Corner - A Monograph of Clavaria and Allied Genera - 1950

J
A MONOGRAPH Of
CLAVARIA
AND ALLIED GENERA
E. J. H. Corner
ANNALS OF BOTANY MEMOIRS
No. 1
A MONOJRAPH OF
CLAVAR IA AND
ALLIED GENERA
A MON.OGRAPH OF
CLAVA R IA AND
ALLIED GENERA
BY
E. J. H. CORNER
University Lecturer in Botany
Cambridge
BISHEN SINGH MAHENDRA PAL SINGH

23-A, New Connaught Place
Dehra Dun-248 001 (INDIA)
2005
ISBN: 81-211-0460-2
A MONOGRAPH OF
CLAVA R IA AND
ALLIED GENERA
BY
E. J. H. CORNER
University Lecturer in Botany
Cambridge
OXFORD UNIVERSITY PRESS

LONDON: GEOFFREY CUMBERLEGE
1950
FOREWORD
THE publication of scientific contributions that exceed a certain length always
presents a problem. Editors of journals, however great the merits of such
papers, are rightly disinclined to accept them, and publishers are usually not
prepared to undertake their publication without a more or less considerable
subvention. It is hoped that, in the botanical field, the Annals of Botany
Memoirs will serve to satisfy the need for a medium for the publication of
longer papers and monographs, and that the Memoirs will render available to
the botanical public the fruits of scientific investigation of the highest standard.
The Annals of Botany Company has been fortunate in being able- to accept
Corner's Monograph of Clavaria and Allied Genera for publication as the first
of these Memoirs. Representing as it J:loes the fruits of many years' work in
the tropics and in temperate regions, and likely to become an important work
of reference, there can be no doubt that it will set a standard for subsequent
publications in this series. It may, however, be well to add that it is not
anticipated that most of the Memoirs will in any way reach the length of this
first one, nor is it expected that more than a small proportion of them will be
taxonomic in their approach.
The Memoirs will be published as and when the funds at the disposal of the
Company are adequate for the purpose and suitable material becomes avail
able. Should this present Memoir find the market it deserves, the first of the
conditions just mentioned will be satisfied.
The Company has to acknowledge the receipt of a grant of £150 towards
the costs of production of this Memoir from the Parliamentary Grant-in-aid
for Scientific Publications administered by the Royal Society. Mr. Corner has
himself borne the cost of several of the coloured plates. The Company is also
greatly indebted to the Clarendon Press for the care taken in the publication of
this Memoir.
F. E. FRITSCH
Chairman, Annals of Botany Company
July 1949
ACKNOWLEDGEMENTS
THE monographs of Cotton and Wakefield on the Clavariaceae of Great
Britain (1919), of Burt (1922) and of Coker (1923) on those of. Canada and
the -United States, and the-floristic works of Rea for Great Britain (1922),
Bourdot and Galzin for France (1927-8), Donk for Holland (1933), van
Overeem for Java (1923), Petch fo,r Ceylon (1925), and Cleland (1916-35) for
South Australia, have made the Clavarioid fungi one of the best-known groups
of Basidiomycetes. They have brought, and will continue to bring, many
successors into the field. As one who would hand on the results of their work
to tropical mycologists, I have made the freest use of their publications and
express my gratitude that it is possible to build up one branch of systematic
mycology on such sure ground. Except where required for taxonomic reasons,
I have omitted references to earlier works, because most students will not
have access to them or, if so fortunate, repetition would be unnecessary.
I wish to thank, also, all the mycologists whose works I have cited for the
privilege of having used their data. I am particularly grateful to Mr. A. D.
Cotton, who lent me his notes and specimens on British Clavarias for leisured
study at my house in 194-6; to Miss E. M. Wakefield for extending to me
similar courtesy for the microscopic study of specimens in the Kew Her
barium; to Mr. M. A. Donk for lending me his unpublished notes on Typhula
and Pistillaria and for many hours of incessant argument in Singapore and
England on matters of nomenclature and generic limitation; to Dr. R. Heim
for the loan of specimens from the Paris Herbarium: to Professor G. W.
Martin; Professor J. B. Cleland, Dr. L. Wehmeyer, Dr. R. Singer, Dr. R.
Heim, and Dr. M. S. Doty for sending me their papers on Clavarioid fungi:
to the late Mr. Carleton Rea for the privilege of many mycological excursions
in his company: to Professor F. T. Brooks for his advice on the preparation
and publication of the manuscript, and, lastly, to Professor Kwan Koriba,
Professor Hidezo Tanakadate, and Marquis Yositika Tokugawa- for having
conserved my mycological notes and collections during the Japanese occupa
tion of Malaya, 194-2-5, and for having enabled me to continue my scientific
work under their protection.
E. J. H. C.
GREAT SHELFORD, ENGLAND
6 October I946
CONTENTS
INTilODUCTION
Object-scope-material-method Xl
GEl'J..ERAL CONSIDERATIONS
Description of the Clavarioid fruit-body I
Affinity of Clavarioid genera 22

Geographical distribution 27
Collection and study 30
Identification 31
TAXONOMIC AND MORPHOLOGICAL NOTES ON GENERA 32

Aphelaria - Araeocoryne - Caripia - Ceratellopsis - Clavaria
- Clavariachaete - Clavariadelphus - Clavicorona - Clavulina -
Clavulinopsis - Dendrocladium - Lachnocladium - Lentaria -
Mucronella - Physalacria (and Hormomitaria) - Pistillaria - Pistillina
- Pteruloid series (Deflexula, Dimorphocystis, Pterula; Pterulicium) -
Ramaria - Ramariopsis - Typhula
CLASSIFICATION
Natural key
Artificial key
Descriptions of genera and species in alphabetical order
Aphelaria - Araeocoryne - Caripia - Ceratellopsis - Chaetotyphula
- Clavaria - Clavariachaete - Clavariadelphus - Clavicorona -
Clavulina - Clavulinopsis - Deflexula - Dimorphocystis - Hormo
mitaria - Lachnocladium - Lentaria - Mucronella - Myxomycidium
- Physalacria - Pistillaria - Pistillina - Pterula - Pterulicium -
Ramaria - Ramariopsis - Scytinopogon - Typhula
List of Clavarioid genera, generic synonyms, and type-species
Novitates
GENERAL BIBLIOGRAPHY
INDEX TO GENERA, SPECIES, AND SYNONYMS
GLOSSARY 737
LIST OF PLATES
At end
PLATE 1. Clavaria helicoides, C. incarnata, C. straminea, and C. Zollingeri.
PLATE 2. Cl.avaria acuta, C. argillacea, C. Gibbsiae, and Clavulinopsis spiralis.
PLATE 3. Clavariadelphus pistillaris.
PLATE 4. Cl.avulina cinerea and C. cristata.
PLATE 5. Clavulina cartilaginea and Lentaria surculus.
PLATE 6. Clavulina gracilis, C. Leveillei, and Clavulinopsis arenicol.a.
PLATE 7. Clavulinopsis alcicornis and C. miniata.
PLATE 8. Clavulinopsis amoena, C. luteo-alba, and C. pulchra.
PLATE 9. Cl.avult'nopsis amoena and C. luteo-tenerrima.
PLATE 10. Clavulinopsis corniculata, C.fruticula, C. helvol.a, C. spiralis, C. umbrinella,
and Ramariopsis pulchella.
PLATE 11. Deflexul.a fascicularis, Lachnocladium fulvum, and L. zonatum.
PLATE 12. Ramaria fragillima, R. Invalii, and R. Zippelii var. gracilis.
PLATE 13. Ramaria gracilis, R. luteo-flaccida, R. ochraceo-virens, and R. Zippe/ii var.
gracilis.
PLATE 14. Ramaria cyanocephala, R. Zippe/ii, and Aphelaria dendroides.
PLATE 15. Clavaria fossicola, C. ornatipes, Clavicorona candelabrum, Deflexula
lilaceo-brunnea, and Hormomitaria sulphurea.
PLATE 16. Chaetotyphula hyalina, Pterula typhuloides, and Pterulicium xylogenum.
INTRODUCTION
Object. The classification of Basidiomycetes is artificial. I have always con
sidered that the larger groups should be taken, one at a time, and their
particular kinds of fruit-body described in terms of hyphal properties. By
studying as wide a range of species as possible, from all parts of the earth,
one should then be able to see the group in its own light, appreciate the
resemblances and differences revealed by this analysis, and so arrive at a
natural and fungal (that is, hyphal) rearrangement. Eventually, on synthesis
of all groups, it should be possible to trace phyletic lines through the homo
plastic levels which are the present concepts of the groups and so to discover
the natural classification. The field is too vast for random investigation.
ln 1925 I began to study Clavariaceae, as the well-defined group of Homo
Basidiomycetes fundamental to the whole, yet so rarely mentioned in text
books as to seem without importance. Since then I have made many sallies
into the unknown, all of which have convinced me of one preliminary, but
prime, essential. Before any such undertaking something must be done to
facilitate systematic mycology in the tropics. To omit the tropical element is
certain failure: to include it supposes not just names but correct names, and
to get any name at all for a tropical basidiomycete is so hard that most
mycologists do not try. Tropical libraries, with a few notable exceptions, are
quite inadequate and the very descriptions of tropical fungi which have been
published have, for the most part, been drawn from dried specimens, in days
of microscopic carelessness, and are practically unrecognizable. It is admitted
the larger fungi, more than any other, should be studied in the living state,
but what can the tropical mycologist do without books or type-specimens,
and what incentive has he to collect the material which will be the remedy?
One is tempted to compare the systematics of higher fungi with a classifica
tion of flowering plants based on the floras of Europe and temperate North
America, but that would be incorrect. In the rain-forest of Singiipore Island,
almost on the equator, there occur, for instance, typically or in perplexing
resemblance, Amanita rubescens, Amanitopsis vaginata, Lepiota acutesquamosa,
Collybia maculata, Hygrophorus puniceus, Omphalia muralis, Laccaria laccata
(never var. amethystina), Pleurotus ostreatus, Russula pectinata, Lactarius
piperatus, Pluteus cervinus, Entoloma lividum, Cortinarius armillatus, Naucoria
cidaris, Psathyrella disseminata, Marasmius foetidus, Crinipellis stipitarius,
Boletus scaber, Hydnum repandum, Craterellus canthar.:llus, and Clavaria ver
micularis. In this work I record the N. American Clavaria ornatipes and C.
angulispora from Malaya. In the tropical mountains it is impossible to say
what well-known temperate fungi may not be found, as Cortinarius torvus,
C. elatior, C. vibratilis, Armillaria mellea, and Peziza aurantia in Pahang.
But, side by side with them, can be found stupendous problems which fit no
genus-I have one which combines the characters of Cantharellus, Stereum,
Xll INTRODUCTION
and Asterostromella, and another whir.h links a group of Omphalia with
rigid, Stereum-like brackets. The position of the tropical mycologist is simply
that, if he would name his 'higher fungi', he must be prepared to do so on a
world-basis, for they may have been described from Patagonia or .Japan,
Ecuador or New Guinea, the Belgian Congo or California, Europe or New
Zealand. Unlike the flowering plants, where interdigitation of temperate and
tropical species in micro-genera is the exception, in the higher fungi it _is" the
rule. As the temperate mycologist would extend his knowledge to the tropics,
so the tropical mycologist must have the most detailed cognizance of tem
perate mycology: he cannot do this, nor the temperate mycologist that, until
there are handy and comprehensive monographs.
My intention, therefore, has given first place to pure systematy. To name
what I would describe I have revised the Clavariaceae of the world. In thi�
work I hope to give tropical mycologists the means of identifying Clav;J.rioid
fungi to the extent that what cannot be named from its pages with certainty
should be described as new: and the full synonymy is intended to facilitate
the coining of new names without the bugbear of homonyms.
Scope. In writing about Clavarioid fungi I have excluded Heterobasidio
mycetes. Strictly, Thelephora ( = Phylacteria) should be included but,
except for indication in the general key, I have reserved it for another mono
graph. The name Clavariaceae is undesirable because many of the Clavarioid
genera are not nearly related, e.g. Typhula, Pterula, and Lachnocladium, and
it is impossible to know what are the limits of true Clavariaceae in the
taxonomic sense of the type-genus Clavaria s. str. I have included Physalacria
and Mucronella because the new genera Hormomitaria and Deflexula show
that the first must be regarded as Clavarioid and that the second includes both
Clavarioid and Hydnoid fungi.
Material. I have collected myself about 120 species of Clavarioid fungi,
mostly in England and Malaya. I have studied the type-collections and
authentic herbarium-material of some sixty more species, excluding those
which I have reduced as synonyms. This is 180 out of a total of 540
species which I recognize, but, of these, only some 170 are too inadequately
known for certain identification. That is to say, it should be possible to
identify from the descriptions 370 species or three-quarters of the world
flora. For the rest, type-material may exist for verification: if not, the species
may be recognized by mycologists visiting the 'type-localities'.
Method. Through the hyphal analysis of my own material I have been
able to discover a number of natural groups which must be separated as new
genera from the artificial assemblages of Clavaria, Lachnocladium, Pterula,
and Pistillaria: some already have generic names, as Clavulinopsis v. Ov.,
Ceratellopsis Konr. et Maubl., and Scytinopogon Singer. In the case of the
north temperate and S. Australian species it has generally been possible
from recent descriptions to ascertain their position among the new genera and
I have, accordingly, transferred them. Doubtful sp1cies have been left in the
genus where they were described, as incertae sedis., Thus, in place of the
INTRODUCTION xiii
customary 11 or 12 genera, I recognize twenty-seven. Clavaria has been
pruned from 620 species and synonyms to 56 species (30 of which are doubt
ful): Lachnocladium has been pruned from 112 to 20, and Pterula from
c. 80 to 52 (of which 30 are still doubtful). The most disturbing innovation
will be the genus Clavulinopsis (66 species) for many of the common north
temperate species usually regarded as typical Clavaria. A world-wide study
of Chvarioid fungi, however, is much more involved than might be expected.
A white, branched 'Clavaria', for instance, may belong to one of several micro
scopically distinct groups (Clavaria s. str., Clavulinopsis, Clavulina, Rama
riopsis, Scytinopogon, Aphelaria, Pterula, or Lachnocladium): their hyphal
peculiarities must be distinguished if identification is to be accurate, or one
will revert to the I<'riesian attitude of the pocket-lens, which named all simple,
yellow, non-caespitose Clavarias as C. inaequalis. There are, certainly, still
more genera to be recognized. Actually, no one can now force the old genera
to fit the species of the world, so that Lachnocladium has become the
receptacle for any dried branched 'Clavaria' from the tropics.
In introducing the microscopic method of classification I have been careful
to be consistent and to avoid using characters for which I have insufficient
explanation: that is, I have employed as fundamental the one method of
hyphal analysis. The modern tendency in the systematy of Basidiomycetes is
to split the old genera into small genera on microscopic details without full
microscopical examination. In effect, macroscopic direction has given way
to microscopic confusion: macroscopic terms mingle ambiguously with micro
scopic and, in the mixture of macroscopic and microscopic genera, there has
appeared not one guiding sign of understanding. We seem to be in mid
stream, progressing, yet through our microscopes unable to see the banks or
·, to tell where we are. So it is possible to use in one case the direction of the
nuclear spindle to distinguish two genera and in the next the form of the
fruit-body: it is possible to describe cystidia and striae on spores, and not to
realize that the 'sterile' growing-point of the fruit-body is a universal Clava
rioid (and Basidiomycetous) feature, not peculiar, as widely supposed, to
Ceratellopsis (Ceratella auctt.): it is possible to emphasize as specific the
shapes of branches, whether acute or blunt, yet not to realize that every
Clavarioid branch becomes blunt before, and acute after, it subdivides: it is
possible to distinguish some genera on spore-colour, others on macroscopic
texture. When there is such admixture of ideas the science is illogical and
unreal.
Nevertheless, macroscopic terms are the present bar _to progress. A few
examples will explain what I mean. The term 'pileus' is used for any expansion
of the fruit-body with or without a stalk. In this sense it is useful, but it does
not imply a morphological entity. Microscopically and morphologically, a
pileus is a dorsiventral hyphal tissue with marginal, or centrifugal, growth
and sterile upper and fertile lower sides. At present it is not known whether
Craterellus clavatus and C. unicolor (which I treat, here, as Clavariadelphus)
have true morphological pilei as Cantharellus or are truly apileate as all
xiv INTRODUCTION
Clavarioid fungi. The terms 'tough' and 'coriaceous' are useful in their proper
place but not for generic definition. Toughness may be caused (a) merely by
the drying· of thin-walled hyphae, as in herbarium-specimens, so that the
tissue is composed only of adherent hyphal walls; (b) by lack of inflation of
thin-walled hyphae; (c) by thickening of the walls of inflated hyphae; (d) by
the presence of skeletal hyphae; (e) by the presence of dichophyses; and (J) by
the presence, also, of binding hyphae (truly coriaceous as in Polystictus versicolor,
in contrast to the fibrous-coriaceous P. perennis). For all of the first five
reasons Clavarioid fungi have been placed in Lachnocladium, yet only the
fifth (e) is valid and it has been suggested as the basis of a new genus.
Each of the six reasons represents a different hyphal construction and an
essentially different manner of fungous growth: or, on the principle of hyphal
analysis, toughness covers six generic categories. Lastly, I would mention
spore-colour. It may reside in the wall, the cytoplasm, or the oil-drops of the
fresh spore, or it may be an after-effect of drying. Thus, among Clavarioid
fungi, Ramaria has ochraceous spore-walls; Clavulinopsis fusiformis, C. luteo
tenerrima, and C. pulchra may have yellow oil in their old spores lying on the
hymenium; Clavulina and Lentaria have colourless fresh spores, the walls of
which may turn yellowish on drying: Clavariadelphus spp. have pale yellowish
cytoplasm, as Clavaria helicoides has pink cytoplasm. The inexperienced may
wonder why spore-colour can be a generic character and that of the fruit-body
only specific, though caused by similar microscopic appearances. There is no
reason: at most, it can be replied that mycologists have, through lack of know
ledge, not yet been able to consider the matter. Nevertheless, hyphal analysis
may supply the answer: the ochrosporous agarics have yellow or brown spore
walls which merely continue their common hyphal character of yellow or
brown hyphal walls (often incrusted), expressed in the prevalent ochraceous
and brown colours of the fruit-bodies of Cortinarius, Inocybe, Hebeloma,
Naucoria, Galera, and so on: on the other hand, the pink cytoplasm of the
spores of the rhodosporous agarics seems to express the cytoplasmic pigmenta
tion of the hyphae of their fruit-bodies: the purple and black spore-walls of
the Porphyrosporace and Melanosporae appear to be the effect of the oxidation
of melanins in the walls, as expressed also in the reddening and blackening· of
the tissue of the fruit-body in Psalliota and Lepiota. Not until the hyphae
have been traced through their elaborate organization in fruit-bodies from the
mycelium to the spores which are finally abstricted from their tips can the
true problem of the higher fungi be seen. This is the principle which I call
hyphal analysis and which I consider to be the only means that human beings
can have of understanding fungi.
Because Clavarioid fungi are manifestly polyphyletic I have used no
systematic units of higher rank than the genus. Groups of allied genera I
have referred to as series, e.g. the Pteruloid series. I have even been chary of
the use of subgenera. It is obvious that, at the present time when most
organisms have disappeared or have evolved into others, genera must be
recognizable as absolutely distinct units, having no living links with other
INTRODUCTION xv
organisms. Subgenera, to be natural, however, imply that one understands
the manner of evolution of modern species within a genus, and in our present
state of mycological knowledge this is hardly possible.
Thie monograph is, thus, the systematic introduction to a later work on
the morphology of Clavarioid fungi. For practical purposes it is better to
separate a volume for laboratory and field use from one intended for the desk
and arm-chair. I have spared Iio pains in either, having revised them wholly
four times in the last three years, and in part considerably more.
GENERAL CONSIDERATIONS
DESCRIPTION OF THE CLAVARIOID FRUIT-BODY
Development-Branching-Additional Form-factors-Simple Fruit-bodies
Hymenium-Hyphae-Cystidia-Habitat-Colour
DEVELOPMENT
THE fruit-body begins as a small subglobose primordium, 0·2-I·O mm. wide,
produced directly from the mycelium or, as usual in species with massive
growth, from a rhizomorph. Apical growth soon develops at the distal end
a.:-i.d the spindle-shaped primordial shaft is initiated to form the stem, or
trunk, of the fructification. Clavariadelphus junceus is unique in that its
rhizomorphs are mostly independent of the fruit-body and only rarely become
fertile. The earliest stages in the development of Clavariadelphus pistillaris
are angiocarpic (p. 45) and, perhaps, the fruit-body of Caripia is angiocarpic.
So far as is known, Clavarioid fruit-bodies are typically gymnocarpic. The
apical growing-point of th,� primordial shaft may persist without modification
to build the simple fruit-body or it may divide to form the branched fruit
body. In most cases the cells of the hyphae inflate acropetally immediately
behind the growing-point, which is thus rapidly projected. In some reduced
fruit-bodies, as Physalacria, Pistillaria, and Typhula, apical growth may be
arrested very early and inflation may be so prolonged that the fruit-bodies
appear to emerge as though from a button-stage, as in typical agarics, but
development is never really indirect with a period of apical growth followed
by a separate period of expansion. In other cases the cells of the hyphae do
not inflate and the fruit-body grows in. height merely by apical growth, e.g.
Pterula, Lachnocladium, Aphelaria, and Scytinopogon.
Details of development are known in few species, and I have described
them, mostly, under the generic discussions.
BRANCHING
There are two kinds of apical branching, the radial and the flattened. In
active growth the primordial shaft, or a branch, is more or less acute. Before
branching it thickens in the case of radial branching, or it broadens out in the
case of flattened branching, and becomes blunt. Thus, according to its state
of growth, a branched fruit-body may have blunt tips or acute _tips, and the
difference is often so marked, especially in species with flattened branching,
that they have erroneously been considered Stereoid in the early primordial
stages of branching, e.g. Scytinopogon and Aphelaria dendroides. It seems
that, preliminary to branching, apical growth declines first · in the central
leading hyphae of the conical apex so that the outer or peripheral hyphae
catch up with them, and then apical growth is renewed at evenly spaced points
Sl19 B
2 GENERAL CONSIDERATIONS
of the dying-away field of growth to form either a circle or a fan of new
branches.,
A third 1orm of branching, which is not apical, is the adventitious. There
is no truly monopodial branching with spiral or acropetal 'phyllotaxie'.
Radial Branching. On the broadly rounded or truncate apex of the pri
mordial shaft many new radially disposed points of outgrowth arise to give a
• 0
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o ····..oo o o
0 9.-· .-�
0 �--....... � ... C?.
0 0 O\··� 0 0o o
.. c) O O O
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o····..
°
0 Q/
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'I'Ex.T-FIG. 1. Diagram of a radialJy branched Clavarioid
fruit-body in longitudinal section, with cross-sections to
show the arrangement, increasing number, and attenuation
of the branches: the tracts of the primary branches shown
by dotted lines in the cross-sections.
polychotomous state. As the process is repeated the branchlets become

thinner and polychotomy gives place to dichotomy. Thickening of the
branches by inflation of the hyphae or by thickening of the hyrnenium soon
obscures their m3nner of origin as minute points of outgrowth, and their
bases become confluent. Generally successive branches lie in alternating
planes, as multiple decussation, and typically, as the branches subdivide and
become attenuate, so their internodes lengthen. Abnormalities of suppression
and accentuation so often intervene through the position of the branches,
whether central or peripheral, that great regularity is seldom noticeable: it
seems, however, that under perfect conditions the manner of growth and time
of subdivision is the same in all branches of a fruit-body which is thus con-
DESCRIPTION OF THE CLAVARIOID FRUIT-BODY 3
structed in tiers, as pulses of growth. An instance is the special case of the
pyxidate branching of Clavicorona (Text-Figs. ZF, 17-19): the main axis or a
branch is acerose or filiform at first, but slowly becomes obconic until the
apex is truncate or subcyathiform: then, on the periphery, arise 3-8, com
monly 4, new branches which repeat the process, but the bases of the branches
A s
r E
'I'ExT-FIG. 2. Radial branching: A, symmetrical dichotomy in Ramaria
fragillima: B, decussation in Ramaria Zippelii: C, unequal dichotomy
in Ramaria fragillima: D, polychotomy in Ramaria Zippe/ii, and E,
with unequal development of the branches: F, pyxidate branching in
Clavicorona candelabrum, with 4-chotomy (in longitudinal section):
G, attenuate tips of Lentaria surculus: mag.
thicken very little and the original apex becomes a shallow, sterile cup: very
regularly tiered fruit-bodies, with sets of alternating branches, are thus often
constructed, as ideal models of a Clavarioid fungus.
Radial branching occurs in all Clavarioid genera with inflating hyphae, as
Clavaria, Clavulina, Clavulinopsis, Lentaria, Ramaria, and Ramariopsis.
Usually only the initial set, or second set, of branches are polychotomous,
even in massive fruit-bodies, and slender fruit-bodies are usually dichotomous
throughout, or at first apparently polychotomous through the early condensa
tion of the internodes. Possibly some massive species of Ramaria, as R.
botrytis, are polychotomous throughout: the persistent, pyxidate, polycho
tomy of Clavicorona seems related to the lack of inflation or the long-delayed
inflation of its hyphae.
Flattened Branching. 1 Before subdivision the apex of the primordial shaft
or of a branch flattens and widens, as the tip of a screwdriver, becoming
ligulate, spathulate, and cuneate or flabelliform, and along the knife-edge
TllxT-FIG. 3. Diagrams of successive stages in the

growth of a radially branched fruit-body with in
flating hyphae: polychotomy followed by dicho
tomy: the diagram on the right in longitudinal
section.
'growing-point' so formed new growing-points appear at regular intervals

to give a palmate or flabellate polychotomy. The new branches quickly
become cylindric through branching of their hyphae and through concrescence
of adjacent hyphae on both sides of the knife-edge (and this influence of the
new branches, extended over the sides of the knife-edge, generally causes
short slight ridges to extend from them down the flattened sides of the dilated
�pex of this parent branch). In turn, the new branches flatten and repeat the
process but, as with ·radial branching, attenuation sets in and polychotomy
soon gives way to a characteristic and rather forcipate, or fish-tailed, dicho
tomy. Successive branchings may be in alternating planes, as in Thelephora,
Aphelaria, Pterula, and Lachnocladium, but in Scytinopogon (and some
' I have avoided the word dorsiventral because it implies dissimilar surfaces, as a sterile
upper and fertile lower surface, and in this sense it is needed to describe branches with uni
lateral hymenium whether of radial origin (Ramaria, Clavulina) or flattened (Thelephora,
Scytinopogon).
DESCRIPTION OF THE CLAVARiOID FRUIT-BODY 5
sryecies of Clavulina) it very remarkably persists in one original plane, and
though, through mutual pressure and unequal hymenial growth, thP. branches
become distorted, it is possible to flatten the fruit-body out so that ;t resembles
a much-divided, bilateral seaweed as Cutleria, Halymenia, or Gigartina.
Thus, in its early stages of growth, Scytinopogon is exactly Stereoid (as
Stereum multizonatum), and it is easy to see how, without ultimate subdivision,
a Clavarioid fungus with a few sets of flattened branching can produce a
multipileate Stereum or Sparassis: indeed, in Thelephora ( = Phylacteria)
___ ...
. � ----·
C B ------·e
A
TEXT-FIG. 4. Diagrams of successive stages in flattened branching:
A, a young fruit-body 4- to :z-chotomous, with inflating hyphae:
B, branch-tips with uninflating hyphae: C, branch-tips with inflating
hyphae.
there are all transitions between the two forms, leading eventually to the
mesopodal, agaric-like, pileus as a webbed, dorsiventral, Clavarioid fruit-body.
Flattened branching occurs typically in genera \Yithout inflation of the
hyphae, namely, Lachnocladium, Pterula, ApheJaria, Scytinopogon (and
Thelephora). Commonly, in the absence of marked thickening of the branches,
the residual tissue of the knife-edge 'growing-point' between the new branches
persists as a fine web or it becomes fertile and, with thickening hymenium,
unites the bases of the branches to form the palmate or flabellate axils of
specific descriptions. Exceptions seem to occur in some species of Clavulina
and Ramaria with inflating hyphae: in them the inflation of.the hyphae is slow
or slight and so they appear as intermediates:
In some species of Pterula there is false monopodial branching, as de
scribed on page r ro. Two or more condensed, alternating dichotomies,
without internodes, occur and one branch of each dichotomy overtops the
other to maintain an axis.
Adventitious Branching. An adventitious branch arises by a local prolifera
tion of hyphae at any part of the surface of the fruit-body and grows out as a
new branch. Such false branching occurs in some species of Typhula, Ceratel
lopsi,s, Pi�illaria, and Pterula. It is normal in Pterulicium and in Deflectula,
in which it: produces by basal proliferation the Mucronelloid tufts of spines
characteristic of one section of the genus. It occurs in the rhizomorphs of
Clavariadelphus junceus and occasionally on its fruit-bodies as abortive pro
liferations. It occurs also in some species of Clavulina and is often confined
TEXT-FIG. 5. Flattened branching in one plane in Scytinopogon angulisporus, the

hyphae not inflating: polychotomy followed by dichotomy: (the earth shown in
black): nat. size.
to the expanding, but attenuate, dying-away apices and so helps to produce

their characterist� cristate tips. The strigose processes on the stems of
Clavulina ornatipes and some species of Aphelaria are comparable sterile
innovations.
ADDITIONAL FORM-FACTORS IN BRANCHED FRUIT-BODIES
Abortion of Branches. In polychotomous tips some of the newly formed
branches abort or develop but little, especially on the outside of the fruit
body. Similarly, if the primordia of the fruit-bodies are caespitose, the peri-
pheral ones mostly abort while a few centrally placed ones develop fully.
There is a centripetal tendency in typical-branched Clavarioid fruit-bodies
which is shown by their conical outline. In Thelephora, in contrart, there is a
centrifugal tendency, the outer branches tending to develop as much, or more,
than the central, which may abort; thus Thelephora, alone among Clavarioid
fungi, has transitional forms to the pleuropodal and mesopodal Stereoid
condition.
Suppression of Branches. In effect this is indistinguishable from abortion
but the cause of suppression is not a fundamental centripetal tendency; it
results from overcrowding and interference of the branches, particularly in
the centre of the fruit-body, where they have no room for development.
Suppressed internal branches are commonly seen in the fastigiate fruit-bodies
of Lachnocladium and Ramaria.
Successive Branching. In regularly formed fruit-bodies, at least in their
early stages, branching is synchronous: they are crowned with acute or blunt
tips, as the case may be. Sooner or later abortion and suppression introduce
irregularities, and the branches begin to grow at different rates, corresponding
doubtless with the number of branchlets which are to be fed through the
separate channels from the stem, and so they divide at different times.
Attenuation. Branches become attenuate by reduction in the number of
hyphae composing their tips. The farther they grow from the food-supply,
or the more tissue they build with a given food-supply, the more attenuate
the branches become. So ultimately, in most species, the branches taper to
undivided, subulate, or filiform tips, which may be only 20-50 µ, wide in those
without inflation of the hyphae, and even reduced to a single leading hypha.
Such slender tips have often given an erroneous idea of the genus Pterula,
the branches of which derive their filiform shape chiefly from their dimitic
construction without inflation. The phenomenon shows how the apical growth
of the branch works: on reduction of food-supply, apical growth in the hyphae
is not diminished (until the very end), but the branching of the hyphae falls off
so that the branch of the fruit-body is less able to thicken, is no longer able to
divide, even by dichotomy, and finally tapers.
Lengthening Internodes. The massive main branches are short and
polychotomous. The intermediate branches are l0nger and progressively
dichotomous. The ultimate branchlets are elongate, tapered, and simple.
Obviously this progressive elongation of the internodes, so implicit in the
branched Clavarioid fruit-body as to be detected only with difficulty, is the
expression of attenuation. The farther from the food-supply, or the shorter
it becomes, the longer time it takes for the new branches to thicken and reach
the threshold of division. The falling ramification, the lengthening internodes,
and the thinning tips are clearly a geometrical expression of the multiaxial
growth of a saprophytic soma expanding away from its point of attachment.
Cristation. Branches tipped with many small rudimentary branchlets are
called cristate. Instead of dividing regularly the tips of the branches ate
resolved into many small, new growing-points, and others may be produced
adventitiously on the sides. The phenomenon is not simple attenuation but
attenuation at the limit of food-supply, coupled with efforts to branch, as
though the controlling factors of the fruit-body had broken down. It occurs
in radial as well as flattened branching and, particularly, in Clavulina.
Torsion. Unequal inflation of the hyphae and unequal thickening of the
hymenium may result in curving or twisting of the branches. It is generally
nnr1� A
f1Q�
Ar1�
B
C D
TEXT-FIG. 6. Cristate tips: A, Clavulina Leveillei: B, Clavulina cartilaginea:
C, Ramaria polypus: D, Clavulina gigartinoides: mag.
limited to a few, often partially aborted, peripheral branches, but in Scytino
pogon and several species of Clavulina it is very pronounced and may twist
the branches out of their original plane of origin. Normally interference of
branches is avoided by their attenuation, or decussation, in successive tiers,
but this end may also be achieved by distortion. Quite different is the
squarrose appearance of many dried specimens caused by unequal shrinking
in drying, yet fal�ly indicating an irregular state for the living fruit-body,
e.g. the twisted branches of Aphelaria and Pterula, when dried, and the
curled, entangled masses of Lachnocladium (caused by the unequally
thickened cortex or hymenium).
Sagging. In many fruit-bodies, small and large, the lower main branches
usually sag under the weight of those superposed on them: particularly the
outer branches, without mutual support, become outcurved. Large, caespi
tose fruit-bodies of Ramaria and Clavulina may almost fall apart for this
reason, and then develop strongly upcurved new branches and assume an
unusual form. Sagging of the outer branches may emphasize the conical
outline of the whole fruit-body.
Flattening. Excessive expansion of the hymenium may cause the branches
to become hollow or, more often, to become flattened and canaliculate, but
the feature is commoner in the simple fleshy fruit-bodies with less entangled
hypha_e. Flattening may also result from excessive thickening of the unilateral
hymenium in such genera as Clavulina, Lachnocladium, and Ramaria.
Phototropism. In some species the branch-tips are positively phototropic
and, if their fruit-bodies develop with strongly unilateral lighting, an obliquely
fastigiate arrangement of the branches may result. It seems, also, that the
length of the stem depends considerably on the depth of the origin of the
P.rimordium in humus, under logs, or in crevices in wood, and on the amount
of apical growth of the primordial shaft before it reaches illumination suitable
for the development of the fruit-body. Indeed, for some species of Typhula
(with simple fruit-bodies), correct illumination is essential for the production
either of the fruit-body or of a fertile hymenium. Little, however, is known of
the action of light on Clavarioid fungi.
Angular Divergence. Such expressions as acute, semilunar, wide, or
flattened to describe the r'orm of the axils need explanation and precision of
reference to young or old axils which may differ much in form. Older axils
are generally wider and rounder, and often somewhat flattened whether
branching is radial or not.
The shape of the axil depends, firstly, on the divergence of the new branches
which are directed obliquely outward and later become vertical and nega
tively geotropic. The primary divergence may continue merely the oblique
direction of the outer hyphae in the expanded apex of the parent branch or it
may be a �utual repulsion, comparable with suppression, followed by rec
tification. A wide divergence with slow rectification, as by gradually increased
inflation of the abaxial hyphae, would give the semilunar axils of Clavulinopsis
corniculata. Generally the divergence is slight and the rectification rapid, as
in Ramaria stricta. On the other · hand the inflation of the hyphae and the
thickening of the hymenium both approximate the bases of the branches and
tend to obscure the axils.
Obliqueness. Unless deviated by light, Clavarioid branches grow vertically
upright after their initial divergence. If branching is rapid at first, with
short internodes, the lower branches may remain permanently oblique,
there having been no time for rectification. In such ca�,es the upper side
remains sterile and the hymenium is unilateral. In this respect Thelephora
again offers contrast, for in the species intermediate in form between the
Clavarioid and the Stereoid the growth of the branches remains oblique: they
become narrowly spathulate or cuneate and develop the hymenium only on
the lower side.
Thus the branched Clavarioid fruit-body consists of a series of growth-

IO GENERAL CONSIDERATIONS
pulses. The first forms the stem or axis, subsequent pulses ithe successive
tiers of branches. The perfect form, so nearly achieved in Clavicorona, is
modified by peripheral abortion, internal suppression, successive branching,
torsion, and sagging, and, through the manner of food-supply, it is gradually
attenuate. It is a very remarkable geometrical form to be analysed in terms of
continued discretion of the growing-point on reaching a threshold of size,
and thus it differs profoundly from the coherent centrifugal growths of e5ther
basidiomycetes.
SIMPLE FRUIT-BODIES
The simple fruit-body corresponds with the first growth-pulse of the
branched fruit-body, and is therefore homologous with the stem. It intro
duces no other feature than limitation of growth to the extent that the
threshold of branching at the primordial apex is never reached. Accordingly,
as a quantitative difference, many intermediate stages occur between sparingly
branched and simple fruit-bodies in the genera Clavaria, Clavulina, Clavuli
nopsis, Pterula, and Clavicorona, and, even within specific limits, as Clavaria
Zollingeri, Clavulinopsis corniculata, and Pterulicium xylogenum. Nevertheless,
it should not be overlooked that simple fruit-bodies are exceptional or absent
in Aphelaria, Scytinopogon, Clavarioid Thelephora, Lachnocladium, Rama
ria, and Lentaria, and that branched fruit-bodies are exceptional in the whole
Clavariadelphus-series, ev.en though the primordia of Clavariadelphus itself
seem massive enough. The transitional states need closer analysis: in
Pterula it appears that apical branching gives place to adventitious, and in
Clavulinopsis that apical branching gives place to the caespitose habit.
Similarly in Ramaria, reduction in size of the primordium produces caespitose
fruit-bodies with slender stems, e.g. R. conjunctipes and R. sinapicolor, and
may lead the way to simple caespitose states.
The caespitose character is often emphasized as if specific. Under good
conditions for growth it may be useful, no Clavariadelphus, for instance,
being caes_pitose, but fruit-bodies which are normally caespitose may be
developed from more or less impoverished mycelia and show all transitions
through the subfasciculate state. to the gregarious and solitary, e.g. Clavaria
vermicular.is, Clavulinopsis amoena, and many species of Ramaria.
HYMENIUM
Position. The stem and branches have typically a radial construction and
are covered entir..ely by the hymenium. When the branches are oblique,
however, either through their eai:ly divergent growth or through displacement
by crowding or phototropic direction, the hymenium develops only on the
underside, and on the upperside there is an incipient sterile hymenium, visible
under a hand�lens as a finely villous or subtomentose surface. The unilateral
hymenium has been considered a generic feature, as for Lachnocladium, but
it has no such value: it occurs in species of Ramaria, Ramariopsis, Clavulina,
Scytinopogon, and Lentaria and may not be even a constant specific character.
DESCRIPTION OF THE CLAVARIOID FRUIT-BODY n
The hymenium does not develop on the lower part of the stem where it is
still more or less immersed in the humus and, even in primordia Cwhich are
free from the first, it may not appear until the primordium is aJvanced in
growth and a sterile stem has been formed, e.g. Typhula, Clavulinopsis, &c.
Only some simple epiphytic species of Pterula, Deflexula, Ceratellopsis, and
Mucronella have truly sessile fruit-bodies, fertile from the very base.
Oh cessation of apical growth the hymenium usually develops over the tips
of the branches or of the simple fruit-body, even in the remarkable Ramaria
capitata the tips of which expand into Leotia-like knobs. The filiform tips of
Pterula and its allies, of Aphelaria, Ceratellopsis, Mucronella, some species
of Typhula, and, perhaps, Lachnocladium are persistently sterile. In Clavi
corona the expanded, subcyathiform tops of the branches are sterile, and in
tli.e Clavariadelphus-series there are some remarkable subpileate forms. Thus
Clavariadelphus truncatus and C. unicolor, Araeocoryne and Caripia have
truncate sterile tops, but no marginal or centrifugal growth to form a pileus.
In Hormomitaria and some species of Physalacria the base of the head is
sterile. In other species of Physalacria the head is wholly fertile, but in P.
infiata it seems that sterile patches of hymenium develop on the uppersides of
the ageotropic fruit-bodies, and may thus be basal, apical, or lateral according
to the direction of the fruit-body.
Occasionally and abnormally the hymenium may be absent or patchily
developed and some simple filiform fruit-bodies of Pterula and Ceratellopsis
may be completely devoid of hymenium.
Development and Thickening. The hymenium develops shortly behind the
growing-point by the sudden outward turning of the tips of the superficial
hyphae. A vacuole forms at or near the hyphal tip and distends a protube
rance to the exterior. The protuberance lengthens and becomes cut off by a
septum: either it becomes a basidium ( or cystidium) immediately or it pro
duces a few subhymenial cells and then becomes the basidium. After the apex
of the superficial hypha has turned outward its subterminal cells likewise
branch to the exterior and end in basidia: laterals may also be contributed by
the longitudinal hyphae which lie at short distances from the surface. The
hymenium is soon compacted into a continuous layer by the addition of these
laterals and by branches from the subhymenial cells. In some cases the
hymenium remains as a single layer of basidia, more especially in small
fruit-bodies as those of Typhula, Pistillaria, simple species of Pterula, and
so on. In many species, perhaps even in rriost, the hymenium continually
thickens throughout the life of the fruit-body by sympodi'11 branching from
the subhymenial cells. The branches shortly exceed the preceding basidia
and so build out the hymenium and, in these cases, not only do the old
collapsed basidia remain visible at various depths in the layer but numerous
spores, which have fallen on to the surface, become embedded.
The systematic importance of these two states of the hymenium has yet to
be evaluated because there are still many species in which it is not known
if thickening occurs. They are variously present in Clavaria, Clavulinopsis,
Ramaria, and Lachnocladium, but the thickening hymenium is a constant
feature of Clavulina and Clavariadelphus, in which it affords the chief dis
tinction fram Typhula and Pistillaria. A difference between Clavariachaete
TEXT-FIG. 7. Apex of a branch of Pterulicium xylogenum,

showing the origin of the hymenium, X r,ooo.
and Lachnocladit1..m may also be the absence of thickening in the former,

which seems to develop a single layer of basidia on a thick subhymenium.
In Lachnocladium the hyme:iium is often discontinuous for a "long while
until it begins to thicken. In Caripia it seems to be endogenous, but the
details of development are not known. In Chaetotyphula it arises first at the
apex of the minute fruit-body and then develops basipetally on to the stalk:
the genus seems to offer a limiting condition to that of Pistillaria, in which
some bas1petal extension may take place at the base of the fertile head.
DESCRIPTION OF THE CLAVARIO ID FRUIT-BODY 13
HYPHAE
Most fruit-bodies are monomitic, or with the hyphae of one kind, but the
distinguishing feature of Pterula and its allies is a dimitic state with thick-
: I
I !
.
�
.
±
�
TExT-FIG. 8.
�±
Hyphae and hymenium of Clavulinopsis amoena:
longitudinal hyphae inflating, interweaving hyphae not inflating:
hymenium beginning to thicken, X 500: clamps, X 1,000.
walled, more or less unbranched, colourless, skeletal hyphae. 1 Among the

monomitic states it is necessary to distinguish those
(i) with normal inflation and with clamp-connexions, as Ramaria, Rama
riopsis, Lentaria, Clavulinopsis, Clavariadelphus, &c., and most species of
Clavulina;
(ii) with inflation and secondary septation without clamps, as in Clavaria
and some species of Clavulina;
1
The tenns monomitic, dimitic, and trimitic for fruit-bodies composed of one kind of
hyphae, two kinds (skeletal and generative), and three kinds (skeletal, binding, and generative)
were introduced by me in 1932 (Tr. Brit. Myc. Soc. 17, p. 51). The trimitic state does not
occur in Clavarioid fungi.
(iii) without inflation, as in Aphelaria, Scytinopogon, Thelephora,
Clavariachaete, Ceratellopsis, and Mucronella;
(iv) without inflation, but with dichophyses, as in Lachnocladium sensu
stricto.
TEXT-FIG. 9. Basidia and clamps of Clavulinopsis fusiformis, X 1 ,ooo.
Inflation means that the cells of the hyphae begin to enlarge behind the
growing-point, and become wider and much longer, thus hastening the
apparent growth, or increase in size, of the fruit-body. It is the usual method
of growth in fleshy fungi. Usually the hyphae remain thin-walled, but in
Lentaria and the Stricta-group of Ramaria they become distinctly, and
characteristically, thick-walled.
In Clavaria the hyphae of the fruit-body have no clamp-connexions and
their cells, on inflation, form secondary septa which, as broad thin membranes,
joint them into shorter sections which, in their turn, enlarge and bec'c)me
septate. Thus typically in the genus, instead of the usual long-celled longi
tudinal hyphae or more or less entangled hyphae, there are parallel longitu-
dinal rows of short, sub-cylindric, inflated cells which cause the very brittle
texture diagnostic of the genus. In some species of Clavulina the hyphae also
lack clamp-connexions and become, in small degree, secondarily £eptate.
TExT-FIG. 10. Basidia and hyphae, without clamps, of Clavaria Zollingeri,

the longitudinal hyphae secondarily septate: basidia and spores,
X 1,000: hyphae, X 500.
In the genera with uninflated hyphae, the hyphal walls usually become more
or less thickened and so give support in the absence of turgidity. The growth
of their fruit-bodies is entirely apical, and therefore slower and needing a
longer period of suitable weather to reach as large a size. lt is not surprising
that they should belong to the tropical rain forests rather than to the tem
perate forests where the opportunity for prolonged apical growth is much
less frequent.
The dichophyses of Lachnocladium, described on page 8 5, are dichoto
mizing hyphae with limited growth and have thickened yellow-brown walls.
They are tissue-elements, not hymenial elements as are the simple setae of
Clavariachaete.
In the dimitic Pteruloid-series the skeletal hyphae are accompanied by
uninflate<;l., thin-walled, generative hyphae, mostly with clamps. The gene
rative hypnae build the hymenium, and the skeletal hyphae form the bulk
of the longitudinal, or medullary, hyphae and provide the mechanical support.
. :
TExT-FIG. 11. Basidia and subhymenial hyphae of Clavaria Gibbsiae (Clavaria subg.
Holocoryne): basidia with wide loop-like clamps, X 1,000.
The skeletals are mostly well differentiated, yet there are numerous transitions
between them and the generative hyphae. The dimitic fruit-bodies have also
only apical growth and their large bushy states are tropical.
In the little-known Lachnocladium reticulatum there appears to be another
dimitic state with much narrower skeletal hyphae.
There are three noteworthy anomalies. In the dimitic Pterulicium there
are skeletal hyphae in the erect Pteruloid fruit-body, but none in the fertile
resupinate or Corticioid patch which develops round the bases of the erect
fructifications. In contrast, the erect fruit-bodies of Ramaria stricta are mono-
mitic, but there are skeletal hyphae in the sterile floccose subiculum by which
they are attached to the wood. In Clavicorona candelabrum there arr.: 'skeletal
TEXT-FIG. 12. Hyphae (uninflated) and hymenium of Aphelaria dendroides:

no clamps, X 500.
hyphae' of limited growth in the erect fruit-body, where they are probably
modified gloeocystidia, but the mycelial subiculum, attaching the fruit
bodies as in Ramaria stricta, is monomitic. Such details need elucidation and
comparison with other species.
Agglutination. On the surface of the slender stems of many of the small
5119 C
GENERAL CONSIDERATIONS
fruit-bodies in the Clavariadelphus-series the hyphal walls become slightly
thickened and stuck together, or agglutinated. Thus a supporting cylinder
.
'
\. · .
TEXT-FIG. 13. Dimitic hyphae of Pterula verticillata: normal skeletal-hyphae,

on the left: a, branching tip of a skeletal hypha: b, generative hyphae arising
from skeletal�: c, a slightly inflated generative hypha: d, a generative hypha
widening into a skeletal: e, a generative hypha with thick-walled portion, x 500.
is formed to bear the weight of the head, e.g. Clavariadelphus fistulosus and
C. junceus, Typhula, Pistillaria, Physalacria, &c. In Typhula the agglutina
tion extends also to the sclerotium, as explained on page 169. In Clavaria
chaete the peripheral longitudinal hyphae develop dark-brown walls which
become agglutinated to form an internal black crust (below the hymenium),
and such is a common feature of the Hymenochaetoideae, to which it belongs.
, .-

Gelatinization. The gelatinous texture, caused by the gelatinization of the
hyphal walls, is often used as a generic character. Thus I have �spected
Myxomycidium, but I am doubtful if it has such value in other cases. Ramaria
gelatinosa is a typical Ramaria and its gelatinous state is bridged by R. sub
gelatinosa, which connects it with the dry-fleshed R. formosa. Similarly R.
capitata has capitate gelatinous tips to the branches, but R. ochraceosalmoni
color is said to connect it with the normal state of the genus. Clavicorona
j(lf)anica has translucent white fruit-bodies suggesting Tremellodendron, but
it has all the characteristics of its genus: it is also unusual in that its gelatinous
hyphae inflate, this modification of the wall usually preventing the develop
ment of turgidity. Hence I do not consider that the gelatinous texture of
Aphelaria flabellata, Clavulinopsis calocera, C. hastula, and C. spathuliformis,
Lentaria coronilla, and some species of Typhula and Pistillaria deserves
generic rank, though additional reasons may be found when the species are
better understood.
Clamp-connexions. It is often assumed that clamps occur on the hyphae of
all Homobasidiomycetes. This is by no means the case. The absence of
clamps so distinguishes the Xanthochroic series (p. 22) that their presence
is proof that a species does. not belong. In other groups the criterion is not
so precise and it can be used only after critical study of allied species. Thus
Clavaria appears to be characterized by the absence of clamps, for which
reason I separate C. fossicola from Lentaria mucida, but in the subgenus
Holocoryne there is a wide loose clamp at the base of the basidium. Clamps
occur in most species of Clavulina, Ramaria, Pterula, and Typhula, but are
absent from others: on the other hand they are typically absent from Aphe
laria and Ceratellopsis. In a few species of Clavulina and Typhula clamps
may occur rarely on some hyphae and not on others. Whether there are
many instances of clamped mycelia and fruit-bodies without clamps (as in
many species of Coprinus), I do not know, for I have studied chiefly the fruit
body, but Typhula trifolii and Clavulina cartilaginea are certainly examples.
There are no instances of multiple clamps as in Coniophora cerebellR. I have
given the details, so far as known, under each genus, but have limited them
mainly to my own observations: published records of the presence or absence
of clamps in Clavarioid fungi are not always correct.
The shape of the clamp varies considerably and may even have some
generic value. In many species of Clavulinopsis, for instance, it is a distinct
loop very much as the wide loops which give the forked base to the basidium
in Clavaria subgen. Holocoryne. In some cases the clamp inflates as well as
the rest of the cell and can, then, be detected only with care. Critical study of
the clamps in these genera may throw some light on the problem of clamps in
general.
CYSTIDIA
Most Clavarioid fungi have no cystidia. They occur as thin-walled c;:ysti
dioles in a few species of Pterula, Clavaria, Caripia, and, perhaps, Pistillaria,
Ramaria, and Clavulinopsis. Clavulina Leveillei is unique in having very
large thi<.:k-walled cystidia, unknown in other species of the genus, yet it has
a variety without them. Thick-walled cystidia occur in some species of
Physalacria and they �istinguish Chaetotyphula and Dimorphocystis, which is
a remarkable Pteruloid genus of tropical Pistillaria-like species with two kinds
of cystidia. · Hymenochaete-setae, with thick brown walls, mark Clavaria
chaete as a unique Clavarioid genus of the Hymenochaetoideae.
A B C r C
TExT-FIG. 14. Spores: A, brown-walled guttate spore of Ramaria Zippelii
with hyaline spines: B, thin-walled granular spore of Clavariadelphus pistillaris:
c, thick-walled multiguttulate spore of Clavaria Gibbs;ae: D, thin-walled vacuo
late spore of Physa[acria tropica: E, brown-walled 1-guttate spore of Ramaria
ochraceo-virens, with colourless warts: F, thick-walled 1-guttate spore of Clavu
lina: G, thin-walled aguttate spore of Clavaria with opalescent cytoplasm, mag.
Lachnocladium and Clavicorona have gloeocystidia, which are somewhat

enlarged sterile basidia with oily contents: they are generally thin-walled but,
as they elongate through the thickening hymenium, they may become thick
walled in their older, and usually vacuolated, proximal parts.
Physalacria and Hormomitaria have oleocystidia, which are clavate, often
capitate, cystidia with an oily resinous exudation at the apex. The exudation
is interlamellar and covered by the thin outer layer of the cystidium-wall: it
dissolves in dilute alkali.
The slender fruit-bodies of the Clavariadelphus-series and the corre
sponding ones of the Pteruloid-series often have characteristic caulocystidia.
In the other simple fruit-bodies, as those of Clavaria, Clavulina, and Clavuli
nopsis, the stem, if not glabrous, has merely scattered rudimentary basidia
or fi:liform hyphal ends which merge downwards into the secondary mycelial
hyphae at the base of the stem.
HABITAT
Most species with large fruit-bodies grow in the humus of forests. Many
with simple fruit-bodies, but few of those with branched fruit-bodies, occur
in open grassland, heaths, moors, or mossy places. In most humicolous
genera there occur also a few lignicolous species, but this habit seems to
distinguish Lentaria, Clavicorona, Deflexula, Caripia, and Mucronella. 1
1
Trunks of monocotyledons, as well as dicotyledons, must be regarded as comprising
the lighicolous habitat.
The small fruit-bodies of Typhula, Pistillaria, Ceratellopsis, &c., in the

Clavariadelphus-series and those of the simple Pterulas grow on dead leaves,
herbaceous stems, twigs, dead fruits, and so on, and may be usefully described
as epiphytic in the saprophytic sense. There are very few instances of the
epiphytic habit in humicolous genera, e.g. Clavulinopsis miniata var. san
guinea, C. luteo-tenerrima, Clavariadelphus fistulosus, and C. junceus.
A peculiar habitat is the phycophilous. The mycelia of a few species with
small, simple or very sparingly branched fruit-bodies grow in films of green
or blue-green algae which coat the soil or the surface of rotting logs (Lentaria
mucida). So far as known, they are:
Clavaria fossicola and C. helicoides, Clavulinopsis hastula and C. vernalis,
Lentaria mucida, L. coronilla, and, perhaps, L. affiata.
They appear to be more or less parasitic on the algae, for their hyphae are
closely applied to the algal cells, even with haustoria (C. vernalis), or they
may kill groups of them (C. fossicola). The species resemble each other very
much in the small simple form of the fruit-body and the small spores, as well
as in this peculiar habitat, but hyphal details show that this similarity is an
effect of convergence. Thus Clavulinopsis constans is non-phycophilous but
in other respects almost identical with C. vernalis; Lentaria mucida (= Cla
varia mucida) seems to be connected with the Lentaria-Ramaria-series; and
Clavaria helicoides is a true Clavaria with secondarily septate hyphae. The
habitat is clearly as secondary as the bryicolous or the habitat of certain
Pyrenomycetes on marine algae. In no case do the algal cells enter the con
struction of the fruit-body.
The only Clavarioid fungi known to be parasitic on higher plants a.re a
few species of Typhula.
COLOUR
It is generally recognized that the colour of the fruit-body is much more
important in the systematy of fungi than of flowering plants and that the
Linnean dictum Ne crede colori should not apply in mycology. Very little is
known about the microscopic occurrence of pigments in the fruit-body, yet it
may be as useful a diagnostic character as any. I have discussed the ,Problem
under the different genera, particularly Ram.aria, and give here only some
concise terms for description.
Euchroic: with true pigmentation in coloured granules, crystals, guttulae, vacuoles,
and so on.
Achroic: without true pigmentation.
Hysterochroic: mature fruit-bodies becoming slowly discoloured acropetally (from
the base of the stem upward) by coloration of the hyphal walls, probably caused by
slow oxidation in senescent tissue (e.g. the blackening, browning, or yellowing of
fruit-bodies of Marasmius, Fornes, Lentinus, and Clavulinopsis).
Epichroic: discoloration on bruising or cutting, as reddening, yellowing, blueing,
or blackening: an oxidation as in hysterochroism, but more rapid and rarely developed
spontaneously, though often shown also in old fruit-bodies as a result of battering by
rain or of drying.
PARTICULAR KINDS OF EUCHROISM
Acrochroic: coloured specially at the hyphal tips in the growing-point (e.g. species
of Ramaria)..
Metachroic: changing colour by the appearance of a new pigment in the older tissue
(not merely epichroic or hysterochroic), as in most acrochroic species of Ramaria.
Ectochroic: with the pigment excreted on the outside of the hyphae as a granular,
amorphous, annular, or plaque-like incrustation, e.g. most ochrosporous agarics,
Marasmius, Boletus, Lachnocladium aurantiacum, Scleroderma.
Endochroic: with the pigment 'inside the cell.
Mesochroic: with the pigment in the hyphal wall, as in the Xanthochroic-series of
Lachnocladium, Hymenochaete, Fornes, &c.
PARTICULAR KINDS OF ENDOCHROISM

Cytochroic: the pigment diffused in the cytoplasm.
Cystochroic: the pigment in the cell-vacuoles.
Lipochroic: the pigment in oil-drops, or guttulae.
AFFINITY OF CLAVARIOID GENERA

(vide Natural Key, p. 172)
Xanthochroic-series. Uninflating hyphae without clamp-connexions are a
constant feature of a series of basidiomycetes which I call the Xanthochroic
because the fruit-bodies are typically yellowish, ochraceous, or brownish
yellow. The hyphae may be monomitic, in which case their walls generally
thicken and become pale yellow or brownish ochraceous under the micro
scope, as in Polystictus perennis and Inonotus (Polyporus) hispidus, or
they may be dimitic, in which case the skeletal hyphae have the strongly
thickened, yellow-brown walls, as in many species of Fornes (F. senex, F.
pachyphloeus). Further, the hyphae may develop, in the hymenium, cystidia
with thick brown walls, usually called setae, as in Hymenochaete and Fornes
spp., and they may also develop in the flesh stellate setae (Asterodon, Astero
stroma) or: dichophyses (Astrostromella, Hymenochaete pr. p.). In all cases
the thickened, yellowish or brownish walls darken perceptibly and charac
teristically under the action of alkali. This darkening and the absence of
clamps always enables one to recognize a member of the series in the absence
of se�ae, dichophyses, or skeletal hyphae. Further, the yellow-brown colour
and direct growth of the fruit-bodies consequent on the colour of the hypha
walls and the absence of inflation usually enable one to distinguish the series
macroscopically. )t has many polyporoid members, a few hydnoid (Astero
don), a few stereoid (Hymenochaete), a few corticioid (Asterostroma, Astero
stromella), and, now, I add two clavarioid genera, Lachnocladium and
Clavariachaete. It is one of the most natural and well-defined series of the
Aphyllophorales (gills are not known in it, but concentric lamellae occur in
Cyclomyces), but what its total range and, therefore, what its correct name
cannot yet be stated. It is the group referred to by Donk as the Hymeno
chaetoideae: eventually it must be raised to the rank of a primary series of the
AFFINITY OF CLAVARIOID GENERA 23
Homobasidiomycetes, and the name Xanthochroales would seem more
suitable.
Lachnocladium belongs to the subseries with dichophyses, instPad of s-etae
or skeletal hyphae. Asterostromella is the Corticioid member and I have
found in Malaya a terrestrial Stereum form with pleuropodal, flabellate,
ascending pileus. Besides the dichophyses, gloeocystidia are characteristic,
andthe hymenium, initially, is discontinuous and consists of scattered basidia
and gloeocystidial tips projecting from the forest of dichophyses: if the hy
menium thickens, as usual in Lachnocladium, it develops a eontinuous layer
of basidia into which the gloeocystidia extend, but without dichophyses. The
spores are colourless, though the wall may turn yellowish in drying, and the
hyphae are monomitic. No instance is known of the Clavarioid state of
Lachnocladium passing into the Stereoid as occurs in Thelephora.
Clavariachaete belongs to the true Hymenochaetoideae with thick-walled
setae, but without dichophyses or gloeocystidia: it appears, also, that the
hymenium does not thicken in this group. The hyphae are monomitic, as in
Clavariachaete and some Polyporoid forms (as Polystictus), or dimitic as in
many Fornes forms and, perhaps, as in Hymenochaete. Mesopodal species of
Hymenochaete are known from tropical America and, possibly, intermediates
may be found there to connect with Clavariae:haete. The carbonaceous
cylinder in the flesh of Clavariachaete confirms its affinity with Hymenochae
toideae.
As yet, I have recognized no Clavarioid form of the Xanthochroic-series
without dichophyses or setae. Possibly Aphelaria aurantiaca, also from tropi
cal America, is such. Lachnocladium reticulatum, also from tropical America,
may be a dimitic Clavarioid state with skeletal hyphae but, so far as I could
make out from the type-specimen at Kew, its hyphae were colourless and did
not darken appreciably with dilute potash, though the spores were brown (cf.
xanthochroic Fornes): I could not find the generative hyphae to see if they
had clamps. But, clearly, in tropical America one must expect some remark
able additions to our knowledge of the Xanthochroic-series.
Pteruloid-series. The slender branches and the filiform simple fruit-bodies
of this series owe their shape to the narrow growing-points and the dimitic
construction without inflation. The skeletals have colourless walls and the
generative hyphae are typically clamped. Thus the series differs fundamen
tally from the Xanthochroic and appears related to another Stereum-Poly
porus-series of dimitic Aphyllophorales. I have seen no connecting form
and the Pteruloid-series seems as isolated as any among Homobasidiomycetes.
(See p. n5.)
Thelephoroid-series. Scytinopogon is so similar to Thelephora s. str.
(= Phylacteria) that, if its spores were fuscous, it would be congeneric. The
angular shape of the spore is characteristic of both, but the feature is less
marked in Scytinopogon which also has less strongly echinulate spores.
Peculiar of Scytinopogon, also, is the branching in one plane: it seems another
variation of the tendency in Thelephora to form palmate-flabellate and
Stereoid fruit-bodies. Thelephora degenerates so readily into resupinate forms
(Tomentella) that it is practically impossible to separate them generically.
I have seen no such forms of Scytinopogon, but there are several Corticioid
fungi with ellipsoid echinulate spores which may belong. Polyozellus
is another allied genus with large, erect, Stereoid fruit-bodies and colour
less spores, _;is in Scytinopogon. As Donk has shown, under the Phylacter
oideae Thelephora is closely related with some Hydnoid genera as Phellodon,
Hydnella, and Sarcodon, and perhaps v.1ith some peculiar Polyporoid forms
(Boletopsis). They may represent another separate series of Homobasidio
mycetes to which, it seems, the name Thelephoraceae (or Thelephorales)
must be restricted.
The position of Aphelaria is problematic. It has not the spores of the
Thelephoroid-series and, as typically defined, the uninflated hyphae without
clamps suggest affinity with the Xanthochroic-series, especially A. aurantiaca,
as already mentioned, for its hyphal walls show the characteristic darken
ing in dilute potash. The genus may be composite. I have placed it with the
Thelephoroid-series because it resembles Thelephora in texture and general
form rather than Clavaria.
Ramaria-series. Ramaria is remarkable among the more typical Clavarioid
genera, as Clavaria, Clavulinopsis, and Clavulina, because it has no simple
derivative. It closely resembles Clavulinopsis in general construction, but the
coloured spore-wall is distinctive and holds throughout the great range of
specific detail, for Ramaria is the largest Clavarioid genus. This is an impor
tant fact which convinces me that, in spite of the apparent resemblance to
other genera (cf. Clavulinopsis corniculata), Ramaria is fundamentally differ
ent. Thus, as Donk remarks, the Cantharelloid genus Gomphus (and Can
tharellus fioccosus) with oblong ochraceous spores may be allied. I have also
found in Malaya simple and compound pleuropodal Stereoid forms with the
same spore. Thus I think that future discoveries may prove that Ramaria
belongs to a subseries, at least, of the Homobasidiomycetes with inflating
monomitic hyphae.
The position of Lentaria is not clear. There is a great resemblance between
L. byssiseda or L. surculus and the Stricta-group of Ramaria and, if it implies
affinity, then the character of the coloured spore for the Ramaria-series breaks
down because L. surculus, if not L. byssiseda, has pure white, thin-walled
spores. The connexion through such species as L. epichnoa with L. mucida,
however, indicates that Lentaria may be a lignicolous parallel to Ramaria and
derived possibly from a branched Clavarioid state ancestral to the Clavaria
delphus-series. Thick-walled hyphae, as in Lentaria and the Stricta-group of
Ramaria, are common in lignicolous fungi, e.g. Pleurotus, Stereum, Poly
porus, and do not necessarily indicate affinity.
Clavariadelphus-series. Contrasted with the preceding series, this is remark
able for the absence of branched fruit-bodies, the few instances of branching
being adventitious proliferation, not apical branching. Its characters appear
negative, merely Clavarioid, and thus it is difficult to define; in all probability
it is a mixed group derived by convergence to a- reduced Clavarioid state
from several sources. Until, therefore, the position of the Clavar_ioid state
in the evolution of fungous fruit-bodies is known, it will be difficuit to assign
the genera to their rightful position. There appear to be thre� elements, at
least, in the series.
Firstly, the genera Clavariadelphus, Typhula, Pistillaria, and Pistillina are
so dosely linked that their limits are almost arbitrary whatever system is
followed for their classification. Clavariadelphus junceus is the remarkable
link with Typhula and Pistillaria: it has the slender form of fruit-body, the
agglutinated superficial hyphae of the stem, and the epiphytic habitat of
Typhula and Pistillaria, but it has the thickening hymenium of Clavariadelphus
and it is connected by C. fistulosus with the typical massive form of the genus:
yet in its rhizomorphs and basal hairs it has the makings of the sclerotium of
Typhula. Pistillina is one extreme with discoid hymenium. At the other
extreme are C. pistillaris, C. truncatus, and C. unicolor, which connect with
Cantharellus. The problem to be solved is whether the pileate Cantharellus
has evolved from Clavariadelphus by the evolution of margin growth and so,
perhaps, produced an agaric series, or whether Clavariadelphus represents
the Cantharelloid state reduced to the primordial shaft which has lost the
faculty of marginal growth to extend the hymenium. If the first alternative
is true, then one must look for a branched ancestral Clavarioid form, such as a
thin-walled humicolous Lentaria or an aguttate Clavulinopsis. If the second
alternative is true, the few branched Cantharelloid fruit-bodies which occur
in some tropical species may represent the ancestral form. In either case this
main subseries is parallel with the independent Clavaria-series; such products
as Lentaria mucida, Clavariadelphus fistulosus var. contortus, or Pistillaria
setipes, and Clavaria filiola resemble each other through convergence.
Typhula, as explained on page 169, presents the extreme, xerophytic and
low-temperature, modification of this mainly temperate subseries.
Secondly, Physalacria and Hormomitaria represent an end-product of a
Marasmioid-series or of some unknown Clavarioid-series, as discussed on
page 97. The characteristic cystidia should prove their .:onnexion with any
other fungi.
Thirdly, Mucronella seems to be, atleast taxonomically; a reduced Hydnoid
genus, but it has been confused with Pteruloidfungi (Deflexula) and, probably,
with Ceratellopsis and Hormomitaria. Ceratellopsis is truly Clavarioid, with
erect fruit-bodies: it may represent the simplified, epiphytic derivatives of
Aphelaria (though this genus is tropical and Ceratellop5is appears to be
temperate), or it may be derived from the Clavariadelphus-series, as Pistillaria
without inflation. It offers a most remarkable instance of homoplasy or of
convergent evolution, for, so far as can be seen, its fruit-bodies differ from
those of Mucronella only in direction of growth.
The remaining four genera are problematic. Caripia may be reduced Len
taria with sterile top or it may be derived from some Cantharelloid genus and
have independently developed thick-walled hyphae, as a lignicolous genus.
Similarly the sterile top of Araeocoryne suggests a Cantharelloid ongm,
but its texture is that of Omphalia. Myxomycidium may be related with
Clavariadnlphus, Lentaria, or Clavulinopsis, as a simple gelatinous derivative,
or it may be the relic of gelatinous Clavarioid fungi, if that was their primitive
condition. Chaetotyphula appears to be Pistillaria with basipetal head and
precocious cystidia, very much as Dimorphocystis is to the simple species of
Pterula.
The problems of this series, or of these subseries, hinge on those of Can
tharelloid fungi with which one must consider at the same time the fleshy
Stereums which occur in strange, if rare, variety in the tropics.
Clavaria-series. The four genera Clavaria, Clavulina, Clavulinopsis, and
Ramariopsis have hitherto been regarded either as one, that is, the typical
state of Clavaria, or as two, with Clavulina distinguished by its 2-spored
basidia. It is clear from their hyphal construction that there are three sub
series, Clavaria, Clavulina, and Clavulinopsis with Ramariopsis. Each ranges
from branched to simple forms of fruit-body with considerable specific
variation.
Clavulinopsis and Ramariopsis have unspecialized, monomitic, clamped,
inflating hyphae and white, usually 1-guttate, spores. They appear as a central
group of humicolous Clavarioid fungi from which lignicolous and epiphytic
Pistillaria-like forms may have been evolved (e.g. C. luteo-tenerrima on leaves),
but, if so, such forms do not appear to be common or numerous. Clavuli
nopsis, however, contains a variety of subseries, some of which may belong
rather to the Clavariadelphus-series. Ramariopsis differs only in the rough,
verruculose, or echinulate spore, and so stands to Clavulinopsis as the
Echinospora- and Verrucispora-groups of Ramaria do to its Decolorans
group: but taken as a whole Ramariopsis, also, appears as a natural genus
with little, if any, tendency to produce simple fruit-bodies. It seems to con
nect the Clavaria- and Ramaria-series to a common origin, from a radially
branched, inflated fruit-body with ornamented spores. The similarity in the
colours, particularly yellow, of the fruit-bodies of Clavulinopsis, Ramariopsis,
and Ramaria may be no coincidence.
Clavaria assumes a special position because of its secondarily septate
hyphae without clamps, The feature occurs in several genera of Homobasi
diomycetes (Tricholoma, Agaricus, Coprinus, Lepiota pr. p., Mycena pr. p.,
Clitocybe pr. p., and Craterellus pr. p.), so that it does not seem to be a prime
indication of affinity: its recurrence in Hormomitaria confirms this view.
One must, therefore, regard Clavaria as a specialization from a branched
Clavarioid stock which seems to be that, also, of Clavulinopsis. The presence
of wide, loop-like, clamps at the base of the basidia of some species of Clavaria
also indicates that they have been derived from a state with clamp-connexions.
Yet these �lamps are so oddly formed and so curiously placed at the base of
the sporangium, that I cannot help wondering if, in Clavaria, there is not a
relic of the primitive basidiomycete with incipient clamps and that its secon
dary septation may also be a primitive feature. In the other genera which I
have mentioned with secondarily septate hyphae there may be normal clamps
on the hyphae before secondary septation and on the basidia, or there may be
no clamps at all: I have never seen in them the loops of Clavaria.,
Finally the genus Clavulina introduces the problem of the direction of the
nuclear spindle in the basidium. I must defer the discussion of the Clavulina
basidium until it is possible to review the whole problem of the basidium
as a sporangium conformable to a hypha: this cytological distinction will then
be seen to have only minor importance. vVhen it is disregarded Clavulina
resembles Clavulinopsis in form, hyphae, and spores, but it tends to the
colouring of Clavaria and some of its species have secondarily septate hyphae
(but without the basidial loops of Clavaria). If a common ancestor can be
postulated for Clavulinopsis and Clavaria, then Clavulina unites them even
more closely.
Conclusion. The Clavarioid form of fruit-body is not, in itself, an indication
of affinity. The primary index is similarity in hyphal construction. The
Clavarioid form is a homoplastic state assumed by several, if not all, series
of Basidiomycetes (cf. Tremellodendron, Calocera). Thus the Clavarioid
Homobasidiomycetes assume a fundamental importance in the classification
of the group because their striking form must be linked through Thelephora,
Stereum, Cantharellus, and Hymenochaete with Hydnoid, Corticioid, Poly
poroid, and Agaricoid genera. Affinity between Clavariachaete and Fames
ferruginosus would never have been expected without hyphal analysis, yet
it is paralleled by Thelephora and Tomentella, by Lachnocladium and
Asterostromella and, most remarkably, by Pterulicium. Thus the secondary
septation of Clavaria, Craterellus sinuosus, Agaricus, and Tricholoma requires
as careful investigation and comparison as does the resemblance between
Physalacria and Marasmius or Clavariadelphus and Cantharellus.
GEOGRAPHICAL DISTRIBUTION
Except in central and western Europe and in parts of the United States,
collections and accurate determinations of these fungi have been too few for
one to give more than a rough indication of their distribution. As with most
saprophytic fungi they are influenced primarily by the effect of temperature
ar.d rainfall on mycelial growth, secondarily by habitat-specialization, and
not at all by any relation which the fruit-body bears to the climate. Thus the
subgenera, genera, and higher ranks are mostly cosmopolitan while the
microgenera comprise closely allied temperate or tropical species with or
without differences of habitat: for example, Clavaria acuta, Clavulinopsis fusi
formis, C. pulchra, and C. aurantio-cinnabarina, Clavulina rugosa and C. amethy
stinoides, and Ramaria formosa and R. stricta have their immediate tropical
allies just as Amanita muscaria, A. excelsa, A. rubescens, or A. vaginata have
each their own tropical allies, but their genera and subgenera are cosmopolitan.
Thus the distribution of the fungi differs essentially from that of the flowering
plants, the somatic growths of which are profoundly influenced by climate:
there are very few genera of higher fungi confined to the tropics, compared
with those of Angiosperms, and probably no families.
Regarding specific distribution one finds, accordingly, many species wide
spread throughout the north temperate zone or throughout the tropics and
that many of these north temperate species occur in south temperate regions,
at least according to the recent works on South Australian fungi. However, I
think the bizonal records need critical examination, because man has greatly
altered the natural vegetation in large areas of south temperate countries,
particularly in the neighbourhood of towns and other places most readily
accessible to the mycologist, and he may either have introduced north tem
perate species or have so altered the vegetation as to allow their spores to
settle and colonize new areas: one wants the assurance that such species
occur in virgin vegetation. A few species are widely tolerant and are cosmq
politan, as Clavaria vermicularis and C. Zollingeri, which occur in the lowland
tropics and are also characteristic temperate species. One can readily under
stand that short-lived tropical species can fruit in temperate summers, but I
am doubtful whether any tropical species with massive fruit-bodies are nor
mally temperate as well and that their mycelia can perennate through a
winter. Thus Lachnocladium and most bushy species of Pterula are tropical,
and there seems to be no authentic record of the temperate Clavariadelphus
or Typhula in the lowland (or mountain) tropics.
The ease with which fungous spores can be blown about implies wide
spread occurrence, though in detail distribution may be sporadic because of
particularities of habitat. Nevertheless it is not proven that all fungous spores
can withstand long periods of desiccation and insolation and, when one con
siders the primeval, densely forested state of most of the world suitable for
fungous growth, it is not obvious that the spores, once lifted up, would ever
have come down alive on suitable ground: particularly is this so in the very
mixed rain-forest of the tropics. Under such conditions it seems that the
spread of fungi dwelling on the forest floor is more likely to have been gradual.
As instan<:es of in�bility to spread widely through the air one may give
Caripia, so common, apparently, on wood and twigs in the American tropics,
but clearly absent from the Old World. Even in the north temperate zone there
are considerable differences between the saprophytic fungous flora of North
America and western Europe. Thus Clavulina ornatipes, widely distributed in
the United States, has been reported from Japan and I have found it in
Singapore: it is one of the most distinct species of Clavarioid fungus, so that
one can be sure that it does not occur in western or central Europe. The
Atlantic Ocean and the forestless tracts of outer China and the Middle East
have .clea�ly been as effective barriers by air as the Siberian forests have been
too cold: on the other hand, only the Behring Sea is the obstacle between
North America and Malaya. Similarly Deflexula ulmi of North America is
very near D. lilaceo-brunnea of Malaya and Sumatra: the genus occurs in
North and tropical America, in tropical Asia and Australasia, and in the Bonin
Islands, but not in central or western Europe. Again, Ramaria Zippe/ii of
'
GEOGRAPHICAL DISTRIBUTION 29
tropical Asia is very near R. grandis of North America, but has no such near
ally in Europe. Myxomycidium has also the distribution Tennessee, British
Guiana, and Tasmania.
The most obvious specialization in habitat of the hig�er fungi is that of
coniferous humus in contrast with dicotyledonous. The species of northern
coniferous forests, as Clavariadelphus ligula and Ramaria ochraceo-virens,
range widely, as one would expect, through the palaearctic coniferous belt:
they have not been found in the tropics, though they may be expected in the
mountain forests of Pinus in northern Sumatra, and their occurrence in
South Australia may be by introduction. It is unknown if there are any
species similarly related to the south temperate and tropical conifers. Whether
there is really any other host-restriction among, Clavarioid fungi is also un
certain because of paucity of collections, but there appears to be such among
the parasitic species of Typhula.
Genera. Clavariadelphus, Typhula, Pistillina, Ceratellopsis, and most
species of Pistillaria are north temperate (I have seen only two species of
Pistillaria in Malaya though I have often looked for them and found their
Pteruloid analogues). Typhula, in fact, seems the most highly specialized,
xerophytic, and low-temperature modification of any Clavarioid fungi and
thus fits physiologically, as well as morphologically, with Clavariadelphus.
Lachnocladium and most species of Pterula and Aphelaria are tropical.
Clavariachaete and Caripia occur in tropical America. Araeocoryne, Chaeto
typhula, Dimorphocystis, and Hormomitaria have been discovered, as yet,
only in south-east Asia.
Clavaria, Clavulina, Clavulinopsis, Lentaria, Mucronella, Ramaria, Ramari
opsis (and Thelephora) are cosmopolitan, but Physalacria, Scytinopogon,
Clavacorona, and, perhaps, both Deflexula and Myxomycidium are mainly
tropical.
Species. Clavaria Zollingeri, C. vermicularis, and possibly C. fumosa and
C. acuta are cosmopolitan. Clavulinopsis amoena and C. miniata may be, to9.
Pan.tropical are the very varied Scytinopogon angulisporus (? lso in the
southern United States), Clavulinopsis amoena and C. miniata, Clapulina
cartilaginea (West Indies, Malaya), Lentaria surculus, and, probably, Ramaria
cyanocephala or R. Zippe/ii. R. Moelleriana occurs in tropical Africa and
Asia and is to be expected in tropical America: perhaps, also, the Australasian
Aphelaria dendroides.
The north temperate Ramaria fiaccida, R. gracilis, R. stricta, R. fiavobrun
nescens, R. fumigata, &c., Clavulina cinerea, Lentaria mucid;i, Clavariadelphus
junceus, Clavulinopsis corniculata, C. pulchra, C. subtilis, and C. umbrinella,
Ramariopsis Kunzei, R. pulchella, and R. crocea, and Clavicorona pyxidata
have been found in South Australia.
Many species of Ramaria and Clavulinopsis, to a less extent of Clavaria
and Ramariopsis, have been found in North America but not in Europe,
though they may be expected in temperate Asia.
A few spectacular species appear to be local. Thus Clavariachaete and
Caripia occur in tropical America, Ramaria capitata and Ramariopsis lori
thamnus b South Australia, Ramaria gelatinosa and Clavariadelphus unicolor
in North America, and the. lignicolous Clavulina Sprucei, C. connata, and C.
delicia in Brazil.
The following north temperate species have extremely similar tropical
allies or representatives (named in brackets):
Clavaria acuta (C. Gibbsiae), Clavulina rugosa (C. gracilis), C. amethysti
noides (C. cavipes), Clavulinopsis fusiformis (C. amoena), C. aurantio
cinnabarina (C. miniata), C. pulchra (C. luteo-tenerrima), Lentaria byssiseda
(L. surculus), Ramaria nigrescens (R. fragillima), R. grandis (R. Zippelii),
R. stricta (R. Moelleriana), R. formosa (R. subgelatinosa).
Exploration. The tropical snow-mountains should yield a rich harves�.
Probably the Himalayas, uniting north temperate and tropical, palaearctic
coniferous and Indo-Malaysian broad-leafed forests, will prove the richest
collecting-ground. Very little is yet known of the fleshy Clavarioid fungi of
South America, Africa, and New Zealand. Araucaria-, Podocarpus-, and
Nothofagus-forests may have their peculiarities (as there is, for instance, a
Clavarioid Thelephora associated with the roots of Casuarina equisetifolia).
COLLECTION AND STUDY

Fleshy fungi, such as the large Clavarioid fruit-bodies, are often supposed
to be rare or poorly represented in the tropics. This is a mistake arising from
the paucity of collections. They are, at the right season, very abundant and
even more varied than in temperate countries, but they are a nuisance for the
ordinary collector to deal with and the resident mycologist has no means of
identifying them. I offer these notes to those who may wish to study the
fleshy fungi where the flora is still unknown, particularly to tropical myco
logists.
Firstly, Living specimens must be studied repeatedly. In the tropics the
fleshy fungi develop in great abundance all at once, very quickly, and decay
rapidly. The mycologist can rarely deal with all that he finds before they
have decomposed: it is certainly impossible to describe their macroscopic
and microscopic characters, to paint the living specimens, and to make satis
factory dried material at one time. Accordingly one must plan to study over
several seasons, if not years, in order not to miss the opportunities which, if
only for their bre.;ity, must be considered rare. I collect in the morning as
much as I can deal with before the light fails in the evening. I write colour
notes and other field-data (size, texture, smell, taste, &c.), make a few micro
scopic observations, then make spore-prints and dried specimens, and always
preserve a suitable specimen in alcohol-formalin: painting I leave for subse
quent seasons when I have more time and, being better acquainted with the
specimens, can select suitable ones with some leisure. That is to say, I
concentrate on the material for adequate description, particularly micro-
COLLECTION AND STUDY 31
scopic; for a painting in itself, however elegant, is not worth the time spent
upon it when it must be taken from the precious hours of collection.
The microscopic features which should be studied in fresh material are the
spore-contents, especially the presence of oil-drops, which cannot be seen
correctly in preserved specimens, and the microscopic distribution of the
pigments, particularly if ectochroic, mesochroic, or endochroic.
Dried material is more convenient to keep than that preserved in liquid, but
is far more difficult to re-examine microscopically, and often inadequate for
hyphal details or for dimensions of basidia, cystidia, and so on. For this
purpose I use the mixture of 25 per cent. alcohol and 5 per cent. formalin in
water which was recommended by Lagarde in his studies on Discomycetes
(Ann. Myc. 4, 1906, 125): formalin, by itself, causes too much gelatinization
of the hyphal walls ( cf. the 'formalin-genus' of Boleti, Ixechinus), and alcohol�
by itself, causes too much contraction: the mixture counteracts both effects
and keeps the material in good condition for 20 years, at least, in my ex
perience. When travelling it is convenient to have square wood-block cases
for the tubes of preservative. In any case, if material is to be preserved in
alcohol-formalin, it should be done as soon after collection as possible to
prevent damage to delicate tissues and to ensure that the basidia are sporing:
it often happens that Cla;arias, Stereums, Hydnums, and Polypores stop
sparing a few hours after collection, especially if not kept in their normal
upright position. A note of warning must be given with Pterulas: it seems
that they may spore only at night, so that specimens should be kept on a glass
slide for spore-prints overnight.
IDENTIFICATION
IN the main key on page 172 I have arranged the genera in what seem to be
their natural groups. The key will enable mycologists to arrive with certainty
at the species which are adequately known. There are, however, many
species of uncertain microscopical character and, though I have refer-red some
of them to particular genera on shreds of evidence, there is a tiresome remain
der, and much error. The second key is, therefore, artificial and, in the old
style, supplementary. It might have been extended to species, but it would
have been cumbrous and clogged with those very uncertainties which it was
intended to avoid: many of the dubious species have been misdescribed.
From the illustrations and a knowledge of common species one can soon
comprehend the genera and refer to them directly.
New Species. If a species cannot be identified, one must look through the
residual species incertae sedis in the genera Clavaria, Pterula, and Lachno
cladium and, for small species, in the genus Pistillaria. One of these doubtful
species may then be recognized from locality, habitat, or some other detail.
Failing this the species should be described as new.
TAXONOMIC AND MORPHOLOGICAL NOTES
ON GENERA
A PHELARIA gen. nov.
THIS genus includes the most Stereum-like of Clavarioid fungi. The narrow,
uninflated, and slightly thick-walled hyphae give the tough fibrous texture,
and the resemblance to Stereum is enhanced by the unilateral tendency,of the
hymenium. Nevertheless the branching is Clavarioid and ends with filiform,
often almost Pteruloid, attenuate tips.
The four species which I regard as typical of the genus, namely, A. den
droides, A. brunneola, A. incarnata, and A. spiculosa, have no clamps. It
appears also that A. fiabellata has no clamps, but it is difficult to make sure
with the dried material. On the other hand, A. pusio and A. tuberosa have
clamps at all the septa. A. tuberosa, as mentioned under its description, is
highly problematic and is referred by Donk to Tremellodendron (in litt.);
A. pusio may well be a related species: possil:>ly, Clavulinopsis pogonati
belongs also with them.
A. fiabellata raises the vexed question of the generic value of gelatinous
hyphae in contrast to d ry-walled hyphae. I have discussed this point on
page 19.
The affinity of the genus is obscure. Possibly it connects the monomitic
pre-Pteruloid state with some Stereoid-group. An ancestral state with
clamped inflating hyphae and white, smooth, aguttate spores would represent
the branched ancestor of the Clavariadelphus-Pistillaria-Typhula-series:
without clamps it would be near the ancestor of Clavaria; and the derivative,
epiphytic, unbranched state, adapted to inferior and ephemeral habitats, is
to be seen, perhaps, in Ceratellopsis. I have not detected any intermediate
'Pistillarid-form' in Malaya, unless the phycophilous Clavaria fossicola is such,
but there are certainly many discoveries yet to be made in tropical and sub
tropical mycology.
ARAEOCORYNE gen. nov.

This genus seems near to Clavariadelphus and Pistillaria. In the small size
and the slender stem it resembles Pistillaria, but differs in the thickening
hymenium and the absence of agglutination from the superficial hyphae of
the stem. The sterile apex, which is the expanded top of the primordial shaft,
resembles that of Clavariadelphus truncatus and C. unicolor, but one can
hardly believe that these large and almost Cantharelloid fungi are congeneric
with Araeocoryne. Possibly it connects with some other pseudo-Cantharel-
ARAEOCORYNE 33
loid series. The thickening hymenium and �bsence of marginal growth show
that it is not related to the degenerate, gill-less Omphalias.
Superficially Clavicorona (Clavaria) Mairei is similar, but its glol:!ocystidia
and 1-guttate spores are fundamentally different. Likewi�e Caripia differs in
its firm flesh, composed of thick-walled and scarcely inflated hyphae, its
cystidia, and its angiocarpic development. The similarity in form of the
fruit-body in all three fungi is convergent through reduction.
0000
Tl!lrr-Fm. 15. Aphelaria spiculosa: longitudinal hyphae, young hymenial hyphae, hymenium
and spores, X 1,000.
I have not seen young primordia of Araeocoryne, but there can be no

doubt that they are essentially Clavarioid. Instead of the apex becoming
fertile, however, it remains steriJe and the longitudinal hyphae composing it
inflate and become pulled apart by the expanding flesh and hymenium of the
lower part of the head, in the same way as the pulling out of the legs of a
folding chair will flatten its top. The papilla in the centre of the sterile apex
is the vestige of the tip of the primordial shaft, and the ce!Js of the hyphae
composing it are progressively more inflated the greater their distance from
the apex of the papilla. There is no rudimentary hymenium and there are no
cystidia on the sterile apex. The cells of the flesh are mostly 15-40X 3-6 µ,
in the flesh of the head before it begins to enlarge, but they are probably
smaller still on delimitation at the apex of the primordial shaft.
The thickening of the hymenium causes the slight longitudinal folds which
may develop towards the apex of the fruit-body.
5119 D
34 TAXONOMIC AND MORPHOLOGICAL NOTES
CARIPIA O.K.
This name was proposed in place of Hypolyssus B. and Perona Fr. which
are later homonyms of two genera of Persoon's. Hypolyssus Pers. (Myc. Eur.
2, 1825, 6) was based on two English agarics illustrated by Sowerby and,
. i!l \Jll\lljl l\l\l1 \l l1\ litf1 f1\ \1 11\ll'JI l�i

•11 !11111 1)(/I\1 111!1 1 1 1 1/1i1i1,,
, i\ Jililij1)/li !1l/j1/J 1rII 11 1
ij� 1 \ i\1l1i1 1 t\1 :1 i1 1 1 :i1 :it11 /1 1tl

1
1
1
- . �"".....-�....:,�':,-�-"t1�:::::;�----==---=---::=.-�-a---
TEXT-FIG. 16. Aphelaria spiculosa: diagram of the
structure of the stem to show the cristation by rudi
mentary primordia giving the strigose appearance.
according to Berkeley and Fries, they are agarics parasitized by Hypocrea. In

1842 Berkeley amended Hypolyssus to .cover the little-understood H. Mon
tagnei, but this was tantamount to using Hypolyssus Pers. in an illegitimate
sense. Fries, therefore, proµosed Perona as a new name, but it, too, is ante
dated. Martin has suggested that Hypolyssus sensu B. should be conserved
against Perona Fr. (Mycol. 30, 1938, 440), but Donk sees no reason for con
serving the obvious misnomer (Bull. Bot. Gard. Buit. 3, 1941, 17). It follows
CARIPIA 35
that one must use Caripia: actually two other names, Podostrumbium and
Heringia, had been proposed, but neither was validly published acrnrding to
the Rules of Nomenclature. As there is only one species which, until very
recently, has scarcely appeared in mycological literature, there can be little
inconvenience in the rigid application of the Rules.
Note. Caripia has been included in this monograph because, at the time of
writing, I could find no evidence, published or in herbarium-material, that it was
Cantharelloid as usually regarded. Examination of living material in all stages of
development in 1948, however, has shown me that the young fruit-bodies have very
slight marginal growth to form a very rudimentary pileus. Caripia is, therefore,
Cantharelloid though the mature fruit-bodies appear Clavarioid, as if reduced mem
bers of the Lentaria-group. The new findings will be reported in a monograph on
Cantharelloid fungi.
CERATELLOPSIS Konr. et Maubl.

This name was proposed, in 1937, to substitute Ceratella Pat. (1883),
which is a later homonym of Ceratella Hook. f. (1845). Ceratellopsis must,
therefore, be defined and typified in the original basis of Ceratella Pat.
Unfortunately the generic definition covers at least two kinds of minute
Clavarioid fungus, and Patouillard did not indicate a type-species. He in
tended the genus for small, simple, or subsimple, filiform fruit-bodies with
sterile tips and white spores: as examples he gave, in 1887 (Hym. Eur.),
'C. Queletii, C. acuminata, C. aculina etc.': in 1900 (Ess. Tax.), he named
also C. aculeata, C. Helenae, C. macrospora, and C. uncialis, though the first
two must have been intended in the original term 'etc.', as they had already
been described in his Tabulae Analyticae.
Now C. aculina ( = Pistillaria aculina Quel.) is a Pterula with dimitic
hyphae and it must be identified with Typhula gracilis B. et Desm. as Pterula
gracilis (p. 514). C. aculeata, in contrast, is monomitic with very narrow,
uninftated hyphae. It cannot be classified generically with C. acuJina for it
probably represents the depauperate 'epiphytic' state of Aphelaria, with simi
lar hyphae, or, perhaps, Lentaria with inflating hyphae. It is, nevertheless,
one of a group of species, which probably includes C. acuminata, and which
needs a generic name, at least until it can be attached to its less reduced
generic state. It is in this sense, based on C. aculeata and probably C. acumi
nata, that I have used Ceratellopsis. Of the other species referred by Patouil
lard to the genus, C. macrospora is almost certainly Pterul;:, (under which I
have described it), C. uncialis is a Pistillaria, and both C. Queletii and C.
Helenae appear Pteruloid, though their structure is not known in sufficient
detail to decide. C. Queletii, however, is the species chosen by Konrad and
Maublanc for the lecto-type of Ceratellopsis, apparently because it was the
first to be listed by Patouillard. If it has the same structure as C. aculeata
there will be no difficulty, but if it is a Pterula, then Ceratellopsis will become
merely a synonym of Pterula and a new generic name will have to be made for
C. aculeata. Thus I consider it preferable to change the lecto-type of Cera
tellopsis so that the name can be used with certainty. Proposals for lecto
types in the genera of these very small reduced fungi can hardly be accepted
without microscopic analysis. The sterile apex, supposed to define the genus,
is present in every Clavarioid fungus and is noticeable in this particular case
because the small size of the fruit-body necessitates greater magnification
even for superficial macroscopic study. Beside making a lecto-type, one must
prove that a genus is a morphological entity.
CLAVARIA Fr. s. str.

DEFINITION
This genus is the original receptacle for all Clavarioid basidiomycetes. It
has thus reached unwieldy dimensions and covers many microscopic differ
ences of generic, if not family, rank, e.g. Clavaria botrytis, C. vermicularis,
C.fusiformis, C.pyxidata, C.angulispora, C.cristata, C. byssiseda, C.pistillaris.
Its restriction is simplified by Bonorden's removal (1851) of the Friesian
subgenera Ramaria and Holocoryne, leaving only Syncoryne. This subgenus
includes the three common north temperate specirs C.fusiformis, C. fumosa,
and C.vermicularis, of which either C.fusiformisor C.vermicularis presents the
obvious lecto-type. Donk has chosen C. vermicularis (1933) and has been
followed by Konrad and Maublanc (1937). The decision is important because
C. vermicularis (with C. fumosa) has a microscopic structure which I consider
generically different from that of C. fusiformis. I have, therefore, accepted
C. vermicularis as the lecto-type and, so far as possible, limited the genus to
its affinity. There remain still many species of uncertain position, mainly
extra-European, which I have been obliged to retain in Clavaria s. lato
because their microscopic structure is unknown. Thus, instead of the grand
total of more than 500 species placed in Clavaria, I have reduced the number
to 24 certain species, 16 possible species, and 22 incertae sedis. My definition
will appeir too narrow until it is realized that the natural classification of the
higher fungi must be as microscopic as that of the lower fungi and that of all
classes of algae. A typical Clavaria s. str. has a structure immediately recog
nizable under the microscope: as proof, I was able to detect from the minute
dried specimens of C. scabra, which were 90 years old, that it was a true
Clavaria, and so this very obscure, neglected species should now be recog
nizable: the definition also removes the violet or amethyst C. Zollingeri, as a
true Clavaria, frnm the neighbourhood of Clavulina amethystina with which
it has been often confused, and it is as useful in distinguishing the pink species
of Clavaria s. str., as C. incarnata, from those of Clavulinopsis, as C. miniata.
Concerning Holocoryne Bon., which is merely the Friesian subgenus
raised to generic rank without explanation or indication of a type, it is neces
sary to follow Donk on grounds of priority. Of the twelve original species of
Holocoryne, Donk (1933) has taken C. acuta and its synonym C. falcata as
typifying Holocoryne and has, therefore, reduced the genus as a synonym of
CLAVARIA 37
Clavaria (C. acuta has essentially the same hyphal construction as C. vermicu
laris). The other ten species must therefore be distributed as follows:
Clavariadelphus Donk: C.pistillaris, C. ligula, C. contorta, C. macrorhiza,
C. fistulosa, C. ardenia, C.juncea.
Clavulina Schrot.: C. canaliculata.
Lentaria Corner: C. mucida.
Pistillaria Fr.: C. uncialis.
One might have wished that C. pistillaris could have ,become the type of
Holocoryne, thus preserving a generic name, but this is no longer possible as
it has been excluded by Donk. Based on C. acuta, however, it becomes a
distinct subgenus of Clavaria s. str. (p. 39).
Concerning Stichoclavaria it is difficult to decide. The genus was based on
Juel's interpretation of C.falcata Fr., as a stichobasidial fungus with 6-7-8-
spored basidia and long-ellipsoid spores. This is certainly not C. falcata Fr.
(= C. acuta), and it remains to be rediscovered and identified or described
as new (cf. Clavulinopsis coronilla). Stichoclavaria, based on C. falcata by
misnomer, must therefore be illegitimate or a direct synonym of Clavaria.
Such a puzzle was introduced by way of elementary science (Lindau, Krypt.
FI. f. Anf. 1, 1928, 3rd ed.).
STRUCTURE AND DEVELOPMENT OF CLA VARIA ZOLLINGERI

This is the one fully branched species of the genus and is, therefore, the
most representative.
Flesh. The hyphae are mainly longitudinal, inflated, and short-celled, with
very copious secondary septation (see p. 14), but among these occur narrow
uninflated hyphae, 2·5-5 µ, wide, which are less septate, and thus longer-celled.
Next the subhymenium, the tissue is compact: the cells are short and often
subglobose, or pyriform, 20-50 X 5-20 µ,. Internally the tissue is loose, being
pulled out by the expansion of the hymenium.
Hymenium and Subhymenium. The hymenium thickens by th� gradual
sympodial development of basidia and eventually reaches treble its original
thickness as a single layer of basidia. During this outgrowth the first sub
hymenial cells inflate and round off, but the intercalary expansion of the
hymenium prevents them from forming a pseudoparenchyma. The sub
terminal cells, immediately below the basidia, are 8-12 X 2· 5-3·5 µ,. The tissue
of both layers is rather loose and easily displayed under the microscope by
light tapping on the coverslip.
Stem. The structure is that of the flesh in the fertile branches, but at the
surface there is a narrow layer of uninflated hyphae 1· 5-3·5 µ, wide, from
which project similar uninflated hyphal ends up to 100 µ, long. The ends may
be short and scattered or long and numerous, but they do not form a regular
pile, at most a thin tomentum which can be seen only after drying the fruit
body.
j
Growing-points. A t the tip of the primordial shaft or at the tips of the
branches the tissue is rather loose and composed of uninflated longitudinal
hyphae 1·5-3 µ, wide. The subterminal cells are 15-30 µ, long on delimitation
and most begin to inflate directly. The larger ones, on reaching nearly their
full size, become secondarily septate by forming one new septum at a time,
less often 2-3 almost simultaneously. The hymenium develops 200-500 µ,
behind the .apex in the usual way by the out-turning of the superficial hyphal
ends and by sympodial branching. Some of the first hymenial cells become
basidia, but most cut off 2-3 subterminal cells before maturing: thus, the sub
hymenium has a considerable thickness, 20-30 µ,, from its beginning.
VARIATION AMONG THE SPECIES
Habit. Some species, as C.fumosa and C. purpurea, are typically caespitose,
others are variable and have gregarious and solitary states, as C. vermicularis:
a few are never caespitose, as C. helicoides and C. fossicola. This character is,
however, rarely diagnostic, for the caespitose species may occur in im
poverished habitats not allowing their full and copious growth. The scattered
growth of the non-caespitose species, even under most favourable conditions,
seems, however, diagnostic.
Form. Most species are simple or have an occasional branch, which may
be merely an early connation carried up by intercalary growth. A few, as C.
fossicola, have some fruit-bodies sparingly branched. C. Zollingeri is typically
much branched, but it has sparingly branched forms and even almost simple
caespitose tufts.
Size. The large size of some fruit-bodies, up to 15 cm. high, as in C.
fumosa and C. purpurea, is caused by secondary septation and inflation of
the cells rather than by prolonged apical growth, which is generally restricted.
The small size of C. helicoides, C. fossicola, and C. filiola is caused by the
small amount of secondary septation and the slight inflation. Thus, analysis
of the size of the fruit-body means its resolution into (i) the amount of apical
growth, (ii) the amount of secondary septation, and (iii) the amount of infla
tion. Th� variation of C. vermicularis and C. Zollingeri in this respect is prob
ably very complicated.
Hymenium. In C. Zollingeri, C. helicoides, C. incarnata, C. fumosa, C.
Gibbsiae, and C. fossicola the hymenium thickens and in C. fossicola the
thickening is most pronounced, reaching 350 µ,, if not more. In C. vermicu
laris, C. filiola, C. acuta, C. argillacea, and perhaps C. tenuipes, I have found
that the hymenium does not thicken. The state of the other species is not
known. The subhymenal cetls inflate only in the species with thickening
hymenium.
Spore-contents. Typically the spores have either cloudy-vacuolate contents,
opalescent-vitreous contents, or, as in the subgen. Holocoryne, minutely
granular-guttulate contents, but there is a large gutta in the spores of C. pur
purea and C. vermicularis var. singaporensis. The differences seem to be only
matters of degree, that is, of the state of dispersal of the oil in the cytoplasm.
If the oil is in ultra-microscopic droplets, the cytoplasm appears cloudy:
CLAVARIA 39
if in fine microscopic droplets 0·2-1 /L wide, it appears granular or multi
guttulate: if these droplets coalesce, it appears 1 -guttate. The cf:l:!gree of
dispersion may depend on temperature, so that the same species may vary in
this respect in different parts of the world.
Secondary Septation. The cells of C. vermicularis, C. acuta, C. fumosa, C.
incarnata, C. argillacea, and C. Zollingeri develop many secondary septa.
The cells of C. filiola have 3 secondary septa, those of C. helicoides 1-2, and
those of C. fossicola and C. scabra 0-1. The secondary septation therefore
seems degenerate in these four and the failure of the mechanism probably
explains their small size. On the other hand, neither the large C. Gibbsiae
nor the small C. tenuipes have secondary septa and their variation in size seems
to depend on the abundance of primary septa.
Texture. The characteristically brittle flesh of Clavaria is caused by the
inflation and secondary, or close, septation of the thin-walled hyphae which
are not constricted by clamps. That of Clavulinopsis and Clavulina is slightly
tough because the cells usually taper to the clamps or are prevented by them
from swelling at the septa, apart from the presence of narrow, uninflated
hyphae or of slightly thickened walls, and thus their fruit-bodies do not snap
easily across the broad hyphal tubes. C. fossicola with rather narrow hyphae
and tough fruit-bodies is exceptional.
Nuclei. Concerning the cytological structure of C. vermicularis, I quote
these lines from Coker (p. 47): 'Rosenvinge finds that in C. vermicularis the
cells of the flesh may contain one to four nuclei, and that the mature spores
have regularly two nuclei (Ann. Sci. Nat. Bot. Ser. 7, 3, 1886, 75, pl. 1, figs.
1-3). He finds that the spore of C. fragilis has but one nucleus (no figures)
and thinks that in these cases as in others the number of nuclei in the spore is
of systematic importance. One cannot be sure what plants he determines as
C. vermicularis and C. fragilis. Juel (Nova Acta Reg. Soc. Sci. Upsal. scr. 4,
4, 1916, 1), on the contrary, finds the spore of what he calls C. fragilis to have
two nuclei (pl. 3, figs. 87-90), resulting from a division in the maturing spore.
He further finds the basidia 4-spored, the basidia and hymenium much as in
C. muscoides and C. subtilis. The latter author has sent us (as C. fragilis) a
collection from Upsala which is like ours: spores smooth, oval, 3-3 ·7 X 3·8-
5"4 µ,.'
CLASSIFICATION OF THE SPECIES
I have been able to retain and emend the two Friesian subgenera Syncoryne
and Holocoryne because of a remarkable microscopic duference between
their lecto-types, C. vermicularis and C. acuta respectively. The emended
diagnoses are given on page 217. The important feature is the presence of a
wide, loop-like clamp at the base of the basidium in C. acuta and in other
species which I place with it in Holocoryne: C. vermicularis and its allies in
Syncoryne have clamps neither on the basidia nor on the hyphae of the fruit
body. The Holocoryne-clamp (Text-Figs. II, 95) is difficult or impossible to
make out in dried material, but it is clearly indicated in living material because
the densely guttulate or oleaginous base of the basidium contrasts with the
watery-J;iyaline subhymenial hyphae and shows up the more or less widely
furcate bi.:se. If fragments of the living hymenium of a species of Holocoryne
are carefully teased apart and splayed asunder by tapping on the coverslip,
then, under an immersion lens, one arm of the forked base of the basidium
can often be traced, as a hyaline hypha, to the subterminal cell of the basidial
hypha. This is the backward branch of the clamp, and the interval between
its point of origin and its connexion with the subterminal cell may be 5-15 µ,.
The subterminal cell, as usual, branches one to three (to four) times and each
branch ends in a basidium with similar loose clamp. The branch may arise
from the body of the subterminal cell or, and apparently as often, from any
part of the clamp proximal to its septum, and the basidium may then seem
merely to be joined by an H-connexion to a subhymenial cell. All the loose
clamps which I managed to trace in C. Gibbsiae and C. tenuipes connected
the basidium with its own subterminal cell but in several instances the base of
the basidium was so widely furcate that I could not see how the clamp was,
or could have been, connected to its own subterminal cell and it seemed
possible that the clamp was connected to another hypha. It is not improbable
that in the confusion of hyphae the long backward branch of the clamp may
frequently unite with any subterminal cell in its brief haploid phase, not
necessarily with its own subterminal cell which then becomes recipient for
another clamp: the subhymenium may, thus, be a network of hyphae joined
by the basidial clamps.
I first discovered these clamps in C. Gibbsiae and in alcohol-formalin
material of C. tenuipes, in 1942, since when I have been able to confirm the
presence of identical basidia in living material of C. acuta, C. argillacea, and
C. incarnata, thus making a remarkable assemblage of nearly allied species.
All of them have finely multiguttulate spores, more widely ellipsoid than usual
in Syncoryne, and so, by analogy, it seems that C. alliacea and C. Daulnoyae,
if not C. fuscata, C. rosea, and C. straminea, belong with them. No branched
Holocory-9e seems to exist.
On page 217 I have arranged the species in small and, apparently, natural
groups, though most are still too little known for any but a tentative scheme.
The cystidia are but slightly developed and, probably, of no great importance:
thus C. purpurea, C. atrobadia, C. atroumbrina, and C. fumosa seem to be
variations on a brown-purple pigmentation and C. vermicularis to be the
albino state. It is noteworthy that the colour of C. argillacea, C. fumosa, and
C. incarnata lies in the longitudinal or medullary hyphae rather than in the
hymenium as is generally the case in Ramaria, Clavulinopsis, and Clavulina.
Whether the other coloured species, as C. purpurea and C. Zollingeri, also
have colourless hymenium and pigmented medulla remains to be seen. In
C. straminea it appears that a medullary yellow becomes masked by a trau
matic browning, which may proceed hysterochroically with age: in other
respects it is close to C. acuta (Holocoryne). In C. helicoides, the pink colour
occurs in the basidia as well as in the medullary hyphae and is ultimately
CLAVARIA 41
injected into the spores: it is the only Clavarioid fungus that I have found to
be so thoroughly pigmented.
C. helicoides and C. fossicola are phycophilous and agree in their reduced
secondary septation and somewhat tough texture, but whether they are allied
or spuriously similar is far from clear. C. scabra may be a phycophilous ally
of C.fossicola in the New World.
C. Zollingeri, C. vermicularis, C. argillacea and, possibly, C. purpurea are
very variable species needing careful description and record of individual
collections before they can be fully understood. There appears to be a con
tinuous series from the fully branched to the simple state in C. Zollingeri,
and in C. vermicularis from the massive caespitose state to the solitary
Pistillaria-form with slender pellucid stem. The other two have almost as
great a range as C. vermicularis.
EVOLUTION
Clavaria must be allied with the other white-spored monomitic genera with
inflating hyphae, namely, Clavulinopsis, Ramariopsis, Clavulina, and, perhaps,
Clavariadelphus, but I have met no species connecting with them. If it
exists, it will be found among the branched species of Clavulinopsis, but I
doubt the likelihood for Cfavaria is a natural genus of much-reduced species,
the ancestral state of which has probably disappeared.
On the principle of the degeneration of the fruit-body, C. Zollingeri with
its branched, pigmented fruit-body and thickening hymenium is the nearest
approach to the ancestral form: indeed, it shows every gradation from the
highly branched to the simple state. Such an ancestral form differs only in
its secondary septation, it� lack of clamps, and its small spores from the 4-
spored ancestral state which must be postulated for Clavulina, and it is note
worthy that secondary septation, lack of clamps, and similar violet, purple,
brown, and grey pigmentation occur in Clavulina. This agreement suggests
a common ancestor for Clavaria and Clavulina with normally clamped hyphae
such as the branched species of Clavulinopsis, e.g. C. cineroides. Tpe secon
dary septation of the old basidia, characteristic of Clavulina, occurs also in
Clavaria helicoides.
The white species of Clavaria with the smallest, unpigmented fruit-bodies
and the smallest spores represent the most degenerate state of this hyphal
construction. They resemble Pistillaria but are humicolous, not epiphytic,
though even this condition may be reached in C. tenuipes.
Problematic is the Holocoryne-group of simple fruit-bodies. The clamp
on the basidium would appear to be a relic from the normally clamped state
of Clavulinopsis, and I have seen normal clamps very occasionally on some
hyphae near the base of the stem of C. tenuipes. The curious form of the
Holocoryne-clamp would, therefore, seem derivative rather than primitive,
but it needs cytological and cultural investigation before it can be understood.
Indeed, the question of clamps in all Basidiomycetes needs much more
detailed investigation and elucidation (see p. 19).
The discovery of other branched species of Clavaria s. str. will be awaited
with grt>at interest.
CLAVARIACHAETE gen. nov.

This is the Hymenochaete-representative among Clavarioid fungi. Only
two species are known, as a single collection of each, from tropical America,
and they may be growth-forms of one species. It is a most interesting and,
phylogenetically, important genus.
Structure of the Fruit-body of C. rubiginosum. A medulla of longitudinal
hyphae traverses the whole fruit-body. The stem and lowest parts of the
main branches have a thick, subtomentose, sterile cortex: the remainder of
the branches has a thick subhymenium pierced by the setae and faced on all
sides with the hymenium. The medulla consists of a wide, rather compact
core of hyphae with pale-yellowish, scarcely thickened walls and a peripheral
tube of closely entwined hyphae with strongly thickened (1-2 µ,), dark-brown
walls. In the stem and lower branches the tissue of the peripheral tube is
agglutinated into a carbonaceous cylinder, forming a black ring in transverse
section, but in the distal half of the branches it is not agglutinated. The core
of the medulla is c. 300-400 µ, wide in the stem, with the carbonaceous layer
c. 200 µ, thick, and it tapers into the branchlets with a gradual thinning of both
parts: ultimately the peripheral tube may be only 30 µ, thick. The cortex
consists of thick-walled, dark-brown hyphae as those of the peripheral tube,
but they are widely divergent from the axis, rather loosely interwoven, and
sparingly branched or not at all: they have simple ends and are without setae
except at the transition to the hymenium. The hyphae of the subhymenium
resemble those of the cortex but are frequently branched at a wide angle, often
pseudo-dichotomous, have rather short cells, and become gradually thin-walled
towards the hymenium. The hymenium, c. 25 µ, thick, appears to consist of a
single compact layer of colourless basidia. The setae arise from all depths in
the subhymenium as ventricose-lanceolate cells which become progressively
thick-walled acropetally on cessation, or decline, of apical growth.
The medulla is formed by the apical growth of the fascicle of hyphae at
the growing-tip of the branch and both cortex and hymenium are formed by
the outcurving of the superficial lateral hyphae, many of which enlarge
directly into the setae.
CLAVARIADELPHUS Donk
This genus was proposed in 1933 for Clavaria pistillaris, C. ligula, and C.
truncata, on the grounds that the long basidia, spongy flesh, simple fruit-body,
and large, smooth, white, ellipsoid spore were distinctive. It certainly repre
sents the nucleus of a group, but I am unable to exclude C. fistulosa (retained
in Clavaria as an appendix, by Donk), C. juncea (placed in Typhula by most
authors), and the North American Craterellus unicolor which appears as the
CLAVARIADELPHUS 43
transition to Cantherellus. The four species C. pistillaris, C. ligula, C. fistulosa
(incl. C. contorta), and C. juncea were placed by Fries in the subgenus Holo
coryne of Clavaria (1874) and my own researches have convinced nie that no
hard-and-fast line can be drawn between them, particularly in view of the
recent Clavaria maricola described from North America.
In Table I I have compared the main features of nine species which I refer
to Clavariadelphus. In every case there is a transition from one extreme to the
other. There appear to be three subgenera which I have called Eu-Clavaria
delphus, Cantharellopsis, and Typhulopsis (p. 271 ), and they indicate the
unique range of this Clavarioid-group from the Agaric form to the Typhuloid:
none can be clearly demarcated from the others. Thus C. pistillaris is very
nearly related with C. truncatus, and C. unicolor, with rudimentary pileus, has
been mistaken for C. truncatus and may have to be transferred to Cantharellus:
TABLE I. SPECIFIC DIFFERENCES IN CLAVARIADELPHUS
Macroscopic Differences
Species Size (cm.) Apex Shape Habitat
C. unicolor 3-5X1·5-5 sterile, often per- subpileate humus, mixed
forate woods
C. truncatus 6-15 X2-9 sterile clavate- coniferous
truncate needles
C. pistillaris 7-30X2-6 acute, then obtuse, clavate humus, frondose
fertile
- -
woods
var. americanus often flattened ,, ,,
C. ligula 3-1oxo·5-1·5 obtuse, fertile ligulate coniferous
needles
C. sachalinensis " " " " " ,, "
C. fistulosus 10-30X0·2-1·3 acute, then blunt, filiform, sub- on sticks and
fertile clavate humus
var. contortus 0·5-3X0·2-0·6 blunt, fertile cylindric- on sticks
clavate
C.junceus 3-15X0·05-0·2 acute, then blunt, filiform on sticks, leaves,
fertile &c.
C. mirus 8-15 xo·6-1·1 obtuse or truncate cylindric on the ground
C. maricolus 7-10X0·2-0·6 blunt, fertile subcylindric, on coniferous
subligulate humus
Microscopic Differences
Species Spores(µ} Basidia (µ) Hyphae
C. unicolor 8-12X4-6 ? ?
C. truncatus 9-14X5-8 6ox12 ?
C. pistillaris II-16X6-10 90-12ox11-13 3-16 /L wide, cells -600 /L long
var. americanus 7-12·5X3'7--'7'5 ? ?
C. ligula 8-15X3-6 40-65X6-9 3-16 /L wide
C. sachalinensis 17-22x 5-5·5 ? ?
C. fistulosus 10-18·5X4·5-9 40-65x8-u·5 1·5-15 /L wide, some laticiferous
var. contortus 14-23X7--9 45-8ox6-10 2-20 /L wide, some laticiferous
C.junceus 6-12X3·5-5 30-55X6-10 3-25 /L wide, some laticiferous
C. mirus 8-10X4-5 30X9 ?
C. maricolus 10-14X4-5 108-12x4-5 ?
C.- mirus is still problematical, possibly with a sterile apex, which may occur
also in �he extreme forms of C. fistulosus with truncate top ( = C. ardenia):
C. maricdus, in size, at least, connects Eu-Clavariadelphus with Typhulopsis,
and even C.fistulosus ranges from forms almost as slender as C. junceus to
others nearly as massive as C. ligula or C. pistillaris var. americanus.
The gem:1s must be defined by exclusion and affinity rather than by peculiari
ties. The spongy flesh, so generally emphasized for C. pistillaris, is caused
mainly by the intercalary growth of the hymenium which pulls apart the much
interwoven hyphae of flesh without forming a clear hollow. Hence, the flesh
of C. ligula, with much less intercalary growth, but the same construction,
is firm, whereas in C. junceus and C.fistulosus, with little intercalary thicken
ing and strictly longitudinal hyphae (consequent on their relatively small
primordia), the flesh is rigid and becomes hollow. For comparison, Clavulin
opsis amoena or C. spiralis may have spongy flesh in old fruit-bodies with
much-thickened and rugulose hymenia, whereas C.fusiformis has not. Such
slight, mainly mechanical, differences have not generic rank.
The long basidia, emphasized by Donk, seem to occur in C. maricolus
which, in other respects, is very near to C. fistulosus with fairly short basidia:
indeed, C. ligula has basidia only about a quarter as long again as those of
C. junceus which approximate to the short basidia of Typhula. The great
length of the basidia of C. pistillaris is reflected in the length of the hyphal
cells, which may reach 600 p, long, and it seems to be a hyphal character
rather than a peculiar reproductive one. Clavulinopsis umbrinella has almost
as long basidia compared with others of its genus.
As regards the size, or massiveness, of the fruit-body, there is every grada
tion from C.pistillaris to C.juncea and, when estimated in numbers of hyphae
composing the fruit-body, there is found to be a greater difference between
C.fistulosus and C.junceus than between C.fistulosus and C. pistillaris. Thus : 1
C. pistillaris: No. of hyphae in cross-section, 5-10 million
C. fistulosus: ,, ,, ,, ,, ,, 1 /4 ,,
C.junceus: ,, ,, ,, ,, ,, 4,000
Their ratios are, in order, 30: 1: 1/60. The difference is no greater than
occurs in most large natural genera of basidiomycetes, e.g. Amanita, Pluteus,
Coprinus, Agaricus.
As regards the habitat, there is every transition from the terrestrial C.
pistillaris to the 'epiphytic' habitat of C.fistulosus and C. junceus on sticks or
leaves: indeed, q. fistulosus may grow in humus and C.junceus I have found
in England on bare .soil.
Concerning the presence of laticiferous or gloeocystidia-like hyphae, Donk
emphasizes their absence from Eu-Clavariadelphus and their presence in
C.junceus as indicating affinity with Typhula. They occur also in C.fistulosus,
1
If r is the radius of cross-section of a cylindric fruit-body of average size and w is the
width of a hypha of average size for the fruit-body, the number of hyphae in cross-section is
'1TT2
roughly
w2•
CLAVARIADELPHUS 45
which cannot be regarded as anything but the link between C. pistillaris and
C. junceus. Moreover, scattered hyphae with oleaginous contents occur in
many unrelated basidiomycetes and may be present or not in many natural
genera, as Mycena, Russula, Entoloma, Clavaria, and so on.
Accordingly, I regard Clavariadelphus as a peculiarly indistinctive, central
group of unbranched Clavarioid fungi with white, ellipsoid, smooth, aguttate
spores, with predominantly yellowish or brownish pigmentation, and with
monomitic, clamped hyphae. On the one hand, it connects through the sub
genus Typhulopsis with Typhula and Pistillaria, but it lacks the simplified
hymenium and specialized habitat of these genera, as well as the sclerotium
of the first. One is apt to overlook the importance of this feature, but no large
Clavarioid fungi are known with a sclerotium and it would be almost as sur
prising to find C. fistulosus or C. junceus developing from one as C. pistillaris
or C. unicolor. On the other hand, it seems to connect through the subgenus
Cantharellopsis with Cantharellus, which differs in the fully developed pileus
with marginal growth and sterile upper surface, the sterile stem with hymen
ium restricted to its apex and to the underside of the pileus and, possibly, in
the 'stichic' basidia. Those of Eu-Clavariadelphus are chiastic (and 4-spored),
according to Juel, whereas the basidia of Cantharellus cibarius, C. cinereus,
C. tubaeformis, and C. lutescens are stichic (subcylindric, with the first spindle
of the fusion nucleus longitudinal). However, the importance of this feature
has yet to be explained: it seems to indicate, merely, that stichic basidia are
degenerate and hypha-like and too narrowly clavate to allow a transverse
spindle: I will refer to the point at greater length in the discussion of the
basidium. Nevertheless, the fact remains that in Cantharellus there is a
tendency to produce 5, 6, 7, or 8 spores on the basidium, and when, to this, is
added the transition in development of tht! pileus via Cantharellopsis to
Eu-Clavariadelphus, the question must be raised whether Clavariadelphus
represents a Cantharellus reduced to the primordial shaft or Cantharellus
represents a pileate Clavariadelphus. The absence of true apical branching
in Clavariadelphus, Typhula, and Pistillaria (even in the rhizomorphs of
C. junceus) suggests that they are fundamentally monopodial and that their
polypodial, or branched, state is to be found among branched Cantharelloid
fungi. Alternatively, a branched Clavariadelphus would resemble Lentaria
with thin-walled hyphae. I conclude that the position, and circumscription,
of Clavariadelphus must await a monograph of Cantharelloid fungi, as well
as the possible discovery of new species of the genus from the mountains of
Asia. The pigmentation of the genus is probably different from the yellow
pigments of Clavulinopsis, which has no tendency to pileate or Cantharelloid
forms: the brown pigment of Typhulopsis is in the hyphal walls, as in the
brown species of Typhula.
DEVELOPMENT AND STRUCTURE OF C. PISTILLARIS

Development. C. pistillaris is locally common in the beech-woods of
Buckinghamshire, in England, and when the first precocious fruit-bodies
A B
'I'ExT-FIG. 17. Clavariadelphus pistillaris: diagrams of five stages in the development of the
fruit-body before the appearance of the hymenium, showing the inception of the primordium
inside the mass of mycelium; A, a young fruit-body beginning to develop a hymenium, x 1:
B, four primordia before the appearance of the hymenium, x s'.
CLAVARIADELPHUS 47
appear above the humus in autumn many young developing stages can be
found by gently brushing aside the dead leaves. Very often the frui<:-bodies
grow from the firm earthy walls of mice-burrows tunnelled just below the
layer of humus, less frequently from the earth in any part below .the humus,
and occasionally from bare ground. It would seem that the mycelium forms
a mycorrhiza, unless it permeates the soil, for the fruit-bodies do not spring
from the humus itself. The primordia are white, fluffy, mycelial patches,
1-3 mm. wide, with a little denser core which enlarges and becomes dome
shaped, then conical, and soon emerges from the mycelial patch as the first
rudiment of the primordial shaft. By continued apical growth and vacuola
tion of the older cells, proceeding acropetally from the base, the primordium
develops into a white cylinder, or slender finger, with a rather blunt point.
After reaching a height of 7-10 cm., occasionally as much as 20 cm., during
which time it has been thickening by inflation as well as lengthening, apical
growth ceases and the fruit-body becomes obtusely clavate as the hymenium
develops over the apex and its intercalary growth widens the cylindric stalk.
Apical growth generally carries the tip of the fruit-body above the humus.
Its tip may then be eaten by slugs and the top of the mature fruit-body
becomes deformed, even sterile, and simulates the truncate apex of C. trun
catus (indeed, one must inquire to what extent this may be the reason for
sterile truncate apices in general, as in C. truncatus and also in C. fistulosus,
where they seem to be exceptional). The fruit-body gradually colours from
below upward, though the buried or unilluminated part of the stalk remains
white. Until the intercalary growth of the hymenium dilates the inner tissue
the flesh is firm and solid and, certainly until the primordium is 4-6 cm. high,
devoid of air-spaces. Then intercalary growth and secondary thickening of
the hymenium pulls apart the hyphae of the flesh, causing the development
of air-spaces and, thus, the spongy-floccose mature texture: the hymenium
also becomes wrinkled as its expansion is restrained by the closely interwoven
hyphae of the subhymenium. The flesh does not become hollow because the
hyphae are much entangled, in a homogenous tissue, though mainl� longitu
dinal, and their cells elongate enormously.
Secondary Mycelium. Hyphae are continually added to the original mycelial
patch by outgrowth from the base of the fruit-body, where the hymenial factor
does not operate: thus a web of guy-ropes holds up the enlarging fruit-body.
Mature fruit-bodies look top-heavy, yet fallen ones are seldom, if ever, seen:
they will derive some support also from the humus through which they rise.
Endogenous Origin. The mycelial hyphae are 2-3 µ, wick and sparingly
branched: they have clamps and minutely rough walls. The white mycelial
primordia of the fruit-bodies are formed by a general increase of the mycelial
hyphae by branching on the surface of the soil; but the basidiocarp arises
from the internal hyphae of these patches, as shown in the Text-Fig. 17.
Inside the mycelial patches the hyphae branch and interweave profusely,
forming a compact tissue which at first expands equally on all sides as a
hemispherical core, but soon its growth is mainly upward, away from the
substratum, and a conical apex is formed. The hyphae of the basidiocarp have
smooth.walls and rather dense contents, by which means they are easily
distinguishable from the original mycelial hypha in the tangled tissue of the
primordium; even in sections of mature fruit-bodies these rough mycelial
hyphae may be seen in the central tissue at the extreme base of the stem.
Text-Fig. 17 shows firstly a longitudinal section of the mycelial patch in
which there is the first sign of the basidiocarp as a denser region of branching
hyphae at the base. Secondly, there is the basidiocarp forming as a hemi
spherical core .still enclosed by the mycelium. Thirdly, apical growth. has
begun: the basidiocarp has alm::>st emerged from the mycelial patch, many
original hyphae of which have been pushed outward at the sides or incorpor
ated in the general tissue; and in the basal part, the hyphae have begun to
enlarge, 3-4 µ wide, and their cells to straighten, the remainder of the tissue
of the basidiocarp being composed of 2-3 µ wide, contiguous, and densely
interwoven hyphae. In the fourth stage the basidiocarp has emerged from the
mycelium, which remains on the sides at its base, and an apical growing
point has arisen so that its shape is conical: the hyphae in the lower part have
enlarged to 4-6 µ wide with scattered cells, or unicellular subclavate branches,
as much as 16 µ wide. Lastly there is a section of a young fruit-body about
3 cm. high. It is colourless and its tissue is still firm and solid, without air
spaces, but the hymenium is already incipient. The internal hyphae are
3-16 µ wide, mostly 5-7 µ,, but towards the surface the hyphae are narrower,
3-4 µ wide, and mostly have grown up as laterals on the surface of the hymen
ium. At the extreme base, the hyphae are 3-6 µ wide and, as usual in large
fruit-bodies, they do not enlarge further. Apical growth is in full vigour.
From the base of the primordium secondary mycelial hyphae, also with rough
walls, have grown out and strengthened the attachment to the soil. Above
the base, the excrescent hyphae from the surface are shorter: they do not
reach the mycelium and are interspersed with others in which the terminal
cell has enlarged somewhat into a caulocystidium. These caulocystidia vary
much in. shape from subclavate to subventricose and occasionally they have
1-4 short curved processes at the apex: downwards there are all transitions
to excrescent rough-walled hyphae; upwards there are all transitions to fertile
basidia. The caulocystidia are, in fact, sterile basidia in the narrow transitional
zone from the mycelial base to the fertile hymenium, their apical processes
being abortive sterigmata. New caulocystidia are intercalated as branches
from the subterminal cells or from hyphae situated internally to a depth of
150 µ, or more. uA small number of these terminal cells mature into normal
basidia at a height of some 20 mm. above the base of the primordium, though
the surface-tissue is loose, minutely subtomentose under a lens, and not
compacted into a hymenium. But from this fifth stage onward the young
basidiocarp becomes fertile, though it has not the shape, colour, or consistency
of the mature fructification. In later stages, from 4 cm. long, the distal half
of the primordium becomes light yellow-ochre and smooth from the continu
ous hymenium.
TExT-FIG. 18. Clavariadelphus pistillaris: part of a fruit-body cut across to show the rugose hymeniwn,
X 1 : diagrams of the structure of the apex of a mature fruit-body and of the thickened hymc:niwn (in
t.s.), to show the ridges.
Apical Growth. The conical apex of the primordium is composed not of a
sheaf of longitudinal hyphae, as might have been expected, but of extremely
numerous, very closely interwoven hyphae with many free tips at the surface.
They are 2-3 µ, wide. The base of the growing-point is a circle about 4 mm.
in diameter, in primordia of average size, so that a simple calculation shows
there must be over a million hyphae in the growing-point. There is little
regularity in the direction of the individual hyphae: some grow upward and
then more or less obliquely: others take an irregular course, insinuating
themselves as best they can between the preformed hyphae: others outstrip
the rest and project some 50 µ,: lateral hyphae may grow beyond their parent
hyphae and all are so inextricably interlaced that it is extremely difficult to
trace them for greater distances than 200 µ,. Nevertheless, the growth of the
whole apex is co-ordinated to form a regular fruit-body. The hyphae grow
most vigorously in the central part. The numerous branches, which they
there produce, in reaching to the surface push the outer hyphae still farther
aside and these hyphae, in receding from the centre, grow more slowly and
drop behind. Hence there is a continuous splaying aside of peripheral hyphae
through more rapid branching in the centre, to give the spring-brunnen effect
of Oltmanns. Even in the mature expanded fruit-body this can be seen on
stripping the flesh longitudinally: the strips peel obliquely towards the
hymenium, a piece of which, as the part formed by this cone of hyphae, being
removed with the strip.
Hymenium. The tips of the deflected outer hyphae are gradually rotated
at right angles to their original direction. During this time they mature into
basidia. Thus the first hymenium is constructed by an extremely gradual
transition in the outer zone of the growing apex. The hymenium thickens
throughout the life of the fruit-body, not merely by upgrowth of laterals
from the subterminal cells of the basidia, but also by laterals from the outer
layer of the flesh which worm their way to the exterior. (The thickening is
even over the whole surface and not papillose or localized as in some Thele
phoras.) _ At first the vacuolation and elongation of the hyphae of the fle'sh
accommodates the expansion of the hymenium, and the fruit-body remains
cylindric, as typical of Clavaria. But the intercalary growth of the hymcnium
soon becomes excessive and pulls apart the hyphae of the flesh, the texture of
which thus loosens gradually to the finally spongy appearance with abundant
air-spaces. Expansion of the hymenium is greatest in the upper part of the
fruit-body, which therefore becomes clavate and even somewhat turbinate.
Through symp�dial growth the hymenium continues to thicken and, because
of the tensions exerted in the densely interwoven subhymenium where the
basidial hyphae are attached to the flesh, the hymenium. becomes longitu
dinally rugulose and the tissue of the flesh is loosest in the layer immediately
beneath the subhymenium. (Text-Fig. 18.) When apical growth ceases all
the hyphal tips become basidia and the hymenium extends over the top of the
fruit-body: if this is sterile, as rarely happens, it is merely the persistence of
the juvenile or developmental state.
CLAVARIADELPHUS 51
The main points in the development of the fruit-body are:
1. The primordium is endogenous in a small loose mass of mycelial hyphae.
2. The growing-point is as massive as that of the primordial shaft of a large
Ramaria, but does not divide.
3. The excessive thickening uf the hymenium and the elongation of the
hyphae of the flesh cause the distention of the fruit-body, leading to its
clavate or subturbinate shape and its spongy-floccose texture: eventually
hymenial thickening causes irregularly anastomosing longitudinal
wrinkles.
4. The apex of the fruit-body is sterile until it has reached about half its
full height, when the hymenium develops over the top.
Cytology. According to Juel the spores are uninucleate and the basidia
contains 4 or, not rarely, 8 nuclei, of which the supernumerary ones remain in
the basidia.
CLAVICORONA Doty
This genus contains the pyxidately branched species C. candelabrum, C.
Colensoi, C.javanica, and C. pyxi.data. They are a natural group distinguished
by the peculiar branching, the small white spores, the more or less toughly
gelatinous hymenium, and the gloeocystidia. Such branching does not occur
in any other Clavarioid fungi, nor do gloeocystidia, except in Lachnocladium,
which has a totally different hyphal construction. Nevertheless, gloeocystidia
occur in four simple fungi which have been described as Craterellus cristatus,
Gr. taxophilus, Clavaria Mairei, and Physalacria tuba. Their fruit-bodies are
small (7�30 mm. high) and narrowly obconic with sterile cyathiform tops;
their spores are small and white; two of them, Gr. cristatus and Cl. Mairei,
have incipient or rudimentary branching; and two of them, Gr. cristatus and
P. tuba, are,lignicolous and have the rather tough, dry consistency of C. pyxi
data. They present, in fact, what one would expect for simple species of
Clavicorona, just as the simple species of Clavaria, Clavulina, Clavolinopsis,
and Pterula; and, as evidence that the branched species of Clavicorona can
be reduced, Coker describes simple and subsimple states of C. pyxi.data
without proper cups. None of them, moreover, can be retained in the genus
to which it has been referred. There can be little doubt over the affinity of
Gr. cristatus, Gr. taxophilus, and,P. tuba, but Cl. Mairei has soft, waxy flesh,
unlike that of the other species, so that its inclusion, while convenient, is
tentative: it cannot be referred to Araeocoryne or Caripia, which have no
gloeocystidia and differ also in spores or texture. The eight species of Clavi
corona, thus constituted, are rare or uncommon but, possibly, very widely
distributed: thus C. pyxidata appears to occur throughout the north tem
perate region and in temperate Australasia; C. Colensoi appears to be pan
tropic and to extend into North America and New Zealand; and it is difficult
to find a structural difference between C. Mairei of France and C. taxophila
of North America.
Among the branched species of the genus there are remarkable differences
in text1..:re and hyphae:
C. javanica; monomitic with gelatinous inflating hyphae: without a peripheral
mechanical cylinder Qf longitudinal hyphae: the gloeocystidia and imperfectly formed
gloeocystidial hyphae forming a network throughout the medulla: colourless: without
cystidia.
ToxT-FIG. 19. Clavicorona candelabrum: full-grown and developing fruit-bodies, x 2.

C. pyxidata; monornitic with dry, thin, or slightly thick-walled, inflating hyphae:
with a peripheral cylinder of longitudinal and slightly thick-walled hyphae surrounding
the looser medulla : ? with cytoplasmic pigmentation: with cystidia.
C. candelalwum; dirnitic with thick-walled, limited, tapering, skeletal hyphae in the
peripheral mechanical cylinder of the flesh, the generative hyphae not inflating:
pigment in the walls of the skeletal hyphae: without cystidia.
C. Colensoi: a!. C. candelabrum but with the hyphae of C. pyxidata.
Such a range from gelatinous inflating hyphae to dry, dimitic, uninflated
hyphae suggests that the living species are diversified relics of a large group,
. the members of which were superficially alike but plastic in hyphal construc
tion. When one looks for allied genera it is impossible to find any with cer
tainty. Some species of 'Stereum' with gloeocystidia,1 as Podoscypha, may
' Martin, G. W., Lloydia 7, 1944, p. 74.
CLAVICORONA 53
be allied and, perhaps, Masseeola (= Sparassis), the gloeocystidial hyphae
of which are mentioned by Donk (Rev. Niederl. Homobas. 2, 70). If so,.the
long-standing problem of Masseeola will be solved, together with that of
Clavicorona, as relics of a small series of Homobasidiomycetes distinguished
mainly by the gloeocystidia, the smooth hymenium, the small white spores,
and the clamped hyphae. C. javanica points to an ancestral state with gela
tinous hyphae, which lingers in all species as the more or less gelatinous
hymenium traversed by the thin-walled gloeocystidia.
A B C D
TEx.T-FIG. 20. Clavicorona candelabrum: A-B, stages of the normal division of a
branch-tip into 4 branches: C, with 3 abortive branchlets: D, with 5 branches
(cut off), X 10.
TEXT-FIG. 21. Clavicoronajavanica: polychotomous tips, X 15.
Dimitic State. Skeletal hyphae are typically modified longitudinal hyphae

of unlimited growth, though they may become limited in derived states, e.g.
Fornes. Those of C. candelabrum are of limited growth and they occupy the
position of the longitudinal gloeocystidia and central gloeocystidia-like hypha
in C. javanica and C. pyxidata. In shape and size they agree also with the
gloeocystidia. Therefore I consider them to be not true skeletal hyphae
homologous with those of Pterula and Polyporoid and Stereoid genera but
transformed gloeocystidia which have developed thick brownish walls instead
of oleaginous contents. Thus the surprisingly advanced dimity of one species
can be explained as a not unlikely modification of a generic feature.
Spores. Despite the small size there are considerable differences. C. can
delabrum, C. tuba, and C. pyxidata var. asperospora have very finely punctate
rough spores. C. Mairei has 1-guttate spores, which rather confirm the
suggestion from the waxy texture that its affinity is elsewhere. C. pyxidata,
C. cristata, and C. tuba have distinctly ellipsoid spores: in the other species
they are subglobose.
Cystidia. C. pyxidata is the only species with hymenial cystidia in addition
to gloeocystidia. They may, perhaps, be regarded as incipient gloeocystidia
which abort as the hymenium fails to thicken.
Taste. The contents of the gloeocystidia may cause the acrid taste which
has been noted for C. candelabrum, C. pyxidata, and C. Colensoi (if Clavaria
piperata is rightly referred to it).
Specific Names. In the dried state the species are very similar and one must
await the description of living material before the synonymy can be decided.
Thus Clavaria radiata Lev. is undoubtedly a Clavicorona, but to which
species it should be referred it may be impossible to say until re-discovered
in the type-locality. C. candelabrum may be only a variety of C. Colensoi.
The tropical records of C. pyxidata may be C. Colensoi, C. candelabrum, or
C. javanica.
THE FRUIT-BODY OF C. CANDELABRUM

Structure of the Stem and Branches. (Text-Figs. 19-23.) The tissue is
composed of four concentric layers, or three tubes round the core. There is a
wide, fibrous-spongy, and elastic pith of interwoven hyphae, forming the
greater part of the flesh. Outside it there is a narrow layer, 50-170 µ, thick,
of mainly longitudinal, compact, skeletal hyphae forming a skeletal tube.
Around this is the original subhymenium, 10-20 µ, thick, composed of genera
tive hyphae and gloeocystidia obliquely divergent into the fourth layer, the
hymenium, which thickens from 20 to 300 µ,. Old fruit-bodies have a fifth
layer, the epihymenium, as a thin levigating tissue, 3-8 µ, thick, on the surface
of the hymenium.
Pith. The tissue is aerenchymatous and composed of loosely interwoven
generative and• skeletal hyphae. Because of their limited growth and the
looseness of the tissue, the fine, tapered ends of the skeletals are readily seen.
In the development of the pith the mediate hyphae, producing the skeletals,
seem to grow in any direction, except vertically downward, and hence the
tissue becomes interwoven. In the trunk and older branches there are many
narrow, 3-4 µ, wide, much interwoven and often kinked, skeletals with almost
linear lumen: whether they are later outgrowths of the generative hyphae or
merely modifications of existing hyphae, I do not know.
CLAVICORONA 55
Skeletal Tube. Most of the hyphae are compact, longitudinal skeletals, but
a few generative hyphae occur among them and, more frequently, the longi
tudinal stalks (or proximal parts) of the more deeply pla.ced gloeocystidia. The
layer serves as a mechanical cylinder round the spongy pith, which it braces
into the rigid, fibrous-elastic fruit-body. Externally it is sharply delimited
from the subhymenium but internally it passes gradually into the pith.
�11\/jl'
1/lf 1
1//-�1
lji/1'/i I
�J,
,\ I�\ Jr. /'/
., ,(f/j/0.
�,�\�-"
��- '--- -_.. .r�..L -';ff,jlg,
'i�,�;\}�\Li:0�/�\ljit Ji, /�/1f·\Jfyffl1�
i
1
/
l\'�� \-)' \ f ( 1-j;lfljp

I (/ 11(1/fj
u
�\\ I i
\1ih\ ·{ · i.\Ii I /1'1f/1iii1\
11
�,\d.,·, , !,.U1I
1
TExT-FIG. 22. Clavicorona candelabrum: diagram of a growing branch-tip and of one that has
branched: the skeletal hyphae shown as heavy lines, mag.
Subhymenium. The hyphae are short branches, 1·5-4 µ, wide, of the super
ficial generative hyphae at the growing apex. They have mucilaginous walls
and between them pass the divergent portions of the gloeocystidia.
Hymenium. From a discontinuous layer of basidia and short excrescent
gloeocystidia the hymenium compacts and thickens by sympodial-- growth.
Slender laterals, c. 1 · 5 µ, wide, arise from the subterminal cells below the
basidia and grow out shortly beyond the mature basidia: they often cut off a
subterminal cell before becoming basidia themselves. The gloeocystidia
lengthen apically at the same rate and project little, if at all, beyond the general
level of basidia. All the excrescent hyphae have rather toughly mucilaginous
walls so that the hymenium has no air-spaces and, macroscopically, a grey,
rubbery-gelatinous appearance. It seems that few basidia mature, becaus�
most, in the many sections which I studied, were sterile: spore-prints, too, ·
after 12 hours, were thin.
Epihymenium. Eventually the tips of the sympodial hyphal branches
which thicken the hymenium lose their palisade arrangement and become
closely interwoven to form a thin, mucilaginous plectenchyma of sinuous
irregular cells 1 · 5-3 µ, wide. The formation of the epihymenium apparently
renders the hymenium sterile.
Structure of the Cyathiform Tips. The shallowly cup-shaped or funnel
.U:
shaped ends of the branches are sterile. They are lined by the interwoven
ends of the central hyphae of the branches and become thinly levigate with a
sterile submucilaginous layer, 5-15 µ, thick, just as the epihymenium. There
are no gloeocystidia. The depressions slowly fill with fallen spores.
·C
TEXT-FIG. 23. Clavicorona candelabrum: skeletal and generative hyphae and gloeocystidia:
a, the tip of a skeletal hypha become secondarily septate: b, an unusual branche1 tip of a skeletal
hypha: c, a generative hypha with thick-walled portion: d, longitudinal gloeocystidia developing
near the apex of a branch before curving into the hymenium, X 500.
Structure of the Secondary Mycelium. At the base of the fruit-body branches

of the generative hyphae grow out as secondary mycelial hyphae to form the
CLAVICORONA 57
compact disk of tissue closely applied to the wood. The hyphae are strictly
radiating and little interwoven. There are no skeletal hyphae or gloeocystidia.
Development of the Fruit-body. A conical primordium, 1 X o· 5 mm., develops
on the surface of the wood. It dilates upward, as it elongates, by multiplica
tion of the hyphae, and when it reaches a certain height, which is that of the
stem of the fruit-body, apical growth stops in the centre of the growing
points (in this immature state the fruit-body corresponds with the mature
state in the simple species). The cessation of growth proceeds centrifugally
to give the first subcyathiform apex, but it continues at four, rarely more or
fewer, points round the edge which form the first branches. During this
upgrowth of the primordial shaft the secondary mycelial disk begins to form
at the base and it continues its radial outgrowth slowly throughout the
development of the fruit-body. The branches develop exactly as the pri
mordial shaft and, on cessation of growth, produce new branches, peripher
ally, in the same way. They begin as minute points which become subulate
and then clavate-truncate, before the tops become cyathiform. The first
branches of the fruit-body generally diverge considerably from each other
before turning vertically upward, but the divergence gets less with subsequent
branching so that the branches become more and more erect or strict:
thus the main branches have wide axils, which are often emphasized by
the sagging of the outer branches under the weight of the superstructure.
Only in the first one or two tiers do all the branches develop equally:
subsequently, only one or two branches develop fully and the others,
particularly on the inner or central and crowded side of the whole fruit
body, abort in varying degree. There is no proliferation of adventitious
branches, nor is there any dichotomy or irregular polychotomy. The tiers
tend to be at the same levels, but many irregularities develop.
Microscopically the growth is as simple and direct. There is no secondary
elongation of the hyphae. The growing-point of the primordial shaft, or of a
branch, consists of a dense sheaf of longitudinal hyphae, 1·5-3 µ, wide, which
are much entwined. They are the ends of generative and skeletal-hyphae,
and through their apical growth the fruit-body lengthens. The skeletal
hyphae arise as laterals, or terminals, of the generative hyphae in the tissue
near the growing-point: their mediate portions are short but gradual. Some
of the cells of both kinds of hypha swell laterally to their mature size behind
the growing-point, but they do not elongate.
I have not been able to study in detail the microscopic origin of the branches,
but it seems that they arise from localized regions of continued upgrowth
rather than from new points of excrescence on the rim of the apical cups after
apical growth has been wholly arrested.
The hymenium begins to develop about 250-300 µ, behind the apex. As
usual, the apical growth of the external hyphae slows down; their ends are
left behind and turn out at right angles to the surface to form the first hy
menial elements. They branch or lobe irregularly and profusely, and the
lobes grow out as a barely continuous layer to form the first hymenium.
Basidia begin to mature 500-800 µ, from the growing-point. The hyrneniurn
develops on all sides of the primordial shaft and so the stem is fertile except
at the base, where there is a brief transition to the secondary rnycelium. The
hyrneniurn develops also on all sides of a branch, but on the uppersides of the
axils it does not proceed beyond the initial stages and soon becomes thinly
levigate and sterile, as on the tops of the branches.
The gloeocystidia are restricted to the peripheral layers of the fruit-body.
They arise as laterals, rarely terminals, of the generative hyphae in the outer
part of the growing-point, distal to the hyrn enium and before it in point of
time: thus they are absent from the sterile tops of the branches which are the
residues of the central or axial tissue of the effete growing-points. The rudi
ments of the gloeocystidia grow longitudinally and form firstly their longi
tudinal, cylindric and immersed, subhyrnenial parts. As they are shifted to
the outside of the growing-point, through multiplication of its central hyphae,
their apical growth slows down. They come into the region of the incipient
hyrneniurn and their tips develop a bulge at right angles to the surface. The
bulge grows into the distal, or hymenial, part of the gloeocystidium and the
flexure swells into the ventricose base. Some gloeocystidia never enter the
regions of the incipient hyrnenium and remain 51s longitudinal elements in
the outer part of the skeletal cylinder. It is possible, also, that others may
develop in the hyrneniurn as it thickens, though most of them, certainly, are
primary and rooted in the subhyrnenium. The transversely elongated vacuoles
of the mature gloeocystidia make them appear banded.
I have no observations on the rate of growth of the fruit-body or on its
longevity but, from general considerations of weather and occurrence, it
seems that at least 2 weeks must elapse before the primordial speck develops
into the full-grown fruit-body and that there is a further 1-2 weeks of sporing.
The fruit-bodies are not readily putrescent. Spores develop on the stern
shortly after the origin of the first tier of branches.
Comparison with other Species. The construction of C. javanica is much
simpler. ... The flesh is almost homogeneous: there is no skeletal tube, but
where it should be there are merely numerous longitudinal gloeocystidia.
In the central tissue, corresponding with the pith of C. candelabrum, there
are scattered irregular gloeocystidia in all stages of development, and many
elongate cells of the hyphae have more or less oleaginous contents so that
there is a diffuse 'gloeocystidial network' (the fluid in which the fruit-bodies
are preserved and aqueous mounts become very oily). Many gloeocystidia
are added in the-hyrnenium as it thickens. Whether the inflation of the hyphae
behind the growing-point is longitudinal as well as transverse in this species,
and in C. pyxidata, I have not been able to determine.
In C. pyxidata and C. Colensoi there is an incipient skeletal tube formed
rather compactly of longitudinal hyphae and gloeocystidia, but it merges
gradually into the pith in which gloeocystidia and oleaginous hyphae evidently
oc-ur as in C.javanica.
( 59)
CLAVULINA Schroet.
Definition. This genus, proposed in 1888 by Schroeter for the three com
mon north temperate fungi Clavaria cinerea, C. cristata, and C. rugosa,
has met with little acceptance. Karsten added a fourth species from Finland,
C. odorata (1889), which is still doubtful. Van Overeem added a fifth, C.
Leveillei (1923), from Java. Then Dank, 1933, drew attention to its impor
tance by withdrawing it from the Clavarioideae and placing it in the Can
tharelloideae: he added C. amethystina. I now bring the number of species
to thirty-two, by adding
2 species from Europe: C. Bessonii,* C. gallica.*
4 species from N. America: C. amethystinoides, C. ornatipes, C. fioridana, C. casta
neipes.*
9 species from S. America: C. cartilaginea, C. chondroides, C. cirrhata,* C. connata,
C. delicia, C. pampaeana,* C. panurensis, C. pilosa,* C. Sprucei.
1 species from tropical Africa: C. Vanderystii.*
5 species from Malaya: C. cavipes, C. decipiens, C. gigartinoides, C. gracilis, C. ingrata.
5 species from Australasia: C. complanata,"' C. humilis, C. subrugosa,"' C. tasmanica,
C. vinaceo-cervina."'
Ten of these, marked *, are dubious to the extent that I have not examined
specimens, but the descriptions clearly indicate this genus which is well
characterized by the spore, the basidium, and the usually flattened, often
cristate, branching. In place, therefore, of a small north temperate group,
there appears a tropical genus of considerable size, to which many new species
will probably be added. One species, C. umbrina (Lev.) van Ov., I have
excluded as a Clavulinopsis. Six of the species, which I place in Clavulina,
have been referred to Lachnocladium s. lat., on the evidence of the texture of
dried specimens, and it may be remarked that if C. cinerea and C. cristata had
been described from dried material they would have suffered the same fate.
The peculiarity of the Clavulina-basidium is its subcylindric form, the
presence of 2, often strongly incurved, sterigmata (occasionally only I sub
median straight sterigma), and a fine secondary septum which appears in it
after spore-discharge, about half-way or two-thirds along, and by which the
residual cytoplasm is evacuated from the effete basidium. A further point is
the relation of the width of the spore to the width of the basidium which, in
other Clavarioid fungi, is about o·6 (the spore being narrower than the
basidium), but in Clavulina is about 1 ·25, the spore being generally wider
than the basidium. The detailed analysis of the Clavulina-b:::sidium must be
deferred until the basidia of Clavarioid fungi can be reviewed in full. It must
suffice to affirm that Clavulina has a peculiar basidium which at once dis
tinguishes its species.
Exceptions are few. C. decipiens, from Malaya, may have brown spores,
for the spore-wall, like the hyphal walls, is slightly thickened and pale brown
under the microscope. C. ornatipes and C. Leveillei may have multiguttulate
spores instead of the characteristic single large gutta. Dank mentions that
3-4 sterigmata occur occasionally in C. amethystina, but this fungus has been
much lX>nfused with Clavaria Zollingeri (2-4 sterigmata). Kauffman, also,
reports 3 sterigmata occasionally in C. ornatipes. In the hundreds of basidia
of the twenty-one species which I have studied, I have never seen more than
2 sterigmata·: hence I feel that even such exceptions need confirmation and
illustration: The only instance consistently without the secondary septum in
the basidium that I have met is C. cristata var. incarnata, but it may also be
absent from many basidia of C. dnerea f. sublilascens.
FORM AND STRUCTURE OF THE FRUIT-BODY
Form. In the different species the fruit-body may be highly branched,
sparsely branched, or simple. Many of the branched species produce in a
depauperate state simple or subsimple fruit-bodies, so that the simple con
dition is obviously reduced. C. cristata, C. amethystinoides, C. cavipes, C.
rugosa, and C. floridana have such a gradation, C. rugosa being particularly
difficult to classify in this respect. On the other hand, C. ingrata, C. gracilis,
and C. tasmanica appear to be invariably simple. The branching is radial in
C. cinerea, C. amethystina, and, perhaps, in some of the South American
species, but in the others it is typically flattened and palmate. In C. cartila
ginea, C. ornatipes, and some states of C. Leveillei, the flattened branching
continues in one plane, as in Scytinopogon, though the branches become
twisted secondarily into different planes. C. cristata and C. Leveillei may have
radial branching during the early stages of development, which passes into
flattened branching in the later stages. The details for most species require
elucidation. Most of the smaller species are simple, noticeably the minute
C. Bessonii and C. pampaeana, with fruit-bodies scarcely 1 cm. high, but C.
humilis and, particularly, C. delicia, though less than 2 cm. high, are branched.
The cristation of the branches, which occurs in many species, as C. cristata, is
a peculiar feature of the genus not found in other Clavarioid fungi: it indicates
a loss of control over apical growth resulting in the outgrowth of minute
fascicles of hyphae, as rudimentary primordia, from the ends of the branches
and is comparable with the production of strigose fibrils on the stem in C.
ornatipes and C. cartilaginea.
Colour. Little is known of this fundamental character beyond its macro
scopic aspect.. C. ornatipes and C. decipiens have their brown colour in the
hyphal walls, as in Lachnocladium. According to Singer, this is also the
position of the grey colour of C. floridana, which turns brown with alkali.
The purple-brown of C. Leveillei, however, is a thin incrustation on the
basidia: the pale yellow of its fruit-bodies is diffused in the cytoplasm (not
in the vacuoles) and, apparently, in the walls of the superficial hyphae. In
C. cinerea and its form sublilascens the colour is caused by the grey or purplish
vitreous cytoplasm at the apex of old collapsed basidia, and hence the older,
thickened hymenia are the darkest. Bright yellow, orange, and red, as in
Clavulinopsis, are noticeably absent.
Hyphal Construction. The construction is simple and normal for a Clavarioid
CLAVULINA 61
fruit-body, such as Clavulinopsis or Ramaria. Growth is by a fascicle of
narrow hyphae at the ends of the branches and the hymenium is com:ructed
by the out-turning, or unilateral inflation, of the hyphal tips which lie on the
sides of the growing-point: thus the subhymenial cells lie at right angles to
'i
TEXT-Frc. 24. Clavulina ornatipes: hyphae from the incipient hymenium

(left): superficial hyphae of the stem (centre): inflated medullary hyphae
of the flesh (right): only the basidia and subhymenial hyphae have clamps,
X 500.
the longitudinal cells of the flesh. All the medullary hyphae inflate, but in
most cases the central hyphae inflate more than the peripheral which they
stretch: thus the core loosens but presses outward and the cells with greatest
elongation often become curved or bent double to give the false appearance
of interwoven hyphae. In C. gigartinoides the outer hyphae on the upper
branches inflate as much as the inner, and the subhymenial hyphae also inflate.
In C. ornatipes, in contrast, with little inflation of any of its hyphae, the
peripheral cylinder is rather wide and dense and, in the stem, it thickens by
upgrmuth of narrow hyphae appressed over the surface. C. decipiens, also with
little inflation, has the most compact structure and 'Pteruloid' form, so that
at first sight it does not appear to be a Clavulina.
Stem. In C. ornatipes and some specimens of C. cartilaginea (at least in
Malaya), the stem becomes covered with short, conical, ascending, strigose
processes consisting of fascicles of narrow hyphae, just as the fas�icles com
posing the cristate tips of the branches. In both species the stem is thickened
TExT-FIG. 25. Clavulina cartilaginea: hyphae constructing the hymenium: without clamps,
X 1,000.
by the upgrowth of narrow hyphae on the surface and, when a sufficient

thickness (or excrescent field) is formed, the outgrowth becomes loc�lized
by cristation because there is no directive leader to control the field. The
process�s may reach 1 -2 mm. long, when their internal hyphae begin to
inflate, as at the growing-tips of the branches, and occasionally a process may
grow out into a slender, fertile, adventitious branch of normal construction.
At the base of the fruit-bodies there are all transitions from microscopic
processes, consisting of a few concrescent hyphae, to abortive primordia,
each composed of several thousand hyphae.
In C. cartilaginea the stem of Malayan specimens is generally covered by a
cortex composed of rows of globose inflated cells, forming an almost pseudo
pa..�nchy.matous layer on the surface. It resembles greatly the cortex of a
multifilamentous alga, e.g. Mesogloia or Halymenia. The cortex passes
gradually into the hymenium on the branches by the appearance of narrow
hyphae bearing basidia at the surface. The narrow hyphae may arise from
inflated cortical hyphae or from the outermost longitudinal hyphae of the
medulla. The algal appearance is heightened by the slow outgrowth of the
cortical filaments during the life of the fruit-body.
CLAVULINA 63
Hymenium. The presence of the hymenium varies in different species and
may be of systematic importance when the details are better known In at
..... ........ .,..,_.,.---------"'""'""�-"'""�--------A..-

._._____.._.·;;,;;;?........
- ..
TEXT-FIG. 26. Clavulina cartilaginea: the pseudo-parenchymatous cortex on the stem shortly
below the hymenium, with a few precocious oasidia, x 1,000.
least eight species it is unilateral, the upper sides of the branches being covered
with a sterile hymenium. The following arrangements occur:
(i) Hymenium covering the whole fruit-body except the growing-tips
and the extreme base of the stem (for 2-5 mm.): C. cinerea, C. cristata, C.
rugo:;a.
(ii) Hymenium absent from a distinct, sterile stem: C. floridana, C. graci
lis, C. ingrata, and C. cavipes (stem without a cortex).
(iii) J:.Iymenium absent from the stem (or the greater part of it) and
absent from the upper sides of the main, oblique, branches and the. upper
sides of the axils: C. cartilaginea, C. connata, C. cristata var. incarnata, C.
delicia, C. decipiens, C. gigartinoides, C. Leveillei, C. ornatipes, C. Sprucei.
The dorsiventrality of the third group is associated with the flattened and in-
clined, or oblique, branching, which is a deflection from negative geotropism.
It appears that the hymenium thickens in all species of the genus.
Subhymenium. In most species the tissue is filamentous, but in C. cartila
ginea, C. decipiens, and C. ingrata its cells inflate and become pseudoparenchy
matous. In C. gracilis it is nearly pseudoparenchymatous. In C. gigartinoides
it is filamentous on the lower branches but becomes pseudoparenchymatous
on the upper. The filamentous subhymenium appears as a reduction from the
pseudoparenchymatous by loss of inflation normal to the hyphae. The
difference is important in as much as the rugose surface develops in the species
without inflating subhymenium to compensate the intercalary expansion of
the hymenium, e.g. C. rugosa, C. cinerea. The inflation of the subhymenial
cells is comparable with that of the cortical cells on the stem of C. decipi"ens
and C. cartilaginea.
Sterile Hymenium. The construction of the sterile hymenium on the upper
sides of the dorsiventral branches is similar to that of the hymenium, but the
tissue does not thicken so much and the basidia are never fertile though they
may bear 1-2 abortive sterigmata without spores. On the stem the sterile
hymenium is generally loose and may consist only of divergent hyphal ends
with subclavate terminal cells, between which and fertile basidia there is
every transition at the apex of the stem where the hymenium begins. C.
decipien� is excepti9nal in the compact, thickening, sterile hymenium on the
stem, comparable with the cortex of C. cartilaginea. The sterilization of the
basidia on the upper sides of oblique branches shows that a negative geo
tropism inhibits the formation of the spores, and this is the mark of the
'inferior hymenium'. That on the stem, in contrast, indicates rather the time
when the primordial shaft begins to develop the hymenium in the diageo
tropic position normal for the Clavarioid fruit-body (I have seen no humi
colous, or terrestrial, Clavarioid fruit-body which is fertile from the very
beginning, as happens in some of the small 'epiphytic' species of Pterula).
Spores. The spores are mostly subglobose, 7-10 /.I- long, with the proportion
of length to width (E) as 1·1-1·3, rarely 1·5 in C. rugosa. They have, thus,
little diagnostic value for most species, but the little C. delicia, of Brazil,
is peculiar in having distinctly ellipsoid spores, 9·5-u X 5-6 /1- (E about 1·9):
nevertheless, it is connected with the normal state by C. connata (E 1·3) and
C. Sprucei (E 1 ·2). The-smallest spores, c. 5 X 4 /.I-, occur in C. gallica (which
CLAVULINA 65
may be a 2-spored Clavulinopsis), and the largest in C. rugosa (9-14X 8-11 µ).
In all species the wall is slightly thickened and in C. decipiens it evide.1tly be
comes pale brown, as do the hyphal walls (but I have not seen a fresh spore
print). Dried spores, particularly in the herbarium, may turn yellowish or
ochraceous, e.g. C. cinerea. The oil-drop generally fills most of the spore:
Malayan specimens of C. ornatipes and C. Leveillei may be exceptional with
multiguttulate spores.
TEXT-FIG. 27. Clavulina cristata: diagrams of the structure of a cristate

branch-tip and of the mature tissue of a branch, showing the loose central
and dense peripheral parts of the medulla, and the thickened hymenium.
Basidia. The secondary septum, which cuts off the completely vacuolate
distal half, or third, of the basidium after spore-discharge, is a constant and
noteworthy feature. In some species it is followed by several more septa
formed successively in the proximal part as the protoplasm withdraws. It
is reported for the three common European species that the fusion-nucleus
of the basidium divides into 6 or 8 nuclei of which I enters each spore and
Sll9 F
4 or 6 remain in the basidium. 1 What happens to these is not known, but the
cytoplaam of the basidium withdraws into its proximal part and may pass into
the subterminal cell carrying the residual nuclei with it. I have· seen no
hyphae develop directly from the proximal part of the old basidium.
The shape of the basidium is generally described as subcylindric, but it is
really very narrowly clavate, being at its greatest width nearly twice the
diameter of its base. In C. gracilis and C. ingrata it is distinctly clavate.
The short measurements given by several authors for the basidium probably
refer to the distal part of the basidium beyond the secondary septum.
The sterigmata are characteristically horn-shaped, outcurved and then
incurved, as in Craterellus and Cantharellus. In C. decipiens and C. cartila
ginea they may be nearly straight.
Cystidia. C. Leveillei is the only species which certainly has cystidia. They
are well developed and grow out continuously with the hymenium which the
primary ones traverse. Nevertheless, they seem to be disappearing because
they may be very few and even absent (var. atricha). C. pilosa, which I have
placed in Clavulina by analogy, is also said to have cystidia, but they are
inadequately described and may be young emergent basidia. In C. cristata
var. incarnata there may be rudimentary cystidia developed under very moist
conditions, but they are hardly comparable with those of C. Leveillei which
are to be interpreted as overgrown basidia. Possibly cystidia were a normal
feature of ancestral Clavulinas.
Inflation. In most species the hyphae inflate. In C. cartilaginea the cells
may inflate irregularly, one or more parts of the cell inflating appreciably, the
rest of the cell little, if at all. In C. decipiens and C. ornatipes inflation is slight,
and this feature, coupled with the small growing-points, gives the Pteruloid
appearance to C. decipiens. The inflation mechanism seems to be degenerat
ing in several species, and the different appearances of the cells suggest that
several processes are involved-deficient turgor-pressure, loss of interstitial
growth of the cell-wall, or deficient swelling of the wall, loss of elasticity of
the wa!l, and increased rigidity by thickening of the wall. The irregular
curvature of the branches in such species as C. cartilaginea and C. Leveillei
shows the irregular inflation of the hyphae.
Clamp-connexions, Secondary Septation. The presence or absence of
clamps is remarkable in this genus and seems to have, at least, specific
importance. Most species have clamps at all septa, indicating that the fruit
bodies are diploid, not haploid as the 2 sterigmata suggest. The following
are exceptiona�:
C. ornatipes has no clamps on the longitudinal hyphae of the fruit-body,
but most of the basidia and subhymenial hyphae are clamped.
C. cartilaginea, C. connata, C. delicia, C. fioridana, and C. Sprucei have
no clamps, not even on the basidia (very occasionally there may be a clamp
on a medullary hypha in C. cartilaginea).
1
Maire (1902), Juel (1916), Bauch (1927).
CLAVULINA
In these six species without clamps, the medullary hyphae are more or less
secondarily septate. This septation is very slight in C. ornatipes with .:hicken
ing hyphal walls, fairly frequent in C. cartilaginea and abundant in C.
fioridana, but never so copious as in typical Clavaria s. str.: it does not occur
in any species with clamps. Whether the absence of clamps and the tendency
to secondary septation indicates affinity of the species is difficult to say, because
(i) C. ornatipes is most nearly related in other respects with the clamped
C. decipiens,
(ii) C. connata is so similar to the clamped C. panurensis as to appear
almost identical in other respects,
0 0 C)
- -
TExT-FIG. 28. Clavulina ornatipes: stages in the flattened branching of a branch-tip,
the two figures on the right showing polychotomy, X 3.
(iii) C. fioridana and C. cartilaginea are very different from each other
in branching, texture, and microscopic structure of the stem. C. fioridana
may be related to C. Leveillei, and C. cartilaginea possibly with C. chon
droides.
DEVELOPMENT OF THE FRUIT-BODY OF C. ORNATIPES
Development. A cylindric primordial shaft, constructed exactly as the tip
of a branch, produces the stem. Its apex is conical and acerose until it
broadens, flattens, and breaks up into several new growing-points like minute
teeth, which are the origins of the primary branches. The strigose fibrils
develop from below upwards, as concrescent fascides of narrow superficial
hyphae and, in longitudinal section, they have exactly the construction of
small primordia. At the base of the stern, indeed, there are all transitions from
these conical fascicles to abortive primordia and on the stem itself one or two
of the fascicles occasionally enlarge into a short branch which may even
become fertile and branched itself. The hymenium begins t9 form 2-6 mm.
behind the growing-point of a branch and its construction is equally gradual,
taking place over a length of 1-2 mm. before the first compact layer of basidia
is formed. It is constructed in the usual way by the out-turning of the super
ficial hyphal tips, the first cells of which become basidia, and then, by sym
podial branching and outgrowth of laterals, it thickens continuously: it does
not reach the same thickness as in other species because of the shortness of the
basidia.
Branching. In the Malayan specimens and, as it seems from the photo
graphs,•,in the North American specimens, branching is flattened or palmate
multifid in one plane , though by unequal g rowth of the t issues the branches
c ome to lie in different planes, and only the attenuate final tips are dichoto
mous . The apex of the primordial shaft and of the branches is conical and
acute while actively growing. The coni-
cal shape is caused by the production of
,fi�lj\\J/ 1
lateral hyph�e e q�ally all round the apex
i1mltl l\\ \ 1i
/ 1 and by the mfla t10n of the cells a crope-
$//1�//'11 \\\\\
��ii�11l \· (i
t ally. At c ertain periods the conical apex
if1i1l 1 mt
{,N/ii1i /11/ j
i ! t :�:.��:�·::;��::�i::��:=�
becomes blunt, then flattened and t�inn�d
l 1 \)��
11 1 1
t�Jfi/1;1�(/1/
i\11 \
�ii11 U; i/fl il\��l .

short spac� of longitudinal growth 3-'/
s a
��:�:::: �o ;i:;:::::r:�e 'fn��:i:

#J1J)I;1aI/. 1j111 l \ \\1\\\i\
JiJ;iJ/f/Mr/ /.1i;
ma� cylindric al bra�ches wh ich, af�e: a
/I I
I \( \\ \ \\��\I r::�: ::m::::. t�/�:::�e� :::� !;
i 0 u i
\\\\\\\�\Ii\\
I
j branches are thickened by the up-

the
. ii/�/ I· / \ \ \i\\\\\�11,.
growth of hyphae over their surface and
1� /t / 1 / /) 1 then by the development of the hymen-
\ \'\1\r,
;�l;/l;l 111 \\�1\\1tr:

1
1/J
j j 1 \ � ch al
��:e fr::::t:!:o�;z;· !�: �:�t:s :�
)f/J/;;f() ft/(
! IJ I 11 0f
\\1l\t,
{.
;1 rf
/ �;:��:ia:��:: �:1�: ;� :�;e, fr�:
jJi'1
(I \ / \\
j/
\\ hyphae which are out of range of the co -
1
. Jif;f//( / / ( \ \ .
/
ordin ati ng force of the tr�e ap ex: bu t
! II / {
/ \ \�
, /if ( ! /r I {/ j / 1 /) ·/ 1 // \'\ \i\ I§:
� ·. such branches are sca. rce, like those. on
;f/Ji//,/t// ,/ / / I/ (\ f
/ / / / the s te1:1. _I have �ot s<:�n sufficient
/
J
1 j f / I /, / 1 1 / , I / , l ! i youn? _ t�ps m the bn<:f critical st_a�e. of
· subdivis10n to be certam that subdivis10n
TExT-Fm.29. Clavulinaornatipes: diagram is a lways by repeate d dichotomy: in some
of the structure of a branch-tip. cases it seems that the spathulate e dge
is simultaneously multifid. The adventitious branches indicate incipient
cristation.
CLASSIFICATION AND AFFINITY
Most tropical species are so little known that it is imposs ible yet to formu
late a natural classification . The specific key on page 295 is largely artificial .
The following groups appear to be natural :
1. C. cristata-group, discussed on pages 70 and 3 18, i ncludes C. cinerea,
C. amethystina, C. amethystinoides, C. subrugosa, C. gracilis, C. ingrata,
C. cavipes, C. gigartinoides, and, probably, C. Bessonii, C. pampaeana,
CLAVULINA
and C. gallica, if not most of sections C and D of the specific kr-y. The
tropical C. gigartinoides appears as the best-developed member, but it is
practically colourless.
TEXT-Fie. 30. C/arnli11a ornatipes: diagram of the structure of the base of die
stem showing the rudimentary primordia forming the strigose processes: one
process on the left has developed inflated hyphae in the medulla, as in the main
stem of the fruit-body.
2. C. ornatipes and C. decipiens with brown-walled, little-inflated hyphae,

and short basidia.
3. C. connata, C. delicia, C. panurensis, and C. Sprucei seem a group from
tropical South America with rather loosely fibrous flesh, at least when
dry. Possibly C. cirrhata belongs here. It is the only group with a
lignicolous tendency (C. delicia, C. Sprucei): possibly, therefore, C.
humilis of New Zealand also belongs.
4. C. Leveillei (with cystidia) and C. floridana (without clamps) may be
related as branched and simple species, for their coloration is similar.
But C. Leveillei, in most other respects, is near C. cristata; it retains a
trace of yellowish, possibly ancestral, pigmentation.
5. C. cartilaginea and C. chondroides have similar and distinctively tough
flesh, drying cartilaginous.
Regarding the affinity of the genus, it is remarkable that in form and colour
the branched Clavulina amethystina and Clavaria Zollingeri are so much alike
that microscopic examination may be necessary to distinguish them by their
basidia. Similarly with the grey Clavulinopsis cineroides and Clavulina cinerea,
and the white Ramariopsis Kunzei, Clavulinopsis subtilis, and non-cristate
forms of Clavulina cristata. In other words, Clavaria, Clavulina, Clavulin
opsis, and Ramariopsis seem referable to a common, branched, lilac ancestor
with grey and colourless derivatives. The oddity of Clavulina is its narrow,
consistently 2-spored, subcylindric and 'stichic' basidia. Donk has par
ticularly emphasized the stichobasidial character (with the spindle of the
first division of the fusion-nucleus longitudinal), for which reason he has
placed Clavulina near Cantharellus, Craterellus, and Hydnum s. str. (H.
repandum). It is not obvious, however, why a narrow basidium, unable to
accommodate for this reason a transverse spindle, is a sign of affinity and I must
leave the point until a fuller discussion of the basidium is possible. It is note
worthy that the old basidia of Clavaria helicoides may also empty by secondary
septation.
GEOGRAPHICAL DISTRIBUTION
The genus is clearly tropical in that most species with the greatest variety of
structure occur in the tropics. Doubtless the Andes, the Himalayas, and the
mountains of tropical Africa and Polynesia will add more species. The com
mon north temperate species of the C. cristata alliance have not been found
in the tropics and only C. cristata, C. cinerea, and, possibly, C. rugosa have
been reported from the south temperate region (as yet little known mycolo
gically): however, in Malaya C. cavipes, C. gigartinoides, C. gracilis, and C.
ingrata represent the group. C. cartilaginea is apparently world-wide in the
0
tropics, but the C. connata-group seems characteristic of South America.
Most remarkable at present is C. ornatipes, first described from the United
States where it appears to be temperate or montane, then from Japan, and
now I record finding it in one place on Singapore Island in the lowland
tropical forest: there can be no doubt of the identity of the Malayan fungus,
and. its apparent rarity shows how little is yet known of tropical fungi. Indeed,
in all its aspects, Clavulina awaits interpretation from tropical material.
C. CRISTATA ALLIANCE
The five north temperate species, C. cristata, C. cinerea, C. rugosa, C.
amethystina, and C. amethystinoides are so nearly related that their distinction
and classification are difficult and unsatisfactory. No two authors agree on
their specific limits, seldom even on their denotation, and the classification
which I offer is yet different from the previous. The variability of the first
CLAVULINA 71
three is apt to be ignored because they are common: that of the last two is
scarcely known because they are rare or uncommon. Coker regarJed the
first four as variations of C. cristata: he cited Romell as having come to the
same conclusion in Sweden, and Wehmeyer has, more recently, supported it.
These authors have not analysed the variation thus incorporated in C. cristata
or explained how a simple white C. rugosa can be conspecific with the branched,
purple C. amethystina. I am sure this extreme view is unacceptable. On the
other hand, some mycologists recognize such species as C. canaliculata, C.
albida, C. coralloides, and C. grossa, not to mention Britzelmayr's host, which
others regard as varieties of the three common ones. I have attempted a
compromise with as much precision as possible so as to guide future investi
gators. What is needed is detailed descriptions, microscopic as well as macro
scopic, of the troops 1 of C. cristata, C. cinerea, and C. rugosa which show
distinguishing characters, however trivial they may seem: and, also, detailed'
descriptions of all collections of the rarer two. By this means only can myco
logists profit from each other's collections, for no one can personally become
acquainted with all their variations even in a lifetime devoted to the purpose,
and by such co-operation knowledge can be advanced rapidly. Macroscopic
descriptions and paintings have no use without adequate microscopic details
which an hour's study of fresh material will supply: the confusion among the
species has arisen largely from the attempt to fit the old illustrations to modern
microscopic diagnoses-a procedure little more than guess-work.
In the north temperate region the species are widely distributed except C.
amethystinoides, which appears limited to North America. C. cristata and
C. cinerea are the only species reported with certainty from the south tem
perate region (Australia, New Zealand, South Africa), and there is no certain
record of any from the tropics: neither Petch nor van Overeem nor myself
has found them in tropical Asia.
As a group they are distinguished by their clamped hyphae with colourless
walls, uninflated subhymenial hyphae, absence of cystidia, and absence of
dorsiventrality, so that the hymenium covers the whol<:! surface of the fruit
body except the branch-tips and the base of the stem. They differ specifically,
or at least appreciably, in manner of branching, colour, and spore-size. In a
critical study the following considerations should be borne in mind.
Colour. The grey, pink, lilac, and purple colours of C. cinerea, C. amethys
ti'noides, and C. amethystina may be variants of the same pigment, or there
may be a grey pigment distinct from a pinkish-purple. The distribution of
the pigment in the tissue may be important. It is not even known whether
the grey colour which appears in some varieties of C. cristata is the same as
that of C. dnerea.
Branching. Each species can have its simple form, either as the initial
developmental state, or as a juvenile form arrested in this state or as a de
pauperate state. The simple states of C. cristata and C. rugosa, of C. amethys
tina and C. amethystinoides, or of C. cinerea and C. rugosa var. fuliginea may
1
A troop is generally derived from one mycelium which perpetuates the peculiarities.
be indistinguishable macroscopically. Thus, C. grossa is a mixture of simple
and subsimple states of C. cristata and C. rugosa which can be distinguished
from their spores. Some of the simple states which have been described are
clearly only young stages, e.g. Coker's figures of C. amethystinoides. The
manner of branching may also be important: C. cinerea and C. amethystina
appear to -have radial branching, the others radial followed by flattened
branching.
Cristation. It seems that cristation is typical of C. cristata and that it occurs
also in some branches of C. cinerea and C. amethystinoides. The opposite
extreme with blunt radial tips may also occur in these species, as it does in
C. cartilaginea which has 'cristata' and 'rugosa' forms mixed in the same
troop. It is not known how the cristate and blunt forms of fruit-body occur
in the other species.
Size. This is a variable feature and of little systematic value unless analysed
in terms of cell-inflation: a small size may be caused by early maturity or
insufficient cell-inflation, and both may be peculiarities or effects of depaupera
tion or desiccation. It is generally sufficient to determine the average length
of the main inflating cells half-way between the hymenium and the centre of
the stem or branch to analyse the size of the fruit-body, but if the hyphae
become secondarily septate there is no ready method.
Spores. Coker pointed out that some of the collections which he referred
to his macro-species, C. cristata, had slightly larger spores than the others,
and that they generally had the simple or sparingly branched, blunt fruit
bodies of C. rugosa rather than the more slender cristate fruit-bodies of C.
cristata, but that there were many intermediates confusing the distinction. I
have emphasized this difference as the mairi distinction between C. rugosa
(large spores) and C. cristata because I think it is more important than the
other characters of the fruit-body. Thus I resolve C. grossa auctt. into a
subsimple, bluntly branched state of C. cristata (small spores) and a sparingly
branched state of C. rugosa (large spores). Coker appears not to have dis
tinguished young or depauperate, subsimple fruit-bodies from adult and
typical fruit-bodies, and thus has arisen, no doubt, the confusion. I do not
find any critical evidence that this spore-difference is not constant, and the
list of authentic measurements given in Table II certainly indicates slightly
larger spores in C. rugosa than in the other four. It should be noted, also, that
fruit-bodies of C. rugosa generally turn lurid yellow in drying.
Habitat. Bresadola says that C. cinerea in coniferous woods has smaller,
less-branched, stouter, and violaceous-cinereous fruit-bodies with whitish
tips, and in frondose woods taller, more-branched, slender, and grey fruit
bodies with blackening tips. The first state suggests C. amethystinoides.
The short and much cristate varieties of C. cristata appear to develop in
pastures and heaths, the more elongate and less cristate in woods. C. rugosa
in sphagnum-bogs is certainly stouter and more often branched than in dry
places. C. cristata is more or less caespitose but C. rugosa is solitary or
gregarious.
CLAVULINA 73
Rosellinia Clavariae. This pyrenomycete, and its imperfect state ( Scoleto
trichum Clavariarum), attacks the fruit-bodies of C. cristata, C. cinerea, and
C. rugosa, perhaps also the other species. It turns them grey, dark fuliginous,
or blackish, and alters, perhaps, the shape of the branches, so that the parasi
tized fruit-bodies of the three species superficially resemble each other, and
thus it has added very materially to their confusion.
Geographical Distribution. In subtropical countries or in the subtropical
and temperate zones of tropical mountains these temperate species may mix
with allied tropical species.· Thus, in Malaya C. gigartinoides and C. cavipes
are certainly allied, and perhaps also C. cirrhata (Brazil) and C. Vanderystii
(Congo), though the tropical species generally have some dorsiventrality of
the branches. From the Himalayas we may expect critical collections of all
the species.
TABLE II. SPORE-SIZE IN CLAVULINA CRISTATA, C. CINEREA, C. RUGOSA,

C. AMETHYST/NA, C. AMETHYSTINOIDES
Species
Spore-si::e in µ Authority (Synonyms in brackets)
9-12X7-9 France Bourdot and Galzin C. rugosa
9-12X7-10 f. mitruloides
8-12X7-9 (C. canaliculata)
9-12x6-9 " " (C. grossa)
9-11X8-10 Gt. Britain Cotton and Wakefield C. rugosa
9-13X7-10 Corner
9-13·5x7·5-12 " "
11-12x7 Rea (C. canaliculata)
9-12X 6-8 " " (C. grossa)
12-14x8-10 Germany Britzelmayr var. macrospora
8-12X7-10 Holland Dank C. rugosa
8-10 7-9 Italy Bresadola
7-10 China Teng
7-fJX6-8 France Bourdot and Galzin C. cristata

6-8 4-5 (C. coralloides)
6-10 5-'7'5 " (C. Lappa)
7-<1'5 x6-7·5 " Konrad and Maublanc C. cri,tata
7-<)(-12) 6-8 Gt. llritain Cotton and Wakefield
7-8 6-7 Rea
7-10 )( 6·5-8 Corner
7-10·5'<'6-9 var. incarnata
7-10x6-8·s " " var. subrugosa
8-12 .6-8 IIolland Dank C. cristata
7-9 , 6-8 , Italy Bresadola "
7-9 .< 6-7 " (C. albida)
7·5-9X6-7· 5 " Coker (Herb. Bres.) "
7-8X6-7 Asia Minor Pilat C. cristata
9-10X6-8 Algeria Patouillard (C. comosa)
7'5-9X5·5-7·5 Australia Cleland C. cristata
7-8·5 China Teng
7-1ox5-7 Rhodesia v. der Bijl "
8-10 U.S.A. Lloyd (C. laciniata)
7-8X6-7·5 Coker (C. sphaerospora)
TABLE II (continued)
Species
Spore-size in I-' Country Authority (Synonyms in brackets)
7-1ox6-9 France Bourdot and Galzin C. cinerea
8-10x7-7·5 var. odorata
7-12X6-7·5 " (C. fuliginea)
7-1ox7-8 " Konrad and Maublanc C. cinerea
7-1ox6-8 Gt. Britain Cotton and Wakefield
7-11x7-8 Rea "
9x8 " var. gracilis
8·5-1 l X7-10 Cornei· C. cinerea
8·5-10·5X6·5-8 " " ,, f. sublilascens
7-1ox6-8 Holland Donk C. cinerea
7--9 Italy Bresadola "
7·5-1 l X6·5-9·5 Europe Coker (Herb. Pers.) (C. fuliginea)
8-11x7·5 Australia Cleland C. cinerea
6·5-8·5 China Teng
7·7-10X7 U.S.A. Povah
7-1ox6-8 France Bourdot and Galzin C. amethystina

10-12X8 Germany Britzelmayr "
9X7·5 " var. lilacina
9-12x7 Killermann C. amethystina
7-8 " var. lilacina
1ox7 " Schroeter C. ametl,ystina
10-12x7-8 Italy Saccardo "
8-10X6-8 var. lilacina
7·5-8·5 X 6-7·5 U.S.A.(S.Car.) Coker C. amethystinoides

7·5-9 U.S.A.(N.Car.) ,,
7·5-1ox6-8
7·5-8·5X5·5-7·5
10·6X8·2 Average C. rugosa

9·9x7·3 C. amethystina
9·0 X 7·6 C. cinerea
8·3 X6·8 C. amethystinoides
8·2 X6·7 C. cristata
CLAVULINOPSIS van Ov.

