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Preface

This book has been prepared in an attempt to fill a long-standing


gap in the series of semi-popular illustrated books available to British
naturalists. Several fairly up-to-date volumes are available to help
those interested in the larger Basidiomycetes but no-one has attempted
a comprehensive account of the Ascomycetes in English since the
appearance of M. C. Cooke's 'Handbook of British Fungi, with full
descriptions of all the Species and Illustrations of the Genera' in 1871.
Cooke recognised 95 Ascomycete genera, some very large; his Peziza
with 162 species included not only the present Pezizales but the
greater part of the Helotiales as well; his Sphaeria with 193 species
covered most of non-stromatic Sphaeriales and Loculoascomycetes.
In 1895 George Massee dealt comprehensively with the Pezizales,
Helotiales, non-lichenised Lecanorales and Ostropales in the fourth
and last volume of his 'British Fungus Flora'. In this restricted field
he recognised 112 genera, most of which are retained in the present
work. Massee's flora has been much ridiculed but his critics have been
careful not to expose themselves by attempting a better work to re-
place it. After 60 years his is still the only English book available for
identification of these groups of Ascomycetes. In fact his descriptions
are usually graphic and accurate and will bear comparison with the
best published by contemporary continental authors. The system he
adopted was, however, already obsolete in t h a t he rejected the dis-
tinction between Pezizales and Helotiales, based on the presence or
absence of an operculum in the ascus, "as the character can only be
distinctly observed in the fresh ascus" and his genus Humaria in-
cluded both operculate and inoperculate species. Unfortunately a com-
panion volume on the Sphaeriales and Loculoascomycetes was never
issued and for these fungi British mycologists have remained de-
pendent on the comprehensive German floras of Winter, Migula (1913)
and Kirschstein, the last never completed.
It is at present quite impracticable to compile a critical British
Ascomycete flora, largely because of the lack of collections on which
such a work could be soundly based. Too often species have been
briefly reported as occurring in the British Isles on the strength of a
single collection, which has not been kept or the fate of which is now
uncertain. Nearly all the larger genera require critical revision before
one can have a clear idea how many of the described species are mor-

IX
phologically distinct or even belong in the genus to which they have
been assigned. Most of the genera said to occur in Britain are illustrated
in this book. Some are omitted because their ascocarps are too in-
conspicuous or too ill-defined to be likely to come to the attention of
the amateur naturalist. A few have been omitted because there was
no suitable material available to be figured or because their presence
in the British Isles seemed to call for confirmation. Some old generic
names will be found to be absent because they have been used in a
very wide sense and it is not clear which of the original species should be
selected as lectotype to define the use of the name in the modern
system.
From the standpoint of the Ascomycetes the British Isles are still
only very imperfectly explored. Many collections have been made
over the past century and a half in the Home Counties and in scattered
districts fortunate enough to have possessed a resident mycologist at
some time. The Highland areas still remain almost untouched and so
do some distinctive habitats elsewhere; a "rare" fungus is often found
to be common in some narrowly circumscribed habitat. As Bisby
(1953) has justly said "The Ascomycetes are a vast and confusing
assemblage. . . . T h e y are a challenge to an army of students". So far
the army wanting. If this book helps to recruit for it a few volunteers
it will not have been written in vain.
THE HERBARIUM,

January 1956. ROYAL BOTANIC GARDENS, KEW.

Preface to the second edition


The last decade has seen great activity in Ascomycete studies and
in preparing a second edition I have endeavoured to incorporate as
much new material as possible within the old framework and to elim-
inate the grosser errors of the first. In deference to Dr. Grove's
opinion Scleroderris and Crumenula have been united under Godronia
while publication of Dr. Booth's critical studies of Nectria has made
it possible greatly to expand the treatment of t h a t economically im-
portant genus, with elimination of the segregates Dialonectria and
Lasionectria. The British species of Leptosphaeria have also been more
adequately covered though without a complete rewritting of the
Pleosporaceae it has not been possible to take full advantage of Dr.
Holm's revision of this and allied genera. In view of their abundance
on a few common British plants it has seemed best to include the
Protomycetaceae even though they produce no ascocarps. There are

x
unfortunately many name changes, some for taxonomic reasons, in-
cluding synonymy with older genera, others on purely nomenclatorial
grounds, like the substitution of Cudoniella and Hymenoscyphus for
Helotium.
Twentyeight additional genera and 154 additional species are de-
scribed while references to many others, including a few found in
Europe but not as yet in Britain, have been inserted in the keys or as
brief notes in the text. In order to retain the number of 40 coloured
plates, convenient for the printer, six of the plates of the first edition
have been withdrawn and replaced by six new ones to figure especially
Protomycetaceae, Taphrinaceae, Helotiaceae and Nectria. There are
eleven new line drawings of which six replace the lost coloured plates
and five are entirely new, three of them devoted to Leptosphaeria.
THE HERBARIUM,

24th June 1967 ROYAL BOTANIC GARDENS, KEW.

XI
INTRODUCTION
In all but the simplest fungi, the plant body consists of very slender
much branched threads, individually called hyphae, collectively the
mycelium. Reproduction takes place by the production of microscopic
bodies adapted for dissemination by air currents, less often by water
or by insects. These are of many kinds but all differ from seeds in not
containing a differentiated embryo and are conveniently referred to
by the comprehensive term "spores". Fungi cannot be named from
their mycelia alone and all but the broadest groupings of the species
are based on the nature of the spores and the organs in or on which
they are borne.
One group of fungi, the Phycomycetes, can be recognised by its
mycelium, which lacks transverse walls or septa. Only a very few of
these produce complex fruit bodies; the majority are plant parasites
(Downy mildews) or microscopic moulds inhabiting water or soil.
None of them is dealt with in this book.
The remaining fungi have mycelia composed of septate hyphae and
fall into three main classes according to the manner in which their
spores are produced.
In the Basidiomycetes, the large group which comprises the familiar
toad-stools, bracket-fungi, coral-fungi, puff-balls, stink-horns and their
allies, the spores are borne, usually in groups of four, on the outside of
a cell called a basidium, hence they are basidiospores. In most species
these are shot into the air from the surface of the basidium and are
dispersed by air currents. As a rule they have a slightly asymmetrical
outline and the point at which they were attached to the basidium,
the apiculus, remains visible, so t h a t basidiospores are fairly easy to
recognise as such even when detached. The basidia are normally
packed close together in a continuous layer, the hymenium, covering
the gills on the under side of the toadstool's cap or the pores beneath
the bracket fungus. There are many popular books about the common
Basidiomycetes and they too fall outside the scope of this book.
The second great group of fungi with septate hyphae is called the
Ascomycetes because its distinctive spores are formed inside a charac-
teristic cell called an ascus, and are hence called ascospores. Except in
the simplest forms, the asci are produced in large numbers, orderly
arranged like basidia in a hymenium or in smaller clusters enclosed
in a common wall or sheath. A very large number of Ascomycetes are

XIII
known only in close association with microscopic algae, building up
often large and characteristically shaped vegetative plant bodies
commonly called lichens, much more complex than the inconspicuous
thread-like mycelium of the non-lichenised ascomycetes. Technically
the lichen fungi have to be inserted in their appropriate places among
the orders and families of the Ascomycetes but there are such enor-
mous numbers both of lichenised and non-lichenised species t h a t a con-
vention has grown up by which they are usually the province of
different specialists and are dealt with in different books. This arti-
ficial separation of often closely allied organisms has had most un-
fortunate results on both the classification and the nomenclature of
the Ascomycetes but it will be adhered to here for the traditional
reason, viz. the enormous number of species and genera involved in
a rational amalgamation of the lichenised and non-lichenised fungi.
No fungus is known to produce both basidia and asci and it is
believed t h a t these are mutually exclusive alternative products of a
nuclear fusion corresponding in function to the sexual reproduction of
other organisms. Both Basidiomycetes and Ascomycetes, however, and
especially the latter, may produce alternative spore forms, borne often
on specialised hyphal branches or in specialised receptacles but neither
on basidia nor in asci. Spores of this kind are collectively called conidia
and are regarded as vegetative reproductive bodies and usually treated
as of secondary importance from the standpoint of classification.They
are, however, often more abundant than the ascospores in many
Ascomycetes and there is still a vast array of fungi in which only
conidia are known. They form the third and last category of fungi with
septate mycelium and are not discussed further in this book. The
conidial fructification of the Ascomycetes described in the text is,
however, occasionally referred to if it is particularly characteristic or
is invariably found in close association with the ascus fructification.

T H E S T R U C T U R E A N D C L A S S I F I C A T I O N OF T H E ASCOMYCETES

The main subdivisions of the Ascomycetes are based on the struc-


ture of the ascus itself and of the fructifications within or upon which
they are assembled. The primary division depends on the nature of
the ascus wall, which may be composed either of one or of two layers,
i.e. may be unitunicate or bitunicate.
A bitunicate ascus normally opens by a transverse split in the outer
wall, a short distance below the tip, which is then pushed off like a
thimble by the rapidly expanding inner wall. The ascospores then

XIV
escape through a pore at the tip of the inner wall. This process can
only be observed when fresh ripe material is mounted in water but
fortunately there are usually associated features which help one to
recognise an ascus as probably bitunicate. Usually it is very short-
stalked with the stalk rather sharply delimited from the body of the
ascus containing the spores. It is also usually distinctly thick-walled
towards the tip and very commonly a well defined dimple in the
inner wall can be seen at the tip of the ascus, i.e. on the inner surface
of the thickened wall. The tips of bitunicate asci do not stain blue
with iodine. There is also a tendency for the spores in bitunicate asci
to be multicellular and coloured but this not invariably true, nor are
all asci with coloured septate spores bitunicate. Septate coloured
ascospores in unitunicate asci occur e.g. in Amphisphaeria, Apiorhyn-
chostoma, Ceriophora, Clypeosphaeria, Diapleella, Massariella, Melan-
coniella and Valsaria. Bitunicate asci are usually found within minute
closed fructifications but again there are a few exceptions, e.g. Patellaria.
Unitunicate asci are of several kinds. In the large order Pezizales
they open by a lid or operculum at the tip, which is pushed open when
the ascospores are shot into the air (Fig. 1A). In the remaining orders
there is no operculum and the ascospores escape through an apical
pore or, less often, are not discharged at all, as in the Tuberales
(Fig. ID). In the Helotiales, with a fully exposed hymenium, there is a
simple apical pore, usually penetrating a somewhat thickened and
more or less conical portion of the ascus wall (Fig. lc). Until the asco-
spores are ready for expulsion the pore is blocked, usually with some
substance different in its staining properties from the rest of the wall.
In the majority of species this pore plug stains blue with iodine, either
as a whole or round its edges. In the Sphaeriales there is often a ring-
like thickening on the inner surface of the ascus wall at the tip. This
is visible as two minute highly refractive dots in optical section and
sometimes also stains blue with iodine.
Most unitunicate asci are accompanied by sterile hyphae, called
paraphyses, which occur between them and apparently afford some
protection to the developing asci, at least in species with an exposed
hymenium. The precise shape of the paraphyses is often a useful
character in differentiating species or even genera of Pezizales and
Helotiales; examples include the curved-tipped paraphyses of Otidia
and Pulvinula, the usually lanceolate paraphyses of Dasyscyphus and
the knob-headed paraphyses of Orbilia. In species with a brightly
coloured hymenium the colour is usually contained in the somewhat
swollen tips of the paraphyses. In some genera the paraphyses cohere
at their tips to form a compact layer above the asci, often of a distinc-

VI
tive colour and then called an epithecium, as in the Lecanorales. In
such apothecia the asci and paraphyses are usually enveloped in a
gelatinous substance which often makes it difficult to separate them
satisfactorily in mounts for microscopic examination. In the Sphaeria-
les, with enclosed hymenium, paraphyses are also usually present but
they are of less distinctive appearance and little used in classification,
except as a feature distinguishing the Sphaeriales from the Loculo-
ascomycetes.
In the Taphrinales the fructification consists merely of a layer of
asci, of indefinite extent, spread over the surface of some portion of
the host plant and not accompanied by paraphyses. This is a very
distinctive order, having little in common with the other Ascomycetes
and probably akin rather to the yeast fungi. In the Protomycetales
the asci arise from scattered cells embedded in plant tissue.
In all the remaining orders dealt with here there is a frucfication of
characteristic shape and size which either carries or encloses the asci.
Any such fruit body is conveniently known as an ascocarp and its
shape is of major importance in classification.
An ascocarp bearing a layer of asci in a hymenium fully exposed to
the air at maturity is called an apothecium (Fig. 2A). This is usually
more or less cup-shaped or saucer-shaped—hence the name "Cup
fungi" often applied to the Pezizales and larger Helotiales in which it
occurs—and may be carried upon a longer or shorter stalk or be
completely sessile. In both the orders mentioned the outside of the
apothecium is often ornamented with characteristic hairs, especially
around the margin, and the shape and colour of these must be care-
fully noted before an attempt is made to name any species in which
they occur. In both orders also there are families in which the hyme-
nium is not concave but covers a conical or rounded apex to a stalk,
i.e. in the Morchellaceae and Helvellaceae (Pezizales) and the Geoglossa-
ceae {Helotiales). If this conical apex has an extensive free margin or
is extensively wrinkled or thrown into honeycomb-like pits it is some-
times referred to as a pileus, by analogy with the cap of a toadstool
which is also so called. It should be noted, however, t h a t whereas in
a toadstool the hymenium is always on the under surface of the cap,
in the pileate Ascomycetes it is on the upper and more exposed surface.
As indicated above, the two main groups of apothecial fungi, the
Pezizales and Helotiales, are distinguished by their asci being oper-
culate and inoperculate respectively. This distinction is usually easily
made in fresh material but in immature or dried apothecia it is often
difficult to see an operculum and it is useful to remember t h a t these
primary characters are usually associated with others, less decisive but

XVI
often useful in difficult cases. The first step should be to treat the asci
with iodine; if a blue pore plug is then revealed one need look no
further, the asci are obviously inoperculate. Operculate asci usually
contain symmetrical ascopores and where, as in some Sarcoscypha-
ceae, the spores are asymmetrical this is about a line approximately
parallel with the long axis of the ascus. Ascospores of operculate fungi
are not septate but they very often have roughened or warted walls.
Nearly all soft-fleshed cup fungi more t h a n 1 cm. across belong to the
Pezizales—exceptions include Coryne, Discinella, Leotia, Neobulgaria,
Bulgaria—but so do many smaller species. Most, but not all, of the
Pezizales grow on the ground or on dung.
The Helotiales, with inoperculate asci, commonly have ascospores
t h a t are slightly asymmetrical about an axis approximately at right
angles to the long axis of the ascus, i.e. the upper half of the ascospore
is slightly broader than the lower half or the latter is distinctly more
tapering in outline. Alternatively the ascospores may be long and
slender or be septate. Ascospores of inoperculate asci do not have
warted or otherwise ornamented walls. The apothecia are usually
small to very small and occur as a rule on wood, bark, stems, leaves
or other plant debris. A few genera have smooth spherical ascospores
and with them it may indeed be difficult to see if the asci are oper-
culate or inoperculate unless fresh ripe apothecia are available.
In the small order Ostropales there are a few apothecial genera
distinguished by their exceptionally long, narrow and parallel-sided
asci (Fig. 1E), often with a cap-like thickening at the tip, enclosing long
thread-like ascospores. Most genera of the Lecanorales can be rec-
ognised by their gelatinous hymenia, which stain blue as a whole
with iodine; the asci are usually thick-walled at the tip. Most of the
genera are lichenised but a few are not or include some non-lichenised
species. In the Tuberales various stages can be traced between a much
incurved apothecium or one in which the hymenium has been thrown
into closely spaced ridges and folds and fructifications in which the
asci are completely enclosed. These fungi have a more or less sub-
terranean habit where the ascospores could not be ejected for dis-
persal by air currents ajid the asci are, in fact, found to be indehiscent.
The ascospores, however, show features in common with those of the
Pezizales and the two orders are commonly regarded as being closely
related.
There is also a vast number of Ascomycetes with ascocarps closed
or nearly so but from which the ascospores are nevertheless ejected
at maturity. Such a closed fruit body containing unitunicate asci is
called a perithecium (Fig. 2B). It is usually small, seldom much over

XVII
1 mm. across, spherical or somewhat flask-shaped, with a pore or
ostiole at the tip through which the ascospores are shed. The asci
develop in a hymenium at the base of the perithecial cavity and are
usually interspersed with slender paraphyses. Expulsion of the asco-
spores may be brought about by elongation of the asci in turn so t h a t
their tips reach the ostiole one by one and shoot out their contents.
In other genera the asci become loose in the perithecial cavity and
ooze in succession up to the ostiole. This occurs especially in peri-
thecia which have the ostiole located at the tip of a long neck or beak.
In a few genera the asci break down within the perithecium and a
slimy mass of free ascospores is produced within the perithecial cavity.
The enveloping slime swells and oozes in droplets out of the ostiole so
t h a t the ascospores can no longer be dispersed by air currents but are
presumably transported by insects which rub against the protruding
sticky drops of spores or are splashed about by raindrops.
A perithccium without an ostiole is called a cleistocarp. This type of
fructification is common in the Plectascales, where the asci are usually
few, not arranged in a hymenium and not accompanied by paraphyses.
Bitunicate asci occur in a hymenium in a few families, either in an
apothecium as in Patellaria or in a massive elongated ascocarp opening
by a longitudinal slit (Hysterothecium) as in the Hysteriales. Usually,
however, they are found within closed ascocarps superficially resem-
bling perithecia but without true paraphyses. Instead the hyphae
between the asci are commonly attached to the ascocarp wall at top
and bottom and are often branched or form a network among the asci.
These interascal threads are called pseudoparaphyses or paraphysoids
and the ascocarp is a pseudothecium (Fig. 2c). In practice it is often
very difficult to decide whether the threads among the asci in a ripe
closed ascocarp are attached at top and bottom or at the bottom only
as would be the case with true paraphyses. Hence the interascal fila-
ments are not as a rule mentioned in the descriptions of Sphaeriales
and Loculoascomycetes printed below. It will be understood, however,
that the former normally have paraphyses and the latter pseudo-
paraphyses.
Both perithecia and pseudothecia are individually small and re-
latively inconspicuous but they are often found in more or less massive
compound fructifications. A mass of fungus tissue in which either
type of closed ascocarp is embedded or upon which it is seated is called
a stroma. Stromata vary in size and shape from the massive club-
shaped bodies of Xylosphaera polymorpha, carrying hundreds of em-
bedded perithecia,to minute cushions of hyphae embedded in bark
and easily overlooked unless the bark is carefully sliced with a razor

XVIII
blade. Wherever perithecia or pseudothecia are found to occur in
clusters the presence of a stroma should be suspected and evidence
of it sought for; the presence or absence of a stroma is often an
essential character of a genus. Often perithecia embedded in bark in a
small cushion-like stroma have long slender necks which emerge
together through a slit in the bark and have their ostioles surrounded
by a more or less circular pad of fungus tissue, the disc of the stroma.
Such a habit is described as valsoid, from the old genus Valsa in which
it is well developed. In other genera the presence of a stroma may be
inferred by the presence of a black line in the substrate, usually wood
or the woody portion of a herbaceous stem, surrounding a group of
perithecia, e.g. Diaporthaceae, or even a single apothecium as in some
species of Rutstroemia. A thin shield like patch of stromatic tissue in
host bark or epidermis surrounding an ostiole is called a clypeus. A
web or loose mat of hyphae around the base of an apothecium or peri-
thecium is called a subiculurn and where it is present the ascocarps
usually occur in swarms upon a common subiculum. Presence of a
subiculum is also often utilised as a generic character but it may
easily be overlooked, especially if made of brown hyphae growing
over a dark brown or grey substrate. Where a large number of super-
ficial ascocarps are found scattered evenly over patches of some dead
plant organ a subiculum should be suspected though one is not always
to be found in such cases.
In the Myriangiales almost spherical asci occur scattered singly
through an undifferentiated mass of fungus tissue. In the Pseudo-
sphaeriales club-shaped asci with distinctly bitunicate walls are some-
what similarly disposed in ascocarps superficially like small perithecia.
Finally, in the peculiar Hemisphaeriales the asci are enclosed in
flattened ascocarps, called thyrioihecia, usually very minute and only
just visible to the naked eye as dark specks on the surface of some
plant organ (Fig. 2D).
Most Ascomycetes have 8-spored asci, though one can often find in
them a few asci in which some of the ascospores are imperfect or have
not been differentiated at all. Consistent production of some larger or
smaller number of spores per ascus is usually regarded as a good spe-
cific or even generic character. Several species have uniformly 4-spored
asci; a few have 8-spored asci in which the spores are of two different
sizes or even different colours, as in Bulgaria inquinans. Occasionally
eight ascospores are differentiated but four degenerate, as in Octospora
carneola. The asci of Schizothyrioma ptarmicae and Ptychoverpa
bohemica are consistently 2-spored. One-spored asci occur in some
genera of lichenised ascomycetes not described here, such as Bombylio-

XIX
spora, Lopadium, Mycoblastus, Pertusaria, Umbilicaria and Varicel-
laria. A similar effect of a single massive projectile shot from the ascus
is obtained by a different method in the non-lichenised genus Sacco-
bolus, where eight ascospores are differentiated b u t remain firmly
stuck together.
Asci with more t h a n eight ascospores are developed in two ways.
More t h a n eight nuclei m a y be present as a result of successive nuclear
divisions before any ascospores are delimited. In t h a t case the final
number of ascospores in each ascus is likely to approximate to some
multiple of eight. Thus 16-spored asci are characteristic of Rhyparobius
sexdecemsporus, 32-spored asci of Thecotheus pelletieri, the asci of
Ascozonus woolhopensis contain about 64 ascospores and much higher
numbers occur in some other species of Rhyparobius. The phenomenon
is met with again in the Sphaeriales, e.g. in Podospora, and in the
Loculoascomycetes, as in Rehmiellopsis.
Alternatively only eight ascospores m a y be differentiated b u t t h e y
proceed to bud off secondary spores or conidia while still within the
ascus. In this case the ascus m a y ultimately be packed with innumer-
able minute subglobose or rod-like spores little bigger t h a n bacteria.
This method again crops up repeatedly in quite unrelated genera, for
example Taphrina (Taphrinales), Rutstroemia, Corynella, Tympanis
(Helotiales), Scoleconectria, Rhamphoria (Sphaeriales).
The arrangement of the spores within the ascus is also a fairly
reliable specific character, whether in one row (uniseriate), two rows
(biseriate) or in a parallel fascicle if the ascospores are very elongated
and narrow.

CLASSIFICATION OF T H E L A R G E R ASCOMYCETES

The modern system of classification based on the criteria described


above m a y be summarised as follows:
I. Asci unitunicate or else arranged in a hymenium fully exposed on an
apothecium EUASCOMYCETES (p. 1)
A. Asci neither in a hymenium nor a fascicle but produced singly from
thick-walled chlamydospores produced within the tissue of vas-
cular plants Protomycetales
B. Hymenium of indefinite extent, a mere film of asci covering the
substrate:
1. Asci operculate, covering patches of more or less hypertro-
phied tissue in some living organ of a vascular plant, para-
physes absent Taphrinales (p. 76)
2. Asci inoperculate, in a thin film on bark
Helotiales: Ascocorticium (p. 84)

XX
G. Hymenium of limited extent, borne in or on some receptacle formed
of fungus tissue, usually including paraphyses, or asci not in a
hymenium:
1. Asci operculate Pezizales (p. 1)
2. Asci inoperculate:
a. Asci indehiscent or at least without a definite apical de-
hiscence mechanism:
+Asci in a hymenium, ascocarps ± subterranean
Tuberales (p. 71)
+ +Asci not in a hymenium Plectascales (p. 334)
b. Asci with an apical pore:
+Asci very long, narrowly cylindrical, with thread-
like ascospores:
fAsci in isolated apothecia Ostropales (p. 221)
tfAsci in perithecia, aggregated in a stroma or
on a common subiculum, perithecia not
long-beaked, usually light or bright co-
loured Clavicipitales (p. 226)
+ +Asci not strictly cylindrical, not capitate, asco-
spores sometimes needle-shaped but not thread-
like:
fAsci in apothecia:
§Hymenium blued by iodine or asci clearly
bitunicate . . . Lecanorales (p. 213)
§§Hymenium not blued by iodine, asci uni-
tunicate:
*Apothecia ± immersed in host tissue,
usually with a dark covering layer
opening by a slit or by lobes
Phacidiales (p. 198)
•Apothecia usually superficial or be-
coming so, without a dark covering
layer Helotiales (p. 83)
f t Asci in perithecia:
§ Paraphyses absent, perithecial cavity
formed by breakdown of the tissue
within an originally solid body
Coronophorales (p. 331)
§§Paraphyses usually present at first but
soon vanishing in genera where the
asci become loose and fill the cavity at
maturity . . . Sphaeriales (p. 232)
II. Asci bitunicate, not in apothecia LOCULOASCOMYCETES (p. 353)
A. Asci subglobose, scattered singly through a stroma
Myriangiales (p. 353)
B. Asci in hysterothecia or in ascocarps without a proper margin
Hysteriales (p. 422)
C. Asci in thyriothecia Hemisphaeriales (p. 428)
D. Asci in pseudothecia:
1. Pseudothecia containing a few large asci separated by belts of
hyphae Pseudosphaeriaceae (p. 356)

xxt
2. Not as above:
a. Pseudothecia without pseudoparaphyses
Dothideales (p. 356)
b. Pseudothecia containing pseudoparaphyses
Pleosporales (p. 369)

The position of some genera within the system is still doubtful,


usually because adequate material has not been available for study
by modern methods or because developmental studies of the asco-
carps have not been made. Hence the reader should not expect to be
able to assign every collection which comes into his hands to its
appropriate genus merely by running quickly through the keys.
Moreover the keys have in some cases been simplified so t h a t , t h o u g h
they m a y apply to the British species or to those described t h e y will
not invariably be found applicable to all known species assigned to
the genera. Some experience of a group of organisms is also usually
required before any such keys can be operated profitably. Hence if
one does not quickly arrive at a description which fits the material in
hand one should not at once assume it to belong to a genus omitted
from the book. The keys m a y be faulty, the material may be in some
way atypical, it is even possible you m a y have misunderstood the key
or have overlooked some essential feature of the fungus. If you cannot
m a t c h your fungus in the order you at first r u n down to, t r y the descrip-
tions and figures in other superficially similar orders. You m a y be able
to demonstrate, for example, t h a t a species supposed to have unituni-
cate asci has in fact bitunicate asci and t h u s contribute to a clearer
understanding of its systematic position. Remember especially t h a t
ascospore septation and colour, t h o u g h still much used in delimiting
genera, are r a t h e r unreliable characters and t h a t both m a y develop
very slowly; in some genera, such as Phaeangellina and Echidnodes,
the ascospores often only develop their full colour after expulsion from
the asci.

TAXONOMY AND NOMENCLATURE

Taxonomy is the art of classifying organisms: not a science b u t an


art, for its t r i u m p h s result not from experiment b u t from disciplined
imagination guided by intuition. This is an intellectual activity in
which the English have seldom distinguished themselves, we t a k e our
mycological systems ready-made from the Continent. T h a t adopted in
this book was proposed in its main outline by Nannfeldt in 1932 and

XXII
subsequently modified as regards the Pezizales by Madame Le Gal,
and as regards the Sphaeriales and Loculoascomycetes by Luttrell,
Munk, von Arx and Miiller.
The basic unit of taxonomy is the species, a concept which obsti-
nately defies precise definition. Few persons are convinced to day t h a t
existing species represent static units whose pattern has remained un-
changed since they were made by the Almighty during one eventful
week in 4004 B.C. There is adequate evidence both for the mutability
of species and for the vastly greater age of the inhabited earth but
those who reject the ancient and convenient concept of fixed species
find great difficulty in supplying an acceptable concept to take its
place. Most of those proposed involve some dynamic factor, often with
a genetic basis, but such criteria are of little value to the hardpressed
mycologist, faced with the task of classifying an enormous number of
fungus collections and usually without any experimental evidence to
guide him. For him, indeed, it is fair to say t h a t a species consists of a
number of collections which appear to him to agree tolerably in
visible characters with one specified collection on which a specific
name has been based, i.e. a type collection, and to be visibly different
from the members of all other such groups of collections known to him
at t h a t time. Thus examination of the collections at Kew shows t h a t
for George Massee the fungi described in this book as Octospora humosa,
Neottiella rutilans and N.vivida represented a single species; for
Berkeley and Cooke 0.humosa was easily separated from the others
by its more cylindrical smooth ascospores; for Boudier not only was
this so but N.rutilans and N.vivida were clearly distinct in their spore
ornament. The writer agrees with Boudier, who was a much greater
taxonomic artist t h a n Massee, but there is no experimental evidence
to show t h a t he is right and t h a t these three forms are not in fact
interbreeding states of a single species. Every taxonomist hopes, of
course, t h a t his taxonomic species correspond to real discrete living
entities in nature but he is seldom in a position to state categorically
t h a t this is so. There thus remains a large subjective element in taxo-
nomy and attempts to eliminate it have seldom been particularly
helpful. From this angle the position has been well summed up by an
eminent mycologist t h u s : "Now it is perfectly possible to believe t h a t
a natural species of plants exists in the field, or if you prefer it in
nature, quite independently of the human race; t h a t it was there
before the hills in order stood, and may be there long after we are gone.
But a taxonomic species cannot exist independently of the human
race; for its constituent individuals can neither taxonomise them-
selves into a species, nor be taxonomised into a species by science in

XXIII
the abstract; they can only be grouped into species by individual
taxonomisers".
Nevertheless there is a comforting degree of agreement among
practising taxonomists as to which units are recognisably distinct,
though they often disagree as to the relative status of particular units,
whether species, subspecies, varieties or forms and as to what consti-
tutes a valid generic criterion. An optimist m a y regard this wide field
of agreement among taxonomists as an indication t h a t taxonomic
species do correspond to real n a t u r a l units; a pessimist m a y dismiss
it as merely a reflection of the influence of tradition on h u m a n activity
or as an indication of the degree to which every taxonomist copies the
writings of his predecessors.
As regards units of classification higher t h a n species there are also
two schools of t h o u g h t . One contends t h a t , whatever may be true of
species, genera and families are purely h u m a n inventions. The other
believes it possible to recognise natural genera, each containing only
species descended from a common ancestor. For this school the genus
of t o d a y is derived from the species of yesterday and will in t u r n give
lise to the family of tomorrow. In their view "Le b u t de la systemati-
que n'est pas de dresser un simple catalogue des etres vivants, mais de
les ranger dans un ordre qui retrace la phylogenie. L'espece constitue
une unite trop petite pour etre utilisable aisement; l'unite pratique
est le genre".* In practice, whatever one's views, one finds it most
satisfactory to group species according to their least variable charac-
ters or by those one considers of most f u n d a m e n t a l importance and so
most likely to be stable. Often these are relatively insignificant or
apparently trivial features, such as the operculum of an ascus or the
blue stain of the ascus tip with iodine characteristic of the Pezizeae.
It is in recognition of the significance of such characters t h a t the
taxonomist's " i n t u i t i o n " comes into play. It m a y well be t h a t these
characters are reliable because t h e y have been inherited from a com-
mon ancestor and in practice the most dependable classification often
proves to be one which is acceptable from the standpoint of theoretical
phylogeny. In fact one of the early arguments in favour of a theory of
evolution of living organisms was based on the prior existence of a
" n a t u r a l " system of classification with its implied interrelationships
of t h e species classified. Nevertheless it must be realised t h a t the pur-
pose of a classification is to enable one to arrange a vast n u m b e r of
units in an intelligible order, so t h a t the same name shall always be
applied to the same unit and to make it possible to discover whether

* Doguet in Le Botaniste, 39, 1955.

XXIV
a previously unfamiliar organism has already received a name from
some other worker. One is not primarily concerned with its pedigree,
nor, in the absence of an adequate fossil record of fungi, is there any
convincing means of verifying the often very diverse speculative lines
of descent and interrelationship which have been proposed, usually on
very flimsy grounds. The real difference between an "artificial" and a
"natural" classification may be better expressed by saying that in the
former one fits collections to the system and in the latter the system
is made to fit the collections.
The names used for fungi, like those for all other organisms, are
constructed in accordance with the binomial system of nomenclature
introduced over 200years ago b y t h e Swedish naturalist Carl von Linne.
Whatever language the names may be derived from they are converted
to a Latin form and the name of each species consists of two parts,
the generic name, written first, and the specific epithet. Thus, in
Morchetta esculenta Persoon the generic name, Morchella, is common
to all the Morels, the specific epithet esculenta, designates the particular
species. The name Persoon indicates the mycologist who first applied
the name to the species in question. In this instance the name has re-
mained unchanged since 1822 but the majority of old established
species have been transferred from the genus to which their founder
first assigned them to some other considered more appropriate by a
later author. In such cases the name of the author who first named
the species is written first, in brackets, and the name of the author
who made the transfer is added after it, for example Trichophaea
abundans (Karsten) Boudier, first described by Karsten as a species of
Peziza and later transferred by Boudier to Trichophaea. Botanical
nomenclature is governed by an elaborate code of rules, agreed to by
International Botanical Congresses, and under this code the nomen-
clature of Ascomycetes begins with the year 1821, with the proviso
that names proposed in 1821 shall not be used if they conflict with
those used by E. M. Fries in the second volume of his 'Systema Myco-
logicum', published in 1822. Hence names proposed by authors in
works published prior to 1821 can only be used if they have been
adopted by some author after 1st January of t h a t year. Thus the name
Helvetia lacunosa Afzelius ex Fries was proposed by Afzelius long prior
to 1821 and accepted by Fries in 1822.
It will be seen t h a t there can be no finality about the name applied
to any collection. The only name which is certainly correct is the
specific epithet as applied to the type collection on which the taxo-
nomic species was based but even t h a t is liable to be displaced if it is
considered that the type collection in question is conspecific with t h a t

XVI
on which an earlier validly published specific epithet was founded.
The appropriate generic name can never be more than a matter of
opinion.
TECHNIQUE

In the field.
The equipment required for collecting ascomycetes is of the simp-
lest, a few tins of assorted sizes for the larger or more fragile specimens,
a bundle of used envelopes for the minute fungi on stems and leaves.
Methods of collecting will depend partly on the collector's special
interests, more on his degree of patience. Only a few large or brightly
coloured or exceptionally gregarious species will be found by walking
briskly through the countryside; the smaller fungi are only to be dis-
covered by crouching or kneeling in a suitable spot, scanning the sur-
rounding herbage closely and turning over fallen leaves, twigs and
even logs one by one. One of our most successful collectors of Helo-
tiales makes a practice of spreading a macintosh in an attractive-
looking ditch, lying upon it and not stirring further for an hour or more
while he methodically scrutinises every piece of vegetable debris
within reach. As far as possible one should endeavour to sort the
material obtained into species by eye, aided by a pocket lens, as it is
collected and make sure one has secured an adequate collection of
every species found. A single apothecium or perithecium of an almost
microscopic fungus is of little value as it will be destroyed during the
subsequent examination when one attempts to name it. Pezizales and
most Helotiales are short-lived and some part of any ample collection
is likely to be in a state of maturity in which it can be recognised
specifically or at least referred to its approximate systematic position.
The Sphaeriales have tough perithecial walls which often persist long
after their contents have been dispersed; moreover they often mature
very slowly and many collections will be found to contain no mature
asci. The larger species should be sliced with a sharp knife in the field;
if the perithecia are found to have sticky contents recognisable asco-
spores are likely to be present, if no contents are seen the collection
will probably prove to be worthless. Care should be taken to note m
the field the habit and substrate of each collection, remembering,
however, t h a t fallen twigs have not necessarily come from the tree
under which they were lying when found.
The collector of ascomycetes has one great advantage over his
colleagues interested in toadstools, one can find ascomycetes of some
kind at all seasons. Some species, for example of Octospora, occur only
in mid-winter and many Sphaeriales are at their best in winter and

xxvi
early spring. The latter is also a good season for Sclerotiniaceae and
m a n y other Helotiales and some Pezizales. The a u t u m n , the tradi-
tional " F u n g u s season", also yields a good harvest of b o t h large and
small species.

At home.
The preliminary examination of any ascomycete should be made
under a hand-lens of magnification about x 10 or a binocular dissecting
microscope at a magnification of about x20. The earlier mycologists
relied entirely on this method and were able to distinguish a sur-
prisingly large n u m b e r of species. It is t r u e t h a t few ascomycetes can
be assigned to their appropriate place in the system without micro-
scopic examination b u t m a n y essential characters are still best seen at
low magnifications. The plates in this book figure nearly all the smaller
species as seen x l O or x20, only a few of the most minute are figured
x 100. E. H. A's dictum t h a t " t h e microscope comes from micros small
and scopein to see and no-one who relies on it can grasp a large idea"
was in p a r t a jest b u t the jester's folly is spiced with wisdom. Many
characters easily seen when ascocarps are viewed in situ and whole are
obscured when t h e y are sliced or crushed for examination under a
compound microscope. I t is a useful routine practice to make a
coloured sketch of all small apothecia as seen under a hand-lens while
t h e y are still fresh. This is less necessary with most Sphaeriales and
Loculo-ascomycetes as t h e y seldom change their shape and colour
greatly when dried for t h e herbarium. Attention should also be
directed to the habit of the ascocarps and their position in or on the
substrate, secondly to their gross morphology, shape, size, presence or
absence of hairs—often very easily overlooked in the smaller Pezi-
zales—presence or absence of a subiculum or a stroma. Only t h e n is
one ready to extract a few asci or to section the flesh for examination
under high magnification, preferably xlOOO or over. Details of asci
and ascospores figured in this book were drawn with a camera lucida
at x 1000 and reduced b y one-third in preparation of the plates so t h a t
the final magnification is about x660 unless otherwise stated. Critical
details of ascospore ornament in species of Peziza m a y only be ap-
preciable with oil immersion lenses at magnifications in the region
of x2000.
Asci, paraphyses and hairs m a y be examined microscopically when
mounted in water and this medium will give one the truest colours of
spore walls and sap. It is usually desirable, however, to employ also
some staining and clearing agent in critical studies and with the
ascomycetes the first stain to be used should always be iodine. This is

XXVII
best supplied in the form of Melzer's reagent, which also contains
chloral hydrate to act as a clearing agent. Melzer's own formula was:
Potassium iodide . 1.5 gr.
Iodine 0.5 gr.
Distilled water . . . 20.0 ml.
Chloral hydrate . . 20.0 ml.
Some authors prefer to use a smaller q u a n t i t y of iodine.
Iodine gives a blue stain to the hymenium of most Lecideaceae and
its action in similarly staining the ascus tip is an essential character
of the Pezizeae, at once separating t h e m from the superficially rather
similar species in Pustularia, the Otideae and the Humariaceae. The
blue reaction of the pore plug is a useful feature of m a n y Helotiales,
though this m a y depend on the stage of development of the ascus,
and a blue stain of the apical a p p a r a t u s in the ascus is sometimes a
help in separating Sphaeriales from Loculoascomycetes. Within the
Sphaeriales it distinguishes the Families Xylariacae and Amphi-
sphaeriaceae. Iodine also usually imparts a yellowish-brown colour to
the non-gelatinised hyphal walls in general and may help one to
decide if thin septa are present in ascospores or if there was merely an
appearance of septa due to two large vacuoles in apposition.
For critical studies of ascospore ornament, especially in the Peziza-
les, a different stain is required. This is prepared by dissolving 0.05 gm.
Cotton Blue or Aniline Blue in 30 ml. lactic acid and filtering to re-
move any undissolved residue or precipitate. Best results are said to be
obtained by allowing the Cotton Blue to dissolve slowly at room
t e m p e r a t u r e instead of boiling it in the lactic acid. Thin sections of
fragments of hymenium picked out with a mounted needle are placed
in a drop of the solution on a microscope slide, a coverslip added and
the slide then gently heated until the drop begins to boil. W a r t s , spines
or reticulated thickenings on the outer surface of ascospores stain
deeply while the ascospore wall itself remains colourless. Cotton Blue
m a y also occasionally be useful in detecting ascospore septa as it stains
protoplasm and leaves the septa as hyaline lines between the proto-
plasts b u t it should not be used as a routine stain for ascomycetes as
it is far inferior to Melzer's reagent for t h a t purpose.
It is often necessary to re-examine asci or tissue from specimens
which have been dried and kept in a herbarium. It is t h e n often
desirable to soak the material for a short time in 10% ammonia or 5 %
KOH solution to soften it and swell the tissues to their former dimen-
sions before staining. A m o u n t m a y be transferred directly from am-
monia to Melzer's reagent b u t it should be rinsed with water before
cotton blue is applied.

XXVIII
F I E L D S OF W O R K E S P E C I A L L Y S U I T A B L E FOR AMATEURS

One great attraction of the ascomycetes for the enthusiastic ama-


teur naturalist should be t h a t the British flora is so little known that he
has an excellent chance of making some worth-while contribution to
knowledge. Most amateurs start by collecting indiscriminately any-
thing t h a t attracts their attention and eventually, if their collections
are numerous enough, aspire to publish one or more lists of the fungi
found in a particular county or parish. The broad experience thus ob tam-
ed maybe very helpful to the collectorandlocallistsofthiskindarenotto
be despised, provided care is taken to see the names used are reason-
ably accurately applied and to preserve the collections on which the
records are based so t h a t subsequent workers can check them if
necessary.Lists of records that cannot be verified are mere waste paper.
Rather more precise and valuable information is likely to be sup-
plied by an amateur who first obtains a general acquaintance with
the fungus flora of his district and then proceeds to specialise in some
way. He may, for example, endeavour to make a detailed study of the
fungi found in some distinctive habitat or associated with a particular
host or substrate. Even city dwellers may be able to contribute
something on these lines. It is a fortunate borough whose pavements
are not daily fouled by dogs yet little is known of the fungi which
colonise this substrate. Nearly half a century ago Grossland observed
interesting species of Rhyparobius and other genera on "canine ex-
crement" but hardly anyone has looked at it since his day. A rich
harvest may well await the man who cares to devote his leisure hours
or his declining years to the study of stale dog dung. In general, how-
ever, dung fungi are not restricted to the product of any particular
animal and 110 such inference is to be drawn from such familiar epithets
as "bovina", "canina", uequina", "felina", llhumana", uporcina" etc.
Thus Fimaria humana Velenovsky is said to be a synonym of F. theio-
leuca (p. 52) which leads a less opulent existence on rabbit droppings
in Britain, also in short supply since the introduction of myxomatosis.
Because of the inconspicuous and often ephemeral character of
many ascocarps as well as their sporadic and unpredictable occurrence
in nature, it is more difficult for an amateur to specialise in the study
of any particular genus with a view to preparing a definitive mono-
graph. There are, however, very many genera which urgently require
intensive study and some, like Leptosphaeria, include so many wide-
spread species t h a t an amateur can reasonably expect to obtain a
considerable number by his own efforts. After he has become expert in
manipulating small amounts of valuable material and has learned to

XXIX
recognise the reliable specific characters in the genus he is likely to
receive a sympathetic hearing if he applies to one of the larger herbaria
for access to additional species he has not encountered in the field.
One great obstacle to all such monographic studies, indeed, lies in the
lack of adequate collections of most species of ascomycetes in the
national herbaria and the dispersal of what few collections have been
left by past generations of mycologists.
There are also a few genera with clearly defined habitats which an
amateur may reasonably hope to be able to monograph completely
by his own efforts. One such which comes readily to mind is Ascozonus.
In Journal of Botany 12:353-357 (1874), Renny described six species
of this genus which according to him are "found on the dung of rab-
bits and hares, birds and mice. At most seasons of the year they may
be met with but chiefly in winter". Some of Kenny's species have not
been met with again and though he figured them well, he did not in-
dicate the dimensions of the ascospores. Here then is a compact genus,
easily recognised by its peculiar method of ascus dehiscence (Fig. In),
said to be common, which would well repay critical study.

ECONOMIC IMPORTANCE

Only a few ascomycetes are directly consumed as food by European


peoples, mainly species of Tuber and Morchella, and even these are
seldom eaten in England. Species of the curious genus Cyttaria, para-
sitic on Nothofagus trees, are said, however, to have formed part of
the diet of the natives of Tierra del Fuego until recent years. Baker's and
brewer's yeasts (strains of Saccharomyces cerevisiae Hansen) are asco-
mycetes belonging to the family Endomycetaceae, not described in
this book.
A few ascomycetes cause diseases of economic plants and others
which normally grow on fading plant parts may become injurious to
crops grown under unsatisfactory cultural conditions. The most de-
structive ascomycete known is no doubt Endothia parasitica (p. 322),
not yet introduced into the British Isles, cause of the disastrous canker
disease of sweet chestnut trees in North America. A less dramatically
effective pathogen of British trees is Ceratocystis ulmi (p. 337), cause
of the Dutch Elm Disease which occasionally attracts public attention
when its presence requires the felling of some well-known avenue. More
important are the parasites of the apple tree, Nectria galligena (p. 244),
cause of canker and of a minor fruit rot, Venturia inaequalis (p. 375),
cause of scab on leaves and fruit, Monilinia laxa, cause of Blossom
Wilt and Spur canker and Monilinia fructigena, the common brown

xxvi
rot fungus. All the species of Sclerotinia are plant parasites and some,
like S.trifoliorum (p. 93), cause of clover rot and S.sclerotiorum (p. 92),
capable of rotting almost any fleshy plant tissue, especially in storage,
occasionally cause serious losses. The Erysiphaceae (p. 345) are also
obligate plant parasites b u t only Apple Mildew, Podosphaera leuco-
tricha, is usually a serious problem to British horticulturalists. Among
the feebly parasitic fungi which sometimes become destructive to
crops, especially under conditions of low t e m p e r a t u r e or high humidity,
the best known is Sclerotinia juckeliana, with its ubiquitous conidial
state formerly known as Botrytis cinerea.
For a useful summary of the prevalence of plant pathogenic ascomycetes
and other fungi in the British Isles the reader should consult Ministry of
Agriculture Bulletins 126 and 139, Diseases of Crop Plants 1933-42 and
1943-46 respectively, both compiled by Mr. W. C. Moore of the Plant patho-
logical Laboratory, Harpenden.

ANNOTATED LIST OF U S E F U L BOOKS

Critical papers on individual genera are cited in the appropriate


places in the text. The following books and papers are of more general
application.
VON A R X , J. A . & M t i L L E R , E. (1954) Die Gattungen der Amerosporen Pyreno-
myceten. Beitrage zur Kryptogamen-Flora der Schweiz, 11 (1).
A critical illustrated account of all known genera of Sphaeriales and
Loculoascomycetes with aseptate ascospores.
B E R L E S E , A . N . (1890-1905) Icones Fungorum ad usum sylloges Saccardianae
ac'comodatae. Excellent figures of Sphaeriales and Loculoascomycetes,
mostly taken from authentic specimens.
B E R T H I E R , P. (1964) Essai Biotaxonomique sur les Discomycetes. Univ. Lyon,
These 340.
B I S B Y , G . ( 1 9 5 3 ) An introduction to the taxonomy and nomenclature of fungi.
Edition 2. Commonwealth Mycological Institute, Kew.
Contains a summary of the International code of botanical nomen-
clature and many useful hints on collecting, recording and preserving
fungi.
B O U D I E R , E . ( 1 9 0 5 - 1 0 ) Icones Mycologicae ou Iconographie des Champignons
de France. Tome 2 (Pezizales), 3 (mainly Helotiales) and 4 (text).
Very expensive and now rarely obtainable but important as con-
taining the finest coloured plates of fungi ever issued, especially of Pezi-
zales.
D E N N I S , R . W . G . ( 1 9 4 9 ) A revision of the British Hyaloscyphaceae with notes
on related European species. Comm. Mycol. Institute Mycol. Paper 32.
— ( 1 9 5 6 ) A Revision of the British Helotiaceae in the Herbarium of the Royal
Botanic Gardens, Kew, with notes on related European species. Comm.
Mycol. Institute Mycol. Paper 62.
These two papers summarise current knowledge of about three-
quarters of the recorded British Helotiales.

XXXI
INGOLD, C. T. (1953) Dispersal in Fungi. Oxford, Clarendon Press.
Describes and figures the diverse methods of ascus dehiscence.
K I R S C I I S T E I N , W. (1911-38) Sphaeriales in Kryptogamen Flora der Mark
Brandenburg, 7 Pts. 2 and 3.
Unfinished but includes also some Loculoascomycetes.
LE GAL, M. (1947) Recherehes sur les Ornamentations sporales des Discomy-
cetes opercules. Thesis, Series A450, No. 474, Paris.
Contains highly technical descriptions of the diverse types of asco-
spore ornament found in the Pezizales and discussion of their taxo-
nomic significance.
— ( 1 9 5 4 ) Les Discomycetes de Madagascar. Prodrome a une Flore mycologique
de Madagascar. Paris. Vol. 4.
A brilliant account of Pezizales and Helotiales received from Mada-
gascar, including many cosmopolitan species; essential to the serious
student of the Pezizales in any part of the globe.
L U T T R E L L , G . S. ( 1 9 5 1 ) Taxonomy of the Pyrenomycetes. University of Mis-
souri Studies, 24 (3).
Draws attention to the significance of the bitunicate ascus.
— (1955) The Ascostromatic Ascomycetes, in Mycologia 47:511-532.
A brief survey of the Loculoascomycetes.
M U L L E R , E . & VON A R X , J . A . ( 1 9 6 2 ) Die Gattungen der didymosporen Pyreno-
myceten. Beitrage zur Kryptogamenflora der Schweiz 11 (2), 922
pages.
Continuation of von A r x & Muller (1954), essential to the student of
Sphaeriales and Loculoascomycetes.
M U N K , A. (1953) The system of the Pyrenomycetes, in Dansk Botanisk Arkiv
15, No. 2.
— (1957) Danish Pyrenomycetes, Ibid. 17, 491pp.
An account in English, of the Sphaeriales and Loculoascomycetes
known in Denmark.
N A N N F E L D T , J. A. (1932) Studien iiber die Morphologie und Systematik der
nicht-lichenisierten inoperculaten Discomyceten, in Nova Acta Regiae
Societatis Scientiarum Upsaliensis. Series 4, 8, No. 2.
The work in which the modern system of ascomycetes was formulated,
deals in detail only with genera of Ostropales and Helotiales.
P E T C H , T. (1938) British Hypocreales. Cambridge University Press.
A condensed but useful summary of the then known British Clavi-
cipitales, Hypocreaceae, Nectriaceae and Melanospora, etc. For
Nectria replaced by Booth ( 1 9 5 9 ) , s. p. 240.
R E H M , H. ( 1 8 8 7 - 9 5 ) Ascomyceten: Hysteriaceen und Discomyceten. Published
as Volume 1, Abt. 3, of Rabenhorst's Kryptogamen Flora, Edition 2.
Some of the descriptions of Helotiales are still useful.
T R A V E R S O , J . B. ( 1 9 0 6 - 1 3 ) Flora Italica Cryptogama 2 (1) Pyrenomycetae.
Unfinished b u t covers the greater part of the Sphaeriales.
W I N T E R , G . ( 1 8 8 4 - 8 7 ) Ascomyceten: Gymnoaseeen und Pyrenomyceten. Pub-
lished as Volume 1, Abt. 2, of Rabenhorst's Kryptogamen Flora,
Edition 2.
Still useful for individual descriptions.

XXXII
DESCRIPTIONS OF GENERA AND SPECIES

EUASCOMYCETES

Order 1. PEZIZALES

The members of this large order are characterised by having asci


which dehisce b y an apical lid, the operculum. Operculate asci are
usually large with symmetrical elliptical or spherical ascospores. Sep-
t a t e ascospores are unknown b u t the ascospore walls are commonly
ornamented with external warts, spines, crests or ridges, the shape and
distribution of which afford valuable specific characters. The hymeni-
um is either on a club-shaped and honeycombed receptacle (Morchella-
ceae), on a stalked pileate receptacle whose upper surface is often con-
voluted or saddle-shaped (Helvellaceae) or, usually, on t h e more or
less concave upper surface of a sessile cup-shaped apothecium. In most
genera the apothecia are relatively large and light coloured or tinted
with shades of brown, red or yellow. A few black species occur, how-
ever, and in some genera, especially in Ascobolaceae, the apothecia
m a y be minute or even microscopic. Few species are of economic im-
portance but the Morels are commonly eaten on the Continent, as are,
less often, some large species of Peziza and Aleuria. In spite of its name
Gyromitra esculenta is not recommended for eating and Sarcosphaera
eximia is said to cause gastric troubles, nausea, vomiting and diar-
rhoea in some people especially if consumed raw. Rhizina undulata has
been regarded as a parasite of conifers.
The arrangement adopted here is in the main t h a t of Le Gal accord-
ing to whom the order contains six families, distinguished as follows:

I. Receptacle stalked (if cup-shaped with slender stalk and red disc see
Geopyxis, Sarcoscypha):
A. Ascospores elliptical, without internal oil drops but having a num-
ber of external granules adhering to each end Morchellaceae
B. Ascospores elliptical to fusiform, with large internal oil drops
Helvellaceae (p. 5)

II. Receptacles usually sessile, cup-shaped or button-shaped:


A. Asci turn blue, at least at the tip, with iodine:
1. Asci narrowly cylindrical, ascospores uniseriate
Pezizaceae [Pezizeae] (p. 4)

1
2. Asci broad, ascospores commonly biseriate, apothecia minute,
see Pseudoascoboleae (p. 60)
B. Asci not blued at the tip by iodine:
1. Apothecia very minute, containing a single mature ascus
which remains enclosed until dehiscence Thelebolaceae (p. 70)
2. Apothecia containing asci in a hymenium or (in Ascodesmis)
in a fascicle:
a. Asci broad, protruding beyond the general level of the
hymenium at maturity, often on dung
Ascobolaceae (p. 55)
b. Asci narrow, not protruding:
*Apothecia large, one-sided, yellow or brown, asco-
spores smooth Pezizaceae [Otideae] (p. 25)
**Apothecia often large, stalked, texture fibrous,
lignicolous . . . . Sarcoseyphaceae (p. 66)
***Apothecia small, often hairy, texture soft and cel-
lular, mostly on the ground Humariaceae (p. 30)

Family 1. Morchellaceae

This family is characterised b y its smooth ascospores, without pro-


minent internal oil drops b u t bearing a n u m b e r of small external
granules on each end of t h e spore. It includes four British genera of
rather diverse external appearance, readily distinguished as follows:

I. Hymenium lining numerous shallow broad pits, separated by sterile


ridges, hymenophore always stalked . . . . Tribe Heterogeneae
A. Hymenophore club-shaped, only separated from the stalk by a
shallow furrow Morchella (p. 3)
B. Hymenophore hanging free in its lower part and forming a loose
collar round the upper part of the stalk Mitrophora (p. 4)

II. Hymenium smooth or veined but not interrupted by sterile ridges


Tribe Homogeneae
A. Hymenophore conical or cylindrical, detached from but surround-
ing a long stalk, from the apex of which it is dependent:
1. Hymenial surface gyrosely furrowed, asci 2-spored
Ptyclioyerpa
2. Hymenial surface smooth or only vertically furrowed, asci
8-spored Verpa (p. 4)
B. Hymenophore broad, flattened or convex, often veined, almost
sessile Disciotis (p. 5)

Ptychoverpa bohemica (Krombholz) Boudier is a central european


fungus, not known in Britain.

2
MORCHELLA St. A m a n s
(The Morels)

Morchella St. Amans, Flore Agenaise:591 (1821).


Hymenophore conical to almost spherical in broad outline b u t very
variable in shape and size, pitted b y broad shallow depressions lined
with hymenium and separated b y narrow, paler, sterile ridges, delimi-
ted below by a shallow furrow and seated on a tall, whitish, cylindrical,
hollow, somewhat scurfy stalk. Numerous species have been described
in Europe but most modern authors seem agreed to regard these as
variants of two, M.esculenta with light coloured reticulating ridges
and M.elata with dark, more or less parallel, longitudinal ridges.

Morchella esculenta Persoon ex St. Amans, Flore Agenaise: 591 (1821).


Common Morel (Fig. 3A).
Fruit bodies very variable in size and shape, up to 20 cm. high, the
fertile upper p a r t covered b y a honeycomb-like arrangement of nar-
row ridges surrounding angular and often slightly elongated shallow
pits, at first greyish, becoming yellowish-brown; stalk white to cream-
coloured, cylindrical or slightly enlarged at the base, brittle, hollow,
t h e surface even or marked by longitudinal furrows, minutely scurfy.
Asci up to 330x20 (i; spores broadly elliptical, 1 6 - 2 3 x 1 1 - 1 4 (j., almost
colourless under the microscope b u t deep cream coloured in the mass;
paraphyses cylindrical or slightly enlarged upwards, multiseptate,
8 - 1 7 (x broad.
Solitary under trees (Acer, Corylus, Fagus, Fraxinus, Ulmus) in open
woods, orchards, roadside banks or in open grassland, April and May.
Widespread, especially in southern and eastern England, but seldom
abundant.

Morchella elata Fries, Systema mycologicum 2 : 8 (1822). Fig. 3B.)


Fruit bodies smaller and narrower t h a n in the preceding, up to
10 cm. high, the fertile upper portion cylindrical to slightly conical,
with yellowish-brown alveoli arranged in rows between dark grey or
black vertical ribs; stalk white or yellowish, cylindrical, hollow, sur-
face coarsely mealy. Asci up to 3 0 0 x 2 0 JA; spores mostly 18-25 x
11-15 |j.; paraphyses as in M.esculenta, spores cream-coloured in t h e
mass.
In coniferous woods in Scotland and northern England, May. Rare.
A variety purpurascens Boudier in Bull. Soc. mycol. France 13:148 (1897),
differing from the type only in its pale violaceous colour, has been reported
from Dunbartonshire.

3
Morchella hortensis Boudier in Bull. Soc. mycol. France 13:145 (1897).
Fruit body similar to t h a t of M. esculenta b u t with the fertile portion
more elongated, olive-brown, with a predominance of more or less
parallel vertical ribs joined b y cross-septa as in M.elata b u t without
the black rib-edges of the latter. (Fig. 31 A.)
In gardens, May. Rare.
Though M.hortensis is the accepted name it would appear to be
illegitimate as Boudier reduced the older M.vaporaria de Brondeau
to a variety of his new species at the time of publication.

MITROPHORA Leveille

Mitrophora Leveille in Ann. Sci. Nat. Bot. I l l , 5 : 2 4 9 (1846).


Differs from Morchella, with which it is often united, in the lower
half of the fertile head being free from the stalk. There is one well
defined British species:

Mitrophora semilibera (de Candolle ex Fries) Leveille, op. c i t . : 2 5 0


(1846). (Fig. 3c.)
F r u i t bodies up to about 13 cm. high, fertile head small, more or less
conical, with yellowish-brown alveoli arranged in vertical rows be-
tween slightly anastomosing blackish ribs, free from the stalk for the
lower half or third of its length: stalk cylindrical or slightly enlarged
at the base, hollow, often becoming furrowed with age, white or
yellowish with a scurfy surface. Asci up to 4 5 0 x 2 5 jx; spores 2 2 - 3 0 x
14-18 (i, cream-coloured in t h e mass; paraphyses slightly clavate, sep-
t a t e , 12-18 [a wide.
In damp woods, April and May. Not uncommon. The figure is based on a
collection from Woodbridge, Suffolk, 5.V.1949. Very large specimens have
been distinguished as M.gigas (Persoon ex Fries) Leveille, op. cit.: 250.

YERPA Swartz ex Persoon


Ycrpa Swartz ex Persoon, Mycologia Europaea 1 : 2 0 2 (1822).
Fertile head campanulate, smooth or slightly furrowed, pendulous
from the apex of a cylindrical stalk; asci and spores like those of
Morchella, paraphyses more slender.

Yerpa conica Swartz ex Persoon, Mycologia Europaea 1:204 (1822).


(Fig. 3 D . )
Fertile head olive-brown to dark brown, c a m p a n u l a t e ; stalk cylin-
drical, whitish to cream-coloured, up to 1 cm. thick, hollow, surface

4
ornamented with slightly darker granules arranged in irregular hori-
zontal bands. Asci up to 3 5 0 x 2 3 jx; spores elliptical, 20-24x 12-14 jx;
paraphyses slightly clavate, septate, up to 12 jx thick at the tip.
Amongst grass and along roadsides, May. Uncommon.
Specimens with the fertile head more cylindrical and fitting closely round
the apex of the stalk are often distinguished as V. digitaliformis Persoon,
Mycologia europaea 1:202 (1822). The figure is from a drawing by Crossland
of a collection made among roadside grass at Masham, Yorkshire, 22. v. 1908.

DISCIOTIS Boudier

Disciotis Boudier in Bull. Soc. mycol. France 1:100 (1885).


Fruit body an apothecium, very large, becoming fully expanded or
even reflexed and lobed, the hymenial surface some shade of brown,
often veined or convoluted; stalk short and broad or almost obliter-
ated. Distinguished from the Pezizaceae b y its asci not being blued b y
iodine and by its broadly elliptical spores often bearing polar granu-
les; paraphyses broad, septate.

Disciotis venosa (Persoon) Boudier, Icones mycologicae II, Livraison


10 (October 1906). (PI. IIA.)
Apothecia large, up to 15 cm. across when fully expanded, hymeni-
u m dark brown, in large specimens always more or less deformed b y
ribs and furrows radiating from the centre, exterior whitish or cream,
minutely scurfy; stalk short and stout, sometimes sunk in the soil.
Asci about 320x20 [x; spores broadly elliptical, 19-25 x 12-15 fx, with-
out conspicuous internal oil drops; paraphyses septate, with brown
sap, slightly clavate and up 12 jx broad at the tip. The flesh has an
odour of E a u de Javelle, or hypochlorite solution, and is very brittle,
so t h a t large expanded apothecia often split extensively in situ and
are not easily collected intact. L u x u r i a n t forms with strongly con-
voluted hymenia have been called var. reticulata (Greville) Boudier,
Hist. Class. Discom. d ' E u r o p e : 4 2 (1907), b u t are scarcely w o r t h a
distinctive name.
On soil in woods, on shady banks and lawns, March to May. Common.
The figure represents part of a collection from the moist shaded bank of a
sunken road nearWotton-under-Edge, Gloucestershire, 22.iv.1953.

Family 2. Helvellaceae

The Helvellaceae include genera with usually more or less stalked


hymenophores, varying in form from cup-shaped to saddle-shaped,

5
often lobed, veined, contorted or intricately folded. All are terrestrial
b u t Rhizina m a y be in close contact with dead wood or buried roots.
The asci do not stain blue with iodine, the ascospores contain con-
spicuous internal oil drops and bear no polar granules. In Discina and
Rhizina the spores are fusiform, apiculate and minutely roughened,
in all the other genera t h e y arc elliptical and smooth. Eight genera
have been recognised in Britain b u t it must be admitted t h a t the
distinctions between Helvetia, Leptopodia, Cyathipodia and Paxina are
ill-defined and probably u n n a t u r a l . Modern Scandinavian authors
reduce Leptopodia, Cyathipodia and Paxina to synonymy under Hel-
vetia. For a recent monograph of this enlarged genus see Dissing, " T h e
Genus Helvetia in E u r o p e " in Dansk. Bot, Arkiv 25 (1): 1-172, 1966.

I. Ascospores smooth and elliptical:


A. Fruit body convoluted and brain-like, brown Gyromitra
B. Fruit body saddle-shaped or cup-shaped . . . Tribe Helvelleae
1. Fruit body saddle-shaped, on a long stalk:
a. Stalk stout, ribbed, furrowed and chambered
Helvella (p. 7)
b. Stalk slender, surface even leptopodia (p. 9)
2. Fruit body cup-shaped:
a. Stalk long and slender Cyathipodia (p. 10)
b. Stalk short, stout, more or less ribbed and furrowed
Paxina (p. 11)
c. Stalk rudimentary or absent, see Pustularia in Peziza-
ceae: Otideae, p. 27
II. Ascospores minutely rough, apiculate in British species
Tribe Discineae
A. Fruit body tough and persistent, attached by numerous rooting pro-
cesses
Rhizina (p. 13)
B. Fruit body fleshy, attached by the centre only Discina (p. 12)

GYROMITRA Fries

Gyromitra Fries, S u m m a Veg. Scand. Sect. P o s t . : 3 4 6 (1849).


Large species with an ovoid fruit body which becomes irregularly
lobed and convoluted b u t not alveolate, the fertile head adhering
closely to the stalk and sometimes united with it at intervals; stalk
short, stout, hollow or chambered, often furrowed or ridged externally;
spores elliptical; paraphyses slightly clavate, broad, septate.

Gyromitra csculenta (Persoon) Fries, op. cit.:346 (1849). (Fig. 4D.)


Fruit bodies up to 12 cm. high, fertile portion subglobose, more or
less lobed, wrinkled and convoluted, surface yellowish-brown to red-

6
dish-brown; stalk short and stout, becoming hollow or chambered,
surface slightly floccose, pale flesh-coloured to greyish-lavender. Asci
Tip to 325 x 18 [j.; spores elliptical, 18-22 (-24) x 9-12 jx, usually contain-
ing two or more small yellowish oil drops; paraphyses cylindrical, en-
larged to 6 - 8 [x near the tip.

Under conifers, especially on sandy soils, April. Not uncommon, especially


in the north. The figure is from a collection on coarse sandy soil under Pinus
silvestris, Aviemore, Inverness-shire, 12.iv. 1949.
In spite of its name this species is said to be poisonous, at least to some
people, and collectors are advised not to eat it. It fruits only in spring but
there is another autumn-fruiting species, G. infula (Schaeffer ex Fries)
Quelet.

Gyromitra infula (Schaeffer ex Fries) Quelet Enchiridion Fungorum:


272 (1886). (Fig. 30c.)
F r u i t bodies similar in size and colouring to G.esculenta, but with
a more saddle-shaped, less convoluted, darker brown head, up to
8 cm. wide. Asci 300-325x 18 |x; ascospores biseriate, narrowly ellip-
tical, 19-23x7-8(x, with two oil-drops; paraphyses often forked near
the tip, apex enlarged up to 10 [x wide.
In pine woods, especially in the Highlands. October and November.
Uncommon. Figured from Rannoch, Perthshire, 12. xi. 1962.
Pseudorhizina sphaerospora (Peck) Pouzar is a somewhat similar
fungus of coniferous woods in central Europe, with spherical ascospores
8 - 1 2 (x diameter.
Gyromitra gigas (Krombholz) Cooke, Mycographia: 191 (1878), is a
rare species of coniferous woods in Europe. It has apiculate spores and
is not a t r u e Gyromitra b u t should be referred to the genus Neogyro-
mitra Imai in the Tribe Discineae. It has been twice reported from
Britain b u t one of the records was certainly based on an overgrown
specimen of G.esculenta for its discoverer figured the spores as elliptical
and non-apiculate.

HELVELLA Linna3us ex St. A m a n s

Helvella Linnaeus ex St. Amans, Flore Agenaise:537 (1821).


Fruit bodies long stalked, the fertile head more or less saddle-
shaped, with two or three lobes strongly reflexed and adpressed to the
stalk, hymenial surface usually whitish or some shade of grey, often
becoming more or less convoluted; stalk cylindrical, hollow or cham-
bered, with the surface strongly ribbed and furrowed. Spores r a t h e r

7
broadly elliptical with one large central oil drop; paraphyses slender,
slightly clavate.

Helvella crispa Fries, Systema mycologicum 2 : 1 4 (1822). (Fig. 4c.)


F r u i t bodies up to about 10 cm. high, fertile head saddle-shaped,
2-lobed, often somewhat convoluted at the centre and w a v y at the
margin, thinfleshed, white to cream-coloured; stalk cylindrical, stout,
hollow, surface smooth b u t very uneven and deeply fluted with longi-
tudinal furrows, white or cream. Asci up to 300x 18 n; spores elliptical,
1 8 - 2 0 x 1 0 - 1 3 [A; paraphyses cylindrical, enlarged to 6 - 8 JA at the tips.
In damp woods of deciduous trees, August to November and occasionally
also in spring. Common. The figure is from a collection from Brockley
Combe, Somerset, 9.xi.l929.

Helvella lacunosa Afzelius ex Fries, Systema mycologicum 2:15 (1822).


(Fig. 4 A . )
F r u i t bodies up to a b o u t 10 cm. high, fertile head saddle-shaped
w i t h its lobes soon becoming u n d u l a t i n g or convoluted, arched away
f r o m the stalk, then sharply reflexed towards it at their margins, flesh
t h i n , dark grey or sometimes with lavender tints, under surface free
from ribs; stalk cylindrical or slightly swollen below, hollow or cham-
bered, deeply furrowed, surface smooth, grey or slightly olivaceous.
Asci up to 3 5 0 x l 8 [ i ; spores 1 7 - 2 0 x 1 1 - 1 3 [A; paraphyses cylindric-
clavate, up to 8 (A thick at the tip.
In woods, especially on burnt soil, September and October. Probably
common though said to be less so in France than H. sulcata. Figured from
a collection among fallen Castanea leaves, Englefleld Green, Surrey, 14.x.
1944.

Helvella sulcata Afzelius ex Fries, Systema Mycologicum 2:15 (1822).


(Fig. 4 B . )
Fruit bodies seldom more t h a n 6 cm. high, fertile head saddle-shaped,
somewhat convoluted towards the centre, one lobe often compressed
and pointing upwards, hymenial surface very dark grey to almost
black, under surface light grey and more or less veined; stalk short
and stout, hollow or chambered, deeply longitudinally furrowed, light
grey. Asci up to 3 2 0 x 1 8 jx; spores 17-18x 10-12 n; paraphyses cylin-
dric-clavate, up 8 [A thick at t h e tip.
In woods, September and October, probably common but less often re-
corded than H.lacunosa with which it has no doubt been confused. Dissing
considers that this cannot be distinguished effectively from H.lacunosa.
Figured from Tollymore Park, County Down, 13.ix. 1948.

8
LEPTOPODIA Boudier

Leptopodia Boudier in Bull. Soc. mycol. France 1:99 (1885).


Small fruit bodies with a fertile head varying from cup-shaped to
saddle-shaped, t h i n fleshed, hymenium whitish, grey, brown or black,
not noticeably veined or convoluted; stalk slender, hollow or stuffed,
without prominent longitudinal furrows. Asci and spores as in Hel-
vetia. In practice it is difficult to find a satisfactory division between
this and the following genus, Cyathipodia. The species fall into two
series, Levipedes, including L.elastica, with stalk and under side of
the head smooth, and Villipedes with these surfaces downy of finely
warted. The species have been m u c h confused b u t the following are
fairly easily recognisable:

Leptopodia elastica (Bulliard ex St. Amans) Boudier, Hist. Class.


Discom. d ' E u r o p e : 3 7 (1907). (PI. I A.)
F r u i t bodies large for the genus, up to 10 cm. high, head saddle-
shaped, irregularly 2-3-lobed, upper surface smooth or slightly con-
voluted, smokegrey or yellowish, drying dark brown, under surface
smooth, drying ochraceous; stalk slender, cylindrical, less t h a n 1 cm.
thick, hollow, smooth, whitish, sometimes laterally compressed b u t not
longitudinally fluted. Asci up to 3 3 0 x 2 0 [x; spores 19-22x 11-13 fx;
paraphyses cylindric-clavate.
Woods in summer and autumn, fairly common. Figured specimen from
Whitby, Yorkshire, 15. ix. 1946.

Leptopodia atra (Konig ex Fries) Boudier, op. cit.:37 (1907). (PI. 1B.)
Fruit bodies up to 5 cm. high, head becoming flattened with deflexed
margin, dark brown or blackish above, light grey and scarcely downy
b e n e a t h ; stalk slender, cylindrical, slightly enlarged at the base, very
minutely downy, clay coloured, tough, stuffed with a white pith. Asci
about 2 5 0 x 1 5 [x; spores broadly elliptical, 1 7 - 1 8 x 1 1 - 1 2 jx; paraphyses
cylindrical, slightly enlarged to 5 - 6 fx at the tip. Excipular hairs short,
terminal cell clavate and up to 20 jx across, cohering to give a minutely
warted appearance to the surface as seen under a hand-lens. According
to Madame Le Gal the young spores are often warted b u t the w a r t s
become hydrolysed and disappear at m a t u r i t y .
In woods, summer and autumn, not uncommon. Figured specimen from
Ruislip, Middlesex, 6.ix.l948.

L. pezizoides differs in the brown villose under surface of the pileus;


its shape is variable to a similar degree in t h e two species.

9
Leptopodia pezizoides (Afzelius ex Fries) Boudier, op. cit. :37 (1907).
(Fig. 6 J . )
Fruit body about 5 cm. high, head cupshaped to saddle-shaped,
hymenium black, under surface reddish-brown, downy; stalk black,
solid, distinctly velvety with black hairs. Asci up to 250x18 |x; spores
elliptical, 18-20 x 10-12 [A ; paraphyses cylindric-clavate with dark
brown contents.

In woods, August, rare.

Leptopodiaephippium (Leveille) Boudier, op. cit.:37 (1907). (Fig.31B.)


Fruit bodies up to 5 cm. high, head irregularly saddle-shaped, pale
gray to gray-brown, paler and villose beneath, stalk cylindrical or
slightly compressed, hairy, pale gray-brown to yellowish. Asci about
250x15 (x, spores 19-21x11-12 (x.
On soil in deciduous woods, August to October. Not uncommon.
Differs from L. pezizoides in the pale hymenium; L. murina Boudier
is a synonym.

Leptopodia stevensii (Peck) Le Gal in Rev. de Myc. 2:156 (1937).


Fruit bodies up to 5 cm. tall but usually shorter, head cup-shaped
then more or less deflexed but rarely saddle-shaped, hymenial surface
whitish to cinnamon, drying pale ochraceous, under side whitish,
finely hairy; stalk cylindrical, concolorous, hairy. Asci about 250x 15 [x,
spores 18-19x12-13 ix. (Fig.3lD.)
On soil of deciduous woods, June to October. Not uncommon.

CYATHIPODIA Boudier
Cyathipodia Boudier, Hist. Glass. Discom. d'Europe:39 (1907).
Differs from Leptopodia in the hymenophore being permanently
cup-shaped; stalk long and slender; asci and spores as in Helvetia.
Macropodia Fuckel is not separable from Cyathipodia and is an illegiti-
mate name, a later homonym of Macropodia R. Br.

Cyathipodia macropus (Persoon ex Fries) Dennis Ed. I : 7 (1960)


Peziza macropus Persoon ex Fries, Systema mycologicum 2:57 (1822)
(PI. Ic.)
Cup 3-4 cm. across, thin-fleshed, grey, downy beneath, seated on a
long slender stalk which tapers slightly upwards, solid, sometimes
furrowed below, grey, covered throughout with downy grey hairs that

10
cohere in minute t u f t s , flesh white. Asci up to 3 5 0 x 2 0 [i; spores
elliptic-fusiform, distinctly more pointed at each end t h a n in most
Helvellaceae, usually with a large central oil drop and a smaller one
at each end, 2 0 - 3 0 x 10-12 (/.; paraphyses cylindrical, rather sharply
enlarged up to 10 jx at the tip.
In woods, both of conifers and of deciduous trees, July to October. Com-
mon. Figured specimen from Pseudoisuga plantation, Glenariff, County
Antrim, 12.ix.1948.

Cyathipodia villosa (Hedwig ex O. Kuntze) Boudierop. cit.: 39 (1907).


Fruit bodies up to 6 cm. tall, head cup-shaped, sometimes becoming
flattened and lobed with age, gray, outer surface concolorous, hairy;
stalk cylindrical or tapered upwards, concolorous, hairy, base white.
Asci 250x14(1, spores 17-21x9-12,5(1. (Fig. 31c.)
On soil in deciduous woods. June-October. Not uncommon.
Superficially this resembles small specimens of C.macropus from
which it is easily separated b y the broadly elliptical spores with only
one large guttule.
Helvella cupuliformis Dissing & Nannfeldt is a similar b u t smaller,
short-stalked, species with light brown hymenium and broader spores
1 8 - 2 0 X 1 1 . 5 - 1 3 [I.

Cyathipodia corium (Weberbauer) Boudier, Hist. Class. Discom. d ' E u -


rope:39 (1907). (Fig. 4E.)
Cup usually less t h a n 2 cm. diameter, occasionally up to 3 cm. when
fully expanded, thin-fleshed, black, downy b e n e a t h ; stalk slender,
about as long as the diameter of the cup, black and downy with thin-
walled septate hairs which v a r y from nearly hyaline to dark grey-
brown. Asci up to 3 0 0 x l 8 [ x ; spores broadly elliptical with one large
oil drop, 2 0 - 2 2 x 1 2 - 1 4 |x; paraphyses r a t h e r a b r u p t l y clavate, sooty
brown, 6 - 7 [x wide at the tip.

On sandy soils, especially of coastal dunes,. July to October, occasionally


in April and May. Locally abundant. Figured from Braunton Burrows,
North Devon, 16.x. 1946.

PA XINA O. Kuntze

Paxina O. Kuntze, Revisio genera p l a n t a r u m Pars 2:864 (1891).


Rather large cup-shaped apothecia with short stout base, usually
more or less ribbed and furrowed, the under side of the cup sometimes
veined; asci, paraphyses and spores as in Helvella.

11
Paxina acetabulum (Linnaeus ex St. Amans) 0 . Kuntze. op. cit.:864
(1891). (PI. 1E.)
Cup large, up to 6 cm. across, h y m e n i u m dark brown, outer surface
paler, minutely downy; stalk short and stout, conspicuously ribbed
and furrowed, t h e ribs often passing upwards into prominent forked
veins on t h e under side of t h e cup, whitish or clay-coloured; flesh
white, hollow or chambered. Asci up to 4 0 0 x 2 0 [x; spores broadly
elliptical, 18-22 x 12-14 ^; paraphyses somewhat clavate, up to 6 n at
t h e tip.
Open woodlands and heaths, April to June and occasionally until August,
uncommon.
Some authors recognise also P. sulcata (Persoon) O. Kuntze, op. cit.: 864
(1891) differing in its smaller size with the ribs of the stalk scarcely encro-
aching on the under side of the cup but this may be only an undeveloped
state of P. acetabulum.
Paxina costifera (Nannfeldt) Stangel is a similar b u t a u t u m n - f r u i t -
ing species, distinguished b y its grayish h y m e n i u m and broader ribs,
connected by low anastomosing folds. Helvella queletii Bresadola,
variously referred to Paxina and Cyathipodia, is like a slender-stalked
P. acetabulum with a closely ribbed stalk, from which the ribs do not
spread on to the under side of the cup. This appears to be rare in
Britain, where it has formerly been referred to as H.bulbosa (Hedwig
ex Fries) Massee. According to Dissing the true H. bulbosa was based
on Cyathipodia macropus.

Paxina leucomelas (Persoon) 0. Kuntze, op. c i t . : 8 6 4 (1891). (PI. ID.)

Cups rather small for the genus, up to 3 cm. across, not expanding
b u t sometimes splitting slightly at the margin, hymenium rather dark
grey-brown, outer surface paler grey, minutely tomentose, not ribbed;
stalk short and stout, slightly ribbed and furrowed, whitish, hollow.
Asci up to 3 3 0 x 2 0 n; spores 18-23x 12-13 [x; paraphyses cylindrical
with grey-brown contents.

On soil under conifers, December to March, uncommon. Figured specimen


from i a r i ^ p l a n t a t i o n , Horsley, Surrey, 12.xii.1948.

DISCINA Fries

Discina Fries, S u m m a Veg. Scand. Sect. P o s t . : 3 4 8 (1849).


Apothecia cup-shaped, becoming expanded, h y m e n i u m some shade
of yellow or brown, stalk short and stout, sometimes furrowed. Asci

12
not blued by iodine, spores with a prominent hyaline apiculus at each
end.

Discina perlata (Fries) Fries, S u m m a Veg. Scand. Sect. P o s t . : 3 4 8


(1849). (PL IIB.)
Apothecia large, up to 8 cm. diameter, soon expanded or reflexed
with an undulating margin, hymenium bay-brown, often wrinkled,
under surface flesh-coloured or pale ochraceous, often veined; stalk
short, stout, solid, furrowed. Asci up to 4 5 0 x 1 2 jx; spores fusiform with
a hyaline apiculus at each end, minutely warted, with three oil drops,
30-35 x 12-13 jx; paraphyses cylindrical, slightly clavate at the tip and
about 8 - 9 |i wide, with brown granular contents.

Coniferous woods, especially in the Highlands, May. Rare. I have seen


no British collections and the figure is based on continental material.
There are two additional rare species in the Alps, D.leucoxantha
Bres., with yellowish h y m e n i u m and ascospores 3 2 - 3 6 x 1 2 - 1 5 [x with
t r u n c a t e appendages, and D.Melaleuca Bres., with blackish-brown
hymenium, white receptacle and flesh and fusiform ascospores,
1 7 - 2 1 x 8 - 9 [x, w i t h o u t appendages.

RHIZINA Fries

Rhizina Systema mycologicum 2 : 3 3 (1822). (PL IF.)


F r u i t body with dark brown to black h y m e n i u m and tough brown
flesh, attached to the soil or wood b y numerous, cylindrical, branched,
root-like processes; spores fusiform, apiculate, rough.

Rhizina undulata Fries, op. cit. 2 : 3 3 (1822). (PI. IF.)


Hymenophore flat or convex, often irregularly lobed, hymenial sur-
face undulating, dark brown to black with a paler margin, flesh red-
dish-brown, tough, fibrous; under surface pale ochraceous, bearing
numerous cylindrical, branched, whitish, root-like structures, 1 - 2 m m .
thick. Asci about 400x20jx; spores fusiform, with two or more oil
drops and with a hyaline apiculus at each end, 2 2 - 4 0 x 8 - 1 1 jx; p a r a -
physes slightly clavate, hyaline b u t with their tips covered b y a t h i n
amorphous brown crust.

On debrisJn coniferous woods, especially in clearings and after fires but


said to be parasitic on seedling pines, June to October. Common.

13
Family 3. Pezizaccae

Fruit body an apothecium, cup-shaped, sometimes flattened at


m a t u r i t y but t h e n relatively thin-fleshed, sessile or only short-stalked,
of medium to large size, i.e. usually more t h a n 1 cm. diameter, hymenium
usually some shade of brown, olive, mauve, purple or black, sometimes
cream-coloured b u t seldom bright yellow (see Sowerbyella), never red,
nor bright orange unless the apothecium is ear-shaped and split down
one side, outer surface smooth, scurfy or minutely tomentose b u t not
bearing distinct hairs or bristles. The family comprises two tribes:

1. Asci stained blue at the upper end in iodine solution . . . Pezizeae


2. Asci not so stained, paraphyses strongly curved at the tips
Otideae (p. 25)
The tribe Pezizeae includes three British genera:
1. Spores globose Plicaria
2. Spores elliptical:
A. Apothecia at first subterranean, then exposed, very large
Sareosphaera (p. 15)
B. Apothecia cup-shaped, exposed from the first on the surface of soil,
plant debris, plaster or dung (partly sunk in sand in P. ammophila)
Peziza (p. 16)

PLICARIA Fuckel

Plicaria Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:325


(1870).
Apothecia cup-shaped to flattened, dark coloured; asci blued at the
tip b y iodine, spores globose, smooth or rough. (For other terrestrial
species with globose spores b u t asci which do not t u r n blue with iodine
see Barlaeina p. 29 and Lamprospora p. 53.

Plicaria leiocarpa (Currey) Boudier in Bull. Soc. mycol. France 1:102


(1885). (PL l i e . )
Apothecia up to 6 cm. across, soon expanded and flattened, hymeni-
u m sooty brown, under surface slightly paler, minutely scurfy, flesh
grey-brown, becoming yellowish when cut and exuding a yellow juice.
Asci about 200xl0[i.; spores uniseriate, smooth, hyaline or faintly
brownish, containing numerous small oil drops, 8 - 9 ^ diameter; para-
physes septate, slender, slightly clavate and up to 5 n thick at the tip,
with yellowish sap.

14
On burnt ground, especially sandy heaths, from October to April, mainly
from November to February. Common. Figured material from burnt ground
where a forest fire had occurred the previous autumn in sandy pine woods,
Peaslake, Surrey, 15.ii. 1948.

Plicaria trachycarpa (Currey) Boudier in Bull. Soc. myc-ol. France 1:


102 (1885). PI. IID.)
Similar to the preceding b u t smaller and thicker, the disc at first
brown, becoming blackish, the under surface dark grey and finely
scurfy. Asci about 2 0 0 - 2 5 0 x 1 4 - 1 8 ji; spores hyaline then pale brown,
l l - 1 5 n diameter, covered with minute hyaline w a r t s ; paraphyses
slender, slightly clavate and up to 8 ji thick at the tip, apical cell with
brown oily contents and coated with a thin brown epithecial crust.

On burnt ground and on sterilised soil, October to May. Uncommon.


In the var. muricata Grelet in Bull. Soc. Bot. Centre Ouest:44 (1937), the
low warts on the spores are replaced by more prominent spines. The figure
shows a spore of this variety on the left and one of the typical variety on
the right.

SARCOSPHAERA Auerswald

Sarcosphaera Auerswald in Hedwigia 8 : 8 2 (1869).


There is a single British species:

Sarcosphaera eximia (Durieu & Leveille) R. Maire in Bull. Soc. Hist,


nat. de L'Afrique du Nord 8 : 7 9 (1917). (PI. IIF.)
Apothecium developed under the soil surface as a hollow, smooth,
white ball, which splits open in a stellate m a n n e r to expose t h e violet-
coloured hymenium. Mature specimens m a y be very large, up to
18 cm. diameter. Flesh white, fragile; under surface white to cream-
coloured, minutely tomentose, usually with fragments of soil or
h u m u s attached. Asci about 3 0 0 - 3 6 0 x 1 2 - 1 3 ji, stained blue at the tip
with iodine, spores broadly elliptical with r a t h e r a b r u p t l y t r u n c a t e d
ends, hyaline, 13-15x7-8(1, with two oil drops; paraphyses septate,
constricted at t h e septa, often once or twice branched above, the
apical cell up to 7 (j. wide, with brownish sap.

In woods, especially of Fagus, on chalk and limestone, May. Locally


common. Figured material from the downs near Chisledon, Wiltshire, 31. v.
1948.
This magnificent species is commonly known as S.coronaria (Jacquin ex
Cooke) Boudier which appears to be an untenable name under the modern
Code of Botanical Nomenclature.

15
P E Z I Z A St. A m a n s

Peziza St. Amans, Flore Agenaise:530 (1821).


Apothecia cup-shaped, becoming flattened in some species, sessile
or very short-stalked, receptacle usually thin, brittle-fleshed and often
more or less scurfy externally. Asci blued at the tip by iodine, asco-
spores r a t h e r large, usually elliptical, occasionally somewhat spindle-
shaped. The species fall into two series, one with smooth spores
without oil drops, the other with spores ornamented by w a r t s or ridges
and usually containing two large oil drops. The second series includes
a few species whose flesh yields a coloured fluid and was formerly given
generic rank under t h e name Galactinia. The two groups are not com-
pletely clear cut, however, and it is generally agreed t o d a y t h a t t h e y
cannot be separated generically. The spore ornament is a valuable
specific character b u t is difficult to see clearly in unstained spores or
in those stained with iodine. It is best demonstrated by staining with
Cotton Blue in lacto phenol. Presence of a coloured juice is most easily
detected by touching the freshly broken flesh with a clean white
handkerchief. The h a b i t a t of the species is often characteristic, several
occur on or beside rotting wood, others on b u r n t ground, one on dung,
another partly buried in sand and the remainder mostly on soil in
woods, fields and gardens. There are about three dozen British species,
some highly critical and only the commonest or most easily recognised
are described below. For keys to the western European species the
reader is referred to a valuable paper by Madame Le Gal, published
as a supplement to Revue de Mycologie (N.s.) 6 (1941),or to Moser,in
Gams Kleine Kryptogamenflora I l a : 95-102 (1963). Several species
are known to have an QSdocephalum conidial state, which confirms
the affinity with Iodophanus (p. 64) indicated b y the iodine reaction.
See Berthet in Bull. Soc. mycol. France 80:125-149 (1964).

Peziza ammophila Durieu & Montagne, Exploration scientiflque de


l'Algerie Botanique, Atlas, t a b . 28, fig. 2 (1847). (PI. IIIA.)
Apothecia cup-shaped and semi-immersed in sand, finally splitting
in a stellate m a n n e r and becoming flattened on the sand surface, disc
dark brown, receptacle pale ochraceous, seated on a pseudostipe of
sand grains firmly bound together b y mycelium. Asci about 2 0 0 x 1 5 jx;
spores elliptical, smooth, 14-16 x 10 n; paraphyses straight, slightly en-
larged to 7 (1 at the tip.

Coastal sands, often amongst Ammophila, September to December,


locally abundant. Boudier referred the species to Geopyxis but its asci give

16
a strong blue reaction to iodine. Figured material from sand dunes of
Gibraltar Point, on the Lincolnshire coast, September and October 1948.

Peziza repanda Persoon, Mycologia europaea 1:231 (1822). (PI. 11 ID.)


Apothecia large, occasionally up to 12 cm. diameter, cup-shaped,
becoming expanded and undulating, sessile, h y m e n i u m light chestnut-
brown; flesh whitish or fawn; outer surface whitish or pale fawn to-
wards t h e often crenate margin, finely scurfy. Asci up to 3 0 0 x 1 3 [x;
spores elliptical, smooth, 1 5 - 1 6 x 9 - 1 0 fx; paraphyses very slender,
slightly clavate at t h e tip.
On the ground in woods, luxuriant forms on sawdust heaps. The flesh is
stratified like that of P. varia but the disc is a different shade of brown and
the peculiar moniliform paraphyses of that species are lacking. Figured
material from Gerrard's Cross, Buckinghamshire, 5. v. 52.

Peziza micropus Persoon, op. cit. 1:227 (1822). (PL I l l s . )


Apothecia of medium size, seldom over 4 cm. diameter, solitary or in
small groups, cup-shaped, often with a small stalk in a crack of the
bark, disc fawn to pale chestnut-brown, margin prominently d e n t a t e ;
flesh thin, fawn, not stratified like t h a t of P. varia-, outer surface
whitish to pale fawn, scurfy, especially near the margin. Asci about
250x14 [x; spores elliptical, smooth, 1 5 - 1 9 x 9 - 1 1 [x; paraphyses slen-
der, straight, scarcely clavate, 4 - 5 jx thick at t h e tip.
On rotting timber and bark of fallen trees, especially Fagus according
to Boudier, September. Perhaps uncommon but much' confused with
P. varia, P. repanda and P. ampliata. Figured material on rotting timber,
Rotherham, Yorkshire, 19. ix. 1947.

Peziza varia (Hedwig) Fries, Systema mycologicum 2 : 6 1 (1822). (PL


IIIE.)

Apothecia of medium size, seldom more t h a n 5 cm. diameter when


fully expanded, cup-shaped b u t soon flattened or even convex, often
with a rudimentary stalk, hymenium light grey-brown at first b u t
soon becoming dark greyish-brown; flesh w a t e r y grey to f a w n ; outer
surface whitish to fawn, scurfy. Asci up to 2 8 0 x 1 4 (x; spores elliptical,
smooth, 1 4 - 1 6 x 9 - 1 1 [x; paraphyses mostly with the lower or middle
cells much inflated, up to 20 fx wide, apical cells slender, tips slightly
clavate.
On soil rich in humus and in contact with rotting wood, decaying tree
roots and the like, common in June but also found in autumn. Figured
material from Nottingham, 27. vi.1951.

17
The flesh of the cup is conspicuously stratified in five layers, beneath the
asci is a subhymenium of small cells, then follow a zone of large subglobose
or vertically elongated cells, a zone of slender woven hyphae, a lower zone
of large globose cells and finally a surface zone of smaller cells and hyphae.
The central hyphal band is often visible as a dark line when a section of
the flesh is viewed under a hand lens.

Peziza ampliata Persoon ex Persoon, Mycologia europaea 1 ;227 (1822).


Apothecia sessile, usually r a t h e r small, 1 - 3 cm. diameter, soon
widely expanded with flattened disc, hymenial surface light brown to
yellowish-brown, receptacle a little paler and pruinose, margin often
dentate, flesh thick and w a t e r y . Asci about 250 x 15 jx, spores elliptical,
smooth, 1 7 . 5 - 2 0 x 1 0 - 1 1 n, w i t h o u t guttules; paraphyses straight,
cylindrical or with some segments slightly inflated, apex rounded,
5 - 8 [x wide. The flesh is not stratified b u t contains numerous subglobose
cells up to 150 (x across, joined b y short lengths of hyphae 6-12 [x wide.
On very rotten wood and on soil containing decayed woody debris. May.
The species seems distinct from other brown lignicolous Pezizas in its
larger spores, relatively thick flesh, peculiar unstratified flesh and probably
more yellowish hue but the traditional identification with Persoon's fungus,
on bark and tree-roots in autumn, is somewhat dubious. (Fig. 31 G.)

Peziza arvernensis Boudier in Bull. Soc. bot. France 26: X X V I (1879).


Apothecia large, up to 4 cm. diameter, light yellowish-brown, cup-
shaped, t h e n expanded and undulating, fragile, receptacle paler, al-
most smooth. Spores elliptical, 1 7 - 1 9 x 9 - 1 0 [x, without guttules b u t
with a fine p u n c t a t e ornament on the wall; paraphyses straight,
rounde at the tip, g u t t u l a t e . (Fig. 31 H.)
On the ground in woods, May to July.
Distinguished from P. repanda and its allies by the very finely verrucose
spores.

Peziza sterigmatizans Phillips apud Cooke, Mycographia: 175 (1879).


Apothecia shallow-cupshaped, sessile, soon expanded and even
repand, about 2 cm across, disc dark brown, flesh thin, whitish, re-
ceptacle pale. Asci 2 5 0 - 3 0 0 x 1 3 - 1 5 jx, spores elliptical, 17-20(-23)x
9-10(-13) [x, smooth, without guttules; paraphyses rather thick closely
septate, straight b u t commonly forked near the apex or with a short
lateral branch, apex clavate, 7-10 [x wide, with brownish oily contents.

18
On wet ground. Autumn and winter. Uncommon.
The thick branched paraphyses, for which it was named, appear cha-
racteristic. (Fig. 31 E.)

Peziza vesiculosa Bulliard ex St. Amans, Flore Agenaise:534 (1821).


(PL I I I G . )
Apothecia large, up to 8 cm. diameter, often clustered, p e r m a n e n t l y
cup-shaped, often with strongly incurved margin, hymenium light
yellowish-brown; flesh very brittle, pale f a w n ; outer surface pale fawn,
coarsely scurfy. Asci up to 3 8 0 x 2 5 jx; spores elliptical, smooth,
2 0 - 2 4 x 1 1 - 1 4 [x; paraphyses slender, straight, scarcely clavate.

On manure heaps and richly manured soil, mushroom beds, etc., August
to April, common. Figured material from Gorstorphine, Edinburgh.
The hymenium often becomes detached from the flesh to form a blister
in the centre of the cup, hence the name "vesiculosa". Luxuriant forms in
which this process has been carried to extreme lengths and the hymenium
has become strongly convoluted have been called var. saccata Fries, Systema
mycologicum 2:53 (1822).

Peziza echinospora Karsten in Not. Sallsk. F a u n a et Flora fennica 10:


115 (1869). (PI. I l l F.)
Apothecia up to 8 cm. diameter b u t usually m u c h smaller, cup-
shaped, sessile, disc dark brown, with a conspicuously incurved den-
t a t e margin; flesh thin, b r o w n ; outer surface light brown or almost
white below, scurfy. Asci about 3 0 0 x l 4 ; x ; spores r a t h e r oblong-ellip-
tic, finely warted, 1 4 - 1 8 x 7 - 9 . 5 y.; paraphyses straight, slightly clavate.
On burnt soil and charcoal, May to October. Common. Figured material
on charcoal, Berkhamsted Common, Hertfordshire, 3.x. 1948.
It has been suggested that this fungus is the old Peziza pustulata (Hed-
wig) Persoon but modern authors are agreed in regarding Hedwig's species
as a synonym of Pustularia catinus. Boudier called the present species
Aleuria umbrina but it is not an Aleuria and this epithet cannot be trans-
ferred to Peziza as there is already a P. umbrina Persoon.

Peziza sepiatra Cooke in Grevillea 3 : 1 1 9 (1875) (Fig. 31 F.)


Apothecia sessile, about 1 - 2 cm. diameter, shallow cup-shaped, t h e n
flattened, disc very dark-brown, receptacle concolorous, scurfy, with
surface cells 14-20 fx diameter. Asci 280-300 x 16-18 jx, spores elliptical,
18—20x 11—12 [x, smooth, without guttules; paraphyses straight, simple,
apex rounded, 5 - 8 fx wide.
On soil, charcoal and rotting textiles. July. Apparently uncommon.

19
Peziza linteicola Phillip sand Plowright, described on rotting cloth,
seems to have been similar b u t with smaller spores, 1 2 - 1 5 x 8 - 1 0 fx, yet
Plowright has endorsed a collection of Crossland's with spores 18-19x
11-12 (x "This is m y idea of P. linteicola—I have no doubt about it
being the right t h i n g . " There appears, however, to be a terrestrial
Peziza of the general appearance of P. sepiatra b u t with smooth
elliptical spores 1 5 - 1 6 x 7 . 5 - 8 fx so t h a t P. linteicola m a y yet prove a
good species. It can scarcely be restricted to rotting textiles.

Peziza violacea Persoon, Mycologia europaea 1 : 2 4 2 (1822). (PL IIIJ.)

Apothecia seldom more t h a n 3 cm. across, cup-shaped t h e n soon


fully expanded, sessile, h y m e n i u m dark violet or purplish-brown;
flesh r a t h e r thin, pale p u r p l e ; outer surface greyish to light-brown,
pruinose. Asci up to 3 0 0 x 1 5 fx; spores elliptical, smooth, 1 3 - 1 5 x 7 - 9 |x;
paraphyses slender, distinctly clavate and often somewhat curved at
t h e tip, which contains purplish-brown sap.
On burnt ground, often amongst Funaria or with Plicaria leiocarpa, in
spring and autumn. Uncommon. Figured material from burnt ground at the
head of Glen More, Aviemore, Inverness-shire, 14.iv.1949.
The flesh is composed of large globose cells interspersed with hyphae and
with a narrow hyphal stratum near the outer surface. The violet colour may
be almost imperceptible and the curved paraphyses are then a useful check
character.

Peziza cerea Sowerby ex Merat, Nouvelle Flore Paris 1: 25 (1821).


(PL IIIc.)
Apothecia cup-shaped, becoming expanded, up to 5 cm. across,
h y m e n i u m pale ochraceous or yellowish-brown; flesh white, firm;
outer surface white, minutely scurfy. Asci to 3 5 0 x 1 6 n; spores ellipti-
cal, smooth, 1 4 - 1 7 x 8 - 1 0 [x; paraphyses scarcely clavate, straight.
On woody debris, rotting sandbags and on the soil between paving stones
in damp cellars, at all seasons. Common. Figured material from Ilighgate,
London, 3.vii.l947.

Peziza tectoria Cooke, Mycographia, fig. 263 (1897), described from


d a m p plaster and m o r t a r , is probably a synonym.

Peziza emileia Cooke, Mycographia, fig. 379 (1879). (PI. IVA.)


Apothecia gregarious, cup-shaped or finally expanded with an un-
dulating margin, sessile, h y m e n i u m varying from violaceous with
yellowish tints to fawn; flesh t h i n ; outer surface whitish, scurfy and

20
often yellowish towards the margin. Asci a b o u t 250-12 [x; spores
elliptical, with two oil drops, finely warted, 1 7 - 2 2 x 8 - 1 1 n; paraphyses
slender, slightly clavate, straight, colourless or containing violaceous
granules.
On lawns and in woods, June to October. Uncommon. Figured material
from a lawn, Canwick Hall, Lincoln, 9.x. 1954.
There is considerable variation in the tint of the hymenium and the name
Peziza howsei Boudier in Bull. Soc. Bot. France 26:LXXV (1879), has been
given to a distinctly violaceous form.

Peziza praetervisa Bresadola in Malpighia 11: 266 (1897). (PI. IVB.)

Apothecia up to 3 cm. diameter, often clustered, cup-shaped t h e n


expanded, violet, sometimes with brownish t i n t s ; flesh thin, m a u v e ;
outer surface pale violaceous, minutely scurfy. Asci about 250x10 jx;
spores with two oil drops, finely warted, 11-13 (-15) x 6 - 8 [x; para-
physes slightly clavate, mostly slightly curved at t h e tip, which is filled
with purplish granules.
On burnt ground in woods and heaths, occasionally on sawdust, October
to May. Common. Figured material from burnt ground, Peaslake, Surrey,
15.ii.1948.

Peziza petersii Berkeley & Curtis in Grevillea 3: 150 (June 1875). (PI.
IVc.)
Apothecia up to 5 cm., often clustered, cup-shaped, hymenium
brown with greyish t i n t s ; flesh thin, grey, outer surface lead-grey
below, becoming brownish towards t h e margin, scurfy. Asci about
200x10 [x; spores with two oil drops, finely warted, 1 0 - 1 2 x 5 . 5 - 6 (x;
paraphyses somewhat clavate, up to 7 [x thick at the tip, which con-
tains brown globules and is often curved.
On burnt ground in woods and on charred stumps, June to October. Un-
common. Figured from burnt ground, Odell Great Wood, Bedfordshire,
27.ix.1953.
Galactinia sarrazinii Boudier in Bull. Soc. mycol. France 3: 147
(1887), is a synonym.

Peziza badia Persoon ex Merat, Nouvelle Flore Paris 1: 24 (1821). (PI.


IVD.)

Apothecia up to 8 cm. across, sessile, more or less cup-shaped b u t


irregularly undulated in old specimens, h y m e n i u m olive-brown, drying
darker; flesh thin, pale reddish-brown with w a t e r y juice; outer surface

21
reddish-brown, scurfy, especially towards t h e margin. Asci up to
330 x 15 [x; spores elliptical, mostly with two oil drops but one is often
larger t h a n the other, ornamented with short ridges which tend to
unite to form an irregular reticulum, 1 7 - 2 0 x 9 - 1 2 jx; paraphyses
straight, slightly clavate.
On the ground in woods, especially on sandy soil, August to October.
Probably fairly common but much confused with other species.
Old published records of P. badia cannot be accepted unless they are
substantiated by recognisable specimens. Several related species are best
distinguished by details of spore ornament, as shown by Madame Le Gal
in the paper cited above. Amongst these are Peziza badioconfusa Korf in
Mycologia 46:838 (1954) ( = Galactinia olivacea Boudier in Bull. Soc. mycol.
France 13:14 [1897]) with narrower, finely warted, spores, 17-21x8-10^
and Galactinia limosa (Grelet) Le Gal & Romagnesi in Revue de Mycologie
4:176 (1939), which has spores with warts elongated into short ridges but
not forming a network and grows on bare clay round the margins of pools.

Peziza michelii (Boudier) Dennis E d . 1:15 (1960)


Galactinia michelii Boudier in Bull. Soc. mvcol. France 7: 215 (1891).
(PL I V F . )
Apothecia up to 5 cm. diameter, sessile, cup-shaped, then somewhat
expanded with a slightly raised even margin, hymenium reddish-
b r o w n ; flesh thin, yellowish; outer surface fawn, minutely scurfy to-
wards the margin. Asci up to 2 8 0 x 1 5 jx; spores elliptical, with two oil
drops, ornamented b y well defined low warts, 1 3 - 1 7 x 7 - 9 [x; para-
physes slightly clavate, straight, with yellowish contents.
On bare soil in woods, September. Uncommon. Figured from bare soil of
a shaded bank by the Tarras Water, Eskdale, Dumfries-shire, 20 .ix. 1948.
Madame Le Gal has noted the occurrence of a violet form of this species
in France, both in spring and autumn.

Peziza ionella Quelet in Bull. Soc. bot. France 24: 328 (1877). (PL IVE.)
Apothecia less t h a n 0.5 cm. diameter, gregarious, violaceous
t h r o u g h o u t , disc soon flattened, with a minutely denticulate margin,
seated on a m i n u t e central stalk; outer surface scurfy. Asci about
3 0 0 x 1 8 jx; spores fusiform, with one to three large oil drops and
numerous small ones, smooth, 2 5 - 2 8 x 8 - 9 |x; paraphyses straight,
slender, colourless.
On heavy soil in woods, in summer. Rare.

Peziza succosa Berkeley in Ann. Mag. n a t . Hist. 6 : 3 5 8 (1841). (PL IVG.)


Apothecia 2 - 5 cm. across, disc grey-bistre with a slight olivaceous
tint, permanently cup-shaped; flesh thin, whitish, yielding copious

22
bright yellow juice; outer surface grey, sometimes yellowish at the
margin. Asci about 3 5 0 x l 8 j x ; spores with two large oil drops, orna-
mented b y coarse w a r t s and short ridges, 1 7 - 2 2 x 9 - 1 2 |x; paraphyses
straight, colourless, slightly clavate.
On the ground in woods, July to September. Common. Figured material
from beech woods, Arundel, Sussex, 3. viii. 1954.

Peziza saniosa Schrader ex Fries, Systema mycologicum 2: 65 (1822).


(PI. IILH.)

Apothecia about 1-1.5 cm. across, soon expanded and flattened,


disc very dark blue-grey; flesh concolorous, yielding bluish juice; outer
surface dark grey-brown, scurfy. Asci about 2 5 0 x 1 3 ;x, contents
brownish; spores with two oil drops and r a t h e r coarsely warted,
1 4 - 1 6 . 5 x 7 - 9 [A; paraphyses straight, colourless, slightly enlarged up-
wards.
On the ground in woods, October. Rare. Figured material from forestry
nurseries, Wykham, Yorkshire, 7.x. 1945,

Peziza badiofusca (Boudier) Dennis E d . 1:16 (1960)


Galactinia badiofusca Boudier, Hist. Class. Discom. d ' E u r o p e : 48 (1907).
(PI. I I J . )
Apothecia about 1-1.5 cm. across, soon e x p a n d e d and flattened, disc
bay-brown; flesh concolorous, yielding copious opalescent juice; outer
surface paler, scurfy, margin minutely crenulate. Asci about 300x 15 [x,
contents yellowish; spores 1 3 - 1 5 x 9 - 1 0 (x, usually with one large cen-
tral oil drop, ornamented b y numerous small regularly arranged w a r t s ;
paraphyses straight, colourless, slightly clavate, their tips united b y a
thin, amorphous, brown, epithecial crust.
On bare ground in woods, September to November. Uncommon. Figured
from amongst moss on bare clay under Carpinus, Watery Grove, Stevenage,
Hertfordshire, 30.ix.1951.

Peziza brunneoatra Desmazieres in Ann. Sci. Nat. Bot. II, 6 : 244


(1836). (PI. IIG.)
Apothecia up to 1 cm. diameter, disc soon flattened, with a slightly
raised even margin, blackish-brown; flesh r a t h e r thick, reddish-brown;
cup shallow, sessile, dark brown, almost smooth. Asci up to 3 0 0 x 1 7 jx;
spores elliptical, finely warted, packed w i t h numerous small oil drops,
1 4 - 1 8 x 9 - 1 1 |x; paraphyses slender, slightly thickened above, with oily
brown contents.

23
On the bare ground, rare. There is no British material of this species at
Kew but it should be easily recognisable and it has therefore been figured
from Desmazieres original material collected at Lille.

Peziza atrovinosa Cooke & Gerard in Bull. Buffalo Acad. Sci.: 288
(1875). (PI. IIH.)
Apothecia 1 - 2 cm. diameter, shallow cup-shaped becoming flatten-
ed, sessile, disc dark brown to blackish; flesh relatively thick, com-
posed throughout of large rounded cells, yielding no juice. Asci
2 6 0 x 1 1 jx; spores elliptical, 1 3 - 1 4 x 7 . 5 - 8 (x, containing two oil drops,
ornamented with a more or less continuous network of prominent
ridges, paraphyses cylindrical, 3 (x thick. Plicaria fulva R. Schneider in
Z. B a k t . Abt. 2, 108: 153 (1954), is a synonym.

Figured material from sterilised soil casing a mushroom bed, Worthing,


Sussex, 5.V.55.

Peziza fimeti (Fuckel) Seaver, N.Amer. Cup Fungi (Operculates):


232 (1928), is a small brown species found on cow dung, with spores
1 5 - 1 6 x 7 - 8 [x and smooth.

Peziza proteana (Boudier) Seaver in Mycologia 9 : 1 (1917) F o r m a spa-


rassoides (Boudier) Korf in Mycologia 48: 714 (1956). (PI. IVL.)
F r u i t b o d y v e r y large, becoming subglobose or elongated and up to
25 cm. high, extensively convoluted like a Gyromitra b u t sessile,
h y m e n i u m whitish flushed with pink, lilaceous or pale ochraceous
tints. Asci up to 3 0 0 x 1 1 [x; spores with two oil drops, 1 0 - 1 3 x 6 - 7 jx,
minutely w a r t e d ; paraphyses straight, slightly clavate, slender,
colourless or containing yellowish drops.

On the ground in woods, especially of Fagus, often after fires, September


to October. Rather frequent. Figured from Norbury Park, Mickleham,
Surrey, 14. xi. 1952.
Because of its superficial resemblance to a Gyromitra this remarkable
fungus has been classed in the Ilelvellaceae under the names Gyromitra
philtipsii Massee, British Fungus Flora 4:478 (1895), Underwoodia camp-
beltii Saccardo in Annales mycologicum 7:433 (1909), U. sparassoides
(Boudier) Banhegyi Index Horti Bot. Univ. Budapest 3:19 (1937),Durandio-
myces philtipsii (Massee) Seaver, N.Amer. Cup Fungi (Operculates):242
(1928), Daleomyces phitlipsii (Massee) Seaver N.Amer. Cup Fungi (Oper-
culates) Supplement:337 (1942) and Daleomyces gardneri Setchell in Myco-
logia 16:241 (1924). It is, however, clearly distinguished from the Helvella-
ceae by the blue reaction of its asci to iodine and in all microscopic features
it agrees with the normal cup-shaped form of P. proteana (= Galactinia
proteana [Boudier] Saccardo Sylloge Fungorum 16:709 [1902].) The latter
occurs on burnt ground in woods and on charred stumps in April and May.

24
Tribe Otideae
This contains five British genera:
I. Apothecia splitting down one side, often ear-shaped Otidea
II. Apothecia cup-shaped:
A. Spores globose Barlaeina (p. 29)
B. Spores elliptical:
1. Spores smooth:
a. Apothecia sessile, dark-fleshed. . . . Pscudotis (p. 27)
b. Apothecia often distinctiy stipitate, pale-fleshed
Pustularia (p. 27)
2. Spores rough Sowerbyella (p. 28)

OTIDEA Fuckel
Otidea Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:329 (1870).
Apothecia of medium to large size, always terrestrial, sessile or with
only a r u d i m e n t a r y stalk, markedly eccentric, split down one side and
often elongated vertically so as to resemble a r a b b i t ' s ear, colour some
shade of yellow, brown or ochre. Asci not blued b y iodine, spores
elliptical to elliptic-fusiform, with two conspicuous oil drops, smooth,
paraphyses strongly curved at t h e tip, slender, often forked. The
species are separated largely b y colour and hence are difficult to re-
cognise when not in prime condition. T h e y have a somewhat uniform
a n a t o m y , a flesh composed of r a t h e r closely woven hyaline hyphae,
covered on the outer surface b y a more or less pseudo-parenchymatous
layer of large thin-walled cells, which tend to lie in rows at right
angles to the surface. Usually the terminal cells p r o t r u d e in short
chains, which often cohere in t u f t s to give a minutely mealy appearance
to the surface. In one group of species these superficial cells and most
of those in the underlying pseudoparenchyma are covered b y large
granules of some yellow or orange substance not dissolved in Potassium
hydroxide solution. The specific nomenclature is highly confused and
the names adopted below must be regarded as subject to revision.

Otidea alutacea (Persoon) Massee, British Fungus Flora 4: 446 (1895).


(PI. V B . )
Apothecia of moderate size, often in clusters, 3 - 5 cm. high and
2 - 4 cm. across, cup-shaped or somewhat elongated on one side, split
down t h e other, subsessile, h y m e n i u m clay colour or greyish-brown;
flesh yellowish, r a t h e r t h i c k ; outer surface fawn, finely scurfy. Asci
about 2 5 0 x 1 5 (j.; spores 1 2 - 1 5 x 6 - 7 [x, with two oil drops; paraphyses
slender, curved at the tip and sometimes slightly lobed.

25
On the ground in woods, August to September. Not common. Figured from
Clandeboye, County Down, ix. 1948.

Otidea bufonia (Persoon) Boudier, Hist. Class. Discom. Europe 52 (1907)


(pl-Vc-) ^ J f t > i u A
Apothecia up to 7 cm. diameter, subsessile, cup-shaped, split down
one side, becoming u n d u l a t i n g and irregular, hymenium dark b r o w n ;
flesh thin, with hyaline hyphae 5 - 7 jx thick, covered b y a layer of three
to four large cells, 50-60 jx deep, bearing superficial 2-celled hairs
coated with ochraceous granules; outer surface light brown, minutely
downy. Asci 160-200x 10-12 jx; spores elliptic-fusiform, with two oil
drops, 1 3 - 1 5 x 6 . 5 - 7 [j.; paraphyses 3 [x thick, closely septate, strongly
curved at the tip.
In woods, August. Uncommon. Figured from Frensham, Surrey, 26. viii. 50.

OtMeacgchlmta (Linnaeus ex St. Amans) Fuckel, op. cit.: 329 (1870),


is a somewhat similar species with larger spores, 1 6 - 1 8 x 7 - 8 (x. O. um-
1
brina (Persoon) Bresadola is a s y n o n y m . y f ^ ^ - ^ ^ ^ v ^
t L- ^/t^^W if W / i ^ o A. -
Otidea onotica (Persoon) Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e
2 3 - 2 4 : 329 (1870). (PI. VA.)
Apothecia large, up to 10 cm. high and 6 cm. broad, often growing in
clusters, soon becoming irregular and "ear-shaped", short-stalked,
h y m e n i u m somewhat ochraceous b u t delicately flushed with pink, the
pink tint becomes more pronounced when the fruit body is dried;
flesh thin, white; outer surface ochraceous, sometimes flushed slightly
with pink b u t drying ochraceous, finely scurfy; stalk white at the base.
Asci up to 2 5 0 x l 0 f x ; spores rather broadly elliptical, (10)-12-13x
5 - 6 (x, with two oil drops; paraphyses slender, 3 (x thick, curved at
the tip.
On the ground in woods, especially of Querents, September. Uncommon.
Figured from Murthly, Pertshire, ix. 1953. The superficial layers of the flesh
and the hairs may be so densely encrusted with orange granules as to be
opaque even in thin sections.

Otidea concinjia (Persoon) Saccardo, Sylloge F u n g o r u m 8: 96 (1889).


(PI. VD.) ' M I J ^U^JI^-

Apothecia more or less "ear-shaped", up to 5 cm. high, bright


sulphur yellow, whitish at the base; flesh t h i n , white. Asci about
150x10 jx; spores elliptical, with two oil drops, 1 0 - 1 2 x 5 - 6 (x; para-
physes slender, 3 - 4 (x t h i c k ; surface layers of t h e flesh encrusted with
yellow granules.

26
On the ground in mixed woods and under Fagus, August to September.
Rare. Figured from under Pseudotsuga, Aviemore, Inverness-shire ,26. ix.
1950.

PSEUDOTIS Boudier

Pseudotis Boudier in Bull. Soc. mycol. France 1: 102 (1885).


This small genus was proposed to accommodate fungi anatomically
like Otidea b u t with regular apothecia not consistently split down one
side. There is one British species:

Pseudotis apophysata (Cooke & Phillips) Boudier, Hist. Class. Discom.


d ' E u r o p e : 52 (1907). (PL IVJ.)
Apothecia gregarious, sessile, cup-shaped, up to 2 cm. diameter,
hymenium dark grey-brown; flesh rather thin, black; outer surface
brown. Asci about 2 5 0 x l 8 [ x ; spores elliptic-fusiform with two large
oil drops, 22-26x 10-11 [x; paraphyses slender, curved at the tip,
contents greyish.
On wet soil, August to October. Extremely rare and figured from Phillips
watercolour sketches of the type collection, at Kew.

PUSTULARIA Fuckel

Pustularia Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 2 3 - 2 4 : 3 2 8


(1870).
Apothecia cup-shaped, entirely pale ochraceous, greyish or pale
pink, sessile or only minutely and imperfectly stipitate, externally
tomentose; asci not blued b y iodine, spores large, elliptical, biguttu-
late, paraphyses slender and colourless, sometimes lobed at t h e tip.
Where there is a r u d i m e n t a r y stalk it is usually hidden in t h e soil.
Hence the beginner will probably seek the species of this genus first
in Peziza, from which it differs in the iodine reaction of the asci, or in
Aleuria, which has ornamented spores. Large specimens of P. catinua
might even be sought in Otidea, which has narrow curved paraphyses
and smaller spores.

Pustularia catinus (Holmskjold ex Fries) Fuckel, op. cit.: 328 (1870).


(PL I G . )
Apothecia solitary or in clusters, 2 - 5 cm. diameter, cream to pale
ochraceous, permanently cup-shaped b u t with a somewhat crenate
margin, thinfleshed, brittle, externally downy with short hyaline hairs;

27
cups either sessile with the base slightly sunk in the ground or seated
on a short stout stalk. Asci about 350x20 jx; spores elliptical, 20-24x
11—13 fx, with two large oil drops; paraphyses slender, septate, often
branched below,tips not clavate but often lobed, 3 - 4 jx wide.
Woods, especially of Fagus, June to September. Not uncommon.

Pustularia cupularis (Linnaeus ex Fries) Fuckel, op. cit.: 328 (1870).


(PI. I J . )
Apothecia usually less t h a n 2 cm. diameter, gregarious, sessile or
nearly so, permanently cup-shaped with a crenate margin, greyish
ochraceous throughout, the exterior minutely downy. Asci about
300x15 fx; spores 19-21x13-15 jx, broadly elliptical, with two large oil
drops; paraphyses slender, septate, colourless, slightly enlarged up-
wards.
Damp places in woods, June to October. Not uncommon.

Pustularia gaillardiana Boudier in Bull. Soc. mycol. France 18: 141


(1902), is a similar but smaller species, usually less than 1 cm. across,
with grey hymenium, growing on bare soil of damp woods in autumn.
Pustularia patavina (Cooke &Saccardo) Boudier, Hist. Class. Discom.
d'Europe: 53 (1907), is equally small but has a light orange hymenium.

Pustularia rosea Rea in Trans. Worcestershire Nat. Club 8: 20 (1924).


(PI. I H . )
Apothecia up to 1.5 cm. diameter, gregarious, permanently cup-
shaped, brittle, the margin splitting slightly, bright pink throughout,
the exterior minutely downy with loose hyaline hyphae. Asci about
250x13 jx; spores broadly elliptical with two large oil drops, 17-20 x
9-11 [x; paraphyses cylindrical, very slightly enlarged upwards but
not clavate, about 6 jx thick, with pinkish granules in their upper parts.
On burnt ground, September to October. Uncommon.

SOWERBYELLA Nannfeldt

Sowerbyella Nannfeldt in Svensk Bot. Tidskrift 32: 118 (1938).


Apothecia cup-shaped, stipitate, yellow, outer surface downy, asci
not blued by iodine, spores with two oil drops, warted. There is one
British species:

28
Sowerbyella radiculata (Sowerby ex Fries) Nannfeldt, op. cit.: 119
(1938). (PI. IVH.)

Apothecia cup-shaped, up to 4 cm. diameter, hymenium lemon-


yellow; flesh thin, white; outer surface cream coloured, downy, seated
on a long rooting stalk clothed with dense white hairs. Asci up t o
250x12 [a; spores broadly elliptical, coarsely warted, 1 2 - 1 4 x 7 - 7 . 5 fx;
paraphyses slender, 3 - 5 jx thick, strongly curved at the tip and sometimes
lobed below the apex, with light yellow sap.

On the ground under conifers, September to November. Uncommon.


Figured material from under Cupressus macrocarpa, Friston Forest, Sussex,
4. ix. 1952. Boudier recorded it also under Sambucus in France. This species
was formerly erroneously named Otidea alutacea by British authors.

BARLAEINA Saccardo

Barlaeina Saccardo, Sylloge F u n g o r u m 14: 30 (1899), emend Le Gal


in Revue de Mycologie 18: 80 (1953).

As explained above under Plicaria this genus has been adopted for
Pezizaceae with globose spores in asci which do not t u r n blue with
iodine. The paraphyses have the strongly curved apices characteristic
of the Otideae.

Barlaeina amethystina (Quelet) Saccardo & Traverso, Sylloge Fungo-


r u m 19: 137 (1910). (PI. IIE and Fig. 1A.)

Apothecia rather small, about 1 cm. across, with flat violet disc,
sessile; flesh thick, soft; outer surface paler. Asci about 2 0 0 x 1 5 fx;
spores globose, coarsely warted, 10-13 y. diameter, hyaline, containing
one large oil drop; paraphyses slender, septate, scarcely enlarged up-
wards and about 5 n thick at t h e tip, strongly curved above and united
by a slight purplish epithecial crust of amorphous m a t t e r .

On humus and roots in swamps, under Gh^oen^Jjuicm and Filipendula,


also on bare soil under Tussilqgo, July to October. Uncommon. Figured
material from Wheatfen Broad, Norfolk, 16. viii. 1947.

Barlaeina persoonii (Crouan) Saccardo & Traverso, Sylloge Fungo-


rum 19: 140 (1910), is a very similar species b u t has perfectly smooth
spores.

29
Family 4. Humariaceae

Fruit body sometimes cup-shaped b u t more often relatively thick-


fleshed and flattened or convex, usually small, often bearing hairs or
bristles on the outer and under surface (Fig. 5),hymenium often bright
red or orange, sometimes white or pallid, in a few species brown, asci
not blued by iodine, contents of t h e paraphyses often turned greenish
in brightly coloured species, spores smooth or ornamented, colourless.
The family contains three tribes:

A. Paraphyses not green with iodine, hymenium usually whitish or


pale ochraceous, outer surface clothed with white or brown hairs
Lachneae
B. Paraphyses usually green in iodine, hymenium red, orange, yellow
or rarely light brown:
1. Margin and outer surface conspicuously hairy
Ciliarieae (p. 37)
2. No distinctive hairs present Aleurieae (p. 46)

Tribe Lachneae

In Boudier's system this tribe comprises seven genera represented in


Britain b u t it m u s t be a d m i t t e d t h a t several of these are separated on
r a t h e r ill-defined grounds:

I. Apothecia subterranean, bursting through the soil surface at maturity,


becoming competely superficial in S. tenuis, clothed with undulating
brown hairs, scarcely differentiated from mycelial hyphae
Sepultaria (p. 31)
II. Apothecia superficial from the first, on wood, leaves or soil, hairs mostly
straight, stiff and tapering, clearly differentiated from mycelial
hyphae, at least towards the margin of the disc:
A. Hairs hyaline, receptacle white Leucoscypha (p. 32)
B. Hairs coloured:
1. Brown setae scattered through the hymenium, on pine needles
Desmazierella (p. 33)
2. No setae in the hymenium:
a. Fruit body cup-shaped:
*Spores without conspicuous oil drops
Tricharia (p. 34)
**Spores with two large oil drops Humaria (p. 35)
b. Fruit body flattened or cushion-like:
*Hairs long, stiff and pointed Trichophaea (p. 35)
**Hairs short, obtuse, thin-walled, in clusters
Pseudombrophila (p. 37)

30
SEPULTARIA (Cooke) L a m b o t t e
Sepultaria (Cooke) L a m b o t t e in Mem. Soc. roy. sciences Liege II, 14:
301 (1887).
Apothecia at first completely closed and subterranean, then splitting
irregularly in a star-shaped m a n n e r b u t remaining cup-shaped except
in S. tenuis, which becomes lenticular, outer surface completely clothed
with long, septate, undulating, brown hairs with obtuse tips. These are
scarcely differentiated from mycelial h y p h a e and m a y even be
branched. Spores large, elliptical or elliptic-fusiform, smooth, usually
with one or two large oil drops.

Sepultaria sumneriana (Cooke) Massee, British Fungus Flora 4 : 3 9 1


(1895). (PL VE.)
Apothecia up to 7 cm. across, developing as a hollow sphere just
below the soil surface, the upper portion t h e n protrudes and splits into
numerous lobes which bend back to expose the light yellowish hymeni-
u m ; flesh thick, white, brittle; outer surface light brown, densely
clothed with long dark brown, branched hairs, whose walls are usually
finely encrusted with colourless granules. Asci about 350 fx long; spores
elliptic-fusiform, 30-37 x 14-16 fx, usually with two large oil drops;
paraphyses slender, colourless, branched below.
On lawns, under Cedrus, January to May. Not uncommon. Figured
material from Kew, iii~195(5.
Sepultaria foliacea (Schaeffer ex Boudier) Boudier, Icones myco-
logicae II, Livraison 7 (March 1906), is a very similar b u t slightly
smaller fungus with paler h y m e n i u m and broadly elliptical spores,
25-28 x 15-18 jx, containing one or two large oil drops.

Not uncommon in heavy soil in woods.

Sepultaria arenosa (Fuckel) Boudier, Hist. Class. Discom. d ' E u r o p e :


59 (1907), sensu Seaver, N. Amer. Cup Fungi (Operculates): 151
(1928). (PL VF.)
Apothecia seldom more t h a n 1 cm. diameter, buried in sand, t h e n
splitting in a stellate m a n n e r with only the tips of the lobes p r o t r u d i n g
above the sand surface, h y m e n i u m cream-coloured, drying pale
ochraceous; outer surface light brown, clothed with undulating, ob-
tuse, m a t t e d hairs, about 6 [x thick, with smooth, hyaline to reddish-
brown walls (Fig. 5g). Asci up to 2 3 0 x 1 8 fx; spores elliptical, 2 0 - 2 5 x
10-14 jx with one or two large oil drops; paraphyses slender, slightly
clavate, about 6 fx thick at the tip.

31
Common in wet slacks of coastal dunes but also found in sandy soils in-
land, September to June. Figured from Hightown, Lancashire, 4. vi. 1952.
Sepultaria arenicola (Leveille) Massee, British Fungus Flora 4: 390
(1895), occurs in similar situations b u t m a y be distinguishable b y its
larger apothecia, with spores 2 3 - 2 8 x 1 4 - 1 6 JA.

Sepultaria tenuis (Fuckel) Boudier, Hist. Class. Discom. d ' E u r o p e : 59


(1907). (PL VG.)
Apothecia 1 - 1 . 5 cm. across, cup-shaped t h e n soon expanded and
finally convex, h y m e n i u m light grey; flesh white, splitting irregularly
into lobes at t h e margin; outer surface light brown, clothed with long
brown hairs, surface cells globose, brown-walled, 30 JA across, hairs
undulating, thick-walled, smooth, septate, 7 - 8 [A thick. Asci about
2 0 0 x 2 0 [A; spores broadly elliptical with one or two large oil drops and
often also several smaller ones, 2 0 - 2 4 x 1 1 - 1 3 [i; paraphyses straight,
somewhat clavate, up to 7 JA thick, containing colourless granules.

On heavy soils in woods and on humus under Salix, August. Uncommon.


Figured material from Wheatfen Broad, Norfolk, 17. viii. 1947.

LEUCOSCYPHA Boudier

Leueoscypha Boudier in Bull. Soc. mycol. France 1: 104 (1907).


Apothecia small, terrestrial, white, clothed with white septate hairs,
spores elliptical, with two oil drops. There is one British species, which
closely resembles, a p a r t from colour, some species of Neottietta in the
Ciliarieae. Le Gal in Bui. J a r d . Bot. Bruxelles 27: 719 (1957) has
united these two genera.

Leucoscypha leucotricha (Albertini & Schweinitz ex Fries) Boudier,


Hist. Class Discom. d ' E u r o p e : 57 (1907). (Fig. 6i.)
Apothecia up t o 5 m m . across, sessile, cup-shaped, white t h r o u g h o u t ,
outer surface completely concealed b y long, tapering, white hairs,
which curve over to cover t h e disc when dried. Hairs thin-walled at
t h e slightly swollen base, contracted to a long, tapering, smooth, very
thick-walled, septate limb with slightly rounded tip, the base some-
times curved b u t the limb usually stiff and straight, up to about
450x 10-15 [A at the b a s e , 2 - 4 JA at the tip (Fig. 5H). Flesh composed of
stout, hyaline, irregularly swollen, septate, interwoven hyphae, 8 - 2 0 [a

32
wide. Asci about 3 0 0 x 2 0 [x; spores elliptic-fusiform, 25-34x 12-16 [x,
containing two oil drops, covered with minute, closely spaced, hyaline
warts; paraphyses slightly clavate, 6 jx at t h e tip, hyaline.
On the ground in woods and on decaying leaves, July to September. Un-
common. Figured material from Norland Clough, Halifax, Yorkshire,
25. vii. 1897.
British material called Leucoscypha nivea (Romell) Boudier, op. cit.: 57,
is the above species, as is that called Leucoscypha fossulae (Limminghe ex
Cooke) Boudier, op. cit.: 57. The type of L. fossulae is a Sepultaria according
to Le Gal (op. cit.).

DESMAZIERELLA Libert

Desmazierella Libert in Ann. Sci. Nat. 17: 82 (1829).

Apothecia like those of a Trichophaea b u t distinguished by the


presence of dark brown setae in the h y m e n i u m ; paraphyses much
branched at the tip. There is one species:

Desmazierella acicola Libert, op. cit.: 8 3 (1829). (PI. VJ.)

Apothecia u p to 5 m m . across, sessile, saucer-shaped with a flat or


slightly convex disc, hymenium whitish or slightly glaucous; flesh
thick, white, formed of loosely woven h y p h a e 5 - 6 jx thick; outer sur-
face light brown, r a t h e r sparsely 'set with stiff, very dark brown,
bristle-like hairs, up to 1000 jx long b y 20 [x at the base, tapering to
about 5 tx at the tip, with minutely roughened walls, closely septate.
Asci about 400x 15 (x, spores elliptical, smooth, with a rounded hyaline
apiculus about 1.5 jx thick on each end, 1 5 - 1 7 x 8 - 1 0 tx, containing two
oil drops; paraphyses slender, much branched near the tip to form
small fascicles of brown, septate, finger-like bodies, 3 - 5 fx thick, with
minute granules on their walls, some protruding as bristles above the
general level of the hymenium. There are also slender simple p a r a -
physes, hyaline below b u t produced above into brown, rough-walled,
septate bristles which protrude above the general level of the disc and
are conspicuous under a hand-lens. The apothecia are surrounded b y
a small and inconspicuous subiculum of undulating, dark-brown
hyphae, 4 ;x wide, creeping over the surface of the needle.
On fallen needles of Pinus silvestris, F e b r u a r y to April.
Apothecia seem to be rare or, at least, seldom collected, but Gremmen
has shown that the fungus has a conidial state which Hughes has identified
with Verticicladium trifidum Preuss. This is common on fallen pine needles
throughout England and Wales and at all seasons. Hence the fungus is

33
actually extremely abundant though its apothecia are so seldom developed.
According to Berthet its true affinities are with the Urnulae in Sarco-
scyphaceae.

TRICHARIA Boudier

Tricharia Boudier in Bull. Soc. mycol. France 1: 104 (1885).


Apothecia always small,less t h a n 1 cm. across, sessile, saucer-shaped,
disc white, grey or grey-brown in British species, hairs stiff, brown,
septate, rather thin-walled; spores elliptical, without oil drops, smooth
or rough, paraphyses slender, straight, colourless or nearly so.

Tricharia gilva Boudier, Icones mycologicae, Livraison 2 (September


1904). (PI. VIA.)
Apothecia up to 5 m m . diameter, sessile, saucer-shaped, disc flat,
pale fawn; outer surface concolorous, covered with stiff, light reddish-
brown hairs which are smooth, thin-walled, sparingly septate, up to
2 0 0 x 1 0 [x, tapering to a rounded point. Asci up to 2 5 0 x 1 5 jx; spores
elliptical, 1 4 - 1 9 x 9 - 1 0 jx, smooth; paraphyses slender, scarcely clavate,
about 4 (x thick.
On burnt ground and cinder heaps, especially after they have become
overgrown with moss, April. Uncommon. Figured from a cinder heap,
Clouside, Staffordshire, iv. 1945, leg. W. D. Graddon.

Tricharia cretea (Cooke) Boudier, Hist. Class. Discom. d ' E u r o p e : 58


(1907). (PI. VIb.)
Apothecia about 5 m m . diameter, sessile, saucer-shaped, disc flat,
white, outer surface reddish-brown, covered with stiff red-brown hairs
which tend to occur in bunches towards the margin and give it a
dentate appearance. Individual hairs are up to 300 (x long, 12-14 jx
thick at the base, tapering to about 3 jx at the tip, with smooth, light
reddish-brown, walls, about 1 jx thick, and numerous septa (Fig. 5i);
the surface cells are obovate with t h i n reddish-brown walls. Asci up
to 250x10 jx; spores elliptical, 1 6 - 1 9 x 9 - 1 2 jx; paraphyses 3 - 4 [x thick,
scarcely enlarged at the tip.

On plaster, decaying leather and soil, October to June. Uncommon.


Figured material from the type collection, on the wet plaster ceiling of a
bedroom in the ruins of Inverleith House, Edinburgh, vi. 1877.

Tricharia fimbriata (Quelet) Boudier, op. cit.: 57 (1907) is probably


a synonym.

34
Lachnea lojkaeana Rehm in Rabenhorst's Kryptogamenflora Ed. 2 , 3
(1): 1045 (1895 belongs in Tricharia b u t has apparently not been
transferred because of uncertainty regarding its synonymy. It is
distinguished b y its deep yellow or orange disc and somewhat tapering
ascospores 1 4 - 1 8 x 8 - 1 0 jx, without guttules. T.praecox (Karsten)
Boudier op. cit. 57 (1907), on b u r n t ground in May, m a y be the corrcet
name for it.

HUMARIA Fuckel
Humaria Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24: 320
(1870.)
Apothecia cup-shaped, clothed with brown, septate, pointed hairs;
hymenium usually light coloured, spores elliptical, usually with two
oil drops and rough. A n u m b e r of species have been referred here b u t
only one is common or well known.

Humaria hemisphaerica (Wiggers ex Fries) Fuckel, op. cit. : 322


(1870). (PI. Vic.)
Apothecia up to 3 cm. across, sessile, permanently cup-shaped,
hymenium white or whitish; outer surface entirely covered with stiff,
smooth, dark brown, thick-walled, septate hairs, about 500 [x long
and 20 [x thick at the base, tapering evenly to a sharp point. Marginal
hairs m a y be 1 m m . long. Asci about 3 5 0 x 2 0 jx; spores broadly ellipti-
cal, with two conspicuous oil drops, coarsely warted, 20-24x 10-12 jx;
paraphyses clavate, 7 - 8 jx thick at t h e tip, colourless.

On soil in woods or, occasionally, on very rotten wood in wet places, July to
October. Common. Figured from soil and sticks, in Fen Carr, Wheatfen
Broad, Norfolk, 27. vii. 1946.

TRICHOPHAEA Boudier
Trichophaea Boudier in Bull. Soc. mycol. France 1: 105 (1885).
Receptacle thick and lenticular, disc white or pale grey, hairs long,
brown, pointed and septate, spores containing large oil drops, smooth
in most species.

Trichophaea woolhopeia (Cooke& Phillips) Boudier, Hist. Class. Discom.


d ' E u r o p e : 6 0 (1907). (PI. VIE.)
Apothecia up to 4 m m . diameter, sessile, disc light grey, drying
cream, flat or slightly concave; outer surface dark brown, clothed with
short dark brown hairs which often occur in clusters. Hairs up to

35
250x 20 /i, fusiform, distinctly bulbous at the base, wall 1 - 2 /« thick and
dark brown below, paling to a subhyaline tip, with from one to five
septa; surface cells of the cup pyriform, dark brown (Fig. 5E). Asci
220-300x15-18/«; spores broadly elliptical, 20-22x13-15/«, smooth,
containing one large oil d r o p ; paraphyses clavate, 7 - 8 n thick at the
tip.

On burnt ground and charcoal, September to October. Uncommon.


Figured material from the type collection, Downton, Herefordshire, 1877.

Trichophaea gregaria (Rehm) Boudier, Hist. Class. Discom. d ' E u -


r o p e : 6 0 (1907), differs in its distinctly fusiform spores, 2 0 - 3 0 x 9 - 1 1 /<
and in var intermedia Le Gal these are finely w a r t e d .

Trichophaea hemisphaerioidcs (Montagne) Graddon in Trans. Brit.


mycol. Soc. 4 3 : 6 8 9 ( 1 9 6 0 ) . (PI. V I D . )
Apothecia u p to 15 m m . diameter, usually smaller, sessile, saucer-
shaped, disc concave, white; outer surface light brown, clothed with
short, stiff, light brown hairs, up to 450 n long, 6-10/« wide below,
tapering to a sharp point, walls 1 - 2 /« thick, smooth, septa numerous
(Fig. 5D). Asci up to 170x10/«; spores elliptical, smooth, 1 3 - 1 6 x 7 - 8 /«,
with a small oil drop near each end; paraphyses slender, slightly cla-
v a t e and up to 6 /t thick at the tip, colourless.

On burnt ground amongst Funaria in woods, April to September. Un-


common. Figured material among i ^ r e a r i ^ L f e ^ T w n ^ ^ , Pass of Killie-
crankie, 27. viii. 1953.

Trichophaea abundans (Karsten) Boudier, op. cit. 6 1 ( 1 9 0 7 ) is a very


similar b u t smaller-fruited fungus growing on charred wood.

Trichophaea bicuspis (Boudier) Boudier, Icones mycologicae, Livraison


1 4 (July 1 9 0 7 ) . (PI. V I F . )

Apothecia up to 4 m m . diameter, disc white, flat, cup obconical,


seated on a small base, clothed with r a t h e r sparse, long, stiff, reddish-
brown hairs. Hairs up to 600/« long, 15/« thick, tapering evenly to a
sharp point, walls 4 - 5 /« thick, septa numerous, thin, base of the hair
often produced into a similar b u t downward pointing appendage, up
to 350/« long (Fig. 5B). Asci about 250x18/«; spores broadly elliptical,
smooth, 17-20x10-11/«, containing two large oil drops.

On decorticated sticks in swamps and on damp soil, September. Rare.


Figured material from ltoyston, near Barnsley, Yorkshire, 20. ix. 1897.

36
PSEUDOMBROPHILA Boudier

Pseudombrophila Boudier in Bull. Soc. mycol. France 1:106 (1885).


Apothecia lenticular with a short stalk, light coloured, outer surface
ornamented with clusters of short, coloured, obtuse, septate hairs; asci
small, spores smooth, without oil drops, paraphyses slender, not cla-
vate.

Pseudombrophila deerata (Karsten) Seaver, N. Amer. Gup Fungi (Oper-


culates): 1 4 1 ( 1 9 2 8 ) . ( P I . V I I F . )
Apothecia up to 10 m m . diameter, short-stalked, disc light grey with
pink or purplish tints, convex, margin often undulating, reflexed;
outer surface concolorous b u t dotted with the dark brown t u f t s of
downy hairs. Hairs obtuse, cylindrical, thin-walled, 1- or 2-septate,
8 0 - 1 2 0 x 4 - 5 y. Asci up to 150x14,«; spores elliptical, smooth, 12-16x
8 - 9 y , without oil drops; paraphyses slender, colourless, scarcely
thickened above.

On plant debris, including decaying bracken, flax, rotting stems of


Heracleum._ Cirsium. Lujmms and Brassisa and on rotting swedes, May to
November. Seldom recorded but no doubt easily overlooked. Figured
material from rotting sacking in a wood near Hebden Bridge, Yorkshire,
28. vi. 1897.

Tribe Ciliarieae
Six genera are conventionally recognised in this tribe b u t the
distinctions between Scutellinia, Cheilymenia and Neottiella are not
altogether satisfactory. Melastiza m a y have affinities with Aleuria in
the next tribe.
I. Ascospores globose:
A. Ascospores ornamented with warts or spines, disc red
. Scutellinia (p. 38)
B. Ascospores smooth, disc brown . . . . Sphaerosporella (p. 39)
II. Ascospores elliptical:
A. Hairs stiff and pointed, often thickwalled:
1. Hairs brown, with a forked base rooting in the flesh of the
receptacle, ascospores usually more or less rough-
walled and filled with oil drops . . . Scutellinia (p.38)
2. Hairs yellowish or hyaline, ascospores smooth in Cheily-
menia, ornamented in some species of Neottiella:
a. Ascospores without oil drops, disc often yellow
Cheilymenia (p. 40)
b. Ascospores containing oil drops, disc often red
Neottiella (p. 42)

37
B. Hairs obtuse, without forked rooting base, short and easily over-
looked :
1. Terrestrial species with coarsely ornamented ascospores
Melastiza (p. 43)
2. Species of burnt ground or charred wood, ascospores smooth
Anthracobia (p. 44)
If the outer surface of the cup bears only a few broad thin-walled, ob-
tuse, hyaline hairs or anchoring hyphae see Aleurieae.

SCUTELLINIA (Cooke) L a m b o t t e

Scutellinia (Cooke) L a m b o t t e in Mem. Soc. roy. sciences de Liege II,


14:299 (1887), emend Le Gal, Discom. de Madagascar: 116 (1954).
This genus has been generally called Ciliaria Quelet, a name which
has to be rejected as a later h o m o n y m of Ciliaria Haw. It has apo-
thecia often 1 cm. across, saucer-shaped with flat, reddish-brown,
brown or more often bright red disc; hairs brown, thick-walled, with
t h i n septa, with forked bases rooting in the flesh; spores spherical
to elliptical, often with m a n y oil drops and usually more or less
o r n a m e n t e d ; paraphyses clavate with the sap turned green with iodine
when the hymenium is red. The species with elliptical spores and
bright red disc have generally passed under the collective name
"Lachnea" scutellata. In fact t h e y include a highly critical assemblage
of t a x a , separable b y details of spore shape and ornament or b y the
nature of the hairs and it is still uncertain how m a n y good species
there are. T h a t described below as S. scutellata is probably the most
a b u n d a n t member of the series in England.
For a key to the european species with elliptical ascospores see Le
Gal in Bull. Soc. mycol. France 82:301-334 (1966). Those with spheri-
cal ascospores are often separated in the genus Sphaerospora (Sac-
cardo) Saccardo, Sylloge F u n g o r u m 8 : 1 8 8 (1889).

