis DC. 


De Candolle, Fl. fr. II., p. 597 (1805). 
Cooke, Handb. Austr. Fung., p. 341 (1892) 
Sacc. Syll. VII., p. 809 (1888). 

I. Aecidia on upper surface of leaf, gregarious, disposed without order 

on rounded spots, or in concentric zones ; pseudoperidia scutelliform, 
margin scarcely prominent, entire, or irregularly laciniate. 

Aecidiospores orange-yellow, angular, delicately warted, 12-20 
ju diam. 

On Limnanthemum indicum Thw. 

Queensland — Near Brisbane, 1884 (Bailey 

This aecidium was first found on the leaves of Limnanthemum nymphoides 
Hoff. and Link and its connexion with Puccinia scirpi DC, was first sug- 
gested by Chodat 1 . Then Bubak 1 , by means of cultures, proved the 
connexion, but until the teleuto stage is found 
aecidium stage will be given. 


in Australia only the 

Specimen not seen. 


135. Aecidium cymbonoti Thuem. 


Thuemen in Muell. Supp. Phvt. Austr. XI., p. 96 (1880). 
Sacc. Syll. VII., p. 833 (1888). 

0. Spermogonia minute, honey-coloured, in groups, surrounded 

aecidia, 130-160 /i diam. 

Spermatia minute, hyaline, ellipsoid, 3 fx long. 

1. Aecidia on upper surface of leaves, markedly circinate, about ^ mm. 

diam.; pseudoperidia yellowish, with fimbriate margin ; peridial cells 
considerably overlapping each other, elongated oblong to slipper- 

ma rgm 

and reticulate all over, 28-38 X 

shaped, with striated 
16-17 ^. 

Aecidiospores orange-yellow, ellipsoid to sub-globose, finely 
echinulate, 13-18 x 13-15 fi. 



Aecidium — Compositae. 107 

On Cymbonotus lawsonianus Gaudich. 

Victoria— Dookie, Jan., 1892 (Pye). Dimboola, May, 1896 

(Reader). Preston, April, 1900 (C. French, jun.). Myrniom 
(C. C. Brittlebank). J * 

Some of the specimens were overrun with Darluca filum Cast. 
There is no clue to the description of this species, but I have drawn oul 
the above from abundant material. 

(Plate XXXIX., Fig. 290 

136. Aecidium monocystis Berk. 


Berkeley, Flor. N.Z. II., p. 196 (1855). 

McAlp., Proc. Roy. Soc. Vic. VII., N.S., p. 218 (1894) 

Sacc. Syll. IX., p. 319 (1891). 

I. Aecidia solitary, relatively large (J mm. diam.) persistent on the upper 

surface of the leaves towards their tips, surrounded with a tough 
border ; pseudoperidia white, with margin toothed ; peridal cells 
somewhat wedge-shaped to angular oblong, appearing hexagonal 
when united, covered with projecting points, 50 x 22 p 

Aecidiospores pale orange-yellow, ellipsoid, finely verrucose, 

25-30 x 19-22 H . 

On leaves of Abrotanella J or sterioides Hook. f. 

Tasmania — Summit of Mt. Wellington, Feb. 1891 (Rodway). 

This is a very characteristic species, occurring solitary at the tips of the 
leaves. It was first recorded on Phyllachne (H el ophyllum) coleusoi Berggr. 
from New Zealand, but L. Rodway, Government Botanist of Tasmania, has 
grave doubts as to the correct determination of the host-plant. He is of 
opinion that this rust is attributed to New Zealand by mistake, and that 
it was really found in Tasmania. He informs me that this Aecidium is 
abundant on every tuft of A. for sterioides, which grows in profusion near 
Hobart, and is not likely to have been overlooked by the early botanists. 
The coloured drawing of the plant in the New Zealand Flora certainly closely 
resembles Abrotanella, and I have forwarded a specimen of the plant to the 
Director of the Royal Gardens, Kew, for comparison with the original. It 
is not likely that the same Aecidium occurs on a Composite and a 

Stylidiaceous plant. 

(Plate XXXIX., Fig. 291.) 


137. Aecidium oleariae McAlp 

I. Aecidia crowded on stems and leaves, generally running in lines 

lengthwise, about 1 mm. ; pseudoperidia round to elongated, with 
lacerated margins ; peridial cells hexagonal, striated, 35-42 p long. 
Aecidiospores orange, elliptic to oblong, with granular contents, 
22-26 x 14-17 /a. 

On stems and leaves of Olearia axillaris F.v.JVL 

Victoria — Port Fairy, June, 1899 (Robinson). 

The aecidia occur most commonly on the stem, and are deeply imbedded 
in the tissues of this coast shrub. 


Aecidium — Rubiaceae, Leguminosae. 


Can thium 

138. Aecidium plectroniae Cooke. 

Cooke, Grev. X., p. 124 (1882) 

Bailey, Queensland Agr. Journ. IV., p. 284 (1899). 
Sacc. Syll. VII., p. 795 (1888). 

O. Spermogonia on upper surface of same spots which bear the aecidia, 

sometimes accompanied by one or a few aecidia. 

I. Aecidia hypophyllous, rotund, in small clusters on orbicular dis 

coloured spots ; pseudoperidia somewhat prominent, whitish, mar 

ovate to somewhat quadrate 

gins sub-entire ; peridial cells 
punctate all over, and with striate mar 


Aecidiospores yellow, finely 
polygonal, 25-' 

ub-globose to ellipsoid 

6 x 19 



2-28 u diam 

On leaves of Canthium coprosmoides F. v. M. 

Queensland — Endeavour River (Roth) (Bailey, 21, 25 

The host-plant is sometimes placed under the genus Plectronia. 

This species was first found on Canthium (Plectronia) guenzii Sond., 
from Natal, and the spores were undescribed, but I have been able to 
redescribs it from specimens kindly sent from Queensland by Bailey. 



139. Aecidium eburneum McAlp. 

McAlpine, Proc. Rov. Soc. Vic. VII., N.S., p. 218 (1895). 
P. Hennings, Hedw. XL., p. 352 (1901). 
Sacc. Syll. XIV., p. 375 (1899). 

A. bossiaeae P. Hennings, Hedw. XL., p. 352 (1901). 

I. Aecidia mostly hypophyllous or on stems and legumes, scattered or in 

dense clusters, bright orange, average £ mm. diam.; pseudoperidia 

cup-shaped, becoming ivory-coloured then brownish, with reflected 

finely toothed margin ; peridial cells polygonal, reticulated, 24-31 /x 

Aecidiospores ellipsoid, bright orange, very finely echinulate, 
18-28 x 12-17 /i. 

On Bossiaea drier ea R. Br., stems, leaves, flower-stalks, calyx, and 


Victoria — Near Melbourne, Sept.-Nov. (Barnard, Robinson, &c). 
Tasmania — Bellerive swamp (Rod way !), and Dec, 1905. 

On leaves of Bossiaea linophylla R. Br. 

West Australia 

— King George's Sound, July, 1901 (Pritzel). 

On fruit of Bossiaea rhombifolia Sieber and B. microphylla Sm. 

New South Wales — Richmond, Nov., 1896, and Falconbridge, 

Oct., 1904 (Maiden). 

On fruit of Bossiaea heterophylla Vent. 

New South Wales, Sep., 1896 (Maiden). 
I have revised the description of A. eburneum from fresh specimens, and 

have no doubt but it is the same as Hennings' species, a specimen 






Aecidium — Leguminosae. 190 

he kindly sent me. The spores in both cases are very finely eehinulate, and 
in the West Australian specimens they are 18-23 x 15—18 /u, while in the 
other they are 18-28 x 12-17 p. 

On the stems of B. cinerea there are oval to lenticular swellings, some- 
times half an inch long, separate or close together, with dark purplish 
border, and often becoming depressed in the centre. On the leaf-stalks the 
swelling may completely surround them, and on the leaf there are brown 
raised orbicular spots, with aecidia on both surfaces, but sparingly on upper. 
On the flower-stalks the swellings are similar to those on the leaf-stalks, and 
on the legumes large circular or irregular patches are formed on margins and 
both valves. 

(Plate XXVII., Fig. 239.) 


140. Aecidium platylobii McAlp. 

0. Spermogonia honey-coloured, minute, scattered, either associated 

with or on opposite side to aecidia. 

Spermatia very numerous, hyaline, cylindrical, 3 X 1 jx. 

1. Aecidia on pallid spots or on swellings on pods, amphigenous, in 

small groups or in dense clusters, deep orange ; pseudoperidia 
cylindrical, white, with small portion inserted in matrix, margin 
finely toothed, up to 1 mm. long. 

Aecidiospores orange, elliptic to sub-angular, finely warted, 
21-25 x 14-18 /u. 

On leaves and legumes of Platylobium formosum Sm. 

Victoria — Murramurrangbong Ranges, Nov., 1902, Dec. 1903, and 

Jan. 1905 (Robinson). 

It is closely related to Aecidium solenii/orme Berk., but until further 
stages are found it will be retained as a distinct species. 

While the bright orange aecidia are very conspicuous and very plentiful 
on both surfaces of the legumes, they are very rare upon leaves, ha\ 


hitherto only been found on the first or second pair of leaves of young 


The mycelium bearing the aecidia penetrated the fruit cover and entered 
seed, so that the young plant'is probably infected from the start. 

(Plate XXVII., Figs. 237, 238.) 


141. Aecidium soleniiforme Berk. 

Berkeley, Fl. Tasm. II., p. 270 (1860). 
Cooke, Handb. Austr. Fung., p. 340 (1892). 
Sacc. Syll. VII., p. 788 (1888). 

I. Aecidia on brown orbicular spots, sometimes densely crowded, par- 
ticularly when on fruit ; pseudoperidia cylindrical, elongated up to 
^ mm., white, radiately laciniate at margin. 

Aecidiospores rather angular, orange, 25-26 /u diam. 

On leaves and fruits of Goodia lotifolia Salisb. 

Victoria— Nov., 1895 (French). Gellibrand River, Dec, 1895 

Tasmania— 1 8 6 (Archer). (Berkeley 1 ) 

The aecidia, as the specific name denotes, are rather tubular at first. 

(Plate XXXIX., Fig. 293.) 


Aecidium — Amarantaceae, Hyfericaceae 


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 fi diam. 

On living leaves of Deeringia relosioides R. Br. 

Queensland — Gladfield (Gwyther), (Bailey. 12 ) 

The colour of the spores could not be determined in the dried specimens. 

Only the name of this fungus was given in the Annual Report, and Dr. 
Cooke has kindly furnished me with the description in MS. 

Uromyces deeringiae Syd. occurs on Deeringia indica in Java and the 
Philippine Islands, but the aecidiospores are 22-28 /x diam., so that they 
are quite distinct from the present form unless there has been a mistake in 
the transcription of the size of the spores. The host in both cases is the 
same according to the Index Kewensis. 

(Plate XXXIX., Fig. 294.) 


1 1:3. 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 fi. 

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). 

Mc Alpine, Proc. Linn. Soc. N.S.W., XXIV., p. 301 (1899). 

Sacc. Syll. VII., p. 602 (1888). 

0. Spermogonia arranged in small groups, honey-coloured. 

1. Aecidia distributed on upper surface of leaf and often confluent, abou 

mm. diam. ; pseudoperidia white, margin laciniate ; peridial 

cells pentagonal or hexagonal, margin finely streaked, 28-32 ^i. 

Aecidiospores bright orange, angular to sub-globose or oval, finely 
echinulate, average 22-23 x 17-20 ^. 

On living leaves of Caltha introloba F. v. M. 

New South Wales — Mt. Kosciusko, Jan., 1S99 (Maiden). 

I have provisionally named this as above, although Greville describes the 
aecidium as hypophyllous and on the petioles, while here it was epiphyllous, 
and only on the blade of the leaf. However, the spores and pseudoperidial 
cells quite agree with British specimens. The host-plant is indigenous to 
Australia, and was found in a rocky creek, on eastern side of Mueller's 
Peak, Mt. Kosciusko, at a height of about 6,500 feet. This was the only 
micro-fungus found there by Mr. Maiden. The Puccinia-stage of this 
fungus occurs in Europe and America, and has hitherto only been found on 
the marsh marigold (Caltha palustris L.) ; until that stage has been dis- 
covered here, it cannot be definitely stated that the two fungi are the same. 

(Plate XXXIX., Fig. 295.) 



145. Aecidium ranunculacearum DC. 

De Candolle, Fl. fr. VI., p. 97 (1805). 
Cooke, Handb. Austr. Fung., p. 340 (189 
Sacc. Syll. VII., p. 776 (1886). 

I. Aecidia hypophyllous or surrounding lobes of leaf and on stalk, 

densely crowded, very bright orange, often causing a swelling ; 
pseudo-peridia cup-shaped to cylindrical, whitish, with margin 
brittle and lacerated, 1 mm. high ; peridial cells somewhat quadrate 
and tapering to a point where they dovetail into each other, 
with striated margin, 27-28 x 18-22 ^u. 

Aecidiospores orange-yellow, polygonal to oblong or ellipsoid, 
very finely echinulate, 17-30 x 14-20 /u, average 22 x 17 yu. 

On Ranunculus parvijlorus L., R. rivularis Banks and Sol., R. lappaceus 
Sm. and R. gunnianus Hook. 

Victoria— Mount Emu Creek, 1854. Omeo, Nov., 1882 (Stirling). 

Ardmona, 1896 (Robinson). Near Dimboola, Nov., 1897 
(Reader). Murramurrangbong Ranges, Nov., 1902 (Robin- 
son). Buffalo Ranges, Dec, 1904 (C. French, jun.). 

On Ranunculus sp. 

New South Wales — Guntawang and Mudgee (Hamilton). Rich- 
mond (Musson). 
Tasmania— St. Patrick's River, Nov., 1844 (Gunn), (Berkeley 1 ). 

varies some- 


This aecidium occurs on several species of Ranunculus, and varies 
what in general characters, although the aecidiospores all agree in be v 
very finely echinulate. As the result of cultures, this has been found to be 


02 Uredo — Gramineae, Cy-peraceae. 

a composite species, the aecidiospores infecting various grasses on which 
they produce uredo and teleutospores. Until similar experiments are 
carried out here, the different aecidia, occurring on species of Ranunculus 
must be grouped together. 

Darluca Jilum Cast, was found associated with the aecidia. 

(Plate XXXIX., Fig. 296.) 

UREDO Pers. 

This stage probably belongs to teleutospore-forms which have not been 
observed, and which may only rarely be produced. But it is also quite 
possible that some may possess a perennial mycelium, and dispense with the 
necessity of forming teleustopores . Several in Dr. Cooke's Handbook of Aus- 
tralian Fungi have been referred to their proper genera. U. notabilis Ludw., 
belongs to Uromycladium notabile, and U. armillata Ludw., to Puccinia 
juncophila. U. clematidis Berk., has already been shown to be a Caeoma, and 
U. cichoracearum DC, as determined by Cooke, belongs to U. bidentis P. 

General characters. — Sori without pseudoperidium, covered or erumpent, 
for the most part orange-yellow, rather pulverulent, and spores produced 
singly on the terminal ends of hyphae, never smooth. Germination by germ- 
tube, which enters the host-plant through stomata. 

Uredo-forms, 13. 




146. Uredo kuehnii Krueg 


Krueger, Bericht. Zuck. Java IX., p. 117(1891). 
Cobb, Agr. Gaz. N.S.W. IV., p. 799 (1893). 
Sacc. Syll. XL, p. 182 (1895). 

Uromyces kuehnii Krueg. 

II. Sori mostly hypophyllous, linear, at length bursting through epidermis 

and forming elongated brownish or blackish narrow streaks, single 
or confluent, with clavate, brown paraphyses. 

TJredospores globose to ellipsoid or piriform, brown or orange, 
rather thin-walled, echinulate, 47-53 x 28-35 /j, ; pedieels hyaline, 
clavate, somewhat shorter than spore. 

Common on leaves of Saccharum ojjfiicinarum L. 

New South Wales — Clarence River (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 u. 

O" *"" "~& 



147. Uredo scirpi-nodosi McAlp 


II. Sori bullate, elongated, splitting lengthwise, running in lines on stem, 

1^-2 mm. long. 

TJredospores brown, subglobose, ellipsoid to oblong, variable in size, 
25-32 jx diam., or 27-36 X 19-25 /u; epispore echinulate, often 

3 a thick. 




Uredo — Liliaceae. 


On Scirpus nodosus Rottb. 

j Victoria— Mordialloc, Dec, 1885 (Reader) (Winter 2 ). 

[ This specimen is given as Puccinia rimosa (Link) Wint., by Winter, but 

I on carefully examining some of the original material, there are coarsely 

I echinulate uredospores and nothing else. It is evidently not that species 

described by Winter in Hedwigia, p. 28 (1880), and since it does not agree 

with the uredo stage of Puccinia scirpi DC. of which I have examined 
specimens from Sydow's Ured. Exs. 685 on Scirpus lacustris L., I have 

ed it as abo\ 

(Plate XXVIII., Fig. 241.) 


148. Uredo anguillariae Cooke. 


Cooke, Grev. XIV., p. 11. (1885). 

Cooke, Handb. Austr. Fung., p. 343 (1892). 

Sacc. Syll. VIL, p. 840 (1888). 

II. Sori on both surfaces, gregarious, elliptic, bullate, long covered by the 


Uredospores globose or globosely oval, smooth, brown, epispore 
thin, 20-22 x 20 /u, on short deciduous hyaline pedicels. 

On leaves of Wurmbea (Anguillaria) dioica F. v. M. 

New South Wales — Guntawang (Hamilton). 

It is curious to note that Cooke and Massee have also described a 
Puccinia with uredo and teleutospores on this plant, the uredospores being 
25-28 x 15-18 p. 


149. Uredo geitonoplesii Mc Alp 

G eitonoplesium 

II. Sori on under surface of leaves on vinous patches, in small groups 

minute, elliptical, ultimately rupturing epidermis. 

Uredospores yellowish to yellowish brown, elliptical to ovoid, 
very finely echinulate, 19-27 X 17-19 p. 

On G eitonoplesium cymosum A. Cunn. 

Victoria — Orbost, July, 1901 (Pescott). 


150. Uredo schelhammerae McAlp. 

II. Uredosori epiphyllous, yellowish-brown, round to ellipsoid, compact, 

scattered or in groups, occasionally confluent, soon naked and sur- 
rounded by ruptured epidermis, ^ mm. diam. or up to 1 mm. long. 
Uredospores golden yellow, ellipsoid to obovate, very finely 
echinulate, usually with two germ-pores on one face, 22-26 x 
15-20 ju, average 24 x 18 yu. 

On Schelhammera undulata R. Br. 

New South Wales— Kurrajong Heights, 1892 and 1903 (Musson). 

204 Uredo — Rhamnaceae, Stylidiaceae, Compositae 


Spyridium, Pomaderi 

151. Uredo spyridii Cooke and Mass. 

Cooke and Massee, Grev. XV., p. 99 (1887). 
Cooke, Handb. Austr. Fung., p. 343 (1892) 
Sacc. Syll. VII., p. 861 (1888;. 

II. Sori hypophyllous, punctiform, generally scattered, sometimes con- 
fluent, ochraceous to snuffy brown, soon bursting through epidermis, 

Uredospores yellowish to orange-yellow, ellipsoid to obovate, 
distinctly echinulate, with coarsely granular contents, 22-31 x 
17-22 /u; paraphyses enveloping sori, long, curved, tufted, hyaline 
to pallid, generally slightly swollen at apex. 

On leaves of Spyridium parvifolium F.v.M. 

Victoria— Oakleigh, June, 1886 (Watts). Sept., 1887 (Mrs. 

Martin). Murramurrangbong Ranges, Dec, 1903, and Jan., 
1905 (Robinson). 

On Pomaderris apetala Labill. 

Victoria — Murramurrangbong Ranges, Jan., 1905 (Robinson). 
Tasmania — Devonport, Jan., 1906 (Robinson). 

On both genera of host plants the sori are surrounded by paraphyses. 
The rust is very plentiful on the leaves, and, although a large quantity of 
material has been examined at different seasons of the year, only the uredo- 

ge has been found 

(Plate XXVIII., Fig. 245.) 


152. Puccinia stylidii McAlp. 


II. Sori on both surfaces of leaf, solitary or sub-gregarious, elliptic, 

bullate, and splitting raised epidermis, sometimes confluent later- 
ally, reaching 1 mm. or more in length. 

Uredospores orange-yellow, elliptic to subglobose, echinulate, with 
solitary germ-pore on one face, 22-24 X 16-18 ju, or 22-24 jx diam. 

III. Teleutosori, p. 210. 

On leaves of Stylidium gramini folium Sm. 

Tasmania — New Waterworks, Hobart, Nov. 1892, II. (Rod way). 
Darluca filum, Cast., literally covered some of the sori. 

(Plate XXVIII., Fig. 242.) 



153. Uredo bidentis P. Henn. 

Hennings, Hedw. XXXV., p. 251 (1896) 

Sacc. Syll. XIV., p. 395 (1899) 

II. Sori scattered or densely gregarious, amphigenous, most numerous 

on under surface, minute, pulvinate, then flattened or somewhat 
patelliform, ochraceous, surrounded by the epidermis. 

Uredospores ovoid, ellipsoid or subglobose, yellow or pale brown, 
epispore pale chestnut-brown, 2-3 li thick, finely echinulate or almost 
smooth, 25-35 X 22-27 fx, in exceptional cases 41 il long; para- 
physes numerous, hyaline, clavate, 50-60 u long. 


Urcdo — Co?npositae 


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 
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 /t thick, 
14-18 n diam. 

On leaves of Crepis japonica Benth. 


Specimen not seen. 


155. Puccinia oleariae McAlp. 

Mc Alpine, Agr. Gaz. N.S.W. VI., p. 756 (1895). 
Sacc. Syll. XIV., p. 278 (1899). 

Uromyces aster is McAlp . 

II. Uredosori hypophyllous, pulvinate, scattered, bright orange. 

Uredospores oval or irregularly pear-shaped to spathulate, 
stalked, bright orange, apiculate at apex and somewhat thickened, 
with three equatorial pores on one face, 39-51 x 19-24 fx; epispore 
longitudinally striate, average thickness 1 J f ; pedicels relatively 
short, hyaline, and continuous with spore. 

III. Teleutosori similar but brownish. 

Teleutospores intermixed with uredospores, bright orange, 
oblong to oblong-clavate, smooth, fragile, constricted at septum, 
rounded at apex, generally tapering slightly towards base, 62-90 
x 18-28 fx; pedicel hyaline, persistent, short, up to 56 it long. 

X. Mesospores occasional, similarly coloured, elongated ellipsoid, 

rounded at both ends, 60-68 x" 18-22 p. 

On leaves of Olearia aryophylla F. v. M. (Aster argophyllus Labill.). 

Tasmania — Near Hobart, November, II. (Rodway 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 Fuccinia. 

(Plate XXVIII., Fig. 249.) 