DEFINITION
In this genus I have placed Clavarias with white spores and clamped,
inflating, thin-walled, but not secondarily septate, hyphae. They centre
round C. corn£.,--ulata, C. fusiformis, and C. subtilis, and thus form the largest
and central group of white-spored Clavarias. The selection of Clavaria
vermicularis as .the type of Clavaria s. str. necessitates this new genus which
seems to embrace most, if not all, the yellow or orange, but white-spored
Clavarias. The earliest generic name for the group is Cornicularia Bon.
(1851) based on Clavaria corniculata Fr., but it is a later homonym. Ramaria
S. F. Gray (1821) should strictly have precedence, as it included C. cornicu
lata, but the definition of Ramaria in this sense would be highly confusing
CLAVULINOPSIS 75
and, as discussed on page 124, is not to be recommended. It happens that
van Overeem described in 1923 aJavanese Clavaria as Clavulinopsis f'ulcata
which belongs to this affinity and, as a monotypic genus, the name Clavulin
opsis must be accepted as validly published in a combined generic-specific
diagnosis, though no generic character was clearly indicated.
C. sulcata has simple, reddish-pink fruit-bodies and smooth, white, globose
spores c. 5 µ, wide. It is identical with Clavaria miniata B. (non Petch), which
is a common species of tropical Asia, but it often has abnormal or unusual
basidia which appear to have been the basis of van Overeem's new genus.
He emphasized the unusual and often very unequal length of the sterigmata
(8-20 µ,) on the same basidium, their irregular number (1-4, mostly 3, often
2 or 4, rarely 1), and their irregular position (in the usual subapical whorl or
at different levels or even in a straight row, ? transverse). The basidia of
typical C. miniata have 2-3-4 sterigmata, and some collections have pre
dominantly 3-spored basidia, but abnormalities, as described by van Overeem,
are not infrequent. In all other respects C. sulcata is identical with C. miniata.
Van Overeem's figures, also, hardly bear out his contention that the abnormal
basidia are characteristically preponderant: of ten drawings of basidia, six
have equal sterigmata of c9mparable length and, of these six, one has 2
sterigmata, two have 4 sterigmata, and three have 3 sterigmata, exactly as
normal basidia of C. miniata. Furthermore, such abnormally elongate
sterigmata occur in other species of the same alliance as C. arenicola, C.
brevipes, and C. aurantio-cinnabarina: they are sometimes natural abnormali
ties, but it must be remembered that they may also be unnatural and developed
after collection of the fruit-bodies when they have been shut up for some time
in a tin: monosterigmatic abnormalities then appear as cystidioles. Van
Overeem appears to have been obsessed with the idea that the unusual
sterigmata were as diagnostic as those of Clavulina and of Heterobasidio
mycetes, but he admits that Clavulinopsis is not nearly related to Clavulina.
In employing Clavulinopsis, therefore, for this group of Clavarias, one is
merely adhering to the rules of nomenclature, in honouring a mo...-i.otypic
genus, without accepting its definition, for the author seems to have been
unaware of the close affinity between C. sulcata and C. fusiformis. The type
species of the genus becomes Clavulinopsis miniata, as.the earliest valid name
(so far as known) for C. sulcata van Ov. So far as concerns the hyphal
structure, C. miniata, C. fusiformis, and C. corniculata are practically identical:
as a branched species C. corniculata would make a better type. Most of the
species of Clavulinopsis are, in fact, so nearly related that thflir classification
is difficult.
STRUCTURE OF THE FRUIT-BODY

Branching. About twenty species have branched fruit-bodies, the majority
being simple. Few are highly branched, as C. alcicornis, C. cineroides, and
C. subtilis, and several branched species have transitions to simple forms, as
C. biformis, C. corniculata, C. luteo-ochracea, C. rufipes, C. similis, and C.
tenella. The normally simple, if caespitose, species C. amoena and C. spiralis
occasionally have fruit-bodies with 2-4 simple or bifid, elongate branches.
Rudimentary branching seems to occur also in C. citrino-alba and C. fusi
formis var. ceranoides, which have usually shortly bifid tips on many of the
simple caespitose fruit-bodies. There is no evident relation between the size
of the fruit-body and branching: C. fruticula has small, much-branched fruit
bodies 1-2 cm. high, in contrast with the large simple fruit-bodies of C.
spiralis. There is also every gradation in the simple fruit-bodies to small
Typhula-like specimens as in C. brevipes and C. luteo-tenerrima, little epiphyl
lous specimens of which are distinguishable from Pistillaria only by such
microscopic details as the oil-drop in the spore, the yellow pigment, and the
superficial hyphae on the stem. The branching and the size of the fruit-body
have, therefore, minor classificatory value.
Flattened branching and cristate tips seem not to occur. The tips are
normally bifid in alternating planes and the two branches, initially divergent,
incurve to give the crescentic or rounded axils so characteristic of C. cornicu
lata. The early branching of some species may be polychotomous, but I have
seen clear evidence only in C. dichotoma. The siJnple branching is connected
with the comparatively small size of the primordial shaft: excepting the
doubtful C. Fleischeriana (stem 2 cm. thick) the stems of Clavulinopsis
are slender, 1-2 mm. thick, as the branches, and therefore unlikely to be
polychotomous.
Stem. A sterile stem, with scattered sterile basidia or scattered excrescent
hyphae, seems a typical feature of the genus. It is more apparent in the small
simple fruit-bodies such as C. filipes and C. brevipes, but I doubt if it is longer
in them than in the branched fruit-bodies. It represents the initial sterile
phase of the primordial shaft.
Habit. For many species the caespitose habit has been emphasized, e.g.
C. fusiformis, but I doubt whether it indicate.s any more than the richness of the
substratum. In C. amoena, C. miniata, C. spiralis, C. luteo-alba, C. luteo
ten�rrima, C. pulchra, C. corniculata, and, even, probably C. fusiformis, there
is every gradation from densely caespitose fruit-bodies, through gregarious
or 2-4 fasciculate fruit-bodies, to solitary ones. However, the caespitose
habit may be useful in distinguishing a typical state of a species, such as C.
spiralis, from that of the subsolitary C. brevipes, but more critical field
observations are needed to establish the character.
Habitat. A few species occasionally grow on rotten wood, as C. alcicornis,
C. appalachiensis, C. depokensis, C. globospora, C. miniata var. sanguinea, and
C. rufipes. C. luteo-tenerrima and C. miniata var. sanguinea may even be
epiphyllous, as in Typhula and Pistillaria. But the normal habitat of these,
as that of the other species, is on the ground or humicolous. Thus the genus
agrees with Clavaria, Clavulina, and Ramariopsis and contrasts with the
lignicolous Lentaria. Two species are phycophilous, namely, C. hastula and
C. vernalis: they are exceptional, as discussed on page 81, but C. calocera
CLAVULINOPSIS 77
and C. constans seem to connect them with the more normal, if reduced, state
of the genus.
Spores. The large oil-drop in the fresh spore is most characteristic. It
generally fills the spore almost completely and, on slight drying, the spore-wall
shrinks on to the gutta and the contents appear wholly vitreous. The feature
can be seen only in fresh specimens; the protoplasm of dried sp0res or of
those preserved in alcohol-formaljn undergoes some physical alteration causing
the gutta to disappear. If a fresh spore is not 1-guttate, the structure of the
fruit-body must be examined very carefully: an aguttate spore suggests
Clavaria; a multiguttulate spore suggests the subgenus Holocoryne. In some
species, such as C. fuszformis, the gutta may be yellowish and the spore-print
pale yellow. The old spores of C. luteo-tenerrima, C. pulchra, and C. cornicu
lata, lying on the hymenium, often appear yellow or orange, but the fresh
spore-prints are white and remain so for several days at least: it seems that
the old spores on the hymenium become coloured by absorbing the yellow
cytoplasmic colour of the old basidia. In the same way, I think, the old spores
of C. depokensis become reddish. The spore-wall is never coloured as it is in
Ramaria.
The subgenus Acularia has spinous spores: its position is discussed on
page 8,1.
Basidia. Most species have normal, clavate, 4-spored basidia. Many have
2-spored basidia occasionally among the normal 4-spored ones, but it is doubt
ful if any species is entirely 2-spored. C. amoena, C. spiralis, and C. miniata
seem to produce 2-spored, 3-spored, and 4-spored basidia indiscriminately
and often in equal numbers on the same fruit-body, though generally 4-spored.
In some species abnormal basidia with a single, very long, simple or shortly
bifurcate sterigma are formed, bearing. 1 or 2 spores, e.g. Text-Figs. 149,
151, but they are exceptional. As already explained, the genus Clavulin
opsis was based on the abnormal basidia of C. miniata, with 2-3-4 sterig
mata of varying length and arrangement at the apex of the basidium. The
2-spored states need to be carefully distinguished from Clavulina: the species
of Clavulinopsis have distinctly clavate basidia which do not become second
arily septate after discharging the spores, but generally collap!'le, and their
sterigmata are not so distinctly outcurved, though the difference is slight:
the absence of cristate-multifid and flattened branching will also distinguish
any 2-spored, branched species of Clavulinopsis.
The size of the basidium varies, as usual, with the size of the spores and the
basidia tend to be longer in thickened hymenia. However, the species allied
with C. amoena, C. fusiformis, and C. corniculata (spores 5-7 µ, wide, sub
globose) have large basidia 40-65 X 7-10 µ,, whereas those allied with C.
arenicola, C. similis, and C. fruticula have smaller basidia 20-40 X 5-7 µ,, though
their spores are but little smaller. Thus the simple white fruit-bodies of C.
brevipes have the large basidia of the C. amoena-group and the similar fruit
bodies of C. filipes have the small basidia of the C. similis-group. The
distinction may be important, but existing data on these basidia, which often
have fong stalks, are probably inaccurate. The longest basidia, 70-95 µ,,
occur in C. umbrinella.
The cytoplasm of the basidium is generally multiguttulate. The oil seems
to be converted into colloidal solution in passing up the sterigmata, to be
recondensed as the gutta in the spore. In some species the oil appears to be
in colloidal solution also in the basidium, the cytoplasm of which appears
vitreous-opalescent, e.g. C. fruticula.
Cystidia. Only C. luticola has been described with cystidia. None of the
30 species I have studied in the living state has cystidia, and it is improbable
that th1::y occur in any of the well-known north temperate species. Those
of C. luticola may be abnormal basidia, e.g. C. brevipes (Text-Fig. 151):
if so C. luticola comes naturally in the C. amoena-group, instead of occupying
an isolated position.
Hymenium. In the following 17 species the hymenium thickens:
C. alcicornis, C. amoena, C. Archeri, C. arenicola, C. aurantio-cinnabarina, C.
brevipes, C. cineroides, C. corniculata, C. dichotoma, C.Jruticula, C.Jusiformis, C. luteo
alba, C. miniata, C. pulchra, C. spiralis, C. subtilis, C. umbrinella.
In C. luteo-tenerrima it does not thicken; possibly, also, in C. similis, but I

have not seen old specimens for comparison. The remaining species have yet
to be examined for this feature, which is probably very general.
Hyphae. The hyphal structure of C. amoena (p. 79) is typical of the genus:
among the inflating longitudinal hyphae occur numerous narrow interweaving
hyphae. In C. fruticula, C. similis, C. luteo-alba, and C. pulchra there are no
narrow interweaving hyphae: that is to say, all the hyphae of the primordial
shaft inflate. In C. minutula, C. propera, and C. tenuicula the hyphae seem not
to inflate, being 2-4(-5) µ, wide in full-grown fruit-bodies. This is· the
condition, also, in C. hastula (with gelatinous hyphal walls), C. constans, and
C. pogonati (which may be an Aphelaria). It is yet impossible to say to what
extent the hyphal details are useful in classifying the species, but they are
0
important in analysing the size of the fruit.,body. Thus, the small fruit-bodie�
of C. brevipes with greatly inflated hyphae must have much more restricted
apical growth than those of C. propera: similarly C. calocera and C. hastula,
both with gelatinous texture. (The notes of previous authors on clamp-
connexions need verification: cf. C. constans.)
Colour. Excluding Ramaria and Clavariadelphus most of the yellow or
orange Clavaijas belong to Clavulinopsis. Thirty-one species are yellow or
orange; 13 are white, or colourless; 5 are pink or red; 2 are grey; 5 (or 6) are
brownish to umber: and 5 are hysterochroic, discolouring acropetally with
age, probably through slow oxidation of the hyphal walls. Yellow seems to be
the basic coloration, as a diffuse pigmentation in the cytoplasm of the sub
hymenial hyphae and of the old effete basidia, as in C. amoena, C. corniculata,
C. helvola, and C. pulchra: rarely are the medullary hyphae yellow. A few
yellow-orange species occur in the closely related Ramariopsis, and Clavaria
CLAVULINOPSIS 79
argillacea (with yellow flesh and colourless hymenium) is the sole instance in
its genus, excluding the hysterochroic C. straminea.
The red and pink colours seem to be caused by carotinoid guttulae in the
medullary hyphae, especially near to the subhymenium, as in C. miniata and
C. luteo-alba. The intermediate orange and flame colours are caused either
by the presence of both yellow hymenial pigmentation and red or pink medul
lary pigmentation, as in C. amoena, or by another rich orange-brown pigment
in resinous masses in the subhymenial hyphae and old basidia, as in C. pulchra.
Indeed, analysis of the colour-variation in the parallel series C. amoena
C. miniata and C. pulchra-C. Archeri-C. corallino-rosacea will be instructive.
Possibly C. luteo-alba introduces a third parallel similar to C. amoena-C.
miniata, but pure yellow and deep red forms are not known.
Concerning the brown and grey pigments, little is known, but in C. um
brinella a pale-brown colour develops slowly in the cytoplasm of the sub
hymenial and outer medullary hyphae.
Apart from the doubtful C. umbrina, C. Holmskjoldii appears to be the only
metachroic species, having purple tips to the brownish branches. It suggests
the coloration of Clavulina with similar transitions to grey states, as in the
closely allied Clavulinopsis griseola and C. cineroides.
The colourless species are: presumably, achroic derivatives of the coloured
and their affinity must be discovered from microscopic indications: they are
certainly not a natural group. Thus C. subtilis connects with C. corniculata or
C. umbrinella, C. spiralis with C. amoena and C. miniata, and C. similis perhaps
with C. alcicornis.
Smell. C. Holms�;oldii is said to smell of aniseed: C. luteo-ochracea is said
to be fetid: C. citrino-alba is said to have an oily smell: C. corniculata smells
of meal. The other species appear to be inodorous.
DEVELOPMENT OF THE FRUIT-BODY OF CLAVULINOPSIS AMOENA

The acute apex of the actively growing young fruit-body is composed of a
tapering fascicle of narrow hyphae 1 · 5-2· 5 fL wide. Some of these hyphae,
in their proximal parts, become the wide longitudinal hyphae of the
mature flesh; others do not inflate, or hardly at all, and remain as the
narrow interweaving hyphae. On the surface, the hyphal ends which lag
behind the apex turn out to form the first basidia and then by profuse sym
podial branching, which begins as soon as the diageotropic curvature of the
hyphal ends, the interwoven subhymenium is constructed. In the mature
fruit-body the subhymenial hyphae are connected with both the inflated and
the uninflated hyphae, so that the difference in inflation does not indicate a
sterile and a fertile system of hyphae. When a basidium has been formed
from the apical cell, a narrow cylindric blunt process, 1·5-2 fL wide, grows
from the subterminal cell and becomes the next basidium: it is cut off by a
septum, with clamp, at a short distance from its origin and then this new apical
cell enlarges into a basidium: it does not become clavate until it has nearly
reached its full length. By repetition of the process, the hymenium is gradually
extended outward, the old basidia being overtopped by the excrescent young
basid�I processes. Sometimes the new process cuts off a subterminal cell
before becoming a basidium and thus increases the outgrowth and adds to the
irregularity of the hymenium. The processes arise from various depths in
the subhymenium, occasionally from the outer longitudinal hyphae of the
flesh, and- they grow out to various lengths, and hence the variable length of
the basidia: generally, the basidia are longer in the old hymenia which have
thickened considerably. There are no paraphyses or true cystidia, but a
basidium sometimes becomes clavate without producing sterigmata and it
remains, embedded in the old hymenium, as a gloeocystidium-like cell with
oily-vacuolate contents. The old basidia collapse and never become secon
darily septate as in Clavulina. The hymenium is not stratified.
On the stem, in place of the hymenium, there is a sterile, finely villous
layer composed of very narrow, distinctly thick-walled hyphae 1-1·5 µ, wide,
simple or sparingly branched, and interwoven but directed more or less
perpendicularly from the surface. These fine hyphae, up to 300 µ, long,
resemble the mycelial hyphae. They arise mostly from the narrow hyphae of
the flesh by profuse branching in the region corresponding with the initial
subhymenium. The sterile villous layer passes gradually into the hymenium
by the development of basidia, fertile and sterile, among the narrow hyphae.
It thus appears as a surface-tissue .inteqnediate between the secondary
mycelium at the base of the stem and the hymenium.
CLASSIFICATION OF THE SPECIES
Apart from the subgenus Acularia, the species differ so little in spore
charactets, the common species are so variable in shape, size, and colour of
the fruit-body, and there are still so many which are practically unknown
microscopically, that their classification is extremely difficult. Important
characters appear to be the shape of the spore (ellipsoid or globose), it"s size
(3-5 µ, or 5-8 µ, long, or wide) and the colour of the fruit-body. On these
princip,Jes I have arranged the species in the Key, the subdivisions ofwhich
seem to, be natural except for the white species. The shape and size of the
fruit-body has little significance, as already mentioned, though the branched
fruit-body is clearly the primitive state and the small simple fruit-body the
derivative. It follows, therefore, that until the branched species are adequately
known there can be no proper understanding of the genus. At present
C. corniculata (yellow), C. alcicornis (pink), C. cineroides (grey), C. umbrina
(with C. umbrjnella), and C. propera (brown, uninflated hyphae) suggest
five series for which simple eoloured and both branched and simple colour
less derivatives may be expected: possibly the hysterochroic C. semivestita
introduces a sixth series. The C. spiralis-C. amoena-C. miniata alliance shows,
however, that the colour may vary through yellow to orange and red, or
to pink or white, without variation in any other respect. C. luteo-alba sug
gests that the spores may vary much in size (5-15 X 3-6 µ,) without material
change in shape or colour of the fruit-body. There is no guiding rule for the
CLAVULINOPSIS 81
genus. But when there is more accurate knowledge of the spores and their
contents (when living), of the size of the basidia, of the thickening hymenium,
of the inflation of the hyphae, and of the pigmentation, then I think the inter
relations of the species will become clear.
A detail which may be important is the prominent apiculus on the spores of
C. corniculata, C. fusiformis, C. pulchra, C. Archeri, and C. luteo-tenerrima.
It seems to separate them from the other yellow-orange species and its
occurrence in the brown C. umbrinella and the white C. dichotoma may indi
cate their affinity.
ANOMALOUS SPECIES
Subgen. Acularia. At first I was inclined to place these species in Ramari
opsis and to regard them merely as having more str.oAgly echinulate spores.
The distinction between Clavulinopsis and Ramariopsis would then have
been easier to state but, as Donk pointed out to me, the uniformity of Ramari
opsis would have suffered. It is a homogeneous genus of branched species
with very shortly echinulate spores, R. asperulospora being the one simple
species. The affinity of Acularia, as indicated by the yellow C. helvola
( = C. dissipabilis), is clearly with C. fusiformis-C. amoena, and the variable
number and length of the rather stout spines in both C. helvola and C. astero
spora do not suggest the same kind of spore-ornamentation as occurs in
Ramariopsis. The angular echinulate spores of Scytinopogon show, also,
how careful one must be in assuming affinity from verbal descriptions,
because this genus is allied with Thelephora rather than with Clavulinopsis
Ramariopsis.
C. luteo-ochracea, C. spathuliformis. Both are said to have rough spores,
not echinulate. The first may belong in Ramariopsis, but C. spathuliformis
is, at present, impossible to classify. It may be noted that, in Clavaria, C.
occidentalis was supposed to have rough spores, as P. sasae is in Physalacria.
Roughness of the spore is a particularly difficult character to describe, figure,
and assess.
C. pogonati. The large spores, the flattened branching, and the narrow
hyphae suggest Aphelaria very strongly, but Coker describes the hyphae as
clamped. Perhaps it is a 'Tremellodendron' like the anomalous Aphelaria
tuberosa and A. pusio.
C. calocera, C. constans, C. hastula, C. vernalis. For C. hastula I had intended
a new genus based on the phycophilous habit, the gelatinous texture, the
narrow aguttate spores, and the 4-6-spored basidia. It is certainty far removed
in these respects from C. fusiformis and C. corniculata. However, the other
three species, with which I group it in the Key, destroy very largely this
generic character. C. calocera appears to be a true yellow Clavulinopsis with
cylindric spores and toughly gelatinous flesh: it may even be phycophilous.
C. vernalis seems to be an aberrant Clavulinopsis with pale-yellowish colour,
tough, but not gelatinous flesh, and narrow, elongate, aguttate spores: it is
phycophilous. C. constans seems a colourless non-phycophilous state of C.
Sl 19 G
vernalis. If C. vernalis and C. hastula are separated because of their ellipsoid
aguttate spores and narrow hyphae, then one must consider the branched C.
propera and Lentaria subgen. Lentariopsis (p. 94). Possibly C. hastula
should belong to the Clavariadelphus-Typhula-Pistillaria series. I leave it,
therefore, in Clavulinopsis until new generic criteria can be established or
new species can be found to link it with others.
C. Miehe/ii. This is possibly a yellow variety of Clavaria vermicularis:
its spores are much too small for the Clavaria argillacea series.
C. minutula, C. tenuicula, C. propera. The hyphae of the fruit-bodies of
these species are not inflated, but they are clamped, unlike Aphelaria, and
the branching of the fruit-bodies does not seem to be flattened.
C. pusilla. The position of this species is wholly problematical. If really
Clavulinopsis, it will be unique in its habitat.
AFFINITY OF CLAVULINOPSIS
Clavulinopsis and Ramariopsis form a main group of Clavarioid fungi with
white, typically guttate, spores. It is paralleled by Ramaria with branched fruit
bodies and brown or ochraceous spores and by Clavariadelphus-Typhula
Pistillaria with simple, often epiphytic, fruit-bodies and white aguttate spores.
Presumably Clavaria belongs to the series, as derivative with peculiar hyphae,
but no intermediate is yet known. Donk separates Clavulina widely from
Clavulinopsis, placing it near Craterellus and Cantharellus because of the
stichic basidia. The Clavulina-basidium is certainly different from that of
Clavulinopsis but I do not think it has the importance which Donk gives to it:
I must leave the point, however, until the form, size, and construction of the
basidium can be discussed in greater detail. There is a curious resemblance in
colour and form between the fruit-bodies of Clavulinopsis umbrina (4-spored)
and Clavulina Leveillei (2-spored) which, together with their subglobose
1-guttate spores, suggests real affinity though the flattened multifid branching
of Clavulina does not occur in Clavulinopsis.
Leiltaria has the same hyphal construction as Clavulinopsis but the hyphae
tend to be thick-walled. Its habitat is lignicolous and its spores narrowly
elliptic and aguttate. Its affinity seems more with Ramaria subgen. Stricta or
with the Clavariadelphus-series than with Clavulinopsis.
DENDROCLADIUM Lloyd
Myc. Notes 'oo, 1919, 870, and 66, 1922, IIII, f. 2068: Stevenson, J. A., and Cash,
E. K., Bull. Lloyd Libr. Mus. 35, 1936: Patouillard, Journ. de Bot. 3, 1889, 23, 33,
and Ess. Taxon. 1900: Killermann, Nat. Pfl. Fam. 2nd ed., 6, 1928, 150.
Lloyd intended this generic name to cover tropical species of Pterula s. lat.
and Lachnocladium s. lat. which had coloured spores, whether they were
smooth or rough. He reserved Lachnocladium for species with colourless
spores (or with spores unknown, on the assumption that coloured spores
could always be found on herbarium-material) and, thus, Dendrocladium
DENDROCLADIUM
became Lloyd's working name for Lachnocladium sect. Coniocladium Pat.
( spores smooth) and sect. DendrocladiumPat. ( spores verrucose or echinulate).
'We adopt a name proposed by Patouillard as a section.' (Lloyd.) Both sec
tions I consider to be only tropical species of Ramaria which have found their
way into Lachnocladium because of the apparent toughness of the fruit-body
when dried and studied in European herbaria. Thus Dendrocladium is a
synonym of Ramaria and, if separable as a genus for the Echinospora-group
of Ramaria, it is antedated by Phaeoclavulina Brinkm. (1897). ·
Lloyd also referred to Dendrocladium the dubious species Phaeopterula
hirsuta P. Henn. and P. juruensis P. Henn., with neither of which he seems to
have been personally acquainted. Thus, he followed P. Hennings in regarding
their spores as brown, which v. Hoehne! has since shown to be an error. Had
Lloyd known this, he would have referred them to Pterula or Lachno
cladium, not to Dendrocladium. Accordingly, one cannot agree with Steven
son and Cash that D. hirsuta (P. Henn.) Lloyd should be the lecto-type and
that Dendrocladium is a synonym of Pterula. (I am indebted to M. A. Donk
for explaining to me this misinterpretation.)
Lloyd made eight combinations under Dendrocladium but there is no
evidence that he regarded al}y of them as a type-species, particularly as he
would put in Dendrocladium 'most of the species included in Patouillard's
synopsis as Lachnocladium'. The eight species belong as follows:
(i) as Thelephora Fr. ( = Phylacteria Pat.}, D. Archeri (B.) Lloyd and D. congestum
(B.) Lloyd;
(ii) as Pterula Fr., D. hirsutum (P. Henn.) Lloyd, D.juruense (P. Henn.) Lloyd and,
perhaps, D. fructicolum (B.) Lloyd (which may be a Pyrenomycete);
(iii) as Ramaria S. F. Gray emend. Donk., D. guadelupense (Lev.) Lloyd;
(iv) as Lachnocladium Lev. s. str., D. furcellatum (Fr.) Lloyd;
(v) as Clavariachaete gen. nov., D. Peckoltii Lloyd.
Killermann interpreted the genus on D. Peckoltii which would have been •
satisfactory had Lloyd not whimsically suggested that it indicated yet another
genus 'Clavariachaeta McGinty', thereby excluding as typical of Dendro
cladium the one species which needed a new genus.
I conclude that Dendrocladium was based, taxonomically, on Lachno
cladium sect. Dendrocladium Pat. and is, therefore, a synonym of Ramaria,
the reference to Pterula or Phaeopterula being an error.
LACHNOCLADIUM Lev.
Taxonomy. In a note concerning Clavaria tubulosa and C. furcellata, in
1830, Fries remarked that tropical Clavarias seemed to differ from the
European in the velutinate hymenium (when dried), and that, when they were
better known, this character might be generic. In 1846, Leveille took up the
idea and made the genus Eriocladus for the more or less coriaceous, tropical
species of Clavaria with minutely villous or subtomentose branches. He
listed the following seven species and, in a supplement, added E. setulosus (B.):
E. acicularis Lev., E. brasiliensis Lev., E. compressus (B.) Lev., E. funalis
Lev., E.furcellatus (Fr.) Lev., E. scoparius Lev., and E. tubulosus (Fr.) Lev.
(E. fistulosus by error).
In 1849 Leveille substituted the name Lachnocladium for Eriocladus which
he regard_ed as a later homonym of Eriocladium Lindl. In 1889 Patouillard
narrowed the genus to species with brown spores, compressed axils, and
longitudinally sulcate branches (when dried), but most authors have followed
Leveille's definition to the present time. The distinguishing features of
texture and surface are, however, so vague, the first being of very uncertain
use with dried specimens (on which nearly all tropical species have been
based) and the second graded in all degrees to the glabrous state, that the
genus has come to be the repository for all apparently tough tropical Clavarias.
The genus is, in fact, one of the great 'uncombed assemblages' of early pocket
lens systematy, which must be resolved by the choice of a type-species and the
use of the microscope. 1
The type must be one of the original seven species of Eriocladus and,
preferably, L. tubulosum or L. furcellatum, as the two Friesian species which
provided the idea of the genus. Neither has been satisfactorily identified.
L. tubulosum, as understood by Patouillard (18�9, p. 33), is a Ramaria of the
group Echinospora and, if selected as the type because of its earlier publica
tion (1828), it would make Ramaria a synonym of Lachnocladium. L. furcel
latum (1830) has, however, been regarded for many years as a common
tropical fungus, and accordingly it has been chosen as the type by Clements
and Shear (1931) and Donk (1941). L. furcellatum was based on collections
from Guinea, Bourbon, and Brazil, aµd the type-specimens are presumably
in the Friesian herbarium. There is none at Kew, but most of the many
specimens referred there to this species have a very characteristic structure
which also occurs in L. brasiliense (one of the seven, of which there is a frag
ment of the type at Kew) and, probably, in the eighth species L. setulosum.
This structure is
monomitic, uninflating hyphae without clamps, with pale yellowish
brown walls (darkening in dilute alkali), and with abundant gloeocystidia
and dichophyses.
This hyphal construction distinguishes these species from all other
'Clavarias' and relates them to the resupinate genus Asterostromella in the
Xanthochroic-series of Basidiomycetes (the Hymenochaetoideae and the
Asterostromel \oideae). Together with the clavarioid form it is a generic state
to which the name Lachnocladium may satisfactorily be applied. When so
defined there appear to be some twenty species of the genus and nearly a
hundred to be excluded and referred variously to other genera of branched
Clavarioid fungi, even to Thelephora and Tremellodendron. Only three of
1
If Clavulina cristata, Ramariopsis Kunzei, or Ramaria strict; had been described from
dried specimens, they would have been put in Lachnocladium. The villous or subtomentose
appearance of the branches of Lachnocladium is caused by the sterile hymenium on their
uppersides.
LACHNOCLADIUM 85
Leveille's eight species of Eriocladus belong to Lachnocladium, n;imely, E.
brasiliensis, E. furcellatus, and E. setulosus: of the other five, E. acicularis,
E. funalis, and E. scoparius are probably Aphelaria dendroides, E. compressus
probably Lentaria surculus, and E. tubulosus is most problematic (though
possibly a Ramaria).
Concerning a lecto-type, however, I am yet doubtful. My studies of the
Malayari species of Lachnocladium s. str. have shown that their differences
are mainly anatomical and microscopic. In the Kew herbarium, moreover,
I do not find any American or African species which are conspecific with the
Asiatic, unless perhaps L. divaricatum Pat. s. lat. L. brasiliense resembles
the Asiatic species macroscopically but appears to differ markedly in its wide
medullary hyphae, and under L. furcellatum there is a mixture of several
American, African, and Asiatic species. I doubt, therefore, whether the
original three collections of L. furtellatum are conspecific, and one cannot
decide until the living species of America and Africa have been adequately
described and the Friesian specimens have been examined microscopically.
On the other hand, L. brasiliense is based on a single collection from Brazil and
appears to be well-defined microscopically, though this point must. be
checked from more ample, modern collections, so that it seems less proble
matic and, therefore, a better type-species. Singer has also suggested that it
should be the type (Mycologia 1944, 553), but I prefer to wait until the
American species are better known and to take the xanthochroic construction
with dichophyses as the generic distinction.
Of the two sections, or subgenera, of Lachnocladium proposed by Patouil
lard, namely, Dendrocladium (asperate, verrucose, or echinulate spores) and
Coniocladium (smooth spores), the first must be referred to Ramaria but the
second, though a mixtum compositum (with species of Pterula and Clavulina),
must be retained for 'Eu-Lachnocladium' if the genus is subdivided, for
Patouillard placed in it both L. furcellatum and L. brasiliensf!.
Dichophyses. These curious organs, found also in the resupinate Astero
stromella and in some tropical spycies of Stereum and Hymenochaete; may
be likened to dichotomizing setae in a diffuse hymenium. They arise either
on lateral hyphae of limited growth, or terminally. The lateral hypha grows
for some 20-100 µ,, when it is cut off by a septum near its origin and its apex
enlarges slightly into the central part of the dichophysis and then produces
two to five bulges, generally at a wide angle to the original direction of growth.
These bulges are the beginnings of the main arms of the dichophysis (Text
Fig. 31). Each grows outward and soon dichotomizes, and its divisions grow
on apically to dichotomize in their turn. As the process is repeated, the arms
become more numerous and narrower and the intervals between the dicho
tomies lengthen: they end in very fine, acute tips o· 5-1 µ, wide. 1 Meantime,
1
There is clearly a mathematical spacing-formula to be derived for the dichophysis, just as
for the Clavarioid fruit-body, which will relate the width of the apex of the arm to the fre
quency of division or the length of the internode. Measurements of the parts of symmetrical
or regular dichophyses would provide interesting data, but the organs are usually too involved
for convenient study.
in the centre of the dichophysis, or in the distal part of its stalk, the hyphal
wall has begun to thicken and turn brown: the thickening proceeds acropetally
along the arms but the tips generally remain colourless. Careful study of the
ToxT-FIG. 3 I. Lachnocladium zonatum: full-grown and developing dichophyses, X 500.
very young stages shows that even the initial bulges are to be regarded as the
first, second, or third dichotomies of a rapid succession. The arms of the
dichophyses never elongate into ordinary hyphae but are always of limited
'die-away' growth: very occasionally they are trichotomous.
LACHNOCLADIUM
Structure of the Fruit-body. A medulla of longitudinal uninflated hyphae
traverses the stem and branches to end at each growing-tip in a fasc1cle of
if;;,
]' �t)r:; Gh W\ u :a-·"
e
, �·
t
-
.
�
�v•,11::, ··,·.��. ' }
\i� ·
�� r �
�twj -�u6
---·
TEXT-FIG. 32. Lachnocladium zonatum: dichophyses from the medulla (top right), from the
hymenium (bottom row), the remaining three from the cortex, X 500.
hyphae. Scattered in the medulla, mostly towards the outside, are a few
dichophyses which are commonly misshapen and sparingly branched with
elongate and mainly longitudinal arms (Text-Fi,g. 32). The medulla is sur
rounded by a more or less tomentose or spongy, but firm, cortex composed,
at first sight, entirely of dichophyses which decrease in size from the medulla
to the..surface until the outermost may be only 20 µ, wide (overall) and appear
as minute masses of coralloid divisions: actually the whole tissue is pervaded by
.,,._>t':'/.,'f:,Y �·
f¥·1Y
0t.�·
�· y-,""'P-"l-:!i'1--
fi!�J�?'�- ·�;i••.
- -,�·�:�v.#.t�
.n�-�.(t.�:�� C],'Af.!(�·
��- · · · · .;vli%,�}...'-
��·,}�if�,.:,,_.(iR1 ��.
��,�j}iitisi��.:
�;_-1:'�0(f��-
··�t?\f.{�1'i��1ft8!
\,.. fl�llt,•/
TEXT-FIG. 33. Lachnocladium: diagrams of the structure of the base of the stem (below, in
t.s. and Ls., x 10) and of a fertile branch (above, in t.s., X 50): L. divaricatum var. cinnamo
meum (left): L. molle (top right) with wide medulla: L. divaricatuni (bottom right) with very
narrow medulla.
fine, colourless, generative hyphae concealed among the densely felted arms
of the dichophyses which they produce. Thus, a transverse section of a
branch always shows a pale medulla in a browner, looser cortex and generally
the medulla is relatively slender, though in L. molle it is wi.der than the cortex.
In most, if not all, species the cortex thickens gradually throughout the
growth of the fruit-body, and thus it is wider in the stem and lower parts of
the branches than in their distal parts.
LACHNOCLADIUM 89
Apical Growth of the Branches. The growing-tips are more or less filiform,
as in Pterula, 40-500 µ, wide, and consist of a fascicle of entwining, col0urless
hyphae 2-3 µ, wide. The apical growth of these hyphae builds up the medulla,
TEXT-FIG. 34. Lachnocladium divaricatum

var. cinnamomeum: longitudinal section of a
branch-tip showing the origin of medullary
and cortical dichophyses, X 500.
,. which undergoes no subsequent elongation or thickening because the cells do

not inflate. At distances of o· 5-2 mm. from the tip the medulla becomes
corticated very gradually by the development of dichophyses from ex
crescent lateral hyphae and from the outcurved hyphal ends on the sides of
the fascicle. Other lateral hyphae continue to grow out into this incipient
cortex to produce more dichophyses and, as they project beyond the general
level, l!O they extend the cortex which continues to thicken by this superficial
outgrowth. As already mentioned, the dichophyses become smaller the far
ther they are situated from the medulla (but they never form undivided, or
®GO 0
TEXT-FIG. 35. Lachnocladium zonatum: the single-layered hymenium with excrescent

dichophyses, x r ,ooo.
simply asteroid, setae as in Asterostroma and Asterodon). The cortex
develops equal!y round the vertically erect stem or main branches but, on the
inclined branches, it develops more on the under than the upper side, so that
the branch becomes elliptic in section. The upperside remains sterile (without
gloeocystidia), and the hymenium develops on the underside. The hyphae
which form the cortical dichophyses are always directed more or less perpen
dicularly from the medullary hyphae and are never increscent.
Hymenium. After a certain specific period of outgrowth, the gen�rative
hyphae in the cortex on the undersides of the branches produce gloeocystidia
LACHNOCLADIUM 91
and basidia. They are at first scattered but soon become compacted into a
thin waxy layer beyond the tips of the coralloid dichophyses which are in
vested by the plexus of subhymenial hyphae. In some species, as L. fiavidum
and L. fulvum, this appears to be the mature state, for even the oldest branches
of their largest fruit-bodies have but a single layer of basidia. In others, as
L. divaricatum and L. molle, the hymenium thickens continually by gradual
excrescence of the subhymenial branches until a waxy layer, 100-200 µ, thick,
is formed: it consists of sympodial rows of effete basidia terminating in the
active basidia at the surface and it is traversed by the continually elongating
gloeocystidia at right angles to the medulla (Text-Fig. 36): during the process
new gloeocystidia may be added at various stages but no dichophyses, which
are restricted at the inmost part of the subhymenium. In L. zonale there is
a third complication in that the simple hymenium becomes overgrown by
another layer of cortex with simple hymenium and this overgrown in its turn
until the tissue appears stratified (Text-Fig. 191). Its generative hyphae grow
through the simple layer of effete basidia to produce a new set of dichophyses
and then again a limiting and simple layer of basidia, to be superseded by a
third and even a fourth layer: so the cortex becomes stratified by layers
indicating periods of active outgrowth terminating in fertile palisades.
In all cases the gloeocystidia are the first hymenial elements to be formed
and are thus deeply rooted in the subhymenium, and, in the species with
continuously thickened hymenium, they generally become more or less
attenuated during the excrescence and appear as subcylindric bodies with
subventricose bases.
The spores are produced abundantly, unlike so many tropical Pterulas, but
they are extremely minute and offer little value in specific determination.
They generally turn pale yellowish on prolonged drying. It seems that the
hymenium may not develop, or not become fertile, until the fruit-body is
several centimetres high.
Branching. The fruit-body is built up by the successive dichotomies of the
filiform tips of the branches, though it may not be obvious in mature fruit
bodies, for the outgrowing cortex webs the branches together at their bases.
During the greater part of their growth the ends of the branches are terete
and taper to a more or less filiform point. At certain times preparatory to
branching, but for no evident reason, the apices flatten and broaden into
spathulate knife-edges, 0·5-1·5 mm. wide, on the two sides of which the arms
of the new dichotomy arise, leaving the central part as an arrested webbing
between them. The new arms appear to be regions of residual apical growth
after that of the central part has been arrested, rather than regions of new
outgrowth, and thus they diverge slightly on origin and later curve upward
and become parallel through geotropic or phototropic direction. The new
branches are first distinguishable as slightly thickened and more opaque
regions along the lateral margins of the spathulate apex, where the hyphae
have begun to proliferate on all sides to regain the terete form. At first the
spathulate tips and new arms consist only of longitudinal medullary hyphae,
TEXT-FIG. 36. Lachnocladium divaricatum var. cinnamomeum: longitudinal
section of the thickened hymenium (simplified), showing the extended
gloeocystidia, the absence of dichophyses, and the small coralloid
subhymenial dichophyses, X 1,000.
LACHNOCLADIUM 93
but they soon become corticate with dichophyses and the separate cortices of
the new arms unite across the axil to web them together. In the case of the
early division of the primordial shaft and main branches in the young fruit
body, 2-4 dichotomies in the same or slightly different planes may follow in
such close succession that, on becoming webbed together by the cortex, there
is the false appearance of 3-8, or more, palmate branches as if produced by
the simultaneous polychotomy of a greatly dilated apex: in transverse section,
however; the true positions of the dichotomies can be seen from the positions
of the medullary strands to each branch, for they may be separated by as
much as 1-2 mm. in the flattened 'palm' of these spurious polychotomies.
Later divisions of the thinner branchlets are well-spaced dichotomies. Thus
the lower branches of the fruit-bodies are commonly described as polychoto
mous or multifid, and the distal branches as dichotomous or bifid. Slender
or depauperate fruit-bodies may be simply dichotomous throughout, e.g.
L. Hisingeri. Actually the full details of the manner of branching have yet
to be made out from the study of young fruit-bodies.
It will be seen that according to the state of the branch-tips, whether about
to divide or having just divided, so they are obtuse and spathulate, or acute
and subulate. In regular fruit-bodies developed without crowding the
branches generally divide r'nore or less simultaneously and then the fruit
bodies may appear very different in form, being Thelephoroid when the
branches are spathulate and Pteruloid when subulate. The differences have
been mistaken as specific.
Texture. Because of the lack of inflation and because of the thick cortex
of thick-walled dichophyses with very fine interlacing arms, the texture is
rather drily coriaceous with a more sappy medulla; dried specimens are light
and rather hardly coriaceous and often somewhat brittle.
Colour. The colour of the fruit-body resides mainly in the walls of the
dichophyses. The oleaginous contents of the gloeocystidia may, however,
be yellowish or pale olivaceous. The growing uncorticated tips of the
branches are white, and just behind the tips, where the dichophy$eS begin to
form, the colour is generally bright ochraceous or mustard-yellow.
Mycelium. The dichophyses occur abundantly in the general mycelium
and they corticate the slender mycelial strands or rhizomorphs which charac
teristically attach the fruit-bodies. Sometimes the mycelium at the base of the
fruit-body expands into a thin felt or subiculum which consists mainly of
dichophyses.
initiation of the Fruit-body. I have not been able to find th� earliest stages
of development. It appears that the mycelial strands first originate in the
substratum and anastomose at the surface and that from this plexus the
primordial shafts arise to form the individual fruit-bodies, which are typically
caespitose.
LENTARIA gen. nov.

This new genus is intended for two groups of Clavaria s. lat. which do not fit
into the microscopic characterization of other genera. I have called the groups
subgen. Eu-Lentaria and subgen. Lentariopsis. The Stricta-group of Ramaria
seems to be their nearest connexion, particularly for Eu-Lentaria which has
the same kind of thick-walled hyphae, tough, radially branched, yellowish
ochraceous fruit,.bodies, lignicolous habit, and abundant development of the
secondary mycelium. But Lentaria has smooth, thin-walled spores which, in
most cases, are white and which tend to be very narrowly ellipsoid or elongate,
whereas the spores of the Stricta-group are rough (or nearly smooth), rather
shortly ellipsoid, and with yellow, ochraceous, or nearly ferruginous, thickened
walls. The exceptions in Lentaria are L. byssiseda and L. Micheneri, the
spore-prints of which are said by different authors to be white, cream, pale
ochraceous, or even deep ochraceous (L. Micheneri sensu Martin). They
seem to bridge the gap between Lentaria and Ramaria. On the other hand it
is very difficult to draw a line between the Stricta-group of Ramaria and the
rest of that genus, for there are several intermediate species. One must, there
fore, either conceive Ramaria as covering R. botrytis, R. Zippelii, and L. sur
culus (smooth white spores), which seems an unnaturally wide range, or, as I
prefer, separate Lentaria. I have a feeling, also, that some of the Stricta
group of Ramaria have been mistaken for species of Lentaria. Moreover,
6-spored basidia, as occur in L. surculus, do not occur in Ramaria. Doubtless
the two genera have a common Clavarioid ancestor from which Lentaria and
the Stricta-group are separate branches.
The species of Eu-Lentaria are so much alike that it is possible there are
really but two cosmopolitan species differing in the size of spore (10-18 X 3-6 µ.
for L. surculus: 7-1 o X 2 · 5-4 µ. for L. Micheneri).
The position of the subgenus Lentariopsis is uncertain. L. affiata, L.
delicata, L. epichnoa, and probably L. virgata are very closely related and truly
lignico!ous on coniferous wood. Unfortunately they seem very rare and are
but little known. They appear to be depauperate or reduced derivatives of
Eu-Lentaria with small narrow spores. Their hyphae have only slightly
thickened walls, at least in the branches, though they may be thicker in the
main stem, as in Eu-Lentaria. The lignicolous habitat prevents them from
being placed in Clavulinopsis, as does the absence of large guttae in the spores:
their hyphal structure also prevents them from being retained in Clavaria
s. str.
L. mucida and L. coronilla appear as phycophilous derivatives, yet further
reduced with fruit-bodies which are generally simple. Nevertheless, if their
relation with wood is secondary and dependent on the lignicolous habitat of
the algae with which they grow, they may belong to the affinity of Clavuli
nopsis vernalis and C. hastula, both of which are phycophilous, but terrestrial.
Certainly it is remarkable that L. coronilla (6-8-spored) and C. hastula (4-6-
spored) have supernumerary sterigmata, but they occur also in L. surculus
LENTARIA 95
(2-6-spored). Such reduced featureless fungi are extremely difficult to
classify.
L. corticola is problematic, but its habitat suggests this genus.
MUCRONELLA Fr.
This genus is usually regarded as Hydnoid, but several species are certainly
Pteruloid, e.g. M. fasicularis, M. pacifica, and M. ulmi, which I have trans
ferred to Deflexula.
The Hydnoid view, which began with Fries, regards the fruit-bodies as
Hydnoid spines borne directly on the rudimentary mycelium which does not
form a stroma or subiculum. Doubtless Fries was influenced by the analogy
of Solenia, the caespitose fruit-bodies of which he regarded as the tubes of an
astromatous Polypore. Structurally the European species M. aggregata and
M. calva, and the tropical 1\1. brasiliensis and M. pusilla, seem to have fruit
bodies identical with Hydnoid spines but this, in itself, is not proof. Their
structure is the same as in the truly Clavarioid fruit-bodies of Ceratellopsis,
which differ in growing upward instead of downward as those of Mucronella.
The Clavarioid genera Deflexula and Hormomitaria show that not even the
direction of growth can distinguish Hydnoid from Clavarioid fungi. What is
lacking is clear evidence that Hydnoid fungi degenerate in this manner. In
most of their resupinate genera, as Odontia, Grandinia, Radulum, and Acia,
the spines degenerate, and even disappear, while the stroma or resupinate
Corticioid state persists. A Hydnoid fungus can degenerate into a Corti
cioid, but it has not been shown that it can become a minute spine, as a
juvenescent primordial shaft. Nevertheless, in favour of the Hydnoid theory,
Bourdot and Galzin note that Mucronella calva sensu Pat. is the young state
of Odontia arguta which, they imply, may develop spines first and stroma
second (the opposite of the usual procedure for resupinate 'Hydna'). The
observation introduces a curious and, as yet, morphologically inexplicable
feature of some resupinate fungi of very diverse origin: normally res·Jpinate
fungi grow by marginal growth, and spines or pores are formed in centrifugal
order just behind the margin and before the advancing hymenium: in some
Porias, as P. sanguinolenta, many minute primordia arise to coalesce and,
ultimately, to give the appearance of a single fruit-body of one origin; the same
occurs in some species of Sebacina, Tulasnella, and Corticium, e.g. C. aegeri
toides. If, then, this proliferation of many intermediate primordia occurred
in extremis in a resupinate Hydnum, the condition of Mucrcnella would be
obtained.
Hennings and Lloyd regarded Mucronella as Clavarioid, but their evidence
is not clear and they certainly confused Clavarioid with Hydnoid fungi. For
myself, I regard a spine of such a species as M. aggregata as equivalent not to
a Hydnoid spine but to the primordium of a Hydnoid fruit-body, in the same
way as a single fruit-body of a minute and gregarious Pterula, Pistillaria, or
Ceratellopsis is the equivalent of the primordium of any Clavarioid fungus.
TAXONOMIC AND MORPHOLOGICAL N,OTES
Thus, I hold the single fruit-body of M. pusilla (Text-Fig. 200) equivalent to
that of Ceratellopsis aculeata (Text-Fig. 65), but whether it has come by way
of Clavarioid or Hydnoid ancestry I cannot decide. If Hydnoid, Mucronella
and Ceratellopsis provide a most extraordinary instance.of convergent evolu
tion, as the effect of degeneration along two widely separate lines, resulting
in an end-term distinguishable only by the direction of growth.
DEVELOPMENT OF THE FRUIT-BODY OF Af. PUSILLA
There is a very slight, diffuse mycelium on the surface of the wood. The
fruit-bodies develop from small localized ramifications of two or three mycelial
hyphae, occasionally from a single hypha. The lateral hyphae grow out from
· the surface of the wood and then turn vertically down. They branch freely,
the branches at first rather spreading but soon coming into alinement as the
apical growing-point. The primordium passes through a brief pulvinate
stage before it becomes conical and develops by apical growth into the pri
mordial shaft. The ptilvinate part becomes the base of the fruit-body with
short, almost pseudoparenchymatous, cells. The hymenium soon begins to
form on the primordial shaft even when it is only 70 µ, long. It develops in the
usual way by the out-turning of the hyphal tips on the outside of the primordial
shaft and about 10-30 µ, from the apex. The out-turned hyphal ends cut off
· 2-3 short subhymenial cells before becoming basidia. Apical growth of the
primordial shaft is very limited but the apex never becomes fertile. Spores
begin to mature before apical growth has ceased. The primordial shaft may
be arrested in any stage of development, even the microscopic pulvinate
stage, and then mature basidia over its surface, except for the apex and the
extreme base: in this respect it seems truly Hydnoid.
PHYSALACRIA Pk. AND HORMOMITARIA gen. nov.

The systematic position of Physalacria has been much disputed. From his
study of the type-species, P. infiata, Peck (1882) recognized that it was a
basidiomycete and placed the new genus in the Clavariaceae. Krieger (1923)
saw in this species a primitive agaric and, without taxonomic reason, changed
the generic name to Eogaricus and made the new family Eogaricaceae.
McGuire (1939) decided that it belonged to the Thelephoraceae because of
the sterile upperside of the head, and that it was not an agaric because the
folds on the fertile underside of the head were not gills, as Krieger thought,
but simple buc,klings of the hymenium. Most recently Baker ( I 941) has re
affirmed its position in the Clavariaceae because of the irregularity in position
of the sterile areas on the head. None of these reasons will stand inquiry:
there are many Clavariaceae with sterile areas on the upperside of the branches
and there are degenerate agarics without gills. All such views, based on gross
appearance, are artificial and do not penetrate the real nature of the fungus.
As defined by Peck and subsequent mycologists, Physalacria is indistin
guishable frorp Pistillaria with globose hollow head ( = Sphaerula Pat. and
PHYSALACRIA AND HORMOMITARIA 97
Pistillina auctt., e.g. Pistillaria capitata). The resemblance is heightened by
the indirect development of the fruit-body in both cases (cf. Text-Figs. 39,
42, 201) and by recent discoveries of very small epiphyllous species of the
genus in tropical America, though the almost equally small P. Langloisii
(1888), P. andina (1893), P. bambusae (1909), and P. villosa (1917) had been
described much earlier and misunderstood. Further, in Pistillaria there is
the same slight agglutination of the hyphae as in Physalacria. But, in contrast,
Physalacria has three, if not four, peculiarities absent from Pistillaria, thus:
1. oleocystidia;
2. subacerose or subfusiform young basidia (with paraphysis-like transi
tions to oleocystidia or fusiform cystidia);
3. late development of the spores, the fruit-bodies appearing sterile until
full-grown (those of Pistillaria, as in most Clavarioid fungi, maturing
basidia from the earliest inception of the hymenium);
4. partial sterility of the hymenium, the sterile areas being composed
mostly of fusiform cystidia.
Thus, aside from the small, monaxial, and unbranched form of the fruit
body, the definition of Physalacria is microscopic.
The second and third peculiarities are typical of many Marasmioid genera,
which have also indirect gymnocarpic development, spores, basidia, and
caulocystidia, as in Physalacria, together with slight agglutination of the
hyphae in the stem, which may also extend to the flesh of the pileus and render
it waxy-tough or subgelatinous. Indeed the stem-structure of some species of
Marasmius or Skepperia is so like that of Physalacria tropica and P. infiata,
and the pileus-structure of Marasmius Menieri, 1 for instance, so like that of
the head of these Physalacrias, that one is tempted to consider the Physalacria
fruit-body as that of Marasmius reduced to an apileate convex proliferation
at the top of the stem-as an antithesis, perhaps, to stem-less Cyphelloid
derivatives. The sterile hymenium of Physalacria would then appear as a
relic of the palisade of pileocystidia in such Mantsmioid fungi. However,
none of these has the characteristic oleocystidia of Physalacria, nor can one
satisfactorily explain the basal position of the sterile hymenium in the head
of many species: even the lnocybe-like cystidia of Skepperia andina are hardly
comparable.· The Marasmioid affinity, therefore, seems unproven, if possible;
and once fungi are reduced to their primordial state it is easy to be misled by
comparable simplicity.
The genus Hormomitaria, which I describe on page 41 o, brings a new stand
point to the subject. It has in common with Physalacria the peculiar oleo
cystidia, the small white spores, the sterile base to the fertile head, indirect
development of the stem, Marasmioid caulocystidia, and similar hyphae.
Therefore, it is undoubtedly allied with Physalacria. But it differs very
strangely in the shape of the fruit-body which is an inverted, positively. geo
tropic spine with conico-cylindric, solid, fertile head, and it has complicated
1
Tr. Brit. Myc. Soc. 19, 1934, 285.
5119 H
secondary septation of the hyphae. Hormomitaria can be interpreted only
as an inverted Clavarioid fungus such as Deflexula, and the relation of
Physalacria to it is that of an ageotropic juvenescent fruit-body with apical
growth restricted to the formation of the stem and with intercalary hymenial
growth dilating the apex of the stem into the hollow head. Secondary septa
tion has disappeared from Physalacria, but the sterile base of the head is
retained in several species, though in some, as P. infiata, it seems to be con
nected rather with the disappearing geotropism than with the developmental
construction. Physalacria thus appears as a limiting state of Hormomitaria,
as P. capitata is to Pistillaria and D. capitata to Dimorphocystis and Pterula,
and the problem of the affinity of Physalacria is shifted to Hormomitaria, which
is as unlike a Marasmioid fungus as possible for a monopodial basidiomycete.
Some species of Physalacria with shortly conical head, as P. andina, seem to
be intermediate in retaining, if slightly, the prolonged apical growth of Hor
momitaria and in having but slightly developed the intercalary expansion of
Physalacria (cf. Text-Fig. 202).
The apical growth of Hormomi .ria shows that it is a Clavarioid fungus.
The secondary septation suggests a link with Clavaria, but the lignicolous
habit is that of the Clavariadelphus-series. Possibly it is a highly modified
member of the Marasmioid series of Basidiomycetes: in any case the resem
blance between Physalacria and Pistillaria is an instance of convergent evolu
tion through degeneration of the fruit-body. Mucronella tenuipes, of tropical
Africa, may belong to the same series.
PISTILLARIA Fr.
Definition. As explained on page 145, the definition of Pistillaria is bound up
with that of Typhula. The limitation of Typhula to species with sclerotia, as
I have used it, defines Pistillaria negatively as Typhuloid fungi without
sclerotia, and this was Greville's use of the genus in 1828. The genus. im
pingeE..on Clavariadelphus, Clavulinopsis, and, perhaps, Lentaria, from all of
which it differs in the simple hymenium (not thickening) and in the aggluti
nated superficial hyphae of the stem-the character which, I believe, has led to
the evolution of the sclerotium, with its agglutinated cortex, as an abortive
fruit-body deformed through adverse environmental conditions (temperature,
light, and so on). Pistillaria resembles in other respects, particularly the
spores, Clavariadelphus (as C. junceus) and it must represent, in the evolu
tionary sense 1 the 'epiphytic' modifications of Clavariadelphus and the
tendencies towards Typhula. Unfortunately the descriptions of most species
are so imperfect that, in practice, it is impossible to follow such a principle
and the genus must still remain the receptacle for most diminutive, epiphytic,
simple, white-spared Clavarioid fungi. The Friesian view ( 1874), adopted by
Patouillard, Bourdot, and Rea, that the slight distinction between a fertile
head and a sterile stem defines Pistillaria, in contrast· with Typhula, is
untenable because, actually, there are very few species without a distinct stem,
PISTILLARIA 99
at least microscopically, and they seem to belong mostly in Ceratellopsis or
Pterula.
I have listed in the index 114 specific names for Pistillaria. From these I
have removed 2 to Clavaria, 9 to Pterula, 16 to Ceratellopsis, 13 to Typhula,
4 to Pistillina, 1 to Chaetotyphula, 2 as Fungi Imperfecti, and 12 as synonyms
of Pistillaria spp., while 2 belong to the Tremellaceae. This total is 61, and
leaves '53 species in Pistillaria, 24- of which I have transferred to the genus,
mostly from Typhula and Clavaria. Of these 53 species, about 20 are doubtful,
in that they may belong to some other genus, as Pterula or Ceratellopsis, and
a few may be synonyms in Pistillaria itself. There is thus a considerable
remainder of some thirty species which centre round
P. capitata, P. diaphana, P. micans, P. pusilla, P. setipes, and P. uncialis.
These six species have essentially the same construction, which I describe
in detail for P. capitata and P. setipes (pp. 104--7). I regard them as the
nucleus of Pistillaria in the sense in which I use it. None of them has a
sclerotium and three, namely, P. diaphana, P. micans, and P. pusilla belong to
the original species of the genus. Fries (1821) placed in Pistillaria these seven
species:
P. diaphana, P. micans, P. ovata, P. pusilla, P. quisquiliaris, P. sclerotioides, and P.
muscicola.
P. ovata I regard as identical with P. pusilla. P. quisquiliaris and P. sclero
tioides have sclerotia and belong in Typhula emend. Karst., .as I have usect it,
while P. muscicola is Eocronartium (Tremellales). Thus, four of the original
seven species remain in Pistillaria as a natural group and from them I would
choose P. micans or P. pusilla as a lecto-type, if such is required. Actually I
think the group itself satisfactorily defines the genus. I have explained this
point of view because it is in opposition to that of Donk (1933). He has chosen
P. quisquiliaris as the type of Pistillaria and, chiefly for this reason, has used
Cnazonaria Cda. (1829) for the group which I call Pistillaria. The reason for
the choice of P. quisquiliaris (see p. 14-5) is the horny texture of the fruit-body,
especially when dried, but the character is shown as well' by P. micans and
P. puiilla. My chief objection to this definition is that I cannot see how to
separate P. quisquiliaris generically from Typhula phacorrhiza (lecto-type of
Typhula): they have the same sclerotial structure and very similar spores, and
the difference in shape, size, and texture of the fruit-body is bridged by
T. gyrans and T. incarnata, both with the same sclerotial structure. So
Pistillaria or Typhula would have to become a synonym of the other. It is
extremely undesirable to reduce either of these foundation-genera, especially
when both are needed, at least for practical expediency in separating sclero
tium-bearing species from those without sclerotia. Further, Fries never saw
the sclerotium of P. quisquiliaris, for which reason alone, as a species mis
understood by him or imperfectly known, it is unsuitable as a lecto-type. To
what extent, however, the species which I place in Pistillaria must be relegated
to Typhula as sclerotium-free derivatives (and so, secondarily, returned to
Pistillaria, as it were), as Donk's classification indicates, remains to be worked
out when they are known in full microscopical detail (see p. 102). The crux
seems to be whether Typhula subgen. Subcarnosae is identical with Pistillaria,
as I have used it.
TEXT-FIG. 37. Diagrams of the structure of the fruit-body of Pistillaria

uncialis (left), P. capitata (centre), and P. setipes (right), showing· the
transformation from the cylindric to the clavate and subglobose head by
reduction of apical growth.
Classification of the Species. There appear to be two main groups, the

P. micans-setipes group, itself consisting of the pink-purple series of P. micans
and the colourless or white series of P. setipes, and the P. uncialis-group. The
first group is distinguished by the waxy texture when fresh, only the super
ficial hy�hae of the stem having agglutinated or toughly mucilaginous walls:
the species certainly belonging to it are
P. setipes-ser�es: P. albobrunnea, P. Boudieri, P. capitata, P. cytisi, P. diaphana,
P. maculaecola, P. pusilla, P. setipes, P. typhicola.
P. micans-series: P. ampelina, P. bellunensis, P. elegantula, ? P. fulgida, P. granu
lata, P. lividula, P. micans, P. purpurea, P. rhodocionides, ? P. syringae, P. uliginosa.
The P. setipes-series may be subdivided into a small section with globose

heads, representing the last term in reduction, or limitation, of the hymenium
to the apex of the primordial shaft immediately on cessation of upgrowth, and
consisting of P. Boudieri and P. capitata: and a main section with clavate,
PISTILLARIA IOI
ovoid, or ellipsoid heads in the formation of which apical growth of the

primordial shaft continues for some time after the initiation of the hymenium.
The P. uncialis-group is distinguished by the waxy-tough or rather horny
texture of the fruit-body, nearly all the medullary hyphae of which have more
or less toughly mucilaginous, subagglutinated, walls. Typically, also, the
fruit-b_odies are yellowish or pale ochraceous. It comprises:
P. oleae, P. subuncialis, P. Todei, P. typhuloides, P. uncialis, and, perhaps, 'P.
epiphylla and P. lignicola.
TEXT-FIG. 38. Pistil/aria capitata: part of a hypha from the head, X 500.
Now, all of these four subdivisions have a generic name, but I do not
regard them as generically distinguishable for there appear to be intermediate
states. Thus:
Cnazonaria Cda. (1829), for the P. setipes-series, with P. setipes as type
species.
Sphaerula Pat. (1883), for the P. capitata-section, with P. capitata as the
type-species.
Scleromitra Cda. (1829), for the P. micans-series, with P. micans as the
type-species.
Gliocoryne Maire (1909), for the P. uncialis-group, with P. uncialis as
the type-species.
Each main group shows a considerable range in spore-size and in the
reduction from the cylindric to the clavate or ovoid head, and, ultimately, in
the P. setipes-series to the capitate head. Sessile derivatives have not been
found, as occur in Pterula and Ceratellopis. The P. uncialis-group has the
longest heads, presumably with least reduced apical growth, and it is note
worthy that this group shows in size, colour, and texture of the fruit-body the
nearest approach in Pistillaria to Typhula phacorrhiza: even the large spores of
P. subuncial£s closely resemble those of T. phacorrhiza. Further, Donk says
that P. Todei is very near to T. quisquiliaris and that P. uncialis sensu Maire
(non Grev.) is the sclerotium-free state of T. sclerotioides. The resemblance of
the species to Typhula and the possibility of their having sclerotia, under
some circumstances, requires to be looked into with greater care. On the other
hand, Pistillina, especially P. hyalina, has the same texture as the P. uncialis
group but is certainly without sclerotium. This genus seems to stand to the
P. uncialis-group very much as Sphaerula to the P. setipes-series: unfortu
nately the species are very rare, very minute, and extremely little known. It is
certainly remarkable, and in need of explanation, that in both Typhula and
Pistillaria there appears to be a comparable division into a group with waxy,
non-agglutinated fruit-bodies (Typhula subgen. Subcarnosae, P. micans
setipes group) and a group with waxy-tough, more xerophytic fruit-bodies
with more or less agglutinated medullary hyphae (Typhula subgen. Phacor
rhizae, P. uncialis-group and Pistillina).
Among the species which I have placed for convenience or through
ignorance in Pistillaria, there is a group with yellowish, erumpent, corticolous,
often deformed fruit-bodies recalling those of Clavariadelphus fistulosus var.
contortus: they are P. alnico/a, P. fuegiana, P. paradoxa, and P. spathulata.
They may belong, rather, to Clavariadelphus, but compare also the curious
tropical assemblage of Lachnocladium aurantiacum.
Distribution. Most species belong to the north temperate region and, as
they appear mostly in late autumn and early winter, they seem to be low
temperature fungi, as do the species of Typhula. Unlike this genus, however,
a few genuine species occur in the tropics, as P. subpellucida (Cuba), P.
rhodocionides (Malaya), and another white species near P. pusilla which I have
found ;n Singapore (but without spores). Nevertheless the genus is certainly
not common in the lowland tropics, but much is to be expected from the
mountains and the south temperate regions.
Structure and Specific Details. The structure of the fruit-body is essentially
as in Typhula and the specific variations need similar elucidation. Thus the
shape of the head may depend merely on intercalary extension of the hymen
ium, on inflation of the subhymenial cells, or on the presence of a wide cortex
of divergent hyphae between the longitudinally medullary hyphae and the
subhymenium (cf. P. setipes, Text-Fig. 41, and T. gyrans, Text-Fig. 47).
The presence or absence of oxalate crystals may have specific value, but the
records are, as yet, too desultory for use.
Conidia are said to occur sometimes on the fruit-body of P. micans.
Clamp-connexions seem generally to be present in the fruit-body, but they
are absent from P. rhodocionides and from some fruit-bodies, at least, of
P. uncialis.
TEXT-Frc. 39. Pistil/aria setipes: two early stages of development before the appearance of the
hymenium: a diminutive fruit-body with the basidia beginning to form acropetally (right) and
the fertile head of another (left): t.s. of a mature stem with agglutinated superficial hyphae
(centre), / 500.
"STRUCTURE AND DEVELOPMENT OF PISTILLARIA SETIPES
At certain points in the leaf the mycelial hyphae branch profusely and
become concrescent to form a mass of hyphae 20-50 µ, wide. On the side of
this primordium next the epidermis, the hyphal tips, 2-3 µ, wide, come into
alinement as the incipient primordial shaft and, as they grow out, they rupture
the epidermis and project at right angles from the leaf. These minute white
points are the first external signs of the fruit-body which is initiated within
Trurr-FfG. 40. Pistillaria setipes: initial stage in the emergence of the fruit-body from the tissue
of a dead leaf (palisade-cells with black contents): iron-alum haematoxylin preparation,
X 1,000.
the leaf and without sclerotium. The primordial shaft soon thickens by
branching of its own hyphae and by upgrowth of new hyphae from the
mycelium in the leaf and round the base of the shaft. The enlargement soon
ceases and the shaft grows out as a cylinder of hyphae which, to judge from
the curvature of the stems of mature fruit-bodies, is positively phototropic.
The apical cells of the hyphae are long and flexuous and intertwine, sometimes
regularly to g�ve all the hyphae of the fruit-body a 'spiral twist'. The sub
terminal cells branch from their distal ends or from the middle, at first one
branch from each cell and then a second branch from some of the cells.
As the primordial shaft grows apically the cells at the base begin to inflate
to form the cushion-like, turgid support and before the head is formed they
may have become thick-walled and coloured. The base also thickens for some
time through branching from the cells, particularly at the surface, and by
upgrowth of short hyphae from the mycelium (as in Pterula gracilis). The
PISTILLARIA 105
caulocystidia develop from below upwards as outgrowths, from the superficial

cells, mainly from their distal ends but also from the middle or proximal part.
They are eventually cut off by a septum, but towards the apex of the shaft they
remain as rudimentary protuberances.
When the primordial shaft has reached 1 -4 mm. long the head begins to
develop. It is initiated by protrusions to the exterior from the tips of the
superficial hyphae on the outside of the growing-point. The protrusions look
like young caulocystidia, but they become clavate, not attenuate, and they are
developed from every hyphal tip so that the distal end of the primordial shaft
becomes closely covered by them, though at first not contiguously (Text-Fig.
39), and the apical hyphae continue to grow longitudinally for a short space.
The protrusions become cut off by a septum when about 10 µ, long. They may
enlarge directly into basidia or cut off one or two short cells c. 5-8 µ, long before
the end-cell becomes a basidium. Thus is built up acropetally the first hy
menium, which is loose because enlargement of the longitudinal hyphae in the
head separates the young basidia at the surface. Into the spaces between them
g1ow branches from the subterminal cells and these branches also cut off one
or two cells before the end-cell becomes a basidium. By such intercalary
growth the continuous hymenium is formed and the subhymenium is con
structed from the short subt�rminal, branching, and interlocking cells. Even
tually the hyphal tips in the growing-point become arrested: they septate
down to cells c. 10 µ, long and the end-cell becomes a basidium. The growing
point may be transformed into the hymenium soon after the inception of the
head, which is then short, or not until it has grown for an appreciable distance,
in which case the head is elongate-clavate: the apex never remains sterile.
The head increases to its mature size by intercalary growth of the hymenium
and by inflation of the longitudinal hyphae: at first the internal inflation
exceeds the intercalary growth, but soon the intercalary growth becomes
excessive and the longitudinal hyphae are pulled somewhat apart, though the
head never becomes hollow as in P. capitata. Laterals from the outer longi
tudinal hyphae may also grow into the hymenium and produce basidi:i. But
the elongation of the longitudinal cells near the apex is always less than in
those near the base of the head, so the head is clavate and blunt. The incep
tion of the head is sudden and, though the hymenium has slight basipetal
extension through the development of basidia from the surface-hyphae at the
ape'{ of the stem (Text-Fig. 41 ), the head is generally sharply delimited from
the slender stem. In some cases, however, particularly in small fruit-bodies
2-3 mm. high, the head appears to taper gradually into the stem, but I think
the appearance is caused by the shortness of the stem compared with the head,
rather than by any less definite formation of the hymenium.
The inflation of the cells of the longitudinal hyphae begins as soon as they
are cut off from the apical cell, and continues until the hymenium has deve
loped to the apex of the head. The elongation of the stem, however, has
generally ceased by this time and further growth in length is by enlargement of
the head.
Variation in the size of the fruit-body is mainly in length of the stem and is
caused by the·varying extent of apical growth. It may be controlled photically
so that fruit-bodies developed on the upperside of a leaf have shorter stems
than those from the underside. Variation in width of the fruit-body is slight
;:J+.
TEXT-FIG. 41. Pistillaria setipes: longitudinal sections of a diminutive head on cessation of

upgrowth and d the basal part of a normally elongate head: iron-alum haematoxylin
preparation, X 1,000.
and is doubtless determined by the number of hyphae in the growing-point,

which will depend on the food-supply in the mycelium: a poor food-supply
will give attenuate fruit-bodies. Variation in length of the head is more
complicated and will depend, mainly, on the extent of apical growth after
inception of the head or, conversely, the rate at which the hymenial factor
extends over the apex.
PISTILLARIA 107
The cells of the fruit-body are binucleate. The fusion-nucleus of the

basidium evidently has more than two divisions because more than 4 nuclei
can be seen in the full-grown basidia. The spores are uninucleate, as found
also by Kuhner (Le Bot. 17, 1926, 30). Sometimes a branch arises from the
base of a basidium before a septum has formed; the basidium then has two
pairs of nuclei, one in the upper part and one at the base, and the basal pair
will move into the branch: but it seems that in such cases no clamp is formed.
The caulocystidia are binucleate, but the reduced caulocystidia at the apex of
the stem have no nuclei. It appears that the apical growth of hyphae and the
inception of branches are not immediately dependent on the nuclei, which are
never situated at the apex of the hypha and which do not enter the branch
until it has reached a considerable length (cf. Noble's observations on Typhula
Trifolii).
STRUCTURE AND DEVELOPMENT OF PISTILLARIA CAP/TATA
The general structure and development of the fruit-body is the same as in
P. setipes. There are slight differences. The fruit-bodies are superficial, not
erumpent, but whether they arise from single hyphae (as in Pterulagracilis) or
from a tuft of hyphae as in P. setipes I could not determine for lack of the
earliest stages. The primordial shaft reaches o· 5-2 mm. long, when the head
is formed by the transformation of the apical cells into young basidia
simultaneously over the whole apex of the shaft. The apical cells become
subclavate and cut off 1-3 short subhymenial cells before they mature
into basidia. The initial hymenium is small and narrow, at the tip of the
primordial shaft, but it enlarges greatly by intercalary growth to form the
subglobose head, the underside of which becomes sharply recurved to form
the characteristic 'umbilicus' in which the stem is inserted. At the same time
there is slight inflation of the longitudinal hyphae of the head, but for the most
part they are pulled apart and stretched to form the air-hollow in the head.
The stem continues to elongate by cell-inflation after the head is formed and
may even double its length. The caulocystidia arise as in P. setipes and
diminish upward in the same way: at the stem-apex there are a few scattered
sterile basidia, as transitions to the abruptly delimited hymenium. A few
secondary mycelial hyphae grow from the base of the stem into the superficial
mycelium.
The cells of the fruit-body are binucleate. The fusion nucleus divides with
its spindle inclined at a slight angle to the long axis of the basidium, and so do
the two daughter nuclei. A third division must occur, though I did not see it,
for full-grown basidia have 6-8 nuclei. One nucleus enters each spore, which
remains uninucleate, and the others become clumped together near the middle
of the basidium.
PISTIL LINA Quel.

This genus has been considered doubtful. I have been fortunate in finding
the type-species and can give, therefore, a fairly adequate description. The
QOo
PISTILLINA 109
genus is near to Pistillaria from which it differs mainly in the extreme limita
tion of the hymenium to the apex of the primordial shaft, with so little
intercalary growth that the outer hyphae of the apex remain as a narrow
margin round the discoid or pulvinate hymenium. Thus the fruit-body
resembles the apothecium of Helotium or Phialea with turgidly convex
hymenium. P. hyalina also differs in its remarkably long basidia, but those
of P. Thaxteri seem to be much shorter. In both the hyphal mechanism is
degenerating: the hyphae of P. hyalina inflate but slightly and those of P.
Thaxteri, apparently, not at all. The superficial hyphae are characteristically
agglutinated, as in Typhula and Pistillaria, and the central hyphae have
submucilaginous walls. Thus, the genus seems to be a limiting term to the
P. uncialis-series of Pistillaria, very much as P. capitata is to the P. setipes
series; possibly the agglutination of the hyphae in the P. uncialis-series
prevents their separation to form a spherical head in Pistillina, as can occur
in Pistillaria capitata in the P. setipes-series which is without agglutination in
the head. Thus, I have excluded Pistillaria capitata ( = Pistillina Patouillardii
Quel.) because it lacks the peculiarities of Pistillina and its resemblance is
homo-plastic, not homologous.
DEVELOPMENT OF THn FRUIT-BODY OF PJSTILLINA HYALINA

Development is essentially the same as in Pistillaria setipes (p. 104), but
there is little inflation of the hyphae, their cells evidently being of considerable
length on delimitation, and the hymenium is limited to the apex of the blunt
primordial shaft. The hymenium dilates the apex of the primordial shaft by
intercalary growth, and thus widens it into the solid cup-shaped head: many
lateral hyphae grow up from the internal tissue, thus preventing it from
becoming hollow, and they terminate with clusters of basidia in the hymenium.
There is no extension of the hymenium on to the sides of the primordial shaft,
the outer apical hyphae of which seem to bound the hymenium from its origin.
According to the amount of its intercalary growth, so the hymenium is discoid,
convex, or almost hemispherical, and the head is narrowly to widely dilated,
but the outside is never reflexed or recurved as in many other capitate fungi
(e.g. Mitrula, Leotia, or Physalacria).
PTERULOID SERIES
General Structure of the Fruit-body. The fruit-body is typically Clavarioid
with apical growth and superficial development of the hymecium from the
out-turned hyphal tips below the growing apex. Development is direct and
gymnocarpic without inflation of the hyphae.
Inversion of the Fruit-body. In Pterula and Pterulicium the fruit-bodies
are erect and, presumably, negatively geotropic as well as positively photo
tropic. In Deflexula the fruit-body is positively geotropic and, perhaps,
positively phototropic, though the adventitious branches seem ageotropic at
first. In one species of Dimorphocystis the fruit-body is erect, like that of a
simple Pterula, but in D. subcapitatus and D. capitatus it seems both ageotropic
and aphototropic. Pterulicium is peculiar in its resupinate patch which
develops ageotropically and aphototropically, but is not fertile unless facing
obliquely or vertically down. The inversion of the fruit-body is a rare pheno
menon among basidiomycetes. It occurs in three other Clavarioid genera,
namely, Physalacria, Hormomitaria, and Mucronella, and elsewhere, perhaps,
only in degenerate Cyphellaceae and some other Thelephoraceae. It is a
problem for experiment, for which the Pteruloid fruit-bodies with only
apical growth and no secondary means of orientation are well fitted.
Size of the Fruit-body. The greatest height recorded for Pterula is 12 cm.
for the fruit-bodies of P. Bresadoleana. P. brunneola, in Malaya, has fruit
bodies reaching 9 X 12 cm. The smallest fruit-body of Pterula, excepting
species of uncertain position, is that of P. nana, 1-1·5 mm. high (as a pseudo
Pistillaria): in Dimorphocystis the fruit-bodies reach 18 mm. high in D. laevis
and only o·6 mm. in D. capitatus. This range of 12-0·06 cm. (200: 1) is
entirely a difference of apical growth, without cell-inflation, and is perhaps
as big a difference as occurs in any two related genera of homologous plants.
In bulk, there seems as great variation but it is more difficult to estimate
because little is known of primordia or their rate of attenuation. The primordia
of Pterula brunneola have a conical growing-point about 0·5 mm. wide at its
base, and it must consist of some 50,000 hyphal tips. P. verticillata has almost
as large primordia and the tips of its branches attenuate to a single leading
hypha. In such Pistillarioid species as P. epiphylla, the growing-point of the
primordium consists of 50-100 hyphal tips. The fruit-body of P. gracilis
begins as a single hypha (Text-Fig. 43).
The variation in size, considered as numbers of hyphae and extent of apical
growth, is thus as great in the Pteruloid-series as in any natural series of
Basidiomycetes.
Branching of the Fruit-body. Branching of the large fruit-bodies of Pterula
is primarily flattened-dichotomous in successively alternating planes. Secon
darily, the main branches may appear polychotomous through condensation
of 2-3 dichotomies with very short intervals. I have not seen true polychotomy,
and the apices of the branches become only narrowly spathulate before
dividing. The branches of the dichotomy may develop equally to give a
multaxial fruit-body, as in the large Pterula brunneola, or one branch may
overtop the other to give an apparently monaxial and monopodial fruit-body.
P. verticillata has such a monopodial axis with false whorls of 3-4 branches:
its main bran�hes repeat the same form and, normally, only the ultimate
branchlets are equally dichotomous (Text-Fig. 235). The effect is produced
by regular overtopping in two or three dichotomies which follow in rapid
sequence in alternating planes so as to maintain an axis.
From the photographs given by Lloyd, this form of fruit-body appears
common in the genus. I have not found such regular and alga-like somata in
any other Basidiomycetes. Different ways of branching may occur in other
species of Pterula, but there is no exact information about this feature, though
PTERULOID SERIES III
it is one of the more important, for on it depends very largely the specific shape
and appearance of the fruit-body.
The species with small short fruit-bodies and microscopically attenuate
apices, as P. epiphylla, P. gracilis, Deflexula, and Dimorphocystis have no
apical branching. Others, with less reduced fruit-bodies, such as P. typhuloides
and Pterulicium, have a few dichotomies often with overtopping of one branch,
and they connect the simple Pistillarioid forms with the bushy ones.
The loss of branching seems connected directly with the attenuation of the
primordia, for the reduced fruit-bodies are exactly like the ultimate branches
of the bushy Pterulas. It seems as if the relation between the number of
hyphae composing the growing-point and the signal for branching has re
mained constant throughout the Pteruloid-series so that unless this number is
reached the hyphal tips remain concrescent in a single fascicle: the number
seems to be of the order of 10,000. In other words, if the species with simple
fruit-bodies could be induced to form stouter primordia, they would develop
larger, ramified fruit-bodies. Actually, when better fed by growing on a
richer or more extensive substratum, they produce not so much larger
primordia as more numerous ones about the same size. Loss of branching
in these fruit-bodies seems, therefore, to be a secondary phenomenon ac
companying reduction in size of the primordia, which is to be explained by
diminution of the value of the factor causing the. original concrescence of
mycelial branches to form the fruit-body: this is true degeneration, not
depau peration.
Data on the size of the primordia and of the branching tips, and on the
number of hyphae composing them, are required to elucidate this problem.
Adventitious branching. In Pterula di<bilis, Pterulicium, and Deflexula
(probably in many other small fruit-bodies, too) the branching is adventitious,
There is no flattening of the apex of the axis, which merely becomes attenuate
as it grows farther from the origin, but new fascicles of hyphae arise at various
points on the axis, breaking through the incipient hymenium, and growing out
into adventitious branches. In Pterula and Pterulicium, these branche$ arise
mostly near the growing apex of the parent branch or axis, for adventitious
branching may be repeated to the third and, even, fourth degree. In Deflexula
the adventitious branches are generally restricted to the base of the first spine
(or primordium) and so produce the tufts of decurved spines, but in D.
fascicularis branching to the second degree occurs from almost any part of the
axis. Even the minute Pterula gracilis may have traces of adventitious branch
ing (Text-Fig. 43). Clearly the phenomenon is homologous with the cristation
of the branches in Clavulina or of the stem, as in Clavulina ornatipes and
species of Aphelaria: it represents excrescent hyphae escaped from the control
of the main axis. Sometimes, in Pterulicium, the main axis itself expands and
becomes cristate with adventitio�s branchlets.
Hyphae. The dimitic state is curiously arid characteristically imperfect.
The skeletal hyphae are mostly unbranched and of unlimited growth, so
that they are easily recognized, but in almost every fruit-body a few can be
found with somewhat dilated tips from which 1-5 skeletal or generative
hyphae arise as terminal lobes (Text-Figs. 13, 169, 246). The generative
hyphae, also, may assume the form of skeletals for a short time and then revert
to the generative state, so that a few elongate, thick-walled, intercalary segments
always occur in the generative system. Detailed though it appears, the
imperfection is normal and also significant: it may be upgrade, as the beginning
of true dimity, or downgrade as its degeneration: it shows how one form of
hypha can change into the other. The only variations that I have seen are in
Pterula gracilis, which has no apical lobing of its distinct skeletals, and in
Dimorphocystis, which has skeletals of limited growth and which, in the most
reduced D. capitatus, produce conical cystidia from their terminal lobes. In
some species the skeletal hyphae are locally inflated to 15 µ wide: whether
this represents abortive inflation of the apex before lobing or an ancestral state
with fully inflated skeletals cannot yet be decided.
Clamp-connexions. The Pteruloid-series seems, primarily, to be possessed of
clamps and to have lost them in some species. Thus there are these exceptions:
Pterula brunneola: 4-spored, occasional septa without clamps.
P. vinacea: 4-spored, occasional septa without clamps.
P. typhuloides: 4-spored, without clamps.
var. minor: 4-spored, clamps at some septa.
P. subtyphuloides: 4-spored, clamps at some septa.
(P. gracilis: 2-spored, dikaryotic, with clamps.)
· Dimorphocystis laevis: 2-spored, no clamps (? haploid).
D. subcapitatus: 2-spored, no clamps (? haploid).
(D. capitatus: 4-spored, with clamps.)
Branching from the clamp occurs normally in Pterulicium, often in

Pterula debilis and P. epiphylla, occasionally in P. verticillata, P. brunneola,
P. vinacea, and Deflexulafascicularis, but it can hardly be regarded as a normal
feature of the series.
Sterile Hymenium and Caulocystidia. The Pistillarioid species have usually
distiru:tive caulocystidia or a distinctive sterile covering on the stem, the origin
of which must be traced to the sterile hymenium on the stem and the upper
sides of the main branches of the bushy Pterulas: this seemingly unimportant
tissue therefore requires careful analysis. In Pterula verticillata, P. brunneola,
and P. vinacea (Text-Fig. 239) the sterile hymenium consists of subcylindric
or subfusiform caulocystidia, loose sterile basidial branches, and increscent
generative hyphae which may become irregularly lobed at their tips. It seems
that the hymeriial factor, in its inception, was imperfect and had affected, in the
stem-region where it was developing, the superficial hyphae sufficiently to
cause the beginning of the hymenium but insufficiently for its completion:
thus the hymenial hyphae are started and abandoned, and aimless laterals
from adjacent generative hyphae in the medulla grow into the loose sub
mucilaginous cortex. The tissue is full of interest, for its very imperfection
reveals both the action of the hymenial factor and the futility of the hyphae
when left to themselves.
PTERULOID SERIES IIJ
Pterulicium has a similar sterile hymenium without caulocystidia. In De

flexula it is more reduced and limited to scattered sterile basidia.
In Pterula typhuloides, P. subtyphuloides, P. debilis, P. tenerrima, and others
the sterile hymenium is modified into a dark-walled 'pavement-epithelium'
like the surface of a Typhula-sclerotium (Text-Figs. 231, 232). In P.
subtyphulo£des the superficial generative hyphae on the stem become short
celled and develop excrescent lobes, some of which grow into caulocystidia
but most of which intertwine, without proper direction, to form. a compact
tissue 15-30 µ thick: the outer walls of the superficial hyphae then thicken
and become dark brown and agglutinated. In P. typhuloides, P. debilis, and
others there is little or no excrescence, save in the formation of the caulo
cystidia, and the 'pavement-epithelium' has no distinct 'subhymenium'. This
structure may be useful for the subgeneric, if not the generic, distinction of
these species.
In Pterula epiphylla, Dimorphocystis laevis, and D. subcapitatus, the stem
bears only scattered conical caulocystidia. In P. gracilis the caulocystidia are
reduced to slight projections. In D. capitatus the stem is truly glabrous.
In its reduction the sterile hymenium passes from a loose palisade or pile
with caulocystidia, through a state with scattered caulocystidia or sterile
basidia, to that without trace of lobing from the surface hyphae: macroscopi
cally, the stem changes from ·villous or subtomentose to puberulous then
glabrous. Now, assuming the presence of caulocystidia, 1 it is easy to under
stand this sequence. The caulocystidia are formed directly from the out
turned hyphal tips, as the first hymenial structures: the sterile basidia are
formed mostly from the subsequent sympodial branchings of these tips. The
degeneration of the sterile hymenium now becomes a natural sequence in
the diminishing effect of the hymenial factor in its incipient phase. In P.
verticillata its effect is almost normal. In P. subtyphuloides and P. typhuloides
it is weaker and modified by the phenomenon of agglutination at the surface:
there is little sympodial branching after the out-turning of the tips to form
caulocystidia. In P. epiphylla, D. laevis, and D. subcapitatus its effect ie- still
weaker and causes only the primary excrescence into caulocystidia. In P.
gracilis its effect is barely observable. In D. capitatus there is no visible effect,
and one can assume that the factor no longer operates in the short initial
phase of the primordium.
Hymenium. In the bushy Pterulas and some of the Pistillarioid species as
P. subtyphuloides, in Pterulicium and in Deflexula, the hymenium thickens by
ordinary sympodial outgrowth. In most species with simple fruit-bodies the
hymenium does not thicken, e.g. P. gracilis, P. epiphylla, P. Gordius, P.
typhuloides, and Dimorphocystis. In P. gracilis it is so short-lived that it
becomes effete at the base of the fruit-body before apical growth is complete.
In P. typhuloides var. minor the hymenium may be absent from parts of the
fruit-body and, exceptionally, its fruit-body is wholly sterile. Such failures
also show the degeneration of the hymenial factor in the series.
1
See the paragraph on Cystidia, p. 114.
5119 l
u4 TAXONOMIC AND MORPHOLOGICAL NOTES
Cystidia. Several species with Pistillarioid fruit-bodies have characteristic
cystidia, e.g. P. gracilis, P. epiphylla, and Dimorphocystis: others have none,
e.g. P. typhuloi.des, Pterulicium, and Deflexula. Whether they occur in any
large bushy Pterula is not known. I found none in P. verticillata, P. brunneola,
and P. vinacea, but they occur feebly developed in P. debilis. The cystidia
are tyP.ically conical or subfusiform, like the caulocystidia but often larger and
more ventricose, and they are likewise formed directly from the out-turned
hyphal tips in the incipient hymenium. The peculiar lobed cystidia of
Dimorphocystis, which are formed from the later hymenial branches, I have
discussed under that genus (p. 117). Until it is known whether any bushy
Pterulas have cystidia it is impossible to say whether the Pistillarioid species
have inherited them or evolved them independently by introducing into the
hymenium the sterilizing factor which evokes the caulocystidia. The study
of cystidia may be important in tracing the lines of evolution in the series and
it may throw light on the origin of cystidia in general. 1
Sterility. In several collections of the bushy species, as Pterula verticillata
and P. brunneola, which I made, every fruit-body was entirely sterile, though
the hymenium was normally developed and thickened. Similarly I have
found completely sterile fruit-bodies of P. debilis, P. typhuloides, and Deflexula
fascicularis: occasionally those of P. typhuloides var. minor fail to develop any
hymenium. In the case of P. debilis, P. typhuloides, and D.fascicularis, however,
these fruit-bodies, sterile in the day-time, gave spore-prints overnight. The
observations suggest that sparing in some species of the Pteruloid-series is
nocturnal, but more experiments are needed for satisfaction.
Colour. The fruit-bodies are mainly white or lightly tinged yellowish,
pinkish, lilaceous, vinaceous, or brownish. The yellowish and brownish
tinges are probably caused by the colour of the walls of the skeletal hyphae,
and the basidia may also have slightly tinted cytoplasm. There are certainly
no conspicuous pigments. The brown colour of the stems in P. typhuloides,
P. subtyphuloi.des, and others is caused by the agglutinated brown walls of the
supe�cial hyphae.
Habitat. The mycelium of the bushy Pterulas grows in humus or on rotting
trunks, which it often permeates with slender rhizomorphs. That of Dimor
phocystis and P. gracili's, in contrast, with minute fruit-bodies, is limited to
single leaves, even parts of leaves, and, in comparison, is depauperate. The
larger Pterulas are found only in the extensive humus and debris of old or
primeval forest, never in the habitats of secondary or ephemeral nature where
the Pistillaridd species occur.
Spores. There is doubt concerning the colour of the spores. They have
been .considered white in P. subulata and P. multifida, but Konrad and
Maublanc have recently described those of P. multifida as slightly brownish
in the mass. Those of P. verticillata, P. brunneola, P. vinacea, P. gracilis,
1
The fascicle of hyphal tips composing the growing apices have been called paraphyses or
cystidia in misunderstanding of their nature: on cessation of growth their terminal cells may
become cystidia.
PTERULOID SERIES 115
P: epi,phylla, P. debilis, P. typhuloides, and P. subtyphuloides I found to be

white. P. hirsuta was said by Hennings to have brownish spores, and on this
character he separated the genus Phaeopterula. He also described Phaeopterula
juruensis with brown spores, but v. Hoehnel examined the type-specimen
and found them to be white. Pterula timon'ensis (placed by Killermann in
Phaeopterula as a subgenus) was said to have spores with a slightly ochraceous
centre. Hence I consider Phaeopterula a doubtful genus of uncertain descrip
tion, because it is not known whether the colour, if it is present in the living
spores, is in the wall or the cytoplasm.
The small white aguttate ellipsoid spores are remarkably similar in the
genus Pterula, the species with 2-spored basidia having more elongate spores.
Dimorphocystis agrees with Pterula in this respect, but Pterulicium and
Deflexula have much larger spores, those of Deflexula being either globose or
broadly ellipsoid. In all, they are aguttate.
Evolution and Affinity. In the main the fruit-body has progressively degen
erated from the branched Clavarioid state to the simple Pistillarioid. The
mycelium, too, has degenerated and become adapted to the secondary, limited,
or ephemeral habitats. The scene of evolution has probably been the tropical
rain forests, whence comparatively few species have migrated to the temperate
regions. No parasitic species have been found.
The fruit-body has degenerated through juvenescence and progressive
attenuation of the primordium with consequent loss of branching. There has
also been loss of thickening of the hymenium and of such details as clamp
connexions and, perhaps, cystidia. As special developments there are the
resupinate patch of Pterulicium, which may have led through loss of the
Clavarioid fruit-body to Corticioid derivatives, and the adventitious branching
of Deflexula with inversion of the fruit-body to give Hydnoid forms.
Degeneration has proceeded, perhaps, on several different lines. There may
be cystidiate lines giving P. debilis, P. epiphylla, and P. gracilis, acystidiate
lines giving P. typhuloides, P. subtyphuloides, Pterulicium, and Deflexula, and
a dimorphously cystidiate line giving Dimorphocystis. So few spedeS are
sufficiently known that these points cannot be decided and it is therefore
impossible to frame natural genera or subgenera for most Pistillarioid species,
which must be placed provisionally in Pterula.
There are no other Clavarioid fungi comparable with the dimitic Pteruloid
series, allies of which may be found among the dimitic Thelephoraceae and
Polyporaceae, but it is possible that the dimity of Pterula has had an indepen
dent origin. Corticioid and Hydnoid derivatives of Pterulicium and Deflexula
must be expected among the resupinate Thelephoraceae and degenerate
Hydnaceae.
DEFLEXULA gen. nov.

Superficially this genus appears Hydnoid and to resemble, particularly,
Mucronella, in which the three species D.fascicularis, D.pacifica, and D.ulmi
have been placed. Mucronella differs entirely, however, in its small spores,
narrow basidia, monomitic hyphae, and simple spines not proliferating from
the base. The affinity of Deflexula is undoubtedly with Pterula and Pteru
licium. Their fruit-bodies have identical hyphal construction, but those of
Pterula and Pterulicium are erect, with apical branching, whereas those of
Deflexula are downcurved, apparently positively geotropic, with wholly
adventitious branching. This manner of proliferating new spines from the
base of the initial one, or primordial shaft, and thereby forming a tuft of simple
Clavarioid fruit-bodies, seems unique and is very different from either the
caespitose habit of true Clavarioid fungi, in which each fruit-body is a separate
primordium, or the formation of spines in the Hydnaceae, which follows
marginal growth of the receptacle, the spines being positive homologues of
Polyporaceous tubes. The cristation of the stem in Pterula verticillata, Clavu
lina ornatipes, and Aphelaria spiculosa is analogous, if not directly comparable.
Between Pterulicium and Deflexula there seems to be a close connexion,
though at first sight they appear far apart. Both have large, slightly thick
walled spores, large basidia, and no caulocystidia or properly developed sterile
hymenium on the stem. The spores of P. pacifica seem identical with those of
P. xylogenum, if not so amygdaliform. The primary distinction lies in the
direction o(growth of the Clavarioid fruit-body,_yet both genera show the same
deviation from the condition in Pterula in that they have become positively
geotropic. In Deflexula, the Clavarioid state has been affected so as to develop
into decurved spines but, in Pterulicium, this state is unaffected and a new
Corticioid state expresses the positive geotropism. The branching of Pteru
licium is either apical and flattened, as in the bushy Pterulas, or adventitious
as in Deflexula, but subapical rather than basal. The adventitious branches
of Deflexula resemble the Corticioid patch of Pterulicium, however, in. being
at first ageotropic or diageotropic, especially in D.fascicularis (Text-Fig. 162).
The genus is mainly tropical, D. ulmi being the most northerly representa
tive, in the United States, and D. pacifica in the Bonin Islands. It is remark
able, in showing how little is yet known of tropical fungi, that the nearest, and
very "Close, ally of D. ulmi is D. lilaceo-brum:iea from Malaya and Sumatra.
The nearest ally ·of D. pacifica is the tropical Australasian D.fascicularis, with
which D. nivea from Guadeloupe is closely allied. Possibly Pterula tahitensis,
P. Vanderystii, and P. Winkleriana belong to this genus.
The five species form two natural groups as given in the Specific Key.
There is a group with short lilaceous or brownish fruit-bodies, reaching 3 mm.
long, comprising D. ulmi and D. lilaceo-brunnea, and a group with much larger
white or pale>ochraceous fruit-bodies reaching 1·5-2·5 cm. long, which com
prises the other three. Globose and ellipsoid spores occur in both groups, thus
providing an instance where spore-shape is a less useful guide to affinity than
the general characters of the fruit-body.
DIMORPHOCYSTIS gen. nov.
Macroscopically the species of this genus would pass for Typhula, Pistillaria,
Ceratellopsis, or Pistillina in their broad sense. Microscopically, the dimitic
....
DIMORPHOCYSTIS 117
uninflated hyphae and the dimorphic cystidia distinguish them. Their true
affinity is shown by Pterula epiphylla with conical cystidia of one kind, which
connects the genus with Pterula. From the Pistillarioid Pterulas, Dimorpho
cystis seems to have evolved by further limitation of apical growth of the axis,
by limitation of apical growth of the skeletal hyphae, and by development of
lobed cystidia in the hymenium. The first and second points are not difficult
to appreciate, and there is a gradation from D. laevis to D. capitatus, without
which it would have been difficult to classify D. capitatus: indeed, very few
such perfect gradations can be known. Thus:
Height of fruit- Apical growth Length of

Species body in mm. of fertile part skeletal hyphae
D. laevis -r8 prolonged 700-2,000 µ.
D. subcapitatus -r(-4)• 'little or none 200-800 µ.
D. capitatus -0·6 none 300-500 µ.
•- The greater height caused by elongation of the sterile stem.
The branching of the skeletal tips in D. subcapitatus and D. capitatus is also

understandable because, on appearance of the factor causing tne development
of the hymenium, it extends over the whole apex of the primordium and thus
provokes the skeletal tips, as well as the generative, to branch: in D. capitatus,
with most abruptly limited apex, the skeletal tips even partake in the regular
formation of the hymenium by developing some, if not most, of the conical
cystidia and by producing generative hyphae ending in lobed cystidia and
basidia.
The lobed cystidia, however, appear as a new hymenial element. There
may be bushy Pterulas with lobed cystidia, for which reason I have made
D. laevis the type of Dimorphocystis as the least degenerate, or the lobed
cystidia may have been evolved only in the Pistillarioid species. Some of the
irregularly excrescent hyphae in the sterile hymenium on the stem of Pterula
verticillata, P. brunneola, and P. vinacea become lobed, and they may" have
been converted into regular lobed caulocystidia while the more elongate
hyphal ends may have become the conical-ventricose caulocystidia. If regular
caulocystidia are formed, the minor factors evoking them may affect also the
primary hyphae of the fertile hymenium into which the cystidia may then be
regarded as intrusions. Alternatively, both kinds 0f cystidia may have been
truly hymenial in evolution, and the caulocystidia their degeneration. As I
have noticed on page 114, the study of the cystidia in the Pteruloid-series may
throw light on the origin of cystidia in general.
The caulocystidia of Dimorphocystis are monomorphic, and this, as I have
explained on page 113, is explicable on the assumption that they are the relics
of a sterile hymenium, which has completely disappeared in D. capitatus
without caulocystidia.
If Dimorphocystis is degenerating by restriction of apical growth and the
loss of the form-factors of the fruit-body, is D. capitatus the last term? Its
fruit-body is restricted to one set of skeletal 4yphae of limited growth: it
has no trace of sterile hymenium on the short stem: and when the factor
evoking the hymenium appears, it instantly extends over the whole apex and
prevents further outgrowth. There can be only one further qualitative step
in degeneration and that is the loss of the stem so that the hymenium appears
as soon as the primordium is initiated: in other words, reduction to a sessile
pulvinate fruit-body 50-200 µ, wide. Such a state must resemble Wiesnerina
which, I am informed by Donk, is Cyphelloid. Nevertheless, relatives of
Dimorphocystis may be expected in this group rather than among the
Corticioid to which Pterulicium leads.
PTERULA Fr.
Definition. Fries founded this genus, in 1825, ori the rare P. subulata of
Europe. In 1830 he added the second European species, P. multifida, for
which Chevalier in 1826 had made the genus Penicillaria, and also the North
American P. plumosa. The genus soon became the same receptacle for all
tropical 'Clavarias' with filiform branches (some, indeed, are the conidial
states of larger Pyrenomycetes and others seem to be rhizomorphs, as of the
'horse-hair fungi').
My studies of the tropical species and of P. multifida have shown me that
the peculiarity which at once distinguishes the true species as a natural genus1s
the dimitic state with colourless skeletal hyphae and, of course, uninflated,
typically clamped, generative hyphae. The construction explains very largely
the slender branches which, in the absence of inflation, become no thicker
after they are laid down at the slender tips (unless the hymenium thickens
appreciably).
At first, Pterula s. str. will seem an awkward genus including large, highly
branched fruit-bodies and almost microscopic, simple, filiform fruit-bodies
apparently identical, even under a pocket-lens, with those of Ceratellopsis.
But they are merely the extremes of apical growth and branching of fruit-bodies
with practically identical construction, and I can find no difference between
P. verticillata, for instance, and P. gracilis which is not bridged by intermediate
species. The future may distinguish cystidiate and acystidiate series, and even
separate the pseudo-Typhuloid species with sclerotial surface to the stem.
DEVELOPMENT OF THE FRUIT-BODY OF PTERULA GRACILIS

Mycelial hyphae, 1·5-2·5 µ, wide, grow on the surface of the substratum as
well as within it. Over certain small areas the superficial hyphae branch and
interweave more profusely to form a thin white subiculum from which the
fruit-bodies develop. A single hypha, 2-3 µ, wide, turns away from the
subiculum into the air and grows negatively geotropically. When 50-100 µ,
long, a septum with a clamp is formed about the middle and at the same time
a few more hyphae turn outwards from the subiculum round the base of the
initial hypha. Soon a branch arises from the subterminal cell of the initial
ThxT-FIG. 43. Pterula gracilis: stages in the development of the fruit-body arising from a
single initial hypha: the centre figure showing subsidiary primordia developing from the
basal disk of the fruit-body and basidia initiated as out-turned hyphal tips, X 500.
hypha at its distal end, and similarly from the subterminal cells of the other
hyphae when they have formed their first septa; the fascicle of hyphae so
formed grows upward as _the primordial shaft, indeed as the axis and substance
of the whole fruit-body, so simple is it. The fascicle thickens at the base by
the addition of more hyphae from the subiculum, and the apex widens by the
continued branching and upgrowth of the lateral hyphae. The hyphae inter
twine, often with a spiral direction so that the mature fruit-body appears to
have a half-twist under the microscope. Growth in length of the fascicle is
TExT-FIG. 44. Pterula gracilis: t.s. of a fruit-body, x 500.
entir-ely apical, the cells not enlarging after being cut off, and growth in
breadth occurs by branching and addition of more hyphae from the subiculum.
Many such primordia develop from one mycelial patch: most of them abort
at some early stage and comparatively few, 5-20, develop further. Some of
the primordia, if close together, may coalesce and form one large primordium,
but this rarely happens. The skeletal hyphae are formed early in the develop
ment of the fascicle and they continue unbranched until the cessation of
apical growtk, but their tips are not the leading hyphae.
The apex of the primordial shaft varies from blunt and wide and composed
of many hyphal tips about the same level to filiform and attenuate to a single
leading hypha. The two extremes are doubtless caused by the early or late
appearance of the secondary hyphae round the base of the- initial one: if late,
the initial hypha leads; if early, the secondary hyphae catch it up. In no case
did I see a primordium arise from two or three hyphae simultaneously.
When it is 0·5-2 mm. long, the primordial shaft becomes invested with the
PTERULA 121
hymenium from the base upward. It is formed in two ways. Either, in the
ordinary· way, by the out-turning of the tips of the generative hyphae on
the sides of the growing apex of the primordial shaft, or from the tips of other
hyphae which have grown up later from the mycelium on the outside of the
shaft. In either case these superficial hyphae become closely septate towards
their ends before they turn out, and, on turning out, the tip becomes cut off
by a septum and develops into a cystidium or a basidium, in which case it
may grow out farther and cut off a short subterminal cell. Then by branching
from the subterminal cells new basidia are intercalated and the hymenium
becomes continuous. The hymenium forms I -2 mm. from the apex in
young fruit-bodies, but the distance closes as they grow older until, in some
cases, the hymenium forms over the apex by the conversion of the hyphal
tips into basidia. Many fruit-bodies become arrested, however, before they
are fully developed and their tips remain sterile: whether it is frost or tempor
ary drying or some internal factor which arrests them, I do not know.
As the hymenium develops by intercalary growth, it dilates the primordial
shaft and pulls the longitudinal hyphae apart, so that the tissue becomes loose.
The hymenium occasionally develops from the very base of the primordial
shaft, but usually at a distance of 100-500 µ, from the base, in which case the
short stem bears scattered processes like rudimentary basidia or cystidia from
its superficial hyphae: they must be regarded as rudimentary caulocystidia
which are never close enough to form a palisade or sterile hymenium. Old
fruit-bodies may appear to have a longer stem, even several millimetres in
length, but it is secondary and appears, as Patouillard pointed out, through
the collapse of the matured basidia in the older basal part of the fruit-body.
The feature emphasizes the limited growth of the hymenium which does not
thicken in these reduced species of Pterula. As a further complication, some
hyphae may grow up very late from the subiculum and form clusters of
basidia on the old and collapsed hymenium.
Lateral hyphae may also grow out from the stem and downwards into the
subiculum where they ramify among the mycelial hyphae and form a secrmdary
support for the enlarging fruit-bodies. They may produce a few lateral
subclavate processes, like sterile basidia, as do some of the mycelial hyphae
round the base of the fruit-body, but I never saw that any of these processes
developed into fertile basidia such as covt:r the resupinate patch of Pterulicium.
Occasionally these excrescent hyphae from the stem grow upward and, on
being supplemented by others, grow into a small adventitious basidiocarp
like th� normally adventitious ramifications of Deflexula.
Cytology. 1 The cells of the generative hyphae are binucleate. The fusion
nucleus of the basidium divides three times to form 8 daughter nuclei, one of
which enters each spore. The remaining 6 nuclei apparently degenerate.
Thus the fruit-body is diploid, in Buller's sense, but the basidia have only
2 sterigmata. The cystidia are persistently binucleate.
1
These notes were made in 1927 from sections of fixed material stained with iron-alum
haematoxylin.
122 TAXONOMIC AND MOEPHOLOGICAL NOTES
Spore-germination. The spores germinate readily by a single germ-tube
from one end, rarely with a second tube from the other end.
PTERULICIUM gen. nov.

I have separated this from Pterula because of the surprising resupinate
structure, like a Corticium, which develops from the bases of the Cl�varioid
fruit-bodies. The widely effused mycelial patches which occur in other
TEXT-FIG. 45. Pterula gracilis: l.s. of part of a fruit-body showing the skeletal and generative
hyphae: iron-alum haematoxylin preparation, X 1,000.
Clavarioid fruit-bodies, as Lentaria surculus and Clavicorona pyxidata, are, so

far as-I have observed, invariably without trace of hymenium and similar to
the mycelial expansions of many basidiomycetes, as Geaster and Marasmius,
being superficial mycelial growths preceding the development of tlie fruit
bodies. In Pterulicium the resupinate patch has the construction of a
Corticium and is formed by the outgrowth of generative hyphae from the
base of the Clavarioid primordia, unless in their absence it develops directly
from the mycelium in the substratum. The genus thus presents the remark
able instance of a Clavarioid fruit-body degenerating into a Corticioid through
geotropic response. The transformation is not gradual by the concrescence
of the branches into effuso-reflexed Stereoid form as in Thelephora ( = Phylac
teria), but by a special resupinate outgrowth which directly substitutes the
Clavarioid and differs from it in lacking skeletal hyphae (if such hyphae occur
in Stereum and Polyporus they are generally retained in their resupinate
derivatives). If, therefore, as seems likely, there are truly Corticioid derivatives
of Pterulicium, they will be extremely difficult to recognize and assign to the
PTERULICIUM 123
Pteruloid-group, unless they have such hereditary features as the characteristic

amygdaliform spores, the acute young basidia, and the tendency to branch
from the clamp. This possibility must be borne in mind when analysing the
resupinate Thelephoraceae. Pterulicium shows from what diverse sources
Corticia may be derived and that the final classification of the resupinak
genera must wait until the structure of the higher Basidiomycetes is known.
For this reason, and because P. xylogenum would have to become a subgenus
if plar,ed in Pterula, I think it preferable to emphasize it as the type of a genus.
The name will surely be useful. Other features which may be generic are the
shape and thickened wall of the spore (those of the bushy Pterulas being small,
ellipsoid, and thin-walled), the tendency to branch from the clamp (rarely
seen in Pterula), the acute young basidia, and the absence of caulocystidia.
The young basidia resemble those of Marasmiwi, Crinipellis, Physalacria, and
Hormomitaria. Branching from the clamp is an uncertain feature, occurring
in many unrelated fungi as species of Sebacina, Polyporus, and Stereum. It is
noteworthy that the adventitious basal branching of Deflexula is comparable
with the resupinate patch of Pterulicium, as an outgrowth of generative
hyphae from the base of the Clavarioid fruit-body, but in Deflexula the
Clavarioid form has itself become negatively geotropic.
DEVELOPMENT OF THE FRUIT-BODY
The primordia of the Clavarioid fruit-bodies arise as conical outgrowths,
0·5-1 X 0·1-0·2 mm., on the surface of the substratum, singly or in clusters:
they are not erumpent. They gradually lengthen to form the stem and then
develop the branches either by slight and almost microscopic flattening of the
apex, followed by polychotomy or dichotomy (Text-Fig. 244), or adventi
tiously as acicular outgrowths, scattered over the stem or grouped to give a
false appearance of polychotomy, in which case the main stem and the branches
have persistently filiform tips. At the same time as the Clavarioid primordia
are developing, generative hyphae grow out from their bases over the surface
of the substratum to form the Corticium-like patch, which extends radhlly by
marginal growth. If the primordia of the Clavarioid fruit-bodies are crowded
they may coalesce at the base and appear caespitose. Apical growth is
typically Clavarioid and the hymenium develops in the normal way by out
turning of the ends of the superficial hyphae on the sides of the apex (Text
Fig. 7). Their ends turn out to form narrowly clavate, then subventricose
cells which may immediately become basidia or not until they have cut off
1-2 short subhymenial cells. The hymenium develops on .the resupinate
patch in the same way, but unilaterally: the narrow basal layer of hyphae,
parallel to the substratum, in this patch corresponds with the longitudinal
medullary hyphae of the Clavarioid fruit-body, but without the formation of
skeletals (these appear restricted to apices with directed growth in length).
The compactness of the subhymenium is caused by outgrowth of lateral
hyphae to the surface of the hymenium from the deeply situated subhymenial
hyphae, and thus they fill the gaps caused by the collapse of the old basidia.
RAMARIA S. F. Gray, emend. Donk

Definition. The problem of Ramaria versus Clavariella has been explained
by Donk (1933, 1941), with whose conclusions Maire has agreed (1937).
Ramaria Holmsk. (1790, Beata Ruris), for branched Clavarias, has been taken
up on three occasions, by S. F. Gray (1821), Bonorden (1851), and, Quelet
(1888). For many reasons it is unsatisfactory and certainly not a natural
genus; thus Fries never recognized it except as a section of Clavaria s. lat.
The name must, however, be accepted in some sense when Clavaria is split
up into several genera. This Karsten failed to do when he separated the
branched, ochraceous-spored species as Clavariella (1881) : he ignored Ramaria
Bon. which was a valid name already proposed for all the Friesian species of
Clavariella. Thus, Clavariella must become a synonym of Ramaria Bon. and,
delimited in this way, Ramaria becomes a natural genus. Quelet and Ricken
(1918-20), however, included in their concept of Ramaria all the branched
species of Clavaria, which is as unacceptable to-day as it was to Fries. Donk,
therefore, emended Ramaria Bon. with R. botrytis as lecto-type, and gave it
the present satisfactory circumscription. When he wrote, however, it had not
been realized that S. F. Gray's Natural Arrat}.gement of British Plants was a
valid, nomenclatural work published later than Fries's Systema, vol. 1, though
in the same year (1821). Ramaria S. F. Gray was taken directly from Ramaria
Holmsk. and, thus, can be defined in the same way by taking, as Donk
approves, one of the original species described by Holmskjold, as lecto-type,
namely, R. botrytis. Admittedly, the eight species 1 of Ramaria given by Gray
as British do not belong to Ramaria emend. Donk, but they were obviously
just Gray's local floristic use of Ramaria Holmsk., and any other interpretation
of Ramaria (for Clavulina or Clavulinopsis) would result in such hopeless
confusion of nomenclature that the name would either have to be abandoned
or conserved in Donk's sense. Actually, it seems that all the uses of Ramaria
have been exactly the same, if independent, being based on the same pre
Friesian concept, until Donk's emendation. Acceptance of this point of view
disposes of the very undesirable Cladaria Ritgen, which seems also to have
been based on Ramaria Holmsk., and which, deservedly, has never been
used.
Phaeoclavulina (1896-7) was made in imitation of Clavulina (1888) but has
no such value. The 2-spored condition in the species of Ramaria resembles
that in most genera of Homobasidiomycetes and is not connected with
peculiarities in form or function of the basidium, as it is in Clavulina. Van
Overeem used Phaeoclavulina for species which grade into 3-4-spored states
(Ramaria, group Echinospora) and there is yet no evidence that these are
nearly related with the type of the genus R. macrospora.
The question of Dendrocladium is discussed on page 82.
1 R. amethystea, R. ceranoides, R. cinerea, R. coriacea, R. corniculata, R. pratensis, R. rugosa,
and R. tuberosa.
RAMARIA 125
FRUIT-BODY
Size. The largest of all Clavarioid fruit-bodies belong to species ofRamaria.
At present those of R. formosa and R. Zippelii appear the tallest, reaching as
much as 30 cm. high, though bigger specimens of other species may yet be
discovered. In massiveness and abundance of branching there is little to
choose, however, between the fruit-bodies of many species such as R. botrytis,
R.flava, R. aurea, R.formosa, R. Mairei, R. Zippelii, and so on. At the other
extreme occur such as R.pusilla, R. flaccida, and R. ochraceo-virens with fruit
bodies as small as 1·5 cm. high. But in each natural group of species there is
every gradation from the massive to the small and slender, e.g. R. botrytis var.
parvula (2 cm. high), R. Zippelii var. gracilis (2-8 cm. high), R. conjunctipes
(allied with R. formosa), and in troops of large fruit-bodies it is generally
possible to find reduced and abortive specimens in all stages of development
from I cm. high (these abortive specimens must not be mistaken for young
states of the normal fruit-bodies, the primordia of which are more massive).
While size, in itself, is thus no indication of affinity, small or medium-sized
fruit-bodies, up to 10 cm. high, characterize the natural groups Verrucispora
(e.g. R.ochraceo-virens) and Stricta (e.g. R. stricta). The discovery of a really
massive specimen of either of these groups will be extremely interesting.
Shape. The most strikingly different shapes are (i) the Leotia-like masses
of R. capitata, with viscid, conjoint, subcapitate tops (-I cm. wide) on the
mature branches, (ii) the cauliflower-like masses of R. aurea, R. botrytis, R.
botrytoides, R. rufescens, and so on, with massive fleshy stems and stout,
condensed, polychotomous branches, (iii) the strict tufts of R. stricta and R.
Zippelii with elongate, parallel, dichotomous branches, (iv) the laxly divaricate
tufts of R. polypus and R. crispula with crescentic axils (as in Clavulinopsis
corniculata), and (v) the caespitose tufts of R. conjunctipes and R. sinapicolor
with many· slender fasciculate, often connate, stems in place of the usual
massive trunk. Strangely enough, the simple fruit-body seems never to have
been evolved, unless the dubious R. cyatheae is such: but even amor..g the
rudimentary abortive primordia at the bases of large fruit-bodies I have not
seen a fertile, undivided, primordial shaft. The remarkable condition of the
Australian R. capitata appears to be connected with the normal state by the
Australian R. salmonicolor which develops somewhat nodular expansions on
the branch-tips. R. Invalii, R. gracilis, and R. palmata have a marked
tend.ency to flattened branching.
The different shapes occur in most of the natural groups and appear to be
varietal rather than specific. Thus R. formosa may have the cauliflower-form,
the strict-form, the laxly divaricate, or the caespitose with slender stems, and
R. ochraceo-virens, in miniature, has the same diversity. Both are common
species and their behaviour suggests that if the others were as abundant they
would also be as variable. In Malaya the commonest species is R. cyanocephala:
its characteristic form is that of R. stricta with crowded parallel branches,
dichotomous throughout in alternating planes, and with elongate internodes,
but it varies into a shorter and shrubbier polychotomous form and an irregular,
laxly divaricate form, though I have never seen it with the true cauliflower
form of R. botrytis. Therefore, at present, I do not regard the shape of the
fruit-body as indicating affinity and, in any case, the shape must be analysed,
as I have explained in the introductory chapter, before it can be understood
in a single species. Thus, the branching of the stem, whether almost from the
base or from the apex of a distinct trunk, is often emphasized, but it is never
considered whether basal branching is the appearance of early division of the
primordial shaft or of the concrescence of the bases of crowded primordia.
Similarly, outcurving outer branches or wide-spreading basal branches have
been emphasized without considering that the inner branches are mutually
supporting and that the outer and lower branches necessarily sag under the
weight of the upper branches: actually, the branches of the primordia grow
vertically, or phototropically, and never at the wide angle which their older
internodes assume.
Colour. This is, perhaps, the most important feature in the natural classifi
cation of the species but the one of which there is the least knowledge. At
present only the macroscopic appearance of the colour is known: more im
portant will be its microscopic distribution: most important, its chemical
analysis and comparison. Every species is pigmented to some extent, with the
possible exception of R. stricta var. alba, and this pigmentation seems resolvable
into four basal colours, namely, yellow, brown-ochre, red, and blue. So far as
I have observed the colours are primarily lipochroic (dissolved in the oil
guttulae of the cytoplasm), though they may be cytochroic (diffused in the
cytoplasm): they are not water-soluble and cystochroic like the similar
bright colours in Agarics (Hygrophorus, Russula, Amanita). In most species
the colouring is generally diffused, but many are characteristically metachroic
and acrochroic (with distinctively coloured tips to the branches). The colour
is made in the hyphae on the surface near the tips of the branches, and
the acrochroism is caused either by the appearance of another pigment in
the oHer cells or in the basidia (e.g. R. formosa with yellow tips), or by the
disappearance or dilution of the original colour which may or may not be
substituted in the older cells (e.g. brown R. cyanocephala with blue tips, the
blue colour disappearing, and the pallid yellowish R. decolorans or R. obtusis
sima with brighter yellow tips, or the pallid yellowish-tan R. botrytis with
rose-purple tips). R. subbotrytis and R. botrytoides give an instructive example
of the doubts which may arise from macroscopic description. R. subbotrytis
is normally co=al pink all over but may be pale cream-ochraceous with pink
tips, the discoloration being caused by bleaching through exposure to sun
and wind (Coker, Kauffman): the bleached state resembles the normal state of
R. botrytoides which appears to be truly acrochroic like R. botrytis, yet the
similarity between the two species is so great that R. botrytoides may be merely
a state of R. subbotrytis with even more unstable pigment: if so, then R.
apiahyana may be the most intensely coloured state of this paling series. But,
whereas metachroism is so well developed in the genus, I have detected no
RAMARIA 127
case of the hysterochroism which confuses macroscopic description in

Clavulinopsis: there is a spurious hysterochroism, however, in all the species,
particularly those with thickening hymenium, caused by the accumulation of
the coloured spores on the surface or in the thickening hymenium, so that the
fruit-bodies becoming darker brownish or ochraceous from below upwards as
they mature: the dark fuscous-ferruginous colour of R. fragillima, R. cyano
cephala; and R. Zippelii is caused mainly by the mature spores trapped in the
thickening hymenium.
The combinations and variations in intensity of the four basal pigments,
which certainly occur in Ramaria (see the list on p. 129), will give all the variety
of colours which have been described, but I am not certain whether the violet,
purple, or lavender colour of the nine species listed on page 128 is caused by
mixtures and variations in intensity of the basal colours (pink or red with blue)
or whether there is a distinct violaceous colour. In any case the species do
not appear to be so closely allied as to form a natural group: R. fennica, R.
fumigata, R. Mairei, and R. testaceo-violacea are rough-spored: R. subdecurens
has verruculose spores: R. cacao and R. subspi11ulosa have smooth spores: and
R. stricta var. violacea-tincta belongs to the separate 'Stricta' -group with
thick-walled hyphae. The green tips of R. testaceo-viridis and the faint
greenish tint in the tips of R. ·apiculata may indicate traces of the blue pigment
of R. cyanocephala; and the existence of this pigment, exceptional as it may
seem, for no other Clavarioid-fungus is blue (unless the doubtful Typhula
caespitosa), may be of great importance in the phylogeny of the genus. In
contrast with R. cyanocephala, the white tips of the closely allied R. Zippe/ii
and R. grandis are caused by the absence of colour in the hyphal tips. The
ochraceous and brown colours of R. Zippe/ii, R. gracilis, and R. lnvalii are in
guttulae in the subhymenial hyphae and in the basidia, the hyphae of the flesh
being colourless. It appears, from comparison of these species, that acrochroism
is caused by the pigment of the sterile hyphal tips at the apices of the branches
and that the general colour of the fruit-body is caused by pigment in the
hymenium (and subhymenium) which invests the branches. Thus the .,viola
ceous, lavender, and vinaceous colours of R. fennica, R. Mairei, &c., appear
as hymenial pigments, distinguishable morphologically, if not chemically,
from the blue and purple or vinaceous acrochroic pigments of R. cyanocephala
and R. botrytis.
Yet another colour-complication is the epichroism of many species (listed
on p. 128) in which the tissues change colour through oxidation on bruising,
cutting, and incipient drying. The colour-change may be to pi.nk or red, but
more usually to purplish-brown or vinaceous (as in Clavariadelphus). Old
specimens of such species, which have been battered by rain, often appear
dingy brownish, for the pink or vinaceous colour fades to a dull tan or brown,
and they may not be recognizable until examined microscopically. Thus, I
think, R. Bataillei and R. testaceo-fiava are merely epichroic states of R.
formosa. The green discoloration of R. ochraceo-virens is exceptional: it indi
cates cyanescence which occurs elsewhere mainly in the Boletus-Paxillus-series,
Lactarius, and some Gastromycetes, certainly in no other Clavarioid fungi.
It would seem that these chromatic oxydases were developed capriciously
without indication of affinity: nevertheless caprice is seldom found to be true
with better understanding, and 'rufescence' and 'vinescence' are character
istic of the Eu-Ramaria and Echinospora-groups.
FRUIT-BODIES WITH YELLOW TIPS

R. condensata R. obtusissima
R. cystidiophora R. ochraceo-salmanicolor
R. decolorans R. rufo-violacea
R.fennica R. sinapicolor
R.formosa R. stricta
R. luteofiaccida R. subgelatinosa
R. luteofusca R. testaceo-fiava
FRUIT-BODIES WITH RED, PURPLE, OR PINK TIPS

R. apiahyana R. holorubella
R. botrytis R. rufescens
R. botrytoides R. secunda (pinkish tips)
R. conjuncta (pinkish-yellow tips)
(R. Mairei, R. subdecurrens, R. subspinulosa, suffused lavender or purplish over the
branchlets.)
FRUIT-BODIES WITH ORANGE OR RUFLESCENT-ORANGE TIPS

R. aurea var. australis (form 1), R. nigrescens, R. fragillima
FRUIT-BODIES WITH GREEN TIPS

R. apiculata (occasional), R. testaceo-viridis
FRUIT-BODIES WITH BLUE TIPS

R. cyanocephala
BRANCHES OR STEM LILACEOUS, AMETHYST, VIOLET, OR

VINACEOUS
R.fennica R. Mairei
R. fumigata } viola- R. cacao lilaceous or lavender-
R. rufo-violacea ceous R. subspinulosa } pink branches or
R. stricta var. violaceo R. subdecurrens branchlets.
tincta R. testaceo-violacea
FLESH TURNING PINK, RED, PURPLISH, OR VINACEOUS ON

BRUISING
R. fiava; reddening in some collections.
R. sanguinea: blood-red to vinaceous.
? R. fumigata; slowly rose-red.
R. macrospora; reddening or blackening.
R. aurea var. australis; stem vinaceous brown.
R. Bataillei; brownish-violet.
R. brunnea; vinescent.
R. cacao; stem vinaceous.
RAMARIA 129
R. cyanocephala: purplish vinaceous.

R. decolorans: violet.
R. Jormosa: vinaceous brown or violet in some collections.
R. fragillima: purplish vinaceous.
R. grandis: purplish vinaceous.
? R. holoruhella: stem often stained dull vinaceous rose.
R. longicaulis: mycelium and stem pink: flesh sometimes vinaceous.
R. nigrescens: purplish vinaceous.
R. obtusissima: rufescent or vinaceous in some collections.
R. pusilla: purplish pink (var. australis-pink).
R. secunda: often reddish brown or dingy vinaceous.
R. stricta: vinaceous brown.
R. testaceo-fiava: vinaceous.
R. testaceo-viridis: turning umber.
R. xanthosperma: vinaceous or reddening.
R. Zippelii: purplish vinaceous.
FLESH TURNING GREEN ON BRUISING(? CYANESCRNT)

R. ochraceo-virens
DISTRIBUTION OF PIGMENTS IN THE TISSUES

Blue or greenish-blue guttulae in the hyphal tips: R. cyanocephala.
Reddish-brown or orange-brown guttulae in the hyphal tips: R. fragillima.
Brownish-ochraceous cytoplasm and guttulae in the basidia: R. cyanocephala, R.
fragillima, R. gracilis, R. Zippelii, R. ochraceo-virens, R. polypus, R. Invalii.
Longitudinal hyphae with yellowish or brownish guttulae: R. cyanocephala, R.
fragillima.
Hyphal walls pale ochraceous: R. cyanocepha[a.
Texture. The flesh varies from brittle to tough and gelatinous, with every
gradation. Brittleness is caused by turgid, thin-walled, short-celled hyphae
and therefore in most, if not all, species the branchlets are brittle. As the
hyphae inflate in the older branches, one would expect the flesh to become
more brittle: actually, it becomes less brittle because the cell-walls thicken
very slightly (0·3--0·5 µ.) in most, if not all, species, and the intercalary growth
of the hymenium slightly dilates the flesh and makes it spongy: moreover,
the lengthening cells snap less readily. Nevertheless some species, as R. aurea,
R. flava, R. formosa, R. botrytis, and particularly R. divaricata, remain fairly
brittle. In species with thickening hymenium and, thus, greater intercalary
growth, the flesh becomes more spongy, fibrous-spongy, or flaccid, as in the
group Echinospora (R. Zippe/ii, R. gracilis, &c.) and the group Verrucispora
(R. ochraceo-virens, R. flaccida, &c.). Toughness is caused by narrowness of
the hyphae, or lack of inflation, and by thickening of the hyphal walls. Narrow
hyphae with scarcely thickened walls (--0·5 µ.) cause the toughness in the
Verrucispora-group and in the miscellaneous species R.fennica, R.fumigata,
R. suecica, R. condensata, R. xanthosperma, and R. fiavo-alba. Strongly
thickened hyphal walls (0·5-2 µ.) causes the toughness of the Stricta-group
(R. stricta, R. gracilis, &c.).
Among the large fleshy species of Europe, Huber emphasizes th� watery
marbled (semi-hygrophanous) character of the flesh in such as R. aurea, R.
5119 K
fiava, R. botrytis, R. rufescens, and R. Strasseri in contrast with the non
hygrophanous R. formosa, R. fumigata, and R. Mairei. To what extent this
· is a useful distinction further field studies must decide: it would seem to
separate easily R. formosa from R.fiavobrunnescens var. aurea (watery-marbled),
and R. Mairei from R. paludosa (watery-marbled). For lack of information
the character cannot yet be used in a general key.
The gelatinous flesh of R. gelatinosa is caused by the gelatinization of the
hyphal walls. It is so extensive that, at first sight, one would imagine the
species to belong to another genus, but its affinity is clearly with R. formosa
and, as explained on page 594, R. subgelatinosa is exactly intermediate. R.
ignicolor and R. xanthosperma also have subgelatinous hyphae, particularly
the former which is peculiar in having no clamps. In this respect it may be
related with R. capitata, in which the expanded tips of the branchlets also
become subgelatinous and viscid. Whether the gelatinization is a primitive
or derived feature will be an interesting point to solve: it would appear to be
primitive.
In R. formosa, R. suecica, and R. subspinulosa the flesh, on drying, becomes
peculiarly friable and chalky, so that it is easy to distinguish the species among
herbarium specimens. The reason for this alter;ition is not known and needs
investigation. R. formosa and R. suecica are certainly closely allied, but
R. subspinulosa in its cinnamon-brown fruit-body with lavender-pink branch
lets seems more nearly allied with R. cacao or R. subdecurrens, the precise
affinity of all being obscure.
Smell. As a chemical indication, the smell of the fruit-body is often helpful
in recognizing species, if not groups of species (e.g. the species of Ganoderma,
the pungent-fragrant Hydnums, the Tricholoma sulfureum-group, the rapha
noid Cortinarii, and the fragrant Inocybes). In Ramaria there are three
distinct smells, the Ganoderma or paracresol smell in R. cyanocephala, R.
fragillima, and R. Zippe/ii (Echinospora-group), the fragrant or aniseed-like
smell of R. gracilis, R. palmata, R. capucina, R. cystidiophora, R. sinapicolor,
and R. conjunctipes var. odora (the last three doubtfully allied with the first
three), and the peculiar cacao-butter smell of R. cacao which becomes more
pronounced on drying. Coker also mentions a smell of 'old ham' as charac
teristic of certain species, e.g. R. obtusissima, R. secunda, and R. Strasseri:
it may be a faint mixture of the paracresol and aniseed smells.
HABITAT
All the species are forest-fungi and, excepting the lignicolous Stricta-group,
all are humicolous. The only records of species occurring in fields, as those of
Ramariopsis, Clavulinopsis, and Clavaria so commonly do, is Maire's R.
luteofusca and some of the South Australian species, but they may have grown
from forest-relics. A few species are characteristic of coniferous forests;
most grow in frondose forests, and so many common species have been re
corded from both frondose and coniferous forests that, excepting the Ver
rucispora and Stricta groups, there is probably no such distinction of habitat.
RAMARIA 131
The species of the Echinospora-group have been recorded only from dico
tyledonous forests. The species of the Verrucispora-group have been recorded
mostly from coniferous forests. Among the lignicolous species there is still
doubt whether those on coniferous wood can be distinguished from those on
dicotyledonous. Coker says that R. stricta grows only on dicotyledonous
wood in. North America, but Doty has questioned this for the western states
and it certainly grows as well on coniferous wood in Europe. R. apiculata, on
the other hand, does seem restricted to coniferous wood, as R. crispula and
R. Bourdotiana are to dicotyledonous. Whether the fem-trunk habitat of
R. filicicola, R. filicina, and R. cyatheae is peculiar, I am doubtful in view of the
varied habitat of the similar R. polypus of Malaya (where terrestrial Hygro
phori may be found in the fibre of leaf-bases on palm-trunks).
CONIFEROUS FRONDOSE AND CONIFEROUS
R. apiculata R. aurea
R.condensata R.fiava
R.cystidiophora R.fiavobrunnescens
R.eumorpha R.formosa
R.fennica ? R.fumigata
R.fiaccida (rarely frondose) R. Mairei
R.gracilis R.obtusissima
R.lnvalii R.secunda
R. myceliosa R. Strasseri
R. ochraceo-virens R.stricta
R.palmata R. subbotrytis
R.pusilla R.subspinulosa
R. subdecurrens
R.suecica (also frondose, Wehmeyer).
HYPHAE
It appears that in most species the hyphae are unspecialized, being rather
long-celled, moderately inflated, and with thin or very slightly thickened walls.
There is no secondary septation and only in R. ochraceo-virens have I found
any considerable development of narrow interweaving hyphae· among the
inflated longitudinal hyphae, as occurs also in many species of Clavulinopsis.
Clamps almost certainly occur in most species, but there have been several
errors in observation: they are said to be absent from R. capitata and R.
luteo-fusca, and they are certainly absent from R. ignicolor, the first and third
species being somewhat allied in other respects and, perhaps, representing a
special subseries.
In several species (R. cyanocephala, R. fragillima, R. luteo-fiaccida, R.
ochraceo-virens, R. Zi,ppelii) I have noticed the short clavate processes, like
sterile basidia, on the longitudinal hyphae (Text-Figs. 260, 26!), which occur
also in some species of Clavulinopsis and Ramariopsis, but what they repre
sent is impossible to say: they are generally too scattered to be considered
more than abortive hymenial branches in the tissue of the flesh. The two
special developments of the hyphae are the gelatinization of the walls, in
R. gelatinosa, &c., and the firm thickening of the walls in the Stricta-group. I
have discussed these features in the paragraph on Texture (p. 130). Both seem
to indicate close affinity of the species possessing them, the gelatinous species
being particularly near to each other and to the common R. formosa. The
thick-walled hyphae of the Stricta-group inflate in the ordinary way and then
thicken in the older tissue: they are thus normally monomitic (as in Lentaria).
But, in the mycelium which develops round the base of the fruit-body of
R. stricta, there are many narrow, thick-walled, and unbranched hyphae, like
skeletal hyphae: they are, perhaps, only thick-walled endings of mycelial
hyphae, rather than special structural elements, and they do not occur in the
secondary mycelium of the related R. polypus.
Some species have been described with the hyphae only 3-5 µ, wide, as if
they did not inflate, but the measurements may refer only to the longitudinal
hyphae next the hymenium which are generally less inflated than the inner
hyphae of the flesh.
HYMENIUM
The hymenium develops in the ordinary way by the out-turning of the
hyphal tips shortly behind the growing-points of the branches. It begins to
form on the primordial shaft before it subdivides: thus, it covers the fruit
body from the upper part of the stem, if not almost from the base, to the tips
of the branches, which eventually become fertile on cessation of upgrowth
unless, of course, as often happens, the tips are destroyed by adverse weather.
The extent of the hymenium on the stem varies considerably: thus R.
ochraceo-virens and R. cyanocephala appear to have completely sterile stems;
R. gracilis, R. Invalii, R. polypus, R. luteo-fiaccida, and R. stricta are fertile
almost from the base; R. aurea, R.fiava, R.fiavo-alba, R.formosa, R.fragillima,
R. Mairei, R. subgelatinosa, and R. Zippelii have the lower part of the stem
sterile, where it is more or less immersed in the humus, and this is clearly
the usual state. In R. cyanocephala the greater part of the stem is covered with
a sterile hymenium (see below), and in R. Zippelii and R. fragillima the lower
part of the stem (except the basal few millimetres) is covered by a sterile
hymenium which gradually gives place to the fertile hymenium in the upper
part. In the other species the sterile part of the stem is cove.red simply with
narrow longitudinal hyphae with a few abortive hymenial processes to the
exterior. The secondary mycelial felt at the base of the stem in the Stricta and
Verrucispora groups is sterile.
On the branches of some, if not many species, the development of the
hymenium is unilateral in that it is sterile and poorly developed on the upper
sides of the larger inclined branches (Text-Figs. 253, 265). Such is the case
in the following species:
R. apiculata (according to Coker), R. cyanocephala, R. fragillima, R. gracilis, R.
luteo-fiaccida, R. Molleriana, R. polypus, R. stricta, R. subgelatinosa, and R. Zippelii.
I did not find such dorsiventrality in my specimens of R. formosa, R. aurea,
R. fiava, R. Mairei, R. Invalii, R. ochraceo-virens, and R. fiavo-alba. The
distinction may be important and, though dorsiventrality in some degree
RAMARIA 133
undoubtedly occurs in a number of species, it should be carefully studied. A
pocket-lens will suffice in the first place, for the sterile hymenium is not waxy
and even but finely velutinate or subtomentose. It seems that the dorsiven
trality is absent from most species of Eu-Ramaria and present in most species
of the Echinospora, Verrucispora, and Stricta groups.
A second and more important point is that the hymenium thickens in
some species but not in others. Coker has recorded the thickening in thirteen
North American species, but I am doubtful if it really occurs in all of them.
Coker based his conclusions mainly on the presence of embedded spores
among the basidia, but a steady breeze might have the same effect, so that it is
necessary to observe the embedded basidia. In some species Coker found the
spores and basidia stratified in 2-4 layers, indicating that the thickening had
been intermittent, probably during wet weather, but I have not seen such
stratification in the tropical species. The following are the species in which it
occurs, or may occur:
Hymenium thickening
R. apiculata, R. cyanocephala, R.fragillima, R. gracilis, R. grandis, R. lnvalii (very
slight), R. luteo-fiaccida, R. polypus, R. stricta, R. subdecurrens, R. subspinulosa,
R. Zippelii.
Hymenium possibly thickening
R. fiaccida, R. fiavo-alba, R. fumigata, R. gracilis, R. obtusissima, R. ochraceo
virens, R. pusilla, R. secunda, R. Strasseri, R. subgelatinosa.
In contrast, the hymenium does not thicken in the following species:
R. aurea, R. botrytis, R. fiava, R. formosa, R. Mairei.
It seems, therefore, that the thickening hymenium, like the dorsiventral
hymenium, is characteristic of the Echinospora, Verrucispora and Stricta
groups and is absent from most of Eu-Ramaria.
BASIDIA
In most species there are 4, rarely 2, sterigrnata. In none has more than 4
been seen, but several species of Echinospora, R. luteo-fusca (Verrucispora)
and R. macrospora (Decolorans), have normally 2, sometimes 3, rarely 1. Thus:
R. lnvalii (2-3-) 4
R. longicaulis (2-3-) 4
R. fragillima (1-) 2-3-4, mostly 4
R. nigrescens 2
var. americana 4
R. cyanocephala (1-) 2 (-3)
R. grandis, R. macrospora 2
R. luteo-fusca 2
R. Zippelii 1-2
var. gracilis 2-3-4, mostly 3 .:..4
Except R. luteo-fusca and R. macrospora, the other seven species are nearly
allied and form the natural Echinospora-group. R. luteo-fusca is said to have
only verruculose spores, but it probably belongs to the same group. R.
macrospora has merely rough spores and seems allied with the 4-spored species
R. decolorans and R. xanthosperma. In R. fragillima the number of sterigmata
is most variable and many basidia have sterigmata of unequal length, 1-3 of
which may be abortive in all stages from bearing a rudimentary spore to being
a rudimentary knob on the basidium. It is the only Basidiomycete with
active basidia in which I have seen such a remarkable transition from 4-spored
to 1-spored basidia, and it shows how very gradually the evolutionary process
can take place, though the transition from the 4-spored to the 2-spored state
would appear a simple mutation. In these species also, with reduced numbers
of spores, the symmetry of the basidial apex may be so disturbed that the
spores develop asymmetrically on the sterigmata, having the �piculus directed
laterally or abaxially instead of adaxially (Text-Figs. 257, 271) ..
CYSTIDIA
Kauffman has described cystidia in one species, R. cystidiophora. Coker
did not find cystidia in the North American species which he studied, nor have
they been reported by European mycologists. I saw no cystidia in the twenty
species which I examined. It may be doubted, therefore, whether Kauffman's
observation was correct and did not refer to excrescent hyphae or basidia in a
thickening hymenium. Admittedly, however, there is a comparable case in
Clavulina, where only C. Leveillei has cystidia, and C. cristata var. incarnata
may develop cystidioles in exceptionally humid surroundings: perhaps, too,
with Clavaria purpurea.
SPORES
These spores provide four diagnostic features which, in order of impor
tance, are the surface-marking, the shape or ellipticity, the colour, and the
actual size: a fifth, perhaps, should be added in the spore-contents, whether
guttate or aguttate. About all there is much still to be learnt, though in
recent years many accurate data on size and colour have been accumulated.
I have made free use of these in my treatment of the genus, and I gladly
express my obligation to the mycologists from whom I have borrowed.
Colour. In most species the spores are ochraceous. In the Echinospora
group occur the most deeply pigmented spores, fuscous ferruginous, in the
species which also have the mostly deeply pigmented fruit-bodies. Like
wise, in the species with pale fruit-bodies, the spores are generally pale
ochraceous or pale yellow, as in several of the Stricta and Decolorans groups,
but there are exceptions, as R. polypus with pale fruit-bodies and cinnamon
spores. In a�few species the spores are brownish pink or cinnamon rather
than yellowish, but they do not seem to be nearly related, e.g. R. polypus,
R. subbotrytis, and R. suecica. The colour is in the spore-wall, and the species
with distinctly warted or echinulate spores have colourless projections from
the ochraceous or brown spore-wall: the pinkish tinge, giving the cinnamon
spore, probably occurs in the cytoplasm. Mycologists do not usually take
spore-prints of Clavarias, but there is sufficient variety of colour in Ramaria
to make the study worthwhile.
RAMARIA 135
Surface marking. The spore-wall may be smooth or.it may be marked with
one of these features:
Echinulate with discrete, acute, hyaline spines o· 5-3 µ,high: e.g. Echinospora-group.
Verruculose with discrete, blunt, hyaline warts o· 5 µ, high or less: e.g. Verrucispora
group.
Rough with minute raised points of immeasurable height (minutely rough) or with
very slightly raised, irregular, small patches like vague tesselations or smears (coarsely
rough): e.g. R. Mairei (minutely rough) R. ftava, R. aurea, R. formosa (coarsely to
minutely rough).
Rugulose with vaguely confluent raised points or patches: e.g. R. polypus, R. sub
sigmoidea.
Striate with faint longitudinal striae on the smooth wall, the striae apparently innate
and not raised or sunken: e.g. R. botrytis, R. holorubella.
The surface-markings appear to give the most important indication of
specific affinity in the genus and, though accurate information has accumu
lated in the last 20 years through the stimulus of Coker's work, there is still
need for thorough research into almost every species. Needless to say, the
research must be made with the highest powers of the microscope for the
markings are very minute, especially in their degradation: and in the case of
dried spores it is necessary to say in what medium thay have been examined,
for the swelling of the epispore may cause the finer points to disappear: I
prefer, and always use, dil. KOH (5 per cent.), because it not only swells but
slightly, but may have an interesting chemical action, and it is easy to prepare.
Each kind of marking grades into another, but there are probably fewer
species intermediate in this character than in any other. The echinulate
spores are easily recognized, though the shorter spines of R. Invalii may be
mistaken for the blunt warts of R. ochraceo-virens or R. fiaccida: thus R.
luteo-fusca, described as verrucose, may be echinulate. It seems that the
smaller the spore the shorter the spines. Thus the echinulate state passes
into the verruculose state of the relatively small-spared species of the
Verrucispora-group. The one species with large verruculose spores appears
to be R. apiahyana, but its spores may really be echinulate OJ.° co�sely
rough. And, therefore, what is the connexion between rough and echinulate
verruculose spores has yet to be made out. Many rough spores are large
and coarsely rough, as . in R. fiava and R. formosa: others are minutely
rough, as in R. Mairei: and both states occur also among small spores,
e.g.· R. subbotrytis and R. botrytoides or R. gracilis and R. stricta. The
roughness is exceedingly difficult to define in words or in drawing, because
of the vague outline of the raised areas, and it is impossible _at present to
analyse the feature in detail from the published descriptions: a comparative
study of the spore-prints of many species, the fresh prints collected on
microscope-slides, would be the ideal method of research. The rugulose
spore may be only a variation of the rough spore, but the rugulose and
aguttate spores of R. polypus, R. fiavo-alba, and R. subsigmoidea suggest a
special state.
The smooth spore may be large or small and, a priori, it may be related to
either rough or verruculose or echinulate spores. But if the relation between
the length of the spore and the length of the spine is true, only short smooth
spores can be related to verruculose and echinulate spores: such may explain,
in part, the smooth spores of the Stricta-group, some species of which appear
allied with the Verrucispora-group. In Eu-Ramaria, however, the smooth
spore is undoubtedly related to the rough spore, but whether the relation
indicates affinity between the smooth-spored species or has evolved inde
pendently on different occasions it is hard to decide. Thus R. conjunctipes
var. odora, with small smooth spores, is closely related to R. formosa with
coarsely rough spores. But in the Decolorans-group, on the other hand, the
smooth spores seem to indicate affinity of the species, e.g. R. cacao, R.
Strasseri, R. obtusissima, R. decolorans, R. xanthosperma, and R. verna.
The striate spores of R. botrytis and R. holorubella are merely a special case
of the smooth spore. The markings indicate, in some way, the micella
structure of the spore-wall and they can be seen in immature spores of
R. formosa, R. flava, R. aurea, and R. Mairei, for example, before they are
obscured by the roughening of the epispore.
Size. In most cases the size of the spore does not indicate affinity. Thus,
in R. formosa the spores vary, in different collections, from 7-9 X 4 µ, to 13-
17 X 5-7 µ,, and the closely allied R. conjunctipes var. odora has spores 6 X 4· 5 µ,.
A glance through the key will show that species with spores about the same
size are by no means nearly related. The two notable exceptions are the
Verrucispora and the Stricta groups, both with comparatively small spores,
5-1ox3-5 µ,.
Shape. (I defer this subject for a later work: see Corner, New Phyt. 1948.)
Contents. In most species the spores contain 1-4 large oil-drops or several
smaller ones. In R. polypus and R. flavo-alba I found the fresh spores were
aguttate, with slightly opalescent cytoplasm. Whether this distinction has
much meaning in the genus is impossible to say, but it should not be forgotten.
Coker gives R. pusilla as aguttate and R. subsigmoidea also appears to be
agutt.11te: it has the same brownish rugulose spores as R. polypus.
CYTOLOGY
There appears to be only Juel's observation on R. Invalii (Clavaria
abietina), in which he found the first nuclear spindle in the basidium to be
transverse and the spores to be uninucleate.
CLASSIFICATION
After many attempts to arrange the species according to their natural
affinities I have arrived at the classification set out in the Key on page 545.
The four main groups are certainly natural and might well be reduced to
three, by the combination ofEchinospora with Verrucispora into one subgenus,
and perhaps to two, by the combination of Stricta withEu-Ramaria. However,
the exact position of several species in both Verrucispora and Stricta is un
certain, through lack of precise observation, and it is simpler at the present
RAMARIA 137
stage of knowledge to keep apart the better-known Echinospora and Eu
Ramaria. The five subdivisions of Eu-Ramaria also seem natural, though
part of Formosa may belong to Flava and parts of Decolorans to Botrytis and,
even, Stricta or Verrucispora. Now, because I regard this classification as
tentative, being the first essay in arranging the species of the world, I have
refrained from making subgenera, sections, or any other systematic division
to which laws of nomenclature may be applied. The nine names which I have
used are fancy words of a nature quickly to call to mind the content of the
group, in place of numbers which are meaningless. Later classifications may
re-arrange so many of the species as to render the names inapposite and it
is time that we sought to avoid the sorry burden of nonsensical terms which
the legalization of the historical growth of systematy has forced upon us. My
notions of fungi may seem, in a hundred years, as infantile as those of Fries.
The correct grouping of species in subgenera is far more difficult than the
recognition of species or genera, for it supposes (what is seldom realized)
the understanding of the evolution of the species: and that must be left for
those who can arrive. At present, we can only indicate.
The first division of the Key is on the spore-markings. The members of the
Echinospora-group are at once distinguished by the sharp hyaline spines on
their spores. The larger their spores, the stronger the spines, as I have already
remarked; the smaller their spores, the shorter their spines and the less their
ellipticity until they become like the spores of the Verrucispora-group.
Indeed, R. lnvalii is placed in Echinospora, and not in Verrucispora, merely
because its spines are acute, and R. luteo-fusca is placed in Verrucispora,
rather than Echinospora, because its warts appear to be blunt: yet, in their
extremes, such as R. cyanocephala and R. myceliosa, the two groups are far
apart. There is no evidence that the spores of either may become merely
rough or smooth, like those of Stricta and Eu-Ramaria.
The second division of the Key is a combination of the characters of the
habitat and the hyphae, whereby the mainly lignicolous Stricta with thick
walled hyphae is separated from the terrestrial Eu-Ramaria with thin-walled
hyphae. Thus the large, fleshy-brittle, clear, or brightly coloured species
with simply rough or smooth spores are separated as the main group of the
genus. The third and fourth divisions of the Key distinguish.the subgroups of
Eu-Ramaria by means of the colour of the fruit-body and the roughness or
smoothness of the spore. Unfortunately neither character is absolute and
these subgroups will doubtless undergo much revision.
I will now consider the groups in detail, to show their nu,dei and their
anomalies.
ECHINOSPORA. This group comprises the following eight species, the
first five of which are the most representative:
R. cyanocephala, R. fragillima, R. grandis, R. nigrescens, R. Zippelii; R. Invalii, R.
longicaulis, R. Murrillii.
(Doubtful species: R. albida, R. apiahyana, R. guadelupensis, R. guayanensis, R.
luteo-fusca, R. cervicornis, R. tubulosa.)
The fruit-bodies vary from massive to small and from polychotomous to
dichotomous. Their hyphae are thin-walled and the texture more or less
fleshy or rather fibrous, when the hyphae are little inflated (R. Zippelii). The
predominant colour is brownish ochre, caused mainly by the brownish ochre
cytoplasm and yellow.ish guttulae of the basidia, but in the first five species,
particularly, the colo�r is intensified to rich brown, ferruginous, fuscous
brown or umber.• Acrochroism is vivid and characteristic of R. cyanbcephala
(blue), R. fragillima and R. nigrescens (orange-rufous): in contrast R. Zippelii
and R. grandis are two examples of the unusual state of a highly pigmented
mature fruit-body wit.h colourless tips. The clear, bright colours of Eu-Ramaria
are absent, even the \miform pale yellow of Stricta, but the three species
R. Invalii, R. longicaulis,. am! R. Murrillii grade both in colour and size of
the fruit-body into the small ochraceous fruit-bodies characteristic of Ver
rucispora. The first five species and R. longicaulis have the vinescent epi
chroism characteristic of Ramaria but it is absent from R. Invalii (perhaps
also R. Murrillii) and from R. nigrescens var. americana, as it is generally from
Verrucispora. The strong smell of Ganoderma (like paracresol) occurs in
R. cyanocephala, R. fragillima, and R. Zi'ppelii, but appears to be absent from
the others, even from R. Zippelii var. gracil£s. All are humicolous forest
dwellers and, probably, in the main tropical.
The hymenium is dorsiventral and thickens more or less strongly in all
except, possibly, R. nigrescens and R. Murrillii, in which the feature may have
been overlooked.
A peculiarity is the degeneration of the basidium from the 4-spored to
the 2- and 1-spored state. The 2-spored state gave van Overeem the idea of
Phaeoclavulina in imitation of Clavulina. Such a generic distinction may be
incipient in Echinospora, but it is impracticable for it cuts a species from its
variety (var. americana from R. nigrescens) and 1-4-spored basidia occur on
the same fruit-body in R. fragillima. Moreover, the type-species of Phaeo
clavulina (R. macrospora) does not belong to Echinospora.
Cop.cerning the spores, there are two important indications. Firstly, the
spores are broader than usual in Ramaria. Secondly, as I have remarked, the
longer spores have longer spines (e.g. 1·5 µ, long in R. cyanocephala, 0·5-1 µ,
long in R. Invalii; 1-2 µ, long in R. Zippelii, 0·5-1 µ, long in var. gracilis).
Neither in spore-shape nor spore-markings, nor in the general colour of the
fruit-body, nor in the thickening of the hymenium does Echinospora grade
into any other group of Ramaria except Verrucispora.
VERRUCJ..SPORA. This group comprises eleven species, of which
R. ochraceo-virens, R. fiaccida, and R. luteo-fiaccida are the most typical
representatives:
R. capucina, R. eumorpha, R. filicicola, R. fiaccida, R. luteo-fiaccida, (R. luteo-fusca)
R. megalorhiza, R. myceliosa, R. ochraceo-virens, R. pusilla, (R. subdecurrens).
(Doubtful species: R. subfennica, R. subdecurrens, R. subspinulosa, R. crispula.)
Excluding the doubtful species Verrucispora consists of nine species with
small fruit-bodies (rarely above 5 cm. high), with pallid yellowish to ochraceous
RAMARIA 1 39
or brownish ochraceous colour and small verruculose spores (mostly less than
8 µ, long). There is no acrochroism (unless in the brighter yellow tips of
R. luteo-fiaccida) and epichroism occurs only in the vinescent R. pusilla and
the cyanescent R. ochraceo-virescens. In habitat there are the following
differences :
CONIFEROUS DICOTYLEDONOUS
R. ochraceo-virens R. eumorpha
R. fiaccida (R. fiaccida ( ?))
R. pusilla R. pusilla var. australis
R. myceliosa R. capucina
R. megalorhiza
R. luteo-fiaccida ( ?)
(R. subdecurrens) (R. luteo-fusca)
(R. subspinulosa) (R. subspinulosa)
(R. gracilis) (R. crispula)
(R. subfennica)
The hyphae appear to be thin-walled and the texture fibrous-tough through
the slight inflation of the hyphae (? thick-walled in R. megalorhiza which may
belong to Stricta). The hymenium is dorsiventral and thickening in R. luteo
fiaccida: the state of the other species is uncertain (though it thickens in
R. subspinulosa and R. subdecurrens of doubtful position): it possibly thickens
in R. ochraceo-virens and R. fiaccida. The basidia are 4-spored in all but
R. luteo-fusca, which may belong to Echinospora.
STRICTA. I have placed eleven species in this group:
R. apiculata, R. Bourdotiana, R. crispula, R. durbana, R. gracilis, R. Moelleriana,
R. nguelensis, R. ochracea, R. palmata, R. polypus, and R. stricta.
The temperate R. apiculata, R. stricta, and R. Bourdotiana and the tropical
R. Moelleriana form the nucleus, being lignicolous with thick-walled hyphae.
R. crispula is lignicolous but its hyphae appear to be thin-walled. R. gracilis
and R. palmata have thick-walled hyphae but are terrestrial. R. polypus agrees
with the nucleus of Stricta but it has rugulose aguttate spores like R. fiavo-alba
and R. subsigmoidea and it may conceivably resemble R. stricta by convergence.
R. nguelensis is a dubious African species and R. durbana may be Scytinopogon.
R. megalorhiza (placed in Verrucispora) may belong to Stricta, at least in the
affinity of R. polypus.
If the group, therefore, may be varied in origia as the final term of several
convergent series, it is superficially uniform in the small to medium-sized,
tough fruit-bodies (-10 cm. high), with pallid white, tan, yellowish, ochraceous,
or cinnamon-ochraceous colouring, and small, rugulose, rough or nearly, or
quite, smooth spores (-10 µ long). R. stricta is acrochroic with yellow tips,
the other species have pallid concolorous or white tips (e.g. R. polypus).
R. gracilis and R. palmata have a characteristic fragrance which may help to
place them with allied species, as I have remarked under R. gracilis. The
hymenium is dorsiventral and thickening in R. apiculata, R. gracilis, R. polypus,
and R. stricta, but its state is not known in the others.
FORMOSA. This group includes the species of Eu-Ramaria which have
pink or pink and yellow pigmentation and rough or smooth (but not striate)
spores. There are eighteen species, two of which (in brackets) doubtfully
belong:
(R. apiahyana), R. Bataillei, R. botrytoides, R. brunnea, R. conjuncta, R. conjunctipes,
R. divaricata, R. formosa, R. gelatinosa, R. ignicolor, R. ochraceo-salmonicolor, (R.
rufescens), R. secunda, R. subbotrytis, R. subgelatinosa, R. suecica, R. testaceo-fiava, R.
testaceo-violacea.
The fruit-bodies vary from massive to rather small and slender in R.
conjunctipes. The tips are bright yellow or pink in some species and the
yellow and pink colours may fade more or less from the older parts. R.
formosa, R. Bataillei, R. testaceo-flava, R. brunnea, and R. secunda are more
or less vinescent. The texture of the flesh varies considerably, even among
closely allied species, and it needs microscopic analysis for proper under
standing. Thus R. gelatinosa and R. subgelatinosa, with gelatinous hyphal
walls, are closely allied with typical R. formosa. R. suecica has softly coriaceous
flesh which dries chalky-friable like that of R. formosa. R. conjunctipes has
rather tough flesh (? thick-walled or narrow hyphae). The other species are
fleshy-brittle, like R. formosa, but their flesh does not dry chalky-friable,
and is, perhaps, watery-marbled in the living state. The hymenium does not
thicken, unless slightly in R. subgelatinosa.
The species fall into two series:
Formosa-series: predominantly yellow tips and extensive yellow coloration.
R. formosa, R. subgelatinosa, R. gelatinosa, R. conjunctipes, R. ochraceo-salmoni
color, R. Bataillei, R. testaceo-fiava, R. ignicolor, R. brunnea.
Subbotrytis-series: predominantly pink coloration.
R. subbotrytis, R. conjuncta, R. divaricata, R. suecica, R. secunda, R. botrytoides,
R. rufescens, R. testaceo-violacea, (R. apiahyana).
The larger spores in this group are the rougher, the smooth spores occurring
in the comparat,ively small-spored species R. conjunctipes, R. subbotrytis,
R. botrytoides, R. rufescens, and R. secunda.
FLAVA. I have placed eight species in this group:
R. aurea, R. capitata, R. cystidiophora, R. fiava, R. fiavobrunnescens, R. madagascari
ensis, R. sanguinea, R. sinapicolor.
The group is very near Formosa, but lacks the pink colour and has, on the
whole, slightly narrower spores. The yellow forms which have been ascribed
to R. formosa, R. divaricata, and R. obtusissima probably belong to R. flavo
brunnescens. All the species appear to be fleshy-brittle and watery-marbled
when fresh. The hymenium does not thicken and is not dorsiventral.
R. capitata is remarkable for the capitate, viscid, subgelatinous tops of the
branches which become confluent into Leotia-like masses. The hymenium
develops over the viscid tops as well as over the normal branches. R. ochraceo
salmonicolor (Formosa) seems to be allied, so that the species may really belong
to the gelatinous group of Formosa.
R. cystidiophora is peculiar in combining rather massive fruit-bodies with
unusually small spores.
RAMARIA
VIOLACEA. There are four species in this group, the last being doubtful:
R. fennica, R. fumigata, R. Mairei, R. rufo-violacea.
The group can be described as Flava suffused violet. R. Mairei is the least
characteristic, for its fruit-bodies are yellow suffused pale lilac or pale purple
on the branches: it may even be entirely yellow and thus it has certainly been
confused with R. fiavo-brunnescens. R. fennica has a violet stem with yellow
branches which become suflused violet acropetally (and therefore appear
olivaceous). R. fumigata is entirely violet or lilac and, thus, appears to be
R. fennica without the yellow pigment, or even R. Mairei without its paler
yellow pigment.
DECOLORANS. This is the most unsatisfactory group and is more a
collection of misfits than a natural assemblage. I have placed in it the
fourteen species:
R. cacao, R. condensata, R. decolorans, R. fiavo-alba, R. Karstenii, R. macrospora,
R. obtusissima, R. spinulosa, R. Strasseri, R. subsigmoidea, R. subspinulosa, R. testaceo
viridis, R. verna, R. xanthosperma.
The group is distinguished more by the lack of characters, e.g. pale colour
and smooth spores, than by their presence and, therefore, a priori, its species
might be derived by loss of character from any of the other groups of Eu
Ramaria. If, for example, it were not for R. subgelatinosa, it would have been
tempting to add R. gelatinosa to Decolorans.
There appears to be a narrow-spored series consisting of R. xanthosperma,
R. obtusissima, R. verna, and R. decolorans (perhaps identical with the first),
A broad-spored series seems to consist of R. Strasseri, R. spinulosa, and R.
condensata.
R. cacao and R. subspinulosa seem allied in their brownish colour with pale
lavender-pink branchlets, but R. subspinulosa is the only other species known
which agrees with R. formosa and R. suecica in the chalky-friable texture of the
dried flesh. Possibly R. brunnea (Formosa) connects them.
R. testaceo-viridis seems allied with R. ochraceo-virens (Verrucispora).
R. fiavo-alba and R. subsigmoidea, with rugulose aguttate spores, may be
allied with R. polypus (Stricta).
On the other hand, if the group is natural, there is the peculiar feature that, in
contrast with Echinospora, Formosa, and Flava, the small epores are rough and
the large spores smooth. Thus Decolorans appears to stretch into Stricta with
small spores and to the next group l3otrytis with large smooth spores, this
anomaly being inexplicable in any other group. T_he relation with Botrytis
would explain the evanescent lilac pigment of R. cacao and R. subspinulosa, the
origin of which must elsewhere be sought in Violacea
BOTRYTIS. This is the nomenclatorial nucleus of the genus, but only
two species are known with the same persistently smooth, finely striate spores,
namely R. botrytis and R. holorubella. In their reddish-purple tips they
resemble the Subbotrytis-series of Formosa but differ in the smooth spores,
the large size of which suggests, as just mentioned, affinity with Decolorans.
EVOLUTION
The classification which I have given is intended to be natural in that I
have endeavoured to bring together related species in one group. Such a
classification supposes that the groups have a common origin in an ancestral
form. The discussion in the preceding section may have been so bewildering
that I now give a diagram of the possible phyletic relations of the.groups.
The diagram is meant to be useful, to provide hypotheses which can be
tested, rather than supposedly real, for in the present state of ignorance over so
many living species one cannot hope to see clearly into the past.
ANCESTRAL RAMARIA
�
Eu-RAMARIA
ECH1N0SP0RA
FORMOSA
V10LACEA
'\
BoTnvT1s
VERRUCJSPORA
narr?w�spored
'
LEN;:-ARIA STRICTA
\/
Su�U\otrytis•
series
Formosa ..
series
The only primitive feature which indicates the evolution of the species is
the thickening hymenium which is so extensive in the Echinospora-group and
so generally absent from Eu-Ramaria. It occurs also in the Stricta-group
and probably, to a slight degree, in some species of Decolorans, wherefore I
think these two groups may have diverged from the Eu-Ramaria stock before
it evolved its modern series of species, particularly the Formosa-group. On
RAMARIA 14 3
the other hand the advanced feature which indicates most clearly the evolution
of the species is the thick wall of the hyphae in Stricta.
These two features present ( 1) the picture of a primitive species with
massive, heavily pigmented, fleshy terrestrial fruit-body with thin-walled,
inflating hyphae and large, broad, dark brown, echinulate spores: (2) the
picture of the advanced, or most derivative, species with small, slender, pale or
colourless, tough, lignicolous fruit-body with thick-walled, uninflating hyphae
and small, pale yellow or nearly colourless, smooth spores. Thus, as a general
principle in the genus, the species with small degenerate fruit-bodies have
small degenerate spores. The Verrucispora-group appears as the natural
product of the degeneration of the Echinospora-group: its features could,
indeed, have been predicted. Stricta appears as the degeneration of Eu
Ramaria, though its exact affinity is uncertain, possibly with Decolorans.
R. conjunctipes appears as the end-product of the Formosa-series with small
fruit-bodies developed from diminutive, caespitose primordia and with short
smooth spores.
The new features, indicative of new genera, evolved by the modern species
of Ramaria are the 2-spored 'Clavulina' tendency in Echinospora, the thick
walled lignicolous tendency in Stricta (which may have produced Lentaria,
see p. 94), the gelatinous tendency in Formosa, the capitate tendency in
R. capitata (Flava), and the softly coriaceous tendency in R. suecica of the
Subbotrytis-series. The most curious feature of the whole genus, however, as
already mentioned, is the total absence of simple monaxial fruit-bodies. Henn
ings has described a simple filiform Clavaria with smooth, subglobose, brown
spores (4X3 µ) from New Guinea, namely, C. subfistulosa, which I have left
among the doubtful species of Clavaria: unfortunately, it must be regarded
with suspicion, like all of Hennings's description, until it has been redescribed.
Now the hypothetical ancestor of Ramaria assumes a bizarre form, the
discovery of which in a tropical forest would throw a mycologist into ecstasy.
Yet, is it not almost exactly the description of R. apiahyana? It appears to be
a tacit assumption that primitive spores are small, smooth, and colourless, as
a corollary of the more explicit idea that large, coloured, and ornamented
spores are advanced. I can discover no evidence for the assumption and,
considering how common are small, smooth, white spores among modern
fungi compared with the large, coloured, ornamented spores, I would infer
from the general principles of evolution that the first were derived, or advanced,
and the second primitive.
NAME-CHANGES
To assist reference to previous works, particularly of Coker, I give the

following list of changes or substitutions of specific epithets.
Name used in this work
C. abietina auctt. R. ochraceo-virens
'non-virescent form of pines' R. lnvalii
'Small non-virescent form' R. fiaccida
Name used in this work
C. botrytis sensu Coker R. botrytoides
C. botrytis sensu Kauffm. R. holorubella
C. botrytoides sensu Coker R. botrytoides, R. conjuncta
C. Broomei R. nigrescens
C. condensata sensu Bourd. et Galz. R. Bourdotiana, R. stricta
C. decurrens sensu Coker R. pusilla
C.dendroidea auctt. R. condensata
sensu Bourd. et Galz. R.stricta
C. fennica sensu Coker R.fumigata
C. fiava sensu Coker R. fiavobrunnescens
var. subtilis Coker R. divaricata
C. rufescens sensu Coker R. holorubella
sensu auctt. ? R. botrytis (non R. rufescens)
C. stricta sensu Bourd. et Galz. R. Bourdotiana
sensu Coker R. stricta var. concolor
var. condensata Lundell R. stricta
C. versatilis R.fumigata
C. Zippe/ii auctt. R. cyanocephala, R. Zippe/ii
RAMARIOPSIS Donk emend.

