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THE STUDY OF FUNGI
INTEODUCTION
TO
THE STUDY OF FUNGI
THEIR ORGANOGRAPHY, CLASSIFICATION, AND
DISTRIBUTION
FOR THE USE OF COLLECTORS
M. C. COOKE, M.A., LL.D., A.L.S.
adthor of
"fungi: their nature, uses, etc."; "handbook of British fungi";
illustrations of british fungi " ; "microscopic fungi " ; "british edii
fungi": "handbook of Australian fungi," etc. etc.
LONDON
ADAM AND CHARLES BLACK
1895
PREFACE
The Introduction to Cryptogamic Botany, puljlishcd by
Berkeley in 1857, was for a long time the only volume, in
English, which could introduce the inquiring student to a
systematic knowledge of Fungi. Later on, this work was
discovered to be insufficient, inasmuch as it was more suited
to the requirements of an advanced student than an inquirer ;
so that the field was left open for a more popular and ele-
mentary work, which, under the title of Fungi : their Nature,
Influence, and Uses, appeared in 1875, subsequently passing
through several editions. The rapid advance in knowledge of
the life-history and development of these organisms during the
past ten years, and especially the large scheme of classification
carried out by Professor Saccardo, made it essential that, in
order to keep pace with the times, a guide and introduction
should be prepared and pul)lished for the use of students,
which, whilst not superseding the volume of 187 5 as a popular
instructor, should treat the subject more after the manner of
a text-book, adapted to the illustration of recent discoveries,
and an explanation of the methods of classification. The
following pages are the result of an effort to supply an acknow-
ledged want, which I have executed under the impression that
it is probably my last contribution of any importance to
British ]\lycology.
For many of the illustrations the publishers and myself
duly acknowledge the kindness with which they have been
placed at our disposal by the publishers of the works from
77736
vi INTRODUCTION TO THE STUDY OF FUNGI
whence they are taken. A large number of the woodcuts
will be recognised as formerly belonging to my Hanclhooh of
British Fungi, the use of which has now been granted by
Messrs. Swan Sonnenschein and Co. We are also under
obligations to the publishing committee of the Society for
Promoting Christian Knowledge, to the proprietors of the
Gardeners Chronicle, to Messrs. Kegan Paul, Trench, Trubner,
and Co., Messrs. Macmillan and Co., and Messrs. Chatto and
Windus ; for which they will please accept our acknowledg-
ments.
Beyond this brief prelude I need not advance, as, under
any circumstances, readers would prefer taking their evidence
from the chapters which follow than from any professions or
explanations in a preface. To the student who seeks for
assistance I have offered all that I had to give, in the hope
that it will be found sufficient for his need.
M. C. COOKE.
London, 1895.
CONTENTS
CHAPTER I
PAGE
Introduction ....... 1
PART I
OEGANOGEAPHY
CHAPTER II
The Mycelium ....... 9
CHAPTER III
The Carpophore . . . . . .22
CHAPTER IV
The Receptacle . . . . . .31
CHAPTER V
The Fructification . . . . . .41
CHAPTER VI
Fertilisation . . . . . . .53
viii INTRODUCTION TO THE STUDY OF FUNGI
CHAPTER VII
PAGE
DiCHOCAUPISM . . . . . . .64
CHAPTER VIII
Saprophytes and Parasites . . . . .73
CHAPTER IX
Constituents . . . . . . .84
PART II
CLASSIFICATION
CHAPTER X
Fungi in General . . . . . .95
CHAPTER XI
Naked -Spored Fungi — Basidioiitcetes . . .119
CHAPTER Xn
Hymenomycetes . . . . . . .126
CHAPTER XIII
Puff-Ball Fungi — Gasthomycetes . . . .149
CHAPTER XIV
AsciGERous Fungi — Ascomycetes . . . .164
CHAPTER XV
Discoid Fungi — Discomycetes . . . . .173
CONTENTS
CHAPTER XVI
PAGE
Subterranean Fungi — Tureraceae . . . .189
CHAPTER XVH
Capsular Fungi — Pyrenomycetes . . . .197
CHAPTER XVm
Gaping Fungi — Hysteriaceae . . . . .222
CHAPTER XIX
Conjugating Fungi — Phycomycetes . . . .227
CHAPTER XX
Rust Fungi — Uredineae . . . . .242
CHAPTER XXI
Smut Fungi — Ustilagines . . . . .251
CHAPTER XXII
Imperfect Capsular Fungi — Sphaeropsideae . . . 259
CHAPTER XXIII
Moulds — Hyphomycetes . . . . .277
CHAPTER XXIV
Microbes — Schizomycetes and Saccharomycetes . . 290
CHAPTER XXV
Slime Fungi — Myxomycetes . . . . .304
X INTRODUCTION TO THE STUDY OF FUNGI
PART III
DISTEIBUTION
CHAPTER XXVI
PAGE
Census of Fungi . . . . . .319
CHAPTER XXVn
Geographical Distribution . . • . . . 324
CHAPTER XXVm
Appendix on Collecting ..... 345
GLOSSARY . . . . . . .353
INDEX ........ 358
CHAPTER I
INTRODUCTION
The student will expect to find in an Introduction to the study
of any subject some definition and delimitation of that subject
— a task difficult in all cases to accomplish with brevity, and
within the compass of a technical description, but one of in-
creased difficulty when the subject is so extensive and com-
plicated as Fungi. In past times definitions have been hazarded
which appeared at the time to be incontestable and complete,
but within a short period they became insufficient. That they
are plants of a low organisation must be conceded, and also that
they belong to the lowest section, or the Cryptogamia, in which
the reproductive organs are more or less concealed ; but the old
characteristics of Algae as cellular plants subsisting in water ;
of Lichens as subsisting in air, and not upon the matrix on
which they flourish ; and, finally, of Fungi, which derived their
sustenance from the matrix, have had to be discarded as in-
sufficient. It is now known that aquatic Fungi are not an
knpossibility, that Algae may grow in a damp atmosphere, and
that some portion of the substance of Lichens may be derived
from their matrix. Seeing the difficulty of obtaining positive
characters, negative ones have been tried ; but these again have
failed to give satisfaction. In one of the most recent works
which has attempted to deal with this difficulty we meet with
the following as one of tlie " leading characters." It is to the
effect that " Chlorophyll, the green colouring matter so general
in the vegetable kingdom, is entirely absent from fungi." ^
Admitting this to be true, may it not be maintained that there
1 Massce, British Fungi (1891). ]>. 1.
1
c. r
No..:. college
2 INTRODUCTION TO THE STUDY OF FUNGI
are probably some lichens or some algae in which true chloro-
phyll is not present ?
But speaking of them as a whole, we are justified in saying
of Fungi that they are " cryptogams without chlorophyll," and
in this we shall embody the most important characters of the
group " — a general definition which may be accepted without
reservation. Except for one or two small fiimilies, we could
add also " without determinate sexuality."
Previous to this, Berkeley had pointed out that the
definition was imperfect which described Fungi as " deriving
nourishment by means of a mycelium from the matrix, and
never producing from their component threads green bodies
resembling chlorophyll," -^ for, he goes on to observe, " it is true
that a few Algae, such as Botrydium, do probably imbibe some-
thing from the soil by means of their rootlets, which can
scarcely be mere holdfasts " ; and again, " When we examine
Fungi more closely, we shall have reason to believe that there
are exceptions here also as to their deriving nutriment from
their matrix. I have, for instance, found a Cyphella on the
hardest gravel stones, where the fine mycelioid threads, by
which it was attached, could not possibly derive any nutriment
except from matters conveyed to it by the air or falling
moisture." To the latter portion of the paragraph, giving the
negative character of the absence of chlorophyll, Berkeley,
however, gives his adhesion.
A logical definition, therefore, so commonly fails, that we
shall excuse ourselves from attempting a new one, simply
indicating a few points to be borne in mind whilst perusing
the following pages, from whence alone a general idea can be
obtained of such a polymorphous group. Lindley divided all
the Cryptogamic plants into two sections, the Acrogens, growing
at the summit, including the Ferns, Mosses, and their allies,
and the ThaUogens, which embraced Algae, Fungi, and Lichens.
Hence we conclude that Fungi are not only Cryptogams, but
of that section in which there is no true root or distinct stem
with foliaceous appendages. Although the Rev. M. J. Berkeley
was, in the main, responsible for Lindley's classification of the
Cryptogams, it is out of date and inapplicable in the present
^ Introduction to Cryjriogamic Botany, p. 235.
INTRODUCTION
day, when other and improved methods have been brought into
use. Instead of the term Thallogcns for the celhilar Crypto-
gamia, it would be preferable to call them Thallojihytes, and,
for the rest, Brj/ophytes would include Mosses and their allies,
whilst Fteridophytes would be represented by the Ferns.
Thallophytes, in a general sense, which will be sufficient for
practical purposes, consist of those plants which grow in water,
and obtain their sustenance therefrom, conmionly known as
Algae ; and those which Hourish in the air, l^eing sustained by
the decomposition of the matrix on which they flourish, as
Fungi ; or drawing their sustenance from the air, and rarely, or
but slightly, from the matrix, as in Lichens. Here again a
negative feature may be interposed with advantage, to the
effect that Lichens are not of a Heshy or putrescent, but of a
dry and leathery consistence, whereas in the bulk of Fungi
the substance is, either entirely or in the early stage, soft and
fleshy, becoming indurated or putrescent with age. The
distinctions between Algae and Fungi will never cause any
practical difficulty, because the Saprolegnieac, which are
aquatic, and approach Algae most nearly in habit, derive their
sustenance from the matrix on which they are parasitic by
means of penetrating mycelial threads, whereas Algae are
simply attached by root-like or sucker-like extensions to the
matrix, from which nothing is absorbed. The relations between
Fungi and Lichens are much more intimate, and in extreme
cases approach each other so closely as to be distinguished
with difficulty even by experts. Whole genera are still
claimed by mycologists on the one hand, and by lichenologists
on the other. Still it must be remembered that these are
extreme cases, and that amongst the larger Fungi, especially in
the great group of Hymenomycetal Fungi, there is but little
suggestion of Lichen affinity except in such genera as Cora,
Dictyonema, Pavonia, etc. Another exceptional case may be
found in the Nostoes, which are Algae, simulating or so closely
resembling Tremella, a genus of Fungi, that microscopical
examination may be necessary for their discrimination.
Habitat may in some cases serve to indicate the character
of the Thallophyte. For instance, all the parasites on living
leaves which are not of insect origin are Fungi, such as the
4 INTRODUCTION TO THE STUDY OF FUNGI
smuts and rusts which are so destructive to crops. There are
often to be seen discoloured spots upon living leaves which
cannot be attributed either to insects or Fungi. It is a common
mistake with novices to infer that all leaf spots are necessarily
caused by parasitic Fungi. Again, all the fixed or vegetable
parasites on insects are Fungi, whether moulds or the larger
club-shaped species of Cordyceps. Another caution becomes
necessary lest the pollinidia of Orchids, which are sometimes
seen temporarily attached to living insects, should be con-
founded with true parasites. Putrescent vegetable, and some-
times animal substances, give rise to Fungi, but dead wood and
the bark of trees may also support Lichens as well as the
living bark, on which Fungi are rarely found, except in cases of
incipient decay. No difficulty need be anticipated in respect to
Thallophytes found growing upon the ground, inasmuch as the
Lichens which have a terrestrial habit would scarcely be con-
founded with Fungi at any time, but especially after the
perusal of succeeding chapters on the details of structure in
the several orders. It should be understood that the above-
named distinctions are not so much of scientific value as they
may be useful as guides to collectors.
The mycelium of Fungi is so general, although at times
nearly obsolete, tliat it must be regarded as somewhat analogous
to the thallus of Lichens, but not to be confounded therewith.
In the Agarics this mycelium is commonly termed the
" spawn," and consists of delicate threads, which traverse the
soil or the rotten wood upon which the Fungus is grown. In
some instances a strong mycelium is developed, but no perfect
Fungus is produced upon it. An instance may be found in the
substance called Xylostroma giganteum, which forms thick sheets
like leather, destructive of wood of solid texture. It is doubt-
less a degraded form of wood-destroying Hymenomycete.^ The
moulds and the mucors produce at first decumbent barren
threads, which constitute the mycelium out of which sub-
sequently the fertile threads arise. In the " rusts " or Uredines
the cushion-like base of the pustules is surrounded by the
delicate threads of mycelium. Going back to its origin, we
discover that the initial spore, or spores, upon germination pro-
^ See next cliapter, p. 10.
INTRODUCTION
duces a delicate thread wliicli, either directly or indirectly, origin-
ates the mycelium upon wliicii the Fungus of the new generation
is developed. In the Agarics it is held that a large numl)er
of spores germinate, and produce the mycelium from which
a single individual or a cluster of young Agarics are evolved.
In some instances the mycelium is undoubtedly perennial, and
produces a crop of Agarics in successive years, but much remains
still to be known of the life -history of the Agarics in the
interval between the maturity of the spore and the first
evidence of the genesis of a new plant.^ In some of the smut
Fungi the germinating spore produces a thread which develops
secondary spores, and these in their turn produce tertiary spores
before the true mycelium-forming spores are developed. In the
Uredincs the earliest spore-forms, called "pro-mycelial spores," are
produced from the germinating threads of the latest spore-forms
or teleutospores, which in turn give rise to the mycelial threads ''
that enter and form a new mycelium within the tissues of the
invaded host-plant. In the Erysiphei the mycelium forms an
external coating on the surface of the living leaves, producing
at first conidia, and ultimately the perithecia or spore^capsules
of the perfect Fungus. Indeed, as a rule, the mycelium repre-
sents the vegetative system of the Fungus upon which, under
varied forms, the reproductive organs with their appendages
are produced. The universality of this mycelium in Fungi
was formerly held to be as certain an indication of distinction
between a Fungus and a Lichen as the production of a thallus
was then held to be a sufficient distinct,ion between a Lichen
and a Fungus. In later times it has come to be understood
that the hyphal elements in Lichens and Fungi are virtually the
same.
' In this connection may be consulted Brefeld's researches into the life-liistory
of Coprimis stereo rarius.
- See j)ust, chap. xx.
BIBLIOGRAPHY
Beiikeley, M. J. Outlines of British Fungology — Introductory Matter. 8vo.
Plates. London, 1860.
• Introduction to Cryptogamic Botany. Roy. 8vo. Cuts. London, 1857.
Articles on " Vegetable Pathology," in Gardener s Chronicle, from 1855.
LiNDLEY, J. Vegetable Kingdom. 8vo. Cuts. London. 2nd Edition. 1847.
CouKE, M. C. Fungi: their Nature, Influence, and Uses. Sm. 8vo. Cuts.
London, 1875.
A plain and easy Account of British Fungi. Sni. 8vo. Plates. London.
Itomance of Low Life amongst Plants. 8vo. Cuts. Loudon, 1893.
BouDiER, E. "Considerations sur I'fitude microscopique des Champignons."
Bulletin de la Socitti Mycologique de France, vol. i., 1885, p. 134.
De Bary, a. Comparative lUorphology and Biology of the Fungi, Mycetozoa, and
Bacteria. English Translation. Roy. 8vo. Cuts. Oxford, 1887.
Tavel, F. von. Vcrgleichende Morphologic der Pilze. Jena, 1892.
PAKT I
OKGANOGEAPHY
CHAPTER II
MYCELIUM
In such an imineiise group as the present, incUiding ahnost an
infinite variety of form, it is extremely difficult to generalise,
even the morphology, beyond that of the mycelium, which re-
presents the vegetative system, the carpophore, which supports
the fructification, and, finally, the fructification itself, with the
organs associated therewith. This, in fact, reduces the whole
scheme of structure to its lowest terms, that of the vegetative
system and the reproductive, since the carpophore is but a
development of the vegetative, and a link between that and the
fructification which it is intended ultimately to bear. The
details of the varied modifications, which are to accomplish the
two purposes of growth and rejuvenescence, can only be de-
scribed under the great primary divisions of Fungi, where
both vegetation and reproduction conform to some definite
type. As a whole, the organisms which are associated to-
gether under the common denomination of Fungi are the most
protean and polymorphic in the entire vegetable kingdom,
and present great difficulties in the way of generalisation.
If we gather a mushroom from a mushroom bed, as usually
cultivated, we shall discover, if we remove it carefully, that
the stem, which represents in this instance the carpophore, or
fruit-bearer, is attached to the soil by a mass of delicate white
hyphae, or threads, which are the mycelium, or spawn. And
if we remove the soil anywhere, we shall find that it is per-
meated in all directions with these white threads. The
artificial "bricks," or spawn, which were employed in the
construction of the bed, consisted of this mycelium in a
quiescent condition, and by supplying sufficient moisture and
lo INTRODUCTION TO THE STUDY OF FUNGI
heat, with a suitable matrix, the filaments of this mycelium
have been revivified, and by a profuse and rapid growth they
have spread over, and penetrated the whole of the soil of which
the mushroom bed is composed, and constitute the vegetative
system of the mushrooms which afterwards appear on the
surface. This mycelium represents an important element in
the morphology of all Fungi. It is rarely reduced to such
small proportions as not to be recognised, but, practically, it
is possibly never wholly absent. We may start, therefore,
with what we may regard as an essential attribute of Fungi,
and the representative of the purely vegetative system. Where-
ever we see a Fungus of the mushroom type, whether we please
to call it a mushroom or a toadstool, we may find the mycelium
in the soil from which it springs. In the autumn, if we stir
up and turn over any clump of dead leaves or other decaying
vegetable matter in a damp situation, we shall be sure to find
a profusion of this mycelium, even though no perfect Fungus
makes its appearance, and it is at work on every dead stump
and every fragment of rotten wood.
Mycelium consists of hyphae or threads, usually septate,
sometimes simple, but mostly branched, increasing by growth
at the extremities, and gregarious, so as to form reticulated
interwoven masses, either in a thin network or a densely-felted
mass. In a certain sense they are analogous to the roots of
flowering plants, and, like them, draw moisture and inorganic
constituents from the soil or other matrix on which they may
be developed. It is another axiom with Fungi that, by means
of the mycelium, they derive their sustenance from the matrix
on which they grow. It cannot be doubted that the growing
points of the mycelium possess the power of penetration by
the production of a ferment, since they are capable of pene-
trating the hardest wood, disintegrating the cells, and reducing
it to powder. The ordinary mycelium found amongst dead
leaves has a power of disintegration, and soon reduces them,
as well as twigs and stems, to a condition of humus ; but the
progress of mycelium in a dead trunk is quite as definite and
certain. Who can doubt the disintegrating power of the
mycelium of the " dry rot," and it must be borne in mind
that it is the mycelium in this instance which works the
M YCELIUM
mischief. Tlie hyphae forming this vegetative system may be
immersed, and prohaljly are so in the majority of instances,
but they may also form a stratum on the surface, and adhere
by haustoria or suckers, which are short brandies designated
for the purpose, or for that combined with the absorption of
nutriment. Tlie hyphae when young have colourless cell walls,
but as they grow older the walls thicken and acquire colour,
sometimes with an appearance of stratification. In some
cases cross branches anastomose, or form clamp
connections (Fig. 1). Gardeners are well aware
that immense masses of white mycelium are some-
times met with in turning over tlie soil. These
mark the site of an old tree when part of the
stump or dead roots have been left to rot in
the ground. Numerous instances are on record
in which trees or shrubs, when planted in soil
overrun in this manner with mycelium, have
been killed, and when taken up the roots Ibund ^^^ ^ _ (^;^
to be enveloped in mycelium. I'ractical men connections.
are quite aware that this has occurred over ^^"^^ ^ '''^^'"
and over again, and yet some theorists contend that it is
not possible, because the mycelium is a saprophyte, that
is to say, flourishes upon dead organic matter, and not
upon living tissues. Observation has nevertheless decided
that in some instances a saprophyte may become parasitical,
and a parasite may acquire the habits of a saprophyte. The
instances may not be common, but are not impossible. Against
theory we are prepared to contend, from experience, that we
have seen plants killed, after planting in a soil overrun with
mycelium, from no other assignal)le cause, and afterwards dug
up with the roots enveloped in mycelium.
It has been stated above that mycelium is usually colour-
less, and it is generally so with the Basidiomycetes, but there
may be exceptions, as in Corticium sangitineum, with its
mycelium of a blood red, in Elaphoviyces LcveiUci, of a yellowish
green, in Chlorosplenium aeruginosum, of a verdigris green, and
in many of the Dematiaei, of a dark brown, or almost black.
And so also it may differ considerably in quantity, from a few
scattered threads to a dense matted felt. Even in tliis country
12 INTRODUCTION TO THE STUDY OF FUNGI
it sometimes forms thick Laminae many inches broad in old
trunks, but in warmer countries it develops into a spongy-
mass, called Xylostroma in past times. These masses will fill
up holes in a log nearly as thick as the wrist, and a foot or
two in length, or they will spread in layers of an inch in
thickness, a foot in breadth, and several feet long. The whole
mass is made up of interwoven threads, almost as dense as cork
when felted together, but wholly barren, so that it is uncertain
whether they are the mycelium of a Polyporus or an Agaric.
Having been furnished with a redundant supply of nutriment,
they never advance beyond the vegetative stage. In the case
of entomogenous Fungi, the mycelium will replace the whole
of the tissues, even to the legs and feet, so as to form a com-
plete cast of the insect, of which only the dermal covering
remains unchanged. In such genera as Cordyceps and Empusa
the filamentous structure is only seen in the earliest stage ;
this soon gives way to a compact granular mass. The mycelium
of the Mucedines, or moulds, when abundantly supplied with
moisture, develop rapidly and vigorously, but do not proceed
with the fructification whilst the nutriment supplied is abnor-
mally great. The conidial stage of the Erysvphei will furnish
examples of a thin superficial mycelium adhering by haustoria.
In these cases a thin white web runs over the surface of living
leaves, as in the vine mildew, and a mildew on the leaves of
the maple, but it does not penetrate deeply into the tissues of
its host, which it injures by choking up the stomata. Another
superficial mycelium is that of Fumago, which forms black
patches on the leaves of the lime and other trees, being especi-
ally vigorous on those subject to honey-dew. In the sphaeria-
like or capsular Fungi, the mycelium is confined, usually, to a
few delicate threads at the base of the perithecia, but there
are exceptions to this in some superficial species, where a
subiculum or conidia-bearing mycelium is present.
Another form assumed by mycelia is that condition whicli
has long been known under the name of Rhizomorpha, when it
was suspected to be an independent Fungus, although no form
of fruit had been discovered. It is now admitted that the
several species are only the vegetative condition of other Fungi.
One kind may be seen running between the bark and wood oi
MYCELIUM 13
dead trunks in long brown or black cords, as thick as whip-
cord, flexible, but tirm, and either branched or joined by
cross connectives of the same substance, into a coarse net-
work. These Ioiil;' cords may be many feet in length, and
whitish internally, with a dark-coloured outer coat or skin.
The tips of the growing branches are paler. This peculiar
growth is very connnon in mines and other dark places, and
glows sometimes with a phosphorescent light. Tulasne
examined some specimens with a view to the discovery of
the cause of their luminosity, of which Humboldt, amongst
others, had given such an elaborate account. He found that
all the young branches brightened with a uniform phosphoric
light the whole of their length, and also the surface of some
of the older branches. The latter when split open were dull,
but after exposure for a time to the air they also became
luminous. By keeping them moist, they preserved their phos-
phorescence for several days. He also states that branches
which had been dried for more than a month, when plunged
into water, revived, and began to vegetate afresh, in a few days,
by sending forth numerous branches, but they were only
luminous on the surface of the new parts. One of our
commonest Agarics, to be found on nearly every rotten stump,
Agaricus melleus, is credited with being the complete develop-
ment of one of the species of Rhizomoiyha, which may be
stated in this way — the cord-like Rhizomorjpha is simply the
persistent mycelium of Agaricus melleus, which grows on
rotten stumps. We have no doubt that another form or
variety of lihizoviorpha is the mycelium, or vegetative con-
dition of Polyporus squamosus, and others, with more or less
certainty, are referred to other species of Agaric and Polyponis.
Something of the nature of Rhizomorpha is found amongst
dead leaves, mostly in long, simple, rigid black threads, which
in size and appearance are not unlike horse hair. These are
believed to be the mycelium — or rather, we should say, the
permanent mycelium, to distinguish it from the filamentouf.
white mycelium — of some species of Marasmiits. In tropical
and sub-tropical regions these horse- hair filaments are very
common amongst dead leaves, and are known to be sterile
conditions of several species of Marasmiits.
INTRODUCTION TO THE STUDY OF FUNGI
There is another condition which the mycelium of some
Fungi assume that is something of a resting stage, and in
former days these were classed under a genus called Sclerotium.
They are in the form of hard, compact, irregular nodules, from
the size of a pin's head to that of a child's head, according to
the species. They are mostly dark coloured externally, and
nearly white and horny within, with a firm cellular substance.
We will conmience with one that is well known, under the
name of Ergot, and occurs on the spikes of rye, wheat, and
many grasses, converting the
ovary into a Sclerotium, or
replacing the ovary with a
Fungus growth, whichever
view pleases best. These
abnormal growths are three
times as long as the ordinary
seed of the rye or grass,
elongated, and a little curved,
so that they look like a horn
or spur projecting from the
ear of grain. If a thin slice
of the substance is placed
under the microscope, it will be seen to consist of a densely
compact mass of cells, somewhat irregular from mutual pressure,
with thick walls and rather oily contents. At last they are liable
to become dissociated from the spike, and fall to the ground,
where they will lie quiescent and unchanged through the winter.
When spring arrives, active vitality recommences, and the sclero-
tium germinates by producing one, two, or more — generally several
— little slender twisted stems, with a globose head, reminding
one of a pin. The stem is whitish, and the head of a pale purple.
It is within the globose head that the fructification is produced.
This latter is the mature Fungus of the Ergot sclerotium, and
is then called Claviceps jmrpurea, one of the Sphaeriacei (Fig. 2).
By way of illustration, we have indicated briefly the history
and development of the Ergot sclerotium, but it will scarcely
be consistent with the design of this chapter to repeat the
process for other species. Another example may be found
inhabiting the dead haulms of potato. . First of all the haulms
FiQ. 2. — Ergot Sclerotia germinating.
After Tulasne.
MYCELIUM
15
are covered with a dense felted mass of wliite mycelium.
When this is fully developed, scores and hundreds of little
black points appear in the midst of the hyphae, averaging in
size I'roni a u;rain of sand to that of a small bean. These
become indurated and hard, and, in fact, properly-constituted
sclerotia, internally composed of polygonal cells. These also
subside into a con-
dition of rest, in
which they spend the
winter, and germinate
in spring. The re-
sulting Fungus in this
instance consists of
similar slender twist-
ed stems, but the
head, instead of being
globose, is cup-shaped,
then flattened, bear-
ing the name of
Sclerotinia Libertiana ,
or, as known in former
years, a species of
Peziza, one of the Dis-
comycetes (Fig. 3).
Several other instances might be quoted in which the Sderotium,
when germinating, produced a species of Peziza, especially a large
one common in company with the roots of the Wood Anemone.
A large species of Russula, common in the woods, turns quite
black when dead and decaying. On the gills of these decaying
Bussulae many sclerotiae will be seen, resembling in form small
grains of barley. These germinate speedily, and produce a little
Agaric {Collyhia tuherosa). In Australia a sclerotium nearly as
large as the fist develops a tough gill-bearing Fungus, shaped
like a wine-glass, named Lcniinits ajathus. But a still larger
sclerotium, which has been known for years as " Native liread,"
and grows as large as a cliild's bead, has been recently found
to develop a white Polyponts with a central sttMu, and has
been named Pulyporus MyLiltac}
1 Orcvillca, Dec. 1892, \\ 37.
Fig. 3. — Sclerotia gerniiuating and producing Pezizae,
A to D. Ascus and sporidia, E. Gard. Vhron.
1 6 INTRODUCTION TO THE STUDY OF FUNGI
We have biietly directed attention to the ordinary develop-
ments of mycelium, whether filamentous or sclerotioid, but
there are still one or two special modifications which must
obtain a passing reference. Of these the mycelium of the
Uredineae is deserving of mention, being formed within the
tissues of living plants, and often starting centrifugally from a
definite point of infection. The hyphae resemble ordinary
mycelial filaments, but, like all internal mycelia, are delicate,
branching and anastomosing so as to form compact cushions or
spore beds, or, in other cases, much diffused and scattered. In
annuals or upon deciduous parts the mycelium is, of course, only
annual, but if it passes into perennial parts, as it may readily
do in shrubs and other perennials, the mycelium becomes
perennial. Take, for instance, the Juniper, in which the
Crymnosporangium may be sought for and expected regularly
year after year. In such species as have a scattered mycelium
there is not much difference between the mycelium of these
and that of other endophytes, but when the mycelium is
circumscribed, the tissues are hypertroj)hied, and starch seems
to be accumulated in the deranged cells. Leaves thus attacked
never repair the injury, and the diseased spots are the first to
die, and occasionally drop out, as we have seen with the large
clusters of perfect-spored pustules in Puccinia deanthi. The
mycelium of the Feronosporeae is more diffused than that of
the Uredines, commonly penetrating the whole plant, descend-
ing into the stem and roots, and in the stems producing oospores,
as the result of a sexual conjugation. In the Ustilagineae the
mycelium is much more diffused than is usual in the Uredines,
permeating the entire plant, and in perennial hosts producing
fruit regularly year after year.
The Phycomycetcs, which include those mould-like Fungi
which bear inflated sacs at the apices of their fertile branches
containing numerous spores (Ilucors), have a mycelium without
septae. The Mucors themselves are mostly saprophytes, and
some of them have a profuse mycelium. The reproduction is
both asexual and sexual, the sexual being developed from the
mycelium ; hence the mycelium in these Fungi, although at first
only vegetative, becomes finally reproductive, and thus assumes
a higher function. The process is after this kind : two short
MYCELIUM
17
branches of the mycelium approach each other until they touch
at their apices. The branches swell and become club-shaped,
including a rich store of protoplasm. At length the upper
portion of each club is cut off by a transverse septum, and the
two apical segments are fused into a globose body, the walls at
the point of contact being dissolved. Thus we have two thick
supporters, with a globose body suspended between them, which
is to become the zygospore, resulting from the conjugation and
coalition of the club-shaped branches (Fig. 4). The succeeding
steps need not be detailed ; the zygospore acquires a thick outer
coat, and then becomes a resting spore, which only germinates
after a period of rest. The same mycelium therefore produces
Fia. 4. — Zygospore of Mucor in course of formation. After De
erect carpophores, or couidiophores, surmounted by an inflated
vesicle containing conidia, an asexual generation, and also pairs
of nearly sessile branches, which collide and form a zygospore
between them, a sexual generation. Similarly, in other families
the two kinds of reproduction are developed, asexual and sexual,
from different parts of the same mycelium, but not precisely
in the same way, yet the details do not affect the mycelium
greatly, except in the family to be presently alluded to.
The Entomophthoraccac^ are those Fungi which are parasitic
and destructive to insects, including the ordinary fly mould,
' The Enlomophthorcac of the United States, by Roland Thaxter, 1888,
Boston, U.S.
i8
INTRODUCTION TO THE STUDY OF FUNGI
Fig. 5. — Hyplial bodies
in Entomophthoraceae.
After Thaxter.
Empusa iiuiscae. When the spore of one of these moulds alights
upon the body of its favourite host-insect it sends out a germ
tube, which enters the body at any favourable spot, and when
this is once accomplished, it develops rapidly, at the expense of
the tissues it replaces. It does not form a branched mycelium,
but grows by the production of hyphal
bodies (Fig. 5), which are short, thick
bodies of variable size and shape, and these
continue to nmltiply, by budding or gem-
mation, until they hll the insect. It is
possible that in some cases a mycelium
of the ordinary kind may be produced.
When the whole interior is absorbed,
and of course the insect is dead, the vegetative stage ends, and
the raproductive begins, by the protrusion into the atmosphere
of conidiophores terminated by conidia, either singly or in
bundles, until the body is covered with the conidia, ready to
be dispersed. This is the asexual reproduction of conidia, but
resting spores are also formed, which may be sexual or asexual,
according to the species. In some cases
the conjugation of two threads of the
mycelium, and in others the conjugation
of two of the hyphal bodies (Fig. 6), results
in the formation of a zygospore. There is
a peculiarity about the conidia which may
be noticed, which is, that should a ripe
conidium not be able to find or enter
a host-insect, it can proceed to germinate
and form a secondary conidium, which
has the same power of infection, and
may be more fortunate. If this also
fails, the secondary conidium may pro-
duce a third, so that the vigour of the
conidia is kept preserved until able to
infect a host. Possibly the Isaria moulds, in the interior of
insects, extend their mycelium in a similar manner by budding,
as they are also granular rather than filamentous.
In opposition to the views of some mycologists of ex-
perience and repute, we still remain persistent in our adhesion
Fig. 6. — Hyphal bodies in
Entomophthoraceae con-
jugating. ; After Thaxter.
MYCELIUM 19
to the vegetable nature of the Myxomycetes, and consequently
regard the vegetative condition as upon an equality in function,
if not in structure, with the mycelium. This was clearly the
view of ]\I. Luveille, who termed it " a malacoid or pulpous
mycelium." We see no objection to its being called a Plas-
modium— the name does not alter its character or functions.
Dc Bary was content to admit that the wall of plasmodia, as
well as the cell-walls of spores and other parts, gave a distinct
cellulose reaction, and possil.)ly cellulose in some form is general
in ]\Iyxomycetes. And further, according to the same eminent
authority, the presence of cellulose is the only character show-
ing that these organisms are in touch with the vegetable
kingdom. This question we are not anxious to discuss further
here. Swarm-cells with the power of movement are produced
on germination from the spores of ]\Iyxomycetes ; these swarm-
cells ultimately coalesce and form a plasmodium, which is capable
of passing into a resting stage, and sometimes to become sur-
rounded by a colourless membrane. There are no threads or
filaments, as in a filamentous mycelium, neither in the sclerotioid
mycelium is there a similar resemblance ; in fact, there is no
greater difference between a plasmodium and a sclerotioid my-
celium than there is between a sclerotioid and a filamentous my-
celium. From the plasmodium are differentiated the carpophores,
the receptacles, and the fructification of a Fungus, even although
the Plasmodium or analogue of mycelium is not filamentous, but
rather resembles a sclerotiuni in a soft and pulpy condition.
Theoretically, mycelium originates with the germ -tubes
which are protruded by spores or conidia upon their germina-
tion. It is easy enough to observe the process thus far,
produced artificially by placing the spores in a nutritive fluid,
but in the case of the larger Fungi the operation cannot be
carried much further under ordinary circumstances. In the
case of Agarics it is concluded that the mycelium produced by
a number of spores unite in the production of a single Agaric,
so that one specimen is the produce of several germinating
spores. We know that the soil contains a great mass of
mycelium in places where Fungi are found growing. Worth-
ington Smith says ^ that the Agarics of the autumn spring up
' Reproduction in Coprinus, Grevillca, iv. (1876), p. 53.
20 INTRODUCTION TO THE STUDY OF FUNGI
from the mycelium formed during the fall of the previous year,
and this mycelium has rested in the ground for twelve months.
In digging up old pasture ground, or the dead leaves of an
autumn which has passed, mycelium in a resting state is
invariably found. We can hardly conceive of the preservation
of the spores of an Agaric through the winter and an entire
year, until the succeeding autumn, in any other way than by
the production of a hibernating mycelium. The spores them-
selves have too delicate an epispore to resist the effects of cold,
and we know from analogy that the resting spores of Algae
and Fungi, when known to be such, are provided with a special
thick outer envelope. The spores of Agarics are not thick
coated, and are incapable of hibernation ; hence we are driven
to the alternative of a perennial mycelium. A theory was
once propounded that a conjugation takes place in the threads
of mycelium which results in the production of a fertile Agaric,
the whole of whose fructification is thereafter rendered fertile,
but this view has never been accepted. Notwithstanding all
the theories, w^e are still in search of the process of fecundation
in Hymenomycetal Fungi. All that we can contend for is the
persistency of the mycelium as the means whereby the Mush-
room Fungi are carried through the winter and reproduced in
the succeeding year.
There is a prevalent opinion, in Germany at least, that
" root fungi " are not always injurious to trees, but sometimes,
on the contrary, beneficial. Frank ^ states that certain trees
are unable to derive nutriment direct from the soil, but do this
by means of a mass of Fungus hyphae which entirely invests
the root, to which he gives the name of Mijcorhiza. It makes
its appearance first on young seedlings, and is replaced by
fresh formations on older roots. He found it on the roots
of every tree examined belonging to the Cupuliferae, and
occasionally on willows and conifers, but considers it may
only be formed in soils which contain a large amount of
humus, or undecomposed vegetable remains. Through the
Mycorhiza the tree absorbs not only water and mineral con-
stituents, but organic substances derived from the humus.
Two or three other authors have since confirmed this in
1 Journ. Roy. Micr. Soc, vol. v. (1885), p. 844 ; vol. vi. (1886), pp. 113, 663.
MYCELIUM
the most important particulars, but not as to its constant
presence.
The mycelium is the active a«,fent l)y wliich Fungi disin-
tegrate decaying organic matter, or prey upon and destroy the
living, and so far as they derive nourishment from the sub-
stratum, their nutrition resembles that of flowering plants, but
beyond this the mycelium is active in decomposing the organic
matrix, the product of which is not required or taken up by
the Fungus. Hence there are forms which are satisfied with
taking up from living or dead substrata only so much as is
needed for the construction of their bodies, as well as those
which in addition produce copious decompositions in the
substratum and destroy it. We may assume that the mycelium
exerts a ferment action upon the matrix, although the quantity
of the ferment may be small, and that these ferment actions
first take place in order to convert a portion of the substratum
into a form which is capable of nourishing the Fungus.
In the headings of the several chapters we have used terms
in their general sense, representing the mycelntm as equivalent
to the vegetative system, the carpopliore as the supporter of
the fructification, or intermediary between the vegetative and
reproductive systems, whilst receptacle is employed in a sense
different from that which it holds in other branches of botanical
science, and should be accepted literally as representing the
envelope of the fructification, whatever its form may be, when
any envelope is present. This definition is necessary so as to
prevent confusion of the terms we have employed in a general
sense, with their special application elsewhere.
CHAPTEK III
THE CARPOPHORE
The mycelium, in all its forms and variations, is but the
prelude and preparation for the development of such parts or
organs as may be necessary for the subsequent processes of
reproduction. The production of the carpophore is, in itself,
only a continuation of the process of vegetation, but that
vegetation is no longer subterranean, subcuticular, or creeping ;
invariably it is more or less, in development, at right angles to
the mycelium, and may be accomplished by the production of
special erect branches, or a stem compounded
of an indefinite number of erect threads,
agglutinated and consolidated together ;
whether it is to be the conidiophore of a
mould, the stroma of a Cordyceps, the club of
a Gcoglossum, or the stalk of an Agaric or
Boletus, it is the fruit-bearer, or carpophore,
which is destined to bear the fructification of
-Unbranched the spccics. It may be reduced to its lowest
carpophore of terms, and be practically obsolete, so that the
RhojKi.lomyces. ^ "
receptacle is sessile, or nearly sessile, upon
the mycelium ; still there is normally and technically a
carpophore, which supports the organs of reproduction.
In the larger moulds generally the ascending hyphae are
branches of the mycelium, and do not alter much in character
except in being rather thicker and with more rigid walls, so as
to maintain an erect position. These erect threads are in
most cases clustered together, and are modified in ramification
^ The term " carpophore," in its special sense, is usually restricted to forms of
a distinct fruit, consisting of an aggregate of reproductive organs.
THE CARPOPHORE
according to the different genera. In certain cases, as in
Aspcrcjillus and Iiho2)alomyccs, tliey are simple and unbranched
up to the top (Fig. 7), but in the hirger number of genera they
are branched in the upper portion. Very often a great
number of these car-
pophores are produced
in a large woolly -
looking patch, not
rarely for an inch or
two in length. Endo-
genous moulds, which
produce mycelium in
the interior of the
tissues, send up little
tufts of carpophores
through the stomata,
and these grow in
patches. Well-known
examples are to be
found in the genus
Peroiiospora, such as
the mould on parsnips and onions (Fig.
Raimdaria the mycelium is internal, and the conidiophores pass
in the same manner out into the atmosphere ; but they are
usually short, often unbranched, with a single conidium. In
Oidium the mycelium is external, and the erect hyphae are
simple, but it is only the short lower portion which is truly a
carpophore, for the upper portion is constricted successively,
and the joints fall off as they are formed, and become
conidia.
There are also genera in which the carpophore is compound
— that is to say, a number of threads are combined so as to
form a common stem, which is consequently thicker and more
permanent. Either these individual hyphae diverge at the
apex, or they remain united and form a ca]»itulum, as in Stilhum.
When- the combined threads form only .1 sliort erumpent
stroma, as in Tiibercularia (Fig. 9), the carpophore is reduced
nearly to its lowest denomination, and is scarcely more than
an erumpent pustule. .Ml the foregoing forms are repeated in
Fi>i. 8.
Br.iiiehed c-uriiopliorf of l't:ronospora.
8). In the genus
INTRODUCTION TO THE STUDY OF FUNGI
"xjI.
— Compound carpophore
of Tubercularia.
the Dematioei, or black moulds, the chief difference being in the
dark-coloured, more rigid, and carbonised hyphae.
Eesembling the moulds in external habit, the Mucors
resemble them also in the carpophore, which is sometimes
forked two or three times, but not dendritically branched. In
Pilobolus the carpophore is curiously inflated, like a bladder
(Fig. 10).
We have in remembrance a pseudo-analogy which some
few years since became current — that
the type of organisation in a Muce-
dinous mould was repeated, with
modifications, in the structure of
Agarics. The mycelium, it was con-
tended, was common to both. From
the mycelium arose the carpophore,
which was a compound stalk, in
which a myriad of erect hyphae
were combined ; in the pileus the combination was continued
of the branches, and then down to the basidia, which
were the terminals of the branchlets, with the spores, or
conidia on spicules, as in such a genus as Bhiiiotrichinn.
This was a fanciful representation, since the
analogy, even if it held good elsewhere, was
broken at the hymenium, and the basidia were
therefore not in continuity with the trama. In
the Hymenomycetcs, or at least in the Agaricini,
the stem is continued from the mycelium at
right angles, as in the moulds, and is compounded
of an infinity of elongated parallel cells ; these
are sometimes deficient in the centre, and the
carpophore, or stem, becomes hollow. Leaving
the appendages to the stem out of question, it
is still an erect carpophore, and hence its func-
tions are the same — that of elevating the re- fig. lo.— inflat-
productive organs into the atmosphere. In ed carpophore
the same manner also the materials of nutrition,
derived by the mycelium from the soil, are conveyed upwards
to the residue of the plant. The veil, where it exists, is
jupplementary appendage, not found in the moulds, and
THE CARPOPHORE
simply an extension of the margin of the receptacle, or pileus,
for the projection of the young hymenium. In Boletus and
the stipitate Polyporci, Htjdnci, etc., the carpophore is of the
same type. In such of the species of Flcurotus, Fomes, etc.,
as have no stem the pileus, or receptacle, is sessile, and the
carpophore is reduced to a mere disc, or is obsolete.
The external surface of the stipe or stem is sometimes
glutinous, as in the section Myxacium of the genus Cortinarius ;
or it is velvety, as in such species as Collyhia longipes and C.
vehitipes ; or it may be woolly, chiefly at the base, or broken up
into scales ; and all these conditions doubtless serve in some
way to fulfil some purpose. AVorthington Smith has suggested
that they are probably of service to arrest the spores as they
fall from the hymenium, and, as he thinks, also the deciduous
cystidia. Of the internal structure M. de Seynes remarks
that the collective cells, which form the stipe, and afterwards
expand into the cap, are generally rather uniform, long, fibrous,
often much separated, rarely ramified, presenting at times in
their distance from each other, at others in their dimensions,
differences which, on the fissure of the stipe, present an aspect
either fibrous, granulated, spongy, or woolly. The cellular
fibres are always closer and more compact at the cortical part.
Those peculiar lactiferous vessels which convey the milk, so
conspicuous in Ladarius, are not confined to the cap, but are
present also in the stem, although possibly not quite so
abundant, but they must be very numerous in the stems of
one section of the genus Mycena where the milk is almost
confined to the stem. In Mycena leucogalus it is quite white,
in Mycena haematopus, of a blood red ; in Mycena crocatus it is
saffron yellow, and in Mycena yalo2ms it is described as white,
but it is often watery, or with a tinge of white, like milk and
water. The quantity of milk depends much on the dampness
of the habitat.
In such degraded forms as Cortirium, liachdum, etc., the
carpophore is obsolete, and the receptacle is reduced to a
fibrous stratum, which is seated directly upon the mycelium,
and only the hymenium receives its proper development.
Other genera require little observation, since in some forms of
Theleplwra, in Lachnocladiuvi, and in Clavaria and its allies,
D. . -r
Nortli i^c. o..r.- , .. College
26
INTRODUCTION TO THE STUDY OF FUNGI
we have the closest resemblance to the carpophores of the
moulds, even to dendritic branching, but of a larger and more
robust habit. Isaria is often closely imitated in external
appearance by Clavaria. The most anomalous of all groups in
respect to the carpophore is the Tremellini, but even in this
there is a link in Chicjnnia, Gyrocephalus, and Ditiola. We
have not forgotten that in some of the Basidiomycetes the
whole of the Fungus, in its earliest stage, when seated upon the
mycelium, and before the development of the carpophore, is
enveloped in a volva. This is not, however, more than a
generic distinction, in any case, and reaches its highest de-
velopment in Amanita, Volvaria, Ithyphallus, Clathrus, etc.
It might be compared to the calyptra in mosses and liver-
worts, but is by no means so general, and without so much
significance.
The Gastromycetes are not so well provided with a carpo-
phore as the Symenomycetes, but at the same time
there is no degradation to resupinate or overturned
forms. The majority of the Phalloid Gastromycetes
have a distinct carpophore, which is functionally the
same as in Agarics, but the structure is more loosely
cellular (Fig. 11), and, from rapidity of growth,
lacunose. It is only in Fungi like these, which
are quick to decay, that we encounter such a loose
texture of cells in the carpophore. In Podaxis
the form of the entire plant resembles that of
Coprinus, but the carpophore is rigid, almost
it is also in Batarrea, Xylopodium, and
The carpophore in Secotium approaches
of the coriaceous Agaricini, such as
There are no other genera which call
hhyphaiius for Special notice respecting the carpophore, which
impudicus. ^g short, and almost spurious in Scleroderma and
Polysaccum, rare in Lycoperdon, and then only a prolongation
of the spongy base of the receptacle. In nearly every genus
except Gyrophragmmm, Podaxis, and Secotium, it expands into,
and is confluent with the receptacle. The subterranean species,
like the truffles, have no carpophore.
It is not difficult to comprehend the functions of the
woody, as
Tylostoma.
the type
Lentinus.
THE CARPOPHORE
27
carpophore in the llymcnoiiiycctcs, in till of which the hymenium
is inferior, and therefore it is essential to its development that
the receptacle should be raised sufficiently above the matrix
to permit of a free development of the hymenium. The
carpophore is only suppressed or obsolete when the receptacle
grows out at riifht anj^des to the matrix, and then no stem is
essential. Furthermore, so many species grow on vegetaljle
deln-is and dead leaves, hence a stem is necessary to push the
pileus into the light. All collectors know how much the
stems are lengtliened beyond their normal proportions when
the mycelium is deeply imbedded in the loose soil, and that
the hymenium is not developed until the pileus is elevated
into the light. In the Gastromycetes the hymenium is not
inferior, so that it is sufficient if the receptacle is just above
the soil, and hence the carpophore is short. The Phcdloichi
are exceptional, as the hymenium is not concealed, but must
be well exposed, in order to mature speedily.
All the remainder of the large fleshy Pungi belong to the
Discomycetes, in which the hymenium is exposed on the upper
surface, and therefore, as might be expected, the carpophore is
often short or absent. The Morels
and Hclvellas are all stipitate, and
the receptacle is like a cap or hood ;
l)ut as they are terrestrial, often grow-
ing in loose soil and amongst debris, a
carpophore long enough to bring the
receptacle into the light is essential. In
this case the substance scarcely differs
from that of the cap, but it is robust,
and the external stratum is not car-
tilaginous. In several genera of ter-
restrial habit tlie form is clavate, with
a carpophore long enough to l)ring the
hymenium through the short grass into the light, just as in
simple club - shaped forms of Clavaria. We need only to
allude to the old genus Pcziza, whatever the modern designation
may be, for all of the species are cup-shaped in form (Fig. 1 '1), and
the liymenium is turned to the light, hence all the carpophore
which is necessary is that wliicli is suflicient for such a purpose.
Fig. 12. — Receptacle or cup
of Pcziza.
28 INTRODUCTION TO THE STUDY OF FUNGI
SO that usually the carpophore is short, and often reduced to
a mere point. Species such as that which grows on the
sclerotium of anemone roots are variable in the length of the
carpophore in proportion to the depth at which the sclerotium
is buried, or of such as grow on acorns or beech-mast lying
on the ground the carpophore is long enough to bring the
hymenium to the light. Pcziza aurantia or Peziza ladia,
growing on naked soil, are fully exposed, and hence are sessile.
Wherever, from its matrix or peculiar habit of growth, a species,
if sessile, could not expose its hymenium to the light under
ordinary circumstances, a carpophore is usually present.
Species which grow beneath the bark of branches, and break
through, have invariably a short carpophore to raise the disc
to the surface.
Some special forms of carpophore are to be found in the
Pyrcnomycetes, where the fructification is capsular, and the
receptacle small and simple. In this case the carpophore is
not, except rarely, that of a single individual, but of a colony
or an agglomeration of individuals, each individual
being represented by the fruit receptacle, the
carpophore being a vegetative branch, developed
from the mycelium, specialised to carry the fruc-
tification, as the conidiophore of a mould is
specialised to carry a great number of conidia. For
example, the pupa of a moth becomes filled with
mycelium, which, in the first instance, developed
^ ,„ , conidia under the form of Isaria farinosa (Fig. 13) ;
Fig. 13. — Isa- "^ \ g / '
riafarinosa finally, a club-shapcd fleshy protuberance called
on pupa of ^ stroma grows from the surface of the pupa in
connection with and continuation of the internal
mycelium. This fleshy stroma is at first only a sterile branch
from the mycelium, like the stem of an Agaric, but ultimately
the whole of its upper surface is covered with an indefinite
number of minute receptacles, which are developed in a
colony at the apex of a carpophore. The insects, M'hether
larvae or pupae, on which these Fungi are developed are
at the time buried in the soil, and the function of the
carpophore is to carry the fructification into the light, so
that sometimes it has to be prolonged several inches before
THE CARPOPHORE
29
Fio. 14.— Clav-
ate stroma of
Cordyceps.
the fructifying suiface is sufficiently raised above tlie soil to
attain its development (Fig. 14). The carpophores vary not only
in length for the same species, according to
circumstances, Init also in form, according to the
species. In some it is simple, and in others
branched, but the receptacles are always densely
accumulated al)out the apices in this genus of
Cordyceps. Hence we recognise again that a
carpoi)hore is a contrivance which is resorted to
in order to bring the fructification into the air
and liglit, and is lengthened or sliortened in con-
formity with that o])ject. In the genus Xylaria
the form of carpophore is similar, but its texture
different. The colour is normally black exter-
nally, white antl corky within, autl it is wholly
tough and hard. The species grow on putrid
wood and rotting leaves. In an allied genus,
Thamnomyces, the carpophore is very long and
thin, often like horse hair, running amongst dead leaves and
vegetable debris.
It is notable how some of the simplest forms of carpophore
are repeated in different groups of Fungi far removed from
each other in structure. This is the
case where the whole Fungus is club-
shaped, as it is in Clavaria jnstillaris,
and again, even as to colour, in Xylaria
invohUa. Others of a smaller size, but
of a like form, will be found in Clavaria
ligula, Zcptoglossum olivaccum ; Xylaria
rhopaloidcs ; Geoglossum, hirsutum, and
Hypocrca ophioglossoides.
No one can doubt, after tracing the
gradations of form in Xylaria, that the
spherical carpophores, not only in
Xylaria, but also in Daldinia, Glaziclla, Sarcoxylon, and the
Sphaeroxylon section of Hypoxylon, are of the same character,
and have a similar purpose to the foregoing (Fig. 15). Possibly
the globose forms may primarily serve to expose the largest
surface of immersed receptacles to the light, rather than elevate
Fig. 15. — Globose stroma of
Hypoxylon.
30 INTRODUCTION TO THE STUDY OF FUNGI
them from darkness into light, as was seen to be the first
function of the long - stemmed carpophores. Everything
indicates in the Pyrenomycetes that there is some necessity
for exposing the fructifying surface to the light ; whether on
carpophores or effused in a stroma, only one stratum of
perithecia is the rule, a double series the very rare exception.
CHArTEK IV
THE KECEI'TACLE
Having descril)ed the mycelium, already designated as the
vegetative Fungus, and which is always present, we passed
to an outgrowth of the mycelium (soiuetimes suppressed),
which, as the carpophore, is destined to support the rcceiitadc.
It is the latter which contains the fructitication, or, in the
ease of naked fruits, supports the fructification itself. It may
be urged that the receptacle is part and parcel of the fructifica-
tion, but it is really no more intimately so than are the
receptacle or the calyx and corolla in flowering plants. This,
however, need not be discussed, as it is only a question of
analogies. What we desire to include under the present
designation is the development, or modification, of the superior
continuation of the carpophore, which encloses or supports the
essentials of fructification. In the absence of any distinct or
evident carpophore, it is still the immediate supporter or
envelope of the fructification, which in that case is sessile upon
the mycelium.
This receptacle may be varial)lc in form, and be known
under different designations, but its function is the same —
that of supporting or enclosing the hymenium, wherever a
hymenium or its analogue is present. This organ will be
represented in some cases by a pileus, in others by a peridium,
an e.xcipulum, a perithecium, a sporangium, or even a
proliferous stratum.
The best-known form, because the largest and most con-
spicuous, is the pileus, which is characteristic of the Hijmnio-
mycetes. It forms the cap in Agarics and the pileus in
Fomcs, Polystictns, etc. This cap in Agarics and allied genera
32 INTRODUCTION TO THE STUDY OF FUNGI
surmouuts the stem or carpophore, and bears on its under sur-
face the hymenium or fructiferous surface. It follows the
same type in the Agaricini, Boleti, some Pohji^orei, and Hydnei.
This type is a convex, or primarily convex, orbicular expansion
of the apex of the stem, with a more or less distinct pellicle on
its upper surface, an intermediate stratum, and its lower sur-
face covered by the hymenium. In this type, then, it is a
hymenophore, as it is in other forms of Hymenomycetcs. This cap
(Fig. 16) not only surmounts, but it is continuous with the stem,
and for the most part conforms to it in texture. In some
cases it passes down in plates on the under surface between
Fig. 16. — Agaric. A, young ; B, mature ; C, section ; p, pileus
s, stipe ; v, volva ; g, lamellae or gills ; a, annulus or ring.
the folds of the membrane which bears the hymenium, and is
the trama. The superior cuticle or pellicle is sometimes so
distinct from the subjacent stratum that it may be stripped
off, but in other species it is so intimately incorporated with
the substratum that it is inseparable. From the margin of
the cap this cuticle is sometimes extended inwards and united
to the stem, covering the young hymenium, and forming a veil.
Externally the cuticle may be quite dry or viscid, or even
covered with a slimy gluten, as in Agaricus mucidus and
Agaricus aeniginosus. In some cases the cuticle is compara-
THE RECEPTACLE 33
tively thick, and consists of an outer and inner layer, the
former breaking up as the cap expands, and adhering in
patches or scales, as in A(jaricus procerus, whilst the inner
silky, fibrous layer is closely adnate to the flesh. Not uncom-
monly the cuticle, without breaking up, is finely striate with
innate silky fibrils, or shining with a satiny lustre. The flesh
beneath the cuticle differs somewhat in different species, in
texture and in comparative thickness, but is always thinnest
about the margin, sometimes scarcely exceeding that of the
cuticle. The cells of the fleshy substance of the pileus in
Agarics are more branched than those of the stipe. They will
form, by anastomosing and crossing each other, a sort of poly-
gonal trellis-work, and in the meshes so formed there is a
second system of larger cells. Corda alludes to them, especially
in the Biissulac, and he says, " These two forms are not always
neatly separated, but pass, as the organ requires, more or less
rapidly, one into the other, or, what is more rare, they are sub-
stituted the one for the other. These two forms of tissue take
part, generally both together, in the structure of the hymenium,
each giving birth, or botli together, to one or many organs of
the hymenium." ^
The lactiferous tubes in such a genus as Ladarius are often
of larger dimensions than the ordinary tissue, and M. de
Seynes protests against their being called " vessels," because, if
the cells are very long, yet it is possible to see that they
are divided " transversely. In Fistulina, which contains an
abundant red juice but more fluid, it is contained in special
varicose and sinuous tubes, like the laticifers, but furnished
with transverse divisions. On approaching the gills the same
series of cells are curved and recurved, showing that the milky
secretion is there more abundant. As to the functions or
import of this juice, that is still an obscure point, for there are
so very many Agarics which do not possess it at all, or if pre-
sent, it is in smaller quantity, and not equally visible. It is
well enough known that in Lactarius the milky juice descends
into the hymenium, for if the gills are cut or bruised, it oozes
out, and hangs suspended in drops. "When dried it is readily
seen to be resinous.
* Corda, Iconcs FioKjuriDn.
3
)^
34 INTRODUCTION TO THE STUDY OF FUNGI
Xo oue has yet paid much uttentiun to the coloration of
the pileus in Agaricini, which is subject to much variation,
due in part to external circumstances, as was pointed out by
]\I. de Seynes. Albinism is one of the variations which he
observed in well-recognised species. It occurs in Hyyropliorus
calyptraeformis, Amanitoims vaginata, Russula fragilis, and
some others. " By the side of this fact there is another quite
opposite, the greater intensity of coloration, according to the
temperature." For instance, he found during the winter
Tricholoma nuda, according as the temperature falls, of a dark
violet, almost black, or a deep brown. When spring arrives
it is found almost white, shaded with lilac or fawn colour.
Tricholoma terreus and Collyhia dryophilus will present the
same phenomena. He found also, in a cold December, Volvaria
media of which the pileus was almost black ; ordinarily it is
nearly white. Upon microscopical examination he found that
there was no new production of cellular elements, but simply
a greater agglomeration of pigmentary granules. He also
indicates that he has often been struck with the deep colora-
tion of Armillaria mellea and Hyplwlowxi siQAateritius, which
were seen by hundreds during a low temperature (41° to 42°
Fahr.), the aspect of which differed very much from the same
species found in the woods during the fine days of autumn.^
The same writer adds that, from numerous observations, he is
certain that, although the cold has an influence upon the
intensity of coloration amongst Agarics, it does not follow that
in the middle of winter specimens may not be found of the
normal colour, either by being shaded, or in proximity to heat.
The effects of external circumstances on the variation of the
fleshy Fungi deserves more serious attention.
In some few instances the fleshy stratum is almost obsolete
over the entire cap, as in Hiatula, some species of Coprinus,
Bolhitius, etc., and in some exotic species of Marasmius. In
such cases the cap is so thin — like a membrane — that when
moist the gills may be seen through the substance. In
Boletus the cuticle is sometimes distinctly velvety, and the
flesh is comparatively thicker than in Agarics. In Polyporus,
Fomes, Polystidus, Hydnum, etc., the entire substance is more
^ Grevilka, vol. ii. p. 12.
THE RECEPTACLE
35
woody, contains less moisture, and consequently dries with but
little shrinking or change of loriii.
As the carpophore is souietinics obsolete in the ILjmcno-
mycetcs, so also is the receptacle or pileus reduced to a simple
stratum, which intervenes between the mycelium and the
spore- bearing suriace. These are undoubtedly rudimentary
forms, but they are very numerous, sometimes constituting
entire genera, as in Poria, Co7iiophora, Corticium, etc., besides
numerous species in other genera. For the most part a thin
tibrous stratum, difl'erentiated from the fibres of the mycelium,
forms, and supports the hymenium. Possibly the old genus
Ozonium consists entirely of these suppressed pilei, which never
form a hymenium. The supporting stratum is very peculiar
in Astcrosfroma, where the hyphae are stellate, and in Thele-
phora pcdiccUata they assume a dendritic form. It is not
uncommon to find specimens of Corticium in which the
hymenium is only in patches, or, in some cases, never formed at
all, so that the whole Fungus remains in the vegetative stage,
that is to say, mycelium, and a sterile fibrous stratum to
represent suppressed carpophore and atrophied receptacle.
The second type is deficient in any appreciable carpophore
or stem, and con-
sists of a pileus of
a semicircular out-
line, attached at its
base to the matrix
and its own my-
celium (Fig. 17).
In these also there
is a superior stra-
tum, which may be
thicker than in the
preceding, an inter-
mediate substance,
and an inferior hy-
menium. Tlie upper
stratum in Foli/purus and Fisitdina is hardly distinct from
the intermediate ; but in Fames it usually forms a firm
hard crust, very hard and Ikhmv in Fnurs ai/s(ralis and
Flo. v.— Fust III Ilia hepatici, .sessile iiileiis.
36 INTRODUCTION TO THE STUDY OF FUNGI
Fomes cornu-hovis, Lut smooth, and mostly shining, often
laccate, as if varnished. According to Wettstein,^ this is
due to the secretion of resin which oozes from pecviliar
hyphae and flows over the surface of the pileus. The exterior
of the pileus exhibits deep concentrated channels, which mark
the annual additions at the circumference. The substance is
often very thick and fibrous, the fibres radiating in every direction
from the base. They may continue growing by the addition
of external zones for many years, always the oldest posteriorly.
From these Polystictus differs in being much thinner, and the
cuticle is fibrous, hairy, woolly, or strigose, and concentrically
zoned. The substance is dry, tough, and leathery, usually
flexible. In so far as these features are concerned, Stereum not
only resembles Polystictus in appearance, but also in texture,
and so does Hymenochacte ; whilst Hexagona differs more in the
hymenium than in anytliing else. These, therefore, may be
accepted as representing the two forms of the sessile receptacle
in Hymenomycetal Fungi — the woody by Fomes, to which might
be added Daedalea, and the coriaceous by Polystictus, and the
others above named.
The next form of receptacle to be adduced is the periclium,
which completely encloses the reproductive organs, and may
also be supported on a distinct carpophore, or it may be sessile
on the mycelium, or invested by it, as in some subterranean
species. The Gastromycetes furnish this kind of receptacle,
which is very often double, typically
globose, the outer coat or exoperidium
being a continuation of the cortex
uf the carpophore when the latter
exists. The internal cavity is filled
with the reproductive bodies, which
are only liberated by the rupture of
the coat of the peridium. The inner
Fig. 18.— Lycoperdo7i, peridium coat or endoperidium is often thin and
and section. membranaceous, and may either be
wholly separated from the outer or adnate therewith (Fig. 18).
A prolongation from the carpophore sometimes protrudes into the
central cavity in the form of a columella. The outer coat or
1 Verhand. Zool. Bot. Gescll., Wien, xxxv. (1886), p. 29.
THE RECEPTACLE
37
exoperidium may be tough and leathery, and in Geastcr it sphts
downwards from the apex into several triangular lobes. In
Bovifitd it is fragile and evanescent. In Lycopcrdon it breaks
uj) into gruimlcs, warts, or spines, wliich adhere for some time
to the inner and persistent peridium. In Polysaccnm and
Scleroderma the periderm is not differentiated into two coats,
but in the latter the exterior cracks into warts or frustules.
In this form the receptacle is an entirely closed envelope, in
which the fructification is completely concealed until it is
quite mature, and then it either opens with a small orifice or
is irregularly ruptured. Hence the light is not essential to
the perfection of the fruit, and the peridium might almost as
well remain in the soil, which it has a tendency to do in some
species of Scleroderma, and does completely in the Hyj)0(jaei.
In the majority of species the substance of the peridium is
tough and leathery, and so persistent that it often remains
behind long after dehiscence and the dispersal of the spores.
A third form of receptacle is the excipulum or cup-shaped
receptacle, which, although often closed when young, is soon
expanded so as to expose the disc or
hymenium to the full light ; in fact
they are heliotropic, for they turn the
disc as much as possible towards the
sun. The type of this form is to be
found in the old genus Peziza, now split
up into many genera, but the form and
structure of the excipulum is the same
throughout (Fig. 19). The external
stratum of cells does not form a
separate cuticle, but is continuous wuth
the subjacent cells, and usually consists 1"'ig.19.-
of smaller or elongated cells, w-liich
may be coloured, and either mixed with or prolonged into
hairs, usually most strongly developed about the margin of the
excipulum. Within the cuticular layer lie the subhymenial
cells, on which rests the hymenium or fruit-bearing surface.
The attributes of this form, therefore, are a cup -shaped
receptacle, with the mouth turned to the light, and composed
of an external and internal series of cells, the latter su]>porting
section aiul ascus.
38 INTRODUCTION TO THE STUDY OF FUNGI
a compact hymenium. The substance of the cup and entire
Fungus is most often soft and fleshy, and therefore they are
more or less hygrometric, closing when dry and expanding
when moist. Whether naked or clothed with hairs, the
exterior is usually dull and sombre-coloured, so as scarcely to
be distinguished from the matrix on which they grow. A
similarly-shaped receptacle is to be found in CypheUa amongst
the Hymenoraycetal Fungi, in Cyathus and Crucihuluon
amongst the Gastromycetes, and in Aecidium amongst the
Uredines ; besides partial resemblances amongst the Siihaerop-
sidcae, and a few of the compact moulds, such as Volutclla and
Chactostroma. The closing of the margins of the cups, and
the long marginal hairs which in dry weather cover the
hymenium, serve as a protection against evaporation, to which,
from their fleshy substance and exposure, they are peculiarly
liable. Some minute species, which are erumpent, retract
themselves within shelter of the cuticle as they lose their
moisture, and can only be distinguished with difficulty.
Another form of receptacle is represented by thousands of
species, and that is the ijcrithccnim. This is a minute form,
#^^ seldom exceeding a rape-seed in size,
yT ;= ^^V ^^^^^ usually very much less. It may
C-'^/:!^! be characterised as a globose flask
\&^iii^^g^ with a very short neck, but this
Fio. 20.— Peritiieciuiii witii iomi is variously modified (Fig. 20).
'''''^^*'°"- Sometimes it is seated directly upon
its mycelium, and sometimes simk in a stroma which
arises from a mycelium. These receptacles may be superficial,
on the surface of the matrix, or either wholly or partially
immersed. The globose form may be flattened at the
base, and the neck be very much elongated, like a horn, or
absent altogether ; and. they may grow singly or in company.
In some genera the perithecium is soft and fleshy, and then
pale or brightly coloured, as in Nedria. It may be thin and
membranaceous, as in Sphaerella ; or it may be coriaceous and
tough, as in Botryosphaeria ; or carbonaceous and brittle, as in
Rosellinia. Externally it may either be smooth and shining,
or mealy, or warted, or bristly, or woolly. The apex is always
closed, except for a minute pore or ostiolum, and this is only
THE RECEPTACLE 39
absent in a lew genera. In one group only, the Lophiostomaceac,
the mouth is broad and compressed. The fructification there-
fore is always enclosed, as it is in the case of a peridium, but
this is their only point of agreement. Whatever the ibrra of
the carpophore, the perithecia are always crowded together on
the upper portion, and even when the carpophore is globose it
is flattened at the base, and as the perithecia approach the
base they are smaller, less numerous, and often imperfect. As
already stated, the perithecia are always peripherical, and in a
single series ; but if, in rare cases, there is a second series, the
necks of the perithecia are elongated so as to reach the surface
of the stroma. When a great number of perithecia are collected
together, and immersed in the matrix, a kind of stroma is formed
from the matrix, as in EiUypa ; but when on the surface, a
regular stroma is formed upon the mycelium, which is fleshy
in Hi/pocrca and carbonaceous in Hypoxylon, and the perithecia
are closely packed and immersed in the stroma, which is per-
forated by the ostiola. The densely
aggregated perithecia may sometimes
be fused together so as to resemble
an effused stroma. There are some
genera in which the perithecia seem |v
to be obsolete, or only formed from
the stroma, in which the perithecia
appear only as cells, as in Phyllaclwra ;
but the walls of the perithecia are
possibly fused with the stroma and M^ ^ ' ' ^ ^^^ (
not wholly absent. The character of
the fructification may be entirely fig. 21. -Receptacle of sporo-
different, while that of the perithecia "'ega in Hysteriacme, with
. m • ascus.
remanis the same. The same peri-
thecium, to all external appearances, may belong either to
the Pyrenomycctcs or tlie Sphacropsidcac. It may be a Diplodia
or a Plaeospora.
It may be necessary to refer incidentally to a modification
which is almost intermediate between an excipulum and a peri-
tliecium, which is prevalent in the comparatively small family,
the Hystcriaccac. Here the excipulum closes from two opposite
sides, leaving: a slit down the centre. When well moi-stened
40 INTRODUCTION TO THE STUDY OF FUNGI
these excipuli expand so as to be nearly cup-shaped, and a com-
pact disc is exposed (Fig. 21). This corresponds to the cups
of the Discomycctes, but the substance is leathery or carbonaceous,
and when closed the receptacles approach the Zophiostomaceae,
which is sphaeriaceous. Nevertheless the most evident affinity
is with an excipulum, in the broad expanding mouth and the
definite disc, very evident in Tryhlidiella mfula, and its
relationship with the Discomycctes would be through Phacidiacei.
Other modifications need not be particularised, since these
principal types will be sufficient to indicate the character
of the receptacle in the majority of Fungi. We have the
vegetative system represented in the mycelium, which sends up
erect or compound branches in the form of a carpophore, for
the support of the receptacle in all cases where the receptacle
is stipitate, or, where the receptacle is wanting, then to support
the naked fruit.
CHAPTKU V
THE FRUCTIFICATION
The contents of the various forms of receptacle already
described, and those forms of fructification which are capable
of being produced nukcil, withoiiL a receptacle, next demand
attention. The best-known to the general public, and there-
fore the most interesting, are those large and conspicuous Fungi
which pass under the name of Mushrooms or Agarics, and
the woody Polypores, with the spore-bearing surface on the
under side. In more scientific language, these are the
Hymenomycetal Fungi, and so called because the hymenium
or fructifying surftice is naked, and produces naked spores.
From what has preceded it will be remembered that a fleshy
or woody pileus or receptacle, sometimes with, and sometimes
without a stem, is the supporter of this kind of fructification.
To the eye it presents the appearance of a continuous surface
extending over plates or gills in the Agaricini, lining the
interior of parallel tubes in the Polyporci, covering the outer
surface of teeth or spines in the Hydnci, disposed over a nearly
even plane in the Thdcphorci, effused over an erect, simple, or
branched carpophore, but without receptacle in the Clavarici,
and immersed in a gelatinous stratum in the Trcmcllinci.
Under all these modifications the primary elements of the
hymenium are the same, or chiefiy so ; that is to say, there are
one, two, or three kinds of elongated cells, packed side by side
and called respectively hasidia, cystidia, and sterile cells.
Only the first kind are fertile, and bear at the apex four
spores, surmounted on short slender spicules ; the cystidia ^ are
' The usual interpretation of the function of cystidia is, that they arc simply
mechanical contrivances projectinj,' fruni the surface of the hjTiionium, and thus
keeping the gills or lamellae apart.
42
INTRODUCTION TO THE STUDY OF FUNGI
usually present, mixed with the basidia but rather larger, and
the sterile cells are smaller and almost of the nature of, or
analogous to, paraphyses (Fig. 22). De Seynes regards all
three forms as modifications of the same organ, -i.e. the
basidia, of which the spore-bearing
are the fertile basidia, the cystidia
are hypertrophied basidia, and the
sterile cells atrophied basidia. All
these cells are continuations and
terminations of the tissues of the
receptacle, sometimes with three or
four subspherical cells intervening.
The basidia are elongated clavate
cells, or sporophores, filled with a
granular fluid, surmounted by four
short slender tubes, or spicules,
each of which expands at the apex
and becomes a spore ; into which
FiQ. 22.-Ba,sidia (6) and cystidia t^g contents of the basidium pass,
leaving the basidium empty, so
are completed it collapses and
shrivels, then falls away. The spores thus formed by
budding or gemmation, as far as known at present, are
asexual and only gemmae. Many efforts have been made
to prove them otherwise, but none of these have been con-
firmed. The spores themselves are unicellular (except in the
Tremdlini), and may be colourless or coloured. Modifications
seem to take place in the cystidia, in different genera, inde-
pendent of any difference in size. In the genus Peniophora
they evidently become encrusted with lime and granular, so "as
to present quite a distinct appearance ; in this condition they
have been called "metuloids." In Hymenochaete and some
species of Fomes the normal cystidia are replaced by rigid
coloured setae, which may be modifications of cystidia.
Corda regarded these peculiar cells as representatives of
male organs, and called them antheridia ; and a similar
interpretation has been given to their functions by Worth-
ington Smith. Most mycologists coincide in the opinion
that a sexual apparatus has not yet been discovered in the
(c) of Agaricus ; {a) jiaraphyses.
that when its duties
t
THE FRUCTIFICATION 43
Hymenomycdcs, and tliat it is scarcely probable that sexuality
exists.
The contents of the closed receptacle of the second type
enumerated in the preceding chapter — the peridium — differ in
some features from the foregoing, although they accord in the
spores being i)roduced upon basidia. In this case there is no
effused and exposed hymenium, but the interior of the peridium
is occupied and filled with the gleba, or entire reproductive
mass, which is at first homogeneous. Afterwards minute rifts
are to be observed in tlie gleba, which increase in size, and
ultimately form a labyrinth of cavities. The walls of these
cavities are composed of hyphae, and the inner face is converted
into a hymenium, the basidia of which are the
terminations of the hyphae of the walls. These
basidia are more variable than in the Hymeno-
mycetes, and the number of spores not so
constant (Fig. 23) : in the Eymcnogastrcac from
one to four ; in the Phalloidcac from four to
eight ; in Bovista and Lycojicrdon four and _^ ^^^
terminal ; but in TuJostoma four and lateral. "^
Hence it will be observed that the spores are Fui. 23.— Basi-
, , , . ,. ... . , . . , (liuiu ami spores
produced on basidia, within special cavities and oiLi/coperdon.
lining the walls ; but the entire mass, or
gleba, is contained within a closed peridium, wliicli is not
ruptured until the spores are mature. When this takes
place the entire gleba will be found in most cases (exclusive
of the Phalloidcac and the Nidulariacci) to be converted into a
finely pulverulent mass, mixed with fine fibres. The powdery
mass consists of the ripe spores, and the fine fibres are the
remains of the internal hyphae, now called the capillitiuvi.
In the Hymcnogastreac, which are the subterranean Gastro-
mycetcs, the walls of the cavities are more persistent, and
therefore there is no capillitium, and the peridium is not
ruptured when mature. One or two features of the spores are
in contrast with those of the Hymcnomycctcs, that they are for
the most part coloured, often warted, or spinulose, whereas the
majority are also globose in form, except in the subterranean
species. The basidia can be seen only whilst the glelia is
young, for before the spores are mature they are dissolved away
44
IXTRODUCTION TO THE STUDY OF FUNGI
The Elapliomyceteae correspond to the Hymenogastreae in
being subterranean, similar in form, and alike enclosed within
an indehiscent peridium ; but they differ in the spores being
enclosed within asci, instead of being produced on basidia, and
in this respect are allied to the Pyrenomycdes.
In so far as their final, and reproductive, stage is concerned,
the Myxomycetes resemble the Gastromycetes ; they are some-
times stipitate, possess a distinct peridivim, in which the spores
are enclosed until maturity, and the latter are mostly coloured,
globose, sometimes rough, mixed with the threads of a
capillitium. On the other hand, the early or vegetative stage
is so different, that the ancient notion of their affinity must
be abandoned, although they are entitled to mention in this
place as Fungi which produce their fructification enclosed
within a peridium. Notwithstanding this, thei*e are those who
regard it as heresy to mention the Myxomycetes on the same
page as the Gastromycdcs.
The fructification, which is produced witliin an open cup-
is of more than one kind, but the most
important is that of
shaped excipuh
6—/
Fig. 24. — Section of hyiuenium iu Peziza.
the Discomycetes, in
which the spores are
ascomycetous — that is
to say, they are pro-
duced within asci. In
describing the recep-
tacles it was stated that
the fructiferous surface
was a compact stratum
or hymenium, which
overspread the interior of the expanded receptacle. In this
instance the fruit-bearing surface is superior, and soon fully
exposed to the light. It is plane or slightly convex when moist,
depressed and concave when dry, from the contraction of the
receptacle,and often brightly coloured (Fig. 24). The hymenium,
or disc, is composed of elongated cylindrical or clavate cells,
which are formed of a delicate hyaline membrane, splitting at
the apex, or opening with an operculum. These cylindrical
cells are closely packed side l:)y side and constitute the asci,
THE FRUCTIFICATION
45
which are generally mixed with slender threads termed
})arap]iy.ses, and these two bodies together compose the
hymcnium. Each ascus when mature encloses
eight, more rarely Iniir, or sixteen, sporidia,^
often globose or cIlipLical, and uncoloured.
The paraphyses may or may not be abortive
asci, the apex may be attenuated, or it may
be thickened in various ways, and in the latter
case often replete with a coloured protoplasm
which imparts the colour to the disc. We
arc only desirous of explaining the normal
form and structure, without regard to the
minor differences which enter into the char-
acters of the different genera. We may term
the above the Pezizaeform type, which is repre-
sented by some two thousand species.
The question as to the nature and mode \ I u^
"of derivation of the ascospores is at present
scarcely more than problematical. No male
organs have yet been found in consort with
the thecae, and there is no reason to assume
that ascospores are the result of sexual union.
The only evidence is that offered of conjuga-
tion in the earliest stage of the receptacle, by
means of which the entire cup and its contents fig
is the sequence of a sexual act. De Bary,
Woronin, and Tulasne are the observers on
whom this jihenomcnon rests. It is to the effect that
25. — Ascus
with sporidia and
parajibyses.
^ It will be well to indicate here the names which are api)lied by tlie best
authorities to the spore in its relation to the different families of Fungi. Although
these names are somewhat arbitrary, the student will find them employed almost
universally in systematic books.
H-porc, without asci, in perfect Fungi, such as the Basidioraycetes.
Sporklium, enclosed in asci, in perfect Fungi, such as the Ascomycetes.
Sporulc, without asci, in imi)erfect Fungi, enclosed in perithceia, such as the
Sphaerojisideae.
ConuHum, without asci, in inijierfect Fungi without a perithccium, such a.s the
moulds, or Ilyphomycetes, and Melancuniaceac.
In Uredines and Phycomycetcae special modifications arc employed which
have reference to their development.
Spermatia, stijlosporcs, and clinosporcs are merged in sporulc.
46
INTRODUCTION TO THE STUDY OF FUNGI
specialised branch of the mycelium, which is thicker than the
rest, is always to be found in close proximity to certain fila-
ments, the short curved branches of which rest their ex-
tremities on the turgid branch first named, and which is termed
the vermiform body or scolecite. At the point where these
two organs meet there is a circular perforation, and one of the
cells appears to transfer to the other a portion of its contents.
The scolecite is stated to be the rudiment of the fertile cup, to
become septate, and to
Fig. 26.— Scolecit(
Bary.
collect around itself
other filaments which
grow and develop into
a perfect cvip (Fig. 26).
This process is reported
to have taken place in
Ascoholus furfuraceus
and in Pyroncma om-
phcdodes, and it is as-
sumed to be general
throughout all the fleshy
Discomycetes. It is
very evident that such
a conclusion cannot be
accepted, so that the
above remarks must
illustrating the nearest
observers have obtained
be received as historical, and as
approach which some of the best
towards establishing sexuality in the higher Fungi, such as
Hymenomycetes and Ascomycetes.
The Gasteromycetal Fungi also include a small family in
which the receptacle assumes a cup shape, especially in the
genera Cyathus and Crucibulnm ; but here again there is great
divergence in the character of the fructification. In the latter
genera the cups are at first covered with a tympanum or
membranaceous veil, and when this is ruptured, are seen to
contain a small number of lentil -shaped bodies, which are
attached by a slender elastic cord to the inner surface of the
cup. These are the peridiola which enclose spores produced
upon basidia within the firm interior (Fig. 27).
rilE FRUCTIFICATION
47
The excipulum, or cup-shaped receptacle, is also the form
whicli is assumed in some genera of the Sphacroimdeae in
which the external resemblance is again to tliat of the Dis-
comycetes. The cups are sessile, often erunipent on the stems
of herbaceous plants, and externally smooth or covered witli
bristles — for example, in Excipula, Discella, and U^^helis, and
many others. The spores are produced on short sporophores
which grow side by side from the inner surface of the recep-
tacle, as the asci are produced in Przizrt, Init without forming
a compact hymenium.
From these brief notices it is evident that fructification
of widely diverse types may be found to be produced within
receptacles which are
open above and
therefore cup-shaped.