Scutellinia scutellata (Linnaeus ex St. Amans) Lambotte, op. cit. :299


(1887). (PI. VIG.)
Apothecia up to 10 m m . diameter, saucer-shaped, disc bright r e d ;
outer surface light brown, clothed with stiff dark-brown or blackish
hairs. Hairs up to 1000 /t long with forked bases rooting in the flesh,
40fi thick below, tapering to a point, wall 4 - 1 0 / t thick, septa t h i n ;
interspersed with these are short, obtuse, 1- to many-celled hairs.
Asci up to 3 0 0 x 2 5 / / ; spores elliptical, 1 8 - 1 9 x 1 0 - 1 2 ^ , packed with
small oil drops, minutely and irregularly roughened; paraphyses cla-
vate, up to 10 fi thick at t h e tip, sap green in iodine.

38
On wet ground or on sodden wood, May to November. Common. Figured
material from Widnes, Lancashire, 1. xi. 1954.
Scutellinia stenosperma Le Gal, Discom. de Madagascar: 146 (1954),
is one of the more distinct related species, with narrowly elliptical
warted spores, 2 0 - 2 5 x 9 - 1 1 . 5 p (Fig. 5A.)

Scutellinia trechispora (Berkeley & Broome) L a m b o t t e , op. cit.:299


(1887). (PI. VIH.)
Apothecia 5-10 m m . diameter, cup-shaped t h e n flattened, disc flat,
bright red; outer surface pale reddish-brown, clothed with stiff brown
hairs. Hairs up to 1000x45/«, with brown walls 6-7/« thick, smooth,
forked or simple at the base, limb slightly fusiform below, tapering to
a sharp point, with up to seven thin septa. Interspersed with these are
other short 1-septate hairs and elongated, brown, obtuse cells up to
75x16 /«, some thin-walled, others thick-walled. Asci up to 4 0 0 x 3 5 /«;
spores globose, ornamented with low round bosses, 18-22/« over all;
paraphyses clavate, up to 10/« thick, contents green with iodine.

On damp soil, often among mosses, all the year. Common. Figured
material, the upper spore, from Midford, Somerset, 26. iii. 1950.
Boudier recognised a variety paludicola with larger spores, 22-26ju.
Scutellinia asperior (Nylander) Dennis in Kew Bulletin 1955:571
(1956), occurs in similar situations b u t differs in its spores being orna-
mented by spines in place of the low w a r t s of S. trechispora. See
PI. VIH, t h e lower spore.

SPHAEROSPORELLA Svrcek & Kubicka

Sphaerosporella (Svrcek) Svrcek & Kubicka in Geska Mykologie 15:66


(1961).
The genus contains a few species separated from Scutellinia sub-
genus Sphaerospora b y their p e r m a n e n t l y smoothwalled ascospores,
brownish or yellowish discs and nonrooting hairs.

Sphaerosporella brunnea (Albertini & Schweinitz ex Fries) Svrcek &


Kubicka, op. cit. 65 (1961). (PI. VIi.)
Apothecia up to 5 m m . diameter, saucer-shaped, disc flat, reddish-
brown; outer surface appearing almost smooth under a lens b u t
marked b y lighter or darker vertical striae formed b y the adpressed
downy hairs. Marginal hairs very closely set, minute, almost hyaline,

39
up to 5 0 x 1 0 n , somewhat fusiform and tapering to a sharp point, wall
about 1 n thick, 1- or 2-septate; hairs on the sides of the cup dark
brown, cylindrical, obtuse, thin-walled, 1- or 2-septate, up to 100x12 /«
more or less fasciculate and closely adpressed to the surface. Asci
about 200x16/«; spores globose, smooth, 13-15 /«; paraphyses clavate,
6 ft thick at the tip, contents reddish-brown.

On burnt ground and amongst mosses, August to September. Uncommon.


Figured material from a charcoal heap, Birnam, Perthshire, 12. viii. 1954.

Sphaerosporella hinnulea (Berkeley & Broome) Rifai differs in grow-


ing on wet ground.

CHEILYMENIA Boudier

Cheilymenia Boudier in Bull. Soc. mycol. France 1 : 1 0 5 (1885).


Apothecia small, usually some shade of yellow or orange, light red
in C. rubra, hairs hyaline or yellowish, occasionally light brown, spores
elliptical without oil drops, usually smooth, paraphyses slender. The
hairs are often lobed at the base and in a few species some of t h e m
assume characteristic forked or stellate shapes; t h e y are often in-
conspicuous and easily overlooked except as a fimbriate margin to
the disc. Several common species occur on dung or on rotting p l a n t
debris, a few on bare soil.

Cheilymenia fibrillosa (Currey) Le Gal Recherches—ornamentation


sporales des Discomycetes opercules 287. These, Paris (1947).
Apothecia large for the genus, 1 - 2 cm. diameter, cupulate, disc
bright orange-yellow, receptacle paler, white-fibrillose. Asci 2 0 0 x 1 1 /i,
ascospores elliptical, smooth, 15-16x8-9/«, without guttules; para-
physes slender, straight, suddenly enlarged to subglobose tips 5-8/«
diameter, with yellow oily contents. Hairs of two kinds, relatively thin-
walled, closely septate, obtuse, brown hairs 100-160x10-20/« largely
concealed by adpressed, m a t t e d , fibrils of much more slender hyaline
hyphae (Fig.31j)
On soil. August to October. Rare.
C. cornubiensis (Berkeley and Broome) Le Gal is similar b u t with
ascospores 17-22x11-12/«, bearing a few scattered granules on their
outer walls.

40
Cheilymenia vitellina (Persoon ex Fries) Dennis E d . 1:27(1960)
Peziza vitellina Persoon, Mycologia europaea 1 : 2 5 7 (1822). (PI. VIK.)
Apothecia up to 10 m m . diameter, sulphur-yellow t h r o u g h o u t , disc
concave; outer surface concolorous, clothed with short yellowish hairs.
Hairs up to 1000x40/«, lobed at the base, walls smooth, 1-3/« thick,
tapering to a rounded tip, septa thin. Asci 250x14/«; spores 14-17x
6.5-9/«, smooth, without oil drops; paraphyses slightly enlarged to
4-6 n at the tip.
On soil, July to November. Not uncommon. Figured from bare soil beside
a stream, Cookham Moor, Buckinghamshire, 14. x. 1951.

Peziza dalmeniensis Cooke in Grevillea 3 : 6 6 (December 1874), is a


synonym.

Cheilymenia crucipila (Cooke & Phillips) Le Gal, Discom. de Madagas-


car: 1 1 1 ( 1 9 5 4 ) . ( P L V I J . ) Peziza crucipila Cke. & Phil., Mycographia:
136 (1876).
Apothecia up to 6 m m . diameter, cup-shaped, t h e n expanded, disc
flat, bright orange to scarlet; outer surface m u c h paler, sparsely
clothed with minute pale brown hairs. Hairs simple, forked from t h e
base or, less often, 3-5-rayed with arms up to 280 x 10 /«, slightly taper-
ing to a narrowly rounded or pointed tip, wall up to 3 /«thick, yellow-
ish-brown, paler at the tip, lumen up to 5-septate (Fig. 5c). Asci up
to 220x17/«; spores elliptical, 15-18 (-20) x 8.5-10 (-12)/«, smooth,
without oil drops; paraphyses slightly clavate, up to 7/« at the tip,
filled with light yellow granules t u r n e d grey with iodine.

On damp ground amongst small mosses, May to September. Not common.


Figured from Tumby, Lincolnshire, 18. vii. 1908.

C. calvescens Boudier, Icones mycologicae, Livraison II (December


1906), is a synonym.

Cheilymenia coprinaria (Cooke) Boudier, Hist. Class. Discom. d ' E u -


rope:63 (1907). (PL VIN.)
Apothecia 3 - 4 m m . diameter, saucer-shaped, disc flat, reddish-
orange; outer surface paler, sparsely clothed with reddish-brown hairs.
Hairs up to 560x30/«, walls 3/« thick, smooth, up to 10-septate, base
often lobed. Asci up to 200x24/«; spores elliptical, smooth, 16-23x
9-13/«, without oil drops b u t with a mucilaginous coating; paraphyses
often enlarged to 7 /« at the tip.

41
On cow dung, less often on horse dung and rabbit droppings, throughout
the year but especially in spring and autumn. Common. Figured from cow
dung near Halifax, Yorkshire, 29. ix. 1896.
Cheilymenia stercorea (Persoon) Boudier, op. cit.:63 (1907), is very
similar and also common on cow dung in spring b u t differs in having
stellate 3-5-armed hairs on the lower p a r t of the cup.

Cheilymenia theleboloides (Albertini & Schweinitz ex Fries) Boudier,


Icones mycologicae, Livraison 2 (September 1904). (PL VIM.)
Apothecia up to 6 m m . diameter, saucer-shaped, disc flat, egg-
yellow; outer surface paler, clothed with inconspicuous, scattered hya-
line to yellowish-brown pointed hairs up to 2 2 0 x 2 0 /i, the wall 1 /.<
thick, 1-3-septate (Fig. 5J). Asci up to 2 5 0 x l 4 / < ; spores elliptical,
12-18x7-10/», smooth, without oil drops; paraphyses slender, slightly
clavate, yellowish at t h e tip.

On spent hops, less often on leaf-mould or lime waste, April to May and
October to November. Rather common. Figured on spent hops, Skircoat,
Halifax, Yorkshire, 15. xi. 1897.
Cheilymenia rubra (Cooke) Boudier, Hist. Class. Discom. d ' E u r o p e :
63 (1907), closely resembles C. theleboloides and also occurs on spent
hops b u t has a light red disc.

NEOTTIELLA (Cooke) Saccardo

Neottiella (Cooke) Saccardo, Sylloge F u n g o r u m 8:190 (1889), emend


Boudier, Hist. Class. Discom. d ' E u r o p e : 6 4 (1907).
Apothecia small, sessile, disc red or reddish-orange; outer surface
pale and clothed with inconspicuous, slender, hyaline hairs; spores
smooth or rough, with oil drops. All the species are closely associated
with Bryophyta. T h e y fall into two series, one with rough spores and
showing affinity with Aleuria, the other with smooth spores and hav-
ing affinity with Cheilymenia on the one hand and Octospora on the
other. The former occur especially with Polytrichum, the latter with
other Bryophyta.

Neottiella rutilans (Fries) Dennis E d . 1:28 (1960)


Peziza rutilans Fries, Systema mycologicum 2 : 6 8 (1822). (PL VIw.)
Apothecia sessile or with a short stalk often buried in the soil, up
to 1.5 cm. diameter, disc concave, reddish-orange; outer surface cup-

42
shaped, whitish, clothed with thin-walled, septate, undulating and
entangled, downy, hyaline hairs, which t a p e r to a rounded tip. Asci
up to 300x20/«; spores broadly elliptical, 22-25x13-15/«, containing
one or two large oil drops, ornamented with a more or less continuous
colourless reticulum; paraphyses slightly clavate, with orange con-
tents t h a t t u r n green in iodine.

On sandy soils amongst Polytrichum, October to January. Common.


Figured from Formby, Lancashire, upper spore only.
N. rutilans has often been called Peziza polytrichi Schumacher ex Fries,
Systema mycologicum 2:70 (1822), perhaps correctly, but Madame Le Gal,
following a tradition founded by Nyfander in 1865, prefers to interpret that
as a Lamprospora with finely reticulated spores.

Neottiella vivida (Nylander) Dennis E d . 1:28. ( = Peziza vivida


Nylander in Flora 48:467 [1865]), is a very similar fungus, distinguish-
able only by its spores being warted instead of reticulated, PL VIw,
the lower spore, from the Isle of Yell, Zetland, 1. ix. 1952.

Neottiella hetieri Boudier in Bull. Soc. mycol. France 12:12 (1896).


(Pl. V I R . )
Apothecia up to 2 m m . diameter, cup-shaped, sessile, disc flat, bright
orange; outer surface white, clothed with delicate white hairs which
also form a fine marginal fringe. Hairs fusiform below, tapering to a
point, up to 200x 15 /<, wall smooth, 1-1.5 /< thick, lumen 0-3-septate.
Asci up to 200x17 /«; spores elliptical, smooth, 13-17x8-9/«, contain-
ing one large oil drop; paraphyses slightly clavate, up to 7 /« thick at
the tip, with orange contents.
Amongst Funaria on burnt soil and charcoal, spring and autumn. Com-
mon. Figured from burnt ground at the head of Gfen More, Aviemore,
Inverness-shire, 14. iv. 1949. The hairs are easily overlooked and the fungus
is then often misdetermined as Octospora leucoloma.

MELASTIZA Boudier

Melastiza Boudier in Bull. Soc. mycol. France 1:106 (1885).


Apothecia sessile, terrestrial, disc red or orange; outer surface
bearing clusters of short, brown, obtuse, septate hairs; spores elliptical,
with oil drops, coarsely ornamented, paraphyses clavate, their con-
tents green in iodine. The hairs are easily overlooked as to the unaided
eye t h e y merely give a brown pruinose appearance to the upper p a r t
of the cup.

43
Melastiza chateri (W. G. Smith) Boudier, Hist. Class. Discom. d ' E u -
rope: 6 4 ( 1 9 0 7 ) . P I . V I L . )

Apothecia up to 1.5 cm. diameter, exceptionally even larger, cup-


shaped then expanded, disc bright orange to vermilion; outer surface
paler, dotted towards the margin b y minute bunches of downy brown
obtuse hairs, 6 0 - 1 5 0 x 1 2 - 1 8 ^ with one to four septa and walls
thick (Fig. 5F). Asci up to 3 0 0 x 1 5 / , ; spores elliptical but ornamented
with a coarse raised reticulum which.often has a spine-like projection
at each end of the spore, 1 7 - 1 9 x 9 - 1 1 paraphyses clavate, up to 1 0 „
thick at the tip, which is filled with orange granules.

T i ^ f 8 0 from
c o m m c tlugured
common,
118
^ 0 1 5 6 1 ;Lancashire,
^ Widnes, 1 0 March
" L1.
° Cxi.
al, abundant
y1954. but
scarcely

Mdmtiza miniata (Fuckel) Boudier, Icones mycologicae, Livraison 5

Q i1SManSr°nym'
i 4 y - t o 4 (1958).
S E E ALS0 L E G A L IN B U L SOC
- M
Y C O L
France 74:

Melastiza scotica Graddon in Trans. Brit, mycol. Soc 44-609 (1961)


9Q a o« U P ro r f f Similar Speci6S w i t h
very coarsely warted ascospores
23-28x 12.5-14 fi.

ANTHRACOBIA Boudier

Anthracobia Boudier, in Bull. Soc. mycol. France 1:106 (1885).

Apothecia like those of Melastiza b u t always on b u r n t ground or


charred wood, disc less red, often yellowish to brown, in one species
nearly black, spores elliptical, with oil drops, permanently smooth
paraphyses clavate. The hairs are very short and easily overlooked'
In doubtful cases the marginal p a r t of the cup should be carefully
examined under a binocular microscope or a portion detached and
mounted for low power examination in water. The "hairs" of A macro-
cyshs are so r u d i m e n t a r y t h a t the fungus will usually be s o u g h t f i r s t
among Octospora b u t it is obviously a near relative of A. melaloma
Species of Anthracobia are also commonly determined as Pyronema
omphalod.es, because of their occurrence on b u r n t ground, b u t there is
little excuse for this. T h e y are gregarious b u t do not form the packed
more or less confluent, crusts of t h a t species, t h e y are less pink and
lack its conspicuous white cobwebby subiculum.

44
Anthracobia mclaloma (Albertini & Schweinitz ex Fries) Boudier, Hist.
Class. Discom. d ' E u r o p e : 6 5 ( 1 9 0 7 ) . ( P L V I P . )
Apothecia 2 - 5 m m . diameter, seldom larger, cujj-shaped, sessile, disc
concave then flattened, ochraceous;orangey outer surface slightly
paler, dotted brown with the bunches of downy hairs, particularly
towards the margin and hence often appearing striate when dried.
Hairs up to 110x10/«, cylindrical, obtuse, septate, with thin, smooth,
light brown walls (Fig. 5K). Asci up to 200x15/«; spores oblong-
ellipsoid, smooth, 14-22x7-11/«, with two oil drops; paraphyses
slender, apex clavate and up to 6 /«thick.

On burnt wood and on the sites of bonfires, July to January. Very com-
mon. Figured from soil where brushwood had been burned, Hackhurst
Downs, Surrey, 14. vii. 1946.

Anthracobia maurilabra (Cooke) Boudier, Hist. Class. Discom. d ' E u -


r o p e : 6 5 (1907). (PI. VIQ.)

Apothecia 3 - 6 m m . diameter, sessile, disc fawn, concave t h e n flat or


slightly convex; outer surface concolorous b u t marked towards t h e
margin by vertical bars and dots formed b y the bunches of adpressed
downy hairs. Hairs 50-130x7-20/«, thin-walled, dark brown, septate,
obtuse. Asci up to 200x15/«; spores oblong-ellipsoid, smooth, 19-22x
8-9/«, with two oil drops; paraphyses slender, only gradually enlarged
upwards and not markedly clavate, up to 5/« thick at the tip.

On burnt ground, June to September. Not uncommon but rather in-


conspicuous. Figured from Norbury Park, Mickleham, Surrey, 20. vi. 1948.

Anthracobia macrocystis (Cooke) Boudier, Hist. Class. Discom. d ' E u -


rope:65 (1907). (PL VIo.)

Apothecia 2 - 3 m m . diameter, sessile, disc concave t h e n flattened,


reddish-orange; outer surface paler, dotted and barred with brown to-
wards the margin where the surface cells are subglobose with t h i n
brown walls and bear short, broad, thin-walled, 1-2-celled hairs, a few
up to 70x23(x, the m a j o r i t y only 1-celled and about 25xl5|j. (Fig. 5l).
Asci up to 200X15|A; spores oblong-ellipsoid, smooth, 1 6 - 1 8 x 7 - 8 ^ ,
with two oil drops; paraphyses slender, clavate, enlarged to l\x at the
apex.

On burnt ground. July to October. Less common than A. melaloma.


Figured from Ilighgate, London, October 1878.

45
Anthracobia nitida Boudier, op. cit.:65 (1907), is probably a syno-
nym.

Anthracobia humillima Malencon in Bull. Soc. mycol. France 4 3 : 9 8


(1927). (Fig. 6G.)
Apothecia up to 2 m m . across, cup-shaped, sessile, disc concave t h e n
flattened, dark grey-brown to almost black, fringed with paler downy
hairs; outer surface dark grey with short dark hairs. Hairs along the
margin closely set and almost colourless, thin-walled, septate, up to
80 X12[A, tapering to a point, those of the flanks of the cup similar b u t
dark brown and up to 150X7(A. Asci about 125X11(A; spores elliptical,
11-15 x6.5—7[x, with two oil drops; paraphyses slender, clavate, about
5jx thick at the tip.

On^burnLgmund, June. Apparently uncommon but so difficult to see


that it is no doubt usually overlooked. Figured from burnt ground in pine
wood, Woburn Sands, Bedfordshire, 18. vi. 1954.

Tribe Aleurieae
This is a residual assemblage of genera, defined largely b y negative
characters, the absence of a blue ascus tip in iodine, the absence of
clearly differentiated hairs and the absence of broad asci protruding
conspicuously above the hymenial level at m a t u r i t y . A few species
might be sought among the Pseudoascoboleae, which is a similar
residual assemblage in the next family.

I. Spores elliptical:
A. Apothecia bright orange or yellow, spores conspicuously warted or
reticulated Aleuria (p. 47)
B. Apothecia smooth-spored or, if the spores are finely warted, then
very small and pulvinate:
1. Apothecia medium size, cup-shaped, stalked, spores smooth,
without oil drops Geopyxis (p. 48)
2. Apothecia small, lenticular or pulvinate:
a. Spores without oil drops, on dung or manured soil:
*Receptacle granulose with large spherical cells,
without adpressed hyphae, paraphyses swollen at
the tip Coprobia (p. 49)
**Receptacle often downy-fibrillose with slender ad-
pressed hyphal tips, paraphyses not enlarged at
the tip Fimaria (p. 51)

46
b. Spores contain oil drops:
*Disc orange or yellow, on soil or on living mosses
Octospora (p. 49)
**Disc brown, on wood Psilopezia (p. 55)
II. Spores globose:
A. Spores ultimately brownish, coarsely reticulated, disc dark brown
Boudiera (p. 52)
B. Spores permanently hyaline, disc orange or reddish:
1. Apothecia large, cupulate, orange, staining greenish
Caloscypha
2. Apothecia small, pulvinate or discoid, not staining greenish
when bruised:
a. Paraphyses simple, clavate . . . . Lamprospora (p. 53)
b. Paraphyses much forked, slender and curved at the tip
Pulvinula (p. 54)

Caloscypha fulgens (Persoon) Boudier, common in coniferous woods in


central Europe, with smooth globose ascospores 5-6 n, is not known from
the British Isles.

ALEUKIA Fuckel

Aleuria Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:325 (1870).


Apothecia terrestrial, often large, cup-shaped, thin-fleshed and
brittle, hymenium bright yellow or orange; outer surface smooth or
minutely downy under a lens, with colourless anchoring h y p h a e ; spores
elliptical, with two oil drops, ornamented with a conspicuous reticu-
lum or warted, paraphyses clavate, their contents turned green by
iodine. There are two British species.

Aleuria aurantia (Fries) Fuckel, op. cit.:325 (1870). (PI. VIIA.)

Apothecia gregarious, sessile, cup-shaped, t h e n undulating and


often flattened, up to 10 cm. diameter, sometimes irregular from m u t u a l
pressure and sometimes split so as to simulate an Otidea, h y m e n i u m
bright orange, flesh thin, fragile, white; outer surface minutely downy,
whitish. Asci up to 2 2 0 x 1 3 ^ ; spores elliptical b u t ornamented with
a coarse reticulum which often projects like an apiculus at each end,
1 7 - 2 4 x 9 - l l | j . , containing two oil drops; paraphyses slightly clavate,
up to 7[x thick at the tip which is filled with orange granules.

On bare soil in woods and open places, also among grass of lawns and road
verges, September to January. Very common. Figured from Auchincruive,

47
Ayrshire, 25. x. 1934. In spite of its large size, abundance and great beauty
it does not appear that this species has any traditional English name. The
term "Orange peel Peziza", however, though no doubt originally a book
name, appears to have become current of recent years and is undoubtedly
apt though somewhat prosaic.

Aleuria rhenana Fuckel, op. cit.:325 (1870) with similar b u t cluster-


ed stalked apothecia is a rare species of beech woods, doubtfully
British.

Aleuria lutconitens (Berkeley & Broome) Gillet, Champignons d<


France, Les Discomycetes:205 ( 1 8 7 9 ) . ( P I . V I I B . )
Apothecia 1 - 2 cm. diameter, sessile, saucer-shaped, disc brigh
golden-yellow, flesh t h i n ; outer surface smooth, pale. Asci up to
150xl0jjt; spores oblong-ellipsoid, 1 0 - 1 5 x 6 - 8 ^ ornamented at m a t u -
rity with conspicuous pointed warts, containing two large oil drops;
paraphyses slender, not clavate b u t slightly enlarged up to by. thick
at the tip which m a y be slightly curved, contents yellow.

On the ground, September to January. Uncommon. Figured from Berke-


ley's sketch of the type collection, Apethorpe, Northamptonshire.

GEOPYXIS (Persoon ex Fries) Saccardo

Geopyxis (Persoon ex Fries) Saccardo, Sylloge F u n g o r u m 8 : 6 3 (1889),


emend Boudier, Hist. Class. Discom. d ' E u r o p e : 5 3 (1907).
Apothecia terrestrial, of medium size, cup-shaped, sessile or more
often stalked, glabrous or nearly so; spores elliptical,smooth,without
oil drops, paraphyses slender. The genus was originally founded for a
miscellaneous assemblage of stalked terrestrial species but it was re-
defined by Boudier to include only those akin to Peziza carbonaria
Albertini & Schweinitz. Species of Pustularia might be sought here b u t
should be easily separated by the large oil drops in their spores. Peziza
ammophila is often placed in Geopyxis b u t should be excluded because
of the strong blue reaction of its ascus tip with iodine.

Geopyxis carbonaria (Albertini & Schweinitz ex Fries) Saccardo,


Sylloge F u n g o r u m 8 : 7 1 (1889). (PL V I I E . )
Apothecia cup-shaped, seated on a slender whitish stalk, disc con-
cave, up to 1.5 cm. diameter, light red, margin minutely crenulate;
outer surface whitish to flesh colour, minutely scurfy. Asci up to

48
200x10^; spores narrowly elliptical, r a t h e r pointed at each end,
smooth, 1 3 - 1 6 x 6 - 8 n , without oil drops; paraphyses slender, slightly
clavate.

On burnt ground and attached to charred wood, September. Uncommon.


In the figure the upper cup is on charred sticks, Scarborough, Yorkshire,
29. ix. 1948, the lower from burnt ground, Virginia Water, Surrey, 10. x. 1954.

COPROBIA Boudier

Coprobia Boudier in Bull. Soc. mycol. France 1 : 1 0 7 (1885).


Apothecia small, saucer-shaped, sessile, externally scurfy, spores
elliptical, smooth, without oil drops, growing on dung. There is one
species:

Coprobia granulata (Bulliard ex Fries) Boudier, Hist. Glass. Discom.


d ' E u r o p e : 6 9 (1907). (PI. Vis.)
Apothecia 1 - 3 m m . diameter, saucer-shaped on a broad base, disc
slightly concave, deep ochraceous-orange; outer surface slightly paler,
appearing coarsely granular owing to the presence of protruding, thin-
walled, subglobose cells, IOOJJ. or more in diameter; flesh thick, soft,
pale yellow, formed of elliptical or globose cells throughout. Asci up
to 190xl5(i, spores smooth, elliptical, 15-18X7-8IA, without oil drops;
paraphyses slender, apex strongly clavate and up to 14[x thick, filled
with orange granules. The tips of the ripe asci protrude somewhat
above the general level of the hymenium and t h e species has, on t h a t
account, sometimes been referred to Ascophanus.

On cow dung, June to September. Very common. Figured from cow dung,
Halifax, Yorkshire, 22. viii. 1898. Care must be taken not to confuse this in
the field with the species of Cheilymenia also common on cow dung.

OCTOSPORA Hedwig ex S. F. Gray

Octospora Hedwig ex S. F. Gray, Natural Arrangement British Plants


1:666 (1821), emend Korf. Mycologia 4 6 : 8 3 8 (1954).
Apothecia terrestrial or growing on mosses, rarely on bark, usually
much less t h a n 1 cm. diameter, disc orange or red, sessile, externally
nominally smooth b u t often with a prominent sterile dentate margin
and minutely downy under a lens; spores elliptical to fusiform, con-
taining oil drops, in most species smooth, in a few finely warted, con-

49
t e n t s of the paraphyses commonly green or blue-green with iodine,
their tips often strongly curved.

Octospora humosa (Fries) Dennis E d . 1:33 (1960)

Peziza humosa Fries, Systema mycologicum 2 : 7 1 (1822). (PI. V l x . )


Apothecia 5-10 m m . diameter, sessile, disc bright orange, flat or con-
vex with undulating margin; outer surface cup-shaped, pale ochra-
ceous, downy below. Asci up to 250x20/«; spores uniseriate, elliptic-
cylindric with rounded ends, 19-22x11-12/«, containing one large oil
drop and usually several small ones; paraphyses curved at t h e tip,
somewhat clavate and up to 9 /«thick.

On sandy soil amongst Polytrichum and other mosses, especially in au-


tumn and winter. Not common. Figured amongst Polytrichum, Shefford,
Bedfordshire, 15. x. 1950.

Octospora carbonigena (Berkeley) Dennis E d . 1 : 3 3 (1960)

Peziza carbonigena Berkeley, Flora Tasmaniae 2:274 (1860). (PI. Vlv.)


Apothecia 1 - 3 m m . diameter, densely gregarious, sessile, marginate,
cylindrical or t u r b i n a t e below, disc concave or flat, bright yellowish-
orange; outer surface paler, downy below. Asci up to 220x15/«; spores
uniseriate, elliptic-fusiform, 2 0 - 3 0 x 8 - 9 (-10)/«; paraphyses curved at
the tip, clavate, up to 5/« thick.
On plant debris on moors and wet ground, throughout the year. Uncom-
mon. Figured from peaty soil near Hebden Bridge, Yorkshire, 13. xii. 1902.

Peziza roumegueri Karsten in Grevillea 7 : 6 4 (December 1878), its


variety carnosissima Phillips in Grevillea 10:67 (December 1881), and
possibly also Humaria fusispora Rabenhorst appear to be synonyms.

Octospora tetraspora (Fuckel) Korf in Mycologia 46:838 (1954), is


a similar species b u t w i t h only four or six spores in each ascus, spores
with two oil drops, 20-30x10-12/«.

Octospora deformis (Karsten) Gamundi in Lilloa 30:300 (1960),


found occasionally in swarms on bare soil, has bright orange apothecia
with small elliptical ascospores, 15-16x8-9/«, usually filled with
granular cytoplasm rather t h a n with a distinct guttule.

The rather common fungus figured b y Boudier in Icones mycologi-


cae t a b . 397 as uHumaria rubricosa (Fuckel) Cooke" is a small orange
species of Octospora with strongly curved paraphyses and ascospores

50
18-25 x 9 - l l /«, apparently confined to cushions of Grimmia on old
walls at the end of December and early J a n u a r y . U n f o r t u n a t e l y there
seems little reason for identifying it with Peziza rubricosa Fries,
Systema mycologicum 2 : 7 2 (1822), or even with Leucoloma rubricosa
(Fries) Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:318 (1870)
and it will probably require a new name when a critical revision of
Octospora is undertaken.

Octospora leucoloma Hedwig ex S. F . Gray, Nat. Arrangement. Brit.


Plants 1:667 (1821). (PI. V I I I Q . )
Apothecia 1 - 3 m m . diameter, saucer-shaped, sessile, marginate, disc
orange; outer surface pale ochraceous with a denticulate margin and
downy with hyaline anchoring hyphae below. Asci up to 250x20/«;
spores uniseriate, elliptic-fusiform or somewhat ovoid and pointed at
one end, 20-22x10-12/«, with one or two large oil drops; paraphyses
straight or slightly curved, clavate, 5-6/« thick at the tip. Spores with
one and two oil drops occur in the same ascus.
On Bryum argenteum and other small mosses, October to May. Fairly
common. Figured from Kimbolton, Huntingdonshire, 15. iii. 1953.

Octospora carneola (Sauter) Dennis E d . 1 : 3 4 (1960)


Peziza carneola Sauter in Hedwigia 20:130 (1881). (PI. VIIIP.)

Apothecia up to 1 m m . diameter, sessile, saucer-shaped, disc light


orange, flat or convex; outer surface paler with a denticulate margin.
Asci up to 160x12/«; spores uniseriate, at first eight per ascus b u t the
lower four degenerate, upper spores broadly elliptical, 15-19 (-20) x
8-10/«, minutely b u t regularly warted, containing one or occasionally
two large oil drops; paraphyses curved at the tip, 4/« thick.
In cushions of Tetraplodon mnioides, July to August. Uncommon. Figured
from Quinag, Sutherland, vii. 1899. This is an alpine species characteristic
of high altitudes in the Highlands and is mentioned here to illustrate the
small section of Octospora with ornamented spores.

Peziza wrightii Berkeley & Broome in Ann. Mag. nat. Hist. I l l , 15:
444 (1865), is a rare species with subglobose warted spores, 13-15x
11-13/«, found on Amblystegia serpens on bark of trees in Wales and
the West of England.

FIMARIA Velenovsky
Fimaria Velenovsky, Monogr. Discom. Bohem^331 (1934).
Small apothecia, on dung or manured soil, sessile, cupshaped, often
with a toothed margin, receptacle parenchymatous, usually some

51
shade of brown or yellow-brown. Asci iodine-negative, ascospores
elliptical, without oil drops, smooth, contents often yellowish; para-
physes slender, simple, often embedded in brownish m a t t e r at the tip.

Pimaria hepatica (Batsch ex Persoon) v a n Brummelin in Persoonia 2:


322 (1962). (PL V I I I i ^
Apothecia lenticular, up to 4 m m . across, disc flat, pale reddish-
brown, p u n c t a t e with the asci, margin denticulate; outer surface
smooth or downy with colourless hyphae, purplish-brown. Asci up to
2 3 0 x 2 5 a s c o s p o r e s elliptical, smooth, uniseriate, 20-25(-38)x 1 0 -
13//, contents yellowish at m a t u r i t y ; paraphyses simple or branched,
enlarged up to 8// wide at t h e tip which is covered by amorphous
vinaceous pigment.
On dropping of rabbits and mice and on surrounding soil. Autumn and
winter. Not uncommon.
Ascobolus vinosus Berkeley, English Flora 5 (2): 209 (1836) is a
synonym and the figure is copied from Berkeley's own drawing of
t h a t species.

Fimaria theioleuca (Rolland) van Brummelin in Persoonia 2:329


(1962). (Fig. 29II.)
Apothecia small, yellowish-brown, turbinate, minutely downy, disc
pale yellowish. Asci 155-180x13-15/«, ascospores 13-15.5x 7.5-8.5 ft,
ellipsoidal, finally with yellowish contents; paraphyses filiform, up to
2/t wide, not enlarged at the tip and not united b y external pigment.
On dung of rabbit, deer and sheep. Winter and Spring. Not uncommon.

Fimaria cervina (Phillips) v a n Brummelin has a chestnut disc and


ascospores 1 4 - 1 7 x 7 - 8 . 5 ,«.

F. lcporum (Albertini & Schweinitz ex Persoon) Velenovsky has a


yellowish disc and ascospores 1 3 - 1 6 x 7 . 7 - 8 . 3 f t .

Fimaria porcina S v r c e k & Kubicka in Ceska Mykologie 19:214 (1965)


is distinguished b y its small ascospores 10-13x7-9/«.

B O U D I E R A Cooke
Boudiera Cooke in Grevillea 6 : 7 6 (December 1877).
Apothecia terrestrial, sessile, pulvinate, asci large, spores globose,
ornamented with a deep reticulum, ultimately t u r n i n g brown. The

52
genus is sometimes referred to the Ascobolaceae. There is only one
British species:

Boudiera areolata Cooke & Phillips, op. cit.:76 (December 1877).


(PI. VIIIK.)
Apothecia up to 3 m m . diameter, sessile pulvinate, disc convex,
purplish-brown, flesh t h i c k ; outer surface smooth, paler. Asci 8-spored,
up to 4 5 0 x 6 0 / t ; spores globose, ornamented with a deep large-meshed
reticulum, at first hyaline, contents becoming brown at m a t u r i t y ,
30-40/, diameter, including the reticulum, 2 5 - 3 0 / , without i t ; para-
physes branched, straight, slightly clavate.

On damp ground. Very rare. Figured from Phillips' sketches of the type
collection.

LAMPROSPORA De Notaris

Lamprospora De Notaris in Commentario della Societa Crittogamo-


logica Italiana 1:388 (1864).
Differs from Octospora only in its globose spores, which are usually
conspicuously ornamented by a reticulum or b y large spines. Though
nominally hairless, the apothecia tend to have a dentate-fimbriate
margin and their outer surface often bears delicate white fibrils. The
species occur on b u r n t ground and amongst small mosses.

Lamprospora crouani (Cooke) Seaver in Mycologia 6 : 8 (1914). (PI.


VIu.)
Apothecia up to 5 m m . diameter, saucer-shaped, sessile, disc convex,
orange, margin d e n t a t e ; outer surface light orange, pruinose. Asci up
to 2 6 0 x 2 5 / , ; spores uniseriate, globose, ornamented with a coarse-
meshed reticulum, the meshes 3 - 6 / , across, spores 17-19/t diameter
over all, containing one large oil d r o p ; paraphyses straight, slightly
enlarged upwards to 5/, at the tip.

Among Tortula ruraliformis, T. intermedia etc., on sandy soils and old


walls, November to April. Not uncommon. Figured from Cirencester,
Gloucestershire, 26. xii. 1951.
The valid name for this species may be Lamprospora miniata de Notaris
in Comm. Soc. Critt. Ital. 1:388 (1864). Peziza crouani Cooke in Journal of
Botany 2:151, was proposed in the same year, 1864, and it is not clear
which epithet has priority.

53
Lamprospora astroidea (Hazslinzky) Boudier, Hist. Class. Discom.
d ' E u r o p e : 6 8 (1907), is a v e r y similar species with smooth spores 1 4 -
16/« diameter and occurs on b u r n t ground.
Lamprospora dictydiola Boudier, Icones mycologicae, Livraison 5
(1905), has spores 12-15 /t across with a fine-meshed reticulum, the
meshes about 2 - 3 /« diameter.
Lamprospora polytrichi (Schumacher ex Fries) Le Gal in Bull. Soc.
mycol. France 5 6 : 3 9 (1940), has still more finely reticulated spores
only 10-13 /« diameter.

Lamprospora crec'hqueraultii (Crouan) Boudier, Icones mycologicae,


Livraison 13 (May 1907). (PI. VIY.)
Apothecia up to 5 m m . diameter, sessile, saucer-shaped, disc convex,
light orange, margin scarcely t o o t h e d ; outer surface concolorous,
minutely downy with thin-walled, obtuse, 2-3-celled hairs up to
60x15/« long. Asci up to 300x26/«; spores globose, 16-19/« diameter,
ornamented with spines 1 - 2 fi long, containing several oil drops; para-
physes straight, slightly enlarged upwards to about 5 - 6 /i at the tip.

On wet clay soils, June to October. Not common.

The var. macrantha Boudier, op. cit. (May 1907), differs in the m u c h
larger and longer spines, up to 5 ft long, on spores 20-25 /t diameter.

PULVINULA Boudier

Pulvinula Boudier in Bull. Soc. mycol. France 1:107 (1885).


Differs from Lamprospora in its more pulvinate apothecia and
especially in its m u c h branched paraphyses, strongly curved at the
tip and not clavate; spores smooth, with several oil drops, typically
globose, b u t see note below.

Pulvinula constellatio (Berkeley & Broome) Boudier, Hist. Class. Dis-


com. d ' E u r o p e : 7 0 (1907). (PI. VIT.)
Apothecia up to 10 m m . diameter, sessile, thin-fleshed, pulvinate
with an undulating margin, disc bright reddish-orange, outer surface
pinkish. Asci up to 250x22/«; spores globose, smooth, 13-15/« dia-
meter, crowded with small oil drops; paraphyses slender, branched,
strongly curved towards the tip, which is only 3 n thick and filled with
reddish granules.

54
On burnt ground and on sandy soil amongst mosses, September to May.
Not uncommon. Figured from burnt ground, Worcestershire, 28. v. 1912.

Pulvinulacinnabarina{Fuckel) Boudier, op. cit.:70 (1907), is doubt-


fully distinct and is an older name. An almost identical fungus b u t
with broadly elliptical spores 1 2 x 8 f t has been called Ascophanus
globoso-pulvinata (Crossland) Boudier in Trans. Brit, mycol. Soc. 4 : 3 6 9
(1914), a disposition which well illustrates t h e artificial character of
genera based on spore shape.

PSILOPEZIA Berkeley

Psilopezia Berkeley in Hooker's London J o u r n a l of B o t a n y 6 : 3 2 5


(1847).
Apothecia sessile, lenticular, firmly adhering to wood over a broad
base, disc flat or convex, yellow or b r o w n ; asci 8-spored, spores
elliptical, smooth, hyaline, containing oil drops, paraphyses simple,
clavate.

Psilopezia babingtonii (Berkeley) Berkeley, Outlines of British Fungo-


logy:373 (1860). (PI. IVK.)
Apothecia gregarious, pulvinate, sessile on a broad base, disc up to
1 cm. across, reddish-brown; flesh translucent, gelatinous, formed of
undulating, slender, loosely woven hyphae, surface layer three to five
cells thick, formed of globose cells 40-60 /« across; outer surface smooth,
pale, ochre-brown. Asci up to 390x20/«; spores broadly elliptical,
smooth, hyaline, 19-21x12-13/«, containing two large oil drops in a
granular cytoplasm; paraphyses slender, simple, apex clavate, up to
8 fi thick, brown.

On wet dead wood, often lying in water, apparently rare. Figured from
wet wood near Halifax, Yorkshire, 10. vi. 1899.
This peculiar species has been recorded also under the names Psilopezia
myrothecioides Berkeley & Broome in Ann. Mag. nat. Hist. IY, 15 :39 (1875);
Pachyella depressa (Phillips) Boudier, Hist. Class. Discom. d'Europe:51
(1907) and Humaria oocardii (Kalchbrenner) Saccardo, Sylloge Fungorum 8:
144 (1889).

Family 5. Ascobolaceae

The Ascobolaceae are characterised mainly b y their relatively broad


asci which protrude above the general level of the hymenium as t h e y

55
ripen. The spores commonly lie in two or three irregular rows in the
ascus instead of in a single vertical row as in other families. The apo-
thecia are pulvinate, usually very small, and occur mainly on dung, a
few species on b u r n t ground, soil or debris. There are two tribes:

I. Ascospores dark-coloured at maturity, elliptical or, if globose, there is a


purplish outer layer to the spore wall soluble in alkalis Ascoboleae
II. Ascospores permanently colourless to yellowish or, if dark then globose
with an ornament of spines and ridges and the apothecium is reduced
to a fascicle of asci Pseudoascoboleao (p. 60)

Tribe Ascoboleae

The spores of the Ascoboleae have a double wall, a hyaline persistent


inner layer like the spore wall of other species, and a dark brown or
purple outer layer, t h e epispore, which commonly cracks into a
characteristic p a t t e r n . T h e epispore swells up and disappears i n K O H
solution and dried specimens of t h e Ascobolaceae should never be
soaked in this reagent to soften t h e m for microscopic examination. In
most genera the spores are ejected singly from the ascus in the normal
way b u t in Saccobolus t h e y are firmly united to form an irregular mass
which is ejected at a single operation. The tribe comprises 12 British
genera:

I. Spores not cohering within the ascus but ejected singly:


A. Apothecia not hairy, though often furfuraceous Ascobolus
B. Apothecia inconspicuously hairy, spores 50 [J. or more long
Dasyobolus (p. 59)
II. Spores ejected together as a single projectile . . . Saccobolus (p. 59)

ASCOBOLUS Persoon ex Fries

Ascobolus Persoon ex Fries, Systema mycologicum 2:161 (1822).


Apothecia cup-shaped, smooth or scurfy, disc often dark or yellow-
ish-olive, sometimes with a slightly greenish hue, dotted with the
almost black protruding tip of ripe asci. Ascospores four or eight,
purplish, elliptical, the epispore either verrucose or cracking into striae
which are usually more or less interconnected, rather variable in shape,
size and ornament. Most species occur on dung or richly manured soil,
a few on b u r n t ground, bare soil or decaying plant debris. About 14
British species have been recognised b u t the genus has now been

56
critically revised by van Brummelen as Persoonia Suppl. Vol. I 1967
wich m a y stimulate collecting and reveal others.

Ascobolus furfuraceus Persoon ex Fries, op. cit.: 163 (1822). (PL VIIH.)

Apothecia cup-shaped, up to 5 m m . diameter, disc yellowish, be-


coming dark brown with age, flat; outer surface yellowish then brown,
coarsely scurfy or mealy. Spores elliptical, violet, t h e n brown, orna-
mented with longitudinal occasionally anastomosing lines, about eight
or nine of which are visible on each view of the spore, 22-28 x 11-13 /<;
paraphyses slender, embedded in yellow mucus.

j3n dung, especially of the cow. May to October. Common. Figured from
Kew, Surrey, 25. x. 1954.

Ascobolus stercorarius (Bulliard) Schroeter, K r y p t . Flora Schlesien 3


(2): 56 (1893), emend Olive in Mycologia 46:105 (1954), is very similar
but may be distinguished by its shorter ascospores, 17-22x9.5-12.5 n,
on which about five to seven freely anastomosing striae are visible at
one time. Van Brummelen thinks t h e m indistinguishable.

Ascobolus viridis Gurrey in Trans. Linn. Soc. London 24:154 (1863).


(Pl. V I I J ) .
Apothecia saucer-shaped, up to 5 m m . diameter, disc concave or
flat, dingy yellowish-green with a prominent crenulate margin, outer
surface light brown, coarsely mealy. Spores markedly fusiform, viola-
ceous, 3 0 - 3 5 x 1 3 - 1 5 /i, ornamented with repeatedly anastomosing or
reticulating striae; paraphyses somewhat clavate, up to 10/, thick at
the tip, embedded in yellowish-green mucus.

On bare clay soil, August to October. Uncommon. Figured from near


Hebden Bridge, Yorkshire, 3. viii. 1898, with a spore from the type collection.

Ascobolus dcnudatus Fries, Systema mycologicum 2:162 (1822). (PL


VIIK.)
Apothecia cup-shaped, up to 5 m m . diameter, disc concave t h e n flat,
yellowish-green, becoming grey-brown; outer surface almost smooth.
Spores narrowly elliptical, 1 7 - 2 0 x 8 - 9 ft, violet t h e n brown, ornament-
ed with closely spaced subparallel lines t h a t only occasionally anasto-
mose; paraphyses slender, embedded in yellowish mucus.

On plant debris, October to November. Uncommon. Figured from com-


posted bracken, Nottingham, 25. xi. 1939.

57
Ascobolus crouani Boudier in Ann. Sci. Nat. Bot. V, 10:216 (1869),
is said to differ mainly in its mealy outer surface.

Ascobolus albidus Crouan in Ann. Sci. Nat. Bot. Ser. 4 , 1 0 : 1 9 3 (1858).


(PI. V I I L . )
Apothecia obconical, 0.3-1 m m . diameter, disc flat, dingy ochraceous
with a few blackish asci protruding, outer surface smooth, fawn,
orange, reddish or purplish. Spores elliptical, 22-26x11-13/«, violet,
ornamented with rather close anastomosing striae; paraphyses slender,
up to 5 fi thick at the tip.
On dung, especially of cow but also that of horse, sheep and rabbit,
throughout the year. Common. Figured from cow dung, Crimsworth Dean,
Wadsworth, Yorkshire, 30. i. 1898. This is Ascobolus glaber of most authors
but according to Dr. van Brummelin (in litt.) it is not A. glaber Persoon ex
Fries 1822. The form on rabbit droppings has been confused with Ascophanus
vinosus (see p. 52).

Ascobolus stictoideus Spegazzini in Michelia 1:474 (September 1879),


is an equally minute species found on dog dung, in which the spore
ornament is finely areolate.

Ascobolus carbonarius Karsten in Not. Sallsk. F a u n a Flora Fennica


11:202 (1870). (PI. VIIi.)
Apothecia cup-shaped, up to 5 m m . diameter, disc concave t h e n
flattened, with a slightly crenulate margin, dark brown to almost
black; outer surface finely mealy, blackish-brown. Spores elliptical,
19-22x11-15/«, purplish-brown, irregularly w a r t e d ; paraphyses
branched, slender, 3 /< thick, embedded in yellowish-green mucus.
On burnt ground, March to August. Not uncommon but not easily seen.
Figured from Norbury Park, Mickleham, Surrey, 20. vi. 1948.

Ascobolus atrojuscus Phillips & Plowright in Grevillea 2:186 (June


1874), is a synonym.
Ascobolus leveillei Boudier in Ann. Sci. Nat. Bot. V, 10:225 (1869),
is a moderately sized species found on dung, with a greenish disc up
to 1 m m . across and spores 2 0 - 2 6 x 1 2 - 1 3 / « which appear smooth or
only very finely p u n c t a t e . The correct name is A. boudieri Quelet.

Ascobolus crosslandii Boudier in Bull. Soc. mycol. France 14:126


(1898). (PI. VIIIE.)
Apothecia cupshaped, disc up to 2 m m . across, concave or flat, pale
yellowish-green; outer surface concolorous, very minutely scurfy.

58
Ascospores globose, 10-12/* diameter, violet, t h e n purplish-brown,
ornamented with anastomosing striae; paraphyses hyaline, clavate,
6 - 8 fi thick at the tip.
dog dung. October to November. Apparently very rare but the sub-
strate is generally neglected by collectors. Figured from Crossland's sketches
of the type, Yorkshire 1. xi. 1897. The epithet was published by Boudier as
"Crowslandi" and the collector as "Crowsland", an obvious error for Cross-
land which has been corrected in all subsequent publications.

DASYOBOLUS (Saccardo) Saccardo

Dasyobolus (Saccardo) Saccardo, Sylloge F u n g o r u m 11:421 (1895).


Apothecia like those of Ascobolus b u t with a downy margin. There
is one British species, referred to Ascobolus b y v a n Brummelen.

Dasyobolus immersus (Persoon ex Fries) Saccardo, op. cit. :421 (1895).


(PI. VIIIc.)
Apothecia subcylindrical or hemispherical, immersed in dung up
to the margin, disc concave or flat, up to 1 m m . across, yellowish-green,
margin fringed with fascicles of non-septate brownish hairs a b o u t
2 5 - 5 0 x 3 - 6 /x. Spores elliptical, 5 0 - 7 5 x 3 0 - 4 0 /*, violaceous t h e n brown,
smooth or with a single longitudinal split in the epispore, less often
with two or three anastomosing splits; paraphyses slender, colourless,
up to 3 /«thick. The marginal hairs are often absent.

On cow dung, less often on that of sheep or horse, especially in autumn.


Common. Figured from Hardcastle, Hebden Bridge, Yorkshire, 1. xi. 1897.

SACCOBOLUS Boudier

Saccobolus Boudier in Ann. Sci. Nat. Bot. V, 10:228 (1869).


Apothecia minute, seldom exceeding 1 mm., cup-shaped to lenti-
cular, sessile. Asci short and broad, 4- or 8-spored, spores firmly united
in a single cluster and ejected together, purplish or brown, smooth,
finely granulate or with loose granules deposited on their exposed
faces. All occur on dung except S. obscurus. See v a n Brummelen 1967.

Saccobolus glaber (Persoon) L a m b o t t e . (PL VIIi.)


Apothecia up to 1 m m . diameter, cup-shaped, disc flat or convex,
ochraceous; outer surface concolorous, smooth. Asci 120-160x20-33/«;
spores subfusiform with rounded ends, smooth to finely p u n c t a t e ,

59
20-27x10-12/«, violaceous or brown; paraphyses slender, slightly cla-
vate and curved at the tip, yellow.
Ondung of cow andsheep, June to September. Uncommon. Figured from
cow dung, near HucktersJield, Yorkshire, 24. ix. 1906.

S. granulospermus Soppitt&Crossland in The Naturalist: 30 (1899),


is a synonym, as is S. Kerverni (Crouan) Boudier.

Saccobolus obscurus (Cooke) Phillips, Manual of British Discomy-


cetes: 2 9 5 ( 1 8 8 7 ) . (PI. V I I I B . )
Apothecia lenticular, disc up to 0.5 m m . diameter, flat, t h e n convex,
grey-brown; outer surface smooth, paler. Asci 90-100x22/«; spores
oval, pointed at the ends, 14-16x7-9/«, purplish-brown, verrucose;
paraphyses much branched, tips slightly clavate.
On rotting sacking, June. Uncommon. Figured from old sacking, Book-
ham Common, Surrey, 26. vi. 1947.

Saccobolus quadrisporus Massee & Salmon in Annals of B o t a n y 15:


329 (1901) was distinguished by its 4-spored asci.
Saccobolus depauperatus (Berkeley & Broome) Hansen in Vid. Meddr.
dansk n a t . Foren.
87 (1876), on dung, is less t h a n half a millimetre across, with spores
only 13-15x6-7/«. Saccobolus neglectus Boudier is a synonym.

Tribe Pseudoascoboleae

The following British genera m a y be recognised:

I. Apothecia setose Lasiobolus (p. 62)


II. Apothecia glabrous:
A. Asci blued by iodine, genera accordingly referred to Pezizeae by
some authors:
1. Asci 8-spored . Iodophanus (p. 64)
2. Asci polysporous Thecotheus (p. 65)
B. Asci not blued by iodine:
1. Asci 8-spored:
a. Apothecia densely gregarious or confluent, seated on a
white subiculum covering burnt soil or damp plaster
Pyronema (p. 62)

60
b. Apothecia without a subiculum, on dung, soil or debris:
*Apothecia relatively large, often downy, asci nar-
rowly cylindrical, see Fimaria, p. 51.
**Apothecia small to very small, asci broad, clavate:
fAscospores elliptical . . . Ascophanus (p. 63)
tfAscospores globose:
§Apothecia discoid with a normal hymenium,
ascospores smooth, pale
Sphaeridiobolus
§§Apothecia globose, very minute, reduced to
a fascicle of asci and paraphyses, asco-
spores dark brown, ornamented by spines
and ridges . . . . Ascodesmis
2. Asci polysporous:
a. Apothecium (or cleistocarp?) containing a single ascus,
see Thelebolus, p. 70
b. Apothecium minute but with a hymenium and several
asci:
*Asci opening by a normal operculum
Rhyparobius (p. 65)
**Asci opening by a vertical slit as far down as an an-
nular thickening Ascozonus (p. 66)
The species of Ascodesmis have minute fructifications too small to be figured
here. For an account of the genus see Obrist in Canadian Journal of Botany
39:943-953 (1961).

SPHAERIDIOBOLUS Boudier
Sphaeridiobolus Boudier in Bull. Soc. mycol. France 1:108 (1885).
The species with globose spores having a purplish-brown outer wall
like t h a t of other Ascobolus species have been transferred to t h a t
genus. The type of Sphaeridiobolus is Boudier's concept of Ascobolus
hyperboreus Karsten, which Madame Le Gal has shown to be the same
as Peziza albofusca Grouan, see Revue de Mycologie 18, 90-91, 1953.

Sphaeridiobolus albofuscus (Crouan) Boudier, Hist. Classif. Discom.


d'Europe 7 4 ( 1 9 0 7 ) . ( P I . V I I I H . )
Apothecia discoid, about 1 m m . across, whitish. Asci 110-160 x
18-22//, spores biseriate, 12.5-14/t diameter, colourless to yellowish,
finely warted when mature, paraphyses slightly curved, enlarged to
4/, wide at the tip.
On dung, especially of rodents. Apparently uncommon.
Plicaria murina Fuckel seems to be the same and offers the oldest
name for the species.

61
LASIOBOLUS Saccardo

Lasiobolus Saccardo in Botanisches Centralblatt 18:220 (1884).


Apothecia sessile, minute, cup-shaped or obconical, somewhat
sparsely set with stiff, thick-walled, hyaline setae; asci not blued by
iodine, 8-spored, spores elliptical, smooth, hyaline. It seems probable
t h a t there is b u t a single variable British species:

Lasiobolus ciliatus (Schmidt ex Fries) Boudier, Hist. Class. Discom.


d ' E u r o p e : 78 (1907). (PL V I I I A . )
Apothecia sessile, obconical, disc flat, about 0.5-0.75 m m . diameter,
varying from yellowish to brick colour, soft-fleshed; outer surface con-
colorous, beset with stiff, smooth, thick-walled, colourless, pointed,
nonseptate setae, usually up to about 400 /< long and 30/« thick at the
base. Asci cylindric-clavate, short-stalked, up to 300x35/«, 8-spored;
spores uniseriate, elliptic-cylindric, 20-25x 11-14/«, without oil drops;
paraphyses slender, forked, 2 - 3 /« thick, with more or less orange
contents.

On_dung of all kinds, throughout the year. Very common. Figured from
sheep dung, Isle of Yell, Zetland, 5. ix. 1952. Forms with unusually short
hairs have been called L. oligotrichus A. L. Smith & Ramsbottom in Trans.
Brit, mycol. Soc. 5:238 (1916), those with very long hairs L. macrotrichus
Rea in Trans. Brit, mycol. Soc. 5:440 (1917) and those with spores over 25/<
long L. ruber (Quelet) Saccardo, Sylloge fungorum 8:537 (1889).

PYRONEMA Carus

Pyronema Carus in Nov. Act. Acad. Caes. Car. Leop.-Nat. Cur. 17:370
(1835).
Apothecia pulvinate, densely gregarious, often becoming confluent,
seated on a conspicuous, superficial, white mycelium; asci 8-spored,
not blued by iodine, spores elliptical, smooth, hyaline, w i t h o u t oil
drops; paraphyses simple.

Pyronema omphalodes (Bulliard ex St. Amans) Fuckel in J a h r b . Nass.


Vereins f. N a t u r k u n d e 23-24:319 (1870). (PL V I I I L . )
Apothecia sessile, disc convex, up to 1 m m . diameter b u t often be-
coming confluent in large patches, light red to reddish-orange, without
a raised margin, surrounded b y a conspicuous, cobweb-like, white
mycelium. Asci up to 150x15/«; spores elliptical, smooth, l l - 1 5 x

62
6.5-8.5 /t, without oil drops; paraphyses simple, slightly clavate, 3-4,«
thick at the tip.
On burnt ground, throughout the year. Common but apparently less so
than the following species. Figured specimen, upper spore, from burnt
ground with Anthracobia melaloma, in beech woods, Norbury Park, Mickle-
ham, Surrey, 19. vi. 1950.

Pyronema domesticum (Sowerby ex Fries) Saccardo, Sylloge Fungo-


r u m 8:109 (1889), differs only in its larger spores, 15-18x9.5-12/«, see
the lower spore in PI. V I I I L , and is very common on wet plaster, b u r n t
soil and especially on steam-sterilised soil.
Pyronema glaucum (Boudier) Saccardo, Sylloge F u n g o r u m 8:110
(1889), is a colourless state not infrequent on steam-sterilised soil.
Pyronema buchsii Hennings in Hedwigia 41:164 (1902), a species of
mossy ground in coniferous forests of central Europe, appears to be
an Octospora, as indicated b y Boudier, who referred it to " H u m a r i a
F r " . , i . e . Octospora S. F. Gray. Though its apothecia are densely gre-
garious on a white subiculum t h e y have a distinct denticulate margin
and fusoid ascospores 19-22x6-9/«, with 2 - 3 oil drops.

ASCOPHANUS Boudier

Ascophanus Boudier in Ann. Sci. Nat. Bot. V, 10:241 (1869).


This genus is ill-defined, largely b y negative characters, and contains
a residue of species not yet disposed in b e t t e r understood genera. Its
apothecia are small, seldom more t h a n 1 m m . across, usually m u c h
less, pulvinate or lenticular, externally smooth to minutely mealy, the
hymenium typically papillate with protruding asci at m a t u r i t y ,
spores elliptical, smooth, without oil drops, asci not blued b y iodine,
paraphyses usually slender, sometimes forked. Most species occur on
dung or freshly manured ground. (See also Coprobia and Fimaria.)

Ascophanus argenteus (Currey) Boudier in Ann. Sci. Nat. Bot. V, 10:


245 (1869). (PI. V I I I J . )
Apothecia subpyriform, sessile, up to 250 fiacross, white t h r o u g h o u t .
Asci 40-50x20/«; spores elliptical, 10-12x6-7/«, clustered and ejected
in a mass; paraphyses clavate.
•, December. Uncommon. Figured from Halifax, Yorkshire,
27. xii. 1898.

63
Ascophanus lacteus (Cooke & Phillips) Phillips, Manual of British
Discomycetes:306 (1887), is similar b u t with spores 6 - 8 x 3 - 4 / « .

Ascophanus microsporus (Berkeley & Broome) Phillips, Manual of


British Discomycetes:307 ( 1 8 8 7 ) . (PI. V I I I G . )
Apothecia gregarious, sessile, disc flat or convex, about 250 /« dia-
meter, yellowish-brown; outer surface smooth. Asci clavate, a b o u t
50x14/«; spores biseriate, elliptical, 7 - 8 x 3 . 5 - 4 / « ; paraphyses slender
with much swollen apices 5 - 6 /«thick, containing greenish sap.

On cow dung in autumn. Rare. Figured from Halifax, Yorkshire, 17. x.


18987~This is perhaps the same as A. subfuscus (Crouan) Boudier in Ann. Sci.
Nat. Bot. Y, 10:242 (1869).

IODOPHANUS Korf

Iodophanus Korf in American J o u r n . B o t a n y 5 4 : 1 8 (1967).


Apothecia minute, pulvinate, pinkish, asci clavate, their walls blued
b y iodine, ascospores hyaline, very finely warted, without oil drops.

Iodophanus carneus (Persoon) Korf. op. cit. 19 (1967). (PI. VIIIF.)

Apothecia lenticular, up to 1.5 m m . diameter, sessile, pink, salmon


or flesh colour, disc convex and papillate with the protruding asci. Asci
broadly clavate, up to 225x30/«; ascospores 1-2-seriate, elliptical,
18-22 (-30) x 10-12 (-18)/«, very finely p u n c t a t e and usually described
as smooth; paraphyses stout, simple or forked, septate, up to 8/« thick
at the tip. The whole h y m e n i u m stains blue with iodine.

On dung, rotting vegetable matter, including textiles and rope, and on


soil, throughout the year. Very common. Figured from rotting hay, Hills-
borough, County Down, 17. ix. 1948. There is an CEdocephalum conidial state.

Ascophanus salmonicolor (Berkeley & Broome) Boudier, Hist, class.


Discom. d ' E u r o p e : 7 6 (1907), A. saccharinus (Berkeley & Curtis)
Boudier, in Ann. Sci. n a t . Bot. V. 10:251 (1869), and A. testaceus
(Mougeot ex Fries) Phillips, Manual of British Discomycetes:310
(1887), are probable synonyms and A. keithii (Phillips) Boudier in
Trans. Brit, mycol. Soc. 4:369 (1914), seems to be only a narrow-spored
variant with ascospores 19-21x8-9/«.

64
T H E C O T H E U S Boudier

Thecotheus Boudier in Ann. Sci. Nat. Bot. V, 10:235 (1869).


Apothecia shortly cylindrical or pulvinate; asci very large, p r o t r u d -
ing beyond the disc at m a t u r i t y , blue in iodine, polysporous; growing
on dung.

Thecotheuspelletieri(Crouan) B o u d i e r , o p . c i t . : 2 3 6 (1869). (PI.VIIIM.)

Apothecia 0.5-1 m m . across, disc-like, flat, white to faintly lilaceous;


outer surface minutely pruinose. Asci 370-430x40-80/«; 32-spored,
spores elliptic-fusiform, smooth, hyaline, 3 0 - 3 5 x 13-17 /,, thick-walled;
paraphyses slender, forked, slightly enlarged to 5 - 7 /, thick at t h e tip.
On dung of cowjamQiorse, July to August. Rare. Figured from cow dung
fromBelgmm

R H Y P A R O B I U S Boudier

Rhyparobius Boudier in Ann. Sci. N a t . Bot. V, 10:237 (1869).


Apothecia minute, seldom as m u c h as 0.5 m m . across, white,
yellowish or brown, asci few, relatively short and broad, with 16 or
more spores per ascus, in some species 100 or more; spores elliptical,
smooth, hyaline; paraphyses scanty. A b o u t t e n species have been re-
cognised in Britain b u t the genus is m u c h in need of revision. The
following are representative:

Rhyparobius soxdecemsporus (Crouan) Saccardo, Sylloge F u n g o r u m 8:


541 (1889). (PI. V I I I o . )
Apothecia scattered or gregarious, sessile, subglobose, disc finally
flat or slightly convex, whitish t h e n cream; a b o u t 0.5 m m . across;
outer surface smooth. Asci cylindric-clavate, 50-100x 15-22 /,; rounded
above, containing 16 spores in two rows, spores broadly elliptical,
8 - 1 2 x 5 - 7 /,; paraphyses slender, often branched, tips curved b u t
scarcely thickened, 2.5 /, wide.
On_rlmio- nf Hngj hnryp fltp. in summer and autumn. Uncommon. Figured
from dog dung, Hardcastle, Yorkshire, 15. vii. 1899.

Rhyparobius cookei (Crouan) Boudier in Ann. Sci. Nat. Bot. V, 10:238


(1869). (PL VIIIis.)
Apothecia gregarious, cup-shaped, disc flat, a b o u t 200/, across,
yellowish-brown; outer surface smooth, darker. Asci broadly clavate,

65
60-70x20-27/«, containing up to 64 spores, spores elliptical, 8 - 1 0 x 4 -
5/i, smooth; paraphyses scanty, slender.
On dung. Uncommon. Figured specimen from dog dung, Halifax, 14. ii.
1899.

ASCOZONUS (Renny) Boudier


Ascozonus (Renny) Boudier in Bull. Soc. mycol. France 1:109 (1885).
Apothecia minute, less t h a n 1 m m . across, whitish to yellowish,
margin fringed with downy white hairs. Asci not blued b y iodine, poly-
sporous, not t r u l y operculate b u t having an annular thickening near
the tip and opening b y a bilabial split down to this ring; spores ellipti-
cal or fusiform, smooth, hyaline, paraphyses slender. The species all
occur on dung, especially t h a t of birds, in winter. Seven British species
are recognised b u t t h e y have been m u c h neglected and the best ac-
count of the genus is still t h a t of Renny in Journal of Botany 12 (1874).
The following species is fairly representative b u t in others the n u m b e r
of spores per ascus ranges from 16 to about 128.

Ascozonus woolhopensis (Berkeley & Broome) Boudier, Hist. Glass.


Discom. d ' E u r o p e : 7 9 (1907). (Fig. 6K and Fig. 1B.)
Apothecia gregarious, t u r b i n a t e , disc flat, about 0.5 m m . across,
whitish becoming yellowish, fringed with downy hairs which are uni-
cellular, thinwalled, obtuse, hyaline, about 25-30x 7 - 8 n; outer surface
granulate below with large globose protruding cells. Asci clavate, about
9 0 x 3 0 /x, containing 64 spores; spores fusiform, 1 2 - 1 3 x 3 - 3 . 5 /i, slightly
yellowish.
OnjluggJif rabbit, etc. in autumn and winter.

Family 6. Sarcoscyphaceae

Fruit bodies developed on woody substrata, either tomentose or


hairy, flesh largely or entirely formed of filamentous hyphae, some-
times very t o u g h ; asci not blued b y iodine, usually very long-c,ylindric
with elongated flexuous stalks and r a t h e r thick-walled; spores ellip-
tical or globose, colourless, smooth in the British species. The family
is p r e d o m i n a n t l y tropical in distribution and contains only five British
members, belonging to as m a n y genera, representative of two tribes:

I. Outer surface never black but red-brown, yellowish-brown, bright red,


yellow or pink; paraphyses of some species green with iodine
Sarcoscypheae (p. 67)

66
II. Outer surface never bright coloured, generally blackish and seated on
a dark brown mycelium enveloping the substrate, or on a sclerotium.
See also Desmazierella p. 33. Urnuleae (p. 68)

Tribe Sarcoscypheae
The more i m p o r t a n t genera are restricted to the tropics and only the
following arc known to occur in Britain:
I. Small species on branches of conifers, spores globose . . . . Pithya
II. Apothecia larger, stalked, on dead wood, spores elliptical:
A. Apothecia devoid of gelatinous tissue Sarcoscypha
B. Apothecia with a gelatinous layer in the flesh Microstoma (p. 68)

PITHYA Fuckel

Pithya Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:317 (1870).


Only one British species is k n o w n :

Pithya cupressina (Fries) Fuckel op. cit.:317 (1870). (PI. VIIG.)

Apothecia solitary, about 1 m m . broad and high, disc flat, bright


orange; outer surface smooth, paler, with a short stout stalk, downy
at t h e base. Asci 2 1 0 x 1 2 8 - s p o r e d , spores uniseriate, globose,
smooth, 9-10,« diameter; paraphyses cylindrical, very slightly en-
larged up to 4/« at the tip.
On dead leaves of Jjmi^erus, July. Rare. Figured on Juniper from East
Mailing, Kent, 17. vii. 1943.
A very similar species P. vulgaris Fuckel, op. cit.: 317 (1870), occurs on
leaves of Abies and Picea in Europe.

SARCOSCYPHA (Fries) Boudier

Sarcoscypha (Fries) Boudier in Bull. Soc. mycol. France 1:103 (1885).


Apothecia stipitate to substipitate, subglobose then cup-shaped
with a more or less incurved margin, outer surface downy. Asci 8-spor-
ed, spores elliptical or oblong, smooth, paraphyses forked or sparingly
branched. There is only one British species:

Sarcoscypha coccinea (Fries) L a m b o t t e in Mem. Soc. roy. sciences


Liege II, 14:302 (1887). (PI. VIIc.)
Apothecia gregarious, subsessile to short-stalked, cup-shaped, up to
4 cm. across, margin somewhat inrolled, h y m e n i u m scarlet; outer sur-

67
face white, floccose with more or less m a t t e d hyaline hairs. Asci up to
400x16,«; spores uniseriate, hyaline, elliptic-cylindric with obtuse
ends, 24-32 (-40) x 12-14/«, usually with small oil droplets cohering in
two clusters, one at each end of the spore; paraphyses slender, forking
near the base, not enlarged upwards, 3 p thick, with red granular con-
tents. Forms with a white h y m e n i u m occasionally occur.

On fallen sticks in woods and hedgerows, December to February, oc-


casionally as late as March. Locally abundant. Even this, perhaps the most
beautiful of British ascomycetes, is without any generally recognised verna-
cular name. In North Somerset it was formerly known to country children
as "Moss cups" but this term seems to have been applied more widely to the
podetia of species of Cladonia.

MICROSTOMA Bernstein

Microstoma Bernstein in Nov. Acta Acad. Caes. Leop.-Carol N a t u r . 23,


2:649 (1852).
Apothecia clustered, stalked, arising from a hard black pseudorhiza
which is usually attached to buried wood, cup-shaped then expanded
with the margin often lobed, outer surface downy. Flesh of cup and
stalk contains a gelatinous layer. There is one rare British species:

Microstoma protracta (Fries) Kanouse in Mycologia 40:486 (1948).


(PI. V I I D . )
Apothecia arising from an immersed, hard, elongate pseudorhiza,
attached to buried sticks or roots, aerial stalk long, slender, often
branched, lower half dark coloured, upper half whitish, downy with
hyaline hairs, cup up to 2 cm. across, cup-shaped t h e n flattened and
torn at t h e margin, h y m e n i u m vivid rose-red, outer surface orange.
Asci up to 275x23/«, 8-spored, operculum somewhat laterally placed;
spores uniseriate, elliptic-fusoid, 24-45x10-14/«, containing several
oil drops; paraphyses slender, dichotomously branched, only slightly
enlarged at the tips, with red contents which t u r n green with iodine.

Scottish Highlands, April. Apparently very rare or possibly merely


ignored because there are no local mycologists to report it.

Tribe Urnuleae

Apothecia blackish, seated on a brown mycelium, paraphyses never


green with iodine.

68
I. Ascospores globose Pseudoplectania
II. Ascospores elliptical:
A. Apothecia large, globose, then turbinate, flesh watery-gelatinous
and very thick when fresh Sarcosoma
B. Apothecia cupulate to discoid, relatively thinfleshed:
1. Apothecia small, subsessile, receptacle dusted with red-lead-
coloured granules Plectania
2. Apothecia very large, stipitate, dark gray throughout
Urnula
Pseudoplectania and Plectania occur rarely in Britain; Sarcosoma
globosum (Schmidt ex Fries) Rehm and Urnula craterium (Schweinitz)
Fries are european species not yet known here.

PSEUDOPLECTANIA Fuckel

Pseudoplectania Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:


324 (1870).
Apothecia sessile, cup-shaped t h e n flattened, black throughout, ex-
ternally hairy, asci not blued with iodine, 8-spored, spores globose,
hyaline, smooth.

Pseudoplectania nigrella (Persoon ex Fries) Fuckel, op. cit.:324 (1870).


(Fig. 6 H . )
Apothecia gregarious, sessile, up to 3 cm. across, cup-shaped or ex-
panded, black t h r o u g h o u t , exterior tomentose with black u n d u l a t i n g
or contorted hairs. Asci up to 3 5 0 x 1 6 / , ; spores uniseriate, globose,
smooth, filled with numerous small oil drops, 10-12 /t diameter; para-
physes slender, forked, not expanded at the tip, brown.
On the litter in coniferous woods. March. Rare.
Pseudoplectania vogesiaca (Persoon) Seaver ( = Melascypha melaena
(Fries) Boudier) is a european species with larger, stipitate, cupulate
apothecia, found on coniferous logs.

PLECTANIA Fuckel

Plectania Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 2 3 - 2 4 : 3 2 3


(1870).
Apothecia more or less stalked with a subiculum of dark brown
hyphae, cup-shaped, very dark olive-grey to black, downy; flesh con-
taining a well developed gelatinous layer. Asci not blued with iodine,
8-spored, spores elliptical, hyaline, smooth, paraphyses slender, branch-

69
ed. The genus is separated from Urnula by the gelatinous layer in the
flesh. (But see Le Gal in Bull. Soc. mycol. France 74:155-177 (1958).

Plectania melastoma (Sowerby ex Fries) Fuckel, op. cit.:324 (1870).


(PI. V H . )
Apothecia cup-shaped, up to 2 cm. across, seated on a r u d i m e n t a r y
stalk, disc black, p e r m a n e n t l y concave; outer surface black, minutely
downy with closely septate hairs whose t h i n walls are often encrusted
with red granules, especially near the margin; subiculum of slender,
wiry, black mycelium. Asci up to 450x12/«; spores uniseriate, fusi-
form, smooth, 2 3 - 2 8 x 10-11 /«, at first containing m a n y oil drops which
disappear at m a t u r i t y ; paraphyses slender, repeatedly forked, brown.

On debris in woods, April to May. Apparently common a century ago but


not reported for manjf years.

Family 7. Thelebolaceae

There is a single genus, Thelebolus, which is referred b y some authors


to Erysiphales. The ascus structure and the closed ascocarp do suggest
a remote affinity with t h a t group b u t the saprophytic habit and lack
of appendages to the ascocarp wall indicate t h a t the relationship, if it
exists, must have been at some remote date. For a detailed account
of the genus see Cooke and Barr in Mycologia 56, 763-769, 1964.

THELEBOLUS Tode ex Fries

Thelebolus Tode ex Fries, Systema mycologicum 2:306 (1823).


Apothecia minute, light coloured, ovoid to turbinate, enclosing a
single elliptical polysporous ascus, or at least only one ascus m a t u r i n g
at a time, spores elliptical, colourless, nonseptate.

Thelebolus stercoreus Tode ex Fries, Op. cit. 307 (1823). (Fig. 29M)
Apothecia yellowish-brown, ovoid, less t h a n y 2 m m . broad, smooth,
partially embedded in dung. Ascus very large, elliptical, up to 230 x
140 fi, with wall 5-10 /«thick, having a thinner annular zone near the
apex along which it breaks open at m a t u r i t y , spores about 1,000 per
ascus, 5 - 7 x 3 - 4 /«.
There is no hymenium and the ascus is p e r m a n e n t l y enclosed b y the
ascocarp wall.

70
Order 2. TUBERALES

A subterranean habit has been adopted b y fungi of the most diverse


affinities including Phycomycetes (Endogone), Basidiomycetes (four
families of "Gasteromycetes") and at least two orders of Ascomycetes,
the Tuberales and the Plectascales. The former are probable derivati-
ves of the Pezizales, in which t h e asci have lost their opercular de-
hiscence and are enclosed in more or less p e r m a n e n t l y closed fruit
bodies. In some genera the fructification has become so elaborately
convoluted and infolded as to be fused together into a solid mass,
with or without narrow cavities lined b y the h y m e n i u m . In t h e most
advanced genera the asci are no longer even arranged in a hymenium
at all b u t are dispersed through the flesh. The British species are
assigned to 11 genera, representing four families. T h e y have recently
been ably monographed by Dr. L. E. Hawker in Phil. Trans. Roy. Soc.
London 237 (1954) and b y Lange in Dansk. Bot. Arkiv. 16 (1956), so
it will suffice to indicate the distinctive characters of the genera b y
illustrating a few representative species.

I. Spores smooth:
A. Asci cylindrical, fruit body with a single cavity opening by a broad
pore Stephensia (p. 73)
B. Asci elliptical, fruit body closed:
1. Flesh containing many small cavities . . Balsamia (p. 74)
2. Flesh solid, see Diehliomyces (p. 345)
II. Spores ornamented at maturity:
A. Asci arranged in a regular hymenium:
1. Paraphyses fused at the tip to form an epithecium
Genea (p. 73)
2. Paraphyses not fused at their tips:
a. Fruit body much convoluted but thin-fleshed and hollow
Gyrocratera (p. 72)
b. Fruit body becoming solid and chambered Hydnotrya
B. Asci not in a regular hymenium:
1. Flesh of the fruit body marbled by darker and lighter veins:
a. Asci clavate, 8-spored Pachypliloeus (p. 74)
b. Asci more or less elliptical, less than 8-spored
Tuber (p. 74)
2. Flesh of the fruit body not veined:
a. Spore ornament a reticulum . . . Hydnobolites (p. 76)
b. Spores warted or spiny:
*Fruit body with a sterile base . . Terfezia (p. 76)
**Fruit body without a sterile base
Choiromyces (p. 76)
If the spores are nearly black and warted or spiny see Elaphomyces (p. 344).

71
Family 1. Pseudotuberaccae

GYROCRATERA P. Hennings

Gyrocratera P. Hennings in Verh. Bot. Ver. Prov. Brandenburg 41:ix


(1899). (PI. IXK.)

There is a single British species:

Gyrocratera ploettneriana P. Hennings, op. cit.:ix (1899). (PI. IXK.)

F r u i t bodies up to 3 m m . diameter, irregular, hollow, opening by


one or more pores or cup-shaped, outer surface fawn to flesh colour or
ochraceous, minutely downy, becoming smooth; inner, hymenial, sur-
face flesh coloured to light red, more or less intricately convoluted;
flesh t h i n and brittle. Asci in a well developed hymenium, cylindrical,
up to 3 6 0 x 3 5 / / ; 4- to 8-spored; spores uniseriate, ellipsoidal or sub-
globose, 2 0 - 3 4 x 1 6 - 2 7 / / , reddish-brown, ornamented with irregular
w a r t s or flanges up t o 5 n high; paraphyses colourless, slender,
scarcely swollen at the tip, m u c h longer t h a n the asci.

Developed 2-3 inches below the surface of sandy soil under conifers. Rare.

HYDNOTRYA Berkeley & Broome

Hydnotrya Berkeley & Broome in Ann. Mag. nat. Hist. 18:78 (1846).
There is a single British species:

Hydnotrya tulasnei Berkeley& Broome, op. cit.:78 (1846). (PI. IXH.)

Fruit b o d y irregularly globose, up to 4 cm. diameter, with a wrin-


kled and pitted surface, reddish-brown, downy; flesh permeated by
labyrinthine cavities communicating with the exterior, pinkish or
russet with a m u s t y odour. Asci in a h y m e n i u m lining the cavities,
clavate, 8-spored, up to 2 3 0 x 7 0 //; spores usually irregularly arranged,
spherical, 20-35 /« diameter, reddish-brown, with large and irregular
w a r t s up to 7// high; paraphyses slender, longer t h a n the asci and
sometimes meeting across the cavities.

Developed in the surface layers of woodland soils, both under conifers


and deciduous trees. Not uncommon.

Family 2. Geneaceae

There is a single British genus:

72
G E N E A Yittadini

Gtenea Vittadini, Monographia T u b e r a c e a r u m : 27 (1831).


There are four British species t h e commonest of which is p r o b a b l y :

Gene a hispidula Berkeley & Broome apud Tulasne, Fungi hypogaei:


121 (1851). (PI. IXD.)
F r u i t b o d y small, scarcely more t h a n 1 cm. across, rounded, wrin-
kled, opening b y an apical pore, dark reddish-brown, drying black,
covered with chestnut-brown hairs, underlying surface somewhat
w a r t e d ; inner surface lining a single large chamber, brown, flesh thin,
whitish. Asci cylindrical, arranged in a h y m e n i u m which is sealed
above b y the fused u p p e r p a r t s of the paraphyses, up to 250x30/«,
8-spored; spores ellipsoidal, 2 8 - 4 0 x 1 9 - 2 8 /«, at first hyaline, becoming
dull yellow, ornamented with low rounded w a r t s or bosses; para-
physes slender, septate, closely crowded and firmly united in a layer
above the asci.

Developed near the surface of humus or under moss, especially in beech


woods. Not uncommon.

Family 3. Eutuberaceae

This central family of the Tuberales contains most of the British


species, distributed in four genera characterised b y t h e veined flesh of
t h e fruit body.

STEPHENSIA Tulasne

Stephensia Tulasne, in G.R. Acad. Sci. Paris 21:1433 (1845).


There is one British species:

Stephensia bombycina (Vittadini) Tulasne, op. cit.: 1433 (1845). (PI.


IXc.)
Fruit body up to 2 cm. across, light brown, externally felted with
short encrusted hairs, flesh cream with straw-coloured veins, smell
strong and unpleasant. Asci cylindrical, up to 270x30/«, 8-spored;
ascospores globose, hyaline, thin-walled, smooth. 19-26/« diameter;
paraphyses slender and colourless.

Developed at or near the soil surface under conifers. Not uncommon.

73
P A C H Y P H L O E U S Tulasne

Pachyphlocus Tulasne in Parlatore, Giornale botanico italiano 1 (2): 60


(1844).
There are three British species, all uncommon or rare.

Pachyphloeus citrinus Berkeley & Broome in Ann. Mag. nat. Hist. 18:
79 (1846). (PI. IXI.)
F r u i t b o d y up to 1.5 cm. across, round or lobed, with an apical
opening, externally warted, yellow, blackening with age; flesh white
with yellow and grey veins, drying grey-brown, smell unpleasant.
Asci clavate, up to 200x50/«, 8-spored; ascospores globose, 13-15/«
diameter, becoming yellowish-brown, ornamented with numerous low
rounded warts.
Developed in humus under trees. Rare.

BALSAM IA Vittadini

Balsainia Vittadini, Monographia T u b e r a c e a r u m : 3 0 (1831).


There are three British species:

Balsamia platyspora Berkeley & Broome in Ann. Mag. n a t . Hist. 13:358


(1844). (PI. IXF.)
F r u i t body 1 - 2 cm. across, irregular, externally covered with chest-
nut-brown w a r t s separated by yellowish furrows; flesh brittle, white,
containing scattered, narrow, sinuous cavities, smell becoming un-
pleasant. Asci lemon-shaped, 60-80x30-40/«, 8-spored; ascospores
broadly elliptical, 19-28x12-16/«, thin-walled, smooth, hyaline.
Developed in humus under deciduous trees, especially Fagus.

B. fragiformis Tulasne, Fungi Hypogaei:125 (1851), is a smaller


species with asci ovoid, not lemon-shaped.
B. vulgaris Vittadini, op. cit.:30 (1831), has cylindrical ascospores
25-40x10-18/«.

TUBER Micheli ex Fries

Tuber Micheli ex Fries, Systema mycologicum 2 (2): 289 (1823).


This is the largest British genus and the one which includes the
well-known edible truffles. The Perigord truffle, T. melanosporum

74
Vittadini, Monographia T u b e r a o e a r u m : 3 6 (1831), is not found in this
country b u t the somewhat similar T. aestivum Vittadini, op. c i t . : 3 8
(1831), occurs not uncommonly in beech woods on calcareous soils in
the south of England and was formerly collected and marketed.
This species m a y be as much as 9 cm. across b u t the m a j o r i t y are
much smaller. Twelve species are recognised in England b u t t h e y are
differentiated with difficulty and for their determination the reader
is referred to the detailed descriptions and comparative table in Dr.
Hawker's monograph cited above. Two of the smaller species re-
presentative of the series with reticulated and spiny spores respective-
ly, will be briefly described here.

Tuber excavatum Vittadini, Monographia T u b e r a c e a r u m : 49 (1831).


(PI. I X A . )
Fruit body subglobose, up to 3 cm. diameter, with a well-defined
basal cavity, externally smooth or finely scurfy, not downy, yellowish;
flesh hard, at first white, becoming purplish-brown, permeated b y
cream-coloured veins radiating from the basal cavity. Asci subglobose
1-6-spored, mostly 4-spored, up to a b o u t 100x90/«; ascospores ellipti-
cal, 22-55 x 16-32/<, golden-brown to dark reddish-brown, wall thick
and ornamented with a deep reticulum with very large meshes. The
fewer the spores in an ascus the larger t h e y tend to be, which accounts
for the very great variation in spore size.

Developed in the upper layer of light soils of beech woods. Not uncommon
in southern England.

Another common species is T. puberulum Berkeley & Broome in


Ann. Mag. nat. Hist. 18:81 (1846), which has downy fruit bodies with
globose reticulated ascospores in 1-3-spored asci.

Tuber rufum Pico ex Fries, Systema mycologicum 2 (2):292 (1823).


(PI. I X B and Fig. I D . )
F r u i t body globose or irregularly lobed, up to 2 cm. diameter, outer
surface finely cracked, foxy red; flesh hard, white with cream veins or
becoming red-brown. Asci clavate, up to 80x70/« with a distinct stalk
20-60/« long in addition; 1-5-spored; ascospores elliptical to fusiform,
17-46 x 14-27 /«, size varying with the n u m b e r of spores per ascus, light
brown, covered with numerous obtuse spines.

Developed in light soil in woods of deciduous trees and conifers. Fairly


common.

75
Family 4. Terfeziaceae

The family includes two British genera, characterised by chambered


fruit bodies in which the flesh is not permeated by veins from t h e sur-
face. An old British record of Terfezia leonis (Tulasne) Tulasne, Fungi
Hypogaei:173 (1851), requires confirmation.

HYDNOBOLITES Tulasne

Hydnobolites Tulasne in Ann. Sci. Nat. Bot. II, 19:378 (1843).


There is a single British species:

Hydnobolites cerebriformis Tulasne, op. cit.:379 (1843). (PI. IXG.)


F r u i t body m u c h lobed and infolded, up to 2 cm. across, outer sur-
face downy, white at first, becoming yellowish; flesh concolorous or
greyish with a few chalky-white streaks. Asci ovoid, up to 100x75//,
mostly 8-spored; ascospores globose, 19-22/« diameter, yellowish,
o r n a m e n t e d with a largemeshed fragile reticulum.

Developed in soil under Fagus. Not common.

CHOIROMYCES Vittadini

Choiromyces Vittadini, Monographia T u b e r a c e a r u m : 50 (1831).


There is only one British species:

Choiromyces meandriformis Vittadini, op. cit.:51 (1831). (Pi. IXJ.)


This is the largest of the British Tuberales, with irregular knobbly
fruit bodies up to 12 cm. across, outer surface yellowish-brown, smooth;
flesh soft, drying hard, white, becoming yellowish and streaked with
numerous irregular w a v y ochraceous lines; smell strong and aromatic.
Asci clavate, up to 180x70/«, 8-spored; ascospores globose, yellow,
16-21 /< diameter, bearing irregular, blunt, often curved spines up to
4 /« long.

Developed on the soil surface in deciduous woods. Rare.

Order 3. TA PHRIN ALES

The Taphrinales are a very isolated and highly specialised group,


perhaps related r a t h e r to the yeasts t h a n to other Ascohymeniales.

76
There is a single family, Taphrinaceae, containing only one large
genus, Taphrina, of which Exoascus, Ascomyces and Magnusiella are
synonyms.

TAPHRINA Fries

Taphrina Fries, Systema Orbis Vegetabilis 1 : 3 1 7 (1825).


The fructification consists of a compact pallisade-like layer of asci
spread over the surface of some living organ of the host plant. In most,
b u t not all species, the ascus is seated on a small basal cell more or less
sunk in the host tissue. There are no paraphyses; the ascospores are
small, globose or elliptical, hyaline, and often bud within the ascus to
give a large n u m b e r of similar, yeast-like, secondary spores. Often in-
fection b y these fungi causes extensive malformation of the host such
as blister-like outgrowth of the leaves, greatly swollen or malformed
carpels, swollen stems or stimulation of lateral b u d s to grow out into
t u f t s of short branches known as "Witches Brooms". Ninety-eight
species of Taphrina are recognised b y Mix, of which 21 occur in the
British Isles. Those described below have been selected to afford some
indication of their effect on the host plants. A full account of t h e
Taphrinae of the world will be found in t h e monograph by Mix in t h e
University of Kansas Science Bulletin 33 (1949), and a useful t r e a t -
m e n t of the British species b y D. M. Henderson in Notes from t h e
Royal Botanic Garden, E d i n b u r g h 21 (1954).

Taphrina populina Fries, Systema mycologicum 3 : 5 2 0 (1832). (PL XB.)

Asci forming a golden-yellow layer over the under surface of blisters,


70-90x18-22/«, with or without a basal cell, packed with numerous
bud-spores about 2 x 1 ju.

P n jeaves of Populus, especially P. nigra and its allies. Common.


The species is better known by its illegitimate synonym T. aurea Fries.

Taphrina johansonii Sadeback in J a h r b . H a m b u r g Wissensch. Anst. 8:


67 (1890). (PL XF.)
Asci forming a golden-yellow layer over t h e surface of swollen car-
pels of Populus tremula and a few other species, clavate, w i t h o u t a
stalk cell but with a long tapering base which is sometimes forked,
60-140x12-27/«, packcd with innumerable bud-spores 4 - 1 0 x 1 . 5 - 4 / « .
LTncommon.

77
T. rhizophora Johanson in Bih. Svenska Yetcnskakad. Handl. 13 (3):
18 (1887), causes similar malformation in Populus alba.

Taphrina amentorum (Sadeback) Rostrup in Vidensk. Meddel. N a t u r h .


Foren K j o b e n h a v n : 2 4 6 ( 1 8 9 0 ) . ( P I . X E . )
Asci covering b o t h surfaces of swollen catkin scales, about 30-50 x
10-20 /«; ascospores 3 - 5 x 4 - 5 / « , often budding in the ascus.
On scales of female catkins of Alnus, infected scales become curved,
tongue-like, outgrowths which whdTT iresfi are light green and more or less
stained or streaked with red so as to be quite conspicuous. Rare, mostly in
the north and west.

Taphrina bctulae (Fuckel) Johansson in Ofvers K. Svensk. Vet.-Akad.


Handl. 4 2 ( 1 ) : 4 0 ( 1 8 8 5 ) . ( P I . X I B . )
Asci cylindrical, 1 7 - 4 6 x 8 - 1 8 / / , with stalk cell 7-17x8-30/«,
8-spored, spores 4 - 6 x 3 . 5 - 5 / « , sometimes budding in the ascus. On
small, circular, pale-green to bright golden yellow spots on leaves of
Betula spp. June to September. Not common.

Taphrina betulina Rostrup i n T i d s s k r . f. Skovbrug 6:199 (1883).


Asci cylindrical, 23-73x10-26/«, with stalk cell 7 - 2 7 x 1 0 - 3 0 /«,
spores soon budding to fill the ascus, 3 . 5 - 6 . 5 x 2 - 5 . 5 /«. On small, rather
pale, leaves of relatively short shoots produced on witches' brooms of
Betula spp. June. (PI. X I D.)
Witches' brooms are exceedingly a b u n d a n t on_^£tida b u t some are
induced b y infestation with mites. Leaves bearing the Taphrina are
soon shed and the fungus-brooms cannot t h e n be distinguished with
certainty, though the mite-brooms bear normal leaves and their shoots
are not negatively geotropic.

Taphrina carpini (Rostrup) Johansson in Ofvers K. Svensk. Vet.


Akad. Handl. 42 (1):41 (1885). (PL X I E . )
Asci cylindrical, 2 0 - 3 0 x 7 - 1 4 //, without a stalk cell, 8-spored, spores
3 . 5 - 5 x 3 - 4 . 5 /«. Covering the surface of slightly deformed pale greenish
leaves on long lax shoots of witches' brooms of Qarpirmsj>etul3s and
Carpinus spp. May to July. Uncommon.

Taphrina tosquinetii (Westendorp) Magnus in Hedwigia 29:25 (1890).


Asci cylindrical, 17-40x7-13/«, stalk cell 7-17x8-18/«,8-spored b u t
often budding in the ascus, spores 2.5-5.5x2.5-5/«. On b o t h surfaces

78
of buckled and deformed leaves or p a r t s of leaves on Alnus glutinosa.
J u n e to September. Very common, by far the most a b u n d a n t of the
three species of Taphrina on this host. (PI. X I c.)

Taphrina potentillae (Farlow) Johanson in Ofvers. Kongl. Svenska


Vetenskakad Forhandl. 42 (1):35 (1885). (PL Xc.)
Asci forming a light yellow layer over swollen stems and thickened
p a r t s of leaves of Potentilla erecta, clavate, w i t h o u t a basal cell, about
2 0 - 5 0 x 7 - 1 5 «, containing about eight ascospores 2-4/« in diameter,
which sometimes b u d in the ascus.
Very common in the wetter districts, especially on plants in ditches and
bogs.

Taphrina pruni Tulasne in Ann. Sci. Nat. Bot. V , 5 : 1 2 9 ( 1 8 6 6 ) . ( P L X D .)

Asci forming a layer over the surface of swollen fruit, " P o c k e t


p l u m s " , cylindric-clavate, with a large stalk cell, a b o u t 2 0 - 5 0 x 5 - 1 5 /«;
ascospores 4 - 7 x 3 - 6 / « , commonly budding freely in t h e ascus.
On young fruit of Prunus domestica and P. spinosa, in June. Uncommon.

Taphrina deformans (Berkeley) Tulasne in Ann. Sci. Nat. Bot. V, 5 : 1 2 8


(1866). (PL XA.)
Asci coating the surface of swollen and convoluted areas of leaves,
cylindric-clavate, a b o u t 20-50x7-15/«, with stalk cells; ascospores
3 - 7 /« diameter, budding in the ascus.
On conspicuous bright red deformed portions of leaves in Prunus persica
and P. amygdalus. Not uncommon.

Taphrina cerasi (Fuckel) Sadebeck in J a h r b . H a m b u r g Wissensch.


Anst. 8 : 8 4 (1890). (PL X I A.)
Asci clavate, 17-53x5-15/«, stalk cell 5 - 2 6 x 4 - 1 2 j x , 8-spored b u t
often budding in the ascus, spores 3 . 5 - 9 x 3 - 6 /«. On bright pink, some-
w h a t buckled and distorted, portions of leaf in Prunus avium, P. cera-
sus &c. Common on P. avium in S.E.England, sometimes associated
with witches' broom formation.

Taphrina athyrii Siemaszko in Arch. Nauk Biolog. Warsaw 1 (14): 17


(1923). (PL XIG.)
Asci 20-30x7-8/«, with stalk cell 5 - 9 x 5 - 7 / « , 8-spored, ascospores
elliptical, 3 - 6 x 2 - 3 / « , sometimes budding in the ascus. On small

79
angular dark-brown lesions in pinnules of Athyrium filix-femina,
covered with a whitish bloom of asci. August and September. Rare or
overlooked.

Taphrina filicina Rostrup ex Johansson in Bibang till K. Vet.-Akad.


Handl. 13 Afd. ILL No. 4:21 (1887). (PI. X I I G . )
Asci 18-46x6-10/«, clavate, without a stalk cell, 8-spored, spores
3.5-6.5x2-3.5 /,. On small yellowish thickened spots in pinnules of
Gryopteris spinulosa. July. Uncommon.

Taphrina vestergrenii Giesenhagen in Bot. Zeit. 59:125 (1901).


Asci 23-50x6-10/«, with stalk cell 10-23x4-10/<, 8-spored, spores
3.5-6x2.5-4/«, often budding in the ascus. On small, irregular, slightly
thickened, yellowish blotches in pinnules of Dryopteris jjlix^Bias and
D. borreri. April to July. Uncommon. ( P I . X I I F . )

Other British species are:


Taphrina sadebackii Johanson in Ofvers Kongl. Svenska Vetensk-
akad. Forhandl. 42 (1):38 (1885), causing small bright yellow spots on
the under side of Alnus glutinosa leaves.
Taphrina ulmi (Fuckel) Johanson, op. cit.:43 (1885), causing yellow
spots on leaves of Ulmus.
Taphrina caerulescens (Montagne & Desmazieres) Tulasne in Ann.
Sci. Nat. Bot. V, 5:127 (1866), causing spots on leaves of Quercus.
Taphrina bullata (Berkeley) Tulasne, op. cit.: 127 (1866), causing
blisters on leaves of Pyrus communis.
Taphrina crataegi Sadeback in Jahrb. Hamburg. Wissensch. Anst. 8:
79 (1890), causing reddish-yellow blisters on leaves of Crataegus but
not to be confused with the much more plentiful marginal inrolling
caused by aphides and mites.
Taphrina padi (Jaczewski) Mix in Trans. Kansas Acad. Sci. 50:77
(1947), causing elongated swollen fruit in Prunus padus.

Order 4. PROTOMYCETALES
There is one family:

Protomycetaceao

The spore sac, variously interpreted as an ascus or a synascus, re-


presenting a number of fused asci, is produced singly by germination

80
of a thickwalled chlamydospore. E a c h sporesac contains numerous,
small, ellipsoid, hyaline spores, which congregate in a ball at the tip
of the sac and are shot out in a mass. A f t e r ejection t h e y copulate in
pairs and germinate to give a mycelium which reinfects t h e host.
There are 3 genera:

I. Chlamydospores scattered irregularly through a somewhat hypertro-


phied portion of host tissue:
A. Chlamydospores smooth-walled, intercalary in the mycelium
Protomyces
B. Chlamydospores finely warted, terminal on hyphal branches
Protomycopsis (p. 82)
II. Chlamydospores forming a single continuous layer beneath the host epi-
dermis Taphridium (p. 82)

PROTOMYCES Unger
Protomyces Unger, Die E x a n t h e m e der Pflanzen 341 (1833).
The mycelium is localised in leaves and stems of herbaceous plants,
in which it induces slight hypertrophy, with formation of inconspicu-
ous galls. There are three very common British species:

Protomyces macrosporus Unger, op. cit. 343 (1833). (PI. XIIB.)

Chlamydospores more or less spherical, 50-70 ,« diameter, with wall


5 n thick, in leaves and stems of Umbelliferae. Ascus subglobose,
similar to the chlamydospore b u t thinwalled, produced in spring from
overwintered chlamydospores, endospores broadly elliptical, 4 . 5 x 3 fi.
Especially common on Mgopodium podagraria and Anthriscus sil-
vestris. Von Biiren in Switzerland records races also on Carum,
Chaerophyllum hirsutum, Cicuta, Daucus, Heracleum, Meum, QZnanthe,
Pastinaca, Peucedanum, Pimpinella, Slum &c.

Protomyces inundatus Dangeard in Le Botaniste 9, 274 (1906).


Chlamydospores in small brownish w a r t s in leaves of Helosciadium
nodiflorum, similar to those of P. macrosporus in size and shape b u t
with wall about 3 fi thick, asci globose, produced at once t h r o u g h o u t
summer and a u t u m n . PI. X I I D.)
Very common and strictly confined to this host. Distinguished from
P. macrosporus mainly by the immediate germination of the summer and
autumn chlamydospores.

Protomyces pachydermus von Thuemen in Hedwigia 13:98 (1874).


Chlamydospores in small often purplish swellings in leaves and
peduncles of Taraxacum officinale, spherical, 30-45 /i diameter, wall

81
4.5 /« thick, germinating only after overwintering. Asci cylindrical,
mostly 100-200x40-60/«. (PI. X I I c . )
Very common.

PROTOMYCOPSIS Magnus
Protomycopsis Magnus Pilzflora von Tyrol 322,1905.
Differs from Protomyces in the chlamydospores being formed singly
a t tips of hyphal branches and having t h e wall finely p u n c t a t e at
m a t u r i t y . The small, round, yellowish leaf-spots induced b y Proto-
mycopsis species m u s t be carefully distinguished from those induced
b y species of t h e Ustilaginaceous genus Entyloma, in which also t h e
host tissue is filled with pale spherical chlamydospores. The vital
difference between t h e two genera depends, of course, on the different
product of chlamydospore germination b u t a useful supplementary
distinction can be found in t h e size of the chlamydospores. It would
be unexpected to find an Entyloma with chlamydospoies more t h a n
25 /« diameter. Resting sporangia of the Phycomycete genera Synchy-
trium and Physoderma m a y also be mistaken for chlamydospores of
Protomycetaceae b u t t h e y lie within the host cell and are not produced
on a septate mycelium. T h e y are commonly embedded in smaller
wart-like galls, often brightly coloured.

Protomycopsis leucanthemi Magnus, op. cit. 323 (1905). (PI. XIIE.)


Chlamydospores 40-50x35-45/«, in small, slightly thickened, yel-
lowish spots in leaves of Chrysanthemum leucanthemum. Asci broadly
clavate, endospores a b o u t 4x3.5/«. J u l y to September. Apparently
rare.
In Switzerland similar lesions in Bellis perennis are caused b y
P. bellidis (Krieger) Magnus and in Leontodon autumnalis b y P. leon-
todontis von Biiren.

TAPHRIDIUM Lagerheim & Juel


Taphridium Lagerheim & Juel in Bihang till K. Sv. Vet. Akad. Handl.
Band 27 Afd. I l l , No. 16:7 (1902).
There is a single British species:

Taphridium umbelliferarum (Rostrup) Lagerheim and Juel, op. cit. 7


(1902). (PI. XIIA.)
Chlamydospores 4 5 - 7 5 x 3 0 - 6 0 / « in a single compact layer beneath
t h e upper epidermis of the host and commonly covering the greater

82
p a r t of a leaf surface to give it a peculiar grey-green appearance, con-
trasting with islands of dark-green healthy lamina. Asci produced at
once, within or p r o t r u d i n g from the chlamydospore, endospores
elliptical, 4 - 6 x 3 - 3 . 5 ft.

Locally abundant on Heracleum sphondylium from late April to June,


recorded elsewhere on Peucedanum palustris.

Order 5. HELOTIALES

This is probably the largest and most diverse order or Ascomycetes


and some of its families as listed here m a y well be worth higher
rank—indeed the Caliciaceae are sharply separated from all other as-
comycetes described in this book b y their peculiar method of asco-
spore dispersal. On t h e other h a n d the family Sclerotiniaceae, re-
cognised here in deference to current trends in systematics, is in fact
not easily separated from the Helotioideae, and it might be wiser to
return to Nannfeldt's earlier and narrower concept of t h e Helotia-
ceae: Ciborioideae, based on anatomical characters of the apothecium.

I. Asci soon disappearing and leaving a mass of loose dry ascospores held
in a cupshaped ascocarp Caliciaceae (p. 104)
II. Asci persistent, ejecting the ascospores through an apical pore:
A. Asci in a pallisade of indefinite extent, without clearly defined
paraphyses, like a thin bloom over patches of dead bark
Ascocorticiaceae (p. 84)
B. Asci in a hymenium, usually with distinctive paraphyses, on a well
defined ascocarp:
a. Ascocarp long-stalked, club-shaped, fan-shaped or pileate,
mainly on the ground Geoglossaceae (p. 84)
/S. Ascocarp ± cup-shaped or immersed in plant tissue, seldom
on the ground:
1. Hymenium waxy owing to close adherence of asci and
paraphyses, the latter usually with knob-shaped tips,
apothecia small, subsessile . . . Orbiliaceae (p. 166)
2. Not as above:
a. Outer tissue of the apothecium (excipulum) soft-
fleshed and composed of subglobose cells, sclerotia
absent Dermateaceae (p. 169)
b. Excipulum formed of elongated or prismatic cells:
*Apothecia beset with characteristic hairs, ex-
cipulum of soft prismatic cells
Hyaloscyphaceae (p. 144)
**Apothecium smooth or merely minutely downy
with simple protuberances from some surface
cells or apothecia arising from sclerotial tissue:

83
t Apothecia arising from sclerotia or stromatised
patches of host tissue Sclerotiniaceae (p. 91)
f t Apothecia not arising from sclerotia, excipulum
usually clearly composed of parallel hyphae
Helotiaceae (p. 105)

Family 1. Ascocorticiaceao
There is a single British genus:

ASCOCORTICIUM Brefeld

Ascocorticium Brefeld Untersuchungen—Mykologie 9:145 (1891).


The only recognised species is:

Ascocorticium anomalum (Ellis and Harkness) Schroeter apud Engler


and P r a n t l Nat. Pflanzenfamilien 1 Abt. 1 : 1 6 1 (1894). (PI. X I r . )
Asci 1 5 - 2 4 x 6 - 8 n , 8-spored, spores 4 - 6 x 2 - 2 . 5 / t , hyaline, elliptical.
Paraphyses have been described b u t seem no more t h a n sparse, un-
differentiated, h y p h a l tips. Forming an evanescent whitish to pinkish
bloom over patches of bark, especially on logs of Qj^us^silvestris b u t
also on T-nnvuTid rinJlupfi August to J a n u a r y . Uncommon. A. albidum
Brefeld is a synonym.

Family 2. Geoglossaceae

The Geoglossaceae include eight British genera in which the fruit


b o d y is relatively large and club-shaped on a long slender stalk. T h e y
correspond morphologically in the Helotiales to the Morchellaceae and
Helvellaceae in the Pezizales and were formerly united with t h e m ,
until the f u n d a m e n t a l distinction between operculate and inoperculate
asci became appreciated. The British species are all terrestrial and of
no economic importance b u t plant pathogenic species of Mitrula are
known in Europe. The genera m a y be distinguished as follows:

I. Fruit body black or very dark purplish-brown, drying black:


A. Conspicuous dark-brown pointed setae present in the hymenium
Trichoglossum (p. 85)
B. No setae in the hymenium:
1. Ascospores brown at maturity . . . . Geoglossum (p. 86)
2. Ascospores permanently hyaline . . . . Corynetes (p. 87)

84
II. Fruit body yellow, brown, olive or green, ascospores permanently hya-
line :
A. Fertile head gelatinous, olivaceous Leotia (p. 87)
B. Head not gelatinous, if olivaceous much more elongated than in
Leotia:
1. Fruit body green or olive-green . . . Microglossum (p. 88)
2. Fruit body yellow or brown:
a. Ascospores 0-1-septate Mitrula (p. 89)
b. Ascospores multiseptate:
*Fertile head flattened, fan-shaped
Spathularia (p. 90)
**Fertile head more or less hemispherical
Cudonia (p. 90)
The ascospores in species of Geoglossum are apt to protrude from the ripe
asci giving the hymenium a downy appearance, liable to be mistaken under
a hand-lens for the setose hymenium of Trichoglossum. The setae of the
latter are at once discernible if a fragment is squashed and examined under
a compound microscope.
A convenient account of all the British species will be found in Prof.
J. A. Nannfeldt's monograph of the Geoglossaceae of Sweden, in Arkiv for
Botanik 30A (4) (1942).

TRICHOGLOSSUM Boudier

Trichoglossum Boudier in Bull. Soc. mycol. France 1:110 (1885).


Fruit bodies tall and slender, stalked, black throughout, velvety;
asci 4- or 8-spored, ascospores cylindric-fusiform, brown, multiseptate,
paraphyses slender, brown at the tip, interspersed with long, stiff,
pointed blackish setae. The species are remarkably uniform in external
appearance and are distinguished b y the n u m b e r of septa in the asco-
spores and the n u m b e r of spores in the ascus.

Trichoglossum hirsutum (Persoon ex Fries) Boudier, Hist. Class.


Discom. d ' E u r o p e : 8 6 (1907). (Fig. 6A.)
F r u i t body variable in shape, usually slender and somewhat
cylindric-fusiform b u t occasionally with a distinctly clavate head, up
to 8 cm. tall, fertile portion thicker t h a n and sharply delimited from
t h e slender velvety stalk. Asci cylindric-clavate, up to 220x20/«, pore
blue with iodine, 8-spored; ascospores lying parallel in the ascus,
cylindric-clavate, brown, normally 15-septate at m a t u r i t y , 100-150x
6-7/«; paraphyses very numerous, slender, septate, slightly enlarged
upwards b u t not m a r k e d l y clavate, apex brown and more or less
curved; setae numerous, thick-walled, almost black and opaque, stiff,
pointed, projecting well beyond the surface of the hymenium.

85
In acid grassland and amongst Sphagnum, August to November. Fairly
common.

Trichoglossum tetrasporum S i n d e n & Fitzpatrick in Mycologia 22:60


(1930), differs in its 4-spored asci.
Trichoglossum walteri (Berkeley) D u r a n d in Annales mycologici 6:
440 (1908), has 7-septate ascospores less t h a n 100 p long.

GEOGLOSSUM Persoon ex Fries

Geoglossum Persoon ex Fries, Systema mycologicum 1:487 (1821).


This, t h e largest genus in the family, is separated from the preceding
b y absence of setae from the h y m e n i u m . There are a t least six British
species, intensive collecting, especially in highland areas, will probably
reveal more. I m p o r t a n t characters are t h e n u m b e r of septa in t h e
ascospores, the rapidity with which these become brown and t h e
n a t u r e of the paraphyses, especially t h e shape of t h e apical cells and
w h e t h e r these are agglutinated or not. P r o b a b l y the commonest
British species is:

Geoglossum cookeianum Nannfeldt in Arkiv for Botanik 30A (4): 22


(1942). (Fig. 6B.)
F r u i t bodies up to 7 cm. high, fertile portion about half the length
or more, somewhat narrowly fusiform, slightly compressed, black,
smooth, d r y ; stalk slender, cylindrical, minutely scurfy or nearly
smooth. Asci up to 180x20/«, 8-spored; ascospores lying parallel in
t h e ascus, cylindricclavate, brown, 7-septate, usually slightly curved,
5 0 - 9 0 x 5 - 7 / « ; paraphyses numerous, slender, slightly longer t h a n the
asci, t e r m i n a t i n g in a short chain of almost globose or barrel-shaped
brown cells up to 8 /< wide.

Common in grassland on sandy soils, especially near the sea. September


to November.

Geoglossum glutinosum Persoon ex Fries, Systema mycologicum 1:


489 (1821), has viscid fruit bodies with ascospores 55-110/« long and
3-7-septate.
Geoglossum fallax Durand in Annales mycologici 6 : 4 2 8 (1908), has
d r y fruit bodies with ascospores 80-100/« long, u p to 12-septate b u t
only slowly turning brown. Its paraphyses are a b r u p t l y swollen up to
10/« at the tips which are almost colourless b u t firmly agglutinated by
a dark brown m a t r i x . (Fig. 6c.)

86
Geoglossum starbaeckii Nannfeldt in Arkiv for Botanik 30A (4): 41
(1942), common on acid Highland soils, has similar b u t shorter asco-
spores, seldom over 85 /« long, with slender curved tips to the para-
physes.
Geoglossum nigritum Cooke, Mycographia: 205 (1879), has uniformly
7-septate ascospores, 50-100 /< long, which t u r n brown relatively early.
One commonly finds two or three species of Geoglossum growing in
the same h a b i t a t and fruiting a t the same time. As t h e y are scarcely
distinguishable to t h e eye mixed collections are therefore often made
and it is necessary to check the determination of each fruit b o d y se-
parately b y microscopic examination.

CORYNETES Hazslinszky
Corynetes Hazslinszky in Mag.Tud. Akad. E r t . ATermes.-Tud. Kor. 11:
1 9 : 8 (1881).
Differs from Geoglossum only in t h e ascospores remaining perma-
nently uncoloured. There are two British species, only one of which is
common:

Corynetes atropurpurcus (Batsch ex Fries) D u r a n d in Annales myco-


logici 6:414 (1908). (Fig. 6D.)
F r u i t bodies growing singly or in small clusters, up to 6 cm. high,
very dark purplish brown to black, fertile portion smooth, narrowly
clavate, seated on a slender, minutely squamulose, cylindrical stalk.
Asci 100-120x10-12/«, 8-spored; ascospores almost cylindrical or
slightly curved, with rounded ends, finally 6-septate, 2 0 - 3 5 x 5 - 6 / « ,
permanently hyaline; paraphyses slender, with swollen hyaline tips
firmly agglutinated b y purplish-brown amorphous m a t t e r .
In acid grassland, often with species of Geoglossum, September to Decem-
ber. Common in western districts.
Corynetes arenarius (Rostrup) D u r a n d in Annales mycologici 6:417
(1908), is a rare species, apparently confined to coastal sand-dunes,
recognised by its dark-brown tipped paraphyses.

LEOTIA Persoon
Leotia Persoon Mycologia E u r o p a e a 1 : 1 9 8 (1822).
F r u i t bodies with a hemispherical or lobed subgelatinous head on a
long slender stalk, ascospores hyaline, multiseptate. There are two
British species:

87
Leotia lubrica Persoon op. cit. 1 : 2 0 1 (1822). (PI. XJ.)

F r u i t bodies often clustered, up to a b o u t 6 cm. high, fertile head


convex with a lobed overhanging margin, yellowish-green to oliva-
ceous, somewhat viscid, flesh gelatinous; stalk cylindrical or tapering,
less t h a n 1 cm. thick, ochraceous, dotted with small greenish granules.
Asci a b o u t 1 5 0 X 1 0 - 1 2 /I, 8-spored, the pore not blued by iodine; asco-
spores slightly fusiform with rounded ends, often slightly curved,
becoming 5 or 7-septate, about 2 0 - 2 5 X 5 - 6 / « , hyaline; paraphyses
slender, branched with clavate tips, hyaline.

In deciduous woods and under Pleridium, especially in wet places, August


to October. Common.

Leotia atrovirens Persoon, op. cit. 1 : 2 0 2 , is a similar fungus, dark


blue-green throughout, found occasionally in d a m p woods in a u t u m n .
The genus is possibly related to Coryne (Helotiaceae).

MICROGLOSSUM Gillet

Microglossum Gillet, Champignons de France, Discomycetes: 25 (1879).


F r u i t bodies similar in shape to the species of Geoglossum b u t with
green or olive-green colouring and hyaline ascospores. There are two
British species.

Microglossum viride (Persoon ex Fries) Gillet, op. cit.:25 (1879). (PI.


XK.)

F r u i t bodies often densely crowded, slender, clavate, up to 5 cm.


high, fertile portion smooth, sharply delimited from the stalk, usually
longitudinally furrowed, green; stalk slender, cylindrical, paler, dis-
tinctly scurfy or scaly. Asci up to about 150x10//, the pore blued b y
iodine, 8-spored; ascospores cylindric-fusiform, about 15-20x5-6/«,
with large oil drops, p e r m a n e n t l y hyaline and only when fully m a t u r e
becoming about 3-septate; paraphyses slender, branched, slightly
thickened at the tips.
In deciduous woods, often amongst moss or attached to debris, September
to November. Common.

Microglossum olivaceum (Persoon ex Fries) Gillet, op. cit.: 25 (1879).


(PI. X M . )
Distinguished from the preceding b y the browner colour of the fer-
tile head and by the smooth glossy stalk.

88
In grassland, on lawns and in deciduous woods, September to November.
Less common than M. viride.

MITRULA Persoon ex Fries

Mitrula Persoon ex Fries, Systema mycologicum 1:491 (1821).


This is a small genus akin to Microglossum b u t with the ascospores
at most 1-septate and with the fertile portions brown or orange in the
three British species. In Scandinavia there occur two parasitic species
which occur on sclerotia in t h e tissues of flowering plants, M. brassicae
H a m m a r l u n d i n A r k i v For Botanik 25A (3): 59 (1932), on Brassica and
M. sclerotiorum (Rostrup) Rostrup in Medd. bot. Foren. 2 : 9 3 (1888),
in underground p a r t s of Trifolium, Lotus and Medicago.

Mitrula paludosa Fries, Systema mycologicum 1 : 4 9 1 (1821). (PI. XG.)

F r u i t bodies solitary or in small clusters, up to about 4 cm. high,


fertile portion club-shaped, smooth, clear orange-yellow, often furrow-
ed towards the base, sharply delimited from t h e more slender pure
white stalk. Asci up to 150x8,«, 8-spored; ascospores biseriate,
cylindrical with rounded ends, hyaline, non-septate, 1 0 - 1 5 x 2 . 5 - 3 / / ;
paraphyses slender, branched.

On rgttjng leaves, especially of Quercus, and sticks in ditches and running


water, also amongst Sphagnum, April to July. Common in suitable situations.

Mitrula abietis Fries, Systema mycologicum 1:492 (1821). (PI. XH.)

F r u i t bodies occurring singly on fallen needles of conifers, up to 3 cm.


high, fertile head short, conical or ovoid, smooth, light yellowish-
brown, seated on a relatively long and very slender, smooth, conco-
lorous stalk. Asci about 5 0 - 7 0 x 6 / / , 8-spored, t h e pore blued b y iodine;
ascospores cylindric-fusiform, hyaline, non-septate, 1 2 - 1 5 x 2 - 3 / « ;
paraphyses slightly enlarged upwards, 3 - 4 n thick, brownish.

On fallen needles of Pinus, Picea etc. September to November. Not un-


common but easily overlooked. Raitviir in Floristic Notes (Tartu) 1 (3)
1961 and Scripta botanica (Tartu) 2, (1962) maintains that M. pusilla Fries
on Pinus needles is distinct from M. abietis on Picea needles.

Mitrula cucullata Fries, Elenchus F u n g o r u m 1 : 2 3 3 (1828), is a


synonym.
Mitrula sclerotipus Boudier in Bull. Soc. bot. France 24:309 (1877),
is a similar slender ochraceous species, up to 2 cm. with ascospores

89
1 0 - 1 3 x 3 / / , borne on small brown sclerotia in wet deciduous woods in
October and November.
Compare also Verpatinia, p. 96.

SPATHULARIA Persoon ex Fries

Spathularia Persoon ex Fries, Systema mycologicum 1:490 (1821).

A small genus easily recognised b y t h e flattened fan-shaped portion


of the fruit body. There is one British species:

Spathularia flavida Persoon ex Fries, Systema mycologicum 1 : 4 9 1


(1921). (PI. XI.)

Fruit bodies solitary, up to 10 cm. high, fertile head flattened, form-


ing a broad flange round the upper p a r t of the stalk, ochraceous, often
slightly u n d u l a t i n g ; stalk slender, tapering at t h e base, m u c h paler
t h a n the head, smooth or mealy. Asci up to 120x14/«, 8-spored, pore
not blued b y iodine; ascospores lying parallel in the ascus, cylindric-
clavate, hyaline, a b o u t 4 0 - 5 0 x 2 - 3 / « , becoming multiseptate; p a r a -
physes slender, branched, hyaline, curled at the tips.

On the groundjn coniferous woods, August to October. Uncommon.

CUDONIA Fries

Cudonia Fries, S u m m a Veg. Scand. Sect. P o s t . : 3 4 8 (1849).

F r u i t bodies resembling Leotia in shape b u t not gelatinous, devoid


of olive t i n t s and with long slender ascospores which bud off swarms
of minute elliptical conidia while still within t h e ascus. There are two
British species, distinguished b y the colour of t h e stalk.

Cudonia circinans (Persoon) Fries, op. cit. 348 (1849). (PI. XL.)

F r u i t bodies solitary or in small clusters, up to 4 cm. high, fertile


head thin, convex and recurved like t h a t of a Helvetia, whitish or
cream, becoming ochraceous; stalk dark grey-brown, downy; flesh
hollow and violaceous in the stalk, white in t h e head. Asci up to
1 5 0 X 1 0 / « , 8-spored, pore not blued b y iodine; ascospores lying parallel
in t h e ascus, cylindric-clavate, 3 0 - 4 5 X 2 / « , becoming multiseptate and
budding in the ascus; paraphyses slender, curled at t h e tip.

On the ground in coniferous woods in the Highlands, August to Septem-


ber. Rare.

90
Cudonia confusa Bresadola, Fungi Tridentini 2 : 6 7 (1892), differs in
having t h e stalk concolorous with the head.

Family 3. Sclerotiniaceae

This family is often united with t h e Helotiaceae from which it m a y


be distinguished by t h e parasitic habit of most of its members and b y
their apothecia arising either directly upon or in association with t h e
sclerotia or from dense aggregations of h y p h a e within the tissues of
the host, often with a blackened surface. The apothecia are often r a t h e r
large, soft-fleshed, long-stalked and frequently have a characteristic
yellowish-brown colour. Their asci tend to be large, broad and thin-
walled and the ascospores are sometimes almost symmetrical about
the short axis, like those of the Pezizales. There are seven British
genera distinguished mainly b y the place of origin of the apothecium.
In m a n y of the parasitic species there is an asexual conidial state and
the apothecia are exceedingly rare or are known only in carefully
m a t e d cultures. These will not be dealt with here b u t a general ac-
count of t h e m will be found in Commonwealth Mycological I n s t i t u t e
Mycological Paper 62.

I. Apothecia arising directly from well developed black sclerotia:


A. Spores colourless:
1. Apothecia cupshaped Sclerotinia (p. 92)
2. Apothecia campanulate "Verpatinia (p. 96)
B. Spores coloured, sclerotia minute, embedded in dung or rotten
bark Martinia (p. 95)
II. Apothecia not arising from sclerotia:
A. Apothecia arising from mummified fruits or catkins:
1. On caryopses of grasses, associated with a slime-spore conidial
state, excipulum of parallel hyphae . . Gloeotinia (p. 98)
2. On other host families, without slime-spores, excipulum formed
of short or subglobose cells:
a. Without macroconidia Ciboria (p. 99)
b. Macroconidia in the form-genus Monilia
Monilinia (p. 96)
B. Apothecia arising from other parts of the host:
1. Minute black microsclerotia present in the host tissue but not
directly associated with the apothecia . . . Stromatinia
2. No microsclerotia present:
a. Apothecia often associated with blackened stromatic
portions of the host or from areas of the substrate delimited
by black lines, ascospores relatively large, elliptical to reni-
form, often becoming septate and/or budding microconidia,
receptacle prosenchymatous (but there is an outer zone of
globose cells in if. americana) . . . Rutstroemia (p. 100)

91
b. Apothecia not associated with stromata, ascospores
small, nonseptate, not budding, outer tissue of re-
ceptacle parenchymatous . . . Ciboriopsis (p. 103)
C. Apothecia arising from blackened stromatic areas in dung, re-
ceptacle prosenchymatous Coprotinia (p. 103)
The known British species of Stromatinia, S. narcissi Drayton & Groves,
is one of those whose apothecia have only been produced artificially and are
not described here. Its microsclerotia are common in the outer papery scales
of Narcissus bulbs. For a full account of this species see the paper by Drayton
& Groves in Mycologia 44 (1952). Two other species, S. rapulum (Bulliard
ex Fries) Boudier and S. paridis Boudier occur in France on rhizomes of
Polygonatum and Paris respectively and may yet be discovered in England.

SCLEROTINIA Fuckel

Sclerotinia Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:330


(1870).
Apothecia arising from well developed tuber- or seed-like sclerotia
with a black smooth rind and white flesh. The sclerotia are formed on
or in tissues of the host b u t these usually decay or break a p a r t before
apothecia develop so t h a t t h e latter are usually found on soil or loose
among plant debris. Care must be t a k e n not to overlook the sclerotia
when making collections, especially when t h e y are buried in soil. Apo-
thecia cup-shaped, sometimes expanding and becoming flat b u t t h e n
usually with a central dimple, some shade of brown. Asci 8-spored;
ascospores hyaline, non-septate. There are about two dozen British
species, m a n y narrowly restricted in host range, and it is likely t h a t
several more remain to be discovered, in association with moulds of
the form-genus Botrytis especially on monocotyledonous hosts. Only
common species whose apothecia are regularly found in nature will be
described here.

Sclerotinia sclerotiorum (Libert) de Bary, Vergl. Morph. Biol. Pilzer: 56


and 236 (1884). (PL X I I I c . )
Apothecia arising singly or in small groups from cushion-like or
short-cylindrical sclerotia which are at first white, finally black, and
formed inside decaying tissues of herbaceous plants or amongst dense
cottony white mycelium on their surface. Apothecia cup-shaped, with
concave, yellowish-brown disc, up to 10 m m . across, on a slender light-
brown, cylindrical, smooth, minutely downy stalk. Asci cylindric-
clavate, up to 130x10/«, 8-spored, pore blued b y iodine; ascospores
uniseriate, elliptical, 9 - 1 3 x 4 - 6 . 5 paraphyses cylindrical, 1.5 / / t h i c k .

92
Common on Helianthus, Daucus and Pkaseolus, on Solanum tuberosum in
wet western districts and associated in the wild with Carduus and Petasites
but parasitic from time to time on a very wide range of hosts. Apothecia
develop from April to July on overwintered sclerotia.

Sclerotinia minor Jagger in J o u r n . Agric. Res. 20:333 (1920), is simi-


lar b u t with very m u c h smaller sclerotia, up to 2 m m . across, developed
on rotting vegetable crops.
Sclerotinia serica Keay in J o u r n a l of B o t a n y 75:132 (1937), occurs
on Caryophyllaceae, with ascospores 14-26x7-17/«.

Sclerotinia trifoliorum Eriksson in K. L a n d t b r a k a d . H a n d b . (1880)


(PI. XIIIB.)
Apothecia arising, usually singly, from irregular black sclerotia up
to 2 0 x 8 mm., commonly embedded in soil. Apothecia brown, concave,
up to 8 m m . across, smooth, seated on a long cylindrical stalk. Asci
200/«, or m o r e x 14/«, 8-spored, pore blued b y iodine; ascospores uni-
seriate, elliptical or oval, 1 2 - 1 8 x 6 - 1 0 / / ; paraphyses cylindrical, en-
larged upwards.

Parasitic especially on leguminous plants such as Trifolium, Vicia, Ono-


brychis, Medicago and Anthyllis but also able to attack Daucus, Pisum and
Lacluca. Apothecia appear from August to November.
There is a variety fabae Keay in Annals of Applied Biology 26:244 (1939),
with ascospores 10-28 x 6.5-15[x, originally described from Faba vulgaris
but also parasitic to Trifolium species.
Some authors regard all the above as mere states of S. sclerotiorum.

Sclerotinia tuberosa ([Hedwig] Fries) Fuckel, op. cit.:331 (1870). (PI.


XIIIA.)
Fructifications arising singly or in groups from large irregular black
sclerotia in rhizomes of Anemone. Apothecia chestnut-brown, cup-
shaped t h e n expanded, up to 3 cm. across, externally paler, smooth,
seated on a cylindrical stalk up to 10 cm. long. Asci cylindric-clavate,
up to 170x10/«, 8-spored, the pore blued b y iodine; ascospores uni-
seriate, ellipsoidal, with a small oil drop at each end, 1 2 - 1 7 x 6 - 9 / « ;
paraphyses cylindrical, slightly enlarged to 3 /< at the tip.

In woods amongst Anemone nemorosa, also on cultivated A. blanda and


A. ranunculoides, March to April. Uncommon.

Sclerotinia candolleana (Leveille) Fuckel, op. cit.:330 (1870). (PI.


XIIIG.)
Fructifications arising singly from small, black, hemispherical or
lenticular sclerotia, 1 - 3 m m . across, amongst rotting leaves of Quercus.

93
Apothecia concave, t h e n flat, up to 4 m m . across, yellowish-brown, ex-
ternally smooth w i t h a slender stalk whose length depends on the
d e p t h of the sclerotium below the surface of the litter. Asci cylindric-
clavate, up to 1 0 0 x 7 / i , 8-spored, t h e pore blued b y iodine; ascospores
uniseriate, elliptical or slightly inequilateral, 6 - 1 0 x 3 - 4 / e , paraphyses
cylindrical, slightly enlarged to 3/j, at the tip.

The fungus causes circular brown spots on leaves of Quercus from July
onwards, sclerotia develop in the fallen leaves and produce apothecia from
May to July the following year. Very common.
The species with small lenticular sclerotia developed in leaves have been
segregated in Ciborinia Whetzel in Mycologia 37:667 (1945). The genus
may be tenable and if so this becomes C. candolleana (Leveille) Whetzel.

Sclerotinia hirtella Boudier, Hist. Class. Discom. d ' E u r o p e : 1 0 7


(1907), is a very similar b u t paler species, with long hairs on t h e stalk
and cup, found on fallen leaves, twigs and bud scales of Castanea in
south-east England. None of the above have macroconidia.

Sclerotinia curroyana (Berkeley) Karsten in Acta Soc. F a u n a Flora


Fennica 2 (6): 123 (1885). (PI. X I I I E . )
Fructifications arising in small clusters from more or less cylindrical
sclerotia embedded in dead stems of Juncus. Apothecia r a t h e r dark
brown, concave t h e n flattened, up to 5 m m . across, externally paler,
smooth, stalk short or almost absent. Asci cylindric-clavate, up to
9 0 x 5 - 6 / « , 8-spored, the pore blued b y iodine; ascospores uniseriate,
narrowly elliptical, straight or slightly curved, 7 - 1 5 x 1 . 5 - 3 / « ; para-
physes cylindrical, slightly enlarged above.

On dead stems in the base of Juncus conglomerates and J. effusus clumps,


March to June. Fairly common. Microcomdial sporodochia develop in living
Juncus stems from July to September and the sclerotia arise in the dead
stems from October onwards.

Sclerotinia sulcata Whetzel in Mycologia 21:15 (1929). (PI. XIIID.)

Fructifications arising singly from fusiform, longitudinally furrowed


sclerotia, up to 2 5 x 3 - 4 m m . b u t varying greatly in size according to
the host, formed inside the culms of Carex spp., from which t h e y are
liberated a t m a t u r i t y . Apothecia fawn coloured, concave, up to 10 m m .
across, externally smooth with a slender, smooth, cylindrical stalk.
Asci cylindric-clavate, up to 180x11/«, 8-spored, t h e pore blued b y
iodine; ascospores uniseriate, ovoid or inequilateral, 9 - 1 7 x 5 - 8 / « ; para-
physes cylindrical, slightly enlarged at the tip.

94
On Carex brizoides, C. disticha, C. gracilis, C. hudsonii, C. inflata, C. nigra,
C. paniculata, C. paradoxa, C. riparia and C. vulpina. Microconidial sporo-
dochia appear scattered over the upper parts of infected culms in late
summer and are followed by the sclerotia, as in S. curreyana. Apothecia
appear from the end of April to June the following year. Common.

Sclerotinia duriaeana (Tulasne) Rehm in Hedwigia 21:66 (1882), is


a very similar species found on Carex disticha, C. brizoides, C. chordor-
rhiza and C. paniculata and distinguished from S. sulcata mainly b y
the microconidial sporodochia which occur as dull black circular spots
1 - 3 m m . across, in paired groups at regular intervals of 5 - 1 5 m m . along
the Carex culms in late summer.
Sclerotinia scirpicola Rehm in Rabenhorst's K r y p t . Flora Deutsch-
land 1(3):822 (1893), forms its apothecia on sclerotia liberated from
t h e culms of Scirpus. The sclerotia are liberated from the dead culms
in winter, fall into t h e water and float until t h e y are blown ashore or
otherwise stranded. Apothecia appear in May.
A species of Sclerotinia very like S. sulcata also occurs in culms of
Eriophorum angustifolium. This is generally identified with S. vahliana
Rostrup in Meddel. om Groenland 3:607 (1891), b u t has been dis-
tinguished as S. dennisii Svrcek in Ceska mykologi 15:37 (1961). The
species on Juncaceae & Cyperaceae have been referred t o Myrio-
sclerotinia Buchwald in Friesia 3 : 2 8 9 (1947).

Sclerotinia fuckeliana (De Bary) Fuckel in J a h r b . Nass. Vereins f.


N a t u r k u n d e 23/24:330 (1870) is the apothecial state of the ubiquitous
gray mould Botrytis cinerea Persoon ex Fries and t h o u g h its apothecia
are seldom seen m n a t u r e t h e y can be produced b y the m a t i n g of
appropriate pure cultures in t h e laboratory. The t y p e host was Vitis
b u t Groves and Loveland in Mycologia 45:415-425 (1953) have
shown t h a t the species is not narrowly host limited and have provided
a detailed description of the apothecia. For related species with
Botrytis conidial states on Caltha and Ranunculus in Europe see Herme_-
bert & Groves in Canadian J o u r n . Bot. 41:341-370 (1963). ""

MARTINIA Whetzel

Martinia Whetzel in Mycologia 34:585 (1942).


The only known species is cosmopolitan b u t seldom collected, per-
haps because of its inconspicuous character. Tropical collections are all
on r o t t e n bark, t e m p e r a t e ones on dung so there m a y be two separable
t a x a passing under the name. However, there are precedents in other

95
groups of fungi for the same species occurring in dung and on wood.
Martinia Whetzel is a later h o m o n y m of Martinia Vaniot, a mono-
typic genus of Compositae, 1903. I have not proposed a valid name
for it here because t h e desirability of distinguishing Helotiaceous
genera on spore colour alone is at least dubious and f u t u r e mono-
graphers m a y well decide t h a t Martinia merits at most sectional rank
in Sclerotinia.

Martinia panamaensis Whetzel, op. cit. 586 (1942). (PI. X I V c . )


Apothecia up to 3 m m . diameter, shallow cupshaped, disc pale
olivaceous to gray, receptacle smooth, reddish-brown, with long slen-
der stalk arising from a subglobose black sclerotium 2 - 3 m m . across.
Asci 30-40x4-5/«, 8-spored, spores biseriate, elliptical, gray-brown,
4-4.5x2/«, paraphyses cylindrical 2/« wide.
Sclerotia in dung. November. Rare.

VERPATINIA Whetzel & D r a y t o n


Yerpatinia Whetzel & D r a y t o n in Mycologia 37:690 (1945).
Apothecium with a long slender stalk bearing a campanulate head
with t h e hymenium covering its outer surface, arising from small,
elongated, black sclerotia formed within t h e tissues of leaves or herba-
ceous stems.

Verpatinia spiraeicola Dennis in Comm. Myc. Inst. Myc. Pap. 62:162


(1956). (PI. XIIIH.)
Sclerotia elliptical, about 2 - 2 . 5 x 1 m m . ; apothecium campanulate,
its surface marked b y a few longitudinal ridges and furrows, with a
pendent margin forming a collar round the tip of the stalk, brown, up
to 2 x 1 m m . ; stalk long, slender, light brown, smooth. Asci cylindric-
clavate, about 35x4/«, 8-spored, the pore blued b y iodine; ascospores
biseriate, elliptic-fusiform, non-septate, 5 - 6 . 5 x 1 . 5 - 2 / « ; paraphyses
scanty, cylindrical, slender.

On rotting leaves ofJFUipendula and stems of Calystegia. Uncommon.


May to July.

MONILINIA Honey
Monilinia Honey in Mycologia 20:153 (1928).
Fructifications developed from mummified fruits, especially fleshy
fruits of Rosaceae and Ericaceae. Apothecia like those of Sclerotinia.

96
Conidial states in the form genus Monilia. A useful account of the
species which cause fruit rot, leaf spots and dieback in economic
plants, will be found in Wormald, " T h e brown rot diseases of fruit
trees", Ministry of Agriculture Bulletin 88. Apothecia of these are
seldom seen.

Monilinia johnsonii (Ellis and E v e r h a r t ) Honey in Amer. J. Bot. 23:


105 (1936). (PI. XIVF.)
Apothecia usually solitary, disc 3 - 8 m m . wide, light brown, re-
ceptacle shallow cupshaped, concolorous, smooth, on a long slender
stalk. Asci 1 5 0 - 1 7 0 x 8 - 1 0 /«, 8-spored, spores elliptical or inequilateral,
10.5-14x5-6,«, paraphyses cylindrical.
Apothecia develop on fallen mummified fruit in late March or April
and shoot ascospores which infect the young leaves to cause large dark
brown or blackish patches, on which is borne a sweet-smelling gray to
buff coloured mould, composed of chains of spherical conidia, 10-13 /«
diameter, separated b y short rod-like disjunctors. These reinfect the
young fruit. On m a n y species of Crataegus, conidia April to J u n e . Very
common in S.E.England b u t apothecia seldom collected because of
their short life.
Sclerotinia crataegi Magnus is a synonym.

Monilinia mespili (Schellenberg) Whetzel in Mycologia 37:673 (1945).


Apothecia very similar to those of M. johnsonii, from which the
species m a y not be t r u l y separable. There is a similar sweet-smelling
conidial state as a whitish mould on very large dark-brown lesions on
leaves of the host. ( P I . X I V N . )
On Mespilus germanicus. Apothecia on mummified fruit in April, conidia
in May. Rare.

Monilinia urnula (Weinmann) Whetzel in Mycologia 37:373 (1945).


Apothecia solitary, receptacle shallow cupshaped, up t o 1 cm. dia-
meter, dark brown, smooth, long-stalked. Asci 190-200x12.5-13.5/«,
8-spored, spores elliptical, uniseriate, 15-18x7.5-9/«, equal in size,
paraphyses cylindrical. ( P I . X I V R . )
On fallen mummified fruit of Var.r. inium. viris-idam. May and June.

Ascospores shot from apothecia on overwintered fruit infect and kill


the shoot tips and a pale buff mould of conidia develops, mainly on
the axis of the dead shoot. The conidia are top-shaped, 2 5 - 3 0 x 1 5 - 1 7 /«,
in short chains.

97
M. megalospora (Woronin) Whetzel is a similar species on V. uligi-
nosum, with ascospores 25-26x17-19/«.
M. oxycocci (Woronin) Honey occurs on V. oxycoccos.

Monilinia baccarum (Schroter) Whetzel in Mycologia 37:672 (1945).


(PL X I I I J . )
Apothecia arising in groups of two or three from mummified berries,
disc concave, t h e n flattened, up to 10 m m . across, light brown, with a
smooth, slender, cylindrical stalk. Asci cylindric-clavate, up to 160x
12 /«, 8-spored, the pore blued b y iodine; ascospores uniseriate, ellipti-
cal b u t usually only four m a t u r e , these 16-18x6-8/«, the other four
a b o u t 1 0 - 1 3 x 4 - 5 / « ; paraphyses cylindrical, 3/« thick.

On Vjmciniu.rn. mi/rliUus.
Conidia occur "on young twigs of the current season's growth and infect
the berries which become mummified and whitened by mycelium. Apo-
thecia develop the following May. Probably widespread in moorland country
but never very abundant. Two other European species, M. ledi (Nawaschin)
Whetzel, op. cit.:673 (1945), and M. rhododendri (Fischer) Whetzel, op. cit.:
673 (1945), are of special biological interest because their annual life cycle
involves two alternate hosts; the conidial state is on species of Vaccinium
in spring and this infects fruit of Ledum and Rhododendron respectively.
From apothecia developed on the overwintered fruits the fungus reinfects
Vaccinium shoots the following spring.

GLOEOTINIA Wilson et al.

Gloeotinia Wilson et al. in Trans. Brit, mycol. Soc. 37:31 (1954).

Apothecia developed on mummified caryopses of grasses, in as-


sociation with a slime-spore conidial state. Structure of the apothecial
flesh resembling t h a t of t h e Helotiaceae, with slenderparallel hyphae
t h r o u g h o u t . Only one species is yet recognised:

Gloeotinia temulenta (Prillieux & Delacroix) Wilson et al., op. cit. 31


(1954). (PL X I I I P . )

Apothecia solitary or in small groups, up to 3.5 m m . diameter, disc


becoming flat, cinnamon, stalk up to 8 m m . long, slender, smooth.
Asci cylindric-clavate, up to 116x7/«, 8-spored, the pore not blued b y
iodine; ascospores uniseriate, ellipsoidal with pointed ends, 7.5-12x
3-6/«, b i g u t t u l a t e ; paraphyses cylindrical, slender.

On fallen caryopses otLolium and other grasses in June. Probably common.

98
CIBORIA Fuckel

Ciboria Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:311 (1870).


Apothecia arising from mummified fruits or catkins, brown, asci 4-
or 8-spored, otherwise as in Sclerotinia. There are eight British species:

Ciboria amentacea (Balbis ex Fries) Fuckel, op. cit.: 311 (1870). (PI.
XIIIL.)
Apothecia occurring singly on fallen male catkins of Alnus and
Salix, disc cup-shaped, light brown, up to 10 m m . broad, externally
smooth, with a long slender stalk. Asci cylindric-clavate, up to 1 3 5 x 9 //,
8-spored, pore blued b y iodine; ascospores uniseriate, elliptical or
slightly inequilateral, 7 . 5 - 1 0 . 5 x 4 . 5 - 6 / t ; paraphyses cylindrical, up to
5 // thick at the tip.
On fallen catkins in March and April. Common.

Ciboria caucus (Rebentisch ex Persoon) Fuckel, op. cit.:311 (1870),


is an almost indistinguishable species on fallen catkins of Populus.
Ciboria acerina Whetzel & Buchwald in Mycologia 28:516 (1936), on
fallen catkins of Myrica, is easily recognised b y its 4-spored asci.

Ciboria batschiana (Zopf) Buchwald in Friesia 3 : 2 5 5 (1947). (PI. XIIII.)

Apothecia arising from blackened cotyledons of fallen acorns, cinna-


mon-brown, concave t h e n flattened, up to 15 m m . across, externally
smooth or slightly downy, on slender stalks. Asci cylindric-clavate, up
to 1 5 0 x 8 / / . 8-spored, the pore blued b y iodine; ascospores uniseriate,
elliptical with slightly pointed ends, 6 - 1 1 x 4 - 6 / < ; paraphyses cylindri-
cal, up to 3.5// thick.

On fallen fruit of Quercus, September to November. Uncommon.

Ciboria aschersoniana (Hennings & Ploettner) Whetzel in Mycologia


37:676 (1945). (PI. X I I I F . )
Apothecia solitary on fallen fruits of Carex spp., disc up to 2 m m .
across, light brown, with a long slender stalk. Asci up to 150x10//,
8-spored, pore not blued b y iodine; ascospores uniseriate, elliptical or
pointed at one end, 8 - 1 2 x 3 . 5 - 5 / « ; paraphyses cylindrical, about 2//
thick at the tip.
On Carex echinata, C. hudsonii, and C. paradoxa, May to June. Probably
common but not easy to find.

99
Ciboria viridifusca(Fuckel) von Hohnel Mitt. bot. I n s t . T e c h n . Hochsch.
Wien 3 : 1 0 1 (1926)
Apothecia scattered, cupshaped, 1 - 4 m m . diameter, grayish to
olivaceous, receptacle dark brown, smooth, or downy at the margin,
with a short slender stalk. Asci 6 0 - 7 0 x 6 - 8 / « , 8-spored, spores uni-
seriate, elliptical, 6 - 9 x 3 - 4 /,, paraphyses cylindrical, obtuse, 2 /t wide.
On fallen female catkins of Alnus. September to November. Common.