2o6 Uredo — Proteaceae, Crassulaceae, Leguminosae. 



156. Uredo angiosperma Thuem. 

Thuemen, Symb. Myc. Austr. IV., No. 95 (1880). 
Cooke, Handb. Austr. Fung., p. 343 (1892). 
Sacc. Syll. VIL, p. 840 (1888). 

II. Sori on both surfaces, large, commonly disposed about a circle, 

covered by the splitting, torn, and elevated epidermis, powdery, 


Uredospores oval or ellipsoid, rounded at the apex, somewhat 
narrowed at the base, 45 x 30 fi ; epispore smooth, pale brownish, 
5-7 f.i thick. 

On living leaves of Hakea sp. 

West Australia— Oct., 1877 (F. v. Mueller) (Thumen 2 ). 

Specimen not seen. 



157. Uredo tillaeae McAlp, 



158. Uredo bossiaeae McAlp. 

II. TJredosori on both surfaces of leaves but mostly on under, chocolate 

brown, erumpent, surrounded by ruptured epidermis, compact, 
scattered or aggregated, sub-globose, 1-1^ mm. diam. 

Uredospores elliptic, finely echinulate, golden-brown, epispore 
about 3 fA thick, 24-31 x 17-21 /u. 

On leaves of Bossiaea prostrata R. Br. 

Tasmania — Hobart, April, 1893 (Rodway). 

An Aecidium has been found on species of Bossiaea in West Australia, 
Victoria, New South Wales, and Tasmania, but there is no evidence to con- 
nect it with this form. It is also noteworthy that a Cronartium is found on 
species of Bossiaea. 

(Plate XXVIII., Fig. 246.) 

159. Uredo pallidula Cke. and Mass. 


Cooke and Massee, Grev. XXII, p. 37 (1893). 
Sacc. Syll. XI., p. 222 (1895). 

II. Sori pallid, convex, gregarious, splitting irregularly and then girt by 

the ruptured epidermis, on both surfaces. 

Uredospores pulverulent, tawny in the mass, pale-yellow by 
transmitted light, elliptical to ovoid, finely echinulate, 16-19 X 
12-13 u. 


McAlpine, Agr. Gaz. N.S.W. VI., p. 757 (1895). 
Sacc. Syll. XIV., p. 390 (1899). 

II. Sori hypophyllous, minute, round to oval, sometimes confluent. 

Uredospores variable in size and shape, usually globose or some- \ 

what oval, smooth, orange-yellow, about 19 li diam. or 20 X 16 p.. 

On leaves of Tillaea sieberiana Schult. 

Victoria — Studley Park, near Melbourne (Barnard). 




Excluded or Doubtful Sp 


On leaves, twigs, and legumes of Cassia sp 


In the original description, the spores are given as smooth, and 12-14 

Queensland — Gladfield (Gwyther) (Bailey 

8-10 ;/, but on examining material kindly supplied by Mr. Bailey, of 
Brisbane, they were found to be as above. 

(Plate XXVIII., Fig. 247.) 


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 /j from 
Watts' specimen from Botanical Gardens, Melbourne. I have been able to 
examine part of that same material returned by Cooke and Massee to the 
National Herbarium and find the spores to measure 24-30 x 17-22 /u. The 
rust is fairly abundant along the eastern shores of Port Phillip, and in fresh 

material examined the spores have the same measurements, roughly about 
27 x 20 H . 

Darluca filum Cast, is often found on the sori. 

(Plate XXX., Fig. 267.) 


When one considers that specimens of the various Rusts have hitherto 
been mostly sent to Britain or Germany for determination, it can readily be 
understood that sometimes the material was in an imperfect condition or not 
in the best possible state for proper examination. Hence in some cases the 
species were wrongly determined, and there are several recorded in Cooke 

Handbook of Australian Fungi, which on further investigation must be 

removed from the list of Australian species. Sometimes, although rarely 

happened that the host-plant of the 

ly given, and 

new species has been created which afterwards turned out to be a k 
On these and other grounds the following are excluded 

1. Uromyces amygdali Pass. — Cooke determined this on Peach and 
Almond leaves from Victoria, New South Wales, and Queensland, but when 
complete specimens were examined it was found to be merely the uredo- 
stage of Puccinia pruni Pers. 

2. Uromyces junci (Desm.) Wint. — This species has not been found in 
Australia, as it was wrongly determined at Kew Herbarium for Pziccinia 
juncophila Cooke and Mass. 

3. Puccinia acetosae (Schum.) Koern.— This rust was determined by 
Cooke on a native species of Rumex, but on the examination of a specimen 

the National Herbarium, Melbourne, named in his own handwritin 


2o8 Excluded or Doubtful Species. 

was found to agree with P. ludwigii Tepper. P. acetosae and its synonym, 
P. rumicis Lasch. are therefore excluded. 

4. Puccinia aegra Grove. — P. depauperaiis (Vize) Syd. — The rust on 
the native violets ( Viola hederacea and V. bet onicif olio) is comparatively 
common, and has been collected at various localities in Victoria, New South 
Wales, and Tasmania. It was at first determined by Winter as this species, 
and Cooke has also recorded it, but the examination of numerous specimens, 
including part of the same material forwarded to Dr. Cooke, showed the 
teleutospores to be warted, not smooth, and aecidiospores likewise finely 
warted, and therefore not this species. 

5. Puccinia apii Desm. — The rust found on Celery, both native and 
imported, in Australia, is found to be quite distinct from this species. 

6. Puccinia caulincola Corda is given by Cooke on Hypochoeris glabra, 
but the species on this host-plant is undoubtedly P. hypochoeridis Oud. 

7. Puccinia geranii Corda was the name given by Cooke to the rust 
occurring on Pelargonium australe, but it is now found to be a different 
species, P. morrisoni McAlp. 

s . Puccinia investita Schw. was determined bv Dr. Morrison on 

species of Gnaphalium, but on investigating the original material I found it 
to be Pn gnaphalii (Speg.) P. Henn. 

9. Puccinia microseris McAlp. — Since the host-plant of this species 
was really Hypochoeris radicata it is now excluded. 

10. Puccinia phragmitis (Schum.) Koern. — This species has not yet 
been found in Australia, only P. magnusiana Koern and 1 \ tepperi Ludw. 

11- Puccinia rimosa (Link.) Wint. — This species is given by Winter 2 
as occurring on Isolepis nodosa R. T$r.=Scirpus 7iodosus Rottb. in Victoria, 
but an examination of the original material only shows uredospores which 
do not agree with those of this species, and Sydow * has arrived at the same 
conclusion. The local specimen I have named Uredo scirpi-nodosi. 

12. Puccinia violae (Schum.) DC. — This species is also given by Cooke 
as occurring on violets in Victoria, but the history of the specimens shows 
how necessary it is to guard against recording species as Australian which 
may have been imported in the dry condition. The original material, with 
the name in Cooke's handwriting, is accompanied by the following note 
from the collector, Mr. Reader: — "On leaves of what appears to be Viola 
sp. found in an old book." I was so convinced, however, that P. violae did ; 
not occur on our native violets, and had not been found growing on culti- 
vated species, that I communicated with the collector, who informed me 
that the leaves on which this species was found had been brought from 
Europe and forwarded with other specimens for determination to Dr. 
Cooke. Unfortunately, on the strength of this determination, Mr. Tryon, in 
Queensland, and others, have also credited Australia with this species. Mr. 
Reader, who is a chemist, also sent myself some jaborandi leaves 
(Pilocarpus pennatifolius, Lem.) with a rust upon them, and it was found 
to be Puccinia pilocarpi Cooke, but of course it had been imported. 

13. Cronartium asclepiadeum (Willd.) Fries. — This widely distributed 
species is recorded on a leguminous plant from Queensland, and the name is 
given in Berkeley's handwriting on the specimen in the National Herbarium. 
Dietel, however, regards it as a new species, and has named it C. jacksoniae. 

14. Melampsora nesodaphnes B. and Br. — This species was found on 
the fruit of Beilschmiedia {Nesodaphne) obtusifolia Benth., sent by Bailey to 
Berkeley from Queensland, and I also found it on the fruit of Cinnamomum 
oliveri Bail., sent by Baker from New South Wales. There is a mass of 
powdery spores on the surface of the fruit which is swollen and distorted. 




Excluded or Doubtful Species. 209 

These spores are exceedingly numerous, and very variable in size and shape. 
They are hyaline individually, although ochraceous in the mass and finely 
granular. They vary from elliptic to ovoid or pear-shaped, and are 
occasionally slightly curved. There is a clear central axis to the spore, and 
it is generally broken in the centre, or may be several times transversely 
divided. Their average size is 24-32 x 9-10 ^. They are usually borne on 
short hyaline filaments, and occasionally a long filament is adherent to a 
detached spore. 

There is nothing in the appearance or nature of this fungus to indi- 
cate that it is a Uredine ; it rather suggests some form of Hyphomycete. 

15. Aecidium apocyni Schwein. — The specimen referred by Cooke to 
this species is Caeoma apocyni. 

16. Aecidium barbareae DC— The Cruciferous host-plant is not a 

Barbarea, and a portion of 

specimen shows 

aecidia. I have named the rust Puecinia cruciferae. 

17. Aecidium bellidis Thuem., was determined by Cooke on Bellis 
perennis from Victoria. This is considered to represent the aecidial stage 
of Puecinia obscura Schroet. which occurs on Luzula, but since teleuto- 
spores have been found from year to year on Victorian specimens along 
with the aecidia, it is evidently quite a distinct species. I have, therefore, 
named it Puecinia distinct a. 

Mr. Cheeseman, one of our leading nurserymen, informs me that the 
seeds of Bellis perennis are imported from England and Germany, mostly 
from the latter country. He also states that some have had to give up 
growing the plants on account of the rust (Puecinia distincta), since it 
attacks the youngest seedlings sometimes, and at other times the plants 
remain healthy till late in spring and then become badly attacked. One 
would naturally conclude that the rust had been imported with the seed, but 
it is strange that no teleutospores have been found accompanying the aecidia 
in the countries named. Although I have provisionally recorded it as a 
native rust, the probabilities are that it will be found to exist in the 
countries from which the seed has been derived. 

18. Aecidium COmpOSitarum Mart. — All the species occurring on 
Compositae have been distributed among their respective host-plants. 

19. Aecidium plantaginis Ces. — The species on the native Plantago 
varia is distinct from this, the spores being echinulate instead of 
tuberculate, and considerably larger. It has been named A. plantaginis variae. 

20. Aecidium seuecionis Desm. — This has also been determined in the 
absence of complete specimens, but since the teleutospores have been found 
associated with the aecidia, it has been determined as a new species by 
Dietel, and named P. tasmanica. It is a striking fact that on three such 
common imported plants as the garden daisy, Bellis perennis, groundsel, 
Senecio vulgaris, and marigold, Calendula officinalis, there should hav^e been 
new species of Puecinia discovered. As regards the groundsel, aecidia have 
been found on native species of Senecio, probably belonging to the same rust. 

21. Aecidium Violae Schum., is also recorded, but this belongs to P. 
violae, and is therefore excluded. 

22. Uredo Cichoracearum DC. originally determined by Dr. Cooke 
on Bidens pilosa is now found to be U. bidentis P. Henn. 

23. Uredo leguminum Desm. — This species originally occurred on 
Phaseolus, and is recorded by Cooke as being found on pods of Acacia in 
Queensland, but since there is every reason to believe that the rust on a bean 
is distinct from that on a wattle it is excluded from the list. Probably it is 
a stage of a Uromycladium. 





161. Uromyces polycnemi McAlp. 


I. Aecidia bright orange, generally covering lower surface of leaves, 

crowded, cylindrical, surrounded at base by ruptured epidermis, up 
to \\ mm. long; pseudoperidia colourless, with toothed margin; 
peridial cells oblong to somewhat lozeuge-shaped, with moderately 
thick striated margins, 37-40 /x long. 

Aecidiospores orange, ellipsoid to oblong or sub-globose, very 
finely echinulate, 22-23 X 17-20 ^ or 20-22 /x diam. 

II. Uredo-sori on both surfaces of leaf, bullate, at first covered by 

leaden-coloured epidermis, then splitting, scattered or gregarious, 
sometimes confluent, up to 2 mm. in diam. 

Uredospores ellipsoid to oval, orange coloured, finely echinulate, 
with three equatorial germ-pores on one face, 28-32 X 18-22 \x. 

III. Teleuto-sori similar to uredo-sori, but generally larger and darker in 


Teleutospores dark golden-brown in mass, tawny brown indivi- 

dually, ellipsoid to 

elongated ellipsoid, rounded 

at both ends, 

occasionally slightly thickened at apex and germ-pore prominent, 
smooth, 34-46 X 17-25 /x; pedicel hyaline, persistent, compara- 
tively short. 

On Polycnemum pentandrum F.v.M. 

Victoria — Port Fairy. Dec, 1905. 

The aecidia were very conspicuous, often covering the under sin 

every leaf, and as the host pla 
near the coast, it was difficult 

surface of 
dense masses, in the saline marshes 

A Tuberculina was frequently parasitic on the aecid 

(Plate XLIIL, Fig. 319.) 


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 ^), 
rounded or attenuated at base, not infrequently three-celled, very 
variable in shape and size, 40-62 x 17-28 /jl; pedicel hyaline, 
persistent, up to 45 fx long. 

X. Mesospores common, similarly coloured to teleutospores, thickened 

at apex, oval to ovoid or elongated ellipsoid, 30-40 X 15-23 /x. 

On leaves of Stylidium gramini folium Sm. 

Tasmania — Devonport, Jan., 1906 (Robinson). 

Only the uredospores were at first sent by Rodway in Nov., 1892, and 

the discovery of the final stage shows that Uredo stylidii is a Puccinia, the 
description of which is now completed. 








Abstriction. — The formation of a spore by pinching off the end of a spore-forming 

hypha, without the production of a septum. 

Aecidiosfore. — A spore formed in an aecidium, serially and successively abstricted. 

Aecidium. — A spore-body consisting of a cup-shaped envelope or peridiuni with a 

hymenium at the bottom of the cup, from which aecidiospores are successively 
detached; also the generic name originally applied to the aecidiospores. 

'Amfkigenous. — Growing on both surfaces of a leaf. 

Amfhisfore. — A modified uredospore with thickened wall, and only germinating 

after a resting period. 

Affressorium. — The vesicle formed by the tip of a germ-tube over the stoma of a 

leaf before entering it. 


Autoecious. — Applied to a parasitic fungus of which all the spore-forms are capable of 

being produced on the same host. 

Basidium. — The mother-cell or hypha from which spores are given off. 

Basi fetal. — Growing in the direction of the base from above downwards. 

Bullate. — Raised like a blister. 

Caeoma. — A spore-layer in which the spores are formed in chains but without an 

envelope of peridial cells; an aecidium without a peridium. 

Caeomosfore. — A spore formed in a caeoma. 

Ca fit ate. — Ending in a head, as when the free end of an upright hypha enlarges in 

a spherical manner. 

Chemotaxis. — A form of sensitiveness which certain organisms possess towards cer- 
tain chemical substances. 

Circinate. — Arranged in a circular manner or nearly so. 
Clavate. — Club-shaped, enlarging towards the apex. 

Deciduous. — Falling away, applied to the pedicel of a spore when it does not re- 
main attached to it. 

E chinul at e . — Covered with short spines. 
Elliftical. — Shaped like an ellipse or oval. 

Endochrome. — The peculiar colouring matter in the cells, and usually applied to the 

coloured cell-contents of spores. 

Endosfore. — The innermost coat of a spore. 

Efifhyllous. — Growing on the upper surface of a leaf. 

Efiteosfore. — A name suggested for a uredospore which functions as an aecidiospore, 

and the sorus contains or is surrounded by prominent paraphyses. 

Erumfent. — Bursting through the surface of the host-plant. 

Fimbriate. — Fringed, when the margin is bordered by slender processes. 

Form-genus. — A genus in which the species are only represented by single stages of 

what is believed to be an incomplete life-history, as Uredo, Aecidium. 

Gall. — A morbid enlargement or hypertrophy of the cells of a plant, due to para- 
sitic agency. 
Gametofhyte. — The stage in the life-cycle of the plant which bears the sexual organs. 

Germ-fore.— An opening on the surface of a spore through which the germ-tube 

Germ-tube.— A tubular process developed from the spore in germination, and may 

either become a mycelium or promycelium. 

Gibbous. — Swollen at the base. 

Globose. — Spherical. 

Haustorium.—A short lateral branch of a hypha, penetrating a cell of the host-plant 

and acting as a sucker as well as an organ of attachment. 
Htteroccious.— Applied to a parasitic fungus of which certain spore-forms occur on 

one host and others on plants widely remote. 
Host. — A plant which nourishes a parasite. 
Hyaline. — Colourless or translucent. 



Hymenium. — A spore-bearing layer of hyphae. 

Hypertrophy. — An abnormal enlargement or excessive growth of particular parts,. 

which may be caused by excess of nourishment or induced by parasites. 

Hypha. — A tubular thread-like cellular structure, collectively constituting the vege- 
tative body or mycelium of the fungus. 

—Growing on the under surface of a leaf. 


Indigenous. — Native to a country, and not introduced. 

Intercellular. — Between the cells of the host-plant. 

Intracellular. — Within the cells of the host-plant. 

Laciniate. — Cut up into narrow lobes. 

Lumen. — The cavity bounded by the walls of an organ. 

Mesospore. — A single-celled spcre-form related to the teleutospore, and probably 

representing a transition from the unicellular Uromyces. 

Micron or "Micro millimetre. — Indicated by the 


ji y is the standard unit for 

microscopical measurements, and equals i-ioooth of a millimetre, or i-25,oooth 
of an inch. 

Mutation. — That kind of 

equilibrium of the organism is disturbed and 

variation produced by internal influences, in which the 

a new position of equilibrium 

found strikingly different from the original one, sometim-es called spontaneous 
variation. , 

Mycelium. — The vegetative portion of a fungus composed of one or more hyphae. 

Nucleus. — An organized proteid constituent of the cell, necessary for its continued 

growth and life. 

Obligate parasite. — One in w T hich a parasitic mode of life is indispensable for com- 

plete development. 

Orbicular. — Applied to a flat body with a circular outline. i 

Papillate. — Covered with protuberances or terminating in a papilla. 

Paraphyses. — Barren filajnents associated with the spoies, either slender or variously 

swollen, hyaline or coloured. 

Parasite. — An organism living on or in another living organism, and at its expense. 

Pedicel. — Spore-bearing stalk. 

Persistent. — Lasting, not soon falling away, applied to the pedicel of a spore. 
Piriform. — Pear-shaped. 

Plasmodium. — A mass of naked protoplasm, containing many nuclei and exhibiting 


Plurivorous. — Applied to a parasite which can exist on a variety of host-plants. 

Predisposition. — The tendency to disease exhibited by an organism when the con- 
ditions are favorable to the development of the parasite. 

Primordia. — The beginnings of any structure, such as the aeciiium. 

Promycelial spore. — The spore produced by a promycelium, also known as a sporid- 


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. 


■The living substance in plants, consisting of albuminoid matter. 

False tissue or compact mycelium, so called because it is 
formed by the union of previously separate elements or hyphae into a dense 



mass, while true tissue is 

forms a coherent, more or less firm, mass of cells. 

derived from a single cell, which divides 


— The outer tnvelopc of the aecidium, consisting of sterile cells. 

Pulverulent. — Powdery 
Pulvinate. — Cushion-shaped. 

Covered with lines or ridges crossing each other so as to form a net- 


Saccate. — Bag-shaped? 

Saprophyte. — A fungus which preys upon dead organic matter only, in contrast to 

a parasite. 

Septate. — Partitioned off into distinct divisions. 

Sorus. — A cluster of spores constituting a spore-bed, as uredosorus, teieutosorus. 



Sfermatium. — A minute detachable cell produced in a spermogonium, and formerly 

considered to have a male sexual function. 

Sfermogonium. — The receptacle in which spermatia are produced and detached. It 

is sometimes called a pyenidium, although that term is generally reserved for 
a spore-receptacle in another division of Fungi. 

Spore. — A detachable cell, capable of reproducing the fungus. 

Sporidiolum. — Spore produced by the germ-tube of a teleutospore. Also called 

sporidium, but since this term is already used as equivalent to an ascospore, 
Saccardo has proposed the present name. 

Sporophyte. — The stage in the life-cycle of a plant which bears the spores. 

Sterigma. — A delicate stalk from which a spore is detached and arising from a 


Stoma. — A breathing pore or aperture in the epidermis forming a means of communi- 
cation between the inside of the plant and the outer air. 

Stroma. — A cushion or receptacle bearing reproductive bodies. 
Symbiosis. — The living together of dissimilar organisms, both contributing to their 

mutual welfare. It differs from parasitism' in which the benefit is all on the 

side of the parasitic organism. 
Teleutos<pore.—Genera\\y regarded as the final spore in the life cycle of the Rusts, 

which germinates and produces a promycelium and promycelial spores. 

Truncate. — Cut off abruptly. 

Uredo. — The generic name originally applied to the uredospores. 
Vredosfore.—A spore detached from the apex of a pedicel and producing a 

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. 

Problems in the Study of Plant Rusts. Bull. Torr. Club, Vol. 30. P- 





Taxonomic Importance of the Spermogonium. Ibid. Vol. 31, p. H3- 

-. The Aecidium of Maize Rust. Bot. Gaz., Vol. 38, p. 64. 1904. 

J. Amphisporesi of the Grass and Sedge Rusts. Bull. Terr. Club, \ol. 32, 

6 . P " 3 A N 9 ew' Uredineous Fungus from Mexico— Baeodromous holwayi Arth. 


Ann. Myc, Vol. 3, p. 18. 1905. 

- - • J 


7. Cultures ot ureameac m iyu 4 . j«-u. MJ ~. .—. --, r ^ ' ^ v , 

8. Terminology of the Spore-Structures in the Uredinales. Bot. Gaz., vol. 

39, P- 2I 9- i9°5 

1. Arthur, J. C, and Holway, E. W. D.-Descriptions of American Ured.neae. 

Bull. Lab. Nat. Hist., Univ. Iowa, Vol. 5, P- 3*9- If ><> 2 - 
1. Atkinson, J.-An account of Agriculture and Grazing in New South ttales. 

London. 1826. 


Brisbane. 1883. 

1. Bailey, F. M.— Synopsis of the Queensland Mora pp. 774"Z7»- ""g 

2. Contributions to the Queensland Flora. Proc. Roy. Soc, Q 

Vol. 1, Part I., p. 18. 1884. 

* =1 l^psU of *e, Firs, Supplement, pp. 84, 85. Bri, 

bane. 1886. -n • u .~ T coo 

5. Ibid, Second Supplement, pp. 125-127-. . Bnsban |- lb88 ' 

6- Ibid, Third Supplement, p. 123. Brisbane. 1890. 

