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Title: Fungi: Their Nature and Uses

Author: M. C. Cooke

Editor: M. J. Berkeley

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THE INTERNATIONAL SCIENTIFIC SERIES.

FUNGI:

THEIR

NATURE AND USES.

BY
M. C. COOKE, M.A., LL.D.

EDITED BY
The Rev. M. J. BERKELEY, M.A., F.L.S.

NEW YORK:
D. APPLETON AND COMPANY,
549 AND 551 BROADWAY.
1875.

PREFACE BY THE EDITOR.

As my name appears on the title-page of this volume, it is necessary that I
should exactly state what part I had in its preparation. I had no doubt
originally engaged to undertake the work myself; but finding, from
multiplicity of engagements and my uncertain health, that I could not
accomplish it satisfactorily, I thought the best course I could take was to
recommend Mr. Cooke to the publishers; a gentleman well known, not only
in this country, but in the United States. The whole of the work has
therefore been prepared by himself, the manuscript and proof sheets being
submitted to me from time to time, in which I merely suggested such
additions as seemed needful, subjoining occasionally a few notes. As the
work is intended for students, the author has had no hesitation in repeating
what has been stated in former chapters where it has been thought to prove
useful. I have no doubt that the same high character will justly apply to this
as to Mr. Cooke’s former publications, and especially to his “Handbook of
British Fungi.”
M. J. BERKELEY.
Sibbertoft,
November 23, 1874.

CONTENTS.

PAGE
I. Nature of Fungi. 1
II. Structure. 17
III Classification 64
IV. Uses. 82
V. Notable Phenomena. 105
VI. The Spore and Its Dissemination. 119

VII. Germination and Growth. 137
VIII. Sexual Reproduction. 163
IX. Polymorphism. 182
X. Influences and Effects. 209
XI. Habitats. 233
XII. Cultivation. 253
XIII. Geographical Distribution. 266
XIV. Collection and Preservation. 287
Index. 295

LIST OF ILLUSTRATIONS.

FIG. PAGE
1. Agaric in Process of Growth. 18
2. Section of Common Mushroom. 19
3. Sterile cells, Basidia, Cystidium, from Gomphidius. 21
4. Polyporus giganteus (reduced). 23
5. Hydnum repandum. 24
6. Calocera viscosa. 25
7. Tremella mesenterica. 25
8. Basidia and spores of Phallus. 28
9. Basidia and spores of Lycoperdon. 30
10. Threads of Trichia. 32
11. Arcyria incarnata, with portion of threads and spore. 33
12. Diachæa elegans. 34
13. Cyathus vernicosus. 34
14. Cyathus, Sporangia and spores. 35
15. Asterosporium Hoffmanni. 36

16. Barren Cysts and Pseudospores of Lecythea. 37
17. Coleosporium Tussilaginis. 37
18. Melampsora salicina, pseudospores of 37
19. Cystopus candidus, conidia of 38
20. Xenodochus carbonarius, pseudospore. 39
21. Phragmidium bulbosum, pseudospores. 39
22. Pseudospores of Puccinia. 40
23. Thecaphora hyalina, pseudospores. 41
24. Æcidium Berberidis, peridia of 41
25. Helminthosporium molle, threads and spores. 43
26. Acrothecium simplex. 44
27. Peronospora Arenariæ. 44
28. Polyactis cinerea. 45
29. Peziza Fuckeliana, with ascus and sporidia. 48
30. Penicillium chartarum. 50
31. Mucor mucedo, with sporangia. 51
32. Small portion of Botrytis Jonesii. 53
33. Section of cup of Ascobolus. 57
34. Asci, sporidia, and paraphyses of Ascobolus. 59
35. Perithecium of Sphæria. 61
36. Uncinula adunca, conceptacle with appendages. 62
37. Agaricus nudus. 66
38. Scleroderma vulgare, Fr. 69
39. Ceuthospora phacidioides. 70
40. Rhopalomyces candidus. 74
41. Mucor caninus. 75
42. Sphæria aquila, cluster of perithecia. 78
43. Morchella gigaspora, from Kashmir. 99
44. Cyttaria Gunnii 101
45. Spores of Agarics 121
46. Spores of Lactarius 121
46a. Spores of Gomphidius 122
47. Spores of Polyporus, Boletus, and Hydnum. 122
48. Diachea elegans, capellitium of 123

49. Spore of Hendersonia polycystis. 124
50. Spores of Dilophospora graminis. 124
51. Spores of Discosia. 124
52. Spore of Prosthemium betulinum. 124
53. Spore of Stegonosporium cellulosum. 125
54. Stylospores of Coryneum disciforme. 125
55. Spores of Asterosporium Hoffmanni. 125
56. Spores of Pestalozzia. 126
57. Bispora monilioides, concatenate spores 126
58. Pseudospores of Thecaphora hyalina. 127
59. Pseudospores of Puccinia. 127
60. Pseudospores of Triphragmium. 127
61. Pseudospores of Phragmidium bulbosum. 127
62. Winter spores of Melampsora salicina. 127
63. Spores of Helicocoryne. 129
64. Sporidium of Genea verrucosa. 130
65. Alveolate sporidium of Tuber. 130
66. Asci, sporidia, and paraphyses of Ascobolus. 131
67. Sporidium of Ostreichnion Americanum. 132
68. Ascus and sporidia of Hypocrea. 133
69. Sporidium of Sphæria ulnaspora. 133
70. Sporidia of Valsa profusa. 133
71. Sporidia of Massaria fœdans. 134
72. Sporidium of Melanconis bicornis. 134
73. Caudate sporidia of Sphæria fimiseda. 134
74. Sporidia of Valsa thelebola. 134
75. Sporidia of Valsa taleola. 135
76. Sporidium of Sporormia intermedia. 135
77. Asci and sporidia of Sphæria (Pleospora) herbarum. 135
78. Sporidium of Sphæria putaminum. 135
79. Basidia and spores of Exidia spiculosa. 139
80. Germinating spore and corpuscles of Dacrymyces. 140
81. Germination of Æcidium Euphorbia. 142
82. Germinating pseudospores of Coleosporium Sonchi. 144

83. Germinating pseudospore of Melampsora betulina. 144
84. Germinating pseudospore of Uromyce appendiculatus. 145
85. Germinating pseudospore of Puccinia Moliniæ. 146
86. Germinating pseudospore of Triphragmium Ulmariæ. 146
87. Germinating pseudospore of Phragmidium bulbosum. 147
88. Germinating pseudospores of Podisoma Juniperi. 148
89. Germinating pseudospore of Tilletia caries. 150
90. Pseudospore of Ustilago receptaculorum in germination, and
secondary spores in conjugation. 151
91. Conidia and zoospores of Cystopus candidus. 151
92. Resting spore of Cystopus candidus with zoospores. 152
93. Zygospores of Mucor phycomyces. 158
94. Sporidium of Ascobolus germinating. 161
95. Zygospore of Mucor. 165
96. Zygospore of Rhizopus in different stages. 167
97. Conjugation in Achlya racemosa. 169
98. Conjugation in Peronospora. 171
99. Antheridia and oogonium of Peronospora. 172
100. Conjugation in Peziza omphalodes. 175
100a. Formation of conceptacle in Erysiphe. 176
101. Tilletia caries with conjugating cells. 178
102. Aspergillus glaucus and Eurotium. 189
103. Erysiphe cichoracearum, receptacle and mycelium. 191
104. Twig with Tubercularia and Nectria. 193
105. Section of Tubercularia with conidia. 194
106. D. Nectria with Tubercularia, ascus and paraphyses. 195
107. Cells and pseudospores of Æcidium berberidis. 201
108. Cells and pseudospores of Æcidium graveolens. 201
109. Torrubia militaris on pupa of a moth. 243

FUNGI
THEIR NATURE, USES, INFLUENCES, ETC.

I.

NATURE OF FUNGI.

The most casual observer of Nature recognizes in almost every instance that
comes under his notice in every-day life, without the aid of logical
definition, the broad distinctions between an animal, a plant, and a stone. To
him, the old definition that an animal is possessed of life and locomotion, a
plant of life without locomotion, and a mineral deficient in both, seems to
be sufficient, until some day he travels beyond the circuit of diurnal routine,
and encounters a sponge or a zoophyte, which possesses only one of his
supposed attributes of animal life, but which he is assured is nevertheless a
member of the animal kingdom. Such an encounter usually perplexes the
neophyte at first, but rather than confess his generalizations to have been
too gross, he will tenaciously contend that the sponge must be a plant, until
the evidence produced is so strong that he is compelled to desert his
position, and seek refuge in the declaration that one kingdom runs into the
other so imperceptibly that no line of demarcation can be drawn between
them. Between these two extremes of broad distinction, and no distinction,
lies the ground occupied by the scientific student, who, whilst admitting
that logical definition fails in assigning briefly and tersely the bounds of the
three kingdoms, contends that such limits exist so positively, that the
universal scientific mind accepts the recognized limit without controversy
or contradiction.
In like manner, if one kingdom be made the subject of inquiry, the same
difficulties will arise. A flowering plant, as represented by a rose or a lily,
will be recognized as distinct from a fern, a seaweed, or a fungus. Yet there
are some flowering plants which, at first sight, and without examination,
simulate cryptogams, as, for example, many Balanophoræ, which the
unscientific would at once class with fungi. It is nevertheless true that even
the incipient botanist will accurately separate the phanerogams from the
cryptogams, and by means of a little more, but still elementary knowledge,
distribute the latter amongst ferns, mosses, fungi, lichens, and algæ, with
comparatively few exceptions. It is true that between fungi and lichens
there exists so close an affinity that difficulties arise, and doubts, and

disputations, regarding certain small groups or a few species; but these are
the exception, and not the rule. Botanists generally are agreed in
recognizing the five principal groups of Cryptogamia, as natural and
distinct. In proportion as we advance from comparison of members of the
three kingdoms, through that of the primary groups in one kingdom, to a
comparison of tribes, alliances, and orders, we shall require closer
observation, and more and more education of the eye to see, and the mind
to appreciate, relationships and distinctions.
We have already assumed that fungi are duly and universally admitted, as
plants, into the vegetable kingdom. But of this fact some have even
ventured to doubt. This doubt, however, has been confined to one order of
fungi, except, perhaps, amongst the most illiterate, although now the animal
nature of the Myxogastres has scarcely a serious advocate left. In this order
the early condition of the plant is pulpy and gelatinous, and consists of a
substance more allied to sarcode than cellulose. De Bary insinuated
affinities with Amœba,[A] whilst Tulasne affirmed that the outer coat in
some of these productions contained so much carbonate of lime that strong
effervescence took place on the application of sulphuric acid. Dr. Henry
Carter is well known as an old and experienced worker amongst amœboid
forms of animal life, and, when in Bombay, he devoted himself to the
examination of the Myxogastres in their early stage, and the result of his
examinations has been a firm conviction that there is no relationship
whatever between the Myxogastres and the lower forms of animal life. De
Bary has himself very much modified, if not wholly abandoned, the views
once propounded by him on this subject. When mature, and the dusty
spores, mixed with threads, sometimes spiral, are produced, the
Myxogastres are so evidently close allies of the Lycoperdons, or Puffballs,
as to leave no doubt of their affinities. It is scarcely necessary to remark that
the presence of zoospores is no proof of animal nature, for not only do they
occur in the white rust (Cystopus), and in such moulds as Peronospora,[B]
but are common in algæ, the vegetable nature of which has never been
disputed.
There is another equally important, but more complicated subject to
which we must allude in this connection. This is the probability of minute
fungi being developed without the intervention of germs, from certain
solutions. The observations of M. Trécul, in a paper laid before the French

Academy, have thus been summarized:—1. Yeast cells may be formed in
the must of beer without spores being previously sown. 2. Cells of the same
form as those of yeast, but with different contents, arise spontaneously in
simple solution of sugar, or to which a little tartrate of ammonia has been
added, and these cells are capable of producing fermentation in certain
liquids under favourable conditions. 3. The cells thus formed produce
Penicillium like the cells of yeast. 4. On the other hand, the spores of
Penicillium are capable of being transformed into yeast.[C] The
interpretation of this is, that the mould Penicillium may be produced from a
sugar solution by “spontaneous generation,” and without spore or germ of
any kind. The theory is, that a molecular mass which is developed in certain
solutions or infusions, may, under the influence of different circumstances,
produce either animalcules or fungi. “In all these cases, no kind of
animalcule or fungus is ever seen to originate from preexisting cells or
larger bodies, but always from molecules.”[D] The molecules are said to
form small masses, which soon melt together to constitute a globular body,
from which a process juts out on one side. These are the so-called Torulæ,
[E] which give off buds which are soon transformed into jointed tubes of
various diameters, terminating in rows of sporules, Penicillium, or capsules
containing numerous globular seeds, Aspergillus (sic).
This is but another mode of stating the same thing as above referred to by
M. Trécul, that certain cells, resembling yeast cells (Torula), are developed
spontaneously, and that these ultimately pass through the form of mould
called Penicillium to the more complex Mucor (which the writer evidently
has confounded with Aspergillus, unless he alludes to the ascigerous form
of Aspergillus, long known as Eurotium). From what is now known of the
polymorphism of fungi, there would be little difficulty in believing that
cells resembling yeast cells would develop into Penicillium, as they do in
fact in what is called the “vinegar plant,” and that the capsuliferous, or
higher condition of this mould may be a Mucor, in which the sporules are
produced in capsules. The difficulty arises earlier, in the supposed
spontaneous origination of yeast cells from molecules, which result from
the peculiar conditions of light, temperature, &c., in which certain solutions
are placed. It would be impossible to review all the arguments, or tabulate
all the experiments, which have been employed for and against this theory.
It could not be passed over in silence, since it has been one of the stirring
questions of the day. The great problem how to exclude all germs from the

solutions experimented upon, and to keep them excluded, lies at the
foundation of the theory. It must ever, as we think, be matter of doubt that
all germs were not excluded or destroyed, rather than one of belief that
forms known to be developed day by day from germs should under other
conditions originate spontaneously.
Fungi are veritably and unmistakably plants, of a low organization, it is
true, but still plants, developed from germs, somewhat analogous, but not
wholly homologous, to the seeds of higher orders. The process of
fertilization is still obscure, but facts are slowly and gradually
accumulating, so that we may hope at some not very distant period to
comprehend what as yet are little removed from hypotheses. Admitting that
fungi are independent plants, much more complex in their relations and
development than was formerly supposed, it will be expected that certain
forms should be comparatively permanent, that is, that they should
constitute good species. Here, also, efforts have been made to develop a
theory that there are no legitimate species amongst fungi, accepting the
terms as hitherto applied to flowering plants. In this, as in allied instances,
too hasty generalizations have been based on a few isolated facts, without
due comprehension of the true interpretation of such facts and phenomena.
Polymorphism will hereafter receive special illustration, but meantime it
may be well to state that, because some forms of fungi which have been
described, and which have borne distinct names as autonomous species, are
now proved to be only stages or conditions of other species, there is no
reason for concluding that no forms are autonomous, or that fungi which
appear and are developed in successive stages are not, in their entirety, good
species. Instead, therefore, of insinuating that there are no good species,
modern investigation tends rather to the establishment of good species, and
the elimination of those that are spurious. It is chiefly amongst the
microscopic species that polymorphism has been determined. In the larger
and fleshy fungi nothing has been discovered which can shake our faith in
the species described half a century, or more, ago. In the Agarics, for
instance, the forms seem to be as permanent and as distinct as in the
flowering plants. In fact, there is still no reason to dissent, except to a very
limited extent, from what was written before polymorphism was accredited,
that, “with a few exceptions only, it may without doubt be asserted that
more certain species do not exist in any part of the organized world than
amongst fungi. The same species constantly recur in the same places, and if

kinds not hitherto detected present themselves, they are either such as are
well known in other districts, or species which have been overlooked, and
which are found on better experience to be widely diffused. There is
nothing like chance about their characters or growth.”[F]
The parasitism of numerous minute species on living and growing plants
has its parallel even amongst phanerogams in the mistletoe and broom-rape
and similar species. Amongst fungi a large number are thus parasitic,
distorting, and in many cases ultimately destroying, their host, burrowing
within the tissues, and causing rust and smut in corn and grasses, or even
more destructive and injurious in such moulds as those of the potato disease
and its allies. A still larger number of fungi are developed from decayed or
decaying vegetable matter. These are found in winter on dead leaves, twigs,
branches, rotten wood, the remains of herbaceous plants, and soil largely
charged with disintegrated vegetables. As soon as a plant begins to decay it
becomes the source of a new vegetation, which hastens its destruction, and
a new cycle of life commences. In these instances, whether parasitic on
living plants or developed on dead ones, the source is still vegetable. But
this is not always the case, so that it cannot be predicated that fungi are
wholly epiphytal. Some species are always found on animal matter, leather,
horn, bone, &c., and some affect such unpromising substances as minerals,
from which it would be supposed that no nourishment could be obtained,
not only hard gravel stones, fragments of rock, but also metals, such as iron
and lead, of which more may be said when we come to treat of the habitats
of fungi. Although in general terms fungi may be described as
“hysterophytal or epiphytal mycetals deriving nourishment by means of a
mycelium from the matrix,”[G] there are exceptions to this rule with which
the majority accord.
Of the fungi found on animal substances, none are more extraordinary
than those species which attack insects. The white mould which in autumn
proves so destructive to the common house-fly may for the present be
omitted, as it is probably a condition of one of the Saprolegniei, which
some authors include with fungi, and others with algæ. Wasps, spiders,
moths, and butterflies become enveloped in a kind of mould named Isaria,
which constitutes the conidia of Torrubia, a genus of club-shaped Sphæriæ
afterwards developed. Some species of Isaria and Torrubia also affect the
larvæ and pupæ of moths and butterflies, converting the whole interior into

a mass of mycelium, and fructifying in a clavate head. It has been subject
for discussion whether in such instances the fungus commenced its
development during the life of the insect, and thus hastened its death, or
whether it resulted after death, and was subsequent to the commencement
of decay.[H] The position in which certain large moths are found standing on
leaves when infested with Isaria resembles so closely that of the house-fly
when succumbing to Sporendonema Muscæ, would lead to the conclusion
that certainly in some cases the insect was attacked by the fungus whilst
still living; whilst in the case of buried caterpillars, such as the New
Zealand or British Hepialus, it is difficult to decide. Whether in life or death
in these instances, it is clear that the silk-worm disease Muscardine attacks
the living insect, and causes death. In the case of the Guêpes végétantes, the
wasp is said to fly about with the fungus partially developed.
In all fungi we may recognize a vegetative and a reproductive system:
sometimes the first only becomes developed, and then the fungus is
imperfect, and sometimes the latter is far more prominent than the former.
There is usually an agglomeration of delicate threads, either jointed or not,
which are somewhat analogous to the roots of higher plants. These delicate
threads permeate the tissues of plants attacked by parasitic fungi, or they
run over dead leaves forming whitened patches, formerly bearing the name
of Himantia, but really the mycelium of some species of Marasmius. If
checked or disturbed, the process stops here, and only a mycelium of
interwoven threads is produced. In this condition the mycelium of one
species so much resembles that of another, that no accurate determination
can be made. If the process goes on, this mycelium gives rise to the stem
and cap of an agaricoid fungus, completing the vegetative system. This in
turn gives origin to a spore-bearing surface, and ultimately the fruit is
formed, and then the fungus is complete; no fungus can be regarded as
perfect or complete without its reproductive system being developed. In
some this is very simple, in others it is as complex. In many of the moulds
we have miniature representatives of higher plants in the mycelium or roots,
stem, branches, and at length capsules bearing sporidia, which correspond
to seeds. It is true that leaves are absent, but these are sometimes
compensated by lateral processes or abortive branchlets. A tuft of mould is
in miniature a forest of trees. Although such a definition may be deemed
more poetic than accurate, more figurative than literal, yet few could
believe in the marvellous beauty of a tuft of mould if they never saw it as

exhibited under the microscope. In such a condition no doubt could be
entertained of its vegetable character. But there is a lower phase in which
these plants are sometimes encountered; they may consist only of single
cells, or strings of cells, or threads of simple structure floating in fluids. In
such conditions only the vegetative system is probably developed, and that
imperfectly, yet some have ventured to give names to isolated cells, or
strings of cells, or threads of mycelium, which really in themselves possess
none of the elements of correct classification—the vegetative system, even,
being imperfect, and consequently the reproductive is absent. As already
observed, no fungus is perfect without fruit of some kind, and the
peculiarities of structure and development of fruit form one of the most
important elements in classification. To attempt, therefore, to give names to
such imperfect fragments of undeveloped plants is almost as absurd as to
name a flowering plant from a stray fragment of a root-fibril accidentally
cast out of the ground—nay, even worse, for identification would probably
be easier. It is well to protest at all times against attempts to push science to
the verge of absurdity; and such must be the verdict upon endeavours to
determine positively such incomplete organisms as floating cells, or hyaline
threads which may belong to any one of fifty species of moulds, or after all
to an alga. This leads us to remark, in passing, that there are forms and
conditions under which fungi may be found when, fructification being
absent—that is, the vegetative system alone developed—they approximate
so closely to algæ that it is almost impossible to say to which group the
organisms belong.
Finally, it is a great characteristic of fungi in general that they are very
rapid in growth, and rapid in decay. In a night a puffball will grow
prodigiously, and in the same short period a mass of paste may be covered
with mould. In a few hours a gelatinous mass of Reticularia will pass into a
bladder of dust, or a Coprinus will be dripping into decay. Remembering
this, mycophagists will take note that a fleshy fungus which may be good
eating at noon may undergo such changes in a few hours as to be anything
but good eating at night. Many instances have been recorded of the rapidity
of growth in fungi; it may also be accepted as an axiom that they are, in
many instances, equally as rapid in decay.
The affinity between lichens and fungi has long been recognized to its
full and legitimate extent by lichenologists and mycologists.[I] In the

“Introduction to Cryptogamic Botany,” it was proposed to unite them in one
alliance, under the name of Mycetales, in the same manner as the late Dr.
Lindley had united allied orders under alliances in his “Vegetable
Kingdom;” but, beyond this, there was no predisposition towards the theory
since propounded, and which, like all new theories, has collected a small
but zealous circle of adherents. It will be necessary briefly to summarize
this theory and the arguments by which it is supported and opposed,
inasmuch as it is intimately connected with our subject.
As recently as 1868, Professor Schwendener first propounded his views,
[J] and then briefly and vaguely, that all and every individual lichen was but
an algal, which had collected about it a parasitic fungal growth, and that
those peculiar bodies which, under the name of gonidia, were considered as
special organs of lichens, were only imprisoned algæ. In language which
the Rev. J. M. Crombie[K] describes as “pictorial,” this author gave the
general conclusion at which he had arrived, as follows:—“As the result of
my researches, all these growths are not simple plants, not individuals in the
usual sense of the term; they are rather colonies, which consist of hundreds
and thousands of individuals, of which, however, only one acts as master,
while the others, in perpetual captivity, provide nourishment for themselves
and their master. This master is a fungus of the order Ascomycetes, a
parasite which is accustomed to live upon the work of others; its slaves are
green algæ, which it has sought out, or indeed caught hold of, and forced
into its service. It surrounds them, as a spider does its prey, with a fibrous
net of narrow meshes, which is gradually converted into an impenetrable
covering. While, however, the spider sucks its prey and leaves it lying dead,
the fungus incites the algæ taken in its net to more rapid activity; nay, to
more vigorous increase.” This hypothesis, ushered upon the world with all
the prestige of the Professor’s name, was not long in meeting with
adherents, and the cardinal points insisted upon were—1st. That the generic
relationship of the coloured “gonidia” to the colourless filaments which
compose the lichen thallus, had only been assumed, and not proved; 2nd.
That the membrane of the gonidia was chemically different from the
membrane of the other tissues, inasmuch as the first had a reaction
corresponding to that of algæ, whilst the second had that of fungi; 3rd. That
the different forms and varieties of gonidia corresponded with parallel types
of algæ; 4th. That as the germination of the spore had not been followed
further than the development of a hypothallus, it might be accounted for by

the absence of the essential algal on which the new organism should
become parasitic; 5th. That there is a striking correspondence between the
development of the fruit in lichens and in some of the sporidiiferous fungi
(Pyrenomycetes).
These five points have been combated incessantly by lichenologists, who
would really be supposed by ordinary minds to be the most practically
acquainted with the structure and development of these plants, in opposition
to the theorists. It is a fact which should have some weight, that no
lichenologist of repute has as yet accepted the theory. In 1873 Dr. E.
Bornet[L] came to the aid of Schwendener, and almost exhausted the
subject, but failed to convince either the practised lichenologist or
mycologist. The two great points sought to be established are these, that
what we call lichens are compound organisms, not simple, independent
vegetable entities; and that this compound organism consists of unicellular
algæ, with a fungus parasitic upon them. The coloured gonidia which are
found in the substance, or thallus of lichens, are the supposed algæ; and the
cellular structure which surrounds, encloses, and imprisons the gonidia is
the parasitic fungus, which is parasitic on something infinitely smaller than
itself, and which it entirely and absolutely isolates from all external
influences.
Dr. Bornet believed himself to have established that every gonidium of a
lichen may be referred to a species of algæ, and that the connection between
the hypha and gonidia is of such a nature as to exclude all possibility of the
one organ being produced by the other. This he thinks is the only way in
which it can be accounted for that the gonidia of diverse lichens should be
almost identical.
Dr. Nylander, in referring to this hypothesis of an imprisoned algal,[M]
writes: “The absurdity of such an hypothesis is evident from the very
consideration that it cannot be the case that an organ (gonidia) should at the
same time be a parasite on the body of which it exercises vital functions; for
with equal propriety it might be contended that the liver or the spleen
constitutes parasites of the mammiferæ. Parasite existence is autonomous,
living upon a foreign body, of which nature prohibits it from being at the
same time an organ. This is an elementary axiom of general physiology. But
observation directly made teaches that the green matter originally arises
within the primary chlorophyll- or phycochrom-bearing cellule, and

consequently is not intruded from any external quarter, nor arises in any
way from any parasitism of any kind. The cellule at first is observed to be
empty, and then, by the aid of secretion, green matter is gradually produced
in the cavity and assumes a definite form. It can, therefore, be very easily
and evidently demonstrated that the origin of green matter in lichens is
entirely the same as in other plants.” On another occasion, and in another
place, the same eminent lichenologist remarks,[N] as to the supposed algoid
nature of gonidia—“that such an unnatural existence as they would thus
pass, enclosed in a prison and deprived of all autonomous liberty, is not at
all consonant with the manner of existence of the other algæ, and that it has
no parallel in nature, for nothing physiologically analogous occurs
anywhere else. Krempelhuber has argued that there are no conclusive
reasons against the assumption that the lichen-gonidia may be self-
developed organs of the lichen proper rather than algæ, and that these
gonidia can continue to vegetate separately, and so be mistaken for
unicellular algæ.” In this Th. Fries seems substantially to concur. But there
is one strong argument, or rather a repetition of an argument already cited,
placed in a much stronger light, which is employed by Nylander in the
following words:—“So far are what are called algæ, according to the turbid
hypothesis of Schwendener, from constituting true algæ, that on the
contrary it may be affirmed that they have a lichenose nature, whence it
follows that these pseudo-algæ are in a systematic arrangement to be
referred rather to the lichens, and that the class of algæ hitherto so vaguely
limited should be circumscribed by new and truer limits.”
As to another phase in this question, there are, as Krempelhuber remarks,
species of lichens which in many countries do not fructify, and whose
propagation can only be carried on by means of the soredia, and the hyphæ
of such could in themselves alone no more serve for propagation than the
hyphæ from the pileus or stalk of an Agaric, while it is highly improbable
that they could acquire this faculty by interposition of a foreign algal. On
the other hand he argues: “It is much more conformable to nature that the
gonidia, as self-developed organs of the lichens, should, like the spores,
enable the hyphæ proceeding from them to propagate the individual.”[O]
A case in point has been adduced[P] in which gonidia were produced by
the hypha, and the genus Emericella,[Q] which is allied to Husseia in the
Trichogastres, shows a structure in the stem exactly resembling Palmella

botryoides of Greville, and to what occurs in Synalyssa. Emericella, with
one or two other genera, must, however, be considered as connecting
Trichogastres with lichens, and the question cannot be considered as
satisfactorily decided till a series of experiments has been made on the
germination of lichen spores and their relation to free algæ considered
identical with gonidia. Mr. Thwaites was the first to point out[R] the relation
of the gonidia in the different sections of lichens to different types of
supposed algæ. The question cannot be settled by mere à priori notions. It
is, perhaps, worthy of remark that in Chionyphe Carteri the threads grow
over the cysts exactly as the hypha of lichens is represented as growing over
the gonidia.
Recently, Dr. Thwaites has communicated his views on one phase of this
controversy,[S] which will serve to illustrate the question as seen from the
mycological side. As is well known, this writer has had considerable
experience in the study of the anatomy and physiology of all the lower
cryptogamia, and any suggestion of his on such a subject will at least
commend itself to a patient consideration.
“According to our experience,” he writes, “I think parasitic fungi
invariably produce a sad effect upon the tissues they fix themselves upon or
in. These tissues become pale in colour, and in every respect sickly in
appearance. But who has ever seen the gonidia of lichens the worse for
having the ‘hypha’ growing amongst them? These gonidia are always in the
plumpest state, and with the freshest, healthiest colour possible. Cannot it
enter into the heads of these most patient and excellent observers, that a
cryptogamic plant may have two kinds of tissue growing side by side,
without the necessity of one being parasitic upon the other, just as one of
the higher plants may have half a dozen kinds of tissue making up its
organization? The beautifully symmetrical growth of the same lichens has
seemed to me a sufficient argument against one portion being parasitic upon
another, but when we see all harmony and robust health, the idea that one
portion is subsisting parasitically upon another appears to me to be a perfect
absurdity.”
It appears to us that a great deal of confusion and a large number of
errors which creep into our modern generalizations and hypotheses, may be
traced to the acceptance of analogies for identities. How many cases of
mistaken identity has the improvement of microscopes revealed during the

past quarter of a century. This should at least serve as a caution for the
future.
Apart, however, from the “gonidia,” whatever they may be, is the
remainder of the lichen a genuine fungus? Nylander writes, “The
anatomical filamentose elements of lichens are distinguished by various
characters from the hyphæ of fungi. They are firmer, elastic, and at once
present themselves in the texture of lichens. On the other hand, the hyphæ
of fungi are very soft, they possess a thin wall, and are not at all gelatinous,
while they are immediately dissolved by the application of hydrate of
potash, &c.”[T]
Our own experience is somewhat to the effect, that there are some few
lichens which are doubtful as to whether they are fungi or lichens, but, in by
far the majority of cases, there is not the slightest difficulty in determining,
from the peculiar firmness and elasticity of the tissues, minute peculiarities
which the practised hand can detect rather than describe, and even the
general character of the fruit that they differ materially from, though closely
allied to fungi. We have only experience to guide us in these matters, but
that is something, and we have no experience in fungi of anything like a
Cladonia, however much it may resemble a Torrubia or Clavaria. We have
Pezizæ with a subiculum in the section Tapesia, but the veriest tyro would
not confound them with species of Parmelia. It is true that a great number
of lichens, at first sight, and casually, resemble species of the Hysteriacei,
but it is no less strange than true, that lichenologists and mycologists know
their own sufficiently not to commit depredations on each other.
Contributions are daily being made to this controversy, and already the
principal arguments on both sides have appeared in an English dress,[U]
hence it will be unnecessary to repeat those which are modifications only of
the views already stated, our own conclusions being capable of a very brief
summary: that lichens and fungi are closely related the one to the other, but
that they are not identical; that the “gonidia” of lichens are part of the
lichen-organization, and consequently are not algæ, or any introduced
bodies; that there is no parasitism; and that the lichen thallus, exclusive of
gonidia, is wholly unknown amongst fungi.
The Rev. J. M. Crombie has therefore our sympathies in the remark with
which his summary of the gonidia controversy closes, in which he

characterizes it as a “sensational romance of lichenology,” of the “unnatural
union between a captive algal damsel and a tyrant fungal master.”
[A]
De Bary, “Des Myxomycètes,” in “Ann. des Sci. Nat.” 4 sér. xi. p. 153; “Bot. Zeit.”
xvi. p. 357. De Bary’s views are controverted by M. Wigand in “Ann. des Sci. Nat.” 4
sér. (Bot.) xvi. p. 255, &c.
[B]
De Bary, “Recherches sur le Developpement de quelques Champignons Parasites,” in
“Ann. des Sci. Nat.” 4 sér. (Bot.) xx. p. 5.
[C]
“Popular Science Review,” vol. viii. p. 96.
[D]
Dr. J. H. Bennett “On the Molecular Origin of Infusoria,” p. 56.
[E]
They have, however, no close relation with real Torulæ, such as T. monilioides, &c.—
Cooke’s Handbook, p. 477.
[F]
Berkeley’s “Outlines of British Fungology,” p. 24.
[G]
Berkeley’s “Introduction to Cryptogamic Botany,” p. 235.
[H]
Gray, “Notices of Insects which form the Basis of Fungoid Parasites.”
[I]
On the relation or connection between fungi and lichens, H. C. Sorby has some
pertinent remarks in his communication to the Royal Society on “Comparative
Vegetable Chromatology” (Proceedings Royal Society, vol. xxi. 1873, p. 479), as one
result of his spectroscopic examinations. He says, “Such being the relations between
the organs of reproduction and the foliage, it is to some extent possible to understand
the connection between parasitic plants like fungi, which do not derive their support
from the constructive energy of their fronds, and those which are self-supporting and
possess true fronds. In the highest classes of plants the flowers are connected with the
leaves, more especially by means of xanthophyll and yellow xanthophyll, whereas in
the case of lichens the apothecia contain very little, if any, of those substances, but a
large amount of the lichenoxanthines so characteristic of the class. Looking upon
fungi from this chromatological point of view, they bear something like the same
relation to lichens that the petals of a leafless parasitic plant would bear to the foliage
of one of normal character—that is to say, they are, as it were, the coloured organs of
reproduction of parasitic plants of a type closely approaching that of lichens, which,

of course, is in very close, if not in absolute agreement with the conclusions drawn by
botanists from entirely different data.”
[J]
Schwendener, “Untersuchungen über den Flechtenthallus.”
[K]
Crombie (J. M.) “On the Lichen-Gonidia Question,” in “Popular Science Review” for
July, 1874.
[L]
Bornet, (E.), “Recherches sur les Gonidies des Lichens,” in “Ann. des Sci. Nat.”
1873, 5 sér. vol. xvii.
[M]
Nylander, “On the Algo-Lichen Hypothesis,” &c., in “Grevillea,” vol. ii. (1874), No.
22, p. 146.
[N]
In Regensburg “Flora,” 1870, p. 92.
[O]
Rev. J. M. Crombie, in “Popular Science Review,” July, 1874.
[P]
Berkeley’s “Introduction to Cryptogamic Botany,” p. 373, fig. 78a.
[Q]
Berkeley’s “Introduction,” p. 341, fig. 76.
[R]
“Annals and Magazine of Natural History,” April, 1849.
[S]
In “Gardener’s Chronicle” for 1873, p. 1341.
[T]
“Grevillea,” vol ii. p. 147, in note.
[U]
W. Archer, in “Quart. Journ. Micr. Sci.” vol. xiii. p. 217; vol. xiv. p. 115. Translation
of Schwendener’s “Nature of the Gonidia of Lichens,” in same journal, vol. xiii. p.
235.

II.
STRUCTURE.

Without some knowledge of the structure of fungi, it is scarcely possible to
comprehend the principles of classification, or to appreciate the curious
phenomena of polymorphism. Yet there is so great a variety in the structure
of the different groups, that this subject cannot be compressed within a few
paragraphs, neither do we think that this would be desired if practicable,
seeing that the anatomy and physiology of plants is, in itself, sufficiently
important and interesting to warrant a rather extended and explicit survey.
In order to impart as much practical utility as possible to this chapter, it
seems advisable to treat some of the most important and typical orders and
suborders separately, giving prominence to the features which are chiefly
characteristic of those sections, following the order of systematists as much
as possible, whilst endeavouring to render each section independent to a
considerable extent, and complete in itself. Some groups naturally present
more noteworthy features than others, and will consequently seem to
receive more than their proportional share of attention, but this seeming
inequality could scarcely have been avoided, inasmuch as hitherto some
groups have been more closely investigated than others, are more intimately
associated with other questions, or are more readily and satisfactorily
examined under different aspects of their life-history.

Fig. 1.—Agaric in Process of Growth.

Agaricini.—For the structure that prevails in the order to which the
mushroom belongs, an examination of that species will be almost sufficient.
Here we shall at once recognize three distinct parts requiring elucidation,
viz. the rooting slender fibres that traverse the soil, and termed the
mycelium, or spawn, the stem and cap or pileus, which together constitute
what is called the hymenophore, and the plates or gills on the under surface
of the cap, which bear the hymenium. The earliest condition in which the
mushroom can be recognized as a vegetable entity is in that of the “spawn”
or mycelium, which is essentially an agglomeration of vegetating spores. Its
normal form is that of branched, slender, entangled, anastomosing, hyaline
threads. At certain privileged points of the mycelium, the threads seem to be
aggregated, and become centres of vertical extension. At first only a small
nearly globose budding, like a grain of mustard seed, is visible, but this
afterwards increases rapidly, and other similar buddings or swellings appear
at the base.[A] These are the young hymenophore. As it pushes through the
soil, it gradually loses its globose form, becomes more or less elongated,
and in this condition a longitudinal section shows the position of the future
gills in a pair of opposite crescent-shaped darker-coloured spots near the
apex. The dermal membrane, or outer skin, seems to be continuous over the
stem and the globose head. At present, there is no external evidence of an
expanded pileus and gills; a longitudinal section at this stage shows that the
gills are being developed, that the pileus is assuming its cap-like form, that
the membrane stretching from the stem to the edge of the young pileus is
separating from the edge of the gills, and forming a veil, which, in course of

time, will separate below and leave the gills exposed. When, therefore, the
mushroom has arrived almost at maturity, the pileus expands, and in this act
the veil is torn away from the margin of the cap, and remains for a time like
a collar around the stem. Fragments of the veil often remain attached to the
margin of the pileus, and the collar adherent to the stem falls back, and
thenceforth is known as the annulus or ring. We have in this stage the fully-
developed hymenophore,—the stem with its ring, supporting an expanded
cap or pileus, with gills on the under surface bearing the hymenium.[B] A
longitudinal section cut through the pileus and down the stem, gives the
best notion of the arrangement of the parts, and their relation to the whole.
By this means it will be seen that the pileus is continuous with the stem,
that the substance of the pileus descends into the gills, and that relatively
the substance of the stem is more fibrous than that of the pileus. In the
common mushroom the ring is very distinct surrounding the stem, a little
above the middle, like a collar. In some Agarics the ring is very fugacious,
or absent altogether. The form of the gills, their mode of attachment to the
stem, their colour, and more especially the colour of the spores, are all very
important features to be attended to in the discrimination of species, since
they vary in different species. The whole substance of the Agaric is cellular.
A longitudinal slice from the stem will exhibit under the microscope
delicate tubular cells, the general direction of which is lengthwise, with
lateral branches, the whole interlacing so intimately that it is difficult to
trace any individual thread very far in its course. It will be evident that the
structure is less compact as it approaches the centre of the stem, which in
many species is hollow. The hymenium is the spore-bearing surface, which
is exposed or naked, and spread over the gills. These plates are covered on
all sides with a delicate membrane, upon which the reproductive organs are
developed. If it were possible to remove this membrane in one entire piece
and spread it out flat, it would cover an immense surface, as compared with
the size of the pileus, for it is plaited or folded like a lady’s fan over the
whole of the gill-plates, or lamellæ, of the fungus.[C] If the stem of a
mushroom be cut off close to the gills, and the cap laid upon a sheet of
paper, with the gills downwards, and left there for a few hours, when
removed a number of dark radiating lines will be deposited upon the paper,
each line corresponding with the interstices between one pair of gills. These
lines are made up of spores which have fallen from the hymenium, and, if
placed under the microscope, their character will at once be made evident.

If a fragment of the hymenium be also submitted to a similar examination,
it will be found that the whole surface is studded with spores. The first
peculiarity which will be observed is, that these spores are almost uniformly
in groups of four together. The next feature to be observed is, that each
spore is borne upon a slender stalk or sterigma, and that four of these
sterigmata proceed from the apex of a thicker projection, from the
hymenium, called a basidium, each basidium being the supporter of four
sterigmata, and each sterigma of a spore.[D] A closer examination of the
hymenium will reveal the fact that the basidia are accompanied by other
bodies, often larger, but without sterigmata or spores; these have been
termed cystidia, and their structure and functions have been the subject of
much controversy.[E] Both kinds of bodies are produced on the hymenium
of most, if not all, the Agaricini.
The basidia are usually expanded
upwards, so as to have more or less of a
clavate form, surmounted by four slender
points, or tubular processes, each
supporting a spore; the contents of these
cells are granular, mixed apparently with
oleaginous particles, which communicate
through the slender tubes of the spicules
with the interior of the spores. Corda states
that, although only one spore is produced at
a time on each sporophore, when this falls
away others are produced in succession for
a limited period. As the spores approach
maturity, the connection between their
contents and the contents of the basidia Fig. 2.—Section of Common Mushroom.
diminishes and ultimately ceases. When the
basidium which bears mature spores is still well charged with granular
matter, it may be presumed that the production of a second or third series of
spores is quite possible. Basidia exhausted entirely of their contents, and
which have become quite hyaline, may often be observed.
The cystidia are usually larger than the basidia, varying in size and form
in different species. They present the appearance of large sterile cells,
attenuated upwards, sometimes into a slender neck. Corda was of opinion

that these were male organs, and gave them the
name of pollinaires. Hoffmann has also
described[F] both these organs under the names of
pollinaria and spermatia, but does not appear to
recognize in them the sexual elements which
those names would indicate; whilst de Seynes
suggests that the cystidia are only organs returned
to vegetative functions by a sort of hypertrophy
of the basidia.[G] This view seems to be supported
by the fact that, in the section Pluteus and some
others, the cystidia are surmounted by short horns
resembling sterigmata. Hoffmann has also
indicated[H] the passage of cystidia into basidia.
The evidence seems to be in favour of regarding Fig. 3.—a. Sterile cells. b.
the cystidia as barren conditions of basidia. There Basidia. c. Cystidium. From
Gomphidius (de Seynes).
are to be found upon the hymenium of Agarics a
third kind of elongated cells, called by Corda[I] basilary cells, and by
Hoffmann “sterile cells,” which are either equal in size or smaller than the
basidia, with which also their structure agrees, excepting in the
development of spicules. These are the “proper cells of the hymenium” of
Léveillé, and are simply the terminal cells of the gill structure—cells which,
under vigorous conditions, might be developed into basidia, but which are
commonly arrested in their development. As suggested by de Seynes, the
hymenium seems to be reduced to great simplicity, “one sole and self-same
organ is the basis of it; according as it experiences an arrest of development,
as it grows and fructifies, or as it becomes hypertrophied, it gives us a
paraphyse, a basidium, or a cystidium—in other terms, atrophied basidium,
normal basidium and hypertrophied basidium; these are the three elements
which form the hymenium.”[J]
The only reproductive organs hitherto demonstrated in Agarics are the
spores, or, as sometimes called, from their method of production,
basidiospores.[K] These are at first colourless, but afterwards acquire the
colour peculiar to the species. In size and form they are, within certain
limits, exceedingly variable, although form and size are tolerably constant
in the same species. At first all are globose; as they mature, the majority are
ovoid or elliptic; some are fusiform, with regularly attenuated extremities.
In Hygrophorus they are rather irregular, reniform, or compressed in the

middle. Sometimes the external surface is rough with more or less
projecting warts. Some mycologists are of opinion that the covering of the
spore is double, consisting of an exospore and an endospore, the latter being
very fine and delicate. In other orders the double coating of the spore has
been demonstrated. When the spore is coloured, the external membrane
alone appears to possess colour, the endospore being constantly hyaline. It
may be added here, that in this order the spore is simple and unicellular. In
Lactarius and Russula the trama, or inner substance, is vesicular. True latex
vessels occur occasionally in Agaricus, though not filled with milk as in
Lactarius.
Polyporei.—In this order the gill plates are
replaced by tubes or pores, the interior of which
is lined by the hymenium; indications of this
structure having already been exhibited in some
of the lower Agaricini. In many cases the stem is
suppressed. The substance is fleshy in Boletus,
but in Polyporus the greater number of species
are leathery or corky, and more persistent. The
basidia, spicules, and quaternate spores agree
with those of Agaricini.[L] In fact there are no
features of importance which relate to the
hymenium in any order of Hymenomycetes (the Fig. 4.—Polyporus giganteus
Tremellini excepted) differing from the same (reduced).

organ in Agaricini, unless it be the absence of cystidia.
Hydnei.—Instead of pores, in this order
the hymenium is spread over the surface of
spines, prickles, or warts.[M]
Auricularini.—The hymenium is more
or less even, and in—
Clavariei the whole fungus is club-
shaped, or more or less intricately
branched, with the hymenium covering the
Fig. 5.—Hydnum repandum. outer surface.
Tremellini.—In this order we have a great departure from the character
of the substance, external appearance, and internal structure of the other

orders in this family. Here we have a gelatinous
substance, and the form is lobed, folded,
convolute, often resembling the brain of some
animal. The internal structure has been specially
illustrated by M. Tulasne,[N] through the common
species, Tremella mesenterica. This latter is of a
fine golden yellow colour, and rather large size. It
is uniformly composed throughout of a colourless
mucilage, with no appreciable texture, in which
are distributed very fine, diversely branched and Fig. 6.—Calocera viscosa.
anastomosing filaments. Towards the surface, the
ultimate branches of this filamentous network give birth, both at their
summits and laterally, to globular cells, which acquire a comparatively large
size. These cells are filled with a protoplasm, to which the plant owes its
orange colour. When they have attained their normal dimensions, they
elongate at the summit into two, three, or four distinct, thick, obtuse tubes,
into which the protoplasm gradually passes. The development of these tubes
is unequal and not simultaneous, so that one will often attain its full
dimensions, equal, perhaps, to three or four times the diameter of the
generative cell, whilst the others are only just appearing. By degrees, as
each tube attains its full size, it is attenuated into a fine point, the extremity
of which swells into a spheroidal cell, which ultimately becomes a spore.
Sometimes these tubes, or spicules, send out one or two lateral branches,
each terminated by a spore. These spores (about ·006 to ·008 mm. diameter)
are smooth, and deposit themselves, like a fine white dust, on the surface of
the Tremella and on its matrix. M. Léveillé[O] was of opinion that the
basidia of the Tremellini were monosporous, whilst M. Tulasne has
demonstrated that they are habitually tetrasporous, as in other of the
Hymenomycetes. Although agreeing in this, they differ in other features,
especially in the globose form of the basidia, mode of production of the
spicules, and, finally, the division of the basidia into two, three, or four cells
by septa which cut each other in their axis. This division precedes the
growth of the spicules. It is not rare to see these cells, formed at the expense
of an unilocular basidium, become partly isolated from each other; in
certain cases they seem to have separated very early, they then become
larger than usual, and are grouped on the same filament so as to represent a

kind of buds. This phenomenon usually takes place below the level of the
fertile cells, at a certain depth in the mucous tissue of the Tremella.
Besides the reproductive system here
described, Tulasne also made known the
existence of a series of filaments which produce
spermatia. These filaments are often scattered and
confused with those which produce the basidia,
and not distinguishable from them in size or any
other apparent characteristic, except the manner
in which their extremities are branched in order
to produce the spermatia. At other times the
spermatia-bearing surface covers exclusively
certain portions of the fungus, especially the
inferior lobes, imparting thereto a very bright Fig. 7.—Tremella mesenterica.
orange colour, which is communicated by the
layer of spermatia, unmixed with spores. These spots retain their bright
colour, while the remainder of the plant becomes pale, or covered with a
white dust. The spermatia are very small, spherical, and smooth, scarcely
equalling ·002 mm. They are sessile, sometimes solitary, sometimes three or
four together, on the slightly swollen extremities of certain filaments of the
weft of the fungus.[P] Tulasne found it impossible to make these corpuscles
germinate, and in all essential particulars they agreed with the spermatia
found in ascomycetous fungi.
In the genus Dacrymyces, the same observer found the structure to have
great affinity with that of Tremella. The spores in the species examined
were of a different form, being oblong, very obtuse, slightly curved (·013 -
·019 × ·004 - ·006 mm.), at first unilocular, but afterwards triseptate. The
basidia are cylindrical or clavate, filled with coloured granular matter; each
of these bifurcates at the summit, and gradually elongates into two very
open branches, which are attenuated above, and ultimately each is crowned
by a spore. There are to be found also in the species of this genus globose
bodies, designated “sporidioles” by M. Léveillé, which Tulasne took
considerable care to trace to their source. He thus accounts for them:—Each
of the cells of the spore emits exteriorly one or several of these corpuscles,
supported on very short and very slender pedicels, which remain after the
corpuscles are detached from them, new corpuscles succeeding the first as

long as there remains any plastic matter within the spore. The pedicels are
not all on the same plane; they are often implanted all on the same, and
oftenest on the convex side of the reproductive body. These corpuscles,
though placed under the most favourable conditions, never gave the least
sign of vegetation, and Tulasne concludes that they are spermatia,
analogous to those produced in Tremella. The spores which produce
spermatia are not at all apt to germinate, whilst those which did not produce
spermatia germinated freely. Hence it would appear that, although all spores
seem to be perfectly identical, they have not all the same function. The
same observer detected also amongst specimens of the Dacrymyces some of
a darker and reddish tint, always bare of spores or spermatia on the surface,
and these presented a somewhat different structure. Where the tissue had
turned red it was sterile, the constituent filaments, ordinarily colourless, and
almost empty of solid matter, were filled with a highly-coloured
protoplasm; they were of less tenuity, more irregularly thick, and instead of
only rarely presenting partitions, and remaining continuous, as in other
parts of the plant, were parcelled out into an infinity of straight or curved
pieces, angular and of irregular form, especially towards the surface of the
fungus, where they compose a sort of pulp, varying in cohesion according
to the dry or moist condition of the atmosphere. All parts of these reddish
individuals seemed more or less infected with this disintegration, the
basidia divided by transverse diaphragms into several cylindrical or oblong
pieces, which finally become free. Transitional conditions were also
observed in mixed individuals. This sterile condition is called by Tulasne
“gemmiparous,” and he believes that it has ere now given origin to one or
more spurious species, and misled mycologists as to the real structure of
perfect and fruitful Dacrymyces.
Phalloidei.—In this order the hymenium is at first enclosed within a sort
of peridium or universal volva, maintaining a somewhat globose or egg-
shape. This envelope consists of an outer and inner coat of somewhat
similar texture, and an intermediate gelatinous layer, often of considerable
thickness. When a section is made of the fungus, whilst still enclosed in the
volva, the hymenium is found to present numerous cavities, in which
basidia are developed, each surmounted by spicules (four to six) bearing
oval or oblong spores.[Q] It is very difficult to observe the structure of the
hymenium in this order, on account of its deliquescent nature. As the
hymenium approaches maturity, the volva is ruptured, and the plant rapidly

enlarges. In Phallus, a long erect cellular stem bears the cap, over which the
hymenium is spread, and this expands enormously after escaping the
restraint of the volva. Soon after exposure, the hymenium deliquesces into a
dark mucilage, coloured by the minute spores, which drips from the pileus,
often diffusing a most loathsome odour for a considerable distance. In
Clathrus, the receptacle forms a kind of network. In Aseröe, the pileus is
beautifully stellate. In many the attractive forms would be considered
objects of beauty, were it not for their deliquescence, and often fœtid odour.
[R]

Podaxinei.—This is a small but very curious group of
fungi, in which the peridium resembles a volva, which is more
or less confluent with the surface of the pileus. They assume
hymenomycetal forms, some of them looking like Agarics,
Boleti, or species of Hydnum, with deformed gills, pores, or
spines; in Montagnites, in fact, the gill structure is very
distinct. The spores are borne in definite clusters on short Fig. 8.—Basidia
pedicels in such of the genera as have been examined.[S] and spores of
Phallus.
Hypogæi.—These are subterranean puff-balls, in which
sometimes a distinct peridium is present; but in most cases it consists
entirely of an external series of cells, continuous with the internal structure,
and cannot be correctly estimated as a peridium. The hymenium is sinuous
and convolute, bearing basidia with sterigmata and spores in the cavities.
Sometimes the cavities are traversed by threads, as in the Myxogastres. The
spores are in many instances beautifully echinulate, sometimes globose, at
others elongated, and produced in such numbers as to lead to the belief that
their development is successive on the spicules. When fully matured, the
peridia are filled with a dusty mass of spores, so that it is scarcely possible
in this condition to gain any notion of the structure. This is, indeed, the case
with nearly all Gasteromycetes. The hypogæous fungi are curiously
connected with Phalloidei by the genus Hysterangium.
Trichogastres.[T]—In their early stages the species contained in this
group are not gelatinous, as in the Myxogastres, but are rather fleshy and
firm. Very little has been added to our knowledge of structure in this group
since 1839 and 1842, when one of us wrote to the following effect:—If a
young plant of Lycoperdon cœlatum or L. gemmatum be cut through and
examined with a common pocket lens, it will be found to consist of a fleshy

mass, perforated in every direction with minute
elongated, reticulated, anastomosing,
labyrinthiform cavities. The resemblance of these
to the tubes of Boleti in an early stage of growth,
first led me to suspect that there must be some
very close connection between them. If a very
thin slice now be taken, while the mass is yet
firm, and before there is the slightest indication of
a change of colour, 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 young hymenium Fig. 9.—Basidia and spores of
Lycoperdon.
of an Agaric or Boletus. Occasionally one or two
filaments cross from one wall to another, and once I have seen these
anastomose. At a more advanced stage of growth, four little spicules are
developed at the tips of the sporophores, 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. It is clear that we have here a structure identical
with that of the true Hymenomycetes, a circumstance which accords well
with the fleshy habit and mode of growth. There is some difficulty in
ascertaining the exact structure of the species just noticed, as the fruit-
bearing cells, or sporophores, are very small, and when the spicules are
developed the substance becomes so flaccid that it is difficult to cut a
proper slice, even with the sharpest lancet. I have, however, satisfied myself
as to the true structure by repeated observations. But should any difficulty
arise in verifying it in the species in question, there will be none in doing so
in Lycoperdon giganteum. In this species the fructifying mass consists of
the same sinuous cavities, which are, however, smaller, so that the
substance is more compact, and I have not seen them traversed by any
filaments. In an early stage of growth, the surface of the hymenium, that is
of the walls of the cavities, consists of short threads composed of two or
three articulations, which are slightly constricted at the joints, from which,
especially from the last, spring short branchlets, often consisting of a single
cell. Sometimes two or more branchlets spring from the same point.
Occasionally the threads are constricted without any dissepiments, the
terminal articulations are obtuse, and soon swell very much, so as greatly to
exceed in diameter those on which they are seated. When arrived at their

full growth, they are somewhat obovate, and produce four spicules, which
at length are surmounted each with a globose spore. When the spores are
fully developed, the sporophores wither, and if a solution of iodine be
applied, which changes the spores to a rich brown, they will be seen still
adhering by their spicules to the faded sporophores. The spores soon
become free, but the spicule often still adheres to them; but they are not
attached to the intermingled filaments. In Bovista plumbea, the spores have
very long peduncles.[U] As in the Hymenomycetes, the prevailing type of
reproductive organs consisted of quaternary spores borne on spicules; so in
Gasteromycetes, the prevailing type, in so far as it is yet known, is very
similar, in some cases nearly identical, consisting of a definite number of
minute spores borne on spicules seated on basidia. In a very large number
of genera, the minute structure and development of the fructification
(beyond the mature spores) is almost unknown, but from analogy it may be
concluded that a method prevails in a large group like the Myxogastres
which does not differ in essential particulars from that which is known to
exist in other groups. The difficulties in the way of studying the
development of the spores in this are far greater than in the previous order.