Ombrophila baumleri Rehm and Ciboria amenticola (Karsten) Bou-


dier are synonyms. (PI. X I V A . )

Ciboria juncorum Velenovsky, Monogr. Discom. Bohem. 220 (1934).


Apothecia solitary, disc concave, yellowish, up to 2 m m . wide, re-
ceptacle cupshaped, concolorous to cream, smooth, with slender stalk.
Asci 1 1 0 - 1 5 0 x 7 - 1 0 / , , apex t r u n c a t e , 8-spored, spores ovoid, uni-
seriate, 8 - 1 1 x 4 - 5 / / , without guttules, paraphyses cylindrical, 2 / ,
wide, with granular contents. ( P I . X I V T . )
On fallen fruit of Juncus sp. June. Rare.
This species, referred to Ciboria because of its occurrence on a fruit,
is not well placed there because of its prosenchymatous excipulum,
composed of thinwalled parallel hyphae like those of a Hymenoscyphus.
The truncate-tipped asci certainly look more Sclerotiniaceous t h a n
Helotiaceous and it m a y be b e t t e r referred to Rutstroemia b u t I do
not propose a transfer until it is known whether or not any stromatic
tissue is produced in t h e host or in artificial culture.
Ciboria betulae (Woronin) W h i t e in Lloydia 4 : 1 7 1 (1941), is a
minute species on fallen fruit of Betula.
Other species occur in Europe on fallen catkins of Corylus and on
the fruit of Alnus.

RUTSTROEMIA Karsten
Rutstroemia Karsten, Mycologia Fennica 1:12 (1871), emend W h i t e in
Lloydia 4 : 1 6 9 (1941).
Apothecia usually arising from stromatised p a r t s of the host tissue,
either blackened or delimited b y a black line, cup-shaped, w a x y in
t e x t u r e and often r a t h e r large. Asci 8-spored, the ascospores in several
species become septate and m a y bud off secondary spores. The genus
is far from homogeneous and grades into Helotium and Chloroscypha
in the Helotiaceae. There are about ten British species.

100
Rutstroemia firma (Persoon) Karsten, Mycologia Fennica 1:108 (1871).
(PI. X I I I k . )
Apothecia solitary, on decorticated twigs with a blackened stroma-
tic surface, cup-shaped t h e n flattened, up to 12 m m . across, yellowish-
brown, externally smooth or slightly wrinkled, stalked. Asci up to
150x12//, 8-spored, the pore blued b y iodine; ascospores uniseriate,
narrowly elliptical and slightly inequilateral, 14-19x4.0-6.5/«, be-
coming 3-5-septate at m a t u r i t y and budding minute secondary spores,
especially from the terminal cells; paraphyses cylindrical, brown and
up to 3.5 /«thick at the tip.
On fallen _twigs of Quercus^ September to December. Widespread but
scarcely common.

Rutstroemia hercynica (Kirschstein) Dennis in Kew Bulletin 16:317


(1962). (PI. XIVD.)
Apothecia 2 m m . diameter, cupshaped, short-stalked, like a small
R. firma. Asci 110x11/«, spores elliptical to reniform, 13-14x5-6/«,
paraphyses simple, 3 /« wide. Apothecia arise from areas of the sub-
strate delimited b y black lines.
On dead stems of Chamaernerion angustiiolium. September. Uncommon.

Rutstroemia sydowiana (Rehm) W h i t e in Lloydia 4 : 2 0 0 (1941). (PI.


XIIIM.)
Apothecia solitary, erumpent from petioles or main veins of fallen
leaves, disc up to 3 m m . across, yellowish-brown, margin prominent
and t o r n into small pointed red-brown teeth, externally paler, smooth,
longitudinally striate, stalked. Asci up to 130x12 /«, 8-spored, the pore
blued b y iodine; ascospores somewhat kidney-shaped, uniseriate,
1 2 - 1 6 x 5 - 6 . 5 ji., usually with a large central oil drop, not becoming sep-
t a t e ; paraphyses cylindrical, up to 4/« thick at the tip.
On fallen leaves of Quercus, July to September. Not uncommon.
Rutstroemia petiolorum (Roberge) W h i t e in Lloydia 4 : 1 9 7 (1941),
is a very similar species with longer and narrower kidney-shaped asco-
spores, 14-17x4.5-5.5/«, sometimes becoming septate, found on fallen
leaves of Fagus and Quercus b u t much less common t h a n R. sydowiana.

Rutstroemia luteovircscons (Roberge) W h i t e in Lloydia 4 : 2 1 1 (1941).


(PI. X I I I n . )
Apothecia solitary, arising from blackened patches on petioles of
fallen leaves, disc concave, t h e n flattened, pallid greenish-yellow, up

101
3 m m . across, exterior smooth, slightly striate, with a long slender
stalk. Asci cylindric-clavate, up to 150x12//, 8-spored, the pore blued
b y iodine; ascospores uniseriate, elliptical, slightly pointed at each end,
often with two oil drops, 1 2 - 1 5 x 5 - 7 / « ; paraphyses cylindrical, about
4// thick towards t h e tip.
On leaves of Acer, September to October. Not uncommon.
The species is said to occur also on leaves of Tilia.

Rutstroemia conformata (Karsten) Nannfeldt, Fungi Exsic. Suec.: 1174


(1942). (PI. XIIIQ.)
Apothecia erumpent from small blackened areas on the veins of
fallen leaves, disc concave or flat, light brown, up to 2 m m . across,
exterior smooth, with a short stalk. Asci cylindric-clavate, up to
1 1 0 x 6 - 7 / ( , 8-spored, the pore blued by iodine; ascospores uniseriate,
elliptical, slightly pointed at each end, 10-12x5-5.5/«; paraphyses
cylindrical, 4// thick at the tip.
On fallen leaves of Alnus, April to May. Common.

Sclerotinia nervisequa Schroeter, K r y p t . Flor. Schles. 3 (2): 65 (1893),


is a synonym.

Rutstroemia cchinophila (Bulliard ex Merat) von Hohnel in S.B. Akad.


Wiss. Wien 126:340 (1917). (PI. X I I I s . )
Apothecia solitary, arising from blackened areas on old Castanea
husks, disc cup-shaped, flat or slightly convex, up to 7 m m . across,
reddish-brown or occasionally with a purplish t i n t , exterior smooth,
with a slender stalk. Asci cylindric-clavate, up to 130x12 /«, 8-spored,
ascospores uniseriate or irregularly biseriate, narrowly elliptical and
usually somewhat curved, 16-20x4.5-6/«, becoming 3-septate with
age and t h e n budding off minute, spherical, secondary spores; para-
physes cylindrical, 4.5 // thick at the tip, with reddish-brown contents.

On fallen empty cupules of Castanea saliva, September to October. Not


uncommon.

Rutstroemia americana (Durand) White in Lloydia 4 : 1 8 8 (1941).


Apothecia scattered, u p to 5 m m . wide, dark brown, with long
slender stalk, outer surface minutely scurfy. Asci 7 0 - 8 0 x 8 - 9 / / ,
8-spored, spores ellipsoid, nonseptate, 7.5-11x3.5-4.5/«. ( P I . X I V P . )
On old fallen involucres of Castanea saliva. September to November.
Common.

102
Rutstroemia lindaviana (Kirschstein) Dennis E d . 1:65. ( = Sclero-
tinia lindaviana Kirschstein in Verh. Bot. Ver. Brandenburg 48:42
(1906). (PI. X I I I o . )
Apothecia erumpent separately from blackened areas of dead leaves,
disc flat or convex, up to 2 m m . across, yellowish-brown, exterior
smooth, short stalked. Asci cylindric-clavate, up to 40x4/«, 8-spored,
the pore blued b y iodine; ascospores uniseriate or biseriate, elliptical,
4 - 5 x 1 . 5 - 2 / / ; paraphyses cylindrical, 1.5 /t thick.

On dead leaves of Phragmites, less often on Glyceria and Carex, May to


September. Common.
This fungus keys to Rutstroemia but in structure it rather resembles a
Hymenoscyphus

COPROTINIA Whetzel

Coprotinia Whetzel in Farlowia 1:484 (1944).


The type species, on dung in North America, had very longstalked
apothecia with small hyaline ascospores 5 - 6 x 1 . 5 - 2 / « . The British
species referred here because of the development of stromatic tissue in
dung differs in having very short-stalked apothecia with much larger
ascospores. In b o t h the outer tissue of t h e receptacle is composed of
parallel, broad, thinwalled, r a t h e r closely septate h y p h a e and the genus
is separated from Rutstroemia largely on habit. Elliott in Canad.
J o u r n . Bot. 45:521 (1967) considers this an inadequate distinction and
made the combination Rutstroemia cuniculi (Boudier) Elliott.

Coprotinia cuniculi (Boudier) Dennis in Kew Bulletin 16:323 (1962).


Apothecia scattered, 2 m m . diameter, disc light brown, receptacle
concolorous, cup-shaped, smooth, with a short cylindrical stalk. Asci
150x10/«, 8-spored, spores uniseriate or biseriate, elliptical or inequi-
lateral, with two large oil drops, 13-15x4-5/«, sometimes becoming
1-septate, paraphyses cylindrical, 3// wide. ( P I . X I V E . )
On rabbit dropping. March. Uncommon.

CIBORIOPSIS Dennis

Ciboriopsis Dennis in Kew Bulletin 16:319 (1962).


Apothecia small, discoid, with long slender stalk, outer tissue of re-
ceptacle parenchymatous, asci 8-spored, spores small, narrowly ellip-

103
tical to clavate, hyaline, n o n s e p t a t e ; on dead plant remains, without
a sclerotium or distinctly stromatic base. The genus is separated from
Lambertella von Hohnel b y the lack of pigment in t h e ascospores, a
distinction traditionally accepted as of generic value but doubtfully
so in the light of modern experience.

Ciboriopsis bramleyi Dennis, op. cit.: 3 1 9 (1962). (PI. XIVB.)

Apothecia scattered, disc flat, yellowish, scarcely 1 m m diameter,


receptacle light brown, pruinose, stalk long and slender. Asci 45x5,«,
8-spored, spores narrowly ellipsoid, 6 - 8 X 2 - 3 p, paraphyses cylindrical,
1.5 /i wide.

On dead stems and leaves of Chamaenerion angustifolium, Filipendula


ulmaria and Potentilla palustris, the first named is the type host. June to
September. Not uncommon.

C. advenula (Phillips) Dennis is a minute white species on fallen


needles of Larix, with spores 9 - 1 0 x 2 . 5 - 3 , « .

Family 4. Caliciaceae

The members of this family have fructifications with a parallel


h y p h a l structure somewhat resembling t h a t of the Helotiaceae b u t are
readily distinguished b y the evanescent character of their asci, which
break down and completely disappear, leaving a globose head of loose
dry ascospores, held together b y u n d u l a t i n g h y p h a l threads. This t y p e
of apothecium is called a mazaedium, compare also Onygena p. 217.
Most Caliciaceae are more or less intimately associated with unicellular
green algae and are conventionally classed as lichens; an exception is
the genus Roesleria, found on b a r k , especially of roots.

ROESLERIA Thiimen & Passerini

Roesleria Thiimen & Passerini in Oesterreichische Botanische Zeit-


schrift 27:270 (1877).

Apothecia stalked, head cup-shaped, breaking down into a globose


powdery mass of loose spores, asci cylindrical, 8-spored, ascospores
colourless, non-septate. Some mycologists unite Roeslaria w i t h the
lichen genus Coniocybe Ach.

104
Roesleria pallida (Fries) Saccardo in Michelia 2 : 2 9 9 (1881). (PI. XLK.)

Apothecia solitary on bark, head breaking down into a light grey


powdery spore mass scarcely 1 m m . across, on a slender, smooth, light
yellow stalk composed of compact, parallel, thin-walled hyphae about
3 - 4 fi wide, which become loose and radiating in the head. Asci
cylindrical, up to 5 5 x 1 0 n , 8-spored, soon vanishing; ascospores uni-
seriate, broadly elliptical, colourless to light greenish-grey, 5 - 6 x 4 - 5 n,
smooth. There are slender h y p h a e among the asci which have been
called paraphyses b u t there is no compact h y m e n i u m .
On bark, especially of dead roots, often well below soil level, October to
March. Widespread and probably common.
This peculiar fungus has received many names and its true systematic
position is not at all clear. Its asci have been claimed as operculate, in which
case it would be referred to the Sarcoscyphaceae in the neighbourhood of
Pithya. The figure is drawn from the type collection of Roesleria hypogaea
Thiimen & Passerini, op. cit.: 270 (1877), one of the many synonyms of
R. pallida and the one on which the genus was founded.

Family 5. Helotiaceae

This, the largest and most heterogeneous family in the Helotiales,


is characterised b y t h e parallel h y p h a l structure of the outer layers
of t h e flesh or b y a thick flesh of woven h y p h a e covered b y a com-
paratively thin zone of short, cuboid, or globose cells. There is a ten-
dency for the fruit body, or at least t h e disc, to be light or bright
coloured, with cream or yellow t i n t s predominating. Chlorosplenium and
Corynella are green or dark olive, m a n y Ombrophiloideae are p u r p l e ;
black or dark brown members are usually gelatinous, as in Bulgaria
and Bulgariella, subgelatinous as in Chloroscypha, or tough as in
Durella and Scleroderris, not soft and brittle-fleshed like m a n y Derma-
teaceae. The asci are usually 8-spored b u t in Corynella t h e y m a y be-
come packed with rod-like secondary spores and t h e n invite confusion
with Tympanis. In m a n y genera t h e apothecia are distinctly stalked.
Externally t h e y are typically smooth and t h o u g h some members are
minutely downy or scurfy with loose superficial cells t h e y never bear
well differentiated hairs like those of the Hyaloscyphaceae. Where a
downy surface is observed it will be found to be formed b y free un-
specialised tips of t h e surface h y p h a e of stalk or cup. Seven tribes
m a y be distinguished:

I. Apothecium gelatinous Ombrophiloideae


II. Apothecium not gelatinous though the hyphal walls may be thick and
glassy in some tissues:

105
A. Apothecium covered with a layer one cell thick of cuboid dark
brown cells over a flesh of hyaline thick-walled hyphae or else
the ascospores are muriform . . Heterosphaerioideao (p. 141)
B. Ascospores not muriform, apothecium not constructed as above:
1. Surface tissue formed of several layers of cuboid to globose
cells, often becoming loose and mealy, growing on wood or
bark Encoelioideae (p. 136)
2. Surface layers of more or less parallel hyphae:
a. Apothecia small, tough, blackish, smooth, sessile, on
wood or woody stems Durelloideae (p. 141)
b. Apothecia usually more or less stalked, usually light
coloured, disc not conspicuously pruinose:
* Apothecia tough, dark brown, often downy, erum-
pent from bark, ascospores usually elongated and
multiseptate Scleroderridoideae (p. 139)
**Apothecia softer, light-coloured, mostly superficial
or growing on other substrata Helotioideae (p. 113)
c. Apothecia sessile, disc pruinose, asci and ascospores large,
excipular hyphae very slender Polydesmioideae (p. 143)

Tribe Ombrophiloideae

The genera in this tribe all have very soft flesh, either completely
gelatinous or with only thin non-gelatinised tissue. Nine British
genera can be distinguished:

I. Ascospores non-septate:
A. Ascospores hyaline, or at least never dark brown:
1. Apothecia small, solitary, mostly on soil or debris:
a. Gelatinous tissue internal . . . . Ombrophila (p. 107)
b. Gelatinous tissue external, margin sometimes toothed
Sphagnicola (p. 107)
2. Apothecia large, turbinate, clustered on bark
Neobulgaria (p. 108)
B. Ascospores, or four of them, dark brown or blackish:
1. Apothecia large, turbinate Bulgaria (p. I l l )
2. Apothecia small, sessile Bulgariella (p. I l l )
II. Ascospores septate:
A. Apothecia sessile, olivaceous or almost black Corynella (p. 110)
B. Apothecia more or less stalked or turbinate, purple
Coryne (p..109)
III. Primary ascospores simple or septate but soon disappearing and re-
placed by innumerable secondary spores filling the ascus:
A. Secondary spores globose (Compare also Biatorella, p. 215)
Retinocyclus (p. 112)
B. Secondary spores rod-shaped:
1. Apothecia soft-textured, superficial on dead wood
Corynella (p. 110)
2. Apothecia leathery, erumpent usually in clusters from bark
Tympanis (p. 112)

106
For fungi with subgelatinous apothecia on leaves or herbs see Chloro-
scypha, Phaeangellina and Phialea.

OMBROPHILA Fries

Ombrophila Fries, Summa Veg. Scand. Sect. Post.: 357 (1849).


Apothecia superficial, sessile or short-stalked, light coloured, of
small or medium size, soft-fleshed with a least p a r t of the tissue highly
gelatinised, disc convex when fresh, exterior smooth. The gelatinous
tissue is usually in the interior of the flesh. Asci 8-spored, pore blued
by iodine, ascospores elliptical, hyaline, non-septate, paraphyses
simple, cylindrical.

Ombrophila violacea Fries, op. c i t . : 3 5 7 (1849). (PI. XVD.)

Apothecia scattered, obconical, violaceous throughout, soft, disc


flat or convex, up to 4 m m . across, occasionally larger, exterior smooth.
Flesh composed of slender hyaline h y p h a e in a gelatinous matrix,
surface tissue formed of large, elongated, non-gelatinised, thin-walled
cells. Asci cylindric-clavate, up to 80x7-8/«, 8-spored; ascospores uni-
seriate, elliptical, 6 - 1 1 x 3 - 4 / « , hyaline; paraphyses cylindrical, up to
2 /«, thick, colourless.

On decaying leaves on boggy places under trees, August. Rare.

Ombrophila ambigua von Hohnel in Sitz.ber. K. Akad. Wiss. Wien,


Math.-nat. Kl. Abt. I, 127:363 (1918). (PI. XIVQ.)
Apothecia scattered, superficial, disc convex, 1 - 2 m m . broad,
whitish to light gray, receptacle saucershaped, smooth, concolorous,
with a short slender stalk, flesh gelatinous. Asci 4 5 - 5 5 x 5 - 7 / / , pore
blued by iodine, spores biseriate, hyaline, fusiform 9 - 1 6 x 2 - 2 . 5 //, para-
physes slender, 1.5/« wide.
On rotting haulm of Glyceria aquatica. July to September. Uncommon.

SPHAGNICOLA Velenovsky

Sphagnicola Velenovsky, Monographia Discomycetum Bohemiae: 111


(1934).
This is a small genus separated from Ombrophila b y having a flesh
of large thin-walled cells covered b y a gelatinous sheath of slender
hyphae embedded in mucilage.

107
Sphagnicola ciliifera (Karsten) Velenovsky, op. c i t . : I l l (1934). (Fig.
6l.)

Apothecia gregarious, cup-shaped, sessile, watery-white t h r o u g h o u t ,


disc flat, up to 4 m m . across, surrounded b y a narrow erect rim from
which arise narrow pointed t e e t h about y 2 m m . long. In d a m p weather
the outer gelatinous sheath m a y b y readily seen b y the naked eye
surrounding the apothecium. Asci cylindric-clavate, up to 115x8/«,
8-spored; ascospores usually in a single row, elliptical, non-septate,
1 0 - 1 5 x 3 . 5 - 4 . 5 / t ; paraphyses numerous b u t very slender.

On the ground amongst Sghagnum and Carex, especially in pine woods.


Rare.

NEOBULGARIA P e t r a k

Neobulgaria P e t r a k in Annales mycologici 19:42 (1921).


Apothecia superficial, t u r b i n a t e or confluent and convoluted, large,
light coloured, t e x t u r e almost completely gelatinous, asci 8-spored,
ascospores hyaline, unicellular, paraphyses filiform. Two British spe-
cies have been recognised.

Neobulgaria pura (Fries) P e t r a k , op. cit.:45 (1921). (PI. XVB.)


Apothecia 1 - 2 cm. across, t u r b i n a t e in dense clusters, disc slightly
concave becoming flat, pale violaceous or flesh-coloured, exterior
smooth, concolorous. Asci cylindric-clavate, up to 70x9/«, 8-spored,
t h e inner surface of the pore blued b y iodine; ascospores uniseriate,
elliptical, 6 - 9 x 3 - 4 / e , hyaline, containing two small oil drops; para-
physes numerous, filiform, 1 - 2 /« thick.

On fallen trunks, especially of Fagus, June to December. Not uncommon.


The flesh is almost entirely composed of slender colourless hyphae permeat-
ing a gelatinous matrix but a little way beneath the surface is a thin belt
of large, thin-walled, non-gelatinised hyphae. Neobulgaria is scarcely se-
parable from Ombrophila and the two genera may well be reunited in future.
Peziza pura Persoon was "ad truncos abietinos" and presumably some other
species.

Neobulgaria foliacea (Bresadola) Dennis, Comm. Myc. Inst. Myc. Pap.


62:166 (1956). (PI. XVc.)
Fructification extensive, convex, convoluted,resembling the basidio-
mycete genus Tremella, raisin-coloured to dull reddish-purple, very
soft and gelatinous, drying t h i n and horny. Asci up to 100x8/«,

108
cylindric-clavate, 8-spored, the pore blued b y iodine; ascospores uni-
seriate, elliptical, 6 - 1 0 x 4 / * , hyaline, containing two oil drops; para-
physes filiform, 1 - 2 /, thick.

On fallen trunks, August to December. Rare.


In microscopic characters this is indistinguishable from the preceeding
and there remains a possibility that N. foliacea is no more than an over-
developed state of N. pura. More collections and especially field observations
are needed to clarify the relationship between these two species and it is
largely in the hope of encouraging such studies that N. foliacea is figured
here.

CORYNE Tulasne

Coryne Tulasne, Selecta F u n g o r u m carpologia 3:190 (1865).*


Apothecia clustered, of medium size, gelatinous, bright purple, asci
8-spored, ascospores becoming septate, hyaline. There are two British
species, easily distinguished b y their ascospores. F r o m the anatomical
standpoint Coryne is an Encoelia with gelatinised flesh.

Coryne sarcoides (Jacquin ex S. F. Gray) Tulasne, op. cit.: 190 (1865).


(PL X V A . )
Apothecia superficial, usually in clusters, gelatinous, cup-shaped,
sessile or short-stalked, disc concave or flattened, often irregular and
lobed, up to 10 m m . across, reddish-purple, exterior usually minutely
scurfy, concolorous. The flesh is composed of loosely woven h y p h a e in
a gelatinous m a t r i x , with a surface tissue formed of non-gelatinised
thin-walled globose cells. Asci cylindric-clavate, up to 1 6 0 x 1 0 / , ,
8-spored, the pore slightly blued by iodine; ascospores often biseriate,
elliptical to inequilateral, 1 0 - 1 9 x 3 - 5 / , , at first non-septate with two
oil drops, ultimately 1-septate, occasionally 3-septate, sometimes
germinating in the ascus; paraphyses slender, often a b r u p t l y swollen
to 4/< thick at the tip. PL XVA, spore on the left.

On stumps and fallen logs, especially of Fagus, August to December.


Common.
The apothecia are almost invariably accompanied by a similarly coloured
conidial state which has been called Pirobasidium sarcoides von Hohnel in
S.B. Akad. Wiss. Wien 111:1002 (1902).

*The name Coryne applied originally to a conidial fungus. Groves & Wil-
son in Taxon 16:35-41 (1967) have proposed the name Ascocoryne to re-
place Coryne Tulasne, with type species A. sarcoides (Jacquin ex Gray)
Groves & Wilson.

109
Coryne cylichnium (Tulasne) Boudier, Hist. Class. Discom. d ' E u r o p e :
97 (1907), is very similar b u t often larger and is distinguished b y its
long ascospores, 1 8 - 3 0 x 4 - 6 //, which become multiseptate and bud off
spherical secondary spores while still within the ascus. PI. XVA, spore
on the right.
Coryne turficola Boudier in Bull. Soc. mycol. France 21:71 (1905)
with non-septate ascospores 15-18x5.5-6.5//, found on p e a t y soil
amongst Sphagnum has large purple apothecia with olive-brown disc
and tapering rooting base.

CORYNELLA Boudier

Corynella Boudier in Bull. Soc. mycol. France 1:114 (1885).


Apothecia sessile, gelatinous, greenish, dark olivaceous or nearly
black, asci at first 8-spored, ascospores hyaline, multiseptate, some-
times budding very freely in the ascus.

Corynella prasinula (Karsten) Boudier, Hist. Class. Discom. d ' E u r o p e :


99 (1907). (PL XVE.)
Apothecia sessile, gelatinous, convex, up to 400 fi across, emerald
green, smooth, usually associated with a slender erect greenish conidial
state called Dendrostilbella prasinula von Hohnel in Oest. Bot. Zeits.
5 5 : 2 2 (1905). Asci clavate, up to 90x8/«, rounded above, the pore not
blued b y iodine, 8-spored; ascospores irregularly biseriate, cylindric-
fusiform, 1 0 - 1 3 x 3 / / , 3-septate; paraphyses very slender, repeatedly
forked towards the tip.

On decaying wood, June to October. Fairly common.

Corynella atrovirens (Persoon) Boudier, Hist. Class. Discom. d ' E u r o p e :


99 (1907). (PI. XVF.)
Apothecia solitary or in small groups, sessile, dark olive-green to
nearly black, drying black, lenticular, smooth, gelatinous. Asci clavate,
up to 120x12 fi, rounded at the tip, the pore not blued by iodine, at
first 8-spored; ascospores biseriate, cylindric-fusiform or curved, up
to 7-septate or even more, budding off inumerable rod-shaped se-
condary spores about 2 - 3 x 1 / / which ultimately fill the ascus; para-
physes very slender, forked above and swollen to 2 // thick at the tip.

On rotting wood, especially in spring. Common.

110
BULGARIA Fries

Bulgaria Fries, Systema mycologicum 2:166 (1822).


Apothecia large, t u r b i n a t e , gregarious, gelatinous, black, ascospores,
or some of them, dark brown. There is only one British species:

Bulgaria inquinans Fries, Systema mycologicum 2:167 (1822). (PI.


XVH.)

Apothecia erumpent, often clustered b u t not becoming confluent,


t u r b i n a t e , gelatinous, disc slightly concave, up to 4 cm. across, smooth,
black, exterior sooty brown, scurfy. Asci cylindric-clavate, 8-spored,
up to 200x9/«, the pore blued by iodine; ascospores of two kinds, the
upper four somewhat kidney-shaped, 11-14x6-7/«, very dark brown
and opaque, the lower four similarly shaped b u t hyaline, and smaller,
containing several small oil drops; paraphyses very slender, with
slightly swollen brown tips.

On fallen trunks and branches, especially of Quercus, occasionally also of


Betula, Castanea, Carpinus and Ulmus. Common.
Amateurs often confuse B. inquinans with the equally common gelatin-
ous black basidiomycete Exidia glandulosa, which also favours branches of
Quercus. The apothecia of Bulgaria have smooth discs, free from the minute
pimples that give the name to E. glandulosa. If laid overnight on a sheet of
paper the Bulgaria fruit bodies will be found next morning surrounded by
a sooty zone of extruded ascospores; no such reaction is possible with
Exidia, which has hyaline spores.

BULGARIELLA Karsten

Bulgariella Karsten in Acta Soc. F a u n a Flora Fennica 2 (6): 142 (1885).


Apothecia small, superficial, solitary, lenticular, blackish, gelatinous,
asci 8-spored; ascospores elliptical, all dark brown. There is one British
species:

Bulgariella pulla (Fries) Karsten, op. cit.: 142 (1885). (PI. X V G . )

Apothecia lenticular, sessile on a small base, gelatinous, black, or


very dark olive-green, up to 3 m m . across. Asci cylindric-clavate, up
to 150x10/«, 8-spored, the pore not blued b y iodine; ascospores uni-
seriate, broadly elliptical, 10-15x6.5-8.5/«, dark sooty brown; para-
physes slender, simple, about 2 /« thick, with dark granular contents.

On rotting decorticated wood, especially of Betula, September to Oc-


tober. Rare.

Ill
The apothecia are indistinguishable in the field from those of Corynella
alrovirens.

RETIN 0 CYCLUS Fuckel

Retinocyclus Fuckel in J a h r b . Nass. Vereins N a t u r k . 25/26:332 (1871).


Two species are recognised by Groves and Wells in Mycologia 48:
865-871 (1956), one of which is as yet recorded in t h e British Isles.

Retinocyclus abietis (Crouan) Groves and Wells, op. cit.: 869 (1956).
Apothecia scattered, discoid, sessile, disc flat or convex, black,
y 2 - 1 m m . wide, receptacle fleshy, outer tissue of parallel h y p h a e with
strongly gelatinized walls, at right angles to t h e surface. Asci 4 5 - 6 0 x
12-15 /x, spores 2 - 3 ji diameter, paraphyses cylindrical, 2 - 3 n wide,
tips united in a brownish gelatinous m a t r i x . (Fig. 29B.)
On resin of Pinus silvestris and other conifers. August. Uncommon.

TYMPANIS Tode ex Fries

Tympanis Tode ex Fries, Systema mycologicum 2:172 (1822).


Apothecia e r u m p e n t , often in dense clusters, cup-shaped, subsessile,
black, smooth or pruinose, tough fleshed, composed of interwoven
brownish hyphae, 2.5-3 n wide, with gelatinised walls, asci at first con-
taining eight p r i m a r y ascospores which are evanescent, being replaced
b y innumerable minute rod-shaped secondary spores; paraphyses
slender, their tips cohering above t h e asci. Conidial states occur, con-
taining conidia similar to the secondary ascospores. The species are
m u c h alike and have been greatly confused so t h a t t h e n u m b e r occur-
ring in t h e British Isles is uncertain. For a thorough unravelling of
this most difficult genus see a p a p e r b y J . W. Groves in Canadian
J o u r n a l of B o t a n y 30:571-651 (1952). Its t r u e affinities m a y lie with
Pragmopora and Durandiella in Dermateaceae for in some species t h e
ephemeral primary ascospores are elongate-fusoid and multiseptate.
The best known British species is:

Tympanis conspersa Fries, op. cit.: 175 (1822). (Fig. 9A.)


Apothecia e r u m p e n t in dense clusters from cracks in the bark, cup-
shaped, often distorted b y m u t u a l pressure, subsessile, up to 0.8 m m .
across, black b u t usually more or less covered with a whitish pruina,
disc flat with a thick inrolled margin. Asci cylindric, r a t h e r thick-

112
walled, up 180x22//, with eight p r i m a r y ascospores, uniseriate,
broadly elliptical, 4 - 8 x 3 - 6 //, non-septate; secondary ascospores filling
the ascus, rod-shaped or slightly curved, hyaline, 2 - 4 x 1 - 1 . 5 / / ; p a r a -
physes hyaline, septate, simple or branched, slender, their tips slightly
swollen and firmly united b y a brownish gelatinous matrix.

On dead twigs of Pyrus, Sorbus and Prunus, especially on Pyrus malus


and Sortyus__aucuparia. February to June, occasionally in autumn. Fairly
common.

Tympanis alnea (Persoon) Fries, op. cit.: 174 (1822), is a very similar
fungus on Alnus and Betula.
Tympanis ligustri Tulasne, Selecta F u n g o r u m Carpologia 3 : 1 5 4
(1865), occurs on Ligustrum.
Tympanis laricina (Fuckel) Saccardo, Sylloge F u n g o r u m 8 : 5 8 3
(1889), occurs on Larix, numerous other species are known on various
conifers in Europe.

Tribe Helotioideae
Several of the genera in this tribe are still ill-defined. Helotium as
formerly interpreted was so widely conceived as to be almost indefinable
so t h a t the divisions between it and Discinella on the one h a n d and
Pezizella on the other were difficult to draw. Many authors consider
Calycella also inseparable from Helotium. The following artificial key
is applicable to the British species:

I. Ascospores more than 1-septate; or at least very long and slender:


A. Apothecia stalked, on grasses, herbs, etc. Belonioscypha (p. 127)
B. Apothecia sessile or nearly so, on other substrata:
1. On dead leaves of deciduous trees, ascospores filiform
Pocillum (p. 114)
2. On other substrata, ascospores clavate or long fusiform:
a. Apothecia associated with Dematiaceous mould on wood
Strossmayeria (p. 116)
b. Apothecia not associated with a mould:
*Fructifications on pine needles and cones
Pseudohelotium (p. 115)
**Fructiflcations on wood Gorgoniceps (p. 115)
II. Ascospores non-septate, occasionally 1-septate:
A. Apothecia sessile:
1. Fructifications cream or yellow, on wood:
a. Outer tissue of receptacle formed of undulating hyphae
at a high angle to the surface Calycella (p. 118)
b. Outer tissue or receptacle formed of thinwalled isodia-
metric cells Phaeohelotium (p. 118)
2. Fructifications developed on soil, very soft Discinella (p. 135)

113
3. Fructifications dark purplish, on Empetrum leaves
Phaeangellina (p. 116)
4. Fructifications dark olive-green to black, on conifer leaves
Chloroscypha (p. 117)
5. Fructifications on other substrata, not coloured as 3 or 4
Hymenoscyphus (p. 121)
B. Apothecia stalked:
1. Apothecia green or olive-green Chlorosplenium (p. 133)
2. Apothecia pink, with a deep marginal collar, on Equisetum
Stamnaria (p. 127)
3. Apothecia other colours or lacking the marginal collar:
a. Surface hyphae broad, thin-walled, sometimes short-
celled, lying at an angle to the surface, cup seldom
downy:
* Ascus pore not blued by iodine, disc usually convex,
white to lilaceous, on woody substrata or debris
of marsh plants Cudoniella (p. 119)
**Ascus pore blued by iodine, disc usually concave,
white to yellow, often on herbaceous plants or
fallen leaves Hymenoscyphus (p. 121)
b. Surface hyphae nearly parallel with the surface or the
cup covered with minute downy hairs:
*Surface hyphae with thickened subgelatinised walls,
outer tissue appearing glassy in section:
-fMargin of the cup toothed Cyathicula (p. 128)
+ +Margin of the cup even, paraphyses often
lanceolate Phialea (p. 128)
**Surface hyphae with normal walls, often running out
into short downy hairs, ascospores usually narrow
and slightly curved Pezizella (p. 130)

POCILUJM de Notaris

Pocillum de Notaris in Commentario della Societa Crittogamologica


Italiana 1 : 3 6 1 (1864).
Apothecia sessile, goblet-shaped, outer tissues formed of parallel
brown hyphae, asci cylindric-clavate, very long, 8-spored; ascospores
filiform like those of t h e Ostropales; paraphyses slender, simple.

Pocillumcesatii (Montagne) de Notaris, op. cit. : 3 6 1 (1864). (PI. XXIJ.)

Apothecia scattered, goblet-shaped, smooth, dark brown below with


a pale margin, about 0.5 m m . high and not as broad, sessile. Asci
cylindric-clavate, u p to 1 5 0 x 7 n, apex rounded, pore not blued b y
iodine; ascospores filiform, u p to 140 x 1 fj. hyaline, lying parallel in the
ascus; paraphyses slender, swollen at the tip.

114
On fallen leaves of Quercus in spring. Rare, the British record needs con-
firmation but this minute fungus is very likely to be overlooked.
Other British fungi sometimes referred to Pocillum obviously belong in
other genera.

PSEUD OHELOTIUM Fuckel

Pseudohelotium Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:


298 (1870).
Apothecia minute sessile or nearly so, t e x t u r e of parallel hyphae,
asci 8-spored; spores needle-shaped, multiseptate. The general aspect
is t h a t of a Pezizella with acicular ascospores.

Pseudohelotium pineti (Batsch ex Fries) Fuckel, op. cit.:298 (1870)


(PI. X X I G . )

Apothecia scattered, superficial, obconical with a small base, disc


flat, whitish to buff, up to 1 m m . across, exterior smooth, brown below,
whitish at the margin which is composed of hyaline, inflated, h y p h a l
tips. Asci cylindric-clavate, up to 7 0 x 9 /«, t h e small pore blued in io-
dine, 8-spored; ascospores narrowly cylindrical or slightly clavate,
20-30x1.5-2/«, 3-septate, lying more or less parallel in t h e ascus;
paraphyses cylindric-clavate, up to 3 /« thick.
On fallen rotting needles and cones of Pinus, in autumn. Common.

GORGONICEPS Karsten

Gorgoniceps Karsten, Mycologia Fennica 1 : 1 5 (1871).


Apothecia sessile or nearly so, obconical, smooth, external tissue
composed of parallel brown hyphae, asci cylindric-clavate, the pore
blued b y iodine, 8-spored; ascospores long and slender, multiseptate,
hyaline; paraphyses slender.

Gorgoniceps aridula (Karsten) Karsten in Not. Sallsk. F a u n a Flora


Fennica 11:247 (1870). (PI. X X I V K . )
Apothecia scattered, superficial, sessile, obconical on a small base,
disc flat, pallid, about 0.5 m m . across, exterior smooth, grey-brown
below, paler at t h e margin. Asci cylindric-clavate, up 130x14/«,
8-spored, conical at t h e tip, the small pore blued b y iodine; ascospores
long and slender, 50-88x2-2.5/«, curved, hyaline, becoming multi-
s e p t a t e ; paraphyses slender, up to 3 n thick.

115
On wood or bark, especially of Pinus, in autumn. Rare, figured from
Karsten's material.
It will be seen that there is little difference between this and Pseudohelo-
lium apart from the number of septa in the ascospores; if they prove to be
identical the name Pseudohelotium has priority.

STROSSMAYERIA Schulzer
Strossmayeria Schulzer von Miiggenburg in Oesterreichische Botani-
sche Zeitschrift 31:313 (1881).
Apothecia minute, sessile, structurally like those of a Durella b u t
soft, light coloured and intimately associated with Dematiaceous
moulds on decorticated wood.

Strossmayeria basitricha (Saccardo) E d . 1 : 7 3 (Fig. 20E.)


Belonidium basitrichum Saccardo in Atti Soc. Venet.-Trent. Sci. Nat.
P a d o v a 4:135 (1875).
Apothecia scattered or in small coherent groups, saucershaped,
sessile or substipitate, whitish, drying amber coloured, disc flat,
0.5 m m . across, receptacle smooth, composed of parallel thinwalled
hyphae 2.5-3/« wide, stained blue by Melzer's reagent. Asci clavate,
100-120x 10-14/.t, 8-spored; ascospores 2 - 3 seriate, fusiform, up to
7-septate and slightly constricted at the septa, 30-40x4-5/«, hyaline;
paraphyses filiform, hyaline.
On decorticated wood of Quercus and Fagus in association with mycelium
of Helminthosporum simplex Kuntze ex Fries.
This is the type species of the genus Leptobelonium von Hohnel in Sitz.
Akad. Wiss. Wien, Mathem.-Nat. Kl. Abt. 1,132:112 (1923) but von Hohnel
unaccountably adopted for it the name Leptobelonium helminthieola von
Hohnel, derived from a manuscript name proposed by Bloxam on a speci-
men in the Berkeley herbarium. Inspection of Batsch's figure suggests that
von Hohnel was right in rejecting any name based on Peziza minutissima
Batsch but there were several validly published names already available
for the species, the earliest of which appears to be B. basitrichum Saccardo.
According to Nannfeldt the type species of Strossmayeria, S. rackii Schulzer
von Miiggenburg op. cit.: 314 (1881) is probably another synonym, though
it was imperfectly described as having elliptical 3-septate spores with a
curious 3-septate cylindrical appendage.

PHAEANGELLINA Dennis
Phaeangellina Dennis in Kew Bulletin.:360 (1955).
Apothecia minute, sessile, subgelatinous, purplish-brown, asci
8-spored, ascospores elliptical, brown, becoming septate. There is one
species:

116
Phaeangellina empetri (Phillips) Dennis, op. cit.:361. (PI. XVJ.)
Apothecia scattered, erumpent, sessile, t u r b i n a t e , disc flat, scarcely
0.5 m m . across, dark olive, exterior smooth, dark purplish-brown,
sometimes vertically ridged and furrowed. Asci cylindric-clavate, up
to 160x20/«, r a t h e r thick-walled, rounded above, not blued b y iodine,
8-spored; ascospores uniseriate, broadly elliptical, 1 7 - 2 1 x 9 - 1 2 /i, with
numerous oil drops, long remaining hyaline t h e n t u r n i n g dark brown,
becoming finally 1-septate, especially after expulsion from the ascus;
paraphyses slender, more or less branched.

On dead leaves of Empetrum, still attached to the plant, August to Sep-


tember. Common. In th'e" field this minute species may be confused with
Sphaeropezia empetri, see p. 212, which is accordingly figured beside it on
PI. XV.

CHLOROSCYPHA Seaver

Chloroscypha Seaver in Mycologia 23:248 (1931).


Apothecia minute, sessile, subgelatinous, yellowish-green to blackish,
on leaves of conifers, asci 8-spored, ascospores broadly elliptical or fusi-
form, p e r m a n e n t l y hyaline and non-septate. Two British species are
known.

Chloroscypha sabinae (Fuckel) Dennis in Kew Bulletin: 410 (1954).


(Fig. 1 8 A . )
Apothecia minute, t u r b i n a t e , sessile on a small base, subgelatinous,
black throughout, disc flat, less t h a n 0.5 m m . across, exterior smooth.
Asci clavate, up to 150x20//, 8-spored, rounded above, r a t h e r thick-
walled, pore reaction inconstant, sometimes blued b y iodine; asco-
spores biseriate, elliptic-fusiform, 1 5 - 2 1 x 8 - 9 f i , hyaline, non-septate;
paraphyses slender, branched, up to 4 /« thick at t h e tips, which are
united by the brownish mucilaginous m a t r i x .

On dead leaves of Ju^^erus_j^mmjjjiis, J. nanus and J. sabina, probably


throughout the year, perhaps common but generally overlooked. Infected
leaves may bear over a dozen apothecia, scattered or in small clusters, on
the concave surface of the needle.

Kriegeria juniperina (Ellis) Seaver in Mycologia 35:493 (1943), is a


synonym.
Chloroscypha seaveri Seaver in Mycologia 23:249 (1931), occurs on
leaves of Thuya and Cupressus lawsoniana.

117
PHAEOHELOTIUM Kanouse

Phacohelotium Kanouse in Papers Michigan Academy Science, Arts,


Letters 20:75 (1935).
Apothecia superficially like those of Calycella b u t with the outer
tissue of the receptacle composed of r a t h e r large-celled, thinwalled,
pseudoparenchyma. The brown colouration of the ascospores, for
which the genus is named, develops tardily, perhaps even after dis-
charge from t h e ascus, and is probably not an essential character.

Phacohelotium monticola (Berkeley) Dennis in Persoonia 3 : 5 4 (1964).


Apothecia solitary or in clusters, disc flat or convex, yellow, 1 - 2 m m .
wide, receptacle sessile, t u r b i n a t e to cupulate, smooth, yellow, soft-
fleshed. Asci 9 5 - 1 0 0 x 7 - 1 0 / / , 8-spored, spores biseriate, elliptic-cylin-
dric, 1 2 - 1 8 x 4 - 5 //, ultimately 1-septate with brown slightly roughened
walls, paraphyses cylindrical 2/i wide. (PI. XIVo.)

On decorticated wood. September and October. Apparently uncommon


but easily mistaken for Calycella citrina.
P. flavum Kanouse is a synonym.

Phacohelotium lilaeinum (Bresadola) Dennis comb.nov.

Pezicula lilacina Bresadola Fungi Tridentini 1 : 6 8 (1884).


Apothecia solitary, disc up to 4 m m . wide, flat, very pale lilac-gray,
receptacle t u r b i n a t e , concolorous, smooth, soft-fleshed. Asci 125x
10-12/t, pore outlined blue b y iodine, 8-spored, spores 1-2-seriate,
14-20x5-8/«, hyaline, nonseptate. (Fig. 29j.)
On wet rotten wood. October.
In spite of its permanently hyaline ascospores the anatomy indicates
Phaeohelotium as an appropriate genus for this species and for several
others referred by Boudier and his followers to Pachydisca Boudier, eg.
P. nobilis (Velenovsky) Le Gal, P. umbilicata Le Gal. Pachydisca was based
on Helotium guernisaci Crouan, which Madame Le Gal has shown to be a
Rutstroemia.

CALYCELLA Boudier

Calycella Boudier in Bull. Soc. mycol. France 1 : 1 1 2 (1885).


Apothecia sessile or nearly so, smooth, tough, the outer tissue of t h e
cup composed of r a t h e r thick-walled h y p h a e undulating and some-
w h a t interwoven b u t in general lying more or less parallel at a high

118
angle to t h e surface, whitish or yellow, asci 8-spored, ascospores often
becoming 1-septate. There are about four British species, of which t h e
following are the best k n o w n :

Calycella citrina ([Hedwig] Fries) Boudier, op. cit.: 112 (1885). (PI.
XVIA.)

Apothecia gregarious, on wood or e r u m p e n t from bark, shallow cup-


shaped, narrowed below to a small base, disc slightly concave, deep
yellow, drying orange-yellow, up 3 m m . across, exterior smooth. Asci
up to 135x10,«, cylindric-clavate, 8-spored, pore not distinctly blued
b y iodine; ascospores biseriate, elliptical, with oil drops at each end,
9 - 1 4 x 3 - 5 / < , often becoming 1-septate; paraphyses slender, slightly
enlarged upwards and containing yellow oil drops.

On dead branches of Alnus, Corylus, Fagus, Quercus etc., September to


December. Very commoiTTyuTTtiTho be~co'nfused with Hymenoscyphus caly-
culus.

Calycella sulfurina (Quelet) Boudier, op. cit.: 112 (1885). (PL XVIB.)

Apothecia gregarious, shallow cup-shaped, about 1 m m . across, disc


flat with a low rim, bright sulphur-yellow, exterior paler and minutely
downy, sessile, sometimes two or three together on a minute stromatic
base. Asci cylindric-clavate, up to 70x4,«, 8-spored, pore not blued by
iodine; ascospores biseriate, elliptic-fusiform, 9 - 1 0 x 2 , « , 1-septate;
paraphyses cylindrical, containing yellow oil drops.

On wood, apparently always associated with members of the Sphaeriales,


either obviously seated on superficial stromata or over the ostioles of immer-
sed perithecia, September to January. Common.

Calycella subpallida (Rehm) Dennis, Comm. Myc. Inst. Myc. P a p . 62:


45 (1956), is a small whitish or cream-coloured species not uncommon
on decorticated wood. Calycella fuscocincta Graddon in Trans. Brit,
mycol. Soc. 5 0 : 9 (1967) is distinguished b y its blackish surface to
the receptacle.

CUDONIELLA Saccardo

Cudoniella Saccardo, Sylloge F u n g o r u m 8 : 4 1 (1889).


Apothecia like those of Hymenoscyphus b u t typically with a convex
hymenium reminiscent of Leotia, often with lilaceous tints. Asus pore

119
neither blued nor outlined b y iodine, ascospores sometimes becoming
1-septate at m a t u r i t y .
%

Cudoniella acicularis (Bulliard ex Fries) Schroeter a p u d Cohn, K r y p t .


Flora Schles. 3 (2):21 (1893). (Fig. 7M and l c . )
Apothecia gregarious, superficial, with long, smooth, cylindrical
stalks, white t h r o u g h o u t , becoming greyish or brownish only with age,
disc convex with a reflexed margin, up to 4 m m . across. Asci cylindric-
clavate, up to 1 2 0 x 1 3 p , apex somewhat conical, 8-spored; ascospores
biseriate, fusiform or inequilateral, 1 5 ^ 2 x 4 - 5 / « , becoming 1-septate
with age, o c c a s i o n a l l y j ^ s e p t a t e ; paraphyses slender, swelling to 3 - 5 fi
thick towards the tip.

On rotting trunks and the cut surfaces of tree stumps, especially of


Quercus. September to December. Common.
This is the type species of Helotium Persoon ex St. Amans 1821, a name
invalidated by Fries adoption in the last volume of Systema mycologicum
1832 of Helotium Tode ex Leman 1821, a genus of Basidiomycetes. Names
adopted by Fries in any volume of his Systema take precedence over all
others proposed or validated during the period 1821-1832 inclusive. Hence
the generic name Helotium Persoon ex St. Amans, adopted in the first
edition of this book, is abandoned here, to be replaced by Cudoniella and
Hymenoseyphus.

Cudoniella clavus (Albertini & Schweinitz ex Fries) Dennis in Per-


soonia 3 : 7 3 ( 1 9 6 4 ) . ( P L X I I I R . )
Apothecia gregarious or solitary, more or less stalked, disc up to
10 m m . across, at first concave, t h e n becoming strongly convex with
reflexed margin, whitish or greyish, usually flushed with violaceous
tints, stalk smooth and white b u t often brown or black at the base.
Asci cylindric-clavate, varying greatly in length, up to 115x10//,
8-spored; ascospores uniseriate or irregularly biseriate, oblong-fusi-
form, often narrower at one end, 1 0 - 1 7 x 3 - 5 /<, nonseptate; paraphyses
slender, cylindrical.

Best developed on rotting twigs and leaves covered with water, as in


ditches and swamps, but also on dead stems of Chamaenerion and the like.
May and June, sometimes in July. Common.
There is a yar. grandis (Boudier) Dennis, op. cit.: 73 (1964), up to 2 cm.
across, subsessile o n T u m u s in swamps in May. In typical C. clavus the
length of the stalk depends on the depth to which the substrate is sub-
merged, it is always long enough to lift the disc clear of the water.
The species is often mistakenly called "Ombrophila" clavus.

120
Cudoniella junciseda (Venenovsky) Dennis comb.nov.
Helotium juncisedum Velenovsky, Monog. Discom. Bohem.: 198
(1934). (PI. XIVJ.)
Apothecia scattered, disc flat, flesh-coloured, 2 m m . wide, receptacle
white, short-stalked. Asci 80-120x6-8/«, 8-spored, spores biseriate,
elliptical to clavate, 9 - 1 4 x 2 . 5 - 4 / / , paraphyses cylindrical 3 y wide.
On dead watersoaked stems and fruit of J uncus effusus. July. Uncommon.

Cudoniella rubicunda (Rehm) Dennis in Persoonia 3 : 7 3 (1964).


Apothecia scattered, superficial, disc convex, 2 - 5 m m . wide, bright
purplish-red, receptacle t u r b i n a t e , short-stalked, pinkish-violaceous.
Asci 4 5 - 5 0 x 6 - 7 / , , 8-spored, spores elliptical, 6 - 8 x 2 - 3 / , , with polar
guttules, paraphyses cylindrical 2 / / wide. ( P I . X I V K . )
On fallen cones of Pinus silvestris. August. Rare.

HYMENOSCYPHUS S. F. Gray

Hymenoscyphus S. F. Gray. Natural Arrangement British Plants 1 : 6 7 3


(1821).

This includes the great m a j o r i t y of species commonly referred to


Helotium Persoon ex St. A m a n s or Helotium Fries 1849 non Fries 1832.
Apothecia of small or medium size, usually more or less stalked,
saprophytic on vegetable debris. Outer tissue of the cup composed of
approximately parallel, rather thin-walled hyphae, lying at a low angle
to the surface and clearly differentiated from the often rather loosely
woven hyphae of the flesh. Asci 8-spored, ascospores hyaline, narrowly
elliptical or somewhat fusiform to navicular, in a few species becoming
1-septate. Paraphyses cylindrical, obtuse, not branched towards the
tip.
In addition to these, thick-fleshed, more or less sessile species on
wood and bark have been called Pachydisca Boudier in Bull. Soc.
mycol. France 1 : 1 1 2 (1885), and there is a small group of species
a p p a r e n t l y parasitic on B r y o p h y t a which m a y prove worth generic
rank when t h e y have been adequately studied. A t e n t a t i v e t r e a t m e n t
of t h e British species will be found in Commonwealth Mycological
Institute Mycological P a p e r 62 b u t the genus is still far from being
adequately studied. The following have been selected as representative
species:

121
Hymenoscyphus vernus (Boudier) Dennis in Persoonia 3 : 7 8 (1964).
(PI. X V I H . )

Apothecia scattered, disc slightly convex, up to 4 m m . across, fawn,


exterior paler, with a short slender stalk which is blackish in the lower
p a r t and more or less downy. Asci cylindric-clavate, up to 90x8/«,
8-spored, the pore not clearly blued b y iodine; ascospores uniseriate
or biseriate, narrowly elliptical, 7 - 1 1 x 3 - 4 / « , non-septate; paraphyses
subcylindric, slightly enlarged to about 3 /«thick towards the rounded
tip, often containing brownish granules.

On twigs and small branches in damp, shady, places, as in the lower part
of heaps of prunings, February to March. Not common. This is Ombrophila
verna Boudier in Bull. Soc. mycol. France 4:77 (1889), but is obviously not
an Ombrophila. There is a different Helotium vernum Velenovsky, Mono-
graphia Discomycetum Bohemiae:196 (1934).

Hymcnoscyphus scutula (Persoon ex Fries) Phillips, British Discom.


137 (1887). (PI. XVIF.)

Apothecia scattered, disc flat with a t h i n low margin, cream to


yellowish, drying ochraceous, up to 4 m m . across, exterior whitish,
appearing fibrous under a lens and often longitudinally streaked with
darker bands of hyphae, seated on a slender cylindrical stalk. Asci
cylindric-clavate, up to 110x11/«, 8-spored, somewhat conical at the
tip with a pore blued b y iodine; ascospores biseriate, somewhat
cylindrical, pointed below and asymmetrically rounded or slightly
beaked above, with a fine hyaline bristle at the lower end and usually
a shorter one a t t a c h e d to the beak, containing two oil drops, some-
times becoming septate, 18-27x3.5-5/«; paraphyses slender, enlarged
u p w a r d s to about 3 /«.

On dead herbaceous stems, especially of Compositae, October to De-


cember. Very common.
There are also allied species or varieties, found especially on stems of
Mentha, Teucrium, Solanum, Campanula and the like with smaller non-
ciliate~~ascospores about 16-18x3-4/« anH a var. fucatus Phillips op. cit. 137
(1887), with ascospores 24-34x5.7/« apparently contined to wet stems of
Polygonum.

Hymenoscyphus vitellinus (Rehm) 0 . Kuntze Rev. Gen. PI. 3 (2): 486


(1898) is an acceptable name for very similar apothecia with aco-
spores only 1 6 - 2 0 x 3 - 3 , 5 /«, without appendages, on herbaceous stems.

122
Hymenoscyphus rcpandus (Phillips) Dennis in Persoonia 3 : 7 5 (1964).
Apothecia like those of H. scutula b u t smaller, pale yellow through-
out, asci 6 0 - 7 0 x 5 / / , spores narrowly elliptical, 8 - 1 3 x 2 - 2 . 5 / / , without
appendages. ( P L X I V G . )
On dead stems of Cirsium, Epilobium, Filipendula, Halimione, Mercuria-
lis, XJlex &c, dead Iris, Juncus, capsules olJEsculus &c. May to November.
Common. ~

Hymenoscyphus calyculus (Sowerby ex Fries) Phillips, op. cit.: 136


"(1887). (PL XVIG.)
Apothecia scattered or in small clusters, cup-shaped with a concave
disc about 2 m m . across and a cylindrical stalk, which m a y be downy
especially towards the base. Asci cylindric-clavate, up to 125x10 //,
8-spored, the pore blued b y iodine; ascospores irregularly biseriate,
cylindrical, rounded above and tapering below, 1 5 - 2 2 x 3 - 4 . 5 / / , some-
times appearing 1-septate; paraphyses cylindrical, up to 3// thick.
On dead wood and fallen twigs, September to January, occasionally in
spring. Widespread but seldom abundant.

Helotium virgultorum and H. sublenticular are ill-defined names


often used for this species.
H. conscriptum (Karsten) Karsten in Nov. Sail. Sk. F u n a Flora
Fennica 11:236 (1870), is a name used for a form on Salix, with asco-
spores towards the lower end of the range in length. This is t h e form
figured here.

Hymenoscyphus fructigenus (Bulliard ex Merat) S. F . Gray, op. cit. 673


(1821). (PL XVIM.)
Apothecia superficial, solitary or in clusters, cup-shaped with a flat
yellowish disc up to 3 m m . across and a slender smooth cream-coloured
stalk. Asci cylindric-clavate up to 2 0 0 x 8 / / , 8-spored, the pore blued
b y iodine; ascospores biseriate, oblong-fusiform or slightly inequi-
lateral, 13-21x3-4/«, occasionally appearing 1-septate; paraphyses
cylindrical, a b o u t 2 // thick.
On fallen nuts of Corylus, cupules of Fagus, acorns of Quercus and stones
of Prunus, August to November. Common.

Hymenoscyphus caudatus (Karsten) Dennis in Persoonia 3 : 7 6 (1964).


(PL XVIi.)
Apothecia scattered, white, drying yellowish, disc flat, about 1 m m .
across, stalk smooth and slender. Asci cylindric-clavate, up to 95 x 10 //,

123
8-spored, pore blued by iodine; ascospores irregularly biseriate,
cylindrical or slightly inequilateral, somewhat pointed below, 16-20 x
4-5/«, non-septate; paraphyses slender, cylindrical.
On decaying fallen leaves of Acer, Alnus, Berberis, Corylus, Fagus, Popu-
lus, Quercus, Tilia and Salix, August to November. Common.
A very similar fungus on blackened patches of fallen Fraxinus petioles,
with ascospores 13-17 x 3-5// has been distinguished as H. albidus (Roberge)
Phillips op. cit. 138 [1887]) ( = Helotium albidum [Roberge & Desmazieres)
Patouillard, Tab. Anal. Fung. 4:173 [1885], non H. albidum Crouan Florule
du Finistere: 47 [1867]).

Hymenoscyphus phyllogenus (Rehm) 0 . Kuntze, op. cit. 485 (1898).


Apothecia scattered, white t h r o u g h o u t , drying yellowish, disc flat,
up to 0.5 mm. across, stalk long, smooth, slender. Asci up to 75 x 10 //,
8-spored, the pore blued b y iodine; ascospores uniseriate or irregularly
biseriate, subcylindric, 1 2 - 1 6 x 4 - 6 n , non-septate; paraphyses slender,
about 3 // thick. ( P I . X V I L . )
On fallen leaves of Populus and Quercus, in autumn. Apparently not
common.

Hymenoscyphus phyllophilus (Desmazieres) 0 . Kuntze, op. cit. 485


(1898). (PI. X V I k . )
Apothecia scattered, white t h r o u g h o u t , drying yellowish, disc
slightly concave, up to 0.4 m m . across, exterior smooth or minutely
downy, with a short cylindrical stalk. Asci cylindric-clavate, up to
7 5 x 9 / / , 8-spored, the pore not blued b y iodine; ascospores biseriate,
cylindric-fusiform, 1 2 - 1 4 x 3 - 4 / / , always distinctly 1-septate; para-
physes cylindrical, 2 - 3 // thick.
On veins of fallen leaves, September to October. Common in leaf beds in
woods of Fagus, occasionally on leaves of Quercus.
Other small whitish species on fallen leaves are Hymenoscyphus immu-
tabilis (P'uckel) Dennis op. cit. 76 (1964), on Populus, Quercus, Bobinia and
Vlmus, short-stalked, with ascospores 10-13 x 4-4.5 //; H. nitidulus (Berkeley
& Broome) Phillips op. cit. 141 (1887) on leaves of Deschampsia.

Hymenoscyphus rhodoleucus (Fries) Phillips, op. cit. 131 (1887). (PI.


XVID.)
Apothecia scattered, pale pink throughout, drying yellowish-brown,
disc flat, up to about 1 m m . across, stalk smooth and slender. Asci up
to 7 0 x 6 / / , cylindric-clavate, 8-spored, the pore blued b y iodine; asco-
spores uniseriate or biseriate, narrowly elliptical, 9 - 1 2 x 5 - 6 / / , non-
septate ; paraphyses slender, about 2 /< thick.

124
On dead stems of Equisetum, in May and September. Fairly common, not
to be confused with Stamnaria persoonii on the same host, figured alongside
for comparison.

Hymenoscyphus robustior (Karsten) Dennis in Persoonia 3 : 7 5 (1964).

Apothecia scattered, disc concave, up to 2 m m . wide, yellow, re-


ceptacle cupshaped, light yellow, with a cylindrical stalk pink when
fresh, drying yellowish brown. Asci 90-120x7-9/«, 8-spored, spores
obtusely elliptic-cylindric, 7 - 1 4 x 3 - 4 / « , paraphyses cylindrical, 2-2.5 /«
wide. (PI. XIVii.)

On dead stems of marsh plants, Cladium, Galium, Ph.ragm.iles, Scirpus,


Typha. June-July. Probably common.

Hymenoscyphus rumicis (Velenovsky) com.nov. (PI. XIVs.)

Septatium rumicis Velenovsky Monog, Discom. B o h e m . : 2 1 3 (1934).


Apothecia solitary, disc yellowish, up to 2 m m . wide, receptacle
shallow cupshaped, cream coloured, drying brownish, with very long
slender stalk. Asci 60-85x6-7/«, pore blued b y iodine, spores elliptic-
cylindric, 9 - 1 3 x 3 - 4 /«, mostly 1-septate, paraphyses cylindrical, 2-2.5 y
wide.

On fallen fruit of Rumex acetosa. June to August. Uncommon.


Septatium Velenovsky was defined as "Ilabitu et notis plene Helotium
quadrat, sed sporae semper cylindrico-elongatae 2-5 cellulares praestant."
This is not for me a generic character, scarcely diagnostic at species level.

Hymenoscyphus herbarum (Persoon ex Fries) Dennis, op. cit. 77 (1964).


(PI. X V I E . )

Apothecia gregarious, superficial, disc up to 3 m m . across, often


slightly convex, yellowish to ochraceous, exterior paler, minutely
downy, seated on a short stalk. Asci cylindric-clavate, up to 90x8/«,
8-spored, pore blued b y iodine; ascospores often biseriate, cylindric-
fusiform, 13-17x2.5-3/«, 1-septate; paraphyses cylindrical, about 2 y
thick.

On dead herbaceous stems, especially of Urtica, October to January.


Very common.

Hymenoscyphus separabilis (Karsten) Dennis, op. cit. 77 (1964) is a


very similar fungus on dead canes of Rubus idaeus.

125
Hymenoscyphus epiphyllus (Persoon ex Fries) Rehm apud K a u f f m a n
in Papers Michigan Acad. Science, Arts, Letters 9 : 7 7 (1929). (PL
XVIJ.)

Apothecia superficial, scattered, bright yellow to orange through-


out, cup-shaped, subsessile on a cylindrical smooth base, disc up to
about 2 m m . wide. Asci up to 120x12 /«, cylindric-clavate, 8-spored,
pore blued by iodine; ascospores often becoming biseriate, oblong-
fusoid, 1 5 - 1 8 x 3 . 5 - 5 ju, often pointed below, sometimes appearing
1-seriate; paraphyses cylindrical, 2 - 3 /« thick, sometimes branched.

On fallen leaves, September to November. Common.

A scarcely distinguishable fungus on fallen needles of Pinus has


been called Helotium sulphuratum\ H. carpinicolum Rehm in Hed-
wigia 35:146 (1896), on fallen Carpinus leaves has been claimed to be
distinct, with smaller ascospores 1 3 - 1 6 x 4 - 4 . 5 /«.
Hymenoscyphus fagineus (Persoon ex Fries) Dennis, op. cit. 74
(1964), 356 (1849), is a r a t h e r similar b u t white fungus, drying reddish-
brown, found on fallen cupules of Fagus, with ascospores non-septate,
8-16x4-5/«.

Hymenoscyphus marchantiae (Berkeley) Dennis, op. cit. 77 (1964).


(PL X X I V l . )
Apothecia solitary, subsessile, smooth, soft-fleshed, light yellowish-
brown t h r o u g h o u t , disc concave, scarcely 2 m m . across. Asci cylindric-
clavate, u p to 80x10/«, 8-spored, pore blued b y iodine; ascospores
sometimes biseriate, elliptic-fusiform, 14-18x3-4/«, non-septate;
paraphyses numerous, slender, 2.5 /«thick.

On dead patches in thalli of Marchantia and Conocephalus, April and


May. Not uncommon.

H. vasaensis (Karsten) Dennis, op. cit.: 77 (1964) is a somewhat simi-


lar species on Sphagnum, w i t h whitish apothecia and ascospores
10-18x2-4.5/«.

The following species, with general appearance and structure of a


Hymenoscyphus, is of uncertain systematic position because its ascus
pore appears constantly iodine-negative. Its affinity m a y lie r a t h e r
with Rutstroemia b u t there is a small group of similar species parasitic
on B r y o p h y t a for which a separate genus m a y perhaps be needed.

126
Helotium fulvum Boudier, in Bull. Soc. mycol. France 13:16 (1897).
Apothecia solitary, disc flat, dark brown, about 1 m m . wide, re-
ceptacle cupshaped with a stout cylindrical stalk, smooth, concolorous,
softfleshed. Asci 150-180x13-16/«, 8-spored, spores elliptical to
oval, with numerous large oil drops, 16-21x6-9/«, paraphyses
cylindrical, 3/« wide. (PI. X I V u . )

In leaf axils of Dicranella heteromera and "muscos minores" (Boudier).


October to February. Rare.

STAMNARIA Fuckel

Stamnaria Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:309


(1870).
Apothecia e r u m p e n t , with soft flesh composed of broad parallel
hyphae, stalked, disc with a deep cylindrical collar, asci 8-spored,
ascospores broadly elliptical, hyaline, non-septate. There is only one
species:

Stamnaria persoonii (Mougeot ex Persoon) Fuckel, op. cit.: 309 (1870).


(PI. XVIc.)
Apothecia scattered or in small groups, e r u m p e n t , with a slender,
smooth cylindrical stalk, above cup-shaped, scarcely 1 m m . broad, disc
concave, surrounded b y a translucent white cylindrical collar u p to
1 m m . deep, otherwise pink t h r o u g h o u t . Asci cylindrical, rounded
above, up to 130x12/«, 8-spored, the pore not distinctly blued b y io-
dine; ascospores uniseriate, broadly elliptical with rounded ends,
15-18x5-8/«, hyaline, usually with a central oil drop, n o n - s e p t a t e ;
paraphyses cylindrical, enlarged to about 4/« near the tip.

On stems olJiguis£lum, May and June. Not common. Compare Hymeno-


scyphus rhodoleucus.

BELONIOSCYPHA Rehm

Belonioscypha Rehm, Rabenhorst's K r y p t . Flora Deutschland 1 (3):


743 (1893).
Apothecia superficial, cup-shaped, smooth, stalked, light-coloured,
outer tissue of glassy-walled h y p h a e nearly parallel with t h e surface,
asci 8-spored, ascospores multiseptate, hyaline, paraphyses cylindrical.
There is one British species:

127
Belonioscypha culmicola (Desmazieres) Dennis, Comm. Myc. Inst.
Myc. P a p . 6 2 : 3 9 (1956). (PI. X V I I G . )
Apothecia scattered, superficial, cup-shaped, about 1 m m . broad,
disc concave with a toothed margin, ivory-white or yellowish through-
out, exterior smooth with a short cylindrical stalk. Asci cylindric-
clavate, up to 140x 14 8-spored, the pore blued b y iodine; ascospores
sometimes biseriate, cylindric-fusiform, 1 9 - 3 3 x 3 - 4 with a thick
hyaline gelatinous coating, 3-septate; paraphyses cylindrical, about
3 [i thick.

On dead stems, especially of grasses, also on Juncus and dictotyledonous


herbs, Jufy to September. Fairly common.

CYATHICULA de Notaris

Cyathicula de Notaris in Commentario della Societa Crittogamologica


Italiana 1:381 (1864).
Apothecia like those of Belonioscypha b u t with prominent marginal
t e e t h and with the ascospores 0-1-septate.

Cyathicula coronata (Bulliard ex Merat) de Notaris apud Karsten


Fungi Fenn. Exsic. 146 (1865). (PI. X V I I F . )
Apothecia scattered, superficial, cup-shaped with a slender cylindri-
cal stalk, smooth, whitish to cream-coloured, disc up to 3 m m . broad,
concave, sometimes with a pinkish flush, surrounded b y a ring of erect,
narrow, pointed, whitish teeth. Asci cylindric-clavate, up to 110x9/«,
8-spored; ascospores biseriate, cylindric-fusiform, 1 5 - 2 0 x 3 - 4 . 5 fi, with
two large oil drops, non-septate; paraphyses slender, 2 fi thick.

Fairly common on dead herbaceous stems, September to November, rare


on decorticated wood.

PHIALEA (Fries) Gillet

Phialea (Fries) Gillet, Champignons de France, Discomycetes: 93 (1879).


Apothecia small, cup-shaped, more or less stalked, usually smooth
b u t sometimes powdered with minute crystals, outer flesh composed
of parallel hyphae with very thick glassy walls and thin septa, con-
t r a s t i n g strongly in longitudinal sections with the thin-walled h y p h a e
of the inner flesh, asci narrow, 8-spored; ascospores narrowly fusiform,

128
non-septate, hyaline; paraphyses often relatively broad, pointed above
and longer t h a n t h e asci.

Phialea cyathoidea (Bulliard ex Merat) Gillet, op. cit.: 106 (1879). (PI.
XVIIA.)
Apothecia scattered, superficial, smooth, cup-shaped with a slender
cylindrical stalk, whitish or cream-coloured throughout, disc up to
2 m m . across, usually smaller, concave. Asci cylindric-clavate, up to
5 0 x 4 - 5 / I , 8-spored, t h e small pore blued b y iodine; ascospores bi-
seriate, narrowly fusiform, 6-12x1.5-2.5/«, often with a minute oil
drop at each e n d ; paraphyses about 2 /i thick.
On dead stems of herbaceous plants, throughout the year. Extremely
common everywhere. This is no doubt a collective species composed of many
races difficult to distinguish morphologically, a rather distinctive large
whitish form on stems of Cnicus is sometimes called var. albidula (Hedwig
ex Karsten) Rehm.

Phialea egenula Rehm, Rabenhorst's K r y p t . Flora Deutschland 1 (3):


726 (1893). (PI. X V I I b . )
Apothecia scattered, superficial, smooth, pale pinkish-brown, with a
short slender cylindrical stalk, disc concave, yellowish, about 0.5 m m .
across. Asci cylindric-clavate, up to 60x6/«, 8-spored, the small pore
blued b y iodine; ascospores biseriate, fusiform, 10-12x2-2.5/«, non-
septate ; paraphyses cylindrical, 3 /« thick, rounded above, with oily
contents.
On dead stems of Rumex acetosa, September. Uncommon.

Phialea turbinata Sydow in Annales mycologici 5:397 (1907). (PI.


XVIIn.)
Apothecia scattered, superficial, smooth, whitish at the margin
shading to smoky-brown below, cup-shaped, thin-fleshed, with a
cylindrical brown stalk and concave disc up to 0.5 m m . across. Asci
cylindric-clavate, up to 50x5/«, 8-spored, apex conical; ascospores
biseriate, narrowly fusiform, 6 - 9 x 1 - 1 . 5 / « ; paraphyses lanceolate, 2/«
thick, longer t h a n the asci.
On dead stems of Ranunculaceae in ditches and swamps, April to June.
Probably common but seldom collected.

Phiala pteridicola (Crouan) Gillet, op. cit. 104 (1879). (PL X V I I c . )


Apothecia scattered or in small groups, superficial, cup-shaped with
a short, stout, cylindrical stalk, grey, vertically streaked with brown,

129
1

disc concave, up to 1 m m . across. Asci cylindric-clavate, up to 5 0 x 5 /«,


8-spored, t h e pore blued b y iodine; ascospores biseriate, elliptic-fusi-
form, 8 - 1 0 x 1 . 5 - 2 / / w i t h a small oil drop at each end, non-septate;
paraphyses cylindrical, up to 3// thick towards the tip, with greenish
contents.
On dead stems of PteridiMtr^a^ilinum, in spring. Common.

Phialea subhyalina Rehm, Rabenhorst's K r y p t . Flora Deutschland 1


(3): 1233 (1896). (PL XVIIE.)
Apothecia scattered, superficial, cup-shaped, smooth, translucent,
whitish, stalked, disc concave less t h a n 0.5 m m . across. Asci cylindric-
clavate, up to 110x10//, 8-spored, pore blued b y iodine; ascospores bi-
seriate, fusiform, 1 4 - 1 8 x 4 - 5 / / , non-septate; paraphyses slender, 2//
broad.
On petioles of fallen leaves of Acej^pseudaplMamis in autumn. Common.

Phialea incertella Rehm in Annales mycologici 4 : 4 8 5 (1906). (PL


XVIIIK.)
Apothecia e r u m p e n t , cup-shaped with an inrolled margin, about
300 // broad and high, subsessile, externally greenish-grey and smooth,
drying black. Asci cylindric-clavate, up to 4 0 x 5 //, 8-spored, the small
pore blued by iodine; ascospores biseriate, narrowly fusiform, 6 - 8 x
1.5-2 /(, non-septate; paraphyses thin-walled, hyaline, about 4// broad,
lanceolate and as m u c h as 25 // longer t h a n the asci.
On dead leaves of Koeleria in summer. Perhaps rare or overlooked.
Though the species is so seldom seen it is included here as a British re-
presentative of a small group of Phialea species found in Europe on leaves
of various grasses, including Phleum. It is not unlikely that others of the
group will be found in Britain when diligently sought for.

PEZIZELLA Fuckel

Pezizella Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:299


(1870).
Apothecia small, light coloured, erumpent or superficial, outer flesh
composed of parallel, firm-walled, usually colourless hyphae, exterior
often minutely downy, asci 8-spored; ascospores often narrowly
elliptic-cylindric or slightly curved, non-septate; paraphyses slender,
not lanceolate. It is difficult to find a clear-cut distinction between

130
Pezizella and Hymenoscyphushi\imihe\a.tteTihe asci tend to be longer,
70 i-i long and more, and the ascospores are longer, over 12 /« long, or
relatively broader.

Pezizella vulgaris (Fries) von Hohnel in Mitt. bot. Lab. tech. Iloehsch.
Wien 3 : 5 4 (1926). (PL X V I I i . )
Apothecia erumpent from bark, usually in clusters, cup-shaped with
a short stout stalk, whitish or cream-coloured, downy with short uni-
cellular hairs, disc about 1 m m . across, concave, often flushed pink.
Asci cylindric-clavate, up to 45x5/«, 8-spored, pore not blued b y
iodine; ascospores biseriate, cylindrical with rounded ends, straight or
slightly curved, 6-10x1.5-2/«, hyaline, non-septate; paraphyses
cylindrical, 2 - 3 /< thick.

On dead twigs, as of Corylus, Rosa and Salix, July to October. Uncommon.

Pezizella parile (Karsten) Dennis, Comm. Myc. Inst. Myc. Pap. 62:49
(1956), is a similar species, with longer asci in which the pore is blued
by iodine and 1-septate ascospores 6 - 1 4 x 1 . 5 - 2 / « , on twigs of Betula.

Pezizella chionea (Fries) Dennis, Comm. Myc. Inst. Myc. P a p . 6 2 : 5 3


(1956). (Fig. 7J.)
Apothecia superficial, solitary or in small clusters, cup-shaped, ses-
sile on a broad base, whitish, minutely downy, disc up to 1 m m . across,
concave, pale buff. Asci up to 60x5/«, cylindric-clavate, 8-spored, pore
blued by iodine; ascospores elliptic-cylindric, straight or slightly
curved, 6-8x1.5/«, non-septate; paraphyses cylindrical, 2 n thick.
On fallen cones and needles of Pinus, in spring. Fairly common.

Pezizella alniella (Nylander) Dennis, op. cit. 62:56 (1956). (PL X V I I n ) .


Apothecia scattered, superficial, saucer-shaped with a very short
slender stalk, cream-coloured, downy with white hairs, disc pale
yellow, flat, 0.5 m m . across. Asci cylindric-clavate, up to 55x7/«,
8-spored, pore blued by iodine; ascospores elliptic-cylindric, 8 - 1 l x
2.5-3/«, non-septate; paraphyses cylindrical, 2/« thick.
On fallen female catkins of Alnusin spring and autumn. Common.

Pezizella discreta (Karsten) Dennis, op. cit. 6 2 : 5 7 (1956). (PL XVIIJ.)

Apothecia scattered, superficial, cup-shaped with a short stalk,


smooth, soft, white, drying cream-coloured, disc concave 0.5 m m .

131
across. Asci cylindric-clavate, about 5 0 x 6 / i , 8-spored, pore blued b y
iodine; ascospores biseriate, elliptic-cylindric with rounded ends,
sometimes slightly curved, 4 - 8 x 1 - 1 . 5 / / .
On dead herbaceous stems, as Artemisia and Hamulus, in autumn, some-
times in company with Hymenoscyphus sculula. Uncommon.

Pezizella amenti (Batsch ex Fries) Dennis, op. cit. 62:55 (1956). (PI.
XVIIM.)
Apothecia scattered, superficial, cup-shaped with a short cylindrical
stalk, whitish, minutely downy, disc concave up to 1 m m . across. Asci
cylindric-clavate, up to 50x5/«, 8-spored, the pore blued b y iodine;
ascospores often biseriate, rounded above, tapering below, often curved
and comma-like, 6 - 1 2 x 3 - 4 / / , non-septate; paraphyses cylindrical, 2/«
thick.
On fallen female catkins of Salix and occasionally of Populus, March to
April. Common. This species is sometimes referred to Ciboria but its struc-
ture is that of a Pezizella or a Hymenoscyphus.

Pezizella gemmarum (Boudier) Dennis, op. cit.:55 (1956), is a rather


similar little fungus, with elliptic-fusiform ascospores 6 - 8 x 2 - 2 . 5 / / , on
overwintered fallen budscales of Populus.

Pezizella chrysostigma (Fries) Saccardo, Sylloge F u n g o r u m 8 : 2 8 8


(1889). (PL XVIIL.)
Apothecia scattered, superficial, cup-shaped with a short cylindrical
stalk, white, minutely downy, disc about 0.6 m m . across, fringed with
white downy hairs. Asci cylindric-clavate, up 30x3/«, 8-spored, the
pore blued by iodine; ascospores biseriate, narrowly fusiform or cla-
vate, 4-7x1-1.25/«, non-septate; paraphyses filiform.
On dead petioles of Nephrodium filix-mas, November to May. Common.

Pezizella campanulaeformis (Fuckel) Dennis, Comm. Myc. Inst. Mvc.


Pap. 62:60 (1956), is a somewhat similar b u t smooth fungus with asci
up to 7 0 x 7 //, on petioles of Nephrodium and Asplenium.

Pezizella eburnea (Iloberge) Dennis, Comm. Myc. Inst. Myc. Pap. 62:
61 (1956), (Fig. 7H.)
Apothccia scattered, superficial, cup-shaped with a short stalk, very
minutely downy towards t h e margin, white throughout, disc about
0.2 m m . across. Asci cylindric-clavate, up to 4 0 x 5 / / , 8-spored, pore
blued b y iodine; ascospores fusiform-clavate, biseriate, 6 - 8 x 1 - 1 . 5 / / ;
paraphyses slender.

132
On dead grass leaves and culms and on cereal stubble, May to August.
Fairly common.

Pezizella punctoidea (Karsten) Rehm, Rabenhorst's K r y p t . Flora


Deutschland 1 ( 3 ) : 6 6 8 ( 1 8 9 2 ) . ( P I . X V I I K . )
Apothecia gregarious, erumpent, cup-shaped, narrowed to a minute
peglike stalk, white, drying yellowish, pruinose, disc flat, less t h a n
0.5 m m . across. Asci cylindric-clavate, up to 4 0 x 6 / / , 8-spored, the
minute pore blued b y iodine; ascospores biseriate, fusiform-clavate,
5 - 7 x 1 . 5 - 2 / / non-septate; paraphyses cylindrical, 2/j, thick, often
branched at the septa.
On dead leaves and stems of Chamaenerion angustijolium, and Epilobium
hirsutum, July to October. Not common.

Pezizella subtilis (Fries) Dennis, Comm. Myc. Inst. Myc. Pap. 6 2 : 5 8


(1956), is a minute whitish-stalked fungus with ascospores 5 - 8 x 1 - 1 . 5 //
found in a u t u m n on fallen needles of Picea excelsa.

Pezizella eriophori Dennis in Kew Bulletin 16:323 (1962). (PI. XIVL.)


Apothecia scattered, sessile, disc convex, bright yellow, up to 1 m m .
diameter, receptacle cupshaped, concolorous, smooth. Asci 75-90x
6-7,«,.pore blued by iodine, 8-spored, spores elliptic-clavate, 8 - 1 2 x
2.5-3'//, paraphyses cylindrical with oily yellow contents.
On rotting leaves of Eriophorum, and Carex spp. June-July. Uncommon.

Pezizella polytrichi Dennis in Kew Bulletin 16:324 (1962).


Apothecia scattered, sessile, disc convex, whitish, y 3 m m . diameter,
receptacle whitish above, brown at the base and seated on a m a t of
brown hyphae. Asci 5 0 X 7 / « , spores 8 - 1 2 X 1 . 5 - 2 / / . ( P I . X I V M . )
On dead leaves of Pohjtrickum. April. Uncommon.
Compare also "Mollisia" chionea, p. 186.

CHLOROSPLENIUM Fries

Chlorosplenium Fries S u m m a Veg. Scand., Sect. Post.:356 (1849)*.


Apothecia developed on wood or herbaceous stems, always green
or olive throughout, outer tissue composed of hyphae almost vertical

* Chlorociboria Seaver, the name formerly adopted for this genus, cannot
be used since it has been made illegitimate bv Korf et al., see Mvcologia 51,
p. 208, 1959.

133
to the surface. Asci 8-spored, ascospores hyaline, non-septate. There
are four British species:

Chlorosplenium aeruginascens (Nylander) Karsten in Not. Sallsk.


F a u n a Flora Fennica 1 1 : 2 3 3 ( 1 8 7 0 ) . ( P I . X V I I I F . )

Apothecia scattered, superficial, cup-shaped, becoming flattened,


with a short slender stalk, smooth, thin-fleshed, blue-green throughout,
sometimes assuming yellowish tints, disc up to 5 m m . across. Asci
cylindric-clavate, up to 7 0 x 5 / / , 8-spored, the small pore blued by io-
dine; ascospores biseriate, fusiform, 6 - 1 0 x 1 . 5 - 2 / / , with a small oil
drop at each end, non-septate; paraphyses cylindrical, slender,
branched at the base, longer t h a n the asci, about 1.5 /< thick.

On rotting logs and decorticated wood, especially of Quercus, in damp


woods, April to November but chiefly in autumn. Very common. Wood
permeated by the mycelium is itself stained blue-green.

Chlorosplenium aeruginosum (Fries) de Notaris in Comm. Soc. Critt.


Ital.:376 (1864) as interpreted b y continental authors, differs from
the above in its larger ascospores, 9 - 1 5 x 1 . 5 - 2 . 5 / / , and in the disc
usually drying yellow. /

Chlorosplenium versiforme (Persoon ex Fries) de Not., op. cit.:376


(1864). (PI. XVIIIG.)

Apothecia scattered or in small clusters, superficial, more or less


cup-shaped but often becoming one-sided and somewhat spathulate,
short-stalked, externally pruinose, yellowish olive-green, drying with
a golden sheen or blackening, disc often convex, commonly 1 cm. or
more across. Asci cylindric-clavate, up to 8 0 x 7 / / , 8-spored, the pore
blued by iodine; ascospores biseriate, cylindrical with rounded ends,
straight or slightly curved, 9 - 1 3 x 3 / , ; paraphyses slender, 2// thick.

On rotting wood of various kinds, Fraxinus, Quercus, Salix, not stained


blue-green, September to November. Rare.
This species shows affinity with the Encoelioideae.

Chlorosplenium aeruginellum (Karsten) Karsten. Mycol. F e n n . l : 1 0 4


(1871), is a small blue-green fungus on rotting stems of Filipendula
ulmaria.

134
DISCINELLA Boudier

Discinella Boudier in Bull. Soc. mycol. France 1 : 1 1 2 (1885).


Apothecia terrestrial, sessile or nearly so, cup-shaped, often com-
paratively large and liable to be mistaken for members of the Pezi-
zales, soft-fleshed, asci 8-spored, the pore not distinctly blued by io-
dine, ascospores hyaline, non-septate. Sphagnicola is differentiated by
its gelatinous outer coating.

Discinella boudieri (Quelet) Boudier, Icones mycologicae, Livraison 6


(December 1905). (PL XVM.)
Apothecia terrestrial, often clustered, cup-shaped then flattened,
closely adpressed to the ground, smooth or finely tomentose and scurfy
near t h e margin, seated on a small stem-like base, disc somewhat urn-
bilicate, purplish-bay, or browner in the var. spadicea Boud., flesh
formed of broad interwoven hyphae, those of t h e outer tissue more
slender, thin-walled. Asci cylindric-clavate, rounded above up to
120x 13 /«, 8-spored, the sides of the pore outlined blue in iodine; asco-
spores uniseriate, elliptical, 10-15x4-5/«, non-septate; paraphyses
slender, slightly enlarged to 3 /«thick at the tip.

On sandy soils in autumn. Rare.

Discinella margarita Buckley in Trans. Brit, mycol. Soc. 6:347 (1920).


(Pl. XVx.)
Apothecia scattered or in small groups on soil, cup-shaped, sessile,
white or cream-coloured throughout, soft-fleshed, disc 1 - 4 m m . across,
slightly concave, margin fringed with very delicate, short, pointed
teeth, formed of cohering, thin-walled, septate hyphae, t e e t h dis-
appearing with age. Outer layers of the flesh composed of large, almost
isodiametric, thin-walled cells, which are replaced by parallel h y p h a e
at the margin. Asci cylindric-clavate, up to 100x10/«, 8-spored, the
pore not blued by iodine; ascospores sometimes biseriate, elliptic-
cylindric, 8 - 1 4 x 3 - 5 / « ; paraphyses slender, 1-2/« thick.

On soil under conifers, May to September. Uncommon.

Discinella menziesii (Boudier) Boudier in Trans. Brit, mycol. Soc. 4:


323 (1914), is a pale pink terrestrial species, up to about 1 cm. across
with ascospores 15-17x4-5/«.

135
Tribe Encoelioideae
The members of this tribe are characterised b y their relatively large
clustered apothecia, erumpent from bark, with the outer tissue com-
posed of cuboid or rounded cells, generally brown-walled, which be-
come loose at the surface to give a mealy or scurfy appearance to the
outside of the cup.
I. Flesh containing numerous large cells, quite 30 // across, filled with
greenish-brown sap Yelutarina
If. Flesh without large greenish cells:
A. Ascospores 2-celled Encoeliopsis (p. 137)
B. Ascospores non-septate:
1. Ascospores broadly elliptical, apothecia on conifers
Cenangium (p. 137)
2. Ascospores narrowly elliptic-cylindric, often slightly curved:
a. Flesh yields a crimson stain in KOH solution, ascospores
become brownish Phaeangella (p. 138)
b. No such colour reaction, ascospores colourless
Encoelia (p. 138)
For superficially somewhat similar fungi with elongated multiseptate
ascospores see the Scleroderridoideae.

YELUTARINA Korf

Yelutarina Korf in Mycologia 45:476 (1953).


Apothecia sessile, cup-shaped, flesh containing numerous vesicular
cells with greenish contents, outer surface powdery, asci 8-spored, the
pore blued by iodine, ascospores non-septate, brown with age. There
is one British species:

Velutarina rufo-olivacea (Albertini& Schweinitz ex Persoon) Korf, op.


cit.:476 (1953). (PI. X V I I I n . )
Apothecia solitary or in small clusters, erumpent, cup-shaped with
an inrolled margin, subsessile, outer surface covered with a rustv-
brown or tan-coloured powder formed of thick-walled, loose, ir-
regularly lobed cells, disc up to 3 m m . across, olive-green or almost
black. Asci cylindric-clavate, up to 160x 12 //, 8-spored, the broad pore
blued by iodine; ascospores uniseriate, elliptical with obtusely rounded
ends, 1 0 - 1 4 x 6 - 7 / / , hyaline at first b u t ultimately becoming pale
brown, containing one or two large oil drops; paraphyses 3,u thick,
swelling to 5 n at the greenish-brown tip.

On dead twigs and branches, as of Acer, Fagus, Fraxinus, Rosa, Rubus,


Ulex, December to May. Not uncommon but seldom plentiful.

136
ENCOELIOPSIS Nannfeldt
Encoeliopsis Nannfeldt in Nova Acta Reg. Soc. Sci. Uppsala IV, 8:306
(1932).
Apothecia coriaceous, erumpent, asci 8-spored, ascospores hyaline,
fusiform, 1-septate. There is a single British species:

Encoeliopsis johnstoni (Berkeley) Dennis, Comm. Myc. Inst. Myc. P a p .


62:120 (1956). (PI. X V I I I c . )
Apothecia scattered, superficial, cup-shaped with a short stem-like
base, dark brown, finely scurfy, striate towards the margin, disc con-
cave, yellowish, drying almost black. Asci clavate, up to 110x12/«,
8-spored, the broad pore blued by iodine; ascospores biseriate, fusi-
form, straight or slightly curved, 1 8 - 3 3 x 6 - 8 / / , hyaline, becoming
1-septate; paraphyses cylindrical, up to 2/« thick at the yellowish tip.
On dead branches of Betula covered with stromatic Sphaeriales. Very rare.

CENANGIUM Fries

Cenangium Fries, Systema mycologicum 2:177 (1822).


Apothecia coriaceous, erumpent, asci 8-spored, ascospores hyaline,
non-septate, broadly elliptical, paraphyses swollen at the tip, on
branches of conifers or coniferous debris. Structurally Cenangium is
very like Encoelia b u t Gremmen who has recently studied members
of the two genera in culture considers t h e y m a y be distinguished b y
their microconidial states. Cenangium ferruginosum forms black sper-
mogonia with t i n y rod-shaped hyaline spermatia whereas slimy sporo-
dochia are formed on the mycelium of Encoelia fascicularis, bearing
globose microconidia like those found in species of Sclerotinia. See
Acta Botanica Neerlandica 2:226-241 (1953).

Cenangium ferruginosum Fries ex Fries, op. cit.: 187 (1822). (PL


XVIIIE.)

Apothecia erumpent singly or in small clusters, at first globose and


closed, t h e n t u r b i n a t e and opening b y an irregularly torn aperture,
exterior dark brown and pruinose or mealy, disc deeply concave, up
to 3 m m . across, ochraceous, completely concealed b y the inrolled
margin when dry. Asci cylindric-clavate, up to 80x15/«, 8-spored,
broadly rounded above, the pore not blued by iodine; ascospores

137
sometimes biseriate in the upper p a r t of the ascus, broadly elliptic-
fusiform, 1 2 - 1 4 x 5 - 6 / / , hyaline, non-septate; paraphyses slender,
hyaline, about 2/t thick, septate, swollen to 4 - 5 / / at the clavate tips
which are embedded in a layer of firm mucilage, contents yellowish,
oily.
On dead twigs and small branches of Pinus, April to June and August to
December. Fairly common.
The fungus is not parasitic on the pine but a saprophyte on dead branches.

Cenangium abietis (Persoon) D u b y is a synonym.


Cenangium acicolum (Fuckel) Rehm in Rabenhorst's K r y p t . Flora
Deutschland 1 (3): 228 (1889), is an allied species on dead pine needles.

PHAEANGELLA Massee

Phaeangella Massee, British Fungus Flora 4:136 (1895).


This is a genus of very doubtful value, possibly distinguished from
Encoelia b y its ascospores becoming brown and by the flesh yielding
crimson colour in K O H solution. Both characters require checking in
fresh collections. There is one British species:

Phaeangella ulicis (Cooke) Massee, op. cit.: 136 (1895). (PI. X V I I I j . )


Apothecia erumpent, solitary or in clusters, cup-shaped with a stem-
like base, dark brown, scurfy, disc up to 1.5 m m . across, deeply con-
cave with an inrolled margin, brown. Asci cylindric-clavate, up to
60x8/<, rounded above, the pore not blued by iodine, 8-spored; asco-
spores biseriate, narrowly elliptical, straight or slightly curved,
6 - 1 1 x 2 . 5 - 3 //, colourless b u t said to become brownish; paraphyses
slightly clavate, up to 3// wide at the tip, the contents of which be-
come wine-red in K O H solution.

On dead branches of Ulex, perhaps also on Sarothamnus, April. Very rare.


This fungus, the type species of Phaeangella, is known only from the
original collection made in 1870; it is most desirable that it be collected
again and fully studied when fresh.

ENCOELIA (Fries) Karsten

Encoelia (Fries) Karsten, Mycologia Fennica 1 : 1 8 (1871).


Apothecia erumpent from bark, leathery, medium to large, cup-
shaped, brown and scurfy, disc concave, asci 8-spored, ascospores
narrowly elliptical, non-septate.

138
Encoelia furfuracea (Roth ex Persoon) Karsten in Not. Sallsk. F a u n a
Flora Fennica 1 1 : 2 5 3 ( 1 8 7 0 ) . ( P I . X V I I I B . )
Apothecia erumpent singly or in small clusters, sessile, cup-shaped,
covered with a rust-coloured meal composed of globose cells 10-15 //
across, disc up to 15 m m . across, cinnamon coloured, drying black.
Asci clavate with a slender stalk, up to 120x7//, 8-spored, the pore
blued b y iodine; ascospores biseriate, cylindrical with rounded ends,
slightly curved, 6 - 1 1 x 2-2.5 /«, hyaline, non-septate; paraphyses slender,
swollen to 5 or 6 n thick at the tip.

On living branches of Alnus and Corylus, December to May. Uncommon.

There are also the following British species:


Encoelia fascicularis (Albertini & Schweinitz ex Persoon) Karsten
in Not. Sallsk. F a u n a Flora Fennica 11:253 (1870), on Fraxinus and
Populus.
Encoelia siparia (Berkeley & Broome) Nannfeldt in Trans. Brit,
mycol. Soc. 20:196 (1936), on Ulmus.
Encoelia tiliacea (Fries) Karsten, Mycologia Fennica 1:218 (1871) on
Tilia.

Tribe Scleroderridoideae

The Scleroderridoideae superficially resemble the Encoelioideae but


lack the latter's powdery outer surface; most species have greatly
elongated multiseptate ascospores. Two British genera are usually
recognised, Scleroderris Bonorden 1851 and Crumenula de Notaris
1864, b u t Groves considers t h e m better united under the older name
Godronia. On a different basis, however, two genera m a y still be
distinguished:

I. Ascus pore blued by iodine Godronia


II. Ascus pore not blued by iodine Ascocalyx

Ascocalyx abietis N a u m o v Bolesni Rast. 14:138 (1925), common in


northern Europe and North America on Abies, has the conidial state
Bothrodiscus pinicola Shear, with locules in a stalked discoid stroma
with membranous margin. This species is not known from t h e British
Isles b u t Groves has suggested t h a t Scleroderris lagerbergii Gremmen
in Sydowia 9:232 (1955) ( = Crumenula abietina Lagerberg 1913), the
ascus state of Brunchorstia pinea (Karsten) von Hohnel, common on
pines in Britain, is "closer to Ascocalyx t h a n to Godronia." For a full
account of Ascocalyx see Groves in Mycologia 28:451-462 (1936).

139
GODRONIA Mougeot & Leveille

Godronia Mougeot & Leveille apud Mougeot Consid. generales Veg.


spont. Dep. Vosges: 355 (1846).
Adoption of the old name Godronia has been deferred because of un-
certainty regarding the identity of its t y p e species, G. muhlenbeckii, on
culms of Phragmites. This has now been fully redescribed b y Groves,
in Ganad. Journ. Bot. 43:1195-1276 (1965), where a key to all the
other recognised species can be found. Of British species formerly re-
ferred to Scleroderris, Groves would transfer S. ericae (Fries) Nann-
feldt to Encoeliopsis because of its 1-septate ascospores and S. bacilli-
fera (Karsten) Saccardo on Picea to Pragmopora as P. bacillifera
(Karsten) Rehm. According to Groves in Canadian J o u r n . Bot. 45:173
(1967) British material referred to this last species was mis determined.

Godronia ribis (Fries) Seaver in Mycologia 3 7 : 3 3 9 (1945). (PL XVIIIA.)

Apothecia erumpent in small groups from a common base, very


dark brown or almost black, with short stout stalks, finely rough, disc
concave, yellowish or greyish-brown, margin toothed. Asci cylindric-
clavate, up to 120x8/«, 8-spored, the pore blued b y iodine; ascospores
biseriate, elongated clavate, 25-35 x 3 - 4 /«, becoming 3-5-septate; para-
physes filiform, scarcely 1.5// thick at the tip.
On dead branches of Ribes, in spring. Uncommon.

Godronia uberiformis Groves in Ganad. J o u r n . Bot. 43:1245 (1965).


(Fig. 8c.)
Apothecia erumpent, usually in small clusters, urn-shaped, up to
1 m m . tall, externally almost black, covered with longitudinal fibrils
which p a r t to expose light grey or cream-coloured tissue round the
m o u t h , disc light grey, margin usually r a t h e r toothed. Asci cylindric-
clavate, up to 145x10 //, 8-spored; ascospores lying parallel, rod-shap-
ed, pointed below, up to 95x2/« and up to 11-septate; paraphyses
filiform.

The British material figured is on dead twigs of Ribes nigrum.


Related species to be sought in Britain are G. urceolijormis (Karsten)
Karsten in Acta Soc. Fauna Flora Fenn. 2 (6): 144 (1885), on Vaccinium
myrtillus & V. uliginosum and G. urceolus (Schmidt ex Fries) Karsten, op.
cit.: 144 (1885) on Betula. G. cassandrae Peck is regarded by Groves as a
plurivorous species with forms specialised on diverse host genera. Of these
f. callunae Groves occurs on Calluna in Scotland, f. betulicola Groves on
Betula in Germany and f. spiraeicola (Ilennings) Groves appears to be
widespread in northern Europe on Spiraea.

140
The remaining British species m a y be summarised as follows:
On Salix, with ascospores up to 60x3.5/«, G. juliginosa (Fries)
Seaver in Mycologia 37:344 (1945).
On C'alluna, with ascospores 60 jx or more x 2,5 [x G. callunigera
(Karsten). Karsten in Acta Soc. F a u n a Flora Fenn. 2 (6): 144 (1885).

Tribe Durelloideae
There is one genus:

DURELLA Tulasne

Durella Tulasne, Selecta F u n g o r u m Carpologia 3:177 (1865).


OC Lf >\fS ?
Apothecia small, saucer-snaped, sessile, black, formed of parallel
broad hyphae with brown thickened walls, asci 8-spored, ascospores in
most species 1-septate or multiseptate, hyaline paraphyses often
brown and swollen at the tip or embedded in a brownish m a t r i x . The
species are all either on, or immersed in, wood or woody stems. The
commonest British species is:

Durella commutata Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e


23-24:281 (1870). (Fig. 6E.)
Apothecia superficial, seated on a small circular base, smooth, black,
about 0.5 mm. across, disc black, flat with a low margin. Asci cylindric-
clavate, up to about 6 0 x 6 / / , 8-spored, pore not blued b y iodine; asco-
spores biseriate, elliptical or somewhat narrower below, 7 - 1 2 x 3 - 3 . 5 / / ,
hyaline, ultimately 1-septate; paraphyses cylindrical, about 2// thick,
tips embedded in an olivaceous m a t r i x . The dark parallel excipular
hyphae are shown x 6 6 0 in the lower left portion of Fig. 6E.

On decorticated wood, especially of Quercus, in winter and spring.

o .
1
J l n b e Heterosphaerioideae })• W

There are three somewhat doubtfully related genera, superficially


somewhat similar in their sessile cup-shaped blackish apothecia with
toothed margins:
I. Ascospores simple, non-septate Heterosphaeria (p. 142)
II. Ascospores muriform:
A. Apothecia regular, on Quercus Triblidium (p. 142)
B. Apothecia elongated, or Conifers . . . . Pseudographis (p. 143)

141
HETEROSPHAERIA Greville

Heterosphacria Greville, Scottish Cryptogamic Flora 2:103 (1824).


There is a single British Species:

Heterosphacria patella (Tode ex Fries) Greville, op. cit.: 103 (1824).


(PI. XVIIIH.)
Apothecia erumpent, becoming superficial by erosion of t h e host
tissue, sessile, subglobose, opening b y a pore, becoming cup-shaped
with t h e margin much inrolled when dry, purplish-brown to black,
with paler striae towards the margin when expanded, disc deeply con-
cave, about 1.5 m m . across, yellowish or grey. Asci cylindric-clavate,
up to 100x13/<, 8-spored, the pore blued by iodine; ascospores bi-
seriate, cylindrical with rounded ends, 12-18x4-5/«, hyaline, non-
septate; paraphyses broad, cylindrical hyaline, somewhat pointed
above.
On dead stems of Umbelliferae, especially in spring and summer. Very
common but more often found in its rather similar conidial state Hetero-
patella bonordenii (Hazslinszky) Lind, Danish Fungi:473 (1913). In Europe
races of this species occur on various other stems and two additional species
are recognised, on Linaria and Clematis respectively.

TRIBLIDIUM Rebentisch & Persoon

Triblidium Rebentisch ex Persoon, Mycologia Europaea 1:331 (1822).


Apothecia erumpent, sessile, leathery, globose, tearing open ir-
regularly and becoming cup-shaped, dark coloured, asci large and
thick-walled b u t not clearly bitunicate, ascospores uniseriate, very
large and multiseptate with b o t h transverse and longitudinal walls,
hyaline to yellowish.

Triblidium calyeiiforme Rebentisch ex Persoon, op. cit.:333 (1822).


(Fig. 8 G . )
Apothecia erumpent singly, leathery, tearing open by several ir-
regular teeth, smooth, black, sessile, disc deeply concave, grey, up t o
3 m m . across. Asci clavate, up to 270x25/«, 4-spored, thick-walled,
rounded above, contents deep red in iodine, no blue reaction; asco-
spores uniseriate, broadly elliptical, with a thick outer wall, about
seven to nine cross walls, each locule f u r t h e r subdivided by up to three
longitudinal walls, 30-48x 12-20 /«, hyaline or slightly yellowish; para-

142
physes slender, forked near the tip, colourless. The iodine reaction
suggests affinity with Pseudographis b u t the relationship of these two
genera to Heterosphaeria is dubious.
On bark of Quercus, in autumn. Rare.

PSEUDOGRAPHIS Nylander

Pseudographis Nylander apud Karsten, Mycologia Fennica 2 : 3 1 (1873).


Apothecia immersed, erumpent, elongated or angular, with black
margin and light coloured disc asci 4-8-spored, ascospores multisep-
t a t e , sometimes with longitudinal septa inserted, stained deep red-
purple by iodine. The place of publication is usually given as Herba-
rium Musei Fennici:96 (1859), b u t the name there is a nomen n u d u m .

Pseudographis pinicola (Rebentisch ex Fries) Rehm in 26th Ber. Natur-


hist. Vereins Augsburg: 10 (1881). (Fig. 11B.)
Apothecia gregarious, elliptical, or triangular, black with a promi-
nent smooth margin, disc up to 2.5 m m . long, flat, yellowish-brown.
Asci cylindric-clavate, not very thick-walled, with a thin-walled
rounded tip, up to 180x20//, 8-spored; ascospores uniseriate or bi-
seriate, elliptic-fusiform, or slightly curved, 2 2 - 3 8 x 7 - 9 / / , 1-7-septate,
hyaline to yellowish b u t instantly stained deep red-purple and opaque
by iodine; paraphyses slender, branched, about 2// wide.
On bark of Pinus and other conifers. Rare.

Pseudographis elatina (Acharius) Nylander apud Karsten, op. cit.: 31


(1873). (Fig. 11A.)
Apothecia gregarious, elliptical, often with pointed ends, black with
a prominent smooth margin, disc yellowish-brown, flat up to 2 m m .
long. Asci cylindric-clavate, not thick-walled, rounded above, up to
300x 15 //, 4-8-spored; ascospores uniseriate, elliptical or often strongly
constricted in the middle, 25-45 x 12-14 //, hyaline or slightly yellowish
b u t instantly stained deep red-brown b y iodine, with up to seven cross
walls and often a longitudinal wall as well; paraphyses slender, 2/<
thick.
On bark of conifers. Rare.

Tribe Polydesmioideae

This tribe is proposed to accommodate a small b u t distinctive genus


for which it is difficult to find a niche elsewhere in the system. It has

143
light-coloured pruinose discs with hymenia composed of relatively
large asci, containing large, hyaline, non-septate ascospores with two
or four oil drops, and very slender paraphyses much branched at their
tips. There is a superficial resemblance to apothecia of Ochrolechia among
the Lecanorales b u t the hymenium does not stain blue with iodine.

POLYDESMIA Boudier

Polydesmia Boudier in Bull. Soc. mycol. France 1 : 1 1 3 (1885).


Apothecia sessile, minutely tomentose, outer tissue composed of
slender interwoven h y p h a e whose free tips give the tomentose ap-
pearance, disc pruinose, asci 8-spored, ascospores large, multiguttulate
b u t not septate; paraphyses slender, branched near the tip. There is
one common British species:

Polydesmia pruinosa (Berkeley & Broome) Boudier, Hist. Class. Dis-


com. d ' E u r o p e : 100 (1907). (Fig. 9D.)
Apothecia sessile, saucer-shaped, white, downy with obtuse, smooth,
r a t h e r thick-walled, h y p h a l tips, about 3 / , wide; disc whitish, con-
spicuously white-pruinose with the protruding tips of the paraphyses,
flat or convex. Asci clavate, up to 100x12 /,, 8-spored, the pore blued
by iodine; ascospores biseriate, elliptic-fusiform or slightly kidney-
shaped, 1 4 - 2 0 x 4 - 6 / , , usually containing about four large oil drops but
not septate; paraphyses filiform, m u c h branched near the tip.

On dead twigs and branches, either visibly seated on large stromatic


Sphaeriales or situated over the ostioles of perithecia immersed in the bark,
throughout the year but especially in the winter months. Very common.

Family 6. Hyaloscyphaceae

The members of the Hyaloscyphaceae are characterised by the


presence of clearly differentiated hairs on the margin or outer surface
of the apothecium and, usually, by a r a t h e r soft texture, with the outer
flesh composed of thin-walled more or less prismatic cells. Some of the
smaller species, however, have the hairs very much reduced and m a y
be confused with the smaller Helotiaceae. It is, in fact, difficult to allot
some of the simplest and smallest of the Helotiales to an appropriate
family.

I. Apothecia without a subiculum:


A. Paraphyses with an abruptly swollen, septate, fusiform tip
Diplocarpa p. 146)

144
B. Paraphyses filiform or lanceolate, not enlarged at the tip:
1. Disc dark olive, surrounded by reddish hairs, apothecium ses-
sile, outer tissue composed of rounded cells
Trichodiscus (p. 146)
2. Not as above:
a. Paraphyses lanceolate, longer than the asci, or apothecia
with septate hairs, encrusted with fine granules, or the
hair walls distinctly brown, species often stalked:
a. Apothecia minute, sessile to short-stalked, hairs
short, white, obtuse, with smooth walls
Psilachnum (p. 155)
fl. Apothecia clothed with hairs in which the walls are
coloured or if hyaline are finely encrusted with
granules or bear crystals or are very long and
pointed.
*Ascus tip conical, pore plug usually blued by iodine
Dasyscyphus (p. 147)
**Ascus tip hemispherical, not blued by iodine
Perrotia (p. 154)
b. Rather similar apothecia but paraphyses filiform or
moniliform, hairs very long and rough, disc orange, on
bark of conifers and more or less erumpent
Lachnellula (p. 154)
c. Paraphyses not lanceolate, hairs usually non-septate,
smooth, usually hyaline, apothecia minute, sessile or
nearly so:
+ Hairs very thick-walled or even solid:
tBrittle glassy hairs, solid except at the base,
adpressed to the side of the cup
Unguicularia (p. 160)
ffSimilar hairs but with a slender lumen,
radiating from the margin of a soft pale
cup Urceolella (p. 160)
t t f H a i r s long, stiff, radiating, with thick outer
walls and numerous thin septa
Hyalopeziza (p. 159)
t t t f H a i r s long, undulating, with a narrow non-
septate lumen Hyalotricha (p. 159)
+ + H a i r s thin-walled, if solid at the tip then only
bristle-like:
fHairs contracted to a slender hooked bristle:
/Spores fusiform-clavate
Unguiculella (p. 161)
//Spores globose . . . Pithyella (p. 162)
t j H a i r s tapering evenly to a fine point
Hyaloscypha (p. 157)
t t f H a r i s cylindrical or clavate:
/Apothecia yellow or ochraceous
Phialina (p. 163)
//Apothecia whitish, grey or brownish:
'Hairs cylindrical or clavate:

145
§Hairs with an upward curve,
sometimes rough at the tip
like minute Dasyscyphus hairs
Cistella (p. 156)
§§Hairs not curved, septate, apo-
thecium brown on ferns
Microscypha (p. 163)
"Hairs thread-like, at the margin only
Mollisina (p. 164)
II. Apothecia seated on a widespreading, usually whitish, subiculum:
A. Tissue light coloured:
1. Ascospores 2-celled, hairs hyaline, thread-like
Arachnoscypha (p. 165)
2. Ascospores 4-8-celled, hairs coloured Arachnopeziza (p. 166)
B. Tissue olivaceous, hairs slender, hyaline Eriopeziza (p. 165)

DIPLOCARPA Massee
Diplocarpa Massee, British Fungus Flora 4:307 (1895).
There is a single British species:

Diplocarpa bloxami (Berkeley) Seaver in Mycologia 29:177 (1937).


(PL X X I K . )

Apothecia cup-shaped, dark olive-brown sometimes with a reddish-


brown margin, distinctly stalked, downy with short brown hairs, disc
concave, up to 3 m m . across, greenish or olive-coloured. Hairs cylindri-
cal, thin-walled, septate, somewhat pointed, smooth, up to about
1 0 0 x 3 / / . Asci cylindric-clavate, up to 7 0 x 7 / ( , 8-spored; ascospores
uniseriate, broadly elliptical, 6 - 9 x 2 . 5 - 3 / / , non-septate; paraphyses
very slender b u t expanded at t h e tip into a distinctive, thin-walled,
septate, fusiform body 2 0 - 3 0 x 3 - 7 / / .
On rotting wood, in spring and autumn. Rare.

TRICHODISCUS Kirschstein
Trichodiscus Kirschstein in Verh. Bot. Ver. Prov. B r a n d e n b u r g 66:25
(1924).
Again there is only one British species:

Trichodiscus prasinus (Quelet) Kirschstein, op. cit.:25 (1924). (PL


XXIL.)
Apothecia saucer-shaped, sessile on a broad base, disc dark olive-
green or nearly black, up to about 2 m m . across, margin and exterior

146
covered b y slender, pointed, thin-walled, reddish-brown hairs, up to
150x5/«, their walls encrusted b y reddish granules. Asci cylindric-
clavate, up to 4 0 x 4 f i , 8-spored; ascospores biseriate, fusiform,
13-15x2/«, non-septate; paraphyses cylindrical, 1.5-2 n wide, scarcely
longer t h a n t h e asci. The outer tissue from which the hairs arise is
composed of rounded dark cells like t h a t of the Dermateaceae.

On rotting monocotyledons in marshes, as on Carex, Digraphis, Glyceria


and Sparganium, May to August. Not uncommon.

DASYSCYPHUS S. F. Gray
Dasyscyphus S. F. Gray, N a t u r a l Arrangement of British Plants 1-670
(1821).
Apothecia usually more or less stalked, cup-shaped, always covered
with well differentiated hairs which fall into several different struc-
t u r a l groups roughly as follows:

1. Hairs thin-walled, obtuse, encrusted throughout with very


minute colourless granules. This is the largest group and the
hairs may be permanently hyaline, appearing white to the
eye, or become brownish with age.
2. Hairs similar to the above but thick-walled.
3. Hairs like group two in the lower part, but more slender, with
a slightly enlarged smooth head which secretes a gummy
substance.
4. Hairs smooth, long, tapering, like those of Hyaloscypha but
septate.
^ 5. Hairs smooth, thick-walled, brown, with a thin-walled glan-
dular tip.

Most species of Dasyscyphus have distinctly lanceolate paraphyses


whose tips protrude beyond the level of the asci b u t in some species
t h e y are slender and cylindrical or so narrow t h a t the lanceolate cha-
racter is difficult to make out. The asci are 8-spored, the pore blued b y
iodine, and in most species t h e ascospores are narrowly fusiform or
fusiform-clavate and non-septate. In a few species the ascospores are
needle-shaped, lying parallel in t h e ascus, and become multiseptate.
These have sometimes been separated to form a genus Erinella
Quelet, Enchiridion F u n g o r u m : 3 0 1 (1886), b u t this seems to be un-
tenable either taxonomically or nomenclatorially. A recent revision of
the British species will be found in Commonwealth Mycological In-
stitute Mycological P a p e r 32 (1949). A n u m b e r of common species
chosen to illustrate the scope of the genus will be described here.

147
Dasyscyphus pygmaeus (Fries) Saccardo, Sylloge F u n g o r u m 8:436
(1889). (PL XIXA.)
Apothecia gregarious or clustered, cup-shaped, becoming expanded
to expose a yellow disc up to 4 m m . across, exterior clothed with fine
downy hairs of type 1, up to 4 0 x 5 / / , stalk usually long and slender.
Asci cylindric-clavate up to 7 0 x 6 / / , 8-spored; ascospores biseriate,
narrowly fusiform, 7-12x1.5-2.5/«, non-septate; paraphyses lanceo-
late, 5 // thick, much longer t h a n t h e asci.

Attached to dead wood or branches just covered by soil or on dead roots


and grass rhizomes, often on Ulex, in spring and autumn. Not uncommon.
This species has been made the type of a genus Helolachnum Torrend in
Broteria 9:53 (1910), but it is a typical Dasyscyphus.

Dasyscyphus virgineus S. F. Gray, op. cit.:671 (1821). (Fig. 7A.)


Apothecia cup-shaped, long stalked, white and covered with white
hairs of type 1, disc cream-coloured, up to 1 m m . across. Asci cylindric-
clavate, up to 5 5 x 5 / / , 8-spored; ascospores fusiform or clavate,
6 - 1 0 x 1 . 5 - 2 . 5 / / , non-septate; paraphyses lanceolate, up*to 5/t wide,
considerably longer t h a n the asci.
On dead twigs, stems of Rubus fruticosus, cupules of Fagus, fallen cones,
scales of Betula catkins, dead stems of Calluna, PteriMum and vegetable
debris generally, especially in spring. Very common.

Dasyscyphus brevipilus Le Gal in Revue de Mycologie N.S. 4 : 2 6


(1939), differs in its slightly shorter hairs and in the paraphyses being
filiform.

Dasyscyphus nudipes (Fuckel) Saccardo, Sylloge F u n g o r u m 8:442


(1889). (PI. XIXB.)
Apothecia cup-shaped, short stalked, white, covered with white
hairs of t y p e 1, disc deep yellow, up to 1.5 m m . across. Asci cylindric-
clavate, up to 5 5 x 5 / / , 8-spored; ascospores biseriate, narrowly fusi-
form, 10-12x1.5-2/«, non-septate; paraphyses lanceolate, up to 5 / ,
wide, considerably longer t h a n the asci.
In swarms on dead herbaceous stems, especially those of Filipendula uL~
maria. May to August. Very common. This species is closely related to
D. virgineus from which it differs in its brighter coloured disc; small forms
with narrow ascospores, 6-11x1-1.5// are sometimes distinguished as var.
minor Dennis, Comm. Myc. Inst. Myc. Pap. 32:90 (1949).

Dasyscyphus controversus (Cooke) Rehm in 26th Ber. n a t u r h . Ver.


Augsburg:31 (1881), on dead culms of Phragmites and Digraphis is

148
distinguished b y turning pink and finally reddish-brown as it ages.
Several other minute species with a t e n d e n c y to redden have been
recognised on culms of the smaller grasses. Dasyscyphus pudibundus
(Quelet) Saccardo, Sylloge F u n g o r u m 8 : 4 3 3 (1889), is a reddening
species found on dead twigs, as of Salix and Crataegus. Dasyscyphus
sulphurellus (Peck) Saccardo is a very pale lemon-coloured species
apparently confined to dead twigs of Myrica gale.

Dasyscyphus palearum (Desmazieres) Massee, British Fungus Flora 4:


347 (1895). (PI. XIXG.)
Apothecia scattered, cup-shaped, long stalked, yellowish-brown,
drying darker, covered with light brown hairs of type 1, disc up to
1 m m . across. Asci cylindric-clavate, up to 5 0 x 4 //, 8-spored; ascospores
biseriate, fusiform, 7 - 1 4 x 1.5-2//, non-septate; paraphyses lanceolate,
5 fi wide, considerably longer t h a n the asci.
On ripe culms and stubble of wheat, and dead grass stems, such as Agro-
pyron and Ammophila, April to July. Fairly common. The hairs usually
have a short apical cell and may carry a few loose tabular crystals at the tip.

Dasyscyphus clavisporus Mouton in Bull. Soc. Roy. Bot. Belg. 56


(Pt. 2): 18 (1897). (Fig. 29G.)
Apothecia scattered, disc y 2 m m . diameter, concave, yellow, re-
ceptacle saucer-shaped, with short cylindrical stalk, buff, clothed with
cylindrical or slightly clavate, thin-walled, septate, white hairs,
45-70 x 4 - 5 //, with finely granulate walls. Asci 7 5 - 1 0 0 x 9 - 1 0 / / ,
8-spored; ascospores cylindric-clavate, pointed below, nonseptate,
1 5 - 2 0 x 3 - 4 / / , paraphyses very narrowly lanceolate, 1.5-2// wide, as
long as the asci.
On Juncus culms, said to be also on Agrostis and Molinia. July. Un-
common.
Distinguished by the broad clavate ascospores.

Dasyscyphus apalus (Berkeley & Broome) Dennis, Comm. myc. Inst.


myc. Pap. 32:25 (1949). (Fig. 7c.)
Apothecia cup-shaped, stalk slender, white, drying yellowish or
pinkish, covered with white hairs of type 1, disc pinkish, up to 1 m m .
across. Asci cylindric-clavate, up to 65x6/«, 8-spored; ascospores lying
parallel, needle-shaped, 3 5 - 4 5 x 1 . 5 / / , 1 - 7 - s e p t a t e ; paraphyses lanceo-
late, 4 (i wide, considerably longer t h a n the asci.
On dead culms and leaves of Juncus. October to April, occasionally in
summer. Very common.

149
During the summer months this species is usually replaced on dead
J uncus by Dasyscyphus diminutus (Roberge) Saccardo, Sylloge Fungorum 8:
449 (1889), which is subsessile with non-septate ascospores 10-15x1.5-2/«.
Dead Juncus also sometimes carries a very minute species, D. fugiens (Buck-
nail) Massee, British Fungus Flora 4:341 (1895), scarcely 200 ft across, with
short downy hairs and ascospores only 6-8x1.5 /«.

Dasyscyphus acuum (Albertini & Schweinitz ex Persoon) Saccardo,


Sylloge F u n g o r u m 8 : 4 4 3 ( 1 8 8 9 ) . ( P I . X X I H . )
Apothecia subsessile, white, drying pinkish-buff, covered with
downy, thin-walled, somewhat clavate, white hairs up to 30x4/«, disc
usually less t h a n 0.5 m m . across. Asci cylindric-clavate, up to 3 0 x 5 /«,
8-spored; ascospores biseriate, clavate, 4 - 5 x 1 - 1 . 5 / « ; paraphyses fili-
form, not noticeably longer t h a n the asci.
On fallen . n e e d l e s of JUaus, also on fallen cones, probably throughout the
year, but especially from autumn to spring. Common.
Clavidisculum acuum (A. & S. ex Persoon) Kirschstein in Ann. mycol. 36-
380 (1938) is a synonym.

Dasyscyphus pteridis (Albertini & Schweinitz ex Persoon), Massee,


British Fungus Flora 4 : 3 6 8 (1895), is an equally small b u t d a r k brown
species found on dead stems of Pteridium.
Dasyscyphus misellus (Roberge & Desmazieres) von Hohnel in S.B.
Akad. Wiss. Wien Abt. 1, 127:606 (1918), is a minute grey-brown spe-
cies found on fallen leaves of Rubus.

Dasyscyphus tricolor (Bulliard ex Merat) Fuckel in J a h r b . Nass. Vereins


f. N a t u r k u n d e 23-24:305 (1870). (PI. X I X c . )
Apothecia subsessile, cup-shaped, white and shaggy with long white
hairs of t y p e 2, disc yellow to orange, up to 2 m m . across, concealed b y
the incurved hairs when dry. Asci cylindric-clavate, up to 60x5/«,
8-spored; ascospores biseriate, fusiform, 6 - 1 0 x 1 . 5 - 2 / « , n o n - s e p t a t e ;
paraphyses lanceolate, 5 /< wide, considerably longer t h a n the asci.
On fallen twigs of Quercus in spring. Uncommon. More abundant is the
var. rubi (Bresadola) Dennis, Comm. Myc. Inst. Myc. Pap. 82:36 (1949),
figured, found on dead canes of Rubus idaeus.

Dasyscyphus fuscescens (Persoon) S. F. Gray, op. cit.:671 (1821). (PL


XIXH.)

Apothecia scattered, cup-shaped, short stalked, dark brown, with


short brown hairs of t y p e 1 b u t often bearing a mass of crystals at the

150
tip, disc dingy yellow, up to 1 m m . across. Asci cylindric-clavate, up to
4 5 x 5 f t , 8-spored; ascospores biseriate, fusiform, 6 - 9 x 2 - 2 . 5 f t , non-
septate; paraphyses lanceolate, 3 - 4 ft wide, m u c h longer t h a n the asci.

On fallen leaves of Quercus and Fagus, May to September. Common.


A var. fagicola (Phill.) Erenrns", op. cit.: 38 (1949), occurs commonly on fallen
cupules of Fagus.
Fallen leaves of Quercus also often bear up to five white species of Dasy-
scyphus, with hairs of types 1 or 2, details of which will be found in the
paper cited above.

Dasyscyphus clandestine (Bulliard ex Merat) Fuckel in J a h r b . Nass.


Vereins f. N a t u r k u n d e 2 3 - 2 4 : 3 0 5 ( 1 8 7 0 ) . (PI. X I X K . )
Apothecia scattered, cup-shaped, short stalked, light brown, covered
with yellowish-brown hairs of t y p e 1, each tipped b y a mass of cal-
cium oxalate, disc dingy yellow, up to 1 m m . across. Asci cylindric-
clavate, up to 50x5,«, 8-spored; ascospores biseriate, fusiform, 5 - 6 x
1.5 ft, non-septate; paraphyses lanceolate, 4/x wide, much longer t h a n
the asci.

On dead stems, as of Arctium, Chamaenerion. Cirsium. Evilobium and


especially Rubus idaeus, April to August but especially in spring. Common.

Dasyscyphus mollissimus (Lasch) Dennis E d . 1 : 9 5 (1960)


Peziza mollissima Lasch in Flora 41:651 (1858). (PI. X I X E . )
Apothecia sessile, completely covered with long yellow hairs, a
modified form of type 1 in which the superficial granules are larger and
easily detached, disc whitish, up to 2 m m . across. Asci cylindric-cla-
vate, up to 5 0 x 5 f t , 8-spored; ascospores biseriate, fusiform, 10-15x
1.5-2fx, non-septate; paraphyses lanceolate, 5 n wide, m u c h longer
t h a n t h e asci. This is the type species of Dyslachnum Clements Gen.
Fungi 1909.

On dead herbaceous stems, especially of Umbelliferae but also on Cam-


panula, Epilobium and Urtica, April to June. Common. Better known under
the later name D. leucophaeus (Persoon) Massee, British Fungus Flora 4:
351 (1895).

Dasyscyphus sulfureus (Persoon) Massee, British Fungus Flora 4 : 3 5 2


(1895), is another sessile yellow species found in a u t u m n on herbaceous
stems and distinguished b y its large ascospores, 2 5 - 3 5 x 2 fx and b y the
sap of the hairs often t u r n i n g purple with K O H solution.

151
Dasyscyphus pulverulentus (Libert) Saccardo, Sylloge F u n g o r u m 8 : 4 6 3
(1889). (PI. XIXD.)
Apothecia subsessile, bright yellow, covered with short yellow hairs,
resembling type 1 b u t often each tipped b y a lump of red resin, disc
yellow, up to 1 m m . across. Asci cylindric-clavate, 40x4/«, 8-spored;
ascospores biseriate, somewhat clavate, 4 - 5 x 1 / « ; paraphyses very
slender b u t usually longer t h a n the asci.
On fallen needles of Pinus, in spring and autumn. Less common than
D. acuum.
As viewed fresh, under a hand-lens, well developed specimens have a
barbaric beauty reminiscent of the enamelled bronzes of the late Celtic
period.

Dasyscyphus cerinus (Persoon) Fuckel in J a h r b . Nass. Yereins f.


N a t u r k u n d e 2 3 - 2 4 : 3 0 5 (1870). (PI. X I X F . )
Apothecia gregarious, globose, t h e n cup-shaped, subsessile, very
dark brown b u t concealed b y the dense golden-brown hairs of type 1,
which yield a purple stain in K O H solution, disc concave, yellowish,
up to 2 m m . across. Asci cylindric-clavate, up to 50x5/«, 8-spored;
ascospores uniseriate, unusually broadly elliptical for the genus,
4 - 5 x 2 - 2 . 5 / « ; paraphyses slender b u t mostly distinctly longer t h a n
t h e asci.

On decorticated wood of deciduous trees and shrubs, in spring and au-


tumn. Common.

Dasyscyphus niveus (Hedwig ex Fries) Saccardo, Sylloge Fungorum 8:


437 (1889). (Fig. 7B.)
Apothecia gregarious, cup-shaped, stalked, white at first b u t t u r n i n g
yellowish or buff with age and when dried, often bearing drops of fluid,
covered with hairs of t y p e 3 which tend to cohere at the tips, gummed
together b y their exudate, disc concave, white, becoming cream, up to
2 m m . across. Asci cylindric-clavate, up to 55x 5 /«, 8-spored; ascospor-
es biseriate, often slightly clavate, 6 - 1 0 x 1 . 5 - 2 / « ; paraphyses fili-
form, scarcely longer t h a n the asci.

On decorticated wood, characteristic of Quercus. October to April.


Common.

Dasyscyphus crystallinus (Fuckel) Saccardo, op. cit.:440 (1889), is a


similar species b u t with very long broadly lanceolate paraphyses and
often with loose balls of crystals among the hairs, found occasionally
on fallen twigs of Quercus.

152
Type 3 hairs are found also in D. distinguendus (Karsten) Saccardo,
op. cit.:434 (1889), on Populus wood and D. dryinus (Karsten) Sac-
cardo, op. cit.:435 (1889), on Betula wood, b o t h of which should be
sought in Britain, especially in t h e north.

Dasyscyphus albotestaceus (Desmazieres) Massee, British Fungus


Flora 4:346 (1895). (PI. X I X M . )
Apothecia erumpent, subsessile, reddish b u t covered with long,
straight, pale pink hairs of type 4, disc concave, flesh-coloured, up to
1 m m . across. Asci cylindric-clavate, up to 5 0 x 5 / / , 8 - s p o r e d ; ascospores
often biseriate, fusiform, 7-10x1.5-2/«, n o n - s e p t a t e ; paraphyses
lanceolate, 4// wide, much longer t h a n the asci.
On dead leaves of small grasses, as Bromus and Holcus. in summer. Un-
common.

Dasyscyphus acutipilus (Karsten) Saccardo, Sylloge F u n g o r u m 8:447


(1889). (Fig. 7D.)
Apothecia cup-shaped, very short stalked, white, covered with deli-
cate, long white tapering hairs of t y p e 4, disc concave white or cream-
coloured, scarcely 0.5 m m . across. Asci cylindric-clavate, up to
50x 5 //, 8-spored; ascospores biseriate, narrowly fusiform, 10-15 x 1.5 //,
non-septate; paraphyses lanceolate, 4/« wide, m u c h longer t h a n the
asci.

On grass culms in wet places, especially on Phragmiles and Digraphis,


May to August. Locally abundant. ~~

Dasyscyphus barbatus (Kunze) Massee, British Fungus Flora 4 : 3 6 1


(1895). ( P I . X I X I . )
Apothecia sessile, cup-shaped, brown, covered with long rust-brown
hairs of type 5, disc concave, up to 2 m m . across. Asci cylindric-clavate,
up to 6 0 x 5 / / , 8-spored; ascospores biseriate, fusiform, 9 - 1 1 x 2 / / , be-
coming 1 -septate; paraphyses lanceolate, 4 // wide, longer t h a n the asci.
On dead stems of Lonicera, throughout the year. Rare.

Dasyscyphus nidulus (Schmidt & Kunze) Massee, op. cit.: 104 (1895).
(PI. XIXJ.)

Apothecia sessile, cup-shaped, brown, covered with stiff brown hairs


of t y p e 5, disc yellowish, concave, about 1 m m . across. Asci cylindric-

153
clavate, up to 5 0 x 5 / , , 8-spored; ascospores biseriate, fusiform, 8 - 1 1 x
1.5/1, non-septate; paraphyses lanceolate, 5 y wide, much longer t h a n
t h e asci.

On dead herbaceous stems, as of Cirsium. Epilobium., Eupatorium, Fili-


penMa^enaM^^ am\_Polygonum, also on Carex and Scirpus, May to June
and also in autumn. Common, the only common British species with type
5 hairs.

PERROTIA Boudier

Perrotia Boudier in Bull. Soc. mycol. France 17:24 (1901).


Apothecia like those of a sessile Dasyscyphus b u t containing asci
with rounded tips in which there is no pore plug blued b y iodine. Hairs
usually brightly coloured, paraphyses filiform.

Perrotia flammea (Albertini & Schweinitz ex Fries) Boudier, Hist.


Class. Discom. E u r o p e : 6 6 (1907). (Fig. 20J.)
Apothecia scattered, saucer-shaped, sessile, 1 - 2 m m . diameter, disc
reddish-brown, receptacle concealed b y a dense coating of cinnabar red
hairs, each rather thick-walled, slightly tapering to a rounded tip, up
to 2 0 0 x 4 - 5 / , , containing a sap which t u r n s purple in potash solution
and having t h e wall encrusted with coarse orange granules. Asci
cylindrical with rounded tips, 8 0 - 1 0 0 x 7 - 1 0 / , , 8-spored; ascospores
biseriate, cylindrical with rounded ends and often somewhat curved,
1 0 - 1 4 x 2 . 5 - 3 / , , becoming 1-septate, paraphyses cylindrical, 2—2.5
thick, no longer t h a n the asci.

On dead twigs and branches, as of Corylus, Ligustrum, Prunus, Salix.


Rare.

LACHNELLULA Karsten

Lachnellula Karsten in Medd. Soc. F a u n a Flora Fennica 11:138 (1884).

Apothecia similar to those of a sessile Dasyscyphus b u t with cylin-


drical or moniliform paraphyses, disc always orange, surrounded b y
dense shaggy hairs of t y p e 1 structure, always on the bark of conifers.
There are about six British species, which look alike to the eye and are
distinguished mainly b y their spores. The t y p e s species, L. suecica
(Fuckel) Nannfeldt, with spherical ascospores 4 - 5 /, diameter, common
on conifers in central and northern Europe is not known in Britain.

154
The name Trichoscyphella Nannfeldt (1932) has been used for those
species with elliptical to acicular ascospores, including:

Lachnellula hahniana (Seaver) Dennis in Persoonia 2:184 (1962).


(Pi. XIXL.) _ i r - . -,
Apothecia erumpent subsessile, often in small groups, at first glo-
bose, becoming cup-shaped, snow-white, shaggy w i t h long, slender,
septate, thin-walled, finely roughened hairs, disc concave or almost
flat, orange, up to 3 m m . across. Asci cylindric-clavate, up to 156x 12 /«,
8-spored; ascospores uniseriate, elliptic-oblong, mostly 13-20x5-6/«,
non-septate; paraphyses filiform b u t mixed with others much swollen
between the septa and "moniliform".

On dead twigs and fallen branches of Larix. throughout the year. Very
common, occasionally on other conifers. The species is a harmless sapro-
phyte.
Lachnellula willkommii (Hartig) Dennis, op. cit.: 184 (1962), is a
morphologically very similar species found around cankers on Larix
and the cause of a serious disease in this tree.
Lachnellula subtilissima (Cooke) Dennis, op. cit.: 184 (1962), though
similar to the eye has very different ascospores, 6 - 1 1 x 2 - 2 . 5 / « and
occurs mainly on Pinus.
* Lachnellula pseudofarinacea (Crouan) Dennis, op. cit.: 184 (1962),
on branches of Pinus, has long needle-shaped ascospores, 7 0 - 8 0 x 1.5 /«.

P S I L A C H N U M von Hohnel

Psilachnum von Hohnel in Mitt. bot. Inst, techn. Hochsch. Wien 3:73
(1926).
Hairs short, thinwalled, cylindrical, obtuse, without superficial
granules or apical granules, paraphyses lanceolate as in Dasyscyphus,
of which this is perhaps a reduced development.

Psilachnum asemum (Phillips) comb.nov.


Lachnella asema Phillips in Grevillea 19:73 (1891). (Fig. 29F.)
Apothecia minute, olive-brown, short-stalked, clothed with white
hairs 3 0 - 4 5 x 3 - 4 / / , 1-3-septate, r a t h e r tapered above to t h e obtuse

* Madame Le Gal in Revue de Mycologie N.S. 18:98 (1953), writes "nous


ne voyons pas qu'on puisse, actuellement, classer l'espece des Crouan ailleurs
que parmi les Trichoscyphella" but did not actually print the necessary
combination.

155
tip. Asci 36-45x5-7/«, spores elliptical 8 - 1 5 x 2 - 3 / « , paraphyses lan-
ceolate, 4-5/( wide longer t h a n the asci.

On dead Carex. Probably common.

Dasyscyphus helotioides (Rehm) Dennis is a synonym.


Psilachnum lateritio-album (Karsten) von Hohnel, op. cit.: 73 (1926)
on Scirpus, differs in having only fine marginal hairs and ascospores
only 6 - 8 x 1.5 /«.
P. granulosellum von Hohnel, op. cit.: 74 (1926) on Carex has very
short clavate hairs, 20-30 x 4 - 6 /«, 0 - 1 -septate, with spores 5 - 7 x 1.5-2 //,
and is white t h r o u g h o u t .
P. inquilinum (Karsten) Dennis in Persoonia 2:182 (1962), also
white, is on Equisetum, with cylindrical obtuse hairs 50x2.5// and
spores 6 - 9 x 1 . 5 - 2 / « .

CISTELLA Quelet

Cistella Quelet, Enchiridion F u n g o r u m : 3 1 9 (1886), emend Nannfeldt


in Nova Acta Reg. Soc. Sci. Upsal. IV, 8:265 (1932).

This is a genus of d o u b t f u l validity*, containing an assemblage of


species some of which appear akin to Pezizella while others are
scarcely separable from minute species of Dasyscyphus, like D. acuum,
except in growing on wood. Two species will suffice to illustrate the
genus here:

Cistella perparvula (Karsten) Nannfeldt, op. cit.: 2 7 0 (1932). (PI. X X K . )

Apothecia superficial, subsessile, shallow cup-shaped, pale buff b u t


covered with m i n u t e downy white hairs towards the margin, disc al-
most flat, cream-coloured, less t h a n 0.5 m m . across. Hairs clavate, thin-
walled, non-septate, hyaline, about 20x4/«; asci clavate, up to 70x8/«,
8-spored; ascospores biseriate, elliptic-cylindric, 7 - 1 1 x 2 - 2 . 5 / / , non-
septate; paraphyses, if any, filiform.

On bark of deciduous trees in spring and autumn. Rare, or at least, seldom


seen.

*It is also a later homonym of Cistella Blume 1825 (Orchidaceae).

156
Cistella piceae (Henn.) comb. nov. var. laricinum (Velenovsky)
Dennis. (Fig. 29c.)
Belonium piceae Hennings in Hedwigia 4 3 : 7 1 (1904) var. laricinum
Velenovsky Monog. Discom. Bohem.: 177 (1934).
Apothecia scattered, superficial, white, disc % m m . diameter, con-
cave, receptacle cup-shaped, sessile on a small brown base, minutely
pruinose with unicellular, clavate, hyaline hairs about 1 5 x 5 /«, with
thin smooth walls. Asci 5 0 - 6 0 x 8 - 1 0 n , pore not blued b y iodine,
8-spored, spores elliptic cylindric, straight or slightly curved, becoming
1-septate, a few 2-septate, not constricted, hyaline 7 - 1 1 x 2 . 5 - 3 . 5 [x,
paraphyses often forked, enlarged up to 5 p at the tip.

On fallen twigs of Larix. March to October. Rather common.


This is distinctive, among the swarm of minute sessile, whitish Hyalo-
scyphaceae, on account of its rather broad regularly septate ascospores.

HYALOSCYPHA Boudier
Hyaloscypha Boudier in Bull. Soc. mycol. France 1 : 1 1 8 (1885), emend
Nannfeldt in Nova Acta Reg. Soc. Sci. Upsal. IV, 8:272 (1932).
Apothecia minute, subsessile, soft t e x t u r e d , clothed with delicate,
thin-walled, tapering, pointed hairs, non-septate or occasionally with
a short basal cell only, asci usually 8-spored, usually with the pore
blued b y iodine b u t this m a y become a p p a r e n t only in dried material
and the character should not be stressed, paraphyses filiform, some-
times very scanty, no longer t h a n the asci. The n u m b e r of British
species is uncertain b u t the following are among the commonest and
most distinctive; see f u r t h e r Commonwealth Mycological I n s t i t u t e
Mycological Paper 32.

Hyaloscyphahyalina (Persoon) Boudier, Hist. Class. Discom. d ' E u r o p e :


127 (1907). (Fig. 7E.)
Apothecia scattered, cup-shaped, sessile on a small base, white or
hyaline, translucent when moist, covered, especially near the margin,
b y slender, pointed, thin-walled, non-septate hairs, up to 3 0 x 3 / / ; disc
concave, up to 0.5 m m . across. Asci cylindric-clavate, up to 35x6/«,
8-spored; ascospores biseriate, elliptical, 6 - 1 0 x 2 - 3 / / ; paraphyses
filiform.

In swarms on dead branches and decorticated wood of Quercus, through-


out the year. Very common. Allied forms occur on bark and wood of other
trees.

157
Hyaloscypha leuconica (Cooke) Nannfeldt in Trans. Brit, mycol. Soc
20:206 (1936). (Fig. 7F.)
Apothecia gregarious, subsessile, cup-shaped on a small base, white,
drying yellowish, covered, especially towards the margin, with long
slender, spreading white hairs, up to about 100x7/«, tapering evenly
to a sharp point, non-septate; disc concave, scarcely 0.5 m m . across.
Asci cylindric-clavate, up to 4 0 x 5 / / , 8-spored; ascospores biseriate,
cylindric-elliptic, 6 - 7 x 2 - 2 . 5 / « , non-septate; paraphyses filiform.

On decorticated wood, typically of conifers and on fallen pine cones, but


scarcely distinguishable fungi occur also on dead twigs of deciduous trees,
Fagus cupules and the like, throughout the year. Common.

Hyaloscypha dematiicola (Berkeley & Broome) Nannfeldt in Trans.


Brit, mycol. Soc. 2 0 : 2 0 5 ( 1 9 3 6 ) . (PI. X X B . )
Apothecia scattered amongst a short black mould, sessile, cup-
shaped, brown, bearing, especially near the margin, pointed non-sep-
t a t e brown hairs, up to 50x4/«; disc concave, pallid, about 0.2 m m .
across. Asci cylindric-clavate, about 40x5/«, 8-spored; ascospores bi-
seriate, elliptic-fusiform, 5 - 8 x 2 - 3 / « , often with two oil drops b u t non-
septate; paraphyses filiform.

On decorticated wood, especially of Quercus, amongst the mould Haplo-


graphium delicatum Berkeley & Broome, which is its conidial state. Probably
common but seldom seen.

Hyaloscypha flaveola (Cooke) Nannfeldt in Trans. Brit, mycol. Soc.


23:252 (1939). (Fig. 29A.)
Apothecia scattered, bright yellow t h r o u g h o u t , about y 3 m m . across,
cup-shaped, sessile, minutely downy with thinwalled hairs 20-35/«
long, having a bulbous base 3-4/« wide tapered to a whip-like apex.
Asci 2 5 - 3 5 x 5 - 7 /«, 4-8-spored, spores fusiform, 10-14x 2 /«, paraphyses
cylindrical.

Amongst the hairs on under sides of rotting fronds of Pteridium aqui-


linum. August and September. Uncommon, much less common than Micro-
scypha grisella on this substrate.

Hyaloscypha curvipila Grelet in Revue de Mycologie N.S. 16:92


(1951), a minute white species on debris of conifers, has curiously
hooked hairs approaching those of Unguiculella, p. 161.

158
HYALOPEZIZA Fuckel

Hyalopeziza Fuckel emend von Hohnel in S.B. Akad. Wiss. Wien


A b t . 1 , 1 1 1 : 1 0 0 3 (1902).

Apothecia similar to those of Hyaloscypha b u t bearing stiff, spread-


ing, r a t h e r thick-walled, multiseptate hairs. There is a single British
species:

Hyalopeziza ciliata Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:


298 (1860). (Fig. 7K.)

Apothecia scattered, sessile on a narrow base, cup-shaped, hyaline-


white, drying yellowish, bearing r a t h e r sparse, stiff, radiating multi-
septate, pointed, hyaline hairs, up to 2 0 0 x 6 / / ; disc concave, up to
0.3 m m . across. Asci cylindric-clavate, up to 3 0 x 5 / / , 8-spored; asco-
spores often biseriate, elliptical, 4 - 5 x 1 . 5 - 2 / / , non-septate; paraphyses
filiform.

On dead and fallen leaves of Carpinus and Acer, in autumn. Perhaps rare.

HYALOTRICHA Dennis

Hyalotricha Dennis, Comm. Myc. Inst. Myc. P a p . 32:75 (1949).

Apothecia small, subsessile, covered with long, flexuous hairs with


thick hyaline walls b u t w i t h t h e lumen still clearly distinguishable,
asci 8-spored, ascospores non-septate, hyaline, paraphyses filiform.
There is one British species:

Hyalotricha corticola Dennis, op. c i t . : 75 (1949). (PI. XXJ.)