7- Additional Fungus Blights observed to have ^ e .l^ 5w * 

' - • - c . of Queensland, p. 45- Brisbane. 1890-01. 

leensland Flora. Botany Bull. 2, p. 34. Brisbane. 

. of Queensland, p. 51. Brisbane. 1891-2. 








10. Bailey, F. M. — A Review of the Fungus Blights which have been observed to in- 
jure living vegetation in Queensland. Rep. Aust. Assoc. Adv. Sci., Hobart, 
Vol. 4, pp. 392, 395, 396, 397, 398, and 400. 















Contributions to the Queensla] 
Ann. Rep. Dep. Agric. of Q 
Contributions to the Queensh 


Botany Bull. 5, p 

p. 40. Brisbane. 

36.. 1892. 

Flora. Botany Bull. 8, pp. 110-113 

Brisbane. 1893. 

Ann. Rep. Dep. 
Contributions to 

Brisbane. 1894. 

Agric. of Queensland, p 
the Queensland Flora. 

Brisbane. 1893-4. 

Ann Rep. Dep. 



of Queensland, p. 

Fungi. Proc. Conf. 

3 1 - 

Botany Bull. 9, pp. 16, 17 

35. Brisbane. 1894-5. 
Austr. Fruit-growers, New 

Zealand, p. 208. Well 


Ann. Rep. Dep. 


of Q 

p. 39. Brisbane. 1897-8. 


Botany Contributions. Queensland Agric. J 

38. Brisbane. 


Ann. Rep. Dep. Agric/ of Q 


Ibid. Vol. 3, p. 205 


Brisbane. 1899. 


p. 33. Brisbane, 1898-9. 
. Agric. Journ. Vol. 4, 


1. Baker, R. T. 

Contributions to a Knowledge of the Flora of Australia. 


Linn. Soc, New South Wales, Vol. 24, p. 447. 1899. 

1. Bandi, W. — Beitrage zur 


der Uredineen (Phragmidium subcorticium 

{Schrank) Winter). Hedw., Vol. 42, p. 118. 1903 

1. Barclay, A. — Aecidiwn 



Schum. var. himalayense. Sci. M 


Off. Army of India, p. 29. Calcutta. 1887. 



On the Life History of a Remarkable Uredine on J 


(Uromyces cunninghamianus, n. sp.). Trans. Linn. Soc, London, 

3, p. 141. 1891. 
On Two Autoecious 

Caeomata in Simla. Scientific M 

by M 

Officers of Army of India, Part VI. 1891. 

Rust and Mildew in India. J 


1. Bary, A. de. — Neue 

Bot, Vol. 30, p. 349 



ueber die Uredineen, insbesondere die 
Entwickelung der Puccinia graminis und den Zusammenhang derselben mit 
Aecidium berberidis. Monatsber. K. Acad. d. Wiss, p. 25. 

Comparative Morphology and Biology of the Fungi, Mycetozoa 

teria. English Edition, Oxford. 1887. 


is d'immunisation des vegetaux contre les maladies crypto- 

e, M. J.- 

gamiques. Compt. rend. Vol. 133, p. 107 

1 901 

1. Berkeley, M. J. — Flora of Tasmania 


Hooker's Botany of the Ant- 

arctic Voyage, Part III., Vol. 2, p. 270. i860. 

& 11 ^^^ a 4 — * 4 ^^^B 





Linn. Soc, London, Vol. 13, pp. 173* ^A* 

1. Berkeley, M. J., and Broome, C. E. — List of 


land. Trans. Linn. Soc, London, Vol. 1, p. 407 

Ibid, Part II., Vol. 2, p. 67. 1882. 

from Brisbane, Q 

Ibid, Part III., Vol. 2, p. 221. 1886. 

3- . 

1. Blackman, V. H. — On the conditions of Teleutospore germination and Spori- 

dia formation in the Uredineae. New Phytologist. Vol. 2, p. 10. 1903. 

On the Fertilisation, Alternation of Generations, and general Cytology 

2. - 


of the Uredineae. Ibid. Vol. 3, p. 23. 1904. 

On the Fertilisation, Alternation of Generations, and 

— -- j 

of the Uredineae. Ann. Bot., Vol. 18, p. 323. 1904 

general Cytology 

1. Bolley, H. L. — The Heteroecismal Puccinieae. Am. Micro. J 

10, p 

169. 1889. 



Bull. 26. Agr. Exp. Station, Indiana. 1889. 


Wheat Rust : Is the infection local or general in origin? 

Vol. 5. 1891. 

Einige Bemerkungen ueber die symbiotische Mykoplasm 

Getrefderost. Centralb. f. Bakt., Vol. 4, p. 855. 1898. 
New work upon wheat rust. Science N.S. Vol. 


bei dem 



22, p. 50 




fi DC. Oesterr. Bot. Zeitschr., Vol. 48, p. 14. 1898. 


Ber. d. Deutsch Bot. Gesellsch., Vol. 21, p. 356. 
Butler, E. J.— The Indian Wheat Rust Problem. 
India. Calcutta. 1903. 


No. 1, Dept. Agric, 





i. Campbell, F. M. — Victorian Fungi. Vict. Nat. Vol. 4, p. 96. 1887. 

1. Carleton, M. A. — Cereal Rusts of the United States. U.S. Department of 

Agriculture, Division of Vegetable Physiology and Pathology, Bull. No. 16. 
Washington. 1899. 

2. Culture Methods with Uredineae. Journ. Applied Microscopy and Lab. 

Methods. Vol. 6, p. 2109. Rochester, N.Y. 1903. 

3. Investigations of Rusts. U.S. Department of Agriculture, Bureau of 

Plant Industry. Bull. 63. Washington. 1904. 

1. Chodat, R. — Identite du Puccinia scirfi DC. avec Aecidium nymfhoidis DC. 

Verh. Schweiz. Nat. Ges., p. 43. 1888-9. 

1 Christman, A. H. — Sexual Reproduction in the Rusts. Bot. Gaz., Vol. 39, p. 

267. 1905. 

1. Cobb 5 N. A. — Peach Rust in our Orchards. Agric. Gaz., N.S. Wales. Vol. i % 

p. 93. 1890. 

2. Contributions to an economic knowledge of Australian Rusts (Uredineae). 

Ibid., pp. 185-214. 1890. 

3. Maize Rust. Ibid. Vol. 2, p. 215. 1891. 

4. Notes on Diseases of Plants. Ibid, pp. 157, 286. 1891. 

5. Report on Rust in Wheat. Proc. Conf. N.S. Wales. 1891. 

6. Contributions to an economic knowledge of Australian Rusts (Uredineae). 

Agric. Gaz., N.S.W., Vol. 3, pp. 44-68. 18Q2. 

7. Fungus Diseases of the Sugar-cane. Ibid. Vol. 4, p. 799. 1893. 

% Host and Habitat Index of the Australian Fungi. Dep. Agric, N.S. 

Wales. 1893. 
9. Notes on Diseases of Plants. Agric. Gaz., N.S. Wales Vol. 5, p. 384. 


10. Letters on the Diseases of Plants. Ibid. Vol. 8, pp. 216, 217, 232-234. 

l8 97- . . . 

11. Comparative Observations on the Brush of about fifty Varieties of 

Wheat. Ibid. Vol. 13, p. 647. 1902. 

12. Letters on the Diseases of Plants. Ibid. Vol. 15, pp. 8-14. 1904 

1. Cobb, N. A., and Oliff, A. S. — Insect Larvae {Cecidomyia sp.) eating Rust 

on Wheat and Flax. Agric. Gaz., N.S.W. Vol. 2, p. 67. 1891. 

1. Connold, E. T. — British Vegetable Galls. London. 1901. 

1. Cooke, M. C— Australian Fungi. Grevillea, Vol. 11, pp. 97, 98. 1883. 

2. Ibid, Vol. 12, p. 20. 1883. 

3. Fungi australiani (reprinted from Grevillea, Vols. 9, 10, 11, 12). 1883. 

4. Australian Fungi. Grevillea, Vol. 14, pp. 11, 12. 1885. 

5. Ibid. Vol. 15, p. 99. 1887. 

6. Ibid. Vol. 16, p. 2. 1887. 

7. Ibid. ' Vol. 16, p. 74. 1888. 

8. Ibid. Vol. 17, pp. 55, 70. 1889. 

9. Ibid. Vol. 19, p. 46. 1890. 

xo. Handbook of Australian Fungi, pp. 330-344. London. 1892. 

1. Corda, A. — Icones Fungorum. 6 Fasc. 1838-56. 

1. Dangeard, P. A., and Sappin Trouffy, P. — Recherches histologiques sur les 

Uredinees. Compt. rend. Vol. 116, p 211. 1893. 

2. Une pseudofecondation chez les Uredinees. Ibid. Vol. 116, p. 267. 



1. Dietel, P. — Beitrage zur Morphologie und Biologie der Uredineen. Bot. 

Centralb. Vol. 32, p. 54, &c. 1887. 

2. Verzeichniss der in der Umgebung von Leipzig beobachten Uredineen. 

Ber. Natf. Ges. Leipzig. 1888-89. 

, 3. Ueber das Vorkommen von zweierlei Teleutosporen bei der Gattung 

Gymnosforangium. Hedwigia, Vol. 28, p. 99. 1889. 

4. Ueber den Generationswechsel von Puccinia agrofyri Ell. and Ev. 

Oesterr. Bot. Zeitschr. Vol. 42, p. 261, 1892. 

5. Die Gattung Ravenelia. Hedw., Vol. 33, pp. 22 and 367. 1894. 

6. Ueber Rostpilze mit wiederholter Aecidienbildung. Flora. Vol. 8i, p. 

394. 1895. 

7. Waren die Rostpilze in friiheren Zeiten plurivor? Bot. Centralb. Vol. 

79, p. 81 and 113. 1899. 





8. Dietel, P. — Uredinaies in Engler and Prantl, Die natiirlichen Pflanzen- 

familien. 1900 





Ueber die biologische Bedeutung der Paraphysen in den Uredolagern 

von Rostpilzen. Hedw., Vol. 41, p. (58). 1902. 

Ueber die auf Leguminosen lebenden Rostpilze und die Verwandt- 

schafts verhaltnisse der Gattungen der Pucciniaceen. Ann, Myc. Vol. 1 
p. 3. 1903. 

Eine neue Puccinia auf Senecio. Ibid., p. 535. 1903. 

Betrachlungen ueber die Vertheilung der Uredineen auf ihrer Nahr- 

pflanzen. Centraib. f. Bakt., Vol. 12, p. 218. 1904 



Ueber die Arten der Gattung Phragmidium. Hedwigia, Vol. 44, p. 122 

P- 33o 


1. Dumee, P., and Maire, R. — Remarques sur les Uredospores de Puccinia fruni 

Pers. Bull. Soc. Myc, France. Vol. 17, p. 308. 1901 


1. Ehrlich, and Huebener. 

Zeitschr. f. Hygiene. 

' - • 

Ueber die Vererbung der Immunitat bei Tetanus. 

Vol. 18. 1894. 


J. — Ueber die Spezialisierung des Parasitismus bei den Getreiderost- 














J 5- 





pilzen. Ber. Deutsch. Bot. Ges. Vol. 12, p. 292 


Ueber die Forderung der Pilzsporenkeimung durch Kalte. Centraib. f. 

Bakt. Vol. 1, p. 557. 1895. 

Ueber die verschiedene Rostempfanglichkeit verschiedener Getreidesorten. 
Zeitschr. f. Pflanzenk. Vol. 5, pp. 80 and 156. 1895. 
1st die verschiedene Widerstands fahigkeit der Weizensorten konstant 

oder nicht? Ibid., p. 198. 

Welche Rostarten zerstoren 




Ibid. Vol. 

6, p. 141. 

Welche Grasarten konnen die Berberitze mit Rost anstecken? Ibid., p. 



Neue Untersuchungen ueber die Spezialisierung, Verbreitung und 

kunft des Schwarzrostes (Puccinia graminis Pers.). Jahrb. f. Wiss. Bot. 



p. 499 



Vol. 15, p. 183. 1897. 

Stand der Getreiderostfrage. Ber. Deutsch. Bot. Ges. 


124. P- 475 . ~ .. 

Ueber die Dauer der Keimkraft in der Wintersporen gewisser Rostpilze. 

Centraib. f. Bakt. Vol. 4, p. 376. 1898. 

Vie latente et plasmatique de certaines Uredinees. Compt. rend. 


Sur l'Origine et la Propagation de la Rouille des Cereales par la Semence. 

Ann. Sci. Nat. Bot., Ser. 8. Vol. 14, p. 107. 1901. 

Ueber die Spezialisierung des Getreidescliwarzrostes in Schweden und 

in anderen Landern. Centraib. f. Bact. Vol. 9, p. 590. 1902. 

The Researches of Professor H. Marshall Ward on the Brown Rust of 

for Botanik udg. of K. 

the Bromes and the Mycoplasm Hypothesis. Arkiv 
Sv. Vet. Akad. Vol. 1, p. 139. 1903. 

Ueber das vegetative Leben der Getreiderostpilze. 

Vol. 77, No. 6, I. 1004. 

Ibid. Vol. 38, No, 3, II. -III. 

K. Sv. Vet. Akad. 



On the Vegetative Life of some Uredineae. Ann. Bot. Vol. 19, p. 55 

Zur Frage der Entstehung und 

Pflanzen. Arkiv. f. Botanik. K. Sv. Vet. Akad. Vol. 5 

Ueber das vegetative Leben der Getreiderostpilze. 

Vol. 39, No. 5, IV. 1905 
Eriksson, J., and Henxing, E. 

ueber die Getreideroste. 

Verbreitune der Rostkrankheiten der 

No. 3. 1905. 
K. Sv. Vet. Akad. 



Die Hauptresultate einer neuen Untersuchung 
Zeitschr. f. Pflanzenk/ Vol. 4, pp. '66 and 140. 

Getreideroste, ihre Geschichte und Natur, sowie Massregeln gegen 

dieselben. Stockholm. 1896. 



1. Farlow, W. G. — Notes on some Species in the Third and Eleventh Centuries of 

Ellis' North American Fungi. 
18. Boston. 1883. 

Proc. Am. Acad. Arts 

d Sci. 


The Conception of Species as affected bv recent Investigations on Fungi 

American Naturalist. 

Vol. 32, p. 675. 





1. Farrer, W.— The making and improvement of Wheats for Australian condi- 

tions, together with Eriksson's General review of the principal results of s» 
Swedish research into the rust of cereals. Agric. Gaz., N.S. Wales, Vol. q* 
p. 239. 1898. 

- Some notes on the Wheat " Bobs"; its Peculiarities, Economic Value, and 


Origin. Agric. Gaz., N.S. Wales, Vol. 15, p. 739 and p. 849. 1904 

1. Fischer, E. — Ueber Gymnosforangium sabinae (Dicks.) Wint. and G. confusum 

Plowright Zeitschr. f. Pflanzenk. Vol. 1, p. 260. x8qi. 

2. Experiences d'infection avec les Puccinia helvetica Schroet. et P. mag- 


Koern. Arch. Sci. Phys. and Nat. Vol. 28, p. 376. 1892. 

3. Die Zugehoerigkeit von Aecidium fenicillatum. Hedw. Vol. 34 p 1. 

. l895 ' 
4- Die biologischen Arten der parasitischen Pilze und die Entstehung neuer 

Formen im Pflanzenreich. Verh. Schweiz. Nat. Ges. Vol. 86. Locarno, p. 49, 

5. Die Uredineen der Schweiz. Beitr. zur Krypt. Flor der Schweiz. Vol. 2 y 

part 2, 1904. 

1. Freeman, E. M. — Experiments on the Brown Rust of Bromes {Puccinia dis- 

fersa). Ann. Bot., Vol. 16, p. 487. 1902. 


1. Galloway, B. T. — Rust of Flax. Journ. Myc, Vol. 5, p. 215. 1889. 

1. Gibson, C. M. — Notes on Infection Experiments with various Uredineae. New 

Phytologist. Vol. 3, p. 184. 1904. 



1. Halsted, B. D. — Mycological Notes — Observation in Wind infection of a Rust- 
Bull. Torr. Bot. Club. Vol. 25, p. 159. 1898. 

1. Henning, E. — Ueber verschiedenartige Predisposition des G°.treides fur Rost. 

Land. Akad. Handl. Tids. 1894. 

1. Hennings, P. — Fungi Warburgiani. Hedw., Vol. 32, p. 216. 1893. 

2. Anpassungsverhaltnisse bei Uredineen bezuglich der physikalischei> 

Beschaffenheit des Substrats. Hedwigia, Vol. 40, p. 125. 1901. 

3. Fungi Australiae occidentalis I. Ibid., pp. 95 and 352. 1901. 

4. Fungi australienses. Ibid., Vol. 42 (p. 73). 1903. 

5. Einige Beobachtungen ueber das Gesunden pilzkranker Pflanzen bei veran- 

derten Kulturverhaltnissen. Zeitschr. f. Pflanzenk. Vol. 13, p. 41. 1903. 

tchcock, A. S., and Carleton, M. A. — Preliminary Report on Rusts of Grain, 
Kans. Agr. Expt. Sta. Bull. No. 38. 1893. 


2. Rusts of Grain, II. Ibid. Bull. No. 46. 1894. 

1. Hooker, J. D., and Jackson, B. D. — Index Kewensis. Oxford. 1895, & c - 


1. Jacky, E. — Die Compositen-bewohnenden Puccinien vom Typus der Puccinia 

hieracii und deren Spezialisierung. Zeitschr. f. Pflanzenk. Vol. 9, pp. 193, 
263, and 330. 18Q9. 

2. Der Chrysanthemum-Rost. Ibid.. Vol. 10, p. 132. 1900. 

3. Der Chrysanthemum-Rost. Centralbl. f. Bakt., Vol. 10, p. 369. 1903. 


1. Kienitz-Gerloff, F. — Die Gonidien von Gymnosforangium clavariaeforme. Bot. 

Zeit. Vol. 46, p. 389. 1888. " , 

1. Klebahn, H. — Die Wirtwechselnden Roslpilze. Berlin. 1904. (This masterly 

work includes references to Klebahn's numerous contributions to the litera- 

ture of the Rusts.) 

2. Kulturversuche mit heteroecischen Uredineen. Zeitschr. f. Pflanzenkr. 

Vol. 2, p. 337. 1892. ^ 

3. Kulturversuche mit Rostpilzen xii. Bericht (1903-1904). Zeitschr. f. Pflan- 
zenkr. Vol. 15, p. 106. 1905. 

1. Kuehn, J.— Der Rost der Runkelriibenblatter, Uromyces betae. Bot. Zeit, p. 

540. 1869. 

H 2 





I. Lagerheim, G. v. — Ueber Uredineen mit variablem Pleomorphismus. Tromso 

Mus. Vol. 16, p. 105. 1894. 

1. Laurent, E. — De Taction interne du sulfate de cuivre dans la resistance de la 

pomme de terre au Phytofhthora infestans. Compt. rend., Vol. 135, p. 




Lindroth, J. I. — Cecidomyia-larver, som ata rostsporer. Medd. Soc. Fauna. 
Fl. Fenn. Vol. 26, p. 25. 1900. 

1. Ludwig, F. — Ueber einen neuen Goodeniaceenrost aus Siidaustralien. Hedw. 

Vol. 28, p. 362. 



Ueber einige neue Pilze aus Australien. Bot. Centralb. Vol. 


43i P- 5 


Ueber neue australische Rostkiankheiten. Zeitschr. f. Pflanzenkrank- 

heiten, Vol. 2, p. 130. 




Ueber einige Rost-und Brandpilze Australiens. Ibid., Vol. 

3. p. 137- 


Lehrbuch der niederen Kryptogamen, p. 455. Stuttgart. 1892. 


1. Magnus, P. — Ueberwinterung der Puccinia caricis. Verh. Bot. Ver. Prov. 


Brand, Vol. 18, p. 27. 1885. 

Ueber das Auftreten der Stylosporen bei den Uredineen. 

Deutsch. Bot. Gesellsch., Vol. 9, p. 85. 1891. 

Ber. der 



Zur Kentniss des Verbreitung einiger Pilze. Ber. Deutsch. Bot. Ges., 

10, p. 195 



Die systematische Unterscheidung nachst verwandter parasitischer Pilze 

auf Grund ihres verschiedenen biologischen Verhaltens. Hedwigia, Vol. 

33> P- 362. 



Fungi Pars. II. in J. Bornmueller, Iter Persico-turcicum. 1892-93. 

Verhandl. Zool.-bot. Gesellsch. Wien, Vol. 49, p. 95. 


Ueber eine Function der Paraphysen von Uredolagern. Ber. Deutsch. 


Bot. Ges., Vol. 20, p. 334 


Kurze Bemerkung zur Biologie des Chrysanthemum-rostes. Centralb. f. 

Bakt. Vol. 10, p. 575. 1903 

1. Maiden, J. H. — Report on Botanic Gardens and Domains, p. 10. N.S. Wales. 


1. Marchal, E. — De la specialisation du parasitisme ohez V Erysifhe graminis. '• 


Compt. Rend., Vol. 135, p. 210. 1902. 

De 1' immunisation de la Laitue (Lettuce) contre le Meunier. Ibid p 



1. Massee, G. — On the Presence of Sexual Organs in Aecidium. Ann. Bot., Vol. 

2, p. 47. 


- Notes on Exotic Fungi in the Royal Herbarium, Kew. Grev., Vol. 21 

p. 4 



Australian Fungi. Grev., Vol. 22, p. 17 



Mycological Flora of the Royal Gardens, Kew. Kew Bull., p. 151. 




Chrysanthemum Rust. Gardeners' Chronicle, 8th Oct., 1898. 

The Cereal Rust Problem. Does Eriksson's Mycoplasma exist in Nature? 

Natural Science, Vol. 15, p. 337. 1899. 

To protect Cucumbers and Tomatoes from fungus. Journ. Roy. Hort. 

Soc, Vol. 28, p. 142. 1903. 

A Text-book of Plant Diseases, 2nd. Ed. London. 1903.. 

On the Origin of Parasitism in Fungi. Phil. Trans. Royal Soc, Ser. B, 

Vol. 197, p. 7. 


cTer Uredineen in ihrer Abhangigkeit von Stand- 
ee o 

1. Mayus, O. — Die Peridienzellen 

ortsverhaltnissen. Centralb. f. Bakt., Vol. 10. p. 644. 1903 
1. McAlpine, D.— The Life-history of the Rust of Wheat. Bull. 14, Dep. Agric, 


Victoria, pp. 22-32, pis. 2. 1891. 

Rust of Wheat. Ibid. pp. 33, 34 



45, 46. 

Beet-leaf Rust or Blighted Mangel leaves 


[Uromyces betae). Ibid, pp 


Report cm Peach and Plum Leaf-Rust {Puccini fruni). Ibid, pp. 138, 

144, pis. 3. 




Report on Rust in Wheat as Victorian Delegate. Proc. Conf., N.S. 

1 891. 