Fig. 10.—a. Threads of Trichia. b. Portion further magnified, with spores. c.
Portion of spinulose thread.

Myxogastres.—At one time that celebrated mycologist, Professor De
Bary, seemed disposed to exclude this group from the vegetable kingdom
altogether, and relegate them to a companionship with amœboid forms. But
in more recent works he seems to have reconsidered, and almost, if not

entirely, abandoned, that disposition. These fungi, mostly minute, are
characterized in their early stages by their gelatinous nature. The substance
of which they are then composed bears considerable resemblance to
sarcode, and, did they never change from this, there might be some excuse
for doubting as to their vegetable nature; but as the species proceed towards
maturity they lose their mucilaginous texture, and become a mass of spores,
intermixed with threads, surrounded by a cellular peridium. Take, for
instance, the genus Trichia, and we have in the matured specimens a
somewhat globose peridium, not larger than a mustard seed, and sometimes
nearly of the same colour; this ultimately ruptures and exposes a mass of
minute yellow spherical spores, intermixed with threads of the same colour.
[V] These threads, when highly magnified, exhibit in themselves a spiral
arrangement, which has been the basis of some controversy, and in some
species these threads are externally spinulose. The chief controversy on
these threads has been whether the spiral markings are external or internal,
whether caused by twisting of the thread or by the presence of an external
or internal fibre. The spiral appearance has never been called in question,
only the structure from whence it arises, and this, like the striæ of diatoms,
is very much an open question. Mr. Currey held that the spiral appearance
may be accounted for by supposing the existence of an accurate elevation in
the wall of the cell, following a spiral direction from one end of the thread
to the other. This supposition would, he thinks, accord well with the optical
appearances, and it would account exactly for the undulations of outline to
which he alludes. He states that he had in his possession a thread of Trichia
chrysosperma, in which the spiral appearance was so manifestly caused by
an elevation of this nature, in which it is so clear that no internal spiral fibre
exists, that he did not think there could be a doubt in the mind of any person
carefully examining it with a power of 500 diameters that the cause of the
spiral appearance was not a spiral fibre. In Arcyria, threads of a different
kind are present; they mostly branch and anastomose, and are externally
furnished with prominent warts or spines, which Mr. Currey[W] holds are
also arranged in a spiral manner around the threads. In other Myxogastres,
threads are also present without any appreciable spiral markings or spines.
In the mature condition of these fungi, they so clearly resemble, and have
such close affinities with, the Trichogastres that one is led almost to doubt
whether it was not on hasty grounds, without due examination or

consideration, that proposals were made to remove them from the society of
their kindred.

Fig. 11.—Arcyria incarnata, with portion of threads and spore, magnified.

Very little is known of the development of the spores in
this group; in the early stages the whole substance is so
pulpy, and in the latter so dusty, whilst the transition from
one to the other is so rapid, that the relation between the
spores and threads, and their mode of attachment, has
never been definitely made out. It has been supposed that
the spinulose projections from the capillitium in some
species are the remains of pedicels from which, the spores
have fallen, but there is no evidence beyond this
supposition in its favour, whilst on the other hand, in
Stemonitis, for instance, there is a profuse interlacing
capillitium, and no spines have been detected. In order to
strengthen the supposition, spines should be more
commonly present. The threads, or capillitium, form a
beautiful reticulated network in Stemonitis, Cribraria,
Diachæa, Dictydium, &c. In Spumaria, Reticularia, Fig. 12.—Diachæa
Lycogala, &c., they are almost obsolete.[X] In no group is elegans.
the examination of the development of structure more
difficult, for the reasons already alleged, than in the Myxogastres.
Nidulariacei.—This small group departs in some important particulars
from the general type of structure present in the rest of the Gasteromycetes.

[Y] The plants here included may be described under three
parts, the mycelium, the peridium, and the sporangia. The
mycelium is often plentiful, stout, rigid, interlacing, and
coloured, running over the surface of the soil, or amongst
the vegetable débris on which the fungi establish
themselves. The peridia are seated upon this mycelium,
and in most instances are at length open above, taking the
form of cups, or beakers. These organs consist of three
Fig. 13.—Cyathus strata of tissue varying in structure, the external being
vernicosus.
fibrous, and sometimes hairy, the interior cellular and
delicate, the intermediate thick and at length tough, coriaceous, and
resistant. When first formed, the peridia are spherical, they then elongate
and expand, the mouth being for some time closed by a veil, or diaphragm,
which ultimately disappears. Within the cups lentil-shaped bodies are
attached to the base and sides by elastic cords. These are the sporangia.
Each of these has a complicated structure; externally there is a filamentous
tunic, composed of interlaced fibres, sometimes called the peridiole;
beneath this is the cortex, of compact homogenous structure, then follows a
cellular thicker stratum, bearing, towards the centre of the sporangia,
delicate branched threads, or sporophores, on which, at their extremities,
the ovate spores are generated, sometimes in pairs, but normally, it would
seem that they are quaternary on spicules, the threads being true basidia.
The whole structure is exceedingly interesting and peculiar, and may be
studied in detail in Tulasne’s memoir on this group.
Sphæronemei.—In this very large and, within
certain limits, variable order, there is but little of
interest as regards structure, which is not better
illustrated elsewhere; as, for instance, some sort
of perithecium is always present, but this can be
better studied in the Sphæriacei. The spores are
mostly very minute, borne on delicate
sporophores, which originate from the inner
surface of the perithecia, but the majority of so-
called species are undoubtedly conditions of
sphæriaceous fungi, either spermatogonia or
pycnidia, and are of much more interest when Fig. 14.—Cyathus. a.
studied in connection with the higher forms to Sporangium. b. Section. c.

which they belong.[Z] Probably the number of Sporophore. d. Spores.
complete and autonomous species are very few.
Melanconiei.—Here, again, are associated
together a great number of what formerly were
considered good species of fungi, but which are
now known to be but conditions of other forms.
One great point of distinction between these and
the preceding is the absence of any true
perithecium, the spores being produced in a kind
of spurious receptacle, or from a sort of stroma.
The spores are, as a rule, larger and much more
Fig. 15.—Asterosporium attractive than in Sphæronemei, and, in some
Hoffmanni. instances, are either very fine, or very curious.
Under this head we may mention the multiseptate spores of Coryneum; the
tri-radiate spores of Asterosporium; the curious crested spores of
Pestalozzia; the doubly crested spores of Dilophospora; and the scarcely
less singular gelatinous coated spores of Cheirospora. In all cases the
fructification is abundant, and the spores frequently ooze out in tendrils, or
form a black mass above the spurious receptacle from which they issue.[a]
Torulacei.—In this order there seems at first
to be a considerable resemblance to the Dematiei,
except that the threads are almost obsolete, and
the plant is reduced to chains of spores, without
trace of perithecium, investing cuticle, or definite
stroma. Sometimes the spores are simple, in other
Fig. 16.—Barren Cysts and
cases septate, and in Sporochisma are at first Pseudospores of Lecythea.
produced in an investing cell. In most cases
simple threads at length become septate, and are ultimately differentiated
into spores, which separate at the joints when fully mature.
Cæomacei.—Of far greater interest are the Coniomycetous parasites on
living plants. The present order includes those in which the spore[b] is
reduced to a single cell; and here we may observe that, although many of
them are now proved to be imperfect in themselves, and only forms or
conditions of other fungals, we shall write of them here without regard to
their duality. These originate, for the most part, within the tissues of living
plants, and are developed outwards in pustules, which burst through the

cuticle. The mycelium
penetrates the
intercellular passages,
and may sometimes be
found in parts of the Fig. 18.—Melampsora salicina.
plants where the fungus does not develop itself.
There is no proper excipulum or peridium, and
the spores spring direct from a more compacted
Fig. 17.—Coleosporium portion of the mycelium, or from a cushion-like
Tussilaginis, Lev. stroma of small cells. In Lecythea, the sub-
globose spores are at first generated at the tips of short pedicels, from which
they are ultimately separated; surrounding these spores arise a series of
barren cells, or cysts, which are considerably larger the true spores, and
colourless, while the spores are of some shade of yellow or orange.[c] In
Trichobasis, the spores are of a similar character, sub-globose, and at first
pedicellate; but there are no surrounding cysts, and the colour is more
usually brown, although sometimes yellow. In Uredo, the spores are at first
generated singly, within a mother cell; they are globose, and either yellow
or brown, without any pedicel. In Coleosporium, there are two kinds of
spores, those of a pulverulent nature, globose, which are sometimes
produced alone at the commencement of the season, and others which
originate as an elongated cell; this becomes septate, and ultimately
separates at the joints. During the greater part of the year, both kinds of
spores are to be found in the same pustule. In Melampsora, the winter
spores are elongated and wedge-shaped, compacted together closely, and
are only matured during winter on dead leaves; the summer spores are
pulverulent and globose, being, in fact, what were until recently regarded as
species of Lecythea. In Cystopus, the spores are sub-globose, or somewhat
angular, generated in a moniliform manner, and afterwards separating at the
joints. The upper spore is always the oldest, continuous production of
spores going on for some time at the base of the chain. Under favourable
conditions of moisture, each of these spores, or conidia, as De Bary terms
them, is capable of producing within itself a number of zoospores;[d] these
ultimately burst the vesicle, move about by the aid of vibratile cilia, and at
last settle down to germinate. Besides these, other reproductive bodies are
generated upon the mycelium, within the tissues of the plant, in the form of
globose oogonia, or resting spores, which, when mature, also enclose great

numbers of zoospores. Similar oogonia are produced amongst the
Mucedines in the genus Peronospora, to which De Bary considers Cystopus
to be closely allied. At all events, this is a peculiarity of structure and
development not as yet met with in any other of the Cæomacei. In
Uromyces is the nearest approach to the Pucciniæi; in fact, it is Puccinia
reduced to a single cell. The form of spore is usually more angular and
irregular than in Trichobasis, and the pedicel is permanent. It may be
remarked here, that of the foregoing genera, many of the species are not
autonomous that have hitherto been included amongst them. This is
especially true of Lecythea, Trichobasis, and, as it now appears, of
Uromyces.[e]

Pucciniæi.—This
group differs from the
foregoing chiefly in
having septate spores.
The pustules, or sori,
break through the
cuticle in a similar
manner, and here also
no true peridium is
present. In
Xenodochus, the
highest development of
Fig. 19.—Cystopus candidus.
joints is reached, each
Fig. 20.—Xenodochus
spore being composed of an indefinite number, carbonarius.
from ten to twenty cells. With it is associated an
unicellular yellow Uredine, of which it is a condition. Probably, in every
species of the Pucciniæi, it may hereafter be proved, as it is now suspected,
that an unicellular Uredine precedes or is associated with it, forming a
condition, or secondary form of fruit of that species. Many instances of that
kind have already been traced by De Bary,[f] Tulasne, and others, and some
have been a little too rashly surmised by their followers. In Phragmidium,
the pedicel is much more elongated than in Xenodochus, and the spore is
shorter, with fewer and a more definite number of cells for each species;
Mr. Currey is of opinion that each cell of the spore in Phragmidium has an
inner globose cell, which he caused to escape by rupture of the outer cell
wall as a sphæroid nucleus,[g] leading to the inference that each cell has its
own individual power of germination and reproduction. In Triphragmium,
there are three cells for each spore, two being placed side by side, and one
superimposed. In one species, however, Triphragmium deglubens (North
American), the cells are arranged as in Phragmidium, so that this represents
really a tricellular Phragmidium, linking the present with the latter genus. In
Puccinia the number of species is by far the most numerous; in this genus
the spores are uniseptate, and, as in all the Pucciniæi, the peduncles are
permanent. There is great variability in the compactness of the spores in the
sori, or pulvinules. In some species, the sori are so pulverulent that the
spores are as readily dispersed as in the Uredines, in others they are so
compact as to be separated from each other with great difficulty. As might

be anticipated, this has considerable effect on the contour of the spores,
which in pulverulent species are shorter, broader, and more ovate than in the
compact species. If a section of one of the more compact sori be made, it
will be seen that the majority of the spores are side by side, nearly at the
same level, their apices forming the external surface of the sori, but it will
not be unusual to observe smaller and younger spores pushing up from the
hymenial cells, between the peduncles of the elder spores, leading to the
inference that there is a succession of spores produced in the same
pulvinule. In Podisoma, a rather anomalous genus, the septate spores are
immersed in a gelatinous stratum, and some authors have imagined that
they have an affinity with the Tremellini, but this affinity is more apparent
than real. The phenomena of germination, and their relations to Rœstelia, if
substantiated, establish their claim to a position amongst the Pucciniæi.[h] It
seems to us that Gymnosporangium does not differ generically from
Podisoma. In a recently-characterized species, Podisoma Ellisii, the spores
are bi-triseptate. This is, moreover, peculiar from the great deficiency in the
gelatinous element. In another North American species, called
Gymnosporangium biseptatum, Ellis, which is distinctly gelatinous, there
are similar biseptate spores, but they are considerably broader and more
obtuse. In other described species they are uniseptate.
Ustilaginei.—These
fungi are now usually
treated as distinct from
the Cæomacei, to
which they are closely
related.[i] They are also
parasitic on growing
plants, but the spores Fig. 22.—Pseudospores of
are usually black or Puccinia.
sooty, and never yellow or orange; on an average
much smaller than in the Cæomacei. In Tilletia,
the spores are spherical and reticulated, mixed
with delicate threads, from whence they spring.
Fig. 21.—Phragmidium In the best known species, Tilletia caries, they
bulbosum. constitute the “bunt” of wheat. The peculiarities
of germination will be alluded to hereafter. In Ustilago, the minute sooty
spores are developed either on delicate threads or in compacted cells,

arising first from a sort of semi-gelatinous, grumous stroma. It is very
difficult to detect any threads associated with the spores. The species attack
the flowers and anthers of composite and polygonaceous plants, the leaves,
culms, and germen of grasses, &c., and are popularly known as “smuts.” In
Urocystis and Thecaphora, the spores are united together into sub-globose
bodies, forming a kind of compound spore. In some species of Urocystis,
the union which subsists between them is comparatively slight. In
Thecaphora, on the contrary, the complex spore, or agglomeration of
spores, is compact, being at first apparently enclosed in a delicate cyst. In
Tuburcinia, the minute cells are compacted into a hollow sphere, having
lacunæ communicating with the interior, and often exhibiting the remains of
a pedicel.
Æcidiacei.—
This group differs
from the
foregoing three
groups
prominently in the
Fig. 23.—Thecaphora hyalina.
presence of a Fig. 24.—Æcidium Berberidis.
cellular peridium, which encloses the spores; hence some mycologists have
not hesitated to propose their association with the Gasteromycetes, although
every other feature in their structure seems to indicate a close affinity with
the Cæomacei. The pretty cups in the genus Æcidium are sometimes
scattered and sometimes collected in clusters, either with spermogonia in
the centre or on the opposite surface. The cups are usually white, composed
of regularly arranged bordered cells at length bursting at the apex, with the
margins turned back and split into radiating teeth. The spores are commonly
of a bright orange or golden yellow, sometimes white or brownish, and are
produced in chains, or moniliform strings, slightly attached to each other,[j]
and breaking off at the summit at the same time that they continue to be
produced at the base, so that for some time there is a successive production
of spores. The spermogonia are not always readily detected, as they are
much smaller than the peridia, and sometimes precede them. The spermatia
are expelled from the lacerated and fringed apices, and are very minute and
colourless. In Rœstelia the peridia are large, growing in company, and
splitting longitudinally in many cases, or by a lacerated mouth. In most
instances, the spores are brownish, but in a splendid species from North

America (Rœstelia aurantiaca, Peck), recently characterized, they are of a
bright orange. If Œrsted is correct in his observations, which await
confirmation, these species are all related to species of Podisoma as a
secondary form of fruit.[k] In the Rœstelia of the pear-tree, as well as in that
of the mountain ash, the spermogonia will be found either in separate tufts
on discoloured spots, or associated with the Rœstelia, In Peridermium there
is very little structural difference from Rœstelia, and the species are all
found on coniferous trees. In Endophyllum, the peridia are immersed in the
succulent substance of the matrix; whilst in Graphiola, there is a tougher
and withal double peridium, the inner of which forms a tuft of erect threads
resembling a small brush.[l]
Hyphomycetes.—The predominant feature
in the structure of this order has already been
intimated to consist in the development of the
vegetative system under the form of simple or
branched threads, on which the fruit is
generated. The common name of mould is
applied to them perhaps more generally than
to other groups, although the term is too
vague, and has been too vaguely applied to be
of much service in giving an idea of the
characteristics of this order. Leaving the
smaller groups, and confining ourselves to the
Dematiei and the Mucedines, we shall obtain
some notion of the prevalent structure. In the
former the threads are more or less
carbonized, in the latter nearly colourless.
One of the largest genera in Dematiei is
Helminthosporium. It appears on decaying
herbaceous plants, and on old wood, forming
effused black velvety patches. The mycelium,
of coloured jointed threads, overlays and
penetrates the matrix; from this arise erect, Fig. 25.—Helminthosporium molle.
rigid, and usually jointed threads, of a dark brown, nearly black colour at
the base, but paler towards the apex. In most cases these threads have an
externally cortical layer, which imparts rigidity; usually from the apex, but
sometimes laterally, the spores are produced. Although sometimes

colourless, these are most commonly of some shade of brown, more or less
elongated, and divided transversely by few or many septa. In
Helminthosporium Smithii, the spores much exceed the dimensions of the
threads;[m] in other species they are smaller. In Dendryphium, the threads
and spores are very similar, except that the threads are branched at their
apex, and the spores are often produced one at the end of another in a short
chain.[n] In Septosporium again, the threads and spores are similar, but the
spores are pedicellate, and attached at or near the base; whilst in
Acrothecium, with similar threads and spores, the latter are clustered
together at the apex of the threads. In Triposporium, the threads are similar,
but the spores are tri-radiate; and in Helicoma, the spores are twisted
spirally. Thus, we might pass through all the genera to illustrate this chief
feature of coloured, septate, rather rigid, and mostly erect threads, bearing
at some point spores, which in most instances are elongated, coloured, and
septate.
Mucedines.—
Here, on the
other hand, the
threads, if
coloured at all,
are still delicate,
more flexuous,
with much
thinner walls,
and never
invested with an
Fig. 26.—Acrothecium simplex. external cortical
layer. One of the most important and highly Fig. 27.—Peronospora Arenariæ.
developed genera is Peronospora, the
members of which are parasitic upon and destructive of living vegetables. It
is to this genus that the mould of the too famous potato disease belongs.
Professor De Bary has done more than any other mycologist in the
investigation and elucidation of this genus; and his monograph is a
masterpiece in its way.[o] He was, however, preceded by Mr. Berkeley, and
more especially by Dr. Montagne, by many years in elucidation of the
structure of the flocci and conidia in a number of species.[p] In this genus,
there is a delicate mycelium, which penetrates the intercellular passages of

living plants, giving rise to erect branched threads, which bear at the tips of
their ultimate ramuli, sub-globose, ovate, or elliptic spores, or, as De Bary
terms them—conidia. Deeply seated on the mycelium, within the substance
of the foster plant, other reproductive bodies, called oogonia, originate.
These are spherical, more or less warted and brownish, the contents of
which become differentiated into vivacious zoospores, capable, when
expelled, of moving in water by the aid of vibratile cilia. A similar structure
has already been indicated in Cystopus, otherwise it is rare in fungi, if the
Saprolegniei be excluded. In Botrytis and in Polyactis, the flocci and spores
are similar, but the branches of the threads are shorter and more compact,
and the septa are more common and numerous; the oogonia also are absent.
De Bary has selected Polyactis cinerea, as it occurs on dead vine leaves, to
illustrate his views of the dualism which he believes himself to have
discovered in this species. “It spreads its mycelium in the tissue which is
becoming brown,” he writes, “and this shows at first essentially the same
construction and growth as that of the mycelium filaments of Aspergillus.”
On the mycelium soon appear, besides those which are spread over the
tissue of the leaves, strong, thick, mostly fasciculate branches, which stand
close to one another, breaking forth from the leaf and rising up
perpendicularly, the conidia-bearers. They grow about 1 mm. long, divide
themselves, by successively rising partitions, into some prominent
cylindrical linked cells, and then their growth is ended, and the upper cell
produces near its point three to six branches almost standing rectangularly.
Of these the under ones are the longest, and they again shoot forth from
under their ends one or more still shorter little branches. The nearer they are
to the top, the shorter are the branches, and less divided; the upper ones are
quite branchless, and their length scarcely exceeds the breadth of the
principal stem. Thus a system of branches appears, upon which, on a small
scale, a bunch of grapes is represented. All the twigs soon end their growth;
they all separate their inner space from the principal stem, by means of a
cross partition placed close to it. All the ends, and also that of the principal
stem, swell about the same time something like a bladder, and on the upper
free half of each swelling appear again, simultaneously, several fine
protuberances, close together, which quickly grow to little oval bladders
filled with protoplasm, and resting on their bearers with a sub-sessile,
pedicellate, narrow basis, and which at length separate themselves through
a partition as in Aspergillus. The detached cells are the conidia of our

fungus; only one is formed on each stalk. When the formation is completed
in the whole of the panicle, the little branches which compose it are
deprived of their protoplasm in favour of the conidia; it is the same with the
under end of the principal stem, the limits of which are marked by a cross
partition. The delicate wall of these parts shrinks up until it is
unrecognizable; all the conidia of the panicle approach one another to form
an irregular grape-like bunch, which rests loosely on the bearer, and from
which it easily falls away as dust. If they be brought into water they fall off
immediately; only the empty, shrivelled, delicate skins are to be found on
the branch which bore them, and the places on which they are fixed to the
principal stem clearly appear as round circumscribed hilums, generally
rather arched towards the exterior. The development of the main stem is not
ended here. It remains solid and filled with protoplasm as far as the portion
which forms the end through its conidia. Its end, which is to be found
among these pieces, becomes pointed after the ripening of the first panicle,
pushes the end of the shrivelled member on one side, and grows to the same
length as the height of one or two panicles, and then remains still, to form a
second panicle similar to the first. This is later equally perfoliated as the
first, then a third follows, and thus a large number of panicles are produced
after and over one another on the same stem. In perfect specimens, every
perfoliated panicle hangs loosely to its original place on the surface of the
stem, until by shaking or the access of water to it, it falls immediately into
the single conidia, or the remains of branches, and the already-mentioned
oval hilums are left behind. Naturally, the stem becomes longer by every
perfoliation; in luxuriant specimens the length can reach that of some lines.
Its partition is already, by the ripening of the first panicle from the
beginning of its foundation, strong and brown; it is only colourless at the
end which is extending, and in all new formations. During all these changes
the filament remains either unbranched, except as regards the transient
panicles, or it sends out here and there, at the perfoliated spots, especially
from the lower ones, one or two strong branches, standing opposite one
another and resembling the principal stem.
The mycelium, which grows so exuberantly in the leaf, often brings forth
many other productions, which are called sclerotia, and are, according to
their nature, a thick bulbous tissue of mycelium filaments. Their formation
begins with the profuse ramification of the mycelium threads in some place
or other; generally, but not always, in the veins of the leaf; the intertwining

twigs form an uninterrupted cavity, in which is
often enclosed the shrivelling tissue of the leaf.
The whole body swells to a greater thickness than
that of the leaf, and protrudes on the surface like
a thickened spot. Its form varies from circular to
fusiform; its size is also very unequal, ranging
between a few lines and about half a millimetre in
its largest diameter. At first it is colourless, but
afterwards its outer layers of cells become round,
of a brown or black colour, and it is surrounded
by a black rind, consisting of round cells, which
separate it from the neighbouring tissue. The
tissue within the rind remains colourless; it is an
entangled uninterrupted tissue of fungus
filaments, which gradually obtain very solid,
hard, cartilaginous coats. The sclerotium, which
ripens as the rind becomes black, loosens itself
easily from the place of its formation, and
Fig. 28.—Polyactis cinerea. a. remains preserved after the latter is decayed.
Apex of hypha.
The sclerotia are,
here as in many other fungi, biennial organs,
designed to begin a new vegetation after a state of
apparent quietude, and to send forth special fruit-
bearers. They may in this respect be compared to
the bulbs and perennial roots of under shrubs.
The usual time for the development of the
sclerotia is late in the autumn, after the fall of the
vine leaves. As long as the frost does not set in,
new ones continually spring up, and each one
attains to ripeness in a few days. If frost appears,
it can lie dry a whole year, without losing its
power of development. This latter commences
when the sclerotium is brought into contact with
Fig. 29.—Peziza Fuckeliana. a.
damp ground during the usual temperature of our Natural size. b. Section enlarged.
warmer seasons. If this occur soon, at the latest c. Ascus and sporidia.
some weeks after it is ripe, new vegetation grows
very quickly, generally after a few days; in several parts the colourless

filaments of the inner tissue begin to send out clusters of strong branches,
which, breaking through the black rind, stretch themselves up
perpendicularly towards the surface, separate from one another, and then
take all the characteristics of the conidia-bearers. Many such clusters can be
produced on one sclerotium, so that soon the greater part of the surface is
covered by filamentous conidia-bearers with their panicles. The colourless
tissue of the sclerotium disappears in the same degree as the conidia-bearers
grow, and at last the black rind remains behind empty and shrivelled. If we
bring, after many months, for the first time, the ripe sclerotium, in damp
ground, in summer or autumn, after it has ripened, the further development
takes place more slowly, and in an essentially different form. It is true that
from the inner tissue numerous filamentous branches shoot forth at the cost
of this growing fascicle, and break through the black rind, but its filaments
remain strongly bound, in an almost parallel situation, to a cylindrical cord,
which for a time lengthens itself and spreads out its free end to a flat plate-
like disc. This is always formed of strongly united threads, ramifications of
the cylindrical cord. On the free upper surface of the disc, the filaments
shoot forth innumerable branches, which, growing to the same height, thick
and parallel with one another, cover the before-named disc. Some remain
narrow and cylindrical, are very numerous, and produce fine hairs
(paraphyses); others, also very numerous, take the form of club-like
ampulla cells, and each one forms in its interior eight free swimming oval
spores. Those ampulla cells are sporidiiferous asci. After the spores have
become ripe, the free point of the utricle bursts, and the spores are scattered
to a great distance by a mechanism which we will not here further describe.
New ampullas push themselves between those which are ripening and
withering; a disc can, under favourable circumstances, always form new
asci for weeks at a time. The number of the already described utricle-
bearers is different, according to the size of the sclerotium; smaller
specimens usually produce only one, larger two to four. The size is
regulated by that of the sclerotia, and ranges, in full-grown specimens,
between one and more millimetres for the length of the stalk, and a half to
three (seldom more) millimetres for the breadth of the disc.[q] For some
time the conidia form, belonging to the Mucedines, has been known as
Botrytis cinerea (or Polyactis cinerea). The compact mycelium, or
sclerotium, as an imperfect fungus, bore the name of Sclerotium echinatum,
whilst to the perfect and cup-like form has been given the name of Peziza

Fuckeliana. We have reproduced De Bary’s life-history of this mould here,
as an illustration of structure in the Mucedines, but hereafter we shall have
to write of similar transformations when treating of polymorphism.
The form of the threads, and the form and
disposition of the spores, vary according to the
genera of which this order is composed. In
Oidium the mostly simple threads break up into
joints. Many of the former species are now
recognized as conditions of Erysiphe. In
Aspergillus, the threads are simple and erect, with
a globose head, around which are clustered
chains of simple spores. In Penicillium, the lower
portion of the threads is simple, but they are
shortly branched at the apex, the branches being
terminated by necklaces of minute spores. In
Fig. 30.—Penicillium
Dactylium, the threads are branched, but the chartarum, Cooke.
spores are collected in clusters usually, and are
moreover septate. In other genera similar distinctions prevail. These two
groups of black moulds and white moulds are the noblest, and contain the
largest number of genera and species amongst the Hyphomycetes. There is,
however, the small group of Isariacei, in which the threads are compacted,
and a semblance of such hymenomycetal forms as Clavaria and Pterula is
the result, but it is doubtful if this group contains many autonomous
species. In another small group, the Stilbacei, there is a composite character
in the head, or receptacle,[r] and in the stem when the latter is present. Many
of these, again, as Tubercularia, Volutella, Fusarium, &c., contain doubtful
species. In Sepedoniei and Trichodermacei, the threads are reduced to a
minimum, and the spores are such a distinctive element that through these
groups the Hyphomycetes are linked with the Coniomycetes. These groups,
however, are not of sufficient size or importance to demand from us, in a
work of this character, anything more than the passing allusion which we
have given to them.
We come now to consider the structure in the Sporidiifera, in which the
fructifying corpuscles or germs, whether called spores or sporidia, are
generated within certain privileged cysts, usually in definite numbers. In
systematic works, these are included under two orders, the Physomycetes

and the Ascomycetes. The former of these consists of cyst-bearing moulds,
and from their nearest affinity to the foregoing will occupy the first place.
Physomycetes include, especially
amongst the Mucorini, many most
interesting and instructive species
for study, which even very lately
have occupied the attention of
continental mycologists. Most of
these phenomena are associated
more or less with reproduction, and
as such will have to be adverted to
again, but there are points in the
structure which can best be alluded
to here. Again taking Professor de
Bary’s researches as our guide,[s]
we will illustrate this by the
common Mucor mucedo: If we
Fig. 31.—Mucor mucedo, with three sporangia. a.
bring quite fresh horse-dung into a Portion of frill with sporangiola.
damp confined atmosphere, for
example, under a bell-glass, there appears on its surface, after a few days,
an immense white mildew. Upright strong filaments of the breadth of a hair
raise themselves over the surface, each of them soon shows at its point a
round little head, which gradually becomes black, and a closer examination
shows us that in all principal points it perfectly agrees with the sporangia of
other species. Each of these white filaments is a sporangia-bearer. They
spring from a mycelium which is spread in the dung, and appear singly
upon it. Certain peculiarities in the form of the sporangium, and the little
long cylindrical spores, which, when examined separately, are quite flat and
colourless, are characteristic of the species. If the latter be sown in a
suitable medium, for example, in a solution of sugar, they swell, and shoot
forth germinating utricles, which quickly grow to mycelia, which bear
sporangia. This is easily produced on the most various organic bodies, and
Mucor mucedo is therefore found spontaneously on every substratum which
is capable of nourishing mildew, but on the above-named the most perfect
and exuberant specimens are generally to be found. The sporangia-bearers
are at first always branchless and without partitions. After the sporangium is
ripe, cross partitions in irregular order and number often appear in the inner

space, and on the upper surface branches of different number and size, each
of which forms a sporangium at its point. The sporangia which are formed
later are often very similar, but sometimes very different, to those which
first appeared, because their partition is very thick and does not fall to
pieces when it is ripe, but irregularly breaks off, or remains entire,
enclosing the spores, and at last falls to the ground, when the fungus
withers. The cross partition which separates the sporangia from its bearers
is in those which are first formed (which are always relatively thicker
sporangia) very strongly convex, while those which follow later are often
smaller, and in little weak specimens much less arched, and sometimes
quite straight. After a few days, similar filaments generally show
themselves on the dung between the sporangia-bearers, which appear to the
naked eye to be provided with delicate white frills. Where such an one is to
be found, two to four rectangular expanding little branches spring up to the
same height round the filament. Each of these, after a short and simple
process, branch out into a furcated form; the furcations being made in such
a manner that the ends of the branch at last so stand together that their
surface forms a ball. Finally, each of the ends of a branch swells to a little
round sporangium, which is limited by a partition (called sporangiolum, to
distinguish it from the larger ones), in which some, generally four, spores
are formed in the manner already known. When the sporangiola are alone,
they have such a peculiar appearance, with their richly-branched bearers,
that they can be taken for something quite different to the organs of the
Mucor mucedo, and were formerly not considered to belong to it. That they
really belong to the Mucor is shown by the principal filament which it
bears, not always, but very often, ending with a large sporangium, which is
characteristic of the Mucor mucedo; it is still more evident if we sow the
spores of the sporangiolum, for, as it germinates, a mycelium is developed,
which, near a simple bearer, can form large sporangia, and those form
sporangiola, the first always considerably preponderating in number, and
very often exclusively. If we examine a large number of specimens, we find
every possible middle form between the simple or less branched sporangia-
bearers and the typical sporangiola frills; and we arrive at last at the
conclusion simply to place the latter among the varieties of form which the
sporangia-bearer of the Mucor mucedo shows, like every other typical
organic form within certain limits. On the other hand, propagation organs,
differing from those of the sporangia and their products, belong to Mucor

mucedo, which may be termed conidia. On the dung (they are rare on any
other substance) these appear at the same time, or generally somewhat later,
than the sporangia-bearers, and are not unlike those to the naked eye. In a
more accurate examination, they appear different; a thicker, partition-less
filament rises up and divides itself, generally three-forked, at the length of
one millimetre, into several series of branchlets. The forked branches of the
last series bear under their points, which are mostly capillary, short erect
little ramuli, and these, with which the ends of the principal branches
articulate on their somewhat broad tops, several spores and conidia, near
one another; about fifteen to twenty are formed at the end of each little
ramulus. The peculiarities and variations which so often appear in the
ramification need not be discussed here. After the articulation of the
conidia, their bearers sink together by degrees, and are quite destroyed. The
ripe conidia are round like a ball, their surface is scarcely coloured, and
almost wholly smooth. These conidioid forms were at first described as a
separate species under the name of Botrytis Jonesii. How, then, do they
belong to the Mucor?[t] That they appear gregariously is as little proof of an
original relation to one another, here as elsewhere. Attempts to prove that
the conidia and sporangia-bearers originate on one and the same mycelium
filament may possibly hereafter succeed. Till now this has not been the
case, and he who has ever tried to disentangle the mass of filaments which
exuberantly covers the substratum of a Mucor vegetation, which has
reached so far as to form conidia, will not be surprised that all attempts
have hitherto proved abortive. The suspicion of the connection founded on
the gregariously springing up, and external resemblance, is fully justified, if
we sow the conidia in a suitable medium, for example, in a solution of
sugar. They here germinate and produce a mycelium which exactly
resembles that of the Mucor mucedo, and, above all, they produce in
profusion the typical sporangia of the same on its bearers. The latter are till
now alone reproductions of conidia-bearers, and have never been observed
on mycelia which have grown out of conidia.
These phenomena of development appear in the Mucor when it dwells on
a damp substance, which must naturally contain the necessary nourishment
for it, and is exposed to the atmospheric air. Its mycelium represents at first
strong branched utricles without partitions; the branches are of the higher
order, mostly divided into rich and very fine-pointed ramuli. In old
mycelium, and also in the sporangia-bearers, the contents of which are

mostly used for the formation of spores, and
the substratum of which is exhausted for our
fungus, short stationary pieces, filled with
protoplasm, are very often formed into cells
through partitions in order to produce spores,
that is, grow to a new fruitful mycelium. These
cells are called gemmules, brooding cells, and
resemble such vegetable buds and sprouts of
Fig. 32.—Small portion of Botrytis
foliaceous plants which remain capable of Jonesii.
development after the organs of vegetation are
dead, in order to grow, under suitable circumstances, to new vegetating
plants, as, for example, the bulbs of onions, &c.
If we bring a vegetating mycelium of Mucor mucedo into a medium
which contains the necessary nourishment for it, but excluded from the free
air, the formation of sporangia takes place very sparingly or not at all, but
that of gemmules is very abundant. Single interstitial pieces of the ramuli,
or even whole systems of branches, are quite filled with a rich greasy
protoplasm; the short pieces and ends are bound by partitions which form
particular, often tun-like or globular cells; the longer ones are changed,
through the formation of cross partitions, into chains of similar cells; the
latter often attain by degrees strong, thick walls, and their greasy contents
often pass into innumerable drops of a very regular globular form and of
equal size. Similar appearances show themselves after the sowing of spores,
which are capable of germinating in the medium already described, from
which the air is excluded. Either short germinating utricles shoot forth,
which soon form themselves into rows of gemmules, or the spores swell to
large round bladders filled with protoplasm, and shoot forth on various parts
of their surface innumerable protuberances, which, fixing themselves with a
narrow basis, soon become round vesiculate cells, and on which the same
sprouts which caused their production are repeated, formations which
remind us of the fungus of fermentation called globular yeast. Among all
the known forms of gemmules we find a variety which are intermediate, all
of which show, when brought into a normal condition of development, the
same proportion, and the same germination, as those we first described.
We have detailed rather at length the structure and development of one of
the most common of the Mucors, which will serve as an illustration of the

order. Other distinctions there may be which are of more interest as defining
the limits of genera, except such as may be noticed when we come to write
more specially of reproduction.
Ascomycetes.—Passing now to the Ascomycetes, which are especially
rich in genera and species, we must first, and but superficially, allude to
Tuberacei, an order of sporidiiferous fungi of subterranean habit, and rather
peculiar structure.[u] In this order an external stratum of cells forms a kind
of perithecium, which is more or less developed in different genera. This
encloses the hymenium, which is sinuous, contorted, and twisted, often
forming lacunæ. The hymenium in some genera consists of elongated,
nearly cylindrical asci, enclosing a definite number of sporidia; in the true
truffles and their immediate allies, the asci are broad sacs, containing very
large and beautiful, often coloured, sporidia. These latter have either a
smooth, warted, spinulose, or lacunose epispore, and, as will be seen from
the figures in Tulasne’s Monograph,[v] or those in the last volume of
Corda’s great work,[w] are attractive microscopical objects. In some cases, it
is not difficult to detect paraphyses, but in others they would seem to be
entirely absent. A comparatively large number have been discovered and
recorded in Great Britain,[x] but of those none are more suitable for study of
general structure than the ordinary truffle of the markets.
The structure of the remaining Ascomycetes can be studied under two
groups, i.e., the fleshy Ascomycetes, or, as they have been termed, the
Discomycetes, and the hard, or carbonaceous Ascomycetes, sometimes
called the Pyrenomycetes. Neither of these names gives an accurate idea of
the distinctions between the two groups, in the former of which the discoid
form is not universal, and the latter contains somewhat fleshy forms. But in
the Discomycetes the hymenium soon becomes more or less exposed, and
in the latter it is enclosed in a perithecium. The Discomycetes are of two
kinds, the pileate and the cup-shaped. Of the pileate such a genus as
Gyromitra or Helvella is, in a certain sense, analogous to the Agarics
amongst Hymenomycetes, with a superior instead of an inferior hymenium,
and enclosed, not naked, spores. Again, Geoglossum is somewhat analogous
to Clavaria. Amongst the cup-shaped, Peziza is an Ascomycetous Cyphella.
But these are perhaps more fanciful than real analogies.
Recently Boudier has examined one group of the cup-shaped
Discomycetes, the Ascobolei, and, by making a somewhat free use of his

Memoir,[y] we may arrive at a general idea of the structure in the cupulate
Discomycetes. They present themselves at first under the form of a small
rounded globule, and almost entirely cellular. This small globule, the
commencement of the receptacle, is not long in increasing, preserving its
rounded form up to the development of the asci. At this period, under the
influence of the rapid growth of these organs, it soon produces at its summit
a fissure of the external membrane, which becomes a more marked
depression in the marginate species. The receptacle thus formed increases
rapidly, becomes plane, more convex, or more or less undulated at the
margin, if at all of large size. Fixed to the place where it is generated by
some more or less abundant mycelioid filaments, the receptacle becomes
somewhat cup-shaped and either stipitate or sessile, composed of the
receptacle proper and the hymenium.

Fig. 33.—Section of cup of Ascobolus. a. External cells. b.
Secondary layer. c. Subhymenial tissue (Janczenski).

The receptacle proper comprehends the subhymenial tissue, the
parenchyma, and the external membrane. The subhymenial tissue is
composed of small compact cells, forming generally a more coloured and
dense stratum, the superior cells of which give rise to the asci and
paraphyses. The parenchyma is seated beneath this, and is generally of
interlaced filaments, of a looser consistency than the preceding, united by
intermediate cellules. The external membrane, which envelopes the
parenchyma, and limits the hymenium, differs from the preceding by the
cells often being polyhedric, sometimes transverse, and united together, and
sometimes separable. Externally it is sometimes smooth, and sometimes
granular or hairy.