The most prevalent
form is the asci-
gerous, in which the
sporidia are produced
in asci, packed close-
ly side by side and
forming a compact
disc, as in the Disco-
mycetes. An ana-
logous genus is found
in Hymenomycetes,
in which the spores
are produced upon
basidia, as they are
in Corticium.
Amongst the Gastro-
mycetes the cup-
shaped receptacles
enclose lenticular
peridiola which con-
tain basidiospores.
Fig, 27. — Vi-ucibulum vuhja
C'hioii.
And in Sphacropsideae certain genera
with an open, cup -shaped receptacle produce naked spores,
or conidia, ujiou short sporopliores. To these might be
added also, I'lnm ihe Uredines, the " cluster-cups " of ^<«(/«<?«,
48
INTRODUCTION TO THE STUDY OF FUNGI
Fig. 28. — PseTuloperidia of
Accidium.
which contain globose spores developed in chains (Fig. 28);
and possibly the Hystcriaccae, in which the receptacles are
often closed and elongated, but in some instances are gaping
in moist weather, so as to expose a compact disc of parallel asci,
combined witli paraphyses, after the manner of Pcziza. This
family falls intermediate between
those with a cup-shaped receptacle
and the following group, in which
the receptacle is a closed perithecium,
but inclined more towards the
former than the latter. It will be
observed, in examining more closely
into the morphology of these groups,
that the preponderance of species which possess a cup-shaped
receptacle are sessile on the mycelium, without the inter-
vention of a carpophore, in the sense to which we have here
limited that term.
We pass now to the representatives of that large mass of
Fungi in which the receptacle is wholly closed, with the ex-
ception of a terminal pore, as included under the general term
perithecium (Fig. 29). In all essential features the fructifica-
tion is the same under the two forms in which it presents
itself, the ascigerous and the stylosporous. The former is that
of the Pyrenomycetes and consists of cylindrical sacs, or asci,
with linear paraphyses intermixed and packed closely side
by side. In this respect the fructification resembles that of
the Discomycetes, only that the disc, or upper surface, is not
exposed, and hence not compacted.
The asci contain four or eight or
sometimes an indefinite number of
sporidia, which are either hyaline
or coloured, and simple or variedly
septate. The summit of the ascus
is normally imperforate. Exceptional instances could be
cited in which the ascus encloses but one or two sporidia, or
where no paraphyses can be detected. Up to the present no
clue has been found to the fertilisation of the sporidia,
whether by the fertilisation of the entire receptacle in its
earliest stage or that of the sporidia. In the case of some of
Fk;. 29. — Peritliueium and
section.
THE FRUCTIFICATION 49
the Perispoi'iacei, after the contact of two hyphae, a process
is evolved from each, which ultimately develops respectively
into oocyst and antheridium. De Bary from this traced the
history of a conceptacle in Erysij)he to its completion ; and
expressed the opinion that the perithecia and asci of many of
the Ascomycctes originated, and were perfected, in the same
manner.
Other perithecia, with the same external form, Iialiit, and
texture, but with stylosporous fructification, are included
systematically in the Sphacropsidcac. There are no asci and no
paraphyses present, but the receptacles enclose an indefinite
number of sporules, which are generated singly at the apices
of very short slender threads. These some have called hasidia,
but that is a misnomer, since in the Basidiomycetes it is
accepted with a different interpretation. In the Sphacropsideac
the delicate supporters of the sporules are simply sporo-
phores. The sporules themselves do not differ materially
in form, size, and appearance from the sporidia which are
generated in asci, and there is a suspicion that some of them
are genetically connected, but in what manner has never been
ascertained.
In some genera the perithecia are nearly obsolete where
the species are immersed, or else they are so fused with the
matrix as not to be distinguishable, or they
may be quite spurious, so that cavities in
the matrix perform all the functions of
perithecia (Fig. 30). The latter condition ^^^
prevails in the Melanconieae, where the yio 30 —Spurious
conidia are produced, as in the Sphaeropsidcac, perithecium of Mel-
on short conidiophores within definite cavities
or cells, the walls of which are differentiated from the matrix.
There is a basal cushion or spore -bed which is formed from
the mycelium, and this spore -bed originates the conidia.
Saccardo and the majority of systematists apply the term
"conidia" to the spore-like vesicles of the Melanconieae as
well as to the H uphomyccteae or moulds.
A small but interesting group of Fungi, having the habit
and appearance of moulds, difter from them very materially
in possessing a receptacle at the apex of the carpophore, which
4
50 INTRODUCTION TO THE STUDY OF FUNGI
is technically a sporangium. This apical receptacle is delicate,
membranaceous, and often transparent, enclosing a number of
small spores or zoospores. The mucors develop only spores
within the sporangia, but the Peronosporaceae sometimes pro-
duce only passive conidia at the ends of the branchlets, in
which case they belong to the naked -spored group, to be
alluded to shortly, in consort with the moulds ; at other times
sporangia are produced, or analogous organs, the contents of
which are differentiated and escape from the vesicle as zoo-
spores. These are the only instances known in Fungi in
wliich the hyphae support a delicate inflated vesicle containing
spore-bodies capable of germination. This is not the only
feature in which they differ materially from the moulds which
they superficially resemble ; but in the mycelium they differ
in the hyphae being for the most part without septa, and
further, in the power which they possess in forming special
branches on the mycelium, which conjugate and form zygospores.
These latter are able to pass through a period of rest, as rest-
ing spores, and form a second form of fruit for the mucors ; as
oospores are produced on the mycelium of the Peronosporaceae
by a similar sexual act, and are a second form of fruit for that
family, capable of passing the winter as resting spores.
Having disposed of the several forms of fructification
which are to be found developed within special envelopes, or
receptacles, according to their kind, we have still to deal with
those less common forms in which the fructification is naked
on the carpophores, and destitute of any kind of receptacle.
The most typical of this system of fructification is that
exhibited by the moulds, or Hyphomycetes, in which the
carpophore or conidiophore is either simple or branched at the
apex, and the spore-bodies, or conidia, are produced singly or
in clusters at the apex of the conidiophore, or at the tips of
its branches, or diffused in any other manner upon its surface.
It may at once be admitted that in these nakedly-disposed
spore -bodies we are unable to recognise any symptoms of
sexuality, and hence they can scarcely come under any other
designation than that of reproductive buds, or if that term
is objected to, then as asexual spores. If we accept as an
example Verticillium agaricinum, the carpophore is an erect,
THE FRUCTIFICATION
Fio. 31. — Conidiophore
of I'enicUliuvi.
septate hypha, proceeding as an assurgent branch from tlie
mycelium. In its upper })ortion it produces, at intervals,
branches one, two, or three, at the same level, and these again
produce branchlets in whorls of three. Each branchlet is
surmounted by an ovate conidium, or sometimes two or even
three together. This, therefore, is a mould,
with somewhat of a dendroid habit, with
verticellate branches and branchlets pro- %\
ducing terminal naked conidia. Take as
another example the common mould, Pcni- ^
cilliam glaucum, or any other Penicillium
(Fig. 31): the erect carpophore divides at the
apex into a cluster of short branches, and
each branch is terminated, not by a single
conidium, but a series of conidia, attached
end to end in a chain, each conidium falling
away successively as it attains maturity.
In other genera the carpophore is very
short and unbranched, either terminating in
a single spore or in a chain of spores, but
the principle is the same — that of naked conidia borne direct
by the carpophore, without receptacle.
We can only recognise in the Uredincs
a modification of the same principle, which
is most strongly manifest in Phragmidium
(Fig. 32). The teleutospores consist of an
elongated simple carpophore, surmounted by
a multiseptate spore-body, and there is no
receptacle. It is similarly manifest in
Uromyccs and Puccinia, for the teleutospores
have a distinct pedicel, which bears the
fruit and is the carpophore. In some
other genera it is less manifest, whilst in
Aecidmm and Racsfelia the distinct recep-
tacle is of the cup -shaped series, open
FiQ. 32. — Teleutospores qVjqvp
of Phragmidium.
In all the instances given in this
chapter we have denominated the bearer of the fructification
by the general term of " carpophore." This is by no means
52 INTRODUCTION TO THE STUDY OF FUNGI
intended to ignore the fact that even as spore-bodies, having
the function of spores, may be designated specifically as spore,
sporidium, sporule, or conidium, so also the carpophore may,
in some cases, more fitly be called a basidium, sporophore, or
conidiophore.
CHAriEE VI
FERTILISATION
The methods by which fertilisation is aceoiuplished in many
forms of Fungi remain still as great a mystery as ever, and
they have only in a comparatively few instances been demon-
strated for certainty. It is not so very long since that all the
lower cryptogamia, at the least, were supposed to be repro-
duced asexually; but this is known to have been too hasty a
conclusion, for the algae present many remarkable instances of
sexual reproduction. In Fungi the examples have been more
isolated, and in some cases still require more certain confirma-
tion, so that it must be confessed, when such an immense
number of species are taken into account, the instances in
which sexual reproduction has been determined are exception-
ally few. One-fourth of the total number of described species
consists of those which are classed as incomplete Fungi, and
hence out of consideration ; whilst another one-fourth consists
of the Hymenomycetal Fungi, and if these are to be excluded,
then one-half are at once to be declared asexual. To these we
shall be compelled to add the Ascomycetes, as without established
evidence. In fact, it is doubtful whether one-fortieth part of
the total number of species can be characterised as possessing
sexuality. Hence the opinion is now very general that sexu-
ality is entirely wanting in all the higher forms of Fungi, and
is only to be found in small families. In this respect Fungi,
as a group, are in strong contrast to Algae.
All previous efforts to establish sexuality in the Hymeno-
mycetes having failed, Worthington Smith endeavoured again
in 1875 to prove it from the Agaricini, and his observations
were made on a minute species of Coprinus. He says that at
54 INTRODUCTION TO THE STUDY OF FUNGI
first the Fungus is composed wholly of simple cells ; no differ-
entiation is seen in infancy when the gills are first formed,
but the basidia and cystidia come into existence only simul-
taneously when the plants approach maturity. This differentia-
tion he distinctly regards as sexual, the basidia representing the
female and the cystidia the male organs. When the contents
of the basidia and the cystidia are interchanged, he says, the
result is a return to another series of cells, which go to form
a new plant. The cystidia are more sparingly produced, and
at first cannot be distinguished from the basidia, though fre-
quently larger, commonly granular within, and sometimes
crowned with granules, but sometimes with four spicules.
Subject to moisture, these spicule-crowned cystidia germinate
at the four points, and produce long threads, which bear at
their tips the granules which are so frequent in typical
cystidia. The " granules " at first are not capable of move-
ment, but they are in reality spermatozoids possessed of a
fecundative power, and after the lapse of a couple of hours
begin to revolve, and ultimately swim about with great
rapidity. These spermatozoids attach themselves to the spores,
pierce the coat, and discharge their contents into the substance
of the spore. From twenty -four to forty -eight hours after
this the spore discharges a cell, which soon becomes free, and
this is the first cell of a new plant, which rapidly produces
others of a like nature. At first the spermatozoids are per-
fectly spherical, when they merely oscillate, then they revolve
slowly, and, as time goes on, a single turn of a spiral makes
itself visible, and the bodies whirl round with great rapidity.
At intervals the motion entirely ceases, and then, after a short
lapse of time, the gyration is again continued. Judging from
the presence of the eddy round these bodies whilst whirling,
they are possibly provided with cilia, but from the extreme
minuteness of the bodies themselves it is not easy to demon-
strate their presence. The whirling of the spermatozoids is
so strong that when they attach themselves to the spores they
twist them round, after the manner of the revolving oosphere
in Fucus. It is also stated that in many cases the cystidia
fall out from the hymenium, and in company with the spores,
and that it is upon the moist eartli that fertilisation is gener-
FERTILISA riON
55
ally carried out. The last observation, if veritied, is rather
strange, as the spores, when fallen, must be regarded as fully
matured ; it seems to be rather an anomaly that a mature
fruit should be fertilised, rather than when in an immature
condition. This much, then, has been related by Smith in a
very circumstantial manner, and from it he argues that in the
Agaricini the cystidia produce the spermatozoids, by means of
which the spores are fertilised, either upon the hymenium, or
after they have fallen to the ground. During twenty years
we have not heard that his observations have been confirmed,
or that the question has been set at re.st.
Long before the above investigations Oersted claimed to
have discovered a sort of conjugation in the filaments of the
Fig. 33. — Developmeut of sporocarp in Podosphaera. After De Bary.
mycelium of Agarics, but this is now regarded as an error of
observation.
In 1872 C. H. Peck supposed that he had found in a
species of Agaricus " spores produced in globose asci, borne on
a thick, tapering, penetrating peduncle, twelve or more spores
in the ascus." This again was doubtless a faulty observation,
for other mycologists failed to find the asci on the gills of the
specimens determined and furnished by the original observer.
M. de Seynes subsequently attributed the assumed asci to
cystidia, and the supposed sporidia to external and internal
granules. Hence it may be affirmed that none of the sup-
posed processes of fertilisation in Basidiomycetes have been
confirmed, and ur.til that is done they must be regarded as
asexual.
Another one-fourth of the total number of species of Fungi
56
INTRODUCTION TO THE STUDY OF FUNGI
is occupied by the Pyrenomycetes, in which the sporidia are
developed in asci ; but, so far as we are aware, it is only in the
family Perisporiaceae that sexual fertilisation has been sug-
gested— i.e. in Erysiphc and Eurotium, by De Bary and sup-
ported by Tulasne. In Erysiplu lamprocarpa the perithecia are
produced where two filaments of the mycelium cross each other
(Fig. 33). They swell slightly, and each emits a process resem-
bling a branch. That from the lower filament soon becomes
Fig. 34. — Development of Eurotium repens. After De Bary.
oval, and is constricted by a septum from the hypha, becoming
a distinct cell, which De Bary calls " oocyst." The process
from the upper filament adheres closely to this cell, and
elongates into a cylindrical tube, which terminates obtusely at
the apex of the cell. This also is divided by a septum from
the parent hypha near the base, and another towards the
apex cuts off a short terminal cell, which is supposed to be the
antheridium. After this eight or nine new tubes spring up
around the base of the oocyst and closely applied to it, which
gradually develop into the wall of the perithecium. In the
FERTILISATION 57
meantime the contpiits of the cell, or oocyst, are being differen-
tiated into an inner wall and the contained ascus. The y-\\-
thecium gradually acquires its brown tint, with the I'orniatiun
of sporidia in the ascus and the rooting filaments at its base,
and is then complete. In ntlior species of Ery^\j)lic oocysts
and aiiLheridia are prodiu imI in a similar manner.
The formation of perithecia in Eurotiuvi is analogous (Fig.
34). The generative filaments twist together at their summit like
a corkscrew, generally presenting six turns closely united to each
other and forming a hollow body. Then follow the production
of a multitude of cells in the central cavity. De Bary thinks
it not impossible that at this time some act of fertilisation
takes place, but tliere is not even as much evidence in favour
as is adduced in the case of Enjsiphc, and hence it remains as
a guess.
The opinion of Ue Bary may have some weight, but it is in-
sufficient without the evidence, which thirty years following has
not produced, in support of this conclusion — that " the pheno-
mena which take place in Erysiphe authorise us to presume
that in others of the Ascomycetes, having isolated perithecia,
the stroma which encloses several conceptacles, or even the
organs of fructification in the Discomycetes, the Tuberaceae, and
other groups, are also the products of a sexual generation."
There is an abundance of instances in which pyrenomycetous
Fungi have two, three, or four distinct kinds of fructification,
but from this fact alone nothing can be concluded as to the
process of fertilisation or the existence of sexuality.
In the Discomycetes, with its four thousand species, the
suggested examples of sexuality are but very few, and these
have not been confirmed. Woronin examined Lachnca 2iul-
cherrima, and succeeded, as he thought, in recognising that
the receptacle derives its origin from a short and fiexible tube,
thicker tlian the other branches of the mycelium, which is
soon divided by transverse partitions into a series of cells,
since denoniinated a " scolecite." He seems also to have
assured himself that there is always in proximity to this body
certain filaments, the curved branches of which, like so many
antiieridia, support their extremities on the " scolecite." This
contact would ajipear to connnunicate to the scolecite a special
58
INTRODUCTION TO THE STUDY OF FUNGI
vital energy, which is immediately applied to the production
of the filanieutous tissue on which the disc is later to be
borne.
Tulasne found the scolecite readily in Ascobolus furftiraccus,
but failed in tracing fertilisation ; but he was rather more
successful with Pyronema melaloma, in which he found that the
scolecite is certainly a
lateral branch of the
mycelium (Fig. 35).
This branch is simple,
or forked at a short
distance from the base,
and its diameter gen-
erally exceeds that of
the filament which bears
it. It is soon bent and
often elongated in de-
scribing a spiral, the
irregular turns of which
are lax or compressed.
At the same time its
cavity is divided into
eight or ten cavities.
Sometimes he had seen this special branch terminated by a
crosier, and interlocked with the bent part of an analogous
crosier terminating a neighbouring filament. In other cases
the growing branch was connected by its extremity with that
of a hooked branch. These contacts, therefore, seemed rather
accidental than constant. There was, however, no room to
doubt that the scolecite was the habitual rudiment of the
fertile cup.
The most complete observations were those on Pyronema
omphalodes. The globose vesicles, or macrocysts, which are
the beginning of the fertile tissues — each of them emits
from its apex a cylindrical tube, always more or less bent
in a crosier shape, so that the vesicles resemble so many
tun-bellied, narrow-necked retorts filled with a roseate plasma.
Out of the same filaments are produced elongated clavate cells,
named imracysts, which soon exceed the macrocysts in height,
Fig. 35.— Scolecite. After De Bary.
FERTILISATION 59
and seem to carry their summit so as to meet the crosier-
sliaped appendages, and they are soon united two and two.
The union or meeting of the extremity of the crosier tube
with the neighbouring paracyst was a constant fact, which he
had observed a hundred times, and leisurely during a few
months. There is no joining of these cells except in the very
limited point where they meet, and there may be seen a
circular perforation at the end, defined by a round swelling.
Elsewhere they may be very near together, but they are
always free from any adherence whatever. One thing can be
Fig. 36. — Formation of zygospore in Mucorini. After De Bary.
affirmed — that the conjugated cells, especially the larger,
wither and empty themselves, while the upright tubes, which
ultimately constitute the asci, increase and multiply.
As to the " scolecite," so called, there can be no doubt
that some such bodies have been seen, but their significance
has been misinterpreted. It is probably the first distinction
of the fertile from the sterile liyphae, and in no sense
represents the female organ.
The above will suffice for the Ascomycdcs ; and we have
left to us the Phycomycctes, as containing the most decided and
definite examples of sexuality amongst the Fungi. The IMucors,
in species already investigated, develop zygospores from the
hyphae of the mycelium (Fig. 36). A short clavate branch
is produced from each of two neighbouring liyphae. These
branches approach each other by their apices until they touch,
and are called the suspenders. Tliey contain an abuiulance of
6o
INTRODUCTION TO THE STUDY OF FUNGI
protoplasm, and gradually enlarge, until they resemble a spindle-
shaped connective of the two hyphae. A septum is soon
formed across the suspenders near the upper extremity, cutting
off a discoid cell from each, which are separated from each
other at the point of contact by the original wall of the con-
jugating suspenders. This division soon becomes perforated,
and at length disappears, leaving the twin discoid cells united
into a subglobose central cell, which is to become the zygospore.
The membrane thickens, becomes warted and of a dark colour,
nearly black, when it reaches maturity, and in this condition
it settles to a period of rest.
In the family of Sa'prolegniaccdc the oogonia, or female
cells, are terminal on short branches of the mycelium (Fig. 37).
They are globose cells, the membrane
of which is soon perforated, and at the
same time the contents are differentiated
into rounded little spheres which float
in the interior. From the pedicel of
the oogonia, or from neighbouring
hyphae, short curved branches arise,
which bend towards the oogonia. These
are the antheridia, which become slightly
swollen at the apex, and closely applied
to the wall of the oogonium. About
the time when the gonospheres are
formed each autheridium projects into
the cavity of the oogonium one or more slender tubes,
but these appear never to open or discharge their contents,
so that they cannot fertilise the gonospheres, which, however,
soon acquire a cellulose membrane, and the process is complete.
This is the general character of the sexual reproduction,
modified somewhat according to the genera and species.
Another family, the Peronosporaceae, possesses an asexual
system of reproduction much resembling that of ordinary
Mucedines, but parasitic on living plants. The mycelium,
which permeates the tissues, usually in the autumn produces
oogonia, which arise from swellings of the mycelial tubes.
These take the form of globose cells, which become divided off
at the base by a septum (Fig. 3 8). Other branches swell at the
Fig. 37. — Oogonium with
two oospheres of Achlya.
After De Bary.
FERTlLISAriON
6i
extremity and become clavate, to form antlieridia, and one of
these applies itself by the obtuse extremity to the face of
each oogonium. With this development the contents of tlie
oogonium become aggregated into a spherical form in the
centre to constitute a gonosphere. A slender tube is projected
from the applied end of the antheridium into the oogonium
until it reaches tlie gonosphere, when it ceases further growth.
After this contact the gonosphere becomes invested with a
membrane of cellulose. Its subsequent progress, as it becomes
mature, is to develop a thick brown epispore, and then come
to rest during the winter months.
In the Entumojyhthoraccae, or tly moulds, the process of
b
Fig. 38. — Sexual orgaus of Peronospora. After De Bary.
sexual reproduction is more shnple. They are produced by
slightly varying modes, as a result of the conjugation of
opposite threads. These hyphae, either within or without the
body of the host, produce lateral outgrowths at opposite points
of two different threads, which meet midway between the two
conjugating cells, and coalesce. The intermediate walls are
absorbed, and a connecting tube is formed, through which tlie
contents are mingled (Fig. 39). A gemma is produced on the
connecting canal, which increases rapidly, appropriating the
contents of the two conjugating cells to form a zygospore.
After this the empty hyphae disappear. This process may be
slightly modified in different species, but it follows the same
principle, and the mature zygospore passes into a period of
rest.^
This much is known of sexual rcproduclicm in Fungi: that
1 Vegetable Wasps and Plant Worms, by M. C. Cooke (18512), p. 11.
62
INTRODUCTION TO THE STUDY OF FUNGI
in the Phycomycetes two bodies are formed, or two specialised
cells come into contact, and the result is a zygospore, which is
nearly always a resting spore, as a
consequence of fertilisation. In the
Disco my cetes it is suggested that in its
earliest stage the elements of the
future cup, or receptacle, become
fertilised by contact with specialised
filaments which represent antheridia.
In the Perisporiaceae special processes
are affirmed to be emitted i'rom two
adjacent hyphae, which are supposed
to become respectively oocysts and
antheridia, and from their contact to
result in the production of fertilised
perithecia. In the whole of the
Basidiomycetes no definite mode of
fertilisation has been confirmed.
And for the rest, we seek for evidence
Fig. 39. — Conjugating hypliae
in EntomojMhora.
In Tilletia, a genus of the Ustilagineae,^ peculiar phenomena
undoubtedly take place in the conjugation of promycelial spores,
but this can scarcely be interpreted as an act of fertilisation.
The facts are simply these : when the spore germinates it pro-
duces a promycelium, a germ tube, which gives origin to bodies
called primary sporidia, or, more properly, promycelial spores
(Fig. 40). "A very remarkable feature about these ' primary
sporidia ' is that they almost invariably conjugate in pairs ; that
is, adjacent pairs become organically united by a short tube grow-
ing from one and becoming blended with the other, thus placing
the protoplasm of the two sporidia in direct communication.
In some instances conjugation takes place before the primary
sporidia break away from the promycelium. After conjugation a
slender germ tube is formed, which receives all the protoplasm
from the two united sporidia, and if developed upon the proper
host plant, penetrates into its tissues and forms a mycelium,
which in turn produces a new crop of resting spores. In some
^ Manograph of British Uredineae and Ustilaginae, by C. B. Plowiight, London
(1889), p. 8ci.
FERTILISATION
63
species the process is more complicated : the germ tubes pro-
duced by the primary sporidia after conjugation give origin to
secondary sporidia ; these in turn produce germ tubes capable
of penetrating the tissues of the host and giving origin to rest-
ing spores." ^ Similar cases occur in Ustilago.
Tulasne was rather sanguine when he wrote ^ that assiduous
observation, and the perfection \\\\\\ wliicli microscopes are
constructed will have enabled
the botanists of this age to
determine that there are no
really agamic plants — that is,
without sex. At any rate they
can from the present time sus-
pect with foundation that in
all vegetables, no matter to
what group they belong, there
exist two distinct orders of
reproductive organs, the rela-
tive values of which may be
compared to that of the two
sexes in animals. Until
latterly, however, tlie Lichens
and Fungi seemed to form ex-
ceptions to this rule, for all
the researches of phytologists could not discover in them that
duality of organs which, after having been for so long the
exclusive privilege of cotyledonous plants, has since been found
to belong to nearly all cryptogams. Experience and investiga-
tion of forty years have shown that Lichens and Fungi still
remain practically exceptions to the rule of sexuality.
^ Massee, British Fungi — Phycomycetes, etc. (1891), p. 166.
^ Comptes Rendus, vol. xxxv. (1852), p. 841.
■TiUetia in germination.
De Bary.
CHAPTEE VII
DICHOCARPISM
By tlie term expressed in the heading to this chapter we intend
to indicate such species of Fungi as present two distinct forms
of fructification, presumably proceeding from the same mycelium
or vegetative system, and hence pertaining to the same species.
It is contended that the word " dimorphous " would have
expressed this, but we cannot assent, because that word vaguely
distinguishes the organism to which it is applied as having
two forms, which might apply to the fruit, to the carpophore,
or to any other organ, whilst we desire its restriction to such
Fungi only as exhibit two forms of fruit. There are a very
large number of Fungi which might be brought under this
designation, and those would fall into three groups. (1) Those
which produce two forms of fruit from the same stroma or
mycelium. (2) Those which are reputed to possess two forms
of fruit, the genetic connection of which has not yet been
clearly demonstrated. (3) And those which produce two
forms of fruit successively or alternately, by an alternation of
generations.
We shall attempt to give only a few illustrations of the
first kind — those which produce two forms of fruit from the
same stroma or mycelium — which will be sufficient to make
clear the purport of our definition, and the first shall be selected
from a genus in which probably all the species are dichocarpous.
Hypomyces is one of the genera of the Pyrenomycetes which
grows upon dead Fungi, chiefly the Hymenornycetes, in broadly
effused patches, the mycelium of which is partly innate. The
woody Fungus Fomes anyiosas may sometimes be found with a
white mycelium running over the hymenium and penetrating
DICIIOCARPISM 65
the substance. Upon this mycelium erect branches are
developed and lurin conidiophores, having the habit and
appearance of an ordinary mould. The conidiophores branch
several times; the branches, usually three, form a whorl, and
these again are similarly branclied in whorls, the branchlets
being each terminated by three shorter branchlets in a whorl,
each branch and branchlet being attenuated upwards. The
terminal branchlets bear one or more small oval conidia. This
has been called VcHicillinm microspcrmum. Upon the same
mycelium grow a number of ovate perithecia, which are clad
with a dense short woo/; and when mature they contain
numerous asci, each enclosing eight lanceolate uniseptate
sporidia. This is Hypomyccs broomeanus, of which the above
mould bears the conidia. Hence this species is dichocarpous,
one form of fruit being the mould, and called the conidia, the
other the Sphaeriaceous Ascomycete with asci and sporidia.
]Maple leaves will furnish another example in the well-
known pitchy blotches which are so common in autumn as
to attract every eye. These black patches, which are closely
adnate to the leaves, are the stroma of Mclasmia acerina, as
it is then called. The perithecia are cells immersed in this
stroma, which contain small hyaline curved sporules borne on
short slender sporophores. After these leaves have fallen on
the ground and lie in damp places during the winter, another
form of fruit is developed within the same stroma ; but in this
instance the sporidia are developed in asci and are nearly ten
times as long as the sporules, thread-like and flexuous. The
stroma itself becomes more corrugated, and splits irregularly to
allow the sporidia to escape. In this state it is called Bhytisma
acerinum, but the two forms are one species, with a stylosporous
and an ascigerous form of fruit, and the latter is never matured
until the leaves have lain for some time upon the ground.
We may select the living leaves of the hedge maple to
furnish our next example, which can be found in summer in
any hedgerow. At this time the leaves are covered or blotched
with white, as if they were whitewashed, but seen under a
microscope this appearance is caused by a dense mycelium of
white threads adhering to the surface of the leaf From these
threads arise short erect branches, which l)ecome constricted
5
66 INTRODUCTION TO THE STUDY OF FUNGI
at short distances, and then divided into cells ; as they mature
the apical cell falls away as a conidium, and then the next,
and the next, in succession, until exhausted ; this is Oiclium
aceris, or the conidial stage of the Fungus which speedily
succeeds it. Little black dots appear to the naked eye to be
sprinkled over the white mycelium in the autumn. These are
the conceptacles which contain the later fruit, forming little
dark brown peridia of a nearly globose shape, and are attached
by delicate threads to the mycelium. Externally the con-
ceptacles are ornamented with a number of projecting thread-
like appendages, curled, or divided and curled, in both directions
at their tips. Internally the conceptacles enclose eight asci,
each holding eight sporidia. In this condition it is known as
Uncinula bicornis, of which the Oiclium produces the naked
conidia.
We may refer incidentally to some Discomycetes, on the
authority of Tulasne,^ who states that in Ocellaria, which occurs
in little tubercles on the twigs of trees,
a great number of the tubercles, which
ought to be transformed into cup-shaped
receptacles, do not pass into this perfect
state until after having produced very
short, narrow conidia, or spermatia, as
he calls them, or else stylospores on
short sporophores, such spores being
equal in size to the true sporidia to
be afterwards developed. Some of the
tubercles confine themselves to this
N^^;/
Fig. 41. Bulgaria stylosporous generation, and always re-
inquDians. ./ i o ' j
main simple " pycnidia" — that is to
say, tubercles or cells enclosing stylospores. The normal
and fully-developed tubercles assume a cup shape, and con-
tain eight-spored asci, as is usual in the Discomycetes. The same
author also cites another species, Bulgaria inquinans (Fig. 41),
which in the adult state represents a very large, deep, black
peziza, is in its extreme youth an obtuse tubercle, the whole
mass of which is divided into ramified lobes, and is of very
irregular form. The extremities of these lobes become, towards
^ Tulasne, Comptes Ilendus, vol. xxxv. (1852), p. 841.
DICHOCARPISM b-j
the surface of the tubercle, recipients from wliich escape for
some time waves either of pure spermutia or of spermatia
mixed with stylospores. Both are ovoid, but the spermatia
are uncoloured and much smaller than the stylospores, which
are as black as the spores of a Melanconiuiii. Tliese two
quotations are given as exhibiting what we have called Dicho-
carpism as it was presented to tlie view of one who accomplished
very much in demonstrating the foct that the same species oi'
Fungus is capable of developing reproductive bodies of more
than one type or form.
Our next example shall be found on a dead twig of birch,
bursting through the bark in black pustules almost as large as
a rape seed, or rather, oozing out in wet weather like thick
black ink. Examined more closely, a mycelium will be found
at the base forming a compact spore-
bed, on which the brown elliptical
sporules grow on short sporophores
closely packed together. When
mature these separate from their
sporophores, and ooze from the apex of
the pustule in an inky mass. In this
condition it is called Melanconmm
hicolor. Later in the season the same
pustules will be found occupied
by a cluster of perithecia, perhaps ,, ,,
f • , 1 T ' r 1 p^^;_ 42. —MdcDicoius.
SIX or eight, placed almost in a circle,
with rather long necks (Fig. 42). Internally these perithecia
contain numerous asci closely packed together, each ascus con-
taining eight sporidia, of an elliptical shape, divided across the
centre into two cells, and known as Melanconis stilhostoma, one of
the compound Sphaeriacei of which the Melanconium bore the
naked conidia, so that we have the same stroma yielding naked
stylospores, and afterwards sporidia enclosed in asci. An
endless variety might be adduced of ascosporous Sphaeriacei
having also a preceding crop of stylospores on tlie same
mycelium.
Here we may cite two examples of another kind which are
described in another chapter. These are — the Mucors, which
bear erect fertile branches surmounted l)v inflated vesicles
68 INTRODUCTION TO THE STUDY OF FUNGI
containing spores and producing from the same mycelium
branches which conjugate and form a zygospore ; also Perono-
spora, the branched carpophores of which sustain zoosporangia,
and from the mycelium produce, sexually, resting spores or
oospores.
Dead box leaves are often to be met with bearing on their
under surface little pink tufts of a delicate mould arising from
a creeping mycelium. Sometimes the fertile hyphae are effused,
and not tufted. The conidiophores are shortly branched, with
the branches in whorls, bearing at the tips of the branchlets
rather spindle-shaped conidia, and then called Verticillium
h(xi. Subsequently from the same mycelium erect branches
form fertile threads, which are themselves sparingly branched,
and bear at their apices small globose sporangia, each enclosing
several minute gonidia. This condition is Mucor hyalinus.
It is nevertheless doubtful if there are any conjugating
branches which form a zygospore, and which would in that
event have been a third form of fruit, but this condition has
never been observed.
The above examples will be sufficient to indicate some of
the forms which, for the want of a better name, we have called
dichocarpous Fungi. They might as truly have been called
dimorphic. In the first we had a Mucedine, or mould, arising
from the same mycelium or vegetative system as a Pyreno-
mycete. That is to say, the same vegetative system produced
two forms of fructification, one having the attributes of a
mould with naked conidia, and the other an Ascomycete with
sporidia enclosed in asci. The second instance was that of a
black effused stroma or cushion-like expansion with the appear-
ance and attributes of one of the Sphaeropsideae, producing within
cells the sporules on short sporophores characteristic of the
family ; but later on the same cells gave origin to sporidia, of
which every eight were enclosed in asci, and the Fungus was in all
respects a Rhytisma, one of the Ascomycetes. In the third we
had a naked mycelium, the erect branches of which produced in
chains the conidia of an Oidium, or white mould ; but at a
later period the same mycelium developed the perithecia of an
Erysiphe, In the fourth instance the pustules of a Melan-
conium gave origin to the conidia characteristic of the genus
DICIIOCARPISM 69
on short sporophores ; but, mixed with these, later on appeared
the peritheeia and ascospores of a compound Sphaeria, an
Ascomycete. In the fifth we had a Mucor producing spores
in terminal sacs, and, by conjugation of other branches,
zygospores. In the sixth a Teronospora with terminal
zoosporangia on branched liyphae, and oospores upon thi;
mycelium. Lastly, a mould, bearing conidia on the branches,
and afterwards from the same mycelium a Mucor with inflated
terminal sacs enclosing spores. All these are examples of a
second form of fruit produced from the mycelium of the first.
We have now to indicate briefly those species which are
reputed to possess two forms of fruit, the genetic connection of
which has not yet been clearly demonstrated. It may be
premised that there are a large number of cases in which an
association of this kin^ has been suspected, but it is needless
to cite more than two or three. In the summer the leaves of
Rumex are often marked on the under surface with mealy
white spots, seated upon discolored blotches of the green
leaves. These spots are caused by a white mould with a short
and simple, rarely branched conidiophore, supporting at the
apex a single elliptical conidium, attached obliquely ; this is
Ovularia ohliqua. In the autumn the same plant, and often
the same leaves, will present similar spots, which do not carry
the mould but clusters of very minute peritheeia, half immersed
in the leaf. These peritheeia contain cylindrical asci, eacli
enclosing eight oblong uniseptate sporidia. It is supposed,
and with some good show of reason, that the mould constitutes
the conidia of this Sphaerdla rumicis, but the connection does
not appear to have been definitely established. We have
observed the two on the same leaf, but not on the same spot.
The leaves of horse - radish {Armoracia) are often seen
covered with whitish circular blotches, upon which are
sprinkled a number of minute black dots, the partially im-
mersed peritheeia of a species of Phijllosticta, having minute
sporules on slender sporophores. Later in the year similar
spots on leaves of the same plant are occupied with peritheeia
of almost identical appearance, but containing the asci, hitherto
immature, of a supposed Sphaerdla armoraciae. There is no
evidence of the relations subsisting between the two Fungi,
70 INTRODUCTION TO THE STUDY OF FUNGI
only a suspicion that the one is a form of the other. ]\Iany
species of Phyllostida growing upon living leaves, as well as
some species of Septoria, are supposed to be in some way
related to corresponding species of Laestaclia and Sphaerella,
but their association has never been determined.
There is scarcely a more common mould on dead herbaceous
plants than Cladosporium herharum, which forms dark olive
patches with a velvety appearance, consisting of flexuous
jointed hyphae and a profusion of long elliptical conidia, at
first simple, and then uniseptate, proceeding from a creeping
mycelium. There is often to be seen, in close proximity, or
even mingled in the same patches, others with similar but
shorter threads and much larger conidia, which are broadly
elliptical, and not only many times septate, but the cells are
again divided at right angles, so as to appear muriform. This
is Macrosporium commune. Some have conjectured that the
one species passes into the other, which is hardly probable ;
others that there is some occult connection between them ; and
it has been intimated that both forms of mould are only
conidial states of a common Sphaeria, with coloured muriform
sporidia, known as Pleospora herbarum. This is another
example of supposed dichocarpism that rests more upon sup-
position than ascertained fact.
Without indicating any particular species, it is generally
believed that the species of Fhoma, which consist of perithecia
enclosing small hyaline sporules borne on short threads, and so
common on nearly every dead twig or herbaceous stem, are
each related to some ascosporous Fungus of similar appearance.
This is probable in at least a great number of instances, but
demonstrated in only a few. What are called "imperfect
Fungi," such as Sphaeropsideae and the Hyphomycetes, are so
called from the impression that they are not autonomous, but
simply forms or conditions of other species. Hence, if the
combined total of these species is accepted at eleven thousand,
there must be an immense number of dichocarpous species of
which we are still in ignorance.
The last section of this subject includes the species which
produce two forms of fruit successively, or alternately, by an
alternation of generations. Although included here for con-
DICIfOCARPISM 71
venience, these are scarcely to be considered as diehocarpous.
The definition would be that each generation or form completes
its career in the same form as it commenced, so that each starts
from a germ, and the cycle is not the career of a single in-
dividual, but of a series of individuals which revert to the
original fonu after one, two, or more intermediate and different
generations. This phenomenon is certainly not common in
Fungi, but it is illustrated in the Uredines, where the series
consists not only of two but of several generations which inter-
vene between those of a like denomination. We will take a
supposititious example, admitting the facts for the sake of illus-
tration. The leaves of pear trees are not uncommonly visited
by a Fungus called RacstcUa cancellata, in which a cluster of
peridia appear on a yellow spot. When mature these peridia
discharge a profusion of subglobose warted brown spores.
This ends the first generation by the production of spores.
According to some these spores are drifted from the pear leaves
to a juniper bush, where they germinate and invest the
bush, producing, as a result, what is assumed to be the same
Fungus, in a different form, upon a new host. In this case
gelatinous cylindrical masses burst through the bark without
any peridium, consisting of elongated, two -celled, hyaline
spores on long pedicels, all agglutinated together by the
gelatine. This is the second generation, different from the
first, but ending in the production of spores, called specially
teleutospores. In the next stage these teleutospores germinate,
and the germ tube produces as buds small promycelial spores,
which are carried by the wind or otherwise back to the leaves
of a pear tree, producing thereon either directly or indirectly
the peridia of RacstcUa cancellata, and so the reversion is made
to the original form, after the intervention of an intermediate
generation as Gymnosporangium sabiiuic. Thus, then, RacstcUa
canccUatcL on pear leaves produces subglobose brown spores,
and these germinating produce Gymnosporangium sahinac, with
hyaline luiiseptate teleutospores on the savin, which is the
second generation. The teleutospores of the Gymnosporangium,
through the intervention of promycelial spores, reappear on
pear leaves as a RacstcUa, and thus the first form of fruit is
reached again after the intervention of a different generation.
72 INTRODUCTION TO THE STUDY OF FUNGI
First generation Eacstdia, second generation Gymnosporangium,
third generation Baestelia, fourth generation Gymnosporangium,
and so on alternately, constituting an alternation of generations.
Technically, there is but one species appearing under two
forms, and the two names represent two conditions of the
same Fungus.
Another illustration may be found detailed in a succeeding
chapter, in which another Uredine is traced through the
Aecidium-form, the Uredo-form, and the Puccinia-form, revert-
ing to the Aecidium-form in the fourth generation. It will be
observed that alternation of generations differs materially from
what we have designated dichocarpism, because in the former
a single generation produces but one form, and the second
generation proceeds directly from the germination of the
first, and consequently upon a new mycelium. In the
latter, dichocarpism, both forms of fruit are produced from the
same mycelium, and belong to one and the same generation.
We have also seen that the number of forms which are possible
to a single species are not confined necessarily to two, but may
extend to three or four, whilst the principle remains the same.
Hence we conclude that the same species of Fungus is capable
of producing, on the basis of its own vegetative system, two or
more forms of fruit, one of which, but not both, may be the
result of sexual fecundation.
CHAPTER VIII
SAPROniYTES AND PARASITES
The student will not proceed far in the investigation of the
nature and relations of Fungi before he is called upon to
recognise the distinctions between saprophytes and parasites —
the species which thrive upon the decay and cause the dis-
integration of dead matter, and those which infest and flourish
upon the tissues of living plants. Saprophytes are numerically
in preponderance throughout the Fungi as a whole, and represent
those forces of rejuvenescence which build up from the ruins
of an old life the forms of a new generation. They are not
only the agents in disintegration, but the immediate con-
sequences of the phenomena of decay. Dead wood, leaves,
fruits, herbaceous stems, and every fragment of dead vegetable
matter, and in some degree of animal, are capable of developing
and supporting new vegetable forms which utilise and assimilate
the chemical products of decay, and inaugurate a new cycle of
activity. In this relation Fungi have been called the scavengers
of vegetation, since if they are powerful in the work of
destruction, they are also the ready agents in regeneration.
The method by which these results are accomplished is some-
what uniform. If a dead log or only a chip of wood lies upon
the ground in a damp situation, it soon becomes permeated by
the delicate, imperceptible threads of Fungus mycelium ; with
4he penetration of these threads the component cells of the
timber become more and more dissociated from each other, a
kind of fermentation softens the material, and it is not long
before the whole mass has become friable and crumbles at a
touch. Before this crisis is reached, and whilst the mass still
adheres together, the Fungus mycelium gives further evidence
74 INTRODUCTION TO THE STUDY OF FUNGI
of its vegetative vitality, and produces here and there external
nodules, which are the commencement of efforts towards repro-
duction. In a comparatively short period of time the surface
of the log is decorated with young Agarics, effused patches of
Corticium or Poria, or other forms of Hymenomycetal Fungi.
Fully thirty-five per cent of the known species of Agarics are
developed in this ipanner from decaying wood and leaves ; and
of the remaining sLxty per cent, allowing five per cent for other
contingencies, it may fairly be assumed that a large proportion
flourish at the expense of the dead roots, vegetable humus,
indistinguishable fragments and remains of vegetation which
are present in the soil, or the dung of animals which are
vegetable feeders. If we turn from the Agaricini to other
of the great groups of the Hymeiiomyceteae, such as the Polyporei
and the Theleplwrei, we shall be still more convinced of the
great, almost overwhelming, preponderance of species which
manifestly flourish upon the remains of previous vegetation.
Of the 9600 known species of Hymenomycetal Fungi, really
parasitic species are almost, if not wholly, unknown.
Although numerically inferior to the saprophytes, the
parasitic Fungi are none the less important, from the sad havoc
they are capable of producing amongst cultivated plants. A
little experience will soon demonstrate that the parasites are of
two kinds — namely, those which establish themselves externally
upon the green parts of growing plants, and do not penetrate
the tissues ; and those which are developed internally, deeply
seated in the tissues of the infested plants, and not making
their appearance externally save for the purpose of fructifica-
tion. The former may be called cjnphytal, growing upon plants,
and the latter endoj^hytal, growing within plants, which may be
illustrated by familiar examples. Common epiphytal parasites
are — the various species of the Erysipliei ; such as the Sphaero-
theca castagnei, or hop mildew ; the Sphaerotheca pannosa, or rose
mildew ; the pea mildew, or Urysiphe Martii ; the mildew of
the maple, Uncimda hicornis ; and many others. These all
appear upon the leaves of living plants in thin effused white
patches, which give the appearance of being dusted with flour.
There are two stages or conditions of the parasite, and both
associated with the web -like mycelium. The earliest stage
SAPROPHYTES AND PARASITES 75
consists solely of the delicate threads which branch and inter-
lace each other, and form a thin white film of mycelium, some-
times on one and sometimes on both surfaces of the leaves, to
which they adhere by means of little projections, or haustoria,
which enter the stomata. From this mycelium arise shorter
and thicker threads, consisting of a chain of oval cells, each of
which ftUls away consecutively from the apex, and becomes a
c'onidium capable of germination. It is these fallen conidia
which mainly give the mealy appearance to the patches. In
former times they were treated as autonomous Mucedines, and
were included in tlie genus Oidium. Later in the year these
same white mouldy patches, when examined with a pocket lens,
will be observed to be sprinkled with
little globose bodies, which are at first
yellow and subsequently dark brown
approaching to black. These are the
perithecia, not larger than small pins'
heads, attached at the base by delicate
filamentsof mycelium (Fig. 43). When Fig. ^z.—Erysiphe lampro-
mature, the membranaceous coat splits '"'''P\ ^'''^^ *^^"^ ^^^
irregularly at the apex, and exposes the
contents, which consist of peai:-shaped asci, or sacs, each enclosing
two, four, or more elliptical hyaline sporidia, which also are
capable of germination and constitute the ascigerous fructifica-
tion of the Erysiphe. From this brief description it will be
evident that this parasite is entirely superficial, or epiphytal,
and that the injury it inflicts is caused by obstructing the
healthy action of the leaves and, in a manner, killing them
by suffocation. When applying external remedies for plant
diseases, such as fungicides, it should first be clearly ascertained
whether the parasite is or is not epiphytal, since fungicides are
more likely to take prompt effect when application can be
made to them direct, and, by removing obstruction, restore the
healthy action of the leaves. Our commonest vine disease,
which attacks the leaves and fruit, is of the present character,
although only the conidial condition is accurately known.
True endophytal parasites are more varied in their character,
and consist primarily of the " rotting moulds " — Fungi which
have the habit and iii)pearance of Mucedines, but with a more
76
INTRODUCTION TO THE STUDY OF FUNGI
complex fructification. The disease which has for some years
attacked the potato, and at one time was called the " potato
murrain," is of this character. This disease is unfortunately
present, and deeply hidden in the tissues of the plant, before
any external evidence is manifest. When the fructification
appears, usually on the under surface of the leaves, it occurs
on vaguely circumscribed patches, which become discoloured
and soon rot away. The mycelium pervades the entire plant
more or less, but especially at the point of issue. The erect
conidia-bearing threads issue singly or in bundles through the
Fig. 44. — Stages of germination of a conidium or sporangium of Phytoph-
thora. a, rii^e condition ; h, contents breaking up into blocks ; which
escape, c, d ; zs zoospores, e ; with two cilia, /, g ; zoospores at rest,
g, h ; and germinating, i, j, k. After Marshall Ward.
stomata, soon becoming branched towards the apex once, twice,
or several times in a furcate manner, the tip of each ultimate
branch bearing a single oval or elliptical hyaline conidium, or, in
the present case, a sporangium (Fig. 44). When mature these
sporangia, for the most part, become granular within, and at
length the granules accumulate in definite spots, and finally
become invested with a delicate membrane ; so that when the
parent membrane ruptures and the contents escape, they do so in
the form of an uncertain number, it may be four or six minute
rounded bodies, each furnished at one end with a pair of
delicate movable hairs. As soon as these are liberated and
they encounter a thin film of moisture, they float away, being
SAPROPHYTES AND PARASITES 77
impelled by the vibrating hairs, as if endowed with animal life,
hence termed zoospores. Thus it will be observed that some
of the species of Fcronospora differ from the true moulds in that
the tlireads do not bear veritable conidia but sporangia, whicli
contain numerous active zoospores which escape on the rupture
of the parent cell and float off on their own account. The
ultimate career of these zoospores is usually brief, for after
floating a short time they settle down to rest, the cilia or hairs
fall away, and a tliin germ tube is projected, which enters a
neighbouring stoma in the leaves of the foster plant, and
originates a new mycelium, and thus extends the action of the
parasite. There is, however, another mode of reproduction
which takes place within tlie tissues of the foster plant, by a
differentiation of the mycelium and the production of oospores,
a kind of resting spores, which hibernate through the winter
and provide for the continuance of the parasite in the spring.
These oospores are of considerable size, and possess a thick
coloured outer coat, and they remain embedded in the old
stems, haulms, petioles, or leaves of the host-plant, quiescent
throughout the winter, and are only liberated by the decay of
the tissues. In the spring, and when uninfected young seedlings
of the host-plant may be supposed to be numerous, these
oospores awaken to activity, the contents become divided in
the same manner as the contents of the conidia were divided,
only much more numerous ; then the thick outer coat is
ruptured, and a great number of active zoospores emerge, ready
to be washed by the spring rains into fovourable positions for
germinating and infesting new plants. In this manner the
parasite is preserved through the winter, and the perpetuation
of the species assured. Whether the zoospores are derived
from the sporangia, which are developed on the aerial branches
of the hyphae, or whether derived from tlie resting spores,
their subsequent history and functions are the same — that is
to say, entering the host-plant by means of a germ tube,
developing a new mycelium, and producing a new infection.
Some species of the Peronosjjoraceac produce simple conidia on
the hyphae, which never develop active zoospores, but ger-
minate at once. Although, as has been shown, infection may
proceed from without inwards, tlie subsequent manifestation of
78 INTRODUCTION TO THE STUDY OF FUNGI
vegetation and reproduction proceeds from within outwards,
and hence these Fungi are enclophytal.
The life-history of some other types of endophytal parasites
is still incomplete. Amongst Mucedines we may instance
Ramularia, with some allied or analogous genera, in which
the mycelium pervades the tissues of living plants, and ulti-
mately hyphae break through the cuticle, and produce conidia
in the air ; such conidia, having the power of germination,
penetrate the host-plant, and cause a new infection. In the
Mclanconicae the species of Gloeosporium produce a plentiful
mycelium within the living plant before localised spore-beds
are formed beneath the cuticle, which latter at length is
ruptured and sporules escape, and these are also capable of
germination and the origination of a new mycelium.
The Hypodcrmeae are, however, amongst the most potent of
endophytal parasites, and of these the Uredineae are almost
ubiquitous. Perhaps no Fungi have been studied more persist-
ently or closely than these, so that the literature would fill
volumes. We shall only select a typical instance from the
genus Fuccinia, as the readiest method of elucidation. The
one which attacks almost all the species of violet, Fuccinia
violae, will answer the purpose. First of all discoloured spots
are observed on the leaves, and then upon these spots small
convex elevations, which are ultimately rather darker in
colour and punctured; these are the spermogonia, which
enclose the minute bodies to which the name of spermatia has
been applied, without any clear idea of their functions, except
that they always are associated with the cluster-cups or aecidia
that appear simultaneously or soon afterwards. Sometimes the
spermogonia are seen on the upper surface of the leaves, with the
aecidia on the under ; or both may be on the same surface, with
the spermogonia in the centre and the cluster-cups surrounding
them. In the present species the aecidium appears in the form
of an orbicular spot on the leaves, or an elongated mass on
the swollen and distorted petioles ; on the leaves the spot is
yellowish, and the cluster-cups are densely packed upon it,
almost touching each other, the margin torn, whitish, and
turned back like a fringe. Within these cups the aecidiospores
are bright orange and in chains, separating themselves at the
SAPROPHYTES AND PARASITES 79
apex when nuiture, and then globose and warted. This stage
was formerly known as Accidium violae. Before either sper-
niogonia or aecidia appear there is always present a plentiful
mycelium in the tissues. The swelling of the petioles is caused
l)y the development of this mycelium, tlie cluster-cups l^eing
developed from the same mycelium as the spermogonia, and
consequently deeply innate and thoroughly endopliytal. The
mature aecidiospores, after voluntary separation from the chain,
will germinate within a few hours, but seldom after a period
of forty -eight hours. Each spore has several germ-pores,
perhaps four or six, but germination seldom proceeds from
more than one. This cylindrical tube continues growing until
it has acquired a considerable lengtli, the coloured contents of
the spore passing meantime along the tube to its extremity,
which finally enters one of the stomata of the proper host-
plant, and there, by branching and progressive growth, con-
stitutes a mycelium, presumably the mycelium which becomes
the spore-bed of the uredospores. If we return to the violet
leaves later in the year, we shall find the under surface
of many leaves exhibiting small raised pustules, which are
scattered all over the surface. These sori, or pustules of the
Uredo, are soon exposed by the irregular splitting of the
cuticle, and the light brown spores, resembling snuff, are freely
distributed. Examined more closely, each pustule will be
found to possess a spore -bed of compacted mycelium, from
which the uredospores grow, at the apices of rather short
hyaline threads or peduncles, which are soon absorbed, leaving
an elliptical pale-brown spore, with a shortly spinulose surface,
as the second stage of an alternation of generations, the problem-
atic spermatia being left out of the question. It must be remem-
bered that the origin of an uredospore-bed is not absolutely
resultant from a germinating aecidiospore, but it may also be
produced by a germinating uredospore, or by the germination
of a promycelial spore. This fact may be associated with the
other fact, that some species of Puccinia are known with whicli
no aecidium has yet been associated. The mature uredospores
have two, three, or four points of germination or germ-pores.
The germination takes place, as in the aecidiospores, within a
few hours, and in liki> manner tlie growing point enters one
So
INTRODUCTION TO THE STUDY OF FUNGI
of the stomata of the host-plant, where it becomes the new
mycelium of a spore-bed, which may either be that of a uredo-
spore or a teleutospore.
The violet leaves which display on their under surface the
pale-brown scattered- pustules of the uredospores, will, later in
the season, exhibit also similar pustules mixed with them, and
nearly of the same size and form, but much darker in colour, or
the leaf may be occupied entirely with these darker pustules,
which contain the teleutospores. Seen under the microscope,
these spores of the third generation will be found to differ
from the aecidiospores and the uredospores in being two-celled
— that is to say, they are divided across the centre by a trans-
verse septum into two superimposed cells of a somewhat hemi-
spherical form, supported upon a
longer and more persistent hyaline
pedicel. They are produced, like
the uredospores, from a spore-bed
of mycelium arranged more or
less compactly side by side. The
apex of the upper cell has gener-
ally a more or less conspicuous
hyaline nipple in the centre. In
this species the coat of the spore
is smooth, but in some others it
is war ted or spinulose (Fig. 45).
The mature teleutospore may
germinate almost immediately, or
in some species only after a con-
siderable period of rest, in which latter case they are practically
resting spores. The germ tube from either cell, projecting through
the germ-pore, is at first a simple tube into which the contents
of the cell pass, and retreat to the upper end, which continues
to grow and become a promycelium. The extremity becomes
divided off from above downwards by one or more septa, and
then each compartment sends off a short pointed branch, which
is soon dilated at the point. This expanded end then assumes
an oval or kidney shape, and receives the contents of the com-
partment to which it belongs. In the course of a few hours
these new bodies are abstricted, and become promycelial spores.
Fig. 45. — Germinating teleutospore
of Picccinia. After Tiilasne.
SAPROPHYTES AND PARASITES
which soon fall away. The germination of teleutosporcs
results therefore in a proniyceliuni, which develops small
secondary or promycelial spores, and these latter are ready to
germinate at once. \\'\\v\\ these promycelial spores are placed
on the damp surface uf the leaves of the host-plant they
germinate, and the growing point enters one of the stomata,
where it forms a mycelium, the contents of the promycelial
spore passing down the tube, whilst the empty spore-case soon
falls away. This new mycelium may produce spermogonia
and aceidiospores, thus reverting to the original point of
departure ; or it may give rise to a crop of uredospores,
without the intervention of aecidiospores ; or it may pro-
duce teleutospores, which are functionally alike or unlike
the parental teleutospores from which tlie promycelium was
derived. Throughout all these mutations there is no diver-
gence from the endophytal character of tlie parasite, which is
of a peculiar and characteristic type. Here, then, we have in
brief the typical life-hist(jry of one of the Uredineae — the
teleutospores in some instances being unicellular, and then
Uromyces ; or bicellular, and then Puccinia ; or multiseptate,
and then Phragmidium ; tlie character of the teleutospore
determining the generic name to be applied to the cycle.
There have from time to time been suggestions of hereditary
transmission in Uredinous infection, but as the frank accept-
ance of such a possibility would weaken the effects of such
results as are claimed to follow upon artificial cultivation, the
advocates of heteroecisni ignore as much as possible all sugges-
tions of hereditary transmission. Analogy nevertheless favours
the probability of inheritance, and some few stubborn facts
seem to support this view. Some years since we had occasion
to examine some celery plants, the leaves of which were badly
attacked by Puccinia, whilst other plants in the same garden
did not show a single diseased leaf Upon inquiry it was
found that the diseased plants were raised from seed which
had been derived from plants badly diseased at the time, but
that the healthy plants were reared from seed which had been
saved from plants without trace of disease, either in the past
year or in their progenitors of preceding years. The foliage of
all the diseased plants was destroyed, and no disease appeared
G
82 INTRODUCTION TO THE STUDY OF FUNGI
in that same garden, upon celery plants, during the succeeding
ten years. The inference certainly must be that the seeds
contained, in some occult manner, the germs of the disease,
transmitted in this way from generation to generation, and
not obtained by local infection of the seedling leaves, from
germinating promycelial spores. If the latter had occurred,
then the infection would not have been confined to plants
descending from infected parents, whilst other plants growing
within less than three feet did not show a spotted leaf; but
both series of plants would, on the contrary, have suffered in
an equal manner.
Another case is related l)y ]\lr. Worthington Smith,
wherein he says it is common to tind hollyhock seedlings show-
ing the Puccinia on their seed leaves. This he had traced to
the presence of pustules of the disease outside the seeds or
carpels, of which he gave a detailed account in the Gardener s
Chronicle. Yet another instance is upon record, in which a
well-known nurseryman had imported Dianthus seeds direct
from Japan. These seeds were carefully grown under glass,
and, immediately they were up in the seed-pans, they were all
attacked and destroyed by the characteristic Puccinia. On
making a microscopical examination of a series of these seeds
mycelium was detected inside the integument which surrounds
the embryo, or infant plant, and within the coat of the seed.
Another and equally conclusive incident has been narrated by
the Eev. M. J. Berkeley, in which plants of Pyracantlia, raised
from seeds imported from Eussia, were all killed by a species
<iiFu%iclaclium,Qx\AduQ}^ mould, whilst old plants of Pyracantlia,
growing at the same place, remained perfectly free from disease.
In this last instance we have corroborative evidence, in which
the parasite was not a Uredine but a mould ; and the doctrine
of inheritance in plant disease is demonstrated to have taken
place with other parasitic Fungi, and is not confined exclusively
to Uredines. It is sometimes objected that these instances can-
not be referred to hereditary transmission, but that they are
simply cases of the transmission of a perennial mycelium. That
does not appear to alter the fact of transmission, for if the parent
transmits disease to its offspring, the disease is inherited from the
parent, whether it has been transmitted by germs or hyphae.
SA /'A'( )/'// ] "FES AND PARASITES 83
It was at one. time regarded as a reproach to those who
studied these endophytes that in very many cases tlie species
were named after the usual host-plant, and it was only neces-
sary t(i kimw the lin.st to Ite able to name the parasite without
trouble ur exumiuaUon. At the present time it is held to be
true that the promycelial spores will not enter by their growing
point, or infect in any way any otlicr plant except the one,
two, or more species of phanerogam u])()ii which it customarily
grows. Such being the case, it was not so very foolish after
all to combine the name of the host with the endophyte ; and
even now we are disposed to doubt if the old grouping of
species, according to the afhnities of the supporting plants, was
not more effective, practicable, and sensible than the more
recent, more complicated, and more unnatural system of the
present day. Tliis, however, is not a point to be discussed
here, inasmuch as it is altogether a question of classification,
when reduced to ]iractice, and Ijclongs to a subsequent cliapter.
CHAPTEE IX
CONSTITUENTS
The chemical composition of Fungi varies considerably as to
quantitative proportions in different families, and, to a less
extent, in different genera and species. The larger Fungi, which
constitute the Hymenomycetes, are those which have generally
been submitted to analysis. Of course water is a considerable
element in fresh specimens of fleshy Fungi. The highest per-
centage is about 9 0 per cent, and the lowest about 9 per cent,
the latter being that of a woody Polyporus. Perhaps a reason-
able mean for Agarics would be about 60 or 70 per cent.^
Taci gives the following analysis of Eussula foetens : —
Water
Maimite
Fibrin (Albuminoids^
Gum
Fungin or Cellulose
Fat .
Ash
67-0
0-G
4-6
1-5
20-0
0-68
5-13
99-51
Acids, etc. undetermined.
The substances allied to sugar are mannite and trehalose,
the latter found in ergot, and elsewhere. Albuminoids are
some form of fibrin or albumen, and distinguished by containing
nitrogen as well as carbon, hydrogen, and oxygen. These
substances are valuable as food, the nutritive value being in
great part indicated by the percentage of nitrogen. This per-
centage is high in Fungi, higher in dried Agarics than in peas
1 Fistidiiia licpaiica contains 86 per cent of water.
CONSTITUENTS 85
and beans. The special kinds of albuminoids are not deter-
mined, but they closely resemble tliose fouml in animal food.
The cell walls consist of meta-cellulose or fungin, but there is
no lignin or woody libre. Some gunnny substance and oil or
fat occur in most species. Mineral salts are found as ash on
burning. Tlie ash or mineral matter varies from 19-8 in
VmUiota ari'cnsu to 3'0 in Fomrs foincntariv!^, calculated on
tlie dried plant.
Vegetable acids of various kinds have been named in con-
nection with Fungi, as citric acid, malic acid, fumaric acid,
oxalic acid — Hamlet and Plowright found "083 per cent in
Fistulina hcpatica — and oxalate of lime or potassium is by no
means uncommon.^ Agaricic acid has been found in Polijporus
officinalis. Other acids of a special nature may be found in
particular species. Some Fungi contain free acetic acid. A
substance called Fungic acid is mentioned by earlier observers,
but this is stated to be a mixture of citric, nudic, and phos-
phoric acids.
The colouring matters of Fungi are still open to investiga-
tion, and especially so by the aid of the spectroscope. By this
means four yellow or orange matters have been determined —
phycoxanthine, which is yellow ; pezizaxanthine, which is the
orange colour of Pcziza aurantia ; and two colours related to
xanthophyll, or the yellow colouring matter of leaves, Phip-
son obtained a red colouring matter from Cortinarius violaccus,
and Stahlsclnnidt a substance whicli constituted 43"5 per cent
of tlie dried Fungus, from what is supposed to have been Poria
purpurea. This substance has been termed " polyporic acid,"
and is soluble in alkalis, with an intense violet colour. Certain
species of Boleti, notably Boldns luridus, contain a yellow
colouring matter which becomes blue on exposure to the air.
riiipson asserted that this was a derivative of aniline, although
neither aniline nor its salts have this property. Stewart -
suggested that indigogen was a yellowish substance which is
' Crystals of oxalate of lime may often be seen upon the surface of the pileus
of Pti/yporus sidfurnis. Oxalic acid in some form has been detected in scores of
species of Hynienomycetes. Hamlet and Plowrij^ht mciiticm a great number
{Jouni. Chcm. Soc., 1877).
- Stewart, Alkaloids in l-'umji. Trans. "Wonlhope Club (1883 1, p. 110.
86 INTRODUCTION TO THE STUDY OF FUNGI
converted to Llue on exposure to air, and possibly the colouring
matter was iudigogen in Boleti. He obtained this pigment,
and found that it rapidly passes from yellow to blue, and from
blue to brown. While in the blue state it can be again reduced
to yellow ; but when it has once become brown it seems to be
destroyed, as the blue colour could not by any means be
restored. The blue matter contains neither indigo nor aniline.
From this it will be seen that he was not successful in deter-
mining this colouring matter in Boletus. It is not by any
means certain that this colouring matter, whatever it may be,
has any relation to the toxicological properties of the Fungus,
as has been generally supposed, although its development is
strong in poisonous species.
Very little can be said of the nature of the odours which
pertain to Fungi, but Stewart has made some suggestions in this
direction. He says that the volatile alkaloid called tri-methyl-
amine is a colourless liquid with a powerful fishy odour, and
is, in fact, the cause of the smell of decayed fish ; it is found also
in the flowers of the hawthorn, and in some Fungi, as in ergot
of rye and putrefying yeast. He also hints at the possibility
of the odour of Phallus impudicus and that of Clathrus cancel-
latus being derived from the same source. There are some
Agarics which possess the odour of putrid fish, but they are
small and not common, hence the source of odour is unknown.
The toxicological ingredients of Fungi have been investigated
several times, but the results have hardly accorded in any two
cases, and are still open to inquiry. Bohm has especially
studied Boletus luridus, and found large quantities of choline,
together with a substance similar to cholesterin, small
quantities of muscarine, and luridic acid, which crystallises in
brilliant red needles, and yields succinic acid on distillation.
Essentially the same substances were found in Amanita
jjantherina, but in that the acid crystallises in yellow crusts.^
The Fly Agaric {Amanita miiscaria) yields two alkaloids —
muscarine and amanitine. Muscarine is a strong narcotic,
and in some respects antagonistic to atropin. Amanitine is
identical with the animal bases choline, mentioned above in
Boletus luridus, and with neurine. An eminent physician and
1 Journ. Chan. Soc, 1885, p. 1008.
CONSTITUENTS 87
surgeon has informed us that upon one or two occasions he
successfully employed hypodermic injections of atropin in
cases of Fungus poisoning. The effects of the amanita on the
human subject are singular. " At first it generally produces
cheerfulness, afterwards giddiness and drunkenness, ending
occasionally in the entire loss of consciousness. The natural
inclinations of the individual become stimulated. The dancer
executes a pas d' extravagance, the musical indulge in a song,
the chatterer divulges all his secrets, the oratorical delivers
himself of a philippic, and the mimic indulges in caricature.
A straw lying in the road may become a formidable object, to
overcome which a leap is taken sufficient to clear a barrel of
ale or the prostrate trunk of an oak." The symptoms are
endless in variety, and justify the arrangement of these agents,
toxicologically, with narcotico-acrid poisons.
It is now conceded that glycogen, or " animal starch," is
not confined to the animal world, but is also found in Fungi,
The asci of the Ascomycetes are completely permeated with it,
and at first it is diffused throughout the whole of the young
plant, but soon accumulates in the asci, where it is utilised
in the development of the spores.^ It has also been found in
the Mucors, in some cases throughout the mycelium and the
young sporangia, especially in Phycomyces. The greater part
of it is taken up by the protoplasmic contents of the spores.^
The same authority has found it also in the Basidiomycetes.
By tracing the passage of glycogen from one part of the plant
to another he convinced himself that it plays the same part in
the economy as starch in other classes of plants, and that it is
the first visible product of the absorption of carbon. It is
usually most abundant towards the base of the Fungus, in the
vicinity of the soil. Its quantity is greatest at early periods
of growth of the Fungus, gradually disappearing with growth,
probably from the effect of respiratory combustion.^ It has
been found plentifully in Feziza vesiculosa, in truffles, and in
Fhalhis impudicus. Errara contends that glycogen plays the
same part in Fungi that starch does in other plants. It is not
1 Jonrn. Roy. Micr. Soc, vol. ii. (1882), p. 824.
2 Ibid., vol. iii. (1883), p. 397.
3 Bid., vol. V. (1885), p. 504 ; vol. vi. (1886), p. 833 : vol. viii. (1888), p. 96.
88 INTRODUCTION TO THE STUDY OF FUNGI
formed, like starch, from the free carhon-dioxide of the atmo-
sphere, hut out of previously existing organic carhon compounds,
especially the products of the decomposition of other food
materials. Examination of the ergot of Claviceps has shown
that the oily material is changed into glycogen in the same
manner that oily material is converted into starch in germinat-
ing seed. There is a special accumulation of glycogen in the
capitulum of the young Claviceps.
The varnished appearance of the pilei in some of the
species of Fomes, such as Fomes laccatus, Fomes australis, and
others, is affirmed by Dr. Wettstein to be due to a secretion of
resin. The special hyphae are of peculiar form, thickened
above clavately, and containing when young an oily yellow
fluid. Eventually protuberances appear at the end of the
hyphae, which gradually increase and exude a cap of resin.
These exudations flow together and form a continuous coat.^
A yellow resin has been found by Zopf in Polyporus hispidus,
which exudes when fresh a plentiful supply of yellow juice
like a pigment, which stains paper of a gamboge yellow.
Zopf reports that there are two substances : one is the yellow
resin, which is insoluble in water, but otherwise resembling
gamboge; the other a soluble yellowish -green pigment with
acid properties. The same author states that the bright
red colour of Polyporus cinnaharinus is due to the mixture
of a substance which forms beautiful cinnabar -red crystals,
denominated xantho-trametin, and also a resin.^ In some
specimens of exotic species of Fomes we have sometimes seen
flakes of resin on the pileus which could be chipped off, but
could never be satisfied how they came there — whether from
the tree trunk which supported the Fungus, or as an exudation
from the Fungus itself. The varnished pileus of Fomes nigro-
laccatus becomes quite sticky soon after the application of
spirit to the surface. Fries mentions Polyporus resinosus as
exuding a resinous juice, and it is quite possible that some of
the woody Polypori which grow on coniferous trees will secrete
a considerable amount of resin, as they have a tendency to
become laccate when old.
1 Juurn. Boy. Mia: Soc, vol. vi. (1886), p. 486.
- Bot. Zcit., vol. xlvii. (1889), p. 54, etc.
CONSTITUENTS
The latex of the Hymeiioiuycetes ^ is of variable composi-
tion, and especially of three types, of which the most numerous
examples are to be found in the genus Lactarius, which is
evidently resinous. There is also the Mycena type, which is
confined to a few species of Mycena, and the Fistulina, which
is rather more fluid and contains tannin. The latex tubes are
large compared with the neighbouring tissue, and much
branched, having occasional septa, enclosing a turbid, granular
fluid variable in colour. In most species of Lactarius the
greatest number of tubes occur in the subhy menial layer and
the periphery of the stem ; the former branches in one direc-
tion into the hymenium, and in the other into the tissue of the
pileus. In Mycena the system is more simple, and the tubes
are extremely long, running through the periphery of the stem
and ending in the pileus. In Fistulina liepatica the tubes are
distributed through the entire receptacle, and are not collected
in definite spots, with very few in the hymenium. The milk
is persistently white in Lactarius piperatus, veUereus, and many
others, sometimes acrid, and in other species quite mild. In
Lactarius scrobiculatus, theiogalus, clirysorrliacus, etc., it is at
first white, then becoming yellow. In L. dcliciosus it is at
first deep yellow, and then green. In L. aspidcus and L.
uvidus the milk, which is white at first, becomes lilac. In
Lactarius acris the white milk turns reddish, and in Lactarius
fidiginosus of a dark yellow, approaching reddish. Not only
does the latex differ in colour, in volume, and in taste, but
also in consistency. In some it is creamy and sluggish, and
in others watery, slightly coloured, and flowing freely. All
these features indicate a variability in composition. The
character of the yellow juice of Pcziza snccosa appears to be
unknown.
The phenomenon of phosphorescence has been so long
known in Fungi and so often alluded to, that only a brief
reference is necessary. Several Agarics have this property of
which the largest number, for any locality, have been met with
in Australia. All of them are species found growing upon
dead wood, and all have white spores. Nearly the same story
is related of all of them — to the effect that they emit a light
1 Journ. Boy. Micr. Soc, vol. vi. (1886), p. 833 ; vol. vii. (1887), p. 627.
90 INTRODUCTION TO THE STUDY OF FUNGI
sufficiently powerful to enable the time on a watch to be seen
by it. The effect produced by it upon the traveller, when on
a dark night he comes sviddenly upon it glowing in the woods,
is startling ; for to a person unacquainted with this phenomenon
the pale, livid, and deadly light emanating from it conveys to
him an impression of something supernatural, and often causes
no little degree of terror in weak minds or in those willing to
believe in supernatural agencies. The kind of light emitted in
all cases is described as shining with a pale, livid, and greenish
phosphorescent glow.-^ This luminosity is not confined to the
Agarics, but may be witnessed in those cord-like Ehizomorphs
which run through rotten wood or bark, and are common in
mines. The thin mycelium which traverses rotten wood in
every direction, under favourable conditions exhibits the same
peculiarity. Kecent observations have determined that in
some cases luminosity is produced by species of bacteria.
Yet, under all these conditions and manifestations, the cause
of the light is still as much of a mystery as ever, despite
all suggestions as to its originating in some connection with
phosphoric acid.
This is merely a chapter of hints and suggestions which
might have been considerably extended, but it will be recog-
nised that in many cases there is no satisfactory explanation at
present to be given for the phenomena alluded to. There is
still ample work for the chemist, but it is not remunerative,
and in many instances necessarily one of much labour and
assiduity. Fungi are themselves so putrescent that their
examination must be commenced at once, when they are col-
lected ; besides, it is not always that a sufficient supply for
investigation can be obtained at one time. When we learn
that 50 lbs. of Agarics are required to obtain a quarter of an
ounce of muscarine, and that the same quantity of Boletus
luridus is necessary to obtain from it any alkaloid it may
possess, or discover truly the source of the blue colour, we are
not sanguine as to much advance in this direction.
1 Romance of Low Life amongst Plants, by M. C. Cooke, London (1893),
p. 208.
CONSTITUENTS 91
BIBLIOGRAPHY!
Bekkeley, M. J. Introduction to Cryptorjamic Botany. 8vo. Cuts. London,
1860.
Brefeld, 0. Untersuchungen iiher Entwickelunggescliichte der Basidiomyccten,
etc. 4to. Plates. Berlin, 1878-94.
De Baky, a. Rccherches sur le Dh-elop2Kment dc quelques Champignons parasites.
8vo. Plates. Paris, 1863.
Comparative Morphology and Biology of the Fungi, Mycetozoa, and Bacteria.
English Translation. Roy. 8vo, Oxford, 1887.
Morphologic und Physiologic der Pilzc, Flcchtcn und Myxomycctcn. Cuts.
8vo. Leipzig, 1866 and 1884.
Masses, Geo, The Evolution of Plant Life, Loiocr Forms. 12mo. London,
1891.
British Fungi— Morphology. Sm. 8vo. London, 1891.
Cooke, JI. C. Fungi: their Nature, Infiuence, and Uses. Sm. 8vo. London,
1875.
De Seynes, J. "Organisation des Champignons superieurs." Ann. des Sci.
Nat., 5th sei'ies, vol. i.
Recherches pour scrvir VHistoire naturcllc des Champignons inferieurs.
4to. Paris, 1874, 1886, 1888.
"Flore Mycologique de la Region de Montpellier, etc." Observations sxtr
les Agaricinds. 8vo. Paris, 1863.
TuLASNE, L. and C. R. "Organisation des Tremellines. " Ann. des Sci. Nat.,
3rd series, vol. xix.
Sachs, J. Lehrhuch der Botanik. 8vo. Leipzig, 1870.
Handbook of Botany. English Translation. Roy. 8vo.
De Candolle, P. IMnoire sur les Sclerotium. Paris, 1813.
CuRREY, F., and Hanbury, D. "Sclerotimu Stipitatum and Pachyma Cocos."
Linn. Trails. London, 1860.
Patouillard, N". Tabulae analyticac Fungorum. Roy. 8vo. Plates. Paris,
1883-89.
Bocquillon, H. Anat. et Physiol, der Organcs rcjjroduct. des Champignons et
des Lichens. 4to. Paris, 1869.
Marsigli, L. F. De Generatione Fungorum. Fol. Rome, 1714.
Smith, W. G. Diseases of Field and Garden Crops. 12mo. Cuts. London,
1884.
Ward, Marshall. Diseases of Plants. 12mo. Cuts. London, 1892.
SOROKIN, N. Mykolog. Untersuchungen. 4to. Kasan, 1872.
VuiLLEMiN, P. Mudes biologiques sur les Gliampignons. 8vo. Nancy, 1887.
Tavel, F. VON. Vergleiclicnde Morphologic der Pilzc. Jena, 1892.
! It has been considered advisable to give only a brief bibliography to this
and the succeeding chapters, of the most important works only, or those accessible
in the English language.
PAET II
CLASSIFICATION
CHAPTEll X
FUNGI IN GENERAL
Concise and accurate definitions are diliicult to construct, and
seldom remain long without gathering about them numerous
exceptions. This is especially the case in botany or zoology,
where the continual accession of knowledge gradually renders
old limitations untenable. All the definitions in vogue with
old authors have one after the other been swept away, and
many of those which succeeded them are either gone or going.
Even the old distinctions between plant and animal are no
longer to be trusted, and subsidiary divisions are either diffuse
or vague. In the lower Cryptogamia there has been a great
shaking amongst the dry bones, so that when Algae, Lichens, and
Fungi are spoken of they no longer convey the same absolute
ideas which the same words represented only half a century
ago. Whether the hypothesis associated with the middle of
these three terms is tenable or not, the affinities between
Lichens and the Algae on the one hand, and with the Fungi on
the other, have been shown to be very intimate, and the
difficulties of delimitation increased.
We have only to concern ourselves directly with Fungi,
and here the difficulty of concise definition soon becomes mani-
fest. We need not go back beyond the year 1835, when
Berkeley contributed a short introduction to the fifth volume
of Smith's English Flora, edited by W. J. Hooker, in which he
defined Fungi as " plants, consisting of cells and fibres, always
springing from organised, and generally decayed or decaying,
substances, not perfected when immersed in water, bearing
reproductive sporidia, either externally or internally, naked or
enclosed in variously formed cells, many of which frequently
96 INTRODUCTION TO THE STUDY OF FUNGI
concur in the reproduction of a single individual." This is not
a very neat definition, and not wholly accurate. " Not perfected
when immersed in water " is contradicted by the Saprolegnieae,
which flourish in water. In a footnote to the same volume
the subject is alluded to in the following terms : — "It is almost
impossible to draw up characters which shall in every case dis-
tinguish the three orders of Lichens, Algae, and Fungi. Indeed,
the more natural such orders are, the more difficult it is to
arrive at anything approaching to mathematical precision.
Thus, in general, Algae grow in water ; Lichens in air, drawing
their nourishment from the medium which surrounds each
respectively, and not from the matrix ; while Fungi are
nourished by dead or decaying organised matter, and have
therefore been styled Usterophytac. Yet, true as these facts are
in general, there are a few instances in which to a certain
extent they will be found incorrect according to the letter."
But a very large number of Fungi are not " nourished by dead
or decayed organised matter," but flourish upon and destroy
living organisms, both vegetable and animal. The most com-
pact definition was that which described Algae as cellular
aquatic plants which derive their sustenance from the water
in which they grow ; Lichens as cellular plants growing in the
air, and deriving their sustenance from the surrounding
medium ; and Fungi as cellular plants which obtain their
sustenance from the matrix on which they flourish. This
was the basis of the definitions given by Berkeley in his
Lntroduction to Cryptogamic Botany, published in 1857,
wherein he defines the Algals as " Thallophytes deriving nutri-
ment from the water in which they are submerged," and the
Mycetales, including both Lichens and Fungi, as " Thallophytes
deriving nutriment from the matrix or the surrounding air ;
mycelium more or less evident." Subdividing the latter, we
have Lichens described as " aerial, nourished by air and not
by the matrix, producing gonidia " ; and Fungi as " hystero-
phytal or epiphytal, nourished by the matrix, never producing
gonidia." If we proceed to apply the last definition— for we
are not concerned with the others — to the whole mass of
organisms now generally grouped under the Fungi, we shall
discover that it is inadequate.
FUNGI IN GENERAL 97
The most recent definition of Fungi with which we are
acquainted is that given by Saccardo in 1889: " Cryptogamic
plants, cellular, destitute of chlorophyll, for the most part
having a mycelium, either parasitic or saprogenous, for the
most part aerial." That is to say, they are " cellular crypto-
gamic plants, chiefly developed in the air, either epiphytal or
saprophytal, and mostly with a mycelium, but destitute of
chlorophyll." Kecognising the difticulty in constructing a
definition which shall approach to mathematical precision, we
may accept the above as the nearest approach to accuracy
which ingenuity has yet devised. Considerable emphasis has
generally been placed on the presence of a mycelium in the
larger majority of Fungi, as the analogue of the protonema of
mosses and the thallus of Lichens. The analogy, however, is
imperfect, and rather superficial in most cases, since a true
mycelium is almost a thing by itself.
The gill-bearing Agarics, such as the common mushroom,
have a conspicuous filamentous mycelium, from which the
stem arises, and which permeates the matrix to such an extent
that, in a compact form under the name of " bricks," it con-
stitutes the medium through which cultivation is maintained.
This mycelium or spawn, although only produced artificially in
the case of a single species, is universal to a greater or less
extent in the Hymenomycetal Fungi. In a certain sense it is
the representative of the root in flowering plants, but may
be better regarded as the vegetative system of the Fungus.