6. McAlpine, D. — Ueber die Verwendung geschrumpfter Koerner von rostigem 

Weizen als Saatgut. Zeitschr. f. Pflanzenkrankh., Vol. 2, p. 193. 1892. 

7. Report on Rust in Wheat as Victorian Delegate. Proc. Conf. S. Aus- 
tralia. i8q2. 

8. The Undescribed Uredospores of Puccinia bur char diac. Vict. Nat., 

Vol. 10, p. 192. 1894. 

9. Report on Rust in Wheat Experiments at School of Horticul- 
ture, Burnley. Illustrated with maps and drawings, pp. 66. Dep. Agric, 
Victoria, 1894. 

10. Report on Rust in Wheat as Victorian Delegate. Proc. Conf., Queens- 
land. 1894. 

11. Australian Fungi. Proc. Roy. Soc, Victoria. Vol. 7, p. 214. 1895. 

12. Systematic Arrangement of Australian Fungi, together with Host Index 

and List of Works on the subject. Dep. Agric, Victoria. 1895. 

13. Notes on Uromyces amygdali — a Synonym of Puccinia fruni Pers. 

Proc. Linn. Soc, N.S. Wales. 2nd series, Vol. 10, p. 440. 1895. 

14. Puccinia on Groundsel with Trimorphic Teleutospores. Ibid., p. 461. 


-15. Australian Fungi. Agric. Gaz., N.S. Wales, Vol. 7, p. 752. 1895. 

16. Ibid, p. 850. 1895. 

17. Ibid. Vol. 7, p. 147. 1896. 

18. Ibid, p. 299. 1896 

19. Two Additions to the Fungi of New South Wales. Proc. Linn. Soc, 

N.S. Wales. Vol. 21, p. 722. 1896. 

20. New South Wales Fungi. Ibid. Vol. 22, p. 36. 1897. 

21. Rust in Wheat Experiments, 1894-1897. Dep. Agric, Victoria, 1S97. 

22. The Fungi on the Wheat Plant in Australia. Agric Gaz., N.S. Wales, 

Vol. q, p. iooq. 1898. 

23. Rust in Wheat during the dry season of 1897. Ibid. p. 1421. 1898. 

24. On a micro-fungus from Mount Kosciusko, and on the first record of 

Uncinula in Australia. Proc Linn. Soc, N.S. Wales. Vol. 24, p. 301. 1899. 

25. Two Mallee Fungi. Vict. Nat., Vol. 16, p. 141. 1899. 

26. Rose Rust. Journ. Dept. Agric, Vic, p. 81. 1902. 

.27. Fungus Diseases of Stone-fruit Trees in Australia, and their Treatment. 

Dept. of Agric, Victoria. 1902. 

28. Cereal Rusts. Joura. Dept. Agric, Vic, p. 425 1902. 

29. Peach and Plum Rust. Ibid. p. 617. 1902. 

30. Australian Fungi, new or unrecorded, Decades V.-VI. Proc. Linn. 

Soc, N.S. Wales. Vol. 28, p. 558. 1903. 

31. Rerraf — A Rust-resisting Wheat. Journ. Dept. Agric, Vic, Vol. 2, 

p. 531. 1904. 

32. Diseases of Cereals, Rust and Take-all in Wheat. Ibid. p. 700. 1904. 

33. Some Misconceptions concerning the Uredospores of Puccinia fruni 

Pers. Ann. Myc, Vol. 2, p. 344. 1904 

34. Note on the Arrangement of Teleutospores in Puccinia fruni Pers. 

Ibid. p. 348. 1904. 

35. Bobs — A Rust-resisting Wheat. Journ. Dept. Agric. Vic, p. 166. 1905. 

36. A new genus of Uredineae — Urotnycladium. Ann. Mvc Vol. 3, p. 303. 


1. Milesi, M., and Tra verso, G. B. — Saggio di una monografia del genere Tri- 

fhragmium. Ann. Myc, Vol. 2, p. 143. 1904. 

1. Molixeux, A. — Some Remarks about Red Rust. Journ. Agric, S. Australia. 
. Vol. 8, p. 95. 1904. 

1. Morrison, A. — New Victorian Micro-fungi. Victorian Nat., Vol. 11, pp. 90, 

91. 1894. 

2. Notices of Victorian Fungi. Ibid. pp. 119, 120. 1894. 

1. Mueller, F. v. — Fragmenta Phytographiae Australiae — Supplement, Vol. 11, p. 

96. 1880. (Fungi enumerated by Dr. Cooke.) 

2. Census of the genera of plants hitherto known as indigenous to Australia. 

Proc. Roy. Soc N.S. Wales, Vol. 15, pp. 254, 255. 1881. 

3. Notes on Victorian Fungi. Victorian Nat., Vol. 2, p. 80. 1885. 

4. Lecture on Rust in Cereals. Sandhurst, Victoria. 1865. 

1. Mueller, Fritz.— Beitrage zur Kentniss <ler Grasroste. Eot. Centralb. Beihefte 

10, p. 181. 1901. 

x. Mueller, J. — Die Rostpilze der Rost-und Rubusarten und die auf ihnen vor- 

kommenden Parasiten. Landw. Jahrb., Vol. 15, p. 719- 1886. 





i. Neger, F. W. — Beitrage zur Biologie der Erysipheen. Flora 90, p. 221 


1. Neumann, R. — Ueber die 

Entwickelungsgeschichte der 

gonien der Uredineen. Hedwigia, Vol. 33, p. 346. 

Aecidien und Spermo- 







i. Pfeffer, W. — Ueber chemotaktische 


von Bakterien, Flagellaten 

und Volvocineen. Untersuch. Bot. Inst., Tubingen. Vol. 2, p. 582. 1888. 

1. Plowright, C. B. 


with the Barberry Aecidium (Aecidium berberidis Gmel.) Gard. Chron. 
Ser. 2, Vol. 18, p, 231. 1882. 

Podisoma junif 

Reproduction of Heteroecious Uredines. Linn. 

Ibid, p. 553. 

Soc, London. 


Vol. 21, p. 368. 



On the Life-history of certain British Heteroecismal Uredines. Quart 

Micro. Sci., Vol. 25, p. 156. 


A Monograph of the British Uredineae and Ustilagineae. London. 1889. 

Barberry and Wheat Mildew. Gard. Chron., Vol. 23, p. 45 


Prain, D. — Rust in Wheat in the Australian Colonies. 

Agric. Ledger No. 

i6 r 



1. Prillieux, E. 

Maladies des Plantes Agricoles. 

Pt. 1, p. 242. Paris, 1895. 








1. Ralph, T. S. — On the Aecidium affecting the Senecio vulgaris or Groundsel. 

Victorian Nat., Vol. 7, pp. 18, 19 


1. Rathay, E. — Untersuchungen ueber die Spermogonien der Rostpilze. Denkschrift. 

Wiss Wien 


1. Richards, H. M. — On some points in the development of Aecidia. Proc. 

Amer. Acad. Sci., Vol. 31, p. 255. 



1. Rodway, L. — Tasmanian 



Proc. Roy. Soc, Tasmania, pp. 142, 143 

1. Roze, M. E. — Le Puccinia chrysanthemi. Bull. Soc. Myc. France, Vol. 16 









1. Saccardo, P. A. — Sylloge Fungorum. Vols. 7, 9, n, 14, 16, 17 1888-1905. 

Mycetes aliquot australiensis. Hedw., Vol. 28, p. 126. 1889. 

Bull. Soc. Myc. France, Vol. 5, p. 118. 


Ibid. Hedw., Vol. 29, p. 155 

Ibid. Hedw., Vol 31, p. 57 



1. Saccardo, P. A., and Berlese, A. N. 

Fungi australiensis. 

Rev. M) 


7> P. 93 



Salmon, E. S. — Mycological Notes. J 

Cultural Experiments with 

Phil. Trans., B., Vol. 197, p. 107 

t c 


184. 1904 

Forms'' of the Erysiphaceae- 


Cultural Experiments with the Barley Mildew, 

'fhe gr a minis 



1. Sappin-Trouffy, P. — La Pseudo-Fecondation chez les Uredinees el les pheno- 

menes qui s'y rattachent. Compt. rend., Vol. 116, p. 1304. 
Recherches histologiques sur la famille des Uredinees. 


Le Botaniste, 

Vol. 5, pp. 59-244 


Shutt. F. T— The effect of Rust on the Straw and Grain of Wheat. Nor'- 

W est Farmer. W 


20th Oct., 1904 

1. Smith, W. G. — Diseases of Field and Garden Crops. London. 


1. Smith, R. E. — The Water relation of Puccinia as far a gi. Bot. Gaz., Vol. 

p. 19 


1. Sorauer, P. — Das. Verhalten des Getreiderostes in trockenen und nassen 

ren. Zeitschr. f. Pflanzenkr., Vol. 4, p. 121. 1894. 

Die Predisposition der Pflanzen fur parasitare Krankheiten. Deutsch.- 

Landwirts. Ges., Berlin. 


1. Sydow, P. and H. — Monographia Uredinearum. Vol. 1. 1Q02-04 

Neue und kritische Uredineen. 





O ^> J 


















i. Tassi, F. — La 


Lab. Bot., Univ. di Siena. November, 1903 
1. Tate, R. — A list of the 

dei Crisantemi {Puccinia chrysanthemi Roze). Bull. 

4, P 

Charas, Mosses, Liverworts, Lichens, Fungi, and Al- 
of Extra-tropical South Australia. Proc. Roy. Soc. S. Australia, Vol. 




J. G. O. 

Red Riist : Its nature, approximate cause and probable cure. 
Proc. Roy. Soc, S. Australia, Vol. 3, 

P- l 3 
Ibid., Vol. 

Contributions to the Fungal Flora of 

Notes on Australian Fungi. 


12, p. 150 

Australia, Vol. 13, p 

1. Tenison-Woods, J. E., 

South Wales and Qu< 

55. 1890. 
and Bailey, 

Proc. Roy. Soc. S. 

F. M. — On some of the 


of New 

Proc. Linn. Soc, N.S. Wales, Vol. 5, p. S3. 

1. Thompson, E. H. — A Hand-book to the Insect Pests of Farm and Orchard 

(including Fungi). 



of Tasmania. 


*> PP- 29-34 


i. Thuemen, F. v. — Symbolae ad floram mycologicam Australiae. I. Grevillea, 


4. P 



Flora No. 28. 1878 

■ 1st der Berberitzenrost 

Oesterr. Landw. Wochenbl 



zur Erzeugung des Grasrostes? 

1. Tranzschel, W 

— Ueber einige auf Grund von irrtlimlicher 
Nahrpflanzen aufgestelle Puccinia Arten. 

Ann. Myc, Vol. 2 



p. 155. 1904 

Moglichkeit, die Biologie wirtswechselnder Rostpilze auf Grun< 
' Merkmale vorauszusehen. Arbeit, der Kais. St. 

Naturf. Gesell. Vol. 35, p. 1. 1904. 
1. Tryon, H. — Report on Insect and Fungus Pests, pp. 97, 196. 1889. 





Vegetable Pathology, pp. 39, 40 

3- " 

4- * 

6. - 

8. - 

Ibid. p. 39. 1895-6. 

Ibid. p. 37. 1890-7. 

Ibid. pp. 43 
Ibid. p. 35 



Ibid, p 



1 899- 1900. 
1 900- 1. 

1. Tubeuf, K. F. V., and Smith, Wm. G. — Diseases of Plants induced by Crypto- 


gamic Parasites. London. 1897. 


Ward, H. Marshall.— Illustrations of the Structure and Life-history of Puc- 

Brit. Assoc. Rep., Glasgow, p. 

cinia graminis. Ann. Bot., Vol. 2, p. 217 
The Bromes and their Brown Rust. 


The Bromes and their Rust-fungus {Puccinia dispersa). Ann Bot., Vol. 

15. P- 5 6 °- 

1 go 1 

On the question of " Predisposition " and " Immunity" in Plants. Proc 

Proc. Roy. Soc, 

Camb. Phil. Soc, Vol. n, p. 326. 1902 

On pure Cultures of a Uredine, Puccinia disp 

Vol. 69, p. 451. 1902. . 

On the Relations between Host and Parasite in the Bromes and their 

Brown Rust, Puccinia dispersa. Ann. Bot., Vol. 16, p. 233. 1902. ^ 

Experiments on the effect of mineral starvation on the Parasitism of the 

Uredine-fungus, Puccinia dispersa, on species of Bromus. Proc Roy. Soc, 
Vol. 71, p. 138. 1902. 

Further Observations on the Brown Rust of the Bromes, Puccinia dis- 
persa, and its adaptive Parasitism. Ann. Myc, Vol. 1, p. 132. i9°3- 

On the Histology of Uredo dispersa Eriks., and the Mycoplasm Hypo- 

thesis. Phil. Trans. B., Vol. 196, p. 29. 1903. 

Recent Researches on the Parasitism of Fungi. Ann. Bot., Vol. 

*9> P 



r. Webber, 


Peridial Cell Characters in the Classification of the Uredineae. 

Amer. Nat., Vol. 24, p. 177. 1890. 

1. Winter, G. — Exotische Pilze II. Hedwigia, Vol. 24, p 

Rev. Myc, Vol 8, p. 208. 

Fungi australienses. 



x. Woroxix, M. — Untersuchungen ueber die Entwickelung des Rostpilzes (Puc* 

cinia helianthi), welcher die Krankheit der Sonnenblume verursacht. 


Zeit., p. 677. 



Explanation of Plates 


(All Figures mat. size.) 


i. Uredosori on upper surface of leaf. 

2. Teleutosori on under surface. 




3. Uredosori on upper and under surface, also on sheath 

4. Teleutosori on stem. 




• I 



- ? ' 



: i 






C C Brzt/JtbanJc.De/,. 





D M:'Alpif?& Dvjc 

• I 








• * 






A c>/ i - :za ^i > ' Pnntr 








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 









JL 1 







if'. * 



D MPJl^ui^Jhrexit 

RSBrauui. u<rvt>Pnn£er. 





4 4 








Explanation of Plates. 


{All Figures nat. size.) 


ii. Uredosori on under surface of leaf. 


. 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. 








• l 



* »■ 

« » ■ 



C C £rUtlehank Del 


I)M c ALpiKt Direxu 

\ S Brow,, (kv 4 * printer 










Explanation of Plates 


{All Figures nat. size.) 

PUCCINIA CICHORII on CHICORY (Cichorium intybus) 

18. Chiefly uredosori on lower surface of chicory leaf. 


P. armeniaca). 

19. Uredo and teleutosori on lower surface of peach leaf. 

20. Uredo and teleutosori on lower surface of apricot leaf 


^M' ( ^k 



DM c Alpuie i Direxiz 

F S Bra j/ w T p -utter 








y - 




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 hederacea 

25. Aecidia on under surface of leaf of V. betonicijolia. 

PUCCINIA DISTINCTA on DAISY (Bellis perennis). 

26. Aecidia and teleutosori on upper surface of leaf, showing the dark teleutofori 

surrounding the other. 



DWAlpuze,, Direxvt 

R. S.Braui GwtPrviUr 




■ ■ 

































- f 











Explanation of Plates 


{All Figures nat. size.) 



27. Aecidia and teieutosori on both surfaces of leaves. 


28. Teieutosori on under surface of leaf and on leaf -stalk. 



29. Aecidia, uredo, and teieutosori on under surface of leaf, and teieutosori on stem 



CC£nttUi?a7ih f Dd 

DM C Alp 1716 Direjcio 

. . "ma ■ . ' r> • v- 






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. 





D AC c llfiVU,, Dtrtxib 

R S Bravi, 6ov t Pnnler. 














Explanation of Plates. 


[Natural Size.) 


BETAE ON MANGEL (Beta vulgaris) 


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. 


* t • •• * 

c I > 


' ' 


• ' •• 




* 9 ,- 


' . 

' ' 





. -* 


2 3 8 

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. 


Y ' BnttUharJi. Dd 

'■' ft ■ ■■ DirexU 

$. /train C*>\* P> 
















Explanation of Plates 





39. Leaves with aecidia in dense clusters 

nat. size 

40. Cluster of aecidia on leaf ... 

X 15 

41. Section of aecidium showing peridial wall and spores in chains 

42. Chain of aecidiospores 

X 50 

.. X 1000 

43. Single spore detached and more or less rounded 

.. X 1000 

44. Pseudoperidial cells seen from the surface 

.• X 














C. CJtrzttUbcuiJi. Dd 

DIFAlputfi, Diructi, 




Explanation of Plates. 



(All Figures X 250 unless otherwise stated.} 



1. Portion of sheath of barley (Hordeu?n vulgare) showing numerous crowded black 

dots, the teleutosori of Puccinia simplex, the 


spring or 

• • • 

orange rust of 
• .. nat. size 

2. Portion of sheath of wheat [Triticum vulgare), the broad dark lines representing 

the uredosori of P. graminis, summer rust of wheat 

nat. size 

3. Portion of flag of wheat, the small dark dots representing the uredo and teleu- 

tosori of P. triticina, the spring or orange rust of wheat 

nat. size 

4. Uredospores of P. simplex from barley, the numerous germ pores appearing as 

paler dots scattered promiscuously over the face of the spore. 

5. Uredospores of P. graminis from wheat, w 7 ith 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. triticifia on wheat, with paraphyses at left. 

Note. — The whole of the photomicrographs in this and the succeeding plates were 
taken with the horizontal laboratory camera made by Messrs. Watson and Sons, of 
London, using their Holoscopic lens of 12 mm. .65 N.A. to obtain the magnifica- 
tion of 250 diams., and the 6 mm. .95 N.A. for 500 diams. 



Platk I 

{,. II. Robinson, Phot. 

Sut. size and v iou 







• r 





Explanation of Plates 


(All Figures X 250.) 




ii. Section of teleutosorus of Puccinia lolii avenae on oat (Avena saliva) with severaE- 

unicellular as well as bicellular teleutospores. 

12. Group of teleutospores of P. beckmanniae on Beckmannia erucaeformis 

13. Section of teleutosorus of P. festucae on Festuca fratensis. 

14. Section of teleutosorus of P. lolii on Lolium -perenne. 

15, 16. Groups of teleutospores of P. tnaydis on Zea mays, including one four 

celled spore. 

17. Teleutospores of P. flavescentis on Stifa flavescens. 

18. Section of teleutosorus of P. magnusiana on Phragmites communis. 

19. Five teleutospores of P. tefferi on Phragmites communis, showing the extremeljT 

long pedicels. 

Plate II. 

G. II. Robinson, Phot. 

x 250 







Explanation of Plates. 


{All Figures X 250.) 


20. Section of teleutosorus of Puccinia anthoxanthi on Anthoxanthum odoraium, one 

spore with a septate pedicel laterally inserted. 

21. Group of teleutospores of the same, one with a much inflated pedicel. 

22. Teleutospores of P. -poarum on Poa annua. 

23* Teleutospores of P. ferflexans on Alofecurus geniculates 

24. Teleutospores of P. cynodontis on Cynodon dactylon. 

25. Section of teleutosorus of P. agrofyri on Agrofyron scab rum 

26. Group of teleutospores of P. imfatientis on Elymus condensatus. 

27. Group of teleutospores of P. agrostidis on Deyeuxia forsteri 

28. Section of teleutosorus of P. bromina on Bromus mollis. 

Platk III. 

G. H. Robinson, Phot. 

x 250 



• * 

' » 







t • 




. ■ 


•J SiiS ,; l i- • v * . « • 




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. longisfora on Carex caesfitosa. 

32. Teleutospores of P. cyferi on Cyferus rotundas. 

33. Group of teleutospores of P. tenuisfora on Luzula eamfestris, Murramurrangbong 

Ranges, Victoria. 

34. Teleutospores of P. obscura on Luzula camfestris, Berlin, Germany (Sydow, Ure- 
dineen, 1076). The spores of this species are differently shaped, much thicker 
in the wall and much darker than those of P. tenuisfora. 

35. Uredospores of P. juncofhila from sorus containing both uredo and teleutospores 

on 1 uncus maritimus. 

36. Teleutospores of the same. 



Plati; I \ . 

G^ H. Robinson, Phot. 

x 250. 



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 Bur char dia umbellata. 

39. Teleutospores, one three-celled, of the same. 


40. Teleutospores and mesospores of P. haemodori on Haemodorum sp. 

41. Teleutospores and mesospores of P. hyfoxidis on Hyfoxis glabella. 

42. Teleutospores, two three-celled, of P. dichondrae on Dichondra re-pens. 

43. Teleutospore and uredospore of P. tnussoni on Ruellia australis^ Richmond River, 

New South Wales. 

44. Teleutospores of P. ruelliae on Ruellia strefens, Columbus, Ohio, U.S.A. (W. 

A. Kellerman, Ohio Fungi, 130). 

Plate V. 

(i. 1J. Robinson, Phot. 

x 2."". 













Explanation of Plates. 



{All Figures X 250.) 


45. Group of teleutospores of Puccinia carissae on Carissa ovata 

46. Section of teleutosorus of P. alyxiae on Alyxia buxifolia, with teleutospores and 


47. Teleutospores of P. gilgiana on Leschenaultia linarioides. 

48, 49. Teleutospores and mesospores of P. saccardoi on Goodenia geniculate 

50. Teleutospores and mesospores of P. brunoniae on Brunonia au straits, with one 

three-celled and one four-celled teleutospore. 

51. Teleutospores of P. aucta on Lobelia ancefs. (From a drawing by C. C. Brittle- 

1 bank from the original material from Berkeley in the National Herbarium, 

Plate VI 

G. H. Robinson, Phot 

x 250 










Explanation of Plates. 


(All Figures X 250.) 



52. Teleutospores and mesospores of Puccinia t as manic a on Senecio vulgaris, one 

teleutospore being three-celled. 

53. Teleutospores and mesospores of P. angustifoliae on Scorzonera angustifolia. 

54, 55. Sections of teleutosori of P. lagenofhorae on Lagenofhora billardieri y show- 

ing mesospores and teleutospores. 

56. Teleutospores and mesospoies of P. helianthi on Helianthus annuus. 

57. Teleutospores of P. gnafhalii on Gnwphalium jafonicum. 

58. Teleutospores and mesospores of /\ calocefhali on Calocefhalus drummondii. 

59. Teleutospores and mesospores of P. erechtitis on Erechtites quadridentata. 

Plate VII. 

G. II. Robinson, Phot 

x 2.")0 







Explanation of Plates 



{All Figures X 250.) 


60. Section of teleutosorus of Puccinia cinerariae on Cineraria sp. cult, with teleuto- 
spores and mesospores. 

61. Three uredospores and several teleutospores of P. cichorii on Cichorium intybus 

62. Uredospores and teleutospores of P. hyfochoeridis on Hyfochoeris radicata. 

63. Teleutospores of the 


64, Two uredospores and numerous teleutospores of P. cyani on Centaur ea cyanus. 


65. One mesospore and several teleutospores of P. calendulae on Calendula officinalis 

66. Mesospores and teleutospores of P. brachycomes on Brachycome ciliaris. 

67. Teleutospores and mesospores of P. distincta on Bellis ferennis. 

Platk VII! 