Apothecia gregarious, cup-shaped with a minute stalk, dark oliva-


ceous b u t concealed b y the dense growth of long white hairs, disc
concave, about 0.2 m m . across. Hairs flexuous, obtuse, smooth, up to
about 120x3/«, t h e lumen stained dark red b y iodine and sometimes
divided b y a few t h i n septa. Asci cylindric-clavate, up to 2 5 x 4 / / ,
8-spored; ascospores biseriate, narrowly fusiform, 4 . 5 - 6 x 1 / / ; p a r a -
physes filiform, 1 ji thick, as long as the asci.

On dead branches of Myrica gale, records on Betula are in error, May to


September. Locally abundant.

159
URCEOLELLA Boudier
Urceolella Boudier, in Bull. Soc. mycol. France 1 : 1 1 9 (1885).
Apothecia small, sessile, yellowish or light brown, ornamented round
t h e margin b y radiating, white, non-septate hairs with very thick
glassy walls, asci 8-spored, ascospores non-septate, paraphyses fili-
form. The hairs are not stained b y iodine.

Urceolella crispula (Karsten) Boudier, Hist. Class. Discom. d ' E u r o p e -


130 (1907). (PI. XXD.)
Apothecia sessile on a broad base, subglobose, white to cream-
coloured, very soft t e x t u r e d , ornamented b y r a t h e r widely spaced
glassy hairs which, towards the margin tend to be arranged tangenti-
ally and to cling together in small groups, giving a curious cog-wheel
appearance, disc up to 0.3 m m . across, deeply concave. Asci cylindric-
clavate, up to 3 0 x 5 n, 8-spored, ascospores fusiform-clavate, biseriate,
6 - 1 0 x 1 . 5 - 2 / « ; paraphyses filiform, as long as the asci.

On dead herbaceous stems in summer and autumn. Uncommon.

Peziza trichodea Phillips & Plowright in Grevillea 3:125 (1875), on


fallen needles of Pinus, belongs in this genus rather t h a n in Hyalo-
tricha, where I formerly placed it, or Dasyscyphus as proposed by
Boudier.

UNGUICULARIA von Hohnel

Unguicularia von Hohnel in Annales mycologici 3:404 (1905).


Apothecia small, sessile, urn-shaped, soft t e x t u r e d , covered with
glassy hairs, a p p a r e n t l y solid except for a small thin-walled basal
portion at which t h e y easily snap, asci 8-spored, ascospores non-sep-
t a t e , paraphyses filiform. The n u m b e r of good species is uncertain as
these minute fungi have been little studied and are usually overlooked
b y collectors.

Unguicularia cirrhata (Crouan) Le Gal in Revue de Mycologie N S 18 •


99 (1953). (PI. XXA.)
Apothecia urn-shaped, sessile on a small base, buff, up to 0.2 m m .
across, covered especially towards the margin b y glassy, brittle, obtuse
hairs, up to 40x5/«, closely adpressed to the cup, with a thin-walled
basal portion and the remainder apparently solid and stained reddish-

160
purple b y iodine; asci cylindric clavate, up to 30x5/«, 8-spored;
ascospores biseriate, fusiform-clavate, 5 - 6 x 1 . 5 / « ; paraphyses filiform,
no longer t h a n the asci.
On dead herbaceous stems, typically on Eupatorium, especially at the
base of those still standing in damp situations. U. scrupulosa (Karsten) von
Hohnel, typically on canes of Rubus idaeus, is probably a synonym. Com-
mon, throughout the year. Similar but possibly distinct fungi occur on
decorticated wood.
It is doubtful if Unguicularia can be maintained as genus distinct from
Urceolella unless the difference in iodine reaction of the hairs is shown to be
constant and significant.

U N G U I C U L E L L A von Hohnel

Unguiculella von Hohnel in S.B. Akad. Wiss. Wien. Abt. 1, 115:281


(1906).
Apothecia subsessile, covered with slender hairs with a thin-walled
cylindrical base and a slender, solid, bristle-like, hooked tip, asci 4- or
8-spored, ascospores non-septate, paraphyses either resembling the
hairs or filiform. There are about three known British species, all
apparently uncommon.

Unguiculella eurotioides (Karsten) Nannfeldt in Trans. Brit, mycol.


Soc. 20:194 (1936). (PI. X X c . )
Apothecia shallow cup-shaped, subsessile on a small base, yellowish,
disc up to 0.3 m m . across, covered externally w i t h hyaline hairs,
cylindrical, thin-walled and 3 - 4 p wide below, suddenly narrowed to
a fine bristle hooked at the tip. Asci cylindric-clavate, about 40x6/«,
8-spored; ascospores biseriate, fusiform, 5 - 8 x 2 - 2 . 5 / « , non-septate;
paraphyses slender, tipped b y hooked bristles like those of the hairs.

On dead herbaceous stems in autumn. Rare.


The Unguiculella hair type seems to be a modification of that found in
Hyaloscypha and the two genera may not be truly separable.

Unguiculella hamulata (Feltgen) von Hohnel in Sitzber. K. Akad.


Wiss. Wien, Math.-nat. Kl. 115 Abt. 1:1280 (1906). (Fig. 29E.)
Apothecia minute, up to 200 /« diameter, whitish, minutely downy,
with hairs 20-30 /« long, having a cylindrical base 3 p. wide a b r u p t l y
contracted to a long hooked apex scarcely 1 /« wide. Asci 20-25x5-6/«,
8-spored, spores biseriate, broadly elliptical, 5 - 6 x 2 - 2 . 5 / « paraphyses

161
filiform. The figure shows the very minute apothecia amongst pseudo-
thecia of Leptosphaeria acuta.

On dead stems of Urtica and Trifolium. May-June. Perhaps not un-


common but seldom seen. This is the same as U nguicularia jalcipila von
Hohnel but not the same as Mollisia hamulata Rehm, which was a much
larger yellow Unguiculella on Cirsium, the attempted transfer of which to
Unguiculella by von Hohnel in 1918 is blocked by the 1906 name above.

Unguiculella robergci (Desmazieres) Dennis in Kew Bulletin (10): 136


(1955). (Fig. 29D.)
Apothecia scattered, sessile, disc dark red, scarcely 1 mm. diameter,
receptacle saucer-shaped, downy with hairs 35-55// long, 3 - 4 / / wide
at the base, a b r u p t l y contracted to a long curved solid tip scarcely 1 /(
wide. Asci 3 5 x 5 - 7 / / , spores biseriate, elliptical, 5 - 8 x 2 . 5 / / , with 2 oil
drops, paraphyses cylindrical 2// wide.
On bark of Lonicera, Rosa & c. April. Rare.

PITHYELLA Boudier

Pithyella Boudier in Bull. Soc. Mycol. France 1:118 (1885).


Apothecia like those of Unguiculella b u t with globose ascospores.

Pithyella ilicincola (Berkeley & Broome) Boudier, Hist. Class. Discom.


Europe: 125 (1907). (Fig. 20B.)
Apothecia scattered, superficial, sessile, shallow-cupshaped, 0.3 m m .
across, pinkish buff with downy white hairs, tissue of the receptacle
composed of subglobose or polygonal cells with t h i n red-brown walls,
bearing hyaline hairs 2 0 - 4 0 x 4 - 6 /«, narrowed above to a solid recurved
beak like those of Unguiculella. Asci clavate, 3 0 - 4 0 x 5 - 7 / / , 8-spored;
ascospores 1 - 2 seriate, globose, hyaline, 4 - 5 // diameter, with a large
central oil drop; paraphyses slender, cylindrical, obtuse.

On stromata of Myriangium and other fungi.


There is some uncertainty about the appropriate generic name for this
fungus. It is commonly referred to Mollisiella Massee, British Fungus Flora
4:221 (1895), but that is a later homonym of Mollisiella Boudier, op. cit.
(1885). The type species of Pithyella is Mollisia hypriina Quelet which I have
not seen but which was figured by Boudier as having thin-walled, slender,
cylindrical, septate hairs. If this be regarded as generically distinct from
P. ilicincola the genus for the latter will be Unguiculariopsis Rehm in An-
nales Mycologici 7:401 (1909).

162
MICROSCYPHA H . & . P. Sydow

Microscypha H. & P. Sydow in Annales mycologici 17:38 (1919).


Apothecia small, dark, soft, short-stalked, bearing short obtuse, thin-
walled, non-septate hairs, asci 8-spored, ascospores non-septate, para-
physes slender. There is one common British species:

Microscypha grisella (Rehm) Sydow, op. cit.:38 (1919). (PL XXG.)

Apothecia cup-shaped with a short cylindrical stalk, brown with a


whitish margin due to the light-coloured fringe of hairs, which measure
up to 6 0 x 3 / ( , disc concave, light grey or yellowish, up to 0.4 m m .
across. Asci cylindrical, pointed above, 8-spored, up to 40x7/«; asco-
spores biseriate, fusiform-clavate, 6 - 1 0 x 1 . 5 - 2 / « , non-septate; para-
physes filiform, as long as the asci.

On the under surface of decaying Pleridium aquilinum fronds but largely


hidden by the hairs of the fern, in spring and summer. Very common.
Microscypha eurhizus Graddon in Trans. Brit, mycol. Soc. 50: 12
(1967) occurred on dead leaves of Ilex aquijolium in J a n u a r y .

PHIALINA von Hohnel

Phialina von Hohnel in Mitt. Bot. Inst. Techn. Hochsch. Wien 3 : 6 1


(1926).
Apothecia sessile or nearly so, yellow or ochraceous, minute, outer
tissue formed of thin-walled prismatic cells which run out into
cylindrical, smooth, thin-walled, non-septate hairs like those of Micros-
cypha, asci 4- or 8-spored, ascospores narrowly fusiform, 0-1-septate,
paraphyses filiform.

Phialina puberula (Lasch) von Hohnel, op. cit.: 106 (1926). (PL XXH.)

Apothecia cup-shaped, sessile on a small base, bright yellow with a


dark basal ring of cells, covered with paler, downy, non-septate, thin-
walled, smooth, obtuse hairs about 25x2-4/«, often containing drops
of yellow oil, disc flat, lemon-yellow, up to 0.5 m m . across. Asci
cylindric-clavate, up to 6 0 x 7 y, 8-spored, ascospores biseriate, narrow-
ly fusiform, 10-14x1.5-2/«, o f t e n slightly curved, 1-septate; para-
physes filiform 1 - 2 /t thick.
Scattered or in groups on fallen leaves of Quercus, in autumn. Common.
This most beautiful and distinctive species has been referred to many
genera and even its specific name is disputed for it seems likely it was this

163
that Greville called Peziza puncliformis in Scottish Cryptogamic Flora 2:63
(1824). If that is so the legitimate name of the genus is probably Calycellina
von Hohnel in S.B. Akad. Wiss. Wien Abt. 1, 127:601 (1918).

Phialina ulmariac (Lasch) Dennis E d . 1:102.

Peziza ulmariae Lasch in Flora 3 6 : 2 0 1 (1853). (PL XXF.)

Apothecia scattered, sessile, saucer-shaped, up to 0.5 m m . across,


ochraceous, minutely downy towards the margin with thin-walled
hairs about 1 5 x 2 / / . Asci cylindric-clavate, up to 3 0 x 5 / / , 4-spored;
ascospores biseriate, narrowly fusiform, 10-13x1.5/«, non-septate;
paraphyses cylindrical, about 2 // thick, usually containing drops of
yellow oil.
On dead stems of Filipendula ulmaria, especially in summer. Very com-
mon.
This species was referred to Hyaloscypha by Nannfeldt.

MOLLIS IN A von Hohnel

Mollisina von Hohnel in Mitt. Bot. Inst. Techn. Hochsch. Wien 3:67
(1926).
Apothecia small, soft-fleshed, t u r b i n a t e , light-coloured, tissue formed
of thin-walled angular cells, bearing at the margin only, slender, short,
hyaline processes, asci 8-spored, ascospores non-septate, paraphyses
filiform.

Mollisina acerina (Mouton) von Hohnel, op. cit.:67 (1926). (Fig. 7L.)
Apothecia scattered, t u r b i n a t e , soft-fleshed, sessile on a small base,
hyaline-white, drying grey-brown with a whitish margin, disc flat, up
to 0.3 m m . across. Marginal processes hyaline, about 5 - 1 0 x 0 . 5 - 1 / / .
Asci cylindric-clavate, up to 4 5 x 7 / / , 8-spored; ascospores biseriate,
elliptic-fusiform, 5 - 8 x 2 . 5 - 4 / « ; paraphyses cylindrical, up to 2/« thick
at the tip.

On rotting leaves, as of Acer pseudoplatanus, in autumn. Apparently un-


common.
There are in nature many minute white virtually hairless Helotiales,
mostly on dead leaves, which may probably be referable to Mollisina but
which have been little studied owing to their minute size and lack of any
very distinctive characters.

164
E R I O P E Z I Z A (Saccardo) Rehm

Eriopeziza (Saccardo) Rehm in Rabenhorst's K r y p t . Flora Deutsch-


land 1 (3): 695 (1892).

Apothecia sessile or nearly so, seated on a thick, cobweb-like,


whitish mycelium, t h e subiculum, downy with slender hyaline hairs,
asci 8-spored, ascospores non-septate, paraphyses filiform, branched.
There is one British species:

Eriopeziza caesia (Persoon) Rehm, op. cit.:696 (1892). (PI. X X i . )

Apothecia cup-shaped with a narrow stem-like base, olive-brown b u t


concealed b y the slender whitish hairs, about 2 - 3 n thick, disc blue-
grey, concave, up to 0.4 m m . across. Asci clavate, up to 35x5/«,
8-spored; ascospores irregularly biseriate, narrowly fusiform, 5 - 6 x
1 - 2 n , non-septate; paraphyses filiform, up to 1.5/« thick, branched.
The subicular hyphae are encrusted with coarse colourless granules.

On rotting decorticated wood of Quercus, in swarms, throughout the year


but chiefly in the winter months. Very common.

ARACHNOSCYPHA Boudier

Arachnoscypha Boudier in Bull. Soc. mycol. France 1:118 (1885).

Apothecia sessile on a fine white subiculum, downy with smooth,


slender, hairs, asci 8-spored, ascospores 1-septate, hyaline, paraphyses
filiform, branched. There is one British species:

Arachnoscypha aranea (de Notaris) Boudier apud Dennis, Comm. Myc.


Inst. Myc. Pa. 32:87 (1949). (Fig. 7G.)

Apothecia cup-shaped, sessile, white, the margin fringed with


smooth, flexuous, thin-walled, hyaline hairs up to 40x2,«, tapering to
a point, disc concave, yellowish, u p to 0.3 m m . across. Asci cylindric-
clavate, up to 6 0 x 6 n , 8-spored; ascospores biseriate, elliptical, be-
coming 1-septate, 7 - 8 x 2 - 2 . 5 / / ; paraphyses very slender, forked.

On the inner surface of decaying cupules of Castanea, in autumn. Un-


common.

165
ARACHNOPEZIZA Fuckel
Arachnopeziza Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24-
303 (1870).
Apothecia gregarious, sessile, cup-shaped, seated on a light-coloured
subiculum, covered with thin-walled septate hairs, asci 8-spored, asco-
spores multiseptate, paraphyses filiform, often branched.

Arachnopeziza aurata Fuckel, op. c i t . : 3 0 4 (1870). (PL XXE.)

Apothecia turbinate, sessile on a narrow base, golden-yellow, covered


with smooth tapering hairs up to 100x6,«, disc up to 0.4 m m . across,
subiculum formed of whitish hyphae encrusted with numerous
granules. Asci clavate, up to 9 0 x 1 1 /<, 8-spored, ascospores needle-like,
lying parallel in the ascus, 55-75x1.5-3/«, multiseptate; paraphyses
filiform, about 1 fi thick, forked.

On dead wood in spring and autumn. Not uncommon. A white state of


this species has been called A. nivea Lorton in Bull. Soc. mvcol France 30
224 (1914).

Arachnopeziza aurelia (Persoon) Fuckel, op. cit.:303 (1870), differs


in its larger apothecia, up to 2 m m . across, and 3-septate ascospores
1 2 - 2 0 x 3 - 5 ,i.
Arachnopeziza obtusipila Grelet in L ' a m a t e u r Champ. 8 (3): 45
(1922), a minute white species with 3-septate ascospores 1 6 - 2 6 x 2 . 5 -
3.5 fi has been found on fallen cones of Larix in Yorkshire. For a full
account of this and other European species see Korf in Lloydia 14
129-180 (1951).

Family 7. Orbiliaceae

Apothecia small, subsessile, mostly on wood or b a r k , occasionally


on the woodier herbaceous stems, light-coloured, having a w a x y
consistency and often translucent when dresh, the flesh composed of
large, rounded or angular, very thin-walled cells, asci exceptionally
small, often with a characteristically forked base and t r u n c a t e tip, not
blued b y iodine, ascospores usually small to very small, hyaline, non-
septate. There are two British genera:

I. Paraphyses abruptly swollen at the tips, apothecial margin quite even


Orbilia (p. 167)
II. Paraphyses not swollen abruptly, apothecial margin dentate with co-
hering downy hairs Hyalinia (p. 169)

166
The genus Habrostictis in the Dermateaceae, easily recognised b y the fusi-
form tips to the paraphyses, has a somewhat Orbilioid aspect and might be
sought in this family.

ORBILIA Fries

Orbilia Fries, Floram Scanicam:343 (1835).


The characters of the genus are sufficiently indicated above and
with a little practice there is no difficulty in recognising an Orbilia in
the field. Differentiation of the species is, however, m u c h more difficult.
The essential characters, a p a r t from colour, are largely those of the
ascospores, which are small and often hard to see, especially as asci
and paraphyses cohere firmly to give the w a x y t e x t u r e peculiar to the
family. Moreover the British species have never been critically studied
and there is confusion about the application of some old specific names
by various continental authorities. The following are among the best
known British representatives b u t the genus is one of those which most
needs monographic s t u d y on modern lines.

Orbilia xanthostigma (Fries) Fries, S u m m a Veg. Scand. Sect. Post.: 357


(1849). (PL XXIA.)
Apothecia subsessile on a small base, waxy, smooth, golden-yellow,
flat or convex when moist, disc about 1 m m . across, margin inrolled
when dry. Asci cylindric-clavate, up to about 4 0 x 4 / / , 8-spored, asco-
spores uniseriate, slightly kidney-shaped, 3 - 4 x 1 - 1 . 5 / « with two oil-
drops; paraphyses slender slightly longer t h a n the asci, a b r u p t l y
swollen to an apical knob 2 - 3 /« across.

In swarms on decorticated wood and on dead bark, throughout the year.


Very common.

Orbilia leucostigma (Fries) Fries, op. cit.:357 (1849), differs only in


being whitish when moist b u t is much less a b u n d a n t .

Orbilia luteorubella (Nylander) Karsten in Not. Sallsk. F a u n a Flora


Fennica 11:248 (1870). (PL X X I B . )
Apothecia subsessile, smooth, disc up to 1.5 m m . across, flat or
convex when moist, pallid, drying yellowish. Asci cylindric-clavate, up
to 50x4.5/«, 8-spored; ascospores biseriate, narrowly fusiform, 5 - l l x
1-1.5/«, non-septate; paraphyses slender, a b r u p t l y swollen to 2.5/« at
the tip.

167
On rotting wood and decaying bracket fungi in summer and autumn.
Fairly common.

Orbilia auricolor (Bloxam ex Berkeley) Saccardo, Sylloge F u n g o r u m 8:


625 (1889). (PL XXIF.)

Apothecia resembling those of O. luteorubella b u t golden-yellow


when d r y and surrounded b y short white anchoring hyphae like a
r u d i m e n t a r y b u t quite conspicuous subiculum. Asci up to 30x4,«,
8-spored, ascospores biseriate, rod-shaped, 4 - 6 x 0 . 5 - 1 / « ; paraphyses
enlarged at the tip b u t not as m a r k e d l y knob-headed as in most species.

On bark of deciduous trees in spring and autumn. Fairly common.

Orbilia inflatula (Karsten) Karsten in Not. Sallsk. F a u n a Flora


Fennica 11:248 (1870), is a synonym.

Orbilia curvatispora Boudier in Bull. Soc. mycol. France 4 : 8 0 (1888).


(PL X X I c . )

Apothecia sessile, white and translucent when fresh, drying yel-


lowish-buff, disc convex when moist, up to 1 m m . across. Asci cylindric-
clavate, up to 4 5 x 4 f i , 8-spored; ascospores biseriate, cylindrical and
distinctly curved, 10-11 x 0 . 5 / t ; paraphyses slender, a b r u p t l y swollen
at the tip.

On bark and rotten wood in summer and autumn. Not uncommon.

Orbilia sarraziniana Boudier in Revue Mycologique 7:221 (1885).


(PL X X I E . )

Apothecia sessile, very pale pink and translucent, often appearing


greyish at the centre over the point of a t t a c h m e n t , where the dark
bark shows through the t h i n flesh, disc convex when moist, up to
1 m m . across b u t usually smaller. Asci up to 40x4/«, 8-spored; asco-
spores biseriate, very narrowly clavate, pointed below, 6 - 8 x 0 . 5 / « ;
paraphyses slender, a b r u p t l y swollen to 3 /t at the tip.

On damp wood or bark of deciduous trees, May to September. Uncommon.

Orbilia vinosa (Albertini & Schweinitz ex Fries) Karsten, Mycologia


Fennica 1:101 (1871), is an uncommon species distinguished b y its
long slender ascospores, 1 1 - 1 6 x 1 - 1 . 5 p.

168
HYALINIA Boudier

Hyalinia Boudier in Bull. Soc. mycol. France 1:114 (1885).


Apothecia like those of Orbilia b u t with the marginal cells running
out into thin-walled cylindrical hairs which cohere in groups to give a
d e n t a t e appearance to the margin as viewed under a hand-lens, para-
physes not knob-headed.

Hyalinia rubella (Fries) Nannfeldt in Nova Acta Reg. Soc. Sci. Upsal.
IV, 8:252 (1932). (PI. XXID.)
Apothecia subsessile, light red or flesh-coloured when moist, disc
concave, up to 2 m m . across, margin flexuous, d e n t a t e with the co-
hering hairs, inrolled, purplish and minutely downy when dry. Asci
cylindric-clavate, up to 4 0 x 4 / / , 8-spored; ascospores biseriate,
narrowly clavate, pointed below and often slightly curved, 8 - 1 2 x 0 . 5 /<;
paraphyses slender, not swollen at t h e tip.

On bark, especially of Salix and Populus, throughout the year. Not common.

According to Nannfeldt Orbilia coccinella (Sommerfelt) Fries,


S u m m a Veg. Scand. Sect. Post.:357 (1849), is probably a synonym
t h o u g h t h e name has been diversely employed for various genuine
species of Orbilia.

Family 8. Dermateaceae

Here are collected a large n u m b e r of genera which have in common


a usually rather soft flesh composed mainly of thin-walled more or less
globose cells. Numerous tribes have been proposed within the family
b u t t h e y are not all clearly distinguished and there are m a n y genera
which cannot be accommodated in any of the existing tribes. Instead
of a t t e m p t i n g a " n a t u r a l " classification, therefore, the genera are
keyed out below in an artificial m a n n e r which will work fairly well, at
least with the representative species described here. Many of the
genera in current use, however, are ill-defined and will probably be
united in the light of more thorough studies of a greater n u m b e r of
species.

I. Apothecia found on wood or bark of trees, shrubs or canes of Rubi:


A. Asci filled with innumerable rod-shaped secondary spores
See Tympanis (p. 112)
B. Asci containing normal ascospores, eight or less:

169
1. Paraphyses tips green, disc greenish-black, on Rubus
Ploettnera (p. 194)
2. Paraphyses tips fusiform, margin toothed, cup pinkish
Habrostictis (p. 176)
3. Paraphyses lacking these distinctive_characters:
*Ascospores becoming smoky-brown, slightly swollen at
each end Catinella (p. 176)
**Ascospores hyaline:
+ Ascospores broadly elliptical to reniform, oflen
multiseptate:
/Apothecia permanently immersed in wood,
pallid:
t Ascospores septate
Cryptodiscus (p. 193)
jjAscospores nonseptate
Propolis (p. 194)
//Apothecia erumpent:
fApothecia concave with a lorn white
margin, not emergent from the bark,
ascospores nonseptate
Ocellaria (p. 175)
t Apothecia more or less stipitate, emer-
gent, often in clusters and accom-
panied by a conidial state:
§Apothocia dark brown to black,
conidia narrowly fusiform:
Ascospores ellipsoid
Dermea (p. 172)
Ascospores fusoid to acicular,
multiseptate:
AAscospores acicular, up
to 5-septate, conidial
state a Chondropo-
dium with filiform co-
nidia in locules in a
stroma Durandiella
A AAscospores fusoid, mul -
tiseptate, conidial
state different or ab-
sent Praginopora
§Apothecia light coloured, disc im-
marginate, often pruinose, coni-
dia oblong-ellipsoid
Pezicula (p. 173)
///Apothecia superficial, lenticular, sessile, as-
cospores filled with numerous small oil
drops, becoming 1-septate
Graddonia (p. 178)
-f+Ascospores clavate, multiseptate, paraphyses
united at the tip in a brown epithecium
Patellariopsis (p. 177)

170
+ + +Ascospores narrowly fusiform, paraphysis tips
not united in an epithecium:
/ Ascospores very long, multiseptate, apo-
thecia with a subiculum
Trichobelonium (p. 180)
//Ascospores 0-1-septate:
tApothecia seated on a subiculum
Tapesia (p. 182)
t j A p o t h e c i a superficial or erumpent but
without a subiculum:
§Apothecia pinkish or pallid through-
out . . . Orbiliopsis (p. 192)
§Apothecia externally dark brown
to blackish-olive:
Apothecia massive, externally
hairy, ascus pore not blued
by iodine Haglundia (p. 180)
Apothecia thinner, smooth or
merely downy near the mar-
gin, ascus pore usually blued
by iodine Mollisia (p. 182)
(On Rubus canes see also Pyrenopeziza)

II. On leaves and herbaceous stems:


A. Apothecia bearing dark septate hairs . . . . Pirottaea (p. 181)
B. Apothecia smooth, or minutely downy at the margin with thin-
walled obtuse elongated cells only:
1. Apothecia superficial:
*Apothecia light coloured:
+Ascospores 1-septate . . . . Calloria (p. 191)
+ + A s c o s p o r e s non-septate with a hyaline sheath
Coleosperma (p. 178)
**Apothecia usually dark coloured externally or asco-
spores nonseptate:
+Ascospores 0-1-septate . . . Mollisia (p. 182)
+ +Ascospores multiseptate Belonopsis (p. 179)
2. Apothecia erumpent:
* Apothecia erumpent in clusters from a stroma, all black
Ephelina (p. 179)
**Apothecia erumpent in rows from a hyphal strand in
leaf tissue Spilopodia (p. 191)
***Apothecia borne neither on a black stroma nor on an
immersed hyphal strand:
/ D a r k mollisioid apothecia:
fApothecia often becoming naked, not on
grasses, sedges etc.
Pyrenopeziza (p. 186)
t j A p o t h e c i a remaining immersed, or with
lanceolate paraphyses, always on
grasses, sedges or rushes
Hysteropezizella (p. 188)

171
//Apothecia not mollisioid:
fApothecia raised above the matrix at
maturity:
§Apothecia minute, turbinate, on
dead leaves
Drepanopeziza (p. 189)
§§Apothecia larger, lenticular, often
on living leaves:
Ascospores non-septate
Pseudopeziza (p. 189)
Ascospores 1-septate
Fabraea (p. 190)
tjApothecia not raised above the sub-
strate, disc often exposed by the rais-
ing of marginal lobes or dehiscence of
a lid:
§Disc circular:
Disc covered by three or four
dark flaps of fungus tissue
Naevia (p. 196)
Disc covered by irregular torn
tissue or by a detachable lid:
ADisc olive coloured
Trochila (p. 195)
A ADisc pallid to pinkish
Laetinaevia (p. 196)
§§Disc elongated Duebenia (p. 197)

III. On soil and humus, margin strongly toothed Podophacidium (p. 193)
For other genera with an immersed erumpent hymenium see Phacidiales,
p. 198.
For monographs of Durandiella Seaver and Pragmopora Massalongo, not
yet known in Britain, see papers by Groves in Canadian Journ. Botany
32:116-144 (1954) and 45:169-181 (1967) respectively.

DERMEA Fries

Dermea Fries, Systema Orbis Vegetabilis 1:114 (1825).

Apothecia erumpent, usually in clusters, cup-shaped, sessile or


shortly stalked, leathery, dark brown to black, asci 8-spored or less,
ascospores ellipsoid, hyaline or yellowish, sometimes septate, para-
physes slender, longer t h a n the asci and cohering above them. Conidial
states very diverse in gross morphology b u t mostly containing elon-
gate-fusiform, often curved, conidia, pointed at the ends. There are
about five British species. For a comprehensive study of the genus see
a paper b y J. W . Groves in Mycologia 38:351-431 (1946).

172
Dermea prunastri (Persoon) Fries, S u m m a Veg. Scand. Sect. Post.: 362
(1849). (PI. X X I I E . )
Apothecia erumpent in small groups from cracks in the bark, cup-
shaped with a short ill-defined stalk, smooth, leathery, dark brown to
black throughout, disc flat, about 1 m m . across. Asci cylindric-clavate,
up to 120x15/«, 8-spored, the pore sometimes feebly blued b y iodine;
ascospores irregularly biseriate, narrowly elliptical, straight or slightly
curved, 15-20x5-7.5/«, usually non-septate; paraphyses slender,
usually branched near the apex, tips closely septate, enlarged to 3 /«
wide and cohering.
The conidial state, often associated with the apothecia, consists of a
cylindrical or conical, smooth, black pycnidium, up to 2 m m . tall,
containing narrowly fusiform, more or less sickle-shaped, hyaline,
nonseptate conidia, 2 0 - 3 0 x 4 - 7 / « .
On dead twigs of Prunus spinosa and P. domestica.

D. padi (Fries) Fries, op. cit.:362 (1849), occurs on the same hosts
and on P. padus and is distinguished mainly b y its narrower conidia,
20-30x4/«.
Dermea cerasi (Persoon ex Merat) Fries, op. cit.:362 (1849), found on
Prunus avium and P. cerasus, differs in its m u c h longer conidia,
40-60x2.5-4.5/«.
Dermea ariae (Persoon) Tulasne apud Karsten, Mycologia Fennica
1:224 (1871), occurs on Sorbus and D. tulasnei Groves on Fraxinus.

PEZICULA Tulasne

Pezicula Tulasne, Selecta F u n g o r u m Garpologia 3:182 (1865).


Apothecia erumpent, usually in clusters from a common stroma,
cup-shaped with a convex, usually pruinose disc, without a raised
margin, brightly coloured, exceptionally darkening when dried, tissue
pseudo-paronchymatous, asci 4- or 8-spored, large, thick-walled with
a broad pore stained deep blue b y iodine, ascospores elliptical or
slightly kidney-shaped, large, hyaline or yellowish, often becoming 3-
or more septate, sometimes budding in the ascus, paraphyses usually
swollen at the tip and more or less coherent. The conidial state usually
belongs to the imperfect genus Cryptosporiopsis, with large, elliptical,
hyaline conidia. There are about a dozen British species. For a useful
account of Pezicula see G. Johansen in Dansk Botanisk Arkiv 13 (3/4):
1949.

173
Pezicula livida (Berkeley & Broome) Rehm in 26th Berichte des
Naturhistorischen Vereins in Augsburg: 112 (1881). ( P I . X X I I F . )
Apothecia erumpent, singly or in small groups, disc light orange-
yellow or brownish, flat or convex, up to 2 m m . across, exterior paler,
smooth, subsessile. Asci clavate, up to 120x20/«, 4-8-spored; asco-
spores uniseriate or biseriate, according to the n u m b e r present, elliptic-
fusiform, 2 2 - 2 4 x 6 - 7 /«, hyaline, often 3- or more septate at m a t u r i t y ;
paraphyses slender, 1.5-2/« thick, swollen at the apex. The ascospores
sometimes bud off cylindrical microconidia.

Saprophytic on branches, twigs, and cones of Abies, Araucaria, Cupressus,


Juniperus, Larix, Picea, Pinus and Pseudotsuga, October to June. Common.

Pezicula rubi (Libert) Niessl apud Rabenhorst, Fungi eruopaei: 2122


(1876). (PI. X X I I n . )
Apothecia erumpent, scattered, usually singly, sessile, lenticular,
very brittle when dry, disc convex, finely pruinose, dull ochraceous
yellow or reddish-orange, up to 1 m m . across. Asci cylindric-clavate,
up to 115x20/«, 8-spored; ascospores biseriate, elliptic-fusiform,
somewhat pointed at each end, 18-26x6-8/«, sometimes becoming
3-septate, hyaline; paraphyses slender, swollen to 5/t at the tips.
On dead stems of Rubus "fruticosus", less often on Rosa, July to Decem-
ber. Common.

Pezicula scoparia (Cooke) Dennis, E d . 1 : 1 0 9


Helotium scoparium Cooke in Grevillea 4: 111 (March 1876).(Pl.XXIIi.)

Apothecia erumpent, solitary or in pairs, sessile, disc convex, pale


dingy buff, up to 0.8 m m . across, minutely pruinose. Asci cylindric-
clavate, up to 90x 20 /«, 8-spored; ascospores biseriate, elliptic-fusiform,
rather pointed at each end, 1 8 - 2 5 x 5 /«, hyaline, occasionally 1-septate;
paraphyses slender, enlarged to 5 /« at the tip.
On dead twigs of Sarothamnus scoparius. Rare. Is has been suggested that
this is only P. rubi, which it greatly resembles, on an unusual host. It is
figured here to draw attention to the species in the hope it may be re-
collected and studied.

Pezicula corticola (C. A. Jorgenssen) Nannfeldt in Nova Acta Reg. Soc.


Sci. Upsal. IV, 8 : 9 4 (1932). (PI. X X I I G . )
Apothecia erumpent in small clusters, sessile, disc convex, pruinose,
bright yellow, up to 0.7 m m . across. Asci cylindric-clavate, up to 110

174
x l 8 / « , 8-spored; ascospores irregularly biseriate, elliptical, somewhat
pointed at the ends 17-29x7-9/.« hyaline, becoming 1 - 3 - s e p t a t e ; para-
physes slender, branched, swollen at the tips.
On bark of Pyrus malus and P. communis, sometimes associated with
cankers, in August. Apothecia are rarely seen but the conidial state, Crypto-
sporiopsis corlicola (Edgert.) Nannf. op. cit. is not uncommon in spring and
autumn.

Pezicula malicorticis (Jackson) Nannfeldt op. cit.:91 (1932), also


occurs on Pyrus malus b u t its apothecia have yet to be found in the
British Isles. The common apple fruit rot fungi Gloeosporium album
Osterw. and G. perennans Zeller & Childs also have species of Pezicula
as their ascus states.
Pezicula carpinea (Persoon) Tulasne apud Fuckel in J a h r b . Nass.
Vereins f. N a t u r k u n d e 23-24:279 (1870), occurs on branches of Car-
pinus and Fagus.
Pezicula cinnamomea (de Candolle ex Persoon) Saccardo in Michelia
2:232 (March 1881), is fairly common on b a r k of Quercus and Dermatea
pseudoplatani Phillips in Grevillea 17:45 (1888), on Acer m a y be a
synonym; Groves t h o u g h t it possibly identical with P. carnea (Cooke &
Ellis) Rehm in Ber. Bayer. Bot. Ges. 13:199 (1912).
Pezicula jrangulae (Fries) Fuckel in J a h r b . Nass. Vereins f. Natur-
kunde 23-24:279 (1870), on Rhamnus frangula is peculiar in its apo-
thecia drying almost black so as to resemble those of a Dermea-, its asci
are 4-spored.
Pezicula houghtonii (Phillips) Groves in Mycologia 38:417 (1946),
occurs on Pruhus lusitanica.
Pezicula myrtillina Karsten, Mycologia Fennica 1 : 1 6 4 (1871), is
found rarely on dead twigs of Vaccinium myrtillus.

OCELLARIA (Tulasne) Karsten

Ocellaria (Tulasne) Karsten, Mycologia Fennica 1:21 (1871).


Apothecia originating below bark, exposed by r u p t u r i n g it b u t not
erumpent, asci and ascospores as in Pezicula and conidial state a
Cryptosporiopsis, as in t h a t genus.

Ocellaria ocellata (Persoon) Schroeter, K r y p t o g a m e n Flora von Schle-


sien 3 (2): 150 (1893). (PI. X X I I J . )
Apothecia sessile, scattered, bursting through the b a r k but not fully
erumpent, saucer-shaped, attached to the substrate by a scanty subi-

175
culum of almost hyaline hyphae 5 - 6 /« wide, disc flat, up to 2 m m .
across, golden-brown, with a prominent margin. Asci clavate, thick-
walled, up to 130x25//, the broad pore deep blue with iodine; asco-
spores uniseriate or biseriate according to their number, four or eight
broadly elliptical, 2 0 - 3 0 ( - 4 0 ) x 8 - 1 2 ( - 1 4 ) /<, hyaline or yellowish; para-
physes slender, branched, 3 /« wide at the tip.

On Salix branches, in spring. Not uncommon. Also on Hippophae, Populus


and Rhamnus in Europe.

HABROSTICTIS Fuckel

Habrostictis Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:249


(1870).
Apothecia sessile, erumpent, brightly coloured, with a laciniate,
fimbriate margin, asci slender, thin-walled, 8-spored, pore not blued by
iodine, ascospores hyaline, non-septate, paraphyses slender with
swollen lanceolate tips.

Habrostictis rubra Fuckel, op. cit.:249 (1870). (PI. XXIIL.)

Apothecia developed beneath the b a r k and exposed b y r u p t u r i n g it,


sessile, scattered or gregarious, disc pink to orange, concave, up to
2 m m . across, margin conspicuously toothed and torn, curving inwards
to conceal the disc when dry. Asci narrowly cylindrical with slender
stalk and t r u n c a t e tip, up to 90x5,«, 8-spored; ascospores biseriate,
narrowly fusiform, 12-15x2/«, non-septate, paraphyses slender, about
1 /t thick, a b r u p t l y enlarged at the tip into a thin-walled lanceolate
swelling about 12x4/«.

On dead branches of Ulex, Ulmus, etc. in spring and autumn. Apparently


rare but the apothecia nestling in cracks of the bark, though obvious enough
when wet, are very hard to see when dry and may easily be overlooked.
The fungus is placed in Dermateaceae to accord with Nannfeldt's system
but the shape of the asci, their negative iodine reaction and the swollen tips
of the paraphyses all recall the Orbiliaceae.

CATINELLA Boudier

Catinella Boudier, Hist. Class. Discom. d ' E u r o p e : 1 5 0 (1907).


Apothecia sessile, soft-fleshed, outer tissue composed of dark thin-
walled cells, staining deep violet with K O H solution, asci 8-spored, the

176
pore not blued b y iodine, ascospores non-septate, slowly becoming
brown, paraphyses gummed together at the tips by an amorphous sub-
stance. The genus is very distinct and its affinities are not clear. The
peculiar colour reaction with Potassium hydroxide and the large-celled
dark outer tissue have suggested to most authors a position near
Karschia but the absence of a blue reaction in the ascus with iodine led
Nannfeldt to reject this and indicate affinity with t h e Dermateaceae.
There is one European species:

Catinella olivacea (Batsch ex Persoon) Boudier, op. cit.: 150 (1907).


(PI. X X I V M . )

Apothecia sessile, saucer-shaped with a broad base and dark-brown


anchoring hyphae, disc up to 10 m m . across, olive-green to blackish-
olive, flat at m a t u r i t y , exterior dark brown, often with a ochraceous
margin, often pruinose and obscurely vertically furrowed. Asci
cylindric-clavate, up to 90x6/«, 8-spored; ascospores uniseriate, ellip-
tical b u t usually slightly constricted in the middle in a very distinctive
manner, 7 - 1 1 x 4 - 5 /«, at first almost colourless, slowly becoming olive-
brown, with a small oil drop in each end b u t never septate; paraphyses
cylindrical, colourless, septate, their tips covered b y amorphous brown
matter.

On rotten wood, especially of very wet logs, throughout the year. Quite
common.

PATELL A R I 0 P S IS Dennis

Patellariopsis Dennis in Kew Bulletin 19:114 (1964).


Apothecia sessile, black, cartilaginous, very like those of Patellaria
b u t with unitunicate asci in which the pore stains deep blue with
iodine.

Patellariopsis clavispora (Berkeley & Broome) Dennis in Kew Bulletin


19:114 (1964). (Fig. 29L.)
Apothecia scattered, superficial, discoid, up to 2 m m . diameter, disc
convex when moist, black, receptacle smooth, black with white flesh
composed of slender radiating h y p h a e which pass into an outer tissue
of subglobose cells whose walls are heavily encrusted with dark brown
pigment. Asci cylindric-clavate, 125x10//, 8-spored, spores biseriate,
clavate, hyaline, 27-38x4.5/«, 3-5-septate, paraphyses cylindrical,
apical cells clavate, dark brown, united in an epithecium.

177
On wood and bark, Acer, Crataegus, Fagus, Ligustrum, Quercus. Oc-
tober-November. Rare.

GRADDONIA Dennis

Graddonia Dennis in Kew Bulletin:359 (1955).


Apothecia sessile, superficial, lenticular, dark coloured, outer tissue
composed of large, dark, thin-walled cells, asci 8-spored, pore not blued
b y iodine, ascospores large, hyaline, becoming 1-septate, paraphyses
slender. There is a single species:

Graddonia coracina (Bresadola) Dennis, op. cit.:359 (PI. X X I Y c . )


Apothecia gregarious, sesslie, saucer-shaped, smooth, soft-fleshed,
dark brown, disc reddish-brown, blackening with age or on drying,
flat, without a raised margin, up to about 1.5 m m . across. Asci cylin-
dric-clavate, thinwalled, up to 170x13,«, 8-spored; ascospores uni-
seriate, elliptic-fusiform, 1 8 - 2 4 x 7 - 9 / / , hyaline, at first non-septate
and packed with small oil drops, tending to become 1-septate; para-
physes slender, slightly enlarged to 3 // at the tip.

On fallen twigs in damp places, February to April. Rare.

COLEOSPERMA Ingold

Coleosperma Ingold in Trans. Brit, mycol. Soc. 3 7 : 9 (1954).


Apothecia aquatic, sessile, superficial, soft-fleshed, light coloured,
asci 8-spored, ascospores non-septate, hyaline, enclosed in a gelatinous
sheath. There is a single species:

Coleosperma lacustre Ingold, op. c i t . : 9 (1954). (Fig. 18B.)


Apothecia superficial, sessile on a broad base, smooth, disc slightly
convex up to about 0.3 m m . across, white, t r a n s l u c e n t ; flesh very t h i n
and soft, composed of small subglobose cells. Asci clavate, up to 140x
18//, 8-spored, t h e pore not blued b y iodine; ascospores usually ar-
ranged in two rows, somewhat overlapping, fusiform, thin-walled,
hyaline, 2 3 - 3 3 x 6 . 5 - 9 //, packed with small oil drops, each surrounded
b y a colourless sheath; paraphyses slender, slightly swollen at the tip,
septate, sometimes branched.
On submerged portions of dead stalks of Scirpus lacustris, May to Sep-
tember. Apparently common.

178
EPHELINA Saccardo
Ephelina Saccardo, Sylloge F u n g o r u m 8:585 (1889).
Apothecia erumpent in clusters from a black subepidermal stroma
on stems, cup-shaped, smooth, black, formed of r a t h e r small more or
less rounded cells, asci 8-spored, pore not blued by iodine, ascospores
hyaline, paraphyses slender.

Ephelina lugubris (de Notaris) von Hohnel in S.B. Ak. Wiss. Wien.
Math.-Nat. Klasse Abt. 1 , 1 2 7 : 5 7 5 (1918). (Fig. 6F.)
Apothecia black throughout, with a pruinose or finely scurfy outer
surface, disc concave, grey, about 1 m m . across. Asci cylindric-clavate,
up to 70 x 7 /t, 8-spored; ascospores biseriate, somewhat clavate, 9-10 x
3 - 4 / i , hyaline, normally non-septate; paraphyses cylindrical, colour-
less.
At soil level on stems of Rhinanthus. Very common.

BELONOPSIS (Saccardo) Rehm

Belonopsis (Saccardo) Rehm, Rabenhorst's Kryptogamen Flora


Deutschland 1 (3): 5 7 1 ( 1 8 9 1 ) .
Apothecia more or less cup-shaped, sessile, superficial, soft-fleshed,
outer tissue formed of brown, thin-walled, globose cells, asci 8-spored,
the pore blued b y iodine, ascospores elongated, 3- or more septate,
hyaline.

Belonopsis excelsior (Karsten) Rehm, op. cit.: 5 7 2 (1891). (PI. X X I I I C . )

Apothecia scattered or in small groups, superficial, saucer-shaped,


sessile on a small base, dark brown beneath, smooth, disc slightly con-
vex, up to 2 m m . across, light grey. Asci cylindric-clavate, up to 130x
10/i, 8-spored; ascospores lying parallel, cylindric-clavate, pointed
below, 50-80x3-4/«, often very slightly curved, septate; paraphyses
slender, enlarged up to 7 p wide at the tip.

In swarms below water level on standing culms of Phragmites, spring to


autumn. Common.

Belonopsis filispora (Cooke) Nannfeldt in Trans. Brit, mycol. Soc. 20:


192 (1936). (PI. XXIIIG.)

Apothecia superficial, saucer-shaped, sessile, attached b y numerous


brown anchoring hyphae, dark brown with a paler margin, smooth,

179
disc slightly concave, light grey, less t h a n 1 m m . across. Asci cylindric-
clavate, up to 85x12/«, 8-spored; ascospores irregularly arranged,
cylindric-clavate, pointed below, 4 0 - 4 5 x 3 - 3 . 5 / / , 3-septate, hyaline^
sometimes very slightly curved.

On culms of Brachypodium, in summer. Apparently uncommon.

Belonopsis graminea (Karsten) Saccardo & Sydow, Sylloge Fungo-


r u m 16:753 (1902), with 3-septate ascospores 20-30x3-4/«, occurs on
culms of small grasses such as Agropyron repens.

Belonopsis iridis (Crouan) Graddon in Trans. Brit, mycol. Soc. 44:


612 (1961) with 3-5-septate ascospores. 44-65x2.5-3/« occurs on
J uncus, Iris and Typha.

TRICHOBELONIUM (Saccardo) Rehm


Trichobelonium (Saccardo) Rehm in Rabenhorst's Kryptogamen
Flora Deutschland 1 (3): 590 (1891).
Apothecia gregarious on a dense, dark-coloured, hyphal web (subi-
culum), more or less cup-shaped, dark coloured, flesh composed on the
outside of brown rounded cells, asci 8-spored, t h e pore blued b y io-
dine; ascospores elongated and 3- or more septate.

Trichobelonium obscurum (Rehm) Rehm, op. cit.: 590 (1891) (PI


XXIVn.)
Apothecia seated on a dense dark brown or almost black m a t of
hyphae 4 - 6 m m . wide, saucer-shaped, externally dark brown with a
pale margin, disc somewhat concave, light grey, up to 2 mm. across.
Asci cylindric-clavate, up to 90x9/«, 8-spored; ascospores irregularly
arranged, cylindric-fusiform, often curved 30-40x2.5-3/«, 3-7-septate,
hyaline; paraphyses 2.5/« thick.

On old stems of Calluna, April to August. Common.

Trichobelonium asteroma (Fuckel) Rehm, op. cit.:592 (1891), occurs


on the larger Carices in spring.

HAGLUNDIA Nannfeldt
Haglundia Nannfeldt in Nova Acta Reg. Soc. Sci. Upsal IV 8-107
(1932).
This genus has been separated from Mollisia mainly on account of
its long, cylindrical septate hairs. The type species is:

180
Haglundia perelegans Nannfeldt, op. cit.: 1 0 8 (1932). (PI. XXIIIA..)

Apothecia crowded or scattered, almost superficial b u t with a short


peg-like base inserted in the surface layers of wood, cup-shaped,
blackish-brown with a grey-brown or whitish margin, clothed with
cylindrical, flexuous, septate, obtuse, grey-brown hairs up to 120/«
long, disc watery-grey to yellowish, up to 3 m m . across. Asci cylindric-
clavate, up to 70x4/«, the small pore not blued b y iodine, 8-spored;
ascospores hyaline, continuous, narrowly fusiform, 7-8x1.5/«; para-
physes cylindrical, 2 - 3 /« wide, longer t h a n the asci.

On rotting stumps and logs of deciduous trees, August to October. Un-


common.

Haglundia elegantior Graddon in Trans. Brit, mycol. Soc. 5 0 : 9 (1967)


differs in its whiter margin and large ascospores 11-13.5x3-4/«.

PIROTTAEA Saccardo

Pirottaca Saccardo in Michelia ,1:424 (November 1878).


Apothecia erumpent or becoming superficial, sessile, cup-shaped,
dark coloured, clothed with stiff, dark, usually pointed hairs, especially
round the margin, asci 8-spored, ascospores 0-3-septate, hyaline. The
species are found on dead herbaceous stems b u t are all apparently un-
common.

Pirottaea brevipila (Roberge) Boudier, Hist. Class. Discom. d ' E u r o p e :


135 (1907). (PI. XXIVG.)
Apothecia erumpent, cup-shaped, dark brown, ornamented with
dark brown, pointed, septate hairs up to 80x7/«, disc concave, light
grey, up to 0.5 m m . across. Asci cylindric-clavate, up to 80x8/«, the
small pore not clearly blued b y iodine, 8-spored; ascospores narrowly
fusiform, tapering below, 24-28x2.5-3/«, becoming 1-septate, said t o
be sometimes 3-septate; paraphyses cylindrical, obtuse, 3 p. thick at
the tip.

On dead stems of Centaurea in summer. Apparently uncommon.

Pirottaea senecionis (Cooke & Phillips) Nannfeldt in Nova Acta Reg.


Soc. Sci. Upsal. IV, 8:134 (1932), occurs on stems of Senecio.
Pirottaea nigro-striata Graddon in Trans. Brit, mycol. Soc. 5 9 : 1 2
(1967) occurs on dead stems of Heracleum in summer.

181
TAPESIA Fuckel

Tapesia Fuckel in Jahrb". Nass. Vereins f. N a t u r k u n d e 23-24:300


(1870).
Tapesia as currently interpreted included those species of Mollisia
in which t h e apothecia are seated on a common subiculum or web of
superficial hyphae. This is not a very reliable character, especially as
m a n y Mollisias have well developed brown anchoring hyphae beneath
the apothecia. U n f o r t u n a t e l y Tapesia is the older name and if the two
genera are united it will t a k e precedence over the more widely used
Mollisia.

Tapesia strobilicola (Rehm) Saccardo, Sylloge F u n g o r u m 8:377 (1889).


(PI. X X I V D . )
Apothecia sessile, gregarious on a dark brown hyphal m a t , cup-
shaped, dark brown, disc concave, pallid to yellowish, up to 2 m m .
across. Asci cylindric-clavate, up to 45 x 6 n, pore blued b y iodine; asco-
spores biseriate, elliptic-fusiform, non-septate, 6 - 9 x 2 . 5 - 3 /i, sometimes
slightly curved; paraphyses cylindrical.

On fallen cones of Larix and Pinus, in spring. Common.

Tapesia fusca (Persoon ex Merat) Fuckel, op. cit.: 302 (1870), occurs
commonly on dead wood and bark, T. rosae (Persoon ex Merat) Fuckel
on dead stems of Rosa and T. melaleucoides Rehm on dead stems of
Calluna.
Tapesia retineola (Rabenhorst) Karsten, Mycologia Fennica 1:209
(1871), with rather bright coloured apothecia on a dense blackish
h y p h a l m a t covering Phragmites culms, just above water level, is
sometimes referred to Trichobelonium b u t its ascospores are normally
non-septate.

MOLLISIA (Fries) Karsten

Mollisia (Fries) Karsten, Mycologia Fennica 1 : 1 5 (1871).


Apothecia superficial, saucer-shaped, often attached b y quite a
small central base, sometimes surrounded b y short dark anchoring
hyphae b u t not grouped on a common hyphal web, typically d a r k
brown to blackish below b u t commonly with a pale margin, nominally
smooth b u t often w i t h short downy elongated cells like small unicellu-
lar clavate hairs towards the margin, asci 8-spored, ascospores 0-1-sep-

182
t a t e , usually narrowly fusiform, hyaline, ascus pore usually blued by
iodine, paraphyses cylindrical, obtuse, often with rather oily contents.
The t e x t u r e is normally soft and the outer tissue composed of rounded
cells with thin dark brown or olive-brown walls. The n u m b e r of good
species, though certainly large, is quite unknown. In the past the genus
has been somewhat broadly interpreted and t h e species described
below have been selected to illustrate its scope in current literature.
For detailed illustrated descriptions of m a n y critical european spe-
cies see a series of papers by Le Gal and Mangenot in Revue de Myco-
logie 23:28-86 (1958), 25:135-214 (1960), 2 6 : 2 63331 (1961) and 31:
3 - 4 4 (1966).

Mollisia cinerea (Batsch ex Merat) Karsten, op. cit.: 189 (1871). (PI.
XXIIIiT~
Apothecia gregarious, superficial, saucer-shaped, becoming flattened,
grey-brown beneath, with short anchoring hyphae, margin whitish,
outline often irregular or undulating, disc light grey, sometimes drying
yellowish, up to about 2 m m . across, if larger t h e n usually irregular.
Asci cylindric-clavate, up to 7 0 x 6 / / , the pore blued b y iodine, 8-spor-
ed; ascospores biseriate, narrowly elliptic-oblong, straight or only very
slightly curved, 7 - 1 0 x 2 - 2 . 5 / / , non-septate; paraphyses cylindrical,
obtuse.
On dead wood and branches, throughout the year. Very common.

Mollisia melaleuca (Fries) Saccardo, Sylloge F u n g o r u m 8:337 (1889).


(PI. X X I I I J . )
Apothecia gregarious, sessile, saucer-shaped, superficial, blackish-
brown and smooth below, sometimes finely wrinkled, margin minutely
downy, disc whitish, up to 2 m m . across. Asci cylindric-clavate, up to
60x6/«, the pore blued by iodine, 8-spored; ascospores biseriate, nar-
rowly elliptical, often slightly curved, 8 - 1 4 x 1.5-2.5 //, non-septate b u t
often containing two small oil drops; paraphyses cylindrical, 2.5//
thick.
On rotting decorticated wood, less common than M. cinerea from which
it is distinguished largeiy by colouring.

Mollisia ventosa (Karsten) Karsten, op. cit.: 188 (1871). (PI. XXIVB.)

Apothecia gregarious, often in small clusters, superficial, saucer-


shaped, often irregularly lobed, smooth, blackish below with a pale

183
margin, disc slightly concave, rather yellowish-grey, up to 2 m m .
across, flesh rather thick, soft, of radiating rows of thin-walled cells
t h a t become rounded towards the outer surface of the cup. Asci
cylindric-clavate, with long stalks, up to 120x7/«, 8-spored, the pore
blued b y iodine; ascospores largely uniseriate, fusiform, straight or
slightly curved, 10-20x2-3.5/«, 1-septate, said to become 3-septate;
paraphyses slender, a b o u t 1 /< thick.
On rotting decorticated wood, in summer. Not common. This is evidently
a species transitional to Belonopsis.

Mollisialigni (Desmazieres) Karsten, op. cit.:204 (1871). (PI. XXIIIK.)

Apothecia scattered, superficial, cup-shaped, dark blackish-brown


and minutely downy all over with dark brown protruding clavate cells,
extreme margin paler, disc concave, dark grey, seldom m u c h over
1 m m . across. Asci cylindric-clavate, up to 70x6/«,8-spored; ascospores
biseriate, elliptic-cylindric, 6 - 1 2 x 2 - 2 . 5 / * , non-septate; paraphyses
cylindrical, 2.5 n thick.
On decorticated wood and chips of deciduous trees and on fallen beech
mast, throughout the year but especially in spring and autumn. Very
common.

Mollisia caespiticia (Karsten) Karsten, op. cit.: 188 (1871). (PI.


XXIVA.)

Apothecia erumpent from b a r k in small clusters, often seated over


some member of the Sphaeriales, cup-shaped, grey, darker towards the
base, smooth, disc slightly concave, dark grey, up to 1.5 m m . across,
often deformed b y m u t u a l pressure. Asci cylindric-clavate, up to
40x4.5/«, 8-spored, the pore not blued by iodine; ascospores biseriate,
narrowly clavate, 4 - 7 x 1 / « , non-septate; paraphyses few, slender,
cylindrical.
On dead branches as of Betula, Ribes, Salix and Sambucus, September to
October. Not common. A collection has also been seen on Crataegus, in
spring, in which apothecia with normal ascospores occur together with
others in which the asci are fdled with innumerable small rod-shaped spores
like those of a Tympanis.

Mollisia discolor (Montagne) Phillips, Manual Brit. Discom.: 175 (1887),


var. longispora le Gal in Revue de Mycologie N.S. 4:57 (1939). (PI.
XXIIIM.)

Apothecia erumpent in clusters from cracks in bark, cup-shaped,


becoming flattened and often undulating, lobed or deformed b y m u t u a l

184
pressure, externally dark brown, disc light grey drying yellow, about
1.5 m m . across. Asci cylindric-clavate, up to 7 0 x 8 / / , 8-spored; asco-
spores biseriate, narrowly elliptical, 1 0 - 1 3 x 2 - 2 . 5 //, with two oil drops,
non-septate; paraphyses cylindrical, a b o u t 2/« thick.

On dead twigs, especially of Quercus, in spring. Common.

The typical variety of M. discolor occurs on twigs of Cornus, with


ascospores 7-9.5x2-3/«.

Mollisia ramealis (Karsten) Karsten, op. cit.: 187 (1871). (PI. XXIIID.)

Apothecia gregarious, erumpent from bark, sessile, cup-shaped, dark


brown, paler at the margin, minutely downy throughout, disc flat,
orange, drying b u r n t sienna, about 1 m m . across. Asci cylindric-
clavate, up to 100x8/«, 8-spored, the pore blued b y iodine; ascospores
biseriate, fusiform, straight or slightly curved, 1 4 - 3 0 x 2 - 3 / / , non-sep-
t a t e or becoming 1-septate; paraphyses slightly enlarged upwards to
3/« thick. The tissue yields a sulphur-yellow stain in K O H solution.

On dead twigs and small branches of Betula &n&Alnus, May to September.


Not uncommon.

Mollisia palustris (Roberge) Karsten, op. cit.: 195 (1871). (PI. XXIIIE.)

Apothecia scattered, superficial, saucer-shaped, flat o r slightly con-


vex when moist, grey-brown beneath, almost white at the margin,
smooth, disc less t h a n 1 m m . across, grey, drying buff, flesh thin, soft,
the outermost cells subglobose to broadly clavate, with thin brown
walls. Asci cylindric-clavate, up to 60x6/«, 8-spored, the small pore
blued b y iodine; ascospores biseriate, narrowly fusiform, more pointed
below, 9 - 1 2 ( - 1 4 ) x 1-2/«, non-septate; paraphyses slightly enlarged
upwards, to about 3 //.

On rotting stems of Juncus and other marsh plants, in summer and au-
tumn. Common.

Mollisia cornea (Berkeley & Broome) von Hohnel in S. B. K.Akad. Wiss.


Wien, Math.-Nat. Klasse. Abt. 1, 1 2 7 : 3 6 7 (1918). ( P I : X X I I I L . )
Apothecia gregarious, superficial, sessile, saucer-shaped, smooth b u t
with a minutely dentate margin, light yellowish-brown, disc flat, con-
colorous, up to 0.5 m m . across, soft-textured when fresh, drying hard
and nearly black. Asci cylindric-clavate, up to 85 x 7 /«, 8-spored; asco-
spores irregularly biseriate, fusiform, usually slightly curved, 12-18x

185
3-3.5//, usually with four oil drops, 1-septate; paraphyses often
branched, somewhat swollen at the tips.

On dead stems of Carex paniculata in February and March. Uncommon.


Although this species was most thoroughly redescribed by Massee from
the type collection at Kew it has been strangely misinterpreted by continen-
tal authors. Hence it has seemed desirable to figure it here from fresh
material kindly supplied by Mr. W. D. Graddon and compared by him with
the type. The outer two or three layers of cells in the flesh are composed
of subglobose, brown, thin-walled elements.

Mollisia chionea Massee &Crossland in Journal of B o t a n y 34:154(1896).


(PI. X X I I I H . )
Apothecia gregarious or scattered, superficial, subsessile on a small
base, cup-shaped, smooth, ivory-white, drying yellowish, disc concave,
white drying ochraceous, up to 1 m m . across, margin minutely fim-
briate. Asci cylindric-clavate, up to 5 0 x 6 / / , pore not blued b y iodine;
ascospores biseriate, cylindric-fusiform, 7 - 9 x 1 . 5 - 2 / / , non-septate;
paraphyses scanty, cylindrical.

On dead stems of Carex pendula in autumn. Not uncommon.


The outer tissue is composed of thin-walled polygonal cells, somewhat
elongated in the direction from base to margin and replaced by short, sep-
tate, obtuse hyphae at the margin. Clearly this is far from being a typical
Mollisia but a more appropriate genus for it is not obvious. Helotium in-
certum Karsten is probably an older name for the species but it is scarcely
a good Helotium or Hymenoscyphus either!

PYRENOPEZIZA Fuckel

Pyrenopeziza Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 2 3 - 2 4 : 2 9 3


(1870).
Here are conventionally placed small, blackish, Mollisia-like fungi
in which t h e apothecia can be seen to originate below the epidermis
of herbaceous stems or dead leaves. In practice it is impossible to draw
a clear distinction between t h e m and Mollisia proper and it is largely
a m a t t e r of tradition to which genus a species is assigned. Nor is at all
clear how m a n y good species there m a y be in this group. Earlier
authors tended to refer most of t h e m to " P e z i z a atrata Pers.", modern
writers are apt to assume t h a t each host genus has at least one
Pyrenopeziza species peculiar to itself, even though their morphologi-
cal differences m a y be hard to define. In these circumstances it will

186
suffice here to describe two very distinct species to give an idea of the
scope of the genus. Students who wish to pursue their studies f u r t h e r
will profit by the rather numerous specific descriptions in Nannfeldt
(1932) and by a series of papers dealing with species on Rubus and on
herbaceous stems by J . Gremmen in F u n g u s 24 (1954) and 26 (1956)
as well as the monograph b y H u t t e r in P h y t o p a t h . Zeitschr. 3 3 : 1 - 5 4
(1958).

Pyrenopeziza arctii (Phillips) Nannfeldt in Nova Acta Reg. Soc. Sci.


Up sal. I V , 8:142 (1932). (PL X X I V E . )
Apothecia erumpent, often elliptical from compression b y the host
cortex, subglobose t h e n cup-shaped, dark brown, pruinose with short,
septate, obtuse hairs, especially near the margin, disc concave, grey-
brown, less t h a n 1 m m . across. Asci cylindric-clavate, up to 8 5 x 1 0 / / ,
8-spored, the small pore blued by iodine; ascospores biseriate, dis-
tinctly clavate, pointed below, 2 5 - 3 0 x 3 / / , 3-septate; paraphyses
cylindrical, about 3 /< wide.

On dead stems of Arctium, overwintered from the previous year, in spring


and summer. Not common. In view of the unsatisfactory value of the
erumpent habit as a generic character this species might more naturally
be sought in Belonopsis.

Pyrenopeziza foliicola (Karsten) Saccardo in Michelia 1 : 6 5 (June


1877). (PL XXIVF.)
Apothecia scattered, erumpent, subglobose t h e n cup-shaped,
blackish-brown with a whitish fimbriate margin, disc concave, grey,
about 0.25 m m . across. Asci cylindric-clavate, up to 4 0 x 6 / / , 8-spored,
the small pore blued by iodine; ascospores narrowly clavate, 6 - 9 x 1.5 /t,
nonseptate, biseriate; paraphyses cylindrical, about 1 // wide.

On rotting leaves of Alnus in April and May. Probably common but very
small, inconspicuous and seldom collected.

P. fuckelii Nannfeldt in Svensk Bot. Tidskr. 25:21 (1931), is a simi-


lar species on rotting leaves of Salix.

P. petiolaris (Albertini & Schweinitz ex Fries) Nannfeldt, op. cit.:


158 (1932), is common on the petioles of fallen leaves of Acer pseudo-
platanus and is worth mentioning because its elongated apothecia
have led it to be placed originally in Hysterium.

187
H Y S T E R O P E Z I Z E L L A von Hohnel

Hysteropezizella von Hohnel in S.B. Akad. Wiss. Wien Math.-Nat.


Klasse Abt. 1, 126:310 (1917).
In this genus have been collected a considerable n u m b e r of very
minute dark coloured Dermateaceae which arise deep in the tissues of
various Gramineae, Gyperaceac and Juncaceae. As a rule t h e y appear
to the eye as minute holes like insect punctures in the surface of leaf or
culm b u t when t h e y are expanded in d a m p weather the hole can be
seen to be occupied b y a minute, usually greyish, soft-fleshed apothc-
cium. Nannfeldt (1932) listed 32 species b u t comparatively few have
yet been recognised in Britain. In the subgenus Hysterostegiella the
paraphyses are broadly lanceolate, longer t h a n the asci, like those of a
Dasyscyphus or some species of Phialea.

Hysteropezizella (Hysterostegiella) valvata (Montagne) Nannfeldt in


Nova Acta Reg. Soc. Sci. Upsal. I V , 8:115 (1932). ( P L X X I I I F . )
Apothecia elliptical, up to 1 m m . long, developed within the tissue
of a dead leaf and covered at first by an elliptical p a t c h of discoloured
grey epidermis. This is t h r o w n off at m a t u r i t y to expose the apothe-
cium which t h e n appears sessile, saucer-shaped, smooth dark grey,
seated in a cavity in the leaf. Asci cylindric-clavate, up to 55x5/«,
8-spored, the small pore blued b y iodine; ascospores biseriate, nar-
rowly clavate, 6 - 7 x 1 . 5 / / , non-septate; paraphyses lanceolate, colour-
less, 5 n wide, sharp-pointed and up to 20 /« longer t h a n the asci.

On dead leaves of Ammophila arenaria, in autumn. Apparently uncom-


mon.

Hysteropezizella exigua (Desmazieres) Nannfeldt, op. cit.: 115 (1932).


(PL XXIIII.)

Apothecia cup-shaped, circular or somewhat angular in outline,


sunken in the leaf tissue and exposed b y a small pore, disc concave,
grey-brown, up to 0.3 m m . across. Asci cylindric-clavate, up to 70x9/«,
8-spored, the pore blued b y iodine; ascospores biseriate, fusiform-
clavate, 15-18x3-4/«, with two oil drops, becoming 1-septate; para-
physes cylindrical, somewhat enlarged at the rounded tip.
On dead stems of J uncus articulatus, and apparently this species on
J uncus subnodulosus, in spring and summer.

Hysteropezizella fenestrata (Desmazieres) Nannfeldt, op. cit.: 114


(1932), occurs on Scirpus lacustris and S. tabernaemontani.

188
DREPANOPEZIZA (Klebahn) von Hohnel

Drepanopeziza (Klebahn) von Hohnel in Annales mycologici 15:332


(1917).

Apothecia dark coloured, very soft-fleshed, erumpent from dead


leaves, not associated with a stroma within the leaf tissue, obconical,
smooth, very minute, asci 8-spored, the pore blued b y iodine, asco-
spores, hyaline, non-septate. There is an allied genus, Diplocarpon
Wolf, in Botanical Gazette 54:231 (1912), in which the ascospores are
1-septate; two species, D. rosae Wolf on Rosa op. cit.:231 (1912), and
D. earliana (Ellis & E v e r h a r t ) Wolf in J o u r n . Elisha Mitchell Sci. Soc.
39:158 (1924), on Fragaria, are common parasites of economic plants
b u t as their apothecia have yet to be recognised in England t h e y will
not be described here. Pseudopeziza differs in its usually much larger
apothecia erumpent from a stroma.

Drepanopeziza ribis (Klebahn) von Hohnel, op. cit. (1917). (PL XXVG.)

Apothecia developed beneath the epidermis of fallen leaves, t h e n


erumpent, obconical with a flat disc and small base, smooth, dark
brown, disc about 0.3 m m . across. Asci cylindric-clavate, up to 7 0 x 1 5 /«,
8-pored, the pore blued b y iodine; ascospores biseriate, elliptical,
10-14x5-6/«, non-septate b u t with an oil drop near each end; para-
physes slender, slightly swollen up to 3.5 fi thick at the tip.

On leaves of Ribes species which have been shed after developing a leaf-
spot disease associated with the conidial state Gloeosporidiella ribis (Libert)
Petrak. For a detailed account of the forms on different species of Ribes see
Miiller et al in Svdowia 12:407-413 (1958).

P S E U D O P E Z I Z A Fuckel

Pseudopeziza Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:290


(1870).

Apothecia erumpent from a stroma within the host tissue, soft-


fleshed, light coloured or with a dark exterior and paler disc, asci
8-spored, the pore i~i most species blued b y iodine, ascospores non-
septate. For a monograph of this and the related genera Leptotrochila
and Pseudorhytisma see Schiiepp in P h y t o p a t h . Zeitschr. 36:213-269
(1959).

189

£
Pseudopeziza trifolii (Bivona-Bernardi) Fuckel, op. cit.:290 (1870).
(PI. X X V E . )

Apothecia erumpent from small dead spots of leaf tissue, saucer-


shaped, exterior brown, closely adpressed to the leaf, smooth, disc flat,
about 0.5 m m . across, yellowish-grey. Asci cylindric-clavate, up to
8 0 x 1 4 / / , 8-spored, the pore not blued by iodine; ascospores irregularly
biseriate, slightly ovoid, 1 0 - 1 2 x 5 - 5 . 5 / / , non-septate b u t with a small
oil drop near each end; paraphyses about 3 - 4 / / wide near the tip.
There is no t r u e cellular excipulum flanking the hymenium as in
Leptotrochila, Fabraea and Pyrenopeziza.

On living leaves of Trifoliurn species, May to November. Very common


on T. repens, less so on T. pratense.
Species of Medicago are attacked by the very similar P. medicaginis
(Libert) Saccardo; in Malpighia 1:455 (1887); P. cerastiorum (Wallroth)
Schroeter in Rabenhorst's Fungi europaei 1705 (1874), is common on leaves
of Cerastium and P. repanda (Fries) Karsten in Acta Soc. Fauna Flora
Fennica 2 (6): 161 (1885), occurs on those of Asperula, Galium and Sher-
ardia. The two last named are referred by Schuepp to Leptotrochila.

Pseudopeziza calthae (Phillips) Massee, Brit. Fungus Flora 4:192


(1895) = Fabraea rousseauana Saccardo & Bommer in Bull. Soc. Roy.
Bot. Beige 25:168 (1886), occurs on leaves of Caltha palustris.

FABRAEA Saccardo

Fabraea Saccardo in Michelia 2:331 (1881).


This genus was erected for species akin to Pseudopeziza b u t with
1-septate ascospores. It is united with Leptotrochila Karsten 1871 b y
Schuepp, which is separated from Pseudopeziza b y having a well
developed p a r e n c h y m a t o u s excipulum and an ascus pore blued by
iodine.

Fabraea ranunculi (Fries) Karsten in Acta Soc. F a u n a Flora Fennica 2


(6): 161 (1885). (PI. X X V J . )
Apothecia crowded on brownish spots of leaf tissue, e r u m p e n t ,
saucer-shaped, blackish brown beneath, disc flat, yellowish, 0.5 m m .
across with a dark brown somewhat dentate margin. Asci cylindric-
clavate, up to 8 0 x 1 4 / / , 8-spored, the ascus pore deep blue in iodine;
ascospores biseriate, distinctly clavate, l l - 1 5 ( - 1 7 ) x 4 - 6 / / , with one
septum nearer the narrow e n d ; paraphyses slender, up to 3// thick.

190
On living leaves of Ranunculus acris, R. repens etc., June to December.
Very common.

SPILOPODIA Boudier

Spilopodia Boudier in Bull. Soc. mycol. France 1 : 1 2 0 (1885).


Apothecia dark coloured like those of a Pyrenopeziza but erumpent
from a hyphal strand associated with a vascular bundle of the host;
asci and ascospores like those of Pseudopeziza.

Spilopodia nerviscquia (Fries) Boudier, Hist. Class. Discom. d ' E u r o p e :


143 (1907). (PI. X X I V J . )
Apothecia erumpent in rows along a m a j o r vein of the host leaf,
sessile, cup-shaped, smooth b u t with a prominently toothed margin,
disc grey, up to 1 m m . across. Asci cylindric-clavate up to 6 0 x 1 0 / / ,
8-spored, the pore blued b y iodine; ascospores biseriate, narrowly
elliptical, 1 0 - 1 2 x 3 - 4 / / , non-septate; paraphyses slender, slightly en-
larged upwards.

On dead leaves of Plantago lanceolata. Rare.

Spilopodia melanogramma Boudier in Bull. Soc. mycol. France 1:120


(1885), occurs on dead stems of Mercurialis perennis.

CALLORIA Fries

Calloria Fries, Floram Scanicam:343 (1835).


Apothecia erumpent, becoming superficial and flattened, light
coloured, tissue composed of thin-walled subglobose cells, asci 8-spored,
t h e pore not blued b y iodine, ascospores at least 1-septate. The apo-
thecia bear some superficial resemblance to those of Orbilia b u t have
quite different hymenial elements.

Calloria fusarioides (Berkeley) Fries, S u m m a Veg. Scand. Sect. P o s t . :


359 (1849). (PL XXVD.)
Apothecia gregarious, e r u m p e n t , soon becoming superficial, saucer-
shaped or discoid, pinkish or orange coloured, disc about 1 m m . across,
soft-fleshed. Asci cylindric-clavate, up to 9 5 x 1 0 / / , 8-spored, the pore
not blued by iodine; ascospores biseriate, elliptic-cylindric, 9 - 1 4 x

191
3.5-4/«, hyaline, with one median s e p t u m ; paraphyses slender, ab-
r u p t l y swollen to about 3/t wide at t h e tip.

On dead stems ot Urtica dioica, in spring. Uncommon though the assumed


conidial state Cylindrocolla urticae (Persoon) Bonorden is very abundant
everywhere on the same substrate. Peziza fusarioides Berkeley was selected
by Boudier 1885 as Neotypus ot Calloria and has been accepted as such by
subsequent authors though Fries clearly did not regard it as a typical re-
presentative of his genus. The name C. carneo-flavida Rehm in Annal. mycol.
3:412 (1905) has been given to a small form, also on Urtica, with apothecia
only y i mm. diameter.

ORBILIOPSIS von Hohnel

Orbiliopsis von Hohnel in Mitt. Bot. Inst. Techn. Hochsch. Wien 3:101
(1926).
Apothecia subsessile, superficial, on wood or bark, light coloured,
soft-fleshed, the outer tissue composed of thin-walled cubical or sub-
globose cells, asci 8-spored, ascospores hyaline, elliptic-cylindric,
0-1-septate. This generic name is a n t e d a t e d b y Orbiliopsis Sydow in
Annales mycologici 22:308 (1934), used in a different sense, and hence
m u s t eventually be either conserved or replaced. In view of the great
confusion t h a t still exists regarding the precise application of generic
names in the ascomycetes it would be rash to risk proposing a new
name for it at present.

Orbiliopsis subcarnea ([Schumacher] Saccardo) von Hohnel, op. cit.:


102 (1926). (PI. XXVK.)
Apothecia subsessile, saucer-shaped with a r a t h e r small base,
smooth, soft, pale purplish-pink t h r o u g h o u t , disc flat, scarcely 1 m m .
across. Asci cylindric-clavate, a b o u t 7 0 x 7 / / , 8-spored, t h e r a t h e r
broad pore blued b y iodine; ascospores irregularly biseriate, elliptic-
cylindric, 9-12(-13) x 2.5-4 fi, with two oil drops b u t non-septate; para-
physes often forked, slender, swollen up to 3/« at the obtuse tip.

On decorticated wood. Not common. This is one of the species which has
featured in British books as "Helotium ferrugineum"; it is clearly no Helo-
tium but forms one of a group of allied species which urgently need study.
The name under which they are to be known is a secondary consideration
which cannot usefully be argued until the species themselves have been
adequately defined. This need for fundamental study based on ample fresh
collections is the justification for drawing attention to the group here.

192
PODOPHACIDIUM Niessl

Podophacidium Niessl in Rabenhorst's Fungi europaei: 1153 (1868).


Apothecia subsessile, terrestrial, externally dark coloured w i t h a
tissue composed of globose cells with dark brown walls, flesh composed
of similar b u t colourless cells, asci 8-spored, pore blued b y iodine;
paraphyses slender, sometimes branched. There is one species:

Podophacidium xanthomelum (Persoon) K a v i n a in Crypt. Czech.


Exsicc. 217 (1936). (Fig. 8F.)
Apothecia gregarious, cup-shaped or obconical with a small base,
soft, externally almost black with a m i n u t e l y scurfy surface, disc flat,
up to 4 m m . across, sulphur-yellow, with a prominent, black, toothed
margin. Asci cylindric-clavate with long stalks, u p to 150x10//,
8-spored, the r a t h e r broad pore deep blue w i t h iodine; ascospores uni-
seriate, elliptic-fusiform, 13-15x5-6/«, with two large oil drops; para-
physes slender, about 2.5/« thick at t h e tip.

On soil and debris, chiefly in coniferous woods, summer and autumn. Un-
common.

CRYPTODISCUS Corda

Cryptodiscus Corda, Icones F u n g o r u m 2:37 (1838).


Apothecia deeply sunk in decorticated wood, flesh very t h i n and
adhering firmly to the wood, composed of a few layers of small, thin-
walled, subglobose cells, asci 8-spored, ascospores hyaline, normally
3-septate, paraphyses slender, forked.

Cryptodiscus pallidus [Persoon] Corda, op. cit.:37 (1838). (PI. XXIIM.)

Apothecia immersed, white or cream-coloured, scattered, disc


elliptical, concave and contracted to a narrow slit when dry. Asci
clavate, somewhat thickened at the broadly rounded apex, up to
60 x l 5 / « , pore not blued b y iodine, 8-spored; ascospores irregularly
biseriate, elliptical with rounded ends, 10-16x5-6/<, hyaline, 3-septate,
occasionally 4-septate; paraphyses v e r y numerous, slender, forked,
slightly enlarged to 2/t at the rounded tip. The h y m e n i u m has a ten-
dency to stain blue as a whole with iodine.

In decaying decorticated wood of deciduous trees. Rare.

193
PROPOLIS (Fries) Fries

Propolis (Fries) Fries, S u m m a Veg. Scand. Sect. Post.:372 (1849).


Apothecia immersed in woody tissues, without a conspicuous mar-
ginal tissue, disc light coloured, pruinose, asci 8-spored, ascospores
large, hyaline, non-septate, paraphyses slender, forked.

Propolis versicolor (Fries) Fries, op. cit.:372 (1849). (Fig. 28E.)


Apothecia scattered, immersed, exposed b y shedding of the epi-
dermis or bark and surrounded b y a t o r n margin of host tissue, disc
circular or elliptical, flat, up to 5 m m . across, hymenium brown b u t
concealed by a white pruina, subhymenial tissue v e r y thin, formed of
v e r y slender, hyaline, interwoven h y p h a e about 1 // thick, marginal
tissue lining and closely adhering to the u p t u r n e d margin of host tissue
and itself forming a vertical rim to the disc, 30-40/i thick, composed of
angular hyaline cells 3-4/« across. There is usually a small a m o u n t of
stromatic tissue beneath the b a r k immediately surrounding t h e apo-
thecium. Asci cylindric-clavate, rounded above, up to 135x18// thin-
walled, the pore not blued b y iodine, 8-spored; ascospores biseriate,
hyaline, slightly kidney-shaped, 2 0 - 2 7 x 6 - 8 / / , non-septate; para-
physes very numerous, thread-like, forked repeatedly near the tip,
their apices sometimes slightly swollen up to 2// thick, longer t h a n the
asci.

On dead wood, branches, woody stems, pine cones etc. throughout the
year. Very common.
There are British collections at Kew on Acer, Chamaenerion, Corylus,
Fagus, Fraxinus, Ilex, Larix, Lonicera, Malus, Pinus, Populus, Prunus,
Quercus, Rhamnus, Salix, Sambucus, Sorbus and Tilia. The form on Pinus
cones is sometimes distinguished as P. rhodoleuca Fr. op. cit.:372 (1849),
and Rehm recognised a var. betulae (Fuckel) Rehm in Rabenh. Krypt. Flora
1 (3): 150 (1888) on Retula.
The affinities of this species are still obscure. The asci and paraphyses
recall those of Polydesmia and the immersed apothecia are suggestive of
Phacidiaceae but it would be quite isolated there also.

PLOETTNERA P. Hennings

Ploettnera P. Hennings in Verh. Bot. Ver. Prov. Brandenburg 41:94


(1899).
Apothecia erumpent, surrounded b y a ring of upraised epidermis,
sessile, disc flat, dark blue-green, marginal tissue reduced to a thin

194
layer of very fine interwoven hyphae, asci clavate, 8-spored, ascospores
hyaline, nonseptate.

Ploettnera exigua (Niessl) von Hohnel in S.B. Akad. Wiss. Wien,


Math.-Nat. Klasse A b t . 1, 1 2 7 : 5 5 6 (1918). (PI. X X I I K . )
Apothecia sessile, erumpent, surrounded b y the t o r n epidermis, disc
flat, dark blue-green, elliptical or nearly circular, up to 0.3 m m . across,
asci clavate, very short-stalked, up to 6 0 x 1 0 / / , broadly rounded
above, the pore not blued b y iodine; ascospores biseriate, elliptical,
1 2 - 1 5 x 5 - 8 / t , hyaline, with two oil drops b u t not septate; paraphyses
very slender, b u t enlarged to 5 ft wide in the blue-green apical cell.

On dead stems of Rubus "frulicosus", in winter. Apparently rare.

TROCHILA Fries

Trochila Fries, S u m m a Veg. Scand. Sect. P o s t . : 3 6 7 (1849).


Apothecia sunk in the tissue of dead leaves, exposed either b y shed-
ding the overlying p a t c h of epidermis or b y its splitting into a n u m b e r
of lobes, which curl back to expose the disc, seated on a basal layer of
dark subglobose cells b u t lacking any comparable marginal tissue, asci
8-spored, the pore blued by iodine, ascospores elliptical, hyaline or
eventually brownish, non-septate.

Trochila craterium Fries, op. cit.:367 (1849). (PI. XXVF.)

Apothecia immersed in dead leaf tissue, circular, opening b y a


variable n u m b e r of irregularly t o r n t e e t h to expose the flat, dark-
brown or greenish-brown disc, up to 0.4 m m . across. Asci clavate, up
to 60x12,«, 8-spored, the small pore blued by iodine; ascospores bi-
seriate, broadly elliptical, 6 - 9 x 4 - 5 / / , hyaline; paraphyses somewhat
clavate, up to 6// wide at the tip.

On the under side of dead leaves of Hedera helix. Fairly common.

Trochila laurocerasi (Desmazieres) Fries, op. cit.:367 (1849), is a


very similar fungus on dead leaves of Prunus laurocerasus.

Trochila ilicina (Nees ex Fries) Greehalgh & Morgan-Jones in Trans.


Brit, mycol. Soc. 4 7 : 3 1 2 (1964). (PI. X X V H . )
Apothecia scattered, immersed in dead leaf tissue, opening b y shed-
ding a circular p a t c h of host epidermis, disc circular or irregular if two

195
or more fructifications have become confluent, up to 1 m m . across,
olive-grey. Asci clavate, up to 8 0 x l 0 / t , the pore blued b y iodine,
8-spored; ascospores biseriate, elliptical, 9 - 1 2 x 3 . 5 - 4 . 5 f i , hyaline, non-
septate; paraphyses clavate, up to b/j, wide at the tip.
On dead and fallen leaves of Ilex aquifolium. Very common.
The name T. ilicis (Chevalier) rouan in general use for this plant is
a later homonym of Trochila ilicis (Schleicher) Fries, op. cit.: 367 (1849)
based on a different type. The fungus is aiso often called Stegia ilicis Fries,
Summa Veg. Scand. Sect. Post.:370 (1849).

NAEVIA Fries

Naevia Fries emend von Hohnel in Annales mycologici 15:300 (1917).


H y m e n i u m subepidermal, seated on a thin colourless tissue com-
posed of subglobose cells, replaced at the margin b y a layer of thin,
dark brown, prismatic cells coherent with the epidermis, which tears
into a n u m b e r of triangular lobes to expose t h e disc, asci small, the
pore not blued b y iodine, ascospores hyaline, non-septate. This is the
concept of the genus accepted b y Nannfeldt (1932) b u t it is difficult to
reconcile w i t h t h e original circumscription b y Fries.

Naevia minutissima (Auerswald) Rehm, Rabenhorst's Kryptogamen


Flora Deutschland 1 (3): 138 (1888.) ( P I . X X V A . )
Apothecia scattered, immersed in t h e tissue of dead leaves, disc flat,
cream-coloured, a b o u t 0.2 m m . wide, covered b y dark brown tissue
which splits into three or four triangular lobes to expose t h e hyme-
nium. Asci elliptic-clavate, short-stalked, u p to 4 0 x 9 / t , 8-spored, pore
not blued b y iodine (Rehm says it is tinged violet); ascospores biseriate,
ovate, 7 - 8 x 3 - 3 . 5 / « , hyaline, non-septate; paraphyses slender, swollen
at the tip.
On the under side of fallen leaves of Quercus. Seldom collected.
Towards the tip of the lobes the brown covering tissue may be composed
of subglobose cells 5-6/« wide.

LAETINAEVIA Nannfeldt

Laetinaevia Nannfeldt in Nova Acta Reg. Soc. Sci. Upsal. IV, 8:190
(1932).
Differs from Naevia in its marginal tissue being composed of thin-
walled, colourless or only slightly brownish cells.

196
Laetinaevia tithymalina (J. Kunze) Dennis comb. nov.
Calloria tithymalina J . Kunze in Hedwigia 1 5 : 1 0 5 ( 1 8 7 6 ) . (PL XXVB.)

Apothecia scattered, immersed in dead stems, disc circular or


broadly elliptical, deep flesh-pink, less t h a n 1 m m . across, exposed b y
the overlying tissue splitting into a b o u t four lobes. Asci cylindric-
clavate, up to 7 5 x 1 2 / / , pore not blued b y iodine, 8-spored; ascospores
somewhat ovate, biseriate, 9 - 1 2 x 5 - 7 / / , hyaline, non-septate; para-
physes slender, swollen at the tip.

On dead stems of Euphorbia amygdaloides, E. cyparissias etc. Uncommon.


Rehm found asci up to 140x18// with the pore blued by iodine.

Laetinaevia tripolii (Berkeley & Broome) Dennis E d . 1 : 1 2 4

Peziza tripolii Berkeley & Broome in Ann. Mag. n a t . Hist. IV, 17:143
(1876). (Fig. 8E.)
Apothecia scattered, covered b y dark brown patches of epidermis,
erumpent by this splitting into from two to four lobes, disc ochraceous,
about 0.6 m m . across, flat. Asci cylindric-clavate, up to 8 0 x 1 1 / / , the
r a t h e r broad pore not blued b y iodine, usually 4-spored; ascospores
uniseriate, broadly fusiform, 1 1 - 1 5 x 6 - 7 / / , hyaline, non-septate; para-
physes slightly clavate, up to 4// wide at t h e tip.

On dead stems of Aster tripolium, in autumn.


Occasional asci contain more than four spores, in which case the additional
ascospores are smaller than the rest.

D U E B E N I A Fries

Duebenia Fries, S u m m a Veg. Scand. Sect. P o s t . : 3 5 6 (1849).


Apothecia immersed in t h e tissue of herbaceous stems, elongated or
flexuous, with a purplish-brown marginal tissue, erumpent, asci 4-8-
spored, ascospores hyaline, non-septate.

Duebenia purpurascens (Rehm) Nannfeldt in Svensk. Bot. Tidsskr. 23:


321 (1929). (PL X X V c . )
Apothecia gregarious, up to 3 m m . long and 0.5 m m . wide, with a
deep purplish-brown marginal tissue which p a r t s to expose a flat
reddish-orange disc. Asci cylindric-clavate, up to 5 5 x 6 / t , pore not
blued b y iodine, 4-7-spored; ascospores elliptic-cylindric or slightly
curved, 6 - 1 0 x 3 / / , uniseriate, hyaline, non-septate; paraphyses slender,
distinctly clavate and up to 3/t wide at t h e tip.

197
On dead herbaceous stems, as of Chenopodium and Leguminosae. Rare.

Order 6. PHACIDIALES

The Phacidiales as here interpreted in a conservative sense are a


heterogeneous group of genera with little in common beyond the pos-
session of apothecia immersed in host tissue, with their flesh reduced
to a minimum and firmly united with t h a t of the host. They include
both ascohymenial and ascolocular genera.
Two fairly n a t u r a l families can be segregated, the Hypodermataceae
and the Cryptomycetaceae b u t there t h e n remains a large residual
group of genera here keyed out together under the Phacidiaceae.
Several of these are still in need of critical study. Those genera of
Dermateaceae with e r u m p e n t much reduced apothecia closely re-
semble the Phacidiales and are also inserted in the key for the sake of
cross reference.

Family 1. Hypodermataceae

Apothecia mostly elongated, embedded in the substrate and firmly


united with the overlying tissue to form a superficial shield or clypeus
of combined host and fungus tissue, which opens b y a longitudinal slit
(by lobes in Coccomyces) to expose the hymenium, ascospores hyaline,
ranging from elliptic-cylindric to filiform. Many species occur on
needles of conifers and these have been ably monographed b y Darker
in Contributions from the Arnold Arboretum 1 (1932).

I. Several hymenia erumpent from a common black stroma Rhyfisma


II. Hymenia developed singly:
A. Apothecia immersed in bark, often curved or forked . . Colpoma
B. Apothecia subcuticular or subepidermal on leaves, herbaceous
stems, grasses etc.:
a. Ascospores filiform, fasciculate, clypeus black
Lophodermium (p. 200)
b. Ascospores elliptical, clavate or cylindrical, rarely fasciculate:
*Ascospores rod-shaped or fusiform, less than one-quarter
the length of the ascus, clypeus black
Hypoderma (p. 202)
**Ascospores clavate, one-quarter to three-quarters the
length of the ascus . . . . Hypodermella (p. 202)
C. Circular or nearly so, opening by teeth . . . Coccomyces (p. 203)
See also Darker in Canadian Journal of Botany 45 :1399-1414 (1967)

198
RHYTISMA Fries

Rhytisma Fries, Systema mycologicum 2:565 (1823).


Hymenia embedded in a flattened sclerotium-like stroma within the
tissue of a leaf, stroma with a compact white flesh and black shiny
crust, asci 8-spored, ascospores filiform, hyaline.

Rhytisma acerinum (Persoon ex St. Amans) Fries, op. cit.: 569 (1823).
(Fig. 9c.)
S t r o m a t a more or less circular, up to 20 m m . across, forming shining
black, slightly raised patches on the upper surface of leaves, containing
numerous elliptical apothecia which become exposed b y longitudinal
or forked fissures in the crust, disc grey, soft. Asci clavate, up to
130x10//, 8-spored, with a conical a p e x ; ascospores fasciculate, fili-
form b u t slightly thicker at the upper end, 6 0 - 8 0 x 1 . 5 - 2 . 5 / / , hyaline;
paraphyses slender, curled and sometimes forked at the tip, about 1.5//
thick.

On leaves of Acer pseudoplalanus, very common, occasionally on A. cam-


pestre. The stromata are abundant on living leaves from July onwards but
then contain only the conidial state, Melasmia acerina Leveille. Apothecia
develop in the stromata on fallen leaves during the winter and can be
collected on these from April to June.

Rhytisma salicinum Fries, op. cit.:568 (1823), is a similar fungus on


leaves of Salices.
Rhytisma andromedae Fries, op. cit.:567 (1823), occurs on Andro-
meda polifolia.

COLPOMA Wallroth

Colpoma Wallroth, Flora Cryptogamica germaniae 2:422 (1833).


Apothecia developed beneath bark, narrow, elongated, often flexu-
ose, black, opening b y a longitudinal slit, asci clavate, 8-spored, asco-
spores fasciculate, filiform, hyaline; paraphyses with filiform curled
tips. The genus is sometimes called Clythris Fries.

Colpoma quercinum (Persoon) Wallroth, op. cit.: 423 (1823). (Fig. 8A.)
Apothecia gregarious, elongated, usually developed transversely to
the long axis of the twig on which t h e y grow, originating beneath the
bark and raising it in straight or curved ridges which split longitudin-

199
ally and p a r t to expose the black fungus tissue. This also splits and
opens in d a m p weather, revealing the light yellow disc which m a y be
up to 1 5 x 2 m m . Asci clavate, up to 150x10/«, 8-spored; ascospores
fasciculate, filiform, obtuse above, pointed below, 80-95 x 1.5 /j,, hyaline,
becoming m u l t i s e p t a t e ; paraphyses slender, curled at the tip, 1.5-2fi
thick, longer t h a n the asci.
On twigs of Quercus, May to July. Common, a weak parasite.

Colpoma degenerans (Fries) de Notaris in Giorn. bot. ital. 2 (2): 41


(1847). (Fig. 8D.)
Apothecia similar to those of C. quercinum b u t arranged longitudin-
ally on t h e twig, e r u m p e n t , black, splitting longitudinally to expose
t h e yellow disc. Asci clavate, apex conical, up to 170x15,«, 8-spored;
ascospores filiform, obtuse above, pointed below, 80-90x1.5-2/«,
hyaline; paraphyses slender.
On dead twigs of Vaccinium uliginosum. Rare. Also called Sporomega de-
generans (Fr.) Corda.

Colpoma juniperi (Karsten) Dennis in Kew Bull. 1957:401 (1958)


occurs on twigs of Juniperus communis and C. crispum (Persoon ex
Fries) Saccardo Sylloge F u n g o r u m 9:1127 (1891) on those of other
conifers.

LOPHODERMIUM Chevalier

Lophermium Chevalier, Flor. Gen.-Paris 1:435 (1826) emend de Nota-


ris in Giorn. bot. ital. 2 (2): 41 (1847).
Apothecia elliptical or elongated, developed either within or beneath
t h e epidermis of flowering plants which t h e y commonly lift to form
small black blisters, w i t h a black covering tissue which splits longi-
tudinally to expose t h e disc, asci cylindric-clavate, 8-spored or 4-spor-
ed, ascospores filiform, non-septate, hyaline, fasciculate, paraphyses
slender. Some species show a peculiarly intricate interlocking p a t t e r n
in the h y p h a l ends of the mycelium forming t h e fringe of the black
covering tissue. Presence of this "aliform mycelium" has been used in
conjunction with t h e place of origin of the apothecium, within or under
the epidermis, to divide this large genus into smaller units. This sub-
division has not met with general acceptance, however, and will not be
adopted here. The student will find useful the "Monographic re-
arrangement of L o p h o d e r m i u m " b y L. R. Tehon, Illinois Biological

200
Monographs 13, No. 4 (1935), b u t should be warned t h a t t h r o u g h mis-
understanding the principles of nomenclature he has absurdly trans-
ferred the name L. pinastri to L. piceae and called the true L. pinastri
by an unnecessary new name.

Lophodermium juniperinum (Fries) de Notaris, op. cit.: 46 (1847).


(Fig. 9 G . )
Apothecia arising beneath the cuticle, usually on the outer surface
of leaves, elliptical, up to a b o u t 1 m m . long b y 0.4 m m . wide, black,
blisterlike, margin composed of aliform mycelium, disc soft, whitish.
Asci cylindric-clavate, up to 130x17/«, 8-spored; ascospores fascicu-
late, filiform, 60-100x2/«, with a t h i n hyaline gelatinous coating; para-
physes slender, curled at the tips.
On dead leaves of Juniperus communis and J. nana. Common.

Other British species include:


L. arundinaceum (Schrader ex Fries) Chevalier, op. cit.:435 (1826),
on Ammophila, Phragmites etc.
L. culmigenum (Fries) de Notaris, op. cit.:47 (1847), on Elymus and
Deschampsia.
L. gramineum (Fries) Chevalier, op. cit.:435 (1826), on Poa and
apparently other species on other grasses.
L. caricinum (Roberge) D u b y in Mem. Soc. Phys. et Hist. N a t .
Geneve 16:59 (1862), in the larger Carices.
L. typhinum (Fries) L a m b o t t e , Flora mycologique de Beige 2:452
(1880), on Typha.
L. macrosporum (Hartig) Rehm in Rabenhorst's K r y p t o g a m e n Flora
Deutschland 1 (3): 45 (1887), on Picea abies needles.
L. piceae (Fuckel) von Hohnel in S.B. K. Akad. Wiss. Wien, Math.-
Nat. Klasse A b t . 1 , 1 2 6 : 2 9 6 (1917), on needles of Abies and Picea.
L. pinastri (Schrader ex Fries) Chevalier, op. cit. 1:436 (1826), on
needles of species of Pinus.
L. maculare (Fries) de Notaris in Giorn. bot. ital. 2 (2): 45 (1847), on
Vaccinium.
L. melaleucum (Fries) de Notaris, op. cit.:44 (1847), on Vaccinium.
L. rhododendri Cesati ex Saccardo, Sylloge F u n g o r u m 2 : 7 9 3 (1883),
on Rhododendron ponticum etc.
L. vagulum Wilson & Robertson in Trans. Roy. Soc. E d i n b u r g h 56:
524 (1947), on Chinese rhododendrons.
L. hysterioides (Persoon) Saccardo, Sylloge F u n g o r u m 2 : 7 9 1 (1883),
on leaves of Crataegus, Berberis etc.
201
HYPOI)ERMA de Candolle

Hypodcrma de Candolle ex Merat, Nouvelle Flore Paris 1:152 (1821),


emend de Notaris in Giorn. bot. ital. 2 (2): 35 (1847).
Apothecia superficially like those of Lophodermium but containing
much shorter spores which are not fasciculate in the ascus.

Hypodcrma dcsmazieri D u b y in Mem. Soc. Phys. et Hist. Nat,. Geneve


16:42 (1862). (Fig. 9E.)
Apothecia usually on the outer surface of needles, developed beneath
the epidermis and raising it into a low black blister, up to 1 m m . long,
surrounded by a narrow grey zone, opening by a longitudinal slit. Asci
about 100x15//, up to 1 5 0 x 1 7 / / according to Darker, 8-spored; asco-
spores biseriate, fusiform-clavate, 2 5 - 3 5 x 4 . 5 - 5 / / , hyaline and en-
veloped in a gelatinous capsule 4 - 5 / / t h i c k ; paraphyses slender, often
curled or forked at the tip.
On needles of Pinus sylvestris and other species. Common.

Hypodcrma virgultorum de Candolle ex St. Amans, Flore Agenaise: 515


(1821). (Fig. 9F.)
Apothecia similar to the preceding species b u t m u c h larger, up to
2 mm. long, % m m . wide and without the surrounding grey zone. Asci
clavate, up to 120x10//, 8-spored; ascospores irregularly arranged in
two or three rows, cylindric-fusiform, 2 1 - 2 4 x 3 - 4 / / , hyaline, containing
two large oil bodies b u t as a rule not septate; paraphyses slender,
flexuous, curled at the tip.
On dead stems of Rubus. Common, also known as Hypodcrma rubi (Per-
soon ex Fries) de Notaris, op. cit.:37 (1847).
H. commune (Fries) Duby, op. cit.:41 (1862), with smaller asci and
ascospores 1 8 - 2 0 x 4 / t is common on dead herbaceous stems.
H. hederae de Notaris, op. cit.:36 (1847), occurs on dead leaves of
Heeler a.
/ / . scirpinum de Candolle ex Merat, op. cit.: 152 (1821), is found on
dead stems of Scirpus lacustris.

HYPODERMELLA von Tubeuf

Hypodermella von Tubeuf in Bot. Centralblatt 61:49 (1895).


Darker differentiated Hypodermella from Hypoderma by its clavate
ascospores, longer in proportion to the length of the ascus. The British

202
species are f u r t h e r distinguished b y having m u c h paler and more in-
conspicuous fructifications.

Hypodermella conjuncta Darker in Gontrib. Arnold Arboretum 1 : 6 0


(1932). (Fig. 8B.)
Apothecia numerous on discoloured patches, especially near the
tips of needles, immersed in the leaf tissue, elliptical, up to 3.75x0.28
m m . b u t often from two to four fuse laterally to make broad com-
pound fruit bodies, opening by a longitudinal slit along the stomatal
lines of the leaf. Asci cylindric-clavate, up to 160x16// conical
above, 8-spored; ascospores long and slender, rounded above and
pointed below, 7 5 - 9 0 x 3 . 0 - 3 . 5 / / , non-septate, hyaline, each enclosed
in a thin gelatinous sheath; paraphyses slender, 1 - 2 / / thick, slightly
swollen at the tip.

On living needles of Pinus sylvestris, in summer. Uncommon.

Hypodermella sulcigena (Rostrup) von Tubeuf, op. cit.:49 (1895),


found on needles of various species of Pinus, differs in its m u c h shorter
ascospores, 2 7 - 3 5 x 4 - 5 / / . Three other species cause needle-cast dis-
eases of Abies and Larix in Europe.

COCCOMYCES de Notaris

Coccomyces de Notaris in Giorn. bot. ital. 2 (2): 38 (1847).


Apothecia scattered, lenticular, embedded in leaf tissue, circular,
with a black covering layer which splits open b y numerous t e e t h to
expose the disc, asci 8-spored, ascospores elongated or filiform, often
multiseptate, hyaline, paraphyses filiform.

Coccomyces coronatus (Schumacher ex Fries) Karsten, Mycologia


Fennica 1:256 (1871). (Fig. 28A.)
Apothecia scattered, circular or broadly elliptical, sometimes slightly
three-cornered, black, shining, up to 3 m m . across, opening b y numer-
ous teeth to expose the soft yellowish disc. Asci clavate, up to 180 x
14/i, 8-spored; ascospores narrowly cylindrical to needle-shaped,
rounded above and pointed below, 3 0 - 7 5 x 2 - 3 . 5 / / , lying parallel in
the ascus, hyaline, multiseptate; paraphyses numerous, slender, 1 - 2 / /
thick, curled at the tip.

On fallen leaves of Betula, Fagus and Quercus, in late summer and au-
tumn. Common. This is probably a collective species in need of critical

203
study. The collection figured corresponds to the var. trigonus (Fries) Kar-
sten, op. cit.:257 (1871), with short ascospores 28-45x2-3.5/«.

Coccomyces dentatus (Kunze & Schmidt) Saccardo in Michelia 1 : 5 9


(June 1877), on dead Quercus and Castanea leaves is distinguished by
its more slender ascospores, 45-55x1.5-2/«, in apothecia up to 1 m m .
across, opening b y only four or five teeth.
Melasmia empetri Magnus in Ber. Deutsch Bot. Ges. 4 : 1 0 4 (1886),
common as a black crust on living twigs of Empetrum, is said to be t h e
conidial state of another member of this family, Duplicaria empetri
(Fries) Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:265 (1870).

Family 2. Cryptomycetaceae

Hymenium developed within a subgelatinous fleshy-stroma beneath


the b a r k of woody plants, ascospores elliptical, non-septate. There are
two genera:

I. Stroma small, covering layer unstratifled, mature fruit-body cup-shaped


Potebniamyces
II. Stroma forming a widespread crust, covering tissue stratified
Cryptomyces (p. 205)

POTEBNIAMYCES Smerlis

Potebniamyces Smerlis in Canad. J o u r n . Bot. 40:352 (1962).


The genus has commonly been called Phacidiella Potebnia in
Zeitschr. f. Pflanzenkr. 22:147 (1912) b u t t h a t is a later homonym of
Phacidiella Karsten in Hedwigia 23:85 (1884). There is a single species:

Potebniamyces discolor ( M o u t o n & Saccardo) Smerlis, op. cit.: (1962).


(PL X X X V I A . )

S t r o m a t a forming small cushions beneath bark, composed of closely


interwoven hyphae 3-4/« wide, w i t h blackish-brown walls 1 /t thick,
apothecia developed singly or in small groups within the stroma, disc
up to 1 m m . across, dark grey, exposed b y the splitting of the stroma
and surrounded b y its light greyish, cushion-like, u p t u r n e d edges
which adhere closely to the under surface of the split bark. Asci
clavate, about 150x15-18/«, 8-spored, the tip not stained b y iodine;
ascospores mostly uniseriate, ellipsoidal, 1 5 - 2 3 x 8 - 1 1 /«, hyaline, often
containing two large oil drops b u t not becoming septate; numerous

204
slender hyphae are embedded in a mucilaginous m a t r i x between t h e
asci.
Erumpent from bark of Pyrus communis and Pyrus malus, associated
with a conidial state which has been called Fuckelia conspicua Marchal, in
autumn. Uncommon.
The fungus causes one of the minor cankers of some fruit trees.

CRYPTOMYCES Greville

Cryptomyces Greville, Scottish Gryptogamic Flora 4, t a b . 206 (1826).


H y m e n i u m developed within a widespreading stroma, t h e u p p e r
portion of which is differentiated into three well defined h y p h a l tissues,
arising beneath the b a r k of twigs and small branches, which is split
and rolled back to expose the fruit body, asci 8-spored, ascospores
elliptical, non-septate.

Cryptomyces maximus (Fries) Rehm in Rabenhorst's K r y p t o g a m e n


Flora Deutschland 1 (3): 107 (1888). (Fig. 27A.)
S t r o m a forming blistered patches up to 10 cm. long on living
branches, blackish-brown with a bright yellow margin at first, soon
becoming shining black all over t h e surface, flesh white, h y m e n i u m
exposed b y the shedding of the covering tissue, yellowish-brown. Asci
cylindric-clavate, up to 250x33/<, 8-spored; ascospores irregularly
biseriate, ellipsoidal or ovoid, 2 0 - 3 0 x 1 0 - 1 3 / / , hyaline or slightly
yellowish, with a t h i n gelatinous coating, interspersed with n u m e r o u s
slender paraphysoidal h y p h a e whose tips are united b y a brown
gelatinous matrix.
On living branches and twigs of Salix. Rare.

Family 3. Phacidiaceae

Here are assembled the remaining genera not segregated in the two
preceding families:
I. Ascospores needle-shaped:
A. Apothecia circular, erumpent from bark . . . Therrya (p. 206)
B. Apothecia oblong, in needles of conifers . Naomacyclus (p. 207)
See also Lasiostictis, p. 224
II. Ascospores clavate, multiseptate Melittiosporium (p. 208)
III. Ascospores elliptical:
A. Ascospores more than 1-septate . . . . Sphaeropezia (p. 212)
See also Cryptodiscus, p. 193.

205
B. Ascospores brown, 1-septate Didymascella (p. 208)
C. Ascospores hyaline, non-septate:
1. Apothecia immersed in wood or bark:
a. With a white-pruinose disc and no distinct sterile mar-
gin. (See Propolis p. 194)
b. With a conspicuous white-pruinose margin
Laquearia (p. 213)
c. With a conspicuous black margin
Pseudophacidium (p. 209)
2. On fronds of Pteridium, ascocarps elongated, black
Cryptomycina (p. 210)
3. On needles of conifers:
a. Opening by a torn margin, ascus pore blued by iodine
Phacidium (p. 210)
b. Opening by a lid, ascus pore not blued by iodine
Rhabdocline (p. 211)
4. On leaves of Dicotyledons:
a. Developed in a stroma, ascus pore blued by iodine
Phacidiostroma (p. 211)
b. Pore not clearly blued by iodine or stroma absent. (See
Trochila (p. 195), Naevia, Laetinaevia (p. 196),
Duebenia (p. 197)
5. W7ith a blue-green disc, on Rubus. (See Ploettnera, p. 194)
D. Ascospores hyaline, 1-septate, asci 2-spored
Schizothyrioma (p. 212)

THERRYA Saccardo & Penzig

Therrya Saccardo & Penzig i s Michelia 2:604 (1882).


Apothecia immersed in b a r k , t h e n erumpent, opening b y several
lobes, more or less circular, asci clavate, 8-spored, ascospores needle-
shaped, pointed at each end, multiseptate, hyaline, paraphyses swollen
at the tip. There is one species:

Therrya pini (Albertini & Schweinitz ex Fries) K u j a l a in Communi-


cationes I n s t i t u t i Forestalis Fenniae 38 (4):49 (1950). (Fig. 28D.)

Apothecia scattered, circular, with a black covering layer which


splits open by from four to six lobes to expose the yellowish-brown
disc, u p to 3 m m . across. Asci clavate, up to 150x15//, the pore not
blued b y iodine, 8-spored; ascospores fasciculate, somewhat spirally
arranged, narrowly fusiform, pointed at each end, somewhat S-shaped,
55-80x3-4/«, hyaline to yellowish, becoming from 3 - 1 1 - s e p t a t e ; p a r a -
physes numerous, slender, swollen up to 6// wide at t h e brownish tip.

206
On dead twigs of Pinus silvestris. Uncommon. Also known as Coccopha-
cidium pini (Fries) Rehm, Rabenhorst's Krypt. Flora 1 (3):98 (1888), and
Therrya gallica Saccardo & Penzig in Michelia 2:604 (1882).

NAE MACYCLUS Fuckel

Naemacyclus Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 2 7 - 2 8 : 4 9


(1873).
Apothecia immersed, elliptic-oblong, opening b y a longitudinal slit
in the host epidermis, without a well developed covering hyphal tissue,
asci 8-spored, ascospores elongated, non-septate, hyaline, paraphyses
slender, branched.

Naemacyclus niveus (Persoon ex Fries) Saccardo in Bot. Centrabl. 18:


251 (1884). (Fig. 28F.)
Apothecia scattered, immersed, erumpent b y a single longitudinal
slit in the host epidermis, which t h e n bends back as a lobe on each side
of the apothecium, disc light grey, elliptic-oblong, up to 0.5 m m . long,
flat, without any clearly defined marginal tissue, delimited simply b y
a few slender hyphae adhering to the upraised epidermis. Asci cylin-
dric-clavate, up to 120x14//, 8-spored, pore not blued by iodine;
ascospores fasciculate, cylindrical, somewhat curved, 75-85x2.523//,
hyaline, non-septate, ends r o u n d e d ; paraphyses slender, about 1 fi
thick, sometimes forked, slightly swollen and rounded at the tips,
which seem to be embedded in an amorphous m a t r i x containing
n u m e r o u s short, slender, colourless rods.

On fallen needles of Pinus silvestris and other pines. Uncommon.

Naemacyclus caulium von Hohnel in Sitzber. K. Akad. Wiss. Wien,


Math.-nat. Kl. 115 Abt. 1:650 (1906). (Fig. 29K.)
Apothecia immersed beneath a dark gray-green p a t c h of host tissue,
exposed by splitting of the latter, usually b y four lobes, disc gray,
elliptical, up to 1 m m . long. Asci 6 0 - 6 5 x 7 / / , 8-spored, spores fascicu-
late, acicular, hyaline, 4 0 - 5 0 x 1 , 5 / / , mostly 7-septate, paraphyses
filiform, 1 // wide, slightly longer t h a n the asci.

On dead stems of Urtica doica. April. Uncommon.

207
Naemacyclus arctostaphyli (Ferdinandsen & Winge) Rehm is com-
m o n on dead leaves of Arctostaphylos uva-ursi in the Highlands, for
description and figures see Miiller et al. in Sydowia 12:416-418 (1958).

MELITTIOSPORIUM Corda

Melittiosporium Corda, Icones F u n g o r u m 2:37 (1838).


Apothecia immersed, t h e n e r u m p e n t , with well developed covering
tissue, asci 1-8-spored, ascospores multiseptate, w i t h both transverse
and longitudinal walls, hyaline to brownish.

Melittiosporium pteridinum (Phillips & Bucknall) Saccardo, Sylloge


F u n g o r u m 8:705 (1889). (Fig. 28B.)

Apothecia scattered, immersed, elliptical, exposed b y splitting t h e


host epidermis into two or more irregular lobes, with a even whitish
margin, disc light brown, u p to 1 m m . long, almost flat. Asci fusiform-
clavate, up to 8 0 x l 2 / i , 8-spored; ascospores 2-3-seriate, clavate,
pointed below, 2 8 - 4 4 x 5 - 9 ( i , hyaline, with up to eight transverse septa
and said also to develop a few longitudinal septa; paraphyses slender,
thread-like, septate, hyaline.

On dead stems of Pteridium aquilinum. March. Apparently rare.


This fungus bears little resemblance to Melittiosporium versicolor (Fries)
Corda, op. cit.:38 (1838), the type species of the genus, and it deserves
further study to elucidate its true relationships, which may lie with Crypto-
discus. C. andersoni Ellis & Everhart in Bull. Torrey bot. Club 24:469 (1897)
is similar but with ascospores only 11-16x2-3/«.

DIDYMASCELLA Maire & Saccardo

Didymascella Maire & Saccardo in Annales mycologici 1 : 4 1 8 (Septem-


ber 1903).

Apothecia immersed, e r u m p e n t , opening b y irregular teeth or b y the


shedding of a lid, asci 2-4-spored, ascospores elliptical, brown, u n -
equally 2-celled, on leaves of conifers. The genus Keithia Saccardo,
Sylloge F u n g o r u m 10:49 (1892), based on Phacidium tetrasporum
Phillips & Keith is a later h o m o n y m of Keithia B e n t h a m , Lab. Gen. et
Spec.:409 (1834).

208
Didymascella thujina (Durand) Maire in Bull. Hist. N a t . Afrique Nord
18:120 (1927). (Fig. 28K.)
Apothecia circular, immersed in leaf tissue, exposed by t h e shedding
of a disc of the covering epidermis, circular, elliptical or irregular in
outline, olive-brown up to about 1 m m . across. Asci clavate, up to
100 x 20 //, 2-spored, the pore not blued b y iodine; ascospores uniseriate,
elliptical, thick-walled, 2 2 - 2 5 x 1 5 - 1 6 / / , w i t h a septum near the upper
end, brown with finely p i t t e d walls; paraphyses branched below,
thickened upwards to 5 - 8 / / wide near the tip, olive. Finally t h e whole
apothecium falls out, leaving an e m p t y pit in t h e leaf.

On dead leaves of Thuja plicata and less often on T. occidentalis, still


ttached to the twigs, summer. Common.

Didymascella tetraspora (Phillips& Keith) Maire, op. cit.: 119 (1927).


(Fig. 28J.)
Apothecia immersed in the tissue on the u p p e r side of leaves, open-
ing by several irregular brown teeth, disc blackish-brown, a b o u t 1 m m .
across. Asci clavate, up to 175x18//, 4-spored, apex rounded, the pore
not blued by iodine; ascospores uniseriate, ovoid, 2 1 - 2 4 x 1 3 - 1 6 / / ,
olive-brown, with a septum near one e n d ; paraphyses slender, swollen
to 8 - 9 / / at the olive-coloured tip.

On living leaves of Juniperus communis. Uncommon.

P S E U D O P H A C I D I U M Karsten

Pseudophacidium Karsten in Acta Soc. F a u n a Flora Fennica 2 (6): 157


(1885).
Apothecia erumpent from woody stems, with a black covering
tissue, disc light coloured, asci 8-spored, ascospores elliptical, hyaline,
non-septate.

Pseudophacidium callunae (Karsten) Karsten, op. cit.: 157 (1885).


(Fig. 8 H . )
Apothecia e r u m p e n t , with a cushion-like stromatic base, black,
opening irregularly to expose the yellow disc, about 2 m m . across.
Asci clavate, up to 100x12//, 8-spored, the pore not blued b y iodine;
ascospores irregularly biseriate, elliptic-cylindric or slightly kidney-
shaped, 9 - 1 3 x 4 - 5 / / , hyaline, non-septate; paraphyses usually absent.
On dead stems of Calluna vulgaris, apparently rare.

209
Yon Arx and Miiller consider the same species occurs also on Betula,
Ledum, Rhododendron, Salix and Vaccinium in which case the legitimate
name for it is P. ledi (Albertini & Schweinitz ex Persoon) Karsten, op. cit.:
157 (1885).

CRYPTOMYCINA von Hohnel

Cryptomycina von Hohnel in Annales mycologici 15:322 (1917).


Apothecia immersed, subepidermal, elongated, black, tearing open
irregularly, asci 8-spored, ascospores hyaline, non-septate.

Cryptomycina ptcridis (Rebentisch ex Fries) von Hohnel, op. cit. 15:


xi (1917). (Fig. 18D.)
Apothecia elongated, subepidermal, h y m e n i u m developing within a
black stromatic tissue, exposed b y the irregular r u p t u r e of the covering
layer on dead overwintered fronds, up to 3 m m . long b y about 0.5 m m .
wide. Asci clavate, up to 6 5 x 1 5 / / , 8-spored; ascospores irregularly bi-
seriate, ellipsoid, 8 - 1 0 x 5 - 6 / « , hyaline, non-septate; a few thread-like
hyphae occur among the asci.
On the under surface of Pteridium aquilinum fronds, developing in late
summer and autumn but not maturing ascospores until May to June in the
following year. Rare. A useful account of the life history of this interesting
parasite will be found in Mycologia 32:214-250 (1940).

PHACIDIUM Fries

Phacidium Fries, Systema mycologicum 2:571 (1823).


Apothecia immersed, circular, opening b y a n u m b e r of teeth, asci
8-spored, pore blued b y iodine, ascospores hyaline, non-septate, more
or less elliptical.

Phacidium lacerum Fries ex Fries, op. cit.:575 (1823). (Fig. 28G.)


Apothecia subepidermal, circular, opening b y from four to six blunt
t e e t h formed b y the radial splitting of the overlying epidermis, disc
grey, up to 1 m m . across, flat. Asci clavate, apex somewhat conical,
70-90 x 8 - 9 n , 8-spored, t h e small pore blued b y iodine; ascospores ir-
regularly biseriate, elliptic-fusiform, 9 - 1 3 x 3 - 4 / / , hyaline, non-sep-
t a t e ; paraphyses simple, slender, slightly swollen towards the tip.

On fallen needles of Pinus sylvestris. Not common.

210
Phacidiurn abietinum Kunze & Schmidt ex Fries, Systema myco-
logicum 2:576 (1823), on needles of Abies differs in its shorter asci,
40-60/jl long; P. infestans Karsten in Medd. Soc. Flora F a u n a Fennica
14:87 (1888), on needles of Pinus sylvestris has asci 130-160/,« long
with ascospores 12-28x4-8.5/«.

PHACIDIOSTROMA von Hohnel

Phacidiostroma von Hohnel in Annales mycologici 15:324 (1917).


Differs from Phacidiurn in possessing a stroma occupying the entire
thickness of the leaf.

Phacidiostroma multivalvc (de Candolle ex St. Amans) von Hohnel,


op. cit.:325 (1917). (Fig. 28II.)
Stroma circular, black, embedded in leaf tissue, opening b y four to
six broad teeth to expose the light, grey-brown disc, 1 - 2 m m . across,
visible as a grey-black ring round t h e base of the t e e t h in the opened
fructification. Asci cylindric-clavate, up to 115xl0/<, somewhat
conical above, the small pore blued b y iodine, 8-spored; ascospores
elliptic-fusiform or slightly clavate, irregularly biseriate, 8 - 1 1 x 3 - 4 f i ,
hyaline, non-septate; paraphyses slender, sometimes distinctly swollen
to 3/t thick near the tip.

On dead leaves of Ilex aquifolium, in spring. Fairly common. The conidial


state is Ceuthospora phacidioides Greville.
As Phacidiurn is regarded as developing its apothecia within a stromatic
tissue most authors regard Phacidiostroma as a redundant genus and call
the above species Phacidiurn multivalve Fries, Systema mycologicum 2:576
(1823).

RHABDOCLINE Sydow

Rhabdocline Sydow in Annales mycologici 20:194 (1922).


Apothecia elongated, subepidermal, without a stroma, erumpent b y
a lid, asci 8-spored, pore not blued b y iodine, ascospores ellipsoid,
hyaline, non-septate.

Rhabdocline pseudotsugae Sydow, op. cit.: 194 (1922). (Fig. 28c.)


Apothecia arranged in rows on each side of the midrib on t h e under
surface of needles, up to 3 m m . long b y about 0.3 m m . wide, opening
b y t h e overlying epidermis folding back in one piece, disc brown,

211
slightly convex. Asci clavate, up to 130x20/«, 8-spored; ascospores ir-
regularly biseriate elliptic-cylindric or often somewhat swollen at each
end, 1 5 - 2 0 x 6 - 9 / t , hyaline, non-septate; paraphyses slender, often
forked. After expulsion from the ascus the ascospores become 1-septate
and one of the two cells usually t u r n s brown, with a thickened wall.

On living needles of Pseudotsuga taxifolia, May and June. Fairly common.

SCHIZOTHYRIOMA von Hohnel

Schizothyrioma von Hohnel in Annales mycologici 15:297 (1917).


Apothecia developed between epidermis and cuticle of living leaves,
minute, black, elliptical, opening irregularly, asci 2-spored, ascospores
hyaline, 1-septate. There is only one British species:

Schizothyrioma ptarmicae (Desmazieres) von Hohnel, op. cit. 15:


xxviii (1917). (Fig. 22c.)
Apothecia scattered, circular to elliptical, up to 1 m m . long, usually
less t h a n 0.5 m m . wide, black, smooth, covering layer splitting ir-
regularly to expose the yellowish-brown disc. Asci clavate, subsessile,
up to 50x8,u, 2-spored; ascospores elliptical, hyaline 1-septate,
1 2 - 1 4 x 5 - 6 / / ; paraphyses cylindrical, slightly enlarged to 3fi wide at
the tip.
On living leaves of Achillea ptarmica, July to September. Uncommon.

SPHAEROPEZIA Saccardo

Sphaeropezia Saccardo in Bot. Centralb. 18:253 (1884).


Apothecia minute, circular to elliptical, on leaves, dark coloured,
at first covered, t h e n with a t o r n central m o u t h , asci 8-spored, thin-
walled, ascospores hyaline 1-3-septate. According to Miiller & von Arx
in P h y t o p a t h . Zeitschr. 24:360 (1955), t h e t y p e species of Sphaero-
pezia, S. alpina (Sacc.) Sacc., is referable to Phacidium so t h a t in their
view the genus is r e d u n d a n t .

Sphaeropezia empetri (Fuckel) Rehm in Rabenhorst's K r y p t o g a m e n


Flora Deutschland 1 ( 3 ) : 7 3 ( 1 8 8 8 ) . ( P L X V K . )
Apothecia scattered, usually several on a leaf, circular, up to 0.4
m m . across, with a dark brown covering layer which splits into three
or four lobes from a central pore. Asci cylindric-clavate, short-stalked,

212
up to about 8 5 x 1 6 / / , 8-spored; ascospores biseriate, elliptic-cylindric,
15-18 x 5 - 6 / t , 3-septate, hyaline; paraphyses numerous, slightly
swollen and brownish at the tip.
On dead leaves of Empetrum still hanging on the stems. Very common
but not to be confused with Phaeangellina empetri, p. 116. According to von
Hohnel in Annales mycologici 15:310 (1917), this is a good Sphaeropezia but
S. vaccinii (Rehm) Rehm, op. cit.: 74 (1888), on leaves of Vaccinium was
made the type of a separate genus, Eupropolella von Hohnel.

LAQUEARIA Fries
Laquearia Fries, S u m m a Veg. Scand. Sect. P o s t . : 3 6 6 (1849).
There is a single species:

Laquearia sphaeralis (Fries) Fries, op. cit.:366 (1849). (Fig. 10c.)


Apothecia immersed in wood, erumpent t h r o u g h the bark, at first
closed b y a blackish tissue composed of more or less rounded purplish-
brown cells up to 10// across, opening b y a pore, t h e n expanded until
the hymenium is fully exposed, disc less t h a n 0.5 mm. across, flat,
brown, with a white pruinose margin. Asci cylindric-clavate, up to
65 x 8 / / , 8-spored, the pore not blued by iodine; ascospores irregularly
biseriate, elliptic-cylindric or slightly curved, 6 - 9 x 1 . 5 - 2 / / , hyaline,
non-septate; paraphyses slender, about 1.5/t thick.

On dead branches of Fraxinus. Rare.


This curious little species might well be sought among the Sphaeriales.
I have seen no British collection though it has long figured in lists of British
fungi. It is illustrated here from Fuckel's Rhineland material in the hope
it may be rediscovered and its true affinities disclosed.

Order 7. LECANORALES

Most members of this order are lichenised and only a few non-
lichenised genera not t r u l y indicative of its scope are briefly mentioned
below. For a full account of the lichenised species see A. L. Smith,
'A Monograph of the British Lichens', second edition, 2 volumes, 1918
and 1926.

Family 1 Lecideaceae
Apothecia circular, light or dark coloured, often more or less
gelatinous in texture, asci and hymenial gelatine blued b y iodine,
except in Mniaecia.

213
I. Ascospores permanently hyaline:
A. Apothecia blue-green or white, on leafy liverworts Mniaecia
B. Apothecia bright-coloured; on wood or bark:
1. Asci 8-spored Agyrium (p. 215)
2. Asci polysporous Biatorella (p. 215)
G. Apothecia black or nearly so:
1. On resinous exudations of coniferous bark see
Retinocyclus (p. 112)
2. On decorticated wood of conifers, apothecia elongated
Placographa (p. 216)
3. Parasites of lichens:
a. Ascospores non-septate Nesolechia (p. 216)
b. Ascospores 1-septate Scutula (p. 217)
c. Ascospores 3- or more septate Mycobilimbia (p. 217)
II. Ascospores coloured:
A. Ascospores 1-septate:
1. Apothecia circular:
a. Apothecia superficial, on wood or on lichens
Iiarschia (p. 219)
b. Apothecia immersed in lichen thalli Abrothallus (p. 220)
2. Apothecia elongated Mycomelaspilea (p. 218)
B. Ascospores 2- or more septate Leciographa (p. 218)
See also Celidium p. 423.

MNIAECIA Boudier

Mniaecia Boudier in Bull. Soc. mycol. France 1:114 (1885).


Apothecia minute, growing on Jungermanniales, hemispherical,
smooth, soft, green or white, asci 8-spored, ascospores large, non-sep-
t a t e , hyaline, paraphyses forked, h y m e n i u m not blued by iodine. The
genus is an isolated one, perhaps b e t t e r transferred to a separate
family.

Mniaecia jungermanniae (Fries) Boudier, Icones mycologicae II,


Livraison 8 (May 1906). (PI. XVo.)
Apothecia sessile, up to 2 m m . across, smooth, soft-fleshed, dark
blue-green t h r o u g h o u t , disc slightly convex. Asci clavate, rather thick-
walled, up to 180x20//, apex rounded and greatly thickened with a
central pore, not blued b y iodine, colourless, 8-spored; ascospores uni-
seriate, elliptical to pyriform, 1 6 - 2 0 x 8 - 1 0 / / , filled with numerous
small oil drops, hyaline to greenish; paraphyses slender, forked, ab-
r u p t l y swollen to 6 - 9 / / wide at the tip, with blue-green contents.

On Aplozia, Calypogeia, Diplophyllum, Jungermannia etc., February to


April. Uncommon.

214
AGYRIUM Fries

Agyrium Fries, Systema mycologicum 2:231 (1822).


Apothecia minute, pulvinate, e r u m p e n t , circular or elliptical, w i t h
flesh greatly reduced, bright coloured, asci 8-spored, ascospores
hyaline, non-septate.

Agyrium rufum (Persoon ex Persoon) Fries, op. cit.:232 (1822).


(PI. X V L . )
Apothecia circular or broadly elliptical, e r u m p e n t , up to 1 m m . long,
gelatinous, disc convex without a sterile margin, reddish-brown. Asci
clavate, rather thick-walled, apex broadly rounded, up to 80x12/«,
8-spored; ascospores biseriate, elliptical, 1 0 - 1 5 x 6 - 8 / « , hyaline, non-
septate, often with a large central oil d r o p ; paraphyses forked,
slender, slightly swollen at the tip, the h y m e n i u m is blued b y iodine.

On decorticated wood, old Calluna stems and the like. Not common.

BIATORELLA de Notaris

Biatorella de Notaris in Giorn. bot. ital. 2 (1):192 (1846).


Apothecia sessile, cup-shaped, superficial, light or dark coloured,
soft-fleshed, asci large, filled with innumerable minute, hyaline,
spherical or elliptical spores.

Biatorella resinae (Fries) Mudd, Manual of British Lichens: 191 (1861).


(PL XVI.)
Apothecia gregarious, sessile, cup-shaped, smooth, disc flat or
slightly convex with a low margin, up to 1.5 m m . across, orange
t h r o u g h o u t . Asci clavate, thick-walled, up to 100x20/«, broadly
rounded above, packed with innumerable globose hyaline ascospores
2 - 3 f i across, slightly yellowish in the mass; paraphyses slender, some-
times forked, swollen at the tips. The asci t u r n deep blue with iodine.

On resinous exudations on the bark on conifers, often associated with the


conidial state, Pycnidiella resinae (Fries) von Hohnel, in S.B. K. Akad. Wiss.
Wien, Math.-Nat. Kl. Abt. 1, 124:91 (1915), and on coniferous stumps, in
winter and spring. Common.
Several other species of Biatorella are regarded as lichens.

Retinocyclus Fuckel in J a h r b . Nass. Vereins N a t u r k . 25/26:332


(1871) was proposed for fungi on resinous exudations of conifers, with

215
apothecia like those of Biatorella in size and shape b u t dark coloured,
having asci packed w i t h small globose secondary spores produced
from septate p r i m a r y ascospores like those of a Tympanis. See
Groves and Wells in Mycologia 48:865-871 (1956).

PLACOGRAPHA Th. Fries

Placographa Th. Fries, Lich. Scand. exs. No. 46 (1860).


Apothecia elongated, black, asci 8- or more spored, ascospores
hyaline, non-septate.

Placographa flexella (Acharius) Th. Fries, Lich. Scand.:637 (1874).


(Fig. 1 1 J . )
Apothecia somewhat immersed, t h e n e r u m p e n t , elliptical to oblong,
up to nearly 1 m m . long, black, with a narrow disc and raised margin.
Asci clavate up to 50x10/«, 8-spored, blued at least in the upper p a r t
by iodine, with a thick-walled a p e x ; ascospores biseriate, elliptical,
6 - 7 x 3 f i , hyaline, non-septate; paraphyses cylindrical, with a r a t h e r
short brown apical cell swollen to 3 or 4fi wide.
On decorticated wood of conifers. Rare.
The species is often classed as a lichen under the name Lithographa flexella
(Acharius) A. Zahlbruckner apud Engler & Prantl, Natiirl. Pflanzenfamil. 1,
abt. 1 *:93 (1903), or is even referred by some authors to Lecidea.

NESOLECHIA Massalongo

Nesolechia Massalongo, Miscell. lichen:43 (1856).


Apothecia sessile, e r u m p e n t , parasitic on lichens, black, sub-
gelatinous, asci 8-spored, ascospores hyaline, non-septate, paraphyses
forked, hymenium blued b y iodine.

Nesolechia oxyspora (Tulasne) Massalongo, op. cit.:43 (1856). (Fig.


LLG.)

Apothecia gregarious, immersed, erumpent, circular, black, pulvi-


nate w i t h a convex disc usually less t h a n 0.5 m m . across. Asci clavate,
thick-walled, up to 6 5 x 2 0 / / , 8-spored; ascospores irregularly arranged,
fusiform, 1 4 - 2 2 x 5 - 7 / t , hyaline, non-septate; paraphyses slender,
swollen to 3 - 5 fi wide at the brown apical cell.

216
On thalli of many species of Parmelia, also of Cetraria glauca and Evernia
furfuracea.

Nesolechia cladoniaria (Nylander) Arnold in Flora 5 7 : 9 9 (1874),


occurs on species of Cladonia.

SCUTULA Tulasne

Scutula Tulasne in Ann. Sci. Nat. Bot. I l l , 17:118 (1852).


Apothecia sessile, erumpent, parasitic on lichens, black, sub-
gelatinous, asci 8-spored, ascospores hyaline, 1-septate, h y m e n i u m
blued b y iodine.

Scutula cpisema (Nylander) Zopf in Hedwigia 35:316 (1896). (Fig. l l r . )


Apothecia usually clustered in groups of from 3 to 14, e r u m p e n t ,
more or less circular or deformed b y m u t u a l pressure, black, about
0.5 m m . across, flat, smooth. Asci clavate, thick-walled, up to 50x 16//,
8-spored; ascospores biseriate, elliptical, 10-14x3.5-4.5/«, hyaline,
1-septate; paraphyses slender, the apical cell a b r u p t l y swollen to 4 - 5 / /
wide, brown.
On thalli of Lecanora calcarea. Also known as Bialorina episema (Nylan-
der) A. L. Smith.

Scutula epiblemastica (Wallroth) Rehm, Rabenhorst's K r y p t o g a m e n


Flora Deutschland 1 (3): 294 (1890), occurs on m a n y species of Pelti-
gera and on Solorina saccata.

MYCOBILIMBIA Rehm

Mycobilimbia Rehm in Rabenhorst's K r y p t o g a m e n Flora E d . 2 , 1 (3):


295 (1890).
Apothecia sessile, circular, light or dark coloured. Asci 8-spored,
ascospores fusiform, multiseptate, colourless. The name applies to
non-lichenised fungi otherwise referable to the lichenised genus
Bilimbia de Notaris in Giorn. Botan. Ital. 2 (1): 190 (1846).

Mycobilimbia killiasii (Hepp) Rehm, op. cit.:327 (1890). (Fig. 19F.)


Apothecia superficial, convex, about 0.75 m m . across, varying from
reddish-yellow to nearly black. Asci clavate, 8-spored, 6 0 - 9 0 x 1 6 - 1 8 / « ;

217
ascospores fusiform, hyaline, 20-40x5-8/<, mostly 3-septate, some-
times 5-septate; paraphyses slender, with swollen yellowish-brown tips
cohering to form an epithecium; h y m e n i u m blued b y iodine.
Scattered over vague grey patches on thalli of Peltigera canina and
P. aphthosa.

MYCOMELASPILEA Reinke
Mycomelaspilea Reinke in J a h r b . f. wissensch. Botan. 28:136 (1895).
Apothecia sessile, superficial, black, elongated, parasitic on lichens,
asci 8-spored, ascospores elliptical, brown, 1-septate. The genus is
distinguished from Abrothallus b y its elongated apothecia and bears
t h e same relationship to the lichenised genus Melaspilea Nvlander in
Act. Soc. Linn. Bordeaux 31:416 (1856), t h a t Karschia does to Buellia.

Mycomelaspilea leciographoides (Vouaux) Keissler, Die Flechtenpara-


siten: 219 (1930). (Fig. 19G.)
Apothecia mostly clustered in small groups, superficial, at first
circular, soon becoming elongated, black. Asci clavate, short-stalked,
8-spored, 6 5 - 8 0 x 2 3 - 2 8 / / ; ascospores biseriate, elliptic-cylindric,
1-septate and constricted at the septum, soon becoming light brown
18-20 x 9 - 1 0 / / ; paraphyses slender, branched, with a b r u p t l y enlarged
brown tips cohering to form an epithecium; hymenium blued b y
iodine.

On sterile thalli of Verrucaria spp. on rocks.

LE CIO GKAPHA Massalongo

Leciographa iMassalongo, Geneac. Lichen: 14 (1854).


Apothecia sessile, e r u m p e n t , parasitic on lichens, black, sub-
gelatinous, asci 8-spored, ascospores becoming brown, 2-7-septate,
paraphyses forked, h y m e n i u m blued b y iodine.

Leciographa parellaria (Nylander) Saccardo & D. Saccardo, Sylloge


F u n g o r u m 18:182 (1906). (Fig. l l n . )
Apothecia scattered or in small groups of two to seven, e r u m p e n t ,
black, smooth, disc flat, up to 0.7 m m . across with a low margin. Asci
clavate, thick-walled, up to 40 x l 5 / « , 8-spored; ascospores irregularly
biseriate, elliptical, light brown, 10-15x3.5-5/«, 1 - 3 - s e p t a t e ; para-
physes slender, with one or two olive-brown apical cells 4 - 6 / / wide.

218
On thalli of Ochrolechia parella, Lecidea platycarpa etc.

Leciographa inspersa (Tulasne) Rehm in Rabenhorst's K r y p t o -


gamen Flora Deutschland 1 (3): 374 (1890), on thalli of Ochrolechia and
Pertusaria, differs only in critical characters.
Lahmia plumbina (Anzi) Keissler in Rabenhorst's K r y p t o g a m e n
Flora Deutschland 8:176 (1930), on thalli of Pannaria plumbea has
hyaline 3-septate ascospores 1 6 - 2 8 x 3 - 4 / / .

KARSCHIA Korber

Karschia Korber, Parerga lichenologica: 459 (1865).


Apothecia sessile, superficial, black or nearly so, subgelatinous, a
few on wood, others parasitic on lichens, asci 8-spored, ascospores
brown, 1-septate, paraphyses forked, h y m e n i u m blued b y iodine;
often united with t h e lichenised genus Buellia de Notaris in Giorn.
bot. ital. 2 (1): 195 (1846).

Karschia lignyota (Fries) Saccardo, Sylloge F u n g o r u m 8 : 7 7 9 (1889).


(Fig. 11c.)
Apothecia scattered, superficial, sessile, shallow cup-shaped, smooth,
black, disc flat or slightly convex, up to 1 m m . across, without a pro-
minent margin. Asci clavate, thick-walled above, up to 40x12/«,
8-spored; ascospores irregularly biseriate, elliptic-clavate, the upper
cell slightly wider t h a n the lower, 9 - 1 2 x 3 - 4 . 5 / / , brown; paraphyses
cylindrical, slightly enlarged towards t h e tip, embedded in yellowish
mucilage, their tips coated b y dark brown amorphous m a t t e r .
On decorticated wood. Common.

The combination in Buellia has been made b y Muller in Beitr.


K r y p t . Flora Schweiz 11 (2): 257 (1962).

Karschia bloxami (Berkeley & Phillips) Saccardo, Sylloge F u n g o r u m 8:


781 (1889). (Fig. 11D.)
Apothecia sessile, superficial, gregarious, black, disc slightly convex,
up to 0.6 m m . across, without a prominent margin. Asci clavate,
thick-walled above, up to 65x12/«, 8-spored; ascospores biseriate
above, elliptical, 1 0 - 1 5 x 5 - 7 / / , brown, 1-septate; paraphyses slender,
their tips coated b y amorphous brown m a t t e r .
On decorticated wood.

219
Karschia advenula (Leighton) Zopf in Hedwigia 35:349 (1896), is
parasitic on thalli of Pertusaria species.
Karschia pulverulenta (Anzi) Korber, Parerga lichenologica: 460
(1865), is parasitic on thalli of Physcia pulverulenta.

Family 2. Patellariaceae

The following genera are alike in having bitunicate asci not blued b y
iodine and are therefore akin to the Dothideales:

I. Ascospores brown, one or more septate:


A. Lichen parasites Abrotliallus
B. Saprophytes on wood and bark Eutryblidiella
II. Ascospores hyaline, multiseptate Patellaria (p. 221)
Eutryblidiella sabinae (de Notaris) von Hohnel in Sitzb. Akad. Wiss.
Wien, Math.-Nat. Kl. 127:564 (1918) occurs on b a r k of Juniperus in
Europe b u t has not yet been found in Britain. See Pirozynski & Reid
in Canad. Journ. Bot. 44:655-662 (1966).

ABROTHALLUS de Notaris

Abrothallus de Notaris in Giorn. bot. ital. 2 (1):192 (1846).


Apothecia sessile, e r u m p e n t , pulvinate, parasitic on lichens, black,
sub-gelatinous, asci 4-8-spored, ascospores becoming brown, 1-septate,
paraphyses forked.

Abrothallus parmeliarum (Sommerfelt) Nylander in Bull. Soc. Linn.


Norm II, 3:12 (1868). (Fig. 11E.)
Apothecia scattered, e r u m p e n t , cushion-shaped with a strongly
convex disc, and no a p p a r e n t margin, black, shining, up to 0.7 m m .
across. Asci clavate, thick-walled, up to 7 0 x 1 6 / / , 8-spored; ascospores
biseriate, somewhat clavate, brown, 1-septate, usually somewhat
constricted at the septum and t h e upper cell a little broader t h a n the
lower, 1 0 - 1 8 x 4 - 6 / / ; paraphyses forked, slightly thickened to 3 - 4 / /
wide at the clavate olive-brown tip. Iodine reaction negative.

On thalli of many species of Parmelia, Cetraria, Sticta, Physcia etc.


Common.

Abrothallus suecicus (Kirschstein) Nordin in Svensk. Bot. Tidskr. 58:


226 (1964), with 3-septate ascospores 1 4 - 2 0 x 5 - 7 / / , occurs on apo-
thecia of Ramalina spp.

220
PATELLARIA Fries
Patellaria Fries, Systema mycologicum 2:158 (1822).
Apothecia sessile, superficial, black, subgelatinous, asci thick-walled,
distinctly bitunicate and dehiscing b y shedding the upper portion of
the outer wall, ascospores elongated, hyaline, multiseptate, h y m e n i u m
not blued b y iodine. For a key to species see Butler in Mycologia 32:
798 (1940).

Patellaria atrata Fries, Systema mycologicum 2:160 (1822). (Fig. H i . )


Apothecia superficial, sessile, saucer-shaped, smooth, black, disc flat
or slightly convex with a low margin, up to 1.5 m m . across. Asci
narrowly clavate, thick-walled, especially in the upper p a r t , up to
150 x20/m, 8-spored, not blued b y iodine; ascospores fusiform-clavate,
usually slightly curved, 3 3 - 4 5 x 8 - 1 0 / « hyaline, 7-11-septate; para-
physes slender, branched, swollen to 3 - 5 / i wide at the olive-brown tips.
On decorticated wood. Uncommon.

Order 8. OSTROPALES

There is a single family, Ostropaceae, with the following genera:

I. Apothecia superficial:
A. Apothecia circular, disc fully exposed:
1. Apothecia stalked Vibrissea
2. Apothecia sessile Apostemidium (p. 222)
B. Apothecia laterally compressed, opening by a pore
Acrospermum (p. 223)
II. Apothecia immersed in plant tissue:
A. Disc fully exposed at maturity:
1. Paraphyses unbranched or only slightly forked at the tip
Stictis (p. 223)
2. Paraphyses highly branched near the tip Schizoxylon (p. 224)
B. Disc exposed by a slit or pore:
1. Disc exposed by a central slit Ostropa (p. 225)
2. Disc exposed by a lateral pore . . . . Robergea (p. 225)

VIBRISSEA Fries

Vibrissea Fries, Systema mycologicum 2:31 (1822).