During the winter, in terrestrial species, this mycelium remains
in the soil in a hibernating condition, so that a crop of the
resultant Fungi may be looked for in the succeeding year. It
is an open question what portion of the Fungus crop in
any given year should be referred to a perennial mycelium,
and what portion to the germination of the spores of the
previous year's crop. Some mycologists contend that in the
majority of instances, as in the commoner species, the continuity
is maintained by means of the mycelium. We have observed
ill the case of Agaricus {Flammula) carhonarms, which occurred
plentifully on charred ground not under cultivation, that, the
ground not being disturbed, the same Agaric made its appear-
ance regularly for four or five years, and then rapidly declined
7
98 INTRODUCTION TO THE STUDY OF FUNGI
in size and number until it disappeared. When the latter
condition occurred, all traces of the carbonisation of the soil
had long vanished, and it was overrun with docks {Rumex),
which superseded and replaced the Agaric. Whilst the
Agarics were in the ascendant, the soil was found, at all
periods of the year, plentifully traversed by mycelium. In the
case of Agarims {Clitocyhe) nebularis we have remembrance
of a mass of decaying leaves, which was undisturbed for
several years, and produced in succession for as many years in
tlie autumn good crops of the Agaric. The inference would be
that the mycelium was perennial, but it is not wholly conclusive,
since the germination of the spores of one season might also
form a new mycelium for the production of the next year's
crop. The generally-accepted theory of the growth of fairy
rings attributes their peripheral extension to the continued
outward growth of a perennial mycelium. Experienced Fungus-
hunters are well aware of the fact that, in the case of such
Fungi as Cortinarms triumjjhans and Agaricus {A^nanita)
muscarius, they may be met with year after year in the same
spot, under the same birch tree, and often in company ; but
this fact would not of itself demonstrate whether the appear-
ance was due to a perennial mycelium or the rejuvenescence
by means of germinating spores. It is not difficult to cause
the spores of Agarics to germinate artificially in a suitable
medium, but it is almost impossible to carry on the process to
the formation of a proper mycelium. This, however, proves
nothing as to what goes on in a state of nature, where the
conditions are such as not to be successfully imitated. We
may be certain that the hibernation in the Agaric does not
take place with the thin-coated spores, but with the mycelial
threads resulting from germination. In other orders of Fungi,
and in the Algae, the resting-spores secrete a thick rigid outer
coating, in which the hibernating season is passed, the conidia
only, or such spores as germinate at once, having but a thin
envelope. It may be inferred, therefore, that whether the
mycelial threads are persistent from a previous season, or but
recently developed by germination of the spores, it is under
the form of mycelium, and in that form alone, that the winter
is passed. Experience has demonstrated that a keen winter.
FUNGI IN GENERAL 99
when the ground is not well clothed with snow, is succeeded
by an autumn in which the fleshy Fungi are considerably
reduced in numbers — that is to say, it is a bad Fungus
year. There may be other conditions — such as general low
temperature, absence of humidity, etc. — which seriously
affect the latent mycelium and, as in the years from 1890
to 1893, largely diminish the production of Agarics in the
autumn.
The indurated tree Fungi, such as the Polyporei, are
also possessed in many cases of a copious mycelium which
penetrates and disintegrates the tissues of the wood. It may
be taken for granted that neither the arboreal Agarics nor the
Polyporei establish themselves, or can be developed from
healthy living tissue. The wood is at first dead at that
particular spot, and traversed by mycelium, before there is
any external appearance of the Fungus. There is no external
Fungus without an internal mycelium, but there may be a
very profuse mycelium and no external Fungus growth. In
this connection we may indicate that condition which is
popularly known as " touchwood," where the whole substance
is permeated by mycelium, and what at one time was hard
and firm oak or ash timber is rendered so friable that it
crumbles beneath the fingers. In tropical regions immense
masses of mycelium are produced from disintegrated timber,
long known under the name of Xylostrovia, extending for
many feet and of variable thickness, with a texture resembling
white leather. The conditions have all been favourable to the
vegetative system rather than the reproductive. As flowering
plants under exceptionally favourable conditions develop a
luxuriant growth without flowers or fruit, so the mycelium of
Fungi may also vegetate luxuriantly without producing organs
of fructification, until the vegetative growth is checked by
deficiency of humidity or some other cause. Another condi-
tion of mycelium was in past times regarded as autonomous,
under the generic name of Rhizomorpha. In this instance the
mycelium partakes of the character of long, branching or
anastomosing, rigid cords, with a dark or black exterior, often
growing between the bark and timber of dead trees, or
penetrating the timber in mines. It may extend for many
loo INTRODUCTION TO THE STUDY OF FUNGI
yards if the conditions are favourable, and cnn only rarely be
traced to the production of any reproductive apparatus, such
as pileus and hymenium. It has been demonstrated that one
form of llhizomorpha may be traced to a development in
Agaricus mellcus, another form in the common Polyporus
squamosus, and yet another in Foines annosus.
The mycelium is also remarkably in evidence in the large
order Hijphomjjcctes, or moulds, in which there is always at
first a creeping mycelium of entangled threads, from which
arise afterwards the erect and sometimes branched conidio-
phores or fertile threads, surmounted by conidia or spores.
Instances will occur in which, owing to a superabundance of
moisture, no fertile threads are produced and the whole Fungus
consists of a mass of sterile mycelium. It is only necessary to
refer to one of the forms of " blue mould," named PeniciUmm
cruataceum, which will produce a profuse mycelium in fluids,
forming an expanded felted mass resembling the substance
known as the " vinegar plant," So long as there is a plentiful
supply of liquid the vegetative system will go on indefinitely,
but a check must take place and the supply of moisture be cut
off before erect fertile threads and spores will be developed.
Not only the saprophytic but also the epiphytic moulds
commence with a mycelium, in the latter instance concealed
within the substance of the host-plant. Thus we have
characteristic examples in the genus Ramularia amongst
the Mucedineae, and in Cercospora amongst the Dematieae.
These have their analogues in the Perunosporaceae, formerly
classed with Mucedines, but now associated with Mucoraceae as
Zygomycetes.
The majority of the genera of the Physomyccteac are on an
equality with the Hyphomyr.etcs in the strong development of
the mycelium ; and it is in connection with this mycelium that
the phenomenon of conjugation takes place which results in
the production of zygospores, or resting spores, by means of
which the species are preserved through the winter. This
takes place not only in the Mucoraceae, which are mostly
saprophytes, but also in the Peronosporaccae, which are destruc-
tive epiphytes. In the former the myceHum is more or less
superficial, and in the latter innate.
FUNGI IN GENERAL
In the Hypodermeae, which include the Ureclineae and the
Ustilagineae, and are consequently epiphytal, the mycelium is
never absent, although innate and concealed.
The Ascomycetes, as well as the Sphaeropsideae, have possibly
a smaller development of mycelium than the other orders
enumerated, but it is seldom even apparently absent. In
the former the Perisporiaceae furnish many examples of an
abundant mycelium, which is superficial on living plants in
the Erysiphei, and consequently easily recognised by the
naked eye. From this cursory view it will be manifest that
the mycelium is an important, and often a very prominent,
attribute of Fungi.
We have not alluded to some of the smaller groups, and
especially to those which are tentatively associated with Fungi,
such as the Saccharomycetes or yeast Fungi, the Schizomycctes or
Bacterial Fungi, and the Myxomycetes — which latter some
authors have struggled to link with the animal kingdom — but
to this we may return hereafter. Neither have we considered
it necessary to allude here to those compact bodies which are
morphologically dense nodules of mycelium, known as Sclerotia.
They are more or less associated with all the principal orders
in their ultimate fructification, and their evidence was not
essential to strengthen the case.
No dissertation is required in support of that portion of
the general definition which characterises Fungi as " destitute
of chlorophyll." In a few instances a green colouring is
present, but it is not due to chlorophyll, nor is it of the colour
of chlorophyll-green, but of a verdigris or aeruginous tint.
When it is contended that chlorophyll is ever present, and
instances are furnished, it will be time enough to contest this
point ; meanwhile nothing will be gained by attempting to
prove a negation.
The popular mind will perhaps seek to be satisfied with a
little more general information than is to be extracted from a
short definition, and to this end we may instance a few of the
typical forms which are assumed by Fungi, which probably are
the most variable in this particular of any of the Cryptogamia.
The most efficacious manner of doing this seems to be an
indication of the most salient features in the different primary
I02 INTRODUCTION TO THE STUDY OF FUNGI
groups into which they are divided, leaving each group to be
analysed and amplified hereafter.^
Let it be remembered that there are two distinct types in
which the fruit, or spores, are produced in the whole of the
Fungi — that is to say, the naked spores are borne at the tips
of threads or basidia, and exposed from the first or soon after-
wards. This is the first type. In other cases the spores are
enclosed from the first, in some definite number, within little
membranous tubes or sacs called asci, and are not set free
until fully mature. This is the second or ascigerous type.
There is a very small intermediate group which seems to
partake of the characters of both primary types — with the
addition of sexual reproduction — represented by the Phyco-
mycetes.
Let us return to the naked-spored Fungi, These we shall
find to constitute three or four groups of a well-defined
character, of which the largest and most important is that
which includes the large fleshy or woody Fungi best known to
the unscientific public as mushrooms and polypores, which
have the naked spores arranged upon a special spore-bearing
surface called the hymenium. In this group the hymenium is
covered with more or less club-shaped, erect cells, or basidia,
which are surmounted by two to four short spicules, or
sterigmata, each surmounted by a spore. These are the Basidio-
mycetes, or Fungi with basidia, and were almost the whole of the
^ I. Spores exposed, or not enclosed in Asci.
p„„_,„__, j a. Hymenoraycetes. Basidiospores.
rERFECT I j_ Uredineae, etc. Spore cycle.
!c. Hyphomycetes. Conidia.
d. Sphaeropsideae. Sponiles at first produced from a
hymenium, which is at first nearly
covered by their receptacle.
This group evidently belongs to the other three, as there are no asci present,
but'are linked to the Ascomycetes by the perithecium, excipulum, or specialised
cavities in which the spores are produced.
II. Sporidia enclosed in Asci.
a. Pyrenomycetes — receptacle perforate or ruptured.
6. Discomycetes — receptacle discoid.
Including Hysteriaceae — receptacle bilabiate,
c. Tuberaceae — receptacle always closed.
FUNGI IN GENERAL 103
Fungi as known to the older botanists of tlie time of Eay and
Hudson. Another group of the naked-spored Fungi are those
parasites of living plants in which for the most part there are
two or three stages of existence, each terminated by a repro-
ductive body having the nature of a spore. These are minute
and pustular Fungi, popularly known as smuts, rusts, and
brands, but called by scientists the Ustilagineae and the
Uredineae, the spores of each generation being borne singly
upon short slender sporophores. The reproductive organs are
at first covered by the cuticle, which splits at maturity and
exposes the naked fruit. The third group are either sapro-
phytes or parasites, but still of minute size, and may be
superficial in the former case, or innate and erumpent in the
latter. The mycelium gives rise to erect threads, either
simple or branched, which produce at the apex, or distributed
over the branches, naked spores, either singly or in clusters,
capable of germination. These are the moulds, or technically,
the Hypliomycetes, which are regarded generally as imperfect
Fungi, a sort of transition stage or conidia of some higher and
more developed forms. The fourth group are also imperfect
Fungi, and may be either saprophytic or parasitic and of
minute size. In this aberrant group the majority do not
produce from the first exposed naked spores, in which feature
they are distinct from the three preceding groups ; but the
spores are enclosed within a globose or flask-shaped receptacle,
which dehisces at the apex when mature and permits the
spores to escape. In certain subsections the receptacle is
spurious or almost obsolete. In all cases the spores are pro-
duced singly at the tips of very short threads, and are
expelled when mature. These Fungi were formerly called
Coniomycetcs, or constituted the bulk of the Coniomycetes, but
are now better known as the S2}haeropsideae and Melanconiaceae.
They are so small as hardly to be visible to the naked eye,
and hence have never acquired a popular name. These groups
represent the naked-spored Fungi, and, it will be observed,
contain only one group in which the individuals are sufficiently
large to attract general attention, and still to some untutored
minds represent all which they recognise as Fungi. Of the
other groups, the rusts and smuts, and the moulds, are
I04
INTRODUCTION TO THE STUDY OF FUNGI
partially and obscurely known, rather from tlieir results than
from their own nature.
The second type of reproductive structure is the ascigerous,
in which the spores are not produced naked at the extremities
of sporophores, but are enclosed within special membranaceous
cylinders or sacs called asci, without sporo-
phores, and are dispersed by rupture of the asci
when mature (Fig. 46). Generally they are all
included under one terra as Ascomycetcs, but
three subdivisions are recognised as furnishing
important distinctions, which may be referred
to with advantage in a general purview. The
largest subdivision is that in which the re-
ceptacle which encloses the fructification is
always closed during growth and development,
and only perforated or ruptured at maturity
for the dispersal of the spores. The typical
form consists of a minute, subglobose receptacle
or perithecium, which may be fleshy, mem-
branaceous, leathery, or carbonaceous, and
either superficial or immersed. Externally
they are often black or dark-coloured, and are
seated upon a more or less distinct mycelium.
In some cases a number of them are associated
together, and sunk beneath the surface of a
common stroma, which is consequently larger
and more conspicuous. In all cases the
interior of the receptacle is filled with a
mass of parallel tubes, closed at each extremity,
and containing normally eight, but sometimes
four or sixteen, or rarely a larger number of minute spores
or sporidia. Side by side with these tubes or asci will
commonly be found a great number of thread-like filaments,
possibly abortive asci, but known as paraphyses. The upper
extremity of the receptacle is sometimes elongated into a neck
which is perforated, or the neck is suppressed and the apex
of the perithecium is perforated to admit of the discharge of
the mature sporidia. This subdivision goes by the name of the
Pyrenomycetes.
-Asci and
FUNGI IN GENERAL 105
The second subdivision of the Ascomycetes differ chiefly
in the character of the receptacle, which is fleshy or leathery
and cup-shaped, and only closed when very young, but soon
expanded, reaching in some cases a diameter of several inches,
but often much smaller or very minute. The disc, or inner
surface, of these cups is compact, rarely gelatinous, and consists
of the hymenium, or fruit-bearing surface, which is composed
of asci and paraphyses, closely packed side by side. The great
feature, then, which distinguishes this subdivision from the
last is the open or cup-shaped receptacle and the exposed disc,
or hymenium, and hence called the Biscoviycetcs.
A small section occupies a position somewhat intermediate
between these two large orders, possessing some of the char-
acters of both. The perithecia are black and leathery or almost
carbonaceous, closed at first and when dry, but opening when
moist with a longitudinal fissure, and then gaping and exposing
the hymenium ; so that whilst it approaches the Pyrenomycetes
in texture, and to some of them in the elongated mouth, it
comes near to Discomycetes in the compact exposed disc, so
that sometimes it has been united with one subdivision and
sometimes the other. In both cases it is maintained as a
distinct order under the name of Hysteriaceac.
In addition to all these are those subterranean Fungi, of
which some are called Truffles, in which, although the spores
are contained in asci, the latter are not enclosed in a peri-
thecium, as in Pyrenomycetes, nor arranged in a disc, as in
Discomycetes, but are dispersed throughout the whole sub-
stance, within folds or cavities, and are not expelled or set
free, except by the decay and dissolution of the entire fabric.
These are the Tuberaceae.
In order to complete this general survey of the chief larger
groups of Fungi, we must allude briefly to a somewhat confused
section, which still maintains the old name of Phy corny ceteae,
somewhat extended in its scope. It will be observed that
although the spores or conidia are naked in some genera, as
they are in the moulds, they are in other cases enclosed within
a common membrane, suggestive of Ascomycetes. The feature
which is relied upon for holding together this rather hetero-
geneous agglomeration of genera, is their sexuality, extending
io6 INTRODUCTION TO THE STUDY OF FUNGI
mostly to the phenomenon of conjugation, and resulting in the
production of zygospores. The polymorphic developments
cannot be described here, so that it must suffice to say that
the ]\Iucors and the fish moulds {Saprolcgniaccae), the rotting
moulds, such as the potato disease {Peronosporaceae), and some
other peculiar forms of Fungus structure, constitute this inter-
mediary group.
What remains of the organisms generally grouped under
the designation of Eungi are rather outside families, the rela-
tionships of which are still somewhat uncertain. Here belongs
the Saccharomyceteae or yeast Fungi, although there is no room
for doubt that they are correctly united with Fungi ; but their
aquatic habit, low organisation, and imperfect fructification
justify their relegation to the lowest place. They are, doubt-
less, to a large extent, imperfect forms of some higher group.
Similarly the microbes, or bacterial organisms, the minute
Schizomyceteae, which in former times were mostly associated
with Algae, find a place in juxtaposition with the Saccharomy-
ceteae. The most aberrant group are the slime Fungi, or Myxo-
myceteae, which some few naturalists still claim for the animal
kingdom. In the early, or vegetative, stage they certainly
conduct themselves in a manner totally different from other
known Fungi, assuming amoeboid forms, and uniting in a Plas-
modium ; but in the final, or reproductive, stage they follow a
completely fungoid type, and produce spores which are not to
be distinguished from the spores of Fungi. Thus much is
sufficient to indicate the predominant features of the chief
groups of Fungi, and to demonstrate what are the kind of
plants which are associated within the limits of the short
definition first given of a Fungus.
In the words of one author, who gave a definition of Fungi
thirty years ago, they " derive nourishment, by means of a
mycelium, from the matrix on which they grow " ; and this
furnishes a clue to their character and functions — i.e. as the
destructive agent in organic nature. That this is really the
case, a few illustrations will suffice to carry conviction. Of
the gill-bearing Fungi nearly 30 per cent grow upon decayed
wood ; the mycelium penetrates the tissues, disintegi-ates the
cells, and produces a condition which we call decay, but which
FUNGI IN GENERAL 107
is in effect reducing it to a pabulum capable of supporting the
life of the Agaric which is to be developed from the mycelium.
All decayed wood is more or less penetrated by Fungus mycelium,
whether the Fungus itself is developed or not, the full develop-
ment depending upon a sufficiency of moisture, or other sur-
roundings. Doubtless continued moisture predisposes the wood
to decay, dissolves what is soluble, softens the cell walls, and
induces a kind of fermentation ; the growing mycelium does the
rest by slow disintegration and the liberation of the chemical
constituents, so that the main factor in the destruction of dead
wood is Fungus mycelium. The destructive process is extended,
in like manner, to dead leaves fallen on the ground, and con-
sequently continually moist, their final reduction to vegetable
humus being expedited by the growth of Fungus mycelium.
There are about 7 per cent of British Agarics which flourish
habitually on dead leaves or the dead stems of herbaceous
plants. We have computed that about 64 per cent of the
British Agaricini are terrestrial, or nominally so, but we cannot
separate those which flourish on old charcoal beds, on decaying
sawdust, or vegetable humus. Undoubtedly many of those
which grow ostensibly upon the ground thrive at the cost of
buried vegetable matter, the sites of decayed stumps, or frag-
ments of old roots. All we can claim for them is that all
these Agarics flourish upon their matrix, deriving their nourish-
ment from the substance upon which they grow, which must
be nitrogenous, and consist more or less of vegetable or animal
matter diffused through the soil, and not its inorganic con-
stituents. Of the residue of the Hymenomycetes little requires
to be said, since nearly all the Polyporei and most of the
Thelefliord grow on rotten wood, which is penetrated by the
mycelium. Need we mention two species as pre-eminent,
Polyporus hylridus and Mcrulius lacrymans, both known as
" dry-rot," which are in evidence for their power of destruction.
In addition to the Saprophytes are all those parasites which
attack living plants and compass their destruction. There can
be no doubt about the whole family of the Uredincae, the rust
and mildew of wheat, the hollyhock disease, the plum-leaf rust
— all determined foes of the plants upon which they flourish.
And there are upwards of twelve hundred different species
io8 INTRODUCTION TO THE STUDY OF FUNGI
known, which attack various plants, and some unfortunate
hosts are the victims of two or three distinct species, all of
which appear to defy the ingenuity of man to eradicate them.
Equally disastrous in their effects, and persistent in their
attacks, are the " rotting moulds," or Pcronos])oraccae, of which
the potato disease is one form, the American vine disease
another, besides many other species which are only of less im-
portance because the plants they attack are less extensively
cultivated, and less associated with the supplies of human food.
No one who has had experience of any of these pests amongst
his lettuces, onions, tomatoes, or in his clover field, would
estimate lightly their powers of destruction. The UvT/sijjhei
also are a family of external parasites, the copious mycelium
of which take possession of living leaves, and destroy by
suffocation, closing and obstructing the air passages, and are
thus conspicuously destructive. Those parasitic Fungi, of which
a considerable number are now known, which establish them-
selves upon the bodies of living insects, and by the penetration
of the mycelium absorb all the tissues, soon cause death, and
then, in most cases, an external fertile manifestation of the
parasite takes place. In like manner the aquatic moulds of
the family Saprolegnieae, of which the iftoving cause of salmon
disease is an example, take possession of fishes and batrachians,
and carry on the work of destruction. There are also amongst
imperfect Fungi many entire genera which attack living plants
and ensure their quick destruction.
Assuming that the power and influence of the Schizomycetes
are not exaggerated, what an agency for destruction must we
recognise in the bacterial Fungi, now so wildly credited with
being the cause of many of the most destructive epidemic
diseases which affect the human subject, as well as inferior
animals ! If we only admit those which are proven beyond
dispute, it is scarcely possible to estimate the full extent of
the marvellous power possessed by organisms so minute in
the destruction of animal, and probably also of vegetable, life.
And so, as we walk through autumnal woods, we see vegetable
matter all around us in a state of decay, with Fungi living and
thriving upon it at the expense of the dissolving tissues, appro-
priating the changed elements of a previous vegetable life to
FUNGI IN GENERAL 109
the support of a new generation, and leaving behind some of
the results of disintegration to assimilate with the soil. The
gardener makes, in a useless corner, his pile of the castaway
twigs and cuttings of his trees and the derelict haulms of herbs
and vegetables. Soon, over the whole putrescent surface,
mycelium and mould proclaim the advent of a new era of
vegetable activity, and anon the whole mass teems with new
life. In this metamorphosis as the Fungi flourish the twigs
decay, for the new life is supported at the expense of the old,
until finally both destroyers and destroyed return again to the
soil from whence they were derived, to form fresh pabulum for
a succeeding season of green leaves and sweet flowers. What
we call decay is mere change — change of form, change of rela-
tionship, change of composition ; and all these changes are
effected by various combined agencies — water, light, air, heat,
these furnishing new and suitable conditions for the develop-
ment of a new race of vegetables. But what a potent agent
have all these in the myriad forms of Fungi, which, above and
beyond all other conditions of vegetable life, deserve the name
of the " Great Destroyer."
We have already intimated that the relationship of Fungi
to Lichens is closer than to any other of the Thallophytes, as
evidenced by their association, in some methods of classifica-
tion, as members of a mycetal alliance. It may be useful to
indicate here some of the general features in which Lichens
differ from Fungi, in addition to those set forth in the short
diagnosis.
Lichens are perennial ; they grow very slowly, but they
attain an extreme age. Some species, grown on the primitive
rocks of the highest mountain ranges in the world, are esti-
mated to have attained an age of at least a thousand years ;
and one author mentions, after the lapse of nearly half a
century, having observed the same specimen of Stida indmon-
aria on the same spot of the same tree. On the other hand,
the Discomycetous Fungi, which are closest in alliance, are
annual, very short-lived, their whole existence being limited
to a few weeks, rapid in growth and rapid in decay, not a
trace of some species remaining after a few days.
Lichens will exist under conditions of aridity which no
no INTRODUCTION TO THE STUDY OF FUNGI
Other vegetables could support. Some are peculiar to cal-
careous rocks ; a few are found on arenaceous rocks ; many are
common on the granitoid series, including micaceous, gneissic,
granitic, and quartzose rocks ; and Lecidea geoyra'phka is
frequent on the smoothest and purest quartz. Fungi, on the
contrary, must have moisture for their very existence, are
mostly found in damp and shaded situations, and could never
exist under the conditions just enumerated for Lichens.
Of all plants Lichens support extreme cold most success-
fully, whilst Fungi succumb at the approach of frost.
Lichens which grow upon the bark of trees may be seen
flourishing in profusion during the life and vigour of the tree ;
whilst Fungi do not and cannot flourish on growing and
vigorous bark, but on diseased, dead, or decaying spots.
Lichens obtain the greater portion of their pabulum from
the atmosphere, and only their mineral constituents from the
matrix. Fungi, on the other hand, obtain their chief support
from the decaying vegetable matter on which they flourish,
gathering up a large store of the nitrogenous results of putre-
faction and decay, and disintegrating the matrix on which
they prey.
Lichens in their chemical composition contain a large
number of substances which are wholly unknown amongst
Fungi, whereas also the most active alkaloids discovered in
Fungi have no analogue in Lichens, notably those of an active
poisonous character.
Lichens contain but a small percentage of water as com-
pared with Fungi, so that in desiccation they do not shrivel,
collapse, or perceptibly diminish in . size ; whereas Fungi
shrivel up and collapse, so as scarcely to be recognised, become
liable constantly to the attack of insects, or, if damp, subject
to the development of mould ; whilst Lichens may be pre-
served for years under like conditions, without fear of insect
or mould.
Lichens, when collected and cast aside without the
slightest care or precaution, do not exhibit the least tendency
to putrefy ; whilst Fungi, with the utmost care in drying, can
scarce be preserved from unmistakable evidences of incipient
putrefaction.
FUNGI IN GENERAL
It is quite erroneous to state, as some have done, that the
gonidia constitute the only difference between Lichens and
Fungi, whereas the presence of gonidia is only one out of
many differences which exist between them.^ The above
comparison is rather suggestive than exhaustive, but it will
be sufficient for our present purpose, and is intended rather
for the assistance of collectors than as a help to scientitic
classification.
There is yet another general aspect to which we may
briefly allude, and that is the recent views which have been
expressed as to the evolution of Fungi. Mr. A. W. Bennett"
traces all the various forms of vegetable life to three lines of
descent, represented by three distinct kinds of cell-contents —
colourless, blue green, and pure green. The first appears to
originate in the Bacteria or Schizomycetes, from which are
derived the whole group of Fungi. The second and third
do not concern our subject. He considers that " too little
importance has at present been attached to degeneration or
retrogression, which may be exhibited in the partial or com-
plete suppression of either the reproductive or the vegetative
parts." Mr. G. Massee, referring to this subject, says : ^ " The
evidence in support of the idea that the fungi are derived
from the algae by retrogression, is the close morphological
agreement of both vegetative and reproductive parts presented
by certain sections of the two groups ; for example, in the
subdivision of fungi called Phycomycetes the vegetative portion
frequently consists of a long, aseptate, variously branched cell
or hypha, similar to the vegetative portion of such algal
genera as Vaucheria ; in the reproductive portion asexually
formed ciliated zoospores occur in Pythium, Saprolegnia,
Cystopus, etc., while oogonia containing one or more oospheres,
fertilised by motile antherozoids, occur in MonoUepharis. In
a second group, the Mucorini, the sexual mode of reproduc-
tion is effected by the conjugation of two similar branches, as
in the Conjugatae, the resulting zygospore becoming invested
^ See The Dual Lichen Hypothesis, by M. C. Cooke, p. 6.
'■^ Journ. Linn. Soc, " Botany," xxvi. p. 49. 1887.
^ Evolution of Plant Life — Lower Forms, by G. Massee, p. 150. Loudon
1891.
112 INTRODUCTION TO THE STUDY OF FUNGI
by a thick cell wall, and forming a resting spore. Many
species belonging to the Phycomyccies are aquatic, being para-
sitic in the tissues of aquatic plants or animals."
A German author, Dr. Oscar Brefeld, who has accom-
plished an immense amount of hard work in investigating
life -histories, is the authority for the classification of Fungi
upon somewhat of this principle. He divides all Fungi into
two primary groups — first, the Phycomyceteae, or Algal-like
Fungi, characterised by sexual as well as asexual modes of
reproduction ; second, the My corny cetes, where the sexual phase
is completely arrested, and consequently they are propagated
exclusively by asexually formed spores. He further divides
the Phycomycctes into Zygomycetes, producing z}'gospores by
conjugation, and Oomycetes, producing oospheres that are ferti-
lised by motile antherozoids, or by transfusion of the contents
of passive antheridia. The other group, or Mycomycetcs, are
also subdivided into Ascomycetcs and Basidiomycetes, with the
Ustiliyineae as a transition group between Phy corny cetes and
the Basidiomycetal group of the higher Fungi.
Earlier than the above, Sachs proposed a very peculiar
arrangement of Thallophytes, which failed to secure general
acceptance. In this arrangement Algae and Fungi were terms
practically abolished, and the series ran in parallel lines.
" The sole character made use of in tlieir primary classifica-
tion was the mode of reproduction. First came the Proto-
phyta, in which no sexual mode of reproduction is known,
followed by three primary classes, the Zygosporeae, Oosporeae,
and Carposporeae, distinguished solely by the degree of com-
plexity of the sexual process. Each of these four classes was
then divided into a series containing chlorophyll, and a series
not containing chlorophyll, the former including the organisms
hitherto known as Algae, the latter those known as Fungi."
The views of Brefeld and his coadjutors seem so likely to
influence the future of mycologic study, that we may be justi-
fied in giving an abstract of them as summarised by Dr. Von
Tavel.^ According to these authorities. Fungi should be classed
in two primary groups: (1) the Phy comy cetes, consisting
typically of a single cell with sexual functions, the nearest
^ Verglcichende Morphologic der Pihe. Jena, 1892.
FUNGI IN GENERAL 113
approach to the Algae ; and (2) the Mesomycetes and Myco-
niycetes, those higher Fungi with a many -celled thallus,
destitute of sexual organs.
The Phycomycetes are subdivided into two distinct sub-
sections, which are termed the Oomycetes and the Zygomycetes.
In each of these the whole plant consists of a sparingly
branched non-septate cell, which reproduces itself sexually by
antheridia and oogonia, and asexually by swarm-spores generated
in sporangia. The Oomycetes exhibit retrogressions, which are
of the nature of adaptations to a more terrestrial mode of life,
accompanied by a progressive loss of sexuality. The families
of the Oomycetes are — the MonoUe2Jharideae, with antheridia
and oogonia in the form of sporangia, and asexual sporangia ;
and the Pcronosporeae and its allies, with the antheridia re-
duced, the oogonia as sporangia, and asexual sporangia or
conidia. These are united to the Zygomycetes by the inter-
mediate family Untomophthoreae, in which both antheridia and
oogonia are reduced and conjugate, and there are asexual
gonidia. The subsection of Zygomycetes has the sexual fructi-
fication in zygospores, and the asexual in sporangia and gonidia.
In this subsection there is a still further degeneracy in sexual
reproduction. Instead of the union of antheridia and oogonia
to produce the zygospore, there is only a conjugation of the
beginnings of such sporangia, because the conjugating threads
are only slightly swollen, and the male and female organs are
not to be distinguished. Asexual sporangia are present, but
the spores have lost their cilia, and are more decidedly adapted
to a terrestrial life. Five families produce sporophores any-
where on the mycelium, the other two bear their sporophores
on specialised threads which arise from the ordinary mycelium.
The second primary group consists of the higher Fungi, in
two subsidiary sections : (1) the Mesomycetes, which connect
the higher Fungi with the Phycomycetes, and (2) the Myco-
mycetes, which include the Ascomycetes and the Basidiomycetes.
The sexual organs entirely disappear in these higher Fungi,
not being found even in the elementary state, whilst asexual
reproduction appears under a multiplicity of forms. The
Mesomycetes include two subsidiary groups : (1) the Hemiasci,
in which the fructification is by sporangia and gonidia, the
1 14 INTRODUCTION TO THE STUDY OF FUNGI
sporangia being asci-like; and (2) the Hemibasidia, whicli
connect the basidia- bearing forms with the simpler Zygomy-
cetes. The former of these, the Hemiasci, consist of three
families: (1) the Ascoideae ; (2) Protomyceteae ; and (3)
Theleboleae. In these simple forms the sporangium becomes
like an ascus, but indeterminate in size or form or the number
of spores, and these latter are usually shot out with consider-
able force. The Hemibasidia fructify by gonidia without
sporangia, but the gonidiophores partake of the character of
basidia. This group includes the XJstilagineae and the Tilletiae.
According to the views of this author all the higher Fungi had
their origin in Zygomycetes, and the two series of Basidio-
mycetes and Ascomycetes were developed in different directions,
the former excluding sporangia, and developing specialised
gonidiophores or basidia, while the latter retained indefinite
gonidiophores, Init developed those special sporangia wliich are
known as asci.
This brings us to the Mycomycetes, which, as already
stated, consist of two series: (1) the Ascomycetes, where the
fructification is by definite sporangia (asci) and conidia ; and
(2) Basidiomycetes, in which the fructification is by gonidia
without sporangia, and the gonidiophores are determinate
(basidia). Ascomycetes are characterised by the presence of
the ascus, which is a sporangium of determinate form, and the
number of its spores. In many cases this form of fructifica-
tion is accompanied by gonidia and chlamydospores. When
mature the spores are often forcibly expelled. Sexual organs
do not occur, since the earlier observations ascribing sexuality
have been found to consist of misinterpretations. The Asco-
mycetes are subdivided into Exoasci and Carpoasci. The
Exoasci have naked asci borne directly on the mycelium. The
Carpoasci have fruit-bodies ; the asci are not naked, but are
contained in special organs composed of fertile hyphae and
sterile threads, which latter form the walls of the envelope.
The simplest forms of ascus fruits are angiocarpous, and in the
highest of the Pyrenomycetes with a special ostiolum. In the
Discomycetes and Hysteriaceae they are gymnocarpous, or
closed at first and afterwards open. The Carpoasci have also
accessory fruit -forms, and in addition to free gonidia and
FUNGI IN GENERAL 115
chlamydospores there are gonidia which are produced within
special fruit-bodies, or perithecia. Another form of fruit is
beheved to be possible, but has not been found, and these are
ordinary sporangia. The simplest form of gonidia in the
Taphrineae is developed directly from the ascospore, or from
another gonidium. A further advance is the production of a
germ tube on which gonidia are borne. From this it is a short
transition to mycelium bearing gonidia on its surface. These
simple forms may be traced through complex stroma-beds into
the highest specialisation of closed fruit -bodies, sometimes
called pycnidia. Free gonidiophores and gonidial fruit bear
usually but one kind of spores, but sometimes the last pro-
duced are of a different shape from the first. Although the
ascospore is the highest form of fructification, the Fungus often
reproduces itself for generations without developing asci ; and
hence many gonidia and chlamydospores have been classed as
" imperfect Fungi " under Hyphomyceteae, Sphaeropsideae, etc.
The Basidiomycetes are a very large group, and their
important character is the possession of basidia, which are
sporophores or gonidiophores, restricted in size, shape, and the
number of spores. Ordinary gonidiophores produce spores one
after the other, indefinitely, but a basidium produces only a
definite number of spores, and only once, and in a particular
place, and then it shrivels up. Most basidia bear four spores,
but some produce two, six, or eight. As a rule basidiospores
are borne on long sterigmata. The Basidiomycetes are arranged
in two groups : (1) the Protobasidiomycetes, in which the
basidia are septate, and (2) the Autobasidiomycetes, in which
the basidia are not septate, and bear a definite number of
basidiospores. The Protobasidiomycetes, or Fungi with a
septate basidium, are ranged in four distinct groups : (1) the
Uredineae have horizontally septate basidia, always free, never
borne in fruit-bodies, and always produced from a chlamydo-
spore or teleutospore ; (2) the Auricularieae have basidia
resembling the Uredineae, but gymnocarpous, viz. having fruit-
bodies which from the beginning form open hymenia ; (3)
Pilacreae, with horizontally septate basidia, but angiocarpous
or closed fruit - bodies ; (4) Tremellineae, having vertically
divided basidia borne in gymnocarpous fruits.
ii6 INTRODUCTION TO THE STUDY OF FUNGI
The Autobasidiomycetes have non-septate basidia, bearhig
spores only at the apex. They include (1) Dacryoniycetes, (2)
Gastromycetes, and (3) Hymenomycetes, of which the last is
by far the largest. In the first of these the basidia split
downwards into two forks, but are not septate. In the second
the Gastromycetes have the basidia borne inside various kinds
of closed fruit-bodies (peridia), whilst Phalloideae are a sub-
section, with the basidia in their early stages borne in a closed
fruit-body, but afterwards pushed into the air and exposed. In
the third, or Hymenomycetes, the basidia are short and simple,
Ijearing normally four spores on delicate sterigmata, but always
finally gymnocarpous or semi-angiocarpous. The simplest
Hymenomycetes are without a fruit-body. The more complex
forms seem to have originated from these. They are followed
by the gymnocarpous Thelephoreae and Clavariae. Then come
the hemi-angiocarpous forms, with the hymenium on the under
surface of the pileus, on the spines of the Tlydnei, tlie walls of
the pores in the Polyporei, and on the gill-plates of Agaricineae.
Accessory fruit-forms are rare in the Polyporei, but oidia or
chains of gonidia occur in some species, while Heterobasidium
bears ordinary gonidia, and Oligoporus and Fistulina bear
chlamydospores. In the Agaricineae no ordinary gonidia
have been found, but oidia occur in certain genera, and
especially in Nyctalis, where chlamydospores are also
abundant.
This is but a brief summary of the views enunciated by
Brefeld and his disciple Von Tavel, which may be further
illustrated by the following table : —
I. PHYCOMYCETES
With a One-celled Thallus and Sexual Organs.
(1. Monoblepharidcae.
( Peronosporeae.
i. uoMYCETEs I Ancylisteae
9 / ^
*" I Saprolegniaceae.
1^? Chytridiaceae.
3. Entomophthoreae.
FUNGI IN GENERAL
117
II. Zygomycetes
A. Exosporangia
B. Carpos'porangia
-. J Mucorineae.
' 1 Thamnidiae.
2. Clioaneplioriae.
o /Chaetocladiaceae.
I Piptocephalideae.
. / Rhizopeae.
( Mortierellaceae.
II. HIGHER FUNGI
With Septate Thallus and no Sexual Organs.
III. Hemiasci
IV. Hemibasidia
A. MESOMYCETES
( I. Exohcmiasci .
\\l. Carpohcmiasci
-. /Ascoideae.
I Protomyceteae.
2. Theleboleae.
/ 1. Ustilagineae.
I 2. Tilletiae.
V. ASCOMYCETES
VI. Basidiomycetes
B. MYCOMYCETES
■ a. Exoasci .
■ h. Carpoasci
■ a. Protohasidiomycetcs
\b. Ai,
itobasidiomycetes
, / Endomyeetes.
I Taplirineae.
r Gynmoasci.
f '^- -{ Perisporiaceae.
I I Pyrenomycetes.
1 r Hysteriaceae.
1^ 3. < Discomycetes.
I Helvellaceae.
, I Uredineae.
I ' I Auricularieae.
"I 2. Pilacreae.
1 3. Tremellineae.
4. Dacryomycetes.
^1. Gastromycetes—
Phalloideae.
6. Hymenomycetes.
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ii8 INTRODUCTION TO THE STUDY OF FUNGI
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MoNTAONE, C. Syllogc Gcnerum Specicrumquc Cryptngamarum . Svo. Paris,
1856.
OicusTEi), A. S. System dcr Pilzc, Lichcnen, und Algen. Leipzig, 1873.
Bkrkeley, M. J. Fungal Alliance. "Arrangement of Families and Genera in
Lindley's Vegetable Kingdom." Svo. London.
Saccardo, p. a. Sylloge Fungorum. 10 vols. Roy. Svo. Padua, 1882-1892.
Gleditsch. Methodns Fungorum. Svo. Berlin, 1753.
Brongniart. Fssai cVune Classification, etc. Paris, 1825.
Hartig, R. Text-Book of Diseases of Trees. English translation. Roy. Svo.
London, 1894.
CHAPTER XI
NAKED-SPORED FUNGI BASIDIOMYCETES
The only safe course in the study of Fungi or any other of the
multitudinous organisms, whether animal or vegetable, with
which the earth teems, is to proceed step by step from the
general to the particular by a systematic sequence. In a few
cases it may be possible by a reference to figures, or from
incidental circumstances, to attach a name with some approach
to accuracy, but such an act is of no service — it teaches
nothing, it avails nothing, it is only a sham, a delusion, and
a snare. The only road to knowledge is a rough one, but it
must be traversed, and all its difficulties surmounted ; there
can be no creeping upwards by a by-path, for all the by-
paths end at a precipice. The most we can do is to tread
firmly, walk circumspectly, and look* upwards. The study of
Fungi is not an easy one, and cannot be got over - empirically,
but with application and perseverance the difficulties, which
seemed at first appalling, become less so at every step.
An effort has been made in the previous chapter to give a
general idea of the scope of the subject, but such efforts are
never thoroughly effective, cannot be final, and at the best
consist only of a shadowy outline. In the present instance
this outline has indicated the existence of a great group or
cluster of groups in which the spores are produced naked —
that is to say, not enclosed in a general vesicle or envelope,
but borne at the apices of spore-bearing threads. The supports
are sometimes highly developed, but in the primary section,
the Basidiomycetes, they are short and thick comparatively,
and as the name indicates, these basidia, or supporters, are
the distinctive feature in the section.
INTRODUCTION TO THE STUDY OF FUNGI
The term Basidiomycdes is a comparatively new one, and
includes the older Hymcnoinycctcs and Gastromycctes, on account
of their agreement in the possession of basidia, although they
still remain distinct on account of other differences ; but
mainly because in the former the hymenium is always naked
and exposed, whereas in the latter it is enclosed within a
peridium during the early stage, and is only exposed by the
rupture of the peridium, when the spores are nearly or wholly
mature. It is easy, therefore, to distinguish at a glance the
true Hymcnomycetcs from the puff-
ball family or the Gastromycetes, so
different are they in external features,
and only correlated by a minute
character which is not to be demon-
strated without the aid of a micro-
scope. As Fries was not expert in
the use of this instrument, and
seldom took into account features
which could not be observed with
a pocket lens, he appears not to
have suspected the presence of
basidia in the Gastromycctes, whilst
he possessed such a remarkable in-
tuition of the relationships of most
of the groups, that he placed these
two orders in juxtaposition. Montague and Berkeley w^ere the
first to indicate the structure of the hymenium in the puff-
balls, and demonstrate the presence of basidia.^
It will facilitate a comprehension of the terms employed in
this connection if we indicate the features of the hymenium
in Agaricus, in its old and broadest sense one of the genera
of the Hymcnomycetcs, which may be accepted as the type of
the rest. The various modifications of the hymenium in
the several families may be reserved for illustration when
the Hymenomycetes come under special notice. It may be
premised that in the Agarics the hymenium or spore-
bearing surface covers completely the thin membrane, which
is pleated and folded on the under side of the cap, and con-
^ Amials of Nat. Hist., iv, (1840), p. 1.55.
Fig. 47. — Hymenial c^Wsoi Agaric,
a, parapliyses ; 6, basidia ; c,
cystidia.
NAKED-SPORED FUNGI— BASIDIOMYCETES 121
stitutes what are popularly termed the gills. The whole of
this surface is covered with a layer of elongated cells, packed
closely side by side, and attached at the base. These hymenial
cells are of three kinds intermixed, although regarded by some
authors as only variations or modifications of one type
(Fig. 47). The most important of these cells are the fertile
ones, or those which bear the spores at the apex, and in fact
are the true hasidia. We may assume it to be generally true
that these basidia are more or less of a clavate, or club-shaped,
form, narrowed a little at the base into the supporting hypha, and
obtuse at the apex, where they are crowned with four delicate,
short, spine-like processes — the sterigmata, each of which is
ultimately surmounted by a spore. Thus, then, each basidium
is normally tetrasporous, producing at its apex four spores.
In rare cases there may be only two, but typically there are
four. The cell of the basidium, as indeed all the cells of the
hymenium in the Agarics, is uncoloured, but the spores may
be either colourless or coloured according to the group to
which the species belongs. The second kind of cells to be
observed on the hymenium are often larger and longer than
the basidia, naked at the apex — that is to say, without sterig-
mata or spores. They also are much fewer in number, and are
called cysticlia. In some genera these cystidia are very large
and conspicuous, and in certain Polyporei and Thcle2ohorei they
undergo considerable modification, and are sometimes coloured.
At least such are the features of the processes which hold an
analogous position to the cystidia, and which are now generally
regarded as modified cystidia. As to the functions of these
cells opinion is still divided. Some have claimed for them the
character and functions of antheridia, but the majority follow
De Seynes in regarding them as hypertrophied basidia, and
possibly their mechanical function is that of keeping the
lamellae apart. The third kind of cells on the hymenium are
rather smaller than the basidia, similar in form, rounded at the
apex, but without sterigmata, and are sometimes called para-
'physcs. There can be no doubt that they are abortive basidia or,
as J>e Seynes terms them, " atrophied basidia." So that the
three kinds of cells on the hymenium are but three forms or
conditions of the same organ — the true basidia, the hyper-
122 INTRODUCTION TO THE STUDY OF FUNGI
tiophied basidia, and the atropliied basidia. It is not
necessary to discuss the question, when we are occupied rather
with the form than the functions of the hymenium. It will,
however, be borne in mind that it has of late been the custom
of some writers to apply the term " basidia " also to the short
sporophores which are present in the Sphaeropsideae and else-
where, which support the solitary spores. Against such a mis-
appropriation of terms it will be advisable to guard ourselves,
and restrict the name of basidia to the spore-supporters in the
Basidiomycdes, where they are not simple sporophores in the
sense that they support a single spore, but tetrasporous
hymenial cells, surmounted by spicular sporophores. The
same term cannot, therefore, be equally applied to the ap-
pendages of a proper and distinct hymenium and the filiform
spore - bearers developed from the base or side walls of a
diminutive receptacle. A true hymenium always consists of
closely -packed hymenial cells; and wt shall find in the
Hyuunomycetes that not only may it assume the form of gill-
plates, as in the Agaricini, but also as a lining to porous
tubes, as in the Polyporei, be diffused over the surface of teeth
or spines, as in the Hydnei, spread over a plane on one side
only, as in the Thelcphorci, or covering both sides of a vertical
hymenopliore, as in the Clavariei.
In the other order, the Gastromycctcs, the basidia closely
resemble those of the Hymenoviycetes, and have the same func-
tion, bearing spicules at the apex, usually quaternary, some-
times in pairs, and each surmounted by a spore. The
examination must be made in a young state to discover the
basidia, because when mature, and the periderm is ruptured,
nothing will be observed except a mass of free spores, some-
times with the spicule attached and a number of threads.
This is the normal condition in the Lycoperdaceae, but varied in
the Phalloideae, and also in the Nididariaceae and Hypogeae.
In all it will be recognised at once that the hymenium is less
highly developed than in the Hyyncnomycetcs, and more fugitive.
Paraphyses may be present as abortive basidia, but tJie
cystidia are scarcely distinct.
Berkeley records the results of his examination of a cut
section of young Lycoperdon. " If a very thin slice be taken,
NAKED-SPORED FUNGI— BASIDIOMYCETES 123
while the mass is yet firm, and before there is the slightest
indication of a change of colonr, the outer stratum of the walls
of these cavities is found to consist of pellucid obtuse cells,
placed parallel to each other like the pile of velvet, exactly as
in the hymenium of an Agaric or Boletus, but without any
trace of those processes which have been regarded by some
authors as male organs {cystidia). Occasionally one or two
filaments cross from one wall to the other, and once I have
seen these anastomose. At a more advanced stage of growth
four little spicules are developed at the tips of the sporo-
phores — all of which, as far as I have been able to observe,
are fertile and of equal height — and on each of these
spicules a globose spore is seated (Fig. 48). It is clear that
we have here a structure identical with that of the true Hy-
menomycetes, a circumstance which accords well
with the fleshy habit and mode of growth." In
his further observations, in reference to the
Phalloidei, he says that " the fructifying mass
consists of a highly sinuated hymenium. The
walls are composed of elongated, somewhat
spathulate, cells, surmounted with from four to
six spicules, each of which bears an oblong ^ig. 48.— Basici-
spore. The sporophores here again appear to ium and spores
be all fertile and of nearly the same height. It ° ycopenon.
will be observed that when the number exceeds four, the
additional spicule is seated between the two, which form one
side of a square, and that if a sixth is present it is placed
opposite to the fifth. Here again we have an Hymeno-
mycetous fungus, and there can be no doubt that the same
structure will be found in all the Phalloidei."
Thus, then, the relationship between Hymenoinycetes and
Gastromycetes may be regarded as established. But Mr. George
Massee, who is nothing if not evolutionist, has some pertinent
remarks on this subject in a recent monograph.^ He observes
that in the Hymenomyceteae " the progressive differentiation of
the sporophore persistently aims at one object, that of con-
cealing the hymenium until the spores are mature — a statement
^ Monograph of the Brilish Gastromycetes, by George Massee, p. 2. London,
1889.
124 INTRODUCTION TO THE STUDY OF FUNGI
which is not in harmony with the general conception that the
hynienium is from tlie first exposed. In the Thelephoreae,
Clavarieae, and Hydneae the hymenium is exposed from the
earliest stage ; whereas in the Polyporeae and the Agaricineae
the hymenium, in the higher forms, is completely differentiated
while yet concealed by a structure known as the veil, which in
many species is only ruptured by the expansion of the pileus
when the spores are mature. The idea of concealing the
hymenium from the light is equally apparent in the various
orders enumerated, except in the Clavarieae. In the Thele-
phoreae the simplest genera — Corticium, Coniophora, and Penio-
phora — have the hymenium covering the whole of the upper-
most or free surface of the hymenophore, and consequently,
from the earliest period of development, exposed to the light ;
whereas in the genera Stereum and Thelephora a portion of
the hymenophore becomes free from the substratum and bends
over, thus turning the hymenium away from the light ; and by
a series of transitions we find the higher species of the two
last-named genera assuming umbrella -shaped forms with a
central stem and inferior hymenium, but not at any period
covered by a veil. In the Agaricineae -we meet with the same
sequence of the evolution. In such low forms as Cantharellus
rctirugis and Agaricus (Pleurotus) hyjjnophilus the plants are
fixed to the substratum by the pileus with the hymenium
uppermost, and may be compared to a Corticium, with the
hymenium imperfectly broken up into gills ; whereas in such
species as Agaricus (Fholiotct) praccox and Agaricus (Amanita)
muscarius the hymenophore is supported on a stem with the
hymenium on the under side, and concealed by a veil until the
spores are mature." It might also have been urged here, in
confirmation, that specimens of Fames and Polystictus from
tropical or sub-tropical regions, where the light is strong,
are constantly to be met with, in which the log has become
turned and the hymenium of the Polypores growing thereon
exposed to the full light. In such cases, and especially
in such common species as Polystictus sangidneus and
Polystictus occidcntalis, the old hymenium gradually becomes
obliterated, and a new hymenium is formed upon what previ-
ously was the upper surface, but accidentally inverted so as to
NAKED-SPORED FUNGI—BASIDIOMYCETES 125
become the under, and consequently turned away from the
light.
" In the Gastromycetes, with the exception of the species
constituting the genus Gaidiera, the hymenium is completely
concealed by a continuous wall or peridium, until the spores
are mature. ... As already stated, two leading features
stand out prominent in the evolution of the Hymenomycctcs :
the conversion of the primitive even hymenial surface into
gills, thereby increasing the spore-bearing area, and secondly,
the gradual concealment of the hymenium until the spores
are mature. In the Gastromycetes these two conditions are
present in the lowest forms, and persist throughout the group,
the very varied forms presented by the different orders being
the outcome of modifications of the sporophore in connection
with spore dissemination."
We may now proceed to a closer analysis of the two
separate orders of Hymcnomycetes and Gastromycetes.
CHAPTER XII
IIYMENOMYCETES
As lately as 1830 the Butanicon Gallicum included under
the term " Fungi " only the Hymenomycetes and the Disco-
mycetes ; whilst at a much later period the ordinary observer
recognised only a few forms, chiefly of the Discomycetes, as
Fungi beyond the Hymenomycetes. The puff-balls came at
length to be included, but even these for a long time were
regarded doubtfully as to whether they were true Fungi or not.
In this country it was not until 1836, when Berkeley's
volume of Hooker's Eiiglisli Flora was published, that the
proper limits of " Fungi," as then known, came to be under-
stood. Under any circumstances the Hymenomycetes have held
the first place amongst Fungi, are usually the first to attract
the attention of students, and stand at the head of every list,
catalogue, or "Fungus flora." In 1825 Elias Fries himself
included Discomycetes within his order " Hymenomycetes,"
and did not practically dissociate them until 1849, when he
constituted the Discomycetes as a distinct and separate order.
At that period the two orders followed each other, whilst the
Gastromycetes were at a distance and scarcely became approxi-
mate until the basidia had been discovered in them.
The limits of the order Hymenomycetes were thus briefly
expressed : " Spores naked. Hymenium free, mostly naked, or
if enclosed at first, soon exposed." This again was more
expanded and rendered clearer by Berkeley, thus : " Mycelium
floccose, giving rise at once to a distinct hymenium or produc-
ing a variously shaped naked or volvate receptacle, even or
bearing on its upper or under surface various folds, plates,
prickles, etc., clothed with fertile hymenial cells." The pre-
H YMENOM YCE TES
127
ceding chapter having prepared us for the general attributes of
the order, we may proceed to indicate the six groups into which
the genera naturally fall. Four of these have the hymeniuin
normally inferior, in the other two either superior or on all sides.
The four first are the Agaricini, Pohjporci, Hyclnci, Thdcpliorci,
and the two latter Clavariei and Trcmellini.
The Agaricini are pre-eminently soft, fleshy, putrescent
Fungi, of the mushroom type, in which the inferior hymenium,
or spore-bearing surface, is spread over folds, or gills, which
^. ^\f^
Fig. 49. — Agaric. A, young state ; B, mature ; C, section,
radiate from a central stem to the circumference of a pileus or
cap. In the typical form there is a floccose mycelium of
delicate threads, surmounted by a stem, more or less developed,
and crowned by a hood or cap of umbrella-like form, with gills
on the under surface. In the earliest condition the form is
nearly globose, enclosed like an egg within a thin shell or
membrane. As it progresses in growth the egg-shaped body
splits round the centre, the upper hemispherical portion being
carried upwards by a quick-growing stem, forming the cap, the
lower half remaining behind to constitute the base. At the
margin of the pileus the jagged remains of the fissured mem-
brane often adhere for some time, and in like manner there
128 INTRODUCTION TO THE STUDY OF FUNGI
may be traces of the fissure on the basal portion. In the
most highly developed forms patches of the broken envelope,
or volva, adhere like warts to the top of the pileus, whilst the
basal portion remains as a loose sheath at the bottom of the
stem, where it is for some time persistent as a kind of sheath,
called the mlva. By cutting through the entire Fungus longi-
tudinally from the apex to the base, the cut section exhibits
the following features — a rooting mycelium, or spawn ; an erect
stem rising therefrom (.s), winch is sometimes solid and some-
times hollow (Fig. 49) ; the base either equal in dimensions or
swollen like a bulb, occasionally with a distinct outer coat or
volva {v), which is adnate below to the bulb and free above ; the
stem is surmounted by a more or less hemispherical or conical
cap or pileus (/?), the under surface covered with parallel plates,
or gills, which radiate from the stem to the margin of the cap
{g). When the cap is hemispherical the gills are often covered
at first by a thin membrane, which extends from the stem to
the edge of the cap ; and, as growth and expansion proceed, this
veil is torn away from the margin of the pileus, and hangs like
a collar or frill around the upper part of the stem, forming an
anuulus or ring (a). When the cap is conical the edges of the
gills are closely applied to the stem vertically, and the edge of
the cap is only slightly attached to the stem, not forming a
ring. When the substance of the cap descends between the
folds of the gills it is the trama, and then the gills do not
part freely from the cap ; but when the trama is thin or
obsolete, the gills part freely from the flesh of the pileus. The
gills are formed by a membrane, which constitutes the hymen-
ium, and is folded like a fim ; so that each gill is a double
membrane, applied back to back, giving as much surface as is
possible for the production of the spores. The appendages of
the hymenium have already been described as basidia, cystidia,
and paraphyses, with the resulting tetraspores. Thus much
may be seen of the structure in a longitudinal section of the
pileus and stem. Modifications in some one or more of these
general details give rise to the different genera into which the
Agaricini are now divided. In former times, when the system
adopted by Fries absolutely prevailed, the greater part of the
species of Agaricini were comprised in one large genus, that of
HYMENOMYCETES 129
Agaricus, in which the gills were membranaceous, and persistent,
— that is to say, not melting or deliquescent when mature ; the
trama was continuous with the substance of the pileus, passing
down between the folds of the hymenium, and the edge of the
gills was acute. The substance was fleshy and putrescent, not
reviving after being dried. In other genera, as in Coprinus
and BolUtius, the gills deliquesced when mature. In such
genera as Panus, Lentinus, and Lenzites the substance was not
fleshy but somewhat leathery, and not putrescent. In the
large genus Marasmius, as well as in Xerotus and Trogia, the
substance was thin but dry, not putrescent ; readily desiccated,
and reviving when moistened. In Bussula the substance was
fleshy and putrescent, but there were peculiar and special
features which severed it from Agaricus, approaching Laetarius,
in which latter a peculiar milky secretion afforded a distinctive
feature. Hijgroplwrus and Cortinarius were two other rather
large genera with distinctive characters, to be alluded to here-
after. Cantharellus and Scliizophyllum afforded prominent
characters in their thickened or splitting gills. Hence it will
be seen that the old genus Agaricus had one or two prominent
characters, which distinguished it from all the other genera of
the Agaricini, and held together one of the largest genera of
Fungi, which at the present time would not number less than
3000 species. For the purposes of classification Fries sub-
divided this genus into five groups, according to the colour of
the spores — the Leucosporac, in which the spores were typically
white or but slightly coloured ; the Hyporhoclii, in which the
spores were pink or salmon-coloured ; the Dermini, in which
the spores were tawny or some shade of rusty brown; the
Pratellac, in which the spores were brownish purple or very
dark brown ; and the Coprinarii, in which the spores were
black. These divisions are substantially maintained in more
recent times, but applied to the whole of the Agaricini.
In the Friesian system each of these groups was sub-
divided into subgenera, which had their analogues in part in
the kindred groups. In a Clavis published by Mr. Worth-
ington Smith he indicated the corresponding subgenera in
each of the five sections, as far as they were represented in the
British flora. All this disappeared when Professor Saccardo
I30 INTRODUCTION TO THE STUDY OF FUNGI
propounded his scheme for the classitication of the Agariciiii,
for wliich previous authors had prepared the way, by at once
elevating the sul)geuera of Fries to the rank of genera, and
ranking them upon an equality with the other old Friesian
genera. This was the inherent weakness of the Saceardiau
arrangement, although it will doubtless come into universal
use where expedients are valued rather for their utility than
their consistency. No one of experience would contend that
Triclwloma and CollyUa bear absolutely the same relation to
each other that Triclwloma would bear to CantharcUas, or
Collyhia to Lentinus.
In the Saccardian system the four primary groups into
which the whole of the Agaricini are divided, depend upon the
colour of the spores. The Leucosporac, having the spores
colourless or but faintly coloured ; the Bhodosporac, in which
the spores are pink or salmon-coloured, corresponding absolutely
to the HyporJiodii of Fries ; the Ochrosporae, in which the
spores are ochraceous, tawny, or some tint of light brown ; and
the Mclanosiwrar, in which the spores appear to be black but
in reality are dark brown, purple brown, or black, and thus
combining the attributes of the Pratcllae with the Coprinarii
of Fries. Of all these groups the first and largest is the
Leucosporae, containing upwards of 3000 species, or more than
half of the whole Agaricincac, which numbers about 5245
species. These are represented by thirty-two genera, and these
latter arrange themselves in two groups or subdivisions — the
larger, or Haplopliyllae, with the edge of the gills entire, and
the Schizophyllae, in which the edge of the gills is split or
appendiculate. The latter is a very small series, consisting of
four genera but numbering in all not more than fifteen species.
Hence, then, the HcqilopliyUae are the only series which require
any special notice in this place ; and these are subdivided once
more into two subsections, in which the chief distinction con-
sists of the texture of the substance. In the first, or Mollcs,
the substance is more or less fleshy, and putrescent, not reviving
after -desiccation. In the second, or Tcnaces, the substance is
tough and persistent, reviving after being dried. These are
useful distinctions to be borne in mind, and will soon
commend themselves to the practical Fungus - hunter. The
// YMENOM YCE TES
131
common mushroom may be taken as an illustration of the
former, and some species of Marasmiiis or Lcntinus of the
latter.
This brings us in face of the fleshy, or Molles, section of
white-spored Agarics, containing in 1893 about 1750 species,
of which not more than 430 are British, having the edge of
the gills acute and the folds of the hymenium separable.
This corresponds therefore with the Leucosporae section of the
old genus Agaricus. Nevertheless there are additional genera
which agree in their fleshy substance but differ in other
particulars. In this category Hygroj^horus is a genus by itself,
with the gills continuous with the pileus, and not separable
from the trania. Thus the gills and the pileus are practically
of one piece. Many, and indeed most of them, are more or
less glutinous when fresh, and perhaps for this reason are
capable of enduring more frost than others of the fleshy Agarics.
In addition to these are two genera which have the substance
of the pileus of a peculiar vesicular character, soft and fragile,
but which have also another remarkable feature of affinity
with each other in the spores being normally globose. These
two genera are Russula and Lactarms,
the latter with and the former tvithout
a milky juice (Fig. 50). In habit and
appearance they most resemble Ti'icho-
loma, but a little experience will soon
distinguish the difference. They are
almost absolutely terrestrial and soli-
tary, with a short robust stem, and
many of the Bussulae have a brightly
coloured pileus. Commonly, but not
universally, the gills in Russula reach
from the margin to the stem without
intervening short gills; or, when
shorter gills are present, these usually anastomose with the long
gills, so as to appear as if the latter were forked. Finally
there is a small group — consisting of four genera, of which
Cantharellus is the chief — in which the edge of the gills is
obtuse or vein-like. All these subsidiary groups together con-
tain about 500 species, bringing the total of fleshy, white-
Fia. 50. — Lactarius deliciosus,
with section and spores.
132 INTRODUCTION TO THE STUDY OF FUNGI
spored Agarics of all kinds up to 2250 species, or nearly one
half of the whole gill-bearing Hijmenomycctcs.
The other section of the HaiTlopliyllac, with a tough per-
sistent substance, includes six genera in which the substance
is at first fleshy, or gelatinous, and then becoming leathery,
and three genera in which the substance is always more or less
corky or woody. As might be anticipated, these are almost the
only representatives of the gill-bearing Hymcnomycetes which
extend into tropical countries. Of the six genera first alluded
to, Marasmius approaches nearest to the soft-fleshed Agarics,
such as Colhjbia and Myccna, and might readily be confounded,
save for their tougher and drier substance. The larger pro-
portion are of a small size, and these affect dead wood and
leaves. Lcntinus often attains a large size, and, with the
exception of a few European species, is a tropical or sub-
tropical genus. The technical distinction between Panus and
Lcntinus is that in the former the edge of the gills is even, and
in the latter toothed or ragged. In order to complete our
numerical estimate, we may add that the Tenaces section of
Ha'ploi^liyllac is represented by about 810 species. The next
order of Hymenomycctcs, the Polyporcae, is approached most
nearly in one direction by Lenzites, which is the analogue in
the Agaricineae of Daedalca in the Polyporcae.
It seems unnecessary in this place to descend any lower
with an analysis of the Lcncosporac. The analytical key to the
genera in any good local flora will indicate the salient features
in each genus, which it would be rather tedious to introduce into
a book having the character of a general introduction, and
would moreover extend this chapter to an inordinate length.
We must now revert to Saccardo's second primary group
of the Agaricineae — that of the PJiodosporae, so called on account
of the spores being pink or of a salmon colour. It may at the
same time be intimated that, although in some instances these
spores are elliptical and smooth, they are often coarsely warted
and angular. The group in itself seems to be a very natural
one, for the species are all soft and fleshy, and even more
putrescent than the softer of the Lcucosjporac. In all countries
they constitute the smallest of the four primary groups of the
Agaricineae, and have often a disagreeable odour. The total
HYMENOMYCETES 133
number of known species does not exceed 366, or about one-
eighth of the number of white-spored species. There are no
genera, recognised as such, in the Sylloge, except such as
were included by Fries as subgenera in his genus Agaricus, and
these correspond to analogous genera in the Leucosporae. For
instance, the recent genus Metraria corresponds to Amanita,
and Volvaria to Amanitopsis ; Annularia is an analogue of
Lepiota, as Pluteus is of Schidzeria, but there is no correspond-
ent to Armillaria. Entoloma is analogous to Tricholoma,
Clitopilus to Clitocyle, Claudopus to Pleurotus, Lcptonia to
Collyhia, Nolanea to Mycena, and Eccilia to Omphalia. In
this way the characteristic features of the genera which have
been learned in connection with the Leucosp)orae may serve
again with the Rhodosporac, conditional upon the difference in
the colour of the spores. It is somewhat remarkable that the
genus Metraria is only known from Australia, and yet there
are known at present only twenty-five other species of the
Rhodosporae in that large continent, which are spread over ten
genera, and fifteen of the species are European. In the
British flora upwards of 100 species are recorded, or about
one to seven of the Leucosporae. In Australia the proportion
is one to thirteen and a half of the Leucosporae. This indicates
that the Rhodosporac prefer a cold to a warm climate, or at
least a moist to a dry one. In Ceylon the proportion is much
the same as it is in Britain.
The third primary group is Oclirosporac, which includes
the Dermini section of Agaricus, with the addition of the large
genus Cortinarius, and the small genus Paxillns — to which, we
contend, should also be added Bolhitius, placed by Saccardo in
Melanosporae, but evidently the colour of the spores is against
this position. The genus Paxillus is distinct from all the rest
in the facility with which the gills separate from the hynieno-
phore ; and Cortinarius differs from all in the universal veil
being of delicate threads like a spider's web. The residue are
the same as the Friesian subgenera of the section Dermini.
Exception might be taken to the term Ochrosporae, as not being
characteristic of the general colour of the spores, which are
much too deep for " ochraceous," approximating more to
ferruginous, and hence the name is misleading. As we have
134 IXTRODUCTIOX TO THE STUDY OF FUNGI
done with the liliodos2)orae, so we might indicate the analogues
of the Lencospcrrde in the Ochrosporde, in so far as they are
represented. There is no genus which corresponds with
Amanita, hut LoccUina or Acdabularia, whichever name we
select, is equivalent to Amanitopsis. Lepioia and Schulzcria
have no analogue, but Armillaria has a correspondent in
Pholiota. Tricholoma is most nearly represented by Hchdoma,
and in some degree by Inocyhc, the essential difference between
these two genera being the fibrillose cuticle in the latter, and
the smooth viscid cuticle in the former, both of which are
represented in subsections of Tricholoma. CUtocyhc is repre-
sented by Flammula in some of its species only, which have
decurrent gills, whilst Pkurotus has its analogue in Crqyidotus.
For Collyhia we have Naucoria ; for Mycena there is Galera ;
and Omphalia is recognised in Tuharia. It is only in Tnocyle
that we meet with irregular spores such as are not uncommon
in the Rhodosporac.
The total number of recorded species is estimated at 1157,
as against 3 6 6 of the Rhodosporac. The large genus Cortinarius
comprises some 400 species, all of which are terrestrial, and
only subtropical at considerable elevations. Should Bolbitius
be included, it differs from all the rest in the thin membra-
naceous pileus, in which respect it is analogous to Hiatula
in the Leucosporae, and to some species of Cop)rinus in the
Melanosporac.
The fourth and last primary division of the Agaricincac,
according to Saccardo, is Melanosporac, in which he combines
the Pratcllae of Fries with the Coprinai^ii, and adds thereto the
genera Coprinus, Bolbitius (ochrosporous), Gomphidius, Anthra-
cophyllum, and Monta.gnitcs. In some cases it is difficult to
distinguish dark purple -brown spores from black, but this is
hardly sufficient reason for combining them. Mr. G. Massee
felt this to be the case, and in his British Fungus Flora he
adopts two divisions instead of one, namely, the Porphyrosporac
and the Melanosporac. After all there is no great principle at
stake, although personally we would rather, if a coalition be
considered advisable, that the two subdivisions were kept
separate as Porphyrosporac and Melanosporac under a common
designation. Following the same course that we have adopted
HYMENOMYCETES 135
in the other primary divisions, we would intimate the analogies
of the several genera of the Forphyrosporac. The representa-
tives of Amanita and Amanitopsis must be sought in a single
volvate genus — that of Chitonia, in which one or two species
are annulate, whilst the residue are not. Lc2nota has its
analogue in Agaricus proper, which includes the old subgenus
Psalliota of Fries, and Sehuhzeria has its correspondent in the
ringless Pilosacc. As for Armillaria, we shall find an analogue
in Stropharia ; and Tricholoma will have its nearest representa-
tive, but not a very perfect one, in Hypiholoma. Up to the
present no purple-spored species with decurrent gills has been
found to occupy the place of Clitocyhe, which is so largely
represented in the Leucosporae, and Pleurotus has also no
analogue. As for the rest, we have Collylia replaced by
Fsilocyhe, Myccna by Psathyra, and Omphalia by Dcconica.
But the same process cannot be applied to the veritable black-
spored species, unless we include the deliquescent Coprinus,
and can, by a little stretch of fancy, find in the pseudo-volvate
species some approach to Amanita, in Anellaria a suggestion
of Lepiota, in Panacolus a suggestion of Collyhia, and in
PsathyreUa of Myccna. At the best these are little more than
fanciful analogies. The combined brown and black -spored
species do not exceed a total of 630 species.
It will be observed that throughout the whole of this long
series of about 5200 species we have but one type, with all
its modifications, of a pileate Fungus, with a stem, sometimes
nearly obsolete, supporting a cap or pileus, bearing on its
inferior surface the radiating folds of a hymenium, on which
the basidia support four naked spores. All the groups,
divisions, subdivisions, genera, and species are dependent upon
the variations in this common type. There need, therefore, be
no surprise that a cultivated eye and experience are essential for
the accurate discrimination of distinctions, often so subtle as to
puzzle the young student and bewilder the casual observer,
whose knowledge has never extended beyond a soft fleshy
thing with a stem supporting a cap with parallel radiating
gills.
The second group of the Hyiiicnomycctcs is the Polyporci, in
which the hymenium is still inferior, but is no longer repre-
136 INTRODUCTION TO THE STUDY OF FUNGI
sented by a folded membrane, in the form of parallel plates or
gills. Instead thereof the gills are replaced by parallel tubes,
more or less adnate to each other, presenting a surface
punctured with an infinity of minute pores, sometimes as small
as if pricked with a pin. The membrane which lines the sides
of these tubes or pores is the hymenium, so that the spores
are produced within the tubes and not so fully exposed as in
the Affaricincae. If we take as a type one of the species of
the soft and fleshy genus Boletus, we shall see that in some re-
spects it resembles the ordinary mushroom, and at the same time
detect its more prominent differences (Fig. 51). In the presence
of mycelium, stem, and cap the Boletus agrees with the Agarics,
but the section will show the parallel tubes replacing the gills.
But this is not so complete a type of
the whole Pohjporei as was the Agaric
of the Agaricini. In the first place
the stem is often absent and the cap
or pileus sessile, attached by the
margin. And, in the second place,
the pileus, or what corresponds to the
pileus, adheres to the matrix by its
whole upper surface, and only the
hymenium, or pore -bearing surface, is
exposed. This resupinate condition is
Fig. 5i.-5o^d«,9, with section ^^^.y common, SO that the essential
and spores. •'
character is a porous hymenium, seated
upon the least possible development of a pileus. And yet,
as far as practicable, the hymenium is inferior, or turned away
from the light. We are prepared, then, to meet with a greater
variety of form than in the Agaricini, as well as greater ex-
tremes of texture.
The nearest approach to Agaricini in habit is to be found
in the four genera Boletus, Strohilomyces, Boleti^ius, and Gyrodon.
All these were, in their earlier history, associated together as
Boletus, but at length came to be dissevered and recognised as
distinct genera. Strohilomyces is Boletus with a scaly pileus ;
Boletinus is Boletus with short, large, radiating pores; and
Gyrodon is Boletus with elongated, sinuate, irregular pores.
All of these are fleshy but firm, soon putrescent, but mostly
HYMENOMYCETES 137
-with the tubes of the hymenium adhering so slightly to the
under side of the cap that they are easily removed. Otherwise
expressed, the trama of the pileus does not descend into the
tubes, the mouths of which constitute the pores. In all of
these a stem is invariably present, and sometimes a manifest
ring, but without a volva. They are usually of a robust habit,
sometimes attain a very large size, and are wholly terrestrial.
Like the fleshy Agaricini, they are most prolific in temperate
regions, being replaced in the tropics by woody Fomes or
leathery Polystidi, as fleshy Agarics are replaced by Lentinus
and Lenzites. The whole total of described species is some-
what under 300.
Fistulina has a similar fleshy substance, but the stem may
be present or absent, and the tubes are not laterally adherent,
and nearly all the species grow upon decayed wood.
The genus Polyporus, as originally characterised by Fries,
was a large one and spread all over the world, but in recent
times it has shared the fate of other large genera, and been
subdivided into the genera Pohjporus, Fomes, Polystidus, and
Porta, as suggested by Fries in one of his latest works. These
appear to be well defined, and no difficulty will be found to
occur practically in their discrimination. The original name is
retained for the section Anodcrmei of the old, undivided genus.
The pileus is at first soft and fleshy but tough, becoming
indurated, rarely fragile, without furrows or zones on the
pileus, and with only a single stratum of tubes, so that practi-
cally they are not perennial. Some of the species have a
central stem, and then resemblf" Boletus, only that the trama
of the pileus is continuous with » ^e tubes, which are not easily
separable from the flesh of the cap. Other species have a
lateral stem, or even a common stem, much divided above, and
bearing several pilei (Fig. 52). Finally, other species have no
stem at all, and the pileus is broadly attached to the matrix, so
as to be semi-orbicular or kidney-shaped. Earely the pileus is
reduced to a thin stratum, adherent by its whole surface, as in
Poria, but with a slightly reflexed margin. In Fomes the
substance is woody from the first, becoming very hard, and
covered with a rigid crust ; not truly zoned, but becoming con-
centrically sulcate. The substance is floccose and interwoven,
138 INTRODUCTION TO THE STUDY OF FCNGI
often zoned, and the tubes are typically stratose, each stratum
being the growth of a year, so that the species are truly per-
ennial. Some have a central, others a lateral stem, but most
commonly they are attached by a broad
'"'* '" H /^ base, where they are very thick, and not
uncommonly of the shape of a horse's
hoof, or more rarely several pilei grow
together in an imbricate manner. The
species of Polyporus, as now restricted,
t^^enerally shrink and become contorted
m the process of drying ; but in Fomes
the substance is so rigid that no shrink-
ing or alteration of form takes place,
and, except for the depredation of in-
FiG. h2.-Poiyporus, with gg^j- ^^^^ |^^ i^g preserved unaltered for
conimoii stem. ° ^
a century. Such species as Fomes
cornubovis, when sawn through, resemble sections of buffalo-
horn, although generally the internal substance is more
fibrous.
Puhjstidus includes thinner, smaller, and more delicate
species, which are of a somewhat tough and leathery consist-
ency, usually flexible, and either hairy or velvety, or becoming
smooth. The surface of the pileus may be concentrically
sulcate, normally zoned, Init not encrusted. The intermediate
stratum is fibrillose, passing down into the hymenophore, so
that the tubes are not separable. The latter are short,
and developed from the centre towards the circumference.
Commonly the whole thickness behind does not exceed a
quarter of an inch, often less, sometimes more ; but the pilei
may be confluent laterally, or densely imbricated, and the
hymenium may run down the matrLx for a considerable dis-
tance. The habit and appearance often closely resemble
species of Stereum. Additional to the sessile species, there are
some which have a central stem, others with a very short
lateral stem, expanded at the base into a sort of disc, for attach-
ment to the matrix ; but most of the European species are
sessile, extended at the base, and more or less imbricate. The
pores are very variable in size in different species ; in some
they are so small as scarcely to be visible to the naked eye,
HYMENOMYCETES
139
\v others they may he broad and shallow. The walls or
dissepiments are normally thin, even so thin as to split down-
wards very readily in process of growth, leaving little appear-
ance of pores, except at the base. In some species the edges
of the pores are fringed and toothed, so as to resemble Ir-pex.
The last group of those which constituted the old genus
Pohfporus, contains the resupinate species, under the name of
Poria. The pileus is reduced to a thin stratum, mostly spread
over, and adhering closely to the matrix, the outer surface
covered with the crowded pores. In habit the resemblance is
to Corticmm, but instead of a smooth horizontal hymenium it
is a porous one. Normally the pores are in a single series,
whilst in resupinate forms, or species, of Fomes they are thicker,
firmer, and stratose.
Allied to Fomes rather than to Polystidus, the genus
Trametes is to be recognised chiefly by the thick obtuse
dissepiments of the pores,
the tubes deeply sunk into
the substance of the pileus,
and not stratose, and with- 1
out the hardened crust to
the pileus. The pores are
rounded and often unequal,
whereas in Sderodepsis they
are large, sometimes an-
gular, with the edge acute
or toothed. Daedalea is
in substance and general
appearance very like
Trametes, with the pores
sinuous or labyrinthiform
(Fig. 53). Hexagonia rather
approaches Polystidus than
Trametes, but the pores ar
with firm entire dissepiments
Fig. 53. — Daedalea quercina.
mostly large and hexagonal,
Favolus differs from Hexagonia
in the pores being less hexagonal, but angular and radiating
from the stem ; most species being substipitate, and fleshy
rather than rigid. In Laschia the substance is still softer,
and more gelatinous, whilst the dissepiments are vein-like
140
INTRODUCTION TO THE SI ^
NCI
and the pores shallower and irregular. The latter gc""\us
leads to Mcridius, with its soft, waxy hymenium, the surface
of which is reticulated with obtuse folds, forming irregular
areolae, the folds sometimes rather toothed. This is possibly
the lowest and most imperfect of the Polyporei. A recent
genus, Campbellia, is a higher development, with a pileus and
stem and more distinct pits or pores. Porothdmm has the
habit of Porta, but the tubes are more scattered, reduced to
papillae, and at length pierced and open. Some authors add
Solenia to the Pohjporei, whilst others have associated it with
Cyphella in the Thelcpliorei.
The third primary group of the Hymcnomyccteac is the
Hydnci, in which the gills of the Agaricincae and the tubes of
the Polyporei are replaced by teeth or spines, the outer surface
of which is clothed with the hymenium, which is therefore
wholly exposed. There is at no time, and in no known species,
any kind of veil covering the hymenium in its early stage.
The most typical genus is Hydnum, which remains much the
same as Fries left it, although there has been more than one pro-
posal to split it up into smaller genera (Fig. 54). In the stipitate
species some have a central stem, others a lateral stem, and in
others the common stem is
branched and subdivided, but
the pilei are imperfect. In
another section there is no
stem, but the pileus is sessile
or imbricate, and there are a
large number of species which
are as entirely resupinate as in
the porous genus Poria, to
which this section is analogous.
There is also considerable
difference in texture, some being fleshy, others waxy or
leathery, and others becoming quite hard and corky. At
one time a gelatinous species was included, but this has
been removed on account of its affinity in fructification with
Trevidla. The teeth are variable in length and thickness in
different species, but they agree in being more or less pointed
at the apex, and free from each other at the base.
o
— Section oi Hydnum repmidui
MENOMYCETES
Fig. 55. — Radulum.
' A nunibe'^'^^^i^^ai'iall genera are associated together in this
family, t\\rciuiiv; 'xcations consisting cliiefly in the teeth or
spines ; for luotance, in Irpex the teeth are flattened at the base,
and connected so as to form irregular pits ; and in Badulum the
teeth more resemble obtuse tubercles,
and are often distorted (Fig. 55).
In Phlebia, an aberrant genus, the
hymenium is corrugated, with fold-like
crests, so as to resemble Auricularia
almost as much as anything else, and
Hydnum scarcely at all. Then in the
wholly resupinate genera, Ghundinia
has the hymenium granular, Odontia
has the granules or warts crested, and
in Knciffia the hymenium is clad with
rigid setae. Except in Mucronclla,
and probably KneiJJia, the basidia are tetrasporous. The whole
family does not include more than about 470 species.
The last of the four primary families of Hymenomyceteae
which have an inferior hymenium, is the Thelephoreae, which
nearly corresponds to the section Auricularini of Fries, with the
exception of the genus Auricidaria, transferred to Tremellineae.
The hymenium is typically even, but rarely rugose, approaching
the Hydneaceae by such genera as Cladoderris and Beccariella,
in which the hymenium is veined, and the veins are warted
or almost aculeate. Mr. G. Massee has intimated ■^ that " the
Thelephoreae constitute the base, and also the starting-point, in
the evolution of the Hymenomycetes, and, further, that from
the Thelejjhoreae all the other orders have directly originated."
In this family, as in the others, the species are variable in
form as well as in texture. Only in Cratcrellus is the substance
fleshy, attenuated in some species to membranaceous, often with
a central stem and a funnel-shaped pileus, the outer or under sur-
face being clothed with a ribbed or rugose hymenium (Fig. 56).