The hymenium is, however, the most, important part, consisting of (1)
the paraphyses, (2) the asci, and sometimes (3) an investing mucilage. The
asci are always present, the paraphyses are sometimes rare, and the
mucilage in many cases seems to be entirely wanting.
The paraphyses, which are formed at the first commencement of the
receptacle, are at first very short, but soon elongate, and become wholly
developed before the appearance of the asci. They are linear, sometimes
branched and sometimes simple, often more or less thickened at their tips;
almost always they contain within them some oleaginous granules, either
coloured or colourless. Their special function seems still somewhat obscure,
and Boudier suggests that they may be excitatory organs for the dehiscence
of the asci. However this may be, some mycologists are of opinion that, at
least in some of the Ascomycetes, the paraphyses are abortive asci, or, at
any rate, that abortive asci mixed with the paraphyses cannot be
distinguished from them.
The mucilage forms itself almost at the same time as the paraphyses, and
previous to the formation of the asci. This substance appears as a colourless
or yellowish mucilage, which envelopes the paraphyses and asci, and so
covers the hymenium with a shining coat.
The asci appear first at the base of the paraphyses, under the form of
oblong cells, filled with colourless protoplasm. By rapid growth, they soon
attain a considerable size and fulness, the protoplasm being gradually
absorbed by the sporidia, the first indication of which is always the central
nucleus. The mucilage also partly disappears, and the asci, attaining their
maturity, become quite distinct, each enclosing its sporidia. But before they
take their complete growth they detach themselves from the subhymenial
tissue, and being attenuated towards their base, are forced upwards by
pressure of the younger asci, to, and in some instances beyond, the upper
surface of the disc. This phenomenon commences during the night, and
continues during the night and all the morning. It attains its height at mid-
day, and it is then that the slightest breath of air, the slightest movement,
suffices to cause dehiscence, which is generally followed by a scarcely
perceptible contractile motion of the receptacle.
There is manifestly a succession in formation and maturity of the asci in
a receptacle. In the true Ascobolei, in which the sporidia are coloured, this

may be more distinctly seen. At first some thin
projecting points appear upon the disc, the next
day they are more numerous, and become more
and more so on following days, so as to render
the disc almost covered with raised black or
crystalline points;[z] these afterwards diminish
day by day, until they ultimately cease. The asci,
after separation from the subhymenial tissue,
continue to lengthen, or it may be that their
elasticity permits of extension, during expulsion.
Boudier considers that an amount of elasticity is
certain, because he has seen an ascus arrive at
maturity, eject its spores, and then make a sharp
and considerable movement of retraction, then
the ascus returned again, immediately towards its
previous limits, always with a reduction in the
number of its contained sporidia.
The dehiscence of the asci takes place in the
Ascobolei, in some species of Peziza, Morchella,
Helvella, and Verpa, by means of an apical
operculum, and in other Pezizæ, Helotium,
Geoglossum, Leotia, Mitrula, &c., by a fissure of Fig. 34.—Asci, sporidia, and
the ascus. This operculum may be the more paraphyses of Ascobolus
readily seen when the ascus is coloured by a drop (Boudier).
of tincture of iodine.
The sporidia are usually four or eight, or some multiple of that number,
in each ascus, rarely four, most commonly eight. At a fixed time the
protoplasm, which at first filled the asci, disappears or is absorbed in a
mucilaginous matter, which occupies its place, in the midst of which is a
small nucleus, which is the rudiment of the first spore; other spores are
formed consecutively, and then the substance separates into as many
sections as there are sporidia. From this period each sporidium seems to
have a separate existence. All have a nucleus, which is scarcely visible,
often slightly granular, but which is quite distinct from the oleaginous
sporidioles so frequent amongst the Discomycetes, and which are
sometimes called by the same name. The sporidia are at first a little smaller

than when mature, and are surrounded by mucilage. After this period the
sporidia lose their nebulous granulations, whilst still preserving their
nucleus; their outlines are distinct, and, amongst the true Ascobolei,
commence acquiring a rosy colour, the first intimation of maturity. This
colour manifests itself rapidly, accumulating exclusively upon the epispore,
which becomes of a deep rose, then violet, and finally violet blue, so deep
as sometimes to appear quite black. There are some modifications in this
coloration, since, in some species, it passes from a vinous red to grey, then
to black, or from rose-violet to brown.
The epispore acquires a waxy consistence by this pigmentation, so that it
may be detached in granules. It is to this particular consistency of the
epispore that the cracks so frequent in the coloured sporidia of Ascobolus
are due, through contraction of the epispore. As they approach maturity, the
sporidia accumulate towards the apex of the asci, and finally escape in the
manner already indicated.
In all essential particulars there is a great similarity in the structure of the
other Discomycetes, especially in their reproductive system. In most of
them coloured sporidia are rare. In some the receptacle is pileate, clavate, or
inflated, whilst in Stictis it is very much reduced, and in the lowest form of
all, Ascomyces, it is entirely absent. In the Phacidiacei, the structure is very
similar to that of the Elvellacei, whilst the Hysteriacei, with greater
affinities with the latter, still tend towards the Pyrenomycetes by the more
horny nature of the receptacle, and the greater tendency of the hymenium to
remain closed, at least when dry. In some species of Hysterium, the sporidia
are remarkably fine. M. Duby[AA] has subjected this group to examination,
and M. Tulasne partly so.[AB]
Sphæriacei.—In this group there is considerable variation, within certain
limits. It contains an immense number of species, and these are daily being
augmented. The general feature in all is the presence of a perithecium,
which contains and encloses the hymenium, and at length opening by a pore
or ostiolum at the apex. In some the perithecia are simple, in others
compound; in some immersed in a stroma, in others free; in some fleshy or
waxy, in others carbonaceous, and in others membranaceous. But in all
there is this important difference from the Ascomycetes we have already
had under consideration, that the hymenium is never exposed. The
perithecium consists usually of an external layer of cellular structure, which

is either smooth or hairy, usually blackish, and an internal stratum of less
compact cells, which give rise to the hymenium.
As in the Discomycetes, the hymenium
consists of asci, paraphyses, and mucilage, but
the whole forms a less compact and more
gelatinous mass within the perithecium. The
formation and growth of the asci and sporidia
differ little from what we have described, and Fig. 35.—Perithecium of Sphæria
when mature the asci dehisce, and the sporidia and Section.
alone are ejected from the ostiolum. We are not aware that operculate asci
have yet been detected. It has been shown in some instances, and suspected
in others, that certain moulds, formerly classed with Mucedines and
Dematiei, especially in the genus Helminthosporium, bear the conidia of
species of Sphæria, so that this may be regarded as one form of fruit.
Perithecia, very similar externally to those of Sphæria, but containing
spores borne on slender pedicels and not enclosed in asci, have had their
relations to certain species of Sphæria indicated, and these are no longer
regarded so much as species of Hendersonia or Diplodia as the pycnidia of
Sphæria. Other and more minute perithecia, containing minute, slender
stylospores in great numbers, formerly classed with Aposphæria, Phoma,
&c., but are now recognized as spermogonia containing the spermatia of
Sphæriæ. How these influence each other, when and under what
circumstances the spermatia are instrumental in impregnation of the
sporidia, is still matter of mystery. It is clear, however, that in all these
conidia, macrospores, microspores, and some spermatia, or by whatever
names they may be called, there exists a power of germination. Tulasne has
indicated in some instances five or six forms of fruit as belonging to one
fungus, of which the highest and most perfect condition is a species of
Sphæria.
Perisporiacei.—Except in the perithecia rupturing irregularly, and not
dehiscing by a pore, some of the genera in this group differ little in structure
from the Sphæriacei. On the other hand, the Erysiphei present important
and very interesting features. They occur chiefly on the green parts of
growing plants. At first there is a more or less profuse white mycelium.[AC]
This gives rise to chains of conidia (Oidium), and afterwards small
sphæroid projections appear at certain points on the mycelium. These

enlarge, take an orange colour, ultimately passing
into brown, and then nearly black. Externally
these perithecia are usually furnished with long,
spreading, intertwined, or branching appendages,
sometimes beautifully branched or hooked at
their tips. In the interior of the receptacles, pear-
shaped or ovate asci are formed in clusters,
attached together at the base, and containing two
Fig. 36.—Uncinula adunca.
or more hyaline sporidia. Other forms of fruit
have also been observed on the same mycelium. In an exotic genus,
Meliola, the fulcra, or appendages, as well as the mycelium, are black,
otherwise it is very analogous to such a genus of Erysiphei as
Microsphæria. In Chætomium, the perithecia bristle with rigid, dark-
coloured hairs, and the sporidia are coloured. Our limits, however, will not
permit of further elucidation of the complex and varied structure to be
found amongst fungi.[AD]
[A]
A curious case occurred some years since at Bury St. Edmunds, which may be
mentioned here in connection with the development of these nodules. Two children
had died under suspicious circumstances, and an examination of the body of the latter
after exhumation was made, a report having arisen that the child died after eating
mushrooms. As certain white nodules appeared on the inner surface of the intestines,
it was at once hastily concluded that the spores of the mushroom had germinated, and
that the nodules were infant mushrooms. This appeared to one of us so strange, that
application was made for specimens, which were kindly forwarded, and a cursory
glance was enough to convince us that they were not fungoid. An examination under
the microscope further confirmed the diagnosis, and the application of nitric acid
showed that the nodules were merely due to chalk mixture, which had been given to
the child for the diarrhetic symptoms under which he succumbed.
[B]
Ehrenberg compared the whole structure of an Agaric with that of a mould, the
mycelium corresponding with the hyphasma, the stem and pileus with the flocci, and
the hymenium with the fructifying branchlets. The comparison is no less ingenious
than true, and gives a lively idea of the connection of the more noble with the more
humble fungi.—Ehrb. de Mycetogenesi.
[C]
In Paxillus involutus the hymenium may be readily torn off and unfolded.
[D]
This was well delineated in “Flora Danica,” plate 834, as observed in Coprinus
comatus as long ago as 1780.

[E]
A. de Bary, “Morphologie und Physiologie der Pilze,” in “Hofmeister’s Handbuch,”
vol. ii. cap. 5, 1866, translated in “Grevillea,” vol. i. p. 181.
[F]
“Die Pollinarien und Spermatien von Agaricus,” in “Botanische Zeitung,” Feb. 29
and March 7, 1856.
[G]
“Essai d’une Flore mycologique de la Région de Montpellier.” Paris, 1863.
[H]
Hoffmann, “Botanische Zeitung,” 1856, p. 139.
[I]
Corda, “Icones Fungorum hucusque cognitorum,” iii. p. 41. Prague, 1839.
[J]
Cooke, M. C., “Anatomy of a Mushroom,” in “Popular Science Review,” vol. viii. p.
380.
[K]
An attempt was made to show that, in Agaricus melleus, distinct asci were found, in a
certain stage, on the gills or lamellæ. We have in vain examined the gills in various
conditions, and could never detect anything of the kind. It is probable that the asci
belonged to some species of Hypomyces, a genus of parasitic Sphæriaceous fungi.
[L]
It is not intended that the spores are always quaternate in Agaricini, though that
number is constant in the more typical species. They sometimes exceed four, and are
sometimes reduced to one.
[M]
The species long known as Hydnum gelatinosum was examined by Mr. F. Currey in
1860 (Journ. Linn. Soc.), and he came to the conclusion that it was not a good
Hydnum. Since then it has been made the type of a new genus (Hydnoglœa B. and Br.
or, as called by Fries, in the new edition of “Epicrisis,” Tremellodon, Pers. Myc. Eur.),
and transferred to the Tremellini. Currey says, upon examining the fructification, he
was surprised to find that, although in its external characters it was a perfect Hydnum,
it bore the fruit of a Tremella. If one of the teeth be examined with the microscope, it
will be seen to consist of threads bearing four-lobed sporophores, and spores exactly
similar to Tremella. It will thus be seen, he adds, that the plant is exactly intermediate
between Hydnei and Tremellini, forming, as it were, a stepping-stone from one to the
other.
[N]

Tulasne, L. R. and C., “Observations on the Organization of the Tremellini,” in “Ann.
des Sci. Nat.” 3me sér. xix. (1853), pp. 193, &c.
[O]

M. Léveillé, in “Ann. des Sci. Nat.” 2me sér. viii. p. 328; 3me sér. ix. p. 127; also
Bonorden, “Handbuch der Mycologie,” p. 151.
[P]
Tulasne, in “Ann. des Sci. Nat.” (loc. cit.) xix. pl. x. fig. 29. Tulasne, “New Notes
upon Tremellinous Fungi,” in “Journ. Linn. Soc.” vol. xiii. (1871), p. 31.
[Q]
Berkeley, M. J., “On the Fructification of Lycoperdon, Phallus, &c.,” in “Ann. Nat.
Hist.” 1840, vol. iv. p. 158, pl. 5. Berkeley, M. J., “Introduction Crypt. Bot.” p. 346.
[R]
Tulasne, L. R. and C., “Fungi Hypogæi.” Paris. Berkeley and Broome, “British
Hypogæous Fungi,” in “Ann. Nat. Hist.” 1846, xviii. p. 74. Corda, “Icones
Fungorum,” vol. vi. pl. vii. viii.
[S]

Tulasne, “Sur le Genre Secotium,” in “Ann. des Sci. Nat.” (1845), 3me sér. vol. iv. p.
169, plate 9.
[T]
Tulasne, L. R. and C., “De la Fructification des Scleroderma comparée a celle des
Lycoperdon et des Borista,” in “Ann. des Sci. Nat.” 1842, xvii. p. 5. Tulasne, L. R.
and C., “Sur les Genres Polysaccum et Geaster,” in “Ann. des Sci. Nat.” 1842, xviii.
p. 129, pl. 5 and 6.
[U]
Berkeley, “On the Fructification of Lycoperdon, &c.,” in “Annals of Natural History”
(1840), iv. p. 155.
[V]

Wigand, “Morphologie des Genres Trichia et Arcyria,” in “Ann. des Sci. Nat.” 4me
sér. xvi. p. 223.
[W]
Currey, “On Spiral Threads of Trichia,” in “Quart. Journ. Micr. Science” (1855), iii. p.
17.
[X]
In some of the genera, as, for instance, in Badhamia, Enerthenema, and Reticularia,
the spores are produced within delicate cells or cysts, which are afterwards absorbed.
[Y]

Tulasne, “Essai d’une Monographie des Nidulariées,” in “Ann. des Sci. Nat.” (1844),
i. 41 and 64.
[Z]
Berkeley, M. J., “Introduction, Crypt. Bot.” p. 330.
[a]
Berkeley, M. J., “Introduction, Crypt. Bot.” p. 329.
[b]
In the Cæomacei and Pucciniæi the term “pseudospore” would be much more
accurate.
[c]
Léveillé, “Sur la Disposition Méthodique des Urédinées,” in “Ann. des Sci. Nat.”
(1847), vol. viii. p. 369.
[d]

De Bary, “Champignons Parasites,” in “Ann. des Sci. Nat.” 4me sér. vol. xx.
[e]
Tulasne, “Mémoire sur les Urédinées, &c.,” in “Ann. des Sci. Nat.” (1854), vol. ii. p.
78.
[f]
De Bary, “Ueber die Brandpilze,” Berlin, 1853.
[g]
Currey, in “Quart. Journ. Micr. Sci.” (1857), vol. v. p. 119, pl. 8, fig 13.
[h]
Cooke, “On Podisoma,” in “Journal of Quekett Microscopical Club,” vol. ii. p. 255.
[i]
Tulasne, “Mémoire sur les Ustilaginées,” in “Ann. des Sci. Nat.” (1847), vii. pp. 12
and 73.
[j]
Corda, “Icones Fungorum,” vol. iii. fig. 45.
[k]
Cooke, “On Podisoma,” in “Quekett Journal,” vol. ii. p. 255.
[l]
It may be a question whether Graphiola is not more nearly allied to Trichocoma
(Jungh Fl. Crypt. Javæ, p. 10, f. 7) than to the genera with which it is usually
associated.—M. J. B.

[m]
Cooke, “On Microscopic Moulds,” in “Quekett Journal,” vol. ii. plate 7.
[n]
See “Dendryphium Fumosum,” in “Quekett Journal,” vol. ii. plate 8; or, “Corda
Prachtflora,” plate 22.
[o]

De Bary, “Champignons Parasites,” in “Ann. des Sci. Nat.” 4me sér. vol. xx.
[p]
Berkeley, “On the Potato Murrain,” in “Journ. of Hort. Soc. of London,” vol. i.
(1846), p. 9.
[q]
De Bary, “On Mildew and Fermentation,” p. 25, reprinted from “German Quarterly
Magazine,” 1872; De Bary, “Morphologie und Physiologie der Pilze,” (1866), 201.
[r]
Cooke, “Handbook of British Fungi,” vol. ii. p. 552.
[s]
De Bary, “On Mildew and Fermentation,” in “Quarterly German Magazine,” for
1872.
[t]
We are quite aware that Von Tieghem and Le Monnier, in “Ann. des Sci. Nat.” 1873,
p. 335, dispute that this belongs to Mucor mucedo, and assert that Chætocladium
Jonesii is itself a true Mucor, with monosporous sporangia.
[u]
Vittadini, “Monographia Tuberacearum,” 1831.
[v]
Tulasne, “Fungi Hypogæi,” 1851.
[w]
Corda, “Icones Fungorum,” vol. vi.
[x]
Berkeley and Broome, in “Ann. of Nat. Hist.” 1st ser. vol. xviii. (1846), p. 73; Cooke,
in “Seem. Journ. Bot.”
[y]

Boudier (E.), “Mémoire sur les Ascobolés,” in “Ann. des Sci. Nat.” 5me sér. vol. x.
(1869).

[z]
Only in some of the Discomycetes are the asci exserted.
[AA]
Duby, “Mémoire sur la Tribu des Hysterinées,” 1861.
[AB]
Tulasne, “Selecta Fungorum Carpologia,” vol. iii.
[AC]
Tulasne, “Selecta Fungorum Carpologia,” vol. i. Léveillé, “Organisation, &c., sur
l‘Érysiphé,” in “Ann. des Sci. Nat.” (1851), vol. xv. p. 109.
[AD]
Other works besides those already cited, which may be consulted with advantage on
structure, are—
Tulasne, L. R. and C., various articles in “Annales des Sciences Naturelles,” série iii.
and iv.
Hoffmann, “Icones Analyticæ Fungorum.”
De Bary, “Der Ascomyceten.” Leipzic, 1863.
Berkeley, M. J., “Introduction to Cryptogamic Botany.”
Seynes, J. de, “Recherches, &c., des Fistulines.” Paris, 1874.
Winter, G., “Die Deutschen Sordarien.” 1874.
Corda, J., “Prachtflora.” Prague, 1840.
De Bary, “Über der Brandpilze.” 1853.
Brefeld, O., “Botan. Untersuch. ü Schimmelpilze.”
Fresenius, G., “Beiträge zur Mykologie.” 1850.
Von Tieghem and Le Monnier, in “Annales des Sciences Naturelles” (1873), p. 335.

Cornu, M., “Sur les Saprolegniées,” in “Ann. des Sci. Nat.” 5me sér. xv. p. 5.

Janczenski, “Sur l’Ascobolus furfuraceus,” in “Ann. des Sci. Nat.” 5me sér. xv. p.
200.
De Bary and Woronin, “Beiträge zur Morphologie und Physiologie der Pilze.” 1870.
Bonorden, H. F., “Abhandlungen aus dem Gebiete der Mykologie.” 1864.
Coemans, E., “Spicilége Mycologique.” 1862, etc.

III
CLASSIFICATION

A work of this kind could not be considered complete without some account
of the systematic arrangement or classification which these plants receive at
the hands of botanists. It would hardly avail to enter too minutely into
details, yet sufficient should be attempted to enable the reader to
comprehend the value and relations of the different groups into which fungi
are divided. The arrangement generally adopted is based upon the “Systema
Mycologicum” of Fries, as modified to meet the requirements of more
recent microscopical researches by Berkeley in his “Introduction,”[A] and
adopted in Lindley’s “Vegetable Kingdom.” Another arrangement was
proposed by Professor de Bary,[B] but it has never met with general
acceptance.
In the arrangement to which we have alluded, all fungi are divided into
two primary sections, having reference to the mode in which the
fructification is produced. In one section, the spores (which occupy nearly
the same position, and perform similar functions, to the seeds of higher
plants) are naked; that is, they are produced on spicules, and are not
enclosed in cysts or capsules. This section is called Sporifera, or spore-
bearing, because, by general consent, the term spore is limited in fungi to
such germ-cells as are not produced in cysts. The second section is termed
Sporidiifera, or sporidia-bearing, because in like manner the term sporidia
is limited to such germ-cells as are produced in cells or cysts. These cysts
are respectively known as sporangia, and asci or thecæ. The true meaning
and value of these divisions will be better comprehended when we have
detailed the characters of the families composing these two divisions.
First, then, the section Sporifera contains four families, in two of which a
hymenium is present, and in two there is no proper hymenium. The term
hymenium is employed to represent a more or less expanded surface, on
which the fructification is produced, and is, in fact, the fruit-bearing

surface. When no such surface is present, the fruit is borne on threads,
proceeding direct from the root-like filaments of the mycelium, or an
intermediate kind of cushion or stroma. The two families in which an
hymenium is present are called Hymenomycetes and Gasteromycetes. In the
former, the hymenium is exposed; in the latter, it is at first enclosed. We
must examine each of these separately.
The common mushroom may be accepted, by way of illustration, as a
type of the family Hymenomycetes, in which the hymenium is exposed, and
is, in fact, the most noticeable feature in the family from which its name is
derived. The pileus or cap bears on its under surface radiating plates or
gills, consisting of the hymenium, over which are thickly scattered the
basidia, each surmounted by four spicules, and on each spicule a spore.
When mature, these spores fall freely upon the ground beneath, imparting to
it the general colour of the spores. But it must be observed that the
hymenium takes the form of gill-plates in only one order of
Hymenomycetes, namely, the Agaricini; and here, as in Cantharellus, the
hymenium is sometimes spread over prominent veins rather than gills. Still
further divergence is manifest in the Polyporei, in which order the
hymenium lines the inner surface of pores or tubes, which are normally on
the under side of the pileus. Both these orders include an immense number
of species, the former more or less fleshy, the latter more or less tough and
leathery. There are still other forms and orders in this family, as the Hydnei,
in which the hymenium clothes the surface of prickles or spines, and the
Auricularini, in which the hymenium is entirely or almost even. In the two
remaining orders, there is a still further divergence from the mushroom
form. In the one called Clavariei, the entire fungus is either simply
cylindrical or club-shaped, or it is very much branched and ramified.
Whatever form the fungus assumes, the hymenium covers the whole
exposed surface. In the Tremellini, a peculiar structure prevails, which at
first seems to agree but little with the preceding. The whole plant is
gelatinous when fresh, lobed and convolute, often brain-like, and varying in
size, according to species, from that of a pin’s head to that of a man’s head.
Threads and sporophores are imbedded in the gelatinous substance,[C] so
that the fertile threads are in reality not compacted into a true hymenium.
With this introduction we may state that the technical characters of the
family are thus expressed:—

Hymenium free, mostly naked, or, if enclosed at first, soon exposed;
spores naked, mostly quaternate, on distinct spicules = Hymenomycetes.
In this family some mycologists believe that
fungi attain the highest form of development of
which they are capable, whilst others contend
that the fructification of the Ascomycetes is
more perfect, and that some of the noblest
species, such as the pileate forms, are entitled to
the first rank. The morel is a familiar example.
Whatever may be said on this point, it is
incontrovertible that the noblest and most
attractive, as well as the largest, forms are
classed under the Hymenomycetes.
In Gasteromycetes, the second family, a true Fig. 37.—Agaricus nudus.
hymenium is also present, but instead of being exposed it is for a long time
enclosed in an outer peridium or sac, until the spores are fully matured, or
the fungus is beginning to decay. The common puff-ball (Lycoperdon) is
well known, and will illustrate the principal feature of the family. Externally
there is a tough coat or peridium, which is at first pale, but ultimately
becomes brown. Internally is at first a cream-coloured, then greenish,
cellular mass, consisting of the sinuated hymenium and young spores,
which at length, and when the spores are fully matured become brownish
and dusty, the hymenium being broken up into threads, and the spores
become free. In earlier stages, and before the hymenium is ruptured, the
spores have been found to harmonize with those of Hymenomycetes in their
mode of production, since basidia are present surmounted each by four
spicules, and each spicule normally surmounted by a spore.[D] Here is,
therefore, a cellular hymenium bearing quaternary spores, but, instead of
being exposed, this hymenium is wholly enclosed within an external sac or
peridium, which is not ruptured until the spores are fully matured, and the
hymenium is resolved into threads, together forming a pulverulent mass. It
must, however, be borne in mind, that in only some of the orders composing
this family is the hymenium thus evanescent, in others being more or less
permanent, and this has led naturally enough to the recognition of two sub-
families, in one of which the hymenium is more or less permanent, thus
following the Hymenomycetous type; and in the other, the hymenium is

evanescent, and the dusty mass of spores tends more towards the
Coniomycetes, this being characterized as the coniospermous (or dusty-
spored) sub-family.
The first sub-family includes, first of all, the Hypogæi, or subterranean
species. And here again it becomes necessary to remind the reader that all
subterranean fungi are not included in this order, inasmuch as some, of
which the truffle is an example, are sporidiiferous, developing their sporidia
in asci. To these allusion must hereafter be made. In the Hypogæi, the
hymenium is permanent and convoluted, leaving numerous minute irregular
cavities, in which the spores are produced on sporophores. When specimens
are very old and decaying, the interior may become pulverulent or
deliquescent. The structure of subterranean fungi attracted the attention of
Messrs. Tulasne, and led to the production of a splendid monograph on the
subject.[E] Another order belonging to this sub-family is the Phalloidei, in
which the volva or peridium is ruptured whilst the plant is still immature,
and the hymenium when mature becomes deliquescent. Not only are some
members of this order most singular in appearance, but they possess an
odour so fœtid as to be unapproached in this property by any other
vegetable production.[F] In this order, the inner stratum of the investing
volva is gelatinous. When still young, and previous to the rupture of the
volva, the hymenium presents sinuous cavities in which the spores are
produced on spicules, after the manner of Hymenomycetes.[G] Nidulariacei
is a somewhat aberrant order, presenting a peculiar structure. The peridium
consists of two or three coats, and bursts at the apex, either irregularly or in
a stellate manner, or by the separation of a little lid. Within the cavity are
contained one or more secondary receptacles, which are either free or
attached by elastic threads to the common receptacle. Ultimately the
secondary receptacles are hollow, and spores are produced in the interior,
borne on spicules.[H] The appearance in some genera as of a little bird’s-
nest containing eggs has furnished the name to the order.
The second sub-family contains the coniospermous puff-balls, and
includes two orders, in which the most readily distinguishable feature is the
cellular condition of the entire plant, in its earlier stages, in the
Trichogastres, and the gelatinous condition of the early state of the
Myxogastres. Both are ultimately resolved internally into a dusty mass of
threads and spores. In the former, the peridium is either single or double,

occasionally borne on a stem, but usually
sessile. In Geaster, the “starry puff-balls,” the
outer peridium divides into several lobes, which
fall back in a stellate manner, and expose the
inner peridium, like a ball in the centre. In
Polysaccum, the interior is divided into
numerous cells, filled with secondary peridia.
The mode of spore-production has already been
alluded to in our remarks on Lycoperdon. All the
species are large, as compared with those of the
following sub-family, and one species of Fig. 38.—Scleroderma vulgare,
Lycoperdon attains an enormous size. One Fr.
specimen recorded in the “Gardener’s Chronicle” was three feet four inches
in circumference, and weighed nearly ten pounds. In the Myxogastres, the
early stage has been the subject of much controversy. The gelatinous
condition presents phenomena so unlike anything previously recorded in
plants, that one learned professor[I] did not hesitate to propose their
exclusion from the vegetable, and recognition in the animal, kingdom as
associates of the Gregarines. When mature, the spores and threads so much
resemble those of the Trichogastres, and the little plants themselves are so
veritably miniature puff-balls, that the theory of their animal nature did not
meet with a ready acceptance, and is now virtually abandoned. The
characters of the family we have thus briefly reviewed are tersely stated, as
—
Hymenium more or less permanently concealed, consisting in most cases
of closely-packed cells, of which the fertile ones bear naked spores on
distinct spicules, exposed only by the rupture or decay of the investing coat
or peridium = Gasteromycetes.

Fig. 39.—Ceuthospora phacidioides (Greville).

We come now to the second section of the Sporifera, in which no definite
hymenium is present. And here we find also two families, in one of which
the dusty spores are the prominent feature, and hence termed Coniomycetes;
the other, in which the threads are most noticeable, is Hyphomycetes. In the
former of these, the reproductive system seems to preponderate so much
over the vegetative, that the fungus appears to be all spores. The mycelium
is often nearly obsolete, and the short pedicels so evanescent, that a rusty or
sooty powder represents the mature fungus, infesting the green parts of
living plants. This is more especially true of one or two orders. It will be
most convenient to recognize two artificial sub-families for the purpose of
illustration, in one of which the species are developed on living, and in the
other on dead, plants. We will commence with the latter, recognizing first
those which are developed beneath the cuticle, and then those which are
superficial. Of the sub-cuticular, two orders may be named as the
representatives of this group in Britain, these are the Sphæronemei, in
which the spores are contained in a more or less perfect perithecium, and
the Melanconiei, in which there is manifestly none. The first of these is
analogous to the Sphæriacei of Ascomycetous fungi, and probably consists
largely of spermogonia of known species of Sphæria, the relations of which
have not hitherto been traced. The spores are produced on slender threads
springing from the inner wall of the perithecium, and, when mature, are
expelled from an orifice at the apex. This is the normal condition, to which
there are some exceptions. In the Melanconiei, there is no true perithecium,
but the spores are produced in like manner upon a kind of stroma or cushion
formed from the mycelium, and, when mature, are expelled through a
rupture of the cuticle beneath which they are generated, often issuing in

long gelatinous tendrils. Here, again, the majority of what were formerly
regarded as distinct species have been found, or suspected, to be forms of
higher fungi. The Torulacei represent the superficial fungi of this family,
and these consist of a more or less developed mycelium, which gives rise to
fertile threads, which, by constriction and division, mature into moniliform
chains of spores. The species mostly appear as blackish velvety patches or
stains on the stems of herbaceous plants and on old weathered wood.
Much interest attaches to the other sub-family of Coniomycetes, in which
the species are produced for the most part on living plants. So much has
been discovered during recent years of the polymorphism which subsists
amongst the species in this section, that any detailed classification can only
be regarded as provisional. Hence we shall proceed here upon the
supposition that we are dealing with autonomous species. In the first place,
we must recognize a small section in which a kind of cellular peridium is
present. This is the Æcidiacei, or order of “cluster cups.” The majority of
species are very beautiful objects under the microscope; the peridia are
distinctly cellular, and white or pallid, produced beneath the cuticle, through
which they burst, and, rupturing at the apex, in one genus in a stellate
manner, so that the teeth, becoming reflexed, resemble delicate fringed
cups, with the orange, golden, brown, or whitish spores or pseudospores
nestling in the interior.[J] These pseudospores are at first produced in chains,
but ultimately separate. In many cases these cups are either accompanied or
preceded by spermogonia. In two other orders there is no peridium. In the
Cæomacei, the pseudospores are more or less globose or ovate, sometimes
laterally compressed and simple; and in Pucciniæi, they are elongated, often
subfusiform and septate. In both, the pseudospores are produced in tufts or
clusters direct from the mycelium. The Cæomacei might again be
subdivided into Ustilagines[K] and Uredines.[L] In the former, the
pseudospores are mostly dingy brown or blackish, and in the latter more
brightly coloured, often yellowish. The Ustilagines include the smuts and
bunt of corn-plants, the Uredines include the red rusts of wheat and grasses.
In some of the species included in the latter, two forms of fruit are found. In
Melampsora, the summer pseudospores are yellow, globose, and were
formerly classed as a species of Lecythea, whilst the winter pseudospores
are brownish, elongated, wedge-shaped by compression, and compact. The
Pucciniæi[M] differ primarily in the septate pseudospores, which in one
genus (Puccinia) are uniseptate; in Triphragmium, they are biseptate; in

Phragmidium, multiseptate; and in Xenodochus, moniliform, breaking up
into distinct articulations. It is probable that, in all of these, as is known to
be the case in most, the septate pseudospores are preceded or accompanied
by simple pseudospores, to which they are mysteriously related. There is
still another, somewhat singular, group usually associated with the
Pucciniæi, in which the septate pseudospores are immersed in gelatin, so
that in many features the species seem to approach the Tremellini. This
group includes two or three genera, the type of which will be found in
Podisoma.[N] These fungi are parasitic on living junipers in Britain and
North America, appearing year after year upon the same gouty swellings of
the branches, in clavate or horn-shaped gelatinous processes of a yellowish
or orange colour. Anomalous as it may at first sight appear to include these
tremelloid forms with the dust-like fungi, their relations will on closer
examination be more fully appreciated, when the form of pseudospores,
mode of germination, and other features are taken into consideration,
especially when compared with Podisoma Ellisii, already alluded to. This
family is technically characterized as,—
Distinct hymenium none. Pseudospores either solitary or concatenate,
produced on the tips of generally short threads, which are either naked or
contained in a perithecium, rarely compacted into a gelatinous mass, at
length producing minute spores = Coniomycetes.
The last family of the sporifera is Hyphomycetes, in which the threads are
conspicuously developed. These are what are more commonly called
“moulds,” including some of the most elegant and delicate of microscopic
forms. It is true of many of these, as well as of the Coniomycetes, that they
are only conidial forms of higher fungi; but there will remain a very large
number of species which, as far as present knowledge extends, must be
accepted as autonomous. In this family, we may again recognize three
subdivisions, in one of which the threads are more or less compacted into a
common stem, in another the threads are free, and in the third the threads
can scarcely be distinguished from the mycelium. It is this latter group
which unites the Hyphomycetes with the Coniomycetes, the affinities being
increased by the great profusion with which the spores are developed. The
first group, in which the fertile threads are united so as to form a compound
stem, consists of two small orders, the Isariacei and the Stilbacei, in the
former of which the spores are dry, and in the latter somewhat gelatinous.

Many of the species closely imitate forms met with in the Hymenomycetes,
such as Clavaria; and, in the genus Isaria, it is almost beyond doubt that the
species found on dead insects, moths, spiders, flies, ants, &c., are merely
the conidiophores of species of Torrubia.[O]
The second group is by far the largest, most typical, and attractive in this
family. It contains the black moulds and white moulds, technically known
as the Dematiei and the Mucedines. In the first, the threads are more or less
corticated, that is, the stem has a distinct investing membrane, which peels
off like a bark; and the threads, often also the spores, are dark-coloured, as
if charred or scorched. In many cases, the spores are highly developed,
large, multiseptate, and nucleate, and seldom are spores and threads
colourless or of bright tints. In the Mucedines, on the contrary, the threads
are never coated, seldom dingy, mostly white or of pure colours, and the
spores have less a tendency to extra development or multiplex septation. In
some genera, as in Peronospora for instance,[P] a secondary fruit is
produced in the form of resting spores from the mycelium; and these
generate zoospores as well as the primary spores, similar to those common
in Algæ. This latter genus is very destructive to growing plants, one species
being the chief agent in the potato disease, and another no less destructive
to crops of onions. The vine disease is produced by a species of Oidium,
which is also classed with Mucedines, but which is really the conidiiferous
form of Erysiphe. In other genera, the majority of species are developed on
decaying plants, so that, with the exception of the two genera mentioned,
the Hyphomycetes exert a much less baneful influence on vegetation than
the Coniomycetes. The last section, including the Sepedoniei, has been
already cited as remarkable for the suppression of the threads, which are
scarcely to be distinguished from the mycelium; the spores are profuse,
nestling on the floccose mycelium; whilst in the Trichodermacei, the spores
are invested by the threads, as if enclosed in a sort of false peridium. A
summary of the characters of the family may therefore be thus briefly
expressed:—
Filamentous; fertile threads naked, for the most part free or loosely
compacted, simple or branched, bearing the spores at their apices, rarely
more closely packed, so as to form a distinct common stem =
Hyphomycetes.

Having thus disposed of the Sporifera, we
must advert to the two families of Sporidiifera.
As more closely related to the Hyphomycetes, the
first of these to be noticed is the Physomycetes, in
which there is no proper hymenium, and the
threads proceeding from the mycelium bear
vesicles containing an indefinite number of
sporidia. The fertile threads are either free or only
slightly felted. In the order Antennariei, the
threads are black and moniliform, more or less Fig. 40.—Rhopalomyces
felted, bearing irregular sporangia. A common candidus.
fungus named Zasmidium cellare, found in
cellars, and incrusting old wine bottles, as with a blackened felt, belongs to
this order. The larger and more highly-developed order, Mucorini, differs in
the threads, which are simple or branched, being free, erect, and bearing the
sporangia at the tips of the thread, or branches. Some of the species bear
great external resemblance to Mucedines until the fruit is examined, when
the fructifying heads, commonly globose or ovate, are found to be delicate
transparent vesicles, enclosing a large number of minute sporidia; when
mature, the sporangia burst and the sporidia are set free. In some species, it
has long been known that a sort of conjugation takes place between
opposite threads, which results in the formation of a sporangium.[Q] None
of these species are destructive to vegetation, appearing only upon
decaying, and not upon living, plants. A state approaching putrescence
seems to be essential to their vigorous development. The following
characters may be compared with those of the family preceding it:—
Filamentous, threads free or only slightly felted, bearing vesicles, which
contain indefinite sporidia = Physomycetes.
In the last family, the Ascomycetes, we shall meet with a very great
variety of forms, all agreeing in producing sporidia contained in certain
cells called asci, which are produced from the hymenium. In some of these,
the asci are evanescent, but in the greater number are permanent. In
Onygenei, the receptacle is either club-shaped or somewhat globose, and
the peridium is filled with branched threads, which produce asci of a very
evanescent character, leaving the pulverulent sporidia to fill the central
cavity. The species are all small, and singular for their habit of affecting

animal substances, otherwise they are of little
importance. The Perisporiacei, on the other hand,
are very destructive of vegetation, being
produced, in the majority of cases, on the green
parts of growing plants. To this order the hop
mildew, rose mildew, and pea mildew belong.
The mycelium is often very much developed, and
in the case of the maple, pea, hop, and some
others, it covers the parts attacked with a thick
white coating, so that from a distance the leaves Fig. 41.—Mucor caninus.
appear to have been whitewashed. Seated on the mycelium, at the first as
little orange points, are the perithecia, which enlarge and become nearly
black. In some species, very elegant whitish appendages radiate from the
sides of the perithecia, the variations in which aid in the discrimination of
species. The perithecia contain pear-shaped asci, which spring from the
base and enclose a definite number of sporidia.[R] The asci themselves are
soon dissolved. Simultaneously with the development of sporidia, other
reproductive bodies are produced direct from the mycelium, and in some
species as many as five different kinds of reproductive bodies have been
traced. The features to be remembered in Perisporiacei, as forming the
basis of their classification, are, that the asci are saccate, springing from the
base of the perithecia, and are soon absorbed. Also that the perithecia
themselves are not perforated at the apex.
The four remaining orders, though large, can be easily characterized. In
Tuberacei, all the species are subterranean, and the hymenium is mostly
sinuated. In Elvellacei, the substance is more or less fleshy, and the
hymenium is exposed. In Phacidiacei, the substance is hard or leathery, and
the hymenium is soon exposed. And in Sphæriacei, although the substance
is variable, the hymenium is never exposed, being enclosed in perithecia
with a distinct opening at the apex, through which the mature spores escape.
Each of these four orders must be examined more in detail. The Tuberacei,
or subterranean Ascomycetes, are analogous to the Hypogæi of the
Gasteromycetes. The truffle is a familiar and highly prized example. There
is a kind of outer peridium, and the interior consists of a fleshy hymenium,
more or less convoluted, sometimes sinuous and confluent, so as to leave
only minute elongated and irregular cavities, and sometimes none at all, the
two opposing faces of the hymenium meeting and coalescing.[S] Certain

privileged cells of the hymenium swell, and ultimately become asci,
enclosing a definite number of sporidia. The sporidia in many cases are
large, reticulated, echinulate or verrucose, and mostly somewhat globose. In
the genus Elaphomyces, the asci are more than commonly diffluent.
The Elvellacei are fleshy in substance, or somewhat waxy, sometimes
tremelloid. There is no peridium, but the hymenium is always exposed.
There is a great variety of forms, some being pileate, and others cup-
shaped, as there is also a great variation in size, from the minute Peziza,
small as a grain of sand, to the large Helvella gigas, which equals in
dimensions the head of a child. In the pileate forms, the stroma is fleshy and
highly developed; in the cup-shaped, it is reduced to the external cells of the
cup which enclose the hymenium. The hymenium itself consists of
elongated fertile cells, or asci, mixed with linear thread-like barren cells,
called paraphyses, which are regarded by some authors as barren asci.
These are placed side by side in juxtaposition with the apex outwards. Each
ascus contains a definite number of sporidia, which are sometimes
coloured. When mature, the asci explode above, and the sporidia may be
seen escaping like a miniature cloud of smoke in the light of the mid-day
sun. The disc or surface of the hymenium is often brightly coloured in the
genus Peziza; tints of orange, red, and brown having the predominance.
In Phacidiacei, the substance is hard and leathery, intermediate between
the fleshy Elvellacei and the more horny of the Sphæriacei. The perithecia
are either orbicular or elongated, and the hymenium soon becomes exposed.
In some instances, there is a close affinity with the Elvellacei, the exposed
hymenium being similar in structure, but in all the disc is at first closed. In
orbicular forms, the fissure takes place in a stellate manner from the centre,
and the teeth are reflexed. In the Hysteriacei, where the perithecia are
elongated, the fissure takes place throughout their length. As a rule, the
sporidia are more elongated, more commonly septate, and more usually
coloured, than in Elvellacei. Only a few solitary instances occur of
individual species that are parasitic on living plants.
In the Sphæriacei, the substance of the stroma (when present) and of the
perithecia is variable, being between fleshy and waxy in Nectriei, and
tough, horny, sometimes brittle, in Hypoxylon. A perithecium, or cell
excavated in the stroma which fulfils the functions of a perithecium, is
always present. The hymenium lines the inner walls of the perithecium, and

forms a gelatinous nucleus, consisting of asci
and paraphyses. When fully mature, the asci
are ruptured and the sporidia escape by a pore
which occupies the apex of the perithecium.
Sometimes the perithecia are solitary or
scattered, and sometimes gregarious, whilst in
other instances they are closely aggregated and
immersed in a stroma of variable size and
form. Conidia, spermatia, pycnidia, &c., have Fig. 42.—Sphæria aquila.
been traced to and associated with some
species, but the history of others is still obscure. Many of the
coniomycetous forms grouped under the Sphæronemei are probably
conditions of the Sphæriacei, as are also the Melanconiei, and some of the
Hyphomycetes. A very common fungus, for instance, which is abundant on
sticks and twigs, forming rosy or reddish pustules the size of a millet seed,
formerly named Tubercularia vulgaris, is known to be the conidia-bearing
stroma of the sphæriaceous fungus, Nectria cinnabarina;[T] and so with
many others. The following are the technical characters of the family:—
Fruit consisting of sporidia, mostly definite, contained in asci, springing
from a naked or enclosed stratum of fructifying cells and forming a
hymenium or nucleus = Ascomycetes.
If the characters of the different families are borne in mind, there will be
but little difficulty in assigning any fungus to the order to which it belongs
by means of the foregoing remarks. For more minute information, and for
analytical tables of the families, orders, and genera, we must refer the
student to some special systematic work, which will present fewer
difficulties, if he keeps in mind the distinctive features of the families.[U]
To assist in this we have given on the following page an analytical
arrangement of the families and orders, according to the system recognized
and adopted in the present volume. It is, in all essential particulars, the
method adopted in our “Handbook,” based on that of Berkeley’s
“Introduction” and “Outlines.”
[A]
Rev. M. J. Berkeley, “Introduction to Cryptogamic Botany” (1857), London, pp. 235
to 372.

[B]
De Bary, in “Streinz Nomenclator Fungorum,” p. 722.
[C]
Tulasne, L. and C. R., “Observations sur l’Organisation des Trémellinées,” “Ann. des
Sci. Nat.” 1853, xix. p. 193.
[D]
Berkeley, M. J., “On the Fructification of Lycoperdon, Phallus, and their Allied
Genera,” in “Ann. of Nat. Hist.” (1840), vol. iv. p. 155; “Ann. des Sci. Nat.” (1839),
xii. p. 163. Tulasne, L. R. and C., “De la Fructification des Scléroderma comparée à
celle des Lycoperdon et des Bovista,” in “Ann. des Sci. Nat.” 2me sér. xvii. p. 5.
[E]
Tulasne, L. R. and C., “Fungi Hypogæi,” Paris, 1851; “Observations sur le Genre
Elaphomyces,” in “Ann. des Sci. Nat.” 1841, xvi. 5.
[F]
Stapeliæ in this respect approach most closely to the Phalloidei.
[G]
Berkeley, in “Ann. Nat. Hist.” vol. iv. p. 155.
[H]
Tulasne, L. R. and C., “Recherches sur l’Organisation et le Mode de Fructification
des Nidulariées,” “Ann. des Sci. Nat.” (1844), i. p. 41.
[I]

De Bary, A., “Des Myxomycètes,” in “Ann. des Sci. Nat.” 4me sér. xi. p. 153; “Bot.
Zeit.” xvi. p. 357.
[J]
Corda, “Icones Fungorum,” vol. iii. fig. 45.
[K]
Tulasne, “Mémoire sur les Ustilaginées,” “Ann. des Sci. Nat.” (1847), vii. 12–73.
[L]
Tulasne, “Mémoire sur les Urédinées,” “Ann. des Sci. Nat.” (1854), ii. 78.
[M]
Tulasne, “Sur les Urédinées,” “Ann. des Sci. Nat.” 1854, ii. pl. 9.
[N]
Cooke, M. C., “Notes on Podisoma,” in “Journ. Quek. Micr. Club,” No. 17 (1871), p.
255.

[O]
Tulasne, L. R. and C., “Selecta Fungorum Carpologia,” vol. iii. pp. 4–19.
[P]
De Bary, A., “Recherches sur les Champignons Parasites,” in “Ann. des Sci. Nat.”
4me sér. xx. p. 5; “Grevillea,” vol. i. p. 150.
[Q]

A. de Bary, translated in “Grevillea,” vol. i. p. 167; Tulasne, “Ann. des Sci. Nat.” 5me
sér. (1866), p. 211.
[R]
Léveillé, J. H., “Organisation, &c., de l‘Érysiphé,” in “Ann. des Sci. Nat.” (1851), xv.
p. 109.
[S]
Tulasne, L. R. and C., “Fungi Hypogæi,” Paris; Vittadini, C., “Monographia
Tuberacearum,” Milan, 1831.
[T]
“A Currant Twig and Something on it,” in “Gardener’s Chronicle” for January 28,
1871.
[U]
Berkeley, M. J., “Introduction to Cryptogamic Botany,” London, 1857; Cooke, M. C.,
“Handbook of British Fungi,” London, 1871 ; Corda, A. C. J., “Anleitung zum
Studium der Mycologie,” Prag, 1842; Kickx, J., “Flore Cryptogamique des Flanders,”
Gand, 1867; Fries, E., “Systema Mycologicum,” Lund, 1830; Fries, E., “Summa
Vegetabilium Scandinaviæ,” 1846; Secretan, L., “Mycographie Suisse,” Geneva,
1833; Berkeley, M. J., “Outlines of British Fungology,” London, 1860.

TABULAR ARRANGEMENT OF FAMILIES AND ORDERS.

Division I. SPORIFERA. Spores naked.

I. Hymenium free, mostly naked, or soon exposed Hymenomycetes.
Hymenium normally inferior—
Fruit-bearing surface lamellose Agaricini.

Fruit-bearing surface porous or tubular Polyporei.
Fruit-bearing surface clothed with prickles Hydnei.
Fruit-bearing surface even or rugose Auricularini.
Hymenium superior or encircling—
Clavate, or branched, rarely lobed Clavariei.
Lobed, convolute, or disc-like, gelatinous Tremellini.

II. Hymenium enclosed in a peridium, ruptured when
mature Gasteromycetes.
Hymenomycetous—
Subterranean, naked or enclosed Hypogæi.
Terrestrial, hymenium deliquescent Phalloidei.
Peridium enclosing sporangia, containing spores Nidulariacei.
Coniospermous—
Stipitate, hymenium convolute, drying into a dusty
mass, enclosed in a volva Podaxinei.
Cellular at first, hymenium drying up into a dusty mass
of threads and spores Trichogastres.
Gelatinous at first, peridium containing at length a
dusty mass of threads and spores Myxogastres.

III. Spores naked, mostly terminal, on inconspicuous
threads, free or enclosed in a perithecium Coniomycetes.
Growing on dead or dying plants—
Subcutaneous—
Perithecium more or less distinct Sphæronemei.
Perithecium obsolete or wanting Melanconiei.
Superficial—
Fructifying surface naked.
Spores compound or tomiparous Torulacei.
Parasitic on living plants—
Peridium distinctly cellular Æcidiacei.

Peridium none—
Spores sub-globose, simple or deciduous Cæomacei.
Spores mostly oblong, usually septate Pucciniæi.

IV. Spores naked, on conspicuous threads, rarely
compacted, small Hyphomycetes.
Fertile threads compacted, sometimes cellular—
Stem or stroma compound—
Spores dry, volatile Isariacei.
Mass of spores moist, diffluent Stilbacei.
Fertile threads, free or anastomosing—
Fertile threads dark, carbonized—
Spores mostly compound Dematiei.
Fertile threads not carbonized—
Very distinct—
Spores mostly simple Mucedines.
Scarcely distinct from mycelium—
Spores profuse Sepedoniei.

Division II. SPORIDIIFERA. Sporidia in Asci.

V. Fertile cells seated on threads, not compacted into a
Physomycetes.
hymenium
Threads felted, moniliform—
Sporangia irregular Antennariei.
Threads free—
Sporangia terminal or lateral Mucorini.
Aquatic Saprolegniei.

VI. Asci formed from the fertile cells of a hymenium Ascomycetes.
Asci often evanescent—
Receptacle clavæform—

Asci springing from threads Onygenei.
Perithecia free—
Asci springing from the base Perisporiacei.
Asci persistent—
Perithecia opening by a distinct ostiolum Sphæriacei.
Hard or coriaceous, hymenium at length exposed Phacidiacei.
Hypogæous; hymenium complicated Tuberacei.
Fleshy, waxy, or tremelloid; hymenium mostly exposed Elvellacei.

IV.
USES.

The rigid utilitarian will hardly be satisfied with the short catalogue which
can be furnished of the uses of fungi. Excepting those which are employed
more or less for human food, very few are of any practical value in arts or
medicine. It is true that imperfect conditions of fungi exert a very important
influence on fermentation, and thus become useful; but, unfortunately, fungi
have the reputation of being more destructive and offensive than valuable or
useful. Notwithstanding that a large number of species have from time to
time been enumerated as edible, yet those commonly employed and
recognized are very few in number, prejudice in many cases, and fear in
others, militating strongly against additions to the number. In Great Britain
this is especially the case, and however advisable it may be to exercise great
care and caution in experimenting on untried or doubtful species, it can only
be regarded as prejudice which prevents good, in fact, excellent, esculent
species being more extensively used, instead of allowing them to rot by
thousands on the spots where they have grown. Poisonous species are also
plentiful, and no golden rule can be established by means of which any one
may detect at a glance good from bad, without that kind of knowledge
which is applied to the discrimination of species. Yet, after all, the
characters of half a dozen good esculent fungi are acquired as easily as the
distinctions between half a dozen birds such as any ploughboy can
discriminate.
The common mushroom (Agaricus campestris) is the best known
esculent, whether in its uncultivated or in a cultivated state. In Britain many
thousands of people, notably the lower classes, will not recognize any other
as fit for food, whilst in Italy the same classes have a strong prejudice
against this very species.[A] In Vienna, we found by personal experience
that, although many others are eaten, it is this which has the most universal
preference, yet it appears but sparingly in the markets as compared with

others. In Hungary it does not enjoy by any means so good a reputation. In
France and in Germany it is a common article of consumption. The
different varieties found, as the results of cultivation, present some variation
in colour, scaliness of pileus, and other minor features, whilst remaining
true to the constituent characters of the species. Although it is not our
intention to enumerate here the botanical distinctions of the species to
which we may call attention, yet, as mistakes (sometimes fatal) are often
being recorded, in which other fungi are confounded with this, we may be
permitted a hint or two which should be remembered. The spores are
purple, the gills are at first delicate pink, afterwards purple; there is a
permanent ring or collar round the stem, and it must not be sought in
woods. Many accidents might have been spared had these facts been
remembered.
The meadow mushroom (Agaricus arvensis) is common in meadows and
lowland pastures, and is usually of a larger size than the preceding, with
which it agrees in many particulars, and is sent in enormous quantities to
Covent Garden, where it frequently predominates over Agaricus
campestris. Some persons prefer this, which has a stronger flavour, to the
ordinary mushroom, and it is the species most commonly sold in the
autumn in the streets of London and provincial towns. According to
Persoon, it is preferred in France; and, in Hungary, it is considered as a
special gift from St. George. It has acquired in England the name of horse
mushroom, from the enormous size it sometimes attains. Withering
mentions a specimen that weighed fourteen pounds.[B]
One of the commonest (in our experience the most common) of all edible
fungi in the public markets of Vienna is the Hallimasche (Agaricus
melleus), which in England enjoys no good reputation for flavour or
quality; indeed, Dr. Badham calls it “nauseous and disagreeable,” and adds
that “not to be poisonous is its only recommendation.” In Vienna it is
employed chiefly for making sauce; but we must confess that even in this
way, and with a prejudice in favour of Viennese cookery, our experience of
it was not satisfactory. It is at best a sorry substitute for the mushroom. In
the summer and autumn this is a very common species in large tufts on old
stumps. In similar localities, and also in tufts, but neither so large, nor so
common, Agaricus fusipes is found. It is preferable to the foregoing as an
esculent, and is easily recognized by the spindle-shaped stem.