I have followed Donk (1933) in regarding Clavaria Kunzei as the type of a
group of fleshy Clavarias characterized by the small, white, finely echinulate or
verruculose spores. Donk made C. Kunzei the type of his subgenus Ramari
opsis of Clavaria, and included in it C. subtilis, C. pyxidata, C. angulispora,
and C. pulchella. I have raised the group to generic rank and excluded
C. subtilis ( = Clavulinopsis), C. pyxidata ( = Clavicorona), and C. angu
lispora ( = Scytinopogon). If it is considered merely subgeneric then it
belongs to Clavulinopsis, comparable with the subgenus Acularia. However,
it is a very homogeneous natural group, the species of which differ mainly in
colour and slight details of the spores, and it must be connected with the
branched ancestral group of Clavulinopsis, but, at present, no bridging species
seems to be known. At first sight R. pulchella, R. crocea, and R. Kunzei seem
entirely different fungi, better to be classified with Clavaria Zollingeri,
Clavulinopsis corniculata, and Clavulina cristata respectively. Microscopic
analysis shows that this view is superficial and emphasizes the fundamental
difference between Clavaria and Clavulinopsis: it reveals also that there has
been independent specific evolution about the Ramariopsis-spore as about the
Clavulinopsis. In each group there has been variation and loss of pigment,
reduction in size of the fruit-body, and degeneration to the simple state. They
must have had a common origin but whether the original spore was echinulate
or smooth is yet impossible to say.
TYPHULA Fr.
Definition.
I'
Most Clavarioid fruit-bodies with small slender fruit-bodies
have been placed in Typhula or Pistillaria. From these heterogeneous
assemblages one can separate the dimitic Pteruloid derivatives and the small
TYPHULA 145
group of Ceratellopsis. There remain about a hundred species the classifica
tion of which has been purely artificial in these two form-genera. So far as
known, this remainder is a natural group characterized by
1. small size, slender form, and limited growth of the fruit-bodies;
2. the more or less distinct stem and fertile head;
3. the simple hymenium;
4. the 'epiphytic' habitat on wood, stems, leaves, &c., rather than the
terrestrial or humicolous;
5. the smooth white ellipsoid spores, aguttate or with a few minute guttulae;
6. the monomitic, inflating, and generally clamped hyphae;
7. the greater or less agglutination of the hyphae on the surface of the stem,
which gives it the sharp form, and which may extend variously to all
parts of the tissue except the hymenium and subhymenium.
The question is whether the two genera should be maintained or, if not,
how they may be distinguished. Unfortunately most species are still very
inadequately known, so that one can do little more than make suggestions and
such alterations as will help in identification or practical classification.
In 1821 Fries distinguished eight species of Typhula from seven species of
Pistillaria by placing in Typhula those with distinct head and stem and those
with indistinct head and stem in Pistillaria: in other words, the taller fruit
bodies were referred to Typhula, the shorter and less developed to Pistillaria.
Thus:
T. erythropus, T. gyrans, T. phacorrhiza, T. villosa, T. filiformis, T. ramentacea, T.
tenuis, and T. Todei;
P. diaphana, P. micans, P. ovata, P. pusilla, P. quisquiliaris, P. sclerotioides, and P.
muscicola.
This definition has been followed by Patouillard (1887, 1900), Rea (1923),
Bourdot and Galzin (1927), Killermann (1928, 1934), and Konrad and
Maublanc (1937). It is, however, unnatural and impracticable, for closely
related species are thereby separated, as T. candida ( = P. setipes) and P.
pusilla or T. phacorrhiza and P. quisquiliaris, and there are many intermediate
states as T. van·abilis or P. uncialis. Actually, apart from Pteruloid species and
Ceratellopsis, none of which seems to ha�e been known to Fries, it should be
realized that all these fungi have a distinct sterile stem though the pocket-lens
may not reveal it, and so this criterion is useless in theory as well as in practice.
Later, Fries continually emphasized the consistency as waxy in Typhula
and waxy-horny in Pistillaria (ceraceo-corneus, FI. Scan.1835; ceraceo-corneus,
induratus, Epicr. 1838; ceraceo-corneus ...sicca indurata, Hym� Eur. 1874).
In this respect he has not been followed, until recently by Donk (1933). In
practice it is very difficult with such small fruit-bodies to decide between
waxy and waxy-horny: the hymenium is waxy in all cases and the stem more.
or less horny, as in T. erythropus Fr. The distinction is as unsatisfactory in
theory and practice as the earlier one.
In 1882 Karsten emended Typhula to cover only the species with sclerotia,
and he placed in it the following eleven species:
S119 L
T. caricina, T. erythropus, T. graminum, T. gyrans, T. incarnata, T. juncea, T. ovata,
T. phacorrhiza, T. sclerotioides (as T. Friesii), T. Muelleri (as T. sclerotioides), and T.
variabilis.
This definition has recently been accepted by Remsberg, who has studied
in detail more species than anyone and has found the sclerotium a constant
feature in fourteen North American species. I have, therefore, carefully
tested the idea and conclude, also, that the species with sclerotia form a natural
group in contrast with all other Clavarioid fungi in this respect and that the
presence or absence of a sclerotium is the most practicable means, at present,
of dividing the species into Typhula and Pistillaria. In favour of the distinction,
one may remark
I. that many species have never been found without a sclerotium, and that
the sixteen species which have been studied in artificial culture invariably
produce them under appropriate conditions;
2. that many species have never been known to produce sclerotia, e.g. the
common P. setipes, P. pusilla, P. capitata, P. culmigena, P. micans, P.
typhuloides, P. uncialis, &c.;
3. that there are very few species known to occur in the natural state both
with and without sclerotia (see below);
4. that the sclerotium is a peculiar morphological feature not lightly to be
dismissed (see below).
In connexion with the third point, it has been stated that some species may
develop fruit-bodies without sclerotia, but it is difficult to find exact informa
tion. Patouillard says this is the case in T.Patouillardii ( = P. Patouillardii)
and Donk assures me that P. uncialis sensu Maire is the sclerotium-free form
of T. sclerotioides (see also p. 149). Further, there is undo'ubtedly a close
resemblance between some species which become generically separated by
this,means, as T.ovata or T.graminum andP.pusilla or P.setipes, T. incarnata
and P. elegantula, T.phacorrhiza and P.oleae, T.graminum and P. graminicola,
T. Patouillardii or T. erythropus and P. albobrunnea, T. quisquiliaris and P.
Todei. Whether these resemblances are real, or but superficial, remains to
be proved, because the species are so little known in hyphal detail (cf. T.
gramin'um), and it is possible that environmental conditions are not always
suitable for the formation of sclerotia, e.g. high temperatures prevent their
development in T. trifolii. Nevertheless, it is possible that Pistillaria, defined
negatively by the absence of the sclerotium, is unnatural and covers both
sclerotium-f�ee Typhula-derivatives and species which have never evolved
the sclerotium. The doubt does not detract from the positive character of
Typhula, defined by the possession of the sclerotium, and from the practical
point of view one must add that sclerotia are often found rather than fruit
bodies, particularly in diseases caused by species of Typhula, so that patho
logists require a sclerotium-genus and a means of identifying sclerotia as
Remsberg has developed.
Concerning the importance of the sclerotium morphologically, it should be
TYPHULA 14 7
noted that it occurs only in these diminutive Clavarioid fungi and is therefore
an indication of natural affinity. It is, in fact, to be interpreted as an abortive
fruit-body convertible to biologic use because of the peculiar agglutination of
the stem-hyphae of these fruit-bodies, and so rendering Typhula pre-emi
nently a low-temperature genus: (see p. 166).
In contrast with this view Donk has developed an independent classification
of Typhula-Pistillaria. He gives the sclerotium minor importance and
emphasizes the size, form, texture, and habitat of the fruit-body, thus:
Typhula: cylindric filiform fruit-bodies 2-10 cm. high, the stem ill-defined, growing
in humus rather than on particular plant-fragments: with or without sclerotium.
T. phacorrhiza, T. juncea, T. incarnata, T. variabilis.
Pistillaria: distinctly stalked, small (-15 mm. high), the head ovate, clavate, or
fusoid-sublinear: soft and gelatinous, with pelliculose hymenium, drying horny, the
hyphae with gelatinous walls: on dead stems: with or without sclerotium.
P. quisquiliaris, P. uncialis, P. typhuloides, P. Todei.
Cnazonaria: as in Pistillaria, but waxy or rather fleshy, not drying horny, the hyphae
thin-walled: on dead stems, leaves, &c., or on wood: with or without sclerotium.
(a) C. inaequalis (with P. diaphana, P. ovata, P. incarnata, P. micans, P. granulata,
P. sagittaeformis, &c.), P. culmigena.
(b) C. setipes (with Typhula gyrans, T. mucor, &c.).
(c) C. erythropus (with Typhula neglecta, &c.).
(d) Ceratella Pat.
There is no doubt that, in the main, these groups are natural, but I think
they have sectional or subgeneric rather than generic rank. Their definition,
however, is difficult to apply in practice: there are many species, particularly
among those described by Remsberg, which seem to connect Typhula-Pistil
laria sensu Donk and Typhula-Cnazonaria sensu Donk, and for many the
critical points, as the details of consistency, have not been accurately described,
if at all. Moreover, as I will show in detail later, the affinity of P. quisquiliaris
and T. gyrans seems to be with T. phacorrhiza and T. incarnata, though these
four are placed in three separate genera by Donk: similarly P. uncialis is
nearer to P. pusilla and P. setipes than to P. quisquiliaris with which Donk
classifies it: and T. juncea I place in Clavariadelphus with C. fistulosus
( = Clavaria fistulosa). Such differences of opinion, or interpretation, show
how much there is yet to be studied in tl:l.e whole group and that it is too early
to hope for a very precise classification.
It remains to point out that Schroeter's distinction between Typhula as
2-spor:ed and Pistillaria as 4-spored is untenable. There is no such peculiarity
as he found in the case of the 2-spored Clavulina. Basidia have- two or four
spores in the same fruit-body in many species of Typhula and Pistillaria: T.
quisquiliaris has 2-spored and 4-spored states; and T. idahoensis 4-6-8-spored
basidia.
Type-species. Donk (1933) and Remsberg (1940) have independently made
T. phacorrhiza the lecto-type. It is the largest and best-known species of the
genus and in all ways the most suitable. Its fruit-body possesses, according to
Donk, gloeocystidia-like (laticiferous ?) hyphae, and its sclerotium, according
to Remsberg, has a distinctive homogeneously horny construction. Of the
other seven original species of Typhula, two must be transferred to Pistillaria
because they have no sclerotia, namely, T. tenuis and T. Todei; two seem to be
merely sterile states of Clavariadelphus junceus, namely, T. filiformis and
T. ramentacea; T. villosa is an lsaria, according to Killermann; and only
T. gyrans and T. sclerotioides remain with T. phacorrhiza in Typhula emend.
Karst.
Synonyms. Phacorrhiza was made by Persoon for T. sclerotiodes because he
thought the fruit-body developed inside the sclerotium as an Amanita inside
its volva. The primordial shaft is actually erumpent from the medulla of the
sclerotium, which thus shows a minute ruptured orifice round the base of
the stem (p. 163): such may be the usual state in Typhula, but the cortex of the
sclerotium is in general too thin to show a rupture under a pocket-lens..
Greville ( 1823-8) redefined Phacorrhiza for the sclerotioid species of Typhula
and Pistillaria, and he clearly stated that the sclerotium distinguished �he
species as a natural genus. He added T. erythropus and T. phacorrhiza (as
the Phacorrhiza filiformis) to Persoon's genus. Thus, with T. phacorrhiza as
the lecto-type of Typhula, Phacorrhiza becomes a synonym of Typhula Fr.
emend. Karst. To Greville, however, must be given the credit of first
discerning the importance of the sclerotium.
STRUCTURE AND HABITAT

Before discussing structural details I give a nominal description to show
the variation of the general construction on the record of which specific
accuracy depends: the list is useful for reference, because it is easy to overlook
points in the moment of discovery:
Fruit-body: height, number per sclerotium, simple or branched, fertile or sterile
(in culture, particularly).
Head: size, shape, colour, apex (fertile or sterile).
Stem: size, shape, colour, direction, origin from sclerotium.
Texture: in stem, in head: (doubtful value).
Habitat: host, locality, date.
Spores: variation in size, spore-contents.
Basidia: size, contents, number of sterigmata, clamped or not.
Cystidia: size, shape, contents.
Hymeniu�: subhymenium.
Caulocystidia.
Hyphae: width, length of cells, thickness and modification of walls, clamps, in-
crustatioq,
Stem-structure: agglutination of hyphae.
Head-structure: inflation of hyphae, cortex, growing apex.
Sclerotium: size, colour, shape, surface-appearance, alteration with age, situation
in host, coalescence (in culture), microscopic structure (section and surface-view).
Stromatic crusts (in culture).
Colour: microscopic basis.
Oidia: on haploid mycelia (in culture).
Fruit-body. To appreciate the proportions of the fruit-body it is necessary
TYPHULA 149
to know the total height as well as the size of the head and stem. There may be
a proportional relation between the length of the head and the stem or the
length of the stem may depend on a phototropic stimulus rather than any
inherent property, in which case the height of the fruit-body will be variable,
according to external conditions, rather than specific.
Generally there is one fruit-body per sclerotium, but there are the following
exceptions:
T. caespitosa: up to 20 per sclerotium.
T. athyrii, T. gyrans, T. idahoensis, T. intermedia, T. latissima, T. nivea, T. phacor
rhiza, T. subulata, T. umbrina, T. variabilis: I-several per sclerotium.
T. erythropus, T. incarnata, T. Patouillardii: occasionally 2-3 per sclerotium.
Whether the number depends on the size of the sclerotium or whether it is

a specific peculiarity, in which case smaller sclerotia will have attenuate fruit
bodies, is not known.
In some species there are occasionally produced stolons, which appear to be
decumbent stems which have not yet become phototropic or geotropic. In
T. gyrans a stolon may grow from the sclerotium for several millimetres before
its apex ascends to form the true stem and head of the fruit-body. In T.
idahoensis stolons may grow from the mycelium in culture and produce fruit
bodies without sclerotia. They are not a regular feature of any species.
Fruit-bodies occasionally develop from the mycelium, without a sclerotium
intervening. Remsberg mentions fruit-bodies of T. idahoensis and T. pertenuis
developed from the mycelium in nature, and other authors mention in passing
'sclerotium-free specimens' of Typhulas, but such are exceptional and need
more careful description (see p. r46). It seems that in culture fruit-bodies are
more commonly developed from the mycelium. Thus, according to Remsberg: ·
Fertile fruit-bodies, in culture, from the mycelium or stromatic crust: T. idahoensis,
T. pertenuis, T. sphaeroidea, T. subulata, T. virgata: T. trifolii (haploid fruit-bodies:
diploid from sclerotia: teste Noble).
Sterile fruit-bodies, in culture, from the mycclium: T. intermedia, T. idahoensis,
T. sphaeroidea, T. subulata, T. viburni.
The sterile fruit-bodies have not been found in nature and it is not stated
whether they have a normal hymenium \, ith sterile basidia or they lack a
culture they seem to be formed at suboptimal temperatures, e.g. at 6-1 s" C. in

hymenium entirely (cf. sterile fruit-bodies of Ceratellopsis aculeata). In
T. intermedia (opt. temp. 12-15 ° C.) and at 9-12° C. in T. sphaeroidea (opt.

temp. 12-18° C.), and these examples suggest that the nuclear prvcesses in the
basidium may be conditioned narrowly by temperature.
Branched fruit-bodies have been reported in the following species, but the
branching is probably spurious or adventitious:
T. athyrii, T. caespitosa var. coacervata, T. corallina, T. intermedia (sterile fruit
bodies in culture at suboptimal temp.), T. ramealis, T. srlerntioides, T. variabilis.
Head. The size of the head appears to be the most constant macroscopic
·;;It"'!. ·:!"H .;!..I.....:
:.-: ::· :::·.�

TEXT· FIG. 46. Typhula sclerotioides: 1.s. of part of the fertile head, x I ,ooo.
TYPHULA
measure of the fruit-body, yet it is often omitted. The width must be measured
microscopically.
The shape of the head is determined by four processes, the first two deter-
mining mainly the length, the second two the width:
r. apical growth;
2. inflation of the hyphae;
3. subhymenial development;
4. intercalary growth of the hymenium.
The interaction of the four processes may be complicated and, in varying
degree, they may produce the same result. The details of the processes are
described later, but I will outline them to show how they affect the shape.
The head is the region of the axis where the hymenial factor develops.
Assuming apical growth and this factor to be the only conditions, the following
two extremes (a, b) and their mean (c) can be distinguished:
(a) prolonged apical growth, with attenuation, the hymenial factor rarely
reaching the apex, which remains sterile: head elongate, cylindric or
filiform, attenuate to the acute sterile tip: e.g. T. phacorrhiza, T. betae,
T. incarnata, T. intermedia, T. pertenuis, T. subulata, T. variabilis,
(b) curtailed apical growth, no attenuation, the hymenial factor affecting
the whole apex: head subglobose or shortly clavate, wholiy fertile,
blunt: e.g. T. abietina, T. caricina, T. euphorbiae, T. graminum, T. gyrans
(occasional), T. ovata, T. quisquiliaris, T. sclerotioides ( occasional),
T. sphaeroidea, T. viburni;
(c) intermediate state, the apex eventually fertile: head subcylindric or
cylindric-clavate, blunt, scarcely attenuate, wholly fertile: e.g. T.
erythropus, T. virgata (and most other species).
These, I think, are the main causes of the different shapes in Typhula-heads,
but other possibilities can easily be seen and, in fact, each species requires
analysis. Thus curtailed apical growth with much elongations of the cells and
little subhymenial development may produce a cylindric head (as in Chaeto
typhula): or much intercalary growth of the hymenium with little elongation
of the cells may produce a subglobose or clavate head in spite of considerable
apical growth. Until the species have been analysed there will be uncertainty
in judging affinity from the shape of the head.
The presence of a sterile apex to the head is emphasized in descriptions, for
instance, of T. phacorrhiza and T. variabilis. It has not been realized that the
sterile apex is the growing-point of the axis and that it is present in -every species
of Typhula, as in every other Clavaroid fruit-body, until the cessation of
apical growth, when the tip may or may not become fertile according to its
state of attenuation. Thus, I think it has yet to be proved that the species in
group (a) (above) do not eventually, and normally, have fertile tips when fully
grown.
Stem. The stem is the part of the axis developed before the appearance of
the hymenial factor. Its length depends on the extent of apical growth and on
the subsequent elongation of the cells, and needs such analysis before differ
ences can be compared. The length may also be controlled by some photic
TExT-FIG. 47. Typhula gyrans: l.s. of the head on cessation

of up growth, X I ,ooo.
stimulus. It is doubtful if much specific value can be attached to the length,

but the width is more constant. It indicates mainly the number of concrescent
hyphae in the primordial shaft, and, secondly, the enlargement of the hyphae.
Some species have rather stout sterns, as T. gyrans, others slender sterns, as
TYPHULA 153
T. sclerotioides and T. erythropus. But before differences in width can be
compared the lateral enlargement of the cells must be known.
The direction of the stem is perpendicularly upright in most species.
Whether the stem is positively phototropic and negatively geotropic is not
known. In T. intermedia the stem appears obliquely erect or decumbent,
perhaps because it is weak and sags under the weight of the long head.
It seems that the stem always arises more or less endogenously from the
sclerotium and that the growing-point of the primordial shaft is formed from
the medullary hyphae of the sclerotium immediately within the cortex. Thus
Persoon first described the endogenous origin of T. sclerotioides but imagined
that the fruit-body was formed entirely in the sclerotium as an Amanita in its
volva. Then Brefeld observed that the fruit-bodies of T. variabilis arose from
the tissue just beneath the cortex of the sclerotium and that those of T. gyrans
were deeply endogenous and made a fairly large split where they had broken
through the cortex. In T. sclerotioides I find that the fruit-bodies arise from
the medullary hyphae immediately internal to its rather wide agglutinated
cortex and are microscopically erumpent (Text-Fig. 51). I think the position
of the primordia may be indicated in the development of the sclerotium.
Texture. This is a fond character of the macroscopic systematy of fungi,
but it has little application in such small fruit-bodies as those of Typhula.
Any differences in texture in the genus depend primarily on the presence or
absence or the relative degree of agglutination of the hyphae, and are therefore
more easily and more precisely expressed microscopically. Waxy, horny, and
cartilaginous are the usual terms. Cartilaginous means with agglutinated
hyphae, as the sclerotium of T. phacorrhiza. Horny probably means less
toughly agglutinated. Waxy means with inflated, closely applied, thin-walled,
and not agglutinated hyphae, the walls being wet and therefore not fibriIIar:
the hymenium and subhymenium are characteristically waxy. Therefore the
stem is always cartilaginous in some degree, because of the peripheral cylinder
of narrow, agglutinated hyphae: but in T. phacorrhiza, T. gyrans, and the
other species which I have placed in the subgenus Phacorrhizae, it is whoily
cartilaginous, because aII the hyphae are somewhat fim1ly agglutinated. The
head, in contrast, is generaIIy waxy : its core, or meduIIa, of longitudinal hyphae
may be more or less horny or cartilaginous according to the modification of
the hyphal walls. The difference may be useful as field-characters in practised
hand.s. The texture is never coriaceous-fibriIIose as in Pterula and Ceratel
lopsis.
Habitat. There is much to be learnt about this fascinating .iubject, but I
think there is no such specialization as the records suggest. So far as known,
Typhulas never develop an extensive mycelium in humus, soil, or wood. Their
mycelia appear restricted to smaII parts of plants as the faIIen leaves, twigs,
petioles, herbaceous stems, inflorescences, fruits, or- pieces of bark: yet a
confined mycelium is not so characteristic as to be the generic distinction
which Donk has suggested, because the parasitic species develop extensive
mycelia when they have invaded their hosts. There are a few records of
r·!
t I :i i !/
/, \..�·
1ii ··
Jt
I�
11: ·
I \,
I .' fii
(i
I
''
�'.
l. l !\'
I
I:
, I
I
:1
I ·1 \
I I ·
I,
1
J
/1 ./
I.
I
I·
I
:·I)
I
.\
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'1
\
'I,
I :1
-� : I:
,I'
·'
J:I
I1,,.:
I l
/···,,
i !
I
TEXT-FIG. 48. Typhula phacorrhiza: l.s. of the hymenium and outer part of the
flesh (the subhymenial crystals omitted for sake of clearness): X 1, 000.
TYPHULA 155
fruit-bodies occuring on the ground or in the soil, but they may have grown
from sclerotia which have fallen from the decaying host and have remained
dormant for some months: thus:
T. cystidiophora, T.jlavescens, T. idahoensis, T. neglecta, T. pertenuis, T. phacorrhiza,
T. variabilis.
There seem to be only two corticolous species, T. caespitosa var. coacervata
and T. ramealis, and two species on woody twigs, T. abietina ( on Abies and Pinus)
and T. erythropus (generally on leaves or herbaceous remains).
The great majority grow saprophytically on herbaceous remains; a few are
facultative parasites. I have listed the records in this way:
SAPROPHYTES
Habitat varied (on leaves, stems, roots, &c., of various herbaceous plants or her
baceous remains of woody plants):
T. erythropus, T. gyrans, T. incarnata, T. intermedia, T. patouillardii, T. phacorrhiza,
T. sclerotioides, T. subulata, T. umbrina, T. variabilis, T. virgata.
Habitat limited (on the remains of one family, genus, or species of plants, so far as
known).
On ferns: T. athyrii (on Athyrium angustum), T. quisquiliaris (on Pteris aquilina).
On grasses: T. idahoensis, T. graminum (T. incarnata).
On Cyperaceae: T. caricina (on Carex vesicaria).
On Typha latifolia : T. latissima.
On Populus: T. ovata.
On Compositae: T. bulbosa (on Eupatorium), T. neglecta (on Helianthus tuberosus,
T. Patouillardii (on Carduus).
On Viburnum: T. elegans (on V. opulus), T. viburni (on V. cassinoides).
On Rubus: T. Laschii, T. sphaeroidea.
On Hedera: T. hederaecola.
On Euphorbia: T. euphorbiae.
On Gentiana: T. Muelleri.
On Mentha: T. corallina, T. nivea.
On Buxus sempervirens: T. buxi.
On Brassica oleracea: T. tochinaiana.
On Olea europea : T. pallens.
On Juglans: T. caespitosa.
On Fraxinus: T. crassipes, T. lutescens.
FACULTATIVE PARASITES
T. betae (on Beta, ? also on Solanum tuberosum).
T. borealis ( on various grasses, cereals, and cruciferous weeds: ? = T. idahoensis).
T. idahoensis (on Triticum vulgare and Agropyrum cristatum).
T. incarnata ( = T. ltoana) (on various grasses and cereals).
T. trifolii (on Trifolium, Anthyllis, Medicago).
T. umbrina (on Brassica napa and Iris).
T. variabilis (on Apium graveolens).
The host-limitation must, however, be queried. The species which are
well known are omnivorous: the restrictions are probably only apparent
because of the paucity of collections. In the case of T. incarnata ( = T.
ltoana), Imai has proved that there is no biological specialization as occurs
among obligate parasites.
The parasitic habit has evidently arisen from the saprophytic. The parasites
are merely facultative and they attack their hosts mainly through injuries
caused by frost, water-logging, or other adverse conditions. The parasites
have lower temperature-ranges and lower optima than the saprophytes, so far
as known, and will grow readily at 0-4° C., when they clearly have the
advantage over their dormant hosts. Entry is through underground parts,
either by hyphae growing saprophytically on dead or moribund tissue, or in
some cases, as T. incarnata, by hyphae which grow directly from dormant
sclerotia into the living tissues of seedlings or young parts. If conditions are
favourable for the parasite it will permeate and kill the underground parts of
the host in two or three weeks. Imai has called the diseases inflicted by
Typhula by the general name Typhula-blights. Their peculiarity is the low
temperature rotting of the host near or under the ground, followed by
superficial white mycelium and sclerotia on the dead tissues. Indeed, it seems
that Typhula-blights are generally winter diseases, and thus absent from the
tropics and sub-tropics.
Spores. Specific descriptions suggest that the spore-size is diagnostic, but
I am doubtful. There appear to be two groups, namely, a large-spored
represented by T. phacorrhiza (spores over 9 µ, long) and a small-spored
represented by T. erythropus or T. sclerotioides (spores under 9 µ, long).
Among the large-spored species the variation in T. phacorrhiza alone (see
Table III) covers almost the entire range of spore-variation in the others, e.g.
T. variabilis, T. quisquiliaris, T. intermedia, T. sphaeroidea, T. subulata,
T. latissima, and T. virgata. In the small-spored group both T. erythropus
(5-II X 2·5-4 µ,) and T. sclerotioides (5-9·5 X 2·5-4 µ,) indicate a similar latitude.
More precise data are required before small differences in spore-size can be
accepted as diagnostic.
The large spores are often rather amygdaliform, as if they passed through
a short broad shape during development. The small spores are oblong
ellipsoid or subcylindric and clearly elongate from an early stage in develop
ment. The smallest spores, if correctly recorded, occur in T. ramealis (3 X 1 µ,).
The spore-contents are generally aguttate or with 1-2 very small guttulae,
but exceptions are said to occur in T. cystidiophora and T. intermedia with
large oil-drops.
Basidia. What little variation there is in size is probably connected with the
variation in spore-size. The smallest occur, as far as is known, in T. buxi, with
small spores and basidia 15-18x4-6 µ,. The largest, or widest, occur in
T. phacorrhiza (20-33 X 9-II µ,), T. quisquiliaris (30-40 X 7--9 µ,), and T.
sphaeroidea (20-45 X 9·5-12 µ,), all of which have large spores. Most records
are about 25-30 X 6-8 µ,. Generally there are 4 sterigmata, 5 µ, long, but only
2 sterigmata are reported in T. corallina, T. sclerotioides, and T. semen sensu
Bourd. et Galz. (see T. variabilis). In T. neglecta and T. erythropus 2-3-4
sterigmata have been reported and in T. idahoensis there are 4-6-8 sterigm ata.
In T. intermedia the sterigmata are 7-10 µ, long according to Appel and Hau
bert, and in T. phacnrrhiza I found them 6-8 µ, long. In T. sphaeroidea
TYPHULA 157
(Text-Fig. 49) the basidia and subhymenial hyphae appear to fragment at
the septa like the mealy powder of the velar hyphae in some species of Lepiota,
Coprinus, and so on. In T. cystidiophora Kauffmann has described the basidia
as unusually long and narrow, 45-50 X 4-5 µ., but I think there may be an
error, unless this species does not belong to Typhula.
TABLE III. SPORE-SIZE IN TYPHULA PHACORRHIZA

(One fruit-body: Wicken Fen, England)
No. of No. of
Size in /1, E* records Size in µ, E* records
IIX 5·0 2'2 I 14·5X6·5 2'2 2
l l X6·5 1·7 l 14·5x6·7 2'2 l
11·5X4·7 2·4 l 15X 5'3 2·8 2
12X4·7 2·6 l 15x5·5 2·7 l
12X 5·0 2·4 2 15X 6·o 2·5 2
12X 5'3 2·3 l 15x6·5 2·3 l
12x6·0 2'0 3 15·5X6·5 2·4 2
12x6·5 1·8 l 15·5X7·0 2'2 l
12x7·5 1·6 l 15·5x7·5 2·1 l
12·3X 7·5 1·6 l 16x6·o 2·6 2
13 x5·0 2·6 2 16X6·5 2·5 l
13X5·3 2·5 l 16x6·7 2·4 3
13X7·0 1·9 l 16x7·0 2·3 2
13·5X6·o 2·3 l 16·5X7·3 2·3 l
13·5X6·5 2·1 3 16·5X 7·5 2'2 l
13·5X7·0 2·0 2 17X6·5 2·6 l
14 X6·o 2·3 3 17x7·0 2·4 4
14X6·5 2·2 I 17X 7·5 2·3 2
14X 7·0 2·0 l 17·5x7·0 2·5 2
14·5X 5·0 2·9 l 18x7·5 2·4 l
14·5X 5·7 2·5 l 2ox7·0 2·9 l
Total range of 64 spores, 11--20X4·7-7·5 µ,, E* 1·6-2·g

Average of 3 spores 11-u·5 µ, long, E 2·1
IQ 12-12·5 µ, ,, E 2'1
IO 13-13·5 µ, " E 2·4
IO 14-14·5 µ, " E 2·3
IO 15-15·5 µ, " E 2·5
10 16-16·5 µ, " E 2·4
10 17-18 µ, " E 2·4
1 spore 20 µ, ,, E 2·9
*E = length-:-width
Cystidia. Only in T. cystidiophora have they been reportecl and, as just

mentioned under the basidia, I think this species requires more critical study
whether or not it really belongs to the genus.
Hymenium. In all species the head is abruptly delimited from the stem and,
at most, only a few scattered sterile basidia intervene. The hymenial factor
develops abruptly and, in species with limited apical growth, it quickly trans
forms the whole head into the hymenium (Text-Fig. 47). In some species the
out-turned hyphal ends immediately become basidia, e.g. T. phacorrhiza, but
in most, perhaps, they grow out for a short distance and cut off 1 -several short
subhymenial cells, e.g. T. gyrans, and T. sclerotioides (Text-Figs. 46, 47).
The extent of the subhymenium varies and largely contributes to the shape and
width of the head. Thus in all species with limited apical growth the sub
hymenium appears to be well developed and to give the clavate or SJ.lbclavate
head. The hymenium is always restricted to a single layer of basidia.
Caulocystidia. Though several species have been described with glabrous
stems, I am not sure that microscopically this is true. In most, certainly, there
fD
are caulocystidia scattered or crowded,
particularly towards the base of the stem
�- and varying from 10-20 µ, long to 200 µ,
and visible as fine hairs to the naked. eye.
They are generally conical to filiform
with thin or slightly thickened smooth
walls, and they may not offer any
striking specific differences. As in the
genera Pterula and Dimorphocystis,
they are to be regarded as the outcome
of the rudimentary working of the hy
� menial factor in the first phase of the
fruit-body, and their elongation at the
oj.., base of the stem is connected with the
tendency to develop a secondary myce
TExT-FIG. 49. Fragmenting subhymenial lium from excrescent hyphae.
hyphae of Typhula sphaeroidea (top left);
4-8 spored basidia of T. idahoensis (top Hyphae. The hyphal tips at the
right): oidia of T. idahoensis: x 300 (from growing-point of the fruit-body are
Remsberg, 1 94 1),
1 · 5-3 µ, wide, and the cells which ·they
cut off inflate directly without secondary septation. Even at the growing-point
the hyphal walls appear to be slightly mucilaginous and the thin-walled hyphal
tips are not therefore as exposed as in the Pteruloid fungi. Behind the growing
point the walls become more mucilaginous and, in the subgenus Phacorrhiza,
more or less agglutinated: the mucilage, however, is always firm and waxy
horny, rather than soft and diffuent as in truly gelatinous fungi. The modifi
cation of the mature walls appears to be characteristic of the species. Thus,
in T. phacorrhiza and T. gyrans, not only are the hyphae of the fruit-booy
more or less agglutinated but also those of the sclerotium and of the mature
mycelium, which becomes in culture 'a tough cartilaginous layer difficult to
cut or break' {Remsberg).
In many species the medullary hyphae of the sclerotiurn become incrusted
with small crystals (Text-Fig. 51), and crystals are often deposited among the
outer hyphae of the stem and head of the fruit-body. The crystals may be
colourless or pale brown as in T. phacorrhiza. They appear to be the residue
of an exudate which evaporates from the growing tissue. Thus, during the
development of the sclerotia, Brefeld and Remsberg have noted_ that drops of
water appear on the surfaces of the sclerotia, and Remsberg found that on
TYPHULA 15 9
drying the drops on a glass slide they left a crystalline deposit. The incrusta
tion of the hyphae seems characteristic of the genus, though Maire distin
guishes T. buxi, among other ways, from the closely allied T. pa/lens by the
absence or paucity of crystals on the stem-hyphae of the former. At least,
the incrustation should not be overlooked in specific descriptions. Oddly
enough the caulocystidia do not become encrusted and appear to take no part
in excretion water as they generally do in other fungi.
Excepting haploid individuals, clamps seem to occur on the mycelial hyphae
of all species, but they are not always present in the fruit-body. Indeed, it
is often not easy to detect whether they occur in the fruit-body, and its hyphae
must be teased apart, so far as agglutination allows, before the point can be
decided. There are the following records:
T. quisquiliaris, T. variabilis, T. buxi, apparently with clamps at most if
not all septa in the fruit-body.
T. erythropus, clamps absent from many septa in the fruit-body, but
many clamped and the basidia mostly clamped.
T. gyrans, T. phacorrhiza (? also T. intermedia and T. sphaeroidea),
without clamps in the fruit-body except a few on some of the hyphae in the
stem, the basidia without clamps.
T. trifolii and T. graminum, without any clamps in the fruit-body: T.
trifolii with clamps in the mycelium.
T. sclerotioides, some fruit-bodies with clamps, others without.
Stem-structure. The hyphae are longitudinal and not interwoven except
at the extreme base. The outer hyphae are narrow, their cells being elongate
but little widened, and their walls become firmly agglutinated to form the
horny or cartilaginous surface characteristic of the genus. In some species,
the agglutinated walls become brown or reddish like the outer surface of the
sclerotium, e.g. T. erythropus. In the subgenus Phacorrhiza the agglutination
. extends to the centre of the stem so that its hyphae are separable with diffi
culty. The pellucid appearance is caused by the tissue being saturated with
water and devoid of air-spaces.
Head-structure. If the stem offers little variation in structure, there are
considerable differences in the head which require careful analysis before
the true affinity of the species can be decided. In all there is a core or medulla
of longitudinal hyphae which is continuous and, at least initially, of equal
diameter with the stem. On it, in T. phacorrhiza, the basidia are borne
directly from the out-turned tips of the superficial hyphae. In �ost species,
however, as already described, a subhymenium of varying thickness develops.
It may be slight as in T. erythropus (cf. Text-Fig. 289), or its inner cells
joining with the medulla may be so inflated and numerous as to appear like an
algal cortex surrounding the axial strand, e.g. T. gyrans and T. sclerotioides
(Text-Figs. 46, 47). In most species the head is dilated by intercalary growth
of the hymenium and in T. variabilis it becomes hollow. The sharp distinction
between the head and the stem, so characteristic of Typhula, is caused by the
sudden development of the hymenial factor and by the appreciable thickness
of the hymenium and subhymenium compared with the axis of longitudinal
hyphae: in the stouter Clavarias, which may have a sterile stem as long or
longer than in Typhula, the thickness of the hymenium and subhymenium is
generally insignificant compared with the width of the stem, e.g. Clavaria
delphus pistillaris or Clavulinopsis fusiformis.
In the sterile fruit-bodies developed in culture it has not been recorded
whether the head is covered by a hymenium of sterile basidia or whether
there is no appearance of the hymenial factor, so that the fruit-body consists
merely of the axis of longitudinal hyphae (cf. Text-Fig. 65, for Ceratellopsis
aculeata).
Conidia. Beside the haploid oidia, T. bulbosa produces from the sclerotium
minute, discoid, conidial stromata, as in a few species of Pistillaria and Cera
tellopsis (P. micans, C. rosella).
Sclerotium. For practical identification, because the sclerotia are found
more often than the fruit-bodies, and for the systematist, because they give
an extra diagnostic character, the structure of the sclerotium is important.
After the early researches of de Bary and Brefeld, little has been added until
Remsberg's work in which it is shown that the sclerotia have specific features
and that, by them alone, at least fourteen North American species of Typhula
can be distinguished. I find the terminology not entirely appropriate and
difficult to remember, so that I have changed it to simpler expressions. The
use of the three seven-syllable words, prosoplectenchymatous, paraplecten
chymatous, and pseudoparenchymatous suggests three tissues profoundly
different; actually they are just stages in the agglutination of a plectenchyma
and grade imperceptibly into each other, and the final state is not pseudo
parenchymatous in the sense of a turgid tissue composed of filaments of
inflated iso-diametric cells, like the pseudoparenchymatous parts of Gas
tromycetes, Hymenomycetes, Discomycetes, and multifilamentous algae: it
is merely an agglutinated mass of compactly interwoven hyphae with con�
tiguous and not always distinguishable walls, very different from the crisp
tissue of the- Phalloid stem or Pezizoid cortex.
In the sclerotium four parts can be distinguished (Text-Fig. 50). There is
the medulla, composed in the simpler cases of loosely interwoven hyphae
which form the central mass and storage tissue of the sclerotium. On the
outside is a cortex of agglutinated hyphae, the outer walls of which may be
thickened and deeply coloured to form a cuticle. Then, in some species, in
stead .of a cuticle there is a superficial layer of narrow hyphae which can be
called the epidermis because it does not seem homologous with the cortex and,
until it has been explained developmentally, it must be distinguished by
another name. These terms are plainly descriptive, in the same sense as
pileus, stipes, pellicle, hymenium, velum, and so on are used in mycology.
Now, according to the development of the four parts, four kinds of sclerotium
can be distinguished and two of these can be subdivided to give six kinds in
all. Thus:
TYPHULA 161
I. M!idullary hyphae free: cortex composed of a single layer of agglutinated hyphae
with cuticle.
(a) Cortical hyphal walls next the medulla but slightly thickened or not at all, the
lumena of the cortical hyphae appearing large compared with those of the thick
walled medullary hyphae (Text-Fig. 50B), or the medullary hyphae also thin
walled (T. graminum).
T. erythropus, T. graminum, T. pertenuis, T. umbrina, T. 'Jariabilis, T. virgata.
A C
CUT
COR
Mto
r
TEXT-FIG. 50. Diagrammatic sections of the surfaces of Typhula-sclerotia: cut = cuticle,
cor = cortex, med = medulla: X c. I ,ooo.
(b) Cortical hyphal walls thickened all round, the lumina of the cortical hyphae
narrow like those of the thick-walled medullary hyphae (Text-Fig. 50A).
T. athyrii, T. idahoensis, T. intermedia, T. latissima, T. subulata.
2. Medullary hyphae free: cortex composed of several layers of agglutinated hyphae,
with cuticle (Text-Fig. 50c).
T. sclerotioides.
3. Medulla wholly agglutinated, indistinguishable from the cortex, with cuticle
(Text-Fig. 500),
T. incarnata, T. phacorrhiza (? T. euphorbiae, T. quisquiliaris, here or in the next
group).
4. Medulla wholly agglutinated, indistinguishable from the cortex, without cuticle
but with epidermis.
5119
M.
(a) Epidermal hyphae thick-walled but not agglutinated, distinguishable (Text
Fig. 50E).
T. gyrans (hyphal outlines invisible in the medulla), T. viburni (hyphal outlines
visible in the medulla).
(b) Epidermal hyphae agglutinated, indistinguishable (Text-Fig. 50F).
T. sphae-roidea.
The first three groups show progressive states in the agglutination of the
sclerotial hyphae, from without inwards, the third (and fourth) groups being
wholly agglutinated though the tissue in the very centre may be more or less
loose. The cortex, therefore, is merely the outer agglutinated layer and not
an outgrowth or investment like the cortex of a multifilamentous soma or
the hymenium and subhymenium of the Typhula-head. The cuticle, on the
other hand, is composed of thickened, perhaps specially secreted, hyphal wall
which becomes characteristically coloured, perhaps through oxidation, so that
it is darker at the surface and gradually paler inwards. 1 During the agglutina
tion of the hyphae they gradually lose their identity as the contacting walls
merge: in T. viburni the hyphal outlines are apparently always distinct, but in
T. gyrans, T. phacorrhiza, T. incarnata, and T. sphaeroidea they disappear,
and the lumina appear embedded in a matrix which is homogeneous save for
the darkened cuticle. The same slight difference in the epidermis appears to
separate T. sphaeroidea from T. gyrans and T. viburni in the fourth group,
but I am uncertain about the exact nature of this epidermis. As a further
complication, perhaps also specific, the cuticle may be smooth or somehow
marked with irregular projections. In T. intermedia it is marked with small
rosettes of tubercles, but whether these tubercles are merely cuticular thick
enings or hyphal processes containing cytoplasm, I Jo not know.
Whether these differences in sclerotial structure are distinguishable in all
species of the genus further research will prove, but I think intermediate
states will be found.
The surface of the sclerotium also varies microscopically in the size and
shape of the cell-cavities and in the thickness and waviness of the radial walls.
Each species has no doubt its specific pattern, like the epidermis .of many
higher plants, but the irregularity of the features makes it difficult to describe.
Remsberg has given drawings of nine species which I have copied in Text
Fig. 286, and I have added those of T. erythropus and T. sclerotioides. For
sclerotial identification, plates showing the patterns in different species may
be sufficient.
The development of the sclerotium appears simple, but it is difficult to
understand how all the details arise and it is to be hoped that full microscopical
drawings will be published. ' A tuft of much branched hyphae arises from the
mycelium in the substratum and becomes a ball of enlarged hyphal cells. It
increases in size by repeated branching of the hyphae as they become inter
twined. The peripheral cells become fused early, and consequently are much
1
In Remsberg's paper, cuticle and cortex are called cortex or rind without distinc,_,1n. I
COrisider the cuticle the non-cellular outer layer of the cortex and composed of more or :,-ss
confluent hyphal walls.
TEXT-FIG. 5 r. Base of the stem of Typhula sclerotioides at its origin from the
sclerotium: X 500.
distorted as the sclerotium increases in size' (Remsberg). This general
description must be translated into terms of hyphal growth. I can never
understand how a ball of hyphae is formed. Either it arises from a cone or
cylinder of more or less parallel, but i_ntertwined, hyphal tips which through
profuse branching and out-turning form a subglobose mass covered with
contiguous and excrescent hyphal tips, as in Text-Fig. 47 (but of this there is
no sign in the structure of the mature sclerotium): or, the hyphae must be
simply concrescent without form-factors and merely grow over each other
repeatedly and in any direction: then, some inflation of the older cells will
produce small gaps between the hyphae into which new branches can push
to confuse still further the manner of growth. If this is the method and the
outer hyphae agglutinate early into a tough rind, their cells must either en
large to accommodate the growth of the internal hyphae or, as seems to be
the explanation, they become pulled out into lobes to give the same 'reticulate'
appearance as the epidermal cells of Dicotyledonous leaves. Compact con
crescence from the start will give a dense structure of contiguous hyphae
through which agglutination can readily proceed (? T. phacorrhiza): loose con
crescence, representing still further degeneration of a somatic tissue, will
give the sclerotium with loose medulla and narrowly agglutinated cortex,
e.g. T. variabilis. It is not until one inquires what the hyphae are doing that
one can understand why an organ has its ultimate structure.
In the species with an epidermis it 'is made up of hyphae wound about
the medulla. The walls of these hyphae become thickened and darkened.
The hyphae are agglutinated but often sluff away from the surface as the
sclerotium enlarges, until but a layer or two is left' (Remsberg). What are
they doing on the surface? Are they subsequent hyphae which through
concrescence invest the original tuft? And how do they come to be regularly
layered, as if they had grown in superposed planes? Thus it seems, as I have
mentioned, that the epidermis differs in development as well as in final
structure from the cortex.
Sclerotia may alter shape somewhat in nature, either through drying and
shrinking or from some other effect of maturity. Thus old dark sclerotia are
generally more or less flattened, the young pale sclerotia more or less globose
or nodular. During their maturation, the sclerotia excrete drops of water
from the surface. In T. sclerotioides, Macdonald found that they took 9-10
days to reach this stage from inception.
The colour of the sclerotium resides in the cuticle, or thickened wall of the
outer layer �f agglutinated hyphae. It develops gradually, and from white
the sclerotia slowly become pale yellow, ochraceous, tawny, or rich brown to
black. The process may take several months as Quelet found in T. semen
( = T. variabilis). The coloration of the wall is like that of the agglutinated
crust of Polypores and Sterea, and its slow darkening from the surface inwards
suggests oxidation of a pigment. The chemistry may well be identical in all
cases. De Bary (T. gyrans) and Macdonald (T. sclerotioides) note that the
rind of the sclerotium cannot be regenerated on cut surfaces.
TYPHULA 165
In culture the sclerotia of some species may coalesce. Remsberg found the
tendency to be characteristic of certain species and it is used in her specific
key to the identification of Typhulas in culture. I am not aware that coale
scence occurs in nature, the natural substrata probably being insufficiently
rich to allow the production of so many sclerotia. Thus, at low temperatures
the sde.rotia grow in piles or masses, but at high temperatures they are
scattered, and when they are heaped the primordia of adjacent sderotia may
. become enclosed in the same rind or more or less fused in masses. Lobed
sclerotia in nature may arise from this cause or merely from the angularity of
their host, cf. T. abietina.
Sclerotia are either superficial or, more commonly, immersed in the tissue
of the host. In some species the immersed sclerotia are erumpent, doubtless
because the dead host-tissues break away readily, but in other species the
host tissues are firmer and the sclerotia permanently embedded. Thus, the
sclerotia of T. quisquiliaris are firmly embedded in bracken petioles and were
never seen by Fries, Karsten, Quelet,. or Patouillard: Tulasne appears to
have been the first to discover them.
Stromatic crusts. These are formed in culture on the surface of the medium
at or near the maximum temperature for growth. They seem not to have been
observed in nature.
Colour. Little is known of the microscopic basis of the colour of the fruit
body. In species with brown or reddish stems, the colour is probably in the
walls of the agglutinated superficial hyphae and the same as the sclerotial
colour, e.g. T. erythropus and haploid T. trifolii. The brownish ochraceous
colour of T; phacorrhiza may be caused by the brownish crystals among the
longitudinal hyphae. The delicate pink qf T. incarnata is surely not caused
in the same way as the pinkish stems of T. erythropus, but is more likely to
be cytoplasmic.
NUCLEAR BEHAVIOUR AND SEXUALITY

On these subjects there is yet little information. Most species are clearly
diploid with clamp-connexions, at least in the mycelium, and it seems that
most are heterothallic and produc� oidia on their initial haploid mycelia. Thus
T. trifolii was found to b.e heterothallic, but it could not be decided whether
it was bipolar or tetrapolar (Noble). It produces haploid oidia, and similar
oidia (presumably haploid) have been reported for T. gyrans (Brefeld), T.
incarnata ( = T. Itoana) (Remsberg), and T. variabilis (Brefeld, Remsberg).
T. sclerotioides is also heterothallic according to Macdonald, but oidia were
not seen. In T. trifolii and T. sclerotioides the fusion-nucleus divides in the
basidium into 4 daughter nuclei, one entering each spore. In T. idahoensis
with 4-8 spores per basidium, there must be more nuclear divisions in the
basidium, giving 4-8 daughter nuclei. For species with 2-spored basidia one
cannot even guess what the nuclear behaviour may be.
T. trifolii produces in culture haploid mycelia and haploid fruit-bodies..
The haploid basidiospores are only half the size of the diploid and-they arc all
of one 'sex'. They are borne on 4-spored basidia, but it is not recorded
whether there is a fusion-nucleus in the basidium, though it seems likely (the
fusion of 2 nuclei in the basidia of haploid fruit-bodies has been found, most
surprisingly, in Coprinus lagopus, Peniophora ludovicianus, Mycena clavicularis,
M. graveolens, M. immaculata, M. margaritispora, and M. viscosa; see Buller,
1941).
The migration of nuclei in the diploidization of the hyphae has been
followed in T. erythropus and T. trifoliz', but there is so much irregularity in
the nuclear behaviour, the-formation and solution of septa, and in the appear
ance of the first clamp, that it is not possible to give a simple statement (see
Buller, 1941, Noble, r937). Hyphal branches develop independently of the
nuclei in T. trifolii and reach a considerable length before the nuclei pass into
them.
FUNCT I O N
Temperature-relations. The researches of Remsberg have shown that at
least fourteen species of Typhula are adapted to low temperatures. To these
may be added the observations of Imai, Volk, and Noble, as I have noted in
Table IV, from which one may reliably conclude that low optimal temperatures
characterize the genus and explain its absence from tropical countries.
TABLE IV. TEMPERATURE-RELATIONS IN TYPHULA
-
Temperature °C.
Species Min. Max. Opt. Authority
T. umbrina 0 18 6-12 Remsberg
T. idahoensis .. 0 18 9-12 Remsberg
T. incarnata 18 9-12 Remsberg (as T. Itoana)
-
0
,, 0 25 Tasugi, Imai (as T. Itoana)
,, 0 25 8 Volle (as T. graminum)
T. intermedia 0 21 12-15 Remsberg
T. phacorrhiza ,, " " ,,
T. subulata, " ,, ,, "
T. variabilis ,, " " ,,
T. virgata ,, " " ,,
T. latissima 0 25 12-15 "
T. gyrans 0 25 12-15 "
T. sphaeroidea ,, ,, " "
T. viburni 0 27 12-18 "
T: sclerotioides
-�
0 25 15-17 Macdonald (as T. gyrans)
T. pertenuis 0 21 15-18 Remsberg
T. athyrii 25 18-21 "
- -
0
T. trijolii - Sclerotia at 13 ° C., not at 23 ° C.
(Noble)
Remsberg says that all the fourteen species studied by her grew well at or
near 0-3° C., and that the three seriously pathogenic species had lower ranges
and optima than the saprophytic.
Production of Sclerotia. They develop readily in cultures from 5 to 14 days
TYPHULA 167
old. At low temperatures they are crowded, even coalesced, and at high
temperatures they are scattered. In general they seem to require no special
conditions beyond food-supply, though Noble found that the sclerotia of
T. trifolii were formed at 13° C., and not at 23° C., as if their development was
dependent on temperature, and Macdonald found 13-15° C. to be the optimal
temperature for their production in T. sclerotioides.
Production of Fruit-bodies. In contrast to the ease with which sclerotia
appear in culture, fruit-bodies require special conditions which also determine
their development in nature. Early attempts to obtain fruit-bodies in culture
generally failed until Yasugi (1929, 1935) and Imai (1936) showed that low
temperatures and bright or full sunlight were necessary for the'development
of fruit-bodies of T. incarnata ( = T. Itoana). Remsberg has corroborated
these results and added more exactly that light of short wave-length (2,700-
3,250 A0) is necessary rather than sunlight: such light is not transmitted by
ordinary window-glass and hence, no doubt, the general absence of fruit
bodies from artificial culture. By applying the requisite conditions of low
temperature and short-wave light, sporophores can be obtained at will in
culture and immediately after the development of sclerotia. ·Nevertheless, there
are exceptions. Thus T. sphaeroidea will produce abundant fruit-bodies at
its optimum temperature of 12-18° C., but sterile white fruit-bodies at
9-12° C.: in T. sclerotioides the optimum temperature for the production of
fruit-bodies is 17-20° C., which is slightly above that for mycelial growth and
that for the production of sclerotia. Possibly the sclerotia, in nature, need a
resting-period before producing sporophores. Thus Noble found that the
sclerotia of T. trifolii would produce fruit-bodies directly after formation, but
if they were removed from the medium to damp sand, then a considerable
resting period intervened. In nature the sclerotia may become detached
physically or, more probably, physiologically through exhaustion of the food
supply in the substratum. In T. sclerotioides, however, Macdonald found no
such dormant period for detached sclerotia.
Growth of the Fruit-body. On this point I can find no other than Tulasne's
observation on T. variabilis, that its fruit-bodies grew slowly and took mostly
15-20 days to reach their full size, in some cases even a month. Further, he
found that the fruit-bodies wilted and became flaccid in the day-time, through
the dry air, but revived at night, and that their growth was thus continually
inhibited during the day. Now this is as long as it takes-the huge Clavaria
delphus pistillaris to grow a fruit-body five-ten times as high and many more
times as massive. The low temperature may retard growth, but the structure of
the fruit-bodies in Typhula with their agglutinated hyphae, at least on the
surface of the stem, is so typical of the xerophytic construction of fungi, that
I think Tulasne's observation is generically applicable. The slow growth
curiously accords with the delayed development as a resting sclerotium.
Seasonal Fruiting. It is known that fruit-bodies of several species can be
found both in the spring and in the autumn, unlike the majority of temperate
fungi, which have a single season. The facts, which have been mentioned in
the preceding paragraph, and the following observations show clearly the
seasonal periodicity of Typhulas in temperate countries.
TABLE V. SEASONAL FRUITINGS IN TYPHULA
Species Season of Sclerotium Season of Fruit-body Authority

--------
T. variabilis winter (sclerotium autumn (sclero- Quelet (for. T. semen)
white, soft, tium black, hard,
smooth) rough, in summer)
T. intermedia spring autumn Remsberg
" winter spring Appel
T. phacorrhiza spring autumn Remsberg
T. idahoensis "
T. incarnata j ,, Imai (as
I T. ltoana)
T. tochinaiana i Imai
Supposing the basidiospores to germinate in autumn, the sclerotia will

form during the winter which, according to its severity, may inhibit mycelial
growth entirely or but little; if at all. Lack of sunlight will prevent the fruit
bodies from forming until spring. Then the basidiospores may grow on fresh
substrata during the remainder of spring and early summer, but high tempera
tures will prevent the appearance of fruit-bodies until autumn. Thus may
arise the double season, as in T. intermedia, and I think also in T. phacorrhiza,
T. gyrans, and, perhaps, T. pertenuis. On the other hand, if the sclerotia do
not mature by spring, as in T. varialnlis, the combination of light and tempera
ture-factors will give the single autumn season.
Longevity of the Sclerotia. The one exact record that I have found is Yolk's
observatio� that the sclerotia of T. incarnata ( = T. Itoana) may survive 25
months in the dried state. Clearly the sclerotia of most species can survive
several months desiccation during a summer.
EVOLU T IO N
Fruit-body. Progressive diminution of apical growth, as a process of ...
juvenescence, and reduction in the number of concrescent hyphae, lead in
Typhula to the ultimate- state of the subglobose head on a short, filiform,
sterile stalk: no species has yet been discovered without a stalk. The sub
hymenium has also dege_nerated, but the two processes have not been coinci
dent fpr, in T. gyrans, there is limited apical growth and well-devel9ped
subhymenium and, in T. phacorrhiza, unlimited apical growth and negligible
· subhymeniu'!,l).. For which reason, species like T. varialnlis and T. intermedia
with elongate heads and moderately developed subhymenium appear to have
the more primitive construction, like a reduced Clavariadelphus (their
sclerotial structure is also simpler than in T. phacorrhiza).
Sclerotium. As a concrescent mass of hyphae without concerted growth of
the hyphal tips, the sclerotium appears as a rudimentary and sterile fruit-body
without the usual form-factors. The stimulus for concrescence, which is the
first indication of the development of a fruit-body from a mycelium, develops
TYPHULA 169
in the Typhula-mycelium, but no other factor. Aimless intricate growth
forms the mass of hyphae; agglutination at the surface gives the cortex;
excretion of water concentrates the medullary hyphae into the storage-tissue;
and the resting sclerotium is formed. Thus a complete failure of the fruit
body has been turned by some new physiological means into a storage organ
from which, under the appropriate conditions of temperature and insolation,
the fruit-bodies will develop. In considering the origin of the sclerotium,
therefore, one must study the degeneration of the form-factors of the fruit
body and the concentration of the cytoplasm in the medullary hyphae.
Concerning the first point, the sterile fruit-bodies which develop in culture
and the special conditions of light and temperature needed to produce fertile
fruit-bodies must be related phenomena: perhaps, also, the formation of
stolons, which appear as sclerotia with apical growth and suggest the penulti
mate state of .degeneration into the sclerotium (cf. Macdonald's observations
on T. sclerotioides, p. 683). Concerning the second point, the excretion leading
to the deposition of crystals between the hyphae of the fruit-body must be a
related phenomenon. The d�velopment of the cortex of the sclerotium is
clearly homogeneous with the agglutination of the peripheral hyphae in the
stalk of the fruit-body; cf. the haploid fruit-bodies of T. trifolii developed
from the mycelium. Thus there appear in the equipment of the Typhula
fruit-body all the factors necessary to make a sclerotium. And the tissue at
the base of the fruit-body of Pistillaria setipes needs only agglutination and
endurance to convert it into a minute sclerotium (Text-Figs. 39, 51).
Agglutination of the Hyphae. In it� simplest form agglutination affects only
the peripheral hyphae of the stem, perhaps only in its lower part, and the
peripheral hyphae of the sclerotium. In T. sclerotioides the agglutination
extends more deeply. In T. gyrans it extends throughout the tissue of the stem
and sclerotium and, in T. phacorrhiza, it extends also throughout the head,
though the texture of the hyphal walls in the head and central part of the stem
is never so tough as in the sclerotium or on the surface of the stem. This
progressive alteration of the hyphal walls has been the. main evolutionary
process within the genus, rendering the mechanism of the sclerotium and the
fruit-body progressively more xerophytic and adaptable to low temperatures.
Affinity. It is in this connexion that the subgenus Typhulopsis of Clavaria
delphus is remarkable as a connecting-link. Enlargement of the fruit-body of
Typhula, or Pistillaria, would produce exactly such growth-forms as C.
fistulosus and C. junceus, and expansion of the slender primordia of these would
produce the massive fruit-body of C. pistillaris. In reverse or4er, one sees
the limitation of the mycelium from unrestricted growth in deep humus
(C. pistillaris) to twigs, leaves, and particular humus (subgen. ,Typhulopiis),
and finally to the specialized and very limited habitats of Typhula and
Pistillaria: their fruit-bodies conform with their mycelial habitat. Then, in
the subgen. Typhulopsis, reduction in size of the primordium, but the reten
tion of the prolonged apical growth and high degree of inflation typical of
Clavariad<:!lphus, results in slender elongate fruit-bodies supported by the
agglutinated surface-layer of the stem: the agglutination of the surface-layer
extends in some degree, even to the apex of the fruit-body, rendering the
hymenium rather firmly, if microscopically, subgelatinous. Then, in C.
junceus, the fruit-body may become sterile, lose the form-factor of the hymen
ium, and develop into a rhizomorph with adventitious branches. Further
reduction of the rhizomorph, perhaps merely through limitation of the
substrate, that is through depauperation, would produce an abortive outgrowth
transformable into a sclerotium. The base of the hairs at the base of the
fruit-body and on the rhizomorphs of C. junceus have exactly the decumbent,
sinuous, applicable form to make the epidermis characteristic of the Typhula
sclerotium. It would seem that there is an. almost unique series from Can
tharellus through Clavariadelphus to Typhula and Pistillaria. Persistence of
such links as C. truncatus and C. junceus indicates a relatively recent evolution
of the series.
Primitive Species. Though it is unlikely now that one species can be
selected as more primitive than another, it is possible to contrast what appear
to have been the characters of the ideally primitive species with those of the
ideally advanced species, even if this has not yet been achieved. T. variabilis
seems nearest to the primitive state. Thus:
TABLE VI
Primitive Typhula Advanced Typhula

Head . With unlimited apical growth: Limited growth: subglobose,
elongate cylindric with fili short, and blunt: agglutinated
form attenuate apex: not ag with softly but firmly mucila
glutinated. ginous hyphal walls (as in
T. quisquiliaris).
Stem . Agglutinated only at the sur- Wholly agglutinated, toughly
face (? in lower part only). (at least in the lower part).
Spores Large. Small.
Subhymenium Well-developed. Well-developed or absent.
Hyphae Agglutinated only on the sur Wholly agglutinated (toughly
faces of stem and sclerotium. in stem and sclerotium).
Sclerotium . Cortex one hypha thick, with Wholly agglutinated, with
cuticle. cuticle (or epidermis?)
As specialities there may be added the lowering of the optimum temperature .

for growth and the parasitic habit.
CLASSIFICATION
The hyphal structure gives a means of dividing the genus into two sub
genera, Phacorrhizae and Subcarnosae, as defined on page 659. It is the most
fundamental property among Homobasidiomycetes, and, as the distinction
applies to the fruit-bodies, the sclerotia, and the mycelia (cf. the tough
mycelium of T. gyrans and T. phacorrhiza in the Phacorrhizae), it seems a
natural partition of the genus. Whether the subgenera are now absolutely
distinct cannot be known until all the species have been examined fully, but
TYPHULA 171
T. sclerotioides suggests in its broad sclerotial cortex an intermediate state.

This main distinction must have occurred, however, very early in the evolu
tion of the genus because species with elongate heads and large spores occur
in both subgenera. Then, within each subgenus, the primitive features must
have been modified independently for there is much variation in detail and,
as yet, t4ere seems none to correspond exactly with the hypothetical advanced
Typhula. Within each, therefore, we may expect to find lesser series recogniz
able from such specific peculiarities as colour, caulocystidia, habitat, and so on,
and in which the head has become progressively shorter or the spores smaller,
or the subhymenium narrower: superficially, members of one series may
resemble those of another through having arrived at the same state of parallel
evolution. The curnplications which may be expected are shown in the
following diagram of the possible evolution of the Phacorrhizae.
In the Subcarnosae, the coloured species T. van"abilis, T. subulata, T.
idahoensis, T. umbrina, and T. latissima may be a natural series with reduction
of the head, in contrast to the colourless species T. intermedia, T. pertenuis,
T. virgata, T. uthyrii, T. trifolii, and T. graminum, but T. erythropus in the
second series is exceptional. So little is known of the microscopic structure of
their fruit-bodies, however, that it is too uncertain to discuss their specific
affinity in detail. It is the subgenus most intimately related with Pistillaria.
AFFINITY OF THE PHACORIUiIZAE
��-1-��--��---��
Large-spared Small-;pored
Ancestor
/�-�
T. phacorrhiza
(reduced subhymenium)
elongate hend
1- - -
I
t shortly cylindric
I- T. gyrans or clavate .head
T. quisqui/iaris
-- � T. viburni
subglobosc
head
T. sphaeroidea
Regarding the spores, the large ones seem pnm1t1ve because they
occur in all species with the primitive elongate head (with the possible
exception of T. betae and T. cystidiophora): they preponderate among those
with primitive sclerotia (filamentous medulla), and the small spores pre
dominate among species with reduced, clavate, or subglobose heads. Analogy
with Clavariadelphus, from which Typhula seems to have been derived,
suggests that the large spore is the primitive state.
CLASSIFICATION
NATURAL CLASSIFICATION OF CLAVARIOID GENERA
Dimitic with colourless skeletal hyphae: generative hyphae usually
clamped: fruit-bodies with very slender filiform branches or the
small fruit-bodies simple: branching flattened or adventitious:
flesh tough, drying horny: sp. white(? brownish in some spp.),
smooth, aguttate: no gloeocystidia Pteruloid-series
Monomitic, or with gloeocystidia: branches rarely filiform

With dichophyses and gloeocystidia, or with Hymenochaete-setae;
without clamps, hyphae not inflating: fruit-bodies with flat
tened branching, never simple, tough, drying coriaceous: sp.
white(? drying yellowish), small, smooth, aguttate Xanthochroic-
series
Without dichophyses or setae
With gloeocystidia: fruit-bodies with pyxidate branching, or
simple, narrowly obconic and with sterile top, tough or
gelatinous, drying horny: hyphae clamped, inflating or not:
sp. white, small, aguttate: lignicolous (rarely terrestrial) Clavicorona
Without gloeocystidia
Fruit-bodies with flattened branching, tough, rarely gela
tinous, drying coriaceous or horny: hyphae not inflating,
often thick-walled: sp. white or coloured, smooth, rough,
or echinulate, often angled, aguttate Thelephoroid-
series
Branching not flattened (or, if so, then 2-spored, with gut
tate spores, Clavulina, or with ochraceous spores), or
simple: usually fleshy, waxy, or gelatinous: hyphae
generally inflating
Branched, rarely simple: sp. ochraceous or brown (the colour
in the wall) or white and then lignicolous or epiphytic and
with thick-walled hyphae: mostly with clamps (at least,
if white-spored): sp. ellipsoid, often elongate, rarely
subglobose, guttulate or not . Ramaria-series
Sp. white (rarely with coloured contents: in some, drying
yellowish: if with brownish wall, then subglobose and
guttate): hyphae thin-walled (if thick-walled, then with
simple fruit-bodies)
Simple, large, and humicolous to very small and epi
phytic (with or without sclerotium), rarely with
adventitious branching: some genera with sterile top or
�inflated head to the fruit-body: fleshy, waxy, or gela
tinous, rarely tough: hyphae usually clamped and
inflating: sp. aguttate, or with vague guttulae . Clavariadelphus-
senes
Branched or simple, mostly terricolous, generally fleshy
or brittle: hyphae typically inflati_ng, sometimes
secondarily septate without clamps: sp. mostly I-gut
tate or multiguttulate or, if aguttate, then with
secondarily septate hyphae . Clavaria-series
,CLASSIFICATION 17 3
p TER ULOID-SERIES
Cystidia dimorphous, conical or lobed-digitate in the same fruit
body: simple, filiform, to 2 cm. high, or minute and capitate:
hymenium not thickening: skeletal hyphae of limited growth:
epiphyllous: trop. Dimorphocystis
Cystidia absent or simply conical-ventricose: hymenium generally

thickening: skeletals of unlimited growth
Fruit-bodies pointing or curving down, as a cluster of inverted
spines, simple or adventitiously branched, to 2·5 cm. long:
cystidia none: sp. large: lignicolous Deflexula
Fruit-bodies erect
Fruit-body simple to rather freely branched, with a resupinate
patch round the base which is fertile when facing down:
lignicolous: Old World tropics Pterulicium
Without such a resupinate patch: fruit-body highly branched

to simple, filiform and minute Pterula
(Sp. ochraceous, 6 X 4/L: fruit-body with dichotomous branch

ing: skeletal hyphae 1 ·5-3 fL wide: Brazil Lachnocladium
reticulatum)
(Skeletal hyphae only in the sterile subiculum at the base of the
lignicolous fruit-body: sp. ochraceous Ramaria,
Stricta-group)
XANTHOCHROIC-SERIES
With Hymenochaete-setae: hymenium amphigenous, ? not thicken-
ing: trop. Amer. Clavariachaete
With dichophyses and gloeocystidia: hymenium unilateral on the

undersides of the branches, generally thickening: pantropical . Lachnocladium
THELEPH OROID-SERIES
Sp. smooth, white: hyphae rather thick-walled, with or, typically,
without clamps; fruit-bodies white, pallid, brownish or yellow�
ish, tough or toughly gelatinous: mainly trop. Aphelaria
Sp. more or less angular, verruculose, echinulate, or tubercular:

hyphae clamped
Sp. white, angular-echinulate: fruit-body white, yellowish,
pinkish, or purplish, branched m one plane, secondarily
distorted Scytinopogon
Sp. fuscous, brown, umber, or purple: fruit-body or hymenium

similarly coloured, or blackish: branched in alternating planes:
flesh generally turning blackish-green with dilute alkali Thelephora'
1
Not further considered in this work.
174 CLASSIFICATION
RAMARIA-SERIES
Sp. yellow, ochraceous, or brown, smooth or variously marked,
mostly guttulate: fruit-bodies often highly coloured: terrestrial
or Iignicolous Ramaria
(With narrow skeletal hyp hae 1·5-3fi wide: Brazil: see Lach-
nocladium reticulatum)
Sp. white or very pale yellowish (? darker on drying), generally
aguttate, smooth, narrowly ellipsoid: hyphae becoming thick
walled: fruit-bodies white, pallid, tan, ochraceous, or brown-
ish, tough, rarely simple (L. mucida); lignicolous or epiphytic Lentaria
(Simple, small, obconic, with sterile top: on sticks, &c.: trop.
Amer. Caripia)
CLAVARIADELPHUS-SERIES
With oleocystidia or thick-walled cystidia: fruit-bodies small or
minute: hyphae agglutinated on the surface of the stem
Inverted, spine-like, solid, with sterile base to the fertile head,
densely gregarious: some hyphae in the head secondarily septate
with moniliform rows of cells: lignicolous: Malaya Honnomitaria
Erect or ageotropic, with suhconic to subglobose, hollow, inflated
head, smooth or plicate or becoming compressed: lignicolous
or epiphytic: mainly trop. Physalacria
(Cystidia thick-walled: head solid, clavate: trop. Chaetotyphula)
Cystidia, if present, as thin-walled cystidioles (except Chaeto
phula)
Hymenium thickening
Pendulous, watery gelatinous, to 3 cm. long: spores 5--'7 x
3-5µ,: Iignicolous: Tasmania, trop. Amer., S. U.S.A. Myxomycidium
Erect, obconic, or turbinate, with sterile top, to 2 cm. high,
firm, tough, drying woody: sp. 5-6 X3-3·5µ,: hyphae thick
walled: lignicolous: trop. Amer. Caripia
Erect, fleshy, waxy or firm, neither gelatinous nor with thick
walled hyphae: sp. larger
-1·5 cm. high, white, brittle, soft, with sterile truncate
apex: sp. 10-12X6-8 µ,: on leaves and twigs: Malaya Araeocoryne
Larger, varying massive, clavate or Cantharelloid (with
sterile truncate top) to slender or filiform (mostly more
than 3 cm. high), yellow, brown, rufescent, umber, or
tinged pink or violet: flesh spongy, firm, or, in the slender
fruit-bodies, rather rigid: mostly N. temp. Clavariadelphus
Hymenium not thickening: small to minute, mostly filiform or
with filiform stem and clavate to subglobose head: epi
phytic, rarely terrestrial
Inverted, as down-pointing spines, white or yellowish: sp.
small: lignicolous Mucronella
Erect
Hyphae not inflating: fruit-bodies mostly minute, filiform
acerose, rarely branched, sessile: temp. Ceratellopsis
CLASSIFICATION 175
Hyphae inflating more or less, agglutinated on the surface
of the distinct stem
With sclerotium: temp. Typhula
Without sclerotium
With thick-walled cystidia: trop. Chaetotyphula
Without thick-walled cystidia
Head discoid, submarginate, fertile only on the upper
surface, minute: temp. Pistillina
Head cyli111dric, clavate or globose, wholly fertile:
mostly temp. Pistillaria
CLAVARIA-SERIES
Basidia 2-spored, cylindric to subclavate, usually secondarily sep
tate aftc;r spore-discharge: sterigmata usually strongly incurved:
sp. smooth, broadly ellipsoid to subglobose, 1-guttate or multi
guttulate: fruit-bodies simple or branched with radial, flattened,
or cristate branching: mostly terrestrial, few lignicolous: temp.
and trop. Clavulina
Basidia clavate with straight sterigmata, mostly 4-spored, not
secondarily septate: branching never flattened or cristate
Hyphae without clamps, mostly secondarily septate: basidia
without clamps or with a wide loop at the base: fruit-bodies
simple or branched, mostly brittle: sp. aguttate or multi-
guttulate, rarely 1-guttate, smooth: temp. and trop. Clavaria
Hyphae clamped, not secondarily septate: sp. mostly 1-guttate
Sp. minutely echinulate or verrucose: fruit-bodies branched:
temp. and trop. . Ramariopsis
Sp. smooth, in a few cases aculeate or rough (the fruit-bodies
then simple): fruit-bodies simple or branched, mostly fleshy
and somewhat brittle, a few gelatinous or tough, white or
variously coloured, often yellow, orange, pink, or red: temp.
and trop. Clavulinopsis
ARTIFICIAL KEY TO CLAVARIOID FUNGI WITH

COLOURED SPORES
Fnµt-bodies branched
Sp. fuscous-umber or vinaceous, shortly ellipsoid or globose,
angular, verrucose or echinulate: fruit-bodies with flattened
branching, purplish, vinaceous, or umber, the tissue generally
blackening or giving a blackish-green colour with dilute alkali Thelephora 1
Sp. globose, smooth, ? pale brown, 1-guttate: basidia 2-spored,
subcylindric: fruit-body brown, with filiform branches:
Malaya . Clavulina
decipiens
1
Not considered further in this work.
CLASSIFICATION
Sp. very pale yellow, elongate, smooth, aguttate: hyphae thick-
walled: on wood or leaves Lentaria
Not so Ramaria
(Hyphae not inflating: branching flattened: see Aphelaria)
Fruit-body simple
Sp. pink (coloured contents), oblong-ellipsoid: fruit-bodies pink,
red, or orange: trop. Asia Clavaria
helicoides
Old spores on the hymenium stained yellow or red Clavulinopsis
Fresh spores light ochraceous, orange, or brown

Sp. ellipsoid
Sp. rough, 7 1 -1X5-6 f-',: fruit-body to 15 mm. high, spathu
late: on earth Clavulinopsis
spathuliformis
Sp. smooth
Fruit-bodies large, clavate, or &ub-Cantharelloid, to
smaller, ligulate, or subcylindric: temp. Clavariadelphus
Very small, to 4 mm. high: sp. 9-10X4/L, yellow: on

wood: China Pterula vitellina
. Sp. subglobose
1-2 mm. high, head inflated: sp. 4-6 f-',: on wood: Africa Physalacria
togoensis
2-4 mm. high, filiform: sp. 4-4·51-'-: on dead leaves: Africa Pistillaria
phaeosperma
17-26 mm. high, white then pale flesh-colour, stem long:
sp. ferruginous: on trunks of Cyathea . Ramaria
cyatheae
9-u cm. high, pale rufescent: sp. 4 x 31-'-: on the ground in
forest: New Guinea . Clavaria
subfistulosa
ARTIFICIAL KEY TO WHITE-SPORED CLAVARIOID FUNGI

Hyphae incrusted with yellow granules: fruit-body 2-10 mm.
high, greyish-yellow, sparingly branched, or orange-red and
simple: sp. 7-u X 3-51-'- (or 4-8X4-51-'-): on bark: trop. Lachnocladium
aurantiacum
Hyphae not incrusted
Basidia with 2 incurved sterigmata, usually secondarily septate
after spore-discharge, subcylindric: fruit-bodies branched or
simple: sp.-1-guttate or multiguttulate, globose to broadly
ellipsoid Clavulina
Basidia clavate, not secondarily septate: sterigmata straight

(2-4, rarely 6 or 8)
Branched fruit-bodies
With dichophyses and gloeocystidia: trop. Lachnocladium
With brown, thick-walled setae: trop. Amer. Clavariachaete

CLASSIFICATION
Not so
Branches filifonn, slender, tough, with elongate, acerose
tips: some with fruit-bodies inverted A
Not filifonn
Branching flattened B
Branching pyxidate with sterile, slightly concave tips:
with gloeocystidia Clavicorona
Branching radial or dichotomous, neither flattened nor
pyxidate . C
Simple (branching exceptional or merely adventitious)
With si.lbglobose, capitate, or discoid head, small or minute:
epiphytic D
With truncate, subpileate, or subcyathifonn sterile apex,
massive to small E
Inverted, pointing down or pendulous, mostly small,
slender, or minute: lignicolous . F
With a sclerotium: small, slender, or minute: epiphytic:
temp. Typhula
Not so
Terrestrial or humicolous, large or small': if lignicolous,
then fairly large (more than 1·5 mm. thick) G
Epiphytic, rarely terrestrial: small, slender, or minure,
less than 1·5 mm. thick . H
A. Branches filiform, tips acerose

Dimitic: drying wiry or horny and, often, brown or blackish
Inverted, pointing down, as a tuft of spines: small or minute:
sp. large: lignicolous Deflexula
Erect
With thin, fertile Corticioid base surrounding the simple or
sparingly branched, white or pinkish-drab frµit-bodies: on
wood and plant-remains: Old World tropics Pterulicium
Without such a fertile base Pterula
Monomitic
Sp. 13-16 X 4µ: hyphae thick-walled: lower branches stout, the
upper often filiform: tropics Lentaria
surculus
Sp. 5-7 X 3-4 fL: hyphae 3-4 fL wide, thin-walled: all the branches
slender: France Clavulinopsis
propera
B. Branches flattened
Sp. rough or echinulate, angular Scytinopogon
Sp. smooth
Orange-yellow, to 11 mm. high, simple or once dichotomous,
the tips 2-6-fid: sp. 4·5-6 µ wide, globose, 1-guttate: hyphae
c. 2 µ wide : on amenta of Rhamnus : Europe Clavulinopsis
pusilla
5119 N
CLASSIFICATION
Not so
Hyphae inflating, becoming thick-walled, clamped: sp.
elongate, aguttate: lignicolous Lentaria
Hyphae not inflating, more or less thick-walled or gelatinous,

with or without clamps: mostly terrestrial and trop. Aphelaria
C. Branching radial
Hyphae thick-walled, clamped: sp. narrowly ellipsoid, often much
elongate and subsigmoid, aguttate: tough, white to ochraceous
or yellow-brown, often with copious byssoid or floccose mycelium:
lignicolous, and on plant-remains . Lentaria
(Hyphae gelatinous, not inflated: New Caledonia . Aphelaria

fiabellata)
Hyphae not thick-walled: sp. round or shortly ellipsoid, mostly
guttate: mycelium rarely conspicuous: mostly terrestrial, a
few lignicolous
Sp. rough, verrucose or echinulate Ramariopsis
Sp. smooth
Hyphae secondarily septate, without clamps: spores aguttate:
brittle: pink, lilac, or violet . Clavaria
Hyphae not secondarily septate, clamped: sp. mostly 1-guttate:

waxy, firm, not very brittle, white or variously coloured,
often yellow Clavulinopsis
D. Simple, small OT minute; head capitate or discoid

With cystidia
Cystidia thick-walled, fusiform or lobed-digitate in the same
fruit-body: hyphae dimitic: head solid, to 1· 5 mm. wide:
trop., on dead leaves Dimorphocystis
Cystidia never lobed-digitate, often resin-incrusted: monomitic:
head mostly hollow, 0·5-10 mm. wide: 1-25 mm. high: mostly
trop. Physalacria
Without cystidia: monomitic: head 0·5-1·5 mm. wide
Head discoid, fertile on the distal side only, solid: Europe,
U.S.A. . Pistillina
Head subglobose, wholly fertile, hollow Pistillaria
E. Simple: apex truncate OT subcyathiform, sterile

Sp. more than 7 µ long
Terrestrial: large, ochraceous, reddish, pinkish, brownish,
blackish, or violaceous, firmly fleshy or spongy: N. temp. Clavariadelphus
On plant-remains: small, slender, white· or cream, narrowly
obconic, fragile: Malaya Araeocoryne
Sp. smaller
With gloeocystidia Clavicorona
CLASSIFICATION 179
Without gloeocystidia
Terrestrial: small, white, soft clubs with thin-walled hyphae:
(? apex not sterile): France . Clavaria
Corbierei
Lignicolous: broadly obconic, white to brownish or ochra-
ceous, tough or woody (on drying): hyphae thick-walled:
with cystidia: trop. Amer. Caripia
F. Simple, inverted, lignicolous

Watery gelatinous, pendulous: Tasmania, trop. and N. Amer. Myxomycidium
Waxy or firm
Dimitic: usually as clusters of small spines: sp. more than 9 µ
long Deflexula
Monomitic: usually densely gregarious
Stem 1-2 cm. long: heads 1-2 cm. long: trop. Africa Mucronella
tenuipes
Stem sherter
Clear yellow, shortly stalked, with sterile base to the conic
spine: with cystidia: Malaya Hormomitaria
White or pallid yellowish, sessile or nearly so, without
cystidia . Mucronella
G. Simple, terrestrial or, if lignicolous, more than I·S mm. wide

Sp. rough, verruculose or echinulate: with clamps Clavulinopsis
Sp. smooth
Less than 1 cm. high, more or less gelatinous: 4-6-spored: trop. Clavulinopsis
Not gelatinous: 2-4-spored
Hyphae without clamps, often secondarily septate: sp. agut
tate or finely multiguttulate, rarely 1-guttate: brittle (except
the tough tropical C. fossicola): rarely yellow . Clavaria
Hyp hae clamped, not secondarily septate: often yellow,
ochraceous, or orange
Sp. subglobose, or shortly ellipsoid, mostly 1-guttate:
fleshy-brittle to rather firm: temp. and trop. Clavulinopsis
Sp. aguttate, rather large, elongate, ellipsoid, or amygdali
form: more or less ochraceous, inclining to pinkish,
orange, tan, rufescent, or brown: spongy or firmly fleshy.:
temp. Clavariadelphus
H. Simple, epiphytic, rarely terrestrial, less than I·S mm. wide

Dimitic (hyphae not inflating)
With thick-walled conical cystidia and smaller lobed-digitate
cystidia: trop. Dimorphocystis
Cystidia absent or simply conical or ventricose
With a fertile Corticium-like patch round the base of the fruit-
body: Old World tropics Pterulicium
Without such Pterula
180 CLASSIFICATION
Monomitic (so far as known)
Hyphae not inflating: stem indistinct or none: temp.
Orange-yellow: sp. 4-6 µ, wide, globose, 1-guttate: on amenta
of Rhamnus : Europe . Clavulinopsis
pusilla
White or yellowish, to 2 cm. high: sp. 5-7 X 2-2·5 µ,: on rotten
··
wood covered with algae Lentaria mucida,
L. coronilla
White, pinkish, or yellowish, linear, filiform-acerose, mostly
very small: sp. aguttate Ceratellopsis
Hyphae inflating: stem distinct
Hyphae on the surface of ·.he stem agglutinated
Long or rather stout, over 2 cm. high, brownish Clavariadelphus
Small
With thick-walled cystidia: trop. Chaetotyphula
Without such . Pistillaria
Hyphae on stem-surface not agglutinated: mostly more than
12 mm. high
Without clamps: white Clavaria
With clamps
Sp. 1-guttate: white or variously coloured Clavulinopsis
Sp. aguttate or vaguely guttulate: mostly very small Pistillaria
APHELARIA gen. nov.

(Gr. apheles, plain: from the lack of character)
Fruit-bodies branched, in a few species sparingly or subsimple, with flattened
multifid then bifid branching, alternating or apparently in some species in one plane,
often with filiform Pteruloid tips, coriaceous or gelatinous (A. flabellata) white, pale
grey, yellowish, brownish, or pale flesh-colour, never clearly pigmented, often Stereum
like at first.
Mostly terrestrial, few lignicolous: 12 spp. tropical, a few temperate.
Spores white, smooth, subglobose to elongate ellipsoid, aguttate (or minutely multi-
guttulate).
Basidia clavate, rather wide: sterigmata 2-4 (sub-Tremellaceous in A. tuber'osa).
Cystidia and gloeocystidia absent.
Hymenium thickening, generally absent from the stem and uppersides of the branches,
or as a rudimentary sterile hymenium.
Hyphae monomitic, not inflated, long-celled, becoming slightly thick-walled, typically
without clamps, colourless.
Type-species: A. dendroides (Jungh.) emend. Corner.
KEY TO THE SPECIES OF APHELARIA

Toughly gelatinous, large: sp. 5-7 X 3·5-5 µ,: New Caledonia A. flabellata
Dry, more or less coriaceous
Hyphae clamped
Sp. 14-23 X 5-8 µ,: N. temp. A. tuberosa
Sp. 10-15X5-7µ,: New Zealand A. pusio
(Sp. u-13 x4-5·5µ,: U.S.A. Clavulinopsis
pogonati)
APHELARIA 181
Hyphae not clamped
-7 cm. high, much branched
Sp. 8-12,X7'5-10·5µ.: trop. Asia, Australasia, Africa A. dendroides
Sp. 6-8X5-6·5µ.: branches very slender: S. Am. A. tropica
Sp. 5-7X5µ.: hyphae darkening in dilute alkali: Brazil A. aurantiaca
Sp. ?: hyphae not darkening: Brazil A. trachodes
-3 cm. high, sparingly branched: sp. ellipsoid
Sp. 13-15X8-10µ.: ?stem naked: Cuba . A. brunneola
Sp. n-15X5-6µ.: stem smooth or slightly strigose at the
base: Malaya . A. incarnata
Sp. 8-10·5X5·5-7µ.: stem strongly strigose-fibrillose:
Malaya, Philippines A. spiculosa
Clamps?
Sp. 16X8-10µ.: Tasmania A. tasmanica
Sp. 4 ·5-6X3-3 · 5µ.: corticolous : Brazil A. defiectens
(See also Lachnocladia incertae sedis, p. 430, and Pterulae incertae sedis, p. 534.)
A. aurantiaca (P. Henn.) comb. nov.

Basinym: Pterula aurantiaca P. Henn., Hedw. 1904, 174.
4-6 cm. high, orange, drying alutaceous, tough: stem 1-2, cm.X1-2, mm., smooth:
branches dichotomous or verticillate, axils compressed, branchlets subcompressed,
tips subulate.
In swampy ground: Amazon (Rio Negro, Ule 3n1).
Spores 5-7X4·5-5·5(-6)µ., white, smooth, subglobose or ovoid, slightly thick
walled.
Hyphae 3-5(-6)µ. wide, monomitic, without clamps, uninflated, longitudinal,
becoming slightly thick-walled, turning dark brown with dilute alkali.
I have, examined the part of the type-collection in the Kew herbarium. Apparently
it belongs to this genus but the darkening of the hyphae in dilute alkali, together with
the absence of clamps, suggests that it may really be a very primitive member of the
Xanthochroic-series: that is, it appears as a Lachnocladium s. str. divested of
dichophyses and gloeocystidia.
Bresadola gave P. aurantiaca as a synonym of Lachnocladium guadelupense (Sacc.,
Sy!!. 2,3, 192.5, 496). Burt interprets thi� species as a Ramaria, however, in which
I follow him. Possibly Pterula squarrosa P. Henn. (Hedw. 190:z, 6), which Bresadola
also referred to L. guadelupense, should also belong here (as the earlier specific
epithet): its description is:
4 cm. high, brownish, cartilaginous, cinereous pruinose: stem 1-:z cm. X. 5-1 mm.;
branches dichotomous once or twice, terete, arcuately curved, elongate-subulate,
squarrose (dried), to :z.·5 cm. long: spores 3-3·5µ. wide, white, smooth, ovoid: on the
ground: Brazil (Para).
Lloyd referred both P. aurantiaca and P. squarrosa to Aphelaria trachodes, but
I cannot find spores or hymenium on the type-collections of this species and its
hyphae do not darken in the same way.
A. brunneola (B. et C.) comb. nov.

Basinym: Clavaria brunneola B. et C., Journ. Linn. Soc. Bot. 10, 1868, 338: Burt,
Ann. Mo. Bot. Gdn. 9, 19:z:z, 2.5, t. 4, f. 2.4: Coker, Clav. U.S. Can., 192,3.
-2,·5 cm. high, pale yellow (? dried): stem cylindric, thin: branches few, flattened,
2,-3-chotomous below, spathulate dichotomous above, divaricate, tips subulate: tough
(dried).
182 APHELARIA
On banks: Cuba (Wright 239).
Spores 13-15X8-10µ., white, smooth, amygdaliform and slightly thick-walled
(Comer, type-specimen in herb. Kew.): 13-15X9-10µ., broadly ellipsoid, pointed
at both ends (Burt, co-type): u·5-12·5 x7·5-9·5µ. (Coker, co-type).
Hyphae 2-4µ. wide, slightly thick-walled, not. inflated, without clamps (Comer,
type in herb. Kew.).
This is very near A. incarnata, but the spores are much wider and have distinctly
thickened walls, as in A. spiculosa and·Pterulicium xylogenum. The fruit-body also
resembles that of A. incarnata. The basidia are probably 2-spored.
A. deflectens (Bres.) comb. nov.

Basinym: Lachnocladium deflectens Bres., Ann. Myc. 18, 1920, 50, f. 4.
Caespitose, moderately large: stems 5-10X2 mm., connate below, rarely solitary,
pale, soon very much branched: branches repeatedly and irregularly branched,
flattened below the arcuate or angular axils, terete or subcompressed, smoky flesh
colour (dried), pallid.and pubescent on the fertile side (dried), apices acute.
On the bark of frondose trees: Brazil (S. Leopoldo).
Spores 4·5-6X3-3·5µ., white, smooth, oblong.
Hyphae 2-4µ. wide.
The flattened branchings, narrow hyphae, and smooth spores indicate Aphelaria.
Superficially it must resemble narrow forms of Scytinopogon. Compare also Lachno
cladium reticulatum.
A. dendroides (Jungh.) comb. nov.

Basinym: Clavaria dendroides Jungh., Verh. Bat. Genoot. 17, 1839, 33, t. 6, f. 20 (non
Fr. 1857 = Ramaria condensata).
Synonyms: Thelephora dendroides (Jungh.) Lev., Ann. Sci. Nat. 3, 2, 1844, 209.
Merisma dendroides (Jungh.) Lev., Ann. Sci. Nat. 3, 5, 1846, 158.
Pterula dendroides (Jungh.) Fr., Nov. Act. Soc. Sci. Up. 3, 1, 1885, u7.
Lachnocladium dendroides (Jungh.) Sacc. et Syd., Syll. 16, 1902, 213.
Clavariafiagelliformis B., Hook. Fl. New Zeal., 1867, 186.
Lachnocladium flagelliforme (B.) Cke., Handb. Austral. Fungi 1892, 179 (quoad
nomen: f. 79 = Lachnocladium sp.).
Clavaria lurida Kalchbr., Proc. Linn. Soc. N.S.W., 1882, 105 (teste Bres., Ann.
Myc. 14, 1916, 233).
Lachnocladium Kurzii Cke., Grev. 20, 1891, 11.
Thelephora bidentata Pat., Ann. Jard. Bot. Buit. Suppl. 1, 1897, II5, t, 24, f. u-13.
Clavaria ornithopoda Mass., Kew Bull. 1901, 154.
Thelephora oubanguiensis Har..et Pat., Bull. Mus. Hist. Nat. Paris, 19u, 365.
? Lachnocladiuin aciculare Lev., Ann. Sci. Nat. 3, 2, 1844, 207.
? Merisma aciculare Lev., Ann. Sci. Nat. 3, 5, 1846, 158.
? Thelephorafunalis Lev., Ann. Sci. Nat. 3, 2, 1844, 208.
? Lachnocladiumfunale (Lev.) Sacc., Syll. Fung. 6, 1888, 739.
? Thelephoriz amboinensis Lev., Ann. Sci. Nat. 3, 2, 1844, 207.
? Thelephora scoparia Lev., Ann. Sci. Nat. 3, 2, 1844, 207.
? Lachnocladium scoparium (Lev.) Sacc., Syll. Fung. 6, 1888, 739.
? Thelephora acanthacea Lev., Ann. Sci. Nat. 3, 5, 1846, 147: Moritzi, Syst. Verz.
1846, 123: Pat., Ann. Jard. Bot. Buit. Suppl. 1, 1897, u5.
Text-Figs. 12, 52-6: Plate 14.
-'] cm. high, solitary, gregarious, or caespitose, generally much branched, branch
ings flattened and polychotomous below, dichotomous upwards, in alternating planes,
fastigiate-erect, spirally twisted on drying, dingy pallid white becoming somewhat clay
APHELARIA
co.lour, dirty buff, or pallid dingy alutaceous ihen greyish or greyish-buff from the base
upwards, tips pallid white, coriaceous, tough: drying horny, twisted, fuscous or brownish,
hymenium pallid white, tips fuscous brown.
ThxT-FIG. 52. Aphelaria dendroides: 'Thelephora form', X 2: branch:.tips, X 5.
Stem 3-22 x 1·5-2·5 mm., generally distinct, smooth (no fibrillar processes), often
attached by a white or pale buff, subtomentose mycelium: branches 1-1·5 mm. wide,
flattened and narrowly flabellate, 2-4 mm. wide, and 2-5-chotomous below, succes
sive branchings alternate, attenuate and dichotomous upwards, the tips often elongate
subulate, erect and rather stiff: hymenium absent from the uppersides of the axils and
branches, generally absent from the stem at least in its lower half: smell rather strong
and sour in some collections.
A�HELARIA
On the ground in the forest, in rubber estates and orchards: tropical Asia and
Australasia, Java, Malaya (frequent in the lowlands), Australia, New Zealand, Africa
(Belgian Congo, Th..oubanguiensis).
Spores 7·5;9.x7..:..8 p.,, or 10-12X9·5-10·5 p., in other collections, white, smooth,
TEXT-FIG. 53. Aphelaria dendroides: 'Lachnocladium form', X 1.
TEXT-FIG. 54. Aphelaria dendroides: 'Lachnocladium form', X 1.
subglobose, thin-walled, contents cloudy-granular, aguttate, apiculus 1 p., long.

Basidia 25-6 5 X 9-12 p.,, rather abruptly clavate, very variable in length in the
narrowed, stalk-like part: sterigmata 2-3-4, rarely 1, some fruit-bodies mainly with
2, 5-12 f1, long, 2-3 p., wide at the base.
Cystidia none.
Hymenium thickening to 200 p.,, the old basidia collapsing: subhymenium at first
APHELARIA 185
15-20µ thick, hyphae thin-walled, the cells 8-15 X 1·5-3µ, generally slightly inflating
4-6µ wide at the base of the old hymenia, rather closely interwoven, without clamps.
Hyphae monomitic, without clamps (not even on the basidia), 2-4µ wide, mostly
with slightly but distinctly thickened walls 0·5-1 µ thick, septa 20-250µ apart, not
secondarily septate, the cells often of rather uneven width, branching normal or
sometimes constricted at the origin.
Flesh with longitudinal, very compact hyphae, slightly looser in the central region
but without a distinct pith, short-celled and very compact next the subhymenium.
Sterile surfaces of the stem and branches glabrous or with rudimentary 'basidia
or short blunt hyphal processes -10µ long, without a pile
or palisade, no caulocystidia.
This variable species, which has gone the round of Clavari
oid genera, greatly resembles Pterula, more so when it is dried,
because it becomes hard, horny, fuscous-brown, and twisted,
with pallid hymenium like a typical Pterula. But it differs
from Pterula in its monomitic hyphae without clamps and its
much stouter branches.
The description which I have given is based on my Malayan
collections (seven, from Singapore and Pahang). 'I'he young
fruit-bodies with rather narrowly flabellate, polychotomous
branches appear very different from the other ones with
elongate-subulate tips (never filiform as in Pterula), and even
the older fruit-bodies with blunt, spathulate, dividing tips
appear different from those with acute growing tips: and no
one would imagine the living plants to be the horny, brown,
twisted objects of the herbarium. The spores vary in different
collections; most have smaller spores 7·5--9µ, but two (from
Pahang) have the larger spores, though their fruit-bodies are
identical in every other way. The length of the basidia and
number and length of the sterigmata also vary much in the
same fruit-body: most fruit-bodies seem to have 2-spored
basidia, but in two collections I found as many 3- and TEXT-FIG. 55· Aphe-
4-spored basidia as 2-spored: in neither case were there laria dendroides: 'Pteru-
clamps. la form', x 1.
I have not seen Junghuhn's type-specimen, but I have no doubt that his Clavaria
dendroides is the same as my Malayan collections. Coker gives the spores of an
apparently authentic specimen of C. d,endroides (from Java) in Persoon's herbarium
as II X 7·4µ, smooth, ovoid, hyaline, and the basidia as 2-spored. Moreover, Lloyd,
Coker, and Hennings say that the species is a Lachnocladium, meaning that the fruit
body is tough or horny when dried.
Thelephora bidentata, from the description and figure, is also identical. Patouillard
describes typical young specimens with blunt dividing tips (spores 6-8µ, globose;
, basidia 30x10-12 µ, 2-4 sterigmata). I found the type of T. oubanguiensis (herb.
Paris) identical with A. dendroides (sp. 7-9 X 6·5-8·5 µ, 2-spored, hyphae 2-4µ, no
clamps).
Thelephora acicularis, T. funalis, and T. scoparia, according to Patouillard, are near
his T. bidentata, having similar spores but subulate, not flattened, tips to the branches.
They are clearly better-developed states of A. dendroides, and the co-types in the
Kew herbarium certainly appear identical. Singer, however, says that the type of
L. aciculare Lev. has spores 15-16x9-11µ, with spines 1·5-3·5µ long (Lloydia 8,
1945, 140): it suggests Ramaria Zippe/ii. T. acanthacea, according to Patouillard,
has similar spores 8 µ wide, but the original description gives the fruit-bodies as
multifid-foliaceous with lateral stem: it may be stereoid.
The types of C. fiagelliformis (New Zealand) and C. ornithopoda (Malaya) in the
186 APHELARIA
Kew herbarium are identical with A. dendroides, with spores 9-10 X 6-7 fL and
9-u X 8-9p, respectively.
The type of L. Kurzii (Java), in the Kew herbarium, is a dwarf bushy form probably
developed in dry or open ground. It is much blackened with corrosive sublimate.
The co-type (or part of the type-collection) of Thelephora amboinensis at the Kew
herbarium may be A. dendroides or A. trachodes, which does not dry so horny and
twisted as A. dendroides and has rather wider hyphae. In these respects, the co-type
of T. amboinensis (on sandy ground, Amboina) agrees with the S. American A. tra
chodes, having hyphae 3-'7 fL wide (with the walls 0·5-2·5 fL thick), without clamps,
TEXT-FIG. 56. Aphelaria dendroides, x 1,000.
and having fibrillose-coriaceous texture. But the specimens of the type-collections

of both A. trachodes and T. amboinensis are completely sterile, without hymenium.
The larger spores of this species are probably borne on the 2-spored basidia.
A. flabellata (Wakef.) comb. nov.

Basinym: Clavariafiabellata Wakef., Journ. Linn. Soc. Bot. 46, 1922, 90.
-15 cm. high, large, white, much branched, drying very hard, horny and fuliginous
fuscous :. branches 1-5 mm. wide, fiabellately divided, flattened, smooth: branchlets
1-1·5 mm. wide,, suberect, slender, cylindric, tips tinged fulvous, often flabellately
expanded (?borny-subgelatinous).
On the ground in the forest: New Caledonia (Mt. Canala, 500 m.): said to be
edible.
Spores 5-7 X 3·5-5 p,, white, smooth, broadly ellipsoid or ovoid, blunt.
Basidia very small, 10-15 X 3 fL (Wakef.) (? 30-40 X 6-7 p,, narrowly clavate, but
immature; Corner).
Hyphae not inflated, apparently without clamps, the walls t\uckened and toughly
gelatinous (2-4 fL thick), the lumen 2-5 fL wide; the subhymeriial · hyphae narrow,
dense, and with thinner walls.
APHELARIA
Hymenium thickening.
This species may prove to be a new genus resembling Aphelaria but with toughly
gelatinous hyphae giving the very horny consistency to the dried fruit-body. The
spores, which are very abundant in the hymenium, resemble those of Aphelaria.
In shape, the fruit-body resembles Scytinopogon and Clavulina cartilaginea. The
basidia are agglutinated in a firm layer in the dried specimen and are difficult to see
in detail, but they do not appear to be like those of Clavulina.
A. incarnata sp. nov.

Text-Figs. 57, 58.
r · 5-2 cm. high, solitary or gregarious, sparingly branched, white becoming very pale
ochraceous flesh colour or pale dingy flesh pink from the base upward, tips white,
fibrous subcoriaceous.
TEXT-FIG. 57. Aphelaria incarnata, X 5: branch-tips, X 10.
Stem 4-10 x 0·5-1 mm., sparsely and shortly spicular-strigose below with upward
pointing subdivergent fibrils, or naked, the extreme base slightly dilated, flattened
upward and divided into 2-3 main branches: branches simple or dichotomous once
188 APHELARIA
or twice, terete but slightly flattened below the axils, tips blunt, becoming subulate
and attenuate, sometimes minutely cristate on branching: hymenium generally absent
from the stem: flesh tough, pliant; without smell.
On the ground in the forest: Singapore (Bukit Timah, 16,4,41).
Spores 11-15 X 5-6µ, white, smooth, subcylindric or elongate pip-shaped, aguttate,
the contents granular-vacuolate.
Basidia 30--42 X 8-10µ, clavate, projecting -20µ: sterigmata 2-3-4, mostly 2, 5-8µ
long.
Cystidia none: hymenium thickening to 120µ: hyphae and structure as in A.
spiculosa.
Stem with loose, unmodified hyphal ends 2-3µ wide, projecting obliquely forward,
not as a regular pile, but here
and there concrescent in the
lower part of the stem to form
the strigose fibrils.
This species is close to A.
spiculosa, having the same
structure and hyphae, but it
differs in the · longer spores,
much less spicular-strigose
stem, and rather more frequent
branching: the hyphae are,
perhaps, not so distinctly thick
walled as in A. spiculosa, but
they are branched in the same
way. The larger size of the
spores may be the result of the
reduction of the basidium to
the 2-spored state.
The branching of the fruit
body is by flattening and slight
TEXT-FIG. 58. Aphelaria incarnata, x 1,ooo. spathulation of the tips which ..
then break up into 2-7 minute
fibrillar points, of which the outer two grow into the new branches and the others
remain microscopic and abort: rarely does the central point grow into a slender branch,
giving an apparent trichotomy, and still more rarely does a fourth branch develop.
Successive branchings are in more or less alternate planes, at right angles. The axils
of the branches are often thinly webbed by the residual hyphae of the spathulate apex.
Compare A. brunneola, which is very close.
A. pusio (B.) comb. nov.

Basinym: Clavariapuszo B., Hook. FI. New Zeal. 1867, 186: Massee, Tr. N. Zeal.
Inst. 39, 1906, 40.
Synonym: Pte rulapusio (B.) Bres. et Syd., Philipp. Journ. Sci. Bot. 9, 1914, 352:
Bres., Hedw. 56, 1915, 304: Lloyd, Myc. Notes 60, 1919, 870.
-3 cm. high, drying brownish: stem slender, thickened upward, dividing into a
few cylindric branches equal in length to the stem, spreading at an acute angle, rarely
branched themselves, acute: drying horny.
On the ground: New Zealand (N. Isl.).
Spores 10--15 X 5-7µ, white, smooth, oblong�ellipsoid, obtuse or subacute, apiculus
1-1·5µ.
Hyphae 2-3·5(-4)µ wide, monomitic, clamped, not inflating, walls c. 0·5µ
thick.
APHELARIA
I have added the details of spores and hyphae from the type at Kew. In form,
texture, spores, and narrow hyphae the species is a typical Aphelaria but it differs
in the clamps. I have nevertheless retained it provisionally in the genus. It is possibly
related with the anomalous A. tuberosa. Pterula Decaryi Pat. (? ined., herb. Paris,
Madagascar) seems identical: (sp. II-14 X 6-7 µ: bas. 50 X 12µ, 4 sterigmata 8 µ long:
hyphae 2-3 ·5µ, clamped at some septa, not at others, basidia clamped: hymenium
thickening).
The Philippine specimen referred to C. pusio by Bresadola is almost certainly
Aphelaria spiculosa, for he describes it as larger and more branched and with rounder
spores 8-g X 5-6µ.
A. spiculosa sp. nov.

Synonym: Pierula pusio (B.) Bres. sensu Bres., Hedw. 56, 1915, 304 (quoad specim.
Philipp.).
Text-Figs. 15, 16, 59, 60.
-3 cm. high, scattered, terrestrial, simple or, generally, the stem breaking into 2-4
simple branches or regularly dichotomous 2 4- times with laxly spreading branches,
branching flattened, white then pale drab flesh colour with white tips, tough, pliant.
Stem 5-14 X o·5-1 mm., terete or slightly flattened, generally becoming strigose from
the base upward with upward-pointing fibrils, sometimes wholly strigose when old,
occasionally smooth: branches 0·3--0·8 mm. wide, simple or 1-3 times dichotomous
(occasionally 3- ,hotomous) generally in alternating planes, terete but flattened· and
slightly dilated below the axils, lax, often curved to one side, tips blunt and thick in
young branches, becoming acute and ultimately filiform-attenuate, not cristate:
hymenium covering the whole fruit-body except the tips and the extreme base, or
patchily developed on the lower part of the stem: flesh subcoriaceous, tough, pliant:
no smell.
On bare earth in the forest: Malaya (Singapore, Bukit Timah, 19,4.41, 20.9.41,
4,4.43), Philippine Isl. (Bres. ut P. pusio).
Spores 8-10·5 X 5·5-7µ, white, smooth, ellipsoid, slightly flattened adaxially, blunt
or subacute at the apex, the wall slightly thickened, aguttate, contents vacuo
late.
Basidia 24-32 X 8-10µ, clavate: sterigmata (2-)4, 6-7 /L long: cystidia none.
Hymenium 30µ thick at first, gradually thickening to 200fL-: subh�•menium c. 30µ
wide, of loosely interwoven thin-walled hyphae like the tissue of the thickened
hymenium, without clamps (not gelatinous).
Hyphae 2·5-3·5µ wide, without clamps, monomitic, walls slightly thickened (0·5µ)
and smooth, not inflated, the cells 30-200 µ long, not secondarily septate, colourless,
cylindric or occasionally undulate, even slightly nodose or kinked; lateral hyphae
attached by a constricted base, 1-1·5µ wide, arising as minute, pyriform outgrowths,
apparently without relation to the septa.
Flesh composed wholly of longitudinal, intertwined hyphae without interweaving
laterals and without a compact cortex, all the hyphae slightly thick-walled except
those of the subhymenium.
Strigose fibrils on the stem constructed like the growing tips of the branches of
conical bundles of hyphae, occasionally enlarging into a rudimentary branch.
Sterile hymenium on the stem, irregularly developed, consisting merely of scattered
sterile basidia or rudimentary hymenial hyphae.
Growing tips of the branches 20-100µ wide; hymenium developing 200-500µ
behind the apex by the out-turning of the hyphal tips on the surface followed by
sympodial branches, the first clavate cells generally fertile but some sterile and
producing a narrow excrescent hypha 2-3µ wide from their distal ends.
The branching is normally flattened and dichotomous but sometimes trichotomous
J
\�
ll
TExT-FIG. 59. Aphelaria spiculosa, x 3.
TExT-FIG. 60. Aphelaria spiculosa: young fruit-bodies, x 5.

------------,.
APHELARIA 191
with a small central tip which generally aborts. Successive branches are generally
in planes more or less at right angles. Sometimes there are minute fascicles of hyphae
near the bases of the branches, like incipient strigose fibrils on the stem, but they
are evanescent and soon covered by the thickening hymenium: they may cause the
branching tips to appear microscopically cristate.
A. tasmanica (Lloyd) comb. nov.

Basinym: Pterula tasmanica Lloyd, Myc. Notes 80, 1923, 1227, f. 2539.
Fruit-bodies like A. dendroides and drying in the same way, homy-fuscous with
spirally twisted branches: Tasmania (L. Rodway).
Spores 16x8-10µ, white, smooth, obtuse: basidia?: cystidia none.
Lloyd suggests this may be Tremellodendron. His photograph exactly resembles
A. dendroides, so that I would identify it as such but for the larger and ellipsoid
spores.
A. trachodes (B). comb. nov.

Basinym: Thelephora trachodes B., Hook. Joum. Bot. 1856, 239.
Synonyms: Thelephora actinaeformis B., Hook. Joum. Bot. 1856, 239.
Pterula trachodes (B.) Lloyd, Myc. Notes 60, 1919, 869, f. 1484.
Pterula actinaeformis (B.) Lloyd, I.e., f. 1485.
-5 cm. high, rigid, pale yellow (? dry), terrestrial: stem 1-2 cm.X2-2·5 mm., rather
stout, dividing into 2-several branches, sometimes confluent below: branches rather
dense, dichotomous, rather irregularly branched, multifid with flattened branching
below, tips subulate (not filiform): fibrillose-coriaceous when dried: Brazil.
Hyphae 3-5(-7)µ wide, monomitic, without clamps, the walls thin or variously
thickened 0·5-2µ, scarcely darkening in dilute alkali.
I have examined the types of both species in the Kew herbarium. They closely
resemble A. dendroides in general appearance and hyphae, but I could find no spores
in confirmation: indeed, the specimens seem completely sterile, without trace of
hymenium, and thus like the type of Thelephora amboinensis (see under A. dendroides).
Compare A. aurantiaca.
A. tropica (Mont.) comb. nov.

Basinym: Pterula tropica Mont., Crypt. Guy. 1855, no. 431: Syll. Crypt. 1856,
no. 600: Lloyd, Myc. Notes 60, 1919, 869, f. 1483.
(Pterula acutissima Mont. ined. et Clavaria acutissima Mont. ined., Crypt. Guy.
no. 979B, leg. Leprieur no. 895, in Herb. Paris.)
3-4 cm. high, deep yellow (tawny, Mont.: ? dried), branched almost from the base:
branches mostly dichotomous, divaricate, fine, with rather long filiform tips, drying
horny and twisted.
On the ground: South America (French Guiana).
Spores 6-8X5-6· 5µ, white, smooth, subglobose, thin-walled.
Basidia 40-50X9-10µ: sterigmata 2-4: hymenium -200µ thick: no cystidia.
Hyphae 2-4µ wide, monomitic, without clamps, thin or slightly thick-walled.
This is near A. dendroides, but has much finer branches and smaller spores. I have
examined the type (Leprieur 18, ann. 1850, in Herb. Paris, <let. Pterula tropica) and
the collection of the unpublished Pterula acutissima (Leprieur no. 895). The type is
sterile, but has exactly the macrosopic appearance and hyphal construction of Leprieur
895 ('P. acutissima'), which has abundant spores and basidia and from which, therefore,
I have drawn the microscopic description. The absence of skeletal hyphae in both
collections proves that the species belongs to Aphelaria.
Saccardo's record of P. tropica from Malacca (leg. Scortechini) may be A. dendroides.
192 APHELARIA
A. tuberosa (Grev.) comb. nov.

Basinym: Merisma tuberosum Grev., Scot. Crypt. Fl. 3, 1825, t. 178.
Synonyms.: Thelephora tuberosa (Grev.) Fr., El. 1828, 167.
Stereum tuberosum (Grev.) Mass., Brit. Fung. Fl. 1, 1892, 130.
Thelephora contorta Karst., Not. Sallsk Faun. Fl. Fenn. Forh. 9, 1868, 368: Bourd.
et Galz., Hym. Fr. 1928, 82.
Polyozus contortus Karst., Finl. Nat. o. Folk 37, 1882, 122: Act. Soc. Sci. Fenn.
15, 1888, 185, t. 8: Finl. Nat. o. Folk 48, 1889, 394: Lundell and Nannfeldt, Sv.
Svamp., fasc. 3-4, 1935, 28.
Lachnocladium semivestitum B. et Br., Grev. 1, 1873, 161 (haud Clavaria semivestita
B. et Br. 1874): Burt, Ann. Mo. Bot. Gdn. 6, 1919, 271, t. 5, f. 4: Coker, Clav. U.S.
Can. 1923, 196, t. 78-80, 90: Heim, Puhl. Junt. Cienc. Nat. Barcel. 15, 1934, 44, f. 9.
Clavaria gigaspora Cotton, Naturalist 1907, 97: Tr. Brit. Myc. Soc. 6, 1919, 33.
? Clavaria rivalis Britz., Hym. Stidb. 6, 1891, 33, f. 49: Rev. Hym. 1909, 221.
Text-Fig. 61.
-6·5 cm. high, solitary, gregarious, or caespitose, with flattened branching (apparently
in one plane though actually, by twisting, in several planes), the lower branches broadly
palmately divided, the upper narrow and dichotomous, pale buff-straw, grey or drab
tinged yellow, tips white, tough.
Stem -2 cm.X1·3-3·2 mm., distinct or not, often divided from the base into the
main palmate branches: branches 3-5-chotomous, or more below, dichotomous and
slightly flattened distally, numerous and bushy or rather few and lax, often irregular:
hymenium apparently covering the whole fruit-body except the tips: flesh tough,
subcoriaceous, especially in the stem: without smell or taste.
On the ground in the forest, or among grass, moss, or on bare soil in the open:
Europe (Sweden, Finland, Gt. Britain, Spain): temperate N. America (N. Car., N.Y.,
Pa., U.S.A.): temperate Asia (China, Japan'): Bermuda.2
Spores (12-)14-20(-24)X(4-)5-7(-<))µ,, white, smooth, elongate ellipsoid, sub
fusoid or subcylindric, blunt or subacute, attenuate to the oblique apiculus, granular
vacuolate or minutely guttulate.
Basidia dimorphic: (i) 50-90X8-10µ, (Bourd. et Galz.), 50-70Xu-15 µ, (auctt.),
clavate, with 2-3-4, mostly 4, normal sterigmata 7·5-10µ, long: (ii) 30-60X 13-18 µ,,
sub-Tremellaceous, 2 -4-lobed at the apex, each lobe prolonged into a stout sterigmata
-18X3µ, and often septate about the middle (Bourd. et Galz.).
Cystidia none.
Hymenium thickening to 200µ, (Coker, for C. gigaspora).
Hyphae 2-5µ, wide, cylindric, not inflated, walls slightly thickened, long-celled,
clampecl at every septum (Corner, for C. gigaspora), occasionally clamped (Bourd. et
Galz.), without clamps (Coker).
I have placed this very odd species in Aphelaria because one would look for it in
the neighbourhood of A. dendroides, were it not for the strange basidia ·intermediate
between those of Aphelarja and Tremella. Donk assures me that it belongs to
Tremellodendron, but in some living specimens which I recently examined (from
Cambridge, England, Sept. 1946) I found only one kind of basidia, as shown in
Text-Fig. 61: they were 70-uoX12·5-�4µ. with 2-4 sterigmata 6-10X4-5 µ,. The
immature basidia were normally clavate until the emergence of the stout sterigmata
caused them to become depressed at the apex and to appear shortly longitudinally
(but incompletely) septate. The species certainly needs fuller investigation because
it will be remarkable if there are two kinds of basidium producing the same kind
1
Lloyd, Myc. Notes 1921, 1079, as Polyozus contortus, but needing confirmation.
2
Burt, Ann. Mo. Bot. Gdn. 7, 1920, 115: the collection N.Y. Bot. Gdn. Herb., No. 508
(Church Cove, Bermuda) placed under Stereum proliferum, but with spores 13-16 X 6-'7 µ,
surely belongs to A. tuberosa.
APHELARIA 193
of spore and because it seems to link Aphelaria with Tremellodendron. The species
is widespread in the north temperate region, though rather rare. Cooke's record from
Queensland needs verification, for it does not seem to be tropical.
In using Greville's specific epithet I have followed Coker's suggestion. I think
there can be no doubt that the fungus described and illustrated by Greville is the
same as Polyozus contortus which Coker and Lundell have shown to be the same
as Lachnocladium semivestitum. Greville described the spores as oval and smooth (the
'
!'
�.. ..
······· . .....:i
\·.:·:.i:.:!
>..:····-.:�::
TEXT-FIG. 61. Aplrelaria tuberosa, X 1,000.

spores of true Thelephoras, as T. palmata, Greville described as globose), and he
said the hymenium covered the whole plant except the stalk. Therefore his species,
Merisma tuberosum, cannot be Stereum or Podoscypha, which have the upper sides
of the lobes sterile, and the spores prevent it from being Thelephora: Berkeley seems
to have originated the mistake that M. tuberosum had 'hymenium inferior' (Outl.,
p. 267), and doubtless he was copied by Massee whose spore-data (7-8 X 5 µ,, elliptic,
colourless, smooth) were probably as fictitious or erroneous as most of his authority.
M. tuberosum (Scotland) has never been recognized in Great Britain, and Clavaria
gigaspora (England), which Coker has shown to be identical with L. semivestitum,
is also a rare fungus in England. The only discrepancy in Greville's description is
the appearance of the fruit-body like a Thelephora with incised infundibuliform
pileus. Comparison of Karsten's figure of Polyozus contortus and Coker's photographs
of L. semivestitum show that the broad palmatifid lobes of the primordial fruit-body
S1l9
194 APHELARIA
by twisting and curling round of the sides give the appearance of a bunth of flattened
Clavarioid branches arising from a falsely infundibuliform pileus, as drawn by Greville.
Indeed, the shape of the fruit-body, especially when subsessile and young, is so
Thelephoroid that i�s true nature as a Clavarioid fruit-body with flattened branching,
primarily in one plane, escaped Greville, Fries, Karsten, and Lundell. There appears
to be no Grevillean specimen and, therefore, judging from his description of M. tube
rosum, I agree with Coker and Bresadola (cited by Coker) in their interpretation of it
as L. semivestitum.
Although the colour of the fruit-body is generally pale yellowish, greyish, or drab,
it seems that it may be brownish. Karsten described P. contortus as pale ferruginous
and Greville described M. tuberosum as grey with a slight brownish tinge.
Possibly Cla�aria regularis (p. 264) and C. rivalis' are forms of this species with
abnormally large, or germinating, spores.
The generic name Polyozus Karst., with P. contortus as the type-species, is a later
homonym of Polyozus Willd. (1793, Rubiaceae) and will become a synonym of
whatever genus A. tuberosa is eventually placed in.
TABLE VII. SPORE-SIZE IN APHELARIA TUBEROSA
Spore-size in µ. E• average Country Authority

14-18 X 5-6 2·9 Finland Karsten (type)
(10--)12-16(-20) X 7-9 1·8(-2·0) Gt. Britain Cotton and Wakefield (for
C. gigaspora)
13·5-18 X 6·5-8·5 2·3 " " Coker (for C. gigaspora)
14-19 X 6--7·5 2·4 " " } Corner (for C. gigaspora)
14-23 x6·5-8·0 2·6 " "
• 15-19 X 5--'7 2·8 France Bourdot and Galzin
:15-24x5-6·3 3'4 Spain Heim
17-2zx5·5--7 3·1 Sweden Coker
'I 6-24 X 6--7 3·1 " Coker (as Thelephora tube-
rosa)
15-18·x 6-6·8 2·4 U.S.A. (N. Car.) Coker (4514)
i5-20·2 X 4·8-6 3·3 " " " (2789)
1.5 X 6·6 2·3 " " " (4620)
14-15·2 X 5·2-6 2·6 " " " (4626)
17-22 >< 5·5-7 3·1 " " " (4631)
13-16·7 X 4-6·6 2·8 " " " (5270)
II-i9.X-4-5·5 3·2 " " " (5304)
15-20 X 5-7 2·9 " (N.Y.) " (Atkinson, Ithaca)
• E = length-;-width.
ARAEOCORYNE gen. nov.

(Gr. areios, slender)
\
Fruit-body simple, erect, small, slender, with sterile truncate apex, white: stem
distinct: flesh rather brittle, soft.
On dead �!ant-remains: I sp., tropical Asia.
Spores white, smooth, broadly and obtusely ellipsoid, aguttate.
Basidia with 4 sterigmata.
Cystidia none.
Hymenium thickening: subhymenium composed of uninflated hyphae.
' Clavaria rivalis, I.e.: -6 cm. high, branched, white or whitish, with yellowish or greyish
spots when moist: stem 4 mm. thick: branches short, clavate, 'tuberiformly' divided: spores
16-18 X 8-10 µ, white, asperulate, broadly ellipsoid, slightly curved, subattenuate at one end:
on the ground in woods, Germany (Bavaria).
ARAEOCORYNE 19 5
Hyphae monomitic, mostly inflating strongly, clamped, thin-walled, not secondarily
septate.
Sterile apex devoid of basidia.
TEXT-FIG. 62. Araeocoryne elegans, X 5: diagram of the structure of the fruit-body.
A. elegans sp. nov.

Text-Figs. 62, 63.
8-15 mm. high, simple, erect, solitary or gregarious, superficial, white: head
2·5-7 X 1-2 mm. at the apex, narrowly clavate-obconic, rarely subventricose, the apex
more or less abruptly truncate, often with a small papilla, sterile and minutely radially
silky-fibrillose: hymenium covering the sides of the head, abruptly delimited from the
stem, smooth and even or with 2-9 faint longitudinal wrinkles towards the apex, pale
cream to pale buff: stem 5-9·5 X 0·5-0·7 mm., slender, cylindric, straight or subflexous,
cften incurved, glabrous or sparsely puberulous, white, base attached by a thin, minutely
byssoid and slightly spreading, pallid white mycelial disk: flesh solid, fleshy-brittle, not
the least tough, white: smell none.
On dead leaves and rotten twigs in the forest: Malaya (Johore, near Mawai, in
swamp forest, 25.1.J1).
Spores 10-12 X 6-8 µ,, white, smooth, ovoid or ellipsoid, blunt, flattened adaxially,
with a stout, subcylindric apiculus, thin-walled, contents rather coarsely granular
vacuolate, not guttate: mature detached spores, and occasionally those on the sterig
mata, sending out several fine, conical or subcylindric, straight or subflexous processes
-5 X 0·5-1 µ,, thin-walled, the spore-contents becoming progressively vacuolate and
the dense cytoplasm passing into the processes.
Basidia 35-47 x 9-11 µ,, clavate, granular-vacuolate: sterigmata 2-4, 4-4·5 X 1 ·5-
2 •5 µ, at the base, the apiculus of the spore seated directly on the sterigma (not
obliquely).
Cystidia none.
Hymenium thickening slightly 35-60 µ, , with included spores, abruptly delimited
at the stem-apex: subhymenium c. 20 µ, wide, composed of narrow interwoven hyphae
3-5 µ, wide, with short cells 9-20 µ, long.
196 ARAEOCORYNE
Hyphae of the head 8-40µ wide (-25 or 30µ, only, in small fruit-bodies), the cells
75-220µ long, thin-walled, clamped, colourless, longitudinal but rather loose· and
fibrillar, with numerous narrow, 3-6µ wide, hyphae interweaving and longitudinal,
the cells regularly inflated and cylindric, or with irregular ampullaeform swellings,
occasionally with a short clavate branch as in the stem.
Hyphae of the stem 8-28µ wide, the cells 25-100µ long, as in the head, but more
compact and less inflated, narrower and more compact, 3-4 µ wide, at the surface
of the stem but not agglutinated, looser towards the centre, with few or many narrow,
longitudinal, or interwoven hyphae: often bearing short, clavate, lateral, I-celled
branches, as blind ramifications: without crystals.
Caulocystidia -40 X 5--7 µ, as short, cylindric, thin-walled, smooth, vacuolate,
e '
Tl!xT-FIG. 63. Araeocoryne elegans: basidia, spores, and three

germinating spores, X 1,000.
scattered processes, appearing as sterile basidia at the apex of the stem and giving
place abniptly to the hymenium.
Mycdial hyphae, at the base of the stem, 2-3 · 5µ wide, with firm, scarcely thickened
walls, clamped.
Sterile apex of the head composed of loosely interwoven hyphae more or less
appressed to the surface, convergent and denser towards the centre or papilla of the
disk, the end-cells of the hyphae 30--70 X 4-15 µ, more or less wholly inflated like the
cells of the flesh, obtuse and cylindric, not modified into basidia.
CARIPIA O.K.
Rev. Gen. Pl. 3, 1898, 451.
Synonyms� Hypolyssus B., Hook. Joum. Bot. I, 1842, 139: (non Pers. 1822 = mis
formations of agarics): Martin, G. W., Mycol. 30, 1938, 440.
Perona Fr., Summ. Veg. Scand. 1849, 333: (non Pers. 1825 = Delicatula Fayod,
1888).
[Podostrumbium Kz. apud Lev. in D'Orbigny, Diet. 8, 1849, 491: nomen invalidum.]
[Heringia Schw. ex B. et C., Exs. Fungi 279, ms: nomen invalidum.]
Fruit-body simple, small erect, turbinate obconic, apex discoid and sterile, solid, with
a distinct short stem; hymenium covering the sides of the obconic head, sharply
delimited, smooth or slightly longitudinally rugulose or plicate: flesh firm, drying hard.
CARIPIA 197
Lignicolous: 1 sp., tropical America.
Spores white, smooth, ellipsoid, aguttate, with a very thin hyaline outer membrane.
Basidia small, subclavate, 4-spored: hymenium thickening.
Cystidia thin- or slightly thick-walled: no gloeocystidia.
Hyphae becoming thick-walled, clamped, slightly inflating, monomitic.
Type-species: C. Montagnei.
The reason for the use of Caripia instead of Perona or Hypolyssus is given on
page 34. The genus was practically unknown until Martin redescribed the type
species, which appears to be common in the American tropics. It has never been
reported from the Old World tropics, where it is safe to assume that it does not occur.
However, there are still several points which must be examined in greater detail
before the genus can be correctly classified. It seemed to me, when writing this
monograph, to be a Clavarioid fungus resembling
Araeocoryne, though with very different hyphae:
those of Caripia are thick-walled and little inflated
(as in Lentaria), those of Araeocoryne thin-walled
.
and very strongly inflated. But, as noted on p. 35, 00
I have since studied living specimens and dis
covered that Caripia has a rudimentary pileus, so
that it should be classified, as hitherto, with Can
oo
tharelloid fungi, Araeocoryne having no marginal
growth to form a pileus (see p. 33). Martin's
description suggests that the fruit-body is endo
genous, which, if true, introduces a new element
into the morphology of Cantharelloid or Craterel
loid fungi. The one Clavarioid fungus with any
suggestion of a veil is Clavariadelphus pistillaris,
the primordium of which develops in the centre of
a small floccose mycelial ball, and such a veil, if it
can be so called, is very different from the caducous
white tomentum which covers the young fruit-
bodies of Caripia. TE�T-FIG. 64. Caripia Mon�afnei:
Clavaria verpiformis, from Brazil, may be allied. frutt-�ody, X 2: spores, bastdtum,
·
· 1e k nown and the type-specimen , cyst1d1a and hypha X 600: (after
It 1s
· very 11tt m the Killermann an'd Mart'm) ·
Kew herbarium is fragmentary. It seems to have
consisted of subgelatinous hyphae and to have no sterile top: yet in form it is similar.
C. Montagnei (B.) 0.K.

Basinym: Hypolyssus Montagnei B., I.e., 1842: Kew Bull. 1934, 251: Martin, I.e.,
f. 29-34.
Synonyms: Hypolyssus Sprucei Mass., Grev. 20, 1891, 33 (fide Martin).
H. foetidus Mass., Journ. Bot. 30, 1892, 197, t. 325, f. 3-5: (fide Martin).
Perona Montagnei (B) Fr., I.e., 1849.
Text-Fig. 64.
5-20 mm. high, 3-6 mm. wide across the apex, arising from a thick, irregular,
ochraceous subiculum, turbinate or urn-shaped, obconic, the apex sterile and convex,
then slightly depressed, when young wholly covered with a white tomentum falling off
and revealing the ochraceous top, brown stem, and creamy hymenium: stem slender,
central, sterile, usually shorter than the head, becoming brown, often with a ring of
mycelium at the base: hymenium covering the sides of the head, more or less abruptly
delimited, smooth or becoming longitudinally rugulose or sublamellate, becoming dingy
white, avellaneous or cinnamon drab with age: flesh firm, drying hard.
On sticks and rotten wood, also on aerial roots of orchids, with conspicuous
198 CARIPIA
mycelium: tropical America (Mexico, Honduras, Panama, Guiana, Bolivia, Brazil,
Trinidad), common.
Spores 5-6 X 3-3·5µ, white, smooth, ovoid to pip-shaped, with a very thin hyaline
outer wall (? aguttate).
Basidia c. 30 X 6 µ, narrowly clavate: sterigmata 2-4, c. 5 µ long.
Cystidia c. 23 X 5·5µ, thin-walled, fusiform, abundant, a few vesicular (-9µ wide)
and slightly thick-walled, both kinds immersed in the sub-hymenium and not
protruding, not encrusted.
Hymenium and subhymenium 100-uoµ thick, dense and abruptly delimited from
the flesh (the hymenium thickening and the cystidia initially hymenial, secondarily
immersed).
Hyphae 7-10µ wide (-15µ in places), with thick irregular walls (1-3µ) and narrow
lumina, clamped, intricately interwoven, irregularly branched: 2-5 µ wide in the
stem, subparallel, siightly thick-walled, with scattered sterile basidia on the surface:
2-4µ wide, appressed and levigate on the sterile top, the walls thin or slightly
thickened.
H. foetidus and H. Sprucei are synonyms, according to Martin, who says that the
spores ascribed to them by Massee (3-4µ, globose) belong to a Eurotium.
CERATELLOPSIS Konr. et Maubl.

..,.
le. Sel. Fung., t. 6, 1937, 502.
Synonym: Ceratella Pat., Hym. Eur. 1887, 157: Ess. Tax. 1900, 49 (non Ceratella
Hook., f. 1845).
Fruit-bodies filiform, very small, simple, in a few species sparingly branched,
0·5-10 mm. long, with a conspicuous, acute, sterile apex, eventually often fertile over
the apex in old specimens : stem very short or none: rather fibrillose-tough, with waxy
hymenium: without sclerotium.
On various plant-remains: north temperate, c. 20 spp.
Spores white, smooth (rough in C. dryopteridis), ellipsoid to globose, aguttate or,
in a few species, guttate.
Basidia 2-4 spored, small.
Cystidia generally absent, present in a few species.
Hymenium continuous or discontinuous, not thickening, generally absent from the
extreme base of the fruit-body, in many species eventually covering the apex on
cessation of growth: subhymenium slightly developed or practically absent.
Caulocystidia mostly absent.
Hyphae monomitic, not inflating, septate with clamps, not secondarily septate, thin
or slightly thick-walled, longitudinal, the cells rather elongate, not agglutinated or
mucilaginous.
Lecto-typus: C. aculeata Pat.
My description of this genus is based on C. aculeata ( = Pistillaria aculeata) and
the species which appear to resemble it. They have very small, simple or subsimple
fruit-bodies which are distinguishable from the corresponding species of Pterula
only by the· simpler hyphal construction. In this they resemble Lentaria and
Aphelaria (which is, typically, without clamps), so that the species of Ceratellopsis
may stand to these genera as Pistillaria and Typhula to Clavariadelphus. Probably
several of the species which I have included may be Pteruloid, e.g. C. Richii, C.
asphodeli, C. Helenae, C. rosella, C. tremula, and C. Queletii, the first two identical,
perhaps, with Pterula gracilis. C. dryopteridis is peculiar in the rough spores, but also
suggests P. gracilis. Their correct classification must await further study.. C. aculeata
shows, however, that a generic name is needed for the very reduced, monomitic,
uninflated fruit-bodies which have been confused with Pistillaria. Ultimately they
CERATELLOPSIS 199
may be related, subgenerically, to other genera of branched fruit-bodies as the simple
species of Pterula.
The lignicolous species differ from Mucronella in the direction of growth of the
fruit-body which is never positively geotropic but, probably, negatively geotropic,
unless predominantly phototropic.
From Pistillaria, Ceratellopsis differs in the absence of inflation of the hyphae,
the absence of a distinct stem, the absence of agglutination of the superficial hyphae
in the stem-like region (when present), and in the filiform shape, which expresses
the absence of inflation as well as the lack of subhymenium and intercalary expansion
of the hymenium.
KEY TO THE SPECIES OF CERATELLOPSIS

(Compare Pistillaria and Pterula)
Sp. rough, 9-12X5-6 µ,: on D ryopteris: Japan. C. dryopteridis
Sp. smooth
Cystidia present, thin-walled: basidia 2-spored: 1-3 mm. high,
sessile
Sp. 6 x 3 µ,: on stems of Mulgedium: France C. Queletii
Sp. 9·5-12X4·5-5·5µ,: on Asparagus-stems: Holland C. Rickii
Sp. 8-10X 5 µ,: on Asphodelus-stems: Tunisia C. asphodeli
Without cystidia
Pink or pinkish
5-6 mm. high, simple or sparingly branched: stem 2-3 mm.
long: 2-spored: on vegetable remains: Europe C. Helenae
2-3 mm. high, caespitose: sp. 9-10X5-5·5µ,: on twigs of
Robinia: Germany . C. Sydowii
1-2 mm. high, often furcate: conidia among the basidia or
on separate brownish tubercles: 4-spored: on plant-
remains: Europe C. rosella
White or pale yellowish
Sp. more than 8 µ, long
(Sp. 15-17 x 7 µ,: on wood: Holland Pterula rigida)
Sp. 9-15X5-6 µ,: 1-3 mm. high, on Equisetum: France C. equiseticola
(Sp. u-15 µ, long: -15 mm. high, filiform: on leaves:
France Clavaria exilis)
Sp. 8-12X3-3·5 µ,
3-5 mm. high: 2-spored: on Trichosanthes-fruits:
Italy C. tremula
1 mm. high or less: ? 4-spored
On twigs of Syringa: Italy . C. Carestiae
On bark of Lonicera: France C. caespitulosa
Sp. mostly less than 8 µ, long
Sp. very small, 3-4µ, long
-10 mm. high, simple or sparingly branched: sp. 3-4 µ,
subglobose: on bark of Thuja: U.S.A. . C. thujicola
-1 mm. high: sp. 2-3 x 1 · 5-2 µ,: on dead bark or pine-
needles: Europe C. acuminata
200 CERATELLOPSIS
Sp. larger, narrowly ellipsoid
0·5-2 mm. high: sp. 4-6 X 2-3 fL: on vegetable remains:
Europe C. aculeata
2-3 mm. high: sp. 6-7 X 4/L: on grasses: France C. graminicola
0·5-1 mm. high
30-40 fL wide: sp. 6-8 X 3-4 fL: on bark: France C. mucedinacea
90-120 µ, wide
Crystalline-encrusted: sp. 6-7 X 3·5 fL: 2-spored:
on mosses and dead leaves; France C. sagittaeformis
? not encrusted: sp. 4 ·5-6 X 2 · 5-3 µ,: 2-4-spored:
on Salix-wood: France. C. acuminata
sensu Bourd.
et Galz.
Sp. unknown
France: 3-5 mm. high, white: ? habitat C. Brondaei
Algiers: 2-3 mm. high, ochraceous: on dead leaves C. aciculata
U.S.A.: 2 mm. high, base subbulbous: on herbaceous stems C. mucosa
C. aciculata (Dur. et Lev.) comb. nov.

Basinym: Pistillaria aciculata Dur. et Lev., FI. Alg. 1846-9, t. 32, f. 4 (sine diagn.):
Sacc., Syll. 6, 1888, 759.
2-3 mm. high, ochraceous, filiform-acicular, tip acute and slightly curved, base
slightly thickened, smooth, glabrous.
On dead coriaceous leaves: Algiers.
Very doubtful.
C. aculeata (Pat.) comb. nov.

Basinym: Pistillaria aculeata Pat., Tab. An. 1883, No. 58: Corner, Tr. Brit. Myc.
So� 19, 1935, 285.
Synonyms: ? Clavaria microscopica Malb. et Sacc., Mich. 2, 1879, 42.
? Pistillaria gracilis sensu Pat., Tab. An. 1887, No. 575.
? Pistillaria culmigena sensu Karst., Quel., Schroet.
Ceratella aculeata Pat., Hym. Eur. 1887, 157: Ess. Tax. 1900, 49.
Text-Fig. 65.
0·5-2 mm. X 40-90 µ,, white, simple, filiform, acerose, minute, tip sterile or becoming
fertile, superficial; stem 200-400 X 20-60 µ,, short, distinct, glabrous, not swollen at
the base.
On dead leaves (Pat.), on dead leaf-bases of Cladium mariscus (Corner): France,
England (common in the fens, July-Nov.).
Spores 4-6 X 2-3µ, white, smooth, ellipsoid, slightly flattened adaxially, contents
cloudy, aguttate (small, ellipsoid, pip-shaped, as figured by Pat.).
Basidia 8-16 X 3·5-5 µ,, subclavate: sterigmata 4, rarely 2, 3-6 µ long.
Cystidia and caulocystidia none.
Hymenium very variable in development, fairly compact or loose or merely as
scattered clusters of basidia, the apex persistently sterile or eventually covered with
basidia, occasionally the whole fruit-body sterile: subhymenium practically absent,
the basidia arising directly from the longitudinal hyphae.
Hyphae 1-2·5 µ, wide, monomitic, uninflated, thin- or distinctly thick-walled,
clamped or some of the thick-walled hyphae septate without clamps, cells 50-350 µ,
long, longitudinal and generally slightly spiral, not agglutinated, branching normal
or, in the thick-walled hyphae, very narrow at the origin: without crystals.
Mycelial hyphae 1-2µ wide, spreading over the surface of the substratum from
r
TEXT-FIG. 65. Ceratellopsis aculeata: a young fertile and a sterile fruit-body, x 500.
202 CERATELLOPSIS
the base of the fruit-bodies and becoming attenuate into fine, apparently homogeneous
threads 0·5-1JJ- wide.
At first sight this fungus seems to be constructed in the same way as Pterula gracilis
( = Pistillaria aculina), but closer inspection will show that it has no typical aseptate
skeletal hyphae coursing throughout the fruit-body as in Pterula. All the thick
walled hyphae of G. aculeata which I traced for more than 200 I-' in length were septate
with clamps and branched, or sparingly septate without clamps and branched. The
construction is the same as in Aphelaria, which, however, has typically no clamps.
The fruit-body develops from single hyphae as in Pterula gracilis or from a group
of hyphae, but there are few or no additions to the primordial shaft after its inception
and the width of the fruit-body depends mainly on the number of initial concrescent
hyphae.
C. mucedinacea and G. acuminata sensu Bourd. et Galz. may be the same as
C. aculeata, but their habitat is different, lignicolous and corticolous respectively.
For P. culmigena Karsten and Schroeter give spores 4-6 X 2 JJ-, not triangular-sub
globose 3-5 JJ, as interpreted by Lundell, so that I think they must have had specimens
of C. aculeata. Quelet gave the spores of P. culmigena as 6-7 JJ-, elliptic-cylindric,
which may refer to G. aculeata or G. graminicola, if there is any difference between
them.
Clavaria microscopica was referred by Patouillard (Tab. An. 1887, p. 29) to Pistillaria
aculina ( = Pterula gracilis of this work). Its description, particularly the short spores,
suggests G. aculeata:
1-1·1.5 mm. high, 6oJJ- wide, shortly attenuate from the base, filiform, gregarious
or scattered, 'sordide cameoalba': spores 6 X 4/L, h.yaline, ovoid: basidia shortly
conoid: on dead.stems of Juncus, France (near Rouen).
Pistillaria gracilis sensu Pat. has a larger fruit-body 2-4 mm. high and 2-spored
basidia, but it has the form of C. aculeata and the small spores 5 X 2 JJ-: it grew on
rotten herbaceous remains in the spring. Possibly it needs a new name, for it is not
P. gracilis B. and Desm. (see under Pterula gracilis).
Killermann records G. aculeata on twigs of Populus in Germany, with white fruit
bodies 2 mm. high, spores 5 X 2·5p., and narrow hyphae, 2-3p. wide, with granular
contents (Zeitschr. f. Pilzk. 81, 1934, 107, f.F). It may be a Mucronella, but the
illustration is crude.
C. acuminata (Fckl.) comb. nov.

Basinym: Pistillaria acuminata Fckl., Symb. 1869, 31, t. 4, f. 39: Pat., Tab. An.
1887, No. 572.
Synonym: Ceratella acuminata (Fckl.) Pat., Hym. Eur. 1887, 157: Ess. Tax. 1900,
49.
Text-Fig. 66.
(a) 0·5-1 mm. high, very small, white, oblong with long, sterile apex, subventricose,
attenuate and very acute: stem distinct, very short, glabrous: hymenium compact.
Spores 2-3p. ovoid, white (Pat.).
Basidia 2-4-spored (Pat.): 1-2-spored (Fckl.): 8-u X 3·5-4p., 2-4-spored (Donk.).
On rotting pine-needles in wet places (Fckl.): on bark of Catalpa (Bignoniaceae)
(Pat.).
(b) C. acuminata (Pat.) sensu Bourd et Galz., Hym. Fr. 1928, 140.
0·3-0·8 x 0·1-0·12 mm., white, filiform, conical or lanceolate, acute, with very
distinct, sterile, hyaline tip at fir$t, becoming blunt and fertile over the apex: stem
0·12-0·2 mm. long.
On rotten wood of Salix viminalis: France.
Spores 4·5-6 X 2·5-3 /L wide, white, oboval.
Basidia 15-20 X 4 · 5-6p.: sterigmata 2-4, 3 /L long.
Hyphae 2-3p. wide, not agglutinated, parallel, thin-walled, without clamps.
CERATELLOPSIS 203
According to Donk's ms. notes, the spores on the type-collection of this species
are extremely minute, c. 3 X 1·7µ,, and a whole fruit-body measured by him had a
stem 75 X 50µ, and clavula 350 X 75µ,. Patouillard's interpretation of the species must
be accepted as correct, therefore, and that of Bourdot may refer,
as he suggested, to C. aculeata, though the habitat is unusual,
or to C. Carestiae.
C. asphodeli nom. nov.

Synonym:. Clavaria microscopica var. Asphodeli Pat., Cat. Pl.
Cell. Tunisie, 1897, 66.
1-2 mm. X 50-80µ, simple, gregarious or scattered, very
slender, subulate, acute, somewhat thickened in the middle, with
an indistinct spurious stem, white then pinkish, stem deeper pink.
On rotting leaves of Asphodelus: Tunisia.
Spores 8-10 X 5 µ,, white, smooth, ovoid.
Basidia 2-spored.
Cystidia ;zo-5 X 6-8µ,, abundant, elongate subfusiform.
This is probably a Pterula and perhaps identical withP.gracilis.
C. Brondaei (Quel.) comb. nov.

Basinym: ClavariaBrondaei Quel., Rev. Myc. 1892, t. 126, f. 3. TEXT-1:IG. 6�.
. · 1e, f ushorm-pomte
·,: ·
· d: stem d"1stmct, Ceratellops1sacum1nata:
3-5 mm. h1gh, wh"1te, s1mp
(from patom.11ard, Ta b.
short, 1 mm. long. An. n. 572).
France.
(Spores and habitat unknown.)
Unless there is a type-specimen, this species is unrecognizable.
C. caespitulosa (Sacc.) comb. nov.

Basinym: Pistillaria caespitulosa Sacc., Atti Congr. Bot. Palermo, 1902.
0·6-0·8 mm. X 100-130µ,, dingy whitish, fasciculate in groups of 4-10, cylindric,
obtuse; hymenium in the upper part of the fruit-body.
On dead bark of Lonicera periclymenum (Caprifoliaceae), France.
Spores 10-12 X 3µ,, white, smooth, oblong-ellipsoid, 2-guttulate.
Basidia 14-16 X 2 µ,, with minute, tooth-like sterigmata.
Hyphae narrow, longitudinal.
Saccardo remarked on the very short, tooth-like sterigmata and suggested that it
might be a stilboid fungus. The structure, nevertheless, seems ider.tical with
C. aculeata, unless the species is a Mucronella.
C. Carestiae (Sacc.) co�b. nov.

Basinym: Pistillaria Carestiae, Sacc., Malpighia rr, 1897, 17.
0·5-1 mm. high, caespitose, minute, cylindric, obtuse.
On dry twigs of Syringa vulgaris: N. Italy.
Spores 9-10 X 3-3·5µ,, white, ovate-oblong, smooth.
Basidia 15-20 X 4µ,: sterigmata 2-4.
This is like C. acuminata sensu Bourd et Galz. with longer spores.
C. dryopteridis (Imai) comb. nov.

Basinym: Pistillaria dryoptendis Imai, Tr. Sapporo Nat. Hist. Soc. 1·3, 1934, 386.
1-5 mm. high, white, gregarious or solitary, linear or filiform, smooth: stem not
distinct.
On dead fronds of Dryopteris dilatata: Japan (Sapporo).
Spores 9-12·5 X 5-6·5µ, white, rough, broadly ellipsoid, apiculate.
204 CERATELLOPSIS
Basiclia 5µ wide, subcylindric, 2-spored.
The rough spores suggest affinity with Clavaria Bourdotii (p. 265); in other respects
it recalls Pterula gracilis.
C. equiseticola (Boud.) comb. nov.

Basinym: Pistillaria equiseticola Boud., Bull. Soc. Myc. Fr. 1917, 13, t. 14, f. 5:
Bourd. et Galz., Hym. Fr. 1928.
1-3 X 0·3-0·5 mm. (100µ wide or less, Donk), acicular, straight or flexuous, white
then yellowish, not swollen at the base, fertile from the base upward, and eventually
over the apex.
On dead stems of Equisetum limosum, singly or 2-3 together: France.
Spores 9-10 X 4·5-5µ, white, smooth, oblong-fusiform, internally granular (Boud.):
10·5-15 X 5·2-()·2µ (Donk).
Basidia 12-18 X 7-8µ, pyriform (Boud.): 18-25 X 7-9µ (Donk).
Hyphae 2-4µ wide (Boud.): 2·5-4·8µ, distinctly thick-walled (Donk).
The larger measurements of spores and basidia are taken from Donk's ms. notes
on the type-collection in herb. Boud. at Paris. They suggest that the species may
be a very depauperate state of Pterula gracilis. Macroscopically it seems identical
with C. aculeata and C. graminicola.
C. graminicola (Bourd. et Galz.) comb. nov.

Basinym: Pistillaria graminicola Bourd. et Galz., Hym. Fr. 1928, 139 (= P. acumi
nata sensu Bourd. et Galz., Hym. Fr., No. 113, haud sensu Fckl.).
? = Pistillaria culmigena sensu Quel., Bull. Soc. Bot. Fr. 26, 1897, 233.
2-3 mm. high, white, then brown, filiform, flexuous, simple, with an acute, sterile
apex: base sterile, hispid with very short hairs like sterile basidia.
On dead grass-remains: France.
Spores 6-'7 X 4µ, white, smooth, oblong-oboval or subpyriform.
Basidia 12-18 X 4-5µ: sterigmata 2-4, 4µ long, filiform.
Cystidia none.
Hyphae 1·5-2·5µ.
This seems to differ from C. aculeata only in the slightly larger spores. Quelet's
interpretation of P. culmigena, with 6-'7µ ovoid spores, may
belong here.
C. Helenae (Pat.) comb. nov.
Basinym: Pistillaria Helenae Pat., Tab. An. 1883,·No. 57:
Quel., Ass. Fr. 1883, 507.
Synonym: Ceratella Helenae Pat., Ess. Tax. 1900, 49.
Text-Fig. 67.
5-() mm. high, white, flesh-coloured or pink, simple or often
sparingly branched, filiform-acute: stem 2-3 mm. long, short,
glabrous, darker pink, then bay or bistre: branches equal,
obliquely ascending, with acute tips, sometimes contort, simple
or once dichtotomous.
On dead plant-remains, gregarious or caespitose: France.
Spores white, ovoid-pruniform (? 3µ, Quel.: ? c. 5µ, Pat.).
TEXT-FIG. 67. Ceratel Basidia 2-spored (Pat.).
lopsis Helenae (from This is very probably a Pterula, but compare C. rosella.
Patouillard, Tab. An. n.
57). C. mucedinea (Boud.) comb. nov.
Basinym: Pistillaria mucedinea Boud., Bull. Soc. Bot. Fr. 1877, 308; Pat., Tab.
An. 1887, No. 571: Bourd. et Galz., Bull. Soc. Myc. Fr. 1910, 222.
Synonym: P. mucoroides Sacc., Syll. Fung. 6, 1888, 757.
CERATELLOPSIS 205
0·6-1X0·03-0·04 mm. (-2 mm. high, Pat.), very slender, minute, white: head
o· 5-0·8 mm. long, linear, fertile to the top, apex always fertile (? how it grows):
stem 0·2 mm. long, short, glabrous, hyaline, distinct.
In the fissures of bark of old trees: France.
Spores 6X1·5-2µ,, white, smooth, subcylindric (Bourd. et Galz.): 7-8x3-3·5µ,
(Boud.): 6X 4µ, (Sacc.).
Basidia 10-12X4µ,: sterigmata 2-4: hymenium discontinuous.
Hyphae 1-2µ,, only 10-20 longitudinal hyphae comprising the fruit-body and
arising from a small mucedinoid patch of mycelium.
I do not know whether the spore-measurements of Bourdot and Galzin are a
correction of Boudier's and Saccardo's, or whether they represent another fungus
macroscopically similar. The species must be very near C. aculeata and C. acuminata
sensu Bourd. et Galz. It is one of the most attenuate, depauperate, Clavaroid fungi.
C. mucosa (Berk. et Curt.) comb. nov.
Basinym: Typhula mucosa B. et C., Grev. 2, 1873, 18.
2 mm. high, simple, filiform, very acute, base slightly thickened, and bulb-like:
on dead herbaceous stems: U.S.A. (S. Car., No. 3832).
This may be Ceratellopsis, Pterula, or a rudimentary Pistillaria.
C. Queletii (Pat.) Konr. et Maubl.

le. Sel. Fung., t. 6, 1937, 502.
Basinym: Pistillaria Queletii Pat., Tab. Ann. 1883, 22, No. 45:
Quel., Ass. Fr. 1883, 507.
Synonym: Ceratella Queletii Pat., Hym. Eur. 1887, 157: Ess.
Tax. 1900, 49.
Text-Fig. 68.
1-3 mm. high, white, sessile, cylindric or abruptly contracted at
the top, apex filiform or subobtuse, the sterile tip -1 mm. long,
fertile from the base.
On dead stems of Mulgedium: France. Q{)
Spores 6 X3 µ,, white, ovoid (Pat.).
Basidia 2-spored (Pat.).
Cystidia elongate (Pat.).
This is said to differ from C. acuminata in the absence of a stalk
and in the larger spores. The cystidia, 2-spored basidia, and
absence of stalk suggest a rudimentary Pterula, as P. gracilis, but TExT-FIG. 68. Cera
with smaller spores. tellopsis Qwfletii:
The form of Pistillaria aculina on dead leaves of Gunnera, with (from Patomllard,
.
spores 6-9 X4·5 µ,, mentioned by Bourdot and Galzin as possibly Tab An. n. 45).
Clavaria microscopica, may belong here (Hym. Fr. 1928, 138).
C. Rickii (Oud.) comb. nov.

Basinym: Mucronella Rickii Oud., Beih. Bot. Centralbl. 11, 1902, 525.
Synonym: Cnazonaria Rickii (Oud.) Donk, Rev. Niederl. Homobas. Aphyll. 2,
1933, 99.
1-3X0·07 mm. white, sessile, solitary or in groups of 2-6, subulate, tapered to the
sterile apex, straight or slightly curved, drying s�iff.
On rotten stems of Asparagus officinalis: Holland.
Spores 9·5-10(-12)X 4·5-5·5 µ,, white, smooth, ellipsoid, aguttate.
Basidia 22-30 X5-6·5 µ,: sterigmata 2, straight, 7'5-8 µ,.
Basidioles ( = cystidia) subacute, projecting to 12 µ,.
Hyphae 2·5-3 · 5 µ, wide, slightly thick-walled, sparsely septate, without clamps,
longitudinal: subhymenial hyphae 2·5-3·5 µ, wide.
206 CERATELLOPS IS
Except for the narrower spores, this seems identical with Pterula gracilis and, like
C. Queletii, it is almost certainly Pteruloid.
C. rosella (Fr.) comb. nov.
Basinym: Pistillaria rosella Fr., Epicr. 1838, 5_87: Quel., Ass. Fr. 1883, 506:
Keissler, Beih. Bot. Centralbl. Sond. Abdr. 29, 1912, abt. 2, 436.
Synonym: Clavaria rosella (Fr.) Karst., Finl. Nat. o. Folk 37, 1882, 178.
1-2 mm. high, rose-pink, subulate, acute or acuminate, often bifurcate, pruinose,
pellucid at the base; [stem -1 mm. long, short, dilated at the base, pellucid white,
(Quel.)].
On dead leaves of Pyrus malus: on leaves of Aquilegia vulgaris (Keissler): Europe. 1
Spores 8 µ,, white, elongate, ellipsoid, pruniform (Quel.).
'Oblong-ellipsoid, white stylospores often accompany or replace the basidia and
form, at the base of the stalk, globular tufts of chamois-colour' (Quel.).
var. ramosa Pat.
Tab. An. 1883, no. 53.
Lanceolate, simple or once furcate, -2 mm. high: sometimes arising from the
conidiophorous tubercles.
On dead stems of Paeonia: France.
Spores 5X2 µ,, white, ovoid.
Basidia 4-spored.
Conidia 10 x 3 µ, on conidiophores among the basidia or at the centre of brown or
alutaceous, sessile, subdiscoid tubercles accompanying the clubs.
It is difficult to see how this differs from minute C. Helenae unless in the formation
of conidia.
C. sagittaeformis (Pat.) comb. nov.
Basinym: Pistillaria sagittaeformis Pat., Tab. An. 1883, 26, n. 56: Bourd. et Galz.,
Hym. Fr. 1928.
Synonym: Pistillaria pusilla var. lanceolata, Quel., FI. Myc. 1888, 452.
0·5-1 x 0·09-0·12 mm., white then cream, fusiform lanceolate, simple or unequally
2-lobed, covered with crystalline incrustation, fertile over the apex, attenuate into a
scarcely distinct stem.
On mosses (Barbula, Hypnum), on dried leaves of Salvia, and dead leaves of
Populus, Ulmus: France.
Spores 6-7X3·5µ, white, ovoid, oblong: (6X3µ, Pat.).
Basidia 15-20X4-5 µ: sterigmata 2, 3 µ long.
Hyphae 2-5-3 µ.
Patouillard's illustration suggests a Pterula with rigid skeletal hyphae.
For var. Rubi.Killerm., see Pistillaria pusilla.
C. Sydowii (Bres.) tomb. nov.
Basinym: Clavaria Sydowii Bres., Hedw. 1896, 6x.
2-3 mm. high, caespitose, filiform, pale flesh pink, entirely covered by the hymenium
except the sterile attenuate tips.
On twigs of Robinia pseudacacia: Germany.
Spores 9-10X5-5·5µ, white, obovate-elongate.
Basidia 24-26X6 µ,. Hyphae 4 µ, wide.
Perhaps this is C. rosella.
C. thujicola (Kauffm.) comb. nov.
Basinym: Pistillaria thujicola Kauffm., Pap. Mich. Ac. Sci. Arts Lett. 9, 1929, 207.
8-10X0·5 mm. white, slender, simple and filiform or forked dichotomously once or
1
On dead leaves: U.S.A. (S. Car.): Berk. in Grev. 2, 1873, 18, needs confirmation.
CERATELLOPS IS 207
twice, entirely fertile except the minute sterile tips, eventually fertile over the tips,
slightly bulbillose at the base and attached by radiating white hairs.
On the inner side of loose bark of Thuja occidentalis: U.S.A. (Mich., Lake Superior).
Spores 3-4µ wide, smooth, oval, subglobose.
Basidia 20-22 X 4-5·µ, subclavate; sterigmata 2, rarely 3.
This seems to be a reduced Pterula or Aphelaria, unless it is related to Lentaria
mucida. Compare Deflexula.
C. tremula (Sacc.) comb. nov.

Basinym: Pistillaria tremula Sacc., Syll. 17, 1905, 202 (ut subsp. nov. Pistillariae
Ferryi = Pterula gracilis).
3-5 mm. X 80-1ooµ,. white, very slender, filiform, often 2-4 fasciculate, wholly fertile
except the short sterile apex and, sometimes, the extreme base, tremulous, twisting
on drying.
On rotten fruits of Trichosanthes anguina (Cucurbitaceae): Italy.
Spores 8-II X3-3·5µ, white, smooth, obovoid, often 1-guttulate.
Basidia 18-20 X 5-6µ, with 2 sterigmata.
Perhaps Pteruloid. Compare G. equiseticola and G. Carestiae.
CHAETOTYPHULA gen. nov.

(As Typhula in form but with bristle-like cystidia)
Fruit-body small, simple, with distinct head and sterile stem: waxy-soft: no
sclerotium.
Hymenium continuous, not thickening, covering the entire head, developing basi
petally from the apex of the primordial shaft downward, the head therefore without a
sterile apex at any stage: subhymenium well developed, short-celled.
On vegetable remains: tropical Asia, 2 spp. (one doubtful).
Spores white, smooth, thin-walled, aguttate or with a few minute guttulae.
Basidia 2-spored, occasionally 1-spored.
Cystidia thick-walled, projecting, the base deeply immersed in the subhymenium.
Caulocystidia like the cystidia but not so thick-walled, scattered.
Hyphae monomitic, inflating, thin-walled, without clamps, not mucilaginous, not
secondarily septate, agglutinated on the surface of the stem, not encrusted with crystals.
Type-species: G. hyalina.
I make this genus for G. hyalina. It is probably a common fungus and, having no
sclerotium, would be referred to Pistillaria, as I use it, or Typhula in the early sense
of Fries and Patouillard. The thick-walled cystidia, however, are distinct and when,
to this feature, the peculiar basipetal development of the hymenium is added, the
species presents a new generic type. Mature fruit-bodies look exactly like those of
Typhula with a white, shortly cylindric head, but whereas in Typhula the head begins
to form before apical growth of the primordial shaft has ceased, and the hymenium
develops acropetally in the normal way, in Chaetotyphula the formation of the
hymenium begins at the apex of the primordial shaft by the conversion of the tips
of the longitudinal hyphae into basidia and proceeds basipetally by the outgrowth
of hymenial branches from the superficial hyphae of the shaft, proceeding downward
or backward from their terminal cells. I think there is also slight basipetal extension
of the hymenium in some species of Typhula and Pistillaria, e.g. Text-Figs. 41 and
47, but it is not so extensive as to construct the entire hymenium. In Chaetotyphula,
moreover, as in Dimorphocystis, the cystidia arise mostly, if not entirely, from the
apical branches of the longitudinal hyphae and not as ordinary hymenial elements
substituting basidia.
The agglutinated surface of the stem suggests affinity or common origin with
208 CHAETOTYPHULA
Typhula and Pistillaria, and thus with Clavariadelphus, but it may be a homoplastic
development of some other, and unknown, Clavarioid line with thick-walled cystidia.
The fruit-body of C. hyalina seems able to mature at any stage in development
and hence its great variability in size. C. actiniceps is a doubtful species which
requires microscopic analysis before it can be classified certainly.
r
TExT-FIG. 69. Chaetotyphula hyalina, X 10.
KEY TO THE SPECIES OF CHAETOTYPHULA

(Compare Pistillaria)
-18 mm. high, mostly more than 2 mm., with stem: sp. ellipsoid,
8-13 X 4-6 p,: on dead wood and leaves: Malaya C. hyalina
-1·5 mm. high, ? no distinct stem: sp. subglobose, 8-12p, wide:
on stems of Pteris and Rubus: Ceylon . C. actiniceps
CHAETOTYPHULA 209
C. actiniceps (Petch) comb. nov.
Basinym: Pistillaria actiniceps Petch, Ann. R. Bot. Gard. Per. 7, 1922, 291.
-1·5 mm. high, 0·3 mm. wide below, o·6 mm. above {apparently clavate), white,
translucent, sometimes slightly lobed above, covered everywhere or sometimes only
in the upper part with upwardly directed or radiating spicules.
'TIDcr-FIG. 76. Chaetotyphula hyalina, x 1,000.
On dead stems of Pteris aquilina and Rubus: Ceylon (Hakgala).

Spores 8-12µ. wide, white, smooth, globose or broadly ovoid.
Basidia 13 X s µ., clavate: sterigmata 2, 8 x 2 µ., conical below, linear and curved
above.
Cystidia (spicules)-200µ. long, hyaline, thick-walled, generally smooth, occasionally
incrusted.
This may be Dimorphocystis (p. 400) or Chaetotyphula, or a new genus. The large
subglobose spores, even one of which must have a much greater volume than the
5119 P
210 CHAETOTYPHULA
basidium producing it, and the curious sterigmata suggest something unusual. Petch
does not say where the hymenium is, or whether there is a stalk. Presumably the
hymenium covers the whole fruit-body: it may be acropetal as in Dimorphocystis
or wholly basipetal from apex to base of the primordial shaft, as may conceivably
happen in diminutive fruit-bodies of C. hyalina.
TExT-F1c. 71. Chaetotyphula hyalina: spores and developing cystidia and

basidia, x 1,000.
C. hyalina (Jungh.) comb. nov.
Basinym: Typhula hyalina Jungh., Crypt. Jav. 1839, 32.
Text-Figs. 69-74: Plate 16.
0·25-18 mm. high, white, solitary or gregarious, simple, very variable in size: head
0·15-8 X 0·09-0·35 mm. puberulous, at least near or at the apex, opaque, cylindric,
blunt or subacute, without a sterile tip: stem 0·1-12 X 0·05-0·18 mm. pellucid, filiform,
puberulous, the base often slightly thickened 0·1-0·2 mm. wide-and brownish: waxy-
soft, solid: without sclerotium.
CHAETOTYPHULA 2II
Oil dead wood and bark of Ficus fistulosa and F. variegata, and on dead remains
of palms: Java, Malaya (Singapore).
Spores 8-10·5 X 4·5-5·5 µ, or 11-13 X 5-6·3µ on 1-spored basidia, white, smooth,
pip-shaped, blunt, slightly curved, with a prominent blunt apiculus 1 µ long, very
slightly thick-walled, aguttate, or with a few irregular minute guttulae.
Basidia 18-36 X 6-8·5µ, subclavate, cloudy-vacuolate, without clamps, not or
scarcely projecting (-5µ): sterigmata 2, 4-5 x
1·5--2µ, the first basidia on young fruit-bodies
often with only 1 sterigma 6-'] µ long: the
spores often set asymmetrically on the sterig�
mata.
Cystidia 4o-<)O X 5-12·µ, cylindric or sub
ventricose at the base, slightly tapered to the
blunt or subacute apex, walls 1-3·5µ thick and
hyaline or pale yellowish, occasionally very
slightly encrusted, the lumen linear except at
the base and the thin-walled, often collapsed,
apex, aseptate, stiff, unbranched, arising deeply
in the subhymenium and projecting to 40µ:
abundant or scattered, mainly in the distal part
of the head.
Hymenium not thickening, soon developed
over the apex of the primordial shaft and ex
tending basipetally: subhymenium 15-35µ
thick, composed of short, compact, interlocked,
uninflated cells 5-12 X 3-4µ, arising directly
from the longitudinal hyphae.
Caulocystidia like the cystidia but shorter,
not so thick-walled, rather more tapered and
narrower, scattered, or abundant at base of
stem, -']Oµ long at the base of the stem, -30µ
long near the apex, 4-'7 µ wide, the walls
0·5-1·5µ thick or scarcely thickened in the
upper part of the stem, arising as outgrowths
from the superficial hyphae, the cytoplasm of
the parent cell contracting in the longitudinal
hyphae and becoming cut off by 2-4 secondary
septa closely set in the apical part of the cell;
if formed from terminal cells, then the tip of
the terminal cell collapsing.
Hyphae thin-walled without clamps, not TExT-FIG. 72. Chaetotyphula hyalina:
caulocystidia, X 500.
secondarily septate, not encrusted or mucila-
ginous, often slightly constricted at the septa, colourless, the cells of the longi
tudinal hyphae 50-350 x 8-25µ, -500µ long in the stem, shorter, narrower, and
more branched next the subhymenium, short and often with brownish walls at the
base of the stem: head composed of homogeneous longitudinal hyphae (without a
core of narrow hyphae, and not hollow): stem as the head, but the superficial hyphae
3-5 µ wide, contiguous and slightly agglutinated, but the walls not thickened.
var. m icrocystis var. nov.

Cystidia 30-50 X 5-7 µ, the walls 1-2 µ thick, generally sparse and limited to the
apex of the head, rarely more than 20 per head, occasionally only 1-3, rarely none.
Caulocystidia -50 X 4-6µ, or not more than 35µ long, walls 0·3-0·5 µ thick, reduced
to thin-walled, incipient processes 5-12µ long at the apex of the stem.
212 CHAETOTYPHULA
On dead leaves of various trees and palms (Ficus variegata, Castilloa elastica, Areca
catechu, Pinanga, Ptychosperma, and others): Malaya (Singapore).
Spores 8-9x4-5µ. (on leaves of Castilloa), 9·5-10·5(-n)x5-5·5(-6)µ. (on Ficus
variegata and palms).
Development of the Fruit-body
The primordial shaft reaches 0·05-1·5 mm. long before apical growth is arrested
and the head develops. Subsequent growth in length is caused entirely by elongation
of the cells. The hyphal tips at the apex of the primordial shaft are 2-3 µ. wide and
'I'ExT-FIG. 73. Chaetotyphula hyalina: developing

caulocystidia with secondary septa formed at the
hyphal tip on retraction of the cytoplasm, X 1,000.
their subterminal cells are 5-12µ. long on delimitation: some of these cells must
therefore elongate thirty to forty times their initial length and, hence, the filiform
stem.
On cessation of upgrowth, the tips of the hyphae at the growing apex expand
slightly, becoming subclavate, and then grow outward to cut off 2-4 short subterminal
cells 5-'7µ. long, which branch freely, and their branches grow out and cut off short
cells in the same way. Some of the end cells, when they have cut off 1-2 cells, dilate
and grow out from the surface more rapidly than the others and become the cystidia:
the apex of the primordial shaft thus soon becomes puberulous. The transformation
of the hyphal tips marks the beginning of the hymenium, the further extension of
which proceeds basipetally from the apex of the primordial shaft (e.g. Text-Fig. 74).
Protrusions develop from the cells of the superficial hyphae progressively more
removed from the apex. They become septate and cut off 1-4 short cells which
branch freely and enlarge the incipient hymenium in the same way. The process
continues, apparently, throughout the life of the fruit-body and such rudimentary
hymenial branches or protrusions can be found at the base of mature heads of maximum
size. Eventually the end-cells mature into basidia and the short rows of 2-4 cells
form the subhymenium and continuously extend the hymenium by inserting new
branches. The cystidia, it seems, develop only from some of the earlier end-cells at
or near the apex of the primordial shaft and not, or rarely, from the later ones
CHAETOTYPHULA 213
developed by basipetal extension of the hymeniuin, and hence their occurrence

mostly in the distal part of the head. The cystidia arise superficially, as the basidia,
TEXT-FIG. 74. Chaetotyphula hyalina: early stages in the basipetal

development of the fertile head, starting with the apical cystidia:
the stem-hyphae beginning to inflate and the caulocystidia just
emerging, X 500.
but the continual outgrowth of the other hyphal ends round them cause them to
have embedded bases in the mature hymenium, as though they had grown up through
214 CHAETOTYPHULA
the mature hymenium from the subhymenium. Thus, in this feature as well as in
its manner of growth, the mature fruit-body presents an erroneous manner of develop
ment: the cystidia develop superficially, though they come to be partly or entirely
immersed, and the head develops basipetally, though it comes to be elongate as if
formed by continued apical growth.
When the normal fruit-body is 2-3·5 mm. high, the incipient hymenium is a
continuous covering over the apex and extending down the sides for 1-1·5 mm.
The first spores then begin to be shed from scattered basidia in all parts of the
hymenium, except the most basal, and as yet rudimentary, zone at the stem-apex.
There is no marked basipetal maturing of the basidiospores, which continue to ripen
all over the head.
The fruit-body grows to its full size by inflation, particularly elongation, of the
cells of the longitudinal hyphae, which accommodates also the intercalary growth
of the hymenium and prevents the head becoming hollow. The inflation proceeds
acropetally from the base of the primordial shaft before its apical growth has been
arrested. There is no secondary septation.
The caulocystidia do not arise until the hymenium has begun to form. They then
develop acropetally from the base of the stem as outgrowths from the superficial
hyphae. Those at the apex of the stem are smallest and least developed and may
become incorporated in the hymenium as it extends downwards.
No disk or support is formed at the base of the fruit-body, which appears held up
by the original mycelial hyphae.
Some fruit-bodies stop growing at a very early stage, ·so that they may be only
250µ. high and consist of a head 150 X 90µ. and a stalk 100 X 50µ.. From these micro
scopic, and uncommon, fruit-bodies there is every gradation to the largest, some
seventy times as long, most being about 10 mm. high with the head 5 mm. X 250µ.
and the stem 5. mm. x 120µ..
Mature fruit-bodies last for 2-3 days. They develop quickly, like epiphyllous
agarics, in the course of a few days, not like the slow-growing species of Typhula,
but I have no detailed observations on their rate of development.
Identification
Typhula hyalina was described thus: 2-3 mm. high, white, subhyaline, simple,
gregarious, erect, glabrous, very delicate: stem filiform, slender, subpruinose: head
cylindric, subdiscrete, as long as the stem: on trunks on Cocos nucifera: Batavia
(Java). It thus answers very well macroscopically to the species which I have described,
even to the 'subdiscrete' head, the basipetal growth of which prevents it from being
normally discrete as in Typhula. Moreover, this is the only common Typhuloid
fungus in Malaya, the fungus flora of which is almost identical with that of Java,
and it is therefore likely to be the species found by Junghuhn.
CLAVARIA Fr. s. str.

Syst. Myc. 1, 1821, 466: Hym. Eur. 1874, subgen. Syncoryne.
Synonyms: Holocoryne Bon., Handb. Myk. 1851, 166, emend. Donk (1933).
? Stichoclavaria Ulbr., Lindau Krypt. FI. f. Anf. 1, 1928, 3rd ed.
Fruit-bodies branched or, generally, simple, often caespitose, brittle, white, pink,
reddish, purple, violet, grey, or brown, rarely yellow: branching radial: stem distinct
or indistinct, noticeable in small fruit-bodies: hymenium extending over the growing
apices.
Terrestrial, in some cases associated with soil-algae: temperate and tropical: 24 spp.,
and 16 doubtful species.
Spores white or pink (C. helicoides), generally smooth, subglobose to cylindric ellipsoid,
CLAVARIA 215
thin-walled, generally aguttate or finely granular guttulate, often with vitreous
opalescent contents, rarely 1-guttate.
Basidia mostly 4-spored, without clamps (Syncoryne) or with a wide, loop-like
clamp (Holocoryne).
Cystidia absent or present as cystidioles.
Hymenium thickening or not, never unilateral: subhymenium well developed, the
hyphae not inflating, or slightly (C. fossicola, C. Zollingeri).
Hyphae thin-walled, inflating, without clamps, generally secondarily septate, compact,
narrow on the surface of the stem, not agglutinated.
Type-species: C. vermicularis Fr.
This original, enormous genus must now be defined from C. vermicularis Fr. as the
lecto-type. It thus represents the Friesian subgenus S yncoryne, excluding the yellow
species of the Clavulinopsis fusifarmis alliance. The main characters are the absence
of clamps from the hyphae of the fruit-body (I have not studied the mycelium), the
secondary septation of the hyphae (which gives the brittle, juicy texture), and the
colourless, aguttatt;, or finely multiguttulate spore. The one constant feature is
the absence of clamps from the medullary hyphae. But on the presence or absence of
a wide loop-like clamp at the base of the basidium, I divide Clavaria s. str. into two
subgenera, Syncoryne (without the clamp) and Holocoryne (with the clamp, and
having C. acuta as its lecto-type, Text-Figs. I I and 95). This character may prove
to be generic, but I have not examined enough species for certainty and it can hardly
be made out in dried specimens because of the contraction and crumpling of the
subhymenium.
Anomalous are the pink spores of C. helicoides, the colour of which is derived from
the pink cytoplasm of the basidium which is injected into the spores. The old basidia
of this species also become secondarily septate as in Clavulina. The spores of C.
vermicularis var. singaparensis have a large gutta, as in Clavulina and Clavulinopsis,
and this feature may occur in some forms of C. purpurea and C. Zollingeri: generally
there is no distinct gutta and characteristically, at \east of the subgenus Holocoryne,
the spore-contents are finely multiguttulate. In some species the secondary septation
is slight (C. filiola, C. helicoides) and even absent (C. fossicola, C. tenuipes), in which
case the fruit-bodies are often small; but in C. Gibbsiae copious primary septation
seems to make up for the lack of secondary septation, so that its fruit-bodies are as
large as, or larger than, those of the closely allied C. acuta, which has abundant
secondary septation.
The genus needs to be elucidated from living material.
C. ROSEA FR. (1821) AND C. INCARNATA WEINM. (1836)

Comparison of the spore-measurements given by,,recent mycologists for C. incarnata
and C. rosea shows that they fall into three groups which I will call the long-spored
species (7-1ox4·5-6·5µ), the subglobose-spored species (6-8x4-5·5µ), and the
narrow-spored species (5-8x 2·5-3·5µ) (Table VIII). The long-spored species has
been called C. rosea, C. incarnata, and C. rubella: the other two have been called
C. rosea. To all three the same macroscopic description applies, thus:
Small, simple, brittle clubs, rose-pink fading yellowish from the tip downward,
with a short white or pallid stem, the flesh paler concolorous, solitary or in small
clusters of two to seven: on the ground in fields and woods, Europe and N. America,
rather rare.
In all cases where the long-spored species has been distinguished (as by Rea,
Bourd. et Galz., Donk, and myself: see under C. incarnata), it is said to be pale
pinkish with a white strigose base to the stem in contrast with the vivid pink, rose
pink, or reddish pink of the other two species for which there is no mention of
such a base to the stem. The subglobose and the narrow-spored species appear
216 CLAVARIA
macroscopically identical. There is no type- or authentic specimen and the
illustration cited by Fries for C. rosea (Obs. Myc. 2, 1818, t. 5, f. 2), according to
Coker, might be either.
C. rosea was first described from Westergotland in Sweden (Dalman ex Swartz,
Acta Holm. 1811, 157): it was then illustrated by Fries (I.e.) and, as C. Swartzii, by
Dalman (Sv. Bot. 1818, t. 558). These names are 'pre-Friesian', but C. Swartzii was
given as a synonym of C. rosea by Fries in 1821. Now the only pink Clavaria of this
kind, so far known from Sweden, is the narrow-spored species, specimens of which
TABLE VIII. SPORE-SlZE INCLAVARIA ROSEA FR.AND c. INCARNATAWEINM.
I Author's Suggested
Spare-size in µ, Authority Country identification identification
7-1ox5-6 Donk Holland C. rosea C. incarnata
England
7-10X 5-6 Cott. Wakef. " " "
6·5-10X3·5-6·5 Comer " C. incarnata ,,
7-8X4·5-6 Bourd. Galz. France " "
7-1ox6-7 Rea England " ,,
7-1ox6-8 Cott. Wakef. " " "
5-8x4-6 Coker (Herb. Pers.) C. rosea C. rosea var.
subglobosa
6-8·5x4-5·5 " Baria, No. 1889, " C. rosea var.
Nice subglobosa
6-6·5X 5 Rea England " C. rosea var.
subglobosa
6-7x5 Ricken Germany " C. rosea var.
subglobosa
5-?'5X2·5-3 Coker Sweden C. rosea C. rosea
5-8X 2·5-3·4 " Colorado " "
6·6-7·5X 3·5 " New York " "
5-8X2·5-3·5 " England " "
(Hebden Bridge)
(6-7X2·5-3•5 Wehmeyer Michigan c. vermicularis ? C. rosea)'
8, globose Quelet France C. incarnata ? Clavulinopsis
miniata
10, pruniform " " C. rubella ? C. purpurea
'
1
Or Clavulinopsis luteo-alba.
were sent by Romell to Coker. Therefore Coker, on the grounds of type-locality,
decided to call the narrow-spored species C. rosea (1923). It would thus seem con
cluded that C, rosea was the deeply coloured, narrow-spored species and C. incarnata
the pale, long-spored species. Quelet, however, interpreted C. incarnata as a species
with 8µ, 1-guttate, globose spores, on the basis of a French collection: what this may
be has yet to be decided, for it may even be Clavulinopsis aurantio-cinnabarina or a
rare northerly stray of the tropical Clavulinopsis miniata. Also, Quelet interpreted
C. rosea (under the invalid name C. rubella Pers.) as a species with ovoid-pruniform
spores 10µ long: it is probably C. purpurea Fr. (see p. 248). Donk, also, has referred
the pale, long-spored fungus to C. rosea rather than to C. incarnata (1933, p. 86),
following the descriptions given by Cotton and Wakefield. It seems to me, however,
that in the absence of a type-specimen or critical illustration one must follow Coker,
who first attempted thoroughly to define C. rosea. Accordingly I use C. rosea, in his
sense, for the narrow-spored species and make the new variety subglobosa for the
subglobose-spored species which is without name and, macroscopically, seems
identical: and the long-spored species I retain as C. incarnata, following Rea, Bourdot
CLAVARIA 217
and Galzin. The descriptions of C. rosea, given by Cotton and Wakefield and by
Donk, I think refer macroscopically to C. rosea and microscopically to C. incarnata. 1
Whether C. Barlae belongs here, as branched forms, or to Clavulinopsis remains to
be seen.
NATURAL GROUPS OF THE KNOWN SPECIES OF CLAVARIA s. STR.