Cladoderris and Beccariella are tough and leathery, mostly fan-
shaped, sometimes funnel-shaped, but with a warted hymenium.
In Thelephora the substance is tough, but softer and more
p. 112.
" Monograph of the Thelephoreae," by G. Massee, in Linnean Journal, xxv.
INTRODUCTION TO THE ^t^uDY OF FUNGI
Si
Alii
1
—Craterellus cornu-
cojdoides.
spongy, without distinct cuticle to the pilei. rpj^^ n Ji|termediate
stratum ; hence homogeneous, the hymeniui,-^^^^ ^^^^ ven or a
little ribbed. The fornl is ariable,
from stipitate and funnel-shaped to
closely adnate and resupinate. It is
notable that in the majority of species
^S^''^^^" ' ' "^"-'' // the spores are globose and rough, mostly
\ // slightly coloured. We are disposed
to place here the genus Lachnodadium,
which some authors include in the
Clavarieae, on account of the erect,
branched habit, resembling some species
of Clavaria, forgetting that there are
also erect, branching species of Tlicle-
pliora, to which these species of Lachno-
dadium are closely allied in texture rather than to fleshy
Clavariae. Stereum in form approaches Thelephora, but the
substance is firmer, more leathery, and the pileus has a distinct
outer stratum analogous to that in
Polystidus, with an intermediate
stratum, and a smooth, even hymenium
(Fig. 57). Closely resembling in
appearance is Hymenochacte, with the
exception that the hymenium is
velvety, with processes resembling
bristles. With the exception of Skep-
peria, in which the pileus is vertical,
most of the remaining genera are
wholly resupinate. These are : Conio-
phora, in which the effused substance
is membranaceous and smooth, with coloured spores ; Corticium ,
in which the effused substance is usually thicker and firmer,
but without an intermediate stratum, the hymenium smooth and
rather waxy, and the spores uncoloured ; Peniophora, with the
habit of Corticium , hut with a velvety hymenium; and Hypochnus,
with the habit and appearance of Corticmm, but with the sub-
stance softer, floccose, and more lax, and the hymenium less com-
pact, but still the spores are uncoloured. To these must be added
the small genera — Aleurodiscns, with a somewhat saucer-shaped
Fig. 57. — Stereum hirsutum.
hymenomycetes 143
pileus, and pec ..j.^Aties of structure which prohibit its unison
with Corticium , ' Michenera, with a placentiform habit, a waxy
JijmeniuE^, and pedicellate spores ; and Exobasidium and Helico-
hasidium, which are encrusting and waxy, growing upon living
plants, and distorting them. Finally, Cyi^hclla, having the form
of Feziza but the fruit of Corticium, being in fact a cup-shaped
Corticium ; and Solcnia, the cups of which are elongated into
tubes, so that it seems doubtful whether they should be placed
in relationship with Poria, in Polyporeae, or with Cyphclla in
Thcleplioreac.
Briefly and succinctly, these are the principal genera of
Thclephoreae, but before dismissing them we must advert to
certain appendages of the hymenium which
distinguish some of the genera above
enumerated. In addition to the basidia
there are to be found in the genus Pcnio-
phora stout projecting cells, which are either
the modified cystidia, or analogues of cystidia,
but which have been called metuloids.
They are fusiform, colourless, and at first
smooth, but afterwards rough and brittle ri«- 58.-Cystidia of
° Peniophora.
from the deposit of oxalate of lime on
their surface. These are conspicuous objects upon the otherwise
smooth hymenium, giving it a velvety appearance, and by
this character separating the species from Corticium. In
another genus, that of Hymenocliaetc, the same place and posi-
tion on the hymenium is occupied by projecting, acute, non-
septate brown bristles, which spring from the hyphae of the
subiculum, and impart also a velvety appearance. Exter-
nally in habit the species resemble Stercum, but they are
readily distinguished by the presence of these brown project-
ing bristles. A similar kind of appendage to the hymenium
has been detected in some species of the Polyporei, for which
the generic distinction of Mueronoporus has been proposed.
In a small section of the genus Hymenochaete, according to
Saccardo, but generically separated by Cooke and by Massee
under the name of Veluticeps, the hairs of the hymenium are
produced generally in bundles, and are flexuous and septate,
in which respect they differ from the setae of Hymenochaete.
144 INTRODUCTION TO THE STUDY OF FUNGI
A type of structure iu the hyphae of thVrrnsal stratum in
some species of Corticium is worthy of note, as aflbrding a
means of discrimination in allied forms ; and this constitutes
the basis of a new genus, proposed by Massee, under the name
of Asterostoma. The species alluded to are distinguished by
the brown stellate hyphae that are present in the subiculum,
"Erect branches at about the level of the base of the basidia
develop at the apex a stellate arrangement of branchlets, all
situated in one plane, parallel to the surface of the hymenium ;
the number of rays varies from three to seven, five being the
most frequent, and differ from the supporting hyphae in being
aseptate, with very thick walls, which soon become bright
brown. When the spores are ripe, the erect hy23hae, sup-
porting both stellate threads and basidia, along with the latter
disappear, leaving the coloured star-shaped bodies mixed with
the spores, resting on the horizontal interwoven basal stratum
of the plant." An analogous differentiation is pointed out as
existing in the basal stratum of Bovista, in the Gastromycetes.
Thus, then, we close our remarks on the first four primary
sections or families of the Hymenomycetes, in which the
hymenium is normally inferior, and either spread over radi-
ating gills, lining the cavities of tubes, investing teeth, warts,
or projections, or finally forming a plane, even, or nearly even,
fructifying surface.
The fifth family, or Clavaricae, has a vertical hymeno-
phore, with the hymenium on all sides, and not distinct from
the stem. Sometimes the entire Fungus is a simple club, and at
other times it is much branched, with the lower portion barren,
forming the stem, and the upper portion fertile, covered with
the even or wrinkled hymenium. In most genera the substance
is either fleshy or waxy, rarely somewhat gelatinous. The most
highly developed genus is Sparassis, in which the branched
hymenophore has the branches flattened into leaf-like expansions.
The largest genus, however, is Clavaria (Fig. 59), in which the
hymenophore is club-shaped and simple, sometimes solitary and
sometimes in clusters or branched, often very much branched,
but always fleshy. In Calocera the form is similar, but the sub-
stance is toughly gelatinous, becoming horny when dry. The
species of Clavaria are for the most part terrestrial, those of
H YMENOM YCE TES
145
Calocera usually growing on dead wood. Saccardo includes also
Zachnpdadium, which resembles a branched Clavaria, but the
substance is coriaceous, and the stem
tomentose. For these and other
reasons we prefer to place it in
ThelepJwreae. In Fterula the sub-
stance is dry and cartilaginous, but
in form resembling very slender
Clavariae. Typhula and Pistillaria
include minute species, mostly waxy
and delicate, in the former with a
very long, and in the latter a very
short stem. Physalacria is Pistillaria
with a subglobose, vesicular head or
capitulum. Most of the species in
this group are white, whitish, or
brightly coloured, and but few of
them attain any considerable size.
The spores are simple, small, and
either uncoloured or yellowisb.
The sixth, and last, family is the
TrcmcUincac, in which the dis-
tinguishing feature is the tremelloid
substance, collapsing when dry and ''^ ■
reviving with moisture, combined with fig.
a peripherical, somewhat peculiar,
basidiosporous fructification. The basidia are not super-
ficial, but immersed, and either undivided or forked at
the apex, or globulose and cruciately divided. The spores
are typically reniform or globose and continuous, and these
on germination give rise to sporidiola. The structure of
this family was investigated at first by Tulasne, but more
recently by Brefeld, and the classification now adopted
is based chiefly upon the records of the latter. In this
manner three subfamilies have been recognised — viz. the
Auricularieac, in which the basidia are elongated or fusoid,
and transversely many-celled ; the Tremcllincae, in which the
basidia are globose, or nearly so, and when mature divided,
in a cruciate manner ; and the Dacnjomyccteac, with the basidia
10
J9. — Clavaria jnstillaris.
146
INTRODUCTION TO THE STUDY OF FUNGI
clavate, forked at the apex, each limb furnished with a single
spicule. Under these three subfamilies the different genera
are located after the following manner. (1) The Auricularieae
includes the typical genus Auricularia
(Fig. 60), in which the Fungi are
leathery and somewhat resemble Stereum,
but with a gelatinous hymenium, which
is veined in a reticulate manner. (2)
Hirneola differs in the substance being
more membranaceous, and often cup-
shaped or ear-shaped, becoming carti-
laginous when dry ; the gelatinous
hymenium being either even or plicate.
Fig. QO.-Auriculariamesen- ^^^ (^) Pl^tVOlaea is wholly gelatinous,
terica, with section and mostly small, erumpent or superficial,
^^°^^^- either wart-like or effused. Perhaps
the nearest relation of this small group will be found in Lascliia,
amongst the Polyporeae.
The most important subfamily is that of the Tremel-
lineae, in which the basidia are subglobose. Of these, Exidia
includes a variety of forms, either discoid, cup-shaped, gyrose,
tubercular, or effused (Fig. 61); some of which are even, and
others papillose or spiculose. The
basidia are rather ovoid, immersed in
the gelatine, partite in a cruciate
manner, and typically tetrasporous.
Spores reniform, and for a long time
continuous ; at length, - preparatory to
germination, two or more celled, each
cell producing a very short filament
crowned with a narrow curved sporidio-
lium. In the genus TremcUa the form
may be pulvinate or effused, often Fig. qi.— Exidia, with section
brain-like, with sinuosities, but with- '^^ spores,
out papillae. The basidia are globose, and divided as in
Exidia, and the spores subglobose. The promycelium result-
ing from germination produces globose or elliptic sporidiola.
Conidia have been observed in some species, but neither
spores, sporidiola, nor conidia are ever septate. In form
HYMENOMYCETES 147
Ulocolla resembles Tremella, but the germinating spores
are bilocular. Naematelia also resembles Tremella, but enclos-
ing a hard central nucleus. The genus Femsjonia presents
cup-shaped or pezizoid forms, with globose basidia and curved
spores. In Craterocolla the form is less cup-shaped, but there
are two kinds — one somewhat tremelliform, bearing basidia ; the
other more regular and rather truncate, bearing conidia. In
Sehacinia the whole Fungus is effused like a Corticmm, bearing
conidia at first, and afterwards reniform spores. The genus
Gyrocephalus is analogous to Gucpinia ; the species are erect in
habit and spathulate, with basidia of the Tremella kind and
pear-shaped spores. The genus Tremellodon, with the form
of Hydnum but the fruit of Tremella, properly belongs here.
The subfamily Baeryomyeeteae includes the lowest Tremel-
loid forms, in which the basidia are clavate, or nearly of the
ordinary Hymenomycetal type, forked above, and each apex
bearing a single spicule. The genus Dacryomyeetes includes
normally small pulvinate species, the spores of which are trans-
versely or muriformly divided when mature, and the conidia
(when present) growing in chains. The genera Arrhyticlia and
Ceraeca are North American, and of minor import. Guepinia
consists mostly of irregularly cup-shaped or spathulate species,
with a more or less developed, and often woolly, stem. The
hymenium is discoid or one-sided, and the basidia linear and
bisporous. The genus Dacryomitra has the fructification of
Dacryomyces, but the form and habit are those of Typhula or
Mitrula, being minute and club-shaped. Collyria is a North
American genus, of a single species, resembling a large Dacryo-
mitra, with an inflated capitulum. Two or three other little-
known genera have been added provisionally to this subfamily,
but their position has not yet been satisfactorily determined.
Thus closes our survey of the groups, and genera, of the
Hymcnomyceteae, in which the most distinctive features have
been indicated ; but there are many cross relationships and
analogies which could scarcely be alluded to. It has been
pertinently observed that no linear arrangement can possibly
illustrate completely the relationship of the families and genera
which approach each other at various points, but it is useful
as a guide to the classification of corresponding forms.
148 INTRODUCTION TO THE STUDY OF FUNGI
BIBLIOGRAPHY
Saccaudo, p. a. "Sylloge Hymcn^niycetum," in S)jllo(je Fungorum, vols.
v., vi. Padua, 1887-88.
Cooke, M. C. Handbook of British Fjiugi. Edit. II. " Hymenomycetes."
8vo. London. 1888-92.
Illustrations of British Fungi. "Hymenomycetes." Roy. 8vo. Coloured
plates, 1882-92.
A plain and easy Account of British Fungi. 12mo. Coloured plates.
London, 1870, etc.
Edible and Poisonous Mushrooms. 12mo. Coloured plates. London, 1894.
British Edible Fungi. Sni. 8vo. Coloured plates. London, 1891.
Cooke, M. C, et Quei.et, L. Claris Synoptica Hymenfymycct^hm Europa^orum.
8vo. London, 1878.
Stevexsox, Jxo. Hymenomycetes Britannici. {British Fungi — Hymeno-
mycetes.) 2 vols. 8vo. Edinburgh, 1886.
Fries, E. Epicrisis Systematis Mycologici, scu Synopsis Hymenomycetum. 8vo.
Upsal, 1836.
Rones Selectae Hymenomycetxim. Fol. 2 vols. Coloured plates. Stock-
holm, 1867-82.
Monographia Hyinencmiycetum Sueciae. 8vo. 2 vols. Upsal, 1857.
Sverigcs atliga och giftiga Svampar. Fol. Coloured plates. Stockholm,
1861.
Gillet, C. C. Lcs Chamjngnons de France. "Tableaux analytiques des
Hymenomycetes." 8vo. Coloured plates. Alen^on, 1878-94.
Kalchbrenker, C. Icones Selectae Hymenomycetum Hungariae. Fol. Coloured
plates. Budapest, 1873.
Massee, Geo. "Monograph of the Thelephoreae, " in Journal Linnean Society.
8vo. London, 1889-91.
Persoon, C. H. Synopisis Methodica Fungorum. 8vo. Gottingen, 1801-8.
QuELET, L. Les Champignons du Jura et des Vosges. 8vo. Plates. Montbellard,
1873-75.
Heese, H. Die Anatomic der Lamelle und ihrc Bedexdung fur die Systematik der
Agaricineen. Roy. 8vo. Berlin, 1883.
Favod, M. V. "Prodrome d'une Histoire Naturelle des Agaricines." Ann. des
Sci. Kat., 7th series, vol. ix.
Patouillard, N. Les Hymenomycetes d' Europe Anat. gener. Paris, 1887.
Britzelmayr, M. Die Hyinenomijceten Augsburgs, 8 (1879), Hymenomyceten aiis
Sildbayern. 8vo. Plates. Augsburg, 1885.
CHAPTER XIII
PUFF-BALL FUNGI GASTROMYCETES
Every schoolboy is supposed to know what a puff-hall is, and
therefore they may be accepted as a type of the peculiar order
of Fungi to which this chapter is devoted. During the summer
the little white puff-balls, growing in the grass of pastures and
on heaths, resemble small snowballs, soft and spongy, and
scarcely tinged with colour in the pulpy interior. As autumn
advances the outer surface at first becomes creamy or ochra-
ceous, covered with small warts or spines, which are readily
rubbed off with the fingers. Later on the colour becomes
brownish, the coating is split irregularly, or opens with a
round mouth, and the interior is seen to be filled with a fine
olive or purplish powder like snuff, mixed with delicate threads,
called the capillitium. Such are the ordinary pufi-balls which
schoolboys puft' in each other's faces, the distinguishing feature
being that the myriads of minute spores are wholly enclosed
at first within the outer case or peridium, and remain so until
mature, when the coating is ruptured. The Gastromycetes,
therefore, are Fungi which, as a rule, produce their spores at
the apex of basidia wholly enclosed within the substance of
the Fungus. They constitute a portion of the Basidiomycctcs,
because the spores are developed on basidia, but are specially
denominated Gastromycetes, because the basidia and spores are
not exposed, as in Hymenomycetes. If an ordinary Lyco-perdon
be cut downwards through the centre, it will be observed that
the basal portion is cellular and does not contain spores ;
moreover, in some species this sterile portion projects upwards
into the interior as a columella — which, however, is not always
present. The peridium or outer coating in this instance is
ISO INTRODUCTION TO THE STUDY OF FUNGI
double, the exterior warty, spinulose, or powdery, and the inner
paper- like. In other genera, as in Bovista, the two coats are
more distinct. In Geaster they are still more distinct, the
outer peridium splitting in stellate rays. Thus much for the
general character, but the varied modifications must be noted
hereafter.
In some concise observations on this group Mr. Massee
remarks that no sexual organs have been observed, but he
alludes to the peculiar form of coalescence between two hyphal
cells under the name of clamp-connections, which are not un-
common (Fig. 62). "A slender lateral branch," he says, " springs
close to a transverse septum separating two superposed cells,
and, after growing for some time, its tip comes in contact with
the wall of the adjoining cell just beyond the
septum, absorption of the walls takes place at
the point of contact, and thus at first an open
communication is established, by means of the
lateral branch, between the two adjoining cells ;
at an early period this channel of communication
is usually interrupted by the appearance of a
septum at the point of origin of the lateral
branch, and a second septum is in some in-
stances formed at the point of contact with
Fig. 62.— Clamp- ^^le sccond Cell. The lateral branch is usually
counections. "^
closely pressed to the hypha from which it
springs, but sometimes becomes arched and free from the
hypha between the two points of attachment." ^
Eeference is also made to the differentiation of the hyphae
which are contained within the peridium. In the gleba, or
hymenial pulp, of the Zycojmxlaceae " at a very early period
two sets of hyphae are present. One, thin- walled, colourless,
septate, and rich in protoplasm, gives origin to the trama and
elements of the hymenium, and usually disappears entirely
after the formation of the spores ; the second type consists of
long, thick-walled, aseptate or sparsely septate, often coloured
hyphae, which are persistent and form the capillitium. The
latter are branches of the hyphae forming the hymenium."
There are three somewhat aberrant groups which offer
^ Massee, Monograph of British Gastromyces, p. 4.
PUFF-BALL FUNGI— GASTROMYCETES 151
consideralile variations in structure from the genuine puff-
balls. These are the Phalloideae, the Nididariaceae, and the
subterranean or Hypogeae. The latter come in for notice
under the head of " subterranean Fungi."
The Phalloideae, or stink-horn Fungi, have mostly a very
fetid odour, and instead of enclosing within themselves pul-
verulent spores mixed with threads, present externally a
gelatinous mass of agglutinated spores, which is collected upon
some superior and exposed surface. The whole number of
described species in this family is about eighty, and they are
most common in warm climates. Some are stipitate and
others clathrate or latticed, but all are at first enclosed in
a general volva of an egg -shape, with a gelatinous inner
stratum. The entire plant is of a soft, watery texture, quick
in growth, and rapid in decay. As the gelatinous dark-
coloured mass of the hymenium is greedily devoured by
insects, it is reasonably assumed that it is by this agency that
the spores are dispersed.
Mr. T. Wemyss Fulton devoted some attention to this
subject,^ and the following is a digest of his observations, en-
tirely confined to the common stink-horn, Itliypliallus impiidicus,
which grows freely in woods and gardens : — " The hymeno-
phore or reproductive portion consists, in its earliest stages, of
minute swellings, which arise on the underground mycelium.
These at first are homogeneous, but gradual differentiation
goes on, so that towards maturity the following parts may be
recognised. (1) An enclosing cortical portion, the volva or
peridium, composed of three layers — an outer firm skin, an
inner thin membrane, and an intermediate gelatinous layer.
At the base there is a cup-shaped portion, which supports the
stem, and is continuous by its margin with the peridial
layers, and below with the mycelium. (2) A central medullary
portion, composed of two very different structures, the gleba
or spore-bearing part, which forms a hollow conical cap, lying
within the inner peridium, and surrounding the upper portion
of the stem, to the apex of which it is attached. Its outer
surface bears the hymenium, and is honeycombed by a number
1 " Dispersion of Spores of Fungi by Agency of Insects, with special reference
to the Phalloidei," by T. Wemyss Fulton, in Annals of Botany, vol. iii., 1889.
INTRODUCTION TO THE STUDY OF FUNGI
of irregular depressions, in which the mass of spores is lodged.
The stem, consisting of a cylinder whose walls at this stage
look firm and solid, is composed of a multitude of small com-
pressed cells filled with jelly.
" The volva is at first concealed beneath the surface of the
soil, but towards maturity it breaks through the ground, and
the exposed part gradually becomes conical, and finally
ruptures, the stem rapidly lengthening and elevating the
gleba in the air. The gelatinous contents of the flattened
cavities disappear, and they become dilated, the previously
compact stem increasing threefold or fourfold in magnitude,
and becoming open and spongy, the cavities .being distended
with air. The elevation of the gleba takes place with great
rapidity, and may be completed in half an hour or from two
to three hours, attaining a height of from six to ten inches.
The utility of this sudden elevation by a mechanical process,
instead of the slower process of simple growth, will hereafter
be evident (Fig. 63).
" At the time of emergence, and for a brief interval after,
the hymenial surface is firm and solid, greenish
gray in colour, and emits a faint, mawkish, but
sweetish and honey-like odour, which is attractive
to house-flies. Very soon, and before the elonga-
tion of the stem is completed, it begins to darken,
the odour becomes fetid, and the consistency
changes so that it gets rather sticky and tenacious.
A little later it is dark green, almost black, the
odour is strong and repulsively fetid, and in con-
sistence slimy or almost fluid. These changes
begin at the apex and proceed downwards ; they
seem to depend largely upon the influence of
light, for if one side be protected from its action
the change in consistency and colour is retarded
i__ui,,j. on that side. When examined microscopically the
dius^ ivi- fg|^j(-[ ^^j(j jg gggjj ^Q contain myriads of spores
(each 3 /x. long). These changes occur during
hot months of the year, from the early part of July till
the end of September, at a time when insect life abounds.
" As soon as the strong, dung-like odour is develoj)ed, the
PUFF-BALL FUNGL—GASTROMYCETES 153
liquefying hymeniuin is visited by large numbers of flies,
which sometimes almost cover it, and suck up the fluid mass
with great avidity during hot sunny days ; but when the
weather is cloudy or cold, fewer flies are to be seen."
Examinations were made of flies taken from the deli-
quescing gleba, and thousands of spores were found adhering
to the feet and proboscis. The flies placed in confinement
showed that their excrements were almost exclusively com-
posed of spores. To determine if the excrementary spores
retained their vitality they were placed in tubes on sterilised
earth. The tubes were then closed with cotton -wool and
buried with the contained spores, and different substances
with them. In about two months the spores had germinated,
and in some produced a plentiful mycelium. Hence it is
clear that the spores after passing through the stomachs of
insects do not lose their power of germination.
This family is remarkable for the prevalence of a bright
red colour near the hymenium, and also for the peculiarity of
many of tlie forms. In Bidyo'pliora., Ithyphallus, and Mutimis
the form is columnar and phalloid ; in Clathrus and Colus it
is clathrate. In Calathiscus and Aseroe the disc is stellate,
and in Kalclibrcnnera it is coralloid. The spores in all the
species are very profuse and minute, generally involved in
mucus.
In the family of Nidulariaceae we meet with other
peculiarities, and of these the common species of Cyatlms or
Crucihulum, called the " bird's-nest Fungus," may be taken as the
type (Fig. 64). There are altogether only about sixty described
species, and the family, under some of its forms, is pretty
widely distributed. When mature the Fungus is not more
than from one to two centimetres high, and resembles in form
little inverted bells, at first covered across the mouth with a
white membrane or operculum, which when ruptured exposes
a number of lentil-shaped bodies, packed like eggs in a little
bird's-nest. These are the peridioles, each of which is
attached to the inner surface of the cup by a long elastic
cord, proceeding from the under face of the sporangiole.
Each of these sporangioles, when cut in section, reveals a
central cavity, into which the basidia project, with their
INTRODUCTION TO THE STUDY OF FUNGI
attached spores. These lentil-shaped bodies are analogous to
the peridiola in such genera as Polysaccvm and Arachnion,
but all the inter-
vening plasma has
been dissolved away,
so that they remain
free within the peri-
dium. In all the
species the sporan-
gioles are very hard
I |-'.^ "^(^Hj^^^ and firm when
mature, and the con-
tents are never
powdery. In some
species the external
peridium has a squa-
mose or hairy surface
but in a few species
it is nearly smooth.
Sometimes the upper
third of its length
is marked with con-
spicuous parallel
channels or striae.
In Cyathus the peri-
dium is composed of three superimposed layers, and in
Crucibulum of two.
Having disposed, in a summary manner, of these two
families, we return to the Trichogasters which form the bulk
of the order, and especially the Zycoperdaceae. Probably the
genera which contain the largest number of known species
are Lycoperdon, Geaster, and Bovista. In these the peridium
is more or less distinctly double, but there are allied genera
in which the peridium is simple. The delicate threads, found
mixed with the spores when mature, forming the capillitium,
are an important element in classification. In the mature
gleba they seem to be entangled, and indefinite as to their
origin in Lycoperdon and Bovista, but in other genera they dis-
tinctly radiate from the columella to the inner wall of the
Fig. 64. — Ciucibuluni tubjait. Aftei Greville
PUFF-BALL FUNGI— GASTROMYCETES
155
Fi(i. 65. — Lycuperdon, with
sterile base and columella.
peridium (Fig. 65). This columella is only a continuation of the
spongy base in Lycoperdon, but in Diplodcrma it is hard and
woody. In some species of Geasfpr
the columella is distinct and club-
shaped, extending half-way up, with
the threads of the capillitium radiat-
ing towards the periphery. De B<iiy
has described the complex peridium
of Geaster in the following terms : —
" Geaster hygrometricus is up to the
period of perfect maturity a roundish
body, which may be of the size of a
hazel-nut, and remains beneath the surface of the ground.
Six layers may be distinguished in the peridium in a vertical
longitudinal section a sliort time before the compound sporo-
phore is mature (Fig. 66). The outermost layer is of a brownish
colour, flaky and fibrous, and is continued on one side into the
mycelial strands which spread through the soil, and on the
other passes into the second layer ; a thick, stout, brown
membrane entirely covering the compound sporophore. This
is followed towards the inside by a white layer, which is
more largely developed at the base of the compound sporo-
phore than elsewhere, and is immediately continuous at that
spot with the inner peridium
and the gleba. Both of these
last-mentioned layers are formed
of stout, closely -woven hyphae
running in the direction of
the surface, and may be com-
bined under the name of the
fibrillose layer. The inner of
the two is lined on the inside
by the collenchyma layer, ex-
cept where its basal portion passes into the gleba. This
layer is cartilaginously gelatinous, and consists of hyphal
branches of uniform height, connected together, without
interstices, which are placed palisade -like vertically to the
surface, and are bent as they spring from the hyphae of the
fibrillose layer. The strongly-thickened stratified walls of the
Fig. 6Q.— Geaster.
156 INTRODUCTION TO THE STUDY OF FUNGI
cc41s of this layer have great capacity for swelUng. Inwards
from the collenchyma is a white layer, the innermost region
of which is the inner peridium, while the outer, which may
be called the split layer, consists of soft, loosely-woven hyphae,
which pass at many points into the inner peridium. AVhen
the fungus is (juite matured, the outer peridium, through the
influence of moisture and the swelling of the collenchyma
layer, bursts outwards from the apex in a stellate manner,
forming several lobes, which turn back, so that the upper
surface, which is covered by the collenchyma, becomes convex.
The split layer is by this means so torn to pieces that its
constituent parts remain hanging as perishable flakes, some to
the collenchyma, some to the inner peridium. It is known
that the collenchyma layer retains its hygroscopic qualities a
long time, and the outer peridium remains a long time lying
on the soil, stellate in shape, spreading out its rays in moist
weather, and bending them inwards when dry. The flaky
investment of the outer peridium is often more strongly
developed in G. fimhriatus and G. fornicatus, and in the latter
it is composed of the finest of hyphae ; it tears away from the
fibrillose layer when the peridium is ruptured, and lies on the
ground, beneath the peridium, as an open empty sac. The
extremities of the lobes remain for the time firmly united
to the margin of this sac, and as the collenchyma layer
expands greatly, the star formed by it and the fibrillose layer,
especially in G. fornicatus, becomes convex upwards, and
carries the inner peridium on the apex of the convexity." ^
The genus Calostoma, when carefully examined, shows many
points of affinity with Geaster. INIassee has given full details
of its morphology,^ w^iich should be perused in extenso, but the
following is a summary. On the authority of Hitchcock it is
stated that in Calostoma cinnaharimim the Fungus on bursting
from the soil is enclosed in a gelatinous envelope, like
IthyphaUus imjmdiciis, nearly a quarter of an inch in thickness.
This immediately bursts, even before the whole body of the
Fungus has risen above the ground, and the exterior pait of it
^ De Bary, Fungi, etc., English edition, p. 316.
- Massee, " Monograj^li of the Genus Calostonui," in Annals of Botany,
vol. ii. p. 25, 1888.
PUFF-BALL FUNGI— GASTROMYCETES 157
falls upon the soil around the Fungus in tlie form of a viscid
jelly, and is ere long absorbed in the earth. The short stem-
like base arises from a few firm white mycelium strands,
composed of thin-walled, sparsely septate, branched threads.
After removal of the external gelatinous volva a vertical
section shows an external colourless zone, separated from the
inner portion, except at the base, by a thin red line. The
outermost zone is composed of thick-walled, mostly aseptate,
densely interwoven hyphae, passing through the red zone into
the central less compact portion, where they are mixed with
thin- walled, septate, branched hyphae having numerous slightly
thickened free tips.
When dry the plant is rigid and cuts like horn ; a median
vertical section in this condition shows the external wall to
consist of three distinct layers — the two outermost confluent at
the base, the innermost free below, but in contact with the
middle layer at the umbonate apex. The external layer or
exoperidium is at first continuous over every part of the plant,
and thinnest at the apex. The red streak is now seen to form
the innermost portion of the exoperidium, and at the present
stage of development exists in the form of red powder. In
the earlier condition the cells forming the red zone are thick-
walled, the substance of the walls being studded with numerous
small red granules. Eventually the walls of the cells
constituting this zone become mucilaginous and disappear,
leaving the red granules in the form of a fine powder, thus
effecting the separation of the exoperidium from the originally
homogeneous spherical weft of hyphae. The innermost portion
of the exoperidium consists of compactly interwoven, thick-
walled hyphae, not at all mucilaginous, and furnished with a
few red granules, which become rarer towards the outside, and
eventually disappear ; the hyphae at the same time becoming
thinner and thinner, owing to the diffluent walls, and at the
outside entirely converted into mucilaginous jelly.
Owing to a slight increase in length of the basal portion,
between the exoperidium and endoperidium, and continued
increase in the size of the latter, the exoperidium is ruptured
at the apex in an irregularly stellate manner, the lobes when
moistened curling inwards, and soon breaking away at the
1 58 INTRODUCTION TO THE STUDY OF FUNGI
base. Ill most species the exoperidium becomes completely-
disorganised, often remaining in the form of warts on the
endoperidium. AMien dry the endoperidium is cartilaginous
and brittle, of a dirty ochraceous colour, becoming much
swollen when moistened. It consists, when young, of thick-
walled, more or less gelatinous hyphae ; later on the thick walls
become disorganised, and present the appearance of a loose weft
of hyphae imbedded in mucilage, but in reality the apparent
hyj)hae are the lumina of the original thick-walled cells.
AVhen young the wall of the endoperidium is of equal
thickness, but during spore-formation local growth takes place
at the apex, forming a cylindrical umbo, the circumference of
which is furnished with several deep vertical furrows. At
this stage a red streak appears in the median line of each
vertical ridge dividing the furrows, these streaks being con-
tinued along the apical portion of the ridge and meeting in the
centre. These streaks extend through the entire thickness of
the wall, and form a central core down the umbo, the hyphae
becoming disintegrated as in the red zone. Eesulting from
this process is the formation of a mouth, the surrounding teeth
remaining closed until the period of dehiscence, when the
separation of the teeth takes place, the margins and inner
surface being covered with red powder. The endoperidium is
not differentiated from the exoperidium at the base. There is
no trace of a columella. The innermost layer, or spore-sac, is
yellowish white and flexible, perfectly free from the endoperid-
ium, except at the apex, where it remains attached to the inner
surface of the teeth. During spore-formation the central mass,
or gleba, is continuous with the inner wall of the spore-sac.
There are irregular cavities, and the basidia produce five or
six spores on wart-like projections at the apex. The spores
are globose at first, and colourless, then elliptical, pale,
ochraceous, and minutely warted.
When the spores are ripe the basidia and the trama dissolve
into mucilage ; the gleba contracts, but still remains attached
to the apex of the peridium ; ultimately the mucilage contracts
and dries into irregular masses, leaving the spores quite free.
In the normal mode of dehiscence the spore -sac and its
contents appear to pass out at the mouth and remain attached
PUFF-BALL FUNGL—GASTROMYCETES 159
to the teeth of the endoperidium ; hut sometimes the spores
are expelled without extrusion of the spore-sac. In all
species every part of the plant, with the exception of the
spore-sac, is perfectly rigid and cartilaginous when dry, every
part except the inner surface of the endoperidium becoming
swollen and more or less mucilaginous when moistened.
The stem-like base increases in growth when the spores are
mature.
In Battarrca De Bary has shown that the whole develop-
ment, up to the maturing of the spores, is passed while still
enclosed in the volva ; and when this is ruptured by elongation
of the stem, a portion of the volva is usually carried up on
the surface of the circular peridium, which is more or less
crescent -shaped in section (Fig. 67). Finally the peridium splits
along the margin, the upper portion falling away, and leaving
the spores exposed on the lower persistent part, from which
they are soon blown away.
In Tylostoma also the differentiation of the gleba takes
place underground. When the spores are mature the stem
elongates. This elongation is due to increase in
length of the central portion, the outer or sheathing
portion being cracked transversely, one portion
remaining below and sheathing the base, the other
forming an abrupt termination of the base of the
peridium like a collar at the apex of the stem.
One of the most interesting genera of the
stipitate forms is Podaxis, which is a native of warm
climates, being particularly associated with the
nests of Termites. In this genus Mr. Massee
contends that the spores are produced in sacs or f\\'\[y\
asci, and infers that therefore they belong system-
atically to the Ascomycetes ; but with this inference
we do not agree. Although the spores are at first
enveloped in cysts, it by no means follows that ^m. 67.—
this establishes an affinity with Ascomycetes, but Battarrm.
only an analogy.
The species of Podaxis bear an external and superficial
resemblance, in size and form, to unexpanded specimens of
Coprinus comcitus : the upper elliptical, spore-bearing capitulum
i6o INTRODUCTION TO THE STUDY OF FUNGI
being borne upon a long cylindrical stem, gradually attenuated
upwards, into and up to the apex of the capitulum, like a
columella ; the differentiation of the gleba, until the maturity
of the spores, taking place while the Fungus remains under
ground. There is an absence of the sinuous cavities, bounded
by well-defined tramal plates, so characteristic of Gadromycdes,
but from the earliest condition, according to Mr. Massee, " the
gleba presents a sponge-like structure, its very irregular walls
consisting of thin-walled, sparsely-septate hyphae, originating
as lateral branches from the hyphae forming the central axis
or the inner portion of the outer protective wall. INlixed with
the colourless, thin-walled hyphae are others which originate
from the hyphae of the axis ; these eventually become coloured,
and form the capillitium ; the thin-walled, colourless hyphae
forming the irregular walls of the gleba send into the inter-
stices numerous long, lateral branches ; these branches — the
ascogenous hyphae — are aseptate, have very thin colourless
walls, are richly supplied with granular vacuolated protoplasm,
and at the tips produce two or more short branches, which in
turn emit short secondary branches, the whole forming a com-
pact tuft ; these terminal branches differ from the parent
hyphae in being broken up into numerous short cells by trans-
verse septa ; each component cell produces a lateral outgrowth,
at first papillaeform, then cylindrical, and eventually broadly
obovate, and attached to the parent cell by a narrow neck ;
these terminal cells — the asci — after receiving all the proto-
plasm from the parent cell, are cut off from the latter by the
formation of a septum across the narrow basal portion. Owing
to the fasciculate arrangement of the terminal branches the
asci are densely crowded, varying in number from ten to fifty,
or even seventy on specially vigorous heads. The asci are
developed in succession, and it is not unusual to meet with
empty shrivelled asci, others with the spores not yet differen-
tiated, and others quite young, in the same cluster. I am
inclined to believe that the short ascigerous branches are also
produced laterally on the aseptate hyphae, but am not certain
on this point. The asci are usually constant in form and size,
but now and again an exceptionally large one may be seen,
and sometimes one or more lateral prominences disturb the
PUFF-BALL FUNGI— GASTROMYCETES i6i
usual symmetry of outline. The asci are normally monosporous,
but occasionally two spores are produced, especially in the
extra large or deformed examples, when the spores are differ-
entiated ; but before attaining their full size, and while yet
quite colourless, they escape from the asci through an irregular
slit, the latter persisting in the shrivelled form seen on examin-
ing the hymenium of mature specimens. The spores, when
mature, are broadly elliptical, or sometimes subglobose (averag-
ing 10 — 12 x 9 yLt.), perfectly smooth, and of a deep translucent
brown by transmitted light, and furnished with a single well-
defined germ-pore. When the spores are first liberated the
colour of the gleba is very pale yellow ; from this condition the
coloration passes through primrose yellow to clear brown, and
eventually dark brown, as seen in the mass. When young
the hyphae of the capillitium are colourless, straight, rarely
branched, and in this condition there is little or no indication
of the spiral marking so conspicuous at maturity ; during the
development of the gleba the capillitium threads pass through
the same sequence of coloration as already described for the
spores, commencing with pale yellow and ending with bright
brown. After the formation of the spores, the compact basal
portion, below the point of attachment of the lower margin of
the peridium to the central axis, elongates into a hollow stem,
eight to ten inches high, elevating the yet closed peridium far
above ground. The ripening of the gleba, as shown by the
progressive coloration of the spores, commences at the base, and
nearest the axis, and progresses towards the apex. When the
spores are mature, and the capillitium fully developed, the
ascogenous hyphae, with the clusters of shrivelled asci, can still
be seen, and although usually colourless, are in some instances
more or less tinged with brown. In the clusters of split
shrivelled asci are others that present no split or fracture in
the wall ; these are homologous with the so-called sterile
basidia or paraphyses. At this stage the peridium breaks
away from the stem at its lower point of attachment, the
margin being irregularly torn, when it resembles a half-
expanded Agaric ; eventually the whole of the dry and brittle
peridium breaks away, and the stem remains with its blackish-
brown mass of spores and capillitium, resembling a bulrush, the
1 62 INTRODUCTION TO THE STUDY OF FUNGI
final dispersion of the spores being efiected by the wind and
rain." ^
In seeking for the affinities or analogues of the species of
PodcLcis, Mr. Massee thinks that these should be traced through
the subterranean Gastromycctcs to the ascigerous Elaphomycctcac.
" I have shown," he writes, " the gradual conversion of the
ascigerous Tuhcraceac into the basidiosporous Hymenogastreae,
due to the changes of asci into basidia, and the subsequent
evolution of the whole of the above-ground Gastromycctcs from
the subterranean ascigerous Tuhcraceac through the Hyincno-
gastreac ; and now we find a second attempt on the part of the
Tuhcraceac to evolve an above-ground branch through the
Maphomycetcae, and continued by the genera Podaxis, Tylostoma,
and possibly Battarrea and Quclctia.
He might further have indicated that in another direction,
through Secotium, Polyjjlocium, and Montar/nitcs, the Gastromycctcs
are linked to Coprinus, and, through that genus, with the
Agaricini. Montagnites, or as sometimes called Gyrophragmium,
has in some systems been included with Hymenomycetes in a
position next to Coprimis, to which it bears some resemblance.
The Sclcrodcrmcae are a group which seem to fall into an
intermediate position between the Lycopcrdaccac and the sub-
terranean Gastromycctcs — a fact which was recognised by Mr.
Massee when he indicated that they differed from the former
in the absence of a capillitium and in the indehiscent peridium ;
and from the latter in not being subterranean, although there
are one or two species in which subterranean individuals are
sometimes to be met with. " As in the Hymcnogastreac, the
peridium is thick, usually warted or rugulose externally, and
but little differentiated, the trama springing from every part of
its inner surface. In Polysaccum the cavities of the gleba are
comparatively large and uniform in shape, being more or less
polygonal in section. The walls of the trama are bright yellow
in most species. In this genus the peridium appears to be
completely formed at a considerable distance underground, as
some species have a stout stem-like base, from eight to ten
inches long and completely buried in the ground, the peridium
alone appearing at the surface. From what is known in other
^ Massee, "Monograpliof the Genus Podaxis," in Jb?/?'. of Botany, March 1890.
PUFF-BALL FUNGI— GASTROMYCETES 163
instances, the stem probably remains rudimentary until the
spores are matured, when it elongates for the purpose of raising
the peridium to the surface, thereby facilitating the dispersion
of the spores." ^
The subterranean Gastromycetes, which technically belong
here, are treated in the chapter on subterranean Fungi, because
of their similarity in habit and appearance ; but for all this
they must not be confounded, and cannot be if the fructification
is properly remembered.
BIBLIOGRAPHY
FrsciiEi;, E., otc. " Gasteromyceteae," in Saccardo, Sylloge Fungorum, vol. vii.
Part I. Padua, 1888.
Kalcubrexner, C. Gastromycetes novi vcl minus cogniti. 8vo. Budapest, 1883.
Phalloidci novi vcl minus cogniti. 8vo. Budapest, 1880.
Massee, Geo. "Monograph of the British Gastromycetes." Annals of Botany,
November 1889.
"Monograph of the Genus Calostoma." Annals of Botany, June 1888.
"Monograph of the Genus Podaxis," in Journal of Botany. London,
February 1890.
"Monograph of the Genus Lycoperdon," in Journal Royal Micro. Soc.
London, 1887.
TuLASXE, L. andC. "Essaid'une Monographic des Nidulariees." Ann. des Sci.
Nat. 8vo. Paris, 1844.
ViTTADiNi, C. Monogra2)Ma Lycoperdineorum. 4to. Plates. Turin, 1842.
TuLASNE, L. " De la Fructification des Sclerodermee comparee a cells des Lyco-
perdon et des Bovista." Ann. des Sci. Nat., 2nd Series, vol. xvii.
" Sur les Genre Polysaccum et Geaster." Ann. des Sci. Nat. 2nd Series,
vol. xviii., 1842.
CoRDA, J. C. Icones Fungorum, vols, ii., v., and vi. Folio. Plates.
SoROKiN, W. " Dt^veloppement du Scleroderma verrucosum." Ann. des Sci.
Nat., 6th Series, vol. iii.
Bambeke, G. Morpliologie du Phalhis impudicus. Gand, 1889.
Hesse, R. Microscop. Unterscheid. der Lycopordaceen-gcnera. Berlin.
Massee, Geo. "A Revision of the Genus Bovista." Journ. Bot. 8vo. London,
1888.
Morgan, A. P. "North American Geasters," in Ainer, Naturalist. Roy. 8vo. 1884.
" The Genus Geaster," in ^?)ier. A'aiifT-a^w^. Roy. 8vo. 1887.
Fries, E. Symbolae Gasteromycorum ad Floram Succiam. Lund., 1817-18.
"On the Genus Queletia," mKon. Vet. Acad. Forhandl. Stockholm, 1871.
De ToNi, G. B. " Revisio Mouogi-aphica generis Geasteris. " Eevue Mycologique,
1887.
Van Bambeke, C. "Recherches sur la Jlorphologie du Phallus." Bull. Acad.
Roy. de Bclg., xxviii. Brussels, 1889.
' Massee, "Monograph of British Gastromycetes," in Annals of Botany, vol.
iv. (1889), p. 11.
CHAPTER XIY
ASCIGEROUS FUNGI ASCOMYCETES
Whatever the form which the receptacle may assume, the
Ascomycetes have always this one feature in common — that the
spores are not naked or exposed, but are always enclosed within
a delicate external membrane or spore-sac, and these latter are
imbedded in the modified hymenium. It is quite true that the
hymenium itself may be exposed, but the spore-sacs, or asci,
are imbedded, and the spores are not visible externally until
they are mature and discharged. It was proposed some years
since that the term sjjore should be applied only to such repro-
ductive bodies as were produced naked, or not enclosed in an
investing membrane, whereas all such reproductive bodies as
were developed within an ascus, or investing sac, should be
termed sporidia. It will be obvious to all who consult the
most recent works, that this distinction has not been maintained,
at least with the old limitation ; so that conidia, spore, and
sporidium are employed without recognised definition, almost,
if not entirely, as if they were synonymous. We still hold
that the spore which is produced naked, whether as a basidio-
spore or stylospore, should possess a name by which it may at
once be distinguished from such as are developed within an
ascus, whether it be ascospore, sporidium, or some equivalent.
When Saccardo elaborated his extensive work Syllogc
Fungorum he recognised this difficulty, and at the commence-
ment of the third volume defined the terms which he should
employ, and accepted siwridia as exclusively applicable to
ascospores. Spiore, simply and without prefix, was practically
the same as basidiospore, for he applied it to all the Hymeno-
mycetes. These were the two distinctive terms for the primary
ASCIGEROUS FUNGI— ASCOMYCETES 165
groups of complete Fungi to recognise the spores. The
Uredinei had special terms for the different stages of the cycle,
as commonly in use. The imperfect Fungi with naked exposed
fructification, as Hyphomycetes, appropriated the term conidia.
The other imperfect Fungi, in which the fructification was more
or less enclosed in a perithecium, cup, or cell, such as the
Sphaeropsideae, have spore-bodies with the name of sijorules.
By adhering to these terms much trouble and confusion will
be spared to the student when he comes to consult systematic
works for himself.
The lowest and simplest form of Ascomycetes is to be
found in the genus Ascomyces or Exoascus, in which the asci
are not compacted into a hymenium, but are loosely arranged
upon a delicate mycelium, without any definite receptacle or
excipulum being present. In the more typical forms the
mycelium gives rise to a receptacle of some kind, either closed
or open, in which a compact hymenium is developed, and the
whole Fungus assumes a definite and determinate form. From
certain features in this receptacle the entire Ascomycetes may
be classed in three or four distinct groups, and are thus char-
acterised : — Pyrenomyceteae, with a distinct perithecium, which
is at first closed, but at length opening by a pore at the apex,
or dehiscing by fracture, so as to allow the mature sporidia to
escape. Discomyceteae, often fleshy or waxy, with a discoid or
cup-shaped excipulum, soon expanded, and exposing a plane or
concave hymenium, from which the sporidia are ejected when
mature. Hysteriaceae, intermediate between Pyrenomyceteae
and Discomyeeteae, substance more or less coriaceous, at first
closed, afterwards dehiscing by an elongated mouth, gaping
when moist, and then exhibiting a compact hymenium. Allied
to Pyrenomyceteae by the coriaceous excipulum and the connivent
lips of the orifice, through such a family as Zojyhiostomciceae ;
but with a tendency towards Discomyceteae in the compact
hymenium, which becomes exposed when moist, and is thus
suggestive of Phacidiaceae. Finally, Tuheraceae, in which the
Fungus is normally subterranean and fleshy, the internal sub-
stance containing irregular cavities or cells, the walls of which
are lined by the hymenium ; never dehiscent, so that the
sporidia are only liberated by the decay of the entire Fungus.
1 66
INTRODUCTION TO THE STUDY OF FUNGI
This group is allied to the Discomyceteac by such genera as
Sphaerosoma and Bcrgrcnnia, and analogous to the Gastromycetes,
especially Scleroderma, through the family of the Hypogeac.
The hymenium consists usually of two kinds of organs,
which stand side by side, closely packed together ; these are
the asci and the paraphyses, but the latter are
sometimes, although rarely, suppressed. The
asci are essentially membranaceous, delicate,
colourless sacs, mostly closed, but occasionally
dehiscing at the apex by an operculum or lid,
more commonly irregularly ruptured, to permit
of the escape of the sporidia. These asci have
either a clavate form, with a more or less
elongated base, or they are cylindrical, of nearly
equal breadth throughout, except at the base,
where they are narrowed downwards to the
dimensions of the supporting hypha. In some
families, such as the Fej^isporiaccae, as well
as in the Tuber aceae, a form of ascus prevails
which approaches to globose or pear-shaped.
All forms of asci are usually very numerous
in each hymenium, but the globose are less so
than the clavate or cylindrical. The form that
is peculiar to any species is persistent in that
species, so that the form and approximate
size are relied upon as having value in the
determination of species. Whatever the form
each ascus assumes, it normally encloses eight
sporidia, or some multiple of eight — as sixteen,
thirty-two, etc. — occasionally only four, more rarely only
two, and very rarely indeed only one. In cylindrical asci
the sporidia may be expected to range themselves in a
single row, but in clavate asci they are either biseriate or
irregularly grouped towards the upper portion of the ascus.
By far the larger number of sporidia are continuous, consisting
of a single cell, and range from a globose to an elHptical form,
especially in the Discomyccteae, whilst in the Pyrenomyceteae
greater variation prevails. Doubtless all the forms of sporidia
are at first continuous, and acquire septa as they approach
Fig. 68.- — Asci aud
paraph yses.
ASCIGEROUS FUNGI— ASCOMYCETES 167
maturity. Thus a sporidium may at first be one-celled, then
it acquires a central septum and is two-celled ; each of these
cells may be again divided, so that the sporidivmi becomes tri-
septate, and by a further process of subdivision the triseptate
sporidium may ultimately become seven-septate. By means of
a less symmetrical subdivision we have also biseptate, four or
five septate, and even muriform sporidia, with the cells divided
in both directions. Whatever the number of cells into which
a sporidium may be divided, each cell appears to be a repro-
ductive unit, capable of germination and producing its kind ;
so that each septate sporidium is in itself compound — that is to
say, each of its component cells acts in the same manner as a
simple, undivided sporidium would act. The forms of sporidia
are so numerous that it would be tedious to enumerate them
here. Some of these are externally rough, but the majority
are smooth, and they may be hyaline or coloured. In the
Discomycdeae coloured sporidia are comparatively rare ; the
coloration is confined to the epispore in all cases, and the
contents remain colourless.
Paraphyses are more slender than asci, with which they
are associated, being placed side by side with them in the
hymenium, and, when present, are more numerous than the
asci, and usually a little longer, but filiform or thread-like.
There has from time to time been much speculation as to
their functions and relationship. Some have regarded them as
abortive asci, amongst whom was the late Eev. M. J. Berkeley,
who wrote : " The essential character of this important division
consists in the development of definite or indefinite sporidia
within certain of the external cells of the hymenium, called
asci, which are frequently accompanied by inarticulate or
septate, simple or branched threads, which are abortive asci,
known under the name of paraphyses." This opinion seems
to have been based chiefly upon the occurrence of organs,
apparently paraphyses, mixed with normal paraphyses and asci
in the hymenium of certain Discomycetes. These bodies pre-
sented inflations at the apex or below it, such inflations enclos-
ing a sporidium resembling the genuine sporidia in neighbouring
asci ; and hence it was concluded that these abnormal bodies
were degraded asci, not yet fully degraded into paraphyses.
1 68 INTRODUCTION TO THE STUDY OF FUNGI
It must be borne in mind that paraphyses are developed first,
and afterwards the asci ; that the paraphyses are often septate,
whilst asci are not; and that they seem to possess functions
of their own. Another theory is that the paraphyses are
sometimes, if not always, styles or peduncles surmounted by
conidia — that is to say, that they are conidiophores. The
hymenium of Tympanis ligustri has been seen bearing normal
asci and paraphyses, but amongst the paraphyses other and
shorter ones, surmounted by brown uniseptate conidia, one to
each filament. The inference that therefore paraphyses are
conidiophores cannot be maintained on such a basis without
stronger corroborative evidence.
Paraphyses are more highly developed in the Biscomyccteae
than in any other of the Ascomyceteae, and the following forms
have been indicated — i.e. Linear paraphyses, which are the
simplest form. They consist of a slender cylindrical cell of
equal thickness throughout. Sometimes they but little exceed
the asci in length, and then remain perfectly straight, but
when considerably exceeding the asci the tips are often more or
less curved as soon as they are set free, so that the retractile
tendency can exhibit itself. Such paraphyses are usually
colourless and without evident contents, but at times a row
of nuclei exhibit themselves, or the threads become septate.
These may be free of each other, or they may be agglutinated
together by hymenial gelatine.
Clavatc paraphyses are those which expand in their upper
portion into a more or less club-shape. Sometimes the expan-
sion is very gradual, occupying the upper half of the para-
physes ; at others it is more abrupt, and at least three-fourths
of the paraphysis remains linear. Gradually this form merges
into the capitate form, and usually the thickened apex is filled
with a granular plasma.
Capitate paraphyses are those in which the apex is
suddenly expanded into a pyriform, obovate, or subglobose
head. This knob contains at times a single large globose
guttule, perhaps an oil-drop ; at others it contains a granular
protoplasm. The colouring of the upper portion of the para-
pliyses may be due to the coloured contents, but in some
instances it is caused by the coloured investing gelatine.
ASCIGEROUS FUNGI— ASCOMYCETES 169
Acuminate IJciraphyses are confined to such minute hairy
Pezizae as formed a portion of the old series Dasyscypha before
the large genus Peziza was broken up into small genera. They
are slender, thickest in the middle, and diminished towards
either extremity, so as to be narrowly fusiform, with the apex
acutely pointed. As they are considerably longer than the
asci, they project on the hymenium and impart to it a velvety
appearance.
Branched p)amphyscs may be met with amongst linear,
clavate, and capitate paraphyses, but not the acuminate, which
last are always simple. Usually the branching is a simple
furcation, with the branch reaching to the same height as
the main stem. Nodulose or inflated paraphyses are rare,
such as are found in Peziza sterigmatizans and Otidea
apophysata ; but in these cases they do not seem to be
accidental, but normal, and therefore incidentally valuable in
the determination of species. In other species abnormal
developments of paraphyses have been seen and figured, but
they are not permanent to the species, and seldom to be met
with, so that they cannot be considered as other than abnormal
developments.
Bissilient paraphyses are those in which the upper joint or
joints when mature break off, and give a pulverulent appear-
ance to the disc. They are not uncommon amongst the
Patellariaceae. There seems to be no valid evidence that the
cast-off cells partake at all of the character of gonidia, or are
capable of germination.
The functions of paraphyses appear to be mainly the pro-
tection of the fructiferous organs. Surrounding the asci, they
seem to stand in a similar relationship to them as in flowering
plants the corolla bears to the essential organs. They con-
stitute in the earlier stage of growth the entire hymenium,
and in this stage form a disc with their upper extremities, as
witnessed in the Discomycetes ; whilst their parallel sides,
immersed in a gelatinous fluid, afford ready channels for the
growth and development upwards of the sporidiiferous asci.
It can readily be imagined that such a structure affords very
great protection for the asci during growth. It can hardly be
supposed that delicate asci could successively be produced on
I70 INTRODUCTION TO THE STUDY OF FUNGI
an entii'ely exposed surface without great risk of destruction ;
but by means of this arrangement they thrust themselves up-
wards through protecting channels, lined everywhere with a
lubricative fluid, so that their movements are facilitated as well
as protected. It is an undoubted fact that all the asci of an
hymenium are not developed at once, but proceed for some
time in a regular succession from the subhymenial tissue. At
first the asci are slender, gradually increasing in volume as
they rise, but until they have attained their full height their
contents are plastic and granular. Having approached their
adult stature, the differentiation of the protoplasm takes place ;
gradually the outline of the sporidia is indicated, commencing
at the summit of the ascus and progressing downwards ; and
finally the sporidia are formed. It is well to bear in mind
that the terminal sporidia are the first to be matured, and this
is conspicuously evident when the sporidia are ultimately
coloured; under favourable circumstances a delicate gradation
of colour will be observable downwards through the whole
series. It has already been remarked that it is of rare
occurrence that the asci should reach by their apices the sur-
face of the disc. As a rule the paraphyses, being the longest,
extend above, and still protect the asci. The swollen or
clavate tips compensate to some extent for the space occupied
below by the asci, and the surface is still maintained imper-
vious. In cases where the tips of the paraphyses are not
clavate but filiform, they are not unusually branched in
their upper portion, which only adds to their volume ; and in
some cases the extremities are bent, curved, circinate, or inter-
woven, so that still the whole disc is covered, and no openings
left above the apices of the rising asci. Undoubtedly the
apices of the asci are always most free from pressure or
restraint, which is essential to the free discharge of the mature
sporidia. It may sometimes be seen on the field of the micro-
scope that, as a mature sporidium is expelled from the apex of
its ascus, the clavate paraphyses which surround it are parted
by the force of the eviction, but immediately resume their
old position again with a jerk, as if impelled by their own
elasticity. These observations have been made, of course, on
such Ascomyceteae as have the disc exposed, but by analogy we
ASCIGEROUS FUNGI— ASCOMYCETES 171
]uay infer that the process is similar, if shghtly modified,
in all.
It has already been intimated that the highest development
of paraphyses is found in Discomyceteae, or such of the Ascomy-
ceteae as have the disc exposed, and we would suggest that
their function in such cases, in part at least, seems to be the
protection of the disc, or rather the apices of the asci, and to
prevent too great evaporation consequent upon the exposure
of the hymenium. In closed perithecia, such as are found in
Pyrenomycetcs, the paraphyses are often insignificant ; and in
the Tubcraccae, which are wholly immersed and preserved from
the light, paraphyses are as nearly as possible obsolete. All
which tends to support the theory of the functions of paraphyses
above suggested.
The relationship of the Ascomyceteae with the other orders of
Fungi has been the subject of some speculation, and has origin-
ated more than one theory, which we need not stay to discuss.
The Ascomyceteae and the Basicliomyceteae may be two parallel
groups, and we will leave them at that, but the Ascomyceteae
have in some of their species been shown to be associated with
such imperfect Fungi as the Hyphomyceteac, the Sphaeropsideae,
and the Melanconieae ; but because some of the species are
known to be so related, it is taking too hazardous a leap to
affirm that all the latter are merely transitional forms of the
former, and should not be regarded as autonomous. Massee
has truly said, in reference to this subject : — " The divisions
called Melanconieae, Sphaerop)sideae, and Hypliomyeeteac include
over eight thousand species from all parts of the world. Out
of this number less than one hundred have been clearly proved
by cultures to be forms of species belonging mostly to the
Ascomyceteae; yet on the strength of this small percentage of
proved cases, the three groups are entirely omitted in the
schemes of classification given by De Bary and Brefeld, imply-
ing that all are considered merely as form-species — a supposition
which may be quite correct, but is far from being proved, and
not altogether countenanced by the investigations of these same
authors, who claim to have shown that in some of the Asco-
myceteae the gonidial stage is completely lost. De Bary and
his followers do not, as a rule, accept the ' special creation '
172 INTRODUCTION TO THE STUDY OF FUNGI
theory, but, judging from their writings, consider that species
are evolved by certain processes of differentiation from previously
existing species. If so, assuming that the gonidial stage of an
originally pleomorphic Fungus alone remains, the ascigerous
condition having been entirely arrested, should the gonidial
form still be considered a phase of a higher form that has no
existence, or, being capable of carrying on an entirely independ-
ent existence, will it ever be entitled to rank as a species ?
If not, then, from the evolution standpoint, all living organ-
isms, from analogy, are merely forms of a primitive progenitor.
From the above it will be seen that in a systematic work
the Sphaeropsideae, Melanconieae, and Hyphomyceteae must be
admitted ; and until their affinities are demonstrated by direct
experiment, not analogy, it will be well to use the terms genera
and species in the ordinary sense." ^
^ British Fungi — Phycomycctcs, etc., by G. Massee, p. 65. London, 1891.
CHAPTER XV
DISCOID FUNGI DISCOMYCETES
This is one of the most interesting groups of the Ascomycetous
Fungi, in which the sporidia are contained in membranaceous
sacs, or asci, and when mature expelled from the apex, often
in a little smoky cloud, under the influence of sunlight. The
normal appearance is that of a cup or saucer, at first deeply
concave, but at length more or less expanded and flattened,
ranging in size from that of a pin's head to several inches.
The hymenium, or spore-bearing surface, is uppermost and soon
exposed, very often of a bright and attractive colour. We may
assume that this bright coloration is of some service to the
plant, but at present that use has not been determined. One
important feature, in which the majority of the discoid Fungi
differ from the majority of the Pyrenomycetal Fungi, is in their
fleshy or waxy substance, which is modified in one direction
until it becomes soft and tremelloid, and in the other direction
it is rather tough and leathery, but never really brittle and
carbonaceous.
It will be better, in the first instance, to attempt a descrip-
tion of a typical discoid Fungus such as was formerly known
by the name of Feziza, although the old genus Pcziza is now
broken up into a number of smaller genera. The general form,
when young, is either globose, or when possessed of a stem,
clavate, or club-shaped, pierced with a pore at the apex. As
growth proceeds, the pore enlarges and the head gradually
becomes cup-shaped, so that the Fungus resembles a wine-glass ;
the disc or lining of the cup flattening with age until it is
almost a plane surface. The outer surface, or cxcipulum, as
174
INTRODUCTION TO THE STUDY OF FUNGI
sometimes called, may be hairy, woolly, granular, or quite
smooth (Fig. 69).
The inner stratum, or disc, has quite a different structure
from the outer stratum, or excipulum, being composed of a
series of elongated, delicate cells like
cylinders, closely packed side by side,
their apices terminating in the disc,
and their bases being seated upon the
inner surface of the excipulum (Fig. 70).
In due time these cylindrical sacs, or asci,
contain four or eight, or some multiple
of that number, of smaller bodies, which
are the spores or sporidia — the repro-
ductive corpuscles of the Fungus. In
some cases the apex of the ascus opens
Fig. 69.— Cup of Peziza, with -^^ means of an operculum, or small
section and ascus. v i t, . . - u i, • i
lid, but at others by an irregular rup-
ture, to permit of the escape of the spores. Mixed with
these spore-bearing sacs will be found a number of long
thread-like bodies of equal length, or longer than the asci, and
these are termed, jparcqjliyses. Some mycologists believe them
to represent abortive
asci, and this is sup-
ported by the fact that
now and then a para-
physe is observed which
encloses one or two
like the normal
of the Fungus.
the Peziza is
and the cup is
spores,
spores
When
younf
Fig. 70. — Section of cup of Ascuhulus.
closed, all the cylindri-
cal cells are narrow, thread-like, and empty ; but as growth pro-
ceeds and sporidia begin to form, the diameter of the cells
increases, and, as a consequence, the disc enlarges and expands
with the lateral pressure so as to occasion the flattening out of
the cup. When there is a great expansion the edges of the cup
are either split or bent back, so that the disc becomes convex,
all these modifications being due to the thickening of the asci.
DISCOID FUNGI— DISCOMYCETES
175
With such a structure it will manifestly be almost im-
possible to trace the development of the spore and to set at
rest the question of sexuality in reproduction. It has been
assumed that there is some form of impregnation in the Dis-
comycetes, either for each individual ascus or for the entire
cup. Those who have advocated the impregnation of the asci,
affect to see in the paraphyses some representative of the male
organs, but in support of this theory there is no evidence. The
granular contents at the apices of the paraphyses do not
suggest spermatia, but mostly colouring matter which imparts
the tone of colour to the disc. Advocates have also been found
for the fertilisation of the entire cup in its most initial stage
of growth. These profess to have found, especially in Ascoholus
fiirfuraccus and in Pyronema conjiuens, all that they require to
establish sexuality. Woronin^ in Lachnca 'pidcherrima (Cr.)
claims to have ascertained that the cup derives its origin from
a short and flexible tube, thicker than the other branches of
the mycelium, and which is soon divided by transverse septa,
or partitions, into a series of cells, the successive increase of
which finally gives to the whole a torulose and unequal appear-
ance. The body thus formed he called a "vermiform body,"
since designated a "scolecite" ^ (Fig. 71). He also seems to have
convinced himself that there always exists in proximity to this
body certain filaments, the short arched or inflexed branches
of which, like so many antheridia, rest their anterior extremities
on the uniform cells. This contact seems to communicate to
the vermiform body a special vital energy, which is immedi-
ately directed towards the production of a somewhat filamentous
tissue, on which the hymenium, or disc, is at a later period
developed.
Tulasne ^ observes that this scolecite can be readily
isolated in Ascoholus furfuraceus. When the young receptacles
are still spherical and white, and have not attained more than
one-twentieth of a millimetre in diameter, it is sufficient to
compress them slightly in order to rupture them at the summit
and expel the scolecite. This occupies the centre of the
1 De Bary, Beitr. zur Morph. der Pihe, 1866.
2 See Fig. 26.
3 Tulasne, Ann. dcs Sci. Nat., Oct. 1866, p. 211.
176
INTRODUCTION TO THE STUDY OF FUNGI
little sphere, and is formed of from six to eight cells curved in
the form of a comma.
In Pyronema mdaloma Tulasne states that the scolecite
in this species is most certainly a lateral branch of the mycelium.
This branch, is isolated,
simple, or forked at a
short distance from its
base, and in diameter
generally exceeding that
of the filament which
bears it. This branch
is soon arched or bent,
and often elongated in
describing a spiral, the
irregular turns of which
are lax or compressed.
At the same time its
interior, at first con-
tinuous, becomes divided
by transverse septa into
eight or ten or more cells. Sometimes this special branch
terminates in a crozier shape, which is involved in the bent
part of another crozier which terminates in a neighbouring
filament. In other cases the growing branch is connected by
its extremity with that of a hooked branch. Of these con-
tacts Tulasne was uncertain whether they were normal or
accidental. But of the importance of the scolecite he conceived
there was no room for doubt, as being the certain and habitual
rudiment of the fertile cup. Inferior cells are produced from
the flexuous filaments which creep about its surface, cover and
surround it on all sides while joining themselves to each other.
At first continuous, then septate, these cells by their union
constitute a cellular tissue, which increases little by little until
the scolecite is so closely enveloped that only its superior
extremity can be seen. These cellular masses attain a con-
siderable volume before the hymenium begins to show itself in
a depression of their summit. So long as their smallness
permits of their being seen in the field of the microscope,
it can be determined that they adhere to a single filament
Fig. 71. — Scolecite. After Kililmaii.
DISCOID FUNGI— DISCOMYCETES 177
of the mycelium l)y the base of the scolecite, wliich remains
naked.^
The same investigator claims to have been more successful
in his search after some act of copulation in his experiments
with Pyronema confluens. As early as 1860 he recognised the
large, globose, sessile, and grouped vesicles which originate the
fertile tissue, but did not comprehend the part which they
were to perform. Each of these emits from its summit a
cylindrical tube, generally flexuous, but always more or less
bent in a crozier shape, sometimes attenuated at the extremity.
Thus provided, these utricles resemble so many tun-shaped,
narrow-necked retorts, filled with a granular, thick, roseate
protoplasm. In the middle of these, and from the same fila-
ments, are generated elongated clavate cells, with paler contents,
and more vacuoles, termed by him paracysts. These, though
produced after the other bodies, or macrocysts, finally exceed
them in height, and seem to carry their summit so as to meet
the crozier-like prolongations. It would be difficult to deter-
mine to which of these two orders of cells belongs the initiative
of conjugation. Sometimes the advance seems to be on one
side and sometimes on the other. However this may be, the
meeting of the extremity of the connecting tube with the
summit of the neighbouring paracyst is a constant fact,
observed over and over again a hundred times. There is no
real junction between the dissimilar cells, except at the very
limited point where they meet, and there a circular perforation
may be discerned at the end, defined by a round swelling,
which is either barely visible or sometimes very decided.'
Everywhere else the two organs may be contiguous, or more or
less near together, but they are free from any adherence what-
ever. If the plastic matters contained in the conjugated cells
influence one another reciprocally, no notable modification in
their appearance results at first. The large appendiculate cell
seems, however, to yield to its consort a portion of the plasma
it contains. One thing only can be affirmed from these
phenomena — that the conjugated cells, especially the larger,
wither and empty themselves, while the upright compressed
^ Cooke, Fungi, their Nature, Uses, etc., p. 174.
12
178
INTRODUCTION TO THE STUDY OF FUNGI
filaments, which will ultimately constitute the asci, increase
and multiply.^
Starting with this idea of the general structure of the
discoid Fungi, it will scarcely be difficult at any time to dis-
tinguish the various genera and species from those of the
residue of ascomycetous Fungi. The latest revision of the
classitication is that by Professor Saccardo, and he has enu-
merated altogether 3450 species, distributed over twelve
families and included under 190 genera.
The most striking divergences from the cup-shaped type
are those of the pileate forms, in which, as in MorchcUa (Fig. 72),
the irregular cups are gregarious upon a
common stroma ; or in others of the
pileate genera in which the hymenium
is spread over the upper surface of
erect clubs, as in Geoglossum, or ex-
panded laminae, as in Hclvella. In
all these cases the fructifying surface
is superior and exposed, and the sporiclia
are contained in membranaceous sacs
or asci, which latter are not enclosed
in closed perithecia. Many of them
Fig. l2.—Morcheila escuienta. are large euough and succulent enough
to be employed as articles of food, and
we do not remember that any one species has been proved to
be poisonous, although doubts have been expressed of the
wholesome character of one or two, and notably of Gyromitra
escuienta.
The largest number inhabit the ground or flourish on
rotten wood, and by far the most part are confined to temperate
climates, species found in tropical or subtropical regions
being chiefly those of a tough and leathery consistence. The
Morels, for instance, wdien they occur in India, are found at
considerable elevations on the Himalayas, where the climate
corresponds to that of temperate regions (Fig. 73). The analogues
of the fleshy Pezizae are found under the aspect of Ccnangium,
JJrnula, or Tympanis in warm climates ; whilst in Mexico and
^ Tulasiie, " On the Plienomena of Copulation in certain Fungi," Aim. dcs Sci.
Nat., 1866, p. 211.
DISCOID FUNGI— DISCOMYCETES
179
Fig. 73. — Himalayan moiel,
with section and s2)oii(Iiuni.
India a large leathery Fungus of the genus Midotis takes the
place of the species of Otidea found in Europe.
The Fungi in this group which exercise a deleterious
influence on growing plants are limited in number, and con-
fined to two or three genera. It has been declared that a
small Feziza, under the name of Pcziza
Willkommi, is the cause of the devas-
tating larch disease ; whereas we are
of opinion that it is really the same
as Dasyscyioha dandestina, and is para-
sitic upon the diseased spots, caused
by resinosis, and is not the source of
the disease. More decided, however,
are the relations between the small
species of Pseudopeziza and the living
plants which they attack. One of these
is common on clover, another on
lucerne, and others on Caltha, Gcdmm,
etc. All these are undoubtedly de-
structive, but outside this genus nearly all the leaf-species only
occur upon leaves subsequent to death or decay. A very
abnormal series of forms, most nearly related to the Discomy-
cetes, and classed with them, are the occasion of peach blister,
the pear -leaf blister, and similar diseases. These Fungi of
the genus U.wascus have no proper excipulum, but consist of
naked asci, placed side by side on a kind of mycelium invest-
ing the blistered spots.
It would be scarcely out of place. to allude here to a few
species of the form of Feziza, but classed together under the
generic name of Sderotinia, which are developed from fungoid
bodies called Sderotia — which are a sort of compact mycelium
— and after a period of rest give rise to species of Feziza or
Sderotinia. One of these is common on a Sderotium found in
company with the roots of the wood anemone (Fig. 74). Another
is developed from a little black Sderotium often common in the
haulms of potatoes and cabbage-stalks. Another, again, occurs
on a Sderotium developed within the substance of rushes.
The injury to the plants is caused in the Sderotium stage, but
the mature Fundus bears the common name of Sderotinia.
I So
IXTRODUCTIOX TO THE STUDY OF FUNGI
Some mycologists have advanced the opinion that at least
many of the species of the genus of moulds named Botrytis
are the conidia of some species of Peziza.
It has been shown by Tulasne, and others, that some of
the discoid Fungi appear under two or more forms or phases,
which resemble each
other in outward ap-
pearance, but differ
in fructification. In
tlie case of Calloria
fusarioicles, on nettle
stems, there is a con-
idial form in which
no asci are developed,
l)ut naked spores are
produced on sporo-
phores. Later on,
and upon the same
stems, in company
with the conidia true
cups are perfected
which contain asci
•-^co and sporidia. Both
are of the same size
and colour, and in
well-developed speci-
mens it is difficult
to distinguish them
without the use of
the microscope. In
the case of Coryne
sarcoides the same re-
semblance exists be-
tween the two con-
ditions. The form
in which conidia
only are produced was previously called Trcmdla sarcoides,
and is of a peculiar reddish -violet colour, bursting in
clusters through fissures in wood or bark. The perfect
Fig. 74. — ^demtinia tvhcrom on anemone.
Ganl. Chron.
DISCOID FUNGI— DISCOMYCETES i8i
condition proliably exhibits a more definite disc, but the asci
are well developed, containing eight sporidia of an elongated
form, at first nucleate but ultimately triseptate. In the
genus Tymrpanis it is not unusual to meet with cups which
have no asci and only bear stylospores or conidia. The various
species of Cyijlidla are suspicious of relations to Pcziza, but not
yet satisfactorily determined. The species in this genus are
imitations of Peziza in form, but the disc is more like the
hymenium of Corticiu'm, and hence the genus is located in the
Hymenomycetes. Many of these were called by the name of
Peziza before the fructification was investigated, and in the
future some of them may have to be restored again as the
stylosporous conditions of true Pezizae. There are still to be
found, in two groups widely apart, the Clavaria nigrita in the
Hymenomycetes and Geoglossum nigritum amongst the Disco-
mycetes, hardly distinguishable in appearance, but bearing in
the former case naked spores and in the latter sporidia enclosed
in asci.
The relations of the discoid Fungi to other groups has
sometimes been matter of speculation. Some of the larger
Pezizae have a subterranean habit in the first instance, and the
cups almost closed, excepting a perforation at the apex ; but
in Berggrenia the species are completely closed and subter-
ranean. In all other respects they are Pezizae, the inner walls
of the receptacle bearing the asci in the form of a continuous
hymenium. Another genus, at present grouped with the
Tiiberaceae} or truffle family, is called Hychiocystis, and the
structure is so similar that it becomes doubtful whether there
is any valid generic difference. At any rate this appears to be
the point where the Tuheraceae are united to the Discomycetes,
and whence they diverge.
If the several genera of Patcllaria, Patinclla, Durella, and
Zecanidion are compared with such genera as Zecidia amongst
^ Berkeley says : ' ' There is a small group of Pezizeae which grow in sand or on
loose earth, in which the cups are more or less buried. These species are scarcely
distinguishable from Hydnocystis. In the species which are more nearly allied
to Peziza, the asci are often cylindrical, and the sporidia of moderate dimensions."
And again : '^Hydnocystis is, in fact, very near to such Pezizae [P. scpulta, etc.],
though essentially distinct and far more neat in habit " {Introduction to Cryido-
gaviic Botany, p. 286).
1 82 INTRODUCTION TO THE STUDY OF FUNGI
the Lichens, it will be observed that here again the discoid
Fungi come almost in touch with another outside group, and
seem to pass almost insensibly into Lichens, destitute of a
visible thallus. Amongst the Stidcac there are such genera as
Xylographa — which some lichenologists still claim, but which
mycologists will not reject — in which the relations of Fungi
and Lichens are most intimate. In Platystida as compared
with Platygrapha, and some species of Stidis with Thdotrcma,
the resemblance, if not affinity, is maintained.
The classification adopted by Fries in this, as in all other
groups, was based primarily on external characters distinguish-
able by the aid of a common lens. Microscopical characters
had none, or but a subsidiary place. Although his success was
very great in appreciating affinities, considering the limits of
his investigations, yet much was left imperfect and undecided
when subsequent observers came to apply the microscope. It
was felt that, however great was the success when applied to
large objects, the system was not sufficient for small ones ; so
that, step by step, alterations and additions were proposed by
Fuckel, Karsten, and others, which culminated in the carpo-
logical system elaborated by Saccardo. Whether, as a whole,
this latter is too artificial to satisfy the aspirations of those
who believe in natural affinities, must be left to individual
judgment. The basis of classification being transferred fioni
external characters to internal fructification, there was a
manifest danger lest external characters should be wholly
ignored ; fortunately, however, this was not the case, so that
the result may be expressed as a combination of the two,
giving the pre-eminence to the carpological.
The fructification of the discoid Fungi, as compacted in the
disc, consists of the asci and their contents and appendages —
that is to say, the asci and their contained sporidia, and the
paraphyses. The asci for the most part are cylindrical, or else
clavately cylindrical, seldom ovate or approaching to globose.
The sporidia, subject to considerable variation in form and
size, are to a greater extent simple and uncoloured than in any
other group of Ascomycetes ; in the larger species elliptical
or globose, in the smaller cylindrical or fusoid — the colour of
the sporidia and their septation being held as of generic value.
DISCOID FUNGI— DISCOMYCETES 183
By this means the larger genera have been divided into
smaller ones, with analogous external characters, as part of the
diagnoses.
The paraphyses are more highly developed, as a rule, than
in the Pyrenomycetes. In many the clavate or swollen ex-
tremities contain coloured granular protoplasm, which contri-
bute to the colour of the disc. The asci themselves are
uncoloured, but the paraphyses being very numerous, and
exceeding the asci in length, determine the colour of the
hymenial surface. In some few instances it has been
suspected, rather than proved, that the paraphyses are capable
of bearing conidia. In some genera the paraphyses are acute
at the tips and thickened downwards, uncoloured, and extend-
ing beyond the asci more than usual, so as to impart a
minutely velvety appearance to the disc. In certain genera
the asci, when mature, are projected beyond the paraphyses
and the surface of the disc, and sometimes are expelled with
the sporidia within them.
It was thought at one time, and perhaps a few faithful
votaries still survive, that the application of iodine to the
hymenium would be a valuable aid in the discrimination of
species. Most practical men have, however, discarded it after
trial, upon the conviction that it is not a trustworthy guide.
Eeagents may be useful with the denser apothecia of Lichens,
but only with such Pezizac as are most nearly allied to Lichens
in texture.
In comparing the sporidia of the discoid Fungi with those
of the Pyrenomycetes it will be observed that, even in cases
where the sporidia are coloured, they are few in number and
not so opaque ; and that as for the form, the most typical in
the Discomycetes is the elliptical, and in the Pyrenomycetes
the fusiform. The clathrate or muriform spore, not uncommon
in SphderiaceA, is almost unknown in the Pczizci. Successful
artificial culture or germination of spores in the Discomycetes
has been accomplished in a few genera.
It seems worse than folly to attribute, as some do, every
peculiarity of structure, habit, or coloration to some special
purpose, such as protection, attractiveness, etc., without having
any basis of fact for their conclusions. There are plenty of
1 84 INTRODUCTION TO THE STUDY OF FUNGI
such problems to solve in connection with discoid Pungi, but
the facts are few and the inferences are not self-evident. In
certain isolated cases they may be, but they concern mostly
only the species in question, and are not of general application.
Take, for instance, the case of Geoinjxis ammoi^liila. Here is a
Peziza of considerable size, of the colour of sea-sand, the
exterior usually covered with particles of sand, and the base
attenuated into a tapering root three times in length that of
the cup. When it is known that this species is found only on
loose sandy dunes, it is at once concluded that the long rooting
base serves a useful purpose in attaching the Fungus well into
the loose sand and thus preventing its drifting away to
destruction, whilst the inconspicuous colour masks its presence
and prevents its being designedly uprooted. But it is not
always evident, as in this case, what is the purpose of the
peculiarity. The semi-subterranean species of SepuUaria are
mostly of some shade of brown, and the exterior is covered
with a dense matting of interwoven hairs. They affect a
globose form, with the disc but slightly exposed, except when
saturated with moisture. Blown away from their attachment,
their form assists them in being transported from place to
place, until a wet locality is reached, when the cup more or
less expands, and they drift no more until again dried and
closed, when the globose form is resumed. The colour, ap-
proximating that of the soil, may be deemed a protection, and
the dense woolly coat a means of retaining moisture, as well as
preventing rapid evaporation and consequent desiccation. But
that which may be true enough of one species, or of one series
of species, is not necessarily a theory of universal application.
It is true that nearly all the species of Ascoholus which grow
upon dung are so inconspicuous that they can only be
distinguished from the matrix by close observation. It might
be inferred that this accommodation of colour was intended
for the preservation of the individual and the perpetuation of
the species. This would not apply, however, to Humaria
gramdata, which is very common in large patches, of a bright
orange colour, on the same matrix, very conspicuous at a
considerable distance. In this instance colour can be no
protection, but what is the purpose of the bright colour ?
DISCOID FUNGI— DISCOMYCETES 185
Then there is the well-known Peziza aurantia, three inches in
diameter, growing amongst grass, several orange si^ecies of
Humaria scarcely a quarter of an inch in diameter, growing on
the naked soil, and in both cases with a smooth external
surface. Compare these with the orange or red species of
Scutellinia which flourish on wood and have the exterior
covered with brown bristles. Does the colour serve the same
purpose in all, and, if so, why are the species of Humaria
smooth and those of Scutellinia covered with bristles ? These
are problems not easily solved, because the substratum of fact
is imperfect. It will not serve to assume that the bright
colour is an attraction to insects, since it is not known that
the visits of insects would be of much service to the Fungus.