Apothecia small, distinctly stalked, r a t h e r tough, disc usually
convex and light coloured, covered in t h e fresh state with partially

221
extruded bundles of ascospores like fine white downy hairs, asci v e r y
long and cylindrical, ascospores thread-like, hyaline, multiseptate.
N a n n f e l d t placed this and Apostemidium with the Ostropales because
of their long cylindrical asci and thread-like ascospores b u t the asci
are not capitate like those of Stictis and other authors refer these two
genera to the Geoglossaceae.

Vibrissea truncorum Fries, Systema mycologicum 2:31 (1822). (PI.


XXIIA.)

Apothecia scattered or in small clusters, disc convex, yellowish


orange, u p to 5 m m . across, margin often strongly curved under, stalk
up to 12 m m . long, tapering downwards, pubescent with short, downy
blackish hairs. Asci cylindrical, a b o u t 300x7-9/«, 8-spored; ascospores
thread-like, lying parallel in the ascus, about 200x1.5,w, colourless,
becoming m u l t i s e p t a t e ; paraphyses slender, simple or forked near
t h e tip.

On fallen twigs or small roots and branches lying submerged in streams.


May. Not uncommon.

APOSTEMIDIUM Karsten
Apostemidium Karsten in Not. Sallsk. F a u n a Flora Fennica 11:243
(1870).

Apothecia resembling those of Vibrissea b u t sessile on a broad base,


not always brightly coloured. For a key see Graddon in Trans. Brit,
mycol. Soc. 48:639-646 (1965).

Apostemidium leptospora (Berkeley & Broome) Boudier, Hist. Glass.


Discom. d ' E u r o p e : 9 1 ( 1 9 0 7 ) . ( P I . X X I I B . )
Apothecia scattered, saucer-shaped, sessile on a broad base, disc
convex, up to 2 m m . across, pale yellow to orange, outer surface
smooth, black. Asci cylindrical a b o u t 3 0 0 - 3 3 0 x 6 / / , 8-spored; asco-
spores thread-like, lying parallel in the ascus, a b o u t 2 5 0 - 3 0 0 x 2 - 2 . 5 / / ,
not distinctly septate b u t breaking into two about the middle after
being discharged into w a t e r ; paraphyses slender, simple or branched
near the tip and there septate, slightly swollen, with yellowish con-
tents.
On submerged twigs, in spring. Probably not uncommon but much con-
fused with other species. The discs of apothecia freshly drawn from a stream
become covered with white partially extruded bundles of ascospores like
those of Vibrissea.

222
A. guernisaci (Crouan) Boudier op. cit. 90 (1907) differs in its shorter
asci 160-190/t long, with spores only 100-145 n long.
A. postemidium fiscellum (Karsten) Karsten in Not. Sallsk. F a u n a
Flora Fennica 11:243 (1870), is a similar fungus with light greyish disc,
asci 1 5 0 - 2 0 0 x 7 / / and ascospores almost as long, found on fallen Salix
twigs in d a m p woods b u t not submerged.

ACROSPERMUM Tode ex Fries,

Acrospcrmum Tode ex Fries, Systema Mycologicum 2:244 (1822).


Apothecium club-shaped, short-stalked, laterally compressed, open-
ing b y an apical pore, dark coloured, asci cylindrical, very long and
narrow, bitunicate, ascospores thread-like. Referred to Acrosperma-
taceae in Loculo ascomycetes b y Eriksson (Act. Univ. Upsal. 88,1967).

Acrospcrmum compressum Tode ex Fries, op. cit.: 245 (1822). (Fig.


10A.)

Apothecia scattered, club-shaped w i t h a conical tip, narrowed below


to a short cylindrical stalk, t h e upper fertile portion laterally com-
pressed, dark brown to almost black t h r o u g h o u t , up to 3 m m . high.
Asci narrowly cylindrical, 200-500x4-6/«, 8-spored (Fig. 10, left
margin); ascospores thread-like, non-septate, 1 0 0 - 4 0 0 x 1 / / , colourless;
paraphyses very slender, unbranched, colourless, longer t h a n the asci.
On dead herbaceous stems, especially Urtica, March to June. Uncommon.
A similar fungus on grass culms is called Acrospermum graminum Libert.

STICTIS Persoon ex S. F. Gray

Stictis Persoon ex S. F. Gray, Natural Arrangement of British Plants 1 :


663 (1821).
Apothecia immersed in plant tissue, t h e n erumpent and opening to
expose a circular disc surrounded b y a broad white sterile margin
which m a y be split into lobes, asci elongated, ascospores long and
slender b u t less thread-like t h a n in t h e other genera, multiseptate,
paraphyses slender, simple or slightly forked at the tip only. W h e n
dry the hymenium separates from the margin.

Stictis stcllata Wallroth, Flora Cryptogamica Germaniae 2:444 (1833).


(PL X X I I c and Fig. 1E.)
Apothecia usually occurring in swarms, deeply immersed in woody
tissue, about 0.5 m m . across, with a soft yellow disc surrounded b y a

223
r a t h e r broad snow-white border which is usually split into a n u m b e r
of obtuse lobes. Asci narrowly cylindrical, up to 2 5 0 x 7 / / , apex rounded
and thick-walled with a narrow central pore, 8-spored; ascospores
lying parallel in the ascus b u t often somewhat spirally arranged,
narrowly cylindrical, 1 5 0 - 2 0 0 x 1 . 5 - 2 / / , multiseptate, with individual
cells 4 - 5 / / long; paraphyses slender, simple or slightly lobed at the
tip. The hymenium is not blued b y iodine. The snow-white colour of
the marginal tissue is due to a large n u m b e r of crystal masses
embedded amongst slender colourless hyphae.

On dead herbaceous stems, especially of Epilobium hirsulum, Eupatorium


and Oenanthe; May to September. Fairly common.

Stictis radiata Persoon ex S. F. Gray, op. cit.:663 (1821), is a very


similar fungus on woody twigs, distinguished b y its broader asco-
spores, 2-2.5 u wide, and b y t h e upper portion of the hymenium stain-
ing deep blue with iodine.

Stictis fimbriata Schweinitz in Trans. Amer. Phil. Soc. Philadelphia


N.S. 4 : 1 7 9 (1834). (Fig. 10F.)
Apothecia scattered, immersed in woody tissue, erumpent, a b o u t
0.5 m m . across, disc whitish, surrounded b y a broad snow-white
border. Asci cylindrical, up to 1 0 5 x 9 / / , apex conical and not thick-
walled, 8-spored; ascospores narrowly fusiform, somewhat curved,
5 0 - 6 0 x 1 . 5 - 2 . 5 / / , septate; paraphyses slender, simple.

On scales of fallen cones of Pinus silvestris. Uncommon.


A separate genus, Lasiosliclus Saccardo in Atti R. Inst. Venet. Sci. VI, 6,
2:475 (1884), said to differ from Stictis in having a conspicuously hairy
margin, has been founded on this species. This seems to have been due to an
error of observation and Lasiostictis has been united with Stictis both by
Rehm and by Nannfeldt. It may possibly be tenable on the grounds of the
much shorter ascospores and unthickened tip to the ascus, suggestive of the
Phacidiales. See Petrak in Sydowia 1:89-93 (1947).

SCHIZOXYLON Persoon ex Chevalier

Schizoxylon Persoon ex Chevalier, Flore Generale Environs de Paris 1 :


544 (1826), emend Tulasne, Selecta F u n g o r u m Carpologia 3 : 1 4 8
(1865).
Apothecia erumpent, becoming superficial, sessile, disc dark coloured
with a broad sterile border, asci cylindrical, ascospores thread-like,
multiseptate, paraphyses v e r y slender, m u c h branched at the tip.

224
Schizoxylon berkeleyanum (Durieu & Leveille) Fuckel in J a h r b . Nass.
Vereins f. N a t u r k u n d e 2 3 - 2 4 : 2 5 7 (1870). (PI. X X I I D . )
Apothecia scattered, erumpent becoming superficial, at first hemi-
spherical, then saucer-shaped on a broad base, up to 1 m m . across, disc
flat, blackish-olive surrounded b y a broad, light yellow, powdery
margin composed of very slender h y p h a e with innumerable irregular
yellow inorganic granules interspersed amongst t h e m . Asci narrowly
cylindrical, up to 2 8 0 x 8 / / , 8-spored, apex thick-walled; ascospores
thread-like, 1 4 0 x 2 5 0 x 1 . 5 - 2 . 5 / / , multiseptate, the individual cells
5 - 8 / / long; paraphyses slender, threadlike, m u c h branched at t h e tip
and there m a t t e d to form an epithecial layer which t u r n s blue w i t h
iodine.
On dead herbaceous stems, Epilobium etc. Rare, apparently more frequent
in the last century.

OSTROPA Fries

Ostropa Fries, Systema Orbis Vegetabilis 1 : 1 0 9 (1825).


Apothecia immersed in bark, dark coloured, e r u m p e n t b y a broadly
rounded apex which opens b y a transverse slit, asci cylindrical, asco-
spores thread-like.

Ostropa barbara (Fries) Nannfeldt in Nova Acta Reg. Soc. Sci. Upsal. IV,
8 : 7 8 (1932). (Fig. 10B.)
Apothecia gregarious, subglobose or pear-shaped, immersed in b a r k
b u t with a broad p r o t u b e r e n t apical papilla, dark grey or blackish
with a whitish bloom, about 1 m m . across, opening b y a transverse
slit to expose the yellowish disc. Asci narrowly cylindrical, up to
250x10//, 8-spored, apex thick-walled w i t h a central pore; ascospores
thread-like, 180-200x1.5//, multiseptate, lying parallel in the ascus;
paraphyses slender, branched at the tip.

On dead twigs of shrubs and small trees. Very rare in the British Isles.
Also known as Ostropa cinerea (Persoon) Fries, op. cit.: 109 (1825).

ROBERGEA Desmazieres

Robergea Desmazieres in Ann. Sci. Nat. Bot. I l l , 8:177 (1847).


Apothecia completely immersed in woody tissue, flask-shaped,
having their long axis parallel with t h a t of the twig, ending in a curved
neck which opens b y a pore, asci cylindrical, ascospores thread-like.

225
Robergea cubicularis (Fries) Rehm in Ber. Bayer. Bot. Ges. 13:163
(1912). (Fig. 10E.)
Apothecia immersed in woody tissue, flask-shaped, up to 2.5 m m .
long, lying almost parallel to t h e surface, with a short curved neck
arising at one end and p e n e t r a t i n g the b a r k to open b y a round pore
with a narrow whitish margin. Asci cylindrical, up to 8 0 0 x 1 0 / / ,
8-spored, the apex thick-walled with a central pore; ascospores thread-
like, lying parallel in t h e ascus, 3 0 0 - 6 0 0 x 1 . 5 - 2 / / , multiseptate; para-
physes very slender.

On dead twigs of shrubs and small trees in autumn. Apparently rare but
inconspicuous and easily overlooked.
Also known as Robergea unica Desmazieres, op. cit.: 178 (1847).

Order 9. CLAVICIPITALES

The British species can be assigned to seven genera, distinguished


from those of the Ostropales b y the asci being contained in small flask-
shaped perithecia with light or bright coloured walls.

I. Perithecia gregarious, more or less immersed in a common stroma:


A. Stroma stalked, arising from a sclerotium which originated in floral
structures of Monocotyledons Clayiceps
B. Stroma stalked, arising from the body of a mummified insect or
from a fructitificafion of an Elaphomyees:
1. Ascospores breaking up into their individual cells
Cordyceps (p. 227)
2. Ascospores not fragmenting Ophiocordyceps (p. 230)
C. Stroma sessile:
1. Stroma a hollow cylinder surrounding living grass stems
Epichloe (p. 231)
2. Stroma small, conoid, scattered over grass leaves
Oomyces (p. 232)
II. Perithecia gregarious, seated on a subiculum:
A. Growing on Claviceps purpurea Barya (p. 231)
B. Growing on spiders Torrubiella (p. 230)

CLAYICEPS Tulasne

Claviceps Tulasne in C.R. Acad. Sci. Paris 33:646 (1851).


S t r o m a t a borne on an elongated black sclerotium, club-shaped with
globose heads, perithecia completely sunk in the head, asci cylindrical,
ascospores thread-like.

226
Claviceps purpurea (Fries) Tulasne in Ann. Sci. Nat. Bot. I l l , 20:45
(1853). (PI. X X V I J . )
Sclerotia elongated, more or less cylindrical with rounded ends b u t
usually with slight longitudinal grooves and ridges, formed beneath
the ovary of a grass flower and growing out to replace it, varying
greatly in size and shape according to the species of grass infected,
black externally with hard white flesh. S t r o m a t a arising singly or in
small clusters on t h e fallen sclerotia, stalk slender, smooth, cylindrical,
pale purple, head globose, commonly about 2 m m . across, ground colour
from cream to purple, dotted with the darker slightly p r o t r u d i n g
ostioles of the perithecia. Asci narrowly cylindrical, very long, a b o u t
5// wide, apex thick-walled with a central pore, 8-spored; ascospores
thread-like, lying parallel in the ascus, about 100x1//, becoming
septate a f t e r expulsion.

Sclerotia ("ergots") in inflorescences of Agropyron, Alopscurus, Ammo-


phila, Anthoxanthum, Arrhenatherum, Brachypodium, Bromus, Dactylis,
Deschampsia, Elymus, Festuca, Glyceria, Holcus, Hordeum, Loliutn, Molinia,
Nardus, Phalaris, Phleum, Phragmites, Poa, Secale and rarely Triticum\
stromata on overwintered sclerotia from April to June.

Claviceps nigricans Tulasne in Ann. Sci. Nat. Bot. I l l , 20:51 (1853),


occurs on Eleocharis, see Trans. Norfolk, Norwich Nat. 18 (3): 18 (1956).
See f u r t h e r two short papers on Claviceps b y Petch in The Naturalist
1935: 151-155 and 1937:25-28.

CORDYCEPS (Fries) Link


Cordyceps (Fries) Link, H a n d b u c h zur E r k e n n u n g der . . . Gewachse
3:347 (1833).
S t r o m a t a stalked, fertile head club-shaped, ovoid or subglobose,
fleshy, perithecia more or less immersed, asci narrowly cylindrical,
ascospores thread-like, breaking up into short rod-like part-spores
while still within the ascus. The species are all parasitic, either on in-
sects or on the subterranean fructifications of Elaphomyces. In collect-
ing a Cordyceps care m u s t be t a k e n to dig down sufficiently to discover
from w h a t organism the fructifications are springing.

Cordyceps militaris (Linnaeus ex St. Amans) Link, op. cit.:347 (1833).


(PI. XXVIA.)
S t r o m a t a usually solitary, up to 5 cm. high, with a slender flexuous
stalk which passes imperceptibly into the cylindric-fusiform fertile

227
head, the latter about 5 m m . thick and minutely roughened with the
perithecial ostioles, red or orange-red t h r o u g h o u t . Perithecia com-
pletely immersed, asci very long, a b o u t 4// wide, the apex thick-
walled, 8-spored; ascospores thread-like, as long as the ascus, breaking
up into part-spores which become slightly barrel-shaped, 3 . 5 - 6 x
I-1.5//.
On larvae and pupae of Lepidoptera, usually buried in soil, September to
November. Common, especially in pastures.
There is no indication that the common white Isaria found on the same
hosts is genetically connected with C. militaris; the conidial state of the
latter has been shown to be a much less conspicuous mould belonging to the
form genus Cephalosporium.
Petch points out that the part-spores of CorcLyceps are best obtained by
keeping a fresh mature specimen on a glass slide in a chamber overnight,
when it will deposit a spore print. Measurements made on undischarged
ascospores are apt to be misleading.

Cordyceps gracilis Montagne & Durieu, Exploration Scientifique de


L'Algerie, Sci. Nat. Bot. 1:449 (1869). (PI. X X V I B . )
S t r o m a t a usually solitary, with a long, straight, cylindrical, smooth
stalk which expands a b r u p t l y into an ovoid fertile head about 7 x 5
m m . , total height above ground up to 4 cm., stalk yellowish, head pale
chestnut-brown, smooth b u t finely dotted with the darker ostioles.
Perithecia completely immersed, asci very long, 4 - 5 / t wide, apex
thick-walled; ascospores threadlike, part-spores cylindrical, 5 - 9 x
1.5-2 /t.
On buried larvae of Lepidoptera, April to June. Not uncommon.

Cordyceps forquignoni Quelet in C.R. Assoc. Frang. Avanc. Sci. 16th


Sess.: 592 (1887). (PI. X X V I c . )
S t r o m a t a usually solitary, up to 3.5 cm. high, with a slender stalk,
smooth pale ochraceous, a b o u t 0.5 m m . thick, crowned by a sub-
globose orange-brown head 1 - 2 m m . across, finely dotted with the
perithecial ostioles. Perithecia completely immersed, asci narrowly
cylindrical, about 5// wide; ascospores thread-like, part-spores
I I - 1 2 x 2 fi.
On flies, May to August. Not uncommon, especially on the ground in
woods, but very inconspicuous.

Cordyceps sphecocephala (Klotzsch ex Berkeley) Berkeley & Curtis in


J o u r n . Linn. Soc. B o t . : 3 7 6 ( 1 8 6 8 ) . ( P I . X X V I F . )
S t r o m a t a usually solitary, up to 6 cm. high, with a slender, flexu-
ous, smooth, whitish stalk and short club-shaped head, about 5 X 2 - 3

228
mm., yellow, marked with a network of furrows around the perithecia
and minutely dotted with their darker yellow ostioles. Perithecia
immersed but lying at an angle to the surface of the head and only
partially overlapping one a n o t h e r ; asci cylindrical, about 2 5 0 x 8 / / ,
8-spored; part-spores narrowly elliptical, 8 - 1 5 x 1 . 5 - 2 . 5 / / .

On Ilymenoptera in autumn. Rare.

Cordyceps tuberculata (Libert) Maire in Bull. Soc. Hist. Nat. Afrique


du Nord 8 : 1 6 5 ( 1 9 1 7 ) . ( P I . X X V I G . )
S t r o m a t a gregarious, up to 1.5 cm. high, stalk slender, whitish, head
narrowly club-shaped, whitish b u t dotted with the yellow p r o t r u d i n g
ostioles of the more or less immersed perithecia, 1 - 2 m m . thick. Asci
narrowly cylindrical, about 5/t thick, 8-spored; ascospores thread-like,
part-spores 4 - 8 X 1 / / .
On pupae of Lepidoptera, sometimes also on the mature insect, in spring
and autumn. In woods, uncommon.

Cordyceps ophioglossoides (Ehrenberg ex Fries) Link, op. cit.:347


(1833). (PI. XXVID.)

S t r o m a t a solitary or a few together, up to 10 cm. high, club-shaped


with a rather stout, smooth, yellow stem u p to 3 m m . thick which
gradually enlarges into a cylindric-ovoid fertile head up to 2.5 x 1.3 cm.
which is at first yellow and smooth, becoming black and minutely
roughened by the p r o t r u d i n g ostioles. Perithecia immersed; asci
narrowly cylindrical, 7// wide, 8-spored; ascospores thread-like, p a r t -
spores elliptic-cylindric, 2 . 5 - 5 X 2 / / .

Parasitic on Elaphomyces, on the ground in woods, September to October.


Not uncommon.

Cordyceps capitata (Holmskjold ex Fries) Link, op. cit.:347 (1833).


(PI. X X V I E . )
S t r o m a t a solitary or in small groups, up to 9 cm. high, with a r a t h e r
stout, smooth or furrowed, yellow stalk up to 1 cm. thick below b u t
tapering upwards, a b r u p t l y enlarged into an ovoid or subglobose d a r k
brown or black head which is finely roughened b y the ostioles when
dry. Perithecia immersed; asci cylindrical, a b o u t 15/t wide, 8-spored;
ascospores narrowly cylindrical, part-spores cylindrical, thinwalled,
varying greatly in length, (7-) 1 4 - 2 0 x 2 - 3 / / .

Parasitic on Elaphomyces, September to October. Less common than the


preceding.

229
Cordyceps canadensis Ellis & E v e r h a r t in Bull. Torrey bot. Club 2o:
501 (1898), superficially like the above, has been distinguished b y its
part-spores being more or less fusoid, 2 0 - 5 0 x 3 - 5 ( - 7 . 5 ) / / , with solid
ends.

OPHIOCORDYCEPS Petch

Ophiocordyceps Petch in Trans. Brit, mycol. Soc. 16:73 (1931).


Differs from Cordyceps in the ascospores not breaking up into p a r t -
spores. There is only one British species:

Ophiocordyceps clavulata (Schweinitz) Petch in Trans. Brit, mycol.


Soc. 1 8 : 5 3 (1933). (PL X X V I H . )
S t r o m a t a gregarious, arising from a common subiculum of whitish
mycelium, each about 2 m m . high with a short, slender, whitish or ash-
coloured, cylindrical stalk and an ovoid head, less t h a n a millimetre
across, rough with the black protruding tips of the perithecia. Asci
cylindric-clavate, up to 100x10//, thick-walled at t h e tip, 8-spored;
ascospores cylindric-clavate, 50-80x2/«, 7- or 8-septate.

Parasitic on scale insects in spring and autumn. Very rare in southern


England or perhaps extinct, not collected for nearly a century and figured
here from North American material.

TORRUBIELLA Boudier

Torrubiella Boudier in Revue Mycologique 7:226 (1885).


Differs from Cordyceps in the perithecia being free, though often
gregarious, and seated on a subiculum of m a t t e d hyphae. The British
species are parasitic on spiders and flies.

Torrubiella albolanata Petch in Trans. Brit, mycol. Soc. 27:85 (1944).


(PL X X V I i . )
Perithecia gregarious, almost buried in a dense white mycelial m a t
with only their yellowish ostioles protruding, conoid, 0.5 m m . high by
0.25 m m . wide. Asci narrowly cylindrical, 4 - 5 / / wide, apex thick-
walled ; ascospores lying parallel, often somewhat spirally arranged in
the ascus, thread-like, part-spores cylindrical, 5 - 7 x 1 - 1 . 5 / / .
On small spiders among debris of marsh plants in summer. Probably not
uncommon in suitable localities.

230
Torrubiella aranicida Boudier, op. cit.:227 (1885), has yellowish
perithecia scattered over a sparse whitish subiculum on body and legs
of m u c h larger spiders.
Torrubiella albotomentosa Petch in Trans. Brit, mycol. Soc. 27:86
(1944), occurs on Dipterous pupae, in marshes.

BARYA Fuckel

Barya Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 2 3 - 2 4 : 9 3 (1870).


Differs from Torrubiella in the ascospores not becoming septate or
breaking into part-spores. There is one very rare British species:

Barya aurantiaca Plowright & Wilson in Gardener's Chronicle 21:176


(February 1 8 8 4 ) . (PI. X X V I M . )
Perithecia gregarious on a sparse whitish subiculum, almost super-
ficial, flask-shaped, up to 0.3 m m . high, yellow, drying orange. Asci
cylindrical up to 250x3/n, thick-walled at the tip, 8-spored; ascospores
thread-like, as long as the ascus.
Parasitic on germinating sclerotia of Claviceps purpurea, July. Very rare,
only known from a single collection made on ergots from Glyceria fluitans
in 1882.

EPICHLOE (Fries) Tulasne

Epichloe (Fries) Tulasne, Selecta F u n g o r u m Carpologia 3 : 2 4 (1865).


Perithecia immersed in a stroma surrounding the culms of grasses,
asci cylindrical very long, 8-spored, ascospores thread-like, multi-
septate b u t not breaking up into part-spores. There is a single British
species:

Epichloe typhina (Persoon ex Fries) Tulasne, op. cit.:24 (1865). (PI.


XXVIk.)
Stroma sessile, cylindrical, sheathing the stems of grasses, up to
5 cm. long b u t usually considerably shorter, at first smooth and white,
becoming golden-yellow and minutely roughened with the tips of t h e
perithecia when m a t u r e . Perithecia oval, 0.25 m m . across; asci nar-
rowly cylindrical, 6-8(M wide, 8-spored; ascospores thread-like, nearly
as long as the ascus, 1.5-2/t wide, individual cells 8 - 1 2 fi, long.

On living culms of Agrostis, Bromus, Dactylis, Deschampsia, Festuca, llol-


cus, Koehleria, Phleum and Poa, May to September. Very common. Affected
shoots seldom flower.

231
OOMYCES Berkeley & Broome

Oomyces Berkeley & Broome in Ann. Mag. Nat. Hist. 11,7:185 (1851).
Stroma erumpent b u t becoming almost superficial, conoid, small,
light coloured and soft-fleshed, containing few perithecia, asci nar-
rowly cylindrical, bitunicate, ascospores thread-like, multiseptate. Re-
ferred to Acrospermataceae in Loculo ascomycetes b y Eriksson 1967.
There is a single species:

Oomyces carneo-albus (Libert) Berkeley & B r o o m e , op. cit.: 185 (1851).


(Plate X X V I L . )
S t r o m a t a scattered, e r u m p e n t , conoid, smooth, up to 0.75 m m .
high b y 0.5 m m . wide, broadly rounded above and there p u n c t a t e with
the m o u t h s of the completely immersed perithecia, pale flesh-coloured.
Perithecia three to seven in a stroma, flask-shaped; asci narrowly
cylindrical, 5 - 6 / / wide; ascospores lying parallel in the ascus, t h r e a d -
like, multiseptate, 1.5// wide.

On dead leaves of Deschampsia caespitosa, January to May. Apparently


rare.

Order 10. SPHAERIALES

There is as yet no general agreement amongst mycologists regarding


the classification of the very large and varied assemblage of species
t h a t m a y be assigned to t h e Sphaeriales. The families adopted below,
therefore, are to be regarded as t e n t a t i v e groupings subject to ex-
tensive revision as t h e species become more adequately known.
I. Perithecia usually brightly coloured, their walls somewhat soft-fleshed,
the ascospores usually hyaline, occasionally greenish or light brown,
often septate, asci persistent:
A. Perithecia immersed in a stroma or the asci 16-spored at maturity
Hypocreaceae (p. 233)
B. Perithecia free though sometimes seated on a stromatic base or on
a subiculum, asci 8-spored Nectriaceae (p. 237)
II. Perithecia usually dark coloured, olive-brown to black, or the ascospores
black or the asci dissolving at maturity. In Selinia the perithecia are
bright orange, in Endothia they are at first red. If they appear dark
blue to purple see Gibberella in Nectriaceae:
A. Asci dissolving at maturity, ascospores emerging in gummy masses
Melanosporaceae (p. 262)
B. Asci persistent:
1. Ascospores 1-septate, hyaline, with a long slender tail, aquatic
species Loramyces (p. 328)
2. Ascospores very large, non-septate, hyaline, perithecia bright
coloured, in dung Seliniaceae (p. 256)

232
3. Ascospores without the above distinctive characters:
a. Ascospores hyaline, non-septate, perithecia immersed
either in a stroma or in plant tissue:
*Ascus pore blued by iodine Amphishaeriaceae
* Ascus pore not blued by iodine Polystigmataceae
b. Ascospores black or dark brown, non-septate:
*Perithecia immersed in a large stroma or beneath a
clypeus or, if free, then usually smooth, not in
dung, ascospores without long hyaline appendages
Xylariaceae (p. 274)
**Perithecia not in a stroma, often hairy or on dung,
ascospores sometimes with cylindrical or tapering
hyaline appendages Lasiosphaeriaceae (p. 266)
c. Ascospores hyaline to yellowish, often septate; or kidney-
shaped:
*Ascospores worm-like, septate or not
lasiosphaeriaceae (p. 266)
**Ascospores not worm-like:
a. Ascus pore or apical ring blued by iodine
Amphisphaeriaceae (p. 287)
Not so:
fPerithecia usually immersed in a cushion-
like stroma within plant tissue or at least
in circular clusters with the ostioles
emerging in a compact group, ascospores
sausage-shaped, non-septate except in
Valsaria . . . Diatrypaceae (p. 296)
tfPerithecia not in a stroma or if a stroma is
present the ascospores are not sausage-
shaped or are septate:
§Perithecia sunk in host tissue or
erumpent, often with a stroma:
.Ascospores hyaline, uniseriate in
cylindrical short-stalked asci
Endodothella (p. 261)
.•/Ascospores biseriate, in long-stalk-
ed asci or ascospores coloured
Diaporthaceae (p. 305)
§§Perithecia immersed singly in wood
or bark, without a stroma, long-
beaked, ascospores hyaline, septate
Ceratostomataceae (p. 326)
§§§Perithecia superficial, without a stro-
ma . . . . Sphaeriaceae (p. 328)

Family 1. Hypocreaceae

Nearly all the members of this family are characterised b y having


narrowly cylindrical asci in which the ascospores each separate into

233
two similar subglobose portions so t h a t the m a t u r e ascus contains a
single row of 16 spores. The exception to this rule is Hypocreopsis.
This is traditionally placed here because of its highly developed stroma
b u t it might find a more n a t u r a l position in t h e next family. Hypo-
creaceae and Nectriaceae were traditionally ranked as an order, Hypo-
creales, and a useful account of the species of b o t h families will be
found in Petch, 'British Hypocreales,' 1938. As here restricted Hypo-
creaceae includes five genera:

1. Ascospores dividing into two and mature asci therefore 16-spored:


A. Stroma erect, club-shaped Podostroma
B. Stroma sessile, cushion-shaped or flattened:
1. Ascospores colourless Hypocrea
2. Ascospores greenish to olive Creopus (p. 235)
C. Stroma reduced to a dense white or cream-coloured subiculum
Protocrea (p. 236)
II. Ascospores 1-septate, not dividing, stroma large and lobed
Hypocreopsis (p. 236)

PODOSTROMA Karsten

Podostroma Karsten in Hedwigia 31:294 (1892).


There is a single British species:

Podostroma alutaccum (Persoon ex Fries) Atkinson in Botanical


Gazette 40:416 (1905). (PI. X X V I I c . )
Stroma erect, club-shaped, up to 4 cm. high, smooth, whitish to pale
yellow throughout, stalk stout, cylindrical, imperceptibly enlarged
into the fertile head which is minutely dotted with the ostioles of the
completely immersed perithecia. Asci narrowly cylindrical, 8 0 - 9 0 x
4//, 16-spored, part-spores globose, to broadly elliptical, 4-4.5x3-4/.;,
hyaline, minutely w a r t e d .
On the ground in coniferous woods or attached to rotting wood, Sep-
tember to October. Rare.

HYPOCREA Fries

Hypocrea Fries, Systema Orbis Vegetabilis 1 : 1 0 4 (1825).


S t r o m a sessile, flat or cushion-shaped, dotted with the ostioles of the
completely immersed perithecia, asci ultimately 16-spored, part-spores
subglobose, hyaline or yellowish.

234
Hypocrea rufa (Persoon ex Fries) Fries, S u m m a Veg. Scand. Sect.
P o s t . : 3 8 3 (1849). (PI. XXVIN.)

Stroma usually convex or cushion-shaped with a broad flat base, up


to 10 m m . across, chestnut-brown with a whitish margin, finally light
reddish-brown all over, smooth, minutely dotted with the ostioles,
outline circular or slightly lobed, flesh firm, whitish. Asci narrowly
cylindrical, about 7 5 x 5 / t , ultimately 16-spored; part-spores globose,
or broadly elliptical, 3 . 5 - 6 x 3 . 5 - 4 . 5 / / , minutely punctate.

On dead wood, bark, old fungus fructifications and other debris, July to
November. Common. The conidial state is the very common blue-green
mould Trichoderma lignorum (Tode ex Fries) Ilarz in Bull. Soc. fmp. Moscou
44:116 (1871).

Hypocrea pulvinata Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e


23-24:185 (1870), is common on old fructications of Polyporus betu-
linus, the common bracket fungus of Betula t r u n k s .
Hypocrea lactea (Fries ex Fries) Fries, S u m m a Veg. Scand. Sect.
P o s t . : 3 8 3 (1849), is a flat yellowish species which forms crusts on
dead wood, soil or debris, see Rifai & Webster in Trans. Brit, mycol.
Soc. 49:303-306 (1966).

CREOPUS Link

Creopus Link, H a n d b . z. E r k e n n t n . d. Gewachse 3 : 3 4 9 (1833).


Distinguished from Hypocrea b y the greenish ascospores. Chromo-
crea Seaver in Mycologia 2 : 5 8 (1910) is a synonym.

Creopus gclatinosus (Tode ex Fries) Link, op. cit.: (1833). (PI. XXVIP.)

Stroma hemispherical or cushion-shaped, up to 3 m m . across,


smooth, light yellow, the upper surface dotted with the ostioles, flesh
translucent when moist so t h a t the perithecial contents appear as
dark dots. Asci narrowly cylindrical, up to 9 5 x 6 / / , 16-spored, p a r t -
spores unequal, t h e upper globose, 4// diameter, t h e lower broadly
elliptical, 5 - 6 x 3 - 4 / / , finely warted, dark green.

On decaying wood, in autumn. Fairly common.

Chromocrea spinulosa (Fuckel) P e t c h in Trans. Brit, mycol. Soc. 33:


350 (1950), is a similar species found on rotting marsh plants.

235
PROTOCREA Petch

Protocrea Petch in J o u r n a l of B o t a n y 75:219 (1937).


Perithecia gregarious, seated on or p a r t l y immersed in a c o t t o n y
hyphal web, asci and ascospores as in Hypocrea. The genus Arachno-
crea Moravec in Bull. Soc. mycol. France 72:161 (1956), has been pro-
posed for similar fungi with fusiform ascospores.

Protocrea farinosa (Berkeley & Broome) Petch, op. cit.:219 (1937).


(PI. X X V I I B . )
Subiculum white, cottony, becoming compact and tomentose or
mealy, wide-spreading, t h i n ; perithecia subglobose, about 0.25 m m .
across, w a t e r y white t h e n yellowish. Asci narrowly cylindrical, up to
9 0 x 4 / / , 16-spored; ascospores dividing into two, part-spores subglo-
bose or the lower one broadly elliptical, 3 - 4 . 5 x 3 - 4 / t , hyaline, minutely
punctate.

On dead wood of deciduous trees, throughout the year. Not uncommon.

Protocrea delicatula (Tulasne) Petch op. cit. (1937) is a similar fungus


of coniferous wood.

HYPOCREOPSIS Karsten

Hypocreopsis Karsten, Mycologia Fennica 2:251 (1873).


Stroma flattened, very large, divided into radiating strap-like lobes,
asci 8-spored, ascospores 1-septate, not dividing into part-spores.
There is a single British species:

Hypocreopsis lichenoides (Tode ex Fries) Seaver in Mycologia 2 : 8 2


(1910). (PI. XXVIIA.)
S t r o m a flattened and conforming approximately with the surface
of the substrate, cut up into strap-like branching lobes radiate from a
common centre, the whole mass up to 10 cm. across, light yellowish-
brown to orange-brown, paler at the margin, minutely dotted with the
ostioles of the immersed perithecia. Asci narrowly cylindrical up to
180x10//, 8-spored; ascospores uniseriate or slightly overlapping,
fusiform, 2 4 - 3 0 x 8 - 9 / / , hyaline, smooth, with a median septum.

On dead branches of Salix\ September to May. Evidently rare as such a


large and conspicuous species could scarcely be overlooked and it has been
very seldom recorded.

236
Family 2. Nectriaceae

The genera of Nectriaceae are to some extent ill-defined and se-


parated b y artificial or inconstant characters. The traditional scheme
is as follows:

I. Ascospores non-septate, hyaline:


A. Perithecia superficial:
1. Perithecia seated on a subiculum Byssonectria (p. 238)
2. Perithecia without a subiculum Pseudonectria (p. 240)
B. Perithecia embedded in the tissue of a Tuber fructification
Battarrina (p. 238)
II. Ascospores 1-septate:
A. Ascospores brown:
1. Asci unitunicate Nectria (p. 240)
2. Asci bitunicate, pseudothecia minute, brown, amongst coni-
diophores of a Dematiaceous mould
see Letendraea (p. 386)
B. Ascospores hyaline:
1. Ascospores pointed at the ends, rough:
a. Cells of the ascospores approximately equal in size
Hypomyces (p. 239)
b. Cells of the ascospores markedly unequal
Apiocrea (p. 239)
2. Ascospores rounded at the ends, usually smooth, occasionally
rough:
a. Perithecia at first immersed in the host
Nectriella (p. 252)
b. Perithecia superficial, sometimes seated on a stromatic
base:
+ Perithecia clustered in small compact groups:
*Perithecia seated on a small stromatic base
Nectria (p. 240)
**Perithecia at the base of a club-shaped coni-
dial body . . . . Sphaerostilbe (p. 250)
+ + Perithecia scattered:
^Perithecia seated on a subiculum
Nectriopsis (p. 251)
**Perithecia without a subiculum:
jPerithecia smooth or bearing short
downy white hairs Nectria (p.240)
ttPerithecia crowned by a fringe of tri-
angular teeth Neohenningsia (p.251)

III. Ascospores with two or more transverse septa, elliptical to oblong:


A. Perithecia crowned with spreading teeth Actiniopsis (p. 253)
B. Perithecia without teeth, surface smooth:
1. Ascospores drawn out into a cilium at each end
Paranectria (p. 254)

237
V
2. Ascospores without cilia: V"
a. Perithecial walls reddish or yellowish .._.- /
(<aIonectria (p. 252) A
b. Perithecial walls dark blue, purplisn-bfack or nearly
black Gibberella (p. 253)
c. Perithecial walls brown, immersed in the host
see "Gripliosphaeria" nivalis (p. 290)
IV. Ascospores worm-shaped with many transverse septa
A. Perithecia smooth Scoleconectria (p. 254)
B. Perithecia hairy Trichonectria (p. 255) v<2 .ft
C. Perithecia with a small crown of short processes ^
(See Rubeufia, p. 421) wXO
V. Ascospores with both transverse and longitudinal septa
Thyronectria (p. 255)

The genus Battarrina (Saccardo) Clements & Shear, Genera fo Fungi: 279
(1931), is represented in Britain by B. inclusa (Berkeley & Broome) Clements
& Shear, op. cit. (1931), the perithecia of which occur completely embedded
in the flesh of Tuber puberulum and will not be further described of figured
here.

BYSSONECTRIA Karsten

Byssonectria Karsten in Medd. Soc. F a u n a Flora Fennica 6 : 6 (1881).


Perithecia small, subglobose to pear-shaped, smooth, bright co-
loured, seated on a common dense h y p h a l web, asci 8-spored, asco-
spores fusiform, non-septate.

Byssonectria viridis (Albertini & Schweinitz ex Berkeley & Broome)


Petch in Journal of B o t a n y 75:220 (1937). (PI. X X X I J . )
Perithecia ovoid, smooth, about 0.3 m m . across, pallid, darkening to
nearly black, gregarious on a dense, thin, h y p h a l web which is at first
egg yellow, becoming greenish. Asci cylindrical, about 1 6 0 x 8 / / ,
8-spored; ascospores uniseriate, narrowly fusiform, with solid pointed
tips which are often slightly hooked, finely warted, 2 7 - 4 5 x 5 - 6 / / ,
hyaline.

On various decaying agarics, especially species of Lactarius, July to Sep-


tember. Uncommon.

Byssonectria lateritia (Fries) Petch in J o u r n a l of B o t a n y 75:220


(1937), also occurs on Lactarii b u t has a white to light brown subiculum
and much smaller ascospores, 1 5 - 2 5 x 4 - 5 / / .

238
HYPOMYCES Tulasne

Hypomyces Tulasne in Ann. Sci. Nat. Bot. IV, 13:11 (1860).


Perithecia gregarious on a common h y p h a l web, asci usually
8-spored, ascospores fusiform, pointed at each end, with a median
septum.

Hypomyces aurantius (Persoon ex Fries) Tulasne in Ann. Sci. N a t . Bot.


IV, 13:12 (1860). (PI. X X X I B . )
Perithecia densely crowded, globose to slightly pear-shaped,
0.3 m m . across, smooth, golden-yellow to orange-red, seated on a dense
h y p h a l web which is usually ochraceous b u t whitish at the margin.
Asci cylindrical, up to 1 4 0 x 6 / / ; ascospores uniseriate, fusiform,
pointed at each end, often slightly curved, finely warted, hyaline,
18-27 x 4 - 6 f i , with a single transverse septum at or near the middle.

On old fructifications of species of Polyporus, Polystictus, Panus etc.,


throughout the year. Fairly common.

Hypomyces ochraceus (Persoon) Tulasne, Selecta F u n g o r u m Carpo-


logia 3 : 4 1 ( 1 8 6 5 ) . (PI. X X X I i . )
Perithecia pear-shaped, 0.3 m m . across, smooth, yellow to reddish,
scattered over a dense white or ochraceous h y p h a l web and often half
buried in it. Asci cylindrical, up to 200x 7 / t ; ascospores uniseriate,
cylindric-fusiform with solid pointed tips, often slightly constricted at
the single median septum, finely warted, 2 7 - 4 0 x 6 - 7 / / , hyaline.

On the ground in woods where infected species of Russula and Lactarius


have disintegrated, often associated with the tiny white brush-like conidial
state, Verticillium agaricinum Corda; September to October. Not common.

Hypomyces rosellus (Albertini & Schweinitz ex Fries) Tulasne in


Ann. Sci. Nat. Bot. IV, 13:12 (1860), is not uncommon on decaying
Stereum hirsutum.
Hypomyces broomeanus Tulasne, Selecta F u n g o r u m Carpologia 3:
108 (1865), occurs on Fomes annosus and H. asterophorus Tulasne in
Ann. Sci. Nat, Bot. IV, 13:14 (1860), on Nyctalis.

APIOCREA Sydow

Apiocrea Sydow in Annales mycologici 18:186 (1920).


Separated from Hypomyces only b y t h e unequal-celled ascospores.

239
Apiocrea chrysosperma (Tulasne) Sydow, op. cit.: 187 (1920). (PL
XXXIA.)

Perithecia gregarious, subglobose to pear-shaped, about 0.4 m m .


across, smooth, at first hyaline, t h e n orange-yellow, seated on a golden-
yellow hyphal m a t . Asci cylindric-clavate, up to 2 0 0 x 1 0 / / ; ascospores
fusiform, with solid pointed tips, finely warted, 2 1 - 3 0 x 5 - 6 / / , w i t h a
single septum near one end, hyaline; according to Petch smaller
ascospores with two almost equal cells m a y also be produced.

On Boleti, especially species of Xerocomus, Paxillus involutus and Rhizo-


pogon, in autumn. Very common in the mycelial state, which bears enormous
numbers of globose, yellow, warted chlamydospores and is then called
Sepedonium chrysospermum Fries (see lower spore in PL XXXIA) but rarely
found with perithecia.
Apiocrea tulasneana (Plowright) Petch, in J o u r n a l of B o t a n y 75:220
(1937), also on Boleti, is a similar fungus w i t h r a t h e r greenish subi-
culum and perithecia and oblong chlamydospores.

PSEUDONECTRIA Seaver
Pscudonectria Seaver in Mycologia 1 : 4 8 (1909).
Perithecia scattered, superficial, without a common h y p h a l web,
bright coloured, small, asci 8-spored, ascospores hyaline, non-septate,
elliptical.

Pseudonectria rousseliana (Montagne) Wollenweber, Fusaria auto-


graphice delineata No. 665 (1930). ( P L X X X I M . )
Perithecia gregarious, superficial, subglobose to subconical, up to
0.3 m m . across, straw-coloured to light red, covered with stiff, scatter-
ed, spreading, septate, white, cylindrical hairs up to 100// long. Asci
cylindric-clavate, up to 6 0 x 8 f i , 8-spored; ascospores becoming bi-
seriate, elliptical, 9 - 1 6 x 3 - 5 / / , hyaline.

On leaves of Ruxus, in spring and autumn. Not common. The conidial


state is Volutella buxi (De Candolle ex Fries) Berkeley & Broome, see Acta
Bot. Neerlandica 12:58-63 (1963).

NECTRIA Fries

Nectria Fries, S u m m a Veg. Scand. Sect. Post.:387 (1849).


Perithecia occurring in clusters on an erumpent stromatic base,
often in association with a conidial state, or scattered on t h e surface

240
of b a r k or stems, small, subglobose to pear-shaped, with an apical
ostiole, sometimes on a distinct papillae or in the centre of a flattened,
discoid, apex, often collapsing when dry and t h e n appearing cup-
shaped like members of the Helotiales; wall smooth, finely warted,
sometimes covered with loose light-coloured, scurfy cells, brightly
coloured, usually some shade of yellow or red, in a few species (sub-
genus Lasionectria Saccardo) bearing downy white hairs. Asci cylin-
drical, 8-spored, at least at first; ascospores uniseriate or irregular,
elliptical, with a more or less median septum, hyaline or in several
species very pale brown at m a t u r i t y , smooth or v e r y finely verrucose,
longitudinally striate in N. peziza and numerous tropical species,
p r i m a r y spores budding at t h e poles to fill t h e ascus with allantoid
secondary spores in N. aquifolii and N. coryli. The British species,
are all described in detail, w i t h particulars of their conidial states,
in a monograph b y C. Booth, Commonwealth Mycological I n s t i t u t e
Mycological P a p e r : 73 (1959) on which t h e following summary is bas-
ed.

Dialonectria (Saccardo) Cooke in Grevillea 12:109 (June 1884) has


been distinguished from Nectria solely b y its perithecia occurring
scattered over the substrate instead of being clustered u p o n a stroma-
tic base. The distinction is difficult to apply and probably inconstant
so t h a t modern authors do not a t t e m p t to separate the two genera.
Lasionectria (Saccardo) Cooke in Grevillea 12:111 (June 1884) is a
small genus proposed for species with perithecia similar to those of
Nectria b u t scattered and covered with short downy hairs, not re-
cognised b y Booth.

Nectria cinnabarina (Tode ex Fries) Fries, S u m m a Veg. Scand. Sect.


P o s t . : 3 8 8 (1849). (PL X X X I r . )
Perithecia densely crowded on a small e r u m p e n t , cushion-like,
stromatic base, 0.4 m m . across, cinnabar-red, darkening with age,
minutely warted, each with a small, apical, papillate ostiole. Asci
cylindric-clavate, up to 70x12,«, 8-spored; ascospores biseriate, ellip-
tic-cylindric, 1 2 - 2 5 x 4 - 9 f i , with a median transverse septum, hyaline;
paraphyses colourless, branched.

On dead sticks of all kinds, throughout the year. Very common, usually
associated with the pink to light red cushions of its conidial state Tuber-
cularia vulgaris Tode ex Fries.

241
Nectria cinnabarina is one of the commonest fungi on dead sticks of
broad-leaved trees and shrubs, especially in its Tubercularia conidial
state, known from its bright coral-red colour as "Coral S p o t " . This
has an extensive synonymy, fully discussed b y Petch in Trans. Brit,
mycol. Soc. 2 4 : 3 3 - 5 8 (1940), where 40 names are cited. It m a y occa-
sionally be weakly parasitic, as when it infects a short dead stub and
grows back from this into the uninjured branch behind. The fungus
is particularly a b u n d a n t on Ribes and the race on this host m a y be
separable at the varietal level on account of its slightly larger asco-
spores and conidia, see H. A. Jorgensen in K. vet. Hojsk. A a r s k r :
57-120 (1952).

Nectria ralfsii Berkeley & Broome in Ann. Mag. n a t . Hist. Ser. 2,13:467
(1854). (PI. XXXA.)
Perithecia gregarious round the edge of a dark conidial sporo-
dochium erumpent from b a r k , subglobose, flattened above, less t h a n
% m m . diameter, light yellow to orange, with a finely warted surface.
Asci 7 5 - 9 6 x 1 2 - 1 6 / / , spores biseriate, 18-23x 6 - 8 / t . Conidial state a
Myrothecium, w i t h dark green conidia 1 0 - 1 4 x 7 - 8 / / .
On Acer, Pyrus and Ulmus. September to January. Uncommon.

Nectria aquifolii (Fries) Berkeley, Outlines of British Fungology:


393 (1860). (PI. X X V I I I H . )
Perithecia clustered on a yellow to red tuberculate stroma erumpent
from bark, about y 3 m m . diameter, globose, collapsing when dried,
with a red wall covered b y loose, mealy, greenish-yellow cells. Asci
7 0 - 8 0 x 6 - 7 jU, with 8 ascospores 9-12 x 3 - 5 / t which bud off innumerable
allantoid microconidia, 3 - 5 x 1 - 1 . 5 / / , filling the ascus and expanding
it to 9 0 - 1 0 0 x 8 - 9 / / . Similar conidia are produced b y phialides in
culture.
On bark of Ilex aquifolium, throughout the year. Uncommon.

Nectria coryli Fuckel, Fungi Rhenani Exs. Suppl. 1, No. 1582 (1865).
Perithecia clustered on a tuberculate stroma erumpent through
bark, globose, collapsing when dry, a b o u t y 3 m m . diameter, red,
without a mealy surface layer. Asci 60-65 x 6 - 8 |x, with 8 ascospores
1 1 - 1 3 x 3 - 3 . 5 / / , which b u d off innumerable allantoid microconidia
3 - 5 x 1 - 1 . 5 / t to fill the ascus and expand it to 7 5 - 1 0 0 x 9 - 1 2 / / . Similar
conidia are borne b y short phialides on the stroma surface.
On Corylus and Salix in Winter. Rare. (PI. XXVIIIJ.)

242
Nectria sinopica Fries, op. cit.:388 (1849). (PI. XXVIIIA.)

Perithecia clustered on a yellow to red tuberculate stroma erumpent


t h r o u g h bark, globose, collapsing when dry, m m . diameter,
light red but more or less covered b y loose, mealy, greenish-yellow
cells. Asci 5 5 - 7 0 x 6 - 8 / / , spores uniseriate, 9 - 1 2 x 4 - 6 / / . Globose red
mm
pycnidia, - diameter, containing phialides bearing conidia
2 - 3 x 1 //, usually precede perithecial production.
On dead stems of Hedera helix. September to January. Not uncommon.

Nectria pallidula Cooke in Grevillea 1 7 : 3 (1888). (PI. X X X c . )


Perithecia clustered on a yellow stroma erumpent from b a r k ,
globose, m m . diameter, w i t h minutely verrucose wall, straw
yellow with a apical brown papilla. Asci 5 5 - 8 0 x 7 - 1 2 / / , spores 10-14x
3.5-5/t, light brown with finely w a r t e d wall at m a t u r i t y . The conidial
state is a Gliocladium.
On Acer, Aesculus, Fagus, Robinia, Solarium, Vlmus throughout the year.
Uncommon.

Nectria keithii Berkeley & Broome in Ann. Mag. n a t . Hist. Ser. 4, 17:
144 (1876). (PI. XXXD.)
Perithecia solitary or in small clusters amongst whitish Gliocladium
sporodochia, ovoid to globose, % m m . diameter, yellow to light brown
with finely verrucpse wall, ostiole surrounded b y a collar of short
hyphae, giving a flattened top to the perithecia. Asci 5 0 - 7 0 x 7 - 1 0 / / ,
spores 8 - 1 2 x 3 . 5 - 5 / « .
On old stalks of cultivated Brassica in spring. Rare.

Nectria solani Reinke & Berthold, Zersetz. d. Kartoffel:39 (1879).


Perithecia clustered on a yellow e r u m p e n t stroma, associated with
white sporodochia of a Gliocladium, ovoid to globose, about % m m .
tall, yellowish-brown, with minutely verrucose wall. Asci 6 0 - 7 5 X 7 - 8 / / ,
spores 1 0 - 1 4 X 3 . 5 - 4 / / , hyaline with a p u n c t a t e wall. (PI. X X X B . )
On dead stems of Solarium tuberosum. May to September. Rare.

Nectria coccinea (Persoon ex Fries) Fries, op. cit.:388 (1849).


Perithecia scattered over a tuberculate stroma developed in b a r k
or scattered over bare wood without a stroma, ovoid, about mm.
diameter, bright red, with a darker apical papilla, smooth walled. Asci

243
7 5 - 1 0 0 x 7 - 1 0 / / , spores 1 2 - 1 5 x 5 - 6 / / , hyaline to very pale brown. The
conidial state is a Cylindrocarpon, with macroconidia 2 0 - 8 0 x 5 - 7 / / ,
0-7-septate. ( P I . X X V I I I D . )

Common on Fagus but also reported on Acer, Alnus, Betula, Carpinus,


Castanea, Corylus, Fraxinus, Hedera, Ilex, Morus, Populus, Quercus, Rham-
nus, Sambueus, Taxus, Tilia and Ulmus. Throughout the year.

Nectria galligena Bresadola apud Strasser in Verh. K . K . Zool. Bot. Ges.


Wien 51:413 (1901). (PI. X X I X A . )
Perithecia scattered or in small clusters, 1 / 4 r 1 / 3 m m . diameter,
ovoid, bright red, smooth to very finely verrucose. Asci 75-95 x 12-15//,
spores 1 4 - 2 2 x 6 - 9 / / , hyaline. The conidial state is Cylindrocarpon mali
(Allescher) Wollenweber, with conidia 1 0 - 6 5 x 4 - 7 / / , 1-5-septate.

On cankers of deciduous tress, especially Fraxinus, Malus, Pyrus and


Salix, reported also on Acer, Aesculus and Betula. Throughout the year.
Common.

Nectria ditissima Tulasne, Selecta F u n g o r u m Garpologia 3 : 7 3 (1865).


Perithecia clustered on bark, ovoid to globose, dark red, mm.
diameter, smooth. Asci 8 5 - 9 5 x 15-18//, spores 14-21x5-8/z, hyaline.
Conidial state a Cylindrocarpon with macroconidia 5 0 - 9 0 x 3 - 6 / / ,
4-7-septate. (PI. X X I X B . )

On Fagus, associated with canker and dieback. October to May. Not


common.

Nectria punicea (Schmidt ex Fries) Fries apud Rabenhorst Herb. Myc.


No. 634 (1858).
Perithecia crowded on an erumpent stroma, ovoid to globose, with
a m a r k e d apical papillae, a b o u t % m m . diameter, reddish-brown,
smooth. Asci 9 0 - 1 2 0 x 1 1 - 1 2 / / , spores 14-16x6/u.

Var. punicea on Rhamnus frangula in W i n t e r and Spring. Un-


common.
Var. ilicis Booth, op. cit.:54 (1959) with larger ascospores 14-24x
7-10/«, on branches of Ilex aquifolium, t h r o u g h o u t the year. Not un-
common. (PI. X X V I I I B . )
The conidial state is a Cylindrocarpon with 7-9-septate conidia,
90-115x6-8/«.

244
Nectria fuckeliana Booth, op. c i t . : 5 6 (1959). (PI. XXVIIIG.)

Perithecia clustered on a stroma erumpent t h r o u g h bark, ovoid,


about % m m . diameter, red to maroon, smooth. Asci 86-120x8-12/«,
spores 1 3 - 1 6 x 5 - 6 / / , finally pale brown and finely verrucose. Macro-
conidial state a Fusarium, with orange sporodochia and 3-septate
spores 3 3 - 4 0 x 4 - 5 / / .
On Abies, Larix and Picea in Spring and Summer. Not common.

Nectria hederac Booth, op. c i t . : 5 9 (1959). (PI. XXIXJ.)

Perithecia scattered, on bark or cut ends of stems, ovoid with


flattened ostiolar disc, m m . diameter, dark red, smooth. Asci
8 0 - 9 5 x 7 - 9 / / , spores 1 2 - 1 5 x 4 . 5 - 6 / / , finally pale brown and finely
verrucose. Macroconidial state a Cylindrocarpon, conidia mostly
5-septate, 5 2 - 7 8 x 5 - 7 // b u t there is also a distinctive microconidial
state in t h e form-genus Paecilomyces, with chains of conidia 6 - 9 x
2-3 ju,.
On Hedera helix. Winter. Rare.

Nectria mammoidea Phillips and Plowright in Grevillea 3:126 (1875).


Perithecia scattered, subglobose, m m . diameter, with convex
apical disc, orange to dark reddish-brown, blackish at the apex, wall
smooth. Asci 9 0 - 1 3 0 x 8 - 1 0 / / , spores 1 6 - 1 7 x 6 . 5 - 7 / / , hyaline to faintly
brownish. Conidial state a Cylindrocarpon, with conidia 5 4 - 7 2 x 6 - 7 / / ,
3-7-septate. (PI. X X I X C . )
On dead twigs and stems of Acer, Brassica, Fagus, Populus, Quercus,
Smyrnium, Ulex and Ulmus.

Var. rubi (Osterwalder) Weese in Zeitschr. Garungsphysiol. 1 : 1 2 9


(1912), with ascospores 1 2 - 1 4 x 5 - 6 . 5 / / occurs on roots of Rubus idaeus.
Var. rugulosus Weese in Sitzber. Akad. Wiss. Wien Math. Nat. Kl.
125, A b t . 1:552 (1916) with more verrucose perithecia and ascospores
1 8 - 2 1 x 6 - 7 / t is recorded on Acer and Quercus.

Nectria veuillotiana Roumeguere & Saccardo in Rev. Mycol. 2:189


(1880). (PI. XXXIG.)
Perithecia mostly in groups on an erumpent stroma, globose with
mm
a flat apical disc surrounding the minutely papillate ostiole, -
diameter, yellow, t h e n bright red, finally brownish, wall finely warted.
Asci 1 0 8 - 1 3 0 x 9 - 1 2 / / , spores uniseriate, hyaline, 17-22x7-9/«. Conidial
state a Cylindrocarpon with conidia 3 0 - 6 0 x 5 - 8 / / , up to 7-septate.

245
On twigs of Fagus, Populus and Salix, the type collection was on Gle-
dilschia in France. October to March. Uncommon.

Nectria pinea Dingley in Trans. Roy. Soc. New Zealand 79:198 (1951).
Perithecia scattered, globose, with a conical apical papilla, up to
% m m . diameter, orange to reddish-brown, smooth. Asci 8 0 - 1 1 0 x 8 -
11 //, spores 1 6 - 2 2 x 7 - 1 0 //, light brown. The conidial state is a Cylin-
drocarpon with conidia 3 0 - 7 5 x 3 - 7 / / , 1-5-septate. (PI. X X V I I I F . )
On bark of conifers, Abies,, Larix, Pinus, Pseudotsuga. Spring and Autumn.
Uncommon.

Nectria episphaeria (Tode ex Fries) Fries, op. cit.:388 (1849).


Perithecia scattered, ovoid, with an apical papilla, 1 I S - 1 I 5 m m . dia-
meter, bright red, smooth. Asci 5 6 - 7 0 x 5 - 6 / / , spores 7 - 1 1 x 3 . 5 - 5 / / ,
brown and verrucose at m a t u r i t y . Conidial state a Fusarium with
spores 3 5 - 5 0 x 2 . 5 - 3 . 5 n, 2-5-septate. (PI. X X V I I I E . )
On effete perithecia and stromata of other ascomycetes, especially Qii-
trygaceae and Xylariaceae. Throughout the year. Common.
This and similar species in which there is no basal stroma have been
referred to a separate genus, Dialonectria (Saccardo) Cooke in Grevillea 12:
109 (1884) but there is no clearcut distinction between them and typical
species of Nectria.

Nectria desmazierii Beccari & de Notaris, E r b . Critt. Ital. No. 983


(1863). (PI. X X X E . )
Perithecia clustered on the remains of a conidial sporodochium,
ovoid to globose, deep yellow to orange, smooth, about 1 / 8 m m . dia-
meter. Asci 6 5 - 8 0 x 7 - 9 / / , spores 1 2 - 1 6 x 5 - 7 / / , hyaline. Conidial state
Fusarium buxi Saccardo with spores 4 5 - 6 0 x 3 - 5 //, 3-6-septate.
On twigs, associated with leafscars, and bark of Buxus sempervirens in
Autumns. Rare.

Nectria magnusiana Rehm apud Saccardo in Michelia 1:294 (1878).


Perithecia clustered, globose with a flat apical disc, red, darkening
mm
with age, - diameter, smooth. Asci 7 2 - 8 4 x 9 - 1 2 / / , spores
1 0 - 1 5 x 4 . 5 - 6 / / , finally light brown. Conidial state gelatinous sporo-
dochia of Dendrodochium epistroma (Sydow) von Hohnel, with allan-
toid conidia, 4 . 5 - 6 x 1 . 5 - 2 / / . (PI. X X I X D . )
On stromata of Diatrypella. August to April. Uncommon.

246
Nectria purtonii (Greville) Berkeley, Outlines of British Fungology:
394 (1860). (PI. XXXM.)
Perithecia clustered, pyriform, with a flat apical disc, yellow to red,
1
lsr1A m m - diameter, smooth. Asci 5 5 - 7 0 x 6 - 9 / / , spores 8 - 1 1 x 3 . 5 -
4.5//, finally light brown. The conidial state is a Fusarium with
1-septate conidia 2 0 - 2 4 x 1 . 5 - 2 / / .
On perithecia of Valsa spp. on twigs of deciduous trees. March to June.
Rare.

N. applanata Fuckel, Symb. Myc. Nacht. 1:22 (1871) is a synonym.

Nectria leptosphaeriae N'iessl apud Krieger Fungi Saxonici 4, No. 165


(1886). (PI. XXIXG.)
Perithecia scattered, each with a small m a t of white anchoring
hyphae, pyriform, orange to red, a b o u t % m m . diameter, smooth.
Asci 85 x 15/i, spores 1 8 - 2 6 x 6 - 8 / / , brownish at m a t u r i t y . The conidial
state is Fusarium sphaeriae Fuckel.
On and amongst pseudothecia of Leptosphaeria acuta in Winter. Fairly
common.

Nectria polyporina Petch in Trans. Brit, mycol. Soc. 25:169 (1941).


Perithecia clustered, pyriform, 1 / 5 m m . diameter, light red, smooth.
Asci 6 0 - 7 0 x 7 - 8 / / , spores hyaline, 1 2 - 1 4 x 3 - 4 . 5 / / . (PL X X I X L . )
On old carpophores of Fomes annosus. Rare.

Nectria flavoviridis (Fuckel) Wollenweber, Angew. Bot. 8:186 (1926).


Perithecia scattered, pyriform, red, papilla becoming v e r y dark,
about m m . diameter, wall smooth. Asci 8 5 - 1 0 4 x 7 . 5 - 9 / / , spores
1 0 - 1 6 x 5 - 8 / / , brownish at m a t u r i t y . There is an associated greenish-
yellow stilboid conidial state bearing Fusarium-like conidia 12-17 x
2 - 3 / / , 1-septate; larger 3 - 5 - s e p t a t e conidia develop in culture.
On bark of Acer, Betula, Fraxinus, Pinus, Quercus and Salix. Spring and
Autumn. Uncommon. (PL XXXj.)

Sphaerostilbe flavoviridis Fuckel in J a h r b . Nass. Vereins f. N a t u r -


kunde 25/26:310 (1871) is a synonym.

Nectria viridescens Booth, op. cit.:89 (1959). (PL XXIXE.)

Perithecia scattered, pyriform, with an apical disc, yellow, reddening


with age, 1 / 8 - 1 /s m m . diameter, smooth. Asci 6 8 - 7 6 x 6 - 9 / / spores

247
7 . 5 - 1 0 x 4 - 5 f i , brown and verruoose at m a t u r i t y . The conidial state
develops s y n n e m a t a bearing clusters of phialides with conidia 4 - 8 x
2 - 3 / / , pink in the mass.
On bark of Acer, Betula, Fagus, Fraxinus, Pinus and Salix. April to
June. Apparently not uncommon. The specific epithet refers to a charac-
teristic green pigment developed in agar culture.

Nectria rishbethii Booth, op. cit.:92 (1959). (PI. XXXK.)


mm
Perithecia pyriform, - diameter, ochraceous to brownish,
minutely verrucose. Asci 60-110x6-11/«, spores 8 - 1 2 x 3 . 5 - 5 / « , brown
and finely verrucose at m a t u r i t y . Conidial state like t h a t of the pre-
ceding, with conidia 2 . 5 - 4 x 1 - 2 / « . Immersed in resin exuded from
Pinus t r u n k s and stumps, superficially like the commonly associated
pycnidia of Pycnidiella resinae (Fries) von Hohnel, as in the figure.

Nectria brassicac Ellis & Saccardo in Michelia 2:374 (1881).


Perithecia scattered, globose, with an apical papilla, yellow to
orange, 1 / i 0 ~ % m m . diameter, smooth. Asci 40-65x7.5-14/«, spores
10-16x4-5/«, finally brownish. Associated conidial state a Fusarium
with 4-6-septate spores, 4 0 - 4 5 x 4 - 4 . 5 / / . (PI. X X I X F . )
On rotting stems of cultivated Brassica. Spring and Autumn. Rare.

Nectria peziza (Tode ex Fries) Fries, op. cit.:388 (1849).


Perithecia scattered or gregarious on a hyphal web, globose b u t
collapsing to appear cupulate when dry, pale to deep yellow,
m m . diameter, smooth. Asci 75-90x8-10/«, spores 12-15x5-7/«,
hyaline, wall with fine longitudinal striations when mature. Conidia,
elliptical, hyaline, 5 - 1 1 x 2 . 5 - 4 / « , are produced on scattered phialides.

On very rotten wood, decaying polypores & c., especially in Summer and
Autumn. Common. (PI. XXVIIIc.)

Nectria arenula (Berkeley & Broome) Berkeley, Outlines of British


Fungology:394 ( 1 8 6 0 ) . ( P I . X X X F . )
Perithecia scattered, ovoid, collapsing when dry, yellow to yellowish
brown, up to % m m . diameter, smooth to minutely verrucose. Asci
45-60x8-10/«, spores 15-18x3.5-4/«, hyaline, faintly striate.
On dead herbaceous stems and debris of Monocotyledons, Carex, De-
schampsia, Glyceria, Helianthus, Heracleum, Typha. September to February.
Uncommon.

248
Nectria citrino-aurantia Delacroix apud Desmazieres PI. Crypt. Franco
Ser. 2, N o . 7 2 8 (1860). (PI. XXXL.)

Perithecia densely crowded on a tuberculate stroma erumpent from


b a r k , subglobose, 1/10—1/8 m m . diameter, yellow, smooth. Asci 3 0 - 3 8 x
5//, spores hyaline, 7 - 8 x 2 . 5 - 3 / / .

On Salix. December. Rare.

Nectria inventa Pethybridge in Trans. Brit, mycol. Soc. 6:104 (1919).


m m
Perithecia clustered, ovoid, - diameter, yellow-brown,
bearing stiff hyaline hairs on the upper p a r t . Asci 60-100x4-6/«,
spores hyaline, 8 - 1 2 x 3 - 3 . 5 / / . The conidial state is Verticillium cinna-
barinum (Corda) Reinke and Berthold. (PI. X X I X I I . )

On decayed Brassica and Solarium tuberosum. Rare.


The remaining species have minutely downy perithecial walls and have
been segregated on that account in a genus Lasionectria (Saccardo) Cooke
in Grevillea 12:111 (1884), not recognised at generic level by Booth.

Nectria ellisii Both, op. cit.: 101 (1959). (PI. XXXG.)

Perithecia scattered or gregarious, globose, % m m . diameter,


yellow, clothed with downy white hairs up to 7 0 x 4 / / . Asci 35-55 x
6 - 8 / / spores hyaline, 1 1 - 1 2 x 3 - 4 / / . Conidial state a Cephalosporium.

On dead herbaceous stems, Epilobium, Eupatorium, Filipendula, Lychnis,


Phragmites and Rubus. Spring and Autu mn. Not common.

This is the fungus which has been called Lasionectria jlavida (Corda)
Cooke in Grevillea 12:112 (1884) in British literature but according
to Booth it does not agree w i t h the description or figures of the basi-
n y m Nectria flavida Corda, which was on Alnus.

Nectria funicola (Berkeley & Broome) Berkeley, Outlines of British


Fungology: 393 (1860). (PL X X X H . )

Perithecia scattered, pyriform, Y ^ U m m - diameter, yellow to


orange, clothed with downy white hairs 40-55x4/n. Asci 70-95 x
12-17//, spores hyaline, 1 6 - 2 0 x 6 - 8 / / .

249
On decaying rope, cardboard and paper. Autumn. Rare.

Nectria aureola W i n t e r in Hedwigia 2 4 : 2 6 1 (1885). (PI. XXIXK.)

Perithecia scattered, globose, clothed with rigid white hairs on a


light yellow wall. Asci 47-53x7/«, spores hyaline, 14x2.5 fi.
On mycelium of Meliola niessleana. Summer. Very rare.

Nectria lecanodes Cesati in Rabenhorst, Herb. myc. E d . 2, 6, No. 525


(1857). (PI. X X X I E . )
Perithecia superficial, scattered or in small groups, subglobose, be-
coming flattened when dry, pale pink, about 0.25 m m . across, clothed
with fine white downy hairs up to 40x3/«. Asci cylindric-clavate, up
to 7 0 x 6 / i , 8-spored; ascospores biseriate, narrowly elliptical, 1-sep-
t a t e , 9 - 1 2 x 3 - 4 / / , hyaline, finely p u n c t a t e .
On thalli of frondose lichens, especially of Peltigera canina, in autumn.
Not common. The hairs of the perithecia may easily be overlooked as they
are apt to become matted and prostrate.

SPHAEROSTILBE Tulasne

Sphaerostilbe Tulasne, Selecta F u n g o r u m Carpologia 1:130 (1861).


Here again the perithecia and their contents resemble those of
Nectria b u t t h e y occur in small clusters around a club-shaped struc-
t u r e which carries conidia.

Sphaerostilbe aurantiaca Tulasne, op. cit.: 130 (1861). (PI. XXXIH.)

Conidial body cylindrical or compressed, up to 2 m m . tall, reddish-


brown, paler in the upper p a r t , swelling into a subglobose head which
bears innumerable elliptic-oblong, hyaline, non-septate conidia 14-18x
6 - 8 f i . Perithecia clustered round the conidial body, globose, almost
smooth, red, up to 0.4 m m . across. Asci clavate, u p to 130x22/t,
8-spored; ascospores elliptical, 1-septate, hyaline, rather thick-walled
and smooth, 2 0 - 2 9 x 8 - 1 0 / / .

On bark of Ulmus in autumn and winter. Rare.

Sphaerostilbe flammea Tulasne, op. cit.: 130 (1861), is a v e r y rare


species occurring on scale insects on bark of Fraxinus and Salix.

250
NEOHENNINGSIA Koorders

Neohenningsia Koorders, Botanische Untersuchungen . . . J a v a . . .


Pilze: 164 (1907).
Perithecia scattered and like those of a Nectria b u t bearing a
crown of thin, spreading, triangular teeth. There is a single British
species:

Neohenningsia suffulta (Berkeley & Curtis) von Hohnel apud Petch in


Trans. Brit, mycol. Soc. 2 1 : 2 6 8 ( 1 9 3 8 ) . ( P L X X V I I L . )
Perithecia superficial, more or less gregarious, subglobose b u t some-
w h a t flattened above, up to 0.5 m m . across, orange-yellow, smooth
a p a r t from the ring of flat, spreading, triangular processes up to 110/<
long which fringes the flattened upper surface. Asci cylindric-clavate,
up to 75x11//, 8-spored; ascospores elliptic-oblong, 1-septate, 12-14x
4 - 5 / / , hyaline, smooth.

This species is common on vegetable debris throughout the tropics but


is a doubtful native of the British Isles where it is known only from a single
collection on plant fibres mixed with horse dung at Kew.

NECTRIOPSIS Maire

Nectriopsis Maire in Annales mycologici 9 : 3 2 3 (1911).


Perithecia scattered over a common h y p h a l web, otherwise like
those of a Nectria.

Nectriopsis aureonitens (Tulasne) Maire, op. cit.:323 (1911). (PL


XXXIc.)
Perithecia subglobose, less t h a n 0.2 m m . across, golden-yellow with
a dark dot at the top marking the ostiole, smooth or minutely downy,
gregarious on a whitish to yellow h y p h a l m a t . Asci cylindric-clavate,
up to 90x7/«, 8-spored; ascospores uniseriate, oval, 9 - 1 3 x 3 - 4 / « ,
1-septate and constricted at the septum, t h e u p p e r cell broader t h a n
the lower, hyaline, finely p u n c t a t e .
The hyphal m a t also bears the small, white, brush-like conidio-
phores of the conidial state Gliocladium penicillioides Corda.

On old fructifications of Stereum in Autumn. Uncommon.

251
Nectriopsis violacea (Schmidt in Fries) Maire, op. cit.:323 (1911),
with a violet h y p h a l m a t , occurs occasionally on the large spreading
fructifications of the Myxomycete Fuligo septica.

NECTRIELLA Nitschke

Nectriella Nitschke apud Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e


23-24:175 (1870).
Perithecia subglobose, smooth, brightly coloured, erumpent but
otherwise like those of Nectria.

Nectriella robergei (Montagne & Desmazieres) Weese in Annales myco-


logici 1 2 : 1 3 8 ( 1 9 1 4 ) . ( P I . X X X I D . )
Perithecia scattered, immersed within the host tissue, t h e n splitting
it to expose their apices, ovoid, smooth, up to 0.25 m m . across, light
red. Asci clavate, up to 8 5 x 1 3 / / , 8-spored; ascospores biseriate,
elliptic-cylindric, 10-15x4-7/«, 1-septate b u t only slightly constricted
at the septum, smooth, hyaline, containing several oil drops.
Parasitic on the thallus of the lichen Peltigera canina and not to be con-
fused with Nectria lecanod.es on the same host, in autumn. Not uncom-
mon.

CALONECTRIA de Notaris

Calonectria de Notaris in Commentario della Societa Crittogamologica


italiana 2:477 (1864).
Perithecia superficial, scattered, brightly coloured, not hairy, asco-
spores hyaline, elliptical or fusiform, with more t h a n one septum. A
n u m b e r of species have been reported from the British Isles b u t the
only one at all common or well known is:

Calonectria ochraceopallida (Berkeley & Broome) Saccardo in Atti Soc.


Venet.-Trent. Sci. Nat. 4 : 1 2 4 (1875). (PI. X X V I I M . )
Perithecia scattered or gregarious, superficial, globose, up to 0.35
m m . across, slightly flattened above with a minute apical papilla,
smooth, ochraceous to reddish-orange. Asci clavate, up to 100x22//,
8-spored; ascospores more or less biseriate, narrowly fusoid, mostly
2 5 - 5 0 x 4 - 6 / « , 3-8-septate, slightly constricted at the septa, hyaline.

On bark of various trees and shrubs and on woody stems, in winter. Un-
common.

252
GIBBERELLA Saccardo

Gibberella Saccardo in Michelia 1 : 4 3 (June 1877).


Perithecia superficial, clustered on an erumpent stroma or scattered,
soft textured b u t very dark blue, bluish-purple or black, ascospores
with two or more septa, hyaline.

Gibberella cyanogena (Desmazieres) Saccardo, Sylloge F u n g o r u m 2:


555 (1883). (PI. X X V I I I . )
Perithecia superficial, scattered or in small clusters with an in-
conspicuous stromatic base, ovoid or subcylindrical with rounded tips,
about 0.25 m m . across, coarsely warted, black when viewed whole b u t
t h e wall appears blue when a crushed perithecium is viewed b y trans-
mitted light. Asci clavate, up to 100x18,«, 8-spored; ascospores often
biseriate, narrowly elliptical, straight or slightly curved, 2 0 - 3 6 x 5 - 7 //,
usually 3-septate, sometimes 4-6-septate, slightly constricted at the
septa, hyaline.

On decaying herbaceous stems, especially of Brassica, less often on dead


twigs, autumn, to spring. Common.

Gibberella pulicaris (Fries) Saccardo in Michelia 1 : 4 3 (1877), found


on similar substrata, has shorter, relatively broader ascospores,
16-24x6-9//.
Gibberella zeae (Schweinitz) Petch in Annales mycologici 34:260
(1936), is common on dead culms and stubble of cereals, less often on
the ears, especially in d a m p fields or on laid crops or plants damaged
by other fungi. It occurs commonly on grain in its conidial state
Fusarium graminearum Schwabe.
Gibberella moricola (de Notaris) Saccardo, Sylloge F u n g o r u m 2 : 5 5 3
(1883), occurs occasionally on dead twigs of Ficus and Morus.
Gibberella buxi (Fuckel) Winter, Rabenhorst's K r y p t o g a m e n Flora
Deutschland 1 (2): 103 (1887), occurs rarely on twigs and dead leaves
of Buxus.

ACTINIOPSIS Starback

Actiniopsis Starback in Bihang K. Svensk. Vet.-Akad. Handl. 25 (3):


54 (1899).
Perithecia very similar to those of Neohenningsia b u t containing
asci with multiseptate ascospores. There is only one British species:

253
Actiniopsis peristomialis (Berkeley & Broome) Petch in Trans. Brit,
mycol. Soc, 2 1 : 2 8 2 ( 1 9 3 8 ) . (PI. X X V I I K . )
Perithecia superficial, globose or subcylindrical, about 0.2 m m .
across b u t up to twice as tall, somewhat flattened at the apex which
is surrounded b y a ring of spreading whitish triangular teeth, re-
mainder of the wall smooth, yellowish. Asci cylindric-clavate, up to
90x10/«, 8-spored; ascospores often biseriate, elliptic-fusiform, 2 0 - 2 7 x
4 - 5 / t , 3-5-septate, hyaline.
On dead bark of Ilex aquijolium. Rare, reported only from Cornwall.

PARANECTRIA Saccardo

Paranectria Saccardo in Michelia 1 : 3 1 7 (July 1878).


Perithecia superficial, brightly coloured, smooth, soft, ascospores
hyaline, 3-septate, ciliate at each end. There is a single British species:

Paranectria affinis (Greville) Saccardo, op. cit.:317 (1878). (PL


XXVIIII.)

Perithecia superficial, scattered or in small clusters, globose, an-


chored b y a fringe of basal hyphae, smooth, pinkish-orange, up to
0.3 m m . across. Asci cylindric-clavate, up to 4 5 x 1 8 f i , 8-spored; asco-
spores irregularly arranged or lying parallel, fusiform, 24-34x6-8/«,
3-septate, hyaline, drawn out at each end into a slender bristle up to
15/f long.
On thalli of the lichen Ephebe lanata. Apparently rare but likely to be
overlooked.

SCOLECONECTRIA Seaver
Scoleconectria Seaver in Mycologia 1 : 1 9 7 (1909).
Perithecia clustered on a stroma, resembling those of Nectria b u t
containing long, cylindrical or worm-like multicellular primary asco-
spores, which m a y b u d off m i n u t e conidia to fill the ascus. The genus
is a segregalfi-4r©^TOphionectrid) Saccardo, the t y p e species of which
J i a r S o l i t a r y perithma-ftftd-a^cospores which do not bud in the ascus.

Scoleconectria cucurbitula (Tode ex Fries) Booth, Comm. Myc. Inst.


Myc. P a p . 72:15 (1959). (Pl. X X V I I J . )
Perithecia borne in clusters on a feebly developed stromatic base
which emerges from cracks in bark, globose, up to 0.3 mm. across, red

254
blackening with age, commonly collapsing when dry so as to resemble
one of the Helotiales, smooth or minutely scurfy. Asci clavate, up to
9 0 x 2 0 / / containing either eight ascospores or innumerable minute
rod-like secondary spores 2 - 4 x 0 . 7 5 / / ; p r i m a r y ascospores cylindrical
or worm-like, often curved, 2 2 - 4 0 x 3 - 4 / / , hyaline, multiseptate at
short intervals.

On twigs of Pinus, in spring. Not common.

Ophionectria cylindrospora (Sollmann) Berlese & Voglino, Syll. Fung.


A d d . : 2 1 7 (1886) is a synonym.
Ophionectria cerea (Berkeley & Curtis) Ellis & E v e r h a r t , N. Amer.
P y r e n o m . : 118 (1892), produces minute yellowish-brown perithecia on
old s t r o m a t a of Diatrype stigma, and is a Tubeufia, T. cerea (Berkeley
& Curtis) Booth.

TRICHONECTRIA Kirschstein
Trichonectria Kirschstein in Verhandl. Bot. Ver. Prov. B r a n d e n b u r g
4 8 : 6 0 (1906).
Perithecia superficial, scattered, soft-textured, brightly coloured,
bearing stiff, spreading white hairs, ascospores elongated, multi-
septate, hyaline.

Trichonectria hirta (Bloxam) Petch in The Naturalist:282 (1937).


(PI. X X V I I G . )
Perithecia superficial, scattered, globose, up to 0.25 m m . across,
pinkish-salmon colour, wall smooth b u t bearing rather sparsely dis-
t r i b u t e d , stiff, white hairs up to 110// long b y 8ju at the base. Asci
clavate, 8-spored; ascospores cylindric-fusoid, straight or worm-like,
closely multiseptate, 50-95x5-9/«, hyaline.
On decorticated wood, in autumn. Uncommon.

THYRONECTRIA Saccardo

Thyronectria Saccardo in Grevillea 4 : 2 1 (September 1875).


Perithecia clustered on a stromatic base or scattered and super-
ficial, brightly coloured, ascospores hyaline, with b o t h transverse and
longitudinal septa, sometimes budding off minute secondary spores
within the ascus. Species with perithecia wholly superficial were se-

255
gregated in Pleonectria Saccardo in Nuovo Giornale Botanico italiano
8 : 1 7 8 (1876) b u t the distinction is considered untenable by Booth and
others.

Thyronectria berolinensis (Saccardo) Seaver in Mycologia 1:205 (1909).


(PI. X X V I I F . )
Perithecia erumpent in dense clusters, globose, smooth, up to 0.5
m m . across, red or reddish-brown, often collapsing and appearing cup-
shaped. Asci cylindric-clavate, up to 1 0 0 x l 0 / t , 8-spored; ascospores
uniseriate, elliptical, 1 8 - 2 0 x 8 / / , hyaline, with seven transverse septa
and one longitudinal wall.
On dead branches of Ribes. Rare. The common Nectria on this host is
N. cinnabarina,.

T. lamyi (Desmazieres) Seeler in J. Arnold Arboretum 21:449 (1940)


is on Berberis.
T. balsamea (Cooke & Peck) Seeler, op. cit.: 442 (1940) is on b a r k of
conifers.

Family 4. Seliniaceac

There is only one British genus:

SELINIA Karsten

Selinia Karsten in Medd. Soc. Flora F a u n a Fennica 1 : 5 7 (1876).


Perithecia in small clusters embedded in a light coloured stroma
sunken in dung of herbivorous animals, asci large, 8-spored, ascospores
large, elliptical, thick-walled, hyaline. There is a single British species:

Selinia pulchra (Winter) Saccardo, Sylloge F u n g o r u m 2:457 (1883).


(PL X X V I I D . )
S t r o m a cushion-like, up to 3 m m . across, subglobose or irregular and
enclosing one or a few flask-shaped perithecia up to 0.7 m m . across,
with protruding ostioles; flesh of the stroma whitish or yellowish,
upper exposed surface smooth or downy, reddish-orange. Asci clavate,
up to 220x60//, usually with eight ascospores, sometimes less; asco-
spores biseriate, elliptic-fusiform, 4 8 - 6 4 x 2 0 - 2 6 / / , thick-walled, non-
septate, or yellowish, with minutely granular contents.

In dung of cows, horses and sheep, in autumn and winter. Not common.

256
Family 5. Polystigmataceae
Here are assigned the genera of Sphaeriales with hyaline non-sep-
t a t e ascospores contained in perithecia immersed either in a stroma or
directly in tissue of a host p l a n t . The stromatic genera bear superficial
resemblance to those of the Dothideales, from which t h e y are dis-
tinguished by t h e structure of their asci. For a modern account of t h e
family see yon Arx & Muller in Beitrage zur Kryptogamenflora der
Schweiz 11 (1): 157-267 (1954) on whose t r e a t m e n t the following key
is based.
I. Stroma or clypeus absent, perithecia embedded singly in the substrate,
saprophytes or parasites which mature their perithecia in dead tissue
only:
A. Asci large, usually broadest about the middle, rather thick walled,
ascospores over 20 pt long, with milky, yellowish, greenish or
pinkish granular contents; perithecial wall fleshy, ostiole usually
darker, often furnished with bristles Physalospora
B. Ascospores hyaline, transparent, seldom yellowish and then less
than 20/« long
Asci cylindrical or slightly clavate, only short-stalked; perithecia
globose or only slightly flattened, with dark parenchyma-
tous walls Glomerella (p. 258)
II. Perithecia embedded singly or in groups within a stroma or beneath a
shield (clypeus) of stromatic tissue pierced by the ostioles, mostly
parasites:
A. Stromatic tissue light coloured, without an epidermal clypeus:
1. Associated with local thickening of the leaf, on Rosaceae
Poly stigma (p. 259)
2. On grass, ascospores yellow at maturity Gibeilina (p. 262)
B. Stromatic tissue or clypeus dark coloured, at least on the outer
surface:
1. Saprophytes or parasites forming perithecia in dead spots,
stromata small or merely represented by a number of con-
fluent perithecia which are individually less than 200 y
across with dark walls Glomerella (p. 258)
2. Parasites with a subcuticular, subepidermal or intra-epidermal
stroma or a clypeus, with a smooth black upper surface:
a. Perithecium completely immersed in a small stroma in
leaf of Dryas Isothea (p. 260)
b. Perithecia developed beneath a subcuticular or epidermal
clypeus which is perforated by the ostioles
Endodothella and Phyllachora (p. 261)

PHYSALOSPORA Niessl
Physalospora Niessl in Verhandl. N a t . Ver. Briinn 14:170 (1876).
The european species retained in Physalospora b y von Arx and
Muller are mostly on moorland plants, Andromeda, Carex, Kobresia,

257
Luzula, Salix, Tofieldia, Vaccinium and should bo sought on these hosts
in Scotland and northern England. One subarctic species first reported
from the Faroes has been found in the Hebrides, viz:

Physalospora empetri Rostrup, Botany Faroes:310 (1901).


Perithecia scattered, subglobose, immersed in the dead leaf, with a
short erumpent neck, smooth, black. Asci clavate, short-stalked,
90-120 x20//, 8-spored, with a hyaline apical ring not blued by
Melzer's reagent; ascospores biseriate, ellipsoidal or slightly kidney-
shaped, 20-24x8-9//, hyaline. (Fig. 25T.)
On Empetrum nigrum.

GLOMERELLA von Schrenk & Spaulding


Glomcrella von Schrenk & Spaulding in U.S. Dept. Agric. Bureau of
Plant Industry Bull. 44:29 (1903).
Perithecia embedded in plant tissue, scattered or in small clusters
united by a rudimentary stroma or with slight development of a
clypeus round the mouth, ostiole well developed, sometimes papillate
or even beaked, wall composed of a few layers of angular brown cells,
asci very short stalked, 8-spored as a rule, ascospores hyaline, non-
septate, paraphyses slender, hyaline. Some species are known to have
conidial states in the form genera Colletotrichum and Gloeosporium and
m a y be parasitic on fleshy tissues of a wide range of plants, especially
in warm climates. Best known of these is the cosmopolitan Glomerella
cingulata (Stoneman) Spaulding & von Schrenk, which sometimes
causes rotting of grapes, apples and pears in England, see for example
Wormald in Gardener's Chronicle Ser. 3, 88:498-500 (1930).

Glomerella cingulata (Stoneman) Spaulding and von Schrenk, op. cit.:


751 (1903). (Fig. 25v.)
Perithecia flask-shaped, dark brown to black, about 100-300/t dia-
meter and up to 350// tall, often hairy on the ostiolar papilla, immersed
or slightly erumpent from dead plant tissue. Asci 55-70x10-14/«,
spores irregularly biseriate, ellipsoidal, straight or slightly curved,
12-28x4-7//, with densely granular protoplasm at the poles and a
clear central area, paraphyses thread-like, soon disappearing. As-
sociated with pale pink slimy acervuli of the conidial state Colleto-
trichum gloeosporioides Penzig, bearing ellipsoidal conidia mostly
12-20x4-6//.

258
The t y p e host of the ascus state was Ligustrum vulgare and t h a t of
C.gloeosporioides was Citrus b u t the host range is enormous, especially
in w a r m e r climates and m a n y races of the conidial state have been
described on a host basis mainly in the genera Colletotrichum and
Vermicularia when setae are well developed and in Gloeosporium when
t h e y are absent. Von Arx in P h y t o p a t h . Zeitschr. 29:413-468 (1957)
cited 592 names of conidial states which he believed t o be morpho-
logically inseparable from t h a t of G. cingulata. These were mostly
associated with leaf-spot, dieback and canker of woody plants or with
rotting of fruit. The fungus is figured f r o m a lesion on an orchid leaf.
G. cingulata has been extensively used in genetical studies, for an
account of its sexual mechanism see McGahen and Wheeler in Amer.
J . Bot. 38:610-617 (1951).
G. miyabeana (Fukushi) von Arx in P h y t o p a t h . Zeitschr. 29:448
(1957) causes leaf-spot and canker in Salix spp. b u t seems to be little
more t h a n a host-specialised race of G. cingulata.

Glomerella montana (Saccardo) von Arx & Miiller in Beitrage zur


Kryptogamenflora der Schweiz 11 (1):196 (1954). (Fig. 28E.)
Perithecia scattered or a few together, completely embedded in the
mesophyll, visible to the unaided eye as minute blackish dots, sub-
globose, scarcely 0.2 m m . across, with a well developed ostiole often
surrounded by a r u d i m e n t a r y clypeus of dark brown hyphae within
the leaf epidermis; asci pinkish in t h e mass, cylindric-clavate, short-
stalked, up to 7 0 x 1 4 / / , thin-walled, w i t h a minute ring of apical
thickening blued by iodine, 8-spored; ascospores sometimes partially
biseriate, elliptical, 1 2 - 1 8 x 5 - 7 / / , hyaline, non-septate; paraphyses
numerous, slender.

On dead leaves of the grass Sesleria coerulea in Spring. Uncommon.

POLYSTIGMA de Candolle ex Saint-Amans

Polystigma de Candolle ex Saint-Amans, Flore Agenaise:516 (1821).

S t r o m a t a soft t e x t u r e d , occupying slightly swollen portions of leaf


blades, brightly coloured at first, parasitic, perithecia crowded, globose
to pear-shaped, with compact colourless walls and well developed
ostioles, asci 8-spored, ascospores hyaline, non-septate. The British
species are parasitic on leaves of Rosaceae b u t exotic species a t t a c k
Leguminosae also.

259
Polystigma rubrum (Persoon) Saint-Amans, op. cit.:516 (1821). (PI.
XXVIIE.)

S t r o m a t a a few millimetres across, slightly thicker t h a n the un-


affected portions of t h e leaf blade, slightly arched, smooth, reddish-
orange, convex to t h e lower leaf surface, perithecia numerous, a b o u t
0.2 m m . across, with papillate ostioles; asci clavate, up to 9 0 x 1 0 / / ,
8-spored; ascospores elliptical, 11-13x4-5,«, hyaline, non-septate;
paraphyses thread-like. Only pycnidia containing slender curved
pycnospores can be found in t h e brightly coloured stromata on t h e
living leaves. On fallen leaves the s t r o m a t a darken, becoming almost
black, and in these the perithecia m a t u r e , ejecting their ripe ascospores
in spring.

On leaves of Prunus. Common in England on Prunus spinosa, especially


near the sea but in central and south-eastern Europe the fungus causes a
serious disease of the domestic plum.

Polystigma julvum de Candolle ex Saint-Amans, op. cit.:516 (1821),


causes similar b u t at first golden-yellow s t r o m a t a on leaves of Prunus
padus in northern England and Scotland.

ISOTHEA Fries

Isothea Fries, S u m m a Veg. Scand. Sect. P o s t . : 4 2 1 (1849).


S t r o m a t a embedded in leaf tissue, small, containing usually a single
perithecium which is subglobose with ostiole minute and not protrud-
ing, asci 8-spored, ascospores hyaline, non-septate. There is only one
British species:

Isothea rhytismoidcs (Babington ex Berkeley) Fries, op. cit.:421


(1849). (Fig. 18F.)
Perithecium subglobose, about 0.25 m m . across, embedded in leaf
tissue and surrounded b y a small stroma the upper surface of which
forms a black shining clypeus like a t i n y black pimple on the upper
surface of the leaf. Asci clavate, up to 60x15/«, 8-spored; ascospores
irregularly biseriate, elliptical, 1 0 - 1 5 x 5 - 7 / / , hyaline, non-septate;
paraphyses numerous, slender.

On living and dead leaves of Dryas octopetala.

260
PHYLLACHORA Nitschke

Phyllachora Nitschke apud Fuckel in J a h r b . Nass. Vereins f. Natur-


k u n d e 23-24:216 (1870).
S t r o m a t a embedded in leaf tissue, commonly with loose white flesh
b u t a black outer crust, perithecia subglobose, with well developed
ostioles, asci usually with eight uniseriate spores, ascospores hyaline
to yellowish, non-septate.

Phyllachora graminis (Persoon ex Fries) Fuckel, op. cit.: 216 (1870).


(Fig. 1 5 J . )
S t r o m a t a scattered, elongated elliptical, white-fleshed with a shining
black crust occupying the upper and lower epidermis of the leaf, com-
monly about 2 - 3 x 1 mm., sometimes larger. Perithecia numerous,
crowded, 0.2-0.3 m m . across, with minutely papillate ostioles; asci
cylindrical, almost sessile, up to 90x10/«, 8-spored; ascospores uni-
seriate, elliptical, 7 - 1 5 x 4 - 7 / « , hyaline, non-septate; paraphyses
numerous, slender.

On living and dead leaves of various grasses, especially Agropyron repens,


Bromus ramosus and Dactylis glomerata, in A u t u m n and winter. Very com-
mon.

Phyllachora sylvatica Saccardo & Spegazzini in Michelia 1:410 (No-


vember 1878), occurs on leaves of Festuca rubra and F. ovina with
ascospores 15-22x7-8/«.
Young stromata of Phyllachora contain pycnidia with thread-like,
hyaline pycnospores.

ENDODOTHELLA Theissen& Sydow

Endodothella Theissen & Sydow in Annales mycologici 13:582 (1915).


Like Phyllachora b u t with t h e ascospores ultimately appearing
1-septate. There is only one British species:

Endodothella junci (Fries) Theissen & Sydow, op. cit.:586 (1915). (PI.
Fig. 2 2 H . )
S t r o m a t a gregarious, embedded in the host tissue, scarcely 1 m m .
long, with a black crust and yellowish flesh enclosing minute perithecia
1
/g m m . across. Asci cylindrical, short-stalked, up to 85x8/«, 8-spored;
ascospores uniseriate, elliptical, 10-15x3-3.5/«, with two large oil

261
bodies, ultimately appearing obscurely 1-septate b u t not constricted
at the septum.
On dead leaves of the larger species of Juncus. Very common.
Though the ascospores are nominally 1-septate the septum is usually
rather vague and hard to see and the species is better referred to Phylla-
chora, as P. junci (Fries) Fuckel in Jahrb. Nass. Vereins f. Naturkunde
28-24:216 (1870).

GIBELLINA Passerini

Gibellina Passerini in Revue Mycologique 8:177 (October 1886).


Perithecia associated with a mycelial felt, gregarious, asci 8-
spored, ascospores yellowish-brown, 1-septate. There is only one
species:

Gibellina cerealis Passerini, op. cit.: 177 (1886). (PI. XXXVIIG.)

Perithecia gregarious in the lower leaf sheaths of Triticum, as-


sociated with and partially embedded in a greyish mycelial felt which
permeates the host tissue and also forms a stromatic layer in the inter-
spaces between the leaf sheaths. Perithecia up to about y 2 m m .
across, light coloured below b u t with black erumpent beaks. Asci
cylindrical, up to 125 x 18//, wall slightly thickened at the tip, 8-spored;
ascospores biseriate, narrowly elliptic-fusiform, 2 3 - 3 6 x 7 - 1 1 //, 1-sep-
t a t e , scarcely constricted, yellowish-brown at m a t u r i t y .

On dark-edged lesions at the base of Triticum culms, from June onwards.


Rare.

Family 6. Melanosporaceae

Perithecia superficial, light or dark coloured, often beaked or hairy,


with an apical ostiole; asci cylindrical or clavate, breaking down to
liberate the spores within the perithecium, ascospores non-septate,
brown or olive-brown, extruded in masses from the ostiole under d a m p
conditions. Species of Neurospora somewhat resemble Sphaeroderma
b u t their asci are not diffluent, see p. 267.

I. Perithecia smooth or tomentose, bristles round the ostiole minute:


A. Perithecia beaked Melanospora
B. Perithecia not beaked Sphaeroderma (p. 264)
II. Perithecia crowned by a conspicuous tuft of hairs, often as large as the
body of the perithecium, hairs often forked or spirally coiled:

262
A. Perithecia spherical, ascospores commonly lemon-shaped
Chaetomium (p. 265)
B. Perithecia pear-shaped, ascospores often discoid Ascotricha (p. 266)
For similar fungi with hyaline ascospores see Sphaeronaemella p. 335.

MELANOSPORA Corda

Melanospora Corda, Icones F u n g o r u m 1 : 2 4 (1837).


Perithecia scattered, superficial, subglobose with a long slender beak
terminated by the ostiole which is surrounded b y a ring of short hyaline
bristles. Occasionally the beak m a y be greatly reduced and the genus
is then to be separated from Sphaeroderma b y the presence of the
ostiolar bristles. Asci usually 8-spored, soon disappearing, ascospores
non-septate, dark brown or black. A full account of the British species
will be found in Petch (1938) and a more comprehensive monograph
b y Doguet in Le Botaniste 39 (1955).

Melanospora chionca (Fries) Corda, Icones F u n g o r u m 1 : 2 4 (1837).


(PI. X X X V i . )
Perithecia gregarious, superficial, subspherical, minutely tomentose,
light yellowish-brown, up to 0.5 m m . across with a slender beak. Asci
clavate, long stalked, about 70 x 15//, 8-spored; ascospores biseriate,
oval, discoid, at first hyaline, soon becoming dark brown and t h e n
liberated into the perithecial cavity b y dissolution of the ascus,
10-16 x 8 - 1 2 x 4 - 6 fx.
On fallen needles of Pinus, in spring and autumn. Uncommon.
The figure shows an immature ascus on the right and two views of a
mature ascospore above.

Melanospora zamiae Corda, op. cit.:24 (1837), common on rotting


vegetation, has lemon-shaped ascospores 15-23x11-16/«.

Melanospora parasitica Tulasne, Selecta F u n g o r u m Carpologia 3 : 1 0


(1865). (PI. X X X V J . )
Perithecia scattered, base somewhat sunken among the loose h y p h a e
of the host, globose, about 0.2 m m . across, beak very long and slender,
brown throughout. Asci clavate, 8-spored, about 3 0 x 5 / / ; ascospores
elliptic-cylindric with t r u n c a t e d ends, 5-8x2-2.5/<, non-septate, dark
brown to black. The asci soon disappear and the ascospores become
visible as a dark mass through the translucent perithecial wall, filling

263
the cavity and t h e canal in the beak and emerging as a blackish glo-
bule from the ostiole.

Parasitic on the entomogenous moulds Isaria farinosa and Beauveria


bassiana. Not common.

Melanospora lagenaria (Persoon ex Fries) Fuckel, in J a h r b . Nass.


Vereins f. N a t u r k u n d e 23-24:126 (1870), with slightly apiculate
elliptical ascospores 13-22 x 7 - 1 4 / t occurs on decaying fructifications
of Polyporus, Polystictus and Stereum.
Melanospora brevirostris (Fuckel) von Hohnel in S.B. K. Akad. Wiss.
Wien, Math.-Nat. Kl. A b t . I, 123:94 (1914), parasitises the disc of
species of Sepultaria.

SPHAERODERMA Fuckel

Sphaerodcrma Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 29-30:22


(1875).
Perithecia scattered, superficial, more or less globose, soft-fleshed,
without a beak and without bristles round the ostiole, asci 8-spored,
soon disappearing, ascospores non-septate, dark brown, sometimes
ornamented by lines. Doguet regards t h e absence of a beak as of little
taxonomic importance and unites the genus with Melanospora.

Sphaeroderma episphacria (Phillips & Plowright) Saccardo, Sylloge


F u n g o r u m 2:460 (1883). (PI. X X X I i . )
Perithecia gregarious, superficial, globose with an apical papilla
t r u n c a t e d b y the ostiole, colourless, drying brown or appearing black
when moist when the enclosed spore mass becomes visible through t h e
translucent wall, about 0.3 m m . across. Asci cylindric-clavate, 8-spor-
ed, soon disappearing; ascospores lemon-shaped, 2 5 - 3 4 x 1 2 - 1 8 / / , soon
becoming dark brown and ornamented b y a wide-meshed network of
lines.

On the hyphal mat of Hypomyces ochraceus, see p. 239, in autumn. Un-


common.

Sphaeroderma fusisporum P e t c h in The Naturalist: 58 (1936), occurs


on Isaria farinosa, distinguished from Melanospora parasitica on t h e
same host not only b y lacking the beak b u t also b y its very much
bigger ascospores, 2 0 - 2 4 x 6 - 9 / / .

264
CHAETOMIUM Kunze ex Fries

Chaetomium Kunze ex Fries, Systema mycologicum 3:253 (1829).

Perithecia superficial, spherical and translucent when young, sub-


globose, opaque and dark coloured at m a t u r i t y , bearing bristle-like
appendages which are most numerous round the ostiole where t h e y
usually form a t u f t about as large as the b o d y of the perithecium. The
shape of t h e bristles is usually highly characteristic and one of t h e best
diagnostic characters of the species. Asci clavate, thin-walled, 8-spored,
soon breaking down and disappearing, ascospores non-septate, olive-
brown, usually lemon-shaped. The species are saprophytes on dung
and decaying vegetable m a t t e r of all kinds. The most convenient
monograph is t h a t of Ames in U.S. A r m y Res. Dev. Ser. 2 (1963).
There m a y be about a dozen British species one of the commonest of
which is:

Chaetomium elatum Kunze & Schmidt ex Fries, op. cit.: 254 (1829).
(Fig. 17D.)

Perithecia shortly ovoid, superficial, scattered over a loose m a t of


dark olive-brown hyphae, up to about 0.5 m m . high and not quite as
wide, bearing numerous stiff h y p h a l appendages, those on t h e basal
p a r t spreading, unbranched and bristle-like, those on the upper p a r t
of the perithecium repeatedly dichotomously branched, their branches
hence interlocked to form a kind of loosely meshed cage above the
ostiole. The walls of the appendages are olive-brown or almost black
and coarsely roughened, only the tips m a y be almost hyaline. Asci
broadly clavate, up to 75 x 17//, 8-spored, soon disappearing; ascospores
lemon-shaped, 1 2 - 1 3 x 8 . 5 - 9 . 5 / / , somewhat flattened in one plane,
d a r k olive-brown.

On decaying vegetable matter of all kinds and on dung, throughout the


year. Very common.

In the equally common and ubiquitous Chaetomium globosum Kunze


ex Fries, op. cit.: 255 (1829), the apical hairs are undulating and loosely
intertwined b u t not dichotomously branched. In some species, such as
C. cochliodes Palliser in N. Amer. Flora 3 : 6 1 (1910), C. bostrychodes
Zopf, in Abh. Bot. Ver. Prov. B r a n d e n b u r g 19:173 (1877), C. spirale
Zopf in Nova Acta Leop. Carol A k a d . 42:275 (1881), the appendages
form beautiful and characteristic spiral coils.

265
ASCOTRICHA Berkeley

Ascotricha Berkeley in Ann. Mag. n a t . Hist. 1 : 2 5 7 (1838).


This genus is very closely allied to Chaetomium and is united with
it b y some authors. It is retained here for species in which the peri-
thecium has a distinct narrow neck. The only British species is:

Ascotricha chartarum Berkeley, op. cit.:257 (1838). (Fig. 17B.)


Perithecia scattered, superficial, globose, about 0.2 mm. across,
with a distinct apical neck, black t h r o u g h o u t ; hairs stiff, blackish,
straight, branched, b u t not dichotomously, variable in length, those
on t h e lower p a r t of the perithecium spreading, those round the ostiole
forming a tall open cage within which the spores are extruded. Asci
cylindric-clavate, 6 5 x 1 1 / / , 8-spored; ascospores uniseriate, soon
liberated from the asci, discoid, circular in face view and about 8/«
across, 4.5-5.5/« thick, olive-brown to black.

On rotting paper and cardboard. Not common.

Family 7. Lasiosphaeriaceae

This family is not sharply differentiated from the Xylariaceae,


especially as hyaline appendages occur on the ascospores of some
species of Rosellinia. The following genera have been assigned to it:
I. Perithecia produced on dung:
A. Ascospores without an appendage:
1. Ascospore marked by a lateral furrow or spfit in the dark coat
Coniochaeta (p. 273)
2. Ascospore without a furrow Sordaria (p. 268 )
B. Ascospore with one or more hyaline appendages:
1. Ascospore at first worm-like . . . . Lasiosordaria (p. 270)
2. Ascospore at first club-shaped . . . . Podospora (p. 269)
If. Perithecia produced on other substrata:
A. Parasitic on Basidiomycetes . . . . Helminthosphaeria (p. 272)
B. Saprophytic on wood, bark and vegetable debris, or parasitic on
lichens:
1. Ascospores at first hyaline and greatly elongated:
a. Mature ascospore without a swollen dark-brown cell
Lasiosphaeria (p. 271)
b. Mature ascospore differentiated into an elliptical dark-
brown cell and a cylindrical hyaline appendage
Bombardia (p. 271)
2. Ascospores more or less elliptical, dark brown:
a. Ascospores unornamented, cohering in groups in the ascus
Synaptospora (p. 274)

266
ft. Ascospores not so cohering:
a. Ascospores smooth, with a lateral furrow
Coniochaeta (p. 273)
b. Ascospores dotted with hyaline pits in the wall
Gelasinospora
c. Ascospores ornamented with lines Neurospora

NEUROSPORA Shear & Dodge

Neurospora Shear & Dodge in J o u r n a l of Agricultural Research 34:


1025 (1927).
Perithecia more or less scattered, superficial or surrounded by a
loose hyphal weft, flask-shaped, dark coloured, smooth, asci 4- or
8-spored, ascospores elliptic-fusiform, non-septate, dark brown,
ornamented with fine longitudinal ribs. Saprophytes on vegetable
m a t t e r , often after it has been charred by fire. For a full account of t h e
genus see Journal of Agricultural Research 34:1019-1042 (1927).

Neurospora sitophila Shear & Dodge, op. cit.: 1026 (1927). (Fig. 10i>.)
Perithecia flask-shaped with an apical papilla, up to 0.3 m m . across,
brown to black, smooth or downy with loose h y p h a e . Asci cylindrical,
short-stalked, up to 1 6 0 x 1 4 t h i n - w a l l e d with a ring-like thickening
at the tip, 8-spored; ascospores uniseriate, elliptic-fusiform, 20-26x
10-15//, olivaceous, nearly black at m a t u r i t y , ornamented with 16 or
17 somewhat anastomosing, fine, longitudinal ribs; paraphyses a b s e n t
from the ripe perithecia.
The conidial state : s the salmon-coloured mould Monilia sitophila Mon-
tagne, not uncommon on charred wood and grain but the perithecia normally
develop only in artificial culture. They are, therefore, unlikely to be met
with by the field mycologist but the genus is worth mentioning here because
it has been the subject of extensive genetical studies, especially by North
American workers, and has hence become familiar by name to many stu-
dents who have never seen it in the flesh.

Neurospora tetrasperma Shear & Dodge, op. cit.: 1027 (1927), has
4-spored asci, as have two additional species not yet recognised in
Britain.

G E L A S I N O S P O R A Dowding

Gelasinospora Dowding in Canadian J o u r n a l of Research C9:294


(1933).
Perithecia very similar to those of Neurospora, asci 4- or 8-spored,
ascospores non-septate, more or less elliptical, with a dark brown outer

267
coat which is perforated b y numerous small round pits, exposing t h e
hyaline inner wall. The pits are best seen in crushed and broken
ascospores.

Gelasinospora cerealis Dowding, op. cit.:295 (1933). (Fig. 12M.)


Perithecia scattered, subglobose with a small apical papilla, dark-
brown to black, smooth, a b o u t 0.4 m m . across. Asci cylindrical, up to
2 6 0 x 3 3 / / , 8-spored; ascospores uniseriate, broadly elliptical, almost
black at m a t u r i t y b u t dotted with small round hyaline pits, 26-32 x
23-25//.
As in Neurospora the perithecia are met with in artificial culture, isolated
from soil, plant remains and dung, but in Gelasinospora there is no mould
stage in the life history.

Gelasinospora tetrasperma Dowding, op. cit.: 294 (1933), differs from


the above in its 4-spored asci.

SORDARIA Cesati & de Notaris

Sordaria Cesati & de Notaris in Comm. Soc. Crittogamologica italiana


1:225 (1863).
Perithecia pear-shaped, d a r k coloured, more or less sunk in dung,
smooth, at least at t h e tip, asci cylindrical with a ring-like thickening
of the wall at t h e apex, usually 8-spored, ascospores subglobose to
elliptical, non-septate, almost black and opaque at m a t u r i t y , with a
germ-pore at the lower end of t h e spore as it lies in t h e ascus. For a
modern revision of the genus see C. Moreau, Les genres Sordaria et
Pleurage. Encyclopedic mycologique 25 (1953).

Sordaria humana (Fuckel) Winter, Abhandl. d. Naturf. Ges. Halle 13


(1):24 (1873). (Fig. 12i.)
Perithecia pear-shaped, scattered with t h e base more or less sunken
in the substrate, smooth above, often with a few anchoring h y p h a e
below, black, u p to 0.4 m m . across. Asci cylindrical, up to 2 0 0 x 1 9 / / ,
8-spored; ascospores uniseriate, broadly obovoid with a small papilla
at the lower end, 2 0 - 2 5 x 1 3 - 1 7 / / , black, smooth, enclosed in a narrow,
colourless, gelatinous sheath; paraphyses absent.
On dung of man and animals. Not uncommon.
Sordaria fimicola (Roberge) Cesati & de Notaris, op. cit.: 276 (1863),
is very similar b u t has elliptical ascospores, 1 9 - 2 5 x 1 0 - 1 3 / / .

268
Sordaria macrospora Auerswald in Hedwigia 5 : 1 9 2 (1866), is dis-
tinguished by its larger ascospores 2 5 - 2 8 x 1 7 - 2 0 / / .

PODOSPORA Gesati

Podospora Cesati in Hedwigia 1 : 1 0 3 (1856).


Perithecia scattered, more or less sunken in the substrate, pear-
shaped or subconical, smooth, hairy or scaly, d a r k coloured, asci
cylindrical, clavate or fusiform, containing from 4 to 512 ascospores,
spores at first club-shaped and hyaline, the swollen upper p a r t be-
coming dark olive-brown whilst t h e lower p a r t remains hyaline and
e m p t y , forming the p r i m a r y appendage. Secondary appendages of
various shapes and with a fibrillose structure m a y develop at each end
of the ascospore. The species are v e r y numerous, Moreau, who calls
the genus Pleurage Fries, offers a key to 17 studied b y him in F r a n c e .

Podospora curvula (de Bary) Niessl in Hedwigia 22:156 (1883). (Fig.


12K.)

Perithecia scattered, pear-shaped, r a t h e r elongated, up to 1 m m .


tall and 0.4 m m . wide, dark brown to blackish, ornamented especially
round the base of the neck b y short septate hairs which commonly
cling together in clusters to form small somewhat triangular scales.
Asci clavate, up to 2 2 0 x 3 0 / / , 8-spored; ascospores biseriate, elliptical,
23-33x13-16/«, blackish and opaque, with a slender, cylindrical, hya-
line basal appendage 1 0 - 2 0 x 2 - 3 / / plus a long tapering secondary
appendage at each end of the spore. There m a y also be one or more
short, slender, secondary appendages a t t a c h e d to the side of the
primary appendage.

On dung of various kinds. Common.

Podospora minuta (Fuckel) Niessl in Hedwigia 22:156 (1883), is very


similar b u t with smaller perithecia and ascospores 1 7 - 2 3 x 1 1 - 1 5 / / ;
there is also a form with 4-spored asci.
In Podospora anserina (Cesati) Niessl in Hedwigia 22:156 (1883),
with similar b u t very large ascospores, 3 4 - 4 0 x 18-20 [/, only 4-spored
asci are known.
Podospora setosa (Winter) Niessl in Hedwigia 22:156 (1883), has
bristly perithecia with polysporous asci. Moreau recognises races with
32-, 64-, 128- and 512-spored asci, the individual spores 1 5 - 2 0 x 9 - 1 2 / « ;

269
P. curvicola (Winter) Niessl in Hedwigia 22:156 (1883), differs in
having scales like P. curvula instead of spreading bristles.
Podospora decipiens (Winter) Niessl in Iledwigia 22:156 (1883), is
distinguished by having secondary appendages attached in a ring near
the top of the p r i m a r y appendage; according to Moreau it includes
races with 8-, 16-, 32- and 64-spored asci.
Podospora fimiseda (Cesati& de Notaris) Niessl in Hedwigia 22:156
(1883), has 8-spored asci with very large ascospores 5 0 - 6 0 x 2 8 - 3 2 / / .
Podospora appendiculata (Auerswald) Niessl in Hedwigia 22:156
(1883), has ascospores 2 5 - 3 2 x 1 3 - 1 7 / i , more fusiform t h a n those of
S. curvula, and its perithecia are sparsely hairy, not scaly.
Podospora neglecta (Hansen) Niessl in Hedwigia 22:156 (1883), with
ascospores 4 3 - 5 2 x 2 2 - 2 8 / / , is peculiar in having no p r i m a r y append-
age, though there is a long secondary appendage at each end of the
spore.

LASIOSORDARIA Chenantais

Lasiosordaria Chenantais in Bull. Soc. mycol. France 35:77 (1919).


Fructifications like those of Podospora except t h a t the ascospores
are at first cylindrical, not club-shaped. These spore characters are
those of Bombardia, with which Lasiosordaria is accordingly united b y
Moreau and other authors.

Lasiosordaria coprophila (Fries) Chenantais, op. cit.:78 (1919). (PI.


XXXVc.)
Perithecia r a t h e r elongated pear-shaped, neck short, base slightly
sunken in the substrate, about 0.5 m m . wide and up to 1 m m . tall,
dark brown to black, covered with a white t o m e n t u m formed of
slender thin-walled septate hairs. Asci cylindrical, long-stalked, with
a well marked apical ring, up to 250x20//, 8-spored; ascospores bi-
seriate, at first cylindrical, somewhat undulating, hyaline, containing
from four to eight large oil drops, bearing a small, pointed, easily over-
looked appendage at each end, m a t u r e ascospore with an elliptical dark
brown body 1 7 - 3 0 x 9 - 1 2 / / and a long cylindrical, hyaline basal
primary appendage. The secondary appendages soon disappear.

On dung. Fairly common.

Lasiosordaria lignicola (Fuckel) Chenantais op. cit.:78 (1919), is a


similar species with the perithecia immersed in very rotten wood.

270
BOMBARDIA Fries
Bombardia Fries, S u m m a Veg. Scand. Sect. Post.:389 (1849).
As indicated above this genus is doubtfully distinct from the pre-
ceding. It is separated b y Munk on account of the structure of the
perithecial wall which is made up of two thin brown layers with a
thick hyaline tissue of interwoven h y p h a e between t h e m . In this
restricted sense it includes only one British species:

Bombardia fasciculata Fries, op. cit.:389 (1849). (Fig. 12jr.)


Perithecia superficial, occurring in dense clusters seated on a com-
mon base, rather narrowly club-shaped or bomb-shaped, shining
black, up to 1.5 m m . tall and about 0.6 m m . wide. Asci cylindrical,
long-stalked, up to 160x12//, 8-spored; ascospores biseriate, at first
cylindrical, curved or undulating, hyaline, about 40x4/<, with a
minute pointed appendage at each end, finally with a dark brown
elliptical body 1 2 - 1 4 x 8 / / in the upper p a r t and a curved, cylindrical,
hyaline primary appendage below; paraphyses numerous, hyaline.

On dead wood, in Autumn. Uncommon.

LASIOSPHAERIA Cesati & de Notaris

Lasiosphaeria Cesati & de Notaris in Comm. Soc. Crittogamologica


italiana 1:229 (1863).
Perithecia superficial, gregarious, on wood or bark, more or less
globose with a papillate ostiole, black, smooth or hairy, asci 8-spored,
ascospores cylindrical, often sharply bent or undulating, hyaline or
ultimately light brown, septate or not, sometimes slightly enlarged at
the upper end b u t never forming a separate dark brown spore b o d y
there. ~~

Lasiosphaeria spermoides (Hoffmann ex Fries) Cesati & de Notaris,


op. cit:229 (1863). (Fig. 12L.)
Perithecia superficial, densely crowded, often forming large patches,
individually subglobose, about 0.6 m m . across, rounded above with a
scarcely perceptible papillate ostiole, black, glistening, hairless. Asci
cylindric-clavate, long-stalked, u p to 11// wide, 8-spored; ascospores
cylindrical, usually bent near the middle like a sausage, with rounded
ends, 1 9 - 2 2 x 3 - 4 / / , hyaline, non-septate b u t usually with a clear space

271
in the middle separating two large oil drops or aggregations of granules ;
paraphyses numerous.
On logs, especially of Fagus, in late autumn and early spring. Common.

Lasiosphaeria hirsuta (Fries) Cesati & de Notaris, op. cit.:229 (1863).


(Fig. 12N.)
Perithecia superficial, gregarious, associated with brown hyphae,
globose, about 0.5 m m . across, black, covered with short, spreading,
blackish hairs. Asci cylindric-clavate, stalked, up to 160x14//, 8-spor-
ed; ascospores biseriate, cylindrical b u t usually strongly curved,
rounded at each end, 55-65x6-7/«, hyaline to yellowish, with five to
seven septa and slightly constricted at the septa; paraphyses nu-
merous.
On logs, of Fagus, Salix and other trees, in spring and autumn. Common.

Lasiosphaeria ovina (Fries) Cesati & de Notaris, op. cit.:229 (1863),


is easily recognised b y its coating of pure white hairs.
Descriptions of other British species are easily accessible in W i n t e r
(1887).

HELMINTHOSPHAERIA Fuckel

Helminthosphaeria Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 2 3 -


24:166 (1870).
Perithecia superficial, scattered, subglobose, hairy, parasitic on
species of Clavariaceae and Thelephoraceae, asci 8-spored, ascospores
non-septate, dark brown. This and t h e following genera form a transi-
tion to Xylariaceae, from which t h e y are separated b y their super-
ficial, scattered, hairy perithecia. B o t h are united with Rosellinia b y
some authors.

Helminthosphaeria clavariarum (Tulasne) Fuckel, op. cit.: 166 (1870).


(Fig. 12G.)
Perithecia scattered over a web of dark brown hyphae, superficial or
somewhat embedded in the m a t , subglobose, about 0.3 m m . across,
without a distinct papilla, black, covered with spreading hairs, some
tapering to a point, others cylindrical and bearing elliptical, 1-septate,
brown conidia. Asci cylindrical, stalked, up to 90x8/«, 8-spored; asco-
spores uniseriate, broadly fusiform or one-sided, 10-14x6-7/«, dark
brown, non-septate; paraphyses numerous.

272
On fructifications of Clavariacinerea and C. cristata, in autumn. Uncommon.
The conidial state, which may be found without the perithecia, has been
called Spadicoides clavariarum, (Desmazieres) Hughes in Canad. Journ. Bot.
36:806 (1958) illustrated above the ascospore.

Helminthosphaeria corticiorum von Hohnel in S.B. K. Akad. Wiss.


Wien, Math.-Nat. Kl. A b t . 1 , 1 1 6 : 1 0 9 (1907), occurs on fructifications
of Peniophora.

CONIOCHAETA (Saccardo) Massee

Coniochaeta (Saccardo) Massee in Grevillea 16:37 (December 1887).


Perithecia similar to those of Helminthosphaeria b u t not bearing
Scolecotrichum conidiophores, hairy, at least on t h e upper surface,
ascospores dark brown and non-septate b u t bearing a fine crack or
furrow in the dark coat along one side of t h e spore, asci 8- to m a n y -
spored. Several species occur on dung, others on wood.

Coniochaeta ligniaria (Greville) Massee in Grevillea 16:37 (December


1887). (Fig. 12E.)
Perithecia gregarious or scattered, superficial or with the base sunk
in the substrate, subglobose, slightly pointed above, about 0.3 m m .
across, blackish-brown, covered with spreading, dark brown, pointed
hairs up to 5 0 x 4 f i . Asci cylindric-clavate, up to 130x14//, 8-spored;
ascospores uniseriate, broadly ellipsoidal b u t flattened, mostly 12-15x
8 - 1 2 x 4—6 //, nearly black, with a distinct furrow along one narrow side,
enclosed in a thin, colourless, gelatinous sheath; paraphyses numerous.

On dead wood, in spring. Not common on wood but according to Munk


C. discospora (Auerswald) Gain in Univ. Toronto Biological Studies 38:62
(1934), is the same species growing on dung, where it is not uncommon.

Coniochaeta velutina (Fuckel) Munk, in Dansk. Bot. Arkiv. 12:11


(1948), with ascospores 7 - 8 x 5 - 5 . 5 / t occurs on logs.
Coniochaeta malacotricha (Auerswald) Traverso in Flora Italica
Cryptogama 1 (2):473 (1907), with ascospores 10-12x8-10/u, occurs
on coniferous logs.
Coniochaeta pulveracea (Ehrenberg ex Fries) Munk, op. cit. :9 (1948),
has smooth perithecia, with ascospores 1 0 - 1 2 x 7 - 8 / / , on logs of deci-
duous trees.
Coniochaeta leucoplaca (Berkeley & Ravenel) Cain with discoid asco-
spores 9 x 7 x 4 / t and Coniochaeta scatigena (Berkeley & Broome) Cain,

273
with ovoid to discoid ascospores 1 7 - 2 3 x 1 0 - 1 9 / / or 17x17//, occur
on dung.

SYNAPTOSPORA Cain

Synaptospora Cain in Beihefte zur Sydowia 1 : 4 (1957).


Perithecia superficial or emergent from a lichen thallus, globose,
black, with an irregularly thickened wall. Asci narrowly cylindrical,
thin-walled, with or without a distinct apical ring; ascospores uni-
cellular, dark-brown, cohering in groups of two to four.

Synaptospora tartaricola (Nylander apud Leighton) Cain, op. cit.:6


(1957). (Fig. 30G.)
Perithecia globose, without a distinct ostiole, with thick, rough,
black wall. Asci 120x10-11 //, without an apical ring; ascospores uni-
seriate, at first elliptical, hyaline, 1 0 - 1 2 x 8 - 9 . 5 / / , becoming dark brown
and cohering in groups of from 2 to 5 spores, sometimes eventually
becoming unevenly 1-septate.
On thallus of Ochrolechia lartarea. August. Rare.
The ascospores do not all cohere and one might dismiss this feature as
an abnormality had not Gain recorded it in a second species, S. petrakii Cain,
saprophytic on wood in North America.

Family 8. Xylariaceae

The sphaeriaceous genera with dark coloured non-septate ascospores


produced in smooth perithecia embedded in a stroma, developed
beneath a clypeus or on a subiculum m a y be distinguished as follows:

I. Perithecia immersed in a massive stroma or seated on a cord-like rhizo-


morph. (The stroma is greatly reduced in Hypoxylon semi-immersurn):
A. Stroma stalked, club-shaped, strap-like or thread-like, often forked:
1. Stroma finally black, perithecia not confined to the tip
Xylosphaera (p. 284)
2. Stroma whitish, perithecia inserted in a flat apical disc
Poronia (p. 286)
3. Stroma brown or black, with a small apical cluster of peri-
thecia on a stalk rooted in dung Podosordaria (p. 287)
B. Stroma sessile:
1. Stroma hemispherical:
a. Flesh of the stroma concentrically zoned Daldinia (p. 283)
b. Flesh of the stroma not zoned Hypoxylon (p. 279)

274
2. Stroma discoid, irregularly cushion-shaped or a spreading
crust:
a. Perithecia narrowly cylindrical, closely crowded and
more or less hexagonal in cross-section
Camarops (p. 282)
b. Perithecia flask-shaped or subglobose:
* Ascospores over 25 p long . . . Ustulina (p. 282)
**Ascospores seldom over 20,« long:
fPerithecia seated immediately beneath the crust
of the stroma or protruding from it, hence
with the ostioles at most papillate
Hypoxylon (p. 279)
tjPerithecia seated in the base of the stroma with
long tubular necks:
§§Perithecia in a single stratum
Nummularia (p. 283)
§Perithecia at different levels in the stroma
Bolinia (p. 282)
II. Perithecia superficial, free, or immersed in dung or plant tissue, with a
subiculum or beneath a clypeus or clustered in a rudimentary stroma
only:
A. Immersed in dung Hypocopra (p. 278)
B. Superficial, often with a subiculum of dark hyphae, on wood
Rosellinia (p. 285)
C. Embedded in plant tissue, or on burnt ground:
1. Perithecia solitary or a few together beneath a clypeus:
a. Ascospores without equatorial germpores:
*Ascospores with a small rounded hyaline basal
appendage Entosordaria
**Ascospores without such an appendage:
Perithecia in herbaceous stems and leaves, not
beaked Anthostomella (p. 276)
Perithec ia in rotten wood and bark, long-beaked
Anthostoma (p. 277)
b. Ascospores with four equatorial germpores
Amphisphaerella (p. 277)
2. Perithecia in small valsoid clusters in bark
Lopadostoma (p. 278)
See also Clypeosphaeria, p. 295.

ENTOSORDARIA (Saccardo) von Hohnel

Entosordaria (Saccardo) von Hohnel in Sitzber. Akad. Wiss. Wien,


Math.-Nat. Kl. 129, A b t . 1:167 (1920).
Perithecia immersed, with papillate ostiole, ascospores uniseriate,
with a large brown cell and a minute hyaline, rounded, basal append-
age, ascus with an apical ring.

275
Entosordaria tomicoides (Saccardo) von Hohnel, op. cit.: 166 (1920).
(PI. X X X V A . )
Perithecia less t h a n 0.5 m m . across, black, immersed in dead stems
with the papillate ostioles exposed, each surrounded b y a small p a t c h
of dark brown hyphae. Asci cylindrical, with a distinct apical ring, up
to 110x10//, 8-spored; ascospores uniseriate, fusiform, flattened on
one side, 1 2 - 1 7 x 5 - 8 / / , dark brown, with a minute, rounded, colourless
papilla at the lower end. There is usually a longitudinal furrow which
appears as a colourless line down one side of the spore but this is not
always easy to see.
On dead stems of Eupatorium cannabinum, in spring and summer. Not
uncommon.

Entosordaria ammophilae (Phillips & Plowright) von Hohnel, op. cit.:


166 (1920). (Fig. 25x.)
Perithecia scattered, immersed in the host, mostly solitary beneath
an elliptical clypeus of grey epidermis, less often in groups of two or
three. Asci cylindrical, 8 0 x 8 - 1 0 / t , 8-spored; ascospores elliptical,
8 - 1 0 x 5 - 6 / / , with a minute basal appendage.
In Ammophila arenaria. Common.

Anthostomella lagabris (Roberge) Saccardo, Sylloge F u n g o r u m 1:


278 (1882), on the same host, also has appendaged ascospores 20-25x
8.5-10//, b u t apparently has not been transferred to Entosordaria.

ANTHOSTOMELLA Saccardo

Anthostomella Saccardo in Nuovo Giorno Botanico Italiano 8 : 1 2


(1878).
Perithecia minute, immersed in leaves and herbaceous stems,
scattered, usually with at least a trace of a b l a c k i s h d y g e u s around the
ostiole, ascospores as in Anthostoma, with which it is united b y von Arx
and Miiller.

Anthostomella punctulata (Roberge) Saccardo, Sylloge F u n g o r u m 1:


278 (1882). (Fig. 25Y.)
Perithecia scattered, 150-200// diameter, ostiole opening in the
centre of a small, black, discoid clypeus within the host epidermis.
Asci cylindrical, 4 5 - 6 0 x 6 - 7 / / , 8-spored; ascospores elliptical, 7 - 1 0 x
3-4//.
In dead leaves of Carex spp.

276
A. phaeosticta (Berkeley) Saccardo in Michelia 1:374 (1879) is a
synonym according to von Arx & Muller.
A. taxi Grove in Journ. Bot. 71:253 (1933), with ascospores 12-14x
7-10//, occurs in dead leaves of Taxus baccata.

Several other species have been described on other grasses and


sedges. The fungi called Sordaria carbonaria (Phillips & Plowright)
Saccardo, op. cit.:233 (1882), with ascospores 1 8 - 2 2 x 8 - 9 / / , and
S. ustorum Mouton in C.R. Bull. Soc. Roy. Bot. Beige 36 (2): 11 (1897),
with ascospores 17-19x 10-13//, b o t h found on b u r n t ground, evidently
belong here r a t h e r t h a n in Sordaria.

ANTHOSTOMA Nitschke

Anthostoma Nitschke, Pyrenomycetes Germanici: 110 (1867).


Perithecia gregarious in a stroma in rotten wood or bark, long-
beaked; ascospores elliptical, brown, without an appendage.

Anthostoma decipiens (De Candolle ex Fries) Nitschke, op. c i t . : I l l


(1867). (Fig. 25z.)
Perithecia densely crowded in a stroma with whitish flesh and thin
black rind, in bark, m m . diameter, with cylindrical neck emerg-
ing as a smooth, black, papillate ostiole. Asci very numerous, cylindri-
cal, 6 0 - 8 0 x 6 - 9 / / ; ascospores elliptic-cylindric, 5 - 1 0 x 3 - 5 / « .

Typically in bark of Carpinus, also in rotten wood of other deciduous trees.

AMPHISPHAERELLA (Saccardo) Kirschstein

Amphisphaerella (Saccardo) Kirschstein in Trans. Brit, mycol. Soc. 18:


306 (1934).
Perithecia like those of Anthostomella and surrounded b y a similar
clypeus b u t with ascospores broadly elliptical, dark brown, with equa-
torial germ pores. There is only one British species:

Amphisphaerella xylostei (Persoon ex Fries) Munk in Dansk. Bot.


Arkiv. 15 ( 2 ) : 8 9 ( 1 9 5 3 ) . (PL X X X V D . )
Perithecia less t h a n 1 m m . across, black, scattered, immersed in dead
twigs with the ostioles slightly protruding and surrounded b y a small
p a t c h of dark brown superficial hyphae. Asci cylindrical, up to 150x

277
15/«, without a distinct apical ring, 8-spored; ascospores uniseriate,
broadly elliptical, 15-24x10-14/«, dark brown with four clear spots
(germ-pores) arranged along the equatorial zone.
On dead branches of Lonicera periclymenum and its allies, August to May.
Not uncommon.

HYPOCOPRA Fries

Hypocopra Fries, S u m m a Veg. Scand. Sect. Post.:397 (1849).


Perithecia globose, r a t h e r large, immersed in dung, on the surface of
which a small, thin, black stroma develops around the ostiole. Asco-
spores large, dark coloured, non-septate, with a gelatinous coat.

Hypocopra merdaria Fries, op. cit.:397 (1849). (PI. X X X V n . )


Perithecia gregarious, globose, up to 1 m m . across, brown to black,
sunk in dung with the ostiole exposed and surrounded b y a thin super-
ficial stroma, which has a white flesh and thin blackish crust. The
clypei of two or more perithecia m a y become confluent. Asci cylindri-
cal, up to 250x30/«, with a strongly developed apical ring-like thicken-
ing which stains blue with iodine, 8-spored; ascospores uniseriate,
elliptical, 27-35x16-20/«, dark brown with a longitudinal colourless
furrow, surrounded b y a gelatinous coat which soon dissolves in water.
The figure shows an ascus tip stained with iodine.

On dung of rabbit, sheep, etc. Not uncommon.


Many other species have been described, on dung of various animals.

LOPADOSTOMA (Nitschke) Traverso

Lopadostoma (Nitschke) Traverso in Flora Italica Cryptogama 1 (2):


169 (1906).
Perithecia globose, immersed in plant tissue in small clusters whose
ostioles emerge together, embedded in a very r u d i m e n t a r y stroma b u t
without a clypeus, ascospores brown, non-septate.

Lopadostoma turgidum (Persoon ex Fries) Traverso, op. cit.: 170 (1906).


(Fig. 27c.)
Perithecia about 0.5 m m . across, black, in clusters of three to six
united b y a r u d i m e n t a r y black stroma, raising the bark in a large
hemispherical pimple the tip of which is pierced b y a small black disc

278
formed b y the cohering ostioles. Asci cylindrical, up to 110x8/t, with
a thickened ring at the apex, 8-spored; ascospores uniseriate, elliptic-
cylindric, 1 0 - 1 3 x 5 - 6 / / , dark brown, with a furrow down one side.

Occurring in swarms on fallen branches of Fagus. Common.

HYPOXYLON Bulliard ex Fries

Hypoxylon Bulliard ex Fries, Systema Mycologicum 3; Index alpha-


beticus:103 (1832), non Hypoxylon Jussieu ex Merat, Nouvelle Flore
de Paris 1:137 (1821).
Perithecia globose to slightly flask-shaped, clustered in a common
stroma which m a y have a black, brown, reddish or even whitish sur-
face and m a y v a r y in shape from a hemispherical cushion to a t h i n
spreading crust, on dead wood or b a r k ; ascospores elliptic-fusiform to
bean-shaped, dark brown non-septate, with a distinct colourless furrow
down one side. The ostioles m a y open flush with the surface of the
stroma, appearing only as m i n u t e pores, or m a y protrude above it as
small papillae and this is an i m p o r t a n t character in differentiating t h e
species.

Hypoxylon fragiforme (Persoon ex Fries) Kickx, Flore Cryptogamique


environs Louvain:116 (1835). ( P L X X X I I A . )
Stroma hemispherical, up to 1 cm. across, salmon-pink at first, be-
coming brick-red at m a t u r i t y , blackening with age, the surface
papillate with the protruding tips of the perithecia. Flesh hard, black
with a single layer of small slightly flask-shaped perithecia just beneath
the surface; the ostiolar necks do not p r o t r u d e above the crust of the
stroma. Asci cylindrical, with a small apical ring, about 150x8/«,
8-spored; ascospores uniseriate, subfusiform with one flattened side,
dark brown, 1 1 - 1 5 x 5 - 7 / / , with one to three oil drops.

On bark of dead or dying branches of Fagus, usually in swarms. Very


common.

Hypoxylon rutilum Tulasne, Selecta F u n g o r u m Carpologia 2 : 3 8


(1863), is a very similar species which also occurs on Fagus b u t has
protruding ostioles and smaller ascospores, 7 - 1 0 x 3 . 3 - 4 / / .
Hypoxylon howeianum Peck in 24th Rept. N.Y. State M u s e u m : 9 8
(1871), which occurs on various deciduous trees, is like H. fragiforme,
with umbilicate ostioles, b u t has small ascospores, 6 - 9 x 3 - 3 . 5 / t .

279
Hypoxylon cohaerens (Persoon ex Fries) Fries, S u m m a Veg. Scand.
Sect. P o s t . : 384 (1849), sometimes occurs with H. fragiforme, from
which it is distinguished b y its smaller mahogany-brown stromata, up
to 4 m m . across, papillate ostioles and ascospores mostly 9 - 1 0 x 4 - 5 //.

Hypoxylon rubiginosum (Persoon ex Fries) Fries, S u m m a Veg. Scand.


Sect. P o s t . : 3 8 4 ( 1 8 4 9 ) . (PI. X X X I I B . )
Stroma flattened, forming a t h i n crust on decorticated wood, vary-
ing in colour from bright brick-red to reddish or purplish-brown,
blackening somewhat with age, surface smooth, minutely dimpled b y
the ostioles which m a y appear as white dots under a lens. Flesh very
thin, black, brittle. Asci cylindrical, about 1 5 0 x 8 / / , 8-spored; ascospo-
res uniseriate, elliptical with one flattened side, 9 - 1 2 x 4 - 6 //, dark brown.
On dead wood of all deciduous trees, seldom on bark. Common.

Hypoxylon multiforme (Fries) Fries, S u m m a Veg. Scand. Sect. Post.:


384 (1849). (PI. X X X I I D . )
Stroma cushion-shaped or forming an extensive and somewhat
flattened crust, black and shining, covered with small pimples formed
b y the protruding upper p a r t s of the perithecia, their ostioles distinctly
papillate. Flesh up to 7 m m . thick, black and brittle. Asci cylindrical,
about 150x6//, 8-spored; ascospores uniseriate, elliptical with one
flattened side, 9 - 1 1 x 3 . 5 - 5 / / ; dark brown.
On bark and decorticated wood of dead branches of Betula. Common.

Hypoxylon serpens (Persoon ex Fries) Fries, S u m m a Veg. Scand. Sect.


P o s t . : 3 8 4 (1849). (PI. XXXIIF.)

Stroma a thin irregular cushion or widespread crust on wood or bark,


black, shining, roughened by the tips of the perithecia and their
papillate ostioles. Asci cylindrical, about 150x8/<, 8-spored; ascospores
elliptic-fusiform to bean-shaped with one flattened side, 1 0 - 1 5 x 5 - 7 / / ,
dark brown.
On decaying wood of any deciduous tree, very variable in external
appearance and extremely common.

Hypoxylon argillaceum (Persoon) Berkeley, Outlines of British Fungo-


logy:387 (1860); non Hypoxylon argillaceum (Fries) Fries, op. cit.:
384 (1849). (PL X X X I I H . )
Stroma a hemispherical cushion on bark or decorticated wood, up
to 5 m m . across, light grey-brown, with slightly protruding perithecia

280
b u t with their ostioles visible as minute pores, not protruding above
the surface. Flesh black, rather soft. Asci cylindrical, about 200x13//,
8-spored; ascospores uniseriate, elliptical to bean-shaped, 17-20x
8—10jM, dark brown.

On dead branches of Fraxinus. Not uncommon.


The valid name for this common species remains uncertain.

Hypoxylon fuscum (Persoon ex Fries) Fries, S u m m a Veg. Scand. Sect.


Post.:384 ( 1 8 4 9 ) . (PI. X X X I I c . )
S t r o m a t a varying from a small hemispherical cushion up to 4 m m .
across on bark to a crust 1 m m . thick on decorticated wood, the outer
surface purplish-brown to purplish-grey, smooth, the ostioles appear-
ing as pores in the crust only. Asci cylindrical, up to 150x8//, 8-spored;
ascospores uniseriate, elliptical with one flattened side, 1 2 - 1 5 x 5 - 7 / / ,
dark brown.

On dead branches of Corylus and Alnus. Very common.

Hypoxylon semi-immersum Nitschke, Pyrenomycetes Germanici:50


(1867). (PI. X X X I I G . )
Stroma forming a t h i n whitish layer on the surface of decorticated
wood around a small cluster of large protruding perithecia, blackening
with age. The perithecia have their lower halves immersed in the wood
and m a y be mistaken for those of a Rosellinia when the surrounding
stroma is feebly developed. Asci cylindrical, about 180x 12//, 8-spored;
ascospores uniseriate, elliptic-fusiform with one flattened side, 17-20x
10-12//, dark brown.

On rotting wood of deciduous trees, especially of Quercus, Fagus and


Ulmus. Not uncommon. Also known as Hypoxylon confluens (Tode ex Fries)
Cooke in Grevillea 11:139 (June 1883), a name of uncertain application.

Hypoxylon nummularium Buillard ex Fries, S u m m a Veg. Scand. Sect.


P o s t . : 3 8 4 (1849). (PI. XXXIIE.)

Stroma forming a more or less elliptical crust on wood or b a r k , at


first light brown, black and shining at m a t u r i t y , minutely roughened
with the papillate ostioles. Asci cylindrical, about 125x 12/t, 8-spored;
ascospores uniseriate, broadly elliptical to subglobose, 1 1 - 1 4 x 7 - 1 0 / / ,
dark brown to black.
On dead branches of Fagus, usually erumpent from beneath the bark.
Uncommon.

281
USTULINA Tulasne

Ustulina Tulasne, Selecta F u n g o r u m Carpologia 2:23 (1863).


Differs from Hypoxylon in its large perithecia and ascospores and
united with it b y m a n y authors.

Ustulina deusta (Fries) P e t r a k in Annales mycologici 19:279 (1921).


(PI. X X X I I M . )
Stroma an irregular cushion or thick crust, up to 3 m m . thick,
greyish-white when young, soon becoming black and t h e n very brittle,
easily detached from the substrate. Perithecia large, with papillate
ostioles, immersed in a whitish flesh which tends to break down in old
stromata, which are t h e n easily crushed in the fingers. Asci cylindrical,
about 300x15//, 8-spored; ascospores uniseriate, fusiform, with one
side somewhat flattened, 2 8 - 3 4 x 7 - 1 0 / / , black.
On stumps, butts and dead roots of deciduous trees, especially Fagus.
Common.

CAMAROPS Karsten

Camarops Karsten, Mycologia Fennica 2:53 (1873).


Stroma a superficial cushion with a thin reddish to black crust en-
closing numerous t u b u l a r perithecia; asci and ascospores as in Hypo-
xylon. There is only one British species:

Camarops polyspcrmum (Montagne) Miller in Trans. Brit, mycol. Soc.


15:151 (1930). (PI. X X X I I K . )
Stroma a somewhat flattened cushion, irregular in outline b u t up to
8 m m . thick, with a reddish to black crust enclosing a very large n u m -
ber of narrow cylindrical perithecia, so tightly packed as to appear
hexagonal like cells of a honeycomb in a cross-section, perithecial walls
thin, pinkish, with a narrow white zone at the top beneath the crust,
ostioles scarcely protruding. Asci cylindrical, about 8 0 x 4 / / , 8-spored;
ascospores uniseriate, elliptical, olive-grey to black, 5 - 6 x 2 - 3 . 5 / / .
On dead wood. Rare in England, common in the tropics.

BOLINIA (Nitschke) Saccardo

Bolinia (Nitschke) Saccardo, Sylloge F u n g o r u m 1:352 (1882).