Subgen. Syncoryne: basidia without clamps: spores aguttate (rarely 1-guttate or
multiguttulate): hyphae generally secondarily septate: fruit-bodies branched or simple.
Subgen. Holocoryne: basidia with a wide loop-like clamp at the base (thus appearing
shortly furcate at the base): spores multiguttulate: hyphae secondarily septate or not:
fruit-bodies simple.
Syncoryne
Sp. with pink contents C. helicoides
Sp. white
With cystidia: ? always· under conifers C. nebulosoides,
C.purpurea
Without cystidia
Violet, purple, or lilac C.xylarioides,
C. Zollingeri
Red or pink C.rosea
Dark brown C.atrobadia,
C. atroumbrina,
C. Pullei
[Yellowish, discolouring brownish from the base: sp. sub
globose C. straminea]
Alutaceous, then fuliginous C.fumosa
White
Hymenium thickening: hyphae not, or rarely, secondarily
septate: phycophilous C. Jossicola,
? C. scabra
Hymenium not thickening: hyphae freely or sparsely
secondarily septate: not phycophilous . C. filiola,
C. vermicularis
(Probably also C. citriceps, C. Corbierei, C. Guilleminii, and C. subacuta)
(Note: C. straminea probably belongs to Holocoryne)
Holocoryne
Hyphae not secondarily septate: wholly white, drab, or greyish C. alliacea,
? C. Daulnoyae,
C. Gibbsiae,
C. tenuipes
Hyphae secondarily septate
Sp. globose or broadly ellipsoid: fruit-body white: hymenium
not thickening C. acuta
Sp. ellipsoid
White with yellow stem or wholly yellow: hymenium not
thickening (or very slightly) C. argillacea
Pale pink: hymenium thickening C. incarnata
1
Cotton and Wakefield give the spores of C. rosea as 7-10 X 5-6 µ, as in C. incarnata, .but
in the collection of Crossland's from Hebden Bridge, cited by them, Coker found the spores
to be 5-7·5 X 2·5-3 µ, as in C. rosea sensu Coker.
218 CLAVARIA
ARTIFICIAL KEY TO THE SPECIES OF CLAVARIA

(Dubious species, see p. 221)
Branched A
Simple
Violet, purple, or lilac B
Pink, red, or rufous . C
Brown or brownish black D
Greyish white, grey, fuscous, fuliginous, or blackish E
Yellow or with yellow stem, or brownish yellow F
White, cream, or isabelline . G
A. Branched fruit-bodies
White
Mostly simple, -12 mm. high: sp. 5-6·5X2-2·7µ,: trop. C. fossicola
Sparingly branched, -8 cm. high: sp. 5·5-7·5x3-3·7µ,: U.S.A. C. cretacea
Red or pink, mostly simple, -6 cm. high: sp. 5-6X3·5-5µ,: Europe C. Barlae
Amethyst, lilac, purple, vinaceous, or pinkish magenta, -8 cm. high:
sp. 4-7 X 3-5µ,: temp., trop. C. Zollingeri
B. Simple, violet, purple, or lilac

Sp. 5·5-9X3-5 µ, (some -15 X5 µ,), ellipsoid: caespitose, purple, or
brown or grey tinged purple: N. temp., under conifers C. purpurea
Sp. broadly ellipsoid or subglobose
Sp. 9-10 X 7-8µ,: violet-black, -4 cm. high: Ceylon C. xylarioides
(Sp. smaller: see C. Zollingeri)
C. Simple, pink, red, or rufous

Sp. yellowish pink, 9-14 X4-5·5µ,: -3 cm. high, pink, orange-pink,
reddish orange: on bare soil in the open: trop. Asia C. helicoides
Sp. white
Sp. 7-10X4-6·5µ,, ellipsoid-oblong: -8 cm. high, pale pink,
fading yellowish or greyish white from the tip, solitary or
gregarious: Europe C. incarnata
Sp. smaller or rounder
Pale lilac-pink or amethyst: sp. 5-6· 5X4-5 µ, C. Zollingeri
Coral-red or rosy pink
N. temp.
Sp. 5-8X2·5-3·5 /L C. rosea
Sp. 5-8 X 4-6 µ, var. subglobosa
Sp. 5-6 X4-5µ,: often sparingly branched C. Barlae
Africa: sp. 6-8X3·5-4µ,: pale cinnabar C. Schwein-
furthiana
(Sp. unknown: see dubious species, p. 221)
(Sp. 8-11 x 6-9µ,: pinkish with brown stem: U.S.A. Clavulina
castaneipes)
CLAVARIA 219
D. Simple, brown or brownish black

(Spores ellipsoid to oblong)
Hymenium more or less longitudinally rugulose
Sp. 5-7 X 4·5 fL, 1-guttate: -4 cm. high, dark brown:? 2-spored:
Holland C. Pullei
Sp. 4·5-6·5 X 2-3 fL, aguttate: -7 cm. high, umber then blackish
brown, soft, elastic: U.S.A. C. atroumbrina
Hymenium smooth
Sp. smooth
With cystidia: sp. 5-6 X 2·5 fL: -8 cm. high, wood-brown or
fawn: under conifers: Oregon C. nebulosoides
Without cystidia
Solitary, wholly pale brown: Tonkin C. tonkinensis
Caespitose
Reddish brown or rufous-bay, then blackish, brittle:
sp. 5 x 2-2·5 fL: smell faint, of meal: Europe C.atrobadia
Rufous-fuliginous, rather tough: sp. ?: Europe C. tenacella
Natal brown: sp. 4-5 X 3-3·5 /L: U.S.A. C. rufobrunnea
(Pale alutaceous then fuliginous, brittle: N. temp. C.Jumosa)
(Sp. subglobose, see E)
E. Simple, greyish white, grey, Juscous, fuliginous, or blackish

Caespitose
2-14 cm. high, pale yellowish or alutaceous, then fuliginous:
sp. 6x3·5 /L: N. temp. C.fumosa
3-5 cm. high, longitudinally striate, greyish bistre, then fuli-
ginous: sp. 3-4 X 2-3 fL: Europe C. striata
-4 cm. high, clear grey: sp. 7-9 X 5-6 fL: France C.Daulnoyae
Solitary or gregarious
Sp. subglobose
Sp. 8-10µ (? -15µ): -2 cm. high, rugose, cinereous: Brazil C.cinereo-atra
Sp. 7-10 fL wide: -5 cm. high, fuscous, fuliginous, or blackish:
Europe C. Greletii
Sp. 4-6 /L wide: -5 cm. high, avellaneous, then blackish: Japan C. avellaneo-
nigrescens
Sp. ellipsoid
Sp. 4-5 X 2·5-3 /L, pip-shaped: -3 cm. high, greyish white or
grey, with darker flesh: Gt. Britain C. Crosslandii
Sp. 6-r 2 X 4-5 · 5 fL: -6 cm. high, greyish or greyish yellow: on
the ground, sticks, or humus: Europe C. tenuipes
(See also C. Daulnoyae, above)
F. Simple, yellow, or with yellow stem, or brownish yellow

Sp. ellipsoid
Sp. 6-r 3 X 3·5-7 /L, multiguttulate: with yellow stem or wholly
yellow: N. temp. C. argillacea
Sp. 7-10 x 4-6 fL: 2-spored: white then pale yellow: stem white:
smell of garlic: N. temp. C. fuscata
220 CLAVARIA
Sp.globose or subglobose
Sp.5--J"J p, wide: -5 cm. high, straw colour then brownish with
age or on handling: stem cinnamon-yellow: Europe C. straminea
Sp. 4-4·5X3-3·5 µ,: -4 cm. high, white, becoming yellowish:
U.S.A. . C. citriceps
G. Simple, white, cream, or isabelline
15 mm. high or less, not caespitose: white
Trop.
Sp. 4· 5-5X4 p,: stem pellucid: soft, brittle: hymenium not
thickening: Malaya C.filiola
Sp.5--6·5 X2-2·7 µ,: simple or little branched: stem indistinct:
firm, subcartilaginous: hymenium thickening: phycophilous
on bare soil, in large troops: Malaya C.fossicola
Sp. ?: filifonn acuminate: ? phycophilous: Brazil C. scabra
Temp.
Fruit-bodies rather truncate-clavate: sp.less than 6 p, long
Sp.4 X2 •5 p,: soft, often compressed: France . C. Guillemin:i
Sp.3-4·5X3-4 µ,, subglobose: often umbilicate: France C. Corbierei
Fruit-bodies cylindric, acute or subclavate: sp. larger
Sp. 6-12X4-5·5 p,: whitish, drab, greyish, or yellowish,
often flattened: on the ground, sticks, and humus:
Europe C. tenuipes
Sp. II-15 p, long, elongate ellipsoid: cream-white, filifonn:
on the ground and on tan-bark: Europe C. exilis
Larger, caespitose or not
Smell of garlic: 2-spored
N. temp.: sp. 7-10X4·5--6 µ, C.fuscata
Malaya: sp. 9-u X9µ, C. alliacea
Not smelling of garlic: mostly 4-spored
Sp. 6-12X4-9 µ,, multiguttulate: basidia shortly furcate at the
base with a loop-like clamp
Trop. (New World?): sp. 5·5-8 µ, wide . C. Gibbsiae
Temp.
Often 2-spored: sp. 7-10X 5-9 p,: -8 cm. high, often
caespitose: on the ground C. acuta
Mostly 4-spored: sp. 6-12X4-5·5 p,: -6 cm. high, often
dingy: on earth, sticks, or humus C. tenuipes
(2-spored: white with brown stem: sp. 8-II x6-9µ,: U.S.A.: see Clavulina
castaneipes)
Sp.smaller, usually not guttulate: basidia without clamps
Stem 6-15 mm.long, pellucid
Bonin Is!. : sp. 5 X3 p, C. subacuta
France: sp. 5-8X3-4 f£ C. a/finis
Stem indistinct or shorter
Subsolitary, -4 cm. high, becoming yellowish: sp.
4-4·5X 3-3·5 µ,: U.S.A.. C. citriceps
Generally caespitose, 3-12 cm.high, sometimes ageing
yellow: sp. varying ellipsoid, 5-8· 5 X3--6 µ,, or sub
globose, 3-5 p,: very variable: cosmopolitan C. vermicularis
CLAVARIA 221
DUBIOUS SPECIES OF CLAVARIA

Branched
Europe (including Canary Is!.)
Pinkish white, spinous: on trunks: Poland C. aculeata
Rose-pink, fleshy: on roots of trees: Canary Is!. . C. rhodochroa
White, then pale ochre flesh-colour: slender, funbriately
digitate: sp. 8-9 µ long: in pine-woods: France C. fimbriata
Yellow: in a hot-house: Germany C. compressa
N. Amer.
Yellowish with red tips: on trunks: Ohio, Columbia C. Berkeleyi
White: Mexico C. polita
Much branched, fuscous: branches subulate: on rotten
trunks: Mexico C. fuscescens
S. Amer.
On Euphorbiaceous seeds: -1 I mm. high, subsimple: Brazil C. seminicola
Africa
Ferruginous ochraceous, rooting: branches flattened: S. Africa C. Cladoniae
Asia
White: Hong Kong C. decolor
White, caespitose, much branched: Kashmir C. Jacquemontii
Austral.
White, much branched from a massive stem: among leaves:
Queensland C. portentosa
White, much branched, cristate: in sandy soil C. plebeia
Simple
Europe
Flesh-colour, caespitose: in oak-woods: Sweden C. anomala
Greyish yellow, caespitose: (sp. 18-31 X 11-17µ): in woods:
Germany C. regularis
S. Amer.
Black: head ovate-clavate, rugulose-plicate: sp. 10 x 6µ: Chile C. nigricans
On Euphorbiaceous seeds: -11 mm. high, simple or sparingly
branched: Brazil C. seminicola
Subsessile, 2-4 mm. high: stem thick, short: head rugulose
plicate: white: on bark or wood: Brazil C. verpiformis
(p. 267)
Africa
Subrufous: on bare earth: Congo C. sibutiana
Pale cinnabar, broadly compressed C. Schwein-
Jurthiana
-r 5 x 0·3 mm. setaceous: mycelium membranous: on bark: C. Lecomtei
Asia, New Guinea
-30 cm. high, 5-7 mm. wide: on the ground: India C. Gollani
Small, ? pale brown: on the ground: Tonkin C. tonkinensis
-r r cm. x r mm., filiform, pale rufescent: sp. ? brown, 4 X 3µ:
on the ground: New Guinea C. subfistulosa
222 CLAVARIA
C. acuta Fr.
Syst. Myc. 1, 1821, 485.
Synonyms: C. falcata Fr., ibid. (fide Donk).
C. Jragilis Fr. sensu Rea (non vera = C. vermicularis).
C. filipes B. et Rav., Grev. 2, 1873, 17 (non sensu Coker= Clavulinopsis filipes).
Text-Fig. 75, Plate 2.
1-8 cm. high, simple, solitary or gregarious in small groups, or densely fasciculate,
shining white, very brittle: smell and taste none, or ill-defined: 1-3 mm. wide,
cylindric and acute, becQming compressed, hollow and obtuse, slender, delicate:
stem 0·5-2 cm. X 1-1·5 mm., translucent, usually distinct.
Among grass or on bare earth in fields and woods, or in flower-pots: Europe, Japan,
U.S.A., S. Austral. (Victoria): not common.
Spores 7-10 X 5-9µ, white, smooth, broadly ellipsoid to subglobose, thin-walled,
with granular-guttulate contents, the marked apiculus about 1 µ. long.
Basidia 25-4ox8-10µ, or 45-58x8-10µ, with 2-4 sterigmata 5-10µ long; with
a short, wide, loop-like clamp at the base.
Cystidia none.
Hymenium not thickening: subhymenial hyphae 1·5-2·5µ wide, interwoven.
'
Hyphae of the flesh 4-12µ wide, up to 20-30µ wide near the centre of the flesh,
the cells 30-150µ long, short-celled, secondarily septate, slightly constricted at the
septa, without clamp-connexions, longitudinal, compact.
'
.
TExT-FIG. 75. Clavaria acuta: above, ellipsoid

spores: middle, subglobose spores: below,as figured
from U.S.A. (from Coker): x 1,000.
This species belongs in the affinity of C. Gibbsiae and C. tenuipes, and differs
fundamentally from C. vermicularis in the loop-like clamp at the base of the basidium
and in the broad multiguttulate spore: it shows how superficial are the macroscopic
resemblances of Clavarias. Donk has made C. acuta the lecto-type of Holocoryne
Bon. (Clavaria sect. Holocoryne Fr.), and thus it confers the subgeneric name Holo
coryne on this group of Clavaria. C. acuta may grade into C. Gibbsiae, but I could
find no evidence of secondary septation in the hyphae of C. Gibbsiae, though it has
the primary septa almost as close as the ultimate secondary septa of C. acuta.
The spores vary much in size and shape, even on the same fruit-body. Measure
ments of 43 spores from one fruit-body are given in Table IX: their range is 6·5 x 5µ
to 10 X 8µ, or 10·3 X 6µ (E 1·1 to 1·8). In this collection and in another of mine from
Hertfordshire (Tring) in England, the spores were broadly ellipsoid rather than sub
globose. Such collections have been referred, I think, to other species (even of
CLAVARIA 223
Clavulina) in the belief that C. acuta has only globose spores: thus C. fragilis sensu
Rea (sp.. 8-9 X 5-6 µ,) and C. falcata sensu Bourd. et Galz. (sp. 7-10 X 5-9 µ,). Com
parison of the total spore-volume with basidium-volume suggests that most basidia
of C. acuta are 2-spored.
The fruit-body is acute when young, but generally becomes blunt as the hymenium
develops over the apex. Some fruit-bodies are slender, and resemble C. tenuipes,
others so thick as to suggest Clavulina rugosa, which is never so brittle.
TABLE IX. CLAVARIA ACUTA (ONE FRUIT-BODY)
Spore-size in µ. Length ---:- width ( = E) Number of records

6·5 X5 1·3
7x6 1·2 3
7x6·5 1 ·1 2
7·5X5·5 1·4
7·5x6 1·3
7'5X6·5 1·2 3
8x5·5 1"5
8x6·5 1·2 7
8·5x6 1·4
8·5x6·5 1 ·3 3
8·5X6·7 1·3
8·5X7 1·2
8·5X7·5 1 ·1
9X6·5 1·4
9x7 1·3 5
9X8 I 'I
9·5X6·5 1·5 I
9·5x8 1·2 2
10X5·5 1·8
IOX6 1·7
10X6•5 1·5 I
10x7 1·4 2
1ox8 1·3
10·3x6 1·7
6·5-10·3X5-8 1·1-1·8 43
7-7·5 µ. long 1·18 Average of 11 spores
8-8·5 1·26 15
9-9·5 " 1·28 10
10-10·3 1·52 7
8·5X6·6 1•3 43
(Collection from Chippenham, Suffolk, England_,,)
Note: apiculus omitted from measurements.
I have examined a fragment,of the type of C. filipes in the Kew herbarium and find
that it has the typical hyphae of Clavaria, short-celled without clamps, and spores
7·5-10X 5·5-7·5 µ,. It is clearly a slender form of C. acuta and very different from
Coker's interpretation (see Clavulinopsis filipes).
In reducing C. falcata as a synonym, I have followed Dank. This species, as
interpreted by Quelet, Patouillard, Bourdot and Galzin, is certainly C. acuta. 1 How
ever, C. falcata var. citronipes Quel. has a yellowish stem and may be-different, thus:
2-3 cm. high: stem yellowish, pellucid: head white: solitary among turf (France):
spores 7 µ,, globose: basidia 2-spored.
1
Recently Coker has distinguished C. falcata in U .S.A. by its prevailing falcate shape, far
less delicate texture, and flesh with duplex structure ('stuffed within a firmer rind'). Its white
spores are said to turn yellowish on drying. (Journ. El. Mitch. Sci. Soc. 63, 1947, 52, pls. 6, 14).
224 CLAVARIA
C. falcata var. tenuissima Weirun. (Karst., Finl. Nat. o. Folk 37, 1882, 176) is de
scribed as: head 2-5 mm. long, stem slender, passing gradually into the club: Russia.
Compare also C. Crosslandii.
The species, studied cytologically by Juel as C.falcata, had 6-7-8 sterigmata, long
ellipsoid spores, and stichobasidial nuclear division (Nova Act. Reg. Soc. Sci. Ups.
Ser. 4, 4, 1916, No. 6). It was certainly misidentified, but I cannot say to what it
should be referred. C.falcata sensu Juel is the basis of Stichoclavaria Ulbrich which,
taxonomically, is a synonym of Clavaria. Compare Clavulinopsis hastula and Lentaria
coronilla.
C. affinis Pat. et Doass.
Tab. An. 1886, 205, No. 470.
1-4 cm. high, simple, solitary or gregarious, whitish or cream, drying pale yellow,
obtuse, often compressed, truncate or retuse at the apex: stem 5-10 mm. slender,
opaque, drying brownish.
On earth, in hedges and woods: France (uncommon).
Spores 5-8 X 3-4,_.., white, smooth (punctulate-Pat.), ovoid or oblong, with
granular or 1-guttate contents.
Basidia 21-40 X 5-7,_...
Hyphae 4-9(-30) fA,, thin-walled.
No difference from a subsolitary variety of C. vermicularis appears from the descrip
tion, but the hyphae are unknown in detail.
C. alliacea sp. nov.

Text-Fig. 76.
-7 cm. X 2·5-3·5 mm., simple, scattered or caespi
tose in small tufts, cylindric, obtuse or subacute,
white: stem 10-15 X 1-2 mm., distinct; waxy, brittle,
solid: smell and taste strong, of garlic.
On the ground in the forest: Malaya (Singapore,
Bukit Timah, 10.11.29).
Spores 9-11 X 8·5-9· 5 fk, white, smooth, broadly
ellipsoid or subglobose granular-guttulate.
Basidia 50-55 X 8-9 fA,: sterigmata 1-2, 10-20 fA,
long.
Cystidia none.
Hyphae 5-20 µ wide, thin-walled, without clamps,
compact, longitudinal, with a few narrow inter
weaving hyphae.
This is my original description of the fresh
specimens. At the time I did not understand the
hyphal differences in Clavarioid fungi and I have,
unfortunately, mislaid the specimens, so that I have
been unable to check them. The brittle flesh, wide
hyphae, and absence of clamps indicate Clavaria
rather than Clavulina or Clavulinopsis and the
multiguttulate spores suggest the subgenus Holoco
ryne. The fungus, indeed, seems distinguishable
TExT-FIG. 76. Clavaria alliacea, from C. Gibbsiae only by the smell and taste, the
X 1, ooo. slightly wider spores, and the 1-2, long sterigmata.
Such sterigmata appear abnormal, but they were present in large numbers on all the
fruit-bodies which I examined only a few hours after collection.
The description of Clavaria fuscata is remarkably like this. Both have a garlic
smell, 2-spored basidia, and ellipsoid spores, but C.juscata has narrower spores, much
CLAVARIA 225
shorter basidia (22-30µ,), short sterigmata, and, apparently, 1-guttate spores, in which
it resembles Clavulina ingrata.
C. argillacea Fr.
Syst. Myc. 1, 1821, 482: Boud., le., t. 175 (as C. ericetorum Pers.): Konr. and
Maubl., le. Se!. Fung., t. 494: Favre, Bull. Soc. Myc. Fr. 55, 1939, 217, f. 10.
Synonyms: C. fiavipes Fr., Syst. Myc. 1, 1821, 483 (teste auctt.).
C. ericetorum Pers., Myc. Eur. 1, 1822, 100 (teste auctt.).
C. argillosa Britzm., Hym. Siidb. 8, 1891, f. 32: Rev. Hym. 1909, 223.
C. pallescens Pk., Bull. N.Y. St. Mus., 131, 1909, 34 (teste Coker).
C. lanceolata Imai, Tr. Sapporo Nat. Hist. Soc. II, 1930, 71: ibid. 12, 1931, 18
(? = var. brevispora).
C. meakanensis Imai, ibid. II, 1930, 71: ibid. 12, 1931, 18 (? = var. obtusata or var.
sphagnicola).
? C. subfalcata Atk:, Ann. Myc. 6, 1908, 58: Coker, Clav. U.S. Can. 1923, 21, t. 2,
f. 81, 91: (non C. subfalcata Karst. 1882 = Pistillaria typhu(oides): Hemmi and
Kurata, Act. Phytotax. Geobot. 2, 1933, II2.
Plate 2.
3-8 cm. high, simple, solitary or gregarious or in small tufts of 2-10, whitish yellow,
alutaceous, cream, yellowish clay, pale citron-yellow, or greenish yellow: stem -1 ·5 cm.
long, distinct, more deeply coloured than the fertile part, clear yellow to sulphur-yellow:
rather brittle: smell none, taste like tallow, or ill-defined, or none.
Fruit-bodies 2-8 mm. wide, cylindric and subacute, becoming clavate and obtuse;
often compressed and with 1-2 longitudinal furrows, sometimes truncate, rarely
branched, gradually attenuate into the stem: flesh pallid white or yellowish, often
becoming hollow.
Terrestrial, on moors, heaths, peat-bogs, or among grass on hillsides: Europe
(rather common in western Europe, from Spain to Sweden): Japan: U.S.A. (Mass.
rare): (Austral.?, teste Cooke).
Spores 9-12 x 4·5-6 µ, (8-14 x 4-7 µ,), white, smooth, ellipsoid or subcylindric, often
slightly curved (flattened adaxially), thin-walled, multiguttulate, occasionally with
several guttulae (Donk), apiculus o· 5 µ, long.
Basidia 55-74 X 6-8·5 µ,, multiguttulate, with a wide loop-like clamp at the base:
sterigmata 2-4, mostly 4, 7-12 µ, long.
Hymenium not thickening, or very slightly, fertile over the truncate head: cystidia
none.
Hyphae 3-25 µ, wide, thin-walled, secondarily septate without clamps, the cells
20--120µ long (-300µ in the scarcely inflated medullary tissue).
Yellow colour of the flesh and stem diffused in the sap of the longitudinal hyphae.
forma citrina Bourd. et Galz.

Hym. Fr. 1927, 116.
Synonym: C. citrina Quel., Bull. Soc. Bot. Fr. 1876, 330 (non sensu Hemmi et
Kurata, see Clavulinopsis luteo-alba).
Fruit-bodies small, -2 cm. high, cream-citron, slightly greenish towards the base,
subclavate, obtuse, very fragile; mycelium pulverulent, white.
Solitary on the ground in heaths: France (Vosges).
Spores 10 /l. long, ellipsoid (Quel.).
var. brevispora var. nov.

Bourd. et Galz., Hym. Fr. 1927, II6 ut forma sine nomine.
Spores 6-9 x 4-5 µ,, shorter than typical.
France.
5119 Q
CLAVARIA
var. dispar Pers.
Myc. Eur. 1, 1822, 180: Bourd. et Galz., Hym. Fr. 1928, II6.
Fruit-bodies 4-6 cm. high, clear sulphur yellow, then cream-alutaceous or cream flesh
colour, acute or obtuse.
Lawns and pastures: France (uncommon).
Spores 6-10X4·5-6 µ,.
var. gracillima var. nov.

Basinym: Clavaria gracillima Pk., Rept. N.Y. St. Mus. 28, 1876, 53, t. 1, f. 9: Burt,
N. Am. Clav. 54, t. 9, f. 87: Coker, Clav. U.S. Can. 1923, 23, t. 91: (non C.
gracillima Wakker 1898, see Pterula Oryzae).
2·5 cm. high, simple, very slender, rather tough, acute or acuminate, pale yellow, a
little thicker than the long, slender, bright yellow, shining, and distinct stem.
Among moss in a pasture: U.S.A. (N.Y.).
Spores 7·5-10X3·5-4 µ,, white, smooth, pip-shaped (Coker, for Peck's type):
7-8X3·5-4·5 µ,, rugulose in aqueous mount, smooth in glycerine mount (Burt).
Basidia 30X6 µ,: sterigmata 4.
var. obtusata Favre.

Bull. Soc. Myc. Fr. 55, 1939, 216, f. 11.
Synonym: C. obtusata Boud., Bull. Soc. Myc. Fr. 33, 1917, 12, t. 4, f. 2.
Spores 8-10·5 x 5-6·5 µ, (9-10X5-6 µ,, Bourd. et Galz.), slightly shorter and broader
than typical.
Fruit-bodies 1•5-7 cm.X2-4·5 mm., slender, obtuse, pale ochraceous grey: stem
indistinct, pale sulphur yellow: fragile, stuffed.
On peaty ground: France (Jura, uncommon).
Basidia 64-75X7 µ,, with 4 sterigmata 13 µ, long (Favre): 50-60X7-10 µ, (Bourd. et
Galz.).
Hyphae -15 µ, wide.
var. sphagnicola var. nov.

Synonym: C. sphagnicola Boud., Bull. Soc. Myc. Fr. 33, 1917, 12, t. 4, f. 3: Favre,
Bull. Soc. Myc. Fr. 55, l939, 215, f. 9.
Growing among Sphagnum: spores slightly broader, 9-II·5X5·5-6·8 µ, (Favre)
(7-10X5-6 µ,, Boud.), ovoid or ellipsoid, rarely with 1 large gutta.
Fruit-bodies 3·5-11 cm.X2-3·5 mm., or flattened up to 7 mm., simple, scattered,
or 2-3 together, subcylindric, subacute then blunt, often compressed and longitudinally
furrowed, pale yellow or pale argillaceous yellow: stem indistinct, clear pale citron yellow,
with white mycelium at the base: flesh white, fragile, insipid.
Among Sphagnum: France (frequent).
Basidia 60-75X7�8 µ, with 4 sterigmata II µ, long (Favre): 50-55 X6-7 µ, (Bourd. et
Galz.).
KEY TO THE VARIETIES

Sp. 9-13 µ, long
Sp. 4·5-6 µ, wide, elongate-ellipsoid or subcylindric
-8 cm. high: pale yellowish, stem brighter yellow C. argillacea
typica
-2 cm. high: pale citron: very fragile f. citrina
Sp. 5·5-7 µ, wide, 9-II ·5 µ, long, ovoid-ellipsoid: -II cm. high,
often flattened: among Sphagnum var. sphagnicola
CLAVARIA 227
Sp. 6-10µ. long
Wholly clear sulphur yellow at first
Sp. 4·5-6 µ. wide: Europe var. dispar
Sp. 3·5-4·5µ. wide: U.S.A. var. gracillima
Coloured as in the typical state
Sp. 8-10·5x5-6·5µ. var. obtusata
Sp. 6-9 X 4-5µ. var. brevispora
This seems to be a fairly common, but very variable, species of the north temperate
parts of the world. It requires much more study before it can be circumscribed
satisfactorily, but it appears to be most nearly related with the tropical C. Gibbsiae.
The predominance of the yellow colour in the stem and flesh distinguishes it from the
yellow species of Clavulinopsis, which mostly have the hymenium and subhymenium
more deeply coloured than the flesh.
Favre has maintained C. sphagnicola as a species with larger, paler fruit-body than
usual, broader spores, and sphagnicolous habit. The first two pqints are generally to
be observed in sphagnicolous forms of fungi and indicate the abundant water for
development rather than specific peculiarities, e.g. Psilocybe uda, Tubaria spp.,
Ombrophila spp. C. argillosa appears to be identical with spores 10-12 X 6-8 µ..
The descriptions of the Japanese species are:
C. lanceolata: as C. argillacea, but with smaller, lanceolate fruit-bodies, smaller
spores less than 10µ. long: clubs lanceolate, cream colour: stem lemon yellow: among
grass in woods: Japan.
C. meakanensis: as C. lanceolata but with compressed, clavate, or subcylindric clubs
and larger spores: cream-white with lemon-yellow stem: on wet humus in the moun
tains: Japan.
Regarding C. subfalcata Atk., the only difference that I can find is the presence of a
gutta in the spore, according to Atkinson, and it is a detail that needs corroboration.
Favre maintains that it is specifically distinct from C. argillacea without clear explana
tion of his reasons: if so, it will need a new specific epithet, as the binomial is a later
homonym. The following is Coker's description:
1-5·7 cm. X 0·7-2·5 mm., simple, solitary or gregarious, slender, cylindric, acute or
blunt, often curved, smooth, white or pale creamy white becoming pale yellowish (occa
sionally faintly vinaceous at the apex): stem 0·8-2·8 cm. X 1-1·5 mm., distinct, pale
yellow to lemon yellow, sometimes white when young: rather brittle: no smell or taste.
On the ground: U.S.A. (N. Car., N.Y.).
Spores 7-IIx4·5-7µ., white, smooth, oblong-ellipsoid, 1-guttate.
Basidia 7µ. wide, 4-spored.
Wehmeyer's collection from Nova Scotia, doubtfully referred to C. subfalcata, may
be Clavulinopsis appalachiensis: the peculiarities of the collection were: clubs sub
fusiform, smooth, primrose yellow, pale yellow, or white, stems deeper yellow,
basidia 2-spored, spores 6-7 X 5-6 µ., subglobose-pip-shaped, terrestrial (Pap. Mich.
Ac. Sci. Arts Lett. 20, 1934, 252). Hemmi and Kurata's collection of C. subfalcata
from Japan had spores 5·7-10x 5-7µ. as in C. argillacea var. obtusata. The nature
of the hyphae will settle any doubts of the genus of these collections.
Regarding C. gracillima Pk., I have followed Coker's suggestion that it is a narrow
spored variety of C. subfalcata. Burt indicates, however, that it may have rugulose
spores and Wehmeyer refers to it a collection from Nova Scotia which has rough
spores, as follows:
1·5-4 cm. X 1-2 mm., simple, scattered, rarely with a short branch, pale orange
yellow to orange buff or deep chrome, acute: stem 5-14xo·5-1 mm., distinct, rather
fibrous, paler, shining yellow: flesh yellow, fleshy-tough, elastic: hymenium paler than
the flesh, in section: on decayed or buried wood among moss, Canada (Nov. Scot.):
CLAVARIA
spores 7-8·5 X 3·5 µ,, white, minutely asperulate when mature, ellipsoid, apiculate:
basidia 7-9 µ, wide, 4-spored. (Pap. Mich. Ac. Sci. Arts Lett. 20, 1934, 253, t. 5 r. f. r.)
Possibly this is a rough-spored Clavulinopsis, as indicated by the deeper coloration
of the club.
TABLE X. SPORE-SIZE IN CLAVARIA ARGILLACEA
Author Spore-size in I-' Country Remarks

Favre 9-12·5X4·5-5·5 France typical C. argillacea
Konrad et Maublanc 8-11X4-6 " C. argillacea
Donk 9-13·5X4-6 Holland
Corner 10-12·5x4·5-5 England ,, (in herb. Kew)
8-12X4·5-6 (Brandon, Suffolk)
Cotton et Wakefield 10-1 l X5-6 C. argillacea
Rea 10X5-6 " "
Coker 9-12X5-7 Germany Sydow Myc. Germ. 453, as
C. argillacea
France Collection ofBoudler's in Bresadola's
herbarium, as C. argillacea
Bourdot et Galzin 10-12(-14)X5-'7 C. argillacea
6-10X4·5-6 forma dispar
6-9X4-5 forma brevispora
9-10X 5-6 var. obtusata
Favre 8-10·5 X 5-6·5 " "
Coker 8-10X4-5·5 C. obtusata (Savigne, Nov. 1913)
Boudier 7-1ox5-6 C. argillacea var. sphagnicola
Favre 9-11·5X5·5-6·8 " " ,,
Coker 9·3-12·2X4·2-5·4 U.S.A. type of C. pallescens
Peck 9-12·7X6·3-7 " ,, "
Britzelmayer 10-12X6-8 Germany type of C. argillosa
Quelet 1o I-', ellipsoid France C. argillacea forma citrina
Atkinson 7-1ox5-7 U.S.A. C. subfalcata-Atk. (type), 1-guttate
Coker 8-10·5X4·5-6 ,, (Coker 5630,
10689) 1-guttate
7-9X5-6 C. subfalcata (Coker 14108)
8-9·3X5-5·5 (Coker 13460)
7·8-11X4·5-6·5 (Coker 5573)
10·5X5 (Harper s.n.)
C. atrobadia nom. nov.

Synonym: C. nigrita Bres., Fungi Trident. 1881, 62, t. 67, f. 4: le. Myc., t. 1105:
Bourd. et Galz., Hym. Fr. 1928: (haud C. nigrita Fr., Syst. Myc. 1, 1821, 483:
Epicr. 1, 1836, 578: Hym. Eur:.-1874, 676 = Geoglossum nigritum (Fr.) Cke., Myc.,
t. 96, f. 345).
6-10 cm. high, simple, caespitose in small tufts of five to nine, reddish brown or
rufous bay then blackish, fragile: smell faint, of meal (Bres.): 4-5 mm. wide, cylindric
and acute, becoming compressed and longitudinally sulcate, straight or curved, hollow,
drying black: flesh bistre.
Among grass: Europe (France, N. Italy: rare).
Spores 5 x 2-2·5 µ,, white, smooth, ovoid, aguttate.
Basidia 20-25 X 5-6 µ,.
This species is closely allied with C. fumosa, from which it is said to differ in the
darker richer colour, smaller tufts, smaller spores, and faint smell. Though I have
not examined it myself, I refer it without hesitation to Clavaria. But C. nigrita sensu
Coker seems to be a different species with elastic flesh and narrower hyphae, and I
CLAVARIA 229
have separated it under the new name C. atroumbrina. I think it necessary to name
both fungi anew in order to clear up the confusion over C. nigrita.
On the evidence of specimens of C. nigrita Fr., in Berkeley's herbarium, which had
been named and sent to Berkeley by Fries, Cooke decided that C. nigrita was a Geo
glossum, for such the specimens were, and he transferred the species as G. nigritum
Pers. It seems that Fries never discovered the error and, apart from Bresadola's
mention of it, it seems to have escaped other mycologists. If C. nigrita of Persoon was
a true Clavaria, it was misinterpreted by Fries, but his misinterpretation (of 1821) must
stand according to the International Rules of Nomenclature. Hence, C. nigrita of
Bresadola and Coker is a later homonym.
C. atroumbrina nom. nov.
Synonym: C. nigrita sensu Coker, Clav. U.S. Can. 1923, 43, t. 1, 82 (baud
C. nigr.ita Fr. = Geoglossum nigritum (Fr.) Cke.: baud C. nigrita Bres. = C. atrobadia
Corner).
2-7 cm. high, simple, solitary or subcaespitose in small groups of two to seven, not
connate, umber then dusky sepia or blackish brown, smooth then wholly,,closely and finely,
longitudinally rugulose, soft and elastic, smell none, taste none: 1-2·5 mm. wide,
cylindric and subacute, becoming somewhat flattened, the apex sometimes knobbed
or dilated and occasionally with very short antler-like branches, sometimes contorted:
stem short, indistinct, white tomentose in the lower half.
Among grass and moss: U.S.A. (N. Car., N.Y., rare).
Spores 4·5-6·5 X 2·2-3·2µ., white, smooth, ovate-elliptic, aguttate.
Basidia 9µ. wide, with 4 sterigmata 7 µ. long.
Hyphae 6-9·5µ. wide.
Coker referred this species with doubt to C. nigrita Bres. ( = C. atrobadia), from
which it differs in the duller colour, the longitudinally rugulose mature hymenium,
the elastic flesh, and the absence of smell. The elastic flesh does not agree with
Clavaria but suggests affinity with Clavulinopsis. There has been confusion over
C. nigrita, as I have explained under C. atrobadia, and in giving this fungus a new
name I hope to make clear the path of inquiry. Has C..atroumbrina the hyphal struc
ture of Clavaria or Clavulinopsis?
The spore-measurements given by Coker for C. atroumbrina are: 5·5-6·3 X 2·7-3·2µ.
(N. Car.), 4·5-6·6 X 2·2-3µ. (N.Y.), 5-6 X 3µ. (N.Y., Atk.).
C. avellaneo-nigrescens Imai.
Tr. Sapporo Nat. Hist. Soc. 12, 1931, 20.
1·5-5 x 0·2-0·4 cm., solitary or gregarious, simple, elongate-clavate, obtuse, avella
neous when young, becoming blackish when adult, drying blackish: stem indistinct,
white tomentose at the base.
On the ground among moss in woods: Japan (Prov. Ishikari: Nopporo).
Spores 4-6µ., whiie, smooth, globose, apiculate.
This is said to be allied with C. Greletii. It may be Clavulinopsis.
C. Barlae Bres.
Sacc., Fl. It. Crypt. Hymen. 1916, 1230: Syll. 23, 1925, 490 (ut C. Bariae).
Simple or sparingly branched and 1-1·5 cm. wide, solitary or caespitose, rose white
or subcinnabar: stem 10-15 x I mm., slender, glabrous, dilated at the apex or .shortly
branched: branches difformed, obtuse.
On the ground: France (Nice).
Spores 5-6 x 4-5µ., white, smooth, subglobose, apiculate, 1-guttate.
Basidia 32-36 X 7-8µ..
This seems the branching form of C. rosea, but until its hyphal structure is known
it cannot be classified with certainty.
230 CLAVARIA
C. cinereo-atra Rick.
Brot. 5, 1906, 12.
2-3 cm. X 2-3 mm., simple, solitary, rugose, cmereous black, white pruinose: stem
white then straw-colour, glabrous
On the ground: Brazil.
Spores 15µ, wide, white, subglobose: (8·5-10 X 7"5-8·8 µ,: Singer, Myc. 37, 1945,
425).
Perhaps allied with C. Greletii, but with very large spores. See also C. nigricans.
? Clavulina (cf. C. fioridana).
C. citriceps Atk.
Ann. Myc. 6, 1908, 56: Coker, Clav. U.S. Can. 1923, 49, t. 1, 82.
1·5-4 cm. X 1·5-3 mm., solitary or in groups of two to three, simple, milk white
becoming pale yellow towards the tip and thence yellowing downward over the greater
part of the club, tip acute, soon withering: stem 1 cm. long, indistinct: solid or partly
hollow: smell and taste none.
On the ground in woods: U.S.A. (N.Y., Mich., Conn.).
Spores 4-4·5 X 3-3·5µ,, white, smooth, subglobose, 1-guttat� (Atkinson), aguttate
(Coker).
Basidia 4-spored.
Coker says this is very near C. vermicularis and perhaps only a variety. It may be
a Clavulinopsis, however, allied with C. amoena or C. spiralis. Compare also Clavaria
falcata sensu Pat. The yellow discoloration seems to be basipetal, unlike the acropetal
browning of C. straminea.
C. Corbierei Bourd. et Galz.
Hym. Fr. 1928, u2, f. 46.
7-12 X 1 mm., simple, solitary or in small groups, white, clavate, rather truncate or
umbilicate at the apex, attenuate to the barely distinct stem 2 X 0·3--0·5 mm.
On bare earth in a rotten stump: France (rare).
Spores 3-4·5(-5) X 3-4µ,, white, smooth, subglobose or broadly ellipsoid, 1-guttate.
Basidia 15-18 X 3-4·5µ,.
Hyphae 2·5-6 µ, wide, thin-walled.
This may belong to the subgenus Holocoryne, though the small spores suggest
alliance with a variety of C. vermicularis and the narrow hyphae suggest Clavulinopsis.
Compare C. Guilleminii.
C. cretacea Coker.
Joum. El. Mitch. Sci. Soc. 63, 1947, 44, pl. 1 (lower figure), pl. 12 (ff. 7-10).
-8 cm. high, solitary, simple, or sparingly branched once or twice, with somewhat'
dilate<;!. axils 4·5-7 mm. wide, branches short, blunt and antler-like, smooth, chalk
white tinged stramineous: stem indistinct: flesh chalk white, fibrous-brittle, easily
splitting, rather dry, becoming hollow: smell and taste none.
On the ground in mixed woods: U.S.A. (N. Car., Coker 13045, Bear Pen Moun-
tains).
Spores 5·5-7·5 X 3-3·7(-4)µ,, white, smooth, subcylindric-ellipsoid.
Basidia 5-6 µ, wide, 4-spored.
Hymenium 33-40µ, thick, 'with numerous inflated, hyaline, often irregular cells, or
cystidia, up to 14µ, thick, completely imbedded or extending to the surface'.
Hyphae 3-7µ, wide: 'no clamp-connexions seen'.
This may be Clavulinopsis filipes.
C. Crosslandii Cotton.
Naturalist 1912, 86: Moller, Fung. Faeriies 1945, 109, f. 40B.
2-3 X o· 1-0·3 cm., simple, solitary or in small groups, cylindric, acute, brittle, stem
CLAVARIA 231
hardly distinct, greyish white or grey, darker with age: flesh darker than the hymenium:
smell and taste pleasant.
In short grass: Gt. Britain (Yorkshire, rare), Faeroes.
Spores 4-5 X 2·5-3µ., white, smooth, pip-shaped.
Basidia 20-25 X 4-5µ.: 4 sterigmata.
Hyphae 5-8 µ. wide, closely compacted.
Apparently near C. Jumosa or C. affinis, but compare C. striata.
A co-type collection (Kew, No. 479, Yorks., 1911) is typical C. acuta with spores
8-10 x 7-8 µ. and short-celled hyphae without clamps. Moller records a single
specimen from the Faeroes as pale umber-grey to pale olive-brown with white base,
fusiform with hardly distinct stem and subglobose spores 4 x 3 · 5µ.
C. Daulnoyae Quel.
Ass. Fr. 1895, 470, t, 3, f. 36: Bourd. et Galz., Hym. Fr. 1928.
(Dedicated to Mme V. Daulnoy, a friend of Quelet's.)
2-4 cm. X 2-7 mm., simple, caespitose, clavate, obtuse or thickened and rounded at
the apex, becoming hollow, clear grey becoming greyish bistre: s�em -10 mm., distinct,
slender, hyaline white, villous at the base, the white mycelium effused and cottony.
On lawns: France (rare).
Spores 7-9 X 5-6µ, white, ellipsoid, smooth (8-9µ, ellipsoid, Quel.).
Basidia 30-45 X 6-9µ.: sterigmata 2-4.
Hyphae 3-12µ. wide.
Quelet says this has the form of Clavaria argillacea and is allied with C. falcata.
Bourdot and Galzin say that it is very close to C. tenuipes and, perhaps, the same: the
only difference seems to be the caespitose habit of C. Daulnoyae. Clavaria argillacea
var. dispar has identical spores.
C. exilis Pers.
Myc. Eur. 1, 1822, 186: Quel., Fl. Myc. 1888, 460: Beeli, Bull. Soc. Roy. Bot. Belg.
42, 1930, fasc. 2, 127.
1-1·5 cm. high, simple, cylindric, filiform, cream white: stem very slender.
On the ground in coniferous forests and on tan bark: France, Belgium, rare.
Spores 11-15 µ. long, white, elongate-ellipsoid, punctate (Quel.).
Little known and, perhaps, near C. tenuipes: placed under Clavaria acuta by Fries
(Hym. Eur. 679). Beeli merely records the species for Belgium.
<::. filiola spec. nov.
(Like a small C. acuta.)
Text-Figs. 77, 78.
7-i 5 mm. high, very small, white, simple, cylindric and subacute to subclavate:
stem 3-7 x 0·3-0·5 mm., subtranslucent, finely puberulous downwards, the base abrupt,
subfibrillose, distinct from the fertile head: soft, brittle, without smell.
On bare soil in the forest, solitary or in small troops: Malaya (Singapore, Botanic
Garden's Jungle 11.11.34).
Spores 4 ·5-5 ;< 4 µ,, white, smooth, subglobose, with vacuolate-oleaginous or cloudy
contents, not guttate.
Basidia 20-34 x 6-7µ,, without clamps: sterigmata (2-)4, 5µ, long.
Cystidia none.
Hymenium not thickening, abruptly delimited at the apex of the stem or with a few
clusters of sterile basidia -r 2 X r ·5-4µ,, fertile over the top of the club: subhymenium
c. 20µ, thick, composed of closely interwoven, uninflated hyphae r · 5-3µ, wide.
Caulocystidia -15ox 1·5-2·5µ,, filiform, unbranched, aseptate, thin-walled, scat
tered, more numerous near the base of the stem, absent from the upper part.
Hyphae 4-8 µ, wide in the flesh, the cells 30-350µ, long, thin-walled, slightly inflated,
232 CLAVARIA
without clamps, narrowed to the primary septa, with 0-3 secondary septa, not con
stricted at the secondary septa, longitudinal, very compact, 1 · 5-3 µ. wide on the surface
of the stem: a few ·hyphae, 2-4 µ. wide, with resinous oleaginous contents like latici
ferous hyphae, simple or occasionally branched.
;: : :.: :: ;
__
;·· ·····,.
':. l
:_:� � .
TEXT-FIG. 77. Clavaria
jiliola, X 1 o.
TEXT-FIG. 78. Clavariajiliola: sterile hymenium at the stem-apex,
hyphae, basidia, and spores, X r ,ooo: caulocystidia, X 500.
This small species suggests a white Typhula or Pistillaria, but its structure is
essentially that of Clavaria. It comes very near C. vermicularis var. sphaerospora and
C. subacuta, both of which appear to have much larger fruit-bodies, and C. subacuta
has a longer stem. C. filiola is the smallest and most-reduced Clavaria. Its hyphae
are rather scarcely secondarily septate, probably through degeneration rather than any
primitive faculty. The deficient septation probably causes the small size of the fruit
body as much as restricted apical growth.
CLAVARIA 233
C. fossicola sp. nov.
Text-Figs. 79A, 79B: Plate 15.
5-12 X 1-1·5 mm., white, dingy cream white or drab with age, very small, simple,
subulate then fusiform-obtuse, acerose with acute apex at first, then obtuse and even
subclavate, often with 2-3 short branches near the apex, occasionally with 2-5 equally
developed branches themselves simple or branched once: stem 1-2·5 X 0·5-0·8 mm.,
scarcely distinct, sterile, subpruinose: flesh firm, cartilaginous, seemingly waxy: smell
none.
TEXT-FIG. 79 A. Clavaria fossicola, X 5.
On bare clay soil covered with Protococcoid slime, gregarious in large troops (not
caespitose), especially in ditches and on banks, generally in the shade of the forest:
Malaya, abundant from sea-level to 1,500 m. alt. (at least).
Spores 5-6·5 X 2-2·7 µ,, white, smooth, thin-walled, oblong-ellipsoid, generally
slightly curved, apex blunt or occasionally subacute, base obliquely apiculate, vacuolate,
aguttate or occasionally 1-2-guttulate.
Basidia 15-25 x 5-7 µ,, subclavate, projecting up to 10µ,, without clamps: sterigmata
4, occasionally 2 or 3, 3·5-4·5µ, long.
Cystidia none.
Sterile basidia 15-25 X 3-4µ,, often very abundant, subclavate or often more or less
capitate, the apex forming a subglobose body 5-7 µ, wide, slightly thick-walled, smooth,
vacuolate, attached by a broad or narrow and sterigma-like apex (? not abstricted and
not conidia).
Hymenium at first 25 µ, thick, gradually thickening to 350µ,, often mostly or entirely
sterile, but generally with some included spores, not stratified, eventually covering the
apex of the fruit-body, submucilaginous at the surface: subhymenium composed at
first of narrow, interwoven hyphae 2-3·5µ, wide, becoming almost pseudoparenchyma
tous with short, rather barrel-shaped cells, 8-25 X 4-12µ,, at the base of the old
thickened hymenia.
Hyphae 3-10µ, wide, the cells 25-150µ, long, without clamps, without secondary
septa, or occasionally a single one f9rming in a cell, the walls firm and becoming
slightly thickened (-0·5µ,), scarcely inflated but generally constricted at the septa,
oocasionally swollen and trumpet-like, often irregular, with H-connexions: loose in
the medulla of old fruit-bodies.
234 CLAVARIA
Stem often with minute blunt processes, 2-6 µ, long, from the narrow superficial
hyphae and with clusters of sterile basidia near the apex.
V
Tmcr-FIG. 79 B. Clavaria fossicola: thickened hymenium, hyphae of the flesh, spores, sterile
hyphal tips of the hymenium, and mycelial hyphae with algal cells: X 1,000.
The soil-alga with which this species is associated is probably a Gloeocystis. It

forms a filmy scum, one cell thick, of Protococcoid cells on the yellow clay, so common
CLAVARIA 235
in Malaya. '.fhe mycelial hyphae are 1·5-2µ wide and have toughly mucilaginous
walls. They are intricately and contiguously interwoven with the algal cells, many of
which die off in patches, but I could not discover that they were penetrated by the
hyphae. The fruit-bodies are devoid of algae. The mycelium may be perennial, for
colonies of fruit-bodies have developed in the same, very localized, patches in the
Reservoir Jungle in Singapore, two or three times a year for some 10 years. The
fruit-bodies appear 2 or 3 weeks after the rains have begun, following a spell of dry
weather, and they last for 1 or 2 months. Generally they develop in thousands, most
commonly on new banks or drains cut by roads in the forest, and they, or their mycelia,
appear shade-loving in contrast with the light-loving C. helicoides.
Such a long life is unusual for small Clavarias. The fruit-bodies of C. Jossicola
persist through the greater part of a rainy season in Malaya, during which the
hymenium slowly thickens and transforms them from sharp, almost filiform upgrowths
to blunt, often misshapen, clubs. A cross-section of' an old fruit-body shows the
original core. of the primordial shaft, 200----'700µ wide, surrounded by the firm, dense,
and much-thickened hymenium. Yet, strangely enough, spores are few: spore-prints
are thin, and many fruit-bodies are permanently sterile, though they have normally
thickened hymenia. Possibly they are sensitive to slight variations in the weather.
The sterile basidia have often capitate tips, suggesting conidia, but I saw no evidence
of their abstriction and they appear wholly vacuolate.
The systematic position of the species is problematic. On the one hand, it is allied
with C. helicoides and appears as a degenerate, colourless, small-spored derivative with
loss of septation of the hyphae, cf. C. filiola and C. acuta. Both are exceptional in
Clavaria for their tough, long-lived fruit-bodies and reduced septation, and both
features are probably connected with the thickening and reduced extensibility of the
hyphal wall, itself connected with the habit of the species. On tl\e other hand, C.
jossicola could easily be mistaken for one of the phycophilous species of Clavulinopsis,
particularly C. constans, or for the lignicolous Lentaria mucida.
C. fumosa Fr.
Syst. Myc. 1, 1821, 483: Bres., le. Myc., t. IIOl: v. Ov., le. Fung. Mai. 1923, t. 3.
1·5-14 cm. high, simple, densely caespitose, pale cream, whitish, or greyish alutaceous,
then pale mouse-grey or fuliginous, rarely deeply coloured or fuscous, 1·5-7 mm. wide,
cylindric then subclavate, becoming compressed and more or less fusiform, often twisted,
acute then obtuse, slender or rather stout, whitish towards the sterile base, becoming
hollow, stem indistinct or none: flesh whitish, brittle: smell none: taste marked,
pleasant, or none.
Among grass in fields, rarely in woods, occasionally on bare ground: Europe
(common), N. America (common), Siberia, Java.
Spores 5-8 x 3-4µ, mostly 5·5-6·5 x 3·5, white, smooth, thin-walled, ellipsoid or
rather pip-shaped, blunt, with a distinct apiculus, aguttate or with granular contents
(not guttate): 5-6 x 2·5-3µ, granular or with a large gutta (Moller, Faeroes).
Basidia 30-45 x 6-7 µ, without clamps: sterigmata (2-)4, 4-5µ long.
Cystidia none.
Hymenium thickening -100µ: subhymenial hyphae 2-3µ wide, loosely interwoven,
secondarily septate.
Hyphae exactly as in C. vermicularis, 3-30µ wide, the cells 25-80(-120)µ long,
without clamp-connexions, constricted at the septa, the cells often barrel-like.
This common north-temperate species appears to be-remarkably constant, so that
all mycologists are agreed on its determination. Yet it seems no more than a large
colour-variety of C. vermicularis, with which it agrees in all microscopic details.
Lundell says the spores of C. vermicularis are shorter, but he was considering, evidently,
only a small-spored variety of C. vermicularis (Sv. Svamp., fasc. 11-12, 1938, 19).
The only disagreement in the spore-measurements of ten authors, to whom I have
236 CLAVARIA
been able to refer, is Coker's records of 5·5-8(-10) X3·4-4µ. for Sydow's collection
Myc. Germ. 454 in the Kew herbarium. It suggests that giant spores may be found
in C. Jumosa, as in C. purpurea, unless this collection is C. purpurea. In colour and
form C. fumosa comes between C. vermicularis and C. purpurea, wanting only the
purple tinge and cystidia to be identified with C. purpurea. In Table XI is given the
fluctuation in spore-size in a single fruit-body of C. Jumosa.
Van Overeem's record from Java is the only one from the tropics, yet there seems
no doubt from his description and figure. Petch does not mention it for Ceylon and
I have not seen it in Malaya. Nor has it been found in China or Japan, though the
record from Siberia suggests that it surely occurs.
TABLE XI. CLAVARIA FUMOSA (ONE FRUIT-BODY, ENGLAND)
Spore-size in µ. Length -+- width ( = E) Number of records

5X3 1·7 7
5 X3·5 1·4
s·sx3 1·8 4
s·sx3· s 1·6 8
6x3 2'0
6X3·5 1·7 5
6x3·8 1·6
6X4 I'S I
6·5X3 2'2 2
6·5X3·5 1·9 4
6·5X3·7 1·8 3
6·5X4 1·6 3
6·8x4 1·7
7x3·5 2·0 3
7x3·7 1·9 4
7x4 I ·8 2
50
s -s· s µ. long 1·66 Average of 20 spores
6-6·5 µ. long 1·75 20
6·7-'7 µ. long 1·90 10
6x3·5µ. 1·70 50
Note: apiculus excluded.
C. fuscata Oud.
Arch. Neer!. 2, 1867, 35, t. 1, f. 1: Coker, Clav. U.S. Can. 1923, 23, t. 81: Donk,
Rev. Nederl. Homobas. Aphyll. 2, 1933, 78.
Synonym: Clavaria foetida Atk., Ann. Myc. 6, 1908, 56 (teste Coker).
1·5-6 cm. X1·5-4 mm., simple, solitary or gregarious, opaque white then pale yellow,
or dull creamy yellow, the tip fuscous with age, blunt: stem I mm. thick, as long as or
longer than the club, translucent white, yellowish upward, with a mycelial film at the
base: brittle: smelling strongly of garlic on crushing.
On the ground and on sandy soil: Europe (Holland), U.S.A. (N.Y., N. Car.).
Spores 7-10X4·5-5µ. (Donk), 8-10 X4·5-6µ. (Coker), white, smooth, ovoid-elliptic,
1-guttate, occasionally merely granular guttate.
Basidia 22-3ox6·5-7·5µ.: sterigmata 2, 6·5µ. long.
Hyphae 3·5-12µ. wide.
This is said to be allied with C. subfalcata (see C. argillacea), but to differ in the
smell and the 2-spored basidia. Its natural position will be uncertain until its hyphal
structure is known.
Coker gives the spores as 8-10 X 4·5-6 fL (No. 3459) and 7·5-9·8 X4·8-6·5 fL
CLAVARIA 237
(No. 3485). Atkinson gives the spores of C. foetida as 6-9 X 5-7µ,, and 1-guttate,
which suggests Clavulina.
C. Gibbsiae Ramsb.
Gibbs, Contr. Phytog. Arfak Mtns. 1917, 187.
Text-Figs. 11, 14c, 80-2.
TEXT-FIG. 80. Clavaria Gibbsiae: stages in the development

of a tuft of fruit-bodies, X 1.
-10 cm. high, 2·5-4 mm. wide, or flattened -8 mm. wide, caespitose, rarely solitary
or gregarious, simple or once furcate, subclavate, generally blunt, rarely subacute even
when young, white, yellowish with age: stem II-22 x 1·5-2·5 mm., distinct, glabrous,
drying cottony: flesh waxy-firm, slightly brittle, solid: smell none.
On earth in the forest or in shady places, and on burnt ground: New Guinea
(Manokoeari), Africa, Malaya (Pahang, 7.n.30: Sing. Field, No. 24165).
Spores 8-10·5 X 7-8 µ, (7-11 X 4-6µ,, Ramsbottom), white, smooth, broadly ellipsoid
to subglobose, densely granular guttulate.
Basidia 4-spored, furcate at the base: (40 X 8-10µ, with 2 sterigmata; Ramsbottom).
Hymenium thickening to 130µ,.
Hyphae as in var. megaspora.
var. tenuis var. nov.

1-5·5 cm. X 1-2·5 mm., or flattened to 3·5 mm., solitary or gregarious;not caespitose,
slender, cylindric and subacute, becoming blunt and subclavate or subfusiform, white,
pale yellowish with age: stem 3-II xo·5-1 mm., distinct, glabrous (even when dry),
subhyaline: waxy-fragile, solid: smell and taste none.
On burnt ground in the forest: Malaya (Singapore, Bukit Timah, every fungus
season).
Spores 8·5-10 X 7-8·5µ,, as in var. subglobosa.
238 CLAVARIA
Basidia 40-66 X 9-10 ft, clavate, furcate at the base: sterigmata 4, 6-8 ft long,
Hymenium not thickening: subhymenial cells inflated, 8-20 X 4-10 ft, not pseudo
parenchymatous.
'I'ExT-FIG. 81. Clavaria Gibbsiae: left, the superficial hyphae of the stem: centre, subhymenial
hyphae: right, longitudinal hyphae of the stem: X 500.
Hyphae as in the typical state but with many narrow hyphae 1·5-5 ft wide, inter
weaving and longitudinal in the fertile part of the fruit-body, the longitudinal hyphae
often irregularly inflated with long and short cells in the same hypha.
Stem microscopically glabrous.
forma microspora f. nov.

Text-Fig. 86.
Spores 6-7"5 X 5·5-7 µ., broadly ellipsoid, ovoid or subglobose. 1
1
These measurements were taken from many full-grown but attached spores in alcohol
formalin material. On the stems of the specimens were numerous slightly larger spores,
7-8·5 µ, wide, set with many hyaline very slender spines -1 µ, long. 'I saw no such spores
attached to the basidia and there were many smooth spores on the stems. Hence I conclude
the echinulate. spores must have belonged to another fungus. But compare Clavu/inopsis
asterospora and Araeocoryne.
CLAVARIA 239
On the ground in the forest: Malaya (Pahang, Tembeling, 6.11.30).
As var. tenuis but with slightly narrower hyphae 3-13p. wide (-25 p. near the sub
hymenium) and slightly smaller basidia 40-50 X 7-8 p. (4 sterigmata, 6-7p.).
I
TEXT-FIG. 82. Clavaria Gibbsiae: TExT-FIG. 83. Clavaria Gibbsiae
fruit-bodies separated from a var. tenuis, X I.
tuft, X I.
var. megaspora var. nov.

Plate 2.
Spores 9-13 X 6-8 p., broadly ellipsoid, blunt at both ends, slightly flattened
adaxially, densely granular-guttulate, rarely with a large gutta.
On bare earth under trees: Malaya (Singapore, Bot. Gdns., 21.12.40: Pahang,
Tembeling, 3.11.30).
-8 cm. x 2-3 mm., or flattened -5 mm. wide, simple, densely caespitose, cylin<lric,
generally blunt, rarely subacute even when young, often becoming flattened, sulcate,
sometimes hollow, white, tips withering brown: stem 8-15 X 0·7-1·5 mm., distinct,
appearing glabrous when moist, finely cottony when dry, substrigose at the base: flesh
white, waxy-soft, brittle: smell and taste ill-defined.
Basidia 50-70 x 9-r r p., clavate with a narrow stalk, furcate at the base with very
lax clamp-connexions, densely oleaginous-granular: sterigmata 4, occasionally 2 or 3,
7-9p. long.
Hymenium c. 70 p. thick, not thickening, abruptly delimited at the stem-apex,
eventually covering the whole club: subhymenium c. 30 p. thick, composed of densely
interwoven, narrow hyphae r ·5-4 p. wide, not pseudoparenchymatous: cystidia none.
Hyphae 5-25p. wide, the cells 30-250 p. long, thin-walled, much inflated, without
CLAVARIA
clamps, constricted at the septa or not (not secondarily septate), the cells more or less
fusiform, loose in the central tissue, compact near the subhymenium and shorter
(30-100µ long) and more frequently branched, the cells often forming submoniliform
rows and almost pseudoparenchymatous: some of the longitudinal hyphae not or
little inflated, 2-5µ wide, but without narrow interweaving hyphae: at the base of the
stem the hyphal walls slightly thickened.
Stem with narrow superficial hyphae 1·5-2 · 5 µ wide, very compact but not
agglutinated, giving off scattered, obliquely excrescent hyphae -150 X 1·5-2·5µ, not
modified into caulocystidia.
Hyphal tips at the growing apex 1·5-2·5µ wide, i.,itertwining, the subapical cells
12-20µ long on delimitation: hymenium developing 200-500µ from the apex by the
out-turning and sympodial branching of the hyphal tips, and by the excrescence of
numerous laterals from the more deeply situated hyphae.
KEY TO THE VARIETIES OF CLAVARIA GJBBSIAE

Generally caespitose: -9 cm. X 2-3·5 mm., or flattened to 7 mm.,
rather large and stout.
Sp. 8-10·5 X 7-8µ: Malaya, New Guinea,? Africa . C. Gibbsiae
Sp. 9-13x6-8µ: Malaya var. megaspora
Solitary or gregarious: -5·5 cm. X 1-2·5 mm.: slender.
Sp. 8·5-1ox7-8µ: Malaya var. tenuis
Sp. 6-7·5 X 5·5-7µ, subglobose: Malaya f. microspora
TABLE XII. SPORE-SIZE IN CLAVARIA GIBBSIAE VAR. MEGASPORA
No. of
Spore-size in µ, E' records
----
Spore-size in µ, E' No. of records
7·7X6·o 1·6 l 1o·ox7·5 I ·3 3
8·oX 5·5 1·5 I 10·3X7·5 1·4 I
8·ox6·o l ·3 2 10·5x7·0 I. 5 6
8·ox6·5 1·2 2 10·5X7·5 1·4 4
8·3X6·o 1·4 I 10·5X8·o 1·3 I
8·5X6·o 1·4 7 11·ox6·5 1·7 I
8·5X6·5 I·3 3 11·ox7·0 1·6 2
8·7X6·o 1·5 I 11·0X7·3 1·5 3
9·ox6·3 1·4 I 11·ox8·5 1·3 I
9·ox6·5 1·4 2 n·5X7·0 l ·6 2
9·0X6·7 1·3 l 11·5 x7·5 l ·5 l
9·0x7·0 1·3 2 12·ox7·5 1·6 I
9·5x6·5 1·5 3 12·5X7·5 1·7 l
9·5X6·7 1·4 l
9·5x7·0 1·4 2 7·7-12·5X 5·5-8·5 l ·2-1·7 73
9·5x7·5 l ·3 l 8-8·3 long I·33 Average of 6 spores
9·7X7·3 I ·3 l 8·5-8·7 " 1·40 " II "
10·ox6·5 1·5 3 9-9·5 " 1·40 " 13 "
10·0X6·7 1·5 3 9·7-10 " 1·45 " 18 "
1o·ox7·0 1·4 7 10·0-10·5 " 1·45 " 12 "
1o·ox7·3 1·4 l II " 1·53 " 7 "
11·5-12·5 " 1"62 " 5 "
9·8X6·8 1·44 " 72 "
' E = length -:- width. Data from I fruit-body, Singapore.
This is probably a widespread and common tropical fungus which may well be
mistaken for Clavaria vermicularis, C. acuta, Clavulinopsis spiralis, or Clavulina rugosa.
CLAVARIA
The large fruit-bodies are nearly always blunt, even when young, and so can be
distinguished from those of Clavaria and Clavulinopsis which are acute when young
or in full growth. Var. tenuis greatly resembles the north temperate Clavaria tenuipes,
but it is never dirty white or greyish and the spores are wider: it seems, at first sight,
very different from the caespitose var. megaspora, but the typical state of the species
connects them.
The original description of C. Gibbsiae gives the spores as 7-II X 4-6 /L and the
basidia as 2-spored. On the co-type collection of Miss
Gibbs in the Kew herbarium, I found abundant spores
9-10·5 X 7-8 /L and many, clearly 4-spored, rather
broadly clavate basidia: the hyphae were rather short-
celled, -15 /L wide, without clamps. This collection
is certainly identical with my Pahang collection (S.F.,
No. 24165). Whether the discrepancy is an error of
observation or caused by a mixture of specimens I do
not know.
It is noteworthy that the variation in spore-size in
the species is entirely one of length, as in C. tenuipes.
Cooke's record of Clavaria fragilis from Australia
may belong here: 3-8 cm., fasciculate, very fragile,
obtuse, hollow, wholly white or yellow above: spores
10-12 X 4-5/L: on the ground: Victoria (Handb.
Austral. Fung. 1892, 202).
C. Greletii Boud.
Bull. Soc. Myc. Fr. 33, 1917, 13, t. 4, f. 4: Rea,
Tr. Br. Myc. Soc. 12, 1927, 226: Bourd. et Galz.,
Hym. Fr. 1928.
3-5 cm. x 1-2 mm., simple, solitary or in pairs,
slender, obtuse, elongate, entirely brownish bistre to
dark fuliginous or black: stem white at the base: flesh
fuliginous.
On the ground, sandy lawns and old charcoal heaps:
Europe (France, Gt. Britain: rare).
Spores 7-10 /L wide, white, smooth, subglobose,
r-several-guttulate or multiguttulate 6-8 /L wide,
r-guttate (Rea).
Basidia 16-30 X 5-6 /L, 4-spored: 20-35 X 6-8/L
(Rea).
Hyphae 3-rzµ..
The genus of this species is doubtful. Compare C.
avellaneo-nigrescens and Clavulina tasmanica. There
appears to be no authentic specimen preserved.
TEXT-FIG. 84. Clavaria Gibbsiae
C. Guilleminii Bourd. et Galz. var. tenuis, X r ,ooo.
Hym. Fr. 19z8, r 12, f. 47.
ro-Iz mm. high, simple, rather soft, gypsum white, drying brownish, clavate or
truncate-compressed, rather cuneiform, attenuated to the base, without a distinct
stem.
On bare earth, solitary or gregarious, not caespitose: France (uncommon).
Spores 4 x 2·3-z·8/L, white, smooth, elliptic-subcylindric, aguttate.
Basidia 18-24 X 4/L·
Hyphae 3-1zµ. wide, thin-walled.
This is distinguished by its small spores and small, truncate fruit-body. Its position
5119 R
CLAVARIA
will be uncertain until its hyphal structure is known, but it appears to be allied with
C. acuta and C. vermicularis, as C. filiola. Compare C. Corbierei with subglobose
spores.
TExT-F!G. 85. Clavaria Gibbsiae var. tenuis: left, a superficial hypha of the stem:
centre, sub-hymenial hyphae: right, hyphae of the flesh: X 500.
C. helicoides Pat. et Demange.

Bull. Soc. Myc. Fr. 1910, 44.
Synonym: C. miniata Berk. sensu Petch, Ann. Roy. Bot. Gard. Per. 9, 1925, 329
(non C. miniata Berk. = Clavulinopsis miniata).
Text-Fig. 87: Plate I.
1-3 cm. high, simple, gregarious, or solitary, clear pink to orange-pink or even orange
red (especially when dry), waxy-fleshy, rather brittle, drying tough: 1-2·5 mm. wide,
or flattened to 3·5 mm. wide, cylindric and acute, becoming subclavate, obtuse and
often flattened above, commonly fiexuous, especially on drying, often longitudinally
sulcate, occasionally with a short lobe near the apex: stem 1-5 x 1 mm., indistinct,
concolorous, glabrous (without strigose processes), sterile: flesh paler concolorous:
smell none, taste slowly subacrid.
On bare yellow clay or hard soil, generally where trodden or beaten by rain, in the
CLAVARIA 243
open: Indo-China, Ceylon, Malaya: common m the lowlands and mountains,
-1,500 m. alt.
Spores 9-12 X 4-5·5µ. (8-14 X 3·5-5µ., Petch), yellowish pink in the mass, smooth,
thin-walled, ellipsoid, subclavate or subcylindric, often slightly waisted, the apex blunt
or subtruncate, the cytoplasm somewhat oleaginous and pale pink (the walls colourless),
not guttate.
Basidia 40-50 x 8-10µ., clavate, with pale-pinkish oleaginous cytoplasm, without
clamps: sterigmata 2-3-4, mostly 4, 7-10µ. long: (basidia 24-36 X 4-6µ., with 4 sterig
mata 6µ. long: Petch).
Cystidia none: stem with scattered filiform processes -60 X 2-3µ..
Hymenium at first 45µ. thick, gradually thickening to I 20µ.,
covering the apex of the fruit-body, the old effete basidia
secondarily septate: subhymenium 2'--30µ. thick, composed
of 2-5µ. wide, short-celled hyphae compactly interwoven
and almost pseudoparenchymatous, slightly inflated.
Hyphae of the flesh 3-9µ. wide, the cells 40-180µ. long,
with firm, often undulate walls, not or very slightly thickened,
without clamps, the cells often with 1-2 secondary septa,
generally constricted at the primary septa, compactly longi
tudinal: the hyphae next the subhymenium narrow and with
short cells I 5-40 X 3-5µ.: H-connexions frequent.
This is probably a phycophilous species, but I have not been
able to study the exact relation between its hyphae and the
slippery film of algae and lichens which cover the bare laterite
clay on which it grows. In dry weather, in Malaya, the
surface of such soil on hills which have been more or less
denuded, or on paths and banks in the open, rarely in the
light shade of exploited forest, is covered by a dirty greenish
brown or blackish felt or crust which cracks into small patches
with upturned edges. In wet weather the felt revives into a
slippery scum of 1nicro-organisms among which the hyphae
of C. helicoides grow, particularly round cells of Gloeocapsa. TEXT-FIG. 86. Clavaria
Gibbsiae var. tenuis f.
Thus the fungus is characteristic of 'new clearings' planted
microspora, X I ,ooo.
with pineapple or rubber-trees in their early years. The
fruit-bodies appear 7-10 days after the beginning of the rains which terminate a dry
spell, and they last for several weeks, though not so long as those of C. fossicola. They
can withstand incipient desiccation and, in drying, become c&rf�rted, as do most
Clavarias, and this is probably the condition in which Demange collected them in
Indo-China. Petch describes the fungus from similar plac:es in Ceylon and notes
that it does not grow on loose or gravelly soil. Probably it is widespread in the Eastern
tropics, if not throughout the tropics, cf. C. sibutiana.
In general structure the species agrees with C. vermicularis, but its hyphae are .less
inflated, with much fewer secondary septa, and their walls become slightly thickened
so that the fruit-bodies are not brittle. The undulate hyphal walls, but partly distended,
suggest that the inflation mechanism is defective. Sometimes the basidia are sterile
and produce one or two hyphae from the apex which contribute to the thickening of
the hymenium. The colour is caused by the pale-pinkish, oleaginous cytoplasm of the
young basidia, of the subhymenial hyphae, and, to a lesser degree because of their
vacuolation, of the medullary hyphae: that of the basidium passes into the spores and
colcmrs them. The hyphal tips at the apex of the fruit-body also have pinkish cyto
plasm: hence the monotony of the whole fruit--body.
Demange saw living specimens of Petch's C. miniata B. in Ceylon and concluded
that they differed from C. helicoides of Indo-China merely in being less curled. That
was surely caused only by the state of the weather. Petch's C. miniata is certainly
244 CLAVARIA
identical with the Malayan fungus I have described, but it is not the true C. miniata
which was described from S. Africa 'on sandy ground', and is a Clavulinopsis with
subglobose spores (5·5-7·5µ wide in the type, according to Coker).
TExT-FIG. 87. Clavaria helicoides: spores and basidia, x 1,000: subhymenial

hyphae, X 500.
C. incarnata Weinm.
Hym. et Gastrom. 1836 , 510.
Synonym: C. rosea Cott. et Wakef., et Donk, pro parte microscopico.
Text-Fig. 88: Plate 1.
1-8 cm. high, simple, solitary or gregarious, never caespitose, flesh pink discolouring
yellowish, livid white or greyish from the apex downward, the base white strigose hairy:
1-5 mm. wide, subcylindric becoming flattened, narrowly fusiform and slightly
grooved longitudinally, straight or flexuous, apex acute then more or less obtuse,
often hollow when old, the short, pallid, stem-like base 3-8 mm. long, strigose with
white hairs, distinct: flesh concolorous, brittle: smell and taste none or slight and
inparticular.
In grass or on bare soil in woods: Europe, rather rare.
Spores 6·5-10 X 3·5-6·5µ, mostly 7-9 X 4·5-5·5µ (E. 1·6, varying 1·5-2·1), white,
smooth, thin-walled, ellipsoid, blunt, rarely pip-shaped, finely multiguttulate (or some
spores with one gutta?): apiculus 1-1 ·5µ long.
Basidia 50-70 X 7-9µ, multiguttulate, with a wide loop-like clamp as in C. Gibbsiae
(Text-Fig. u), colourless: sterigmata (2-)4, 5-9µ long.
CLAVARIA 245
Hymenium thickening by sympodial branching to 100µ: subhymenium composed
of 1·5-2·5µ wide interwoven hyphae: cystidia none.
Hyphae 3-15µ wide, the cells 25-180µ long, secondarily septate as in C. vermicu
laris, without clamp-connexions: slightly thick-walled in the stem: the pink colour
in the sap of the longitudinal hyphae.
Stem strigose with slightly thick-walled, narrow hyphae 1·5-3µ wide, flexuous and
somewhat intertwined.
This is a rare or uncommon species
distinguished by the pale colour and the
narrowly ellipsoid spores. My description
is based mainly on four collections which
I made in 1924 and 1946 (in fields, Great TEXT-FIG. 88. Clavaria incarnata, X1,000.
Missenden, Bucks., and Cambridge, Eng-
land). It agrees with that given by Rea, Bourdot, and Galzin, and Cotton and Wake
field, except that Cotton and Wakefield give the spores as 7-10 X6-8 µ, which is
unusually wide. T}:ie Clavaria from Mulgrave Woods, England, referred by Donk to
C. rosea (1933, p. 86) is also the same.
My reasons for distinguishing this species from the smaller-spored C. rosea are
given on page 215.
The variation in spore-size from a single fruit-body is given in Table XIII.
Compare the tropical C. helicoides which differs radically in the pink hymenium
and spores, the absence of clamps from the basidia, and the slight secondary septation.
TABLE XIII. CLAVARIA INCARNATA (ONE FRUIT-BODY, ENGLAND)
Spore-size in µ, Length 7 Width Number of records

6·5X3·5 1·9 l
6·5X4·0 1·6 l
7·0X4·0 1·8 I
7·0X4·5 1·6 5
7·5X3·5 2·1 l
7·5X4·5 I"7 3
7·5X5·0 l "5 2
8·oxs·o 1·6 9
8·5X4·5 1·9 4
8·5X5·0 1·7 9
8·5x5·5 1·5 5
9·ox5·0 1·8 3
9·0X5'5 1·6 2
10·0X5·0 2·0 l
10·0X5·5 l ·8 l
10·0X5·7 1·8 l
10·0X6·o 1·7 2
1o·ox6·5 1·5 3
-
6·5-10X3·5-6·5 1·5-2·1 54
6·5�·5 µ, long l ·61 Average of 14 spores
8-8·5 µ, long 1·65 " 27 "
9-10µ, long 1·70 " 13 "
8·3X5·0 1·65 " 54 "
C. isabellina Bres.
Ann. Myc. 9, 1911, 272.
1·5-2X 0·2 cm., simple, clavate or fusoid, glabrous, rugulose, subcompressed,
becoming hollow, bright isabelline, acuminate in fusiform specimens, gradually
attenuate into a concolorous, slightly pruinose stem: flesh pallid, spongy.
CLAVARIA
On the ground: tropical Africa (Congo, Kisantu).
Spores 8-9 X 3·5-4·5 µ., white, smooth, oblong, apex depressed.
Basidia 30-35 X 7-8 µ..
Hyphae 3-9 µ..
Said to be allied with Clavariadelphus ligula, but surely not!
C. nebulosoides Kauffm.
Pap. Mich. Ac. Sci. Arts Lett. 5, 1925, 118: Coker, Journ. El. Mitch. Sci. Soc., 63,
1947, 56, pl. 13, f. 5.
4·5-8 X 0·15-0·2 cm., simple, fleshy, cylindric, wood-brown to fawn: stem 1·5-3 cm.
long, minutely pruinose.
On mosses in coniferous forest: U.S.A. (Mt. Hood, Oregon).
Spores 5-6(-7·5) X 2·5 µ., white, smooth, ellipsoid: (5·5-)6-7'5 X 2·2-3·5 fL (Coker,
type).
Basidia 30 X 5 fL: sterigmata 4.
Cystidia 65-70 X 7-12 µ., lanceolate, hyaline, thin-walled, fairly abundant, with long
slender stalk.
Hyphae -20 µ. wide (Doty).
This seems very near C. purpurea, but it is without purple colour and has smaller
spores and lanceolate cystidia: perhaps, also, it is not caespitose.
C. Pullei Donk.
Rev. Nieder!. Homobas. Aphyll. 2, 1933, 86: FI. Bat. 1929, t. 2156b (ut C. tenacella
Pers.).
-4 cm. X 1·5-3 mm., simple, solitary or gregarious, cylindric, sometimes dilated
upward, subacute or blunt, slightly longitudinally rugulose, dark brown, rufous, or
blackish brown: stem indistinct, short, paler brown, finely cottony, base white: flesh
greyish brown.
Among grass: Holland.
Spores 4·7-6·7 X 4·3 µ., white, smooth, ovoid, mostly 1-guttate.
Basidia 20-36 X 5-6·5 µ. : sterigmata 2 (? always, few basidia seen).
Hyphae 4·5-9 µ. wide, -16 µ. in the central tissue, thin-walled, no clamps seen.
The absence of clamps clearly indicates Clavaria, but the guttate spore suggests
Clavulinopsis. The affinity of the species seems to be with C. atro-umbrina (narrower
spores, more rugulose hymenium). Donk says it is relate.cl with C. Greletii rather than
C. tenacella, but it is not easy to appreciate the difference from this last, of which only
the original description appears to be known.
C. purpurea Fr.
Syst. Myc. 1, 1821, 480: emend. Kauffmann, Pap. Mich. Ac. Sci. Arts Lett. 8, 1928,
144: Wehm., ibid. 20, 1935, 255: Coker, Journ. El. Mitch. Sci. Soc. 63, 1947, 61,
pl. 13, f. 6-15.
Synonyms: C. nebulosa Pk., Bull. Torr. Bot. Cl. 25, 1898, 326: Burt, Ann. Mo.
Bot. Gdn. 9, 1922, 47, t. 8, f. 72 (teste Coker et Wehm.).
C.fumosoides Kauffm., Pap. Mich. Ac. Sci. Arts Lett. 8, 1928, 145 (teste Wehm. et
Coker).
C. fumosa Kauffm., ibid. 1, 1923, 124 (non Fr.).
C. rosea sensu Quel., Bull. Soc. Bot. Fr. 26, 1879, 433 (ut C. rubella Pers.) (non
Fr. emend. Coker).
C. occidentalis Zeller, Mycol. 21, 1929, 97, f. 1: (f. Coker, 1947).
C. purpurea var. australis Coker, Journ. El. Mitch. Sci. Soc. 63, 1947, 63, pl. 13,
f. 16 (sine diagnosi): the form with g(ant spores.
2·5-12 cm. high, simple, generally caespitose, up to twenty in a tuft, sometimes
CLAVARIA 247
gregarious or solitary, purple, greyish purple, brownish purple, smoky brown tinged
purple, often pale or merely brownish, isabelline or clay tinged purplish, lavender or
amethyst, discolouring light ochraceous buff to pinkish buff, white-villous at the base,
1·5-5 mm. wide, subcylindric, slender, acute, then fusiform, compressed, often
longitudinally sulcate, obtuse and hollow, occasionally forked (? connate) near the
base: stem scarcely distinct, paler or light creamy, white-villous: flesh brittle, white or
paler concolorous, the hymenium often peeling off: smell and taste none, or slightly
fishy or nauseous.
Among grass or on bare soil, generally (? always) under conifers: Europe (Gt.
Britain, France, Germany, Denmark, Sweden, Finland), U.S.A. (N. Car., Colorado,
Utah, New Brunswick, Mich., Id., Cal., Or., Wash.), Newfoundland (C. nebulosa).
Spores 5·5-9 X 3-5 µ, sometimes also with 'giant spores' 8·5-13(-15·5) X 3·5-5 µ,
white, smooth, thin-walled, ellipsoid to oblong, slightly flattened adaxially, blunt,
slightly apiculate, granular-guttulate (? not guttate).
TABLE XIV. SPORE-SIZE IN CLAVARIA PURPUREA FR.
Size in µ. Authority Remarks

5·5-7 X3-4 Small and giant spores of C. purpurea from
} Wehmeyer
8-10X4-5 Michigan
5-7X3·5 Kauffmann C. fumosoides from Washington
8-9X3·2-4 Doty C. fumosoides (type)
6-7·5X3-4 Peck C. nebulosa from Newfoundland
6·5-7·5X3·7-4 Coker C. purpurea from Sweden (1923, p. 39)
7-8x4-5 Cotton and Wakefield, C. purpurea from Gt. Britain and France
Rea, Bourdot, and
Galzin
7-g(-10) X3·5-4·5 Kauffmann C. purpurea
7·5-9·2X3-4·5 Wehmeyer C. purpurea from Germany
7·4-1 I X3·6-4 Coker, Kauffmann C. nebulosa (type)
8-9X3 Coker C. purpurea (? C. fumosa) from Colorado
8·5-9X3·5-4 C. purpurea from Utah
8-1 l X3·7-4·8 C. purpurea from Sweden, Finland, and
N. Carolina
9-10X5 Kauffmann C. purpurea (as C. fumosa, 1923)
8·5-13(-15) X3·8-4·5 Coker C. purpurea from New Brunswick
6·5-9•3 X3·5-4 Coker C. occidentalis (type)
Basidia 70-75 X 6-8 µ, with 4 sterigmata (Kauffm. for C. fumosoides).

Cystidia 45-130 X 5-9µ, cylindric or slightly tapered downward, thin-walled,
colourless, obtuse, with granular contents, very abundant, projecting.
Hyphae 3-15 µ wide, cells 50-95 µ long, without clamp-connexions (Coker, for
C. nebulosa) or with very few clamps (Coker, for C. purpurea, but? what is meant),
constricted at the septa.
This seems to be a rare species very variable in size, habit, colour, and spores.
Many more collections are needed before it can be understood. Probably it occurs
regularly L if sparsely, in the northern coniferous forests.
European specimens are said to be purple, purple grey, or brownish purple. In
N. America they are dull brown tinged smoky purple with white flesh (Coker); light
greyish vinaceous, smoky grey with a tinge of purple, concolorous within, lighter
creamy towards the base, becoming pale ochraceous buff to pinkish buff, often with
shades of pale amethyst (Wehm.): isabelline or clay, clouded with darker hues (Peck,
for C. nebulosa): grey-lavender, concolorous within (Kauffm., for C. fumosoides).
Thus there appear to be pale and dark specimens, with or without a purplish hue,
varying to brown (as in C. nebulosoides and C. occidentalis) or grey (as in C. /umosa).
The spore-data are given in Table XIV. At one time it was thought that there were
CLAVARIA
two species, one with spores 7-9 X 4-5µ, the other with long spores 9-13 X 3·5-51i
(Coker, Kauffm.). Wehmeyer found both spores on _some fruit-bodies whereas others
had no 'giant spores'. What proportion they bear to the small spores and whether
there are fruit-bodies with only giant spores are not known. For a discussion of the
two kinds, see Comer (1948).
The presence of cystidia, first reported by Kauffmann, has been confirmed by
Wehmeyer. They are but slightly differentiated from the basidia and, in most cases,
do not exceed them in length, though abundant enough. They occur also in the nearly
allied species C. nebulosoides.
The habitat, according to Coker, is coniferous, as that of C. nebulosoides. If true, it
should suffice to distinguish them among true Clavarias.
C. rosea sensu Quel. ( = C. rubella Pers. ap. Quel.) was described as: 2-3 cm. high,
compressed, fusiform, optuse, hollow, fragile, purple then ochraceous: stem fistular,
3 mm. long, rooting, undulate, satiny, rosy, pruinose and white at the base: spores
10µ long, ovoid-pruniform, white: in troops in mossy pastures, Jura. The colour and
habit agree with C. purpurea rather than C. incarnata.
C. rosea Fr.
Syst. Myc. 1, 1821, 482: emend. Coker, Clav. U.S. Can. 1923, 40: (C. rosella of
Bourd. et Galz., in error).
(C. rosea Fr. sensu Clel. et Chee! = Clavulinopsis corallino-rosacea Clel.: sensu
Quel. = ? C. purpurea Fr.)
2-5 cm. high, simple, solitary or in groups of two to seven, bright rose-pink, clear pinh,
or reddish pink, discolouring yellowish at the apex, 1-5 mm. wide, cylindric, acute then
fusiform and blunt, sometimes flattened and shortly bifurcate or dentate (as a cocks
comb): stem short, fairly distinct, pallid(? not strigose): flesh brittle, paler concolorous
or white· and deep rose beneath the hymenium: smell and taste none, or slight and
inparticular.
On the ground among grass and in woods: Europe(Sweden to France and England),
U.S.A. (N.Y., Colorado), rare.
Spores 5-8 X 2·5-3·5µ, white, smooth, ellipsoid, blunt, slightly flattened adaxially
(? guttate).
Basidia 7 µ wide: sterigmata 4 (Coker).
Hyphae 11µ wide, without clamps (Coker).
var. subglobosa var. nov.
= C. rosea Fr. sensu Bres., Rea, Ricken, et sensu Pers. (fide Coker).
Spores 5-8 x 4-6µ, subglobose or broadly ellipsoid.
Germany, England(? France, Baria, No. 18�9, Nice, with sp. 6-8·5 X 4-5·5 µ), rare.
As explained on page 215, I have followed Coker's interpretation of the species.
Cooke's record "from Australia, and his variety attenuata, need to be checked: the
spores are given as 2µ wide, which is surely in error (Handb. Austral. Fungi 1892,
202). Compare Clavulinopsis miniata with var. subglobosa.
C. rufobrunnea Coker.
Journ. El. Mitch. Sci. Soc. 63, 1947, 43, pl. 13 (f. 1, 2).
-5 cm. high X 2-4 mm., simple, caespitose, terete, natal brown (Ridg.), drying dark
brown to nearly black and wrinkled, the stem indistinct, paler at the base: smell none,
taste inparticular.
On earth under conifers: U.S.A. (California, A. H. Smith 9403, type: Smith 9467).
Spores 3·8-5 X 3-3·5 µ, white, smooth, ovoid.
Basidia 7·5-8 µ wide, 4-spored.
Hymenium c. 70µ thick (very dark in dried specimens).
Said to be near C.fumosa and C. nebulosoides, but compare C. Pullei also with ovoid
and slightly larger spores.
CLAVARIA 249
C. scabra B.
Hook. Joum. Bot. 8, 1856, 277.
Burt, Ann. Mo. Bot. Gdns. 7, 1922, 71, t. 11, f. II2.
-8 mm. high, gregarious, subcaespitose, simple, erect, acuminate, pale umber (? dried),
scabrous with little rough granules: base tuberous, clothed with white or pallid erect
bristles.
On the ground: Brazil (Panure, Spruce r 57).
Spores ?
Hyphae 2-5(-7)µ, wide, sparsely secondarily septate, without clamps, with thin or
slightly thickened walls, rather long-celled and scarcely inflated.
I have examined the type at Kew. It must be closely allied with C. fossicola and
C. helicoides. Burt found smooth, white, subglobose spores, 1·5-2µ, wide on the type,
but doubted whether they belonged to the fungus. 'There is another simple Clavaria
in the collection, growing on a green substance, which appears to be an anamorphosis
of some lichen. The specimens are, however, too imperfect to afford much informa
tion.' (Burt.)
C. straminea Cott.
Tr. Brit. Myc. Soc. 3, 1910, t. I r (labelled C. persimilis in error).
Plate r.
1·5-5 cm. X 1-4 mm. simple, solitary or tufted, cylindric or compressed, acute,
straw colour, brownish with age or when handled: stem -1 cm. long, usually very distinct,
cinnamon-yellow: flesh somewhat darker than the hymenium, brittle: smell and taste
inparticular.
Among short grass: Gt. Britain, Holland (rare).
Spores 5-7µ, wide, white, smooth, globose, granular (Cott.), occasionally 1-guttate
(Donk).
Basidia 40-60 x 7-9µ,, 4-spored (Cott.): 25-50 X 8·5-10µ,, with 2-4 sterigmata
8-10µ, long (Donk).
Hyphae 4-12µ, wide, without clamps (Donk): short-celled and without clamps
(Comer, co-type in herb. Kew.).
From the macroscopic description this appears to be a yellow Clavulinopsis, but the
hyphae are typical of Clavaria. The colour is probably an acropetal oxydative dis
coloration and not a true pigmentation. Coker compared it with the simple form of
Clavulinopsis corniculata, which has different hyphae, but Donk noted the brittle
flesh characteristic of Clavaria. It is either a yellowing C. acuta or a globose-spored
C. argillacea.
C. striata Fr.
Syst. Myc. 1, 1821, 483: Pers., le. et Descr., t. 3, f. 5.
3-5 x 0·3-0·4 cm., subfuliginous or bistre greyish, white at the base, cylindric, here
and there longitudinally striate, subobtuse, becoming hollow, caespitose, rather brittle.
Among grass under Fagus: Gt. Britain (rare), Holland.
Spores 3-4 X 2-3 µ,, white, smooth, subglobose.
This is Rea's description of the species. Cotton and Wakefield suggest C. striata is
a discoloured C. vermicularis. Bourdot and Galzin regard C. striata as a form of
C. fumosa. Rea's fungus has smaller spores (the locus of which on the sporograph lies
on the backward extension of the C. fumosa-line). Compare C. Crosslandii.
C. subacuta Ito et Imai.
2-4 X o· 1-0· 15 cm., simple, fasciculate, white, terete, attenuate or obtuse at the apex:
stem 0·6-1·5 cm. long, distinct, more slender than the club, transparent white.
On the ground in shady woods: Bonin Is!.
CLAVARIA
Spores 5 x 3µ, white, smooth, ellipsoid or subovoid.
But for the long slender stem I would regard this as C. vermicularis. It may be
related to C. filiola which has smaller fruit-bodies and rounder spores.
C. tenacella Fr.
Syst. Myc. 1, 1821, 472.
Simple, fasciculate, obtuse, rufous Juliginous, rufous bistre or greyish, base whitish,
erect, subobtuse, rather tough.
On earth and heaths: France, Sweden.
Little known. Compare C. Pullei. The tough flesh is more like Clavulinopsis than
Clavaria.
Joachim's collection from France, identified as C. tenacella by Boudier, must have
been a Clavulina, possibly C. cinerea f. sublilascens (see p. 311). Perhaps this may be
the true explanation of C. tenacella.
C. tenuipes B. et Br.
Ann. Mag. Nat. Hist. 2, 1848, 266, t. 9, f. 2.
Pistillaria tenuipes (B. et. Br.) Mass., Brit. Fung. FI. 1, 1892, 91.
? Pistillaria furcata W. G. Sm. ex Cke., Handb. Brit. Fungi 1870, 343.
Text-Figs. 89-<)2.
1-6 cm. X 1-3 mm., simple, solitary or gregarious, cylindric then subclavate, acute
then blunt, often compressed 3-10 mm. wide, solid and fleshy or eventually hollow,
whitish, pallid clay, greyish yellow, pale grey, or drab: stem 5-20 X o·5-3 mm., distinct,
subtranslucent, pallid whitish: flesh soft and brittle, becoming rather firm: smell none.
On the ground, on charcoal, sticks, or humus: Europe (Gt. Britain, France).
Spores 6-12 X 4-5·5 µ, very variable in size, white, smooth, oblong ellipsoid, blunt,
often subventricose, thin-walled, tnultiguttulate.
Basidia 30-40 X 8-9 µ, clavate, multiguttulate, shortly but distinctly bifurcately
clamped at the base: sterigmata 4, 5 µ long.
Cystidia none.
Hymenium c. 35µ thick, not thickening: subhymenium 3-40µ thick, composed of
narrow, loosely interwoven hyphae 3-5 µ wide. Hyphae
7-25µ wide (-30µ in the stem), the cells 100-250µ
Jong, shorter-celled (20-120µ), narrow, and more fre
quently branched near the subhymenium, tapered
to the septa, very occasionally with a clamp, not
ThxT-FIG. 89. = Clavaria tenuipes, secondarily septate, thin-walled, smooth, with fairly
x 1• 000 •
numerous, generally interweaving, narrow hyphae
1·5-5 µ wide, some with oleaginous contents like laticiferous hyphae: the cells short,
20-50 X 8-25µ, and pseudoparenchymatous at the base of the stem.
(Caulocystidia?: see below.)
I have had only dried material of this species for examination, but I am satisfied
that it belongs to the subgen. Holocoryne. Indeed, it resembles the scattered or
solitary states of C. Gibbsiae so closely that it would seem distinguishable only by the
drab or greyish appearance, the narrow spores and smaller basidia, and, as in C.
Gibbsiae, the spores are variable. Rea gives them as 6-9 x 4-5 µ, but I have seen them
also 9-12 X 4·5-5·5µ in collections identical in all other respects.
Massee placed the species in Pistillaria, but it has none of the characters of that genus:
its hyphae are not agglutinated at the surface of the stem, its spores are multiguttulate,
and only its basidia are clamped.
Pistillaria furcata is poorly described, but may be C. tenuipes. Its description is
merely: 2·5-3 cm. high, white or yellowish, compressed, broad at the apex, attenuate
downward, generally furcate, waxy then tough: caespitose, in cold greenhouses (? on
the ground): England.
CLAVARIA 251
I have also found in England, on the dead leaf-bases of Cladium, a funguf which
seems to be C. tenuipes but which has longer cells and distinct caulocystidia, which
I have been unable to make out in the dried specimens of C. tenuipes. Should it be
another species, I give its description separately so as to avoid confusion:
C. tenuipes?: superficial on dead leaf-bases of Cladium mariscus: England (Wicken
Fen, 22.8.26). (Text-Figs. 96-8.)
FIG. 90. FIG. 91.

TEXT-FIG. 90. Clavaria tenuipes: fruit-bodies from Wicken Fen, England, on Cladium
(22.viii.26), X 5.
TEXT-FIG. 9r. Clavaria tenuipes: collection from Wicken Fen, England (22.viii.26): X1,000.
9-15 x 1-1·3 mm., simple, fusoid and acute, then clavate and blunt, white: stem
3-5X0·4-0·7 mm., distinct, puberulous.
Spores 8-10X4·5-5·5µ, as in C. tenuipes.
Basidia 32-40X8-9µ, as in C. tenuipes: 4 sterigmata 5µ long.
Hymenium as in C. tenuipes, not thickening.
Hyphae as in C. tenuipes, but the cells 100-500µ long.
Caulocystidia -Sox 5-8µ, subcylindric to clavate or ventricose, obtuse or subacute,
walls thin or very slightly thickened, smooth, aseptate, colourless, as the out-turned
ends of sterile hyphae, numerous but not forming a close pile, reduced at the apex of
the stei;n to blunt processes 3-15 x 3-5µ, at the base of the stem elongating iuto
filiform hairs, -200X1·5-2·5µ, with slightly thickened walls.
C. vermicularis Fr.
Syst. Myc. r, 1821, 484: Konr. et Maubl., le. Se!. Fung., t. 493: Bres., le. Myc. 22,
1932, t. I 100 (ut C. fragilis): v. Ov., le. Fung. Mai. 4, 1923 (ut C. vermiculata Mich.).
CLAVARIA
TEXT-FIG. 92. Clavaria tenuipes: collection from Wicken Fen, England (28.viii.26); middle
and basal parts of the stem with caulocystidia: X 500.
Synonyms: C. fragilis Fr., Syst. Myc. 1, 1821, 484 (teste Cott. et Wakef., Bourd.
et Galz., Konr. et Maubl., Coker, Donk).
C. cylindrica S. F. Gray, Nat. Arr. 1, 1821, 656.
C. solida S. F. Gray, ibid. 656.
C. alba Pers., Myc. Eur. 1, 1822, 175 (teste auctt. cit.): (non C. alba Pers., ibid.,
p. 161 = Clavulina cristata var. coralloides).
CLAVARIA 253
C. eburnea Pers., Myc. Eur. 1, 1822, 183 (teste auctt. cit.).
C. pistilliforma Pers., Myc. Eur. 1, 1822, 183 (teste auctt. cit.).
C. vermiculata Pers., Myc. Eur. 1, 1822, 184 (teste auctt. cit.).
corynoides Pk., Rept. N.Y. St. Mus. 31, 1879, 39 (teste Coker).
,.1·
C.
' .
\: !
.. ,
;:�-·
·r··.,.
TEXT-FIG. 93. Clavaria vermicularis: spores and basidia, X 1,000: hyphae of flesh, X 500.
C. simplex Karst., Hattsv. 2, 1882, 172 (teste Ito et Imai).

C. gracilior Britzm., Hym. SUdb. 7, 1891, 15, f. 74: ibid. 9, 22, f. 84: Rev. Hymen.
1909, 223.
C. Muelleri B., Grev. 20, 1891, 10. ra(;} /Q)
'8 D· ·.
C. nivea Quel., Ass. Fr. 1901, 496, t. 3, Q @ C Q
f. 11 (non Pers.).
C. vermiculata Mich. ap. v. Ov., Bull.
Jard. Bot. Buit. Ser. 3, 5, 1923, 268: (non
sensu Petch = Clavulinopsis spiralis).
Misinterpretation: C. vermicularis sensu
v. Ov., Bull. Jard. Bot. Buit. Ser. 3, 5,
�
o.·> (�
1923, 268: le. Fung. Mai. 4, 1923: ( = TEXT-FIG. 94. Clavaria vermicularis: above,
spores of a Javanese collection (from van
0\'ereem): below, var. singaporensis: x r ,coo.
Clavulinopsis spiralis).
Text-Figs. 93, 94.
6-12(-15) cm. high, simple, densely caespitose, white, very brittle, without smell or
taste: 3-5 mm. wide, cylindric then elongate fusiform, becoming flattened and sulcate,
often curved or flexuous, occasionally once furcate, solid then generally hollow, acute
becoming obtuse, often yelloic:ish towards the tip, wholly pale yellowish with age: stem
indistinct, as a short, slightly narrower, sterile basal part of the fruit-body: occasionally
gregarious, or in small fasciculate clusters of three to six, rarely solitary.
Among grass or on bare earth in fields and woods: Europe, temperate N. America,
China, Japan, Bonin Is!., Ceylon, Java, S. Africa, Australia (Queensland, N.S.W.,
Victoria): generally common in temperate regions, edible.
Spores 5-7 X 3-4µ., white, smooth, ellipsoid or pip-shaped, generally aguttate or
very finely granular guttulate, sometimes 1-guttate, thin-walled, shortly apiculate.
254 CLAVARIA
Basidia 30-45 X 6-8 /J,, clavate, finely multiguttulate, without clamps: sterigmata 4,
4-5 fL long: (34-54 X 5-6 fL for Faeri:ies, 38-42 X 7-8 fL for Denmark, according to
Moller).
Cystidia none.
Hymenium not thickening: subhymenial hyphae I· 5-2·5 fL wide, closely interwoven,
not inflating.
Hyphae of the flesh 3-16 fL wide, the cells 25-80 fL long (occasionally up to 150 fL
long), short-celled, the narrow hyphae with longer cells, slightly constricted at the
septa, secondarily septate, without clamps, thin-walled, longitudinal, compact.
f. fasciata Bourd. et Galz.

Hym. Fr. 1, 1928, I 10.
Caespitose, 3-:5 cm. high, cylindric, mostly connate forming once- or twice-forked
branches.
Spores 4·5-6 X 2·5-4,_,,.
France (uncommon).
var. gracilis Bourd. et Galz.

Hym. Fr. 1, 1928, no.
Synonym: C. gracilis Sow. ex Pat., Tab. An., t. 33 (non Fr.).
Smaller, less fasciculate, gregarious or occasionally solitary, with a distinct pellucid
stem: spores slightly smaller.
Fruit-bodies 2-5 cm. high: spores 4·5-5 X 3-4,_,,, broadly ellipsoid or ovoid.
France (frequent): U.S.A.: Malaya (Singapore, No. 5929, Bot. Gdns. 24.7.20).
var. singaporensis var. nov.

Spores 7-8· 5 X 5-6 /J,, considerably larger than typical spores, white, smooth, thin
walled, pip-shaped, attenuate strongly to the apiculus, with I large gutta 3-4,_,, wide.
3-4 cm. X 2 mm., simple, gregarious, not fasciculate, white, cylindric, apex acute,
waxy brittle: stem I cm. X 1·5 mm., more or less distinct.
Basidia c. 40 X 7-8 /J,, without clamps at the base: sterigmata 2-4, 6-7 fL long.
Cystidia none.
Hyphae 4-13 fL wide, cells 20-70 fL long, as in the typical state, without clamps.
On mossy soil in the shade of trees: Singapore, Bot. Gdns., May 1929.
var. sphaerospora Bourd. et Galz.

Hym. Fr. 1, 1927, no (ut forma): Teng. Sinensia 7, 1936, 254 (ut C. vermicularis).
3-4(-7) cm. high: spores 3-6 1-L wide, subglobose, a few pyriform 4·5-5 X 3/J,:
(sp. 4-5 X 3-41-L, Teng).
Europe, U.S.A., China (Kiangu, Chekiang, Yunnan), Japan, Bonin Isl., S. Austral.
(Victoria).
KEY TO THE VARIETIES

Caespitose: stem indistinct
6-12 cm. high: sp. 5-7 X 3-41-L C. vermicularis
-5 cm. high, fasciate and branched: sp. 4·5-6 X 2·5-4/J, f. fasciata
Gregarious, solitary or subfasciculate, 2-6 cm. high: stem generally
distinct, pellucid
Sp. 7-8·S X 5-6 /J,, pip-shaped, I -guttate var. singaporensis
Sp. 4·5-5 X 3-41-L, broadly ellipsoid var. gracilis
Sp. 3-6 µ, subglobose var. sphaerospora
CLAVARIA 255
This common and, apparently, cosmopolitan species is the type of the genus. It is
variable in many respects and needs much detailed study.
Colour. The fruit-bodies often acquire yellowish tips as they age, but they have no
yellow pigment. Coker describes them as often becoming creamy flesh colour in the
lower part, which suggests faint pigmentation as in C. incarnata. Tropical records
require verification because the species has been confused with Clavulinopsis spiralis
which varies into the yellow C. amoena and the pink or red C. miniata.
Size. Typically, in Europe, the fruit-bodies are 6-10 cm. high. The N. American
specimens described by Coker are mostly 1-6 cm. high, though Wehmeyer describes
others as 4-9 cm. high and Doty gives the extraordinary limits of 15 and 25 cm.,
which may refer to the typical state or to var. sphaerospora. The size of the fruit-body
has little meaning, however, until analysed into the causal factors.
Habit. Typically the fruit-bodies are densely caespitose in tufts of 20-50, but there
are gradations to small tufts of 3-6 fruit bodies, gregarious and solitary fruit-bodies.
Generally the smaller fruit-bodies are the less caespitose, as if developed on a poorer
substratum. The caespitose habit has not the importance given to it by Fries in his
subgenus Syncoryne.
Spores. The data assembled in Table XV lead to these conclusions:
1. There is a gradation from the short subglobose spores of var. sphaerospora to the
most elliptic spores of var. singaporensis.
2. The N. American collections have slightly smaller spores than the European,
though both represent, apparently, the typical form of the species.
3. There may be a variety with unusually narrow spores, as shown by the last four
measurements in the table.
TABLE XV. SPORE-SIZE IN CLAVARIA VERMICULARIS
Mean Mean length

Spore-size in µ, length Mean width
=
E Authority I Remarks
7-8·5X5-6 7·8 1·4 Corner var. singaporensis
6-9X3-4 7·5 2·1 Boudier France
6-8X3-5 7·0 1·8 Lindau Germany
6x3·5 6·o 1·7 Barbier France
4·5-7X2·8-4·5 6·o 1·6 Comer Gt. Britain
(Table XVI)
5-7X3·5-4·5 6·o 1· 5 Konrad, France
Maublanc
5-?X3-4 6·o 1·6 Rea Gt. Britain
5-6·5X3·5-4·3(-4·7) 5·8 1·5 Donk Holland
5-6(-7'5)X3-4(-5) S'5 1·6 Bourdot, France
Galzin
(Average for European collections, 6X3·8, E = 1·6)
5-6X3 5·5 1·8 Petch Ceylon
5-6X3-4 5·5 1·6 S. Africa
v. d. Bijl
4·5-6X2·5-3 5·3 1·9 Coker U.S.A.: No. 2335
( = C. corynoides Pk.)
4-6·6X2·2-2·9 5·3 2·1 U.S.A.: No. 2759
4·4-6X 2·6-3·6 5·2 1·7 U.S.A.: No. 2751
4·5-5·5X2·8 5·0 !'8 U.S.A.: No. 4932
5X3 5·0 1·7 California, s.n.
4·3-5·5X2·3-3·3 4·9 1·8 U.S.A.: No. 28
4·4-5·2X2-2·6 4·8 2·1 U.S.A.: No. 2773
4·4-5X2·5-2·9 4·7 1·7 U.S.A.: leg.Earle (N.Y.)
4-5X 2·5-2·8 4·5 1·7 U.S.A.: No. 1786
(Average for N. American collections, 5·0X2·74, E = 1·82)
CLAVARIA
TABLE XV (continued)
Mean Mean length
Spore-size in µ. length Mean width
--- =E Authority Remarks
4·5-6X2·5-4 5·3 1·6 Bourdot, f. fasciata: France
Galzin
4•5-5X3-4 4·8 1·4 Bourdot, var. gracilis: France
Galzin
4·5-5·5X3·5 5·0 1·4 Wehmeyer var. gracilis: U.S.A.
4-6X3-4 5·0 1·4 Bresadola var. gracilis: Italy
(Average for var. gracilis, 5·ox3·5, E = 1·43)
5-6x4-5 5·5 1·2 Ito, Imai var. sphaerospora:
Bonin Isl.
3-6, subglobose 4·5 I "I Bourdot, var. sphaerospora:
Galzin France
3-6X3-4 4·5 1"3 Imai var. sphaerospora:
Japan
4-5X3·5-4 4·5 1·2 Lundell var. sphaerospora:
Sweden
4-5X3-4 4·5 1'2 Teng var. sphaerospora: China
3-5X3-4 4·0 1·14 Cotton, var. sphaerospora:
Wakefield Gt. Britain
4·5-5 4·8 I' I Doty var. sphaerospora:
Oregon
3-5X3-4 4·0 1·14 Fawcett var. sphaerospora:
Victoria
(Average for var. sphaerospora, 4·6X3·9, E = I· 18)
5·6-6·7 X2·2-2·6 6·2 2·6 Coker U.S.A.: No. 2434
5X2 5·0 2·5 Ricken Germany
5·5X2 5·5 2·8 Cleland S. Australia
5-5·7 X2·3-3 5·35 2·0 van
Overeem Java
The normal variation in the spores of a single fruit-body is shown in Table XVI.
Synonymy. The Friesian name is C. vermicularis, the alternative C. vermiculata
being pre-Friesian and invalid. C. fragilis is generally accepted as a synonym, but
specimens of C. acuta have been misnamed C. fragilis. It appears that Schroeter first
reduced the two species and gave C. vermicularis as the synonym (Krypt. FI. Schles.,
p. 445). Massee's record of C.fragilis probably refers to C. Gibbsiae (Kew Bull. 1901,
154). Regarding C. Muelleri, from Victoria and Queensland, I have examined the
type at Kew and find that it is typically C. vermicularis with spores 6-7 X 3-3·5 µ.
Concerning the species in Ceylon and Java there has been much confusion with
Clavulinopsis spiralis, which may be almost identical macroscopically. Thus,
C. vermiculata Mich. sensu v. Ov. is C. vermicularis Fr. (? var. singaporensis).
C. vermiculata Mich. sensu Petch is Clavulinopsis spiralis.
C. vermicularis Fr. sensu v. Ov. is Clavulinopsis spiralis.
I doubt if C. affinis, C. citriceps, and C. subacuta are separable specifically from
C. vermicularis.
C. xylarioides Petch
Ann. Roy. Bot. Gdns. Per. 7, 1922, 291.
-4 cm. high, 4 mm. wide above, 2 mm. below, simple, clavate, terete, obtuse, violet
black, black at the base: flesh purple brown with violet black outer zone.
On the ground: Ceylon (Peradeniya, Dewalakanda).
Spores 9-10 x 7 µ, broadly ovoid; or 8-9 µ, globose, white, smooth.
CLAVARIA 2 57
Basidia -40 X 8µ below, rzµ above: sterigmata 4, roµ long, conical for half their
length, hair-like distally.
Hyphae inflated, closely septate, without clamps (Petch 4389, 6096 in herb. Kew.).
I have examined the co-type and a later collection of this in the Kew herbarium.
TABLE XVI. CLAVARIA VERMICULARIS (ONE FRUIT-BODY, ENGLAND)
Spore-size in µ. Length -:- Width ( = E) Number of records

4·5x2·8 1·6 l
4·8 X3·0 l ·6 2
5·0X3·0 1·7 6
5·ox 3·5 1·4 7
5'3X3'5 1·5 3
°
5·5xJ -5 1·6 5
5· 5x4·0 1·4 l
6·ox3·5 1·7 6
6·ox3·7 1·6 l
6·0X4·0 1·5 6
6·5X3·5 1·9 2
6·3X3·5 1·8 l
6·5X3·0 2·2 l
6·5X3·5 1·9 5
6·5x3·7 1·8 l
6·5X4·0 1·6 4
6·5X4·5 1·4 l
6·7 X4·0 1·7 4
6·7 X3·7 1·8 l
7·0X4·0 1·8
-2
60
5·3-5·5 µ.
6·0-6·3 µ.
...
4'$-5·0 µ. long 1·53
1·50
1·62
.."
Average of 16 spores
9
14
.."
6·5 µ. long
6·7-7·0 µ. "
7·ox4·3
1·76
1·70
1·6 3
,,
"
"
14 "
7
60
....
ThxT-FIG. 95. Clavaria Zollingeri: various forms of fruit-bodies

(after several authors): X I.
SIIQ S
258 CLAVARIA
It is a true Clavaria and appears to be near to C. Zollingeri but simple and with larger
spores.
C. Zollingeri Lev.
Ann. Sci. Nat. Ser. 3, 5, 1846, 155: emend. v. Ov., Bull. Jard. Bot. Buit. Ser. 3, 5,
1923, 270: le. Fung. Mai. 4, 1923, f. 4.
Synonyms: C. amethystina Zoll., Geneesk. Arch. Neer!. Ind. 1, 1844, 381: (non
Fr., 1821 = Clavulina amethystina).
? C. lilacina Jungh. ex Lev., Ann. Sci. Nat. Ser. 3, 2, 1844, 216.
? C. arborescens B., Fl. N. Zeal. 2, 1867, 186.
C. lavendula Pk., Bull. N.Y. St. Mus. 139, 1910, 47: Burt, Ann. Mo. Bot. Gdn. 9,
1922, 47, t. 9, f. 73: Donk, Rev. Nieder!. Homobas. 2, 1933, 23, 75: Wehm., Pap.
Mich. Ac. Sci. Arts Lett. 20, 1935, 256.
C. Nymaniana P. Henn., Monsunia 1, 1900, 9 (teste v. Ov.): Fawcett. Proc. Roy.
Soc. Viet. 51, 1939, 8, t. 4, f. 2.
C. bicolor Mass., Kew Bull. 1901, 54.
C. rosalana Petch, Ann. R. Bot. Gdn. Per. 7, 1922, 290.
C. violacea Petch, ibid. 290: ibid. 9, 1925, 332: (fide Petch, v. Ov.).
Misinterpretation: C. amethystina Fr. sensu Coker, Clav. U.S. Can. 1923, 90, t. 24,
25, 28, 84: sensu Cott. et Wakef., Tr. Br. Myc. Soc. 6, 1919, 180: sensu Ito et Imai,
Tr. Sapporo Nat. Hist. Soc. 15, 1937, 52-3: ? sensu Quel., Bres., Rea, vel pr. p.
1·5-7·5 cm. high, generally caespitose, varying gregarious or solitary, copiously
branched or sparingly, even almost simple, deep amethyst or
violet, varying greyish or brownish violet, lilac pink, or
vinaceous: stem 0-3 cm. x 2-3 mm., variable, often branched
from the base, paler, often greyish or pallid white, yellowish
with age: branches 1-3 mm. wide, or 3-5 mm. wide below
on massive specimens, 1-4 times divided, polychotomous below
in massive specimens, blunt or becoming subulate above,
often few and irregular, rarely absent, more or less fastigiate,
axils rounded: flesh concolorous, brittle, yellowing on drying:
taste and smell none, or slightly farinaceous (Wehm.), or smell
strong and taste of tallow (Cott. et Wakef.).
On the ground in woods or under trees in the open: Europe
(France, Gt. Britain, Italy), U.S.A., Canada (Nov. Scot.),
Ceylon, Malaya, Java, Bonin Isl., S. Austral. (Victoria).
Spores 4-7 X 3-5µ., white, smooth, broadly ellipsoid or
subglobose, thin-walled, aguttate (1-guttate, Fawcett).
Basidia 40-60 X 6-9µ. (28-50 X 4·5-7µ., Bourd. et Galz:
50-60 X 7-10µ., Cott. et Wakef.), clavate, multiguttulate:
sterigmata 4, rarely 2 or 3, 4-7µ. long.
Hymenium thickening to 100µ., at first c. 40µ. thick:
subhymenium c. 20µ. thick, the cells at first narrow,
10-25 X 1·5-4µ., but gradually inflating, 4-10µ. wide, as the
hymenium thickens, and often subglobose, never pseudo
parenchymatous.
Hyphae monomitic, without clamps, becoming frequently
TEXT-FIG. 96. Clavaria secondarily septate, the cells 20-120 x 4-25 µ, more or less
Zollingeri: subsimple constricted at the primary septa, not at the secondary septa,
form (Malayan form A,
of the text): x 2. walls firm and very slightly thickened, smooth, H-connexions
frequent.
This appears to be a cosmopolitan fungus and one of great variability. It is not
common and there is need for the detailed description of nearly every collection before
CLAVARIA 259
one can divide it satisfactorily into varieties. It is the only branched species of
Clavaria s. str., other than C. fossicola, yet it varies into the almost simple caespitose
state of C. xylarioides.
Shape. Typically it is fairly massive, possibly reaching 10 cm. high, if Henning's
measurements for C. Nymaniana are correct. Slender fruit-bodies are regularly and
alternatingly dichotomous throughout. Simple or sparingly branched fruit-bodies
often become fusiform and compressed.
TEXT-FIG. 97. Clavaria Zollingeri, X 500.
Colour. Deep amethyst violet seems to be the typical full pigmentation, but
brownish and greyish tints creep into the paler forms. How the pigment is distributed
is not known.
Spores. The measurements of different authors are given in Table XVII. Their
shape, whether subglobose, ovoid, or broadly pip-shaped, may be varietal, but the
different shapes and sizes seem variously combined with the shapes and colours of the
fruit-body. Van Overeem's specimens from Java have very narrow spores.
Malayan forms. I describe these two, depauperate, states for reference.
A. 2-4 cm. high, 2-4 mm. wide, simple or occasionally with one branch, brownish
· lilac, or brownish violet to violet, often flattened and rather fusiform: stem 2-5 X 1 · 5 mm.,
short, indistinct: spores 5-6·5 X 3-3·5 µ, ellipsoid, blunt, vacuolate: hyphae -25 µ wide:
caespitose, on the ground in virgin forest, Bukit Timah, Singapore (1935-41): (Text
Fig. 96: Plate 1).
B. -3·5 cm. high, sparingly branched, pale lilac pink, apices pallid whitish, withering
260 CLAVARIA
brownish: stem practically none: branches 2-3 · 5 mm. wide, cylindric, clavate or
flattened, once or twice dichotomous, rather stout, arising from a common base: spores
5-6·5 x 4-5 µ., broadly pip-shaped to subglobose, with wholly opalescent vitreous con
tents: hyphae -13 µ.wide: caespitose, in grass under Dacrydium elatum (Coniferae),
Penang Hill, 700 m.: (Plate 1),
TExT-Fm. 98. Clavaria Zollingeri: part of an old hymenium with

inflated subhymenial cells: x 1,000.
Form A approaches C. xylarioides but is paler and has much smaller spores. It is
possibly C. bicolor Massee, which was said to reach 7-8 cm. high and be pallid umber
with amethyst tips (Gardens' Jungle, Penang).
Form B may be C. rosalana Petch, described as reaching 4 cm. high, pale rosalane
purple (Ridgway 69' d) (or deep magenta pink) with spores 5-7 X 3-4µ. (in the type
in herb. Kew., not 3-4µ., subglobose, as stated by Petch).
Nova Scotia form. The specimens described by Wehmeyer from Nova Scotia
reached 5 cm. high and were sparingly branched, deep amethyst violet, caespitose,
with ellipsoid spores 5-6 x 3·5-4µ.. They resemble the English specimens macro-
CLAVARIA 261
scopically, but the spores of the English specimens are larger, 5-7 p, long, and sub---·
globose.
Synonymy. This species has been described from Europe as C. amethystina, and
has been confused with the true C. amethystina Fr.which is a Clavulina. It seems also
to have been mistaken for Ramariopsis pulchella. From N. America it has been
described as C. lavendula, from Java as C. Zollingeri, from Malaya as C. bicolor, and
from Ceylon as C. rosalana and C. violacea. Because of its variability and because of
the inadequacy of the older descriptions, the correct name is still uncertain. I have
followed van Overeem, with whom Donk agrees. Van Overeem recognized the violet
Clavaria at Buitenzorg as C. Zollingeri, which was described from Java as C. amethy
stina Zoll. (non Fr.). Petch queried this identification but agreed that his C. violacea
was the same· .as van Overeem's fungus. Petch remarked that Leveille wrote of
C. Zollingeri 'les rameaux ...d'une belle couleur rosee tirant. sur le violet', and that
such was not the colour of C. violacea or of C. Zollingeri sensu v. Ov. However, in
view of the variation in colour of C. lavendula given by Coker and Wehmeyer, and of
the Malayan specimens which I have studied, there seems little force in Petch's
objection.
TABLE XVII. SPORE-SIZE IN CLAVARIA ZOLLINGER!
Mean
Size in µ, length E Remarks
4-'7X3-4 5·5 I"57 France: Bourdot et Galzin
5·5-6·5X5 5·75 1· l 5 Gt. Britain: Comer
5-7, subglobose 6·o l· l ))
Cotton et Wakefieid
5-6X 3·5-4 5·5 1·47 Nova Scotia: Wehmeyer
5·5-6·7X3·3-3·7 6·1 1·74 North Carolina: Coker (2622)
5·5-6·5X 3·5-4·5 6·o 1·60 )) )) ))
(4363)
6X3·5 6·o l '71 Alabama: Coker (leg. Earle)
7·2X3 7·2 2·40 Mass.: Coker (leg. Davis)
3·7-6X3·3-4 4·85 1·33 U.S.A.: Coker (as C. amethystina)
6x3 6·o 2·0 (type of C. lavendula)
(Average for N. American collections 6·1X3·5, E = 1·74)
5-7 X 3-4 6·o l ·71 Ceylon: Corner (C. rosalana)
4·5-6·5 X2·5-3 5·5 1·95 Java: van Overeem
4·5-5, subglobose 4·7 ;/.'I ,, Hennings (C. Nymaniana)
5-6·5X3-3·5 5·8 1·77 Malaya: Corner (form A)
5-6·5X4-5 5·8 1·28 ,, ,, (form B)
4-6X3·5-4 5·0 I '33 Bonin Is!.: Ito and Imai
3-5·5, subglobose 4·3 I· I Victoria: Fawcett (C. Nymaniana)
The earliest legitimate name may be C. lilacina Jungh., described as 'Pallide violacea,
stipite brevi nudo, ramis ramulisque compressis, dichotomis sulcatis patulis, axillis
dilatatis, apicibus bifidis. Hab. Java, ad truncos.' Except for the lignicolous habitat,
it answers well to C. Zollingeri, but I think it must be proved that C. Zollingeri may
be lignicolous before the identity can be clinched.
I have examined the type-specimens of C. bicolor, C. rosalana, and C. violacea in
the Kew herbarium, and also authentic specimens of C. amethystina sensu Cott. et
Wakef. At most they are form- or colour-varieties.
C. arborescens seems identical. Its description is:
Amethyst: stem 2-3 cm. high, slightly wavy, slightly thickened upward, slender,
dividing at the apex into a few main branches with short fastigiate branchlets at the
tips: spores 6 x 4 µ,, hyaline, elliptic: New Zealand (N. Is!.: Bay of Islands): on the
ground.
262 CLAVARIA
Similarly C. Nymaniana from Java seems identical, though said to reach 8-10 cm.
high. The S. Australian specimens referred to C. Nymaniana by Fawcett are typical
C. Zollingeri.
DUBIOUS SPECIES
C. aculeata Blonski.
Wyniki. Posz. Floryst. 1890, 17.
Very much branched, delicate, pinkish white: branches terete, smooth, erect:
branchlets attenuate, mucronate, toothed with spines on all sides, concolorous or
pinker, dense and regular.
On rotten trunks, in the mountains: Poland.
Spores white.
Perhaps related to Clavulinopsis virgata. ? Hydnoid.
C. anomala Fr.
Syst. Myc. 1, 1821, 480.
Connate-caespitose, single, strongly clavate, subfusiform, large, flesh colour
becoming pale, fleshy, fragile.
On the ground in oak-woods: Sweden.
This has not been recognized since the time of Fries. It seems to be a caespitose
state of C. incarnata, or a pale C. purpurea.
C. Berkeieyi Mont.
Syll. Crypt. 1856, 180.
6-8 cm. high, pallid then yellowish, tips red, drying black: stem rooting: branches
numerous, fastigiate, as thick as a crow's quill, repeatedly trichotomous, ultimate
branchlets capitate-fasciculate: fragile.
On trunks in wet places: Ohio, Columbia.
Coker suggests this may be Ramaria stricta.
C. Cladoniae Kalchbr.
Grev. 10, 1882, 105.
2·5 cm. high, solitary, ferruginous ochraceous, divided from the base into short
flattened branches, subtuberculate in parts: mycelium white, rooting.
On the ground: S. Africa (Somerset-East).
Said to resemble somewhat the podetia of Cladonia pyxidata. ? Lachnocladium.
C. compressa Schroet.
Pilz. Schles. 1888, 447 (non B., non Schw.).
Synonym: C. Schroeteri P. Henn., Sacc., Sy!!. 14, 1891, 238.
-2·5 cm. high, yellow, the tips becoming brown, fleshy firm: stem I cm. high:
branches few, 2-4-chotomous, compressed, tips more or less flattened, rounded, or
obtuse: (sp. ? white).
In the palm hot-house of Breslau Botanic Garden: Germany.
Probably unidentifiable, as an introduced species. Lachnocladjum or Clavulinopsis?
The name is a later homonym.
C. decolor B. et C.
Proc. Am. Ac. Arts Sci. 4, 1858, 124: Burt, Ann. Mo. Bot. Gdn. 7, 1922, 70, t. 11,
f. 109.
White, soon turning brown or black, branched: stem cylindric, arising from
branched mycelial fibrils: branches short, subdichotomous.
On hillsides: Hongkong.
'Allied to C. abietina' (B. et C.). Burt found white, globose, minutely rough spores,
CLAVARIA
3-4 x 3 µ, on the dried specimen, but was uncertain if they belonged to the fungus.
There is no specimen in the Kew herbarium. Possibly Scytinopogon.
C. fimbriata Quel.
Ass. Fr. 21, 1887, 589, t. 21, f. 12: (non Pers. 1794, nomen invalidum = Clavulina
cristata, teste Coker).
-2 cm. high, tenacious, white then pale ochre flesh colour, slender: stem surmounted
by filiform branches flattened and palmately divided into digitate or fimbriate tips:
spores 8-9 µ, pruniform-oblong, white, punctulate: caespitose on humus in mountain
pine-woods: France.
This is like a slender form of Clavulina cristata, according to Quelet, but with a
different spore. It may be Lentaria Micheneri or L. Patouillardii.
C. fuscescens Fr.
R. Soc. Upps. Acta 3, 1, 1851, 116.
3·5 cm. high, very much and repeatedly branched from a slender base, glabrous
(pallid, fuscous, soft and flaccid: dried): branches multifid, filiform, subulate.
On rotten trunks: Mexico (near Mirador).
Little known and probably unrecognizable. Compare Pterula angustata.
C. Gollani P. Henn.
Hedw. 1900, 151.
-30 cm. high, 5--7 mm. wide, apex c. 1 cm. wide, gregarious, simple, cylindric, pery
long, strict, hollow, pallid, obtuse, base slightly attenuate, subdiscoid.
On the ground: India (Saharanpur).
Spores 4-4·5 X 3·5 µ, white or slightly fuscous, subovoid.
This is said to be near Clavariadelphus fistulosus, but the spores (doubtfully correct)
are very different. It may be an exceptionally large state of Clavulinopsis spiralis.
C. Jacquemontii Lev.
Champ. Ex. 214: Sacc., Syll. 6, 1888, 698.
6-8 cm. high, white, caespitose: stem very short, much branched: branches crowded,
dichotomous, fastigiate, compressed.
On the ground: Kashmir.
Perhaps Scytinopogon or Clavulina.
C. Lecomtei Pat.
Joum. de Bot. 1895, 366.
8-15 mm. X 0·3 mm., simple, filiform, setaceous, acute, ansmg from a tubercle,
scattered or subgregarious, glabrous, rufous, or yellowish, arising from a very thin
crustaceous membranous mycelium, whitish fuscescent, with fibrillose strands, widely
effused, stratified.
On bark: Congo.
The description refers to dried specimens. It suggests Lentaria surculus, Pterula,
or the Tremellaceous Pterula sclerodontia (p. 536).
C. nigricans Lloyd
Myc. Notes 51, 1917, 729, f. 1093, 1094.
1-1·5 cm. high, simple, erect, black: stem 5-7 X 2 mm., distinct: club ovate-clavate,
rugulose-plicate.
On the ground: Chile.
Spores 10 X 6 µ, white, smooth, guttulate.
Superficially like a Geoglossum, but with basidia according to ·Lloyd. It may be
allied with C. cinereo-atra.
264 CLAVARIA
C. plebeia Fr.
Pl. Preiss. 2, 1847, 137.
Tough, white, becoming yellowish, trunk rather thick, branches and branchlets
very much divided, multifid and cristate at the tips: spores white: in sandy soil:
Australia.
Perhaps Scytinopogon.
C. polita Fr.
R. Soc. Sci. Upps. Act. 3, 1, 1851, u6.
Branched from the base, glabrous, white (rigid-fragile when dry): branches few,
hollow, little divided, unequal, acute: drying polished and shiny.
? terrestrial: Mexico (Zuacapa).
Probably unrecognizable, unless there is a type-specimen.
C. portentosa B. et Br.
Tr. Linn. Soc. Ser. 2, 2, 1883, 65, t. 14, f. 15.
u-12 cm. high, whitish, repeatedly much branched: stem 6-7 x 2·5 cm., sub
cylindrit:, rough: branches with elongate, subulate tips: spores?: among leaves,
Queensland.
Perhaps Scytinopogon.
C. regularis Herpell
Hedw. 52, 1912, 392.
2-3 cm. high, -6 mm. wide, simple, caespitose, clavate, shortly acuminate, tapered
into a stem, greyish yellow, tomentose at the base.
In woods: Germany.
Spores 18-31x11-17 µ,, white, smooth, ovoid, inequilateral.
Such large spores hardly seem possible. Compare Clavulinopsis rivalis and Aphelaria
tuberosa.
C. rhodochroa Mont.
Canar. 77, t. 4, f. 3: Sacc., Syll. Fung. 6, 1888, 714.
Rose-pink, tips blackening, fleshy: stem thick': branches terete-compressed,
spreading or deflexed, subdichotomous.
On roots of trees in woods: Canary Isl. (leg. Despreaux). ? a Pyrenomycete.
C. Schweinfurthiana P. Henn.
Hedw. 1898, 284.
3-4 cm. high, 2 cm. wide, simple, clavate, broadly compressed, smooth, pale cinnabar,
obtuse or subacute, attenuate into a cylindric stem.
On the ground under trees: central Africa.
Spores 6-8 X 3·5-4µ,, white, smooth.
? Clavulinopsis.
C. seminicola P. Henn.
Hedw. 1897, 195.
5-11 mm. high, fasciculate, simple and acute or palmatifid at the apex, subspathulate
or subulate-compressed, rugulose (? dried).
On seeds of some Euphorbiaceous plant: Brazil (Blumenau).
Spores 3·5-4·5µ,, white, smooth, subglobose.
Perhaps Pteruloid, if it is a Basidiomycete. ? Xylaria.
C. sibutiana Har. et Pat.
Bull. Mus. Hist. Nat. 1909, 89.
1-2 cm. high, scarcely 1 mm. wide, simple, wholly somewhat rufous, oblong, attenuate
at the apex, glabrous, often channelled: stem 2 x o·5 mm., glabrous, cylindric.
CLAVARIA 265
On bare, earth: tropical Africa (Fort Sibut, Congo). 'C. falcatae affinis sed colore
diversa.'
This may be C. helicoides, but is probably unrecognizable, save from the type
specimen.
C. subfistulosa P. Henn.
Engl. Jahrb. 25, 1898, 498.
9-11 cm. X 1 mm., gregarious, simple, filiform, equal, strict, fistulose, subobtuse,
pale rufescent.
On the ground in the forest: New Guinea (Astrolabe Plain, Kaiser Wilhelmsland).
Spores 4-4 · 5 X 3-3· 5 µ,, brown, smooth, ellipsoid or subglobose.
Basidia 4-spored.
If really brown-spored, it is the only simple Ramaria, but Hennings's spore-data
are not always correct.
C. tonkinensis Pat.
3-4 cm. X 2-3 mm., solitary, simple, straight, cylindric, obtuse, base subinflated,
glabrous, pellucid, fleshy, wholly pallid brown (? dried}.
On the ground: Tonkin.
Perhaps the same as C. helicoides.
EXCLUDED SPECIES
C. Bourdotii Bres.
Ann. Myc. 6, 1908, 45.
C. Bresadolae Quel.
FI. Myc. 1888, 458: Bres., Fungi Trid. 2, 40, t. 146, f. 2: Bres., le. Myc., t. 1105:
(non C. Bresadolae Cav. = Ramariopsis pulchella): (non sensu Bres., Ann. Myc. 1,
1903, 90 = Protohydnum fasciculare).
C. himantia (Schw.) Fr.
Epicr. 1838: Bourd. et Galz., Hym. Fr. 1928, 123, f. 50: Lundell et Nannfeldt, Sv.
Svamp. 1-2, 1934, 17.
Hydnum himantia Schw., Sehr. Naturf. Ges. Leipz. 1, 1822, 104.
Hydnum subjuscum Pk., N.Y. St. Mus. Dept. 40, 1887, 55.
Coniophorafiavomarginata Burt, Ann. Mo. Bot. Gdn. 13, 1926, 311.
Odontia himantia (Schw.) Bres., Ann. Myc. 1, 1903, 85.
Oxydontia himantia (Schw.) Miller, Mycol. 25, 1933, 363.
Mycoacia himantia (Schw.) Miller et Boyle, Univ. Iowa St. Nat. Hist. 18, 1943, 44:
Rogers et Jackson, Farlowia 1, 1943, 278.
These three species are Hydnoid fungi, arising from a more or less extensive
subiculum in the manner of Odontioid spines. Presumably the spines are positively
geotropic, but this point has not been mentioned in descriptions. C. himantia has
narrow, monomitic, uninflated, clamped hyphae 2-5 µ, wide, long-celled, with slightly
thickened walls (o· 5 µ) in the subiculum, and, apparently, subgelatinous in the spines.
The generic position of the three species is not yet satisfactorily determined.
The earliest specific epithet for C. himantia is C. byssiseda Fr. (Syst. Myc. 1, 1821:
Pers., Myc. Eur. 1, 1822). This has been misinterpreted very generally for the true
Clavarioid fungus Lentaria byssiseda, but the original descriptions of Fries and
Persoon leave no doubt that by C. byssiseda they intended what has come to be known,
from N. America, as C. himantia.
266 CLAVARIA
C. coriaria Pers.
Myc. Eur. r, 1822, 171.
'Alba, fragilis caespitosa ramosa, ramis erectis elongatis crassitie aequalibus sub
acutis. Rebent. Flor. Neomarch. p. 378. Cl. subcaulescens.'
'In pulvere coriariorum, magnos cespites format. Altitudine unciali ad rt. Reb.'
'Obs. Ob crescendi locum ulterius inquiri meretur, ita ut descriptio, nimis brevis,
novis suppleatur observationibus.'
I cannot find that this has been identified.
C. ? dubiosa Speg., Fung. Arg. Pug. 3, n. 41: Sacc., Syll. 6, 1888, 714.
3-4 x 0·1-0·2 mm., erumpent through cracks in bark, consisting of tufts of 3-5
simple branches, more or less united at the base, the central one erect, the others
spreading, white or pinkish violaceous, rather fleshy-tough but slender: spores not
seen: on the bark of fallen, rotting trunks of Salix Humboldtiana: Argentina.
Perhaps the conidial state of a Pyrenomycete (Hypocreales). Compare Lachno
cladium aurantiacum and Pistillaria fueggiana.
C. fusco-lilacina B.
Hook. Journ. Bot. 8, 1856, 276: Petch, Tr. Br. Myc. Soc. 15, 1930, 247.
'A dense cluster of stout clavae (or branches) which appear to arise from a common
base but it is possible that they may be merely united at ground level by mycelium.
The branches divide once or twice from about half their height and together form a
l�ose coralloid fructification. The colour is now pale purple brown.' (Petch).
This type-specimen in the Kew herbarium has no trace of hymenium, the surface
being covered by a thin, fuscous, agglutinated crust of hyphal ends, c. 100 µ, thick.
The hyphae appear uninflated, but are much collapsed. It seems to be a sterile
Pyrenomycete.
Van Overeem's interpretation of the species should be referred to Clavulina
Leveillei.
C. ignea Pers.
Myc. Eur. r, 1822, 174.
'Cespitosa flavo-rubra, deorsum ventricosa, ramulis paucis tenuibus brevibus.
Mick. Gen. pl. p. 209, t. 88, f. 2.'
'Hab. in Montis-Boni ericetis et sylvis caeduis, October. Mick.'
I cannot find that this species has been identified.
C. nigra Lev.
Ann. Sci. Nat. 3, 2, 1844, 215.
Caespitose-pulvinate, 2 cm. high, 2-3 cm. thick, tuberculose, black: pedieels sub
fasciculate, dichotomous: branches short, obtuse, umbellate: spores?: Sumatra.
Van Overeem suggests this is a Xylaria, but the microscopic structure of the co-type
fragment at Kew seems parenchymatous ! It is certainly no Clavaria and doubtfully
a fungus.
C. Ridleyi Mass.
Kew Bull. 1899, 172.
Branched, umber: stem I cm. wide, short, thick, solid: branches somewhat crowded,
terete, smooth, dichotomous, lunulate-furcate at the apex, umber-rufescent: flesh
white.
On the ground at the base of trunks: Malaya (Perak, Ridley 19).
The type in the Kew herbarium suggests a branched Pyrenomycete. The surface
has a palisade of short, clavate, sterile cells with brown walls. The spores and basidia
attributed by Massee are erroneous.
CLAVARIA
C. sparassoides Speg.
Fung. Guar. Pug. 1, 1883, n. 86.
Synonym: Thelephora sparassoides Speg. (? ined.).
Fawn-white: stem 5 X 3 mm., arising from a small tubercle, with 4-9 branches at
the apex: branches spathulate-dilated or foliaceous, to 2 X 1 cm., undulate, plicate,
ending in flattened erect laciniae: on rotten wood in the forest: S. America (near
Guarapi).
I have not seen the type-collection, but in the Kew herbarium there is a later collec
tion (Thaxter 7009, from the Argentine) identified as Thelephora sparassoides Speg.
by Spegazzini. Macroscopically it answers to the description of C. sparassoides, but it
is an incipient polypore. The thin, foliaceous, coriaceous pilei appear to have been
7-8 cm. long, tufted from a common base, laciniate distally into narrowly spathulate
or ligulate lobes, with thin, uneven or incised margin. Their uppersides are inoderm
and sterile and marked with narrow, darker, fawn-ochraceous zones 3-6 mm. apart:
on the undersides, towards the margin, there are very shallow irregular pores,
c. 0·2-0·5 x 0·1-0·2 mm., or an interrupted network, as if incipiently irpicoid. The
flesh is very thin in the dried specimen, possibly o· 5 mm. thick or more in the living
state, and is composed of radial, long-celled, uninflated, monomitic hyphae, 3-8 µ,
wide, without clamps and with the walls 0·5-2µ thick, not darkening with dilute
alkali. The hymenium on both the smooth and poroid parts of the underside of the
pilei does not seem to thicken: the subhymenium is narrow and loose: the basidia are
4-spored: there are neither cystidia nor gloeocystidia. The spores are very abundant,
5-6 x 3-3·7 µ,, white, smooth, ellipsoid, blunt, thin-walled, and finely multiguttulate
(dried). Thus from this collection of Thaxter's, the species has the hyphal construc
tion and spores of Aphelaria, but the dorsiventral merismoid pilei and rudimentary
pores of some Polyporoid genus: macroscopically it recalls sterile poreless specimens
of Polystictus Gaudichaudii, which is, however, trimitic.
Compare also C. guarapiensis and C. Spegazzinii mentioned under Scytinopogon
angulisporus: they may be young states of C. sparassoides or of Scytinopogon.
C. verpiformis B. et Cke.
Journ. Linn. Soc. Bot. 15, 1876, 391.
Text-Fig. 99.
-4 mm. high, white: stem 1-2 mm. wide, 0·5-1 mm. high, VeY)' short and thick: head
rugulose-subcerebriform, solid, wholly covered by the hyme
nium: flesh drying friable with a firmer white core, not woody
(? subgelatinous when fresh), solid.
On bark and wood(? erumpent): Brazil (Rio Jurua: Traill
180).
Hyphae 2-6 µ, wide, monomitic, clamped, with thin, sub
gelatinous walls, longitudinal. ThxT-FIG. 99.
Hymenium covered with a mucilaginous layer (more or Clavaria verpiformis:
less disorganized); no basidia seen but many subclavate and dried specimen from the
type-collection, x 2.
often irregularly branched or lobed cells in a loose palisade.
The type-specimens in the Kew herbarium are few, rather battered and broken,
and all detached from the wood so as to leave white disk-like scars where they had
grown. The affinity is probably with Caripia, unless it is a heterobasidiomycete.
I could find no spores.
268 CLAVARIACHAETE
CLAVARIACHAETE gen. nov.
Fruit-body much branched, erect, clavarioid, dark ferruginous when dried, stei:n short,
subtomentose, sterile: (branching apparently flattened, but with little dilation of the
axils): branches rather slender, terete, attenuate, straight, becoming repeatedly dicho
tomous in different planes: hymenium on all sides of the branches, finely brown setose:
flesh tough, coriaceous (? rather dry).
Lignicolous (? also humicolous): tropical America: 2 spp.
Spores and basidia ?
Setae simple, conical, straight, with thick, dark brown walls, smooth, arising deeply
in the thick subhymenium and projecting far.
Gloeocystidia none: dichophyses none.
Hyphae monomitic, not inflating, without clamps, with more or less thickened, pale
yellow-brown walls, not encrusted.
Hymenium not thickening.
TEXT-FIG. 100. Clavariachaete rubiginosum: diagrams

of the structure of the stem with carbonaceous
cylinder and sterile cortex (on the left), and of a
fertile branch (on the right) with hymenium, setae,
and cylinder of thick-walled medullary hyphae: X 25.
Among Clavarioid fungi this genus comes next to Lachnocladium. It differs in the
presence of true setae, in the absence of gloeocystidia and dichophyses, and in the
amphigenous, not unilateral, hymenium, which is continuous from the first.
It seems that only two collections of dried specimens are known-the type of each
species. The colour of the fresh fruit-bodies is probably tawny or light ferruginous
ochraceous, much as in the more richly coloured species of Lachnocladium. Lloyd
very doubtfully ascribed 8 µ,, subglobose, brown spores to C. Peckoltii: I could find
none on the type of C. rubiginosum. The basidia are probably as small as in Lachno
cladium. However, the genus cannot be fully described until fresh specimens have
been studied.
The generic name is based on 'Clavariachaeta McGinty' which Lloyd facetiously
proposed for his Dendrocladium Peckoltii. He emphasized the remarkable 'Hymeno
chaete-setae' as unique among Clavarioid fungi. One can hardly take such whimsi
cality, however, as a valid generic description or proposal.
CLAV ARIACHAETE
KEY TO THE SPECIES
Stem short, c. 1 cm. long or less: branches terete, the axils scarcely
dilated: lignicolous: Venezuela C. rubiginosum
Stem elongate: branches flattened and dilated at the divisions,
antler-like: Brazil C. Peckoltii
C. Peckoltii (Lloyd) comb. nov.
Basinym: Dendrocladium Peckoltii Lloyd, Myc. Notes 66, 1922, 1u1, f. 2068.
-8 cm. high, ferruginous brown (dried): stems 2-4 cm. high, caespitose: branches
palmate, antler-like (? polychotomous below), strongly flattened and dilated at the
divisions, pubescent with dark-brown setae.
Habitat?: Brazil.
[Spores 8 µ. wide, subglobose, deep-coloured, 'but they do not appear to be ordinary
basidial spores for one seen had a long, attached pedicel which is evidently the hypha
on which it grew': Lloyd.]
Setae projecting 60-80 µ., large, intense brown 'typically the setae of Hymenochaete'.
Whether this has the same microscopic structure of the flesh as in C. rubiginosum
remains to be seen.
C. rubiginosum (Cke.) comb. nov.
Basinym: Lachnocladium rubiginosum Cke., Grev. 20, 1891, II.
Text-Figs. 100, 101.
-8 cm. high, rubiginous (dried), very much branched, fastigiate: stem short, sub
tomentose: branches 1-1·5 mm. wide below, tapering to 0·5 mm., terete, repeatedly
dichotomous, dense, with subulate tips, wholly set with dark brown setae, scarcely
dilated at the axils: rigid, coriaceous.
On trunks: Venezuela.
Spores and basidia?
Setae 100-250 X 12-18 µ., projecting -120 µ., very abundant, lanceolate, straight,
with very thick, dark brown, smooth walls.
Hyphae 3-6 µ. wide, long-celled in the medulla and sterile cortex of the trunk,
shorter-celled in the subhymenium.
Medulla 300-400 µ. wide in the trunk, tapering·distally, defined in transverse section
by a thin black carbonaceous circle in the lower part of the fruit-body, the inner
hyphae with scarcely thickened walls, the peripheral with strongly thickened walls.
Subhymenium 100-170µ. thick, the hyphae with distinctly thickened walls, thinner
and paler towards the hymenium, rather loose, branched at a Ytide angle and often
pseudodichotomous.
I have studied the type in the Kew herbarium.
CLAVARIADELPHUS Donk.
Rev. Niederl. Homobas. Aphyll. 2, 1933, 72.
Fruit-bodies typically solitary, occasionally subcaespitose, filiform to clavate, ligulate
or turbinate, in some species truncate and sterile at the expanded apex (even perforate),
without marginal growth, simple (exceptionally with one or a few short branches),
massive to filiform, light yellow or light ochraceous at first, in some species tinged pink
or violet, becoming deep ochraceous, brown, rufescent or ferruginous (persistently pale in
C. contortus; light orange-yellow in C. maricola): flesh rather firm, not brittle, spongy
in large fruit-bodies: hymenium thickening, becoming longitudinally rugulose in large
fruit-bodies, covering the greater part of the fruit-body except the extreme base and,
in a few species, the sterile apex.
Terrestrial in humus of coniferous or frondose trees, or on twigs, leaves, and cones,
not lignicolous on trunks: 9 spp., north temperate (1 in Indo-China).
T'ExT-FIG. 101. Clavariachaete rubiginosum: the hymenium in longitudinal section,
showing the single layer of basidia, the thick subhymenium witb deep-seated setae,
the thick-walled outer medullary hyphae, and the thin-walled inner medullary
hyphae, x 500 (from the type-specimen).
CLAVARIADELPHUS
Spores 9-22 X 4· 5-1o µ, white or pale yellowish, smooth, thin-walled, elongate ellip
soid, pip-shaped or almond-shaped to subcylindric, contents finely granular guttulate,
or merely cloudy vacuolate.
Basidia with 4 straight or slightly curved sterigmata, cytoplasm multiguttulate.
Cystidia none.
Hyphae monomitic, with clamp-connexions, inflating, thin-walled, not secondarily
septate: some species with scattered laticiferous hyphae.
Type-species: C. pistillaris (Fr.) Donk
Subgenus Eu-Clavariadelphus
Fruit-bodies massive, fertile at the apex, clavate or ligulate, becoming spongy:
hyphae much interwoven.
C. pistillaris, C. ligula, C. sachalinensis, C. maricolus, (? C. mirus).
Subgenus Cantharellopsis
Fruit-bodies massive, turbinate with sterile truncate or subpileate apex, becoming
spongy: hyphae much interwoven.
C. truncatus, C. unicolor, (? C. mirus).
Subgenus Typhulopsis
Fruit-bodies slender, fertile at the apex, rigid, not spongy: hyphae strictly longi
tudinal, agglutinated on the surface of the stem: humicolous or on twigs and leaves.
C. fistulosus, C. junceus.
It is likely that several species will be added to this genus when the fungus flora of
the tropical mountain regions is better known. Even the N. American fungi referred
to C. pistillaris, C. ligula, and C. truncatus may need specific rank. The genus has not
been reported certainly from the tropics, and from the southern hemisphere only as
C. junceus.
The spores of several species become pale yellowish on drying: possibly the fresh
spore-print is pale yellow or ochraceous, as in species of Cantharellus, but more precise
information is required.
KEY TO THE SPECIES OF CLAVARIADELPHUS

Fruit-bodies more or less turbinate or subpileate with a sterile
truncate apex, large: sp. 9-14 X 4-8 fL
Yellow or orange ochraceous, then rufescent or cinnamon,
6-15 X 2-9 cm.: on coniferous needles: N. temp. C. truncatus
Chocolate to warm blackish brown: China var. atrobrunneus
Flesh colour tinged violet, then ochraceous buff, often perforate
at the apex and subpileate, 3-5 X 1·5-5 cm.: on the ground in
mixed woods: U.S.A. C. unicolor
(Pale brown, 8-15 X 0·6-1·1 cm.: on the ground in mixed forest:
Indo-China C. mirus)
Fruit-bodies with fertile, round or pointed apex (truncate in C.
mirus from lndo-China)
Fruit-bodies very large, 7-30 X 2-6 cm., on the ground in fron
dose woods
Sp. 11-16 x 6-10 fL: fruit-bodies yellow then ochraceous or
tinged rufescent or brownish, the flesh becoming brownish
vinaceous on bruising: Europe C. pisttllaris
Sp. 7-12·5 X 3·7-7·5 µ.: fruit-bodies flesh colour or rose-pink,
then ochraceous: ? turning brownish vinaceous on bruising:
U.S.A. var. amerzcanus
CLAVARIADELPHUS
Fruit-bodies smaller
Fruit-bodies 0·6-1·6 cm. wide
With obtuse, truncate, or mamillate apex: fruit-body
8-15X6-1·1 cm., cylindric, pale brown: sp. 8-10x
4-5 µ: on the ground in mixed forest: lndo-China C. mirus
Not so: more or less ochraceous then brownish
Fruit-body 10-30X0·2-1·3 cm. filiform to narrowly
clavate: sp. 10-18·5X4·5-9µ: on dead sticks, frondose
and coniferous: N. temp. C.fistulosus
Fruit-body 3-1oxo·5-1·5 cm., more or less ligulate or
spathulate: on coniferous needles
Light ochraceous, then tinged rufescent: sp. 8-15X
3-6µ: Europe C. ligula
Dark ochraceous: sp. 17-22X5-5·5 µ: temp. N. Amer.
and Japan . C. sachalinensis
Fruit-bodies o·6 cm. wide or less
Fruit-bodies 0·5-3x0·2-0·6 cm., cylindric, clavate, short,
thick, blunt, often deformed, pale ochraceous: on dead
frondose sticks: sp. 14-23X7-9µ: N. temp. C.fistulosus var.
contortus'
Fruit-bodies longer, more slender
10-30X0·2-1·3 cm., filiform to narrowly clavate, ochra
ceous to rufescent or dark brown: sp. 10-18·5X4·5-9µ:
on dead sticks, coniferous and frondose: N. temp. C.fistulosus
7-10x0·2-0·6 cm., light orange-yellow to orange-buff,
often subligulate: sp. 10-14X4-5µ: ? very long basidia:
on needles and cones of Western Yellow Pine: U.S.A. C. maricolus
3-15x0·05-0·2 cm., filiform or subclavate, slender, pale
ochraceous to brownish: sp. 6-12X3·5-5µ: basidia
short: on humus and sticks in frondose woods and on
the ground: N. temp. C.junceus
C. fistulosus (Fr.) comb. nov.

Basinym: Clavariafistulosa Fr., Syst. Myc. 1, 1821, 479: v. Hoehne!, Oesterr. Bot.
Zeitschr. 1904, 425: Lind, J., Ann. Myc. 5, 1907, 272: Harper, Mycol. 10, 1918, 53:
Bourd. et Galz., Hym. Fr. 1928, 120, f. 49: Konr. et Maubl., le. Sel. Fung., t. 496:
Konr., Bull. Soc. Myc. Fr. 41, 1925, t. 8.
Synonyms: Clavaria ardenia Fr., Syst. Myc. 1, 1821, 478: Bres., Ann. Myc. 1,
1903, 113: Harper, Mycol. 10, 1918, 53, t. 3.
C. tuberosa Fr., Syst. Myc. 1, 1821, 486.
C. pilosa Pers., Myc. Eur. 1, 1822, 177.
C. strigosa Pers., Myc. Eur. 1, 1822, 177.
C. alnea Schulz., in Kalchbr., le. Hym. Hung. 4, 1877, t. 35, f. 7 (? = var. contortus).
Calocera tuberosa Fr., Hym. Eur. 1874, 680.
Clavariellafistulosa (Fr.) Karst., Rev. Myc. 3, 1881, 21.
10-30 cm.x2-8 mm., -13 mm. in robust specimens ( = C. ardenia), simple, fili
form, cylindric and acute, becoming narrowly subclavate and obtuse, even subtruncate
at first: filiform, always slender, becoming hollow, yellow then tawny, rufescent or ferru
ginous or finally sometimes date-brown, often contorted below, the base often villous
1
Compare also P. alnicola, P. paradoxa, &c., in PistiUaria (Section C of the Key, p. 474).
CLAVARIADELPHUS 273
and rooting or decumbent: flesh yellowish, firm, rather rigid: smell and taste none:
hymenium covering the upper two-thirds of the fruit-body.
Solitary or gregarious, never caespitose, superficial on dead sticks and branches of
frondose and coniferous trees: Europe (uncommon), Japan (Prov. Kushiro), Canada,
U.S.A. (northern states).
Spores 10-18·5 X 4·5-9 f-L (av. 14·5 X 6·7), white, smooth, thin-walled, narrowly
ellipsoid or subfusiform, the apex subacute, with finely granular-guttulate contents.
Basidia 40-95 X 8-u·5 µ,, with 4 sterigmata 10-12 x 3 µ,: cystidia none.
Hyphae 1·5-15 µ, wide, with clamp-connexions, with scattered, frequently branched,
laticiferous hyphae 3-9 f-L wide, ending in the hymenium with round tips like gloeo
cystidia.
forma rnacrorrhizus Bourd. et Galz.
Hym. Fr. 1928, 122.
Synonym: Clavaria macrorrhiza Fr., Syst. Myc. 1, 1821, 478.
Cylindric, obtuse, yellow-brown, with long tomentose-fibrillose root.
Europe.
TEXT-FIG. 102. Clavariadelphus fistulosus var. contortus: fruit-body, X I;

spores, X 500; basidia, x 1,000.
var. contortus var. nov.
Synonyms: Clavaria contorta Fr., Syst. Myc. 1, 1821, 478: Li!:.J, Ann. Myc. 5,
1907, 272: Boud., Bull. Soc. Myc. Fr. 33, 1917, t. 1.,.f. 5: Harper, Mycol. 10, 1918, 53:
Coker, Joum. El. Mitch. Sci. Soc. 63, 1947, 49, pl. 5.
C. brachiata Fr., Syst. Myc. 1, 1821, 478.
? C. bulbosa Pers., Myc. Eur. 1, 1822, 176.
C. fistulosa forma brachiata Bourd. et Galz., Hym. Fr. 1928, 123.
C. fistulosa forma contorta Donk, Rev. Nieder!. Homobas. Aphyll. 2, 1933, 92.
Aleurodiscus helveolus Bres., Mycol. 17, 1925, 21: (fide Rogers et Jackson, Farlowia
I, 1943, 269).
Text-Fig. 102.
0·5-3 cm. x 2-6 mm., generally erumpent, solitary or fasciculate, simple or occa
sionally with 1-2 short branches or deformed and tubercular, cylindric to spathulate,
rather short and thick, generally obtuse, sometimes twisted, becoming wrinkled, watery
yellowish or pale yellowish drab, then ochraceous bistre, often tinged brown: flesh
yellowish, floccose, loose, rather tough, becoming hollow: smell and taste none:
entirely covered with the hymenium.
On dead twigs, especially of Alnus, lying on the ground or on the trees: Europe,
temperate N. America.
5119 T
274 CLAVARIADELPHUS
Spores 14-23 X 7-9µ (av. 18 x 8), white, smooth, narrowly ellipsoid and flattened
on one side, almond-shaped or ovoid, with finely granular-guttulate contents.
Basidia 45-80 X 6-10µ, granular-guttulate, with 4 sterigmata 8-14µ long.
Hyphae 2-20µ wide, widest in the central tissue, narrower to the subhymenial
hyphae 2-5 µ wide, with scattered laticiferous hyphae as in C. fistulosa.
It is now generally agreed that C.fistulosa and C. contorta are forms or states of one
species, C. fistulosa being the normal condition on sticks in humus, with f. macrorrhizus
as a 'deeply rooting' form, and C. contorta being an abnormal condition developed on
TABLE XVIII. SPORE-SIZE IN CLAVARIADELPHUS FISTULOSUS
Size in µ Country Author

14·4-17·5X 6-7 Denmark Lind
12-15X6--'] France Bourdot and Galzin
12-18X 5-8 " Konrad and Maublanc
10-17X 7-9 Gt. Britain Cotton and Wakefield
15X 6 Rea (as C. ardenia)
12X7 " Rea
13-18·5X4·5-7·5 Holland Donk
12-15X5-? Poland Bresadola (as C. ardenia)
10-15X5-8 Japan Imai
14-18X6-9 U.S.A. Harper
12-16X5--'7 " Harper (as C. ardenia)
12-15 X5·5--'7·5 Europe Coker (herb. Bres.)
15-18x6·5-8
18·5X8
10-18·5x4·5-9, av. = 14·6x6·8
var. contortus
17-2ox8-9 Denmark Lind
12-18X6-9 France Bourdot and Galzin
17-23 X8-10 Gt. Britain Cotton and Wakefield
12-14X9 Rea
17-20X7-9 " Comer
14-2ox7-9 Europe Coker (herb. Bres.)
14-18X6-9 U.S.A. Harper
12-23X 6-9, av. 17·5 X8·1 (excluding Rea's data)
sticks in exposed places. Thus C. contorta has a shorter, stemless, and often deformed,
fruit-body of paler colour, and erumpent habit, corresponding with stunting through
exposure. But C. contorta has, also, larger spores than C. fistulosa as shown in
Table XVIII (av. 17·5x8·rµ, compared with 14·6x6·8µ). Further, there is yet no
experimental proof that the mycelium of C. contorta will produce fruit-bodies of
C. fistulosa under the supposedly appropriate conditions, or vice versa. I have, there
fore, retained C. contorta as a variety of Clavariadelphus fistulosus. Neither is really
common enough for close study in the field, and the descriptions of those who unite
them are insufficiently critical of the differences emphasized by those who maintain
them as separate species.
In Denmark, Lind draws these distinctions:
Spores: 17-20 X 8-9µ in C. contorta, 14·5-17·5 X 6-7µ in C. fistulosa.
Habitat: fruit-bodies of C. contorta developed on Alnus incana Oct.-Nov., often on
twigs still attached to their branches at heights above 2 in. from the ground::
those of C. fistulosa developed towards the end of December, only on fallen twigs.
of Fagus silvatica hidden beneath the leaf-mould.
CLAVARIADELPHUS 275
These observations certainly do not hold in other countries. C. contorta occurs on
twigs of Fagus silvatica in England and C. fistulosa in the United States occurs on
coniferous twigs, according to Harper (unless his specimens are referable to C.
maricolus): both have been reported from Betula, Corylus, Populus, Ulmus, Acer,
Quercus, and Castanea in other parts of Europe. Yet, locally, there may well be fairly
constant differences in habitat, with which size of spore, shape of fruit-body, and so
on may be correlated.
C. ardenia is a robust form often becoming truncate or subcyathiform and,
eventually, hollow. It is rare and little studied, but it may be separable as a variety
tending towards C. pistillaris.
The only record of either species from the south temperate region is Massee's for
New Zealand (Tr. N. Zeal. Inst. 39, 1906, 40). As the spores are given as 4-5µ, sub
globose, confirmation is needed. Compare Pistillaria paradoxa.
C. junceus (Fr.) comb. nov.

Basinym: Clavariajuncea Fr., Syst. Myc. 1, 1821, 479: Harper, Mycol. 10, 1918,
56, t. 5: Boud., le., t. 176: Bres., le. Myc., t. 1104.
Synonyms: Typhula juncea (Fr.) Karst., Bidr. Finl. Nat. o. Folk, 37, 1882, 181:
(Schroet., Coker, Donk, Clel.): forma vivipara Fr., ibid.
Typhula filiformis Fr., Syst. Myc. I, 1821.
Clavaria filiformis (Fr.) Karst., ibid. 180.
Clavaria tortilis Pers., Myc. Eur. 1, 1822, 187.
Typhula ramentacea Fr., Syst. Myc. 1, 1821.
Clavaria ramentacea (Fr.) Karst., ibid. 180.
Clavaria hirta Pers., Myc. Eur. 1, 1822, 186.
Clavaria virgultorum Pers., Myc. Eur. 1, 1822, 186.
Clavaria Klotzschii Lasch, in Klotzsch Herb. Viv. Suppl. 240 (cum diagn. 'querci
cola, typhuloides, gracillima').
Text-Fig. 103.
3-15 cm. high, 0·5-2 mm. wide, solitary or gregarious, filiform-acute, then often
narrowly fusiform or subclavate, acute then obtuse, rather stiff and rigid, flaccid with
age, pale brownish ochraceous, becoming pallid ochraceous or brownish drab or somewhat
rufescent: stem distinct, 1-5 cm. long, slightly narrower than the fertile club, attached
by appressed fibrils at the base, sometimes with abortive branches above, appearing
smooth: flesh rather firm and rigid, not brittle,juicy, becoming hollow in old specimens:
smell rank, sour, taste similar (? subacrid).
Among dead leaves and twigs, generally attached to small rotten twigs and petioles
in frondose woods, rarely on coniferous needles (U.S.A.), rarely on bare soil (Eng
land): Europe, N. Amer., China, Japan, N. Africa (Tunisia), S. Austral., Tasmania
(? New Zealand).
Spores 6-12 X 3·5-5·5(-6)µ, white, smooth, more or less amygdaliform, thin-walled,
aggutate.
Basidia 30-55 x 7-10µ, clavate, with a narrow tapered base 2·5-3µ wide: sterig
mata 4, 8-11µ long.
Hymenium thickening to 90µ (? more), with very many small, often brownish,
amorphous or subcrystalline granules and a few larger masses, firmly subgelatinous:
subhymenium c. 10µ thick at first, becoming 20-35µ thick, composed of loosely inter
woven short-celled hyphae 2-3·5µ wide, arising abruptly at right angles from the
outer, narrow, longitudinal hyphae of the flesh: cystidia none.
Caulocystidia as narrow, simple, often flexuous, aseptate hairs 10-300µ long,
2·5-4µ wide at the base, gradually tapered to the blunt apex, the walls slightly
thickened (0·5-1µ), smooth, generally decumbent, shorter and fewer upwards: at the
base of the stem passing into the secondary mycelial hyphae, as stouter, much longer
CLAVARIADELPHUS
hairs, 4-10 µ, wide at the base, mostly 4-5 µ, wide in the middle part, often sinuous,
kinked, zigzag or lobed at the base, aseptate, unbranched, or occasionally at the base.
Hyphae 10-25 µ, wide, strictly longitudinal, compact, clamped, with broad transverse
septa, cells 100-500 µ, long and cylindric, walls firm and brownish, with a few narrow
uninflated hyphae 3-5 µ, wide, narrower to the exterior next the subhymenium and a
few with vitreous or oily-guttulate contents: on the surface of the stem 2· 5-5 µ, wide,
with slightly thickened, agglutinated walls.
Rhizomorphs 0·5 mm. wide, pallid ochraceous, finely hairy or villous, -10 cm. long,
often much branched adventitiously, commonly developed in humus and on rotting
leaves, rarely producing fertile fruit-bodies or short fertile heads, mostly sterile.
TEXT-FIG. 103. Clavariadelphus junceus: normal fruit-bodies (left), and rhizomorphs

(right): X I.
This remarkable species is very widely distributed in temperate c;ountries, where it

is not uncommon in swampy places in the autumn, but it has not been recorded from
tropical mountains. Systematically it comes exactly between C. fistulosus and Typhula
phacorrhiza and a closer link between the two genera could hardly be imagined. It is
so Typhuloid that it is now usually put in that genus. I have removed it because it
has no sclerotium, because the hymenium thickens and has the narrowly stalked
basidia of Clavariadelphus, and because it so much resembles a slender, small-spared
C. fistulosus. Karsten interpreted the slender fibrillose base and rhizomorphs as a
sclerotium; certainly they resemble the rudimentary stolons found in some species of
Typhula and the abnormal rhizomorphs developed by Typhula sclerotioides in culture.
If one imagines a rhizomorph checked at its very outset and clothed with the winding,
lobed, and distorted bases of the hairs (such as occur at the base of the fruit-body of
C. junceus), and this outer layer agglutinated, as happens t6- the superficial hyphae of
the rhizomorphs and stem, one would have a sclerotium with characteristic, if simple,
CLAVARIADELPHUS 277
epidermis just as in Typhula. However, this never happens, and C. junceus is pre
Typhuloid. It might be placed in Pistillaria, but its large size and thickening hymenium
wquld then be as exceptional as its close agreement with C. fistulosus.
The great variation in spore-size is shown in Table XIX. Possibly long-spored and
short-spored varieties may be distinguished, but discussion must be deferred until the
spore-variation in Clavarioid fungi can be examined in greater detail.
Data for the basidium are as follows (but I think most measures omit the slender
base):
18-45 X 5-8 µ,, Bourdot and Galzin,
30-5 X 7-8 µ,, Bresadola,
30-5 X 6-8 µ,, Cotton and Wakefield,
30-40 X 7-10 µ,, Corner (dried specimens),
38-55 X 8·5-10 µ,, Corner (living specim,ens),
22-38 X 6-8 µ,, Donk.
De Bary found that the hyphae (? of the fruit-body) sometimes, but not always, gave
the violet cellulose-reaction of the wall with iodine and sulphuric acid (Comp. Morph.
Fungi. Bact. 8).
TABLE XIX. SPORE-SIZE IN CLAVARIADELPHUS JUNCEUS
Spore-size in µ. Country Author

10-IZX 5 Finland Karsten
7-10X3-5 France Bourdot and Galzin
8-9X4-5 Germany Schroeter
8-11X4-5 Gt. Britain Cotton and Wakefield
8-9X4-5 " Rea
10-IZX4-5 " Corner
6·5-1ox4-5 Holland Donk
7-9X3·5-5 Italy Bresadola
(6-)7-9(-10)X3'5-4·5 Sweden Lundell
8-1zX4-5 U.S.A. Burt
6--9·3X3 ·8-4·8 " Coker
9-1ox3·8-4·5 " Doty
9-xzX4-5 " Harper
8-9X4·5 China Teng
(s-s-7X3·7-4 Australia Cleland)
6-12 X3-5, 'E r8-z·5, Maximum range
9·1X4·4, 'E z·1, I Average (excluding Cleland's figures)
' E = length -;- width.
I have reduced T. filiformis and T. ramentacea as synonyms because they seem to be

merely the rhizomorphs of C. junceus. Usually the rhizomorphs are independent of
the fruit-bodies and very rarely fertile, at the tips, themselves. Their structure is the
same as that of the fruit2body, but more slender, and at the tips the sterile .hairs form
instead of basidia. T. filiformis, as interpreted by Bourdot and Galzin, is certainly the
rhizomorph of C.junceus, and Quelet reduced T. ramentacea to T.filiformis. T. ramen
tacea was said to grow on cow-dung, but it may well have grown through or on to
cow-dung from adjacent humus. If they are not the rhizomorphs of C. junceus they
may be those of a Typhula, as described for T. sclerotioides in culture.
Forma vivipara Fr. refers merely to fruit-bodies with a tendency to produce adventi
tious branches, which may or may not become fertile, or to produce rhizomorphs from
the stalk, which they may do when growing against dead leaves in wet places: neither
condition has any systematic value.
CLAVARIADELPHUS
C. ligula (Fr.) Donk
Rev. Nieder!. Homobas. Aphyll. 2, 1933, 73.
Basinym: Clavaria ligula Fr., Syst. Myc. 1, 1821, 477: Bres., le. Myc., t. II03.
Synonyms: Clavaria luteola Pers., Myc. Eur. 1, 1821, 179.
Clavariella ligula (Fr.) Karst., Rev. Myc. 3, 1881, 21.
Text-Fig. 104.
3-10 cm. high, 0·5-1·5 cm. wide, yellowish cream then ochraceous or somewhat
rufescent, simple, rarely bifid, gradually elongato-clavate, obtuse, often flattened and
ligulate or spathulate, stem indistinct, base white villous: hymenium smooth (? slightly
rugulose): flesh white, rather spongy, floccose and loose.
On coniferous needles, solitary or in troops, attached by white mycelium, occa
sionally on humus in frondose woods: Europe, N. Africa, Japan, U.S.A., uncommon.
Spores 8-15 x 3-6 µ,, white, smooth, narrowly ellipsoid or subcylindric, the apex
blunt or subacute, with granular contents.
Basidia 40-65 x 6-9 µ,, with 4 sterigmata.
Hyphae as in C. pistillaris, with clamp-connexions.
This species is little known. I have examined a collection of young fruit-bodies
from Austria and find the structure identical with that of
f!J � {J y{) ;;;�i t�E��'.!�}:,1:.�!;.!J!��

TExT-FIG. 104.
bodies, its habitat, its slightly smaller but much narrower
spores (E = 2·65, in C. pistillaris 1·78), its smaller basidia,
Clavariadelphus ligula: and, perhaps, in the smooth hymenium and unchanging flesh,
spores, X1 ,coo. but these two points require confirmation.
As in C. pistillaris, the spores are variable, but there is clearly some confusion
between the European and N. American fungi referred to C. ligula. As shown in
Table XX, the N. American specimens have much larger and narrower spores.
TABLE XX. SPORE-SIZE IN CLAVARIADELPHUS LIGULA
Spore-size in µ Country Author

9-13X4-5·5 Austria Corner
11-14X4 Gt. Britain Cotton and Wakefield
IOX5-6 " Rea
10-15X3-5 Italy Bresadola
8-10X5 Japan Imai
10-14X3-4 Europe Burt
8-15X3-6, Av. = 11·2 X4·2 (E = 2·65) 1
15-18·5X4·5-5 U.S.A. (N.Y.) Coker

I
18·5-23x5·5-6 ,, (Utah)
15-23X4·5-6, Av. = 18·8x5·3 (E = 3·55)1
12-15x3-4·5 (E = 3·6)' U.S.A. Doty

7-12X3-4 (E = 2·71) 1 Burt
17·5-22X5-5·5 (E = 2·71)1 Japan
,,
Imai ( C. sachalinensis)
' E = length --,- width.

Burt and Coker describe the fruit-bodies of some N. American collections of C.
ligula as dull pink (Coker) or pinkish buff (Burt), becoming alutaceous or tinged fawn
or buff (Coker), or finally cinnamon or approaching Rood's brown (Burt). A pink
CLAVARIADELPH;US 279
colour seems never to have been noticed in European specimens (cf. C. pistillaris and
var. americdnus), nor do they become so dark brown. On the other hand, C. sacha
linensis is said to differ from the European C. ligula in its darker colour (? brownish or
cinnamon-ochraceous) and its longer spores (as given in Table XX). I suspect that
the American C. ligula is partly C. sachalinensis and partly C. ligula (as shown by
Burt's and Doty's smaller spore-measurements). Both species grow on coniferous
needles, but it seems that C. ligula may occur rarely on humus of frondose trees.
The records of C. ligula from Australia and S. Africa need confirmation.
C. maricolus (Kauffm.) comb. nov.
Basinym: Clavaria maricola Kauffm., Pap. Mich. Ac. Sci. Arts Lett. 8, 1928, 149:
Coker, Journ. El. Mitch. Sci. Soc. 63, 1947, 55, pl. 13, f. 3, 4.
7-10 cm. X 2-6 mm., simple, fleshy, erect, slender, rather ligulate and compressed
above, subcylindric below, tapering at the base, sometimes twisted and longitudinally
sulcate, solid, pale vinaceous buff when young, becoming light orange-yellow to orange
buff (Ridgway): flesh white (only the hymenium coloured), soft at the base with age,
not noticeably brittle: odour and taste mild.
Gregarious, often very abundant, on decaying needles, and, mostly, on decaying
male cones of Western Yellow Pine (P. ponderosa): U.S.A. (Oregon, Mich.).
Spores 10-14 x 4-5 µ,, white or slightly yellowish under the microscope, smooth,
subcylindric, obtuse, minutely granular within: 9·3-12·5 X 4·2-5·2µ, (Coker, type).
Basidia 108-112 X 4-5 µ,, with 4, rarely 2-3, sterigmata.
Cystidia none.
This is said to be near C. fistulosus and C. ligula. The long basidia, if correctly
measured, suggest affinity with C. pistillaris, but the length may refer to the thicken
ing hymenium. From C. fistulosus it seems to differ mainly in the brighrer, clearer
colour, somewhat ligulate shape, and habitat, but Harper has recorded C. fe.tulosus on
coniferous twigs in Michigan (his specimens may have been C. maricoti). From
C. ligula it differs merely in the brighter colour, narrower clubs, and lack of white
villous mycelium from the base of the stem.
C. mirus (Pat.) comb. nov.
Basinym: Clavaria mira Pat., Bull. Soc. Myc. Fr. 23, 1907, 71, cum ic.
8-15 cm. X 6-11 mm. wide at the apex, solitary or 2-3 fasciculate, simple, fleshy,
becoming hollow, cylindric, slightly attenuate at the base, apex obtusely rounded or trun
cate, mamillate: hymenium longitudinally rugose, smooth downward: wholly pale brown
or pale avellaneous.
On the ground in woods: lndo-China.
Spores 8-10 X 4-5µ,, white, ovoid, smooth.
Basidia 30 X 19µ,, clavate.
I have copied this description from Saccardo. The basidia seem extraordinarily·
wide and perhaps should be 30 x 9µ,. The fungus is clearly related to C. pistillaris,
but differs in the more slender, brown fruit-bodies and smaller spores. Thus it seems
to connect C. pistillaris with C. junceus. It shows how much mycologists may hope to
learn when the fungus flora of the Himalayas and the sources of the great continental
rivers of eastern Asia is known.
C. pistillaris (Fr. ) Donk
Basinym: Clavaria pistillaris Fr., Syst. Myc. 1, 1821, 477.
Synonyms: Clavaria herculeana S. F. Gray, Nat. Arr. 1, 1821, 657.
C. spathulata, Chev., FI. Gen. Paris, 1, 1826, 108, t. 8, f. 2.
Clavariella pistillaris (Fr.) Karst., Rev. Myc. 3, 1881, 21.
Text-Figs. 14B, 17, 18, 105: Plate 3.
280 CLAVARIADELPHUS
7-30 X 2-6 cm., subcylindric and acute, then blunt and more or less strongly clavate,
sometimes even turbinate or subtruncate, very massive, often longitudinally rugose,
sometimes very closely, light yellow then more or less deep ochraceous, sometimes rufescent
or brownish, brownishvinaceous on bruising, the hymenium always developing over the apex:
stem indistinct, white villous at the base: flesh firm and dense when young, becoming
soft, spongy-floccose, white, quickly turning purplish brown on cutting or bruising (when
fresh): taste rather bitter, edible.
On the ground in frondose woods, scattered or gregarious, never caespitose: Europe
(frequent, locally common), China (Yunnan).
Spores u-16 X 6-10µ, rarely as small as 9 X 5µ, white or tinged yellowish, smooth,
oblong-ellipsoid, apex blunt, often slightly waisted, thin-walled, with finely granular
vacuolate contents (not guttate).
Basidia 90-12oxII-13µ, rarely as small as 6ox10µ (50-75x8-13µ, Donk), pro
jecting up to 10µ, elongate, clavate with oleaginous contents: sterigmata 4, occasionally
2, 9-IIµ long X 3-3·5 µ wide at the base (5-6· 5µ long, Donk).
Cystidia none.
Hymenium gradually thickening from 100 to 300µ deep in old specimens, very
compact, with many included spores.
Hyphae 3-16µ wide, variable in width even in the same cell, with very long cells
50-600µ, with normal clamps or the clamps swollen up to 15µ wide, occasionally
with short lateral clavate cells up to 30 X 12µ.
var. americanus var. nov.
Synonyms: Clavaria pistillaris sensu Harper, Myc. 5, 1913, 261: sensu Coker,
Clav. U.S. Can. 1923, 83, t. 22, 23, 83, et Journ. El. Mitch. Sci. Soc. 63, 1947, 56-8:
sensu Wehm., Pap. Mich. Ac. Sci. Arts. Lett. 20, 1935, 256.
Differing in colour, form, and small spores from the European species: clubs often
flesh-colour or rose-pink, then ochraceous or alutaceous at maturity, often more or less
flattened or ligulate, occasionally with a few short antler-like branches, solitary or
caespitose in small clusters: spores 7-12·5(-14)x3·7-7·5µ (av. 9·8x5·5µ): basidia
60-70 X 6·5-8µ (Doty).
In frondose woods, rarely coniferous: U.S.A., Canada.
This most striking species of Clavarioid fungus is imperfectly known. The charac
teristic colour-change seems to have been unnoticed as, also, the very long cells of the
hyphae, the length of which is reflected in the basidia: nor has the extreme sponginess
of the texture been explained. On pages 45-51 I have described the structure and
development of the fruit-bodies.
There is no certain evidence, as yet, that the typical C. pistillaris of Europe occurs
in N. America. Var. americanus seems, rather, to connect C. ligula with C. truncatus
in colour, form, and spore-size. Coker says that there is a gradation from typical
clavate fruit-bodies to the truncate fruit-bodies with sterile apex, as in C. truncatus,
and thence to the perforate Craterellus unicolor, which I have called Clavariadelphus
unicolor. Coker (1947) has classed these as f. truncata and f. unicolor of C. pistillaris.
In Europe, C. truncatus seems distinguishable without difficulty from C. pistillaris
and, certainly, among the hundreds of specimens of C. pistillaris which I have seen
in the beech-woods of Buckinghamshire in England (1923-8) there was none which
varied in this manner: nor has such a state as C. unicolor been found in Europe.
Coker's illustrations suggest C. ligula (or a short-spored state of C. sachalinensis), and
the pink or rosy colour in the young fruit-bodies does not occur in the European
C. pistillaris. It seems to be this colour which distinguishes, also, the American C.
ligula and C. unicolor.
In Table XXI, I have listed the spore-measurements of C. pistillaris from Europe
and N. America, and in Table XXII, I have given spore-data from a single fruit-body.
The American collections have consistently smaller spores than the European, but
CLAVARIADELPHUS 281
with the same ellipticity. In contrast, the spores of the one fruit-body of C. pistillaris
vary in length from the upper limits of American collections to double their lowest
limit, yet the shorter spores are relatively wider than the American and the longer
TEXT-FIG. 105. Clavariadelphus pistillaris: part of the thickening hymenium and medullary
hyphae, x 500: spores, x 1 ,ooo.
spores relatively narrower. This is an important difference to which I have referred
elsewhere (1948). Further studies may show that var. americanus is a distinct species.
The records of C. pistillaris from S. Africa and Australia need confirmation. The
species appears to occur only in the frondose woods of Europe and, perhaps, China.
It has not been recorded from Japan.
282 CLAVARIADELPHUS
C. sachalinensis (Imai) comb. nov.
Basinym: Clavaria sachalinensis Imai, Tr. Sapporo Nat. Hist. Soc. II, 1930, 73:
ibid. 12, 1931, 19.
Differs from C. ligula in the darker coloured fruit-bodies and longer, narrower
spores, 17·5-22 X 5--5·5 µ,.
On the ground among Picea-needles: Japan (S. Saghlien, Mt. Kashipo).
TABLE XXI. SPORE-SIZE IN CLAVARIADELPHUS PISTILLARIS
Size in µ, Country Author

·-----
9-16 X5--'7 France Bourdot and Galzin
12-16x7-8 Gt. Britain Cotton and Wakefield
12-13 X7-8 Rea
12-15 X7-10 Comer
11-14·5 X6·7-9·5 " " r
10-16 X5--'7·5 Holland Donk
I 1-13 X6·5-8 China (Yunnan) Teng
9-16 X5-10, av. = 13 X7·3 (E = 1·78),' for Europe
7·4-10 X3·7-4·5 U.S.A. (Conn.) Coker, var. americanus
10-11 • 5 X5 · 5--'7 (N.Y.)
8-11·8 X3·8-5·5
9·3-12·5 X5'5--'7 "
7·2� X5·1-5·5 (N. Car.)
8·5-11·5 X5·6
8-11 X4-5 " ,, ,,
9-12 X4·5-6·5 Burt var. americanus
(9-)10-12(-14) X4·5-6(-7·5) " Doty var. americanus
8·5�·5 X5--'7"5 Canada (Nov. Scot.) Wehmeyer, var. americanus
7·2-12·5(-14) X3·7--'7(--'7·5), av. = 9·8 X5·5 (E = 1·78)' for var. americana
' E = length -:- width.
Though I have not seen the original description, I have indicated on page 278 my
reasons for supposing some, at least, of the N. American fungi, referred to C. ligula,
belong to this species. Here, too, may come the long-spored (18-22 µ,) collection of
C. pistillaris mentioned by Doty (1944, p. 26).
C. truncatus (Quel.) Donk

Basinym: Clavaria truncata Quel., Enchir. 1886, 221: Lloyd, Myc. Notes 64, 1920,
988, f. 1755: Konr., Bull. Soc. Myc. Fr. 39, 1923, 43, t. 3.
Synonyms: Craterellus pistillaris Fr., Epicr. 1838, 534: Harper, Mycol. 5, 1913,
261: Burt, Ann. Mo. Bot. Gdn. 1, 1914, 341: Wehm., Can. Journ. Res. 18c, 1940, 97.
Clavaria pistillaris auctt.: sensu Bres., le. Myc., t. 1102.
Clavaria truncata Lovejoy, Bot. Gaz. 50, 1910, 385: Doty, Clav. Pac. N.W. 1944,
24, t. 3, f. 23, t. l I.
6-15 cm. high, cylindric-acute then clavate, becoming turbinate with a truncate,
uneven, sterile top 2-9 cm. wide, sometimes concave or perfo.rate and more or less
pervious to the base, gradually attenuate downwards, yellowish, 'then the uneven,
rugose, or bullate top becoming bright yellow, ochraceous, orange, or rufescent; hymenium
becoming rugulose, yellow, or ochraceous-orange, then ochraceous rufescent or yellowish
cinnamon: stem 3-8 x 0·4-1·5 cm., sometimes swollen, attached to a white mycelial
network: flesh whitish or ochraceous, slightly brownish violaceous on bruising or cutting,
CLAVARIADELPHUS
fibrillosely fleshy then spongy, the surface-layer (? hymeniurn) brittle: taste sweet:
smell hone or slight: edible.
In coniferous woods: Europe, N. Africa (Algiers), Japan, N. America.
Spores 9-13 X 5-7(-8)µ, very pale ochraceous in the mass, smooth, elliptic-oblong,
thin-walled, with multiguttulate or granular contents: 10-12 x6-8 µ (Burt),
10-rr X 5-5·5 µ (Wehm.), 12-14 X 6-7 µ (Doty).
TABLE XXII. SPORE-SIZE IN A SINGLE FRUIT-BODY OF CLAVARIADELPHUS

PISTILLARIS (BURNHAM BEECHES, ENGLAND, 1927)
Number Number
Size in µ, E' of records Size in µ, E' of records
11·0X6·7 1·6 2 13·0X8·o 1·6 3
l I"OX7·0 1·6 l 13·0X8·3 1·6 l
11·0X7·3 1·5 l 13·0X8·7 1·5 l
11·ox7·5 l. 5 2 13·5X7·0 I"9 l
11·5X 6·7 1·7 l 13·5x7·3 1·9 l
11·5X7·0 1·6 6 13·5X7·5 1·8 5
11·5X7·3 1·6 2 13·5X7·7 1·8 l
n·5X7·5 1·5 2 13·5X8·3 I"6 2
II"5X 7·7 I"5 2 13·5x8·5 1·6 l
u·5X8·o 1·4 l 13·5X8·7 1·6 l
12·ox7·0 1·7 3 14·0X7·0 2·0 2
12·0X7·3 1·6 3 14·0X7·5 I"9 2
12·0X7·5 1·6 5 14·ox7·7 1·8 2
12·ox7·7 1·6 l 14·0X8·o I"8 4
12·5X7·3 1·7 l 14·0X8·5 I"6 2
12·5X7·7 1·6 2 14·0X9·0 I"6 l
12·5X8·o 1·6 4 14·5x7·5 1·9 I
12·5X8·3 l"5 l 14·5X8·o 1·8 3
13·ox7·3 1·8 l 14·5X8·5 I"7 l
13·0X7·5 1·7 l 14·5X87 1·7 l
13·0X7·7 1·7 l 14·5X9·5 1·5 l
I
Total range = u-14·5X6·7-9·5 µ,, E = 1·4-2·0, for 80 spores.