If there is no known process of fecundation there will be no
cross fertilisation to accomplish, and the dispersion of the
sporidia is assured by the elasticity of the asci and the force
with which the sporidia are seen to be expelled in a little
cloud, puffed out at intervals under the influence of sunlight.
It would be folly to dogmatise, and say that coloration of the
disc is only accidental, and of no service to the plant, because
observation has as yet given no clue to the mystery. It would
be far more reasonable to assume that there is a purpose for
everything, and endeavour to ascertain what that purpose may
be. Why are the majority of brightly coloured species of a
soft and fleshy consistency, as in Calloria and Orhilia, and why
are the black, or nearly black, species tough and coriaceous, as in
Ti/mjMnis and Urnula ?
The economic uses of the discoid Fungi are limited to but
a few species, and these comprised within three or four genera.
The Morels are widely known and appreciated, and deserve to
stand at the head of the list. There are in all twenty-four
species of Morchella, all of which may be assumed to be edible.
Eighteen of these are European, two North American, two
Asiatic, one common to Eussia and North America, and one
confined to the Canaries. Two or three of the European
species have a wide distribution, being found also in North
America, Australia, Tasmania, and Kashmir. The nine species
of Gyromitra are rarer and of less importance. Helvetia, though
including forty-five species, contain a large proportion which
1 85
INTRODUCTION TO THE STUDY OF FUNGI
are too small for practical use. The two best known and ap-
preciated are tlie European Hclvclla crispa and Hclvclla lacunosa.
Corresponding to the
true ]\Iorels are the
seven species of Tree-
morels {Cyttaria) (Fig.
75), which grow on the
trunks of various species
of birch in the Southern
Hemisphere, Chili, Tierra
del Fuego, or Fuegia,
Fuegia thev are so im-
-Cijttaria, with section.
In
Tasmania, and New Zealand,
portant as to constitute the staple food of the Fuegians
during many months of the year. In appearance they some-
what resemble the Morels, but are rather more gelatinous
and smaller. " Where Fungi form a large portion of the
food of the people, it is in general a sure indication of
an unproductive climate or
an extremely depressed
peasantry ; but it is possible
that the qualities of Cyttaria
(Fig. 76) may be superior
to those of other Fungi,
arising probably from its
immediate imbibition of the
elaborated gummy sap of
the matrix." ^ The other
esculents remaining are witn sectiou.
nearly all the larger species of Pcziza or Discina, which are
more or less eaten throughout Europe, but have very little to
recommend them. Discina vcnosa is sometimes sold under the
name of Morel.
All the collections of Fungi hitherto made in tropical
countries have exhibited a preponderance of the large woody
species of Fomcs and other Polyporci, which are not only persist-
ent through the year, but are also of a size readily to be seen
by the collector, and giving but little trouble in their preserva-
tion. On this account the minute and inconspicuous species
^ Berkeley, Introduction, p. 293.
Fig.
Cyttaria Gunnii,
DISCOID FUNGI— DISCOMYCETES 187
have been overlooked, so that the catalogues of Fungi from
the tropics are always to be suspected as very incomplete in
their enumeration of the Ascomycctcs, except in a few in-
stances where a competent resident has superintended the
investigation. Hence a comparison of the species recorded from
such places with those found in Europe is hardly just to the
tropics, and would not, at present, offer a fair estimate of
geographical distribution. Out of a total of about 1200 species
collected in Ceylon by Dr. Thwaites there were fifty discoid
Fungi and upwards of 2 0 0 Spliaeriacei. This is a much larger
proportion than usual, but in neither case can it be regarded
as exhaustive. Taking the whole estimate of described
Hymenomycetal Fungi at 9000, those of the Discoid Fungi,
for all the world, are upwards of one-third of that number,
and of Sjjhaeriacei two-thirds more ; so that the proportion for
Ceylon, with 700 Hymenomycetes, would have been some
230 Discomycetes and upwards of 450 Sphaeriacei. Hence
to bring up the proportion to the universal ratio there should
have been nearly five times as many Discomycetes, and more
than twice as many Pyrenomycetes, found in Ceylon. If this
does not indicate that the Discoid Fungi are one-fifth less in
number in the tropics than in an universal total, then the
only conclusion remaining is that the list of Ceylon Discomy-
cetes is still incomplete, and yet Ceylon has been one of the
most favoured localities in having Dr. Thwaites as a resident.
In other tropical countries there is a still greater disparity, so
that it would be premature to draw inferences of geographical
distribution based upon present knowledge.
BIBLIOGRAPHY
Saccardo, p. a. " Sylloge Discomycetmii," in SijUoge Fungorum, vol. iii.
Padua, 1889.
Phillips, W. A Manual of British Discomycetes. Sm. 8vo. Plates. London,
1887.
Cooke, ]\l. C. Mycographia.Yol. \. "Discomycetes." Col. plates. Imp. 8vo.
London, 1879.
BouDiEK, E. " Memoire sur les Ascoboles." Ann. dcs Sci. JS'at., 5tli series,
vol. X. Paris, 1869.
De Notaris, G. Fropostc di Alcune Rcttificazioni al Profile dei Discomiceti.
Imp. 8vo. Genoa, 1864.
iSS
INTRODUCTION TO THE STUDY OF FUNGI
GiLLET, C. C. Champignons dc France— Discomycctcs. Svo. Col. plates.
Alencon, 1879, etc.
Karstex, p. a. Monographia Pczizamm Fennicarum. Svo. Helsin"fors,
1871.
Synopsis Pezizarum et Ascoholorum Fenniae. Svo. Helsingfors, 1861.
Nylander, W. Observationes circa Pezizae Fenniae. 8vo. Helsingfors, 1868.
Janczewski. Morph. der Ascoholus furfuraceus. Botan. Zeitung. 1871.
EiDAM. "Beitr. z. Kenntn. d. Gymnoasceen," in Cohn, Beitr. z. BioL, iii.
Massee, G. British Fungus Flora. Vol. iv. Discomycetes. Svo. London,
1895.
CHAPTEE XVI
SUBTERRANEAN FUNGI TUBERACEAE
Two very similar small groups of Fungi resemble each other
very much in habit and in external appearance, but differ
considerably in their internal structure. These are the some-
what globose balls which are produced under the surface of
the soil, and called Tuberaceous or Subterranean Fungi. One
group has the spores developed in asci, belonging therefore
to the Ascomycetes, and the other has the spores naked on
basidia, and so belong to the Basidiomycetes, or that section
called Gastromycetes. The former are represented by the true
Truffles, and the latter by the false Truffles or Hypogaei.
To begin with the simplest forms, we encounter struc-
tures which resemble small underground species of Scleroderma.
They are mostly somewhat globose in form, of a
dirty colour, with a thick outer coat or peridium.
In one or two genera the outer coat is thin or
obsolete ; when mature, if cut through the centre,
they are seen to be filled with a powdery mass of
dark -coloured spores, but if cut when young
the interior is streaked and mottled, at length
full of small cavities, in which the spores are
produced. The spores themselves are sometimes
elliptical or almond-shaped, with either a rough Odaviana.
or smooth surface, or they are globose and warted (Fig. 77).
It may well be said that these Fungi are underground
puff-balls, which are united to the terrestrial Gastro-
mycetes through the species of Scleroderma, one of which has
often a modified subterranean habit. In former times a species
of Melanogaster was sold and used, under the name of Eed
I
I90 INTRODUCTION TO THE STUDY OF FUNGI
Truffle, as a substitute for the genuine article (Fig. 78). Many of
the species possess a strong penetrating odour, which may be
useful in guiding animals where to search for them, but can
scarcely be protective. From their habit they are very diffi-
cult to find, and hence are regarded as more uncommon than
they probably are. They seem to
prefer a sandy soil, and are to be sought
near the roots of trees. Altogether not
more than seventy -five species are
known, of which no less than sixty
are European. A very few occur in
Australia, about fifteen in America,
one or two in Africa, and about three
in Asia, so that it is pre-eminently a
^''i/efe^^Sr °^ European group. Whether we regard
them as degenerate Truliies or sub-
terranean puff-balls, they appear to be the link which unites
the Basidiomycetes to the Ascomycdes, by means of the
Tuberacei or genuine Trufties. In old age, when the asci are
dissolved, it is difficult to distinguish the species of Elaphomyccti
from the Hypogaci.
In their earlier stage it is not difficult to determine the
character of the fructification. Then the walls of the in-
ternal cavities are lined with basidia or elongated cells, crowned
at the apex with two or four little spicules or sterigmata, at
the tips of which the spores are produced. When mature the
spores fall away, and lie free in the cavities. From the
number of spores that these cavities contain, the Rev. M. J.
Berkeley was led to infer that spores were produced consecu-
tively ; but there is no substantial evidence to support this
view, and no analogy in any other group of Basidiomycetes, so
that we fear the theory is untenable.
Of the germination of the spores, the production of
mycelium, and gradual development of the young plant, we
are not aware that anything positive is known ; and the
Fungi themselves being of no economic value, their cultivation
has not been attempted.
From the Gastromycetal Hypogaei we turn to the more
highly developed Tiiberacci, and here we find in most cases
SUBTERRANEAN FUNGI— TUBERACEAE 191
that the individuals attain a far larger size and a greater
value, from the utilitarian point of view. The interior does
not become pulverulent in the genuine Truffles, although it does
in Elaj^hoviyces, and when cut through in section the flesh is
mottled and veined. Slices of the substance under the micro-
scope exhibit rounded delicate sacs or asci, which enclose the
sporidia lying freely within them. Externally the surface of
the peridium is often warted or rough, and the resemblance to
a subterranean Scleroderma is more remote. The total number
of described species scarcely exceeds one hundred and twenty,
of which fifty belong to the genus Titber, and all except eiglit
are European. Of the extra-European species two belong to
Ceylon,^ one to Malacca, one uo North America, three to South
America, and one to Tasmania. It is somewhat remarkable
that North America is so deficient in examples of this group, as
only two or three of the European species have been found there
in addition to the one indigenous species. The species 0^ My-
litta, or native bread, are not genuine members of this group, •
although often associated with them, but they would seem to
belong rather to that congeries of imperfect forms termed
SdcTotia.
The most prolific country for the production of species of
Truffles is Italy, but the most celebrated of Truffles for the
table are those of France. At one time Truffle hunting was
conducted with success in the southern coun-
ties of England, but for many years the industry
has declined, and is now almost extinguished.
This is said to be due to the importation of
French Truffles of more exquisite flavour, and
at a lower price. All Truffle spores are large, fig. 79. — Aiveo-
those of our common British species being late sporidium of
Truffle.
alveolate, or covered with hexagonal pits, the
walls of which are of transparent membrane (Fig. 79). The
spores of the French Truffles are spinulose (Fig. 80). It is easy,
by aid of the microscope, to determine even by a fragment the
French from the English Truffle, or Tuber aestivum from Tuber
melanosporum, on account of the difference in the sporidia. " The
extent of the trade in Truffles may be estimated from the fact
^ Some kind of Truffles are reported to be found in the Kangra Valley, India.
192
INTRODUCTION TO THE STUDY OF FUNGI
that ill Apt upwards of four thousand pounds are sold every
week durini;- the height of the season, whilst the department
of Vaucluse yields about thirty tons per annum."
V" .; Berkeley says that "though a few species are not
S^X^:^:'\ confined to limestone formations, it may be
assumed in general that Truffles require a cal-
careous soil for their growth, and that they increase
Fig. 80.— ^^^ number of species and individuals as we approach
Spiuulose , . . A -1 1 A 1 • p 1 • •
sporidium the southern limits of our island. And it this is
of Truffle. ^^^^^ ^^ Trufflcs in general, it is more especially so
as regards the esculent kinds, which are alone likely to be
objects of cultivation."
The problem of Truffle cultivation has been discussed
over and over again, and experiments have been made,
with more or less success. MM. Tulasne have shown that
an abundant spawn is produced in several genera. The
common Truffle exhibits in the soil in which it grows, durmg
the month of September, a profuse mycelium of white
cylindrical strings, more slender than sewing -thread, which
themselves consist of multitudes of delicate articulated
filaments, communicating with a kind of byssoid mass, some
lines in thickness, surrounding the young Truffles. This mass
soon disappears, so that only a few isolated filaments remain
attached to the surface of the Truffle.
Attempts have been made from time to time to propagate
Truffles, or to produce a saleable spawn, but hitherto with but
small results. In the south of France one nobleman succeeded
in raising Truffles in his woods by sprinkling the soil with water
in which the parings of Truffles had been rubbed down, and
protecting the ground.
Some trees appear to be more favourable to the production
of Truffles than others. Oak and hornbeam are especially
mentioned, but besides these chestnut, birch, box, and hazel
are alluded to. The old Truflie hunters obtained them chiefly
under beech, and in mixed plantations of fir and beech.
Count de Borch and M. Bornholz wrote the chief accounts of
Truffle culture. " They inform us that a compost was prepared
of pure mould and vegetable soil, mixed with dry leaves and
sawdust, in which, when properly moistened, mature Truffles
5 UB TERRA NEA N FUNGI— TUB ERA CEA E 1 93
were placed in winter, either whole or in fragments, and that
after a lapse of time small Truffles were found in the compost.
But the result was discouraging rather than otherwise. The
most successful plan consisted in sowing acorns over a
considerable extent of land of a calcareous nature ; and when
the young oaks had attained the age of ten or twelve years,
Truffles were found in the intervals between the trees. This
process was carried on in the neighbourhood of Loudun, where
Truffle beds had formerly existed, but where they had long
ceased to be productive — a fact indicating the aptitude of the
soil for the purpose. In this case no attempt was made to
produce Truffles by placing ripe specimens in the earth ; but
they sprang up of themselves from spores probably contained
in the soil. The young trees were left rather wide apart, and
were cut for the first time about the twelfth year from the
sowing, and afterwards at intervals of from seven to nine years.
Truffles were thus obtained for a period of from twenty-five to
thirty years, after which the plantations ceased to be productive,
owing, it is said, to the ground being too much shaded by the
branches of the young trees — a remedy for which might have
been found by thinning out the trees, but this would not be
adopted till all the barren tracks had been planted. The
brushwood by being thus thinned out would be converted into
timber trees ; and the Truffle grounds rendered permanent, like
those of Poitou, which are commonly situated under the shade
of lofty trees. It is the opinion of MM. Tulasne that the regular
cultivation of the Truffle in gardens can never be so successful
as this so-called indirect culture at Loudun ; but they think
that a satisfactory result might be obtained in suitable soils
by planting fragments of mature Truffles in wooded localities,
taking care that the other conditions of the spots selected
should be analogous to those of the regular Truffle grounds ;
and they recommend a judicious thinning of the trees, and
clearing the surface of brushwood, etc., which prevents at once
the beneficial effects of rain and of the direct sun rays. It is
added that this species of industry has added much to the
value of certain districts of Loudun and Civray, which were
previously comparatively worthless, and has enriched many of
its proprietors, who now make periodical sowings of acorns,
13
194 INTRODUCTION TO THE STUDY OF FUNGI
thus bringing in a certain portion of wood as Truffle grounds
each year. At Bouardeline, for instance, the annual return
from Truffles in a plantation of less than half an acre was from
£4 to £5. Another case is adduced in the arrondissement of
Apt, where several proprietors have made plantations ; the
trees are left about five or six yards apart, and so soon as their
branches meet, and shade the ground too much, they are thinned
out." 1
There are several other genera of Tuberaceous Fungi, but
of little or no commercial importance. The species of
Hydnotrya (Fig. 81) have long twisting
cavities in the interior, and the sporidia
are globose and warted. In Hydno-
holites the sporidia are globose and
-r. or rr 7 . rr i ■ alvcoktc (Fig. 82). In Choiromyces
FiG.Sl.— Hydnotrya Tidasnei. \ b / a
the sporidia are also globose, covered
with blunt spines. These Tubers sometimes attain to the size
of a man's fist. The African Truffle is Tcrfezia leonis, which
was discovered about four hundred years ago. For ages Truffles
have been eaten by the natives, but a peculiar interest attaches
to this species from the fact that the
French Academy of Sciences recently
discussed them ; and it is probably this
species of which it is said that they can
be imported from Bagdad and Biskra
so that they can be sold in the markets ^i^- ^'^•-Hydnohoiitcs with
of Paris at about one penny per pound.
This is possibly an exaggeration, but at six times the price
they would have some influence on tlie Truffle trade. One
species is reported to realise twelve shillings a pound in
Italy, but what a comparison !
In the genus Sphacrosoma the hymenium soon becomes
exposed (Fig. 83), the elongated cells or asci are closely packed
together side by side, as in the species of Pcziza ; so that the
genus approaches the character of a subterranean Discomycete,
as also does another genus named Hydnocystis. Indeed, it is
difficult to distinguish this genus from Bcrggrcnia, which is
^ C. E. Broome in Juurn. Roy. Ilort. Society, and in Gardener's Clironicle, 21st
Oct, 1865.
6- UB TERRA NEA N FUNGI- TUB ERA CEA E
195
classed under the Discomycetes, and may possibly be the same
genus with another name. At any rate the Tuhcracci are joined
to the Discomycetes by these genera, which form the connecting
link, and they might almost as well be regarded as aberrant
Pezizae as abnormal Tubers.
One of the largest genera of Tuberaceae, next to the genuine
Truffles, is Elaphomyces, with its
twenty-one species (Fig. 84). The
interior mass soon becomes dusty
and black, like soot, caused by the
disappearance of the thin walls of Fig. %-i.—Sii]Mcmsoma, with section
the cavities and the asci which at '^"^ spores.
first enclose the sporidia (Fig. 85), leaving only the free, dark-
coloured spores to constitute the internal dust. We collected
four or five species on one occasion,
in a young chestnut wood at
Montmorency. All the known
species are European, and only
one or two of these have been
recorded out of Europe. Most of
them are nearly globose, and the
outer coat is harder than in the Truffles. In former times they
had a fanciful reputation in medicine, but have long since gone
out of use.
It will be observed that in the majority of the TiLbcraceac,
where the pressure is equalised during growth, the asci, which
contain the sporidia, approach a globose form,
whilst in the genera where the hymenium is effused
over the interior, as in those which approach the
Discomycetes, the pressure is lateral and the asci
assume a cylindrical form, as typical in the fleshy
Discomycetes. In none of them are paraphyses
present. No facts are known which can lead to
the inference that any kind of sexual reproduction is probable
in this group ; and although it is believed that the germination
of the sporidia results in the production of mycelium, but little
is known of the process of germination. Spores, or sporidia,
consisting of a single cell appear to be universal, and the
form approaches more nearly to the globose than any other.
Fig. 84. — Elaphomyces and section.
Fig 85. —
Spoi idia of
Elaphomyces.
196 INTRODUCTION TO THE STUDY OF FUNGI
nir.LIOGRAPMY
Paoletti, J. "Tuberaceae, Elapliomycetaccae, Onygenaceae," in Saccardo,
Sxjlloge Fungorvm, vol. viii. Imp. 8vo. Padua, 1889.
TuLASNE, L. R. and C. Fmuji Hypogaei. Fol. Col. plates. Paris, 1862.
"Observations sur le Genre Elaiiliomyces," Ann. des Sci. A'^at. Paris,
1841.
YiTTADixi, C. Monogi-apMa Tuhcraccarum. 4to. Plates. Turin, 1831.
— — " Monographia Lyeoperdiueorum," Mem. Acad. Torino. 4to. Turin,
1841.
Hesse, R. Entwickelungs. ges. der Tiibcraceen und Elaphomyceteen. Gassel, 1889.
Enticickchivgs. dcr Hypogcen. Cassel, 1890.
Die Hyimgccn Dcutschl amis. Halle, 1891.
CHAPTER XVII
CArSULAR FUNGI PYEENOMYCETES
Adverting to what has already been written of the Ascomycetcae,
it will be remembered that the largest and most important
group of those there enumerated is that of the Pyrcnomycetcs,
although, perhaps, the Discomycdcs may be considered as the
most highly developed. It may be premised that the total of
described species for the whole world, up to date, is not less
than 10,500, or not less than one thousand more than the
whole known Hymenoniycetes. The first distinct recognition of
the Pyrcnomycetes by Fries was in 1849, and then it was
supposed to include not only the SiJhaeriacei and the Peris-
poriacei, but also the Sphacrojjsidei and Melanconiaccae. In
more recent times, when ascigerous Fungi were separated from
stylosporous Fungi, the Pyrenomycetcs were revised, and the
ascigerous species only retained as a portion of the Ascomycetes.
With this limitation they are included in Saccardo's Sylloge
and universally accepted. Eeduced to its simplest designation,
the Pyrenomycetcs are ascigerous Fungi, having the fructification
enclosed within a perithecium, and growing on vegetable or
animal substances, but are never truly terrestrial. The various
families depend for their most prominent feature upon the
character of this perithecium. The form varies within definite
limits, as well as the texture and the mode of dehiscence.
Normally the form is spherical, or nearly so, and minute,
seldom much larger than a good-sized pin's head, and either
with or without a more or less elongated neck, not unlike a
miniature flask. This may be entirely immersed, or absolutely
superficial, or intermediate. The texture may be membran-
aceous or fleshy, and then brightly coloured ; or tough and
198 INTRO DUCTIOX TO THE STUDY OF FUNGI
coriaceous, or hard and carbonaceous, and usually black.
Dehiscence may take place by means of an apical pore or
mouth, occasionally elongated but more commonly circular and
dot-like ; or the perithecium may be absolutely closed, and
when mature, splitting irregularly.
In all cases the rupture only takes
place at maturity, to permit of the
escape of the sporidia. As to
habit, in some species the individuals
Fig. 86. — Perithecium and section. 11,1-1 i i .
are absolutely isolated and simple,
and either scattered or gregarious ; whilst in others they are
collected upon a pulvinate, or more or less strongly developed
stroma, which may be of variable form and magnitude, and
in this condition are characterised as comjjound. The large
majority are saprophytic, but a few are parasitic, and the
former occur most commonly on dead or decaying wood,
branches, twigs, fruits, leaves, or the dead parts of herbaceous
plants ; the latter are mostly confined to living leaves, or the
green parts of growing plants. Other details will follow more
conveniently under the several families.
The Perisporiaceae may occupy the first place, in that they
are entirely simple, or with the perithecia separate from each
other, and not combined in a common stroma ; sometimes
membranaceous, at others coriaceous or tough, or more rarely
hard and brittle ; but their most distinguishing feature consists
in the wholly closed, or astomous, perithecia, which are
irregularly split when mature to permit of the escape of the
sporidia. It will be most convenient to review the details
under the several subfamilies, the first of which, at least, is a
compact and natural group. The Erysiplicae are of parasitical
habit, and Hourish on the living leaves and green parts of
arborescent and herbaceous plants. In the first instance a
web-like mycelium effuses itself broadly over the parts attacked,
and in this condition the leaves assume a white and mouldy
or chalky appearance, as if powdered with flour. In this stage
the delicate threads of mycelium, like a spider's web, interlace
each other, are repeatedly branched, and adhere closely to the
surface by the means of haustoria or suckers. From this
mycelium arise erect fertile branches, which become differen-
CA FS ULA R FUNGI— P YRENOM YCE TES
199
tiated into chains of hyaline conidia, which fall away and are
capable of germinating and producing a new mycelium. In
this condition the parasite is a mould,
or Hyphomycete, and was formerly in-
cluded under the genus Oidiuin, under
the supposition that it was a complete
and autonomous Fungus. Eecent in-
vestigation has shown that this stage of
mould is only the conidial condition of Fig. 87. — Perithecium and
some species of the Erysiphcae, which "S^i^ '^'°"'^^' '^
the Oidium. In the first in-
FiG. 88.
stance minute spherical yellow bodies appear on the surface of
the mycelium, and these gradually enlarge until they become
just visible to the naked eye, and acquire a dark brown colour
(Fig. 87). These are the perithecia, or
membranaceous capsules, attached at the
base by a copious mycelium and surrounded
by a circlet of free arms, or processes, as
appendages, which vary in the different
genera. In Erysiphe they are thread-like
and flexuous, of equal diameter throughout,
and simple. In Uncinida the arms are
hooked or curved at the tips (Fig. 88), In
Phylladinia the appendages are straight, and often inflated at
the base. In Sphacrothcca they are flexuous and sometimes
vaguely branched. In Podos2jhaera the
appendages are repeatedly forked at the
tips, as they are also in Microsphacra
(Fig. 89). Internally these globose
perithecia are replete with the ascigerous
fructification. The asci are nearly
globose, or pear-shaped, and contain
hyaline elliptic sporidia. In Fodosp)]iacra
and Spkaerotheca each perithecium en-
closes but a single ascus. In the other
genera the asci are numerous in each
number of sporidia in each ascus varies with the genera,
but with the exception of a single genus these sporidia are
ovoid and continuous. In the exceptional genus Saccardia
Fig. 89. — Microsphaera.
perithecium. The
200 IXTRODUCTION TO THE STUDY OF FUNGI
the sporidia are septate in both directions. In all cases
these Fungi are destructive pests, by choking the pores or
stomata, although only flourishing on the external surface.
Such are the rose mildew, Sphaerotheca pannosa ; the hop
mildew, Sphaerotheca castagnei ; the pea mildew, Erysiphe
Martii, and others. The number of known species is about 100.
The next subfamily, Perisporieae, is more numerous, and
whereas the majority of the 3ri/sipheae are confined to cool and
temperate regions, the majority of the Perisporieae are sub-
tropical. The species are not parasitical, or to a limited extent,
and the subiculum, when present, may be either colourless or
coloured, mostly the latter. The perithecia, although most
commonly subglobose, are sometimes depressed and lens-shaped,
and, except in Mcliola and one or two smaller genera, without
appendages. The sporidia are more variable than in the first
subfamily, and hence fall under five sections : Hyalosporae, with
continuous hyaline sporidia ; Phaeosp)orae, with continuous
coloured sporidia ; Didymosporae, with uniseptate sporidia ;
Phra(/mosporae,with. multiseptate sporidia; and Dictyosp)orae,vi\ih.
muriform sporidia. Under these various sections the twenty-
five genera are distributed. This arrangement foreshadows the
principles upon which the several groups of Pyrenomycetcs are
grouped ; that is to say, primarily on the basis of the sporidia,
whether coloured or hyaline, and whether continuous or septate.
It is unnecessary to occupy space by a synopsis or comparison
of the component genera, but simply to make reference to two
or three of the most important. Asterina, in its broadest
sense, is characterised by a flattened or lens-shaped perithecium,
seated on a dark radiating subiculum, with sporidia continuous
or septate, hyaline or coloured ; hence the principle adopted
almost universally by Saccardo, in other genera, of making
spore-characters of generic value is set aside, and only employed
for the distinction of subgenera. We have always contended
in favour of the Friesian system of adopting external characters
in the definition of genera, reserving carpological features for
subsidiary sections ; and hence the genus Aste7'ina, as it finds a
place in Saccardo's Sylloge, will serve as an illustration of our
method, but not of that of Saccardo, to which it does not
conform, and would only do so by elevating the separate
CAPSULAR FUNGI— PYRENOMYCETES 201
sections to the rank of genera, and distributing them over the
Hyalosporae, Phaeosporac, Bidymosporac. That is to say, we
have advocated the negation of carpological characters as of
primary generic importance in favour of their adoption in sub-
sidiary classification. Bimerosporium differs from Asterina in
possessing a globose, and not a flattened perithecium, whereas
the fructification follows the Asterina type. Another import-
ant genus of the Perisporieae is Meliola, which to a certain
extent is the analogue of some of the genera of the Erysiijhcae :
(1) by the possession of an effused mycelium, or conidia-bearing
subiculum, but in this genus more strongly developed, and of
the nature of black moulds, or Dcmaticae, and not of the Muce-
dineae ; (2) by the presence of appendages surrounding the
perithecia ; and (3) sometimes by their parasitic habit on living
leaves, — differing, however, in the fructification, inasmuch as
the sporidia are normally large, septate, and coloured. The
genus Pcrisp)orium has elongated triseptate sporidia, which
break up freely at the joints into the component cells. In
this respect there is analogy to a genus of Sphacriaccac, that of
Sporormia, which latter, except for its peri-
thecia having a distinct mouth, might be
allied with Perisporiicm. It may be stated
in general terms that the Perisporieae in-
cludes all the Perisporiaceae which do not
fall into the first subfamily, that of the
Urysipheae ; for the third subfamily, that
of the Capnodieae, includes only two aberrant
genera which have little definite alliance with
the Perisporiaceae. The genus Capnodium
is distinguished by elongated large peri-
thecia, which are often branched, and usually
opening at the apex with a large fringed
orifice (Fig. 90). These are seated upon and ^^^ ^,,, _|v,itiiecia of
amongst a dense subiculum of closely jointed Cupnndium, with
or moniliform black hyphae, so as to form '''P"^' ^^'
large velvety patches, and are possibly, in some instances, the
more complete developments of moulds belonging to the genus
Fiimago. Scorias is allied to Cajmodiuin, but thicker and more
spongy, and the perithecia somewhat clavate. Asci tetrasporous.
INTRODUCTION TO THE STUDY OF FUNGI
and sporidia triseptate. The genus AnUnnaria is more like
a dense black mould with nioniliform hyphae and apparently
minute perithecia, but the fructification is obscure, and hence
its true place and position is uncertain. The whole family of
Pcrisporiaccae contains about 700 species, and is an outside
group of Pyrcnomyceteae, joining the Hypliomycetcac on the one
hand with the Spliacriaceae on the other.
The next family to be noticed is the Hypocreaceae, in itself
characteristic and distinct, in which most of the typical features
of the Pyrenomyccteae prevail, such as the ostiolate perithecium
and ascigerous fructification, but characterised specially by the
fleshy, or nearly fleshy, perithecia, usually pale or bright
coloured, but never carbonaceous. The stroma, when present,
is soft and between fleshy and waxy, rarely forming a subiculum.
Sporidia for the most part hyaline. Without indulging in too
great prolixity of detail, it may be observed that in our
arrangement,^ which is mainly a regrouping of Saccardo's
genera by external characters, we have recognised three sub-
families. In one of these, Hypocreoideae, the species are com-
j)osite, viz. seated upon or immersed in a stroma. In the
second, Nedrieae, the species are simple, viz. the perithecia are
distinct from each other, although sometimes densely caespitose.
And in the third, Pscudonectrieae, the perithecia are soft and
membranaceous, or rostrate or
elongated, or clavate, some-
times becoming horny, either
whitish or dark coloured, and
in fact verging on Spliacriaceae.
The most highly developed
forms in the Hypocreoideae
are Claviccps (Fig. 91) and
Cordyceps, in which the species
assume a clavate or capitate
form, and the perithecia are
crowded on the upper portion
of a fleshy stroma (Fig. 92). Many of these are found on dead
insects, and their conidial forms were formerly known as species
of Isaria, a genus of Hyphomycetes. These are succeeded by
^ " Synopsis Pyrenoiiiycetuiii," by M. C. Cooke in Grerillca.
Fig. 91. — Claricejis on
CAPSULAR FUNGI— P YRENOMYCETES
203
smaller genera, and at length by Hyijocrm, in which the stroma
is pulvinate or effused, except in a few species with a vertical
stroma, and the sporidia are formed from a pair
of opposed globose cells, which separate at
maturity, and then appear to be sixteen globose
sporidia (Fig. 93). This is the typical form, but
the sporidia vary in most of the subgenera.
There can be no doubt that the eight divisions,
which are called genera by Saccardo, and sepa-
rated widely from each other in his system, on
account of differences in fructification, are naturally
closely allied to each other in structure, habit,
and development. The fundamental problem is
whether the latter are to be accepted as evidences
of close affinity, or the former ; and herein we are '■yi
at issue with Saccardo, contending that his car- fig. 92.— Co/-
pological classification of the Ascomyceteae, as t^Hl^^^ °^
developed in the Syllogc, is artificial, whilst
the method we have adopted is natural. It may be true that
an artificial and mechanical arrangement offers greater facilities
for the novice or the superficial student, but it fails to satisfy
all those who are seeking something higher than a catalogue of
Latin names.
In the second subfamily, the Nectricae, the perithecia are
all free of each other, sometimes scattered and sometimes
caespitose. The old genus Nedria, as
recognised by Fries, included both the
caespitose species, which are analogous
to Cucurhitaria in habit ; and the scat-
tered species, which are analogous to
the denudatae group of superficial
Sphaeriaceae. In our arrangement the
caespitose species are combined under
the old name of Nedria, whilst the
scattered species find their place in
another genus, under the name of
Dicdonedria. We need not repeat that
under each of these genera modifications of the sporidia fall
into place as subgenera. As a matter of fact it is well known
Fig. 93. — Byiwcrm, with
section and sporidia.
204
IXTRODUCTION TO THE STUDY OF FUNGI
that some of tlie species of Ncdria have an early stage in
which the stroma develops only conidia without perithecia, and
that these conidial forms were in earlier times regarded as
autonomous moulds of the genus Tiibcrcidaria} Later on
perithecia appear upon the old stroma, which contain asci and
sporidia (Fig. 94).
There are a few species which resemble, when mature, in
external appearance certain species of Ncdria or Dialoncdria,
hut are accompanied
by capitate conidial
forms which are not to
be distinguished from
species of the Hypho-
raycetal genus Siilhum.
Such species of the
Ncdrieae are associated
under the genus
Sjjhaerostilhc. Other
species, formerly united
with Nedria, have the
perithecia seated upon
a more or less byssoid
subiculum ; these are
now separated from that genus, and united under the name
of Byssonedria, analogous to the Byssosphacria of the
Spliaeriaceae. In another group, the perithecia, which resemble
Nedria, are densely gregarious, and often partially immersed
in a velvety subiculum, transformed from the tissues of
decaying Fungi. This genus is Hypomyccs, each species of
which has also a conidial form, which precedes the
ascigerous, and corresponds to some genus of the Mucedineae.
Some of the species of Ncdria and Dicdoncdria also
have conidial forms, which would be referable to the
Hyphomycetal genus iusarium. In these instances we must
recognise the relationship between the Hypliomycdcae and
the Ascomyceteae, but it would be assuming too much to infer,
from a few examples, that all the species of Stilbum are conidia
of Sphacrostilhe, or Tidjercularia of Nedria, Isaria of Cordyceps,
^ See Gardener's Chronicle, 28th Jan. 1S71.
Fig. 94. — D, Tuberctdaria with Xectria ; E, Nedria,
F, section of stroma ; G, asci and sporidia
Gard. Chron.
CAPSULAR FUNGI— PYRENOMYCETES 205
or various species of Mucedineae of Hypomyces. The genus
Lasionectria includes such species of Ncctria as possess a hairy
perithecium, and in this way are analogous to the Sphaeriaceous
genus Venturia or the section Villosae of the old genus Bphaeria
of Fries. Gibbcrella closely resembles Cucurlitaria in habit,
but the perithecia, although dark, are waxy, and blue or violet.
Hyponcctria again includes species of the old genus Ncctria,
but the perithecia are immersed in the matrix.
The third subfamily is Pseudonectrieae, and, as the name
indicates, links the Nectrioideae with the S])liacriaccac. The
substance of the perithecia is not of the fleshy or waxy con-
sistence of the first two families, but either membranaceous or
becoming horny, and not carbonaceous. The genus Mclcmo-
spora is somewhat analogous to Ccratostoma, the perithecia
furnished in most cases with an elongated beak-like rostrum
and brown sporidia. Another genus, Acrospermum, is placed
by Saccardo in Hysteriaccac, but Fries included it in SpJiacroj)-
sidcae through ignorance of the fructification. The species are
small, blackish, and of a club shape, with no pore at the apex,
otherwise analogous to Pocillum, amongst the Discomycetes, and
with similar long thread-like sporidia. Two or three other
small genera of little importance make up the total of this sub-
family and close the Hypocreaceae.
The remaining families of the Pyrenoviycetcae have in past
times been known as the Sphaeriaceae, but we prefer to treat
them as two large groups, each containing several families.
The Compositae, in which either a few or a great number of
perithecia are collected together upon, or immersed in, a common
stroma ; and the Simjylices, in which the perithecia are distinct
from each other, and either clustered together or scattered.
Normally the colour is black, the substance membranaceous, or
carbonaceous, and dehiscence takes place through an apical
pore or ostiolum. Fries classified them entirely according to
the external features of the perithecia or stroma, and independ-
ently of the fructification. Saccardo classified them primarily
according to the fructification, and secondarily, in great part,
from external features, or these in combination with the
sporidia.
The Compositae, or Compound Sphaeriaceae, contain the
2o6
INTRODUCTIOX TO THE STUDY OF FUXGI
following families : — Xylarieae, Dothideaceac, Melogrammcac,
Diatrypcac, Valscae, and Eiitypeae.
The Xylarieae possess a very definite stroma, which is
either vertical, or pulvinate, or effused ; the
perithecia carbonaceous and somewhat im-
mersed, and the sporidia coloured brown and
unicellular. Xyla.ria is the typical genus,
with an erect, branched, clavate, capitate, or
subglobose stroma, which is white and corky
within, and usually solid (Fig. 95). The
perithecia are peripherical, and immersed
entirely, or partially, in the upper portion
of the stroma, on all sides leaving the stem
sterile. This stem, sometimes very short,
sometimes very long, may be smooth or hairy,
Ijut it is always surmounted by a fertile head,
dotted with the ostiola of the circumambient
perithecia. In some species the entire fungus
scarcely exceeds one-eighth of an inch in
length, in others it attains to six inches or
^%yim-ia, section of i^ioi"©. with a diameter as variable ; and yet
portion and ascus throughout nearly two hundred species the
with sporidia. ^ . , ^ , , ,
essentials are the same — an erect poly-
morphous stroma, white and corky within, and a peripherical
series of immersed, or semi-immersed, perithecia, enclosing
brown continuous or unicellular sporidia. At first, and
before the perithecia are fully formed, the apex of the
stroma is usually pruinose, with pulverulent minute colourless
conidia. The species, with few exceptions, grow on rotten
wood, in damp situations, in almost all the countries of the
world, wherever a timber tree can flourish and decay. In
Thamnomyees the stroma is reduced to long black threads,
upon which the perithecia are clustered or scattered. In
Ehojjalojjsis the clubs are densely caespitose, with a short stem,
or crowded upon a very much branched common stroma. In
Poronia the stroma is almost pezizaeform (Fig. 96), with the
perithecia immersed in the disc, whilst in Camillea the stroma
is subcylindrical and truncate, with the perithecia vertically
immersed about the apex. In Daldlnia the stroma is sub-
CAPSULAR FUNGI— P YRENOMYCETES
207
globose (Fig. 97), concentrically zoned within, and the perithecia
immersed at the periphery. In Ustidina the stroma is pulvinate,
becoming hollow ; and in Nummu-
laria discoid, and plane or concave,
distinctly margined. But in the large
genus Hypoxylon, the stroma is either
subglobose (Fig. 98) or effused, solid
and dark within, closely adnate, of
variable thickness, sometimes re-
duced to little more than a crust of
densely-packed perithecia. All these
genera are allied by the possession
of a stroma and unicellular brown
sporidia, as well as their habit of
growing upon decayed wood and
dead branches, with the exception
of Poronia, most of which flourish
upon old dung.
The family Dothideaceac bear a
superficial resemblance to some
species of Hypoxylon, but differ in the perithecia being
formed from the stroma ; or, in other words, are fertile cavities
excavated in the stroma, without definite ostiola. There are
three subfamilies, viz. Dotliideoideae, Rhytismoidcae, and Stigma-
toideae. In Dotliideoideae and Bhytismoideae the species are
compound, and in Dotliideoideae the carbonaceous or coriaceous
stroma is seldom broadly effused, and the pseudo- perithecia
dehisce when mature by an apical pore. The largest genus is
Pliyllacliora, in which the stroma is either shield-like or shortly
effused and superficial, and the species are most commonly
found growing on leaves, and not rarely whilst they are still
living, but sometimes when dead. The sporidia are uncoloured
and unicellular in typical forms, but in some of the subgenera
they are coloured and continuous, or uniseptate, and in others
uniseptate and hyaline, but rarely triseptate and hyaline or
coloured. In Dothidca the stroma is erumpent and pulvinate,
the sporidia again are variable, according to the subgenera.
The species occur, in most cases, on branches, and rarely
on leaves, in which feature it differs from Pliyllacliora. In
Fig. 96. — Poronia irunctata with
section (enlarged). Gard. Chron,
2oS
IiXTRODUCTION TO THE STUDY OF FUNGI
Euryachwa the stroma is broadly etfused and puiictulate, whilst
in Homoxlcjia the stroma is plane or hemispherical, and the
species are parasitic upon
Lichens. In Illiupogyaphus
the stroma is elongated and
linear, suggesting a resem-
blance to some Hysteriaceous
perithecia. Species of Phyl-
lachora are common on cori-
aceous leaves in tropical coun-
tries, and are sometimes
difficult to distinguish at
first from some of the PJiy-
tisriioideac. In common with
the latter the stroma is often
present for some time before
the fructification is developed,
hence they are often met with
in a sterile condition.
The subfamily Rhytis-
moideae is included by some
on account of the mode of
Flu. 97. — Ihddiiiia, globose stroma and
sectiou. Oard. Chron.
authors with the Discomycdea
dehiscence, which is usually by gaping fissures, so that the
hymenium is more or less exposed ; but this dehiscence does
not take place until the sporidia are
fully mature, and sometimes not until
disintegration has commenced. In
external appearance the species arc
very similar to Phyllachora and Eury-
acliora, and yet in texture of the
stroma, and often in the fructification,
appear to be more closely allied to
the Dothidcaceae than to any family
of the Discomyccteae. Practically, the
only genus is IlhyUsma, for the sporidia
of so many described species are
unknown that no proposals have been
possible to divide them into genera based upon the fructifi-
cation. Such a species as Rhytisma accrinum, which is common
Fig. 98.— Globose stroma of
Jfl/2}oxi/lo)i, with asci and
si:)oridia.
CAPSULAR FUNGI— PYRENOMYCETES 209
on the living leaves of maple, belongs to the Sphaeropsideae
in that condition, but after resting on the ground during the
winter, asci and sporidia are developed.
The third subfamily, Stigmatoideae, includes genera in
vi^hich the perithecia are distinct from each other, and therefore
divergent from the family type, and approaching the Stiper-
ficialcs of the old genus Sphaeria. Hypospila has, however, a
feature which associates it with Dothideoideae, in the definite
stroma in which the perithecia are immersed; and also, on
this account, Trabutia shows a relationship with Phyllachora.
In the genera Stigmatea and Parodidla the perithecia are
superficial and globose, often found growing on living leaves ;
but in Stigmatea there is a very minute ostiolum, and in Paro-
didla none at all ; hence the latter suggests Perisporiaceae.
In the subfamily Melogramiiicae the perithecia are either
formed from the stroma or confluent with it, and are not car-
bonaceous, but tough and coriaceous, sometimes soft, but not
brittle, and occasionally coloured. They are densely caespitose,
but usually almost free at the apex, and destitute of any
definite neck. In most cases the tufts, or clusters, are erum-
pent, connate below, and confluent with the stroma. In habit
approaching to Diatrypeae rather than to Dothideoideae, but
the perithecia are more distinct and clustered, as in Cucur-
bitaria, and not confluent above, so as to form a disc. The
genus Sarcoxylon is rather a remarkable one, as it forms a
globose stroma, in one species as large as an orange, and solid,
with the perithecia sunk in the substance, and over the whole
surface, as in Daldinia ; but the crust is soft and never carbon-
aceous, and the perithecia are thin and membranaceous. The
sporidia are coloured, and hence it is analogous with some
species of Xylaria. Botryospliaeria is a rather numerous genus,
with erumpent, botryoid, or grape-like clusters of perithecia,
analogous to Cucurhitaria, but less distinct ; in the majority of
species the sporidia are large and hyaline, consisting of a single
cell. Endothia resembles Diatrype in habit, with a bright
yellow stroma. Fuchclia contains species which resemble an
erumpent Hypoxylon, having a subglobose stroma with im-
mersed perithecia ; but the substance is tough and rather
flexible, not at all carbonaceous or brittle. The sporidia are
14
2IO INTRODUCTION TO THE STUDY OF FUNGI
brown and without septa. Camarops in habit is just an
effused Hi/poxylon, but the stroma is soft, and the sporidia
septate and brownish. The genus Melofjramma, as limited by
Saccardo, has sporidia which are continuous and brown. The
stroma is erumpent, and then ahnost superficial, and the numer-
ous perithecia are aggregated in a similar manner to Botryo-
sphaeria, although sometimes effused. As expanded by ourselves,
this genus includes as subgenera species which correspond to
the type in habit and general appearance, but vary in the
form and septation of the sporidia. It may be intimated here
that the genus Valsaria, as characterised by Saccardo, which
has coloured and uniseptate sporidia, includes three different
types of stroma — that of Melogramma, that of Biatrype, and
that of Valsa, or rather of Pseudovalsa. In our arrangement
we refer each of these groups, as subgenera, to the species
indicated above. Thus the Melogrammoid Valsariae will be
found under the subgenus Valsariae of Melogramma.
The subfamily Biatrypeae differs at sight from Melogrammcae
in the perithecia being immersed in a stroma of a different
character, and consequently not superficially visible, and in the
substance being carbonaceous. In some cases the species are
broadly effused and crustaceous, and then resembling effused
species of Hypoxylon. In some other cases the stroma is flattened
and discoid, the imbedded perithecia being indicated by the
punctate ostiola ; whilst in a few other cases the erumpent stroma
is wart-like, and the convex surface marked with prominent
ostiola. The typical genus Biatrype has an erumpent stroma,
which is effused, or discoid, or wart-like. Those of the latter
kind, or with a wart-like stroma, but which have asci containing
a great number of sausage-shaped hyaline sporidia, are included
in the subgenus Biatrypella. The species which have similar
sporidia, but only eight of them in each ascus, whether the
stroma be verrucaeform or discoid or effused, constitute the
typical Biatrype. Other species, with the external characters
of Biatrype, but with other than sausage-shaped and hyaline
sporidia, will be found under the several subgenera of the one
genus Biatrype, which represents the sub-family Biatrypeae.
Practically there would be no difficulty for a student in the
way of discriminating species of Biatrypeae from any other sub-
CAPSULAR FUNGI— PYRE NOMYCETES 211
family except the Valscac, next to be enumerated, whilst a little
experience will soon enable him to surmount this temporary
difficulty.
The subfamily Valscac contains an immense number of
species, which are pustular and erumpent. The stroma is
formed from the altered matrix. The perithecia are quite
distinct, and mostly arranged in a circle, with convergent
necks. The principal genus is Valsa, in which the perithecia
are collected in more or less definite clusters, immersed in the
bark of trees or of their branches and twigs, and either disposed
in a simple circle or a circular group, with the necks converging
towards the centre, so as to form an erumpent disc, which splits
the bark (Fig. 99). The sporidia are hyaline, and either con-
tinuous or septate according to the subgenera. The largest
number of species have small hyaline
sausage - shaped sporidia, which is the
typical Valsa. Those in which the sporidia
are more than eight in each ascus are
either Valsclla or CoronojjJiora, as sub-
genera. When the sporidia are only
eight, the species are again subdivided ^"'- 99-— Peritiiecia of
into those in which the ostiolum is sulcate,
as Eutypclla ; and into those in which the ostiolum is not sul-
cate, but the disc is whitish, gray, or yellowish, and then called
Leucostoma ; or the ostiolum is not sulcate, and there is no pallid
disc, which is Euvalsa, or true Valsa. In two other small sub-
genera, with like sporidia, when the perithecia are four in a
group, or a small number, it is Quatcmaria ; and when a larger
number, and loosely disposed, or free, then Calosphacria ; and thus
the series of the species of Valsa with sausage-shaped sporidia
is complete. In a second section with simple hyaline sporidia,
these are of some other form than sausage-shaped, as represented
by the subgenera Oryptosporclla and Cryptospora. In a third
section the sporidia are still colourless, but septate, that is to
say, uniseptate in Chorostate and triseptate in Calospora. Closely
resembling Valsa in habit, or external appearance, is the genus
Mclanconis (Fig. 100), in which the sporidia are uniseptate,
and either hyaline or coloured, accompanied by or associated
with a conidial stage, which resembles the stylosporous genus
INTRODUCTION TO THE STUDY OF FUNGI
Melanconium. In some of the species the ascigerous perithecia
have been found growing in the midst of the pustules of
conidia, and in others closely associated in contiguous pustules.
In another genus, that of Pseudovalsa,
the sporidia are septate and coloured,
but there is no Melanconium to which
the species are related. When the
spuridia are uniseptate they I'all into
the subgenus Valsciria, but when three
or more septate into the subgenus
Aglaospora. Only one other genus of
the Valseae remains to be alluded to,
and that is Fcnestdla, in which the
sporidia are multiseptate, and divided
in both directions so as to be muriform
be intimated that of the species of
Fig. 100. — Stroma aud peri-
thecia of Melanconis.
It
may
or clathrate.
genuine Valsa, with sausage-shaped sporidia, very many of
the species are genetically connected with species of Cyto-
spora, a genus of Sphaei^o'^Jsideae, in which the spores are also
sausage-shaped and hyaline, but without asci, being produced
on short slender threads within a kind of spurious compound
receptacle which greatly resembles Valsa in appearance.
These conidia are often found growing on the same twigs as
Valsa, or upon twigs upon which the Valsa appears sub-
sequently, but the precise influence of the one upon the other
has not yet been demonstrated. Other species of the Valseae
have been named in conjunction with species of other genera
of Sphacropsideae, but less universally. In like manner species
oi Pseudovalsa are related to the similar, but stylosporous, fructi-
fication of species of Coryncum.
The last subfamily is Eidypcae, and here the leading
feature, as distiuct from Valseae, is that the stroma is broadly
and indefinitely effused, being formed from the more or less
changed matrix. The perithecia are immersed in the stroma,
and for the most part are densely and broadly gregarious.
Eidypa is the typical genus, often occurring on naked wood,
the substance of which is transformed into a stroma, in which
the perithecia are immersed. In about half the species the
ostiola are sulcate, and in the other half they are not. The
CAPSULAR FUNGI— P y RENO MYCETES 213
asci contain eight sausage-shaped hyaline sporidia. A closely
allied genus is Cryptovalsa, which conforms externally to
Eutypa, but the asci contain respectively more than eight
similar sporidia.
The other genus contained in this family has the stroma
commonly less broadly effused, and the species are more rare
on wood than on the bark of branches, twigs, and the stems of
herbaceous plants. The ostiola are often very much elongated,
and the surface of the stroma is usually blackened. The great
feature in which this genus differs from the preceding is in
the fruit : although the asci are octosporous, the sporidia are
typically fusoid and colourless, replete at first with four
guttules or minute oil-drops, and at length often uniseptate, or
in some cases triseptate.
The compound Sphaeriaceae for the most part may be
recognised at once by the naked eye, on account of their larger
size, from the agglomeration of perithecia, and the presence of
a stroma, or bed, on which they are placed or immersed, and
which of itself partakes of a definite form. In all systems of
classification prior to Saccardo, the compound and simple
Sphaeriaceae were kept apart, and were recognised as separate
groups, which certainly facilitated identification by the student.
Under the system promulgated by Saccardo, there is no
distinction of that kind, but all are mixed together and
classified according to their sporidia. Convinced that this was
practically an error, we undertook a rearrangement of the
Pyrenomycetes, in which the compound and simple were kept
distinct, whilst most of the new genera were accepted. This
scheme was embodied in Synopsis Pyreiiomycetum, published in
1884 to 1886.
The section of the Spliacriaceae which includes the Sim-
'jMces, or genera in which the perithecia are distinct from each
other, and not combined in or upon a common stroma, is even
more numerous in species, and may be described in like
manner under the several subfamilies. The connecting link
is the subfamily Cucurhitarieae, in which the perithecia are
densely gregarious or caespitose, and for the most part form
large erumpent clusters. In Nitschkia the black perithecia
are mostly clustered upon a thin whitish mycelium, the asci
2 14 INTRODUCTION TO THE STUDY OF FUNGI
are octosporous, tmd the sporidia are small, hyaline, and
cylindrical. In Fracchiaea the perithecia are aggregated upon a
sort of stroma like a crust ; the asci, except in one species, con-
tain numerous simple hyaline sporidia. Gihbera and Gihheridca
have caespitose perithecia, which in the former are persistently
setulose, and in the latter soon become smooth ; in both cases
the sporidia are septate and hyaline or brown. In the two
remaining genera the habit is more typical, the pustules being
erumpent ; and in Otthia the sporidia are uniseptate, whilst in
Cucurhitaria the sporidia are typically muriformly septate and
coloured, whilst in small sul)genera tliey are either continuous
or septate and brown.
The ninth subfamily includes the Su^Krficiales of Fries, in
which the perithecia are quite distinct, and superficial or
nearly so, aggregated together or scattered. These may be
further subdivided into the Byssisedae, Villosae, Bosclliniac, and
Sordariae. In the Byssisedae the perithecia are seated upon a
byssoid or felted stratum called a subiculum (Fig. 101). The
genus Byssosphaeria contains species in which the perithecium is
smooth or naked, with the sporidia
varying according to the several sub-
genera, from simple or continuous to
multiseptate or muriform, and hyaline
or coloured. In Chactospheria the
perithecia are villose, and also seated
,,(f^—^-.^.* ,..,,,;. upon a subiculum. In Villosae there
'^'j^^i*' is no subiculum, l)ut the perithecia
Fig. 101.— Bijssosphaeria, with are woolly, downy, or setulose. The
section of perithecium, ascus, . . , . . ^ . .
and sporidia. principal genus IS Lasiosphaeria, m
which the sporidia are hyaline, or but
slightly coloured. In Coniochacta the sporidia are distinctly
coloured. In Vcnturia the perithecia are membranaceous'
and setulose, growing for the most part on leaves, and
the sporidia are oblong and hyaline, continuous or septate.
Chaetomium is a genus easily recognised, Ijy its i'ragile
bristly perithecia, diflluent asci, and continuous brown
sporidia, which are often almond-shaped. Rosellincae almost
corresponds to the Deniidatae of Fries, with the perithecia
almost or quite superficial, smooth and naked, and mostly
CAPSULAR FUNGI— PYRENOMYCETES 215
carbonaceous. The numerous genera which are recognised
by Saccardo, having hyaline sporidia, have been reunited
under the genus Psilosphacria, the different forms of sporidia
being recognised by subgenera. Another genus, Roscllinia,
includes the species with coloured sporidia without septa ;
and Mclanomma those species with coloured septate sporidia,
in conjunction with the smaller genus Olileria. Sporidia
which are divided in both directions, so as to be muriform,
fall under the genus Strickeria. In appearance the sub-
division of Sordarieae is almost equal to Bosellineae ; but,
instead of being carbonaceous, the perithecia are membrana-
ceous, and nearly all the species flourish on dung. In many
instances the sporidia are involved in a gelatinous coat.
There are really but two distinct genera, although, on account
of modifications in the sporidia, several subgenera are accepted,
which some authors have raised to the rank of genera.
Sordaria is, as a whole, a very characteristic genus, which the
late Dr. Winter made the subject of a monograph. The
sporidia in the majority of species are large, elliptical, and
brown, sometimes with a hyaline tail, but in one subgenus
they are uniseptate. The other genus, Sporormia, has a
peculiar form of fruit which calls to mind Perisporium ; most
of the species are very minute, almost invisible without a
strong lens, and the sporidia are brown and septate, readily
breaking up at the joints.
In the old arrangement of the Simple Sphaeriaceae adopted
by Fries, the Svperficialcs were succeeded by a group of
genera in which the perithecia were smooth and half immersed
in the matrix. The base of the perithecium being flattened,
they possessed a more or less conical form, and were occasion-
ally only adnate, although apparently half immersed. These
were the Fertusae, which we constitute the tenth subfamily ;
and the typical genera are Conisphaeria, in which the sporidia
are hyaline ; and Amphisphacria, in which the sporidia are
coloured. Another genus, under the name of Ticotliecium,
includes minute species, found growing upon Lichens. Such
species as conform in general habit and appearance, but with
muriform sporidia, find a place in the genus Teicliospora.
This last subfamily leads almost imperceptibly to the two
2i6 INTRODUCTION TO THE STUDY OF FUNGI
following, ill which the peritliecia are more and more im-
mersed, until they entirely disappear in the matrix. These
are the Lopliiostomaccac and the Endoxylcac, the Ceratostomeae
being rather a parallel than a consecutive series.
The Lophiostomaceae find a place in the Syllogc as a sepa-
rate group, and are regarded as a link between Hij&Uriaceac
and Sphaeriaccac, on account of the broad compressed mouth of
the perithecia ; there is, however, no real affinity, but rather
analogy, between them. This subfamily includes the Sphaeria
platystomae of Persoon, and consists of species which are simple
in habit, with the base of the perithecium liattened and
adnate, or partially imbedded in the matrix, so as to be sub-
superficial. In texture they are mostly hard and carbon-
aceous, with the broad ostiolum more or less compressed,
opening by a very narrow fissure. The sporidia are very
variable, except that the hyaline continuous form is absent.
There is but one genus in all the seven sections based upon
the character of the sporidia, according to Saccardo's arrange-
ment, but which we unite in three genera — Lophiospliacra, with
hyaline sporidia ; Lopliiostoma, with coloured sporidia ; and
Lophidium, with muriform sporidia.
Parallel with this and the succeeding subfamily must be
placed the Ceratostomaceae, in which the perithecia are usually
immersed, but sometimes subsuperficial, either somewhat car-
bonaceous or almost membranaceous — two genera to the former,
and one to the latter. In Ceratostomella the sporidia are
hyaline, and in Ceratostoma coloured. In their entirety they
were formally included by Fries in his Sp)lici&ria Ceratostomac,
on account of the elongated beak-like ostiolum. The remain-
ing genus G-iiomonia, with submembranaceous perithecia,
contains minute species, usually growing on dead leaves or
petioles, with the long ostiola protruding like bristles. The
habit is the same in the subgenera, but the sporidia are con-
tinuous, uniseptate, or rarely triseptate, or still more rarely
thread-like, but always uncoloured.
In sequence from the Lopliiostomaccac follows the thirteenth
subfamily, Emloxyleae, which corresponds to the Sphacriae
immersae of Fries. The perithecia are immersed in the
matrix, usually rotten wood, with only the short erumpent
I
CAPSULAR FUNGI— PYRENOMYCETES 217
necks of the perithecia visible on the surface. The small
genus Endoxyla includes such species as have sausage-shaped
sporidia, which are slightly coloured. The bulk of the species
are included in Xylospliaeria, having sporidia somewhat
elliptical, either continuous or septate, and brown. Here
_ again the subgenera include the various types of sporidia, for
in Anthostoma they are continuous, in Phaeosperma uniseptate,
in Kalmusia three, or more, septate ; and those with muriform
sporidia are relegated to be a distinct genus under the name
of ThyricUum, on account of a sort of effused woody stroma
in which the perithecia are immersed.
From these lignicolous or wood- inhabiting groups we
pass to the subfamily Ohtedac, which almost corresponds to
the Sphaeriae oUcdae of Fries. The perithecia are innate,
growing on bark, and covered by the cuticle. One of the
most notable genera is Massaria, in which the sporidia exude
from the perithecia, and blacken the matrix around the ostiola.
The sporidia are nearly always involved in a hyaline gluten,
and many of them are very large and beautiful. In the
typical section they are two, or more, septate, and coloured ;
but in the subgenus Massariella only uniseptate, and in the
subgenus Massarina multiseptate, but uncoloured. The sub-
genus Pleomassaria includes such species as have muriform
sporidia. In fructification this genus corresponds to Pseudo-
vcdsa amongst the composite Pyrenomyceteae, but differs, of
course, in the distinctly scattered perithecia and the mucous
envelope of the sporidia. In the genus CnjptosjjJiaeria we
encounter a remote resemblance to the Putypeae ; for the
perithecia are densely gregarious, sometimes in patches of
some inches in length, but without stroma of any kind. The
perithecia are smooth and naked, and the sporidia sausage-
shaped and hyaline. In EncJinoa the perithecia are fewer and
scattered, but externally pilose. Another genus which in-
cludes only species with hyaline continuous sporidia is
Physalospora, in which the perithecia are scattered over
twigs and branches, covered by the cuticle, so as only to be
recognised by the slight elevations with the central dot of the
ostiolum. In Endoplilaea the habit is similar but the sporidia
different, being one or more times septate ; as, for instance, in
2iS INTRODUCTION TO THE STUDY OF FUNGI
the subgenus DUlymdla uniseptate, and in Metasphaeria
multiseptate. The small genus Ophiobolus corresponds to the
genus Rcq^hidospora in the Caulicolae, in which the sporidia
are filitbrni. Of genera with coloured sporidia Anthostoma
includes those which are continuous ; and Didymosphaeria
those in which the sporidia are uniseptate. Leptosphaeria has
the sporidia multiseptate, but the coloration is often very-
slight. Other species of Lcptosphacria which grow on
herbaceous plants will be found under Heptameria in the
Caulicolae ; in fact, the separation of the two families Ohtcdac
and Caulicolae is a purely arbitrary one. Delacourea, again,
includes such species of Plcospora as are found growing on the
twigs of trees and shrubs, having coloured muriform sporidia.
The fifteenth subfamily, Caulicolae, includes such species as
conform in habit to OUectae but are found on the dead stems
of herbaceous plants. In the genus Phomatospora the sporidia
are continuous and hyaline, hence corresponding to Physalo-
spora. In Apiospora the rather unusual form of sporidia is
found in which, though practically uniseptate, the lower cell is
small, consequent on the septum being placed near the base.
In Diclymella the sporidia are uniseptate and hyaline, and
correspond to the subgenus Didymclla of Endoplilaea. Mcta-
sphaeria is a large and important genus of Caulicolae, in which
the sporidia are multiseptate and hyaline. Raphidospora in-
cludes such caulicolous species as possess very long thread-like
sporidia, which may or may not be divided by transverse
septa into short joints. Of species which have coloured
sporidia those in which the sporidia are continuous will be
found in the genus Anthostomella, those with the sporidia
uniseptate in Didymosphacrella, and with the sporidia multi-
septate in Heptameria, corresponding to Lrptosphaeria. This
is one of the largest genera on herbaceous plants. Pleospora
is a well-known genus in which the perithecia are often
large, and become erumpeut. The sporidia are muriform and
coloured. Some species or other may be anticipated on the
dead stem of almost any herbaceous plant. The genus Pyreno-
p)liora differs from Pleosjjora in the perithecia being selulose,
but the sporidia the same (Fig. 102).
The sixteenth subftimily is the Foliicolac, which, as the
CA PS ULA R FUNGI-P YRENOM YCE TES
219
Fig. 102. — Asci and sporidia of
Pleos2)ora.
name suggests, are normally found growing on leaves, some-
times living and sometimes dead. The former are more or
less destructive parasites, and are found not only on the leaves
of trees and shrubs, but also on ^^
herbaceous plants, and on the fronds
of Cryptogams. The perithecia are
immersed in the substance of the
leaf, sometimes scattered, but often
in groups. They are thin and
membranaceous, usually minute, and
perforate at the apex. When grow-
ing upon living leaves they are
mostly seated on discoloured spots,
caused probably by the delicate
mycelium destroying the vitality of
the cells. Similar blanched spots,
with analogous minute perithecia
seated upon them, are to be met
with in several genera of 8'phaeroj}-
sidcae, such as Phyllostida, Scpioria,
etc. ; but the asci are absent, and the minute sporules
are produced at the tips of slender sporophores. In certain
cases these are believed to be in some manner connected
with foliicolous Sphacriae, as a stylosporous or imperfect
condition, but how they are connected has not yet been
determined. Of the genera in this subfamily Laestaclia has
the sporidia continuous, or without septa ; in Sphaerclla they
are uniseptate, and are very numerous in species. In
Sphacrulina the sporidia are either three or many septate.
Under this genus are included the foliicolous species of the
genera Metasp)liaeria and Liptospliacria, so that in some the
sporidia are slightly coloured. In one other genus, that of
Linospora, the sporidia are very long and thread-like.
The last subfamily is that of Microthyriaceae, and this is
somewhat of an outside group, as the perithecia are different in
form and structure, being superficial, or nearly so, either membran-
aceous or somewhat carbonaceous, dimidiate and flattened almost
like a shield, formed of radiating cells, and either pierced with a
pore in the centre or with out one. The genus Microthyrium seems
220 INTRODUCTION TO THE STUDY OF FUNGI
to be almost analogous to that of Asterina in the Perisporiaccac.
In this genus the perithecium is flat and membranaceous.
Amongst the subgenera, J/t/zoco^jj-oti has continuous sporidia; in
the typical subgenus they are uniseptate, and in Seynesia the
uniseptate sporidia are brown. In the genus Clypeolum the
perithecia are carbonaceous, and shield-like, with uniseptate
hyaline sporidia ; or in the subgenus Vizella continuous and
brown. In the subgenus Scutellum they are septate and coloured.
The two remaining genera are Micropeltis and Pcmjjhidium ; in
the former the perithecia are convex and the sporidia multi-
septate and hyaline ; in the latter the perithecia are scutate,
the nucleus gelatinous, and the fusiform sporidia brown.
Polymorphism of a somewhat elaborate form has been
credited to some of the Pyrcnomycctcac. Take, for instance,
the very common Picosjjoiri hcrharum, on dead stems. That
ubiquitous black mould, Cladosporium hcrlarum, has been
recorded as one conidial form. This also has been suspected
of merging into Macrosj)orium commune, which again has been
named as a form of conidia of the same Pleospora. By a
further development the Macrosporium, in some occult manner,
appears with the spores in chains, and then again, under the
name of Alternaria tenuis, is referred to the Pleospora ; so that
three supposed species of Pematieae have been recorded as
conidia of Pleospora herharum. Then again one of the
Spliaerojjsideae, having perithecia, but with stylosporous fruit
and named Phoma herharum, has been called the spermogonia
of the same Pleospora. At the same time it may be asked
what function is ascribed to these spermogonia ; for if, as
M, Cornu has suggested, they are capable of germination, then
tlie small sporules are not spcrmatia, or fecundating bodies, as
the name would imply, but have some other function. What
again are the bodies termed j??/c?iif/ia ? Until the process has
been traced, for at present it is little more than suspected,
accurate phraseology cannot be applied. Conidia, of various
forms, and for the most part of the nature of Hyphomycetcae,
are common enough, and possibly produce a mycelium upon
which perithecia are afterwards developed, but so much is at
present only a matter of faith, which remains for the future to
demonstrate.
CAPSULAR FUNGI— P YRENOM YCE TES
BIBLIOGRAPHY
Saccardo, P. A. " Sylloge Pyrenoniycetuni," iii »S'y//o(;c 7''i(?((/or2t?n, vols. i. ii.
Padua, 1882-83.
" Conspectus generum Pyrenoniycetimi Italicoi'uni Systemate Carpologico
Dispositomm." Atti Soc. Ven. Trent., iv. Padua, 1875.
TuLASNE, L. et C. R. Sclcda Fungorum Carpologia. 3 vols. Folio. Plates.
Paris, 1861-65.
Ellis, J, B., and Everhakt, B. M. The North American Fyrcnomycetcs. Roy.
8vo. Plates. Newfield, 1892.
Berlese, a. N. lames Fimgorum acl usum Syllogcs Saceardimiae — Pyrcnomy-
cctes. Imp. 8vo. Col. plates. Abellini, 1894.
Mo)iografia dei gcneri Plcospoi-a, Glathrospora, c Pyrenopliora. Florence,
1888.
Cooke, M. C. Synopsis Pyrenomycctum. 2 parts. Svo. London, 1884-86.
CuKREY, F. "On the Fructification of certain Spheriaceous Fungi." 4to.
Linn. Trails. London, 1857.
Fabre, J. H. "Essai sur les Spheriacees de Vaucluse." Ann. clcs Sci. Nat.
Paris, 1882.
De Notaris, G. "Prime linee di una disposizione de Pirenomiceti. " Giorn.
Bot. Ital, ii. Florence, 1847.
et Cesati, V. " Schema di Classificazione degli Sferiacei Italici
aschigeri." Co7nm. Soc. Critt. Ital., vol. i.
Winter, G. Die Dcutschen Sorclarieen. 4to. Plates. Halle, 1873.
Von Niessl, G. "Die Arten der Pjrenomycetengattung Sporormia." Ocst.
Bot. Zcit. Vienna, 1878.
KiTSCHKE, T. Pyrenomycctcs Germanici. Svo. Breslau, 1867-70.
PLOVi^RiGHT, C. B. Monograph of British Eypomy CCS. 8vo. Col. plates. Lon-
don, 1883.
GiLKiNET, A. Eecherchcs Morphologiques sur les Pyrenomycctcs — /. Sordariecs.
Brussels, 1874.
ZoPF, W. Zur Fntiuickelungsgeschichte der Ascomyceten — Chaetomium. Halle,
1881.
Leveill:^, J. H. "Organisation et Disposition du genre Erysiphe." Ann, des
Sci. Nat. Paris, 1851,
Jaczewski, a. de. " Classification naturelle des Pyrenomycetes." Bull. Soc.
Myc. de France, vol. x. 1894.
CHAPTEE XVIII
GAPING FUNGI IIYSTERIACEAE
The HystericLceae are related on the one hand to the Pyrcno-
mycdaccac, and on the other to the Discomycctcae, but perhaps
externally most strongly to the former, to which they approach
through the subfamily Lopldostomaccac, whilst their connection
with Discomycctcae must be through such genera as Colpoma
and Triblydium. As the Discomyceteae approach Lichens
through some of the FaUllariacccic, so also do the Hysteriaceae
through Aidoyrapimm and Dichaena. In this flimily the
perithecia are erumpent and then superficial, with a flattened
base, and horizontally extended, either oblong or linear. In
substance they are occasionally membranaceous, but more
often coriaceous, or rather carbonaceous, but very rarely some-
what fleshy. They are for the most part black, and dehiscence
takes place by a long narrow fissure which extends the whole
length of the perithecium. The asci may be tetrasporous or
oetosporous, and sometimes polysporous, but the sporidia vary
in the different sections and genera, from simple and continuous
to multiseptate and muriform. The arrangement proposed by
Saccardo may be accepted as the best yet proposed. The only
objection which could be urged concerns the genera Schizo-
thyrium, Aulograplmm, and Lemhosia, which from our point of
view seem to be scarcely generically distinct, since the differ-
ence lies only in the sporidia, which are respectively con-
tinuous, uniseptate, and hyaline, and uniseptate and coloured.
Great care needs to be exercised, for species of Aulograjjlmm
are at first continuous, as in Schizothyrium, and. may ultimately
be faintly coloured, passing thus into Lemhosia.
The whole family is divided, carpologically , into nine sections.
GAPIA'G FUNGI— HYSTERIACEAE 223
which may be referred to in their order. Hyalosporae, in which
the sporidia are continuous and hyaline, includes the two
genera Schizothyrium and Henriquesia. In the former the
sporidia are minute, and in the latter large ; but the habit is
also different, the species being small and superficial, mostly
occurring on leaves and herbaceous stems in the former ; and
erumpent, Hysterium-like, and arboricolous in the latter. The
analogue of Schizothyrmm, in the Spliaeropsideae, is Ldbrella,
which resembles Schizothyrium in habit and appearance, but
the perithecia are stylosporous, and in some species may prove
to be genetically related. The Phaeosporae differ from Hyalo-
sporae in the continuous sporidia being distinctly coloured. The
only genus is Farlowia, in which the habit is distinctly that of
Hysterium, to which the species were formerly referred, and
the perithecia carbonaceous. The British representatives are
the Hysterium Carmichaelianum of Berkeley and the Hysterium
repajidum of Bloxam, which hitherto are the only species
known. The Hyalodidymae are characterised by hyaline uni-
septate sporidia, and include four genera, in one of which,
Aulographum, the perithecia are membranaceous, already
alluded to ; in Glonimn and Actidium the perithecia are
carbonaceous, in the former being simple or branched, and in
the latter stellate. There is considerable difference in the
habit of the different species of Glojiium, some being linear
and scattered, whilst others are densely agglomerated in compact
heaps. There is still a fourth genus, that of Angclinia, in
which the perithecia are at first somewhat fleshy, and open,
exposing the disc, so that it resembles Ccnavyium. The colour
of the excipulum being reddish, is also abnormal, and the single
species is rather a doubtful member of the Hystcriaccac. The
Phacodidymae, with coloured uniseptate sporidia, includes the
two genera Tryllidium and Le7ubosia. In the former the
perithecia are gaping, with swollen lips, exposing the hymenium
more than usual in this subfamily. Its nearest ally is Try-
hlidiella, in the Pheophragmiae section, and the two supposed
genera differ only in the latter having sporidia with more than
one septum, whilst in the former they are only uniseptate, for
which reason we are disposed to regard the distinctions as only of
subgeneric value. Lemhosia has also been referred to already
2 24 IXTRODUCTION TO THE STUDY OF FUNGI
as representing AulograiJhinm , with coloured sporidia. The
Phaeopliragmiac is the Lirgest section in this subfamily, having
six genera and a large number of species. The genera are
arranged in two groups, in one of which the lips of the peri-
thecia are obtuse and rather distant, and in the other acute
and connivent. In the former is placed TryUidicUa, already
mentioned as a triseptate condition of TryUidium and Hysterium,
the typical genus, in which the lips are so connivent that the
disc is rarely exposed, except when moist. The perithecia are
carbonaceous and even, with three - septate or multiseptate
coloured sporidia. Corresponding species, with sporidia un-
coloured, have been transferred to the genus Gloniella, and
species with muriform sporidia to Gloniopsis and Hystcro-
yraphium. The third genus of Fhaeophraymiae is Bhytid-
hystcrium, in which the perithecia are striate or sulcate, and
resemble a Lichen without a thallus. Hitherto the genus is
confined to South America. In one other genus of the first
division the asci are polysporous, and Bagyca has but one species,
which has been found in Xorthern Europe. The two remain-
ing genera, which constitute the division with thin connivent
lips, are Mytilinidion and Ostreion. In the former the asci are
octosporous and in the latter tetrasporous. Mytilinidioii has the
perithecium, in typical species, vertical and compressed, after
the manner of LojjJiium, with the lips very acute, and firmly
closed. The other genus, Ostreion, originally denominated
Ostreichnion, but, as we think, unwisely changed for insufficient
reasons by Saccardo into Ostreion, has perithecia which in form
resemble an oyster, placed vertically, and resting on the hinge.
The sporidia are four in each ascus, which, in the only recog-
nised species, are very large and multiseptate. The sixth
section is Hycdophragmiae, in which the multiseptate sporidia
are hyaline. As already intimated, Gloniella is analogous to
Hysterium, with hyaline sporidia, and consequently the peri-
thecia are carbonaceous ; luit in Pscudographis the perithecia
are coriaceous, and gaping, with precisely the habit of Try-
Uidium and Tryllidiella. Arranged according to natural
affinities, these three genera would be consolidated, and the
three forms of fruit recognised only as subgeneric distinctions.
The genus Dichaena approaches Lichens in its habit of growing
GAPING FUNGI— HYSTERIACEAE 225
on living bark, aggregated in dense patches, and the perithecia
are elliptical and irregular. A condition may often be met
with in which no asci are developed, but these are replaced by
stylospores. This condition has been referred to Sphaeroiosideac,
under the generic name of Psilospora, whilst other authors
simply mention it as the pyc7iidia of Dichaena. There is little
doubt of its being an imperfect condition of this Hysteriaceous
genus. The section Hyalodidyae includes the one genus
Gloniopsis, in which the sporidia are muriform but hyaline.
The habit and texture is that of Hystermm. In the same
manner, species of the old genus Hysterium which have muriform
coloured sporidia find a place in the section Phaeodictyae under
the genvis Hystcrograplimm,. The ninth section is Scolecospoixic,
in which the sporidia are very much elongated, so as to be
thread-like, or rod-like, and hyaline. The one genus in which
the sporidia are cylindrical, and much shorter than the asci, is
Hypoderma. The perithecia are membranaceous, and flattened,
with a narrow fissure, and the species are most commonly
found on dead leaves, herbaceous stems, and occasionally on
young twigs. Sometimes several perithecia grow on irregular
bleached spots, and these are often accompanied by smaller
perithecia, which contain only minute stylospores, belonging
technically to the Sphaeropsideous genus Leptostroma, but which
are often called the spermogonia of the various species of
Hypoderma. It must not be assumed that the term spermatia,
applied to the minute bodies enclosed in the smaller perithecia,
indicates fecundative functions, since no sexuality has been
proved. In the remaining genera the sporidia are truly filiform.
Lopliodermiuin has the habit and appearance of Hypoderma,
but the sporidia are different, and in like manner the species
are often associated with forms of Leptostroma. Lophium is
a small genus with rather carbonaceous perithecia of a shell-
shape, as mentioned under Mytilinidion, with very acute con-
nivent lips. Sporomega has depressed, and rather coriaceous,
perithecia, with thick gaping lips, which partially expose the
disc (Fig. 103); and Golpoma resembles it in these features,
but differs in being developed beneath the cuticle, which is for
a long time adpressed, or adherent to the lips, and the substance
is softer. In habit it resembles TryUidiuvi rather than
15
226
INTRODUCTION TO THE STUDY OF FUNGI
Sporonuga. The last genuine species is Ostropa, in which the
perithecia are almost sphaeroid, dehiscing above with a
longitudinal fissure, thus suggesting relationship with the
Lo'phiostomaceae. The rather aberrant
genus Rohergea is sometimes placed
here, as having affinity with Ostropa ;
but Acrospermum appears to us to
be more closely related to Hgpocreaceae,
from the absence of any apical fissure.
The total number of species in
this subfamily may be accepted as
450, and of these some have a very
wide geographical distribution. It is
an open question whether the family
is most closely allied to the Pyrcno-
mycctcac or the Discomyceteae. The
consolidation of the hymenium into
a disc, and the strong development of the paraphyses, indicate
relationship with Bisconiycetes, and this is supported by the
almost universal subsuperficial habit. This is not, however,
a question of practical importance, and may be compromised
by placing the Hystcriaccac as an intermediate group between
the Pyrenomycctcs and tlie Discomycetcs.
Fig. 103. — Sporomega with
enlarged perithecium and ascus,
IBLIOGRAPHY
Saccardo, p. a. "Hysteriaceae," in SaccarddlSylloge Fungorum, vol. ii. Imp.
8vo. Padua, 1883.
DuBY, J. E. MimoircsurlatrLhidesHystcrindes. 4to. Plates. Geneva, 1861.
De NoTAiiis, G. "Prime linee di una disposizione de Pirenomiceti Isterini," in
Giorn. Bot. Ital., vol. ii. Florence, 1847.
Billings, J. S. "The Genus Hysterium and some of its Allies." American
Naturalist, vol. v., October 1871. Salem, Mass.
Rehm, H. "Revision der Hysterineen," in Tfcri Diiiy. Dresden, 1886.
Massee, G, " Hysteriaceae," in /.ViYis/i J'»?)r/?/s i^'forff, vol. iv. Svo. London,
1895.
CHAPTER XIX
CONJUGATING FUNGI PHYCOMYCETES '
The comparatively small order of fungi known as the
Phycomycetes has been subjected to more exact examination
and clearer definition than when Berkeley called it Physomycetes
in 1857, and limited it by the definition of "fertile cells,
bladder-shaped, scattered on the threads which are not com-
pacted so as to form a distinct hymenium. Sporidia in-
definite, formed from the protoplasm of the cells." It will be
observed that this diagnosis only takes cognisance of the
superficial, and asexual, reproduction by gonidia, produced
within sporangia, the latter being scattered over the threads,
as in typical forms of Mucor.
Technically, as at present recognised, the Phycomycetes are
characterised by a vmicellular mycelium, often parasitic on
plants or animals, sometimes saprophytic, developed in the air
or in water. Reproduction sexual or asexual. Sexual by
oogonia and antheridia, or by conjugation, producing zygospores.
Asexual by means of gonidia or zoospores. Many of them
resemble, more or less, the moulds in external appearance,
having conspicuous hyphae, arising from a procumbent creep-
ing mycelium — but that the latter threads are without septa,
and the former may either produce sporangia or naked gonidia.
Moreover, they are fm-ther removed from the moulds by
possessing, in addition to the conspicuous agamic reproduction,
a true sexual method by means of oogonia fertilised by
antheridia, or by zygospores resulting from conjugation of
specialised branches.
Undoubtedly there is considerable variability in the
external features of the different families constituting this
228 INTRODUCTION TO THE STUDY OF FUNGI
order, which includes the old typical Mucors and their
allies — almost the sum total of the Physovujcetes of former
times ; and also the Peronosporeae, or rotting moulds, previously
classed with the Hyphomycetes ; as well as the " white rusts "
{Cystopi), formerly united with Uredines. In addition to these
are the fish moulds, or Saprolegniaceae ; and the insect moulds,
or Entomophthoraceae ; so that altogether there are four very
distinct families, with well-developed hyphae ; to which must
be added two other, inferior, groups, in which the hyphae are
obsolete, for a long time regarded as outside families, with
obscure affinities, but remotely associated with the Uredines
under the names of Chytridium, Synchytrium, and Frotomyces.
This association of apparently rather heterogeneous elements
is held together by the conservating bond of a dimorphic
reproduction ; otherwise their relationship is, at first sight, so
obscure that it will be necessary to advert to each family
separately.