Agaricus rubescens, P., belongs to a very suspicious group of fungi, in
which the cap or pileus is commonly studded or sprinkled with paler warts,
the remains of an investing volva. To this group the poisonous but splendid
fly-agaric (Agaricus muscarius) belongs. Notwithstanding its bad company,
this agaric has a good reputation, especially for making ketchup; and
Cordier reports it as one of the most delicate mushrooms of the Lorraine.[C]
Its name is derived from its tendency to become red when bruised.
The white variety of an allied species (Agaricus vaginatus) has been
commended, and Dr. Badham says that it will be found inferior to but few
agarics in flavour.
A scaly-capped fungus (Agaricus procerus), with a slender stem, called
sometimes the parasol mushroom, from its habit, is an esteemed esculent. In
Italy and France it is in high request, and is included in the majority of
continental works on the edible fungi.[D] In Austria, Germany, and Spain, it
has special “vulgar” names, and is eaten in all these countries. It is much
more collected in England than formerly, but deserves to be still better
known. When once seen it can scarcely be confounded with any other
British species, save one of its nearest allies, which partakes of its own
good qualities (Agaricus rachodes), though not quite so good.
Agaricus prunulus, Scop., and Agaricus orcella, Badh., if they be not
forms of the same species (which Dr. Bull contends that they are not[E]),
have also a good reputation as esculents. They are both neat, white agarics,
with a mealy odour, growing respectively in woods and open glades.
Agaricus nebularis, Batsch, is a much larger species, found in woods, often
in large gregarious patches amongst dead leaves, with a smoky mouse-
coloured pileus, and profuse white spores. It is sometimes as much as five
or six inches in diameter, with rather a faint odour and mild taste. On the
continent, as well as in Britain, this is included amongst edible fungi. Still
larger and more imposing is the magnificent white species, Agaricus
maximus, Fr.,[F] which is figured by Sowerby,[G] under the name of
Agaricus giganteus. It will attain a diameter of fourteen inches, with a stem,
two inches thick, and rather a strong odour.
A spring fungus, the true St. George’s mushroom, Agaricus gambosus,
Fr., makes its appearance in pastures, usually growing in rings, in May and
June, and is welcome to mycophagists from its early growth, when esculent

species are rare. It is highly esteemed in France and Italy, so that when dried
it will realize as much as from twelve to fifteen shillings per pound.
Guillarmod includes it amongst Swiss esculents.[H] Professor Buckman says
that it is one of the earliest and best of English mushrooms, and others have
endorsed his opinions, and Dr. Badham in writing of it observes, that small
baskets of them, when they first appear in the spring in Italy, are sent as
“presents to lawyers and fees to medical men.”
The closely allied species, Agaricus albellus,[I] D.C., has also the
reputation of being edible, but it is so rare in England that this quality
cannot be put to the test. The curious short-stemmed Agaricus brevipes,
Bull,[J] has a similar reputation.
Two singularly fragrant species are also included amongst the esculent.
These are Agaricus fragrans, Sow., and Agaricus odorus, Bull. Both have a
sweet anise-like odour, which is persistent for a long time. The former is
pale tawny-coloured, nearly white, the latter of a dirty pale green. Both are
white-spored, and although somewhat local, sufficient specimens of Ag.
odorus may be collected in the autumn for domestic use. We have the
assurance of one who has often proved them that they constitute an
exquisite dish.
A clear ivory-white fungus, Agaricus dealbatus, of which a crisped
variety is occasionally found in great numbers, springing up on old
mushroom beds in dense clusters, is very good eating, but rather deficient in
the delicate aroma of some other species. The typical form is not
uncommon on the ground in fir plantations. A more robust and larger
species, Agaricus geotrupes, Bull, found on the borders of woods, often
forming rings, both in this country and in the United States, as well as on
the continent of Europe, is recognized as esculent.
We may add to these three or four other species, in which the stem is
lateral, and sometimes nearly obsolete. The largest and most common is the
oyster mushroom (Agaricus ostreatus, Jacq.[K]), so universally eaten, that it
is included in almost every list and book on edible fungi; it is the most
common species in Transylvania, tons of it sometimes appearing in the
markets. It does not possess that delicate flavour which is found in many
species, and although extolled by some beyond its merits, it is nevertheless
perfectly wholesome, and, when young and carefully cooked, not to be

despised. It must not be confounded with a very similar species (Agaricus
euosmus, B.), with rosy spores, which is unpleasant. Agaricus tessellatus,
Bull, Agaricus pometi, Fr., Agaricus glandulosus, Bull, are all allies of the
foregoing, and recorded as edible in the United States, although not one of
the three has hitherto been recorded as occurring in Great Britain. To these
may also be added the following:—Agaricus salignus,[L] Fr., which is rare
in England, but not uncommon abroad and in the United States. In Austria it
is commonly eaten. Agaricus ulmarius,[M] Bull, is common on elm trunks,
not only in Britain but also in North America, and is by some preferred to
the oyster mushroom. An allied species, Agaricus fossulatus, Cooke,[N] is
found on the Cabul Hills, where it is collected, dried, and forms an article of
commerce with the plains. Another, but smaller species, is dried in the air
on strings passed through a hole in the short stem (Agaricus subocreatus,
Cooke), and sent, it is believed, from China to Singapore.
The smallest species with which we have any acquaintance, that is edible,
is the “nail fungus” (Agaricus esculentus,[O] Jacq.), scarcely exceeding one
inch in diameter of the pileus, with a thin rooting stem. The taste in British
specimens when raw is bitter and unpleasant, but it is clearly eaten in
Austria, as its name testifies, and elsewhere in Europe. It is found in fir
plantations in the spring, at which season it is collected from the fir woods
around and sent to Vienna, where it is only used for flavouring sauces under
the name of “Nagelschwämme.”
Before quitting the group of true agarics, to which all hitherto
enumerated belong, we must mention a few others of less importance, but
which are included amongst those good for food. Foremost of these is a
really splendid orange species (Agaricus cæsarius, Scop.[P]), which belongs
to the same subgenus as the very deleterious fly-agaric, and the scarcely
less fatal Agaricus vernus, Bull. It is universally eaten on the continent, but
has hitherto never been found in Great Britain. In the same subgenus,
Agaricus strobiliformis,[Q] Fr., which is rare in this country, and probably
also Agaricus Ceciliæ, B. & Br.[R] Besides these, Agaricus excoriatus,
Schæff., Agaricus mastoideus, Fr., Agaricus gracilentus, Kromb., and
Agaricus holosericeus, Fr.,[S] all belonging to the same subgenus as the
parasol mushroom, more or less uncommon in England.
Although the larger number of esculent agarics are white-spored, some
few, worthy of note, will be found in the other sections, and notably

amongst these the common mushroom and its congener the meadow, or
horse mushroom. In addition to those already enumerated, might be
included also the Agaricus pudicus, Bull, which is certainly wholesome, as
well as its ally, Agaricus leochromus, Cooke,[T] both of which have rusty
spores.
The late Dr. Curtis,[U] in a letter to the Rev. M. J. Berkeley, enumerates
several of the fungi which are edible amongst those found in the United
States. Of these, he says, Agaricus amygdalinus, Curt., can scarcely be
distinguished when cooked from the common mushroom. Agaricus
frumentaceus, Bull, and three allied new species, peculiar to the United
States, are commended. Agaricus cæspitosus, Curt., he says, is found in
enormous quantities, a single cluster containing from fifty to one hundred
stems, and might well be deemed a valuable species in times of scarcity. It
would not be highly esteemed where other and better species can be had,
but it is generally preferred to Agaricus melleus, Fr. It is suitable for drying
for winter use. In the same communication, he observes that the imperial
(Agaricus cæsarius, Scop.), grows in great quantities in oak forests, and
may be obtained by the cart-load in its season; but to his taste, and that of
his family, it is the most unpalatable of fungi, nor could he find any of the
most passionate mycophagists who would avow that they liked it. There is a
disagreeable saline flavour that they could not remove nor overlay. In
addition to these, the same authority enumerates Agaricus russula, Schæff.,
Agaricus hypopithyus, Curt., and Agaricus consociatus, Curt., the latter two
being confined to the United States; Agaricus columbetta, Fr., found in
Britain, but not eaten, as well as Agaricus radicatus, Bull. Agaricus
bombycinus, Schæff., and Agaricus speciosus, Fr., are found in Britain, but
by no means common; Agaricus squarrosus, Mull., has always been
regarded with great suspicion in this country, where it is by no means
uncommon; Agaricus cretaceus, Fr., and Agaricus sylvaticus, Schæff., are
close allies of the common mushroom.
Dr. Curtis says that hill and plain, mountain and valley, woods, fields,
and pastures, swarm with a profusion of good nutritious fungi, which are
allowed to decay where they spring up, because people do not know how, or
are afraid, to use them. By those of us who know their use, their value was
appreciated, as never before, during the late war, when other food,
especially meat, was scarce and dear. Then such persons as I have heard

express a preference for mushrooms over meat had generally no need to
lack grateful food, as it was easily had for the gathering, and within easy
distance of their homes if living in the country. Such was not always the
case, however. I remember once, during the gloomy period when there had
been a protracted drought, and fleshy fungi were to be found only in damp
shaded woods, and but few even there, I was unable to find enough of any
one species for a meal, so, gathering of every kind, I brought home thirteen
different kinds, had them all cooked together in one grand pot pourri, and
made an excellent supper.
One important use to which several species of fungi can be applied, is the
manufacture of ketchup. For this purpose, not only is the mushroom,
Agaricus campestris, and the horse mushroom, Agaricus arvensis,
available, but also Agaricus rubescens is declared to be excellent for the
purpose, and a delicious, but pale, extract is to be obtained from Marasmius
oreades. Other species, as Coprinus comatus, and Coprinus atramentarius,
are also available, together with Fistulina hepatica, and Morchella
esculenta. In some districts, when mushrooms are scarce, it is stated that
almost any species that will yield a dark juice is without scruple mixed with
the common mushroom, and it should seem without any bad consequence
except the deterioration of the ketchup.[V] There is an extensive
manufacture of ketchup conducted at Lubbenham, near Market
Harborough, but the great difficulty appears to be the prevention of
decomposition. Messrs. Perkins receive tons of mushrooms from every part
of the kingdom, and they find, even in the same species, an immense
difference in the quality and quantity of the produce. The price of
mushrooms varies greatly with the season, ranging between one penny and
sixpence per pound. Messrs. Perkins are very careful in their selection, but
little discrimination is used by country manufacturers on a small scale, who
use such doubtful species as Agaricus lacrymabundus, with Agaricus
spadiceus, and a host of allied species, which they characterize as
nonpareils and champignons. In the eastern counties Agaricus arvensis has
the preference for ketchup.
The generic distinctions between the genuine Agarics and some of the
allied genera can hardly be appreciated by the non-botanical reader, but we
have nevertheless preferred grouping the edible species together in a
somewhat botanical order; and, pursuing this plan, the next species will be

those of Coprinus, in which the gills are deliquescent after the plant has
arrived at maturity. The maned mushroom (Coprinus comatus, Fr.)[W] is the
best of edible species in this group. It is very common here by roadsides
and other places, and whilst still young and cylindrical, and the gills still
whitish or with a roseate tint, it is highly to be commended. Similar, but
perhaps somewhat inferior, is Coprinus atramentarius, Fr.,[X] equally
common about old stumps and on the naked soil. Both species are also
found and eaten in the United States.
In Cortinarius, the veil is composed of arachnoid threads, and the spores
are rusty. The number of edible species are few. Foremost is the really
handsome Cortinarius violaeus, Fr.,[Y] often nearly four inches in diameter,
and of a beautiful violet colour; and the smaller Cortinarius castaneus, Fr.,
[Z] scarcely exceeding an inch in diameter, both being found in woods, and
common alike to Britain and the United States. Cortinarius cinnamomeus,
Fr., is also a lover of woods, and in northern latitudes is found inhabiting
them everywhere. It has a cinnamon-coloured pileus, with yellowish flesh,
and its odour and flavour is said to partake of the same spice. In Germany it
is held in high esteem. Cortinarius emodensis, B., is eaten in Northern
India.
The small genus Lepista of Smith, (which, however, is not adopted by
Fries in his now edition of the “Epicrisis”) includes one esculent species in
Lepista personata, the Agaricus personatus of Fries.[a] It is by no means
uncommon in Northern Europe or America, frequently growing in large
rings; the pileus is pallid, and the stem stained with lilac. Formerly it was
said to be sold in Covent Garden Market under the name of “blewits,” but
we have failed to see or hear of it during many years in London.
Small fungi of ivory-whiteness are very common amongst grass on lawns
in autumn. These are chiefly Hygrophorus virgineus, Fr.,[b] and although
not much exceeding an inch in diameter, with a short stem, and wide
decurrent gills, they are so plentiful in season that quantity soon
compensates for the small size. Except that it is occasionally eaten in
France, it does not enjoy much reputation abroad. A larger species, varying
from buff to orange, Hygrophorus pratensis, Fr.,[c] is scarcely less common
in open pastures. This is very gregarious in habit, often growing in tufts, or
portions of rings. The pileus is fleshy in the centre, and the gills thick and
decurrent. In France, Germany, Bohemia, and Denmark, it is included with

esculent species. In addition may be mentioned Hygrophorus eburneus, Fr.,
another white species, as also Hygrophorus niveus, Fr., which grows in
mossy pastures. Paxillus involutus, Fr.,[d] though very common in Europe,
is not eaten, yet it is included by Dr. Curtis with the esculent species of the
United States.
The milky agarics, belonging to the genus Lactarius, are distinguished by
the milky juice which is exuded when they are wounded. The spores are
more or less globose, and rough or echinulate, at least in many species. The
most notable esculent is Lactarius deliciosus, Fr.,[e] in which the milk is at
first saffron-red, and afterwards greenish, the plant assuming a lurid
greenish hue wherever bruised or broken. Universal commendation seems
to fall upon this species, writers vying with each other to say the best in its
praise, and mycophagists everywhere endorsing the assumption of its name,
declaring it to be delicious. It is found in the markets of Paris, Berlin,
Prague, and Vienna, as we are informed, and in Sweden, Denmark,
Switzerland, Russia, Belgium; in fact, in nearly all countries in Europe it is
esteemed.
Another esculent species, Lactarius volemum, Fr.,[f] has white milk,
which is mild to the taste, whilst in deleterious species with white milk it is
pungent and acrid. This species has been celebrated from early times, and is
said to resemble lamb’s kidney.
Lactarius piperatus, Fr., is classed in England with dangerous, sometimes
poisonous species, whereas the late Dr. Curtis, of North Carolina, has
distinctly informed us that it is cooked and eaten in the United States, and
that he has partaken of it. He includes Lactarius insulsus, Fr., and Lactarius
subdulcis, Fr.,[g] amongst esculent species; both are also found in this
country, but not reputed as edible; and Lactarius angustissimus, Lasch,
which is not British. Species of Lactarius seem to be eaten almost
indiscriminately in Russia when preserved in vinegar and salt, in which
condition they form an important item in the kinds of food allowed in their
long fasts, some Boleti in the dried state entering into the same category.
The species of Russula in many respects resemble Lactarii without milk.
Some of them are dangerous, and others esculent. Amongst the latter may
be enumerated Russula heterophylla, Fr., which is very common in woods.
Vittadini pronounces it unsurpassed for fineness of flavour by even the

notable Amanita cæsarea.[h] Roques gives also an account in its favour as
consumed in France. Both these authors give favourable accounts of
Russula virescens, P.,[i] which the peasants about Milan are in the habit of
putting over wood embers to toast, and eating afterwards with a little salt.
Unfortunately it is by no means common in England. A third species of
Russula, with buff-yellow gills, is Russula alutacea, Fr., which is by no
means to be despised, notwithstanding that Dr. Badham has placed it
amongst species to be avoided. Three or four others have also the merit of
being harmless, and these recorded as esculent by some one or more
mycological authors: Russula lactea, Fr., a white species, found also in the
United States; Russula lepida, Fr., a roseate species, found also in lower
Carolina, U.S.; and another reddish species, Russula vesca, Fr., as well as
Russula decolorans, Fr. Whilst writing of this genus, we may observe, by
way of caution, that it includes also one very noxious red species, Russula
emetica, Fr., with white gills, with which some of the foregoing might be
confounded by inexperienced persons.
The chantarelle Cantharellus cibarius, Fr., has a most charming and
enticing appearance and odour. In colour, it is of a bright golden yellow, and
its smell has been compared to that of ripe apricots. It is almost universally
eaten in all countries where it is found, England excepted, where it is only
to be met with at the “Freemason’s Tavern” on state occasions, and at the
tables of pertinacious mycophagists.[j] Trattinnick says: “Not only this same
fungus never did any one harm, but might even restore the dead.”[k]
The fairy-ring champignon Marasmius oreades, Fr., though small, is
plentiful, and one of the most delicious of edible fungi. It grows in exposed
pastures, forming rings, or parts of rings. This champignon possesses the
advantage of drying readily, and preserving its aroma for a long time. We
have often regretted that no persistent attempts and experiments have been
made with the view of cultivating this excellent and useful species.
Marasmius scorodonius, Fr.,[l] a small, strong-scented, and in all respects
inferior species, found on heaths and dry pastures, extending even to the
United States, is consumed in Germany, Austria, and other continental
countries, where, perhaps its garlic odour has been one of its
recommendations as an ingredient in sauces. In this enumeration we have
not exhausted all the gill-bearing species which might be eaten, having

included only those which have some reputation as esculents, and of these
more particularly those found in Great Britain and the United States.
Amongst the Polyporei, in which the gill plates are represented by pores
or tubes, fewer esculent species are to be met with than in the Agaricini,
and the majority of these belong to the genus Boletus. Whilst in Vienna and
Hanover, we were rather surprised to find Boletus edulis, Fr., cut into thin
slices and dried, exposed for sale in almost every shop where meal, peas,
and other farinaceous edibles were sold. This species is common enough in
England, but as a rule it does not seem to please the English palate, whereas
on the continent no fungus is more commonly eaten. This is believed to be
the suillus eaten by the ancient Romans,[m] who obtained it from Bithynia.
The modern Italians dry them on strings for winter use, and in Hungary a
soup is made from them when fresh. A more excellent species, according to
our judgment, is Boletus æstivalis, Fr.,[n] which appears in early summer,
and has a peculiar nutty flavour when raw, reminding one more of a fresh
mushroom. Boletus scaber, Fr.,[o] is also common in Britain, as well as the
continent, but does not enjoy so good a reputation as B. edulis. Krombholz
says that Boletus bovinus, Fr., a gregarious species, found on heaths and in
fir woods, is much sought after abroad as a dish, and is good when dried.
Boletus castaneus, Fr.,[p] is a small species with a mild, pleasant taste when
raw, and very good when properly cooked. It is not uncommonly eaten on
the continent. Boletus chrysenteron, Fr.,[q] and Boletus subtomentosus, Fr.,
are said to be very poor eating, and some authors have considered them
injurious; but Mr. W. G. Smith states that he has on more than one occasion
eaten the former, and Trattinnick states that the latter is eaten in Germany.
The late Mr. Salter informed us that, when employed on the geological
staff, he at one time lived almost entirely on different species of Boleti,
without using much discrimination. Sir W. C. Trevelyan also informs us that
he has eaten Boletus luridus without any unpleasant consequences, but we
confess that we should be sorry to repeat the experiment. Dr. Badham
remarks that he has eaten Boletus Grevillei, B., Boletus flavus, With., and
Boletus granulatus, L., the latter being recognized also as edible abroad. Dr.
Curtis experimented, in the United States, on Boletus collinitus, and
although he professes not to be particularly fond of the Boleti, he
recognizes it as esculent, and adds that it had been pronounced delicious by
some to whom he had sent it. He also enumerates as edible Boletus luteus,
Fr., Boletus elegans, Fr., Boletus flavidus, Fr., Boletus versipellis, Fr.,

Boletus leucomelas, Tr., and Boletus ovinus, Sch. Two Italian species of
Polyporus must not be forgotten. These are Polyporus tuberaster, Pers.,
which is procured by watering the pietra funghaia, or fungus stone, a kind
of tufa, in which the mycelium is embedded. It is confined to Naples. The
other species is Polyporus corylinus, Mauri., procured artificially in Rome
from charred stumps of the cob-nut tree.[r]
Of true Polyporus, only two or three species have been regarded
favourably as esculents. These are—Polyporus intybaceus, Fr., which is of
very large size, sometimes attaining as much as forty pounds; Polyporus
giganteus, Fr., also very large, and leathery when old. Both these species
are natives of Britain. Only young and juicy specimens must be selected for
cooking. Polyporus umbellatus, Fr., is stated by Fries to be esculent, but it
is not found in Britain. Polyporus squamosus, Fr., has been also included;
but Mrs. Hussey thinks that one might as well think of eating saddle-flaps.
None of these receive very much commendation. Dr. Curtis enumerates,
amongst North American species, the Polyporus cristatus, Fr., Polyporus
poripes, Fr., which, when raw, tastes like the best chestnuts or filberts, but is
rather too dry when cooked. Polyporus Berkeleii, Fr., is intensely pungent
when raw, but when young, and before the pores are visible, it may be eaten
with impunity, all its pungency being dissipated by cooking. Polyporus
confluens, Fr., he considers superior, and, in fact, quite a favourite.
Polyporus sulfureus, Fr., which is not eaten in Europe, he considers just
tolerably safe, but not to be coveted. It is by no means to be recommended
to persons with weak stomachs. In his catalogue, Dr. Curtis enumerates one
hundred and eleven species of edible fungi found in Carolina.[s]
With Fistulina hepatica, Fr., it is different; for here we encounter a
fleshy, juicy fungus, resembling beefsteak a little in appearance, and so
much more in its uses, that the name of “beefsteak fungus” has been given
to it. Some authors are rapturous in their praise of Fistulina. It sometimes
attains a very large size, Dr. Badham quoting[t] one found by himself nearly
five feet in circumference, and weighing eight pounds; whilst another found
by Mr. Graves weighed nearly thirty pounds. In Vienna it is sliced and eaten
with salad, like beetroot, which it then much resembles. On the continent it
is everywhere included amongst the best of edible species.
The Hydnei, instead of pores or tubes, are characterized by spines or
warts, over which the fructifying surface is expanded. The most common is

Hydnum repandum, Fr., found in woods and woody places in England, and
on the continent, extending into the United States. When raw, it is peppery
to the taste, but when cooked is much esteemed. From its drier nature, it can
readily be dried for winter use. Less common in England is Hydnum
imbricatum, Fr., although not so uncommon on the continent. It is eaten in
Germany, Austria, Switzerland, France, and elsewhere. Hydnum lævigatum,
Swartz, is eaten in Alpine districts.[u] Of the branched species, Hydnum
coralloides, Scop.,[v] and Hydnum Caput Medusæ, Bull,[w] are esculent, but
very rare in England. The latter is not uncommon in Austria and Italy, the
former in Germany, Switzerland, and France. Hydnum erinaceum, Bull, is
eaten in Germany[x] and France.
The Clavarioid fungi are mostly small, but of these the majority of the
white-spored are edible. Clavaria rugosa, Bull, is a common British
species, as also is Clavaria coralloides, L., the former being found also in
the United States. Clavaria fastigiata, D. C., is not uncommon; but
Clavaria amethystina, Bull, a beautiful violet species, is rare. In France and
Italy, Clavaria cinerea, Bull, is classed with esculents; and it is not
uncommon in Britain. Clavaria botrytis, P., and Clavaria aurea, Schæff.,
are large and beautiful species, but rare with us; they extend also into the
United States. Others might be named (Dr. Curtis enumerates thirteen
species eaten in Carolina), which are certainly wholesome, but they are of
little importance as edible species. Sparassis crispa, Fr., is, on the contrary,
very large, resembling in size,[y] and somewhat in appearance, a
cauliflower; it has of late years been found several times in this country. In
Austria it is fricasseed with butter and herbs.
Of the true Tremellæ, none merit insertion here. The curious Jew’s ear
(Hirneola auricula-Judæ, Fr.), with one or two other species of Hirneola,
are collected in great quantities in Tahiti, and shipped in a dried state to
China, where they are used for soup. Some of these find their way to
Singapore.
The false truffles (Hypogæi) are of doubtful value, one species
(Melanogaster variegatus, Tul.) having formerly been sold in the markets
of Bath as a substitute for the genuine truffle.[z] Neither amongst the
Phalloidei do we meet with species of any economic value. The gelatinous
volva of a species of Ileodictyon is eaten by the New Zealanders, to whom
it is known as thunder dirt; whilst that of Phallus Mokusin is applied to a

like purpose in China;[AA] but these examples would not lead us to
recommend a similar use for Phallus impudicus, Fr., in Britain, or induce us
to prove the assertion of a Scotch friend that the porous stem is very good
eating.
One species of puff-ball, Lycoperdon giganteum, Fr,[AB] has many
staunch advocates, and whilst young and cream-like, it is, when well
manipulated, an excellent addition to the breakfast-table. A decided
advantage is possessed by this species, since one specimen is often found
large enough to satisfy the appetites of ten or twelve persons. Other species
of Lycoperdon have been eaten when young, and we have been assured by
those who have made the experiment, that they are scarcely inferior to their
larger congener. Bovista nigrescens, Fr., and Bovista plumbea, Fr., are also
eaten in the United States. More than one species of Lycoperdon and
Bovista appear in the bazaars of India, as at Secunderabad and Rangoon;
while the white ant-hills, together with an excellent Agaric, produce one or
more species of Podaxon which are esculent when young. A species of
Scleroderma which grows abundantly in sandy districts, is substituted for
truffles in Perigord pies, of which, however, it does not possess any of the
aroma.

Fig. 43.—Morchella gigaspora, from Kashmir.

Passing over the rest of the sporiferous fungi, we find amongst the
Ascomycetous group several that are highly esteemed. Amongst these may
first be named the species of morel, which are regarded as delicacies
wherever they are found. Morchella esculenta, Pers., is the most common
species, but we have also Morchella semilibera, D. C., and the much larger
Morchella crassipes, Pers. Probably all the species of Morchella are
esculent, and we know that many besides the above are eaten in Europe and
other places; Morchella deliciosa, Fr., in Java; Morchella bohemica,
Kromb., in Bohemia; Morchella gigaspora, Cooke, and Morchella
deliciosa, Fr., in Kashmere.[AC] Morchella rimosipes, D. C., occurs in
France and Bohemia; Morchella Caroliniana, Bosc., in the Southern United
States of America. W. G. Smith records the occurrence in Britain of
specimens of Morchella crassipes, P., ten inches in height, and one
specimen was eleven inches high, with a diameter of seven and a half
inches.[AD]
Similar in uses, though differing in appearance, are the species of
Helvella, of which several are edible. In both these genera, the individuals
can be dried so readily that they are the more valuable on that account, as

they can be used for flavouring in winter when fresh specimens of any kind
of fungus are difficult to procure. The most common English species is
Helvella crispa, Fr., but Helvella lacunosa, Fr., is declared to be equally
good, though not so large and somewhat rare. Helvella infula, Fr., is also a
large species, but is not British, although it extends to North America, as
also does Helvella sulcata, Afz. Intermediate between the morel and
Helvella is the species which was formerly included with the latter, but now
known as Gyromitra esculenta, Fr.[AE] It is rarely found in Great Britain,
but is more common on the continent, where it is held in esteem. A curious
stipitate fungus, with a pileus like a hood, called Verpa digitaliformis, Pers.,
[AF] is uncommon in England, but Vittadini states that it is sold in the Italian
markets, although only to be recommended when no other esculent fungus
offers, which is sometimes the case in spring.[AG]
Two or three species of Peziza have the reputation of being esculent, but
they are of very little value; one of these is Peziza acetabulum, L., another
is Peziza cochleata, Huds., and a third is Peziza venosa, Pers.[AH] The latter
has the most decided nitrous odour, and also fungoid flavour, whilst the
former seem to have but little to recommend them; we have seen whole
baskets full of Peziza cochleata gathered in Northamptonshire as a
substitute for morels.
A very interesting genus of edible fungi, growing on evergreen beech
trees in South America, has been named Cyttaria. One of these, Cyttaria
Darwinii, B., occurs in Terra del Fuego, where it was found by Mr. C.
Darwin[AI] growing in vast numbers, and forming a very essential article of
food for the natives. Another is Cyttaria Berteroi, B., also seen by Mr.
Darwin in Chili, and eaten occasionally, but apparently not so good as the
preceding.[AJ] Another species is Cyttaria Gunnii, B., which abounds in
Tasmania, and is held in repute amongst the settlers for its esculent
properties.[AK]

Fig. 44.—Cyttaria Gunnii, B.

It remains for us only to note the subterranean fungi, of which the truffle
is the type, to complete our enumeration of esculent species. The truffle
which is consumed in England is Tuber æstivum, Vitt.; but in France the
more highly-flavoured Tuber melanospermum, Vitt.,[AL] and also Tuber
magnatum, Pico, with some other species. In Italy they are very common,
whilst some are found in Algeria. One species at least is recorded in the
North-west of India, but in Northern Europe and North America they appear
to be rare, and Terfezia Leonis is used as an esculent in Damascus. A large
species of Mylitta, sometimes several inches in diameter, occurs plentifully
in some parts of Australia. Although often included with fungi, the curious
production known under the name of Pachyma cocos, Fr., is not a fungus,
as proved by the examinations made by the Rev. M. J. Berkeley. It is eaten
under the name of “Tuckahoe” in the United States, and as it consists
almost entirely of pectic acid, it is sometimes used in the manufacture of
jelly.
In the Neilgherries (S. India), a substance is occasionally found which is
allied to the native bread of southern latitudes. It is found at an elevation of
5,000 feet. The natives call it “a little man’s bread,” in allusion to the
tradition that the Neilgherries were once peopled by a race of dwarfs.[AM]
At first it was supposed that these were the bulbs of some orchid, but later
another view was held of their character. Mr. Scott, who examined the
specimens sent down to him, remarks that, instead of being the product of
orchids, it is that of an underground fungus of the genus Mylitta. It indeed
seems, he says, very closely allied to, if really distinct from, the so-called
native bread of Tasmania.[AN]

Of the fungi employed in medicine, the first place must be assigned to
ergot, which is the sclerotioid condition of a species of Claviceps. It occurs
not only on rye but on wheat, and many of the wild grasses. On account of
its active principle, this fungus still holds its place in the Materia Medica.
Others which formerly had a reputation are now discarded, as, for instance,
the species of Elaphomyces; and Polyporus officinalis, Fr., which has been
partly superseded as a styptic by other substances, was formerly employed
as a purgative. The ripe spongy capillitium of the great puff-ball
Lycoperdon giganteum, Fr., has been used for similar purposes, and also
recommended as an anodyne; indeed formidable surgical operations have
been performed under its influence, and it is frequently used as a narcotic in
the taking of honey. Langsdorf gives a curious account of its employment as
a narcotic; and in a recent work on Kamtschatka it is said to obtain a very
high price in that country. Dr. Porter Smith writes of its employment
medicinally by the Chinese, but from his own specimens it is clearly a
species of Polysaccum, which he has mistaken for Lycoperdon. In China
several species are supposed to possess great virtue, notably the Torrubia
sinensis, Tul.,[AO] which is developed on dead caterpillars; as it is, however,
recommended to administer it as a stuffing to roast duck, we may be
sceptical as to its own sanitary qualities. Geaster hygrometricus, Fr., we
have also detected amongst Chinese drugs, as also a species of Polysaccum,
and the small hard Mylitta lapidescens, Horn. In India, a large but imperfect
fungus, named provisionally Sclerotium stipitatum, Curr., found in nests of
the white ant, is supposed to possess great medicinal virtues.[AP] A species
of Polyporus (P. anthelminticus, B.), which grows at the root of old
bamboos, is employed in Burmah as an anthelmintic.[AQ] In former times
the Jew’s ear (Hirneola auricula Judæ, Fr.) was supposed to possess great
virtues, which are now discredited. Yeast is still included amongst
pharmaceutical substances, but could doubtless be very well dispensed
with. Truffles are no longer regarded as aphrodisiacs.
For other uses, we can only allude to amadou, or German tinder, which is
prepared in Northern Europe from Polyporus fomentarius, Fr., cut in slices,
dried, and beaten until it is soft. This substance, besides being used as
tinder, is made into warm caps, chest protectors, and other articles. This
same, or an allied species of Polyporus, probably P. igniarius, Fr., is dried
and pounded as an ingredient in snuff by the Ostyacks on the Obi. In
Bohemia some of the large Polyporei, such as P. igniarius and P.

fomentarius, have the pores and part of the inner substance removed, and
then the pileus is fastened in an inverted position to the wall, by the part
where originally it adhered to the wood. The cavity is then filled with
mould, and the fungus is used, with good effect, instead of flower-pots, for
the cultivation of such creeping plants as require but little moisture.[AR]
The barren mycelioid condition of Penicillium crustaceum, Fr., is
employed in country districts for the domestic manufacture of vinegar from
saccharine liquor, under the name of the “vinegar plant.” It is stated that
Polysaccum crassipes, D. C.,[AS] is employed in the South of Europe to
produce a yellow dye; whilst recently Polyporus sulfureus, Fr., has been
recommended for a similar purpose. Agaricus muscarius, Fr., the fly-agaric,
known to be an active poison, is used in decoction in some parts of Europe
for the destruction of flies and bugs. Probably Helotium æruginosum, Fr.,
[AT] deserves mention here, because it stains the wood on which it grows, by
means of its diffuse mycelium, of a beautiful green tint, and the wood thus
stained is employed for its colour in the manufacture of Tonbridge ware.
This completes the list, certainly of the most important, of the fungi
which are of any direct use to humanity as food, medicine, or in the arts. As
compared with lichens, the advantage is certainly in favour of fungi; and
even when compared with algæ, the balance appears in their favour. In fact,
it may be questioned whether, after all, fungi do not present a larger
proportion of really useful species than any other of the cryptogams; and
without any desire to disparage the elegance of ferns, the delicacy of
mosses, the brilliancy of some algæ, or the interest which attaches to
lichens, it may be claimed for fungi that in real utility (not uncombined with
injuries as real) they stand at the head of the cryptogams, and in closest
alliance with the flowering plants.
[A]
Badham, Dr. C. D., “A Treatise on the Esculent Funguses of England,” 1st edition
(1847), p. 81, pl. 4; 2nd edition, edited by F. Currey, M.A. (1863), p. 94, pl. 4; Cooke,
M. C., “A Plain and Easy Account of British Fungi,” 1st edition (1862), p. 44.
[B]
Mr. Worthington Smith has published, on two sheets, coloured figures of the most
common esculent and poisonous fungi (London, Hardwicke), which will be found
more useful than mere description in the discrimination of the species.
[C]

Roques, J., “Hist. des Champignons Comestibles et Vénéneux,” Paris (1832), p. 130.
[D]
Lenz, Dr. H. 0., “Die Nützlichen und Schädlichen Schwämme,” Gotha (1831), p. 32,
pl. 2.
[E]
Bull, H. G., in “Transactions of Woolhope Club” (1869). Fries admits them as distinct
species in the new edition of his “Epicrisis.”
[F]
Hussey’s “Illustrations of Mycology,” ser. i. pl. 79.
[G]
Sowerby’s “British Fungi,” pl. 244.
[H]
Favre-Guillarmod, “Les Champignons Comestibles du Canton de Neuchatel” (1861),
p. 27.
[I]
Sowerby, “English Fungi,” pl. 122; Smith, in “Seemann’s Journ. Bot.” (1866), t. 46, f.
45.
[J]
Klotsch, “Flora Borussica,” t. 374; Smith, in “Seem. Journ. Bot.” (1869), t. 95, f. 1–4.
[K]
Krombholz, “Abbildungen der Schwämme,” pl. 41, f. 1–7.
[L]
Tratinnick, L., “Fungi Austriaci,” p. 47, pl. 4, f. 8.
[M]
Vittadini, “Fungi Mangerecci,” pl. 23.
[N]
Cooke, in “Journal of Botany,” vol. viii. p. 352.
[O]
Cooke, M. C., “A Plain and Easy Guide,” &c., p. 38, pl. 6, fig. 1.
[P]
Krombholz, “Schwämme,” t. 8. Vittadini, “Mang.” t. 1.
[Q]
Vittadini, “Mangerecci,” t. 9.

[R]
Berkeley, “Outlines,” pl. 3, fig. 5.
[S]
Saunders and Smith, “Mycological Illustr.” pl. 23.
[T]
Cooke, M. C., “Handbook of British Fungi,” vol. i. pl. 1, fig. 2.
[U]
“Gardener’s Chronicle” (1869), p. 1066.
[V]
Berkeley, “Outlines of British Fungology,” p. 64.
[W]
Cooke, “Easy Guide to British Fungi,” pl. 11.
[X]
Ibid., pl. 12.
[Y]
Hussey, “Mycol. Illust.” pl. 12.
[Z]
Bulliard, “Champ.” t. 268.
[a]
Cooke, “Easy Guide,” pl. 4, fig. 1; Hussey, “Illust.” vol. ii. pl. 40.
[b]
Greville, “Scot. Crypt. Flora,” t. 166.
[c]
Ibid., t. 91.
[d]
Sowerby, “Fungi,” pl. 56; Schæffer, “Icones Bav.” t. 72.
[e]
Trattinnick, L., “Die Essbaren Schwämme” (1809), p. 82, pl. M; Barla, J. B.,
“Champignons de la Nice” (1859), p. 34, pl. 19.
[f]
Smith, “Edible Mushrooms,” fig. 26.

[g]
Barla, “Champ. Nice,” t. 20, f. 4–10.
[h]
Vittadini, C., “Funghi Mangerecci” (1835), p. 209; Barla, “Champ. Nice,” pl. i.
[i]
Vittadini, C., “Funghi Mangerecci,” p. 245; Roques, “Champ. Comest.” p. 86.
[j]
Badham, Dr., “Esculent Funguses of Britain,” 2nd ed. p. 110; Hussey, “Illust. Brit.
Mycol.” 1st ser. pl. 4; Barla, “Champ.” pl. 28, f. 7–15.
[k]
Trattinnick, L., “Essbaren Schwämme,” p. 98.
[l]
Lenz, “Die Nützlichen und Schädlichen Schwämme,” p. 49.
[m]
Badham, “Esculent Funguses of Great Britain,” 2 ed. p. 91.
[n]
Hussey, “Myc. Illus.” ii. pl. 25; Paulet, “Champ.” t. 170.
[o]
Barla, J. B., “Champ. de la Nice,” p. 71, pl. 35, f. 1–5.
[p]
Hussey, “Illustr.” ii. t. 17; Barla, “Champ. Nice,” t. 32, f. 11–15.
[q]
Hussey, “Illustr.” i. t. 5; Krombholz, “Schwämme,” t. 76.
[r]
Badham’s “Esculent Funguses,” 1st ed. pp. 116 and 120.
[s]
Catalogue of Plants of Carolina, U.S.
[t]
Badham, Dr., “Esculent Funguses,” 2nd ed. p. 128; Hussey, “Illustrations,” 1st ser. pl.
65; Berkeley, in “Gard. Chron.” (1861), p. 121; Bull, in “Trans. Woolhope Club”
(1869).
[u]

Barla, “Champ. Nice,” p. 79, pl. 38, f. 5, 6.
[v]
Roques, I. c. p. 48.
[w]
Lenz, p. 93; Roques, I. c. p. 47, pl. 2, fig. 5.
[x]
Lenz, H. O., “Die Nützlichen und Schädlichen Schwämme,” p. 93.
[y]
Berkeley, M. J., in “Intellectual Observer,” No. 25, pl. 1.
[z]
Berkeley, M. J., “Outlines of British Fungology,” p. 293.
[AA]
Berkeley, M. J., “Introduction to Crypt. Bot.” p. 347.
[AB]
Cooke, M. C., “A Plain and Easy Guide,” &c., p. 96.
[AC]
Cooke, M. C., “On Kashmir Morels,” in “Trans. Bot. Soc. Edin.” vol. x. p. 439, with
figs.
[AD]
Smith, “Journ. Bot.” vol. ix. p. 214.
[AE]
Cooke, “Handbook,” fig. 322.
[AF]
Cooke, “Handbook,” fig. 324.
[AG]
Vittadini, C., “Funghi Mangerecci,” p. 117.
[AH]
Greville, “Sc. Crypt. Fl.” pl. 156.
[AI]
Berkeley, in “Linn Trans.” xix. p. 37; Cooke, in “Technologist” (1864), p. 387.
[AJ]
Berkeley, M. J., in “Linn. Trans.” xix. p. 37.

[AK]
Berkeley, M. J., in “Hooker, Flora Antarctica,” p. 147; in “Hooker’s Journ. Bot.”
(1848), 576, t. 20, 21.
[AL]
Vittadini, C., “Monographia Tuberacearum” (1831), pp. 36, &c.
[AM]
“Proceedings Agri. Hort. Soc. India” (Dec. 1871), p. lxxix.
[AN]
Ibid. (June, 1872), p. xxiii.
[AO]
Lindley, “Vegetable Kingdom,” fig. xxiv.
[AP]
Currey, F., in “Linn. Trans.” vol. xxiii. p. 93.
[AQ]
“Pharmacopœia of India,” p. 258.
[AR]
“Gard. Chron.” (1862), p. 21.
[AS]
Barla, “Champ. de la Nice,” p. 126, pl. 47, fig. 11.
[AT]
Greville, “Scott. Crypt. Flora,” pl. 241.