Average for 20 spores 11-u·5 µ, long, E = 1·56
12-12·5 µ, E = 1·62
13-13·5 µ, E = 1·71
14-14·5 µ, E = 1·78
Basidia 6ox 12µ, with 4 sterigmata 10µ long (Harper): 80-12ox6-10µ (Bres., as
Clavaria pistillaris, but possibly intended for Clavariadelphus pistillaris).
var. atrobrunneus var. nov.

Synonym: Craterellus pistillaris sensu Teng, Sinensia 6, 1934, 12.
Chocolate to warm blackish brown at the sterile upper end, gradually paler downward,
yellowish cinnamon on the stem.
6·5-9 cm. high, 1-2·5 cm. wide at the top, stem 0·8-1 cm. wide; clavate, truncate
or slightly umbilicate, the apex sterile, tapered downward, often bulbous at the base,
longitudinally rugose: flesh whitish, spongy.
Spores 10-12 X 6-8µ, white, smooth, ellipsoid, inequilateral.
On the ground in coniferous woods: China (Sikong).
This is so similar to C. pistillaris in size, colour, texture, and spores that they have
often been confused, and until the microscopic structure of C. truncatus is known the
CLAVARIADELPHUS
confusion may continue. It seems that C. truncatus differs from the true, or European,
C. pistillaris in growing in coniferous woods, in its smaller and, perhaps, relatively
narrower spores, and particularly in the turbinate shape of the fruit-body which is
said to be sterile over the apex, as in Cantharellus and Craterellus (sensu lato). It is
generally stated that the truncate apex is paler yellow than the hymenium, and sterile,
but as the sides of the top are often shown in illustrations as rounded it is difficult to
understand what morphological limits the sterile area can have: nor is it known
whether the sterile surface is covered with sterile basidia or unmodified hyphae ends.
For C. pistillaris I have shown (p. 50) that, until full maturity, on the cessation of
apical growth, its fruit-bodies are sterile over their pointed tops: if the apex remained
sterile, the intercalary growth of the hymenium would stretch and flatten the top and,
doubtless, pull the central tissue apart so that the fruit-body would become pervious
and hollow. Is this the state in C. truncatus? Or in C. truncatus is there a true dead
space formed over the apex of the primordial shaft by expansion and slight marginal
growth in the manner of a rudimentary pileus? Such occurs, evidently, in C. unicolor
which seems to bridge the gap between Clavariadelphus and Cantharellus.
As with C. ligula and C. pistillaris, the N. American fungi referred to C. truncatus
seem to be more varied. Burt's description agrees with the European, as does that
given recently by Doty, so that typical C. truncatus certainly occurs there, but Harper
described the apex of the club as reddish brown, the hymenium as often dull violet
with yellow tints towards the apex, and the stem as often bulbous, which points sug
gest transitions to C. unicolor. Coker reduced C. truncatus Quel., C. truncata Lovejoy,
and C. unicolor to Clavaria pistillaris, which is clearly unacceptable. Burt kept C.
unicolor and C. truncata Quel. distinct, but reduced C. truncata Lovejoy to C. truncata
Quel., with which I agree. C. truncata Lovejoy appears as a luridly described specimen
of C. truncata Quel., the sterile top of which may be bright yellow, bright orange, or
reddish orange in N. American specimens, according to Wehmeyer and Doty: C.
truncata Lovejoy was described as d�ll flesh-colour below, reddish orange upward to
the bright red pileate top (under balsam and spruce, Foxpark, Rocky Mountains,
2,900 m. alt.).
var. atrobrunneus is so differently coloured that it may be another species. Teng is
emphatic that the truncate apex is sterile, but its microscopic structure is not described.
The sweet taste of European C. truncatus is caused by mannite (R. Heim, Rev.
Myc. Cours Prat. de Myc. 1938, 68).
C. unicolor (B. et Rav.) comb. nov.
Basinym: Craterellus unicolor B. et Rav., Grev. 1, 1873, 148: Burt, Ann. Mo. Bot.
Gdn. 1, 1914, 340, t. 16: Murrill, Mycol. 32, 1940, 415, with photograph.
Synonyms: Craterellus corrugis Pk., Bull. Torr. Bot. Cl. 26, 1899, 69: (fide Burt et
Coker).
Clavaria unicolor (B. et Rav.) Doty, Clav. Pac. N.W. 1944, 23.
Clavaria pistillaris f. unicolor Coker, Journ. El. Mitch. Sci. Soc. 63, 1947, 58,
pls. 8, 9, 10, and 14 (f. 5, 6).
Fruit-bodies 3-7 cm. high, solitary, gregarious or caespitose, fleshy, at first cylindric
and acute, then becoming clavate or subpileate: pileus I· 5-5 cm. wide, at first cla·vate,
obtuse, soon obconic or turbinate, broadly convex or truncate, often abruptly cerebriforrn,
glabrous, sometimes perforate, flesh colour tinged with violet, then ochraceous buff, the
margin obtuse: stem 1-2·5 cm. x 5-8 mm., short, equal or attenuate downward, con
colorous or a little paler than the pileus: hymenium closely rugose, concolorous with the
pileus; flesh white, not discolouring on bruising, soft, soon shrinking and leaving the
pileus hollow: taste mildly bitter (Coker).
On the ground in mixed woods, or in troops under Picea· Engelmannii (in Oregon):
U.S.A.
Spores 8-12 X 4-6 µ (12-15 x 3·5-5 µ, Doty), white, ellipsoid-oblong, smooth.
CLAVARIADELPHUS 285
Basidia 7"5-9µ, wide, with 4 sterigmata 4·5-5·5µ, (Doty).
Murrill regards this as a good species, but Coker, after reducing it to C. pistillaris,
has recognized it as a form. Burt says that it is coloured very much as C. truncatus,
but that the pileus is more capitate and often cerebriform: whether it is sterile is not
mentioned. Violaceous colouring does not occur in the European C. pistillaris or
C. truncatus, though it may occur in C. pistillaris var. americanus which inclines to
pink in its early stages. Murrill also says that the spores are narrower than in C.
pistillaris and C. truncatus but gives no measurements. However, from Murrill's
photograph it is clear that C. unicolor has a true pileus with rudimentary marginal
growth. In shape it resembles incipient Craterellus cantharellus and, together with it,
is perhaps rather to be placed in Cantherellus: neither belongs with C. cornucopioides
in Craterellus s. str.
Doty's description of the species seems rather different and to indicate a fungus
more similar in form to C. truncatus. It develops as C. pistillaris, but the top of the
club dilates (possibly by intercalary growth of the hymenium), becoming hollow and
even depressed in the centre, where traces of the original acute apex of the primordial
shaft may remain as a mucro. His spore-measurements are longer and narrower than
those of other authors, and agree with Murrill's interpretation. In habit, he compares
it with C. ligula.
Coker regards C. unicolor as a lowland fungus in the southern U.S.A., compared
with the northerly C. pistillaris var. americanus which is montane in the southern states.
The distinction may relate to the forest rather than to the physiology of the fungi.
CLAVICORONA Doty
Lloydia, ro, 1947, 38-44, f. r-6.
Fruit-body erect, clavarioid, pyxidately branched, usually with a distinct stem, or
simple and narrowly obconic with sterile cyathiform apex: branches verticillate, narrowly
obconic, with sterile cyathiform tops, proliferating whorls of 2-8, mostly 3-4, similar
branchlets from the margin of the top, successive whorls more or less alternating: the
lignicolous species attached by a fibrillose-strigose or subtomentose subiculum of
secondary mycelium: white, light yellowish, ochraceous, fuscous-brown or tinged
lilac, growing tips white: hymenium waxy-mucilaginous, smooth, often thickening,
absent from the stem, from the bases of the branches on their upper sides and from
the cyathiform tops, often covered eventually by a thin sterile epihymenium of narrow
hyphae: flesh toughly gelatinous, dry and pliable or fibrous-subcoriaceous (waxy-soft in
two simple species), often acrid.
Lignicolous or terrestrial: 8 spp., temperate and tropical, uncommon.
Spores white, small, 3-6 X 2-3 · 5 µ,, smooth or punctate-rough, ellipsoid or subglobose,
thin-walled, aguttate.
Basidia small, subclavate, 4-spored.
Cystidia present or not: gloeocystidia large, traversing the subhymenium and
hymenium, immersed or with slightly projecting tips, -800 x 8-20 µ,, absent from sterile
surfaces.
Hyphae monomitic, clamped and inflating or dimitic with aseptate skeletal hyphae
of limited growth and clamped uninflated generative hyphae.
Type: C. taxophila (Thom) Doty.
KEY TO THE SPECIES OF CLAVICORONA
Ramosae, lignicolous, > 2·5 cm. high
Toughly gelatinous, white: Java, Sumatra . C. javanica
Fibrous coriaceous or pliable, dry, coloured
-13 cm. high, light yellow, ochraceous, alutaceous or
subrufescent: sp. 4-5 '< 2-3 /1-, smooth, ellipsoid: cystidia
present: mainly temp. . C. pyxidata
286 CLAVICORONA
(sp. rough, 4-5 X 3-4µ: S. Austral. var. asperospora)
-6 cm. high, fuscous-brown with livid greyish white rubbery
hymenium: sp. subglobose: cystidia absent: mainly trop.
Hyphae monomitic.: ? sp. smooth . C. Colensoi
Hyphae dimitic with brown, thick-walled skeletals of limited
growth: sp. 4-4·5 X 3-3·5µ, finely rough: Malaya. C. candelabrum
(Incertae sedis, from Mexico, see C. radiata, p. 294)
Simplices, narrowly obconic, sometimes with a few branches, small
Tough, lignicolous: sp. ellipsoid, 4-6 X 2-3µ
--7 mm. high, white or yellowish: apex entire, 1-2 mm. wide:
sp. subverruculose: Spain C. tuba
10-25 mm. high, vinaceous-buff or purplish grey, sometimes
sparingly branched, cristate round the top with 5-10 short
teeth: sp. smooth: U.S.A. . C. cristata
Waxy-fleshy, soft, white: apex 2-9 mm. wide: sp. subglobose,
3-4·5µ
On dead leaves and twigs under conifers: N. Amer. C. taxophila
Terrestrial: often sparingly branched or toothed at the apex:
sp. 1-guttate: France . C. Mairei
C. candelabrum (Mass.) comb. nov.
Basinym: Clavaria candelabrum Mass., Kew Bull. 1899, 172.
Text-Figs. 2F, 19, 20, 22, 23, 106: Plate 15.
2-6 cm. high, gregarious, fuscous-fawn or fuscous-bay brown with pallid white or
yellowish tips, becoming dark fuscous then fuscous grey or dark grey from the livid
greyish white rubbery hymenium: drying fuscous-brown or blackish.
Stem 5-15 X 1·5-2 mm., thickened upwards, rarely almost absent, smooth, con
colorous, attached by a thin fibrilloselyfioccose or felted fuscous-brown disk with fibrillosely
strigose edge: branches with internodes 7-13 X 0·3-1 mm., about as long as the stem,
obconic, the sterile cyathiform tops 1·5-2 mm. wide, proliferating 1-6, generally 4,
similar branchlets, at first subulate, the inner branches often abortive, generally erect,
in a narrow tuft, sometimes lax and sagging: hymenium thickening (-300µ) into a livid
grey, rubbery-gelatinous layer: flesh concolorous, fibrous-subcoriaceous, pliable: taste
very acrid: smell none or inparticular.
On very rotten trunks in the forest: Malaya (Selangor, Ridley 37, type: Singapore,
Bukit Timah, 4,4,41).
Spores 4-4·5 X 3-3·5µ, white, broadly ellipsoid to subglobose, pip-shaped, very
minutely punctate rough, aguttate (fresh).
Basidia 10-18 X 3-4µ, subclavate, aguttate, hyaline: sterigmata 2-3-4, mostly 4,
2·5µ long.
Cystidia none: gloeocystidia -600 (-800) X 8-20µ, 3-8µ wide at the obtuse apex,
cylindric-ventricose, tapering thin-walled, smooth, with dense granular-guttate cyto
plasm banded by transverse elliptic vacuoles, with a longer or shorter longitudinal
subhymenial part and a hymenial part ventricose below and tapering through a straight
or flexuous neck to the surface, not or scarcely projecting -8µ.
Hyphae dimitic: generative hyphae 1·5-6(--7)µ wide, not or scai;cely inflating,
mostly thin-walled, hyaline, clamped, some becoming thick-walled in older parts of
the fruit-body: skeletal hyphae 3-8µ wide, -600µ long, aseptate (or secondarily
septate near the apex), rarely branched, tapering gradually to the blunt or subacute
hyaline apex 1 ·5-3µ wide, with fuscous-brown smooth walls o·5-2µ thick, the narrower
interwoven skeletals in the older parts of the receptacle with almost linea;: lumen.
CLAVICORONA
Secondary mycelial hyphae at the base of the stem 1 · 5-3· 5 fL wide, distantly septate,
clamped, sparingly branched, mostly straight and radiating, with smooth fuscous
brown walls 0·5-1 µ, thick.
TExT-FIG. 106. Clavicorona candelabrum: the thickened hymenium in longitudinal section,

showing the adjacent medullary hyphae and pervading gloeocystidia: x 500.
C. Colensoi (B.) comb. nov.

Basinym: Clavaria Colensoi B., FI. N. Zeal. 1867, 186: Coker, Clav. U.S. Can.
1923, 94.
Synonyms:? Lachnocladiumfurcellarioides P. Henn., Monsunia 1, 1900, 142.
? C. piperata Kauffm., Pap. Mich. Ac. Sci. Arts Lett. 18, 1928, 14{>: Doty, Clav.
Pac. N.W. 1944, 21, t. 2, f. 6, 12, et Lloydia 10, 1947, 38-44, f. 5: Coker, Journ. El.
Mitch. Sci. Soc. 63, 1947, 56.
? C. candelabrum Mass. sensu Petch, Ann. R. Bot. Gdns. Per. 9, 1925, 338.
-3 cm. high, small, much branched from the compressed base, attached to the soft
decayed wood by short tow-like fibrils, wholly fuscous-brown, opaque and horny
brittle when dry: branches subfastigiate, pyxidately branched.
On decayed wood: New Zealand.
Spores?
Gloeocystidia long, many in the outer layer of the medulla next the subhymenium:
no cystidia.
Hyphae as in C. pyxidata, monomitic, with firm walls.
The type-specimens in the Kew herbarium appear identical with the Malayan
C. candelabrum, but they lack the thick-walled, tapering, skeletal hyphae. I find the
same discrepancy in the Ceylon collections 4064 and 5987 (Hakgala, -6 cm. high)
referred by Petch to C. candelabrum. Petch gives their spores as 3-4 x 2-3 µ,, smooth,
globose or ovoid, and m,ither the New Zealand nor the Ceylon fungus has cystidia.
It seems therefore that 1ere is a species almost identical with C. candelabrum but
CLAVICORONA
with the hyphal construction of C. pyxidata. The structure is difficult to make out in
dried specimens and the distinction needs corroboration from living material, but
until then I do not feel justified in reducing C. candelabrum to C. Colensoi.
C. piperata, on coniferous logs at Lake Quiniault in Washington State (U.S.A.),'
has-the small, but smooth, subglobose spores (4 x 3 µ) of T. candelabrum, the thicken
ing hymenium and the lack of cystidia, but the brighter colour of C. pyxidata-'pallid
to cinnamon-brown'. I refer it to this intermediate species, C. Colensoi, for the time
..-_ being.
Coker's reference of 'dark' ovate spores, 7-8·5 x 5-6·5 /L, to C. Colensoi is mistaken:
I found no distinct spores on the type.
Lachnocladium Jurcellarioides appears from its description identical with this or
with C. candelabrum. Hennings gives the spores as 3·5-4 x 3· 5 /L and likens the plant
to Ceratocladium Clautriavii (about which I can find nothing).
C. cristata (Kauffm.) Doty

Lloydia 10, 1947, 38-44, f. 1.
Basinym: Craterellus cristatus Kauffm., Pap. Mich. Ac. Sci. Arts Lett. 11, 1930,
172, f. 3: Overholts, Myc. 32, 1940, 260, f. 10: Doty, Clav. Pac. N.W. 1944, 21 (ut
C. coronata Schw.): Coker, Journ. El. Mitch. Sci. Soc. 63, 1947, 64.
Text-Fig. 107.
0·7-2·5 cm. high, 2-5 mm. wide at the apex, erect or suberect, narrowly cornucopioid
(but solid), tapering equally to the slender curved inserted
('./\/\ base, truncate-flattened or subconcave at the top, glabrous,
� \/V/V'I cinnamon drab, vinaceous, buff, or purplish grey (Overholts);
margin (at the top) entire then cristate with 5-10 slender fiat
triangular teeth or processes 1-2(-6) x 0·5-0·75 mm., forming
a marginal fringe round the apex of the fruit-body,
If - 0 occasionally with 2-4 short truncate obconic branches from
the margin (Overholts): stem -5 mm. long, short, concolor-
ous, indistinct, naked or slightly hairy at the base, inserted:
hymenium covering the flattened top and extending down
the sides to within about 5 mm. from the base, even, not
ridged, concolorous: flesh slightly tough, solid, paler con
colorous, translucent or hygrophanous (Doty): smell none.
On partly decayed logs of Pseudotsuga Douglasii and other
conifers: U.S.A. (Oregon), Canada (Quebec).
Spores 4-5(-6) x 2-2·3 µ, white, smooth, subcylindric:
5-6 X 2·5-3 /L (Overholts): 3·7-4·8(-6·5) X 2-2·5(-3) µ, white,
smooth, ellipsoid to nearly allantoid (Doty).
'TExT-FIG. r o7. Basidia 25 X 5 /L, ster. 4·5 /L (Doty). Hymenium 60-75 /L
.
Cla�icorona cristata: frmt- thick (Coker).
bodies, X 2: spores, X 1 ,ooo : G
loeocystl'd'1a 3-8 /L w1'de, extend'mg deep1y from the
(from Kauffman). .
subhymenmm to the surface of the hymemum, . flexuous.
The fruit-bodies are said to develop slowly. The larger processes round the margin
of the top may become 2-4-fid and subcristate themselves, or, according to Overholts,
they may elongate into short branches. Kauffman describes the hymenium as extend
ing over the discoid apex of the fruit-body, but this hardly seems possible. The
species is so near to C. pyxidata and C. Colensoi that it may be a juvenile state of one
1
-6 cm. high, pallid to cinnamon-brown, paler upwards: stem 2-3 mm. thick: branches
pyxidate, at times candelabriform, curved spreading, loosely arranged, dilated upwards with
rounded obtuse sinuses, branched 4-5 times, branchlets acuminate: fleshy, slightly tO\<lgh:
spores 4 X 3 µ., 3· 5-4·8 X 2· 5-3· 5 µ. (Doty), white, subglobose, smouth: gloeocystidia projecting
to 15 µ.: taste peppery.
CLAVICORONA
TEXT-FIG. 108. Clavicoronajavanica: young fruit-bodies and branches of old fruit-bodies, x 4.
of them. The length of the basidia, as given by Kauffman, is clearly erroneous and
refers, doubtless, to the thickness of the hymenium.
Doty refers this species to Clavaria coronata Schw. which Coker regarded as a
synonym of C. pyxidata.
C. javanica (Sacc. et Syd.) comb. nov.

Basinym: Clavaria javanica Sacc. et Syd., Syll. Fung. 14, 1890, 238.
Synonym: C. coronata Zipp. ex Lev., Ann. Sci. Nat. 3rd ser. 2, 1844, 2·15 (non
Schw. 1832 = C. pyxidata).
Text-Figs. 21, 108, 109.
Wholly waxy white, toughly gelatinous, drying light brownish ochre and horny-
5119
U
290 CLAVICORONA
membranous: mostly with 5-8 incipient peripheral branchlets at a node, but usually
3-5 developing fully.
-10 X 9 cm.: stem 2-17 X 2-3 mm., sometimes flattened -5 mm., attached by a
shortly spreading, substrigose, white mycelial patch, sometimes several fruit-bodies
I
;\
1' '\
11\
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I 11
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(
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t
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il/tl
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:111
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:.1
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TEXT-FIG. 109. Clavicorona javanica: hyphae of the medulla, some with oily contents and one
(in the centre) becoming secondarily septate: X 500.
arising from a common mycelial patch 6-10 mm. wide, generally distinct: branches
slender, narrowly obconic, 1-1·5 mm. at the thickened apex, in whorls of 3-5 (varying
2-8 but generally 4 5
- abortive), the tips subtruncate then subcyatniform and becoming
spicular-dentate or cristate round the periphery with the new branches, each internode
3-12 mm. long: hymenium covering the branches but absent from the subcyathiform
tops of the internodes and from the greater part of the trunk: taste?
Densely gregarious on very rotten fallen trunks in the forest: Java (type), Sumatra
(Brastagi, c. 1,800 m. alt., 14.9.31, Corner).
CLAVICORONA 291
Spores 3·7-4·5 X 2·5-3·5p,, white, broadly ellipsoid, smooth (? subpunctate), thin
walled (apparently aguttate).
Basidia apparently as in C. candelabrum (mature basidia not seen).
Cystidia none: gloeocystidia mostly 6-12 p, wide, thin-walled, granular-guttulate
as in C. candelabrum, often becoming secondarily septate, smooth, blunt, apex sub
clavate (at least when young), often irregular.
Hymenium thickening up to 250p,: subhymenial hyphae 2-3·5p, wide, with toughly
gelatinous walls, the hymenium covered by a gelatinous film, and developing an
epihymenium as in C. candelabrum.
Hyphae monomitic, 2·5-16 p, wide, very variable in size, the cells up to 4oop, long,
variously inflating, clamped, often variously swollen at the clamps, gelatinous and with
thin or slightly thickened walls (near the base of the trunk), smooth, colourless,
variously and irregularly branched, often with H-connexions, occasionally secondarily
septate, the larger hyphae often with oleaginous contents especially near the subhymenium
and with many transitions to the longitudinal and outcurving gloeocystidia in the
subhymenium and peripheral layer of the medulla.
Secondary mycelial hyphae 2-2· 5 p, wide, with sparse septa, clamped, slightly thick
walled, smooth.
The species is remarkable for its Tremellaceous or Caloceroid appearance, both
when living and dried. The part of the type-specimen of C. coronata Lev. from Java
in the Kew herbarium shows the same gelatinous hyphae and diffuse gloeocystidial
network in the medulla as in my Sumatran collection, so that I am confident in
identifying the two, though the macroscopic appearance of the type, as well as
Leveille's description based on the dried specimen, might suggest one of the three
non-gelatinous species. Coker gives the spores of Leveille's type as 3·8-,r2 x 3-3·5 p,.
C. Mairei (Battetta) comb. nov.
Basinym: Clavaria Mairei Battetta, Bull. Soc. Myc. Fr. 54, 1938, 44, f. 1-3.
Text-Fig. 110.
12-22 mm. high, 2-7 mm. wide at the apex, at first
as a simple, narrowly obconic club with subcyathiform
apex, then sometimes developing 2-5 short branches
themselves cylindric truncate, often undulate or toothed @
at the edge of the cup when not branched, white: stem
one-third to one-half the height of the fruit-body,
finely pilose, sterile, translucent when moist: hymenium
smooth, white: flesh compact, wa:1."Y fleshy, white drying
®
yellowish: taste mild.
On bare earth, solitary or gregarious: France (Bron,
Rhone).
Spores4·5 X 3·5 p,, white, smooth, ovoid-subglobose,
1-guttate, not amyloid.
Basidia (20-)28-35(-40) X 5·5 p,, elongate-clavate,
granular, forming a dense hymenium: sterigmata
4, 3p, long.
Gloeocystidia 72-115 x 5·5 p,, immersed or slightly
projecting -5 · 5 p,, subcylindric, obtuse, thin-walled,
with granular contents, arising deeply in the subhy-
menium or trama, TEXT-FIG. IIO. Clavicorona
Caulocystidia 17-23 X 5·5 p,, subcylindric, thin Mairei: fruit-bodies, X 1: spores,
x 1 ,ooo: gloeocystidium (top left)
walled, vacuolate, blunt, scattered: 18-20 X 2-3p,,
and caulocystidia (top right),
thick-walled processes from the base of the stem. x 500 (from Battetta).
Hyphae 10-20p, wide, narrower at the base of the
stem, thin-walled, longitudinal, forking, inflated (apparently without narrow hyphae),
J
292 CLAVICORONA
? clamps (not described or drawn): 5·5µ wide in the subhymenium, densely inter
woven.
Cyathiform apex of the fruit-body apparently sterile and covered with hyphal ends
and 'hairs' like the caulocystidia.
Battetta says this is near Clavaria Corbierei but differs in its Craterelloid-form,
wider hyphae, sterile apex, and cystidia. The inflated thin-walled hyphae prevent it
from being Podoscypha. It is conceivably a reduced Omphalia, cf. 0. integrella and
0. gibba, but more likely, in view of the incipient branching, a reduced Clavicorona.
C. pyxidata (Fr.) Doty

Lloydia 10, 1947, 38-44.
Basinym: Clavaria pyxidata Fr., Syst. Myc. 1, 1821, 470: Cooke, Handb. Austral.
Fungi 1892, 200: Coker, Clav. U.S. Can. 1923, 92, t. 26, 27, 84: Pilat, Bull. Soc. Myc.
Fr. 49, 1933, 50, t. 5, f. 3: Kauffm., Pap. Mich. Ac. Sci. Arts Lett. 8, 1928, 147:
Wehm., Pap. Mich. Ac. Sci. Arts Lett. 20, 1935, 264.
Synonyms: C. coronata Schw., Tr. Am. Phil. Soc. ii, 4, 1832, 182 (non Lev. = Clavi-
corona javanica).
C. Petersii B. et C., Grev. 2, 1873, 7.
C. laetissima B., Journ. Linn. Soc. Bot. 18, 1881, 386.
Clavicorona coronata (Schw.) Doty, Lloydia 10, 1947, f. 2.
-13 cm. high, copiously pyxidately branched, 4-6 branches in a whorl, clear yellow
becoming dull ochraceous with age or on bruising, sometimes pallid white then tan or
alutaceous, even pale pinkish or somewhat rufescent: stem 1·5-2·5 mm. thick, smooth,
thinly or densely whitish or brownish pink pubescent, springing from an amorphous
base or from a resupinate white or brownish pink pubescent mass one to several centi
metres wide: flesh white or pale, pliable, rather tough, peppery, often slowly, appar
ently slight or mild in some cases.
On dead wood, especially of Populus and Salix in the northern hemisphere,
gregarious or caespitose: Europe (Gt. Britain, France, Sweden, Finland, Czecho
slovakia, rare): N. America (U.S.A., Canada, Cuba, frequent): Asia Minor (Ilgaz
Dagh mountains, on Abies bornmuelleriana): Australia (Queensland, N.S.W.).
Spores 4-5 X 2-3 µ, white, ellipsoid, pip-shaped, aguttate, smooth: (3·3-4·5 X 2-2·5µ,
Wehm.).
Basidia 3·5-4·5µ wide, 4-spored.
Cystidia 18-45 x 4-711-, fusoid, subcylindric or subventricose, thin-walled, acute,
smooth, sometimes long-pointed or acicular, colourless, projecting slightly or -20µ.
Hymenium apparently not thickening (26µ thick; Coker).
Gloeocystidia -500 x 5-8µ, the greater part longitudinal in the subhymenium, with
short outcurving tips reaching the surface of the hymenium.
Hyphae 3-12µ wide, monomitic, inflated, clamped, oftP.n with the clamp much
enlarged, thin-walled or the wall thickening o· 5-1 · 5µ in the older parts, loosely
arranged in the medulla but denser and longitudinal at the periphery next the sub
hymenium.
This is the largest, most brightly coloured and, apparently, the most widely
distributed species. It is also distinguished, rather narrowly, by the more ellipsoid
spores and the presence of ordinary cystidia as well as gloeocystidia. Coker first
described the cystidia, and their presence was confirmed by Pilat and Wehmeyer,
though denied by Kauffman: I find them in two collections from Europe in the Kew
herbarium (Karst., Finland, and Romell, Sweden). If the hymenium does ncit thicken,
that is another important distinction.
Concerning the taste, Coker says it is peppery and Wehmeyer that the pepperiness
may be slight and slow to develop, but Kauffman says it is tasteless, while European
authors are silent. Kauffman suggests that Coker confuses C. piperata (? = C.
CLAVICORONA 293
Colensoi) with a tasteless C. pyxidata, and Doty (1947) separates the subsimple fruit
bodies referred to C. pyxidata by Coker as C. coronata Schw.
The fruit-bodies may develop imperfectly and mature in a rudimentary state.
'Young plants often lack the cups and if halted may reach maturity without their
formation. In such cases the peppery taste, which persists in drying, will usually
prevent confusion.' (Coker.) This may be the explanation of C. cristata.
The type of C. laetissima (Lockyer 477) in the Kew herbarium is identical in all
respects with C. pyxidata, and thus confirms the presence of the species in the southern
hemisphere. The record from Cuba may refer to one of the tropical species.
C. pyxidata var. asperospora Fawcett has the two kinds of cystidia of C. pyxidata,
as well as its large size, bui: it has the broader rough spores of C. candelabrum. Perhaps
it is the fully developed state of C. Colensoi. Its characters are:
Wood-brown to avellaneous, brownish even when young, the base cinnamon
brown: very peppery: cystidia of two kinds, either fusiform, hyaline, and pointed, or
as gloeocystida: spores 4-4·8x3·2-4/L, finely rough: hyphae -II/L wide, clamped:
on decaying wood (Eucalyptus, rarely Acacia), widely distributed in Victoria (Proc.
Roy. Soc. Viet. 51, 1939, 14, t. 4, f. 1, 4, 5).
C. taxophila (Thom) Doty
Lloydia 10, 1947, 38-44.
Basinym: Craterellus taxophilus Thom, Bot. Gaz. 37, 1904, 215, f. 1-8: Doty, Clav.
Pac. N.W. 1944, 22, t. 2 (non t. 6=? C. cristata).
Synonym: Clavaria taxophila (Thom) Lloyd, Myc. Notes 1920.
8-30 mm. high, mostly 14-18 mm., stalked with an elongate obconical truncate head
4-9 mm. wide at the sterile, plane, or slightly concave apex, white becoming ochraceous
with age and even orange upward, paler below: stem 0·5-1 mm. wide, solid, cylindric,
slightly dilating into the head, white pruinose, with scattered white hairs at the base:
margin of the head thin, upturned or expanded, even deflexed and almost involute
at times, wavy or lobed, or with incipient pyxidate branching (Doty): hymenium
covering the sides of the head, smooth or becoming vaguely longitudinally rugulose,
especially when dry: flesh waxy, very soft, hygrophanous.
On moist, very rotten leaves and twigs of deciduous trees and conifers under
prostrate branches of Taxus canadensis and Pseudotsuga taxifolia, solitary or 2-3
together, or subcaespitose (Doty): U.S.A. (Fall Creek Gorge, Ithaca, N.Y.: Or., Cal.).
Spores 3-4/L wide, broadly elliptic to subglobose, smooth, white: 3·2-3·8 X 2·2-3/L,
smooth, oblong-ovoid (Doty): 3-4 X 2-3 /L (Coker).
Basidia 18-24x4/L, clavate, with 4 sterigmata 4/L long: 25-3ox4·5/L (Doty).
Gloeocystidia -70 X 4-6/L, subcylindric, irregularly swollen in places, often curved,
thin-walled, wholly immersed or with the tips slightly protruding.
Hymenium apparently thickening: 25-40 /L thick (Coker).
Hyphae evidently monomitic and thin-walled: clamps?
As interpreted by Doty, this species is certainly a juvenile state or rudimenta ry
species- of Clavicorona, and suggests alliance with C. javanica. However, Thom did
not observe any incipient marginal branching in his specimens. The association with
Taxus, as Thom observed, is ecological, not physiological. C. Mairei may be identical.
C. tuba (Heim) comb. nov.
Basinym: Physalacria tuba Heim, Fungi Iberici 1934, 50, f. 12.
Text-Fig. 111.
5-7 mm. high, 1-2 mm. wide at the apex, clm·ate, shallowly cupuliform at the apex,
the entire margin often slightly repand, ichite then slightly yellowish: stem 2 X o· 5-1 mm.,
hyaline, bulbous at the base, with long, spreading, silky, distant hairs, scarcely distinct
from the clavula: flesh solid.
On rotten wood: Spain (Catalonia).
294 CLAVICORONA
Spores 4·2-5· 5 x 2·5-3µ,, white, slightly verruculose, ellipsoid-cylindric.
Basidia c. 40 X 4-5µ,, subclavate: sterigmata 2-4, c. 4µ, long.
Gloeocystidia 70-110 X 5-11µ,, cylindric, of uneven
width, the tips projecting -20µ, and often subacuminate,
thin-walled, with oleaginous contents, arising deeply
from the subhymenium or from the trama (often with a
longer or shorter longitudinal base).
Hyphae 4-5·5µ, wide, with numerous clamps, walls
0·8-1µ, thick, brownish.
This was said to resemble Physalacria stilboidea and
P. Langloisii, but the solid, narrowly obconic head (?
with sterile apex) and. the gloeocystidia are different
from Physalacria. It seems to be a reduced ally of C.
javanica and C. candelabrum, perhaps even a juvenile
form. Doty regards it as a synonym of C. taxophila,
but the texture and longer ellipsoid spores of C. tuba
seem different.
SPECIESINCERTAE SEDIS
Clavaria radiata Lev.
Ann. Sci. Nat. Bot. Ser. 3, 5, 1846, 156.
©·. 5-6 cm. high, gregarious, branched: stem 1-2 cm.,
0
dilating into a cup and proliferating branches from both
0
the upper- and undersides of the cup, the branches
similarly dilating into cups and branching three to
four times: ultimate branchlets acute: Mexico (Vera
'I'ExT-FIG. I I I. Clavicorvna Cruz).
tuba: fruit-body, X s: spores, Possibly C. pyxidata or C. Colensoi.
x I ,ooo: hymenium, hyphae,
and gloeocystidial tips, X 500:
(from Heim). CLAVULINA Schroet.
Cohn, Krypt. FI. Schles. Pilze 1888, 442.
Synonym: Stichoramaria Ulbrich, Lindau Krypt. FI. f. Anf. 1, 1928, 83, 3rd ed.
( = an exact synonym based on C. cristata, teste Donk, 1933, p. 17).
Fruit-bodies simple or branched, generally with flattened branching and more or less
cristate tips, in some species slightly dorsiventral with a sterile hymenium on the
uppersides of the main branches: white or variously coloured, never bright yellow,
orange, red, or black: flesh tough, waxy-fibrous, or rather brittle: mostly terrestrial,
!l few lignicolous, rarely on herbaceous remains (C. cristata var. incarnata).
Spores white(? brown in C. decipiens), generally ochraceous after prolonged drying
(in the herbarium), subglobose, or broadly ellipsoid, blunt, smooth, with I large gutta
nearly filling the spore, occasionally multiguttulate, 5-14µ, long.
Basidia subcylindric, projecting, usually becoming secondarily septate with one or more
septa after spore-discharge, stichobasidial (? in all species): sterigmata 2, rarely x,
generally incurved and short, in some species nearly straight.
Cystidia present in a few species, generally absent.
Hymenium thickening: subhymenium persistently filamentous or becoming pseudo-
parenchymatous.
Hyphae monomitic, with clamps in most species, generally with colourless, slightly
but distinctly thickened, walls (pale brown in C. ornatipes and C. decipiens), secondarily
septate only in the species without clamps, mostly short-celled (20-120µ, long), more or
less inflated but often irregularly: H-connexions frequent.
CLAVULINA 295
Flesh consisting of a wide loose core (becoming hollow in some species) of more
inflated, loose, and generally contorted or convolute hyphae, and a more compact
peripheral layer of longitudinal hyphae with narrower and generally shorter cells,
particularly next the subhyrneniurn, the outer laying retaining the core and stretched
I,
��
Terrestrial, rarely lignicolous: 32 spp., in temperate and tropical regions.'
Type-species: C. cristata (Fr.) Schroet.
KEY TO THE SPECIES OF CLAVULINA

(Compare Clavaria, Clavulinopsis)
Hyphae of the fruit-body with brown walls, 3-8µ wide, little
inflated: basidia 20-40 X 5-'7µ: hyrneniurn unilateral:. branched:
subgen. Fusco-Clavulina A
Hyphae with colourless walls, usually inflated: basidia usually
larger: subgen. Eu-Clavulina
Hyphae of the fruit-body without clamps, often secondarily
septate: hyrneniurn generally unilateral (amphigenous in the
subsirnple C. fioridana): generally branched B
Hyphae clamped at all septa (so far as known): hyrneniurn
unilateral or amphigenous: branched or simple
With cystidia C
Without cystidia D
A. Fusco-Clavulina
Hyphal walls 0·5-1·5µ thick: -5 cm. high, cinnamon-fawn, slender:
stern -2 cm. X 0·8-1·5 mm., slightly spicular-strigose at the base,
attached by a thin brown mycelial felt: branches tapering to filiform
tips, erect, fastigiate, little divided: sp. ? pale brown: terrestrial or
lignicolous: Malaya C. decipiens
Hyphal walls slightly thickened, 0·5µ: -9 cm. high, Juscous-brown,
robust: stern -6 cm. X 2-8 mm., densely strigose spiculose: branches
rather stout, few, palmate, tips subulate (not filiforrn): sp.
white: terrestrial: U.S.A., Japan, Malaya C. ornatipes
(Compare C. odorata: stern not spicular-strigose, very short: Finland)
B. Eu-Clavulina, without clamps
Coriaceous, drying cartilaginous,, at least in the stem and main
branches: -r I cm. high, pale pinkish drab or pale vinaceous-drab,
tips white: sparingly or much branched, commonly in one plane,
palmato-fiabellate, fastigiate: branches becoming rugulose: sp.
7-9·5 X 6-8·5µ: caespitose, terrestrial: W. Indies, Malaya
(? Africa) . C. cartilaginea
. Fleshy, fibrous or somewhat floccose, not palrnato-flabellate: trop.
Arner.
Simple or sparingly branched, densely caespitose, cinereous,
stern yellowish: sp. 7-8·5 X 6-7·5µ: terrestrial, Florida C. fioridana
Branched more or less profusely: rather fibrous or fibrillose-
floccose when dry: (? white or yellowish)
Sp. 8-10 X 7-8µ, subglobose: -4 cm. high: on trunks: Brazil C. Sprucei
I Edible species are C. amethystina, C. cinerea, C. cristata, C. rugosa,
296 CLAVULINA
Sp. ellipsoid
Sp. 7·5-9 X5 ·5-6· 5 µ.: -5 cm. high: on the ground: Brazil C. connata
Sp. 9·5-11 x5-6 µ.: -1·5 cm. high, in very delicate tree-like
tufts: on dead leaves and twigs: Brazil C. delicia
C. Eu-Clavulina, with clamps and cystidia

-10 cm. high, white, then light yellow, then more or less suffused
fuscous-vinaceous, violet-brown, or fuliginous: branches few and
lax, or many and tufted: stem often well developed: sp.
7-10 X6·5-9 µ.: cystidia cylindric clavate, stout, becoming thick-
walled and incrusted: terrestrial: Malaya, Java C. Lr!veillei
[-2 cm. high, simple, compressed, subfusiform, obtuse, (brownish
when dry): sp. 6-8 X6-7 µ.: cystidia 6 µ. wide, smooth: terrestrial:
Porto Rico C. pilosa]
D. Eu-Clavulina, with clamps, without cystidia

Grey or fuliginous, sometimes tinged purplish
Simple or with few simple branches
Sp. 7·5-9·5 X6·5-8 µ.: simple: on wood, tree-ferns: Tasmania C. tasmanica
Sp. 9-13 x8-u µ.: simple or sparingly branched: N. temp. C. rugosa var.
Much branched: sp. 7-11 X7-10 µ.: N. temp.
White becoming fuliginous, often cristate . C. cristata var.
Grey or fuliginous from the first, rarely cristate C. cinerea
More or less lilac or purple, inclining to greyish
Branches many, obtuse, clear lilac-purple or lilac-violet: -6 cm.
high: sp. 7-11 X6-8µ.: terrestrial: Europe C. amethystina
Simple, sparingly branched or with many branches, pale lilac,
-ro cm. high: sp. 7-1ox6-8 µ.: terrestrial: U.S.A. C. amethystino
ides
White, then yellowish, finally vinaceous, violaceous, brownish, or
fuliginous: sp. 8-10 x7-9 µ.: trop. Asia C. Leveillr!i
..
var. atricha
(Violaceous-grey or fuscous-purple, much branched, stout:
Europe. C. cinerea
var. sublilascens)
Pinkish, pinkish brown, or vinaceous-brown (not white at first)
Branches many, palmate-digitate, pale pinkish tussore, then
brownish salmon: -7· 5 X12 cm.: sp. 5-7 µ.: terrestrial: Austral. C. complanata
Branches irregular, few or many
-5 cm. high, vinaceous-fawn to fawn, with vinaceous-pink tips:
sp. 7·5-1ox6·5-9 µ.: terrestrial: Austral. C. vinaceo-
cervina
-10 cm. high, occasionally simple, pale pinkish drab, pinkish
tan, fawn, or lilac: sp. 7-10 X6-8 µ.: terrestrial: U.S.A. C. amethystino
ides
Simple, stout, often rugulose, pinkish: Europe C. rugosa var.
Simple, slender, with filiform brown stem and pinkish or white
head: sp. 8-11 x6-9µ.: U.S.A. C. castaneipes
CLAVULINA 297
White or yellowish, in some species becoming pinkish, greyish,
vinaceous, ochraceous, or cinnamon: mostly terrestrial
Sp. 5-7µ, long, subglobose
Simple or irregularly branched
-5 cm. high, white, then subcinereous, pale greyish brown or
cartridge-buff: sp. 5·5-7·5µ,: terrestrial: S. Austral. C. subrugosa
-1 cm. high, white: sp. 7x5µ,: terrestrial: France C. Bessonii
Much branched
White, -6 cm. high: sp. 4·5-5 X4µ,: France C. gallica
Ochraceous-white, -15 cm. high: sp. 5-6µ,: Brazil C. cirrhata
Pale alutaceous, then cinnamon, odoriferous: sp. 5-7µ,:
Finland . C. odorata
Sp. larger
Much branched
-15X9 cm., pale yellowish drab with cream-buff tips, then
dingy fawn or subfuliginous: hymenium pale fawn drab,
unilateral: smell of chocolate: sp. 7-8·5x6·5-·r5µ,:
Malaya . C. gigartinoides
-10 cm. high, white, sometimes becoming yellowish or
greyish, often cristate, very variable: hymenium rarely
unilateral: sp. 7-11 X6·5-10µ,: temp. regions C. cristata
-2·5 cm. high, golden(? dried), drying cartilaginous, tough:
sp. 7-8·5x5·5-7µ,: Surinam C. chondroides
Congo: -8 cm. high, alutaceous(? dried): sp. 7-8X6-7 µ, . C. Vanderystii
Brazil: -5 cm. high, ochraceous(? dried), rather fibrillose-
floccose: sp. 7-9 X 6-7µ, C. panurensis
Sparingly branched or occasionally simple: trop., S. temp.
-7"5 cm. high, white, then pinkish grey, vinaceous-drab, or
pale pink, palmately branched, tips subulate or subfili
form: stem becoming hollow: sp. 8-1ox7-8·5µ,: on the
ground or on rotten wood: Malaya C. cavipes
-10 cm. high, white then yellowish, finally often vinaceous,
brownish, or fuliginous: sp. 8-10x7-9 µ,: trop. Asia C. Leveillei
var. atricha
-r5 cm. high(? more), white: sp. 8-1ox7·5-9µ,: on wood:
New Zealand . C. humilis
(See also C. rugosa, sp. 9-14X8-11 µ,: N. temp.)
Simple, sparingly branched in forms of C. rugosa(N. temp.)
Smell strong, of fenugreek: taste bitter, very unpleasant:
-9 cm. X2-6 mm. pale yellow-white, fusiform: sp.
10-12x9·5-10·5µ,: Malaya C. ingrata
(Smell of garlic: -6 cm. high, white to yellowish or fuscous:
sp. 7-10X4·5-6µ,: N. temp. Clavaria fuscata)
Without strong smell or taste
-8 mm. high: sp. 7-11 X6-7µ,: Argentine C. pampaeana
-6 cm. high, very slender: stem brown: head white or
pink: 8-11 x6-9µ,: U.S.A. C. castaneipes
298 CLAVULINA
-6 cm. X 1-2 mm., white then yellowish, slender: sp.
8-10µ., globose: Malaya C. gracilis
Generally larger, often longitudinally rugulose, or sulcate
and hollow, or with few blunt branches, very variable:
sp. 9-14 X 8-rrµ.: N. temp. C. rugosa
C. amethystina (Fr.) Donk
Basinym: Clavaria amethystina Fr., Syst. Myc. 1, 1821, 472: Bourd. et Galz.
Hym. Fr. 1928, jI-06.
Synonyms: Clavaria amethystea Pers., Myc. Eur. 1, 1822, 165.
? Clavaria coerulescens Karst., Myc. Fenn. 23, 2: Sacc., Syll. 1891, 249.
Ramaria amethystea S. F. Gray, Nat. Arr. 1, 1821, 655.
R. amethystina (Fr,) Quel., FI. Myc. 1888, 463 (quoad nomen).
Misinterpretations: Clavaria amethystina sensu Cott. et Wakef., Coker, Rea, Ito et
Imai, (Bres.?) = Clavaria Zollingeri Lev.
Ramaria amethystina sensu Quel., Ass. Fr. 1891 = Ramaria fennica (Karst.), vel
prox.: sensu Quel., FI. Myc. 1888 = ? Clavaria Zollingeri.
2-6 cm. high, solitary or caespitose, wholly lilac-violet, paler or whitish at the base:
stem short and stout, or practically none: branches numerous, cylindric smooth, then
rugulose, obtuse or toothed, not cristate: flesh slightly brittle, concolorous: without
particular smell or taste.
On the ground in frondose woods: central and western Europe, rather rare (France,
Holland, Germany, Italy, Sweden).
Spores 7-rr(-12) X 6-8µ., white, ovoid-ellipsoid or subglobose, with 1 large gutta.
Basidia 42-60 X 4-8µ.: sterigmata 2.
Hyphae 4-18 µ. wide.
var. lilacina Quel.
Clav. Syn. 1878, 223: Rea, Brit. Bas. 1922, 706.
Synonyms: Clavaria lilacina Fr., Hym. Fr. 1874, 667 (non Jungh. 1844).
Ramaria lilacina (Fr.) Quel., FI. Myc. 1888 (quoad nomen: quoad sporae?
= Clavaria Zollingeri).
Clavaria Schaefferi Sacc., Sy!!. 6, 1888, 693.
Clavaria sublilacina Karst., Finl. Nat. o. Folk 48, 1889, 375.
Clavaria amethystina var. purpurea Bourd. et Galz., Hym. Fr. 1928, 106.
Caespitose, lilac-purple, or somewhat brownish, especially on drying: branches
firmer, rigid, linear, less numerous, often twisted.
Spores 7-10 X 6-8µ..
On the ground in frondose woods: central and western Europe, rare.
This species has often been confused with the brittle Clavaria Zollingeri; the
basidia and hyphae are, at once, distinctive under the microscope. Donk was the first
to disentangle them, and I have followed his nomenclature.
C. amethystina seems to differ from C. cinerea only in colour, but there may be
vinaceous tints in C. cinerea, and its f. sublilascens seems to bridge the difference. It is
rare and seems not to occur in the British Isles.
Clavaria coerulescens Karst. was described as: 'Tota coerulescens: forma magni
tudineque cum Cl.fiava conveniens: in paroecia Hollola Fenniae.' It may be Ramaria
fumigata.
C. amethystinoides (Pk.) comb. nov.

Basinym: Clavaria amethystinoides Pk., Bull. Torr. Bot. Cl. 34, 1907, 102: Coker,
Clav. U.S. Can. 1923, 65, t. 9, 14, 82, 83.
3-10 cm. high, solitary or gregarious, with irregular, subsimple, antler-branchings
Cl,AVULINA 299
like C. rugosa, to rather frequently branched, occasionally simple, pale livid flesh colour,
pale drab, pinkish tan, fawn, or very pale lilac, the tips eventually blackening: stem
0·5-3·5 cm. X 1-2·5 mm., short or elongate, rather darker than the branches: branches
simple or palmately branched, and flattened -10 mm. wide, even subcristate, often
irregular, generally few, ascending, the tips blunt or pointed: flesh soft and rather
waxy-brittle in the branches, tough and pliable in the stem: without smell, taste faint
and inparticular.
On the ground in damp, shady places: U.S.A. (N. Car., S. Car., Mass.).
Spores 7-10 X 6-8 µ., white, smooth, subglobose to broadly ellipsoid, 1-guttate.
Basidia (22) X 7-10 µ.: 1 sterigmata 2.
Cystidia none.
Hymenium 37 µ. thick at first, thickening to 130 µ..
Hyphae -9 µ. wide, with rnoniliform rows of short, inflated cells.
This species has been found only in the temperate part of N. America. Coker says
that it is close to C. rugosa and differs mainly in its colour. The spores, however, are
those of C. cristata or C. amethystina, and the palmatifid branching is like that of
C. cristata var. nivea or var. laciniata, rather than the obtuse branching of C. rugosa.
From C. amethystina, to which it seems most nearly allied, C. amethystinoides differs
in its simpler form, palmatifid branching and paler colour. In these respects it
approaches the tropical C. cavipes, under which I have made comparison. Because of
these differences and because C. amethystinoides produces its own series of subsimple
or simple degenerate forms, I think it is a separate species. Compare, also, C. carti
laginea.
Coker says Clavaria ligata Britz. (Hym. Sudb. 5, 1888, 290, f. 37) is very near
C. amethystinoides and may be the same. The description, however, fits subsolitary
Clavulinopsis fusiformis !
C. Bessonii (Pat.) comb. nov.
Basinym: Clavaria Bessonii Pat., Tab. An. 1885, 163, n. 359.
4-10 mm. high, white, simple OT sparingly branched: stem indistinct, rufescent tomen-
tose at the base: apex sterile: hymenium continuous almost to the base.
On bare soil: France (Jura).
Spores 7 X 5 µ., white, smooth, ovoid, 1-guttate.
Basidia 2-spored.
This may be a very juvenile state of C. cristata or of Clavaria acuta, which, however,
does not branch. Donk agrees (in litt.) that it is probably a Clavulina.
C. cartilaginea (B. et C.) comb. nov.
Basinym: Lachnocladium cartilagineum B. et C., Journ. Linn. Soc. Bot. 10, 1868,
330: Pat., Journ. de Bot. 3, 1889, 26, t. 1, f. 4: Burt, Ann. Mo. Bot. Gdn. 6, 1919, 269,
t. 5, f. 2: Coker, Clav. U.S., Can. 1923, 197-8.
Text-Figs. 6B, 25, 26, u2-18: Plate 5.
3-11 cm. high, often densely caespitose, sometimes gregarious or solitary, sparingly
to much branched, generally palmately or flabellately, twisted, fastigiate, rather dry and
tough, pale pinkish drab, pale pinkish cinereous drab, with elongate or cristate white tips,
occasionally blunt.
Stem o· 5-4 cm. X 2-4 mm., or flattened -6 mm. wide, tapered downward, flattened
upward, often becoming rugulose and twisted, generally strigose-fibrillose at the base OT
in the lower part, with abortive filiform primordia and irregular rugulose cristations,
concolorous: branches 1-2 mm. wide, rather narrow, cylindric or ligulate, palmately or
fiabellately polychotomous (3-8-chotomous) below, flattened -10 mm. wide beww the
points of branching, dichotomous or cristate above, branching in one plane, but much
1
The short length, 22 µ,, given by Coker, may refer only to the terminal part of the old
basidiwn cut off by the secondary septum.
-
300 CLAVULINA
twisted, curved, and distorted, µpright and more or less fastigiate, or somewhat spread
ing, not very dense, becoming longitudinally rugulose, the branchlets elongate-ligulate,
or linear and unbranched 3 cm. long or cristate with small points, occasionally blunt,
rugulose or even tuberculate: flesh rather dry, floccose, pliable, elastic, generally break
ing when bent double, distinctly coriaceous or cartilaginous in the stem and main
branches, drying cartilaginous-coriaceous, the hymenium waxy: without smell or
taste.
Tmrr-FIG. n2. Clavulina cartilaginea, X 2.
In the ground in the forest: Cuqa (type): Malaya (Singapore, Bukit Timah,
occasional, 27.8.40, 4.4.41: Negri Sembilan, Pasoh Forest Reserve, 25.6.30): ? Gold
Coast (on dead wood, Kew Bull. 1901, 164): (W. Africa, Hennings, but ?).
Spores 7-10 X 6-8·5 ft', white, smooth, subglobose or broadly ovoid, with slightly
thickened wall, 1-guttate, multiguttulate, or with wholly vitreous contents.
Basidia 30-50 X 5-8µ., rather narrow, subclavate, with oleaginous guttulate contents,
projecting -15µ.: sterigmata 2, 5-7µ. long, varying from distinctly curved to nearly
straight, rarely only 1 sterigma.
Hymenium thickening -180µ.: absent from the base of the stem and the uppersides
of the main oblique branches: subhymenium at first filamentous with cells 8-15 x 3·5-
6·5µ., soon pseudoparenchymatous, the cells 12-30 X 5-15µ., and often somewhat moni
liform: cystidia none.
CLAVULINA 301
Hyphae of the fruit-body mostly without clamp-connexions, a very few clamps on the
hyphae of the flesh, the basidia and subhymenial hyphae without clamps: hyphae of
the flesh longitudinal, often contorted, mostly inflated, a few not or little inflated,
sparingly or rather frequently secondarily septate, more or less constricted at the
septa, often swollen on one or both sides of the septum, the cells 20-150 X 3-13 p,, in places
-20p, wide, often unequally inflated, the walls becoming slightly thickened 0·5-0·8 p,
thick (or more at the base of the stem) and often irregularly undulate, smooth, with
H-connexions: the hyphae narrower, short-celled, and very compact near the sub
hymenium, lobser in the central tissue.
TEXT-FIG. n3. Clavulina cartilaginea, X 1.
Mycelial hyphae 3-4·5 p, wide, the walls 0·5-1p, thick, smooth, with clamp-connexions.
Sterile hymenium, on the uppersides of the oblique main branches, c. 50 p, thick,
the terminal cells 10-20 X 3-7p, as sterile basidia, cylindric or subclavate, blunt, rarely
with abortive sterigmata, set in a fairly regular palisade, the subterminal cells-30 X 10 p,
and pseudoparenchymatous.
Sterile hymenium on the stem, for 1-10 mm. from the base, as a cortex 50-1oop,
thick, composed of radial rows of inflated globose cells, 7-15 p, wide, the rows simple
or branched, often with the cells diminishing in size distally, compact but not truly
pseudoparenchymatous, sterile or, in the upper half of the stem, with scattered sterile
and fertile basidia: passing gradually into the hymenium on the underside of the
main branches or into the sterile hymenium on their uppersides: rarely, the sterile
base of the stem inoderm with narrow hyphae 3-6p, wide and appressed or slightly
divergent tips.
-
Tllx:T-FIG. 114. Clavulina cartilaginea, x 2.
Tllx:T-FIG. u5. Clavulina carti/aginea, X 2.

TExT-FIG. 116. Clavulina cartilaginea: hyphae of the medulla (often secondarily septate),
that on the right being a subhymenial hypha: rarely clamped: X 500.
TExT-FIG. u7. Clavulina ca11tilaginea: the thickening hymenium, X 1, 000 .
CLAVULINA 305
This species is very distinct in its flabelliform branching in one plane (though the
branches become twisted), its tough and almost cartilaginous flesh, the pseudo
parenchymatous subhymenium, the algal-like cortex on the stem, the irregularly
swollen cells, often like 'trumpet-hyphae' at the septa, and the absence of clamps.
TExT-FIG. 1 18. Clavulina cartilaginea: surface of the stem with rudimentary

sterile hymeniwn, X 1,000.
The cells of the medulla become curved and contorted by the small amount of
secondary septation, so that the tissue appears very irregular, at first sight. The
presence of clamps on the mycelial hyphae and their absence from those of the fruit
body have been noted in other fungi, as Coprinus lagopus (by Buller).
The stem-cortex thickens gradually throughout the life of the fruit-body. The
terminal cells of the moniliform rows elongate and cut off clavate or lenticular distal
cells which slowly inflate and, in their tum, send out new apical protrusions. Originally,
5119 X
306 CLAVULINA
on the primordial shaft, the cortex develops as the hymenium from out-turned hyphae
from which 2-4 cells are cut off to inflate acropetally and start the primary cortex.
I have examined the type-specimen in the Kew herbarium. It agrees exactly with
some forms of the Malayan material but has narrow hyphae 2-5(-8) µ wide. The sub
hymenial hyphae are swollen, however, 15-30 x 6-10µ, and there are trumpet
swellings on some of the medullary hypha�, and no clamps. The spores are 7-9 X 6-8 µ
(9 X 6 µ, Burt: 6·2-8·5 X 5·5-7·5 µ, Coker): the basidia are 2-spored, as Coker found:
and the hymenium is thickened to 150µ. The base of the stem is broken off the speci
men and so the presence of the characteristic cortex cannot be checked. My descrip
tion of the species is based on the Malayan collections. It does not seem to be common.
C. castaneipes (Atk.) comb. nov.
Basinym: Typhula castaneipes Atk., Ann. Myc. 6, 1908, 60.
3-6 cm. high, simple, rarely once branched; head 6-20 X 1·5-2·5 mm., white or pale
rose-pink, cylindric-subclavate or cylindric: stem filiform, brown, glabrous, straight, or
curved.
On the ground in woods: U.S.A. (N. Car.).
Spores 8-11 X 6-9µ, white, smooth, obovate-subglobose.
Basidia 2-spored.
The position of this species is doubtful. The large subglobose spores and 2-spored
basidia suggest Clavulina, but the long filiform stem is unusual and suggests Clavuli
nopsis or Clavaria (under which I have also keyed the fungus). As a terrestrial fungus
without sclerotium it cannot be Typhula.
C. cavipes sp. nov.
Text-Figs. n9-21.
-'r5 cm. high, solitary, gregarious or subcaespitose, sparingly branched dichotomously
or palmately, narrow, occasionally simple and even forming small tufts of simple acerose
clubs -3·5 cm. high, white, the branches becoming pinkish grey or vinaceous drab or
merely tinged pink.
TEXT-FIG. 119. Clavulina cavipes, X r.

Stem 0·5-4 cm. X 1·5-3·5 mm. cylindric or flattened, generally distinct, sometimes
forked from the base, dichotomous or flattened, palmate above, hollow in old specimens,
persistently white: branches 0·5-2·5 mm. wide, often flattened 3-6 mm. wide below
the branchings, few, often unbranched, or one to two times dichotomous or palmatijid,
TEXT-FIG. 120. Clavulina cavipes, X 2.
TEXT-FIG. 121. Clavulina cavipes, X 2.

308 CLAVULINA
generally in the same plane, slightly divergent, ascending, lax, never bushy, smooth or
becoming rugulose, the tips acute, subulate or subfiliform or subcristate-palmate, some
times the branches set with short spike-like branchlets: flesh somewhat firm, but easily
snapping: without smell.
On the ground in forest or on very rotten wood: Malaya (Pahang, Tembeling, Sing.
Field, No. 24853, 26.5.31, lowland: Fraser's Hill, 16.5.30, 1,300 m. alt.).
Spores 8-10 X 7-8· 5 µ, white, smooth, subglobose pip-shaped, with 1 large gutta.
Basidia 38-55 X 6-8 µ, subcylindric to subclavate, densely granular-guttulate:
sterigmata 2, 5-6 µ long.
Hymenium thickening to 150 µ (? more), generally absent from the stem, or in the
· upper third only: subhymenium filamentous, interwoven, the hyphae 3-5 µ wide:
cystidia none.
Hyphae as in C. cristata: the cells 25-200 x 3-15 µ, with clamps, shorter and
narrower near the subhymenium.
Surface of stem with narrow, 3-5 µ hyphae, with appressed or somewhat projecting
ends, and bearing sterile basidia near the apex of the stem, the hymenium beginning
rather abruptly.
Microscopically this species is indistinguishable from C. cristata, C. cinerea, C.
amethystina, and C. amethystinoides, unless in the longer sterile portion of the stem.
Therefore it might be reduced to C. cristata, in the wide sense, along with the other
three species. However, such a procedure would be confusing at present. It seems
that all five species can be distinguished macroscopically from their form and colour.
C. cavipes has white fruit-bodies, the branches of which become suffused pink or
vinaceous. Thus it differs from the other four species. In shape it is distinguished
by its sparse palmatifid branching, generally in one plane, the narrow outline of its
fruit-bodies, and the hollow stem of old specimens. In form it comes nearest to
C. amethystinoides, which is a truly coloured species close to C. amethystina. Tufts of
simple fruit-bodies of C. cavipes ·or of young fruit-bodies may easily be mistaken for
Clavulinopsis spiralis.
C. chondroides (B.) comb. nov.
Basinym: Clavaria chondroides B., Hook. Joum. Bot. 1, 1842, 140, t. 6, f. 3.
-2· 5 cm. high, caespitose, golden (? dried), erect: stem, very slender: branches
dichotomous, paniculate and irregularly divided above, tips obtuse: drying carti
lagi�ous.
On the ground: Surinam (leg. Hostmann).
Spores 7-8·5 X .'i'5--'7 µ, white, smooth, subglobose or ovoid.
Basidia as in Clavulina.
Hyphae -8 µ wide, clamped, thin-walled, rather loosely fibrillar.
Macroscopically this species seems identical with C. cartilaginea, though living
specimens may be differently coloured. Microscopically it differs in the presence of
clamps, as I have ascertained from a study of the type-specimen in the Kew herbarium.
C. gigartinoides may be identical.
C. cinerea (Fr.) Schroet.
Krypt. Fl. Schles. Pilze, 1888, 442.
Basinym: Clavaria cinerea Fr., Syst. Myc. 1, 1821, 468.
Synonyms: Ramaria cinerea S. F. Gray, Nat. Arr. 1, 1821, 655.
Ramaria cinerea (Fr.) Quel., Fl. Myc. 1888, 465.
Clavaria grisea Fr., Syst. Myc. 1, 1821, 468 (teste Cott. et Wakef., Bres., Coker,
Donk, Koor. et Maubl.).
Clavariella grisea (Fr.) Karst., Finl. Nat. o. Folk 37, 1882, 186.
Clavaria grisea forma petricola Bourd. et Galz., Hym. Fr. 1928, 108 (attached to
stones by an extensive white mycelium).
CLAVULINA 309
Clavariafuliginea Pers., Myc. Eur. 1, 1822, 166 (teste Pers., Coker, Dank: spores
of Persoon's specimens 7· 5-11 X 6·5-9· 5µ,, teste Coker).
Clavaria sphaerospora E. et E., Joum. Myc. 4, 1888, 62.
Text-Figs. 122, 123: Plate 4.
? Clavaria tenacella Pers., sensu Joachim, Contr. Fl. Myc. Territ. Belfort (These),
1914, 77, t. 5, f. 2: (Boudier, Obs. et Aquarelles ined., p. I08).
2·5-10 cm. high, solitary, gregarious or caespitose, generally much branched witJ,
compact branches, stout and polychotomous below, dichotomous above with blunt tips,
sometimes flattened and toothed but seldom slightly cristate, not fimbriate, occa
sionally subsimple, grey to dark cinereous, often tinged purplish, often brownish with
age.
Stem -3 x 1 cm. thick, sometimes apparently absent, white or concolorous: branches
becoming longitudinally rugulose, rather irregular and unequal: flesh firm, greyish
white: smell and taste inparticular: edible.
On the ground, in woods and fields: north temperate, Brazil, S. Austral.: common.
Spores 6-5-11 X 6-IOµ,, white, yellow, or ochraceous with age or on drying (in
herbarium-specimens), smooth, subglobose or broadly ellipsoid,. 1-guttate.
Basidia 40-70 X 5-12µ,, sterigmata 6-7·5µ,.
Hymenium thickening to 200µ,: subhymenial hyphae 3-5µ, wide: cystidia none.
Hyphae clamped, the cells 25-170 X 3-15µ, (-24µ,, occasionally, Bourd. et Galz.).
Microscopic structure as C. cristata.
Grey colour caused by the fuscous cytoplasm of the effete basidia.
f. suhcristata Bourd. et Galz.
Hym. Fr. 1928, 107.
Smaller, clearer grey: branches flattened, laciniate.
France (common).
f. sublilascens Bourd. et Galz.
Hym. Fr. 1928, 107.
Synonym: ? Clavaria crassa Britz., Hym. Si.idb. 1885, 286.
Plate 4.
More or less fuscous lilaceous: (sp. 8· 5-10 X 6· 5-8µ,: basidia 48-70 x 7· 5-9µ,:
sterigmata 7-8µ,).
France (common), England (Chippenham Fen, Cambridge, Aug. 1946).
The colour is caused by the purplish, vitreous, residual cytoplasm at the apex of
the old collapsed basidia and, thus, is darker in old thickened hymenia.
var. gracilis Rea
Tr. Br. Myc. Soc. 6, 1920, 62, t. 2, f. 4.
Plate 4.
Stem long, slender: branches long, thin, tapering, acute: p::tl0 greyish white,
branchings somewhat palmate-flattened.
Spores 9 X 8 µ,.
Gt. Britain (uncommon).
var. odorata Bourd. et Galz.
Hym. Fr. 1928, 107.
With a strong sweet fragrance of Muscari racemosum or of Mirabelle plum.
Spores 8-10 X 7-7·5µ,.
France (rare).
Concerning the synonym C. grisea, Persoon described the spores as reddish ochre,
and this description has been repeated until Cotton showed that it was mistaken (Tr.
Brit. Myc. Soc. 3, 1909, 184). It is well known that the spores of C. rugosa, C. cristata,
and, especially, C. cinerea become ochraceous after drying in the herbarium.
310 CLAVULINA
Forma sublilascens seems hardly distinguishable from C. amethystina, the colour of
which is clearer. For this reason I have referred C. crassa, described as 'violet or
lilac-grey' to f. sublilascens: it may be a colour-form of C. rugosa, however. Its
description is:
5-7 cm. high, violet or lilac-grey: stem slender, expanding upward and dividing
into several obtuse subcompressed branches: spores 8-10 X 8 µ,, white, subglobose: in
woods: Bavaria.
TExT-FIG. 122. Clavulina cinerea, X 1.
·d·, ·. Q·· ·. �--. �

.
The fruit-bodies shown in Plate 4, f. 2, as var. gracilis, are more narrowly branched
than in Rea's original illustration, but there appear to be intermediate forms. These
fruit-bodies begin as white cristate growths
which slowly become thicker, blunter, and
�. � 0-.,_ _ ..= darker with age. As the hymenium thick-
0, ·.
,. . · · . · ·' ·. · · ens it smooths over the cristations and the
TExT-FIG. 123. Clavulina cinerea, x I,ooo. effete basidia develop the grey colour which
gradually turns the branches from pale
grey to dark cinereous or dull black. Thus, young specimens must be identified with
C. cristata, old specimens with C. cinerea var. gracilis, and sparingly branched
specimens with fuliginous forms of C. rugosa. I think the following seven forms and
varieties may be only differently shaped fruit-bodies of this one variety (or, even,
species), which is intermediate between C. cristata and C. cinerea:
C. cinerea f. subcristata and var. gracilis,
C. cristata var. bicolor and var. subcinerea,
C. rugosa var. fuliginea, var. grisea and var. hercynica.
The variety is connected more closely with C. cristata by C. cristata var. incarnata,
CLAVULINA JII
which has the same form but, instead of the grey pigment, the pinkish or lilaceous
pigment, in slight degree, of C. cinerea f. sublilascens. C. tasmanica may also belong
to the alliance.
Clavaria tenacella, collected by Joachim and identified by Boudier, must have been
a sparingly branched Clavulina 'umbrino-purpurascens' with 2-spored basidia and
spores 10-11 X 9-10 µ. (according to Boudier's unpublished notes). It is clearly C.
cinerea f. sublilascens or a variety of C. rugosa. (I am indebted to M. A. Donk for this
information from Boudier's MS. in Paris.)
Compare Clavulinopsis cineroides with 4-spored basidia.
Doty mentions that the spores of C. cinerea are more greenish yellow fluorescent
under long-wave ultra-violet light than those of C. cristata.
C. cirrhata (B.) comb. nov.
Basinym: Clavaria cirrhata B., Hook. Journ. Bot. 8, 1856, 275, t. 5, f. 5: Burt, Ann.
Mo. Bot. Gdn. 7, 1922, 72, t. II, f. u3.
Synonyms: ? Clavaria pteruloides Pat., Bull. Soc. Myc. Fr. 1888, 41, t. 13, f. 3.
? Lachnocladium hamatum P. Henn., Hedw. 1904, 174.
? L. strictissimum P. Henn., ibid.
-9·5 cm. high, ochraceous white, caespitose, much branched: stem 1-1·5 cm. X
1-2 mm.: branches cylindric, strict, dichotomous or verticillate, narrow, suberect, the
tips straight or curved and acute: rather tough (Burt).
Spores 5-6 µ. wide, white, smooth, globose (Burt}.
On the ground: Brazil (Mt. Cocui).
Clavulina seems the obvious genus for this species, but it may belong to Aphelaria.
The description of C. pteruloides Pat., which appears identical, is: 4-6 cm. high,
branched, dichotomous, glabrous, with acute, 2-3-fid tips, yellowish fuscous, slightly
pink (? colour of dried specimen), the tips rather darker and somewhat compressed,
stem 5 mm. wide: spores 5-7 µ. wide, white, globose, smooth, 1 -guttate: on the ground,
Venezuela. It also suggests Clavulinopsis alcicornis.
Lachnocladium strictissimum is said to reach 15 cm. high, with very strict branches
and short stem c. 1 cm. thick: the colour is given as ferruginous (? when dried):
spores 5-6 µ., subglobose: on the ground in woods: Brazil. Bresadola makes L. hama
tum a synonym (Syll. Fung. 23, 1935, 494): it is described as 3-6 cm. high, alutaceous,
spores 6-7 x 4-7 µ., basidia 2-spored, on the ground in woods: Brazil. No difference
from C. cirrhata is apparent.
C. complanata nom. nov.

Synonym: Clavaria complanata Clel., Tr. Roy. Soc. S. Austral. 55, 1931, 159 (non
C. complanata de Bary = Typhula gyrans Fr.).
-7·5 x 12·5 cm., much wider than high: stem short, divided closely and repeatedly
into slender branches becoming flattened and expanded and ending in slender digitate
processes 6 mm. long, pale pinkish tussore becoming brownish salmon (staining paper
pinkish salmon when fresh).
Spores 5·2-7 µ., white, smooth, subglobose.
On the ground: Australia (N.S.W., Sydney).
Though its basidia are unknown, I have no doubt this is a Clavulina allied, perhaps,
with C. cartilaginea. In transferring the species I have thought it advisable to make a
new name in view of de Bary's much earlier, if little-known, synonym.
C. connata (B.) comb. nov.

Basinym: Clm•aria connata B., Hook. Journ. Bot. 8, 1856, 275.
-5 cm. high, dirty white: stems elongate, many, connate, branched above: branches
here and there connate, tips subacute: fibrillose, floccose.
On the ground in shady places: Brazil (Panure, leg. Spruce).
312 CLAVULINA
Spores 7·5-9 X 5·5-6·5µ,, white, smooth, obovoid, apiculate.
Basidia 2-spored, as in Clavulina.
Hyp hae -15µ, wide, inflating, without clamps.
The microscopic details I have added from a study of the type-specimen in the
Kew herbarium. The species is closely related to C. delicia and C. Sprucei, and all
three were described by Berkeley in the same paper. They are, perhaps, only varieties
of a tropical American species as variable as C. cristata in the north temperate region,
but it seems that no other collections are known. They are distinguished by the
absence of clamp-connexions, in which they resemble C. cartilaginea, but they differ
from this species in the looser, fihrillose texture, at least when dried. C. Sprucei and
C. delicia have very small, possibly depauperate, epiphytic fruit-bodies compared
with the terrestrial C. connata.
The distinctly elliptic spores of C. connata and C. delicia are remarkable in the genus,
and suggest unusually constructed basidia, with a gradual transition to the normal
subglobose spore in C. Sprucei. It is to be hoped that these species will be recognized
and soon described in full detail from living specimens.
C. cristata (Fr.) Schroet.
Krypt. FI. Schles. Pilze, 1888, 442.
Basinym: Clavaria cristata Fr., Syst. Myc. 1, 1821, 473: Rolland, Champ., t. 103:
Bres., le. Myc., t. 1094: Konr. et Maubl., t. 492: Coker, Clav. U.S. Can., t. 17
(immature), t. 18 (lower figure).
Synonyms: Cla.varia cristata var.fallax Fr., Syst. Myc. 1, 1821, 473 (= specimens
attacked by Rosellinia).
C. cristata var. minor Pat., Tab. An. 1884, No. 261.
C. fallax Pers., Myc. Eur. 1, 18:;,.2, 167.
C. trichopus Pers., ibid. 168.
? C. obtusiuscula Britz., Bot. Centralbl. 71, 1897, 95.
Clavariella cristata (Fr.) Karst., Finl. Nat. o. Folk 37, 1882, 188.
Clavariella trichopus (Pers.) Karst. ibid.
Ramaria cristata (Fr.) Quel., FI. Myc. 1888, 464.
Misinterpretation: Clavulina Kunzei (Fr.) Schroet., I.e., quoad descr. (non nomen
= Ramariopsis Kunzei).
Text-Figs. 27, 124, 125, 145.
2·5-8 cm. high, solitary, gregarious or caespitose, much branched, occasionally sub
simple, stem generally distinct, tips acute and generally becoming cristate, white often
becoming tinged yellowish, ochraceous or fuliginous, rarely pinkish white.
Stem 0·5-3 cm. high or practically none, greyish when parasitized by Rosellinia
Clavr,zriae: branches polychotomous or dichotomous below, the tips generally becoming
cristate-fimbriate or spicular-denticulate (20-100µ, wide) or even reduced to single
hyphae, often flattened, sometimes merely subulate: flesh rather tough, or moderately
brittle when turgid, sometimes hollow, drying whitish: smell and taste inparticular:
edible.
On the ground in fields and woods (deciduous and coniferous): temperate regions
of the whole world, common.
Spores 7-11 X 6·5-10µ,, white, smooth, subglobose, slightly thick-walled, 1-guttate.
Basidia 40-60 X 6-8µ,: sterigmata 5-7µ, long.
Hymenium thickening -200µ,: subhymenial hyphae 3-5µ, wide: cystidia none.
Hyphae clamped, colourless, the walls very slightly thickened, the cells
25-200 X 3-15µ,, shorter near the subhymenium.
Sterile base of the stem with narrow hyp hae, 3--'7µ, wide, at the surface and loosely
projecting hyphal ends 30-100µ, long, often with a few subclavate sterile basidia as
a loose and very irregular sterile hymenium passing gradually or suddenly into the
hymenium at a distance of 2-5 mm. from the base of the stem.
·
TEXT-FIG. 124. · tata, X 1.
Clavulina cris
314 CLAVULINA
var. curta Jungh.
Linnea 5, 1830, 407, t. 7, f. 2b.
3-4 cm. high, caespitose, irregular, white: branches very crowded, even concrescent,
dilated palmatifid above and setaceous-fimbriate.
(Merely a small form of typical C. cristata connecting it with var. Zappa.)
var. fimbriata Fr.
Syst. Myc. 1, 1821, 473.
Synonym: Clavaria laciniata Schaeff. ex Pers., Myc. Eur. 1, 1822, 156: Lloyd,
Myc. Notes 65, 1921, 1046, t. 1935, 1936: Coker, Clav. U.S. Can. 1923, t. 18 (upper
figure), t. 19, f. 1.
Stem elongate, distinct, laxly branched at the top: branches palmatifid with subulate,
bifid, or spinulose-cristate tips.
Spores 8-10µ., globose (Lloyd).
Europe, N. America.
(Possibly only a form developed in deep humus or shade.)
var. incarnata var. nov.
White, then pale pink or dingy flesh colour, becoming pale pinkish, brownish, or greyish
drab.
Tl!xT-FIG. 126. Clavulina cristata var. incarnata: young stages on rotting

grass-leaves, X 2.
-9·5 cm. high, gregarious or subcaespitose, much branched, at first cristate, then
the branches elongating: stem 0·2-1·5 cm. X 2-4 mm., often indistinct: branches at
first irregularly cristate-multifid, mostly adventitious, elongating and becoming
CLAVULINA 315
subulate, 1-2 mm. wide, dichotomous: flesh rather brittle, whitish: smell and taste
none: drying greyish drab.
On the ground in swampy places and on rotting stems and leaves of grasses
(Molinia): England (Chippenham Fen, Cambridge, 30.8.46, 12.9.46).
Spores 8-10·5 X 7-9µ, or 7-9 X 6-7·5 µ: basidia 35-50 X 6-7µ, not becoming
secondarily septate: sterigmata 6-8 µ long.
TExT-FrG. 127. Clavulina cristata var. incarnata:

cystidia in fruit-bodies developed on dead grass
leaves in marshy places, X 500.
Cystidia developed in fruit-bodies in very wet places (under tussocks of Molinia),

projecting to 100 µ, 5-10 µ wide below, subventricose, thin-walled, smooth, colourless,
vacuolate except the cylindric, blunt, or subcapitate apex 2-4µ wide, or 4-8µ wide
when capitate, excrescent with the hymenium or becoming immersed, sometimes
1-2 septate with clamps.
In form and colour (when old) this resembles C. ciuerea var. gracilis, as illustrated
by Rea, but it certainly belongs to the cristata-group. Fruit-bodies developed in
rotting grass-remains in the fen have lax, more elongate, and flaccid branches com
pared with those on firmer soil, and they are minutely hispid with ill-defined cystidia
which are absent from fruit-bodies in drier, more normal situations. Possibly the
variety may be distinguishable as a species.
var. lappa Karst.
Hattsv. 2, 1882, 168.
Synonym:? Clavaria alpina Saut., 1 Fl. 24, 1841: Sacc., Syll. 6, 1888, 706.
1
-1·5 cm. high, white, watery, slightly viscid: stem -'7 mm. long; branches subumbellate,
dilated above, obtuse, cristate: near Gaisstein (central Europe).
CLAVULINA
Text-Fig. 128.
1-1·5 cm. high, caespitose, white, very small, much branched from the base:
branchlets thin, cristate, and setaceous-spinulose.
Spores 6-10 X 5-7·5 µ, (Bourd. et Galz.).
TExT-FIG. 128. Clavulina cristata var.

Zappa, x2.
Basidia 27-75 x6-7·5µ,: sterigmata 2: hyphae 2-9µ, wide: (Bourd. et Galz.).
Europe.
Subspecies Coralloides
var. coralloides var. nov.
Synonyms: Clavaria coralloides Fr., Syst. Myc. 1, 1821, 467.
C. alba Pers., Myc. Eur. 1, 1822, 161: (non C. alba Pers., ibid. 175 = Clavaria
vermicularis: non C. alba sensu Bourd. et Galz.= Clavulina gallica: non C. alba
Lloyd= Lentaria mucida).
Ramaria coralloides (Fr.) Quel., FI. Myc. 1888, 465.
Ramaria alba (Pers.) Quel., Ass. Fr. 22, 1893, 488.
5-10 cm. high, white, repeatedly and irregularly much branched: branchlets very
many: generally larger and more brittle than typical C. cristata and not so cristate, if
at all.
Europe.
var. flexuosa Jungh.
Linnea 5, 1830, 407, t. 7, f. 2a.
-12 cm. high, slender, sparsely branched, subincarnate: branches elongate, flexuous,
acute.
Europe.
var. mutans Moller
Fung. Faeri:ies 1945, 117, f. 40a.
Synonym: Clavaria mutans Burt, Ann. Mo. Bot.
Gdn. 9, 1922, 31, t. 6, f. 41.
Text-Fig. 129.
-4 X 3 cm. white, then whitish-ochre, caespitose,
compact: stem I cm. thick, composed of several connate
stems, white tomentose below, irregularly divided at
the base or shortly above into 5-6 branches: branches
somewhat flattened, 4-5 times dichotomous, short,
TEXT-FIG. 129. Clavulina cristata crowded, unequal, often divergent, axils rounded,
var. mutans, X 1 (from Moller). tips blunt, not cristate or fimbriate.
CLAVULINA 317
Among grass: Faeroes (Nolso, leg. N. Peterson 1939), U.S.A.
Spores 6-8x5-7 µ, 1-guttate.
Basidia 24-50 X 6-8µ, mostly 34-38 X 7-8 µ.
var. nivea Bourd. et Galz.
Hym. Fr. 1928, 103.
Synonym: Clavaria albida Sacc., FI. It. Crypt., fasc. 15, 1916, 1232: Bres., le.
Myc., t. 1097: (non Pk., 1887 = Ramaria obtusissima).
4-8 cm. high, white to pallid cream, irregularly branched, di- or polychotomous,
mostly irregularly palmately divided, flattened 0·5-1 cm. wide just below the points of
branching, with simple or bifid, blunt or subulate tips, not cristate: branches lax, rather
numerous: stem distinct or branched from the base.
Spores 7-9 X 6-7·5µ (Coker, Bres., Bourd.).
Basidia 35-45 X 5-6µ (Bres., Bourd.).
Europe, China.
var. subrugosa var. nov.
Synonym: Clavaria grossa auctt. pr. p.: (alt. p. = Clavulina rugosa var. alcyonaria).
TEXT-FIG. 130. Clavulina cristata var. subrugosa, X 2 (coll. from

Callander, Scotland, Sept. 1933: sp. 8-10 X 7-8·5 µ: subhymenial
hyphae 3-5 J.L wide).
-7 cm. high, white, sparingly branched, drying dingy ochraceous (like C. rugosa):
stem 2-4 cm. X 3-5 mm.: branches few, smooth, thick, short, blunt, simple or once
divided, not cristate: solitary or caespitose.
Spores 7-1ox6-8·5µ.
Europe.
Subspecies Cinerascens
var. bicolor Donk
Rev. Nieder!. Homobas. Aphyll. 2, 1933, 19 (ut forma): Coker, Clav. U.S. Can.
1923, t. 9, f. 4 (ut C. cristata).
318 CLAVULINA
Stem slender, 6-15 X 1-3 mm. whitish, yellowish, or isabelline: branches smoky
grey.
Holland, U.S.A.
var. subcinerea Donk

Rev. Nieder!. Homobas. Aphyll. 2, 1933, 19 (ut forma).
Synonym: ? CZavaria comosa Pat., Bull. Soc. Myc. Fr. 22, 1906, 196.1
Seldom crested, dirty white, yellowish, or grey, blackish grey with age or on drying,
brownish where bruised.
Europe (Holland, common), ? Algiers (C. comosa).
KEY TO THE VARIETIES OF C. CRISTATA

Persistently white, yellowish or pinkish, not fuliginous
Branches more or less palmatifid above and becoming cristate,
fimbriate-dentate, or fimbriate-setose at the tips Eu-CRISTATA
1-1·5 cm. high, much branched, with slender branches and
cristate or setaceous spinulose tips . var. Zappa
3-4 cm. high, branches very crowded, dilated palmatifid
above with setaceous fimbriate tips var. curta
Generally larger
Stem elcmgate, distinct, 3-6 cm. long, laxly branched above var. fimbriata
Stem often short or none, generally bushy
White or becoming cream C. cristata typica
Pinkish, flesh colour becoming greyish drab var. incarnata
Branches irregular 2-3-chotomous above, sometimes palmatifid,
not or little cristate CoRALLOIDES
Frequently or much branched
Branches many, crowded, elongate: generally robust,
5-10 cm. high . .var. coralloides
Branches many, short, dichotomous: stems caespitose:
-4 cm. high var. mutans
Branches rather lax, few or many, mostly irregularly pal-
mately divided with simple tips . var. nivea
Branches few
-7 cm. high; branches short, blunt, simple, or once divided var. subrugosa
-12 cm. high, branches elongate, flexuous, acute var. fiexuosa
Branches becoming grey or fuliginous C1NERASCENS
Stem persistently white or yellowish: slender var. bicoZor
Stem and branches becoming blackish grey, brownish where
bruised var. subcinerea
This common species is extremely variable. Some of the varieties may be no more
than fruit-bodies developed in exposed or impoverished places (as var. Zappa, curta,
mutans) or in deep shady humus (var. fimbriata, fiexuosa, nivea)·, but others seem to
' C. comosa Pat.-greyish yellow, -8 X 5-8 cm.: stem 2 X I cm., cylindric, erect, white, then
yellowish, sulcate: branches many, narrow, cylindric, much congested, entire or incised at. the
tips: flesh fibrous, fleshy, firm: spores 9-10 X 6-8 µ., white, ovoid-globose, smooth, 1-guttate:
basidia with 2 elongate sterigmata: terrestrial: Algeria.
CLAVULINA 319
have peculiar branching. Between var. Lappa and typical C. cristata there is every
graduation through var. curta.
Var. coralloides is often maintained as a species, following Quelet, Rea, Bourdot and
Galzin, but Konrad and Maublanc say that wherever they have found it, there are
also transitions to typical C. cristata. Hence they agree with Cotton and Wakefield in
reducing it as a synonym. Bourdot and Galzin give the spores of C. coralloides as
6-8 X 4-5µ, which is smaller than usual in C. cristata and suggests a transition to
C. gallica (sp. 4·5-5 x 4µ), which are the smallest in the genus. I have retained var.
coralloides to indicate the large, not or scarcely cristate, much-branched state of
C. cristata which may have been its ancestral non-attenuate form. It has frequently
been mistaken for Ramariopsis Kunzei, the villous stem and main branches of which
should distinguish it in the field. Coker gives the spores of specimens of C. coral
loides in Persoon's herbarium as 10·5-13 X 4-4·8µ and slightly rough; they suggest a
Ramaria. Var. coralloides is also said to be brittle, compared with the typical state, but
the feature may refer to misidentified specimens of R. Kunzei.
Var. nivea has large, lax, non-cristate fruit-bodies with more or less open, palmate
branching, as if grown in deep shade. Coker, Imai, Konrad, and Maublanc reduce it
to C. cristata but, as a growth-form, it is recognizable and needs further inquiry.
Saccardo suggests that Clavaria pruinella Ces. may belong here: it was described with
ochraceous spore,; and I have referred it to Ramaria stricta.
Var. mutans is Moller's interpretation of Burt's species, which Coker reduces to
C. cristata s. lat. It is to var. coralloides as var. curta is to typical C. cristata: that is, a
stunted form. l n the field it may easily be mistaken for Ramariopsis Kunzei var.
deformis.
Var. subrugosa differs from C. rugosa var. alcyonaria only in its slightly smaller
spores, which are typical of C. cristata. It is almost exactly intermediate between the
two species. Typical specimens, which I found in Scotland (Callander, Sept. 1'933),
had spores 8-10 X 7-8·5µ and subhymenial hyphae 3-5µ wide. Coker describes the
spores of specimens of Clavaria grossa from Bresadola's herbarium as 7-9 x 6-8µ:
hence I refer them also to this variety.
Var. subcinerea differs from C. cinerea, according to Donk, in being whitish at first,
in its small size, less compact habit, and more slender stem, and branches which become
little, if at all, longitudinally rugulose. He says that transitions to typical C. cristata
are common. Var. bicolor is very similar, but still more slender with whitish stem.
One or other, or both, may be var. fuligineo-cinerascens Bourd. et Galz., merely
described as 'passing into C. cinerea' (Hym. Fr. 1928, 103). C. cinerea var. gracili; is
probably identical.
Var. minor Pat. appears to be merely the young state of normal, if depauperate,
fruit-bodies. Clavulina Bessonii may be an even more depauperate state.
C. decipiens sp. nov.

Text-Figs. 131-4.
-5 cm. high, generally more or less caespitose, rarely solitary, with rath"er few, slender,
erect, more or less fastigiate branches, pale cinnamon, dirty fawn, or cinnamon-drab,
with acute, and often filiform white tips, fibrous-tough.
Stem 2-20 X 0·8-1·5 mm., often ill defined, cylindric, smooth or slightly rugulose,
often slightly strigose-spiculose at the base, sometimes connate, soon dividing into the
slender branches, co,ncolorous: branches 0·5-1 mm. wide below, narrower upwards,
2-5-chotomous below, 2-3-chotomous above, branched more or less in alternate
planes, cylindric but slightly flattened below the narrowly palmate or spathulate dichoto
mous branchings, often with uneven outline and often connate, slightly divergent, sub
fastigiate, never numerous (though often apparently so from the caespitose habit),
matt and sterile on the uppersides, the hymenium waxy and even, ultimately tapering to
TExT-FIG. 131. Clavulina decipiens: (A, rudimentary primordia on a living fern-petiole more
or less covered by the brown myceliwn): x 2.
TEXT-FIG. 132. Clavulina decipiens, x 2.

TEXT-FIG. 133. Clavulina decipiens: thickening hymenium (left, top and bottom): young
hymenium (bottom right): sterile surface of a branch (top right): medullary hyphae with
thickened brownish walls: X 1,000.
5II9 y
322 CLAVULINA
subulate or shortly filiform tips 20-200 µ, wide, not cristate: flesh distinctly fibrous and
tough, concolorous: without smell.
On the ground or on rotten wood in the forest, with a thin, brownish felted mycelium
shortly spreading from the base of the fruit-body: Malaya (Pahang, Tembeling,
10.11.30, s.n.: 15.11.30, Sing. Field, No. 24177).
TExT-FIG. 134. Clavulina decipiens: medullary hyphae and sterile surface of the stem:
the hyphae with thickened brownish walls: X r , ooo .
Spores 8-10 x 7-9 µ, (? colour when fresh), smooth, subglobose or broadly pip
shaped, with slightly but distinctly thickened pale brownish walls o· 5 µ, thick, with I large
gutta.
Basidia 22-40 x 5·5-7 µ,, subcylindric, with granular oleaginous contents, becoming
secondarily septate about two-thirds from the base, projecting 12-20 µ,: sterigmata 2,
3·5-5 µ, long, slightly curved or nearly straight.
Hymenium c. 30 µ, thick at first, thickening to 200 µ, with few and scattered fertile
basidia at a time, the old basidia soon collapsing, included spores frequent, absent from
the stem and uppersides of the branches: subhymenium with slightly inflated and
often almost pseudoparenchymatous cells 8-25 x 4-8(-10) µ,, with colourless and thin
or very slightly thickened brownish walls: cystidia none.
CLAVULINA 323
Hyphae of the flesh monomitic, clamped, little inflated, 3--7(-8)µ wide, the cells
30-250µ long, and generally of very uneven width so as to'appear clumsy, sometimes
swollen and trumpet-like at the septa, with distinctly thickened, pale brown, smooth
walls, colourless and thin-walled only at the growing tips, short-celled next the sub
hymenium, not secondarily septate.
Sterile hymenium on the stem and uppersides of the branches constructed like the
fertile hymenium but the terminal cells 8-30 X 5-12µ, the subterminal cells -20µ
wide, more or less clavate and thick-walled, at first one cell thick, but gradually
thickening by outgrowth to 70µ, the cells pointing obliquely upward or perpendicular,
either in a compact palisade or rather loose.
Stem composed of longitudinal hyphae, the central ones paler than the peripheral
but not differing in other respects, becoming corticated by the paler sterile hymenium,
and then old stems appearing, in t.s., to have a darker core.
Growing tips composed of a fascicle of colourless thin-walled hyphae 2-4µ wide,
the subterminal cells 25-80 µ long, and soon becoming thick-walled; the hymenium
developing Sooµ -3 mm. behind the tip and thickening gradually.
In the colour and texture of the fruit-body and in the central core of the medulla,
this species superficially resembles Lachnocladium. It would be difficult to classify
without C. ornatipes which connects it with typical Clavulinas. It appears as one of
the most degenerate species in that the hyphae scarcely inflate and become thick
walled, so causing the slender branches and tough texture. The colour of the hyphal
walls clearly extends to the slightly thickened spore-wall. The basidia are typical of
Clavulina, though the sterigmata are nearly straight.
C. delicia (B.) comb. nov.

Basinym: Clavaria delicia B., Hook. Joum. Bot. 8, 1856, 274: Burt, Ann. Mo. Bot.
Gdns. 7, 1922, 70, t. 11, f. 110.
Synonym: Lachnocladium delicia (B.) Cke., Grev. 20, 1891, 10._
-1 · 5 cm. high, caespitose, ochraceous (dried), forming delicate tree-like tufts: stem
short, cylindric, subtomentose at the base, arising from a white, downy, membranous
disk, forked two to three times, some of the branches spreading, tips very acute.
On dead leaves and twigs: Brazil (Spruce 161).
Spores 9· 5-11 ·ox 5-6µ, white, smooth, oblong ellipsoid, blunt (4-5µ, globose;
Burt) (c. 10-12 x 5-6µ; Wakef. in herb. Kew.).
Basidia 2-spored, becoming septate, as in Clavulina.
Hyphae 3-12µ wide, without clamps (not even on the basidia), the walls firm and
thickened (-1µ) in the lower part of the fruit-body, often rather short-celled (? secondarily
septate), loosely fibrillar.
Hymenium unilateral, the upperside of the branches sterile and minutely vinous.
I have added the microscopical details from an examination of the type-specimens
in the Kew herbarium. The fruit-bodies are very small, slender, and delicate, and
resemble depauperate states of Lentaria byssiseda. The species is related to C. connata
and C. Sprucei (p. 69).
C. floridana (Singer) comb. nov.

Basinym: Clavaria fioridana Singer, Myc. 37, 1945, 425.
5-6·5(-8) cm. high, 2-3(-5) mm. wide, simple or forked below, more often above,
rarely almost cristate, very rarely with 2-6 branches, often compressed or canaliculate,
acute, caespitose, cinereous: stem narrower, yellowish, white below, sterile: flesh white,
rather fragile, solid, without smell, taste mild: dried fruit-bodies dark cinereous with
yellow or ochraceous stem.
On the ground in the forest, occasionally on small rotten sticks and leaves, often
very densely fasciculate as a grey turf: Florida (Matheson Hammock, Dade Co., Sept.
Oct.: Singer, F733, Rei. Farlow. No. 976).
324 CLAVULINA
Spores 7-8·5 X 5·8-7·5µ, white, smooth, subglobose, 1-guttate, non-amyloid.
Basidia 3 8-42 X 6·7-7·5µ: sterigmata 2, curved.
Hymenium thickening: subhymenium with interwoven uninflated hyphae 2-5 µ,
wide, without clamps: cystidia none.
Hyphae 2-15 µ wide, thin-walled, frequently secondarily septate, cells variable in
length 20-150µ or more, without clamps (not even on the basidia).
Sterile surface of the stem with simple appressed hyphae.
Pigment in the hyphal walls, olive, turning honey-brown,in alkali.
In describing this species Singer suggested that it was a Clavulina. I have been
able to examine part of the type-collection in the Kew herbarium and am satisfied
that it belongs to this genus, though the basidia, in other respects typical, do not
appear -to become secondarily septate: the detail, however, is .difficult to make out
from dried material.
In the form of the fruit-body, which is subsimple, the species resembles C. amethy
stinoides and C. cavipes, but the hyphal characters (secondarily septate without clamps)
suggest affinity with C. cartilaginea. and C. ornatipes, perhaps also with C. connata.
Singer suggests it is allied with Clavaria cinereo-atra of Brazil, which may well be ::i
Clavulina.
C. gallica nom. nov.

Synonym: Clavaria alba sensu Bourd. et Galz., Bull. Soc. Myc. Fr. 26, 1910, 211:
Hym. Fr. 1928, 102 (non C. alba Pers. = Clavulina cristata var. coralloides).
4-6 cm. high, much branched,, white: stem thick, short: branches crowded, slender,
flexuous, curved, blunt, or bifid, very fragile when dried.
Spores 4 · 5-5 X 4 µ,, white, smooth, oboval.
Basidia 30-42 X 4·5-5µ, with 2 sterigmata.
Hyphae 2-15µ wide, parallel.
On the ground: France (Alpes Maritimes, and in southern France).
The subglobose spores and 2-spored basidia indicate Clavulina for this species, of
which little is known. The spores are much smaller than those of C. alba which is a
synonym of C. cristata. If C. gallica is correctly placed in Clavulina, it has the smallest
spores of the genus.
C. gigartinoides sp. nov.

Text-Figs. 60, 135, 136.
Up to 15 cm. X 9 cm., caespitose or scattered, much branched, often with irregularly
and shortly spinulose-cristate tips, pallid yellowish drab with cream-buff tips, becoming
dingy fawn-drab or slightly fuliginous: flesh pallid white or pallid cream-buff, turning
slightly pinkish on bruising or cutting, rather brittle: smell not strong, somewhat of
chocolate.
Stem t-2·5 cm.X4-15 mm., more or less buried in the ground, often short and
indistinct or appearing branched from the base, pallid whitish or pale buff-drab:
branches 1·5-3 mm. thick below, 0·5-1 mm. above, often flattened up to IO mm. wide
at the branchings, generally very numerous, interlacing and often connate in places,
ascending-erect, sometimes twisted or curved, mostly regularly di-tri-chotomous below,
and irregularly branched above, often the ends more or less monopodial with short or
long branches or shortly spinulose-cristate, often tuberculate or gnarled-looking, the
tips very acute, and even subfiliform, to blunt: hymenium pale fawn-drab, only on the
underside of the lower branches, the upper branches fertile all round: flesh rather brittle,
dry, not waxy, slightly tough, generally hollow in the larger branches on drying.
On the ground in the forest (? from dead roots): Malaya (Pahang, Sungei Cheka,
11.6.31, 15.6.31, Sing. Field, No. 24867).
Spores 7-8·5 X 6·5-7·5µ, white, globose or subglobose, smooth, thin-walled, with
a small apiculus, with I large gutta, then wholly vitreous-opalescent.
CLAVULINA 325
Basidia 38-55 X 6-7µ, subclavate, projecting up to 20µ, with granular guttulate
contents: 2 curved sterigmata 4-6µ long, forming a subcrescentic apex to the basidium.
Hymenium c. 35µ thick at first, thickening-120µ, developing 0·2-1·5 mm. from the
tips, subhymenium 20-30µ thick at first, composed of 3-5µ wide, uninflated hyphae,
not inflating on the lower branches but with cells 10-20 X 5-12µ on the upper branches:
cystidia none.
T'ExT-FIG. 135. Clavulina gigartinoides, x 1: branch-tips, x 2.
Sterile hymenium on the uppersides of the main branches 50-60µ thick, loose, with
sterile basidia often bearing 1-2 abortive sterigmata: on the stem, very loose and
gradually passing into the fertile hymenium at the top of the stem, the fertile hymenium
even composed of scattered basidia (often with 1-2 short subterminal cells and pro
jecting to 60µ).
Hyphae clamped, not secondarily septate, with thin or very slightly thickened walls,
inflating 4-12µ wide, the cells 20-120µ long and generally somewhat constricted at
the septa.
Flesh with compact longitudinal hyphae near the hymenium and looser hyphae in
. the centre: in the stem and lower parts of the branches the superficial hyphae, or those
next the subhymenium, little inflated, 3-6µ wide, and very compact, but in the upper
branches these hyphae as inflated as the more internal ones.
Colour apparently caused by the granular-guttulate cytoplasm of the basidia and
CLAVULINA
subhymenial hyphae: ? brownish cytoplasm in some of the internal hyphae: (living
specimens not examined).
This may be the problematical Merisma implexa (see Lachnocladium implexus). It
has the largest fruit-body yet known in Clavulina. Compare also C. chondroides.
TEXT-FIG. I 36. Clavulina gigartinoides, X I ,ooo.

C. gracilis sp. nov.
-6 cm. x 1-2 mm., solitary, often in small troops, not caespitose, simple, acute then
blunt, cylindric or subclavate, smooth, even, with a short, distinct, narrower opaque
stem -6 X I mm., white becoming yellowish, the tip becoming brownish: solid, firm,
waxy: without smell or taste.
On the ground in the forest: Malaya (Singapore, Bukit Timah, 27.3.43, 6.3.44).
Spores 8-10µ, wide, white, smooth, nearly or quite globose, with slightly thickened
wall, 1-guttate.
Basidia 40-60 X 8-10µ,, subclavate, becoming secondarily septate, above the middle:
sterigmata 2, 5-7 µ, long.
CLAVULINA 327
Hymenium c. 50µ, thick at first, thickening -250µ,, absent from the stem: sub
hymenium c. 50µ, thick, well-developed, the hyphae 3-5µ, wide at first, the cells
inflating 15-35 x 5-16µ,, not pseudoparenchymatous: cystidia none.
Hyphae with clamps, not secondarily septate, with slightly thickened walls (0·5µ,),
smooth, the cells 30-150 X 4-17µ,, short near the subhymenium.
This species differs from C. rugosa in its small simple fruit-bodies with distinct
slender stem, its smooth hymenium, the expansion of which is accommodated by
the inflating cells of the subhymenium, and the slightly smaller spores. Moreover, as
remarked under C. ingrata, C. rugosa is not known from the tropics. Macroscopically
the species can easily be mistaken for a simple white Clavaria or Clavulinopsis.
Compare Clavulina pampaeana.
C. humilis (Cke.) comb. nov.
Basinym: Clavaria humilis Cke., Grev. 19, 1890, 2.
-15 mm. high, gregarious, very small, white, simple or sparsely ramoso-incised,
smooth, glabrous: stem short, scarcely distinct, attenuate downward, attached by a
small villous pad of mycelium.
On rotten wood and sticks: New Zealand (Namgarosa).
Spores 8-10 X T 5-9µ,, white, smooth, ovoid-subglobose.
Basidia 2-spored, subcylindric.
Hyphae -10µ, wide (but much collapsed), clamped, loosely fibrillar.
This is a very small Clavulina, as I have discovered from the type at Kew. The
specimens may be young, for they have the appearance of an incipient small fruit
body of C. cristata about to branch: they are not delicately and regularly branched as
in C. delicia.
C. ingrata sp. nov.
Text-Fig. 137.
-9 cm. X 2-3·5 mm., sometimes flattened -6 mm. wide, solitary or in small tufts,
simple, straight or flexuous, fusiform, sometimes compressed, tapering gradually to the
acute apex, becoming obtuse, and even somewhat lobed, without distinct stem,
smooth, glabrous, becoming.hollow, white then wholly pale yellowish: flesh waxy-firm,
white: with a strong, very unpleasant bitter taste and peculiar smell, somewhat of
Trigonella foenum-graecum or Hymenogaster tener.
On the ground in t.he forest: Malaya (Johore, Gunong Panti, 8.9.29, 16.4.31).
Spores 10-12 x 9·5-10·5 µ,, white, smooth, subglobose, with slightly thickened wall,
1-guttate.
Basidia 60-75 x 7-9µ,, becoming secondarily septate, clavate, with long tapering
base, projecting 15-20µ,: sterigmata 2, rarely 1, 5-6 µ, long.
Hymenium c. 65µ, thick at first, thickening -200µ,: subhymenium becoming pseudo
parenchymatous, the cells 20-38 x 7-12µ,: cystidia none.
Hyphae with clamps, the cells 20-120 X 6-19µ,, short-celled next the subhymenium, ·
looser and with narrower longer cells in the central tissue.
Stem as a sterile base to the fruit-body, 3-10 mm. long, ill-defined, with scattered
divergent hyphal ends, some with sterile basidia at the apex.
This species is macroscopically like a simple, pale yellow C. rugosa, but can be
distinguished by the smell and taste, and by the smooth surface which does not
become rugulose because the subhymenial cells inflate and accommodate the expand
ing hymenium. For this reason I think it is more closely related to C. gigartinoides,
being a simple deviation with larger spores, peculiar smell, and uniform pale yellowish
colour. C. rugosa has not been reported from the tropics and I have not found it in
Malaya.
The Australian fungus called C. rugosa? by Cleland and Chee! (1916, p. 867) may
be identical with this or another subsimple derivation of C. gigartinoides. The smell
CLAVULINA
and small spores prevent it from being C. rugosa, of which there is no certain record
from Australia: that from Wentworth Falls, N.S.W., requires confirmation in view
of the tropical development of the genus (Proc. Linn. Soc. N.S.W. 1906, 720). I give
the description of Cleland and Cheel's fungus for the benefit of Australian mycolo
gists, who may be able to identify or describe it. 1
TuxT-FIG. 137. Clavulina ingrata (left) and C. gracilis (right): hymenium

and inflated subhymenial cells, X 1 ,ooo.
C. Leveillei (Sacc.) v. Ov.
Bull. Jard. Bot. Buit. Ser. 3, 5, 1923, 260: le. Fung. Mai. 2, 1923.
Basinym: Clavaria Leveillei Sacc., Syll. Fung. 6, 1888, 709.
1
C. rugosa? sensu Clel. et Chee!: -4 cm. high, simple or with several irregular prongs,
occasionally forked into two from near the base, blunt, generally smooth, occasionally slightly
rugose, solid, pure white or cream-white, tips often yellow-brown, somewhat mealy: smell
distinct, slightly foetid (of rotten cabbage): spores white, subglobose, 7-8· S X 7 µ.: N.S.W.
(Sydney, N. Bridge, 1916: Hawkesbury River, 1916: Milson Is!., 1912, with spores 9-11 X
7--'7' S µ., thick-walled).
CLAVULINA 329
Synonyms: Clavaria alcicornis Lev., Ann. Sci. Nat. 1846, 155: (non Zoll. et
Mor. = Clavulinopsis alcicornis).
Clavaria cristatula P. Henn., Monsunia 1, 1899, 8.
Clavaria fusco-lilacina B. sensu v. Ov., Bull. Jard. Bot. Buit. Ser. 3, 5, 1923, 262:
Ic. Fung. Mai. 2, 1923: (non B., 1856 = ? a pyrenomycete).
Text-Figs. 6A, 138-41: Plate 6.
TEXT-FIG. 138. Clavulina Leveillei, X I: branch-tips, X 5.
1·5-10 cm. high, scattered, gregarious or caespitose, branched, often sparingly,

occasionally simple, stem generally distinct, branchlets subulate or cristate, drab white
becoming pale yellowish white, pale sulphur-yellow or pale dingy bistre from the base
upward,finally often (not always) more or less vinaceous drab, vinaceous fuscous, vinaceous
Juliginous, or violet-brownish on the branches, floccose-tough, pliable.
Stem 1-6 cm. X 2-4 mm., or flattened -7 mm. wide, subcylindric or unequal and
flattened, glabrous or finely cottony at the base, persistently pale yellowish, pale dingy
bistre, or somewhat deeper yellow, ochraceous or fuscous ochraceous, polychotomous or
dichotomous, radially or palmately, into the 2-5 main branches, sometimes with one
or a few small subulate abortive branches at any leveL
Branches 2-3 mm. wide below, rather few to many, ascending, branched once or
twice, 2-4-chotomously in the same plane or alternating, then polychotomous and
330 CLAVULINA
often palmate and irregularly and minutely cristate (generally radially cristate
nmbriate), in some cases with rather stout, subulate, entire or bifid tips, blunt in old
specimens, occasionally with only a few blunt stout branches, tips persistently whitish
or pale yellow: flesh white or yellowish, unchanging, the hymenium waxy-soft and
easily separable.
On the ground in the forest, lowland, and mountain -1,500 m. alt.: Java, Malaya
(Singapore, frequent).
\
i!
TExT-FIG. 139. Clavulina Leveillei var. atricha, x 1.
Spores 8-10 X 7-<Jµ, white, smooth, subglobose, with 1 large gutta 6-7·5µ wide,
occasionally multiguttulate.
Basidia 30-50 X 7-8µ, subclavate, densely granular guttulate, the effete basidia
secondarily septate: sterigmata 2, rarely 1, curved, 4-5µ long.
Cystidia 70-300 X 10-15µ, scattered through the hymenium, never abundant,
cylindric, mostly straight, but generally curved and tapered at the base, blunt, the wall
thickened 0·5-1·5µ, generally patchily encruste</, with yellowish granules, aseptate, with
vacuolate colourless contents except at the thin-walled growing tip with dense con
tents, wholly immersed or projecting -50µ, the larger ones extending through the
thickened hymenium.
Hyphae clamped, not secondarily septate, the cells of the flesh 20-150 X 3-18µ,
becoming slightly thick-walled, not or slightly constricted at the septa.
Hymenium thickening -200 or -300µ, absent from the greater part of the stem and
the uppersides of the main oblique branches and axils, developing 150-300µ behind
the tips, old basidia and spores persistent, traversed by the continuously elongating
cystidia or some of these with limited growth and becoming immersed, and others
) TExT-FIG. 140. Clavulina Leveillei: cristate tips, x 5.
TuxT-FIG. 141. Clavulina Leveillei: thickened hymenium with cystidia

and medullary hyphae: X 500.
332 CLAVULINA
arising at various levels from the excrescent hyphae: subhymenium 10-15µ. wide at
first, with short cells z-5-5 µ. wide, filamentous, not inflating.
Flesh composed mostly of longitudinal hyphae with few interweaving hyphae, the
cells 20-70 x·8-18 µ., often in submoniliform rows, with narrower, less inflated or
uninflated hyphae with cells -150 X 3-7 µ.: the central tissue loose: the peripheral
tissue next the subhymenium compact with rather narrower, shorter-celled hyphae,
their cells 20-50 X 3-7 µ..
Stem like the flesh, with a fairly wide layer of narrow hyphae, 3-7 µ. wide, at the
surface, with loosely projecting simple ends 2-3µ. wide, projecting -50µ., thin- or
slightly thick-walled, bearing scattered sterile basidia in the upper part of the stem:
the hyphal ends not forming a regular pile and not concrescent into strigose-fibrils.
Apices of the branches composed of hyphae 1•5-2 · 5µ. wide, in a dense sheaf: the
cristate tips breaking up into short filiform tips 15-80µ. wide.
Colour: yellow colour caused by a yellowish pigment in the cytoplasm of the super
ficial hyphae, sometimes also in hyphal walls: the vinaceous, violaceous, fuliginous
purple, or brownish colour caused by a slight purplish incrustation in very thin small
smears on parts of the basidia (young or old), persisting on the old basidia in the
thickened hymenium (and hence the deeper colour of old specimens, and the absence
of purplish or vinaceous colour from immature fruit-bodies and young tips), the old
basidia and ·cystidia sometimes with brownish-yellow collapsed cytoplasm.
var. atricha var. nov.
Cystidiis deficientibus.
Ad terram in silvis: Malaya (Pahang, Tembeling, Nov. 1930: Fraser's Hill,
1,300 m. alt., 27.11.30).
Differing only in the absence of cystidia: macroscopically indistinguishable.
This is a variable fungus in colour, shape, branching, and development of the
cystidia. Young and old specimens may be so dissimilar in form and colour that they
can be connected only by observing the changes during development. C. fusco
lilacina sensu v. Ov. is certainly the old, highly coloured, state with blunt branches.
It is not impossible that C. pilosa of the western tropics is the same, though it seems
to have smaller cystidia.
The cystidia are normally large and easily discernible from the blunt projecting
ends. It seems that they arise from ordinary hyphal ends on the hymenial surface
which, instead of becoming basidia, swell into the clavate young cystidia and, then,
by continued apical growth they extend with the rest of the hymenium. However,
their bases are curved and sinuous and thus very difficult to trace, but I saw no evidence
of their origin from any unusual hyphae or cells and there are no peculiar hyphae in
the flesh which would produce them as there are in Clavicorona.
C. oporata Karst.
Med. Soc. Faun. Flor. Fenn. 16, 1889, 103: Finl. Nat. o. Folk 50, 1891, 223.
Synonym: Clavaria odorata (Karst.) Sacc., Sy!!. 9, 1891, ·248.
5-6 cm. high, much-branched, fleshy, pale alutaceous, drying cinnamon, odoriferous:
stem very short, whitish: branches numerous, upright, or arcuate, compressed, pal
mate or cristate above.
Spores 5-7µ., white, smooth, globose.
On the ground. Finland.
Nothing is known of this species besides Karsten's original. description. Unless
the brownish pigment is in the hyphal walls, as in C. ornatipes and C. decipiens, the
colour is unusual in the genus and suggests that the species may be allied with
Clavulinopsis corniculata rather than with Clavulina. I retain it in Clavulina on
Karsten's authority and because the branches are palmate-cristate. It may be
Aphelaria.
CLAVULINA 333
C. ornatipes (Pk.) comb. nov.
Basinym: Clavaria ornatipes Pk., Bull. N.Y. St. Mus. 122, 1908, 18: Coker, Clav.
U.S. Can. 1923, 67, t. 9, 15, 83: Kauffm., Pap. Mich. Ac. Sci. Arts Lett. 5, 1925, u8,
t. 2, f. 1 (photo): Imai, Tr. Sapporo Nat. Hist. Soc. 13, 1934, 386.
Synonym: Lachnocladium ornatipes (Pk.) Burt, Ann. Mo. Bot. Gdn. 9, 1922, 65,
t. II, f. 102.
Text-Figs. 24, 28-30, 142, 143: Plate 15.
TEXT-FIG. 142. Clavulina ornatipes: fruit-bodies from a Singapore collection, X J.

2·5-9 cm. high, gregarious or solitary, long-stalked, with flattened, palmate, often
Jew, branchings, rather tough, Juscous-Jerruginous or fuscous-vinaceous, with pale ochra
ceous or whitish tips, drying paler.
Stem 1-6 cm.x2-3 mm. or flattened -8 mm. wide above, cylindric or somewhat
flattened and longitudinally rugulose, dark fuscous-brown or juscous-vinaceous, strigose
hispid in the lower part or throughout and even to the bases of the main branches
with juscous-ferruginous or concolorous, mostly upward-pointing, spicular or jastigiate
fibrils, often spreading somewhat over the ground from the base of the stem: branches
1·5-2 mm. wide, few to fairly numerous, rarely simple, generally palmate and multifid
with 3-7 branches in one plane, less often simply dichotomous, flattened -6 mm. wide
below the points of branching, smooth or longitudinally rugulose, upright, sub
fastigiate, with conical acute, subacute, or shortly filiform tips becoming narrowly
spathulate before branching: hymenium dark vinaceous drab or juscous vinaceous: flesh
concolorous, rather fibrous-tough, especially in the stem: smell none: taste mild or
slowly somewhat bitter.
On the ground in humus in frondose forests and under conifers: U.S.A. (widely
distributed), Canada (Nov. Scot.), Japan (Prov. lburi, Shikotsu), Malaya (Singapore,
Bukit Timah, in lowland Dipterocarp forest, in humus under palms).
Spores 8-10·5x7·5-9·5µ (12x9µ, ovoid, or 12-15µ subglobose, Doty), white,
smooth, subglobose, the wall slightly thickened, 1-guttate or (in Malayan specimens)
multiguttulate.
334 CLAVULINA
Basidia 20-36 X 6-7·5 µ (7·5-9·5 µ wide, Coker: 45-60 X 9-II µ, Doty), projecting
up to 20 µ, subcylindric or subclavate, granular-guttulate, becoming secondarily
septate: sterigmata 2, curved, 5-7 µ long (1-2 sterigmata, rarely 3, Kauffm.).
Cystidia none.
Hymenium c. 30µ thick at first, thickening to 120µ on all sides of the branches, or
occasionally absent from a narrow longitudinal strip on the uppersides, absent from
the trunk or patchily developed near the apex, not developing until 2-6 mm. behind
the tips of the branches: subhymenium c. 30 µ thick, the cells 12-30 x 4-8 µ with thin
colourless walls or those next the medulla with slightly thickened walls, not pseudo
parenchymatous, mostly with clamps, a few without.
TEXT-FIG. 143. Clavulina ornatipes: hymenium with clamps, the medullary

hyphae without clamps: x 1,000.
Hyphae of the flesh monomitic, without clamps, sparingly secondarily septate,
rather short-celled, the cells 20-150 x 3-8 µ, in places swollen irregularly -15 µ, with
slightly, but distinctly, fhickened, pale Juscous-brown walls 0·5--0·7 µ thick; strictly
longitudinal, sparingly branched, without interweaving laterals, often somewhat
curved through inflation: the cells mostly irregularly inflated, narrow in some parts,
wider in others, often bulging on either side of the septum.
s
Growing tips composed of compact hyphae I· 5-2 · µ wide, with thin colourkss
walls, longitudinal but intertwined, the cells inflating irregularly behind the tips, and
sparsely secondarily septate, the subtenninal cells 30-50 µ long.
CLAVULINA 335
Stem with narrow hyphae, 2 4-µ, wide, at the surface, rather long-celled, forming
an outer layer thickening from 50-500µ, thick (by upgrowth of laterals) and passing
gradually into the loose medulla of inflated hyphae; the ends of the superficial hyphae
projecting or more or less appressed: in the upper part of the stem, with elongate
patches of sterile hymenium as a loose palisade 30-50µ, thick, generally compact, and
becoming fertile at the apex of the stem.
Strigose fibrils on the stem 1-4 mm. long, or rudimentary, composed of fascicles
of longitudinal hyphae 2-3µ, wide, like the growing points, the large ones with the
central hyphae slightly inflated, the smaller ones often reduced to microscopic
fascicles.
This species was first described from the United States where it is widely distributed.
It was then reported from Japan, and now I add the tropical record from Malaya.
The specimens which I found in Singapore differ from the American only in having
multiguttulate, instead of 1-guttate, spores: the difference may be climatic, as an
effect of higher temperature on the oil-phase in the cytoplasm. Coker implies that the
species is northerly in distribution by stating that in the southern states it occurs only
in the mountains: this is, perhaps, more a coincidence of mycological activity.
There are slight differences in spore-measurements given by most authors, but
Doty describes them as unusually large (as are the basidia in his collection):
Spore-size in µ Country Authority

8·5-9·5 X 7·5-8·2 U.S.A. Coker (N. Car., 5620)
9·3-10·5 X 8-8·5 (Virginia)
7·5-9·3 " (N.Y., Peck's type}
9-10 " Kauffman
8-10 Japan Imai
9-10·5 Malaya Corner
12 X 9 or 12-15 U.S.A. Doty
Clamp-connexions occur on some of the hyphae of the fruit-body, not on others.

Coker says tl1ere are no clamps. I have examined the Malayan specimens twice, for
this purpose, and find that the hyphae of the flesh (or medulla) have no clamps, but
that most of the subhymenial and most, but not all, of the basidia, have clamps. The
hymenium is compact and it· is difficult to be sure that a basidium has no clamp:
I saw many with them and, certainly, a few without.
The strigose fibrils on the stem resemble rudimentary attenuate primordia, com
parable with ultimate cristate branch-endings, and suggest that the species was
caespitose.
As with C. decipiens, its nearest ally in the genus, the inflation-mechanism is
degenerate.
C. pampaeana (Speg.) comb. nov.

Basinym: Clavaria pampaeana Speg., Fungi Arg. Pug. 3, 1881, No. 40: Sacc.,
Sy!!. Fung. 6, 1888, 731: (non Speg. 1899 = Ramaria pamparum).
5-8 X 1·5 mm., gregarious (2-7), very small, white, waxy-firm; simple, erect, smooth,
blunt, with a very short thick stem.
On the ground among grass and moss: Argentine (in Pampa de Tuyu, and near
Barracas de! Sur).
Spores 7-11 X 6-7µ,, white, globose or ellipsoid, 1-guttate or multiguttulate.
Coker thinks this may be Clavaria acuta, the spores of which, however, are not
1-guttate. It seems near to Clavulina gracilis, from Malaya, but with very small
fruit-bodies, cf. C. Bessonii, C. humilis.
336 CLAVULINA
C. panurensis (B.) comb. nov.
Basinym: Clavaria panurensis B., Hook. Journ. Bot. 8, 1856, z.76.
5 cm. high, orange-ochraceous (? dried): stem slender, four to five times forked:
branches erect, here and there with lunate axils, becoming elongate, cylindric, acute.
On the ground: Brazil (Panure, Spruce 156).
Spores 7-9 X 6-7 µ, white, smooth, broadly ellipsoid.
Basidia z.-spored, subcylindric.
Hyphae somewhat inflated (very collapsed in the dried specimen), clamped.
I have examined the type at Kew. The species is a typical Clavulina very like
C. connata but, possibly, specifically different in the presence of clamps and the colour.
C. pilosa nom. nov.
Basinym: Clavaria pilosa Burt, Ann. Mo. Bot. Gdn. 9, 19z.z., 46, t. 8, f. 70 (non
Pers. = Clavariadelphus fistulosus).
1-z. cm. long, z. mm. thick (when dried), simple, solitary or two to three together
(huffy brown to drab when dry), compressed, thickened in the middle, obtuse.
Spores 6-8 X 6-7 µ, white, smooth, subglobose.
Cystidia 6µ wide, projecting -30µ beyond the basidia, cylindric, hair-like, smooth,
hyaline.
On humus: Porto Rico (Johnston and Stevens 1453).
Though so little is known of it, this species seems undoubtedly a Clavulina near to
C. Leveillei, but simple. Compare also C. fioridana.
C. rugosa (Fr.) Schroet.
Krypt. FI. Schles. Pilze 1888, 44z..
Basinym: Clavaria rugosa Fr., Syst. Myc. 1, 18z.1, 473: Rolland, Champ., t. 103,
No. z.33: Bres., le. Myc., t. 1098: Coker, t. 16, f. 4, t. 19, f. 3 (? old, approaching var.
fuliginea).
Synonyms: Clavaria pseudofiava Britz., Hym. Sildb. 8, 1891, 14, f. 6z.: Rev. Hym.
1909, z.z.o (only a yellowish form).
C. clavaeformis Britz., ibid. (yellowish form).
C. Herveyi Pk., Rept. N.Y. St. Mus. 45, 189z., 84.
C. gigantula Britz., Bot. Centralbl. 71, 1897, 95, f. 98.
C. obtusiuscula Britz., ibid. 19, f. 97.
? Clavaria rubescens (Quel.) Sacc., Syll. 11, 1895, 136.
Ramaria rugosa S. F. Gray, Nat. Arr. 1, 18z.r, 655.
R. rugosa (Fr.) Quel., Fl. Myc. 1888, 464.
? R. rubescens Quel., Rev. Myc. 189z., 65.
4-1z. cm. high, 3-10 mm. thick, solitary, gregarious, or subcaespitose, simple or with
1-3 undivided short branches, often antler-like, cylindric, subacute, even, becoming
more or less clavate, obtuse, and longitudinally rugulose, varying rugoso-bullate, even
cerebriform, often flattened, narrowed downwards into an indistinct stem, white to
cream, rarely yellow, drying light ochre-yellow or antimony-yellow: flesh rather
tough, firm: smell and taste slight, imparticular.
On the ground in woods and pastures: north temperate, common: Australia ?
Spores 9-14 X 8-u µ, white, smooth, broadly ovoid-ellipsoi<l, r-guttate.
Basidia 40-85 x6-9·5µ: sterigmata 6-9µ long.
Hymenium thickening to 150µ: subhymenial hyphae 3-7 /,,t wide, interwoven:
cystidia none.
Hyphae clamped, inflating, becoming slightly thick-walled, the cells 40-120 X 3-13 µ,
H-connexions frequent, not secondarily septate: microscopic structure as in C.
cristata.
CLAVULINA 337
f. mitruloides Bourd. et Galz.
Hym. Fr. 1928, 102.
-7 cm. x 3-4 mm. oblong, undulate, crisped or cerebriform, 2-3 connate at the base,
soft.
Spores 9-12 X 7-10µ.
l\
l(
I
')O
\ r/
\�
'iI
i
diI
t\ I\
I
'
'I
/ I \
I
I \
\ \ (
TEXT-FIG. 144. Clavulina rugosa: typical fruit-bodies (left), var. alcyonaria (right): X 2.
[Basidia 7 5-100 X 9-12 µ, many sterile and septate as gloeocystidia (? only young
basidial branches arising deeply in the thickened hymenium).]
Under Pinus: France (rare).
var. alcyonaria var. nov.

Synonyms: Clavaria grossa auctt. pr. p. (alt. p. = Clavulina cristata var. subrugosa):
Bres., le. Myc., t. 1096 (? 4 sterigmata).
C. incrassata Chev., FI. Gen. Paris 1, 1826, 108.
C. Krombholzii Fr. pr. p. (Krombh., f. 18-20): (non vera = Ramariopsis Kunzei
var. deformis).
C. arctata Britz., Hym. Sudb. 8, 1891, 286.
? C. gregalis Britz., ibid.
5119 z
338 CLAVULINA
? C. unistirpis Britz., Hym. Si.idb. 8, 1891, 287.
Ramaria grossa Quel., FI. Myc. 1888, 464.
Text-Fig. 144.
With several long, stout, blunt branches, simple or one to three times branched them
selves, or merely shortly lobed or toothed at the blunt tips (not cristate), often caespitose.
Spores 9-12 X 7-9 µ,.
Europe.
var. canaliculata var. nov.

Synonyms: Clavaria canaliculata Fr., Syst. Myc. 1, 1821, 485.
C. canaliculata var. ingrica Weinm. 1836.
C. ingrica Weinm. (? nomen validum).
3-20 cm. high, 4-6 mm. wide, s,iow white or greyish (? parasitized), cylindric or
subclavate, smooth, t!Ven, becoming longitudinally sulcate, not rugulose, finally hollow.
Spores 8-12 X 7-9 µ,.
Europe.
var. fuliginea Fr.

Hym. Eur. 1874, 669.
Dark fuliginous.
Gt. Britain, France, Sweden.
var. grisea Fr.

Syst. Myc. 1, 1821, 474.
Caespitose, fuliginous: branches deformed, rugose, subcristate.
var. hercynica Fr.

Syst. Myc. 1, 1821, 474.
Larger, fuliginous white: branches obsolete, obtuse.
var. macrospora var. nov.

Synonym: Clavaria macrospora Britz., Hym. Si.idb. 8, 1891, 287.
Spores 12-14 x 8-10 µ,.
Gt. Britain, France, Germany.
KEY TO THE VARIETIES OF CLAVULINA RUGOSA

Sp. 12-14 X 8-10µ,: otherwise typical var. macrospora
Sp. mostly 9-12 X 7-10µ,
Branches long, stout, blunt, often divided 1-3 times var. alcyonaria
Branches 1-3, short, antler-like or none
-20 cm. high, even then longitudinally sulcate and hollow, not
rugulose or bullate-cerebriform var. canaliculata
-12 cm. high, becoming longitudinally rugulose and more or
less bullate-cerebriform, rarely hollow
Fuliginous
Fuliginous white var. hercynica
Dark fuliginous var. /uliginea
Dark fuliginous, caespitose subcristate var. grisea
White, cream, or yellowish C. rugosa typica
Undulate, crisped, or cerebriform f. mitruloides
CLAVULINA 339
The varieties of this polymorphic species are unsatisfactory. I have listed them in
order to emphasize the need for closer inquiry. Var. fuliginea and var. grisea, if not
merely fruit-bodies parasitized by Rosellinia, seem to grade into C. cinerea var. gracilis
or C. cristata var. bicolor and var. subcinerea. Var. alcyonaria also grades in C. cristata
var. subrugosa, which has the smaller spores and narrower subhymenial hyphae of
C. cristata. Var. canaliculata is the smooth, white form suggesting Clavulinopsis or
Clavaria and the one which connects with the truly simple C. gracilis and C. ingrata
of Malaya. C. rugosa has not been found in the tropics, however, and the records
from Australia must be regarded as doubtful (see C. ingrata and C. subrugosa).
Qoo·G
TExT-FIG. _ 145. Clavulina cristata (right) and C. rugosa (left), showing the difference in size
of the spores and subhymenial hyphae: x 1 ,ooo.
Concerning the synonyms, C. Herveyi is said by Coker to be the rugosa-form of his

C. cristata, though it was described as golden, subsimple, with white flesh and globose,
minutely rugose spores 7· 5 fL wide. C. gigantula (sp. 10-12 X 9-10 µ.) is referred
tentatively to C. amethystinoides by Coker, but this species has not been recognized in
Europt: and C. gigantula appears to be normal C. rugosa, or a colour form, as it was
said to be white then brownish. C. obtusiuscula seems to be a greyish violaceous form,
unless it is parasitized by Rosellinia. C. rubescens appears to be a pinkish variety; its
description is:
Simple or branched, cylindric or compressed, acuminate-obtuse, thick, sometimes
340 CLAVULINA
umbilicate, attenuate and twisted toward the base, whitish rufescent: on the ground
in woods: France. (Possibly a variety of C. rugosa: Saccardo.)
C. arctata (sp. 10-12 X 6-8 µ,), C. gregalis (sp. 12 X 8-9 µ,), and C. unistirpis (sp.
10-12 X 6-8 µ,) have been referred to C. cristata, and macroscopically would fit var.
subrugosa, but they have the larger spores of C. rugosa and, therefore, I refer them to
its var. alcyonaria.
TABLE XXIII. SPORE-SIZE IN CLAVULINA RUGOSA
Onefruit-body: Tring, England, 22.9. 2 5 Onefruit-body: Cambridge, England, r5.ro. 2 5
I
Spore-sizein J-L E' Number of records Spore-sizein J-L E' Number of records
9·0X7·5 1·2 l
9·0X7·0 1·3 l 9·0X8·o 1• l 2
10·0X7·5 1·3 l 9·5X8·3 1 ·1 l
1o·ox8·o 1'3 3 9·5X8·5 I•I I
10·0X 8·3 1·2 2 1o·ox7·5 1·3 2
10·0X8·5 1·2 2 10·0X8·o 1·3 4
10·0X9·0 I•1 4 10·0X8·5 1·2 4
10·0X9·5 l •I I 10·0X9·0 I •1 2
10·3X10·0 1·0 I 10·5X8·5 1·2 3
10·5X8·o 1·3 I 10·5X9·0 1·2 I
10·5X8·3 1·3 l 11·0X8·5 1·3 3
10·5X8·5 1·2 2 11·0X9·0 1·2 2
10·5X9·0 1·2 2 u·ox9·5 I·2 I
10·5X 9·5 I·1 I I I·oX10·0 1·1 2
11·0X8·o 1·4 l n·5x8·5 1·4 l
11·0X8·3 I ·3 l 11•5X9·0 r·3 3
11·ox8·5 1·3 2 11·5X9·5 1'2 2
u·ox9·0 1·2 3 11·5X9·7 1·2 l
II·oX9·5 1'2 l 11·5X10·0 1·2 5
u·ox10·0 I• l l 11·5X10·3 I•I I
11·5x8·5 1·4 2 12·0 X9·5 1·3 2
II·5X9·0 I •3 5 12·0 X10·0 l ·2 6
11·5X 9·3 1·2 I 12·ox10·5 l •I I
11·5x9·5 l ·2 I 12·0X11·0 l·l l
JI·5X 9·7 1·2 3 12·5X10·0 l ·3 3
II·5X10·0 I·2 I I2·5 X10·5 1·2 l
12·0X8·5 1·4 4 12·5X11·0 l·l l
12·0X8·7 1·4 2 13·0 X10·0 l ·3 2
12·0 X9·0 1·3 l 13·0X11·5 l·l l
12·0 X9·3 l ·3 l 13·5X 10·0 1·4 2
12·ox9··5 1·3 3 13·5X12·0 I· I l
12·0X9·7 1·2 4
12·0X10·0 1·2 4
12·5x9·5 1·3 l 9-13·5x7·5-12 1·1-1·4 63 (total)
12·5X10·0 1·3 I
13·ox8·5
---
1·5 I
9-9·5µ long 1·14 Av. of 5 spores
9-13X7-10 1-1·5 66 (total) 10·0 " 1·25 " 12 "
9-10µ long I'18 Av. of 15 spores 10·5 " 1·22 " 4 "
10·5 " I'21 " 7 " l 1·0 " 1·21 " 8 "
I1·0 " r-23 " 9 " l I. 5 " 1·20 " 13 "
I I·5 " 1'25 " 13 " 12·0 " 1·20 " 10 "
12·0
12·5-13 "
" 1·30 "
"
19 "
"
12·5 "
13-13·5 "
1·21
I " 5 "
" 6 "
I
1·37 3 1·25
l 1·2X 9·0 1·25 " 66 " 11·2x9·3 l ·22 " 63 "
1
E = length -,- width.
CLAVULINA 341
C. Sprucei (B.) comb. nov.
Basinym: Clavaria Sprucei B., Hook. Joum. Bot. 8, 1856, 276.
3·5-4 cm. high, gregarious, alutaceous (?dried): stem slender, glabrous, arising from
a small, membranous, byssoid mycelial patch, three to four times branched: branches
cylindric, substrict, obtuse.
On trunks: Brazil (Panure, Spruce, No. 26).
Spores 8-1o x 7-8 µ,, white, smooth, subglobose.
Basidia as in Clavulina, with 2 sterigmata.
Hyphae -12 µ, (? -20µ,) wide, inflating, thin-walled, without clamps, possibly
secondarily septate.
The microscopic details I have added from the type-specimen in the Kew herbarium.
It is related to C. connata and C. delicia (p. 312).
C. subrugosa (Clel.) comb. nov.

Basinym: Clavaria subrugosa Clel., Tr. Roy. Soc. S. Austr. 59, 1935, 220.
-5 cm. high, simple or irregularly branched, white, subcinereous-white, pale greyish
brown, or somewhat cartridge-buff, slender below, somewhat expanded above or even
flattened, irregularly rugulose, with filiform or flattened branches, solid.
Spores 5·5-7·5µ,, white, smooth, subglobose.
On the ground, gregarious: S. Australia (Mt. Lofty).
Without further microscopical details, the position of this fungus must remain
uncertain. The colour and flattened branching indicate Clavulina, but the spores are
rather small. Fawcett regards it as a synonym of C. cristata sensu Coker.
C. tasmanica (B.) comb. nov.

Basinym: Clavaria tasmanica B., Grev. 20, 1891, 10.
-4 cm.high, simple, solitary or two to three together, clavate,fuliginous: stem slender,
paler, subflexuous, expanded into a white floccose mycelium.
On tree-ferns, wood, &c.: Tasmania.
Spores 7·5-9·5 X 6·5-8 µ,, white, subglobose to broadly pyriform, apiculate.
Basidia with 2 sterigmata, subcylindric, evidently withfuliginous sap.
Hyphae -10µ, wide, fibrillose, clamped.
I have examined the type at Kew and found that it is a typical Clavulina. Possibly
Clavaria cinereo-atra and C. Greletii are related.
C. Vanderystii (Bres.) comb. nov.
Basinym: Lachnocladium Vanderystii Bres., Bull.Jard. Bot. Brux. 4, 1913, 25.
6-8 X 3-6 cm. caespitose: stem 8-15 x 2-4 mm., congested, soon branched: branches
repeatedly 2-4-fid, subcompressed, rugulose, glabrous on one side, pubescent on the
other, lurid alutaceous (? the colour when dry), axils angular: branchlets fuscous
rufescent (? dried) with acute 2-4-fid tips.
On the ground: Belgian Congo (Bambuta).
Spores 7-8 X 6-7 µ,, white, smooth, globose or subglobose.
Basidia 20-25 x 6-7 µ,, clavate.
Hyphae 4-12 µ, wide, thin-walled, irregular, colourless.
This must be a Clavulina and n-::ar, perhaps, to C. gigartinoides, but I doubt if it is
recognizable. The short length 01 the basidia may refer only to the part distal to the
secondary septum.
C. vinaceo-cervina (Clel.) comb. nov.
Basinym: Clavaria vinaceo-cervina Clel., Tr. Roy. Soc. S. Austr. 55, 1931, 158:
Toadstools ... of S.Austr. 2, 1935, 266, t. 8, f. 4_.
-5 cm. high, erect or slightly spreading, very irregularly branchedfrom a short trunk,
342 CLAVULINA
sometimes with only a few branches and prong-like divisions, sometimes with a
number of small branches: branchlets short, prong-like, mostly blunt, sometimes aC11te
and thorn-like or subulate or digitate, often fastigiate: branches of ten irregularly flattened
and rugose, usually slender, sometimes stouter and more knobby, vinaceous fawn to
fawn colour, vinaceous pink at the tips, the stem vinaceous fawn and pallid at the base.
Spores 7·5-10 X 6·5-8·8µ (? white), smooth, subglobose.
On the ground under trees: S. Australia.
This may be the same as C. cartilaginea, but it needs more detailed study: Fawcett
regards it as a synonym of C. cristata sensu Coker.
CLAVULINOPSIS van Ov.

Bull. Jard. Bot. Buit., Ser. 3, 5, 1923, 278.
Synonym: Cornicularia Bon., Handb. Myk. 1851, 166 (non Hoffrn. 1789).
Clavaria pr. p.
Fruit-bodies simple or branched, clavarioid, variously coloured, often white, yellow,
orange, or red, rarely brown, grey, or purple: stem sterile, more or less distinct:
branching radial, alternating: fie.shy-waxy to rather tough, never so brittle as in Clavaria,
gelatinous in a few species.
Terrestrial, solitary, gregarious or caespitose, exceptionally lignicolous (or amenti
colous, C. pusilla): about 60 spp.: throughout the world.
�
\�
A (r:i
��
TEx<r-FIG. 146. Spores of the orange-yellow species of Clavulinopsis: a, C. luteo-tenerrima
(normal): b, C. luteo-tenerr.ima (with pronounced apiculus): c, C. Archeri: d, C. pulchra:
e, C. luteo-alba var. latispora: f, C. corniculata: g, C. alcicornis: h, C. helvola: i, C.jusiformis:
X 1,000.
Spores white, or tinged yellow from the oil-drop, smooth or, in a few species, coarsely
echinulate, globose, pip-shaped or ellipsoid, usually 1-guttate, sometimes multi
guttulate.
CLAVULINOPSIS 343
Basidia clavate, not becoming secondarily septate: sterigmata mostly 4, occasionally
2-3, rarely 4-6 (C. hastula).
Cystidia absent (except C. luticola).
Hymenium _generally thickening, absent trom the stem, eventually covering the
apex of the fruit-body: subhymenial hyphae short-celled, generally inflating.
Hyphae monomitic, clamped, mostly inflating, not secondarily septate, often with
narrow interweaving hyphae among the inflated longitudinal hyphae, the walls thin
or very slightly thickened.
Type-species: C. miniata.
This genus differs from Clavaria in the hyphal construction of the fruit-body. It
has the usual construction for a basidiomycete, so that it becomes the largest and
most representative group of white-spored Clavarias. Its nearest ally is Ramariopsis
with minutely echinulate spores; possibly the subgenus Acularia of Clavulinopsis,
including C. helvola ( = C. dissipabilis) and C. asterospora, bridges the gap. Lentaria
is typically lignicolous with narrowly ellipsoid aguttate spores and thick-walled hyphae.
Nevertheless, there are anomalous species of uncertain position which I have placed
in Clavulinopsis for convenience (see p. 81).
Macroscopically species of Clavulinopsis can be distinguished from those of
Clavaria by the less brittle texture, the hyphae not being secondarily septate: it seems,
also, that most of the yellow or orange Clavarias belong to Clavulinopsis. For this
reason I have had to reconsider the nomenclature of the yellow European species,
which the microscopic studies of this century have, I hope, enabled one at last to
decide.
Many additions are to be expected from tropical and subtropical countries.
CLAVARIA INAEQUALIS FR., AS A NOMEN AMBIGUUM

The interpretation of C. inaequalis Fr. is one of the knottiest problems of myco
logical systematy. It has led to much confusion over the names of the non-caespitose
Clavarias of Europe with simple, yellow, or orange fruit-bodies, which does not seem
yet to have been settled. I have, therefore, gone into the matter carefully in view of the
mass of information which is now available and because it is necessary to raise a new
and very important point, that is the identity of C. inaequalis Mull. (in FI. Dan. 1780,
t. 836, f. 1) from which the specific epithet is derived.
Three yellow or orange, simple, gregarious or subcaespitose species are now recog
nized in Europe, chiefly on microscopic characters, namely C. dissipabilis Britz.
(globose, spiny spores), C. luteo-alba Rea (smooth, ellipsoid spores), and C. pulchra
Pk. (smooth, subglobose, strongly apiculate spores). It appears, also, that there may
be another species with larger spores than C. luteo-alba, or at least a variety of it, and
that C. Jusiformis Fr. may have a subcaespitose variety comparable with the tropical
C. amoena. Macroscopically they are so much alike that one may doubt whether they
can be identified with certainty except from their spores. It is impossible, therefore,
to be certain to which species the earlier descriptions of Fries and Persoon are to be
referred, and, in the absence of type-specimens, the evidence is circumstantial.
There have been two interpretations of C. inaequalis, in addition to that of Fries
(Syst. Myc. 1, 2, 1830, 231, which appears also to have been overlooked, vide infra).
I will call them Karsten's and Cotton's.
In 1882, Karsten ascribed a smooth ellipsoid spore, 10 x s µ, to C. inaequalis. Con
sequently, Britzelmayr described the very common species with globose, spiny spores
as C. dissipabilis (1885). Actually, Cooke had previously described the spores of C.
inaequalis in 1871 (Handb. Brit. Fungi _1, _337) as smooth, ellipsoid, 11 ·s X 6 µ, but to
what extent this influenced subsequent mycologists is not known (and such a fungus
has not been found again in Gt. Britain). C. inaequalis has been accepted in this sense,
with smooth, ellipsoid spores, by Quelet, Boudier, Patouillard, Massee, Saccardo,
344 CLAVULINOPSIS
Bresadola, Ricken, Killermann, Rea, Bourdot and Galzin, and Konrad and Maublanc,
who refer to the spinous-spored species as C. dissipabilis or by its synonym C. similis
Boud. et Pat.
In 1906 Cotton showed that the spinous-spored species was probably by far the
commonest in Europe, that it was the one represented in European Exsiccati under the
name C. inaequalis, and that Karsten's own collection (No. 439) had this spore and
not that which he had ascribed to it. Cotton, therefore, gave the name C. inaequalis
to the spiny-spored species, reduced C. dissipabilis as a synonym, and referred the
species with smooth, ellipsoid spores to C. luteo-alba Rea, which had just been pub
lished (1903). With Miss Wakefield he re-affirmed this view in 1919, and it is accepted
by Coker and Donk as the current interpretation.
Typification of C. inaequalis Mull. ex Fr. (1821). There is said to be no type
specimen. Typification, therefore, depends on
(i) the identification of C. inaequalis Mi.ill. (FI. Dan.) from which the specific
epithet was taken;
(ii) Fries's own idea of the species, for he adds '(v.v.)' after his description;
(iii) customary use.
With regard to the first point, it is generally assumed that the illustration represents
C. dissipabilis. Reference to the plate will, however, convince anyone, I think, that
it depicts C. fusiformis: the fruit-bodies are characteristically fusiform, yellow rather
than orange, and hollow, as shown by the unmistakable section. The description states
'Clavaria inaequalis caespitosa compressa, lutea, apice acuminata', which confirms the
caespitose habit suggested by the plate and clinches the identity with C. fusiformis.
It would be strange if this common species had not been illustrated in the Flora
Danica, for it is not represented on another plate, and that Fries was uncertain about
it, too, is shown by the next consideration. The illustration certainly cannot be the
solid obtuse C. dissipabilis.
In 1821 (p. 481) Fries questioned whether C. inaequalis was different from C.
fusiformis; he wrote 'An satis distincta ?' and regarded it chiefly as a less caespitose,
more irregular form. The common C. dissipabilis was undoubtedly described by
Fries, then, as C. aurantiaca, C. helvola and, possibly, C. angustata. However, he
reviewed the problem in 1828 (Elench. 231-2) and used C. inaequalis as a general
specific name to cover C. fusiformis, C. inaequalis (FI. Dan.), C. angustata, C. auraf!
tiaca, C. helvola, and, possibly, C. rufa, which he regarded as varieties. Finally (1874),
he separated C. fusiformis, as the densely caespitose, hollow, fusiform species, from
the subcaespitose or gregarious, solid and usually blunt C. inaequalis, the variability
of which he remarked in giving the other three names as synonyms. Thus, in 1821,
Fries considered C. inaequalis as very near to C. fusiformis, if not identical: in 1828,
he considered it as covering all the simple yellow to orange European species: and in
1874, his definition refers in all likelihood to C. dissipabilis, as Cotton has shown.
Such confusion is to be expected where only the microscope can achieve certainty,
and does away with any hope of typifying C. inaequalis from the Friesian classification
of the living specimens.
With regard to customary use of the name by other mycologists, Cotton's conclusion
must be accepted that it refers to C. dissipabilis.
There are, therefore, these four possibilities:
(i) C. inaequalis sensu Mi.ill. is C. fusiformis;
(ii) C. inaequalis sensu Fr. is indeterminable;
(iii) C. inaequalis sensu auctt. is C. dissipabilis;
(iv) C. inaequalis, sensu Karst. is C. luteo-alba.
I cannot see how the identity can be decided except by an arbitrary decision.
Therefore, I propose to regard C. inaequalis as a nomen ambiguum and to use the names
C. fusiformis, C. helvola ( = C. dissipabilis), C. luteo-alba, and C. pulchra, the identity
of which is acknowledged, except for C. helvola, which I will now consider.
CLAVULINOPSIS 345
C. helvola Pers. ex Fr. (I82I) and C. dissipabilis Britz. (r885). C. helvola has been
interpreted in two ways. It has generally been considered identical with C. inaequalis
in Cotton's sense, that is with C. dissipabilis, and in support Donlc states that authentic
material of C. helvola in Persoon's herbarium has the spores of C. dissipabilis (Rev.
Niederl. Homobas. 2, 1933, 82, 84). Cotton notes, in contrast, that a Clavaria with
smooth, ellipsoid spores, 7 X 3 f-1,, has been identified as C. helvola in Europe (Tr. Brit.
Myc. Soc. 3, 1906, 165, footnote), and in this sense Coker uses the name in the
N. American flora. This smooth-spored species, however, cannot be separated from
C. luteo-alba, as I have discussed on page 375. C. helvola, therefore, on the basis of
Persoon's specimens, must be the earliest name for C. dissipabilis ( = C. inaequalis
sensu Cotton). C. fiammans B. (1875) has, also, the spores of C. dissipabilis and, on
the grounds of type-specimen, is also a prior name. Nevertheless, Britzelmayr was the
first to describe this peculiar spore and it is regrettable that his name must disappear
in synonymy. Persoon, it may be noted, evidently considered C. helvola to cover C.
angustata and C. aurantiaca, which he gave as varieties rather than species under
C. helvola (Myc. Eur. 1, 1822, 181-2), and he also remarked of C. inaequalis Fr. 'An
a Clavaria helvola satis diversa ?' As Cotton has shown, C. helvola sensu Persoon must
have covered mainly, if not entirely, C. dissipabilis.
Exercise. In Table XXIV are listed the spores which have been ascribed to C.
inaequalis, other than those of C. dissipabilis, and in the last column the species to
which they should be referred. It is noteworthy that Britzelmayr, Quelet, and Rea
(following Bourd. and Galz.) attribute the spores of C. fusiformis to C. inaequalis, thus
implying that there is a subcaespitose state of C. fusiformis identical with C. inaequalis
(FI. Dan.) The very closely allied C. amoena has such forms in the tropics and such
a 'var. inaequalis' is to be expected in Europe. Also, it may be noted, Schroeter
must be regarded as the first to discover C. pulchra in Europe: it has since been
recognized by Donlc in Holland and by J osserand in France, though previously
recorded as C. persimilis by Cotton from Great Britain.
TABLE XXIV. SMOOTH SPORES ASCRIBED TO CLAVARIA INAEQUALIS
Author Size in I' New identification

Karsten, 1882 . IOX 5 C. luteo-alba var. latispora
Bourdot, Galzin, 1927 6-9X4-6
Killermann, 1928 10X4-6 "
Schroeter, 1888 5-7X4 C. pulchra
Karsten, 1889 . 5-6X3-4 C. pulchra or C. luteo-a[ba
Saccardo, 1888 . 10-12X5-6 C. luteo-alba var. macrospora
Ricken, 1918 15X6 "
Britzelmayr, 1885 6-8 x6 C. fusiformis
QueJet, I 888 8-10, subglobose
Rea, 1922 7-9 X 6-8 "
Bourdot, Galzin, 1927 4-8X4-6 C. fusiformis, C. pulchra
(as C. aurantia)
KEY TO THE SPECIES OF CLAVULINOPSIS

Sp. spinous or verrucose: simple
Acularia A
Sp. smooth: simple or branched
Eu-Clavulinopsis
Sp. narrowly ellipsoid or allantoid: fruit-bodies very small and
slender, generally 1-2 cm. high, or less, mostly simple: tough
or gelatinous B
CLAVULINOPSIS
Sp. shortly or broadly ellipsoid, subglobose or globose: fruit
bodies medium to large, waxy or brittle, rarely rather tough
Fruit-bodies white, cream, or tinged yellow C
Fruit-bodies white, then yellow, brownish, or rufescent from
below upwards (hysterochroic) D
Fruit-bodies yellow, greenish, orange, or ochraceous •E
Fruit-bodies pink, red, or orange-red F
Fruit-bodie,s brown, umber, grey, or with purplish tips G
A. Acularia. Sp. spinous or verrucose: simple

Sp. verrucose, 8-9µ wide: -4 cm. high, white; Gt. Britain . C. tenerrima
Sp. set with spines 1-3·5µ long
Sp. pruniform: white, filiform: Europe . C. candida
Sp. subglobose
White: Europe C. asterospora
Yellow, orange: N. temp., S. Africa C. helvola
(Wood-brown: sp. minutely echinulate: U.S.A. Ramariopsis
asperulospora)
(? Sp. verruculose, 3·5-5 X2·5-4µ: -5 cm. high, simple or
sparingly branched, yellowish with brownish scurfy stem:
foetid: Europe, U.S.A. C. luteo
ochracea)
(Sp. rough, yellowish, 7-u x5-6µ: -15 x2-3 mm., spathulate,
yellowish clay: on earth in hot-house: Berlin C. spathuli
formis)
(Sp. rough, 7-8·5 X3·5µ: simple, orange: Nova Scotia Clavaria
gracillima,
p. 227)
B. Small, mostly simple, often phycophilous: sp. narrowly

ellipsoid or allantoid
Gelatinous, simple: trop.
-25 mm. high, orange-yellow, toughly gelatinous: sp. 8·5-10 x
4-5µ: basidia 4-spored: Colombia C. calocera
-7 mm. high, pale pinkish-cream, softly gelatinous, flattened:
sp. 5-7 X2·5-3µ: basidia 4-6-spored: Malaya C. hastula
(Sp. 7-7·5 X5-6µ, 3-lobed: trop. Amer. see C. fiavella
sensu Martin)
Tough: temp.
Sp. II-13 X4·2-5·5µ: -23 mm. high, simple or sparingly
branched, white: U.S.A. C. pogonati
Sp. 8-II X1·5-3µ: simple, -12 mm. high
Watery ochraceous to dull orange: phycophilous: U.S.A.,
Europe C. vernalis
.Pale cream-white: not phycophilous: U.S.A. C. constans
CLAVULINOPSIS 347
(Sp. 7-1 I x5�6 µ, yellowish, rough: spathulate, 2-4 mm. wide . C. spathuli
formis)
(Sp. 5-7 X2-3µ.: -2 cm. high, white or yellowish: phycophilous,
on rotten wood Lentaria mucida
L. coronilla)
C. White, cream, or pale yellow: sp. smooth
Sp. globose or subglobose, or (if ellipsoid) more than 4·5 µ. wide
Sp. 4·5-7·5µ. long
Branched
-6 cm. high: branches many, dichotomous, fragile: sp.
4-7 X 3-5µ.: caespitose: France, England C. dichotoma
-4 cm. high: branches often few and irregular, rather
tough: sp. 3·5-4·5(-6) x3-3·5(-4·5) µ.: Europe C. subtilis
-1 ·5 cm. high: 2-5 times dichotomous: base white villous:
sp. 5-7·5µ.: Bonin Is!. C. boninensis
-2·5 cm. high, stem arising from a small tubercular base:
sparingly branched: sp. 5-7µ.: Brazil C. Puiggarii
(-9·5 cm. high, ochraceous-white, caespitose, much
branched: sp. 5-6µ.: Brazil Clavulina cirrata)
Simple
1-14 cm. high, often caespitose, often becoming rugulose
and hollow: sterigmata 6-9 µ. long: sp. subglobose: trop.
Asia C. spiralis
1-7 cm. high, solitary or 2-3 together, solid, smooth:
sterigmata 9-r 2µ. long: sp. broadly pip-shaped: Malaya . C. brevipes
(-2 cm. high: sp. 6-8 X6-7µ.: with cystidia: Porto Rico Clavulina pilosa)
Sp. 2·5-4·5µ. long: branched (C. biformis, branched or simple)
Sp. 2·5-3·5µ. long: hyphae 2-5µ. wide, not inflated
-1·5 cm. high: sparingly branched: sp. 2·5-3 X2·5-2·8 µ.:
France C. minutula
-2·5 cm. high: branches Jax, inclining to greyish purple at
the tips: sp. 3-3·5 x2·5-3µ.: Europe, N. Amer. C. tenuicula
Sp. larger: hyphae inflating
Java: -7 cm. high, pale yellowish-white, much-branched:
stem -2 cm. thick: sp. 4-4·5 X 3·5-4µ. C. Fleischeriana
N. temp.
-4 cm. high, white, rather tough: sp. 3·5-4·5 x3-3·5µ.:
Europe, N. Amer. C. subtilis
-4 cm. high, simple or sparingly branched: tough, white
to dull yellow: sp. 3·5-5 x2·2-3·5µ.: U.S.A. (? Austral.) C. biformis
Sp. ellipsoid, 2-4·5 µ wide
(Sp. u-15µ. long: -15 mm. high, filiform: France . Clavaria exilis)
Sp. 5-8 X 3·5-4·511.: simple, white or cream, -25 mm. high: stem
distinct, cream flesh colour: U.S.A. C. filipes
Sp. 3" 5-5·5 X 2·5-3·5 µ: -4 cm. high, simple or sparingly
branched, white then yellowish
Sp. subglobose or obovoid: U.S.A. C. biformis
CLAVULINOPSIS
Sp. oblong-ellipsoid or pip-shaped: Malaya C. similis
(see also C. dichotoma and C. subtilis, above)
Sp. 3-3·5 X2-3µ.: -25 mm. high, laxly branched: hyphae 2-5 µ.
wide, not inflating: Europe, N. Amer. C. tenuicula
D. White, then yellowish, brownish, or rufescent from below

upwards (hysterochroic): sp. smooth
Branched: trop.
Sp. 5-7 µ., subglobose: -4 · 5 cm. high, white then brownish,
yellow-brown when old, drying villous in the lower half:
Ceylon C. semivestita
Sp. 4-5 x2·5-3·5µ.: -2 cm. high, white then pale ochraceous or
brownish-ochraceous: Malaya . C. fruticula
Simple or sparingly branched
Sp. 6-8 X5-6 µ., tear-drop shaped or subglobose: -2·5 cm. high,
white then buff-ochraceous, compressed-fusiform: smell
strong: Malaya C. arenicola
Sp. 4-6 X3-4µ., or 3-4 X2-2·5µ.: -2 cm. high, white or yellowish,
then brownish or rufescent: stem distinct: often sparingly
branched: U.S.A., France C. rufipes
Sp. 3·5-5 x2·5-4 µ. (? verruculose): -5 cm. high, often sparingly
branched, yellowish or cream, then the scurfy stem brownish-
ochraceous: foetid: Europe, U.S.A. C. luteo-
ochracea
(-2 cm. high: sp. 6-8 X 6-7 µ.: with cystidia: Porto Rico Clavulina pilosa)
E. Yellow, greenish, orange, ochraceous: sp. smooth
Branched
Sp. 9-10 X7-8 µ. : dubious species
-5 cm. high, yellowish or reddish yellow, becoming white:
Germany . C. subfastigiata
-1·5 cm. high, fawn brown then yellowish, tips paler, little-
branched: Cameroons C. Ledermannii
Sp. 6-8 x3-4µ., ellipsoid: -7 cm. high, pale yellow, much-
branched: Germany C. subfiava
Sp. 4-7 µ. wide, subglobose
-8 cm. high, yellow, egg-yellow or ochraceous, smelling of
new meal: bifurcate tips crescentic: temp. C. corniculata
-4 cm. high, pale greyish yellow, sparingly branched, or
simple: France C. tenella
-7 cm. high, pale yellow, much-branched: stem -2 cm. thick:
sp. 4-4·5 X3·5-4 µ.: Java C. Fleischeriana
-11 mm. high, orange-yellow: sp. 4·5-6 µ.: hyphae 1-2·5µ.: on
amenta of Rhamnus: Holland C. pusilla
(N. Amer.: dull yellowish: sp. 3·5-5 X2·2-3·5µ. . C. biformis)
(Trap., sparingly branched: C. amoena, see below)
Simple, rarely branched
With cystidia, 16-18 x 5-6 µ.: -2 cm. high, yellowish then
brownish, base white-villous: sp. 5-6 X4-4·5µ.: Europe C. luticola
CLAVULINOPSIS 349
Without cystidia
Sp. globose or subglobose, or subtriangular
Sp. 4·5-8·5µ. wide
Stem deeper yellow than the pale hymenium
4-spored
Sp. 5-7·5 x4-6·5µ.: U.S.A. C. appalachiensis
Sp. 7·5-10·5X6·5-9 µ. : Austral. C. luteostirpata
2-spored
Sp. 6-7X5-6 µ., globose pip-shaped: Nov. Scot. see Clavaria
argillacea
Hymenium generally deeper-coloured than the stem
Trop.
Sp. 4-7 µ. : 2-16 cm. high, light yellow, orange, or
apricot: Asia . C. amoena
Sp. 7-7·5X5-6 µ., subtriangular: -3 cm. high, egg
yellow, gelatinous: trop. Amer. . C. fiavella sensu
Martin
Temp.
Caespitose, -14 cm. high, fusiform, hollow, bright
yellow: sp. 5-9X4·5-8·5 µ., strongly apiculate:
N. temp. C. fusiformis
Solitary or gregarious
Pale greyish yellow: sp. 5-6 µ. : France C. tenella
Light yellow to ochraceous, rather tough: sp.
5-7µ.: N. temp. . C. corniculata f.
simplex
Tawny ochraceous: hymenium transversely rugu
lose: hollow: sp. 7 µ. : France C. coliformis
-II mm. high, fasciculate, orange-yellow: on
amenta of Rhamnus: Holland C. pusilla
Sp. 2:5-4·5µ. wide
Java: yellow-brown, caespitose: sp. 4-4·5 µ., globose C. liguloides
England: yellow, caespitose, fragile: sp. 3X2 µ. C. Michelii
Europe, N. Amer. : yellowish: stem becoming brown,
furfuraceous: simple or sparingly branched: foetid:
sp. 3·5-5X2·5-4µ. C. luteo
ochracea
Sp. ellipsoid (distinctly longer than broad)
Sp. with a strong apiculus 1-2µ. long
N. temp.: chrome-yellow to orange, sometimes red at the
base: sp. 5-7·5 X3·5-5·5 µ. C.·pulchra
Trop. Asia: citron to egg-yellow, 1-7·5 cm. high: sp.
6-9 X3·5-5·5 µ., becoming stained yellow on the
hymenium . C. luteo-
tenerrima
Austral.: orange: sp. 6-8X4-5 µ., often subacuminate or
amygdaliform C. Archeri
Apiculus slight, less than 1 µ. long
Sp. 4µ. wide or more
N. temp.: golden, orange, apricot, -8 cm. high: sp.
6-10X4-6 µ. C. luteo-alba
var. latispora
Sp. 10-15X5-8µ. var. macrosporq
350 CLAVULINOPSIS
S. Africa: yellow-ochre, -2·5 cm. high: sp. 7-9X4-5µ C. ochracea
Trop. Amer.: yellowish, -2·5 cm. high: sp. 6-8 x 4-5µ C. fiavella
(Sp. rough, 7-u X 5-6 µ: -15 X 2-4 mm., spathulate C. spathuli-
formis)
Sp. 2·5-4µ wide
Greenish yellow, small
Caespitose, -2·5 cm. high, apex often bifid: smell
oily: sp. 5·5--'7 X2·5-3 µ: Faeroes C. citrino-alba
Solitary or gregarious, -2 cm. high: sp. 5-6 X
3-3·5µ: Canada C. Macounii
Not greenish, larger
-3 cm. high, yellow-ochre: sp. 5-6X3-3·5µ, not
guttate: France C. Daigremon-
tiana
-8 cm. high, light yellow, apricot, or orange, the apex
often pallid or white: sp. 5-8X2·5-4µ, guttate:
N. temp. C. luteo-alba
(Sp. rough, 7-8·5X3·5µ: orange-yellow: Nova
Scotia . see Clavaria
gracillima,
p. 227)
Sp. 3 X 2µ: yellow, caespitose, fragile: England C. Miehe/ii
Sp. unknown
-3·5 cm. high, subsimple, yellow, fragile, stem furcate or bifurcate,
quadrangular: U.S.A. C. tetragona
-r · 5 cm.· high, branched, sulphur then fuliginous: stem thickened
and white pruinose at the base, with a byssoid rhizomorph: on
dead leaves: U.S.A. C. sulphurascens
-5 cm. high, much-branched, orange, then ochraceous: S. Amer. C. patagonica
F. Pink, red, or orange-red: sp. smooth

Much-branched, pale pinkish, flesh-colour or pinkish ochraceous,
tips often reddish: sp. 5-6·5 X5 µ, subglobose, pip-shaped: trop.
Asia . C. alcicornis
Simple or sparingly branched

Sp. ellipsoid
Sp. 7-10X3-4·5µ, subfusiform: coral red: Austral. C. corallino-
rosacea
Sp. 4-6X2·5-4·5 µ, stained red in a crust on the old hymenium:
orange, then blood-red: Java C. depokensis
Sp. subglobose
Amer., Japan, China: orange-red: sp. 5-7 x4-5·5µ C. aurantio-
cinnabarina
Trop. Asia, Austral.: pale pink, orange-red, scarlet, or blood-
red, often flattened or misshapen: hymenium colourless, the
flesh intensely coloured: sp. 4-8µ, globose C. miniata
(See also Clavaria Barlae, C. rosea, and C. Schweinfurthiana, p. 218.)
CLAVULINOPSIS 351
G. Brown, umber, grey, or with purplish tips: sp. smooth
White or hazel-brown with purple tips: smell of aniseed: sp. 5-8µ.,
globose: Holland C. Holmskjoldii
Grey
-7 cm. high, much-branched from the base: sp. 4-6 µ., globose:
U.S.A. . C. cineroides
-4 cm. high, shortly and sparingly branched: stem distinct: sp.
5--'7 µ.: Gt. Britain . C. griseola
(Simple, fuliginous: see Clavaria)
Brown or umber (pale or dark)
Simple, or sparingly branched
Java: -6 cm. high, ligulate, caespitose, yellow-brown: sp.
4-4·5µ., globose . C. liguloides
N. temp.: -5 cm. high, simple or forked once or twice,
yellowish with brown furfuraceous stem: sp. 3·5-5 X 2·5-4µ. C. luteo
ochracea
Branched (without simple forms)
Sp. elliptic, 5-7 X3-4µ.: -5 cm. high, pale brown, branches
many, filiform-acute: hyphae 2-4 µ. wide, not inflated:
France C. propera
Sp. broadly ellipsoid to globose: hyphae inflating
Sp. 9 X 7-8µ.: -1 ·5 cm. high, fawn-brown, sparingly
branched: trop. Africa C. Ledermannii
Sp. 4-7 X3-6 µ.
Purplish brown to umber, branches yellow towards the
golden tips: stem yellow: sp. 5-7µ.: Java . C. umbrina
Tips not yellow: rather sparingly branched
White then pale brown or umber: stem short or none:
sp. 4-6·5 X3-5·5µ.: Europe C. umbrinella
Umber or tawny-olive: stem distinct: sp. 5 X 3µ.:Ja pan C. subumbrinella
Sp. 3-4 X2-3 µ.: pinkish buff to cinnamon: Victoria C. cinnamomea
C. alcicornis (Zoll. et Mor.) comb. nov.

Basinym: Clavaria alcicornis Zoll. et Mor., Nat. Geneesk Arch. Ned. Ind. 1, 1844,
382: sensu v. Ov., Bull. Jard. Bot. Buit. Ser. 3, 5, 1923, 274: le. Fung. Mai. 4, 1923
(non C. alcicornis Lev. = Clavulina Leveillei).
Text-Fig. 146g: Plate 7.
2·5-8 cm. high, scattered or caespitose, branched, cream-rose-pink or flesh-colour,
sometimes pale, becoming pinkish ochraceous or yellowish drab with age, the tips con
colorous, pallid white, reddish, or dingy brown, waxy, not or but slightly tough; taste
sweetish: smell none.
Stem 1_-3 xo·1-0·5 cm., short and thick or rather elongate, whitish and sometimes
subvillous at the base: branches 1-1·5 mm. wide below, two to several times dichoto
mous in alternating planes, or irregularly branched, cylindric or flattened, slightly
broadened at the axils, upright but lax, the tips at various levels, dense or loose, sub
acute then blunt.
On the ground· in· the open and in the forest, sometimes on very rotten wood and
earth (termites): Java (common), Malaya (uncommon).
Spores 4-6·5 µ., round (v. Ov.): 5-6 X4·5-5µ., broadly subglobose pip-shaped
(Corner): pale flesh colour (v. Ov.), smooth, 1-guttate, the walls slightly thickened.
352 CLAVULINOPSIS
Basidia 38-48x8-9µ (38-75x5-8µ, v. Ov.), multiguttulate: sterigmata 2-3-4,
(mostly 3, v. Ov.), 5-7µ long (-17µ, v. Ov.).
Hymenium thickening -150µ, with many included spores, the old basidia more or
less persistent: subhymenium narrow, with unin
flated hyphae.
I ; Hyphae 3-12µ wide, the cells 40-350µ long,
,I
,
with thin or slightly thickened walls, clamped,
narrower and shorter celled near the subhy
men1um.
\ Stem with sterile glabrous surface, the super
ficial hyphae 3-5 µ wide, at the base thinly \'illous
with 1-2µ wide hyphae.
This species is closely allied with C. corniculata
and differs mainly in the colour, more numerous
branches, slightly pip-shaped spores, and tropical
habitat. With pale pink or flesh-coloured speci
mens of Clavaria Zollingeri it may easily be
confused, but its branches are not so brittle.
The original description of C. alcicornis gives
the colour as beautiful rose towards the ends of
the branches. Van Overeem did not examine an
authentic specimen when identifying his Javanese
fungus, but his interpretation of C. alcicornis can
be accepted.
C. amoena (Zoll. et Mor.) comb. nov.

Basinym: Clavaria amoena Zoll. et Mor., Nat.
Geneesk. Arch. Neer!. Ind. 1, 1844, 380.
Synonyms: ? Clavaria clara B. et C., Journ.
TEXT-FIG. 1+7· Linn. Soc. Bot. 10, 1867, 338: Burt, Ann. Mo.
C/avulinopsis amoena: variation in Bot. Gdn. 9, 1922, 56, t. 10, f. 91.
size of the fruit-bodies, x r. ? Clavaria laeticolor, B. et B., ibid.: Burt, ibid.
57, t. ro, f. 92.
Clavaria subaurantiaca P. Henn. et Nym., Monsunia 1, 1900, 8: v. Ov., Bull. Jard.
Bot. Buit. Ser. 3, 5, 1923, 266: le. Fung. Mai. 3, 1923: Petch, Ann. R. Bot. Gard.
Per. 9, 1925, 33 I.
Clavaria strigosa P. Henn. et Nym., Mons. 1, 1900, 8: (non Pers. = Clavariadelphus
fistulosus).
Clavaria subargillacea Ito et Imai, Tr. Sapporo Nat. Hist. Soc. 15, 1937, 55.
Clavaria pallidorosea Fawcett, Proc. Roy. Soc. Viet. 51, 1939, 271, t. 20, f. 2.
Misidentifications: Clavaria aurantia Cke. et Mass., sensu Clel., Toadst. S. Austral.
2, 1935, 268: (non vera = Clavulinopsis Archeri).
Clavariafusiformis Fr., sensu v. Ov., Petch, Clel., et Fawcett.
Clavaria inaequalis Fr., sensu Clel., Proc. Linn. Soc. N.S.W. 41, 1916, 869.
Clavaria laeta B. et Br., sensu Lloyd, Myc. Notes 66, 1922, 1128 (non vera = Clavu-
linopsis miniata).
Text-Figs. 8, 147, 148: Plates 8, 9.
Simple, rarely flattened and with a short branch or two, varying solitary to gregarious
or caespitose, pale yellow becoming more or less rich yellow, very variable in form, size,
and habit.
2-16 cm. X 2--'7 mm., or flattened torr mm. wide, at firstfusiform and acute, becoming
cylindric and blunt or subclavate, even subtruncate, often becoming more or less deeply
and extensively longitudinallyrugulose, often twisted, occasionally spathulately flattened
and then generally with 1-3 blunt or subacute points or short branches, rarely truly
CLAVULINOPSIS 353
branched with 2-4 branches 2-3 cm. long and simple or bifid tips, pale lemon-yellow
becoming rich egg-yellow or pinkish-yellow to apricot, varying bright orange-yellow,
ochraceous, or persistently lemon-yellow, paler towards the often wh{tish growing apex,
occasionally pallid white in the distal third: stem 2-15 X 1-2·5 mm., indistinct,
generally shortly white-villous or strigose at the base, paler concolorous: flesh waxy
firm, solid or becoming hollow, paler concolorous: taste and smell none or slightly
fungusy (distinctly acrid in a branched specimen from Cameron Highlands, Malaya,
29.7.34, c. 1,300 m. alt.).
0)
co
(0)
Or
TEXT-FIG, 148. Clilvulinopsis arnoena: young and old hymenia, X 500: spores, X 1,000.
On the ground in the forest and in shady places in cultivat_ed areas: tropical Asia
(Ceylon, Malaya, Java, Bonin Is!., and Australia (? W. Indies): common in Malaya
from sea-level to 1,300 m. alt. (? higher).
Spores 4-7(-8) X4-6·5(-7)µ, white or pale yellowish, smooth, subglobose, thin
walled, with a large gutta 3 · 5-5µ wide, becoming uniformly vitreous opalescent on
drying and often yellowish (the gutta becoming intense greenish-yellow at maturity,
v. Ov.), apiculus 0·5-1 µ long.
Basidia 35-60 x 5-10µ, subclavate with a long narrow cylindric base, generally
longer in old hymenia, projecting -20µ: sterigmata 2-3-4, mostly 4, 5-9µ long.
5119 Aa
354 CLAVULINOPSIS
Hymenium c. 40 µ thick at first, thickening to 300-µ (or more?), covering the whole
fruit-body except the sterile stem: subhymenium composed at first of narrow short
celled hyphae i ·5-2·5 µ wide, gradually inflating 6-15 µ wide at the base of old
hymenia.
Hyphae 7-14µ wide, longitudinal, clamped, with thin or very slightly thickened
walls, the cells 35-200 µ long, the clamps often loose and curiously formed, the cells
often unevenly inflated, shorter near the hymenium, with few or many, narrow, little
inflated, longitudinal and interweaving hyphae, r ·5-7 µ wide, derived from the
longitudinal inflated hyphae and binding them together (but the system monomitic).
Colour. The yellow colour is caused by the yellowish cytoplasm of the basidia and
subhymenial hyphae and by the pale yellow or pale orange walls of the hyphae of the
flesh. The basidia and subhymenial cells may have fine yellow or orange granules or
guttulae but no conspicuous large oil-drops except in the spores where they are
colourless. It seems that, as in C. luteo-tenerrima, the old spores lying on the hymenium
may absorb some of its yellow colour, but a fresh spore-print is white and remains so
for several days.
The small orange granules in the subhymenial hyphae of the orange form of C.
amoena ( = Clavaria subaurantiaca P. Henn. et Nym.) were found by van Overeem to
turn green with iodine but not blue with strong sulphuric acid and, hence, not truly
carotin.
Variation. This common species is ·one of the most variable, so that it resembles at
times Clavulina rugosa, Clavulinopsis Jusiformis, C. pulchra, Clavaria helicoides, or a
sparingly branched Ramaria. Its macroscopic identification must always be checked
microscopically.
The variation in height seems to be mainly a variation in apical growth. That in
thickness depends on the number of concrescent hyphae in the original primordium,
which may depend on the richness of the substratum, and on the extent of hymenial
growth. Thus old fruit-bodies with much-thickened hymenia become dilated, hollow,
and rugulose.
The habit may depend on the substratum, poor soil giving subsolitary fruit-bodies,
rich forest-humus giving densely caespitose.
The slender fusiform young fruit-bodies appe,ar specifically different from the old
clavate, rugulose, and hollow fruit-bodies. Branching is unusual but in some troops of
fruit-bodies it is not infrequent and clearly suggests the factor for branching is
vestigial.
The variation in spore-size is given in Table XXVI. It needs to be correlated with
other features of the fruit-body before varieties can be recognized.
The variation in size, particularly length, of the basidia is usual in species with
thickening hymenium, the old basidia having long stalks formed in their sinuous
upgrowth. Two or three sterigmata occur so often that they appear normal variations.
Typically the fruit-bodies are intense yellow, but some collections vary into the
reddish-pink of C. miniata, others pale into the white of C. spiralis. The variation
seems to depend on the presence or absence of the diffuse yellow pigment and of the
carotinoid granules. The macroscopic vari:ition needs microscopic explanation, as
mentioned under C. miniata.
Identification. The reasons for maintaining this common and variable species are
given under C.jusiformis. I use the name C. amoena because, as van Overeem pointed
out, it is obviously the fungus intended by Zollinger and Moritzi. No spore-characters
were given by them, but there is only one other species, C. luteo-tenerrima, which
might be regarded as C. amoena. The original description' gives the fruit-bodies as
hollow: those of C. luteo-tenerrima are solid; those of the fungus I refer to C. amoena
1
Original description: 'Simple or paired and slightly coherent at the base, 1 in. long, barely
1 line thick, subterete, hollow, fragile, beautiful golden yellow, glabrous: on the ground in
wet woods near Tjikoya, Java.'
CLAVULINOPSIS 355
often become hollow. Indeed, the description applies exactly to small forms of this
common yellow species which can be found every fungus-season in Malaya and
doubtless, too, in Java.
C. subaurantiaca is the orange colour-form, C. pallidorosea the orange-pink form, 1
and C. substrigosa the pale yellow form: both are connected by all gradations. Petch
notes the variation in colour in the Ceylon specimens. C. subargillacea (Naples yellow)
is indistinguishable.
C. clara and C. laeticolor are known mainly from their original collections from Cuba.
Their descriptions are indistinguishable from C. amoena, as small, simple, terrestrial
species, C. clara being pale orange and C. laeticolor deep orange. Burt gives the spores
as 4-4·5 X3-3·5µ, (C. clara) and 4·5-6 X4-4·5µ, (C. laeticolor). Lorrain Smith
recorded C. laeticolor from Dominica. These three appear to be the only records of
C. amoena in the New World. But compare C.fiavella and C. calocera. (Doty informs
me that the collection he referred to C. clara in 1944 may be a new species.)
C. appalachiensis (Coker) comb. nov.
Basinym: Clavaria appalachiensis Coker, C!av. U.S. Can. 1923, 53, t. 10, t. 91.
Journ. El. Mitch. Sci. Soc. 63, 1947, 46, pl. 4.
Synonym: C. globospora Kauffm., Pap. Mich. Ac. Sci. Arts Lett. 8, 1928, 148.
3--9 X0·5--0·15 cm., simple, solitary or gregarious, pale fleshy yellow to light cream or
creamy yellow, straight or curved, terete, sometimes compressed or finely longitudinally
rugulose, obtuse, tips concolorous, withering brownish: stem 1-4 cm., very distinct,
lemon-yellow, base white tomentose: tender and brittle: smell and taste none.
On humus and rotten logs in frondose and coniferous woods, alt. 1 ,ooo m. : U.S.A.
(N. Car., Pa., Mich., Ore.), Canada (Quebec).
Spores 5-7·5 X4-6·5µ,, white, smooth, subg!obose, 1-guttate.
Basidia 7-8· 5µ, wide: sterigmata (2-)4, long and stout.
Hyphae 3-14µ, wide, narrow near the hymenium, clamped.
I have followed Coker's later suggestion (1947) and reduced C. globospora to a
synonym of this species. It is not easy to see how it differs from pale forms of C.
amoena. (See also Wehmeyer's collection, from Nova Scotia, referred to Clavaria
subfalcata, p. 227, under C. argillacea.)
Coker 5861, in the Kew herbarium, shows the typical clamped hyphae of Clavu
linopsis.
C. Archeri (B.) comb. nov.
Basinym: Clavaria Archeri B., FI. Tasm. 1860, t. 188, f. 3.
Synonym: Clavaria aurantia Cke. et Mass., Grev. 16, 1887, 33: (non C. aurantia
Pers. ex Karst., 1882 = Clavulinopsis luteo-alba).
Text-Fig. 146c.
-6 cm. high, simple, gregarious or fasciculate, cylindric, clavate or somewhat
flabellate and rugose, orange.
On the ground: Australia (Queensland, Victoria), Tasmania.
Spores 6-8 X4-5µ,, white, smooth, broadly ellipsoid, obtuse or often subacute or
subacuminate and somewhat amygdaliform, with a prominent apiculus 1-t,·5µ, long,
evidently 1,--guttate.
Basidia 4-spored.
Hymenium thickening to u.oµ,.
Hyphae -8µ, wide, thin-walled, clamped.
I have examined the typ·es of C. Archeri and C. aurantia at Kew and find the same
characteristic spores very abundant on each. The species is very close to C. pulchra,
1
C. pallidorosea Fawcett, I.e.: -II Xo· 5-1 cm., becoming flattened and twisted, often
hollow, longitudinally rugulose, occasionally bifid, orange-pink: taste and smell sweet: ·
sp. 5-7 p,, subgl., 1-guttate: on burnt ground, solitary or subcaespitose: Victoria.
356 CLAVULINOPSIS
but it needs more detailed study in the living state before one can be certain. Fawcett's
record of C. pulchra' from Victoria seems to have the same spores as C. Archeri.
The species which Cleland and Fawcett have identified as C. aurantia is, I think,
C. amoena which has subglobose spores (4-8 µ, wide, in Cleland's collections; 6·2-7·7 X
4- 6 µ, in Fawcett's collections). The wide range in colour observed by Cleland (rich
to pale yellow and ochre) agrees with C. amoena. But, as noted under C. corallino
rosacea, there may be a complex of colour-forms with the subfusiform spore of C.
Archeri.
· C. arenicola spec. nov.