('•. H. Robinson, Phot 

x 250 





2 5 8 

Explanation of Plates 


(All Figures X 250 unless otherwise stated.) 


* • 

' l g- 


68. Three uredospores and four teleutospores of Puccinia thuemeni on Afium fros- 

tratum^ showing thickened apex of uredospore and slight warting of teleuto- 


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- 


• * 

• • • 

• • 

• • 

• * 

x 500 

71. Teleutospores of P. thuemeni on Afium graveolens, from Brighton, Victoria. 

72. Uredospore and teleutospore from the same as 71, showing roughened epispore 

in both forms more clearly 

• • 

• • 

• • . 

X 5°° 

73. Uredospores and teleutospores of P. afii on Afium graveolens, Berlin, Ger- 
many. (Sydow, Uredineen, 558.) 

74. Teleutospores as in Fig. 73, showing distinctly smooth epispore 

X 500 

75. Teleutospores of P. btillata on Aethusa cynafium^ with coarsely warted epispore. 

(Sydow, Uredineen, 1261.) 

• • 

• • 

c • 

X 5°° 

76. Teleutospores of P. xanthosiae on Xanthosia fusilla. 

Platk IX 

G. II. Robinson, Phot. 

x 230 ami 5u«>. 








(All Figures X 250.) 



77. Teleutospore of Puccinia oliganthae on Asferula oligantha 

78. Teleutospores, some having germinated, of P. cofrosmae, on Cofrosma hirtella 

79. Uredospores and teleutospores of P. efilobii-tetragoni on Efilobium glabel/um, 

Murramurrangbong Ranges, Victoria. 

80. Uredospores and teleutospores of P. efilobii-tetragoni on Efilobium montanum. 

(Sydow, Uredineen, 1369.) 

Si. Teleutospores of P. efilobii DC. on Efilobium roseum, the teleutospores having 

much thinner walls, and being more variable in shape than those of P. 
efilobii-tetragoni and also finely verrucose. (Sydow, Uredineen, 1418.) 

82. Teleutospores of P. gei on Geum reni folium. 

83. Uredospores of P. fruni on Prunus fcrsica. 

84. Paraphysis and uredospores of P. fruni on Prunus fcrsica. 

85. Teleutospores of P. fruni on Prunus fersica^ rather smaller than average 

86. Teleutospores of P. fruni on Prunus domestica. 

87. Uredospores and teleutospores of P. zorniae on Zornia difhylla. 


Platk X. 

G. II. Robinson, Phot. 

x 25u 












(All Figures X 250.) 


88, 89. Teleutospores of Puccinia tetragoniae on Tetragonxa imflexicoma, one with 

nearly vertical septum in upper cell. 

90. Teleutospores, strongly warted, of P. ludwigii on Rumex flexuosus. 

91. Teleutospores strongly warted, of P. ludwigii on Rum ex brownii, one spore being 


92. Uredospore and teleutospores of P. acetosae on Rumex arifohus^ all the latter 

with smooth walls. (Sydow, Uredineen, 954.) 

93. Teleutospores of P. mue/ilnibrcfciar on MuehlenbecTzia adfressa. 

94. Teleutospores and mesospore of P. diclsiana on Threlkeldia sp. 

95. Uredospores of P. kochiae on Koclria sedifolia, showing the characteristic nume- 
rous germ pores. 

96. Teleutospores of P. kochiae on Kochia sedifolia. 


Plate XI 

G. II. Robinson, Phot. 

x UoO 




> ' 



Explanation of Plates 


(All Figures X 250.) 



1 or 

97. Teleutospores of Puccinia arenariae on Stellaria media 

98. Teleutospores of /\ -plagianthi on Plagianthus sidoides, the epispore 

slightly channelled. 


99. Teleutospores (stained) of P. malvacearum on Malva rotundij alia. 

100. Teleutospores and mesospore of P. malvacearum on Lavatcra flebeia. 

101. Teleutospores, mostly one-celled, of P. heterosfora on Abutilon crisfum 

T02. Teleutospores and mesospores of P. morrisoni on Pelargonium australe. 

103. Uredospores of P. geranii-filosi on Geranium filosum. 

104. Teleutospores of the 

same, the single germ pore being 

noticeable as a paler 

channel through the apex of the cell wall. 



G. H. Kobinson, Phot. 

x 250 







-. — _ ' 






(All Figures X 250 unless otherwise stated.) 


05. Teleutospores of Puccinia eriostemonis on Eriostemon myo-poroidcs. 

106. Teleutospores of P. correae on Correa lawrenciana. 

107. Teleutospores and mesospores of P. boroniae on Boronia sfinescens. 

108. Teleutospores and mesospores of P. fritzeliana on Tremandra stelligera, some 

teleutospores having germinated. 

109. Teleutospores of P. hederaceae on Viola hederacea. 

no. Teleutospores of the same, more highly magnified, to show more clearly the 

slightly warted epispore 

- . - 

• • • 

• * 

• • 

X 500 

in. Teleutospore of P. aegra on Viola tricolor with smooth epispore. (W. B. Grove 

in Rabh. Fungi Europaei, 31 13) 

• • • 

• • 

X 5°° 

112. Teleutospores of P. violae on Viola arenaria, wtih smooth epispore. (Sydow, 

Uredineen, 286) 

- • • 

• • • 

X 5°° 


L'late Mil 

G. II. Robinson, Phot. 

x 250 ami 5l0 






Explanation of Plates 


(All Figures X 250.) 



113. Teleutospores and mesospores on wild oat, Avena fatua. 

114. Teleutospores on barley, Hordeum vidgare. 

11^, 116. Teleutospores, mesospores, and uredospore, one of the first three-celled, on 

native barley, Echinofogon ovatus. 

117. Teleutospores on cocksfoot, Dactylis glomerata. 

118. Teleutospores and uredospores on silver grass, Festuca bromoides 

ii(). Teleutospores and uredospores on an imported barley grass, Hordeum secalinum 

120. Teleutospores on small canary-seed grass, Phaiaris minor. 

121. Teleutospores on native wheat grass, Agrofyron scabrum 

122. Teleutospores on Amfhibromus neesii. 

I'j.ATK XI \ 


G. H. Robinson, Phot. 

X 2'MI 



I ; 









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 



125. Two germinating teleutospores, in the one on the left only the two median seg- 



ments of the promycelium as yet bear sporidiola. 
Germinating teleutospore with stout elongated and 

contorted filaments 


from the promycelial cells, those from the two median cells ultimately giving 
rise to abnormally small sporidiola. 

127. Germinating teleutospore in w r hich the upper two promycelial cells have produced 

sporidiola, while in the third from the apex the formation of the sporidiolum 

is just commencing. 

Germinating teleutospore, the promycelium bearing three 

sporidiola on stunt 

The sporidiolum second from the base is already 
germinating and putting forth a germ tube extending across to the promy- 

apex has not so far produced a 

elongated sterigmata. 


The segment second from the 

sporidiolum, though a stout elongated filament has been emitted. 

129. Promycelial spores 

130. Germinating promycelial spore 

• - 

- - - 

X 500 

• • 

• • * 

• • • 

X 5°° 

Note. — All the above were grown in droplets of water on slides in a moist cham- 
ber, and prepared for photographing by being first dried, then fixed with a saturated 
solution of mercuric bichloride, washed, rinsed in dilute acetic acid, washed again, 
and after drying stained with alcoholic Bismarck brown, again washed and mounted 
in glycerine and water. 



G. H. Robinson, Phot. 


x 250 & 50 1 








Explanation of Plates 

* • 



[All Figures X 250.) 


131. Group of teleutospores of Uromyces danthoniae on Danthonia semi annularis 

132. Teleutospores of U. ehrhartae on Micro Laena [Ehrharta) stifoides. 

133. Teleutospores of U . tenuicutis on S-porobolus indicus. 

1 ^4. Two one-celled and one two-celled teleutospore of U . tricorynes on Tricoryne 


135. Teleutospores of U . tricorynes on Tricoryne elatior. 

136. Uredospores of U . thelymitrae on Thelymitra antennifera 

137. Teleutospores of the same 

138. Teleutospores of U. orchid earitm on Chilo glottis difihylla, from original type 

material in National Herbarium. 

130. Teleutospores of U . microtidis on Microtis forri folia from type material from 


Platk XVI 

G. H. Robinson, Phot. 

X -■"•" 












Explanation of Plates 



(All Figures X 250.) 


140. Group of teleutospores of Uromyces fuccinioides on Selliera radicans. 

141. Teleutospores of U . as ferulae on As ferula oligantha. 

142. Uredospores and teleutospores of U. trifolii on Trifolium refens 

143. Uredospores of U . hardenbergiae on Hardenbergia monofhylla. 

14a. Teleutospore of the same. 

145. Two uredospores of same at top and one teleutospore below 

146. Section of teleutosorus of U . atriflicis on Atriflex semibaccata 

spores are faintly striate with one prominent germ pore at apex. 

The teleuto- 

147. Two teleutospores of same at left and four uredospores at right. 

148. Uredospores of U. brtae on Beta vulgaris 

149. Teleutospores of same with prominent hyaline apiculus. 

Platk XVII 

G. II. Robinson, Phot. 













Explanation of Plates. 


(All Figures X 250 unless otherwise stated.) 




150. Uredospores of Uromyccs folygoni on Polygonum aviculare 

151. Teleutospores of same. 

152. Uredospores of U. caryofhyllinus on Dianthus caryofhyl/us, with prominent 

scattered germ pores. 

153. Teleutospores and one uredospore of same. 

154. Section through leaf of Dianthus caryofhyllus, showing uredosorus of U. caryo- 

-phyllinus with two pycnidia of Darluca filum growing upon it, and two 

young pycnidia on opposite surface of leaf 

• • • 

155. Uredospores of U. vesiculosus on Zygofhyllum glaucescens. 

156. Teleutospores of same. 

157. Two-celled teleutospore of same 

X 5° 

Platk XVIII 


G. H. Robinson, Phot. 

X 50 & 250 






■ > 











, Si 








{All Figures X 250 unless otherwise stated.) 


158. Teleutospores, seen from above, of Uromyces fusisforus on Acacia nerii folia 

159. Side view of same. 

160. Uredospores of same, with pronounced apiculus and equatorial band of germ 



161. Teleutospores of Uromycladium simplex on Acacia fycnantha, part of the 

pedicel being still attached with the remains of the vesicle. (Stained). 

162. Young teleutospores of the same showing the vesicles still attached. (Stained). 

16}. Portion of flake of gummy material on leaf of Acacia fycnantha, in which nume- 
rous teleutospores of U. simplex are embedded, nearly all germinating and 

producing sporidiola. 

164. Germinating sporidiolum of same. (Stained) 

• • 

x 5°° 

165. Uredospores of same, being very similar to those of Uromyces fusisforus, fig. 

160, though somewhat shorter and broader and' possessing more germ pores. 




G. H. Robinson, Phot. 

X 250 & 500, 











Explanation of Plates. 


{All Figures X 250.) 



j66. Section of teleutosorus of V \ maritimum on Acacia longifolia^ the mature 

teleutospores having the remains of the vesicles attached to the pedicels. 

167. Group of mature teleutospores of the same. 

168. Formation of teleutospore head ; a young cluster, showing the two young spores 

at the apex, and the lateral vesicle below the septum. (Stained.) 

169, 170. Similar heads, more advanced. (Stained.) 

171. Mature teleutospore head, two teleutospores above and one vesicle below. 


Abnormal development of head of teleutospores, the lateral vesicle being re- 
placed by a true spore, and a septum placed in the stalk beneath it. In the 
normal head of two spores and a vesicle there is no septum below the vesicle. 

173. Teleutospores germinating while still attached to each other : two sporidiola on 

one promycelium, others so far undeveloped. 

174. Uredospores with prominent equatorial germ pores and serrated epispore 

thickened and dentate at apex. 

75. One-celled spores (mesospores ?), occasionallv found " intermixed with uredo- 

176. Immature two-celled colourless spore, of same character, from .uredosorus. 


G. 11. Robinson, Phot. 

X 2 50. 






K 2 


Explanation of Plates 


(All Figures X 500.) 

177. Two 



teleutospore heads, 

right being younger than 
T2 and T3 in next figure. 

arising from common stalk, the one on the 
that on the left, and both intermediate between 


178. Main stem, 

bearing teleutospore 

heads in various stages of development; T, 

filament with two septa, the upper two segments destined to become teleuto- 
spores, the lowermost a vesicle ; Tl, similar filament, 9omewhat more ad- 

vanced, the vesicle beginning to push out from the lowermost segment; T2, 
similar filament still more advanced, the uppermost cell distending, in the 

next growth is taking place at the side, and in 

the lowest the vesicle has 

attained considerable development; T3, nearly mature head, with tw r o 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. 

1S1. Teleutospore head in which the vesicle is apparently at the apex, but this is 

probably due to displacement in mounting. 

182, 183. Normal teleutospore heads. 

184. Uredospores with prominent germ pores, serrated epispore thickened and dentate 

at apex. 



(J. II. Robinson, Phot. 

-I- ;uu 







Explanation of Plates. 



(All Figures X 250 unless otherwise stated.) 



1 i ^- 

185, 186. Uredospores of U. robinsoni on Acacia melanoxylon. The solitary large 

smooth-walled spore (?) in Fig. 186 is difficult of interpretation. 

187. Young and mature teleutospores of same, with and without attached vesicles. 


188. 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. 

tefferianum on Acacia salicina^ the striated epispore being clearly shown in 
192. (Stained.) ... ... ... ... ... ... X 500 

193. Single teleutospore of same, seen from above, and showing striated mark- 

• • • ••• ••• • • • ••• ••• ••• ••• ^\ ^^^^^ 


194. Mature teleutospore head of same. 

of poor condition of material) 

(The markings are indistinct on account 

X 500 

• • • 

195. Detached teleutospores of same. 

Platk XXII 


G. II Robinson, 

x 250 & 510. 










Explanation of Plates. 


(All Figures X 250 unless otherwise stated.) 


196. Uredospores of U. notabile on Acacia notabilis. 


Uredo notabilis Ludw. 
caustic potash.) 

type material of 

(Somewhat swollen by lengthened treatment with 

197. Uredospores of the same, showing the net-like surface markings 

X 500 

iq8. Compound stem, with basidia and uredospores of the same. 

199. Compound structure from uredosorus of same, being probably commencement 

of teleutospore formation 

• • • 

• • 

• • 

X 5°° 

200. Uredospores of U. notabile on Acacia dealbata. 

201, 202. Uredospores of the same, showing the net-like surface markings... X 500 

203. Portion of spermogonium of the same, with basidia bearing spermatia in 

chains. (Stained) 

* • • 

• • • 

X 500 

204. Cluster of three immature spores of the same, attached to their stalk. (Stained.) 

205. Mass of mature teleutospores of the same, separated from each other, from 

Acacia decurrens. 

206. Mass of scarcely mature teleutospores, separated from each other, of V . tef- 

ferianum, on Acacia melanoxylon^ from Cheltenham, showing striated mark- 
ings on the surface. These are considerably smaller than the average. 

Plate XX HI 

G. H Robinson, Phot. 

x 250 & 600. 





Explanation of Plates 


(All Figures X ~jo unless otherwise stated.) 



207. Teleutospores of U. bisforum^ on Acacia dealbata, one cluster of t 

the common staik. 

o showing 

208. Teleutospores of the same. 

209. Uredospores of U. alfinum, on Acacia dealbata. 

210, an. Teleutospores of the same. 

212. Five uredospores of U. alpinum on Acacia dallachiana, mixed with teleutospores 

and one mesospore (m). 

13, 214. Successive stages in the development of teleutospore clusters of the same 


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 


• * • 

• • 

# • • 

• • ■ 

X 30 

217. Uredospore of U. maritimum attacked by some Hyphomycete, which has gained 

access to the interior of the spore and grown within it preparatory to forming 
the long septate threads which have pierced the wall near the germ pores. 

Plate XXIV 

(J. II. Robinson, Phot. 

x 30 & L'6U 






Explanation of Plates. 


{All Figures X 250 unless otherwise stated.) 



218. Two uredospores of U. fhyllodiorum (B. aud Br.) McAlp., on Acacia sp. from 


Queensland. (Type material of Melamfsora fhyllodiorum B. and Br., 
from Herbarium of F. M. Bailey, Government Botanist, Queensland.) 

219. One of the very sparse digitate teleutospores present in the same material. 

220. Uredospore from the same material 

• • • 

X 500 

(All the above from old and much faded material.) 

1, 2 

22. Uredospores of U. fhyllodiorum, on Acacia sp. from Queensland. (Type 

material of Uromyces fhyllodiae Ccoke rnd Mass., these uredospores 
described as teleutospores. From Herbarium of F. M. Bailey.) 



223, 224. Two of the few digitate teleutospores present in the same material 

(Material of above old and much faded.) 


225. Uredospores of U . fhyllodiorum on Acacia dallachiana, from Bright, Victoria. 

226. Two of the same, more highly magnified, to show surface markings ... 

X 500 

227, 228. Teleutospores from the same material, some beginning to germinate 


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 Rub us farvifolius, two 

germ pores being often seen on one face of each cell. 

236. Teleutospores, closely attached to each other, of Melamfsora lini on Linum 




G H. Robinson, Phot. 

Nat. size & X 250 







Explanation of Plates 




237. Young 

seedling of Platylobium formosum, with Aecidium flatylobii on leaves 

and leaf stalks 

nat. size 

2i : 8. Fruiting branch of the same, with aecidial cups on the pods 

nat. size 

239. Section through cups of Aecidium eburneum on pod of Bossiaea hetero- 

X 5° 


• • • 

• • 1 

240. Section of aecidial cup of Puccinia erechtitis on Erechtites quadridentata X 50 

t'LA'VK XXV [I. 



G. H. Robinson, Phot 

Nat, size & x 50 








Explanation of Plates 




{All Figures X 250). 



241. Spore of Uredo scirfi-nodosi or Scirfus nodosus 

242. Uredospores of Puccinia stylidii on Stylidium graminijolium. 

243. U. bidentis on Bidens filosa from Queensland. 

244. U. bidentis on Bidens filcsa from Brazil. (Sydow, Uredineen, 1647.) 

245. U . sfyridii on Sfyridium farvifolium 

246. U. bossiaeae on Bossiaea frostrata. 

247. U. fallidula on Cassia sp. 

24S. Uredospores of Uromyces scleranthi on Scleranthus diander. 

249. Uredospores of Puccinia oleariae on Olearia argofhylla^ epispore finely striate. 



Plate XXVI II 

G. H. Robinson, Phot 

X 250 



* t 










Explanation of Plates 



{All Figures X 250.) 


250. Uredospores and leleutospores of Puccinia menthae on Mentha fulegium 

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., 

Japan. (Herbarium A. Ideta.) 

254. Abnormal teleutospore associated with uredospores and teleutospores of same. 

255. Mesospore from same. 

256. Group of teleutospores and mesospores of P. -podolefidis on Podolefis longu 


257. Teleutospore from same, with hyaline germ pore at each side of apex of upper 


258. Teleutospores of P. oferculariae on Ofercularia varia. 


Platk XXIX 

G. H. Robinson, Phot. 

X 250. 







Explanation of Plates 


(All Figures X 250.) 



259. Uredospores of Puccinia cacao on Rollboellia comfressa 

260. Uredospores and one teleutospore of same 

261. Uredospores of P. lolii avenae on Avena saliva, germinating in water after 

24 hours. (Stained.) 

262. Uredospores of P. subnitens on Distichlis maritima 

263, 264. Teleutospores from same. 



266. Teleutospores of P. crucifrrae on an undetermined Crucifer. From type 

material referred to Aecidium barbareac DC, at Kew. 

267. Uredo rhagodiae on Rhagodia billardieri. 

Platk XXX 

I * 

G. H. Robinson, Phot. 

x 2:>0, 


• « 


* * 





302 Explanation of Plates. 


[All Figures X 250 unless otherwise stated.) 


26S. Leaf of Loranthus celastroides^ showing uredosori and teleutosori of Puccinia 


• • • • • • • 

... nat. size 

69. Teleutospores from same. 

270. Uredospores from s^me. 

271. Leaf of Loranthus celastroides, with aecidia of same ... ... nat. size 

272. Teleutospores of Phragmidium longissimum. 

a. Four and five celled spores, before germination. 

b. Spores commencing to germinate, with one germ tube divided by a 


c. Spore cell, giving rise to four-celled promycelium, with loose sporidiola 

around it. 

The material was fourteen years old, and although there were plenty of sporidiola, 
none were found attached to the promycelia. 

Note. — Figs. 272, a, b, c, from drawings. 

Plate XXXI 


G. H. Robinson, Phot 

x 2f>0 & 5<m>. 






> . 






Explanation of Plates. 





273. Leaves of Acacia longifolia, with the tuberculate spermogonial sori of Uromy- 

cladium maritimum 

• • • 

• • • 

• • • 

nat. size 

274. Leaves of Acacia melanoxylon^ the upper with tuberculate spermogonial sori, and 

the lower with minute pulverulent sori, both of U. robinsoni. ... 

nat. size 

275. Leaf of Acacia fycnantha, with the minute powdery sori of U . simflex. nat. size 

Plate XXXII 

O. H. Kobinson, Phot. 


Nat. size 




.. » 

*■ I 

• (- 

-< - - 

4 » 


Explanation of Plates. 




276. Sori of U. bisforum on stems and leaf stalks of Acacia dealbata 

nat. size. 

277. Sori on fruits 

• • • 

• • 

nat. size 


G. H. Robinson, Phot. 

Nat. size 



3°8 Explanation of Plates 




278. Branch of Acacia armata showing the powdery galls caused by the growth of 

U. tefferianum 

• • • 

nat. size 

Plate XXXIV 



■ ■ 


. ■ v» ■ ■ ■ 

■ ■ 


■ ■ 





■ - 





. ■ 


* 4 I* 

G. H. Robinson, Phot 

Nat. size. 









Explanation of Plates 





279. Galls attached to branches of Acacia fycnantha, caused 



• • • 

• • 

U. tef. 

nat. size 

280. Galls detached 

• • • 

• • 

nat. size 

Pi-ATK XX \\ 

G. H. Rot inson, Phot. 

Nat, size 

















» ■ 







Explanation of Plates 



281. Numerous young galls on Acacia dealbata, bearing the uredospores of U. 

« « • • • • ••• • • • ••• llclL.ol L* \Z 


• • • 

• • • 

(Being on the young growth of the current season, the galls are necessarily 
comparatively small, though many of several years 5 growth bearing teleuto- 
spores have been found as large as potatoes, and weighing nearly 1 lb. 

Plate XXXVI 

G. H. Robinson, Phot. 

Nat. size 










• J 

• ■ 





• * 






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 Gomfholobiitm lati folium 

nat. size 

2S6. Normal healthy shoot of Gomfholobium latifolium 

nat. size 


G. H. Robinson, Phot. 

Nat. size 



1 - 

• k 

* - * 


















v - 

* • 

• . 