V.
NOTABLE PHENOMENA.

There are no phenomena associated with fungi that are of greater interest
than those which relate to luminosity. The fact that fungi under some
conditions are luminous has long been known, since schoolboys in our
juvenile days were in the habit of secreting fragments of rotten wood
penetrated by mycelium, in order to exhibit their luminous properties in the
dark, and thus astonish their more ignorant or incredulous fellows
Rumphius noted its appearance in Amboyna, and Fries, in his Observations,
gives the name of Thelephora phosphorea to a species of Corticium now
known as Corticium cæruleum, on account of its phosphorescence under
certain conditions. The same species is the Auricularia phosphorea of
Sowerby, but he makes no note of its phosphorescence. Luminosity in fungi
“has been observed in various parts of the world, and where the species has
been fully developed it has been generally a species of Agaricus which has
yielded the phenomenon.”[A] One of the best-known species is the Agaricus
olearius of the South of Europe, which was examined by Tulasne with
especial view to its luminosity.[B] In his introductory remarks, he says that
four species only of Agaricus that are luminous appear at present to be
known. One of them, A. olearius, D. C., is indigenous to Central Europe;
another, A. igneus, Rumph., comes from Amboyna; the third, A.
noctileucus, Lév., has been discovered at Manilla by Gaudichaud, in 1836;
the last, A. Gardneri, Berk., is produced in the Brazilian province of Goyaz,
upon dead leaves. As to the Dematium violaceum, Pers., the Himantia
candida, Pers., cited once by Link, and the Thelephora cærulea, D. C.
(Corticium cæruleum, Fr.), Tulasne is of opinion that their phosphorescent
properties are still problematical; at least no recent observation confirms
them.
The phosphorescence of A. olearius, D. C., appears to have been first
made known by De Candolle, but it seems that he was in error in stating

that these phosphorescent properties manifest themselves only at the time of
its decomposition. Fries, describing the Cladosporium umbrinum, which
lives upon the Agaric of the olive-tree, expressed the opinion that the Agaric
only owes its phosphorescence to the presence of the mould. This, however,
Tulasne denies, for he writes, “I have had the opportunity of observing that
the Agaric of the olive is really phosphorescent of itself, and that it is not
indebted to any foreign production for the light it emits.” Like Delile, he
considers that the fungus is only phosphorescent up to the time when it
ceases to grow; thus the light which it projects, one might say, is a
manifestation of its vegetation.
“It is an important fact,” writes Tulasne, “which I can confirm, and which
it is important to insist upon, that the phosphorescence is not exclusively
confined to the hymenial surface. Numerous observations made by me
prove that the whole of the substance of the fungus participates very
frequently, if not always, in the faculty of shining in the dark. Among the
first Agarics which I examined, I found many, the stipe of which shed here
and there a light as brilliant as the hymenium, and led me to think that it
was due to the spores which had fallen on the surface of the stipe.
Therefore, being in the dark, I scraped with my scalpel the luminous parts
of the stipe, but it did not sensibly diminish their brightness; then I split the
stipe, bruised it, divided it into small fragments, and I found that the whole
of this mass, even in its deepest parts, enjoyed, in a similar degree to its
superficies, the property of light. I found, besides, a phosphorescence quite
as brilliant in all the cap, for, having split it vertically in the form of plates, I
found that the trama, when bruised, threw out a light equal to that of their
fructiferous surfaces, and there is really only the superior surface of the
pileus, or its cuticle, which I have never seen luminous.
“As I have said, the Agaric of the olive-tree, which is itself very yellow,
reflects a strong brilliant light, and remains endowed with this remarkable
faculty whilst it grows, or, at least, while it appears to preserve an active
life, and remains fresh. The phosphorescence is at first, and more ordinarily,
recognizable at the surface of the hymenium. I have seen a great number of
young fungi which were very phosphorescent in the gills, but not in any
other part. In another case, and amongst more aged fungi, the hymenium of
which had ceased to give light, the stipe, on the contrary, threw out a
brilliant glare. Habitually, the phosphorescence is distributed in an unequal

manner upon the stipe, and the same upon the gills. Although the stipe is
luminous at its surface, it is not always necessarily so in its interior
substance, if one bruises it, but this substance frequently becomes
phosphorescent after contact with the air. Thus, I had irregularly split and
slit a large stipe in its length, and I found the whole flesh obscure, whilst on
the exterior were some luminous places. I roughly joined the lacerated
parts, and the following evening, on observing them anew, I found them all
flashing a bright light. At another time, I had with a scalpel split vertically
many fungi in order to hasten their dessication; the evening of the same day,
the surface of all these cuts was phosphorescent, but in many of these
pieces of fungi the luminosity was limited to the cut surface which
remained exposed to the air; the flesh beneath was unchanged.
“I have seen a stipe opened and lacerated irregularly, the whole of the
flesh of which remained phosphorescent during three consecutive evenings,
but the brightness diminished in intensity from the exterior to the interior,
so that on the third day it did not issue from the inner part of the stipe. The
phosphorescence of the gills is in no way modified at first by immersing the
fungus in water; when they have been immersed they are as bright as in the
air, but the fungi which I left immersed until the next evening lost all their
phosphorescence, and communicated to the water an already sensible
yellow tint; alcohol put upon the phosphorescent gills did not at once
completely obliterate the light, but visibly enfeebled it. As to the spores,
which are white, I have found many times very dense coats of them thrown
down on porcelain plates, but I have never seen them phosphorescent.
“As to the observation made by Delile that the Agaric of the olive does
not shine during the day when placed in total darkness, I think that it could
not have been repeated. From what I have said of the phosphorescence of A.
olearius, one naturally concludes that there does not exist any necessary
relation between this phenomenon and the fructification of the fungus; the
luminous brightness of the hymenium shows, says Delile, ‘the greater
activity of the reproductive organs,’ but it is not in consequence of its
reproductive functions, which may be judged only as an accessory
phenomenon, the cause of which is independent of, and more general than
these functions, since all the parts of the fungus, its entire substance, throws
forth at one time, or at successive times, light. From these experiments
Tulasne infers that the same agents, oxygen, water, and warmth, are

perfectly necessary to the production of phosphorescence as much in living
organized beings as in those which have ceased to live. In either case, the
luminous phenomena accompany a chemical reaction which consists
principally in a combination of the organized matter with the oxygen of the
air; that is to say, in its combustion, and in the discharge of carbonic acid
which thus shows itself.”
We have quoted at considerable length from these observations of
Tulasne on the Agaric of the olive, as they serve very much to illustrate
similar manifestations in other species, which doubtless resemble each
other in their main features.
Mr. Gardner has graphically described his first acquaintance in Brazil
with the phosphorescent species which now bears his name. It was
encountered on a dark night of December, while passing through the streets
of Villa de Natividate. Some boys were amusing themselves with some
luminous object, which at first he supposed to be a kind of large fire-fly, but
on making inquiry he found it to be a beautiful phosphorescent Agaric,
which he was told grew abundantly in the neighbourhood on the decaying
fronds of a dwarf palm. The whole plant gives out at night a bright light
somewhat similar to that emitted by the larger fire-flies, having a pale
greenish hue. From this circumstance, and from growing on a palm, it was
called by the inhabitants “flor de coco.”[C]
The number of recognized phosphorescent species of Agaricus is not
large, although two or three others may be enumerated in addition to those
cited by Tulasne. Of these, Agaricus lampas, and some others, are found in
Australia.[D] In addition to the Agaricus noctileucus, discovered by
Gaudichaud, and the Agaricus igneus of Rumphius, found in Amboyna, Dr.
Hooker speaks of the phenomenon as common in Sikkim, but he seems
never to have been able to ascertain with what species it was associated.
Dr. Cuthbert Collingwood has communicated some further information
relative to the luminosity of a species of Agaricus in Borneo (supposed to
be A. Gardneri), in which he says, “The night being dark, the fungi could be
very distinctly seen, though not at any great distance, shining with a soft
pale greenish light. Here and there spots of much more intense light were
visible, and these proved to be very young and minute specimens. The older
specimens may more properly be described as possessing a greenish

luminous glow, like the glow of the electric discharge, which, however, was
quite sufficient to define its shape, and, when closely examined, the chief
details of its form and appearance. The luminosity did not impart itself to
the hand, and did not appear to be affected by the separation from the root
on which it grew, at least not for some hours. I think it probable that the
mycelium of this fungus is also luminous, for, upon turning up the ground
in search of small luminous worms, minute spots of light were observed,
which could not be referred to any particular object or body when brought
to the light and examined, and were probably due to some minute portions
of its mycelium.”[E] The same writer also adds, “Mr. Hugh Low has assured
me that he saw the jungle all in a blaze of light (by which he could see to
read) as, some years ago, he was riding across the island by the jungle road;
and that this luminosity was produced by an Agaric.”
Similar experiences were detailed by Mr. James Drummond in a letter
from Swan River, in which two species of Agaric are concerned. They grew
on the stumps of trees, and had nothing remarkable in their appearance by
day, but by night emitted a most curious light, such as the writer never saw
described in any book. One species was found growing on the stump of a
Banksia in Western Australia. The stump was at the time surrounded by
water. It was on a dark night, when passing, that the curious light was first
observed. When the fungus was laid on a newspaper, it emitted by night a
phosphorescent light, enabling persons to read the words around it, and it
continued to do so for several nights with gradually decreasing intensity as
the plant dried up. In the other instance, which occurred some years after,
the author, during one of his botanical trips, was struck by the appearance of
a large Agaric, measuring sixteen inches in diameter, and weighing about
five pounds. This specimen was hung up to dry in the sitting-room, and on
passing through the apartment in the dark it was observed to give out the
same remarkable light. The luminous property continued, though gradually
diminishing, for four or five nights, when it ceased on the plant becoming
dry. “We called some of the natives,” he adds, “and showed them this
fungus when emitting light, and the poor creatures cried out ‘chinga,’ their
name for a spirit, and seemed much afraid of it.”[F]
Although the examples already cited are those of species of Agaric,
luminosity is not by any means wholly confined to that genus. Mr.
Worthington Smith has recorded his experiences of some specimens of the

common Polyporus annosus which were found on some timbers in the
Cardiff coal mines. He remarks that the colliers are well acquainted with
phosphorescent fungi, and the men state that sufficient light is given “to see
their hands by.” The specimens of Polyporus were so luminous that they
could be seen in the dark at a distance of twenty yards. He observes further,
that he has met with specimens of Polyporus sulfureus which were
phosphorescent. Some of the fungi found in mines, which emit light
familiar to the miners, belong to the incomplete genus Rhizomorpha, of
which Humboldt amongst others gives a glowing account. Tulasne has also
investigated this phenomenon in connection with the common Rhizomorpha
subterranea, Pers. This species extends underneath the soil in long strings,
in the neighbourhood of old tree stumps, those of the oak especially, which
are becoming rotten, and upon these it is fixed by one of its branches. These
are cylindrical, very flexible, branching, and clothed with a hard bark,
encrusting and fragile, at first smooth and brown, becoming later very
rough and black. The interior tissue, at first whitish, afterwards of a more or
less deep brown colour, is formed of extremely long parallel filaments from
.0035 to .015 mm. in diameter.
On the evening of the day when I received the specimens,[G] he writes,
the temperature being about 22° Cent., all the young branches brightened
with an uniform phosphoric light the whole of their length; it was the same
with the surface of some of the older branches, the greater number of which
were still brilliant in some parts, and only on their surface. I split and
lacerated many of these twigs, but their internal substance remained dull.
The next evening, on the contrary, this substance, having been exposed to
contact with the air, exhibited at its surface the same brightness as the bark
of the branches. I made this observation upon the old stalks as well as upon
the young ones. Prolonged friction of the luminous surfaces reduced the
brightness and dried them to a certain degree, but did not leave on the
fingers any phosphorescent matter. These parts continued with the same
luminous intensity after holding them in the mouth so as to moisten them
with saliva; plunged into water, held to the flame of a candle so that the heat
they acquired was very appreciable to the touch, they still emitted in the
dark a feeble light; it was the same after being held in water heated to 30°
C.; but putting them in water bearing a temperature of 55° C. extinguished
them entirely. They are equally extinguished if held in the mouth until they
catch the temperature; perhaps, still, it might be attributed less to the heat

which is communicated to them than to the deficiency of sufficient oxygen,
because I have seen some stalks, having become dull in the mouth, recover
after a few instants a little of their phosphorescence. A young stalk which
had been split lengthwise, and the internal substance of which was very
phosphorescent, could imbibe olive oil many times and yet continue for a
long time to give a feeble light. By preserving these Rhizomorphæ in an
adequate state of humidity, I have been able for many evenings to renew the
examination of their phosphorescence; the commencement of dessication,
long before they really perish, deprives them of the faculty of giving light.
Those which had been dried for more than a month, when plunged into
water, commenced to vegetate anew and send forth numerous branches in a
few days; but I could only discover phosphorescence at the surface of these
new formations, or very rarely in their immediate neighbourhood, the
mother stalks appearing to have lost by dessication their luminous
properties, and did not recover them on being recalled to life. These
observations prove that what Schmitz has written was not true, that all parts
of these fungi were seldom phosphorescent.
The luminous phenomenon in question is without doubt more
complicated than it appears, and the causes to which we attribute it are
certainly powerfully modified by the general character of the objects in
which they reside. Most of the German botanists give this explanation,
others suppose that it forms at first or during its continuance a special
matter, in which the luminous property resides; this matter, which is said to
be mucilaginous in the luminous wood, appears to be in the Rhizomorpha
only a kind of chemical combination between the membrane and some
gummy substance which they contain. Notwithstanding this opinion, I am
assured that all external mucous matter was completely absent from the
Agaricus olearius, and I neither discovered it upon the branches of
Rhizomorpha subterranea nor upon the dead leaves which I have seen
phosphorescent; in all these objects the luminous surfaces were nothing else
than their proper tissue.
It may be remarked here that the so-called species of Rhizomorpha are
imperfect fungi, being entirely devoid of fructification, consisting in fact
only of a vegetative system—a sort of compact mycelium—(probably of
species of Xylaria) with some affinity to Sclerotium.

Recently an extraordinary instance of luminosity was recorded as
occurring in our own country.[H] “A quantity of wood had been purchased
in a neighbouring parish, which was dragged up a very steep hill to its
destination. Amongst them was a log of larch or spruce, it is not quite
certain which, 24 feet long and a foot in diameter. Some young friends
happened to pass up the hill at night, and were surprised to find the road
scattered with luminous patches, which, when more closely examined,
proved to be portions of bark or little fragments of wood. Following the
track, they came to a blaze of white light which was perfectly surprising.
On examination, it appeared that the whole of the inside of the bark of the
log was covered with a white byssoid mycelium of a peculiarly strong
smell, but unfortunately in such a state that the perfect form could not be
ascertained. This was luminous, but the light was by no means so bright as
in those parts of the wood where the spawn had penetrated more deeply,
and where it was so intense that the roughest treatment scarcely seemed to
check it. If any attempt was made to rub off the luminous matter it only
shone the more brightly, and when wrapped up in five folds of paper the
light penetrated through all the folds on either side as brightly as if the
specimen was exposed; when, again, the specimens were placed in the
pocket, the pocket when opened was a mass of light. The luminosity had
now been going on for three days. Unfortunately we did not see it ourselves
till the third day, when it had, possibly from a change in the state of
electricity, been somewhat impaired; but it was still most interesting, and
we have merely recorded what we observed ourselves. It was almost
possible to read the time on the face of a watch even in its less luminous
condition. We do not for a moment suppose that the mycelium is essentially
luminous, but are rather inclined to believe that a peculiar concurrence of
climatic conditions is necessary for the production of the phenomenon,
which is certainly one of great rarity. Observers as we have been of fungi in
their native haunts for fifty years, it has never fallen to our lot to witness a
similar case before, though Prof. Churchill Babington once sent us
specimens of luminous wood, which had, however, lost their luminosity
before they arrived. It should be observed that the parts of the wood which
were most luminous were not only deeply penetrated by the more delicate
parts of the mycelium, but were those which were most decomposed. It is
probable, therefore, that this fact is an element in the case as well as the
presence of fungoid matter.”

In all cases of phosphorescence recorded, the light emitted is described as
of the same character, varying only in intensity. It answers well to the name
applied to it, as it seems remarkably similar to the light emitted by some
living insects and other animal organisms, as well as to that evolved, under
favourable conditions, by dead animal matter—a pale bluish light,
resembling that emitted by phosphorus as seen in a dark room.
Another phenomenon worthy of note is the change of colour which the
bruised or cut surface of some fungi undergo. Most prominent amongst
these are certain poisonous species of Boletus, such, for instance, as Boletus
luridus, and some others, which, on being bruised, cut, or divided, exhibit
an intense, and in some cases vivid, blue. At times this change is so
instantaneous that before the two freshly-cut portions of a Boletus can be
separated, it has already commenced, and proceeds rapidly till the depth of
intensity has been gained. This blue colour is so universally confined to
dangerous species that it is given as a caution that all species which exhibit
a blue colour when cut or bruised, should on no account be eaten. The
degree of intensity varies considerably according to the condition of the
species. For example, Boletus cærulescens is sometimes only very slightly,
if at all, tinged with blue when cut, though, as the name implies, the
peculiar phenomenon is generally highly developed. It cannot be said that
this change of colour has as yet been fully investigated. One writer some
time since suggested, if he did not affirm, that the colour was due to the
presence of aniline, others have contented themselves with the affirmation
that it was a rapid oxidization and chemical change, consequent upon
exposure of the surfaces to the air. Archdeacon Robinson examined this
phenomenon in different gases, and arrived at the conclusion that the
change depends on an alteration of molecular arrangement.[I]
One of the best of the edible species of Lactarius, known as Lactarius
deliciosus, changes, wherever cut or bruised, to a dull livid green. This
fungus is filled with an orange milky fluid, which becomes green on
exposure to the air, and it is consequently the juice which oxidizes on
exposure. Some varieties more than others of the cultivated mushroom
become brownish on being cut, and a similar change we have observed,
though not recorded, in other species.
The presence of a milky juice in certain fungi has been alluded to. This is
by no means confined to the genus Lactarius, in which such juice is

universal, sometimes white, sometimes yellow, and sometimes colourless.
In Agarics, especially in the subgenus Mycena, the gills and stem are replete
with a milky juice. Also in some species of Peziza, as for instance in Peziza
succosa, B., sometimes found growing on the ground in gardens, and in
Peziza saniosa, Schrad., also a terrestrial species, the same phenomenon
occurs. To this might be added such species as Stereum spadiceum, Fr., and
Stereum sanguinolentum, Fr., both of which become discoloured and
bleeding when bruised, while Corticium lactescens distils a watery milk.
Fungi in general have not a good repute for pleasant odours, and yet it
must be conceded that they are not by any means devoid of odour,
sometimes peculiar, often strong, and occasionally very offensive. There is
a peculiar odour common to a great many forms, which has come to be
called a fungoid odour; it is the faint smell of a long-closed damp cellar, an
odour of mouldiness and decay, which often arises from a process of
eremocausis. But there are other, stronger, and equally distinct odours,
which, when once inhaled, are never to be forgotten. Amongst these is the
fetid odour of the common stinkhorn, which is intensified in the more
beautiful and curious Clathrus. It is very probable that, after all, the odour
of the Phallus would not be so unpleasant if it were not so strong. It is not
difficult to imagine, when one encounters a slight sniff borne on a passing
breeze, that there is the element of something not by any means unpleasant
about the odour when so diluted; yet it must be confessed that when carried
in a vasculum, in a close carriage, or railway car, or exposed in a close
room, there is no scruple about pronouncing the odour intensely fetid. The
experience of more than one artist, who has attempted the delineation of
Clathrus from the life, is to the effect that the odour is unbearable even by
an enthusiastic artist determined on making a sketch.
Perhaps one of the most fetid of fungi is Thelephora palmata. Some
specimens were on one occasion taken by Mr. Berkeley into his bedroom at
Aboyne, when, after an hour or two, he was horrified at finding the scent far
worse than that of any dissecting room. He was anxious to save the
specimens, but the scent was so powerful that it was quite intolerable till he
had wrapped them in twelve thick folds of the strongest brown paper. The
scent of Thelephora fastidiosa is bad enough, but, like that of Coprinus
picaceus, it is probably derived from the imbibition of the ordure on which
it is developed. There needs no stronger evidence that the scent must not

only be powerful, but unpleasant, when an artist is compelled, before a
rough sketch is more than half finished, to throw it away, and seek relief in
the open air. A great number of edible Agarics have the peculiar odour of
fresh meal, but two species, Agaricus odorus and Agaricus fragrans, have a
pleasant anise-like odour. In two or three species of tough Hydnum, there is
a strong persistent odour somewhat like melilot or woodruffe, which does
not pass away after the specimen has been dried for years. In some species
of Marasmius, there is a decidedly strong odour of garlic, and in one
species of Hygrophorus, such a resemblance to that of the larva of the goat
moth, that it bears the name of Hygrophorus cossus. Most of the fleshy
forms exhale a strong nitrous odour during decay, but the most powerful we
remember to have experienced was developed by a very large specimen of
Choiromyces meandriformis, a gigantic subterranean species of the truffle
kind, and this specimen was four inches in diameter when found, and then
partially decayed. It was a most peculiar, but strong and unpleasantly
pungent nitrous odour, such as we never remember to have met with in any
other substance. Peziza venosa is remarkable when fresh for a strong scent
like that of aquafortis.
Of colour, fungi exhibit an almost endless variety, from white, through
ochraceous, to all tints of brown until nearly black, or through sulphury
yellow to reds of all shades, deepening into crimson, or passing by vinous
tints into purplish black. These are the predominating gradations, but there
are occasional blues and mineral greens, passing into olive, but no pure or
chlorophyllous green. The nearest approach to the latter is found in the
hymenium of some Boleti. Some of the Agarics exhibit bright colours, but
the larger number of bright-coloured species occur in the genus Peziza.
Nothing can be more elegant than the orange cups of Peziza aurantia, the
glowing crimson of Peziza coccinea, the bright scarlet of Peziza rutilans,
the snowy whiteness of Peziza nivea, the delicate yellow of Peziza
theleboloides, or the velvety brown of Peziza repanda. Amongst Agarics,
the most noble Agaricus muscarius, with its warty crimson pileus, is
scarcely eclipsed by the continental orange Agaricus cæsarius. The
amethystine variety of Agaricus laccatus is so common and yet so
attractive; whilst some forms and species Russula are gems of brilliant
colouring. The golden tufts of more than one species of Clavaria are
exceedingly attractive, and the delicate pink of immature Lycogala
epidendrum is sure to command admiration. The minute forms which

require the microscope, as much to exhibit their colour as their structure,
are not wanting in rich and delicate tints, so that the colour-student would
find much to charm him, and good practice for his pencil in these much
despised examples of low life.
Amongst phenomena might be cursorily mentioned the peculiar
sarcodioid mycelium of Myxogastres, the development of amœboid forms
from their spores, and the extraordinary rapidity of growth, as the well-
known instance of the Reticularia which Schweinitz observed running over
iron a few hours after it had been red hot. Mr. Berkeley has observed that
the creamy mycelium of Lycogala will not revive after it has become dry
for a few hours, though so active before.
[A]
M. J. Berkeley, “Introduction to Cryptogamic Botany,” p. 265.
[B]
Tulasne, “Sur la Phosphorescence des Champignons,” in “Ann. des Sci. Nat.” (1848),
vol. ix, p. 338.
[C]
In “Hooker’s Journal of Botany” (1840), vol. ii. p. 426.
[D]
Berkeley, “Introduction to Crypt. Bot.” t. 265.
[E]
Dr. Collingwood, in “Journal of Linnæan Society (Botany),” vol. x. p. 469.
[F]
In “Hooker’s Journal of Botany” for April, 1842.
[G]
Tulasne, “Sur la Phosphorescence,” in “Ann. des Sci. Nat.” (1848), vol ix. p. 340, &c.
[H]
Rev. M. J. Berkeley, in “Gardener’s Chronicle” for 1872, p. 1258.
[I]
Berkeley, “Introduction to Crypt. Bot.” p. 266.

VI.
THE SPORE AND ITS DISSEMINATION.

A work of this character would hardly be deemed complete without some
reference to the above subject, which has moreover a relation to some of the
questions discussed, and particularly of spore diffusion in the atmosphere.
The largest spore is microscopic, and the smallest known scarcely visible
under a magnifying power of 360 diameters. Taking into account the large
number of species of fungi, probably scarcely less numerous than all the
flowering plants, and the immense number of spores which some of the
individuals produce, they must be exceedingly plentiful and widely
diffused, though from their minuteness not easy to be discerned. It has been
attempted to estimate the number of spores which might be produced by
one single plant of Lycoperdon, but the number so far exceeds that which
the mind is accustomed to contemplate that it seems scarcely possible to
realize their profusion. Recent microscopic examinations of the common
atmosphere[A] show the large quantity of spores that are continually
suspended. In these investigations it was found that spores and similar cells
were of constant occurrence, and were generally present in considerable
numbers. That the majority of the cells were living, and ready to undergo
development on meeting with suitable conditions, was very manifest, as in
those cases in which preparations were retained under observation for any
length of time, germination rapidly took place in many of the cells. In few
instances did any development take place, beyond the formation of
networks of mycelium, or masses of toruloid cells, but, in one or two,
distinct sporules were developed on the filaments arising from some of the
larger septate spores; and in a few others, Penicillium and Aspergillus
produced their characteristic heads of fructification. With regard to the
precise nature of the spores, and other cells present in various instances,
little can be said, as, unless their development were to be carefully followed
out through all its stages, it is impossible to refer them to their correct
species or even genera. The greater number of them are apparently referable

to the old orders of fungi, Sphæronemei, Melanconei, Torulacei, Dematiei
and Mucedines, while some probably belonged to the Pucciniæi and
Cæomacei.
Hence it is demonstrated that a large number of the spores of fungi are
constantly present in the atmosphere, which is confirmed by the fact that
whenever a suitable pabulum is exposed it is taken possession of by floating
spores, and soon converted into a forest of fungoid vegetation. It is admitted
that the spores of such common moulds as Aspergillus and Penicillium are
so widely diffused, that it is almost impossible to exclude them from closed
vessels, or the most carefully guarded preparations. Special contrivances for
the dispersion of the spores in the different groups follow a few general
types, and it is only rarely that we meet with any method that is confined
only to a species or genus. Some of the more significant forms of spores
may be illustrated, with their modes of dissemination.
Basidiospores is a term which we may employ here to designate all
spores borne at the tips of such supports as are found in the Hymenomycetes
and Gasteromycetes, to which the name of basidia has been given. In fact,
under this section we may include all the spores of those two orders,
although we may be ignorant of the precise mode in which the fruit of most
of the Myxogastres is developed. Guarding ourselves at the outset against
any misinterpretation as to the use of this term, which, in fact, we employ
simply to designate the fruit of Hymenomycetes, we may have excuse in our
desire to limit special terms as much as possible. In the Agaricini the spores
are plentiful, and are distributed over the hymenium or gill plates, the
surface of which is studded with basidia, each of which normally terminates
with four short, erect, delicate, thread-like processes, each of which is
surmounted by a spore. These spores are colourless or coloured, and it is
upon this fact that primary divisions in the genus Agaricus are based,
inasmuch as colour in the spores appears to be a permanent feature. In
white-spored species the spores are white in all the individuals, not mutable
as the colour of the pileus, or the corolla in phanerogamic plants. So also
with the pink spored, rusty spored, black spored, and others. This may serve
to explain why colour, which is so little relied upon in classification
amongst the higher plants, should be introduced as an element of
classification in one of the largest genera of fungi.

Fig. 45.—Spores of (a) Agaricus mucidus; (b) Agaricus
vaginatus; (c) Agaricus pascuus; (d) Agaricus nidorosus; (e)
Agaricus campestris. (Smith.)

Fig. 46.—Spores of (a) Lactarius blennius; (b) Lactarius
fuliginosus; (c) Lactarius quietus. (Smith.)

There are considerable differences in size and form amongst the spores of
the Agaricini, although at first globose; when mature they are globose, oval,
oblong, elliptic, fusiform, and either smooth or tuberculated, often
maintaining in the different genera or subgenera one particular
characteristic, or typical form. It is unnecessary here to particularize all the
modifications which the form and colour of the spores undergo in different
species, as this has already been alluded to. The spores in the Polyporei,
Hydnei, &c., are less variable, of a similar character, as in all the
Hymenomycetes, except perhaps the Tremellini.

Fig. 46a.—(a) Spore of Gomphidius viscidus; (b)
spore of Coprinus micaceus.

Fig. 47.—Spores of (a) Polyporus cæsius; (b)
Boletus parasiticus; (c) Hydnum.

When an Agaric is mature, if the stem is cut off close to the gills, and the
pileus inverted, with the gills downwards on a sheet of black paper (one of
the pale-spored species is best for this purpose), and left for a few hours, or
all night, in that position, the paper will be found imprinted in the morning
with a likeness of the under side of the pileus with its radiating gills, the
spores having been thrown down upon the paper in such profusion, from the
hymenium, and in greater numbers from the opposed surfaces of the gills.
This little experiment will be instructive in two or three points. It will
illustrate the facility with which the spores are disseminated, the immense
number in which they are produced, and the adaptability of the gill structure
to the economy of space, and the development of the largest number of
basidiospores from a given surface. The tubes or pores in Polyporei, the
spines in Hydnei, are modifications of the same principles, producing a like
result.
In the Gasteromycetes the spores are produced in many cases, probably
in most, if not all, at the tips of sporophores; but the hymenium, instead of
being exposed, as in the Hymenomycetes, is enclosed within an outer
peridium or sac, which is sometimes double. The majority of these spores
are globose in form, some of them extremely minute, variously coloured,
often dark, nearly black, and either externally smooth or echinulate. In
some genera, as Enerthenema, Badhamia, &c., a definite number of spores
are at first enclosed in delicate cysts, but these are exceptions to the general
rule: this also is the case in at least one species of Hymenogaster. As the
spores approach maturity, it may be observed in such genera as Stemonitis,
Arcyria, Diachea, Dictydium, Cribraria, Trichia, &c., that they are
accompanied by a sort of reticulated skeleton of threads, which remain
permanent, and served in earlier stages, doubtless, as supports for the
spores; being, in fact, the skeleton of the hymenium. It has been suggested
that the spiral character of the threads in Trichia calls to mind the elaters in
the Hepaticæ, and like them may, by elasticity, aid in the dispersion of the

spores. There is nothing known, however, which will warrant this view.
When the spores are mature, the peridium ruptures either by an external
orifice, as in Geaster, Lycoperdon, &c., or by an irregular opening, and the
light, minute, delicate, spores are disseminated by the slightest breath of air.
Specimens of Geaster and Bovista are easily separated from the spot on
which they grew; when rolling from place to place, the spores are deposited
over a large surface. In the Phalloidei the spores are involved in a slimy
mucus which would prevent their diffusion in such a manner. This
gelatinous substance has nevertheless a peculiar attraction for insects, and it
is not altogether romantic to believe that in sucking up the fetid slime, they
also imbibe the spores and transfer them from place to place, so that even
amongst fungi insects aid in the dissemination of species. Whether or not
the Myxogastres should be included here is matter of opinion, since the
mode in which the spores are developed is but little known; analogy with
the Trichogastres in other points alone leading to the conclusion that they
may produce basidiospores. The slender, elastic stems which support the
peridia in many species are undoubted aids to the dissemination of the
spores.[B]
Under the name of Stylospores may be classed those
spores which in some orders of Coniomycetes are produced at
the apex of short threads, either enclosed in a perithecium, or
seated upon a kind of stroma. These are exceedingly variable,
sometimes large, and multiseptate, at other times minute,
resembling spermatia. In such genera as are chiefly epiphytal,
in Septoria, Phyllosticta, and their allies, the minute spores
are enclosed within membranaceous perithecia, and when
mature these are ejected from the orifice at the apex, or are
exposed by the breaking off of the upper portion of the
perithecia. In Diplodia and Hendersonia the spores are larger,
mostly coloured, often very fine in the latter genus, and
multiseptate, escaping from the perithecia by a terminal pore.
Probably the species are only pycnidia of Sphæriacei, but that
is of no consequence in relation to our present inquiry. Of
stylospores which deserve mention on account of their Fig. 48.—
singularity of form, we may note those of Dilophospora Diachea
graminis, which are straight, and have two or three hair-like elegans.
appendages at each extremity. In Discosia there is a single oblique bristle at

each end, or at the side of the septate spores, whilst in Neottiospora a tuft of
delicate hairs is found at one extremity only. The appendages in
Dinemasporium are similar to those of Discosia. The spores in
Prosthemium may be said in some sort to resemble compound Hendersonia,
being fusiform and multiseptate, often united at the base in a stellate
manner. In this genus, as in Darluca, Cytispora, and the most of those
belonging to the Melanconiei, the spores when mature are expelled from the
orifice of the perithecium or spurious perithecium, either in the form of
tendrils, or in a pasty mass. In these instances the spores are more or less
involved in gelatine, and when expelled lie spread over the matrix, around
the orifice; their ultimate diffusion being due to moisture washing them
over other parts of the same tree, since it is probable that their natural area
of dissemination is not large, the higher plants, of which they are mostly
conditions, being developed on the same branches. More must be known of
the relations between Melanconium and Tulasne’s sphæriaceous genus
Melanconis before we can appreciate entirely the advantage to
Melanconium and some other genera, that the wide diffusion of their spores
should be checked by involving them in mucus, or their being agglutinated
to the surface of the matrix, only to be softened and diffused by rain. The
spores in many species amongst the Melanconiei are remarkably fine; those
of Stegonosporium have the endochrome partite and cellular. In Stilbospora
and Coryneum the spores are multiseptate, large, and mostly coloured. In
Asterosporium the spores are stellate, whilst in Pestalozzia they are septate,
with a permanent peduncle, and crested above with two or three hyaline
appendages.

Fig. 50.—Spores of
Dilophospora graminis.

Fig. 49.—Spore of Hendersonia
polycystis.

Fig. 52.—Spore of Prosthemium
Fig. 51.—Spores of Discosia. betulinum.

Fig. 53.—Spore of Fig. 54.—Stylospores of
Stegonosporium cellulosum. Coryneum disciforme.

Fig. 55.—Spores of Asterosporium
Hoffmanni. Fig. 56.—Spores of Pestalozzia.

Fig. 57.—Bispora monilioides.

The Torulacei externally, and to the naked eye, are very similar to the
black moulds, and the mode of dissemination will be alike in both. The
spores are chiefly compound, at first resembling septate threads, and at
length breaking up into joints, each joint of which possesses the function of
a spore. In some instances the threads are connate, side by side, as in Torula
hysterioides, and in Speira, being concentrically arranged in laminæ in the
latter genus. The structure in Sporochisma is very peculiar, the joints
breaking up within an external tube or membrane. The spores in
Sporidesmium appear to consist of irregular masses of cells, agglomerated
into a kind of compound spore. Most of the species become pulverulent,
and the spores are easily diffused through the air like an impalpable dust.

They form a sort of link between the stylospores of one section of the
Coniomycetes, and the pseudospores of the parasitical section.
Pseudospore is, perhaps, the most fitting name which can be applied to
the so-called spores of the parasitical Coniomycetes. Their peculiar
germination, and the production of reproductive bodies on the germ tubes,
prove their analogy to some extent with the prothallus of other cryptogams,
and necessitate the use of some term to distinguish them from such spores
as are reproductive without the intervention of a promycelium. The
differences between these pseudospores in the several genera are confined
in some instances to their septation, in others to their mode of development.
In the Æcidiacei the pseudospores are more or less globose, produced in
chains within an external cellular peridium. In the Cæomacei they are
simple, sometimes produced in chains, and sometimes free, with or without
a caduceous peduncle. In the Ustilaginei they are simple, dark coloured,
and occasionally attached in subglobose masses, as in Urocystis and
Thecaphora, which, are more or less compact. In the Pucciniæi the
distinctive features of the genera are based upon the more or less complex
nature of the pseudospores, which are bilocular in Puccinia, trilocular in
Triphragmium, multilocular in Phragmidium, &c. In the curious genus
Podisoma the septate pseudospores are involved in a gelatinous element.
The diffusion of these fruits is more or less complete according to their
compact or pulverulent nature. In some species of Puccinia the sori are so
compact that they remain attached to the leaves long after they are dead and
fallen. In the genus Melampsora, the wedge-shaped winter-pseudospores
are not perfected until after the dead leaves have for a long time remained
and almost rotted on the ground. It is probable that their ultimate diffusion
is only accomplished by the rotting and disintegration of the matrix. In the
Cæomacei, Ustilaginei, and Æcidiacei the pseudospores are pulverulent, as
in some species of Puccinia, and are easily diffused by the motion of the
leaves in the wind, or the contact of passing bodies. Their diffusion in the
atmosphere seems to be much less than in the case of the Hyphomycetes. By
what means such a species as Puccinia malvacearum, which has very
compact sori, has become within so short a period diffused over such a wide
area, is a problem which in the present state of our knowledge must remain
unsolved. It may be through minute and plentiful secondary spores.

Fig. 58.—Pseudospores of Fig. 59.—Pseudospores of Fig. 60.—Pseudospores of
Thecaphora hyalina. Puccinia. Triphragmium.

Fig. 61.—Pseudospores of Phragmidium Fig. 62.—Melampsora salicina.
bulbosum. (Winter fruit.)

Spermatia are very minute delicate bodies found associated with many of
the epiphyllous Coniomycetes, and it has been supposed are produced in
conjunction with some of the Sphæriacei, but their real function is at
present obscure, and the name is applied rather upon conjecture than
knowledge. It is by no means improbable that spermatia do exist
extensively amongst fungi, but we must wait in patience for the history of
their relationship.

Trichospores might be applied better, perhaps, than conidia to the spores
which are produced on the threads of the Hyphomycetes. Some of them are
known to be the conidia of higher plants; but as this is by no means the case
with all, it would be assuming too much to give the name of conidia to the
whole. By whatever name they may be called, the spores of the
Hyphomycetes are of quite a different type from any yet mentioned,
approximating, perhaps, most closely to the basidiospores of the
Hymenomycetes in some, and Gasteromycetes in others; as, for instance, in
the Sepedoniei and the Trichodermacei. The form of the spores and their
size differ materially, as well as the manner in which they are produced on
the threads. In many they are very minute and profuse, but larger and less
plentiful in the Dematiei than in the Mucedines. The spores of some species
of Helminthosporium are large and multiseptate, calling to mind the spores
of the Melanconiei. Others are very curious, being stellate in Triposporium,
circinate in Helicoma and Helicocoryne, angular in Gonatosporium, and
ciliate in Menispora ciliata. Some are produced singly and some in chains,
and in some the threads are nearly obsolete. In Peronospora, it has been
demonstrated that certain species produce minute zoospores from the so-
called spores. The dissemination of the minute spores of the Mucedines
through the air is undoubted; rain also certainly assists not only in the
dispersion of the spores in this as in other groups, but also in the production
of zoospores which require moisture for that purpose. The form of the
threads, and the mode of attachment of the spores, is far more variable
amongst the Mucedines than the form of the spores, but the latter are in all
instances so slightly attached to their supports as to be dissevered by the
least motion. This aids also in the diffusion of the spores through the
atmosphere.
Sporangia are produced in the Physomycetes
usually on the tips or branches of delicate threads,
and these when mature dehisce and set free the
minute sporidia. These are so small and uniform
in their character that they require but a passing
mention. The method of diffusion agrees much
with that of the Mucedines, the walls of the
sporangia being usually so thin and delicate as to
be easily ruptured. Other modes of fructification
prevail in some species by the production of

cysts, which are the result of conjugation of the Fig. 63.—Spores of
Helicocoryne.
threads. These bodies are for the most part
furnished with thicker and more resistant walls, and the diffusion of their
contents will be regulated by other circumstances than those which
influence the dispersion of the minute sporidia from the terminal cysts.
Probably they are more perennial in their character, and are assimilated
more to the oogonia of Cystopus and Peronospora, being rather of the
nature of resting spores, inasmuch as the same threads usually bear the
terminal fruits.
Thecaspores is a
term which may be
applied generally to all
sporidia produced in
asci, but these are in
Fig. 64.—Sporidium of Genea turn so innumerable
verrucosa.
and variable that it will Fig. 65.—Alveolate sporidium
of Tuber.
be necessary to treat of some of the groups
individually. The Thecaspores, for instance, of the Tuberacei offer several
features whereby they may be distinguished from other thecaspores. The
asci in which these sporidia are generated mostly partake of a broadly
saccate, ovate form. The number of sporidia contained in an individual
ascus is usually less than in the majority of the Ascomycetes, and the
sporidia approximate more nearly to the globose form. Usually, also, they
are comparatively large. Many have been figured by Corda[C] and Tulasne.
[D] Three types of spores may be said to prevail in the Tuberacei: the
smooth spored, the warted or spinulose, and the areolate. The first of these
may be represented by the Stephensia bombycina, in which the globose
sporidia are quite smooth and colourless. The warted sporidia may be
observed in Genea verrucosa, the spinulose in Tuber nitidum, and the
areolate are present in Tuber æstivum and Tuber excavatum, in which the
epispore is divided into polygonal alveoli, bounded by thin,
membranaceous, prominent partitions. This form of sporidium is very
beautiful. In all no special provision is made for the dissemination of the
sporidia, as, from their subterranean habit, none would be available save the
ultimate dissolution of the external integuments. As they are greedily
devoured by several animals, it is possible that they may be dispersed
through the excrements.

In the Perisporiacei the perithecium has no proper orifice, or ostiolum,
for the discharge of the mature sporidia, which are usually small, and are
disseminated by the irregular rupture of the somewhat fragile conceptacles.
The asci are usually more or less saccate, and the sporidia approximate to a
globose form. The asci are often very diffluent. In Perisporium vulgare the
ovate brown sporidia are at first, and for some time, attached together in
fours in a concatenate or beaded manner. In some species of Erysiphei the
conceptacle encloses but a single sporangium, in others several, which are
attached together at the base. In some species the sporangia contain two, in
others four, in others eight, and in others numerous sporidia. In Chætomium
the asci are cylindrical, and in most cases the coloured sporidia are lemon-
shaped. When the conceptacles are fully matured, it is commonly the case
that the asci are absorbed and the sporidia are free in the interior of the
conceptacles.
Of the fleshy Discomycetes the genus Peziza
may be taken as the type. If the structure which
prevails in this genus be brought to mind, it will
be remembered that the hymenium lines an
expanded cup, and that the asci are packed
together, side by side, with their apices outwards,
and their bases attached to a substratum of cells
which form the inner layer of the receptacle. The
sporidia are usually eight in each ascus, either
arranged in single or double rows, or irregularly
grouped together. The asci are produced in
succession; the later, pressing themselves
upwards between those previously developed,
cause the rupture of the mature asci at the apex
and the ejection of the sporidia with considerable
force. When a large Peziza is observed for a time
a whitish cloud will be seen to rise suddenly from
the surface of the disc, which is repeated again
and again whenever the specimen is moved. This
cloud consists of sporidia ejected simultaneously
from several asci. Sometimes the ejected sporidia
Fig. 66.—Asci, sporidia, and
lie like frost on the surface of the disc. Theories
paraphyses of Ascobolus have been devised to account for this sudden

(Boudier). extrusion of the sporidia, in Ascobolus, and a few
species of Peziza, of the asci also, the most feasible one being the
successive growth of the asci; contraction of the cup may also assist, as well
as some other less potent causes. It may be remarked here that the sporidia
in Peziza and Helotium are mostly colourless, whilst in Ascobolus they pass
through pink to violet, or dark brown, and the epispore, which is of a waxy
nature, becomes fissured in a more or less reticulated manner.
The sporidia in Hysterium proper are usually
coloured, often multiseptate, sometimes
fenestrate, and occasionally of considerable size.
There is no evidence that the sporidia are ever
excluded in the same manner as in Peziza, the
lips closing over the disc so much as to prevent
this. The diffusion of the sporidia probably
depends on the dissolution of the asci, and hence
they will not be widely dispersed, unless,
perhaps, by the action of rain.
Fig. 67.—Sporidium of
In Tympanis, asci of two kinds have been Ostreichnion Americanum.
observed in some species; one kind containing an
indefinite number of very minute bodies resembling spermatia, and the
other octosporous, containing sporidia of the usual type.
The Sphæriacei include an almost infinite variety in the form and
character of the sporidia. Some of these are indefinite in the number
contained in an ascus, although the majority are eight, and a few less. In the
genera Torrubia and Hypocrea the structure differs somewhat from other
groups, inasmuch as in the former the long thread-like sporidia break up
into short joints, and in the latter the ascus contains sixteen subglobose or
subquadrate sporidia. Other species contain linear sporidia, which are often
the length of the ascus, and may either be simple or septate. In Sphæria
ulnaspora the sporidia are abruptly bent at the second joint. Shorter
fusiform sporidia are by no means uncommon, varying in the number of
septa, and in constriction at the joints in different species. Elliptic or ovate
sporidia are common, as are those of the peculiar form which may be
termed sausage-shaped. These are either hyaline or coloured of some shade
of brown. Coloured sporidia of this kind are common in Xylaria and
Hypoxylon, as well as in certain species of the section Superficiales.

Coloured sporidia are often large and beautiful: they are mostly of an
elongated, elliptical form, or fusiform. As noteworthy may be mentioned
the sporidia of Melanconis lanciformis, those of Valsa profusa, and some
species of Massaria, the latter being at first invested with a hyaline coat.
Some coloured sporidia have hyaline appendages at each extremity, as in
Melanconis Berkeleii, and an allied species, Melanconis bicornis, from the
United States, also some dung Sphæriæ, as S. fimiseda, included under the
proposed genus Sordaria.[E] Hyaline sporidia occasionally exhibit a delicate
bristle-like appendage at each extremity, as in the Valsa thelebola, or with
two additional cilia at the central constriction, as in Valsa taleola. A peculiar
form of sporidium is present in certain species of Sphæria found on dung,
for which the generic name of Sporormia has been proposed, in which the
sporidium (as in Perisporium vulgare) consists of four coloured ovate
joints, which ultimately separate. Multiseptate fenestrate sporidia are not
uncommon in Cucurbitaria and Pleospora, as well as in Valsa fenestrata
and some other species. In the North American Sphæria putaminum the
sporidia are extraordinarily large.

Fig. 68.—Ascus and sporidia of Fig. 69.—Sporidium of Sphæria
Hypocrea. ulnaspora.

Fig. 70.—Sporidia of Valsa profusa Fig. 71.—Sporidia of Massaria
(Currey). fœdans. × 400.

Fig. 72.—Sporidium of Fig. 73.—Caudate sporidia of
Melanconis bicornis, Cooke. Sphæria fimiseda.

Fig. 74.—Sporidia of Valsa Fig. 75.—Sporidia of Valsa
thelebola. taleola. × 400.

Fig. 76.—Sporidium of Fig. 77.—Asci and sporidia of Sphæria
Sporormia intermedia. (Pleospora) herbarum.

The dissemination of the sporidia may,
from identity of structure in the
perithecium, be deemed to follow a like
method in all. When mature, they are in a
great measure expelled from the mouth of
the perithecia, as is evident in species with
large dark sporidia, such as exist in the
genera Hypoxylon, Melanconis, and
Massaria. In these genera the sporidia, on
maturity, may be observed blackening the
matrix round the mouths of the perithecia.
As moisture has an evident effect in Fig. 78.—Sporidium of Sphæria
producing an expulsion of sporidia by putaminum. × 400.
swelling the gelatinous nucleus, it may be assumed that this is one of the
causes of expulsion, and therefore of aids to dissemination. When Sphæriæ

are submitted to extra moisture, either by placing the twig which bears them
on damp sand, or dipping one end in a vessel of water, the sporidia will
exude and form a gelatinous bead at the orifice. There may be other
methods, and possibly the successive production of new asci may also be
one, and the increase in bulk by growth of the sporidia another; but of this
the evidence is scanty.
Finally, Oogonia may be mentioned as occurring in such genera as
Peronospora amongst moulds, Cystopus amongst Uredines, and the
Saprolegniaceæ amongst the Physomycetes. The zoospores being furnished
with vibratile cilia, are for some time active, and need only water in which
to disseminate themselves, and this is furnished by rain.
We have briefly indicated the characteristics of some of the more
important types of spores to be found in fungi, and some of the modes by
which it is known, or presumed, that their dissemination takes place. In this
summary we have been compelled to rest content with suggestions, since an
exhaustive essay would have occupied considerable space. The variability
in the fruit of fungi, in so far as we have failed to demonstrate, will be
found exhibited in the illustrated works devoted more especially to the
minute species.[F]
[A]
Cunningham, in “Ninth Annual Report of the Sanitary Commissioner with the
Government of India.” Calcutta, 1872.
[B]
See “Corda Icones,” tab. 2.
[C]
Corda, “Icones Fungorum,” vol. vi. Prague.
[D]
Tulasne, “Fungi Hypogæi.” Paris.
[E]
Winter, “Die Deutschen Sordarien” (1873).
[F]
Corda, “Icones Fungorum,” 6 vols. (1837–1842); Sturm, “Deutschlands Flora,” Pilze
(1841); Tulasne, “Selecta Fungorum Carpologia;” Bischoff, “Kryptogamenkunde”
(1860); Corda, “Anleitung zum Studium der Mykologie” (1842); Fresenius, “Beiträge

zur Mykologie” (1850); Nees Ton Esenbeck, “Das System der Pilze” (1816);
Bonorden, “Handbuch der Allgemeinen Mykologie” (1851).