-2·5 X 0·1-0·15 cm., or flattened -2·5 mm. wide, scattered or gregarious, simple,
cylindric and subacute, then fiattened-ligulate, fusiform or subspathulate and obtuse,
becoming hollow and externally longitudinally sulcate, occasionally once forked, white
then pale buff-ochraceous from the base upward, the tip white, rather brittle: smell and
taste rather strong, of Agaricus campestris.
TEXT-FIG. 149. · Clavulinopsis arenicola: showing the

sterigmata increasing in length as their number
decreases: X r ,ooo.
On sandy soil in forest by the sea, attached to woody rootlets: Malaya (Port Dick
son, Negri Sembilan, 26.1 r.41).
Spores 7-8 X 5-6 /L, pyriform or tear-drop shaped, to subglobose, 6-7 x 5-6 µ,,
white, smooth, 1-guttate..
1 -12 X I cm., -4 mm. thick, usually compressed, becoming longitudinally rugulose,
cadmium yellow to flame-scarlet, orange-chrome or light yellow: taste and smell sweet:
sp. 6·7--7·8 X 4·2-5·3 µ,, oblong-ellipsoid, 1-guttate, with prominent apiculus: Proc. Roy. Soc.
Viet. 51, 1939, 270, t. 19, f. 1.
CLAVULINOPSIS 357
Basidia 30-48 X 6-7 µ., narrowly clavate: sterigmata 3-4, 5-6 µ. long, occasionally
r-2 sterigmata r2-2op., long.
Hymenium c. 30 µ. thick at first, thickening to 60 µ. (at least): subhymenial hyphae
2-3·5µ, wide, closely interwoven as in C. amoena.
Hyphae 3-12µ, wide, the cells 30-150µ, long, very compact, longitudinal, slightly
constricted at the septa, clamped, with fairly numerous, narrow, uninflated, inter
weaving hyphae.
Stem with narrow hyphae 1·5-2·5 µ. wide projecting -roeµ, from the surface as a
thin subtomentum, gradually shorter upward and passing gradually into the hymenium
by the intercalation of basidia, the lower part of the hymenium often with sterile
1·5-2µ, wide, simple or lobed, hyphal ends among the basidia.
This species is near C. amoena but differs in its pale colour, small size, and pointed,
tear-drop shaped spores. C. brevipes has much longer basidia, wider hyphae, and white
fruit-bodies. C. similis has truly ellipsoid spores, narrower hyphae, and still shorter
basidia. C. liguloides is said to have smaller spores.
C. asterospora (Pat.) comb. nov.

Basinym: Clavaria asterospora Pat., Tab. An. Ser. 2, 1887, 568: Locquin, M., Rev.
Myc. ro, 1945, 62, fig.
Text-Fig. r49A.
2-5 cm. x 2-5 mm., simple, solitary or in small groups, slender, cylindric or sub
ventricose, acute then blunt, smooth, hollow, white: stem 5-15 mm., not very distinct,
greenish at the base: brittle: smell and taste inparticular.
On bare soil: Europe (France, Gt. Britain, Holland), rare.
Spores 5-8 µ. (Bourd. et Galz.), 7-8 µ. (Rea), 8-ro(-13) X 6·5-8(-ro)µ, (Donk),
white, subglobose, set with long scattered spines -3·5µ, long, r-guttate (Donk).
Basidia 30-40 X 7-8p.,: sterigmata 4.
Hyphae 4-20 µ. wide, pseudoparenchymatous, with few or numerous clamps (Donk).
TEXT-FIG. r49A. Clavulinopsis asterospora: semi

diagrammatic optical sections of immature
and mature spores stained with cresyl blue: r,
endospore (vinous pink): 2, mesospore (granular
blue, shown in black, often discontinuous): 3,
epispore (pink), producing the spines: 4, exo
spore (bluish): 5, perispore, disappearing at
maturity: X 2,500: (from Locquin).
This seems an albino derivative of C. helvola, but with longer spines to the spores.
C. candida appears to differ only in its pruniform spores. C. tenerrima · may be an
immature state. Donk says that the spines vary greatly in size and number in different
collections. According to Locquin, the spore has five membranes, of which the outer
(or fifth) is extremely tenuous and the third produces the spines by extrusion through
the fourth (or exospore) ..
CLAVULINOPSIS
C. aurantio-cinnabarina(Schw.) comb. nov.
Basinym: Clavaria aurantio-cinnabarina Schw., Tr. Am. Phil. Soc. 2, 4, 1832, 183:
Coker, Clav. U.S. Can. 1923: Wehm., Pap. Mich. Ac. Sci. Arts Lett. 20, 1935, 254:
Martin, Lilloa 5, 1940, 195.
Synonyms: Clavaria Schimadai Imai, Tr. Sapporo Nat. Hist. Soc. II, 1929, 41: 12,
1931, 2I.
Clavaria Miyabeana Imai, ibid. II, 1930, 72: 12, 1931, 16.
1·5-:-7- cm. high, simple, solitary, gregarious or caespitose, deep orange-red, varying
salmon-orange to scarlet, fading huffy orange: flesh deep orange-red, not fading,
moderately brittle: smell rather foetid-aromatic(Coker).
1·2-3 mm. wide, cylindric, subacute or blunt, sometimes flattened and rugulose or
channelled, without distinct stem: drying orange-buff (Coker), reddish brown
(Wehm.): tip of the club generally sterile.
On the ground in the forest and among grass: Canada, U.S.A.,.Panama, Venezuela,
China, Japan(? Philippine Isl.).
Spores 5-7 X4-5·5µ, white or pale yellow, smooth, subglobose, 1-guttate: 7--7·5 X
5·5-6µ(Martin).
Basidia 5·5-7·5µ wide: sterigmata (2-)4, 8µ long: often with elongate abortive
sterigmata(Martin).
Hymenium 100-130µ thick, with many small crystals: cystidia none.
Hyphae 4·5-5·5µ wide, with few clamps(Coker): basidia clamped(Martin).
Coker suggests that C. miniata is synonymous: I have kept them apart for the
reasons given on page 381. I can see no means of distinguishing C. Miyabeana
(scarlet) and C. Schimadai(orange-chrome to deep orange-red). It is curious that the
species has not been recorded from Europe.
Wehmeyer's collections from Nova Scotia had light yellow tips to the clubs when
young(evidently when the growing point is active), strongly or feebly triangular spores
6--7 x 4-5·2µ (with the apiculus at one angle), and dark reddish-brown conducting
organs, 3·5-7µ wide, in the hymenium. The colour and the spores suggest C.pulchra,
but such spores occur also in C. luteo-tenerrima and C. fiavella sensu Martin. There
is no doubt that our concept of these yellow, orange, and red species will be altered
when more is known about them.
C. biformis (Atk.) comb. nov.
Basinym: Clavaria biformis Atk., Ann. Myc. 6, 1908, 56: Coker, Clav. U.S. Can.
1923, 34, t. 91: Burt, N. Am. Clav. 51, t. 9, f. 79: Coker, Journ. Elisha Mitch. Sci.
Soc. 55, 1939, 385, t. 44, and 63, 1947, 46, pl. 5.
1-4 cm. high, simple, once branched or several times dichotomous, dull white to sordid
yellow or light buff with paler tips darkening with age: branches 0·5-1·5 mm. wide,
mostly short, sometimes irregular and crooked, rarely strict: flesh very elastic, not
breaking on doubling: smell none.
On the ground in woods or on rotten wood: U.S.A.(N. Car., N.Y.).
Spores 3·5-5 x 2·2-3·5 µ,, white, smooth, obovoid to subglobose, granular or
1-guttate.
Basidia 20-26 X4-6·5µ: sterigmata 4.
Coker compares this with C. luteo-ochracea, but it seems nearer to C. subtilis and to
differ mainly in the narrower spore, thus connecting C. subtilis with C. minutula and
C. tenuicula. The South Australian specimens referred to C .. subtilis by Fawcett
appear to belong rather to this species: their description is:
-4 cm. high, white then cream, solitary or caespitose: stem usually distinct, glabrous:
branches numerous, often prolonged and subulate, axils wide and rounded, not
flattened: sp. 3·5-4·3 X2·2-3·6µ, ovoid pip-shaped, smooth, 1-guttate: hyphae 4-8µ
wide: taste and smell none: on damp soil: Victoria(Proc. Roy. Soc. Viet. 51, 1939, 9,
t. 3).
CLAVULINOPSIS 359
C. boninensis (Ito et Imai) comb. nov.
Basinym: Clavaria boninensis Ito et Imai, Tr. Sapporo Nat. Hist. Soc. 15, 1937, 53.
-1·5 cm. high, gregarious, branched, white becoming yellowish, especially when
dried, two to five times dichotomously branched, with very short acute tips: base
white and slightly villous.
On the ground in shady woods: Bonin Is!.
Spores 5-7·5 µ, white, smooth, subglobose or broadly ovoid, apiculate.
The authors say this is close to Ramariopsis Kunzei but with smooth spores. It
seems close to C. fruticula and both seem allied with C. subtilis and C. dichotoma.
C. brevipes sp. nov.

1-7 cm. X 1-3 mm., simple, solitary or fasciculate in small groups of two to four,
white then cream colour, cylindric and subacute, then blunt and subclavate: stem
4-6 x 0·7 mm., distinct, white, finely subpruinose, the base subvillous: rather brittle
but firm: smell none: taste rather bitter and unpleasant.
On the ground in the forest: Malaya (Singapore, Bukit Timah, 10.9.39, 19.4.41,
13.6.43).
:•..
;��;
';..�/\
,..
FIG. 150. FIG. 151.

TEXT-FIG. 150. Clavulinopsis brevipes: small fruit-bodies, X 1: spores and basidia, X 1,000.
TEXT-FIG. I 5 I. Clavulinopsis bre-.;ipes: large fruit-bodies, X I: abnormal basidia, X r ,ooo.
Spores 5-7 '5 X 4-5·7 µ, white, smooth, broadly pip-shaped, the wall slightly
thickened, apiculus 0·7 µ long, 1-guttate.
Basidia 50-70 X 9-1 r µ, vacuolate with oleaginous-cloudy cytoplasm or with small
to large oil-drops: sterigmata 4, 9-11(-1:z) X 3 µ, very large, occasionally only r-:z
sterigmata, rarely with bifurcate sterigmata.
360 CLAVULINOPSIS
Hymenium thickening to 180µ: subhymenial hyphae 3-6µ wide, short-celled,
slightly inflated, not pseudoparenchymatous.
Hyphae 6-18µ wide, the cells 25-200µ long, thin-walled, clamped, longitudinal,
with a few narrow interweaving hyphae.
Stem with scattered sterile basidia (often with abortive sterigmata) and divergent
or appressed, 2-4µ wide, hyphal ends, but no palisade: hyphae of the villous layer at
base of stem, as in C. amoena.
This species appears to the naked eye like a white Clavaria tenuipes, but it belongs
to the Clavulinopsis amoena-alliance, and may be only a variety of C. spiralis. It is
close to C. arenicola which has more distinctly coloured fruit-bodies, much shorter
basidia and sterigmata, and somewhat narrower hyphae: whether such differences are
specific can be decided only from further collections. But this, apparently insigni
ficant, species shows in its large basidia and large sterigmata how false is the idea that
small white Clavarias are necessarily allied, cf. Clavulinopsis brevipes, Clavaria vermi
cularis var. gracilis, Clavaria tenuipes.
C. calocera (Martin) comb. nov.
Basinym: Clavaria calocera Martin, Lilloa 5, 1940, 196.
10-25 x 1-2 mm., simple, gregarious, bright orange or golden-yellow, drying orange
brown, fertile throughout except for the very short base, toughly gelatinous: hymenium
dense.
On moss-covered rocks and earth: Colombia (1,250-1,500 m., Hacienda Cincinnati,
Dept. Magdalena, above Santa Marta).
Spores 8·5-10 x 4-5µ, white, smooth, cylindric to suballantoid.
Basidia 25-32 X 5-6µ, clavate, 4-spored.
Hyphae 4-8(-10)µ wide, with very small clamps.
This resembles a simple Calocera in colour and texture. Miss Wakefield suggests
it is near C. clara, which I have placed under C. amoena. C. fiavella sensu Martin
may be allied in spite of its peculiar spore.
C. candida (Weinm.) comb. nov.

Basinym: Clavaria candida Weinm., Hym. Gastr. 1836: sensu Quel., Bull. Soc. Bot.
Fr. 26, 1879, 233.
Simple, gregarious, small, filiform, acuminate, pruinose or puberulous, pellucid white:
stem indistinct, short, hyaline, tomentose at the base.
In swampy woods and greenhouses: Europe (Russia, France).
Spores 10µ long, pruniform, white, aculeolate (Quel.).
Quelet appears to have been the only mycologist to have interpreted this species
microscopically, and his interpretation suggests C. asterospora except for the shape
of the spore.
C. cineroides (Atk.) comb. nov.

Basinym: Clavaria cineroides Atk., Ann. Myc. 7, 1909, 367: Coker, Clav. U.S. Can.
1923, 78, t. 20, 83, and Journ. El. Mitch. Sci. Soc. 63, 1947, 48, pl. 2.
-7 cm. X 5-6 cm., very much branched from the base, without stem, uniformly pale
grey: branches dichotomous, subclavate, axils acute or round, apices bidentate, blunt:
somewhat tough.
On the ground in woods: U.S.A. (N. Car., N.Y.).
Spores 4-6µ wide, white, smooth, globose, 1-guttate.
Basidia 40-45 X 5·5-7µ: sterigmata 4.
Hymenium 75µ. thick (? thickening) (with or without small crystals: ? sand).
Macroscopically this is like Clavulina cinerea, but the small spores and 4-spored
basidia distinguish it. I think it is allied with Clavulinopsis corniculata and perhaps
near the ancestral branched stock from which the other grey species have been derived,
CLAVULINOPSIS
as C. griseola. Coker records a collection from Sweden with white fruit-bodies other
wise identical: it may have been C. subtilis. He gives the spores of four collections of
C. cineroides as 5-6 µ, (Burnham 99, and Vaughns, N.Y.), 4·5-5·5 (Atk., Ithaca), and
4-5·5 µ, (Mrs. Murrill, Chappaqua).
The revised description of Coker's (1947) suggests merely collections of a pale
greyish form of C. dichotoma with caespitose, sparsely branched fruit-bodies.
C. cinnamomea (Fawcett) comb. nov.

Basinym: Clavaria cinnamomea Fawcett, Proc. Roy. Soc. Viet. 51, 1939, II, t. 3, f. 3.
-4 cm. high, mostly branched, rarely simple, solitary or gregarious, cinnamon-buff
to pinkish-buff: stem 1 mm. thick, slender, cylindric, darker brown: branches few, open,
often flattened below, 1-2 ·5 mm. thick, axils rounded and rather wide: rather fleshy.
On the ground in fern-gullies: Victoria (Cockatoo).
Spores 2·8-4·2 X 2-3 µ, (av. 3 ·45 X 2·42 µ,), white, smooth, ellipsoid, 1-guttate, with
prominent apiculus.
This is said to resemble Ramariopsis crocea in form and Clavulinopsis umbrinella in
colour. However, C. umbrinella is never pinkish-buff or cinnamon-buff, but a rather
pale or watery umber which slowly develops on the white branches. The spores also
are much smaller than in C. umbrinella. For these reasons I think that the S. Australian
specimens referred by Fawcett to C. umbrinella must also belong to C. cinnamomea:
the only difference that I can find in their description is the minute apiculus to the
spore, that of C. cinnamomea being prominent:
-5 cm. high, three to four times dichotomous; branches upright 1 ·5 mm. wide, axils
rounded, tips long and blunt, pinkish-buff to cinnamon-buff or browner: base slightly
tomentose: flesh whitish, firm, rather fibrous: sp. 3 ·3-4·2 X 2 ·2-3 ·2 µ,, white, smooth,
apiculus minute: Victoria (Mt. Evelyn) (Proc. Roy. Soc. Viet. 51, 1939, II).
C. citrino-alba (Moll.) comb. nov.

Basinym: Clavaria citrino-alba Moll., Fung. Faeroes 1945, 113, f. 43B, t. 3k.
Text-Fig. 152.
2-2·5 cm. X 2-3 mm., simple, usually with sl:J:.ortly bijid or furcate ·
apex, caespitose, dark greenish-yellow, discolouring white from the
top, base whitish, cylindric-fusiform, curved, very seldom forked
twice, tips acute: flesh whitish-ochre, brittle, fragile, floccose: smell
and taste unpleasant, spermatic or of train-oil (as in Stropharia
coronilla).
TEXT-FIG. 152. Among grass: Faeroes (Sydero, Aug.).
Clavulinopsis Spores 5·5-7 X 2 ·5-3 µ,, white, smooth, oval-ellipsoid, with a small
citrino-alba, x r :
apiculus, 1-2 very small guttulae.
(from Moeller).
Basidia 40-46 X 7-8 µ,, elongate-clavate, 4-spored.
This is, apparently, near C. luteo-alba but with different habit, smell, and lemon
yellow colour. It appears macroscopically as a short, lemon-yellow C. fusiformis, but
with ellipsoid spores. Compare C. Macounii, as well as C. fusiformis var. ceranoides.
C. coliformis (Boud.) comb. nov.

Basinym: Clavaria coliformis Boud., Bull. Soc. Myc. Fr. 33, 1917, 11, t. 3, f. 2:
Sacc., Sy!!. Fung. 23, 1925, 491: Bourd. et Galz., Hym. Fr. 1928.
2-5 cm. X 4-6 mm., simple or scarcely branched, gregarious, tawny ochraceous,
hollow and yellow within, cylindric-clavate, obtuse, transversely plicate, longitudinally
sulcate, white tomentose at the base, the hymenium not distinct from the stem: taste
bitter: smell of tallow.
On the ground: France (Amiens, rare).
Spores 7 µ, wide, white, smooth, globose, 1-guttate.
362 CLAVULINOPSIS
This may be only an unusual state of some well-known species, e.g. C. corniculata
or C.fusiformis.
C. constans (Coker) comb. nov.

Basinym: Clavaria constans Coker, Journ. Elish. Mitch. Sci. Soc. 55, 1939, 383,
t. 44, f. 12 (spores).
9-10 mm. high, narrowly clavate, simple, very pale cream-white, subtranslucent:
stem 2·5-3 X 0·5 mm., watery white, glabrous: rather tough, pliable.
On bare clay, in large troops of single fruit-bodies (not caespitose): U.S.A. (N. Car.).
Spores 7·5-11 X 1·8-2·6µ, mostly 9·0µ long, white, subcylindric, rod-shaped,
slightly curved.
Basidia c. 11 x7·5µ, 4-spored.
Hyphae 3-5 µ wide, long-celled (no clamps seen).
This is said to be near C. vernalis and Lentaria mucida, but it is not associated with
algae. It would appear to be but a pale form of C. vernalis. The basidia seem unusually
wide. On the other hand, if really without clamps, it must be near Clavaria f ossicola.
C. corallino-rosacea (Clel.) comb. nov.

Basinym: Clavaria corallino-rosacea Clel., Tr. Roy. Soc. S. Austr. 55, 1931, 159:
emend. Fawcett, Proc. Roy. Soc. Viet. 51, 1939, 269, t. 22, f. 26: (C. rosea Fr. sensu
C!el. et Chee!, Proc. Linn. N.S.W. 41, 1916, 870).
-12 cm. high, mostly 6-9 cm. X 2-4(-5) mm., simple or, occasionally, branched once
or several times, coral-pink to coral-red, fading with age: stem indistinct: clubs fusi
form, often twisted and compressed, becoming longitudinally rugulose: flesh solid,
concolorous beneath the hymenium, paler in the centre: taste and smell slight,
pleasant, sweet.
On the ground under trees: Australia (N.S.W., Victoria).
Spores 7-10 X 3-4·5 µ (6 X 3·5-4µ, Clel.), white, smooth, rather bluntly fusiform or
(in side-view) amygdaliform, 1-2-guttate.
Basidia with 2-4 sterigmata: hyphae 4-5 µ wide, with many oil-drops.
This species, the considerable emendation of which is confirmed by Cleland, stands
to the Australian C. Archeri as C. miniata does to C. amoena. There may thus be as
difficult a complex of colour-forms with the subfusoid or amygdaliform spore in
Australia as with the subglobose spore in tropical Asia. C. pulchra, in spore-shape, at
least, is their link.
C. corniculata (Fr.) comb. nov.

Basinym: Clavaria corniculata Fr., Syst. Myc. 1, 1821, 471: Konr. et Maubl., le.
Sel. Fung., t. 489: Bres., le. Myc., t. 1096: Coker, C!av. U.S. Can., t. 9, 21, 83 (ut
C. muscoides): Fawcett, Proc. Roy. Soc. Viet. 51, 1939, 5, t. 1, f. 1, 2.
Synonyms: C. pratensis Fr., Syst. Myc. 1, 1821, 471.
C. vitellina Pers., Myc. Eur. 1, 1822, 170.
C. helvola Fr. var. dispar Pers., ibid., 181.
C.fastigiata Fr., Epicr. 1838, 571.
C. muscoides Fr., ibid.
C. similis Pk., Rept. N.Y. St. Mus. 43, 1890, 24 (non C. similis Boud. et Pat.,
1888 = Clavulinopsis helvola).
C. Peckii Sacc., Sy!!. Fung. 9, 1891, 249 (non C. Peckii Sacc. et D. Sacc., 1905 =
Ramariopsis vestitipes).
C.fellea Pk., Rept. N.Y. St. Mus. 51, 1898, 292.
C. caloceriformis Oud., Beih. Bot. Centralbl. 11, 1902, 525.
Cornicularia Bonorden, Handb. Myk. 1851, 166 (non Cornicularia Hoffm., 1789).
Ram.aria corniculata S. F. Gray, Nat. Arr. 1, 1821, 655.
CLAVULINOPSIS
R. pratensis S. F. Gray, ibid.
R. corniculata (Fr.) Quel., Fl. Myc. 1888, 466.
R. fastigiata (Fr.) Quel., ibid.
Text-Fig. 146F: Plate 10.
2-8 cm. high, wholly yellow, egg-yellow, or ochraceous-yellow, finally often brownish
from the base upward, generally branched dichotomously two to three times, occasionally
simple (forma simplex) with incurved crescentic, subulate tips, firm, rather tough,
generally gregarious or caespitose.
Stem 0·6-4 cm. X 1-4 mm., generally short, occasionally none, white subtomentose
at the base: branches cylindric, divaricate with wide lunate axils, sometimes polychoto
mous below, tips concolorous: flesh concolorous: smell of new meal (Bourd. et Galz.,
Corner, Konr. et Maubl., Donk), or none (Cott. et Wakef., Wehm.): taste rank, bitter.
On the ground in open pastures and lawns, and in woods, common: Europe, N.
America (Canada, northern states of U.S.A.), Japan: S. Australia.
Spores 4·5-7 /L wide, white, nearly or quite globose, apiculus 1-1·5 /L long, smooth,
slightly thick-walled, with 1 large gutta.
Basidia 35-60 X 7-9 /L, with 4 sterigmata 6-1011- long.
Hymenium 30-40 /L thick at first, thickening to 100 /L, not stratified: cystidia none.
Hyphae 2-10(-13)11- wide, the cells 60-25011- long, monomitic, septate with clamp-
connexions, not secondarily septate, thin-walled: hyphae of the subtomentum at the
base of the stem 1·5-2·511- thick, with slightly thickened walls, interwoven to form a
loose layer 100-3oo /L thick.
Yellow colour in the cytoplasm of the subhymenial hyphae, old effete basidia and
old spores, little or none in the medullary hyphae.
forma simplex Donk.

Synonyms:? Clavaria austera Britzm., Hym. Stidb. 8, 1885, 289, n. 27.
? Clavaria distincta Britzm., ibid., f. 3 I.
Unbranched, 1·5-2·5 mm. thick, blunt or subacute: stem slender, indistinct:
solitary or in small groups: distributed as the typical state.
This common species, widespread throughout the north temperate region, is
remarkably constant in its colour and microscopic characters but variable in the
development of the fruit-body. When grown in long, damp grass or shady places, the
branching is said to be loose and, in open places, compact with the branchlets forming
a level top, the compact and dwarf form being sometimes distinguished as var.
pratensis ( = C. pratensis and C. fastigiata). What the relation between forma simplex
and the typical state may be I do not know, but forma sir:plex mu�t be either the
ordinary young state before the branches have developed, or a persistently juvenile
state leading to th':: condition of the simple fruit-bodies of C. fusiformis, C. amoena,
and others. It seems that some fruit-bodies may have slightly smaller spores than
others, as can be seen in the accompanying table. The basidia seem also to vary in
size, but their variation is normal: they are short, 35-45 /L long, in the young hymenium,
and lengthen, 50-6011-, in the old, thickened hymenium.
The microscopic structure of the fruit-body is identical with that of C. amoena.
The longitudinal hyphae are but slightly inflated, and rather numerous uninflated
hyphae weave among them to cause the firmness of the flesh.
Juel found two transverse nuclear divisions in the basidium.
Clavaria austera is described as 'resembling C. inaequalis, but: club slightly greenish
then rufescent-yellow, paler below, at length obtusely crenulate above: spores 4-6 /L,
globose: in mountain pastures: Bavaria'. The small spores and rufescent colour
suggest C. corniculata, rather than C..fusiformis. Clavaria distincta has the same spores
and rufous-yellow colour but is merely somewhat curved or twisted.
Whether the Oregon specimens, growing beneath Pseudotsuga taxifolia, described
CLAVULINOPSIS
by Doty, really belong to C. corniculata, I am not sure. Their colour is different
(apricot-yellow to cream-buff); they appear to be larger, -9 cm. high, and more
branched, four to six times; and they have larger spores, 7-7·5(-10)µ,, and larger
basidia, 60-75 X 9-10·5 µ,.
TABLE XXV. SPORE-SIZE IN CLAVULINOPSIS CORNICULATA
Spore-size in µ, Country Author

4"5-'7 France Bourdot and Galzin
5-6·5 " Konrad and Maublanc
6-'7 Gt. Britain Cotton and Wakefield
5-6 Rea
5· 5-'i Corner
5·:;-6 X 5-5·5 " "
5-'7"5 Holland Donk
4·5-6 Sweden Lundell and Nannfeldt
4·5-6 " Coker
4·8-5·5 Europe (Type of C. pratensis)
4-5·5 " ( " C. vitellina)
4-6 U.S.A. (New York, Burnham u8)
S"5 ( " 41)
5-6·5 ( Coker 7)
5-5·5 X4-5 " " ( " Murrill and Volkert)
5-'7 X 5-6 Canada Wehmeyer (Nov. Scot.)
s-s-8 S. Australia Fawcett
C. Daigremontiana (Boud.) comb. nov.

Basinym: Clavaria Daigremontiana Boud., Bull. Soc. Myc. Fr. 33, 1917, 10, t. 4,
f. 3: Sacc., Syll. Fung. 23, 1925, 490: Bourd. et Galz., Hym. Fr. 1928.
1-3 cm. high, simple, solitary or gregarious, ochraceous-yellow, rarely forked, often
compressed and furrowed, acute: stalk indistinct: solid or stuffed.
On burnt earth in swampy ground: France (rare).
Spores 5-6 X 3-3·5 µ, white, smooth, ovoid, vacuolate (not granulate or guttulate).
Basidia 25 X 5-6 µ,: sterigmata 2-4.
The position of this is doubtful. The colour indicates Clavulinopsis, the non
guttate spores Clavaria. It may be C. luteo-alba.
C. depokensis (v. Ov.) comb. nov.

Basinym: Clavaria depokensis v. Ov., Bull. Jard. Bot. Buit. Ser. 3, 5, 1923, 271:
le. Fung. Mai. 3, 1923.
1-8 cm. X 0·5-3 mm., simple, solitary or in small groups, cylindric or subclavate,
becoming compressed and sulcate, filiform when young, stiff and upright, or flexuous,
subacute then blunt, dark orange then blood-red, base slightly white villous and attached
by white mycelial strands.
On dead leaves and twigs in woods: Java.
Spores 4-6 X 2·5-4·5 µ, cylindric, ovoid or pip-shaped, smooth, at first white with
1(-2) golden-yellow oil-drops, finally the old spores on the hymenium deep brown
red and collapsed, forming a granular crust on the hymenium.
Basidia 32-38 X 6-7·5 µ,, clavate, with yellow guttulae: sterigmata 4, -8·5 µ, long.
This may be C. minia,ta var. sanguinea which has the same habit and general appear
ance but subglobose spores. It is not related to Clavaria helicoides as van Overeem
suggests. Old spores lying on the hymenium often becoming coloured with the
hymenial pigment, e.g. C. luteo-tenerrima, and C. pulchra.
CLAVULINOPSIS
C. dichotoma (God.) comb. nov.
Basinym: Clavaria dichotoma God., Gill. Champ. 766: Bourd. et Maire, Bull. Soc.
Myc. Fr. 36, 1920, 72: Bourd. et Galz., Hym. Fr. 1928.
3-6 cm. high, caespitose, branched, white: stem slender: branches dichotomous,
slender, flexuous, obtuse, cylindric or slightly compressed: fragile, becoming rather
tough.
On the ground among moss, in lanes and woods: France, England (near Cambridge,
Oct. 1946), uncommon.
Spores 4-6X3·5-5 µ,, white, smooth, subglobose or obovoid, 1-guttate: 5·5-7µ,,
subglobose, 1-guttate, apiculus 1-1·5µ, (Corner).
Basidia 18-36X4-6 µ, (Bourd.): sterigmata 2-4.
Hyphae 3-10(-15)µ, wide, with thin or slightly thickened walls, clamped, as in
C. corniculata.
This is probably only a form of C. subtilis with the branches somc:,vhat thickened
and compressed near the tips, not merely subulate, and with slightly larger spores.
There seems, however, to be a series from C. dichotoma through C. subtilis and C.
biformis to C. minutula and C. tenuicula with progressively smaller spores and narrower
hyphae: the specific limits, if any, seem arbitrary. C.fruticula, from Malaya, is similar
but distinctly hysterochroic. For a greyish form, see C. cineroides.
Coker reduced C. dichotoma to Clavulina cristata on the evidence of Roum. Fung.
Gall. 3u5 (in the Kew herbarium), which may be C. cristata. I have examined an
authentically named specimen of C. dichotoma of Bourdot's from his collection in the
Paris herbarium and I have recently found almost identical specimens in a wood near
Cambridge.
C. filipes nom. nov.
Synonym: Clavariafilipes B. et Rav. sensu Coker, Clav. U.S. Can. 1923, 20, t. 1,
81: (non vera = Clavaria acuta).
13-24 X 1-1 ·4 mm., simple, solitary or gregarious, cylindric or narrowly clavate,
subacute then blunt, pale whitish cream or greenish cream, the tips yellowish then reddish
with age: stem distinct, as long as or longer than the club, narrower and more trans
lucent, pale creamy flesh colour, flexible: flesh brittle, slightly tougher in the stem: no
smell or taste.
On bare earth: U.S.A. (N. and S. Car., Mich., Alabama).
Spores 5-7·5X3·5-4·5µ,, mostly 6x4µ,, white, smooth, ellipsoid with a small
apiculus, contents granular (rarely 1-guttate).
Basidia 22X5-5 · 5 µ,: sterigmata 4.
Hymenium 35µ, thick, densely set with large and small crystals.
Hyphae -9µ, wide, thin-walled, clamped (Coker).
The type of Clavariafilipes in the Kew herbarium has spores 7·5-10X5·5-7·5 µ, and
secondarily septate hyphae without clamps. It is evidently Clavaria acuta. Coker's
fungus with narrower spores and clamped hyphae must be different and, so, I give
this provisional name. It appears to be a slender, whitish form of C. luteo-alba.
C. flavella (B. et C.) comb. nov.
Basinym: Clavariafiavella B. et Curt., Journ. Linn. Soc. Bot. 10, 1868, 338: Burt,
Ann. Mo. Bot. Gdn. 9, 1922, 57, pl. 10, f. 94: Martin, Lilloa 5, 1940, 194.
2·5 cm. high, simple, slender, cylindric, acute, yellowish, hymenium confluent with
the narrow base, opaque and striate when dried.
On the ground: Cuba (Wright 561).
Spores 7-8·5 x 4-5µ, (Corner), 6-7X4·5-5µ, (Burt), white, smooth, ellipsoid, with
a short apiculus, 0·5-1µ, long.
Hymenium? not thickening.
366 CLAVULINOPSIS
Hyphae -8 µ wide, thin-walled, clamped.
I have examined the type at Kew: it seems very near C. luteo-alba. Martin inter
prets the species rather differently, on the basis of a new collection of Calocera-like
fruit-bodies with oddly shaped spores: his description is:
25-30 x 2-3 mm., simple, Calocera-like egg-yellow, gelatinous, substriate: spores
7-7·5 x 5-6µ, subglobose but triangular or J-lobed in outline, possibly really bluntly
4-lobed: on the ground in mossy litter, W. Panama (c. 2,200 m.).
C. Fleischeriana (P. Henn.) comb. nov.

Basinym: Clavaria Fleischeriana P. Henn., Monsunia 1, 1900, 9.
6-7 cm. high, fragile, branched, pale yellow or yellowish ivhite: stem 2 cm. thick,
fleshy, much branched: branches terete, smooth, fastigiate, crowded, subdivergent,
erect, axils compressed, tips subulate or palmatifid.
On the ground: Java (Tjibodas, 1,300 m. alt.).
Spores 4-4·5 X3·5-4µ, white or pale yellowish, smooth, ovoid or subglobose.
Basidia 5-6 µ wide.
This may be Ramaria, cf. R. fiavo-alba and R. polypus. It was not recognized by
van Overeem.
C. fruticula sp. nov.

1-2 cm. high, solitary or gregarious, once to several times dichotomously branched in
alternating planes, white becoming pale ochraceous from the base upward, then pale
brownish ochraceous on the stem and lower parts of the branches, the acute tips white:
FIG. 153. FIG. 15+.
TEXT-FIG. 153. Clavulinopsisfrutiw/a, X 1.

TEXT-FIG. 154. Clavulinopsisfruticula, X 1,000.
stem 2-7 x 1-1 ·5 mm., often indistinct, sometimes thinly tomentose at the base:
branches 1-1 ·s mm. wide below, few and divergent, or rather numerous, cylindric,
with short internodes, tips acute then blunt: waxy-fleshy, rather fibrous in the stem:
no smell or taste.
On bare earth under hedges and in the forest: Malaya (Singapore, Botanic Gardens,
19.12.41, 18.3.43).
CLAVULINOPSIS
Spores 4-5 X 2 ·5-3 ·5 µ, white, smooth, rather pip-shaped, 1-guttate, the gutta
1·5-2 µ wide.
Basidia 22-28 x 5-6µ, simply vacuolate: sterigmata 4, 3-4µ long, stout, divergent.
Hymenium thickening to 120µ: subhymenium 15-30µ thick, composed of inter
woven short-celled hyphae 2-4µ wide, the cells gradually inflating, 5-16 X 3-14µ,
and almost pseudoparenchymatous in the older hymenia, rather suddenly divergent
from the longitudinal hyphae of the flesh.
Hyphae 2-15 µ wide, the cells -180 µ long, with thin or very slightly thickened walls,
clamped, not or slightly constricted at the septa, longitudinal and. rather compact,
without interweaving hyphae: short-celled and rather more inflated next the sub
hymenium.
Ochraceous and brownish colour of the old branches caused by a pale brown dis
coloration of the basidia of the old hymenium, without definite pigmentation.
This is certainly related to C. corniculata and C. subtilis. The absence of narrow
interweaving hyphae in the flesh of this and other species with pip-shaped spores, as
C. alcicornis and C. similis, may be a distinction from the alliance of C. amoena-C.
corniculata: however, C. arenicola and C. brevipes are intermediate.
C. boninensis has larger spores and may not be hysterochroic.
C. fusiformis (Fr.) comb. nov.
Basinym: Clavariafusiformis Fr., Syst. Myc. 1, 1821, 480: Coker, Clav. U.S. Can.
1923, t. 11, 12, 82: Konr. et Maubi., le. Se!. Fung., t. 494.
Synonyms: Clavaria compressa Schw., Tr. Am. Phil. Soc. 2, 4, 1832, 182 (fide
Coker): (non B., non S::hrot.).
Clavaria platyclada Pk., Bull.Torr. Bot. Cl. 23, 1896, 419.
? C. cochleariformis Pers., Myc. Eur. 1, 1822, 175.
Misidentifications: C.fusiformis sensu v.Ov., Bull. Jard. Bot. Buit. Ser. 3, 5, 1923,
267: le. Fung. Mai. 3, 1923: sensu Petch, Ann. R. Bot. Gdns. Per. 9, 1925, 330:
sensu Clel., Toadst. . .. S. Austral. 2, 1935, 268: to be referred to Clavulinopsis
amoena.
Text-Figs. 9, 146(i).
5-14 cm. high, simple, densely fasciculate, the stems caespitose-connate at the base,
bright yellow to brownish yellow with age, the tips withering brown, becoming fusiform
and hollow, acute, moderately brittle, rather firm: flesh yellow: smell none: taste bitter,
occasionally mild: 2-6 mm. wide, or flattened up to 12 mm. wide, subcylindric or
narrowly fusiform and acute, becoming compressed, often longitudinally sulcate and
distinctly fusiform, rarely bifurcate near the tip: stem indistinct, thinly white villous.
Among grass in fields and woods: Europe (common), Japan, U.S.A. (common),
Canada.
Spores 5-9 X 4·5-8·5µ, white or yellowish in the mass, smooth, slightly thick-walled,
broadly ellipsoid, broadly pip-shaped or subglobose with a strong apiculus 1-2µ long,
occasionally broader than long, globose in end-view, with I large gutta (often slightly
yellowish) or rnultiguttulate.
Basidia 40-65 X 6-9µ (30-45 X 6-8µ, Bourd. et Galz., Donk), multiguttulate with
yellow oil-drops, clavate with a long, filiform-cylindric base 1·5-2·5 µ wide, with a
clamp-connexion (generally rather a wide loop) at the base: sterigmata 4, sometimes
2 or 3, 6-8µ long.
Hymenium thickening up to 100µ, 40-50µ thick at first: subhymenium composed
of narrow, 1 ·5-2·5µ wide, hyphae, closely interwoven, often branching from the
clamps, not inflated, with yellowish oily contents: cystidia none.
Hyphae 2-10 µ wide, thin-walled, rather long-celled, with clamp-connexions (often
with wide or long loops as in C. amoena); with inflated longitudinal hyphae and
narrow, uninflated longitudinal and interwoven hyphae: not secondarily septate:
structure as in C. amoena.
368 CLAVULINOPSIS
Stem with a thin appressed subtomentum or matt superficial layer towards the base
composed of narrow hyphae 1·5-2·5 µ, wide, thin or slightly thick-walled, with clamp
connexions.
var. ceranoides W. G. Sm.
Sow. Brit. Fung. 1908, t. 235.
Synonyms: Clavaria ceranoides Fr., Syst. Myc. 1, 1821, 481.
C. bifurca S. F. Gray, Nat. Arr. 1, 1821, 657.
? C. bifida Chev., FI. Gen. P ar. 1, 1826, 110.
Ramaria ceranoides S. F. Gray, Nat. Arr. 1, 1821, 655.
Apices of the fruit-bodies blunt, often divided, becoming brown: Europe.
TABLE XXVI. SPORE-SIZE IN CLAVULINOPSIS FUSIFORMIS AND C. AMOENA
Author Size in µ,
I Country
I Remarks
C. jusiformis
Bourdot, Galzin 6-8X5-7 France 1-guttate
Corner 6·5-9 X6-8·5 England 1-guttate (see Table XXVII)
Cotton, Wakefield 5--7(-8) " yellow, then colourless
Donk 5-7 Holland pale yellow, multiguttulate
Konrad, Maublanc 6-8X5-7 France 1-guttate
Rea 6-8 England 1-guttate
(Average 7x6 Europe)
Coker 5-6·7 U.S.A. No. 1362: white to cream yellow
5·5--7·5 Vermont (Stratton)
5-6 No. 12
5-6·5 New Hampshire (Farlow)
4·5-5·2X4-4·5 Mass. (Sprague)
6--7X4·8-6 C. compressa (type)
" 4·5-6·5 C. platyclada (type)
Wehmeyer 5-6·5X4·5-6 " Michigan
(Average s·Bx s N. Amer.)
C. amoena
Corner 4·5-7X4-6·5 Malaya 1-guttate, white
van Overeem 4-6 Java 1-guttate (as C. fusiformis)
" 3·5-6·3 " 1-guttate (as C. subaurantiaca)
Petch 4--'7 Ceylon as C. fusiformis
" 4-6 " as C. subaurantiaca
Cleland 4-5·8 Austral. as C. fusiformis
" 4-8 as C. aurantia
Lloyd 6--7 " as C. laeta
Ito, Imai 4·5-5·5 Bonin Isl. as C. subargillacea
Bresadola Java 1 µ, less than in Europe (as C.Jusiformis)
(Average 5·4)
This description is drawn from the writings of European mycologists and from my
notes on English specimens. The species has been described also from N. America,
tropical and subtropical Asia, New Caledonia, and Australia. The N. American fungus
appears to be a variety with slightly smaller spores, possibly the same as the tropical
and Australian fungi which I consider to be C. amoena. The two species are very
closely related.
In Europe C. fusiformis is remarkably constant in form, colour, and habit of the
fruit-body. 1 C. amoena, by contrast, is extremely variable in these respects, so that it
' See p. 345 for the possibility of a gregarious or subcaespitose state which is C. inaequalis
FI. Dan.
CLAVULINOPSIS
grades into the white C. spiralis and the pink or red C. miniata, neither of which is
recorded as temperate (unless C. aurantio-cinnabarina of N. America is the same as
C. miniata). Hence, I cannot believe that the common tropical C. amoena is specifically
identical with C. Jusiformis, though this constant species may be considered in the
evolutionary sense a temperate variety of C. amoena.
TABLE XXVII. SPORE-SIZE IN CLAVULINOPSIS FUSIFORMIS
(One fruit-body, England)
Spore-size in µ. Length -,- width Number of records
6·5x6·o 1· 1 2
7·0X6·o 1·2 5
7·0X6·3 1·1
7·ox6·5 1·1 3
7·3 X6·5 l"I 2
7·5X6·o 1·25 3
7·5X6·5 1·15 2
7·5X6·7 1·1
7·5X7·0 1·1
7·7X7·0 1· 1
8·ox6·5 1·2 2
8·ox6·7 1·2
8·oX7·0 1·1 5
8·0X7·3 1· 1 3
8·ox7·5 1· 1 2
8·3 x7·0 1·2
8·3 x7·7 1· 1
8·5X7·0 1·2 I
8·5X 7'3 1·2 I
8·5x7·5 1· 1 2
8·5X7·7 1· 1
8·5X 8·o 1·06 4
8·7X 8·o 1·1
9·0X7·7 1·17 I
9·0X8·o 1·13 2
9·ox8·5 1·06
6·5-9X6-8·5 1·1-1·3 50
E 1·15 = Average of 21 spores 6·5-7·7 µ. long.
E 1·12 = Average of 25 spores 8-8·9 µ. long.
E 1·12 = Average of 4 spores 9 µ. long.
The spores of C. fusijormis in Europe seem to be slightly larger and more strongly
apiculate than those of C. amoena: Bresadola formed the same conclusion (Ann. Myc.
5, 1907, 240). Those of the American specimens of C. fusiformis are the same size as
in C. amoena yet, from the descriptions given by Coker and Wehmeyer, the fruit
bodies agree with C. Jusijormis of Europe (see Table XXVI).
The spores of C.Jusiformis have a.much stronger apiculus than those of C. amoena,
which suggests that its alliance is, perhaps, rather with C. pulchra and C. luteo
tenerrima (see p. 81).
In so far as the description of C. bifida includes the reference to C. inaequalis of the
Flora Danica, C. bifida is a superfluous name for ·c. inaequalis Fr.; Chevallier's
description and specific epithet suggest rather C. fusiformis var. ceranoides.
Clavaria fusijormis var. congoensis Beeli (Bull. Soc. Roy. Bot. Belg. 58, 2, 1926, 7) is
said to differ from C.Jusijormis in colour, habit, and shape, the fruit-bodies not being
attenuate at the base. It may be a form of C. miniata. The description is:
SII9 B b
370 CLAVULINOPSIS
-8 x 0·3-0·4 cm., simple, solitary, cylindric, undulate, solid, pointed, yellowish white
at the summit, becoming reddish: on the ground in swampy forest: Belgian Congo:
(spores?).
C. griseola (Rea) comb. nov.

Basinym: Clavaria griseola Rea, Tr. Brit. Myc. Soc. 17, 1932, 47.
2-4 cm. high, generally sparingly branched two to three times, grey, drying brownish:
stem expanding into the branches: branches acute, with crescent-shaped tips: flesh
white, firm tough.
On bare soil and among moss in woods: Europe (Gt. Britain, Mulgrave Woods,
near Whitby).
Spores 5-6 µ. wide, white, smooth, globose, 1-guttate: 6·5-7·5 µ. (in co-type, herb.
Kew., Corner).
Basidia 30-5 X 7-8 µ.: sterigmata 4, 8-9 µ. long.
Hyphae 3-8 µ. wide, rather narrow, clamped.
To the naked eye this is like a depauperate Clavulina cinerea, but the small spores
and 4-spored basidia show that it is allied with C. corniculata. It may, indeed, be a
depauperate state of C. cineroides. I have examined the type at Kew.
Donk, who saw the living ·specimens at the Yorkshire foray, thinks it may be C.
Holmskjoldii.
C. hastula sp. nov.

--'7 mm. high, simple, shortly stalked, terete becoming spathulate or lanceolate,
flattened and very thin, 1-2 mm. wide, rarely persistently terete, apex very acute,
sometimes 2-3-fid at the apex or nearly to the base, very pale flesh colour or pale pinkish
cream: stem 1-3 X 0·5-0·7 mm., terete, pallid watery white: flesh very thin, slightly
watery-gelatinous, soft.
Ir1
=
0
TEXT-FIG. 155. Clavulinopsis hastula, X 5.
Scattered singly on sandy, mossy banks in the forest: Malaya (Pahang, Cameron
Highlands, 1,700 m. alt. 29.7.34).
Spores 5-7 X 2·5-3 µ., white, smooth, oblong-ellipsoid, often slightly curved, blunt,
flattened adaxially, vacuolate and apparently aguttate (alc.-form. material).
Basidia 15-25 X 4-6 µ., clavate or pyriform distally, with a slender stalk 2 µ. wide,
rather granular vacuolate, projecting -6 µ., the immature basidia often with subglobose
pyriform distal half: sterigmata 4-5-6, mostly 4 or 6, 3-4·5 µ. long, distinctly outcurved:
rarely with a narrow, uninflated hyphal end, 2-3 µ. wide, abstricting a single spore,
like a 1-spored uninflated basidium.
Cystidia none.
Hymenium c. 20 µ. thick at first, soon thickening -200 µ., the old basidia collapsing,
mucilaginous: subhymenial hyphae 1 ·5-2 µ. wide, the cells 8-16 µ. long, with muci-
TEXT-FIG 156. Clavulinopsis hastula: thickened hymeniwn, subhymeniwn, medullary hyphae, and spores: X 1,000.
372 CLAVULINOPSIS
laginous walls, the older cells at the base of the thickened hymenium gradually inflating
12-25 X 4-7µ: not pseudoparenchymatous.
Hyphae 2-2·5(-3)µ wide, clamped, with slightly thickened, submucilaginous walls,
strictly longitudinal, sparingly branched, cells 40-:woµ long, shorter (30-70µ) and
m9re frequently branched next the subhymenium, not inflated or here and there
irregularly swollen (ampullaeform) 5-12µ wide, the whole cell never inflated, not
secondarily septate, H-connexions frequent: branching normal (not from the clamp):
hyphal tips, at the apex of the fruit-body, 1·5-2·5µ wide, the cells 40-200µ long on
delimitation; not elongating.
Stem with few or numerous, short, divergent, simple hyphal ends -100 X 1·5-2µ,
not as a distinct pile: hymenium starting abruptly at the stem apex.
Mycelial hyphae 2 µ wide, clamped, thin- or slightly thick-walled, loosely twining
round the mucilaginous colonies of unicellular Cyanophyceae (Gloeocapsa) on the
surface of the soil.
This is a phycophilous species with subgelatinous fruit-bodies and narrow hyphae.
Most of the cells do not inflate, though a few of the longitudinal hyphae develop
ampullaeform swellings and the subhymenial cells enlarge slightly. Such lack of
inflation is generally found in hyphae with mucilaginous walls, but the ampullaeform
swellings suggest that the gelatinization of the walls is a modification of an ancestral
state with normal inflation of hyphae with firm walls.
The peculiar flattening of the fruit-bodies must be the mechanical effect of the
thickening hymenium on the softly mucilaginous, strictly longitudinal hyphae of the
flesh. The young fruit-bodies are terete and they broaden and flatten as the hymenium
thickens, exactly as the fruit-bodies of species of Clavaria and Clavulinopsis become
compressed. The hyphae of the flesh of C. hastula offer no resistance to the expand
ing hymenium, but the mucilage is evidently firm enough to prevent the sides of the
hymenium from bulging apart and causing the club to become hollow. However, the
hymenium in expanding cannot raise much lateral pressure because the club does not
become concavo-convex: at most it splits longitudinally through the lack of cohesion
of the longitudinal hyphae, which do not intertwine, or but little, and which have no
interweaving laterals. These clefts in the mature fruit-body falsely suggest that it
may have branched.
C. calocera, C. constans, and C. vernalis must be related: the point is discussed on
page 81.
C. helvola (Fr.) comb. nov.
Basinym: Clavaria helvola F., Syst. Myc. 1, 1821, 482: Pers., Myc. Eur. 1, 1822,
180: (non Coker, 1923 = Clavulinopsis luteo-alba).
Synonyms: Clavaria angustata Fr., Syst. Myc. 1, 1821, 481.
C. aurantiaca Fr., ibid. 482.
? C. rufa Fr., ibid. 480: FI. Dan. 1778, t. 775, f. I.
C. cylindrica Chev., FI. Gen. Paris 1, 1826, 109.
C. fiammans B., Journ. Linn. Soc. Bot. 14, 1875, 350.
C. dissipabilis Britz., Hym. Sudb. 1885, 289, t. 734-28, t. 743-55: Konr. et Maubi.,
le. Se!. Fung., t. 493.
? C. praetervisa Britz., ibid. f. 29.
C. echinospora Baud. et Pat., Journ. de Bot. 2, 1888, 341: (non B. and Br., 1875
Scytinopogon echinosporus).
C. similis Baud. et Pat., ibid. 1888, 406, t. 8, f. 1: Pat., Tab. An. 1889, n. 686: (non
Pk., 1890 = Clavulinopsis corniculata).
C. inaequalis sensu Cott. et Wakef., Coker, Donk.
Text-Fig. 146 h: Plate 10.
3-7 cm. X 1 ·5-4 mm., simple, solitary or in small clusters, not caespitose, subcylindric
or elongate becoming clavate, subobtuse when growing, more obtuse and rounded with
CLAVULINOPSIS 373
age, occasionally with a short branch at the apex, bright yellow to orange-yellow: stem
5-18 X 1-1 ·5 mm., generally distinct as the narrower, sterile, basal part of the fruit-body
vaguely delimited from the hymenium, paler concolorous: flesh white or yellowish,
solid, subfloccose, waxy-firm, scarcely brittle: smell and taste none.
On the ground in woods, pastures, or among moss on heaths: Europe (common),
Teneriffe, Japan, S. Africa (as C. inaequalis), U.S.A. (Mass., Vermont, as C. inaequalis,
teste Burt.).
Spores 4-7 X 3·6-6 µ. (excl. the spines), white or tinged yellow, rather sparsely
verrucose or bluntly echinulate with blunt or subacute processes 1-2µ. long x 1-1 ·5µ.
wide at the base, subglobose, often subangular, 1-guttate (gutta 3-5µ. wide).
Basidia 30-48 X 6-7·5µ., subclavate: sterigmata (2-)4, 5-7 µ. long.
Cystidia none.
Hymeninm 40-5µ. thick, not thickening or very slightly: subhymenium 30-50µ.
thick, composed of closely interwoven hyphae I· 5-5µ. wide.
Hyphae 4-10µ. wide, cells 20-150µ. long, with a few, narrow, interweaving hyphae
i·5-3 µ. wide, thin-walled, clamped, not secondarily septate, shorter-celled and
narrower (2-5µ. wide) near the subhymenium: a few narrow hyphae of the flesh with
oleaginous contents like laticiferous hyphae.
Stem with a compact layer of narrow, longitudinal hyphae 2-3µ. wide at the surface,
with a few scattered abortive basidia and with a gradual transition upwards to the
fertile hymenium: finely villous at the base with 1·5-2·5µ. wide, excrescent hyphae,
thin- or slightly thick-walled.
var. geoglossoides var. nov.

Synonym: Clavaria geoglossoides Baud. et Pat., Bull. Soc. Myc. Fr. 1892, 42, t. 4, f. 1.
Fruit-body compressed, grooved longitudinally, enlarged and sometimes divided at
the apex: stem distinct, paler.
On lawns: France (frequent).
Spores 6-9 X 4-8µ., or 6-8µ. wide and subglobose, ovoid rounded or oblong, with
large conical warts.
The reasons for this use of Persoon's name are given on pages 343-5. Var. geo
glossoides is probably only a form grown on rich soil under wet conditions. Typical
C. helvola has a distinct sterile stalk, though it has generally been overlooked.
Coker doubts if the species occurs in N. America.
C. rufa is a doubtful species, based on the plate in the Flora Danica which may
represent C. helvola, C. luteo-alba, or C. pulchra. In the absence of a type-specimen,
to show spores, a decision seems impossible.
The type-collection of C. fiammans in the Kew herbarium has abundantly the
characteristic spores of C. dissipabilis. It was described as 'caespitose, orange, thickened
and obtuse upward, 4 cm. high: mycelium cinnabar: on the ground, Teneriffe,
3,500 ft. alt.'. Its hyphae are identical with those of C. dissipabilis and the only
discrepancy is the colour of the mycelium, for evidence of which I can find no
collector's note or painting. No other orange Clavaria has such spores and the
occurrence of the European C. dissipabilis in the Canary Is!. is most likely. Hence,
I conclude that C. fiammans is a synonym of C. helvola.
C. Holmskjoldii (Oud.) comb. nov.

Basinym: Clavaria Holmskjoldii Oud., Beih. Bot. Centralbl. 11, 1902, 525: Ned.
Kruidk. Arch. 3, 2, 1902, 672, t. 3, f. 2: Dank, Rev. Nieder!. Homobas. Aphyll. 1933.
2-7 cm. high, branched, solitary or in small groups, white or hazel-brown with dark
purple tips: stem 1 ·5-3 ·5 cm.: branches few, short or well-developed, two to three
times branched, tips blunt, often with a pair of crescentic tips (as in C. corniculata):
smell strong, of aniseed.
On the ground: Holland (rare).
374 CLAVULINOPSIS
Spores 5-8µ, white, smooth, globose, 1-guttate (Donk): 4·2µ, round (Oud.).
Basidia 40-65 x8-10µ: sterigmata 4, -10µ long.
Hyphae 2-6µ wide, thin-walled, clamped.
Compare C. griseola and C. umbrinella.
C. Ledermanni (Bres.) comb. nov.

Basinym: Clavaria Ledermanni Bres., Ann. Myc. 12, 1914, 543.
-1·5 cm. high, small, sparingly branched, fawn-brown turning yellowish, tips paler:
stem 5 X 1 mm.: branches 1 cm. long, few, once or twice forked, apices 2-3-fid.
On the ground under trees: tropical Africa (Cameroons, Ledermann 2893, type).
Spores 9 X 7-8µ, white, smooth, subglobose.
Basidia 28-30 X 7-9µ.
Hyphae 3-10µ wide.
Said to be near C. corniculata but much smaller, with larger spores and without
distinctly crescentic tips. In form the fruit-body must resemble that of C. fruticula.
C. liguloides (P. Henn. et Nym.) comb. nov.

Basinym: Clavaria liguloides P. Henn. et Nym., Monsunia 1, 1900, 8.
-6 cm. X 3-4 mm., simple, caespitose, spongy-fleshy, elongate clavate, cylindric,
stuffed, apex ligulate, broadly compressed, rounded, sulcate, yellow-brown, attenuate to
the smooth base, glabrous.
On the ground in coastal forest (Barringtonia-Terminalia formation): Java (Zandbai).
Spores 4-4·5µ, white, smooth, globose.
Basidia 20-25 x 7-8µ: sterigmata 4, 6 X 2µ, brownish.
This has the same habitat as C. arenicola which has larger spores; they may be
identical.
C. luteo-alba (Rea) comb. nov.

Basinym: Clavaria luteo-alba Rea, Tr. Br. Myc. Soc. 2, 1903, 66, t. 3, f. 2B: ibid. 3,
1907, t. II, c: Coker, Journ. El. Mitch. Sci. Soc. 63, 1947, 54.
Synonym: Clavaria helveola Pers. sensu Coker, Clav. U.S. Can. 1923, 37, t. 1, 4,
81: (non Pers. = C. helvola Pers. = Clavulinopsis helvola).
Plate 8.
-8 cm. x 1-3 mm., simple, occasionally with 2-3 short branches, scattered or caespitose
in small groups, cylindric or narrowly clavate often compressed, rarely flattened to
1 cm. wide, acute then blunt, often curved or flexuous, sometimes becoming finely
longitudinally rugulose, light buff-yellow, often faintly greenish or flesh colour, to deep
rich yellow, orange, or apricot, the tip often paler or white, becoming concolorous or
shrivelling, drying pale ochraceous: stem short, more or less distinct, clearer: flesh
concolorous or orange-yellow, floccose (Rea), brittle (Coker), occasionally with a narrow
hollow: smell none: taste musty (Coker) or of tallow (Rea).
On the ground in pastures, heaths, and woods, rarely on rotten wood: Europe (Gt.
Britain, France, Holland, rather common), U.S.A. (N.Y., N. Car., S. Car., not
uncommon), Japan (as C. citrina Quel.).
Spores 5-8 X 2·5-4·5µ, white, smooth, ellipsoid, flattened adaxially, generally some-
what ovoid, one to several guttulate, the apiculus 0·5µ long.
Basidia 40-60 X 6-7·5µ (25-60 X 4-7µ, Bourd.): sterigmata (2-) 4, 5-7µ long.
Hymenium thickening to 120µ, colourless: cystidia none.
Hyphae 2-8µ wide, narrow, slowly and slightly inflating, long-celled, with few
interweaving hyphae, clamped.
Colour in the flesh only, caused by small pinkish orange guttulae in most of the
longitudinal hyphae.
CLAVULINOPSIS 375
f. montana Coker.
Journ. EI. Mitch. Sci. Soc. 63, 1947, SS, pls. 7 and 14 (f. 4).
Shorter, less caespitose, deeper orange, with relatively thicker clubs and more
distinct stalks.
U.S.A. (N. Car., Highlands, Coker 5571, type).
Spores 6-7·5(-8) X 3·5-3·7µ.
var. latispora var. nov.
Synonym: Clavaria inaequalis sensu Karst. (1882), Bourd. et Galz., Killerm.
C. aurantia Pers. ex Karst., Hattsv. 1882 (non Cke. et Mass., 1887 = Clavulinopsis
Archeri).
Text-Fig. 146 (e).
Spores 6-10 X 4-6µ, wider than typical, 1-guttate.
Basidia 35-60 X 7-9µ: sterigmata 4, 4-5 µ long.
Hymenium thickening to 100µ: subhymenium c. 30µ thick, composed of narrow
hyphae 2-3 · s µ wide: cystidia none.
Hyphae 2·5-8(-u)µ wide, thin-walled, with fairly long cells: structure as in C.
amoena, with many narrow interweaving hyphae.
Stem set with spreading hyphae 1·5-2·5 µ wide, with thin or slightly thickened walls,
forming the villosity.
On the ground in woods (mixed and coniferous) and in pastures: Europe (Scan
dinavia, France, Gt. Britain, Germany).
var. longispora var. nov.
Synonym: Clavaria inaequalis sensu Sacc. (1888), Ricken (1918).
Spores 10-12 X 5-6(-8)µ (Sacc.), 1s X 6µ (Ricken).
Europe (Germany, Italy).
I have examined Coker's collections of Clavaria helveola in the Kew herbarium and
find that they have the same hyphal construction as in Clavulinopsis amoena, C. fusi
formis, C. helvola, and so on. Hence, I agree'with Dank that it must be identified with
C. luteo-alba which has the same spores.
The species is, thus, widespread in the north temperate regions. In Europe it is
generally rich pinkish yellow or orange, often with a white or pallid tip, which may be
concolorous or become so with age. In N. America it varies from pale flesh yellow,
buff-yellow, light egg-yellow, and pale apricot to deeper shades and appears rarely
to be wholly orange. Macroscopically it may be impossible to distinguish from C.
pulchra, C. amoena (tropical), and C. helvola. As mentioned on page 344, it is one of
the species which must have been confused as C. inaequalis.
The typical state has narrowly ellipsoid spores 2·5-4µ wide. There have been
described, however, larger spores, 4-6 µ wide, for collections which, in other respects,
seem identical, namely, C. inaequalis sensu Karst. (1882), Bourd. et Galz. (1927), and
Killerrn. (1928). Cotton and Wakefield (1919) described a large-spared form of
C. luteo-alba from Hazlemere in England, and I have found a similar collection near
High Wycombe (Bucks.). Hence I feel justified in distinguishing the variety latispora.
The evidence for var. macrospora is less satisfactory, though there seems no reason to
doubt the measurements given by Saccardo and Ricken: such long spores may be only
giant spores as occur in Clavaria purpurea along with the normal short spores. The
data are given in Table XXVIII.
The short white tip, characteristic of the fruit-bodies, more especially when young
and beginning to dry, is caused by the colourless apical hyphae and hymenium at the
growing point. The presence of orange-pink guttulae in the cells throughout the flesh
distinguishes C. luteo-alba from other orange-yellow species but brings it into the
complex of C. amoena and C. miniata.
Cooke's and Massee's records of Clavaria inaequalis from Australia and New
Zealand may belong here or to Clavulinopsis Archeri.
376 CLAVULINOPSIS
C. citrina Quel. sensu Hemmi et Kurata (Acta Phytotax. Geobot. 2, 1933, 113-14,
f. 4) must surely belong here. Its description, from Japan, is:
2-7 cm.x2-3 mm., simple, solitary or fasciculate, cylindric to subfusiform, sub
acute or blunt, occasionally once forked, light orange-yellow to mustard yellow; stem
more or less distinct, short: spores 5-8 X 2·5-3·3µ..
TABLE XXVIII. SPORE-SIZE IN CLAVULINOPSIS LUTEO-ALBA
Author Country Size in µ, Remarks

Bourdot, Galzin France 5-6(-8)X3-4
Coker England 5-'7 X3·5
Corner " 5-7·5X3·5-4·5
Cotton, Wakefield " 6-8 X3-4
Rea " 5-8X3 -4
Hemmi, Kurata Japan 5-8 X2·5-3·3 as C. citrina Quel.
Coker U.S.A. 7·5-8·2X 3·7-4 No. 4368
" " 6·3-7·5X 3·7-4 Burnham 6
" " 5·5-6·7X3·7-4 No. 2654
" " 6·2-8·0X2·5-3·7 No. 2770
" " 6·0-6·7X3-3·3 No. 2285
" " 5-7 X3-3·7 No. 5571: f. montana
" " 6·5-7x2·5-3 No. 18
" " 6-7 X2·3-3 No. 2818
" " 5·5-7 X 2·5-3·5 No. 2433
" " 4·8-6X2·5-3·3 No. 2788
Karsten (1882) Finland IOX5 var. latispora
Bourdot, Galzin France 6-9X4-6 "
Killermann Germany 10x46 "
Cotton, Wakefield England 7-10 X4-5 " (Hazlemere, 1919)
Corner " 7-9X4-5 " (High Wycombe)
Ricken Germany 15X6 var. longispora
Saccardo Italy 10-12X5-6 "
C. luteo-ochracea (Cavara) comb. nov.

Basinym: Clavaria luteo-ochracea Cavara, Fung. Long. 1892, Exs., No. 64: Coker,
Clav. U.S. Can. 1923, 32, t. 1, t. 91.
0·6-5 cm. high, simple or flattened above and forked once or twice like antlers, solitary
or in small groups, cylindric, spathulate, pale cream, yellowish, or pale lemon-yellow:
stem concolorous or brownish ochraceous, more or less scurfy-hairy in the lower third: flesh
rather tough: smell foetid (like Phallus impudicus).
On the ground: Europe (France, Italy, rare), U.S.A. (N.Y., N. Car.).
Spores 4-5 X3-4µ. (Cavara), 3·5-4·5 X2·2-3·5µ. (Coker), white, smooth (slightly
warted at maturity, Cavara), ovoid-ellipsoid to pip-shaped, 1-guttate.
Basidia 30 X5-6µ.: sterigmata 4 (Cavara).
Until it has been decided from further collections whether the spores become
verrucose or not, I am uncertain if this is Clavulinopsis or Ramariopsis. If verrucose,
as Cavara gives, then it is near R. asterella, R. crocea, and R. pulchella. If smooth, as
Coker gives both for the American specimens and for Cavara's type, then it s�ems
allied with C. rufipes or, as Coker suggests, C. bijormis. That it is difficult to place in
the key implies that it is imperfectly known.
The following are the spore-measurements given by Coker:
Cavara: 4-5 X3-4µ., slightly warted at maturity (E 1·3).
Coker: 4-4·8 X3-3·5µ., smooth (Cavara's type) (E 1·4).
3·5X2·2, Burnham No. 39 (N.Y.) (E 1·6).
3·5-4·5x2·2-3, Coker No. 16a (N.Y.) (E 1·5-1·6).
CLAVULINOPSIS 377
Coker: 3·7-4·5 X2·5-3·4, Coker No. 2674 ( N. Car.) (E 1·3-1·5).
(E = length -c- width)
C. luteostirpata (Fawcett) comb. nov.
Basinym: Clavaria luteostirpata Fawcett, Proc. Roy. Soc. Viet. 51, 1939, 267, t. 19,
f. 5.
-7 cm. high, simple, rarely bifid, solitary or caespitose in small groups: stem
distinct, smooth, shining yellow (Empire Yellow): club 2-5 cm. X2·5 mm., often
flattened to 4 mm., usually broader to the apex, blunt, pale yellow (Pinard Yellow),
smooth, rather translucent when young: flesh concolorous, soft, brittle, the stem with
a cartilaginous rind: smell and taste none.
On the ground in gullies: Victoria (Cockatoo, Otway Forest).
Spores 7·3-10·5 X6·3-9µ (av. 7·5-8·8), white, smooth, subglobose, with a prominent
apiculus, 1-guttate.
This is said to be readily distinguished by the shining yellow stem and paler, rather
translucent club and to differ from C. appalachiensis in the smooth hymenium and
larger spores. But for the gutta in the spore it could easily be mistaken for a variety
of Clavaria argillacea. Perhaps it is only a form of C. amoena, but the spores have the
larger size of those of C. fusiformis.
C. luteo-tenerrima (v. Ov.) comb. nov.

Basinym: Clavaria luteo-tenerrima v. Ov., Bull. J ard. Bot. Buit. Ser. 3, 5, 1923,
269: le. Fung. Mai. 3, 1923.
Text-Fig. 146 (a, b): Plate 9.
1·5-7·5 cm. X1-2·5 mm., or flattened to 5 mm. wide, simple, occasionally with a
short branch, solitary, gregarious, or caespitose, rather slender, fusiform and acute,
becoming subclavate and blunt or flattened, straight or flexuous, bright citron yellow,
vivid chrome, golden, or rich egg-yellow: stem 5-ro X0·5-2 mm., distinct in small fruit
bodies, indistinct in larger ones, smooth, pale clear yellow, short: flesh paler yellow,
waxy-brittle, solid.
On the ground or in humus, in the open and in the forest: tropical Asia (Java,
Ceylon, Malaya, to 1,300 m. alt.).
Spores 6-9 X3·5-5·5µ, white when fresh, smooth, ellipsoid, pip-shaped, or sub
cylindric, with a prominent apiculus 1-2 µ long, with 1-2 guttae 1 · 5-3·5 µ wide: old
spores lying on the hymenium with homogeneous yellow, vitreous cytoplasm.
Basidia 30-50(-70) X6-8(-9) µ: sterigmata 4, 5-6 µ long.
Hymenium not thickening: subhymenium 30-50 µ thick, composed of 2-5 µ wide,
scarcely inflated hyphae.
Hyphae 2-14µ wide, thin-walled, clamped, the cells of the longitudinal hyphae
30-180µ long, with rather few, narrow, interweaving hyphae 2-4µ wide.
Stem with narrow hyphae 1 ·5-3µ wide at the surface, bearing scattered sterile
basidia near the stem-apex, but the transition to the hymenium abrupt: with very
narrow, excrescent hyphae, 1-2µ wide, at the base of the stem.
Colour caused by a yellow, orange-yellow, or greenish yellow pigment in guttulae
in the young basidia, subhymenial hyphae, and most of the hyphae of the flesh and
also in the vitreous, residual cytoplasm of old, expended basidia and of many narrow
hyphae of the flesh: absent from the mature basidia and spores, the old spores
becoming stained yellow: dried fruit-bodies stain paper yellow.
This tropical species comes very near to C. pulchra. It has more variable fruit-bodies,
slightly larger spores and, perhaps, a differently dispersed yellow pigment which,
however, also stains the old spores in the hymenium: the fresh spore-print is pure
white. The strong apiculus to the spore seems to distinguish the alliance of C. Jusi
formis, C. luteo-tenerrima, C. pulchra, and C. Archeri from that of C. amoena and
C. luteo-alba. In some collections of C. luteo-tenerrima the apiculus may appear to
CLAVULINOPSIS
come from the side of the spore (Text-Fig. 14.6b), but in reality it is the abaxial side of
the spore which bulges below the base as if to form a subglobose spore like that of
C. Jusiformis.
The species is much more variable than van Overeem's description suggests and,
like other simple yellow Clavarias, it cannot be identified with certainty without the
microscope. It does not grow as large as C. amoena and C.fusiformis nor as tufted, but
it becomes much smaller so as to resemble a yellow Pistillaria (Plate 9). The smallest
specimens which I have found were growing on single dead leaves in the forest
(Fraser's Hill, Pahang, 1,000 m. alt.): they were 15-28 mm. high, with the stem 5-10 X
0·5-0·8 mm., and the club 1-1·5 mm. wide. C. miniata var. sanguinea is the only
other species of the genus known to degenerate into the epiphyllous Pistillarioid state.
The spore-data are:
van Overeem, Java: 5·5-8·5 X 3·2-4µ.
Petch, Ceylon: 6-9 X 4-5µ.
Corner, Malaya: 6-9 X 3·5-5·5µ.
C. luticola (Fr.) comb. nov.
Basinym: Clavaria luticola Fr., Hym. Eur. 1874, 678: Rea, Tr.·Br. Myc. Soc. 12,
1927, 226.
5-20 x 0·5-1 mm., simple, gregarious, yellowish then brownish, apex often whitish,
obtuse, attenuate to the white villous base: somewhat fragile.
On bare soil in woods: Europe.
Spores 5-6 X 4-4·5µ, white, smooth, subglobose.
Basidia 8-12 x4µ: sterigmata 2, long.
Cystidia 16-18 X 5-6µ, flask-shaped, apex obtuse, 3µ wide.
Hyphae 2-3µ.
This is Rea's interpretation of the species. I remember finding the fungus with
him on the Malvern Hills in England, but I did not examine it microscopically. The
small size, dingy colour, and gregarious habit of the fruit-bodies will certainly enable
it to be recognized. Its affinity is possibly with C. luteo-ochracea.
C. Macouni (Pk.) comb. nov.
Basinym: Clavaria Macouni Pk., Rept. N.Y. St. Mus. 47, 1894, 150: Burt, N. Am.
Clav. 45, t. 8, f. 69: Coker, Clav. U.S. Can. 1923, 34, t. 91, 92.
1-2 cm. high, simple, solitary or clustered, dingy greenish yellow or pale cinereous,
subacute or obtuse.
Among moss under Cedrus: Canada.
Spores 5-6 X 3-3·5µ, white, smooth, subellipsoid to pip-shaped.
Basidia 5-5 · 5µ wide, 4-spored: hymenium 40µ thick.
Coker suggests that this rare and little-known species may be allied with Clavaria
Crosslandii. Compare Clavulinopsis citrino-alba and C. Daigremontiana.
C. Michelii (Rea) comb. nov.

Basinym: Clavaria Michelii Rea, Tr. Brit. Myc. Soc. 2, 1903, 39.
4-7 cm. X 1-2 mm., simple, caespitose, cylindric, acute, hollow, yellow, white at the
base: flesh yellowish, very fragile.
In grass under a cherry-tree: Gt. Britain (rare).
Spores 3 X 2 µ, white, smooth, subglobose.
The very small spores are the sole distinction of this species. It may be a yellow
variety of Clavaria vermicularis.
C. miniata (B.) comb. nov.
Basinym: Clavaria miniata B., Hook. Journ. Bot. 2, 1843, 416: (non Petch,
1925 = Clavaria helicoides).
CLAVULINOPSIS 379
Synonyms: Clavaria phoenicea Zoll. et Mor., Nat. Geneesk. Arch. Neer!. Ind. 1,
1844, 381.
Clavaria laeta B. et Br., Journ. Linn. Soc. Bot. 14, 1875, 76: Petch, Ann. R.B. Gdn.
Per. 9, 1925, 330: (non Lloyd, Myc. Notes 66, 1922, 1128 = Clavulinvpsis amoena).
? Lachnocladium simplex P. Henn., Monsunia 1, 1900, 142.
Clavulinopsis sulcata v. Ov., Bull. Jard. Bot. Buit. Ser. 3, 5, 1923, 279: le. Fung.
Mai. 2, 1923.
? C. phoenicea var. Ealensis Beeli, Bull. Soc. Roy. Bot. Belg. 58, 2, 1926.
Clavaria caepicolorosa Fawcett, Proc. Roy. Soc. Viet. 51, 1939, 266, t. 19.
f \
TEXT-Fie. 157. Clavulinopsis miniata: abnormally expanded

fruit-bodies, X r.
-10 cm. high, 2-4 mm. wide, often becoming flattened 4-8 mm. wide, simple,
occasionally with a short branch, rarely dilated and fan-shaped 3-5 cm. wide, caespitose
in small groups, rarely solitary, cylindric then fusiform, acute then blunt, soon longi
tudinally sulcate and rugulose, wholly rich orange-pink, rose-pink-apricot, flame-orange
or reddish orange, discolouring yellow at the tips: stem -14 mm. long, short, indistinct,
subcylindric, paler concolorous, whitish at the base, thinly white villous when dried:
flesh paler concolorous, the hymenium colourless, rather waxy, slightly tough, becoming
hollow; smell and taste none.
On the ground in the forest, among grass in the open, on bare earth under trees:
S. Africa, Ceylon, Malaya, Java, Bonin Is!., New Guinea (Carr 14555, Boridi, 1,800 m.
alt., 14.10.35), Australia (Victoria).
Spores 5-7 X 4·5-6·5 µ. (4-5·5 µ., subglobose, Petch), white, smooth, globose or
subglobose, with a short apiculus 0·5-1 µ. long, 1-guttate, often becoming wholly
vitreous on being shed (the wall shrinking).
Basidia 40-90 X 7-8·5 µ. (30-45 X 4-6 µ., Petch), narrowly cylindric clavate, longer
in the old hymenia, with slightly oleaginous cytoplasm, projecting up to 20 µ. from
380 CLAVULINOPSIS
young hymenia, scarcely projecting from the old: sterigmata 2-3-4, mostly 4, 6-10µ.
long.
Hymenium (with subhymenium) at first 60µ, thick, gradually extending to 150µ,
thick: subhymenium composed originally of 2-4µ, wide, narrow, interwoven hyphae
inflating in the older hymenia to 5-8µ, wide, never pseudoparenchymatous: cystidia
none.
Hyphae of the flesh monomitic as in C. amoena: 3-15µ, wide, the cells 30-250µ.
long, with clamp-connexions (often as rather open or distorted loops), slightly con
stricted at the septa, thin-walled, smooth, with pink or reddish minute oleaginous
granules in the cytoplasm.
QQ
TEXT-FIG. x58. Clavulinopsis miniata: spores and basidia, x 1,000:

medullary hyphae, X 500.
var. rosacea var. nov.

Synonym: ? Clavaria rosacea P. Henn., Voelzkow Reise Ostafrica 3, 1908, 19, t. 2,
f. 5: v. Ov., Bull. Jard. Bot. Buit. Ser. 3, 5, 1923, 265: le. Fung. Mai. 3, x923.
Plate 7.
Wholly pinkish flesh colour, clear pink or rose-pink, less intensely coloured than the
typical form: flesh pink, hymenium colourless.
Fruit-bodies 3-15 cm. high, 1·5-4 mm. wide, simple, solitary, gregarious or
caespitose in small or large tufts, cylindric then fusiform, more or less acute, or
flattened up to 8·5 mm. wide and longitudinally sulcate, sometimes rugulose, occa-
CLAVULINOPSIS 381
sionally with a short branch, often twisted, often becoming hollow: stem 0·6-1·8 cm.
long, indistinct, thinly white villous below, sometimes becoming yellowish brown at
the base: without smell or taste.
On the ground in the forest, occasionally among grass in the open: on dead bark of
trees (Madagascar): Madagascar, Java, Malaya (lowlands and mountains -1,500 m. alt.).
Spores 4-7 X4-6µ. (4-5·5µ., v. Ov.), as in the typical state.
Microscopic structure as in the typical state.
var. sanguinea var. nov.
Synonyms: Clavaria miltina B., Hook. Journ. Bot. 1852, 140.
Clavaria cardinalis Boud. et Pat., Journ. de Bot. 2, 1888, 341, t. 8, f. 2: Boud., le.
Myc. 1, t. 174: Josserand, Bull. Soc. Myc. Fr. 53, 1937, 222.
? Clavaria fusiformis var. Antillarum Pat., Doass. Enum. Champ. Guad. 1903, 14.
Clavaria sanguineo-acuta v. Ov., Bull. Jard. Bot. Buit. Ser. 3, 5, 1923, 273: le. Fung.
Mal. 3, 1923.
Plate 7.
7·5 cm. high, deep rose-red, carmine-red, or blood-red, not orange-red or pink: flesh
concolorous, hymenium white.
Fruit-bodies 1·5-3 mm. wide, rather small, solitary, 2-3 together or subcaespitose,
cylindric to narrowly clavate, acute then blunt, or becoming flattened and hollow,
smooth or simple longitudinally sulcate: stem -4 X 2 mm., or indistinct, paler or
yellowish, with white or pale yellowish mycelium at the base: waxy soft: without
smell or taste.
On bare earth in the forest, or attached to single leaves: Jav.., Malaya, India
(Khasya, as C. miltina), Australia (as C. cardinalis).
Spores 6-7 x 5-6µ.: hyphae as in the typical state.
As with all variable species, the interpretation of C. miniata is difficult. It differs
from C. amoena only in the pink or red colour, and intermediate specimens can
occasionally be found which are pinkish yellow or pinkish orange, e.g. C. pallidorosea
(p. 355). I have kept them apart because their typical states are readily distinguished
and because microscopic examination of the colour shows that the apparent transition
needs detailed analysis. The yellow of C. amoena is caused by the yellowish cytoplasm
of the basidia and subhymenial hyphae, as in most yellow species of the genus; the
hyphae of the flesh have also pale yellowish walls: and, in its orange form (Clavaria
subaurantiaca), there are minute orange granules in the cytoplasm of the subhymenial
and medullary hyphae. The red or pink of C. miniata and var. rosacea is caused by
scattered red, or numerous pink, granules in the medullary hyphae, mostly next the
subhymenium, but the basidia and subhymenial hy;:;hae are .::olourless. In the orange
form of C. amoena, van Overeem found the orange granules to be subcarotenoid
(green with iodine, unaltered by sulphuric acid): in C. sanguinea-acuta ( = C. miniata
var. sang11inea), he found the orange-red granules to be truly carotenoid (green with
iodine, blue-violet with sulphuric acid). Thus, it seems that the colour of C. amoena
differs chemically, as well as in position, from that of C. miniata. If their intermediates
are hybrid, they may have both colours in both positions. For the same reason I have
not followed Coker in reducing C. miniata to C. aurantio-cinnabarina because in this
American species the surface (presumably, hymenium) is said to be the most deeply
coloured part: in all forms of C. miniata that I have seen, the hymenium is colourless.
Synonymy. For the type of C. miniata in the Kew herbarium Coker gives the spores
as 5·5-7µ., globose, white. It was described as:
'Parva, c;innabarina, stipite tenui attenuato in clavulam palmato-subbifidam com
pressiusculam, sursum in crassato: on sandy ground among moss: S. Africa.'
These facts agree well with the Ceylon fungus called C. laeta B. by Petch, and with
the Malayan fungus, but the shape, size, and colour of the spores and the habitat
disagree with Petch's interpretation of C. miniata, which is clearly Clavaria helicoides.
CLAVULINOPSIS
C. phoenicea is certainly identical. Its description is:
1-4 in. long, 1 line wide or flattened-connate 2-3 X 1 line, simple, fasciculate,
connate-caespitose, often twinned, becoming flattened, connate and furcate, obsoletely
canaliculate, more rarely simple, fragile, scarcely coriaceous, beautifully phoeniceous,
becoming pale with age, the acute tips blackening: whitish and often villous at the
base: on the ground in a nutmeg-plantation and in the forest: Java. 'Fungus specio
sissimus, Cl. purpureae et anomalae affinis et iisdem intermedius.'
This flame-orange, or phoeniceous, fungus is evidently Clavulinopsis sulcata: its
spores are identical and it agrees in form, habitat, and locality, though the colour is
rather reddish pink. The unusual length and position of the sterigmata in some
basidia, which suggested to van Overeem the new genus, are abnormalities, as
explained on page 75,
C. rosacea sensu v. Ov. is certainly the same as the Malayan specimens on which
I have based the variety, but the type of C. rosacea (on bark of trees, Madagascar) has
not been examined, though the description agrees in other respects.
C. sanguineo-acuta has the colour, small size, and spores of var. sanguinea, which I
consider merely a deeply coloured form of C. miniata with limited apical growth.
C. cardinalis is almost certainly a synonym: the spores of the type are given as 5-7 µ,
wide, globose, by Coker and by Josserand, and the basidia as mostly 3-spored, occa
sionally 2- or 4-, thus recalling C. sulcata. C. cardinalis was described from specimens
on a trunk of Todea barbata in a Paris conservatory: perhaps it came from Australia.
The type of C. miltina in the Kew herbarium has no spores and the hymenium is very
collapsed, but it has the hyphal structure of C. miniata with many narrow interweaving
hyphae between the longitudinal ones: it was described as simple, crimson, fragile,
and gregarious: the habitat, on rotten wood, seems anomalous, but in tropical forests
there is often little distinction between wood and rotten leaves in the last stages of
decay.
C. fusiformis var. Antillarum, also said to be lignicolous, is described as resembling
C. fusiformis but rubescent, less caespitose, and attenuate into a whitish stem. It may
be C. aurantio-cinnabarina, if it is different from C. miniata.
C. phoenicea var. Ealensis is described as:
6 x 0·3-0·5 cm., simple, lanceolate-liguliform, hollow, tapered to the apex, rose
purple: flesh pale rose: spores not seen: gregarious on the ground in swampy forest:
Belgian Congo.
Lachnocladium simplex is described as:
2 cm. x 2 mm., caespitose, pale yellowish red, coriaceous-tough, simple, subulate,
curved, subacute or obtuse, pruinose or minutely setulose: spores 4-4·5µ,, hyaline
fuscidulous, smooth, globose: basidia 18-20 X 5-6 µ, ; on the ground: Java (Tjibodas).
The tough flesh may refer to the dried state: the basidial length may not include the
attenuate base. Simple Lachnocladia s. str. are not known.
Clavaria caepicolorosa, from Victoria, seems to be a massive red state of C. miniata,
reaching 15 cm. X 5-6 mm., or 15 mm. when compressed. It may have larger spores
(7-9µ,), but this measure may include the apiculus.
The Victorian 'Flame Fungu!l', described by Fawcett as Clavaria Miyabeana, also
seems to be a monstrous state of C. miniata which is often more or less flattened and
misshapen. Whether this is a normal variation or an abnormality is not yet clear.
The Flame Fungus is said to reach 14 x 7 cm. and to be variously flattened and mis
shapen, often with the margins inrolled, and longitudinally rugulose, grenadine-red,
with the globose spores 6·5-8·5 µ,: (Proc. Roy. Soc. Viet. 51, r939)..
Variation. Beside the main variation in colour, there is one in structure. Some
collections have many narrow interweaving hyphae in the tissue, as in C. amoena;
others have few and even none. I have not connected the feature with any other.
As mentioned under C. luteo-tenerrima, this is the only other species of Clavulinopsis
which tends to become epiphyllous as Pistillaria.
CLAVULINOPSIS
Spores. In the following table I have listed the spore-data for C. miniata, C. aurantio
cinnabarina, and Clavaria rosea var. subglobosa. The indication is that they are one
universal species, or that the temperate N. American and the European fungi have
subglobose spores compared with the globose tropical C. miniata.
TABLE XXIX. SPORE-SIZE IN CLAVULINOPSIS MINIATA
Author Cou11try :::.pore-size i11 µ. Remarks

Coker S. Africa 5·5-7·4, globose c.
mimata (type)
Corner Malaya 4·5-7X4·5-6·5 C. miniata et var.
Berkeley Ceylon 7·5, globose C. laeta (type)
Petch " 4-5·5, globose C. laeta
Ito, Imai Bonin Is!. 5(�·5), globose · C. laeta
van Overeem Java 4-5, globose C. sanguineo-acuta (type)
" " 4-5·5, " C. rosacea
" " 4-6, " C. sulcata (type)
Coker, Josserand (? Australia) 5-7, " C. cardinalis (type)
Coker
Wehmeyer
Overholts
U.S.A., Canada
Michigan
Venezuela
4·8�X4-5·5
5·5�X4·5-5
5�, globose
.
C. aurantio-cinnabarina
"
"
"
Martin Panama 7�·5 x5·5-6 " "
Hemmi, Kurat;i Japan 5·7-7 X 4·5-5·5 " "
Auctt. Europe 5-8x4-6 Clavaria rosea var. subglobosa
C. minutula (Bourd. et Galz.) comb. nov.

Basinym: Clavaria minutula Bourd. et Galz., Hym. Fr. 1928, 105.
6-15 x 0·2-0·5 mm., very small, white, with 2-3 simple or forked branches arising
about the middle of the fruit-body.
On bare earth: France (rare).
Spores 2·5-3 X2·5-2·8µ, white, smooth, subglobose, 1-guttate.
Basidia 14-20 X4-4·5 µ.
I have examined the type-collection (on loan from the Paris herbarium). The
species is very near C. tenuicula. Both appear related to C. propera in their uriinflated
hyphae which, doubtless, cause their small size. Compare also small states of C.
subtilis.
Forma subasperata Bourd. et Galz. I have transferred to Ramariopsis Kunzei as a
variety.
C. ochracea nom. nov.

Synonym: Clavaria pulchra sensu v. d. Bijl, S. Afr. Journ. Sci. 29, 1932, 320,
t. 3, f. 1.
-2·6 cm. X 1-3 mm., simple, caespitose, cylindric or flattened, blunt, tapered below,
yellow-ochre: stem indistinct, paler.
On the ground: S. Africa (Stellenbosch, Rhodesia, Cape Colony).
Spores 7-9 X4 · 5 µ, white, smooth, elongate-ovoid, with a lateral basal apicuJus,
mostly I sguttate.
This has larger spores and a duller colour than C. pulchra. Perhaps it is C. inaequalis
or C. luteo-tenerrima.
C. patagonica (Speg.) comb. nov.

Basinym: Clavaria patagonica Speg., F. Patagon. 19, n. 53: Sacc., Syll. 6, 1888, 692.
-5 cm. high and wide, orange then ochraceous: stem 15-20 X5-6 mm., white, often
connate at the base: branches numerous, straight or flexuous, crowded, longitudinally
CLAVULINOPSIS
striate or canaliculate, tips botryoidal-ramulose and truncate lobulate: flesh rather
tough.
On the ground among roots of Senecio sp. on dry hill-sides: Patagonia (near
Gregory Bay).
Spores hyaline.
Perhaps only C. corniculata or Ramariopsis crocea.
C. pogonati(Coker) comb. nov.

Basinym: Clavaria Pogonati Coker, Journ. El. Mitch. Sci. Soc. 55, 1939, 384,
t. 41 et 44(f. 14).
10-23 x 3·5 mm., solitary or in small troops, sparingly branched or simple, slender,
pallid white: stem indistinct in the simple fruit-bodies, sometimes forked nearly from
the base: branches somewhat flattened at the forks, filiform: very tough and pliable.
On earth covered with protonemata of Pogonatum brevicaule: U.S.A.(N. Car.).
Spores 11-13 X 4·2-5·4µ,, white, smooth, subellipsoid.
Basidia c. 9·5µ, wide, 4-spored.
Hyphae very slender, clamped.
This is said to belong to the alliance of C. vernalis and C. constans. On the other
hand, it may be a species of Aphelaria.
C. propera(Bourd.) comb. nov.

Basinym: Clavaria propera Bourd., Bull. Soc. Myc. Fr. 48, 1932, 208.
4-5 X 4 cm., branched, pyriform in outline, confluent by the branches, isabelline
hazel, brownish: stem 10 x 1-3 mm., slender, brownish: branches 0·5-1 mm. wide,
much ramified, with very acute axils: branchlets very slender, very pointed, filiform with
elongate tips: brittle, fragile: no smell or taste.
Among grass: France(Bourd. 43366).
Spores 5-'7 X 3-4µ,, white or slightly isabelline, oblong, with one to several small
gu�tulae. Basidia 24-30 X 5-6µ,.
Hyphae 3-5 µ, wide, thin-walled, subequal, longitudinal, little coherent, clamped,
not inflating.
Subhymenium fairly thick(? thickening hymenium).
Bourdot regards this as related to C. umbrinella, C. dichotoma, C. subtilis, and
Ramarwpsis Kunzei. But the narrow uninflated hyphae are peculiar and such as occur
in C. minutula and C. tenuicula. It is difficult to see how the, texture can be brittle
with this construction. I have been able to examine the type-collection, on loan from
the Paris herbarium.
C. Puiggarii(Speg.) comb. nov.

Basinym: Clavaria Puiggarii Speg., Fung. Arg. 3, n. 39: Sacc. Sy!!. 6, 1888, 697.
-2·5 cm. high, slender, white or pallid: stem 15 X 0·3-1 mm., rather long, becoming
fuscous, arising from a tuberculiform base 1·5-2 mm. wide, flattened and expanding
upward, even canaliculate-lacunose, dividing abruptly into 5-6 branches, simple or
with very few branchlets near the acute tips(5-10 X 0·3-0·5 mm.).
On the ground in shady places: Brazil(Apiahy, leg. Dr. J. Puiggari).
Spores 5-7µ,, white, smooth, globose, papillate-apiculate.
Perhaps Clavulina.
C. pulchra(Pk.) comb. nov.

Basinym: Clavaria pulchra Pk., Rept. N.Y. St. Mus. 28, 1876, 53, t. 1, f. 10: Coker,
Clav. U.S. Can. 1923: Donk, Rev. Nieder!. Homobas. 1933, 82: Wehm., Pap. Mich.
Ac. Sci. Arts Lett. 20, 1935, 253: Moll., Fung. Faeroes 1945, 112, f. 42(ut C. rufa Fr.).
Synonyms: C. persimilis Cott., Tr. Br. Myc. Soc. 3, 1909, 182(teste Coker, Donk,
Josserand).
CLAVULINOPSIS
C. inaequalis var. aurantia Bres., le. Myc., t. 1099.
C. inaequalis sensu Schroet. (? sensu Quel., 1888: ? sensu Karst., 1889).
Text-Fig. 146 (d): Plate 8.
1·5-10 cm. X 1·2-6 mm., simple, solitary, gregarious or fasciculate in small tufts, clear
yellow to deep yellow, orange, orflame, drying deep orange, cylindric and acute, becoming
blunt, compressed and sometimes spathulate or rugulose: stem 2-20 X 1-1·7 mm.,
distinct, drying finely subtomentose: flesh pale, often hollow, floccoso-firm, not brittle:
smell and taste none or sweetish.
Among grass in fields, or on the ground in woods: Europe (Gt. Britain, Faeroes,
Holland, France, Germany, Italy, probably common), U.S.A. (widely distributed),
China.
Spores 5-7·5 X 3 ·5-(, µ, white (old spores in or on the hymenium stained yellow or
orange), smooth, slightly thick-walled, ovoid, broadly ellipsoid or subglobose,
1-2-guttate, with a strong, often sublateral, apiculus 1-2µ long.
Basidia 40-(,0 x7-10µ (50-60 x 5-6µ, Josserand: 44-52 x7-7·5µ., Moller:
40-55 x7-8·5µ. and 48-58 X 8-10µ., Corner): sterigmata 2-3-4, 7-10µ. long.
Hymenium thickening to 100µ.: cystidia none.
Hyphae 3-12µ wide, clamped, long-celled, longitudinal, without interweaving
hyphae: a few laticiferous (Josserand): 2-3µ. wide, slightly thick-walled, variously
excrescent on the surface of the stem.
Colour mainly in the hymenium and subhymenium, as a diffuse yellow pigmentation
of the cytoplasm of the subhymenial hyphae, the old effete basidia and the old spores
in the hymenium, also in some of the longitudinal hyphae: orange and flame-coloured
fruit-bodies also with rich orange-brown granular masses in the old immersed basidia
and in some subhymenial hyphae: sometimes also a smeary yellow exudation on some
of the medullary hyphae.
forma coccineo-basalis Josserand, Bull. Soc. Myc. Fr. 53, 1937, 224, f. 16.
Synonym: ? Clavaria bulbosa Wallr., Krypt. FI. 4, 1833, 538.
Base of stem more or less intensely scarlet.
Under Pseudotsuga Douglasii: France (La Tour du Salvagny, Rhone).
There is general agreement on the identity of this fungus.
Schroeter seems to have been the .first to discover it in Europe, under the name
C. inaequalis (see p. 345). It has not been recorded from the tropics or the southern
hemisphere, unless C. Archeri is identical.
· Moller proposes to identify the dubious Clavaria rufa with C. pulchra on the ground
that the illustration in the Flora Danica, typifying C. rufa, is macroscopically identical
with C. pulchra. Cotton considered it to be a distorted form of Clavulinopsis helvola
(Tr. Brit. Myc. Soc. 3, 1907, 33), where I think it may well be forgotten. The species
has never been rediscovered as a new idea, and it has served no pul'pose. Whether it
is C. pulchra, C. helvola, or C. luteo-alba could be proved only from the spores, and
there is no type-specimen for verification. I doubt if any mycologist would stake his
reputation on the correct determination of the simple non-caespitose Clavarias, even
of Europe, without recourse to the microscope.
Donk reduces C. inaequalis sensu Bourd. et Galz. to C. pulchra but, as discussed
on p. 344, I think it must be considered as C. luteo-alba. On the other hand, C.
inaequalis var. aurantia Bres. has spores identical with C. pulchra and must be so
interpreted. C. pulchra sensu v. d. Bijl I have called C. ochracea.
The red base of f. coccineo-basalis needs explanation. C. flammans, which is identical
with C. helvola ( = C. dissipabilis), ;as also described with cinnabar mycelium (see
p. 373). C. bulbosa is a dubious species described thus:
Simple, orange, attenuate into the rufescent stem inflated at the base: Germany.
C. pulchra, C. luteo-tenerrima, C. Archeri, and C. fusiformis have a strong apiculus
to the spore which suggests their alliance, in contrast with C. luteo-alba and C. amoena.
5119 cc
386 CLAVULINOPSIS
C. pulchra and C. luteo-alba have widely different pigmentation, though macro
scopically they may have almost the same colour.
Wehmeyer writes of one collection of C. pulchra, No. 1312, from Michigan:
'Young plants show what appear to be conidia. Long slender hyphae (1-1·5µ wide)
grow out through the hymenial layer to a distance of 20-30µ and cut off from their
apices numerous oblong cylindric conidia which are hyaline and measure 4-11 X
1·5-2µ. Typical basidiospores were found in the same hymenia and the plants appear
normal in all respects, showing no indication of parasitism.'
TABLE XXX. SPORE-SIZE IN CLAVULINOPSIS PULCHRA
Author Spore-size in µ Country Remarks

Coker 6·7X4·4 U.S.A. Type-specimen
" 6-7X4·6 " Nos. 1717, 5877 usually 1-guttate
" 6-6·7X4·5-5 " No. 3149
" 6·3X4·5 " New York (Catskill Mts.)
" 6·5-7x5·5 " N.Y. (Chappaqua)
" 6·6X4·8 " N.Y. (Frostburg)
" 5-7·5X4-4·8 " Conn. (Redding)
Wehmeyer 5·5-7X3·5-4·5 " Michigan
Cotton and Wakefield 5-6x4 England r-guttate (as C. persimilis)
Rea 5-6x4. " " "
Comer 5-'7X4-4·5 " "
" 6-7·5X4·7-6 " "
Bresadola 5-6X4-5 Italy as C. inaequalis var. aurantia
Donk 5-7X3·5-5 Holland r-guttate
Josserand 5-6X4-4·5 France "
Moller 6x4-5 Faeroes " (as C. rufa)
C. pusilla (Coker) comb. nov.

Basinym: Lachnocladium pusillum Coker, Joum. El. Mitch. Sci. Soc. 42, 1927, 252,
t. 47, f. 2: Fl. Bat. 27, 1928, t. 2136b: Donk, Rev. Niederl. Homob. Aphyll. 2, 1933, 72.
10-11 mm. high, formed of 6 or more divergent branches joined at their bases, or
merely fasciculate, orange-yellow: branches subcylindric, somewhat dilated above,
smooth, in places longitudinally rugulose, simple or once dichotomous, tips furcate
or 3-6 toothed: flesh tough, pliable, becoming hard and horny (? on d rying): hymenium
amphigenous.
On an amentum of Rhamnus alnifolius in a plant-house: Holland (Leiden, Botanic
Garden).
Spores 4·5-6µ wide, white, smooth, globose, 1-guttate.
Basicl.ia 20-28 X 5-6·5µ: sterigmata 2-4, 4-6µ long.
Hyphae 1·5-2·5µ wide, with somewhat gelatinous thickened walls, agglutinated,
parallel: the tips of the branches with hyphal ends 2·5 µ wide (i.e. the growing-tips).
The position of this species is problematic. It cannot be Lachnocladium s. str.
The branched fruit-body and the guttate spores do not agree with the Clavariadelphus
Pistillaria series, but suggest Clavulinopsis, as does the orange-yellow colour: the
habitat, however, disagrees with Clavulinopsis. I place it here for convenience and
because it may be related with the other anomalous species, e.g. C. calocera.
C. rufipes (Atk.) comb. nov.

Basinym: Clavaria rufipes Atk., Ann. Myc. 6, 1908, 57: Burt, N. Am. Clav. 33, t. 6,
f. 43: Coker, Clav. U.S. Can. 1923, I06, t. 91: Wehm., Pap. Mich. Ac. Sci. Arts Lett.
20, 1935, 253: Coker, Joum. El. Mitch. Sci. Soc. 55, 1939, 384, t. 42.
Synonym: Clavaria microspora Josserand, Bull. Soc. Myc. Fr. 64, 1942, 31, f. 8
(sine diagnosi).
CLAVULINOPSIS
-2 cm. high, simple and 1-1 ·5 mm. wide, or flattened and sparingly branched with
antler-like arms, solitary or gregarious, slender, delicate, white, cream, straw-colour, or
brownish: stem 6-8 mm. long, generally distinct, concolorous or pale brown or rufescent,
arising from a mycelial patch and often expanding into a few, rather flattened, blunt
branches: flesh pliable, slightly tough, not brittle.
On humus, twigs, and leaves: temperate N. America (U.S.A., N.Y., N. Car.:
Canada, Nov. Scot.), France (? introduced).
Spores 4-6x3-4µ (Coker), 4-6x 2·5-3µ (Atk.), 3·5-4 x2-2·5µ (Wehm.), white,
smooth, pip-shaped to ovoid, 1-guttate.
Basidia 23-30 X4-5µ (Coker): sterigmata 2-4: hymenium 25µ thick.
Coker compares this with Clavaria delicata, which I refer to Lentaria because it is
truly lignicolous and has aguttate, oblong-ellipsoid spores. ;l?ossibly C. rufipes is
related with Clavulinopsis Jruticula or C. luteo-ochracea.
I am unable to distinguish C. microspora Josserand. It has exactly the same shape
with the tendency to produce flattened antler-like forms (-23 mm. high), the same
colour (pale whitish alutaceous, then ochraceous or ochraceous rufescent, then date
brown from below upwards), small 1-guttate spores (3·5-4·5 X2-2·3µ), small basidia
(22-35 X4-6µ), and narrow (apparently clamped) hyphae 4-7µ wide, sometimes
granular-incrusted. It was found in troops in pots of ferns in a conservatory at the
Pare de la T�te d'Or, Lyon, France, and may have been introduced.
C. semivestita (B. et Br.) comb. nov.

Basinym: Clavaria semivestita B. et Br., Journ. Linn. Soc. Bot. 14, 1874, 75: Petch,
Ann. R. Bot. Gard. Per. 9, 1925, 337 (haud Lachnocladium semivestitum B. et Br.
= Aphefuria tuberosa).
-4·5 cm. high, branched, in groups of 3-4 or in tufts -3 cm. wide, white then brownish,
yellow-brown when old, drying minutely tomentose and rufous or pale brown in the lower
half, glabrous and dark brown above: stem -3 mm. wide, cylindric, soon dividing at
a wide angle: branches dichotomous, generally twice, axils rounded, tips acute or
obtuse, usually simple, sometimes -1 cm. long.
On the ground among grass: Ceylon (Peradeniya).
Spores 5--7µ wide, white, smooth, globose.
Hyphae inflated, clamped (Corner).
The tomentose covering cannot be seen until the fruit-bodies have been dried: it is
then very noticeable and extends to an unusual height along the main branches. The
affinity of the species cannot be decided until it is known whether the hyphae are
truly pigmented, as in C. corniculata, or merely discoloured with age, as in C.fruticu/,a,
Van der Bijl has doubtfully recorded the species from S. Africa (Somerset East)
with globose white spores, 4µ wide. Compare C. umbrinella.
C. similis sp. nov.

Text-Fig. 159.
-4-cm., simple or, usually, dichotomous once, rarely twice, with short or long acute
branches, 1 mm. wide, solitary or subcaespitose in small groups, pallid white, yellowish
with age: stem 4-10 X0·5 mm., distinct, straight or subflexuous: waxy-fleshy, rather
tough.
On the ground in the forest: l\falaya (Tembeling, Pahang, 3.12.30).
Spores 4·5-5·5(-6) X2·5-3 µ, white, smooth, oblong-ellipsoid to pip-shaped, thin
walled (? guttate).
Basidia 20-25x5-6µ, slightly projecting: sterigmata 4, 3µ long.
Hymenium apparently not thickening: subhymenium 10-12µ thick, composed of
2-3 µ wide, short-celled hyphae.
Hyphae 2·5-8µ wide, the cells 50-150µ long, short-celled near the subhymenium,.
388 CLAVULINOPSIS
longitudinal, compact, thin-walled, clamped, without narrow interweaving hyphae:
cells 10-25 X 2-3µ on delimitation at the apex.
Stem glabrous, with a few, slightly thick-walled mycelial hyphae 1·5-2µ wide at
the base.
This undistinguished species comes near C.
000000 alcicornis and C. fruticula. It could be mistaken for
Clavaria vermicularis if the clamp-connexions were
overlooked. The spores are truly ellipsoid, not
subglobose as in C. biformis, near which it comes in
the Key.
C. spathuliformis (Bres.) comb. nov.
Basinym: Clavaria spathuliformis Bres., Sacc.,
Syll. Fung. 9, 1891, 250: le. Myc., t. uo5.
1-1·5 cm. X 2-4 mm., simple, clavate-spathulate,
compressed, stuffed, yellowish clay-colour: stem white,
0·7 mm. thick.
On earth in hothouses: Germany (Berlin Bot.
Gard.).
TuxT-FIG. 159. Clavulinopsis Spores 7-11 X 5-6µ, pale yellowish white, rough,
similis, X I: spores, X 1,000. ellipsoid, aguttate.
This is said to be allied with C. luticola. It seems nearer to Clavaria gracillima sensu
Wehm., with smaller asperulate spores. Whether they belong to Clavulinopsis or not,
cannot be decided until they have been thoroughly described. The aguttate spores of
C. spathuliformis do not suggest the genus, and why are they coloured? It may be a
tropical species introduced into the Berlin Botanic Garden.
C. spiralis (Jungh.) comb. nov.
Basinym: Clavaria spiralis Jungh., Verh. Bat. Genootsch. 17, 1839, 32.
Synonyms: C.filiformis P. Henn. et Nym., Mons. 1, 1900, 8: v. Ov., Bull. Jard. Bot.
Buit. Ser. 3, 5, 1923, 265: le. Fung. Mai. 3, 1923.
Pseudonyms: C. vermicularis Fr. sensu v. Ov., Bull. Jard. Bot. Buit. Ser. 3, 5, 1923,
268: le. Fung. Mai. 4, 1923 (non Fr.).
C. vermiculata (Mich.) ex v. Ov., sensu Petch, Ann. R. Bot. Gdn. Per. 9, 1925, 331
(non v. Ov. = Clavaria vermicularis Fr.).
Simple, rarely branched, caespitose or gregarious, rarely solitary, white, the tips
becoming yellowish with age, finally wholly pale yellowish when old and effete, sometimes
pale cream-white or pale yellowish white becoming deeper yellowish from the base upward,
rarely white then wholly pallid ochraceous.
1-14 cm. X 0·7-4 mm., or flattened 6-15 mm. wide, very variable in size, exactly as
C. amoena; cylindric or subfusiform and acute, becoming blunt and often compressed,
often longitudinally sulcate or rugulose, often twisted and flexuous, solid then hollow,
occasionally with 1-3 short branches: stem 2-15 x o· 5-1·5 mm., indistinct, subpellucid,
slightly villous at the base: flesh waxy, rather brittle, without smell or taste.
On the ground in the forest, or in shady places in the open: tropical Asia (Java,
Ceylon, Malaya: common).
Spores 5·5-7 X4·5-6·5µ, white, smooth, subglobose, thin-walled, with one large
gutta 4-5·5µ wide, becoming wholly vitreous opalescent, occasionally wholly vitreous
opalescent from the first.
Basidia 35-65 X 6-10µ clavate with a long stalk, granular-guttulate: sterigmata
2-3-4, 6-<)µ long.
Hymenium 40µ thick at first, thickening to 150µ (probably more in old specimens).
CLAVULINOPSIS
Hyphae, microscopic structure, and development as in C. amoena.
I have used this name, not without misgiving, for the white or pale yellow state of
C. amoena, as C. miniata is the pink or red state. They differ only in colour and can
be considered varieties of one species. But, until their pigmentation is understood
(see p. 381) and they are better and more widely known in other respects, it is more
convenient to separate them : each has its own set of variations which are better made
varieties than forms.
'TExT-FIG. 160. Clavulinopsis spiralis: variation in size and

habit of full-grown fruit-bodies, X x.
It is now impossible to identify Clavaria spiralis with certainty, but I agree with
van Overeem that it probably represents the old, much inflated, and twisted fruit
bodies of the Javanese fungus which he calls C. vermicularis. This is the commonest
simple 'white' Clavaria in Java and Malaya and the only one which assumes such
grotesque forms. It is not C. vermicularis, as Donk had reason to doubt, but a true
Clavulinopsis.
The original description of C. spiralis gives the colour as 'laete sulphurea' and the
spores as ochraceous. Old specimens of this white species often become pale yellow,
but the spores are never ochraceous. No such species has been recognized in recent
years in tropical Asia and one may question the existence of a simple Ramaria.
39CI CLAVULINOPSIS
C. spiralis might be interpreted as C. amoena, but this species (described as 'pulcherrime
aurea') answers better to the middle term of the series of white-yellow-orange-red
pink species of Clavulinopsis, as C. miniata is the end term. C. spiralis fits the first
two terms and, hence, my use of this old name rather than a new one.
C.filiformis is only a slender form of C. spiralis. Van Overeem fj.gures a clamp at the
base of the basidium, which shows that it is not C. vermicularis.
J1
..
' :�.
TExT-FlG. 161. Clavulinopsis spiralis: spores and basidia, X 1,000:

hyphae, X 500.
Clavulinopsis brevipes is, perhaps, a small solitary state of C. spiralis with large
sterigmata and more pip-shaped spores. It differs constantly in Singapore, for both
C. brevipes and C. spiralis (caespitose) occur every fungous-season in Bukit Timah
forest.
Probably most Asiatic records of C. vermicularis are really C. spiralis. It is remark
able that no temperate ally has been found, such as C. fusiformis to C. amoena, and
C. aurantio-cinnabarina to C. miniata.
C. subfastigiata (Britzm.) comb. nov.

Basinym: Clavaria subfastigiata Britzm., Hym. Siidb. 8, 1891, 14, f. 43, 44: Rev.
Hymen. 1909, 220.
4·5 cm. high, branched, yellow or reddish yellow, becoming white: stem distinct:
branches equal, erect or slightly curved, with irregular short or long teeth subfragile.
CLAVULINOPSIS 391
In pastures: southern Germany.
Spores 10 X 8µ, white, smooth, broadly ellipsoid, 1-guttate.
Near to C. corniculata but with slightly larger spores.
C. subflava (Britzm.) comb. nov.

Basinym: Clavaria subfiava Britzm., Hym. Siidb. 8, 1891, 14, f. 61: Rev. Hymen.
1909, 220.
Synonym: Clavaria extans Britzm., ibid.
-'J cm. high, branched, pale yellow: stem short, thick, -1 cm. wide, white, dividing
abruptly into elongate, filiform, furcate, erect branches 4-5 cm. long, with subacute
tips, bifid or multifid.
In woods: southern Germany.
Spores 6-8 X 3-4µ, white, smooth, shortly ellipsoid, aguttate, internally granulate.
I can see no difference in the descriptions of C. subjlava and C. extans except that
the colour of the spores is omitted from that of C. subfiava. They may be forms of
Ramariopsis Kunzei or R. clavuligera. Compare C. biformis (N. America).
C. subtilis (Fr.) comb. nov.

Basinym: Clavaria subtilis Fr., Syst. Myc. 1, 1821, 479: Lundell et Nannfeldt, Sv.
Svamp. 11-12, 1938, 18: (non Bres., le. Myc., t. 1099 = Ramariopsis Kunzei).
Synonyms: Clavariamacropus Fr., Syst. Myc. 1, 1821, 475: Pers., Myc. Eur. 1,
1822, 168: Bourd. et Galz., Hym. Fr. 1928, 104.
Ramaria subtilis (Fr.) Quel., FI. Myc. 1888, 464.
1·5-4 cm. high, branched, slender, caespitose, white then pale yellowish: stem 1-1·5 cm.
X 1-2 mm., equal, with few branches of equal size, sometimes elongate ( = forma
macropus Fr., teste Lundell): branches dichotomous, subfastigiate, subacute: flesh
white, tough.
Among grass and in woods: Europe (Sweden, France, Gt. Britain), U.S.A.
Spores 3·5-4·5 X 3-3·5µ, white, smooth, broadly ellipsoid to subovoid (Lundell):
4-6 X 3·5-4·5µ, subglobose (Bourd. et Galz.): 6 X 3µ, oblong ellipsoid, 1-guttate (Rea).
Basidia 30-36 x4·5-6µ, with 2 sterigmata 6µ long (Bourd. et Galz.).
Hyphae 3-12µ wide, thin-walled (Bourd. et Galz.): clamped (Corner, for C.
macropus, Bourd., No. 25767, herb. Paris.).
I have followed Lundell's redefinition of this species, for which there is no type
specimen. It resembles a white C. corniculata, with slightly smaller spores. C. dicho
toma seems to be identical, or a form with slightly different shape to the branches and
with larger spores than given by Lundell. Clavaria chionea sensu Rea, with tough
flesh, may also belong here. Bourdot and Galzin distinguigh C. ma::ropus because of
its very minutely rough spores. I have examined an authentic collection of theirs
(Bourd. 25767) and could not convince myself that the spores were rough. They
were, as Bourdot gives, 4-5 X 3-3·5µ, as in C. subtilis, but Bourdot says that, when
fresh, they are 7-8 X 4-6µ: I do not understand the difference. It is doubtful, also,
whether the basidia are always 2-spored: the point is not mentioned for C. macropus
sensu Bourd. et Galz., but if they are 2-4-spored, the difference will explain the
variation in spore-size.
C. subtilis sensu Oud. (1902) and sensu Wehm. seem to be C. tenuicula, under
which I have discussed their collections.
Coker found the spores on a Friesian specimen of C. macropus (from Upsala, at
Kew) to be smooth, ovoid, 8-8·5 x6·5-'7µ, as in Clavulina cristata, and he follows
Bresadola in reducing C. macropus Pers. sensu Fr. to Clavulina cristata. Personally,
I can see no value in such a specimen: it is not a type, and it is impossible to see
how Fries, Persoon, or any mycologist could distinguish macroscopically, as was
their means, small, branched, white Clavarias accurately enough for the modern
392 CLAVULINOPSIS
microscopic criteria of species. I am sure that Fries, and many other mycologists,
confused Clavulina cristata, Ramariopsis Kunzei, and Clavulinopsis subtilis.
Compare C. biformis (N. America) and C. Jruticula (Malaya).
C. subumbrinella (Imai) comb. nov.

Basinym: Clavaria subumbrinella Imai, Tr. Sapporo Nat. Hist. Soc. 13, 1934, 386.
-5 cm. high, branched 3-4 times, solitary, umber or tawny-olive: stem distinct, white
tomentose in the subterranean part: branches dichotomous, tips acute.
On the ground in woods: Japan (Nopporo, Prov. Ishikari).
Spores 5 X 3 µ, white, broadly ellipsoid, smooth.
Basidia clavate.
This is said to be near C. umbrinella and to differ in the slightly tomentose, distinct
stem and the mode of branching, perhaps also in the more ellipsoid spore.
C. sulphurascens (Schw.) comb. nov.

Basinym: Clavaria sulphurascens Schw., Tr. Am. Phil. Soc. 4, 1832, 182.
15 mm. high, delicate, sulphur-yellow then subfuliginous: stem cylindric, the base
thickened and white pruinose, with a byssoid rhizomorph: branches subfastigiate,
cylindric, short, corniculate, acute.
On dead leaves: U.S.A.
Doubtful, no specimen existing. Coker suggests Ramariopsis crocea.
C. tenella (Boud.} comb. nov.

Basinym: Clavaria tenella Boud., Bull. Soc. Myc. Fr. 33, 1917, II, t. 4, f. 1: Bourd.
et Galz., Hym. Fr. 1928.
3-4 cm. high, branched, solitary or subcaespitose, slender, pale ochraceous grey,
white at the base: stem 1-1·5 mm. wide, simple or divided at the summit into 2-3
cylindric, pointed, erect branches: hymenium not distinct from the stem.
On the ground: France (rare).
Spores 5-6µ wide, white, smooth, ovoid-spheric, 1-guttate.
Coker suggests this is the subsimple state of C. corniculata.
C. tenerrima (Mass. et Cross!.) comb. nov.

Basinym: Clavaria tenerrima Mass. et Cross!., Naturalist 1904, 2.
1·5-4 cm. X 1-2 mm., simple, gregarious, cylindric, fiexuous, slender, subacute, not
narrowed at the base, stuffed, firm, hyaline or shining white.
On the ground among grass: Gt. Britain (rare).
Spores 8-9µ, white, verrucose, subglobose.
Basidia with 4 sterigmata.
Cotton and Wakefield suggest this may be Clavaria acuta. On the other hand, it
may be an immature state of C. asterospora.
C. tenuicula (Bourd. et Galz.) comb. nov.

Basinym: Clavaria tenuicula Bourd. et Galz., Hym. Fr. 1928, .104.
1-2·5 cm. high, branched, slender, white or tinged purplish grey at the tips: stem
0·5-1 mm. wide: branches from the middle or near the base of the stem, simple or
branched, flexuous, divaricate, acute, few.
On bare earth and humus: France (uncommon: Bourd. 25764), Holland, Canada.
Spores 3-3·5 X 2·5-3 µ., white, smooth, subglobose, r-guttate.
Basidia 14-18 X 4-4·5 µ..
Hyphae 2-5 µ. wide, thin-walled, not inflated, clamped.
I have examined the type-collection, on loan from the Paris herbarium, and find
that it is a true Clavulinopsis with uninflated hyphae as occur in C. minutula and
C. propera. It differs from C. ,minutula only in the slightly larger spores (2·5-3 X
CLAVULINOPSIS 393
2·5-2·8µ in C. minutula) and seems hardly worth distinguishing. They are, perhaps,
diminutive uninflated derivations of C. subtilis, unless the narrow hyphae indicate
affinity with C. propera.
Because of their small spores and small sparingly branched fruit-bodies, I refer
here C. subtilis sensu Oud. and sensu Wehm. Their descriptions are:
C. subtilis sensu Oud., Arch. Nederl. Kruidk. Ser. 3, 2, 1903, 674: 1-z cm. high,
terrestrial, simple or branched once or twice, 0·5-1·5 mm. thick, slender, white or
pale greyish, branches reaching about the same level: spores 3 ·5 X2µ, white, smooth:
Holland.
C. subtilis sensu Wehm., Pap. Mich. Ac. Sci. Arts Lett. 20, 1935, 257: 1-2 cm.
high, white, occasionally simple, generally branched one to three times in antler
fashion: stem 5-12 Xo· 5-1 mm.: branches dichotomous or fascicled, 0·2-1 mm. wide,
often flattened, with rounded axils, lax: spores 3-3· 5 X2,-3µ, white, subglobose,
smooth; on moss or humus: Nova Scotia.
C. tetragona (Schw.) comb. nov.
Basinym: Clavaria tetragona Schw., Naturf. Ges. Leipz. Sehr. 1, 1822, II2.
3·5 cm. high, subsimple, yellow, fragile: stem furcate or bifurcate at the apex,
quadrangular.
On the ground: U.S.A.
Very doubtful, no specimen existing. Coker suggests Ramariopsis crocea.
C. umbrina (Lev.) comb. nov.

Basinym: Clavaria umbrina Lev., Ann. Sc. Nat. Ser. 3, 5, 1846, 155.
Synonyms: Clavariafurcata P. Henn. et Nym., Mons. 1, 1900, 9 (teste v. Ov.).
Clavulina umbrina (Lev.) v. Ov., Bull. Jard. Bot. Buit. Ser. 3, 5, 1923: le. Fung.
Mai. 2, 1923.
--7 cm. high, branched one to three times, often irregularly, solitary, gregarious or
caespitose, purplish or purplish brown to umber, yellowish towards the golden-yellow tips;
stem 1-3 cm., or more, cylindric, often paler or yellowish: branches 1-4 mm. wide,
more or less regularly dichotomous, somewhat divergent with semilunar axils, often
rather flexuous, often flattened at the axils, tips subacute or blunt.
On the ground: Java (Buitenzorg Botanic Gardens).
Spores 5--7(-9) µ. wide, white or stained yellow, smooth, subglobose, thick-walled,
1-guttate.
Basidia 46-58 X5·7-8·5µ, elongate-stipitate below, with yellowish guttulae: sterig
mata 4, occasionally 2, 10-14µ long.
Macroscopically and microscopically this is very like intensely coloured and old
specimens of Clavulina Leveillei, but if it has 4 sterigmata, as van Overeem describes,
it cannot be Clavulina. If Clavulinopsis, it must be allied with C. umbrinella and
C. corniculata.
C. umbrinella (Sacc.) comb. nov.
Basinym: Clavaria umbrinella Sacc., Syll. Fung. 6, 1888, 695.
Synonym: Clavaria umbrina B., Outl. 1860, 279, t. 18, f. 4 (non C. umbrina Lev.
1846): Rea, Tr. Brit. Myc. Soc. 3, 1909, t. u, f. e.
Plate 10.
2-4·5 cm. high, branched, white then pale brown to umber, the tips often darkest:
stem short, white, villous: branches fastigiate below, polychotomous, then irregularly
dichotomous, 1-2·5 mm. wide, blunt, the axils acute below, more or less lunate above:
flesh hard, firm, white then pale brownish: smell none, taste pleasant.
Among grass: Gt. Britain, France (uncommon).
Spores 4-6·7 X3-6µ, white, smooth, subglobose or broadly pip-shaped, 1-guttate,
the apiculus 1-1·5µ long: 4-6x3-4µ (Cotton): 5-6x4-5µ (Coker, type-specimen):
394 CLAVULINOPSIS
4-4·5 X3-3·5µ (Corner, spec. in herb. Kew.): 5·5-6·7 x 5-6µ (Corner, living speci
mens).
Basidia 70-95 X 8-9µ, very long, tapered to a narrow base 2-2·5 µ wide (35-40 X 5--7 µ,
auctt.): sterigmata 4, 8-10 µ long.
Hymenium thickening to-150µ: cystidia none.
Hyphae 2·5-9µ wide, clamped, long-celled, little inflated, with firm, slightly
thickened walls: 1·5-2 µ wide on the surface of the stem, unbranched, aseptate, with
thickened walls.
Brown colour in the cytoplasm of the subhymenial and outer longitudinal hyphae,
very pale.
This species is clearly related to C. corniculata and differs chiefly in the lack of
yellow ground colour and in the slowly developing umber pigment: it also _has much
firmer flesh and remarkably ioi'lg basidia. Possibly it is the coloured precursor of the
C. dichotoma-C. subtilis series.
The S. Australian specimens, \Teferred to C. umbrinella by Fawcett, seem to me to
be the species which she had distinguished as Clavaria cinnamomea (see Clavulinopsis
cinnamomea): their colour is not that of C. umbrinella and their spores are too small,
though there is evidently considerable range in those of C. umbrinella.
C. vernalis (Schw.) comb. nov.
Basinym: Clavaria vernalis Schw., Sehr. Nat. Ges. Leipz. 1, 1822, II2: Coker,
Clav. U.S. Can. 1923, 35, t. 82, 92: Journ. EI. Mitch. Sci. Soc. 55, 1939, 382, t. 41.
Synonyms: Clavaria clavata Pk., Bull. Buff. Soc. Nat. Sci. 1, 1873, 62 (teste Coker).
C. paludicola Lib., PI. Crypt. Arduenn. 1837, No. 322 (teste Coker et Donk).
7-12 X 1-2 mm., simple, rarely with a short branch, gregarious in extensive colonies,
clavate, obtuse or abruptly acute with a sterile tip, watery ochraceous to dull orange
(Coker), yellow drying orange (Peck), pinkish cinnamon above, paler below to the
whitish base (Burt): stem 2-2·5 mm. long, poorly defined, whitish, translucent, glabrous:
flesh pliable, rather tough: smell none: taste slightly mouldy: the sterile tip covered
with crystals when dried.
On bare earth covered with Cyanophyceae and moss protonemata: U.S.A. (N. Car.,
N.Y., Mass., N. Jers.: uncommon): Europe (Holland, France).
Spores 8-1 I X 2-3µ, white, smooth, narrowly ellipsoid, slightly curved, aguttate,
or (Donk) 1-few guttulae.
Basidia 18-25 X 4-5 µ (Coker), 25-35 X 6-8 µ (Donk): sterigmata 2-4, 3-5 µ long.
Hyphae 2--7·5(-u)µ wide, clamped.
This is a phycophilous species. Coker found that it always grew from a mass of
Cyanophyceae (Gloeocapsa, Oscillatoria, &c.) with its hyphae closely applied to
the algal cells. It has clamp-connexions (as I have proved to my satisfaction from
the examination of dried specimens in the Kew herbarium), and thus belongs in the
affinity of C. calocera and C. hastula, also with rather tough or gelatinous texture,
instead of the affinity of Clavaria helicoides or Clavaria fossicola. Whether it is related
to Leiztaria mucida ( = Clavaria mucida) is doubtful, but Clavaria mucida var. rosea
Bres. may be identical. Clavulinopsis constans may be the same: possibly also Pistillaria
gilva.
The sterile tip is the growing point, which may often wither before maturity.
DEFLEXULA gen. nov.

(From the down-curving of the Pteruloid fruit-bodies)
Fruit-bodies as small clusters of simple or sparingly branched, dqwn-pointing spines,
each cluster developing from an initial spine by adventitious proliferation of new spines
from its base, branching wholly adventitious, the subulate or filiform tips not dividing,
without resupinate patch or mycelial subiculum: flesh firm, tough: superficial, ligni
colous.
DEFLEXULA 395
On dead bark and wood: 5 spp., tropical Asia, Australasia, N. and tropical America.
Spores white, smooth, globose or ellipsoid, large, slightly thick-walled, aguttate.
Basidia large, clavate: sterigmata 4.
Cystidia none, but often with narrow excrescent hyphae, 1·5-3 µ. wide, scattered
among the basidia.
Hymenium thickening: subhymenium composed of narrow uninflated hyphae.
Hyphae as in Pterula, dimitic: skeletal hyphae of unlimited growth, thick-walled,
rarely branched, aseptate, occasionally proliferating skeletals or generative hyphae
from the apex: generative hyphae clamped, occasionally with intercalary, thick-walled,
skeletal portions.
Stem with a short sterile base covered with narrow generative hyphae (proliferating
the lateral spines) and with a few sterile basidia in the transition to the fertile region.
Type-species: D. fascicularis.
KEY TO THE SPECIES OF DEFLEXULA

(Compare Mucronella and Pterula)
Spines short, -3 mm. long, often lilaceous or violet
Sp. 14-16 X 10-12µ, ellipsoid: spines dull violet then white:
U.S.A. . D. ulmi
Sp. 12-14µ wide, globose: spines white then yellowish, brownish,
or lilaceous: Malaya, Sumatra D. lilaceo-
brunnea
(Sp. 18-2ox10-12µ: -1 mm. high: Ecuador Pterula macro-
spora)
Spines 1-2·5 cm. long, not lilaceous
Sp. 9-10µ (? -12µ) wide, globose: spines -18 mm. long, often
branched, even to the second degree, white then pallid
ochraceous or tinged flesh colour: Malaya, Philippines, Samoa D. fascicularis
Sp. ellipsoid
Sp. 10-14x8µ: spines -18 mm. longx300µ wide, white:
Guadeloupe D. nivea
Sp. 12-13·5x6-7µ: spines -25 mm. long, white or sub-
ochraceous: Bonin Is!. D. pacifica
D. fascicularis (Bres. et Pat.) comb. nov.
Basinym: Pterula fascicularis Bres. et Pat., in Lloyd, Myc. Writ. I, 1901, 50: Lloyd,
Myc. Notes 60, 1919, 865, f. 1470.
Synonym: Mucronella fascicularis sensu Lloyd, non sensu Bres. [M. fascicularis
Fr. = Protohydnum fasciculare (Fr.) Bres.: Lloyd confused this fungus repeatedly
with his Samoan plant Pterula fascicularis].
Text-Figs. 162, 163: Plate 1 I.
Fruit-body -18 mm. long, 0·5-1 mm. wide at the base, fasciculate in clusters of two
to fifteen, the clusters generally gregarious, rarely solitary, pointing or curving vertically
downward, simple or sparingly branched, white then pale ochraceous, finally pale dingy
ochraceous flesh colour.
Stem (if unbranched) and branches tapering to subulate or shortly filiform tips
15-80µ wide; no mycelial subiculum: branches 1-5, simple or with 1-2 branchlets,
arising from the base or along the sides of the axis and appearing pinnately secund, the
tips of the axis and branches never dividing: hymenium amphigenous, generally
absent from the base of the stem for a length of 1-2 mm.: flesh tough, pliant; no smell.
On dead wood in the forest: Samoa, Philippine Is!., Malaya (Singapore, Bukit
Timah Forest Reserve, 7-4-41).
DEFLEXULA
Spores g-10·5 X9-10µ. (9-12µ., Bres. et Pat.: 8-10µ., Lloyd), white, smooth,
gl.obose, with a short apiculus 1-1·5µ. long, thin- or very slightly thick-walled, contents
cloudy-vacuolate, aguttate.
Basidia 25-40(-50) X 8-II µ., clavate, sometimes with a long narrow stalk (having
grown up from a lower' layer of the subhymenium), vacuolate-cloudy: sterigmata 4
(? mature length).
Cystidia none.
TExT-FIG. 162. Defle:x:ula fascicularis, x 2.
TExT-FIG. 163. Defle:x:ulafascietdaris: spores and sterile(? daytime) hymenium, X 500.

Hymenium thickening -150 µ., with embedded spores: subhymenium c. 30 µ. thick,
composed of closely interwoven or contiguous, thin-walled, uninflated, short-celled
hyphae 1·5-2·5µ. wide.
Hyphae dimitic: skeletal hyphae 3-4µ. wide, -6 µ. in places, with smooth walls
0·5-1 µ. thick, aseptate, often of uneven thickness, often kinked, of unlixnited growth,
occasionally proliferating skeletal or generative hyphae from the apex, longitudinal:
generative hyphae 1·5-3·5µ. wide, thin-walled, clamped, sometimes branching from
the clamp.
Base of the stem, for a length of 1-2 mm., covered with very narrow, shortly
DEFLEXULA 397
excrescent hyphae 1·5-2µ, wide, simple or irregularly branched, with scattered sterile
basidia: no caulocystidia: no palisade-like sterile hym enium.
Mycelial hyphae 1·5-2·5µ, wide, clamped, with thin or slightly thickened walls.
This species agrees exactly with the description and illustration of Pterula fascicularis
given by Lloyd. It has no connexion with Mucronella fascicularis Fr., which Bresadola
has shown to be Tremellaceous and has called Protohydnum fasciculare (Ann. Myc. 18,
1926, 62: le. Myc., t. n17).
The Malayan specimens, which I examined a few hours after collection, had no
sporing basidia, yet they appeared to be in full vigour. During the night they gave a
spore-print on a glass slide, but next day I could again find no sporing basidia.
Unfortunately, at that time, I had no leisure to examine them further. Among the
basidia I saw scattered or fairly abundant, narrow, sterile hyphal tips 1-3µ, wide,
unbranched or branched or irregularly lobed at the apex, reaching the surface of the
hymenium and projecting up to 20µ,, sometimes branched and lobed above the sur
face. They are clearly homologous with the pleurocystidia which Kobaiyashi has
described for D. pacifica, but I think they are merely the excrescent hyphae which will
build a new layer of basidia and not cystidia or paraphyses like those of Aleurodiscus.
The fruit-bodies develop as in D. lilaceo-brunnea, but they often produce adventi
tious branches above the base and the secondary spines seem to have a longer ageo
tropic or diageotropic period of growth.
D. lilaceo-brunnea sp. nov.

-3·5 X 0·2-0·5 mm.,fasciculate in clusters of 2-20 down-curved or downward pointing,
subulate spines, simple or with I-J branches from the base, the larger spines in the centre
of the tuft, the outer ones short and young, entire, very acute, 25-55µ, wide at the
tip, tapered from base to apex, white then yellowish, pale fawn-brown, lilac, or sub
violaceous: flesh tough: no smell.
On the underside of dead branches, densely gregarious: Malaya (Singapore, Bukit
Timah Forest Reserve, 17.4.43), Sumatra (fide Donk, in litt.).
Spores 12-14µ, wide, white, smooth, globose or slightly wider than long, wall
o·8µ, thick, with cloudy vacuolate coments, aguttate.
Basidia 50-75 X 15-18 µ, ; clavate, large, vacuolate-cloudy: sterigmata 4, 9µ, long,
stout.
Cystidia none.
Hym enium thickening slightly -120 µ,, the old basidia collapsing: subhymenium
composed of loosely interwoven, thin-walled, hyphae with cells 10-20 X 2-5µ,, the
layer initially c. 1oµ, thick.
Hyphae dimitic: skeletal hyphae 3-5(-7)µ, wide, with smooth, colourless or pale
yellowish walls 1-2µ, thick, often kinked, occasionally forked, some of limited growth
(300-700µ, long) with simple or slightly tapered ends, others traversing the whole
length of the spine: generative hyphae 2-5µ, wide, thin-walled, rarely thick-walled,
septate with clamps, the cells 15-200µ, long, the branches sometimes constricted at
the. base: with occasional transitions between generative and skeletal hyphae, and the
skeletals occasionally proliferating at the apex.
Base of stem sterile as in D. fascicularis.
This species has essentially the same structure as D. fascicularis but differs in its
smaller, less branched, and differently coloured fruit-bodies with larger spores and
basidia. It is very near D. ulmi, with ellipsoid spores, from N. America.
The fruit-bodies originate as superficial and discrete, conical, down-pointing,
primordia 20-50µ, wide, composed of 2-3µ, wide, generative hyphae which have
emerged from the underside of the branch or twig in which the mycelium was growing.
Each primordium develops slowly by apical growth into a subulate spine which
proliferates new spines adventitiously from the base and thus builds the fasciculate
3 98 DEFLEXULA
cluster. When the initial spine is 1-1·5 mm. long, a second primordium develops
from,the narrow generative hyphae covering its sterile base: it grows outward and then
curves down. Other primordia are continually proliferated from the base of the
initial spine and its branches, and they also grow outwards, sometimes radiating for
V
1-2·5 mm., before they curve down. The projecting part of the spine is generally
V V
TEXT-FIG. 164. Defiexula lilaceo-brunnea, X 10.
simple, but occasionally it branches by the formation of a new growing-point adventi

tiously from the side, and probably from the subhymenial hyphae. Thus each cluster
of spines is built up from an initial primordium. There is no resupinate-patch or
mycelial subiculum at the base of the cluster, and there is no apical branching.
D. nivea (Pat.) comb. nov.

Basinym: Pterula nivea Pat., Bull. Soc. Myc. Fr. 18, 1902.
Spines 10-18 mm. long x 300 µ, wide, simple or fasciculately branched from the
base, cylindric, subulate, white, tough.
DEFLEXULA 399
On rotten stems: Guadeloupe.
Spores 10-14 X 8µ,, white, smooth.
Entirely covered with the hymenium.
Lloyd refers this to D. fascicularis, but the ellipsoid spores and very slender fruit
bodies, which are not branched except adventitiously from the base, indicate another
species.
TEXT-FIG. 165. Defiexula lilaceo-brunnea, X 1,000.
D. pacifica (Kobayashi) comb. nov.

Basinym: Mucronella pacifica Kobayashi, Bot. Mag. Tokyo 53, 1939, 160, f. 4.
Fruit-bodies -2·5 cm. x 0·3-1 mm. wide at the base, fasciculate in clusters of few
to many spines, down-curved without subiculum, arcuate-subulate, simple or forked at
the base, white then subochraceous, drying brownish: flesh firm, solid.
On dead trunks of Callicarpa subpubescens (Verbenaceae): Bonin lsl.
Spores 12-13·5 x 6·5-7µ,, white, smooth, ovoid, apiculate, slightly thick-walled,
aguttate.
Basidia 40-45 X 9-10 µ,, clavate: sterigmata 4.
Cystidia none, but a few filiform hyphae, 2µ, wide, occurring among the basidia
and projecting slightly beyond them.
Hyphae 3·5-4µ, wide, longitudinal.
From the shape and direction of the fruit-body there can be no doubt that this is a
species of Defiexula closely related to D.fascicularis and differing in its almost simple
spines and ellipsoid spores. Kobayashi described the narrow hyphae amon! the
basidia as pleurocystidia but, as I have remarked under D. fascicularis, I think they
are merely excrescent hyphae thickening the hymenium.
400 DEFLEXULA
D. ulmi (Pk.) comb. nov.
Basinym: Mucronella ulmi Pk., Rept. N.Y. St. Mus. 54, 1901, 154: Miller, Mycol.
26, 1934, t. 27, f. 4, 5.
Fruit-bodies consisting of clusters of 2-8 down-turned spines 1-3 X 0·2-0·35 mm.
(? initially single), subulate, acute, soft, dusty dull violet with a white base, soon white
and mealy: subiculum absent.
On bark of living and dead Salix, Quercus, Fraxinus, and Ulmus: U.S.A. (N.Y.,
Pa., Iowa), June-November.
Spores 14-16 x 10-12µ,, white, smooth, obovate, with a prominent apiculus.
Basidia 30-35 X 10-15µ,, clavate: sterigmata 4: accompanied by slender, hyaline,
and slightly projecting paraphysoid hyphae.
Hyphae 2·5-3·5µ, wide, smooth, 'walls thickened, with few septa and clamp
connections'.
This is clearly a Pteruloid fungus very near to D. lilaceo-brunnea, which has globose
spores, and, if Miller's measurements are correct, much shorter basidia. I think, how
ever, they may be found to be varieties of one widespread species. Miller figures the
skeletal and generative hyphae. He says that fresh young fruit-bodies are violet,
though they fade to white as described by Peck and Overholts. The paraphysoid
hyphae among the basidia are probably, as in D. fascicularis, the excrescent hyp hae
which thicken the hymenium.
DIMORPHOCYSTIS gen. nov.

(From the two forms of cystidium)
Fruit-body simple, stalked, white, puberulous: stem generally with scattered caulo
cystidia: fertile part filiform-attenuate to a sterile tip, becoming wholly fertile, or
shortly clavate or globose without a sterile tip, puberulous: flesh tough, rather horny
or cartilaginous: on dead leaves, erumpent.
On dead leaves: 3 spp., tropical Asia.
Spores white, smooth, ellipsoid, aguttate.
Basidia with 2-4 sterigmata.
Cystidia dimorphic: conical cystidia arising from the medullary hyp hae or from the
basal subhymenial hyphae, as the primary hymenial elements, traversing the hymenium
and subhymenium and projecting, thick-walled, often rugose or incrusted; lobed cystidia
more or less superficial and hymenial, slightly thick-walled, the distal half or third
irregularly digitate with simple or shortly lobed processes, often incrusted.
Hymenium not thickening: subhymenium often very slightly developed.
Caulocystidia monomorphic, conical, scattered, or absent.
Stem without a palisade or sterile cov�ring.
Hyphae dimitic as in Pterula: skeletal hyphae of limited growth, with tapered, simple
or branched ends: generative hyphae with or without clamps.
Type-species: D. laevis.
KEY TO THE SPECIES

Fruit-bodies -18 mm. high, erect: fertile part subulate-filiform,
with sterile apex, at l�ast when growing: cystidia smooth, not
incrusted: sp. u-14 X 4-5µ,: basidia 2-spored, occasionally
1-spored . D. laevis
Fruit-bodies -1 mm. high (rarely -4 mm. in D. subcapitatus),
growing perpendicularly from the substratum in any direction:
fertile part globose to shortly clavate, without sterile tip .
DIMORPHOCYSTIS 401
Fruit-body 0·5-1(-4) mm. high: fertile head often shortly
clavate: cystidia slightly incrusted: caulocystidia present:
sp. u-13X 3-4µ.: basidia z-spored D. subcapitatus
Fruit-body -0·6 mm. high: fertile head globose: cystidia thickly
incmsted: caulocystidia none: sp. 6-7Xz·5-3·5µ.: basidia
4-spored D. capitatus
(Compare Physalacria villosa, Ceylon)
D. capitatus spec. nov.

Text-Figs. 166, 167, 169, 170, 171.
Fruit-bodies -0·6 mm. high, gregarious, often closely, always distinct, erumpent:
stem 300-500X40-50µ., cylindric, glabrous, pellucid white, the base inserted (not
darkened): fertile head 140-180µ. wide (excluding cystidia), globose ar subglobose,
shining white, minutely hispid, solid, wholly covered by the hymenium.
c::::::'L1J.J_t,J_fjJj_J_tf.d,-oJJ_lt.-"'
n-.rr1Tr,rJ,,lTfl"'--1nff1T�
D.G<1t,.•�"tu.s
TEXT-FIG. 166. Dimorphocystis: fruit-bodies
on dead leaves (in section), X 5.
On either side of dead leaves of various trees, shrubs, and palms in the forest
(Durio, Eugenia, Agrostistachys, Oncosperma, Pinanga), probably common: Malaya
(Singapore, Bukit Timah Forest Reserve, Selitar Reservoir).
Spores 6-7 X z·5-3·5µ., white, smooth, thin-walled, ellipsoid pip-shaped, with a
slight apiculus, aguttate.
Basidia 18-z4 x 5-6µ., subclavate, vacuolate-opalescent: sterigmata 4, 4µ. long.
5119 D d
DIMORPHOCYSTIS
Conical cystidia 40-95 X 8-15µ, or with long filiform appendages -150µ long
overall, arising deeply in the subhymenium near the stem-apex, radiating through the
tissue of the head, projecting up to 50µ, fusiform, with strongly thickened colourless walls
3-6µ thick, tapered to the subacute thick-walled apex and to the base, the lumen
always open, the distal half or two-thirds rugose and incrusted with small crystals.
TEXT-FIG. 167. Dimorphocystis capitatus: showing the relation of the basidia

and two kinds of cystidia to the generative hyphae and the digitiform tips of
the skeletal hyphae: x 1,000.
Lobed cystidia r 8-40 X 5-8µ, arising from the subhymenium immediately below the
hymenium, not or barely projecting, subclavate or subventricose, with slightly thickened
walls 0·5-1 ·5 µ thick, the distal third set with few or rather many, short, subcylindric,
straight, sinuous or shortly lobed, protuberances 1-3 X r µ, also with slightly thickened
walls, the distal third becoming incrusted with small colourless crystals obscuring the
processes.
Hymenium c. 25 µ thick, not thickening, recurved on to the stem-apex: sub
hymenium composed of short-celled, interwoven, contigu ous, thin-walled hyphae
1·5-3µ wide, not distinguishable from the tissue of the head.
Caulocystidia none: the stem glabrous, without trace of sterile hymenium.
TExT-FIG. 168. Dimorphocystis laevis: a hymenial hypha with a
normal cystidium of each kind and a transitional form (on the
right): Xl,000.
TEXT-FIG. 169. Skeletal hyphal ends of Dimorphocystis:

left, D. laevis; centre, D. subcapitatus; right, D. capitatus:
X 500.
'TEx:T-FIG. 170. Dimorphocystis capitatus, the interhyphal air-spaces
shown in black, X 500.
DIMORPHOCYSTIS
Hyphae dimitic: skeletal hyphae 3-6µ wide, smooth, unbranched, very thick
walled and glassy, the lumen mostly obliterated, aseptate, traversing the stem longitudinally,
ending at the base of the pileus in simple, shortly tapered tips or, usually, with I-several
branches becoming thin-walled, septate and often transformed into short generative
hyphae: generative hyphae 1·5-3µ wide, thin-walled, with clamps, cells -150µ long
in the stem, short and closely interwoven in the head.
TEXT-FIG. 171. Dimorphocystis capitatus: diagrams of stages in the

development of the fruit-body, X c. 250: A, inception in the leaf-tissue:
B, C, the primordial shaft in upgrowth, the skeletal hyphae shown as
heavy lines: D, cessation of up growth, the skeletal tips branching and
producing the first lanceola'te cystidia: E, these first cystidia fulJ-grown,
the hymenium forming: F, the hymenium dilating the head, but yet im
mature: G, lobed cystidia and other lanceolate cystidia beginning to form
in the hymenium: H, fulJy mature head with both kinds of cystidia
displayed by the intercalary development of the hymenium.
Both stem and pileus are solid, without air spaces, and hence their translucency.
The stem consists mostly of skeletal hyphae which appear as homogeneous glassy
rods. The generative hyphae between them are difficult to find until the tissue is
splayed apart by tapping the coverslip. It can then be seen that the hyphae on the
surface are not agglutinated (as is generally their condition in Typhula and Pistillaria).
In the fertile head, the skeletal hyphae mostly revert to generate hyphae by branching
and becoming thin-walled, and these branches and the generative hyphae which have
grown up in the stem form the closely interwoven, thin-walled tissue of the head in
which there are practically no air-spaces.
Development of the fruit-body is direct, without inflation of the hyphae. A knot of
generative hyphae forms in the tissue of the leaf through which the hyphae break.
They then give rise to skeleta 1 hyphae which grow perpendicularly from the leaf
s1,1rface, apparently without geotropic direction, and form the short stem. A small
number of generative hyphae grow closely round the base of the primordial shaft to
form its very slightly dilated support in the tissue of the leaf. On cessation of apical
growth of the stem, the generative hyphae and the branches of the skeletals form a
compactly interwoven, short-celled tissue, and their tips grow out in all directions to
form the fertile head which is, at first, discoid but becomes globose with the sides
DIMORPHOCYSTIS
recurved on the stem as it dilates by branching of the hyphae. The first distinct
elements to be formed in the head are the conical cystidia which arise from inflating
hyphal ends as soon as the tips of the skeletals begin to branch: these hyphal ends
grow out rapidly to 50-150µ, their walls thicken, and they become transformed into
the conical cystidia. The other excrescent hyphal tips grow outward between the
bases of the conical cystidia which are thus splayed apart and become immersed in the
tissue. At a certain period of outgrowth, all the hyphal tips enlarge into the lobed
cystidia or the first-formed basidia, generally in the centre of the head before the sides,
and by ordinary sympodial branching new basidia are intercalated. It is mainly the
inflation of the basidia and the intercalation of new ones that cause the head to become
globose with the sides recurved against the stem, and that cause the conical cystidia
to radiate in all directions by forcing them apart. The conical cystidia may arise
directly from the skeletal branches or after the inter-position of 2-3 short cells, and
as often, perhaps, from the generative hyphae which have grown up in the stem: the
bases of the conical cystidia are sinuous and it is difficult to trace them to their origins.
This remarkable fungus resembles Pistillaria capitata, but microscopically it is
found to be extraordinarily complicated with dimitic hyphae, which do not inflate,
and with dimorphic cystidia. It would be exceedingly difficult to classify, were it not
for the other species D. subcapitatus and D. laevis which show that it is derived from
the Pistillarioid Pterulas by limitation of the axis to a single short set of skeletal hyphae.
Thus D. capitatus and Pistillaria capitata are examples of convergent evolution by
degeneration of the Clavarioid fruit-body. In detail, D. capitatus has a solid head
without the distinct groove between it and the stem which occurs in P. capitata, the
head of which is largely filled by air and traversed by inflated monomitic hyphal
branches.
D. laevis sp. nov.

Text-Figs. 166, 168, 169, 172.
Fruit-bodies -18 mm. high, simple, scattered, white, puberulous, erumpent: stem
2-5 mm. X 200-280 µ wide at the base, subpellucid, inserted at the base, gradually
tapered into the opaque white cylindric fertile part 150-200 µ wide, the tip acute and
sterile, sometimes becoming fertile over the tip: rather tough and horny.
On dead leaves of Eugenia cerina (Myrtaceae): Malaya (Singapore, Botanic
Gardens, 16:3.43, 26.11.43).
Spores 11-14(-16) X 4-5µ, white, smooth, thin-walled, subcylindric, blunt,
obliquely apiculate at the base, cloudy vacuolate, aguttate.
Basidia 18-24 X 7-8µ, subclavate hyaline-vacuolate: sterigmata 2, 6-10µ long, IL
occasionally only 1-sterigma 7-10µ long: (without clamps).
Conical cystidia 40-75 X 10-18 µ, projecting far beyond the basidia, occasionally
only 15-25 x 3-6µ and immersed, conical or ventricose with narrowed or dilated
triangular base, acute or subacute, with slightly or distinctly thickened walls 0·5-2
(-3)µ, smooth, rarely slightly roughened.
Lobed cystidia 20-35 X 5-12µ, cylindric or subventricose below, the distal part (for
10-12µ from the apex) set with short simple or lobed, blunt processes -3 ·5 x 1 ·5 µ,
with slightly thickened walls 0·5-1·5µ, smooth, abundant: with occasional transitions
to the conical cystidia having a few short lobes.
Hymenium not thickening, seated directly on the superficial longitudinal generative
hyphae with short cells, 5-12 X 2-4µ, occasionally with a short subterminal cell below
a basidium or lobed cystidium: subhymenium practically none: developing at the
stem-apex by the gradual enlargement of the short blunt processes on the stem into
basidia.
Caulocystidia -50 X 5-12µ, like the conical cystidia but shorter, narrower, and
slightly thicker-walled, generally acute, scattered, generally absent from the extreme
base of the stem.
ThxT-FIG. 172. Dimorphocystis laevis: parts of the stem and fertil<; head, the air-spaces shown
in black, X 500.
DIMORPHOCYSTIS
Stem with a few short processes from the superficial generative hyphae, as well as
the scattered caulocystidia, but no palisade-like layer or sterile hymenium.
Hyphae dimitic: skeletal hyphae 4-8µ, wide, aseptate, unbranched, colourless, the
walls 0·5-3µ, thick, occasionally nearly solid and appearing glassy, of limited growth,
certainly 1-2 mm. long (even more), but always ending in tapering blunt points 1-3µ,
wide, the tapering part 150-450µ, long: the mediate (basal) parts of the skeletals
4-5µ, wide on origin and thin-walled: generative hyphae 2-5µ, wide, with thin or
slightly thickened (0·5µ,) walls, without clamps, the cells -200µ, long, generally slightly
constricted at the septa, often with slightly undulate walls.
D. subcapitatus sp. nov.

Text-Figs. 166, 169, 173.
Fruit-bodies 0·5-! mm. high, occasionally -4 mm. high, erumpent, rather densely
gregarious, not caespitose, simple, white: stem 200-800 X30-70µ,, occasionally to
3·5 mm. Jong, but equally slender, slightly enlarged at the inserted base, 40-130µ,
wide, straight, cylindric, puberulous, pellucid white: fertile head 90-350 X130-250µ,
wide (excluding cystidia), globose to narrowly clavate, puberulous, opaque shining white.
On the underside of dead leaves of Eugenia cerina (Myrtaceae): Malaya (Singapore,
Botanic Gardens, 17.3.43, 27.3.43).
Spores rr-13 X3-4(-4·5)µ,, white, smooth, thin-walled, cylindric, blunt at the
apex, shortly apiculate at the base, cloudy-vacuolate, aguttate.
Basidia 20-28 X6-7µ,, subclavate, hyaline: sterigmata 2, 5-7µ, long, occasionally
only 1 sterigma 7-10µ, long.
Conical cystidia 30-75 X9-15µ,, conical-ventricose, acute, the walls 1-4µ, thick,
generally thickest in the distal half; the narrow tip often nearly solid, becoming slightly
and finely rugulose, not incrusted, colourless, hyaline, projecting.
Lobed cystidia 18-35 X5-9µ,, cylindric or subventricose, set with simple or lobed
blunt processes -3·5 X1·5µ, in the distal half or third, slightly thick-walled, generally
slightly incrusted.
Hymenium not thickening, sometimes extended shortly down one side of the stem
or Jacking from one side of the head: subhymenium compact, distinct, with short
uninflated cells 5-11 X2-4µ,.
Caulocystidia -70 X4-7µ,, narrowly conical, acute, not incrusted, often very slightly
rugulose, the walls 1-1·5µ, thick, scattered, generally absent from the base of the stem.
Stem without sterile hymenium or a covering of hyphae.
Hyphae dimitic: skeletal hyphae 4-8µ, wide, the walls 0·5-3µ, thick, often nearly
solid and glassy, aseptate, unbranched, of limited growth, traversing the stem longi
tudinally and ending in tapering, 1-3µ, wide, simple or 2-3-fid tips in the head, often
dilated 6-12µ, wide at the base of the stem on origin from the generative hyphae:
generative hyphae 2-4µ, wide, without clamps, cells -150µ, Jong, thin-walled, little
branched in the stem, profusely branched and short-celled in the head.
This species combines the characters of D. laevis and D. capitatus. It resembles
D. laevis in its spores, 2-spored basidia, caulotystidia (which are narrower), the
tendency of the stem to elongate and of the hymenium to develop over the distal part
of the stem, not merely at the apex, so that the hymenium is not discoid at first: its
conical cystidia are also not deeply immersed. On the other hand, as in D. capitatus,
the stem is generally short and limited to the length of one set of skeletals, the fertile
part is also reduced, the conical cystidia are rugulose, and the lobed cystidia slightly
incrusted.
In the occasional fruit-bodies of D. subcapitatus with elongate stem, the skeletals
are more elongate, 1-2 mm. long as in D. laevis, but those which end at the apex of
the fruit-body have the characteristic lobed ends of D. subcapitatus. The elongated
fruit-bodies have a small capitate or subclavate head of normal size and long sterile
stem occasionally with discontinuous patches of hymenium on the sides.
ThxT-FIG. 173. Dimorphocystis subcapitatus: fruit-body with the air-spaces in black, x 500;
spores, X 1,000.
410 DIMORPHOCYSTIS
The fruit-bodies of D. subcapitatus grow on the underside of the same leaf on which
the fruit-bodies of D. laevis grow from the upperside. It is thus possible that they are
two unexpectedly different states of the same fungus, D. laevis being negatively
geotropic, long and of scattered habit, and D. subcapitatus positively geotropic, short
and of rather crowded habit. I saw no intermediate states.
HORMOMITARIA gen. nov.

(Hormos =necklace: mitos =thread: from the moniliform rows of secondarily septate
hyphae in the medulla of the fertile club.)
Fruit-body simple, pendent, with a short stalk and conical fertile club tapering to the
apex, the base of the fertile club slightly expanded and sterile: flesh waxy-firm, solid:
lignicolous.
Basidia small and narrowly clavate, c. 16 X 5 fl,; sterigmata 4, straight.
TEXT-FIG. 174. Hormomitaria sulphurea, X 10:

developmental sections, X 25.
Cystidia as oleocystidia, immersed in the subhymenium or reaching the surface of the

hymenium but not projecting, thin-walled, clavate, capped with an oily-resinous exr:retion.
Hymenium not thickening, eventually covering the tip of the fertile club: sub
hymenium rather thick, waxy, mucilaginous, the hyphae interwoven and not inflated.
Sterile hymenium at the base of the fertile club composed of a palisade of inflated
cells like the caulocystidia gradually passing into the fertile hymenium.
Caulocystidia abundant, as a loose palisade, colourless.
Hyphae monomitic, clamped, inflating, becoming secondarily septate and forming in the
fertile club short curved rows of moniliform cells, with rather toughly mucilaginous walls,
without air-spaces in the tissue.
HORMOMITARIA 4II
Development gymnocarpic and semi-indirect, a minute capitate primordium in

flating into the mature fruit-body with a small amount of concomitant apical growth
of the fertile club.
I 'I
\'.
�\:i<
. .. 1 .
·).....;
. 'j\
1:
I
I
,I
I·
1.
)
TEXT-FIG. 175. Hormomitaria sulphurea: hyphae of the head showing the rows of cells formed
by secondary septation: X 500.
On page 97 I have explained the close relation of this new genus with Physalacria.
They agree in general construction of the fruit-body with more or less indirect develop
ment, in the spores, in the caulocystidia and, particularly, in the subhymenial oleo
cystidia. Hormomitaria differs in the prolonged, positively geotropic, apical growth
of its head, which is therefore not inflated and hollow, and in the secondary septation
of the hyp hae. Without Hormomitaria, the interpretation of Physalacria would be
extremely difficult, if not impossible; yet Hormomitaria itself seems isolated among
Clavarioid fungi. The one species, H. sulphurea, appears to the naked eye as a long
spined Acia; under a hand-lens it is seen to consist of crowded, but separately stalked,
spine-like fruit-bodies, as Mucronella; under the microscope it is found to have the
structure of Pistillaria, though inverted, the secondary septation of Clavaria, the
412 HORMOMITARIA
oleocystidia and sterile base to the hymenium as in Physalacria, and the subagglutinated
hyphae of the Clavariadelphus-series. If its fruit-bodies were erect it would be placed
in Pistillaria. The inversion, therefore, seems analogous with that of Deflexula in the
Pteruloid-series. The resemblance to Hydnoid fungi is entirely superficial for such
degenerate into resupinate Corticioid states (as described under Mucronella). One
must postulate for Hormomitaria, therefore, yet another Clavarioid-series which, on
the evidence of Physalacria, may be the basis of the Marasmioid-series. Possibly
Mucronella tenuipes is another of the 'tropical surprises' belonging to this series.
TEXT-FIG. 176. Hormomitaria sulphurea: base of the stem, x 500.
H. sulphurea sp. nov.

Fruit-bodies -12 mm. long, shortly stip1tate, pendent, densely gregarious: stem
0·3-0·8 Xo·15-0·3 mm., cylindric, slender, abruptly widened into the fertile head,
distinct, puberulous, white: fertile head 0·4-0·8 mm. wide at the base, gradually tapered
TEXT-Ft a. 177. Honnonzitaria sulphurea: base of the head, showing the sterile rim connecting
the stem and hymenium (with oleocystidia), X 500.
HORMOMITARIA
downward to the subacute (never filiform) apex, entire, straight or subsinuous, smooth,
pale clear yellow or sulphur-yellow: flesh waxy-firm, solid: without smell.
On a rotten branch in the forest, on a limestone hill: Malaya (Kedah, Langkawi
Island, Bukit Batu Puteh near Kuah, 12.11.41).
Spores 5-6 X 3-3·5µ, white, smooth, pip-shaped, aguttate.
Basidia 14-18 X 4-5·5µ, narrowly clavate, with yellowish cytoplasm, projecting
3-4µ.: sterigmata 4, 4µ. long, straight, erect, or slightly spreading (sometimes only 2,
elongate and sterile, sterigmata -10µ. long).
Cystidia 30-50(-65) X9-12µ., clavate, like much-enlarged sterile basidia but
immersed, longitudinal in the subhymenium, curving into the hymenium or upright
in the hymenium but not projecting, abundant, thin-walled, capped with a dense oily
resinous yellowish excretion 3-6µ thick (enclosed in a hyaline membrane but often with
irregular processes -12µ. long), contents fairly dense with a few large vacuoles, the
variation in length caused mainly by the elongation of the narrow stalk-like base.
Hymenium not thickening: subhymenium 35-40µ. thick at the base of the club,
narrower towards the apex, composed of 2-3µ wide, closely interwoven, uninflated
hyphae, with rather toughly mucilaginous walls.
Caulocystidia 30-65 X 8-20µ, subcylindric to clavate or subventricose, blunt, not
appendaged, with firm, very slightly thickened, smooth walls, colourless, often somewhat
curved, occasionally branched, the longer ones sometimes I-septate (without a clamp),
occasionally with a small subglobose lobe, as a loose palisade over the whole stem,
crowded and compacted at the apex, looser, narrower, and often irregular at the base
of the stem, with transitions to 2-3µ wide, secondary mycelial hyphae.
Hyphae monomitic, inflating, with rather toughly mucilaginous walls, without air
spaces in the tissue, clamped, becoming secondarily septate in the stem and fertile
head.
Flesh of fertile head composed of longitudinal inflated hyphae, the cells 15-140 X
8-30µ, with a few narrow uninflated hyphae: many of the secondarily septate hyphae
forming curved rows of subglobose or broadly ellipsoid cells 15-30µ wide, 4-IO cells in
a row.
Flesh of the stem composed of longitudinal, secondarily septate hyphae with cells
20-100 x 5-14 µ, not forming moniliform rows, the surface hyphae 5-10 µ. wide.
Sterile base of the fertile head covered with a compact layer of caulocystidia,
gradually passing through a sterile hymenium of short subclavate hyphal ends
20-30 X 6-10µ., into the fertile hymenium, the sterile hymenium covering the base
for a length of 100-200µ. from the apex of the stem.
Growing apex of the fruit-body 50-100µ. wide, composed of very narrow hyphal
tips, 1 · 5-2 ·5µ wide, agglutinated with firmly mucilaginous walls and not projecting
as a transparent fascicle of hyphae, the hymenium developing 30-150µ. behind the
apex, and the apex becoming subacute, 90-150µ wide, on cessation of downgrowth:
subterminal cells 10-20µ. long on delimitation. Colour caused by the yellow cytoplasm
of the basidia and subhymenial hyphae.
Development of the Fruit-body. The fruit-body passes through a minute primordial
stage which gradually enlarges by vacuolation, by secondary septation of the hyphae
of the stem and of the flesh of the head, and by intercalary growth of the hymenium;
apical growth may continue until the fruit-body is 6 mm. long. The primordia are
superficial and discrete, never truly caespitose, though they may grow together in
pairs if too crowded. The earliest stage which I examined was a white conical body
130 X 100µ (at the base). It was composed of narrow hyphae, densely intertwined but
mainly longitudinal, with slight inflation of the cells near the base (4-5µ wide): round
the base some of the superficial hyphal ends had begun to project as the first, rudi
mentary, caulocystidia 8-15 X 4-5µ. This minute primordial shaft evidently grows
apically until it is 250-350µ. long, when a distinct, conical, and pale yellow head
appears, c. 70µ long X 100µ wide at the base, which is slightly wider than the apex of
HORMOMITARIA
the stem. The structure is extremely difficult to make out because the agglutinated,
firmly mucilaginous, hyphal walls prevent the tissue from being displayed by the usual
method of tapping the coverslip. It seems that the head is formed by the sudden
production of many short lateral processes to the exterior or by the sudden out
turning of the tips of the outer hyphae of the growing-point, which until this time
have remained concrescent and longitudinal. When the head appears, the stem is a
short cylindric shaft of hyphae c. 200-300 fL long, with scattered, rudimentary, caulo
cystidia. It is not corticated directly by hyphal ends divergent from the growing
point, but secondarily by the development of the caulocystidia as excrescent lobes
from the superficial hypha. This secondary cortication takes place from belowupwards
and then by intercalary lobing as the superficial hyphae become secondarily septate.
The stem elongates by vacuolation and secondary septation followed by further
vacuolation and enlargement of the cells: the process disrupts the palisade of caulo
cystidia and thus permits the continual intercalation of new ones. Thus the stem is
not mature and full-grown until the head is well formed, perhaps even 5-6 mm. long.
The head is narrowly conical from the first. The base widens by inflation of the
medullary hyphae and by the stretching which results from the great intercalary
growth of the hymenium. The widening proceeds acropetally, but the distal tissue
never becomes as inflated as the proximal, so that the fertile clubs are always tapered
for this reason, if not through slight attenuation of the apex, which, however, seems
equally narrow from its inception.
The hymenium develops by the out-turning of the hyphal tips on the outside of
the growing-point. Subapical branching of the hyphae occurs almost immediately
and so the hymenium is compact from the first and appears continuous almost to the
HORMOMITARIA
apex. Eventually, on cessation of growth, the hymenium develops over the apex,
which is never permanently sterile. The first hymenial elements in the transition
region between the stem and the head are sterile and they gradually pass from typical
colourless caulocystidia into the narrow yellow basidia: thus every fruit-body has this
characteristic, sterile, expanded base to the head. The fertile hymenial elements
develop directly into basidia, and the fruit-bodies, therefore, begin to spore when in
the primordial state, even when they are only 500µ, long.
The cystidia are formed only in the fertile head, but they appear as soon as the
hymenium in the primordia. They are produced either from the ends of longitudinal
hyphae .near the outside of the growing-point, in which case they are longitudinal and
deeply buri�d in the subhymenium, or they are produced from the original hymenial
hyphae and become gradually embedded as the hymenium expands through inter
calary growth, in which case they are more or less perpendicular to the medullary
hyphae and often slightly projecting in the primordia.
The characteristic short moniliform rows of inflated cells arise from the close
secondary septation of some of the inflating hyphae and from the almost spherical
inflation of the cells. The rows become curved as they enlarge more than the sur
rounding tissue.
LACHNOCLADIUM Lev.
Orbigny, Diet. Nat. Hist. 8, 1849, 487: Pat., Journ. de Bot. 3, 1889, 23, 33; Ess.
Taxon. 1900: Burt, Ann. Mo. Bot. Gdn. 6, 1919, 266: Donk, Bull. Bot. Gard. Buit.
Ser. 3, 17, 1941, 1 73: Singer, Mycologia 36, 1944, 553.
Synonyms: Eriocladus Lev., Ann. Sci. Nat. Bot. Ser. 3, 5, 1846, 158 (haud Erio
cladium Lindley).
Stelligera Heim ex Doty, Lloydia II, 1948, 134.
Fruit-bodies highly ramified, often caespitose, attached by yellowish mycelial fibrils,
often with a short sterile, matt or tomentose stem: branching flattened, in alternating
planes, dichotomous or apparently polychotomous from approximate dichotomies:
branches rather slender, terete or ± flattened, obcuneate when young, eventually
tapering, sterile and matt or subvillous on the upperside: hymenium waxy, unilateral on
the underside: more or less yellowish, ochraceous, tawny or subferruginous, often clearer
yellow or ochraceous near the white tips: flesh ± coriaceous, rather dry, pliant, drying
stiff and fairly rigid, not horny.
In humus or lignicolous, in tropical forests: c. 20 spp.
Spores white, smooth, very small, aguttate, often drying pale yellowish.
Basidia small, with (2-)4 subulate sterigmata.
Gloeocystidia always present, thin-walled, colourless, smooth, with oily homogeneous
or guttate contents, mostly immersed, traversing the hymenium in species with thicken
ing hymenia: setae and thick-walled cystidia absent.
Hyphae monomitic, septate, without clamps, not inflating, with thin and colourless or
slightly thickened, pale yellowish or brownish walls, a few with oleaginous contents
(as laticiferous hyphae).
Dichophyses abundant in the mycelium and fruit-body but absent from the thickening
hymenia, arising from a stalk but rarely with a distinct central body, repeatedly
dichotomous, aseptate, with thickened pale yellow or brownish yellow walls and colourless
acute tips (rarely subcircinate).
Stem and branches (and mycelial strands) with a medulla of longitudinal, compact
generative hyphae and few dichophyses, and with a paler looser, often somewhat
spongy cortex with interwoven generative hyphae and very numerous dichophyses.
Hymenium developing as a discontinuous layer of strongly projecting basidia,
. becoming continuous in species with thickening hymenium: without dichophyses:
the subhymenium becoming waxy and traversed by gloeocystidia.
LACHNOCLADIUM
Sterile surfaces covered with a mass of subcoralloid, very small dichophyses,
without gloeocystidia, often thickening slowly throughout the life of the fruit-body.
Colour caused mainly by the yellow or brownish yellow walls of the medullary
generative hyphae and of the dichophyses, darkening with dilute alkali.
In general form, structure, and colour (at least, of dried specimens) the species are
so uniform that I doubt if any can be distinguished macroscopically. Some have
distinctly flattened branches and others subcylindric, yet both extremes may occur
on the same fruit-body. Nevertheless, there may be a species with unusually stout
branches, appearing articulate at the nodes like a Coralline alga, but I have seen it
only in the Kew herbarium. Similarly, the spores, gloeocystidia, and dichophyses
are extremely alike. But important differences appear in the formation of the
hymenium, the relative thickness of the medulla and cortex, and in the size of the
medullary hyphae, e.g. L. brasiliense. These points have not been noted in the previous
descriptions of species which have been based on superficial differences, probably
untenable. It is, therefore, impossible to give a reliable or useful general key to the
species while such doubts exist. I give merely a tentative key to those which I have
studied personally from living specimens. My new species may be identified with the
old when the types have been examined microscopically or when they have been
rediscovered and identified from the type-localities, but, at present, our knowledge of
the genus can best be advanced by the description of living plants, under new names
if necessary, by 'field-mycologists'. Later, one may hope, the synonomy will be
disentangled. The species live mostly in the rain-forests of the tropics and, though
locally common, are seldom met with, and the field-mycologist who can study them
will seldom have access to the types in European herbaria. The herbarium-worker,
on the other hand, can form no idea of specific differences until he has much more
ample material coupled with the descriptions of the living specimens. It is noteworthy
that the specific differences in this peculiarly isolated tropical genus are anatomical
like those of algae.
KEY TO THE MALAYAN SPECIES

Hymenium thickening continuously
Medulla wide, 600-800 µ: flesh softly floccose: branches ±
flattened, divaricate: in humus L. molle
Medulla narrow, 100-350µ wide: flesh firm, tough . L. divaricatum
Irregularly bushy, fawn-ochraceous or cinnamon-ochraceous,
branchlets bright ochraceous: smell of bugs: in humus and
lignicolous : var. cznnamo-
meum
Densely fastigiate, pale fawn drab, branchlets pale yellowish:
smell sour: lignicolous var. sordidum
Hymenium not thickening continuously
Cortex ± zoned, each layer terminated by a single layer of
basidia on the fertile surface: fawn-ochraceous, subferruginous
downward: in humus L. zonatum
Cortex simple, the hymenium not layered
Densely and strictly branched: wholly dingy yellowish:
lignicolous . L. fiavidum
Irregularly bushy, rather spreading: fawn-ochraceous or
tawny, branchlets bright yellowish: humicolous L. fulvum
(Lachnocladia incertae sedis: p. 430.)
SI 19 Ee
418 LACHNOCLADIUM
L. brasiliense Lev.
Ann. Sci. Nat. Bot. Ser. 3, 5, 1846, 159 (as Eriocladus brasiliensis): Pat., Journ. de
Bot. 3, 1889, 26, pl. 1, f. 5: Lloyd, Myc. Notes 59, 1919, 827, f. 1384: Burt, Ann.
Mo. Bot. Gdn. 6, 1919, 268, pl. 5, f. 1, text-fig. 7: Singer, Mycol. 36, 1944, 553.
Synonyms: ? L. ambiginosum Cke. (teste Lloyd:? MS. name).
? L. samoense P. Henn., Engl. Jahrb. 23, 1896, 279 (teste Lloyd).
3-5 cm. high, very shortly stalked, very much branched: branches terete, dichoto
mous, ochroleucous, tips acute: on trunks: Brazil (Bahia). [Lev.]
3-12 cm. high, ochraceous (drying tawny-olive), rigid: stem 0-1·5 cm. X 2-4 mm.,
dividing into 2-several stout branches: branches 1-1 ·5 mm. wide, terete, solid,
divaricate, intertwined, repeatedly 2-3-fid with somewhat flattened axils, highly
branched, tips subulate: spores 3-4·5 X 2--2·5µ., white, smooth: dichophyses with rays
-30 X 3µ., pale brown, slender, abundant in the subhymenium, in the cortex, and on
the sterile sides of the branches: on rotten wood: pantropical. [Burt, Lloyd] : densely
caespitose, southern Florida (Singer).
I am doubtful if this species has been correctly recognized. The fragment of the
type at Kew shows what appear to be very wide (4-10 µ.) medullary hyphae, which
seem distinctive: in all the Malayan species which I have seen they are only 2-5µ. wide.
L. cladonioides P. Henn.
Engl. Jahrb. 25, 1898, 498.
2--2·5 cm. high, branched from the base, pale ochraceous, tomentose: branches
dichotomous, subterete, mostly only once divided: branchlets 1-2 mm. thick, obtuse
or cristate.
Spores 4-5µ., brown, smooth, subglobose.
On the ground in the forest: New Guinea (Neu Pommern).
Said to be near L. furcellatum. Probably dwarfed or young.
L. cristatum Lloyd.
Myc. Notes 61, 1919, 890, f. 1547.
3-4 cm. high, pale ochraceous: stems -1 cm. X 2 mm., or branched from the base:
branches not numerous, axils flattened, tips subcristate, pubescent to the eye: tough.
S. Africa (? terrestrial).
Spores 3-4µ. wide, white, subglobose (but irregular).
'Hairs under the microscope hyaline, sharp, smooth, projecting about 20µ..'
This seems to be an extra-tropical species.
L. divaricatum Pat.
Journ. de Bot. 1889, 6: Bres., Ann. Myc. 9, 19u, 272 (non Calocera divaricata B.,
Journ. Bot. 1842, 140: ? Pterula divaricata Lev., Ann. Sci. Nat. 3rd Ser. 1846, 158).
Synonyms:? L. asterosetosum P. Henn., Hedw. 1897, 196 (teste Bres.).
? L. cervino-album P. Henn., Engl. Jahrb. 28, 1901, 320.
? L. madeirense P. Henn., Hedw. 1904, 174 (teste Bres.).
Text-Figs. 33, 179.
? Stelligera membranacea Heim ex Doty, Lloydia II, 1948, 134: Heim, Com. Rend.
Ac. Sci. Paris 206, 1938, 1920.
I use this name, on the evidence of Bresadola, for a variable species of Africa and
Asia, distinguished by the thickening hymenium and the slender medulla. It may
well be a group of species, for there are considerable differences in manner of branch
ing, colour, smell, habitat, and details of microscopic structure, but collections and
field-notes are far too few to form an opinion. Beside the two Malayan varieties,
which I describe in detail, I have studied also a collection from W. Africa (C. 0. Far
quarson, S. Nigeria, 1914), referred to this species tentatively by Miss Wakefield at
Kew. It differs from the Malayan varieties in the strongly palmate branching and the
hollow medulla. The lower branches are slightly flattened but strongly dilated at the
LACHNOCLADIUM
divisions into 2-6 smaller branches (actually successive close dichotomies): the upper
branchlets are terete and dichotomous. The medulla, in the dried specimen, is hollow
in the larger branches and is probably so in the living state because, in the stem, it
becomes stuffed with a yellowish ingrowth of loose generative hyphae with short
dichophyses (25-100µ. wide), as the mycelial strands. Other notes on this collection
are:
Cortex 1-2 mm. wide on the trunk: medulla 100-250µ. wide, even in the stem:
hymenium thickening to 100µ., unilateral, traversed by
gloeocystidia: spores 3·5-4 X 2·5-3µ.: dichophyses 4-6µ. wide
at the central body.
The descriptions of Hennings's species, the identity of
which needs verification, are:
L. asterosetosum: pale alutaceous (dried), tomentose: stem
1-2 cm. X 1-1·5 mm., terete, stuffed, repeatedly branched:
branches subterete, axils compressed, divergent, tips furcate
or subulate, covered with asteroid setae: spores 3-4µ.,
globose, smooth: on the ground: Brazil (Blumenau).
L. cervino-album: 4-7 cm. high, coriaceous, densely setulose,
fawn-colour (dried): stem -4 cm. x 2-5 mm., subterete or
compressed, ± elongate, subflexuous, cervino-velutinate,
repeatedly branched: branches palmatifid or dichotomous,
with broadly compressed axils, setulose: branchlets subulate
or cristate, white: spores 3�4 X 3-3·5µ., yellowish, ellipsoid or
subglobose: dichophyses with arms -80 X 3·5-4·5µ.: on rotten
leaves in the forest: E. Africa (Uhehe).
L. madeirense: -9 cm. high, alutaceous cinereous (dried):
stems -3 cm. x 2 mm., free or connected: branches repeatedly
dichotomous, axils scarcely compressed, cinerous setulose,
with long subulate tips: spores 2-3µ., subglobose: on the
ground and on wood: Amazon.
It is possible that the African collections of L. furcellatum,
on which Fries based Clavaria furcellata, are this species.
L. divaricatum var. cinnamomeum var. nov.