It has already been intimated that four of the families
possess a conspicuous vegetative system, in a unicellular
creeping mycelium, giving rise to erect, simple, or branched
threads, which bear the conidia, or otherwise asexual repro-
ductive organs. Although agreeing in this, the organs them-
selves differ considerably in the four families, inasmuch as the
Mucoraceae develop cysts, or bladder-like cells (sporangia),
which enclose either many or only one reproductive cell, or
gonidium. These cysts are terminal on the fertile hyphae,
and may be produced singly or in clusters. In the Perono-
sporaceae there are no true cysts, but naked gonidia, which
may be passive, and germinate directly, or their contents may
become differentiated into zoospores, or zoogonidia ; that is to
say, active ciliated zoospores, which at length become passive
and germinate. The Saprolegniaceae are aquatic, and produce
zoospores within the changed hyphae ; and the Entomoph-
tlioraceae are parasitic on insects, and develop single conidia
on short sporophores. Thus it will be seen that there are
distinct features in the asexual reproduction of the four
families sufficient for their discrimination. It may be added,
as a further distinction, that the Mucoraceae are saprophytic
on dead animal or vegetable substances. The Peronosporaccae
CON JUG A TING FUNGI— PHYCOM YCE TES
229
Fig. 104. — Ilucor.
are parasitic on living vegetables. The Saprulcgnimeae are
al^uatic. And the Entomophtlioreae are entomogenous.
The Mucoraceae might be mistaken for moulds if not more
closely examined. The erect threads are not conidia-bearers,
but sporangiophores, because they support
sporangia at their tips ; and these sporangia
are nearly globose cells of thin membrane,
which enclose the spores, or reproductive
bodies (Fig. 104). When fully matured the
sporangium is ruptured and the enclosed spores
escape. This is the ordinary asexual repro-
duction of the Mucors, and all that was really
known of them half a century ago. In some
cases the fertile branch, or sporangiophore, is
prolonged into the interior of the sporangium, and becomes a
columella.
The sexual reproduction of the Mucoraceae is accomplished
by zygospores, resulting from conjugation, and hence they are
sometimes characterised as Zygomycetes. In many of the species
this form of reproduction has never been traced, but has been
accepted from analogy. Two lateral branches resembling each
other, and termed archicarps, are concerned in the process.
They resemble at first ordinary branches, which approach each
other until the tips meet, but as they increase in size they
become clavate, and
are densely filled
with protoplasm.
At length the ex-
treme portion of
each archicarp is
separated from the
basal portion by a
transverse septum,
each portion acquir-
ing a distinctive
name, the basal cell
being termed the
suspensor, and the apical cell the gamete (Fig. 105). At the
point where the two gametes meet the separating cell-walls are
///
Fig. 105. — Formation of Zygospore,
230 INTRODUCTION TO THE STUDY OF FUNGI
soon dissolved, so that the contents coalesce, and a single cell is
constituted from the union of the two gametes, which is, in
effect, the young zygospore. Henceforward the cell-wall of
the zygospore becomes thickened, and coloured, so that it
usually acquires a brown colour, and is warted or spinulose
when mature. The zygospores thus formed are also character-
ised as resting spores, because they are capal)le of resting, or
remaining in a dormant condition for months before germina-
tion takes place, usually tli rough the winter, becoming active
in the spring.
It may sometimes happen that the two gametes, instead of
coalescing, remain distinct ; or in rare cases only one archicarp
is produced ; yet in such instances a body resembling a
zygospore is developed, without conjugation, and therefore the
resultant spore is called an azygospore. In some species the
zygospores are produced freely, and in considerable number,
amongst the ordinary vegetative hyphae, at the same time, or
succeeding, the production of the ordinary gonidia ; but the
latter germinate at once, without any period of rest, and
hence they do not survive through the winter. Usually the
development of gonidia is arrested in the autumn and the
formation of zygospores commences. In the absence of
zygospores the mycelium becomes perennial, and thus survives
the winter, so that the species may be perpetuated. Provision
is thus made for reproduction, asexually, by means of gonidia,
which germinate at once ; l)y azygospores, which germinate
after a period of rest, and sometimes by a perennial mycelium,
which sm-vives the winter ; and sexually by the production of
zygospores, which accrue from the conjugation of two approxi-
mating specialised branches of the hyphae.
The Feronosporaceae are, in their typical forms, more nearly
resembling the Mucedines in habit than are the Mucoraceae.
The erect hyphae, or gonidiophores, are usually furcately
branclied two or three times, and bear the gonidia (Fig. 106), as
more or less elliptical hyaline bodies, at the ends of the branches.
These constitute the means of asexual reproduction, and may
be developed successively or simultaneously. In some cases
the gonidia so produced appear to be simple gonidia, but in
others they undergo transformatit)n ; in the former case they
CONJUGA TING FUNGI--PH YCOM YCE TES 23 1
germinate at once through a lateral pore, but in the latter
case an intermediate stage intervenes. Each gonidium when
mature has more turbid contents, which are seen to accumulate
in several centres, and then to become divided by the growth
of a membrane about each segment, into distinct inner cells,
each with a nucleus.
Soon the wall of the
mother cell is rup-
tured, and the con-
tents escape, now
differentiated into
three or four, or more,
smaller but similarly
shaped bodies, armed
with a pair of vibra-
tile cilia, by means
of which they move
actively in any drop
of moisture with
which they may come
Fig. 106. — Gonidiopliore of Peronaswora.
m contact, trans-
formed into secondary gonidia or zoogonidia, sometimes called
zoospores. In this condition they move about for some time
over the moist surface of the leaf upon which they are
discharged, until at length they come to rest, lose their cilia,
and commence germination, by the production of a delicate
germ-tube which enters the stomata of the fostering plant,
and form a mycelium beneath the surface. From this my-
celium spring erect hyphae, which seek the air, and, becoming
branched, constitute new gonidiophores, and the cycle is
complete.
The sexual reproduction takes place within the host-plant,
by the production of oogonia upon branches of the mycelium.
They originate as spherical swellings at the end, or inter-
calated in the hyphae, and after a time reach a considerable
volume, and contain a dense protoplasm with oil drops. Soon
these swellings are isolated by the production of a septum
across the hypha when terminal, or above and below when
intercalary. After this differentiation of the oosphere takes
232 INTRODUCTION TO THE STUDY OF FUNGI
place, by the concentration of the denser portion in the centre,
in the form of a sphere, involved in a delicate membrane,
surrounded by a hyaline layer of protoplasm. At this time
the antheridmm is being developed from a lateral branch of
the hypha, just below the oogonium (Fig. 107). When fully
developed tliis organ is elliptical or obovate, smaller than the
oogonium, and filled with granular protoplasm ; cut off at the
base from connection with the hypha by a transverse septum.
Assuming that the oosphere is formed, and the antheridium
perfected, the latter comes in contact with the former, and, at
the point of contact a slender tube is projected through the wall
of the oogonium, which grows until it reaches the surface of the
Fig. 107. — Perijaosj)ura. o, young state ; h, formation of oosphere ;
c, after fertilisation. After De Bary.
oosphere. Meanwhile the contents of the antheridium under-
go change : towards the centre they are more dense, and, as the
fertilisation tube becomes complete, this portion passes down
it and mingles with the protoplasm of the oosphere, and the
connection is complete. The oosphere is fertilised, and, secret-
ing a thick wall, becomes an oospore, a resting spore, analogous
to the zygospore of the Mucoraceae. Gradually, by the decay
and dissolution of the hyphae, these oospores become free,
hybernating amidst the decaying tissue of the foster-plant,
and awaiting rejuvenescence in the spring. When the latter
period arrives the contents of the oospore, in most species,
become differentiated into a host of minute active zoospores,
similar to those evolved from the differentiated gonidia, and,
by rupture of the wall of the mother cell, become diffused and
ready to attach and establish themselves upon young seedlings
CONJUGA TING FUNGI— PH YCOM YCE TES
233
of their favourite host. Thus, by asexual reproduction during
the summer, and by hybernation of the oospores through the
winter, provision is made for the continuance of the species.
Incidentally, it may be observed that in one genus, that of
Cystopus, the habit of the parasites is but little like that of the
residue of the Peronosporaceae : the conidiophores are extremely
short and simple, and the con-
idia are produced in chains ;
but the sexual reproduction by
oospores is virtually the same,
and this is almost the only
link which unites them. It is
worthy of remembrance here
that there is manifest in this
group a gradual loss of sex-
uality, although external feat-
ures remain much the same.
The Saprolegniaceae, from
their aquatic habit, would at
first seem to have an afiinity
with Algae, rather than with
Fungi, but this is rather
analogy than affinity. The
hyphae, in most cases, are
modified at their extremity,
and become zoosporangia,
which are elongated cells
separated from the rest of the
hypha by a septum (Fig. 108).
After the zoogonidia have
escaped through an opening at
the apex, the hypha, or stem,
continues to grow up through
the empty sporangium, and
forms a second sporangium, and this, in like manner, when the
zoogonidia are discharged, may enclose a third, so that upon
old threads it is not unusual to see the remains of two or three
empty sporangia, the one within the other. The zoogonidia
are produced in great numbers, in each zoosporangium, having
-Sporangia and zoospores of
Saprolegnia.
234 INTRODUCTION TO THE STUDY OF FUNGI
the common ovate form, with a pair of active cilia at tlie
smaller end. At maturity they escape by an orifice at the
apex, and swim freely and actively in the surrounding water.
In addition to this asexual reproduction, there is a more com-
plex system of sexual reproduction, by means of antheridia and
oogonia, the resultant oospores, or resting spores, serving to
carry the germs through the winter and provide for their
appearance in the spring. In typical species the sexual appa-
ratus is of this kind, the oogonia are globose cells, generally
terminal on short branches of the mycelium (Fig. 109). The
external membrane is absorbed at various points, leaving it
pierced with rounded holes. The pro-
toplasm becomes divided into a greater
or less number of distinct portions,
which become rounded into little spheres
and separate from the walls of the cell
to congregate in the centre, where they
float in a watery fluid. During the
formation of the oogonium, there arise
from its pedicel, or the neighbouring
filaments, thin curved branches, which
^ , „„ ^ . .^, are sometimes twisted round the pedicel.
Fig. 109.— Oogonium with . ^ ^. .
two oospheres of xicWya. but tend towards the oogonium. Their
After DeBary. ^^^^ extremity is closely applied to the
wall, and becomes slightly inflated above, and cut off below by a
septum. It is then an oblong cell, or antheridium, filled with
protoplasm. Each oogonium possesses one or several of these
antheridia. Towards the time when the oospheres are formed,
each antheridium projects into the interior of the oogonium one
or more tubular processes, which are applied by their extremities
to the nearest oosphere. They have not been seen to open,
nor has anything like a discharge of protoplasm been observed.
Afterwards the oospheres become covered with cellulose, and
are converted into so many oospores. When they have arrived
at maturity these oospores possess a tolerably thick integument,
which is double, and, after a considerable period of rest, they
develop germ tubes or sporangia direct.
The Entorrioj)hthoraceae are minute parasites which inhabit
the bodies of small flies and other insects, and are " distin-
CON JUG A TING FUNGI— PH YCOM YCE TES
235
guishecl by the production of numerous hyphae of large diameter
and fatty contents, which ultimately emerge from the host in
white masses of peculiar appearance, producing at their ex-
tremities large conidial spores, which are violently discharged
into the air and propagate the disease.
In addition to these conidia, the propa-
gation of the fungus, after long periods
of rest, may be provided for by the
formation of thick-walled resting spores,
adapted to withstand successfully the Fig. 110.— Hyphai bodies.
most unfavourable conditions. These After Thaxter.
resting spores, which may be either sexual {zijgosiwres) or
asexual (azygospores), finally germinate and produce conidia
that are discharged in the usual fashion, and serve to infect
fresh hosts." ^
Infection results from contact of one of the conidia which
adheres to the surface of the host, germinates there, and the
germ-thread enters the body. After
entering, growth proceeds rapidly
and forms " liyphal bodies " (Fig.
110), which are short thick fragments
of variable size and shape, continually
reproduced by budding, until the
body of the host is more or less
completely filled with them. Having
absorbed the contents of the body,
these hyphal bodies germinate, each
one producing one or more threads,
which proceeds directly into the
outer air, and bears its conidia, or
it branches indefinitely, each branch-
let producing spores at the extremity
(Fig. 111). These are the conidia-
bearers, and their results the simple
asexual reproduction. Conidia are
formed by constriction or budding,
and when fully matured are forcibly ejected to a considerable
distance. The discharged conidium germinates at once, but,
1 Thaxter On the Entomophtlwreac of the United States, 4to, 1888.
Fiu. 111. — Conidiopliores of
Eiitomophtlioi-a. S. P. C. K.
236
INTRODUCTION TO THE STUDY OF FUNGI
failing to reach a suitable host, a secondaiy conidium is formed,
resembling that from whence it was derived (Fig. 112).
Should this second fail in finding a suitable host, a third is
formed from the second in the same manner.
The other form of reproduction is by means of resting
spores, which may be sexual or
asexual, and proceeds also from the
hyphal bodies. The latter, or
azygospores, are formed by the
conversion of a hyphal body into a
resting spore, or by direct budding
tlierefrom (Fig. 113). They are
usually spherical, rather large, sur-
rounded by triple walls. Sexual
resting spores, or zygospores, are
produced as the result of conjuga-
tion of opposite threads. Threads
either within or without the body
of the host produce lateral ovit-
growths, at opposite points of two
different threads, which meet mid-
way and coalesce. The intermediate
walls are absorbed, a connective is
formed, and the contents are mingled. A bud is produced upon
the connecting canal, which appropriates
the contents of the two conjugating
cells, and the zygospore is formed.
After this the empty hyphae disappear.
Earely another modification of conjuga-
tion takes place. The hyphal bodies
join laterally, by means of short pro-
cesses, and ;[)roduce an expansion at tlie
point of union, which enlarges and
absorbs the contents of the two con-
jugating bodies, and thus a resting spore,
or zygospore, results. Some other slight
modifications take place, according to
the species, but the general character is
the same (Fig. 114). The mature resting spores oare spherical.
Fu.. 112. — Secondary simres of
Entomophthora. S.P.C.K.
Fi(i. 113. — Coujugatiug
liyphal bodies. Thaxter.
CONJUGA TING FUNGI— PHYCOMYCETES
^37
smooth, slightly coloured, but their ultimate history and
development are still rather obscm'e.^
By comparison of this family with the preceding it will be
observed that whilst the gonidial reproduction resembles most
that of the Peronosporaceae, it is by no means the same : the
gonidiophores are less highly developed, and active gonidia, or
zoogonidia, would seem to be absent. The conjugation also
differs from that of all the other families, and approaches more
closely to that of the Algoid type, as represented by some of
the filamentous Conjugatae.
From this summary of the main features of the four
normal families, we must turn to the
two remaining families, which are so
far abnormal, or aberrant, as to be
deficient of conspicuous hyphae. The
Chytridieae are mostly very minute,
and either parasitic or saprophytic,
forming sporangia of characteristic
forms, the contents breaking up into
swarm-spores. These zoogonidia, or
swarm-spores, escape from the spor-
angium, through a narrow opening,
usually at the apex. Nowakowski "
has given the life -history of one
species, which is parasitic upon
Euglena. In this species of Poly-
phagus, the swarm-spore, when it has
come to rest in the water, becomes
spherical in shape, and at once puts
out hair-like, tubular rooting (rhizoid) processes in indefinite
directions. If one of these encounters a resting Euglena
it penetrates into its body, destroying and exhausting
it to supply food to the parasite. The parasite then
begins to increase in size, the tubes become larger and
thicker, and new ones are formed which throw out branches,
and attack and destroy any new Euglenae which they encounter.
114. — Conjugating hyphae
in Entomophthora, with
zygospores. After Thaxter.
1 Vegetable Wasps, etc., by M. C. Cooke, 1892, p. 10.
^ Beitrag zur Kenntniss der Chytridiaceen, von Dr. L. Nowakowski, Breslau,
1876.
238 INTRODUCTION TO THE STUDY OF FUNGI
In this way a luueh-branched plant is formed, with hair-like
terminal brauchlets, which connect with the larger main stems,
and through these with the body of the original spore ; the
latter lias grown in the meantime into a large round or elon-
gated vesicle at the expense of the Euglenae, which have been
exhausted by the rhizoids. When it has reached a certain
size, varying according to the food which has been supplied to
it, it shows itself to be a sporangium. It grows out at one
spot into a bluntly and irregularly cylindrical thick tube with
a delicate membrane, into which the whole of the protoplasm
passes, and is at once divided into swarm-spores. This process
of development may be repeated for many generations, and
leads to an immense multiplication of individuals, if there is a
sufficient number of Euglenae within reach. When this has
taken place, the course of events changes. The young plants
remain for the most part small and become gametes, which
conjugate in pairs, each pair forming a zygos'pore, and these
behave as resting spores. Of the two conjugating gametes, the
one which is the supplying gamete has usually a round and
larger body, but shows no other apparent difference before con-
tact with the other, the receptive gamete. The latter usually
continues to be smaller, and often very small, and puts out
rhizoid branches, and if one of these encomiters a supplying
gamete it applies its extremity to it as a conjugating tube, and
increases in thickness, while it ceases to grow in length. The
membrane between the conjugating tube and the supplying
gamete disappears at the point of attachment, and an open
communication between them being thus established, the whole
of the united protoplasm of both gametes passes into an en-
largement of the conjugation tube, close to the point of attach-
ment ; the swelling gradually expands into a spherical vesicle,
and, being delimited by a membrane after receiving the proto-
plasm, becomes a thick-walled zygospore. The outer wall
assumes a pale yellow colour, which is in some cases smooth, in
others spinulose. The whole process of forming a zygospore is
completed in from six to seven hours. This zygospore is a
resting spore, and germinates when its period of rest is over,
producing a zoosporangium like non-conjugating plants.^ The
^ British Fungi — Phycomycctes, etc., by G. Massee, London, 1891.
CONJUGATING FUNGI— PHYCOMYCETES 239
genus Synchytri'um is provisionally included in this family,
although no sexual reproduction is known. When compared
with such families as the Mucoraceae and the Peronosporaceae,
the Chytridieae seem to have little in common, save the pheno-
menon of conjugation, and appear to be, in fact, in conjunction
with Protomyceteae, an outside group, of doubtful natural
affinity.
In the Protomyceteae the mycelium is very fugitive, at first
seated in the tissues of the plants upon which the species are
parasitic, and then septate, contrary to the usual condition in
the Phycomycetes. Conidia are unknown. The entire system
of reproduction consists in the development of thick-walled
resting spores. In germination the endospore escapes through
the rupture of the thick wall, in the form of a sporangium,
filled with minute, motionless spores, which conjugate in pairs.
After conjugation the spores germinate by emitting a slender
germ-tube, which enters the foster -plant, and produces a
mycelium, from which resting spores are developed, and the
cycle is complete.
Strongly impressed with the absence of any true natural
affinity between the last two families and the four preceding
ones with which they have been associated, we have no alter-
native but to include them under protest, and to suggest that
the one fact of conjugation, as feebly carried out, is insufficient,
in the absence of other indications of relationship, to warrant
the retention of these two families with the Phycomycetes. As
evidence that their affinities have always been held in doubt,
it may be added that, until very recently, the genera Chytridium
and Synchitrium have been included with Algae,^ although
subject to the observation that " The genus Synchitrium
appears to be more nearly related to Protomyces, amongst
Fungi, than to Algae." Under any circumstances they can
only be regarded as aberrant families, mechanically and pro-
visionally tacked on at the end of this order, until they may
be assigned to a more fitting place.
1 British Fresh Water Algac, by M. C. Cooke, 18S4, p. 198.
INTRODUCTION TO THE STUDY OF FUNGI
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CONJUGA TING FUNGI— PH YCOM YCE TES 24 1
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Soc. Nat. Hist., vol. iv. 4to. Plates. 1888.
Brefeld, 0. Schimmelpilze, iv. 4to. 1873.
Cooke, M. C. Vegetable JVasiK and Plant Worms. Svo. Plates. London, 1892.
Saprolegnieae
Pringsheim, N. " Entwickelungsgeschichte d. Achlya prolifera." N. Acta
Acad. Loop. Car., xxiii.
" Die Saprolegnieen. " Jahrhilchcr filr Wiss. Bot., i. 1857.
"Naclitrage zur Morphologie d. Saprolegnieen." Jalirh. f. Wiss. Bot., li.
1860.
■ • "Weitere Nachtrage," etc. Jahrb. f. Wiss. Bot., ix. 1874.
Leitgeb. "Neue Saprolegnieen." Jahrb. f. Wiss. Bot. , yH. 1869.
Lindstedt, K. Synopsis d. Saprolegniaceen. Svo. Four plates. Berlin, 1872.
CoRNU, M. " Monographie des Saprolegniees. ' Ann. des Sci. Nat., 5th series,
vol. XV. Paris, 1872.
Pringsheim, N. "Neue Beobacht. u. d. Befruchtungsact. v. Achlya u. Sapro-
legnia." Jahrb. f. Wiss. Bot., xiv. Heft. 1. 1882.
Hesse, R. Pythium de Baryanum, ein Endophylischer Schmarotzcr. Svo. Two
plates. Halle, 1874.
Sadebeck, R. " Untersuchungen liber Pythium Equiseti. " Svo. Two plates.
Cohn, Beitr. 1875.
Chytridieae
FiscH, C. Beitrag zur Kenntniss der Chytridiaceen. Erlangen, 1884.
Braun, a. "Ueber Chytridium." Monatsb. d. Berlin. Acad. June 1855.
CoHN, J. "Ueber Chytridium." Nova Acta Acad. Leop. Car.^ xxiv. 1
De Bary u. Woronin. "Beitr. z. Kenntn. d. Chytridieen." Bcr. d. naturf.
Ges. zu Fi'cibicrg, iii. 1S63.
CoRNU, M. " Chytridinees parasites des Saprolegniees." Ann. des Sci. Nat.,
5th series, t. xv. Paris, 1872.
NowAKOWSKi, L. "Beitr. z. Kenntn. d. Chytridiaceen." Cohn's Beitr. z.
Biol., ii. 1876.
16
CHAPTEE XX
ItUST FUNGI UREDINEAE
None of the primary groups in the division of Fungi, as
adopted by Fries, were so unsatisfactory as that of the Conio-
mycetes, which inchided with the Sphaeropsideae and the 3Ielan-
conieae such heterogeneous elements as the Uredineae and the
Ustilagineae. More recently the two latter were combined
under the name of Hypodermeae, and might still be so retained
without grave objection, although they have little save their
parasitism in common. One of the most important divergences
is to be found in the complex character of the fructification in
the Uredineae, as compared with the comparative simplicity of
that in the Ustilagineae. The species form erumpent pustules
on living plants, being furnished with an innate septate
mycelium, but destitute of perithecia or true asci. The
typical fructification consists of spermogonia, aecidia, uredo-
spores, and teleutospores. The Spermogonia usually accompany
the Aecidia, and are punctiform, yellow, orange, brown, or
turning black ; the sporules are very small, and ovoid or
cylindrical, mostly expelled from a pore or orifice, at the apex
of the pustule, in little tendrils. The Aecidia are pale, and
possess a pseudoperidium, mostly in the form of a little cup,
when mature, with a serrate white margin, popularly termed
" cluster-cups " (Fig. 115). The aecidiospores are simple, rather
large, usually orange and warted, produced in chains within the
cups, or pseudoperidia. The uredosiJoriferons sori are variously
coloured, rarely possessed of a pseudoperidium, and mostly
pulverulent. The uredospores are continuous, subglobose, and
hyaline, yellowish, or pale brown, very rarely catenulate, rather
large, germinating from two to six pores externally, and for
RUST FUNGI—UREDINEAE 243
the most part aculeolate or minutely punctulate. The teleuto-
sporiferous sori are also variously coloured, and very rarely
furnished with a pseudoperidium. The teleutos2Jores are either
continuous or septate, generally sup-
ported on a persistent peduncle,
externally smooth, or ornamented in
various ways by spines, warts, gran-
ules, or other appendages, germinat-
ing through determinate pores. By fig. 115.-" Cluster-cups" of
germination of the teleutospores a Aecidia.
promycelium is developed, which is typically four -septate,
bearing sporidiola at the apices of sterigmata. This, therefore,
is the normal sequence of fructification — spermogonia, aecidia,
uredospores, teleutospores, and sporidiola — but some one or more
of the series is often suppressed.
The arrangement adopted most generally is, to a great
extent, an imperfect one, since it assumes a knowledge of the
most important features in the life -history of each species.
This may be all very well for a local flora, where details may
be determined, but it is of doubtful value in dealing with a
mass of exotic species, where special information is not to be
obtained. It is an open question whether all sound classification
should not be based upon characters which may be determined
directly from the individuals themselves, and should include
nothing which is not present or evident in examples of any
given species. The entomologist finds no difficulty in classify-
ing his Lepidoptera, although they may have passed through
previous stages wholly different from that presented by the
imago, and his classification is based upon features to be found
in the perfect insect, and in those alone. Is it so with the
species of Puccinia, for example ? if recent disclosures are to
accepted. There is a certain species called Puccinia phragmitis,
which is found growing upon Phragmitis communis; and there
is a second supposed species called Puccinia Trailii, which
occurs upon the same host ; and there is still a third species
which has been named Puccinia Magnusiana, having the same
habitat. How are these species proposed to be distinguished,
except by the intuitive knowledge that the Aecidium of
Puccinia 2^^iragmitis is supposed to flourish on the leaves of
244 INTRODUCTION TO THE STUDY OF FUNGI
certain species of Rumex, that of Puccinia Trailii on Rumex
acetosa only, and that of Puccinia Mcignusiana on Ranunculus
repens and bullosa. If the Puccinia itself, which corresponds
to the imago stage, in being the ultimate and most perfect
development of a cycle — is insufficient fur the determination,
or discrimination, of Puccinia p)hragmitis, Trailii, and Magnus-
iana, tlien it may be contended that the basis of classification
is greatly defective for all practical purposes. Yet it is not
uncommon to meet with observations associated with one
species, so called, of Puccinia, that " it cannot be distinguished
morphologically" from another species, but that "its life-history
is different." This is the crucial test of the system, for if the
perfect stage of one species is not to be distinguished from
another, or, it may be, from two others, although the system
may be very philosophical, it is nevertheless impracticable.
Let us proceed, however, to the method set forth in Saccardo's
Sylloge, and now generally adopted.
The primary divisions are again based upon the spore
and its septation. The Amerosporae includes all the genera in
which the teleutospores are unicellidar. Of these, two genera
have no pseudoperidium to enclose the teleutospores, namely
Uromyces and Hemileia. The latter is represented by the
destructive coffee -leaf disease, the assumed teleutospores of
which have half the surface smooth, and half warted. Its
association here is scarcely more than provisional. Uromyces
is the great genus of the section, and the teleutospores
germinate by one pore. In typical species the spermogonia,
aecidia, iiredospores, and teleutospores are all present on the
same host-plant. In others the spermogonia and aecidia are
unknown. In another section the spermogonia, aecidia, and
teleutospores are present on the same host, but the m^edospores
are unknown. In the final section only the teleutospores are
known, the spermogonia, aecidia, or uredospores never having
been discovered or recorded ; this is, in effect, the section which
includes the imperfectly known species. The two genera in
which the uredospores are included in a pseudoperidium are
Melampsora and Melampsorella, very closely related — the sori
of the teleutospores being crust-like, blackish, and determinate
in the former ; flattened, indeterminate, and pallid in the
RUST FUNGI— UREDINEAE 245
latter. In the one remaining genus of the section, that of
Cronartium, the teleutospores surround a vertical columella,
but in external appearance the species resemble Ureclines
as little as possible, and are not unlike persistent tendrils of
some member of the Melanconieae.
The Didymosporae are well typified by the large and
important genus Puccinia, with its transversely septate teleuto-
spores. The grades are like those of Uromyces, viz. spermo-
gonia, aecidia, uredospores, and teleutospores ; but a different
element comes into the subsidiary grouping, since it is con-
tended that in some cases these grades are not all passed upon
the same host. It is perfectly true that in one typical group,
Fig. 116. — Section of aecidia aud spermogonia. S.P.C.K.
that of the Auto-'pucciniae, all the grades are developed on the
same species of plant ; but in the Hetero-pucciniae, although
all the same grades are affirmed to be present, yet the spermo-
gonia and aecidia appear on one species of plant, generally a
Dicotyledon ; whilst the uredospores and teleutospores make their
appearance on a plant belonging to quite a distinct genera of
plants, mostly a Monocotyledon. Now the doctrine which
associates these forms is that designated Heteroecism, and
although some writers contend that the facts are not effectually
proved, and therefore dispute the conclusions, others accept
the inferences derived from artificial cultures as conclusive,
and bow down to Heteroecism. Whichever may ultimately
succeed in persuading the rest, it is manifest that Heteroecism
for the present is the favourite, and consequently the faithful
are happy in finding the four grades separated, two on one
kind of matrix and two on another. To quote a very familiar
example, the spermogonia and aecidia of the berberry are found
246
INTRODUCTIOX TO THE STUDY OF FUNGI
developed on Bcrhcris vul(/aris, Ijut no corresponding iiredo-
spores or teleutospores upon that plant, and therefore they
have to be sought elsewhere (Fig. 117).
On the other hand, the uredospores and
teleutospores of Puccinia graminis flourish
on wheat and other grasses, whilst no
spermogonia or aecidia have been known to
infest the latter plants. Hence it is con-
cluded that the above form the normal series,
with the spermogonia and aecidia on the
berberry, and the uredospores and teleuto-
spores on wheat, which completes the cycle.
This theory is supported by the contention
that the germinating spores of Aecidium
herheridis are capable of producing Puccinia
FiG.117.— AeLKiiospore graminis by artificial inoculation on wheat:
in germiuation. Alter - , , i , . ■,
Tuiasne. ^nd couvcrscly the promycehal spores of
Puccinia graminis (Fig 118) may be used
to inoculate the leaves of the berberry, and produce thereon
Aecidium herheridis. This is the theory and its application,
which we will here leave as it
stands. In another group, the
Braclnj - 'pueciniae, spermogonia,
uredospores, and teleutospores are
found on the same species of plant,
but the aecidia are unknown. To
this succeeds the Hcmi-fuecinia ,
in which uredospores and teleuto-
spores occur on the same plant,
but the spermogonia and aecidia
are unknown. It is here that the
sceptics w^ould place Puccinia
graminis were they not debarred p^^. ns— Puccinia teleutospore
by the anathemas of the votaries germiuating and producing pro-
of Heteroecism. Hereafter follows "^"''''^' '^°"'- ^"''" ^"'^^'^'•
the group Pucciniopsis, in which spermogonia, aecidia, and
teleutospores have been recognised, but not uredospores, so
that in this group of species the uredo stage is deficient.
Of the two remaining groups, Micro-pucciniae includes those
A' ^^-7' FUNGI— UREDI NEAR 247
species in which only teleutospores are known, and these do
not germinate for a long time after the foster-plant is dead ;
and the Lepto-pucciniae, in which also the species only possess
telentospores, but the sori are compact, and germination takes
place at once and whilst the foster-plant is living. Of course,
outside of all these groups there still remain a rather large
number of species, of uncertain place, mostly with only the
teleutospores definitely known, but which it is suspected will
ultimately find a place in some of the foregoing groups, when
their life-history has been ascertained. Other Didymosporous
genera are Uropyxis, which seems to be hardly generically
distinct, in which the teleutospore is involved in a thick per-
manent hyaline integument ; and Diorchidium, which differs
chiefly from Puccinia in the septum being vertical. Gymno-
sporangium is most distinct in the teleutospores being agglut-
inated together in, generally, large tremelloid masses, the
teleutospores themselves being transversely uniseptate, or very
rarely biseptate, nearly hyaline, with long, sometimes very
long, pedicels.
In Phragmosporae the teleutospores are three, or more,
septate, in one direction. In most genera
they are destitute of a pseudoperidium,
whilst in Phragmidium'^ (Fig. 119) and
Xenodochus the uredospores are solitary.
The differences between these two genera
are slight: in the former the teleutospores
are cylindrical, the cells not readily break-
ing up into joints ; in the latter the cells
are more numerous and moniliform, soon
breaking up into the component cells. In
two genera the uredospores are catenulate,
of which Coleosporium is the most import-
ant; in this genus the uredospores are
associated in chains, and the teleutospores ^i«- 1 19— Teleutospores
^ 01 Phragmulmm.
are three, or many, septate. The pro-
mycelium is continuous. In Cltrysomyxa the uredospores are as
in Coleosporium, the teleutospores are multiseptate, and simple or
1 Hamaspora longissima has been included under Phragmidium, but we doubt
if it should not be maintained as a distinct genus.
248 INTRODUCTION TO THE STUDY OF FUNGI
branched, whilst the proiuycelium is tiiseptate. In three small
genera the teleutospores are longitudinally septate ; that is, in
Pucciniastruvi, in which the uredospores are enclosed in a pseudo-
peridium, but the teleutospores are evolved externally to the
matrix ; Thecopsora, in which the uredospores are also enclosed
in a pseudoperidium, and tlie teleutospores are intracellular ; in
Calyptos2')ora there are aecidia but no iu"edospores, and teleuto-
spores, provided it is accurately determined that, in the single
species, the aecidium is found on the leaves of Abies, and the
teleutospores on the branches of Vaceinium. In the remaining
two genera the teleutospores are enclosed in a pseudoperidium —
in Endopliyllum resembling an Aecidium with catenulate spores;
and in Milesia the teleutospores are catenulate within a reticu-
late immersed pseudoperidium. These two genera are outsiders
and only remain here on sufferance.
TheDictyosporae, in which the teleutospores are transversely,
longitudinally, and obliquely septate, contain but two very
distinct but different genera. In Triphragmium the teleuto-
spores are triseptate, or radiately three-celled. In JRavenelia
the teleutospores are many-celled, the cells radiating or con-
centric, often with hyaline basal cells, in surface aspect re-
sembling the glomerules of Sorosporium.
In such an arrangement as the foregoing, wherein so much
depends upon a knowledge of the life-history of every species,
it is but natural to expect that there will be a number of forms
which present the earlier stages of a succession, and are yet
deficient of the requisite determinator, the teleutospore. These
have to be relegated to outside groups under the denomination
of Inferior Uredines, or Imperfect Uredines, and as such find
their places under the following genera. The species having
spermogonia only are ranged under Aecidiolum — the species of
Aecidium, which remain isolated, retain their position under
that old generic name ; the species which are analogous, but
have elongated pseudoperidia, and were formerly known as
Baestelia, still retain that name, producing aecidiospores, as also
does Peridermium, which is analogous to Aecidium, on conifers.
The remaining genus is Uredo, which includes aU unplaced
species of uredosporous Uredines, whether known formerly as
Uredo, or Trichohasis, Lecythea, or Caeoma, the latter having the
RUST FUNGI— UREDINEA E 249
spores catenulate. Here, then, we have the spermogonia, aecidia,
and the uredospores, which may possibly prove to form parts of
unknown cycles, the position, affinity, and association of which
must be left to the chances of future investigation.
During late years there has been no lack of investigation
and observation on the structure, growth, and development of
the Uredines, which have mostly taken the form of artificial
cultures. Whether the same results take place, and in precisely
the same manner, in a state of nature, cannot be affirmed, whilst
some present grave reasons for doubt. It is still possible that,
if the facts are accepted, the inductions may be wrong. When
it is argued that certain experiments succeed in producing upon:
certain plants the identical Uredines which would have been de-
veloped in the ordinary course of nature, a sceptic will naturally
inquire for the evidence which proves that the resulting Uredines
were really produced by inoculation, or whether the elements
were not already present, and that these were simply stimulated
by the introduction, or intervention, of other agencies, and hence
not actually produced by inoculation. Common sense would
allow that, if the resulting Uredine were foreign to the par-
ticular species of plant, the assumption of inoculation would be
more convincing. This is not the place to enter upon a dis-
cussion of the doubts and dangers which attach to inductions
from the results of artificial cultures, hence it must suffice to
suggest that such doubts and dangers may still continue to
exist in the minds of those who venture to hold independent
opinions.
Suggestions have not been wanting of sexuality, or of
fertilisation which implies sexuality, in the Uredines. The
application of the term " spermogonia " to the small pustular
eruptions, which occupy the first place in the cycle, can scarcely
be accepted as a suggestion, but some authors have assigned to
the minute spore-bodies not only the name but the function of
spermatia. Meyen was evidently of this opinion, and he was
not the only one who thought that they played the part of the
male element. Worthington Smith has intimated that he has
often seen the small sporules attached to the exterior of Aecidio-
spores, but he was unable to trace any pollinial tube into the
interior. Against the supposition that they are fecundative
2 50 INTRODUCTION TO THE STUDY OF FUNGI
bodies it may be urged that the globose form, and absence of
movement, are unusual with known spermatia. And again,
the fact of their power of germination, or rather of budding, is
opposed to their possessing the function of spermatia. Cornu
found that the so-called spermatia budded in the manner of
Saccharomyces. In saccharine solutions they behave in a very
similar manner to yeast spores, or analogous to the concatenate
production of cells in the Ustilagineae.
It has also been suggested that in the aecidium stage the
conjugation of two swollen hyphae of the mycelium takes place,
from which the whole aecidial cup is produced as the result of
a sexual act. This again requires careful confirmation before
it can be accepted as more than a hypothesis. In the subse-
quent stages of uredospore and teleutospore no sexual act has
been discovered, and at present we are l)ound to admit that in
the Uredineae sexuality has not been proven.
BIBLIOGRAPHY
De Tom, J. B. " Sylloge Urediuearum," in Saccardo, Syllocje Fungonnn, vol.
vii. pt. ii. Padua, 1888.
Plowri&ht, C. B. a Monograph of the British Uredineae aiul Ustilagineae. Svo.
London, 1889.
De Baby, A. Untcrsuehungen iiber die Brandpilzc. Svo. 1853.
Smith, W. G. Diseases of Field and Garden Crops. 12mo. Cuts. London, 1884.
Unger, F. Die Exantheme der Pflanzen. Svo. Vienna, 1833.
Cooke, M. C. Rust, Smut, Mildew, ami Mould. 12mo. Col. plates. London,
1870.
DiETEL, P. Beitrdge zur Morphologie %ind Biologie der Uredinccn. 1887.
ScHROTER, J. "Die Brand- und Rostpilze Schlesiens," in Ahhand. der Schles.
Gesell. 1869-72.
Winter, Geo., in Rabenhorst's Kryptogamen Flora — PUze. Svo. Cuts. 1884.
Leveill^, J. H. "Sur la disposition des Uredines." Ann. des Sci. Nat., 3rd
series, vol. viii. Paris, 1847.
Fablow, W. G. "The Gymnosporangia or Cedar Apples of the United States."
Mein. Boston Sac. Nat. Hist. Svo. Boston, 1880.
CHAPTEK XXI
SMUT FUNGI USTILAGINES
It was at one time so customary to associate the Ustilagines
with the Uredines that it would have been thought that some
close bond of union existed between them, instead of which
they have really no closer affinity than the fact of their being
alike parasites upon living plants. In the days when the
Coniomycetes were accepted as an order of Fungi, on the basis
that they produced spores, on more or less distinct sporophores,
with the threads, or hyphae, obsolete, or nearly so, then the
Ustilagines and Uredines were associated with the Sphae-
ropsideae as members of that order. It was then contended
that this division was distinguished " by the vast predominance
of the reproductive bodies over the rest of the plant, if not in
size, at least in abundance, and from the ease with which in
general they fall from the point of attachment, in consequence
of which, as the name implies, they have a dusty appearance,
and often soil the fingers of those who handle them."^ No
longer can so artificial an association be recognised, and whilst
the Sphaeropsideae hold lower rank as imperfect forms, the two
groups of Ustilagines and Uredines maintain independent
positions, as autonomous, within certain restrictions. Tulasne
contributed much to the better knowledge of the Ustilagines
in 1847, to which Fischer de Waldheim, with Brefeld and
others, have contributed since. How far these organisms
differ from the Uredines must be gathered from a comparison
of the present with our chapter on the latter group.
It must be premised that these are pustular Fungi, which
^ Introduction to Cnjptugamic Botany, l>y M. J. Berkeley, London, 1S57, p-
315.
2 52 INTRODUCTION TO THE STUDY OF FUNGI
attack growing plants, and produce copious soot-like spores, so
that they have acquired the general name of " Smuts." The
mycelium is deeply seated in the tissues, and the spores are
developed in definite positions on the host : on the stem,
leaves, flowers, ovaries, fruit, and sometimes in the corm, tuber,
or root, but seldom in more than one of these places, and that
one habitual to the species. A good example may be seen in
the smutted ears of corn, or the distorted receptacles of the
goat's-beard.
The mycelium is an important element in this family,
although so delicate and deeply seated that it is often passed
over. The whole substance of the host may be penetrated
and taken possession of by the mycelium before there is any
external evidence of its presence ; therefore when the pustules
are formed it is too late to apply remedial measures, for the
plant has long been doomed. It is always tedious and difficult
to trace the ramifications of mycelium in growing tissue, but
in some of these species it may be seen bearing haustoria or
suckers amongst the cells. Not only does the mycelium
traverse the intercellular spaces, but frequently the branches
pierce the walls of the cells, and though seen most readily in
young plants, it is always manifest about the spore-pustules.
The persistency of the mycelium is one of the agencies by
which the continuity of the species is preserved. When the
foster-plant dies in the winter the mycelium dies with it, but
when the root-stock is perennial the mycelium also remains,
to revive and penetrate the young shoots which are put forth
in the spring. All the leaves of the violet may die year after
year, and still every season Urocystis violae appears again, even
when every infested leaf has been picked and burnt.
The most important function of the mycelium in this
family is its concern in the formation of the fruit. At the
special spot where the development of fructification is to take
place the mycelium undergoes some change in its character : the
walls increase in thickness, and the contents become gelatinised.
Some slight modifications take place in the different genera,
but for the most part the hyphae branch and become en-
tangled so as to form compact knots, or spore -beds. With
this the hyphae gradually swell in places, and it is evident
SMUT FUNGI— USTILAGINES 253
that a change is taking place within. The swellings are
indication of spore - formation, which proceeds sometimes
centripetally, so that . those on the exterior are most com-
pletely developed, the circumference darkens, and an epispore
is formed. In Siihacdotheca part of the hyphae are concerned
in the production of the receptacle and columella, and part in
the origination of the spores. In some genera the production
of spores is centrifugal, and the peripheral spores are sterile.
In Sorosjoorium " the spore-forming hyphae from several con-
tiguous mycelial branches incline together and twist them-
selves into a ball, as happens in the formation of a Lichen
thallus. These convoluted and contorting spore-forming
hyphae, being gelatinous, soon become so entwined and en-
tangled that they cease to be individually recognisable ; to all
appearances they coalesce together in part, if not entirely, and
on the exterior of this gelatinous ball other hyphae are seen
encircling it. These latter also being gelatinous, soon lose
their individuality, although at times traces of their concentric
arrangement can be made out. Spore-formation takes place
only in the central gelatinous ball, in the middle of which it
commences by the central part darkening in colour and
becoming differentiated into spore-like bodies, which vary in
number from four to sixteen. Apparently these bodies again
subdivide, so that when the spores arrive at their maturity
the spore-balls contain sixty to a hundred or more spores. In
the young state the developing spores are polygonal from
mutual pressure ; subsequently the balls increase in size, and
the gelatinous zone swells also. When the spores assume
their dark brown colour the gelatinous zone begins to be ab-
sorbed, and entirely disappears when the spores are fully
matured. In a certain sense the spore formation is centri-
fugal, as it commences in the centre of the gelatinous ball, but
the peripheral spores are the oldest, having been pushed
outward by the formation of younger spores in the centre." ^
The spores of the Ustilagineae are practically teleuto-
spores, and are called such by some writers. They are com-
posed of two membranes — the outer, or exospore, the thicker
and dark coloured ; the inner, or endospore, thin and colourless.
^ British Uredineae aiul Ustilagineae, by C. B. Plowright, London, 1889, p. 64.
254 INTRODUCTION TO THE STUDY OF FUNGI
The surface of the epispore varies in its character, being some-
times (piite smooth, in other species reticulated, in others so
minutely granular as to appear to be smooth until closely
examined, or in others distinctly rough and either obtusely
warted or spinulose. In colour they seem to be black in a
mass, but viewed obliquely sometimes with a yellowish or
olive tinge ; seen under the microscope, by transmitted light
they may be black or brown, violet, olive, or yellowish, and
rarely hyaline. Where colour is present it resides in the
epispore, and is fairly constant in each species when mature.
The form is commonly globose, or approximately so, when
perfectly free, but being usually closely compressed in growth,
is apparently angular. The spore -masses in some genera,
such as Ustilago and Tilletia, are more loosely packed, and the
teleutospores do not adhere to each other in definite clusters,
but are normally free. In Sorosporium and Thecaphora they
form compact clusters, which in the latter genus separate with
difficulty, whilst in Cintractia, although at first agglomerated,
they soon separate. In Urocystis there are a few large, dark-
coloured fertile spores, closely adnate to each other in the
centre, and these are surrounded by hyaline sterile cells, or
pseudospores, which give the appearance of a beaded border.
In Entyloma, Melanotaenium, and Entorrhiza we meet with
aberrant genera, which remind us of Protomyces and Syncliy-
trium, and are probably more closely allied to the latter than
they are to the rest of the Fhycomycetes.
Germination of the teleutospores in this family has often
been observed and watched. In some species a small germ-
pore has been observed, but they are never so distinct as in
the UrecUneae. When the spores germinate they protrude a
germ-tube — usually designated, for reasons hereafter evident,
a promycelium. In a certain sense it may be regarded as
analogous to the protonema of mosses. This promycelium
bears small hyaline bodies, which resemble spores, and are called
by Continental mycologists sj^oridia, a name to which we take
exception as it should be restricted to spores generated in asci.
We will call them, for the time, promycelial spores, as suggested
by Plowright, and much more appropriate. The promycelial
spores will bud and produce secondary promycelial spores,
SM UT FUNGI— US TIL A GINES
'■SS
and these again may continue to multiply themselves many
times by budding, after the manner of yeast-spores, which is
the term applied to them by Brefeld, but liable to misinter-
pretation. In order the better to comprehend the process, it
may be detailed as observed in Tilletia (Fig. 120). This parasite
produces its teleutospores within the grains of wheat, and is
known to farmers as " bunt."
The appearance of the grains
externally is very little
changed, but slightly darkened
in colour, and when crushed
are seen to be filled with
a sooty, rather fetid powder.
These teleutospores are globose,
dark coloured, almost black,
and the surface minutely
reticulated. When placed in
water they germinate in about
forty-eight hours. A germ-
tube is emitted from a very
small germ-pore, but it does
not attain any considerable
length; and this germ-tube
constitutes the promycelium,
into which the contents of the
parent spore pass and retreat
to the extremity, and are
of a transverse septum,
about the summit, and
first promycelial spores,
and colourless, to the
Fig. 120. — Tilletia s^ov^a in germination,
ft, producing a promycelium, ^j ; b,
primary spore with promycelium, 2>i
bearing conidia, of which some are
conjugating ; c, conjugated gonidia in
germination, with secondary gonidium
at s'. After De Barv.
are shut off by the formation
Tuberculations are soon manifest
these by lengthening become the
They are thread - like, curved,
number of from four to a dozen.
When fully developed they are cut off from the pro-
mycelium by a septum at the base. Soon afterwards these
primary spores will be seen to connect themselves, mostly in
pairs, by a transverse connective, performing an act of conjuga-
tion. These conjugated primary spores are often separated
from the promycelium, but they may remain for a long time
attached. In due time liudding takes place, and the buds
become converted into cylindrical curved secondary spores, the
256 INTRODUCTION TO THE STUDY OF FUNGI
second generation of proniycelial spores, which in like manner
are cut off by a septum at the base, and become free inde-
pendent bodies. The act of conjugation, which results in their
production, is not an essential, because solitary primary spores
are equally capable of budding and producing secondary spores,
although tliey are usually smaller than those produced by con-
jugating primary spores. Hence it may be concluded that the
conjugation of the linear spores is not a sexual act. The
secondary spores are usually those which, by germination, enter
the host-plant and form a mycelium, but they are also capable
of budding and forming proniycelial spores of a third, or even
of a fourth, generation, if the conditions are unfiivourable for
infecting a new host-plant.^
The germination in Entyloma is similar to the above, but
less complex. The spores send out a germ -tube in about
twenty-four hours, and this constitutes the promycelium, which
develops several branches at the apex, each of which is cut
off by a septum at the base and becomes a proniycelial spore.
These spores then conjugate in pairs by the formation of a
connective bridge ; afterwards, by a continuation of growth at
the apex, secondary spores are produced, which fall off and
germinate. The growing point enters the host -plant and
forms a mycelium, which starts a new infection, and in com'se
of time teleutospores of a normal kind are developed in
clusters. In this genus conidia are also produced direct from
the mycelium, the conidia-bearers rising to the surface of the
leaf through the stomata. These conidia germinate on the
surface of the leaf upon which they fall, and the germ-tube
enters the stomata and forms a mycelium. They are able to
form secondary conidia, but this seldom takes place under
normal conditions. Hence there are two forms of reproduction
in this genus — that of the germinating teleutospores, forming a
promycelium which gives rise to proniycelial spores, and these
after conjugation developing secondary spores, capable of repro-
ducing the parasite, after an alternation of generations ; and,
secondly, of germinating conidia, which reproduce the parent
Entyloma at once, without an intervening generation.
1 "On Bunt Spores," by M. C. Cooke, Journal of QuebcU Microscojncal Club,
vol. i. p. 167, 1868.
SMUT FUNGI— USriLAGINES 257
The teleutospores in some instances in this family are
resting-spores — that is to say, they are capable of germination
after a period of rest ; but for the most part they germinate
freely when moist, and a delay of germination can only be
secured by maintaining a condition of dryness which does not
obtain in a state of nature. It is uncertain how the interval
is connected between the matm-ity of the teleutospores and the
growth of the seedling hosts, where the entire plant is annual.
In the case of perennial hosts a persistent mycelium removes
all difiiculty, but where mature teleutospores are produced upon
an annual in summer or autumn, and there are no seedlings
until two or three months afterwards, it is not evident how
the continuity of the species is preserved.
It has been shown that, when cultivated in a suitable
medium, the promycelial spores multiply themselves almost
indefinitely by budding, but the nutrient fluid must be
maintained unexhausted. In this condition the growth is
similar to that of yeast, and the term " yeast cells " has been
applied to them. So long as the supply of nutrient fluid is
maintained there is no departure from the budding process.
Brefeld maintains it to be extremely probable that the conidial
fructification, in a toruloid form, occurs in nature in many
species of the Ustilagineae ; that they have the power of pro-
pagating outside the host as " torulae," and develop their spore
fructification only when they penetrate the tissues of the host-
plant by means of germ threads, which takes place when the
supply of nutriment ceases. The extreme assumption on this
basis is that certain forms of Saccharomyces, or indistinguish-
able therefrom, are in fact aquatic forms of the conidia of
Ustilagines, which have become " toruloid " on account of their
surrounding conditions. In fact, that some ferments are not
autonomous, but depraved Ustilagines which have abandoned
their parasitic habit and become saprophytes.
It will therefore be evident by this time that the members
of this family are universally parasitic, and that the hosts are
herbaceous plants. In a great number of instances the
graminaceous plants are the victims, but by no means ex-
clusively. In all cases they are eminently destructive, and,
from their habit, difficult to contend with. Most of the pro-
17
258 INTRODUCTION TO THE STUDY OF FUNGI
posed remedies are preventive and not curative ; they could
hardly be otherwise until that period in their life-history is
more definitely determined which intervenes between the
maturity of the teleutospores and the inoculation of seedlings
of the host-plant.
Assuming the total of known species to be somewhere
about three hundred, upwards of one hundred of these affect
the grasses, and nearly fifty attack other Monocotyledonous
plants, so that scarcely half of the total are found on Dicoty-
ledons. This is a peculiar fact in distribution which is
perhaps without parallel in any other family of the parasitic
Funsi.
BIBLIOGRAPHY
De Toni, J. B. " Sylloge Ustilagiuearum et Uredinearum," in Saccardo, Syllogc
Fungorum. Imp. 8vo. Vol, vii. pt. ii. Padua, 1888.
Fischer de Waldheim. Sur la Structure des Sjwres des Ustilaginics. Moscow,
1867.
Apcrgu systimatique des Ustilaginies. 1877.
"Les Ustilaginees et leiir Plantes nourricieres." Ann. des Sci. Nat.,
6th series, vol. iv. Paris, 1877.
"Winter, Geo., in Rabenhorst's Kryptogamen Flora — Dcv Pilzc. 8vo. Guts.
1884.
WORONIN, M. Bcitrag zur Kenntniss der Ustilaginecn. 1882.
Massee, Geo. British Fu7igi — Phycomycetes and Ustilagineae. 8vo. Plates.
London, 1891.
TuLASNE. " Memoire sur les Ustilaginees comparees aux Uredinees." Ann.
des Sci. Nat., 3rd series, vol. vii. Paris, 1847.
"Second Memoire." Ann. des Sci. Nat., 4tli series, vol. ii. Paris.
ScHROTER, J. "Bemerkungen und Beobachtungen ii. einige Ustilaginecn. "
Cohn, Beitrage z. Biol., ii.
Brefeld, 0," Bot. U7itersuch. ii. Refenpilze. Leipzig, 1883.
CoRNU, M. "Contributions t\ I'^^tude des Ustilaginees." Bullet. Sac. Bot. de
France. August 1883.
CHAPTEE XXII
IMPERFECT CAPSULAR FUNGI SPHAEROPSIDEAE
In the old arrangement by Fries, one of the primary divisions
of Fungi was that termed Coniomycetes, which was interpreted
as " Dust-fungi," and was represented as including those fungi
in which the spores were the principal feature, such spores
appearing like an impalpable dust. It was, perhaps, an odd
mixture, but this group included not only the Si^haero^psideae,
as at present limited, and the Melanconieae, which are closely
related, but also the Urediiieae and the Ustilagineae, which are
not related at all, and are now separated and rank as a distinct
group. It need not be explained here wherefore the Uredineae
and its allies were entirely out of place in an association with
Fungi which either possessed a distinct perithecium, in which
the spores were generated, or a pseudoperithecium formed from
the matrix.
The group now under consideration is analogous, in ex-
ternal features, to the Pyrenomycetes, but wholly deficient of
asci. The perithecia, or pseudoperithecia, include only stylo-
spores, and have been assumed to be imperfect representatives,
or imperfect stages or conditions, of the Pyrenomyceteae, and
hence called " imperfect capsular Fungi." In some instances
this may be undoubtedly true, but we think it assuming
too much to affirm that all are imperfect conditions of higher
Fungi, because it has been demonstrated to be the case in a
comparatively few instances. It is at least premature to
decline acknowledgment of thousands of very distinct forms of
Fungi in a systematic position, simply because a few of them
have been shown to be transitional, whilst the majority may
never be demonstrated to be other than autonomous. This
26o INTRODUCTION TO THE STUD V OF FUNGI
objection luis been appreciated by Saccardo, who includes all
the species in his Sylloge, although he relegates them to an
inferior position as " imperfect fungi."
The Sj)haero23sideae must be considered apart from the
Melanconieae, on the fundamental basis that the former possess
a distinct perithecium, and the latter are only circumscribed
by a modification of the matrix. With this limitation, there-
fore, the Sphaeropsideae correspond to the Pyrenomyceteae,
although dissevered by the absence of asci and paraphyses. It
would be well if authors in future would respect Saccardo's
definitions of the fruit in the different orders, by a restriction
of the terms. Thus in the Ascomycetes, where the representa-
tives of seeds, or the spores, as they are generally termed, are
produced within asci, that they are sporidia. When produced
naked on basidia, as in the Basidiomycetes or Myxomycetes,
they should retain the name of spore. When enclosed in
perithecia, but without asci, as in the Sphaeropsideae, then to
be termed sporules. But when wholly naked, and without
basidia, or receptacle, as in the Hyphomycetes, then to be termed
co7iidia. The only modification to this arrangement which
approves itself to us is the application of the term sporules to
those bodies which are enclosed in a pseudo-perithecium, such
as the Melanconieae, as well as those contained within a definite
perithecium ; and the restriction of conidia to absolutely
naked fruit, in which there is neither perithecium nor semblance
of a perithecium, as in the Hyphoinycetes. Practically this
means the association of the Melanconieae with the Spihaerop-
sideae, in the denomination of sporules, instead of union with
the Hyphomycetes under the denomination of conidia. This
may be a distinction of little importance, but it is one which
appears to commend itself to consideration.
The Sjjhaeropsidcae, therefore, may be thus defined, as Fungi
possessed of a perithecium, but without asci, the sporules, or stylo-
spores, being produced internally at the apex of more or less
distinct supporting hyphae or pedicels, which, for the sake of
distinction, should not be termed basidia, but sporophores.
This would obviate any confusion with the spore-bearers of the
Basidiomycetes, and the definition would be reduced to " peri-
thecia, without asci, enclosing sporules, on more or less distinct
IMPERFECT CAPSULAR FUNGI— SPHAEROPSIDEAE 261
sporophores." The primary families would depend for their
distinctive characters upon the nature of the perithecium.
The first and largest is the Sphaerioideae, in which the
perithecia are membranaceous, coriaceous, or subcarbonaceous,
typically subglobose, and closed ; thus analogous to the old
genus Sphaeria. The second family, Nectrioideae, with the
perithecia similar in form, but fleshy or waxy, and usually
brightly coloured, analogous to the old genus Nedria, or the
more recent family Hypocreaceae. Then the third family, the
Leptostromaceae, has the perithecium more or less dimidiate,
and astomous, or with a longitudinal fissure, and black, corre-
sponding in some respects to Hysteriaceae. Finally, the fourth
family is JExcipulaceae, with the perithecium cup -shaped, or
patellate, at first spherical, then broadly open, and making the
nearest approach to analogy with the Discomycetes. Each of
these families we must therefore analyse a little more in detail,
bearing in mind their distinctive family features.
The SjjJiaerioideae are therefore the Sphaeriaceous, or
Sphaeria-like, SjjJiaerojysideae, with blackish closed perithecia ;
and although we should have preferred grouping them in a
similar manner to our subfamilies of the Sphaeriaceae, we will
rest content with the arrangement proposed in the Sylloge,
which will be the one generally adopted for some time to come.
Of course this method is an artificial one, to a great extent,
being based upon the character of the sporules. The Hyalo-
sporae is again the largest section, including all the genera
with continuous hyaline sporules ; those in which the peri-
thecia are simple or distinct forming one subsection, and
those in which the perithecia are composite or caespitose
forming another. Amongst the simple species the larger
number have the perithecia naked or smooth, and of these one
genus, Fhyllostida, is often parasitic, growing upon leaves, the
depressed and innate perithecia being grouped on discoloured
spots ; the remaining genera have the species not seated on
definite spots, and of these three are very similar to each other ;
that is to say, Phoma, with the perithecia (Fig. 121) covered
by the cuticle ; Aposphaeria, with the perithecia exposed, or
superficial, mostly on dead wood ; and Dendrophoma, which in all
things else resemble PJioma except that the sporophores are
262 INTRODUCTION TO THE STUDY OF FUNGI
branched, instead of remaining simple. More recently the
genus Phoma has been subjected to another mechanical sub-
division into Phoma and Macrophoma, the latter to include all
the species of Phoma which have sporules exceeding a definite
size, so that the determination of the genus may
9 depend upon the difference of a micromillemetre.
^ <^ Another genus, Asteromdla, which is equal to
Asteroma, minus a subiculum, has the minute
perithecia clustered on dendritic spots. There are
five or six smaller genera, consisting of but a
Pig 121 Peri-
thecium of few spccies, whicli Completes the series of genera
Phoma with j^ which the perithecia are not rostrate. In
Sphaeronema the habit is that of Phoma or
Aposphaeria, but the perithecia are rostrate. In addition to
these follows a series of genera in which the perithecia are
seated upon a subiculum of some kind, more or less distinct
and definite. In Chaetophoina the perithecia resemble those of
Phoma, but are innate in a dematioid subiculum resembling
Fumago or Asterina. An allied genus, Asteroma, is the ana-
logue of such genera as Asterina or Dimerosporiitm, the minute
perithecia being seated upon, or amongst, a subiculmn of radiat-
ing black fibrils. In the remaining two genera, Ypsilonia and
Cicinnobolus, each contains but a single species, and the latter
is parasitic upon Oidiuvi. In the three genera which com-
plete those in which the perithecium is bare, Neottiospora has
the sporules cristate, and the other two genera have the
sporules in chains. We pass now to the smaller series, in
which the perithecia are hairy or bristly. Here are four
genera, the most numerous and important of which is
Vermicularia. In habit the perithecia resemble those of
Venturia, or some species of Chaetomium, the long dark
bristles are septate, and the sporules mostly curved. Pyreno-
chaeta is similar, but the hairs of the perithecium are shorter,
and the sporules ovoid or oblong. Muricularia and Staitrochaeta
differ from the foregoing in the character of the external hairs.
This brief review of the simple species leads us to the series
in which the perithecia are composite or caespitose, usually
with a definite stroma. Dothiorella resembles superficially
either Botryosphaeria or Cucurhitaria. The pustules are
IMPERFECT CAPSULAR FUNGI— SPHAEROPSIDEAE 263
erurapent, and consist of a number of perithecia aggregated
upon a basal stroma. Rabenhorstia, and especially Fuchelia,
have a subglobose stroma, in which are fertile cells, so that in
habit and structure there is a similarity to some species of the
ascigerous genus Fuckdia of the Melogrammeae. Flacosphaeria,
on the other hand, has an effused stroma resembling Rhytisma,
or certain species of Phyllaclwra. The four remaining genera
possess a stroma, more or less like Vaha, especially so in
Cytospora, in which the sporules are small and sausage-shaped.
It is probable that all the species in this genus are stylospor-
ous conditions of Valsa. The genus Cytosporella only differs
from Cytospora in having the sporules either globose or ovoid ;
and Fusicoccum again in the sporules being large and straight,
and mostly fusiform ; so that these three genera resemble Valsa
in habit, but differ amongst themselves in the form of the
sporules. Centhospora might be included in the same series,
but the stroma is firmer, and the spore-bearing cells have the
converging necks mostly united in a central orifice. The
sporules are cylindrical and typically straight. It is very
usual, even if not universal, for the mature sporules which are
held together in a pasty mass to be ejected in the form of a
tendril, or a contorted thread. Thus concludes the section
Hyalosporae of tlie Sphaerioid family of the Spthaeropsideae,
including an enumeration of not less than 2625 species.
The Phaeosporae section has the sporules continuous, and
coloured either brown or sooty. The number of genera is
comparatively small, and of these four have the perithecia
subglobose and smooth. Two of these are practically old
genera, since Sphaeropsis was recognised by Fries, although
not limited, as now, to species with coloured sporules. It is
in all respects the corresponding genus to Diplodia, but with
continuous, and not uniseptate, sporules, as in that instance.
Coniothyrium is closely allied to Sphaeropsis, but the perithecia
are normally smaller, as are also the sporules, which are large
and elliptical in Sphaeropsis, small and globose, or ovoid, in
Coniothyrium. The genus Harhnessia closely resembles Melan-
conium in the sporules, and also in their ejection when
mature, and the consequent blackening of the matrix, but
differs in the possession of a distinct perithecium. In
264 INTRODUCTION TO THE STUDY OF FUNGI
Hypocenia the sporules are fusoid and pale browu. Levieuxia
is a South African genus, containing a single species, with a
stipitate perithecium, which is fissured at the apex when
mature. The only genus with hairy perithecia is Chaetoinella,
which is the analogue of Chaetomium, but deficient in asci.
The residue of the section consists of genera in which the
perithecia are compound or caespitose. Haplosporella is, in
fact, a caespitose Sjjhaeropsis, the perithecia being aggregated
in dense erumpent pustules, resembling those of some species
of JBotryosphaeria. Weinmannodora has a stroma which is
hemispherical and carbonaceous, with radiating fertile cells,
containing globose dark sporules. Cyto2)lea consists of a single
species, in which the stroma is at first pulvinate, then con-
fluent and effused. Practically the entire section is repre-
sented in Europe by Sphaeropsis and Coniothyrium, with
smooth perithecia ; and Chaetomella, with setose perithecia ;
and in the compound genera by Haplosporella.
The Phaeodidymae is also a small section, with uniseptate
coloured sporules, and is, in fact, entirely made up of species
which entered into the old genus Diplodia, as recognised by
Fries. Thus Diplodia, as limited, contains species with a
smooth perithecium, and coloured sporules, not having a mucous
envelope ; Macrodiplodia, with similar smooth perithecia, but
coloured sporules having a mucous envelope, as in the sporidia
of Massaria ; Chaetodiplodia, with hairy perithecia, and
sporules as in Diplodia. The remaining genus with simple
perithecia is Diplodiella, in which the perithecia are almost
superficial, and flourish on decaying wood. The single com-
posite genus is Botryodipilodia, with the perithecia densely
aggregated in erumpent pustules as in Haplosporella.
The section Hyalodidymae is characterised by hyaline
uniseptate sporules. Two genera correspond to Phyllosticia in
Hycdosporae, in that the species are mostly parasitic on living
leaves, collected on discoloured spots. In AscocJiyta the sporules
are simply uniseptate, but in Bohillarda the sporules are uni-
septate and crested at the apex with long setae, resembling in
this respect the genus Pestalozzia. In three other genera the
smooth perithecia are scattered and not seated on discoloured
spots. Actinonemu has the perithecia seated upon a radiating
IMPERFECT CAPSULAR FUNGI— SPHAEROPSIDEAE 265
adnate subiculum as in Asteroma, l)iit with dili'erent sporules.
Darluca has no subiculum, but the perithecia are parasitic on
old Uredines, and in Di])lodina the species grow on branches
the perithecia, and even the sporules, resembling Diplodia, save
that they are uncoloured. In Cystotricha the perithecia
dehisce as in Hysterium, with a gaping fissure, and the sporo-
phores are septate and constricted, so as to possess a moniliform
appearance. In the only remaining genus, EhyncJioj^homa ,
the perithecia resemble Fhoma externally, except that they
are rostellate and the sporules are uniseptate.
The succeeding section, Fhragmosporae, has the sporules
multiseptate, and is represented in two divisions, in one of
which the sporules are brown, and in the other hyaline. The
former is the most numerous in genera and species. The old
genus Hendersonia, as interpreted by Berkeley, has been
divided, and is now restricted to such species as possess
coloured sporules. The perithecia are papillate, covered by
the cuticle. In Couturea the species have superficial perithecia,
which are seated on a stellate subiculum, somewhat after the
manner of Asteroma. In the two small genera Angwpovia
and Lichenopsis the perithecia dehisce by an operculum at the
apex. In the former the perithecia are superficial and hairy ;
in the latter immersed and smooth. Cryptostictis somewhat
resembles Pestcdozzia in the septate sporules being furnished
at both extremities with a hyaline bristle, whereas in Pesta-
lozzia the cilia are more than one, and confined to the apex of
the sporule. In another small genus, that of Prosthemium,
the sporules are also peculiar, in being joined together at the
base in a stellate manner (Fig. 122). The compound species
are confined to a single genus, in which the perithecia are
immersed in a stroma, as in Dothideaceac. This genus is
Hendersonula, and is, in fact, a compound Hendersonia.
The Hyalophragmiae, in which the sporules are colourless,
includes but two genera : Stagonospora, which is practically
Hendersonia with hyaline sporules ; and Mastomyces, in which
the perithecia are elongated and superficial, resembling scattered
perithecia of the rather obscure genus Corynelia, which is
ascigerous.
The Dictyosporae, in which the sporules are coloured and
266 INTRODUCTION TO THE STUDY OF FUNGI
muriform, consist almost absolutely of one type, that of
Camarosporium, which resembles Hendersonia in habit and
appearance, growing npon branches, covered by the cuticle.
Cijtosporium only differs in the perithecia being subsuper-
ficial, growing on naked wood. Dichomera, in which the
perithecia are immersed in a stroma, as in Dothideaceae, is
consequently compound. The doubtful genus Endohotrya,
contains but one species, which is North American.
A rather important section is the Scolecosporae, in which
the sporules are very much elongated,
so as to be thread-like, or rod-like, and
either hyaline or faintly coloured. The
principal genus is Septoria, of which
the species are in greater part parasitic,
growing on living leaves or the green
parts of plants. The minute perithecia
are flattened and innate, and typically
Fig. 122.— Prosfhemium . i j- ^ i •
section with sporules. aggregated upon discoloured spots.
This genus is analogous to Pliyllosticta,
from which the species cannot be distinguished except
by the sporules. There is a suggestion of genetic con-
nection between some of the species and the ascigerous
genus Sphaerella, but this has not been demonstrated.
Fhlaeospora includes such species as would otherwise find a
place in Septoria, were not the sporules thickened, and com-
paratively shorter. Hhahdosjjora scarcely differs from Sep)toria
except that the perithecia are not seated on discoloured spots,
and are confined to twigs and the stems of herbaceous plants.
It bears about the same relation to Septoria that Phoma does
to Pliyllosticta. Phlyctaena would otherwise be the same as
Ehahdospora, only that the perithecia split with a fissure, and
become deficient above. In the small genus Gelatinosporium,
the perithecia dehisce broadly and irregularly, the sporules in
the interior forming a gelatinous mass. There are two genera
in which the perithecia are distinctly rostrate as in Sj^haero-
nema, from which genus the species have been separated, on
account of the difference in the sporules : Sphaerograpliium, in
which the sporules are continuous ; and Cornularia, in which
they are septate. Of the three compound genera, Uriosjjora
IMPERFECT CAPSULAR FUNGI— SPHAEROPSIDEAE 267
has the stroma small and depressed, with the sporules con-
nected in bundles of four ; Dilopliospora has a crustaceous
stroma, with the sporules crested with cilia at each end;
whilst Cytosporina accords with Cytospora in general features,
but the sporules are thread-like and curved, or flexuous.
Possibly few of the species are autonomous. There remain
but two aberrant genera to be alluded to, and these are Micula
and Micropera. The species occur on bark and often resemble
lenticels, or are clustered like species of Cenangium, of which
they are said to be the pycnidia ; the sporules are elongated
and nucleate.
The second family, Nectrioideae, bears the same relation to
the Sphaerioideae as the Hypocreaceae to Spliaeriaceae in the
Pyrenomyccteae. The perithecia and the stroma, when present,
are fleshy or waxy, and pale or bright coloured. The arrange-
ment here is the same as in the preceding family, primarily
based on the sporules, so that the sections correspond. The first
section is the Hyalosporae, in which the sporules are continuous
and hyaline. Most of the genera are simple, and only one is
composite. Of the four in which the perithecia are not beaked,
only one needs particular reference, as the residue contain only
a single species. Zythia resembles a scattered Nectria in
appearance, with ovoid or oblong sporules. Sjyhaeronemella is
analogous to Sphaeronema, and has the perithecia rostrate.
The only composite genus is Aschersonia, of which the species
might be mistaken at first for species of Hypocrea, the
structure and habit being that of Hypocrea without asci.
In the only species of the genus Dichlaena the perithecia
have a double tunic, and the sporules are minute and globose.
In Didymosporae the only genus is Fseudodiplodia, which
corresponds to Diplodia in the brown uniseptate sporules, but
differs in the colour and texture of the perithecia. In
Phragmosporae all are hyaline. Stagonopsis corresponds to
Stagonospora, with like sporules ; but in Chiastospiora the
sporules are arranged in four rays. Only one species is known
in each genus. The Scolecosporae has but one genus, with fili-
form sporules. This is Polystigmina, a stylosporous form of
the ascigerous genus Polystigma, in which the perithecia are
immersed in a discoid stroma. A small group, consisting of
268 INTRODUCTION TO THE STUDY OF FUNGI
three genera and three species, has been placed in proximity
to the Nectrioideae, although not quite conforming thereto, on
account of their approach to a cup-shaped receptacle, so that
their position can only be regarded as provisional. Hysteromyxa
combines the habit of a Hysterium with the fruit of a
Myxomycete. Patellina with the habit of a Patellaria has
stylosporous fruit. And Cyphina has the appearance of an
Uxcipula, but is bright coloured and garnished with white
hairs, so as to offer an analogy to Volutella. All of these
require fuller investigation, and none are European.
The family Leptostromaceae diverges from the previous
families in losing much of the Sphaeria-like habit, and
approaching that of some of the Hysteriaceae, with occasional
suggestions of Fhacidiaceae. The perithecia are more or less
distinctly dimidiate, or scutiform, with or without an ostiolum,
or fissured longitudinally, either membranaceous or carbonace-
ous, black, and either erumpent or superficial. Under the
carpological arrangement, the Hyalosporae are again the most
numerous, and resolve themselves into two subdivisions, in one
of which, although the perithecia are destitute of a definite
mouth, they dehisce in diverse ways, but not with a longitu-
dinal fissure. In the other subdivision the perithecia split
longitudinally after the manner of the Hysteriaceae. Lep)totliy-
rium is the chief genus in the first subdivision,
with a dimidiate and shield-like perithecium,
which does not split by a longitudinal fissure, but
soon cracks all round and falls away (Fig. 123).
Some of the species are believed to be the stylo-
,^^=z=^ ^^^ spores of certain species of Coccomyces. The genus
^^"^ -=^ Piggotia has an irregular depressed perithecium
Fig. i2Z.—Lep- -whieh does not fall away. Ifelasmia is allied,
tothynuin. . . ....
but in this genus the perithecia are innate in a
black effused stroma, growing on fading leaves. The species
are the stylosporous condition of Phytisma, which develop
asci in the same stroma after the leaves have fallen and
rested on the ground during winter. Actinothecium has
an orbicular scutate perithecium, which dehisces by several
radiating fissures. In the other group, in which the perithecia
split longitudinally — the chief is Lep)tostroma, which corre-
IMPERFECT CAPSULAR FUNGI— SPHAEROPSIDEAE 269
spends to Lej)totliyri%Lm — the perithecia are elongated and
fissured after the manner of Hysterium, and some of them
are probably the stylosporous condition of species of Hypoderma
or of Zophodermium. Lahrella has nearly circular perithecia,
which are innate, and sometimes formed from the changed
matrix, dehiscing by a longitudinal crack. A final genus,
Sacidium, differs from all the rest in the perithecia not being
distinctly parenchymatous. The sporules are often globose.
The Fhaeosporae, with coloured continuous sporules, consist of
but one small genus, with scutate perithecia pierced in the
centre. Of this single genus, Firostoma, only one species is
known. The Fhragmosporae have all hyaline sporules, which
in Discosia are fusoid and ciliate at each end. In Entomo-
sporium the sporules are two-celled, with a lateral smaller cell
on each side at the septum, so as to be cruciate, each with a
cilium. This is the same genus as is sometimes known under
the name of Morthiera. The Scolecosporae, with long filiform
sporules, include the genus Acti^iothyrium, which has flat
shield-like perithecia, delicately fringed all around the margin ;
and Melop)hia, in which the similar perithecia are corrugated,
but not fringed at the margin. Leptostromella consists of
species of Leptostroma, as originally interpreted, which have
long filiform sporules. In other respects with the habit and
appearance of Leptostroma. The Leptostromaceae are most
common on leaves, or the stems of herbaceous plants, but with
only a few exceptions appear to be saprophytic.
The last family is the Excipulaceae, in which there is a
nearer approach to cupulate forms. In its general character
the perithecium, or receptacle, is cup-shaped, patellate, discoid,
or hysteriform, in all of which it is at first nearly spherical,
but soon open ; and either smooth or hairy, commonly erumpent,
and then superficial, so as almost to resemble minute black
Fezizae, with which some of the species were formerly associated.
The Hycdosporae are again the largest section, and may be
divided into those which have the receptacle smooth and those
in which the receptable is hairy or bristly. The smooth cupped
are of two kinds, namely, those in which the receptacle is cup-
shaped and those in which the receptacle is split longitudinally,
or is valvate. The cup-shaped, smooth-surfaced group contains
270 INTRODUCTION TO THE STUDY OF FUNGI
Godroniella, a genus in which the receptacle is composed of
agglutinated hyphae, in other respects scarcely differing from
the next genus, Excipula, in which the excipulum is cellular,
membranaceous, or tough, and black. The genus Excipula of
Fries, and many subsequent authors, was rather a heterogeneous
one, even including some ascigerous species, and the residue
are now distributed over six different genera. HeteQ'ojmtcIla
differs in the perithecia being thicker and more leathery, with
the mouth always contracted and torn, the sporules fusiform,
growing upon branched sporophores. In Bothichiza the black
receptacles are mostly erumpent, often gregarious, at first closed,
then rather cup-shaped, and are often the spermogonia of species
of Cenangium. Sporules oblong and continuous. In Lemalis
the receptacles are membranaceous, or rather fleshy, coloured,
but not black. Gatinula has the receptacles tough or horny,
and black, rather cup -shaped, disc often bright coloured.
Discula corresponds to Discella, but the sporules are continuous.
The receptacles are discoid or patellate, often imperfect, black or
coloured. Hereafter follow the genera in which the perithecia
are hysteriform or valvate. In Sporonema the receptacles are
valvate, dehiscing with angular teeth, as in Phacidium, of
which they are possibly a stylosporous condition. Fleococcum
is scarcely distinct from Sporonema, although the contents are
assumed to be more mucilaginous. Psilospora closely resembles
Dichaena, of which it is evidently a stylosporous state. The
perithecia dehisce in the manner of Hysterium, with two
lips, and occur upon living bark of trees. The remaining
genera of the Hyalosporae possess hairy perithecia. The genus
Amerosporium has the receptacles cup-shaped, and corresponds
to Excipula ; but the cups are setulose, the sporules are naked
at the ends. In Dinemasporium the habit and external appear-
ance are the same, but the sporules have a hyaline bristle, or
awn, at each end (Fig. 124). Polynema differs in having the
apex of the sporules crowned with about four awns. In
the next section, the Hyalodidymae, there are but two genera.
In Discella the perithecia are normally discoid, sometimes
imperfect, or formed from the matrix, a long time covered by
the cuticle. Sporules oblong, uniseptate, and hyaline. In
Pseudopatella the receptacle is cup-shaped, almost superficial
IMPERFECT CAPSULAR FUNGI— SPHAEROPSIDEAE 271
tough and black, with a pallid disc. Possibly a stylosporous
form of Diirella cotiqjressa is the only recorded species. The
section Phragmosporae contains two genera which are offsets
from the old genus Excipula. These are Exciindina, with
smooth receptacles and multiseptate sporules ;
and Excipularia, with setose receptacles and multi-
septate sporules. In the remaining genus, Pili-
dirnn, the receptacles are erumpent, discoid, and
membranaceous, blackish and torn at the margin,
with a pallid disc. The section Scolecosporae
alone remains, containing three genera in which
no definite stroma is present : Schizothyrella, with
the perithecia hemispherical, then torn at the
margin, the filiform sporules breaking up into Fig.
cylindrical joints ; Protostegia, in which the
receptacles are discoid, at first covered, then
exposed, margin torn or fringed, disc gelatinous, sporules
thread-like, but not breaking up ; and Oncospora, with
discoid or cup-shaped receptacles, usually gregarious, erumpent,
disc gelatinous and the sporules clavate, hamate (curved like a
sickle) or sigmoid (like the letter S). The only compound
genus is Ephelis, in which an effused, sclerotium-like l)lack
stroma bears the cup-shaped receptacles, with filiform sporules.
It is analogous to a genus of Discomyceteae, to which the name
of Ephelina has been given.
This running commentary on the Sp)haerop)sideae has made
it manifest that at many points the genera, or some species,
have a close relationship with some of the Ascomyceteae ; but even
if all these were removed, there would still remain an imposing
array of species against the autonomy of which no word of
calumny has yet been offered. The Sphaeropsideae may not be
so attractive or interesting as the Pyrenomyceteae, but they
equally claim recognition, until the alleged dimorphism can be
proved against them.
The Melanconieae have undou].>tedly a close afSnity with
the SjjJiaeropsideae, with which they have always been associated.
The chief distinction, and the only one which can be insisted
upon, is that the perithecia — so universal, under some form, in
the Sphaeropsideae — are absent in the Melanconieae. The habit
272 INTRODUCTION TO THE STUDY OF FUNGI
is nearly the same, except that the pustules are always erum-
pent, and never supertieial, and the sporules are similar in form
and size, produced similarly at the apex of short sporophores.
The difference therefore is reduced to that of the character of
the walls of the cavities in which the sporules are engendered.
In the Melanconieae there are definite cavities, beneath the
cuticle, which correspond in function to immersed perithecia ;
yet these cells have no heterogeneous walls, but are simply
modifications of the matrix. In many cases they are distinctly
modified so as to appear as pseudoperithecia ; in some there is
merely a compact base, formed by the mycelium into a spore-
bed, upon which the sporules are developed, and when mature
are ejected, in a more or less gelatinous mass, through fissures
or orifices in the covering cuticle. The spore-bodies are termed
conidia by Saccardo, as they are in the Hyphomyceteae, but we
prefer to employ the same term as that adopted in the allied
Sphaeropsideae, and distinguish them as spondes.