* • 








Explanation of Plates. 


(All aecidia X 50, and aecidiosfores X 300.) 



287. A. veronicae on Veronica sp 

288. A. flantaginis-variae on Plantago varia. 

289. A. lobeliae Thuem., on Lobelia fratioides, described in connexion with Puccinia 

aucta. The free margin ruptures irregularly, and it is sometimes difficult to 
detect a peridial wall, so that it partakes of the nature of a Caeoma. 

290. A. cymbonoti on Cymbonotus lawsonianus. 


291. A. monocystis on Abrotanella forsterioides. 

292. A. vittadiniae on Vittadinia australis. 

293. A. soleniiforme en Goodia loti folia. 

294. A. deeringiae on Deeringia celosioides. 

295. A. calthae on Caltha introloba. 

206. A. ranunculacearum on Ranunculus rivularis. 

297. Aecidium of Uromyces folitus on leaves of Muehlenbeckia cunninghami — a 

nat. size, b. aecidial tubes X 5. 

298. Aecidiospores of same(X 300) and pseudoperidial cells (X 150). 

Platk XXXIX 

C. C. Brittlebank, Del. 

X 5, 50, 15U & 30U 



I • . — 







Explanation of Plates 


(All Figures X 500.) 




209. The teleutospores exhibit a great variety of shape and size, and range from one 

to four celled forms. In the two-celled forms the septum may either be 
transverse or longitudinal, and in the three-celled forms the septa may 
either be transverse or oblique, and the upper or lower cell may be divided 
longitudinally. The four-celled forms may, in addition, become somewhat 
spherical, and be divided longitudinally and obliquely, so as to resemble 
a Sfhaero-phragmium. The variation even in the same sorus is sometimes 
greater than it is between some different species. 


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. 


Plate XL 


G. C. Brittlebank. Del. 

X 6UU. 





' ■ 


% * 



Explanation of Plates. 




Large gall on Acacia imflcxa, weight 3 


• • • 

X * 

G IJ. Robinson, Phot. 

X J 











3 2 4 

Explanation of Plates. 





305. Witches' broom on Acacia im-plexa, due to Uromycladium tefferianum 

X i 


306. Teleutospores of Uromyces affendiculatus on Vigna catjang or Cow-pea, from 

Richmond, New South Wales 
307. Teleutospores of Uromyces fabat 

# • 

x 25° 

on Lathyrus venosus. (Sydow, Uredineen, 


• • 

• • 

• • 

• * • 

X 210 



G. 11. Robinson, Phot. 

x { &, 2*>u 





• . ~ 

•if o ■ ■• ; 









3 26 

Explanation of Plates. 


(All Figures X 250.) 



308. Teleutospores of P. vittadiniae, mostly of a short broad type, on Vittadinia 

austral is. 

309. Teleutospores and mesospores of the same. 


310. Teleutospores and mesospores of P. calotidis on Calotis sp 

311. Uredospore of P. 

branching growth at 




on Avena Jatua, germinating in water, showing 
end of twenfy-four hours. ^Stained with Congo 

312. Mesospore of P. calendulae on Calendula officinalis, germinating in water after 

three days, two sporidiola produced, both of which have already germinated. 
(Stained with Bismarck brown.) 


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 show r n 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. folitus, one 


being two-celled, 

on Muehlenbeckia cun- 

318. Uredospores with two bands of germ pores, and teleutospores of U. bicinctus 

on Acacia fasciculifera. 

319. Uredospores and teleutospores of U. folycnemi on Polycnemum fentandrum 


G. II. Robinson, Phot. 

x 250 


• **ir. ' 






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. 


IVvn WAY. 

3-colour process 

I . « . .* 


Straw perfectly clean 

R. S. Hkain, (iovt. l'r 


Straw ba«ll\ ni-' • 1. 






J * 







- * 
















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. 

Uromycladium 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 




Acacia juniperina Willd. 

Uromycladium tepperianum (Sa 

Acacia linifolia Willd. 


Acacia longifolia Willd 




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.) 

( Ludw. ) 

Uromycladium notabile 

Acacia penninervis Sieber. 

Uromyces phyllodiorum (B. and Br. ) 

Acacia pruinosa A. Cunn. 

Uromyces phyllodiorum (B. and Br.) 

Uromycladium 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 Lindl. 

Uromyces fusisporus, Cke. and Mass. 
Uromycladium tepperianum (Sacc) 



Host Index. 

Acacia siculiformis A. Cunn. 

Uromycladium tepperianum (Sacc. ) 

Acacia spinescens Benth. 

Uromycladium tepperianum i 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. 


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 tbuemeni, McAlp. 

Apium prostratum Labill. 

Puccinia thuemeni, McAlp. 

Asperula oligantha F.v.M 

scoparia, Hook. f. ) 

Uromyces asperulae, McAlp. 
Puccinia oliganthae, McAlp. 

Atriplex semibaccata R. Br. 

Uromyces atriplicis, McAlp. 

Avena fatua L. 

Puccinia graminis, Pers. 
P. lolii avenae. 

Avena sativa L, 

Puccinia graminis, Pers. 
P. lolii avenae. 

Beckmannia erucaeformis Host. 

Puccinia beckmanniae, McAlp. 
P. graminis, Pers. 

Bellis perennis L. 

Puccinia distincta, McAlp. 

Beta vulgaris L. 

Uromyces betae (Pers.) Kuehn. 




Boronia spinescens Benth. 

Puccinia boroniae, P. Henn. 

Bossiaea cinerea R. Br. 

Aecidium eburneum, McAlp. 
Cronartium jacksoniae, P. Henn. 

Bossiaea heterophylla Vent. 

Aecidium eburneum, McAlp. 

Bossiaea linophylla K. Br. 

Aecidium eburneum, McAlp. 

Bossiaea microphylla Sm. 

Aecidium eburneum, McAlp. 

Bossiaea prostrata R. Br. 

Uredo bossiaeae, McAlp. 

Bossiaea rhombifolia Sieber. 

Aecidium eburneum, McAlp. 

Brachycome ciliaris Less. 

Puccinia brachycomes, McAlp. \ 

Brachycome diversifolia Fisch. and 

Puccinia brachycomes, McAlp. 

Brachycome pachyptera Turcz. 

Puccinia brachycomes, McAlp. 

Brachycome scapiformis DC. 

Puccinia brachycomes, McAlp. 

Briza minor L. 

Puccinia graminis, Pers. 

Bromus arenarius Labill. 

Puccinia bromina, Eriks. 

Host Index. 


Bromus mollis L. 

Puccinia bromina, Eriks. 

Bromus racemosus L. 

Puecinia graminis, Pers. 

Bromus secalinus L. 

Puccinia graminis, Pers. 

Bromus sterilis L. 

Puccinia graminis, Pers. 

Brunonia australis Sm. 

Puccinia brunoniae, McAlp. 

Bulbine bulbosa Haw. 

Uromyces bulbinis, Thuem. 

Burchardia umbellata R. Br. 

Puccinia burchardiae, Sacc. 

Calendula officinalis L. 

Puccinia calendulae, McAlp. 

Calocephalus drummondii Benth. 

Puccinia calocephali, McAlp. 

Calocephalus lacteus Less. 

Puccinia calocephali, McAlp. 

Calotis R. Br. 

Puccinia calotidis, McAlp 

Calotis cuneifolia P. Br. 

Puccinia calotidis, McAlp. 

Caltha introloba F.v.M. 

Aecidium calthae, Grev. 

Candollea serrulata Labill. = Sty- 

lidium graminifoliura, Sw. 

Canthium (Plectronia) coprosmoides 

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. 


Cassia Tourn. 

Uredo pallidula, Cke. and Mass. 

Centaurea cyanus L. 


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 clematidis DC). 

Clematis microphylla DC. 

Caeoma clematidis, Thuem. 

Coprosma billardieri Hook. 

Puccinia coprosmae, Cke. 

Coprosma hirtella Labill. 

Puccinia coprosmae, Cke. 

Correa lawrenciana Hook. 

Puccinia correae, McAlp. 

Crepis japonica Benth. 

Uredo crepidis-japonicae, Lindr. 


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. 

S3 2 

Host Index. 

Dampiera stricta R. Br. 

Puccinia dampierae, Syd. 

Danthonia DC. 

Uromyces danthoniae, McAlp. I. 

Danthonia semiannularis R. Br. 

Uromyces danthoniae, McAlp. II., III. 

Deeringia celosioides KBr. 

Aecidium deeringiae, Cke. and Mass. 

Deyeuxia forsteri Kunth. [Agrostis 

isolandri F. v. M.) 
Puccina agrostidis, Plow. 

Deyeuxia quadriseta Benth. 

Puccinia graminis, Pers. 

Dianthus caryophyllus L. 

Uromyces caryophyllinus (Schrank) 

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 renif olium P. v. M. 

Puccinia gei, McAlp. 

Glyceria dives F. v. M. 

Puccinia graminis, Pers. 

Glyceria stricta Hook. f. 

Puccinia graminis, Pers. 

Gnaphalium japonicum Thunb. 

Puccinia gnaphalii (Speg.) P. Henn. 

Gnaphalium purpureum L. 

Puccinia gnaphalii (Speg.) P. Henn. 

Gompholobium latifolium Sm. 

Cronartium jacksoniae, P. Henn. 

Goodenia albiflora Schlecht. 

Puccinia saccardoi, Ludw. 

Goodenia geniculata R. Br. 

Puccinia saccardoi, Ludw. 

Goodenia glauca F. v. M. 

Puccinia saccardoi, Ludw. 

Goodenia hederacea Sm. 

Puccinia saccardoi, Ludw. 

Goodenia ovata Sm. 

Puccinia saccardoi, Ludw. 

Goodenia pinnatifida Schlecht. 

Puccinia saccardoi, Ludw. 

Goodia lotifolia Salisb. 

Aecidium soleniiforme, Berk. 


Haemodorum Sm. 

Puccinia haemodori, P. Henn 


Host Index. 




Hakea Schrad. 

Uredo angiosperma, Thuem. 

Hardenbergia monophylla Benth. 

(Kennedy a monophyUa Vent. ) 
Uromyces hardenbergiae, McAlp. 

Helianthus annuus L. 

Puccina helianthi, Schw. 

Helianthus tuberosus L. 

Puccinia helianthi, Schw. 

Helichrysum Vaill. 

Puccinia kalchbrenneri, De Toni. 

Hibbertia sericea Benth. 

Puccinia hibbertiae, McAlp. 

Hibiscus L. 

Puccinia heterospora, B. and C. 

Hordeum inurinum L. 

Puccinia graminis, Pers. 

Hordeum secalinum Schreb. 

Puccinia graminis, Pers. 

Hordeum vulgare L. 

Puccinia graminis, Pers. 

P. simplex (Koern.) Eriks. and Henn. 

Hypericum japonicum Thunb. 

Melampsora hypericorum (DC.) 

Aecidium disseminatum, Berk. 

Hypochoeris glabra L. 

Puccinia hypochoeridis, Oud. 

Hypochoeris radicata L. 

Puccinia hypochoeridis, Oud. 

Hypoxis glabella R. Br. 

Puccinia hypoxidis, McAlp. 

Jacksonia scoparia R. Br. 

Cronartium jacksoniae, P. Henn. 

Juncus effusus L. 

Puccinia juncophila, Cke and Mass. 

Juncus maritimus Lam. 

Puccinia juncophila, Cke and Mass. 

Juncus pallidus R. Br. 

Puccinia juncophila, Cke and Mass. 

Juncus pauciflora R. Br. 

Puccinia juncophila, Cke and Mass. 

Kennedya monophylla, Vent. = Har- 
denbergia monophylla, Benth. 

Kochia sedifolia F. v. M. 

Puccinia kochiae, Mass. 

Kochia villosa Lindl. 

Puccinia kochiae, Mass. 

Lactuca L. 

Puccinia prenanthis (Pers.) Lindr. 

Lagenophora billardieri Cass. 

Puccinia lagenophorae, Cke. 

Lagenophora huegelii Benth. 

Puccinia lagenophorae, Cke. 

Lavatera plebeia Sims. 

Puccinia malvacearum, Mont. 

Leschenaultia linarioides DC. 

Puccinia gilgiana, P. Henn. 

Limnanthemum indicum Thw. 

Aecidium nymphoidis, DC. 

Limosella aquatica L. 

Uromyces limosellae, Ludw. 

Linum marginale A. Cunn. 

Melampsora lini (Pers.) Tul. 

Linum usitatissimum L. 

Melampsora lini (Pers.) Tul. 

Lobelia anceps L. 

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

Lobelia pratioides Benth. 

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



Lolium perenne L. 

Puccinia lolii, Niels. 

Loranthus celastroides Sieber. 

Puccinia loranthicola, McAlp. 

Luzula campestris DC. 

Puccinia tenuispora, McAlp. 

Luzula oldfieldii Hook. f. 

Puccinia tenuispora, McAlp. 

Malva rotundifolia L. 

Puccinia malvacearum, Mont. 

Malva sylvestris L. 

Pucciniainalvacearum, Mont. 

Mentha laxiflora Benth. 

Puccinia menthae, Pers. 

Mentha pulegium L. 

Puccinia menthae, Pers. 

Microtis porrifolia R.Br. 

Uromyces microtidis, Cke. 

Micr olaena s tipoides R. Br. ( Ekrharta 

atipoides Labill . ) 
Uromyces ehrhartae, McAlp. 

Muehlenbeckia adpressa Meissn. 

Puccinia muehlenbeckiae (Cke.) Syd. 

Muehlenbeckia cunninghami F.v. M. 

Uromyces politus (B. and Br.) McAlp. 


Host Index. 

Muehlenbeckia gracillima Meissn. 

Puccinia muehlenbeckiae (Cke.) Syd 


Olearia argophylla F. v. M. (Aster). 

Puccinia oleariae, McAlp. 

Olearia axillaris F.v.M. 

Aeciclium oleariae, McAlp. 

Opercularia aspera Gaertn. 

Puccinia operculariae (Morr. ) Syd. 

Opercularia varia Hook f . 

Puccinia operculariae (Morr.) Syd. 

Pelargonium australe Jacq. 

Puccinia morrisoni, McAlp. 

Phalaris canariensis L. 

Puccinia graminis, Pers. 

Phalaris minor Retz. 

Puccinia graminis, Pers. 

Phragmites communis Trin. 

Puccinia magnusiana, Koern. 
P. tepperi, Ludw. 

Plagianthus sidoides Hook. 

Puccinia plagianthi, McAlp. 

Plagianthus spicatus Benth. 

Puccinia malvacearum, Mont. 

Plantago varia R. Br. 

Aecidium plantaginis-variae, McAlp. 

Platylobium formosum Sm. 

Aecidium platylobii, McAlp. 
Cronartium jacksoniae, P. Henn. 

Plectronia coprosmoides = Canthium 

coprosmoides, F. v. M. 

Poa annua L. 

Puccinia poarum, Niels. 

Poa caespitosa Forst. 

Puccinia poarum, Niels. 

Poa pratensis L. 

Puccinia poarum, Niels. 

Podolepis longipedata A. Cunn. 

Puccinia podolepidis, McAlp. 

Polygonum aviculare L. 

Uromyces polygoni, Fckl. 

Pomaderris apetala Labill. 

Uredo spy rid ii, Cke. and Mass. 

Pratia erecta Gaudich. 

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

Pratia pedunculata Benth. 

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

Pratia platycalyx Benth. 

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

Prunus amygdalus Stokes. 

Puccinia pruni, Pers. 

Prunus armeniaca L. 

Puccinia pruni, Pers. 

Prunus domestica L. 

Puccinia pruni, Pers. 

Prunus persica Stokes. 

Puccinia pruni, Pers. 

Ranunculus L. 

Aecidium ranunculacearum, DC. 

Ranunculus gunnianus Hook. 

Aecidium ranunculacearum, DC. 

Ranunculus lappaceus Sm. 

Aecidium ranunculacearum, DC. 

Ranunculus parviflorus L. 

Aecidium ranunculacearum, DC. 

Ranunculus rivularis Banks and Sol 

Aecidium ranunculacearum, DC. 

Rhagodia billardieri R.Br. 

Uredo rhagodiae, Cke. and Mass. 

Rosa canina L. 

Phragmidium subcorticium (Schrank) 

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 tasmanica, Diet. 

Senecio vulgaris L. 

Puccinia tasmanica, Diet. 

Sorghum halepense Pers. 

Puccinia purpurea, Cke. 

Sorghum vulgare Pers. 

Puccinia purpurea, Cke. 

Sporobolus indicus R. Br. 

Uromyces tenuicutis, McAlp. 

Spyridium parvifolium F. v. 

(Cryptandra hookeri F. v. M.) 
Uredo spyridii, Cke and Mass. 


Stellaria media Cyrill. 

Puccinia arenariae (Schum.) Schroet. 

Stipa flavescens Labill. 

Puccinia flavescentis, McAlp. 

Stipa semibarbata R. Br. 

Puccinia flavescentis, McAlp. 

Stylidium graminifolium Sm. (Can- 

dollea serrulata Labill.) 
Puccinia stylidii, McAlp. 

Tabernaemontana orientalis R. Br. 

Caeoma apocyni, McAlp. 

Tetragonia implexicoma Hook. f. 

Puccinia tetragoniae, McAlp. 

Thelymitra antennifera Hook. f. 

Uromyces thelymitrae, McAlp. 

Thelymitra flexuosa Endl. 

Uromyces thelymitrae, McAlp. 

Threlkeldia drupata Diels. 

Puccinia dielsiana, P. Henn. 

Tillaea sieberiana Schult. 

Uredo tillaeae, McAlp. 

Tremandra stelligera R. Br. 

Puccinia pritzeliana, P. Henn. 

Tricoryne elatior R. Br. 

Uromyces tricorynes, McAlp. 

Trifolium repens L. 

Uromyces trifolii (Alb. and Schw.) 

Triticum polonicum L. 

Puccinia graminis, Pers. 
P. triticina, Eriks. 

Triticum vulgare Vill. 

Puccinia graminis, Pers. 
P. triticina, Eriks. 

Urtica dioica L. 

Puccinia caricis (Schum.) Reb. I 
(Aecidium urticae Schum.) 

Velleia macrocalyx De Vriese. 

Puccinia saccardoi, Ludw. 

Velleia paradoxa R. Br. 

Puccinia saccardoi, Ludw. 

Veronica L. 

Aecidium veronicae, Berk. 

Veronica calycina R. Br. 

Aecidium disc\forme, McAlp. 

Veronica gracilis R. Br. 

Aecidium disciforme, McAlp. 

Vicia faba L. 

Uromyces fabae (Pers.) De Bary. 

Vigna catjang Walp. 

Uromyces appendiculatus (Pers.) 

Viola betonicifolia Sm. 

Puccinia hederaceae, McAlp. 


Host Index. 

Viola hederacea LabilL 

Puccinia hederaceae, McAlp. 

Vittadinia australis Rich. 

Puccinia vittadiniae, McAlp. 

Wurmbea dioicaF.v.M. 

Puccinia wurmbeae, Cke. and Mass 
Uredo anguillariae, Cooke. 

Xanthosia pusilla Bunge. 

Puccinia xanthosiae, McAlp. 

Zea mays L. 

Puccinia maydis, Bereng. 

Zornia diphylla Pers. 

Puccinia zorniae (Diet.) McAlp. 

Zygophyllum billardieri DC. 

Uromyces vesiculosus, Wint. 

Zygophyllum glaucescens F.v.M. 

Uromyces vesiculosus, Wint. 

Fungus Index. 




Synonyms in italics ; 


Species excluded from Australia ; t Doubtful species 

Aecidium Pers. 

f • • 


apocyni, Schwein. (See Caeoma apocyni, McAlp., p 
*barbareae, DC. (SeePuccinia cruciferae, McAlp., p. 184) 
*bellidis, Thuem. (See Puccinia distincta, McAlp., p. 156) 

bossiaeae, P. Henn. = Aecidium eburneum, McAlp. 

cakndulae, McAlp. = Puccinia calendulae, McAlp. 

calthae, Grev. — Caltha introloba 

dematidis, DC. = Puccinia agropyri, Ell. & Ev. I. 

• • 

• • 

• • 

• • 

• • 

• • • 

• • 

ccmpositarum, Mart. (See Puccinia tasmanica, Diet., p. 163) 
cymbonoti, Thuem. — Cy mbonotus lawsonianus ... 
cystoveiroides, Berk. = Puccinia operculariae (Morr.) Syd. I. 

• • • 

• • 


leeringiae, Cke. and Mass. — Deeringia celosioides 
disciforme, McAlp. — Veronica calycina, V. gracilis 
disseminatum, Berk. — Hypericum japonicum 
eburneum, McAlp. — Bossiaea cinerea, B. heterophylla, B. linophylla, B. 

microphylla, B. rhombifolia 

• • 

• • 

goodeniacearum, Berk. = Uromyces puccinioides, Berk. & F.v.M. I., and 












Puccinia saccardoi, Ludw. I. 

• • • 

impatientis, Schw. = Puccinia impatientis (Schw. ) Arth. I. 
lobeliae> Thuem. = Puccinia aucta, Berk. & F.v.M. I. 

microstomum, Berk. = Puccinia aucta, F.v.M. I. 
monocystis, Berk. — Abrotanella forsterioides 
nymphoidis, DC. — Limnanthemum indicum 
oleariae, McAlp. — Olearia axillaris 
perkinsiae, P. Henn. = P. gilgiana, P. Henn. I. 

• • 

• • 

- - . 

• • 

• • • 

• • • 

• • • 

• • • 

• • 

• • 

• • • 

• • 

plantaginis, Ces. (See A. plantaginis- variae, McAlp., p. 195) 

• • • 

plantaginis- variae, McAlp. — Plantago varia 
platylobii, McAlp. — Platylobium formosum 
plectroniae, Cke. — Canthium (Plectronia) coprosmoides 
ranunculacearum, DC. — Ranunculus sp., R. gunnianus, 

• ■ ■ 

• • 

- • ■ 

91, 147 

. . 123 

. . 148 

.. 148 

.. 197 

,.. 196 

... 197 

.. 146 

.. 209 

.. 195 

.. 199 

.. 198 

R. lappaceus, R. 

parviflorus, R. rivularis 

- • ■ 

• • 

• • • 

• • 

senecionis, Desm. (See Puccinia tasmanica, Diet., p. 163) 

• • • 

• • • 

• - • 

soleniiforme, Berk. — Goodia lotifolia 
urticae, DC. = Puccinia caricis (Schum.) Reb. I. 
veronicae, Berk. — Veronica sp. 
*violae, Schum. (See Puccinia hederaceae, McAlp., p. 183) 
vittadiniae, McAlp. = Puccinia vittadiniae, McAlp. 