TExT-FIG. 179.
Text-Figs. 33, 34, 36, 180-3. Medullary dichophysis
-10 x 8 cm., densely gregarious, compact, very bushy, pale of Lachnocladium divari-
fawn-ochraceous to pale cinnamon-ochraceous, dingy or fuscous catum, X 500.
below, bright ochraceous near the extreme white tips; stem
0-5 cm. x 2-4 mm., often indistinct, dividing into a few stout branches, attached by
pallid white mycelial strands: branches c. 1 mm. wide, more or less flattened -2 mm.
wide at the divisions, terete, varying dichotomous or alternate to multifid and
palmate, sterile or matt on the upperside: hymenium on the underside of the branches,
continuous, waxy, rarely extending on to the stem, pale brownish cream or ochraceous
drab: flesh concolorous, tough, pliant, smell peculiar, of bugs.
In deep humus and on rotting wood in the forest: Malaya (Perlis 25.11.29, in
humus of fallen tufts of Asplenium nidus at Bukit Lagi, near Kangar: Pahang, J erantut,
10.6.31, S.F., No. 24865, on dead wood: Pahang, Tembeling, 22.11.30, in humus:
Selangor, Ridley 19, Batu Caves, Dec. 1896, det. L. furcellatum).
Spores 3-3·5 x 2·5µ., white, smooth, ellipsoid, with a minute apiculus.
Basidia 15-24 x 3-4µ., projecting -10µ.: sterigmata (2-3-)4, 3-3·5µ. long, needle-like.
Gloeocystidia 25-200 X 6-18µ., wholly immersed or projecting -15 µ., at first sub-
ventricose or subfusiform, then elongating and traversing the hymenium with a
subcylindric, subflexuous, blunt neck, rarely bifurcate, filled with vitreous-oleaginous
matter, thin-walled, colourless, absent from the sterile hymenium.
420 LACHNOCLADIUM
Dichophyses in the cortex with the stalk 20-50µ, long, the central body scarcely
enlarged, 4-5 µ, wide, the 3-8 branches -150µ, long, one to five times dichotomous,
3-4µ, wide at the base and tapering to acute, divergent tips 0·5-1 µ, wide, the walls
becoming 0·5-1 µ, thick and pale brownish ochre to dark ferruginous, often with slight
irregular processes: in the subhymenium and first-formed hymenium 20-35 µ, wide,
short, dense, and subcoralloid: in the medulla, few, with elongate, lax, more or less
longitudinal, rarely transverse and short, branches.
TEXT-FIG. 180.Lachnocladium divaricatum var. cinnamomeum: fruit-bodies, x 1:

A, immature and, as yet, sterile: branch-tips, X 5.
Hymenium 15-20µ, thick at first, gradually thickening up to 200µ,, very compact
but the thickened part without dichophyses.
Medulla 0·4-1 mm. wide at the base of the stem, 100-350µ, wide in the greater
part of the stem and main branches, tapering to 20-50µ, wide in the branchlets,
relatively slender.
Cortex 1-2 mm. thick at the base of the stem, 200-400µ, thick on the main branches,
tapering to 20-30 I'- thick on the branchlets, fairly compact, relatively thick.
Generative hyphae 2-5 I'- wide, long-celled, not inflated, without clamps, the walls
thin or thickened and pale yellowish brownish: 1 · 5-2·5 µ, wide in the subhymenium:
at the tips of the branches 1·5-2·5 I'- wide, often reduced to a single leading hypha on
the fine, ultimate tips.
p;;;;),��":-.�
� .. ·�c
=.=-.r.,_\ :,'_r'("(;
--),-?,'fr-
,r.i,O,::,..
�(.-.;r\.�
TEXT-FIG. 181. Lachnocladium divaricatum var. cinnamomeum: a fruit TEXT-FIG. 182. Lachnocladium divaricatum var. cinnamomeum,
body growing among the debris of a fallen tuft of Asplenium nidus, X 1 : young fruit-bodies, X 2.
sections, X 2.
422 LACHNOCLADIUM
Both strikingly palmate and scarcely palmate branching may occur even on the
same fruit-body in this variety.
TExT-FIG. 183. Lachnocladium divaricatum var. cinnamomeum: young fruit-bodies

on debris of Asplenium nidus, X 2.
L. divaricatum var. sordidum var. nov.

Text-Fig. 184.
-15 cm. high, densely branched, upright, pale fawn-drab, light yellow towards the
white tips: stem 5-28 X 3-5 mm., short, matt, thick or, if growing in very rotten wood,
thin and rooting, sterile, with yellowish white mycelial strands 0·5 mm. wide: branches
1·5-2·5 mm. thick below, soon narrowed to 1 mm. and tapering to fine acute tips,
terete, slightly flattened 1·5-2 mm. wide at the divisions or not at all, spuriously
polychotomous below, dichotomous above, with rather elongate internodes, erect,
strict, tense, scarcely spreading, sterile and matt on the uppersides: hymenium waxy,
light drab: flesh slightly tough, pliant, drying coriaceous: smell rather strong, sour.
On dead fallen trunks in the forest: Malaya (Pahang, Padang Pio!, 4.12.30).
Spores 3-3·5 X 2·5-3 µ,, as in var. cinnamomea.
Basidia, gloecocystidia, dichophyses, and hyphae as in var. cinnamomea.
Medulla 80-200µ, wide, tapering to 80-120µ, in the branchlets, narrow.
This is very near var. cinnamomeum, but the colour is much paler, the branching
more strict, and the smell different, though perhaps derived rather from the sub
stratum as with many other sour-smelling lignicolous fungi. See also L. fiavidum,
which may be only a state with unthickened hymenium.
L. flavidum sp. nov.

Text-Fig. 185.
-7 cm. high, densely branched, upright, wholly dingy yellowish, tinged fawn below,
the tips pale yellowish white: stem 5-20 X 1·5-3·5 mm., terete, with very thick cortex
and narrow medulla, sometimes branched from the base, with fine yellowish white,
mycelial strands: branches 1-2 mm. wide below, terete, tapering gradually to the acute
LACHNOCLADIUM 423
tips 0·2-0·5 mm. wide, spuriously polychotomous below and flattened 1·5-2·5 mm.
wide at the divisions, curved ascending, crowded, minutely villous on the sterile
uppersides: hymenium waxy, thin: flesh subcoriaceous, sappy, pliant, drying very
brittle and rather spongy: smell ?
On rotten wood in the forest: Malaya (Pahang, Tembeling, 6.11.30).
Spores 3-3·5 X 2·5 µ, as in L. divaricatum var. cinnamomeum.
Gloeocystidia 30-50 X7-12µ, projecting slightly or not at all.
8
C:)
TEXT-FIG. 184. Lachnocladium divaricatum var. sordidum, X r: sections,

X2.
Basidia, dichophyses, and hyphae as in L. divaricatum var. cinnamomeum.

Medulla 100-250µ thick, tapering in the branchlets, narrow.
Hymenium 25-30µ thick, not thickening.
This species is perhaps only a state of L. divaricatum var. sordidum with the
hymenium not thickening, but it may differ also in texture, having looser dichophyses
without such long interweaving tips, and in lacking the strong sour smell.
424 LACHNOCLADIUM
L. fulvum sp. nov.
Text-Fig. 186: Plate 1 I.
-9 cm. high, densely branched, erect, fastigiate, light fawn-ochraceous or tawny,
brighter yellowish towards the white tips: stem indistinct, 1-2·5 mm. thick, with rooting
TExT-FIG. 185. Lachnocladium fiavidum, X 1 : sections, X 2: growing on rotten wood.
yellowish mycelial strands: branches 0·5-1 mm. wide, terete but flattened slightly,
1-1 ·5 mm. wide, at the divisions, tapering to the acute tips: hymenium pale fawn-drab,
waxy, on the underside of the branches: flesh, tough, dry,
light fawn-ochraceous with darker medulla: smell strong, sour,
and musty.
In deep humus and on sticks: Malaya (Singapore, Botanic
Gardens, 22.3.45).
Spores 3·5-,i-3 X3-3·5 µ, white, smooth, pip-shaped with
a prominent blunt apiculus 1 · 5 µ long, with opalescent, sub
vitreous contents.
Basidia 25-35 X4·5-5·3µ, subclavate, projecting -12µ, as
a discontinuous layer: sterigmata 4, 5 µ long, straight, parallel.
Gloeocystidia -100 X6-10µ, cylindric or subventricose,
blunt, thin-walled, with large guttae or vitreous contents,
not or scarcely projecting.
Hymenium, with subhymenium, 60-70µ thick, not
thickening.
TExT-FIG. 186. Lachno- Medulla 150-350µ thick, relatively narrow.
cladiumfulvum, X i, ooo. Dichophyses and hyphae as in L. divaricatum.
This species is near L. divaricatum var. cinnamomea and L. zonatum, but differs in
LACHNOCLADIUM
the larger, strongly apiculate spores with larger basidia. From the first it differs also
in the hymenium not thickening, and from the second in the longer gloeocystidia, the
simple hymenium, and, perhaps, the paler colour and larger size of the fruit-body.
A collection from New Guinea (C. E. Carr u692, Veiya, Papua, in humus and on
rotten twigs in the forest, 12.3.35) appears to be the same species, but I could find no
spores in the dried specimens (see Plate 11, f. 4). The medulla is very slender,
50-130µ thick, even in the short stem (1-2 cm. X 1·5-2 mm.).
L. furcellatum (Fr.) Lev.

Ann. Sci. Nat. Bot. Ser. 3, 5, 1846, 159 (ut Eriocladusfurcellatus): Berkeley, Journ.
Linn. Soc. Bot. 10, 1868, 330: ibid. 14, 1875, 67: Pat., Journ. de Bot. 3, 1889, 26,
pl. 1, f. 3: Burt, Ann. Mo. Bot. Gdns. 6, 1919, 274, pl. 5, f. 7: Beeli, Bull. Soc. Roy.
Bot. Belg. 58, 1926, fasc. 2, 7.
Basinym: Clavariafurcellata Fr., Linnea 5, 1830, 531.
Synonym: Dendrocladiumfurcellatum (Fr.) Lloyd, Myc. Writ. 6, 1920, 958.
As mentioned on page 84, it is as yet impossible to identify this species. It was based
on, at least, three collections (Guinea, Bourbon, Brazil, 'ad ligna putrida in tropicis'),
but it has to be proved that they have the same microscopical construction and are
conspecific. The species has since been discovered in most tropical countries but,
again, more through superficial resemblance than precise anatomy. Certainly in the
Kew herbarium, several species have been placed under this name and the American
specimens appear different from the Asiatic which, again, do not seem homogeneous:
a few clearly do not belong to Lachnocladium s. str. I have not worked through these
collections, partly because they are not authentic, but chiefly because I believe that
the living plants must be described before a clear idea of the specific variation and
limits in the genus can be obtained.
Macroscopically the species is like L. divaricatum which may, in fact, be a synonym.
It is generally described as:
-12 cm. high, pallid ferruginous to ochraceous ferruginous with a vague stem
tapering downward, divided into several large, flattened, multifid main branches, and
dichotomous, terete, elongate branchlets with acute, filiform tips: rather tough, solid,
velvety: on rotting wood.
Burt gives the spores as 7-12 X 4 · 5-6µ, pale ochraceous; Beeli gives them as 8 X 3-4µ,
pale ochraceous: they must refer to specimens of Ramaria misidentified as Lachno
cladium. Yet Berkeley wrote 'the tips of the threads composing the hymenium are
widely forked': he was the first to describe the character of Lachnocladium.
L. galaxauroides P. Henn.
Pilze Ost. Afr. 1898, 55: Sacc., Sy!!. 14, 1899, 240.
2-5 cm. high, caespitose, erect, alutaceous (dried), rather spongy, somewhat stuffed,
setulose-tomentose: mycelium membranous, whitish: stem simple or branched from
the base: branches di- or trichotomous, or irregularly partite, terete, thick, rarely
compressed, subrugose, with subulate-furcate, obtuse or canaliculate tips: subfragile
(dried).
On burnt ground: tropical Africa (leg. Holst.).
Setulae branched.
Possibly a young stage of L. divaricatum.
L. Hisingeri (Karst.) comb. nov.

Basinym: Polyozus Hisingeri Karst., Finl. Nat. o. Folk 48, 1889, 395: Acta Soc.
Sci. Fenn. 18, ·1891, 106, t. 70.
Synonym: Thelephora Hisingeri (Karst.) Sacc., Syll. 9, 1891, 220. 2-4 cm. high,
erect, slender, sparingly and laxly branched, pale yellowish white, drying slightly
LACHNOCLADIUM
fuscous or ferruginous: stem slender, three to four times dichotomous: branches
flattened, slender, with paler, acute tips: coriaceous-soft.
On roots of Cyathea medullaris, in a warm orchid-house (Finland),
Spores 2 µ wide and globose, or 3 X 2 µ, white, smooth.
Karsten's figure shows the colour as pale brownish ochraceous. I think the species
is clearly a depauperate Lachnocladium of doubtful origin.
L. molle sp. nov.

Text-Figs. 3, 187.
--'7·5 cm. high, profusely branched, spreading, pallid dingy ochraceous, tbe growing
tips white: stem 15--22 X 2-3 r.un., closely and minutely villous with dirty ochraceous
'I'ExT-FIG. 187. Lachnocladium molle, X 2, showing the wide medulla (in black).
down, attached by short yellowish mycelial strands o·5-1 mm. wide: branches
1-1·5 mm. wide, more or less flattened particularly at the divisions, dichotomous or
apparently polychotomous at first, divaricate, laxly ascending, the uppersides sterile
and minutely villous, tapering gradually to the acute tips (obtuse in young fruit
bodies with active division), branchlets 0·5 mm. wide: hyrneniurn waxy, palefawn-drab,
LACHNOCLADIUM
on the underside of the branches, often discontinuous: flesh slightly tough, but rather
soft, pliant, dry.
In humus in the forest: Malaya (Pahang, Tembeling, 20. 11.30, Sing. Field, No.
24193).
Spores 3-3·5 X 2·5 µ,, as in L. divaricatum var. cinnamomeum.
Basidia, gloeocystidia, dichophyses, and hyphae as in L. divaricatum var. cznna
momeum.
Cortex 100-200 µ, thick on the sterile parts, 50-80µ, thick on the fertile sides of the
branches (25-35 µ, on the smaller branches), relatively narrow.
Medulla 600-800µ, wide, narrowing to 200µ, in the ultimate branchlets, relatively
wide.
Dichophyses stiffly rayed in the cortex, rather abundant in the medulla.
This species is distinguished by its flattened, lax and spreading, though abundant,
branches and by the very wide medulla which causes the comparatively soft flesh.
Microscopically there is no other difference from L. divaricatum var. cinnamomea.
The medulla at the base of the trunk of this variety may be as wide as in L. molle, but
it very soon tapers to a relatively minute strand through the branches.
L. moniliforme P. Henn.
Hedw. 1904.
Synonyms: Thelephora clavarioides Torr., Brot. Ser. Bot. 12, 1914, 61 (teste Bres.).
Hypolyssus clavarioides (Torr.) Lloyd, Lett. 54, 1915, n. 206.
Stem -3 cm. long, alutaceous pruinose, subligneous, sclerotioid, articulate-monili
form, the joints 4-10 X 3-4 mm. and subovoid or oblong subterete, fragile, constricted:
branches 2-2 · 5 cm. long, ferruginous fuscidulous, fasciculate, subcoriaceous, repeatedly
branched, mostly dichotomous, divergent, compressed, tips subulate.
On the ground: Brazil (Sao Paulo).
Spores 3-4µ,, hyaline-fuscidulous, ovoid, smooth.
L. ramalinoides P. Henn.
Mons. 1, 1900, 143.
-4 cm. high, fasciate-compressed, brown (dried), setulose: stem short: branches
short, repeatedly palmately branched, with flabellate axes: branchlets with compressed,
cristate apices.
On wood: Java (Tjibodas).
Spores 3 ·5-4 X 3 ·5 µ,, white, globose or ellipsoid.
This is said to be near L. cornicularioides, to which Hennings, however, ascribed
Ramaria-spores: those of L. ramalinoides indicate Lachnocladium s. str. In form it is
said to resemble Ramalina farinacea.
L. Sarasinii P. Henn.
Mons. 1, 1900, 7, t. 1, f. 1.
3-4 cm. high, caespitose, ochraceous, densely tomentose: stems confluent, sub
terete, tuberculate, irregularly repeatedly branched, mostly dichotomous: branches
subterete, axils scarcely compressed, straight, tips subulate, erect: coriaceous, fragile
(dried).
On twigs: northern Celebes (Buol., leg. Sarasin).
Spores 3-3 ·5 µ,, white, smooth, globose.
Dichophyses forming a thick felt over the fruit-body, with forked arms -70 X 2-3 µ,.
L. setulosum (B.) Lev.
Basinym: Clavaria setulosa B., Hook. Journ. Bot. 1845, 60.
Synonym: Eriocladus setulosus (B.) Lev., Ann. Sci. Nat. Bot. Ser. 3, 5, 1846, 303.
2·5 cm. high, ochraceous: stem short, irregularly divided: branches compressed,
'� .._.:; __
TEXT-FIG • I g8 . Lacrmocladium
1 zonaturn, X z.
LACHNOCLADIUM 429
forked, obtuse or flabellate, subtomentose and, in places, shortly spiculate (under a
lens).
On the ground: W. Australia (Swan River).
L. subarticulatum P. Henn.
Mons. 1, 1900, 142.
4-6 cm. high, blackish brown (dried), 'albescente setulosum', upright: stem
1 cm. X 2 mm., terete, fasciculate ramose: branches strict, erect, dichotomous, axils
compressed, attenuate, sulcate, tips subulate: coriaceous.
? Habitat.
Spores 3·5-4 X 3-3·5µ, white, smooth, ellipsoid to subovoid (1-guttulate).
Said to be near L. articulatum, but with quite different spores.
TEXT-FIG. 189. Lachnocladium zonatum: branch-tips with flattened branching, X 5.
L. subochraceum Sacc.
Ann. Myc. 4, 1906, 73.
3 cm. high, tawny ochraceous (dried), slightly and finely velutinate: stem very
short, 2-3 mm. thick, dividing into numerous branches: branches many times dicho
tomous, filiform, divergent-ascending, axils flattened and dilated: waxy coriaceous,
but fragile.
? On rotten fallen branches: Congo (Kisantu).
Spores?
Dichophyses with dichotomous branches, 2·5-4µ wide, and acute divergent tips.
Perhaps a dwarf form of L. divaricatum.
L. subpteruloides P. Henn.
Engl. Jahrb. 25, 1898, 498.
(haud L. subpteruloides P. Henn. 1900 = L. pteruliforme.)
7-9 cm. high, densely caespitose, very much branched, ochraceous, tomentose:
stem terete, 2 mm. thick: branches terete, thin, slender, repeatedly dichotomous or
trichotomous, 1 mm. wide, axils hardly compressed, tips subulate, squarrose.
On the ground in the forest: New Guinea (Kabakaul, Neu Pommem).
Spores 3-3 · s µ, globose, smooth, brown.
Dichophyses with arms 3-3·5µ wide.
L. Ulei P. Henn.
Hedw. 1892, 98.
Densely caespitose, ochraceous, ferruginous pruinose: stem 2 cm. X 2 mm., terete,
subflexuous: branches repeatedly dichotomous, axils arcuate and broadly compressed,
tips subulate, -1 cm. long: rather tough.
On dead trunks in the forest: Bra:z.il.
430 LACHNOCLADIUM
Spores 5-6 X4-5µ,, ellipsoid or ovoid, fuscidulous.
The spores seem rather large for Lachnocladium s. str. It may be Ramaria. Hen
nings likened it with Ramaria Moelleriana.
L. zonatum sp. nov.
Text-Figs. 31, 32, 35, 188-92: Plate II.
-5·5 cm. high and wide, much branched, with a distinct stem and laxly spreading
branches, solitary or gregarious, often several springing from a
small pulvinate, tomentose pad or subiculum,fawn-ochraceous,
subferruginous downward, tips white: stem 1-2 cm. X1·5-2·5
mm., finely subtomentose, sterile or fertile upwards: branches
I mm. wide below, tapered to the acute, often almost filiform
tips, more or less flattened below, often with 3-4 branches
about the same level and somewhat antler-like, straight or
outcurving, very finely subtomentose, often concrescent:
hymenium slightly waxy, vaguely delimited, absent from the
uppersides of the branches, concolorous or somewhat olivaceous
ochraceous: flesh paler concolorous, fibrous, sub-coriaceous,
dry.
On the ground in the forest, sometimes attached to sticks
or woody fruits': Malaya (Singapore, Bukit Timah Forest,
10.12.40, 16-4-41).
Spores 3-3·5 X2·5-3·3µ,, white, subglobose, smooth, agut
tate.
Basidia 15-23 X3-4µ,, subclavate, small: sterigmata (2-3-)4,
2-2·5 µ, long.
Gloeocstydia 25-55 X6-15 µ,, subventricose or subfusiform,
blunt, smooth, thin-walled, colourless, immersed or projecting
-15 µ,, with several large colourless oil-globules, hyaline
vacuolate, abundant.
Dichophyses with a central body 3-5 µ, wide and repeatedly
dichotomous rays up to 90 µ, long, tapering to acute or sub
acute tips 0·5-1 µ, wide, with smooth, slightly thickened (0·5-
1µ,), pale ferruginous walls, the tips generally colourless: rays
up to seven times dichotomous, lax and elongate in the
medulla, short and dense with more or less coralloid tips in
the hymenium and often small (even only 30µ, wide, overall):
stalk 10-50 X3-4µ,, slightly thick-walled.
Hyphae 2-4µ, wide, without clamps, with thin, colourless or
very slightly thickened, pale brownish walls, smooth, the cells
20-200µ, long: a few hyphae unbranched, aseptate, laticiferous,
3-4 µ, wide, with resinous-oleaginous contents.
Hymenium and subhymenium 30-40µ, thick, vague,
'TExT-FIG. 190. scarcely continuous, not thickening but developing as more or
Lachnocladium zonatum: less complete, successive zones over the thickening cortex.
diagram of the apex of a Medulla 100-300µ, wide: cortex gradually thickening
branch in l.s. and t.s. (to
the same scale as Text-
throughout the life of the fruit-body.
Figs. 191 and 192).
LACHNOCLADIA JNCERTAE SEDIS
Trop. Amer.
Greyish violet: French Guiana L. violaceum
White, yellowish, or brownish
French Gwana L. clavarioideum
TEXT-FIG. 191. Lachnocladium zonatum: diagram of the structure of a
'
mature upright branch with three cortical layers round the medulla, each
bounded by a hymenium, and with a fourth layer started by excrescent
dichophyses: (to the same scale as Text-Figs. 190 and 192).
TExT-FIG. 192. Lachnocladium zonatum: diagram of a cross-section of an oblique
branch flattened at the region of two successive dichotomies (as shown by the form
of the medulla): the upperside sterile, the lowerside with two cortical layers each
bounded by a hymenium, with a third layer started by excrescent dichophyses:
(to the same scale as Text-Figs. 190 and 191).
LACHNOCLADIUM 433
Brazil . L. manaosense
Brazil: with skeletal hyphae r ·5-3 µ wide L. reticulatum
Trop. Africa
Greyish violet: Cameroons
Sp. 7-8 X 3·5-4µ, fuscidulous L. victoriense
Sp. 4-5 µ, subglobose, white L. Dusenii
Sp. 6-7 X 5-6µ L. quangense
Sp. 5-7·5 X3·5-5µ L. palmatifidum
L. pteruloides
Trop. Asia and Australasia
Sp. 7-8 X 4-5 µ: India L. mussooriense
Sp. 6-'7 X 5-6 µ: lignicolous: Java L. palmatum
Sp. 5-7 X 4-5 µ, rough: Java L. zandbaiense
Sp. 3 ·5-5 µ, subglobose
Java . L. pteruliforme
New Guinea L. ralumense
Sp.?
India: (? a conidial Xylaria) L. himalayense
Java . L. implexum
Greyish yellow, orange, or red, small, -10 mm. high, simple or
sparingly forked: on bark, often erumpent L. aurantiacum
SPECIES lNCERTAE SEDIS

L. aurantiacum (B. et Br.) Petch
Ann. R. B. Gard. Per. 9, 1924, 177.
Basinym: Isaria aurantiaca B. et Br., Journ. Linn. Soc. 1875: Fungi of Ceylon,
No. 860.
Synonym: ? Clavaria Braunii P. Henn., Engl. Jahrb. 17, 1893, 22.
? Clavaria typhoidea P. Henn., Mons. 1, 1900, 141.
? Clavaria rhizomorpha B., Fl. Tasm. 2, 1860, t. 183, f. 4: Cke., Handb. Austr.
Fungi 1892, 203.
Fertile fruit-bodies -10 X o·6 mm., greyish yellow, forked about half their height, the
2 branches forked again into comparatively long, simple, terete, acute tips, with all
transitions to the sterile fruit-bodies, the fertile being rare.
Sterile fruit-bodies -2 X 0·4 mm., orange-yellow, simple or rarely forked, clavate or
fusoid, subacute, more or less tomentose: crowded in small clusters, more or less
radiating from a centre, or in long lines in cracks in the bark, arising from a yellow
mycelium in the bark: always, apparently, sterile.
On the bark of Apodytes Gardneriana (Icacinaceae): Ceylon (Hakgala): (type,
Thwaites 344, at Peradeniya).
Spores 7-11 x 3-5 µ, white (pale yellow under the microscope), smooth, oblong
oval, inaequilateral.
Hyphae 2-4µ wide, incrusted with yellow granules, loosely arranged.
This is Petch's description. The fungus is certainly no Lachnocladium. It seems
to have uninflated hyphae, which with their yellow incrustation and the shape of the
fruit-body suggest a Hydnoid fungus. Unfortunately it is not stated what direction
the clubs or spines take or whether their tips remain sterile.
5119 F f
434 LACHNOCLADIUM
Clavaria Braunii, C. typhoidea and, perhaps, C. rhizomorpha seem the same fungus
as Petch's. These are their descriptions:
C. Braunii: -2·5 X 1-1·5 mm., gregarious, simple, very small, fleshy, liguliform or
spathulate, broadly compressed, sometimes keeled dorsally, obtuse, smooth, orange,
stem short: spores 7-S x 4-5 µ,, hyaline, ellipsoid: on wood: Madagasc_ar (Tocamasino).
C. typhoidea: 4-6 mm. high, very small, simple, scattered, corticicolous, subulate,
intense reddish yellow to red: stem 0·3 mm. wide, slender, cylindric: head 0·5 mm. wide,
compressed, smooth, attenuate to the rather acute apex: spores 4-6 X 4-5 µ,, hyaline
fuscidulous, ovoid: on mossy bark of trees: Java (Gedeh, 2,600 m.).
C. rhizomorpha: 6-12 mm. high, erumpent, confluent, nearly simple, chestnut-red
or with a violet tinge: apices yellowish: on dead bark: Tasmania.
Compare also Pistillaria spathulata and Clavaria? dubiosa (p. 266).
L. clavarioideum Pat.
Journ. de Bot. 3, 1889, 27.
Stem 2-4 cm. X 3-4 mm., compressed, thickened at the base, striate, whitish
rufescent: branches 0·5-1 mm. wide, fastigiate, compressed, canaliculate, 2-3-choto
mous, with long sterile tips, fulvous: hymenium whitish rufescent, unilateral, glabrous.
Spores 4-5 x 3 µ,, scarcely yellowish (dried), ovoid, smooth, 1-guttate.
On the ground: French Guiana.
Perhaps Lachnocladium s. str. Compare L. manaosense.
L. Dusenii P. Henn.
Engl. Jahrb. 22, 1895, 86.
Stalked, much branched, coriaceous, soft: stem -5 X 2 mm., bulbous at base, com
pressed, repeatedly branched, subpalmate: branches broadly compressed, sub
tomentose, greyish violet: apices dichotomous, compressed, subspathulate, obtuse, or
subulate, pallid, pruinose.
Spores 4-5 µ, wide, white, smooth, subglobose.
On sandy ground: Cameroons.
This is said to recall Thelephora palmata in shape and colour. It may be a Clavulina.
L. himalayense Mass.
Kew. Bull. 1898, 114.
Synonym: Pterula himalayensis (Mass.) Lloyd, Myc. Notes 60, 1919, 867.
Stem 8-10 cm. long, rather thick, elongate, pale rufescent: branches and branchlets
rugulose, terete or compressed, axils arcuate, alutaceous then cinnamon fuliginous,
apices pallid white, blackening with age: spores 4-5 µ, wide, subglobose, white,
smooth: on the ground in fir-forests: Himalayas (Sikkim, 3,000 m. alt., Gamble 99).
There is nothing in the description to suggest that this is a Pterula, and Pterula
Mannii, with which Lloyd compares it, is a dubious species suggesting a Xylaria.
Perhaps this is also a conidial Xylaria, cf. Xylaria nigripes.
L. implexum (Lev.) P. Henn.

Mons. 1, 1900, 143.
Basinym: Clavaria implexa Lev., Ann. Sci. Nat. Ser. 3, 5, 1846, 154: [haud sensu
v. Ov. 1923 = Scytinopogon echinosporus].
Synonym: Merisma implexa Lev. (? ined.).
4-8 cm. high, branches 2 mm. wide at the base, much branched from the base,
branches and lower branchlets compressed, dichotomous or cylindric above, here and
there tubercular cinereous and cohering together.
On the ground: Java.
This is the original description, beyond which nothing is known. Van Overeem
identified with it a Javanese fungus which is Scytinopogon echinosporus, but whether
LACHNOCLADIUM 435
the original L. implexum is the same is impossible to decide without examination of
the type. It may even be a Clavulina with flattened branching.
L. manaosense P. Henn.
Hedw. 44, 1905, 59.
-6 cm. high, brownish cinereous (? dried), branched, stalked, caespitose, coriaceous:
stems subcylindric, variable in length, somewhat bulbous at the base, connate:
branches 0·5-0·8 mm. thick, palmatifid or dichotomous, repeatedly subverticillate,
subcylindric, axils not compressed, subulate, sometimes cirrate, acute, smooth.
Spores 5-6 X 4·5-5 µ,, white, smooth, ovoid or subellipsoid.
On damp earth in woods: Brazil (Rio Negro, Amazon).
This fungus seems related to Aphelaria defiectens, except in the colour, which may
be only that of a dried or preserved specimen. Compare L. clavarioideum.
L. mussooriense P. Henn.
Hedw. 1901, 324.
6-8 cm. high, subrufous (dried), tuberose, fasciculate-ramos: branches fasciate
compressed, sulcate, flexuous (dried), repeatedly dichotomous or trichotomous,
branchlets compressed and squarrose, not flabellate, axils hardly compressed, tips
subulate, bifid or denticulate.
On the ground: India (Mussoorie, Arnigadh).
Spores 7-8 X 4-5 µ,, white, smooth, ovoid.
? Aphe!aria dendroides.
L. palmatifidum P. Henn.
Engl. Jahrb. 23, 1897, 544.
-6 cm. high, white, erect, coriaceous, tough: stems branched, or united at the base,
subterete or compressed, tomentose: branches with broadly flattened, flabellate axils,
palmately branched: branchlets repeatedly dichotomous or palmatifid, with squarrose,
subulate-elongate tips 0·5-1·5 cm. long: spores 6-7·5 x 3·5-4·5 µ,, slightly yellowish
under the miscroscope (dried specimen), smooth, ellipsoid: among fallen leaves,
Cameroons (Bipindi).
? Scytinopogon or Ramaria Molleriana.
L. palmatum P. Henn.
Mons. 1, 1900, 142.
-4 cm. high, coriaceous, caespitose, stalked, pale yellow: branches subterete, pal
matifid, or fasciculate at the apex, broadly compressed: bases of the branches confluent,
apices subulate or cristate.
Spores (>-;; µ, wide, yellowish white, smooth, globose.
Basidia with 4 sterigmata 6-8 µ, long.
On stumps: Java (Tjibodas, c. 1,500 m. alt.).
This is said to be near Lachnocladium galaxaurioides. It may be a true Lachno
cladium, but the spores are larger and the sterigmata longer than in that genus.
L. quangense may be the same.
L. pteruliforme P. Henn.
Mons. 1, 1900, 173.
Synonym: Lachnocladium subpteruloides P. Henn., Mons. 1, 1900, 143 (non L.
subpteruloides P. Henn. 1898).
-5 cm. high, horny cartilaginous, tough, cinereous white, becoming brown: stem
thickened at the base and clothed with pallid membranous mycelium: branches
subligneous, compressed, sulcate, repeatedly fasciately branched, 'setulose', axils
436 LACHNOCLADIUM
compressed, subflabellate: branchlets erect, subulate or cristate, reddish brown,
glabrous, naked, often dilated.
Spores 3·5-4µ, white, subglobose, smooth.
On the ground: Java (Buitenzorg Botanic Garden).
If it were not for the small spores, I would identify this with Aphelaria dendroides.
The description refers to dried specimens. The broadly flattened branching is not
like Pterula. L. ralumense seems identical.
L. pteruloides P. Henn.
Pilze Ost-Afr. 1898, 56: Sacc., Syll. Fung. 1899, 240.
2-4 cm. high, caespitose, pale ochraceous, confluent, subpulvinate, erect, setose:
stems very much branched, fastigi'ite: branches repeatedly dichotomous or fasciculate,
axils widely compressed, strict or divergent, rather tough, tips subulate, fastigiate or
cristate: mycelium white, membranous: spores 5-6 X 3·5-4· 5µ, fuscidulous, smooth,
ellipsoid: under rotten leaves: tropical Africa.
Compare L. palmatifidum, which seems identical. ? Ramaria Molleriana.
L. quangense P. Henn.
Engl. Jahrb. 22, 1895, 86.
-9 X 5-8 cm., coriaceous, tough, pallid, finely tomentose (dried): stems simple or
branched from the base, -3 cm. X 5 mm.: branches with broadened compressed axils;
branchlets repeatedly dichotomous, apices mostly multipartite, subulate, glabrous,
pale yellow.
Spores 6-7 X 5-6 µ, yellowish, smooth, ovoid.
On the ground in the forest: Angola (Quango River): Cameroons.
This may be a true Lachnocladium. It seems identical with L. palmatum. ? Ramaria
Molleriana.
L. ralumense P. Henn.
Engl. Jahrb. 25, 1898, 489.
-4 cm. high, white: stem short, terete or subcompressed, 2-3 times dichotomous or
subpalmatifid: branches terete, with scarcely compressed axils, strict or divergent,
with furcate acute tips.
Spores 4-4·5µ wide, white, smooth subglobose.
On the ground: New Guinea (Ralum, Neu Pommern).
This is said to be little branched and compressed. See L. pteruliforme.
L. reticulatum B. et C.
Journ. Linn. Soc. 15, 1876, 388.
Synonym: L. Hoffmanni P. Henn., Hedw. 1904 (teste Bres., Sacc., Syll. 23, 1925,
495).
-2·5 cm. high, caespitose, dichotomous, the branches divaricate-reticulate: stem
very short, 6 mm. long.
On the ground: Brazil (Traill, No. 127, Rio Jurua).
On the type-specimen in the Kew herbarium there are abundant pale yellowish
brown, punctately rough spores 5·5-7 X 3·5-4·5µ, which suggest that the species
belongs to the Stricta-group of Ramaria. Indeed, in habit and spore-size it is very
like my R. polypus. But the hyphal construction is quite different: there are many,
very narrow, 1·5-3µ wide, skeletal hyphae with thickened, colowless walls, and linear
lumen. I could find no generative hyphae, which had probably collapsed, but the
construction is clearly dimitic and, thus far, suggests the Pteruloid-series. But the
dichotomous branching of the fruit-body, which does not have subulate tips, and
the coloured spores indicate a different affinity which is borne out by the extreme
LACHNOCLADIUM 437
narrowness of the skeletals, those of the Pteruloid-series being typically rather wide
and often ill-defined. The species suggests a most interesting new genus.
Whether L. Hoffmanni is really a synonym remains to be proved. Its description is:
-12 cm. high, flesh colour, very much branched, scarcely stalked: branches
repeatedly dichotomous, broadly compressed, divergent, tips subulate: spores
4-5 X 3-4µ, smooth: on rotten wood: Costa Rica.
L. victoriense P. Henn.
Engl. Jahrb. 23, 1897, 543.
5-6 cm. high, fusco-violaceous, branched: stem 0·5-1 cm. thick, short, irregularly
branched: branches often confluent, terete or subcompressed: branchlets repeatedly
dichotomous, smooth, not compressed, divergent, with subulate tips.
On the ground: tropical Africa (Cameroons, Victoria Station).
Spores 7-8 x 3·5-4µ, fuscidulous epispore, smooth, ellipsoid or subovoid.
Dubious, possibly Thelephora. Said to be distinguished by the colour and the.
cylindric branches bent here and there (? the effect of drying).
L. violaceum Pat.
Journ. de Bot. 3, 1889, 27.
5-8 cm. high: stem 1-5 cm. X 2-3 mm., tuberous at the base, compressed, whitish:
branches 1 mm. wide, greyish violet, fastigiate, 2-3-chotomous, compressed·, canalicu
late, acute: hymenium unilateral, glabrous, whitish: tough.
Spores 4-5 X 3µ, yellowish, ovoid, smooth.
On the ground in the forest: French Guiana.
This may be Clavaria Zollingeri.
L. zandbaiense P. Henn. et Nym.

Mons. 1, 1900, 142.
-6 cm. high, stipitate, coriaceous, pale yellow: stem -2 cm. X 2-3 mm., terete,
fasciculately branched: branches terete with slightly thickened and scarcely compressed
axils, divergent, with setulae: tips long-subulate, dichotomous.
(Habitat?, terrestrial, tropical Asia).
Spores 5-7 X 4µ, yellowish white, asperate, ovoid.
Basidia 22-26 x 4-5 µ.
This was said to be allied with L. brasiliense but with rough spores. Ir. this respect
it suggests Scytinopogon.
LENTARIA gen. nov.

(Lentus, pliant: from the texture)
Fruit-bodies radially branched, polychotomous then dichotomous, varying sparingly
branched to subsimple in Lentariopsis, the uppersides of the main branches and axils
sterile in Eu-Lentaria, white, cream, alutaceous, tan, flesh-colour, ochraceous, or brownish,
often arising from a spreading villous or subtomentose, white or yellowish, sterile mycelial
felt spreading over the substratum: flesh tough, pliant: taste often bitter.
Lignicolous or on leaf-mould: 12 spp., temperate and tropical.
Spores white (to pale ochraceous ?), smooth, narrowly ellipsoid, aguttate.
Basidia clavate, with 4, in some species 2-8, sterigmata.
Cystidia none.
Hymenium usually thickening, the old basidia collapsing or, if abortive, becoming
rather thick-walled: subhymenial hyphae not inflating:
Hyphae monomitic, clamped, inflating, becoming thick-walled, not secondarily
septate.
438 LENTARIA
Colour in the hyphal walls, particularly those near the subhymenium.
T ype: L. surculus.
KEY TO THE SPECIES OF LENTARIA

Sp. 7µ or more long; fruit-bodies small to large, much branched:
hyphal walls becoming strongly thickened: Eu-Lentaria
Sp. ro-18x 3-6µ, elongate or subsigmoid
N. temp.: fruit-bodies often small L. byssiseda
Trop. L. surculus
Sp. 7-10X2·5-4µ
N. Amer. L. Micheneri
N. temp.:? sp. 2-2·5µ wide L. Patouillardii
Java L. tJibodensis
Sp. 4-7X2-4µ: fruit-bodies small, slender, often simple or
sparingly branched: hyphal walls slightly thickened: subgen.
Lentariopsis
Simple or sparingly branched: waxy-tough
Phycophilous, on sodden logs coated with algae
Sterigmata 4: sp. 4·5-7·5X1·8-3µ, mostly 5-6x2-2·5µ
(? 1-2 guttulate): -2 cm. high, white, yellowish, or
pinkish: cosmopolitan L. mucida
Sterigmata 6-8: sp. 5-7X2·5-3 µ: -4 mm. high, white to
watery ochraceous: Colombia L. coronilla
Not phycophilous: -3 mm. high, white or yellowish, 2-3 times
furcate: France . L. corticola
Mostly branched, often profusely: on coniferous wood: Europe
-7 cm. high, white then fuliginous, much branched: sp. ?:
Sweden, Finland L. virgata
-3 cm. high, often caespitose
White then greyish or chestnut-brown, or with fuscous
violaceous tips, sparingly branched, caespitose: sp.
5-6x2·7-3µ, I-several guttulate: 2-spored: on rotten
pine-wood: Europe . L. affiata
White or yellowish, much branched, attached by a byssoid
or floccoso-membranous mycelium
White to cream-citron, with many slender branches:
sp. 5-6 X3-4µ, aguttate: on rotten coniferous wood:
Europe L. epichnoa
White: sp. 5-6 X2·5-3 µ: rotten coniferous wood: Europe L. delicata
L. afflata (Lagger.) comb. nov.

Basinym: Clavaria affiata Lagger., Flora 19, 1836, 231: sensu Bourd. et Galz.,
Hym. Fr. 1928 (haud C. affiata sensu Bres. = Clavulina cristata).
0·5-2·5 cm. high, caespitose, rarely solitary, with 3-4 branches from.the base or at the
end of a short stem, the branches simple or once forked, rarely with simple fruit-bodies,
white then clear chestnut-brown, finally greyish pruinose, almost scurfy:(subdiaphanous,
white then greyish with fuscous violaceous tips, Fries).
On rotten trunks of Pinus: Sweden, France (rare).
Spores 5-6 X2·7-3µ, white, smooth, oblong, subarcuate, with I-several guttulae.
LEN'rARIA 439
Basidia 18-24 X 4-5µ: sterigmata 2, 3-4µ long.
Hyphae 4-8µ wide, thin-walled, sometimes rugose, without clamps: agglutinated
and subgranular in the subhymenium.
L. epichnoa may be identical, for they have the same habitat and spores. The
microscopical details are taken from Bourdot: possibly, the statement that there are
no clamps is erroneous, as in L. epichnoa. L. virgata seems also very similar, but with
longer branches as in L. delicata.
L. byssiseda nom. nov.
Synonyms: Clavaria byssiseda sensu auctt.: Bres., Ann. Myc. 1, 1903, 112: Coker,
Clav. U.S. Can. 1923, 152, t. 28, 59, 60, 87: Wehm., Pap. Mich. Ac. Sci. Arts Lett.
20, 1935, 260: Lundell et Nannfeldt, Sv. Svamp. fasc. 3-4, 1935, 20: (non Fr., 1821,
vide p. 265).
Clavariella byssiseda (Fr.) Karst., Finl. Nat. o. Folk 37, 1882, 190.
Ramaria byssiseda (Fr.) Quel., Fl. Myc. 1888, 462.
Clavaria pinophila Pk., Rept. N.Y. St. Mus. 35, 1884, 136.
? Clavaria soluta Karst. (see Ramaria soluta, p. 621).
Misidentifications:
Clavaria byssiseda sensu Karst., and Ramaria byssiseda sensu Quel. are probably
Ramaria crispula or R. stricta.
-6 X 4·5 cm., sometimes only 0·5-2·5 cm. high and wide, densely gregarious or
caespitose, with a short stem and more or less fastigiate branches, few or many, the tips
short and very acute or elongate-filiform, pale cream or yellowish white, becoming some
what ochraceous or tan, often tinged pinkish flesh-colour, then brownish, hazel, or
rufescent when old, the tips cream-white, arising from a small or spreading, white or
cream, tomentose mycelial patch and often with slender rhizomorphs permeating the sub
stratum: tough, pliable.
Stem 3-14 X 1-4 mm., or branched from the base, pallid white then concolorous,
rough, scurfy, or puberulous from the mycelium: branches 2-4-chotomous below and
radially branched, dichotomous above and often narrowly flattened palmate, numerous,
dense, and fastigiate, or short, few, and divaricate, the tips short and very acute,
becoming attenuate and elongate-filiform, divaricate or curved, in older or better
developed specimens (the tips often tinged green, Coker): hymenium sometimes absent
from the uppersides of some axils, generally covering the whole fruit-body almost
from the base: flesh pallid white or pallid ochraceous, tough, pliable, or fleshy-fibrous,
sometimes quickly turning brown on cutting or bruising (Coker).
On twigs, leaves, cones, and rotten wood of various deciduous and coniferous trees,
generally connected by slender rhizomorphs: north temperate region (Sweden,
Holland, France, Germany, Poland, U.S.A., C anada).'
Spores 10-18x3-6µ, mostly 12-16x3·5-5µ, white, cream, or pale ochraceous,
smooth, cylindric oblong, often sinuous or subsigmoid, blunt at the apex, acute at the
oblique base, thin-walled, aguttate.
Basidia 30-48 X 6-8µ, with 2(-4) sterigmata 7-9 µ long (Bourd. et Galz.): 25-30 X
8-10µ (Bres.): 7·5µ wide, with 4 sterigmata, hymenium 40-50µ thick (Coker).
Cystidia none.
Hyphae 3-5(-8)µ, thin-walled (Bourd. et Galz.): 4-6µ (Bres.): 4-5µ, clamped
(Coker).
The type-specimen of C. byssiseda Fr., according to Donk in litt., is Clavaria
himantia (Schw.) Fr., but for convenience I have retained the name in the sense of
Bresadola, Coker, Bourdot and Galzin, and Lundell. It is so near the tropical L.
surculus that it appears to be only a temperate variety with small fruit-body. Either
it has usually been collected while young and short, with congested branches, as it is
generally described from Europe, or it may mature in this state. Larger fruit-bodies,
1
Berkeley's record from Ceylon is Ramaria megalorhi::ra.
440 LENTARIA
corresponding with small specimens of L. surculus, have been found in N. America.
Lundell says that it grows only on dead needles of Picea in Sweden, but other authors
give a variety of coniferous and dicotyledonous hosts, as Salix and Populus in Poland.
Bresadola and Wehmeyer give the spores as white: Coker as cream, faintly brown,
or pale ochraceous: Bourdot and Galzin as white then pale cream: Lundell as pale
ochraceous. They appear white under the microscope. (The white spore-print of
L. surculus becomes yellowish on drying, and so do the spores when examined in dilute
potash.) In the following Table it will be seen that the spores are absolutely and
relatively wider in the European collections than in the American, which agree almost
exactly with those of L. surculus. Wehmeyer's collection from Nova Scotia, with very
narrow spores, may be L. Patouillardii, which is said to be common in Manitoba.
Ramaria crispula, R. stricta, R. apiculata, R. gracilis, and R. suecica have been con
fused with this species, but they have shorter, distinctly ochraceous, verruculose or
rough spores.
TABLE XXXI. SPORE-SIZE IN LENTARIA BYSSISEDA
E
Spore-size in µ. (average) Country Authority
(8-)12-18 X4--6 3·0 France Bourdot and Galzin
13-18 X4-6 3·1 Poland Bresadola
10-14X 4·5-5·5 2·4 Sweden Coker
(10-)12-16(-18) X (3•5-)4·5 3·1 Sweden Lundell
12-13·7 X3·7 3·5 U.S.A. (N. Car.) Coker (4395)
11-16·6X 3·4-3·7 3·9 " " " (5302)
12-15 X3·4-4·4 3·5 " " " (leg. Schallert).
10-14X 2·9-3·7 3·6 " (N.Y.) " (Burnham 90)
12-16·6 X 3·5-4 3·8 " " " (leg. Underwood,
Jamesville)
13-15X3·8 3·7 " (Conn.) " (s.n.)
12·5-14·5 X3·5-4 3·6 " " (type of C. pinophila)
10-14X2·5-3·5 4·0 Canada (Nov. Scot.) Wehmeyer
[13-16X3·5-4·5 3·6 Malaya Corner (L. surculus)]
L. coronilla (Martin) comb. nov.

Basinym: Clavaria coronilla Martin, Lilloa 5, 1940, 194.
2-4 X 0·2--0·5 mm., very small, simple or forked once, gregarious, white to pale watery
ochraceous, drying pale brownish ochraceous and horny: stem 0·5-1·5 mm. high,
distinct: club fusiform broadly clavate or subpileate: gelatinous-tough.
On rotten wood coated with blue-green algae: Colombia (1,250-1,500 m. alt.,
Hacienda Cincinnati, Dept. Magdalena, above Santa Marta).
Spores 5-7 X 2·5-3 µ, white, smooth, suballantoid.
Basidia 10-12 X 5-6 µ, elongate-urniform, with 6-8 sterigmata.
This differs from L. mucida in the more numerous sterigmata. Possibly it is more
nearly related with Clavulinopsis hastula with 4-6 sterigmata, but L. surculus may also
have 6 sterigmata.
L. corticola (Quel.) comb. nov.
Basinym: C/,avaria corticola Quel., Bull. Soc. Bot. Fr. 24, 1877, 326.
2-3 mm. high, white, yellowish at the base, very small, two to three times furcate,
mealy and white at the apex.
Lignicolous: France (Paris, Jura, spring).
Spores 5 µ, ovoid, white.
Such a small branched Clavaria is remarkable. It may be related to L. coronilla,
L. delicata, or L. epichnoa. Quelet's spore-measurements are not always reliable.
LENTARIA 441
L. delicata(Fr.) comb. nov.
Basinym: :][avaria delicata Fr., Syst. Myc. 1, 1821, 475: Burt, Ann. Mo. Bot.
Gdn. 7, 1922, 71, t. II, f. I.
Branched, white, lignicolous, -3 cm. high:stem short or none, attached to the wood
by a villous white mycelium:branches slender, elongate, fastigiate, acute.
On rotten wood(coniferous):Europe (rare:Sweden, ? France).
Spores 4 ·5-6 X 2·5-3µ, white, smooth, ellipsoid (Burt, for Friesian specimen in
Curtis's Herb.; Coker for Friesian specimens in herb. Kew.): 9 µ, ovoid-spheric,
white (Quel. FI. Myc. 462).
Hyphae -15 µ wide, inflated, clamped, with slightly, but distinctly, thickened walls
(Corner: co-type in herb. Kew.).
This little-known species appears not to have been collected since Fries described
it, unless by Quelet. It is certainly allied with L. epichnoa but has slender, Pteruloid
branches. On the co-type in the Kew herbarium there are many white spores of two
kinds, 9-12 X3·5-5 µ and 5·5-7 X3-3·5 µ, both distinctly apiculate as basidiospores,
but there are also many other foreign spores.
L. epichnoa (Fr.) comb. nov.
Basinym: Clavaria epichnoa Fr., Hym. Eur. 1874, 670: Quel., Ass. Fr. 1885, 450:
Bourd., Bull. Soc. Myc. Fr. 48, 1932, 208: Lundell et Nannfeldt, Sv. Svamp. fasc.
3-4, 1935, 20.
1-2·5 cm. high, white, branched, lignicolous, densely clustered or caespitose: stem
0·5-1 mm. wide, much branched above, arising from a byssoid white mycelium: branches
slender, filiform, divaricate or recurved, or fastigiate, acute, becoming flattened and
sulcate, white then &eam citron.
On dead coniferous wood: Europe (Sweden, France:rare).
Spores 5-6·5 X2·5-4µ, white, smooth, ellipsoid, aguttate.
Basidia 18-30 X4-6µ (Bourd.).
Hyphae 3-12µ wide, inflated, clamped, with slightly thickened(? submucilaginous)
walls, long-celled (Corner, from Lundell's collection in herb. Bourd., at Paris).
This species appears as the branched antecedent to L. mucida: the texture is similar
and the branches of L. epichnoa resemble single specimens of L. mucida. Whether it
is distinct from L. affiata needs further inquiry.
Bourdot and Lundell both give the spores as aguttate. They measure 5-6 X3-4µ
according to Lundell, and 4 ·5-6 X 2· 5-3 µ according to Bourdot: I found them to be
5·5-6·5 X2·5-3µ on Lundell's collection (18.9.1930) in herb. Bourd. Quelet gives
them as 11µ, 1-guttate, ovoid spheric(Ass. Fr. 1885), and 6 11,, ovoid (FI. Myc. 1888,
463): it is difficult to know what fungus he was dealing with.
Bourdot gives the hyphae as without clamps, which is an error.
L. Micheneri (B. et C.) comb. nov.
Basinym: Lachnocladium Micheneri B. et C., Grev. 1, 1873, 161: Burt, Ann. Mo.
Bot. Gdn. 6, 1919, 270:Coker, Clav. U.S. Can. 1923, and Journ. Elisha Mitch. Soc.
42, 1927, 253, t. 47, 49: Martin, Iowa Acad. Sci. 49, 1942, 149.
Synonym:? L. Atkinsonii Bres., Journ. Myc. 8, 1902, II9.
-4 cm. high, solitary or gregarious, with a short stem arising from a white byssoid
or tomentose mycelial patch, drying drab-grey, brownish, or subochraceous: stem
2-3 mm. wide, irregularly polychotomous: branches becoming dichotomous above,
rather congested with many acute tips: hymenium in patches on the stem, absent
from the uppersides of the branches, the sterile parts subtomentose in dried specimens:
tough, bitter (Burt).
On dead leaves, often of Fagus or Quercus:U.S.A., Canada.
Spores 8-9 X 3-4µ, white, slightly rough (Coker, for the co-type Curtis 3534):
6-8·5X3·6-4µ,, white, smooth (Coker, 1927): 6-12X3-3·5µ, white smooth (Burt):
7-9 X3·5-4µ,, white, smooth (Corner, for the type in herb. Kew.).
442 LENTARIA
Hyphae 3-6µ wide, monomitic, clamped, the walls becoming slightly or rather
strongly thickened, scarcely inflated (Corner, for the type in herb. Kew.).
The form and microscopic structure of the type-specimen in the Kew herbarium
is exactly that of Lentaria byssiseda, from which it seems to differ only in the shorter
spores. This was Bresadola's opinion. Coker suggested, at first, that it was near to
Ramaria stricta, pallid forms of which could be distinguished by their rough ochraceous
spores: in 1927 he decided that it had smaller and more delicate fruit-bodies and
truly hyaline, or white, thin-walled spores. On the other hand, Martin refers to
L. Micheneri a fairly abundant fungus in the neighbourhood of Iowa City which has
a brownish ochraceous spore-print, though almost colourless spores under the micro
scope, and a slightly nauseous, if tardy, taste, not bitter: (he also found thick-walled
cystidium-like structures, which may be only abortive basidia which have become
thick-walled in the thickening hymenium, as in L. surculus). I conclude that two
species must have been confused under this name and that the true L. Micheneri is a
white-spored ally of L. byssiseda and L. surculus very near to L. Patouillardii, if not
identical, and that there is a Ramaria of very similar appearance, perhaps R. apiculata
or R. stricta. Possibly proof is to be found in Lachnocladium Atkinsonii which seems
intermediate in size of fruit-body, between L. byssiseda and L. surculus, and to have
the white smooth spores of L. Micheneri. There is no type-specimen, according to
Coker. Its description is:
Lachnocladium Atkinsonii Bres., I.e.: 11-13X5-6 cm., somewhat coriaceous: stem
6-7X1 cm., elongate, compressed-canaliculate, pallid, tomentose, somewhat 4-fid at
the apex: branches compressed, sulcate, repeatedly verticillate or dichotomous,
tomentose on the sterile side, lurid ochraceous: branchlets subterete, furcate at the
apex, straw-yellow: stem somewhat waxy: spores 9-10X4·5-5µ (7-10X3·5-4·5µ,
teste Atk.), white or pale straw-colour, smooth, subcylindric to amygdaliform-oblong:
U.S.A. (N. Car.).
L. mucida (Fr.) comb. nov.

Basinym: Clavaria mucida Fr., Syst. Myc. 1, 1821, 476: Lind, Bot. Gaz. 37, 1904,
62, f. 16, 17, and Rostrup's Herb. Fungi 1913, 368: Coker, Clav. U.S. Can, 1923:
Lundell, Sv. Svamp. fasc. 3-4, 1935, 20: Martin, G. W., Lilloa 5, 1940, 191, and
Lloydia 7, 1944, 78.
Synonyms: C. mucida var. Curtisii B., Grev. 2, 1873, 17 (teste Coker).
? C. albipes Mont., Ann. Sci. Nat. 2, 18, 1842, 244: [ = Calocera albipes (Mont.) B.,
Grev. 2, 1873, 18).
? C. alba Lloyd, Myc. Notes 67, 1922, II53, f. 2246: (non Pers.).
0·3-2 cm. high, mostly simple, sometimes forked into z-6 linear curved ascending
branches, or incised or minutely cristate at the apex, slender, 1 mm. thick, gregarious,
often in large colonies, not fasciculate, solid, acute, rarely clavate-obtuse, cylindric
fusiform, attenuate into a scarcely distinct stem, white, yellowish, pale cream, or even
pinkish, apex sometimes becoming lateritious, brownish or blackish: delicate but
tough, not breaking on bending, waxy-tough: smell none, taste woody.
On rotten wood (ad terram, Fries!), associated with a film of Chlorococcoid algae
on the surface of the wood: Europe (Sweden, France, rare): Siberia (on Populus
tremula): U.S.A. (widely distributed, uncommon, often on Nyssa), Canada: Japan:
West Panama (Chiriqui), Columbia (Sierra Nevada de Santa Maria, c. 1,400 m.):
[? Australia (N.S.W.), New Zealand, Tasmania].
Spores 4·5-7·5X1 ·8-3 µ white, smooth, narrowly or oblong ellipsoid, 1-2 guttulate
(Lundell), aguttate (Coker).
Basidia 15-21X4-5µ (Bourd.): sterigmata 4 (Bourd., Coker, Martin).
Hymenium extending over the tip of the fruit-body (Coker), ? not thickening:
cystidia none.
LENTARIA 443
Hyphae 3-6µ wide, often dilated to 8µ at the septa, scarcely inflated, clamped
(Coker, Comer, Martin), with few or no interweaving hyphae.
var. rosea Bres.
Sacc., FI. It. Crypt. Hymenom. 1916, 1251: Syll. Fung. 23, 1925, 490.
Fusiform, attenuate to each end, simple or apex bifid, or with 2-3 short lateral branches,
smooth, glabrous, sometimes flattened, 1-2·5 mm. wide in the middle, white below,
rose in the middle, pale rose at the apex, yellow or greenish with age: flesh white,
�tuffed.
In mossy places under Alnus, northern Italy (Trentino): widely gregarious, scarcely
visible, very small.
This is a phycophilous species. The mycelium grows in a scum of Chlorococcus,
forming a green film on well-rotted wood, as Fries, Persoon, and Schweinitz noted.
The hyphae grow so intricately among the algal cells that Lind and Coker have sug
gested that the fungus is a lichen rather than a true basidiomycete: however, there are
no algae in the fruit-body and it is a case of semiparasitism rather than symbiosis.
According to Coker it grows only on logs of Nyssa in the United States, but in Sweden
and Siberia it grows on Populus tremula according to Lundell and Pilat. If the wood
in any way determines the occurrence of the fungus, it seems to be by way of the alga,
as host, rather than a direct lignicolous effect: Doty is the only recent author to report
the species as growing on the ground: (and var. rosea is possibly a form of Clavulinopsis
vernalis). In Sweden, according to Fries, the fungus fruits throughout the year.
The spores seem to vary in size. Lundell is emphatic that they are 5·5-6 X2-2·5 µ
and 1-2 guttulate, in contrast with the rounded-ellipsoid, aguttate spores of L. epichnoa,
5-6 X3-4µ. Other authors give these data:
Saccardo: 5-6 X2-3µ, 6--7 X3-4µ.
Bourdot: 6--7·5 X3-3·5µ (Sweden).
Pilat: 5·5-6 X2-2·2µ (Siberia).
Coker: 5·2--7·4 x 1·8-2·3µ (U.S.A.).
Martin: 4·5-6 X2·5-3µ (Panama, Colombia).
Doty: 7--7·5 X2-3µ (Oregon).
Possibly Bourdot's specimens were very young or depauperate L. epichnoa, as they
are given as 3-6 X1 mm., simple (Bull. Soc. Myc. Fr. 48, 1932, 209).
There seems to be no type-specimen of Clavaria albipes, but its description indicates
L. mucida: gregarious, simple, strict, attenuate at each end, glabrous, acute, pale
rufescent (? dried): stem attached by a dilated white base: on semi-putrid and slimy
wood: Ohio.
Clavaria alba, named provisionally by Lloyd for specimens from Tasmania, seems
identical, with spores 6 X2µ. It is the only reliable record of L. mucida from the
Australasian region.
The systematic position of L. mucida is debatable. By analogy with L. epichnoa,
L. delicata, and L. aifiata, I have placed it in Lentaria: they are lignicolous, with
small white, often sparingly branched fruit-bodies and similar small spores, so as to
suggest the antecedent of the simple L. mucida. On the other hand, if the lignicolous
habitat of L. mucida is secondary, dependent on the lignicolous habitat of the Chloro
coccus, it may be related with Clavulinopsis vernalis. L. coronilla, with 6-8 sterigmata,
is very similar and, by analogy with the 2-6-spored L. surculus, confirms the position
of L. mucida in Lentaria.
Martin suggests the species is related with Pistillaria uncialis (= Gloeocoryne
uncialis), but the stem hyphae are not agglutinated, as in.Pistillaria, and the incipient
branching of the fruit-body does not accord. In the tissue of three collections from
tropical America, Martin found abundant chlamydospores, 10-13 x 5-6µ, thick
walled, ovate, or suballantoid, possibly apiculate at one end: they may have been
transformed basidia, but none were seen attached.
444 LENTARIA
. Compare Clavaria fossicola which is truly terrestrial, has a thickening hymenium,
and is without clamps.
L. Patouillardii (Bres.) comb. nov.
Basinym: Clavaria Patouillardii Bres., Fung. Trid. 2, 39, t. 146, f. 1: le. Myc.,
t. 1093: Coker, Clav. U.S. Can. 1923, 156, t. 60, 88: Buller, Bisby, Dearness, Fungi
of Manitoba 1929, 92.
-6·5 cm. high, much branched, drab-white or dingy clay-white, becoming pale tan or
pale alutaceous, with yellowi.sh-white tips, rather slender, flaccid, pliant, arising from a
conspicuous white tomentose or farinoso-flocculose incrusting mycelium.
Stem 1-1·5 cm. x 2-5 mm., breaking up at the top into numerous branches: branches
rather crowded but lax, many, ascending, polychotomous then dichotomous: flesh
white, not changing colour: taste bitter, or none (Coker): no smell.
On dead leaves and rotten wood in coniferous and frondose woods: France, Italy,
Tunisia, U.S.A. (Pa., N.Y.), Canada (Man., common).
Spores 7-8(-10) X 2-2·5 µ., white, smooth, ovoid-elongate, often subsinuous,
granular-guttulate: (8-IO X 2-2·5 µ., Bres.: 6·5-8 X 2·2-2·5µ., Coker).
Basidia 40-50 X 7-8µ., with 4 sterigmata.
Hyphae 2--7µ. wide, monomitic, clamped, the walls 0·5-1·5µ. thick, becoming thick
walled mostly in the lower part of the fruit-body.
The hyphal details I have added from a collection of Bresadola's in the Kew
herbarium, which may be a co-type, and from another collection by R. Heim (on
larch-needles, France, Sept. 1929). In both, I find spores 7-10 X 3-4µ., and only
immature spores, attached to basidia, as narrow as Bresadola and Coker give (2-2·5 µ.).
The species is certainly allied with L. byssiseda and L. surculus and, perhaps, it is
identical with L. Micheneri.
L. surculus (B.) comb. nov.
Basinym: Clavaria surculus B., Hook. Journ. Bot. 1, 1842, 154, t. 6, f. 5.
Synonyms: ? C. compressa B., Ann. Mag. Nat. Hist. rn, 1842, 383, t. 12, f. 16 (non
C. compressa Schw., 1832 = Clavulinopsis fusiformis): = Lachnocladium compressum
(B.) Lev., Ann. Sci. Nat. Ser. 3, 1846, 153: Rick, Ann. Myc. 5, 1907, 335.
? C. trichotoma Lev., Ann. Sci. Nat. Ser. 3, 2, 1844, 216: Sacc. et Pao!., Att. R.
Inst. Ven. 6, Ser. 6, 1888, 14.
C. trichoclada Sacc. et Paol., Att. R. Inst. Ven. 6, Ser. 6, 1888, 14, t. 5, f. 6: Syll.
Fung. 6, 1888, 703.
? Lachnocladium tonkinense Pat., Joum. de Bot. 1891, 314, t. 4, f. 5.
Text-Figs. 2G, 193-9: Plate 5.
-11 cm. high, solitary, gregarious or subcaespitose, with a distinct stem and narrowly
fastigiate branches with progressively lengthening internodes and filiform or subulate,
elongate tips, tough, elastic, white then pale cream, alutaceous, or pale ochraceous, more or
less tinged pinkish flesh-colour or pale fawn, rather pale clear ochraceous flesh-colour in
full growth, tips white, paler and dingy with age and becoming somewhat cinereous at
the base: arising from a white, pale yellowish, or cream, villous, sterile mycelial felt,
-25 cm. wide, incrusting the substratum, and with fine white mycelium strands
permeating the substratum.
Stem 0·5-4·5 cm. X 1·5-5 mm., sometimes very short or absent, base villous, else
where smooth or thinly felted, dividing into 2-3-4 branches at the apex: branches
1-2 mm. wide and 2-4-chotomous below, becoming dichotomous distally, radially
and alternately branched, not or slightly flattened below the rounded axils, terete,
smooth, glabrous, at first lax and divaricate, becoming repeatedly dichotomous, the
branches narrowly fastigiate, often strictly parallel, erect, with lengthening internodes and
long, tapered, subulate or filiform tips, old fruit-bodies often with the branches gnarled
from the irregularly thickened hymenium: hymenium waxy-smooth, pale drab-grey or
TExT-FIG. 193. Lentaria surculus, X 1: growing from old Bauhinia-pods TExT-FIG. 194. Lentaria surculus: young fruit-bodies with mycelial
in the ground. felt, on a twig, x 2.
LENTARIA
pale fawn-drab, continuous almost from the base of the stem, on all sides of the
branches or poorly developed and sterile on the uppersides: flesh tough, pliable,
elastic, slightly floccoso-felted, very tough and coriaceous at the base of the stem,
pallid white: smell often slightly fruity and pleasant (like plums): taste somewhat
bitter or subacrid.
TEXT-FIG. 195. Lentaria surculus: old fruit-body, the mycelial felt

completely mummifying the twig, x 2.
On dead sticks, branches, palm-fronds, woody fruits, and so on in the forest:

apparently pan-tropical, common in tropical Asia.
Spores 13-16 X 3·5-4·5 /L, white, smooth, elongate pip-shaped, often subsigmoid,
rounded and blunt at the apex, subacute at the base, aguttate: (curiously rare and
difficult to find in dried specimens): young spores very narrow and elongate, 10 x 2·5 /L·
Basidia 30-50 X 7-9 /L, clavate, with densely granular-guttulate cytoplasm: sterig
mata 2-3-4-5-6, mostly 4, 5-7 X 1-2 /L, slender erect and slightly curved, occasionally
spreading almost at right angles from the apex of the basidium: occasionally young
basidia producing 1-2 sterile sterigmata 8-30 fL long.
I
I. •
TExT-FIG. 196. Lentaria surculus: medullary hyphae of the fruit-body: on the right,
two very thick-walled hyphae from the stem, and the tips of two hyphae in the
growing point, x 500.
LENTARIA
Cystidia none.
Hymenium c. 35-40µ thick at first, thickening to 400µ, not layered, the old basidia
.:ollapsing and leaving gaps: subhymenium composed of loosely interwoven hyphae, 2-4µ
wide, with firm dry walls (scarcely thickened), short-celled, not inflating, with abrupt
transition from the longitudinal hyphae of the flesh to the perpendicular hyphae of
the subhymenium.
Hyphae monomitic, clamped, not secondarily
septate, 3-10 µ wide, occasionally with local
subglobose swellings -15 µ wide, the walls pale
ochraceous or pale tan and distinctly thickened
0·6-1·5µ (2-3µ at the base of the stem), smooth,
the cells 40-300 µ long, generally unequally and
irregularly inflated in different parts, often with
undulate walls, often kinked or somewhat con
volute, the narrower hyphae often with linear
lumen, the clamps often swollen, -15µ wide, and
variously shaped, not or slightly constricted at
the septa.
Flesh composed of longitudinal hyphae
without interweaving laterals, the hyphae
narrower, denser, slightly thicker-walled and
shorter-celled in a layer 30-40µ thick next the
subhymenium.
Growing tips with hyphae 2-2·5µ wide, the
clamps forming 40-100µ behind the tips, the
subterminal cells 30-100 µ long on delimitation
(shorter on the subhymenial hyphae): the
hymenium developing shortly behind the apex
and very gradually.
Stem mostly covered with fertile hymenium,
but the tomentose surface at the base composed
of 2-4µ wide, interwoven, cylindric, branched
hyphae, clamped and with slightly thickened
walls, like the mycelial hyphae, this loose
floccose layer creeping up the stem and covering
the old hymenium.
TEXT-FIG. 197. Lentaria surculus:
Sterile hymenium on the uppersides of the
clamp-connexions on medullary hyphae
of.the fruit-body, x 1,000. branches as a rudimentary, loose hymenium
with sterile basidia, not or little thickened.
Colour caused by the pale ochraceous or pale tan walls of the hyphae, especially
those next the subhymenium.
This fungus is common in Malaya, but it is not infrequently sterile and the spores
are in all cases difficult to find on dried specimens. The type of C. surculus in the
Kew herbarium, from the Philippines (Cuming 2042), has exactly the same structure,
with thickening hymenium, but I could find no spores. It was said to be growing
from the ground, but the fruit-body may well have been attached to a buried stick or
woody fruit. C. trichoclada was described from Malaya, on fallen rotten branches;
the illustration and description both indicate L. surculus: the spores were said to be
14 X 9 µ., fuscous ochraceous, but this is probably erroneous. C. trichotoma from
Borneo also seems identical: its description is:
Caespitose, fleshy-coriaceous, glabrous, fuliginous, branches trichotomous, fasti
giate, terete, thickened upward, tips bifid: Borneo (Korthals, in Herb. Lugd. Bat.).
Saccardo and Paoletti refer to this dubious species a collection from Malaya
described thus: wholly brown, rather rigid, 3-4-chotomous, repeatedly and regularly
LENTARIA 449
branched, the branches ascending, arcuate from the base, thickened above: on trunks,
Malaya: rather a Pterula.
Lachnocladium tonkinense may also be this species, with wrongly ascribed spores:
the description, of the dried specimen, is:
TEXT-FIG. 198. Lentaria su.rrn/11s: young hymenial hyphae from near the growing point
of a branch, an<l the outt'r and inner parts of the thickened hymenium: , �oo.
Stem erect, rufescent, dichotomous, surrounded by abundant white floccose

mycelium: branches cylindric, yellowish brownish, smooth on one side, ochraceous
pulverulent on the other: spores yellowish, ovoid, smooth, on dead stems of mono
cotyledons: Tonkin. (Said to be near Septobasidium rameale.)
su9 Gg
450 LENTARIA
Of Lachnocladium (Clavaria) compressum B., Rick says it is the commonest and most
variable species of Lachnocladium in S. America and that it is easily known from the
abundant development of the mycelium. This is the macroscopic character of Len
taria surculus in Malaya, its mycelium often forming cream or pale ochraceous felts
for 20-30 cm. over rotting wood.
See also the remarks under L. byssiseda. Pterula pungens and P. Janseniana may also
be synonyms.
s�
f'
�
�
TEXT-FIG. 199. Lentaria surculus: basidial variation, 2-3-4-6-spored, with erect and dis
traught sterigmata: two abnormal sterile basidia (bottom right): developing and mature spores:
X r,ooo.
L. tjihodensis (P. Henn.) comb. nov.

Basinym: Clavaria tjibodensis P. Henn., Mons. 1, 1900, 141.
-7 cm. high, fleshy, very much branched, Ju.rid yellow-brown: stem thick, covered
with membranous white mycelium: branches fastigiate, terete, smooth, repeatedly
dichotomous, axils compressed, sulcate, tips subulate or subcristate.
On stumps: Java (Tjibodas, c. 1,500 m. alt.).
Spores 7-9 X 3-4µ, white, smooth, ellipsoid.
The position of this fungus is doubtful. The membranous mycelium, the colour
and branching of the fruit-body, the white spores, and the habitat indicate Lentaria.
Compare also Ramaria Moelleriana.
L. virgata (Fr.) comb. nov.

Basinym: Clavaria virgata Fr., Syst. Myc. 1, 1821, 472.
-7·5 cm. high, much branched, white then fuliginous: branches elongate, sulcate,
acute.
On rotten pine-wood: Europe (Sweden, Finland).
Little known. Coker suggests it is Ramaria apiculata. On the other hand it seems
like a large L. epichnoa or L. delicata.
MUCRONELLA 4 51
MUCRONELLA Fr.
Hym. Eur. 1874, 629: Lloyd, Myc. Notes 39, 1915, 531: Miller, Mycol. 26, 1934,
215.
Synonym: Mucronia Fr., Summ. Veg. Scand. 1849, 329: (non Mucronea Benth.,
1836).
Fruit-bodies simple, spine-like, cylindric, subulate, or conical, small, scattered or
gregarious (in a few, perhaps exceptional, species fasciculate), individually distinct,
without subiculum,pointing vertically down: apex sterile, acute.
Lignicolous and corticolous: temperate and tropical, 9 spp.
Spores white, smooth, thin-walled, ellipsoid to globose,aguttate.
Basidia small, with 4 sterigmata.
Cystidia none.
Hyphae monomitic,clamped,not secondarily septate,slightly inflating,thin-walled.
Compare Deflexula, to which several species of Mucronella may belong, and
Ceratellopsis.
KEY TO THE SPECIES OF MUCRONELLA

2-4 cm. long: stems 1-2 cm., slender, densely fasciculate: trop.
Africa M. tenuipes
-5 mm. long, white or yellowish, with a very short stem or sessile
2-5 mm. long,caespitose: sp. subglobose
Sp. 8 x 6µ.: U.S.A. M. alba
Sp. 4-6 µ.: trop. Africa M. togoensis
Shorter,not caespitose: sp. ellipsoid
Temp.
Sp. 2-4 x 2 µ: Finland . M. subtilis
Sp. 4-6·5 X2·5-3·5µ.
Gregarious in small patches M. aggregata
Scattered M. calva
Trop.
Brazil: 0·5-1 mm. long: sp. 4-6x2·5-4 µ. M. brasiliensis
Malaya: -o·5 mm. long: sp. 4-4 · X µ.
7 3 . M. pusilla
Sp. unknown
-2· 5x0·3 mm., 3-8 fasciculate, arcuate-pendulous,cinereous
white: Argentine M. argentina
M. aggregata Fr.
Hym. Eur. 1874,629: Lloyd, Myc. Notes 39,1915,531,f. 724,725: Miller,Mycol.
26, 1934, 215.
Synonyms: Mucronia aggregata Fr., Monogr. 2,280.
Mucronella minutissirna Pk.,Rept. N.Y. St. Mus. 44, 22.
M. abnormis P. Henn, Verh. Bot. Ver. Pr. Brand. 40, 122, t. 1, f. 2.
? M. ramosa Lloyd, Myc. Notes 66, 1922, 1107, f. 2036.
0·5-1·5 mm. long,white then pallid, subulate, short,free,in dense clusters 0·5-2 cm.
wide,individually separate, subiculum none or of scattered hyphae.
On old rotten wood: Europe, N. America.
Spores 4-6·5(-7) X 2·5-3·5(-4)µ., white, smooth,ellipsoid.
Basidia 10-20 X 3-5 µ.
Hyphae 2-4µ. wide,in places 6-7 f' wide,thin-walled,clamped,with oxalate crystals
(Miller).
452 MUCRONELLA
Miller says the three synonyms appear indistinguishable from M. aggrega a: it is
possible, however, that there is some difference, whether their spores are amyloid
or not.
M. minutissima: 0·5 mm. long or less: spores 4 >, 2·5 µ. (Pk.), 6 x 4µ. (Lloyd): on old
pine-wood: Canada, U.S.A.
M. ramosa: fruit-bodies fasciculate so as to appear branched: spores 6 X 3µ.(Lloyd):
on wood: Alabama.
M. abnormis: yellow spines 0·5-1 mm. long, gregarious or crowded: spores? 4-5X
3-4µ.: on a trunk from the Cameroons, in the Bot. Gdn., Berlin.
Specimens of M. aggregata, which I collected in the neighbourhood of Cambridge
(England) in 1925-8, had spores 4-6X3µ., basidia 10-16X3·5-4·5µ., 4 sterigmata
3µ. long, no cystidia, and thin-walled hyphae 3-6µ. wide. The structure of their fruit
bodies was the same as that shown for M. pusilla on p. 453, with the hyphae forming
a short-celled and almost pseudoparenchymatous tissue at the base of the very short
stem, which was attached by few narrow hyphae, 2-3 µ. wide, to the wood without a
subiculum. Young fruit-bodies, therefore, differ from those of M. pusilla only in their
slightly longer spores, though old fruit-bodies grow 3-4 times as long. M. aggregata,
in England, fruits from autumn throughout the winter to early spring (September to
March) and is, thus, a low-temperature fungus compared with the tropical M. pusilla.
M. alba Lloyd
Myc. Notes 61, 1919, 880, f. 1509.
-5 mm. long, white, caespitose, clustered on bark, without subiculum.
Spores 8X6 µ., subglobose: basidia clavate.
On bark: Washington.
This is said to be like M. aggregata but much larger. If correctly described the
spores are distinctive, and suggest Deflexula.
M. argentina Speg.
Fg. Arg. 1899, 178.
Subiculum none: spines 2-2·5X0·3 mm., solitary or 3-8 fasciculate, arising from
a minute tuberculiform ·common base, distinct, free, fusoid-terete, cinereous, arcuate,
arcuate-pendulous, fleshy-soft.
On bark of living mossy trunks of Celtis: Argentine.
? Deflexula.
M. brasiliensis nom. nov.
Synonym: Clavaria mucronella Bres., Hedw. 1896, 290.
0·5-1 mm. high, densely gregarious, subulate, simple or rarely forked at the tip,
white, subpruinose, filiform, slightly white fibrillose at the base.
On rotten wood: Brazil (Blumenau).
Spores 4-6X2·5-4µ., white, smooth, obovoid.
Basidia 15-20X6-7 µ., 4-spored.
This may be M. pusilla, but I have always found it to be smaller and with rounder
spores.
M. calva Fr.
Hym. Eur. 1874, 629: Bres., Ann. Myc. 1, 1903, 90.
Basinym: Isaria calva Fr., Syst. Myc. 3, 1832, 277.
Synonym: Mucronia calva Fr., Summ. Veg. Scand. 1849, 329.
1-3 mm. long, white, then pale, scattered, rigid, thin.
On rotten pine-wood: Europe, U.S.A.
Spores 4-6 X 3µ., oblong, hardly depressed: 4-6 X2-3µ. (Bres.).
Basidia 12-18X4µ.: 15X4µ. (Bres.).
MUCRONELLA 453
Hyphae 3-6 µ wide, thin-walled, emerging in a sterile bundle at the tip of the fruit
body: 3-7µ (Bres.).
Except for the scattered habit, this seems identical with M. aggregata. Bourdot and
Galzin say the species are too close. Miller thinks them identical, but Lloyd describes
M. calva with spines 6-25 mm. long. According to Bourdot and Galzin, M. calva
sensu Pat. (Bull. Soc. Myc. Fr. 5, p. 32) is a young state of Odontia arguta with
practically no subiculum, which grades into the normal mature state of Odontia.
According to Bresadola, in some forms there may be a slight sterile subiculum com
posed of hyphae 4-12µ wide, joining the spines.
M. pusilla spec. nov.
Text-Fig. 200.
Fruit-bodies -500 X 50-130µ, pale yellowish white, dingy, or ochraceous with age,
conical, straight, slightly flexuous, pointing vertically down, rather waxy, solid, with
a short, sterile stem 10-75 X 30-50 µ, apex acute, simple, persistently sterile, densely
gregarious in small patches -I5 mm. wide, but individually distinct, without subiculum.
On decorticated branches and rotten wood in the forest: Malaya (Johore, Mawai,
Sept. 1934: Singapore, Nov. 1943: probably common).
'TEXT-FIG. 200. Stages in the development of the fruit-body of Mucro

nella pusilla: X 500.
Spores 4-4 ·7 X 2 ·7-3 ·3 µ, white, smooth, ellipsoid, flattened adaxially, -0btuse, thin
walled, aguttate.
Basidia u-15 x4-5µ, subclavate: sterigmata 4, 1·5-2µ long.
Cystidia none: the apex of the spine composed of a fascicle, I 5-35µ wide, of narrow
hyphae, projecting 15-50µ.
Hymenium continuous except at the sterile base and growing apex of the fruit-
454 MUCRONELLA
body, the stem with a few scattered sterile basidia but no caulocystidia: subhymenium
4-8µ thick, composed of short uninflated cells 3-9·5 X 2·5-3µ, 2-3 cells thick.
Hyphae monomitic, septate, clamped, slightly inflating, thin-walled (becoming
slightly thick-walled at the base of the fruit-body), with numerous tetrahedral crystals
or irregular crystalline masses, 4-18 µ wide, among the hyphae of the flesh, mostly in
the subhymenium: hyphae 2-3µ wide at the growing apex, the cells 6-12µ long on
delimitation and gradually enlarging to 9-45 X 4-7µ, the hyphae longitudinal with
the cells shortest at the base of the fruit-body, slightly divergent into the hymenium,
the tissue compact: clamp-connexions relatively large, one at every septum, sometimes
branching from the clamp: H-connexions frequent.
Mycelial hyphae with short cells 4-16 X 2-3·5 µ, walls firm and slightly thickened,
the septa sometimes without clamp,:;, apparently without H-connexions.
Yellow colour of the fruit-body in the hyphal walls, especially where slightly
thickened at the base of the stem.
This seems identical with M. aggregata or M. minutissima, but until their hyphal
construction is adequately known it seems unwise to identify the tropical species
with them.
M. subtilis Karst.
Symb. Myc. Fenn. 23, 1: Sacc., Syll. 9, 1891, 218.
Spines -1 mm. long, crowded, very slender, white, acuminate, drying yellowish,
without subiculum.
On rotten wood of Pinus: Finland (Mustiala).
Spores 2-4 X 2µ.
Perhaps only M. aggregata, but the spores very small.
M. tenuipes Lloyd.
Myc. Notes, 55, 1918, 789, f. 1187.
Fertile heads 1-2 cm. long, unbranched, pendulous, densely fasciculate, with
slender, hair-like stems 1-2 cm. long, the heads drying yellowish brown and angular
flattened: subiculum none.
Spores ? 4 X 1µ, white, allantoid.
? on wood: tropical Africa (Angola).
Certainly not Mucronella: possibly Hormomitaria.
M. togoensis P. Henn
Engl. Jahr. 23, 1897, 544.
Fruit-bodies 3-3·5 X 0·5 mm., white, pendulous, curved, subulate, connate at the
base in fascicles of 5-16.
On tree-bark: tropical Africa (Togo).
Spores 4·5-6µ, white, smooth, globose.
Basidia 12-15x7-8µ.
This may be Mucronella with round spores, as M. alba, or Deflexula with small
spores.
MYXOMYCIDIUM Mass.
Kew. Bull. 1899, 179: Linder, D. H., Mycologia 26, 1934, 332.
Fruit-body simple, pendulous, watery-gelatinous, with short, firm, sterile stalk:
hymenium covering the more or less obconic head.
On dead wood: 3 spp., Tasmania, British Guiana, Tennessee.
Spores white, smooth, ellipsoid (? aguttate).
Basidia clavate, sterigmata 2-4.
Cystidia none.
MYXOMYCIDIUM 455
Hymenium evidently thickening.
Hyphae monomitic, with or without clamps: with gelatinous walls, at least in the
head, probably not inflated.
Type-species: M. pendulum Mass.
KEY TO THE SPECIES

Basidia clavate, not conspicuously narrowed below, 25-28 X 5-7 µ,:
sterigmata 2-4, 2·5-4µ, long: sp. 6-7X4·5-5µ: Tasmania M. pendulum
Basidia conspicuously narrowed below: sterigmata 4: sp. 5-6· 5 X
3-3·5 µ,
Clamps absent: sterigmata. short, 2 ·5-4 µ, long M. guianense
Clamps present: sterigmata 5·5-9µ, long M. nodosum
This genus has been revised by Linder, who has added the two species M. guianense
and M. nodosum. I have copied his specific descriptions and key.
Linder considers the genus allied with Vuilleminia, which may be true, but I do
not agree with his opinion that it cannot belong to the Clavarioid fungi. Thus, Linder
gives as distinctive of Clavariaceae the absence of gelatinous fruit-bodies, the narrowly
clavate form of the basidium, and the absence of proliferating basidia, that is, the
absence of a thickening hymenium. But these statements are not true. If Myxo
mycidium grew erect on the ground, it would be identical generically with the
gelatinous Clavulinopsis hastula, so I consider it a member of the Clavariadelphus
series. It is possible that the fruit-bodies of Myxomycidium are not truly inverted
but ageotropic, as in Physalacria, and pendulous through the weight of the watery
head. The absence of cystidia precludes affinity with Physalacria and Hormomitaria.
�QDOO
TEXT-FIG. 201. i\,lyxomycidium: a, M. guianense, spores and

basidia, x c. 1 ,ooo; fruit-bodies, X t: b, M. pendulum, X c. 1 ,ooo:
c, M. nodosum, x c. 1,000: (from Linder.)
Linder emphasizes the division of many basidia of Myxomycidium into the so-called
hypobasidium and epibasidium. I doubt if the words have any morphological signi
ficance in this case, because the elongation of the basidium to form an epibasidium
may be only the resumption of apical growth to project the tip of the basidium beyond
MYXOMYCIDIUM
the jelly of the head, and thus, perhaps, dependent on the weather: similar, though
less conspicuous, proliferations occur in the subgelatinous Clavariafossicola, which is
possibly allied.
In M. guianense, Linder found the division of the fusion-nucleus to be transverse in
the clavate apex of the basidium, and the divisions of the daughter-nuclei to be
longitudinal or oblique in the proximal part of the clavate head of the basidium. If
an 'epibasidium' is formed, it develops during the division of the daughter-nuclei.
The haploid number cif chromosomes is four. There appears to be no connexion
between the daughter-nuclei and the development of the sterigmata. Only daughter
nuclei are formed and they become smaller as they pass into the spores; it is possible
that the mature spores become binucleate.
M. guianense Linder
I.e. p. 340, f. 3-4, pl. 39.
Text-Fig. 201.
1-3 cm. long, pendulous, watery gelatinous, shortly stipitate, lanceolate, the apex
acute at first, hyaline or milky white, tinged ochraceous when old: becoming very
softly watery at maturity.
On a decaying fallen tree-trunk: British Guiana (Bartica).
Spores 5·5-6·5X2·5-3·5µ,, white, smooth, ovoid or ellipsoid.
Basidia 15-23X4·5-5·5(-7)µ,, clavate, constricted towards the base (1·5-2µ, wide),
frequently with swollen hypobasidium: sterigmata 4, 2·5-4µ, long.
Hyphae slender (? uninflated), without clamps: longitudinal in the stem and
medulla of the head: with a cortex, 100-150µ, wide, of radiating hyphae between the
medulla and hymenium in the head: hymenium and subhymenium 25-40µ, thick
(? thickening), covering the entire head.
M. nodosum Linder
I.e. f. 1-2.
Text-Fig. 201.
As M. guianense, but tinged greenish ochraceous at maturity.
On decaying wood: Tennessee (Burbank).
Spores 5-6X3-3·5 /L·
Basidia 16·5-25·5X4-5µ,, rarely with a hypobasidium: sterigmata 5·5-9µ, long.
Hyphae with clamps.
Doubtfully separable from M. guianense.
M. pendulum Mass.
Linder, I.e., f. 3-4.
Text-Fig. 201.
1-1·5 cm. long, pendulous, watery gelatinous, st1p1tate, lanceolate with an acute
apex, white, hyaline, or pale ochraceous at the base.
On rotten wood: Tasmania.
Spores (5-) 6-7X4·5-5µ,, white, smooth, subglobose or ovoid: (8-9µ,, Mass.).
Basidia (13·5-)25-28 X 5-'7µ,, 3·5-4µ, wide at the base: sterigmata 2-4, 2·5-4µ, long:
apparently without hypobasidia.
Hyphae apparently without clamps (? wider than in M. guianense).
PHYSALACRIA Pk. 1
Bull. Torr. Bot. Cl. 9, 1882, 2.
Synonyms: Baumanniella P. Henn., Engl. Jahrb. 23, 1897, 543.
Eoagaricus Krieger, Bull. Md. Ac. Sci. 3, 1923, 7.
Fruit-body simple, small, 1-25 mm. high, stalked-capitate, white or yellowish: head
1
Compare Pistillaria, Dimorphocystis.
PHYSALACRIA 457
globose, conical or variously folded, cerebriform or flattened, inflated, hollow: stem
cylindric, generally (if not always) with caulocystidia, solid: texture waxy-firm, the
wall of the head thin.
Lignicolous or epiphyllous: temperate and, mainly, tropical, 20 spp.
Spores white, smooth (rarely rough, P. sasae); ellipsoid-fusiform to subglobose,
? aguttate or merely with a few guttulae.
Basidia small, narrowly clavate, the young basidia narrow and subacerose: sterig
mata 2-4, nearly straight.
Cystidia characteristically oleocystidia, clavate, or ventricose-rostrate, subhymenial or
shortly projecting, and capped with a yellowish or brownish amorphous, resinous exudation
enclosed in a hyaline membrane (the resinous matter wholly soluble in dilute KOH),
contents granular and rather dense: sometimes also with fusiform hymenial cystidia,
capped with an exudation or not, in sterile hymenia.
Hymenium covering the whole head, absent from the stem, either wholly fertile or
sterile over the proximal part of the head or over the physical upperside of the head
or in irregular patches, not thickening: subhymenium composed of narrow uninflated
hyphae.
Caulocystidia generally (? always) present, scattered or very abundant, in some
species as oleocystidia.
Hyphae monomitic, clamped, inflating, not secondarily septate, those of the stem
elongating considerably, those of the head elongating slightly and soon pulled apart
to form the thin wall surrounding the central air-cavity: the walls of the hyphae slightly
but firmly submucilaginous in the head and forming a wall of subagglutinated tissue
without air-spaces: in the stem similarly modified or not.
Development gymnocarpic but indirect.
Type-species: P. infiata.
The affinity between this genus and Hormomitaria is explained on page 97.
Physalacria is the last term of reduction of a series of Clavarioid fungi, related, perhap,,
with Marasmius, Skepperia, and so on, and it stands to Hormomitaria as Dimorpho
cystis to Pterula, or as Pistillaria to Typhula and Clavariadelphus. The apex of the
small, indirect, primordial shaft is transformed into a convex hymenium which
enlarges by intercalary growth into a shortly conical, ellipsoid or globose, often plicate
head with a large air-space surrounded by the very thin, submucilaginous wall of
tissue supporting the hymenium: the air-space is caused by the disruption of the
internal tissue at the apex of the primordial shaft, or stem, which thus appears to end
abruptly in the head. Accordingly, there are no massive fruit-bodies in the genus.
A microscopic distinction, common with Hormomitaria, is the oleocystidia. They
are developed mostly from the primary hyphal ends of the incipient hymenium on
cessation of upgrowth of the primordial shaft. Typically they are clavate or ventricose
with a short blunt apical process: they have rather dense, finely granular contents, and
as they become slightly thick-walled distally, they exude a colourless, yellowish, or
brownish resinous-vitreous matter between the outer hyaline membrane of the wall
and the thickened middle layer. The exudate is very soluble in dilute alkali and such
swelling reagents as used in studying dried fungi, and is thus easily overlooked unless
fresh material or that preserved in alcohol-formalin is examined. On solution, the
distended outer pellich! of the apex of the cystidium can be seen surrounding the
space occupied by the exudate.
The development of the fruit-body is described under P. tropica. That of P. infiata,
P. bambusae, P. corticola, and P. Decaryi is essentially similar. In P. andina, apical
growth appears to be shortly prolonged after the acropetal inception of the hymenium,
and intercalary extension is comparatively slight, so that the head is conical and
resembles the rudimentary state in Hormomitaria.
Most species are still inadequately known. They have been described mainly from
dried material, so that many critical features may have been overlooked. Hence it is
PHYSALACRIA
impossible yet to define them accurately or to classify them naturally. I mention what
seem the more important structural differences.
Shape of the head. If apical growth is prolonged, the conical or ellipsoid head with
obconic base is formed, as in P. aggregata, P. andina, P. Langloisii, P. orinocensis, and
P. Sanctae-Martae. In contrast, abrupt cessation of growth gives the initially globous
head which becomes puckered or folded with a rounded or more or less umbilicate
base, as in P. tropica, P. infiata, P. Decaryi, P. bambusae, P. corticola, and P. clusiae.
Sterile hymenial areas. Typically they occur, as in Hormomitaria, around the base
of the head, e.g. P. tropica, P. bambusae, P. aggregata, P. Sanctae-lvlartae and, perhaps,
in P. andina. In P. infiata their disposition appears to be negatively geotropic instead of
structural. In P. Decaryi and P. corticola, however, the heads are wholly fertile and
there are no fusiform cystidia.
Oleocystidia. In the species which I have examined, namely P. tropica, P. Decaryi,
P. bambusae, and P. corticola, the oleocystidia have characteristic shapes, though little
if any specific differences in size. They are largest in P. tropica and P. Decaryi. Their
study is difficult, however, in dried material, but they are certainly absent from
P. stilboidea.
Caulocystidia. P. tropica, probably also P. infiata, have subcylindric caulocystidia
developing pale brownish walls exactly as in many species of Marasmius. In P.
Decaryi, P. bambusae, P. corticola, .P. australiensis, and, probably, most species with
slender stems, the caulocystidia are oleocystidia, as in the hymenium, though often
much reduced. This difference seems to be fundamental.
Spores. P. sasae, if correctly described, is at once distinguished by its rough spores.
P. andina, P. clusiae, and P. stilboidea have long fusiform spores, as in P. sasae, but
they may be bisporous effects rather than indications of the elongate Marasmioid
spore. The bisporous P. australiensis and P. Sanctae-Martae, however, have the
smallest spores of the genus, and it is remarkable that they should be dispores, for
such are usually larger than tetraspores.
Colour. Most species are colourless. The browning of the lower part of the stem
in P. tropica is caused by the coloration of the slightly thickened walls of the caulo
cystidia as they mature. The yellowish tinge in the head of several species is probably
caused by the colour of the resinous-oily exudate of the oleocystidia. Whether any
have cytoplasmic pigmentation as in Hormomitaria is not known.
Conclusion. The two species, P. infiata and P. tropica, with largest fruit-bodies,
seem to form a well-defined subgenus with relatively stout stems, Marasmioid caulo
cystidia, and much-folded hymenium. The remaining species appear as degenerate
and juvenescent forms. Thus, in P. tropica, with prolonged outgrowth of the initial
hymenium, the oleocystidia become more or less deeply immersed: in P. bambusae,
P. corticola, and P. Decaryi with precocious hymenium, the oleocystidia project con
siderably. Similarly, the few and imperfect caulocystidia of P. bambusae and P.
corticola are an effect of shortening the time of development of the stem: in P. tropica,
with longer period of extension of the stem, the caulocystidia are perfected and
multiplied while it is growing. The transformation of the Marasmioid caulocystidia
into hymenial oleocystidia signifies the precocity of the hymenium.
Whether there are sessile derivatives of the genus remains to be discovered. The
oleocystidia should enable such to be recognized.
P. stilboidea is problematic: it may have lost the oleocystidia or, as seems less likely,
have never developed them.
KEY TO THE SPECIES OF PHYSALACRIA

Head> 2·5 mm. wide: stalks conspicuous
10-30 mm. high, densely caespitose: head 2-10 mm. wide, much
folded: sterile hymenium extensive: sp. 4-6 x 2-3 ·s µ
PHYSALACRIA 459
Head with 3-6 elliptic holes at the plicate, sterile base: oleo
cystidia 10-18 µ wide: trop. (Malaya) P. tropica
Head entire, the sterile parts variously placed: oleocystidia
6-10µ wide: temp. (N. Amer.) P. inj/.ata
Smaller; head 3-5 mm. wide: (sterile hymenium?)
Sp. 10-13X3-5 µ, asperulate:-4 mm. high:on bamboo:Ja pan P. sasae
Sp. smaller, smooth
Sp. 6X3 · 5µ: -8 mm. high: on wood: Brazil P. brasiliensis
3p. 3·5 X2µ: - 7 mm. high: on wood: Brazil P. rugosa
Sp.?: yellow-brown: on wood: Siam P. changensis
Sp. 3-4X2-3 µ: 2-spored: 4-10 mm. high: on wood: Vic
·:oria P. · australiensis
Head < 2·5 mm. wide: mostly very small, with filiform stem
Basidia 2-spored: trop. Amer.
Sp. 7-12X3-4·5 µ: -3·5 mm. high, head conical: (? Asia) P. andina
Sp. 3-4X1·5-2µ: -4 mm. high, irregularly plicate, ellipsoid . P. Sanctae
Martae
Basidia (2-)4-spored
Sp. 13-17µ long
Cystidia thick-walled, not resin-capped: sp. 4-5µ wide:
erumpent, on leaves of Panax: New Zealand P. stilboidea
Cystidia resin-capped:sp. 2·5-3µ wide:on leaves of Clusia:
Venezuela P. clusiae
Sp. 6-9µ long
Sp. 6-7 X3·5-4·5µ:-4 mm. high:on wood:trop. Amer. and
Asia P. orinocensis
Sp. 6·5-9X3·5-4µ: -1·5 mm. high: on bamboo: S.E. Asia P. bambusae
Sp. 6-8x3-3·5µ: -1 mm. high: epiphyllous: Venezuela . P. concinna
Sp. 6µ long or less
2-6 mm. high: head 1-2 mm., without sterile base: sp.
3·5-4·5 X2·5-3µ: on dead wood: Madagascar, Malaya . P. Decaryi
1·5-4 mm. high: head 0·5-1 mm. wide, with sterile base:
sp. 4·5-6 X3-3·5µ: Panama P. aggregata
0·5-2 mm. high:head 0·25-1 mm. wide:sp. 4-5X t·5-2·5µ:
on wood, N. Amer. (Louisiana, Florida) P. Langloisii
0·5-1·2 mm. high: head 0·3-0·7 mm. wide, wholly fertile:
sp. 5·5-6·5 X2·5-3µ: on dead wood: Mal�ya P. corticola
2-3 mm. high: head 1-1·5 mm. wide: sp. 4-6µ, subglobose,
? fuscous: Togoland P. togoensis
Spores unknown: incertae sedis
1 mm. high: head 0·4 mm. wide: epiphyllous: Ceylon P. villosa
(2-4 mm. high: stem short, thick: on bark: Brazil Clavaria verpi-
Jormis)
P. aggregata Martin et Baker
Bull. Torr. Bot. Cl. 68, 1941, 280, f. 88-93.
1·5-4 mm. high, gregarious or crowded, white, d rying ochraceous: head 0·5-2·4X
0·4-1·1 mm., subglobose, rather irregular or somewhat lobed, hollow, sometimes
PHYSALACRIA
wrinkled at the sterile base: stem 0·9-1·8 mm. high, slender, puberulous, stem apex
projecting shortly into the head.
Habitat (? dead wood), Panama (G. Vv. Martin 2260, 2261, 2321, 232m, 2586, in
herb. Univ. Iowa).
Spores (3·5-)4·5-6·0 X (2-)3-3·5µ,, ovoid-ellipsoid with a distinct apiculus.
Basidia (10-)12-14 X 2·5-4µ,: sterigmata (2-)4, 3 µ, long.
Cystidia 50-54 X 9-12µ,, ventricose-rostrate, subhymenial, projecting 8-10 µ,, m
crusted.
Paraphyses 22 X 3 µ,.
Hymenium 10-15µ, thick, sterile at the base of the head and in irregular patches.
P. andina Pat.
Ess. Tax. 1900, 50: Syd., Ann. Myc. 28, 1930, 33: Baker, Bull. Torr. Bot. Cl. 68,
1941, 284, f. 69-75.
Synonyms: P. orinocensis Pat. et Gail!. var. andina Pat., Bull. Soc. Myc. Fr. 9,
1893, 136.
P. tenera Syd., Ann. Myc. 28, 1930, 33 (teste Baker, I.e., 286).
Text-Fig. 202.
0·75-3·5 mm. high, solitary to sparsely gregarious, �umetimes 2-3 joined at the
base, white, drying yellowish: head 0·5-2·5 X
0·3-2 mm., more or less conical, ovoid or obtuse
to subacuminate, rounded at the base, hollow
in the proximal part, sometimes subundulate:
stem 0·3-1·4 mm. ·75-100µ,: hymenium
25-30 µ, thick, subhymenium 15-20µ,,
apparently sterile rnuml the base of the head.
On dead wood, leaves, bamboo: Ecuador,
Columbia, Venezuela (amphigenous on dead
hanging leaves of Podocarpus cMiaceus, Sydow
360, ut P. tenera) (Philippine ls!., on bamboo,
PsiJium, Alangium, but ? identification).
Spores 7-10 ·.. · 3·5-4·5 µ, (Baker), i2-r5
3-4µ, (Syd.), 6-u(-12);. 2·5-3·5(-4)µ, (Syd.,
for P. lenera), oblcmg ellipsoid or fusiform.
13asidia 20-24 · +·-Sfl (Baker), 18-24 ·,;7-81•,
(Syd.), 20-25 >: 5-6 µ, (Syd., for P. tenera),
2-spored (Baker), 4-sporcd (Syd.).
Cystidia --70 ><. 12µ, (Syd.), 30-40 · · 12-15µ,
(Baker), projecting -30 µ,, ventricose-rostrate,
with a thick dense oleaginous resinous cap,
abundant.
Hyphae 1·5-4µ, wide in the head, here and
there swollen to 8µ,, clamped, thin-walled: in
TEXT-FIG. 202. Ph-ysalacria a11dilla: the stem 3-6 wide, somewhat thick-walled,
fruit-bodies, X 14: diagram of the µ,
structure of the incompletely hollow the superficial hyphae closely set with short,
head with sterile base: (from Baker, obtuse processes 1-2µ, high.
19+1). According to Baker, the basidia are typically
2-spored. A mean dispore, 8·5 X 4/L (Baker),
has a volume of c. 70 c.µ,, and an equivalent tetraspore _of half the volume
would be 6·5 X 3·25µ, (36 c.µ,): thus, the species appears like a 2-spored state of
P. orinocensis. On the other hand, it appears to have a peculiar structure more like
that of Hormomitaria than any other of the genus. The conical shape of the head is
caused, apparently, by the usual Physalacria-inflation occurring only in the proximal
part, just above the stalk, while the distal part retains the compact structure of the
PHYSALACRIA
apex of a primordial shaft. Thus, the fruit-body of P. andina may be likened to the
juvenile state of Hormomitaria (cf. Text-Fig. 178, third stage) which has matured as
a Physalacria: and, from the diagram given by Baker, there appears to be a sterile base
to the head as in Hormomitaria (Text-Fig. 202).
Baker also describes the dispores as binucleate and suggests that two of the normal
four nuclei of the basidium pass into each spore.
P. australiensis sp. nov.
Synonym: Physalacria infiata (Schw.) Pk. sensu Fawcett, Proc. Roy. Soc. Viet. 52,
1940, 157, t. 7, f. 2-3, text f. IA, B, c.
4-ro(-13) mm. high, singly or two to three together, gregarious, rarely caespitose,
white then yellowish or brownish with age: stem 3-6x1-1·5 mm., puberulous: head
3-5 mm. wide, irregularly globose, hollow, collapsing, sometimes plicate at the base,
minutely puberulous.
On dead wood: Victoria.
Spores 3-4 X 2-3 µ, white, smooth, pip-shaped, aguttate.
Basidia 2-spored.
Cystidia subventricose, thin-walled, evidently with a small resin-cap, projecting:
caulocystidia similar.
This species differs from P. infiata in its smaller size, smaller spores (on 2-spored
basidia), and its caulocystidia. P. infiata has a loose palisade of ill-defined, subcylindric
to subclavate, caulocystidia as in P. tropica, and these two species seem to be distin
guished by this feature. P. australiensis, as described and illustrated by Miss Fawcett,
clearly belongs to the same group as P. Decaryi and P. corticola, but it has larger
fruit-bodies. It may be considered only a variety of P. Decaryi, but at present it is
advisable to keep them apart.
P. bambusae v. Hoehn. emend.

Sitzb. K. Ak. Wiss. Wien II8, 1909, 290.
Text-Fig. 203.
-r ·5 mm. high, gregarious, generally subcaespitose, wholly cream-white or the
extreme base of the stem brownish: head -1 mm. wide and long, globose or more or less
bluntly conical, hollow, very thin, the wall 50-60µ thick, generally dilated at the base
or subumbilicate, not lacunose: stem roo-4oox60-80µ, roo-150µ wide at the base,
solid, very slender, inconspicuous.
On dead rotting culms of bamboo: Java, Malaya (Singapore, on Bambusa vulgaris,
Jan. 1944, Oct. 1945).
Spores 6·5-9 X 3·5-4µ, white, smooth, oblong, blunt, opalescent, aguttate.
Basidia 18-23 -:<.5-6 µ, the immature basidia acerose: sterigmata 4, 3 µ long.
Cystidia 30-45(-55) X 8-14µ, flask-shaped with a short neck -9 X 3-5 µ, becoming
slightly thick walled, at first wholly finely granular-vacuolate, then widely vacuolate,
the neck capped with a yellowish or colourless resinous globule 9-15µ wide: abundant,
projecting -r6 µ.
Hymenium c. 20µ thick, not thickening: sterile at the base of the head and composed
there of many cystidia (with and without exudation) and of many, somewhat enlarged,
vacuolate sterile basidia grading into the cystidia.
Caulocystidia as the hymenial cystidia, but smaller and ·.vith all abortions to slight
processes, very scattered.
Hyphae thin-walled, clamped, colourless: in the stem 3-6 µ wide at the surface and
subagglutinated, the inner hyphae 3-roµ wide with cells 30-80(-roo)µ long, not
secondarily septate, the basal hyphae often with pale brownish walls: in the head,
2-5 µ wide, the inner ones stretched apart.
Macroscopically this is indistinguishable, unless by habitat, from P. corticola, but
it has a sterile base to the head and larger spores. The fruit-bodies grow on the
PHYSALACRIA
underside of prostrate bamboo-stems and the sterile base of the head thus faces
upward, as in Hormomitaria.
The description which I give is based on living Malayan specimens. It differs from
von Hoehnel's in the spores and the cystidia, which von Hoehne) says are absent. Yet
in all other respects the Malayan and Javanese fungi agree so closely that I cannot
believe that this common little Physalacria on dead bamboos in Malaya can be other
than that which grows in Java. This is von Hoehnel's description:
-0·8 mm. high, gregarious, white, d rying yellowish: head -330µ wide, globose or
ovoid, vesicular, hollow: stem 360-450 X 40-60µ, cylindric, stuffed: spores 3-4µ
wide, white, globose, punctate-asperulate: basidia 12-14 X 3-4 p,: sterigmata 4:
cystidia nor..e: hyphae of stem 3-5 p, wide: on dead culms of bamboo, Java (Buiten
zorg).
TExT-FIG. 203. Phy•·alacriabambusae: fruit-bodies in section (the hollow in black), x 5: spores,

sterile hymenium (left) and fertile hymenium (right), X 1,000. (Malayan specimens.)
It should be noted that, if dried material is examined in dilute swelling and clearing
reagents, the very thin spore-coat may become so transparent that it is overlooked and
the shrunken spore-contents may then be mistaken for the spore: the reagent will also
dissolve the resinous exudation of the cystidia which renders them so conspicuous in
fresh material. This may explain the discrepancies in von Hoehnel's description.
His measurements for the basidia refer to immature cells.
P. brasiliensis (Rick) comb. nov.

Basinym: Baumanniella brasiliensis Rick, Brot. 5, 1906, 11.
-8 mm. high, gregarious, pale stramineous white, drying reddish brown: head 3-5 mm.
wide, globose, often depressed, hollow: stem -3 X 0·5 mm., at first white pruinose, then
dingy.
On wounds of trees of Anchietea salutaris (Violaceae): Brazil (Rio Grande do Sul).
Spores 6 X 3 ·5 µ, white, smooth, ovoid, apiculate.
Basidia monosporous.
PHYSALACRIA
Cystidia flask-shaped.
This appears to be related to P. infiata and P. tropica. The 'monosporous' basidia
are, perhaps, ventricose-rostrate cystidia.
P. changensis Rostr.
Bot. Tijdsskr. Copenh. 24, 1902, 206.
Yellow-brown, caespitose: head 3-4 mm. wide, globose, hollow, membranous,
perforate at the base: stem I mm. long, slender.
On trunks, Siam.
Spores ellipsoid, minute.
Cystidia 32 X 12µ, clavate.
The colour, exceptional in the genus, may refer to dried specimens, in which case
they may be only small specimens of P. tropica mihi.
P. clusiae Syd.
Ann. Myc. 28, 1930, 35.
0·5-1 mm. high, gregarious in patches 1-4 cm. wide, laxly or densely arranged,
amphigenous or, mostly, hypophyllous and breaking through to the upperside,
ochraceous yellow: head 400-800µ wide, globose or slightly depressed: stem 150-220 X
80-140µ, short, slightly widened at the base.
On dead, fallen leaves of Clusia rosea (Guttiferae): Venezuela (Puerto La Cruz,
Syd., n. 105).
Spores 13-17 X 2·5-3µ, white, smooth, more or less fusiform, flattened adaxially,
subacute at each end, with numerous small and large guttulae.
Basidia 25-30 X 7-8µ, clavate: sterigmata 2-4, -10µ long.
Cystidia 20-50 X 10-12µ, more or less prominent, clavate .or cylindric-clavate, with
a short, thick-walled stalk, capped with a yellowish oleaginous resinous exudation.
Hyphae of stem 2-4µ wide, irregular, rigid.
Superficially this must resemble Dimorphocystis unless it has the delicate, semi
translucent head of Physalacria.
P. concinna Syd.
Ann. Myc. 28, 1930, 36.
0·5-0·8 mm. high, sparsely or laxly gregarious, often solitary, mostly hypophyllous,
whitish or yellowish white: head 350-600µ wide, globose or subglobose, somewhat
collapsed in the centre on drying, very finely mealy: stem 140-180 x 200µ, very short
and thick.
On dead, hanging leaves of Podocarpus coriaceus (Coniferae): Venezuela (between
El Limon and Colonia Tovar, Syd. n. 361).
Spores 6-8 X 3-3·5 µ, white, smooth, obliquely ovoid, inequilateral, rounded-obtuse
at the apex, obliquely apiculate at the base.
Basidia 18-22 X 5-6·5µ, clavate: sterigmata 2-4, 3-5µ long.
Cystidia -70 X 6-8(-10)µ, scattered, clavate or cylindric-clavate, mostly 5-6µ wide
at the capitulate apex covered by the reddish brown oleaginous-resinous exudatiqn
(12-15µ wide), projecting for about half their length.
Cystidioles -28 X 5-6µ, acute, very abundant among the basidia.
Hyphae 1-3µ wide.
This may be only a form of P. andina with unusually thick stem. The variability in
stature of these small Basidiomycetes is little realized, cf. Pistillaria capitata and
P. setipes. The form suggests Dimorphocystis, but the cystidia are those of Physalacria.
The cystidioles may be immature basidia or paraphyses as described for P. infiata.
P. corticola spec. nov.
Text-Fig. 204.
0·5-1·2 mm. high, solitary, or gregarious, not caespitose, pendent, superficial, white:
PHYSALACR IA
head 0·3-0·7 mm. wide, the wall 40-50µ thick, subglobose, often flattened or slightly
plicate, hollow, slightly umbilicate at the base: stem 200-500 X80-150µ, cylindric or
slightly thickened upward, solid, very sparsely and minutely puberulous.
On dead bark and wood: Malaya (Singapore, Botanic Gardens, 22.3.42, 11.11.43).
Spores 5·5-6·5 X2·5-3µ, white, smooth, narrowly
ellipsoid, obtuse or subacute, thin-walled, aguttate.
Basidia 12-18 X 4-5µ, narrowly clavate, the young
basidia subacerose-acute: sterigmata 2-4, 3 fL long.
Cystidia 35-50 X8-17µ, some only 20-30 X5-'7 µ,
projecting -20µ, arising deeply from the sub
hymenium, clavate and very obtuse, mostly with a short,
blunt, subconical tip, thin or slightly thick•walled,
sm;>oth or becoming capped with amorphous, resinous
matter (wholly soluble in dilute KOH), with dense
granular cytoplasm (the young cystidia not incrusted).
Hymenium c. 20µ thick, not thickening: sub
hymenium c. 15 fL thick, cornpos.ed of narrow hyphae
1·5-2·5µ wide.
Caulocystidia -30 X3-7µ, as scattered, colourless,
clavate, or subventricose thin-walled processes like
blunt, sterile basidia or as small cystidia with a small
resinous-oleaginous apical exudation.
Hyphae of the head 3-5 µ wide, thin-walled,
clamped: of the stem 5-12µ wide, the cells 40-100µ
long, not secondarily septate, clamped, thin-walled,
those at the surface 2-5 fL wide and slightly agglutinated
into a pellicle 2-'7 µ thick.
The fruit-bodies hang from the vertical and lower
sides of fallen trunks and branches and are clearly
ageotropic. Superficially they resemble P. aggregata
and P. bambusae, but they have no sterile base to the
head. Possibly they are only small specimens of P.
orinocensis.
TEXT-FIG. 204. Physalacria
corticola: fruit-bodies, X 5: P. Decaryi Pat.
section of the fruit-body (the
hollow in black), X 10: spores, Mero. Acad. Malgache 6, 1927, 10.
X 1 ,ooo: hymenium, oleocystidia, Text-Fig. 205.
and caulocystidia (right), X 500. 2·5-6 mm. high, scattered or gregarious, not
caespitose, white: head 1-2 mm. wide, subglobose or
bluntly conical with broad base, often three-angled, sometimes somewhat flattened on
the top, inflated, hollow, without lacunae, wholly fertile: stem 1·5-4 mm. X100-250µ,
cylindric, minutely puberulous.
On dead wood in the forest: Madagascar, Malaya (Pahang, Tembeling, 6.11.30).
Spores 3·5-4·5 X 2·5-3µ, white, smooth, ellipsoid pip-shaped.
Basidia 18-22 X3·5-4·5µ, subclavate, the young basidia subacerose: sterigmata 2-4,
2·5-3µ long.
Cystidia 35-5s :< 9-22 /J·, projecting for l-;\ their length, clavate with a short, blunt
cylindric or slightly tapered appendage, thin or slightly thick-walled, capped with a
resinous-oleaginous amorphous mass (whoJly soluble in dilute KOH), contents densely
granular, arising deeply ·frum the subhymenium.
Hymenium c. 20µ thick, not thickening, fertile over the whole surface of the head.
Caulocystidia 25-50 x 8-18 µ, scattered but abundant, not forming a palisade,
ventricose or conoid-ventricose with a short, blunt apical process capped with resinous
matter like the hymenial cysti<lia, thin- or slightly thick-walle<l, contents vacuolate.
PHYSALACRIA
Hyphae 2-5µ. wide in the head, 1·5-2·5µ. wide in the subhymenium, clamped: in
the stem 5-12µ. wide, the cells 40-100µ. long, the outer hyphae 3-5µ. wide and
agglutinated slightly into a pellicle 2-7 µ. thick.
This description is drawn from the Malayan specimens. That of Patouillard's
agrees so closely that I have no hesitation in identifying the Malayan and Madagascan
fungi. The species comes near to P. aggregata, which seems to have smaller fruit
bodies with a distinct sterile base to the head. P. corticola differs in the very small
fruit-bodies, longer spores, and less-developed caulocystidia. P. australiensis is similar,
but larger.
J .
.
<
0 V
A �
A LY
0, ()·...'
TE.xT-FIG. 205. Physalacria Decaryi: fruit-bodies, X 5: spores,

.< 1,000: hymenium, oleocystidia, and caulocystidia (right), X 500.
(Malayan specimens.)
Patouillard's description is:

3-4 mm. high, in small tufts, white: head 1 mm. high, cylindric, truncate at the
top, hollow: stem 150-200 fL thick, slender, puberulous: on dead wood, Madagascar:
spores 3-4 µ., ovoid, hyaline: cystidia 25 X 9-10 µ., hyaline, with an apical beak, project
ing: caulocystidia 45 X 12-15 µ., also beaked.
P. inflata (Schw.) Pk.

Bull. Torr. Bot. Cl. 9, 1882, 2, t. 9, f. 1-5: Lloyd, Myc. Notes 1, 1908, 4, f. 216-18:
ibid. 682, f. 1020: Krieger, Bull. Md. Ac. Sci. 3, 1923, 7: Rogers, Iowa Stud. Nat.
Hist. 15, 1933, 20: McGuire, Mycol. 31, 1939, 433: Baker, Bull. Torr. Bot. Cl. 68,
1941, 279, f. 1-68: (non Fawcett = P. australiensis).
Basinym: lvlitrula infiata Schw. ex Fr., El. 1, 1828, 234.
5119 H h
PHYSALACRIA
Synonyms: Spathularia infiata (Schw.) Cke., Mycogr. 1879, t. 344.
Eoagaricus infiatus (Schw.) Krieger, Le.
10-27 mm. high, densely gregarious or caespitose, white then yellowish: head
1-8 mm. wide, subglobose or flattened, often irregularly undulate and plicate, mem
branous, smooth; hollow: stem 4-20 X 0·5 mm., slender, firm, pruinoso-puberulous.
On rotten wood and dead leaves: Canada (Ontario), U.S.A. (N.Y., Md., Pa., Wis.,
Neb., Iowa).
Spores 4-6 X 2-3 ·5µ,, white, smooth, ellipsoid pip-shaped, apiculate.
Basidia 18-25 X 4-6µ, (15-18 X 5·5µ,, Baker): sterigmata (2-)4.
Cystidia dimorphic: ventricose-rostrate, arising deeply in the subhymenium, project
ing 5-15µ, (-30µ,, Baker) or wholly immersed, heavily resin-incrusted, thin-walled,
40-50 X 8-toµ, (Rick), 26-52 x 4-10µ, (Baker), scattered in the fertile hymenium:
fusiform, acute or subacute, rarely branched, vacuolate, hymenial, projecting 20-50µ,,
smooth or thinly incrusted, 24-50 X 4-9µ, (Baker), mostly in the sterile areas of the
hymenium.
Paraphyses 27-31 X 2-3 µ,, subfusiform cylindric (Baker).
Hyphae narrow, clamped.
Hymenium with sterile areas of fusiform cystidia, mainly(? always) on the uppersides
of the folds: 20µ, thick in the fertile areas with the subhymenium 10-15µ, thick (the
underlying tissue 50-150µ, thick): 30-40µ, thick in the sterile areas, without sub
hymenium.
Structure and Development of the Fruit-body. According to McGuire, the growth of
the fruit-body may take any time from I to 2 weeks, though why there should be such
variation and how long are the periods of development and maturity are not stated.
Development is essentially as in P. tropica (Text-Fig. 210) and the stem becomes
clothed with similar caulocystidia (not yet described in detail). The fruit-body grows
in any direction but the head, according to McGuire, is geotropic to the extent that
the upper part is sterile, and the remainder, with a downward component, is fertile.
Thus, if the primordial shaft has grown vertically upward, the distal half of the head
is sterile and the proximal half, round the stem, is fertile: if the primordial shaft has
grown vertically downward, the distal half of the head is fertile and the proximal is
sterile: Baker, however, finds an irregular distribution of sterile areas over the folds
of the head. The point has yet to be settled by careful study of the fruit-bodies in
nature: the sterile areas can be distinguished macroscopically by being more evenly
rounded and less folded. The stem is central, not lateral as described by Krieger, and
the folds of the hymenium are formed by uneven intercalary growth, not as rudi
mentary gills with geotropic downgrowth or outgrowth, as suggested in his false idea
of Eoagaricus. The cells of the hyphae are binucleate.
Hymenial elements. Baker distinguishes four elements beside the basidia, namely
ventricose-rostrate, incrusted cystidia of subhymenial origin, fusiform hymenial
cystidia of two sizes, and paraphyses. I think, however, that it has not been satisfac
torily shown that the narrower or smaller fusiform cystidia and the paraphyses are
more than transitions from typical fusiform cystidia to sterile basidia(as in P. tropica
and Hormomitaria). The sterile parts of the hymenium consist of fusiform cystidia
with many transitional forms and few incrusted cystidia, whereas the fertile parts
consist of basidia, abundant incrusted cystidia, and few fusiform cystidia with transi
tional forms as 'paraphyses'. The cystidia of all kinds and the paraphyses, according
to Baker, are binucleate at first, then uninucleate from nuclear fusion, and finally
binucleate again, but these two nuclei never divide into the 4-nucleate stage as in the
basidium. The spores are uninucleate: they may be formed before the nuclei pass
into them.
Distribution. The species has been recorded from Brazil and the Philippine Isl.,
but the specimens from these countries may be P. tropica. Thus P. infiata seems to
be a temperate N. American species.
PHYSALACRIA
P. Langloisii E. et E.
Joum. Myc. 4, 1888, 73: Baker, Bull. Torr. Bot. Cl. 68, 1941, 284, f. 83-7.
0·5-2 mm. high, white or yellowish: head 0·25-1 mm. wide, subglobose, somewhat
elongate or depressed, solid then hollow: stem 0·3-1 mm.X50-70µ, relatively long,
puberulous.
On rotten wood: U.S.A. (La., Fla).
Spores 4-5X2·5µ (4·2-5X1·5-2µ, Baker).
Basidia u-13X2·5-3µ, 4-spored.
Cystidia 30-35X15µ (22-28x5-7·5µ, projecting 7-13µ, Baker), clavate to
ventricose-rostrate.
According to Baker the hymenium is not wholly fertile, but whether there is a
sterile base, as in the very similar 2-spored P. Sanctae-Martae, is uncertain. From
P. orinocensis it seems to differ only in the slightly smaller spores.
P. orinocensis Pat. et Gaill.
Bull. Soc. Myc. Fr. 1888, 41, t. 13, f. 2: Baker, Bull. Torr. Bot. Cl. 68, 1941, 279,
f. 77-82.
3-4 mm. high, whitish: head 0·58-1·6X0·55-2 mm., oblong-ellipsoid, subplicate,
blunt, base not or scarcely depressed: stem 0·5-0·75(-3·5) mm., rather short.
On rotten wood: Venezuela, lndo-China, Philippine Isl.
Spores 6'-7·5X3·5-4·5µ, ovoid-ellipsoid.
Basidia 18-22x4-5µ: sterigmata 2-4, 3-3·5µ long.
Cystidia abundant, ventricose-rostrate, often incrusted, with small fusiform, not
incrusted, cystidia 12-13X3µ, not projecting.
Hymenium 15-24µ thick, with sterile areas: subhymenium c. 10µ thick.
This appears to be a pantropic species, yet without more microscopic details I
cannot identify it with any of the Malayan species. P. bambusae and P. corticola seem
to have smaller fruit-bodies, though P. bambusae has almost identical spores. P. Lang
loisii seems to differ only in the slightly smaller spores.
P. rugosa Rick.
Brot. 5, 1906, 12.
-7 mm. high: head 5 mm. wide, hemispherical, straw-yellow, gyroso-rugulose, inflated,
hollow, waxy-soft, rather fleshy: stem 5X2 (? 0·2) mm., fuliginous, white pruinose,
running through the pileus.
On pieces of dead wood: Brazil (Rio Grande do Sul).
Spores 3·5X2 µ, white, smooth, pip-shaped.
Basidia 25µ long: sterigmata 3-6µ long.
According to Baker (Le. 287) two authentic collections of Rick'!; examined by her
did not belong to Physalacria. The description of the stem running through the
pileus is odd unless it indicates somewhat the same condition as .in P. andina. Com
pare Clavaria verpiformis.
P. Sanctae-Martae Martin et Baker
Bul). Torr. Bot. Cl. 68, 1941, 286, f. 94-9.
1-4 mm. high (rarely more), white, gregarious, pendent: head 1 ·4-2·5 X 0·8-1·5 mm.,
ellipsoid to subconic, hollow, conspicuously and irregularly plicate with short, shallow,
subgyrose folds, with a sterile basal zone: stem 0·8-1·6(-5) mm., puberulous.
Habitat: ?Columbia (G. W. Martin, 3485, 3654, 3704, in herb. Univ. Iowa).
Spores 3-4X 1·5-2 µ, very small, elliptic-ovoid to pyriform, apiculate.
Basidia with 2 slender sterigmata, 10-12 X1·5-3µ.
Cystidia 35-46 X8-ro µ, projecting -12µ, thick-walled distally.
Hymenium 10-15µ thick, sterile round the base and in patches on the sides: sub
hymenium 5µ thick, verf slight.
PHYSALACRIA
The short gyrose puckerings of the head and the very small dispores seem charac
teristic.
P. sasae Imai
-4 mm. high, solitary or gregarious, white, drying straw-colour: head -3 mm. wide ,
subglobose or ovoid, hollow: stem 1 X0·2 mm., cylindric, stuffed.
On dead culms of Sasa kurilensis (bamboo): Japan (Prov. Ishikari).
Spores 10-13 X3-5µ,, white, ellipsoid, asperulate.
Basidia cylindric, 4-spored.
Cystidia 60-70 X10-13µ,, fusiform, hyaline.
The asperulate spores are peculiar in the genus.
P. stilboidea (Cke.) Sacc.

Syll. Fung. 9, 1891, 256.
Basinym: Pistillina stilboidea Cke., Grev.
Text-Fig. 206.
-2·5 mm. high, erumpent, gregarious, minute, wholly pallid ochraceous (? white
when living), glabrous: head 0·2-1·5 mm.
wide, globose-depressed, hollow: stem -1 ·s
mm. xo·1-0·2 mm., filiform, pruinose, appear
ing cupular-volvate from the erumpent tissue of
D000
the leaf, inserted into the head.
On blackish spots on leaves of Panax: New
Zealand (Berggren 275, Colenso 423).
1
Spores 13-16 X 4-5µ,, white, smooth, elon
gate pip-shaped.
Basidia c. 30-40 X8-9 µ, (? mature), clavate
cylindric.
Cystidia 45-90 X9-18µ,, abundant ventri
coso-fusiform, subacute or blunt, projecting
-50µ, with a more or less prolonged apex, walls
1-3µ, thick, smooth, not incrusted, colourless,
the base deeply immersed: oleocystidia absent.
TEXT-FIG. 206. Ph:ysalacria stilboidea: Caulocystidia -55 X7-12µ,, as the cystidia
fruit-bodies erumpent from the black- but shorter, more acute, and often subtriangu
ened leaf-spots, X 5: spores, cystidia,Jar at the base, scattered on the stem, crowded
and caulocystidia (bottom right), X 500:
above and compacting into a narrow sterile
_
(type-collectwn, Herb. Kew.). hymenium at the base of the head.
Hyphae 2-4µ, wide on the surface of the stem and agglutinated: internal hyphae
2-5µ, wide, long-celled, the walls rather firm, o· 5µ, thick: apparently with many
granules or crystals in the tissue, often aggregated into subglobose lumps: clamped.
I have examined the collection Berggren 275, from Waitaki, in the Kew herbarium.
The species is peculiar in lacking oleocystidia but in having abundant ordinary thick
walled cystidia corresponding with the fusiform cystidia of P. tropica and, apparently,
forming in the same way a narrow sterile hymenium at the base of the head. The
species should easily be recognized from the habitat, the erumpent habit, and the
large spores.
P. togoensis (P. Henn.) comb. nov.
Basinym: Baumanniella togoensis P. Henn., Engl. Jahrb. 23, 18<)7, 543, t. r4, f. 7a-c.
Head 1·5-2 x 1-1·5 mm., pale yellow, subglobose or ovoid-clavate, hollow: stem
barely 1 mm. long, solid, hyaline, slender.
On dead wood: Togoland.
PHYSALACRIA
Spores 4-6 µ,, clear brown or fuscidulous, smooth, subglobose.
Cystidia 23-30 X14-16µ,, much projecting, flask-shaped, sparse.
According to von Hoehne!, the basidia are normal, not 1-spored as stated by
Hennings, and the species is a true Physalacria: apparently Hennings mistook the
oleocystidia for basidia with spores, hence the spore-data may not be correct.
P. tropica sp. nov.
Text-Figs. r4 (D), 207-10.
7-22 mm. high, caespitose, superficial, white, the base of the stem eventually pale
brown: head 4-10 mm. wide, the wall c. 100µ thick, subglobose, inflated, hollow,
becoming irregularly plicate, flattened, lobed, or somewhat cerebriform, generally puckered
round the more or less umbilicate base and often with 3-6 small oval or ellipsoid lacunae
radiating from the stem-apex: stem 3-r 2 mm. X200-600µ,, cylindric, solid, wholly
minutely {>ruinoso-puberulous.
TEXT-FIG. 207. Physalacria tropica: the hollow of the head shown in black, >-: +
On fallen trunks in the forest: Malaya (Pahang, Tembeling, 6. 1r .30: Negri Sem
bilan, Pasoh Forest Reserve, 25.6.30).
Spores 4 · 5-5·5 X2· 5-3µ,, white, smooth, ellipsoid pip-shaped, thin-walled, aguttate.
Basidia 18-25 X4·5-5·5 µ,, narrowly clavate, the young basidia subacerose, 2-3·5µ
wide: sterigmata 4, 3µ, long.
Cystidia of two kinds: oleocystidia 35-65 X I 0-18µ,, stout, blunt, or subtruncate, the
wall 0·5-1µ, thick, with a yellowish brown, irregular, often some,<-·hat cristate, resinous
vitreous cap (wholly soluble in dilute KOH), contents yellO'l.vish resinous-vitreous,
arising deeply in the subhymenium or trama, immersed (-20µ below the surface) or
reaching the surface or slightly projecting, abundant but scattered: fusiform cystidia
30-70 X7-16µ, ventricoso-fusiform, subacute, thin-walled, vacuolate-hyaline, not
incrusted, hymenial, projecting far, abundant near the stem-apex and at the base of
the head in the sterile hymenium, absent from the fertile hymenium.
Hymenium c. 25 µ, thick, covering the whole head, mostly fertile with basidia and
immersed clavate cystidia, but more or less extensively sterile over the puckered and
lacunose base of the head, the sterile hymenium consisting of fusiform cystidia with
sterile basidia and vny scattered immersed clavate cystidia, or consisting solely of
fusiform cystidia (at the stem-apex and along the edges of the lacunae): subhymenium
15-20µ, thick, composed of closely interwoven hyphae 2-2·5µ wide.
Caulocystidia -80 X 3-7 µ,, occasionally -13 µ, wide, cylindric, clavate, or ventricose,
variable in shape, often flcxuous, sometimes branched or lobed, blunt, forming a fairly
470 PHYSALACRIA
close, but not contiguous, palisade, mostly with slightly thickened, yellowish walls,
smooth, with rather dense cytoplasm: abundant in the lower part of the stem, fewer
upward, at the base of the stem elongating into short, divergent 2-3-septate hyphal
ends -120 X 2-3 µ.
TEXT-FIG. 208. Physalacria tropica: the fertile wall of the head with immersed and projecting
oleocystidia (left), and the sterile basal part of the head with lanceolate cystidia (right), X 500:
spores, X I ,ooo.
Hyphae monomitic, slightly inflating, clamped, not secondarily septate, with thinly,
but toughly, mucilaginous walls, forming a firm tissue without air-spaces (except the
central hollow of the head): hyphae of the stem 2-7 µ wide, the cells 45-300 µ long,
longitudinal, sometimes convolute, very compact, thin-walled, a few narrow hyphae
with dense resinous-oleaginous contents like laticiferous hyphae, the superficial
hyphae narrow and with slightly thickened, yellowish walls like the caulocystidia but
not agglutinated: hyphae of the head 4-7 µ wide, the cells short, 15-35 µ long, thin
walled, colourless, rather loosely branched and pulled apart: all the cells of the fruit
body having rather dense cytoplasm, those of the wall of the head being the most
vacuolate.
Development of the fruit-body. Development is superficial, gymnocarpic, and in
direct, apical growth ceasing in the primordial stage long before the fruit-body has
become full-sized. A minute, conical, primordial shaft, composed of narrow longi
tudinal hyphae and rather loosely corticated with incipient caulocystidia, grows
apically to a height of 0·5-1·2 mm., when apical growth ceases and the apex of the
shaft is converted into the primordial hymenium. About this time the cells of the stem
enter their grand period of extension, the process passing acropetally from the base
to the apex of the primordial shaft, and, thus, as the head develops it is projected
farther and farther from the substratum. The cells of the primordial shaft are 8-12 µ
long on delimitation, and their inflation begins slowly at the base of the shaft when it
is only 200-300 µ high. As the stem elongates, more cau!ocystidia are intercalated by out
growths from the superficial hyphae, but the process is restricted mainly to the proxi
mal part of the stem. The head is formed from the primitive hymenium over the
convexo-discoid apex of the primordial shaft by a short petiod of outgrowth of the
hyphal tips, cutting off 2-4 subhymenial cells, and then by sympodial branching and
slight inflation of the internal cells. The hymenial surface increases to a much greater
extent than the slight inflation of the internal cells permits, so the longitudinal hyphae
become pulled apart and a hollow appears in the primordial head when it is about
400 µ wide. The head enlarges and becomes folded th,;ough inequalities of hymenial
PHYSALACRIA 471
expansion and unequal tension or resilience of the internal hyphae which now form
the thin submucilaginous wall supporting the hymenium. The sterile hymenium
forms over the base of the head and the
fertile hymenium over the rest of its surface.
The first hymenial elements to mature are
the oleocystidia. They are modifications of
many of the initially excrescent hyphal ends
which compose the primordial hymenium
when the head is 200-400 µ, wide. The other
hyphal ends and most of their sympodial
branchings form 2-4 subterminal cells and
then their terminal cells become basidia.
Thus the clavate cystidia have a deep origin
in the tissue, which shows their early
formation, and they are more or less over
topped by the basidial hyphae. Sparing
seems to begin very late and not until the
fruit-body is almost or quite full grown.
Thus, most fruit-bodies appear sterile, the
hymenium consisting of closely packed,
subacerose young basidia.
The primordial shafts grow from the
wood in any direction, though mainly per
pendicularly. Thus the fruit-bodies project
in all directions and there is no evident
geotropic or phototropic curvature of the
stem. In P. inflata it is said that the fertile
hymenium develops only on the physical
underside of the heads, which are similarly
projected in any direction, but in P. tropica
I could find no such geotropic relation:
its sterile hymenium is always at the mor
phological base of the head (as in Hormo
mitaria). However, I have not been able to
study this feature in living specimens, and
it certainly needs more thorough investiga
tion.
Systematic position. At first I thought
that the common large Malayan species
must be P. inflata, but the extended descrip
tions recently given of this North American
species by McGuire and Baker reveal con
sistent differences. Thus the perforations
between the thickened sterile ridges at the TEXT-FIG. 209. Physalacria tropica: surface
base of the head in P. tropica are not of the stem with Marasmius-like
caulocystidia, X 500.
mentioned for P. in,+I
J ,ata, nor the characteris-
tic, Marasmius-like browning of the stem. The cystidia of P. tropica are larger and
somewhat differently shaped and, perhaps, the distribution of the sterile hymenium
is different.
P. villosa Petch
Ann. R. Bot. Gard. Per. 6, 1917, 206.
1 mm. high, white: head 0·4 mm. wide, globose: stem 0·7 mm. X 100µ, at the base,
40 µ, at the apex, twisted at the apex, fibrous, puberulous.
472 PHYSALACRIA
On dead leaves: Ceylon (Hakgala).
Spores?
Basidia 4-spored.
( Cystidia?)
Caulocystidia -30 X 8 µ,, conic or ventricose, rigid, thick-walled, attenuate above the
middle, the apex generally truncate or subcapitate.
TExT-FIG. 21 o. Physalacria tropica: diagrams of the development of the fruit

body: A, the young primordial shaft corticated with caulocystidia; B, inception
of the head on cessation of upgrowth, with the primary oleocystidia: c,
expansion of the head: D, the full-grown head with air-space and sterile
region round the base, the primary oleocystidia immersed in the wall of the
head.
It is impossible to decide whether this is Dimorphocystis, Pistillaria, or Physalacria,
though probably it belongs to the first genus. Such descriptions, without adequate
structural and hyphal details, are useless and confusing.
PISTILLARIA Fr.
Syst. Myc. 1, 1821, 496: Killerm., Zeitschr. f. Pilzk. 18, 1934, 98-108, 137-9.
Synonyms: Typhula subgen. Leptorrhizae Fr., Hym. Eur. 1874 (pr. p. maj.).
Cnazonaria Cda., Sturm. Deutsch!. FI. Pilze 7, 1829, SS: Donk, Rev. Niederl.
Homobas. Aphyll. 2, 1933, 96.
PISTILLARIA 473
Scleromitra Cda., ibid. 1829, 59.
Sphaerula Pat., Tab. An. 1883, 27.
Gliocoryne Maire, Bull. Soc. Bot. Fr. 55, 1908, 120 ( = Gloiocoryne auctt.).
Fruit-body generally small or slender, simple, or occasionally branched, with a filiform
sterile stem and a subglobose, clavate, or cylindric head, rarely elongate filiform, without
sclerotium, the head with a sterile apex at first, sooner or later fertile over the apex:
texture waxy-soft to rather horny, toughly mucilaginous or cartilaginous, especially
in the stem.
On dead plant-remains (a few possibly exceptional species terrestrial): north
temperate, few tropical or south temperate, c. 53 spp.
Spores white, smooth, ellipsoid to subglobose, thin-walled, aguttate, or with a few
minute guttulae.
Basidia with 2-4 sterigmata, rarely 1 (P. maculaecola ?)
Cystidia absent, except as cystidioles in P. subuncialis.
Hymenium not thickening; as in Typhula.
Caulocystidia generally present.
Hyphae monomitic, inflating, septate with clamps, or the clamps more or less absent
from the fruit-body, not secondarily septate, hyphae walls thin or slightly thickened and
toughly submucilaginous, agglutinated on the surface of the stem: generally with crystals
on or among the hyphae.
Lecto-type: P. pusilla or P. micans (see p. 99).
Most species of this genus are imperfectly known. Small, filiform, subsessile whitish
species must be sought also in Ceratellopsis. -
KEY TO THE SPECIES OF PISTILLARIA

Sp. brown, 4-5 µ, subglobose: on leaves of Solanum: Nyasaland P. phaeosperma
Sp. white
Head pink, red, orange, orange-yellow, or purple, or the stem
pink A
Head brown, fuscous-rufous, bistre, grey, or blackish B
Head white, yellowish, or ochraceous (stem brownish or con-
�m� C
(Compare Ceratellopsis)
A
On the ground
r 5-30 mm. high, or more: stem white: Europe P. limicola
5-6xo·5-1 mm., pinkish cinnamon then orange-pink: sp.
6 x3 µ: Montevideo P. montevidensis
On dead plant-remains
7-20 mm. high, rose-pink: head 2·5-4 mm. long: sp. 6-9 X 3-4µ.:
on Triticum repens: Europe P. elegantula
(-12 mm. high, orange to red and simple, or greyish yellow and
branched: sp. 7-r IX 3-5 µ: trop. Lachnocladium
aurantiacum)
0·3-5 mm. high: stem short
Scarlet, stem yellow: sp. 6-8 x 5-6 µ: on leaves of Syringa:
Europe P. syringae
Olivaceous, then whitish, stem pink: sp. 8 X 3-3 ·5 µ: on Helle-
borus : Corsica P. lividula
474 PISTILLARIA
Purple to purplish black, or blackish pink
Sp. 7-11 X 6-8µ,: 2-spored: on stems of Vitis: France P. ampelina
Sp. 4 X 3 µ,: stem whitish: on leaves: England . P. purpurea
Sp. 2-3µ globose: blackish pink: on Callirhoe: U.S.A. P. Bartholomaei
Sp. globose: concolorous purplish: on herbaceous stems:
France P. uliginosa
Rose-pink, either wholly or on the stem or head
Trop.: sp. 7-9 X 4·5-5µ,: 4-spored: on pods of Cassia alata P. rhodocionides
Temp.
Sp. 8-13 X 5-7 µ: mostly 2-spored: Europe, N. Amer. P. micans
Sp. 6 X 3 µ,: 4-spored: on Populus-leaves: France . P. granulata
Sp.?: stem pink, head white: on stems of Vitis: Italy P. bellunensis
B
Europe
Head cinnamon-yellow: stem bistre to blackish, long: on twigs P. Juscipes
Bistre then blackish, 4-6 mm. high: on wood P. tenuis
Head tawny or tawny-orange, 1-2 X o·1-0·25 mm.: stem whitish
or concolorous, -5 X 0·04-6·2 mm.: sp. 8-10 X 4-5µ,: on her-
baceous stems P. fulgida
Head honey-yellow, 5-6 X 0·3-0·8 mm.: on the ground P. gilva
America
Brown-ochraceous, then reddish bay: -4 mm. high, sessile, erum-
pent: sp. 10-15 X 6-7·5µ,: on branches of Alnus: U.S.A. P. alnicola
Blackish pink, 1 mm. high: sp. 2-3µ,, globose: on stems: U.S.A. P. Bartholomaei
Head greyish, stem black, o· 5 mm. high: on twigs: Cuba P. cupressiformis
Africa
Fuscous-rufus, -5 mm. high: sp. 5-6x3 µ,: on fallen leaves:
Cameroons P. bipindiensis
C
On the ground: doubtful: Europe
Honey-yellow: head 5-6 mm. long P. gilva
Head white: stem with moniliform, glandular hairs P. glandulosa
On mosses: doubtful: Amer.
-15 mm. high, white: sp. 5-7x3-3·5µ,: Cuba P. misella
-6 mm. high, whitish: among moss on dead wood: U.S.A. P. tenera
Erumpent from bark, yellowish, often deformed or flattened
Finland: sp. 4-6 X 2µ,: on Pinus P. paradoxa
U.S.A.: -4 mm. high
Yellowish: on Carya alba P. spathulata
Brownish: sp. 10-15x6-7·5µ,: on Alnus P. alnicola
Tierra del Fuego: sp. 2·5-4 X 1·5µ,: -8 mm. high: on Nothofagus P. fuegiana
PISTILLARIA 475
Not so: typical species on dead plant-remains
Head very short, globose or subglobose, 0·3-1 mm. long
Sp. 17 x 4µ: on Carex-leaves: France P. Boudieri
Sp. 7·5-12X3·5-5µ: base of head umbilicate: on Phragmites,
rarely Rubus or Juncus P. capitata
(Compare Physalacria, Pistillina, Dimorphocystis)
Head cylindric, elongate, 4-20 mm. long (see below)
White: stem short: dubious
France: -ro mm. high: stem 1-2 mm.: sp. 6µ long: on
Alnus-leaves P. epiphylla
U.S.A.
-12 mm. high: on Zea P. trichomorpha
-30 mm. high: stem long: head 2-4 mm., short: sp.
ro-r4X2·5-3µ: on leaves P. cylindrospora
White, then yellowish or pale ochraceous
Sp. 4X3µ: -15 mm. high: on Quercus-leaves: Germany. P. epiphylla
Sp. 4-7X2-3µ: -15(-25) mm. high: on sticks and her
baceous stems: Europe P. uncialis
Sp. 7-9X 3-4µ: -30 mm. high: on Ulex-leaves: Mediter
ranean P. olea
Sp. 10-13X5-6µ: -15 mm. high: on Juncus-stems: Eng
land P. subuncialis
Head shortly cylindric, clavate or ovoid, 0·5-5 mm. long
Sp. subglobose or subcordate
Sp. 8-roµ wide: very slender: Finland . P. anceps
Sp. 7 · 5 X6µ: -5 mm. high: on coniferous wood: Oregon P. fusiformis
Sp. 3-5X 3-4µ: -4 mm. high
Sp. triangular-cordate: on grass-leaves: Europe, Canada . P. culmigena
Sp. subglobose: on leaves of deciduous trees: Europe,
U.S.A. P. diaphana
(See also P. uliginosa, var. albo-lutea)
Sp. ellipsoid
Sp. 3-4X1-2µ, very small: N. temp. P. albobrunnea
Sp. 4-7X 2-3µ: glabrous
On Abies-twigs: -3 mm. high, yellowish or greyish: sp.
7x2µ: Germany P. lignicola
Usually on leaves of deciduous trees: white: sp. 4-6X
2-3µ: Europe P. pusilla
(On fem-stems: Cuba see P. subpellucida)
Sp. larger
Sp. 1-3µ wide, very ,1arrow
10-30 mm. high: head 2-4 mm.: sp. ro-14x2·5-3µ:
on dead leaves: U.S.A. P. cylindrospora
0·5-1 mm. high: sp. 10-20X1-2µ: on twigs of
Callirhoe: U.S.A. P. Batesii
2-5 mm. high: sp. 6-rox2-3µ
On stems of Epilobium: gelatinous-viscid, tough:
Europe, N. Amer. . P. typhuloides
On leaves of trees: Europe (? S. Amer.): see P. setipes
(Typhula mucor)
PISTILLARIA
Sp. 3-5 µ, wide
Sp. 9·5-12·5X3-3·5µ,: 2-spored: -4 mm. high: stem
pilose: on Juncus and Typha: France . P. typhicola
Sp. 9·5-11 x4·5-5µ,: 1-2-spored: -2 mm. high,
yellowish: on spots of leaves of Populus: Europe P. maculaecola
Mostly 4-spored
On ferns: -10 mm. high, yellowish: sp. 8-9X3-4µ,:
Europe P. Todei
On Epilobium-stems: -5 mm. high, rather gelatinous
viscid, tough: sp. 6-10X2·5-3·5µ,: Europe, N.
Amer. P. typhuloides
On twigs of Cytisus: -5 mm. high: sp. 10 X5µ,:
Tunis P. cytisi
On Rhizopogon (Gasteromycete): -6 mm. high: sp.
6-8X4µ,: Germany P. mycophila
On leaves of deciduous trees, perhaps also on twigs
and bark: --12 mm. high: stem hairy or glabrous:
sp. 6-12X3-4·5µ,: N. temp.(? S. Amer.) P. setipes
Sp. unknown or uncertain
10-25 mm. high: on cones of Pinus: France P. peronata
-2 mm. high: on fern-stems: Cuba, Montevideo P. subpellucida
-2 mm. high: on bark of Ulmus: Canada P. subfasciculata
P. albobrunnea Pat.
Tab. An. 1883, No. 52.
Synonyms: Pistillaria diaphana var. albobrunnea Quel., Ass. Fr. 1883, 506.
? P. inaequalis sensu Pat., Tab. An. 1883, No. 46.
? Clavaria quercicola Imai, Tr. Sapporo Nat. Hist. Soc. 11, 1929, 41.
2-3 mm. high: head elongate, ovoid, white, sometimes forked: stem 1-2 mm., slender,
glabrous, brown, distinct, with oxalate crystals.
On dead leaves of Pyrus: France.
Spores 4X1·5-2µ,, subcylindric.
From P. pusilla, this seems to differ only in the brown stem and narrow spores
(? correct): possibly it is only a variety, for the base of the stem may be brownish in
P. pusilla. This was Quelet's interpretation, because P. diaphana sensu Quel. (non
sensu Pat.) is evidently the same as P. pusilla. Compare TyphulaPatouillardii, which
may be the scleroticolous form.
Patouillard's description of P. inaequalis differs in the white, hairy stem and the
habitat (grass-leaves), but it has the same very small spores, for which reason it cannot
be P. inaequalis Lasch(see P. setipes):
-2 mm. high, white: head ovate-ellipsoid: stem finely hairy, longer or generally
shorter than the head: spores 3-3·5 X1·5µ,: on dead grass-leaves: France.
The description of Clavaria quercicola agrees very closely with that of P. inaequalis
sensu Pat., but the habitat, on Quercus-bark, is again different, or indicates P. sub
fasciculata:
Solitary or gregarious, small, white, subcylindric, truncate, simple: spores 3-3 ·5 X
1·5-2·5µ,, white, smooth, ellipsoid: on decaying bark of Quercus, on the ground in
woods, Japan.
PISTILLARIA 477
P. alnicola Pk.
Rept. N.Y. St. Mus. 42, 1890, 29, t. 2, f. 22-4.
2-4 mm. high, sessile or scarcely stalked, ovoid or oblong, obtuse, sometimes com
pressed or irregular, erumpent glabrous, brownish ochraceous then reddish bay, internally
white and spongy.
On branches of Alnus incana: U.S.A. (Cascadeville, Adirondack Mts.).
Spores 10-15 x 6-7·5µ,, white, smooth, ovoid, acute at one end.
Basidia 4-spored.
Perhaps Clavariadelphus fistulosus var. contortus.
P. ampelina Bourd. et Galz.
Hym. Fr. 1928, 136.
Head 0·4-1 X 0·07-0· 1 mm., cylindric, obtuse, simple or forked,purplish then blackish
purple: stem 0·2-0·6 mm. long, short, distinct, glabrous, concolorous.
On dead stems of Vitis: France.
Spores 7-11 X 6-8µ,, white, subglobose.
Basidia 25-30 X 9-10 µ,: sterigmata 2.
Hyphae 2-3 µ, wide, with clamps.
Bourdot and Galzin say this is near P. uliginosa, the microscopic characters of which
are not known. But, except for the darker colour, I do not see how it differs from
P. micans, for which Bourdot and Galzin have described almost as narrow hyphae,
3-4µ, wide, as in Ceratellopsis or Pterula, rather than Pistillaria.
P. anceps (Karst.) comb. nov.

Basinym: Typhula anceps Karst., Finl. Nat. o. Folk 48, 1889, 386.
Synonym: Clavaria gracilis B. sensu Karst., ibid. 37, 1882, 181 (non B. et Desm.
= Pterula gracilis).
Club very slender, white, puberulous with projecting sterigmata: no sclerotium:
spores 8-10µ,, round, white: Finland.
Little known. ? reduced Clavaria acuta.
P. Bartholomaei E. et E.
Proc. Nat. Ac. Sci. Philad. 1893, 441.
1 mm. high, gregarious, blackish pink, subcylindric or compressed, obtusely
accuminate, stem indistinct.
On dead stems of Callirhoe involucrata (Malvaceae): U.S.A. (Rockport, Kan.).
Spores 2-3µ,, globose.
Basidia 10-12 x 2-2·5µ,.
The microscopic details seem doubtful.
P. Batesii Pk.
Journ. Myc. 14, 1908, 3.
0·5-1 mm. long, very small, densely gregarious or caespitose: head ovate or oblong,
obtuse, sessile or narrowed into a short stem-like base, soft, pallid when fresh, drying
greyish cinnamon.
On twigs of Callirrhoe involucrata (Malvaceae): U.S.A. (Neb.): apparently parasitic.
Spores 10-20 X 1-2µ,, hyaline, filiform, often slightly curved.
Little known. The spores do not suggest basidiospores, though such acicular ones
occur in some species of Peniophora.
P. bellunensis Speg.
Michelia 2, 1879, 244.
1-2 mm. high: head 0·3-0·35 mm. wide, white, minutely granular: stem 0·1 mm.
wide, pinkish white, villous.
PISTILLARIA
On dead stems of Vitis: Italy (common near Belluno).'
? P. micans.
P. bipindiensis (P. Henn.) comb. nov.
Basinym: Typhula bipindiensis P. Henn., Engl. Jahrb. 30, 1902, 42.
3-5 mm. long, 160-220µ, thick, simple, fleshy, filiform, clavate, substipitate, fuscous
rufous, rounded at the apex, attenuate to the base.
On the underside of fallen leaves, gregarious: Cameroons (Bipindi).
Spores 5-6X3µ,, white, fusoid: basidia 4-spored.
Hennings says it seems to arise from a sclerotium.
P. Boudieri Pat.
Tab. An. 1887, 573.
Text-Fig. 21 I.
4-6 mm. high, white: head subglobose, not depressed or umbilicate
at the base: stem slender, finely villous, long.
On dead leaves of Carex: France.
Spores 17X4µ,, white, oblong, slightly curved.
I doubt if this is any more than P. capitata with long spores.
P. capitata (Pat.) Sacc.

Syll. Fung. 6, 1888, 759.
TEXT-FIG. 2II Basinym: Sphaerula capitata Pat., Tab. An. 1883, No. 60.
Pistillaria Synonym: Pistillina Patouillardii Quel. Ass. Fr. 1883, 506 (haud
Boudieri Pistillaria Patouillardii Quel. 1883 = Typhula Patouillardii).
(from Patouil Text-Figs. 37, 38, 42, 212.
lard, Tab. An. 1-6 mm. high, superficial, scattered, gregarious or subcaespitose,
n. 573).
white, the base of the stem often brownish: head 0·3-0·6 mm. wide,
globose or subglobose, hollow, fragile, minutely umbilicate at the base: stem 0·8-5·5
mm. long, 40-50 µ, wide at the apex, 80-100µ, wide at the base, capillary, tapering
slightly, diaphanous, sparsely pilose.
On dead stems and leaves of Phragmites communis (Quel., Pat., Rea, Corner), rarely
on dead leaves of Rubus (Rea) or dead stems of Jltncus obtusifiorus (Corner), late
summer, autumn: France, England (probably common locally).
Spores 7 ·5-13X3·5-5µ, (Corner), 7 ·5-10X3·5-4µ, (Rea), 6X3µ, (Bourd. et Galz.),
white, smooth, ellipsoid, blunt, flattened adaxially, obliquely apiculate, aguttate, the
contents cloudy-vacuolate.
Basidia 24-35x7-11 µ,, clavate, but broadest shortly beyond the middle: sterigmata
2-3-4, 5-10µ, long.
Cystidia none.
Hymenium covering the whole head, abruptly delimited at the stem-apex by a
shallow groove, not thickening: subhymenium I 2-20µ, thick, composed of 3-5µ, wide,
thin-walled hyphae with short, interlocking cells 5-10µ, long.
Caulocystidia -rooµ, long, 2-4µ, wide at the base, narrowed rather abruptly just
beyond the base and tapering gradually to the subacute apex, 1µ, wide, simple or
sparingly branched, aseptate, thin-walled, colourless, smooth, scattered, shorter
towards the apex of the stem.
Hyphae thin-walled, inflating, clamped at all septa: hyphae in the stern,. 3-12 µ, wide,
forming a compact cylinder of longitudinal hyphae, the surface hyphae with slightly
thickened, agglutinated walls, the central hyphae 3-6µ, wide and often pulled some
what apart (but rarely leaving a hollow as in P. setipes), sometimes the hyphae within
the agglutinated sheath elongating excessively and becoming coiled like a spring: the
surface-hyphae narrower, 3-5µ, wide, towards the apex of the stem: short-celled,
5-17µ, wide, at the base of the stem, with slightly thickened, brownish walls: hyphae
PISTILLARIA 479
of the head 3-8µ, wide, long-celled, and pulled apart by the enlargement of the head,
a few hyphae traversing the hollow but most lying against the subhymenium, the walls
not agglutinated or mucilaginous, the cavity in the head filled with air.
Mycelial hyphae 2-3 µ, wide, slightly thick-walled, clamped.
This fungus is very common in the fen-districts of
Cambridgeshire and, probably, wherever its usual host,
Phragmites communis, occurs. In the shape of the fruit
body, with subglobose head, it bears the same relation to
P. setipes as Typhula quisquiliaris to T. phac01-rhiza or T.
gyrans and as Dimorphocystis capitatztS to D. laevis; that
is, it presents the extreme form of limited apical growth,
the head becoming wholly fertile as soon as the hymenial
factor develops and approaching a spherical form through
intercalary growth of the hymenium. In P. capitata the
hymenium is so sharply delimited, or initiated, and its
intercalary expansion is so great in the confined space at
the apex of the primordial shaft, that it becomes pressed
down over the top of the stem to form the minute, but TEXT-FIG. 212. Pistil/aria
characteristic, groove. Thus it differs from Pistillina capitata, X 10.
hyalina in which intercalary growth is so slight that the
hymenium, formed in the centre of the top of the primordial shaft, remains
surrounded by a sterile margin, as in an apothecium.
P. culmigena Mont. et Fr.

Ann. Sci. Nat. 1836, t. 12, f. 2: Donk, Rev. Nieder!. Homobas. Aphyll. 2, 1933, 96:
Lundell, Sv. Svamp. II-12, 1938, 20: (non Quel., Karst., Schroet. = Ceratellopsis
aculeata; non Pat., Tab. An., No. 265 = Pistillaria pusilla).
Synonyms: Clavaria culmigena (Fr.) Karst., Finl. Nat. o. Folk 37, 1882, 177.
Pistillaria cardiospora Quel., Ass. Fr. 1883, 507: Pat., Tab. An. 1883, No. 55 (sec.
Donk, Lundell).
Typhula culmigena (Fr.) Schroet., Pilzfl. Schles. 1888, 439: (quoad nomen).
0·5-3 mm. high, 0·2-0·4 mm. wide, white: head ellipsoid to ovoid, obtuse, com
pressed: stem 0·2-0·5 mm. long, very short, distinct, glabrous or puberulous, pellucid.
On dead grass-remains, perhaps on other herbaceous remains: Europe, Canada
(Man.).
Spores 3-5 X 3-4µ, white, irregularly triangular, almost cordate (Lundell).
Basidia 20-35 X 3-7 /.L: sterigmata 2-4, 5-6 /.L long.
Hyphae 1·5-4·5µ,, thin-walled, clamped (Bourd. et Galz., Donk): with abundant
calcium oxalate crystals in the tissue.
Donk examined the type-collection of P. culrnigena in Paris and found the charac
teristic spores of P. cardiospora. Quelet interpreted P. culmigena as a species with
ellipsoid-cylindric spores 6-7µ, long: Karsten and Schroeter ascribed to it ellipsoid
spores 4-6 X 2µ. Their fungi suggest to me Ceratellopsis aculeata. Spegazzini's
record of P. culmigena from the Argentine, with subglobose spores 4-5 µ,, needs con
firmation. It may be P. diaphana, which differs mainly in the more regularly sub
globose spores, and its habitat on leaves.<of trees.
P. culmigena may be a Ceratellopsis.
P. cupressiformis B. et C.
0·5 mm. high: head cupressiform (? narrowly conical), greyish pulverulent: stem
short, setiform, black.
On twigs: Cuba.
This may be a Myxomycete, such as Comatricha, or a Hyphomycete.
PISTILLARIA
P. cylindrospora (Atk.) comb. nov.
Basinym: Typhula cylindrospora Atk., Ann. Myc. 6, 1908, 60.
1-3 cm. high, white, filiform: head 2-4 mm. long, cylindric, with a few crystals
among the hyphae: stem 8-26 mm. long, solid, glabrous except the villous base.
On rotting leaves: U.S.A. (Ithaca, N.Y.).
Spores 10-14 X 2·5-3µ, white, oblong-cylindric, straight or slightly curved.
Basidia 25 X 5-6µ, 4-spored.
This seems close to P. setipes, but with a longer stem and narrower spores such as
described for Typhula hirsuta and T. mucor (see under P. setipes).
P. cytisi Pat.
Joum. de Bot. 1894, 220.
3-5 mm. high, white, gregarious: head globose, ovoid, or spathulate: stem indistinct:
sclerotium none.
On dead twigs of Cytisus trifiorus (Leguminosae): Tunis.
Spores 10 X 5 µ,, white.
Perhaps only a state of P. setipes.
P. diaphana Fr.
Syst. Myc. 1, 1821, 498: sensu P11t., Tab. An. 1883, No. 51: (non sensu Quel., Ass.
Fr. 1883, 506 = P. pusilla): = P. pusilla sensu Kauffm., Pap. Mich. Ac. Sci. Arts
Lett. 9, 1929, 207.
Synonym: Clavaria diaphana (Fr.) Karst., Finl. Nat. o. Folk 37, 1882, 178.
1-4 mm. high, white: head 1-2 mm., 0·3--0·4 mm. wide (Kauffm.), subcylindric and
acute, then ellipsoid and blunt: stem 1-2 mm. long, slender, pellucid, glabrous,
becoming yellowish at the minutely bulbous base, mostly shorter than the club.
On dead leaves of Alnus, Fagus, &c.: Europe, U.S.A. (Mich.).
Spores 4"x 3 µ, (Pat.), 3-3·5µ (Kauffm.), white, smooth, ovoid, or subglobose.
Basidia 18-20 X 3·5-4µ, subclavate, with 1-2-3-4 sterigmata (Kauffm.).
Abundant oxalate-crystals in all parts (Pat.).
I have interpreted this as a species growing, like P. setipes and P. pusilla, on dead
leaves of deciduous trees but with small subglobose spores. It may not be specifrcally
distinct from P. culmigena. As described by Fries, P. diaphana could be either P.
setipes or P. pusilla, and rather than dismissing the name as doubtful and substituting
a new one, I have taken it in Patouillard's sense, as first indicating a specific distinction.
P. elegantula (Karst.) comb. nov.
Basinym: Typhula elegantula Karst., Not. Soc. Fenn. II, 1871, 222: Act. Soc. Sci.
Fenn. 15, 1888, 185, t. 7: Imai, Jap. Joum. Bot. 8, 1936, 1 I.
Synonym: Clavaria elegantula Karst., Finl. Nat. o. Folk 37, 1882, 179.
Head 2·5-4 mm. long, rose-pink, cylindric, attenuate at both ends, or subclavate
and obtuse: stem 5-16 mm. long, puberulous, filiform, subflexuous, pellucid, pallid.
On dead stubble and roots of Triticum repens: Finland.
Spores 6--g X 3-4µ, white, smooth, ellipsoid.
This species in colour and habitat resembles Typhula incarnata, but it has a shorter
head, smaller spores, and no sclerotium. I think it is more likely to be a pink Pistillaria
setipes than a 'sclerotium-free' form of Typhula, and P. granulata is, perhaps, the
connecting link.
P. epiphylla (Quel.) comb. nov.

Basinym: Clavaria epiphylla Quel., Ass. Fr. 1883, 505, t. 6, f. 16: Pat., Tab. An.
1884, n5, No. 259.
IO mm. high, pellucid white, cylindric, filiform, pointed or obtuse, simple: stem
1-2 mm. long, very short, glabrous.
PISTILLARIA
On dead leaves of Alnus, solitary, gregarious, or subcaespitose: France.
Spores 6 µ,, pruniform, white: basidia 4-spored (Pat.).
Quelet thought this might be P. aculina (which is Pterula gracilis), but the spores
are too small. It may be an unusual form of P. setipes which is very variable, but the
following notes of Killermann's, on a collection which he referred to Pistillaria
Patouillardii ( = Typhula Patouillardii, p, .677, with very small fruit-bodies), suggests
that there is a distinct epiphyllous species with long heads as in P. uncialis:
-15 mm. high: head c. 10 mm. long, filiform, linear, yellowish: stem arising from
a whitish disk: spores 4 X 3 µ,, ellipsoid, 2-guttulate: on leaves of Quercus: Germany
(Zeitschr. f. Pilzk. 18, 1934, rn7, f. E).
P. fuegiana (Speg.) comb. nov.

Basinym: Clavariafuegiana Speg., Fungi, n. rn6, p·. 42: Sacc., Syll. Fung. 6, 1888,
720.
5-8 mm. high, 1-2 mm. wide above, erumpent in groups of 3-5 through bark, yellow,
obovate-spathulate above, obtuse, undulate-rugulose, attenuate cuneately into a smooth
glabrous stem 4-6 X 0·4-0·5 mm.
On fallen branches of Nothofagus: Tierra de! Fuego (common).
Spores 2·5-4 X 1-5·5 µ, (? 1-1·5 µ,), white, smooth, cylindric or suballantoid, obtuse,
rounded or subtruncate.
This is doubtfully a Basidiomycete, but it may be related to Lachnocladium aurantia
cum or Pistillaria spathulata. Compare also Clavaria? dubiosa (p. 266).
P. fulgida Fr.
Epicr. 1838, 587: Quel., Ass. Fr. 1883, 506: Pat., Tab. An. 1883, No. 47.
Synonyms: ? Clavaria minuta Pers., Myc. Eur. 1, 1822, 189.
Pistillaria fulvida Karst., Hedw. 1892, 301.
Head 1-2 X o· 1-0·25 mm., tawny or tawny-orange, cylindric or lanceolate, subacute,
flexuous, smooth: stem -5 mm. X 0·04-0·2 mm. (very short, Bourd. et Galz.), whitish,
yellow, or concolorous, equal or constricted at the apex.
On dead stems of Dipsacus pilosus and Helianthus tuberosus, on dead leaves (Quel.):
western Europe.
Spores 8-10 X 4-5 µ,, white, oblong or subcylindric (Bourd.): 8 µ,, ovoid-oblong,
hyaline (Quel.).
Basidia 18-25 X 6-9 µ,: sterigmata 2-4, 4-6 µ, long (Bourd. et Galz.): sterigmata I,
rarely 2 (Quel., Gilbert).
Hyphae 2-8 µ, wide, slightly thick-walled, with masses of calcium oxalate among
the longitudinal hyphae.
This species has been recorded from Malaya, thus: 'A typo·_ differt quia adhuc
minor et sursum non vel vix clavata: sporae 7 x 4 µ,, ellipsoid, hyalinae: in fructibus
putridis Nephelii glabri' (Sacc. et Paol., Att. R. Inst. Venet. 6, Ser. 6, 1888, 15).
I have not found the fungus in Malaya, but feel sure it is Pteruloid: cf. P. bipindiensis.
The· description of P. fulvida seems indistinguishable: 2 mm. high, cylindric
clavulate, smooth, glabrous, fulvescent, base becoming brown, stem indistinct: on
leaves of Cornus sanguinea, Finlar1d.
It would seem that Persoon's epithet has priority.
P. fuscipes (Pers.) comb. nov.

Basinym: Clavariafuscipes Pers., Myc. Eur. 1, 1822, 158.
Synonym: Typhula fuscipes (Pers.) Fr., Epicr. 1838, 586.
Head subflexuous, subsulcate, obtuse, simple, glabrous, incrassate, cinnamon-yellow:
stem bistre, thenfuscous-black, long: on twigs (France).
A dubious species, recalling Clavariadelphus fistulosus or C. junceus.
5119 I i
PISTILLARIA
K.illermann's record of it seems to be Typhula erythropus (Zeitschr. f. Pilzk. 18,
1934, 138, f. C.).
P. fusiform.is Kauffm.
Pap. Mich. Ac. Sci. Arts Lett. 5, 1925, II9, t. 2, f. 1 r.
3-5 mm. tall, white: head subfusiform acuminate, somewhat curved, subcompressed,
sometimes furrowed on the flattened side, fleshy: stem 1 mm. long, slender, terete,
puberulous; no sclerotium: rather spongy-fleshy.
On decayed coniferous wood, gregarious: U.S.A. (Oregon, Mt. Hood).
Spores 7·5 X 6µ, white, smooth, ovoid-ellipsoid to subglobose.
Basidia 45 x 8--gµ, 2-3-spored.
Cystidia none.
In shape and appearance this is said to be like Ceratellopsis rosella var. ramosa Pat.
It may be a true Clavaria, Mucronella, or Deflexula, but cannot be classified cor
rectly until the hyphae are known. Compare P. subfasciculata.
P. gilva (Lasch) comb. nov.

Basinym: Typhula gilva Lasch in Rabenh. Fung. Eur. 619 (? 419): Winter, Pilze 300.
Head honey-yellow, obovate-oblong, 5-6 X 0·3-0·8 mm. (Donk MS.): stem paler:
no sclerotiurn.
On the ground among moss: Europe.
Spores ovoid.
Donk examined the type-collection but was unable to make out any details.
P. glandulosa (Preuss) comb. nov.

Basiriym: Typhula glandulosa Preuss, Linn. 1853, 725.
Head turgid, glabrous, pallid white: stem long, pubescent with moniliform glandular
hairs.
Spores inaequilateral: basidia forked with long sterigmata.
On the ground in damp places: Europe.
Sourdot and Galzin suggest this may be Hirsutella.
P. granulata Pat.
Tab. An. 1884, 118, No. 266: (? = Clavaria granulata Pers., Myc. Eur. 1, 1822, 192).
1-2 mm. high, scattered or gregarious, glabrous; head white, then reddish or flesh
colour, blunt, cylindric or subclavate, granular: stem very short, 0·2-0·3 mm. long,
distinct, of clearer colour.
On rotting leaves of Populus : France.
Spores 6 X 3µ, ovoid: basidia 4-spored.
Quelet treated this as a variety of P. micans (FI. Myc. 1888, 400), but it seems only
a pirik form of P. pusilla, connecting it in this respect with P. elegantula.
P. lignicola (K.illerm.) comb. nov.