The technical definition of the Melanconieae is to the effect
that they are Fungi without perithecia or asci, forming sub-
cuticular pustules, which are partially erumpent, discharging
the sporules through openings in the cuticle, such sporules
being produced on a proligerous stratum, growing upon distinct
or obsolete sporophores, and either in themselves continuous
or septate, either hyaline or coloured.
The arrangement adopted is similar to that of the Sphaerop-
sideae, the primary sections having relation to the character
of the sporules. The Hyalosporae include those which have
oblong, or shortly cylindrical continuous hyaline sporules,
whether solitary on the sporophores or produced in chains.
Four genera are indicated in which the sporules are solitary
on the sporophores, two being found mostly growing on
leaves, and two upon branches. The distinctions between
Hainesia and Gloeosporuim, which are the two genera that
flourish for the most part on living leaves or succulent fruits,
are scarcely sufficient, since they resolve themselves into this,
that in Hainesia the pustules are brightly coloured, and in
Gloeosporium gray, pallid, or dull coloured. Hence we can
treat them both as a single genus. These parasites are
amonffst the most destructive with which the horticulturist
IMPERFECT CAPSULAR FUNGI— SPHAEROPSIDEAE 273
has to contend, or, at the least, the most insidious, and least
subject to control. They do not spread over such tracts as the
potato disease and the hop mildew, but the infected plants upon
which they appear are doomed, and these often the rarest
and most valuable. The first external indication is usually in
the form of small elevations of the cuticle, or little warts,
which cover the concealed pustules ; for a long time these re-
main unbroken, but when the sporules are mature the cuticle
is ruptured, and a globule, or tendril, of agglutinated sporules
emerge through the orifice. These sporules are either elliptical
or elongated, usually much longer than broad, and often of
considerable size, but without septum or colour. Including
the two supposed genera, not less than some 230 species are
known, to say nothing of Marsonia, which is a corresponding
genus with uniseptate sporules, and similar habit and propen-
sities. The two corticolous genera Myxos2Jorium and Melano-
stroma are not clearly distinct from each other. The habit
is similar to Gloeos]Jormm, but the species are found chiefly on
dead bark, and therefore not parasitic, or destructive. Many
of the species are credited with being stylosporous conditions
of various ascigerous Fungi. The sporules resemble those of
Gloeosporium. The series of genera in which the sporules are
produced in chains is represented by only a few species.
Hypodermium has black pustules, which, being elongated, re-
semble the perithecia in Hypoderma, a genus of the Hysteri-
aceae. Myxosporella is simply Myxosporium with the sporules
catenulate. Blennoria has discoid pustules, which bear a re-
semblance to Puccinia, to which Agyriella is closely allied ; but
the pustules are at first gelatinous, becoming hard and shining.
In Trulhda the pustules are compact and erumpent, often
having the appearance of perithecia ; the sporules are sometimes
coloured. In the two genera Myxormia and Bloxamia the
pustules are apparently pezizoid ; that is to say, the form re-
sembles a shallow cup, or concave disc, without a receptacle.
In Myxormia the sporules are joined in a chain, by a narrow
isthmus ; and in Bloxamia they are truncate, and closely applied
to each other. There are two other genera, Colletotrichum,
which is simply a Gloeosporium, with the margin of the
pustules hairy ; and Pestalozziella, in which the sporules are
18
274 INTRODUCTION TO THE STUDY OF FUNGI
cristate, and therefore analogous to Bolillarda but with con-
tinuous sporules, or a dwarfed form of Pestalozzia without
colour and without divisions. The next section is a modified
one, or at least the Scoleco-aUantosporae combines Scolecosporae
and Allantosporae in a single section. In one genus, that of
Cylindrosporium, the sporules are really filiform ; in Crypto-
sporium and Libertella elongated and falcate, but scarcely fili-
form ; and in Nemaspora they are allantoid, or sausage-shaped.
In Cylindrosporium the species are parasitic on living leaves,
and thus correspond to Gloeosporium. In the other three
genera they are saprophytic, chiefly affecting the bark of dead
branches. In Cryptosporium the sporules are mostly rather
large and robust, but in Libertella slender, oozing out in
brightly coloured tendrils. Nemaspora somewhat resembles
Lihertella, but the sporules are shorter, and allantoid. In all
three genera there are many species which are regarded as
stylosporous forms of ascigerous Fungi, and suggest analogy to
Cytospora in the Sphaeropsoideae.
The section Phaeosporae is the most typical, and includes
the genus Melanconium, which is almost the same as Fries left
it, with subglobose or oblong dark -coloured sporules, often
oozing out and blackening the orifice of the pustules. Some
of the species are associated with Sphaeriaceae of the genus
Melanconis, but others may prove to be autonomous. Crypto-
mela is analogous to Cryptospoi'ium, but with coloured sporules.
Thyrsidium is, however, a genus by itself, in which the con-
tents of the pustules are gelatinous, and the sporules are
minute, but clustered in chains at the apex of elongated
sporophores, in a capitate manner, involved in a mucous
envelope.
The Didymosporae include four genera, in which the
sporules are uniseptate, and in two of them coloured, whilst in
other two they are hyaline. Of the former, Didymosporium
corresponds to Melanconium, but with two-celled sporules ; and
Bullaria, with a single species, has the conidia connected in
chains by a narrow hyaline isthmus. Of the two genera with
hyaline sporules, it has already been intimated that Marsonia
is the analogue of Gloeosporium, with the same habit and the
same parasitic character, but the sporules are septate. Se2)to-
IMPERFECT CAPSULAR FUNGI— SPHAEROPSIDEAE 275
viyxa, in like manner, corresponds to Myxosporium, growing on
dead branches, but with uniseptate sporules.
The section Phragmos])orae includes such species as have
sporules with two or more septa, whether hyaline or coloured ;
and thus we have two subsections, the Phaoeophragmiae and
the Hyalojphragmiae. In the former Stilhosijora is the ana-
logue of Melanconium and Didymosporium, with sporules soon
oozing out and blackening the orifice ; whilst Coryneum forms
compact pustules, in which the sporules are for a long time
attached to their pedicels, and do not ooze out and blacken the
matrix. In habit the species are more pulvinate and erumpent,
being held together almost as compactly as if enclosed in a
perithecium. Scolecosporium resembles Coryneum, but the
sporules are beaked at the apex. Asterosporium has more the
habit of Stilhospora, but the sporules are compound, or rather
triradiate, resembling three sporules of Stilbosptora grown to-
gether at the base and diverging above in three rays on the
same plane. In another genus, Seiridium, the septate brown
sporules are united to each other by a hyaline isthmus, so as
to form a chain. The two remaining genera, having ciliate
sporules, are Hyaloceras, in which the
multiseptate brown sporules have a single
curved awn at each extremity ; and Pes-
talozzia, in which the sporules are crested
by one or more hyaline cilia, which are
usually divergent when more than one,
and the central cells of the sporules are
commonly coloured (Fig. 125). The
Hyalophraginiae includes but three genera;
that of Bhopalidmm, with one little-known ^^«- ^p%'^^l°f^^^^^ "^
species, has clavate, multiseptate, hyaline
sporules, aggregated in little innate brown pustules on the
leaves of plants ; and Scptoglaemn, w^hich is practically
Gloeosporium, or llarsonia, with more than one septum to
the sporules. The remaining genus is Prosthemiella, which is
the analogue of Prosthemium, but without a perithecium,
and the stellate sporules are hyaline. It will facilitate
determination to remember the instances, which are so con-
stantly recurring, in which Fungi possessing the same habit
276 INTRODUCTION TO THE STUDY OF FUNGI
and external appearance find their place in sections widely
removed from each other, as the consequence of the septation,
or multiseptation, of the sporules. Thus we have, for instance,
Gloeosporium with continuous sporules in Hyalosporae ; with
filiform sporules, as Cylindrosporium, under Scoleco - allanto-
sporae ; with uniseptate sporules, as Marsonia, under the
Bidymosporae ; and with multiseptate sporules, as Septoglaeum,
under Hyalophragmiae.
There remains but one small section to notice, and that is
the Dictyosporae, in which the sporules are divided in both
directions, so as to be muriform. Of the two genera, Stegano-
spormm is the analogue of Coryneum, having compact pulvinate
pustules, but with mm'iform, coloured sporules ; and Phrag-
motrichum, in which the sporules are concatenate, or in chains,
as in Myxormia and Seiridium, and is practically Seiridium
with the sporules septate in both directions.
BIBLIOGRAPHY
Saccabdo, p. a. "Sylloge Sphaeropsidearum et Melanconiearum," in Syllogc
Fungomm, vol. iii. Imp. 8vo. Padua, 1884.
Berlese, a. N., et Voglino. Un nuovo genere di Pirenomiceti d. Funghi
Sferopsidei. Padua, 1866.
Crie, L. a. Recherclies sur Ics Pyrenoviycetes infirieurs du group de Bepazics.
8vo. Paris, 1878.
CoRDA, J. C. Icones Fiingorum. Fol. 6 vols. Prague, 1837-54.
AnUitung zum Stvdium der Mycologie. 8vo. Plates. Prague, 1842.
BoNORDEN. Handhuch dcT Mycologie. 8vo. Plates. Stuttgart, 1851.
Zur Kcnntniss der Coniomyecten m. Cryptomyceten. 4to. Halle, 1860.
CHAPTEE XXIII
MOULDS HYPHOMYCETES
In their internal relations to each other, and their external
relations to the remaining orders, the Hyphomycetes are un-
doubtedly a well-defined and natural group. It may be, and
probably is, too rash an assumption to contend that all the
species are form-species, and only represent the conidial stage
of more perfect Fungi ; nevertheless a large nmnber of them
have been demonstrated to be merely transitionary, although
the precise mode of continuity has not been made clear. In
such a case the only reasonable course to adopt is to recog-
nise their morphological distinctions, and treat them, for all
purposes of classification, on the supposition that they may be
autonomous, and leave to the future, when their life-histories
are thoroughly known, to develop their true affinities and
relationships. The number of described species falls but little
short of 5000, and such a number is too large and important
to remain unrecognised, or without definite classification, within
the limits of present knowledge. Because many of the species
of Isaria have been ascertained to represent the conidia of
Cordyceps ; because certain of the subgenus Polyactis may be
the conidia of Sclerotinia; or even the entire genus Zygodesmium
may be so intimately related to resupinate Thelepliorae that
definite limit cannot be assigned between them, it would be
folly to expunge the whole upon suspicion, and thus increase
the difficulties in the way of the student in the pursuit of
knowledge along a path already sufficiently thorny and stony.
The general characteristics of the order are, tliat the spores,
or conidia, are naked or free, as they are in no other order,
except the Eymenomycetes and some of the Phijcomycdcs ;
278 INTRODUCTION TO THE STUDY OF FUNGI
whilst they differ from the former in having no hymeuium
and being deficient in true basidia, and from the latter in the
absence of sexual reproduction. The Fungi themselves are
either superficial or subsuperficial, and the hyphae or conidia-
bearers are, for the most part, strongly developed. Typically
there is a creeping, septate mycelium, seated upon or pene-
trating the matrix, which gives rise to erect, more or less
developed, hj^Dhae, or spore-bearers, which produce terminally
or laterally naked spores or conidia. Most of them are
saprophytic on dead animal or vegetable substances, whilst a
limited number are parasitic upon living plants.
The fom^ families are the Mucedineae, or " white moulds,"
with the threads colom^less, pale, or brightly coloured, often
fasciculate, but not coherent, with conidia of the same colour ;
the Dematieae, or " black moulds," having the hyphae brown,
or black, rather rigid, and not coherent, rarely pale and then
with the conidia blackish ; the Stilbeae, with the hyphae either
pallid or brown, densely cohering in long stem-like fascicles ;
and the Tubercularieae, with the hyphae pallid or brown,
densely conglutinate in wart-like pustules, or sporodochia or
spore-beds, often forming a rather thick stroma at the base.
Thus, it will be observed that in two of the families the
hyphae are free and distinct from each other, being typically
pale in the one and dark coloured in the other ; whilst in the
other two families the hyphae are closely coherent and elongated
in the one, and shortened, conglutinate, and stromatic in the
other. The latter are for the most part erumpent and pustular.
This, then, is the primary division of the Hypliomycetes, of
which the largest, and most typical, of the two great sections
is that in which the erect threads or conidiophores are free of
each other, and not united in a common stem. These are the
moulds, which as yet are not known to possess any but a
simple and asexual reproduction, by means of conidia, but
which are considered to be genetically connected, by some
means not clearly manifest, with species belonging to other
orders of Fungi, and especially of the Ascomycetes. For the
better understanding of the principles on which the classifica-
tion of these imperfect Fungi has been reduced to a system, we
must examine the sections in further detail, commencing with
MO ULDS—H YPHOM YCE TES
279
the Mucedines. In this family, as in all the primary divisions
of the orders devised by Saccardo, the spores, or conidia, hold
the first place, so that not only are the genera limited by the
septation, or non-septation, of the spores, but this also forms
the basis of the first subdivision into the Amewsporae (Fig.
126), in which the conidia are spheroid or shortly cylindrical ;
the Didymos2)orae, in which the conidia are
oblong or fusoid, and uniseptate; the Phragmo-
sporae, in which the more elongated conidia are
two, three, or many septate ; the Staurosporae,
in which the conidia are stellate, radiate, or
trifurcate ; and the Helicosporae, in which the
elongated conidia are spirally convolute. It is
not clear that the last is a necessary or homo-
geneous section, or that it is at best any other
than a subsection of the Phragmosporae, with
the elongated and septate conidia, instead of
being simply curved or flexuous, curved more
strongly so as to be spirally convolute. Thus,
then, having discovered that any given mould
has a simple or compound stem, it is incumbent to ascertain,
on the assumption that the stem is simple, whether the threads
are carbonised, or only hyaline, or bright coloured, and thus dis-
cover the one of the four families in which its place has to be
found. It being determined, for example, that the mould in
question is a Mucedine, the next step is to find the conidia, and
ascertain if they are continuous, or in what manner they are
septate. Up to this point it may be possible to place a sterile
mould, but from this point forwards it is manifestly impossible
to proceed, in the absence of all fructification. This leads us to
observe how utterly futile it is to attempt the determination
of even the genus, much less the species, of any mould, in the
absence of conidia. Novices are apt to infer that it is not
only possible, but easy, to give a name to any mouldy tuft
which presents itself as such to the naked eye, but possessing
only mycelium and threads, without any indication of the
character of the spore. The labour which is expended in any
such endeavour is wasted, and it is always better to abandon
the task at once, not only in this but in other orders, unless
FiG.126.— i^/em■-
sjiora lucida,
one of the
Amerosjmrac.
:8o
INTRODUCTION TO THE STUDY OF FUNGI
conidia, or spores, of some kind can be detected, otherwise the
endeavour can only terminate in vexation of spirit.
Eesuming our survey of the system at the point where it
is necessary to determine the character of the spore, or conidium,
and if it is uniseptate to seek it in the Bidy-
mosporae, but if further septate in the Phrag-
mosjoorae, we shall soon discover that the
greater number of species have conidia which
are not septate at all, and therefore belong to
the section Amerosporae. At this point we
may leave the conidia and revert to the
hyphae or threads which bear them. In some
cases we shall observe that the conidiophores,
the Macroneineae
branched once,
myces representing or conidia-bearers, are long threads, which
are sometimes simple, but in most cases
twice, or many times ; these generally form
large, conspicuous woolly tufts, easily recognised by the naked
eye, and constitute the subsection Macronemeae, or, as we might
say, the subsection in which the threads or hyphae are strongly
developed, and quite distinct from the conidia (Fig. 127). Then
there is another and smaller section, the Micronemeae, in which
the threads are very short, and mostly
unbranched, so short, indeed, as only
just to be recognised, and, at times,
scarcely different from the spores or
conidia themselves (Fig. 128). In
nearly all the subdivisions of the various
families of the Hyphomycetes, such
subdivisions being based upon the char-
acter of the conidia, the genera are
associated in these two groups of
Macronemeae or Micronemeae, according as the conidia-bearing
threads are long and well developed or short and almost
obsolete. It would be wearisome and unnecessary here to
detail all the varied modifications of the conidia-bearers, or
the conidia, which are taken advantage of in the construction
of genera, or groups of genera. It must suffice to say that
most of the distinctions are based upon the form, or mode of
arrangement, of the conidia about the threads. For instance.
Fig. 128.— One of
the Microneineae Aegerita.
MO ULDS—H YPHOM YCE TES
Fig. 129.— FcnicilUum
with the conidia iu
chams.
a distinction is made between such conidia as are solitary and
those which are produced in chains, or catenulate (Fig. 129) ;
between those which are solitary and those which are clustered
at the apex of the hyphae, or its branches,
so as to form more or less dense heads, or
clusters of conidia ; between those in which
the conidia are terminal and those in which
they are lateral or dispersed. Other dis-
tinctions are derived from the hyphae them-
selves, whether simple or branched ; or if
simple, whether inflated at the apex or not ;
and if branched, whether simply furcate,
repeatedly divided, or if the branches are
arranged in whorls or verticillate. All
these are details which are readily gathered
from the diagnoses of the separate genera,
and we have said sufficient to indicate the
principal features which have to be taken
into account in the determination of the genus to which any
particular mould may belong.
Although the above observations apply in the first instance
to the 31'ucedines, they apply also generally to the Dematieae,
with the exception that in the divisions based on the forms
of the spores, or conidia, there will be found an additional
division, the Dictyosporae, in which the conidia are divided in
both directions, so as to be clathrate or muriform. Some of
these conidia will therefore present the appearance of twenty
or more simple cells, aggregated into one large complex
conidium. Judging from the facility with which each cell of
these compound conidia germinates, it may be inferred that
each cell is a reproductive unit, and is in itself a perfect
conidium, capable of reproducing the species. So in respect
to uniseptate or multiseptate conidia, in a linear series, each
cell is capable of germination, and even, in some instances, of
separating itself from its sister cells, when arrived at maturity
(Fig. 130). Mr. Worthington Smith, in his observations on
Fusarium solani, has intimated that, although some of the
segments of the conidia germinate at once, others are capable of
undergoing a period of rest. He says, " Sometimes these little
INTRODUCTION TO THE STUDY OF FUNGI
Lodies do not germinate at once, but hibernate for a short time,
generally varying from three weeks to three months, commonly
two months, and during this period they become slightly
spinulose and faintly tinted with a brownish hue. These
little bodies, there-
fore, hibernate after
the manner of rest-
ing spores, and it
is possible that
many of them rest
during the entire
winter.
Assum-
FiG. 130.
Conidia of Fusarium. B, mature coniJium ;
C, cells germinatiug ; D, cells separating, and be-
coming rounded ; E, separated cell after a period of
rest-germinating. After Smith. Macmillan and Co.
ing that the seg-
ments of the conidia
of Fusarium are
capable of forming
a thicker integu-
ment, and hibernat-
ing through the
winter, there is no
reason why, from
analogy, other
conidia, belonging to other genera, may not be capable of
a like modification, and thus aid in the perpetuation of the
species. It is almost certain that the thin -walled conidia
are unable to survive the winter, and hence the question
arises as to how the rejuvenescence of the Jlyj^Jiomycetes is
assured; for, although in some cases a perennial mycelium may
explain the difficulty, it cannot do so in the parasitic species,
such as Bamularia, Ovularia, and Cercospora, where the
destructive fungus appears as a pest on the living leaves, year
after year. As an example, the leaves of the " ground ivy "
{Gleclwmd) through the autumn will present hundreds of leaves
with the white blotches of Ramularict calcea, sometimes every
leaf more or less affected, and during the winter most of these
leaves will die and decay. With the spring there will be a
carpet of green leaves again, without a spot of Ramularia; but
as summer advances the pest appears as profusely as ever, and the
1 Diseases of Field and Garden Croi^s, by "\V. G. Smitli, London (1884), p. 33.
MOULDS— HYPHOMYCETES 283
leaves are blotched with white. Can we answer the question
satisfactorily and confidently as to how the continuance of the
parasite has been secured ? It is possible that a perennial
mycelium within such of the plant's tissues as have survived
the winter may be a sufficient cause, but it is doubtful whether
this is the only method in which the perpetuation of the
Bamularia has been assured. If it should be contended that
the decaying leaves and petioles of the previous year must
contain the germs of the parasite, and that the young leaves
are infected thereby, this only removes the difficulty a step
further, for it has to be shown in what form the germs have
been preserved, as it must have been by some form of resting
spores or a resting mycelium capable of producing germinating
bodies. This is one of the problems which is left for the
future to solve.
So many of the Mucedines and " black moulds " have been
ascertained to have relationships with the higher Fungi that it
is impossible to do more than briefly allude to a few. In the
genus Oidium a number of the species are the conidia of species
of the Urysiphei, such as Oidium leucoconium, the rose mildew,
of Sphaerotheca pannosa ; Oidium erysiphoides of Erysiphe
Ifartii ; Oidium monilioides of Erysiphe graminis. The
common fruit mould Aspergillus glaucus has the reputation of
being the conidia of Eurotium lierlariorum. Some of the
species Botrytis, of the subgenus Polyactis, are the conidia of
small species of Peziza, such as Sclerotinia sclerotiorum. The
bright yellow mould Sepedoniumchrysospermum, vfhioh attacks
decaying Boleti, and converts them into a mass of golden
powder, develops Hypomyces chrysospermum, one of the
Sp)hcieriaceae, of which the mould constitutes the conidia. If
we investigate the British species of this genus Hypomyces}
we shall find that all have their conidia in some of the moulds.
As, for instance, Verticillium agaricinum of Hypomyces
ochraceus, and Verticillium lactescentium of Hypomyces
terrestris ; also Verticillium microspermum of Hypomyces
hroomeanum ; Diplocladium p)enicilloides of Hypomyces
aurantitts ; Diplosporium album of Hypomyces violaceus, etc.
The same kind of association prevails also amongst the
1 Monorjra'ph of British Hypomyces, by C. B. Plowright.
284 INTRODUCTION TO THE STUDY OF FUNGI
Demuticae, for TrickosjJorinm fuscum is found forming the
subiculum of Bosellinia aquila, and the common Bispora
monilioides is reputed to constitute the conidia of a small
Peziza, hence called Bisporella monilifera ; but this appears to
us a doubtful case. FusiclacUum clepressum is reported to be
the conidia of Phyllacliora angelicas, and Pohjthrincium
trifolii of Phyllachora trifolii. Species of Cladosporium,
of Cladotrichum, and Hehnintliosporium respectively are be-
lieved to be related genetically to various species of the
Sphaeriacei, and especially species of Macrospoj'ium to certain
species of Phospora. It is sufficient for our purpose to suggest
these relationships as indicating the evidence on which the
Hyphomycetes are concluded to be imperfect Fungi, and
principally conidial forms of Ascomycetes.
After this digression we may return to the two inferior
families of the order, in which
the hyphae are fused into a com-
, mon stem. The Stilheae (Fig.
■^ 131) are of a more imposing
.^ appearance than the Tubercu-
larieae, and perhaps of a higher
^J' \\Y / ' , /,,' development. There are not
more than about five hundred
Fig. 131. — bliibum vulyare. .
described species, and these are
grouped in two parallel sections : the Hyahstilheae, in which the
hyphae and conidia are pallid ; and the Phaeo stilheae, in which
the hyphae and conidia are typically dusky coloured. Thus
these two sections correspond to the Mucedineae and the
Dematieae.
The Hyalostilheae, as far as at present known, are less
variable in fructification than the Phaeostilbeae, it being found
necessary to recognise but two of the subsections : the
Amerosporae, in which the conidia are globose or oblong,
and continuous ; and the Phragrnosporae, in which the conidia
are septate. The latter is a very small section, of some seven
or eight species, so that practically the Hyalostilheae have small
and continuous spores, or conidia. The subsidiary arrangement
is very much on the same lines as in the moulds. The
principal genera are the old ones of Stilhum and Isaria, with
MOULDS— HYPHOMYCETES 285
their allies. The former has a capitate form, typically a com-
pound stem, with a globose head ; and the latter assumes a
cylindrical or club-shaped form, the stem and head being
continuous. The surface is generally powdery with the
minute conidia. As to their autonomy, it is known that,
in several instances, the species of Stilhum represent the
conidia of a peculiar genus of the Sphaeriaceae, that of
SpJiaerostilhe, whilst others have given no indication of such
an association. Of Isaria the greater proportion, probably all
which flourish on dead insects, are the conidia of Cordyceps}
The Phaeostilbeae are more variable in their conidia, being
grouped in five sections, as in Dematieae, of which it is the
analogue, and they represent a somewhat higher develop-
ment. Some of the genera exactly correspond to genera of
Dematieae, but with a compound stem, as for example Simroctjhe
and Periconia, Fodosporium and Helmintliosporium, Sclero-
graphmm and Mystros2')orium. Instances of undoubted rela-
tionship with the higher Fungi are rare, but in some cases
it is suspected.
The family of Tulercularieae includes genera which recede
from the moulds in their compact form, thickish stroma-like base,
more or less pustular, erumpent habit, and somewhat gelatinous
consistency, which suggest analogies with such genera as Dacryo-
myces, amongst the Tremellinae. Here again are two parallel
sections, the Mucedineae and ihe Dematieae, in the former of which
the colour is whitish, or brightly coloured, and in the latter
dusky or black. The subdivisions follow the same plan as in
the preceding families, firstly into sections based on the septa-
tion of the conidia, and afterwards into genera, or groups of
genera, according to the character of the stroma. The typical
genus, Tuber eularia, with some sixty species, is composed
chiefly of the conidia of corticolous species of Nectria, of
which a familiar example may be found upon nearly every
dead twig of currant bush lying on the ground^ (^ig- 132).
The whole surface of the twig will be found to be covered
from end to end with little bright pink prominences, bursting
^ See Vegetable Wasps, etc., by M. C. Cooke, London (1892), p. 189.
2 "A Currant Twig and Something on It," by M. C. Cooke, in Gardener's
Chronicle, 28tli Jan. 1871.
!86
IXTRODUCTIOX TO THE STUDY OF FUNGI
through the bark at regular distances, scarcely a quarter of an
inch apart. Towards one end of the twig the prominences
will doubtless appear of a darker colour, almost
blood-red, and, intermediate between the two,
pink pustules sprinkled with red dots. The dark
red pustules are composed of a number of minute
red bodies clustered together, the perfected condition
of the parasite (Fig. 133). By removing the bark
it will be seen that the pink bodies have a paler
stem, which expands above into a rather globose
head, covered with a mealy bloom. This is the
Tuhercularia, which at its base penetrates to the
inner bark, and there the threads of mycelium
branch in all directions, within the bark, but do
not extend to the woody tissues beneath. The
head, more closely examined, will be found to
consist of delicate parallel threads, which are
compacted together into a common stem, with its
head. Some threads are simple, others branched,
bearing here and there little bodies, easily detached,
which are the conidia, and form the mealy bloom
134). The darker clusters,
when examined in the same manner, will present,
instead of one uniform head, a cluster of smaller globose bodies,
closely packed together, or, in some cases, a circle of these dark
bodies around a smooth pink centre. These darker bodies are
the mature Nectria, which grow at length upon the same stroma,
and are the ultimate development of the pink pustules which
produce the conidia. Each of the dark bodies is a perithecium,
or receptacle, which encloses the fruit, consisting of sporidia, con-
tained in asci (Fig. 1 3 3 at G). Here, then, we have the Tuhercti-
laria in the first instance, as a smooth, compact, pink, erumpent
pustule, the stem composed of numerous delicate threads con-
glutinated together, and sprinkled with minute conidia ; then
the darker capsular Nectria originates from the same stroma,
these capsules containing the fully-developed sporidia enclosed
in asci, — the first stage representing the Tiibercularieae family
of the Hyphomycetes, the last stage belonging to the Hypocreaceae
family of the Pyrenomycetes. Hence, as the first is an im-
low. Gard. ^f ^j^g gurfacc (Fig.
MO ULDS—H YPHOM YCE TES
287
Fig. 133.— Tubercularia, D ; with Nectria, E ;
section, F ; and asci, G. Gard. Chron.
perfect condition, the Fungi to which it belongs are characterised
as imperfect Fungi.
We might follow the same process with one or other of
the species of i^Msa?'MMn,
which is a genus in the
present family, the con-
idia of which are com-
paratively large, fusi-
form, and mostly three
or five septate. Some
of them are, in like
manner, only the con-
idia of some more
highly developed Fun-
gus, and often a species
of Nectria. The pus-
tules are not so com-
pact, sometimes effused,
seldom with a determinate stroma, and rarely with the hyphae
much developed. The genus altogether is much more variable
than Tubercularia,
and not so well con-
stituted, so that pos-
sibly it will be broken
up into more homo-
geneous genera in the
near future. On the
faith of some obser-
vations made by Mr.
Worthington Smith,
the conidia must be
regarded as bodies of
a much higher order than their analogues in Tubercularia.
Not only are they capable of dividing at the joints, and
each segment vegetating as a separate unit, but these may
be converted into chlamydospores, or at least have a thickened
epispore, capable of hibernation. When this is confirmed it
will go far towards necessitating a revision of the classification,
so far as an association with Tubercularia is concerned.
Fig. 134.— B, section of Tubercularia ; C, conidia.
Gard. Chron.
288 INTRODUCTION TO THE STUDY OF FUNGI
Tor the purposes of classification, the genera of the Tuber-
culariae are grouped according to the general principles adopted
in the Mucedines and Dematiaei, and in fact throughout the
Saccardian system — that is to say, the sections are based on the
septation of the conidia, whether unicellular, bilocular, multi-
cellular, or with stellate or helicoid forms. In each of the
sections the genera are characterised by the features presented
by the sporodochium, or spore-bed, and the development of the
gonidia, whether produced singly or in chains. There are
some forty-two genera in all, which it would be somewhat
tedious to describe in detail.
The Tuherculariae Bematieae contain such genera as possess
the habit and development of Tuherculariae, but with coloured
hyphae, and similarly coloured, or rarely of hyaline, gonidia.
They are less numerous in genera and species than the previous
section, but many of the genera correspond in habit and appear-
ance, differing only in the coloured hyphae. Not long ago,
when the septation of conidia was not held to be of generic
importance, or the coloured or uncoloured hyphae a fact of
moment, the few genera which were contained in the Tuber-
cularieae were rather a heterogeneous collection of species, held
together by some superficial character, and embracing forms
which are now dispersed through several genera. The large
increase of genera which has resulted from the adoption of a
more precise method of classification is therefore something
more than a numerical gain, since it is the result of a closer
investigation, and the application of a more uniform and
scientific system, which in the end must conduce to the
benefit of the student, and, encouraging a more rigid examina-
tion of species, tend to the advancement of this branch of
biological study.
BIBLIOGRAPHY
Saccardo, p. a. Sylloge Fungorum, vol. iv. — " Hyphomyceteae. " Padua, 1886.
CoRDA, J. C. Prachtflora Euro}). Schimmdhild. Fol. Col. plates. Leipzig, 1839.
The same in French as Flore illuslrte cle Mucedinies d' Europe. 1840.
"Die Pilze Deutschlands," in Sturm, De^itscldamls Flora. 12nio.
1829-41.
Anleitung zum Studium dcr Mjkologic. 8vo. Prague, 1842.
MOULDS— HYPHOMYCETES 289
Hakz, C. 0. Einicjc neue Hyxihomycctcn u. Beitr. z. Systcmatik dcrsdben.
Moscow, 1871.
BoNOiiDEN, H. F. Ilandbuch ckr Allgcmeincn Mykoloyic. 8vo. Plates.
Stuttgart, 1851.
Link, H. F. " Observatioues in Ordines Naturales. " Berlin Magazine. 4to.
Berlin, 1809.
"Species Hyphomycetnra et Gyninomycetum," in Willdenow's SiKcies'
Plantamm, vol. vi. 8vo.
Fresenius, G. Beitrdge zur Mykologic. 4to. Frankfurt, 1850-63.
Cooke, M. C. The Hyplwmycetous Fungi of the United States. 8vo. 1877.
MiYABE, KiNGO. "The Life History of Macrosporium parasiticuni." 8vo,.
Annals of Botany. 1889.
CONSTANTIN, J. Les Muccdinees simj)les. Paris, 1888.
CHAPTEE XXIV
MICROBES SCHIZOMYCETES AND SACCHAROMYCETES
The recognition of the Schizomycetes, or " splitting Fungi,"
as an order, is of comparatively recent date, and the entire
study, notwithstanding all that has been done, is still in an
elementary condition. The very minute organisms of which
the group is composed have long been recognised, but even now
it is open to doubt whether they should be associated with
Algae or with Fungi, or outside of both. As part of the
Infusoria, Ehrenberg made the first attempt at their classifica-
tion in 1838. Then they were transferred, almost bodily, to
Algae, in 1872, whilst, more recently, they have been held to
be most closely related to Fungi, and united to Fungi by
Saccardo in 1889. It is of but small import whether they
should, technically, be regarded as Fungi or only as allies : they
evidently are closely related, and, notwithstanding their minute
size, are of too great importance to be practically ignored. They
are defined as " unicellular plants, which multiply by repeated
subdivision, in one, two, or three dimensions of space, and also
frequently reproduce themselves by spores, which are formed
endogenously." Mr. Grove points out that they differ from
Algae : " On account of their want of chlorophyll they are
reduced to live on ready-organised substances, as are Fungi
generally. The Schizomycetes, therefore, produce in their sub-
stratum, or in the fluid which they inhabit, very considerable
and striking decompositions. They perish in pure water con-
taining no decomposable substance. They grow, therefore,
exclusively in organic liquids, or in water, or on damp spots,
where there is an abundance of organised matter." ^
^ Synopsis of the Bacteria and Yeast Fungi, by W. B. Grove, B.A., London,
SCHIZOMYCETES AND SACCHAROMYCETES 291
The term " Microbe " has been employed, in a general sense,
by the French, and adopted from them to indicate all the
minute organisms which are now recognised under Schizo-
mycetes and Saccharomycetcs, /,v';"wr--V'v.
whilst in many cases the
former are often spoken of
simply as " Bacteria " and the
latter as " ferments." These
" bacteria " appear " in liquids
examined under the micro-
scope as small cells of a spheri-
cal, oval, or cylindrical shape,
sometimes detached, some- Fig. 12,0.— Bacterium termo. Cliatto and
times united in pairs, or in Wmdus.
articulated chains and chaplets (Fig. 135). The diameter of
the largest of these cells is two micromillimetres and that of
the smallest is a fourth of that size, so that at least 5 0 0 of the
former and 2000 of the latter must be placed end to end in
order to attain the length of a millimetre. It is therefore
plain that a magnifying power of 500 to 1000 diameters, or
even still higher, is required to make these beings clearly
visible under the microscope." ^
Besides the vegetative multiplication of these cells in one,
two, or three directions, there is a double method of formation
of spores, which must be described. So long as all the con-
ditions remain favourable to growth and vegetative develop-
ment, only vegetative multiplication prevails. If the cells
can obtain sufficient food, and the food is of exactly the right
kind, the rate at which they grow is marvellous. " Colin cal-
culated that a single germ could produce by simple fission
two of its kind in one hour, in the second hour these would be
multiplied to four, and in three days they would, if their
surroundings were ideally favourable, form a mass which can
scarcely be reckoned in numbers — or if reckoned, could scarcely
be imagined — 4772 billions. If we reduce this number to
weight, we find that the mass arising from this single germ
would in three days weigh no less than 7500 tons. Fortu-
nately for us, they can seldom get food enough to carry on this
^ Microbes, Ferments, and Moulds, by E. L. Trouessart, Loudon, 1889.
292 INTRODUCTION TO THE STUDY OF FUNGI
appalling rate of development, and a great number die both for
want of food and because of the presence of other conditions
unfixvourable to their existence. Vegetative multiplication
only takes place when the conditions are extremely favouraljle
to the growth of the organism. If nutrition is interfered
with in any way, or if the removal of excretionary products is
obstructed, or if there be a large amount of oxygen present,
marked changes may at once be observed in the appearance of
the protoplasm of the micro-organism. It becomes granular,
then a small bright point appears in each cell ; this point
gradually increases in size until its diameter may be greater
than that of the original organism. This large, clear, rounded,
ovoid, or rod-shaped node is known as a spore, or resting spore,
by which the species may be continued although the parent
should perish. The shape varies slightly in different species,
but in every case it has a dark limiting outline ; it is devoid
of colour, and is highly refractile. The dark outline of the
spore is usually surrounded by a pale, soft, gelatinous envelope,
the substance of which may, in some cases, be accumulated in
rather larger quantity near the two poles of the refractile body.
As soon as these bodies make their appearance, degeneration
of the protoplasm of the bacteria in which they are found
immediately follows, but the period at which the death of the
protoplasm actually takes place varies in different cases.
Where the spores are small they may lie for some time im-
bedded in the protoplasm of the cell, which, as it degenerates,
leaves the resting spore free to be carried about from place to
place, by currents of air or water, to be developed when the
conditions of moisture, temperature, and food supply again
become sufficiently favourable. Where the diameter of the
spore exceeds that of the bacterium, it may be situated in the
centre, giving rise to a spindle-shaped organism ; or it may be
at one end, when the organism becomes clubbed or pendulum-
shaped. The spore in this case appears to escape more readily.
This method is that which De Bary has called endospore
formation." ^
Another kind of spore is called artlirospore, which is also
1 Bacteria and their Products, by G. S. Woodhead, M.D., London (1891),
p. 33.
SCHIZOMYCETES AND SACCHAROMYCETES 293
defined by De Bary. In this there is a combination of spore
formation and of fission ; the mother cell undergoes division into
a series of daughter cells, a few of which differ from the rest
in very important and essential points. There appear to be
two kinds of anthrospores : one form, met with in Leuconostoc,
for example, where simple vegetative division of small round
bacteria goes on regularly, so long as the conditions are favour-
able, and a regular chain is formed. In this chain there
appear at intervals micrococci, which differ from the remainder
of the elements of the chain in the following points. As soon
as the conditions of nutrition are altered they do not, like the
other parts of the chain, die off, I)ut they become somewhat
larger than the rest, acquire a more distinct outline, become
thicker- walled, and their protoplasm grows darker. Eventually
they become free by the deliquescence of the gelatinous envelope,
and may claim the name of spores, because, when placed in the
fresh nutrient solution, they develop into new rows of beads
like those of , the mother plant. This body has most of the
characteristics of the resting spore, but it is not formed within
the protoplasm of the vegetative organism, but by a process of
fission, and as a result of vegetative division. It is possible
that there is as much differentiation of the protoplasm as there
is where the spore is formed within the cell, the only distinc-
tion being that the separation between the spore and the
vegetative element of the chain takes place at an earlier stage,
and more completely, than in endospore production. The
reverse takes place in Bacterium Zopjii, which, during the
vegetative stage, consists of short rods, then of motionless
filaments, and, if the temperature be lowered, of short motile
rods. As soon as conditions become unfavourable the rods,
apparently by a simple process of fission, are divided into
short roundish cells, which retain their vitality for a consider-
able time, and, when placed under favourable conditions, act as
spores — that is to say, they develop into the original charac-
teristic rod-shaped bacteria.
The functions of the Schizomycetes have been described
as exciting peculiar decompositions, and transforming com-
plicated chemical combinations into simpler ones. This
chemical action consists in the production and excretion of
294 INTRODUCTION TO THE STUDY OF FUNGI
colouring matters, such species being distinguished as chromo-
genous ; in the exciting of various fermentations, and hence
called zymogenous ; and in the decomposition of the humours
of animal and human bodies, whereby diseases arise, and these
are 2^a'i^'ogenous species. Some authors prefer to group them
as pathogenous, zymogenous, and saprogenous.
The classification of this order must still be regarded as
imperfect and transitionary, and will be the subject of much
change in proportion to the development of knowledge which
experience will afford. There are some who are prepared to
accept all the morphologically or physiologically distinct forms
as different species, and with them the number of genera and
species would be large. There are others who hold that most
of the Schizomycetes pass through a series of adaptive forms,
influenced by surrounding circumstances, and modified by
external conditions, so that at one time it may have the form
of a Bacillus or of a Bacterium, of a Micrococcus or a Spiro-
chaete. In this latter case the number of genera and species
would be reduced to their lowest expression. Perhaps, in the
present state of knowledge, the wisest course is to accept the
various forms as they appear to be, on the presumption that
they are autonomous, and leave condensation and reduction to
the gradual operations of the future, and the verification of
facts or assumptions, in the light of experience. The arrange-
ment adopted by Saccardo recognises three primary groups, or
families. The Tricliogenae, with three evokitionary states — the
filament, the rod, and the coccus — of which the filament is the
primary condition, vaginate or evaginate, fixed at the base or
radiating from a central point, rarely entirely free ; rods and
cocci included in the filaments. The second family, Baculogenae,
also with three evolutionary states — rods, filaments, and cocci.
In this group the rod is the primary state, the filament
secondary, never vaginate, or fixed, or radiating, formed by the
indefinite prolongation of a single rod or the union of many.
The third family, Coccogenae ; there is but one state, that of
the coccus. Beyond this it would not be profitable to follow
the subdivisions.
No one can doubt for a moment that the pathogenous
species are of immense importance as objects of study and
SCHIZOMYCETES AND SACCHAROMYCETES 295
investigation, in face of the contention that in men and
animals, and probably plants, they are the associates, and in
many cases the causes, of disease. Since =, [1
the discovery of the Bacillus of anthrax, y^<=^^^^^J
or splenic fever (Fig. 136), facts have 'MT^j
rapidly developed in the association of -^^'
microbes with contagious diseases, which ""^^^
previously were theoretically attributed to V^^"^^ S^
many sources. That which at first was «'^W['''^%'/' /''
an hypothesis is now an ascertained „, 51, „ ...
'' ^ _ 1*10. 136. — Bamlus an-
fact ; but before an infectious disease can thrads. Chatto and
be considered due to the presence of a '^Vmdu.s.
specific microbe, it must submit to the test of the four
rules established by Koch. (1) " The microbe in question
must have been found either in the blood or tissues of the
man or animal which has died of the disease. (2) The
microbe taken from this medium, and artificially cultivated
out of the animal's body, must be transferred from culture to
culture, for several successive generations, taking the precautions
necessary to prevent the introduction of any other microbe
into these cultures,
-=— SO as to obtain the
specific microbe, pure
from every kind of
matter proceeding
Fig. 137. — Development of the Bacillus anthmcis. fi^-nm the bodv of the
After Ewart. "^
animal whence it
originally came. (3) The microbe thus purified by successive
cultures, and reintroduced into the body of a healthy animal,
capable of taking the disease, ought to reproduce the disease
in question in that animal, with its characteristic symptoms
and lesions. (4) Finally, it must be ascertained that the
microbe in question has multiplied in the system of the
animal thus inoculated, and that it exists in greater number
than in the inoculating liquid." These conditions have been
fulfilled in the case of a large number of diseases, such as
anthrax, swine-fever, smallpox, erysipelas, etc., and the microbe
theory of the origin of contagious diseases is, in principle,
accepted as fact.
296 INTRODUCTION TO THE STUDY OF FUNGI
The close study and prolonged investigations of patho-
genous species led to important results in the practice of a
system of vaccination, which has been adopted not only in
anthrax but also in other contagious diseases. It has long
been known that in a number of diseases of this class one
attack carries some immunity against that particular disease
in the future. The process had been employed in smallpox,
and it was found that vaccination of a mild form commonly
ensured the individual, at least for a lengthened period, against
subsequent attacks from the virulent form. It was thought to
apply this to animals in the case of anthrax, if a mild form
could be obtained for the purpose. In 1880 Greenfield
announced the first indication of the modification of anthrax
virus, and from that time forward there was a steady advance
in the production of a protective vaccinal fluid for anthrax.
Pasteur attributed the diminution of the virulence of the
bacillus to the action of heat, in the presence of oxygen, but
Chauveau contended that heat alone was sufficient. By culti-
vating the bacilli successively in a temperature of from 42°
to 43° Cent., they were found to lose all their vitality in about
six weeks, this loss going on progressively with the rise of
temperature. Sheep inoculated with the culture, after twelve
days' heating, only succumbed to the extent of one half.
After twenty-four days of heating, inoculation did not cause
the death of a single animal. After twelve days more, inocula-
tion with virulent anthrax blood only caused slight febrile
conditions. Absolute protection could only be secured by a
second vaccination with the attenuated lymph. It was demon-
strated further that the modified action of the bacilli was
transmitted to their spores, and that, when produced, these
sprouted, not into virulent anthrax bacilli, but into modified
anthrax bacilli, suitable for vaccination.
It follows, then, that when animals are inoculated with a
liquid containing bacilli, of which the virulence has been
attenuated by culture, carried as far as the tenth generation,
their lives are preserved ; they have the disease in a very mild
form, and, as a result of this treatment, they are henceforward
safe from a fresh attack of the disease, — they are vaccinated
af^ainst anthrax. Other methods have been tried for the
SCHIZOMYCETES AND SACCHAROMYCETES 297
purpose of modifying the virulence of the original virus, and
with more or less of success. As, for instance, a solution of
carbolic acid, of one part in six hundred, destroys the
microbes, whilst a solution of one part in nine hundred
attenuates the virulence without producing spores. What-
ever the means, the principle is the same — the reduction of
virulence in the bacilli, so as to produce by inoculation only
a mild form of the disease.
We have now, writes Dr. Woodhead, " a whole series of
diseases from which immunity may be conferred by the
inoculation, or introduction into the tissues of an animal
of the soluble products of pure cultures of micro-organisms.
In America hog -cholera has been vaccinated against, the
vaccinator using the sterilised cultures of the hog -cholera
organism as his protective virus. Wooldridge, who was
the first to adopt this principle in connection with anthrax,
was followed by Pasteur and Perdrix, and by Hankin. Fowl-
cnolera, certain forms of septicaemia, and a number of other
diseases, amongst which may be mentioned hydrophobia — in
which, however, the facts do not belong to quite the same
order — all were brought within the same zone, when it was
found that the introduction of the sterilised products of a
specific organism, first in minute doses and then in gradually
increasing doses, could confer a protection against the subse-
quent action of even the most virulent organism that, under
ordinary circumstances, gives rise to the same products as
those injected."
The discovery of bacteria in plant diseases is more recent,
although Bechamp noticed the presence of microzyma, or
l)acteria, in the affected parts as long since as in 1869. Still
at that time, and long after, they were held to be the
associates, and not the cause, of disease. In 1880 Dr. Burrill
declared tlie shrivelling of pears to be due to a species of
bacterium, and in 1882 Wakker of Amsterdam attributed the
jaundice of hyacinth bulbs to the same cause. In 1885 a
bacterium was detected in vines said to be diseased by
Phylloxera, and affirmed to be the true cause of the disease.
More recently still, and the California vine disease ^ was
^ Gardener's Chronicle, July 1893.
298 INTRODUCTION TO THE STUDY OF FUNGI
attributed to the presence of bacteria. In 1891 Dr. Halsted ^
apparently determined that a rotting disease of cucumbers
and melons was caused by microbes, and that not only could
healthy plants be infected, but the virus could be transferred
to tomato plants, rapidly producing decay. The destructive
" Peach yellows," which long baffled all efforts to discover its
cause, has been found to contain these organisms, and efforts
are being made to trace its bacteriological relationships.
Finally, the pear blight which Dr. Burrill investigated in
1880, and which is sometimes called "fire blight," was finally
determined in 1884 to be the result of the attacks of Micrococcus
amylovorus, otherwise named Bacillus amylovorus. In this
species, althougli the formation of zooglaea has never been
observed in the tissues of tlie tree, or upon solid media, they
occur with much regularity in fluid cultures, when placed
under favourable concUtions for rapid growth. They are
produced to some extent throughout the fluid, but are most
abundant in the thin pellicle which forms upon the surface.
They often appear the more distinctly by being surrounded by
a colourless layer, free of bacteria, which is an extension of
the basal stratum of the zooglaea mass. This branch of the
inquiry is, however, of such recent origin, and is in such
elementary condition, that it would be imprudent to affirm too
much, or indulge too freely in speculation.
Thus much, then, for the Microbes, which are regarded
generally, and spoken of, as the organisms which are instru-
mental in producing putrefaction. It is remarkable wliat a
voluminous literature has already accumulated, within a few
years, which may be accepted as some evidence of its im-
portance. The subject may not affect business interests so
much as the cognate one of the Fungi of fermentation, but it
is more than suspected that it has a very intimate relation to
life and death.
The yeast Fungi are very simple and low forms of vegetable
life, although of a more imposing size than the Schizomycetes,
or Microbes, to which we have given brief attention. The
yeast Fungi, which are the agents of fermentation, are repre-
sented in old books under the name of Torula cerevisiae, and
• Gardener's Chronicle, 3rd June 1S93.
SCHIZOMYCETES AND SACCHAROMYCETES 299
the generic name of Torida prevailed, in all notices of yeast
plants, for very many years after it was demonstrated and
known that Torulct had nothing whatever to do with them.
But faith, fanatic faith, once wedded fast
To some dear falsehood, hugs it to the List.
The cells, in budding, give rise to similar cells, attached to
each other in chains, resembling the conidia in Oidium, or,
less closely, those of some species of Torida, in which latter
genus the cells are dark coloured, almost black. The similarity
of form led to the confusion of names ; whilst in point of fact
the yeast Fungi have no affinity with Torula.
The technical, or scientific, description of the yeast Fungi
is " Unicellular plants, which multiply themselves by budding,
and reproduce themselves by endogenous spores. They live
singly or united in bud colonies, chiefly in saccharine solutions,
where they excite alcoholic fermentation." For the purpose
of illustration, the yeast which causes fermentation in beer
may be taken as a type of these organisms. Primarily they
consist of a single cell, which is round or elliptic, but
occasionally becomes elongated, and parted off by transverse
divisions. In order to multiply themselves, the simple cells
produce an outgrowth from the periphery, which gradually
enlarges, absorbs a portion of the
contents of the parent cell — ■ t\> ^ i) ^Y^
which it ultimately resembles in '^ ^[V\ r\
form and size — then the con- J^ ^ cQ fV- v--, VJ
uection between them is cut off ^ 0 ^ Q
by a transverse wall or partition, •x-c^^ x^-^-s^
and two cells occupy the place Fig. \2,i.—Saccharomyces ellipsoideus.
,. , 1 ,. -T71 1 p Chatto and Wiudus.
01 the lormer one. Each 01
these cells is capable, in like manner, of budding and producing
daughter cells, and so the course goes on (Fig. 138).
Increase by budding, or gemmation, goes on most rapidly
under the influence of moisture, mostly immersed in a
saccharine solution, and in this position the sugar of the fluid
is decomposed, resulting in alcoholic fermentation. Spores
may be developed on a moist substratum, by the contents of a
cell dividing itself into two or four portions, each of which
300 INTRODUCTION TO THE STUDY OF FUNGI
surrounds itself with a proper membrane, and becomes a spore,
\vhicli is capable of budding, like the vegetative cells.
AVhether the so-called species of Saccharomyces are autono-
mous, or only stages in the development of some higher forms,
need not be discussed here. " Brefeld considers that the
conidia of A-arious species of Ustilagineae exactly resemble in
mode of growth many of the forms of the so-called Saccharomyces.
It is well known that the spores of the smuts, in germinating,
protrude a thread, from which spring tufts, or clusters, of
sporules ; these unite with one another by short transverse
processes, and then give rise to sporules, or conidia of the
third generation, and these to even a fourth kind. Brefeld's
theory is that these successive generations of conidia do not
merely resemble Saccharomyces, but are identical with them.
He cultivated the spores of many Ustilagineae in nutrient
fluids, and found that the conidia to which they gave rise were
in form and dimensions similar to those of the various species
of yeast Fungi — those of one being ovate, of another oblong-
ovate, of another fusiform, of another cylindrical, of another
small and roundish, and of another filiform, and so on. More-
over, he cultivated these sporules in suitable media for numerous
generations, and found that they reproduced themselves, so long
as^the conditions remained unaltered, with unfailing certainty
the whole year through. A pair of smut spores was in-
duced to germinate, and the conidia which they produced
were transported, with due precautions, into a drop of
nutrient fluid, in which they continued to bud till the nutri-
ment was exhausted. A few of these were then removed to
another drop of the same fluid, and the process was continued
for nearly thirty times, extending over a space of twelve
months. The author considers that he has thus proved that
these conidia can propagate themselves indefinitely by budding,
just like the cells of Saccharomyces, and he asks — If we had
commenced this series of cultivations, not with the smut spores,
but with the conidia which arise from them, should we have
been able to distinguish their mode of growth from that of
the yeast of beer ? " ^
^ Syncqisis of the Bacteria ami Yeast Fungi, by W. B. Grove. B.A. (1884),
p. 81.
SCHIZOMYCETES AND SACCHAROMYCETES 301
All saccharine fluids which contain glucose, or grape sugar,
or a sugar which can be changed into glucose, and also
all nitrogenous substances, phosphates, and ammoniacal salts,
produce alcohol at a given temperature. The process of con-
version is by fermentation. Pasteur states that every
fermentation has its specific ferment ; in all fermentations in
which the presence of an organised ferment has been ascer-
tained that ferment is necessary. This minute being produces
the transformation which constitutes fermentation, by breath-
ing the oxygen of the substance to be fermented, or by
appropriating for an
instant the whole _Q q^^ ^ ^V)
substance, then de- 0 C5 © @^ ^ Q-^y rp
stroying it, by what ^<||) © ^ cC^p^^^ ^>' ^b
may be termed the ^ ^J O
secretion of the fer- ^ ,„„ ,,„. , ,„ ,,, ^, , ,„. ,
Fig. 139. — ' High yeast. Chatto and Wmdus.
mented products.
Three things are necessary for the development of the
ferment — nitrogen in a soluble condition, phosphoric acid,
and a hydrocarbon capable of fermentation, such as grape
sugar. The common ferment of wine has elliptical cells, but
there are other forms, or species, which are capable of pro-
ducing fermentation in wine. The yeast of beer has round or
oval cells,^ and so on, through the range of species, of which
Saccardo enumerates thirty-one (Fig. 139).
It was contended at one time that these ferments were
derived from moulds and Mucors, which under favourable
conditions continued to increase themselves by budding — viz.
simple vegetation — but, if deprived of nutrition, produced the
fructification of a mould. De Bary, whilst controverting this,
suggests that some yeast cells have probably been mixed with
the spores sown in a nutritive fluid. He thus describes the
development of yeast, beyond the ordinary vegetation in a
fermentable solution : " If we bring living cells of yeast out of
the fluid, on the moist surface of a succulent part of a plant —
for example, a piece of carrot — the sprouting goes on slowly for
some time, and entirely ceases after some days. About the
sixth day, we remark how some of the cells wither and others
1 Microbes, Ferments, ami Moulds, by E. L. Trouessart, London, 1889.
302 INTRODUCTION TO THE STUDY OF FUNGI
become larger ; the greater part of the latter form spores in
their inner space through the free formation of cells, like those
of an asciis, and then becoming thicker at the cost of the
protoplasm, at last entirely fill the membrane of the utricle.
We can produce the same phenomenon if we thoroughly wash
fresh yeast, and, mixing a little clear water with it, let it stand.
The formation of the spores here follows, by a sufficient supply
of water, at the cost of the organic substance, which has
assimilated during the fermentation ; we must seek it in the
yeast which is used technically when, after its fermentation is
complete, it is laid aside clear and wet. The spores begin,
when they are brought into a suitable liquid, to sprout like
the vegetating cells, in order to produce new repeated genera-
tions of the latter. No other forms of development are known
for the Fungus which is found in yeast." ^
The yeast Fungus is the principal promoter of the
alcoholic fermentation which appears in practical life, especi-
ally the greater part of beer and spirit fermentation. That
which is distinguished by the name of harm, and the yeast
deposited at the bottom of the cask, are in many cases — not in
all — the same Fungi, which in a lower temperature remains at
the bottom, and collects as under-yeast ; by higher temperature
it accumulates in the froth on the surface of the fluid, and is
called harm. There is a slight difference in the form of the
yeasts, but the one form can be transferred to the other, by
changing the temperature of the fermentation. Further
details will have to be sought in some work dealing specially
with the subject, as we are only interested in furnishing an
outline of the organisms concerned in the processes of putre-
faction and fermentation.
BIBLIOGRAPHY
Saccardo, p. a. " Saccharomycetaceae," by J. B. de Toui, and "' Scliizoniy-
cetaceae," by De Toni et V. Trevisan, in Syllogc Fungurum, vol. viii. Padua,
1889.
TiiouEssAiiT, E. L. Microbes, Ferments, and Moulds. Cuts. Sni. Svo. London,
1889.
Grove, W. B. A Synopsis of Bacteria and Yeast Fungi. Sm. Svo. Cuts.
London, 1884.
1 De Bary On Mildew and Fermentation, Berlin (1872), p. 61.
SCHIZOMYCETES AND SACCHAROMYCETES 303
WooDHEAD, G. S. Bacteria and their Prod^ids. Sm. 8vo. Cuts. London, 1891.
and Hare, A. W. Pathological My cologie. Roy. Svo. Cuts. Edinburgh,
1885.
De Bary, a. Morphologic iiiid Physiologic der Pilze, etc. 8vo. Leipzig, 1866.
Lectures on Bacteria, translated into English. Svo. London, 1887.
Pasteur, iltudcs sur le Vin (1866). £tudcs sur Ic Vinaigre (1868). Etudes
sur la Biere (1876). Paris.
Schutzenberger, Pruf. Fermentation. Sm. Svo. Cuts. London.
Bastian, C. H. The modes of Origin of Lowest Organisms. Svo. London, 1871.
The Beginnings of Life. 2 vols. Post Svo. London, 1872.
CiENKOWSKi, L. " Zur Morphologie der Bacterien." Monoirs Acad., x's.y. St.
Petersburg, 1877.
Reess, M. Bot. Untersuch ii. d. Alkuholgahrungsinlzc. Leipzig, 1870.
MoRiNi, F. Gli Schizomiceti. Milan, 1883.
Alcune Considcrazioni sugli Schizomiceti. Milan, 1882.
Crookshank, E. M. Manual of Bacteriology. Svo. Col. plates. London, 1886.
ScHENK, S. L. Elements of Bacteriology. English Translation. Roy. Svo.
London, 1893.
CHAPTEE XXV
SLIME FUNGI :\rYXOMYCETES
The Myxomycetes, or Myxogasters, are an extraordinary group,
which have been the subject of much discussion, on account of
some peculiar features which characterise them, and separate
them from Fungi generally, and all other of the Cryptogamia.
On this account some have advocated their exclusion from the
vegetable kingdom altogether, whilst zoologists have been in
no hurry to accept them. The common error of accepting
analogy for affinity is one which even scientific minds are
occasionally betrayed into committing, and yet, apparently,
unconscious of their own failing. We liave, during a period
of half a century, seen several hypotheses started on similar
unstable bases, flourish awhile, and then come to nought. For
a long time, and up to a very recent date, the Myxogasters
were classed with the Trichogasters, as two groups of the order
of Gasteromycetes. Without a knowledge of their life-history,
and but little of their microscopical structure, this assumed
alliance was a natural one, but it has come to be renounced.
It was only in 1864 that the position was assailed by De Bary,
who changed the name to Mycetozoa, and claimed for them a
position as nearly related to the animal as the vegetable world.
" I have," he says, " placed the Myxomycetes, under the name
of Mycetozoa, outside the limits of the vegetable kingdom, and
I still consider this to be their true position." Strangely
enough, however, in all his subsequent botanical works, he
continues to include the Mycetozoa, as if he lacked the courage
of his opinions; and other botanical writers and compilers of
text-books have continued the same course. In this group the
two stages or phases of life, the vegetative and the reproductive.
SLIME FUNGI— MYXOMYCETES 305
are sharply defined and distinct. It is in the vegetative stage
that all the supposed affinities with the animal world are
encountered, and in the reproductive everything is suggestive
of Fungi, even to the terminology which is borrowed from, and
represents identical structure with what is familiar in Gas-
tromycetes. The outer wall is either a sporangium or peridium,
the threads of the interior still compose a capillitium, the con-
tinuation of the stem into the interior is a columella, and the
reproductive units are not ova, but the spores. This is accounted
for by De Bary from the " close agreement in structure and in
biological characters between their organs of reproduction and
the spores of Fungi." As Mr. Massee has lately pointed out,
it is clear that De Bary derived all his reasons and his evidence
against the vegetable nature of the Myxomycetes from the early,
or vegetative, phase. On the other hand, it seems to have been
suggested that in the later, or reproductive, phase the disparity is
so great between the structure and biological characters of the
Mycetozoa and those of any of the lower animals, that he was
compelled to use the terms, in describing them, which belong
also to the Gastromycetes. Here, then, we are supposed to
come face to face with a problem — certain organisms in their
early, or vegetative, stage belonging to the animal kingdom, and
subsequently in their final, or reproductive, stage undoubtedly
vegetable, — a worse example of a dual-hypothesis than that
which combines an Alga with a Fungus to produce a Lichen.
Taking away all expletives, and reducing the indictment
to its simplest form, it remains as a specific reason that " the
characteristic mark of separation lies in the formation of Plas-
modia, or aggregation of swarm-cells." In his recent monograph
of this group,^ Mr. Massee has faced and combated the position
occupied by De Bary, step by step. " In the Myxomycetes,"
he says, " the spores on germination give origin to one, two, or
more naked cells, which possess the power of movement, due
to the protrusion of pseudopodia, or the presence of a cilium ;
these cells are known as swarm-cells. The swarm-cells possess
a nucleus, multiply by bi-partition, and eventually coalesce to
form a plasmodium in the following manner. After the pro-
duction of numerous swarm-spores by repeated bi-partition, little
^ Massee, Monograph of the Myxogastres, London (1892), p. 5.
20
3o6 INTRODUCTION TO THE STUDY OF FUNGI
groups are formed by the close approach of two or more of
these bodies ; these groups often disperse again, but eventually
the components of a group coalesce, and lose their individuality ;
this coalescence and loss of individuality results in the forma-
tion of a small plasmodium, which, in some unknown way,
possesses the power of attracting surrounding free swarm-cells ;
these at once coalesce and add to the bulk of the plasmodium.
The nuclei of the component swarm -cells retain their in-
dividuality in the plasmodium, the latter retaining the power
of motion originally possessed by its components, and represent
the vegetative phase of a Myxogaster. Under certain con-
ditions, unfavourable for active vegetative work, plasmodia
possess the power of passing into a temporary sclerotioid, or
resting stage ; the preliminaries for this condition are the
breaking up of the protoplasm into innumerable roundish or
polyhedric cells. In some species the cells become surrounded
by a distinct, colourless membrane, which shows the reaction
of cellulose."
From the above account we learn that the coalescence of
naked motile cells, or even the aggregation of naked motile
cells without loss of individuality, is, from De Bary's standpoint,
the proof that the Myxogasters are not plants.
After comparison of these phenomena with similar analogous
instances in the Phycomycetes and other Fungi, the following
reasons are adduced in support of the vegetable, rather than
animal, nature of these organisms : —
(1) Frequent presence of cellulose in the general membrane
protecting plasmodia, cell-walls of spores, sporangia, and walls
enclosing the protoplasm in the sclerotioid, or resting stage of
plasmodia. (2) Presence of germ -pores in the cell- walls of the
spores, of some species. (3) The frequent separation of lime
from the protoplasm at the commencement of the reproductive
phase. (4) The frequent separation of a substance from the
protoplasm during the period of spore -formation, homologous
with the substance separated during the same period in the
Ascomycetes, etc. This substance in the Myxogasters forms
the capillitium. (5) The agreement with many Fungi in the
contrivance for spore dissemination. (6) The production by
free cell-formation of spores protected in the early stage with
SLIME FUNGI— MYXOMYCETES 307
a wall of cellulose, which eventually becomes differentiated, and,
as stated by De Bary, " behaves towards reagents in a similar
manner to cuticularised plant cell-membranes, and to spore-
membranes as in the Fungi. (7) Presenting analogy with
undoubted members of the vegetable kingdom, as Hydrodictyon,
where the naked motile swarm-cells coalesce to form a caenobium,
which eventually becomes invested with a membrane. (8) In
the close affinity with Ceratium (but, as Ceratium has been
included by some with Myxogasters, this will not carry so much
weight). (9) In the coalescence of the naked cells to form a
Plasmodium, being the result of conjugation between the com-
ponent cells, thus presenting features in common with the
primitive forms included in the group Zygosporeae."
It may be added that, in this country, Mr. Saville Kent,
as a zoologist, espoused De Bary's views, and even went beyond
him, in his Manual of Infusoria, for he included the
Mycetozoa, and suggested their affinity with sponges. These
views were contested at the time,^ but really no fresh evidence
was produced in support of the views of De Bary, who was the
great authority cited.
The only addition necessary to quote, or allude to, in
support of the animal nature of the Myxomycetes, in the
vegetative stage, is the evidence of Mr. Lister ; but these obser-
vations extend no further than the vegetative stage, and do
not furnish any convincing proof that the phenomena are in-
compatible with a condition of vegetable organisms, any more
than the amoeboid forms in such Algae as the Volvocineae.
" I have repeatedly seen bacteria taken by swarm-cells of
Chondrioderma difforme in the manner described, and it would
appear that bacteria form their principal food. On one
occasion I had a favourable opportunity for observing the
digestion of bacilli on account of the quiescent state assumed
by a swarm-cell, which remained with little active movement
for an hour and a half On the previous evening I had placed
some spores of Chondrioderma difforme in water, under a thin
cover -slip; on the following morning swarm-cells were in
great abundance in the pure water. I introduced a drop con-
taining multitudes of bacilli from a glass in which a piece of
^ "Animal Nature of Myxomycetes," in Grcvillca, vol, ix. (Dec. ISSO), p. 41.
3oS INTRODUCTION TO THE STUDY OF FUNGI
Stereum hirsutum had been soaking for several days. In a
short time a number of the swarm-cells were seen, attended by
bacilli, some of which were attached to their pseudopodia, and
some were already enclosed in vacuoles. The swarm-cell in
question had taken an amoeboid form, occasionally producing
and again withdrawing the cilium, while from time to time
thin pseudopodia were extended from the opposite end, but
more frequently the posterior region expanded into a
somewhat funnel-shaped mouth. Into such an expansion a
stout bacillus was seen to enter ; in the course of a few seconds
it was enclosed with a noticeable amount of water, by the
folding over of the lips of the funnel, and conveyed into the
body -substance ; a few minutes after, another bacillus was
taken in, much in the same manner, but no globule of water
was introduced. Ten minutes later a large bacillus was caught
by a prolongation of one side of the funnel, and in the course
of half a minute a tube-like extension of protoplasmic substance
invested the bacillus, and it was drawn in. It remained for a
short time in direct contact with the granular matter of the
body, but was soon surrounded by an oval vacuole. The
swarm -cell continued inactive for nearly an hour, when it
assumed an extended form, and shortly after swam away with
rapid jogging movement. Constant observation was maintained
during this hour, and the bacilli were seen gradually to dissolve
in the vacuoles in which they lay, until at length all trace of
them had disappeared, together with their containing vacuoles,
and only the contracting vacuole remained in the homogeneous
granular substance of the swarm-cell.
" At the commencement of the observation this granular
protoplasm was much more turbid than at the close, when it
was remarkably hyaline ; the swarm-cell appeared also to have
increased in size, though it was difficult to determine by
measurement in consequence of its changing form, No re-
jection of refuse matter took place while the observation lasted.
" In the same preparation I watched a swarm-cell creeping
in a straight line, with the strange snail-like movement so
difficult to understand. In its course it came to a small group
of motionless bacilli lying against the glass ; immediately it
changed its linear form and spread itself out, covering four of
SLIME FUNGI— MYXOMVCETES 309
the bacilli. In about two minutes it resumed its former shape
and movement, and crept away, carrying off two of the bacilli
in vacuoles.
" These observations seem to confirm the opinion of De Bary
that the organisms under consideration should be classed
among the animal rather than the vegetable kingdom. When
a creeping swarm-cell is watched, with the projecting cilium
placed immediately in advance of the nucleus, which never
shifts its position, and when we note the manner in which the
vibrating extremity of the cilium appeared to detect the
presence of the bacilli, before the swarm-cell spread itself over
them ; again, when we observe the
creeping action suddenly change,
and raising itself from the decum-
bent attitude, with a few lashing
strokes of the cilium the swarm-cell
releases its foothold and swims Fig. 140.— Cluster of TubuUna
away ; and when to these remarkable cyimdnca.
movements is added the process of ingestion, we cannot but
feel the force of the conclusion at which De Bary arrived, if
indeed a distinct line of demarcation between the two kingdoms
can be said to exist." ^
Nearly all the species in this grc^^p are minute, and when
not so are composite, several individuals being united in a
cluster (Fig. 140). Most of them are more or less gregarious,
and sometimes covered with a shiny envelope, of which portions
extend to the matrix, and resemble when dry a sort of mem-
branaceous thallus. The tendency is certainly towards the
globose in form, now and then attenuated into the cylindrical.
If we were to attempt a kind of typical description, we should
say that they are small globose or pear-shaped bodies, with or
without a stem, scarcely exceeding a millimetre in diameter,
variable in colour, sometimes shining and sometimes covered
with white chalky granules ; at first pulpy, then dry and
fragile, filled within with a mass of pulverulent spores, often
mixed with threads of a capillitium. Into this interior the
stem is continued as a columella, which is connected with the
1 " Xotes on Chondrioderma difforme," etc., by A. Lister, in Annals of
Botany, iv. (May 1890), p. 281.
3IO INTRODUCTION TO THE STUDY OF FUNGI
walls of the sporaugiiuii Ly the radiating threads uf the
capillitium. It will be observed how closely these details
accord with those of the Lyco'perdaceae amongst puS-balls, so
that we would seem to be describing very microscopical puff-
balls. To a certain extent this is correct, but with the ex-
ception that the early condition is slimy, there is often when
mature a thin film of dried mucilage derived from the envelope,
and the capillitium is sometimes very highly developed. We
must, however, guard against the inference that there is any
true affinity between Myxomycetes and Gastromycetes, as old
authors believed. They are mostly developed upon dead leaves,
or very rotten wood, in damp places ; and though probably
most common in temperate regions, a few species extend into
the tropics.
The total number of recorded species, included in the most
recent work on the subject, is 425 ; but no extended analysis
of their geographical distribution has been attempted, as it is
not clear that species recorded from distant stations twenty
or thirty years ago were determined with sufficient accuracy.
It is unnecessary to allude at any length to the classifica-
tion adopted for the arrangement of this group. Previous to
the Monograph by Eostafinski ^ all the species were arranged
according to external characters, determined by the aid of a
pocket lens. Although it must be admitted that Fries
exhibited a remarkable insight into the relations and affinities
of the various groups of Fungi, yet his method was insufficient
for the minute species, and the Myxogasters, amongst others,
received only inadequate treatment at his hands, so that a
revision, with the aid of the microscope, became an absolute
necessity. De Bary evidently intended to do this, but never
accomplished it, although subsequently one of his pupils, who
had the benefit of becoming acquainted with his views, pro-
duced, in Polish, the Monograph above alluded to. Ptostafinski
accepted, in name, the Mycetozoa of De Bary, but with a more
restricted application, and his classification proceeded on a
botanical basis, since it was the reproductive phase, or
completed condition, which he recognised as the individual.
■ Sluzowce, a Monograj^h of the Ilycdozoa, by Dr. Joseph Eostafinski (in
Polish), 1875.
SLIME FUNGI— MYXOMYCETES 311
Hence we find that the primary division into two sub-
divisions was based upon spore characters. In the first
section, the Amaurosporeae, the spores were violet, or brownish
violet ; and in the second subdivision, the LamjjrosjJoreae, the
spores were variously coloured, but never of any tinge of
violet. The next feature which seemed to him most im-
portant, or at least most fitting for the purpose of classifica-
tion, was the presence or absence of a capillitium. Sub-
sidiary to these two features, the presence or absence of lime
in the sporangium or capillitium, the production or suppression
of a columella, and the perforation of the walls, were employed
in the delimitation of families ; after which followed the genera,
with their varied predominant characteristics.
Subsequent writers, having Eostafinski's work as a basis,
have proposed alterations and emendations, whilst the
majority of mycologists have felt that, although it did much
to direct inquiry into a new channel, and classify on sounder
principles, the Monograph did not exhaust the subject, but
left many occasions for improvement. As we are writing this
chapter, the latest attempt at a revised classification has issued
from the press.^ In this arrangement the primary subdivision,
as to spore coloration, is abolished, and another central idea
established, which is thus explained : — " The most pronounced
feature in the evolution of the Myxogastres is in connection
with spore dissemination, and the following arrangement is
based on the relative development of the capillitium, which is
seen in its most perfect form in the genera Trichia and
Arcyria."
The entire group is subdivided into four orders, in the
following sequence: — (1) Wall of sporangium without lime;
capillitium absent, or formed from the wall of the sporangium.
(2) Wall of sporangium still without lime ; capillitium
originating from a central columella. (3) Wall of sporangium
with an external deposit of lime ; capillitium present. (4)
Wall of sporangium without external deposit of lime ; capil-
litium present, but not springing from a columella. In the
introduction the above is the sequence of orders, but in the
1 A Monograph of the Myxogastres, by G. Massee, Loudon, 1892 : and
subsequently that by A. Lister in 1895.
312 INTRODUCTION TO THE STUDY OF FUNGI
subsequent elaboration tbe last two orders are transposed.
For the most part the genera are the same as in Eostafinski,
with the exception of two or three instances in which con-
tiguous genera are amalgamated.
It must be expected that an evolutionist, such as Mr.
Massee confesses himself to be, would have decided ideas as
to the evolution of this group. " I consider," he says, " the
Myxogastres as illustrating one of the earliest known attempts
at differentiation in the direction that has eventually resulted
in the mass of organisms constituting the vegetable kingdom ;
but having originated from the Flarjellatae, a group more in
touch with the animal side of life, the work of developing
individuality has been slow, as illustrated by the tardy
appearance of cellulose cell-walls, which, as would be expected,
is most complete in the newly evolved reproductive phase,
itself to a great extent the outcome of a gradual change of
environment from aquatic to aerial ; but the radical mistake,
after having adopted the plant line of development, consisted
in the non-development of chromatophores, and retention of
the animal mode of nutrition, which in the plant world means
parasite, or sapr ophite. The fungi, a later group, differentiated
from ancestors that had already evolved the leading plant
characteristics, including cell-walls, chlorophyll, starch, hence
in this respect are more typical plants than the Myxogastres ;
but in the fungi, the check to progress was due to the
degeneration of the chromatophores, already evolved by their
ancestors, whereas, in the Myxogastres, the check was due to
their inability to differentiate these essentials."
As for ourselves, we are by no means disposed to dogmatise
on any speculations of this kind, which seem to have such a
slight basis of solid fact, and permit such a free scope to
inference. Neither are we content to exclude Myxomycetes
from Fungi, as the above quotation suggests, since their
strongest affinities when mature appear to be with Fungi ; but
we confess to a predilection for regarding them as a peculiar
and aberrant group, which, by reason of their vegetative phase,
do not fall well into place with our present arrangement of
Fungi.
It is incumbent upon us to append a brief synopsis of the
SLIME FUNGI— M YXOM YCE TES
;i3
classification which has been adoi^tecl for these singular
organisms, the characters for which are derived from the
final and reproductive condition.
The first of the four orders, into which the entire group is
subdivided, is the Peritrichiaceae, in which the wall of the
sporangium is not encrusted with lime, and the capillitium is
either absent or formed from the wall of the sporangium.
This order is again subdivided into two suborders — that of
the Tululinae, in which the wall of the sporangium is not
perforated ; and the Crihrariae, in which the wall of the
sporangium is perforated.
The principal genus in the Tubulinae is that of Tuhulina,
in which the sporangia are crowded
together so as to form an aethalium,
which term is applied to an ag-
glomeration of sporangia. The Crih-
rariae includes the genera Enteridium,
Clathroptychium, Cribraria, and Dic-
tydium, in all of which the perforated
sporangia are very elegant objects.
The subsidiary characteristics of the
several genera have reference chiefly
to the manner of the perforations.
In Cribraria the permanent upper
portion of the sporangium forms a
Fig. 141. — Cribraria intricata.
kind of network (Fig. 141), and in
Dictydium the permanent radiating
ribs are united by transverse bars
(Fig. 142).
The second order, ColumelUferae,
with the walls of the sporangium
not containing lime, has for its chief
character a central columella, from
which the capillitium originates. The
two suborders into which this group
is divided are the Stemoniteae, in
which the capillitium springs from
every part of an elongated columella,
and the Zamprodermeae, in which the capillitium springs from
Fig. 142. — Dictydium. natural
size and magnified.
314
INTRODUCTION TO THE STUDY OF FUNGI
the upper portion
Fig. 143.— ,S<ewi'.
fusca.
of the columella. lu the Stemoniteae the
typical genus is Stemonitis (Fig. 143), in
which the sporangia are free, whilst in the
other genera, as Amaurochaete, Brefeldia,
and Eeticularia, the sporangia are combined
into an aethalium. In the other section,
called Lamprodermeae, there are some half
dozen genera, in all of which the sporangia
are free, but in the most numerous and
typical genus, Lamproderma, the threads
of the capillitium arise from the abrupt apex
of a short columella.
The third order is the Zithodermeae,
which includes a great number of species,
and is subdivided into two sections, in both
of which there is an external deposit of lime
on the wall of the sporangium. The
Didymeae have a capillitium which is
wholly without lime (Fig. 144), and the
Physareae a capillitium which encloses lime.
We need not stay to analyse the different
genera in these two sections, inasmuch as
they will offer no difficulty to the student.
The Didymeae includes such genera
Didymium, Lepidoderma, Spumaria, and
145). In
as Chondrioderma,
Diachaea (Fig.
Spumaria only are the spor-
angia combined in an aethal-
ium, in each of the other
genera they are free. The
typical genus, Didymium, has
the sporangium encrusted with
a powdery coating of lime, and
the distinguishing feature of
the three other genera consists in the coating of the sporangia.
The Physareae include eight genera, only one of which, Ftdigo,
has the sporangia combined into an aethalium, as a generic
character, although in other genera some of the species
may form an aethalium. Physaruin is a large genus in which
Didymiuvi farinaceum.
SLIME FUNGI^MYXOMYCETES
315
ym
the capillitium is much swollen at the nodes, enclosing lime ;
whilst Tilmadoclie has small nodes containing
lime, and Badhamia has thick threads in the
capillitium, containing lime throughout. In Cra-
terivm the form of the sporangium more or less
resembles a wine-glass, closed by a lid or operculum
(Fig. 146). The re-
maining genera are
small, consisting of a
single species in each.
The last of the four
orders is the Calotricheae,
in which the capillitium
is for the most part
highly developed, and
the sporangia have no
external deposit of lime. Fig. 145.—
The two subdivisions are ^yj^-ij capii-
Tricheae and Arcyriae ; in the former the threads of ^itium aud
the capillitium are free, and do not anastomose, exposed.'
whilst in the latter they are attached by one end,
or combined into a network. The Tricheae include two genera,
11
Fig. 146. — Cmtcrium, natural size
and masnilied.
the principal being Trichia, in which the threads are spiral
Fig. 147.— Threads and
spores of Trichia.
Fig. 148. — Arcyria, with portion of
capillitium magnified.
(Fig. 147), and Oligonema, in which there are no distinct spirals.
The seven genera of the Arcyriae are partly known by the char-
acter of the capillitium, of which the largest genus is Arcyria.
having the threads combined into a network which becomes
naked or protruded at maturity (Fig. 148). Two other genera,
3i6 INTRODUCTION TO THE STUDY OF FUNGI
such as Lycogala and Perichaena, have elementary threads,
and the remainder are of secondary importance. This is,
briefly, the basis of classification in the Myxomycetes, and
is dependent, as in other groups, upon the full and mature
development of the individuals for their identification.
BIBLIOGRAPHY
RosTAFiNSKi, J. Sluzoicce, Mycetozoa Monografia. Plates. 4to. Paris, 1875.
Massee, G. A Monograph of the Myxogastres. Col. plates. Roy. 8vo. London,
1S92.
"A Revision of the Trieliiaceae," \n Journ. Royal Micr. Soc. London,
1S89.
Cooke, M. C. The Myxomycetes of Great Britain. Plates. 8vo. London, 1877.
The Myxomycetes of tlie United States. Roy. 8vo. New York, 1877.
Lister, A. " Monograph of the Mycetozoa. " 8vo. 78 plates and cuts. London,
1894. {British Museum Catalogue.)
De Bary, A. " Die Mycetozoon " (.Sc/i^eiwy/j/^ie). Plates. 8vo. Leipzig, 1864.
CiENKOWKSi, L. "Zitr Entwickelungsgeschichte der Myxomyceten." Prings-
heim Jahrb. , iii.
Stahl, E. "Zur Biologic der Myxomyceten." Bot. Zeit. 1884.
Saccardo, p. a. "Myxomyceteae." Sylloge Fungorum, vol. vii. pt. i. 1888.
Blytt, a. Norges Myxomyceter. Christiania, 1892.
Raciborski, M. Sluzowce. Cracow, 1884.
Bemerkungen iiher einigc heschrcib. Myxomyceten. Dresden, 1887.
Raunkier, C. Myxomycetes Daniae. Copenhagen, 1888 ; in English, 1889.