• • 

• ■ • 

• • 

• • 

Caeoma Link 

• • • 

• • • 

• • 

• • 

• • 

apocyni, McAlp. — Tabernaemontana orientalis 
clematidis, Thuem. - Clematis aristata, C. microphylla 

• • • 

• * • 

• • • 

Cronartium Fr. 

• • 

♦ • • 

• • • 

• • • 

p. 190) 

asclepiadeum (Willd.) Fries (See C. jacksoniae, P. Henn., 
jacksoniae, P. Henn.— Jacksonia scoparia, Aotus villosa, Bossiaea cinerea, 

Gompholobium latifolium, Platylobium formosum ... 






Hamaspora Koern. 

longissima, Koern 

Melampsora Cast 

Phragmidium longissimum, Thuem 

- - • 

• • 

• • • 

• • • 

hypericorum(DC.)Schroet.— Hypericum japonicum ... ••• 

lini (Pers.) Tul.-Linum marginale, L. usitatissimum. (Introduced.) 
tnesodaphnes, Berk, and Br. Probably a Hyphomycete ... - .•• 

phylMwrum, Berk, and Br. = Uromyces phyllodiorum (B. and Br.) 

Mc Alp. 






• • 




Fungus Index. 

Phragmidium Link 

• • I 

• • • 

• t 

• t • 

• • 

- - • 

• • • 

• • • 

barnardi, Plow, and Wint. — Rubus parvifolius 

longissimum, Thuem. — Rubus moluccanus 

potentillae (Pers.) Karst. — Acaena ovina, A. sanguisorba ... 

subcorticium (Sclirank) Wint. — Rosa canina, R laxa, R. rubiginosa. 


• • » 

• • • 

• • • 





Puccinia Pers. 

• • • 

• • 

» • • 

• • • 

• • • 

*acetosae (Schum.) Koern. (See P. ludwigii, Tepp. p. 174) 
*aegra, Grove. (See P. hederaceae, McAlp., p. 183) 

agropyri, Ell. and Ev. — Ayropyron scabrum II., III.; Clematis aristatal. 

agrostidis, Plow. — Deyeuxia forsteri ... 

altera, McAlp. = P. cynodontis, Desm. ... 

alyxiae, Cke. and Mass. — Alyxia buxifolia 

angustifoliae, McAlp. — Scorzonera augustifolia 

anthoxanthi, Fckl. — Anthoxanthum odoratum. (Introduced.) 
*apii, Desm. (See Puccinia thuemeni, McAlp., p. 168) ... 

arenariae (Schum.) Schroet. — Stellaria media. (Introduced.) 

aucta, Berk, and F.v.M. — Lobelia anceps, L. pratioides, L. purpurascens, 

• • t 

• • • 

• • • 

» • • 

• • • 

- - . 

• • 

Pratia erecta, P. pedunculata, P. platycalyx 

• • • 

• • • 

• • • 

• • • 

beckmanniae, McAlp. — Beckmannia erucaeformis. (Introduce!.) 

berkekyana, De Toni = P. dichondrae, Mont. ... 

boroniae, P. Henn. — Boronia spines cens 

brachycomes, McAlp. — Brachycome ciliaris, B. diversifolia, B. pachyptera, 

B. scapiformis 

• • 

• • 

• • 

• • • 

• * 

• • 

• • • 

9 • 

• • • 

• • • 

# • • 

• • 

• ■ ■ 

bromina, Eriks. — Bromus arenarius, B. mollis 

brunoniae, McAlp. — Brunonia australis 

burchardiae, Sacc. — Burchardia umbel lata 

cacao, McAlp. — Rottboellia compressa 

calendulae, McAlp. — Calendula officinalis 

calocephali, McAlp. — Calocephalus drummondii, C. lacteus 

colotidis, McAlp. — Calotis sp., C cuneifolia 

caricis (Schum.) Reb. — Carex alsophila, C. breviculmis, C. gunniana, C. 

inversa, C. paniculata, C. pedunculata, II., III. ; Urtica dioica I. 
carissae, Cke. and Mass. — Carissa ovata 
castagnei, Thuem. = P. thuemeni (Thuem.) McAlp. ... . 

*caulincola, Cda. (See P. hypochoeridis, Oud., p. 159.)... 
chrysanthemi, Roze — Chrysanthemum indicum. (Introduced.) 
cichorii (DC.) Bell. — Cichorium intybus. (Introduced.)... 
cinerariae, McAlp. — Cineraria sp. cult, 
coprosmae, Cke. — Coprosma billardieri, C. hirtella 
coprosmatisy Morr. = P. coprosmae, Cke. 
coronifera, Kleb. =P. lolii, Niels. ... ... 

correae, McAlp. — Correa lawrenciana » 

cruciferae, Mc^lp. — Crucifer unknown 

cyani (Schleich.) Pass. — Centaurea cyanus. (Introduced.) 

cynodontis, Desm. — Cynodon dactylon 

cyperi, Arth. — Cyperus rotundus 

dampierae, Syd. — Dampiera alata, D. stricta 

dichondrae, Mont. — Dichondra repens. 

dielsiana. P. Henn. — Threlkeldia drupata, Diels. 

dispersa Eriks. (See P. bromina, Kriks. and P. triticina, Eriks.)... 

distincta, McAlp.— Bellis perennis 

epilobii-tetragoni (DC.) Wint — Epilobium sp., E. billardieranum, E. 

• • • 

• • 

• • • 

• * 

« • • 

• • • 

• • 

1 • • 

• • • 

• - 

• • • 

• ■ 

• • t 









... 144 

... 168 

... 208 

... 153 

... 154 

... 155 

... 165 

... 165 

... 123 

... 181 

... 184 

... 156 

... 118 

... 134 

... 146 

... 142 

... 176 

116, 132 

... 156 


• * ■ 

• • • 

• • • 


erechtitis, McAlp. — Erechtites sp., E. arguta, E. prenanthoides, E. quad- 


• - - 

• • 

• • • 

• * • 

• • • 

• • • 

- ■ - 

• • • 

• • 

• • 

eriostemonis, McAlp. — Eriostemon myoporoides 

festucae, Plow. — Festuca ovina, F. rigida. (Introduced.) 

flavescentis, McAlp. — Stipa flavescens, S. semibarbata 

gei, McAlp. — Geum renifolium 

geranii, Corda — (See P. morrisoni, McAlp., p. 180)... 

geranii-pilosi, McAlp. — Geranium pilosum 

gilgiana, P. Henn. — Leschenaultia linarioides 

gnaphalii, (Speg.) P. Henn. — Gnaphalium japonicum, G. purpureum 

gnaphaliicola, P. Henn. = P. gnaphalii (Speg.) P. Henn. 

. . . 

• • 

• • 

• • • 

• • 



Fungus Index. 



Puccinia Pers. — continued. 


graminis, Pers.— Agropyron divergens, A. scabrum, Alopecurus geniculates, 

Amphibromus neesii, Avena fatua, A. sativa, Beokmannia erucae- 
formis, Briza minor, Bromus racemosus, B. secalinus, B. sterilis, Dactylia 
glomerata, Deyeuxia quadriseta, Echinopogon ovatus, Elymus striatus, 
E. virginicus, Festuca bromoides, Glyceria dives, G. stricta, Hordeum 
murinum, H. secalinum, H. vulgare, Phalaris canariensis, P. minor, 
Secale cereale, Triticum polonicum, T. vulgare. (Introduced.) ... 120 

haemodori, P. Henn. — Haemodorum sp. ... ... ... ... 139 

McAlp. — Viola betonicifolia, V. he deracea ... ... ... 183 

helianthi, Schwein. — Helianth 

(Introduced.) ... 158 

heterospora, Berk, and Curt. — Abutilon avicennae, A. crispum. Hibis 

cus sp 

• • • ••• ••• ••• 

McAlp. — Hibbertia sericea 

• . . 

• - • 

McAlp. — Hypoxis glabella 

• • • • • • ••• - • « • • - 

loranthicola, McAlp. — Loranthus celastroides 
ludwigii, Tepp. — Rumex brownii, R. flexuosus... 

• • • 

• • • . ■ • 

• • 

• • • • • • • 

macalpini, Syd. =P. lagenophorae, Cke. 
magnusiana, Koern. — Phragmites communis 

Mont.— Althaea rosea, Lavatera plebeja, Malva rotundifolia, 

Plagia nthus spicatus . ( Introduced. ) 

• ■ 

••• ••• ••• 

• • • 

• • • • • • 

mussoni, McAlp. — RuelJia australis 

• * • 

McAlp. = P. saccardoi, Ludw. 
McAlp. — Olearia argophylla 

• • • • • 

• • 

• • 

• • • • • 

perplexans, Plow.— Alopecurus geniculatus 

• * • 



hypochoeridis, Oud. — Hypochoeris glabra, H. radicata. (Introduced.) ... 159 
hypochoeridis, McAlp. — (P.macalpini, Syd.) = P. lagenophorae, Cke. ... 161 


impatientis, (Schw.) Arth. — Elymus condensatus. (Introduced.) ... 123 

*investita, Schw. — (See P.gnaphalii (Speg.) P. Henn. ... ... ... 158 

juncophila, Cke. and Mass. — Juncus effusus, J. maritimus, J. pallidus ... 136 

mm art. ... ... ... ... 160 

kalchbrenneri, De Toni — Helichry 

kochiae, Mass. — Enchylaena tomentosa, Kochia sedifolia, K. villosa ... 176 

lagenophorae, Cke. — Lagenophora billardieri, L. huegelii ... ... 161 

lolii, Niels. — Avena fatua, A. sativa, Lolium perenne. (Introduced.) 123 

longispora, McAlp. — Carex sp., C. caespitosa, C. vulgaris ... ... 135 




maydis, Bereng. — Zea mays. (Introduced.) 

menthae, Pers.— Mentha laxiflora, M. pulegium. (Introduced.)... ... 140 

microseris, McAlp. = P. hypochoeridis, Oud. 


morrisoni, McAlp. — Pelargonium australe 

muehlenbeckiae (Cke. ) Syd.— Muehlenbeckia adpressa, M. gracillima ... Uo 

munita, Liidw. = P. dichondrae, Mont — ••• 142 



McAlp. — Asperula oligantha ... ... 

Morr.) Syd.— Opercularia aspera, O. varia . ... ... loo 


phragmitis (Schum.) Koern.— (See P. magnusiana, Koern, p. 1^5) ... **> 

plagianthi, McAlp.— Plagian thus sidoides ... # ... ••• ••• {£? 

poarum, Niels.— Poa annua, P. caespitosa, P. pratensis. (Introduced.) ... j^» 

podolepidis, McAlp.— Podolepis longipedata ... •• — *" i«o 

prenanthis(Pers.)Lindr.— Lactuca sp. (Introduced.) J?f 

pritzeliana, P. Henn.— Tremandra stelligera ... ... «• »• 

pruni, Pers.— Prunus amygdalus, P. armeniaca, P. domestica, P. persica. ^ 

(Introduced.) ... ... ••• ••• ••• , ,*V '*' i 4 >q 

purpurea, Cke. -Sorghum halepense, S. vulgare. (Introduced.) ... i-» 

*rimosa, Link.— (See Uredo scirpi-nodosi, McAlp., p. 203) £° 

*rumicis, Lasch.— (See P. ludwigii, Tepp., p. 174) ... - , - n5 

rwmcis-sciUati var. muehlmbeckiae, Cke. =P. muehlenbeckiae (Cke ) Syd. ... uo 

saccardoi, Ludw. - Goodenia albiflora, G. geniculata, G. g»«ca. u - - 

hederacea, G. ovata,G. pinnatifida, Velleia macrocalyx, \ . pa"£ox» •» J* 

simplex (Koern.) Eriks. and Henn. -Hordeum vulgare. (/ntr^ucwU •» ™ 
sorqhi Schw. = P. maydis, Bereng., and P. purpurea, Cke. (Introduced.) 127, 1£ 



subnitens, Diet.— Distich) is maritima ... ... ••• vpllpinidea' 

tasmanica, Diet. - Senecio brachyglossus, S. pectmatus, S. n elleioides, 


... 163 

S. vulgaris ... ••• ••• ••• . ••' 137 

McAlp.— Luzula campestns, L. oldneidn ... ••• ••• 

tepperi, Ludw.— Phragmites communis... ••• •• 173 

tetragoniae, McAlp.— Tetragonia implexicoma "/r.^ n An^A \ '" 168 

thuemeni, McAlp.-Apium graveolens A. prostratum. (Introduced.) ... ig 

triticina, Eriks.— Triticum vulgare. (Introduced.) ... 


Fungus Index. 

Puccinia Pers. — continued. 

violae, (Schum. ) DC. 


McAlp. — Vittadinia australis 
Ike. and Mass. — Wurmbea dio] 
McAlp. — Xanthosia pusilla 

McAlp., p. 183) 

• • • 

• • 

• • • 


Zornia diphylla 

■ • - 

• • 4 

» • • 


• • • 

t • * 

» • • 

* • 

• • 4) 

• • • 

• t • 

angiosperma, Thuem. — Hakea sp. 
anguillariae, Cke. — Wurmbea dioica 
armillata, Ludw. 

bidentis, P. Henn. — Bidens pilosa * 
bossiaeae, McAlp. — Bossiaea prostrata 
*cichoracearum, DC. 
clematidis, Berk. = Caeoma clematidis, Thuem. 
crepidis-japonicae, Lindr. — Crepis japonica 

• • 

• • • 


• • • 

« • • 

• - . 



• • • 

• - • 

• • 

chrhartae, McAlp. 


gnaphalii Speg. 

McAlp — Geitonoplesium cymosum 

- - • 

• • • 

• t 

- t 

(Speg. ) 

• • 

kuehnii, Krueg. — Saccharum officinarum. (Introduced.) 

leguminum, Desm. — Acacia sp. 

• • • 

• • 

notabilis, Ludw. 


Cassia sp. 


• • 

. - • 

• • t 

• . 

• • 

rhagodiae, Cke. and Mass. — Rhagodia billardieri 
rottboelliae, Diet. = Puccinia cacao, McAlp. 
schelhammerae, McAlp. — Schelhammera undulata 
scirpi-nodosi, McAlp. — Scirpus nodosus 
sorghi, Pckl. = Puccinia purpurea, Cooke, 
spyridii, Cke. and Mass. .....__ 

tillaeae, McAlp. — Tillaea sieberiana 

♦ • * 

• • • 

• • • 

• • • 

• ^ 

• • • 

• • • 

• • 

• • • 


• • 

• • • 

Uromyces Linl 

• • • 

• • • 

• • • 

• • 

• • 

• • • 

• • 

• • • 

• • » 

• • • 

• • • 

amygdali, Pass. (See Puccinia pruni, Pers., p. 171) 
appendiculatus, (Pers.) Link — Vigna catjang. (Introduced.) 
asperulae, McAlp — Asperula oligantha 

asteris, McAlp. = Uredo dubia, McAlp 

atriplicis, McAlp. — A triplex semibaccatum 

betae (Pers.) Kuehn. — Beta vulgaris. (Introduced.) 

bicinctus, McAlp. — Acacia fasciculifera 

bulbinis, Thuem. — Bulbine bulbosa 

caryophyllinus (Schrank) Schroet. — Dianthus caryophyllus, D chinensis 

• • 

• • • 

• • • 

• • 

* • 

• • 


• • 

• * • 

• • 

• • 

danthoniae, tylcAlp. — Danthonia sp., D. semiannularis ... 
digit at us, W int. = Uromyces phyllodiorum (B. and Br.) McAlp. 
diploglottidis, Cke. and Mass. — Diploglottis cunninghamii 
ehrhartae, McAlp — Microlaena (Ehrharta) stipoides 

• • 

• • 

• ■ 

• • 

• • 

» • * 

• • • 

• • 

* • • 

• • « 

• 9 

fabae (Pers.) De Bary. — Vicia faba. (Introduced.) 

fusisporus, Cke. a&d Mass. — Acacia neriifolia, A. salicina 

hardenbergiae, McAlp — Hardenbergia [Kennedya) monophylla 

junci (Desm.) Tul. — (See Puccinia juncophila, Cke. and Mass. p. 136.) 

kuehnii, Krueg. =Uredo kuehnii, Krueg. 

limosellae, Ludw. — Limosella aquatica... 

microtidis, Cke — Microtis porrifolia 

orchidearum, Cke. and Mass — Chiloglottis diphylla, C. gunnii 

phaseoli (Pers. ) Wint. = Uromyces appeadiculatus (Pers.) Link. ... 

phyllodiae, Cke. and Mass. = Uromyces phyllodiorum (B. and Br.) McAlp. 

phyllodiorum (B. and Br-) McAlp. — Acacia sp., A. dallachiana, A. micro- 

botrya, A. neriifolia, A notabilis, A. penninervis, A. pruinosa 
politus (Berk) McAlp. — Muehlenbeckia cunninghami 
polycnemi, McAlp. — Polycnemum pentandrum 

• • • 

• f 

polygoni (Pers.) Fckl. — Polygonum aviculare. (Introduced.) 
puccinioides, Berk, and P. v. M. — Selliera radicans, Scaevola sp. 
scleranthi, Rostr. — Scleranthus diander ... ... 

tenuicutis, McAlp. — Sporobolus indicus 

tepperianus, Sacc. = Uromycladium tepperianum (Sacc. ) McAlp. 

thelymitrae, McAlp — Thelymitra antennifera, T. flexuosa 

• • 

• • 

• • 

• • • 

• • 

• • • 

• • « 

tricorynes, McAlp. — Tricoryne elatior ... 

trifolii (Alb. and Schw.) Wint. — Trifolium repens. (Introduced.) 

vesiculosus, Wint. — Zygophyllum billardieri, Z. glaucescens 

• • • 

< • • 

• • 



























Fungus Index. 


Uromycladium McAlp. 

■ - • • - ' • • 

• - • ••• •• 



alpinum, McAlp. — Acacia buxifolia, A, dalluchiana, A. dealbata, A. implexa, 

A. linifolia 
bisporum, McAlp. — Acacia dealbata 
maritimum, McAlp. — Acacia longifolia 
notabile (Ludw.) McAlp. — Acacia binervaba, A. dealbata, A. decurrens, A. 

9 w » ••• ••• ••• • • • 

• • • • . • • • • • ■ . ••• 

• • • • • • • ■ • • • • 

elata, A. notabilis, A. pruinosa 

• •• ••• . . . 

robinsoni, McAlp. — Acacia melanoxylon 

simplex, McAlp. —Acacia pycnantha 

tepperianum (Sacc.) McAlp. — Acacia armata A. diffusa, A. erioclada, A 

glaucoptera, A. hakeoides, A. implexa, A. juniperina, A. longifolia, A. 

melanoxylon, A. myrtifolia, A. pycnantha, A. rigens, A. salicina, A. 

siculiformis, A. spinescens, A. siricta, A. verniciflua, A. verticiliata, 





A. vomer if or mis . . . 

• • 







General Index. 



• • 

• • i 

Acacia, gall fungi 

Acacia armata 




Aecidiospores, binucleate 

• • • 

a • 

• • 

• « • 

• • ■ 

• • 

• • I 

- • » 

• • • 

• I • 

coloured membrane in Gymno- 

colouring matter 
distinction from uredospores 
germination in Puccinia 






• • • 

• • • 



partaking of 

repeated formation... 

• - 

• • • 




... 17, 36 

character of 



• • 


power . . . 
Aecid ium, colour 

invigorating power 

in heteroecious species 
on grass ... 

on seeds ... 

t • • 

• • i 

• * • 

paraphy ses 
peridiurn ... 
primary and secondary 
pseudoperidial cells 
repeated formation 
sterile cells 
A ecidium abietinum 

* * ■ ■ ■ 


berberidis, spore germination 

platylobii, mycelium in seed 


Alopecurus pratensis with Puc- 






... 17, 75 
22, 36 





14, 17, 75 







cinia gramims 

- • * 

• • • 

• • • 


• • 

• • • 

• • t 

Anagallis arvensis, importation 

• • • - - ■ 


Apium, graveolens (Puccinia) 

proHtratum (Puccinia) 
Asclepia'leae, Cronartium 
Ascomycete8 ... 

Asparagus rust (Puccinia axpar- 

agi), distributed by wind 

53, 80 


water relation 

• * 

• • 



• • • 

• • 

Atmospheric influences on spore 

A uric xdariaceae 
Australian rusts, number 





Autoecious species 

most numerous 

repetition of aecidia ... 

A vena elatior with Puccinio 


Baeodromus ... 

• • • •• • 

Barberry, connexion with wheat 


), 55 




• • t 

• • 

ft • ft 

• ft ft 

in Australia 

infection experiments, local 

in Kew gardens 
Barberry rust (A ecidium ber- 
beridis) ... 

absent in Australia 

connexion with wheat rust ... 

increasing vigor of wheat rust 

local attempts at inoculation 
Barclay e lla 

Basidia in aecidia 


55, 58 



• • • 

— * 

formis, impor 

• • • 

• • 


• • 

• • 

tion of rust on seed 
Beet rust ( Uromyces betae) 
Binucleate spores 

Biologic forms, and classification 

evolution of 
Bismarck brown 
Black wattle gall fungus (Uro- 

mydadium notabile) 
Br achy puccinia 
Bremia lactuca % effect of copper 

Sell Lo ♦ • • ••• . . , 

Bridging species 

Brome rust (Puccinia bromina) ... 
infection by uredospores 
starvation of host ... 
uninfluenced by structural 
peculiarities of leaf 

Brown Rust (Puccinia dispersa) ... 
Burning stubble, effect on wheat 











ft • 

■ • ft 

t • • 

• € • 

• • • 


Caeomospores ... 

Carnation rust (Uromyces caryo- 




Caustic potash for softening tissues 

Cecidomyia, eating rust spores ... 

Cedar apples ... 

Celery rust (Puccinia thwmeni) ... 

Cereal rusts not liable to affect 

other cereals indiscrimin- 
ately ... ••• ••• 

Chemotaxis, parasitism due to 

positive and negative 

Chick weed rust (Puccinia aren- 

Chicory rust (Puccinia cichorii) . . 

Chrysanthemum rust (Puccinia 

germination of uredospores 











I I 



General Index. 

Chrysomyxa, absence of fir trees 
germination of teleutospore 
origin of species in Alps 

» • • 

• i • 

• • 

uredospores in chains 
Chrysomyxa, abietis 

ICCuw • • « •• • 


Chrysopsora, teleutospore of ex- 
ceptional form ... 

Classification and biologic forms 

of Uredines 




• • • 


trifolii) ... 18, 84 



Cluster cups ... 

Coleosporium, function of paraphyses 30 



teleutospore in 
uredospores in chains 

Coleosporium senexionis, and fir trees 58 

Collema, fertilisation in ... 37 




• • • 

Compositae, native rusts on 
Compound teleutospores 
Copper salts, influence on fungi... 
Cronartiaceae ... 
Cronartium>asclepiadeum on varied 



• • 


jacksoniae producing witches' 
brooms ... 