VII.
GERMINATION AND GROWTH.

In describing the structure of these organisms in a previous chapter, the
modes of germination and growth from the spores have been purposely
excluded and reserved for the present. It may be assumed that the reader,
having followed us to this point, is prepared for our observations by some
knowledge of the chief features of structure in the principal groups, and of
the main distinctions in the classification, or at least sufficient to obviate
any repetition here. In very many species it is by no means difficult to
induce germination of the spores, whilst in others success is by no means
certain.
M. de Seynes made the Hymenomycetes an especial object of study,[A]
but he can give us no information on the germination and growth of the
spore. Hitherto almost nothing is positively known. As to the form of the
spore, it is always at first spherical, which it retains for a long time, while
attached to the basidia, and in some species, but rarely, this form is final, as
in Ag. terreus, &c. The most usual form is either ovoid or regularly elliptic.
All the Coprini have the spores oval, ovoid, more or less elongated or
attenuated from the hilum, which is more translucent than the rest of the
spore. This last form is rather general amongst the Leucospores, in Amanita,
Lepiota, &c. At other times the spores are fusiform, with regularly
attenuated extremities, as in Ag. ermineus, Fr., or with obtuse extremities, as
in Ag. rutilans, Sch. In Hygrophorus they are rather irregular, reniform, or
compressed in the centre all round. Hoffmann[B] has given a figure taken
from Ag. chlorophanus, and Seynes verified it upon Ag. ceraceus, Sow.
(See figures on page 121.)
The exospore is sometimes roughened, with more or less projecting
warts, as may be seen in Russula, which much resembles Lactarius in this
as in some other particulars. The spores of the Dermini and the Hyporhodii
often differ much from the sphærical form. In Ag. pluteus, Fr., and Ag.

phaiocephalus, Bull, there is already a commencement of the polygonal
form, but the angles are much rounded. It is in Ag. sericeus, Ag. rubellus,
&c., that the polygonal form becomes most distinct. In Dermini the angles
are more or less pronounced, and become rather acute in Ag. murinus, Sow.,
and Ag. ramosus, Bull. The passage from one to the other may be seen in
the stellate form of the conidia of Nyctalis.
It is almost always the external membrane that is coloured, which is
subject to as much variation as the form. The more fine and more delicate
shades are of rose, yellow-dun or yellow, violet, ashy-grey, clear fawn
colour, yellow-orange, olive-green, brick-red, cinnamon-brown, reddish-
brown, up to sepia-black and other combinations. It is only by the
microscope and transparency that one can make sure of these tints; upon a
sufficient quantity of agglomerated spores the colour may be distinguished
by the naked eye. Colour, which has only a slight importance when
considered in connection with other organs, acquires much in the spores, as
a basis of classification.
With the growth of Agarics from the mycelium, or spawn, we are not
deficient in information, but what are the conditions necessary to cause the
spores themselves to germinate before our eyes and produce this mycelium
is but too obscure. In the cultivated species we proceed on the assumption
that the spores have passed a period of probation in the intestines of the
horse, and by this process have acquired a germinating power, so that when
expelled we have only to collect them, and the excrement in which they are
concealed, and we shall secure a crop.[C] As to other species, we know that
hitherto all attempts to solve the mystery of germination and cultivation has
failed. There are several species which it would be most desirable to
cultivate if the conditions could be discovered which are essential to
germination.[D] In the same manner the Boleti and Hydnei—in fact, all other
hymenomycetal fungi, with the exception of the Tremellini—still require to
be interrogated by persevering experiment and close inquiry as to their
mode of germination, but more especially as to the essential conditions
under which alone a fruitful mycelium is produced.
The germination of the spore has been observed in some of the
Tremellini. Tulasne described it in Tremella violacea.[E] These spores are
white, unilocular, and filled with a plastic matter of homogeneous
appearance. From some portion of their surface an elongated germ filament

is produced, into which the contents of the
reproductive cell pass until quite exhausted.
Other spores, perhaps more abundant, have a
very different kind of vegetation. From their
convex side, more rarely from the outer edge,
these particular spores emit a conical process,
generally shorter than themselves, and directed
perpendicularly to the axis of their figure. This
appendage becomes filled with protoplasm at
the expense of the spore, and its free and
pointed extremity finally dilated into a sac, at
first globose and empty. This afterwards admits
into its cavity the plastic matter contained in its
support, and, increasing, takes exactly the form
of a new spore, without, however, quite
equalling in size the primary or mother spore. Fig. 79.—(a) Basidia and spores of
The spore of the new formation long retains its Exidia spiculosa; (b) Germinating
spore.
pedicel, and the mother spore which produced
it, but these latter organs are then entirely empty and extremely transparent.
Sometimes two secondary spores are thus engendered from the same spore,
and their pedicels may be implanted on the same or on different sides, so as
to be parallel in the former case, and growing in opposite directions in the
latter. The fate of these secondary spores was not determined.
In Dacrymyces deliquescens are found mingled
amongst the spores immense numbers of small
round or ovoid unilocular bodies, without
appendages of any kind, which long puzzled
mycologists. Tulasne ascertained that they are
derived from the spores of this fungus when they
have become free, and rest on the surface of the
hymenium. Each of the cells of the spore emits
exteriorly one or several of these corpuscles,
supported on very short slender pedicels, which
remain after the corpuscles are detached from Fig(a). 80.—Germinating spore and
corpuscles of Dacrymyces
them. This latter circumstance evidences that new deliquescens.
corpuscles succeed the firstborn one on each
pedicel as long as there remains any plastic matter within the spore. The

latter, in fact, in consequence of this labour of production, becomes
gradually emptied, and yet preserves the generative pedicels of the
corpuscles, even when it no longer contains any solid or coloured matter.
These pedicels are not all in the same plane, as may be ascertained by
turning the spore on its longitudinal axis; but it often seems to be so when
they are looked at in profile, on account of the very slight distance which
then separates them one from another. It will also be remarked that they are
in this case often implanted all on the same side of the reproductive body,
and most often on its convex side. Their fecundity is exhausted with the
plastic contents of the spore. The corpuscles, when placed in the most
favourable conditions, have never given the least sign of vegetation; they
have also remained for a long time in water without experiencing any
appreciable alteration.
All the individuals of Dacrymyces deliquescens do not produce these
corpuscles in the same abundance; those which bear the most are
recognizable by the pale tint of the reproductive dust with which they are
covered; in others, where this dust preserves its golden appearance, only a
few corpuscles are found. The spores which produce corpuscles do not
appear at all apt to germinate. On the other hand, multitudes of spores will
germinate which had not produced any corpuscles. Tulasne remarks on this,
that these observations would authorize us to think that all spores, though
perfectly identical to our eyes, have not, without distinction, the same fate,
nor doubtless the same nature; and, in the second place, that these two kinds
of bodies, if they are not always isolated, yet are most frequently met with
on distinct individuals. This author claims for the corpuscles in question
that they are spermatia, and thinks that their origin is only so far unusual in
that they proceed from veritable spores.
The whole of the Gasteromycetes have as yet to be challenged as to the
mode and conditions of germination and development. It is probable that
these will not materially differ from those which prevail in Hymenomycetes.
The germination in Æcidium has been followed out by Tulasne,[F] either
by placing the pseudospores in a drop of water, or confining them in a moist
atmosphere, or by placing the leaves on which the Æcidium flourishes upon
water. The pseudospores plunged in water germinated more readily than the
others. If the conditions were favourable, germination would take place in a
few hours. Æcidium Ranunculacearum, D. C., on leaves of figwort, gives

rarely more than one germinating filament, which soon attains three times
the length of the diameter of the pseudospore. This filament generally
remains simple, sometimes torulose, and distorted in a long spire.
Sometimes it has been seen divided into two branches, nearly equal to each
other. The spore in germinating empties itself of its plastic contents,
contracts, and diminishes in size. The pseudospores of Æcidium crassum,
P., emit three long filaments, which describe spirals, imitating the twistings
of the stem of a bean or bindweed. In Æcidium Violæ, Schum, one filament
is produced, which frequently rolls up its anterior extremity into a spire, but
more often this same extremity rises in a large ovoid, irregular vesicle,
which continues the axis of the filament, or makes with it a more or less
decided angle. In whatever manner placed, this vesicle attracts to it all the
orange protoplasm, and hardly does this become settled and complete
before the vesicle becomes the starting point of a new development, for it
begins to produce at its apex a filament, more slender than the previous one,
stiff, and unbranched.
According to M. Tulasne, the germination of
the pseudospores of Æcidium Euphorbiæ on
Euphorbia sylvatica differ in some respects from
the preceding. When dropped upon water these
spores very soon emit a short tube, which
ordinarily curves in an arch or circle, almost from
its origin, attaining a length of from three to six
times the diameter of the spore; then this tube
gives rise to four spicules, each of which
produces a small obovate or reniform sporule; the
generation of these sporules absorbs all the
plastic matter contained in the germ-tube, which
permits of the observation that it was divided into
four cells corresponding with the number of Fig. 81.—Germination of
spicules. These sporules germinate very rapidly Æcidium Euphorbia (sylvaticæ),
Tulasne.
from an indefinite point of their surface, emitting
a filiform process, which is flexuous and very delicate, not extending more
in length than three times that of the long axis of the sporule, often less,
reproducing at its summit a new sporule, differing in form and size from
that which preceded it. This sporule of the second formation becomes at its
apex a vital centre, and sprouts one or more linear buds, of which the

elongation is occasionally interrupted by the formation of vesicular
swellings. As Tulasne observes, the pseudospores of the Æcidium and the
greater number of Uredines are easily wetted with water before arriving at
maturity; but when they are ripe, on the contrary, they appear to be clothed
with a greasy matter which protects them from the liquid, forcing them
almost all to rest on the surface.
The pseudospores of Rœstelia are produced in strings or chaplets, as in
Æcidium, with this difference, that instead of being contiguous they are
separated by narrow isthmuses. The ripe pseudospores are enveloped in a
thick tegument, of a dark brown colour. They germinate readily on water,
producing a filament fifteen times as long as the diameter of the spore. This
filament is sometimes rolled or curved. Towards its extremity it exhibits
protuberances which resemble the rudiments of ramuli, or they terminate in
a vesicle which gives rise to a slender filament. The tegument of these
pseudospores, above all in those which have germinated, and have
consequently become more transparent, it is easy to see has many pores, or
round ostioles.
In Peridermium the pseudospores, when dropped upon water, germinate
at any point of their surface. Sometimes two unequal filaments issue from
the same spore. After forty-eight hours of vegetation in the air, the greater
part had already emitted a multitude of thick little branches, themselves
either simple or branched, giving to the filaments a peculiar aspect. Tulasne
did not on any occasion observe the formation of secondary spores.
In the Uredines proper the germination seems
to be somewhat similar, or at least not offering
sufficient differences to warrant special reference
in Uredo, Trichobasis, Lecythea, &c. In
Coleosporium there are two kinds of spores, one
kind consisting of pulverulent single cells, and
the other of elongated septate cells, which break
up into obovate joints. Soon after the maturity of
the pulverulent spores, each begins to emit a long
tube, which is habitually simple, and produces at
its summit a reproductive cellule, or reniform
sporule. The orange protoplasm passes along the
colourless tubes to the terminal sporule at the end

Fig. 82.—Germinating of its vegetation. The two forms of spores in this
pseudospores of (b)
genus are constantly found on the same leaf, and
Coleosporium Sonchi; (s s)
in the same pulvinule, but generally the
secondary spores, or sporules
(Tulasne). pulverulent spores abound at the commencement
of the summer. The reniform sporules begin to germinate in a great number
as soon as they are free; some few extend a filament which remains simple
and uniform, but more commonly it forms at its extremity a second sporule.
If this does not become isolated, to play an independent life, the filament is
continued, and new vesicles are repeated many times.
In Melampsora the summer spores are of
the Lecythea type, and were included in
that genus till their relation with
Melampsora was clearly made out. The
winter spores are in solid pulvinules, and
their fructification takes place towards the
end of winter or in the spring. This
phenomenon consists in the production of
cylindrical tubes, which start from the
upper extremity of the wedge-shaped
spores, or more rarely from the base. These
tubes are straight or twisted, simple or
bifurcated, and each of them very soon
emits four monosporous spicules, at the
same time that they become septate. The
sporules are in this instance globose.
In Uromyces germination follows
precisely the same type as that of the upper
cell of Puccinia; in fact, Tulasne states that
it is very difficult to say in what they differ Fig. 83.—Germinating pseudospore (b)
from the Pucciniæ which are accidentally of Melampsora betulina (Tulasne).
unilocular.
In Cystopus a more complex method prevails, which will be examined
more closely hereafter.
In Puccinia, as already observed when describing their structure, the
pseudospores are two-celled. From the pores of each cell, which are near

the central
septum,
springs a
clavate tube,
which attains
two or three
times the total
length of the
fruit, and of
which the very
obtuse
extremity
curves more or
less in the Fig. 85.—Germinating pseudospore of
Puccinia Moliniæ. (Tulasne.)
manner of a
crozier.[G] This tube, making a perfectly
uncoloured transparent membrane, is filled with a
granular and very pale plastic matter at the
Fig. 84.—Germinating expense of the generative cell, which is soon
pseudospore of Uromyce
appendiculatus. (Tulasne.) rendered vacant; then it gives rise to four
spicules, usually on the same side, and at the
summit of these produces a reniform cellule. The four sporules so
engendered exhaust all the protoplasm at first contained in the generative
cell, so that their united capacity proves to be evidently much insufficient to
contain it, the more so as it leads to the belief that this matter undergoes as
it condenses an elaboration which diminishes its size. In all cases the
spicule originates before the sporule which it carries, and also attains its full
length when the sporule appears. The form of the latter is at first globular,
then ellipsoid, and more or less curved. All these phases of vegetation are
accomplished in less than twelve hours, and if the spore is mature and ready
for germination, it is sufficient to provoke it by keeping the pseudospores in
a humid atmosphere. During this process the two cells do not separate, nor
does one commence germination before the other, but both simultaneously.
When the sporules are produced, the protospore, somewhat analogous to a
prothallus, has performed its functions and decays. Towards the time of the
falling of the sporules they are nearly all divided into four unequal cells by
transverse and parallel septa. These sporules in time produce, from any

point on their surface, a filament, which reproduces a new sporule,
resembling the first, but generally smaller. This sporule of the second
generation ordinarily detaches itself from its support before germinating.
The pseudospores of Triphragmium ulmariæ
have been seen in April germinating on old leaves
of the meadowsweet which survived the winter,
whilst at the same time new tufts of the spores
were being developed on the leaves of the year.
These fruits of the spring vegetation would not
germinate the same year. Each cell in germination
emits a long cylindrical filament, containing a
brownish protoplasm, on which four spicules,
bearing as many sporules, are generated.
The germination of
the black fruits of
Fig. 86.—Germinating Phragmidium only
pseudospore of Triphragmium
ulmariæ (Tulasne.) appears to take place in
the spring. It greatly
resembles that in Puccinia, except that the
filament is shorter, and the sporules are spherical
and orange-coloured, instead of being kidney-
shaped and pale. In the species found on the
leaves of the common bramble, the filament
emitted by each cell attains three or four times the
length of the fruit. The granular orange
protoplasm which fills it passes ere long into the
sporules, which are engendered at the extremity
of pointed spicules. After the long warty fruits are
emptied of their contents they still seem as dark
as before, but the pores which are pierced in the
sides, through which the germinating filaments Fig. 87.—Germinating
have proceeded, are more distinctly visible. pseudospore of Phragmidium
bulbosum. (Tulasne.)
It will be observed that throughout all these
allied genera of Uromyces, Puccinia, Triphragmium, and Phragmidium the
same type of germination prevails, which confirms the accuracy of their
classification together, and renders still less probable the supposed affinity

of Phragmidium with Sporidesmium, which was at one time held by very
astute mycologists, but which is now abandoned. This study of germination
leads also to a very definite conclusion with regard to the genus Uromyces
—that it is much more closely related to Puccinia and its immediate allies
than to other unicellular Uredines.

Fig. 88.—Germinating pseudospores of Podisoma Juniperi.
(Tulasne)

The germination of the pseudospores of the gelatinous Uredines of the
genus Podisoma was studied by Tulasne.[H] These pretended spores, he
writes, are formed of two large conical cells, opposed by their base and
easily separating. They vary in length. The membrane of which they are
formed is thin and completely colourless in most of them, though much
thicker and coloured brown in others. It is principally the spores with thin
membranes that emit from near the middle very obtuse tubes, into which by
degrees, as they elongate, the contents of the parent utricles pass. Each of
the two cells of the supposed spore may originate near its base four of these
tubes, opposed to each other at their point of origin, and their subsequent
direction; but it is rather rare for eight tubes, two by two, to decussate from
the same spore or basidium. Usually there are only two or three which are
completely developed, and these tend together towards the surface of the
fungus, which they pass, and expand at liberty in the air. The tubes
generally become thicker by degrees as they elongate, some only slightly
exceeding the length of the protospores. Others attain three or four times

that length, according to the greater or less distance between the protospore
and the surface of the plant. In the longest tubes it is easy to observe how
the colouring matter passes to their outer extremity, leaving the portion
nearest to the parent cell colourless and lifeless. When nearly attaining their
ultimate dimensions, all the tubes are divided towards their outer extremity
by transverse septa into unequal cells; then simple and solitary processes, of
variable length and form, but attenuated upwards, proceed from each
segment of the initial tube, and produce at their extremity an oval spore
(teleutospore, Tul.), which is slightly curved and unilocular. These spores
absorb all the orange endochrome from the original tubes. They appear in
immense numbers on the surface of the fungus, and when detached from
their spicules fall upon the ground or on any object which may be beneath
them. So freely are they deposited that they may be collected on paper, or a
slip of glass, like a fine gold-coloured powder. Again, these secondary
spores (teleutospores) are capable of germination, and many of them will be
found to have germinated on the surface of the Podisoma whence they
originated. The germ filament which they produce springs habitually from
the side, at a short distance from the hilum, which indicates the point of
attachment to the original spicule. These filaments will attain to from fifteen
to twenty times the diameter of the spore in length before branching, and
are in themselves exceedingly delicate. The tubes which issue from the
primary spores (protospores, Tul.) are not always simple, but sometimes
forked; and the cells which are ultimately formed at their extremities,
though producing filiform processes, do not always generate secondary
spores (teleutospores) at their apices. This mode of germination, it will be
seen, resembles greatly that which takes place in Puccinia.

The germination of the Ustilagines
was in part examined by Tulasne, but
since has received accessions through
the labours of Dr. A. Fischer von
Waldheim.[I] Nothing, however, of any
importance is added to our knowledge of
the germination of Tilletia, which was
made known as early as 1847.[J] After
some days a little obtuse tube is
protruded through the epispore, bearing
at its apex long fusiform bodies, which
are the sporules of the first generation.
These conjugate by means of short
transverse tubes, after the manner of the
Fig. 89.—Germinating pseudospore (g) of threads of Zygnema. Afterwards long
Tilletia caries with secondary spores in elliptical sporules of the second
conjugation. (Tul.)
generation are produced on short
pedicels by the conjugated fusiform bodies of the first generation. (Fig. 89,
ss.) Ultimately these sporules of the second generation germinate, and
generate, on short spicules, similar sporules of a third generation. (Fig. 89,
st.)
In Ustilago (flosculorum) germination takes
place readily in warm weather. The germ tube is
rather smaller at its base than further on. In from
fifteen to eighteen hours the contents become
coarsely granular; at the same time little
projections appear on the tube which are
narrowed at the base, into which some of the
protoplasm passes. These ultimately mature into
Fig. 90.—Pseudospore of
sporules. At the same time a terminal sporule Ustilago receptaculorum in
generally appears on the threads. Secondary germination, and secondary
sporules frequently grow from the primary, which spores in conjugation. (Tul.)
are rather smaller, and these occasionally give rise to a third generation.
In Urocystis (pompholygodes) the germinating tubes spring exclusively
from the darker central cells of the clusters. From these are developed at
their extremity three or four linear bodies, as in Tilletia, but after this no

further development has as yet been traced. It may be remarked here that
Waldheim observed similar conjugation of the sporules in some species of
Ustilago as have been remarked in the sporules of the first generation in
Tilletia.
Returning to Cystopus, as the last of the
Uredines, we must briefly recapitulate the
observations made by Professor de Bary,
[K] who, by the bye, claims for them an
affinity with Peronospora (Mucedines but
too well known in connection with the
potato disease), and not with the Uredines
and their allies. In this genus there are two
kinds of reproductive organs, those
Fig. 91.—Conidia and zoospores of produced on the surface of the plant
Cystopus candidus; a. conidium with the
plasma divided; b. zoospores escaping; c. bursting through the cuticle in white
zoospores escaped from the conidium; d. pustules, and which De Bary terms
active zoospores; e. zoospores, having lost conidia, which are generated in chains,
their cilia, commencing to germinate.
and certain globose bodies termed
oogonia, which are developed on the mycelium in the internal tissues of the
foster plant. When the conidia are sown on water they rapidly absorb the
moisture, and swell; the centre of one of the extremities soon becomes a
large obtuse papilla resembling the neck of a bottle. This is filled with a
granular protoplasm, in which vacuoles are formed. Soon, however, these
vacuoles disappear, and very fine lines of demarcation separate the
protoplasm into from five to eight polyhedric portions, each presenting a
little faintly-coloured vacuole in the centre (a). Soon after this division the
papilla at the extremity swells, opens itself, and at the same time the five to
eight bodies which had formed in the interior are expelled one by one (b).
These are zoospores, which at first take a lenticular form, and group
themselves before the mouth of the parent cell in a globose mass (c.) Very
soon, however, they begin to move, and then vibratile cilia show themselves
(d), and by means of these appendages the entire globule moves in an
oscillating manner as one by one the zoospores disengage themselves, each
becoming isolated and swimming freely in the surrounding fluid. The
movement is precisely that of the zoospores of Algæ.

The generation of the zoospores commences
within from an hour and a half to three hours
after the sowing of the conidia on water. From the
oogonia, or resting spores, similar zoospores, but
in greater number, are generated in the same
manner, and their conduct after becoming free is
identical. Their movements in the water usually
last from two to three hours, then they abate, the
cilia disappear, and the spore becomes
immovable, takes a globose form, and covers
itself with a membrane of cellulose. Afterwards Fig. 92.—Resting spore of
the spore emits, from any point whatever of its Cystopus candidus with
zoospores escaped.
surface, a thin, straight or flexuous tube, which
attains a length of from two to ten times the diameter of the spore. The
extremity becomes clavate or swollen, after the manner of a vesicle, which
receives by degrees the whole of the protoplasm.
De Bary then proceeds to describe experiments which he had performed
by watering growing plants with these zoospores, the result being that the
germinating tubes did not penetrate the epidermis, but entered by the
stomates, and there put forth an abundant mycelium which traversed the
intercellular passages. Altogether the germination of these conidia or
zoospores offers so many differences from the ordinary germination of the
Uredines, and is so like that which prevails in Peronospora, in addition to
the fact of both genera producing winter spores or oogonia, that we cannot
feel surprised that the learned mycologist who made these observations
should claim for Cystopus an affinity with Peronospora rather than with the
plants so long associated with it amongst the Coniomycetes.
In passing from these to the Mucedines, therefore, we cannot do so more
naturally than by means of that genus of white moulds to which we have
just alluded. The erect branched threads bear at the tip of their branchlets
spores, or conidia, which conduct themselves in a like manner to the organs
so named in Cystopus, and oogonia or resting spores developed on the
mycelium within the tissues of the foster plant also give origin to similar
zoospores.
The conidia are borne upon erect, elongated filaments, originating from
the creeping mycelium. These threads are hollow, and rarely septate; the

upper portion divided into numerous branches, and these again are
subdivided, the ultimate ramuli each terminated by a single conidium. This
body when mature is oval or elliptical, filled with protoplasm, but there is a
diversity in their mode of germination. In the greater part, of which P. effusa
may be taken as an example, the conidia have the function of simple spores.
Placed in favourable conditions, each of them puts forth a germ-tube, the
formation of which does not differ in any essential point from what is
known of the spores of the greater part of fungi.
The short oval conidia of P. gangliformis have little obtuse papillæ at
their apex, and it is at this point that germination commences.
The conidia of P. densa are similar, but the germination is different.
When placed in a drop of water, under favourable circumstances, the
following changes may be observed in from four to six hours. The
protoplasm, at first uniformly distributed in all the conidia, appears strewn
with semi-lenticular, and nearly equidistant vacuoles, of which the plane
face is immediately in contact with the periphery of the protoplasm. These
vacuoles number from sixteen to eighteen in P. macrocarpa, but are less
numerous in P. densa. A short time after the appearance of the vacuoles the
entire conidium extends itself so that the papilla disappears. Suddenly it
reappears, elongates itself, its attenuated membrane vanishes, and the
protoplasm is expelled by the narrow opening that remains in place of the
papilla. In normal cases the protoplasm remains united in a single mass that
shows a clear but very delicate outline. When it has reached the front of the
opening in the conidium, which is thus emptied, the mass remains
immovable. In P. densa it is at first of a very irregular form, but assumes by
degrees a regular globose shape. This is deprived of a distinct membrane,
the vacuoles that disappeared in the expulsion again become visible, but
soon disappear for a second time. The globule becomes surrounded with a
membrane of cellulose, and soon puts out from the point opposite to the
opening of the conidium a thick tube which grows in the same manner as
the germ-tube of the conidia in other species. Sometimes the expulsion of
the protoplasm is not completely accomplished; a portion of it remaining in
the membrane of the conidium detaches itself from the expelled portion,
and while this is undergoing changes takes the form of a vesicle, which is
destroyed with the membrane. It is very rare that the protoplasm is not
evacuated, and that the conidia give out terminal or lateral tubes in the

manner that is normal to other species without papillæ. The germination just
described does not take place unless the conidia are entirely surrounded by
water; it is not sufficient that they repose upon its surface. Besides, there is
another condition which, without being indispensable, has a sensible
influence on the germination of P. macrocarpa, and that is the exclusion of
light. To ascertain if the light or the darkness had any influence, two equal
sowings were placed side by side, the one under a clear glass bell, the other
under a blackened glass bell. Repeated many times, these experiments
always gave the same result—germination in from four to six hours in the
conidia under the blackened glass; no change in those under the clear glass
up to the evening. In the morning germination was completed.
The conidia of P. umbelliferarum and P. infestans[L] show an analogous
structure. These bodies, if their development be normal, become
zoosporangia. When they are sown upon water, one sees at the end of some
hours the protoplasm divided by very fine lines, and each of the parts
furnished with a small central vacuole. Then the papilla of the conidium
disappears. In its place appears a rounded opening, by which the parts of the
protoplasm are expelled rapidly, one after the other. Each of these, when
free, immediately takes the form of a perfect zoospore, and commences to
agitate itself. In a few moments the sporangium is empty and the spores
disappear from the field of the microscope.
The zoospores are oval or semi-oval, and in P. infestans the two cilia
spring from the same point on the inferior border of the vacuole. Their
number in a sporangium are from six to sixteen in P. infestans, and from six
to fourteen in P. umbelliferarum. The movement of the zoospores ceases at
the end of from fifteen to thirty minutes. They become motionless, cover
themselves with a membrane of cellulose, and push out slender bent germ-
tubes which are rarely branched. It is but seldom that two tubes proceed
from the same spore. The same development of the zoospores in P. infestans
is favoured by the exclusion of the light. Placed in a position moderately
lighted or protected by a blackened bell, the conidia very readily produced
zoospores.
A second form of germination of the conidia in P. infestans, when sown
upon a humid body or on the surface of a drop of water, consists in the
conidium emitting from its summit a simple tube, the extremity of which
swells itself into the form of an oval vesicle, drawing to itself, little by little,

all the protoplasm contained in the conidium. Then it isolates itself from the
germ-tube by a septum, and takes all the essential characteristics of the
parent conidium. This secondary conidium can sometimes engender a third
cellule by a similar process. These secondary and tertiary productions have
equally the character of sporangia. When they are plunged into water, the
ordinary production of zoospores takes place.
Lastly, there is a third mode of germination which the conidia of P.
infestans manifest, and which consists in the conidium emitting from its
summit a simple or branched germ-tube. This grows in a similar manner to
the conidia first named as of such species as P. effusa. The conditions which
control this form of germination cannot be indicated, since some conidia
which germinate after this manner will sometimes be found mixed with
others, the majority of which furnish zoospores. It may be that the conidia
themselves are in some sort of abnormal condition.
In all the species examined the conidia possess the power of germination
from the moment of their maturity. The younger they are the more freely
they germinate. They can retain this power for some days or weeks,
provided they are not entirely dried. Dessication in an ordinary temperature
seemed sufficient to destroy the faculty of germinating in twenty-four
hours, when the conidia had been removed from the leaves on which they
were produced. They none of them retained the faculty during a few
months, hence they cannot preserve it during the winter.
The germs of Peronospora enter the foster plant if the spores are sown
upon a part suitable for the development of the parasite. It is easy to
convince one’s self that the mycelium, springing from the penetrating
germs, soon takes all the characters that are found in the adult state.
Besides, when cultivated for some time, conidiiphorous branches can be
seen growing, identical with those to which it owes its origin. Such
cultivation is so readily accomplished that it can be made upon cut leaves
preserved fresh in a moist atmosphere.
In the species of Peronospora that inhabit perennial plants, or annual
plants that last through the winter, the mycelium hidden in the tissues of the
foster-plant lasts with it. In the spring it recommences vegetation, and emits
its branches into the newly-formed organs of its host, there to fructify. The
Peronospora of the potato is thus perennial by means of its mycelium

contained in the browned tissue of the diseased tubers. When in the spring a
diseased potato begins to grow, the mycelium rises in the stalk, and soon
betrays itself by blackish spots. The parasites can fructify abundantly on
these little stalks, and in consequence propagate themselves in the new
season by the conidia coming from the vivacious mycelium.
The diseased tubers of the potato always contain the mycelium of P.
infestans, which never fructifies there as long as the skin of the tuber is
intact. But when, in cutting the tuber, the parenchyma occupied by the
mycelium is exposed to the contact of the air, it covers itself with conidia-
bearing branches at the end of from twenty-four to forty-eight hours.
Analogous results are obtained with the stalks of the potato. It is evident
that in these experiments nothing is changed except the contact of the air;
the specific conditions particularly remain the same. It appears, therefore,
that it is this contact alone which determines generally the production of the
conidiiferous branches.[M]
The mode of germination and development in the Mucors has been
studied by several observers, but most recently by Van Tieghem and Le
Monnier.[N] In one of the common forms, the Mucor phycomyces of some
authors, and the Phycomyces nitens of others, the process is given in detail.
In this species germination will not take place in ordinary water, but it
readily takes place in orange juice and other media. The spore loses colour,
swells, and absorbs fluid around it until double its original size and ovoid.
Then a thick thread is emitted from one or both extremities, which
elongates and becomes branched in a pinnate manner. Sometimes the
exospore is ruptured and detached loosely from the germinating spore. After
about forty-eight hours from the first sowing, the mycelium will send
branches into the air, which again become abundantly branched; other short
submerged branches will also remain simple, or have tuft-like ramifications,
each terminating in a point, so as to bristle with spiny hairs. In two or three
days abruptly swollen branches, of a club shape, will make their appearance
on the threads both in the air and in the fluid. Sometimes these branches are
prolonged into an equal number of sporangia-bearing threads, but most
frequently they divide first at their swollen summits into numerous
branches, of which usually one, sometimes two or three, develop into
sporangia-bearing threads, while the rest are short, pointed, and form a tuft

of rootlets. Sometimes these rootlets reduce themselves to one or more
rounded protuberances towards the base of the sporangia-bearing threads.

Fig. 93.—Zygospores of Mucor phycomyces.
(Van Tieghem.)

There are often also a certain number of the branches which had acquired
a clavate shape, and do not erect themselves above the surface, instead of
producing a fertile thread, which would seem to have been their first
intention, become abruptly attenuated, and are merely prolonged into a
mycelial filament. Although in other species chlamydospores are formed in
such places on the mycelium, nothing of the kind has been traced in this
species, more than here indicated. Occasionally, when germination is
arrested prematurely, certain portions of the hyphæ, in which the
protoplasm maintains its vitality, become partitioned off. This may be
interpreted as a tendency towards the formation of chlamydospores, but
there is no condensation of protoplasm, or investiture with a special
membrane. Later on this isolated protoplasm is gradually altered, separating
into somewhat regular ovoid or fusiform granules, which have, to a certain
extent, the appearance of spores in an ascus, but they seem to be incapable
of germination.
Another method of reproduction, not uncommon in Mucorini, is
described by Van Tieghem in this species. Conjugating threads on the

substratum by degrees elaborate zygospores, but these, contrary to the mode
in other species, are surrounded by curious branched processes which
emanate from the arcuate cells on either side of the newly-developed
zygospore. This system of reproduction is again noticed more in detail in
the chapter on polymorphism.
M. de Seynes has given the details of his examination of the sporidia of
Morchella esculenta during germination.[O] A number of these sporidia,
placed in water in the morning, presented, at nine o’clock of the same
evening, a sprout from one of the extremities, measuring half the length of
the spore. In the morning of the next day this sprout had augmented, and
become a filament three or four times as long. The next day these elongated
filaments exhibited some transverse divisions and some ramifications. On
the third day, the germination being more advanced, many more of the
sporidia were as completely changed, and presented, in consequence of the
elongation, the appearance of a cylindrical ruffle, the cellular prolongations
arising from the germination having a tendency towards one of the
extremities of the longer axis of the sporidium, and more often to the two
opposed extremities, either simultaneously or successively. Out of many
hundreds of sporidia examined during germination, he had only seen a very
few exceptions to this rule, among which he had encountered the
centrifugal tendency to vegetate by two opposed filaments, proving that if it
bears a second by the side of the primal filament situated at one of the
poles, a second would also be seen from the side of the filament coming
from the opposite pole.
Before being submitted to the action of water, the contents of the sporidia
seemed formed of two distinct parts, one big drop of yellow oil of the same
form as the sporidium, with the space between it and the cell wall occupied
by a clear liquid, more fluid and less refractive, nearly colourless, or at
times slightly roseate. As the membrane absorbed the water by which it was
surrounded, the quantity of this clear liquid was augmented, and the rosy
tint could be more easily distinguished. All the contents of the spore, which
up to this time remained divided into two parts, presented altogether one
aspect, only containing numerous granulations, nearly of equal size,
completely filling it, and reaching the inner face of the sporic membrane.
After this time the sporidium augments in size very rapidly, becoming at
times irregular, and sometimes even as much as from two to three times its

original dimensions, then there appears at the surface, usually at one of the
poles of the ellipse, a small prominence, with an extremely fine membrane,
which does not appear to separate itself from that which surrounds the
sporidium, and it is difficult to say whether it is a prolongation of the
internal membrane going across the outside, or simply a prolongation
caused by a continuation of tissue of an unique membrane. Sometimes there
may be seen at the point where the primal filament issues from the
sporidium a circular mark, which appears to indicate the rupture of the
external membrane. From this time another change comes over the
contents. We again find the yellow oily liquid, now occupying the external
position, with some drops of colourless or roseate liquid in the centre, so
that the oily liquid and the more limpid fluid interchange the positions
which they occupied previous to the commencement of germination.
Whether these two fluids have undergone any change in their constitution is
difficult to determine, at all events the oily liquid appears to be less
refractive and more granular, and it may be that it is a product of new
formation, containing some of the elements of the primitive oily drop.
Having regard to the delicate character of the membrane of the germinating
filaments, De Seynes supposed that it might offer greater facility for the
entrance of water by endosmose, and account for the rapid enlargement of
the sporidia. By a series of experiments he became satisfied that this was
the case to a considerable extent, but he adds:—“I cannot help supposing
that a greater absorption of greasy matter in the cell which is the first
product of germination raises an objection to an aqueous endosmose. One
can also see in this experience a proof of the existence of two special
membranes, and so suppose that the germinative cell is the continuation of
the internal membrane, the external membrane alone being susceptible of
absorbing the liquids, at least with a certain rapidity.”
In other Discomycetes germination takes place in a similar manner.
Boudier[P] narrates that in Ascobolus, when once the spore reaches a
favourable place, if the circumstances are good, i.e., if the temperature is
sufficiently high and the moisture sufficient, it will germinate. The time
necessary for this purpose is variable, some hours sufficing for some
species; those of A. viridis, for example, germinate in eight or ten hours,
doubtless because, being terrestrial, it has in consequence less heat. The
spore slightly augments in size, then opens, generally at one or other
extremity, sometimes at two, or at any point on its surface, in order to pass

the mycelium tubes. At first simple, without
septa, and granular in the interior, above all at the
extremity, these tubes, the rudiment of the
mycelium, are not long in elongating, in
branching, and later in having partitions. These
filaments are always colourless, only the spore
may be coloured, or not. Coemans has described
them as giving rise to two kinds of conidia,[Q] the
one having the form of Torula, when they give
rise to continuous filaments, the other in the form
of Penicillium, when they give birth to
partitioned filaments. De Seynes could never
obtain this result. Many times he had seen the
Penicillium glaucum invade his sowings, but he
feels confident that it had nothing to do with the Fig. 94.—Sporidium of
Ascobolus. M. Woronin[R] has detailed some Ascobolus germinating.
observations on the sexual phenomena which he has observed in Ascobolus
and Peziza, and so far as the scolecite is concerned these have been
confirmed by M. Boudier.
There is no reason for doubt that in other of the Discomycetes the
germination of the sporidia is very similar to that already seen and
described, whilst in the Pyrenomycetes, as far as we are aware, although the
production of germinating tubes is by no means difficult, development has
not been traced beyond this stage.[S]
[A]
Seynes, J. de, “Essai d’une Flore Mycologique de la Montpellier,” &c. (1863), p. 30.
[B]
Hoffman, “Icones Analyticæ Fungorum.”
[C]
The spores of Agarics which are devoured by flies, however, though returned in their
dung in an apparently perfect state, are quite effete. It is, we believe, principally by
the Syrphidæ, which devour pollen, that fungus spores are consumed.
[D]
All attempts at Chiswick failed with some of the more esculent species, and Mr.
Ingram at Belvoir, and the late Mr. Henderson at Milton, were unsuccessful with
native and imported spawn.

[E]

Tulasne, “On the Organization of the Tremellini,” “Ann. des. Sci. Nat.” 3me sér. xix.
(1853), p. 193.
[F]
Tulasne, “Mémoire sur les Urédinées.”
[G]
Tulasne, in his “Memoirs on the Uredines.”
[H]
Mr. Berkeley has lately published a species under the name of P. Ellisii, in which the
gelatinous element is scarcely discernible till the plant is moistened. There are two
septa in this species, and another species or form has lately been received from Mr.
Ellis which has much shorter pedicels, and resembles more closely Puccinia, from
which it is chiefly distinguished by its revivescent character.
[I]
Von Waldheim, on the “Development of the Ustilagineæ,” in “Pringsheim’s
Jahrbucher,” vol. vii. (1869); translated in “Transactions of N. Y. State Agricultural
Society for 1870.”
[J]
Berkeley, on the “Propagation of Bunt,” in “Trans. Hort. Soc. London,” ii. (1847), p.
113; Tulasne, second memoir, in “Ann. des. Sci. Nat.” ii. (4me sér.), p. 77; Cooke, in
“Journ. Quekett Micro. Club,” i. p. 170.
[K]

De Bary, “Recherches,” &c. in “Annales des Sciences Naturelles” (4me sér.), xx. p. 5;
Cooke in “Pop. Sci. Rev.” iii. (1864), p. 459.
[L]
This is the mould which produces the potato murrain.
[M]

De Bary, “Champignons parasitiques,” in “Annales des Sci. Nat.” (4me sér.), xx. p. 5;
Cooke, “Microscopic Fungi,” cap. xi. p. 138; “Popular Science Review,” iii. 193
(1864).
[N]
Van Tieghem and Le Monnier, “Researches on Mucorini,” in “Ann. des Sci. Nat.”
(1873), xvii. p. 261; Summary in “Quart. Journ. Micro. Science” (2nd ser.), xiv. p. 49.
[O]
Seynes, “Essai d’une Flore Mycologique.”

[P]
Boudier, “Mémoire sur l’Ascoboles,” pt. i. iv. f. 13–15.
[Q]
Coemans, “Spicilége Mycologique,” i. p. 6.
[R]
Woronin, “Abhandlungen der Senchenbergischen Naturfor. Gesellschaft” (1865), p.
333.
[S]
In the very important observations made by Dr. Cunningham at Calcutta, on
substances floating in the atmosphere, it appeared that the sporidia of many Sphæriæ
actually germinated after being taken up by the air. The multitude of fungus spores
which were observed in every case was quite extraordinary.

VIII.
SEXUAL REPRODUCTION.

The existence of some sort of sexual reproduction in Fungi has long been
suspected, although in earlier instances upon insufficient grounds; but of
late years observations have multiplied and facts accumulated which leave
no doubt of its existence. If the Saprolegniæ are left out of the question as
disputed Fungi, there still remain a number of well authenticated instances
of the phenomena of copulation, and many other facts which indicate some
sort of sexual relationship. The precise manner in which those minute
bodies, so common amongst the Sphæronemei, which we prefer to call
stylospores, perform their functions is still to a great extent a mystery; yet it
is no longer doubted that certain species of Aposphæria, Phoma, Septoria,
&c., are only conditions of some species of Sphæria, often developed and
matured in close proximity to them on the same host. In Æcidium, Rœstelia,
&c., spermogonia are produced plentifully on or near the same spots on
which the fructification appears, either simultaneously or at a later period.
[A] The relation of Cytispora to Valsa was suspected by Fries very many
years ago, and, as since demonstrated, with very good reason. All attempts,
however, to establish anything like sexual reproduction in the higher forms
of Hymenomycetes have at present been unsuccessful; and the same may be
said of the Gasteromycetes; but in Ascomycetes and Physomycetes instances
abound.
We know not whether any importance is to be attached to the views of M.
A. S. Œrsted,[B] which have not since been confirmed, but which have been
cited with some approval by Professor de Bary, as to a trace of sexual
organs in Hymenomycetes. He is supposed to have seen in Agaricus
variabilis, P., oocysts or elongated reniform cells, which spring up like
rudimentary branches of the filaments of the mycelium, and enclose an
abundant protoplasm, if not even a nucleus. At the base of these oocysts
appear the presumed antheridia, that is to say, one or two slender filaments,

which generally turn their extremities towards the oocysts, and which more
rarely are applied to them. Then, without ulteriorily undergoing any
appreciable modifications, the fertile cell or oocyst becomes enveloped in a
network of filaments of mycelium which proceed from the one which bears
it, and this tissue forms the rudiments of the cap. The reality of some kind
of fecundation in this circumstance, and the mode of the phenomena, if
there is one, are for the present equally uncertain. If M. Œrsted’s opinion is
confirmed, naturally the whole of the cap will be the product of
fecundation. Probably Karsten (Bonplandia, 1862, p. 62) saw something
similar in Agaricus campestris, but his account is obscure.

Fig. 95.—Zygospore of Mucor phycomyces.

In Phycomyces the organs of reproduction have been subjected to close
examination by Van Tieghem,[C] and although he failed to discover
chlamydospores in this, he describes them in other Mucors. In this species,
besides the regular sexual development, by means of sporangia, there is a
so-called sexual reproduction by means of zygospores, which takes place in
this wise. The threads which conjugate to form the zygospores are slender
and erect on the surface of the substratum. Two of these threads come into
close contact through a considerable length, and clasp each other by
alternate protuberances and depressions. Some of the protuberances are
prolonged into slender tubes. At the same time the free extremities of the

threads dilate, and arch over one towards the other until their tops touch like
a vice, each limb of which rapidly increases in size. Each of these arcuate,
clavate cells has now a portion of its extremity isolated by a partition, by
means of which a new hemispherical cell is formed at the end of each
thread at its point of junction with the opposed thread. These cells become
afterwards cylindrical by pressure, the protoplasm is aggregated into a
mass, the double membrane at the point of first contact is absorbed, and the
two confluent masses of protoplasm form a zygospore invested with a
tubercular coat and enveloped by the primary wall of the two conjugating
cells. During this formation of the zygospore, the two arched cells whence
the zygospore originated develop a series of dichotomous processes in close
proximity to the walls which separate them from the zygospore. These
processes appear at first on one of the arcuate cells in successive order. The
first makes its appearance above upon the convex side; the succeeding ones
to the right and left in descending order; the last is in the concavity beneath.
It is only after the development of this that the first process appears on the
opposite cell, which is followed by others in the same order. These
dichotomous processes are nothing more than branches developed from the
arcuate, or mother cells. During all these changes, while the zygospore
enlarges, the wall of the arcuate cells becomes coloured brown. This
colouring is more marked on the convex side, and it shows itself first in the
cell on which the dichotomous branches are first produced, and which
retains the darker tint longer than the other. The zone from whence the
processes issue, and also the processes themselves, have their walls
blackened deeply, while the walls of the conjugated cells, which continue to
clothe the zygospore during the whole of its development, are bluish-black.
By pressure, the thin brittle coat which envelopes the zygospore is ruptured,
and the coat of the zygospore exposed, formed of a thick cartilaginous
membrane, studded with large irregular warts.
The germination of the zygospores in this species has not as yet been
observed, but it is probably the same or very similar to that observed in
other species of Mucor. In these the rough tuberculate epispore splits on one
side, and its internal coat elongates itself and protrudes as a tube filled with
protoplasm and oil globules, terminating in an ordinary sporangium.
Usually the amount of nutriment contained in the zygospore is exhausted by
the formation of the terminal sporangium, according to Brefeld;[D] but Van
Tieghem and Le Monnier remark that in their examinations they have often

seen a partition formed at about a third of the length of the principal
filament from the base, below which a strong branch is given off, and this is
also terminated by a large sporangium.

Fig. 96.—Zygospore of Rhizopus in different stages. (De Bary.)

De Bary has given a precise account of the formation of the zygospore in
another of the Mucors, Rhizopus nigricans, in which he says that the
filaments which conjugate are solid rampant tubes, which are branched
without order and confusedly intermingled. Where two of these filaments
meet each of them pushes towards the other an appendage which is at first
cylindrical and of the same diameter. From the first these two processes are
applied firmly one to the other by their extremities; they increase in size,
become clavate, and constitute together a fusiform body placed across the
two conjugated filaments. Between the two halves of this body there exists
no constant difference of size; often they are both perfectly equal. In each
there is collected an abundance of protoplasm, and when they have attained
a certain development the largest extremity of each is isolated by a septum
from the clavule, which thus becomes the support or suspender of the
copulative cell. The two conjugated cells of the fusiform body are generally
unequal; the one is a cylinder as long as it is broad, the other is disciform,
and its length is only equal to half its breadth. The primitive membrane of
the clavule forms between the copulative cells a solid partition of two
membranes, but soon after the cells have become defined the medial
partition becomes pierced in the centre, and then soon entirely disappears,
so that the two twin cells are confounded in one single zygospore, which is

due to the union of two more or less similar utricles. After its formation the
zygospore still increases considerably in size, and acquires a diameter of
more than one-fifth of a millimetre. Its form is generally spherical, and
flattened on the faces which are united to the suspenders, or it resembles a
slightly elongated cask. The membrane thickens considerably, and consists
at the time of maturity of two superposed integuments; the exterior or
epispore is solid, of a dark blackish-blue colour, smooth on the plane faces
in contact with the suspenders, but covered everywhere else with thick
warts, which are hollow beneath. The endospore is thick and composed of
several layers, colourless, and covered with warts, which correspond and fit
into those of the epispore. The contents of the zygospore are a coarsely
granular protoplasm, in which float large oleaginous drops. While the
zygospore is increasing in size, the suspender of the smaller copulative cell
becomes a rounded and stipitate utricle, often divided at the base by a
septum, and which attains almost to the size of the zygospore. The
suspender of the larger copulative cell preserves its primitive form and
becomes scarcely any larger. It is rare that there is not a considerable
difference of size between the two conjugated cells and the suspenders.[E]
Similar conjugation with like results also takes place in Syzygites
megalocarpus. In this species the germination of the zygospores has been
observed. If, after a certain time of repose, these bodies are placed on a
moist substratum, they emit a germ-like tube, which, without originating a
proper mycelium, develops at the expense of the nutritive material stored in
the zygospore into a carpophore or fruit bearer, which is many times
dichotomously branched, bearing terminal sporangia characteristic of the
species.
It has already been remarked by us that the Saprolegnei are claimed by
some authors as Algæ, whilst we are more disposed to regard them as
closely allied to the Mucors, and as they exhibit in themselves strong
evidence in support of the existence of sexual reproduction, we cannot
forbear giving a summary of what has been observed by De Bary and others
in this very interesting and singular group of plants, to which M. Cornu has
recently dedicated an exhaustive monograph.[F]
In Saprolegnia monoica, and others, the female organs consist of oogonia
—that is to say, of cells which are at first globose and rich in plastic matter,
which most generally terminate short branches of the mycelium, and which

are rarely seen in an interstitial position. The constitutive membrane of the
adult oogonia is reabsorbed in a great many points, and is there pierced with
rounded holes. At the same time the plasma is divided into a larger or
smaller number of distinct portions, which are rounded into little spheres,
and separate from the walls of the conceptacle in order to group themselves
at the centre, where they float in a watery fluid. These gonospheres are then
smooth and bare, with no membrane on their surface of the nature of
cellulose.
During the formation of the oogonia
there arise from its pedicel or from
neighbouring filaments slight cylindrical
curved branches, sometimes turned round
the support of the oogonia, and which all
tend towards this organ. Their superior
extremity is intimately applied to its wall,
then ceases to be elongated, becomes
slightly inflated, and is limited below by a
partition; it is then an oblong cell, slightly
curved, filled with protoplasm, and
intimately applied to the oogonia—in fact,
an antheridium or organ of the male sex.
Each oogonium possesses one or several Fig. 97.—Conjugation in Achlya
racemosa. (Cornu.)
antheridia. Towards the time when the
gonospheres are formed it may be observed that each antheridium sends to
the interior of the oogonia one or several tubular processes, which have
crossed its side wall, and which open at their extremity in order to discharge
their contents. These, while they are flowing out, present some very agile
corpuscles, and which, considering their resemblance to those in Vaucheria,
to which the name of spermatozoids are applied, ought to be considered as
the fecundating corpuscles. After the evacuation of the antheridia the
gonospheres are found to be covered with cellulose; they then constitute so
many oospores, with solid walls. De Bary considers that, bearing in mind
analogous phenomena observed in Vaucheria, and the direct observations of
Pringsheim,[G] the cellulose membrane on the surface of the gonospheres is
only the consequence of a sexual fecundation.

In Achlya dioica the antheridium is cylindrical, the plasma which it
encloses is divided into particles, which attain nearly the size of the
zoospores of the same plant. These particles become globose cells, grouped
in the centre of the antheridium. Afterwards the contents of these latter cells
become divided into numerous bacillary spermatozoids, which first break
the wall of their mother cell, and then issue from the antheridium. These
rod-like corpuscles, which resemble the spermatozoids in Vaucheria, have
their movements assisted by a long cilium. It is presumable that here, as in
the Algæ, the spermatozoids introduce themselves into the cavity of the
oogonium, and unite with the gonospheres.
Amongst obscure and doubtful bodies are those described by Pringsheim,
which have their origin in thick filaments or tubes, similar to those which
form the zoosporangia, and represent so many distinct little masses of
plasma within an homogeneous parietal ganglion. The contour of these
plastic masses is soon delineated in a more precise manner. We see in their
interior some homogeneous granules, which are at first globose, then oval,
and finally travel to the enlarged and ampullæform extremity of the
generating tube. There they become rounded or oval cells covered with
cellulose, and emit from their surface one or several cylindrical processes,
which elongate towards the wall of the conceptacle, and pierce it, without,
however, ever projecting very far beyond it. At the same time the lacunose
protoplasm of each cell becomes divided into a number of corpuscles,
which escape by the open extremity of the cylindrical neck. They resemble
in their organization and agility the spermatozoids of Achlya dioica. They
soon become motionless in water, and do not germinate. During the
development of these organs, the protoplasm of the utricle which contains
them offers at first completely normal characteristics, and disappears
entirely by degrees as they increase. De Bary and Pringsheim believe that
these organs constitute the antheridia of the species of Saprolegnia to which
they belong.
The oospores of the Saprolegniæ, when arrived at maturity, possess a
tolerably thick double integument, consisting of an epispore and an
endospore. After a considerable time of repose they give rise to tubular or
vesicular germs, which, without being much elongated, produce zoospores.
[H]

De Bary has claimed for the oogonia in Cystopus and Peronospora a kind
of fecundation which deserves mention here.[I] These same fruits, he says,
which owe their origin to sexual organs, should bear the names of oogonia
and antheridia, according to the terminology proposed by Pringsheim for
analogous organs in the Algæ. The formation of the oogonia, or female
organs, commences by the terminal or interstitial swelling of the tubes of
the mycelium, which increase and take the form of large spherical or oboval
cells, and which separate themselves by septa from the tube which carries
them. Their membrane encloses granules of opaque protoplasm, mingled
with numerous bulky granules of colourless fatty matter.
The branches of the mycelium which do not
bear oogonia apply their obtuse extremities
against the growing oogonia; this extremity
swells, and, by a transverse partition, separates
itself from the supporting tube. It is the
antheridium, or male organ, which is formed by
this process; it takes the form of an obliquely
clavate or obovate cellule, which is always
considerably smaller than the oogonium, and Fig. 98.—Conjugation in
Peronospora; a. antheridium.
adheres to its walls by a plane or convex area. (De Bary.)
The slightly thickened membrane of the
antheridia encloses protoplasm which is finely granular. It is seldom that
more than one antheridium applies itself to an oogonium.
The two organs having together achieved their development, the large
granules contained in the oogonium accumulate at its centre to group
themselves under the form of an irregular globule deprived of a proper
membrane, and surrounded by a bed of almost homogeneous protoplasm.
This globule is the gonosphere, or reproductive sphere, which, through the
means of fecundation, should become the reproductive body, vegetable egg,
or oospore. The gonosphere having been formed, the antheridium shoots out
from the centre of its face, close against the oogonium, a straight tube,
which perforates the walls of the female cell, and traversing the protoplasm
of its periphery, directs itself to the gonosphere. It ceases to elongate itself
as soon as it touches it, and the gonosphere becomes clothed with a
membrane of cellulose, and takes a regular spheroidal form.

Considering the great resemblance of these
organs with the sexual organs of the Saprolegniæ,
which are closely allied to the Algæ, and of
which the sexuality has been proved, De Bary
adds, we have no doubt whatever that the
phenomena just described represent an act of
fecundation, and that the tube pushed out by the Fig. 99.—Antheridia and
antheridium should be regarded as a fecundating oogonium of Peronospora. (De
tube. It is remarkable that amongst these fungi the Bary.)
tube projected by the antheridium effects fecundation only by contact. Its
extremity never opens, and we never find antherozoids; on the contrary, the
antheridium presents, up to the maturity of the oospore, the appearance
which it presented at the moment of fecundation.
The primitive membrane of the oospore, at first very thin, soon acquires a
more sensible thickness, and becomes surrounded by an external layer
(epospore), which is formed at the expense of the protoplasm of the
periphery. This disappears in proportion as the epispore attains maturity,
and finally there only remains a quantity of granules, suspended in a
transparent watery fluid. At the period of maturity, the epispore is a slightly
thickened, resistant membrane, of a yellowish-brown colour, and finely
punctate. The surface is almost always provided with brownish warts,
which are large and obtuse, sometimes isolated, and sometimes confluent,
forming irregular crests. These warts are composed of cellulose, which
reagents colour of a deep blue, whilst the membrane which bears them
preserves its primitive colour. One of the warts, larger than the rest, and
recognizable by its cylindrical form, always forms a kind of thick sheath
around the fecundating tube. The ripe endospore is a thick, smooth,
colourless membrane, composed of cellulose containing a bed of finely
granulated protoplasm, which surrounds a great central vacuole. This
oospore, or resting spore, may remain dormant in this state within the
tissues of the foster plant for some months. Its ultimate development by
production of zoospores is similar to the production of zoospores from
conidia, which it is unnecessary to repeat here. The oospore becomes an
oosporangium, and from it at least a hundred germinating bodies are at
length expelled.