There is a single British species:

282
Bolinia lutea (Albertini & Schweinitz ex Fries) Miller, Monogr. World
Species Hypoxylon: 138 (1961). (PI. X X X I I i . )
Stroma a circular or elliptical cushion up to 20 m m . long and 6 m m .
thick, with a flat or slightly concave upper surface, light yellowish,
darkening with age; flesh woody, light yellowish, with numerous flask-
shaped perithecia inserted at different levels in it, their ostioles opening
on the upper flat surface of the strome b u t not protruding. Asci
cylindrical about 8 0 x 5 / / ; ascospores elliptical, 5 - 7 x 3 - 4 / / , uniseriate,
dark olive-brown.

On dead branches and stumps, usually of Buxus in England. Uncommon.

NUMMULARIA Tulasne

Nummylaria Tulasne, Selecta F u n g o r u m Carpologia 2 : 4 2 (1863),


emend Miller in Trans. Brit, mycol. Soc. 17:132 (1932).
Stroma a more or less circular disc or flattened cushion w i t h the
perithecia seated in its base instead of just b e n e a t h the crust as in
ra; perithecia therefore flask-shaped with long t u b u l a r necks,
ascospores as in Hypoxvlnn with which the genus is often united.

Niiininularia discreta (Schweinitz) Tulasne, Selecta F u n g o r u m Carpo-


logia 2 : 4 5 (1863). ( P I . X X X I I L . )
Stroma circular or elliptical, up to 5 m m . across, arising beneath the
surface of wood or b a r k and becoming exposed b y the falling away of
the overlying tissue, grey or black, w i t h a slightly concave upper sur-
face and well defined rim. Flesh grey, surrounding the perithecial
necks, lower p a r t of t h e perithecia sunk in t h e b a r k or wood, ostioles
minute, not protruding. Asci cylindrical, about 200x15//, 8-spored;
ascospores subglobose, 12-24//, across, black at m a t u r i t y .

On Pyrus malus. Rare.

DALDINIA Cesati & de Notaris

Daldinia Cesati & de Notaris in Comm. Soc. Crittogamologica Italiana


1 : 1 9 7 (1863).
Stroma like t h a t of a Hypoxylon b u t comparatively large, hemi-
spherical, with distinct concentric zones of growth in its fibrous flesh,
perithecia in a single layer just beneath the crust.

283
Daldinia concentrica (Bolton ex Fries) Cesati & de Notaris, op. cit.: 198
(1863). (PI. X X X I I J . )
Stroma hemispherical, the base somewhat incurved, up to 4 cm.
across, at first reddish-brown, soon becoming black and somewhat
shiny, smooth, dotted with minute pores formed by the ostioles; flesh
dark purplish-brown, fibrous, with darker concentric zones; perithecia
small, crowded, in a single layer beneath t h e thin crust. Asci about
200x12//, cylindrical, 8-spored; ascospores uniseriate, elliptic-fusiform
with one flattened side, 1 2 - 1 7 x 6 - 9 / / , black.

On dead branches, especially of Fraxinus but also on other trees and a


small state is sometimes found on dead Ulex and Betula after heath fires.
Very common.

XYLOSPHAERA Dumortier

Xylosphaera Dumortier, Commentationes botanicae:91 (1822).


(= Xylaria Hill ex Greville in Flora Edinensis:355 (1824).
Stroma erect, more or less stalked, usually more or less club-shaped,
cylindrical or fusiform, sometimes forked, black or nearly so b u t
sometimes p a r t l y covered with light coloured conidia when immature,
perithecia in a single layer, usually inserted beneath a black crust, in
a few species superficial, flesh of the stroma usually white, asci and
ascospores as in Hypoxylon.

Xylosphaera polymorpha (Persoon ex Merat) Dumortier, op. cit.:92


(1822). (Fig. 12A.)
Stroma more or less club-shaped with a short cylindrical stalk,
sometimes irregular or even slightly lobed at the tip, solitary or in
clusters, seldom more t h a n 8 b y 2.5 cm.; flesh tough, solid, white, crust
thin, black, not shining, papillate with the protruding tips of the peri-
thecia or wrinkled somewhat between them, surface of the crust
appearing finely granular under a lens, ostioles prominent, light
coloured. Asci cylindrical, over 200x10//, 8-spored; ascospores uni-
seriate, fusiform with one side flattened 2 0 - 3 2 x 5 - 9 / / , black. Young
stromata are covered with a layer of light brown conidia.

Usually in small clusters at about soil level on dead stumps, especially of


Fagus. Common.

Xylosphaera longipes (Nitschke) Dennis in Kew B u l l : 104 (1958)


Fig. 19c), is a similar b u t usually more slender species with ascospores
12-16 x5—7//; it is usually attached to fallen branches of Acer.

284
Xylosphaera hypoxylon [Linnaeus] Dumortier, op. cit.:91 (1822).
(Fig. 12B.)
Stroma slender, up to about 8 cm. tall, subcylindrical to strap-
shaped and usually forked, often repeatedly at t h e tip, stalk black and
hairy, upper fertile portion at first with the powdery conidia, darken-
ing as the perithecia develop and t h e n papillate with their protruding
tips. There are also commonly separate unbranched s t r o m a t a with
pointed tips, bearing perithecia only. Asci cylindrical, about 1 0 0 x 8 / / ,
8-spored; ascospores uniseriate, slightly bean-shaped, 11-14x5-6/«,
black.

Very common everywhere on dead wood throughout the year.

Xylosphaera carpophila [Persoon] Dumortier, op. cit.:92 (1822), is a


slender species found on fallen beech mast.
Xylosphaera oxyacanthae (Tulasne) Dennis in Kew Bull.: 105 (1958),
springs from fallen or buried fruits of Crataegus.

Xylosphaera hippotrichoides (Sowerby ex Fries) Dennis in Kew Bull.


104 (1958). (Fig. 12F.)
Stroma very long and slender, like black horse-hair, with small
black flask-shaped perithecia scattered along it. Asci cylindrical,
8-spored; ascospores fusiform with one flattened side, 1 4 - 2 0 x 7 - 1 0 / / ,
black.
On decaying sawdust, rush matting and other debris, especially in cellars
and damp churches. Rare or perhaps not native. This species is sometimes
called Thamnomyces hippotrichoides (Sowerby) Ehrenberg but the genus
Thamnomyces Ehrenberg ex Fries in Linnaea 5:534 (1830), was based on a
tropical American fungus of very different character. The genus Simoninus
Roumeguere Fung. Sel. Exs.:211 (1879), was proposed to accommodate
fungi like X. hippotrichoides.

ROSELLINIA Cesati & de Notaris

Rosellinia Cesati & de Notaris in Giorno Botanico Italiano 1 : 3 3 4


(1844).
Perithecia superficial, usually on bark, not embedded in a common
stroma b u t often occurring in dense swarms upon a common blackish
hyphal m a t (subiculum), perithecia subglobose, smooth, black, ostiole
papillate; asci with a well developed apical ring, ascospores black,
non-septate, often with minute colourless appendages.

285
Rosellinia aquila (Fries) de Notaris, op. cit.:334 (1844). (Fig. 12c.)
Perithecia subglobose, 1-1.5 m m . across, somewhat flattened round
the prominent papillate ostiole, smooth, black, seated in dense swarms
on a dark brown h y p h a l m a t . Asci about 2 0 0 x 1 0 f i , cylindrical,
8-spored; ascospores uniseriate, elliptical,black, 17-27x7-8,«, one side
often somewhat flattened and bearing a longitudinal furrow; there is
usually a minute round hyaline appendage at each end of the ascospore.
On fallen branches of various deciduous trees, especially in spring.
Common.

Rosellinia thelena (Fries) Rabenhorst, Fungi europaei No. 575 (1865),


differs in its more pointed ascospores with a slender pencil-like hyaline
appendage at each end.

Rosellinia mammiformis (Persoon ex Fries) Cesati & de Notaris in


Comm. Soc. Crittogamologica Italiana 1:227 (1863). (Fig. 12D.)
Perithecia subglobose, about 1 m m . across, smooth, black and shiny,
w i t h a small apical papilla, scattered and without a subiculum. Asci
cylindrical, about 110x10/«, 8-spored; ascospores uniseriate, cylindric-
fusiform with a small b u t distinct beak at each end b u t no hyaline
appendage, 20-26x7.5-9/«, black with a distinct longitudinal furrow.
On dead stems and branches especially of Hedera but also on Fraxinus,
Salix etc. In spring and summer. Common.
For similar fungi with hairy perithecia see Coniochaeta and Helmintho-
sphaeria.

PORONIA Willdenow ex S. F. Gray

Poronia Willdenow ex S. F. Gray Nat. Arrang. Brit. Plants 1 : 5 1 5


(1821).
Stroma light coloured, nail-shaped with a flattened apical disc
beneath which is inserted a single layer of perithecia, asci and asco-
spores as in other genera of the family.

Poronia punctata (Linnaeus ex Fries) Fries, S u m m a Veg. Scand. Sect.


Post.:382 (1849). (Fig. 12II.)
S t r o m a nail-shaped, with a long cylindrical stalk buried in the dung,
expanded to a flat more or less circular disc at the surface, stalk black,
disc white, up to 15 m m . across, d o t t e d with the minute black ostioles

286
of the perithecia. Asci cylindrical, about 180 x 18/i, 8-spored; ascospores
uniseriate, somewhat bean-shaped, 1 8 - 2 6 x 7 - 1 2 / / , blackish-brown,
with a gelatinous coat.

On horse dung. Evidently abundant in the nineteenth century, now ex-


tremely rare but apparently not quite extinct.

PODOSORDARIA Ellis & Holway

Podosordaria Ellis & Holway in Botanical Gazette 24:37 (1897).


Fungi somewhat intermediate in character between Xylosphaera
and Poronia, having a small n u m b e r of r a t h e r p r o t u b e r a n t perithecia
clustered in the subglobose tip of a small stalked stroma arising from
dung. The crust of the perithecia is brown or black and the flesh of the
stroma white and solid.

Podosordaria tulasnei (Nitschke) Dennis in Kew Bulletin:306 (1957).


(Fig. 19A.)
Stroma with a head 2 - 3 m m . across, its surface occupied b y protrud-
ing hemispherical perithecia with small discoid ostioles; stalk cylin-
drical, rooting deeply in the dung. Asci cylindrical, about 160x20/<,
with rounded tips and massive apical ring stained deep blue by iodine,
8-spored; ascospores uniseriate, elliptical, 2 1 - 2 4 x 1 1 - 1 2 / / , dark brown
with a thin hyaline mucilaginous sheath.

Rooting in old rabbit droppings.

Podosordaria leporina (Ellis & E v e r h a r t ) Dennis in Kew Bulletin:


306 (1957), also on rabbit droppings, differs in its brown perithecia
with ascospores 1 2 - 1 7 x 6 - 9 / / .
Podosordaria pedunculata (S. F . Gray) Dennis loc. cit. has larger
s t r o m a t a with ascospores 4 0 - 6 0 x 2 0 - 3 0 / / , deeply rooting in dung or
richly manured soil.

Family 9. AMPHISPHAERIACEAE

Here are assembled perithecial fungi in which the ascus has an apical
ring blued by iodine or a pore plug whose inner surface reacts in the
same way, b u t with ascospores unlike those of the Xylariaceae.

287
I. Ascospores hyaline, non-septate, inner surface of the ascus pore blued by
iodine:
A. Perithecia small, upper part of the wall composed of large angular
cells, asci small, ostiole minute, inconspicuous, in leaves oiQuercus
Anisostomula (p. 289)
B. Perithecial wall colourless with an olive-brown ostiole, in leaves of
Buxus Hyponectria (p. 289)
II. Ascospores septate:
A. Ascospores hyaline to yellowish:
1. Ascospores 1-septate:
a. Ascospores very large, dumb-bell shaped, on Ilex
Vialaea (p. 293)
b. Ascospores elliptical, smaller:
*Ascospores with a median septum:
Perithecia with a clypeus, in herbaceous stems
Paradidymella (p. 292)
Perithecia without a clypeus, ascospores with a
germ-pore at each end, in leaves of Dryas
Cainiella
**Ascospores with septum near one end:
Perithecia in aseries in a stroma on leaves of grass
Apiospora (p. 292)
Perithecia solitary, in twigs or dicotyledonous
leaves:
Ostiole surrounded by bristles Chaetapiospora
No so Pseudomassaria (p. 291)
2. Ascospores with more than one septum
Gripliosphaeria (p. 290)
B. Ascopores brown:
1. Is leaves of pmonocotyledons:
na. Ascosore with a hyaline appendage at each end
Ceriophora (p. 294)
b. Without such an appendage, on grass and sedges in
Europe Cainia
2. In or on wood or bark or in dicotyledonous stems and leaves:
a. Ascospores 1-septate:
*Perithecia embedded in bark, forming pustules
Massariella (p. 293)
**Perithecia with the upper part protruding
Amphisphaeria (p. 294)
***Perithecia in leaves of Dryas, see Cainiella above
b. Ascospores finally with more than one septum:
*Ascospores with a hyaline cell at one end
Apiorhynchostoma (p. 296)
**Not as above Clypeosphaeria (p. 295)
***3-septate Diapleela (p. 295)

I have not noted a blue reaction with iodine in Apiorhynchostoma


b u t the genus is referred here b y Miiller and von Arx and it certainly
appears akin to Clypeosphaeria. Chaetapiospora islandica (Johanson)

"288
P e t r a k in Sydowia 1:87 (1947) should be sought in Britain in leaves of
Dry as octopetala, as should Cainiella johansonii (Rehm) Miiller in
Sydowia 10:121 (1956). Chaetapiospora rhododendri (Tengwall von Arx
in Ber. Schweiz. Bot. Ges. 62:359 (1952) should be looked for in leaves
of Rhododendron ponticum.

ANISOSTOMULA von Hohnel

Anisostomula von Hohnel in Annales mycologici 16:49 (1918).


Perithecia scattered, completely immersed in the mesophyll of dead
leaves, subglobose, flattened above and below, perithecial wall com-
posed of a few layers of dark brown angular cells, asci 8-spored, asco-
spores hyaline, non-septate.

Anisostomula cookeana (Auerswald) von Hohnel, op. cit.:49 (1918).


(PI. X X X V g . )
Perithecia in swarms, immersed individually in the leaf tissue and
visible as minute dark dots between the veins on t h e under side of t h e
leaf, about 150// across and 110// high, ostiole very minute, not
papillate. Asci cylindric-clavate, distinctly stalked, up to 50x10/«,
8-spored, the inner surface of t h e pore blued b y iodine; ascospores
biseriate, elliptic-fusiform or inequilateral, 8 - 1 0 x 2 . 5 - 3 / / , hyaline, non-
septate; paraphyses few and slender.

On dead and fallen loaves of Quercus. Common


There is said to be a second species on oak leaves, A. areola (Fuckel) von
Hohnel, op. cit.: 49 (1918), with ascospores 7-10x3-4/« and with the brown
perithecial wall developed on its upper and lower surfaces only.

HYPONECTRIA Saccardo

Hyponectria Saccardo in Michelia 1 : 2 5 0 ( J a n u a r y 1878).


Perithecia embedded in leaf tissue, scattered, somewhat flattened,
wall composed for the greater p a r t of slender almost colourless hyphae,
with a ring of dark brown angular cells around the ostiole, asci 8-spor-
ed, ascospores hyaline, non-septate. There is a single British species:

Hyponectria buxi (Desmazieres) Saccardo, op. cit.:250 (1878). (PI.


XXXIL.)
Perithecia gregarious, embedded in the mesophyll and visible as
dark olive dots on the under surface of the leaf, distinctly flattened,

"289
about 0.3 m m . across and 0.15 m m . high, ostiole conspicuous b u t not
protruding. Asci clavate, stalked, up to 8 5 x 1 4 / / , 8-spored, the inner
surface of the pore blued b y iodine; ascospores biseriate, elliptical,
1 2 - 1 8 x 4 - 5 / / , hyaline, non-septate.
On leaves of Buxus, throughout the year. Very common.

GRIPHOSPHAERIA von Hohnel

Griphosphaeria von Hohnel in Annales mycologici 16:87 (1918).


Perithecia embedded in b a r k in small clusters often covered b y a
small m a t of stromatic tissue in t h e form of a clypeus, asci cylindrical,
with an apical thickening which stains blue with iodine on its inner sur-
face, ascospores uniseriate, 3-septate, hyaline. The genus has been
united with the older Clathridium Berlese, Icon. Fung. 2:110 (1894)
by Shoemaker & Miiller in Canad. J o u r n . Bot. 42:403 (1964).

Griphosphaeria corticola (Fuckel) von Hohnel in Annales mycologici


16:87 (1918). (PI. XXXVH.)
Perithecia less t h a n % m m - across, black, inserted often in pairs
beneath a small grey clypeus in the surface layer of bark, ostioles
visible on the surface as m i n u t e black dots, not protruding. Asci
cylindrical, short-stalked, up to 9 0 x 8 / / , 8-spored; ascospores uni-
seriate, elliptic-cylindric, 1 1 - 1 4 x 5 - 6 / / , 3-septate, thin-walled, hyaline.

On dead twigs of Salix and Rosa, sometimes parasitic on the latter.


Uncommon.
The condial state is Seimatosporium lichenicola (Corda) Shoemaker &
Miiller op. cit.: 405 (1964).

Griphosphaeria nivalis (Schaffnit) E. Miiller & von Arx in P h y t o p a t h .


Zeitschr. 24:356 (1955). (Fig. 25AA.)
Perithecia scattered or in small groups, immersed, globose, dark-
brown, smooth, a b o u t 150-200// diameter, thinwalled. Asci 4 0 x 6 / / ,
8-spored, inner surface of t h e pore blue in iodine; ascospores biseriate,
elliptic-fusoid, straight or slightly curved, 1 0 - 1 8 x 3 - 3 . 5 / / ,
Abundant in dead leaves of grasses and cereals. The conidial state is
Fusarium nivale (Fries) Cesati, and because of this very different conidial
development it is unlikely that G. nivalis would be referred to Clathridium.
Booth in Comm. myc. Pap.: 94 (1964) suggested an appropriate genus for it
would be Micronectriella von Hohnel but did not make the necessary re-
combination. Micronectriella agropyri Apinis & Chesters in Trans. Brit.

"290
mycol. Soc. 47:432 (1964) has been described from leaves of Agropyron
pungens with ascospores 3-5-septate, 16-22 x 4-6 ji.

PSEUDOMASSARIA Jaczewski

Pscudomassaria Jaczewski in Bull. herb. Boissier 2:662 (1896).


Perithecia scattered, embedded in b a r k of twigs, not beaked, stroma
absent or represented merely b y a thin belt of hyphae around and
above the individual perithecia, asci clavate, short-stalked, thin-
walled, ascospores hyaline, ellipsoidal with a septum near the lower
end as t h e y lie in the ascus.

Pscudomassaria sepincolaeformis (de Notaris) von Arx in Ber. Schweiz.


B o t . Ges. 6 2 : 3 5 0 (1952). (PI. XXXVIIIH.)

Perithecia gregarious, immersed in cortical tissue, subglobose,


broader t h a n high, up to 0.3 m m . wide, with short ostioles perforating
the epidermis, visible under a hand-lens as small grey patches on dead
twigs. Asci up to 110x20//, cylindric-clavate, 8-spored, the inner sur-
face of the apical pore blued b y iodine; ascospores more or less bi-
seriate, ovoid, the broader end uppermost in the ascus, 17-22x7-10/«,
hyaline, with a septum near the narrow end.

On dead twigs of Rosa. Uncommon. Also known as Didymella sepincolae-


formis (de Notaris) Saccardo, Sylloge Fungorum 1:551 (1882), and Apio-
sporella sepincolaeformis (de Notaris) Theissen in Annales mycologici 15:275
(1917).

Pscudomassaria chondrospora (Cesati) Jaczewski, op. cit.:663 (1894).


Perithecia immersed, flattened, about 250-350/« wide and up to
200// tall, brown. Asci 6 5 - 9 5 x 2 0 - 3 0 / / , spores biseriate, 1 8 - 3 5 x 8 - 1 2 / / ,
with a primary septum 4-6/« from the lower end, long remaining hya-
line b u t ultimately light brown and sometimes developing a f u r t h e r
approximately median septum. (Fig. 25w.)
In dead twigs of Tilia. Rare.

Pseudomassaria corni Sowerby ex von Arx, op. cit.:349 (1952), is a


similar fungus, with r a t h e r less eccentrically septate ascospores 18-24
x 6 - 8 / / on dead twigs of Cornus.
For an account of several other European species of Pseudomassaria
see a paper by J . A. von Arx in Schweiz. Bot. Ges. Ber. 62 (1952).

"291
APIOSPORA Saccardo

Apiospora Saccardo in A t t i Soc. Venet.-Trent. Sci. Nat. 4 : 8 5 (1875).


Perithecia similar to those of Pseudomassaria b u t arranged in rows
within a small elongated blackish stroma.

Apiospora montagnei Saccardo in Nuovo Giornale Botanico Italiano


7:306 (1875). (PI. X X X V I I I D . )
S t r o m a t a embedded beneath the epidermis of the larger grass culms,
up to 5 m m . long b y 0.6 m m . wide, forming raised grey spots on the
surface which splits longitudinally above t h e m , perithecia closely
packed, in rows, black, about 0.2 m m . across, globose with short
papillate ostioles. Asci cylindric-clavate, up to lOOx 17//, short-stalked,
8-spored; ascospores biseriate, ellipsoidal, 2 0 - 3 0 x 6 - 1 0 jx, hyaline, with
a septum near the narrower end.

Common in the Mediterranean region and once recorded in England on


Cortaderia selloana. I have seen no British material and have figured the
species from an Algerian collection.

PARADIDYMELLA P e t r a k

Paradidymella P e t r a k in Annals mycologici 25:237 (1927).


Like Sydowiella b u t without beaks to the perithecia, which are each
covered b y a small grey clypeus. The genus has been united with
Griphosphaeria and Lejosphaerella von Hohnel under the older name
Clatkridium Berlese b y Miiller& Shoemaker in Ganad. Journ. Bot. 43:
1343 (1965).

Paradidymella tosta (Berkeley & Broome) P e t r a k , op. cit.:238 (1927).


(Fig. 26 G.)
Perithecia scattered or sometimes in pairs, subglobose, blackish, up
to 0.3 m m . wide, immersed in the host cortex and each covered b y a
p a t c h of dark brown hyphae, forming a clypeus visible as a minute
grey stain on the surface of the stem. Asci cylindrical, up to 8 0 x 6 / t ,
with a small apical ring which stains dark blue with iodine, 8-spored;
ascospores mostly uniseriate, elliptic-fusiform, 10-14x 3 - 4 / t , with a
median septum, not constricted, hyaline.

On dead stems of Chamaenerion, angustifolium & Epilobium spp., often


in company with Sydowiella fenestrans, in spring. Common.
The conidial state is Seimatosporiurn Kriegerianum (Bresadola) Morgan
Jones & Sutton; Lejosphaerella vexata (Saccardo) Miiller is a similar fungus,
with ascospores 21-30 x 9-12 p on twigs of Cornus.

"292
YIALAEA Saccardo

Vialaea Saccardo in Hedwigia 35 R e p e r t o r i u m : x x x i (1896).


Perithecia scattered, immersed in cortical tissues, without a stroma,
not beaked, asci cylindric-clavate, ascospores very large, swollen to-
wards each end with a single septum in the central slender part, hyaline
to brownish.

Yialaea insculpta (Fries) Saccardo, op. c i t . : x x x i (1896). (Fig. 23c.)


Perithecia scattered, globose to subconical, up to 0.4 m m . across,
black, seated on the wood and perforating t h e bark b y a short black
ostiole which appears on t h e surface as a dot surrounded b y a brown
halo. Asci narrowly cylindrical, apex rounded and rather thick, the
inner surface of t h e pore stained blue b y iodine, up to 170x20//,
8-spored; ascospores irregularly biseriate, elongated, narrow in the
middle and inflated towards each end, tips pointed, about 80// long,
3// wide in the centre, 7 - 8 / / wide at the broadest parts, 1-septate,
hyaline.

On dead twigs of Ilex aquijolium. Not uncommon. Afso known as Boydia


remuliformis A. L. Smith.

MASSARIELLA Spegazzini

Massariella Spegazzini in Ann. Soc. Cient. Argentina 4 (9), Table


following p. 192 (1880).
Perithecia scattered, immersed in b a r k , not beaked, asci cylindric-
clavate thin-walled, ascospores 1-septate, brown. The genus is united
with Amphisphaeria b y Miiller & von Arx (1962).

Massariella bufonia (Berkeley & Broome) Spegazzini op. cit. (1880).


(Fig. 2 3 B . )
Perithecia scattered, black, immersed in the outer layers of bark and
raising it into small pustules, scarcely 1 m m . across, with a low
papillate ostiole just reaching the surface. Asci narrowly cylindric-
clavate, short-stalked, up to 250x16/«, 8-spored, the broad pore
stained deep blue b y iodine; ascospores uniseriate, elliptic-cylindric
b u t m a r k e d l y constricted at the single septum, 22-28x9-12/«, the
lower cell sometimes slightly larger t h a n t h e upper, dark brown, with a
gelatinous envelope.

"293
Oil dead branches of Quercus, in winter. Not uncommon.
Massariella vibratilis (Fuckel) Saccardo, Sylloge F u n g o r u m
1:716 (1886), with ascospores 17-24x7-9//, occurs on Prunus.

AMPHISPHAERIA Cesati & de Notaris

Amphisphaeria Cesati & de Notaris in Commentario della Societa


Crittogamologica Italiana 1 (4): p. 223 (1863), emend P e t r a k in
Annales mycologici 13:33 (1923).
Perithecia large, erumpent from bark, smooth, papillate, asci nar-
rowly cylindrical, with apical ring blued by Melzer's reagent, asco-
spores 1-septate, brown, paraphyses numerous, slender.

Amphisphaeria umbrina (Fries) de Notaris, Sferiacei italici (1863) (n.v.).


Perithecia scattered or in rows, semi-immersed in bark, about 1 m m .
diameter, upper p a r t protruding, dull black, smooth, with a small apical
papilla. Asci thin-walled, 2 0 0 x 8 / / ; ascospores uniseriate, elliptic-
cylindric, 1-septate b u t scarcely constricted, brown, 1 7 - 2 6 x 6 - 8 / / ;
paraphyses numerous, slender. (Fig. 20D.)

In thick bark of Ulmus.

Amphisphaeria millepunctata (Fuckel) P e t r a k in Ann. Mycol. 21:329


(1923) with ascospores 1 2 - 1 8 x 5 - 7 / / occurs on dead twigs of Acer,
Carpinus, Corylus, Prunus, Quercus, Salix, Sorbuslk Viburnum.

CERIOPHORA von Hohnel

Ceriophora von Hohnel in S.B. K. Akad. Wiss. Wien, Math.-Nat. Klasse


Abt. 1 , 1 2 8 : 5 8 6 (1919).
Perithecia scattered, immersed in leaves of Monocotyledons, not
beaked, asci cylindric-clavate, thin-walled, ascospores 1-septate,
brown, with a conspicuous hyaline appendage at each end.

Ceriophora palustris (Berkeley & Broome) von Hohnel apud Clements


& Shear, Genera of F u n g i : 2 6 8 (1931). (PI. X X X V I I n . )
Perithecia scattered, globose, black, up to % m m - across, immersed
in the mesophyll, with a small papillate ostiole visible as a minute

"294
black dot on the leaf surface. Asci narrowly cylindrical, short-stalked,
about 180x10//, thin-walled with a conspicuous apical ring-like
thickening which stains deep blue in iodine; ascospores uniseriate,
elliptic-fusiform, 1-septate and slightly constricted at the septum,
1 6 - 2 4 x 6 - 7 / / , brown, with a gelatinous coating which is prolonged
into a conspicuous rounded appendage at each end of the spore.

On dead leaves of Carex, Iris etc., in winter.

DIAPLEELLA Munk

Diapleella Munk in Dansk Bot. Ark. 15 (2): 125 (1953).


There is a single species, superficially resembling a Leptosphaeria, to
which genus it has long been referred, b u t with longstalked thinwalled
asci. Compare Chitonospora p. 403.

Diapleella clivensis (Berkeley & Broome) Munk, op. cit.:75 (1953).


(Fig. 2 6 F . )
Ascocarps scattered, immersed, globose, about y 3 m m . across, black,
with papillate erumpent ostioles. Asci narrowly clavate, a b o u t 95 x
16//, 8-spored; ascospores biseriate, elliptical or slightly fusiform,
1 8 - 2 4 x 6 - 8 / / , 3-septate, somewhat constricted at the septa, yellowish-
brown.
On dead herbaceous stems. Not uncommon.

CLYPEOSPHAERIA Fuckel

Clypeosphaeria Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:


117 (1870).

Perithecia scattered, embedded in bark beneath a small dark stroma


(clypeus), not beaked, asci cylindric-clavate, with a well developed
apical ring, ascospores brown, ultimately 3-septate. Though the genus
is placed here on account of its septate ascospores its affinities appear
to lie r a t h e r with Anthostomella.

Clypeosphaeria mamillana (Fries) L a m b o t t e in Mem. Soc. royale


Sciences Liege I I , 1 4 : 1 2 8 ( 1 8 8 7 ) . ( P L X X X V E . )
Perithecia scattered, subglobose, about y 2 m m . across, black, each
embedded in bark beneath a small p a t c h of blackish-brown mycelium

"295
which forms a conspicuous clypeus, ostiole shortly conical, not pro-
truding beyond t h e surface. Asci narrowly cylindric-clavate, short-
stalked, u p to 1 5 0 x 9 / / , 8-spored, apical ring blued b y iodine; ascospores
uniseriate, elliptic-cylindric, usually slightly curved, 1 8 - 2 4 x 5 - 6 / / ,
with a r a t h e r thick outer wall and ultimately three thin transverse
septa, dark brown.
On dead twigs of various deciduous trees and shrubs such as Acer, Cornus
and Quercus.

Clypeosphaeria notarisii Fuckel, common on dead Rubus, m a y be


only a form of the same species.

APIORHYNCHOSTOMA P e t r a k

Apiorhynchostoma P e t r a k in Annales mycologici 21:185 (1923).


Perithecia clustered in small groups, embedded in dead wood,
without a distinct stroma, asci cylindrical, ascospores brown, with a
small hyaline cell at one end, ultimately 2-septate.

Apiorhynchostoma curreyi (Rabenhorst) Muller in Miiller & von A r x :


707 (1962). (PI. X X X V I I K . )
Perithecia embedded in t h e outer layers of decorticated wood, in
small clusters, associated with a grey stain b u t without a defined
stroma, flask-shaped, their ostioles opening independently, black,
cavity of the perithecium about 0.4 m m . across. Asci narrowly
cylindric-clavate, stalked, up to 160x11//, thin-walled, 8-spored;
ascospores uniseriate, elliptic-cylindric, sometimes slightly curved and
sausage-shaped, 1 8 - 2 7 x 5 . 5 - 9 / / , brown, with a small hyaline cell cut
off at t h e lower end and ultimately a second septum towards the upper
end of t h e spore.
In decorticated coniferous wood; the type collection came from wooden
railings beside the railway near Weybridge, Surrey, a century ago and it has
been seldom collected since.

F a m i l y 10. Diatrypaceae

As here interpreted the family includes a r a t h e r artificial assemblage


of genera, brought together b y possessing small hyaline to light brown
sausage-shaped ascospores, similar to those of the Coronophorales. The

"296
beginner will probably find it easier to trace genera in this w a y b u t
most modern authorities transfer Peroneutypa, Valsa and their allies
to the Diaporthaceae, with which t h e y have more in common in the
arrangement of the perithecia within the stroma. The very peculiar
Enchnoa and Calosphaeria have no obvious close relationships and
ought perhaps to be relegated to separate families.

I. Perithecia embedded in a crust-like or cushion-like stroma or immersed


in swarms over a wide expanse of wood or bark, orientated verti-
cally, with the ostioles emerging singly:
A. Stroma cushion-like, erumpent:
1. Asci 8-spored:
a. Perithecia not beaked Diatrype (p. 298)
b. Perithecia beaked Eutypella (p. 300)
2. Asci many-spored Diatrypella (p. 299)
B. Stroma completely embedded in the wood or rudimentary:
1. Perithecia embedded in wood beneath a blackened surface
layer Eutypa (p. 298)
2. Perithecia without a stroma or blackened surface layer:
a. Perithecia immersed in bark, scattered evenly over wide
areas Cryptosphaeria (p. 301)
b. Perithecia immersed in wood, singly or in small groups
Endoxyla (p. 304)

II. Perithecia in small circular clusters, often lying obliquely with their
ostioles converging at the centre and erupting in a common stromatic
disc:
A. Perithecia with free ostioles on the ends of long protruding beaks,
interspersed with conidial stalks . . . Peroneutypa (p. 303)
B. Perithecial necks not extruded beyond an ostiolar disc on the bark
surface:
1. Asci with eight spores or less:
a. Ostiolar disc white Leucostoma (p. 302)
b. Ostiolar disc grey or black:
fPerithecia in large clusters, long-necked, often as-
sociated with a Cytospora conidial state
Valsa (p. 301)
tfPerithecia in groups of four or five, short-necked
Quaternaria (p. 300)
2. Asci with very numerous spores . . . . "Valsella (p. 303)

III. Perithecia arranged radially beneath bark, like spokes of a wheel, their
ostioles converging at the centre and erumpent through a slit in the
bark Calosphaeria (p. 305)

IV. Perithecia large, without prominent necks, in a cluster beneath bark


and surrounded by a web of loose brown hyphae Enchnoa (p. 304)

"297
EUTYPA Tulasne
Eutypa Tulasne, Selecta F u n g o r u m Garpologia 2:52 (1863).
Stroma widely extended in the surface layers of wood or bark which
t h u s become blackened, perithecia scattered evenly in a single layer
just beneath t h e surface, flask-shaped, the ostioles often prominent,
sometimes distinctly beaked, black, sometimes ornamented with radial
furrows. Asci r a t h e r small, clavate, stalked, with a well developed
apical ring in some species, 8-spored; ascospores sausage-shaped,
usually light brown.

Eutypa acharii Tulasne, op. cit.:53 (1863). (Fig. 18H.)


Stroma wide-spreading on the surface of decorticated wood, grey,
smooth between the ostioles which are minute, black and slightly pro-
jecting. Perithecia flask-shaped, about 0.5 m m . across. Asci clavate,
about 5fi wide above, with a slender stalk; ascospores eight, in a
cluster about 20-25/« long, individually sausage-shaped, 5 - 7 x 1 / « ,
with a minute oil drop at each end, almost colourless b u t appearing
light brown in the cluster.

On dead branches, especially of Acer. Not uncommon.

Eutypa spinosa (Persoon ex Fries) Tulasne, op. cit.:59 (1863), has a


wide-spreading black stroma on wood of Fagus, thickly set with short,
black, perithecial beaks each ornamented b y four to five furrows
radiating from the ostiole.

DIATRYPE Fries
Diatrype Fries, S u m m a Veg. Scand. Sect. P o s t . : 3 8 4 (1849).
Stroma cushion-shaped, discoid or forming a widespread crust,
erumpent from beneath bark of dead branches, asci and ascospores as
in Eutypa.

Diatrype disciformis (Hoffmann ex Fries) Fries, op. cit.:385 (1849).


(Fig. I S G . ) ^ ^

Stroma up to 3 m m . across, elliptical or circular, flat topped, at first


whitish dotted with the black ostioles, soon developing a black outer
surface while the flesh remains whitish, ostioles p r o t r u d i n g slightly
above the general level of the crust. Perithecia in a single layer, up to

"298
3/4 m m . across; asci club-shaped, with long slender stalks, the upper
part about 5/< wide, 8-spored; ascospores sausage-shaped, 5 - 8 x 2 / « ,
almost colourless, appearing slightly brown in the cluster.

In swarms on dead branches of Fjjgus, occasionally on those of other trees.


Very common.

Diatrype bullata (Hoffmann ex Fries) Fries, op. cit.:385 (1849),


forms similar s t r o m a t a on dead branches of Salix.

Diatrype stigma (Hoffman ex Fries) Fries, op. cit.: 385 (1849). Fig. 27 F.)
Stroma a wide-spreading crust, erumpent from the b a r k of dead
twigs and branches, about 1 m m . thick, often cracked and sometimes
breaking away in patches, surface smooth a p a r t from the innumerable
slightly protruding ostioles, at first pale purplish-brown, soon becom-
ing black. Asci club-shaped, about 8/« thick in the upper p a r t , long-
stalked, 8-spored; ascospores sausage-shaped, 6 - 1 0 x 2 / / , very pale
brown.
On dead wrood of deciduous trees, perhaps especially abundant on
Crataegus. Very common.

DIATRYPELLA (Cesati & de Notaris) Cooke

Diatrypclla (Cesati & de Notaris) Cooke in J o u r n a l of B o t a n y 4 : 9 9


(1866). Differs from Diatrype in the many-spored asci.

Diatrypclla quercina (Persoon ex Fries) Cooke, op. cit.:99 (1866).


(Fig. 13K.)
Stroma about 2 m m . across, erumpent from bark, cushion-shaped,
with a dark brown to black crust and hard greyish flesh, the crust often
cracked and slightly roughened by the somewhat protruding ostioles.
Perithecia in a single layer, each about % m m . wide; asci cylindric-
clavate, about 120x12/« containing very m a n y spores; ascospores
sausage-shaped, strongly curved, 8 - 1 3 x 2 . 5 - 3 / / , very pale brown.

On dead branches of Quercus. Very common in winter and spring.

Diatrypella javacea (Fries) Saccardo, Sylloge F u n g o r u m 1:201


(1882), is a similar fungus with ascospores 5 - 6 x 1 / / on dead branches
of Betula and Alnus.

"299
EUTYPELLA (Nitschke) Saccardo

Eutypella (Nitschke) Saccardo in Atti Soc. Venet.-Trent. Sci. Nat. 4 : 8 0


(1875).
Stroma usually small, cushion-shaped, r a t h e r feebly developed and
often enclosing fragments of bark, perithecia not necessarily in a single
layer, with columnar strongly protruding ostioles, usually ornamented
by several superficial longitudinal furrows; asci and ascospores as in
Eutypa.

Eutypella prunastri (Persoon ex Fries) Saccardo, Sylloge F u n g o r u m 1:


147 (1882). (Fig. 13D.)
Stroma cushion-shaped, up to 8 m m . across, developed in the upper
layers of the b a r k and erumpent t h r o u g h a crack, black, rough,
crowned b y a dense cluster of columnar or conical ostiolar necks, each
marked b y three to five deep longitudinal furrows in its upper p a r t .
Perithecia numerous, closely crowded and irregularly arranged, black;
asci club-shaped, long-stalked, the upper p a r t about 4// thick, 8-spored;
ascospores sausage-shaped, 6 - 8 x 1 / / , very pale brown.

On dead branches of Prunus spinosa, occasionally on the domestic plum,


especially on trees killed by Bacterial Canker. Not common.

Eutypella sorbi (Schmidt ex Fries) Saccardo, Sylloge F u n g o r u m 1:


148 (1882), occurs occasionally on dead Sorbus aucuparia.
Eutypella stellulata (Fries) Saccardo, Sylloge F u n g o r u m 1:149 (1882),
occurs on Ulmus.

QUATERNARIA Tulasne

Quaternaria Tulasne, Selecta F u n g o r u m Carpologia 2:104 (1863).


Stroma small, cushion-shaped, immersed in bark, containing up to
five perithecia whose ostioles converge and become erumpent in a
small stromatic disc; asci and ascospores similar to those of Eutypa
b u t m u c h larger.

Quaternaria quaternata (Persoon ex Fries) Schroeter, Kryptogamen


Flora von Schlesien 3 (2): 451 (1897). (Fig. 27i.)
Stroma circular, cushion-shaped, developed in the upper layers of
the bark and erumpent as a smooth black disc about y 2 mm. across,
bearing the slightly protruding ostioles, usually four in number. Peri-

"300
thecia three to five in a stroma, closely packed, each about % m m .
across, with black walls and short necks; asci cylindric-clavate, long-
stalked, about 7/t wide above, apex flattened, with a delicate ring-like
thickening, 8-spored; ascospores sausage-shaped, 1 3 - 2 1 x 2 . 5 - 3 . 5 / t ,
light brown.
In dense swarms, giving a greyish pimply surface to large areas of bark
on dead and fallen branches of Fagus in winter and spring. Very common.

Quaternaria dissepta (Fries) Tulasne, op. cit. 2:107 (1863), common


on Ulmus, is a somewhat similar fungus with m u c h bigger ascospores,
2 4 - 3 2 x 6 - 8 /<.

CRYPTOSPHAERIA Greville

Cryptosphacria Greville, Scottish Gryptogamic Flora t a b . 13 (1823)


emend Flora Edinensis:359 (1824).
Stroma wide-spreading b u t feebly developed, entirely within the
bark and lacking a black outer crust, perithecia scattered, in a single
layer, asci and ascospores like those of Quaternaria.

Cryptosphaeria eunomia (Fries) Fuckel in J a h r b . Nass. Vereins f. Na-


t u r k u n d e 23-24:212 (1870). (Fig. 27K.)
Stroma very wide-spreading within the bark, the outer surface of
which is scarcely discoloured b u t is closely and minutely dotted with
the scarcely protruding black ostioles, the lower surface of the stroma
forms a dark grey to black discoloration on the inner surface of the
bark. Perithecia spherical, about y 2 m m . across, with short necks,
black; asci club-shaped, long-stalked, up to 1 4 0 x l 2 / i , 8-spored, with
a distinct apical ring; ascospores sausage-shaped 1 3 - 1 6 x 2 - 3 / / ,
yellowish-brown.

On dead twigs and small branches of Fraxinus. Extremely common


everywhere.

YALSA Fries

Yalsa Fries, S u m m a Veg. Scand. Sect. Post.:410 (1849).


Stroma rudimentary, a mere web of hyphae between the perithecial
necks, perithecia clustered in small groups within dead b a r k , flask-
shaped, their long necks converging and erumpent together, usually
in a small grey pad of stromatic tissue, asci club-shaped, with eight

"301
spores or less, ascospores sausage-shaped, colourless. The perithecia
are usually preceded by a conidial stage belonging to the form-genus
Cytospora, characterised b y chambered pycnidia immersed in the bark
and containing innumerable, minute, hyaline, sausage-shaped conidia
which emerge in glutinous coils in d a m p weather.

Valsa ambiens (Persoon ex Fries) Fries, op. cit.:412 (1849). (PI.


XXXIIIA.)
Perithecia in clusters of up to 12, each about y 2 m m . across, ar-
ranged in a ring within the bark, their long slender necks converging
and penetrating a pad of grey stromatic tissue which becomes erum-
pent as a small grey disc, dotted with a ring of black ostioles. Asci
club-shaped, about 60x10/«, mostly 8-spored; ascospores sausage-
shaped, 1 5 - 2 0 x 3 - 4 f i , hyaline.

On dead twigs and small branches of deciduous trees, especially of Rosa-


ceae. Common. The form figured is that on Pyrus malus.
Many species of Valsa appear in the literature but the genus is in urgent
need of revision and it is not at all clear at present how many of the recorded
species are distinct or how many are based simply on differences in the host
on which they were described.

LEUCOSTOMA (Nitschke) von Hohnel

Leucostoma (Nitschke) von Hohnel in Ber. Deutsche Bot. Ges. 35:637


(1917).
Similar to Valsa, with which it is usually united, b u t separated by
von Hohnel on account of t h e b e t t e r developed stroma which envelops
the perithecia and is delimited b y a black line in the bark.

Leucostoma niveum (Persoon ex Fries) von Hohnel in Mitt. bot. Lab.


techn. Hochsch. Wien 5 : 5 8 (1928). (PL X X X I I I B . )
Stroma small, cushion-like with a flat base, developed within t h e
b a r k and raising it slightly into a pimple, outlined b y a thin black
layer, flesh grey, enclosing up to ten flask-shaped black perithecia,
apical portion of the stroma e r u m p e n t as a small flat white disc, on
which the ostioles appear as black dots. Asci clavate, about 60x9/«,
mostly 4-spored; ascospores sausage-shaped, 1 2 - 1 6 x 3 f i , hyaline.

On dead twigs of Populus tremula and P. nigra. Not common.

"302
VALSELLA Fuckel

Valsella Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:203


(1870).
Similar to Leucostoma b u t with very m a n y spores in the ascus. Ac-
cording to some authors the species of Valsella are mere polysporous
states of species of Leucostoma b u t this suggestion has yet to be de-
monstrated experimentally.

Yalsella salicis Fuckel, op. cit.:203 (1870). (PI. X X X I I I c . )


Stroma small, cushion-shaped, developed within the bark and rais-
ing it into a distinct pustule, about 1 m m . across, outlined b y a thin
black layer, flesh light brown, enclosing up to 12 perithecia, apex of
the stroma erumpent through a small split in the bark and exposed as
a grey disc dotted with the black ostioles. Asci cylindric-clavate, about
50x8/4, polysporous; ascospores slightly sausage-shaped, 5 - 8 x 1 . 5 / / ,
hyaline.

On dead twigs of Salix, in winter. Apparently rare.

PERONEUTYPA Berlese

Peroneutypa Berlese, Icones F u n g o r u m 3 : 8 0 (1902).


Stroma wide-spreading within bark or wood, like t h a t of an Eutypa,
with a blackish upper surface, perithecia inserted in a single layer,
scattered or more often in small valsoid clusters, with long, protruding,
cylindrical, black necks; asci clavate, long-stalked, 8-spored, asco-
spores sausage-shaped, slightly yellowish.

Peroneutypa heteracantha (Saccardo) Berlese, op. cit.:81 (1902).


(Fig. 2 7 J . )
Stroma widespread in wood or b a r k and sometimes causing a slight
greying of the surface, perithecia mostly clustered in groups of up to
five, black, minute, flask-shaped, with long, slender, cylindrical,
smooth, black beaks. Asci clavate, long-stalked, about 7 0 x 5 / / ,
8-spored; ascospores sausage-shaped, 5 - 7 x 1 . 5 - 2 / / , almost colourless.
On dead branches of many deciduous trees, including Acer, Sambucus and
Ulmus. Fairly common. The perithecia are commonly accompanied by a
slender, black, cylindrical, conidial state which has been called Graphium
fasciculatum Saccardo. The figure shows this conidial state among the peri-
thecial necks.

"303
ENDOXYLA Fuckel

Endoxyla Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 25-26:321


(1871).
Perithecia flask-shaped, immersed singly or in clusters in decorti-
cated coniferous wood or in t h a t of Quercus, asci clavate, long-stalked,
8-spored, ascospores sausage-shaped, non-septate, dark brown. The
genus was described b y its a u t h o r as intermediate between Anthostoma
and Valsa and its systematic position is still not clear.

Endoxyla operculata (Albertini & Schweinitz ex Fries) Saccardo,


Sylloge F u n g o r u m 1:181 (1882). (Fig. 27D.)
Perithecia scattered, occasionally in pairs, blackish, about y 2 m m .
across, immersed in wood, flask-shaped with short stout necks whose
rounded tips slightly p r o t r u d e above the surface of the wood, each
pierced b y a r a t h e r large central ostiole. Asci clavate, about 6 0 x 7 / / ,
8-spored; ascospores sausage-shaped, 1 2 - 1 4 x 3 / / , dark brown. Figured
from Fries' own material distributed as Scleromyceti Sueciae 268 in
1822.

On decorticated wood of Abies and Pinus. Apparently rare. Compare


Apiorhynchostoma, p. 296.

ENCHNOA Fries

Enchnoa Fries, S u m m a Veg. Scand. Sect. P o s t . : 3 9 3 (1849).


Stroma reduced to a m a t of loose dark hyphae enveloping the peri-
thecia, more or less widespread, beneath the surface layers of bark,
perithecia black, globose, without necks; asci clavate, 8-spored; asco-
spores sausage-shaped, slightly coloured.

Enchnoa lanata (Fries) Fries apud Cesati & de Notaris in Comm. Soc.
Crittogamologica Italiana 1 : 2 3 0 (1863). (Fig. 14G.)
Perithecia in swarms beneath the outer layer of bark, their position
often indicated b y a dark grey p a t c h on the inner surface of the b a r k
b e n e a t h the swarm, globose, about 1.5 m m . across, surrounded b y and
seated upon a dense m a t of dark brown hyphae. The overlying b a r k
is slightly raised and u n d u l a t i n g and m a y show small cracks b u t there
are no p r o t r u d i n g ostioles so t h a t the fungus is hard to find unless the
surface of the b a r k is sliced away with a sharp knife or razor blade.

"304
Asci clavate, stalked, about 7// wide in the u p p e r p a r t , 8-spored; asco-
spores sausage-shaped, 9 - 1 1 x 2 . 5 / / , colourless.
On dead branches of Betula. Probably not uncommon.
Echnoa infernalis (Kunze ex Fries) Fuckel in J a h r b . Nass. Vereins f.
N a t u r k u n d e 25-26:302 (1871), common on small dead branches of
Quercus, is more valsoid, with ostioles exposed in clusters in cracks of
the bark. It has brownish ascospores 2 0 - 3 5 x 5 - 7 / / and m a y be mis-
t a k e n for Valsa ambiens.

CALOSPHAERIA Tulasne

Calosphaeria Tulasne, Selecta F u n g o r u m Carpologia 2:108 (1863).


Perithecia without a stroma, seated beneath bark, flask-shaped,
radially arranged with their ostioles converging towards the centre
and erumpent in a cluster through a common crack; asci borne in
clusters on a common stalk, clavate, long-stalked, eight or more
spored, ascospores sausage-shaped, hyaline.

Calosphaeria pulchella (Persoon ex Fries) Schroeter, Kryptogamen


Flora von Schlesien 3 (2): 451 (1897). (Fig. 14H.)
Perithecia aggregated in dense clusters beneath bark, lying on their
sides in several ranks, black, smooth, less t h a n 1 m m . across, each with
a long, slender, somewhat flexuous, cylindrical neck pointing towards
the centre of the cluster, tips of t h e necks erumpent together through
a slit in the bark. Asci club-shaped, stalked, about 4 5 x 5 / / , 8-spored;
ascospores sausage-shaped, about 6 x 1 / / , hyaline. The u p p e r p a r t of
the figure shows a cluster of erumpent ostioles, below is a ring of peri-
thecia exposed b y slicing away the bark.
On dead trunks and branches of Prunus, especially P. cerasus. Not
common.
C. wahlenbergii (Desmazieres) Nitschke, Pyrenomycetes Germanici:
92 (1867), is a very similar species, b u t with larger ascospores, on dead
Betula.
Calosphaeria dryina (Currey) Nitschke, op. cit.:94 (1867), occurs on
Quercus.

Family 11. Diaporthaceae

The 38 genera dealt with under this heading are a somewhat hetero-
geneous residual assemblage of Sphaeriales susceptible, no doubt, to

"305
f u r t h e r subdivision, for example, b y separation of the stromatic and
non-stromatic genera. The reader will, however, probably find it most
convenient to have t h e m keyed out together, with a primary division
based on ascospore characters, which are the easiest to define and to
observe.

I. Perithecia developed in wood or bark of trees or shrubs:


A. Ascospores colourless or yellowish:
1. Ascospores non-septate:
a. Ascospores small, elliptical, appearing pseudoseptate
Ditopclla (p. 319)
b. Ascospores very large:
* Ascospores cylindrical and sausage-shaped
Cryptospora (p. 317)
**Ascospores fusiform . . Cryptosporella (p. 317)
2. Ascospores 1-septate:
a. Perithecia clustered, often with a well developed stroma,
their ostioles erumpent through a stromatic disc or
in a cluster through a crack or hole in the bark;
*Ascospore septum eccentric:
+Perithecia sunk side by side in a wide flat
stroma Anisogramma (p. 321)
+ +Perithecia in small valsoid groups
Apioporthe (p. 310)
**Ascospore septum median:
-fStroma extensive, reddish, at least externally
when young Endothia (p. 322)
+ -fPerithecia in small clusters, stroma externally
greyish or black:
"Ascospores with thread-like appendages
Caudospora (p. 312)
" "Ascospores at most apiculate:
x Blackened zone present in the sub-
strate both above and beneath the
perithecial clusters
Diaporthe (p. 308)
x x Blackened zone present above the
perithecia only or absent:
fAsci still attached to the peri-
thecial wall at maturity,
ostioles erumpent in clu-
sters:
V- C- " §Qstioles erumpent through \\
. ^ \ rf a white disc "pui y v ® VI
\ ' / \ t \F!iIniinnniLi
MManconis I v> 313)
(p. O4O\
§§Ostioles erumpenffhrough
a blackjiisc
Hercospora (p. 314)

"306
§§§Ostioles erumpent through
a crack without a
common stromatic disc
Prosthecium (p. 314)
t f A s c i loose within the perithe-
cium at m a t u r i t y :
§Ostiolar disc strongly de-
veloped
Discodiaporthe (p. 312)
§§Ostiolar disc feebly devel-
oped or perithecia scat-
tered
Cryptodiaporthe (p. 310)
b. Perithecia solitary: Ascospores elliptical. Ascospore
septum median, see Ditopella, Cryptodiaporthe.
3. Ascospores with more than one septum:
a. Ascospores elliptical: Perithecia clustered in a circular
stroma, with convergent ostioles Calosporella (p. 316)
b. Ascospores cylindric-fusiform . . . . Sillia (p. 319)
B. Ascospores brown at maturity, all septate or appearing so:
1. Ascospores 1-septate:
a. Septum median. Perithecia clustered in a circular stroma
*Ascospores fusoid, often with appendages
Melanconiella (p. 315)
**Ascospores broadly elliptical, without appendages
Valsaria (p. 315)
b. Septum cutting off a
hyaline cell at one end
Apiorhynchostoma (p. 296)
2. Ascospores 2-septate,
end cell hyaline

3. Ascospores 3-septate:
a. Perithecium with a clypeus Clypeosphaeria (p. 295)
b. Perithecia clustered, without a clypeus
Prosthecium (p. 314)
4. Ascospores with more than three septa or apparent septa,
perithecia in a stroma:
a. Ascospores truly septate, narrowly cylindrical
Melogramma (p. 318)
b. Ascospores pseudoseptate, elliptic-fusiform
Pseudovalsa (p. 318)

II. Perithecia developed in herbaceous stems, fern rhachides or grass culms:


A. Ascospores hyaline to yellowish:
1. Ascospores non-septate:
a. Stromatic zones present in the substrate as in Diaporthe
Diaporthopsis (p. 311)
b. No stromatic tissue present Phomatospora (p. 320)
2. Ascospores 1-septate:
a. Septum at one end of the spore, perithecia in rows in a
stroma Apiospora (p. 292)

"307
b. Septum median:
*Perithecia enclosed by a black line through the sub-
strate Diaporthe (p. 308)
**Perithecia without a clypeus, beaked:
f Ostiolar papilla at one end of the perithecium
Plagiostoma (p. 324)
fjOstiolar papilla central . . . Sydowiella (p. 320)
cf. also Gnomonia, below.
3. Ascospores very long, slender, multiseptate
Gaeumannomyces (p. 321)
B. Ascospores brown, 3-septate Diaplcella (p. 295)
See also Chitonospora p. 403.

III. Perithecia developed in leaves of Dicotyledons, with or without stro-


mata:
A. Ascospores thread-like, very long, on Salix and Populus
Linospora (p. 325)
B. Ascospores elliptical to needle-shaped:
1. Perithecia developed in a stroma:
a. Ostiole at the end of a long beak, stroma on living leaves
Mamiana (p. 323)
b. Ostiole papillate, on dead and fallen leaves
Hypospilina (p. 323)
2. Perithecia without a stroma:
a. Ascospores septate Gnomonia (p. 324)
b. Ascospores non-septate . . . . Gnomoniella (p. 325)
The first 16 genera are obviously closely interrelated and form the core of
the family.

DIAPORTHE Nitschke

Diaporthe Nitschke, Pyrenomycetes Germanici: 240 (1870).


Perithecia scattered or clustered b u t enclosed within wide-spreading
blackened zones of the substrate, both above the level of the peri-
thecial bodies (dorsal zone) and beneath t h e m (ventral zone). These
stromatic crusts appear as fine black lines when the substrate is cut
at right angles to the surface. Perithecia small, black, commonly in
small groups with their ostiolar beaks erumpent together. Asci clavate
to cylindric-clavate, with a distinct apical ring-like thickening, freed
from the perithecial wall and lying loosely in the cavity at m a t u r i t y ;
ascospores cylindric-fusoid, with one approximately median septum,
often slightly eccentric or curved, hyaline, without appendages.
Diaporthe is a large cosmopolitan genus and some species occur on
a wide range of host plants. Most have conidial states belonging to the
form genus Phomopsis, characterised b y somewhat chambered pycni-

"308
dia embedded in host tissue and yielding innumerable fusiform
hyaline conidia, usually with a small oil drop at each end. Many have
also a second conidial state, the so-called B-spore, thread-like and
often hooked like a walking stick.
For a thorough t r e a t m e n t of this difficult genus see L. E. Wehmeyer,
" T h e genus Diaporthe Nitschke and its segregates", University of
Michigan Scientific Series 9 (1933).

Diaporthe arctii (Lasch) Nitschke, op. cit.:268 (1870). (Fig. 13F.)


Stromatic area wide-spreading, the dorsal zone darkening the sur-
face of the host stem in large dark grey or blackish patches, ventral
zone sunk deeply in the wood. Perithecia scattered or in small groups,
less t h a n y 2 m m . across, black, inserted just beneath the surface,
ostioles rather long, conical, protruding, black. Asci clavate, up to
6 0 x 1 0 / / , 8-spored; ascospores biseriate, fusiform-cylindric, slightly
constricted at the median septum, with two oil drops in each cell,
mostly 1 2 - 1 5 x 2 . 5 - 4 / / , colourless.

On dead herbaceous stems of the woodier type, figured on Arctium, but


according to Wehmeyer also on many other host genera, such as Brassica,
Rumex and various Umbelliferae. Common. Closely related species occur
on Atropa (D. chailletii Nitschke, op. cit.:276 (1870), Daucus and Vinca
(D. eumorpha [Durieu & Montagne] Maire, Myc. Bor. Afr.: 246 [1917]), and
on Epilobium, Euphorbia and Polygonatum (D. pardalota [Montagne]
Nitzschke apud Fuckel in Jahrb. Nass. Vereins f. Naturkunde 23-24:206
[1870]).

Diaporthe eres Nitschke, op. cit.:245 (1870). (PL X X X I V c . )


Stromatic area usually wide-spreading, the dorsal zone usually
blacking the surface of the bark, at least around the ostioles, ventral
zone always present at the margin of the stromatic area and usually
more or less traceable b e n e a t h it. Perithecia in small clusters, within the
bark, raising it into small pimples through the tips of which the clu-
sters of black ostioles emerge. Asci clavate, up to 6 0 x 8 / t , 8-spored;
ascospores biseriate, narrowly fusiform, constricted at the single
median septum, 9 . 5 - 1 4 x 2 . 5 - 4 / / , hyaline.

On dead twigs of very many genera of deciduous trees and shrubs but
figured on the type host, Ulmus. British collections have been recorded also
on Acer, Alnus, Betula, Brassica, Buxus, Corylus, Euonymus, Fagus, Fraxi-
nus, Hibiscus, Ilex, Jasminum, Liriodendron, Lonicera, Lycium, Phillyrea,
Populus, Prunus, Quercus, Bhus, Spiraea, Symphoricarpus, Syringa, Tilia,
Ulex, Veronica and occasionally on conifers. Under the name D. perniciosa

"309
Marchall in Bull. Soc. Roy. Bot. Belg. 54:117 (1921), this common fungus
was formerly regarded as the cause of a serious die-back disease of plum
trees but this is now known to have been an error, it is not an active parasite.
There are about 18 other British species on woody plants.

CRYPTODIAPORTHE P e t r a k
Cryptodiaporthe P e t r a k in Annales mycologici 19:118 (1921).
Similar to Diaporthe b u t entirely without blackened zones in the
substrate. There is often a small stroma developed as a disc about the
ostioles or as a web of hyphae which r u p t u r e s the bark. Perithecia
scattered or in small groups, asci and ascospores as in Diaporthe.

Cryptodiaporthe salicella (Fries) Petrak, op. cit.: 182 (1921). (PI.


XXXIVB.)
Perithecia scattered or a few together, slightly raising and splitting
the bark through which the slender black ostioles emerge, b o d y of the
perithecium small, about % m m . across, just below the surface of the
bark. Asci clavate, up to 5 0 x 1 2 / / , 8-spored; ascospores biseriate,
narrowly fusiform and only slightly constricted at the septum,
15-22 x 2-2.5//, hyaline. There is no trace of a blackened zone either
above or below the perithecia b u t there m a y be a small greyish stroma-
tic cushion around the ostioles.
On dead twigs of Salix. Not very common.

Cryptodiaporthe salicina (Gurrey) Wehmeyer in Univ. Michigan Sci.


I X : 194 (1933), on the same host genus and on Populus, differs in its
wider ascospores, 1 5 - 2 0 x 4 . 5 - 7 . 5 / / .
There are about six other British species.

APIOPORTHE von Hohnel

Apioporthe von Hohnel in S.B. K. Akad. Wiss. Wien, Math.-Nat.


Klasse Abt. 1, 126:381 (1917).
Similar to Diaporthe b u t with less developed black zones in the host
tissue and with the septum near one end of the ascospore.

Apioporthe vepris (de Lacroix) Wehmeyer in Univ. Michigan Sci. I X :


221 (1933). (PI. X X X I V F . )
Perithecia in small dense clusters within the bark, usually forming
small elongated slits in its surface to expose blackened discs containing

"310
the protruding ostioles. Perithecia up to 0.2 m m . across, black. Asci
clavate, up to 40x8/<, 8-spored; ascospores biseriate, ellipsoid, nar-
rowed towards the lower end at which the septum develops, 5-10 x
2-2.5 /i, with two oil drops in the larger cell and usually one in the
smaller. There m a y be a slight bristle-like appendage at each end of
the spore.
On dead stems of Rubus. Common.

DIAPORTHOPSIS F a b r e

Diaporthopsis Fabre in Ann. Sci. Nat. Bot. VI, 15:35 (1883).


Perithecia embedded in superficial tissues of the host, which are
often blackened externally as in Diaporthe, perithecia scattered, se-
parately erumpent. Asci and ascospores as in Diaporthe, except t h a t
no septum develops in the spore.

Diaporthopsis angelicac (Berkeley) Wehmeyer in Univ. Michigan Sci.


I X : 2 2 8 (1933). (PL X X X I V D . )
Perithecia scattered, black, about % m m . across, ostioles short, only
just erumpent and not at all conspicuous; host surface eventually be-
coming greyish or somewhat violaceous. Asci clavate at first, t h e n
cylindrical, about 5 0 x 7 f i , 8-spored; ascospores biseriate or uniseriate,
elliptic-fusoid, 9 - 1 5 x 4 - 7 / / .
The habit of this inconspicuous species was well described over a
century ago by Desmazieres, who knew it as Sphaeria berkeleyi. He
wrote (translated) " T h e Sphaeria of Berkeley is, without doubt, one
of the most curious in t h e genus; it is also one of the smallest and its
position renders it one of the hardest to find, when one has not seen it
before. Its perithecia, which are only visible under a good lens, are
usually sunk in the tissue of the host, either in t h e wood or between
it and the pith. . . . T h e y are black, globose and a little depressed,
topped b y a straight ostiole, the length of which depends on the thick-
ness of the host tissue it has to traverse. W h e n the ostiole is short it is
somewhat pyramidal, when long it is cylindrical. The tip of t h e ostiole
does not always reach the surface b u t when it does it appears there,
even under a lens, only as a very minute black dot, scarcely percep-
tible." He goes on to say t h a t to find the perithecia it is necessary to
split t h e stems longitudinally and examine the broken edges under a
lens.

"311
On dead stems of Umbelliferae, typically on Angelica but also on Daucus,
Eryngium, Foeniculum, Heracleum and Peucedanum. Common and not to be
confused with Diaporthe arctii, in which the septum may be slow to develop
in the ascospores, but which has rather larger perithecia, a blacker dorsal
zone and usually a black ventral zone such as is lacking in D. angelicae.

Diaporthopsis pantherina (Berkeley) Wehmeyer, op. cit.: 232 (1933)


occurs on dead Pteridium aquilinum.

CAUDOSPORA Starback

Caudospora Starback in Bihang t . K. Svenska Vet.—Akad. Handl. 15


(3), No. 2 : 1 1 (1889).
Perithecia like those of a Diaporthe, to which genus the species is
commonly assigned, without a dorsal zone b u t with a strongly
developed ventral zone surrounding each cluster of perithecia. Asci
not becoming freed from the perithecial wall, 8-spored; ascospores
1-septate, with lateral and terminal appendages.

Caudospora taleola (Fries) Starback, op. cit.: 11 (1889). (PI. XXXIVA.)

Perithecia 0.4 m m . across, in small clusters within the bark and


raising it into small pustules which crack at their tips to expose small,
black, ostiolar discs; ventral black zone well developed, completely
enclosing each cluster of perithecia and sometimes visible as a slight
ridge where its u p t u r n e d edge reaches the surface of the bark. Asci
cylindrical, up to 150x16//, with four or eight spores; ascospores
elliptical b u t distinctly constricted at the septum, 1 7 - 2 5 x 7 - 9 . 5 / / ,
hyaline, with a cylindrical hyaline appendage 6 - 1 0 x 1 - 1 . 5 / / at each
end and two or three similar b u t longer appendages attached at the
level of the septum.

On dead branches of Quercus, on which it is said to be parasitic. Not un-


common.

DISCODIAPORTHE P e t r a k

Discodiaporthe P e t r a k in Hedwigia 62:293 (1921).


Differs from Diaporthe in having neither dorsal nor ventral blacken-
ed zones, from Cryptodiaporthe in its well developed disc and from
Melanconis, with which it is usually united, in the asci becoming loose
in the perithecium.

"312
Discodiaporthe sulphurea (Fuckel) P e t r a k , op. cit.:294 (1921). (PI.
XXXIVE.)
Perithecia up to % m m . across, in compact clusters of five to ten
within the bark and raising it into small pustules which crack to ex-
pose the strongly developed stromatic discs, dotted with the black
ostioles. Asci clavate, up to 110x15/t, 8-spored; ascospores biseriate
elliptic-fusiform, with a single median septum, 17.5-27 x 6-9.5,«, r a t h e r
thick-walled, with a t i n y pointed appendage at each end, hyaline. The
surface of the disc is sometimes somewhat yellowish.

On dead branches of Corylus. Not common. Also known as Melanconis


flavovirens (Otth) Wehmeyer, in Mycologia 29:602 (1937).

MELANCONIS Tulasne

Melanconis Tulasne in Ann. Sci. Nat. Bot. IV, 5 : 1 0 9 (1856).


Perithecia clustered within bark, without associated blackened
zones b u t with a well developed stromatic disc enclosing the ostioles,
usually white, grey or greenish at the surface. Asci cylindric-clavate,
ascospores 1-septate, hyaline or nearly so. According to Wehmeyer
the asci become detached from the perithecial wall, according to Munk
t h e y do not.

Melanconis stilbostoma (Fries) Tulasne, op. cit.: 109 (1856). (PI.


XXXIIIE.)
Perithecia black, about % m m . across, in circular clusters of 5 - 1 5 to
each pustule, splitting the bark to expose the massive whitish stroma-
tic discs, dotted with the black ostioles. Asci narrowly clavate, up to
120x16/i, 8-spored; ascospores irregularly biseriate, narrowly ellip-
tical, mostly 1 5 - 2 0 x 5 - 8 / / , slightly constricted at the septum, r a t h e r
thick-walled, hyaline.

On twigs and branches of Betula. Common.


Two species of Melanconis occur on Alnus, M. Alni Tulasne in Ann. Sci.
Nat. Bot. IV, 5:109 (1856), with ascospores 13-22x3.5-7// having minute
appendages or none at all and M. thelebola (Fries) Saccardo, Sylloge Fungo-
rum 1:605 (1882), in which the ascospores are 24-42x8-10// with long
slender appendages at each end.

Melanconis chrysostroma (Fries) Tulasne, Selecta F u n g o r u m Carpo-


logia 2:125 (1863), occurs on Carpinus.

"313
Melanconis modonia Tulasne in Ann. Sci. Nat. Bot. IV, 5 : 1 1 1 (1856),
on Castanea has ascospores which become brown with age and differs
from Pseudovalsa only in its 1-septate spores.

HERCOSPORA Fries

Hercospora Fries, S u m m a Veg. Scand. Sect. Post.:397 (1849), emend


P e t r a k in Annales mycologici 21:319 (1923).
Perithecia in small compact valsoid clusters, with a well developed
stroma surrounding their necks and carried down laterally where it is
defined b y a black line in the b a r k ; asci cylindrical, ascospores ellip-
tical, 1-septate, hyaline. Differs from Melanconis in its better devel-
oped stroma and from Diaporthe in the stroma being limited to t h e
individual pustule and in the short-stalked cylindrical asci.

Hercospora tiliae (Persoon ex Fries) Tulasne, Selecta F u n g o r u m Car-


pologia 2:154 (1863). (PI. XXXIVG.)
Perithecia about 0.4 m m . across, black, flask-shaped, in clusters of
two to eight in small s t r o m a t a immersed in b a r k ; stroma cushion-
shaped, with a conical grey upper p a r t which emerges from a crack in
the b a r k as a dark grey disc dotted with the ostioles, in the lower p a r t
delimited b y a black line. Asci cylindrical, short-stalked, up to 100x
12/z, mostly 8-spored; ascospores uniseriate, elliptic-cylindric, 12-18x
6 - 8 / / , w i t h a median septum b u t not noticeably constricted, hyaline.

On dead twigs of Tilia. Not uncommon.

PROSTHECIUM Fries

Prosthccium Fries, S u m m a Veg. Scand. Sect. Post.: 416 (1849).


Perithecia in small clusters within bark, without a distinct stroma
b u t with their ostioles erumpent together from a crack in the surface;
asci clavate, 8-spored, ascospores large, usually 1-septate, sometimes
with more septa when old, hyaline or browning with age, with append-
ages at each end.

Prosthecium auctum (Berkeley & Broome) P e t r a k in Annales myco-


logici 2 1 : 3 2 5 (1923). (PI. X X X I I I J . )
Perithecia in clusters of three to seven, rather large, up to 0.8 m m .
across, black, with short, stout, conical ostioles erumpent from cracks

"314
in the bark. Asci broadly clavate, short-stalked, up to 110x36//,
8-spored; ascospores elliptic-cylindric, in two or three rows in the
ascus, 2 7 - 3 8 x 1 0 - 1 4 / / , r a t h e r thick-walled, with one median septum,
scarcely constricted, bearing a small rounded appendage at each end,
hyaline b u t occasionally becoming brown and 3-septate when old or
after extrusion from the ascus.

On dead branches of Alnus. Common.


The systematic position of this species raises considerable difficulties; it
has been placed in Pseudovalsa because of the colour and septation of the
over-ripe ascospores, Wehmeyer refers it to Melanconis and reserves
Prosthecium for species with hyaline or coloured multiseptate ascospores,
separated from Pseudovalsa by their conidial states belonging not to Cory-
neum but to Slilbospora and Steganosporium.

MELANCONIELLA Saccardo

Melanconiella Saccardo, Sylloge F u n g o r u m 1:740 (1882).


Perithecia in clusters within bark, embedded in a feebly developed
stromatic cushion which emerges as an ostiolar disc, asci cylindric-
clavate, short-stalked, 8-spored, ascospores 1-septate, brown. Like
Melanconis but with brown ascospores.

Melanconiella spodiaea (Tulasne) Saccardo, op. cit.:740 (1882). (PI.


XXXIIIF.)
Perithecia densely crowded in clusters of up to 20 in a small powdery
stroma within the bark, erumpent as a dark grey disc dotted with the
ostioles. Asci cylindric-clavate, short-stalked up to lOOx 16/<, 8-spored;
ascospores 1-2-seriate, elliptic-fusiform, constricted at the median
septum, 1 6 - 2 0 x 7 - 9 / / , olive-brown, sometimes with a minute hyaline
appendage at each end.

On dead twigs of Carpinus betulus. Uncommon.

VALSARIA Cesati & de Notaris

Yalsaria Cesati & de Notaris in Commentario Societa Crittogamologica


Italiana 1 : 2 0 5 (1863).
Pseudothecia clustered within a stroma embedded in bark, super-
ficially like a Valsa; asci cylindrical, 8-spored, ascospores uniseriate,
1-septate, brown.

"315
ValsariainsitivaCesati& de Notaris, op. cit.: 205 (1863). (PI. X X X V I I H ) .

Pseudothecia flask-shaped, with long cylindrical necks, black, up


to y 3 m m . across, in small groups clustered within a circular brownish
stroma with a t h i n black limiting layer and erumpent by a dark grey
disc on which the clusters of ostioles appear as minute black papillae.
Asci cylindrical, short-stalked, up to 130x14//, the wall thickened at
the tip, 8-spored; ascospores uniseriate, broadly elliptical and strongly
constricted at the single median septum, brown, 1 6 - 1 8 x 9 - 1 0 / / .

On dead branches of many deciduous trees, figured on Quercus.

CALOSPORELLA Schroeter

Calosporella Schroeter, K r y p t o g a m e n Flora von Schlesicn 3 (2): 442


(1897).
Perithecia in clusters within the bark, surmounted b y a stromatic
disc b u t without any enveloping black zone, asci clavate, not becoming
loose in the perithecium, ascospores 3-septate, hyaline to very pale
brown.

Calosporella innesii (Gurrey) Schroeter, op. cit.: 442 (1897). (PI.


XXXIIIH.)

Perithecia about y 2 m m . across, black, in dense clusters of up to 20


in small grey cushion-like s t r o m a t a without any black delimiting zone,
erumpent b y grey discs studded with the black ostioles. Asci broadly
clavate, up to 100x20/i, 8-spored; ascospores in two or three rows,
elliptic-cylindric, 2 0 - 3 9 x 6 - 9 / / , 3-septate, scarcely constricted at the
septa, hyaline, occasionally with a minute tapering hyaline append-
age at each end.

On dead twigs of Acer pseudoplatanus in winter. Common.


There is considerable confusion over both the name and the generic posi-
tion of this species and it will be encountered in the literature under the
names Calospora innesii (Currey) Saccardo, Sylloge Fungorum 2:231 (1883),
Prosthecium innesii (Currey) Wehmeyer in Univ. Michigan Studies Sci. Ser.
14:98 (1941), Calospora platanoides (Persoon) Niessl in Hedwigia 14:130
(1875), Pseudovalsa platanoides (Persoon) Winter, Rabenhorst's Krypt.
Flora 1 (2):790 (1887), and Calosporella platanoides (Persoon) von Hohnel
in Annales mycologici 16:116 (1918). It is a typical species of Prosthecium
in the interpretation of WTehmeyer.

"316
CRYPTOSPORA Tulasne

Cryptospora Tulasne, Seleeta F u n g o r u m Carpologia 2:144 (1863).


Perithecia embedded in bark in small dense clusters, with a feebly
developed stroma which m a y appear only as a discoloured area of
bark, without black zones, asci cylindric-clavate, not loose in the peri-
thecium, ascospores non-septate, cylindrical, very large, hyaline.

Cryptospora suffusa (Fries) Tulasne, ip. cit.: 145 (1863). (PL XXXIIIG.)

Perithecia inserted in groups of up to ten, surrounded b y lighter-


coloured portions of bark, forming distinct pustules at the tips of
which the black ostioles break through, without an enveloping stro-
matic disc. Asci cylindric-clavate, up to 9 0 x 2 0 / / , 8-spored; ascospores
irregularly arranged, narrowly cylindrical with rounded ends and
usually somewhat curved, 5 0 - 6 0 x 4 - 5 / / , hyaline, packed with small
oil drops.

On twigs of Alnus in winter and spring. Common.

Cryptospora corylina (Tulasne) Fuckel in J a h r b . Nass. Vereins f.


N a t u r k u n d e 23-24:192 (1870), occurs on Corylus, with ascospores up
to 1 0 0 x 6 / / .
Cryptospora betulae Tulasne, op. cit.: 149 (1863), with ascospores
3 3 - 3 4 x 3 . 5 - 4 / / occurs on Betula.

CRYPTOSPORELLA Saccardo

Cryptosporella Saccardo in Michelia 1 : 3 0 (June 1877).


Similar to Cryptospora, with which it is sometimes united, b u t with
pointed fusiform ascospores.

Cryptosporella hypodermia (Fries) Saccardo, op. cit. 1 : 3 0 (1877).


(PL X X X I I I K . )
Perithecia about % m m - across, black, inserted in bark in small dense
clusters of up to ten, without a stroma b u t w i t h their ostioles erum-
pent together in a compact group which projects a little above the sur-
face. Asci cylindrical, short-stalked, up to 120x20//, 8-spored; asco-
spores irregularly biseriate, fusiform with pointed ends, often asym-
metrical, 2 0 - 6 0 x 8 - 9 / / hyaline or nearly so, containing small oil drops
and granules.

On dead twigs of Ulmus in winter and spring. Not uncommon.

"317
Cryptosporella umbrina (Jenkins) Jenkins & Wehmeyer in P h y t o -
pathology 25:888 (1935), with ascospores only 8 - 1 1 x 2 . 5 - 4 / / , is oc-
casionally encountered as a cause of canker in cultivated roses.

MELOGRAMMA Fries

Melogramma Fries, S u m m a Veg. Scand. Sect. Post. : 386 (1849), emend


Tulasne, Selecta F u n g o r u m Carpologia 2 : 84 (1863).
Perithecia inserted in a small b u t well developed stroma, asci clavate,
short-stalked, ascospores elongated, multiseptate, brownish b u t with
t h e terminal cells colourless.
<

Melogramma spiniferum (Wallroth) de Notaris in Comm. Soc. Critto-


gamologica Italiana 1 :203 (1863). (Fig. 26J.)
Stroma up to 3 m m . across, black, hemispherical, soon breaking
through the jbark to expose the whole of its upper surface, containing
up to 12 flask-shaped perithecia whose stout cylindrical black ostioles
project well above the surface of the stroma. Asci narrowly clavate,
up to 220x18//,. 8-spored, with a distinct apical ring; ascospores ir-
regularly biseriate, narrowly cylindrical with rounded ends, 55-75x
7-8/«,'7-septate, the terminal cells hyaline, t h e remainder light brown.

On bark of Fagus, often in densely crowded swarms, especially at the


base of trunks and on the exposed parts of the larger roots. Common.

Melogramma vagans de Notaris, Micromycetes Italici 9 : 2 (1855),


with smaller 3-septate ascospores, occurs on dead branches of Car-
pinus.

rSEUDOYALSA Cesati & de Notaris

Pseudovalsa Cesati & de Notaris in Comm. Soc. Crittogamologica Ita-


liana 1:206 (1863).
Similar to Melogramma b u t with more broadly fusiform ascospores
which are only falsely septate.

Pseudovalsa lanciformis (Fries) Cesati & de Notaris, op. cit.: 206 (1863).
(Fig. 26 K . )
S t r o m a cushion-shaped, up to 3 m m . across, containing up to ten
perithecia each about % m m . wide, erumpent b y a slit in the bark to
expose a black lancet-shaped disc on which the ostioles open without

"318
noticeably projecting. Asci cylindric-clavate, short-stalked, up to
2 0 0 x 3 0 / / , 8-spored; ascospores biseriate, narrowly ellipsoidal, not
constricted, 3 0 - 5 0 x 1 1 - 1 6 / / , wall thick, olive-brown b u t colourless at
the ends of the spore, contents with four or six large oil bodies b u t ap-
parently without t r u e septa.
On twigs and small branches of Betula. Not uncommon. According to
Wehmeyer P. umbonata (Tulasne) Saccardo in Atti Soc. Yenet. Trent. Sci.
Nat. 4:91 (1875), on Quercus is probably the same fungus.

Pseudovalsa longipes (Tulasne) Saccardo, op. cit.:91 (1875), which


differs in its narrower up to 10-celled ascospores, 3 5 - 8 0 x 6 - 1 1 / / , also
occurs on Quercus.

SILLIA Karsten

Sillia Karsten, Mycologia Fennica 2 : 2 0 (1873) (as " H i l l i a " , corrected


to Sillia p. 251).
Perithecia inserted at different levels in a well defined cushion-like
stroma with a well-developed apical disc, ascospores very long and
narrow, multiseptate, hyaline.

Sillia ferruginea (Persoon ex Fries) Karsten, op. cit.: 159 (1873). (PI.
XXXIVH.)
Stroma up to 4 m m . long, somewhat conical with a broad flat base,
flesh bright r u s t y brown, enclosing numerous, scattered, black, flask-
shaped perithecia up to 0.4 m m . across, upper portion of the stroma
erumpent as a black powdery disc, from which project the numerous
black conical ostioles. Asci oblong, sessile, up to 112x16//, 8-spored,
with a small apical ring; ascospores more or less parallel, very nar-
rowly fusiform and often somewhat curved, 6 0 - 7 5 x 3 / / , a b o u t 5-sep-
t a t e , hyaline, containing numerous small oil drops.

On dead twigs of Corylus. Not uncommon.

DITOPELLA de Notaris

Ditopella de Notaris, Sferiacei Italici: 42 (1863).


Perithecia inserted singly in bark, with short stout ostioles, asci be-
coming loose in the cavity, with numerous spores, ascospores non-sep-
t a t e or falsely 1-septate, hyaline.

"319
Ditopella ditopa (Fries) Schroeter, Kryptogamen Flora von Schlesien 3
(2):388 (1897). (PI. X X X I I I D . )
Perithecia black, up to 0.8 m m . across, singly within bark and raising
it into small pustules which are pierced b y the short black ostioles.
Asci cylindric-clavate, sessile, up to 9 0 x 1 6 / / , polysporous; ascospores
elliptic-cylindric, 1 3 - 1 6 x 2 - 3 . 5 / / , non-septate or with an .obscure p a r t -
ing across the middle, hyaline, sometimes with a minute appendage at
each end.
On twigs of Alnus in winter and spring. Common.

SYDOWIELLA P e t r a k

Sydowiella P e t r a k in Annales mycologici 21:30 (1923).


Perithecia scattered, without a stroma, embedded in bark or cortex,
with short erumpent ostioles, asci subcylindrical, short-stalked, with
an apical ring-like thickening, ascospores 1-seriate, hyaline, with one
median septum. The species superficially resemble those of Didymella
in the Loculoascomycetes b u t have thin-walled asci.

Sydowiella fenestrans (Duby) Petrak, op. cit.:30 (1923). (Fig. 261.)


Perithecia scattered, subglobose, up to 0.7 m m . across, black, sub-
epidermal, each with a short, cylindrical, erumpent, protruding ostiole.
Asci cylindrical, up to 120x15//, subsessile, 8-spored; ascospores
1-seriate, elliptical to bean-shaped, 1 9 - 2 4 x 8 - 9 / / , hyaline, with a
median septum and usually slightly constricted at t h a t level.
On dead stems of Chamaenerion angustifolium, in spring. Common.

PHOMATOSPORA Saccardo
Phomatospora Saccardo in Nuovo Giornale Botanico Italiano 7-306
(1875).
Perithecia scattered, subepidermal, not beaked, without a stroma,
r a t h e r thin-walled; asci cylindrical, short-stalked, with a small apical
ring-like thickening, ascospores ellipsoidal, usually biguttulate, hya-
line, non-septate.

Phomatospora berkeleyi Saccardo, op. cit.:306 (1875). (Fig. 26H.)


Perithecia scattered, globose, up to 0.2 m m . across, blackish, ostiole
short, perforating the epidermis and appearing as a minute dot on the

"320
outer surface. Asci narrowly cylindrical, thin-walled, short-stalked, up
to 9 0 x 6 / t , 8-spored; ascospores uniseriate, elliptic-cylindric, 6 - 1 1 x
2 - 3 / L i , hyaline, with a conspicuous oil drop at each end.

Figured from the type collection, on dead stems of Solanum tuberosum in


spring, but said to occur commonly on dead herbaceous stems and grass
culms of all kinds. This inconspicuous little species is very similar in habit to
Diaporthopsis angelicae with which it was, in fact, compared by its dis-
coverer.

GAEUMANNOMYCES von Arx & Olivier

Gaeumannomyces von Arx & Olivier in Trans. Brit, mycol. Soc. 35:32
(1952).
Perithecia scattered, subepidermal, more or less beaked, on culms of
grasses and sedges, asci subcylindrical, with a minute apical ring,
8-spored, ascospores very long and slender, multiseptate, hyaline. The
genus is superficially similar to Ophiobolus in t h e Loculoascomycetes.

Gaeumannomyces graminis (Saccardo) von Arx & Olivier, op. cit:32


(1952). (Fig. 26L.)
Perithecia scattered within the basal discoloured portions of dead
grass culms and surrounded b y a sparse network of dark brown to
blackish hyphae, perithecia black with short, conical, erumpent, black
beaks. Asci narrowly cylindrical, short-stalked, 8-spored, about 110 x
13/t; ascospores very narrowly cylindric-fusiform, lying parallel in
the ascus, about 7 0 - 1 0 0 x 4 / / in the typical variety, 8 5 - 1 3 0 n long in
t h e var. avenae, multiseptate, hyaline.
On basal parts of culms of grasses and cereals, in summer. Not uncommon.
The typical variety causes a serious disease, "Take-all" or "Whiteheads"
of wheat and barley but does not attack oats; the var. avenae (E. M. Turner)
Dennis* attacks all three cereals as well as various wild grasses.

ANISOGRAMMA Theissen & Sydow

Anisogramma Theissen & Sydow in Annales mycologici 14:451 (1916).


Perithecia embedded in an extensive stroma within bark, their
ostioles opening individually, asci clavate, thin-walled, ascospores
ovoid, hyaline, with a single septum near the lower end.

* Ophiobolus graminis var. avenae Turner in Trans. Brit, mycol. Soc. 24:
279 (1940).

"321
Anisogramma virgultorum (Fries) Theissen & Sydow, op. cit.:451
(1916). (Fig. 27B.)
Stroma an elongated crust, seated on the wood and exposed b y rup-
ture of the bark, dark brown to blackish, containing numerous flask-
shaped perithecia whose ostioles open separately as small papillae on
the stroma surface. Asci clavate, up to 7 5 x l 2 / t , 8-spored, thin-walled;
ascospores biseriate, ovoid, the broad end uppermost in the ascus, with
a single septum near the narrower lower end, 10-12x4-5/«, hyaline.

On twigs of Betula. Apparently rare, I have seen no British material and


have figured the species from Fries own material, distributed in 1819 as
Scleromyceti Sueciae number 2. It is said to occur on still living twigs so
may be a parasite like Endothia.

E N D O T H I A Fries
Endothia Fries, S u m m a Veg. Scand. Sect. P o s t . : 3 8 5 (1849).
Perithecia flask-shaped with long slender necks, embedded in a
massive reddish stroma, asci and ascospores similar to those of Dia-
porthe.

Endothia parasitica (Murrill) P. J. & H. W. Anderson in P h y t o p a t h o -


logy 2 : 2 1 0 (1912). (PI. X X X I V J . )
Stroma a cushion or elongated crust, erumpent through cracks in
the b a r k , with a reddish outer surface, perithecia up to 60 in a stroma,
usually from 15 to 30, seated towards its base, flask-shaped, up to
0.4 m m . across, with long, slender, somewhat undulating necks which
emerge as small papillate ostioles on the stroma surface. Asci cylindric-
clavate, up to 5 0 x 9 / t , thin-walled with a well developed apical ring-
like thickening, 8-spored; ascospores biseriate or uniseriate, elliptical,
9 - 1 0 x 4 - 5 fi, usually constricted at the single m e d i a n septum, hyaline.

On bark of Castanea, associated with a pycnidial state containing hyaline


non-septate conidia 3-4 x 1-1.5 fi.
This is the only fungus described in the present volume which has not
yet been reported to occur in the British Isles. It is included because of its
great economic importance and because there is a possibility of its intro-
duction into this country from Europe. The species is apparently native to
China, where it causes cankers on Castanea mollissima, and was introduced
into North America about 1904. There it spread with startling rapidity and
has since destroyed the formerly extensive forests of Castanea dentata in
the eastern United States. In recent years it has appeared in Europe and
spread through Spain, Italy, Switzerland and Jugoslavia, causing a destruc-
tive disease of the European sweet chestnut, Castanea vesca.

"322
HYPOSPILINA (Saccardo) Traverso

Hypospilina (Saccardo) Traverso Flor. Ital. crypt. 1:495 (1913).


Perithecia inserted singly or in small groups within a black stroma
in tissue of fallen leaves, not beaked, asci clavate, ascospores hyaline
1-3-septate.

Hypospilina bifrons (De Candolle) Traverso, op. cit.:495 (1913).


(Fig. 26B.)
Stroma small, circular or occupying an angular area delimited b y
leaf veins, black, embedded in the leaf tissue, containing from one to
three subglobose perithecia, about % m m . across with minute papillate
ostioles. Asci clavate, about 5 0 x 1 0 / / , 8-spored; ascospores biseriate,
rather broadly fusiform with a septum cutting off a small cell near the
lower end, 1 1 - 1 3 x 4 / / , hyaline.

On dead leaves of Quercus in spring. Not uncommon.

MAM IANA Cesati & de Notaris

Mamiana Cesati & de Notaris in Commentario Societa Crittogamo-


logica Italiana 1:210 (1863).
Stroma well developed in tissue of leaves, containing several long-
beaked perithecia, asci cylindric-clavate, ascospores non-septate to
1-septate, hyaline.

Mamiana fimbriata (Persoon ex Fries) Cesati & de Notaris, op. cit.: 211
(1863). (Fig. 26A.)
Stroma developed as a black thickened spot up to 4 mm. across on
living leaves, perithecia up to 25 in a stroma, flask-shaped, a b o u t
0.3 m m . across, each with a long cylindrical black neck erumpent from
the under surface of the leaf. Asci cylindric-clavate, up to 5 0 x 1 0 / / ,
with a well developed apical ring, 8-spored; ascospores broadly ellip-
tical, 8 - 1 1 x 3 - 4 / / , with one septum cutting off a small cell near the
lower end, hyaline.

On leaves of Carpinus betulus rare; the perithecia mature in the following


spring, on the fallen leaves.

Mamiana coryli (Batsch ox Fries) Cesati & de Notaris, op. cit.:211


(1863), is a similar fungus with perithecia solitary or in much smaller

"323
groups in living leaves of Corylus, with ascospores similar b u t non-
septate.

GNOMONIA Cesati & de Notaris

Gnomonia Cesati & de Notaris in Commentario Societa Crittogamo-


logica Italiana 1:231 (1863).
Perithecia scattered, without a stroma, in tissue of leaves or herba-
ceous stems, long-beaked, asci small, clavate, becoming loose, asco-
spores 1-septate, hyaline.

Gnomonia vulgaris Cesati & de Notaris, op. cit.:232 (1863). (Fig. 26E.)
Perithecia subglobose, dark brown, up to % m m - across, thin-walled
and collapsing when dry, embedded in dead leaf tissue, each with a
long, very slender, bristle-like beak erumpent from the under side of
the leaf, the ostiole surrounded by a fringe of short whitish cells. Asci
somewhat clavate, up to 3 5 x 8 / / , with a minute apical ring, 8-spored;
ascospores lying parallel in the ascus, very narrowly fusiform, 17-22 x
1.5//, with a single median septum, hyaline.

On fallen leaves of Corylus, in spring. Common; also known as Gnomonia


gnomon (Tode ex Fries) Schroeter, Kryptogamen Flora von Schlesien 3 (2):
390 (1897).

PLAGIOSTOMA Fuckel

Plagiostoma Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23/24:118


(1870).
Perithecia immersed in dead stems or leaves, like those of a Gno-
monia except t h a t the neck is situated at one end instead of in the
middle of the upper surface.

Plagiostoma devexa (Desmazieres) Fuckel, op. cit.: 119 (1870). (Fig.


20H.)

Perithecia scattered, subepidermal, elliptical and somewhat flatten-


ed, about % m m . long, neck black, at one end, short and erumpent
through t h e host epidermis. Asci elliptical, about 4 0 x 8 / / , 8-spored;
ascospores narrowly elliptical, 1-septate, not constricted, hyaline,
8 - 1 0 x 2 . 5 - 3 / / , with a short curved appendage at each end.
On dead stems of Polygonum in winter.

"324
Plagiostoma pustula (Persoon ex Fries) von Arx in Antonie v.
Leeuwenh. 17:264 (1951), on dead oak leaves, differs in its longer
ascospores, 17-23x4/« with either one median septum or three equally-
spaced septa. The legitimate name of the species would be Phoma
pustula Fries, Systema mycologicum 2:546 (1823), b u t this name has
been rejected to permit the conventional use of Phoma as the name for
a form-genus of pycnidial fungi.

GNOMONIELLA Saccardo

Gnomoniella Saccardo in Michelia 2:312 (March 1881).


Similar to Gnomonia b u t with non-septate ascospores and usually
shorter stouter beaks to the perithecia.

Gnomoniella tubiformis (Fries) Saccardo, Sylloge F u n g o r u m 1 : 4 1 3


(1882). (Fig. 26D.)
Perithecia scattered, embedded in the mesophyll and causing a
distinct swelling on b o t h surfaces of the leaf, brown, about % m m .
across, somewhat flattened, each with a subcylindrical brown beak,
usually shorter t h a n the diameter of the perithecium. Asci clavate,
becoming loose, up to 70x15/«, with a well developed apical ring,
8-spored; ascospores irregularly biseriate, somewhat lemon-shaped or
flattened on one side, 13-17x5-6/«, hyaline, slightly olivaceous in the
mass.

On fallen leaves of Alnus, in spring. Common.

LINOSPORA Fuckel
Linospora Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:123
(1870).
Perithecia inserted in small stromata, usually singly b u t sometimes
in pairs, in dead leaf tissue, long-beaked, asci cylindrical, ascospores
thread-like, hyaline.

Linospora capreae (de Candolle) Fuckel, op. cit.: 124 (1870). (Fig. 26c.)
Stroma small, subcircular, black, occupying the thickness of t h e
mesophyll and forming a black spot on the upper surface of the leaf,
concealed by the hairs on the lower surface, containing usually one
perithecium, about 0.4 m m . across, with a long cylindrical black beak

"325
attached at one end and erumpent among the hairs of the lower sur-
face of the leaf. Asci cylindrical, lying horizontally in the perithecium,
up to 160x9//, with a minute apical ring. 8-spored; ascospores lying
parallel in the ascus, thread-like up to 120x2/«, hyaline.
On dead leaves of Salix caprea and its allies, in spring and summer.
Common.

Linospora ceuthocarpa (Fries) Munk, Dansk. Bot. Arkiv. 15 (2): 76


(1953), forms gregarious black spots on fallen leaves of Populus
tremula.

Family 12. Ceratostomataceao

This small family contains genera with smooth carbonaceous peri-


thecia, more or less sunk in rotten wood b u t without stromata, with
well developed often very long beaks and colourless ascospores. The
genera are separated on ascospore characters:

I. Ascospores non-septate Ceratostomella


II. Ascospores with several cross septa . . . . Ceratosphaeria (p. 327)
III. Ascospores with transverse and longitudinal septa, budding off in-
numerable minute secondary spores within the ascus
Rliamphoria (p. 327)

CERATOSTOMELLA Saccardo

Ceratostomella Saccardo in Michelia 1:370 (November 1878).


Perithecia often gregarious but not within a common stroma, flask-
shaped, black, immersed in rotten wood, with long erumpent beaks,
asci cylindric-clavate, thin-walled, ascospores elliptical, hyaline, non-
septate.

Ceratostomella ampullasca (Cooke) Saccardo, Sylloge Fungorum 1 : 4 0 9


(1882). (Fig. 13H.)
Perithecia scattered, partially embedded in rotting wood, black,
flask-shaped, about % m m . across, with long, slender, often undulating,
cylindrical, black beaks protruding above the surface. Asci clavate,
stalked, up to 8 0 x 1 2 / / , 8-spored; ascospores irregularly biseriate,
elliptic-cylindric, 1 0 - 1 2 x 3 - 4 . 5 / / , with a large oil drop near the end,
hyaline.

"326
On rotting wood of Quercus and Acer. Common. Ceratostomella ligneola
(Berkeley & Broome) Cooke in Grevillea 17:49 (March 1889), is said to be an
earlier name for this species.

CERATOSPHAERIA Niessl
Ccratosphaeria Niessl in Verh. Naturforsch. Ver. Brunn. 14:208 (1876).
Perithecia like those of Ceratostomella b u t ascospores multiseptate.

Ccratosphaeria lampadophora (Berkeley & Broome) Niessl, op. cit.


(1876). (Fig. 13i.)
Perithecia gregarious, black, lower portion subglobose, about % m m
across, more or less immersed in r o t t e n wood, beak long and stout,
often curved, protruding. Asci cylindric-clavate, up to 130x12//,
8-spored; ascospores biseriate, fusiform, 5 5 - 6 2 x 4 / i , multiseptate,
hyaline.

RHAMPHORIA Niessl

Rhamphoria Niessl in Verh. Naturforsch. Ver. Brunn. 14:209 (1876).


Perithecia solitary or gregarious, more or less immersed in rotting
wood, less beaked t h a n in the preceding genera, asci clavate, at first
8-spored, ascospores with transverse and longitudinal septa, budding
secondary spores within the ascus.

Rhamphoria pyriformis (Fries) von Ilohnel in S.B. K. Akad. Wiss.


Wien, Math.-Nat. Klasse Abt. 1, 122 (1913). (Fig. 13 G.)
Perithecia gregarious b u t without a stroma, more or less immersed
in rotting wood b u t sometimes almost superficial, pear shaped, about
0.2 m m . across, with a short stout beak, black. Asci cylindric-clavate,
up to 1 6 0 x 1 3 / / ; primary ascospores 8, irregularly biseriate, fusiform,
broadest above the middle, 1 5 - 3 5 x 3 - 8 / / , with up to 15 transverse
septa and ultimately a few longitudinal septa, constricted at each sep-
t u m , hyaline, budding off large numbers of broadly elliptical se-
condary spores about 2 x 1 p, while still within the ascus. Mature asci
are t h u s filled with countless very minute spores, like those of a
Tympanis or Corynella.

On very wet rotting wood of Quercus, in winter and spring. Uncommon.

"327
Family 13. Sphaeriaccae

The genera described below all have scattered, superficial, sub-


globose, black or dark brown perithecia with hyaline or yellowish
ascospores.

I. Ascospores 1-septate:
A. Ascospores with a long basal appendage, perithecia on marsh plants
Loramyces
B. Not as above:
1. Parasitic on lichens Rhagadostoma (p. 329)
2. Not so:
a. Ascospores narrowly cylindrical, perithecia spiny, on
leaves and twigs Niesslia (p. 329)
b. Ascospores elliptical, on decorticated wood:
*Perithecia subglobose, not collapsing, ascospores
minute Chaetosphaeria (p. 330)
**Perithecia collapsing and appearing concave above,
ascospores over 10 p long Melanopsamma (p. 330)
II. Ascospores multiseptate Zignoella (p. 331)

LORAMYCES Weston

Loramyces Weston in Mycologia 21:72 (1929).


Perithecia subspherical, dark coloured, minutely hairy, soft-fleshed,
opening b y a broad ostiole, asci 8-spored, thin-walled, ascospores fusi-
form, septate, hyaline, furnished with a long basal appendage and a
gelatinous sheath. For a full description of this unique genus see
Weston in Mycologia 2 1 : 5 5 - 7 6 (1929).

Loramyces juncicola Weston, op. cit.:72 (1929). (Fig. 17A.)


Perithecia scattered, superficial, sessile on a broad base, somewhat
flattened spheroidal, up to 2 m m . across, dark brown to blackish, soft-
fleshed and covered with short, septate, downy hairs. Asci cylindric-
clavate, up to 175x11//, thin-walled, w i t h o u t any annular thickening
or conspicuous pore at the tip, not blued by iodine; ascospores eight,
uniseriate, with a fusiform 1-septate apical portion 1 7 - 2 2 x 3 . 5 - 6 / /
and a slender basal appendage 5 0 - 6 5 x 0 . 5 - 0 . 7 5 / / ; paraphyses slender
with rounded tips. There is a colourless gelatinous sheath round the
head of the ascospore.
On dead culms of Juncus spp. and other marsh plants, below water level,
in summer. Not uncommon.

"328
RHAGADOSTOMA Korber

Rhagadostoma Korber, Parerga lichenologica: 473 (1865).


Pseudothecia massive, erumpent, asci thick-walled, surrounded b y
pseudo-paraphyses, 2- or 4-spored, ascospores very large, hyaline
1-septate, parasitic on lichens. There is only one species:

Rhagadostoma lichenicola (de Notaris) Keisslerin Rabenhorst's Kryp-


t o g a m e n Flora Deutschland 8 : 3 2 0 (1930). (Fig. 11M.)
Pseudothecia scattered, erumpent, rough, black, subglobose, about
l
/ 2 m m . across. Asci clavate, short-stalked, thick-walled above, up to
120x16//, containing two or four spores; ascospores biseriate, nar-
rowly elliptic-fusiform, 30-50x 7-10/t, 1-septate at m a t u r i t y , said
occasionally to become 3-septate, hyaline.

On the thallus of the Arctic-Alpine lichen Solorina crocea. Rare.

NIESSLIA Auerswald

Niesslia Auerswald apud Gonnermann & Rabenhorst, Mycologia euro-


p a e a : 30 (1869).
Perithecia superficial, globose, covered with stiff short bristles,
without a papillate ostiole, asci cylindric-clavate, thin-walled, with a
m i n u t e apical ring-like thickening, ascospores narrow, 1-septate,
hyaline.

Niesslia pusilla (Fries) Schroeter, K r y p t o g a m e n Flora von Schlesien 3


(2): 294 (1894). (PI. X X X V l . )
Perithecia gregarious, superficial, minute, about 150// across, glo-
bose, r a t h e r soft and thin-walled and hence often collapsing to become
concave above when dry, dark brown to black, covered with stiff,
radiating, dark brown hairs up to 5 0 x 4 / t . Asci cylindric-clavate, short
stalked, about 4 0 x 5 / / , with t r u n c a t e d conical tips, 8-spored; ascospo-
res irregularly biseriate, cylindric-fusiform, 9 - 1 1 x 2 - 2 . 5 / / , 1-septate,
hyaline.

On leaves and twigs of Pinus and Picea, also on Rubus canes, in spring.
Uncommon; known also as Niesslia exilis (Albertini& Schweinitz ex Fries)
Winter, Rabenhorst's Kryptogamen Flora Deutschland 1 (2): 196 (1885).

"329
C H A E T O S P H A E R I A Tulasne

Chaetosphaeria Tulasne, Selecta F u n g o r u m Carpologia 2:252 (1863).


Perithecia often gregarious b u t without a stroma, more or less
globose or pear-shaped b u t not long-beaked, superficial, smooth or
hairy, asci cylindric-clavate, ascospores hyaline, usually 1-septate.

Chaetosphaeria myriocarpa (Fries) Booth in Comm. Myc, Inst. Myc.


P a p e r 6 8 : 5 (1957). (Fig. 14A.)
Perithecia gregarious, superficial on wood, globose with a minute
papillate ostiole, black, smooth, shining, up to 0.2 m m . across. Asci
narrowly cylindric-clavate, short-stalked, up to 45x4/u, thin-walled,
8-spored; ascospores uniseriate, elliptic-cylindric, 5 - 6 x 2 . 5 / t , 1-sep-
t a t e , not constricted, hyaline.

On wood of various kinds. Very common.


This common species has long been known as Trichosphaeria myriocarpa
(Fries) Petrak & Sydow in Annales mycologici 22:331 (1924).

MELANOPSAMMA Niessl

Melanopsamma Niessl apud Saccardo in Michelia 1:347 (July 1878).


Perithecia superficial, globose b u t typically collapsing inwards in
the upper half when dry, with a papillate ostiole, asci cylindrical,
ascospores elliptical, 1-septate, hyaline.

Melanopsamma pomiformis (Persoon ex Fries) Saccardo, op. cit.:347


(1878). (PI. X X X V F . )

Perithecia gregarious, superficial, globose about 0.3 mm. across, the


upper half collapsing when d r y ; black, with a small apical papilla,
smooth or bearing a few colourless septate hyphae. Asci cylindric-
clavate, short-stalked, up to 8 5 x 1 5 / / , 8-spored; ascospores irregularly
biseriate, elliptic-cylindric, 1 2 - 1 6 x 5 - 7 / / , slightly constricted at the
single septum, hyaline, with a conspicuous oil drop in each cell.

On wood of deciduous trees, especially Fagus, in dense swarms. Un-


common.
Von Hohnel thought the affinities of this species lay with Nectria.

"330
ZIGNOELLA Saccardo

Zignoella Saccardo in Michelia 1 : 3 4 6 (July 1878).


Perithecia superficial, more or less globose with a papillate ostiole
smooth, black, asci clavate, ascospores multiseptate, hyaline.

Zignoella ovoidea (Fries) Saccardo, op. cit.:346 (1828). (Fig. 14B.)


Perithecia gregarious, superficial, on wood, globose a b o u t 0.3 m m
across with a low papillate ostiole, black, somewhat shining, smooth.
Asci clavate up to 140x14//, 8-spored; ascospores biseriate, fusiform,
one side often flattened, 2 4 - 2 8 x 4 - 5 / t , 3-septate, hyaline.
On decorticated wood of various deciduous trees. Common.
Many other species of Zignoella have been described but it is uncertain
how many of them are distinct as the genus has never been critically revised.
It occupies a very isolated position in the Sphaeriales and bears considerable
resemblance to Melanomma in Pleosporales but its asci are not clearly
bitunicate.

Order CORONOPHORALES

There is only one family in the order:

Family 1. Coronophoraceae

In this pecular group of genera the cavity of the perithecium origi-


nates by breakdown of the inner portion of an originally solid sclero-
tium-like body and the hymenium develops subsequently within the
cavity. The asci are rather thin-walled, clavate, with long slender
stalks like those of Diaporthe; the ascospores m a y be hyaline or brown,
septate or not and are often sausage-shaped b u t in some genera are
elliptical or even minutely spiny. The known British genera m a y be
distinguished as follows:

I. Perithecia large, immersed in bark, asci polysporous Coronophora


II. Perithecia superficial, asci 8-spored:
A. Perithecia large, subcylindrical, wall thick with a furrowed surface
Bertia (p. 332)
B. Perithecia usually less than 1 mm. across, wall thin and smooth:
1. Ascospores hyaline:
a. Ascospores sausage-shaped, non-septate
Nitschkia (p. 333)
b. Ascospores straight, 1-septate Calyculosphaeria (p. 333)
2. Ascospores coloured, non-septate Tympanopsis (p. 334)
Compare also Enchnoa, p. 304.

"331
CORONOPHORA Fuckel

Coronophora Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:229


(1870).
Perithecia large, solitary or in small compact groups without a
stroma, embedded in bark, globose, without an ostiole, asci clavate,
polysporous, ascospores sausage-shaped, hyaline.

Coronophora gregaria (Libert) Fuckel, op. cit.:229 (1870). (PI


XXXVK.)
Perithecia large, nearly 2 m m . across, black, smooth, globose, with-
out an ostiole, immersed in bark, singly or in compact clusters of up
to eight, erumpent at the tips. Asci clavate with long slender stalks,
about 7 0 x 1 6 / / containing a large n u m b e r of spores; ascospores ir-
regularly clustered, sausage-shaped, 7 - 9 x 1 - 2 / / , hyaline, non-septate.
On bark of various deciduous trees, apparently especially on Sorbus
aucuparia but not confined to Rosaceae. Uncommon.

Coronophora angustata (Nitschke) Fuckel, op. cit.:229 (1870), on


bark of Fagus, has smaller perithecia with ascospores only 4 - 6 // long.

BERTIA de Notaris

Bertia de Notaris in Giornale Botanico Italiano 1 : 3 3 4 (1846).


Perithecia superficial, subglobose to subcylindrical, black, wall thick
and conspicuously furrowed, asci clavate, 8-spored, ascospores very
large, hyaline, 1-septate.

Bertia moriformis (Tode ex Fries) de Notaris, op. cit.:335 (1846). (Fig.


14E.)

Perithecia densely gregarious b u t without a subiculum, superficial,


seated on a broad base, subcylindrical, up to 1.5 m m . high and % m m .
wide, black, the outer surface m a r k e d with coarse w a r t s and furrows
so as to resemble a blackberry or mulberry, without an ostiole. Asci
clavate, stalked, thin-walled, up to 105x13//, 8-spored; ascospores ir-
regularly biseriate, sausage-shaped, 30-50x4-5.5/«, hyaline, 1-septate.
On decorticated wood, especially of Fagus but occasionally even on
Pinus. Common.
See also Rhagadostoma, p. 329.

"332
NITSCHKIA O t t h

Nitschkia O t t h apud Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e


23-24:165 (1870).
Perithecia becoming superficial, gregarious, seated on a subiculum
of dark hyphae or even on a definite stroma, black, collapsing when
dry, asci clavate, long-stalked, 8-spored, ascospores sausage-shaped,
hyaline, non-septate.

Mtschkia cupularis (Persoon ex Fries) Karsten, Mycologia Fennica 2:


81 (1873). (Fig. 14c.)
Perithecia gregarious, seated within b a r k and exposed b y shedding
of the surface, subglobose, about 0.4 m m . across, with a prominent
papillate ostiole, the upper half collapsing inwards when dry, surface
minutely roughened, black. Asci clavate, long-stalked, about 70x12,«,
with a thickened cap at the apex, 8-spored; ascospores irregularly
arranged, sausage-shaped, 9 - 1 6 x 2 / / , non-septate. The perithecia
normally have a t u f t of brown hyphae at the base, forming a more or
less well developed subiculum.

On dead branches of Acer, Aesculus, Prunus etc. Not common.

CALYCULOSPHAERIA Fitzpatrick

Calyculosphaeria Fitzpatrick in Mycologia 15:45 (1923).


Perithecia gregarious, seated on a subiculum, black, papilla minute
or absent, asci clavate, long-stalked, ascospores eight, fusiform,
1-septate.

Calyculosphaeria tristis (Fuckel) Fitzpatrick, op. cit.:48 (1923). (Fig.


14D.)

Perithecia superficial, gregarious, seated on a blackish somewhat


iridescent subiculum composed of hyphae 7 - 9 / / wide, black, globose,
up to % m m . across, without an obvious ostiole, collapsing and
appearing cup-shaped when dry, surface minutely warted. Asci cla-
vate, long-stalked, up to about 5 0 x 7 / / , with a distinct apical thicken-
ing, 8-spored; ascospores more or less biseriate, straight, fusiform,
5 - 9 x 1 . 5 - 2 . 5 / / , 1-septate at m a t u r i t y , hyaline.

On wood, normally on that bearing members of the Diatrypaceae. Not


common.

"333
Calyculosphaeria collapsa (Romell) Fitzpatrick, op. cit.: 52 (1923),
has larger ascospores, 12-17x3.5-7/1.

TYMPANOPSIS Starback

Tympanopsis Starback in Bihang K. Svenska Vet.-Akad. Handl. 19 (3)


No. 2:24 (1894).
Perithecia superficial, seated on a subiculum, black, collapsing, asci
clavate, 8-spored, ascospores elliptical, brownish, non-septate, smooth
or minutely spiny.

Tympanopsis euomphala (Berkeley & Curtis) Starback, op. cit.:24


(1894). (Fig. 14F.)
Perithecia gregarious, superficial, seated on a dense blackish-brown
subiculum formed of iridescent hyphae, black, up to y 2 m m . across,
without an obvious ostiole, collapsing and appearing cup-shaped when
dry, surface minutely w a r t e d . Asci clavate, thin-walled, long-stalked,
about 5 0 x 1 2 / / ; ascospores irregularly clustered, elliptical or flattened
on one side, 7 - 9 x 4 - 5 / « , smoky to yellowish, usually with two oil drops.
On b a r k &c.

Order 11. PLECTASCALES

The Plectascales are characterised b y having thin-walled asci, often


subglobose or relatively short, not arranged in a h y m e n i u m ; in most
families the ascocarp is globose, without an ostiole. The British species
are referred to nine families, viz.:

I. Ascocarp with a long slender beak, asci diffluent and ascospores emerging
in a drop of mucilage at the apex of the beak
Ophiostomataceae (p. 335)
II. Ascocarps more or less globose, without ostioles, opening by splitting:
A. Saprophytes:
1. Ascocarp stalked, on animal debris, hoofs, horns, hair, etc.
Onygenaceae (p. 343)
2. Ascocarps sessile:
a. Ascocarps globose or ellipsoidal, asci enclosed within a
membrane:
+ Minute fungi on plant debris above ground:
t Ascocarps breaking open along predetermined
lines . . . . Cephalothecaceae (p. 342)

"334
f f N o so dehiscing Eurotiaceae (p. 337)
+ +Large more or less subterranean fruit bodies
Elaphomycetaceae (p. 344)
b. Clusters of asci enclosed by a hyphal web which does not
form a continuous membrane
Gymnoascaceae (not further described here)
B. Parasites, especially on leaves and other green parts of flowering
plants:
1. Mycelium without hyphopodia, conidia hyaline, abundant
Erysipliaceae (p. 345)
2. Mycelium bearing lateral swellings (hyphopodia), no conidia
Meliolaceae (p. 351)
C. Parasites on larvae of bees or on pollen in bee hives, asci produced
in dense clusters within a large spherical cyst-like cell, after fu-
sion between hyphae of male and female mycelia
Ascos phaeraceae

The Ascosphaeraceae are too minute to be dealt with here, see


Spiltoir and Olive in Mycologia 47:238-243 (1955).
For keys to the genera and species of Gymnoascaceae see H. H.
K u e h n in Mycologia 50:417-439 (1958) and a revision of 38 British
species in 9 genera b y A. E. Apinis, Comm. Myc. Inst. Myc. P a p e r : 96
(1964).

Family 1. Ophiostomataceae

The fungi referred here should be compared with those described


under Melanosporaceae, p. 262. Sphaeronaemella is indeed often
treated as a hyaline-spored member of t h a t family.

I. Perithecia dark coloured, on wood or bark . . . Ceratocystis (p, 336)


II. Perithecia pale, on dung or hyperparasitic on Gyromitra infula
Sphaeronaemella

Some species of Ceratocystis are of economic importance because of


the blue-gray stain caused b y their mycelia growing in coniferous
timber.

SPHAERONAEMELLA Karsten
Sphaeronaemella Karsten in Hedwigia 23:17 (1884), emend Seeler in
Farlowia 1:127 (1943).
Perithecia pink to cream coloured, with very long thread-like necks,
t e r m i n a t e d by a fringe of hairs which hold the globule of escaped asco-
spores. Asci diffluent, ascospores clliptical, non-septate, hyaline. The

"335
t y p e species is S. helvellae Karsten, hyperparasitic on ascocarps of
Gyromitra infula, in Europe.

Sphaeronaemella fimicola Marchal in Bull. Soc. roy. Bot. Belg. 30:143


(1891). (Fig. 30L.)
Perithecial venter sunk in the surface of dung, about % m m . dia-
meter, beak protruding, pale pink, up to 2 m m . long b u t very fine and
thread-like, about 20fi wide. Asci diffluent so t h a t t h e fungus long
passed as pycnidial; ascospores elliptical, hyaline, 5-9x2-4.5/u, emerg-
ing as a white globule at the tip of the beak. See Pease in Mycologia 40:
122 (1948).

On rabbit droppings.

CERATOCYSTIS Ellis & Halstead

Ceratocystis Ellis & Halstead apud Halstead & Fairchild in Journal of


Mycology 7 : 1 (1891).
Perithecia minute, on wood, bearing a very long slender beak, dark
coloured, asci irregularly arranged, 8-spored, subglobose, becoming
loose in a mucilaginous m a t r i x within the perithecium and soon dis-
appearing, liberating the ascospores which ooze out of the ostiole in a
drop of mucilage, ascospores small, hyaline, non-septate.

Ceratocystis piceae (Munch) Bakshi in Trans. Brit, mycol. So. 33:113


(1950). (Fig. 17c.)
Perithecia scattered, superficial, black, body globose, about % m m .
across, with a bristle-like beak up to 1 m m . long and 20-30fj, thick,
ostiole surrounded by a fringe of short hairs. Asci numerous, sub-
globose, 6 - 7 n across, 8-spored, soon disappearing, ascospores cylindric-
fusiform, often slightly curved, 3 . 5 - 4 . 5 x 1 . 5 - 2 h y a l i n e , emerging
from t h e ostiole in a drop of mucilage which is held in place b y the
fringe of short hairs.

On coniferous wood, both on logs and sawn timber, associated with a


greyish-blue stain. Common; also known as Ophiostoma piceae (Munch)
H.& P. Sydow in Annales mycologici 17:43 (1919). A monograph of Cerato-
cystis by J. Hunt will be found in Lloydia 19:1-58 (1957). The type species,
C. fimbriata Ellis & Halstead, was first described as a pycnidial fungus as
the diffluent asci were overlooked. Hence the genus Ophiostoma H.& P. Sy-
dow was proposed for species in which the asci had been seen.

"336
Ceratocystis ulmi (Buisman) C. Moreau in Revue de Mycologie,
Suppl. Colonial 17:22 (1952), causes a common and fatal disease of
Ulmus species.

Family 2. Eurotiaceao

The following simplified key will serve to differentiate the British


genera discussed below:

A. Ascospores large, brown, 1-septate:


1. Fruit bodies black Zopfia (p. 338)
2. Fruit bodies light coloured Heleococcum (p. 338)
B. Ascospores non-septate:
1. Ascospores globose, not merely circular and biconvex:
a. Ascospores smooth, uniseriate in cylindrical asci
Orbicula (p. 338)
b. Ascospores warted, irregularly arranged in elliptical asci
Lilliputia (p. 339)
2. Ascospores not globose, asci subglobose (except in Allescheria):
a. Ascospores biconvex, usually girdled by a furrow:
*Conidial state an Aspergillus:
•fAscocarp wall with a single layer of cells,
usually yellow . . . . Eurotium (p. 339)
ttAscocarp wall composed of several layers of
hyphae, dark-coloured, surrounded by loose,
globose, thickwalled cells
Emericella (p. 341)
**Conidial state Penicillium:
T
t Ascocarp w all composed of interwoven hyphae
Talaromyces (p. 338)
ttAscocarp wall of very thickwalled cells
Carpenteles
b. Ascospores not as above:
*Ascospores brown:
tAsci clavate Allescheria (p. 340)
ttAsci globose:
§Ascospores over 10/i long, with one or more
minute germ-pores Thielavia
§§Ascospores smaller, without germ-pores
Pseudeurotium (p. 341)
**Ascospores hyaline to yellowish, rough
Anixiopsis (p. 342)

The last three genera have affinity with Cephalotheca b u t lack the
sutures in the ascocarp wall. For descriptions and figures of four
British species of Thielavia isolated from soil and rotten wood see
C. Booth, Comm. Myc. Inst. Myc. P a p . : 8 3 (1961). The ascocarps of
Carpenteles Langeron in Compt. Rend. Soc. Biol. Paris 87:344 (1922)

"337
and Talaromyces B e n j a m i n are too small to be figured here. For an
account of these genera see B e n j a m i n in Mycologia 47:669-687 (1955).

ZOPFIA Rabenhorst

Zopfia Rabenhorst in Hedwigia 13:44 (1874).


There is only one species:

Zopfia rhizophila Rabenhorst, op. cit.:44 (1874). (PI. XLH.)

Fructifications superficial or partially embedded in the loose surface


tissue of the host, subglobose, without an ostiole, black, about % m m .
across, smooth. Asci broadly clavate, subsessile, 4- to 8-spored; asco-
spores very large, 6 5 - 7 5 x 3 2 - 4 5 / / , 1-septate and slightly constricted
at the septum, dark brown, r a t h e r thick-walled and with a pro-
nounced papilla at each end. Note t h a t the ascospore is figured at %
the magnification of most others on the plate.

On decaying roots of Asparagus and formerly regarded as parasitic; prob-


ably not uncommon.

HELEOCOCCUM J0rgensen

Heleococcum Jorgensen in Bot. Tidsskrift. 37:417 (1922).


Again only one species is k n o w n :

Heleococcum aurantiacum J0rgensen, op. cit.:417 (1922). (PL XLJ.)


Fructifications superficial, globose, smooth, without an ostiole, up
to y 3 m m . across, whitish, becoming light orange colour, associated
with a sparse white mycelium. Asci almost spherical, numerous,
tightly packed, 8-spored, about 4 5 x 3 5 / / ; ascospores elliptical, 25-30x
10-15/t, 1-septate b u t not constricted, r a t h e r thick-walled, yellowish-
brown, with from one to three large oil drops in each cell.
On humic soil and on mushroom compost. Apparently rare but easily
overlooked, the only British material seen was on compost.

ORBICULA Cooke

Orbicula Cooke, Handbook of British Fungi 2:926 (1871).


Perithecia subglobose, superficial, dark coloured, without an ostiole,
smooth, asci cylindrical, ascospores spherical or nearly so, yellow or
brown.

"338
Orbicula parietina (Schrader ex Fries) Hughes, Comm. myc. Inst. myc.
P a p . 4 2 : 1 (1951). (PL XLF.)
Perithecia often gregarious, superficial, subglobose, up to 1.3 m m .
across, with a thin purplish-brown wall, without an ostiole b u t brittle
and easily broken. Asci arising from a cushion at the base of the fructi-
fication, narrowly cylindric-clavate, up to 150x17//, thin-walled,
8-spored; ascospores uniseriate, spherical or nearly so, 7-13// diameter,
thin-walled, hyaline to yellow. The asci break down at m a t u r i t y ,
liberating the ascospores as a yellow powder filling the cavity of the
perithecium.
On compost, decaying bark, straw, rotting wood, decaying paper and
cardboard and similar substrata. Not uncommon.
This beautiful little fungus occupies rather an isolated position on
account of its cylindrical asci which may by some be regarded as arranged
in a hymenium, for they are interspersed with slender paraphysis-like
hyphae. It is not surprising that it has been repeatedly described as new and
has from time to time been assigned to nine different genera!

LILLIPUTIA Boudier & Patouillard


Lilliputia Boudier & Patouillard in Bull. Soc. mycol. France 16:144
(1900).
There is only one species:

Lilliputia rufula (Berkeley & Broome) Hughes in Comm. myc. Inst.


myc. Pap. 4 2 : 2 (1951). (PL XLG.)
Fructifications subglobose, superficial, associated with a sparse
white mycelium, about % m m . across, pale ochraceous or white at
first, becoming yellowish-brown with age, without an ostiole. Asci
numerous, embedded in thin-walled tissue which fills the centre of the
fruit body, 8-spored; ascospores globose, 13-24/t diameter, wall about
iju thick, densely studded with short b l u n t warts, yellow, usually with
a large central oil globule. The ascospores are liberated at m a t u r i t y b y
a break down of the asci and their associated tissue and the powdery
spores t h e n fill the cavity of the fructifications.
On dung of herbivorous animals and birds, mushroom compost and rot-
ting paper. Probably not uncommon but seldom collected.

EUROTIUM Link ex Fries


Eurotium Link ex Fries, Systema mycologicum 3 : 3 3 1 (1832).
Fructifications minute, light coloured, globose, without an ostiole or
other specialised means of dehiscence, superficial, on mycelium which

"339
boars conidiophores belonging to the form-genus Aspergillus Micheli
ex Fries, op. cit.:383 (1832); asci subglobose, irregularly arranged,
8-spored, ascospores colourless, usually more or less biconvex, like
two saucers placed together with their concave surfaces inwards, in
most species marked with a more or less ornamented furrow along the
line of junction of the "saucers". Because of the great economic im-
portance of the Aspergillus moulds in industrial mycology m a n y of
the species have merely been described in the form-genus Aspergillus,
w i t h o u t "legitimate" names in Eurotium. Their differentiation is a
highly critical m a t t e r and only one ubiquitous species which forms
ascocarps very freely will be mentioned here. Those who wish to pur-
sue f u r t h e r studies in the genus should consult the excellent mono-
graph b y T h o m & Raper, ' A M a n u a l of the Aspergilli', Baltimore (1945).

Eurotium herbariorum Link ex Fries, op. cit.:332 (1832). (PL XLE.)


Fructifications scattered on a sparse superficial mycelium amongst
the stalked greenish conidial heads of Aspergillus species in the group
of A.glaucus Link, globose, light yellow, about 100// diameter, smooth,
without an ostiole. Asci irregularly arranged, subglobose, 10-12fi
across, thin-walled, 8-spored; ascospores irregularly arranged, bi-
convex, with a more or less defined equatorial furrow, about 5 - 7 / / dia-
meter, hyaline.

On dead plant debris, herbarium specimens, jam and other preserves etc.
Common.
The above brief description applies to a collective species. Of the com-
ponent critical species perhaps the commonest is that associated with
Apergillus repens (Corda) de Bary, in which the ascospores measure 4.8-5.6 fi
in their broadest diameter and the marginal furrow is very feebly developed.
Thom & Raper, however, apply the name Eurotium herbariorum in a re-
stricted sense to a group of species characterised by large ascospores,
measuring from 6.5-10/« in long axis and with well developed, sometimes
minutely ornamented, furrows. Unfortunately these large ascospored strains
appear to be relatively rare and hence hardly likely to be the true E. her-
bariorum of the older mycologists.

ALLESCHERIA Saccardo & Sydow

Allescheria Saccardo & Sydow, Sylloge F u n g o r u m 14:464 (1899).


Fructifications globose, with a thin brown wall, often bearing hyphae
from surface cells b u t no specialised appendages. Asci clavate, 8-spor-
ed; ascospores elliptical or inequilateral, somewhat pointed at the
ends, brown.

"340
Allescheria terrestris Apinis in Nova Hedwigia 5 : 6 8 (1961).
Cleistocarps globose, black, embedded in or beneath a t h i n web of
brown hyphae, 1 h 0 - 1 L m m - diameter, wall thin and smooth a p a r t from
the emergent hyphae. Asci clavate, 1 4 - 2 0 x 6 - 8 / t , in clusters on asco-
genous hyphae, 8-spored; ascospores ovoid, brown, smooth, 4 - 6 x 3 - 4 [x.
On dead grass stems and in soil. (Fig. 30E.)

EMERICELLA Berkeley & Broome

Emericella Berkeley & Broome, Introduction to Gryptogamic B o t a n y :


340 (1857).
Ascocarps as in Eurotium b u t with a wall composed of more t h a n
one layer of hyphae and surrounded by looser hyphae bearing globose
thickwalled cells ("Hiille cells"). The ascospores are reddish-purple or
blueish instead of hyaline. The t y p e species is E. variecolor Berkeley &
Broome, with conidial state Aspergillus variecolor Thom & Raper.

Emericella nidulans (Eidam) Vuillemin in Gomptes Rendus Academie


des Sciences (Paris) 184:137 (1927). (Fig. 30F.)
Ascocarps with a dark purplish-brown wall, concealed b y loose
hyphae bearing numerous, more or less globose, glistening, white,
thickwalled Hiille cells, about 20/« diameter. Asci evanescent, 8-spor-
ed; ascospores lenticular, smooth, with two equatorial sinuous crests,
reddish-purple, 4 - 4 . 5 x 3 . 5 - 4 / / .
In dung, decaying vegetation and soil.

The conidial state is Aspergillus nidulans (Eidam) Winter.


Cleistocarps with a similar multilayered dark wall b u t without Hiille
cells are referred to Sartorya Vuillemin, op. cit.: 136 (1927), with type
species S. fumigata Vuillemin, ascus state of the common Aspergillus
fumigatus Fresenius.

PSEUDEUROTIUM van Beyma

Pseudeurotiuin van Beyma in Zentralbl. f. B a k t . II, 96:411 (1937).


Fructifications superficial, globose, smooth, dark coloured, without
an ostiole, thin-walled, not associated with Aspergillus, asci subglobose,
with eight or more spores, ascospores globose or elliptical, smooth,
brown.

"341
Pseudeurotium ovalis Stolk in Antonie van Leeuwenhoek 21:78 (1955).
(PI. XLc.)
Fructifications globose, dark brown smooth, up to 180// diameter,
wall formed of a single layer of polygonal brown cells. Asci globose to
broadly elliptical, 7 . 5 - 9 x 6 . 5 - 8 / / , thin-walled, 8-spored; ascospores
ellipsoid, non-septate, 5 . 5 - 6 x 3 . 5 - 4 / / , smooth, hyaline at first, be-
coming brown at m a t u r i t y .

Figured from material kindly communicated by Miss Stolk, isolated from


cysts of the nematode Heterodera rostochiensis collected in the Isle of Jersey.
There is an associated conidial state belonging to the form-genus Sporo-
trichum.
For descriptions and figures of two other British species see Booth,
Comm. myc. Inst. myc. Paper 83 (1961).

ANIXIOPSIS Hansen

Anixiopsis Hansen in Bot. Zeit. 55 (1):27 (1897).


This genus is similar to Eurotium b u t lacks the Aspergillus conidial
state and its ascospores lack the characteristic equatorial furrow of
the latter.

Anixiopsis stercorarea (Hansen) Hansen, op. cit.:27 (1897). (PI. X L n . )


Fructifications borne on a white superficial mycelium, globose,
smooth, without an ostiole, up to about % m m - across, dark brown.
Asci numerous, irregularly arranged and filling the centre of the fruit
body, at first embedded in a thin-walled ground tissue as in Lilliputia,
subglobose, thin-walled, 6 . 5 - 7 . 5 x 5 - 6 . 5 / / , 8-spored; ascosporcs broadly
elliptical, 2 . 5 - 3 x 2 - 2 . 5 / / , minutely spiny, hyaline at first, ultimately
light yellow. The asci break down and liberate the ascospores within
the cavity of the m a t u r e fructification.

On dung and in soil, sometimes in cysts of the eelworm Heterodera rosto-


chiensis; figured from a culture kindly communicated by Miss Stolk.

Family 3. Cephalothecaceae

There is only one British genus with sutures in the ascocarp wall,
along which it breaks open at m a t u r i t y . There is, however, an allied
tropical genus, Batistia Ciferri, with similar ascocarps borne singly
on long stalks and Booth holds t h a t a n u m b e r of genera with dark
ascospores without distinct sutures in the wall are akin to Cephalo-

"342
theca viz. Anixiopsis and Pseudeurotium, here left at the end of
Eurotiaceae, Emericellopsis, Pyenidiophora, Saturnomyces, Thielavia
and Westerdykella.

CEPHALOTHECA Fuckel

Cephalotheca Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 25-26:297


(1871).
Fructifications superficial, dark coloured, subglobose, without an
ostiole b u t having the wall ornamented with regularly arranged fur-
rows along which it shatters at m a t u r i t y , asci numerous, subglobose,
irregularly arranged, ascospores small, elliptical or kidney-shaped,
brown.

Cephalotheca sulfurea Fuckel, op. cit.:297 (1871). (PI. XLi.)


Fructifications gregarious, superficial, subglobose, up to % m m .
across, black, their surfaces divided into regular polygonal areas b y a
network of furrows, otherwise smooth, each seated on a small dense
m a t of sulphur-coloured hyphae. A t m a t u r i t y the perithecia walls
break open along the furrows until there remains only a basal cup
containing the loose spores. Asci subglobose, about 10// across, thin-
walled; 8-spored; ascospores broadly elliptic-fusiform, 4 - 6 x 3 - 4 / / ,
light brown.
On rotting wood, typically on Quercus. Rare.
Cephalotheca purpurea (Shear) Chesters in Trans. Brit, mycol. Soc.
19:266 (1935), and C. reniformis Saccardo & T h e r r y in Michelia 2:312
(1881), have kidney-shaped ascospores, measuring 2 . 5 - 3 x 1 . 5 - 2 / / and
4 - 5 x 3 - 3 . 5 / / respectively.

Family 4. Onygenacoae

The single small genus Onygena is usually separated from the Euro-
tiaceae on account of its stalked ascocarps and its peculiarly restricted
h a b i t a t , on animal remains.

ONYGENA Fries
Onygena Fries, Systema mycologicum 3:206 (1829).

"343
Onygena equina (Willdenow) Persoon ex Fries, op. cit.:206 (1829)
(PI. X L l . )
Fructifications solitary or in small clusters, with a subglobose fertile
head and a slender sterile stalk. Head 2 - 4 m m . across, light brown,
smooth, without an ostiole b u t eventually bursting to liberate the
spores, wall rather thick, enclosing a loose weft of hyphae and, at first
asci, later loose ascospores, in a yellowish mass. Asci subglobose, thin-
walled, 14-22 x 10-14/«, 8-spored; ascospores elliptical, 6—9x4—5.5//,
light brown, smooth. The stalk of t h e fructification is cylindrical,
smooth, whitish, up to 6 x 2 m m .

On shed horns of cow and sheep, on rotting hooves and similar substrata.
Common.

Onygena corvina Albertini & Schweinitz ex Fries, op. cit.: 208 (1829)
is a similar fungus found on cast pellets of owls and similar aggrega-
tions of rotting animal hair.

Family 5. Elaphomycetaceae

Two British genera m a y be placed here.


I. Ascospores brown, warted Elaphomyces
II. Ascospores hyaline, smooth Diehliomyces

ELAPHOMYCES Nees ex Fries

Elaphomyces Nees ex Fries, Systema mycologicum 3:57 (1829).


Fructifications large, tuber-like, subterranean or at least buried
under leaf litter, with a thick rind enclosing a powdery spore mass,
dark in colour b u t permeated b y light coloured veins.

Elaphomyces granulatus Fries, Systema mycologicum 3 : 5 8 (1829).


(PL IXk.)
F r u i t bodies globose to ovoid, up to 4 cm. across, pale ochraceous,
the outer surface covered with small somewhat pyramidal warts. The
rind consists of two layers, an outer thin yellowish stratum which in-
cludes the w a r t s and an inner m u c h thicker compact white zone.
Within this is a mass of purplish-black spores, at first divided into
compartments b y t h i n whitish bands of sterile tissue. Asci globose to
pear-shaped, 35-45/t across, thin-walled, usually 6-spored, soon

"344
breaking down and disappearing; ascospores spherical, 24-32/t dia-
meter, blackish-brown, covered with low irregular warts.

In surface layers of soil in woods of Pinus, less often under deciduous


trees. Common in suitable situations.
There are a number of additional British species of Elaphomyces, whose
characteristic features are fully described in Miss Hawker's monograph,
cited on p. 71.

DIEHLIOMYCES Gilkey

Diehliomyees Gilkey in Mycologia 46:789 (1954).


There is one British species.

Diehliomyees microsporus ( D i e h l & L a m b e r t ) Gilkey op. cit. (PI. IXL.)


Fruit bodies subglobose to irregularly lobed, up to 3 cm. across,
outer surface cream-coloured to pinkish-brown, finely scurfy; flesh
firm, cream to yellowish veined. Asci irregularly arranged, subglobose,
mostly 8-spored, up to 2 5 x 1 5 / / ; ascospores globose, 5 - 7 / t diameter,
smooth, hyaline, thin-walled. Formerly referred to the genus Pseudo-
balsamia Fischer.

Occurs as a weed in beds of the cultivated mushroom.


The fruit bodies develop mainly on the surface of the soil. This species is
not dealt with in Dr. Hawker's monograph but a full description will be
found in Mycologia 22 (1930), and an account of control measures in Phyto-
pathology 34 (1944). The wall of the ascocarp consists of undifferentiated
woven hyphae instead of being stratified as in Elaphomyces and Gilkey
accordingly suggested a relationship with Gymnoascaceae.

Family 6. Erysiphaceae

The Erysiphaceae comprise the well-known plant parasites com-


monly called " P o w d e r y Mildews" because of the powdery white
coating formed on green plants by their superficial mycelium and its
innumerable chains of conidia. Except in Leveillula and Phyllactinia
the mycelium is purely superficial, creeping over t h e host and merely
sending small branches into the epidermal cells. The mycelium bears
large numbers of short erect branches each of which terminates in a
chain of relatively large, thin-walled, cylindrical or barrel-shaped,
hyaline conidia. In Phyllactinia and Leveillula the conidia are ovoid

"345
and borne singly on the tips of erect hyphae. On this account, and be-
cause t h e y possess also a mycelium which penetrates f u r t h e r into t h e
host t h a n the epidermal cells, these two genera are sometimes segre-
gated from the others to form a distinct subfamily, Phyllactinieae.
Ascocarp s develop on the same mycelium as the conidia, often late in
t h e season or on fading leaves, and are dark coloured at m a t u r i t y . Seven
genera are recognised, distinguished p a r t l y b y the nature of the h y p h a l
appendages which spring from the wall of the ascocarp and p a r t l y b y
the n u m b e r of asci it contains.

f. Ascocarp containing a single ascus:


A. Appendages attached to the base of the fructification or absent
merely hyphoid and not characteristically branched
Spliaerotheea
B. Appendages characteristically branched at the tip or arising from
the apex of the ascocarp Podosphaera (p. 347)
II. Ascocarp containing several asci:
A. Appendages hooked at the tip Uncinula (p. 348)
B. Appendages repeatedly forked at the tip Microsphaera (p. 348)
C. Appendages of two types, a ring of long, stiff, pointed bristles round
the equatorial zone and a crown of short sfender ones over the
summit Phyllactinia (p. 351)
D. Appendages hyphoid, unbranched or irregularly so:
1. Conidia in chains, mycelium purely superficial
Erysiphe (p. 349)
2. Conidia solitary, mycelium largely within the host
Leyeillula (p. 350)

There are two valuable and comprehensive monographs of the Erysipha-


ceae: E. S. Salmon, A Monograph of the Erysiphaceae, in Memoirs of the
Torrey Botanical Club 9 (1900), which includes all the species known at that
time, and: S. Bliimer "Die Erysiphaceen Mitteleuropas", in Beitrage zur
Kryptogamenflora der Schweiz 7 (1) (1933), which deals only with European
species.
Bliimer adopts a narrower species concept than Salmon and hence re-
cognises a greater number of species. Salmon's concepts are still generally
followed in England and North America and are adopted here.

SPHAEROTHECA Leveille

Sphaerotheca Leveille in Ann. Sci. Nat. I l l , 15:133 (1851).


The characters of the genus are sufficiently indicated in the key. In
Salmon's system there are four British species, with two named
varieties; in Bliimer's there are 14, all with host plants t h a t occur in
the British Isles.

"346
Sphaerotheca humuli (de Candolle ex Merat) Burrill in Bull. Illinois
State Lab. Nat. Hist. 2:400 (1887).. (Fig. 17J.)
Ascocarps gregarious or scattered, globose, up to % m m . across,
dark brown to black, smooth above b u t giving off numerous h y p h a l
appendages from the lower half, these h y p h a e are brown, unbranched,
m a t t e d and interwoven with the mycelium and with those of n e a r b y
ascocarps. Asci broadly elliptical, 4 5 - 9 0 x 50-72/x, 8-spored; ascospores
elliptical, hyaline, 2 0 - 2 5 x 1 2 - 1 8 / / .

On living leaves and other green parts of many plants, especially of


Rosaceae, also on Dipsacus, Epilobium, Humulus, Phlox, Ribes, Viola etc.
The race figured is that on Filipendula ulmaria; in Bliimer's system this
race on Rosaceae is called S. macularis (Wallroth ex Fries) Jaczewski,
Karmanyi opredielitel gribov: 65 (1927).
Salmon recognised also a variety fuliginea (Schlechtendal ex Fries) Sal-
mon in Mem. Torrey Bot. Club 9:49 (1900), on a wide range of hosts,
distinguished by having the cells of the ascocarp wall 20-30/« wide, instead
of 10-20/i, as in the typical form. Other British species are:

Sphaerotheca euphorbiae (Castagne) Salmon in Bull. Torrey Bot.


Club. 29:95 (1902), on Euphorbia.
Sphaerotheca mors-uvae (Schweinitz) Berkeley & Curtis in Grevillea
4 : 1 5 8 (1876), the cause of American Mildew on Ribes.
Sphaerotheca pannosa (Wallroth ex Fries) Leveille in Ann. Sci. Nat.
Bot. I l l , 15:138 (1861), on Rosa, with a variety persicae Woronichin
on Prunus persica.

PODOSPHAERA Kunze

Podosphaera Kunze, Mykol. Hefte 2:111 (1823).


Distinguished from Sphaerotheca b y its equatorially or apically
attached, repeatedly forked appendages. Salmon recognised four
species with one variety; Bliimer distinguished eight species on
British plants b u t two of these, though acknowledged by Salmon, have
not been found here.

Podosphaera oxyacanthae var. tridactyla (Wallroth) Salmon in Mem.


Torrey Bot. Club 9 : 3 6 (1900). (Fig. 17L.)
Ascocarps scattered or gregarious, on a scanty mycelium, globose,
a b o u t 100/t across, dark brown to blackish at m a t u r i t y , with from two
to eight erect appendages springing from the top of the ascocarp, each
with a long stalk, brown below shading t o hyaline at the tip, forked

"347
three to five times nearly at right angles near the apex. Ascus sub-
globose, 6 0 - 7 8 x 6 0 - 7 0 / / , 8-spored; ascospores 2 0 - 3 0 x 1 3 - 1 5 / / .
On species of Prunus.
Typical Podosphaera oxyacanlhae, on Crataegus, Prunus, Pyrus, Sorbus,
Spiraea and Vaccinium, differs in its more spreading appendages, attached
equatorially.

Podosphaera leucotricha (Ellis & E v e r h a r t ) Salmon in Mem. Torrey


Bot. Club 9 : 4 0 (1900), causes mildew of the cultivated apple tree and
other species of Malus and Pyrus.

UNCINULA Leveille

Uncinula Leveille in Ann. Sci. Nat. Bot. I l l , 15:151 (1851).


The ascocarps of this genus are readily recognised by their pe-
culiarly hooked appendages, which are unbranched, except in U. bi-
cornis. Salmon and Bliimer recognise four British species.

Uncinula bicornis (Fries) Leveille, op. cit.: 153 (1851). (Fig. 17G.)
Ascocarps mostly scattered, on a sparse mycelium, subglobose,
somewhat flattened above, up to 1 / e m m . across, blackish, bearing over
the upper half a large n u m b e r of short spreading appendages, mostly
not longer t h a n the diameter of the ascocarp, each forked near the tip
with the branches strongly recurved to form a double hook, hyaline.
Asci 4-12, more or less pear-shaped, 7 0 - 9 5 x 4 5 - 5 5 / / , 8-spored, asco-
spores 2 2 - 2 6 x 1 3 - 1 5 //.
On species of Acer.