Crossbreeding wheats for rust- 

Crown or coronate rusts 

Cucumber diseases and copper salts 

Cycle of development, in heteroe- 



cious species 
in rusts 

• • 

• • 

Cuperaceae, number of native rusts 

, 58 
2, 10 


• • • 

Dactylis glo merata 

Daisy rust (Puccinia distincta) ... 

introduced on seed ... 
Danthonia aec 
Darluca Mum 



... 18, 57 
parasite 22 
mistaken for spermogonia 22, 158, 165 


Development of fungi ... 
Dew necessary for infection 
Diorchidium ... 

doubtful genus 

• • 

• • • 

• • 

* • • 

Distribution of Australian species 
Distribution of spores 
Doubtful species 

• • • 

• # • 

Drainage, effect on rust 

Dry atmosphere and spore ger- 

Early maturing wheat escaping rust 

Echinulate uredospores... 

Effect of rust on straw and grain 

Elymus condensatus, importation 

of rust on seed ... 

Endochrome ... 

Endophyllum, teleutospores 

Endospore, in aecidiospores 
in teleutospores 

Enzymes in leaf cells ... 

Epispore of teleutospore 

Epiteospores .... 

Erysiphaceae, specialisation 









# • 

• • • 

• • « 





• • • 


• • • 




• • 1 

• • • 

Euphorbia, aecidium of Pea rust 

Gu&fvLCLLCL ••• ••• ••• 

rust ( Uromyces euphorbiae) . . . 
Excluded species 
Fallowing, effect on rust 
Fertilisation, in aecidium 

in Uredineae ... 

Fir trees and rust 
Flax rust (Melampsora lini) 

first record in Australia 
Formalin treatment of seed wheat 
Frangula alnus 

Fungi, origin and development . . . 
Fusion of nuclei 
Gall formation... 





... 17, 75 
... 14, 75 


% • • 

... 43, 60 



... 14, 75 

• - * 

Gall fungi, on wattles (Acacias) ..• 
Genera, Australian, and number of 

OlJL/V-/lt>0 »•• ••• ••• 

General parasites 

Germinating power, duration, in 



in teleutospores 
in uredospores 

• • • 

• • • 

• • 

• * • 


8, 21 

Germinating spores, methods for 

aecidiospores and uredo- 

bUOI vS ••• ••• ••• 

Germination, of aecidiospores of 



• • • 

Puccinia tasmanica 
of amphispores 
of teleutospores, factors in- 


of Puccinia graminis 

of P. malvacearum 


8, 24, 66 
... 24, 27 

of Phragmidium rubi abnor- 


• • ♦ 


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 

• • • 

. • • 

• • 

• • # 

• • 

■ ■ ■ 

3, 8 




solitary in Puccinia mono- 


• • • 

Glycerine, and water ... 

I Li i y ••• ••• • • • 

Golden rust (Puccinia glumarum) 

Golden wattle galls 

Gradations of specific variation ... 

Gramineae, native rusts on 

Grass aecidium 

Grasses, and heteroecious rusts ... 


and wheat rust 

• • • 

• •• lo, O/. 


Groundsel rust (Puccinia tas- 


• • 

Gymnoconia, characters... 
Gymnosporangium, aecidiospores 

colored membrane 

cTclllS . . • ••• ••• 

insects conveying sporidiola 
sculpturing of peri dial cells ... 

... 16, 49 





General Index. 




mosporangium clavariaeforme 9 

direct infection by teleuto- 

spores ... 
confusum, promycelial 


teleutospores, thickness of 




cell wall 

• • 

• • • 

« • 

Hap a 

Hard tissues, softening... 

Harvesting methods and rust 



Helianthus anmtus, aecidi 

// em ibasidii ... 
Hemipuccinia ... 
Heteroecious species 


regular development 

• - » 

• # 

• • • 

• • • 

• - * 

• • • 

• - • 

• • • 

• ■ • 

• • • 

• • 

. . . 



beneficial ... 

causing increased vigor 

discovery ... 


when possible 

• • • 

• • 

♦ • • 

• * • 

- - - 











arum ) 

• • • 

wide distribution 

- - • 

• • 

• • • 

• • « 

• ■ • 


Honeydew of spermogonia 
Host and parasite, relatior 
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- 





62, 75 


• • 

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 ... 

15y sporidiola 

germ tubes entering stomata 

by teleutospores 
by uredospores 
germ tubes piercing epider- 












9, 60 







mal cells 

• • I 

dew necessary for ... 
experiments, with barberry in 



• • • 

with foreign host plants 

factors influencing 

• t • 


Injuries, influence on infection 
Inoculation, protective... 

• ■ • 


Insects attracted 

colour of 


• • ■ 

• • • 

♦ • • 

by honeydew 

by scent of spermogonia 

conveying sporidiola 

eating spores 
Intermediate hosts of wheat 
Internal developmental tendencies 
Introduced species 


• • . 

• • # 

• • • 

how introduced 

on native hosts 
Investigation of spores ... 
Irrigation and wheat rust 

Kangaroo thorn, gall fungus 

K not- wet d rust ( Uromyces polygoni) 

Lactic acid showing up germ pores 
Ledum palustre 
Leguminosae, native rusts on 
Leptopuccinia ... 

species with war ted 
Lettuce, attempt to confer immu- 

• • • 

• • • 

• • • 

















Liability to disease 

influenced or not by structural 

Lichen-fungi ... 

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, rus 
Maize rust (Puccinia maydis) 
Mallow rust (Puccinia malva* 
Manures and wheat rust 

M ■ 

• • • 

• • • 

• - • 













Marigold rust {Puccinia calendulae) 10, 42 
Measuring spores by photography 12 
Melampsora, germination of teleu- 

tospore ... ••• ••• 

Melampsora eucalypti, merely a 



• • • 


• • 

• • • 


M elampsoraceae^ 


• • • 

■ - • 


in Uromycladium 



22, 49 


25, 26 


Mint, native, rust (Pucctma 

Mounting spores 

Mycelium, development of 
in Refill of annuals 

. • - 

• • • 

localized, causi w 
perennial causing galls 


• • • 

no evidence in wheat grain 
of Puccinia arrhenatheri .•• 

of Uromyces euphortriae 

t ♦ t 






General Index. 

Mycoplasm theory 

• • • 

• ft 

• • • 

• . • 

• • • 

bearing on rust in wheat 
Native celery ... 



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 

in uredosori 

- ■ . 

• • 

• • • 

• • 

• • 

• • # 

• • 

• * 

• • 

• ♦ ' 

• • • 

• • • 


4, 74 



42, 44 






14, 75 






• • 




13, 29 

• • • 

Australian species with 

• • # 

Parasitism, origin 

Pea rust ( Uromyces pisi) 
Peach or prune rust {Puccinia 

22, 24, 75 



pruni ) 

• • • 

• • 

Pedicel of teleutospore ... 
Peltandra virginica 
Perennial mycelium 
Periderm ium ... 
Peridial cells ... 

• • • 

* • • 

Peridium, =Pseudoperidium 

often wanting 

varied forms 
Phalaris, Puccinia 
Photography in measuring spores 
Phragmidium bamardi, germ pores 




rubi 9 teleutospore germination 

ft • 4 

paraphyses, in aecidia 

in uredosori 

• . • 

perennial mycelium 

... 19, 44 


• • • o 







5, 36 



* • 

• • 

• * • 

• • g 

Phragmites communis, Puccinia ... 29, 42 
Phragmopyxis ... ... 84 

Phy corny cetes^ ... ... ... 40 

Phytopthora infestans, and copper 

ScHuS ••• mm* 

Picea excelsa ... 


Ploughing, influence on wheat rust 

Plum or prune rust (Puccinia 

pruni) ... 

Poa annua 


pratensis . . . 

Polygonum aviculare 
Potash, caustic, 


• • • 



for softening 

r • • 


• • * 

• • • 

• • • 

Promycelial spores (see Spori- 






• . . 

• • 

cells of, separating 


in various genera 

• • 

• - . 

• • • 

• • 

• • 

• * 

• • 

• • 

• « 

Protective inoculation 




Pseudoperidial cells (see Peridial cells) 

Pseudoperidium (see Peridium) 

Puccinia, introduced species 

variation in teleutospores ... 
Puccinia agropyri, heteroecious 







agrostidis ... 
arenariae, introduced 

* • • 

* • • 

• • 

• • • 

• • * 

• • 







sporidiola germ-tubes enter- 
ing by stomata 

arrhenatheri, perennial myce- 


• • • 





asparagi, influence of wind ... 

water relation ... 
beckmanniae, imported 
bromina contrasted with P. 

triticina ... 80 

paraphyses in teleutosori ...24, 75 
uredospore, duration of 

3, 8 

• ft * 

germinating power 
germination and infection 
calendidae, marigold rust 
Australian species 
short cycle 

caricis, heteroecious 

chrytanthemi, germination of 

two-celled uredospores 
cichorii, introduced 
cinerariae . . . 
coronata . . . 

• • • 

« * • 

♦ • • 

C ft ft 

cyani, introduced ... 
dianthi, sporidiola germ-tubes 


entering stomat a 

diihondrae , 



• • I 

ft • • 


• • • 


dispersa, biologic forms 


• • 



distincta, daisy rust, Aus- 

tralian species 
introduced on seed 
short cycle 

♦ • • 

• • 

glumarum, mycoplasm theory 
graminella> with aecidium on 

££^ciSS ... ... • ■ . 

graminis, biologic forms 

carried over from year to 







in Australia 
collective species 
cycle of development 
forms on special hosts 

• • • 

• * # 

21, 69 





General Index. 



Puccinia graminw, importation on 


• • • 

• • 

injurious rust in Australia 
no aecidia in Australia ... 
on grasses, wheat, &c. ..• 
repeated formation of 

reproduced by uredo - 





spores in Australia 

• • * 

specialisation of forms ... 
teleutospores,from Australia 
not germinating in Eng- 
land, and vice virsd ... 
germinating power 
germination, exposure to 
cold not necessary ... 





in Australia... 
in water 
time for 

• • • 

... 24, 67 


. . . o 


uredospores, germinating 

on old straw 
germination ... 
persist through winter . . . 
graminis and barberry rust, 
connexion first discovered 



in Australia 
in Kew Gardens ... 
helianthi, germination 


• • ■ 

... 58, 66 


• • • t/ 

heterospora, one celled teleuto- 

spores most common 

* • • 

transition form 

• • 



hieracii, spermogonia with 
subdivision of 

• • • 

impatientis, introduced 
liliacearurti) spermogonia with 

lolii, introduced 



paraphyses in uredosori ... 
magnusiana, heteroecious ... 
paraphyses, in teleutosori 
and uredosori 
malvacearwn, causing "shot- 

... 13, 37 







germination of teleutospores 24, 27 
importation and widespread 



suppression of spermogonia 

maydis, introduced... 
menthae, introduced 
obtegens, spermogonia with 




paraphyses in uredosori ... 22, 29 
uredo withstanding cold ... 22 

perplexans, on native host ... 
paraphyses in uredosori ... 

plagianthiy epispore 
poarum, introduced 

podolepidis, two germ pores in 

upper cell of teleutospores 

prainiana, abnormal teleuto- 
spore germination 
prenanlhis, introduced 


peridium often wanting ... 



cinia prunt, causing "shot-hole 
first record in Australia ... 
producing uredospores 

within fruits ... 
supposed spermogonia 
purpurea f introduced 
rubigovera, germinating power 
of uredospores 
paraphyses in teleutosori .. 





• • • 

senecionisy repetition of aecidia 

simplex, aecidia unknown ... 


suaveoleris, scent 

• • • 





tasmanica, germination oi 

• • * 


... 42, 49 

groundsel rust 
thuemeniy introduced 

attacking native host .. 42 
triticina, aecidia unknown 


comparatively harmless ... 64, 66 

contrasted with P. bromina 




■— ' — — ----- »— - •— - ■ -w ^ -*^ «^— — v v ~ w w« w v «-r -v a. m www 

preference for certain parts 
vexansy amphispores 


• • • 

Queen's Jubilee wheat, rus 

on grain ... 
fianuculaceae, Cronartium 

• • • 



• t 



infection ex- 


compared with Uromycladium 

Repetition, of aecidia ... 

of spore forms 

" spermogonia... 

of uredospores 

• • 








Rerraf, a rust-resisting wheat ... 64, 74 


• • • t)*9 



at hart tea 

Rhododendron, Chrysomyxa 


Rose rust (Phragmidium subcorti- 


• • 

• • 

Rotation of crops and rust 
Rubiaceae, native rusts... 
Rust Conference 
Rust of wheat ... 

• • 


* • 




connexion with rust on grasses 
early theories 
effect of, drainage ... 

early maturing wheats ... 

* • ■ 




muggy weather 


rotation of crops 


seed treatment 

stubble burning 
effect on straw and grain 
first appearance in season 
first record in Australia 





... 67| 72 








• • • 

• • • 

• • • 


General Index. 



Rust of wheat, how carried over 

from year to year 

• # 

• • • 

• • 

how spread 

infection from within, myco- 

plasm theory 
influence on yield ... 
losses from 
mycelium not in seed 
mycoplasm theory 
not spread from other cereals 
prevention and mitigation ... 

question, present position in 

spores, in soil 

21, 68 


4, 70 


on seed 

to destroy 

• * 

• • 

• • • 

• • • 

• • 

• • 

spraying impracticable 

• • 

• • % 

Rust-liable soils 

Rust -proof wheat unknown 

Rust-resistance and structural 

• • 

• • 


8, 20 




♦ • • 

• • 

Rust-resisting wheats ... 

Rusts, Australian, and their hosts 
cycle of development 

in heteroecious species 
indigenous and introduced ... 
influence of parasitic habit ... 
relation to other fungi 



2, 10 

14, 17, 75 

Rusty straw and grain analyzed 
Rye, rust 

• • 

• ■ • 

• • 

• • 

Saccoblastia ovispora 
Saprophytes becoming parasitic 
Scarlet pimpernel 
Scent of spermogonia ... 
Scrophulariaceae, Cronartium . . . 

kD tSC/Cvt'O ••• ••• ••• 

Seed, perennial mycelium 

not in wheat 

Seed-bed and rust 

Seed-treatment for rust 

Seeds with aecidia 

Selection and crossbreeding wheat 

Self-sown wheat and rust 
Senecio vulgaris 

21, 69, 






• • • 

- ■ . 

formation in promycelia 
Sexuality and nuclei 

of aecidia 

• • 



... 14, 75 
... 15, 17 

Shot-hole caused, by Puccinia mal- 

by P. pruni 

Silver wattle gall fungus ( Uromy- 

cladium notabile) ... 

Softening tissues for microscopical 

Soil moisture, influence on rust ... 

Soils, rust-liable and rust- free ... 
treatment with copper salts 






to prevent disease 

Sorus, uredo ... 

Special corpuscles 

Specialisation of parasitism 
in Erysiphaceae 

9 • • 

• • • 

• • • 





Specific variation , gradations 
Spermatia, embedded in sugary 


formation ... 
not male cells 

■ • • 

• • 


• . . 1 , ' o 
. . . 1 «"> , . > / 



Spermogonia, associated with cer- 

tain spore forms ... 
attractive to insects 
Darluca filura mistaken for... 

... 13, 37 

... ±0) oo 


• • • 


• • • 

• • • 


repetition of spore forms 

* • 

Spore -bed 


origin of principal 

Spores, distribution 

• • 

V • 

* • 

• * 

13, 37, 39 





11, 18, 19 


• • 

• • • 

• • 

• • • 

germination, of aecidiospores 
of amphispores 
of spermatia 
of sporidiola 
of teleutospores 

of uredospores 

• • 

• • • 

• • • 

« • • 

• • • 

7, 16 



*.. 13, 3/ 


8, 24, 27, 66 


on seed 
parasites ... 

wintering, of teleutospores 
of uredospores 

7, 19, 21, 27 


• • 

• • • 

• • • 

• • • 


8, 20 

Sporidiola, air necessary for for- 

• • • 

conveyed by insects 
earliest spore form ... 
formation ... 
germination ... 

incapable of infecting grasses 

bearing them 
infection ... 

• • • 

• • • 






• • • 


• • • 





Spraying for rust of wheat im- 
Staining spores 


Starvation of host and infective 

power of rust 

• • • 

Stellar ia media 

Structural characters of host and 

susceptibility, to disease ... 

to wheat rust 
Stubble burning and rust 
Sugar, influence on germ tubes of 






• • • 

• • • 

... 52, 62 

Sulphate of copper, in seed treat- 

ment for rust 

• • 

• • 

treatment of soil with 
Suppression of spore forms 
Surface markings of spores 

• • t 

• • « 





General Index. 




Susceptibility of host, due to 

physiological peculiarities 
influence of structural charac- 



Teleutospores ... 

• • • 

• • • 

• • 

direct infection by germ tube 

function ... 

germination, factors influenc- 

61, 62 



in Phragmidiumrubi 

in Puccinia graminis 
in P. malvacearum 

• * 

• • • 

origin and development 


- • - 

• « • 

• • 

simple or compound 

two kinds... 
variability, of form 

of germination 

• ■ • 

Teleutsori paraphysate... 

Tomato, copper salts and disease 


Trifolium repens ... 

Triphragmium ulmariae 


8, 24, 66 

.. 24, 27 









33, 35 




• • ♦ 

• • t 

primary and secondary uredo 


4 * 

• • • 

• • • 

• • • 

spermogonia with uredo 

Uninucleate spores 

Uredineae in relation to other 



Uredo dispersa, histology 

kuehnii, paraphyses 

spyridii, paraphyses 

20, 36 


# •-• 

• • • 

• • 


• • 

Uredosori, paraphyses ... 

parasites ... 
Uredospores, binucleate 

derived from teleutospores . . . 



function ... 
germinating power 

duration of 
germination, in Puccinia chry- 

sait themi 
in P. rubigovera ... 
germ tubes, piercing cells ... 
infection by germ tubes 


primary and secondary 

produced within fruit in Puc- 




22, 29 

22, 29 












cinia pruni 

• ft • 

repeated formation 

two celled, in Puccinia chry- 


wintering ... 



• • 

20, 21 
24, 83 

distinction between uredo- 
spores and teleutospores . . 





Uromyces dactylidis, paraphyses 
danthoniae. with aecidium ( 



... 18, 57 


euphorbiae, perennial mycelium 5, 70 



fabae in Ecuador 
limosellae ... 
orchidearum, two-celled teleu- 
tospores ... ... 39 

phyllodiorum , paraphyses ... 24, 29 

pisi ... ... ... 84 

politus, two-celled teleuto- 

spores ... 
polygoni ... 


• • • 

... 43, 55 

proeminens transition forms 
from teleutospore to uredo- 

scutellatus, origin of uredo- 




• ■ 

uredo with teleuto spore ... 

solid aginis 

tricoryiies y two-celled teleuto- 
spores ... 

trifolii — complete cycle 
perennial mycelium 



• • • 

. . . 39, 83 




idosus. two-celled teleu- 

Uromycladium . . . 

• 9 • 

forming galls often 





• • • 

• • • 

... 3*7, Ot> 

24, 83, 104 


... 25, 2li 


Uromycladium maritimum, occur- 
rence of spermogonia 


perennial mycelium and 

galls ... 

... 13, 37 


• • • 


robiwoni, spermogonia 
tapper ianum, spermogonia ... 13, 37 
perennial mycelium and 

galls ... 

• • • 

• • 

• • • 

• ■ • 



Vegetative organs, mycelium 
Verrucose uredospores . . . 
Vesicle in Uromycladium 
Violet rust, native 

Water relation, for Puccinia on 









• • • 

asparagus ... 

to infection, direct and m- 



• • • 


Wattles, gall- fungi on ... • • • 

Wheat rust. See Rust of Wheat. 

• • • D^j # ^ 

Wheat, rust -resisting ...' 
self-sown and rust ... 
Wild oats and wheat rust 
Wind distribution of spores 
Wintering of rust spores 
Witches' brooms 

21, 69, 72 


... 8,20 


Brain. Government Printer, Melbou 











A.' J> 



















, I*** 




I ■* 


t rl 






* . 



JAN 2 3 1963 







' V 








f U 

h ^*» - 


- ^^h^^^^jj»W& ^< 

lr> - X 






• < 











i * 















LD 21 3m-8,'3° 




fr? ^ 


s . 



• • 

• , 


- ri> 












' 5fc> 







. »> 




F >- 




• fc 

/ - 






. ,& 




1 i 



«• » 


■ * 



: ---':-. 

, — 

* • *v 

V V ^B 

i * 

1^ t ' ■ - - 

• < « ~ « - * 

• •-•.#•-'•4. . « i k * . . \ 1^. 114 t # 1 ? , 1 91 1 • 1 

... . ■ 

' , ■ : 

!«4> ; " J • 

; :. .... ; r 1 _ ; . - .! ; : r 


P ; "-it -OBI;; 


Bt2 *; ;**£*}'•{ §t;; 'iliffJrl ,::. 

i: ? |M |j{f iSiii jmiiiflAMfe* 

4 • 

- j 

- I 

< ' 


• . f 

«• » 

I * 1 • ■ • * ? ♦ % 4f 14**' 'iflUC 

tifUf —***ttfr vV 

f 1 TYlf .««.»• t^»f>i| -i. j *4*f 

-,;>;:,; •;:.;„. — • 

i :■•::■■ ■, ; I 

• 1 ■ • - 4 ... 

- *>— • • • •* 4 

r « 

• :?»i:Mi ij^iiitiSKiti 



* * • til 

• 1*M 1 t K *■ "• • » 1 ,•»*•-* » % * 



■ » 

Pl^Hlii^^pj^HpHpfp^H - * - - 

-* « t 

^ « 


*1 T 

* * * 

» — 

4 v 

• * » « 

* fk 

ID; Hi- »J- .:i - IHilll. ilxr, tJuHi-. «•;;« 

- t 


■ ■ 

- * -> 



3: r 


« *• * 



■ V 


- ■Bl?9>4f •" "4%™- * 

*. •• .--. . . m * . • 

* *•-— 

^ * f 


. N , , 



* 4 

•~ — » ■ 

• • 

A I 

* 7 -- 

• t 

41 * *■ 

A • <■ 

■ * * 

7 .: : i 




*t t 

• * 1 

t# • 

4 • •.-" « 


* * 


■• I * 


m m v 

1 I 


* - 4 

-*l *— * 




1 . 





i ' • . 

tt ■ 





ii fl 

'HpJHWtt^n RfflUIBffl 

m\V •::ft|Hif!- '-•'{'; i : 


in iffl n 



* * a X ^ w iVl r 4 J I l&l iff 1 tMlllf llll'ijtlf 15 12t IU 

' • ill il n 1 III 1 it il ! Oiliiiifttfl 


■ • 


' • rlf r ll#I ill 24 it !tI§j llnlii^pitp^Bi^HI i 


if j 

! I f 


i : 

- • 


• * 


. . 

_, • 






. ' • 








I li' 


, * # 


rt ■ •'" # »r*