Amongst the principal observers of certain phenomena of copulation in
cells formed in the earliest stages of the Discomycetes are Professor de
Bary,[J] Dr. Woronin,[K] and Messrs. Tulasne.[L] In the Ascobolus
pulcherrimus of Crouan, Woronin 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 into a series of
cells, the successive increase of which finally gives to the whole a torulose
and unequal appearance. The body thus formed he calls a “vermiform
body.” The same observer also seems to have convinced himself that there
exists always in proximity to this body certain filaments, the short arched or
inflected branches of which, like so many antheridia, rest their anterior
extremities on the utriform cells. This contact seems to communicate to the
vermiform body a special vital energy, which is immediately directed
towards the production of a somewhat filamentous tissue, on which the
hymenium is at a later period developed. This “vermiform body” of M.
Woronin has since come to be recognized under the name of “scolecite.”
Tulasne observes that this “scolecite” or ringed body can be readily
isolated in Ascobolus furfuraceus. When the young receptacles are still
spherical and white, and have not attained a diameter exceeding the one-
twentieth of a millimetre, 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 little sphere, and is formed of from six to eight cells, curved in
the shape of a comma.
In Peziza melanoloma, A. and S., the same observer succeeded still better
in his searches after the scolecite, which he remarks is in this species most
certainly a lateral branch of the filaments 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
arcuate 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
continuous, 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 a neighbouring
filament. In other cases the growing branch is connected, by its extremity,
with that of a hooked branch. These contacts, however, did not appear to
Tulasne to be so much normal as accidental. But of the importance of the

ringed body, or “scolecite,” there was no room for doubt, as being the
certain and habitual rudiment of the fertile cup. In fact, 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 considerable 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 of the mycelium by the base of the scolecite
which remains naked.
Although Tulasne could not satisfy himself
of the presence of any act of copulation in
Ascobolus furfuraceus, or Peziza
melanoloma, he was more successful with
Peziza omphalodes. As early as 1860 he
recognized the large globose, sessile, and
grouped vesicles which originate the fertile
tissue, but did not comprehend the part which
these macrocysts 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
Fig. 100.—Conjugation in Peziza
attenuated at the extremity. Thus provided, omphalodes. (Tulasne.)
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 filaments, are generated elongated
clavate cells, with paler contents, more vacuoles, which Tulasne names
paracysts. These, though produced after the 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 determine 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 above

described, 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 whatever. 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 filaments, which will ultimately
constitute the asci, increase and multiply.[M]
Certain phenomena concerned in
the development of the Erysiphei
belong also to this connection. The
mycelium of Erysiphe cichoracearum,
like that of other species, consists of
branched filaments, crossed in all
directions, which adhere as they climb
to the epidermis of the plant on which
the fungus lives as a parasite. The
perithecia are engendered where two
filaments cross each other. These
swell slightly at this point, and each
emits a process which imitates a Fig. 100a.—Formation of conceptacle in
nascent branch, and remains upright Erysiphe
on the surface of the epidermis. The
process originating from the inferior filament soon acquires an oval form
and a diameter double that of the filament; then it becomes isolated from it
by a septum, and constitutes a distinct cell, which De Bary[N] terms an
oocyst. The appendage which proceeds from the inferior filament always
adheres intimately to this cell, and elongates into a slender cylindrical tube,
which terminates in an obtuse manner at the summit of the same cell. At its
base it is also limited by a septum, and soon after another appears a little
below its extremity at a point indicated beforehand by a constriction. This
new septum defines a terminal short obtuse cell, the antheridium, which is
thus borne on a narrow tube like a sort of pedicel. Immediately after the

formation of the antheridia new productions show themselves, both around
the oocyst and within it. Underneath this cell eight or ten tubes are seen to
spring from the filament which bears it; these join themselves by the sides
to each other and to the pedicel of the antheridium, while they apply their
inner face to the oocyst, above which their extremities soon meet. Each of
the tubes is then divided by transverse septa into two or three distinct cells,
and in this manner the cellular walls of the perithecia come into existence.
During this time the oocyst enlarges and divides, without its being
possible precisely to determine the way in which it happens, into a central
cell and an outer layer, ordinarily simple, of smaller cells, contiguous to the
general enveloping wall. The central cell becomes the single ascus, which is
characteristic of the species, and the layer which surrounds it constitutes the
inner wall of its perithecium. The only changes afterwards observed are the
increase in size of the perithecium, the production of the root-like filaments
which proceed from its outer wall, the brown tint which it assumes, and
finally the formation of the sporidia in the ascus. The antheridium remains
for a long time recognizable without undergoing any essential modification,
but the dark colour of the perithecium soon hides it from the observer’s eye.
De Bary thinks that he is authorized in assuming the probability that the
conceptacles and organs of fructification of others of the Ascomycetes,
including the Discomycetes and the Tuberacei, are the results of sexual
generation.
Certain phenomena which have been observed amongst the
Coniomycetes are cited as examples of sexual association. Amongst these
may be named the conjugation of the slender spores of the first generation,
produced on the germinating threads of Tilletia,[O] and similar acts of
conjugation, as observed in some species of Ustilago. Whether this
interpretation should be placed on those phenomena in the present condition
of our knowledge is perhaps an open question.
Finally, the spermogonia must be regarded as in some occult manner,
which as yet has baffled detection, influencing the perfection of sporidia[P]
In Rhytisma, found on the leaves of maple and willow, black pitchy spots at
first appear, which contain within them a golden pulp, in which very slender
corpuscles are mixed with an abundant mucilage. These corpuscles are the
spermatia, which in Rhytisma acerinum are linear and short, in Rhytisma
salicinum globose. When the spermatia are expelled, the stroma thickens for

the production of asci and sporidia,
which are afterwards developed during
the autumn and winter.
Several of the species of Hysterium
also possess spermogonia, notably H.
Fraxini, which may be distinguished
from the ascigerous perithecia with
which they are associated by their
smaller size and flask-like shape. From
these the spermatia are expelled long
before the maturity of the spores. In
Hypoderma virgultorum, H. commune,
and H. scirpinum, the spermogonia are
small depressed black capsules, which
contain an abundance of minute Fig. 101.—Tilletia caries with conjugating
cells.
spermatia. These were formerly
regarded as distinct species, under the name of Leptostroma. In Stictis
ocellata a great number of the tubercles do not pass into the perfect state
until after they have produced either linear, very short spermatia, or
stylospores, the latter being reproductive bodies of an oblong shape, equal
in size to the perfect sporidia. Some of the tubercles never pass beyond this
stage.
Again, there is a very common fungus which forms black discoid spots
on dead holly leaves, called Ceuthospora phacidioides, figured by Greville
in his “Scottish Cryptogamic Flora,” which expels a profusion of minute
stylospores; but later in the season, instead of these, we find the asci and
sporidia of Phacidium ilicis, so that the two are forms and conditions the
one of the other.
In Tympanis conspersa the spermogonia are much more commonly met
with than the complete fruit. There is a great external resemblance in them
to the ascigerous cups, but there is no evidence that they are ever
transformed into such. The perfect sporidia are also very minute and
numerous, being contained in asci borne in cups, which usually surround
the spermogonia.

In several species of Dermatea the stylospores and spermatia co-exist,
but they are disseminated before the appearance of the ascigerous
receptacles, yet they are produced upon a common stroma not unlike that of
Tubercularia.
In its early stage the common and well-known Bulgaria inquinans, which
when mature looks like a black Peziza, is a little tubercle, the whole mass of
which is divided into ramified lobes, the extremities of which become,
towards the surface of the tubercle, receptacles from whence escape waves
of spermatia which are colourless, or stylospores mixed with them which
are larger and nearly black.
Amongst the Sphæriacei numerous instances might be cited of minute
stylosporous bodies in consort with, or preceding, the ascigerous
receptacles. A very familiar example may be found at the base of old nettle
stems in what has been named Aposphæria acuta, but which truly are only
the stylospores of the Sphæria coniformis, the perithecia of which flourish
in company or in close proximity to them. Most of these bodies are so
minute, delicate, and hyaline that the difficulties in the way of tracing them
in their relations to the bodies with which they are associated are very great.
Nevertheless there is strong presumption in favour of regarding some of
them as performing the functions which the name applied to them indicates.
Professor de Bary cautiously refrains from accepting spermatia other than
as doubtful or at least uncertain sexual bodies.[Q] He says that the Messrs.
Tulasne have supposed that the spermogonia represented the male sex, and
that the spermatia were analogous to spermatozoids. Their opinion depends
on two plausible reasons,—the spermatia, in fact, do not germinate, and the
development of the spermogonia generally precedes the appearance of the
sporophorous organs, a double circumstance which reminds us of what is
known of the spermatozoids and antheridia of other vegetables. It remained
to discover which were the female organs which underwent fecundation
from the spermatia.
Many organs placed at first amongst spermatia have been recognized by
M. Tulasne as being themselves susceptible of germination, and
consequently ought to take their place among legitimate spores. Then it
must be considered that very many spores can only germinate under certain
conditions. It is, therefore, for the present a doubtful question whether there

exist really any spermatia incapable of germination, or if the default of
germination of these corpuscles does not rather depend on the experiments
hitherto attempted not having included the conditions required by the
phenomena. Moreover, as yet no trace has been discovered of the female
organs which are specially fecundated by the spermatia.
Finally, there exist in the Ascomycetes certain organs of reproduction,
diverse spore-bearing apparatus, pycnidia, and others, which, like the
spermogonia, usually precede ascophorous fruits. The real nature of the
spermogonia and spermatia should therefore be regarded as, at present, very
uncertain; as regards, however, the spermatia which have never been seen to
germinate, perhaps it is as well not to absolutely reject the first opinion
formed concerning them, or perhaps they might be thought to perform the
part of androspores, attributing to that expression the meaning which
Pringsheim gives it in the Conferoæ. The experiments performed with the
spermatia which do not germinate, and with the spermogonia of the
Uredines, do not, at any rate, appear to justify the reputed masculine or
fecundative nature of these organs. The spermogonia constantly accompany
or precede fruits of Æcidium, whence naturally follows the presumption
that the first are in a sexual relation to the second. Still, when Tulasne
cultivated Endophyllum sempervivum, he obtained on some perfectly
isolated rosettes of Sempervivum some Æcidium richly provided with
normal and fertile spores, without any trace of spermogonia or of spermatia.
[A]
M. Tulasne has devoted a chapter to the spermogonia of the Uredines in his memoir,
to which we have already alluded.
[B]
Œersted, in “Verhandl der König. Dän. Gesell. Der Wissensch,” 1st January, 1865; De
Bary, “Handbuch der Physiol. Botanik” (1866), p. 172; “Annales des Sci. Nat.” (5me
sér.), vol. v. (1866), p. 366.
[C]
Van Tieghem and Le Monnier, in “Annales des Sci. Nat.” (1873), vol. xvii. p. 261.
[D]
Brefeld, “Bot. Unt. uber Schimmelpilze,” p. 31.
[E]

De Bary, “Morphologie und Physiologie der Pilze,” cap. 5, p. 160; “Ann. des Sci.
Nat.” (1866), p. 343.
[F]

Cornu, in “Ann. des Sci. Nat.” (5me sér.), vol. xv. p. 1 (1872).
[G]
Pringsheim’s “Jahrbucher,” vol. ii. p. 169.
[H]

De Bary, in “Annales des Sciences Naturelles” (5me sér.), vol. v. (1866), p. 343;
Hoffmeister’s “Handbook” (Fungi), cap. v. p. 155.
[I]

De Bary, in “Annales des Sci. Nat.” (4me sér.), vol. xx. p. 129.
[J]

De Bary, in “Annales des Sciences Naturelles” (5me sér.), p. 343.
[K]
Woronin, in De Bary’s “Beitr. zur. Morph. und Physiol. der Pilze,” ii. (1866), pp. 1–
11.
[L]

Tulasne, “Ann. des Sci. Nat.” (5me sér.), October, 1866, p. 211.
[M]
Tulasne, “On the Phenomena of Copulation in certain Fungi,” in “Ann. des Sci. Nat.”
(1866), p. 211.
[N]
De Bary, “Morphologie und Phys. der Pilze,” cap. v., p. 162.
[O]

Berkeley, in “Journ. Hort. Soc.” vol ii. p. 107; Tulasne, “Ann. d. Sc. Nat.” (4me sér.),
vol. ii. tab. 12.
[P]
Tulasne, “New Researches on the Reproductive Apparatus of Fungi;” “Comptes
Rendus,” vol. xxxv. (1852), p. 841.
[Q]
De Bary, “Morphologie und Physiologie der Pilze,” cap. v. p. 168.

IX.
POLYMORPHISM.

A great number of very interesting facts have during late years been brought
to light of the different forms which fungi assume in the course of their
development. At the same time, we fear that a great many assumptions have
been accepted for fact, and supposed connections and relations between two
or three or more so-called species, belonging to different genera, have upon
insufficient data been regarded as so many states or conditions of one and
the same plant. Had the very pertinent suggestions of Professor de Bary
been more generally acted upon, these suspicions would have been
baseless. His observations are so valuable as a caution, that we cannot
forbear prefacing our own remarks on this subject by quoting them.[A] In
order to determine, he says, whether an organic form, an organ, or an
organism, belongs to the same series of development as another, or that
which is the same is developed from it, or vice versâ, there is only one way,
viz., to observe how the second grows out of the first. We see the
commencement of the second begin as a part of the first, perfect itself in
connection with it, and at last it often becomes independent; but be it
through spontaneous dismembering from the first, or that the latter be
destroyed and the second remains, both their disunited bodies are always
connected together in organic continuity, as parts of a whole (single one)
that can cease earlier or later.
By observing the organic continuity, we know that the apple is the
product of development of an apple-tree, and not hung on it by chance, that
the pip of an apple is a product of the development of the apple, and that
from the pip an apple-tree can at last be developed, that therewith all these
bodies are members of a sphere of development or form. It is the same with
every similar experience of our daily life, that where an apple-tree stands,
many apples lie on the ground, or that in the place where apple-pips are
sown seedlings, little apple-trees, grow out of the ground, is not important

to our view of the course of development. Every one recognizes that in his
daily life, because he laughs at a person who thinks a plum which lies under
an apple-tree has grown on it, or that the weeds which appear among the
apple seedlings come from apple-pips. If the apple-tree with its fruit and
seed were microscopically small, it would not make the difference of a
hair’s breadth in the form of the question or the method of answering it, as
the size of the object can be of no importance to the latter, and the questions
which apply to microscopical fungi are to be treated in the same manner.
If it then be asserted that two or several forms belong to a series of
development of one kind, it can only be based on the fact of their organic
continuity. The proof is more difficult than in large plants, partly because of
the delicacy, minuteness, and fragility of the single parts, particularly the
greater part of the mycelia, partly because of the resemblance of the latter in
different species, and therefore follows the danger of confusing them with
different kinds, and finally, partly in consequence of the presence of
different kinds in the same substratum, and therefore the mixture not only
of different sorts of mycelia, but also that different kinds of spores are
sown. With some care and patience, these difficulties are in no way
insurmountable, and they must at any rate be overcome; the organic
continuity or non-continuity must be cleared up, unless the question
respecting the course of development, and the series of forms of special
kinds, be laid on one side as insolvable.
Simple and intelligible as these principles are, they have not always been
acted upon, but partly neglected, partly expressly rejected, not because they
were considered false, but because the difficulties of their application were
looked upon as insurmountable. Therefore another method of examination
was adopted; the spores of a certain form were sown, and sooner or later
they were looked after to see what the seed had produced—not every single
spore—but the seed en masse, that is, in other words, what had grown on
that place where the seed had been sown. As far as it relates to those forms
which are so widely spread, and above all grow in conjunction with one
another—and that is always the case in the specimens of which we speak—
we can never be sure that the spores of the form which we mean to test are
not mingled with those of another species. He who has made an attentive
and minute examination of this kind knows that we may be sure to find
such a mixture, and that such an one was there can be afterwards decidedly

proved. From the seed which is sown, these spores, for which the
substratum was most suitable, will more easily germinate, and their
development will follow the more quickly. The favoured germs will
suppress the less favoured, and grow up at their expense. The same relation
exists between them as between the seeds, germs, and seedlings of a sown
summer plant, and the seeds which have been undesignedly sown with it,
only in a still more striking manner, in consequence of the relatively quick
development of the mildew fungus.
Therefore, that from the latter a decided form, or a mixture of several
forms, is to be found sown on one spot, is no proof of their generic
connection with one which has been sown for the purpose of experiments;
and the matter will only be more confused if we call imagination to our aid,
and place the forms which are found near one another, according to a real or
fancied resemblance, in a certain series of development. All those
statements on the sphere of form and connection, which have for their basis
such a superficial work, and are not based on the clear exposition of the
continuity of development, as by the origin of the connection of the Mucor
with Penicillium, Oidium lactis and Mucor, Oidium and Penicillium, are
rejected as unfounded.
A source of error, which can also interfere in the last-named superficial
method of cultivation for experiments, is, viz., that heterogeneous
unwished-for spores intrude themselves from without, among the seed
which is sown, but that has been until now quite disregarded. It is of great
importance in practice, but in truth, for our present purpose, synonymous
with what we have already written. Those learned in the science of this kind
of culture lay great stress on its importance, and many apparatuses have
been constructed, called “purely cultivating machines,” for the purpose of
destroying the spores which are contained in the substratum, and preventing
the intrusion of those from without. The mixture in the seed which is sown
has of course not been obviated. These machines may, perhaps, in every
other respect, fulfil their purpose, but they cannot change the form of the
question, and the most ingeniously constructed apparatus cannot replace the
attention and intellect of the observer.[B]
Two distinct kinds of phenomena have been grouped under the term
“polymorphy.” In one series two or more forms of fruit occur consecutively
or simultaneously on the same individual, and in the other two or more

forms appear on a different mycelium, on a different part of the same plant,
or on a matrix wholly distinct and different; in the latter case the connection
being attested or suspected circumstantially, in the former proved by the
method suggested by De Bary. It will at once be conceded that in cases
where actual growth and development substantiate the facts the polymorphy
is undoubted, whilst in the other series it can at best be little more than
suspected. We will endeavour to illustrate both these series by examples.
One of the first and earliest suspected cases of dualism, which long
puzzled the older mycologists, was observed amongst the Uredines, and
many years ago it was held that there must be some mysterious association
between the “red rust” (Trichobasis ruligo vera) of wheat and grasses and
the “corn mildew” (Puccinia graminis) which succeeded it. The simple
spored rust first makes its appearance, and later the bilocular “mildew.” It is
by no means uncommon to find the two forms in the same pustule. Some
have held, without good reason, that the simple cells became afterwards
divided and converted into Puccinia, but this is not the case; the uredo-
spores are always simple, and remain so except in Uredo linearis, where
every intermediate stage has been observed. Both are also perfect in their
kind, and capable of germination.
What the precise relations between the two forms may be has as yet
never been revealed to observers, but that the two forms belong to one
species is not now doubted. Very many species of Puccinia have already
been found associated with a corresponding Trichobasis, and of
Phragmidium with a relative Lecythea, but it may be open to grave doubt
whether some of the very many species associated by authors are not so
classed upon suspicion rather than observation. We are ready to admit that
the evidence is strong in favour of the dimorphism of a large number of
species—it may be in all, but this awaits proof, or substantial presumption
on good grounds. Up to the present we know that there are species of
Trichobasis which have never been traced to association with a Puccinia,
and doubtless there will be species of Puccinia for which no corresponding
Uredo or Trichobasis can be found.
Tulasne remarks, in reference to Puccinia sonchi, in one of his memoirs,
that this curious species exhibits, in effect, that a Puccinia may unite three
sorts of reproductive bodies, which, taking part, constitute for the
mycologists of the day three entirely different plants—a Trichobasis, a

Uromyces, and a Puccinia. The Uredines are not less rich, he adds, in
reproductive bodies of divers sorts than the Pyrenomycetes and the
Discomycetes; and we should not be surprised at this, since it seems to be a
law, almost constant in the general harmony of nature, that the smaller the
organized beings are, the more their races are prolific.
In Puccinia variabilis, Grev., it is common to find a unicellular form,
species of Trichobasis, in the same pustules. A like circumstance occurs
with Puccinia violarum, Link., and Trichobasis violarum, B.; with Puccinia
fallens, C., and Trichobasis fallens, Desm.; also with Puccinia menthæ, P.,
and Trichobasis Labiatarum, D. C. In Melampsora, again, the prismatic
pseudospores of Melampsora salicina, Lev., are the winter fruits of
Lecythea caprearum, Lev., as those of Melampsora populina, Lev., are of
Lecythea populina, Lev. In the species of Lecythea themselves will be
found, as De Bary[C] has shown, hyaline cysts of a larger size, which
surround the pseudospores in the pustules in which they are developed.
A good illustration of dimorphism in one of the commonest of moulds is
given by De Bary in a paper from which we have already quoted.[D] He
writes thus:—In every household there is a frequent unbidden guest, which
appears particularly on preserved fruits, viz., the mould which is called
Aspergillus glaucus. It shows itself to the naked eye as a woolly floccy crust
over the substance, first purely white, then gradually covered with little fine
glaucous, or dark green dusty heads. More minute microscopical
examination shows that the fungus consists of richly ramified fine
filaments, which are partly disseminated in the substratum, and partly raised
obliquely over it. They have a cylindrical form with rounded ends, and are
divided into long outstretched members, each of which possesses the
property which legitimatizes it as a vesicle in the ordinary sense of the
word; it contains, enclosed within a delicate structureless wall, those bodies
which bear the appearance of a finely granulated mucous substance, which
is designated by the name of protoplasm, and which either equally fills the
cells, or the older the cell the more it is filled with watery cavities called
vacuoles.
All parts are at first colourless. The increase in the length of the filaments
takes place through the preponderating growth near their points; these
continually push forward, and, at a short distance from them, successive
new partitions rise up, but at a greater distance, the growth in the length

ceases. This kind of growth is called point growth. The twigs and branches
spring up as lateral dilatations of the principal filament, which, once
designed, enlarges according to the point growth. This point growth of
every branch is, to a certain extent, unlimited. The filaments in and on the
substratum are the first existing members of the fungus; they continue so
long as it vegetates. As the parts which absorb nourishment from and
consume the substance, they are called the mycelium. Nearly every fungus
possesses a mycelium, which, without regard to the specific difference of
form and size, especially shows the described nature in its construction and
growth.
The superficial threads of the mycelium produce other filaments beside
those numerous branches which have been described, and which are the
fruit thread (carpophore) or conidia thread. These are on an average thicker
than the mycelium threads, and only exceptionally ramified or furnished
with partitions; they rise almost perpendicularly into the air, and attain a
length of, on an average, half a millimetre, or one-fiftieth of an inch, but
they seldom become longer, and then their growth is at an end. Their free
upper end swells in a rounded manner, and from this is produced, on the
whole of its upper part, rayed divergent protuberances, which attain an oval
form, and a length almost equal to their radius, or, in weaker specimens, the
diameter of the rounded head. The rayed divergent protuberances are the
direct producers and bearers of the propagating cells, spores, or conidia, and
are called sterigmata. Every sterigma at first produces at its point a little
round protuberance, which, with a strong narrow basis, rests upon the
sterigma. These are filled with protoplasm, swell more and more, and, after
some time, separate themselves by a partition from the sterigma into
independent cells, spores, or conidia.
The formation of the first spore takes place at the same end of the
sterigma, and in the same manner a second follows, then a third, and so on;
every one which springs up later pushes its predecessor in the direction of
the axis of the sterigma in the same degree in which it grows itself; every
successive spore formed from a sterigma remains for a time in a row with
one another. Consequently every sterigma bears on its apex a chain of
spores, which are so much the older, the farther they stand from the
sterigma. The number of the links in a chain of spores reaches in normal
specimens to ten or more. All sterigmata spring up at the same time, and

keep pace with one another in the formation of the spores. Every spore
grows for a time, according to its construction, and at last separates itself
from its neighbours. The mass of dismembered spores forms that fine
glaucous hue which is mentioned above. The spores, therefore, are
articulated in rows, one after the other, from the ends of the sterigmata. The
ripe spore, or conidium, is a cell of a round or broadly oval form, filled with
a colourless protoplasm, and, if observed separately, is found to be provided
with a brownish, finely verruculose, dotted wall.

Fig. 102.—a. Aspergillus glaucus; b. conidia; c.
germinating conidium; d. conceptacle of Eurotium; e.
ascus.

The same mycelium which forms the pedicel for the conidia when it is
near the end of its development, forms by normal vegetation a second kind
of fructification. It begins as delicate thin little branches, which are not to
be distinguished by the naked eye, and which mostly in four or six turns,

after a quickly terminated growth, wind their ends like a corkscrew. (Fig.
102.) The sinuations decrease in width more and more, till they at last reach
close to one another, and the whole end changes from the form of a
corkscrew into that of a hollow screw. In and on that screw-like body, a
change of a complicated kind takes place, which is a productive process. In
consequence of this, from the screw body a globose receptacle is formed,
consisting of a thin wall of delicate cells, and a closely entwined row of
cells surrounded by this dense mass (d). By the enlargement of all these
parts the round body grows so much, that by the time it is ripe it is visible to
the naked eye. The outer surface of the wall assumes a compactness and a
bright yellow colour; the greater part of the cells of the inner mass become
asci for the formation of sporidia, while they free themselves from the
reciprocal union, take a broad oval form, and each one produces within its
inner space eight sporidia (e). These soon entirely fill the ascus. When they
are quite ripe, the wall of the conceptacle becomes brittle, and from
irregular fissures, arising easily from contact, the colourless round sporidia
are liberated.
The pedicels of both kinds of fruit are formed from the same mycelium in
the order just described. If we examine attentively, we can often see both
springing up close to one another from the same filament of a mycelium.
This is not very easy in the close interlacing of the stalks of a mass of fungi
in consequence of their delicacy and fragility. Before their connection was
known, the conceptacles and the conidia pedicels were considered as organs
of two very different species of fungi. The conceptacles were called
Eurotium herbariorum, and the conidia bearers were called Aspergillus
glaucus.
Allied to Eurotium is the group of Erysiphei, in which well-authenticated
polymorphy prevails. These fungi are developed on the green parts of
growing plants, and at first consist of a white mouldy stratum, composed of
delicate mycelium, on which erect threads are produced, which break up
into subglobose joints or conidia. The species on grass was named Oidium
monilioides before its relationship was known, but undoubtedly this is only
the conidia of Erysiphe graminis. In like manner the vine disease (Oidium
Tuckeri) is most probably only the conidia of a species of Erysiphe, of
which the perfect condition has not yet been discovered. On roses the old
Oidium leucoconium is but the conidia of Sphærotheca pannosa, and so of

other species. The Erysiphe which
ultimately appears on the same
mycelium consists of globose
perithecia, externally furnished with
thread-like appendages, and internally
with asci containing sporidia. In this
genus there are no less than five
different forms of fruit,[E] the
multiform threads on the mycelium,
already alluded to as forms of Oidium,
the asci contained in the sporangia,
which is the proper fruit of the
Erysiphe, larger stylospores which are Fig. 103.—Erysiphe cichoracearum. a.
produced in other sporangia, the Receptacle; o. mycelium. (De Bary.)
smaller stylospores which are
generated in the pycnidia, and separate sporules which are sometimes
formed in the joints of the necklaces of the conidia. These forms are figured
in the “Introduction to Cryptogamic Botany” from Sphærotheca Castagnei,
which is the hop mildew.[F] The vine disease, hop mildew, and rose mildew,
are the most destructive species of this group, and the constant annoyance
of cultivators.
When first describing an allied fungus found on old paper, and named
Ascotricha chartarum, the Rev. M. J. Berkeley called attention to the
presence of globose conidia attached to the threads which surround the
conceptacles,[G] and this occurred as long since as 1838. In a recent species
of Chætomium found on old sacking, Chætomium griseum, Cooke,[H] we
have found tufts in all respects similar externally to the Chætomium, but no
perithecium was formed, naked conidia being developed apparently at the
base of the coloured threads. In Chætomium funicolum, Cooke, a black
mould was also found which may possibly prove to be its conidia, but at
present there is no direct evidence.
The brothers Tulasne have made us acquainted with a greater number of
instances amongst the Sphæriacei in which multiple organs of reproduction
prevail. Very often old and decaying individuals belonging to species of
Boletus will be found filled, and their entire substance internally replaced,
by the threads and multitudinous spores of a golden yellow parasite, to

which the name of Sepedonium chrysospermum has been given. According
to Tulasne, this is merely a condition of a sphæriaceous fungus belonging to
his genus Hypomyces.[I]
The same observers also first demonstrated that Trichoderma viride, P.,
was but the conidia-bearing stage of Hypocrea rufa, P., another
sphæriaceous fungus. The ascigerous stroma of the latter is indeed
frequently associated in a very close manner with the cushions of the
pretended Trichoderma, or in other cases the same stroma will give rise to a
different apparatus of conidia, of which the principal elements are acicular
filaments, which are short, upright, and almost simple, and which give rise
to small oval conidia which are solitary on the tips of the threads. Therefore
this Hypocrea will possess two different kinds of conidia, as is the case in
many species of Hypomyces.
A most familiar instance of dualism will be found in Nectria
cinnabarina, of which the conidia form is one of the most common of
fungi, forming little reddish nodules on all kinds of dead twigs.[J]
Almost any small currant twig which has been lying
on the ground in a damp situation will afford an
opportunity of studying this phenomenon. The whole
surface of the twig will be covered from end to end with
little bright pink prominences, bursting through the bark
at regular distances, scarcely a quarter of an inch apart.
Towards one end of the twig probably the prominences
will be of a deeper, richer colour, like powdered
cinnabar. The naked eye is sufficient to detect some
difference between the two kinds of pustules, and where
the two merge into each other specks of cinnabar will
be visible on the pink projections. By removing the
bark it will be seen that the pink bodies have a sort of
paler stem, which spreads above into a somewhat
globose head, covered with a delicate mealy bloom. At
the base it penetrates to the inner bark, and from it the Fig. 104.—Twig with
threads of mycelium branch in all directions, confined, Tubercularia on the
however, to the bark, and not entering the woody upper portion, Nectria
on the lower.
tissues beneath. The head, placed under examination,
will be found to consist of delicate parallel threads compacted together to

form the stem and head. Some of these threads are simple, others are
branched, bearing here and there upon them delicate little bodies, which are
readily detached, and which form the mealy bloom which covers the
surface. These are the conidia, little slender cylindrical bodies, rounded at
the ends.
Passing to the other bodies, which are of a deeper colour, it will soon be
discovered that, instead of being simple rounded heads, each tubercle is
composed of numerous smaller, nearly globose bodies, closely packed
together, often compressed, all united to a base closely resembling the base
of the other tubercles. If for a moment we look at one of the tubercles near
the spot where the crimson tubercles seem to merge into the pink, we shall
not only find them particoloured, but that the red points are the identical
globose little heads just observed in clusters. This will lead to the suspicion,
which can afterwards be verified, that the red heads are really produced on
the stem or stroma of the pink tubercles.

Fig. 105.—Section of Tubercularia. c. Threads with conidia.[K]

A section of one of the red tubercles will show us how much the internal
structure differs. The little subglobose bodies which spring from a common
stroma or stem are hollow shells or capsules, externally granular, internally
filled with a gelatinous nucleus. They are, indeed, the perithecia of a
sphæriaceous fungus of the genus Nectria, and the gelatinous nucleus
contains the fructification. Still further examination will show that this
fructification consists of cylindrical asci, each enclosing eight elliptical
sporidia, closely packed together, and mixed with slender threads called
paraphyses.

Here, then, we have undoubted evidence of Nectria cinnabarina, with its
fruit, produced in asci growing from the stroma or stem, and in intimate
relationship with what was formerly named Tubercularia vulgaris. A fungus
with two forms of fruit, one proper to the pink, or Tubercularia form, with
naked slender conidia, the other proper to the mature fungus, enclosed in
asci, and generated within the walls of a perithecium. Instances of this kind
are now known to be far from uncommon, although they cannot always, or
often, be so clearly and distinctly traced as in the illustration which we have
selected.

Fig. 106.—D. Nectria surrounding Tubercularia; E. tuft of Nectria
cinnabarina; F. section of stroma; G. ascus and paraphyses.

It is not uncommon for the conidia of the Sphæria to partake of the
characteristics of a mould, and then the perithecia are developed amongst
the conidial threads. A recently recorded instance of this relates to Sphæria
Epochnii, B. and Br.,[L] the conidia form of which was long known before
the Sphæria related to it was discovered, under the name of Epochnium
fungorum. The Epochnium forms a thin stratum, which overruns various
species of Corticium. The conidia are at first uniseptate. The perithecia of
the Sphæria are at first pale bottle-green, crowded in the centre of the
Epochnium, then black green granulated, sometimes depressed at the
summit, with a minute pore. The sporidia are strongly constricted in the
centre, at first uniseptate, with two nuclei in each division.

Another Sphæria in which the association is undoubted is the Sphæria
aquila, Fr.,[M] which is almost always found nestling in a woolly brown
subiculum, for the most part composed of barren brown jointed threads.
These threads, however, produce, under favourable conditions, mostly
before the perfection of the perithecia, minute subglobose conidia, and in
this state constitute what formerly bore the name of Sporotrichum fuscum,
Link., but now recognized as the conidia of Sphæria aquila.
In Sphæria nidulans, Schw., a North American species, we have more
than once found the dark brown subiculum bearing large triseptate conidia,
having all the characters of the genus Helminthosporium. In Sphæria pilosa,
P., Messrs. Berkeley and Broome have observed oblong conidia, rather
irregular in outline, terminating the hairs of the perithecium.[N] The same
authors have also figured the curious pentagonal conidia springing from
flexuous threads accompanying Sphæria felina, Fckl.,[O] and also the
threads resembling those of a Cladotrichum with the angular conidia of
Sphæria cupulifera, B. and Br.[P] A most remarkable example is also given
by the Brothers Tulasne in Pleospora polytricha, in which the conidia-
bearing threads not only surround, but grow upon the perithecia, and are
crowned by fascicles of septate conidia.[Q]
Instances of this kind have now become so numerous that only a few can
be cited as examples of the rest. It is not at all improbable that the majority
of what are now classed together as species under the genus of black
moulds, Helminthosporium, will at some not very distant period be traced
as the conidia of different species of ascomycetous fungi. The same fate
may also await other allied genera, but until this association is established,
they must keep the rank and position which has been assigned to them.
Another form of dualism, differing somewhat in character from the
foregoing, finds illustration in the sphæriaceous genus Melanconis, of
Tulasne, in which the free spores are still called conidia, though in most
instances produced in a sort of spurious conceptaculum, or borne on short
threads from a kind of cushion-shaped stroma. In the Melanconis
stilbostoma,[R] there are three forms, one of slender minute bodies, oozing
out in the form of yellow tendrils, which may be spermatia, formerly called
Nemaspora crocea. Then there are the oval brown or olive brown conidia,
which are at first covered, then oozing out in a black pasty mass, formerly
Melanconium bicolor, and finally the sporidia in asci of Sphæria

stilbostoma, Fries. In Melanconis Berkeleii, Tul., the conidia are
quadrilocular, previously known as Stilbospora macrosperma, B. and Br. In
a closely-allied species from North America, Melanconis bicornis, Cooke,
the appendiculate sporidia are similar, and the conidia would also appear to
partake of the character of Stilbospora. We may remark here that we have
seen a brown mould, probably an undescribed species of Dematiei, growing
in definite patches around the openings in birch bark caused by the
crumpent ostiola of the perithecia of Melanconis stilbostoma, from the
United States.
In Melanconis lanciformis,[S] Tul., there are, it would appear, four forms
of fruit. One of these consists of conidia, characterized by Corda as
Coryneum disciforme.[T] Stylospores, which are also figured by Corda
under the name of Coniothecium betulinum; pycnidia,[U] first discovered by
Berkeley and Broome, and named by them Hendersonia polycystis;[V] and
the ascophorous fruits which constituted the Sphæria lanciformis of Fries.
Mr. Currey indicated Hendersonia polycystis, B. and Br., as a form of fruit
of this species in a communication to the Royal Society in 1857.[W] He says
this plant grows upon birch, and is in perfection in very moist weather,
when it may be recognized by the large black soft gelatinous protuberances
on the bark, formed by spores escaping and depositing themselves upon and
about the apex of the perithecium. This I suspect to be an abnormal state of
a well-known Sphæria (S. lanciformis), which grows upon birch, and upon
birch only.
We might multiply, almost indefinitely, instances amongst the
Sphæriacei, but have already given sufficient for illustration, and will
therefore proceed briefly to notice some instances amongst the
Discomycetes, which also bear their complete or perfect fruit in asci.
The beautiful purple stipitate cups of Bulgaria sarcoides, which may be
seen flourishing in the autumn on old rotten wood, are often accompanied
by club-shaped bodies of the same colour; or earlier in the season these
clavate bodies may be found alone, and at one time bore the name of
Tremella sarcoides. The upper part of these clubs disseminate a great
abundance of straight and very slender spermatia. Earlier than this they are
covered with globose conidia. The fully-matured Bulgaria develops on its
hymenium clavate delicate asci, each enclosing eight elongated hyaline
sporidia, so that we have three forms of fruit belonging to the same fungus,

viz. conidia and spermatia in the Tremella stage, and sporidia contained in
asci in the mature condition.[X] The same phenomena occur with Bulgaria
purpurea, a larger species with different fruit, long confounded with
Bulgaria sarcoides.
On the dead stems of nettles it is very common to meet with small orange
tubercles, not much larger than a pin’s head, which yield at this stage a
profusion of slender linear bodies, produced on delicate branched threads,
and at one time bore the name of Dacrymyces Urticæ, but which are now
acknowledged to be only a condition of a little tremelloid Peziza of the
same size and colour, which might be mistaken for it, if not examined with
the microscope, but in which there are distinct asci and sporidia. Both forms
together are now regarded as the same fungus, under the name of Peziza
fusarioides, B.
The other series of phenomena grouped together under the name of
polymorphism relate to forms which are removed from each other, so that
the mycelium is not identical, or, more usually, produced on different
plants. The first instance of this kind to which we shall make reference is
one of particular interest, as illustrative of the old popular creed, that
berberry bushes near corn-fields produced mildewed corn. There is a village
in Norfolk, not far from Great Yarmouth, called “Mildew Rollesby,”
because of its unenviable notoriety in days past for mildewed corn,
produced, it was said, by the berberry bushes, which were cut down, and
then mildew disappeared from the corn-fields, so that Rollesby no longer
merited its sobriquet. It has already been shown that the corn-mildew
(Puccinia graminis) is dimorphous, having a one-celled fruit (Trichobasis),
as well as a two-celled fruit (Puccinia). The fungus which attacks the
berberry is a species of cluster-cup (Æcidium berberidis), in which little
cup-like peridia, containing bright orange pseudospores, are produced in
tufts or clusters on the green leaves, together with their spermogonia.
De Bary’s observations on this association of forms were published in
1865.[Y] In view of the popular belief, he determined to sow the spores of
Puccinia graminis on the leaves of the berberry. For this purpose he
selected the septate resting spores from Poa pratensis and Triticum repens.
Having caused the spores to germinate in a moist atmosphere, he placed
fragments of the leaves on which they had developed their secondary spores
on young but full-grown berberry leaves, under the same atmospheric

conditions. In from twenty-four to forty-eight hours a quantity of the
germinating threads had bored through the walls and penetrated amongst
the subjacent cells. This took place both on the upper and under surface of
the leaves. Since, in former experiments, it appeared that the spores would
penetrate only in those cases where the plant was adapted to develop the
parasite, the connection between P. graminis and Æcid. berberidis seemed
more than ever probable. In about ten days the spermogonia appeared. After
a time the cut leaves began to decay, so that the fungus never got beyond
the spermogonoid stage. Some three-year-old seedlings were then taken,
and the germinating resting spores applied as before. The plants were kept
under a bell-glass from twenty-four to forty-eight hours, and then exposed
to the air like other plants. From the sixth to the tenth day, yellow spots
appeared, with single spermogonia; from the ninth to the twelfth,
spermogonia appeared in numbers on either surface; and, a few days later,
on the under surface of the leaves, the cylindrical sporangia of the Æcidium
made their appearance, exactly as in the normally developed parasite,
except that they were longer, from being protected from external agents.
The younger the leaves, the more rapid was the development of the parasite,
and sometimes, in the younger leaves, the luxuriance was far greater than in
free nature. Similar plants, to the number of two hundred, were observed in
the nursery, and though some of them had Æcidium pustules, not one fresh
pustule was produced; while two placed under similar circumstances, but
without the application of any resting spores, remained all the summer free
from Æcidium. It seems, then, indubitable so far that Æcidium berberidis
does spring from the spores of Puccinia graminis.
It has, however, to be remarked that De Bary was not equally successful
in producing the Puccinia from the spores of the Æcidium. In many cases
the spores do not germinate when placed on glass, and they do not preserve
their power of germinating very long. He reverts then to the evidence of
experiments instituted by agriculturists. Bönninghausen remarked, in 1818,
that wheat, rye, and barley which were sown in the neighbourhood of a
berberry bush covered with Æcidium contracted rust immediately after the
maturation of the spores of the Æcidia. The rust was most abundant where
the wind carried the spores. The following year the same observations were
repeated; the spores of the Æcidium were collected, and applied to some
healthy plants of rye. After five or six days these plants were affected with
rust, while the remainder of the crop was sound. In 1863 some winter rye

was sown round a berberry bush, which in the following year was infested
with Æcidium, which was mature in the middle of May, when the rye was
completely covered with rust. Of the wild grasses near the bush, Triticum
repens was most affected. The distant plants of rye were free from rust.

Fig. 107.—Cells and pseudospores of Æcidium berberidis.

The spores of the Æcidium would not germinate on berberry leaves; the
berberry Æcidium could not therefore spring from the previous Æcidium.
The uredospores of Puccinia graminis on germinating penetrate into the
parenchym of the grass on which they are sown; but on berberry leaves, if
the tips of the threads enter for a short distance into the stomates their
growth at once ceases, and the leaves remain free from parasites.

Fig. 108.—Cells and pseudospores of Æcidium graveolens.

Montagne has, however, described a Puccinia berberidis on leaves of
Berberis glauca from Chili, which grows in company with Æcidium
berberidis. This at first sight seems to contradict the above conclusions; but
the Æcidium which from the same disc produces the puccinoid resting
spores, appears to be different from the European species, inasmuch as the
cells of the wall of the sporangium are twice as large, and the spores
decidedly of greater diameter.[Z] The resting spores, moreover, differ not
only from those of Puccinia graminis, but from those of all other European
species.

From this account, then, it is extremely probable that the Æcidium of the
berberry enters into the cycle of existence of Puccinia graminis, and, if this
be true, wherefore should not other species of Puccinia be related in like
manner to other Æcidia? This is the conclusion to which many have
arrived, and, taking advantage of certain presumptions, have, we fear, rashly
associated many such forms together without substantial evidence. On the
leaves of the primrose we have commonly a species of Æcidium, Puccinia,
and Uromyces nearly at the same time; we may imagine that all these
belong to one cycle, but it has not yet been proved. Again, Uromyces
cacaliæ, Unger, Uredo cacaliæ, Unger, and Æcidium cacaliæ, Thumen, are
considered by Heufler[a] to form one cycle. Numerous others are given by
Fuckel,[b] and De Bary, in the same memoir from which we have already
cited, notes Uromyces appendiculatus, Link., U. phaseolorum, Tul., and
Puccinia tragopogonis, Ca., as possessing five kinds of reproductive
organs. Towards the end of the year, shortly stipitate spores appear on their
stroma, which do not fall off. These spores, which do not germinate till after
a shorter or longer winter rest, may conveniently be called resting spores,
or, as De Bary calls them, teleutospores, being the last which are produced.
These at length germinate, become articulated, and produce ovate or
kidney-shaped spores, which in their turn germinate, penetrating the cuticle
of the mother plant, avoiding the stomates or apertures by which it breathes.
After about two or three weeks, the mycelium, which has ramified among
the tissues, produces an Æcidium, with its constant companion,
spermogonia—distinct cysts, that is, from which a quantity of minute
bodies ooze out, often in the form of a tendril, the function of which is
imperfectly known at present, but which from analogy we regard as a form
of fruit, though it is just possible that they may be rather of the nature of
spermatozoids. The Æcidia contain, within a cellular membranous sac, a
fructifying disc, which produces necklaces of spores, which ultimately
separate from each other in the form of a granular powder. The grains of
which it is composed germinate in their turn, no longer avoiding the
stomates as before, but penetrating through their aperture into the
parenchym. The new resultant mycelium reproduces the Uredo, or fifth
form of fructification, and the Uredo spores fall off like those of the
Æcidium, and in respect of germination, and mode of penetration, present
precisely the same phenomena. The disc which has produced the Uredo
spores now gives rise to the resting spores, and so the cycle is complete.[c]

The late Professor Œrsted, of Copenhagen, was of opinion that he had
demonstrated the polymorphy of the Tremelloid Uredines, and satisfied
himself that the one condition known as Podisoma was but another stage of
Rœstelia.[d] Some freshly gathered specimens of Gymnosporangium were
damped with water, and during the night following the spores germinated
profusely, so that the teleutospores formed an orange-coloured powder. A
little of this powder was placed on the leaves of five small sorbs, which
were damped and placed under bell-glasses. In five days yellow spots were
seen on the leaves, and in two days more indications of spermogonia. The
spermatia were discharged, and in two months from the first sowing, the
peridia of Rœstelia appeared, and were developed. “This trial of spores,”
says Œrsted, “has conduced to the result expected, and proves that the
teleutospores of Gymnosporangium, when transported upon the sorb, give
rise to a totally different fungus, the Rœstelia cornuta, that is to say, that an
alternate generation comes between these fungi. They appertain in
consequence to a single species, and the Gymnosporangium ceased to be an
independent species, and must be considered as synonymous with the first
generation of Rœstelia. The spores have been transported upon young
shoots of the juniper-tree, and have now commenced to produce some
mycelium in the bark. There is no doubt that in next spring it will result in
Gymnosporangium.”
Subsequently the same learned professor instituted similar experiments
upon other hosts, with the spores of Podisoma, and from thence he
concluded that Rœstelia and Podisoma, in all their known species, were but
forms the one of the other. Hitherto we are not aware that these results have
been confirmed, or that the sowing of the spores of Rœstelia on juniper
resulted in Podisoma. Such experiments should be received always with
care, and not too hastily accepted in their apparent results as proven facts.
Who shall say that Rœstelia would not have appeared on Sorbus within two
months without the sowing of Podisoma spores?—because it is not by any
means uncommon for that fungus to appear upon that plant. It is true many
mycologists write and speak of Rœstelia and Podisoma (or
Gymnosporangium) as identical; but, as we think, without the evidence
being so complete as to be beyond suspicion. It is, nevertheless, a curious
fact that in Europe the number of species of Rœstelia and Podisoma are
equal, if one species be excluded, which is certainly not a good Podisoma,
for the reception of which a new genus has been proposed.[e]

Amongst the ascigerous fungi will be found a curious but interesting
genus formerly called Cordyceps, but for which Tulasne, in consequence of
the discovery of secondary forms of fruit, has substituted that of Torrubia.[f]
These curious fungi partake more or less of a clavate form, and are parasitic
on insects. The pupæ of moths are sometimes seen bearing upon them the
white branched mould, something like a Clavaria in appearance, to which
the name of Isaria farinosa has been given. According to Tulasne, this is the
conidia form of the bright scarlet, club-shaped body which is also found on
dead pupæ, called Torrubia militaris. An American mould of the same
genus, Isaria sphingum, found on mature moths,[g] is in like manner
declared to be the conidia of Torrubia sphingum; whereas a similar mould,
found on dead spiders, called Isaria arachnophila,[h] is probably of a
similar nature. An allied kind of compact mould, which is parasitic on
Cocci, on the bark of trees, recently found in England by Mr. C. E. Broome,
and named Microcera coccophila,[i] is said by Tulasne to be a condition of
Sphærostilbe, and it is intimated that other productions of a similar
character bear like relations to other sphæriaceous fungi. For many species
of Torrubia no corresponding conidia are yet known.
Some instances might be noted, not without interest, in which the facts of
dimorphism or polymorphism have not been satisfactorily proved, but final
judgment is held in suspense until suspicion is replaced by conviction.
Some years since, a quantity of dead box leaves were collected, on which
flourished at the time a mould named Penicillium roseum. This mould has a
roseate tint, and occurs in patches on the dead leaves lying upon the ground;
the threads are erect and branched above, bearing chains of oblong,
somewhat spindle-shaped spores, or, perhaps more accurately, conidia.
When collected, these leaves were examined, and nothing was observed or
noted upon them except this Penicillium. After some time, certainly
between two and three years, during which period the box remained
undisturbed, circumstances led to the examination again of one or two of
the leaves, and afterwards of the greater number of them, when the patches
of Penicillium were found to be intermixed with another mould of a higher
development, and far different character. This mould, or rather Mucor,
consists of erect branching threads, many of the branches terminating in a
delicate globose, glassy head, or sporangium, containing numerous very
minute subglobose sporidia. This species was named Mucor hyalinus.[j] The
habit is very much like that of the Penicillium, but without any roseate tint.