Uncinula necator (Schweinitz) Burrill apud Ellis & E v e r h a r t , North


American Pvrenomycetes: 15 (1892), is the common mildew of Vitis
b u t the ascocarps are very seldom seen.
Uncinula prunastri (de Candolle ex Merat) Saccardo, Sylloge Fungo-
r u m 1 : 7 (1882), occurs on Prunus.
Uncinula salicis (de Candolle ex Merat) Winter, Rabenhorst's
Kryptogamen Flora Deutschland 1 (2): 40 (1884), occurs on Salix.

MICROSPHAERA Leveille

Microsphaera Leveille in Ann. Sci. Nat. Bot. I l l , 15:154 (1851).


Differs from Uncinula in its repeatedly forked appendages, with the
branches not hooked b u t forming a compact head on a long stalk.
Salmon recognised seven British species.

"348
Microsphaera grossulariae [Wallroth] Leveille, op. cit.: 160 (1851).
(Fig. 17K.)
Ascocarps scattered or gregarious, on a sparse mycelium, globose,
about % mm. across, black, bearing from 5 to 22 appendages, not
much longer t h a n the diameter of the ascocarp, each from four to five
times regularly forked close together near the tip. Asci from four to
ten per ascocarp, ovate, short-stalked, 4 6 - 6 0 x 2 8 - 3 8 / / , 4- to 6-spored;
ascospores 2 0 - 2 8 x 1 2 - 1 5 / / .

On Ribes, especially R. grossularia, but confined to the leaves, whereas


Sphaerotheca mors-uvae forms dense mycelial mats on fruits and shoot-tips
also.

Microsphaera alphitoides Griffon & Maublanc in Bull. Soc. mycol.


France 28:100 (1912), is the very common powdery mildew of Quercus
b u t ascocarps are only occasionally found, usually in very hot sum-
mers, as in 1959.
Microsphaera astragali (de Candolle ex Merat) Trevisan, Spighe e
Paglie 1 : 3 9 (1853), occurs on Astragalus.
Microsphaera baumleri P. Magnus in Ber. Deutsch Bot. Ges. 17:148
(1899), is found occasionally on Vicia sylvatica.
Microsphaera berberidis (de Candolle ex Merat) Leveille, op. cit.: 159
(1851), is common on Berberis vulgaris and occurs also on Mahonia
aquifolium.
Microsphaera euonymi (de Candolle ex Merat) Saccardo, Sylloge
F u n g o r u m 1 : 1 1 (1882), occurs on Euonymus europaeus.
Microsphaera mougeotii Leveille, op. cit.: 158 (1851), is found on
Lycium.

ERYSIPHE Hedwig ex Merat

Erysiphe Hedwig ex Merat, Nouvelle Flore de Paris 1:131 (1821).


Ascocarps like those of Sphaerotheca b u t containing numerous asci;
borne on a purely superficial mycelium with a b u n d a n t chains of coni-
dia belonging to the form-genus Acrosporium Nees es Sumstine. Sal-
mon recognised five British species, two of t h e m with very wide host
ranges; Bliimer recognises about two dozen species within the field
covered by Salmon's E. polygoni and E. cichoracearum.

Erysiphe polygoni de Candolle ex Merat, op. cit.: 132 (1821). (Fig. 17i.)
Ascocarps scattered or gregarious, globose, dark brown to blackish,
seldom more t h a n 100// across, bearing numerous horizontally spread-

"349
ing hyaline or brown h y p h a l appendages, which m a y branch b u t , if so,
not in any distinctive p a t t e r n and often become interwoven with the
mycelium. Asci usually from two to eight, sometimes more, ovate,
sometimes short-stalked, 46-72x30-45/«, each containing from three
to eight spores; ascospores 1 9 - 2 5 x 9 - 1 4 f i .

Very common on herbaceous plants of many families. The form illustrated


is that on Heracleum which Bliimer differentiates as Erysiphe umbelliferarum
de Bary in Beitr. z. Morph. u. Phys. Pilze 1: (1870).

Erysiphe tortilis [Wallroth] Fries, System mycologicum 3:243 (1829),


characterised b y its very long u p s t a n d i n g appendages, is confined to
Cornus.
Erysiphe cichoracearum de Candolle ex Merat, op. cit.: 132 (1821),
with almost as wide a host range as E. polygoni, is distinguished b y its
asci regularly containing only two ascospores.
Erysiphe galeopsidis de Candolle ex Merat, op. cit.: 134 (1821), found
on m a n y Labiateae, is separated from E. cichoracearum by its asco-
spores not developing until after the leaves on which it grew have died
and b y its hyphal branches within the epidermal cells being lobed.
Erysiphe graminis de Candolle ex Merat, op. cit.: 133 (1821), is the
very common powdery mildew of cereals and wild grasses.

LEVEILLULA A r n a u d

Leveillula Arnaud, Annales des Epiphyties 7 : 9 4 (1921).


Ascocarps like those of Erysiphe b u t associated with conidia borne
singly on short h y p h a l branches (form-genus Oidiopsis Scalia) and
with a b u n d a n t mycelium permeating the tissues of the host. There is
only one species:

Lcveillula taurica (Leveille) A r n a u d , op. cit.:94 (1921). (Fig. 17F.)


Ascocarps scattered or gregarious, often somewhat embedded in a
dense superficial mycelium, up to % m m . across, globose or becoming
concave, densely fringed with simple appendages m a t t e d into t h e
mycelium. Asci usually about 20, ovate distinctly stalked, 75-110 x
28-40/u, each with two large spores; ascospores 28-40x14-22jx.

On a wide range of host plants but mainly in the Mediterranean region,


Central Europe and the Near East; found in England on species of Helian-
themum.

"350
PHYLLACTINIA Leveille

Phyllactinia Leveille in Ann. Sci. Nat. Bot. I l l , 15:144 (1851).


Salmon recognised b u t a single species of Phyllactinia, which Bliimer
has subdivided into ten. The following description is based on t h a t by
Salmon:

Phyllactinia guttata (Fries) Leveille, op. cit.: 144 (1851). (Fig. 17H.)
Ascocarps usually on the under side of leaves, associated with a very
sparse mycelium, usually scattered, up to about y 3 m m . across, sub-
globose to somewhat flattened, black at m a t u r i t y , bearing round t h e
equatorial zone a ring of from 5 to 18 stiff, pointed, colourless append-
ages, from one to three times as long as t h e ascocarp is wide, each with a
subglobose hollow swelling at its base. The apex of the ascocarp is
covered with a mop of short, slender, more or less mucilaginous h y p h a l
branches. Asci numerous, up to 45, ovate, more or less stalked, 60-105
x 2 5 - 4 0 / / , normally 2-spored; ascospores 3 0 - 4 0 x 1 6 - 2 5 / / .
On leaves of many plants but mainly on those of trees and shrubs;
especially common on Corylus.

Family 7. Meliolaceao

This is another family of obligate parasites on green plants, whose


members superficially somewhat resemble t h e Erysiphaceae in t h a t
t h e y have black subglobose ascocarps borne by myelium restricted to
the surface of living p l a n t organs. The mycelial mat, however,
appears black to the eye and very dark brown under the microscope,
when it is seen to bear numerous very short lateral branches with club-
shaped or lobed tips (hyphopodia) attached to the host cells. I t does
not carry chains of aerial conidia b u t in some genera it is studded with
stiff erect bristles (setae). The family includes an enormous n u m b e r of
species, mostly with very restricted host ranges, b u t nearly all are
confined to plants of tropical or subtropical regions. Only two are
known to occur in the British Isles b u t these represent two of t h e
widespread tropical genera, viz.:

I. Mycelium beset with setae Meliola


II. Mycelium without setae, often having curious worm-like appendages on
the ascocarp Irene (p. 352)
For an account of the family as a whole see Hansford, The Meliolineae.
Beihefte zur Sydowia 2, 1961.

"351
MELIOLA Fries

Meliola Fries, Systema Orbis Vegetabilis 1:111 (1825).


There is only one British species:

Meliola niessleana Winter in Hedwigia 2 4 : 2 6 0 (1885). (PI. XLA.)

Mycelium forming small irregular black patches on both surfaces of


leaves, hyphae dark brown, about 7/t wide, thick-walled, septate,
branched to form a close network over the surface, bearing numerous
short hyphopodia, arranged alternately on opposite sides of a h y p h a ,
each with a short basal cell and a swollen, club-shaped, angular
or slightly lobed head, total dimensions, 2 1 - 2 5 x 1 4 - 1 7 / / . The mycelium
is also studded with stiff, erect, black setae up to about 1 m m . high,
and with the globose black ascocarps, about y 3 m m . across. Asci ob-
long, up to 100x30//, 2- to 4-spored; ascospores brown, elliptic-fusi-
form, slightly curved, 4 7 - 6 4 x 1 4 - 1 6 / / , 3-septate and slightly cons-
tricted at the septa.

On Vaccinium vitis-idaea. Uncommon.

IRENE Theissen & Sydow

Irene Theissen & Sydow in Annales mycologici 15:194 (1917).


Again there is only a single British species:

Irene calostroma (Desmazieres) von Hohnel in Annales mycologici 16:


213 (1918). (PI. XLB.)
Mycelium forming rather sparse networks on leaves and canes of
Rubus, dark brown, septate, bearing alternate capitate hyphopodia
which are slightly lobed at the t i p ; no mycelial setae. Ascocarps glo-
bose, with a tuberculate surface, black, about % m m - across, bearing
appendages about 100// long. Ascospores elliptic-fusiform, often curved,
brown, about 4 5 - 5 0 x 1 5 / / , 3-septate, slightly constricted at the septa.

Reported once, on Rubus idaeus in County Kerry. It is desirable that the


species be searched for again in the west of Ireland and south-west England.
According to Ilansford the legitimate name is Appendiculclla calostroma
(Dosm.) v. Hohn.

LOCULOASCOMYCETES

The five orders represented in the British flora m a y be distinguished


as follows:

"352
I. Asci subglobose, occurring separately in cavities of a stroma
Myriangiales
II. Asci clavate or cylindrical, or the ascocarp is shield-shaped and dehis-
cent:
A. Ascocarps shield-like (Thyriothecia) on superficial mycelium
Hemisphaeriales (p. 428)
B. Ascocarps elongated or branched, opening by a slit (Hystero-
thecia), usually with massive black walls Hysteriales (p. 422)
C. Ascocarps compound, containing several perithecium-like cavities
Dothideales:Dothideaceae (p. 358)
D. Mycelium a sooty mould or dense hyphal mat on leaves:
1. Mycelium a sooty mould, ascospores brown, muriform
Dothideales:Capnodiaceae (p. 366)
2. Ascospores septate, hyaline, mycelium bearing setae, forming
a superficial membrane on living leaves
Dothideales:Chaetothyriaceae (p. 367)
3. Ascospores septate often coloured, mycelium not membranous,
not setose in British genus Dothideales:Dimeriaceae (p. 368)
D. Ascocarps globose to pyriform, not on sooty moulds, often immersed
or erumpent:
1. Asci large, few, separated by stromatic tissue
Dothideales:Pseudosphaeriaceae (p. 356)
2. Asci numerous, not distinctly separated by stromatic tissue,
ascocarps like those of Sphaeriales but with bitunicate asci
Pleosporales (p. 369)
3. Asci subsessile, clustered on a basal cushion, ascocarps minute
Dothideales:Mycosphaerellaceae (p. 362)

Order 1. MYRIANGIALES

Two families are represented in the British Isles, easily distinguish-


ed t h u s :

1. Asci embedded in a gelatinous black or olive-brown thallus of a honey-


dew saprophyte, usually on the surface of leaves, thallus hyphae form-
ing characteristic moniliform branched chains of cetls embedded in a
gelatinous matrix, multicellular asexual propagules commonly pro-
duced in specialised portions of the thallus surface Atichiaceae
II. Parasites, thallus without the distinctive structure described above
Myriangiaceae (p. 354)

Family 1. Atichiaceae

There is only one British genus, much better represented in the wet
tropics.

"353
ATICHIA Flotow

Atichia Flotow in Linnaea 23:149 (1850).


One European species has long been known t h o u g h described at
first as a lichen because of its resemblance in colour, habit and t e x t u r e
to some minute m e m b e r of Collemaceae.

Atichia glomerulosa (Acharius ex Mann) Stein, K r y p t . Flora Schlesien


2 (2):356 (1879). (Fig. 30J.)
Thalli superficial, originating as thin discoid plates, in size, shape
and colour like droppings of flies, b u t soon becoming stellately lobed,
olive-black, up to 8 m m . across. The tips of thallus lobes are differen-
tiated into cupulate asexual reproductive organs containing numerous
black, roughly globular, multicellular propagules. Asci subglobose,
8-spored, spores 1 2 - 1 8 x 6 j i , hyaline, 1-septate.
On evergreen leaves covered with honeydew, often with Chaeto-
thyriurri babingtonii, on coniferous bark in Europe. Apparently con-
fined to the humid west b u t probably generally overlooked or passed
as an alga by mycologists.

Family 2. Myriangiaceae

The family is best developed in the tropics; its British genera m a y


be easily differentiated t h u s :
I. Ascocarp erumpent or superficial, with sterile and fertile portions
Myriangium
II. Ascocarp undifferentiated, immersed in surface tissue of the host:
A. Parasites, spores multiseptate Elsinoe
B. Saprophytes, spores becoming muriform Dermatina

MYRIANGIUM Montagne & Berkeley

Myriangium Montagne & Berkeley in Hooker's London J o u r n . Bot. 4:


72 (1845).
Stroma mostly superficial, cushion-shaped or with a basal cushion
bearing crowded, short, nail-shaped outgrowths with flattened tops
which m a y break down to become slightly cup-shaped, usually black;
asci embedded singly in p a r e n c h y m a t o u s tissue of the outgrowths,
ascospores multiseptate, hyaline.

"354
Myriangiumduriaei Montagne& Berkeley, op. cit.:73. (PI. X X X V I c . )
Stroma up to 5 m m . across, with a flattened base bearing crowded
obconical tubercles with flat or concave tops which contain the fertile
tissue, externally black, flesh whitish or greenish. Asci globose, or
nearly so, up to 50// across, 8-spored; ascospores elliptic-cylindric,
1 4 - 3 7 x 6 - 1 5 / / , usually with 7 transverse septa and one or more
longitudinal ones, hyaline.

Parasitic on scale insects on bark, especially of Fraxinus. Uncommon,


only in the south-west of England, Ireland and west coast of Scotland.

ELSINOE Raciborski

Elsinoe Raciborski, Parasitische Algen und Pilze J a v a ' s 1 : 1 4 (1900).


Ascocarp a pad of undifferentiated tissue inserted immediately
beneath the surface of the host, containing isolated cavities, each with
a single ascus, ascospores multiseptate, hyaline. There is usually an
associated conidial state in the form-genus Sphaceloma de Bary emend
Jenkins in Mycologia (1941).

Elsinoe veneta (Burkholder) Jenkins in J o u r n . Agric. Res. 44:696


(1932). (PI. X X X V I B . )
Stroma minute, developed in greyish patches of slightly sunken host
tissue, usually with a purplish border, individual stromata subepider-
mal, up to y 3 m m . across, appearing as a grey dot to the naked eye,
forming a pad of pseudoparenchymatous tissue within which the asci
are arranged at random, each in a separate cavity. Asci globose, thick-
walled, up to 30// diameter, 8-spored; ascospores elliptic-cylindric,
1 8 - 2 1 x 6 . 5 - 8 / / , 3-septate and slightly constricted at the septa, hyaline.
The lower half of the figure shows a vertical section through an
ascocarp.

On canes and, less well developed, on leaves of Rubus species. Common,


causing the disease known as Cane Spot.

DERMATINA Almquist

Dermatina Almquist in Kgl. Svensk. Vet.-Akad. Handl. 17 (6) :8 (1880).


Stroma minute, with black outer layer and pale flesh, subdivided by
black partitions, erumpent from bark. Asci and spores as in Elsinoe or

"355
becoming brown and muriform. Most species are associated with algal
cells and hence "lichens"; those devoid of algal cells are sometimes
placed in a separate genus, Cyrtidula Minks in Rev. Mycol. 13:61
(1891).

Dermatina quercus (Massalongo) Zahlbruckner, Catalogus lichenum


universalis 1 : 5 5 1 (1922). (Fig. 19b.)
S t r o m a t a very minute, just visible to the naked eye as small black
dots, usually in groups on a slightly paler p a t c h of bark, irregularly
lobed; showing from beneath, when picked off and viewed under a
strong lens, a n u m b e r of brownish areas separated b y black partitions.
Asci broadly elliptical to subglobose, 2 0 - 2 5 x 1 7 - 2 5 / / , stained reddish-
brown by Melzer's reagent, embedded singly in a small celled, light
brown tissue; ascospores eight, closely packed, ellipsoidal, colourless
or pale brownish, with usually four cross septa and one longitudinal
septum, 1 5 - 1 8 x 5 - 8 / « on young bark of trees especially Quercus.

This species is usually classed as a "lichen" but both Vainio and Watson
were unable to find associated algal cells and considered it to be a non-
lichenised fungus. The name Cyrtidula quercus (Massalongo) Minks in Rev.
mycol. 13:61 (1891), is available for those who prefer to assign lichenised
and non-lichenised fungi to different genera.

Order 2. DOTIIIDEALES

The families included in this order on a basis of developmental stu-


dies are of widely divergent external appearance and have hence been
keyed out separately on p. 353. T h e y are characterised especially by
the absence of paraphyses and pseudoparaphyses from their asco-
carps.

Family 1. Pseudosphaeriaceae

Pseudothecia small, immersed in plant tissue, containing relatively


few large asci, each within a separate cavity in stromatic tissue b u t
clavate and arranged side b y side, not globose and scattered as in
Myriangiales. Many genera of Pleosporales have been included here
from time to time b u t as redefined by Luttrell the family contains two
British genera:
I. Ascospores with transverse septa only, distinctly fusiform Wettsteinina
II. Ascospores ultimately with transverse and longitudinal septa, or broadly
elliptical with transverse septa only Pseudoplea (p. 357)

"356
WETTSTEININA von Hohnel

Wettsteinina von Hohnel in S.B. K. Akad. Wiss. Wien, Math.-Nat.


Klasse Abt. 1, 116:126 (1907).
Ascocarps small, globose to flask-shaped, embedded in plant tissue,
usually stems or leaves of herbs, with imperfectly developed ostioles,
asci few, clavate, relatively large, separated from one another by con-
tinuous strips of pseudoparenchymatous tissue, ascospores large,
usually over 40// long, ultimately yellowish or brownish, with one or
more transverse septa and often with an outer gelatinous coat.

Wettsteinina niesslii E. Miiller in Sydowia 4 : 2 0 4 (1950). (Fig. 10G.)


Ascocarps scattered or in small clusters, hemispherical with a flat-
tened base and minute apical papilla, black, about % m m . across.
Asci clavate, up to 110x30/«, 8-spored, separated b y strips of sterile
tissue; ascospores irregularly biseriate, narrowly fusiform, 5 2 - 6 2 x
7 - 9 / / , at first l - s e p t a t e , slightly constricted at t h e septum, hyaline,
with a distinct gelatinous sheath, ultimately becoming 3-septate and
yellowish.

On dead culms of Phragmites communis below the water line.

PSEUDOPLEA von Hohnel

Pseudoplea von Hohnel in Annales mycologici 16:162 (1918).


Ascocarps embedded in leaf tissue, minute, globose, with im-
perfectly developed ostioles, asci few, very large, broadly clavate,
separated by belts of sterile tissue, ascospores seldom more t h a n 40/«
long, ultimately b o t h transversely and longitudinally septate, hyaline
or brownish, with gelatinous outer coats.

Pseudoplea myrtillina (Fautrey & Saccardo) P e t r a k in Sydowia 1 : 6 0


(1947). (Fig. 21E.)
Fungus causing brown spots with purple borders on living leaves
and tips of twigs, ascocarps scattered, black, erumpent, up to 120//
across. Asci only two to four per fruit body, broadly oblong, up to
200x 60//, very thick-walled, 8-spored; ascospores irregularly biseriate,
elliptic-cylindric, 3 5 - 5 0 x 1 5 - 1 8 / / , with three transverse septa and
noticeably constricted at the middle one, hyaline, with a t h i n ge-
latinous outer coat.

"357
On Vaccinium myrtillus, in Autumn. Uncommon.
It will be seen that this is not a typical Pseudoplea in that its ascospores
lack longitudinal septa. Nevertheless Petrak transferred it to that genus on
account of the general shape of the ascospore, which resembles that of
Pseudoplea rather than that of Wettsteinina. It has also been called Pseudo-
sphaeria myrtillina (Fautrey & Saccardo) von Hohnel but Pseudosphaeria is
now regarded as synonymous with Wettsteinina.

Pseudoplea trifolii (E. Rostrup) P e t r a k in Annales mycologici 19:29


(1921), found in leaf-spots of Trifolium and Medicago, is a typical
Pseudoplea with transverse and longitudinal septa in ascospores
30-40 x 14-16//.

Family 2. Dothideaceae

Traditionally this is a very large family b u t in the modern concept


it is greatly reduced and it is not even certain t h a t all of the few
species still referred to it here are correctly assigned.

I. Ascospores yellow or brown at maturity:


A. Ascospores 1-septate:
1. Ascocarps associated with a Polythrincium state, on Trifolium
(See Mycosphaerellaeeae p. 362)
2. Not associated with a Polythrincium, often on woody organs
Dothidea
B. Ascospores with three or more septa:
1. On lichen thalli Homostegia (p. 359)
2. On ferns Rhopographus (p. 360)
II. Ascospores permanently hyaline, 1-septate:
A. Stroma developed beneath epidermis of grasses and Equisetum
Scirrhia (p. 360)
B. Cushion-shaped stroma subcuticular or subepidermal on dead leaves
Euryachora (p. 361)
C. Thick cushion-shaped stroma erumpent from dead twigs
Plowrightia (p. 362)

DOTHIDEA Fries ex Fries

Dothidea Fries ex Fries, Systema mycologicum 2 : 5 4 8 (1823).


S t r o m a t a erumpent, cushion-shaped, with numerous cavities con-
taining thick-walled asci with. 1-septate brown ascospores.

Dothidea tetraspora Berkeley & Broome in Ann. Mag. n a t , Hist. I l l , 3:


37 (1859). (Fig. 15K.)
Stroma erumpent from beneath bark, hemispherical or irregular,
about 2 m m . across, black, smooth, containing numerous small asco-

"358
genous cavities. Asc icylindric-clavate, 9 0 x 1 4 / / , 2-4-spored;ascospores
uniseriate, elliptic-cylindric, 2 0 - 2 7 x 8 - 1 1 / / , dark brown.

On dead stems of Daphne laureola, said to be also on Ulex. Figured from


the type collection.

Dothidea sambuci Fries, Systema mycologicum 2 : 5 5 1 (1823). (Fig.


18c.)
S t r o m a t a gregarious, erumpent, cushion-like, black, about 1 m m .
long, containing numerous very small ascogenous cavities. Asci
cylindric-clavate up to 9 5 x 1 6 / / , 8-spored; ascospores irregularly bi-
seriate, 19-21x8-9/«, 1-septate with the lower cell smaller t h a n the
upper, olive-brown.

On dead twigs of Sambucus and other shrubs. Not common.

HOMOSTEGIA Fuckel

Homostegia Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 2 3 - 2 4 : 2 2 3


(1870).
Stroma black, embedded in lichen thalli, containing numerous
cavities filled with thick-walled asci and paraphysis-like hyphae,
ascospores 3-septate, brown. The genus presumably should be trans-
ferred elsewhere because of the a b u n d a n t " p a r a p h y s e s " b u t develop-
mental studies are desirable before an a t t e m p t is made to redispose it
appropriately.

Homostegia piggotii (Berkeley & Broome) Karsten, Mycologia Fen-


nica 2:222 (1873). (Fig. 11L.)
Stroma immersed, causing small, black, cushion-like swellings of the
thallus, up to 3 m m . across, with a slightly tuberculate surface, asco-
genous cavities numerous, about % m m . across. Asci broadly clavate,
short-stalked, thick-walled, up to 7 0 x 2 4 / / , 8-spored, surrounded by
numerous slender hyaline h y p h a e ; ascospores elliptic-fusiform, straight
or slightly curved, 1 9 - 2 3 x 7 - 8 / / , 3-septate and slightly constricted at
the septa, dark brown.

On the thallus of the lichen Parmelia saxatilis.

"359
RHOPOGRAPHUS Nitschke

Rhopographus Nitschke apud Fuckel in J a h r b . Nass. Vereins f. Natur-


kunde 23-24:219 (1870).

S t r o m a t a subepidermal, elongated, black, asci thick-walled, asco-


spores 3- to multiseptate, brown. British species on ferns.

Rhopographus filicinus (Fries) Nitschke apud Fuckel, op. cit.:219


(1870). (Fig. 22F.)
S t r o m a t a subepidermal, elongated, a b o u t 3 m m . long, black, form-
ing small blisters on the surface of the host, each containing numerous,
small, crowded, ascogenous cavities. Asci broadly cylindric-clavate,
almost sessile, thick-walled, up to 8 0 x 2 1 //, 8-spored; ascospores in
two or three rows, closely packed, elliptic-cylindric, 2 8 - 3 6 x 7 - 8 / / ,
usually slightly curved, 3-7-septate, brown.

In swarms on dead leaf stalks of Pteridium aquilinum. Very common


everywhere.

SCIRRHIA Nitschke

Scirrhia Nitschke apud Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e


23-24:220 (1870).

S t r o m a t a elongated, subepidermal, containing rows of ascogenous


cavities, asci thick-walled, ascospores 1-septate, hyaline. The British
species is on grass b u t there is a S. castagnei (Montagne) Fuckel on
Equisetum in Europe.

Scirrhia rimosa (Albertini& Schweinitz ex Fries) Fuckel op. cit. : 221


(1870). (Fig. 22J.)
S t r o m a t a subepidermal, elongated, sometimes up to 3 cm. long,
black, containing rows of ascogenous cavities, splitting the epidermis
longitudinally. Asci cylindric-clavate, thick-walled, up to 110x14//,
8-spored; ascospores biseriate, elliptic-cylindric or slightly clavate,
narrowed below, 1 7 - 2 4 x 5 - 6 / / , 1-septate, hyaline.

Forming grey spots and stripes on dead leaf-sheaths of Phragmites.


Common.

"360
EURYACHORA Fuckel

Euryachora Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:220


(1870).
S t r o m a t a cushion-shaped, black, subcuticular or subepidermal in
dead leaves, containing numerous ascogenous cavities, asci thick-
walled, ascospores hyaline, 1-septate, usually unequally so. Miiller and
von Arx (1962) restrict Euryachora to species in which the ascospore
has a median septum, with t y p e species E. sedi (Fries) Fuckel on
Sedum spp.

Euryachora betulina (Fries) Schroeter, K r y p t o g a m e n Flora von Schle-


sien 3 (2):474 (1897). (Fig. 22G.)
S t r o m a t a gregarious, subcuticular, about % m m . across, black,
containing several m i n u t e ascogenous cavities. Asci cylindric-clavate,
thick-walled, up to 4 4 x 1 2 / / , 8-spored; ascospores biseriate, elliptic-
cylindric, 1 0 - 1 4 x 4 - 5 / / , with one septum towards the upper end of the
spore, not constricted, hyaline.

In clusters on fallen leaves of Betula. Very common. Also known as


Atopospora betulina (Fries) Petrak in Annales mycologici 23:101 (1925), and
transferred by von Arx to the Yenturiaceae.

Euryachora ulmi (Schleicher ex Fries) Schroeter, K r y p t o g a m e n Flora


von Schlcsien 3 (2):473 (1897). (Fig. 22D.)
S t r o m a t a subepidermal, cushion-shaped, black, 2 or 3 m m . across,
ultimately shedding t h e epidermis and appearing somewhat tuber-
culate, containing numerous small ascogenous cavities about % m m .
across. Asci narrowly cylindrical, not very thick-walled, sessile up to
6 0 x 8 / / , 8-spored; ascospores uniseriate, ovoid, 1 0 - 1 2 x 4 - 4 . 5 / / , with
a septum very near the lower end, hyaline or nearly so.

On fallen leaves of Ulmus from January onwards. Very common every-


where, preceded on the living leaves by a conidial state, Septogloeum ulmi
(Wallroth) Diedicke, which produces 3-septate hyaline spores, 20-58 x 2-6 //
in small acervuli.
The ascospores are said to become pale olive-brown when fully mature
and the species is then referred to Dothidea or its synonymous genus
Systremma Theissen & Sydow in Annales mycologici 13:330 (1915), by some
authors. It is, however, the type species of Platychora Petrak in Ann. mycol.
23:103 (1925) as P. ulmi (Fries) Petrak, referred by Miiller and von Arx to
the Venturiaceae.

"361
PLOWRIGHTIA Saccardo
Plowrightia Saccardo, Sylloge F u n g o r u m 2:635 (1883).
S t r o m a t a cushion-shaped, erumpent, usually on twigs, containing
numerous ascogenous cavities, ascospores 1-septate, hyaline. The
genus is reunited with Dothidea by Muller and von Arx.

Plowrightia ribesia (Persoon ex Fries) Saccardo, op. cit.:635 (1883).


(Fig. 15L.)
S t r o m a t a scattered or gregarious, cushion-shaped with flat tops, up
to 3 m m . across, erumpent through bark, black; flesh brown, enclosing
numerous small cavities in a single layer across the top. Asci cylindric-
clavate, stalked, up to about 100x14//, 8-spored; ascospores biseriate,
elliptic-cylindric or slightly clavate, 1 6 - 2 2 x 5 - 6 / / , 1-septate r a t h e r
below the middle, only slightly constricted, hyaline.
On dead twigs of Ribes. Not uncommon. Dothidea ribesia Persoon ex
Fries, Syst. myc. 2:550 (1823) is a synonym.

Family 3. Mycosphaerellaceae

In this family the stromata are minute and usually contain only a
single ascogenous cavity. The ripe ascocarp therefore resembles a
perithecium and is distinguished from the pseudothecia of the Pleo-
sporales b y the absence of hyphal threads among the asci and b y the
latter's clustered arrangement.

I. Ascospores 1-septate:
A. Asci 8-spored Mycosphaerella
13. Asci many-spored Rehmiellopsis (p. 366)
II. Ascospores with several cross-septa Sphaerulina (p. 365)
III. Ascospores occasionally developing a longitudinal septum in addition
to several cross-septa, asci clustered on a hemispherical cushion
Saceothecium (p. 365)
Compare also Pharcidia, p. 371.

MYCOSPHAERELLA Johanson
Mycosphaerella Johanson in Svenska Vetensk.-Akad. Ofvers. 9 : 1 6 3
(1884).
Pseudothecia minute, usually black, more or less globose, asci
relatively large, often ventricose, without pseudoparaphyses, asco-

"362
spores 1-septate, hyaline or nearly so. Many species have been des-
cribed, mainly on dead herbaceous stems and dead leaves or leaf-spots,
b u t it is probable t h a t a large proportion of the "species" are based on
chance occurrence on different hosts. See especially J . A. von Arx,
"Beitrage zur Kenntnis der G a t t u n g Mycosphaerella" in Sydowia 3:
28-100 (1949). The following species are representative of the genus as
it occurs in Britain.

Mycosphaerella lineolata (Roberge & Desmazieres) Schroeter, K r y p t o -


gamen Flora von Schlesien 3 (2): 339 (1894). (Fig. 21A..)
Pseudothecia immersed, usually in long rows, erumpent by their
ostioles, about 100// across, black; asci few, pyriform with a very thick-
walled apex, subsessile, about 40 x 20//, 8-spored; ascospores irregularly
arranged, elliptic-cylindric or very slightly clavate, 1 6 - 2 2 x 4 - 5 / / , with
a single median septum, not constricted, hyaline.
On dead grasses, especially Phragmites and Ammophila, also on Carices.
Very common.

Mycosphaerella iridis (Desmazieres) Schroeter, op. cit.:339 (1894).


(Fig. 21c.)
Pseudothecia scattered, immersed, black, up to 125// across, their
ostioles erumpent on either side of the leaf; asci few, fusiform-clavate,
subsessile, thick-walled at the top, up to 50x20//,, 8-spored; ascospores
irregularly arranged, slightly clavate, rounded at the ends, 16-21 x
4.5-6//, 1-septate, often with two oil drops in each cell, hyaline.
In dead brown patches on living and fading leaves of Iris. Common.
This is the type species of the genus Didymellina von Hohnel but it is
generally agreed that this is not generically distinct from Mycosphaerella.

Mycosphaerella punctiformis (Persoon ex Fries) Starback in Bihang


till K. Svenska Vet.-Akad. Handl. 15 (3) No. 2 : 9 (1889). (Fig. 21B.)
Pseudothecia scattered in the mesophyll of dead leaves, their
ostioles erumpent mostly on the under surface, black, up to 140//,
across; asci cylindrical, subsessile, thick-walled at the tip, up to 5 0 x 8 / / ,
8-spored; ascospores mostly uniseriate, narrowly ovate, 7 - 8 x 2 - 3 / / ,
1-septate, hyaline.
On fallen leaves of various deciduous trees, figured on Quercus. Very
common.

Mycosphaerella maculiformis (Persoon ex Fries) Schroeter, Krypto-


gamen Flora von Schlesien 3 (2): 333 (1894), is differentiated b y its

"363
pseudothecia occurring in compact groups up to 1 m m . across and by
having ascospores 9 - 1 4 / / long. Among other notable species m a y be
mentioned:
Mycosphaerella brassicicola (Duby) Oudemans, Rev. Champ. P a y s
Bas 2:210 (1897), common in large dark grey spots in living and dead
leaves of cultivated Brassicas.
Mycosphaerella carinthiaca J a p p in Annales mycologici 6:210 (1908),
in marginal leaf-spots of Trijolium species.
Mycosphaerella fragariae (Tulasne) Lindau, Kryptogamen Flora f u r
Anfanger 2:111 (1912), in leaf-spots of Fragaria.
Mycosphaerella primulae (Auerswald & Heufler) Schroeter, op. cit.:
338 (1894), on leaves of Primula auricula.
Mycosphaerella recutita (Fries) Johanson, op. cit.: 166 (1884), on
Dactylis and other grasses, with ascospores 11-14x3.5-4/«.
Mycosphaerella tassiana (de Notaris) Johanson, op. cit. (1884), on
innumerable dead herbaceous plants, grasses and sedges, with asco-
spores mostly 18-25x6-8/«.

Mycosphaerella killiani P e t r a k in Annales mycologici 39:324 (1941).


(Fig. 2 2 E . )
S t r o m a t a scattered, black, up to 1 m m . across, bearing at
first, in summer, a layer of short brown conidiophores, each with a
zig-zag outline owing to its sympodial development, bearing olive-
brown, ovate, 1-septate conidia 20-22x11-15/« belonging to the form-
genus Polythrincium Kunze & Schmidt ex Fries; in a u t u m n the stro-
m a t a develop round their margins small black globose spermogonia,
containing hyaline spermatia 3 - 5 x 1 . 5 - 2 / / , which emerge from the
ostioles as minute pale mucilaginous droplets. On overwintered leaves
ascocarps develop in minute cavities within the stroma, each con-
taining a few clavate asci, each with eight ascospores which are bi-
seriate, narrowly ovoid, 1-septate, 2 0 - 2 6 x 8 - 9 / / , at first hyaline, ulti-
mately pale yellow.

On living and faded leaves of Trifolium. Very common. Although this


fungus is so abundant there is no pseudothecial material at Kew or at the
Commonwealth Mycological Institute, and although it has been described
independently by three authors none has adequately described the asci.
Hence Fig. 33D represents the conidial spermafial states only, the ascospore
has been drawn from the published descriptions.
Mycosphaerella killiani is commonly known as Cymadothea trifolii Wolf,
in Mycologia 27:71 (1835) but Cymadothea differs from Mycosphaerella only
in its conidial state and is treated as a section of the latter genus by modern
authors. There is already a different Mycosphaerella trifolii (Karsten) Moesz
in Magyar Tud. Akad. Balkan. Kutat. tud. eredm. 8:138 (1925).

"364
SPHAERULINA Saccardo
Sphaerulina Saccardo in Michelia 1 : 3 9 9 (November 1878).
Similar to Mycosphaerella b u t with multiseptate ascospores.

Sphaerulina taxi (Cooke) Massee, Diseases of Cultivated P l a n t s : 2 2 0


(1910). (Fig. 21F.)
Pseudothecia scattered, minute, immersed with erumpent ostioles,
black; asci clavate, thick-walled at t h e tip, up to 7 5 x 1 3 / / , 8-spored;
ascospores biseriate, narrowly elliptical, 2 0 - 3 5 x 6 . 5 - 9 / / , 3-septate,
hyaline.
On dead leaves of Taxus baccata, associated with a pycnidial state, Cyto-
spora taxifolia Cooke & Massee which contains rod-shaped conidia 3 - 5 x 1 //.
Common and capable of killing both leaves and twigs of Yew.

SACCOTHECIUM Fries

Saccothecium Fries, S u m m a Veg. Scand. Sect. P o s t . : 3 9 8 (1849).


Saccothecium is said to differ from Sphaerulina in its ascospores
ultimately developing longitudinal as well as transverse septa, in oc-
curring on woody stems and in having darker walls to the ascostroma.
In the common species figured here, however, t h e longitudinal septa
are usually hard to find, t h e y were not seen in the material drawn for
PL X X X V I D . ) The genus is t h e n b e t t e r recognised by the brown
hemispherical cushion upon which the asci are seated, which readily
comes away bearing all its asci with it when a pseudothecium is
crushed in water on a slide. This mass of asci, rather like an old-
fashioned straw skep beehive, is shown at a moderate magnification in
the upper part of the figure.

Saccothecium sepincola (Fries) Fries, S u m m a Veg. Scand. Sect. P o s t . :


398 (1849). (Fig. 21D.)
Pseudothecia gregarious, usually in short rows, globose, black,
immersed in bark, with erumpent ostioles, about J / 5 mm. across. Asci
seated upon a hemispherical or short cylindrical basal cushion of small
brown cells, rather numerous, clavate, subsessile, up to 7 0 x 1 6 / / , very
thick-walled above, 8-spored; ascospores irregularly biseriate, elliptic-
cylindric 1 6 - 2 0 x 5 . 5 - 7 / / , with usually 5-transverse septa, markedly
constricted at the middle septum, hyaline, occasionally with a longi-
t u d i n a l septum inserted as well.

"365
On dead twigs of Rosa, the host figured, also on Cornus and Rubus.
Common. Other names current for this peculiar little fungus are Prings-
heirnia sepincola (Fries) von Ilohnel in Annales mycologici 18:97 (1920),
and Sphaerulina intermixta (Berkeley & Broome) Saccardo.
See Wehmeyer in Mycologia 49:83-94 (1957).

REHMIELLOPSIS Bubak & K a b a t


Rehmiellopsis B u b a k & K a b a t in N a t u r w . Zeits. f. Land- u. Forst-
wirtsch. 8:320 (1910).
Pseudothecia minute, immersed or erumpent, asci with more t h a n
eight spores, ascospores 1-septate, hyaline. There is only one British
species:

Rehmiellopsis abietis (E. Rostrup) 0 . Rostrup in Dansk. Bot. Arkiv. 2


(5): 16 (1916). (PI. X X X V I I c . )
Pseudothecia scattered, mostly on the upper surface of the needle,
immersed then erumpent, globose, up to 200// across, black. Asci few,
clavate, thick-walled, up to 9 0 x 2 2 / / , with 16-24 spores; ascospores
irregularly arranged, distinctly clavate, 1-septate, hyaline, 11-21 x
4-6.5 //.

On living needles of species of Abies; known also as Rehmiellopsis bo-


hemica Bubak & Kabat, op. cit.: (1910).

Family 4. Capnodiaceae

The Capnodiaceae are "Sooty moulds" with masses of dark brown


mycelium which form a black coating over the surface of leaves and
twigs of plants infested b y plant lice. The fungi grow in the sugary
secretions of the insects and are not themselves plant parasites though
t h e y m a y be injurious to leaves by cutting t h e m off from the light.
The family thrives mainly in w a r m climates and is poorly represented
in the British Isles; sooty moulds on leaves in this country are usually
mere mycelial growths without perithecia and in most cases are pro-
bably not connected with the t r u e Capnodiaceae.
Only two genera will be mentioned here:

I. Ascospores multiseptate, brown Metacapnodium (p. 367)


II. Ascospores with both transverse and longitudinal septa
Capnodium (p. 367)

"366
METACAPNODIUM Spegazzini

Metacapnodium Spegazzini in Physis 4 : 2 8 8 (1918).


There is a single British species; the type of the genus.

Metacapnodium juniperi (Phillips & Plowright) Spegazzini, op. cit.:


288 (1918). (Fig. 30H.)
Mycelium forming a black felt on small patches of bark, pseudo-
thecia ovoid, half embedded in loose hyphae, smooth, black, about
1
/ 5 m m . tall. Asci clavate, about 6 0 x 2 2 - 2 4 / / , 8-spored; ascospores
elliptical, 1-3-septate, brown, 2 0 - 3 3 x 8 - 1 4 / / . The associated mycelium
consists of chains of dark brown subglobose cells, about 17-20// wide,
becoming progressively smaller down to about 5/t at the tips of the
hyphae.

On twigs of Juniperus communis.

CAPNODIUM Montagne

Capnodium Montagne in Ann. Sci. Nat. Bot. I l l , 11:233 (1849).


Pseudothecia usually somewhat pear-shaped, with a short stout
stalk, black, seated on a superficial, short-celled, blackish-brown
mould; asci thick-walled, ascospores dark brown, with both transverse
and longitudinal septa.

Capnodium salicinum Montagne apud Berkeley & Desmazieres in


J o u r n . Soc. Hortic. London 4 : 2 5 1 (1849). (Fig. 17E.)
Pseudothecia globose to pear-shaped with short stout stalks,
minute and hard to find amongst the mycelial m a t , about x / 6 m m .
across, black, smooth. Asci few, almost sessile, ventricose, up to
6 0 x 2 5 / / , very thick-walled above, up to 8-spored; ascospores elliptical
or slightly clavate, 1 6 - 2 6 x 9 - 1 3 / / , usually with four transverse septa
and constricted at the middle one, also with one longitudinal septum,
dark brown.
On living leaves and young twigs of Salix and Alnus. Uncommon, at least
with pseudothecia.

Family 5. Chaetothyriaceae

In this family the mycelium is pale olivaceous to dark brown,


entirely superficial, without hyphopodia b u t often with erect bristles.

"367
The pseudothecia are globose and superficial, containing thick-walled
asci without interascal hyphae. There is one British genus:

CHAETOTHYRIUM Spegazzini

Chaetothyrium Spegazzini in Ann. Soc. Cientifica Argentina 26 (1888).


Mycelium usually scanty b u t in the British species forming a con-
spicuous dark film on leaves, bearing erect dark bristles; pseudothecia
globose, dark, covered by the mycelial m e m b r a n e asci 8-spored, asco-
spores hyaline, multiseptate.

Chaetothyrium babingtonii (Berkeley) Keissler in Rabenhorst's Kryp-


togamen Flora Deutschland Abt. 1 , 9 ( 2 ) : 4 4 6 ( 1 9 3 7 ) . ( P L X X X V I E . )
Mycelium mainly on t h e u p p e r surface of leaves, forming a small
olive to blackish film, studded with erect, dark-brown, pointed bristles
up to 210/JL long and 8// thick. Pseudothecia subglobose, smooth, black,
about 1 / 6 m m . across, with a m i n u t e apical pore. Asci very numerous,
cylindric-clavate, up to 6 3 x 1 7 / / , subsessile, thick-walled at the tip,
8-spored; ascospores irregularly biseriate, elliptic-fusiform, sometimes
slightly curved, 1 4 - 2 3 x 5 - 7 / / , hyaline, up to 6-septate.

On leaves, especially of evergreens such as Buxus, Ligustrum, Prunus


laurocerasus, Rhododendron but also sometimes on those of deciduous trees
or even herbs, mainly in western districts but extending northwards to the
Hebrides. It is noteworthy that both the ascocarps and the bristles arise
beneath or within the mycelial mat, push it up and remain largely covered
or sheathed by it. The species has been much misunderstood and was first
described as a lichen but it appears to have no algal cells regularly associated
with it. It does not appear to be parasitic on the leaves or to grow on insect
exudates so that its mode of nutrition in such an aerial habitat is difficult
to understand.

Family 6. Dimeriaceae

This is a very large family of leaf-parasites in the tropics b u t is re-


presented by a single British genus, occurring on moss:

LIZONIA Cesati & de Notaris

Lizonia Cesati & de Notaris, Sferiacei Italici 71 (1863).


Pseudothecia gregarious, free on a dark coloured mycelium, smooth,
asci large, cylindric-clavate, 8-spored, ascospores 1-septate, yellowish

"368
brown; parasitic on mosses. Except for the difference in host and the
less developed mycelial m a t there is little to differentiate this from
Gibellina. There is only one species:

Lizonia emperigonia (Auerswald) de Notaris, Sferiacei Italici 71 (1863).


(PI. X X X V I I i . )
Pseudothecia gregarious, slightly elongated, about % m m . across,
black, with a minute conical papilla, associated with a sparse m a t of
brown mycelium 2 f i thick. Asci cylindric-clavate, short-stalked, up
to 130x23//, thick-walled above, 8-spored; ascospores biseriate,
narrowly elliptic-fusiform, 3 2 - 4 3 x 1 2 - 1 4 1 - s e p t a t e , slightly con-
stricted at the septum, the upper cell slightly broader t h a n the lower,
yellowish-brown.

In the antheridial cups of Polytrichum, in spring. Rare, or seldom seen.

Order 3. PLEOSPORALES

This order includes t h e great m a j o r i t y of Loculoascomycetes. T h e y


usually have spherical to flask-shaped pseudothecia, superficially like
the perithecia of the Sphaeriales b u t containing bitunicate asci, inter-
mingled with pseudoparaphyses.
The British representatives are distributed in seven families:

I. Pseudothecia with a prominent elongated ostiole at the tip of a flattened


papilla Lophiostomataceae (p. 379)
If. Ostiole a circular pore:
A. Ascospores non-septate, hyaline . . Botryosphaeriaceae (p. 373)
B. Ascospores 1-septate:
1. Ascospores brown:
a. Ascocarps on dung (see Trichodelitschia, p. 412)
b. Ascocarps lichens Verrucariaceae (p. 370)
2. Ascospores not brown:
a. Ascospores first hyaline to greenish, becoming greyish or
pale olivaceous, ascocarps often bristly
fr x I'm i Venturiaceae (p. 374)
^ b . Ascospores hyaline JH/'). / / .y'/J^'PIeosporaceae (p. 382)
C. Ascospores. 2-or more seplate: ' f f i t ^ i / * • /-)ccLa/VK.
1. Ascospores grey-green, small ascoca/ps hairy ^
Herpotrichiellaceae (p. 370)
2. Ascospores hyaline to brown:
a. Pseudothecia with an ostiole . . Pleosporaceae (p. 382)
b. Pseudothecia without an ostiole Perisporiacoae (p. 422)

"369
F a m i l y 1. Herpotrichiellaceae

Pseudothecia small, scattered or clustered, externally bristly, at


least round the pore, asci clavate or oblong, small, ascospores small,
olive-grey, septate, on rotten wood. This is a small family and only
two genera need be mentioned here:
I. Pseudothecia scattered, ascospores with cross and longitudinal septa,
—>asd^ithJJLspores Capronia (not further described)
II. Pseudothecia clustered on a basal stroma, asci 8-spored, ascospores as
above Berlesiella
Capronia will not be further described though there is said to be one little
known British species, Capronia sexdecemspora (Cooke) Saccardo, Sylloge
Fungorum 2:289 (1883).

BERLESIELLA Saccardo

Berlesiella Saccardo in Revue mycologique 10:7 ( J a n u a r y 1888).


There is one British species:

Berlesiella nigerrima (Bloxam ex Currey) Saccardo, op. cit.:7 (1888).


(Fig. 1 6 E . )
S t r o m a t a more or less hemispherical, up to 1 m m . across, covered
with numerous partially immersed black pseudothecia so as to look
like minute blackberries; minutely downy, individual pseudothecia
are only about 60fi across. Asci cylindric-clavate, short-stalked, up to
6 0 x 1 7 / / , rather thick-walled above, 8-spored; ascospores biseriate or
irregularly arranged, somewhat clavate, 1 5 - 2 2 x 5 - 6 / / , with up to five
cross septa and a longitudinal one, light olive-grey.

On old stromata of Eutypa in spring. Not common.

Family 2. Yerrucariaceae

The family contains lichenised fungi and a few lichen parasites, viz.:

I. Asci 8-spored:
A. Ascospores 1-septate, brown Discothecium (p. 372)
B. Ascospores multiseptate or hyaline:
1. Ascospores hyaline Pharcidia (p. 371)
2. Ascospores brown Phaeospora (p. 371)
II. Asci with more than 8-spores:
A. Ascospores 1-septate Tichothecium (p. 372)
B. Ascospores small and nonseptate (not described here)
Muellerella

"370
The shape of the asci and their greatly thickened walls will inevi-
t a b l y lead the student to seek these fungi among the Loculoascomy-
cetes. It appears, however, t h a t the asci are unitunicate, perhaps a
specialised form associated with the lichenised habit, compare
Lecanorales.

P H A R C I D I A Korber

Pharcidia KSrber, Parerga lichenologica: 469 (1865).


Pseudothecia small, immersed in lichen thalli or apothecia, asci
thick-walled, 4- or 8-spored, ascospores hyaline, 1-3-septate.

Pharcidia epicymatia (Wallroth) W i n t e r in Rabenhorst's Krypto-


gamen Flora Deutschland 1 ( 2 ) : 3 4 2 ( P L X X X V I I B . )
Pseudothecia scattered or crowded, subglobose, black, up to 100/t
across, immersed in or erumpent from the hymenium of lichen apo-
thecia. Asci clavate or ventricose, thick-walled, subsessile, u p to
4 5 x 1 5 / / , 8-spored; ascospores in two or three rows, elliptic-cylindric,
with four oil drops, at first 1-septate, sometimes ultimately 3-septate,
9.5-14x2-4.5//.
On the apothecia of many species of Caloplaca and Lecanora, figured on
apothecia of Lecanora subfasca; also found on thalli of other lichen genera.
The systematic position of the genus is uncertain, it might possibly be better
referred to the Mycosphaerellaceae.
For descriptions of the numerous other species see Keissler in Raben-
horst's Kryptogamenflora 8 (1930).

P H A E O S P O R A Hepp

Phaeospora Hepp, Die Flechten E u r o p a s 16, No. 947 (1867).


Pseudothecia scattered, minute, immersed, asci thick-walled, 4- or
8-spored, ascospores 3- or more septate, b r o w n ; parasitic on lichens.

Phaeospora parasitica (L5nnroth) Arnold in Flora 5 7 : 5 1 (1874). (PL


XXXVIIF.)

Pseudothecia scattered, immersed with erumpent ostioles, about


% m m - across, subglobose, black. Asci cylindric-clavate, subsessile,
thick-walled, up to 7 0 x 2 0 / t , 8-spored; ascospores biseriate, elliptical,
1 0 - 2 3 x 4 - 9 / t , mostly 3-septate, brown.
On thalli of species of Bacidia, Biatorella, Buellia, Lecanora, Lecidia and
Bhizocarpon; figured on thallus of Lecidia granulosa.

"371
For other species of Phaeospora see the monograph by Keissler cited
under Pharcidia.

DISCOTHECIUM Zopf
Discothecium Zopf in Nova Acta Leop. Carol. Acad. 70:131 (1897).
Pseudothecia more or less immersed, spherical to obconical, minute,
dark coloured, asci 8-spored, ascospores brown, 2-celled; parasitic on
lichen thalli.

Discothecium gommiferum (Taylor) Vouaux in Bull. Soc. mycol.


France 2 9 : 4 6 ( 1 9 1 3 ) . ( P I . X X X V I I A . )
Pseudothecia scattered, globose with erumpent papillate ostioles,
black, up to 7 5 m m . across. Asci clavate, subsessile, up to 5 0 x 1 2 / / ,
thick-walled above, 8-spored; ascospores irregularly biseriate, broadly
elliptical, 8 - 1 2 x 4 - 6 / / , with one median septum, not constricted,
brown.
On thalli of many genera of crustose lichens. The two large black fructi-
fications in the figure are apothecia of the lichen Lecidia crustulata.
Several other species are recognised, on various lichen genera; see
especially the monograph of this and other lichen parasites by Keissler, "Die
Flechtenparasiten", issued as Volume 8 of Rabenhorst's Kryptogamen-
Flora (1930).

TICHOTHECIUM Flotow
Tichothecium Flotow apud Massalongo Neogenia L i c h e n u m : 8 (1854).
Differs from Discothecium only in its many-spored asci. The two
genera are frequently united, in which case the name Tichothecium has
priority.

Tichothecium pygmaeum Koerber in Denkschr. Feier 50jahr. Best,


schles. Gesellsch. f. vaterl. Cult.:236 (1853). ( P I . X X X V I I E . )
Pseudothecia scattered, globose, black, immersed in the host, vary-
ing greatly in size from 50 to 400/< across. Asci cylindric-clavate, also
very variable, up to 9 5 x 2 3 / / , thick-walled above, subsessile, containing
up to 100 spores; ascospores irregularly arranged, broadly elliptical,
4 - 1 2 x 2 - 6 / / , with one median septum b u t only slightly constricted,
brown.
On thalli of many genera of crustose lichens; figured on Caloplaca auran-
tiaca, with an apothecium of the lichen in the upper segment.

"372
Family 3. Botryosphaeriaceae

This is a large family, characterised b y having hyaline or coloured


non-septate ascospores in bitunicate asci, enclosed in pseudothecia
which m a y be scattered or aggregated in stromata. Many of the genera
are exotic or even tropical and only the type-genus Botryosphaeria will
concern us here. For a thorough t r e a t m e n t of the family see J. A. von
Arx & E. Muller, "Die G a t t u n g e n der amerosporen P y r e n o m y c e t e n "
issued as Beitrage zur Kryptogamenflora der Schweiz 11 (1) (1954).

BOTRYOSPHAERIA Cesati & de Notaris

Botryosphaeria Cesati & de Notaris in Commentario Societa Critto-


gamologica Italiana 1:211 (1863).

Pseudothecia 1 / 5 m m . across or larger, globose, scattered or aggregat-


ed in large or small cushion-like stromata, ostioles papillate, asci
clavate, bitunicate, with numerous pseudoparaphyses, ascospores
hyaline or nearly so, non-septate, usually averaging over 18// long
(similar fungi with smaller ascospores m a y be sought in Guignardia,
not f u r t h e r dealt with here).

Botryosphaeria dothidea (Mougeot ex Fries) Cesati & de Notaris, op.


cit.:212 (1863). (PI. X X X V I I I E . )

S t r o m a t a forming blackish, erumpent, circular or elliptical crusts,


up to 1.5 cm. long, usually more or less distinctly marked with con-
centric ridges and furrows. Pseudothecia very numerous, usually in
small groups, rather less t h a n % m m . across, black, globose, embedded
in the brown tissue of the stroma and erumpent by their papillate
ostioles. Asci cylindric-clavate, subsessile, up to 110x22//, thick-walled
above, 8-spored; ascospores irregularly biseriate, elliptical, 20-24x
8 - 1 0 / / , hyaline to faintly yellowish, non-septate.

On bark of various woody plants, figured on its type host, Rosa, on which
it is common, but according to von Arx & Muller the same species occurs on
many trees and shrubs, sometimes almost without a stroma.
The rather similar Rotryosphaeria quercuum (Schweinitz) Saccardo,
Sylloge Fungorum 1:456 (1882), has larger ascospores 24-42x10-18//; it
has an equally wide host range on woody plants but mainly in warmer
climates and probably does not occur in the British Isles.

"373
Botryosphaeria festucae (Libert) von Arx & Miiller in Beitrage zur
Kryptogamen Flora der Schweiz 11 (1):38 (1954). (PI. X X X V I I I c . )
Pseudothecia scattered, about % m m - across, black, spherical with
conical erumpent ostiole, thick-walled, immersed in the mesophyll of
dead leaves and accompanied by a web of olive to dark brown hyphae.
Asci clavate, up to 100x20//, rather tliick-walled at the tip, 8-spored;
ascospores biseriate, narrowly elliptical, 2 2 - 3 5 x 8 - 1 3 / / , hyaline or
slightly yellowish, non-septate.

In dead leaves and culms of various grasses and sedges. Common, figured
on Molinia.

Botryosphaeria foliorum (Saccardo) von Arx & Miiller, op. cit.:42


(1954). (PI. X X X V I I I J . )
Pseudothecia scattered, globose, black, scarcely % m m . across,
ostiole not noticeably papillate. Asci clavate, subsessile, up to 100x
25//, thick-walled above, 8-spored; ascospores uniseriate or irregularly
biseriate, slightly ovate and narrowed below, 15-24x7-12/«, hyaline,
non-septate.
On dead leaves and twigs of Taxus baccala, erumpent mainly on the upper
surface of the leaf. Not uncommon, parasitic; also known as Physalospora
gregaria Saccardo var. foliorum Saccardo, Sylloge Fungorum 1:435 (1882).

Family 4. Yenturiaceae

This family, as defined b y von Arx, includes Loculoascomycete


genera with 1-septate ascospores which are at first hyaline to greenish,
eventually olive-brown or greyish-green b u t never opaque; the asco-
spores usually have the septum a little displaced from the middle, so
t h a t one cell is longer t h a n the other; the asci are usually rather small,
bitunicate but without a massive apical thickening, and are inter-
spersed with pseudoparaphyses, at least in the unripe ascocarp. Some
b u t not all of the species have short dark hairs, especially round t h e
ostiole. Von Arx included 12 genera in the family b u t only six of these
concern us here, all parasites of flowering plants:

I. Pseudothecia sunken in the host tissue, subepidermal, scattered, ostioles


papillate with or without hairs Venturia (p. 375)
II. Pseudothecia subcuticular, erumpent or superficial:
A. Pseudothecia subcuticular Stigmatea (p. 378)
B. Pseudothecia erumpent or superficial and associated with subcuti-
cular mycelium:

"374
1. Pseudothecia small, smooth, seated on stomata of conifer
needles Phaeocryptopus (p. 378)
2. Pseudothecia not on conifers, not scattered or not smooth:
a. Pseudothecia in a ring round a sclerotium-like stroma
Lasiobotrys (p. 379)
b. Pseudothecia without a sclerotium-like stroma; hairy:
+Pseudothecia associated with a single layer of
hyphae beneath the host cuticle, on Rosaceae
and Geraniaceae Coleroa (p. 376)
+ +Pseudothecia associated with a thicker hyphal
crust, British species on Ericaceae and Capri-
foliaceae Gibbera (p. 377)

Compare also Euryachora betulina & E. ulmi, p. 361, which von Arx re-
gards as stromatic members of the Venturiaceae, in the genera Atopospora
and Platychora respectively.

VENTURIA de Notaris

Venturia de Notaris in Giornale Botanico Italiano 1:332 (1844),


emend Saccardo, Sylloge F u n g o r u m 1:586 (1882).
The essential generic characters have been sufficiently indicated in
the key. The species fall into two series, treated b y some authors as
independent genera; Endostigme in which there is a conidial state
growing parasitically on living leaves whereas the pseudothecia
develop in the fallen overwintered leaves, shooting out their ascospores
in time to reinfect the new season's leaves as t h e y unfold in spring.
Spilosticta in which there is no conidial state b u t pseudothecia develop
in spots on living leaves. The t y p e species of Venturia de Notaris was
V. rosae de Notaris, which Saccardo excluded from his emended con-
cept of the genus and which has been renamed Gibbera rosae (de
Notaris) Miiller & Menon in Phytopathologische Zeitschrift 25:194
(1955).

Venturia ditricha (Fries) Karsten, Mycologia Fennica 2:188 (1873),


on Betula, with ascospores 1 2 - 1 6 x 5 - 6
Venturia inaequalis (Cooke) W i n t e r apud Saccardo, Sylloge F u n -
gorum 1:587 (1882), emend Aderhold in Hedwigia 36:81 (1896), on
Malus, with conidial state Spilocaea pomi Fries on the living leaves
and fruit, causing t h e common disease "Apple Scab".
Venturia pirina Aderhold in Ldw. J a h r b . 25:875 (1896) and Hed-
wigia 36:80 (1896), on Pyrus, with conidial state Fusicladium pirinum
(Libert) Fnckel causing " P e a r Scab" on living leaves and fruit.

375
These two well-known plant parasites have been described too
often to be dealt with in detail or figured here.

Venturia (Spilosticta) maculaeformis (Desmazieres) Winter in Raben-


horst's K r y p t . Flora Deutschland 1 (2): 435 (1885). (Fig. 21H.)
Pseudothecia clustered in the tissue of small, dead, brown spots with
narrow, purple margins, on still living leaves, black, very minute, with
papillate ostioles fringed with short brown bristles. Asci clavate or
slightly ventricose, subsessile, up to about 4 0 x 1 2 / / , thick-walled
above, 8-spored; ascospores irregularly biseriate, slightly clavate,
9 - 1 2 x 3 . 5 - 4 / / , pale greenish, 1-septate.

On leaves of Epilobium montanum. Common.


Also called Sphaeria microspila Berkeley & Broome in Ann. Mag. nat.
Hist. I l l : 455 (1861).

Venturia geranii (Fries) W i n t e r in Rabenhorst 1 (2): 434 (1887) is


common on leaves of Geranium, sylvaticum.

Venturia rumicis (Desmazieres) Winter, op. cit.:435 (1885). (Fig. 21G.)


Pseudothecia clustered in small, dead, brown spots, often outlined
b y a purple line, black, u p to 140// across, with erumpent ostioles
fringed with short brown bristles. Asci clavate to ventricose, up to
55 x l 7 / / , thick-walled above, 8-spored; ascospores biseriate, slightly
clavate, 1 2 - 1 5 x 4 - 7 / / , 1-septate, grey-green.

On living and faded leaves of species of Rumex. Very common everywhere.

COLEROA (Fries) Rabenhorst

Coleroa (Fries) Rabenhorst in Botanisch Zeitung 9, Column 180 (Fe-


b r u a r y 1851).
Pseudothecia like those of Venturia b u t superficial and bearing
bristles over their entire surface.

Coleroa chaetomium (Kunze ex Fries) Rabenhorst, op. cit. (1851).


(Fig. 2 1 J . )
Pseudothecia scattered or in small clusters, often on small brown
spots, superficial, black, about 150// across, covered with short, stiff,
spreading, brown bristles. Asci clavate, subsessile, up to 6 0 x 1 2 / / ,

"376
thick-walled above, 8-spored; ascospores biseriate, slightly clavate,
1 1 - 1 3 x 5 - 6 / / , 1-septate, grey-green.
On living leaves of Rubus idaeus and R. caesius. Not uncommon.

Coleroa alchemillae (Greville) W i n t e r in Rabenhorst's K r y p t o g a m e n


Flora Deutschland 1 (2): 199 (1885). (Fig. 21K.)
Pseudothecia gregarious, usually arranged in rows radiating from a
common centre, black, superficial, up to 110// across, bearing short
brown bristles. Asci clavate, subsessile, up to 4 5 x 1 0 / / , thick-walled
above, 8-spored; ascospores biseriate, slightly clavate, 8 - 1 1 x 3 - 4 / i ,
1-septate, hyaline.
On living leaves of Alchemilla vulgaris agg. Common.
Coleroa potentillae (Sowerby ex Fries) Winter, op. cit.:99 (1885),
occurs on leaves of species of Potentilla.
Coleroa circinans (Fries) W i n t e r in Rabenhorst's Kryptogamenflora
1 (2): 200 (1887), of which Venturia glomerata Cooke in Grevillea 3 : 6 9
(September 1874) is a synonym, occurs in dead spots on living leaves of
Geranium dissectum, G. molle, G. pusillum etc., with ascospores 10-12x
5 fi.

GIBBERA Fries

Gibbera Fries Floram Scanicam:347 (1835).


Pseudothecia clustered on an erumpent black stroma, bristly, asci
cylindrical, ascospores 1-septate, yellowish. There are two British
species:

Gibbera vaccinii (Sowerby ex Fries) Fries, S u m m a Veg. Scand. Sect.


Post.:402 (1849). (Fig. 21L.)
Stroma forming a dark-brown erumpent crust, covered with densely
clustered black pseudothecia, each globose, up to 300// across, covered
with dark brown spreading bristles up to 60// long. Asci narrowly
cylindrical, short-stalked, up to 130x12/t, the wall not thickened at
the apex, 8-spored; ascospores uniseriate-elliptical, 1 2 - 1 8 x 5 - 8 / / ,
1-septate, yellowish.
On leaves and twigs of Vaccinium vitis-idaea. Uncommon.

Gibbera myrtilli (Cooke) P e t r a k in Sydowia 1:200 (1947). (Fig. 211.)


Pseudothecia scattered, superficial, subglobose or slightly flattened
above, about 150// across, black, thickly beset with short, stiff,

"377
pointed bristles. Asci cylindric-clavate, subsessile, not very thick-
walled, up to 5 0 x 1 4 / / , 8-spored; ascospores biseriate, slightly clavate,
1 0 - 1 4 x 4 - 5 f i , 1-septate, pale greenish.
On dead leaves ot Vaccinium myrtillis and V. uliginosum.

Gibbera dickiei (Berkeley & Broome) von Arx in Tijdschrift over


Plantenziekten 58:265 (1952), is a similar fungus on Linnaea borealis.

STIGMATEA Fries

Stigmatea Fries, Floram Scanicam:347 (1835).


Differs from Venturia and Coleroa in its hemispherical pseudothecia
surrounded by a rim of dark, radiating, compact hyphae beneath the
cuticle of the living host leaf.

Stigmatea robertiani (Fries) Fries, S u m m a Veg. Scand. Sect. Post.: 421


(1849). (Fig. 22A.)
Pseudothecia clustered, in small spots on the u p p e r surface of living
leaves, hemispherical, superficial, black, perfectly smooth, up to 140/<
across. Asci usually somewhat ventricose, up to 4 5 x 1 4 / / , thick-walled
above, 8-spored; ascospores irregularly biseriate, somewhat clavate,
1 0 - 1 4 x 4 - 5 / / , 1-septate, the lower cell considerably shorter t h a n the
upper, pale olive. A portion of the basal flange of the pseudothecium is
illustrated separately x660.
On Geranium robertianum. Very common. Transferred to Coleroa by
Miiller in Miiller & von Arx 1962. This must not be confused with a species
of Venturia found on other species of Geranium.

PHAEOCRYPTOPUS N a u m o v

Phaeocryptopus N a u m o v in Bull. Soc. Oural d'Amis de Sci. Nat. 35:20


(1915).
There is only one British species:

Phaeocryptopus gaeumannii (Rohde) P e t r a k in Annales mycologici 36:


22 (1938). (PL X X X V I I I B . )
Pseudothecia scattered, superficial, globose, up to 80/t across, black,
smooth, each seated on a t i n y peg of tissue inserted in a stoma on the

"378
under side of a needle of the host and attached by this to a slender
olive-coloured mycelium within the leaf. Asci clavate to slightly
ventricose, subsessile, up to 4 0 x 1 5 / / , wall slightly thickened above,
8-spored; ascospores irregularly biseriate, slightly clavate, 1 0 - 1 5 x 3 . 5 -
5/<, 1-septate, hyaline.
On living and fallen needles of Pseudotsuga. Not uncommon.

LASIOBOTRYS Kunze

Lasiobotrys Kunze, Mykologische Ilefte 2 : 8 8 (1823).


Pseudothecia attached to a sclerotium, hairless, sclerotia clustered
on a p a t c h of dark superficial mycelium. There is probably only one
species:

Lasiobotrys lonicerae Kunze, op. cit.:89 (1823). (PI. XXXVIIIA.)

Mycelial m a t black, superficial, communicating with mycelium


within the leaf, in approximately circular patches up to 4 m m . across.
Sclerotia top-shaped, top nearly flat, smooth, black, up to J / 3 m m .
across, covering the surface of the mycelial m a t and anchored to it by
numerous dark brown hyphae descending from the upper edge like the
guy ropes of a t e n t ; pseudothecia globose, blackish-brown, smooth,
about 75// across, attached laterally in a ring round the side of the
sclerotium. Asci few, cylindric-clavate, sessile, up to 5 0 x 1 4 / / , thick-
walled above, 8-spored; ascospores slightly clavate, 8 - 1 0 x 4 - 5 / / ,
1-septate, hyaline. The sclerotia, with pseudothecia attached, are shot
off at m a t u r i t y so t h a t it is difficult to find m a t u r e asci in those still
on the host leaf.
On living leaves of Lonicera. Fairly common.
Though fully exposed at maturity the mycelial mat can be seen to
originate beneath the cuticle as in so many Venturiaceae. This extra-
ordinary genus is traditionally placed with the Erysiphales but its asci are
clearly bitunicate.

Family 5. Lophiostomataceae

This is a small family in which the more or less immersed pseudo-


thecia are easily recognised b y their elongated slot-like ostioles. The
genera are distinguished b y the conventional ascospore characters:
1. Ascospores non-septate, brown Lophiella
II. Ascospores 1-septate:
A. Ascospores hyaline Lophiosphaeria

"379
B. Ascospores brown:
1. Pseudothecia crowded on a subiculum . . . Byssolophis
2. Not so Ostropclla
III. Ascospores with several cross septa:
A. Ascospores hyaline Lophiotrema
B. Ascospores brown Lophiostoma (p. 381)
IV. Ascospores brown with both transverse and longitudinal septa
Lophidium (p. 381)
V. Ascospores needle-shaped or worm-like Lophionema

Only four of these concern us here, Lophiotrema Byssolophis, Lophiostoma


and Lophidium.

LOPHIOTREMA Saccardo

Lophiotrema Saccardo in Michelia 1 : 3 3 8 (July 1878).


The characters of the genus are sufficiently indicated in the key. The
n u m b e r of species is uncertain as the genus has not been revised in
modern times:

Lophiotrema praemorsum (Lasch) Saccardo, op. cit.: 338 (1878).


(Fig. 13A.)
Pseudothecia immersed in dead wood, flask-shaped with the upper
half erumpent, up to % m m . across, smooth, black, neck short, laterally
compressed with a slot-like ostiole almost as long as the pseudo-
thecium is wide. Asci cylindric-clavate, short-stalked, up to 115x12//,
thick-walled, 8-spored; ascospores biseriate, fusiform, 2 4 - 2 8 x 5 - 7 / / ,
noticeably constricted at the median septum, commonly only 1-sep-
t a t e b u t said to develop up to five septa, hyaline. Figured on Ulex.

On dead branches of woody plants, especially Ulex, on Rubus and on


dead herbaceous stems and grasses.
In the absence of a modern critical revision I have quoted the wide inter-
pretation of the species suggested by Berlese; the form on Ulex is sometimes
distinguished as L. angustilabrum (Berkeley & Broome) Saccardo, op. cit.: 338
(1878). This fungus well exemplifies the unsatisfactory character of genera
based solely on ascospore septation for most of the British material under
this name would on that basis be referable to Lophiosphaeria, though the
constant presence of two or more large oil drops in each cell may lead one
to suspect additional septation at maturity.

Lophiotrema nucula (Fries) Saccardo, op. cit.:338 (1878), on bark of


Quercus and other trees, has 3-septate ascospores, 1 8 - 2 0 x 6 / / .
Lophiotrema vagabundum Saccardo, op. cit.:338 (1878), on herba-
ceous stems, has 3-septate ascospores, 2 2 - 2 5 x 4 - 6 / / .

"380
LOPHIOSTOMA (Fries) Cesati & de Notaris

Lophiostoma (Fries) Cesati & de Notaris in Commentario Societa


Crittogamologica Italiana 1:219 (1863).
Differs from Lophiotrema in its brown ascospores.

Lophiostoma caulium (Fries) Cesati & de Notaris, op. cit.: 219 (1863).
(Fig. 13B [lower spore].)
Pseudothecia immersed, scattered, flask-shaped, a b o u t % m m .
across, smooth, black, with erumpent slit-like ostioles. Asci narrowly
clavate, short-stalked, up to 9 0 x 1 2 / / , 8-spored, thick-walled; asco-
spores biseriate, fusiform, 1 8 - 2 2 x 6 - 7 / / , 5-septate, slightly constricted
at the septa, olive-brown.

On dead herbaceous stems. Not uncommon.


Many other species have been described but it is not clear how many are
truly distinct. One with 7-septate ascospores about 26x6// has been con-
fused with L. caulium; this may be L. niessleanum Saccardo in Michelia 1:
447 (November 1878), see Fig. 13B (upper spore).

LOPHIDIUM Saccardo

Lophidium Saccardo in Michelia 1:340 (July 1878).


There is only one common British species:

Lophidium compressum (Persoon ex Fries) Saccardo, op. cit.:340


(1878). (Fig. 13c.)
Pseudothecia scattered, immersed, flask-shaped, about % m m -
across, with short erumpent necks and long slit-like ostioles, black.
Asci cylindric-clavate, short-stalked, up to 110x16//, 8-spored; asco-
spores elliptic-fusiform, often slightly constricted in the middle,
20-32x7-10/«, with up to nine transverse septa and one longitudinal
septum dividing at least some of the transverse segments, brown.

On decorticated wood, especially of Salix. Not uncommon on that host.

BYSSOLOPHIS Clements

Byssolophis Clements Genera of Fungi 286 (1931).


There is one British species separated from Glonium by its asco-
spores becoming brown at m a t u r i t y .

"381
Byssolophis sphaerioides (Karsten) Miiller in Beitr. Krypt.-Fl. Schweiz
11 (2) 341 (1962). (Fig. 15 H.)
Pseudothecia usually crowded, r a t h e r broadly elliptical, up to
1.4x0.5 mm., smooth, black, associated with a thin m a t of brown
hyphae. Asci cylindric-clavate, short-stalked, up to 120x13/«, thick-
walled, 8-spored; ascospores mostly uniseriate, elliptical, 15-19x5-6/«,
1-septate and constricted at the septum, long remaining hyaline b u t
becoming light brown at m a t u r i t y .

On wood or bark. Apparently very rare. Also known as Glonium amplum


(Berkeley & Broome) Duby in Mem. Tribu Hyst.:37 (1861).

Family 6. Pleosporaceae

Here are collected the remaining genera of Pleosporales, a hetero-


geneous assemblage no d o u b t capable of f u r t h e r subdivision when
more adequately studied. For the present it m u s t suffice to arrange
t h e m along the traditional lines dictated b y the usual ascospore
characters.

I. Ascospores 1-septate, hyaline (see also Venturiaceae):


A. Pseudothecia superficial on a subiculum Horpotrichia (p. 410)
B. Pseudothecia immersed Didymella (p. 385)
(For apparently 1-septate, small, hyaline ascospores in pseudo-
thecia on lichens see Pharcidia, p. 371)
II. Ascospores 1-septate, dark brown:
A. Ascospores unevenly septate, lower cell smaller
Pteridiospora (p.407)
B. Ascospores with a median septum:
1. Pseudothecia erumpent in clusters from bark, associated with
Diplodia pycnidia Otthia (p. 415)
2. Not as above:
a. Pseudothecia on herbaceous stems
Didymosphaeria (p. 385)
b. Pseudothecia on dung:
Ascospores very large, with germ slits
Delitschia (p. 412)
Ascospores small, without germ slits
Trichodelitschia (p. 412)
c. Pseudothecia superficial on wood:
/Ascospores over 20/« long, pseudothecia conical on a
broad base Astrosphaeriella (p. 405)

"382
//Ascospores up to 15//, pseudothecia spherical,
amongst conidiophores of Dematiaceous moulds
Letendraea (p. 386)

Ascospores 2- or more septate:


A. Parasitic on lichens, see Verrucariaceae
B. Not on lichens:
1. On dung; ascospores dark brown, multiseptate
Sporormia (p. 411)
2. On other substrata:
a. On herbs:
:Ascospores with two septa towards one end
Buergenerula (p. 387)
::Ascospores with three or more septa:
*Ascospores hyaline, pseudothecia with hairs
round the ostiole Keissleriella (p. 388)
**Ascospores yellow or brown:
tAscospores with a hyaline appendage or
terminal hyaline cells:
§Ascospores with a hyaline basal ap-
pendage Rebentischia (p. 402)
§§Ascospores with rounded hyaline ter-
minal cells Passeriniella (p. 403)
t f Ascospores without an appendage:
§Ascospores with a thick double wall
Chitonospora (p. 403)
§§Ascospores not as above
Leptosphaeria (p. 389)
b. On wood or b a r k :
:Pseudothecia immersed or partly so:
f Ascospores hyaline:
§Ascospores 2-septate, small
Melomastia (p. 406)
§§Ascospores 3-septate, large
Massarina (p. 406)
ttAscospores coloured:
*Ascospores 3-septate, end cells hya-
line with appendages
Broomella (p. 387)
**Not as above:
§Pseudothecia erumpent from
dead wood, ascospores without
a gelatinous coat
0Ascospores over 50 long,
markedly apiculate
Caryospora (p. 405)

"383
0 0 Ascospores much smaller,
not apiculate
Trematospliaeria (p. 404)
§§Pseudothecia immersed in bark,
ascospores very large, with a
gelatinous coat
0 Pseudothecia free, associat-
ed with long, multi-sep-
tate brown conidia
Asteromassaria (p. 408)
0 0Pseudothecia in a stroma,
often clustered
Massaria (p. 407)
: :Pseudothecia erumpent in clusters, enveloped
in a hyphal web Thyridaria (p. 409)
:::Pseudothecia superficial:
f W i t h a subiculum, ascospores well over
20 fi long or fusiform:
§Ascospore end cells hyaline
Thaxteria (p. 410)
§§Entire ascospore hyaline
Herpotrichia (p. 410)
t f W i t h o u t a subiculum or only a slight hy-
phal mat, ascospores elliptical, less than
20 p long . . . Melanomma (p. 409)
IV. Ascospores with both transverse and longitudinal septa; always brown:
A. Pseudothecia scattered on herbaceous plants:
1. Pseudothecia without bristles . . . . Pleospora (p. 413)
2. Pseudothecia with bristles . . . . Pyrenophora (p. 414)
B. Pseudothecia in or on wood or bark:
1. Pseudothecia in or on a stroma:
a. Pseudothecia enclosed in a small stroma, their ostioles
collectively erumpent Fenestella (p. 416)
b. Pseudothecia clustered on a basal stroma
Cucurbitaria (p. 414)
2. Pseudothecia not in or on a stroma, immersed in bark:
a. Clustered and enveloped by a dark mycelium
Karstenula (p. 417)
b. Subsolitary, ascospores enormous with gelatinous coats
Pleomassaria (p. 417)
c. Not clustered, ascospores small Thyridium (p. 418)
3. Pseudothecia gregarious, more or less superficial on wood
Teichospora (p. 416)
V. Ascospores needle-shaped or worm-like:
A. Pseudothecia clustered on a stromatic base Naumovia (p. 421)
B. Pseudothecia solitary, without a stroma:
1. Pseudothecia with apical appendages, associated with a
helicosporous conidial state, on marsh plants
Tubeufia (p. 421)
2. Pseudothecia covered with stiff bristles
Acanthophiobolus (p. 420)

"384
3. Pseudothecia without appendages or bristles:
a. Ascospores long and very slender, without a swollen cell
Leptospora (p. 418)
b. Ascospores having one or more cells broader than those
adjacent Ophiobolus (p. 419)
C. Pseudothecia parasitic on scale insects, gregarious on a byssoid
stroma Podonectria (p. 420)

DIDYMELLA Saccardo

Didymella Saccardo in Michelia 1 : 3 7 7 (November 1878).


Pseudothecia small, scattered, immersed, mainly in dead herbaceous
stems, ascospores 2-celled, hyaline; there is often an associated conidial
state in the form-genera Ascochyta, Diplodina or Phoma.

Didymella applanata (Niessl) Saccardo, Sylloge F u n g o r u m 1:546


(1882). (PL X X X V I I I G . )
Pseudothecia scattered, immersed with erumpent ostioles, sub-
globose, about y 4 m m . across, almost black. Asci cylindric-clavate,
short-stalked, up to 70x13/«, thick-walled above, 8-spored; ascospores
biseriate, slightly clavate, 12-18x5-6/«, 1-septate, hyaline.

On grey patches of skin, especially at the nodes, on canes of Rubus idaeus,


associated with the disease called "Spur Blight". Common in spring. The
conidial state is a Phoma.

Didymella pinodes (Berkeley & Bloxam) P e t r a k in Annales myco-


logici 22:18 (1924), with ascospores 1 5 - 2 0 x 6 . 5 - 8 / / , occurs on Pisum
sativum as one of the causes of foot-rot, leaf-spot and pod spot.

DIDYMOSPHAERIA Fuckel

Didymosphaeria Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:


140 (1870).

Pseudothecia scattered, immersed, subglobose, dark coloured, some-


times associated with slight development of a clypeus around the
ostiole, asci 8-spored, ascospores uniseriate, 1-septate, brown; on dead
twigs or herbaceous stems.

"385
Didymosphaeria diplospora (Cooke) Rehm in Hedwigia 18:167 (1879).
(PI. X X X V I I I F . )
Pseudothecia scattered, subglobose, up to % m m . across, immersed,
ostioles erumpent and each surrounded b y a small brown-stained patch
of epidermis. Asci cylindrical, subsessile, up to 8 5 x 9 / / , wall somewhat
thickened above, 4- to 8-spored; ascospores uniseriate, elliptic-
cylindric, 9 - 1 2 x 4 - 7 / / , 1-septate and usually slightly constricted at
the septum, brown, the wall minutely w a r t e d .
On dead canes of Iiubus fruticosus, said to occur also on Cornus. Not un-
common.

Didymosphaeria futilis (Berkeley & Broome) Rehm in Hedwigia 18:


167 (1879) with ascospores 8 - 1 5 x 3 - 5 / / , occurs on herbaceous stems
and Typha.
Didymosphaeria conoidea Niessl in Osterr. Bot. Zeitschr. 25:202
(1875) has ascospores 8 - 1 2 x 4 - 5 / / , in pseudothecia superficial on
herbaceous stems, often amongst Leptosphaeria doliolum.
Didymosphaeria winteri Niessl, op. cit.: 165 (1875), with ascospores
1 1 - 1 4 x 4 - 6 / / , develops in s t r o m a t a of other ascomycetes, especially
Phyllachora and Ophiobolus.
Many other species have been described but some, at least, belong in very
different genera, such as Ceriophora,Didymella, Leptosphaeria and Sydowiella-,
for a modern treatment see Scheinflug in Schweiz. Bot. Ges. 68, 1958.

LETENDRAEA Saccardo

Letendraea Saccardo in Michelia 2 : 7 3 (April 1880).


Perithecia superficial, scattered, brightly coloured, smooth, soft,
ascospores 1 -septate, becoming brown. There is a single British species:

Letendraea helminthicola (Berkeley & Broome) Weese in Mitt. Bot,


Techn. Hochsch. Wien 1 : 5 9 ( 1 9 2 4 ) . (PL X X X I K . )
Perithecia globose to pear-shaped, solitary or in small groups, flesh-
coloured or light yellow, less t h a n 0.2 m m . across and scarcely visible
to the naked eye. Asci clavate, up to 7 0 x 1 4 / / , 8-spored; ascospores
biseriate, 1 2 - 1 5 x 4 . 5 - 6 / / , oblong-fusoid, 1-septate and markedly con-
stricted at the septum, smooth, light brown at m a t u r i t y .
Amongst the densely crowded dark brown to blackish conidiophores of
the mould Helminthosporium appendieulatum Cda. in winter. Inconspicuous
and seldom recorded.

"386
BROOMELLA Saccardo

Broomella Saccardo, Sylloge fungorum 2:557 (1883). (Fig. 20c.)


Perithecia in small clusters on a basal tissue, subepidermal; asco-
spores brown, 3-septate, with terminal appendages. The conidial
states belong to the form-genus Pestalotia de Notaris, see Shoemaker
& Miiller in Canad. J o u r n . Bot. 41:1235-1243 (1963).

Broomella vitalbae (Berkeley & Broome) Saccardo, op. cit.:558 (1883).


Perithecia subepidermal, in small clusters on a basal stromatic
tissue, causing vague pimples in the host epidermis, each with a central
black dot formed by the erumpent ostiole, dark brown, thin-walled,
subglobose, smooth, about y 3 m m . across. Asci narrowly cylindrical,
thin-walled, apex not blued b y Melzer's reagent, 110-175x10/«,
8-spored; ascospores overlapping, spindle-shaped, 26-36x5-7/«, 3-sep-
t a t e , the central cells light brown, the end ones colourless, bearing a
slender hyaline bristly appendage about 10fi long at each end.

On dead stems of Clematis vitalba in winter.


This is the type species of the genus and also the same as that of Keissleria
von Hohnel in Annales Mycologici 16:93, 1918, under the name K. xantha
(Saccardo) von Hohnel.

BUERGENERULA Sydow

Buergenerula Sydow in Annales mycologici 34:392 (1936).


Pseudothecia immersed, scattered, minute, asci thick-walled,
8-spored, ascospores with two septa towards one end. There is only
one species:

Buergenerula biseptata (E. Rostrup) Sydow, op. cit.:392 (1936). (Fig.


10H.)

Pseudothecia gregarious on dead patches on leaves, immersed, glo-


bose, up to 1 / 5 m m . across, with erumpent ostioles, each with a rudi-
m e n t a r y clypeus. Asci clavate, short-stalked, up to 120x23/«, thick-
walled, 8-spored; ascospores more or less biseriate, elliptic-fusiform,
21-30x7-10/«, with two septa near the lower end, not constricted,
hyaline.
On Carices, typically on Carex vesicaria, in winter.

"387
K E I S S L E R I E L L A von Hohnel

Keissleriella von Hohnel in Sitzber. K. Akad. Wiss. Wien, Math. Nat.


Kl. 128 Abt. 1 : 5 9 2 (1919).
Pseudothecia subepidermal, less often on b a r k , minute, with small
ostiolar papilla bearing short dark brown bristles. Ascospores 1 - 3 or
more septate, hyaline.

Keissleriella culmiflda (Karsten) Bose in P h y t o p a t h . Zeitschr. 4 : 1 8 8


(1961). (Fig. 20G.)
Pseudothecia scattered, subglobose, black, scarcely y 2 m m . across,
with erumpent black ostiolar papillae. Asci cylindric-clavate, 65-100x
10-15//, 8-spored, occasionally only 4-spored; ascospores biseriate,
elliptic-cylindric, slightly curved, 3-septate and constricted at the
septa, 2 2 - 2 6 x 5 - 6 / / , hyaline. The papilla is covered with straight or
slightly curved, dark brown, thick-walled, pointed bristles 2 0 - 4 4 x
3 - 5 //,

On dead grass culms, typically on Phleurn, in late autumn.

Trichometasphaeria culmifida (Karsten) Munk in Dansk Bot. Ark.


17:445 (1957) is a synonym.

Keissleriella linearis Miiller in Kew Bull. 19:120 (1964). (Fig. 25u.)


Pseudothecia immersed, often in short rows, in darkened patches
of the host culm, subglobose or compressed, up to y 2 m m . diameter,
black, bearing brown bristles up to 4 0 x 4 - 5 / / round the ostiole. Asci
numerous, cylindric-clavate, 130-150x18-20//, 8-spored; ascospores
elliptic-fusiform, 4 0 - 4 5 x 7 - 9 / / , 3-septate, slightly constricted at the
septa, hyaline, with a thick mucilaginous coat.

On Phragmites communis.

Keissleriella gallica (Miiller) Bose, op. cit.: 189 (1961). (Fig. 25R.)
Pseudothecia scattered, immersed in the substrate beneath a slightly
blackened piece of epidermis, subglobose, about % m m . diameter, the
ostiolar papilla clothed w i t h short, dark brown, thick-walled bristles
about 2 5 x 4 / / . Asci cylindric-clavate, 9 0 - 1 0 5 x 1 2 / / , 8-spored, asco-
spores irregularly biseriate, elliptic-fusoid, straight or slightly curved,
1 9 - 2 1 x 4 . 5 - 5 / / , 3-septate, hyaline.
On stem of Sparteum junceum and on Rumex acetosa.

"388
Keissleriella subalpina (Rehm) Bose, op. cit.: 183 (1961). (Fig. 25s.)
Pseudothecia scattered, superficial or with the base partly sunk in
the substrate, about % mm. diameter, obconical on a broad base,
thick-walled, black. Asci cylindrical, 40x12fi, 8-spored; ascospores
irregularly biseriate, 12-14x4.5-5jn, 1-septate, the upper cell shorter
and broader than the lower.
On old stems of Calluna vulgaris.

Keissleriella ocellata (Niessl) Bose, op. cit.: 184 (1961). (Fig. 25Q.)
Pseudothecia scattered, subepidermal, subglobose, % mm. dia-
meter, without apical papilla or bristles. Asci numerous, cylindric-
clavate, 60-90x11-12fi, 4-8-spored; ascospores biseriate, ellipsoid,
15-21x6-7/«, 2-septate, long hyaline but finally brownish. On stems
of Hypericum spp.
Metasphaeria hyperici (Phillips & Plowright) Grove is a synonym.

LEPTOSPHAERIA Cesati & de Notaris

Leptosphaeria Cesati & de Notaris in Commentario Societa Crittogamo-


logica Italiana 1:234 (1863).
Pseudothecia scattered, immersed or becoming superficial by shed-
ding of the host tissue, globose to conical with well developed ostioles,
asci thick-walled, 8-spored, ascospores usually somewhat fusiform,
with from two to many cross septa, usually yellowish to yellowish-
brown. This is a large genus, found usually on dead herbaceous stems
and grass culms, Carices and the like. One group of species is charac-
terised by having one cell in the upper half of the ascospore distinctly
broader than the others and on this basis is sometimes treated as a
distinct genus, Nodulosphaeria Rabenhorst. The genus has never been
monographed in the British Isles and the number of British species
is unknown. There is, however, an excellent monograph of the Swiss
species by E. Muller in Sydowia 4:185-319 (1950) (in German) and a
useful summary of the species known on herbaceous plants in Sweden
by L. Holm in Svensk Botanisk Tidskrift 46:18-46 (1952) (in Eng-
lish). Muller described 114 species, most of them figured, and Holm has
62 species; most British collections should be identifiable from their
papers. For fungi on Helianthemum, Iris, Leguminosae or Plantago
with 3-septate ascospores paler at the ends see Trematosphaeria
p. 404.

"389
More recently Holm, in Symb. Bot, Upsal. 14 (3) (1957) has divided
the traditional Leptosphaeria into four genera, with L. doliolum re-
cognised as type species of Leptosphaeria sensu stricto. Nodulosphaeria
Rabenhorst, Herb. Myc. Ed. 2, No. 725 (1858) has for type species t h a t
described below as L. derasa\ a large group of species is segregated in
Phaeosphaeria Miyake, in J . Coll. Agric. Univ. T o k y o : 2 (1910), with
P. oryzae Miyake as type and a small group of species on Leguminosae
is referred to Entodesmium Riess in Hedwigia 1 : 2 8 (1854), with
E. rude Riess as type. He has also transferred several species to
Melanomrna and Trematosphaeria, genera traditionally separated
from Leptosphaeria by growing on wood. Generic distinctions based
on substrate and position of the fruit body in relation to the substrate
are no doubt u n n a t u r a l b u t t h e y arc easy for the inexperienced
student to grasp. I have accordingly retained most of these species
under Leptosphaeria here, with the necessary cross-references. There
remain a few species for which no modern disposition has been pro-
posed, some of which possibly require still other segregate genera in
the future.

Leptosphaeria doliolum (Fries) de Notaris, op. cit.:234 (1863). (PI.


XXXIXG.)

Pseudothecia usually somewhat gregarious, at first immersed, soon


exposed b y shedding of the host epidermis, conical on a broad base,
up to y 2 mm. across, with a prominent apical ostiole, thick-walled,
black, smooth. Asci numerous, cylindric-clavate, up to 140x10//,
short-stalked, 8-spored; ascospores uniseriate, fusiform, usually with
one flattened side or slightly curved, 1 6 - 2 8 x 3 . 5 - 7 / / , 3-septate and
slightly constricted at the septa, yellowish-brown.

On dead herbaceous stems, figured on Angelica. Common.

Leptosphaeria acuta (Fries) Karsten, Mycologia Fennica 2:98 (1873).


(Fig. 16G.)
Pseudothecia scattered or somewhat gregarious, soon exposed by
shedding of the cortical tissues, conical on a broad base, up to 2 / 5 m m .
across, thick-walled, smooth, black, with a prominent apical ostiole.
Asci numerous, cylindric-clavate, short-stalked, up to 140x10//,
8-spored; ascospores biseriate, narrowly fusiform, usually slightly
curved, 3 6 - 5 0 x 5 - 6 / / , with from six to ten cross walls but scarcely
constricted at t h e septa, yellowish.

"390
At the base of dead stems of Urtica dioica. Very common from October
to May but sometimes replaced by the externally similar Phoma acuta
Fuckel, which contains only hyaline elliptical conidia 4-5 x 1-2 p and is the
conidial state, see Miiller in Phytopath. Zeitschr. 29:291 (1957).

Leptosphaeria lycopodina (Montagne) Saccardo Sylloge F u n g o r u m 2 : 8 1


(1883). (PI. X X X I X D . )
Pseudothecia scattered, immersed, globose, with erumpent papillate
ostioles, black, up to 150// across. Asci few, clavate, subsessile, up to
75x15/«, 8-spored; ascospores closely packed in two or three rows,
elliptic-fusiform, 20-26x7-10/«, 3-septate, slightly constricted at the
septa, yellowish.
On dead leaves and bracts of Lycopodium alpinum and L. annotinum;
figured from a collection on the latter host from Meall Gorm, above the
Bealach nam Bo, Applecross, West Ross. Leptosphaeria crepini (Westen-
dorp) de Notaris is a synonym.

Leptosphaeria dolioloides (Auerswald) Karsten, op. cit,: 106 (1873).


(Fig. 16j.)
Pseudothecia scattered, immersed, sometimes becoming exposed,
subglobose with a flattened base, dark brown, up to 2 / 5 m m . across.
Asci numerous, clavate, short-stalked, up to 130x16/«, 8-spored;
ascospores biseriate, narrowly fusiform, 33-55/i long, with from 6 - 1 1
cross walls, the fourth cell from the top distinctly wider t h a n the rest,
a b o u t 5/1 wide, yellowish throughout.

On dead stems of Tanacetum and other Compositae. Common.

Leptosphaeria agnita (Desmazieres) Cesati & de Notaris, op. cit.:236


(1863). (Fig. 24A.)
Pseudothecia gregarious, developed beneath the host epidermis,
pear-shaped, up to 2 / 5 m m . wide and y2 m m . tall, with thick, smooth,
black wall. Asci subcylindric, numerous, 100-120x8-10/«, ascospores
biseriate, subcylindric, 3 0 - 3 7 x 3 . 5 - 4 . 5 / / , 6-septate, the third cell from
the top slightly enlarged, pale yellowish-brown.
The typical host is Eupatorium cannabinum but it occurs on many other
hosts, not confined to Compositae.

Leptosphaeria ogilvicnsis (Berkeley & Broome) Cesati & de Notaris,


op. cit.:235 (1863). (Fig. 24B.) "
Pseudothecia scattered or gregarious, developed beneath the host
epidermis, subglobose, about % m m . diameter, black, wall smooth,

"391
only 3 cells thick. Asci cylindric-clavate, 90-100x9-10/«. 8-spored;
ascospores biseriate, fusiform, 30-38x3.5-4.5/«, 5-septate, constricted
at the median septum, third cell from the top slightly enlarged,
yellowish-brown.

Typically on Senecio jacobaea, also on other Compositae, Valeriana & c.

Leptosphaeria maculans (Desmazieres) Cesati & de Notaris, op. cit.: 253


(1863). (Fig. 24c.)
Pseudothecia gregarious, subepidermal, hemispherical, black,
smooth, thick-walled, up to % m m . diameter. Asci numerous, sub-
cylindric, 100-150x12-15/«, 8-spored; ascospores irregularly arranged,
fusoid, 35-60x4.5-6/«, without an enlarged cell, yellowish-brown.
On dead stems of Cruciferae, especially Alliaria officinalis and Brassica
spp.

The conidial state is Phoma lingam (Tode ex Fries) Desmazieres,


commonly associated with a destructive rot in swedes see Muller,
P h y t o p a t h . Zeitschr. 29:291 (1957).
L. alliariae Auerswald is a synonym.

Leptosphaeria galiorum (Roberge) Cesati & de Notaris, op. cit.:235


(1863). (Fig. 24D.)
Pseudothecia scattered, subepidermal, pear-shaped, black, clothed
with sparse hyphae, about % m m . diameter. Asci numerous, cylindric-
clavate, 8-spored, 1 0 0 - 1 2 5 x 1 2 - 1 5 / / , ascospores biseriate, fusoid,
3 2 - 4 5 x 6 - 9 / / , 3-septate, light yellowish-brown.
On Galium aparine. Common.

L. aparines (Fuckel) Saccardo is a synonym.

Leptosphaeria macrospora (Fuckel) von Thuemen, Mycoth. univ. No.


1359 (1879). (Fig. 24E.)
Pseudothecia gregarious, subepidermal, more or less flattened, about
y 3 m m . wide, with a short neck, smooth, black, thick-walled. Asci
numerous, subcylindric, about 100-120x12/«, 8-spored; ascospores
biseriate, fusiform, pointed at the ends, 3 0 - 4 0 x 4 . 5 - 6 / / , 3-septate,
second cell from t h e top m a r k e d l y enlarged, pale yellowish-brown.
On Cirsium and other Compositae, Rumex & c.
L. nitschkei Rehm is a synonym. The conidial state is Rhabdospora
bernardiana Saccardo, see Sydowia 7:327.

"392
Leptosphaeria planiuscula (Riess) Cesati & de Notaris, op. cit.:236
(1836). (Fig. 24F.)
Pseudothecia gregarious, subepidermal, hemispherical, with a short
neck, y 3 m m . diameter, blackish, thick-walled. Asci numerous,
cylindric-clavate, 100-120x13-15//, 4-spored; ascospores biseriate,
fusiform-cylindric with rounded ends, 3 5 - 5 6 x 6 - 8 / / , 5-septate, con-
stricted at the septa, especially the median one, light brown.
On stems of Solidago virgaurea.

Leptosphaeria praetermissa (Karsten) Saccardo, Sylloge fungorum 2:


26 (1883). (Fig. 24G.)
Pseudothecia gregarious, immersed in the substrate, pear-shaped,
y 2 m m . diameter, thick-walled. Asci numerous, subcylindric, 8-spored;
ascospores uniseriate, ellipsoid, 1 8 - 2 4 x 6 - 8 / / , 3-septate, olivaceous.
On canes of Rubus idaeus. The British record needs confirmation.

Leptosphaeria cesatiana (Montagne) Holm, op. cit.:39 (1959).


Pseudothecia embedded in the host cortex, beneath a blackened
portion of epidermis, pear-shaped with a well developed neck, y 3 - %
m m . diameter, thick-walled. Asci numerous, cylindric-clavate, about
150x10//, 4-spored; ascospores parallel, cylindrical, 110-120x3.5-4//,
15-septate, yellowish, constricted at the median septum. (Fig. 24n.)

On dead stems, typically on Echium vulgare but also on Brassica, Hyperi-


cum and Lycopus.

Ophiobolus nigrificans Saccardo is a synonym.


Holm (1959) refers a couple of dozen other european species to
Leptosphaeria sensu stricto, most of which are to be sought on common
British plants.

Leptosphaeria modesta (Desmazieres) Auerswald, General-Doubl.


Verzeichn. Leipz. Tausch-Ver. :4 (1866). (Fig. 24i.)
Pseudothecia scattered, subepidermal, pear-shaped, up to % m m .
diameter, blackish hairy. Asci numerous, 8 0 - 1 0 0 x 1 0 - 1 2 / / , cylindric-
clavate, 8-spored; ascospores irregularly arranged, subcylindric,
2 5 - 4 0 x 4 . 5 - 5 / / , mostly 4-septate, occasionally 6-septate, pale olive,
ends bearing a small rounded hyaline appendage, second cell from the
top shorter and broader t h a n the rest.

Typically on Scabiosa columbaria, also on Aconitum, Bupleurum, Phy-


teuma, Sanguisorba, Scrophularia, Seseli, Tofieldia and many other plants.

"393
Nodulosphaeria modesta (Desmazieres) Munk apud Holm, op. cit.:
80 (1959).

Leptosphaeria derasa (Berkeley & Broome) Auerswald, op. cit.: 4 (1866).


Pseudothecia gregarious, embedded in the host cortex, pear-shaped,
mm
black, hairy, - diameter, thick-walled. Asci numerous,
cylindric-clavate, 9 0 - 1 2 0 x 1 1 - 1 4 / / , 8-spored; ascospores irregularly
arranged, subcylindric, 3 5 - 5 2 x 4 - 4 . 5 / / , mostly 8-septate, pale olive,
f o u r t h cell from the top slightly enlarged, with a short curved append-
age at each end. (Fig. 24J.)

On Senecio jacobaea, occasionally on Picris.

Nodulosphaeria derasa (Berkeley & Broome) Holm, op. cit,: 89


(1959).

Leptosphaeria dolioloides common on various genera of Composi-


tae, is very similar to the above b u t with more pointed ascospores,
lacking the appendages and 8-11-septate.

Leptosphaeria silenes-acaulis de Notaris in Comm. Soc. crit. Ital. 2:485


(1867). (Fig. 24K.)
Pseudothecia scattered, immersed in the host, conical, 50-70// dia-
meter, smooth, black, thin-walled. Asci few, ellipsoidal, 45-60x 12-16//,
8-spored; ascospores irregularly arranged, cylindric-fusoid, 30-38x
5 - 6 / / , 3-septate, yellowish-brown.
In leaves and capsules of Silene acaulis.

Phaeosphaeria silenes-acaulis (de Notaris) Holm, op. cit.: 128 (1959).

Leptosphaeria sowerbyi (Fuckel) Saccardo, Sylloge f u n g o r u m 2 : 7 8


(1883). (Fig. 24L.)
Pseudothecia gregarious, immersed in the host, dark brown, globose
without a neck, 75-100// diameter, thin-walled. Asci few, broadly
cylindrical, 6 0 - 7 0 x 1 8 - 2 0 / / , 8-spored; ascospores fasciculate, sub-
cylindric, 42-50x5-6/1, 6-septate, pale yellow, third cell from the top
very slightly enlarged.
On dead Scirpus lacustris.

Phaeosphaeria sowerbyi (Fuckel) Holm, op. cit.: 130 (1959).

"394
Leptosphaeria juncina (Auerswald) Saccardo, Sylloge fungorum 2:66
(1883). (Fig. 24M.)
Pseudothecia in swarms embedded in the substrate, subglobose,
75-100/t diameter, without a neck, wall smooth, thin, brown. Asci
few, subcylindric, 6 0 - 7 0 x 1 4 - 1 5 / / , 8-spored; ascospores irregularly
arranged, cylindric-fusiform, often slightly curved, 3 0 - 4 0 x 4 . 5 - 6 / / ,
3-septate, yellowish, end cells shorter t h a n the others.
In dead culms of Juncus spp.

Phaeosphaeria juncina (Auerswald) Holm, op. cit.: 127 (1959).

Leptosphaeria juncicola Rehm apud W i n t e r in Hedwigia 19:167


(1880). (Fig. 24N.)
Pseudothecia gregarious, immersed in the host, subglobose, 100-
200/< diameter, brown. Asci few, cylindrical, 60-80x 12-15//, 8-spored;
ascospores irregularly disposed, cvlindric-fusoid, often curved, 33-45 x
4-4.5//, 3-septate, yellowish, second cell often slightly enlarged.
In Juncus trifidus and leaves of Carex spp.

Phaeosphaeria juncicola (Rehm) Holm, op. cit.: 129 (1959).

Leptosphaeria typharum (Desmazieres) Karsten, Mycologia fennica


2:100 (1873). (Fig. 24o.)
Pseudothecia gregarious, immersed in the substrate, subglobose,
smooth, 150// diameter, brown, thin-walled. Asci few, cylindric-
clavate, 70-100x20-25//, 8-spored; ascospores irregularly arranged,
broadly ellipsoid, 2 0 - 3 3 x 9 - 1 3 / / , 3-septate, brownish, broadest at the
second cell from the top.
In dead leaves of Typha.

Phaeosphaeria typharum (Desmazieres) Holm, op. cit.: 126 (1959).


There is also on Typha a L. typhae (Karsten) Saccardo with ascospores
1 6 - 2 3 x 4 - 5 . 5 / / , which Holm regards as no more t h a n a race of L. eu-
stoma.

Leptosphaeria epicalamia (Riess) Cesati & de Notaris, op. cit.:236


(1863). (Fig. 24P.)
Pseudothecia scattered, immersed in the substrate, pear-shaped,
y 4 m m . diameter, brown. Asci numerous, cylindric-clavate, 7 0 x 1 0 / / ,

"395
8-spored; ascospores biseriate, cylindric-fusoid, straight, 2 1 - 2 5 x 5 - 6 / / ,
5-septate, second cell from the top markedly enlarged.
On Luzula.
Phaeosphaeria epicalamia (Riess) Holm, op. cit.: 114 (1959).

Leptosphaeria littoralis Saccardo in Michelia 1 : 3 8 (1877). (Fig. 24Q.)


Pseudothecia gregarious, immersed with erumpent ostioles, pear-
shaped, about % m m . diameter, smooth, black, moderately thick-
walled. Asci few, cylindric-clavate, 150-190x30-35//, 8-spored; asco-
spores biseriate, narrowly ellipsoid, 4 2 - 4 6 x 1 3 - 1 5 / / , 6-7-septate, light
brown, broadest at the middle.
On Ammophila arenaria and other sanddune grasses.

Phaeosphaeria littoralis (Saccardo) Holm, op. cit.: 121 (1959). The


older Metasphaeria sabuletorum (Berkeley & Broome) was based in
p a r t on this species b u t the name is rejected b y Holm as having
originated in confusion of several fungi.

Leptosphaeria nardi (Fries) Cesati & de Notaris, op. cit.:236 (1863).


Pseudothecia scattered, immersed in the host, subglobose, up to
1
j 6 m m . diameter, smooth, black, wall moderately thick. Asci numer-
ous, 7 0 - 8 0 x 1 0 - 1 2 / / , 8-spored; ascospores biseriate, subcylindric,
23-25 x4—4.5//, 5-septate, the f o u r t h cell from the top enlarged and
longer t h a n the third, hyaline at first, finally pale yellow. (Fig. 24R.)
On Nardus stricta.

Phaeosphaeria nardi (Fries) Holm, op. cit.: 124 (1959).


L. fuckelii Niessl in Oesterr. Bot. Zeit. 32:357 (1882), common on
Calamogrostis, Deschampsia and especially on Phalaris and Phragmites,
is very similar b u t has the f o u r t h cell of the ascospore distinctly
shorter. Phaeosphaeria fuckelii (Niessl) Holm, op. cit. 123 (1959). (Fig.
24v.)

Leptosphaeria nigrans (Roberge) Cesati &de Notaris, op. cit.: 235 (1863).
Pseudothecia scattered, immersed in the substrate, more or less
pear-shaped with a short neck, dark brown, about % m m - diameter,
smooth to downy. Asci numerous, cylindric clavate, 7 5 - 9 0 x 6 - 8 / / ,
8 spored; ascospores biseriate, subcylindric, 18-23x 3-4/t, 5-septate,
second cell from the top distinctly enlarged, yellowish. (Fig. 24s.)
Very common on dead culms and leaves of many genera of grasses.

"396
Phaeosphaeria nigrans (Roberge) Holm, op. cit.: 112 (1959). The
species is best known as L. culmicola (Fries) Auerswald but the identity
of Sphaeria culmicola Fries is now uncertain.

Leptosphaeria herpotrichoides de Notaris, Sferiacei italici:80 (1863).


Pseudothecia scattered to clustered in rows, immersed in the host,
pear-shaped or flattened with a short neck, or laterally compressed,
1
U~1//3 m m . diameter, usually clothed with loose hyphae, blackish, wall
moderately thick. Asci numerous, cylindric-clavate, 9 0 - 1 2 0 x 9 - 1 2 / / ,
8-spored; ascospores biseriate, subfusiform, 2 2 - 4 2 x 4 - 6 / / , 6-10-sep-
t a t e , mostly 7-septate, usually the t h i r d cell from the top enlarged,
yellowish. (Fig. 24T.)

Common on a great many genera of grasses, occasionally on Carex and


Luzula.

Phaeosphaeria herpotrichoides (de Notaris) Holm, op. cit.: 115 (1959).


The species has commonly been called L. culmifraga (Fries) Cesati &
de Notaris but according to Holm it is not Sphaeria culmifraga Fries.

Leptosphaeria graminis (Fuckel) Saccardo, Sylloge fungorum 2 : 7 6


(1883). (Fig. 24u.)
Pseudothecia usually in groups, immersed in the host, subglobose
or laterally compressed, sometimes with a neck up to 100// long,
% m m . diameter, smooth or covered with loose hyphae, blackish. Asci
numerous, 1 0 0 - 1 2 0 x 9 - 1 2 / / , cylindric-clavate, 8-spored; ascospores
biseriate, subcylindric, 3 2 - 4 5 x 4 - 5 / / , 10-12-septate, usually the third
cell from the top enlarged, yellowish-olivaceous.
On culms and leaf-sheaths especially of Phragmites communis, also on
Elymus, Phalaris and Secale.

Phaeosphaeria graminis (Fuckel) Holm, op. cit.: 118 (1959).

Leptosphaeria eustoma (Fuckel) Saccardo, Fungi italici t . : 497 (1879).


Pseudothecia scattered, immersed in the substrate, subglobose,
100-200/« diameter, dark brown, wall smooth, of medium thickness.
Asci numerous, cylindric-clavate, 7 5 - 1 0 5 x 1 5 - 2 0 / / , 8-spored; asco-
spores biseriate, ellipsoid, 1 8 - 3 3 x 5 - 1 0 . 5 / / , 3-septate, the second cell
from the top slightly enlarged, yellowish. (Fig. 24w.)
Very common on leaves and culms of a great many genera of grasses and
also on other Monocotyledons, Carex spp., Hydrocharis morsus-ranae, Iris,
Juncus, Luzula, Scirpus, Sparganium and Typha.

"397
Phaeosphaeria eustoma (Fuckel) Holm, op. cit. 109 (1959).
L. microscopica Karsten and L. ciilmorum Auerswald are synonyms.
The race on Typha tends to have n a r r o w ascospores and has been
distinguished as L. typhae (Karsten) Saccardo b u t Holm thinks it not
distinct; nor perhaps is L. parvula Niessl, on Iris with ascospores
3-septate, 1 8 - 2 1 x 4 / / .

Leptosphaeria tritici (Garovaglio) Passerini apud de Thuemen Myco-


theca universalis No. 859 (1877). (Fig. 24x.)

Pseudothecia scattered, immersed in the substrate, subglobose,


black 7 5 m m . wide, smooth. Asci numerous, cylindric-clavate, 50x
15-16//, 8-spored; ascospores biseriate, straight or slightly curved,
1 8 - 2 5 x 4 - 5 . 5 / / , 3-septate, the second cell from the top slightly en-
larged, pale yellowish-brown.

On dead leaves of Triticum. Holm thinks this scarcely morphologically


distinct from L. eustoma and the same applies to L. avenaria Weber on
Avena saliva.

Leptosphaeria eustomoides Saccardo in Nuovo Giorn. bit. Ital. 7:319


(1875) is a synonym according to Miiller b u t it is difficult to say how
m a n y of these graminicolous species can eventually be distinguished.
The t y p e was on Andropogon isehoemi and the name has been adopted
for a Leptosphaeria on Dactylis, see Webster in Trans. Brit, mycol. Soc,
40:519 (1957).

Leptosphaeria nodorum Miiller in P h y t o p a t h . Zeitschr. 19:409 (1952).


Pseudothecia immersed in the culm, gregarious, often connected by
a stromatic tissue, subglobose, up to % m m . diameter, wall thin, dark
brown. Asci numerous, cylindric-clavate 6 0 - 7 0 x 9 / / , 8-spored; asco-
spores fusoid, 2 0 - 2 6 x 4 / / , 3-septate, broadest at the second cell from
the top, yellowish. (Fig. 24Y.)

On Triticum vulgare. The conidial state is Septoria nodorum. Berkeley.

Leptosphaeria luctuosa Niessl apud Saccardo in Nuovo Giorn. bot.


Ital. 7:321 (1875). (Fig. 24z.)
This species with 5-septate ascospores 28-36x5.5-6.5//, has been
reported on Dactylis, see Webster, op. cit.:511 (1957).

"398
Leptosphaeria pontilormis (Fuckel) Saccardo, Sylloge F u n g o r u m 2:78
(1883). (Fig. 24cc.)
This species with ascospores 32-50x4-4.5/« up to 12-septate, with
the third or fourth cell enlarged, has been reported on Agropyron by
Webster, op. cit.:517 (1957).

Leptosphaeria arundinacea (Sowerby ex Fries) Saccardo in Nuovo


Giorn. bot. Ital. 7:320 (1875). (Fig. 24AA.)
Pseudothecia gregarious, even confluent, erumpent from dead
culms, more or less globose, smooth, black, % m m - diameter, with an
apical papilla. Asci numerous, cylindric-clavate, 7 5 - 9 0 x 1 0 - 1 3 / / ,
8-spored; ascospores biseriate, cylindric-fusoid, 21-30x4-5/«, 3-sep-
t a t e , the second cell from the top slightly enlarged, finally brownish.
On Phragmites communis. The ascospores remain for a long time colour-
less and in this state were described as a separate species, L. godeni (Des-
mazieres) Auerswald.

Leptosphaeria michotii (Westendorp) Saccardo, Fungi Italici t a b . : 2 7 9


(1878). (Fig. 24BB.)
Pseudothecia gregarious, immersed in the substrate, globose, a b o u t
1
/ 5 m m . diameter, with t h i n dark-brown walls. Asci cylindric-clavate,
55-70x10-15/«, 8-spored; ascospores biseriate, cylindrical with round-
ed ends, 14-22x3.5-6/«, 2-septate, constricted at the lowest s e p t u m ,
brown. The conidial state is Coniothyrium scirpi Trail.
On Calamagroslis, Dactylis, Juncus, Scirpus and Typha.
Paraphaeosphaeria michotii (Westendorp) S h o e m a k e r & E r i k s s o n in
Canadian Journal of B o t a n y 45:1605 (1967).

Leptosphaeria rusci (Wallroth) Saccardo, Syllogte F u n g o r u m 2:74


(1883). (Fig. 25A.)
Pseudothecia in clusters, embedded in dead cladodes, flattened-
globose, up to 150/t diameter, with conical ostiolar papilla, smooth,
black. Asci numerous, cylindric-clavate, 60-80x10/«, 8-spored; asco-
spores biseriate, subcylindric, 15-23x3.5-4.5/«, 4-5-septate, broadest
at t h e second or third cell, light brown.
On Ruscus aculeatus. The conidial state is Haplosporella ruscigena Bubak.
Paraphaeosphaeria rusci (Wallroth) Eriksson, Ark. F. Bot. 6:406.
Leptosphaeria niessleana Rabenhorst apud Niessl in Verhandl. Nat.
Ver. Briinn 14:179 (1872). (Fig. 25B.)
Pseudothecia scattered, immersed in the host, % m m - broad, flask-
shaped with a long neck, smooth, black. Asci numerous, cylindric-

"399
clavate, 8 0 - 9 0 x 9 - 1 0 / / , 8-spored; ascospores biseriate, subcylindric,
2 2 - 3 2 x 3 . 5 - 5 / / , 4-septate, the apical cell long and slender, subapical
cell slightly enlarged, yellowish.

On Lathyrus spp.

Entodesmium niessleanum (Rabenhorst) Holm, op. cit.: 133 (1959).

Leptosphaeria vectis (Berkeley & Broome) Cesati & de Notaris, op. cit.:
236 (1883). (Fig. 25c.)
Pseudothecia gregarious, immersed in dead leaves, globose, black.
Asci cylindric-clavate, up to 100x12//, 8-spored; ascospores biseriate,
subcylindric, 20-22 x 6-7/z, eventually light-brown and 4-septate.

On Iris foetidissima.

Leptosphaeria scitula Sydow in Ann. mycol. 36:321 (1938). (Fig. 25n.)


Pseudothecia scattered, subepidermal, subglobose, 1 j 5 m m . dia-
meter, thin-walled and collapsing when dry, black. Asci numerous,
clavate, 5 5 - 6 5 x 7 - 8 / / , 8-spored; ascospores irregularly arranged, nar-
rowly fusoid, 2 2 - 3 0 x 3 - 3 . 5 / / , 3-septate, light yellowish-brown.
On Galium aparine.

Melanomma scitulum (Sydow) Holm, op. cit.:66 (1959). Holm notes


t h a t the substrate is occasionally tinged red, as in other species he
transferred to Melanomma.

Leptosphaeria haematites (Roberge) Niessl apud Rabenhorst Fungi


Europaei No. 2761. (Fig. 25E.)
Pseudothecia gregarious in reddened patches of the substrate, sub-
epidermal, subglobose, % m m . diameter. Asci numerous, cylindric-
clavate, 55-70x8-9 ( M, 8-spored; ascospores biseriate, fusiform, 21-27x
4-5//, 3-septate, the second cell slightly enlarged, light brown.

On dead stems of Clematis vitalba and Eupatorium.

Leptosphaeria purpurea Rehm, Ascomycetes Lojkani lecti:54 (1882).


Pseudothecia gregarious, subepidermal beneath reddened patches
of the host, subglobose, 1 / i m m . diameter, flattened or collapsed when

"400
dry, black, covered with m a t t e d hyphae. Asci numerous, cylindric-
clavate, 7 0 - 9 0 x 7 - 9 / / , 8-spored; ascospores biseriate, cylindric-fusoid,
2 4 - 3 3 x 4 . 5 - 5 / / , 3-septate, the second cell from the top markedly en-
larged, yellowish-brown. (Fig. 25r.)

On dead stems of Betonica, Chamaenerion, and especially of Compositae.

Melanomma purpureum (Rehm) Holm, op. cit.: 64 (1959).

Leptosphaeria coniothyrium Saccardo in Nuovo Giorn. bot. Ital. 7:317


(1875). (Fig.25o.)
Pseudothecia gregarious, immersed in t h e host cortex, subglobose,
y4~y 3 m m . diameter, black, covered b y a small blackish clypeus. Asci
numerous, cylindric-clavate, 7 0 x 6 / / , 8-spored; ascospores biseriate,
ellipsoidal, 12-15x3.5-4.5//, 3-septate, pale olive-brown.

In canes of Rubus spp.

Melanomma coniothyrium (Saccardo) Holm, op. cit.:57 (1959).

Leptosphaeria albopunctata (Westendorp) Saccardo, Sylloge Fungo-


rum 2:72 (1883). (Fig. 25J.)
Pseudothecia scattered, subglobose with an erumpent papillate
ostiole, smooth, black, % m m . diameter. Asci few, cylindric-clavate,
135-190x11-30//, 8-spored; ascospores biseriate, elliptic-fusoid, 30-55
x 10-15//, 5-7-septate, slightly constricted at the septa, yellowish-
brown.

On dead culms of Spartina townsendii and on driftwood.

Leptosphaeria macrosporidium Gareth Jones in Trans. Brit., mycol.


Soc. 45:103 (1962). (Fig. 25K.)
Pseudothecia scattered, erumpent from culms or superficial on
mm
wood' subglobose, - diameter with an apical papilla, smooth,
black. Asci cylindric-clavate, 140-220x20-28//, 8-spored; ascospores
elliptic-cylindric, 48-72x 10-14//, 3-septate, constricted at the septa,
hyaline.
Apinis & Chesters think this to be synonym of L. marina Ellis and
E v e r h a r t in J . mycol. 1 : 4 3 (1885).
On Spartina townsendii and driftwood.

"401
Leptosphaeria maritima (Cooke & Plowright) Saccardo, Sylloge F u n -
gorum 2 : 7 3 (1883). (Fig. 25i.)
Pseudothecia scattered, erumpent from culms, globose with a
papillate ostiole, % m m . diameter, smooth, black. Asci few, cylindric-
clavate, 120-140x14-18//, 8-spored; ascospores biseriate, ellipsoid,
3 2 - 4 0 x 8 - 1 2 / / , 3-5-septate, yellowish-brown.
On Spartina townsendii.

Leptosphaeria orae-maris Linder in Farlowia 1 : 4 1 3 (1944). (Fig. 25L.)


Pseudothecia scattered, erumpent from culms, % m m . diameter,
subglobose with an apical papilla, black, smooth. Asci few, cylindric-
clavate, 1 0 0 - 1 4 0 x 8 - 1 2 / / , 8-spored; ascospores biseriate, ellipsoidal,
1 8 - 2 5 x 5 - 8 / / , one t h e n three-septate, constricted at the middle, brown.
On Spartina townsendii and driftwood.

Leptosphaeria pelagica Gareth Jones in Trans. Brit, mycol. Soc. 45:105


(1962). (Fig. 25M.)
Pseudothecia scattered, e r u m p e n t or superficial, subglobose with an
apical papilla, black, % m m - diameter. Asci few, cylindric-clavate,
1 0 0 - 1 5 0 x 1 6 - 2 3 / / , 8-spored; ascospores uniseriate, ellipsoidal, 28-40x
8 - 1 2 //, 3-septate, constricted at the septa, hyaline.
On Spartina townsendii and driftwood.

The following species has been transferred to Trichometasphaeria b y


Holm. It lacks apical bristles, however, and has not been recombined
in Keissleriella, which is regarded as an older name for Trichometa-
sphaeria.

Leptosphaeria scirpina Winter in Hedwigia 11:146 (1872). (Fig. 25H.)


Pseudothecia scattered, subepidermal, globose, 1 / 6 m m . diameter,
smooth. Asci numerous, cylindric-clavate, 100-125x 15-29//, 8-spored;
ascospores biseriate, subfusoid, 3 0 - 3 6 x 6 / / , hyaline, 5-septate.
Typically on Scirpus lacustris but also on Carex and Molinia according
to Holm.

REBENTISCHIA Karsten
Rebentischia Karsten, Mycologia Fennica 2 : 1 4 (1873).
Differs from Leptosphaeria in the ascospores having a hyaline basal
appendage. There are two British species:

"402
Rebentischia typhae H. F a b r e in Ann. Sci. Nat. Bot. Ser. VI, 9 : 8 8
(1878). (PL X X X I X i . )
Pseudothecia scattered, immersed b u t forming small pustules, sub-
globose, black, about % m m - across, with erumpent ostioles. Asci
cylindric-clavate, stalked, up to 125x20//, 8-spored; ascospores bi-
seriate, slightly clavate, 3 0 - 4 6 x 8 - 1 0 / / , 5-septate, somewhat constrict-
ed at the septa, brown, with a pointed, hyaline, basal appendage 4 - 6 / /
long.
On dead culms of Phragmites in the north. Apparently uncommon.

Rebentischia unicaudata (Berkeley & Broome) Saccardo in A t t i Soc.


Veneto-Trentini Scienze Naturali 4 : 8 8 (1875), occurs on dead stems of
Clematis vitalba.

PASSERINIELLA Berlese

Passeriniella Berlese Icones F u n g o r u m 1 Fasc. 2 : 5 1 (1902).


Pseudothecia in stems, ascospores transversely septate, with brown
central cells and hyaline terminal ones.

Passeriniella discors (Saccardo&Ellis) Apinis&Chesters in Trans. Brit,


mycol. Soc. 47:432 (1964). (Fig. 25N.)
Pseudothecia erumpent, black, smooth, subglobose, about y 3 m m .
diameter, with a broad apical papilla. Asci cylindric-clavate, 150-180x
15-16/«, 8-spored; ascospores ellipsoidal, 2 4 - 3 3 x 8 - 1 3 / t , 3-septate,
constricted at the septa, the central cells brown, the end ones colour-
less.

On stems of Spartina townsendii and driftwood. Common.

Didymosphaeria spartinae Grove is a synonym.

CHITONOSPORA Bommer, Roussel & Saccardo

Chitonospora Bommer, Roussel & Saccardo, Sylloge Fungorum 9:797


(1891).
Differs from Leptosphaeria in the distinctly 2-layered wall of the
ascospores and according to Eriksson the asci are unitunicate.

"403
Chitonospora ammophilae Bommer, Roussel & Saccardo, op. cit.:797
(1891). (PI. X X X I X H . )
Pseudothecia scattered immersed, globose with erumpent ostioles,
about i/ 3 mm. across, black. Asci narrowly clavate, thick-walled, sub-
sessile up to 120x20//, 8-spored; ascospores more or less biseriate,
elliptic-fusiform, 23-29x 10-14/.«, 3-septate, brown, t h e wall very thick
and distinctly 2-layered.
In dead leaves of Ammophila arenaria; apparently not uncommon but
only recently recognised in the British Isles.

TREMATOSPHAERIA Fuckel

Trematosphaeria Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:


161 (1870).
Pseudothecia partially immersed in wood, with thick, opaque, black
walls, asci thick-walled, 8-spored, ascospores more or less biseriate,
becoming coloured and more t h a n 1-septate.
Holm, seeking to define the genus by more fundamental characters,
stressed especially the ascospores remaining paler at their tips and the
anatomy of the pseudothecial wall. The latter is described as strong, com-
posed of numerous layers of cells, usually thin-walled, very variable in
shape and size, but heavily pigmented in an irregular manner so that in
section the wall appears remarkably variegated. On this basis he added to
the lignicolous species a number on herbaceous stems commonly referred
to Leptosphaeria, notably L. circinans and L. heterospora.

Trematosphaeria pertusa (Persoon ex Fries) Fuckel, op. cit.: 162 (1870).


(Fig. 16D.)
Pseudothecia scattered, subglobose or obtusely conical with rounded
bases, their lower portions sunk in decorticated wood, about % m m .
across, black, smooth. Asci narrowly clavate, short-stalked, up to
140x20//, 8-spored; ascospores mostly biseriate, fusiform, 20-27x
5 - 7 / / , at first hyaline, 1-septate and markedly constricted at the sep-
t u m , finally brown with two additional septa.
On decorticated wood of Fagus, Quercus and other deciduous trees. Not
uncommon.

Trematosphaeria circinans (Saccardo) W i n t e r in Rabenhorst's K r y p t .


Flora 1 (2): 277 (1885). (Fig. 29o.)
Pseudothecia gregarious, base sunk in the substrate, pearshaped,
1/3 m m . diameter, smooth, black. Asci numerous, cylindric-clavate,

"404
112-130x20//, 8-spored; ascospores biseriate, broadly fusoid, 26-35x
10-11//, 3-septate, median cells dark brown, terminal cells paler, wall
finally verrucose.
On rootstock of Medicago sativa, also on Helianthemum and twigs of
Ligustrum according to Holm.

Trematosphaeria heterospora (de Notaris) Winter, op. cit.:277 (1885).


Pseudothecia scattered, erumpent, pearshaped, y 3 m m . diameter,
smooth, black. Asci numerous, cylindric-clavate, 110-120x20-23//,
8-spored; ascospores biseriate, broadly fusoid, 3 6 - 4 5 x 1 0 - 1 2 / / , 3-sep-
t a t e , long remaining hyaline b u t eventually dark brown. (Fig. 29P.)
On rhizomes of Iris germanica, also on Planlago maritima according to
Holm.

ASTROSPHAERIELLA Sydow
iisuosphacriella Sydow in Annales mycologici 11:260 (1913).
Pseudothecia almost superficial, carbonaceous, smooth, on wood or
bark, like those of Trematosphaeria b u t with 1-septate brown asco-
spores.

Astrosphaeriella applanata (Fries) Scheinpflug in Ber. Schweiz. Bot.


Ges. 68:368 (1958). (Fig. 20K.)
Pseudothecia scattered, almost superficial, subconical, each with a
very broad base and prominent apical papilla, smooth, black, % to
% m m . across. Asci thick-walled, 8 5 - 1 1 0 x 1 8 - 2 2 / / , 4 or 8-spored;
ascospores irregularly biseriate, ellipsoidal b u t more pointed above,
1-septate, constricted at the septum and the upper cell broader t h a n
the lower, 2 4 - 2 9 x 8 - 1 1 / / , brown.
On wood of Alnus, Carpinus and Quercus in winter and spring.

Kirschsteiniella applanata (Fries) P e t r a k in Ann. mycol. 21:331 (1923)


is a synonym.

CARYOSPORA de Notaris

Caryospora de Notaris Mycromyc. Ital. Dec. 9:7 (1856).


Pseudothecia like those of Trematosphaeria b u t ascospores very
large, dark brown and markedly apiculate.

405
Caryospora callicarpa (Currey) Nitschke, apud Fuckel in J a h r b . Nass.
Yer. N a t u r k . 23/24:163 (1870). (Fig. 20F.)
Pseudothecia erumpent, semi-immersed in decorticated wood, each
about y 2 to % m m . diameter, smooth, black, with low ostiolar papilla;
asci about 250x65//, 8-spored; ascospores irregularly biseriate, with
2 large, dark brown, central cells and one or two small paler cells at
each end, 6 0 - 7 5 x 2 8 - 3 2 / / .
On Quercus in spring, apparently very rare.

MELOMASTIA Nitschke

Melomastia Nitschke apud Fuckel in J a h r b . Nass. Vereins f. N a t u r -


kunde 25-26:306 (1871).
Pseudothecia immersed, at least at first, black, asci 8-spored, asco-
spores uniseriate, 3-celled, hyaline. There is one British species:

Melomastia mastoidea (Fries) Schroeter, K r y p t o g a m e n Flora von


Schlesien 3 (2): 320 (1894). (Fig. 23D.)
Pseudothecia scattered or gregarious, immersed, raising b a r k into
small pustules pierced b y the ostioles, later often exposed, globose
with an apical papilla, from y 2 to % m m . across, black, with a thick
opaque wall, smooth. Asci narrowly cylindrical, short-stalked, up to
1 5 0 x 8 / / , 8-spored; ascospores uniseriate, oblong with rounded ends,
1 4 - 2 0 x 4 . 5 - 7 / / , hyaline, with two septa and slightly constricted at the
septa.

On dead twigs and branches of many shrubs, figured on Syringa, also


reported on Fraxinus and Populus. Not uncommon.

MASSARINA Saccardo

Massarina Saccardo, Sylloge F u n g o r u m 2:153 (1883).


Pseudothecia gregarious, immersed in bark, each with a feebly
developed clypeus, asci large, thick-walled, ascospores large, trans-
versely septate, hyaline, with gelatinous outer coats.

Massarina eburnea (Tulasne) Saccardo, op. cit.: 153 (1883). (Fig. 23A.)
Pseudothecia gregarious, immersed in bark, subglobose, flattened
above and there covered by a small p a t c h of black stromatic tissue,

"406
forming a small clypeus within the outer layer of b a r k , about % m m .
across, the wall towards the base thin and almost colourless. Asci
cylindric-clavate, short-stalked, thick-walled, up to 170x21/«, 8-
spored; ascospores more or less biseriate, narrowly elliptical, 30-37 x
8 - 9 / / , 3-septate, strongly constricted, hyaline, with a gelatinous outer
coat.

On thin dead twigs of Fagus, in spring.

PTERIDIOSPORA Penzig & Saccardo

Pteridiospora Penzig & Saccardo in Malpighia 11:393 (1897).


Pseudothecia gregarious in bark, subspherical, with thick black wall.
Ascospores subclavate, large, dark brown, septate in the lower third,
with a gelatinous coat. The genus is very near Massaria, from which it
differs in the 1-septate ascospores.

Pteridiospora scoriadea (Fries) comb, no v. (Fig. 30D.)


Sphaeria scoriadea Fries, Elenchus F u n g o r u m 2:87 (1828).
Pseudothecia gregarious, erumpent from extensive patches of b a r k ,
about 1 mm. diameter, smooth, black, subglobose or mutually com-
pressed, with an ostiole b u t no beak. Asci clavate, 8-spored, soon dif-
fluent; ascospores 5 3 - 6 5 x 1 7 - 2 4 / / , the upper cell much longer and
darker t h a n the lower.
In bark of Beiula twigs. Autumn. Uncommon.

Pteridiospora currcyi (Tulasne) Muller in Beitr. K r y p t . Flora


Schweiz 11 (2): 281 (1962) with ascospores 3 0 - 4 0 x 1 2 - 1 8 / / , occurs on
Tilia.

MASSARIA de Notaris

Massaria de Notaris in Giornale Botanico Italiano 1 : 3 3 3 (1844).


Pseudothecia large, immersed in bark, sometimes clustered and sur-
rounded b y a r u d i m e n t a r y stroma, asci large, thick-walled, ascospores
very large, brown, with more t h a n one cross septum and a thick
hyaline gelatinous coat.

"407
Massaria inquinans (Tode ex Fries) de Notaris, op. cit.:333 (1844).
(Fig. 2 3 F . )
Pseudothecia about 1 m m . across, solitary or in small clusters, im-
mersed and lifting t h e b a r k into pustules, each pseudothecium or
group of pseudothecia surrounded b y a zone of blackish stromatic
tissue, pseudothecia slightly flask-shaped with short ostiolar necks,
black, smooth. Asci cylindric-clavate, short-stalked, up to 3 5 0 x 6 0 / / ,
8-spored; ascospores biseriate, fusiform, 75-100x17-21/«, dark brown,
with a well marked median septum and ultimately two other septa,
containing large oil drops, enclosed in a thick coat which is also usually
constricted at the middle.

On dead branches of Acer pseudoplatanus; in spring.

Massaria pupula (Fries) Tulasne, Selecta F u n g o r u m Carpologia 2:225


(1863). (Fig. 23H.)
Pseudothecia gregarious, somewhat flattened, up to % mm. across,
immersed in bark with very small apical papillae which appear as
m i n u t e black erumpent dots. Asci cylindric-clavate, thick-walled,
stalked, up to 2 3 0 x 4 5 / / , 8-spored; ascospores biseriate, elliptic-fusi-
form, 4 0 - 5 5 x 1 0 - 1 8 / / , dark brown, 3-septate, appearing minutely
punctate, sheathed b y a thick, hyaline, gelatinous coat which is
strongly constricted at the level of t h e middle septum.

On dead branches of Acer pseudoplatanus. Also known as Stigmato-


massaria pupula (Fries) Munk in Dansk. Bot. Arkiv. 15 (2): 127 (1953).

Massaria foedans (Fries) Fries, S u m m a Veg. Scand. Sect. Post.:396


(1849), on Ulmus, has clavate brown ascospores 4 8 - 5 2 x 1 9 - 2 4 / / with
two septa towards the lower narrower end.

ASTEROMASSARIA von Hohnel

Asteromassaria von Hohnel in Sitzb. Akad. Wiss. Wien, Mathem.-Nat.


Kl. A b t . 1, 126:368 (1917).
Like Massaria b u t with the pseudothecia free and seated on a com-
mon stromatic base, erumpent through slits in bark.

Asteromassaria macrospora (Desmazieres) von Hohnel, op. cit. (Fig.


20i.)
Pseudothecia in small clusters, subglobose, about y 2 to % m m .
across, smooth, black, each with an apical minutely papillate ostiole.

"408
Asci thick-walled, very large, short-stalked, 8-spored; ascospores 1 - 2 -
seriate, ellipsoidal, for some time hyaline and 1-septate, finally brown
and 3-septate, 45-55x16-20/«. Associated with the pseudothecia there
are usually long curved brown conidia, 100-150x12-15//, with
numerous transverse septa.

Erumpent from branches of Fagus, in winter.

MELANOMMA Nitschke

Melanomma Nitschke apud Fuckel in J a h r b . Nass. Vereins f. Natur-


kunde 23-24:159 (1870).
Pseudothecia gregarious, superficial, at least at m a t u r i t y , black, asci
cylindrical, ascospores uniseriate or biseriate, brownith, with more
t h a n one septum.
Holm, seeking more natural characters to define genera than those based
on substrate and position of growth, has enlarged Melanomma to indued a
number of species on dead herbaceous stems traditionally referred to
Leptosphaeria, notably L. haematites, L. purpurea and L. scitula, see p. 400.

Melanomma pulvis-pyrius (Persoon ex Fries) Fuckel, op. cit.: 160


(1870). (Fig. 16c.)
Pseudothecia crowded, often seated on a thin black mycelial crust,
subglobose, up to y 2 m m . across, without a distinct apical papilla,
black, almost smooth, dull or shining. Asci narrowly cylindrical, short-
stalked, thick-walled at the tip, up to 110x9//, 8-spored; ascospores
uniseriate b u t usually somewhat overlapping, elliptic-fusiform,
straight or slightly curved, 12-20x4-6/«, 3-septate and slightly con-
stricted, especially at the middle septum, the second cell from the top
usually slightly wider t h a n the others, pale olive-brown.

On dead wood and bark, mostly of deciduous trees and shrubs. Very
common.
Melanomma fuscidulum Saccardo in Michelia 1:450 (November
1878) has shorter asci with biseriate ascospores 1 1 - 1 6 x 3 - 4 / / .

THYRIDARIA Saccardo

Thyridaria Saccardo in Grevillea 4 : 2 1 (September 1875).


Pseudothecia gregarious in an erumpent stroma, asci and ascospores
like those of Melanomma. There is one British species:

"409
Thyridaria rubro-notata (Berkeley & Broome) Saccardo, Sylloge F u n -
gorum 2 : 1 4 1 ( 1 8 8 3 ) . ( P I . X X X I X B . )
Stroma a compact cushion of interwoven brown hyphae, immersed
in bark and erumpent to expose the ostioles, up to about 3 m m . across,
pseudothecia globose, about %-Mt m m . across, with short stout necks
and well developed ostioles, surrounded by more or less conspicuous
reddish mealy hyphae on the surface of the stroma. Asci narrowly
cylindric-clavate, up to 120x10fi, 8-spored; ascospores uniseriate,
elliptical or slightly clavate, 14-19x5-6/«, 3-septate and slightly
constricted at the septa, dark brown. The stroma also contains small
pycnidia among the pseudothecia, with elliptical yellowish to oliva-
ceous conidia 5 - 8 x 2 . 5 - 4 / « , figured above the ascus on the plate.
On dead bark of Acer and lllmus. Not common.

HERPOTRICHIA Fuckel
Herpotrichia Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:146
(1870).
Pseudothecia not embedded in host tissue b u t gregarious and sur-
rounded b y a m a t of interwoven brown hyphae, asci thick-walled,
ascospores 2-celled, sometimes finally 4-celled, hyaline or nearly so.

Herpotrichia macrotricha (Berkeley & Broome) Saccardo, Sylloge


F u n g o r u m 2:213 (1883). (Fig. 16A.)
Pseudothecia gregarious, subglobose with an apical papilla, about
Vz m m - across, covered with spreading brown hairs and embedded in a
dense blackish subiculum. Asci cylindric-clavate, very short stalked,
up to 115x12fi, 8-spored; ascospores biseriate, fusiform, 3 0 - 4 0 x
4.5-5.5fi, at first 1-septate and distinctly constricted at the septum,
containing from four to six oil globules and ultimately sometimes
developing further septa between the globules, hyaline, with a small
hyaline appendage at each end.
On plant debris of various kinds, rotting Carices, fallen twigs and the like.
Not common.

THAXTERIA Saccardo
Thaxteria Saccardo, Sylloge F u n g o r u m 9:687 (1891).

Pseudothecia gregarious on a dark subiculum as in Herpotrichia b u t


containing 3-septate ascospores with the two middle cells brown and
the end cells colourless or nearly so.

"410
Thaxteria phaeostroma (Durieu& Montagne) G. Booth in the Naturalist
866:88 (1958). (Fig. 16B.)
Pseudothecia gregarious, subglobose or slightly ovoid without a well
defined apical papilla, up to % m m . across, blackish, surrounded b y a
dense blackish subiculum composed of dark brown septate hyphae
7-10// wide.Asci cylindric-clavate, sessile, up to 110x20//, 8-spored;
ascospores elliptic-cylindric, usually slightly curved, biseriate, 28-40x
7-9//, 3-septate, the middle cells dark brown, the end cells short and
hyaline or nearly so.

On wood and bark of fallen branches of deciduous trees, in association


with various Diatrypaceae. Not uncommon.
This common species has long been mistakenly referred to the genus
Chaetosphaeria Tulasne (see p. 330). The pseudothecia are preceded by a
patch of black mycelium, bearing conidia very like the ascospores and
formerly called Cladotrichum triseptatum Berkeley & Broome in Ann. Mag.
Nat. Hist. Ser. 2, 7:98 (1851).

SPORORMIA de Notaris

Sporormia de Notaris in Mem. Acad. Torino II, 10:342 (1849).


Pseudothecia scattered, small, more or less immersed in dung, asci
large, ascospores dark brown, with several cross septa and showing a
tendency to break up at the septa, surrounded by a hyaline gelatinous
sheath. This is a large genus containing m a n y cosmopolitan species.
Only one can be dealt with here b u t a convenient t r e a t m e n t of the
genus as a whole by R. F. Cain, will be found in "Studies of Copro-
philous Sphaeriales in Ontario", published as University of Toronto
studies, Biological series 38 (1934).

Sporormia intermedia Auerswald in Hedwigia 7:67 (1868). (PL


XXXVIIL.)
Pseudothecia ovoid, up to % m m . across, largely immersed in dung,
black, smooth. Asci cylindrical, short-stalked, up to 2 3 0 x 2 8 / / ,
8-spored; ascospores arranged in two or three rows, cylindrical,
straight or slightly curved, 40-60x8-12/<, 3-septate, strongly con-
stricted at the septa and readily separating under pressure into uni-
cellular segments, dark brown, enclosed in a hyaline gelatinous sheath
3 - 4 / / thick.

On dung of all herbivorous animals. Very common.

"411
Sporormia minima Auerswald, op. cit.:66 (1868), differs in its small
ascospores, 2 8 - 3 2 x 5 - 6 / / .
Sporormia ambigua Niessl in Osterr. Bot. Zeitschr. 28:165 (1878),
has clavate longer-stalked asci containing ascospores 3 5 - 4 5 x 7 - 9 / / .
Sporormia megalospora Auerswald in Hedwigia 7 : 6 8 (1868), has
ascospores 7 0 - 8 5 x 1 5 - 1 7 . 5 / / .
Sporormia pascua Niessl in Osterr. Bot. Zeitschr. 28:165 (1878),
differs from the preceding species in having 7-septate ascospores
33-40x7-9//.