PAKT III
DISTRIBUTION
CHAPTEE XXVI
CENSUS OF FUNGI
The estimated number of species in any department of natural
history, at any given time, is of passing interest, although
necessarily it is always changing, and must, to a great extent,
be only an estimate. It is a very long time since any estimate
of the number of described species of Fungi could have been
made upon an equally satisfactory basis to the present. This
is due to the recent publication by Professor Saccardo of a
Sylloge, which was presumed to contain an enumeration of
all species described up to date, and this Sylloge must there-
fore be taken as the basis of our calculations.
The last previous attempt at a full enumeration of species
was that of Streinz' Nomendator, dated 1862, in which the
total number of species was 11,893; and besides that we had
only vague estimates to guide us, such as that expressed by
De Bary in 1872, when he said, " It is no exaggerated estimate,
if we place the number of the species of living Fungi on an
equality with that of the floriferous plants, viz. about 150,000."
Probably his intention was not to include merely the described
species, which had been discovered, but to estimate the entire
number of species, known or unknown, which might be in
existence on the surface of the globe. After all, such an
estimate could only have the value of an individual opinion.
An estimate which we ventured to give in about 1872 placed
the number of known species at 20,000 ; whilst, some fifteen
years afterwards, we intimated an opinion that they must
approach to nearly double that number; whereas a clear total,
according to Saccardo, on a determinate basis, is 40,000 up to
1892. It is interesting to revert to the opinions and estimates
320 INTRODUCTION TO THE STUDY OF FUNGI
of some still earlier writers than we have mentioned, Hum-
boldt, for instance, three-quarters of a century ago wrote : " If
we estimate the whole number of the Cryptogamia hitherto
described at 19,000 species, as has been done by Dr. Klotsch,
a naturalist possessing a profound acquaintance with the agamic
plants, we shall have for the Fungi 8000 (of which the
Agarics constitute the eighth part)." It is a remarkable co-
incidence that in Saccardo's enumeration the Agarics still
constitute the eighth part of the whole. As to the Hymeno-
mycetes, which include the Agarics, Fries, in his Hipncnomycctes
Euroimci, gives a total number of species for the whole of
Europe as 2778. Before the publication of the Sijllorje, we
remarked on this fact : " It may fairly be concluded that the
total number of species of the Hymenomycetes is not less than
5000." Subsequently the Syllogc extended that number to
9634, the proportion of which that are confined to Europe we
have not ascertained, but it is considerably beyond that
enumerated by Fries. The total of species of Hymenomycetal
Fungi now known reaches to nearly one-fourth of the total
of described species of Fungi. If we analyse these results still
further, we find that of the Hymenomycetes not less than 5245
belong to the Agaricini, or gill-bearing series, and 2200 to the
Polyporei, or pore-bearing series, leaving only rather more than
2000, or about equal to the whole of the Polyporei, for the
remainder of the Hymenomycetes, i.e. the Hydnei, Thelephorei,
Clavariei, and the Tremellini.
The next group of importance, as to number, is that of the
Pyrcnomycetes, formerly termed the Sphacriaceae, in which the
spores are contained in asci and enclosed in a perithecium.
Placing the total at 10,500, we see at once that it is more
numerous in species than the whole of the Hymenomycetes,
and more than one-fourth the total of all known Fungi. It
must be remembered that the largest perithecium known is
not much larger than a grain of mustard seed, or, at any rate,
not so large as the seed of a vetch, although in some compound
species, in which some hundreds of perithecia are collected
in a single stroma, that stroma may attain the size of a
man's fist. Hitherto the number of British species has always
been less than the total of British Hymenomycetes, perhaps
CENSUS OF FUNGI 321
not more than three-fourths, but I'ecently the dih'erence has
diminished.
Closely allied to the above are the Discomycetcs, with the
sporidia also enclosed in asci, but with the fertile disc exposed.
The species enumerated are about 3800, with which there are
no previous lists for comparison, and no estimate, save that of
twenty years ago, when we estimated the total number at some-
thing like 2000 species. The Systema of Fries only contains
about 430 species for 1822 ; whilst for British Discomycetes
alone, Phillips, in 1887, records 607 species. A large
majority of the species are fleshy, and hence almost confined to
temperate regions, but it is only during recent years that they
have been studied seriously and effectively. Until about
twenty-five years ago they were absolutely neglected, and we
are indebted chiefly to Fuckel, Nylander, and Karsten for indi-
cating the lines upon which future studies should be pursued.
The remaining groups are comparatively small, and none
are of more importance than the Gastromycetes, which are of
considerable size, so as not to be easily overlooked, and dis-
tributed over a wide geographical range ; yet the number of
species has hardly increased in proportion to those in other
groups. The present total number of species, of all kinds,
does not exceed 720, and of these no less than 173 are
represented in Australia, which seems to be the happy land
for the Gastromycetes, not half that number being found in
Britain.
The Hypodermei include the two smaller groups which are
better known as the Uredinei and the Ustilaginei, to both of
which large additions have been made in recent years, not so
manifest in a catalogue on account of the union of the members
of several so-called genera under one designation. Some of
the present species are by no means stable, which go to make
up the total of 1750, The Ustilaginei only number about
320 of these, leaving 1430 species for the Uredinei. The
total number of British species, according to the latest mono-
graph, was 261. The previous catalogue for 1878 included
293 species, but this is accounted for by the new arrangement
placing the Aecidium, Uredo, and Teleutospores under one
generic denomination. This makes it difficult to compare
21
322 INTRODUCTION TO THE STUDY OF FUNGI
recent lists with old ones, although practically a considerable
increase is inevitable.
The Phjjcomiicdes are interpreted now in a broader sense
than they were a few years ago, which renders comparison
with the older authors ditticult ; nevertheless we must accept
the 686 species which are included in the total of the Sylloge.
We have still upwards of 10,000 species to deal w1!lh,
which belong to the imperfect Fungi, and these include 6865
which are classed under the Spliaeropsidcae and Melanconiae ;
and 4760 moulds or Hypliomycetes. Although these are sus-
pected to represent imperfect states of other Fungi, they must
retain a place as species until their affinities are determined.
Undoubtedly the number of these form-species has increased
enormously of late years ; they have always held a subsidiary
place in the estimation of mycologists. Certainly a combined
total of not less than 1400 species may be set down as British,
against 489, the total number recorded in the Handlool- for
1871, or nearly treble within about twenty years.
Whatever position posterity may assign to the Microbes,
they cannot be excluded from our census. Hence we have a
record of no less than 689 species of Saccharomycetes and
Schizomycetes, otherwise known as yeast Fungi and Bacteria,
which are absolutely the growth of the past few years. There
are some who are ready to contend that the bacteria are all, or
nearly all, simply the modifications of a single species ; but
there is such a thing as rushing to extremes, so that whilst we
may cherish the belief that more experience and closer observa-
tion will tend rather to diminish than increase the number,
we must accept the total as it stands. There are no standards
for comparison that are twenty years old, and in Britain the
first attempt to construct a synopsis was not made until 1884,
and even in this the indigenous species are not indicated.
Finally, the slime Fungi, or Myxomycetes, have to be
included, for we do not hesitate to regard them as more
closely related to Fungi than to anything else, and of these
there are 450 species. They do not increase so rapidly in
numbers as some other groups, and we have a very good
standard of comparison in Eostafinski's Monograph of 1875,
which enumerated 178 species. Of these no less than 100
CENSUS OF FUNGI
323
were ideiititied as British in 1877, and that number was
increased to 144 by Massee in his 3Ionograph of 1892.
There are one or two smaller groups which could scarcely be
included in any of the principal groups, but they do not
altogether exceed more than about some 200 species.
From the foregoing, then, we gather the following con-
clusions— that the total numljer of described species of Fungi
to 1892 was about 40,000.
Of the H_ymenomycetes we accept a total of
For the Pyrenomycetes, or Sphaeriaceae
To these add for the Discoraycetes
And for the Gastroinycetes
The Hypodermei, or Rust and Smut Fungi
The Phyconiycetes in its broadest sense
The Sphaeropsideae and ]\Ielanconiaceae
The Hyphomycetes, or Moulds
The Saccharomycetes and Scllizomycetes
The Myxomycetes, or Slime Fungi
Tuberaceae and others not specialised
9,634
10,500
3,800
720
1,750
686
6,865
4,760
689
450
145
CHAPTEE XXVII
GEOGRAPHICAL DISTRIBUTION
The facts from which a satisfactory account of the distribution
of Fungi over the world could be constructed are, even now,
too fragmentary for the purpose. For the more civilised and
best known countries there is not much difficulty, but there
are still immense tracts over which no mycologist has ever
passed, and for which no catalogue of species is known. When
we attempted a survey of this kind twenty years ago, we were
perfectly conscious of this difficulty, and in that interval very
few of the difficulties have been removed. Although the
materials are more complete than they have ever been for
generalisation in respect to well -explored countries, it is un-
fortunately true that very few of the countries then imperfectly
known, or wholly unknown, in this respect are in a better
position now than they were then. Even in Europe we are
still compelled to confess ignorance, almost as great as it was
then, of the whole of European Turkey, a great part of Eussia,
and the Spanish Peninsula. And this forms the stronger
contrast on account of the better development of our knowledge
respecting the remaining countries. In the northern parts of
the New World there has been continued activity, excepting in
those parts which are under British rule, where no progress
has been made. Of all the vast continent of Asia we are
nearly as ignorant as we were a quarter of a century ago.
For Japan there is a prospect of a better future through the
exertions of a few intelligent natives who are cultivating this
branch of botany, but China is still an unknown land, and the
accessions to our knowledge of British India, in its broadest
sense, are but few and far between. The islands are still
GEOGRAPHICAL DISTRIBUTION 325
almost in the same position as they were. The southern
hemisphere exhibits some improvement, but this is principally
around old centres. In South America activity has been con-
fined chiefly to the eastern side, south of 20°, and for about
twenty degrees southward, but beyond that all is silence. The
hopes that the Dark Continent, which has evinced so much
vitality in other directions, would furnish good botanical
records have not been fulfilled, and even the temporary activity
at the Cape has subsided into stagnation. From our point of
view the whole of Africa is nearly as it was in 1874. The
colonies of Australia have, nevertheless, added much to our
knowledge, through the efforts of a few local botanists, and
acquired the distinction of possessing a combined Flora of their
own, for the Fungi of five of the colonies. Other islands of
the Pacific are much as they were, and for the rest of the
world we can recognise no alteration, except perhaps some
additions to our knowledge of parts of Northern Asia, and a
little more of Egypt.
Even in our own country we are conscious that Fungi are
more erratic in their appearance and disappearance than
flowering plants, and even than other cryptogams. It is in
the experience of every one that a species, or even an entire
genus, which is common in one year becomes scarce in the
next ; or that a comparatively common species may gradually
become rare in certain localities, through a series of years, and
at length vanish altogether. General conditions of temperature,
or humidity, affect the appearance of fleshy Fungi much more
than it does that of any other plants, and sometimes it is im-
possible to account for the fluctuation. For instance, in 1893
there were generally more of the common mushroom to be
found in England than in any period during the previous
thirty years, and yet all other Agarics were remarkably scarce.
The fleshy Hymenomycetal Fungi, of which the mushroom
is the type, belong almost exclusively to temperate regions ; as
warmer countries are approached, they are only found at high
elevations, whilst their representatives near the sea level
belong to genera in which the substance is tough and leathery,
and the proportion of water in their composition is compara-
tively small. Hence we find that nearly all the Fungi of the
326 INTRODUCTION TO THE STUDY OF FUNGI
Agaric type to be met with constantly in the tropics belong to
such genera as Marasmius, Schizopliyllum, Lentinus, and the
almost woody Lenzites. As we approach the cold polar regions,
lieshy Fungi gradually disappear in the face of frost and snow.
If we accept the number of gill-bearing Fungi as 5200
species, we shall find that those genera in which they are
tough and elastic, rather than brittle and fleshy, contain about
800 of that number, and all of these have white spores. So
that not more than one-seventh of the total number of gill-
bearing Fungi can be regarded, generally, as capable of support-
ing a tropical climate. Then, again, of this number of 800 a
certain proportion will be found in temperate regions, not less
than 320 of this total being recorded, so as to leave only 480
as exclusively tropical or subtropical amongst the Tenaces
genera of Agaricini. But to these must be added 550 white-
spored species, of the fleshy kind, that have at some time
or other been recorded for some tropical locality, including all
those which may have been found at a great elevation, and
consequently in a temperate region; and, finally, 450 species
with coloured spores; making a total of 1480 species
which have been found in tropical or subtropical countries.
It must, however, be remembered that of these 1000 species
of Agarics, of the more fleshy kind, which have been found in
the tropics, a great many of them are really species which
belong to a temperate zone, and it would be difficult to
estimate how many of them liave been found only at a con-
siderable elevation, as on the slopes of the Himalayas and the
Andes. Whilst a proportionately large number of species of
Lepiota have been found in warm countries, as in Ceylon, it is
remarkable that of Cortinarius, Bicssula, and Ladarius, which
number some 626 species, only 12 have been met with in
tropical regions.
It may fairly be estimated that not less than 4000, but
possibly more, species of Agaricini have been recorded in
temperate climates. By far the largest number of these
belong, either exclusively or conjointly, to Europe and North
America. Of these 2800 belong to Europe, many of them
extending into the United States, whilst 505 are found in the
United States which do not occur in Europe, which leaves only
GEOGRAPHICAL DISTRIBUTION 327
700 species to be distributed over all other temperate regions
of the world. Thus we arrive at the conclusion that not more
than one -tenth of known species of Agaricini are tropical,
whilst from circumstances of locality, elevation, etc., as many
as three-tenths have occurred in tropical countries ; that more
than half of the total number of Agaricini occur in Europe,
and nearly two-thirds in Europe and North America. There-
fore the northern temperate zone is the most favourable for the
Agaricini, and there is no reason to doubt that the temperate
regions of Asia will nearly equal those of Europe and America
when they are properly explored.
If we take two genera which systematically follow each
other, Amanita and Lcpiota, we shall find remarkable diver-
gences in their distribution, an explanation of which we
discover in the fact that in the former the species are large,
soft, and fragile, containing much water, whilst in the latter
a great number are small, and all are dry and tough, as
compared with other true Agarics. Hence the former genus
is essentially that of the temperate, and the latter of the sub-
tropical zone. In Amanita we reckon 80 species, of which
61 are European and North American, and 9 Australian.
The four Indian species only occur high up on the Himalayas,
and the one South American on the slopes of the Andes.
Hence the only tropical species to be accounted for are two
in Ceylon, one in Algeria, one in Java, and one in Cuba ; the
Javanese is doubtless not an Amanita at all. In this case
seven -eighths are distinctly located in the temperate zone,
one - twentieth at a temperate elevation, and only one-
twentieth presumably tropical. On the contrary, in Lefiota,
with a total of 225 species, there are 118 belonging to tem-
perate regions, and 107, or nearly one-half, to the tropical.
This is a greater proportion than occurs in any other genus
of the fleshy Agarics. Those of the temperate zone are 88
for Europe, 1 6 wholly United States — adding of course a great
number of European — and 13 Australasian (out of a total of
33) and 1 Siberian. Those of the tropical zone are — 68 for
Ceylon, 6 for India, 11 for South Africa, 15 for South
America, 3 for Cuba, and 4 for Bonin Island, Java, and Hong-
Kong. Circumstances like these render it extremely ditticult
328 INTRODUCTION TO THE STUDY OF FUNGI
to elaborate any scheme of general distribution. In the case
of the two genera given above, it should have been stated
that both are wholly terrestrial, and similar in their habitats.
The genus Cortinarius is one of considerable interest, not
only for the beauty of many of the species, but also on
account of its distribution. The number of described species
is 391, of which 371 belong to Europe and the United
States, and of these 68 are confined to America. The residue
include 14, chiefly from the most southern part of South
America, really temperate, 1 from Tasmania (nine other
European species occur in Australia), 3 species from a tem-
perate elevation on the Himalayas, and 1 species each from
Japan and the Canaries. The only tropical species is one
from Brazil. This is, therefore, a genus of strictly temperate
regions, not a single species being found in Ceylon, the West
Indies, or Africa. Two hundred of the 371 European species
are found in Sweden, and about 180 in Great Britain.
The two closely allied genera, Lactarius and Bussula,
belong also to the northern parts of the temperate zone. The
119 species of Lactarius include 85 European and 27 North
American species, one each from Madeira, Tasmania, temperate
Himalayas, and Japan. This leaves only the three Algerian
species outside of the temperate zone. For Eussula 112
species are recorded, which are thus distributed^ 9 6 to
Europe, 12 to the United States, and 2 to Australia. This
leaves only two tropical species — one to Ceylon and one to
Venezuela. Australia contains 5 species of Lactarius, of which
4 are European ; and 1 0 species of Eussula, of which 8 are
European.
The only remaining genus which we purpose to analyse
is that of Coprinus, in which the pileus is usually very thin,
the gills deliquescent, and the spores black. The number of
described species is l72, of which 117 are European, and 19
peculiar to the United States. The one species from the
Canaries is almost temperate, and also the three Australian
species. For tropical regions — 12 species for South America,
3 for the West Indies, 3 for Egypt and Mesopotamia, and 9
for Ceylon; 1 for Bonin Island, 1 for Java, and 3 for South
Africa. Many of the species are widely distributed, and
GEOGRAPHICAL DISTRIBUTION 329
are not averse to heat, so long as there is plenty of moisture,
yet four-fifths of the species belong to a temperate climate.
The next group in importance is that of the Polyporci,
with its 2200 species. This includes some genera which are
fleshy, and delight in a temperate climate ; but the majority
are of a leathery or woody substance, and can flourish in any
climate, but in many cases require a hot one. We may com-
mence by excluding 257 species of Boletus and allied genera
which are fleshy, and partake of the character of the fleshy
Agarics, for their love of a temperate region. The old genus
Pohjiwriis now constitutes four genera, of which Polyporus is
retained as the name of one genus, and inckides the annual
species, which are at first soft, and prefer a temperate or warm
temperate climate. Of the 403 species, 210 are Enropean or
North American, but chiefly in the southern parts, whilst
more than half the remainder enter the subtropical region.
The genus Fomes includes the hard woody species, which are
nominally tropical or subtropical, although a few will inhabit
temperate countries. One species, Fomes lucidus, is one of
the most cosmopolitan of Fungi, and is found all over the
world, except in the Arctic zone. Several other species have
a very wide range. The remaining genera, such as Polystictus,
Trametes, Daedalea, Hexagona, etc., extend through similar
countries to the equator, and together constitute the bulk of
the tropical Hymenomycetes.
Very little more needs to be said respecting the Hymeno-
mycetes, since the remaining 2000 species follow the same
law of distribution, the fleshy to the temperate, the leathery
and woody to the subtropical and tropical regions. We
remarked of this, a quarter of a century ago, that when the
majority of the species of a genus are of a fleshy consistence,
it may generally be concluded that it belongs to a northern
region, even if it should have some representatives in lands
which enjoy more sunshine. Thus the species of Hydnum
are the principal ornaments of northern forests, where they
attain so luxuriant a growth and beauty that every other
country must yield the palm to Sweden in respect to them.
In Irpex and Badulum the texture is more coriaceous, and
hence we find the species more commonly inhabiting warmer
330 INTRODUCTION TO THE STUDY OF FUNGI
climates. The Thclcphorci have a very wide range, and some
species of Stercum are almost cosmopolitan, or are represented
by very close allies, whilst Corticium affects generally a more
temperate region. Allied genera are distributed in con-
formity with their texture. The Clavariei are all more or
less fleshy, and have their home in temperate regions, being
represented in the tropics by Lachnocladiiim, which is of a
dry and leathery texture. Of the total of 240 species of
Clavaria there are 146 indigenous to Europe and the United
States, and 20 others to a temperate climate, whilst probably
10 of the original number of species belong to Lachnodadinm
or Caloccra, which would account for three-fourths as inhabit-
ants of the temperate zone, and only one -fourth to be
accounted for in warmer countries. Of the Tremcllini only
some of the Hirneolae belong to a subtropical climate.
The Gastromycetes, or puff-ball family, is a comparatively
small one, with about 720 species, and these are subdivided
into four distinct sections. The Fhalloideae are fleshly fetid
Fungi, which prefer a warm climate, although a few species
reach the south temperate zone. There are only about 93
species, of which 50 at least are tropical. The Nididariaccae
are small, tough species, widely distributed, and of the 65
species about one-third of them are subtropical. The chief
section, the Lycoperdaccae, contains about 480 species, of
which rather more than one-third belong to Europe and North
America. Australia is the richest country in the world for
these Fungi, possessing not less than one-fourth of the total
number of described species, whilst Great Britain has only
about one-sixth. About one-fourth of the whole are tropical
or subtropical. The subterranean family, the Hyporjad, is
only a small one, containing about 85 species, but there is
hardly a record of a subtropical species, and 68 are recorded
for Europe, so that it is almost a European family, for hitherto
it is not well represented in the United States. From the
above we may conclude, in general terms, that the Phcdloidci
are subtropical ; that the Nidulariaceae are generally dis-
tributed ; that the Lycoiperdacme prefer a warm temperate
climate, especially when dry and sandy; and that the Hyiwgaci
are absolutely of a temperate zone, and chiefly European,
GEOGRAPHICAL DISTRIBUTION 331
The following are recent estimates of the number of Gastro-
mycetes in the countries of Western Europe: — Britain, 78;
France, 85; Belgium, 31; Netherlands, 39; Scandinavia,
50. Also in the middle and south — Germany, 75 ; Italy, 80 ;
and Austro-Hungary, 40. The Hypodermci follow mostly the
distribution of the host-plants, as they are all parasitical, and
the greater portion inhabit a temperate zone. Estimating the
Uredines at about 1430 species, their chief home is in Europe
and North America, but follow their host-plants, when those are
cultivated, wherever they go. Of these, about 370 are exclu-
sively subtropical, or nearly one-fourth of the whole ; the
residue may be assumed to belong to the temperate or warm
temperate zone. The Ustilagines, about 330 species, have a
similar distribution. Bunt, Tilldia caries, and smut, Ustilago
segctum, have followed the wheat and oat plant to Australia,
as well as has Fuccinia graminis, and in some cases cause
more mischief than in their original home.
We pass now to the Ascomycetes, of which the principal
features of structure have already been given, but we do not
intend to attempt any elaborate account of their distribution.
The really fleshy species are for the most part in the Disco-
mycetes, and we may repeat that the fleshy species, such as
Morchella, Hclvdla, and the old genus Peziza, are exclusively,
or nearly so, inhabitants of temperate regions. The species of
Morchella found in the north of India are from temperate
elevations, and always of small size. The Triclioscyphae- ixre,
almost the only Pezizae of hot climates, and they are of a
peculiar tough substance. The finest species of fleshy Pezizae
are to be found in the North of Europe and America. The
tree-morels, or Cyttariae, are confined to the temperate zone of
the southern hemisphere. Out of a total of 3800 species,
about two-thirds are soft and fleshy ; whilst the residue are
fleshy when moist, or have a fleshy disc. If we accept
Ceylon as an example of a tropical climate, we shall discover
that with its 700 species of Hymenomycetal Fungi, the same
list contains but 50 Discomycetes. If we separate the
strictly fleshy species of Cyttariaceae, Helvellaceae, Pezizaceae,
and Ascobolaceae from the rest, v/e shall have 2390 species,
of which no less than 207 G are to be found in Europe and
332 INTRODUCTION TO THE STUDY OF TUNC, I
the United States. Of the remaiuiiig 3 14 the majority will
be found inhabiting the temperate zone of the southern
hemisphere. Hence this portion of the Discomycetes must be
accepted as confined in a remarkable manner to a temperate
climate.
The whole of the Pyrenomycetes, according to the latest
enumeration, are not less than 10,478, and of these a large
proportion belong to Europe and North America, — probably
not so much on account of their actual preponderance in
nature, as because of the greater attention which has been
paid to their collection and investigation. The distribution is
rather unequal in such a large group, some large genera being
almost tropical, while others are nearly wholly temperate. For
example, the Hjjpocrcaccac are fleshy, and hence a large pro-
portion occur in temperate regions. The Dothideaceae and
Microthyriaceae, on the other hand, are tropical, or sub-
tropical, and so also are some genera of the Sphaeriaceae.
This will appear more clearly if we divide the w^hole into
subsidiary groups, and first examine into the Perisporiaceae,
with about 770 species. These again consist of the Ery-
sipheae and the Perisporieae, the latter subtropical, the former
temperate. The few species of Erysipheae not found in
Europe or North America will be found in temperate Asia or
in the temperate zone of the south hemisphere. With the
other group it is the reverse, for the European species of
Perisporieae are few, and in some genera none, whereas
in North America they are found in the southern states.
Meliola is really the tropical, or subtropical, analogue of
Erysiphe, and with Astcrina, Dimerosporium, and Capnodium
rarely found, and only in a depraved state in Southern
Europe.
The Hyjwcrcaceae number nearly 900 species, and these
preponderate in temperate regions, but some species extend
into the subtropical. The remarkable genus Cordrjceps, the
species of which possess a fleshy stroma, growing mostly on
dead insects, has some 50 species, of which 14 are European,
8 North American, 5 Australian — or 27 temperate against
23 subtropical. Again in Hijpomyces, with 54 species, all
except six are found in Europe or North America. And also
GEOGRAPHICAL DISTRIBUTION 333
in Hypocrca there are 102 species for Europe and North
America, against 54 for all other localities. This will be
sufficient to show that the majority are in favour of a temperate
climate.
Taking the Dotliideaceae and Microthyriaceae together, the
number of species would be about 650. A large number of
the species appear as shining black dots or patches on living
or fading leaves, and especially the leaves of forest trees.
Some, of course, are erumpent on twigs. We have only been
able to trace 116 European species, or about one-sixth of the
whole, and there are certainly not so many more in the United
States, so that two-thirds of the total number will be tropical
or subtropical.
Of the 7500 species of the 8pliaeriaceae we cannot
attempt an analysis. Ellis gives 1680 North American
species, which is two-ninths of the whole, and many of these
are European also. Some of the species are cosmopolitan,
such as Daldinia concentrica ; and some, such as Xylaria
polymorpha and Xylaria hypoxyloii, are found almost every-
where, even in the tropics. Xylaria and Eypoxylon have their
representatives all over the world, amid heat or cold, but
with an evident preference for the former. In Cuba we find
20 species of Xylaria and 30 of Hypoxylon, in Ceylon
nearly the same number of both ; but of the simple scattered
Sphaeriaceae the number of species is very small, — probably
only a very few collectors would observe them or hunt for
them, and tliey require looking after, — yet there is no reason
why they should not be as common in the north of Africa or
South America as in the United States or the south of Europe.
The 300 species of Lophiostomaceae might practically be united
with the above, as they follow the same distribution. Of
these 170 are European, and only about 20 subtropical.
Another small group consists of the Hysteriaceae, in which the
texture is that of Sphaeriaceae, but with the habit and com-
pact disc of the Discomycetes. The number of species is also
about 300, and some of these are widely distributed; they are
capable of bearing a subtropical climate, although only about
84 occur outside of Europe and North America, and not more
than half of these are subtropical.
334 INTRODUCTION TO THE STUDY OF FUNGI
The home of the Tnheraceac is in the south of Euro})e, and
of the 145 species 138 are European. The Phycomycetes,
which inchide the Mucors ; the aquatic moulds (the Sapro-
Icrjniaccac) ; the Peronosporeae, which are plant parasites ; the
Entomopkthoreae, insect parasites ; and a few small groups, are
chiefly European or North American.
Of imperfect Eungi, the Sjjhaerojisidcac almost follow the
distribution of the Sphaeriaceae ; and the moulds, or Hyplio-
mycetcs, prefer a warm damp atmosphere in the warm temper-
ate zone to a hotter region. The Dematiaei reach farther
towards the equator than the Mucedines. About 30 species
are recorded for Cuba and 50 for Ceylon, and of these the
Mucedines are of a low type ; hence the 4800 species recorded
must be sought in temperate regions.
The yeast Fungi and Microbes, or SaccJtaromycetes and
Schizomycetes, depend so much upon their surroundings that no
scheme of geographical distribution can be propounded. The
Myxoriiycdcs are much the strongest in Europe and America,
and with a few exceptions are almost entirely confined to
those regions. A few of the widely diffused species, such as
Stemonitis fusca, Physarum cinereum, and Spumaria alba,
sometimes appear at remote places, but they seem to be more
scarce than would be anticipated in subtropical localities.
Although Thwaites found some 50 species in Ceylon, and 30
species were collected in Cuba, they are rarely to be met with
in tropical collections. Out of a total of 450 described species,
we can only find 67 that are not represented in Europe or
North America.
This appears to be the most complete general survey which
we can arrive at with our present knowledge of the Fungi of
tropical and subtropical regions, in which the information is
most fragmentary. In many cases our knowledge of the Fungi
of any given country depends on the work of a single col-
lector, and in no single instance has a tropical country been
thoroughly investigated. For the larger, woody, or otherwise
persistent species there is no difficulty, as they are conspicuous
objects, readily seen, easily collected, and can be conveyed with-
out much difficulty ; but the fleshy species, which soon decay
or deliquesce, and the minute species, only to be seen with a
GEOGRAPHICAL DISTRIBUTION
335
lens, never find their way to the places where they could be
identified and recorded. Hence the advantages which the
Mosses, the Lichens, and even the Algae possess are denied to
the Fungi, so that the complete history of their distribution
can never be written.^
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De Lamarck et De Candolle, A. P. La Flore Frangaise. 8vo. 6 vols.
' 1805-15.
Duby, J. E., in De Candolle, Botanicon Gallicum. 8vo. Paris, 1828.
Switzerland
Secretan, L. Mycographie Suisse. 3 vols. 8vo. Geneva, 1833.
Trog, J. G. Tabula Analytica Fungorum. 8vo. Berne, 1846.
Verzeichniss Schweizerischer Schwamme. Berne, 1846.
Austria
Bresadola, J. Fungi Tridentini. 8vo. Col. plates. Trent, 1881.
NiKSSL, G. VON. Zur Pilzfiora Niederosterreichs. 8vo. Vienna, 1857.
Beck, G. Zur Pilzflora Niederosterreichs. 8vo. Vienna, 1881-86.
Opiz, F. M. Seznam Rostlin Kveteny Ceske. 8vo. Prague, 1852.
Krombholz, J. V. Ahbildungen und Beschreibungen der Schwamme. Fol.
Col. plates. Prague, 1831.
Hungary
ScHULZER VON MuGGENBURG. Enumeratio Syst. Fungoru,m Hungariae. Vienna,
1857.
Italy
BizzozERO, G. Flora Vencta Crittogamica. Parti., " Fungbi." 8vo. Padua,
1885.
22
338 INTRODUCTION TO THE STUDY OF FUNGI
Inzenga, G. Fuiujhi Skiliani. 4to. Col. plates. Palermo, 1869, etc.
Passerixi, G. Elcnco di Funghi Parmensi. Genoa, 1867.
Venturi, a. / miceti dell' agro Bresdano. Fol. Col. jjlates. Brescia,
1845-60.
Saccardo, p. a. Fungi Italici. Fol. Col. plates. Padua, 1877-86.
et Berlese, a. N. Catalogo dei Fnnghi Italiani. 8vo. Varese, 1885.
Spain and Portugal
Thuemex, F. de. Contribution's ad Floram Mycologicam Lusitmiicmn. 8vo.
Coimbra, 1879. Continued by Dr. G. von Niessl. Coimbra, 1883.
ASIA
Siberia
Saccardo, P. A. Mycetes Sibirici. Brussels, 1889.
Thuemen, F. de. Beitragc zur Pilz-Flora Sibiriens. 8vo. Moscow, 1882.
China
Kalchbrenner et Thuemen, Fungorum in Itincre Mongolico a Potaninet in
China boreali cnumeratio. St. Petersburg, 1880.
Patouillard, W. CJiampignons de la Chine. Toulouse, 1886.
Quelqucs Champignons de la Chine. Toulouse, 1S90.
Japan
Zollinger, H. System. Verzeichniss der in Indischcn Archipcl. und aus Japon
empfangenen Pflanzen. 8vo. Zurich, 1854.
Cooke, M. C. "Fungi of Japan." Grevillca, -s.. 1882.
Berkeley, M. J. "Fungi of Challenger Expedition." Linn. Journ., xvi.
1878.
India
Berkeley, M. J., and Currey, F. A. "A Collection of Indian Fungi made
by S. Kurz." Linn. Trans., 2nd series, vol. i. London, 1876.
"Decades of Fungi," xxv.-xxx. " Sikkim Himalayan Fungi." Hook.
Journ. London, 1850.
"Decades of Fungi," xxxii.-l. "Sikkim Himalayan Fungi." Hool:
Journ., vols, iii.-vi. 1851-54.
Barclay, A. Descriptive List of the Uredineac of Simla, West Himalaya.
8vo. 3 parts. Calcutta, 1888-90.
Additional Uredineae from Simla. 8vo. Calcutta, 1891.
Montagne, C. "Cryptogamae Nilgherrensis." Ann. des Sci. Nat. Roy.
8vo. Paris, 1842.
Cooke, M. C. "Some Indian Fungi." Grevillca, vol. iv. 1876; vi. 1877.
London, 1876-77.
"Himalayan Fungi." Grcvillea, vii. 1878, 1879 ; viii. 1880.
GEOGRAPHICAL DISTRIBUTION 339
Ceylon
Berkeley, M. J. "Decades of ruiif,n," xvii.-xx. — "Ceylon." Hook. Jour n.,
vol. vi. London, 1847.
and Broome, C. E. "Fungi of Ceylon," in Journ. Linn. Sac. 8vo.
London, 1870-71.
Malay Peninsula
Saccardo, p. a. Mycetcs Malaccensis. 8vo. 3 col. plates. Venice, 1888.
Cooke, M. C. " Fungi of Perak. " GreviUea. 8vo. xii. 1884 ; xiii. 1884.
"Malacca Fungi." GrcviUca, xi. 1883 ; xiv. 1885.
EAST INDIA ISLANDS
Borneo
Cesati, V. Mycetum in itincrc Bornecnsi lectorum Bcccari. 4to. Naples,
1878.
Java
JuNGHUHN, F. Praemissa in Floram Cryptogamicam Javac Insulae. 8vo.
Batavia, 1838.
Nees von Esenbeck. "Fungi Javanici," in Acta Nova Acad. Ccs. Lcop. Carol.
4to. Vol. xiii. Jena, 1826.
Zollinger, H. Ohscrvationcs Phytograpliicae. 8vo. Batavae, 1844,
Fungi Archijxlagi Malaijo-Neerlandici Novi. 8vo. Batavia, 1845.
L^veill:^, J. H. "Champignons exotiques," in Ann. des Sci. Nat., 3rd series,
vols. ii. iii. 1844-45.
MoNTAGNE, C, and Berkeley, M. J. "Decades of Fungi," ii. Hooker's
Journ., vol. iii. London, 1844.
Cooke, M. C. "Fungi of Java," in GreviUea, xviii. 1889.
Philippines
Berkeley, M. J. " Enumeration of Fungi collected by Cuming in the
Philippines." Hook. Journ,, i. 1842.
" Ynngi oi Challenge)- Ex-pedition." Linn. Journ., xvi. 1878.
New Guinea
Hennings, p. Fungi Novo-guincensis. Berlin, 1892.
Cooke, M. C. " Fungi of New Guinea. " GreviUea, xiv. 1886.
Aru Islands
Berkeley, M. J. "Fungi of Challenger Expedition." Linn. Joxirn., xvi.
1878.
Pacific Islands
Berkeley, M. J. "Description of Fungi collected by R. B. Hinds in Islands
of Pacific." Hook. Journ., i. 1842.
" Fungi of C/taZZejigfcr Expedition. " Linn. Jou7-n., xvi. 1878.
Richards, H. W. Beitrcige zur Flora der Hawaiischen Inscln. Vienna. 1878.
340 INTRODUCTION TO THE STUDY OF FUNGI
AUSTRALASIA
Australia
Bailfa', F. M. List of Queensland Plants, with Sup2)lements. 8vo. Brisbane,
V. d.
Berkeley, M. J. "Australian Fungi," in Jour. Linn. Soc. 8vo. London,
1872.
"Fungi of the Challenger Expedition." Journ. Linn. Soc. 8vo.
1875-78.
"Decades of Fungi," iii.-vii. "Australian." Hook. Journ., vol. iv.
1845.
and Broome, C. E. " Fungi of Brisbane," in Trans. Linn. Soc. London,
1879-87.
Bresadola, J., and Saccardo, P. A. Pugillus Mycetum Australiensium, in
Malpighia. Genoa, 1890.
Cooke, M. C. Fimgi Australiani, reprinted from Grsvillea. 8vo. London and
Melbourne, 1883.
Handbook of Atistralian Fungi. 8vo. Col. plates. Melbourne, etc.,
1892.
Fries, E. Plantae Preissianae in Australasia Coll. 8vo. Hamburg, 1844-48.
Saccardo, P. A. "Mycetes Aliquot Australiensis." Bulletin dc la Soc. Mycol.
France, v, Paris, 1890.
" Fungi Aliquot Australiensis." Hedwigia. Vol. xxix. Dresden, 1890.
M 'Alpine, D. Systematic Arrangement of Australian Fungi. 4to. Melbourne,
1895.
New Zealand
Berkeley, M. J., in Hooker's i^to-a o/ iV«<; Zealand. 2 vols. 4to, London,
1853-55.
in Handbook to the Flora of Neiv Zealand, by J. D. Hooker. Svo.
London, 1864.
Cooke, M. C. "New Zealand Fungi," in Grevillea, viii. 1879.
Tasmania
Berkeley, M. J., in Hooker's Flora of Tasmania. 4to. 2 vols. Col. plates.
London, 1860.
Kerguelen Lsland
Berkeley, M. J. "Fungi of Kerguelen Land Transit Expedition." Linn.
Journ., XV. 1877.
Auckland and Campbell Lslands
Berkeley, M. J., in Hooker's Botany of the Antarctic Foyagc of H.M.S.
" Erebus" ami " Terror." 4to. Loudon, 1845.
GEOGRAPHICAL DISTRIBUTION 341
AFRICA
Algeria
DuEiEU et MoNTAGNE, C. Florc d' Algiric—Cnjptogames. Fol. Col. plates.
Paris.
RouMEGUERE, C, et Saccardo, p. a. "Fungi Algeriensis Trabutiani." Revue
Myc. 1881.
Saccaedo, p. a. F%(,ngi Algeriensis. Toulouse, 1886.
Fuiigi Algeriensis, Tahitensis et Gallici. Toulouse, 1885.
Egypt
Barbey, W. Champignons rapp. cCEgypte et de Palestine. 8vo. Paris, 1881.
De Thuemen, F. "Fungi Aegyptiaci," in Grevillea. 1879.
AscHERSON, P. Beitrag zur Flora Acgyptens. Berlin, 1879.
Abyssinia
Saccaedo, P. A. Fungi Ahyssinici. 8vo. Genoa, 1891.
Natal
Feies, E. Fungi Natalensis. 8vo. Stockholm, 1848.
Cooke, M. C. "Fungi of Natal," in Ch-cvillea, viii. 1880 ; ix. 1880 ; x. 1881.
"African Fungi," in Grevillea, xix. 1890.
Cape Colony, etc., South Afeica
Kalchbeennee, C. " Fungi Macowaniana," in Grevillea, ix. 1880 ; xi. 1882.
and Cooke, M. C. "South African Fungi," in Grevillea, ix. 1880-81.
Fayod, V. " Beitrage zur Kenntuiss der Flora von Deutsch Sudwest Afrika."
Ahhand. der Bot. Vcreinsf. Brandenbcrg, xxxi.
Berkeley, M. J. "Fungi, collected by Zeyher in Uitenhage." Hool:. Journ.,
voh ii. 1843.
"Decades of Fungi," No. 1. Hook. Journ., vol. iii. 1844.
MoNTAGNE, C. Enwneratio fungorum d M. Dregc in Afrika Mcrid., etc.
Paris, 1847.
Madagascae
Cooke, M. C. "Fungi of Madagascar, " in Grevillea, xviii. 1890.
West Coast
Welwitsch, F., and Cuerey, F. "Fungi Angolensis." Trans. Linn. Soc,
vol. xxvi. 4to. London, 1867.
Beesadola, G. Fungi Kamcrunenses. 1890.
Cooke, M. C. " Gaboon Fungi, " ffremY^ea, xv. 1887.
Afzelius, A. Reliquiae Afzelianae. Fungorum ah Afzelio in Guinea Coll.
Fol. Upsal, 1860.
342 INTRODUCTION TO THE STUDY OF FUNGI
St. Thomas Island
Bresadola, G., and Roumeguere, C. "Nouvelles Contributions a Flore Myco-
logique des lies St. Thome et des Princes." Revue Mycol. 1890.
Berkeley, M. J. "Fungi of Cliallenger Expedition." Lliin. Journ., xiv.
London, 1875.
NORTH AMERICA
Canada and Arctic America
Berkeley, M.J. "Fungi of Arctic Expedition, 187.5-76." Linn. Journ., xvii.
1880.
"Fungi of M'Clintock's Expedition." Linn. Journ., v. 1861.
Ellis, J. B. "Canadian Fungi." Journ. Mycol. , vol. i. Manhattan, 1885.
Watt, D. A. P, " Provisional Catalogue of Canadian Cryptogams." Canadian
Naturalist, vol. ii. Oct. 1865.
United States
Schweinitz, L. de. Synopsis fungorum in Amcr. Borcali media dcgentium.
4to. Philadelphia, 1831.
■ Synopsis fungorum CaroUnae Superioris. 4to. Leipzig, 1822.
Berkeley, M. J., and Curtis, M. A, "North American Fungi," in Grevillca,
vols. i.-iv. London, 1871-75.
"Centuries of North American Fungi." Ann. Nat. Hist., 2nd series,
vol. xii., and 3rd series, a'oI. iv.
Curtis, M. A. "Contributions to the Mycology of North America." Silliman
Journ. 8vo. 1848.
Catalogue of the Plants of North Carolina. 8vo. Raleigh, 1867.
Berkeley, M. J. "Decades of Fungi," viii.-x., in Hook. Journ., vol. iv.
London, 1845.
"Decades of Fungi," xii. -xiv. "Ohio Fungi." Hoolc. Journ, vol. vi.
London, 1847.
"Decades of Fungi," xxi.-xxiv. "North and South Carolina." Hook.
Journ., vol. i. 1849.
" North American Fungi, " in (?rm^te, vols. i. and ii. 8vo. London.
Harkness, H. W. Pacific Coast Fxmgi, i.-iv. San Francisco, 1885-87.
Lea, T. G. Catalogue of the Plants of Cincinnati. 8vo. Philadelphia, 1849.
Peck, C. H. Picports of the New York Museum of Natural History. Albany,
1872-94.
Cooke, M. C. "Fungi of Texas." Linn. Journ., vol. xvii. 1880.
and Ellis, J. B. "New Jersey Fungi," in Grevillca. — Various.
and Harkness. " Californiau Fungi," in (?rCTi7^ea. 1878-80.
Ellis, J. B., and Everhart, B. "New American Fungi," in Journal of Mycology.
8vo. 1885-95.
Farlow, W. G., and Trelease, W. List of Works on North American Fungi.
Svo. Cambridge, 1887.
Bermuda
Berkeley, M. J. " Fungi of Challenger Expedition." Linn. Journ,, xiv. 1875 ;
XV. 1877.
GEOGRAPHICAL DISTRIBUTION 343
Central America
Fries, E.' Novae. Symholae Mycologicae {F%mgi Mexicani). 4to. Upsal, 1851.
KiCKX, J. Note sur quclques Champignons du Mexique. 8vo. Brussels, 1841.
Berkeley, M. J. " On some new Fungi from Mexico." Lhm. Journ., ix. Svo.
London, 1867.
WEST INDIAN ISLANDS
Cuba
Berkeley, J. M., and Curtis, M. A. "Fungi of Cuba," in Journ. Linn. Soc.
Svo. London, 1867.
MoNTAGNE, C. , in Historia fisica dc la isla de Cuba par Ramon de la Sagra —
Planies Cellulaires. Svo. Paris, 1838-42.
Jamaica
SwARTZ. Flora Indiac Occidentalis.
Bahamas
Berkeley, M. J., in " Fungi of Challenger Expedition." Joimi. Linn. Soc. Svo.
1875-78.
SOUTH AMERICA
Guiana
MoNTAGNE, C. " Cryptogamia Guyanensis. " Ann. des Sci. Nat. Paris, 1855.
Cooke, M. C. "Cocoa-palm Fungi." Grevillea, vol. v. London, 1877.
Berkeley, M. J. "Enumeration of Fungi collected by Dr. Hostmann in
Surinam." Rook. Journ., vol. i. London, 1842.
and Curtis, M. A. ExoticFungi{Schweinitz),principally from Surinam.
Roy. Svo. Philadelphia, 1854.
Venezuela
Berkeley, M. J. Some new Fungi from Venezuela. Svo. London, 1857.
Cooke, M. C, in Grevillea, ix. 1880.
Patouillard, N. " Champignons de Venezuela. " Bull. Soc. Myc. Fr. 1888.
Equator
Patouillard, X. , et Lagerheim. " Champignons de I'Equateur," in Bidl. Soc.
Myc. Fr., vii. 1891.
Brazil
Berkeley, M. J. "Fungi Brasiliensis," in Videns. Meddels. Naturhist. for
Kjobenharn. 1879-80.
"Notices of some Brazilian Fungi." Rool\ Journ., vol. ii. London,
1843.
Decades of Fungi, xxxi. 1851.
344 INTRODUCTION TO THE STUDY OF FUNGI
Berkeley, M. J. Decades of Fungi, li.-lxii. "Rio Negro Fungi." A'ol. viii.
London, 1856.
and Cooke, M. C. "The Fungi of Brazil." 8vo. Linn. Journ., vol.
XV. London.
Henkings, p. Fu7igi Brctsilicnsis. Leipzig, 1892.
MoxTAGNE, C, in Ann. des Set. Nat., 4th series, vol. v. Paris.
Ann. clcs Sci. Nat. for July 1839.
Chili and Peru
MoNTAGNE, C. "Fungi de Gaudichaud," in Ann. des Sci. Ned., 2nd series, vol.
ii. Paris, 1834.
in Gay, Hist, fisica y politica de Chile. 1845.
Cooke, M. C. " Fungi of Peruvian Andes," in Grevillea, xiii. 1884.
Argentina, etc.
Spegazzini, C. Fungi Guaranitici. i. Buenos Ayres, 1883 ; ii. 1888 ; iii. 1891.
Fungi Fuecjiani. 8vo. Buenos Ayres, 1887.
Fungi Patagonici. Svo. Buenos Ayres, 1887.
Fungi Fuiggccriani. 8vo. Buenos Ayres, 1889.
Juan Fernandez
MoNTAGNE, C. " Prodromus Florae Fernandesianae." Ann. des Sci. Nat.
June 1835.
Berkeley, M. J. "Fungi of Challenger Expedition." Linn. Journ., xvi.
1878.
CHAPTEE XXVIII
APPENDIX ON COLLECTING
It will be manifest from the foregoing chapters that the im-
portance of obtaining mature and perfect specimens for
examination and determination cannot be too highly estimated.
It is not only essential for the determination of any species,
but in many cases even the genus, that fructification should
be present. In classification nearly everything depends upon
the spore, and if no spores are present, and only the vegetative
system is developed, any identification is the merest chance.
The true relations of an Agaric can only be sought after the
colour of the spores has been determined. This is readily
done by cutting off the stipes, and inverting the pileus with
the gills downwards upon a piece of paper, and allowing it to
remain all night in that position. In the morning the spores,
if mature, will have fallen upon the paper, in radiating lines,
corresponding to the gills. If it is suspected that the spores
are white, it is preferable to invert the pileus on a piece of
black paper, but, if they are presumed to be coloured, then
white paper will suffice. When the colour of the spores has
been determined, it can be seen to which of the primary
groups the species must be referred, whether Leucosporae,
Ehodosporae, or any other. This method may be resorted to
with all the Hymenomycetes with advantage, although it is
nowhere so important as with the Agaricini.
The value of the spores in classification is not confined to
the Hymenomycetes, but pervades the whole of the Fungi.
Those minute species in which the fructification is enclosed in
a perithecium, having the habit of a Sphaeria, must, in the first
instance, exhibit fruit before it can be affirmed whether, by
346 INTRODUCTION TO THE STUDY OF FUNGI
virtue of the presence of asci, it should have its place in the
Pyrenomycetes, or whether, on account of their absence, it must
be relegated to the Sphaeropsideae. External appearance will
furnish no direct evidence as to its place in the system, and
hence a knowledge of the spores is again the first step in
identification, without which it is impossible to proceed.
The moulds, or Hyphomycetes, again, when devoid of
conidia, are no better than a condition of mycelium. A mass
of sterile hyphae is only equivalent to mycelium, even though
some of the branches may be erect, as if they exhibited the
intention of producing gonidia. When the gonidia are present
upon the branches, then the fertile threads possess a new im-
portance, as they become gonidiophores, and in connection with
the gonidia determination of the species is probable.
Spotted leaves are often collected by the inexperienced
on the assumption that, whenever a living leaf has become
spotted in a particular manner, the spotting is due to the
presence of a Fungus. In many instances the assumption will
prove to be correct, but even then it will not be sufficient to
know that mycelivmi, or even small perithecia, are present, if no
mature fruit can be found. Many a weary hour of fruitless
labour may be expended in the examination of spotted leaves
which do not furnish the organs essential to an accurate
diagnosis.
Hence it will be evident that the collector, even if he
aspires to be nothing more, must acquire sufficient elementary
information to guide him, and prevent the accumulation of a
store of waste material, in which a pocket lens will give no
evidence of Fungus growth in the condition of fructifica-
tion. A little knowledge and experience may be sufficient to
determine whether a Hymenomycete is mature, or whether
there is any ground for the belief that in other cases repro-
ductive organs are present, in some form or other, whilst a
larger experience and a more extended knowledge may be
necessary for an accurate determination.
When it has been ascertained that a Fungus has all the
appearance of possessing mature fructification, the question is
sure to arise as to the best method of preserving it for future
examination, although it may be premised that fleshy Fungi
APPENDIX ON COLLECTING 347
can never be examined so satisfactorily as in the fresh state.
Wherever it is impossible at once to examine and determine
the name of any given Fungus, some effort must be made for
its preservation. With the soft and fleshy Agarics no
amount of careful desiccation will be satisfactory alone, as they
will soon shrink out of all recognition, change colour, and be-
come liable not only to decay, but also to quick destruction
by insects. Some persons have suggested the immersion of
the fresh specimen in some preserving fluid, such as Goadsby's
solution, methylated spirit, glycerine, etc., but none of these can
be employed, because the colour of the Agaric will be destroyed,
and, worse than all, the spores will be washed away from their
sporophores and disseminated through the fluid, suffering de-
coloration in the process.
The only method which we are prepared to recommend for
Fungi of this kind is to make a sketch, or drawing, of the
Agaric, with the form, size, and colour as in life. It is not
absolutely essential that they should be coloured, although that
is best, but the colours should always be stated explicitly upon
the drawings. To assist those who are not facile with the
pencil, it is recommended that the specimen collected should
be divided longitudinally through the cap, and down the centre
of the stem. When this is done, one half should be laid on a
sheet of white paper, with the cut surface downwards, and the
outline traced carefully upon the paper with a sharp-pointed
pencil. On removing the specimen there will be left upon the
paper an outline of the form of the Agaric, natural size. This
may be completed by hand, drawing in the line marking the
margin of the pileus, indications of scales (if any exist), the
character of the ring (if present), and the scales, lines, or mark-
ings of the stem. Another copy of the section, made side by
side on the same paper, would give the outline of the gills,
and by a little care and practice it would be found easy to
draw the line from the stem to the edge of the cap, indicating
the point of junction of the gills with the flesh of the cap.
This should be done very carefully and accurately, as it must
be depended upon to show whether the gills are quite free from
the stem at their inner extremity, or whether they are adnexed,
or whether they are decurrent, and to what extent they run
348 INTRODUCTION TO THE STUDY OF FUNGI
down the stem. Then, also, it should be shown if the stem is
solid or hollow. A little colouring, even if not artistic, would
be more useful than mere description of general appearance.
Of no less importance is the addition of notes, giving such
particulars as cannot be conveyed by the sketch, and these
would embrace a statement of habitat, whether growing on the
ground or on wood. Amongst other details it should be stated
whether the pileus was dry or moist and glutinous, whether
the odour was agreeable or fetid or indistinct, whether the
taste was mild or acrid and pungent, and whether the gills
exhibited any tendency to deliquesce. Finally, if the drawing
was not coloured, then the colour of the pileus and stem must
be indicated as explicitly as possible, and not vaguely, as red,
brown, or gray, but what particular tone of each colour, whether
bright red or dull red, dark red or light red, vermilion or
crimson, and so on, with any other colour, so that at any time
the sketch might be completed in colour and made to represent
the species.
Having done this, the next step would be to utilise the
specimen itself, supposing it to be an Agaric, by cutting a thin
slice from the section of the pileus and stem, laying it upon
blotting-paper to dry ; the half stem may then be removed and
laid to dry, in order to represent the outer surface of the stem ;
and then the gills and flesh of the half pileus may be cut away
so as only to leave a little of the flesh adhering to the cap.
By this means we should have three pieces to represent the
half Agaric, viz. the section of the pileus and stem, the half
stem, and the half pileus, in order to show its external surface.
When this is done, the three pieces are to be placed on blotting-
paper, covered by one or two thicknesses of paper, and sub-
mitted to a gentle pressure, so as to prevent curling, and
allowed to dry. At first the papers must be changed every
two or three hours, because of the moisture they will absorb,
but later on less frequently, until the specimens are quite dry.
It will soon become manifest that all trace of the original colour
will disappear, and the fragments shrink from loss of moisture ;
so that, without the precaution of making a previous drawing,
there would be little chance of identification. The above
suggestions as to drying apply only to species of a compara-
APPENDIX ON COLLECTING 349
tively large size. There will be hundreds of species so small
that they cannot be manipulated further than by taking a
section through the cap and stem, after which the specimens
will dry up readily in their entirety, and may be kept in small
envelopes attached to the drawing. It may be asked. Of what
use are these dried fragments, if they are insufficient to
determine the species ? Granted, that they are only acces-
sories to the sketches, yet they will be sufficient to indicate
clearly the colour, size, and shape of the spores, the mode of
attachment of the gills to the stem, and the nature of the
scales, warts, or silkiness of the surface of the pileus ; but beyond
this they can teach very little, nor by any other method yet
devised can fleshy Fungi be preserved, so as to retain the form,
colour, and size of their natural condition. In a few genera of
the Agaricini, such as Lentinus, Lenzites, Schizophyllum, and
even Marasmius, where the substance is dry and tough, the
species will be readily dried in their entirety, and by aid of a
few brief notes may be determined without difficulty. Fleshy
species of Boletus and Hydnum will have to be subjected to
the above-named process of drawing and desiccation.
The large woody Fomes, and the smaller leathery Polystictus,
with the resupinate Poriae and nearly all the Thelephorei,
require only to be dried in the air, in some cases under pressure
to keep them flat, and in this condition they do not lose much
either in colour or form. These are, consequently, the most
commonly selected species which are collected by travellers in
foreign countries, whilst the smaller or more fragile are neglected.
The whole of the Gastromycetes, excepting the Phalloidei,
require little or no preparation. They only need be collected
when mature, and dried in the air. The same may be said of
the Myxomycetes, which only require to be placed in small
pill -boxes, and secured by pins or otherwise, as insects are
secured, so as to prevent injury in transit. In no case should
more than one species be placed in a single box, or the spores
will be transferred and confusion result.
As to the collection of moulds and mucors little can be said,
as it seems scarcely possible to carry such delicate objects, even
for short distances, without injury. For home purposes we
have used small boxes, with fragments of cork glued to the
350 INTRODUCTION TO THE STUDY OF FUNGI
bottom. It is possible sometimes to obtain some mould upon
its matrix, which can be placed in such a box, and pinned to
the cork, and thus the conidiophores will remain intact; but
the conidia are attached so slightly that very few of them will
be found in situ. Compact species of such families as Tuber-
culariae and Stilbaceae are easily transported and preserved.
Parasitical species such as the Uredines and all leaf Fungi
are the easiest to collect and preserve, and for them no
instructions are required except, perhaps, the suggestion that
the leaves should be pressed and dried flat in all cases, as they
consequently will occupy much less room, and can be
examined more readily when dry.
The Discomycetes, although some of them are large and
fleshy, such as Morels and the more imposing Pezizeae, may
be dried in the air, taking care to note always the colour of
the disc when fresh. After being dried they will resume
their old form and dimensions when placed in water, although
they will never regain the lost colour. The only disadvantage
which results from reviving them in this way is, that when
they dry again they are liable to become hard, horny, and
brittle, except in the tough and leathery species.
It is only necessary now to allude to the largest and
most widely distributed group of Fungi, containing not less
than 17,000 species. These are the Pyrenomycetes, with
the Sphaeropsideae, which latter resemble the Pyrenomycetes
in form and habit, but differ in not producing ascospores.
These Fungi are to be found on dead wood, branches, twigs,
leaves, herbaceous stems, dung, and almost every kind of
vegetable debris, the smaller species like little black dots, no
larger than a small pin's head, but the largest compound
species reaching the size of a man's fist. All of these suffer
nothing in the process of drying, and may be as readily
determined in five or ten years as on the day in which they
were collected. It will be necessary to note the locality and
date, and then each specimen can be folded in paper and put
away to await a more convenient season. It would be an
advantage, when this season arrives, that in every instance,
where possible, the name of the host should be indicated — as,
for instance, oak, beech, or elm stump, maple branch, or dead
APPENDIX ON COLLECTING 351
stem of Angelica, Eumex, or other herbaceous phmt. For the
minute species the pocket lens will consequently he in
requisition, but minute and exhaustive research amongst dead
vegetable matter, in damp situations, is almost certain to be
well rewarded.
We have possibly passed over, in this brief generalisation,
small and interesting subsidiary groups, which are technically
included under the larger ones, to which attention might have
been profitably directed. If we were to advise a young
collector as to the course he shovild pursue with the greatest
profit and interest to himself, it would be, that, after making
himself generally acquainted with the characteristics of the
primary groups, as we have indicated them, he should select
for himself a compact family of moderate size, and devote
himself to that group alone until he is familiar with all the
details ; after this course of practical education, he might with
advantage widen his field of operation and extend his patron-
age to other groups. By concentration of his thoughts and
energies he will be the better able to cope with the difficulties,
and master the details, of a comparatively small group, than
by attacking a large one. There are several of such groups
available — as, for instance, the Uredines, the Ustilagines, the
Myxomycetes, the Gastromycetes, or even the Hyphomycetes.
In whatever direction his inclination may lead him, the
student will find peculiarities, and adaptations of methods of
examination and study, applicable to the special objects of
his research. In none will he be able to proceed far without
the use of the microscope, and we would strongly urge upon
him the necessity of cultivating the power of the hand in
making sketches and drawings, either with or without the use
of the camera lucida, or some form of substitute. Accurate
drawings, made to scale, of reproductive bodies, structural
details, modes of development, and other minutiae will
always prove a source of satisfaction in the future, and a help
towards progress.
Finally, we would urge also upon the young and inex-
perienced never to rest content with being mere collectors,
since the knowledge so obtained is liable to become super-
ficial and empirical ; but, on the contrary, to examine for
352 INTRODUCTION TO THE STUDY OF FUNGI
himself, as thoroughly aud completely as possible, every
organism which he acquires in his own selected group, and
endeavour to ascertain all that is possible of its life-history.
The whole history of one species, worked out with persever-
ance and intelligence, will present the key to a knowledge of
many kindred species, and always prove to be a valuable
contribution to science, when the names of species are
changed or forgotten.
GLOSSARY
AcROGENOUS — produced at tlie sum-
mit.
Ac7-o(joni(lium—gomi)i\\xVi\ at tlie sum-
mit of a gonidiophore.
Acrosporc — spore formed at the summit
of a sporophore.
AecicUospore — spores formed in an
Aecidium, serially and successively
abstricted.
Aecidium — cup - shaped receptacle in
the Uredines, enclosing a hymenium
producing Aecidiospores.
Aethalium — body formed in Myxo-
mycetes from a large combination of
Plasmodia.
Alveolate — pitted like honeycomb.
Amoeboid — like an Amoeba ; applied to
a protoplasmic body which creeps
by putting out and retracting
pseudopodia.
Angiocar2}oiis—h.a.vmg the hymenium
developed within the sporophore, and
covered from the first by a special
envelope.
Annulus — in Hymenomycetes, portion
of the veil, or tissue of the stipe
forming a collar or ring.
Anfheridiuvi — male sexual organ.
ArcMcarp — cell, or group of cells, fertil-
ised by a sexual act.
ArthrosjMrous — such Schizomycetes as
have no endogenous spore-formation.
Asciferous, Ascigerous — bearing asci.
Ascocarp — a sporocarp bearing asci and
sporidia, or ascospores.
Ascogenous — producing asci.
Ascophore — sporophore bearing an
ascus.
Ascospore — spore contained in an ascus
= sporidium.
Ascus, Theca — large cell or sac in
which ascospores are developed,
typically eight.
Autoecious, or Autoxenous — a parasite
which goes through the whole course
of its development on a single host.
Autonomous— \Aaj\\ls that are perfect
and complete in themselves, not
Ibrmiug part of a cycle.
Basidiophore — sporophore bearing a
basidium.
Basidiospore — spore produced at the
apex of a basidium.
Basidium — mother - cell from which
spores are abjointed. In Hymeno-
mycetes, a sporophore bearing from
one to four spores on short sterig-
mata.
Brood-cell — same as gonidium or coni-
dium.
Cap — in Hymenomycetes, same as
pileus.
Capillitium — sterile threads or tubes,
often branched, mixed with the
spores in the spore-masses of some
Gasti'omycetes and Myxomycetes.
Carpophore — generally, the support of
the fructification ; specially, the stalk
of a sporocarp.
Carposporc — spore formed in a sporo-
carp.
Chlamydosporc — spore with a very thick
spore-membrane.
Chlorophyll — the green colouring
matter in plants, absent in all
Fungi.
Clwmp-comiection — small semicircular
protuberance attached through its
length, or leaving an eyehole, to the
walls of two adjoining cells of a
septate hypha, and stretching over
the septa between them, communicat-
ing with one or both, or cut off from
both, and forming a clamp-cell.
Clcistocarp — ascocarp forming a com-
pletely closed cavity, which is finally
ruptured to permit the ascospores to
escape.
Columella — sterile central body in a
sporangium.
Concatenate — linked together in a
chain.
23
354
IIVTRODUCTION TO THE STUDY OF FUNGI
Conceptaclc — a superficial cavity ojien-
ing outwards within Avhich coiiidia
or sporulcs are produced.
ConiiUo2)horc — same as gonidiopliore.
Conidinm — same as gonidium or brood-
cell.
Cortina, or curtain — in Hymenomy-
cetes, marginal veil, ruptured from
the stipe and lianging from the edge
of the pileus, or around the stipe in
threads.
Cryptogamia — apjjlied to the lower
orders of plants in which there are
no conspicuous flowers, as there are
in the Phanerogamia.
Cuticle, or ikIUcIc — the separable outer
layer.
Cijst — a bladder specially applied to the
terminal sporangia of Mucors.
Cystidium — in Hymenomycetes, large
])rojecting cells of the hymenium,
extending beyond the basidia and
paraphyses.
Dichotomy — branching in pairs in a
forked manner.
Disc — the hymenium of a discocarp.
Discocar}) — an open ascocarji in which
the hymenium is ex2iosed whilst the
asci mature.
Elater — in Myxomycetes, a free
capillitium thread, mostly sjiirally
marked or warted.
Endogonidium — gonidium formed with-
in a recei)tacle.
Endophytal — growing within another
plant.
Endos2}orium, E^idospore — innermost
coat of a spore.
Entomogcnous — growing upon or within
insects.
EiJiphytal — growing upon another
jilant.
Epi.iporiu7)i, Episporc — outer coat of
spore.
Excipulum — outer envelope of a disco-
carp developed as part of the recep-
tacle.
Facultative parasite — an organism
which normally goes through its
whole course as a saprophyte, but
which may also go through its course
either wholly or in part as a parasite.
Facultative sajn-opJtytc ■ — an organism
wliich normally goes through its
whole course as a parasite, but
which can vegetate at certain stages
as a saprophyte.
Flagdlum — whip-like process of a
swarm - spore, a single or solitary
long cilium.
Funiculus — in Nidulariaceae, the cord
of hyphae attaching a peridiolum to
the inner wall of the peridiimi.
Gamete — sexual protoplasmic body,
wliich on conjugation with another
gamete gives rise to a body called a
zygote or zygosi)ore.
Germ-cell — first product of commencing
germination of a spore.
Glcha — chambered spore - producing
tissue within a sporophore. As in
Gasti'omycetes.
Go7iidiophore — sporophore bearing a
gonidium.
Gonidium = conidium, or hrood-cell —
propagative cell, ]iroducedasexualh',
separating from the parent and cap-
able of direct development into a
new individual.
Gonoplas7n — portion of protoplasm ot
antheridium in Peronosporeae which
passes through iertilisation tube and
coalesces with the oosphere.
Gonosphcrc — the same as oosphere.
Gymnocarpous — having the hymenium
exposed while tlie spores are growing.
Habitat— the jilace in wliich a plant
grows.
Haustoriuin — special branch of fila-
mentous mycelium wliich serves as an
organ of adhesion and suction.
Heteroecious — forms wdiich jjass through
separate sections of their complete
history on different hosts.
Kctrrosporous — having spores asexually
produced, of more than one kind.
Homosp)orous,Isosporoufi — having spores
asexually produced, of only one kind.
Hymenium — spore mother-cells, aggre-
gated in a continuous layer upon a
sporophore, or that specialised por-
tion termed the receptacle.
Hynienophorc — portion of a sporo^jhore
which bears a hymenium.
Hyplia, as applied to Fungi — a cylind-
rical, thread-like, simple, or branched
body, consisting of a tubular mem-
brane enclosing protoi)lasm, growing
apically, and often becoming trans-
versely septate.
Hyphal bodies — irregular bodies analo-
gous to mycelium in Entomoph-
thoraceae.
Hypothccium — layer of hyphal tissue
immediately beneath a hymenium.
GLOSSARY
355
Intralamellar Ti;isujs — same as
trama in Hymenomycetes.
Isogamy — conjunction of two gametes
of similar form.
Isosporojis — same as Homosporous.
Lactiferous, Latigiferous — Leaving
or conveying latex, or milky lluitl.
Lamella — in Hymenomycetes, the gills
or folds of the hymenium, radiating
from a definite point.
Lipoxenous — applied to a jiarasite
which leaves its host and completes
its development independently.
Macrogox^ihillv, Megalogoxidium —
large gonidium compared with others
produced by the same plant.
Merispore — segment of a sjioridesm.
Metaecious — same as Heteroecious.
Microcyst — in Myxomycetes, applied to
a resting state of swarm-cells.
Microgonidium — small gonidium com-
pared with others in the same species.
See Macrogonidium.
MuUilocular spore — see Sporidesm.
Mutualisjii — symbiosis of two organ-
isms living together and mutually
helping and supporting each other.
Mycelium — vegetative portion of Fungi
composed of one or more hyphae.
Neck, or collum — conical or cylin-
drical prolongation of the apex of
perithecium in Pyrenomycetes.
OiDiUM — a generic term, sometimes
applied to concatenate conidia, which
are successively abstricted at the apex
of hyphae.
Oogamy — conjugation of two gametes
of different form.
Oogonium — female sexual organ, usu-
ally a spherical sac containing one
or more oospheres.
Oospherc — spherical body which de-
velops the oospore as the result of
fertilisation.
Oospore — product of fertilisation in
oosphere.
Ostiolum — in Pyrenomycetes, orifice or
mouth of perithecium, or pyrenocarp
through which the spores are dis-
charged.
Paraphvsis— sterile, thread-like hy-
phal branch accompanying the
mother-cells in a hymenium.
Parasite — organism living in or upon,
and at the expense of, anotlier.
Pathogen o us — producing disease.
Pcnicillate — like a pencil of hairs.
Pcridlolum — little lenticular bodies in
Nidularieae, which are free, or at-
tached by a funiculum to the inner
wall of the jjeridium. Each peri-
diolum enclosing a mass of spores.
Peridiuvi — the enveloping coat of a
sporophore, or receptacle in which
the spores are developed in a closed
cavity. In Gastromycetes sometimes
called the uterus, the contents being
the gleba.
Perithecium, or Pyrenocarp — ascocarp
with the margin incurved so as to
form a narrow-mouthed cavity. A
more or less globose receptacle, per-
forated at the apex.
Pilcus — in Hymenomycetes, the conical
or dome -shaped cap bearing the
hymenium on the under surface.
Extended also to other compound
sporophores.
Plasmatoparous — in Peronosporeae,
when in germination the protoplasm
of a gonidium issues as a spherical
mass, which becomes invested with a
membrane and projects a germ-tube.
Plasmodium — in Myxomycetes, the
multinucleate ]>rotoplasm, exhibiting
amoeboid motion.
Pleomorphism — when more than one
independent form in the life-cycle of
a species occurs it is called pleo-
morphy.
Pleicroblastic — in Peronosporeae, those
forms which produce vesicular lateral
outgrowths that serve as haustoria.
Pore — in Pyrenomycetes it is the ostio-
lum ; in Polyporei the mouth of the
tube which encloses the hymenium.
Promycelium — the product of tube
germination of a spore which con-
stricts off a number of spores, unlike
the mother spore, and then dies.
Psrudoperidium — the cup, or recep-
tacle, in Aecidium.
Pscitdopodium — in Myxomycetes, the
protruded and I'etracted protoplasm
of amoeboid forms, imparting motion.
Pycnidium — in Ascomycetes, a cavity
resembling a perithecium containing
gonidia, which are termed pycno-
gonidia.
Receptacle — general term for hol-
lowed-out body, containing otlier
bodies.
356
INTRODUCTION TO THE STUDY OF FUNGI
Jlcsti/nj-sjJorc^VL .sjiore which lies dor-
mant or rests lor a period before
germination. A hibernating spore.
nesting stage, resting 'period — stage or
period of quiescence or dormancy.
Eesupinatc — attached to the matrix by
the back.
niiizoid, or Bhizine — thread-like deli-
cate organs of attachment.
SAPROPnYTE — a plant living and
thriving on dead organic matter.
Sclerotium — hard tuber-like body tilled
with reserve material, of the nature
of a compact mycelium, which re-
mains dormant tor a time, and then
develops sporophores.
Scokcitc — peculiar rudimentary bodies
in Discomycetes which are jirobably
the first distinction of fertile from
sterile hypha, doubtfully described
as sexual.
Sorus — a heap, or aggregation, chiefly
of reproductive bodies.
Spcrmatiuni — male gamete cell which
conjugates with a trichogyne.
Spermatozoid — thread-like bodies, pos-
sessed of motion, and supposed to be
fecundative.
Sporangiole — small sporangium, pro-
duced in some genera of Mucors,
supplementary to large sporangium.
Sjmrangiophorc — the sperojihore of a
sporangium.
Sporangium — envelope or sac in which
spores are produced.
Spore — in a general sense it is a repro-
ductive cell, which becomes free, and
is capable of developing into a new
plant ; in a special sense, restricted
to the Hymen omycetes.
Sporidesm — multicellular spore-body,
becoming free, of which each cell is
an independent spore.
Sporidiolum — diminutive of sporidium
— applied to promycelial spores in
the Uredines.
Sporidium — in Ascomycetes, a spore
developed in an ascus. In Uredines
a. spore abjointed on a promycelium.
Sporifcrous — bearing spores.
Sporocarp — multicellular body, de-
veloped sexiially from an archicarp,
unlike the body which produced the
archicarp, and serving to form
spores.
Sporogcnous — producing spores.
Sporophore — branch which bears spores,
or mother-cells.
Sjiorule — designation for spore, en-
closed in a perithecium, in imperfect
Fungi, such as Sphaeropsideae, with-
out asci.
Sterigma, Sjncule — slender stalk-like
branch of basidium bearing a spore.
Stipe — general term for the stalk of a
sporophore, usually applied to the
stem of Agarics.
Stroma — Fungus body with the form of
a cushion, crust, club, or branched
exjiansion ; usually supporting cora-
]iound fructitication.
Stglospore — sjiore borne on a filament.
Suspcnsor — in Mucors ; club-shaped or
conical portion of hypha, adjoining a
gamete cell after its differentiation.
Sivarm-cell — naked motile protoplasmic
body.
Symbion — an organism living in a state
of symbiosis.
Symbiosis — the living together of dis-
similar organisms.
Televtospore — in Uredines, the ulti-
mate spore of the cycle which is
cai>able of germinating and producing
a promycelium.
Tliallopkytes — cellular Cryptogamia.
Includes Algae and Fungi, where
there is no ditferentiation into stem
and leaf.
Thallus — the vegetative body of a
Thallo])hyte.
Theca — the same as ascus.
Tliccaspore — synonymous with asco-
spore.
Trama — in Basidiomycetes, the middle
tissue of the gill plates, or other
projections of the receptacle which
bears the hymenium.
Trichogyne — female receptive portion
of an archicarj) to which the sper-
matia become attached.
Tubulus, Tube — in Hymenomycetes,
the tube lined with the hymenium,
as in Polyporei.
Uredo — hymenium producing uredo-
spores.
Uredospore Urcdogoniditim — in Ure-
dineae, spore formed upon a sporo-
phore from which it separates at
maturity, and on germination pro-
duces a mycelium bearing uredospores
or teleutospores, or both.
Uterus — same as peridium in Gastro-
mycetes.
Veil, Velum — in Hvmenomycetes,
special envelope in which the growth
GLOSSAJ^y
357
of the whole or part of the sporo-
phore takes place.
Volva, Velum universale — in Hymeiio-
mycetes, sac enclosing the whole of
a sporophore at first, but ultimately
ruptured at the apex by the expand-
ing pileus.
Veast fungus — species of Saccharo-
myces.
ZooGLAEA — in Schizomycetes, a colony
imbedded in a gelatinous stratum.
Zoogonidium — active gonidium. See
Zoospore.
Zoosporangium — sporangium contain-
ing zoospores.
Zoospore — motile spore.
Zygote, Zygospore — spore resulting from
the conjugation of two similar
gametes.
INDEX
Aecidium, 243
Agaric, parts of, 32
Agaricus, section of, 127
Alternation of generations, 70
Anthrax, 295
Appendix on collecting, 345
Archicarps, 229
Arthrospore, 292
Artificial cultures, 249
Asci and parapliyses, 104, 166
Ascomycetes, 164
Ascus and sporidia, 45
Aseptate mycelium, 16
Atrophied basidia, 121
Bacteria, 291
Basidia, 41, 120
Basidiomycetes, 115, 119
Beech Morels, 186
Bird's-nest Fungus, 153
Black moulds, 278, 283
Bunt spores, 255
Calostoma, development of, 156
Capillitium, 43, 309
Capsular Fungi, 197
Carpophore, 22
Car})ophores agglomerated, 28
Census of Fungi, 319
Clamp-connections, 11, 150
Classification, 93
Clavate carpophore, 29
Clinospore, 45
Cluster-cups, 48, 242
Collecting, instructions, 345
Coloration of pileus, 34
Colouring matters, 85
Columella, 36, 313
Compound carpophore, 23
Sphaeriaceae, 213
Conidiojihore, 51
Conidium, 45
Coniomycetes, 259
Conjugating Fungi, 227
hyphae, 62
Conjugation in Peziza, 58
Constituents of Fungi, 84
Crested sporules, 275
Cup-shaped receptacle, 37
Cuticle of pileus, 33
Cystidia, 41, 121
Cyttaria, 186
Definition of Fungi, 1
Dematiaei, 278, 281
Destructiveness, 107
Development in Erysiphe, 55
of Eurotium, 56
Diatrype and Valsa, 211
Dichocarpism, 64, 286
Dimorphic Fungi, 64, 286
Discoid Fungi, 172
Discomycetes, 173
Disintegration by mycelium, 21
Distribution, 317, 324
Dothideaceae, 207
Dust Fungi, 259
Elaphomyces, 195
Empusa, development, 236
Endopliytal parasites, 75
Entomophthoraceae, 235
Entyloma, germination, 256
Epiphytal parasites, 74
Ergot and Claviceps, 202
of grains, 1 4
Erysipheae, 199
Evolution of Fungi, 111
Excipulum, 37
Fertilisation, 53
Fish-moulds, 232
Flies and Fungi, 152
Formation of zygospore, 17, 59
Fructification, 41
Fungi, census of, 319
and Lichens, 109
in general, 95
the Great Destroyer, 109
Fungus poisons, 86
INDEX
359
Gaping Fungi, 222
Gastromycetal carpophore, 26
Gastromycetes, 149
Geographical distribution, 32 -i
Germination of zoospores, 76
Gleba, 43
Globose carpophore, 29
Gloeosporiura, 272
Glossary, 353
Glycogen in Fungi, 87
Growth of Bacteria, 291
Haplophyllae, 132
Hereditary transmission, 81
Heteroecism, 245
Hymenium, 42, 102, 124
of Peziza, 44
Hymenomycetes, 126
Hypertrophied basidia, 122
Hyphal bodies, 18, 235
Hyphomycetes, 277
Hypocreaceae, 202
Hypodermeae, 78, 242
Hysteriaceae, 222
analogues, 268
Imperfect capsular Fungi, 259
Imperfect Fungi, 103
Introduction, 1
Lactiferous vessels, 25, 33
Lichens and Fungi, 3, 109
Lycoperdaceae, 154
Macrocysts, 58
Macronemeae, 280
Melanoonieae, 260, 271
]\Ielanconis and conidia, 67
Melasmia and Rhytisma, 65
Mesomycetes, 113
Metuloids, 42, 143
Microbes, 290
Micronemeae, 280
Microthyriaceae, 219
Microzyma, 297
Milky juice, 25, 33, 89
Morels, 178
Moulds and their conidia, 50, 277
Mucedines, 278
Mucor, development, 229
Mucoraceae, 228
Mushroom spawn, 9, 97
Mycelium, 9, 97
Mycomycetes, 113
Mycorhiza, 20
Myxomycetes, 304
Myxomycetes not animal, 19, 304
Nakkd hymenium, 41
Naked-spored Fungi, 102, 119
Nidulariaceae, 153
OiDiuM and Uncinula, 66
Old definitions, 96
Oocyst, 56
Oogonia, or female cells, 60
Oogonium and oosphere, 232
Organography, 7
Ostiolum, or mouth, 38
Paracysts, 58
Paraphyses, 42, 121, 168
Pathogenous microbes, 296
Perennial mycelium, 20
Peridiola, 46, 153
Peridium, 36
Perisporiaceae, 198
Perithecium, 38, 48, 198
Peronospora, development, 231
reproduction of, 77
Peronosporaceae, 230
Peziza sclerotia, 15, 180
Phalloideae, 151
Phosphorescence, 89
Phyeomycetes, 113, 227
Pileate receptacle, 31
Plasmodia, 305
Podaxis, structure of, 160
Polymorphism, 220
Polyphagus, development, 237
Polyporei, 135
Promycelial spores, 246, 254
Pseudoperidia, 48
Puccinia, life-history, 78
Puff-ball Fungi, 149
Pyrenomycetes, 197
Receptacle, 31
Resin in Pol3-porei, 88
Rhizomorpha, 12, 99
Root Fungi, 20
Rust Fungi, 242
Saccharomycetes, 290, 298
Saprolegnia, development, 233
Saprolegniaceae, 232
Saprophytes and parasites, 73
Schizomycetes, 290, 293
Schizophyllae, 130
Sclerotium, 14
Scolecite, 46, 57, 175
Section of Agaric, 32
Sessile pilei, 35
Sexuality, 54
in Achlya, 60
in Peronospora, 61
in Peziza, 176
Sexual reproduction, 229, 234
Simple Sphaeriaceae, 214
Slime Fungi, 304
36o
INTRODUCTION TO THE STUDY OF FUNGI
Smut Fungi, 251
Spermatia, 45
Sphaeriaceae, 205
analogues, 261
Sphaeiopsiiieae, 259
Sponuigiuiu, 50
Siiorc, naked, 45
division of Agaricus, 130
forms, 45
value, 345
Sporidium, 45
Sporule, 45
Stinkhorn Fungi, 151
Stipe, or stem, 25
Stylospore, 45
Stylosporous fructification, 49
Subterranean Fungi, 189
Suspensor and gamete, 229
Swarm-cells, 305
Table of groups, 117
Teleutospores, 242
Thallogens, 2
Thallophytes, 3
Tilletia, germination, 63, 255
Touchwood, 99
Trama, 32, 128
Tremelloid Fungi, 145
Truffles, 191
TuT)eraceae, 190
Tubercularia and Nectria, 204
Two forms of fruit, 69
Uredineae, 242
Ustilagines, 251
Varnished pilei, 88
Vegetative system, 11
Vinegar plant, 100
White moulds, 278
Xylaria and stroma, 206
Yeast Fungi, 298
Zoosi'OiiEs, ^ogonidia, 227
Zygomycetes, 229
Zygospore formation, 17, 59
Zygospores, 227
THE END
D. H. HILL i_^:rr?4RT
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