It is almost certain that the Mucor could not have been present when the
Penicillium was examined, and the leaves on which it had grown were
enclosed in the tin box, but that the Mucor afterwards appeared on the same
leaves, sometimes from the same patches, and, as it would appear, from the
same mycelium. The great difference in the two species lies in the
fructification. In the Penicillium, the spores are naked, and in moniliform
threads; whilst in Mucor the spores are enclosed within globose
membraneous heads or sporangia. Scarcely can we doubt that the Mucor
alluded to above, found thus intermixed, under peculiar circumstances, with
Penicillium roseum, is no other than the higher and more complete form of
that species, and that the Penicillium is only its conidiiferous state. The
presumption in this case is strong, and not so open to suspicion as it would
be did not analogy render it so extremely probable that such is the case,
apart from the fact of both forms springing from the same mass of
mycelium. In such minute and delicate structures it is very difficult to
manipulate the specimens so as to arrive at positive evidence. If a filament
of mycelium could be isolated successfully, and a fertile thread, bearing the
fruit of each form, could be traced from the same individual mycelium
thread, the evidence would be conclusive. In default of such conclusive
evidence, we are compelled to rest with assumption until further researches
enable us to record the assumption as fact.[k]
Apropos of this very connection of Penicillium with Mucor, a similar
suspicion attaches to an instance noted by a wholly disinterested observer to
this effect. “On a preparation preserved in a moist chamber, on the third day
a white speck was seen on the surface, consisting of innumerable ‘yeast’
cells, with some filaments, branching in all directions. On the fourth day
tufts of Penicillium, had developed two varieties—P. glaucum and P. viride.
This continued until the ninth day, when a few of the filaments springing up
in the midst of the Penicillium were tipped with a dewdrop-like dilatation,
excessively delicate—a mere distended pellicle. In some cases they seemed
to be derived from the same filament as others bearing the ordinary
branching spores of Penicillium, but of this I could not be positive. This
kind of fructification increased rapidly, and on the fourteenth day spores
had undoubtedly developed within the pellicle, just as had been observed in
a previous cultivation, precisely similar revolving movements being also
manifested.”[l] Although we have here another instance of Mucor and
Penicillium growing in contact, the evidence is insufficient to warrant more

than a suspicion of their identity, inasmuch as the equally minute spores of
Mucor and Penicillium might have mingled, and each producing its kind, no
relationship whatever have existed between them, except their development
from the same matrix.
Another case of association—for the evidence does not proceed further—
was recorded by us, in which a dark-coloured species of Penicillium was
closely associated with what we now believe to be a species of
Macrosporium—but then designated a Sporidesmium—and a minute
Sphæria growing in succession on damp wall-paper. Association is all that
the facts warrant us in calling it.
We cannot forbear alluding to one of the species of Sphæria to which
Tulasne[m] attributes a variety of forms of fruit, and we do so here because
we think that a circumstance so extraordinary should be confirmed before it
is accepted as absolutely true. This refers to the common Sphæria found on
herbaceous plants, known as Sphæria (Pleospora) herbarum. First of all the
very common mould called Cladosporium herbarum is constituted as
conidia, and of this again Macrosporium sarcinula, Berk., is considered to
be another condition. In the next place, Cytispora orbicularis, Berk., and
Phoma herbarum, West., are regarded as pycnidia, enclosing stylospores.
Then Alternaria tenuis, Pr.,[n] which is said to be parasitic on Cladosporium
herbarum, is held to be only a form of that species, so that here we have
(including the perithecia) no less than six forms or phases for the same
fungus. As Macrosporium Cheiranthi, Pr., often is found in company with
Cladosporium herbarum, that is also open to suspicion.
We have adduced in the foregoing pages a few instances which will serve
to illustrate the polymorphism of fungi. Some of these it will be observed
are accepted as beyond doubt, occurring as they do in intimate relationship
with each other. Others are considered as scarcely so well established, but
probable, although developed sometimes on different species of plants.
Finally, some are regarded as hitherto not satisfactorily proved, or, it may
be, only suspicious. In this latter group, however much probability may be
in their favour, it can hardly be deemed philosophical to accept them on
such slender evidence as in some cases alone is afforded. It would not have
been difficult to have extended the latter group considerably by the addition
of instances enumerated by various mycologists in their works without any
explanation of the data upon which their conclusions have been founded. In

fact, altogether this chapter must be accepted as illustrative and suggestive,
but by no means as exhaustive.

[A]
De Bary, in “Quarterly German Magazine” (1872), p. 197.
[B]
The method pursued by Messrs. Berkeley and Hoffmann of surrounding the drop of
fluid, in which a definite number of spores or yeast globules had been placed, with a
pellicle of air, into which the germinating threads might pass and fructify, is perhaps
the most satisfactory that has been adopted, though it requires nice manipulation. If
carefully managed, the result is irrefragable, though doubts have been cast, without
any reason, on their observations.
[C]
De Bary, “Uber die Brandpilze” (Berlin, 1853), pl. iv. figs. 3, 4, 5.
[D]
A. de Bary, on Mildew and Fermentation, in “Quarterly German Magazine,” vol. ii.
1872.
[E]
Berkeley, “Introd. Crypt. Bot.” p. 78, fig. 20.
[F]
See also Berkeley, in “Trans. Hort. Soc. London,” vol. ix. p. 68.
[G]
Berkeley, in “Ann. Nat. Hist.” (June, 1838), No. 116.
[H]
“Grevillea,” vol. i. p. 176.
[I]

Tulasne, “On Certain Fungicolous Sphæriæ,” in “Ann. des Sci. Nat.” 4me sér. xiii.
(1860), p. 5.
[J]
“A Currant Twig, and Something on it,” in “Gardener’s Chronicle,” January 28, 1871.
[K]
Figs. 104 to 106 by permission from the “Gardener’s Chronicle.”
[L]
Berkeley and Broome, in “Annals of Natural History” (1866), No. 1177, pl. v. fig. 36;
Cooke, “Handbook,” ii. p. 866.
[M]

Cooke, “Handbook,” ii. p. 853, No. 2549; specimens in Cooke’s “Fungi Britannici
Exsiccati,” No. 270.
[N]
Berk. and Br. “Ann. Nat. Hist.” (1865), No. 1096.
[O]
“Ann. Nat. Hist.” (1871), No. 1332, pl. xx. fig. 23.
[P]
Ibid. No. 1333, pl. xxi. fig. 24.
[Q]
Tulasne, “Selecta Fungorum Carpologia,” ii. p. 269, pl. 29.
[R]
Cooke, “Handbook,” ii. p. 878; Tulasne, “Carpologia,” ii. p. 120, plate 14.
[S]
Tulasne, “Selecta Fung. Carp.,” ii. plate 16.
[T]
Corda, “Icones Fungorum,” vol. iii. fig. 91.
[U]
Corda, “Icones,” vol. i. fig. 25.
[V]
Berk. and Br. “Ann. Nat. Hist.” No. 415.
[W]
Currey, in “Philosoph. Trans. Roy. Soc.” (1857), pl. 25.
[X]
Tulasne, “On the Reproductive Apparatus of Fungi,” in “Comptes Rendus” (1852), p.
841; and Tulasne, “Selecta Fungorum Carpologia,” vol. iii.
[Y]
“Monatsbericht der Koniglichen Preuss, Acad. der Wissenschaften au Berlin,” Jan.
1865; Summary, in “Journ. Roy. Hort. Soc., London,” vol. i. n.s. p. 107.
[Z]
We have before us an Æcidium on leaves of Berberis vulgaris, collected at Berne by
Shuttleworth in 1833. It is named by him Æcidium graveolens, and differs in the
following particulars from Æcidium berberidis. The peridia are scattered as in Æ.
Epilobii, and not collected in clusters. They are not so much elongated. The cells are
larger, and the orange spores nearly twice the diameter. There is a decided, strong, but
unpleasant odour in the fresh plant; hence the name. The above figures (figs. 107,

108) of the cells and spores of both species are drawn by camera lucida to the same
scale—380 diameters.
[a]
Freiherrn von Hohenbühel-Heufler, in “Œsterr. Botan. Zeitschrift,” No. 3, 1870.
[b]
Fuckel, “Symbolæ Mycologicæ” (1869), p. 49.
[c]
Almost simultaneously with De Bary, the late Professor Œrsted instituted
experiments, from which the same results ensued, as to Æcidium berberidis and
Puccinia graminis. See “Journ. Hort. Soc. Lond.” new ser. i., p. 85.
[d]
“Oversigt over det Kon. Danske Videns. Selskabs” (1866), p. 185, t. 3, 4; (1867,) p.
208, t. 3, 4; “Résumé du Bulletin de la Soc. Roy. Danoise des Sciences” (1866), p. 15;
(1867), p. 38; “Botanische Zeitung” (1867), p. 104; “Quekett Microscopical Club
Journal,” vol. ii. p. 260.
[e]
This is Podisoma foliicola, B. and Br., or, as proposed in “Journ. Quekett Club,” ii. p.
267, Sarcostroma Berkeleyi, C.
[f]
Tulasne, “Selecta Fungorum Carpologia,” iii. p. 6, pl. i. figs. 19–31.
[g]
Cramer’s “Papilio Exotic” (1782), fig. 267.
[h]
Cooke, “Handbook,” p. 548, No. 1639.
[i]
Ibid. p. 556, No. 1666.
[j]
Specimens were published under this name in Cooke’s “Fungi Britannici Exsiccati,”
No. 359.
[k]
Cooke, “On Polymorphism in Fungi,” in “Popular Science Review.”
[l]
Lewis’s “Report on Microscopic Objects found in Cholera Evacuations,” Calcutta,
1870.
[m]

Tulasne, “Selecta Fungorum Carpologia,” ii. p. 261.
[n]
Corda, “Prachtflora,” plate vii.

X.
INFLUENCES AND EFFECTS.

It is no longer doubted that fungi exercise a large and very important
influence in the economy of nature. It may be that in some directions these
influences are exaggerated; but it is certain that on the whole their influence
is far more important for evil and for good than that of any other of the
Cryptogamia. In our endeavour to estimate the character and extent of these
influences it will prove advantageous to examine them under three sections.
1. Their influence on man. 2. Their influence on lower animals. 3. Their
influence on vegetation. Under these sections the chief facts may be
grouped, and some approximate idea obtained of the very great importance
of this family of inferior plants, and consequently the advisability of
pursuing their study more thoroughly and nationally than has hitherto been
done.
I. In estimating the influence of fungi upon man, we naturally enough
seek in the first instance to know what baneful effects they are capable of
producing on food. Although in the case of “poisonous fungi,” popularly
understood, fungi may be the passive agents, yet they cannot be ignored in
an inquiry of this nature. Writing of the Uses of Fungi, we have already
shown that a large number are available for food, and some of these real
delicacies; so, on the other hand, it becomes imperative, even with stronger
emphasis, to declare that many are poisonous, and some of them virulently
so. It is not sufficient to say that they are perfectly harmless until
voluntarily introduced into the human system, whilst it is well known that
accidents are always possible, and probably would be if every baneful
fungus had the word POISON inscribed in capitals on its pileus.
The inquiry is constantly being made as to what plain rules can be given
for distinguishing poisonous from edible fungi, and we can answer only that
there are none other than those which apply to flowering plants. How can
aconite, henbane, œnanthe, stramonium, and such plants, be distinguished

from parsley, sorrel, watercress, or spinach? Manifestly not by any general
characters, but by specific differences. And so it is with the fungi. We must
learn to discriminate Agaricus muscarius from Agaricus rubescens, in the
same manner as we would discriminate parsley from Æthusa cynapium.
Indeed, fungi have an advantage in this respect, since one or two general
cautions can be given, when none such are applicable for higher plants. For
instance, it may be said truly that all fungi that exhibit a rapid change to
blue when bruised or broken should be avoided; that all Agarics are open to
suspicion which possess an acrid taste; that fungi found growing on wood
should not be eaten unless the species is well known; that no species of
edible fungus has a strong, unpleasant odour, and similar cautions, which,
after all, are insufficient. The only safe guide lies in mastering, one by one,
the specific distinctions, and increasing the number of one’s own esculents
gradually, by dint of knowledge and experience, even as a child learns to
distinguish a filbert from an acorn, or with wider experience will thrust in
his mouth a leaf of Oxalis and reject that of the white clover.
One of the most deleterious of fungi that we possess is at the same time
one of the most beautiful. This is the Agaricus muscarius, or Fly Agaric,
which is sometimes used as a fly poison.[A] It has a bright crimson pileus
studded with pale whitish (sometimes yellowish) warts, and a stem and gills
of ivory whiteness. Many instances have been recorded of poisoning by this
fungus, and amongst them some British soldiers abroad, and yet it cannot
be doubted that this fungus is eaten in Russia. Two instances have come
under our notice of persons with some botanical knowledge, and one a
gardener, who had resided in Russia and eaten of this fungus. In one case
the Fly Agaric was collected and shown to us, and in the other the figure
was indicated, so that we might be under no doubt as to the species. Only
one hypothesis can be advanced in explanation. It is known that a large
number of fungi are eaten in Russia, and that they enter much into the
domestic cookery of the peasantry, but it is also known that they pay
considerable attention to the mode of cooking, and add a large amount of
salt and vinegar, both of which, with long boiling, must be powerful agents
in counteracting the poison (probably somewhat volatile) of such fungi as
the Fly Agaric. In this place we may give a recipe published by a French
author of a process for rendering poisonous fungi edible. It must be taken
on his authority, and not our own, as we have never made the experiment,
notwithstanding it seems somewhat feasible:—For each pound of

mushrooms, cut into moderately small pieces, take a quart of water
acidulated with two or three spoonfuls of vinegar, or two spoonfuls of bay
salt. Leave the mushrooms to macerate in the liquid for two hours, then
wash them with plenty of water; this done, put them in cold water and make
them boil. After a quarter or half hour’s boiling take them off and wash
them, then drain, and prepare them either as a special dish, or use them for
seasoning in the same manner as other species.[B]
This method is said to have been tried successfully with some of the most
dangerous kinds. Of these may be mentioned the emetic mushroom,
Russula emetica, with a bright red pileus and white gills, which has a clear,
waxy, tempting appearance, but which is so virulent that a small portion is
sufficient to produce disagreeable consequences. It would be safer to
eschew all fungi with a red or crimson pileus than to run the risk of
indulging in this. A white species, which, however, is not very common,
with a bulbous base enclosed in a volva, called Agaricus vernus, should
also be avoided. The pink spored species should also be regarded with
suspicion. Of the Boleti several turn blue when cut or broken, and these
again require to be discarded. This is especially the case with Boletus
luridus[C] and Boletus Satanas,[D] two species which have the under surface
or orifice of the pores of a vermilion or blood-red colour.
Not only are species which are known to be poisonous to be avoided, but
discretion should be used in eating recognized good species. Fungi undergo
chemical changes so rapidly that even the cultivated mushroom may cause
inconvenience if kept so long after being gathered as to undergo chemical
change. It is not enough that they should be of a good kind, but also fresh.
The employment of plenty of salt in their preparation is calculated very
much to neutralize any deleterious property. Salt, pepper, and vinegar are
much more freely employed abroad in preparing fungi than with us, and
with manifest advantage.
It is undoubtedly true that fungi exert an important influence in skin
diseases. This seems to be admitted on all hands by medical men,[E]
however much they may differ on the question of the extent to which they
are the cause or consequence of disease. Facts generally seem to bear out
the opinion that a great number of skin diseases are aggravated, and even
produced, by fungi. Robin[F] insists that a peculiar soil is necessary, and Dr.
Fox says it is usually taught that tuberculous, scrofulous, and dirty people

furnish the best nidus. It is scarcely necessary to enumerate all these
diseases, with which medical men are familiar, but simply to indicate a few.
There is favus or scall-head, called also “porrigo,” which has its primary
seat in the hair follicles. Plica polonica, which is endemic in Russia, is
almost cosmopolitan. Then there is Tinea tonsurans, Alopecia, Sycosis, &c.,
and in India a more deeply-seated disease, the Madura Foot, has been traced
to the ravages of a fungus described under the name of Chionyphe Carteri.
[G] It is probable that the application of different names to the very often
imperfect forms of fungi which are associated with different diseases is not
scientifically tenable. Perhaps one or two common moulds, such as
Aspergillus or Penicillium, lie at the base of the majority, but this is of little
importance here, and does not affect the general principle that some skin
diseases are due to fungi.
Whilst admitting that there are such diseases, it must be understood that
diseases have been attributed to fungi as a primary cause, when the
evidence does not warrant such a conclusion. Diphtheria and thrush have
been referred to the devastations of fungi, whereas diphtheria certainly may
and does occur without any trace of fungi. Fevers may sometimes be
accompanied by fungoid bodies in the evacuations, but it is very difficult to
determine them. The whole question of epidemic diseases being caused by
the presence of fungi seems based on most incomplete evidence. Dr.
Salisbury was of opinion that camp measles was produced by Puccinia
graminis, the pseudospores of which germinated in the damp straw,
disseminated the resultant secondary bodies in the air, and caused the
disease. This has never been verified. Measles, too, has been attributed
freely, as well as scarlatina,[H] to fungal influences, and the endeavours to
implicate fungi in being the cause of cholera have been pertinaciously
persevered in with no conviction. The presence of certain cysts, said to be
those of Urocystis, derived from rice, was announced by Dr. Hallier, but
when it was shown that no such fungus was found on rice, this phase of the
theory collapsed. Special and competent experts were sent from this country
to examine the preparations and hear the explanations of Dr. Hallier on his
theory of cholera contagion, but they were neither convinced nor satisfied.
As long ago as 1853, Dr. Lauder Lindsay examined and reported on
cholera evacuations, and in 1856 he declared—“It will be evident that I can
see no satisfactory groundwork for the fungus theory of cholera, which I am

not a little surprised to find still possesses powerful advocates.”[I] And of
the examinations undertaken by him he writes:—“The mycelium and
sporules of various species of fungi, constituting various forms of vegetable
mould, were found in the scum of the vomit, as well as of the stools, but
only at some stage of decomposition. They are found, however, under
similar circumstances, in the vomit and stools of other diseases, and,
indeed, in all decomposing animal fluids, and they are therefore far from
peculiar to cholera.”
Some writers have held that the atmosphere is often highly charged with
fungi spores, others have denied the presence of organic bodies to any
extent in the air. The experiments conducted in India by Dr. Cunningham[J]
have been convincing enough on this point. This report states that spores
and similar cells were of constant occurrence, and were generally present in
considerable numbers. That the majority of the cells were living and ready
to undergo development on meeting with suitable conditions was very
manifest, as in those cases in which preparations were retained under
observation for any length of time, germination rapidly took place in many
of the cells; indeed, many spores already germinating were deposited on the
slides. In few instances did any development take place beyond the
formation of mycelium or masses of toruloid cells, but in one or two
distinct sporules were developed on the filaments arising from some of the
larger septate spores, and in a few others Penicillium and Aspergillus
produced their characteristic heads of fructification.
With regard to the precise nature of the spores and other cells present in
various instances little can be said, as, unless their development were to be
carefully followed out through all its stages, it is impossible to refer them to
their correct species or even genera. The greater number of them are
apparently referable to the old orders of fungi—Sphæronemei, Melanconei,
Torulacei, Dematiei, and Mucedines, while some probably belonged to the
Pucciniei and Coæmacei. Amongst those belonging to the Torulacei, the
most interesting was a representative of the rare genus Tetraploa. Distinct
green algoid cells occurred in some specimens. Then follow in the report
details of observations made on the rise and fall of diseases, of which
diarrhœa, dysentery, cholera, ague, and dengue were selected and compared
with the increase or diminution of atmospheric cells. The conclusions
arrived at are:—

“Spores and other vegetable cells are constantly present in atmospheric
dust, and usually occur in considerable numbers; the majority of them are
living, and capable of growth and development. The amount of them
present in the air appears to be independent of conditions of velocity and
direction of the wind, and their number is not diminished by moisture.
“No connection can be traced between the numbers of bacteria, spores,
&c., present in the air, and the occurrence of diarrhœa, dysentery, cholera,
ague, or dengue, nor between the presence or abundance of any special
form or forms of cells, and the prevalence of any of these diseases.
“The amount of inorganic and amorphous particles and other débris
suspended in the atmosphere is directly dependent on conditions of
moisture and velocity of wind.”
This report is accompanied by fourteen large and well-executed plates,
each containing hundreds of figures of organic bodies collected from the air
between February and September. It is valuable both for its evidence as to
the number and character of the spores in the air, and also for the tables
showing the relation between five forms of disease, and their fluctuations,
as compared with the amount of spores floating in the atmosphere.
We are fain to believe that we have represented the influence of fungi on
man as far as evidence seems to warrant. The presence of forms of mould in
some of their incipient conditions in different diseased parts of the human
body, externally and internally, may be admitted without the assumption
that they are in any manner the cause of the diseased tissues, except in such
cases as we have indicated. Hospital gangrene may be alluded to in this
connection, and it is possible that it may be due to some fungus allied to the
crimson spots (blood rain) which occur on decayed vegetation and meat in
an incipient stage of decomposition. This fungus was at one time regarded
as an algal, at another as animal; but it is much more probable that it is a
low condition of some common mould. The readiness with which the spores
of fungi floating in the atmosphere adhere to and establish themselves on all
putrid or corrupt substances is manifest in the experience of all who have
had to do with the dressing of wounds, and in this case it is a matter of the
greatest importance that, as much as possible, atmospherical contact should
be avoided.

Recently a case occurred at the Botanic Gardens at Edinburgh which was
somewhat novel. The assistant to the botanical professor was preparing for
demonstration some dried specimens of a large puff-ball, filled with the
dust-like spores, which he accidentally inhaled, and was for some time
confined to his room under medical attendance from the irritation they
caused. This would seem to prove that the spores of some fungi are liable,
when inhaled in large quantities, to derange the system and become
dangerous; but under usual and natural conditions such spores are not likely
to be present in the atmosphere in sufficient quantity to cause
inconvenience. In the autumn a very large number of basidiospores must be
present in the atmosphere of woods, and yet there is no reason to believe
that it is more unhealthy to breathe the atmosphere of a wood in September
or October than in January or May. Dreadful effects are said to be produced
by a species of black rust which attacks the large South of Europe reed,
Arundo donax. This is in all probability the same species with that which
attacks Arundo phragmitis in this country, the spores of which produce
violent headaches and other disorders amongst the labourers who cut the
reeds for thatching. M. Michel states that the spores from the parasite on
Arundo donax, either inhaled or injected, produce violent papular eruption
on the face, attended with great swelling, and a variety of alarming
symptoms which it is unnecessary to particularize, in various parts of the
body.[K] Perhaps if Sarcina should ultimately prove to be a fungus, it may
be added to the list of those which aggravate, if they are not the primary
cause of, disease in the human subject.
II. What influences can be attributed to fungi upon animals other than
man? Clearly instinct preserves animals from many dangers. It may be
presumed that under ordinary circumstances there is not much fear of a cow
or a sheep poisoning itself in a pasture or a wood. But under extraordinary
circumstances it is not only possible, but very probable, that injuries may
occur. For instance, it is well known that not only rye and wheat, but also
many of the grasses, are liable to infection from a peculiar form of fungus
called “ergot.” In certain seasons this ergot is much more common than
others, and the belief is strong in those who ought to know something of the
subject from experience, viz., farmers and graziers, that in such seasons it is
not uncommon for cattle to slip their young through feeding on ergotized
grass. Then, again, it is fairly open to inquiry whether, in years when “red
rust” and “mildew” are more than usually plentiful on grasses, these may

not be to a certain extent injurious. Without attempting to associate the
cattle plague in any way with fungi on grass, it is nevertheless a most
remarkable coincidence that the year in which the cattle disease was most
prevalent in this country was one in which there was—at least in some
districts—more “red rust” on grasses than we ever remember to have seen
before or since; the clothes of a person walking through the rusty field soon
became orange-coloured from the abundance of spores. Graziers on this
point again seem to be generally agreed, that they do not think “red rust”
has been proved to be injurious to cattle. The direct influence of fungi on
quadrupeds, birds, reptilia, &c., seems to be infinitesimally small.
Insects of various orders have been observed from time to time to
become the prey of fungi.[L] That known at Guadaloupe under the name of
La Guêpe Végétale, or vegetable wasp, has been often cited as evidence
that, in some instances at least, the fungus attacks the insect whilst still
living. Dr. Madianna states that he has noticed the wasp still living with its
incumbrance attached to it, though apparently in the last stage of existence,
and seeming about to perish from the influence of its destructive parasite.
[M] This fungus is called by Tulasne Torrubia sphecocephala.[N] About
twenty-five species of this genus of sphæriaceous fungi have been
described as parasitic on insects. Five species are recorded in South
Carolina, one in Pennsylvania, found on the larvæ of the May-bug, and one
other North American species on Nocturnal Lepidoptera, one in Cayenne,
one in Brazil, on the larva of a Cicada, and one on a species of ant, two in
the West Indies, one in New Guinea on a species of Coccus, and one on a
species of Vespa in Senegal. In Australia two species have been recorded,
and two are natives of New Zealand. Dr. Hooker found two in the Khassya
mountains of India, and one American species has also been found at
Darjeeling. It has long been known that one species, which has a medicinal
repute there, is found in China, whilst three have been recorded in Great
Britain. Opinions are divided as to whether in these instances the fungus
causes or is subsequent to the death of the insect. It is generally the belief of
entomologists that the death of the insect is caused by the fungus. In the
case of Isaria sphingum, which is the conidia form of a species of Torrubia,
the moth has been found standing on a leaf, as during life, with the fungus
sprouting from its body.

Other and less perfect forms of fungi also attack insects. During the
summer of 1826, Professor Sebert collected a great many caterpillars of
Arctia villica, for the purpose of watching their growth. These insects on
arriving at their full size became quite soft, and then suddenly died. Soon
after they became hard, and, if bent, would easily break into two pieces.
Their bodies were covered with a beautiful shining white mould. If some of
the caterpillars affected with the parasitic mould were placed on the same
tree with those apparently free from its attack, the latter soon exhibited
signs that they also were attacked in the same manner, in consequence of
coming into contact with each other.[O]
During the spring of 1851, some twelve or twenty specimens were found
from amongst myriads of Cicada septemdecim, which, though living, had
the posterior third of the abdominal contents converted into a dry, powdery,
ochreous-yellow compact mass of sporuloid bodies. The outer coverings of
that portion of the insect were loose and easily detached, leaving the
fungoid matter in the form of a cone affixed by its base to the unaffected
part of the abdomen of the insect. The fungus may commence, says Dr.
Leidy, its attacks upon the larva, develop its mycelium, and produce a
sporular mass within the active pupa, when many are probably destroyed;
but should some be only affected so far as not to destroy the organs
immediately essential to life, they might undergo their metamorphosis into
the imago, in which case they would be affected in the manner previously
described.[P]
The common house-fly in autumn is very usually subject to the attacks of
a mouldy fungus called Sporendonema muscæ, or Empusa muscæ in former
times, which is now regarded as the terrestrial condition of one of the
Saprolegniei.[Q] The flies become sluggish, and at last fix themselves to
some object on which they die, with their legs extended and head
depressed, the body and wings soon becoming covered with a minute white
mould, the joints of which fall on the surrounding object. Examples are
readily distinguished when they settle on windows and thus succumb to
their foe. Mr. Gray says that a similar mould has been observed on
individuals of the wasp family.
A Gryllotalpa was found in a wood near Newark, Delaware, U. S., upon
turning over a log. The insect was seen standing very quietly at the mouth
of its oval cell, which is formed in the earth, having a short curved tube to

the surface. Upon taking it up it exhibited no signs of movement, though
perfectly fresh and lifelike in appearance. On examining it next morning it
still presented no signs of life. Every part of the insect was perfect, not even
the antennæ being broken. Upon feeling it, it was very hard and resistant,
and on making an incision through the thorax it exhaled a fungoid odour.
The insect had been invaded by a parasitic fungus which everywhere filled
the animal, occupying the position of all the soft tissue, and extending even
into the tarsal joints. It formed a yellowish or cream-coloured compact
mass.[R]
The destructive silk-worm disease, Botrytis Bassiana, is also a fungus
which attacks and destroys the living insect, concerning which an immense
deal has been written, but which has not yet been eradicated. It has also
been supposed that a low form or imperfect condition of a mould has much
to do with the disease of bees known as “foul brood.”[S]
Penicillium Fieberi, figured by Corda on a beetle, was doubtless
developed entirely after death, with which event it had probably nothing
whatever to do.[T] Sufficient, however, has been written to show that fungi
have an influence on insect life, and this might be extended to other animal
forms, as to spiders, on which one or two species of Isaria are developed,
whilst Dr. Leidy has recorded observations on Julus[U] which may be
perused with advantage. Fish are subject to a mouldy-looking parasite
belonging to the Saprolegniei, and a similar form attacks the ova of toads
and frogs. Gold fish in globes and aquaria are very subject to attack from
this mouldy enemy, and although we have seen them recover under a
constant change of water, this is by no means always the case, for in a few
weeks the parasite will usually prevail.
The influence of fungi upon animals in countries other than European is
very little known, except in the case of the species of Torrubia found on
insects, and the diseases to which silkworms are subject. Instances have
been recorded of the occurrence of fungoid mycelium—for in most it is
nothing more—in the tissues of animals, in the hard structure of bone and
shell, in the intestines, lungs, and other fleshy parts, and in various organs
of birds.[V] In some of the latter cases it has been described as a Mucor, in
most it is merely cells without sufficient character for determination. It is by
no means improbable that fungi may be found in such situations; the only
question with regard to them is whether they are not accidental, and not the

producers of unhealthy or diseased tissues, even when found in proximity
thereto.
There is one phase of the influences of fungi on the lower animals which
must not be wholly passed over, and that is the relation which they bear to
some of the insect tribes in furnishing them with food. It is especially the
case with the Coleoptera that many species seem to be entirely dependent
on fungi for existence, since they are found in no other situations. Beetle-
hunters tell us that old Polyporei, and similar fungi of a corky or woody
nature, are always sought after for certain species which they seek in vain
elsewhere,[W] and those who possess herbaria know how destructive certain
minute members of the animal kingdom are to their choicest specimens,
against whose depredations even poison is sometimes unavailing.
Some of the Uredines, as Trichobasis suaveolens and Coleosporium
sonchi, are generally accompanied by a little orange larva which preys upon
the fungus; and in the United States Dr. Bolles informs us that some species
of Æcidium are so constantly infested with this red larva that it is scarcely
possible to get a good specimen, or to keep it from its sworn enemy. Minute
Anguillidæ revel in tufts of mould, and fleshy Agarics, as they pass into
decay, become colonies of insect life. Small Lepidoptera, belonging to the
Tineina, appear to have a liking for such Polyporei as P. sulfureus when it
becomes dry and hard, or P. squamosus when it has attained a similar
condition. Acari and Psocidæ attack dried fungi of all kinds, and speedily
reduce them to an unrecognizable powder.
III. What are the influences exerted by fungi on other plants? This is a
broad subject, but withal an important one, since these influences act
indirectly on man as well as on the lower animals. On man, inasmuch as it
interferes with the vegetable portion of his food, either by checking its
production or depreciating its quality. On the lower animals, since by this
means not only is their natural food deteriorated or diminished, but through
it injurious effects are liable to be produced by the introduction of minute
fungi into the system. These remarks apply mainly to fungi which are
parasitic on living plants. On the other hand, the influence of fungi must not
be lost sight of as the scavengers of nature when dealing with dead and
decaying vegetable matter. Therefore, as in other instances, we have here
also good and bad influences intermingled, so that it cannot be said that
they are wholly evil, or unmixed good.

Wherever we encounter decaying vegetable matter we meet with fungi,
living upon and at the expense of decay, appropriating the changed
elements of previous vegetable life to the support of a new generation, and
hastening disintegration and assimilation with the soil. No one can have
observed the mycelium of fungi at work on old stumps, twigs, and decayed
wood, without being struck with the rapidity and certainty with which
disintegration is being carried on. The gardener casts on one side, in a pile
as rubbish, twigs and cuttings from his trees, which are useless to him, but
which have all derived much from the soil on which they flourished.
Shortly fungi make their appearance in species almost innumerable, sending
their subtle threads of mycelium deep into the tissues of the woody
substance, and the whole mass teems with new life. In this metamorphosis
as the fungi flourish so the twigs decay, for the new life is supported at the
expense of the old, and together the destroyers and their victims return as
useful constituents to the soil from whence they were derived, and form
fresh pabulum for a succeeding season of green leaves and sweet flowers.
In woods and forests we can even more readily appreciate the good offices
of fungi in accelerating the decay of fallen leaves and twigs which surround
the base of the parent trees. In such places Nature is left absolutely to her
own resources, and what man would accomplish in his carefully attended
gardens and shrubberies must here be done without his aid. What we call
decay is merely change; change of form, change of relationship, change of
composition; and all these changes are effected by various combined
agencies—water, air, light, heat, these furnishing new and suitable
conditions for the development of a new race of vegetables. These, by their
vigorous growth, continue what water and oxygen, stimulated by light and
heat, had begun, and as they flourish for a brief season on the fallen glories
of the past summer, make preparation for the coming spring.
Unfortunately this destructive power of fungi over vegetable tissues is
too often exemplified in a manner which man does not approve. The dry rot
is a name which has been given to the ravages of more than one species of
fungus which flourishes at the expense of the timber it destroys. One of
these forms of dry rot fungus is Merulius lacrymans, which is sometimes
spoken of as if it were the only one, though perhaps the most destructive in
houses. Another is Polyporus hybridus, which attacks oak-built vessels;[X]
and these are not the only ones which are capable of mischief. It appears
that the dry rot fungus acts indirectly on the wood, whose cells are saturated

with its juice, and in consequence lose their lignine and cellulose, though
their walls suffer no corrosion. The different forms of decay in wood are
accompanied by fungi, which either completely destroy the tissue, or alter
its nature so much by the abstraction of the cellulose and lignine, that it
becomes loose and friable. Thus fungi induce the rapid destruction of
decaying wood. These are the conclusions determined by Schacht, in his
memoir on the subject.[Y]
We may allude, in passing, to another phase of destructiveness in the
mycelium of fungi, which traverse the soil and interfere most injuriously
with the growth of shrubs and trees. The reader of journals devoted to
horticulture will not fail to notice the constant appeals for advice to stop the
work of fungi in the soil, which sometimes threatens vines, at others
conifers, and at others rhododendrons. Dead leaves, and other vegetable
substances, not thoroughly and completely decayed, are almost sure to
introduce this unwelcome element.
Living plants suffer considerably from the predations of parasitic species,
and foremost amongst these in importance are those which attack the
cereals. The corn mildew and its accompanying rust are cosmopolitan, as
far as we know, wherever corn is cultivated, whether in Australia or on the
slopes of the Himalayas. The same may also be said of smut, for Ustilago is
as common in Asia and America as in Europe. We have seen it on numerous
grasses as well as on barley from the Punjab, and a species different from
Ustilago maydis on the male florets of maize from the same locality. In
addition to this, we learn that in 1870 one form made its appearance on rice.
It was described as constituting in some of the infested grains a whitish,
gummy, interlaced, ill-defined, thread-like mycelium, growing at the
expense of the tissues of the affected organs, and at last becoming
converted into a more or less coherent mass of spores, of a dirty green
colour, on the exterior of the deformed grains. Beneath the outer coating the
aggregated spores are of a bright orange red; the central portion has a
vesicular appearance, and is white in colour.[Z] It is difficult to determine
from the description what this so-called Ustilago may be, which was said to
have affected a considerable portion of the standing rice crop in the vicinity
of Diamond Harbour.
Bunt is another pest (Tilletia caries) which occupies the whole
farinaceous portion of the grains of wheat. Since dressing the seed wheat

has been so widely adopted in this country, this pest has been of
comparatively little trouble. Sorghum and the small millets, in countries
where these are cultivated for food, are liable to attacks from allied
parasites. Ergot attacks wheat and rice as well as rye, but not to such an
extent as to have any important influence upon the crop. Two or three other
species of fungi are sometimes locally troublesome, as Dilophospora
graminis, and Septoria nodorum on wheat, but not to any considerable
extent. In countries where maize is extensively grown it has not only its
own species of mildew (Puccinia), but also one of the most enormous and
destructive species of Ustilago.
A singular parasite on grasses was found by Cesati in Italy, in 1850,
infesting the glumes of Andropogon.[a] It received the name of Cerebella
Andropogonis, but it never appears to have increased and spread to such an
extent as was at first feared.
Even more destructive than any of these is the potato disease[b]
(Peronospora infestans), which is, unfortunately, too well known to need
description. This disease was at one time attributed to various causes, but
long since its ascertained source has been acknowledged to be a species of
white mould, which also attacks tomatoes, but less vigorously. De Bary has
given considerable attention to this disease, and his opinions are clearly
detailed in his memoir on Peronospora, as well as in his special pamphlet
on the potato disease.[c] One sees the cause of the epidemic, he says, in the
diseased state of the potato itself, produced either accidentally by
unfavourable conditions of soil and atmosphere, or by a depravation that the
plant has experienced in its culture. According to these opinions, the
vegetation of the parasite would be purely accidental, the disease would be
independent of it, the parasite would be able frequently even to spare the
diseased organs. Others see in the vegetation of the Peronospora the
immediate or indirect cause of the various symptoms of the disease; either
that the parasite invades the stalks of the potato, and in destroying them, or,
so to speak, in poisoning them, determines a diseased state of the tubercles,
or that it introduces itself into all the organs of the plant, and that its
vegetation is the immediate cause of all the symptoms of the disease that
one meets with in any organ whatever. His observations rigorously proved
that the opinions of the latter were those only which were well founded. All
the alterations seen on examining spontaneous individuals are found when

the Peronospora is sown in a nourishing plant. The most scrupulous
examination demonstrates the most perfect identity between the cultivated
and spontaneous individuals as much in the organization of the parasite as
in the alteration of the plant that nourishes it. In the experiments that he had
made he affirms that he never observed an individual or unhealthy
predisposition of the nourishing plant. It appeared to him, on the contrary,
that the more the plant was healthy, the more the mould prospered.
We cannot follow him through all the details of the growth and
development of the disease, or of his experiments on this and allied species,
which resulted in the affirmation that the mould immediately determines the
disease of the tubercles as well as that of the leaves, and that the vegetation
of the Peronospora alone determines the redoubtable epidemic to which the
potato is exposed.[d] We believe that this same observer is still engaged in a
series of observations, with the view, if possible, of suggesting some
remedy or mitigation of the disease.
Dr. Hassall pointed out, many years since, the action of fungous
mycelium, when coming in contact with cellular tissue, of inducing
decomposition, a fact which has been fully confirmed by Berkeley.
Unfortunately there are other species of the same genus of moulds which
are very destructive to garden produce. Peronospora gangliformis, B.,
attacks lettuces, and is but too common and injurious. Peronospora effusa,
Grev., is found on spinach and allied plants. Peronospora Schleideniana, D.
By., is in some years very common and destructive to young onions, and
field crops of lucerne are very liable to attack from Peronospora
trifoliorum, D. By.
The vine crops are liable to be seriously affected by a species of mould,
which is but the conidia form of a species of Erysiphe. This mould, known
under the name of Oidium Tuckeri, B., attacks the vines in hothouses in this
country, but on the Continent the vineyards often suffer severely[e] from its
depredations; unfortunately, not the only pest to which the vine is subject,
for an insect threatens to be even more destructive.
Hop gardens suffer severely, in some years, from a similar disease; in this
instance the mature or ultimate form is perfected. The hop mildew is
Sphærotheca Castagnei, Lév., which first appears as whitish mouldy
blotches on the leaves, soon becoming discoloured, and developing the

black receptacles on either surface of the leaf. These may be regarded as the
cardinal diseases of fungoid origin to which useful plants are subject in this
country.
Amongst those of less importance, but still troublesome enough to secure
the anathemas of cultivators, may be mentioned Puccinia Apii, Ca., often
successful in spoiling beds of celery by attacking the leaves; Cystopus
candidus, Lév., and Glæosporium concentricum, Grev., destructive to
cabbages and other cruciferous plants; Trichobasis Fabæ, Lév., unsparing
when once established on beans; Erysiphe Martii, Lév., in some seasons a
great nuisance to the crop of peas.
Fruit trees do not wholly escape, for Rœstelia cancellata, Tul., attacks the
leaves of the pear. Puccinia prunorum affects the leaves of almost all the
varieties of plum. Blisters caused by Ascomyces deformans, B., contort the
leaves of peaches, as Ascomyces bullatus, B., does those of the pear, and
Ascomyces juglandis, B., those of the walnut. Happily we do not at present
suffer from Ascomyces pruni, Fchl., which, on the Continent, attacks young
plum-fruits, causing them to shrivel and fall. During the past year pear-
blossoms have suffered from what seems to be a form of Helminthosporium
pyrorum, and the branches are sometimes infected with Capnodium
elongatum; but orchards in the United States have a worse foe in the “black
knot,”[f] which causes gouty swellings in the branches, and is caused by the
Sphæria morbosa of Schweinitz.
Cotton plants in India[g] were described by Dr. Shortt as subject to the
attacks of a kind of mildew, which from the description appeared to be a
species of Erysiphe, but on receiving specimens from India for
examination, we found it to be one of those diseased conditions of tissue
formerly classed with fungi under the name of Erineum; and a species of
Torula attacks cotton pods after they are ripe. Tea leaves in plantations in
Cachar have been said to suffer from some sort of blight, but in all that we
have seen insects appear to be the depredators, although on the decaying
leaves Hendersonia theicola, Cooke, establishes itself.[h] The coffee
plantations of Ceylon suffer from the depredations of Hemiliea vastatrix, as
well as from insects.[i] Other useful plants have also their enemies in
parasitic fungi.

Olive-trees in the south of Europe suffer from the attacks of a species of
Antennaria, as do also orange and lemon trees from a Capnodium, which
covers the foliage as if with a coating of soot. In fact most useful plants
appear to have some enemy to contend with, and it is fortunate, not only for
the plant, but its cultivators, if this enemy is less exacting than is the case
with the potato, the vine, and the hop.
Forestry in Britain is an insignificant interest compared to what it is in
some parts of Europe, in the United States, and in our Indian possessions. In
these latter places it becomes a matter of importance to inquire what
influence fungi exert on forest trees. It may, however, be predicated that the
injury caused by fungi is far outstripped by insects, and that there are not
many fungi which become pests in such situations. Coniferous trees may be
infested with the species of Peridermium, which are undoubtedly injurious,
Peridermium elatinum, Lk., distorting and disfiguring the silver fir, as
Peridermium Thomsoni, B.,[j] does those of Abies Smithiana in the
Himalayas. This species occurred at an elevation of 8,000 feet. The leaves
become reduced in length one-half, curved, and sprinkled, sometimes in
double rows, with the large sori of this species, which gives the tree a
strange appearance, and at length proves fatal, from the immense diversion
of nutriment requisite to support a parasite so large and multitudinous. The
dried specimens have a sweet scent resembling violets. In Northern Europe
Cæoma pinitorquum, D. By., seems to be plentiful and destructive. All
species of juniper, both in Europe and the United States, are liable to be
attacked and distorted by species of Podisoma[k] and Gymnosporangium.
Antennaria pinophila, Fr., is undoubtedly injurious, as also are other species
of Antennaria, which probably attain their more complete development in
Capnodium, of which Capnodium Citri is troublesome to orange-trees in
the south of Europe, and other species to other trees. How far birch-trees are
injured by Dothidea betulina, Fr., or Melampsora betulina, Lév., or poplars
and aspens by Melampsora populina, Lév., and Melampsora tremulæ, Lév.,
we cannot say. The species of Lecythea found on willow leaves have
decidedly a prejudicial effect on the growth of the affected plant.
Floriculture has to contend with many fungoid enemies, which
sometimes commit great ravages amongst the choicest flowers. Roses have
to contend against the two forms of Phragmidium mucronatum as well as
Asteroma Rosæ. Still more disastrous is a species of Erysiphei, which at

first appears like a dense white mould. This is named Sphærotheca
pannosa. Nor is this all, for Peronospora sparsa, when it attacks roses in
conservatories, is merciless in its exactions.[l] Sometimes violets will be
distorted and spoiled by Urocystis Violæ. The garden anemone is freely
attacked by Æcidium quadrifidum. Orchids are liable to spot from fungi on
the leaves, and recently the whole of the choicest hollyhocks have been
threatened with destruction by a merciless foe in Puccinia malvacearum.
This fungus was first made known to the world as an inhabitant of South
America many years ago. It seems next to have come into notoriety in the
Australian colonies. Then two or three years ago we hear of it for the first
time on the continent of Europe, and last year for the first time in any
threatening form in our own islands. During the present year its ravages are
spreading, until all admirers of hollyhocks begin to feel alarm lest it should
entirely exterminate the hollyhock from cultivation. It is common on wild
mallows, and cotton cultivators must be on the alert, for there is a
probability that other malvaceous plants may suffer.
A writer in the “Gardener’s Chronicle” has proposed a remedy for the
hollyhock disease, which he hopes will prove effectual. He says, “This
terrible disease has now, for twelve months, threatened the complete
annihilation of the glorious family of hollyhock, and to baffle all the
antidotes that the ingenuity of man could suggest, so rapidly does it spread
and accomplish its deadly work. Of this I have had very sad evidence, as
last year at this time I had charge of, if not the largest, one of the largest and
finest collections of hollyhocks anywhere in cultivation, which had been
under my special care for eleven years, and up to within a month of my
resigning that position I had observed nothing uncommon amongst them;
but before taking my final leave of them I had to witness the melancholy
spectacle of bed after bed being smitten down, and amongst them many
splendid seedlings, which had cost me years of patience and anxiety to
produce. And again, upon taking a share and the management of this
business, another infected collection fell to my lot, so that I have been doing
earnest battle with this disease since its first appearance amongst us, and I
must confess that, up to a very short time back, I had come in for a great
deal the worst of the fight, although I had made use of every agent I could
imagine as being likely to aid me, and all that many competent friends
could suggest. But lately I was reminded of Condy’s patent fluid, diluted
with water, and at once procured a bottle of the green quality, and applied it

in the proportion of a large tablespoonful to one quart of water, and upon
examining the plants dressed, twelve hours afterwards, was delighted to
find it had effectually destroyed the disease (which is easily discernible, as
when it is living and thriving it is of a light grey colour, but when killed it
becomes of a rusty black). Further to test the power at which the plant was
capable of bearing the antidote without injury, I used it double the strength.
This dose was instant death to the pest, leaving no trace of any injury to the
foliage. As to its application, I advocate sponging in all dressings of this
description. Syringing is a very ready means, but very wasteful. No doubt
sponging consumes more time, but taking into consideration the more
effectual manner in which the dressing can be executed alone, it is in the
end most economical, especially in regard to this little parasite. I have
found it difficult by syringing, as it has great power of resisting and
throwing off moisture, and if but a very few are left living, it is astonishing
how quickly it redistributes itself. I feel confident, that by the application of
this remedy in time another season, I shall keep this collection clean. I
believe planting the hollyhock in large crowded beds should be avoided, as
I have observed the closer they are growing the more virulently does the
disease attack them, whereas isolated rows and plants are but little
injured.”[m]
The “Gardener’s Chronicle” has also sounded a note of warning that a
species of Uredine has been very destructive to pelargoniums at the Cape of
Good Hope. Hitherto these plants have not suffered much in this country
from parasites. Besides these, there are many other less troublesome
parasites, such as Uredo filicum, on ferns; Puccinia Lychnidearum, on
leaves of sweet-william; Uredo Orchidis, on leaves of orchids, &c.
If we would sum up the influences of fungi in a few words, it could be
done somewhat in the following form.
Fungi exert a deleterious influence—

On Man,
When eaten inadvertently.
By the destruction of his legitimate food.
In producing or aggravating skin diseases.

On Animals,

By deteriorating or diminishing their food supplies.
By establishing themselves as parasites on some species.

On Plants,
By hastening the decay of timber.
By establishing themselves as parasites.
By impregnating the soil.

But it is not proved that they produce epidemic diseases in man or
animals, or that the dissemination of their multitudinous spores in the
atmosphere has any appreciable influence on the health of the human race.
Hence their association with cholera, diarrhœa, measles, scarlatina, and the
manifold ills that flesh is heir to, as producing or aggravating causes, must,
in the present state of our knowledge and experience, be deemed
apocryphal.
[A]
A detailed account of the peculiar properties of this fungus and its employment as a
narcotic will be found in Cooke’s “Seven Sisters of Sleep,” p. 337. It is figured in
Greville’s “Scottish Cryptogamic Flora,” plate 54.
[B]
Pour chaque 500 grammes de champignons coupes en morceaux d’assez mediocre
grandeur, il faut un litre d’eau acidulée par deux ou trois cuillerées de vinaigre, ou
deux cuillerées de sel gris. Dans le cas ou l’on n’aurait que de l’eau à sa disposition, il
faut la renouveler une ou deux fois. On laisse les champignons macérer dans le
liquids pendant deux heures entières, puis on les lave à grande eau. Ils sont alors mis
dans de l’eau froide qu’on porte à l‘ébullition, et après un quart d’heure ou une demi-
heure, on les retire, on les lave, on les essuie, et ou les apprête soit comme un mets
spécial, et ils comportent les mêmes assaisonnements que les autres, soit comme
condiment.—Morel Traité des Champignons, p. lix. Paris, 1865.
[C]
Smith’s “Chart of Poisonous Fungi,” fig. 10.
[D]
Ibid. fig. 27. It would be well to become acquainted with all these figures.
[E]
“Skin Diseases of Parasitic Origin,” by Dr. Tilbury Fox. London, 1863.
[F]
Robin, “Hist. Nat. des Végétaux Parasites.” Paris, 1853. Kuchenmeister, “Animal and
Vegetable Parasites of the Human Body.” London, Sydenham Society, 1857.