TRICHODELITSCHIA Munk

Trichodelitschia Munk in Dansk. Bot. Arkiv. 15 (2): 109 (1953).


Pseudothecia scattered in or on dung, ovoid, hairy, ascospores uni-
seriate, brown, 2-celled, with a gelatinous sheath. There is one species:

Trichodelitschia bisporula (Crouan) Munk, op. cit.: 109 (1953). (PI.


XXXVIIJ.)
Pseudothecia scattered, flask-shaped with short stout necks beset
with stiff black hairs up to 130// long, more or less immersed in dung,
blackish, about % m m . across. Asci cylindrical, short-stalked, up to
200x18//, 8-spored; ascospores obliquely uniseriate, ellipsoidal, 1-sep-
t a t e and strongly constricted at the septum, dark brown, 18-26x
7-11//, with a hyaline gelatinous sheath up to 5// thick.

On dung of herbivorous animals. Not uncommon.


In Trichodelitschia the inner wall of the ascus breaks through a pore in
the tip of the outer wall when the ascospores are about to be expelled In
the genus Delitschia the outer wall of the ascus fractures transversely some
way below the tip, as in other Loculo'ascomycetes. See Lindquist inSvensk.
Bot. Tidskr. 58:269 (1964).

DELITSCHIA Auerswald

Delitschia Auerswald in Hedwigia 5 : 4 9 (1866).


Pseudothecia usually immersed in dung, black, minute, with well-
developed necks. Spores dark brown, 1-septate, with germ-slits and
thick hyaline gelatinous coats. The t y p e species is D. didyma Auers-
wald, not yet recorded from the British Isles.

"412
Delitschia winteri (Phillips & Plowright) Saccardo, Sylloge F u n g o r u m
1 : 7 3 4 (1882). (Fig. 20L.)
Pseudothecia elongated pyriform, about % m m . long, black, smooth,
immersed in dung with only the tip of the neck showing as a m i n u t e
black dot. Asci about 350x50//, 8-spored; ascospores uniseriate or be-
coming biseriate, elliptical with a single central transverse septum,
slightly constricted, dark brown, 50-65x28-33/«, the gelatinous coat
disappears in old material, and the germ slits are often hard to see.
On dung, typically of rabbit, in winter.

PLEOSPORA Rabenhorst
Pleospora Rabenhorst in Klotzsch Herb. Viv. Mycol. Ed. Nov. No. 547
(1857), see Flora 40:381 (1857).
Pseudothecia immersed in plant tissue, rather small, brown to black,
smooth, asci thick-walled, ascospores yellow to brown, with several
cross septa and one or more longitudinal septa in most or all of the
segments. This is a large genus and the n u m b e r of British species is
uncertain. A helpful t r e a t m e n t of 53 European species b y E . Miiller
will be found in Sydowia 5 : 2 6 0 - 3 0 8 (1951), with supplementary ac-
counts of additional species in volume 7 (1953). Several Rritish species
are described in a series of papers b y Webster in Trans. Brit, mycol.
Soc. 38:347-365 (1955), 40:177-186 (1957), 42:332-342 (1959) & 44:
417-436 (1961).

Pleospora herbarum (Fr.) Rabenhorst, op. cit. No. 547 (1857). (PI.
XXXIXF.)
Pseudothecia scattered, subepidermal, subglobose to ovoid with an
apical papilla, up to % m m . across, smooth, black. Asci cylindric-
clavate, short-stalked, up to 150x40//, 8-spored; ascospores more or
less biseriate, ovoid with the upper half usually slightly wider t h a n the
lower, 24-40 x 11—16/«, with seven transverse septa, always constricted
at the middle septum and often slightly so at other septa, each segment
f u r t h e r subdivided by one or usually two longitudinal septa, yellowish-
brown.
On herbaceous debris of all kinds but especially on Dicotyledons. One
of the commonest of British fungi.

Pleospora discors (Durieu& Montagne) Cesati & de Notaris in Comm.


Soc. Critt. Ital. 1:218 (1863), is a very similar species found on grasses
and sedges.

"413
Pleospora multiseptata Starback in Botanisk. Notiser:207 (1887), is
remarkable for its very large ascospores with up to 17 cross septa.
(Fig. 1 9 E . )

PYRENOPHORA Fries
Pyrenophora Fries, S u m m a Veg. Scand. Sect. Post.:397 (1849).
This is a small genus distinguished from Pleospora, if at all, b y its
thicker-walled pseudothecia, bearing dark brown bristles on their
upper surface. By the most modern authors it is united with Pleospora.

Pyrenophora trichostoma (Fries) Fuckel in J a h r b . Nass. Vereins f.


N a t u r k u n d e 2 3 - 2 4 : 2 1 5 (1870). ( P I . X X X I X J . )
Pseudothecia scattered, subepidermal, subglobose or flattened at
the base, about y 2 m m . across, black, thick-walled, bearing numerous,
stiff, spreading, blackish bristles. Asci few, somewhat ventricose, thick-
walled, short-stalked, up to 2 1 0 x 5 0 / / , containing up>to eight spores;
ascospores irregularly arranged, ellipsoidal, slightly broader above the
middle, 38-50 x20—24/t, with three cross septa and one longitudinal
septum, only slightly constricted, yellowish-brown, with a t h i n
gelatinous sheath.
On dead leaves and leaf-sheaths of grasses and cereals, figured on Bromus.
This is a collective species, associated with conidial states in the form
genus Helminlhosporium. The ascocarps of the component microspecies
are morphologically alike but the Ilelminthosporium states on various wild
grasses and cereals have received different names; they mostly cause leaf
spots or "Stripe" diseases and in some cases are responsible for serious
decay of cereal seedlings in the early stages of growth.

CUCURBITARIA S. F. Gray
Cucurbitaria S. F. Gray, N a t u r a l Arrangement of British Plants 1:519
(1821).
Pseudothecia small, black, subglobose, clustered on a basal stroma,
asci thick-walled, cylindrical, ascospores uniseriate, brown, with trans-
verse and longitudinal septa.

Cucurbitaria laburni (Persoon ex Fries) Cesati & de Notaris in Com-


mentario Societa Crittogamologica Italiana 1 : 2 1 4 (1863). (PI.
XXXIXE.)

Pseudothecia densely crowded on a blackish m a t of woven hyphae,


a b o u t % m m . across, subglobose or angular b y m u t u a l pressure, with

"414
a small apical papilla, upper surface somewhat flattened or even
slightly depressed round the papilla, wall thick, black, with a finely
roughened outer surface. Asci cylindric-clavate, short-stalked, up to
200x15//, up to 8-spored; ascospores uniseriate, oblong-fusiform,
2 4 - 3 6 x 9 - 1 6 / / , with from five to seven transverse septa, markedly
constricted at the middle septum, and with one or two longitudinal
septa in most segments, yellowish-brown.

On dead twigs and branches of Cytisus laburnum, erumpent through the


bark. Not common, much less so than in France.

Cucurbitaria berberidis (Persoon ex Fries) S. F. Gray, op. cit.:519


(1821), occurs on Berberis and Mahonia.
Cucurbitaria dulcamarae (Kunze & Schmidt ex Fries) Fuckel in
J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:175 (1870), is found oc-
casionally on Solarium dulcamara.
Cucurbitaria elongata (Fries) Greville, Scottish Cryptogamic F l o r a :
t a b . 195 (1827), comes on dead branches of Robinia pseudacacia.
Cucurbitaria pithyophila (Schmidt & Kunze ex Fries) Cesati & de
Notaris, op. cit.:214 (1863), is a rare parasite of various coniferous
trees, and C. piceae Borthwick a t t a c k s buds of Picea.
Cucurbitaria spartii (Nees in Fries) Cesati & de Notaris, op. cit.:214
(1863), occurs not uncommonly on Sarothamnus and Ulex.

OTTHIA Nitschke

Otthia Nitschke apud Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e


23-24:169 (1870).
Pseudothecia like those of Cucurbitaria b u t usually in smaller
clusters and containing only 1-septate ascospores, see Scheinpflug in
Ber. Schweiz. Bot. Ges. 68:373 (1958).

Otthia spiraeae Fuckel, op. cit. :170 (1870). (Fig. 18H.)


Pseudothecia in small clusters on a dark brown subiculum, sub-
globose to pyriform with short stout necks and thick dark brown walls,
e r u m p e n t through cracks in bark. Asci short-stalked, cylindrical, about
160x17//, 8-spored; ascospores uniseriate, elliptical to slightly in-
equilateral, 1-septate, 2 4 - 2 8 x 1 1 - 1 5 / / , slightly constricted at the sep-
t u m , dark brown.

On dead twigs or branches of woody Dicotyledons, especially on Rosa-


ceae, associated with pycnidia in the form-genus Diplodia, pycnospores

"415
morphologically very similar to the ascospores. Apparently rare, though the
Diplodia state is common, but inconspicuous and easily overlooked.

FENESTELLA Tulasne

Fenestella Tulasne, Selecta F u n g o r u m Carpologia 2:208 (1863).


Pseudothecia clustered in a stroma and collectively erumpent from
bark, asci large and thick-walled, ascospores uniseriate, large, brown,
with m a n y transverse and longitudinal septa.

Fenestella fenestrata (Berkeley & Broome) Schroeter, Kryptogamen


Flora von Schlesien 3 (2): 435 (1897). (Fig. 23G.)
Stroma cushion-like, blackish-brown, immersed in bark, enclosing
up to 14 crowded subglobose, black pseudothecia, each up to % m m .
across, their ostioles short and collectively erumpent in the disc of the
stroma. Asci subcylindrical, short-stalked, very thick-walled, up to
340x40[i, 8-spored; ascospores uniseriate, r a t h e r variable in size and
shape, from elliptic-cylindric to somewhat clavate, constricted in the
middle, 30-60x11-28/«, composed of very numerous, small, cuboid or
irregular cells, brown, with a somewhat pointed hyaline cell protrud-
ing at each end.
On dead branches and twigs of many deciduous trees, figured on Quercus,
the type host. Fairly common.

Fenestella vestita (Fries) Saccardo in Michelia 1:575 (1879), has much


smaller ascospores, 19-25x10-12/«.

TEICHOSPORA Fuckel

Teichospora Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:160


(1870).
Pseudothecia superficial or nearly so, gregarious but without a
stromatic base, small, smooth, black, asci cylindrical, ascospores uni-
seriate, like those of Cucurbitaria.

Teichospora obducens (Fries) Fuckel, op. cit.: 161 (1870). (Fig. 16K.)
Pseudothecia densely crowded on blackened patches of decorticated
wood, superficial or with the base slightly sunken in the wood, sub-
globose, somewhat flattened above, a b o u t % m m . across, with a minute

"416
apical papilla, black, smooth or slightly rough. Asci cylindric-clavate,
short-stalked, up to 210x20//, 8-spored; ascospores uniseriate, usually
slightly clavate, 1 6 - 2 5 x 8 - 1 1 / / , w i t h up to seven transverse septa,
constricted at t h e middle septum only, with one or two longitudinal
septa in most segments, yellowish-brown.
On wood of various deciduous trees but especially on Fraxinus; not un-
common.

PLEOMASSARIA Spegazzini

Pleomassaria Spegazzini in Ann. Soc. Cientifica Argentina 9: table


following p. 192 (1880).
Pseudothecia r a t h e r large, immersed in b a r k , without a stroma, asci
large, thick-walled, ascospores large, brown w i t h several transverse
walls and one or two longitudinal septa, sheathed with a gelatinous
coat.

Pleomassaria siparia (Berkeley & Broome) Saccardo, Sylloge F u n g o r u m


2:239 (1883). (Fig. 23E.)
Pseudothecia scattered, immersed in b a r k , r a t h e r flattened, up to
1 m m . or even more across, black, smooth, w i t h o u t an apical papilla
b u t erumpent b y a minute slit in the bark. Asci cylindric-clavate,
thick-walled, up to 250x44//, 8-spored; ascospores more or less bi-
seriate, fusiform-clavate, 50-75x15-22/«, with from five to eight
transverse septa, constricted at each of t h e m , and with one or two
longitudinal septa in t h e broader segments, golden-brown, sheathed
with a hyaline gelatinous coat.
On dead twigs of Betula, in winter and early spring. Not uncommon.

KARSTEN UFA Spegazzini

Karstenula Spegazzini in Decades Mycologici Italiani No. 94 (1879) and


Atti Soc. Grittogamologica Italiana 3 : 5 6 (1881).
Pseudothecia immersed in b a r k , in small clusters surrounded b y a
dark brown mycelium, asci cylindrical, ascospores uniseriate, brown,
with transverse and longitudinal septa, erumpent individually b y a
pore in the bark.

"417
Karstenula rhodostoma (Albertini & Schweinitz ex Fries) Spegazzini,
Decades Mycologici Italiani No. 94 (1879). (PL X X X I X A . )
Pseudothecia gregarious beneath b a r k , surrounded b y a dense, dark-
brown, hyphal web, subglobose, blackish, scarcely % m m . across, with
a m i n u t e apical papilla apparent as a pore on the otherwise unbroken
surface of the bark. Asci narrowly cylindrical, short-stalked, up to
150x12//, 8-spored; ascospores uniseriate, elliptic-cylindric, 3-septate
and strongly constricted at the middle septum, 1 8 - 2 7 x 7 - 9 / / , some-
times w i t h a longitudinal septum in the middle segments, dark brown.

On dead twigs of Rhamnus Frangula; uncommon.

THYRIDIUM Nitschke

Thyridium Nitschke emend Saccardo.in Michelia 1 : 5 0 (1877).


Pseudothecia immersed, often with a clypeus, ascospores like those
of Karstenula.

Thyridium lividum (Fries) Saccardo, op. cit. (Fig. 20A.)


Pseudothecia minute, globose, completely immersed in elongated
ridges with blackened surface on decorticated wood; asci cylindrical,
apex rounded, not blued b y iodine, about 1 2 0 x 9 - 1 0 / / ; ascospores uni-
seriate, elliptic-cylindric, 1 5 - 1 8 x 8 - 9 / / , with 3 to 5 cross septa and a
few longitudinal septa, dark brown.

The blackened ridges in which the pseudothecia are embedded resemble


sterile stromata of some Diaporthe. The name commonly adopted for the
genus is unacceptable as Saccardo emended Nitschke's genus by exclusion
of the type species. Unfortunately the alternative name proposed, viz.
Xylosphaeria Otth in Mitteil. Naturf. Ges. Bern 1870, p. 104 emend. Petrak
in Sydowia 4:18, 1950, is equally untenable, as a later homonym of Xylo-
sphaera Dumortier and apparently also of an earlier concept of Xylosphaeria
by Otth himself.

LEPTOSPORA Rabenhorst

Leptospora Rabenhorst in Hedwigia 1:116 (1857).


Pseudothecia on herbaceous stems, often laterally compressed,
beaked. Asci 8-spored; ascospores very slender, equal, multiseptate.
This is a segregate from Ophiobolus.

"418
Leptospora rubella (Persoon ex Fries) Rabenhorst, Herb. Myc. Ed. II
No. 532 (1857). (PI. X X I X c . )
Also known as Ophiobolus rubellus (Persoon ex Fries) Saccardo in Michelia
2:324 (1881).
Pseudothecia usually scattered over reddish-purple patches on dead
herbaceous stems, developed beneath the epidermis but commonly
shedding it and fully exposed at m a t u r i t y , flask-shaped, up to J / 3 m m .
across, with short cylindrical necks, smooth, black. Asci numerous,
narrowly cylindrical, short-stalked, up to 2 0 0 x 7 / / , 8-spored; asco-
spores lying parallel and spirally arranged in the ascus, 80-180 x
0.5-1.5/1, with numerous transverse septa, almost colourless.

On dead stems of many herbaceous plants, Miiller cites collections on


58 genera in Switzerland, common in England especially on Angelica, Arc-
tium, Chaerophyllum, Heracleum, occasionally also on wood of deciduous
trees.

OPHIOBOLUS Riess

Ophiobolus Riess in Hedwigia 1 : 2 7 (1853).


Pseudothecia immersed mainly in dead tissues of herbaceous plants
or becoming superficial by shedding of the host epidermis, subglobose,
usually with a long apical papilla or short beak, asci cylindrical, thick-
walled at the tip, ascospores long and thread-like, multiseptate,
yellowish to brownish. A useful account of 20 European species will be
found in a paper by E. Miiller in Berichte der Schweizerischen Botani-
schen Gesellschaft 62:307-339 (1952).
This traditional wide concept of the genus is rejected by Holm, who
restricts if to species in which the ascospore separates into halves at matu-
rity. He referred O. rubellus with long, equal, spirally coiled spores to Lepto-
spora and those species with a single enlarged cell in the spore to Nodulo-
sphaeria.

Ophiobolus acuminatus (Sowerby ex Fries) D u b y in Flora 38:493


(1855). (Fig. 30A.)
Pseudothecia developed beneath the epidermis of dead herbaceous
stems, subglobose, less t h a n y 2 m m . diameter, black, with a short
apical beak. Asci cylindrical, 130-150x10//, ascospores parallel in the
ascus, yellow-brown, 120-150x3.5-5//, with 18-30 septa, subcylindric
b u t noticeably constricted at the middle and with one cell near the
constriction distinctly enlarged in each half-spore. The free ascospores
separate into two at the constriction.

"419
Common on stems of Compositae, especially Cirsium and Arctium.
0. cirsii (Karsten) Saccardo, on the same substrata, is distinguished
b y its ascospores with a single enlarged cell near the upper end and no
median constriction or separation into half-spores. (Fig. 30B.)

PODONECTRIA Petch

Podonectria Petch. in Trans. Brit, mycol. Soc. 7:161 (1921).


Pseudothecia clustered on a h y p h a l m a t enveloping scale insects,
light coloured, thick-walled, with or without hairs. Asci and ascospores
like those of Ophiobolus b u t hyaline. Some species are plentiful in t h e
tropics b u t only one is known in t h e British Isles.

Podonectria tenuispora Dennis in Kew Bull. 1957:404 (1958). (Fig.


19D.)

Pseudothecia subglobose, a b o u t 250// across, pale pinkish, com-


pletely covered with erect, white, thick-walled hairs 3.5// thick and
up to 200// long. Asci clavate, 1 5 5 - 1 6 5 x 1 1 - 1 2 / / , 8-spored; ascospores
lying parallel spirally twisted in the ascus, long and thread-like, 3 - 4 / /
thick, multiseptate, hyaline; pseudoparaphyses a b u n d a n t , 1// thick.

Clustered on a hyphal mat partially enveloping the common scale


Lepidosaphis ulmi.

ACANTHOPHIOBOLUS Berlese

Acanthophiobolus Berlese in Atti del Congresso botanico internazionali


di Genova:1892, p. 571 (1893).
This is a small genus distinguished from Ophiobolus by its bristly
pseudothecia and united with it b y most authors.

Acanthophiobolus helminthosporus (Rehm) Berlese, op. cit.:571


(1893). (Fig. 16H.)
Pseudothecia superficial, almost globose to pear-shaped, up to
y 3 m m . across, black, bearing r a t h e r sparse, stiff, spreading, pointed,
blackish bristles. Asci narrowly cylindric-clavate, short-stalked, up to
200x12//, 8-spored; ascospores lying approximately parallel b u t
intricately coiled within the ascus, u p to about 170x2.5//, not dis-
tinctly septate b u t containing m a n y oil drops, hyaline.

"420
On plant debris of various kinds. Not common. Also known as Ophio-
chaeta helminthospora (Rehm) Saccardo, Sylloge Fungorum 11:352 (1895).

TUBEUFIA Penzig & Saccardo

Tubeufia Penzig & Saccardo in Malpighia 11:517 (1897).


Pseudothecia superficial, subcylindrical, often crowned b y a ring of
short processes, asci cylindric-clavate, 4-8-spored, ascospores parallel,
greatly elongated, multiseptate, hyaline. There are two British species:

Tubeufia helicomyces von Hohnel in S.B. K. Akad. Wiss. Wien, Math.-


Nat. Klasse A b t . 1, 118:1477 (1909). (Fig. 9B.)
Pseudothecia scattered or in small clusters on a usually r a t h e r sparse
brown mycelial m a t , superficial, r a t h e r barrel-shaped, yellow, becom-
ing brown, smooth a p a r t from a crown of short, stiff, spreading, sep-
t a t e bristles round the ostiole, about 34 m m - across. Asci numerous,
narrowly cylindric-clavate, short-stalked, thick-walled at the tip, up
to 160x14//, 8-spored; ascospores parallel, almost straight, slender
and tapering each end, mostly 9 0 - 1 1 0 x 2 - 3 / e with about seven to nine
transverse septa, hyaline.

On debris of marsh plants and on grasses in damp places, not uncommon.


The pseudothecia occur in association with patches of a pinkish-grey mould,
bearing spirally coiled conidia, Helicosporium phragmitis von Hohnel.

Tubeufia cerea (Berkeley & Curtis) Booth occurs on old s t r o m a t a of


Diatype stigma.

NAUMOVIA Dobrozrakova

Naumovia Dobrozrakova in Morbi P l a n t a r u m 16:197 (1927).


Pseudothecia clustered on a well developed stroma, erumpent, sub-
globose, thick-walled, smooth, asci cylindrical, ascospores slender,
multiseptate, hyaline. There is only one species unless, as most modern
authors prefer, the genus be united with t h e m u c h older Gibberidea
Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e 23-24:168 (1870).

Naumovia abundans Dobrozrakova, op. cit. (1927). (Fig. 161.)


Stroma elongated, immersed in the cortex of the host, cushion-like,
blackish, bearing numerous, crowded, subglobose, smooth, black

"421
pseudothecia, about y 3 m m . across, becoming erumpent through a slit
in the epidermis. Asci narrowly cylindrical, short stalked, thick-walled
at the tip, up to 100x7.5//, 8-spored; ascospores very slender, pointed
below, somewhat rounded above, 3 0 - 4 0 x 1 . 5 - 3 / / , with from one to six
transverse septa, almost hyaline.
On living' stems of Prunella vulgaris. Uncommon. Also known as Gib-
beridea abundans (Dobrozrakova) Shear in Mycologia 29:361 (1937); the
original species of Gibberidea was G. visci Fuckel, op. cit.: 168 (1870), on
Viscum album, which has brownish ascospores about 38x7//, with several
cross septa.

Family 7. Perisporiaceae

This was formerly regarded as a very large family, including genera


here referred to the Plectascales. After these have been removed there
remains only one British genus:

PERISPORIUM Fries

Perisporium Fries, Systema Orbis Vegetabilis 1 : 1 6 1 (1825).


Ascocarps superficial, subglobose, without ostioles, smooth, black,
asci clavate, stalked, ascospores 3-septate, brown, like those of a
Sporormia.

Perisporium vulgare Corda, Icones fungorum 2:27 (1838). (Fig. 16F.)


Ascocarps superficial, gregarious, subglobose, about y 2 mm. across,
smooth, black. Asci clavate, rather thick-walled, up to 8 0 x 2 0 / / , up to
8-spored; ascospores irregularly arranged, elliptic-cylindric, 2 4 - 3 8 x
5 - 7 / / , 3-septate, strongly constricted at each septum and easily frac-
turing into unicellular segments, dark brown to nearly black.

On vegetable debris, as decorticated wood, rotting cloth, sacking and the


like. Not uncommon.

Order 4. HYSTERIALES

In this order the ascocarp is either elongated, with a well developed


black basal and marginal tissue, forming a characteristic fructification
known as a hysterothecium, or it is of indefinite shape, without a n y
sterile marginal tissue at all. On this basis the genera fall into two
families:

"422
I. Ascocarp indefinite, more or less immersed, lichenised fungi or lichen
parasites Arthoniaceae
If. Ascocarp a hyslerothecium, more or less superficial, elongated, forked,
star-shaped, mussel-shaped or wedge-shaped, non-lichenised fungi on
wood and bark Hysteriaceae

Family 1. Arthoniaceae

Most of the genera are lichenised and only one will be briefly men-
tioned here:

CELIDIUM Tulasne

Celidium Tulasne in Ann. Sci. Nat. Bot. I l l , 17:120 (1852).


Ascocarps immersed or erumpent, more or less rounded and cushion-
shaped, black, parasitic on lichens, asci clavate, thick-walled, short-
stalked, 4-8-spored, ascospores multiseptate, usually colourless.

Celidium varians (Davies) Arnold in Flora 44:678 (1862). (Fig. 11K.)


Ascocarps usually gregarious, circular or angular, immersed, becom-
ing erumpent, up to 1 m m . across, black, upper surface slightly convex,
without a definite marginal tissue. Asci broadly clavate, short-stalked,
thick-walled above, up to 5 0 x 2 0 / / , 8-spored; ascospores irregularly ar-
ranged, elliptical or slightly clavate, 1 2 - 1 8 x 4 - 7 / / , 3-septate, not
constricted, hyaline; interascal hyphae numerous, uniting to form a
dark brown covering layer above the asci.

Parasitic in the hymenium of lichen fungi belonging to the genera


Bacidia, Lecanora and Lecidia.
Compare also Agyrium, p. 215.

Family 2. Hysteriaceae

The genera are differentiated largely on ascospore characters b u t


also in p a r t on the shape of the hysterothecium:

Ilysterothecia shell-like or wedge-shaped, taller than broad:


1. Ascospores needle-like, very long Lophium (p. 427)
2. Ascospores elliptic-cylindric, multiseptate, brown
Mytilidion (p. 427)
Hysterothecia star-shaped, ascospores 1-septate, hyaline
Actidium (p. 426)

"423
Hysterothecia elongated, broader than high:
1. Ascospores brown with a minute hyaline basal protuberance
Farlowiella
2. Ascospores brown throughout:
, A. Ascospores 1-septate Aetidium
, ' B. Ascospores with several transverse septa Hysterium (p. 425)
C. Ascospores with both transverse and longitudinal septa
Hysterographium (p. 426)
3. Ascospores hyaline or pale olivaceous throughout:
A. Ascospores 1-septate Glonium
v. B. Ascospores with transverse and longitudinal septa
Gloniopsis (p. 425)
Compare also Pseudographis, p. 143.

F o r a monograph of the family see Zogg in Beitr. K r y p t . Flora


Schweiz: 11 (3) (1962).

FARLOWIELLA Saccardo
Farlowiella Saccardo, Sylloge F u n g o r u m 9:1100 (1891).
There is only one British species:

Farlowiella carmichaeliana (Berkeley) Saccardo, op. cit.: 1101 (1891).


(Fig. 15A.)
Hysterothecia long and narrow with pointed ends, up to 3 x % mm.,
smooth, black, opening b y a longitudinal slit. Asci cylindric-clavate,
short-stalked, up to 90x18/«, 8-spored; ascospores uniseriate, ovoid,
18-27 x 7-12 fi, ultimately black except for the narrow basal end which
remains hyaline.
On wood or bark, associated with conidiophores bearing solitary black
conidia 15-35x13-24,« (Monotospora megalospora Berkeley & Broome). Un-
common.

GLONIUM Fries
Glonium Fries, Systema Mycologicum 2:594 (1823).
There is only one clearly defined British species:

Glonium lineare (Fries) de Notaris in Giornale Botanico Italiano Anno


2, Tome 1, P a r t 3, p. (27) (1846). (Fig. 15F.)
Hysterothecia usually gregarious, lying parallel, up to about 2 m m .
long and very narrow, smooth, black, with a longitudinal slit. Asci

"424
cylindric-clavate, about 100x12//, 8-spored; ascospores uniseriate,
elliptical or slightly tapering below, 1 2 - 1 5 x 5 - 7 / / , 1-septate, hyaline.
On decorticated wood. Uncommon.

HYSTERIUM de Candolle ex Merat

Hysterium de Candolle ex Merat, Nouvelle Flore de Paris 1:153 (1821).


Hysterothecia elongated, smooth or longitudinally furrowed, black,
ascospores brown, with more t h a n one transverse septum. There are
four British species, all on wood or bark.

Hysterium angustatum Albertini & Schweinitz ex Merat, op. cit.: 153


(1821). (Fig. 15D.)
Hysterothecia scattered, up to about 1 m m . long by V3 m m . across,
smooth, black, with a central longitudinal slit, often accompanied by a
parallel furrow in the wall on each side. Asci clavate, short-stalked,
about 90x17ju, 8-spored; ascospores in two or three rows, elliptic-
cylindric, 1 5 - 2 3 x 5 - 7 j u , 3-septate, slightly constricted at the septa,
uniformly brown t h r o u g h o u t .
On wood and bark of deciduous trees, Acer, Alnus, Fagus, Fraxinus etc.
Common.
Hysterium pulicare Persoon ex Merat, op. cit.: 153 (1821), also on
b a r k of deciduous trees, especially Betula, differs in its slightly larger
ascospores 2 1 - 2 6 x 6 - 8 fi with the end cells usually paler t h a n t h e two
middle cells.
Hysterium acuminatum Fries, Systema mycologicum 2:582 (1823),
occurs on bark of conifers, with uniformly brown 3-septate ascospores
12-18x5-6//.
Hysterium insidens Schweinitz in Trans. American Phil. Soc. Phila-
delphia N.S. 4 : 2 4 4 (1834), is distinguished from all the above b y its
4-9-septate ascospores, 2 1 - 3 8 x 6 - 1 2 / / .

GLONIOPSIS de Notaris

Gloniopsis de Notaris in Giornale Botanico Italiano Anno 2, Tome 1,


P a r t 3, p. (23) (1846).
Modern authors recognise only one British species of Gloniopsis, the
current name of which is:

"425
Gloniopsis levantica Rehm in Hedwigia 25:142 (1886). (Fig. 15B.)
Hysterothecia scattered or gregarious, elongated, straight or curved,
up to about 3 x 0 . 8 mm., often longitudinally furrowed, with a central
slit, smooth, black. Asci narrowly clavate, up to 120x20//, 8-spored;
ascospores biseriate, elliptical or the upper half slightly broader t h a n
the lower, 1 7 - 2 7 x 6 - 1 0 / / , hyaline or slightly yellowish, with from three
to seven transverse septa, constricted at the middle septum, and with
one or two longitudinal septa in most segments.
On bark of dicotyledonous trees and shrubs and on the larger herbaceous
stems, very common on old canes of Rubus "fruticosus" agg.

HYSTEROGRAPHIUM Corda

Hysterographiuin Corda, Icones F u n g o r u m 5 : 3 4 (1842).


Differs from Gloniopsis in its distinctly brown ascospores.

Hyst(>rographium fraxini (Persoon ex Fries) de Notaris in Giornalc


Bolanico Italiano Anno 2, Tome 1, P a r t 3, p. (22) (1846), (Fig. 15c.)
Hysterothecia r a t h e r broadly elliptical, up to 2 x % mm., smooth,
black, with a longitudinal slit. Asci clavate, short-stalked, up to
8-spored, 2 0 0 x 4 0 / / ; ascospores irregularly bissriate, ellipsoidal, 30-50
x 12-20//, with from five to ten transverse septa, sometimes constricted
at the centre, and with from one to three longitudinal septa in most
segments, golden to chestnut-brown.
On bark of deciduous trees, especially Fraxinus. Now very rare except in
west Wales.

Hysterographium elongatum (Fries) Corda, op. cit.: 77 (1842), on de-


corticated wood of Salix and Populus, differs mainly in its opaque
ascospores and is also very rare.
Hysterographium mori (Schweinitz) Rehm, Ascomyceten No. 363
(1876), has ascospores 1 5 - 3 0 x 7 - 1 0 / / with from three to five, oc-
casionally 7, transverse septa and has been found rarely in southern
England and the Channel Islands.

ACTIDIUM Fries

Actidium Fries, Systema mycologicum 2:595 (1823).


Hysterothecia star-shaped, black, with a longitudinal slit along each
arm, ascospores 1-septate, hyaline. There is only one British species
b u t Zogg refers here also some european fungi with 1-septate brown

"426
ascospores in simple elongate hysterothecia formerly referred to
Bulliardella Paoli.

Actidium hysterioides Fries, op. cit.:596 (1823). (Fig. 15i.)


Hysterothecia superficial, about 1 m m . across, each like a minute
black starfish with four or five equal arms, each opening b y a longi-
tudinal slit. Asci not seen; ascospores hyaline, narrowly elliptical,
11-15x2-2.5/«, 1-septate.
On wood and bark of conifers. Apparently extremely rare but no doubt
easily overlooked.

LOPHIUM Fries
Lophium Fries, Systema mycologicum 2:533 (1823).
Hysterothecia erect, shaped like bivalve shells set on edge or like
long narrow axe heads standing on their narrower ends, black, shining,
asci cylindrical, ascospores needle-shaped, multiseptate, pale oliva-
ceous. There are two British species:

Lophium mytilinum (Persoon ex Fries) Fries, op. cit.:533 (1823).


(Fig. 15G.)
Hysterothecia scattered, like minute mussel shells set on edge, about
1 m m . long, black, shining, with a longitudinal slit running the whole
length of the sharp upper edge. Asci narrowly cylindrical, short-
stalked, 8-spored, up to 200x10/«; ascospores parallel, very narrowly
cylindrical, up to 180x2.5/,«, ultimately lip to 20-septate, pale oliva-
ceous.
On old wood and bark of conifers. Rare.
Lophium elatum Greville, Scottish Cryptogamic Flora t a b . 177
(1825), differs in its erect wedge-shaped hysterothecia, up to 2 m m . tall
but only % m m . wide. It occurs on bark of deciduous trees and shrubs
b u t appears to be one of the rarest of European fungi, only found a
dozen times in the past century and a quarter, only twice in the British
Isles.

MYTILIDION D u b y

Mytilidion Duby, Mem. Tribu Hysterinees:62 (1862).


Hysterothecia shell-shaped like those of Lophium mytilinum but
ascospores brown and elliptic-cylindric. The three British species are all
on conifers.

"427
Mytilidion gemmigenum Fuckel in J a h r b . Nass. Vereins f. N a t u r k u n d e
25-26:299 (1871). (Fig. 15E.)
Hysterothecia scattered, about 1 - 1 % m m . long, y 2 m m - wide and
up to 1 m m . tall, with a longitudinal slit along the upper edge, smooth,
black. Asci narrowly cylindrical, up to 130x12fx, 8-spored; ascospores
biseriate, elliptic-cylindric, 30-36x6fx, 7-septate, hyaline at first,
brown at m a t u r i t y .
On coniferous bark. Uncommon.

Mytilidion laeviusculum (Karsten) Saccardo, Sylloge F u n g o r u m 2:


761 (1883), with 1-3-septate ascospores only 16-22x2.5-3fx, occurs
occasionally on coniferous b a r k and rotting pine needles.
Mytilidion karstenii Saccardo, op. cit.:763 (1883), has 3-4-septate
ascospores, 28-42x3.5-4fx.

Order 5. HEMISPHAERIALES

This is a large order, best developed in the tropics, containing minute


fungi mostly found on the surface of leaves and other living green plant
p a r t s . It is easily recognised b y its strongly flattened ascocarps (thyrio-
thecia), circular, elongated or irregular in outline, usually with a central
pore or slit. The few British representatives fall into two families,
readily differentiated b y the structure of the upper wall of the thyrio-
thecium:

I. Upper surface formed of hyphae clearly radiating from the centre


Microthyriaceae
II. Upper surface formed of irregular cells, without a radial arrangement,
except sometimes at the margin of the thyriothecium only
Hemisphaeriaceae (p. 432)

Family 1. Microthyriaceae

Mycelium superficial, septate, brown, forming an open network or


a b s e n t ; thyriothecia superficial, the upper layer composed of nearly
parallel brown hyphae radiating from the central pore or slit. The
known British genera m a y be distinguished as follows:

I. Free superficial mycelium absent, or at least scanty, thyriothecia circular:


A. Ascospores non-septate, colourless (not further described here)
Myiocopron

"428
B. Ascospores 1-septate:
1. Ripe ascospores colourless Microthyrium
2. Ripe ascospores brown Seynesiella (p. 430)
I. Free superficial mycelium present around the thyriothecia or the latter
are elongated:
A. Hyphae with hyphopodia like those of Meliola Asterina (p. 430)
B. Hyphae without hyphopodia:
1. Thyriothecia circular, ascospores 1-septate, colourless
Calothyrium (p. 431)
2. Thyriothecia elongated:
a. Ascospores hyaline, 1-septate Aulographuin (p. 431)
b. Ascospores brown Echidnodes (p. 432)

MICROTHYRIUM Desmazieres

Microthyrium Desmazieres in Ann. Sci. Nat. Bot. II, 15:137 (1841).


Thyriothecia superficial, more or less circular with a central pore,
minute, appearing like fly specks on leaves, grass culms and the like.

Microthyrium nigroannulatum Webster in Trans. Brit, mycol. Soc. 35:


208 (1952). (PL X X X V I F . )
Thyriothecia scattered, up to 90/< across, approximately circular,
grey-brown with a ring of darker cells around the ostiole. Asci numer-
ous, ventricose, broadest below the middle and tapering towards the
thick-walled apex, sessile, up to 2 6 x 8 / / , 8-spored; ascospores ir-
regularly arranged, fusiform, 3.5-10.5x1.5-2.5//, hyaline, 1-septate.

On stems and leaf-sheaths of Dactylis. Common.

Microthyrium gramineum Bommer, Roussel & Saccardo, Sylloge


F u n g o r u m 9:1059 (1891), rather similar to the above b u t without the
dark ring round the ostiole and with ascospores 9 - 1 2 x 2 . 5 - 3 . 5 //, occurs
commonly on Ammophila arenaria.
Microthyrium culmigenum Sydow in Annales mycologici 19:140
(1921), common on various grasses, is larger t h a n either of the preced-
ing, with thyriothecia up to 200// across, asci 35-50// long and asco-
spores 1 0 - 1 3 x 2 - 3 (i.
Microthyrium microscopicum Desmazieres, op. cit.: 138 (1841), occurs
on leaves of evergreens.
Myiocopron ilicinum (de Notaris) Saccardo, Sylloge F u n g o r u m 2:
660 (1883), occurs on the upper surface of Quercus leaves.

"429
SEYNESIELLA A r n a u d

Seynesiella Arnaud in Annales de l'Ecole nationale d'Agriculture de


Montpellier, N. S. 16:202 (1918).
Seynesiella is distinguished from Microthyrium by its ascospores
ultimately becoming brown.

Seynesiella juniperi (Desmazieres) Arnaud, op. cit.:203 (1918). (Fig.


19h.)
Thyriothecia up to 250^t across, broadly conical, shining black and
opaque, without surrounding mycelium on the host surface but con-
nected at numerous points with mycelium beneath the host cuticle.
Asci erect, 5 0 - 6 0 x 1 5 - 2 0 / / , 8-spored; ascospores 1-septate, slightly
clavate, at first hyaline with a large oil drop in each cell, but ultimately
becoming brown; pseudoparaphyses a b u n d a n t , branched and t h r e a d -
like.

On living and dead needles of Juniperus communis.

ASTERINA Leveille

Asterina Leveille in Ann. Sci. Nat. Bot. I l l , 3 : 5 9 (1845).


There is only one British species of this very large tropical genus:

Asterina veronicae (Libert) Cooke in Grevillea 5:122 (March 1877).


(PI. XXXVIG.)
Mycelium dark brown, superficial, forming a network visible to the
eye as small irregular, black patches on living leaves, septate, bearing
numerous lobed hyphopodia. Thyriothecia approximately circular, very
d a r k brown, with a central pore, minute. Asci subglobose, up to
3 2 x 2 4 f i , 8-spored; ascospores irregularly arranged, elliptical, or
slightly clavate, 1 5 - 1 7 x 7 - 8 / t , 1-septate, often slightly constricted,
dark brown.

On leaves of Veronica officinalis, mainly on the upper surface. Not un-


common.
Also known as Dimerosporium veronicae (Libert) Arnaud, Les Asterinees:
174 (1918). Dimerosporium Fuckel in Jahrb. Nass. Vereins f. Naturkunde
23-24:89 (1870), is distinguished from Asterina in the restricted sense by
the absence of paraphysoids.

"430
CALOTHYRIUM Theissen

Calothyrium Theissen in Annales mycologici 10:160 (1912).


Distinguished from Microthyrium by the copious brown superficial
mycelium around the thyriothecia. There is one British species:

Calothyrium pinastri (Fuckel) Theissen in Osterr. Bot. Zeitschr. 62:219


(1912). (PI. X X X V I i i . )
Superficial mycelium composed of numerous brown septate hyphae
2 - 3 / / wide, without hyphopodia; thyriothecia approximately circular,
up to 150// across, with a central pore, dark brown. Asci cylindric-
clavate, subsessile, up to 3 0 x 6 / / , 8-spored; ascospores biseriate, fusi-
form, 8 - 1 0 x 1 . 5 - 2 / / , 1-septate, hyaline.

On rotting needles of Pinus silvestris. Seldom seen but easily overlooked.

AULOGRAPHUM Libert

Aulographum Libert, Plantae Cryptogamicae Arduenna collegit No.


272 (1836).
Hysterothecia elongated, opening by a central slit, from which the
h y p h a e radiate to the margin and pass into a brown superficial myce-
lium, ascospores hyaline.

Aulographum vagum Desmazieres in Ann. Sci. Nat. Bot. II, 19:362


(1843). (Fig. 22B.)
Hysterothecia narrowly elliptical, up to 650x120//, black, opening
by a longitudinal slit and running out at the margin into short, ir-
regularly branched, brown hyphae. Asci broadly clavate, sessile, up to
3 0 x 1 2 / / , thick-walled, 8-spored; ascospores fusiform, straight or
slightly curved, irregularly arranged, 9 - 1 2 x 2 . 5 - 3 / / , 1-septate, hyaline.
On fallen leaves, especially of Hedera helix. Not common.

Aulographum fcstucae Libert, Plantes Crypt. Arduenn. Fasc. IY:373


(1837). (Fig. 30K.)
Thyriothecia scattered, superficial, less t h a n 1 m m . long and up to
80// wide, black, usually branched or even stellate, opening by a longi-
tudinal slit. Asci sessile, elliptical, apex thick-walled, 1 9 - 2 2 x 1 1 - 1 2 / / ,
8-spored; ascospores irregularly arranged, clavate, 1-septate, 9 - 1 4 x
4 - 5 //.
On dead leaf sheaths of grasses.

"431
Aulographum filicinum Libert,- op. cit. No. 275 (1834), with asco-
spores 14-22x2-25/«, occurs on dead ferns. It was transferred to
Leptopeltis von Hohnel in Ann. mycol. 15:368 (1917).

ECHIDNODES Theissen & Sydow


Echidnodcs Theissen & Sydow in Annales mycologici 15:422 (1917).
There is only one British species:

Echidnodes aulographoidcs (Bommer, Roussel & Saccardo) Robertson


in Trans. Brit, mycol. Soc. 3 3 : 1 0 8 ( 1 9 5 0 ) . ( P I . X X X V I D . )
Superficial mycelium formed of brown hyphae 2-3pi wide; thyrio-
thecia elongated, up to 350x225//, black, opening b y a longitudinal
slit, sometimes lobed or triangular, surrounded by a narrow belt of
superficial, radiating, brown hyphae. Asci globose, up to 44/« across,
8-spored; ascospores irregularly arranged, slightly clavate, 17-23x
7-12//, 1-septate, constricted at the septum, ultimately pale brown
and punctate.
On the bark of living twigs of Rhododendron. Common.
The common tropical genus Lembosia Leveille in Ann. Sci. Nat. Bot. Ill,
3:58 (1848), differs in having hyphopodia.

Family 2. Hemisphaeriaceae
This is another very large family of tropical "Fly-speck" fungi,
differentiated from the Microthyriaceae b y the absence of radially dis-
posed hyphae running from the ostiole to the margin of the thyrio-
thecium. Two genera have as yet been reported from the British Isles:
I. Upper wall of the thyriothecium formed of pseudoparenchymatous cells,
arranged radially towards the margin, ostiole absent, ascospores
1-septate, hyaline Clypeolura
II. Upper wall of the thyriothecium formed of irregularly interlocked cells,
with no trace of a radial arrangement, ascospores 1-septate, hyaline
Microthyriella
Only the latter is briefly described here as an example of this very in-
conspicuous group of fungi for the type species of Clypeolum has been shown
to be a "lichen".

MICROTHYRIELLA von Hohnel


Microthyriella von Hohnel in Sitz. Kaiserl. Akad. Wiss. Wien, Mathem.-
N a t u r w . Kl. 118:370 (1909).
The following minute species is probably not u n c o m m o n in E n g l a n d :

"432
Microthyriella osmundae Booth in Kew Bulletin 1957:424 (1958).
(Fig. 18G.)
Thyriothecia superficial, scattered, approximately circular, up to
2 0 0 n across, blackish, upper surface composed of intricately inter-
locked dark brown cells, 2-3// wide, running out at the margin into a
sparse mycelium. Asci broadly clavate, sessile, up to 22x15//, thick-
walled above, 8-spored, separated b y narrow strips of interascal tissue;
ascospores irregularly arranged, narrowly clavate, 7 - 1 1 x 3 - 4 / / , 1-sep-
t a t e and slightly constricted at the septum, hyaline.

On dead stems of Osmunda regalis.

"433
Index

abietina, Crumenula, 139 albofuscus, Sphaeridiobolus, 61


abietinum, Phacidium, 211 albolanata, Torrubietla, 230
abietis, Ascocalyx, 139 albopunclata, Leptosphaeria, 401
abietis, Ccnangium, 138 albotestaceus, Dasyscyphus, 153
abietis, Mitrula, 89 albotomentosa, Torrubiella. 231
abietis, Rehmiellopsis, 366 album, Gloeosporium, 175
abietis, Retinocyclus 112 alchemillae, Coleroa, 377
Abrothallus, 220 Aleuria, 47
abundans, Gibberidea, 422 Allescheria, 340
abundans, Naumovia, 421 alliariae, Leptosphaeria, 392
abundans, Trichophaea, 36 alnea, Tympanis, 113
Acanthophiobolus, 420 alni, Melanconis, 313
acerina, Ciboria, 99 alniella, Pezizella, 131
acerina, Melasmia, 199 alphitoides, Microsphaera, 349
acerina, Mollisina, 164 alpina, Sphaeropezia, 212
acerinum, Rhytisma, 199 alutacea, Otidea, 25
acetabulum, Paxina, 12 alulaceum, Podostroma, 234
acharii, Eutypa, 298 ambiens, Yalsa, 302
acicola, Desmazierella, 33 ambigua, Ombrophila, 107
acicolum, Cenangium, 138 ambigua, Sporormia, 412
acicularis, Cudoniella, 120 amentacea, Ciboria, 99
Acrospermum, 223 amenti, Pezizella, 132
Actidiuin, 426 amenticola, Ciboria, 100
Actiniopsis, 253 amentorum, Taphrina, 78
acuminatum, Hysterium, 425 americana, Rutstroemia, 102
acuminatus, Ojihiobolus, 419 amethystina, Barlaeina, 29
acuta, Leptosphaeria, 390 aminophila, Peziza, 16
acuta, Phoma, 391 ammophilae, Chitonospora, 404
acutipilus, Dasyscyphus, 153 ammophilae, Entosordaria, 276
acuum, Dasyscyphus, 150 Amphisphaerella, 277
advenula, Ciboriopsis, 104 Amphisphaeria, 294
advenula, Karschia, 220 ampliata, Peziza, 18
aeruginascens, Chlorosplenium, 134 amplum, Glonium, 382
aeruginellum, Chlorosplenium, 134 ampullasca, Ceratostomella, 326
aeruginosum, Chlorosplenium, 134 andersoni, Cryptodiscus, 208
aestivum, Tuber, 75 andromedae, Rhytisma, 199
affinis, Paranectria, 254 angelicae, Diaporthopsis, 311
agaricinum, Verticillium, 239 angustata, Coronophora, 332
agnita, Leptosphaeria, 391 angustatum, Hysterium, 425
agropyri, Micronectriella, 290 angustilabrum, Lophiotrema, 380
Agyrium, 215 Anisogramma, 321
albidum, Ascocorticium, 84 Anisostomula, 289
albidum, Helotium, 124 Anixiopsis, 342
albidus, Ascobolus, 58 anomalurn, Ascocorticium, 84
albidus, Hymenoscyphus, 124 anserina, Podospora, 269

"434
Anthostoma, 277 Ascophanus, 63
tf Anthostomella, 276 Ascosphaeraceae, 335
Anthracobia, 44 Ascotricha, 266
apalus, Dasyscyphus, 149 Ascozonus, 66
aparines, Leptosphaeria, 392 ascma, Lachnella, 155
Apiocrea, 239 asemum, Psilachnum, 155
Apioporthe, 310 Aspergillus, 340, 341
Apiorhynchostoma, 296 asperior, Scutellinia, 39
Apiospora, 292 Asterina, 4 30
Apiosporella, 291 asteroma, Trichobelonium, 180
apophysata, Pseudotis, 27 Asteroinassaria, 408
Apostemidium, 222 asterophorus, Hypomyces, 239
appendiculata, Podospora, 270 astragali, Microsphaera, 349
appcndiculatum, Helminthosporium astroidea, Lamprospora, 54
386 Astrosphaeriella, 405
Appcndiculella, 352 athyrii, Taphrina, 79
applanata, Astrosphaeriella, 405 Atichia, 354
applanata, Didymella, 385 Atopospora, 361
applanata, Kirschsteiniella, 405 atra, Leptopodia, 9
applanata, Nectria, 247 atrata, Patellaria, 221
aquifolii, Nectria, 242 atrata, Peziza, 186
aquila, Rosellinia, 286 atrofuscus, Ascobolus, 58
Arachnocrea, 236 atropurpureus, Corynetes, 87
Arachnopeziza, 166 atrovinosa, Peziza, 24
Arachnoscypha, 165 atrovirens, Corynella, 110
aranea, Arachnoscypha, 165 atrovirens, Leotia, 88
aranicida, Torrubiella, 231 auctum, Prosthecium, 314
arctii, Diaporthe, 309 aulographoides, Echidnodes, 432
arctii, Pyrenopeziza, 187 Aulographum, 431
arctostaphyli, Naemacyclus, 208 aurantia, Aleuria, 47
arenarius, Corynetes, 87 aurantiaca, Barya, 231
arenicola, Sepultaria, 32 aurantiaca, Sphaerostilbe, 250
arenosa, Sepultaria, 31 aurantiacum, lleleococcum, 338
arenula, Nectria, 248 aurantius, Hypomyces, 239
areola, Anisostomula, 289 aurata, Arachnopeziza, 166
areolata, Boudiera, 53 aurea, Taphrina, 77
argenteus, Ascophanus, 63 aurelia, Arachnopeziza, 166
argillaceum, Ilvpoxylon, 280 aureola, Nectria, 250
ariae, Dermea, 173 aureonitens, Nectriopsis, 251
aridula, Gorgoniceps, 115 auricolor, Orbilia, 168
arundinacea, Leptosphaeria, 399 avenaria, Leptosphaeria, 398
arundinaceum, Lophodermium, 201
arvernensis, Peziza, 18
aschersonia, Ciboria, 99 babingtonii, Chaetothyrium, 368
Ascobolus, 56 babingtonii, Psilopezia, 55
Ascocalyx, 139 baccarum, Monilinia, 98
Ascochyta, 385 baccilifera, Pragmopora, 140
Ascocorticium, 84 baccilifera, Scleroderris, 140
Ascocoryne, 109 badia, Peziza, 21
Ascodesmis, 61 badioconfusa, Peziza, 22
Ascomyces, 77 badiofusca, Peziza, 23
balsamea, Thyronectria, 256

"435
Balsamia, 74 Boydia, 293
barbara, Ostropa, 225 bramleyi, Ciboriopsis, 104
barbatus, Dasyscyphus, 153 brassicae, Mitrula, 89
Barlaeina, 29 brassicae, Nectria, 248
Barya, 231 brassicicola, Mycosphaerella, 364
basitricha, Strossmayeria, 116 brevipila, Pirottaea, 181
basitrichum, Belonidium, 116 brevipilus, Dasyscyphus, 148
batschiana, Ciboria, 99 brevirostris, Melanospora, 264
Battarrina, 238 Broomella, 387
baumleri, Microsphaera, 349 broomeanus, Hypomyces, 239
baumleri, Ombrophila, 100 Brunchorstia, 139
bellidis, Protomycopsis, 82 brunnea, Sphaerosporella, 39
Belonioscypha, 127 brunneoatra, Peziza, 23
Belonopsis, 179 buchsii, Pyronema, 63
berberidis, Cucurbitaria, 415 Buellia, 219
berberidis, Microsphaera, 349 Buergenerula, 387
berkeleyanum, Schizoxylon, 225 bufonia, Massariella, 293
berkeleyi, Phomatospora, 320 bufonia, Otidea, 26
berkeleyi, Sphaeria, 311 bulbosa, Helvella, 12
Berlesiella, 370 Bulgaria, 111
Bulgariella, 111
bernardiana, Rhabdospora, 392
bullata, Taphrina, 80
berolinensis, Thyronectria, 256
Bulliardella, 427
Bertia, 332
buxi, Fusarium, 246
betulae, Ciboria, 100
buxi, Gibberella, 253
betulae, Cryptospora, 317
buxi, Hyponectria, 289
betulae, Taphrina, 78
buxi, Volutella, 240
betulina, Atopospora, 361
Byssolophis, 381
betulina, Euryachora, 361
Byssonectria, 238
betulina, Taphrina, 78
Biatorella, 215
bicolor, Dasyscyphus, 150 caerulescens, Taphrina, 80
bicornis, Uncinula, 348 caesia, Eriopeziza, 165
bicuspis, Trichophaea, 36 caespiticia, Mollisia, 184
bifrons, Hypospilina, 323 Cainia, 288
biseptata, Buergenerula, 387 Cainiella, 289
bisporula, Trichodelitschia, 412 callicarpa, Caryospora, 406
bloxami, Diplocarpa,146 Calloria, 191
bloxami, Karschia, 219 callunae, Pseudophacidium, 209
bohemica, Ptychoverpa, 2 callunigera, Godronia, 141
bohemica, Rehmiellopsis, 366 Calonectria, 252
Bolinia, 282 Caloscypha, 47
Bombardia, 271 Calosphaeria, 305
bombycina, Stephensia, 73 Calosporella, 316
bonordenii, Heteropatella, 142 calostroma, Appendiculella, 352
bostrychodes, Chaetomium, 265 calostroma, Irene, 352
Bothrodiscus, 139 Calothyrium, 431
Botryosphaeria, 373 calthae, Pseudopeziza, 190
Botrytis, 95 calvescens, Cheilymenia, 41
Boudiera, 52 Calycella, 118
boudieri, Ascobolus, 58 Calycellina, 164
boudieri, Discinella, 135 calyciiforme, Triblidium, 142

"436
Calyculosphaeria, 333 cerealis, Gibellina, 262
calyculus, Hymenoscyphus, 123 cerebriformis, Hydnobolites, 76
Camarops, 282 cerinus, Dasyscyphus, 152
campanulaeformis, Pezizella, 132 Ceriophora, 294
campbellii, Underwoodia, 24 cervina, Fimaria, 52
canadensis, Cordyceps, 230 cesatiana, Leptosphaeria, 393
candolleana, Sclerotinia, 93 cesatii, Pocillum, 114
capitata, Cordyceps, 229 ceuthocarpa, Linospora, 326
Capnodium, 367 Ceuthospora, 211
capreae, Linospora, 325 Chaetapiospora, 288
Capronia, 370 Chaetomium, 265
carbonaria, Geopyxis, 48 chaetomium, Coleroa, 376
carbonaria, Sordaria, 277 Chaetosphaeria, 330
carbonarius, Ascobolus, 58 Chaetothyrium, 368
carbonigena, Octospora, 50 chailletii, Diaporthe, 309
caricinum, Lophodermium, 201 chartarum, Ascotricha, 266
carinthiaca, Mvcosphaerella, 364 chateri, Melastiza, 44
carmichaeliana, Farlowiella, 424 Cheilymenia, 40
carnea, Pezicula, 175 chionea, Melanospora, 263
carneoalbus, Oomyces, 232 chionea, Mollisia, 186
carneoflavida, Calloria, 192 chionea, Pezizella, 131
carneola, Octospora, 51 Chitonospora, 403
carneus, Iodophanus, 64 Chlorociboria, 133
Carpenteles, 337 Chloroseypha, 117
carpinea, Pezicula, 175 Chlorosplenium, 133
carpini, Taphrina, 78 Choiromyces, 76
carpinicolum, Helotium, 126
carpophila, Xylosphaera, 285 chondrospora, Pseudomassaria, 291
Caryospora, 405 Chromocrea, 235
cassandrae, Godronia, 140 chrysosperma, Apiocrea, 240
castagnei, Scirrhia, 360 chrysospermum, Sepedonium, 240
Catinella, 176 chrysostigma, Pezizella, 132
catinus, Pustularia, 27 chrysostroma, Melanconis, 313
caucus, Ciboria, 99 Ciboria, 99
caudatus, Hymenoscyphus, 123 Ciborinia, 94
Caudospora, 312 Ciboriopsis, 103
caulium, Xaernacyclus, 207 cichoracearum, Erysiphe, 350
Celidium, '.23 Ciliaria, 38
Cenangium, 137 ciliata, Ilyalopeziza, 159
Cephalosporium, 228, 249 ciliatus, Lasiobolus, 62
Cephalotheca, 343 ciliifera, Sphagnicola, 108
cerasi, Dermea, 173 cinerea, Mollisia, 183
cerasi, Taphrina, 79 cinerea, Ostropa, 225
cerastiorum, Pseudopeziza, 190 cingulata, Glomerella, 258
Ceratocystis, 336 cinnabarina, Nectria, 241
Ccratosphaeria, 327 cinnabarina, Pulvinula, 55
Ceratostomella, 326 cinnabarinnm, Verticillium, 249
cerea, Ophionectria, 255 cinnamomea, Pezicula, 175
cerea, Peziza, 20 circinans, Coleroa, 377
cerea, Tubeufla, 255, 421 circinans, Cudonia, 90
cerealis, Gelasinospora, 268 circinans, Leptosphaeria, 404
circinans, Trematosphaeria, 404

"437
cirrhata, Unguicularia, 160 conjuncta, Hypodermella, 203
cirsii, Ophiobolus, 420 conoidea, Didymosphaeria, 386
Cistella, 156 conscriptum, Helotium, 123
citrina, Calycella, 119 conspersa, Tympanis, 112
citrino-aurantia, Nectria, 249 conspicua, Fuckelia, 205
citrinus, Pachyphloeus, 74 constellatio, Pulvinula, 54
cladoniaria, Nesolechia, 217 controversus, Dasyscyphus, 148
Cladotrichum, 411 cookeana, Anisostomula, 289
clandestinus, Dasyscyphus, 151 cookei, Rhyparobius, 65
Clathridium, 290, 292 cookeianum, Geoglossum, 86
clavariarum, Helminthosphaeria, coprinaria, Cheilymenia, 41
272 Coprobia, 49
clavariarum, Spadicoides, 273 coprophila, Lasiosordaria, 270
Claviceps, 226 Coprotinia, 103
Clavidisculum, 150 coracina, Graddonia, 178
clavispora, Patellariopsis, 177 Cordyceps, 227
clavisporus, Dasyscyphus, 149 corium, Cyathipodia, 11
clavulata, Ophiocordyceps, 230 cornea, Mollisia, 185
clavus, Cudoniella, 120 corni, Pseudomassaria, 291
clivensis, Diapleella, 295 cornubiensis, Cheilymenia, 40
Clypeolum, 432 coronaria, Sarcosphaera, 15
Clypeosphaeria, 295 coronata, Cyathicula, 128
Clythris, 199 coronatus, Coccomyces, 203
coccinea, Nectria, 243 Coronophora, 332
coccinea, Sarcoscypha, 67 corticiorum, Helminthosphaeria,
coccinella, Orbilia, 169 273
Coccomyces, 203
Coccophacidiuin, 207 corticola, Griphosphaeria, 290
cochleata, Otidea, 26 corticola, Hyalotricha, 159
cochliodes, Chaetomium, 265 corticola, Pezicula, 174
cohaerens, Hypoxylon, 280 corvina, Onygena, 344
Coleosperma, 178 coryli, Mamiana, 323
Coleroa, 376 coryli, Nectria, 242
corylina, Cryptospora, 317
collapsa, Calyculosphaeria, 334 Coryne, 109*
Colletotrichum, 258 Corynella, 110
Colpoma, 199 Corynetes, 87
commune, Hypoderma, 202 costifera, Paxina 12
commutata, Durella, 141 crataegi, Sclerotinia, 97
compressum, Acrospermum, 223 crataegi, Taphrina, 80
compressum, Lophidium, 381 craterium, Trochila, 195
concentrica, Daldinia, 284 craterium, tfrnula, 69
concinna, Otidea, 26 crec'hqueraultii, Lamprospora, 54
confluens, Hypoxylon, 281 Creopus, 235
conformata, Rutstroemia, 102 crepini, Leptosphaeria, 391
confusa, Cudonia, 91 cretea, Tricharia, 34
conica, Verpa, 4 crispa, Helvella, 8
Coniochaeta, 273 crispula, Urceolella, 160
Coniocybe, 104 crispum, Colpoma, 200
Coniothyrium, 399 crosslandii, Ascobolus, 58
coniothyrium, Leptosphaeria, 401 crouani, Ascobolus, 58
coniothyrium, Melanomma, 401 crouanii, Lamprospora, 53

"438
crucipila, Cheilymenia, 41 Daldinia, 283
Crumenula, 139 Daleomyces, 24
Cryptodiaporthe, 310 dalmeniensis, Peziza, 41
Cryptodiscus, 193 Dasyobolus, 59
Cryptomyces, 205 Dasyscyphus, 147
Cryptomycina, 210 decipiens, Anthostoma, 277
Cryptosphaeria, 301 decipiens, Podospora, 270
Cryptospora, 317 deerata, Pseudombrophila, 37
Crvptosporella, 317 deformans, Taphrina, 79
Cryptosporiopsis, 173 deformis, Octospora, 50
cryslallinus, Dasyscyphus, 152 degenerans, Colpoma, 200
cubicularis, Robergea, 226 degenerans, Sporomega, 200
cucullata, Mitruia, 89 delicatula, Protocrea, 236
Cucurbitaria, 414 delicatum, Ilaplographium, 158
cucurbitula, Scoleconectria, 254 Delitschia, 412
Cudonia, 90
dematiicola, Ilyaloscypha, 158
Cudoniella, 119
Dendrodochium, 246
culmicola, Belonioscypha, 128
Dendrostilbella, 110
culmicola, Leptosphaeria, 397
dennisii, Sclerotinia, 95
culmicola, Sphaeria, 397
dentatus, Coccomyces, 204
culmifida, Keissleriella, 388
denudatus, Ascobolus, 57
culmifida, Trichometasphaeria, 388
depauperatus, Saccobolus, 60
culmifraga, Leptosphaeria, 397
depressa, Pachyclla, 55
culmifraga, Sphaeria, 397
(lerasa, Leptosphaeria, 394
culmigenum, Lophodermium, 201
derasa, Xodulosphaeria, 394
culmigenurii, Microthyrium, 429
Dermatina, 355
culmorum, Leptosphaeria, 398
Dermea, 172
cuniculi, Caprotinia, 103
Desmazierella, 33
cunicula, Rutstroemia, 103
desmazieri, Ilypoderma, 202
cupressina, Pithya, 67
desmazierii, Nectria, 246
cupularis, Xitschkia, 333
(leusta, Ustulina, 282
cupularis, Pustularia, 28
devexa, Plagiostoma, 324
cupuliformis, Helvetia, 11
Dialonectria, 241
curreyana, Sclerotinia, 94
Diapleella, 295
currevi, Apiorhynchostoma, 296
Diaporthe, 308
curreyi, Pteridiospora, 407
Diaporthopsis, 311
curvatispora, Orbilia, 168
Diatrype, 298
curvicola, Podospora, 270
Diatrypella, 299
curvipila, ltyaloscypha, 158
dickiei, Gibbera, 378
curvula, Podospora, 269
dictvdiola, Lamprospora, 54
cyanogena, Gibberella, 253
didyma, Delitschia, 412
Cyathicula, 128
Didymascella, 208
Cyathipodia, 10
Didymella, 385
cyathoidea, Phialea, 129
Didymellina, 363
cylichnium, Coryne, 110
Didymosphaeria, 385
Cylindrocarpon, 244, 245
Diehliomyces, 345
Cylindrocolla, 192
digitaliformis, Verpa, 5
cylindrospora, Ophionectria, 255
Diinerosporium,430
Cymadothia, 364
diminutus, Dasyscyphus, 150
Cyrtidula, 356
Diplocarpa, 146
Cytospora, 302
Diplocarpon, 189

"439
Diplodia, 415 elatum, Chaetomium, 265
Diplodina, 385 elatum, Lophium, 427
diplospora, Didymosphaera, 386 elegantior, Haglundia, 181
disciformis, Diatrype, 298 ellisii, Nectria, 249
Discina, 12 elongata, Cucurbitaria, 415
Discinella, 135 elongatum, Hysterographium, 426
Disciotis, 5 Elsinoe, 355
Discodiaporthe, 312 Emericella, 341
discolor, Mollisia, 184 Emericellopsis, 343
discolor, Potebniamyces, 204 emileia, Peziza, 20
discors, Passeriniella, 403 emperigonia, Lizonia, 369
discors, Pleospora, 413 empetri, Duplicaria, 204
discospora, Coniochaeta, 273 empetri, Melasmia, 204
Discothecium, 372 empetri, Phaeangellina, 117
discreta, Nummularia, 283 empetri, Physalospora, 258
discreta, Pezizella, 131 empetri, Sphaeropezia, 212
dissepta, Quaternaria, 301 Enchnoa, 304
distinguendus, Dasyscyphus, 153 Encoelia, 138
ditissima, Nectria, 244 Encoeliopsis, 137
ditopa, Ditopella, 320 Endodothella, 261
Ditopella, 319 Endostigme, 375
ditricha, Yenturia, 375 Endothia, 322
Endoxyla, 304
dolioloides, Leptosphaeria, 394
Entodesmium, 390
doliolum, Leptosphaeria, 390
Entosordaria, 275
domesticum, Pyronema, 63
Ephelina, 179
Dothidea, 358
ephippium, Leptopodia, 10
dothidea, Botryosphaeria, 373 epiblemastica, Scutula, 217
Drepanopeziza, 189 epicalamia, Leptosphaeria, 395
dryina, Calosphaeria, 305 epicalamia, Phaeosphaeria, 396
dryinus, Dasyscyphus, 153 Epicliloe, 231
Duebenia, 197 epicymatia, Pharcidia. 371
dulcamarae, Cucurbitaria, 415 epiphyllus, Hymenoscyphus, 126
Duplicaria, 204 episema, Biatorina, 217
Durandiella, 172 episema, Scutula, 217
Durandiomyces, 24 episphaeria, Nectria, 246
Durella, 141 episphaeria, Sphaeroderma, 264
duriaeana, Sclerotinia, 95 epistroma, Dendrodochium, 246
duriaei, Myriangium, 355 equina, Onygena, 344
Dyslachnum, 151 eres, Diaporthe, 309
ericae, Scleroderris, 140
earliana, Diplocarpon, 189 Erinella, 147
eburnea, Massarina, 406 Eriopeziza, 165
eburnea, Pezizella, 132 eriophori, Pezizella, 133
Echidnodes, 432 Ervsiphe, 349
echinophila, Rutstroemia, 102 esculenta, Gyromitra, 6
echinospora, Peziza, 19 esculenta, Morchella, 3
egenula, Phialea, 129 eumorpha, Diaporthe, 309
Elaphomyces, 344 eunomia, Cryptosphaeria, 301
elastica, Leptopodia, 9 euomphala, Tympanopsis, 334
elata, Morchella, 3 euonymi, Microsphaera, 349
elatina, Pseudographis, 143

"440
euphorbiae, Sphaerotheca, 347 fimicola, Sphaeronaemella, 336
Eupropolella, 213 fimiseda, Podospora, 270
eurhizus, Microscypha, 163 firma, Rutstroemia, 101
eurotioides, Unguiculella, 161 flscellum, Apostemidium, 223
Eurotium, 339 flammea, Perrotia, 154
Euryachora, 361 flammea, Sphaerostilbe, 250
eustoma, Leptosphaeria, 397 flaveola, Hyaloscypha, 158
eustoma, Phaeosphaeria, 398 flavida, Lasionectria, 249
eustomoides, Leptosphaeria, 398 flavida, Spathularia, 90
Eutryblidiella, 220 flavovirens, Melanconis, 313
Eutypa, 298 flavoviridis, Nectria, 247
Eutypella, 300 flavoviridis, Sphaerostilbe, 247
excavatum, Tuber, 75 flavum, Phaeohelotium, 118
excelsior, Belonopsis, 179 flexella, Lithographa, 216
exigua, Hysteropezizella, 188 flexella, Placographa, 216
exigua, Ploettnera, 195 foedans, Massaria, 408
exilis, Niesslia, 329 foliacea, Neobulgaria, 108
eximia, Sarcosphaera, 15 foliacea, Sepultaria, 31
Exoascus, 77 foliicola, Pyrenopeziza, 187
foliorum, Botryosphaeria, 374
Fabraea, 190 forquignoni, Cordyceps, 228
fagineus, Hymenoscyphus, 126 fossulae, Leucoscypha, 33
falcipila, Unguicularia, 162 fragariae, Mycosphaerella, 364
fallax, Geoglossum, 86 fragiforme, Hypoxylon, 279
farinosa, Protocrea, 236 fragiformis, Balsamia 74
Farfowiella, 424 frangulae, Pezicula, 175
fascicularis, Encoelia, 139 fraxini, Hysterographium, 426
fasciculata, Bombardia, 271 fructigenus, Hymenoscyphus, 123
fasciculatum, Graphium, 303 Fuckelia, 205
favacea, Diatrypella, 299 fuckeliana, Nectria, 245
Fenestella, 416 fuckeliana, Sclerotinia, 95
fenestrans, Sydowiella, 320 fuckelii, Leptosphaeria, 396
fenestrata, Fenestella, 416 fuckelii, Phaeosphaeria, 396
fenestrata, Hysteropezizella, 188 fuckelii, Pyrenopeziza, 187
ferruginea, Siilia, 319 fugiens, Dasyscyphus, 150
ferrugineum, Helotium, 192 fulgens, Caloscypha, 47
ferruginosum, Cenangium, 137 fuliginosa, Godronia, 141
festucae, Aulographum, 431 fulva, Plicaria, 24
festucae, Botryosphaeria, 374 fulvum, Helotium, 127
fibrillosa, Cheilymenia, 40 fulvum, Polystigma, 260
fllicina, Taphrina, 80 fumigata, Sartorya, 341
filicinum, Aulographum, 432 fumigatus, Aspergillus, 341
fllicinus, Rhopographus, 360 funicola, Nectria, 249
filispora, Belonopsis, 179 furfuracea, Encoelia, 139
Fimaria, 51 furfuraceus, Ascobolus, 57
fusarioides, Calloria, 191
fimbriata, Ceratocystis, 336 fusarioides, Peziza, 192
fimbriata, Mamiana, 323 Fusarium, 245, 246, 247, 248, 253
fimbriata, Stictis, 224 290
fimbriata, Tricharia, 34
fimeti, Peziza, 24 fusca, Tapesia, 182
fimicola, Sordaria, 268 fuscescens, Dasyscyphus, 150

"441
fuscidulum, Melanomma, 409 gnomon, Gnomonia, 324
fuscocincta, Calycella, 119 Gnomonia, 324
fuscum, Hypoxylon, 281 Gnomoniella, 325
Fusicladium, 375 godeni, Leptosphaeria, 399
fusispora, Humaria, 50 Godronia, 140
fusisporum, Sphaeroderma, 264 Gorgoniceps, 115
futilis, Didymosphaeria, 386 gracilis, Cordyceps, 228
Graddonia, 178
gaeumannii, Phaeocryptopus, 378 graminea, Belonopsis, 180
Gaeumannomyces, 321 graminearum, Fusarium, 253
gaillardiana, Pustularia, 28 gramineum, Lophodermium, 201
Galactinia, 16 gramineum, Microthyrium, 429
galeopsidis, Erysiphe, 350 graminis, Erysiphe, 350
galiorum, Leptosphaeria, 392 graminis, Gaeumannomyces, 321
gallica, Keissleriella, 388 graminis, Leptosphaeria, 397
gallica, Therrya, 207 graminis, Phaeosphaeria, 397
galligena, Nectria, 244 graminis, Phyllachora, 261
gardneri, Daleomyces, 24 graminum, Acrospermum, 223
Gelasinospora, 267 grandis, Cudoniella clavus var., 120
gelatinosus, Creopus, 235 granulata, Coprobia, 49
gemmarum, Pezizella, 132 granulatus, Elaphomyces, 344
gemmiferum, Discothecium, 372 granulosellum, Psilachnum, 156
gemmigenum, Mytilidion, 428 granulospermus, Saccobolus, 60
Genea, 73 Graphium, 303
gregaria, Coronophora, 332
Geoglossum, 86
gregaria, Physalospora, 374
Geopyxis, 48
gregaria, Trichophaea, 36
geranii, Venturia, 376
Griphosphaeria, 290
Gibbera, 377
grisella, Microscypha, 163
Gibberella, 253 grossulariae, Microsphaera, 349
Gibberidea, 422 guernisaci, Apostemidium, 223
Gibellina, 262 guernisaci, Helotium, 118
gigas, Gyromitra, 7 Guignardia, 373
gigas, Mitrophora, 4 guttata, Phyllactinia, 351
gilva, Tricharia, 34 Gymnoascaceae, 335
glaber, Ascobolus, 58 Gyrocratera, 72
glaber, Saccobolus, 59 Gyromitra, 6
glaucum, Pyronema, 63
Gliocladium, 243, 251
globoso-pulvinata, Ascophanus, 55 Habrostictis, 176
globosum, Ghaetomium, 265 haematites, Leptosphaeria, 400
globosum, Sarcosoma, 69 Haglundia, 180
Gloeosporidiella, 189 hahniana, Lachnellula, 155
gloeosporioides, Colletotrichum, 258 hamulata, Mollisia, 162
Gloeosporium, 175, 258 hamulata, Unguiculella, 161
Gloeotinia, 98 Haplographium, 158
glomerata, Venturia, 377 Haplosporella, 399
Glomerella, 258 hederae, Hypoderma, 202
glomerulosa, Atichia, 354 hederae, Nectria, 245
Gloniopsis, 425 Heleococcum, 338
Glonium, 424 helicomyces, Tubeufia, 421
glutinosum, Geoglossum, 86 Helicosporium, 421

"442
helminthicola, Leptobelonium, 116 Hydnobolites, 76
helminthicola, Letendraea, 386 Hydnotrya, 72
Helminthosphaeria, 272 Hymenoscyphus, 121
helminthospora, Ophiochaeta, 421 hyperici, Metasphaeria, 389
Helminthosporium, 386, 414 hypnina, Mollisia, 162
helminthosporus, Acanthophiobolus, hyperboreus, Ascobolus, 61
420 Ilypocopra, 278
Helolachnum, 148 Hypocrea, 234
helotioides, Dasyscyphus, 156 Hypocreopsis, 236
Helotium, 113, 120, 121 Hypoderma, 202
Helvella, 7 Hypodermella, 202
helvellae, Sphaeronaemella, 336 hypodermia, Cryptosporella, 317
hemisphaerica, Humaria, 35 hypogaea, Roesleria, 105
hemisphaerioides, Trichophaea, 36 Ilypomyces, 239
hepatica, Fimaria, 52 Ilyponectria, 289
herbariorum, Eurotium, 340 Hypospilina, 323
herbarum, Hymenoscyphus, 125 Hypoxylon, 279
herbarum, Pleospora, 413 hypoxylon, Xylosphaera, 285
Hercospora, 314 hysterioides, Actidium, 427
hercynica, Rutstroemia, 101 hysterioides, Lophodermium, 201
Ilerpotrichia, 410 Hysterium, 425
Hysterographium, 426
herpotrichoides, Leptosphaeria, 397
Hysteropezizella, 188
herpotrichoides, Phaeosphaeria, 397
Hysterostegiella, 188
heteracantha, Peroneutypa, 303
Heteropatella, 142
Heterosphaeria, 142 ilicina, Trochila, 195
heterospora, Leptosphaeria, 404 ilicincola, Pithyella, 162
heterospora, Trematosphaeria, 405 ilicinum, Myiocopron, 429
hetieri, Neotiella, 43 ilicis, Stegia, 196
hinnulea, Sphaerosporella, 40 ilicis, Trochila, 196
hippotrichoides, Xylosphaera, 285 immersus, Dasyobolus, 59
hirsuta, Lasiosphaeria, 272 immutabilis, Hymenoscyphus, 124
hirsutum, Trichoglossum, 85 inaequalis, Venturia, 375
hirta, Trichonectria, 255 incertella, Phialea, 130
hirtella, Sclerotinia, 94 incertum, Helotium, 186
hispidula, Genea, 73 inclusa, Battarrina, 238
Homostegia, 359 infernalis, Enchnoa, 305
hortensis, Morchella, 4 infestans, Phacidiurn, 211
houghtonii, Pezicula, 175 inflatula, Orbilia, 168
howeianum, Hypoxylon, 279 infula, Gyromitra, 7
howsei, Peziza, 21
innesii, Calosporella, 316
humana, Sordaria, 268
inquilinum, Psilachnum, 156
Humaria, 35
inquinans, Bulgaria, 111
humillima, Anthracobia, 46 inquinans, Massaria, 408
humosa, Octospora, 50 insculpta, Vialaea, 293
humuli, Sphaerotheca, 347 insidens, Hysterium, 425
hyalina, Hyaloscypha, 157 insitiva, Valsaria, 316
Hyalinia, 169 inspersa, Leciographa, 219
Hyalopeziza, 159 intermedia, Sporormia, 411
Hyaloscypha, 157 intermixta, Sphaerulina, 366
Hyalotricha, 159 inundatus, Protomyces, 81

"443
inventa, Nectria, 249 lacunosa, Helvella, 8
Iodophanus, 64 lacustre, Coleosperma, 178
ionella, Peziza, 22 Laetinaevia, 196
Irene, 352 laeviusculum, Mytilidion, 428
iridis, Belonopsis, 180 lagenaria, Melanospora, 264
iridis, Mycosphaerella, 363 lagerbergii, Scleroderris, 139
Isaria, 228 Lahmia, 219
islandica, Ghaetapiospora, 288 lampadophora, Ceratosphaeria, 327
Isothea, 260 Lamprospora, 53
lamyi, Thyronectria, 256
johansonii, Gainiella, 289 lanata, Enchnoa, 304
johansonii, Taphrina, 77 lanciformis, Pseudovalsa, 318
johnsonii, Monilia, 97 Laquearia, 213
johnstoni, Encoeliopsis, 137 laricina, Tympanis, 113
junci, Endodothella, 261 laricinum, Cistella piceae var., 157
junci, Phyllachora, 262 Lasiobolus, 62
juncicola, Leptosphaeria, 395 Lasiobotrys, 379
juncicola, Loramyces, 328 Lasionectria, 241, 249
juncicola, Phaeosphaeria, 395 Lasiosordaria, 270
juncina, Leptosphaeria, 395 Lasiosphaeria, 271
juncina, Phaeosphaeria, 395 Lasiostictis, 224
junciseda, Cudoniella, 121 lateritia, Byssonectria, 238
juncisedum, Helotium, 121 lateritio-album, Psilachnum, 156
juncorum, Ciboria, 100 laurocerasi, Trochila, 195
jungermanniae, Mniaecia, 214 lecanodes, Nectria, 250
juniperi, Colpoma, 200 Leciographa, 218
juniperi, Metacapnodium, 367 leciographoides, Mycomelaspilea,
juniperi, Seynesiella, 430 218
juniperina, Kriegeria, 117 ledi, Monilinia, 98
juniperinum, Lophodermium, 201 ledi, Pseudophacidium, 210
leiocarpa, Plicaria, 14
Karschia, 219 Lejosphaerella, 292
karstenii, Mytilidion, 428 Lembosia, 432
Karstenula, 417 leonis, Terfezia, 76
Keissleria, 387 leontodontis, Protomycopsis, 82
Keissleriella, 388 Leotia, 87
Keithia, 208 leporina, Podosordaria, 287
keithii, Ascophanus, 64 leporum, Fimaria, 52
keithii, Nectria, 243 Leptobelonium, 116
kerverni, Saccobolus, 60 Leptopeltis, 432
killiani, Mycosphaerella, 364 Leptopodia, 9
killiasii, Mycobilimbia, 217 Leptosphaeria, 389
Kirschsteiniella, 405 leptosphaeriae, Nectria, 247
Kriegeria, 117 Leptospora, 418
kriegerianum, Seimatosporium, 292 leptospora, Apostemidium, 222
Leptotrochila, 189, 190
laburni, Cucurbitaria, 414 Letendraea, 386
lacerum, Phacidium, 210 leucanthemi, Protomycopsis, 82
Lachnellula, 154 leucoloma, Octospora, 51
lactea, Hypocrea, 235 leucomelas, Paxina, 12
lacteus, Ascophanus, 64 leuconica, Ilyaloscypha, 158

"444
leucophaeus, Dasyscyphus, 151 lubrica, Leotia, 88
leucoplaca, Coniochaeta, 273 luctuosa, Leptosphaeria, 398
Leucoscypha, 32 lugubris, Anthostomella, 276
leucostigma, Orbilia, 167 lugubris, Ephelina, 179
Leucostoma, 302 lutea, Bolinia, 283
leucotricha, Leucoscypha, 32 luteonitens, Aleuria, 48
leucotricha, Podosphaera, 348 luteorubella, Orbilia, 167
leucoxantha, Discina, 13 luteovirescens, Rutstroemia, 101
levantica, Gloniopsis, 426 lycopodina, Leptosphaeria, 391
leveillei, Ascobolus, 58
Leveillula, 350 macrocystis, Anthracobia, 45
lichenicola, Rhagadostoma, 329 Macropodia 10
lichenicola, Seimatosporium, 290 macropus, Cyathipodia, 10
lichenoides, Hypocreopsis, 236 macrospora, Asteromassaria, 408
ligneola, Ceratostomella, 327 macrospora, Leptsophaeria, 392
ligni, Mollisia, 184 macrospora, Sordaria, 269
ligniaria, Coniochaeta, 273 macrosporidium, Leptosphaeria, 401
lignicola, Lasiosordaria, 270 macrosporum, Lophodermium, 201
lignorum, Trichoderma, 235 macrosporus, Protomyces, 81
lignyota, Karschia, 219 macrotrichia, Herpotrichia, 410
ligustri, Tympanis, 113 macrotrichus, Lasiobolus, 62
lilacina, Pezicula, 118 maculaeformis, Venturia, 376
lilacinum, Phaeohelotium, 118 maculans, Leptosphaeria, 392
Lilliputia, 339 maculare, Lophodermium, 201
limosa, Galactinia, 22 macularis, Sphaerotheca, 347
lindaviana, Rutstroemia, 103 maculiformis, Mycosphaerella, 363
lineare, Glonium, 424 magnusiana, Nectria, 246
linearis, Keissleriella, 388 Magnusiella, 77
lineolata, Mycosphaerella, 363
malacotricha, Coniochaeta, 273
lingam, Phoma, 392
mali, Cylindrocarpon, 244
Linospora, 325
malicorticis, Pezicula, 175
linteicola, Peziza, 20
Mamiana, 323
Lithographa, 216
littoralis, Leptosphaeria, 396 mamillana, Clypeosphaeria, 295
littoralis, Phaeosphaeria, 396 mammiformis, Rosellinia, 286
livida, Pezicula, 174 mammoidea, Nectria, 245
lividum, Thyridium, 418 marchantiae, Hymenoscyphus, 126
Lizonia, 368 margarita, Discinella, 135
lojkaeana, Lachnea, 35 marina, Leptosphaeria, 401
longipes, Xylosphaera, 284 maritima, Leptosphaeria, 402
lonicerae, Lasiobotrys, 379 Martinia, 95
Lopadostoma, 278 Massaria, 407
Lophidium, 381 Massariella, 293
Lophiella, 379 Massarina, 406
Lophionema, 380 mastoidea, Melomastia, 406
Lophiosphaeria, 379 maurilabra, Anthracobia, 45
Lophiostoma, 381 maximus, Cryptomyces, 205
Lophiotrema, 380 meandriformis, Choiromyces, 76
Lophium, 427 medicaginis, Pseudopeziza, 190
Lophodermium, 200 megalospora, Monilinia, 98
Loramyces, 328 megalospora, Monotospora, 424
megalospora, Sporormia, 412

"445
melaena, Melascypha, 69 Mitrophora, 4
melaleuca, Discina, 13 Mitrula, 89
melaleuca, Mollisia, 183 miyabeana, Glomerella, 259
melaleucoides, Tapesia, 182 Mniaecia, 214
melaleucum, Lophodermium, 201 modesta, Leptosphaeria, 393
melaloma, Anthracobia, 45 modesta, Nodulosphaeria, 394
Melanconiella, 315 modonia, Melanconis, 314
Melanconis, 313 Mollisia, 182
melanogramma, Spilopodia, 191 Mollisiella, 162
Melanomma, 409 Mollisina, 164
Melanopsamma, 330 mollisimus, Dasyscyphus, 151
Melanospora, 263 Monilia, 97, 267
melanosporum, Tuber, 74 Monilinia, 96
Melascypha, 69 Monotospora, 424
Melasmia, 199, 204 montagnei, Apiospora, 292
Melastiza, 43 monticola, Phaeohelotium, 118
melastoma, Plectania, 70 Morchella, 3
Meliola, 352 mori, Hysterographium, 426
Melittosporium, 208 moricola, Gibberella, 253
Melogramma, 318 moriformis, Bertia, 332
Melomastia, 406 mors-uvae, Sphaerotheca, 347
menziesii, Discinella, 135 mougeotii, Microsphaera, 349
merdaria, Hypocopra, 278 Muellerella, 370
Metacapnodium, 367 muhlenbeckii, Godronia, 140
michelii, Peziza, 22 multiforme, Hypoxylon, 280
michotii, Leptosphaeria, 399 multiseptata, Pleospora, 414
michotii, Paraphaeosphaeria, 399 multivalve, Phacidiostroma, 211
Microglossum, 88 murina, Leptopodia, 10
Micronectriella, 290 murina, Plicaria, 61
micropus, Peziza, 17 Mycobilimbia, 217
microscopica, Leptosphaeria, 398 Mycomelaspilea, 218
microscopicum, Microthyrium, 429 Mycosphaerella, 362
Microscypha, 163 Myiocopron, 428
Microsphaera, 348 Myriangium, 354
microspila, Sphaeria, 376 myriocarpa, Chaetosphaeria, 330
microsporus, Diehliomyces, 345 Myriosclerotinia, 95
microsporus, Ascophanus, 64 myrothecioides, Psilopezia, 55
Microstoma, 68 Myrothccium, 242
Microthyriella, 432 myrtilli, Gibberia, 377
Microthyrium, 429 myrtillina, Pezicula, 175
militaris, Cordyceps, 227 myrtillina, Pseudoplea, 357
millepunctata, Amphisphaeria, Mytilidion, 427
294 mytillinum, Lophium, 427
miniata, Lamprospora, 53
miniata, Melastiza, 44 Naemacyclus, 207
minima, Sporormia, 412 Naevia, 196
minor, Sclerotinia, 93 narcissi, Stromatinia, 92
minuta, Podospora, 269 nardi, Leptosphaeria, 396
minutissima, Naevia, 196 nardi, Phaeosphaeria, 396
minutissima, Peziza, 116 Naumovia, 421
misellus, Dasyscyphus, 150 necator, Uncinula, 348

"446
Nectria, 240 obducens, Teichospora, 416
Nectriella, 252 obscurum, Trichobelonium, 180
Nectriopsis, 251 obscurus, Saccobolus, 60
neglecta, Podospora, 270 obtusipila, Arachnopeziza, 166
neglectus, Saccobolus, 60 Ocellaria, 175
Neobulgaria, 108 ocellata, Keissleriella, 389
Neogyromitra, 7 ocellata, Ocellaria, 175
Neohenningsia, 251 ochraceo-pallida, Calonectria, 252
Neottiella, 42 ochraceus, Hypomyces, 239
nervisequia, Sclerotinia, 102 Octospora, 49
nervisequia, Spilopodia, 191 Oedocephalum, 16
Nesolechia, 216 ogilviensis, Leptosphaeria, 391
Neurospora, 267 oligotrichus, Lasiobolus, 62
nidulans, Emericella, 341 olivacea, Catinella, 177
nidulus, Dasyscyphus, 153 olivacea, Galactinia, 22
Niesslea, 329 olivaceum, Microglossum, 88
niessleana, Leptosphaeria, 399 Ombrophila, 107
niessleana, Meliola, 352 omphalodes, Pyronema, 62
niessleanum, Entodesmium, 400 onotica, Otidea, 26
niessleanum, Lophiostoma, 381 Onygena, 343
niesslii, Wettsteinina, 357 oocardii, Humaria, 55
nigerrima, Berlesiella, 370 Oomyces, 232
nigrans, Leptosphaeria, 396 operculata, Endoxylon, 304
nigrans, Phaeosphaeria, 397 Ophiobolus, 419
nigrella, Pseudoplectania, 69 Ophiochaeta, 421
nigricans, Claviceps, 227 Ophiocordyceps, 230
nigrificans, Ophiobolus, 393 ophioglossoides, Cordyceps, 229
nigritum, Geoglossum, 87 Ophionectria, 254
nigro-annulatum, Micro thyrium, Ophiostoma, 336
429 orae-maris, Leptosphaeria, 402
nigro-striata, Pirottaea, 181 Orbicula, 338
nitida, Anthrocobia, 46 Orbilia, 167
nitidulus, Hymenoscyphus, 124 Orbiliopsis, 192
nitschkei, Leptosphaeria, 392 oryzae, Phaeosphaeria, 390
Nitschkia, 333 osmundae, Microthyriella, 433
nivale, Fusarium, 290 Ostropa, 225
nivalis, Griphosphaeria, 290 Ostropella, 380
nivea, Arachnopeziza, 166 Otidea, 25
nivea, Leucoscypha, 33 Otthia, 415
niveum, Leucostoma, 302 ovalis, Pseudeurotium, 342
niveus, Dasyscyphus, 152 ovina, Lasiosphaeria, 272
niveus, Naemacyclus, 207 ovoidea, Zignoella, 331
nobilis, Pachydisca, 118 oxyacanthae, Podosphaera, 347
nodorum, Leptosphaeria, 398 oxyacanthae, Xylosphaera, 285
nodorum, Septoria, 398 oxycocci, Monilinia, 98
Nodulosphaeria, 389 oxyspora, Nesolechia, 216
notarisii, Clypeosphaeria, 296
nucula, Lophiotrema, 380 pachydermus, Protomyces, 81
nudipes, Dasyscyphus, 148
Pachydisca, 118
Nummularia, 283
Pachyella, 55
nummularium, Hypoxylon, 281
Pachyphloeus, 74

"447
padi, Dermea, 173 petiolorum, Rutstroemia, 101
padi, Taphrina, 80 petrakii, Synaptospora, 274
palearum, Dasyscyphus, 149 Pezicula, 173
pallida, Roesleria, 105 Peziza, 16
pallidula, Nectria, 243 peziza, Nectria, 248
pallidus, Cryptodiscus, 193 Pezizella, 130
paludosa, Mitrula, 89 pezizoides, Leptopodia, 10
palustriis, Ceriophora, 294 Phacidiella, 204
palustris, Mollisia, 185 phacidioides, Ceuthospora, 211
panamaensis, Martinia, 96 Phacidiostroma, 211
pannosa, Sphaerotheca, 347 Phacidiurn, 210
pantherina, Diaporthopsis, 312 Phaeangella, 138
Paradidymella, 292 Phaeangellina, 116
Paranectria, 254 Phaeocryptopus, 378
Paraphaeosphaeria, 399 Phaeohelotium, 118
parasitica, Endothia, 322 Phaeosphaeria, 390
parasitica, Melanospora, 263 Phaeospora, 371
parasitica, Phaeospora, 371 phaeosticta, Anthostomella, 277
pardalota, Diaporthe, 309 phaeostroma, Thaxteria, 411
parellaria, Leciographa, 218 Pharcidia, 371
paridis, Stromatinia, 92 Phialea, 128
parietina, Orbicula, 339 Phialina, 163
parile, Pezizella, 131 phillipsii, Daleomyces, 24
parmeliarum, Abrothallus, 220 phillipsii, Gyromitra, 24
parvula, Leptosphaeria, 398 Phoma, 325, 385, 391, 392
pascua, Sporormia, 412 Phomatospora, 320
Passeriniella, 403 Phomopsis, 308
patavina, Pustularia, 28 phragmitis, Helicosporium, 421
patella, Heterosphaeria, 142 Phyllachora, 261
Patellaria, 221 Phyllactinia, 351
Patellariopsis, 177
Paxina, 11 phyllogenus, Hymenoscyphus, 124
phyllophilus, Hymenoscyphus, 124
pedunculata, Podosordaria, 287 Physalospora, 257
pelagica, Leptosphaeria, 402 piceae, Belonium, 157
pelletieri, Thecotheus, 65 piceae, Ceratocystis, 336
penicillioides, Gliocladium, 251 piceae, Cistella, 157
perelegans, Haglundia, 181 piceae, Cucurbitaria, 415
perennans, Gloeosporium, 175 piceae, Lophodermium, 201
Perisporium, 422 piceae, Ophiostoma, 336
peristomialis, Actiniopsis, 254 piggotii, Homostegia, 359
perlata, Discina, 13 pinastri, Calothyrium, 431
perniciosa, Diaporthe, 309 pinastri, Lophodermium, 201
Peroneutypa, 303 pinea, Brunchorstia, 139
perparvula, Cistella, 156 pinea, Nectria, 246
Perrotia, 154 pineti, Pseudohelotium, 115
persoonii, Barlaeina, 29 pini, Coccophacidium, 207
persoonii, Stamnaria, 127 pini, Therrya, 206
pertusa, Trematosphaeria, 404 pinicola, Bothrodiscus, 139
Pestalotia, 387 pinicola, Pseudographis, 143
petersii, Peziza, 21 pinodes, Didymella, 385
petiolaris, Pyrenopeziza, 187 pirina, Venturia, 375

"448
pirinum, Fusicladium, 375 praemorsum, Lophiotrema, 380
Pirobasidium 109 praetermissa, Leptosphaeria, 393
Pirottaea, 181 praetervisa, Peziza, 21
Pithya, 67 Pragmopora, 140, 170
Pithyella, 162 prasinula, Corynella, 110
pithyophila, Cucurbitaria, 415 prasinula, Dendrostilbella, 110
Placographa, 216 prasinus, Trichodiscus, 146
Plagiostoma, 324 primulae, Mycosphaerella, 364
planiuscula, Leptosphaeria, 393 Propolis, 194
platanoides, Calospora, 316 Prosthecium, 314
platanoides, Calosporella, 316 proteana, Peziza, 24
platanoides, Pseudovalsa, 316 Protocrea, 236
Platychora, 361 Protomyces, 81
platyspora, Balsamia, 74 Protomycopsis, 82
Plectania, 69 protracta, Microstoma, 68
Pleomassaria, 417 pruinosa, Polydesmia, 144
Pleonectria, 256 prunastri, Dermea, 173
Pleospora, 413 prunastri, Eutypella, 300
Pleurage, 269 prunastri, Uncinula, 348
Plicaria, 14 pruni, Taphrina, 79
Ploettnera, 194 Pseudeurotium, 341
ploettneriana, Gyrocratera, 72 Pseudobalsamia, 345
Plowrightia, 362 pseudofarinacea, Lachnellula, 155
plumbina, Lahmia, 219 Pseudographis, 143
Pocillum, 114 Pseudohelotium, 115
Podonectria, 420 Pseudomassaria, 291
Podophacidium, 193 Pseudombrophila, 37
Podosordaria, 287 Pseudonectria, 240
Podosphaera, 347 Pseudopeziza, 189
Podospora, 269 Pseudophacidium, 209
Podostroma, 234 pseudoplatani, Dermatea, 175
Polydesmia, 144 Pseudoplea, 357
polygoni, Erysiphe, 349 Pseudoplectania, 69
polymorpha, Xylosphaera, 284 Pseudorhizina, 7
polyporina, Nectria, 247 Pseudorhytisma, 189
polyspermum, Camarops, 282 Pseudosphaeria, 358
Polystigma, 259 Pseudotis, 27
Polythrincium, 364
polytrichi, Lamprospora, 54 pseudotsugae, Rhabdocline, 211
polytrichi, Peziza, 43 Pseudovalsa, 318
polytrichi, Pezizella, 133 Psilachnum, 155
pomi, Spilocaea, 375 Psilopezia, 55
pomiformis, Melanopsamma, 330 ptarmicae, Schizothyrioma, 212
pontiformis, Leptosphaeria, 399 pteridicola, Phialea, 129
populina, Taphrina, 77 pteridinum, Melittosporium, 208
porcina, Fimaria, 52 Pteridiospora, 407
Poronia, 286 pteridis, Cryptomycina, 210
Potebniamyces, 204 pteridis, Dasyscyphus, 150
potentillae, Coleroa, 377 Ptychoverpa, 2
potentillae, Taphrina, 79 puberula, Phialina, 163
praecox, Tricharia, 35 puberulum, Tuber, 75
pudibundus, Dasyscyphus, 149

"449
pulchella, Calosphaeria, 305 rackii, Strossmayeria, 116
pulchra, Selinia, 256 radiata, Stictis, 224
pulicare, Hysterium, 425 radiculata, Sowerbyella, 29
pulicaris, Gibberella, 253 ralfsii, Nectria, 242
pulla, Bulgariella, 111 ramealis, Mollisia, 185
pulveracea, Coniochaeta, 273 ranunculi, Fabraea, 190
pulverulenta, Karschia, 220 rapulum, Stromatinia, 92
pulverulentus, Dasyscyphus, 152 Rebentischia, 402
pulvinata, Ilypocrea, 235 recutita, Mycosphaerella, 364
Pulvinula, 54 Rehmiellopsis, 366
pulvis-pyrius, Melanomma, 409 remuliformis, Boydia, 293
punctata, Poronia, 286 reniformis, Cephalotheca, 343
punctiformis, Mycosphaerella, repanda, Peziza, 17
363 repanda, Pseudopeziza, 190
punctiformis, Peziza, 164 repandus, Hymenoscyphus, 123
punctoidea, Pezizella, 133 repens, Aspergillus, 340
punctulata, Anthostomella, 276 resinae, Biatorella, 215
punicea, Nectria, 244 resinae, Pycnidiella, 215
pupula, Massaria, 408 retincola, Tapesia, 182
pura, Neobulgaria, 108 Retinocyclus, 112, 215
purpurascens, Duebenia, 197 Rhabdocline, 211
purpurea, Gephalotheca, 343 Rhagadostoma, 329
purpurea, Claviceps, 227 Rhamphoria, 327
purpurea, Leptosphaeria, 400 rhenana, Aleuria, 48
purpureum, Melanomma, 401 Rhizina, 13
purtonii, Nectria, 247 rhizophila, Zopfia, 338
pusilla, Mitrula, 89 rhizophora, Taphrina, 78
pusilla, Niesslia, 329 rhododendri, Chaetapiospora, 289
pustula, Plagiostoma, 325 rhododendri, Lophodermium, 201
pustula, Phoma, 325 rhododendri, Monilinia, 98
Pustularia, 27 rhodoleuca, Propolis, 194
pustulata, Peziza, 19 rhodoleucus, Hymenoscyphus, 124
Pycnidiella, 215 rhodostoma, Karstenula, 418
Pycnidiophora, 343 Rhopographis, 360
pygmaeum, Tichothecium, 372 Rhyparobius, 65
pygmaeus, Dasyscyphus, 148 Rhytisma, 199
Pyrenopeziza, 186 rhytismoides, Isothea, 260
Pyrenophora, 414 ribesia, Dothidea, 362
pyriformis, Rhamphoria, 327 ribesia, Plowrightia, 362
Pyronema, 62 ribis, Drepanopeziza, 189
ribis, Gloeosporidiella, 189
ribis, Godronia, 140
quadrisporus, Saccobolus, 60 rimosa, Scirrhia, 360
Quaternaria, 300 rishbethii, Nectria, 248
quaternata, Quaternaria, 300 Robergea, 225
queletii, Helvella, 12 robergei, Nectriella, 252
quercina, Diatrypella, 299 robergei, Unguiculella, 162
quercinum, Colpoma, 199 robertiani, Stigmatea, 378
quercuum, Botryosphaeria, robustior, Hymenoscyphus, 125
373 Roesleria, 104
quercus, Cyrtidula, 356 rosae, Diplocarpon, 189
quercus, Dermatina, 356

"450
rosae, Gibbera, 375 sambuci, Dothidea, 359
rosae, Tapesia, 182 saniosa, Peziza, 23
rosae, Venturia, 375 sarcoides, Ascocoryne, 109
rosea, Pustularia, 28 sarcoides, Coryne, 109
rosellus, Hypomyces, 239 sarcoides, Pirobasidium, 109
Rosellinia, 285 Sarcoscypha, 67
roumegueri, Peziza, 50 Sarcosoma, 69
rousseauana, Fabraea, 190 Sarcosphaera, 15
rousseliana, Pseudonectria, 240 sarraziniana, Orbilia, 168
rubella, Hyalinia, 169 sarrazinii, Galactinia, 21
rubella, Leptospora, 419 Sartorya, 341
rubellus, Ophiobolus, 419 Saturnomyces, 343
ruber, Lasiobolus, 62 scatigena,'Coniochaeta, 273
rubi, Pezicula, 174 Schizothyrioma, 212
rubicunda, Cudoniella, 121 Schizoxylon, 224
rubiginosum, Hypoxylon, 280 scirpi, Coniothyrium, 399
rubra, Cheilymenia, 42 scirpicola, Sclerotinia, 95
rubra, Habrostictis, 176 scirpina, Leptosphaeria, 402
rubricosa, Humaria, 50 scirpinum, Hypoderma, 202
rubronotata, Thyridaria, 410 Scirrhia, 360
rubrum, Polystigma, 260
scitula, Leptosphaeria, 400
nide, Entodesmium, 390
scitulum, Melanomma, 400
rufa, Hypocrea, 235
rufo-olivacea, Velutarina, 136 Scleroderris, 139
rufula, Lilliputia, 339 Sclerotinia, 92
rufum, Agyrium, 215 sclerotidrum, Mitrula, 89
rufum, Tuber, 75 sclerotiorum, Sclerotinia, 92
rumicis, Hymenoscyphus, 125 sclerotipus, Mitrula, 89
rumicis, Septatium, 125 Scoleconectria, 254
rumicis, Venturia, 376 scoparia, Pezicula, 174
rusci, Leptosphaeria, 399 scoriadea, Pteridiospora, 407
rusci, Paraphaeosphaeria, 399 scotica, Melastiza, 44
ruscigena, Haplosporella, 399 scrupulosa, Unguicularia, 161
rutilans, Neotiella, 42 scutellata, Scutellinia, 38
rutilum, Hypoxylon, 279 Scutellinia, 38
Rutstroemia, 100 Scutula, 217
scutula, Hymenoscyphus, 122
seaveri, Chloroscypha, 117
sabinae, Chloroscypha, 117 sedi, Euryachora, 361
sabinae, Eutryblidiella, 220 Seimatosporium, 290, 292
sabuletorum, Metasphaeria, 396 Selinia, 256
saccharinus, Ascophanus, 64 semi-immersum, Hypoxylon, 281
Saccobolus, 59 semilibera, Mitrophora, 4
Saccothecium, 365 senecionis, Pirottaea, 181
sadebeckii, Taphrina, 80 separabilis, Hymenoscyphus, 125
salicella, Cryptodiaporthe, 310 Sepedonium, 240
salicina, Cryptodiaporthe, 310 sepiatra, Peziza, 19
salicinum, Capnodium, 367 sepincola, Saccothecium, 365
salicinum, Rhytisma, 199 sepincolaeformis, Apiosporella, 291
salicis, Uncinula, 348 sepincolaeformis, Pseudomassaria,
salicis, Valsella, 303 291
salmonicolor, Ascophanus, 64 Septatium, 125

"451
Septogloeum, 361 spiraeicola, Verpatinia, 96
Septoria, 398 spirale, Chaetomium, 265
Sepultaria, 31 spodiaea, Melanconiella, 315
serica, Sclerotinia, 93 Sporomega, 200
serpens, Hypoxylon, 280 Sporormia, 411
setosa, Podospora, 269 Stamnaria, 127
sexdecemspora, Capronia, 370 starbaeckii, Geoglossum, 87
sexdecemsporus, Rhyparobius, 65 Stegia, 196
Seynesiella, 430 stellata, Stictis, 223
silenes-acaulis, Leptosphaeria, 394 stellulata, Eutypella, 300
silenes-acaulis, Phaeosphaeria, 394 stenosperma, Scutellinia, 39
Sillia, 319 Stephensia, 73
Simoninus, 285 stercorarea, Anixiopsis, 342
sinopica, Nectria, 243 stercorareus, Ascobolus, 57
siparia, Encoelia, 139 stercorea, Cheilymenia, 42
siparia, Pleomassaria, 417 stercoreus, Thelebolus, 70
sitophila, Monilia, 267 sterigmatizans, Peziza, 18
sitophila, Neurospora, 267 stevensii, Leptopodia, 10
solani, Nectria, 243 Stictis, 223
sorbi, Eutypella, 300 stictoideus, Ascobolus, 58
Sordaria, 268 stigma, Diatrype, 299
Sowerbyella, 28 Stigmatea, 378
sowerbyi, Leptosphaeria, 394 Stigmatomassaria, 408
sowerbyi, Phaeosphaeria, 394 stilbostoma, Melanconis, 313
Spadicoides, 273 strobilicola, Tapesia, 182
sparassoides, Underwoodia, 24 Stromatinia, 92
spartii, Cucurbitaria, 415 Strossmayeria, 116
spartinae, Didymosphaeria, 403 subalpina, Keissleriella, 389
Spathularia, 90 subcarnea, Orbiliopsis, 192
spermoides, Lasiosphaeria, 271 subfuscus, Ascophanus, 64
sphaeralis, Laquearia, 213 subhyalina, Phialea, 130
Sphaeridiobolus, 61 sublenticular, Helotium, 123
sphaerioides, Byssolophis, 382 subpallida, Calycella, 119
Sphaeroderma, 264 subtilis, Pezizella, 133
Sphaeronaemella, 335 subtilissima, Lachnellula, 155
Sphaeropezia, 212 succosa, Peziza, 22
Sphaerospora, 38 suecica, Lachnellula, 154
sphaerospora, Pseudorhizina, 7 suecicus, Abrothallus, 220
Sphaerosporella, 39 suffulta, Neohenningsia/ 251
Sphaerostilbe, 250 suffusa, Cryptospora, 317
Sphaerotheca, 346 sulcata, Helvella, 8
Sphaerulina, 365 sulcata, Paxina, 12
Sphagnicola, 107 sulcata, Sclerotinia, 94
sphecocephala, Cordyceps, 228 sulcigena, Ilypodermella, 203
Spilocaea, 375 sulfurea, Cephalotheca, 343
Spilopodia, 191 sulfureus, Dasyscyphus, 151
Spilosticta, 375 sulfurina, Calycella, 119
spiniferum, Melogramma, 318 sulphuratum, Helotium, 126
spinosa, Eutypa, 298 sulphurea, Discodiaporthe, 313
spinulosa, Chromocrea, 235 sulphurellus, Dasyscyphus, 149
spiraeae, Otthia, 415 sumneriana, Sepultaria, 31

"452
sydowiana, Rutstroemia, 101 Torrubiella, 230
Sydowiella, 320 tortilis, Erysiphe, 350
sylvatica, Phyllachora, 261 tosquinetii, Taphrina, 78
Synaptospora, 274 tosta, Paradidymella, 292
Systremma, 361 trachycarpa, Plicaria, 15
trechispora, Scutellinia, 39
Talaromyces, 338 Trematosphaeria, 404
taleola, Caudospora, 312 Triblidium, 142
Tapesia, 182 Tricharia, 34
Taphridium, 82 Trichobelonium, 180
Taphrina, 77 trichodea, Peziza, 160
tartaricola, Synaptospora, 274 Trichodelitschia, 412
tassiana, Mycosphaerella, 364 Trichoderma, 235
taurica, Leveillula, 350 Trichodiscus, 146
taxi, Anthostomella, 277 Trichoglossum, 85
taxi, Sphaerulina, 365 Trichometasphaeria, 388
taxifolia, Cytospora, 365 Trichonectria, 255
tectoria, Peziza, 20 Trichophaea, 35
Teichospora, 416 Trichoscyphella, 155
temulenta, Gloeotinia, 98 Trichosphaeria, 330
tenuispora, Podonectria, 420 trichostoma, Pyrenophora, 414
tenuis, Sepultaria, 32 trifidum, Verticicladium, 33
Terfezia, 76 trifolii, Cymadothea, 364
trifolii, Mycosphaerella, 364
terrestris, Allescheria, 341
trifolii, Pseudopeziza, 190
testaceus, Ascophanus, 64
trifolii, Pseudoplea, 358
tetrasperma, Gelasinospora, 268
trifoliorum, Sclerotinia, 93
tetrasperma, Neurospora, 267
tripolii, Laetinaevia, 197
tetraspora, Didymascella, 209
triseptatum, Cladotrichum, 411
tetraspora, Dothidea, 358
tristis, Calyculosphaeria, 333
tetraspora, Octospora, 50
tritici, Leptosphaeria, 398
tetrasporum, Phacidium, 208
Trochila, 195
tetrasporum, Trichoglossum, 86
truncorum, Vibrissea, 222
Thamnomyces, 285
Tuber, 74
Thaxteria, 410
Tubercularia, 241
Thecotheus, 65
tuberculata, Cordyceps, 229
theioleuca, Fimaria, 52
tuberosa, Sclerotinia, 93
thelebola, Melanconis, 313
Tubeufia, 421
theleboloides, Cheilymenia, 42
Thelebolus, 70
tubiformis, Gnomoniella, 325
thelena, Rosellinia, 286
tulasneana, Apiocrea, 240
Therrya, 206
tulasnei, Dermea, 173
^Jiielavia, 337
tulasnei, Hydnotrya, 72
thujina, Didymascella, 209
tulasnei, Podosordaria, 287
Thyridaria, 409
turbinata, Phialea, 129
Thyridium, 418
turficola, Coryne, 110
Thyronectria, 255
turgidum, Lopadostoma, 278
Tichothecium, 372
Tympanis, 112
tiliacea, Encoelia, 139
Tympanopsis, 334
tiliae, Hercospora, 314
typhae, Leptosphaeria, 395
tithymalina, Laetinaevia, 197
typhae, Rebentischia, 403
tomicoides, Entosordaria, 276
typharum, Leptosphaeria, 395

"453
typharum, Phaeosphaeria, 395 variecolor, Aspergillus, 341
typhina, Bpichloe, 231 variecolor, Emericella, 341
typhinum, Lophodermium, 201 vasaensis, Hymenoscyphus, 126
vectis, Leptosphaeria, 400
uberiformis, Godronia, 140 Velutarina, 136
ulicis, Phaeangella, 138 velutina, Coniochaeta, 273
ulmariae, Phialina, 164 veneta, Elsinoe, 355
ulmi, Ceratocystis, 337 venosa, Disciotis, 5
ulmi, Euryachora, 361 ventosa, Mollisia, 183
ulmi, Platychora, 361 Venturia, 375
ulmi, Septogloeum, 361 vepris, Apioporthe, 310
ulmi, Taphrina, 80 Vermicularia, 259
verna, Ombrophila, 122
umbelliferarum, Taphridium, 82
vernum, Helotium, 122
umbilicata, Pachydisca, 118
vernus, Hymenoscyphus, 122
umbonata, Pseudovalsa, 319
veronicae, Asterina, 430
umbrina, Aleuria, 19
Verpa, 4
umbrina, Amphisphaeria, 294 Verpatinia, 96
umbrina, Cryptosporella, 318
umbrina, Otidea, 26 versicolor, Melittosporium, 208
umbrina, Peziza, 19 versicolor, Propolis, 194
Uncinula, 348 versiforme, Chlorosplenium, 134
Underwoodia, 24 Verticicladium, 33
undulata, Rhizina, 13 Verticillium, 239, 249
Unguicularia, 160 vesiculosa, Peziza, 19
Unguiculariopsis, 162 vestergrenii, Taphrina, 80
Unguiculella, 161 vestita, Fenestella, 416
unica, Robergea, 226 veuillotiana, Nectria, 245
unicaudata, Rebentischia, 403 vexata, Lejosphaerella, 292
Urceolella, 160 Vialaea, 293
urceoliformis, Godronia, 140 vibratilis, Massariella, 294
urceolus, Godronia, 140 Vibrissea, 221
Lirnula, 69 villosa, Cyathipodia, 11
urnula, Monilinia, 97 vinosa, Orbilia, 168
urticae, Cylindrocolla, 192 vinosus, Ascobolus, 52
ustorum, Sordaria, 277 violacea, Nectriopsis, 252
Ustulina, 282 violacea, Ombrophila, 107
violacea, Peziza, 20
vaccinii, Gibbera, 377 virgineus, Dasyscyphus, 148
vaccinii, Sphaeropezia, 213 virgultorum, Anisogramma, 322
vagabundum, Lophiotrema, 380 virgultorum, Helotium, 123
vagans, Melogramma, 318 virgultorum, Hypoderma, 202
vaguluin, Lophodermium, 201 viride, Microglossum, 88
vagum, Aulographum, 431 viridescens, Nectria, 247
vahliana, Sclerotinia, 95 viridifusca, Ciboria, 100
Valsa, 301 viridis, Ascobolus, 57
Valsaria, 315 viridis, Byssonectria, 238
Valsella, 303 visci, Gibberidea, 422
vitalbae, Broomella, 387
valvata, Hysteropezizella, 188 vitellina, Cheilymenia, 41
vaporaria, Morchella, 4 vitellinus, Hymenoscyphus, 122
varia, Peziza, 17 vivida, Neotiella, 43
varians, Celidium, 423

"454
vogesiaca, Pseudoplectania, 69 woolhopeia, Trichophaea, 35
Volutella, 240 woolhopensis, Ascozonus, 66
vulgare, Perisporium, 422 wrightii, Peziza, 51
vulgaris, Balsamia, 74
vulgaris, Gnomonia, 324 xantha, Keissleria, 387
vulgaris, Pezizella, 131 xanthomelum, Podophacidium, 193
vulgaris, Pithya, 67 xanthostigma, Orbilia, 167
vulgaris, Tubercularia, 241 Xylaria, 284
Xylosphaera, 284
wahlenbergii, Calosphaeria, 305 Xylosphaeria, 418
walteri, Trichoglossum, 86 xylostei, Amphisphaerella, 277
Westerdykella, 343
Wettsteinina, 357 zamiae, Melanospora, 263
willkommii, Lachnellula, 155 zeae, Gibberella, 253
winteri, Delitschia, 413 Zignoella, 331
winteri, Didymosphaeria, 386 Zopfia, 338

"455
P L A T E I.
A. Leptopodia elastica. B. L. atra. c. Cyathipodia macropus. r>. Paxina
leucomelas. E. P. acetabulum. F. Rhizina undulata. G. Pustularia catinus.
H. P. rosea. J. P. cupularis.
P L A T E II.
A. Disciotis venosa. B. Discina perlata. c. Plicaria leiocarpa. D.P.trachy-
carpa and spore of var. maricata. E. Barlaeina amethystina. F. Sarcosphaera
exima. G. Peziza brunneoalra. N. P. atrovinosa. J. P. badiofusca.
P L A T E III.
A. Peziza ammophila. B. P. micropus. c. P. cerea. D. P. repanda. E. P.
varia. F. P. echinospora. a. P. vesiculosa, it. P. saniosa. .1. P. violacea.
P L A T E IV.
A. Peziza emileia. B. P. praetervisa. c. P. petersii. D. P. badia. E. P.
ionella. F. P. michelii. G. P. succosa. H. Sowrebyella radiculata. J. Pseudotis
apophysata. K. Psilopezia babingtonii. L. Peziz ajproteana var. sparassoides.
P L A T E V.
A. Otidea onotica. B. O. alutacea. c. O. bujonia,. D. O. concinna. E. Sepul-
taria sumneriana. F. S. arenosa. c. S. tenuis. 11. Plectania melastoma.
•J. Desmazierella. acicola.
A. Tricharia gilva. B. T.cretea. c. Humaria hemisphaerica. D. Trichophaea hemisphaerioides.
E. T. woolhopeia. F. T. bicuspis. c. Scutellinia scutellata. H. S. trechispora and spore of S.
asperior. I. Sphaerosporella brunnea. j. Cheilymenia crucipila. K. C. vitellina. L. Melastiza
chateri. M. Cheilymenia theleboloides. N. C. coprinaria. o. Anthracobia macrocystis. p. A.
melaloma. Q. A. maurilabra. B. Neottiella hetieri. s. Coprobia granulata. T. Pulvinula con-
stellatio. u. Lamprospora crouani. v. Octospora carbonigena. w. Neottiella rutilans and spore
of N. vivida. x. Octospora humosa. Y. Ascospore of Lamprospora crec'hquerauilii.
P L A T E VII.
A. Aleuria aurantia. B. A. luteonitens. c. Sarcoscypha coccinea. n. Micro-
stoma protra.cta. E. Geopyxis carbonaria. F. Pseudombrophila deerala.
G. Pithy a cupressina. H. Ascobolus furfuraceus. i. A. carbonarius. s.A.viri-
dis. K. A. denudatus. L. A. albidus.
PLATE VIII.
A. Lasiobolus ciliatus. u. Saccobolus obscurus. c. Dasyobolus immersus.
D. Saccobolus glaber. E. Ascobolus crosslandii. F. Iodophanus carneus.
c. Ascophanus microsporus. H. Sphaeridiobolus albojuscus. I. Fimaria
hepatica. J. Ascophanus argenteus. K. Boudiera areolata.. L. Pyronema
omphalodes. M. Thecotheus pelletieri. N. Hhyparobius cookei. o. H. sexdecem-
sporus. p. Octospora carneola. Q. Octospora leucoloma.
P L A T E IX.
A. Tuber excavatum. B. T. rufum. c. Stephensia bombycina. i>. Genea
hispidula. E. Elaphomyces granulatus. F. Balsamia platyspora. c. Hydnobo-
lites cerebri]ormis. H. Hydnotrya tulasnei. I. Pachyphloeus citrinus. j. Choi-
romyces meandriformis. K. Gyrocratera, ploettneriana. L. Diehliomyces micro-
sporus.
P L A T E X.
A. Taphrina deformans, B. T. populina. c. T. potentillae. D. T.pruni.
E. T. amentorum. F. T. johansonii. G. Mitrula paludosa. H. M. abietis.
I.Spathularia flavida. J. Leotia lubrica. K. Microglossum viride. L. Cudonia
circinans. M. Microglossum olivaceum.
P L A T E XI.
A. Taphrina. cerasi. B. T. betulae. c. T. tosquinetii. D. T. betulina. E. T.
carpini. F. Ascocorticium anomalum. G. Taphrina athyrii.
PLATE XII.
A. Taphridium umbelliferarum. B. Protomyces macrosporus. c. P. pachy-
dermia. D. P. inundatus. E. Protomycopsis leucanthemi. F. Taphrina vester-
grenii. G. T. filicina.
PLATE XIII.
A. Sclerotinia tuberosa. B. S. trifoliorum. c. S. sclerotiorum. i>. S. sulcata.
E. S. curreyana. F. Ciboria aschersoniana. G. Sclerotinia candolleana.
H. Verpatinia spiraeicola. i. Ciboria batschiana. J. Monilinia baccarum.
K. Rutstroemia firma. L. Ciboria amentacea on Alnus. M. Rutstroemia sydo-
wiana. N. R. luteovirescens. o. R. lindaviana. P. Gloeotinia temulenta.
Q. Rutstroemia conjormata. R. Cudoniella clavus. s. Rutstroemia echinata.
P L A T E XIV.
A. Ciboria viridifusca. B. Ciboriopsis bramleyi. c. Martinia panamaensis.
I). Rutstroemia hercynica. E. Coprotinia cuniculi. F. Monilinia johnsonii.
G. Hymenoscyphus repandus. H. II. robustius. J. Cudoniella juncisedum.
K. C. rubicunda. L. Pezizella eriophori. M. P. polytrichi. N. Monilinia
mespili. o. Phaeohelotium monticola. p. Rutstroemia americana. Q. Ombro-
phila ambigua. n. Monilinia urnula. s. Hymenoscyphus rumicis. T. Ciboria
juncorum. v. ,,Helotium" fulvum.
P L A T E XV.
A. Coryne sa.rcoi.des a n d s p o r e of C. cylichnium. B. Neobulgaria. pura.
c. N. foliacea. n. Ombrophila violacea. E. Corynella prasinula. F. C. atrovi-
reris. G. Bulgariella pulla. H. Bulgaria inquinans. I. Biatorella resinae.
J. Phaeangellina empetri. K. Sphaeropezia empetri. L. Agyrium rufum.
M. Discinella boudieri. N. D. margarita. o. Mniaecia jungermanniae.
P L A T E XVI.
A. Calycella citrina. B. C.sulfurina. c. Stamnaria. persoonii. D. Hymenos-
cyphus rliodoleucus. E. H. herbarum. F. II. scutula. G. II. calyculus. H. II.
vernus. I. H. caudatus. J. H. epiphyllus. K. H. phyllophilus. L. II. phylloge-
nus. M. II. fructigenus.
P L A T E XVII.
A. Phialea cyathoidea on Sonchus. B. P. egenula. c. P. pteridicola. D. P.
lurbinata. E. P. su.bhyali.na. F. Cyathicula coronata. G. Belonioscypha
culmicola. u. Pezizella alniella. P. vulgaris. j. P.discreta. K. P. punctoidea.
L. P. chrysostigma. M. P. amenti.
PLATE XVIII.
A. Godronia ribis. B. Encoelia furfuracea. c. Encoeliopsis johnstoni.
r>. Velutarina rufo-olivacea. E. Cenangium jerruginosum. F. Chlorosplenium
aeruginascens. G. C. versijorme. H. Heterosphaeria patella. J. Phaeangella
ulicis. K. Phialea incertella.
PLATE XIX.
A. Dasyscyphus pygmaeus. B. D. nudipes. c. D. bicolor var. rubi. D. D.
pulverulentus. E. D. mollissimus. F. D. cerinus. G. I), palearum. H. D. fus-
cescens. I. D. barbatus. j. D. nidulus. K. D. clandeslinus. L. Lachnellula
hahniana. M. Dasyscyphus albotestaceus.
PLATE XX.
A. Unguicularia cirrhata.. B. Hyaloscypha demaliicola. c. Unguiculella
eurotioides. D. Urceolella crispula. E. Arachnopeziza aurata. F. Phialina
ulmaria. G. Microscypha grisella. H. Phialina puberula. I. Eriopeziza caesia.
.1. Hyalotricha corticola. K. Cistella. perparvula.
PLATE XXI.
A. Orbilia xanthostigma. B. O. luteorubella. c. O. curvatispora. n. Hyalinia
rubella. E. Orbilia sarraziniana. F. O. auricolor. G. Pseudohelotium pineti.
H. Dasyscyphus acuum. s. Pocillumcesatii. K. Diplocarpabloxami. L. Tricho-
discus prasinus. All xlO unless otherwise stated.
PLATE XXII.
A. Vibrissea truncorum. B. Apostemidium leptospora. c. Stictis stellata.
i). Schizoxylon berkeleyanum. E. Dermea prunastri. r. Pezicula livida. G. P.
corticola. n.P.rubi. I. P. scoparia. J. Ocellaria ocellata. K. Ploeltnera exigua.
I.. Habrostictis rubra. M. Cryptodiscus pallidus.
PLATE XXIII.
A. Haglundia perelegans. b. Mollisia cinerea. c. Belonopsis excelsior.
D. Mollisia ramealis. E. M. palustris. F. Hi/steropezizella valvata. G. Belo-
nopsis filispora. H. Mollisia chionea. I. Hysteropezizella exigua. J. Mollisia
Melaleuca. K. M. ligni. L. M. cornea. M. M. discolor var. longispora.
PLATE XXIV.
A. Mollisia caespiticia. B. M. ventosa. c. Graddonia coracina. D. Tapesia,
strobilicola. E. Pyrenopeziza. arctii. F. P. joliicola. G. Pirottaea brevipila.
H. Trichobelonium obscurum. .J. Spilopdia. nervisequia. K. Gorgoniceps
aridula. L. Hymenoscyphus marchantiae. M. Catinella olivacea.
P L A T E XXV.
A. Naevia minutissima. B. Laetinaevia tithymalina. c. Duebenia purpuras-
cens. D. Calloria fusarioides. E. Pseudopeziza trifolii. F. Trochila craterium.
G. Drepanopeziza ribis. H. Trochila ilicina. j. Fabraea ranunculi. K. Orbili-
opsis subcarnea.
PLATE XXVI.
A. Cordyceps militaris. l). C. gracilis, c. C. forquignoni. v. C. ophioglossoi-
des. E. C. capitata. F. C. sphecocephala. o. C. tuberculata. H. Ophiocordyceps
clavulata. i. Torrubiella albolanata. J. Claviceps purpurea, K. Epichloe
typhina. L. Oomyces carneoalbus. M. Barya aurantiaca. N. Hypocrea rufa.
p. Creopus gelatinosus.
PLATE XXVII.
A. Hypocreopsis lichenoides. B. Protocrea. farinosa. c. Podostroma aluta-
ceum. D. Selinia pulchra. E. Polystigma rubrum. F. Thyronectria berolinen-
sis. G. Trichonectria hirta. H. Paranectria affinis. i. Gibberella cyanogena.
J. Scoleconectria cucurbitula. K. Actinopsis peristomialis. L. Neohenningsia
suffulta. M. Calonectria ochraceopallida.
PLATE XXVIII.
A. Nectria sinopica. B. A. punicea var ilicis. c. A. peziza. i>. A. coccinea.
v.. N. episphaeria. v.N.pinea. G. N. fuckeliana. n. A. aquijolii. j. A. coryli.
PLATE XXIX.
A. Nectria galligena. 11. N. ditissima. c. N. mammoidea. D. N. magnusiana.
E. N. viridescens. F. N. brassicae. G. N. leptosphaeriae. H. N. inventa. J. N.
hederae. K. N. aureola. L. N• polyporina.
mm
•Sllisw

PLATE XXX.
A. Nectria ralfsii. n. JV. solani. c. iV. pallidula. D. iV. keithii. E. N. desma-
zierii. F. N. arenula. a. N. ellisii. H. N. funicola. .]. N. flavoviridis. K. N.
rishbethii. L. N. citrino-aurantia. M. N. purtonii.
PLATE XXXI.
A. Apiocrea chrysosperma witli a Sepedonium chlamydospore. B. Hypomy-
ces aurantius. c. Nectriopsis aureonitens. D. Nectriella, robergei. E. Nectria
lecanodes. F. A. cinnabarina. G. A. veuillotiana. H. Sphaerostilbe aurantiaca.
I. Hypomyces ochraceus with a patch of Sphaeroderma episphaeria on the
upper right. J. Byssonectria viridis. k. Letendraea helminthicola. l . Hypo-
nectria buxi. M. Pseudonectria rousseliana.
PLATE XXXII.
A. Hypoxylon fragiforme. B. H. rubiginosum. c. / / . fuscum. D. II. multi-
forme. E. H. nummularium. F. H. serpens. G. II. semi-immersum. H. H. ar-
gillaceurn. I. liolinia lutea. J. Daldiriia eoneentrica. K. Camarops polysper-
mum. L. Nummularia discreta. M. Ustulina deusta.
PLATE XXXIII.
A. Valsa ambiens. B. Leucostoma nivea. c. Valsella salicis. D. Ditopella
ditopa. E. Melanconis stilbostoma. F. Melanconiella spodiaea. G. Cryplospora
sujjusa. H. Calosporella innesii. j. Prosthecium auctum. K. Cryplosporella
hypodermia.
PLATE XXXIV.
A. Caudospora taleola. is. Cryptodiaporthe salicella. C. Diaporthe eres on
Ulmus. D. Diaporthopsis angelicae. F,. Discodiaporthe sulphurea. F. Apio-
porthe vepris. G. Hercospora tiliae. H. Sillia ferruginea. J. Endothia parasi-
tica.
PLATE XXXV.
A. Entosordaria tomicoides. b. Hypocopra merdaria. c. Lasiosordaria
coprophila. i>. Amphisphaerella xylostei. E. Clypeosphaeria mamillana.
F. Melanopsamma pomiformis. G. Anisostomula. cookeana. H. Gripho-
sphaera, corticola. I. Melanospora. chionea. J. Melanospora parasitica, v.. Co-
ronophora gregaria. L. Niesslia pusilla.
PLATE XXXVI.
A. Potebniamyces discolor. B. Elsinoe veneta. c. Myriangium duriaei.
I). Echidnodes aulographoides. E. Chaetothyrium babingtonii. F. Microthy-
rium nigroannulatum. G. Asterina veronicac. H. Calothyrium pinastri.
PLATE XXXVII.
A. Discothecium gemmiferum. B. Pharcidia epicymatia. c. Rehmiellopsis
abietis. D. Ceriophora paluslris. E. Tichothecium pygmaeum. F. Phaeospora
parasitica. G. Gibellina cerealis. H. Valsaria insitiva. t. Lizonia emperigonia.
J. Trichodelitschia bisporula. K. Apiorhynchostoma. apiculata. L. Sporormia.
intermedia.
PLATE XXXVIII.
A. Lasiobotrys lonicerae. B. Phaeocryptopusgaeumannii. c. Botryosphaeria
festucae. I>. Apiospora montagnei. E. Botryosphaeria dothidea. F. Didymo-
sphaeria diplospora. G. Didymella applanata. H. Pseudomassaria sepincolae-
formis. J. Botryosphaeria. foliorum.
PLATE XXXIX.
A. Karstenula rhodostoma. u. Thyridariarubronotata. c. Leptospora rubella.
D. Leptosphaeria lycopodina. E. Cucurbitaria laburni. F. Pleospora herbarum.
G. Leptosphaeria doliolum. H. Chitonospora ammophilae. I. Rebentischia
typhae. j. Pyrenophora bromi.
P L A T E XL.
A. Meliola niessleana. B. Irene calostroma. c. Pseudeurotium ovalis.
D. Anixiopsis stercorarea. E. Eurotium herbariorum. F. Orbicula parietina.
G. Lilliputia rufula. H. Zopfia rhizophila. I. Cephalotheca sulfurea. J. Ileleo-
coccum aurantiacum. K. Roesleria pallida. L. Onygena equina.
FIG. 1. - Some kinds of unitunicate asci. A. Operculate ascus of Barlaeina
amethystina. B. Bilabiate ascus of Ascozonus woolhopensis. c. Inoper-
culate ascus of Cudoniella acicularis. D. Indehiscent ascus of Tuber rufum.
E. Parallel-sided ascus of Stictis stellata. For bitunicate asci see individual
species figured on the plates.
FIG. 2. - Diagrammatic sections of ascocarps, showing the arrangement of
the asci. A. Apothecium. B. Perithecium. c. Pseudothecium. D. Thyrio-
thecium.
FIG. 3. - A. Morchella esculenta. B. Morchella elata. c. Mitrophora semilibera.
D. Verpa conica.
FIG. 4. - A. Helvella laeunosa. B. H. sulcata, c. H. crispa. D. Gyromitra
esculenta. E. Cyathipodium corium.
FIG. 5. - Hairs of the Humariaceae. A. Scutellinia stenosperma, ascospore
oil left. B. Trichophaea bicuspis. c. Cheilymenia crucipila, simple, 3, 4
and 5-armed hairs also occour. D. Trichophaea hemisphaerioides. E. T.
woolhopeia. F. Melastiza chateri. G. Sepultaria arenosa. H. Leucoscypha
leucotricha. I. Tricharia cretea. J. Cheilymenia theleboloides. K. Anthra-
cobia melaloma. L. A. macrocystis. All x500 except B x300.
FIG. 6. - A. Trichoglossum hirsutum. B. Geoglossum cookeianum. c. G. fallax.
D. Corynetes atropurpureus. E. Durella commutata. F. Ephelina lugubris.
G. Anthracobia humillima. H. Pseudoplectania nigrella. I. Leucoscypha
leucotricha. J. Leptopodia pezizoides. K. Ascozonus woolhopensis. L. Sphag-
nicola ciliifera.
FIG. 7. - A. Dasyscyphus virgineus. B. D. niveus. c. D. apalus. D. D. acuti-
pilus. E. Hyaloscypha hyalina. F. H. leuconica. G. Arachnoscypha aranea.
H. Pezizella eburnea. 3. P. chionea. K. Hyalopeziza ciliata. L. Mollisina
acerina. M. Cudoniella acicularis.
FIG. 9. - A. Tympanis conspersa. B. Tubeufia helicomyces. c. Rhytisma
acerinum. D. Polydesmia pruinosa. E. Hypoderma desmazieri. F. II.
virgultorum. G. Lophodermium juniperinum.
FIG. 10. - A. Acrospermum compressum. B. Ostropa barbara. c. Laqueria
sphaeralis. D. Neurospora sitophila. E. Robergea cubicularis. F. Stictis
fimbriata. G. Wettsteinina niesslii. H. Buergenerula biseptata.
FIG. 11. - A. Pseudographis elatina. B. P. pinicola. c. Karschia lignyota.
D. Ascus of K. bloxami. E. Abrothallus parmeliarum. F. Scutula episema.
G. Nesolechia oxyspora. H. Leciographa parellaria. I. Patellaria atrata.
J. Placographa flexella. K. Celidium varians. L. Homostegia piggotii.
M. Rhagadostoma lichenicola.
FIG. 12. - A. Xylosphaera polymorpha. B. X. hypoxylon. c. Rosellinia aquila.
D. R. mammiformis. E. Coniochaeta ligniaria. F. Xylosphaera hippo-
trichoides. G. Helminthosphaeria clavariarum. H. Poronia punctata.
I. Sordaria humana. J. Rombardia fasciculata. K. Podospora curvula.
L. Lasiosphaeria spermoides. M. Gelasinospora cerealis. N. Lasiosphaeria
hirsuta.
FIG. 13. - A. Lophiotrema praemorsum. B. Lophiostoma caulium. c. Lophi-
dium compressum. D. Eutypella prunastri. E. Diatripella quercina.
F. Diaporthe arctii on Arctium, G. Rhamphoria pyriformis. H. Ceratosto-
mella ampullasca. I. Ceratosphaeria lampadophora.
FIG. 14. - A. Chaetosphaeria myriocarpa. B. Zignoella ovoidea. c. Nitschkia
cupularis. D. Calyculosphaeria tristis. E. Bertia moriformis. F. Tympa-
nopsis euomphala. G. Enchnoa lanata. H. Calosphaeria pulchella.
FIG. 15. - A. Farlowiella carmichaeliana. B. Gloniopsis levantica. c. Hystero-
graphium fraxini. D. Hysterium angustatum. E. Mytilidion gemmigenum.
F. Glonium lineare. G. Lophium mylilinum. H. Byssolophis sphaerioides.
I. Actidium hysterioides. j. Phyllachora graminis. K. Dothidea tetraspora.
L. Plowrightia ribesia.
FIG. 16. - A. Herpotrichia macrotricha. B. Thaxteriea phaeostroma. c. Me-
lanomma pulvis-pyrius. D. Trematosphaeria pertusa. E. Berlesiella
nigerrima. F. Perisporium vulgare. G. Leptosphaeria acuta. H. Acantho-
phiobolus helminthospora. I. Naumovia abundans. J. Leptosphaeria dolio-
loides. K. Teichospora obducens.
FIG. 17. - A. Loramyces juncicola. B. Ascotricha chartarum. c. Ceratocystis
piceae. D. Chaetomium elatum. E. Capnodium salicinum. F. Leveillula
taurica. G. Uncinulabicornis. n. Phyllactiniaguttata. I. Erysiphe polygoni.
j. Sphaerotheca humuli. K. Microsphaera grossulariae. L. Podosphaera,
oxyacanthae var. tridactyla.
FIG. 18. - A. Chloroscypha sabinae. B. Coleosperma lacustre. c. Dothidea
sambuci. D. Cryptomycina pteridis. E. Glomerella montana. F. Isothea
rhytismoides. G. Microthyriella osmundae. H. Otthia spiraeae.
FIG. 19. - A. Podosordaria tulasnei, habit sketch natural size, apex*bf stroma
XLO, ascus tip x660. B. Dermatina quercus, stromata natural size on pale
patch of bark and in lens view (one as seen from below), asci and spore
x660. c. Xylosphaera longipes, stroma natural size, portion of surface in
lens view, ascospores x660. D. Podonectria tenuispora, pseudo thecia on
scale insect x60, ascus x660. E. Pleospora multiseptata, pseudothecia on
Ammophila natural size, ascospore x660. F. Mycobilimbia killiasii,
apothecium x20, ascospore x660. G. Mycomelaspilea leciographoides,
apothecia x20, ascus x660. H. Seynesiella juniperi, habit sketch natural
size, thyriothecium xlOO, ascus and ripe ascospore x660.
FIG. 20. - A. Thyridium lividum. B. Pithyella ilicincola. o. Broomella
vitalbae. D. Amphisphaeria umbrina. E. Strossmayeria basitricha. F. Caryo-
spora callicarpa. G. Keissleriella culmifida. H. Plagiostoma devexa.
i. Asteromassaria macrospora. J. Perrotia flammea. K. Astrosphaeriella
applanata. L. Delitschia winteri.
FIG. 21. - A. Mycosphaerella lineolata. B. M. punctiformis. c. M. iridis.
D. Pringsheimia sepincola. E. Pseudoplea myrtillina. F. Sphaerulina taxi.
G. Venturia rumicis. H. V. maculae]ormis. I. Gibbera myrtilli. J. Coleroa
chaetomium. K. C. alchemillae. L. Gibbera vaccinii. Lesions xlO, asci and
spores x660.
FIG. 22. - A. Stigmatea robertiani. B. Aulographum vagum. c. Schizothyrioma
ptarmicae. D. Euryachora ulmi. E. Mycosphaerella killiani. F. Rhopogra-
phus filicinus. G. Euryachora betulina. H. Endodothella junci. J. Scirrhia
rimosa.
FIG. 23. - A. Massarina eburnea. B. Massariella bufonia. c. Vialeia inscu Ipta.
D. Melomastia mastoidea. E. Pleomassaria siparia. F. Massaria inquinans.
G. Fenestella fenestrata. H. Massaria pupula.
galiorum. E. L. macrospora. F. L. planiuscula. G. L. praetermissa. H. L.
cesatiana. i. L. modesta. l.L.derasa. K. L. silenes-acaulis. L. L. sowerbyi.
M. L. juncina. N. L. juncicola. o. L. typharum. p. L. epicalamia. Q. L.
littoralis. R. L. nardi. s. L. nigrans. T. L. herpotrichoides. u. L. graminis.
v. L. fuckelii. w. L. eustoma. x. L. tritici. Y. L. nodorum. z. L. luctuosa.
AA. L. arundinacea. BB. L. michotii. cc. L. pontiformis
FIG. 25. - A. Leptosphaeria rusci. B. L. niessleana. c.L.vectis. D. L. scitula.
E. L. haematites, F. L. purpurea. G. L. coniothyrium. H. L. scirpina.
I. L. maritima J. L. albopunctata. K. L. macrosporidium. L. L. orae-maris.
M. L. pelagica. N. Passeriniella discors. o. Trematosphaeria circinans. P.
T. heterospora. Q. Keissleriella ocellata. R. K. gallica. s. K. subalpina.
T. Physalospora empetri. u. Keissleriella linearis, v. Glomerella cingu-
lata. w. Pseudomassaria chondrospora. x. Entosordaria ammophilae.
Y. Anthostomella punctulata. z. Anthostoma decipiens. AA. Griphosphae-
ria nivalis.
FIG. 26. - A. Mamiana fimbriata. B. Hypospilina bifrons. c. Linospora
capreae. D. Gnomoniella tubiformis. E. Gnomonia vulgaris. F. Diapleella
clivensis. G. Paradidymella tosta. H. Phomatospora berkeleyi. I. Sydo-
wiella fenestrata. j. Melogramma spiniferum. K. Pseudovalsa laneiformis.
L. Gauemannomyees graminis.
FIG. 27. - A. Cryptomyces maximus. B. Anisogramma virgultorum. c. Lopa
dostoma turgidum. D. Endoxyla operculata. E. Ascus of Crytomyces maxi-
mus. F. Diatrype stigma. G. D. disciformis. H. Eutypaacharii. I. Qua-
ternaria quaternata. J. Peroneutypa heteracantha. K. Cryptosphaerio
eunomia.
FIG. 28. - A. Coccomyces coronatus var. trigonus. B. Melittosporium pteridi-
num. c. Rhabdocline pseudotsugae. D. Therrya pini. E. Propolis versicolor.
F. Naemacyclus niveus. G. Phacidium lacerum. 11. Phacidiostroma multi-
valve. J. Didymascella tetraspora. K. D. thujina. All XLO.
FIG. 29. - A. Hyaloscypha flaveola. B. Retinocyclus abietis. c. Cistella piceae
var. laricinum. D. Unguiculella robergei. E. U. hamulata. F. Psilachnum
asemum. G. Dasyscyphus clavisporus. H. Fimaria theioleuca. J. Phaeo-
helotium lilacinum. K. Naemacyclus caulium. L. Patellariopsis clavispora.
M. Thelebolus stercoreus.
FIG. 30. - A. Ophiobolus acuminatus. B. Ascospore of O. (Nodulosphaeria)
cirsii. c. Gyromitra infula. D. Pteridiospora scoriadea. E. Allescheria
terrestris. F. Emericella nidulans, one cleistocarp partly wiped free of
hiille cells to expose the dark wall beneath them. G. Synaptospora tartarica.
H. Metacapnodium juniperi. j. Atichia glomerulosa. K. Aulographum
festucae. L. Sphaeronaemella ftmicola.
FIG. 31. - A. Morchella hortensis. B. Leptopodia ephippium. c. Cyathipodia
villosa. D. Leptopodia stevensii. e. Peziza sterigmatizans. F. P. sepiatra.
G. P. ampliata. H. P. arvernensis. J. Cheilymenia fibrillosa.

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