Basinym: Typhula lignicola Killerm., Zeitschr. f. Pilzk. 18, 1934, 103, f. A.
2-3 mm. high, glabrous, dingy yellowish white, or greyish: stem c. 2 mm. long,
filiform: head I mm. long, cylindric-ovate.
On twigs of Abies: Germany.
Spores 7 X 2-3µ, white, cylindric, smooth, 2-guttulate.
Basidia 20µ long, with 4 sterigmata.
Stem-hyphae 4-5µ wide, with slightly thickened walls.
This grew together with Typhula abietina which had sclerotia, but P. lignicola had
no sclerotia. They do not seem to be forms of the same species because the spores of
T. ab�etina are considerably larger, 8-10 X 4-5µ.
PISTILLARIA
P. limicola (Saut.) comb. nov.
Basinym: Typhula limicola Saut., Hedw. 1876, 150.
Club pinkish rufescent: stem white, setuliform, 1·5-3 cm. long.
On the ground: France, Germany.
P. lividula (Roll.) comb. nov.
Basinym: Typhula lividula Roll., Bull. Soc. Myc. Fr. 14, 1898, 83, t. 9, f. 2.
2-3 mm. high: head olivaceous, then hoary white: stem rose-pink, bulbous at the
base.
Spores 8 X 3-3 ·5 µ, white, smooth, oval-oblong.
On dead remains of Helleborus: Corsica.
Perhaps related to P. elegantula.
P. maculaecola Fckl.
Symb. 31, 1869: Fries, Hym. Eur. 1874, 689: Pat., Tab. An. 1883,' No. 50.
0·5-2 mm. high: head yellowish (jonquil), 0·7--0·8 X 0·35 mm., ellipsoid and sub
acute to subglobose or ovoid: stem o·15 mm. wide, white, sparsely pilose, incrusted
with oxalate-crystals.
On desiccated, sordid spots on living and dead leaves of Populus tremula (Fckl.);
apple, &c. (Quel.): Europe.
Spores 9·5-11 X 4·5-5 µ, ellipsoid-pip-shaped (Donk): 6-7µ, ovoid, pruniform
(Quel.).
Basidia 18-25 X 6-7µ (Donk): sterigmata 1-2, 5 · 5µ long (Donk), 1 only (Quel., Pat).
I have amended this description from Donk's MS. notes on the type-distribution.
Except for the habit and rather wide spores, which may merely reflect their disporous
(monosporous) origin, the species seems to belong to the complex of P. setipes.
P. micans Fr.
Syst. Myc. 1, 1821, 497: Boud., le., t. 177: Wehm., Pap. Mich. Ac. Sci. Arts Lett.
20, 1935, 264.
Synonyms: Stilbum micans Pers., Myc. Eur. 1, 1822, 355.
Clavaria micans (Fr.) Karst., Finl. Nat. o. Folk 37, 1882, 177.
Scleromitra coccinea Cda., Sturm. Deutsch!. FI. Pilze 7, 1829, 59, t. 27.
Pistillaria coccinea (Cda.) Fr., Epicr. 1838, 587.
P. micans var. coccinea Pat., Tab. An. 1883, No. 44.
P. incarnata Desm., Ann. Sc. Nat. 1843, 375: Pat., Tab. An., No. 569.
P. carnea Preuss, Linn. 1851, 15_1: (teste Quel.).
? Mucronella viticola Pass. et Beltr., Fung. Sicul., No. 2.
1-4 mm. high, rose-pink to reddish pink, or the stem whitish: head 0·5-2·5 X 0·5 mm.
(0·6-1 x o·z5-0·35, Wehm.), clavate, obovate or oblong, obtuse, pruinose and glistening:
stem 0·5-0·8 x 0·25 mm., translucent, cylindric: flesh soft, drying hard.
On dead herbaceous stems and leaves: Europe, U.S.A., Canada (Nov. Scot.)
(? New Zealand, Grev. 8, 1879, 57).
Spores 8-13 X 5-7µ, white, smooth, ellipsoid or oblong-ellipsoid, blunt.
Basidia 30-5ox7-10µ: sterigmata 2(-4), 7-9(-r4)x2-3µ.
Hyphae 2-8 µ wide, rather long-celled, thin-walled, inflating, clamped (Corner):
no cystidia or caulocystidia: oxalate crystals on the stem.
var. Sedi Karst.
Finl. Nat. o. Folk 37, 1882, 177.
Very small: head 0·2 x o·1 mm.: stem o· 1 mm. long.
On dead stems of Sedum acre: Finland (Mustiala).
This species is certainly related to P. setipes, though much smaller and brilliantly
coloured: thus it connects P. setipes w:th P. elegantula. The hyphal construction is
PISTILLARIA
identical with that of P. setipes. The colour of the head varies from pale pink to deep
reddish pink (P. coccinea), and the stem may be white or pale pink: the basis of the
coloration is not known: possibly P. ampelina (purple to blackish purple) is only an
extreme state. In size of the fruit-body there appears to be every gradation to the
_minute var. Sedi: usually it is 1-1·5 mm. high.
The spore-data which I have been able to collect are:
8 µ., ovoid pruniform (Quel.).
9-10 X 5-6 µ. (Karst.).
8-12 X 6-7 µ. (Rea).
9-13 X 5-'7 µ. (Bourd. et Galz.).
8·5-10·5 X 5-6 µ. (Wehm.).
10-12·5 x 5·7-7 µ. (Donk, MS.).
The fact that they are mostly di-spores probably explains their greater width and
blunter apex than in the mainly tetrasporic P. setipes. Thus P. granulata may be the
tetrasporic state of P. micans.
De Bary assigned Sclerotium laetum Ehr. to P. micans (Comp. Morph. Fungi 42),
in which he is copied by Killermann, but Wehmeyer says there is no sclerotium, and
it has not been described by other mycologists.
Quelet wrote 'herisee de cystides a la loupe' (Bull. Soc. Bot. Fr. 24, 1877, 326).
He may have mistaken the large sterigmata, as no cystidia have been described.
Concerning P. incarnata, Donk (MS.) maintains that it is a distinct species with more
elongate head and stem (-3 mm. high, overall), 2-4 sterigmata and narrower spores
10-13·5 X4·7-5·3 µ.. It was described originally from leaves of Scirpus (?) in France.
It seems to fall, however, within the range of P. micans though, certainly, it requires
investigation.
P. misella (B. et C.) comb. nov.
Basinym: Clavaria misella B. et C., Journ. Linn. Soc. Bot. 10, 1868, 339: Burt,
Ann. Mo. Bot. Gdn. 9, 1922, 49, t. 9, f. 76.
-1·5 cm. high, white, simple, clavate, obtuse: stem slender, tomentose-dilated at
the base.
Attached to mosses, the stem attached to a cluster of 2-3 moss-leaves: Cuba
(Wright, 222).
Sp?res 5-7 X 3-3·5 µ., white, smo�th (Burt).
Basidia 4-spored. Hyphae 4-5 µ. wide (Burt).
The true position of this little-known fungus remains to he discovered. Clavaria
paupercula B. (MS.), with which Burt compares it, seems to be a true Pistillaria, accord
ing to the 'type-collection' (No. 217, Venezuela) in the Kew herbarium: it has 2-4 µ.
wide, clamped, long-celled hyphae, with submucilaginous walls.
Massee's record of C. misella from New Zealand is certainly erroneous (Tr. N. Zeal.
Inst. 39, 1906, 40).
P. montevidensis Speg.
Anal. Mus. Nac. Buen. Air. 23, 1912, 1 I.
5-6 X 0·5-1 mm., mostly 2-3 caespitose, erect, cylindric then flattened, pinkish
cinnamon, then rose-pink-orange, slightly fleshy, very delicate: no sclerotium.
Spores 6 X 3, white, smooth, ovoid, inaequilateral.
On the ground among mosses: Montevideo.
Perhaps allied with Clavaria helicoides.
P. mycophila (Fckl.) comb. nov.

Basinym: Typhula mycophila Fckl., Symb. 32, 1869.
2-6 mm. high: head oblong-ovate, obtuse, distinct, sordid: stem simple or branched,
glabrous.
PISTILLARIA
On rotten Rhizopogon rubescens (Gasteromycetes): Germany.
Spores 6-8 X4 µ, white, smooth, ellipsoid.
P. oleae (Maire) comb. nov.

Basinym: Typhula Oleae Maire, Bull. Soc. Myc. Fr. 44, 1928, 53.
10-30X0·5 mm., solitary or gregarious, fleshy-elastic, solid glabrous, without
sclerotium: taste mild, smell slight.
Head cylindric, linear, pale ochraceous, soon pruinose with the spores, rather acute,
attenuate or not into the concolorous stem: flesh white.
On dead fallen leaves of Olea europaea: N. Africa (Mauretania).
Spores 7-9 X 3-4 µ, white, smooth, oblong, rounded at the apex, obliquely apiculate,
I-several guttulate.
Basidia 27-35X6 µ, elongate clavate, often flexuous: sterigmata 4, 6 µ long.
Hyphae 5-13µ wide, longitudinal, unequal.
This is said to be near Typhula phacorrhiza and Clavariadelphus junceus. It seems
to me related also to P. uncialis.
P. paradoxa (Karst.) comb. nov.

Basinym: Clavaria paradoxa Karst., Not. Soc. Fenn. 9, 1882, 372.
Synonym: Clavariella paradoxa Karst., Rev. Myc. 3, 1881, 21.
Erumpent, simple, fleshy, solid, variable in shape, spathulate, compressed, truncate
or undulate, smooth, yellowish: stem paler, villous-pruinose, glabrescent.
On bark of Pinus: Finland.
Spores 4-6X2 µ, white, smooth, ellipsoid-cylindric.
Like P. spathulata, this suggests rudimentary Clavariadelp_hus fistulosus var. con
tortus, but the spores are much smaller.
P. peronata (Pers.) comb. nov.

Basinym: Clavaria peronata Pers., Myc. Eur. r, 1822, 190.
Synonym: Typhula peronata (Pers.) Fr., Epicr. 1838, 585.
Head 4 mm. long, fusiform-incrassate, short: stem 10--20 mm. long, pubescent,
attached to a villous white mycelium.
On cones of Pinus sylvestris: France (rare).
P. phaeosperma (P. Henn.) comb. nov.

Basinym: Typhula phaeosperma P. Henn., Engl. Jahrb. 28, 1901, 320.
2-4X0·18--0·2 mm., filiform, fusoid or clavate, obtuse, pallid then brownish,
scarcely attenuate into the stem.
On fallen leaves of Solanum in the forest: Nyasaland (Ukingaberge, Bulongwa,
2,200 m.).
Spores 4-4·5µ, brown, smooth, globose, r-guttulate.
Basidia 18-24X4-6µ: sterigmata 2-4.
Hennings suggested that a new genus Phaeotyphula might be made for'this, but it
has not been described. The brown spores may be as dubious as those of Pterula
hirsuta. P. phaeosperma may b.: Pteruloid.
P. purpurea W. G. Smith
Journ. Bot. 1873, t. 130, f. 10-12.
2 mm. high, purple, obovate: stem whitish, spotted carmine, often growing with a
blood-red mycelium.
PISTILLARIA
Spores 4 X 3µ. white (Rea): 2·5µ., subglobose (Sacc., Syll. 6, 754).
On damp rotten leaves: Gt. Britain (rare).
A dubious species, perhaps P. uliginosa.
P. pusilla Fr.
Syst. Myc. 1, 1821, 498: Pat., Tab. An. 1883, No. 49: Quel., Ass. Fr. 1883, 506:
(non Bourd., Rea, Denk = P. setipes: non Kauffm. = P. diaphana).
Synonyms: Clavaria pusilla (Fr.) Karst., Finl. Nat. o. Folk 37, 1882, 179.
Typhula pusilla (Fr.) Schroet., Pilz. Schles. 1888, 439.
Pistillaria ovata Fr., Syst. Myc. 1, 1821, 497: (non Quel. = P. setipes): Pat., Tab
An. 1883, No. 54.
Clavaria ovata Pers., Myc. Eur. 1, 1822, 190.
Clavaria ovata (Fr.) Karst., Finl. Nat. o. Folk 37, 1882, 178.
Typhula ovata (Fr.) Schroet., Pilz. Schles. 1888, 439: (non Karst.).
? Clavariafruticum Karst., Finl. Nat. o. Folk 37, 1882, 179.
? Typhulafruticum Karst., ibid. 48, 1889, 386.
? Pistillariafruticum (Karst.) Sacc., Syll. 6, 1888.
? Typhula candida var.fruticum Bourd. et Galz., Hym. Fr. 1928, 131.
Pistillaria culmigena sensu Pat., Tab. An., No. 265: (non Fr.).
Pistillaria diaphana sensu Quel., Ass. Fr. 1883, 506: (non Pat.).
? Pistillaria.quercus Oud., Ned. Kruidk. 3, 1903, 674: Donlc, Rev. Nieder!. Homobas.
Aphyll. 2, 1933.
? P. sagittaeformis var. rubi Killerm., Zeitschr. f. Pilzk. 18, 1934, 138, f. d.
Text-Fig. 213.
0·5-6 mm. high, white, linear or subclavate, lanceolate or obtuse, smooth, glabrous:
stem distinct (1-2 mm. long, Quel.) or indistinct, rarely forked.
On dead leaves of Betula (Quel.), Ulmus and Rubus (Fr.): Europe.
SporP-s 6µ., ovoid (Quel.): 5 x 2µ., ovoid (Pat.): 5 x 3 µ. (Massee).
Basidia with 4 sterigmata (Quel., Pat.).
1 have used this name in the sense of Quelet and Patouillard, who first gave micro
scopic precision to the species. It differs from small glabrous specimens of P. setipes
only in the smaller spores (so far as known) and I am
doubtful if it is any more than a variety. Fries
suggested that it might be young Typhula candida ( =
£
P. setipes), but I do not see how Fries could have
critically distinguished the species of Pistillaria without
the microscope. P. ovata is just such a dubious species.
As interpreted by Quelet and Massee it is a glabrous
P. setipes (Table XXXII, p. 491), but if Fries had
intended P. setipes, he would surely not have described
the same fungus twice again as Typhula Grevillei and
/ / ,.,..�., / / . .,-:<
T. candida. According to Fries, P. ovata has an ovoid
TEXT-FIG. 2i3. Pistillariapusilla, hollow head and glabrous stem, so that may be
it P.
x 10·
capitata. It may equally well be old specimens of
P. pusilla, as I reduce it. Thus, I regard P. pusilla as generally a small, always glabrous,
species with small spores in contrast to P. setipes which is generally elongate (at least
the stem), usually hairy, and with considerably larger spores. P. diaphana resembles
P. pusilla but has even shorter, broadly ovoid, spores. Admittedly, they need much
critical study of fresh material.
The description of P. Jruticum is:
2-4 mm. high, white or vellowish, clavate, attenuate into the rather long stalk: head
subclavate or subcylindric: spores 6 X 3µ.: on dead branches of Rosa pimpinellifolia:
Finland. It seems to be a rather elongate state of P. pusilla.
The fungus referred to T. Grevillei by Rea, with spores 4-5 X 2µ., probably belongs
PISTILLARIA
here: he may have combined the macroscopic characters of T. Grevillei with the
microscopic characters of P. pusilla, unless there are states of P. pusilla as well grown
as P. setipes and with hairy stems.
P. quercus may be the rudimentary state of P. pusilla or of some Ceratellopsis.
According to Donk, there is no type-material. Its description is:
800 x120 µ, white, clavate, tubercular, or irregular in form, somewhat floccose,
sessile, without sclerotium, completely fertile over the surface: spores 4 X 2·5µ, ovoid,
slightly constricted in the middle: basidia 8-12 X 6-7µ, with 4 sterigmata 3µ long:
paraphyses abundant, filiform, 1-1· 5 fL wide, longer than the basidia, often forked at
the apex: on dead leaves of Quercus robur: Holland.
P. diaphana sensu Quel., on dead leaves, had ovoid spores 5-6µ long, and clearly
belongs to P. pusilla. Similarly P. culmigena sensu Pat., but its haibitat was on grass
leaves.
I refer here also two collections of Killermann's. Thus:
Pistillaria sagittaeformis var. rubi, I.e.: 1-2 mm. high, 0·2 mm. thick, white: stem
short: hymenium covering the club: spores 6 X 3µ: basidia 30-50µ long, sterigmata
2 4-: on Rubus-twigs, Germany.
P. inaequalis sensu Killerm., I.e., 106, f. D: 3-4 mm. high, white, subclavate, some
what pruinose and swollen below: spores 6 X 3µ, 1 -guttulate: basidia ? 2-spored: on
rotten wood: Germany.
Whether P. pusilla sensu Josserand, with spores (7-)7·7-8·5(--9·3) X 4-5µ, belongs
here or to Ceratellopsis (e.g. C. sagittaeformis), I am uncertain: (Bull. Soc. Myc. Fr. 59,
1943, 31, f. 6.)
It would seem, therefore, that P. pusilla may occur on twigs, wood, and grass-leaves
as well as its usual habitat on the leaves of dicotyledonous trees.
P. rhodocionides sp. nov.

1-2 mm. high, pale rose-pink, rarely whitish, gregarious, often crowded, not
caespitose: head 150-250µ wide, subcylindric, subfusiform or subclavate, solid, more
or less sharply demarcated: stem 100-600 X 100-140·µ, distinct, slender, pink, glabrous.
On dead pods of Cassia alata: Singapore (Apr. 1945).
Spores 7-9·5 X 4·3-5·3µ, white, ellipsoid, smooth, aguttate.
Basidia 21-33 x7·5-9µ, narrowly clavate: sterigmata 4, 6-8µ long, sometimes
10-12µ, parallel or slightly divergent: cystidia none.
Hyphae 3-7(-10)µ wide, thin-walled, without clamps, the contents very pale pinkish,
often slightly swollen at the septa: on the surface of the stem, 2·5-3 · 5 fL wide and
slightly, but firmly, agglutinated: inner medullary hyphae with cells 30-90 µ long,
parallel, fairly compact.
Subhymenium at first narrow, composed of 1-2 layers of cells 8-12 X 2·5-4µ,
thickening to 25-35µ through the enlargement of the cells (15-30 X 3-7-10µ.) on
intercalary growth of the hymenium.
This very small species seems related to the temperate P. elegantula (larger fruit
body) and P. granulata (smaller spores). The hyphal construction is the same as in
P. micans and P. setipes except for the absence of clamps: thus, it is conceivably a very
reduced Clavaria, as C. helicoides. Nevertheless, it has the typical appearance and
habit of Pistillaria.
The blackened, dehisced pods of Cassia alata remain for many weeks on the old
persistent infructescences, at heights of 4-10 ft. from the ground, until they have
almost completely decayed. The fruit-bodies of the Pistillaria can develop on them,
therefore, only in continuously wet weather. I found that they could not surviv
desiccation, to which they are immediately exposed in the day-time, but that they
were able to develop very rapidly in adaptation, as it were, to the few days of suitably.
muggy weather. On pods which I kept in a dish they developed in 24 hours, even less.
PISTILLARIA
P. setipes Grev.
Scot. Crypt. FI. 6, 1828, 61 (index).
Synonyms: Clavaria trichopus Grev., Scot. Crypt. FI. 1, 1823, t. 49 (non Pers.
1822 = Clavulina cristata).
Clavaria setipes Grev., Scot. Crypt. FI. 6, 1828, 61 (index).
Cnazonaria setipes (Grev.) Corda, Sturm, Deutsch!. FI. Pilze 7, 1829, 55: Donk,
Rev. Nieder!. Homobas. Aphyll. 2, 1933.
Typhula Grevillei Fr., Ep. 1838, 585.
T. candida Fr., Monogr. 2, 1863, 282: Bourd. et Galz., Hym. Fr. 1928: Rea, Tr.
Brit. Myc. Soc. 12, 1927, 226: var. populina Killerm., Zeitschr. f. Pilzk. 18, 1934,
138, f. b.
Pistillaria inaequalis Lasch, Rabenh. Exsicc., No. 1930: Fr., Hym. Eur. 1874, 688:
Donk, Rev. Nieder!. Homobas. ApLyll. 2, 1933 (ut Cnazonaria inaequalis): Quel.,
FI. Myc. 1888, 451.
? T. hirsuta Cke., Grev. 8, 1880, 82: Bourd. et Galz., Hym. Fr., 1928.
Clavaria candiaula Karst., Finl. Nat. o. Folk 37, 1882, 180: ibid., 48, 1889, 386.
? Clavaria longipes Karst., Symb. Myc. Fenn. II, 1882, 70.
? Typhula longipes Karst., Finl. Nat. o. Folk 48, 1889, 386.
? Typhula mucor Pat., Tab. An. 1883, No. 472.
? Pistillaria viticola Pk., Rept. N.Y. St. Mus. 42, 1890, 28, t. 2, ff. 25-7.
P. cylindracea Karst., Hedw. 1892, 301.
P. pusilla Fr., sensu Rea pr. p., Bourd. et Galz., Donk.
P. ovata Fr., sensu Quel., Mass., Schroet. (ut Typhula ovata).
Text-Figs. 37, 39, 40, 41, 214-17.
1·5-12 mm. high, white, the base of the stem brownish, ageing yellowish: head
0·8-3 ·5 X 0·2-0·7 mm., ellipsoid-fusiform and subclavate, then elongate-pyriform to
clavate or obovoid, blunt, opaque, solid: stem 0·7-7·5 X o·1-0·25 mm., filiform, pellucid,
generally sp_arsely hairy, varying rather densely hairy to glabrous, the base minutely
erumpent.
On dead leaves of deciduous trees in damp places, gregarious or scattered, autumn
(mostly on Alnus, Betula, Populus, Salix, also on Corylus, Fagus, Fraxinus, Quercus,
Ulmus, and, perhaps, Pyrus): Europe.
Spores 6-12(-14) X 3-4·5(-5)µ, white, smooth, ellipsoid, blunt, flattened adaxially,
contents cloudy vacuolate, aguttate.
Basidia 20-33 x6-8µ, cloudy-vacuolate: sterigmata 2-4, mostly 4, 5-7·5µ long:
cystidia none.
Caulocystidia -120 X 3-7·5 µ wide at the base, tapering gradually to the subacute
apex c. 1 µ wide, shorter towards the apex of the stem and reduced to short lobes
below the head, simple, occasionally with 1-5, often recurved, branches thin-walled,
colourless, smooth: in some collections barely 50µ long: in other collections absent or
merely short, scattered protuberances -15 µ long, as normally occur near the apex of
.he stem.
Hymenium not thickening, soon covering the apex of the head: subhymenium
c. 15µ thick, composed of short-celled interwoven hyphae, the cells 6.-10 x 3-5µ.
Hyphae thin-walled, inflating, colourless, clamped (? without clamps in some fruit
bodies), thin-walled, not mucilaginous, not secondarily septate: with fairly abundant
oxalate crystals among the hyphae of the stem and on its surface, few or none in the
head.
Stem consisting of a more or less disrupted core of uninflated hyphae 3-5 µ wide,
often hollow, and a firm cylinder of inflated hyphae 3-12 µ wide, long.-celled, appear
ing pseudoparenchymatous in transverse section, 3-5 hyphae thick, the outermost
hyphae with firm, agglutinated, but scarcely thickened, walls: the same two layers
short-celled at the base of the stem, the cells of the outer layer with distinctly thickened
PISTILLARIA
yellowish or brownish walls and compacted into a firm pseudoparenchymatous pul
vinate base for the fruit-body.
Head consisting of a loose core of hyphae 4-8 µ wide, continuous with that of the
stem but not hollow, a surrounding cylinder of more compact hyphae, 8-20µ wide,
continuous with the outer cylinder of the stem, and the hymenium over the surface:
the junction between the cylinder of wide hyphae and the subhymenium being rather
loose and composed of branches more or less at right angles to the axis of the head:
none of the hyphae agglutinated.
FIG. :u4. FIG. 215.

Twn-FIG. 214. Pistillaria setipes, X 10.
TllxT-FIG. 215. Pistillaria setipes,., x 10: variations to the·glabrous state.
The description which I have given is drawn from many collections which I made
near Cambridge, in England, between 1924 and 1926. It covers the descriptions of
all the species which I have listed as synonyms, with a few slight exceptions, and I am
unable to see how P. setipes in this sense, admittedly wide, can be subdivided into
smaller species. Indeed, I doubt whether P. pusilla, with smaller spores 4-6 X 2-3 µ,
is really separable. The basis of specification has been the height of the fruit-body,
the size and shape of the head, the hairiness of the stem, and the size of the spores.
The first four are such variable features that it is possible to find nearly-all the 'specific'
states on the same leaf, and there is every gradation between them in different collec
tions, though the extreme states, like T. Grevillei with long hairy stem and P. pusilla
with very short glabrous stem, seem distinct. In the spores there is so much over
lapping in different collections, that it is impossible to draw any specific line.
490 PISTILLARIA
I give in the accompanying list, Table XXXII, the 'diagnostic' data in case any
one can find a true species among them, though I think it would be better to describe
such as new rather than to hope to identify it among the existing confusion of ideas
or interpretations. The original descriptions of these fungi are macroscopic and so
vague that their names can be used scientifically only when they have been defined by
custom or when there is an indisputable type-specimen, i.e. the sole specimen or
-·
��:
,"1.
�
... _- ·.·
FIG. 216. FIG. 217.
TExT-FlG. 216. Pistilfaria setipes: spores of four collections, x 1,000.

TExT-FlG. 217. Pistillaria setipes: caulocystidia, X 1,000.
collection on which the species was based, not a subsequent specimen so identified
macroscopically as most 'Friesian specimens' are.
Among the data in Table XXXII, it seems that:
I. P. inaequalis Lasch has rather wide spores. Donk's measurements were taken
from the type-collection (Rab. 1930) and from one of Oudeman's named P. ovata.
The fruit-bodies in both were very small and nearly glabrous, yet there seems to be a
gradation to typical P. setipes as shown by my dwarf collections. For Patouillard's
interpretation of P. inaequalis, see P. albobrunnea.
2. T. hirsuta Cke. sensu Bourd. et Galz. may have longer narrow spores.
3. T. mucor Pat. may have very narrow spores.
4. T. longipes Karst. has an exceptionally stout stem.
PISTILLARIA 49 1
5. The remammg species and interpretations refer to one species which I have
called P. setipes Grev., because Greville first clearly described a small white Pistillaria
of this size, shape, and habitat with hairy stem: P. ovata and P. pusilla are older names,
but Fries did not describe them with hairy stems, as is certainly the normal state of
P. setipes, and I have followed Quelet and Patouillard in their interpretation of P.
pusilla, as the first mycologists to give it microscopic precision. Thus P. setipes
becomes the commonest white Pistillaria on the dead leaves of deciduous trees in
Europe: it is strange that it has not been recorded from N. America or Japan.'
Whether it grows on sticks, herbaceous stems, or grasses I do not know, though
Bourdot and Galzin give stems of Genista and grass-leaves for P. inaequalis, and
Killermann gives leaves, twigs, and bark for T. candida var. populina. P. typhuloides,
on dead stems of Epilobium, is very similar, but has small narrow spores, as in P.
pusilla, though more elongate, and perhaps more agglutinated or subgelatinous hyphae.
TABLE XXXII. SPECIFIC DATA FOR PISTILLARIA SETIPES
Height Head Stem Spores

Name i mm. l mm. mm. µ. I Stem hairiness· Authority: habitat
P. setipes -3·5/0·3-0·7 ...,,. 5/0·1-0·2 8·5-10/3-4 pilose or not Corner: on Salix
Corner: Salix,
5-11
" 1-7 -2·5/0·7 -4·5/0·15-0·25 8-10/J-4 sparsely hairy
Populus
" 2-3 0·8-1·5/0·2-0·4 0·7-1 ·2,'o· 1-0·2 6-8/3-3·5 " Corner: Alnus,
Popuius
Corner: Betula
.."
" 2-4
2-3
1·5-4
0·8-1·5/0·2
0·6-2·5'io·3-0·6
1-2·5/0·1-0·15
2·5-3/-
6-8/3'5-4
9-12/3·5-4·5
7-9·5/3-4·5
glabrous
pilos� much
I Corner: Salix
Donk: Alnus, Popu-
or little lus, other trees
T. carzdida 1-4 0·5-I'5/0·3-0·7 1-3/- 6-10/3-4 sparse or Bourd. et Gal::.:
glabrous
- -
rotting leaves
" " 6-9/3-4 " " Rea: Alnus, Salix
iI
Fraxinus,
-
Quercus
var. populina -5 1-2/- 7-8/3-4 glabrous Killerm.: Populus
T. pusil/a 0·5-3 - Bourd., Rea
P. ovata 4-7 - I -
indistinct 10/4
12/
"
" Quil•t
" - - - 7-8/3·5 " Massee, Schroeter
- - I - 8· 5-10/4-4· 5 " Sydow 1310 (Donk
P. i�;aequ.alis -1·5 0·8, 0·,t 0·8,'0·15 9'5·-I I/4-5 Dank: various leaves
1
sparse or
i
i esp. Populus
I
glabrous
- - Quilet: on paper
Bourd. et Galz.:
1··2 12/- puberulous
T. !iinula -:·5 0·3-o·s! 1·..:.-z;.- 10-14'3-4 bristly
T. mucor c-3
I II
glabrous
I Fagus, Ulmus
Patouillard: leaves
I I
I
9-10/2-3
Karsten; leaves
1/0·1 1-2/0·05
T. longipes -0·5 0·2/0·13 2/0·65 'I
P. cyli11dracea j 4-5 ! - I
: indistinct 6-10/3-5 " " Ainu,
I
Donk's MS. notes suggest that there is a distinct form or variety on leaves of Syringa
with relatively long, glabrous stalks (-2·6 X 1 mm.), small, pyriform head (1 X 0·3 mm.),
and rather short spores 6-8·5 x 4·5-5 µ. He records it from Holland (leg. C. J. Koning,
Oct. 1901: herb. Oud. as T. Grevillei), and Sweden (Romell, Fung. Exsc. Scand., as
P. inaequalis f. syringae). It seems very near P. ·citicola, the description of which is:
Head ovoid or obovoid, glabrous, white, obtuse: stem 1-2 times as long as the head,
cylindric or attenuate upward, glabrous: spores 6-7·5µ long, ellipsoid: on stems of
Vitis aestirnlis, U.S.A. (Ellenville, Ulster County).
Spegazzini records Typhula candida from the Argentine, on leaves of Salix Hum
boldtiana. He gives the spores as 6-9 x 2-3 µ, and the basidia as 15-18 X 5µ with 3
sterigmata (Sacc., Sy]!. Fung. 6, 749). Possibly the fungus referred by Bresadola to
Typhula Traillii is identical, thus:
9-12 mm. high, head 4-5 x 0·25 mm., cylindric-subfusiform, wholly fertile, often
1
T. Gre1:i/lei on petioles of Liquidambar, S. Car. (Berk., Grev. 2, 1873, 18): T. candida,
Argentine (Speg.). Both records need confirmation. See also P. viticola in the next paragraph.
492 PISTILLARIA
curved: stem 5-7 mm. long, filiform, with a few microscopic cuspidate caulocystidia:
spores 5-6 X 2·5 µ. oblong-obovate: basidia 15-20 X 6-7 µ., 4-spored: on bits of wood:
Brazil (Biumenau, leg. Moeller): Hedw. 1896, 291.
The narrow spores of both suggest a transition to P. pusilla, unless such are
separable with T. mucor: (T. Traillii is a tremellaceous fungus).
P. spathulata nom. nov.

Basinym: Clavaria spathulata Pk., Rept. N.Y. St. Mus. 27, 1875, 100, t. 2, f. 20-21:
Burt, Ann. Mo. Bot. Gdn. 9, 1922, 53, t. 9, f. 84: (non Chev. = Clavariadelphus
pistillaris).
-4 mm. high, simple, pale yellow, compressed, spathulate, tapering into a slender,
slightly furfuraceous stem.
On dead branch(.;s of Carva alba (Juglandaceae): U.S.A. (Greenbush, N.Y.).
Spores?
Burt says the clubs emerge from crevices in the bark, sometimes 2-3 together.
Compare P. fuegiana and P. paradoxa.
P. subfasciculata (E. et E.) comb. nov.

Basinym: Typhula subfasciculata E. et E., Proc. Am. Phil. Soc. 1890, 219.
1-2 mm. high, subfasciculate, cylindric, curved, white-mealy, apex contracted,
acute.
On dead bark of Ulmus: Canada (London).
Spores ? 3 X 2 µ..
Basidia 15-25 X 6-8 µ..
? Mucronella or Defiexula ulmi. Compare Clavaria quercicola (under Pistillaria
albobrunnea).
P. subpellucida B. et C.
White, subpellucid, linear; slightly clavate, base subincrassate, 1-2 mm. high.
On fern-stems: Cuba: (on living fern-stems in a conservatory, Montevideo: Speg.).
Under the name P. quisquiliaris (which grows on petioles of Pteris in Europe),
Donk found in the Paris herbarium a collection of Wright's from Cuba which
probably belongs here. Donk's MS. notes are: 2-4 mm. high: head o·8 mm. wide,
clavate, obtuse: stem 0·3 mm. wide, glabrous, about as long as the head: spores
5·5-8 x 2·5-3·5 µ.: basidia 20-32 x 5-7·5 µ.: hyphae 3-8·5 µ. wide.
It greatly resembles P. pusilla.
P. subuncialis sp. nov.

Text-Fig. 218.
5-15 mm. high, 1 mm. wide, cylindric, rarely slightly clavate,
rarely forked, obtuse, glabrous, pallid dingy yellowish, the stem
Q
indistinct, base inserted, apex fertile.
On dead stems of Juncus: England (Dernford Fen, Wicken
Fen, Cambridgeshire, Nov. 1925, 1927).
'
"
. Spores 10_-13 X 5-6 µ., white, elongate, pip-shaped, aguttate.
Basidia 20-30 X 7-10 µ., mostly rather truncate below, some
times prolonged into a narrow base: sterigmata 4, 7-1o µ. long.
TExT-FIG. 218. Cystidia -35 X 17 µ., as sterile, inflate.cl basidia, ventricose,
Pistillaria subuncialis: blunt, scarcely projecting, if at all.
cystidia and a basidium, This species closely resembles P. uncialis but has much
><500. larger spores and distinct cystidioles.
PISTILLARIA 493
P. syringae Fckl.
Enum. 100, t. 1, f. 24: Exsic. 891, cum. diagn.: Symb. Myc. 1869, 31, no. 6.
2-3 mm. high, scarlet, linear, blunt, glabrous, often curved, a little thickened at the
base: head -1·6 X 0·23 mm.:.stem 0·27 X 0·21 mm., very short, jonquil-yellow.
On dead leaves of Syringa: Germany, France.
Spores 5 ·5-8 X 5-6 µ., broadly ellipsoid, with a small, acute apiculus (Donk).
Basidia 20-35 X 5·5-7·5 µ.: sterigmata 7.-4, 4·5 fL long (Donk).
Hyphae 3·5-5 fL wide (Donk).
I have amended the description of this very striking, but little known, species from
Donk's MS. notes on the type-distribution (herb. Oud.). In colour, size, and shape of
the fruit-body, and in the form of the spores, it seems related to P. micans.
P. tenera nom. nov.

Synonym: Clavaria tenuis Schw., Am. Phil. Soc. Trans. 4, 1832, 182: Burt, Ann.
Mo. Bot. Gdn. 7, 1922, 49, t. 9, f. 75: [baud Pistillaria tenuis (Fr.)].
? Clavaria muscorum Karst., Hedw. 1889.
6 mm. high, mostly simple, occasionally forked at the apex, white or pallid, slender,
tender.
Scattered among moss on dead wood: U.S.A. (N.Y.).
(Spores? 3-3·5µ, subglobose, smooth; doubtful, Burt.)
Hyphae 3-3·5 fL wide, long-celled.
Burt suggests this may be Eocronartium. It may also be Cerate!lopsis, but is prob
ably unrecognizable.
The description of C. muscorum is:
6-8 X 0·2-0·3 mm., scattered, simple, filiform, attenuate upwards or clavate and
obtuse, often fusiform, stem scarcely distinct, whitish: on living Jungermannias,
Minas Geraes, Brazil.
P. tenuis (Fr.) comb. nov.

Basinym: Typhula tenuis Fr., .Syst. Myc. 1, 1821, 495: (Sow., Engl. Fungi, t. 386,
f. 5, as Clavaria tenuis).
4-6 mm. high, bistre then fuscous-black: head thickened, pyriform, smooth: stem
filiform.
Gregarious on rotten wood: Gt. Britain, Switzerland.
Doubtfully a basidiomycete.
P. Todei (Fr.) comb. nov.

Basinym: Typhula Todei Fr., Syst. Myc. 1, 1821, 494: Bourd. et Gatz., Hym. Fr.
1928.
Synonyms: Clavaria chordostyla Pers., Myc. Eur. 1, 1822, 189.
Clavaria filicina Pers., ibid. 190.
Clavaria Todei (Fr.) Karst., Finl. Nat. o. Folk 37, 1882, 189.
Head 2-3 X 0·3-0·5 mm., linear then fusiform, obtuse, pale or clear yellow, or
slightly rufescent: stem 5-8 mm., very slender, capillary, white, shortly pilose towards
the often subbulbous base: (-2·5 cm. long, overall, fide Sacc.).
On fern-remains: Europe.
Spores 8-9 X 3-4µ, white, smooth, often subarcuate, aguttate.
Basidia 18-24 X 5-7µ, with 2-4 sterigmata.
Hyphae of stem 3-6 µ. wide, with irregular swellings -24 µ. wide, without clamps.
Bourdot and Ga!zin, who supply the microscopical details, say that this is not
uncommon in France. It seems very close to P. setipes and P. typhuloides. Fries said
that during 60 years he never saw it with a sclerotium: nevertheless, by analogy with
Typhula quisquiliaris, it may have a sclerotium deeply immersed in the fern-tissue.
494 PISTILLARIA
Donk (MS.) thinks it may be T. quisquiliaris, but the spores (given by Bourd.) are
smaller.
P. trichomorpha (Schw.) comb. nov.

Basinym: Clavaria trichomorpha Schw., Naturf. Ges. Leipz. 1, 1822, 112.
-12 mm. high, simple, densely clustered, white, subpellucid, attenuate at both
ends: on �ems of Zea: U.S.A. (Schw., Car., No. 1699).
No specimen is known to exist. Probably a Pterula.
P. typhicola Bourd. et Galz.

Hym. Fr. 1928, 139: (Typhula gracilis B. et Desm., sensu Bourd. et Galz., Bull.
Soc. Myc. Fr., non B. et Desm. = Pterula gracilis).
Wholly white, subcylindric, straight or slightly fl.exuous: head 0·9-1 · 5 mm. long,
fertile over the apex: stem 1·5-2·5 mm. long, softly hispid, nearly as wide as the head.
On dead remains of Typhula and J uncus: France.
Spores 9·5-12•5X3-3·5 µ, white, elongate-ellipsoid (Donk).
Basidia 18-29X6-10µ: sterigmata 2, 9X3·5µ.
Hyphae 2-4µ wide, slightly thick-walled, incrusted with crystals.
Caulocystidia 20-200X1-3µ, flexuous, hypha-like.
This belongs to the P. pusilla-P. setipes alliance. Donk (MS.) has examined the type
collection (herb. Bourd.) and found two kinds of spores on the fruit-bodies, that
given above being the correct size and that given by Bourdot and Galzin (7-9X3·5-4µ)
being extraneous.
P. typhuloides (Pk.) Burt

Ann. Mo. Bot. Gdn. 7, 1922, 69, t. 11, f. 108: Lundell, Sv. Svamp. 5-6, 1936, 27.
Basinym: Clavaria typhuloides Pk., Rept. N.Y. St. Mus. 30, 1878, 49, t. 2, f. 12-14.
Synonyms: Typhula falcata Karst., Hedw. 1881; Act. Soc. Sci. Fenn. 15, 1888,
180, t. 20.
Clavaria subfalcata Karst., Finl. Nat. o. Folk 37, 1882, 180.
Pistillaria subfalcata Karst., ibid. 48, 1889, 382.
Pistillaria puberula var. viscidula Karst., Med. Soc. Faun. FI. Fenn. 13, 1886, 161
(teste Lundell).
3-5 mm. high, scattered or gregarious, white, yellowish or pinkish yellow with age
(Karst.): simple or 1-2 branches from the base: head 1·5-2 mm. wide, linear and sub
acute then clavate, obtuse, solid: stem shorter or longer than the head, puberulous,
pellucid: flesh rather tough, subdiaphanous, subgelatinous when moist, drying horny
(Lundell), viscid (Karst.): no sclerotium.
On dead stems of Epilobium angustifolium: Europe (Sweden, common; Finland).:
U.S.A. (Adirondack): Canada (Manit.).
Spores 6-8(-1o)X2·5-3(-3·5)µ (Lundell), 5-7·5µ long (Peck), 5...:6x2·5µ (Burt),
6-8X3-4µ (Karst.), white, oblong-ellipsoid, smooth, 1-2 guttulate (Lundell).
With very abundant calcium oxalate crystals among the hyphae of the whole fruit
body (Karst.).
This is probably a common species wherever the Epilobium occurs, but, as Lundell
suggests, it is perhaps mistaken for Helotium cyathoideum. The viscid, subgelatinous
texture, mentioned by all authors, must be caused by the modification of the hyphal
wall, as incipient agglutination extending, perhaps, into the head as well as on the
surface of the stem. The species, therefore, may be alliect with Pistillina hyalina,
though there is only-the habit.at and this slight difference in texture to distinguish it
from Pistillaria setipes. On the whole i.t seems to belong to the P. uncialis-subuncialis
group, as the most reduced member.
PISTILLARIA 495
P. uliginosa Crouan
F. Finistere 1867, 60.
Club cylindric or clavate, purplish: stem short.
Spores spherical.
On various dead herbaceous stems: France, Austria.
var. albo-lutea Keissler
Beih. Bot. Centralbl. Abt. 2, 29, 1912, 437.
Yellowish white.
On dried stems and leaves of Aegopodium podagraria: Yugoslavia (Camiolia).
This species may be the same as P. ampelina. Keissler remarks that other fruit
bodies of this species found by him on Umbelliferae had purple clubs(? heads).
P. uncialis (Grev.) Cost. et Dufour
Basinym: Clavaria uncialis Grev., Scot. Crypt. FI. 2, 1824, t. 98.
= Ty_phula-quis
Synonyms: C. obtusa Pers., Mys. Eur. 1, 1822, 190 (pr. p.:. alt. p.
quiliaris).
? Typhula gracillima White in 13. et Br., Ann. Mag. Nat. Hist., Ser.. 5, 1, 1878,
No. 1699: Grev. 6, 1878, 125.
Ceratella uncialis (Grev.) Quel., FI. Myc. 1888.
Gliocoryne uncialis (Grev.) Maire, Bull. Soc. Bot. Fr. 1909, 120.
Text-Figs. 37, 219.
o·5-2· 5 cm. high, white then yellowish or pale ochraceousfrom below upward, solitary or
gregarious: head 4-20 :l< 0·2-1·S mm., cylindric to subclavate, subacute then blunt: stem
2-7 X o· 15-0· 5 mm., glabrous, more or less distinct: no sclerotium: slightly tough, solid.
On dead herbaceous stems, espectally Umbelliferae (Grev.), Mulgedium Plumieri
(Maire), Juncus-stems (Comer), or on sticks (Comer): Gt. Britain, France.
Spores 4-7 X 2-3µ, white, smooth, ovoid-ellipsoid to ellipsoid, blunt,. obliquely
apiculate, aguttate or 1-plurigu ttulate (Maire).
Basidia 30-35 X 4-5 µ(Maire), 20-28 X 4-5µ(Corner): sterigmata 2-4, .r5-4µ long.
Cystidia none.
Caulocystidia none, the stem microscopically glabrous or with a few -blunt, slightly
divergent, scattered hyphal ends as sterile basidia.
Hymenium not thickening, covering the whole head: subhymenium c. 10µ thick,
composed of narrow, short-celled hyphae 2-3 µ wide, arising directly from the longi-
tudinal hyphae.
This species has been regarded as doubtful. Maire is the only author who has
given adequate microscopic details. He discovered the agglutinated superficial hyphae
of the stem and founded the genus Gliocoryne on P. uncialis, but the feature is
characteristic, in some.degree, of all species of Typhu!a and Pistillaria. Donk says,
however, that the fungus identified by Maire with P. uncialis is the 'sclerotium-free·
form of Typhula sclerotioides and not P. uncialis. I do not find an explanation, either,
that anyone has proved T. sclerotioides to have a 'sclerotium-free' state, or how the
clongate-cylindric head of Maire's fungus(5-15 X 1 mm.) agrees with the short head
(1-5 mm. long) of T. sclerotioides. In contrast, I have made two collections of Pistillaria
which agree well with the descriptions of Greville and Maire in the elongate yellowing
head and short glabrous stem, without sclerotium, which are the chief macroscopic
features of P. uncialis, and their spores agree with those of Maire's fungus. But the
collections differ in microscopic structure, so that I describe them separately below,
and in detail, for they may be two species macroscopically similar.
Regarding T. gracillima the description is:
\Vhite, 15-20 mm. high; stem very slender, curved, smooth: head elongate: on dead
herbaceous stems: Scotland (Perthshire). Massee reported that on an authentic
specimen in Berkeley's herbarium at Kew he could find no trace of hymenial surface
PISTILLARIA
(Brit. Fung. Fl. 1, 1892, 90). The description passes well for Greville's fungus
macrosopically.
P. uncialis: on a dead stick, Wicken Fen, Cambridgeshire, England, 24.10.26:
Text-Fig. 219:
1 cm. high, pale ochraceous, apex whitish: head -7 X 0·2-0·25 mm., cylindric,
subacute (? immature): stem 3 X o· 15-0·2 mm., glabrous: no sclerotium.
Spores 4-5 X 2·5-3 µ., broadly ellipsoid, aguttate.
Basidia 20-28 X 4-5µ.: 4 sterigmata, 3·5-4µ. long.
Subhymenium 10µ. thick, hyphae 2-3µ. wide, interwoven.
=--=-=- ··.:......=
TExT-FIG. 219. Pistillaria uncialis: section of the hymenium, showing

the fractured crystals among the longitudinal hyphae: x 1,000: (collection
from Wicken Fen, England, 24.10.26).
Hyphae 4-7(-12) µ. wide, little inflated, with slightly mucilaginous walls, agglutinated
on the surface of the stem, cells 40-300 µ. long, with clamps at all septa: crystals
10-25 X 2--{) µ., white, oblong-cylindric or with truncate ends, often fractured trans
versely, set longitudinally between the longitudinal hyphae of the head and stem,
very abundant.
P. uncialis: on dead Juncus-stems, Wicken Fen, Cambridgeshire, England, Nov.
1926:
Differs in the short wider head, larger and wider stem, wider hyphae, absence of
clamps and crystals, and in the slightly longer spores.
-12 mm. high, white, then pale yellowish, scattered: head 1-3 X 0'.5 mm.. cylindric
subacute (? immature): stem 5-9-x 0·2-0·4 mm., glabrous: no sclerotium.
Spores 5--{)•5 X 2·5-3 µ., rather narrowly pip-shaped or ellipsoid, aguttate.
Hyphae 3-20µ. wide, cells 50-300µ. long, not mucilaginous but agglutinated and
PISTILLARIA 497
slightly thick-walled on the surface of the stem, no clamps, no crystals: caulocystidia
none.
P. uncialis sensu Maire (as Gliocoryne uncialis):
Head 5-15X I mm.: stem glabrous.
Spores 6-']X2-3µ,, oblong-ellipsoid, 1-pluriguttulate.
Basidia 30-35 x 4-5µ,: 2-4 sterigmata.
Hyphae with thickened gelatinous walls.
P. uncialis sensu Greville (measurements from the illustration):
-2·5 cm. high, white, yellowish with age, cylindric, gradually attenuate downwards,
scattered or gregarious: head 4-20 x 1-1·5 mm.: stem 2-7 x 0·3--0·5 mm.
(Spores 5X3µ,; Mass.)
Whether there is more than one species among these collections can be proved only
from further mate.rial, but the differences may well be varietal unless for the presence
of crystals. The species, or group of species, differs from the.P. setipes-alliance in the
elongate-cylindric head, evidently with considerable apical growth, in the glabrous
stem, and the narrower spores. P. oleae seems to be a similar fungus, with slightly
larger spores and epiphyllous habitat. P. subuncialis, of this alliance, has much larger
spores.
Species Excludenda
P. Johnsonii Mass.
.t(ew Bull. 1901, 165.
Fungus imperfectus: synnemata 1-2 mm. high, very slender, white, composed of
parallel hyphae 2-3µ, wide (without clamps!), conidiophores forming a very dense
'hymenium' of rather barrel-shaped cells, 8-g x 4 µ,, with a single sterigma 3-4µ, long:
conidia 3X2µ,, smooth, colourless, thin-walled, apparently home in rows: on a dead
herbaceous stem: Gold Coa11t (type in herb. Kew: Comer).
PISTILLINA Quel.
Ass. Fr. 1880, 671.
Fruit-body minute, erect, stipitate, without sclerotium: head discoid to solid cyathi
form or subglobose (? always bounded by a narrow sterile margin), the sterile sides
not reflexed: stem slender, sterile, with or without caulocystidia: hymenium covering
only the discoid or convex surface of the head.
On dead plant-remains: north temperate, 4 spp.
Basidia (? long, as in P. hyalina).
Cystidia none.
Hyphae monornitic, clamped (? in all cases), not secondarily septate, slightly inflat
ing or not at all, with firmly submucilaginous walls, the superficial hyphae of the
stem and outside of the head agglutinated.
Type-species: P. hyalina.
KEY TO THE SPECIES

Sp. 12-14 X4-5 µ,: rufous, coronately toothed round the margin: on
twigs: Mediterranean P. ,rubra
Sp. smaller: not toothed round the margin
Lignicolous: c. 0·1 mm. high, glabrous: sp. 5-gX3·5-4·5µ,:
U.S.A. . P. Thaxteri
On grasses: sp. 7-10X3·5-4·5µ,: Europe
White: stem finely pilose: margin of hymenium glabrous P. hyalina
Brown: stem villous: margin of hymenium ciliate P. brunneola
5119 K k
49� PISTILLI NA
P. brunneola Pat.
Tab. An. 1887, 574.
Synonym: Pistillaria brunneola (Pat.) Sacc., Syll. Fung. 6, 1888, 759.
2 mm. high: head clear brown, diaphanous, convex above, flattened and paler
beneath, margin ciliate: stem erect, villous, minutely bulbous, brown.
Gregarious on dead grass-leaves: France (Jura).
Spores 8-9 X 3 µ, white, smooth, ovoid.
Basidia 4-spored.
The description suggests a species allied with P. hyalina, but the ciliate margin,
villous stem, and brown colour also suggest Cyphella.
P. hyalina Quel.
Ass. Fr. 1880, 671, t. 8, f. 12: Rea, Tr. Br. Myc. Soc. 12, 1927, 227.
Text-Figs. 220-2.
1-2 mm. high, white, diaphanous, like a minute, slender-stalked solid cup (or a narrowly
stipitate Helotium): erect: head 300-500 µ wide, solid, cup-shaped, pulvinate or
TExT-FIG. 22 I. P.istillina hyalina: longitudinal section of the margin of the
fruit-body, x 1,000.
500 PISTILLINA
subglobose: stem 1-1·5 mm. long, 70-<;5µ, wide above, 100-130µ, wide at the slightly
dilated base, slender, expanding gradually or rather suddenly into the head, finely
pilose with rather long hairs: without sclerotium.
On dead grass-leaves: France, Gt. Britain (Wicken Fen, Cambridgeshire, 15.2.25).
Spores 7-10 X 3 · 5-4·5µ,, white, smooth, subcylindric, slightly curved, aguttate, thin
walled.
Basidia 40-60 X 6-7µ,, clavate with long narrow base 1 · 5-2µ, wide, clamped:
sterigmata 4, 6--'7µ, long.
Cystidia none.
TEXT-FIG. 222. Pistillina hyalina: base of the stem (the outer hyphac
agglutinated), and a caulocystidial hair (bottom right): >'. 500.
Hymenium not thickening, covering the more or less convex upperside of the head
as a disk, not extending on to the sides : subhymenium well developed, the hyphae
1·5-2·5µ, wide, interwoven, clamped.
Caulocystidia -150 X 1-2µ,, cylindric-filiform, thin-walled, aseptate, abundant,
shorter upward on the stem.
Hyphae 1·5-5µ, wide, scarcely inflating, with thin submucilaginous walls, most
septa with clamps, a few without: hyphae longitudinal, the cells 50-200µ, long in the
stem, 20-150µ, in the head and shorter towards the subhymenium, the central hyphae
of the head and stem 1 ·5-3µ, wide, the outer hyphae 2-5µ, wide, the superficial hyphal
PISTILLINA 501
with slightly, but distinctly, agglutinated walls: margin of the head round the
hymenium thin and entire: with numerous crystals of calcium oxalate on the surface
of the head and stem and in the tissue.
This seems to be a rare fungus, for my collection from Wicken Fen is the second
to be recorded. It fruits at a time when the mycologist has little inclination to search
among wet grass, which may contribute to an apparent rareness, yet I looked for it in
the same place during the next 3 years (1926-8) without success. My description
differs from Quelet's only in the spores and I think his remarks must be erroneous:
he gives the spores as 10-12µ pruniform-elongate, finely aculeolate, 1-2 guttulate.
The texture of the fruit-body is rather tough because of the submucilaginous hyphal
walls and the agglutinated surface-hyphae: thus, they resemble in form, texture, and
colour a Helotium, as H. cyathoideum.
[P. paradoxa (B. et C.) Cke., Grev. 19, 1890, 2: = nomen nudum.
Basinym: Crinula paradoxa B. et C., Rav. Fung. Car. Exs. 3, 35: Rav. Fungi Am.
399: Thuemen, Myc. Univ. 208: Ellis, N. Am. Fung. 23: vel Pistillaria paradoxa
B. et C.: = Cronartium quercuum Schroet., vide Seymour, Host Ind. Fungi N. Am.
1929, 250.]
P. rubra Fautr. et Ferry
Rev. Myc. 1893, 92.
Synonym: Pistillaria rubra (Fautr. et Ferry) Sacc., Syll. 11, 1895, 142.
Head hemispheric, rufous, covered by the hymenium, coronately toothed round the
margin: stem short, solid, dilating gradually into the head.
On twigs of Sarothamnus scoparius (Leguminosae): France (Cote d'Or).
Spores 12-14X4-5µ, white, smooth, pruniform-elongate.
P. Thaxteri (Burt) comb. nov.
Basinym: Pistillaria Thaxteri Burt, Ann. Mo. Bot. Gdn. 3, 1916, 406.
100-uoµ high, gregarious, subcapitate, erect, white then tawny: head 50-110µ
wide, 40-50µ high, almost hemispheric, the hymenium limited to the convex surface:
stem 60X20-50(-80)µ, short and dilating obconically into the head, glabrous; with
out sclerotium.
On rotten wood: U.S.A. (Westhaven, Conn.; Thaxter, 7.u.1888).
Spores 5-9X3·5-4·5 µ, white, smooth, ellipsoid, flattened adaxially, acute at the
base.
Basidia 13-17X4-4·5 µ, subclavate: sterigmata 4.
Cystidia none.
Hyphae 1 · 5-2 µ wide, thin-walled, without clamps, longitudinal in the stem,
divergent in the head.
This seems to be closely allied with P. hyalina and to differ in its minute size,
absence of caulocystidia, and, apparently, its short basidia, though their slender stalks
may have been overlooked. It is, to judge from Burt's figure, a rudimentary primordial
shaft with the hymenium limited to the apex, as in P. hyalina, and perhaps bounded
in the same way by a thin margin of the outer, sterile hyphal tips. The hyphae seem
not to inflate. It is, as Burt says, practically the smallest known Homobasidiomycete.
PTERULA Fr.
Syst. Orb. Veg. 1825, 90: Linnea 5, 1830, 531: Lloyd, C. G., The Genus Pterula,
Myc. Writ. 5, 1919, 863.
Synonyms: Penicillaria Chev., FI. Gen. Paris, 1, 1826, 111.
Phaeopterula P. Henn., Hedw. 43, 1904, 175.
Fruit-body varying much branched and bushy to simple: stem generally distinct,
short, slender, sometimes with a slight byssoid or strigose disk: branches 0·3-1 mm.
wide, slender, terete, attenuate to finely subulate or filiform tips 10-50µ wide (even
502 PTERULA
reduced to a single hypha), with flattened-dichotomous branching in alternating planes
or the main branches apparently polychotomous (through condensed dichotomy), or
adventitious in subsimple species, multaxial with equal dichotomy or monaxial with
pseudo-verticillate branching through overtopping or adventitious branching, often
connate from crowding: hymenium waxy, generally absent from the stem and uppersides
of the main branches: flesh tough, often drying fuscous brown, horny and cartilaginous
with twisted branches.
On humus, wood, and various plant remains: 52 spp., temperate and, mostly,
tropical.
Spores white, smooth (asperulate in P. grandis), ellipsoid to subglobose, aguttate (or
with a minute guttula).
Basidia small, clavate: sterigmata 2-4.
Cystidia present or absent: no gloeocystidia.
Hymenium thickening, or not in a few simple species, the old basidia collapsing:
subhymenium composed of short-celled uninflated hyphae with submucilaginous
walls.
Caulocystidia often present.
Sterile hymenium on the stem and uppersides of the branches cons1stmg of a
rudimentary hymenium, not or little thickened, often with caulocystidia on the stem
but few or none on the branches: modified into a 'pavement epithelium' of hyphae
with agglutinated brownish walls in several subsimple epiphyllous species.
Hyphae dimitic: skeletal hyphae of unlimited growth, thick-walled, colourless, or with
pale brownish or yellowish walls, aseptate, smooth, mostly unbranched, occasionally
branched into 2-5 skeletals apically or producing a lateral generative hyphae: genera
tive hyphae thin-walled, uninflated, septate, generally with clamps, a few species without,
someti.mes branched from the clamp, occasionally with thick-walled, intercalary,
skeletal segments.
Type-species: P. subulata Fr.
Beyond more closely defining the genus on the hyphal structure, excluding several
species, amplifying a few descriptions, and adding seven new species, I have done
little to ascertain the identity of many of the old species. Most of these have been
described from dried specimens, and, in drying, the fruit-bodies become so shrivelled,
twisted, ·and discoloured as to be unrecognizable in their delicate living counterparts.
Even the European and N. American species are so little known, in manner of branch
ing and microscopic detail, that it is impossible to decide whether any of the more
numerous tropical ones are identical, e.g. the record of P. multifida from Java and the
resemblance between P. penicellata and P. verticillata. I have come to the conclusion
that, even if one could examine all the type-specimens, it would still be impossible
to set specific limits. Therefore, I recommend mycologists able to study fresh (or
adequately preserved and annoted, or illustrated) material, to describe as new and as
fully as possible what cannot be identified with certainty. I take a species with white
fruit-bodies to be different from one with yellow, brown, or vinaceous fruit-bodies,
until it is shown that they vary into each other: and, thus, I have described P. verti
cillata, P. brunneola, and P. vinacea as new. Further collections may show them to
be varieties of one species which may already have been described several times. But
only by such means can future monographers be supplied with d:ita for taxonomic
revision.
It should be noted that the sparing of Pterulas may be nocturnal, e.g. P. debilis,
P. typhuloides (and Defiexula fascicularis), so that it is advisable to obtain spore-prints
overnight for future study: specimens dried or preserved in alcohol-formalin in the
day-time may be sterile, and hence the difficulty in determining the specific limits of
existing herbarium material. At present, Pterula is a genus for field-work.
In the specific key it is scarcely possible to indicate more than the barest means of
identifying most species.
PTERULA 503
KEY TO THE SPECIES OF PTERULA�
(Doubtful species, see pp. 534-6)
Fruit-bodies bushy, much branched: in humus or on wood . A
Fruit-bodies simple or sparingly and laxly branched, generally
small or minute: on plant-remains B
A. Bushy Pterulas
Sp. 10-12 X 5-7 µ,: pale ochraceous fruit-bodies: trop. Africa and
Asia (? Amer.) . P. capillaris
Sp. 8-10 x 5--'7 µ,: whitish cinereous, then yellowish, sparsely
branched but densely tufted: on dead wood: Europe P. subulata
(Sp. u-15 X6-7·5µ,: sparingly branched . see Pterulicium)
Sp. smaller (or unknown)
Monopodial with more or less verticillate branching
Temp. N. Amer., in humus P. penicellata
Trop.
Stem tomentose or strigose-hirsute
Viol aceous: on the ground: Java P. hirsuta
On rotten wood: Amazon . P. juruensis
Pale drab flesh colour, yellowish drab, or fawn-drab: sp.
6-7 X 3-3 ·5 µ,: in humus: Malaya P. verticillata
Not strigose or tomentose see P. taxiformis
Multaxial with equal dichotomies, or only a few branches mono
podial
N. temp.
-5 cm. high, white, yellowish, greyish, lilaceous, then
brownish, densely branched and tufted: sp. 5-7 X 3-4µ,:
in coniferous humus: Europe P. multifida
As P. multifida, but branches very dense and compact: on
the ground: Europe, U.S.A. P. densissima
·-4 cm. high, light grey-brown: branches rather few, divari-
cate: sp. 7-8x3·5--4·5µ,: in humus: U.S.A. P. plumosa
-1 ·5 cm. high: on coniferous needles: France P. abietis
Trop. or S. temp.
New World
As P. multifida: sp. 4-4·5 X 3·5-4µ,: on bark: Brazil P. Uleana
Branchlets secund: ? pale rufescent: on rotten wood:
Brazil . P. secundiramea
Cinereous, moss-like, -2 cm. high: on trunks: Brazil P. adustipes
Old World
Sp. 3-4x 1 µ,: yellowish ochraceous, with extensive
mycelial (7lt: on sticks: Java . P. Janseniana
Sp. 4-5 µ,, subglobose, asperulate: -9 cm. high, pale
vellowish brown: on dead stems: Africa, New Guinea P. grandis
1
The colours gi\'en in the Key refer, so far as can be determined, to living specimens.
PTERULA
Sp. 4-5µ, subglobose, smooth: -12 cm. high, branches
rather coarse, axils flattened: trop. Africa P. Bresadoleana
Sp. ellipsoid, smooth (see below)
-3 cm. high: sp. 5-6X3·5-4·5µ: on bark: Timor P. tirrwrensis
Larger: sp. 6-7X3-3·5 µ: terrestrial: -9X12 cm., very
bushy, main branches decumbent, pale fawn-drab,
greyish at the base: Malaya P. brunneola
-7x 6 cm. vinaceous-drab, darker below, pale yellowish
flesh-colour upward, some branches monopodial:
Malaya P. vinacea
(Fuliginous, stem and branches rather stout: on the
ground: New Guinea see P. grandis)
Sp. unknown
Madura: highly branched, axils dilated: on the ground P. dilatata
Mauritius: branches not many P. Commersonii
W. Africa: on rotten wood P. togoensis
S. Africa: �2·5 cm. high
On rotten wood: white P. filaris
On the ground: whitish or rusty P. setacea
B. Simple'or sparingly branched Pterulas'

Sp. subglobose
Branches racemose, apparently monopodial, lax
-2·5 cm. high, pale greyish: sp. 4-5X3-4µ: ? on bark or
wood: S. Amer., France P. taxiformis
Caespitose and subsimple, or simple and minute
Sp. 7-8µ: -14 mm. high, dichotomous, base strigose-hairy:
on wood: Tahiti . P. tahitensis
Sp. 5-7µ
-3·5 ·cm.X o·15--0·2 mm., simple, or rarely 1 branch, base
white byssoid: on palm-sheaths: Congo P. Winkleriana
-3 mm. Xo·12 mm., simple: on wood: Congo . P. Vanderystii
Sp. 3·5-4µ
-s mm. high: on herbaceous stems·: Brazil P. falcatula
-30 mm. high, base thickened, fuscous, subsclerotioid: on
dead leaves: Brazil . P. tenerrima
Sp. ellipsoid (see below)
Sp. 18-2ox 10-12µ: 1 mm. high: on branches: Ecuador P. macrospora
Sp. smaller
On old fruits of Pandanus: -3 cm. high, sparingly branched:
sp. 9-10 X 5-6 µ: Philippine Isl. P. fructico/,a
On dead leaves, stems, twigs, rarely wood
Hymenium with cystidia: base of stem not sclerotioid
-s cm. high, irregularly branched, flaccid, white then pale
brownish below: sp. 7-9X3·5-4µ: on Juncus-stems:
England P. debilis
' Compare Ceratellopsis.
PTERULA 505
-2·5 cm. high: sp. 9X4µ: on leaves: Tasmania P. phyllcphila
Smaller, simple
Temperate: -II mm. high, white: sp. 9-16 X4-7µ:
2-spored:on herbaceous stems and leaves: Europe . P. gracilis
Trop.: -18 mm. high, white then pale ye!lowish, stem
becoming fuscous: sp. 9-13 X4-5µ: 4-spored: with
caulocystidia: on dead leaves: Malaya P. epiphylla
(Sp. II-15x6-7·5µ: see Pterulicium, trop.)
Hymenium without cystidia: base of stem often sclerotioid
Without caulocystidia
-8 cm. high, simple or with 1-3 branches, white, then
pale ochraceous or brownish: sp. 9-11·5 X5-8µ: on
dead leaves of Oryza: U.S.A. P. oryzae
(Sp. u-15x6--7·5µ: on wood, palms, bamboos Pterulicium)
-20 mm. high, mostly simple
Stem becoming fuscous or blackish, levigate with
sterile epithelium
Sp. 9-11X5µ: -14 nun. high: on dead leaves:
U.S.A., S. Amer. P. tenuissi.mo
Sp. 6-8X2·5-4µ: -20 (?'30) mm. high: base of
stem sclerotioid: on leaves: S. Amer. · P. tenerrima
Stem not becoming fuscous
Sp. 15-17x7µ:-3·5 mm. high: on wood: Holland P. rigida
Sp. smaller
-1·5 mm. high: sp. 7-1ox5-{)µ: on twigs:
Guadeloupe P. nana
-4 mm. high, yellowish: sp. 9-10x4µ: on
trunks: China P. vitellina
Larger, white: on leaves
Sp. 8-1ox3·5-4·5µ: Brazil P. Gordius
Sp. 6--7 X4-5µ: Cameroons P. penniseti
Sp. 9X4µ: -2·5 cm. high: on leaves:
Tasmania P. phyllophila
With caulocystidia: stem levigate with a fuscous epithelial
layer: Malaya
Sp. 10-13 X4-5µ: epithelial layer on stem 15-30µ
. thick: -zo mm. high, simple: on twigs, herbaceous
stems P. subtyphuloidet
Sp. 8-10 X3·5-5 µ: epithelial layer on stem 5-10µ thick
-35 mm. Xo·5-1·5 mm., simple or with 1-2 branches,
white to pale ochraceous: on dead leaves, in-
florescences, &c. P. typhuloides
-11 mm. Xo·1-0·25 mm.: on dead grass stems·and
leaves var. minor
Sp. unknown
Trop. and S. Amer.
Cuba, on bark: -20 mm. high, white, base tomentose-affixed:
gen. hyphae without clamps . P. subulaeformis
506 PTERULA
Brazil, Guadeloupe
-15 mm. high, caespitose-simple: on wood: Brazil P. subsimplex
-12 mm. high, sparsely verticillately branched, attached by
mycelial fibrils: on herbaceous remains: Brazil P. pusilla
-5 mm. high, caespitose-simple, pale flesh colour: on wood P. incarnata
-8 mm. high, with a few simple branches, whitish, Guade-
loupe P. laxa
Congo
(-15 mm. high, simple, corticolous, with thin incrusting
mycelium and strands Clavaria
Lecomtei)
Borneo
-8 mm., simple or forked once or twice, attached to a straw
coloured mycelial patch, densely gregarious: on dead her-
baceous stems P. fulvescens
? origin
-15 mm. high, 1-3 times dichotomous (? monopodial): on
wood P. exserta
[Dubious species (p. 534).]
P. abietis Lloyd
Myc. Notes 75, 1925, 1357, f. 3200.
-1 ·5 cm. high, densely caespitose: stem little distinct: branching from the base,
fimbriate above, drying fuscous brownish with paler tips.
On Abies-needles: France.
Without microscopic details the position of this fungus canr.ot be determined.
? Lentaria.
P. adustipes (Speg.) comb. nov.
Basinym: Clavaria adustipls Speg., Fungi Guar. Png. 1, 1883, n. 87: Sacc., Sy!!.
Fung. 6, 1888, 715.
-2 cm. high, cinereous, slender, moss-like tufts, rather rigid and firm: stem 4-10 x
3-4 mm., solitary or fasciculate-connate, fuscous-black, branched apically and on the
sides: branches 5-10 mm. long, sparse, erect or arcuate ascending, simple or with
1-2 branches, elongate, slender, long-attenuate, acute.
On dead mossy trunks in the forest: S, America (near Guarapi).
Compare P. secundiramea and P. Uleana.
P. Bresadoleana P. Henn.
Engl. Jahrb. 17, 1893, 22: Killerm., Nat. Pfl. Fam. 6, 1928, t. 102, f. C-E: Lloyd,
Myc. Notes 60, 1919, 868, f. 1480.
-12 cm. high, very much branched, caespitose, bushy (drying red-brown or
alutaceous, stiff, tough, horny, glabrous, and smooth): stem -2 cm. long, or branched
from the base, not tomentose: branches 1 mm. thick below, rather coarse, somewhat
compressed, forked or bushily divided, axils flattened, acute, tips subulate.
Spores 4-5 µ., white, smooth, subglobose.
On the ground in the forest: central Africa (Wakondjo, Gt. Lakes, 1,950 m. alt.).
This may be Aphelaria.
P. brunneola sp. nov.

Text-Figs. 223, 238a.
-9 cm. high, 12 cm. wide, forming very bushy, multaxial tufts, pale fav.;n-drab,
PTERULA
greyish at the base, pale brown towards the pallid white apices, rather tough and dry,
with a slight fishy smell: drying fuscous brownish, horny, with pallid drab hymenium
and spirally twisted branches.
Stem 3-25 x 1 ·5-2 mm., often indistinct and branched from the base: main branches
c. 1 mm. wide, 2-3-4-chotomous below (actually successive close dichotomies), the
successive branchings in alternating planes, multaxial, the narrower branchlets
dichotomous, strict, dense, very abundant, tapering to fine (but not filiform) tips, some
of the main branches decumbent with upright side branches forming bushy tufts:
hymenium absent from the uppersides of the main branches.
T'ExT-FIG. 223. Pterula brunneola, with apical branching, X 2.
On the gtound in humus, in the forest: Malaya (lowland, Tembeling, Pahang,

21.11.30).
Spores 6-7 X 3-3·5µ,, white, smooth, pip-shaped, thin-walled(? with a small gutta).
Basidia 20-30 X 6->; µ,, clavate: sterigmata 4, 3-4µ, long.
Cystidia none.
Hymenium thickening to 100µ,, the old basidia collapsing: subhymenium composed
of narrow uninflated submucilaginous hyphae.
Hyphae and general construction as in Pterula verticillata: skeletal hyphae 2-5µ,
wide, with walls 0·5-1 µ, thick: generative hyphae qccasionally septate without clamps.
508 PTERULA
P. capillaris (Uv.) Sacc.
Syll. Fung. 6, 1888, 742: Wakef., Kew Bull. 1912, 143: Lloyd, Myc. Notes 60, 1919,
867, f. 1477: (?= Clavaria tenuissima Uv., An. Sci. Nat. Ser. 3, 5, 1846, 156).
Basinym: Merisma capillaris Uv., Ann. Sci. Nat. Bot. 3, 2, 1844, 208.
Synonym:? Pterula arbuscula Bres., Hedw. 1896, 291.
-5 cm. high, with many very fine branches like P. multifida, pale ochraceous: stem
simple, naked: branches spreading, very slender, the tips compressed and acutely
penicillate.
On dead wood and bamboos: Peru, Brazil, Java, Philippine Is!., Nigeria.
Spores 10 x 5 µ. (Lloyd), 11-12 x 6--7µ., mostly 12 x 7µ. (Wakef.), white, smooth,
ellipsoid, inequilateral with rather acute ends, apiculate at the base.
This species was originally described from Java. Lloyd doubts whether the African
plants, so named at Kew, or the Philippine plants, so named by him, are really P.
capillaris, which was described as 'espece capillaire comme Pterula Commersonii'.
An African collection, from northern Nigeria, referred to this species by Miss
Wakefield, has extremely slender, mostly monopodial branches 150-250µ. wide, the
main stem being c. 300µ. wide near the base, at its thickest part, and quite smooth.
The hyphae are typically dimitic with 2-4(-5)µ. wide skeletals. The basidia are
c. 30 X 10µ. and the hymenium appears not to thicken. I found a few spores lo-'l 1 X
6--7µ.. It is certainly distinct from Pterulicium xylogenum, though the large sub
amygdaliform spores suggest affinity.
P. arbuscula (sp. 10-12 X 7µ.) may be identical. The description (of dried specimens)
is indistinguishable. Lloyd suggests that it is P. tropica, :which I place in Aphelaria.
P. Commersonii (Lev.) Lloyd

Myc. Notes 60, 1919, 868, f. 1482.
Basinym: Thelephora Commersonii Lev., Ann. Sci. Nat. Ser. 3, 2, 1844, 208.
-8 cm. high, branched from the base: branches erect, fastigiate, not numerous,
c. 0·5 mm. thick (dried).
Mauritius.
Lloyd says this is like P. trachodes, which I refer to Aphelaria. He suggests P.
secundiramea is identical. It may be Scytinopogon.
P. debilis sp. nov.

-5 cm. high, with few or many, mostly simple, branches, very slender, weak, flaccid,
white � pale brownish from the base upward, irregular, occasionally subsimple or
simple, solitary or gregarious: stem 5-25 mm. long, 0·2-0·4 mm. wide, fertile except
for a short sterile base 1-0 mm. long, seated on the matrix: branches 100-200(-300)µ.
w�de, adventitious, curved ascending, without particular order, lax or subfastigiate on
the main axis, tapering to filiform tips 15-40µ. wide: rather tough.
Gregarious, often densely, on dead, fallen stems of Juncus in bogs: England
(Whittlesford Fen, Cambridgeshire, 24.7.46).
Spores 7-g·5 X 3·5-4µ., white, smooth, narrowly ellipsoid, aguttate or with 1-2
minute guttulae, often 1-guttate on drying, apiculus rather prominent.
Basidia 19-26 X 6--7µ., subclavate, hyaline: sterigmata 4, 4µ. long.
Cystidia 30-40 x 5-8µ,, sparse, thin-walled, hyaline, narrowly flask-shaped or
fusiform-subventricose, apex subacute, often shortly prolonged.
Hymenium not thickening, compact, slowly disorganizing from below upwards on
the main stem: subhymenium 8-20µ, thick, composed of 1·5-3µ, wide, closely inter
woven hyphae.
Caulocystidia sparse, as ill-formed hymenial cystidia, thin-walled, or as scattered
hyphal ends 2·5-5µ, wide at the base, tapering to 100µ, long, thin-walled, not incrusted.
PTERULA 509
Hyphae dimitic: skeletals 3-5µ wide, walls 0·5-1(-1·5)µ.thick, aseptate, unbranched,
unlimited or becoming transformed again into generative hyphae, longitudinal:
generative hyp hae 2·5-4µ wide, thin or very slightly thick-walled, clamped and
usually branching from the clamp.
TExT-FIG. 224. Pterula debilis, with adventitious branching, x 2.
TEXT-FIG. 225. Pterula debilis: cystidia and spores, x 1,000.
Stem becoming thinly levigate from below upward with. an epithelial layer of
flattened, contiguous, irregularly interwoven generative hyphae with slightly thickened,
agglutinated, brownish walls, one cell thick.
The fruit-bodies are usually laxly and irregularly branched, but monopodial. The
510 PTERULA
primordial shaft can be traced as an axis on which the branches develop adventitiously,
there being no apical flattening or dichotomy. Stunted fruit-bodies may remain
simple, and become fertile, and there are all transitions to the typical state. Sparing
seems to occur mainly during the night, or when the fruit-bodies are shut in the dark.
I found this species after I had written this monograph on Pterula, and was able,
theref6re, to study it critically. Among European species it seems to come between
P. taxiformis sensu Bourd. et Galz. and P. gracilis, so far as concerns the form of
the fruit-body and the size of the spores, but it is certainly distinct from both. The
absence of mycelial subiculum, the smaller spores, the 4-spored basidia, and the
levigate stem show that it cannot be interpreted as an overgrown state of the normally
diminutive and simple P. gracilis. Among tropical species it comes near to P. typhuloides
which is less branched, has distinct and abundant caulocystidia, and lacks both clamps
and hymenial cystidia. P. _epiphylla appears to be constantly simple and lacks the
levigation of the stem. P. debilis, therefore, as a summer-fungus in the north tem
perate fens seems to be a link between the busy humicolous species so characteristic
of the tropics and the simple Pistillarioid derivatives with restricted mycelium in
plant-fragments.
P. densissima B. et C.
Grev. 2, 1873, 16: Lloyd, Myc. Notes 60, 1919, 865, f. 1469: ibid. 75, 1358, f. 3213.
Like P. multiji.da but pulvinate-congested, very much branched from a common
base: branches repeatedly and ( ?) equally dichotomous, fine, with penicillate tips:
drying rufous.
On the ground: U.S.A. (Mass.), France.
How this differs, if at all, from P. multifida remains to be shown.
P. dilatata (Mont.) comb. nov.

Basinym: Clavaria dilatata Mont., Bel. Voy. Ind. Or. Crypt. 1851, 152.
Synonym: Calocera dilatata Mont., Syll. Crypt. 1856, No. 602.
Lachnocladium dilatatum (Mont.) (? author).
Ochraceous cinereous, very fragile, highly branched: branches compresst::d,
irregularly dichotomous, tips subulate, fastigiate, axils strongly dilated.
On the ground by mountain woods: Madura Peninsula (Belanger).
The specimen at Kew, ex herb. Mont., but not the type, is a Pterula.
P. epiphylla sp. nov.

Text-Figs. 226 G, 227-8.
Fruit-body -18 mm. high, simple, scattered, filiform, erect: stem 2-4 mm., 200-300µ
wide at the base, 130-170µ wide at the top, gradually tapered from the base, erumpent,
pale yellowish becoming pale fuscous, subpellucid, puberulous: fertile part 120-250 µ
wide, tapered to the persistently sterile apex 20-50 µ wide, opaque white then pale
yellowish.
On dead leaves of Ixonanthes icosandra (Linaceae): Malaya (Singapore, Gardens'
Jungle, 18.3-43).
Spores 9-13(-14) X 4-5µ, white, smooth, ellipsoid, subacute, flattened adaxially,
often becoming slightly thick-walled, aguttate.
Basidia 22-28 x 8-10µ, clavate: sterigmata 4, 5 µ long.
Cystidia 30-50X5-11µ, conical or subventricose, thin-walled, vacuolate, with
obtuse apex 2-3µ wide, frequent, projecting considerably, hymenial (.without digitate
cystidia).
Hymenium not thickening, continuous, often originating unevenly at the upex of
the stem: subhymenium 30-35 µ thick, composed of closely interwoven, narrow
hyphae 2-4µ wide, in places -ro µ wide, with short cells 5-15 µ. long, thin-walled.
PTERULA 511
Caulocystidia -45 X4-IIµ, scattered or abundant, subclavate to subventricose,
blunt, thin-walled and colourless, or with slightly
thickened (0·5µ wide), pale brown walls, smooth,
vacuolate.
Stem without a sterile hymenium, the superficial
longitudinal hyphae with slightly thickened, pale
brown walls.
Hyphae dimitic: skeletal hyphae 3-5µ wide, in
places up to 15µ wide, with colourless thickened walls
0·5-1·5µ wide, often rather flexuous and kinked or
nodose, smooth, aseptate, of unlimited growth, up to
1,500µ long, mostly unbranched but commonly
ending in generative hyphae: generative hyphae 2-5 µ
wide, septate with clamp-connexions, thin or slightly
thick-walled, longitudinal, profusely branched near
the subhymenium; the cells 30-250µ long, but short
near the subhymenium, often but not always branching
from the clamp.
Colour of the fruit-body in the walls of the skeletal
hyphae, of the superficial hyphae on the stem and of
the caulocystidia.
As a simple Pteruloid derivative, P. epiphylla is A •D E r C
similar to P. gracilis, but I think it is on a different
TEXT-FIG. 226. Pterula typhu
line of degeneration as shown by its erumpent habit, loides, A-D: var. minor, E: P. ·
by the brown walls of the superficial hyphae on the subtyphuloides, F: P. epiphylla,
stem, and by the presence of the caulocystidia. G: x2.
Though it has a simple hymenium like P. typhuloides,
its subhymenium is better developed, or less rudimentary. All three appear connected
with P. debilis as the intermediate between the simple and the bushy species.
P. exserta Lloyd
Myc. Notes 60, 1919, 866, f. 1475, 1476.
-1·5 cm. high: stem 5-8xo·5 mm. dividing into 2-4 branches one to three times
dichotomous with flattened axils and flattened subspathulate tips(incurved on drying):
sometimes apparently monaxial.
On wood: ? origin(type-specimen at Berlin, named P. plumosa by Henn.).
This may be the young state of an Aphelaria or even a Xylaria. As published it is
probably unrecognizable, and Lloyd failed to note the country of origin.
P. falcatula(P. Henn.) comb. nov.

Basinym: Clavariafalcatula P. Henn., Hedw. 1897, 195.
3-5 X0·5-0·7 mm., simple, fleshy, subochraceous (? dried), straight or curved,
subulate, pallid at the byssoid base.
. On rotting herbaceous stems: Brazil(Sta. Catherina).
Spores 3·5-4µ, subglobose, smooth(? spore-wall brownish).
Certainly no Clavaria. Compare P. tenuissima.
P. filaris (Kalchbr.) comb. nov.

Basinym: Clavaria filaris Kalchbr., Grev. 10, 1882, 106.
2·5 cm. high, white, arising from a vaguely branched mycelial fibril: branches
slender, elongate, narrower than a thread, acute: on rotten wood: S. Africa(Somerset
East).
Almost certainly Pteruloid. Perhaps Pterulicium.
512 PTERULA
P. fructicola Bres.
Hedw. 56, 1915, 304,
2-3 cm. X 1·5-2 cm., isabelline becoming fuscous (? dried): stem -6 X 0·5 mm.,
sometimes branched from the base: branches few, glabrous, subfiliform: branchlets
bifid or alternate, with acuminate apices.
TEXT-FIG. 227. Pterula epiphylla: hymenium, spores, generative hyphae (often

branched from the clamp), and unusually massive skeletal hyphae: X 1,000.
On old fruits of Pandanus: Philippine Is!. (Luzon).

Spores 9-10 x 5-6µ, white, smooth, subnavicular or sinuous.
Basidia 25-28 X 6-7 µ, clavate-subcapitate.
Hyphae 1·5-3·5µ wide.
This may be P. typhuloides. If the basidia are correctly described they must be
2-spored.
PTERULA 513
P. fulvescens Bres.
Ann. Myc. 9, 1911, 551.
5-8 X 0·25 mm., densely gregarious, simple or very rarely furcate or twice furcate,
subulate, somewhat tawny (? dried), slender, seated on a straw-coloured mycelial crust.
>
; i
··-·i:
TExT-FlG. 228. Pterula epiphylla: hymenial cystidia (above) and caulocystidia,

X 1,000.
On dead herbaceous stems: south-east Borneo (Winkler·2241).
Spores and basidia?
Hyphae 2-4·5 µ., hyaline.
The mycelial crust suggests P. gracilis or Pterulicium, unless it is a very rudimentary
state of Lentaria surculus.
P. Gordius (Speg.) comb. nov.
Basinym: Clavaria Gordius Speg., Fungi Guar. Pug. 1, 1883, n. 88.
Synonyms: Clavaria subulata B. et Cke., Journ. Linn. Soc. Bot. 15, 1876, 391.
? Typhula Uleana P. Henn., Hedw. 1897, 195.
-15 xo·o8 mm., simple,filiform, attenuate acute, erect, base inserted, glabrous, dull
ochraceous white (dried): stem short, sterile, without caulocystidia.
5119 L l
PTERULA
On dead leaves in the forest: S. America (Brazil: pr. Guarapi).
Spores 8-10 X 3·5-4·5µ, white, smooth, oblong ellipsoid, obtuse or subacute,
distinctly apiculate (very abundant in the co-type Balansa, No. 3755, in herb. Kew.):
5-7 X 2·5-3µ (Speg.).
Basidia 15-20 X 5-7µ (Speg.).
Cystidia none: hymenium not thickening.
Hyphae dimitic: skeletals 3-5µ wide, unlimited, aseptate, the walls o· 5-1· 51-' thick:
generative hyphae? (in C. subulata clamped).
Stem not levigate with sterile hymenium.
I have examined part of the type-collection in the Kew herbarium but could find
no spores as small as Spegazzini gives. The type-collection of Clavaria subulata is
also identical, but I could find no spores: its habitat was 'on the midrib of dead leaves,
Brazil (Rio Jurua, Trail, No. 172)'. C. subulata is an earlier name but, if transferred
to Pterula, becomes a later homonym of P. subulata Fr.
The description of T. Uleana is:
10-20 X 0·5 mm., filiform, subulate, drying pallid or brownish, subtomentose, base
scarcely thickened: spores 4-4·5µ, white, subglobose: basidia 10-14x6-7µ: on dead
leaves: Brazil (Sta. Catherina).
Except for the spore-measurements, which are not reliable, T. Uleana is identical.
P. Gordius is probably a widespread common species in tropical America. P. tucu
manensis is very similar but has a fuscous base to the subsclerotioid insertion of the
stem, which suggests fuscous levigation of the stem as in P. debilis, P. epiphylla, and
P. typhuloides and which is lacking from both P. Gordius and Clavaria subulata.
P. gracilis (B. et Desm.) comb. nov.

Basinym: Typhula? gracilis B. et Desm., Ann. Mag. Nat. Hist. 1, 1838, 202, t. 8, f. 1:
(non Pat., Tab. An., No. 575 = ? Hirsutella gracilis Pat.: non Bourd. et Galz. =
Pistillaria typhicola).
Synonyms: Pistillaria gracilis (B. et Desm.) Pat., Tab. An. 1886,. No. 575, quoad
nomen: (quoad spec. = ? Hirsutella gracilis Pat.).
Clavaria aculina Quel., Ass. Fr. 1880, 670, t. 8, f. 11.
Pistillaria aculina (Quel.) Pat., Tab. An., No. 570: Bourd. et Galz., Bull. Soc. Myc.
Fr. 1910, 221: Hym. Fr. 1928, 138: Corner, Tr. Br. Myc. Soc. 19, 1935, 285.
Ceratella aculina (Quel.) Pat., Hym. Eur. 1887, 157.
Cnazonaria aculina (Quel.) Donk, Rev. Nieder!. Homob. Aphyll. 2, 1933, 96.
Cerat_ella Ferryi Quel., Rev. Myc. 1893, 15.
Pistillaria Ferryi (Quel.) Sacc., Syll. rr, 1895, 141.
Pistillaria attenuata Syd., Hedw. 1900, r: Ann. Myc. 19, 1921, 137.
Pistillaria acicula Bourd. et Galz., Hym. Fr. 1928, 139.
Pistillariajuncicola Bourd. et Galz., ibid. 138.
Typhula Brunaudii Quel., Ass. Fr. 1884, 283, t. 8, f. 14.
Text-Figs. 43-5, 229, 230.
3-1r mm. high, o·1-0·3 mm. wide (1-5 X 0·08-0-14 mm., Bourd. et Galz.) acicular,
attenuate to afiliform apex, rarely subcylindric with obtuse, abrupt, or mucronate apex,
sessile or with an indistinct stem 0·1-0·5 mm. long, simple, white, occasionally with an
adventitious branch from the stem, more or less gregarious or subfasciculate, super
ficial on a slight mycelial patch, the base often slightly dilated and subvillous, white
then brownish: hymenium continuous from the base, sometimes lacking from the
extreme base or decomposing from below upward and the stem secondarily denudate,
the sterile apex sometimes becoming fertile on cessation of growth.
On dead stems and leaves of Juncus, Carex, Cladium, Typha, Eupatorium, grasses,
&c., in swampy places: Europe, common.
Spores 9-16 x 4-7µ, white, smooth, ellipsoid-oblong, blunt or subacute, thin-walled,
aguttate, cloudy-vacuolate.
PTERULA
Basidia 20-30 X7-9µ,, clavate: sterigmata 2, 4-5µ, long.
Cystidia 25-40 X 5-8 µ, frequent, varying from sterile basidia, narrowed at the apex,
to subconical or subventricose cells with subacute apex, thin-walled, colourless,
smooth, rarely with a small subglobose stalked appendage as an abortive spore.
Hymenium not thickening: subhymenium very slightly developed, short-celled, or
the basidia arising directly from the longitudinal hyphae.
Caulocystidia none or as scattered, rudimentary
basidia or cystidia on the short sterile. stem,
-20X 6 µ, thin-walled, obtuse.
Hyph?.e dimitic: skeletal hyp hae 2·5-3·5µ,(-4µ,)
wide, the walls o·5-1µ, thick, aseptate, rarely with
a short branch, smooth, traversing the length
of the fruit-body, abundant: generative hyphae
1·5-3µ, wide, thin-walled, clamped.
This fungus is better known as Pistil/aria aculina.
Donk has shown me his ms. notes on Typhula
gracilis, however, and there can be no doubt that
it is the same. Donk examined the type in
Desmazieres's collection in the Paris herbarium,
labelled 'Typhula ? typhae nobis. Clavaria typhae
nob. Berk. det. Typhula ,!racilis' : 1 he noted the
following points:
Receptacles 1·5-u xo·1-0·18 mm., sessile,
slightly swollen at the base, apex acute and sterile
when young, becoming fertile with age: solitary or TEXT-FIG. 229. Pterulagracilis, X 10.
in small groups of 2-3.
Spores 10--13x5-6µ,, oblong pip-shaped, obliquely apiculate, subacute.
Basidia 18-28x 6-7·5 µ, clavate: sterigmata 2.
Cystidioles 25-35X 4-6µ,, subventricose, projecting, with a filiform apex 2·5µ, wide.
Hyphae 3-4·5µ, wide, thick-walled, septa few.
T. gracilis has not been recognized correctly in recent mycological works because of
Patouillard's erroneous interpretation of it as Hirsutella and Bourdot's erroneous
interpretation of it as Pistillaria typhicola, with a hairy stem and much smaller spores,
as quoted by Rea. Desmazieres's collection, as described by Donk, agrees exactly
with the fen specimens from Cambridgeshire which I described in 1935 as Pistil/aria
aculina, and Patouillard's figure of P. aculina fits exactly Donk's sketches of Des
mazieres's specimens except in size. Most authors give the fruit-bodies as very small:
for T. gracilis, Berkeley gives 2-3 mm. high: for P. aculina, Quelet gives 1-2 mm.,
Patouillard 1-5 mm., Bourdot and Galzin 1-4 mm. In the Cambridge fens I found
them to vary 1-10 mm. in height and 0·1-0·3 mm. in width, which measures corre
spond well with Desmazieres's specimens. The characteristics of the species are the
sessile or subsessile, acicular fruit-body, the 2-spored basidia, the cystidia, and the
rather long spores, all of which are given by Patouillard, Bourdot, and Donk.
I agree with Bourdot in reducing Pistillaria acicula and P. juncicola as synonyms,
for there is great variation in the spore-size of P. aculina, which covers the descrip
tions of the two species. Donk's ms. notes, however, suggest that there may be a
form on J uncus which has separate and gregarious, not fasciculate, fruit-bodies
(-5xo·o7-0·1 mm.) and small spores 7·5-12X4·5-5·5µ, (basidia 2-spored, 18-25x
7·5-9µ,: coll. H. Bourd., No. 24978).
Bourdot also suggests that Pistillaria equiseticola, P. graminicola, and P. typhicola
1
Berkeley noted that Desmazieres agreed to change the unpublished name Typhula
? typhae to T. gracilis (presumably to avoid any chance of tautonomy). The new name was
published, therefore, to cover Desmazieres's collection, which must now be regarded as the
type.
516 PTERULA
belong here, but the first two I refer to Ceratellopsis because it is not clear that they
have 2-spored basidia, cystidia, or skeletal hyphae, and P. typhicola I retain in
Pistillaria because of its resemblance to P. setipes. All three have consistently smaller
spores (6-roX3·5-5µ.) than Pterula gracilis. On the other hand, Ceratellopsis Rickii
and C. Asphodeli may well be identical with P. gracilis. It is impossible to decide until
their hyphal construction is known.
The description of Pistillaria Ferryi agrees exactly with that of Pterula gracilis and
such characteristic points as the superficial mycelium and 2-spored basidia are men
tioned, but cystidia are omitted (probably overlooked). It grew on dried culms of
Coix in France. Similarly with Pistillaria attenuata, on leaves of Typha latifolia, for
which Donk's ms. notes give: r-2·5Xo·o6-o·r5 mm.: spores ro-12·5X4-5·5(-6)µ.:
basidia 20-25X7·5µ.: 2-spored: basidioles 28µ..
TEXT-FIG. 230. Pterula gracilis: spores (one germinating), basidia and cystidia, and
young hymenial hyphae (on the right): X 1,000.
P. gracilis differs from other simple Pterulas, especially the tropical ones, mainly in
the superficial, not erumpent, fruit-bodies, in the absence of any thickening or brown
discoloration of the walls of the superficial hyphae on the stem, and in the absence of
caulocystidia. The skeletal hyphae are also narrower in P. gracilis. It is the only
Clavarioid fungus that I have found so reduced that its fruit-body originates from a
single hypha (p. r 19). Doubtless other species related to it will be discovt!red among
'Pistillaria' and Ceratellopsis. Pterula debilis seems the branched, more robust, 4-spored
state which connects it with the bushy Pterulas, though the branching of P. debilis is
only adventitious.
The table on the opposite page gives the data on the spores of P. f;!racilis. It should
be noted that their greater length, compared with other 4-spored species, is to be
related with the 2-spored basidia, dispores having roughly twice the volume of
tetraspores. The largest spores are possibly derived from 1-spored basidia.
P. grandis Syd.
Engl. Jahrb. 54, 1916, 252, f. r E.
-9 cm. high, pale brown or yellowish brown, drying rufescent, glabrous, cartilaginous,
very much branched: stem -I cm. long, short, arising from a mycelial patch: branches
repeatedly divided, subcompressed, tips subulate.
On dead stems: New Guinea (north-east, Malu): Africa (? locality).
Spores 4-5 µ. wide, white, minutely asperulate, globose.
PTERULA 517
Hyphae 3-4µ.
This is said to be near P. pallescens for which Singer has made the genus Scytino
pogon (p. 647). It is the only Pterula with globose asperulate ·spores and it is reason
able therefore to ask for verification. Sydow's figure shows 3 short subcaespitose
stems, the lower branches 2-3-fid, the upper regularly and equally dichotomous, at
an acute angle.
Spore-size in µ. Author Species or Collection

10-13X5-6 Donk Typhula gracilis (co-type)
12-16X5-7 Bourdot Pt. aculina (France)
12-16 X5·5-7 Donk (Bourd., 4,700)
10·5-15X5-6·5 (Bourd., 6,777)
I 1-13·5X5·25-6 (Bourd., 37,376)
10-12·5 X5-5·5 " (Bourd., 37,378)
9-14X5-'7 Corner (England)
12 X5 Patouillard (France)
12, pruniform Quelet ,, "
10-12·5X4-5·5 Donk Pist. attenuata (type-collection)
9-11X4·5-6 Pist. acicula (Bourd., 15,233)
7·5-12X4·5-5·5 Pist. juncicola (Bourd., 24,978)
The collection L. S. Gibbs, No. 6165 (Manokoeari, Dutch north-west New Guinea),
identified as this species, differs in these respects (according to the specimen in herb.
Kew.):
(a) fruit-bodies 'smoke-grey' (living), on the ground;
(b) spores 6-7 X3µ, smooth, ellipsoid (abundant).
The stem and main branches, 0·5-0·7 mm. wide (dried), are rather stout: the
branching is mainly dichotomous: the hymenium is limited to the undersides of
the branches: the basidia are c. 20-25X5-6µ: the skeletal hyphae are 3-5µ wide. The
collection is near P. brunneola and P. vinacea.
P. hirsuta P. Henn.
Mons. 1, 1900, 9.
Synonyms: Phaeopterula hirsuta P. Henn., Hedw. 1904, 175.
Dendrocladium hirsutum Lloyd, Myc. Writ. 5, 1919, index 12.
Very much branched, tough, cartilaginous, reddish brown (? dried), violaceous
strigose hirsute, especially in the lower part, and giving a whitish violaceous appearance:
stem thick, branched: branches verticillately arranged and branched, filiform, very
crowded, erect, with subulate tips.
On the ground: Java (Tjibodas, c. 1,500 m. alt.).
Spores 6-8 x 4 µ, hyaline fuscescent or clear brown, smooth, elliptic-ovoid, apiculate
at the base.
Basidia 30-35 X 7-9µ, fuscous: sterigmata 4.
Because of the brown spores, Hennings fir§t made the subgenus Phaeopterula for
this species and later raised it to generic ran1:. It is doubtful, nevertheless, whether
the spores have been correctly described. Concerning Dendrocladium, see page 82.
P. incarnata Pat.
Bull. Boiss. 1895, 58.
3-5 mm. high, scattered or caespitose, rigid, tough, erect, pale flesh colour: branches
spreading, simple, paler, covered all over with basidia.
Spores not found.
On pieces of wood: S. America.
? Deflexula.
PTERULA
P. Janseniana (Holterm.) comb. nov.
Basinym: Clavaria Janseniana Holterm., Myk. Unters. Trop. 1898, 103, t. 11,
f. 1a-c (ut C. Janseiana, ? lapsus: vide iconem).
Synonym: Lachnocladium Jansenianum (Holterm.) P. Henn., Mons. 1, 1900, 143
(ut L. Janseanum):
Upright, leathery, bushy, with a cylindric stem, repeatedly dichotomously branched
to filiform tips: stem 1-2·5 cm. X 2-6 mm.: yellowish below, brownish above.
Spores 2·8-4xo·6-1·1µ.: basidia with 4 sterigmata.
On living and dead twigs, covering them with a white mycelial weft of clamped hyphae,
mycelium perennating and covering the twigs for a metre length without fruit-bodies:
Java.
The position of this fungus is doubtful. In habit it recalls Lentaria surculus, which is
almost certainly the fungus figured by Lloyd from northern Borneo as L. Jansenianum
(Myc. Notes 66, 1922, uo6, f. 2027). But the spores given by Holtermann are much
too small-the smallest yet recorded in the Clav:ariaceae !-unless they are the very
young spores of L. surculus which are as narrow and elongate. According to Holter
mann's figure the spores are 2µ. wide. Hennings suggested Lachnocladium, but there
is no evidence that the fungus has dichophyses. Hence I transfer it provisionally to
Pterula. Compare P. Lecomtei.
P. juruensis (P. Henn.) comb. nov.
Basinym: Phaeopterula juruensis P. Henn., Hedw. 43, 1904, 175: v. Hoehne!, Ann.
Myc. 9, 1911, 174.
Synonym: Dendrocladium juruense (P. Henn.) Lloyd, Myc. Writ. 5, 1919, 870,
index 13.
1 cm. high, brownish cinereous (? dried), subsimple or fasciculate: stem 2-3 mm.,
terete, hirsute-tomentose, with many filiform, subulate, squarrose, cinereous, pruinose
branches racemosely arranged.
On rotten wood: Amazon (Rio Jurua).
Spores 4-6µ., subglobose, smooth, white.
Hennings gave the spores as brown, and on this point made the genus Phaeopterula.
v. Hoehne! re-examined the type-specimens and found them to be hyaline. Possibly
related to P. taxiformis, P. adustipes, and P. secundiramea. Concerning Dendrocladium,
see page 82.
P. laxa Pat.
5-8 mm. high, solitary, scattered, white or rufescent, simple or with I-J spreading
branches, 250µ. thick, filiform, acute: hymenium covering the middle part.
On rotten fragments of Richeria grandis (Euphprbiaceae): Guadeloupe.
Spores, basidia?
This is said to be near Pterulicium xylogenum.
P. macrospora (Pat.) comb. tiov.
Basinym: Ceratella macrospora Pat., Bull. Soc. Myc. Fr. 1892, u9.
Synonym: Pistillaria macrospora (Pat.) Sacc., Syll. I 1, 1895, 142.
I mm. high, very small, scattered or gregarious, simple or Jurcate, filiform, everywhere
covered with basidia except the sterile acuminate apex, pale brownish: texture
filamentous.
On dead branches: Ecuador (Guapolo).
Spores 18-20 x 10-12µ., hyaline.
Basidia 18-20 X 8-10µ., 2-spored.
This is most probably a reduced Pterula. If the size of the spores is correct, that
of the basidia is wrong. Possibly a very small Deflexula.
PTERULA 519
P. multifida Fr.
Linnaea 5, 1830, 531: Monogr. Hym. Suec. z, 282: Icon., t. 200, f. 2: Quel., Ass.
Fr. 1886, 488: Lloyd, Myc. Notes 60, 1919, 863, f. 1464, 1465: Konr. et Maubl., le.
Se!. Fung., t. 486.
Synonyms: Penicillaria multifida Chev., FI. Gen. Par. 1, 1826, 111.
? Clavaria carnea Wallr., FI. Crypt. Germ. 2, 1833, 541.
1-5 cm. high, very much branched from the base, often caespitose, pallid whitish,
silvery white, pearl-grey, even lilaceous, then dirty pale yellowish, finally brownish,. almost
black when dried, delicate, flaccid, slightly tough or cartilaginous: stem generally
indistinct: branches tense, straight, not much thicker than a hair, heaped as if swept
together or with spreading branches giving a broadly conical outline, tips spear-shaped
or crisped.
On coniferous needles, twigs, and branches: Europe (Sweden, Gt. Britain, France,
Germany, Spain: rare).
Spores 6 x 3 µ, (Rea), 4·5-7·5 X 3-4µ (B. et G.), 5-7 X 3-4µ (K. et M.), 5 µ long
(Quel.), white or slightly brownish in the mass (K. et M.), smooth, pip-shaped or
ellipsoid, contents granular.
Basidia 24-30 X 5-6µ: sterigmata 2-4.
Hyphae 3-6 µ wide, skeletal hyphae slightly thick-walled, generative hyphae
clamped.
I have examined an authentic English specimen of this type-species in the Kew
herbarium and find that it has the typical dimitic structure of the tropical Pterulas,
though the skeletal hyphae have only slightly thickened walls. Hennings has recorded
the species from Java, but until the details of its branching are known it cannot be
identified with certainty.
The description of C. carnea, which seems never to have been rediscovered, is
difficult to follow, but suggests a Pterula. Thus 'Pileis angustissimis linearibus,
cylindricis, tenacellis, carneis, a basi inde in ramos conformes dichotomos divulsis:
ramulis extremis acutissimis, bi-trichotomis, flexuosis, elongatis, raro simplicibus: ad
caudices quercinos muscosos in Thuringia Gennaniae.'
P. nana Pat.
1-1·5 mm. high, scarcely 250 µ, wide, like a Mucronella, drying whitish rufescent,
scattered or subfasciculate: entirely covered by the hymeniurn, except the tip.
Spores 7-10 X 5-6µ, white, smooth, ellipsoid.
On twigs: Guadeloupe.
P. oryzae (Remsberg) comb. nov.

Basinym: Pistillaria oryzae Remsberg, Mycol. 32, 1940, 667, f. 1.
Synonyms: ? Clavaria gracillima Wakker, Die Ziekten v. h. Suckerriet o. Java,
Leiden 1898, 195 (haud C. gracillima Pk. 1876).
? Clavaria Wakkeri Sacc. et Syd., Sy!!. Fung. 14, 1899, 237 ( = C. gracillima
Wakker).
2·5-8 cm. high, white then light ochraceous to pinkish cinnamon, filiform, simple and
gradually tapered from the base to the acerose-filiform sterile apex, or with I-J short
acerose-filiform branches above, wholly covered by the hymenium except for the tip, or
only in the middle part, but not divided into stem and head: without sclerotium.
On dead leaves of Oryza sativa, apparently saprophytic: U.S.A. (Louisiana).
Spores 9-II · 5 x 5-8 µ, (av. 11 x 6·5µ), white, smooth, ovoid to fusiform, truncate
with a prominent apiculus, with a distinct guttula.
Basidia 2 7-3 5 X 8-1o µ,, clavate: sterigmata 4, 3-4 X 1 · 5-3µ.
Cystidia none.
520 PTERULA
Hyphae compact, parallel, sometimes spirally arranged: subhymenium compact:
longitudinal hyphae loosely interwoven, clamped, incrusted with small crystals.
Mycelial hyphae clamped.
Temperature-range 6-33 ° C., opt. 24-27 ° C.
Spore-germination in 24 hours, with 1-2 germ-tubes soon becoming branched
hyphae.
I have little doubt that this is a subsimple Pterula related to P. tenuissima. The high
optimum temperature of growth indicates a tropical affinity.
Remsberg says that successive crops of fruit-bodies were obtained from rice-leaves
in a moist chamber by placing the chamber for 2-3 days in a refrigerator at 3 ° C.: the
process was repeated five times, each time giving 8-10 new fruit-bodies.
The description of C. gracillima Wakker is:
10-30 mm. high, filiform, solitary or 2-5 fasciculate, simple, pale ochraceous, very
acute: spores 8 X 5 µ,, white, ovoid, thin-walled, acute at each end: basidia 4-spored:
on dead sheaths of sugar-cane: Java.
Macroscopically it is indistinguishable from P. oryzae, but these fungi can be
studied accurately only in hyphal detail.
P. penicellata Lloyd
Myc. Notes 60, 1919, 863, f. 1466, 1467: ibid. 74, 1925, 1340, f. 3125 (Indian
specimens).
Rather large, bushy, apparently monopodial with verticillate branches (as P. verti
cillata): stem rather stout: branches 1-1 ·5 mm. wide below, spuriously polychotomous
below, equally dichotomous above, or the main branches repeating the verticillate
0
arrangement of the stem.

On humus in woods: U.S.A., Canada (Man.), India.
From Lloyd's photographs this appears similar to P. verticillata, and the Indian
specimen may be identical with P. verticillata of Malaya. The structure, branching,
spores, and so on of P. penicillata must be described before it can be recognized.
P. penniseti (P. Henn.) comb. nov.

Basinym: Pistillaria penniseti P. Henn., Engl. Jahrb. 22, 1895, 86.
10-20 mm. long, gregarious, filiform, simple or branched, white, base slightly
bulbous, smooth: branches filiform, sparse, with subulate tips.
On fallen leaves of Pennisetum: tropical Africa (Cameroons).
Spores 6-'7 X 4-5 µ, white, ovoid.
Basidia 5-6 µ wide, clavate.
This is almost certainly Pteruloid.
P. phyllophila (Mcalp.) comb. nov.

Basinym: Clavaria phyllophila Mcalp., Ag. Gaz. N.S.W. 7, 1896, 86, f. 7, 8.
-2·5 cm. high, pale ochre, nearly white, slender, simple or with a few irregular
divaricate branches, the tips acute, the base brown.
On decayed leaves of Rhagodia billardieri: Tasmania.
Spores 9 X 4 µ, white, smooth, ellipsoid.
Basidia 4-spored.
From the figure and habitat there can be no doubt that this is a reduced Pterula,
possibly near to P. typhuloides.
P. plumosa (Schw.) Fr.

LinnaP.a 5, 1830, 532: Lloyd, Myc. Notes 60, 1919, 864, f. 1468: Coker, Clav.
U.S. Can. 203, t. 77: Overholts, Myc. Exp!. Venez. 1934, 306.
Basinym: Clavaria plumosa Schw., Fung. Carol. 1822, n. 1089.
Synonym: P. divaricata Pk., Rept. N,Y. St, Mus. 32, 1879, 36,
PTERULA 521
2·5-4 cm. high, light grey-brown, drying paler (whitish or rufescent, Peck), slender,
brush-like: stem 0·5-r mm. thick, slender, dark wood brown, springing from a hair-like,
greyish brown mycelium, several or many stems close together: branches dichotomous,
about four times, with long, slender, acute divaricate branchlets (branching irregular,
Peck): elastic, tough: taste like musty wood.
On twigs and rotting leaves: U.S.A. (N. Car.), Venezuela (Overholts), Brazil
(Berkeley), Africa (Henn.).
Spores 6·5-7·5 X 3-3·7 µ. (Coker, U.S.A.): 7-8 X 3·5-4·5 µ. (Overholts, Venezuela),
white, ellipsoid, smooth.
This has fewer, laxer, and more spreading branches than P. multifida as well as a
different colour. It may have an interesting structure because the macroscopic
description suggests that the superficial hyphae of the stem may have dark walls as in
the reduced P. epiphylla and P. typhuloides. I think the tropical records should be
regarded with doubt until their structure is known. Coker's photograph suggests that
there may be some overtopping of the branches.
P. pusilla Bres.
Ann. Myc. 18, 1920, 50.
ro-12 mm. high and wide, pale, subavellaneous (? dried): stem 3 X 0·5 mm., with
many mycelial fibrils, rugulose, pubescent, verticillately branched at the apex, white
tomentose at the orbicular base with fimbriate margin: branches few, furcate or
vertici!lately branched, tips circinate (dried).
Spores? Basidia 25-30 X 6 µ.: hyphae 2·5-5 µ., the larger ones rather irregular.
On sticks and leaves: Brazil.
This suggests a reduced 'monopodial' Pterula.
P. rigida Donk sp. nov.

r ·2-3·8 X 0·4-0·7 mm., simple, linear, attenuate slightly to the sterile apex, often
with one or two short branches or the apex with a cluster of short branches, white or
yellowish, rigid, drying rather horny, sessile on a small villous subiculum, solitary or
2-4 gregarious.
On rotten wood: Holland (Vogelensang, leg. Swanenburg de Vege, autumn).
Spores (r3-)r5-r7 x (6-)6·5-7·5 µ., white, elliptic, acuminate to the base, smooth.
H yphae 2-3· 5 µ. wide, distinctly thick-walled in the central tissue, sparingly septate
with clamps.
This species is described in Donk's ms. as Pistillaria rigida nov. sp., with the remark
that it may be near Pterula. It is either Pterula or Ceratellopsis; the thickened walls
of the hyphae and the sparse branching of the fruit-body indicate Pterula. It seems to
be near P. gracilis and it may also have cystidioles and 2-spored basidia.
P. secundiramea Speg.
Fungi Puigg. 1883, n. 167: Sacc., Syll. 9, 1891, 255.
Basinym: ? Clm:aria secundiramea Lev., Ann. Sci. Nat. 3, 2, 1844, 216.
Solitary or caespitose with slender, intricately divided branches and secund brancn
lets: stem 10-20 x 0·3-0·5 mm., filiform, fuscous downward, pale rufescent or flesh
colour upward: branches concolorous, filiform, laxly alternate or subopposite:
branchlets 5-10 mm. long, spreading, secund, acute, straight or incurved at the tips,
simple or sparsely branched.
On rotting branches in the forest: Brazil (Apiahy).
The secund branchlets may be the effect of unilateral illumination ur positive
geotropism on outgrowth from a prostrate log.
vVhether this is a new species or new combination for Clavaria secundiramea is
uncertain: Leveille's description is:
Very much branched, yellowish-rufescent: stem slender, simple: branches spreading,
522 PTERULA
branchlets subulate, secund, erect: on bark of Cocos butyracea, New Granada. This
description fits also Lentaria surculus. Lloyd suggests it may be Pterula Commersonii.
Compare P. juruensis and P. adustipes.
P. setacea (Kalchbr.) comb. nov. ..

Basinym: Clavaria setacea Kalchbr., Grev. 10, 1881, 105.
-2 cm. high (12 cm. in Sacc., Syll. 6, 710; ? in error), caespitose, divided into wiry
branches from the base: branches thick or thin, ending in brush-like tufts, with acute
tips and axils, whitish or rusty: base white villous, glabrous and reddish above.
On the ground: S. Africa (Somerset East).
'Resembling Polysiphonia' (v. der Bijl). Evidently very near P. multifida.
P. subsimplex P. Henn.
Hedw. 1897, 197.
1-1·5 cm. X130-200µ, fasciculate-ramose at the base (yellowish flesh colour and
cartilaginous when dry), filiform, simple, subulate.
On trunks in the forest: Brazil (Blumenau).
Spores 3-3 · 5 µ, white, globose.
This may be Pterulicium xylogenum, but it may also be the sterile rudimentary
development of another Basidiomycete.
P. subtyphuloides sp�c. nov.

-2 cm. high, simple, as in P. typhuloides var. minor, but thicker.
On dead twigs and stems of Costus (Zingiberaceae): Malaya (Singapore, Botanic
Gardens, 19.3.43).
TEXT-·FrG. 231. Pterula suhtyphuloides: sclerotioid surface of the stem, x 1,000.

Spores 10-13 X4-5µ, as in P. typhuloides.
Hymenium gradually thickening -120µ.
Cystidia none.
Caulocystidia as in P. typhuloides var. minor.
PTERULA 523
Sterile hymenium on the stem as in P. typhuloides but seated on a thick subhymenium,
15-30µ deep, composed of short-celled, densely interwoven hyphae 2-4µ wide.
Hyphae as in P. typhuloides, the generative hyphae with clamps at some septa, not
at others: the skeletals inflated in places -15µ wide.
This may be only a state of P. typhuloides, which has not lost the factor of thickening
the hymenium, and thus it connects P. typhuloides with the normal condition of
Pterula. It has the distinctive sclerotium-like surface to the stem. On the other hand,
in spores, occasionally inflated skeletals, and macroscopic appearance it resembles
P. epiphylla.
P. subulaeformis (B. et C.) comb. nov.
Basinym: Typhula subulaeformis B. et C., Journ. Linn. Soc. Bot. 10, 1867, 339.
-20 mm. high, white, greyish and subpruinose when dry, filiform, simple, very
acute, the base tomentose-affixed, sometimes 2-3 individuals from the same base.
On bark: Cuba.
Skeletal hyphae 4-5 µ wide, very thick-walled, lumen linear: generative hyphae
with firm walls, without clamps.
The specimens in the Kew herbarium are sterile or, at most, with a few scattered
incipient basidia. They have the typical Pteruloid hyphae and the absence of clamps
may enable one to rediscover the species. The texture is fibrillose-floccose (when
dried}, not horny as in most simple species of Pterula. It may be an incipient Xylaria,
however, for some species of this genus appear to have skeletal hyphae.
P. subulata Fr.
Syst. Orb.·Veg. 1825, 90: Linnaea 5, 1830, 531: Lloyd, Myc. Notes 60, 1919, 863,
f. 1463.
2·5-4 cm. high, whitish cinereous, densely tufted, sparsely branched, tough, straight,
equal, attenuate at the base: branches strict, erect, fasciculate, not spreading, growing
into each other, multifid at the apex, subulate or filiform, smooth.
On wood among pines, on bare earth and in gardens: Europe (Gt. Britain, France,
Italy}, rare.
Spores 8-10 X 51 µ, white, smooth, ovoid (Sacc.).
This has been chosen as the type-species of Pterula. Its structure is unknown and
it may possibly be Aphelaria, in which case it may be advisable to change the type of
Pterula to P. plumosa.
P. tahitensis Reichardt
Novara Exped. Fungi 1866, 136.
12-14 mm. high, caespitose, horny, yellow, strigose at the base with hoary fasciculate
hairs, dichotomous with a stem: branches with obtuse, compressed apex.
On rotten wood in the forest: Tahiti.
Spores 7-8µ, white, smooth, globose: basidia 4-spored.
The spores suggest Deflexula, but the branching appears to be apical as in Pterula.
P. taxiformis Mont.
Sy!!. C rypt. 1856, No. 601: Lloyd, Myc. Notes 60, 1919, 866, f. 1474: Bourd. et
Galz., Hym. Fr. 1928.
Synonym: Lachnocladium taxiforme (Mont.) Sacc., Syll. 6, 1888, 740.
-3 cm. high, bushy, fuscous or greyish, pale: stem straight, villous at the base:
branches short, tense, frequently branched below.
? on wood, S. America (type}: on pine-bark (France).
Spores 4-5 X 3-4µ, white, obovoid, 1-guttulate (Bourd. et Galz.).
Basidia 18-24 X 5-6 µ (Bourd. et Galz.).
Hyphae 2-3 ft \Yide, distantly septate, mostly thick-walled, parallel (Bourd. et Galz.).
PTERULA
Lloyd's figure shows a monaxial fruit-body, c. 2·5 cm. high, with short and simple
or sparingly branched laterals. He suggests that P. tenuissima Lev. and P. humilis
Speg. (p. 535) may be the same, but I am unable to trace the descriptions of either
species. Bourdot and Galzin record the species for France, but I think the identification
requires explanation. P. juruensis may be the same.
P. tenerrima (P. Henn.) comb. nov.
Basinym: Typhula tenerrima P. Henn., Hedw. 1897, 195.
Synonyms: ? Typhula tenerrima Speg., An. Mus. Nac. Buenos Aires, 19, 1909, 280.
? Typhula tucumanensis Speg., ibid.
15-20 X 0·035-0·05 mm., filiform, subulate, drying rufous brown: base thickened,
fuscous, subsclerotioid.
On fallen leaves: Brazil (Sta. Catherina).
Spores 3·5-4µ, white, subglobose.
The description of T. tenerrima Speg. is:
3-5 X 0·05 mm., white, filiform, straight or slightly curved, acute, sometimes
slightly fuscous at the inserted base: spores 6-8 X 2·5-3µ, white, elongate-ellipsoid, with
one large gutta: basidia 25-30 X 10µ, clavate, with 4 sterigmata: on fallen rotting
leaves of Chamissoa altissima (Amaranthaceae): Argentine (Tucuman).
The description T. tucumanensis is: fruit-body 20-30 X 0·2-0·25 mm., erect, simple,
filiform, very acute(? white): sclerotium 0·5 mm. wide, very small, flattened, subdiscoid,
black, surrounded with a narrow white border.
On dead leaves of Bignonia: Argentine.
Spores 6-8 X 3-4µ, white, ellipsoid, with I large gutta.
Basidia 30 X 8-9µ, with 4 very short sterigmata.
Until the type-specimens have been studied in microscopical detail and more
numerous collections are known, it is impossible to regard these three species as more
than a simple epiphyllous Pterula with fuscous levigation at the base of the stem, as
occurs in P. debilis, P. epiphylla, and P. typhuloides. It is their sole distinction from
P. Gordius unless P. •enerrima has really subglobose spores (Henning's spore-data
being often erroneous), and the other two have a large gutta in the spore.
P. tenuissima (Curt.) comb. nov.

Basinym: Typhula tenuissima Curt., Sill. Journ. 2nd ser. 6, 1848, 350.
-14 X 0·25 mm., gregarious, setaceous-filiform, simple, smooth, pallid white (or
wholly pallid fuscous, except the white acerose tip, when sterile): stem fuscous-black,
1--6 mm. long, not expanded at the base.
On dead leaves: U.S.A. (S. Car., Society Hill, Aug.-Sept., on Phaseolus, type),
Cuba (Wright, 563), Venezuela (No. 216, herb. Berk.).
Spores 9-11 X 5 µ, white, smooth, subcylindric, blunt, apiculate.
Basidia in a more or less continuous hymenium, immature, reaching 17-20 x 9-10µ.
Cystidia none.
Skeletal hyphae 2·5--6µ wide, the walls 0·5-1·5µ thick, aseptate, unbranched,
apparently unlimited, the lumen rather wide: generative hyphae clamped.
Stem without caulocystidia, but covered with a pavement-like epithelium (1 hypha
thick) of irregular, rather short, contiguous cells, 4-7µ wide, with pale fuscous, sinuous,
agglutinated walls: (some fruit-bodies wholly sterile and, then, covered to the growing
tip with this epithelium).
I have examined the type-collection at Kew. The collections from Cuba and
Venezuela, added by Berkeley, are identical except that the hyphae are slightly
narrower (skeletals 2-4µ wide, epithelial hyphae on the stem 2-4µ wide). The
species is intermediate between P. subulata, without an epithelium on the stem, and
P. typhuloides with a thicker epithelium. The spores on all three collections were very
few, but corroborative.
PTERULA 525
P. timorensis Torr.
Brot. 9, 1910, 89, f. 4.
2-3 cm. high, caespitose or densely gregarious, cartilaginous-tough, drying reddish
brown: stem 0·5-2x 1-2 mm., very short: branches 0·5 mm. wide, filiform, often
membranous at the axils, subulate, very numerous, often connate.
On tree-bark: Timor.
Spores 5-6 X 3-4µ, slightly ochraceous in the centre, smooth, ovoid-ellipsoid.
Basidia 14-18 X 3-4µ, clavate.
Hyphae 3-4µ.
Killermann places this in the subgenus Phaeopterula, but the brown colour of the
spores seems to be in the cytoplasm and may be only an effect of drying.
P. togoensis P. Henn.
Engl. Jahrb. 23, 1897, 543.
-5 cm. high, repeatedly branched, drying reddish brown and cartilaginous: branches
compressed, fasciculate, strict, slender, filiform, glabrous, tips -2 cm. long and subulate.
Spores?
On rotten wood: W. Africa (Togo).
This description would apply to any bushy Pterula.
P. typhuloides sp. nov.

Fruit-bodies-3·5 cm. high, 0·5-1·3 mm. wide, solitary or gregarious, not fasciculate,
simple, filiform-subulate or with I-2 branches shorter than, or as long as, the axis:
stem 2-4 mm. long, sterile, fuscous, puberulous: fertile part cylindric, tapered at the
apex to the filiform sterile tip, white becoming pale yellowish ochraceous from the base to
the middle, the tips white: flesh tough, white.
TEXT-FIG. 232. Pterula t_iphuloides: sclerotioid surface of the stem, :< 1,000.
On dead petioles and inflorescences (? of Pometia alnifolia: Sapindaceae) in swampy

forest; on dead leaves of Scindapsus (Araceae): Malaya (Singapore: Selitar Reservoir,
6.3.43: Gardens Jungle, 12. r 1.44).
Spores 9-10 x 4-5µ, white, smooth, narrowly ellipsoid, with a short blunt apiculus
o·8 µ long, thin-walled, aguttate.
Basidia 24-30 X 7-9µ, subclavate, vacuolate: sterigmata 4, 4-5µ long.
Cystidia none.
Hymenium not thickening, decaying from beluw upwards on the fertile part of the
526 PTERULA
fruit-body: subhymenium very narrow, 5--7µ thick, composed of closely interwoven
generative hyphae 1-3 µ wide, seated directly on the longitudinal hyphae, with short
cells 8-15µ long.
Caulocystidia -50 x 5-8 µ (at base), subcylindric to conical, mostly acute, or some
times obtuse, very abundant, with slightly thickened, smooth, pale brown walls,
sometimes with a minute hyaline point, occasionally 1-septate (without a clamp).
Stem covered by a thin sterile and abortive hymenium or merely an incipient sub
hymenium 1-2 hyphae thick, the cells contiguous, 4-8 µ wide, and their outer walls
s
slightly thickened, o · µ, agglutinated and paleJuscous-brown, more or less gyrose in surface
view, the caulocystidia innate and with many transitions to incipient protuberances
from the gyrose cells.
Hyphae dimitic: skeletal hyphae 3-5µ wide, with slightly thickened walls 0·5-1·5µ
thick, aseptate, lumen generally wide, of unlimited growth, unbranched, often kinked
or nodose, especially near the base of the fruit-body: generative hyphae 1·5-3·S µ,
wide, without clamps, cells 30-150µ long, shorter near the surface, thin-walled, or
slightly thick-walled, occasionally with intercalary skeletal portions 150-300 µ long.
var. minor var. nov.

Text-Figs. 226, 233, 234: Plate 16.
-II mm. high, simple, filiform, scattered; stem 1·5-3 mm. x 200-250µ, -300µ at the
base, pale fuscous-brown: fertile part 100-140µ wide, with a long, sterile, tapering
apex 10-15µ wide at the tip, occasionally wholly sterile or with the hymenium irregu
larly and patchily developed, pallid white or pale yellowish.
TExT-FIG. 233. Pterula typhuloides var. minor, X 1,000.
On dead grass stems and lt>af-sheaths: Malaya (Singapore, Botanic Gardens,

19.2.43).
Spores 8-10X3·5-4·5µ.
Basidia 19-23X7-9 µ.
Hymenium not thickening: subhymenium very narrow, 5--7µ thick, only 2-3 hyphae
thick, short-celled.
Cystidia none.
Caulocystidia generally blunt.
Sterile agglutinated hymenium on the stem, and hyphae, as in the typical form, but.
the generative hyphae with clamps at some septa and not at others.
The noteworthy peculiarity of this fungus is the agglutinated, fuscous-brown,
sterile hymenium covering the stem. In surface-view the hyphae are short-celled and
gyrose with slightly thickened brown walls, like the surface-view of a Typhula
sclerotium, but the walls are neither undulate nor mucilaginous and many of the cells
PTERULA 527
project as caulocystidia. The layer develops as a rudimentary, incipient hymenium
and slowly becomes transformed from below upward.
The fruit-bodies appear to grow slowly and are long-lived, lasting in the mature
state for 2-3 weeks. The tip never becomes fertile. I found extremely few spores on
the fruit-bodies during the day but obtained fairly copious spore-prints overnight.
·,,'\
ToxT-FIG. 234. Pterula typhuloides var. minor: sclerotioid surface of the stem (above, in
(surface-view), X 1,000.
The absence of clamps is noteworthy.. I carefully examined the fruit-bodies on two

occasions and could find none in the typical form but occasional clamps in var. minor.
The loss of clamp-connexion thus appears to be taking pl::i.ce in thfa species as well as
in P. subtyphuloides.
Compare P. Jructicola and P. phyllophila, which may be the same.
P. Uleana P. Henn.
Hedw. 1897, 197: Lloyd, Myc. Notes 60, 1919, 867, f. 1479.
Macroscopically as P. multifida: caespitose, much branched with filiform, repeatedly
2-3-chotomous, branches.
On bark: Brazil (Sta. Catherina).
Spores 4-4·5 X 3·5-4µ,, subglobose.
P. Vanderystii P. Henn.
FI. du Bas- et Moy.-Congo 2, fasc. 2, 1907, 96.
2-3 mm. X 120µ,, simple,. subulate, often fascicled at the base, pale cinereous,
pruinose, the subacute apices 20-40µ, wide.
Spores 5-6µ, wide, white, smooth, subglobose.
528 PTERULA
On dead wood: Congo (Kisantu).
This may be Deflexula, Mucronella, or a Pistillaroid Aphelaria.
P. verticillata sp. nov.

Text-Figs. 13, 235--9.
-6 cm. high, solitary or caespitose, monopodially branched, the main axis with 2-6
tiers of 3-4 branches, and numerous filiform lateral branches, wholly pallid drab fiesh
colour, pale yellowish drab or pale fawn-drab, the tips white, drying fuscous horny, the
hymenium shrivelling to a broken skin on the hard axis.
TEXT-FIG 235. Pterula verticillata, with apparent monopodial

branching, X 2.
Stem 3-15 X 1-2·5 mm., distinct, more or less strigose, often subdiscoid at the base,
with shortly spreading, dirty whitish fibrils attaching the fruit-body to decaying leaves
and twigs: main branches in alternating tiers, 0·5-r mm. wide, terete: branchlets
0·2-0·4 mm. wide, curved ascending, simple or dichotomous once or twice, rather lax,
tapering very gradually to hair-like tips 5-50 µ. wide (or reduced to a single hypha):
flesh rather tough, pliant, very thin.
On humus, dead twigs and leaves, in the forest: Malaya, not uncommon (Singapore
Field, Nos. 24155, 24873).
Spores 6-'] X 3-3·5 µ., white, smooth, pip-shaped, ? aguttate.
Basidia 15-26 X 5-7 µ., subclavate: sterigmata 4, 4·5 µ. long.
Cystidia absent from the hymenium: the tips of some sterile basidia often with a
PTERULA 529
single subglobose, vacuolate, spore-like body 3·5-4·5/k wide on a short sterigma-like
stalk 2-3/k long, but not abstricted.
TEXT-FIG. 236. Ptentla •1.:erticillata: an old fruit-body with attenuate tips, X 3.

Hymenium thickening to 170µ., often entirely sterile, compact, the terminal cells
of the sterile hymenium 10-26 X 4-7 µ. but lengthening 30-40µ. in old hymenia: sub
hymenium composed of compact hyphae 2·5-6µ. wide, short-celled, with slightly
mucilaginous walls, not inflating, the ends of the skeletals often curved and projecting
-30µ. into the hymenium.
Sterile hymenium on the stem and uppersides of the main branches consisting of
loose, aborti\'e, basidial branches and thin-\valled, vacuolate, subfusiform, smooth
caulocystidia -So Y 5-9 µ., the longer ones sometimes 1-3 septate and branched, the
caulocystidia abundant on the stem but very scattered on the main branches, the
sterile hymenium scarcely thickening, loosely constructed, and all .the hyphae with
slightly mucilaginous walls.
51 IQ Mm
530 PTERULA
Hyphae dimitic: skeletal hyphae 3·5-8(-r'o)µ. wide, with smooth, pale yellowish
walls 0·5-1 ·5 µ thick, occasionally branched in the mediate portion, mostly unbranched,
Tox:r-Fm. 237. Pteru.la verticillata: a, a branch showing overtopping by

unequal dichotomy: b, an abnormal fruit-body with imperfect overtopping:
c, five stages in the successive dichotomy of the apex of a tnain branch
resulting in apparently monopodial growth with a whorl of three overtopped
branchlets: d, an abnormal fruit-body with an equal dichotomy in the main
stem: X 3.
sometimes producing a cluster of 2-5 skeletals from the apex, longitudinal, flexuous,
aseptate, often kinked and of uneven width, always with wide lumen, of unlimited
growth (unbranched for lengths of 1-3 mm.): mediate parts 50-150 µ. -long, aseptate,
gradually enlarging into the skeletal and becoming thick-walled: generative hyphae
2-5 µ. wide, thin-walled, clamped, the cells 10-100 µ. long, occasionally swollen in
places to 12 µ. wide, sometimes (but by no means always) branched from the clamp:
PTERULA 531
occasionally a generative hyphae transformed, for 150-35-0µ, into a skeletal with thick
walls, then reverting to the generative form, and occasionally a generative hypha
arising from a skeletal, or a long skeletal occasionally ending in a generative hypha.
Apices of the branches composed of a sheaf of parallel hyphal tips 2-4µ wide,
eventually tapering macroscopically to a thread, microscopica1ly to a single hypha.
a.
0 .O.. ·:. o:'· o·.>· v
TEXT-FIG. 238. Pterula verticillata: the thickened hymenium, and sterile

basidia (c), X 500: a, the spores of P. bru11neola, x r ,ooo: b, the spores of
P. vinacea, x 1,000.
ToxT-FIG. 239. Pterula verticillata: sterile hymenium on the stem, X 500.

Disk-like base of the stem· sterile, at most with scattered rudimentary basidia, com
posed of generative and skeletal hyphae: the fine mycelial fibrils composed of thin or
slightly thick-walled generative hyphae 1 ·5-3µ wide, concrescent in minute fascicles.
The elegant monopodial form of this fungus is caused by overtopping of one branch
53 2 PTERULA
of a very condensed double or triple dichotomy, which gives a false whorl of 2-3
branches and the leading branch (cf. Text-Fig. 235). The successive dichotomies
alternate more or less at right angles and thus the tiers of branches alternate: there is
no evident twisting of the living axis or branches, although they are spirally twisted
when dried.
P. penicellata is similar, perhaps identical.
P. vinacea sp. nov.
Text-Figs. 238b, 240, 241.
-'] X 6 cm., bushy, multaxial or occasionally with some monopodial branches, vinaceous-
TEXT-FIG. 240. Pterula viuacea. X 2.

drab, darker vinaceous-drab at the base, pale yellozdsh flesh colour 11pzrnrds, dry and
tough: drying fuscous horny, with pallid hymenium, the branches not or but slightly
spirally twisted.
Stem 5-20 x r-r ·s mm., often shortly 'rooting', sometimes decumbent and bearing
PTERULA 533
several erect branches, sterile: main branches 1 mm. wide, dichotomous in alternating
planes, the main axis and a few main branches monopodial (by overtopping) or wholly
multaxial: branchlets 0·5 mm. wide, tapering to filiform tips, the lower dichotomies
close but not whorled, more or less alternating, occasionally palmatifid (through con
densation), ultimate dichotomies distant and equal: hymenium absent from the upper
sides of the main branches.
TEXT-FIG. 241. Pterula vinacea, showing the apparent polychotomy, x 2.

On the ground in the forest: Malaya (Perlis, Bukit Besi Hangit, 17.11.29).
Spores 7-8 X 3-3· 5 µ,, white, smooth, narrowly ellipsoid, thin-walled (? with a small
gutta).
Structure as Pterula brunneola.
P. vitellina (Pat.) comb. nov.
Basinym: Lachnocladium vitellinum Pat., Rev. Myc. 1890, 134, t. 107, f. 3.
-4 x 0·5 mm., simple, rarely dichotomous, very small, yellowish or ochrateous,
elongate, fusiform, slightly canaliculate (? effect of drying), pulverulent with spores,
apex sterile, acute: fibrous, hard (? dried).
534 PTERULA
On trunks in the forest: China (Ma Eul Chan).
Spores 9-10 X 4 µ,, yellow, sµiooth, ovoid.
Basidia 15-20 X 6-8 µ,, 2-4-spored.
This is certainly not Lachnocladium. The sterile apex and fibrous flesh suggest
Pterula rather than any other genus.
P. Winkleriana P. Henn.
Engl. Jahrb. 38, 1905, 121: Lloyd, Myc. Notes 75, 1925, 1359, f. 3216.
-3·5 cm. high, caespitose, branched, or branched-fasciculate at the base, yellowish
cinereous, cartilaginous (? dried), surrounded with white byssoid mycelium: branches
filiform, 150-180 µ, wide, mostly simple, rarely with secondary branches, often cirrate
at the apex.
On palm sheaths: Cameroons (Victoria).
Spores 5-7 µ, wide, yellowish white, smooth, globose, 1-guttate.
Basidia 15-20 x 8 µ,.
This may be Deflexula or Pterulicium.
DOUBTFUL SPECIES OF PTERULA

P. aciculaeformis Lloyd
Myc. Notes 60, 1919, 868, £: 1481: Teodoro, En. Philipp. Fungi 1937 (ut P. aciculae
Lloyd).
-10 cm. high, 0·5 mm. thick, simple, cylindric, slender, rigid, dark brown (dried): ..
sp. ? 4-5 µ,, white, globose: on wood.
Philippine Isl. (0. Reinking).
This may be only the erect rhizomorph of a horse-hair fungus (Marasmius, Crini
pellis, or Polyporoid).
P. angustata (Lev.) Sacc.

Syll. 6, 1888, 742.
Basinym: Merisma angustata Lev., Ann. Sc. Nat. 3, 5, 1846, 158.
6-g cm., shortly stalked: branches fastigiate, compressed, attenuate to the acute
· tips, glabrous, cinereous fuscous (? dried).
On wood: Mexico (Vera Cruz).
var. fodinaria Sacc., Ann. Myc. 12, 1914, 300.
The variety is clearly an abnormal, abortive fruit-body of some Basidiomycete
developed on wood of Quercus used in Mexican mines (? Lentinus). The typical
P. angustata may be the same as Clavaria Juscescens.
P. dicbotoma Sauter
Hedw. 1876, 152.
Cinereous, slender, erect: stem simple, soon divided into repeatedly dichotomous
branches, compressed: tips obtuse, white.
On rotten wood of frondose trees: Austria (Steyer, pr. Salzburg).
Lloyd regards this as doubtful. There are no specimens. It may be P. subulata or
a Lentaria.
Dendrocladium fructicolum Lloyd
Myc. Notes 61, 1919, 942, f. 1739.
Stems -3 cm. tall, brown, with a few lax dichotomous branches above, tips filiform,
axils flattened.
Spores 8 µ,, coloured, subglobose (others more elliptical 12 x 6 µ,).
On fruit of some kind: Africa O. Gossweiler).
PTERULA 535
'We cannot explain the two forms of spore we find. Perhaps one is accidental'
(Lloyd). The fungus may be a Xylaria.
P. fruticum Lloyd
Myc. Notes 59, 1919, 822, f. 1373: 60, 1919, 867, f. 1478.
-8 cm. high, 0·5-1 mm. thick, simple or with one branch, cylindric, with filiform
acute tip: dark brown (dried).
Spores 4-5 X 3 µ, smooth, subhyaline: on a fruit of some kind: W. Africa
(J. Gossweikr).
'An abundant violet colouring matter is given off in an alkaline solution' (Lloyd).
The fungus is certainly the conidial state of some Pyrenomycete.
[P. fusispora Yasuda, Bot. Mag. Tokyo 33, 1919, 297: in Japanese.]
P. humilis var. tucumanensis Speg.

An. Mus. Nac. Buenos Aires 19, 1909, 280: Sacc., Syll. 21, 1912.
Differs from the type in the long stem with distinct, more abruptly branched apex,
white, soft when fresh but tough, drying coriaceous-rigid and flesh-colour: spores
4 X 3 µ, white, ellipsoid or ovoid, 1 -guttate: on rotten branches: Tucuman, Argentine.
Saccardo says he has not discovered the name P. humilis Speg. and suggests that
Clavaria humilis Cke. may be intended. C. humilis Cke. is a Clavulina. I do not know
to what genus var. tucumanensis can belong. Lloyd suggests P. humilis Speg. is P. taxi
formis.
Since writing these notes I have examined the co-type of Pterula humilis Speg
(? ined.) in the Paris herb., namely Balansa 3348, collected from rotten wobd in ,
Paraguay. It is a true, small, branched Pterula with, apparently, a rather lax1y hairy
stem and unilateral hymenium. I could find no spores or cystidia and the fruit-bodies
do not have a brown agglutinated, or sclerotioid, surface to the stem. The branching
seems mainly adventitious, but is not bushy.
P. incisa Lloyd
Myc. Notes 65, 1921, 1057, f. 1974.
-1 ·5 cm. high, white: stem --'J X 1(-1 ·5 ?) mm., cylindric, branched above irtto
numerous filiform branches, axils flattened: spores and basidia not seen: on wood,
Philippine Isl.
Lloyd suggests Tremellodendron for this. It may be a Xyla,:ia.
P. Landolphiae Lloyd
Myc. Notes 67, 1922, 1144, £. 2184 (ut P. Landelphiae).
As P. Jruticum but not giving off a violet colour in alkaline solution: set with
projections 100-150 X 30µ making the stem scurfy: no basidia or spores seen: orange
yellow: on seed of Landolphiafiorida: Africa (Gossweiler).
Lloyd found spores 5 ><4µ, pale, smooth, but said they might not belong. It is
surely the conidial state of a Pyrenomycete, similar to P. subplumosa.
P. luzonensis Lloyd
Myc. Notes 66, 1922, 1106, f. 2026.
Deeply rooting in the ground: stem slender, rigid, terminating in a few spreading
branches: hairs and cystidia none: spores 8 µ, globose, hyaline, smooth: Luzon (Mt.
Maquiling, Reinking 105i1).
Surely the conidial state of Xylaria nigripes.
P. Mannii Lloyd
Myc. Notes 55, 1918, 793, f. 1194: 60, 1919, 867.
-12 cm. high, with a long stem (about three-quarters of the length of the fruit-bOOyJ
536 PTERULA
bearing several lax dichotomous branches with filiform tips, the main axis some
times more or less monopodial, stem becoming blackened at the base, axils flattened
but scarcely dilated, branchlets laxly dichotomous, spreading: spores 6 x 4 µ., white,
smooth: ? on wood: W. Africa (Gossweiler): on the ground (Philippine lsl.): also
a specimen in Kew herb. as Lachnocladium Mannii ms.
Surely a conidial Xylaria, cf. X. nigripes.
P. pungens (Lev.) Sacc.
Syll. Fung. 6, 1888, 741: Lloyd, Myc. Notes 60, 1919, 866, f. 1472: 63, 1920, 982.
Basinym: Merisma pungens Lev., Ann. Sci. Nat. 3, 5, 1846, 157.
Caespitose: stem elongate, unequal: branches cylindric, fastigiate, dichotomous,
glabrous, umber, with acute tips: on fallen wood: Java (Mt. Pangerango): (original
description).
The following quotation is from Lloyd:
'J'ai vu page 866 un Pterula pungens: le dessin que vous donnez est tres different du
veritable Merisma pungens Lev. Cette plante n'est pas un Pterula, ni un Lachno
cladium. Je ne sais mil;me pas ce que c'est, et je l'ai bien etudiee. C'est une grande
plante (10 cm.), rigide, tres glabre, naissant d'un subiculum membraneux. Je n'ai
pas vu de spores, pas de basides, mais je crois sous toutes reserves que le champignon
peut etre une heterobasidee. Je suis peut-etre le se:.il a posseder le type de cette
espece' (Pat.).
Possibly it is Lentaria surculus.
P. sclerodontia (B. et Mont.) Lloyd

Myc. Notes 67, 1922, 1153, f. 2247: 74, 1925, 1340, f. 3123.
Basinym: Hydnum sclerodontium B. et Mont., Hook. Journ. 1844, 333.
The Malayan specimens, so named by Lloyd, are a Tremellaceous fungus, for
which I have not found a genus.
P. subplumosa P. Henn.
Hedw. 1897, 197: Lloyd, Myc. Notes 60, 1919, 865, f. 1471 (ut P. pennata).
Synonym: P. pennata P. Henn., Hedw. 1904, 174: (teste Bres. in Sacc., Syll. 23,
1925, 496).
Hydnum Sprucei Mont., Syll. Crypt. 1856, n. 563: (teste Lloyd).
1 cm. high, yellowish flesh colour: stem terete, ochraceous tomentose, squarrose
ramose: branches repeatedly dichotomous, terete, cartilaginous, smooth, tips subulate:
spores 3·5µ., globose, hyaline: on bark: Brazil.
As figured by Lloyd, this is the conidial state of some Pyrenomycete, cf. Pterula
Landolphiae. Hennings gives for P. pennata: -5 cm. high, cinereous alutaceous:
branches erect, filiform, penniform-ramose on all sides: branchlets recurved, subulate,
simple, 1·5-3 X 0·08-0·12 mm.: spores 10-13 X 7-8 µ., white or yellowish, smooth, ovoid
or subpyriform, subangular, with a large gutta: on rotten wood: Amazon (Rio Jurua).
PTERULICIUM gen. nov.

(From Pterula and Corticium)
As Pterula, but developing a resupinate Corticium-like patch covered with a normal,
thickening, fertile hymenium when facing downward, generally accompanying the
Clavarioid fruit-bodies, sometimes separate; the resupinate patch composed of generative
hyphae without skeletals.
Clavarioid fruit-bodies freely or sparingly branched, the branching apical and
flattened in polychotomy or dichotomy, or adverititious: tough, with waxy, amphi
genous or unilateral hymenium.
On dead plant remains: 1 sp., tropical Asia and Africa.
ThxT-FIG. 242. Pterulicium xylogenum, X 2: a cluster of primordia, • X 5.
TEXT-FIG. 243. Pterulicium xylogenum, subsimple fruit-bodies, X 2.

538 PTERULICIUM
Spores white, smooth, aguttate.
Cystidia none, or as thin-walled basidioles in the young hymenium.
Hymenium thickening: young basidia subacute.
Sterile hymenium at the base of the stem and on the uppersides of the main
branches consisting of rudimentary collapsing basidial branches, loose, not as a
palisade.
Hyphae dirnitic: skeletals unlimited, aseptate, rarely branched: generative hyphae
clamped, usually branching from the clamp.
The Corticium-patch at the base of the fruit-body is normally extensive with
marginal growth, but it may be limited to a rudimentary steJiile or fertile disk,
0·5-2 mm. wide, when it appears as the sterile discoid attachment so common in
fungi which grow on plant remains: rarely is it absent. In other cases the Clavarioid
fruit-bodies are rudimentary, as minute conical primordia, and only the Corticium
patch develops fully: occasionally the Corticium-patch develops without trace of the
Clavarioid primordia, and such forms appear as Corticium sui generis. The fertility of
the patch seems to depend on its position. It is always covered by a thickening
hymenium which is sterile when facing upwards and fertile when facing down, the
basidia and spores being identical with those of the Clavarioid fruit-bodies.
P. xylogenum (B. et Br.) comb. nov.
Basinym: Clavaria xylogena B. et Br., Journ. Linn. Soc. Bot. 14, 1874, 76.
Synonyms: Pterula xylogena (B. et Br.) Petch., Ann. R. Bot. Gard. Per. 9, 1925,
332 (ut P. zylogena).
Pterula simplex Sacc. et Pao!., Att. R. Inst. Ven. 6, Ser. 6, 1888, 15, t. 5, f. 7; sensu
Lloyd, Myc. Notes 60, 1919, 866, f. 1473: 'an vera?
Pterula subaquatica Bres. et Roum., Rev. Myc. 1890, 36, t. 92, f. 7.
TEXT-FIG. 244. Pterulicium xylogenum: irregular polychotomy after attenuate flattening in

young fruit-bodies re mm. high: X 15.
PTERULICIUM 539
Pterula importata P. Henn., Verh. Bot. Ver. Prov. Brand. 40, 1898, 121, t. 1, f. 1:
Kew Bull. 1917, 13.
[? Pterula subsimplex P. Henn. (see under Pterula, p. 522).]
0·8-5 cm. high, solitary or fasciculate, often densely, mostly sparingly branched,
well-grown specimens copiously branched, occasionally simple, white, often becoming
wholly pale livid flesh-colour or very pale vinaceous drab, tips pallid white, then yellowish
or Juscous-brown from the base upwards, drying wholly white pruinose: generally arising
from a waxy crustaceous resupinate patch -3 cm. wide x 100-200µ. thick, separable,
even, minutely pruinose, white then pallid alutaceous or subochraceous drab, margin
pallid white, minutely pruinose-crustaceous, or in full growth, minutely arachnoid,
generally determinate, sometimes with only a small sterile disk, 0·5-2 mm. wide, at the
base of the fruit-body.
TEXT-FIG. 2+5. Pterulicium xylogenum: diagrams of the structure of the fruit-body: above, as
developed from a fertile resupinate patch facing downwards, the Clavarioid prirn ordium
abortive and sterile: beloic, as developed from a sterile resupinate patch facing upwards, the
Clarnrioid fruit-body normal: note the absence of skeletal hyphae (shown as heavy lines) from
the resupinate patch.
Stem 2-15 x 0·2-0·5 mm., dividing into 2-6 branches at the apex, or, occasionally,
continued monopodially (by overtopping) with 1-3 lateral branches: branches Jew,
terete, divaricate, often connate where overlapping, elongate, lax, simple or with adventi
tious acicular branchlets zdth long filifarm acute tips 15-40 µ. wide at the apex, axils
acute, rarely flattened, tips sometimes recurved: hymenium on all sides of the upright
branches, generally only on one side of the stem and proximal parts of the main
branches, or absent from the base of the stem, the sterile uppersides often faintly
longitudinally sulcate, pruinose, extending to the base of the stem and covering the
whole resupinate patch: flesh white, becoming fuscous-brown from the base upwards,
fibrous-subcoriaceous, dry: without smell.
Gregarious or subcaespitose, rarely solitary, often in rows on dead wood, dead
540 PTERULICIUM
bamboo-culms, and dead leaf-sheaths and trunks of various palms (Pinanga, Areca,
Oncosperma, Ptychosperma): Ceylon, Malaya, Philippine Is!., Uganda, (? tropical
America).
TEXT-FIG. 246. Pterulicium xylogenum: generative hyphae (left)

branching from the clamp, skeletal hyphae (one on the right being
secondarily septate), and a thick-walled generative hypha (extreme
right): X 1,000.
Spores n-13 X 6-7·5µ, (12-15 x 6-7 µ,, Petch), white, smooth, elongate amygdaloid,
unequal-sided, generally subacute at the apex, very obliquely and bluntly apiculate at the
base, slightly thick-walled, contents finely granular-clouded with a few small, faint
vacuoles.
Basidia 35-65(-70) X 10--13µ,, clavate, contents dense, subgranular, with a few large
vacuoles, immature basidia pointed: sterigmata 2-3-4, mostly 4, 6-7 µ, long.
Cystidia none, or thin-walled, subventricose, vacuolate, smooth, -40 X 5-8µ, in the
young hymenium.
Hymenium c. 30µ, thick at first, thickening -200µ, (or more): subhymenium com
posed of contiguous, closely interwoven, short-celled, thin-walled, uninflated hyphae,
cells 5-30 X 2-5 µ,.
PTERULICIUM 541
Hyphae dimitic: skeletal hyphae 2·5-5(-6)µ wide, with smooth walls .-:,·5-2µ thick,
colour becoming pale brownish in the older tissue, aseptate or occasionally secondarily
septate, rarely branched, sometimes kinked, apparently of unlimited growth, arising
directly from the generative hyphae, strictly longitudinal: generative hyphae 2-4(-5) µ
wide, mostly thin-walled, occasionally becoming rather thick-walled, septate with
clamps, branching from the clamp and often also from other parts of the cell in the
subhymenium (rarely in the flesh), smooth, colourless, H-connexions occasional:
with occasional intermediates between skeletal and generative hyphae.
TEXT-FIG. 247. Pterulicium xylogenum: the thickening hymenium,

X 1,000.
Growing tips of the branches 20-40 µ wide, consisting of a tapered shea(of genera
tive and skeletal hyphal tips, widening behind the apex by the branching and develop
ment of new hyphae and by the development of the hymenium 100-1,000µ from the
apex.
Sterile hymenium on the stem and uppersides of the branches as a rudimentary
542 PTERULICIUM
hymenium with abortive basidial branches 1·5-6 µ. wide, not thickening and without
cystidia.
Flesh of stem and branches consisting of a homogeneous fibrillose tissue of longi
tudinal generative and skeletal hyphae, without interweaving hyphae.
Resupinate patch constructed, like the hymenium, of generative hyphae, without
skeletal hyphae, consisting of a basal layer 20-50 µ. thick, of radiating generative hyphae
parallel to the substratum and with slightly thickened and agglutinated walls, the layer
thinning off towards the margin, and a hymenium, with narrow subhymenium, thicken
ing -150 µ. exactly as on the branches, the hymeniumfertile when facing downwards but
ster.ile when facing upwards (though normally developed).
Colour of the fruit-body in the brownish walls of the skeletal hyphae, perhaps also
in the pale alutaceous or pale flesh-coloured cytoplasm of the basidia.
This description is drawn entirely from Malayan specimens, which occur commonly
on dead remains of palms and bamboos in the Singapore Botanic Gardens. Petch's
description from Ceylon agrees except in the slightly longer spores, as noted, and the
habitat, which is dead wood, as well as dead bamboos: the existence of the Corticium
patch was omitted, either because it was rudimentary or, as might easily have happened,
because it was overlooked. Dried Clavarioid fruit-bodies of Petch 4310, in the Kew
herbarium, are identical microscopically with the Malayan.
P. simplex, as figured and interpreted by Lloyd, is certainly identical, but P. simplex
was described originally as fuliginous-black and seems to resemble the incipient
rhizomorphs of a horse-hair fungus. It was described from Malaya as growing on the
inside of dead bark (separating from the wood), and without spores. Lloyd says that
P. importata (sp. 12-14 x 8 µ.), P. subsimplex from Brazil, aFld P. subaquatica from
Africa (sp. 12-13 X 6 µ.) are very close, if not identical. I have kept P. subsimplex apart,
under Pterula, because it seems to have different spores.
The collection (Dtimmer II85, Uganda) in the Kew herbarium, identified as P.
importata by Miss Wakefield, is probably identical with P. xylogenum. It consists of
small dense tufts of simple or sparingly branched Pteruloid fruit-bodies 1-1·5 cm.
high on dead bark: spores II·5-14 x 7-8 µ., as in P. xylogenum, but squarish with
rounded angles in optical t.s. (? effect of drying): basidia 50-60 x 10 µ.: skeletal hyphae
3-5 µ. wide. There was no trace of a Corticioid patch, possibly because the fruit
bodies grew on the upperside of a fallen log.
RAMARIA S. F. Gray emend. Donk

Nat. Arr. Br. Pl. 1, 1821, 655: Bonorden, Handb. Myk. 1851, 166: Quel., FI. Myc.
1888: Ricken, Vadem. 1918 et 1920: Huber, Zeitschr. f. Pilzk. 15, 1931, 109, et ibid.
18, 1934, 34: Donk, Rev. Niederl. Homob. Aphyll. 2, 1933, 103, et Bull. Bot. Gdn.
Buit. Ser. 3, 17, 1941, 177: Maire, Fungi Catal. 2, 1937, 31.
Synonyms: Cladaria Ritgen, Sehr. Marb. Ges. 2, 1831, 94.
Clavaria sect. Ramaria Fr., Ep. 1838, 571, pr. p.
Clavarielfa Karst., Rev. Myc. 3, 1881, 21: Schroet., Cohn Krypt.-Fl. Schles. Pilze
1888, 447.
Phaeoclavulina Brinkm., Jahrb. Westfal. Prov. Ver. Wiss. Kunst., Bot. Sect. 1896-7,
197: v. Ov., Bull. Jard. Bot. Buit. Ser. 3, 5, 1923, 276.
Lachnocladium sect. Coniocladium Pat. et sect. Dendrocladium Pat., Journ. de Bot. 3,
1889, 23, 33: Ess. Tax. 1900.
Dendrocladium Lloyd, Myc. Writ. 5, 1919, 870 (see p. 82).
Fruit-bodies massive to small, radially branched, polychotomous or dichotomous,
rarely with flattened branching (R. gracilis), generally coloured, pailid wJ1ite in some
species: flesh brittle, fleshy-fibrous, softly coriaceous, tough or gelatinous, often
vinescent or rufescent (cyanescent in R. ochraceo-virens).
Terrestrial in humus or lignicolous (Stricta-group): cosmopolitan, 97 spp.
RAMARIA 543
Spores pale yellow, ochraceous, cinnamon or ferruginous, the spore-wall coloured,
ellipsoid, large to small, smooth, striate, rough, rugulose, verruculose or echinulate,
generally with one to several guttulae, aguttate in a few species: (subglobose in a few
doubtful species).
Basidia 4-spored (1-)2-3 spored in a few species, not secondarily septate, usually
multiguttulate: sterigmata erect, straight, or slightly curved.
Cystidia and gloeocystidia none (? R. cystidiophora).
Hymenium thick�ning or not, often dorsiventral and sterile on the uppersides of
the branches: subhymenium composed of uninflated hyphae.
Hyphae monomitic, 1 clamped, rather long-celled, not secondarily septate, the walls
thin or slightly thickened, strongly thickened in the Stricta-group, not incrusted:
longitudinal, some species with narrow interweaving hyphae.
{.,,
' ,<;,
\,
i
t"
'.��. .
' ,,
(
TExT-FIG. 248. Ramaria-spores: above, R. botrytis: centre, R.Jormosa:

below, R. fiava (collection from Rekawinkl, Austria): X 1,000.
Mycelium white, or pallid yellowish in the Stricta-group.
Colour chiefly in oil-guttulae in the hyphae and basidia: often acrochroic.
Lecto-type: R. botrytis (Fr.) Rick.
Ramaria and Clavulinopsis are the two main genera of unspecialized Clavarioid
fungi. Ramaria embraces the largest fruit-bodies and, in this respect, is one of the
finest genera of Basidiomycetes. The species have been much collected in recent
years and, as their spores are often large and characteristically marked, they have been
more accurately described than those of other Clavarioid fungi. The result has been
the accumulation of many macroscopic descriptions, paintings (mostly inaccessible),
and spore-records, particularly for the common species, but instead of clarifying the
identification of the species they have rather obscured them, either through mis
identification or through the ·unexpectedly great variation in form, colour, and size of
the fruit-body and of the spores. The tropical species, moreover, have been woefully
hidden in Lachnocladium. I turned to Ramaria with a light heart, after the per
plexities of the smaller 'Clavarias', only to find the hardest task of all. I spent many
weeks in transcribing descriptions and as many more in digesting, assembling, and
' Dimitic, with skeletals, in the secondary mycelium of R. stricta.
� r.,-
TEXT-FIG. 249.
;JI
r:
l�\ �9·, l;;�
�(Q)1 ,{·(()_J 1�1j1 ·'.:�::J t:. 1i.@Jw
...,/ � 'L, I�
Ramaria-spores: a, R.flavo
d_
alba: b, R. stricta: c, R. subgelatinosa: d, R.

fennica: /- 1,000.
TExT-FIG. 250. Ramaria-spores: a, R. fragillima (two collections): b, R. longicaulis

(Malaya, 20.10.30):c, R. lnvalii (two collections): d, R. Zippelii (large-spored state):
X 1,000.
RAMARIA 545
rearranging the data before I could be in any way satisfied with a system of classifica
tion. I have, since, revised the whole on two occasions; firstly to transform Clavariella
into Ramaria, on Donk's advice, and secondly to assimilate the new data of Fawcett's
from Victoria (1939-40) and of Doty's from Oregon (1944). Still there are many
doubts which can be solved only by continued study of living specimens, but I think
one can look forward to progress, at least in the little-known parts of the world, in the
Andes, the Himalayas, and the mountains of Africa, where many new species must
await discovery. The genus holds, unquestionably, primitive fungi of great interest to
evolutionary mycology.
In the following key, the basis of which is explained on pp. 136-41, I have
sought by means of nine headings quickly to bring to mind the content of the group.
An' acquaintance with the common species, about which the others are arranged, is
thus essential. The main groups are not sharply delimited because of several 'bridging
species', and in identifying a species it will always be necessary to go through one or
more adjacent groups: thus a Verrucispora must be compared with Stricta, a Flava
with Formosa, and vice versa: the least satisfactory, as featureless, is Decolorans.
Such double, or triple, effort will be light compared with the previous task of identify
ing, say, a pale ochraceous Ramaria from Africa.
Edible Species
R. aurea, R. botrytis, R. botrytoides, R. cyanocephala, R. Jennica, R. fiava, R.
fiavobrunnescens, R. holorubella, R. Invalii, R. obtusissima, R. paludosa, R. rufescens,
R. Strasseri, R. stricta, R. Zippelii.
Poisonous, though not deadly, are R. formosa and R. Mairei.
KEY TO THE SPECIES OF RA.MARIA

Key to the Groups of Species (Species Incertae Sedis, p. 553.)
Sp. echinulate
Fruit-bodies massive to slender, generally deep ochraceous,
brownish, ferruginous, umber, or fuscous: flesh slightly
tough, often discolouring vinous or pink, often acrochroic:
spores fuscous ferruginous to brownish ochre: hymenium
generally dorsiventral on the main branches, generally thicken
ing: basidia with (1-)2(-3) sterigmata in several species:
terrestrial, rarely lignicolous Echinospora
Sp. verruculose, short, 5-10(-II) X3-5µ.
Fruit-bodies small to medium-size, often slender, pallid white,
pale tan, or cream to ochraceous or brownish, rarely umber,
with lavender tips in R. subdecurrens: flesh slightly tough,
rarely discolouring vinous: sp, ochraceous: hymenium often
thickening slightly, in some unilateral: basidia · 4-spored
(2-spored in R. luteofusca): terrestrial Verrucispora
Sp. rough or smooth: basidia 4-spored
Lignicolous or terrestrial, with abundant mycelium at the base
of the fruit-body: hyphae thick-walled (0·5-1·5µ.) (? in R.
crispula): sp. short 4-11 X2·5-5µ.: fruit-bodies small to
medium-size, pallid white to yellow, ochraceous or brownish,
tips deeper yellow in R. stricta: flesh tough, vinescent in R.
stricta: sp. ochraceous, minutely rough or smooth: hymenium
generally unilateral and thickening Stric(a
SI 19 Nn
RAMARIA
Terrestrial: hyp hae with thin or very slightly thickened walls:
sp. large or small: flesh often vinescent or rufescent: mostly
medium-size to large Eu-Ramaria
Sp. smooth, faintly longitudinally striate: branch-tips reddish
purple or vinaceous Botrytis
Sp. not striate (or, if so, then also rough)
More or less pink or reddish, often yellowish pink or pinkish
orange; tips concolorous, yellow or deeper pink: flesh
brittle, softly coriaceous or gelatinous: often vinescent . Formosa
Bright yellow, ochraceous, or orange, rarely tinged pink:
flesh in some cases rufescent or vinescent, subgelatinous
in the heads of R. capitata Flava
More or less intense violet, at least on the stem, or yellowish
with the branches suffused lilac or purple Violacea
Pallid white, yellowish, pallid ochraceous, tan or pallid
brownish, in some species deeper cinnamon-brown with
age, or suffused lavender-pink or purplish towards the
concolorous or yellow tips: sp. often smooth Decolorans
Echinospora
Sp. 10-20µ long, spines 1-3µ, basidia 2-4-spored: generally mas
sive, the flesh often vinescent
Tips blue: brown-ochre to umber, large: trop. R. cyanocephala
Tips red, rufescent, or orange-ochre
Trop.
(Orange with red tips: sp. 16-17 X 8-9µ, ? verrucose:
S. Amer. R. apiahyana)
Brownish with orange-ochre or rufescent tips: sp. 9-16 X
4-'7'5µ: (2-3-)4-spored: trop. Asia R. fragillima
Temp.
2-spored: vinescent-nigrescent R. nigrescens
4-spored: flesh unchanging var. americana
Tips white
Temp.: deep brick-brown: N. Amer. R. grandis
Trop. Asia
Lignicolous, whitish: Java R. albida
Terrestrial: ochraceous to brownish: smell strong R. Zippelii
Tips?
Yellow: branches hollow (on drying): S. Amer. R. tubulosa
Colour?: branches slender: Guadeloupe R. guadelupensis
Sp. -IIµ long, spines 0·5-1µ: (2-)4-spored: flesh usually un
changing: tips concolorous or paler: small to medium
Sp. 8·5-9·5 X 3-4µ: ferruginous with brown tips: stem long,
slender, flesh-colour: U.S.A., Spain . R. Murrillii
Sp. wider, 7-II X 3-6µ
Under conifers: ochre, then brownish: flesh unchanging: N.
temp. R. Invalii
RAMARIA 547
Under frondose trees: often vinescent
Cinnamon-tawny, then darker brown: mycelium and stem
vinescent, or pale rufescent, sometimes the flesh also:
U.S.A. (? trop. Asia) R. longicaulis
Ochraceous to dingy brown or fuliginous, slender: flesh
sometimes unchanging: trop. Asia R. Zippelii var.
gracilis
Densely caespitose, -10 cm. high: colour ?: French Guiana R. guyanensis
(Sp. 8X6p,: on rotten wood: Dominica . R. cervicornis)
(Tips yellow, branches umber, 4 cm.: sp. verrucose: Spain R. luteo-fusca)
Verrucispora
Tips lavender-pink
Pallid ochraceous, -4 cm. high: sp. 5·5-7 ·5(-9) X2·5-3·5(-4·5)µ:
under conifers: U.S.A. R. subdecurrens
(Cinnamon-brown, -10 cm. high: sp. 7·5-12X4-5·5µ: in mixed
woods: U.S.A. R. subspinulosa)
Tips yellow or paler concolorous
Flesh vinescent
-5 cm. high, cream-white or yellowish: sp. 4-6X2·5-3·5µ:
under conifers: U.S.A. R. pusilla
-IO cm. high, buff-yellow to deeper honey-yellow: in frondose
woods: U.S.A. var. australis
(Umber on bruising: tips green: compact: sp. 9-12x4µ:
U.S.A. R. testaceo
viridis)
Not vjnescent
Greenish on bruising: -4·5 cm. high, very compact, ochra
ceous: sp. 6-9x3-4·5µ: under conifers: N. temp. R. ochraceovirens
Not virescent
Tips yellow, branches umber: 2-spored: sp. 8-10X 3-3·5µ:
in fields: Spain R. luteo-fusca
Tips concolorous or paler: 4-spored
Smell pleasant, of aniseed
(-8 cm., pallid then ochraceous or pale cinnamon:
branching often flattened: sp. rough or nearly
smooth : under conifers R. gracilis)
-3 cm. high, yellow-ochraceous: sp. 6X3µ: Indo-china R. capucina
Smell not of aniseed, or none
Sp. 3·5-5x2·5-3·5µ: -4 cm. high, ochraceous: under
conifers: Pacific N. Amer. R. myceliosa
Sp. larger
Trop. Asia: -4·5 cm. high
Ceylon: white, then yellow-brown, sometimes
purplish: mycelium floccoso-membraneous: sp.
5-7 X 3-5µ: on humus and wood R. megalorhiza
Sumatra: yellowish or pale ochraceous, clearer
yellow near the tips, with slender rhizomorphs:
sp. 5-6·5X3-3·5µ R. luteo-fiaccida
RAMARIA
N. temp., S. Austral.
Sp. 8·5-II x4µ: dark brown: U.S.A. R. subfennica
Sp. smaller
-6 cm. high, pallid then deep ochraceous or
brownish: sp. 5-8 X3-4µ: usually under coni-
� - Rfe�
-9 cm. high, pale ochraceous: sp. 7 X4µ: under
frondose trees R. eumorpha
-4 cm. high, yellow-ochraceous: sp. 5·5-7 X3-4µ:
on Dicksonia-stems: Victoria R. jilicicola
(on dicotyledonous wood: -3 cm. high, pale ochra-
ceous: sp. 5-7·5x3-4µ . R. crispula)
Stricta'
Terrestrial: spores verruculose-rough or rugulose
Not fragrant: (compare Verrucispora)
Sp. rugulose-rough, aguttate, 6-8 X 4-5 µ: Malaya R.polypus
Fragrant: sp. verruculose-rough: branching often flattened
-8 cm. high, white, clay, tan, then pale ochraceous or pale
cinnamon: mycelium floccose, abundant: sp. 5-7 X 3-4 µ,
guttate: under conifers: N. temp., S. Austral. R. gracilis
Similar but with palmate branchinis: Europe R. palmata
(Pinkish, -10 cm. high: sp. 7-11 x2·5-4·5µ: in coniferous woods R. suecica)
Lignicolous
Sp. 7-11 X4-5 ·5 µ, minutely rough or nearly smooth: temp.
Vinescent or browning on bruising
Tips clear yellow: branches pallid yellow to ochraceous, then
brownish, fastigiate, parallel: Europe, Japan R. stricta
Stem tinged violet: France var. violaceo
tincta
Tips concolorous or paler
Wholly cream-white: Scotland var. alba
Pale fleshy tan or pale ochraceous, then cinnamon-brown:
U.S.A., S. Austral. var. concolor
Dingy fuliginous brown: U.S.A. var.fumida
Not vinescent: usually on coniferous wood
-7 cm. high, cream or pale pinkish ochraceous, then deep
ochraceous or brownish: tips paler, then concolorous or
greenish: branches rather lax: N. temp. R. apiculata
-5 cm. high, pale cream ochraceous, tips whitish: branches
very crowded, short: France var. compacta
Sp. 6-9X4-5µ; trop. (see below)
Sp. smooth (or punctate rough): -10 cm. high, caespitose, pale
yellow or ochraceous, much branched: Old World tropics . R. Moelleriana
Sp. rugulose-rough, aguttate, cinnamon-buff: -8 cm. high,
whitish then pale yellowish, flesh colour or dingy ochraceous,
tips whitish: branches abundant, lax, curved ascending: on
rotten wood, palm-stems, and in humus: Malaya R. polypus
1
Compare Lentaria with narrow, elongate spores white or pale ochraceous (? on drying),
8-18x3-6µ.
RAMARIA 549
Sp. 4-7 X 3-4µ: on dicotyledonous wood
Sp. smooth: -6 cm. high, yellow-brown: E. Africa R. nguelensis
(Compare R. Moelleriana, foregoing)
Sp. rough
Europe, N. Africa, S. Austral.
-9 cm. high, pale yellowish, then ochraceous, tips white:
branches fastigiate: sp. 4-6x2·5-3·5µ: Europe, N.
Africa R. Bourdotiana
-5 cm. high, cream or tan, then ochraceous, tips con-
colorous: divaricate: sp. 5-7·5 X 3-4µ: Europe, S.
Austral. R. crispula
Africa: much branched
-6 cm. high; sp. 5-5·5x3-3·5µ: Congo R. ochracea
-2·5 cm. high, flesh-colour, fastigiate: sp. 6 X 3µ R. durbana
(Ceylon: -4·5 cm., white then yellow-brown, or purplish:
sp. 5-7 X 3-5µ, verruculose R. megalorhiza)
Botrytis
Sp. 12-20 X 4-6µ: pale alutaceous, tan, or ochraceous, tips deep
pink to purple, usually massive, congested: Europe, N. Amer. . R. botrytis
Sp. 10-13(-15) X 3-5(-6)µ: as R. botrytis but the tips vinaceous:
? vinescent on bruising: N. Amer., S. Austral. R. holorubella
(Sp. 8-10 x 3·5-4µ: pallid, then somewhat ochraceous, tips rufous
or pinkish: Europe R. rufescens)
Formosa
Flesh more or less gelatinous: not vinescent
Without· clamps: slightly gelatinous: pale orange-yellow: sp.
7-8·5 X 4-5µ, rough: Italy R. ignicolor
With clamps
Flesh wholly firmly gelatinous, translucent: cream-white, then
pinkish or brownish: sp. 7·5-1ox4·5-6µ, rough: U.S.A. . R. gelatinosa
Subgelatinous, not translucent: apricot-pink with honey
yellow tips: sp. 8·5-10·5 x4·5-5µ, rough: Malaya R. subgelatinosa
(With capitate, viscid tips: Austral. R. capitata)
Flesh softly coriaceous (drying chalky friable): not vinescent
Pinkish fading pallid ochraceous: base tomentose with abundant
mycelium: sp. 7-11 X2·5-4·5µ: mostly in coniferous woods:
N. temp. R. suecica
Flesh more or less brittle, sappy, firm
Tips expanding into subglobose knobs or capitate, viscid heads,
becoming cauliflower-like: S. Austral.
(Heads viscid, capitate, fusing in tiers: yellow or tinged
orange: sp. 11-13 x4·2-5µ, rough R. capitata)
Tips knob-like, not viscid, not fusing: yellow to pinkish
orange, tips often yellower: sp. 8-10·5 X4-5µ, rough R. ochraceo
salmonicolor
550 RAMARIA
Tips not expanded
Tips clear, bright yellow: often vinescent
Stems rather slender, connate-caespitose: sp. smooth, or
nearly: salmon-yellow fading cinnamon-buff: rather
tough: U.S.A.
Sp. 7-10X4-5µ: stems 1·5-2·5 mm. wide . R. conjunctipes
Sp. 5·5-6·3 x4-4·8µ: stems -8 mm. wide: fragrant var. odora
Stem massive, distinct: vinescent
7-30 cm. high, pinkish buff to orange-rose: flesh drying
chalky-friable: sp. 8-15X 4-6µ, rough: N. temp., S.
Austral. R. formosa
Similar but rose-orange with greyish violet or fuscous
branches: France R. Bataillei
Similar but small, -5 cm. high, pale cinnamon-rufescent:
Europe R. testaceofiava
(Sp. 6·5-9·5X3-4µ, nearly smooth: not vinescent: N.
temp., S. Austral. R.fiavo-brun-
nescens var.
aurea)
Tips not yellower than the branches: rarely vinescent (R.
brunnea, R. Secunda)
Tips deeper pink, reddish, or rufescent
Tips brick-red, nearly scarlet: branches orange: sp.
16-17X8-9µ, verruculose: S. Amer. R. apiahyana
Tips rose-pink, branches cream then pinkish buff or tan,
finally brownish, massive: sp. 7-11·5X3·5-4·5µ, rough
or nearly smooth: N. Amer., Japan, S. Austral., Tas-
mama . R. botrytoides
Tips rufous or pinkish, branches pale alutaceous, then
goldenochraceous, massive: sp. 8-ro x3·5-4µ, smooth:
Europe R. rufescens
(Tips becoming reddish vinaceous: branches orange-
yellow: sp. 6·5-9·5X 3-4µ, nearly smooth: N. temp.,
S. Austral. R. fiavo-brun-
nescens var.
aurea)
Tips concolorous or paler, not differently or more intensely
coloured: N. Amer.
Reddish or bright pink at first, then fading
-3 cm. high, blood-red brown to light reddish salmon,
becoming lilaceous or violaceous: sp. 9-11 x 4µ,
rough or nearly smooth: Oregon . R. testaceo-
violacea
Larger, coral-pink, fading cream-ochraceous: sp.
cinnamon-ochraceous or pinkish buff
Sp. 7-9· 5X3-3·7 µ, rough or nearly smooth . . R. mbbotrytis
Sp. 8·5--11 X 3·5-4·5 µ, nearly smooth var. i11termedia
(Pink or.flesh colour: sp. 10-14X3-4·5µ, smooth R. obtusissima,
pink form)
RAMARIA 551
Not so brightly coloured

Vinaceous on bruising
Cream-white to pale tan, tinged pink, or flesh-colour,
tips concolorous, paler or pinkish: sp. 8-12·5x
4-5·5µ, nearly smooth R. secunda
Apricot-buff, then brown: sp. 10·5-12X4·5-5µ,
rough: Oregon R. brunnea
Not vinescent
Yellowish flesh-colour, branches divaricate: very
fragile: sp. 9-12·5X3·7-5µ, rough R. divaricata
Cream to light ochraceous�yellow, deeper upwards,
tips pinkish yellow, soon yellow, massive: sp.
9-12X3·5-4µ, rough R. conjuncta
Flava
Reddening on bruising
Sp. 10-18 X4-6·5µ, coarsely rough: sulphur or lemon-yellow:
Europe, Japan R. fiava
Sp. 6·5-12X3-5µ, finely rough or nearly smooth: pale cream
white, tips yellowish, then deeper yellow or pinkish, dull red-
brown with age: N. Amer., S. Austral. R. sanguinea
(Vinescent: orange-buff to cream or pinkish tan: U.S.A. R. aurea var.
australis)
(Dingy ochre, reddening or blackening: 2-spored: sp.
14-18X5-7 µ,: Germany R. macrospora)
Not reddening
Branch tips becoming viscid, capitate, and fused in tiers: yellow
to orange-yellow, tops honey-colour: sp. II-I 3X4·2-5µ,
rough: S. Austral. R. capitata
(Tips knob-like, not viscid, not fusing: yellow to pinkish
orange: sp. 8-10·5X4-5µ, rough: S. Austral. R. ochraceo
salmonicolor
Tips not dilated
Massive with very compact, short branches, rich yellow to
orange: sp. 8-15 X3-6µ, minutely rough or nearly smooth:
N. temp. R. aurea
Branches elongate
Stems rather slender, caespitose-fasciculate: light yellow,
mustard- or straw-yellow, or tinged orange: smell sweet,
strong with age: sp. 6-10X3-4·5µ, rough: S. Austral. . R. sinapicolor
Stem generally distinct, single, massive
Sp. 5·5-8x3·5µ, rough or nearly smooth: smell of
aniseed: buff-yellow to warm yellow: U.S.A. R. cystidiophora
Sp. larger: no smell of aniseed
Madagascar, ? trop. Asia
Sp. 11-14X4-7µ, rough: ochraceous R. madagas
cariensts
Temp.
Pale cream, rich yellow, or orange-ochre: sp.
7·5-II(-14) X3-4·5µ, rough: N. temp., S. Austral. R. fiavo
brunnescens
552 RAMARIA
More orange or pinkish yellow: sp. 7-9X3-4·5µ.,
rough or nearly smooth . var. aurea
(Lemon-yellow to ochraceous: sp. 9-16x4-6·5µ., rough:
Europe, N. Africa R. Mairei)
Violacea
Wholly lilac or violet, becoming fuliginous ochraceous or bistre
from the spores: sp. 8·5-12·5X3·7-5·5µ., rough: N. temp., S.
Austral. R.fumigata
Branches yellow, olive, or with yellow tips
Tips rusty-yellow, branches amethyst: sp.?: Europe R. rufoviolacea
Tips yellow, branches olivaceous: stem violet: sp. 8-12·5X
4-5·5 µ., rough: Europe R.fennica
Pale yellow or ochraceous-cream, the branches suffused lilac or
purplish, finally wholly ochraceous: sp. 9-16X4-6·5 µ.,rough:
Europe, N. Africa R. Mairei
(Blood-red brown or reddish pink, branches becoming lilac or
violet: sp. 9-11X4µ., rough or nearly smooth: Oregon . R. testaceo
violacea)
(Tan or cinnamon with lavender branchlets: see Decolorans)
Decolorans
Tan or brownish with lavender-pink branches or branchlets: U.S.A.
Sp. 13-16:5 X3 ·5µ., smooth: pale lavendf'r pink, becoming pale
tan or buff, vinescent on bruising: smell fragrant, medicinal . R. cacoa
Sp. 7·5-12X4-5·5µ, rough or nearly smooth: cinnamon-brown,
paler to the lavender-pink branchlets: not vinescent: smell
faint, musty . R. subspinulosa
Branches not lavender-pink
Spores generally more than 1 1µ. long
Vinescent or blackening
Dingy ochraceous: 2·-spored: sp. 14-18 X5-7µ, rough:
Germany R. macrospora
4-spored: sp. smooth, or nearly so: pallid white to light
yellow,· tips often brighter yellow
Sp. 10-2ox4-6 µ.: Finland R. decolorans
Sp. 12-15·5x2·5-5µ: N. Amer. R. xanthosperma
(Sp. 10-13x 3-4µ.: N. Amer. R. obtusissima)
Not vinescent
Pale alutaceous or tan, then brownish to cinnamon-umber:
sp. 12-18·5x4-7µ., smooth: Europe, N. Amer. R. Strasseri
Whitish to buff-yellow or ochraceous: sp. 9-15 X3-5µ.,
smooth: N. Amer. R. obtusissima
(Sp. rough: see R. Mairei, R. divaricata)
(Sp. striate: wholly white: see R. botrytis var. alba)
Sp. mostly less than 11µ. long
Tips greenish: -5 cm. high, alutaceous to pinkish cinnamon,
umber on bruising, compact: sp. 9-12X3-4·5µ, rough: in
coniferous woods: U .S.A. R. testaceo-
viridis
RAMARIA 553
Tips not greenish: flesh unchanging (or somewhat ochraceous
in R. verna)
Brownish or rufescent: sp. smooth
Tips clear yellow: branches alutaceous then tan- or
cinnamon-rufescent, reddish brown in age, crowded,
parallel: sp. 8-10(-12) X4-5µ: mostly in coniferous
woods: Europe R. condensata
Tips paler or concolorous, not yellow
Cinnamon or brownish umber, with spinulose tips: sp.
8-12 X4-6·5 µ: Europe R. spinulosa
Rufescent, branchlets alutaceous: sp. 6-10 X4-6µ: in
coniferous woods: Finland . R. Karstenii
Pallid: sp. more or less rough
N. Arner.: sp. 6·5-10 X3-4µ, rough or nearly smooth:
-10 cm. high, pale tan, pale buff, or pale flesh tan:
vernal . R. verna
Trop. or S. Amer.: sp. rugulose:
Sp. 6-7 X4-4·5 µ: -3 cm. high, dingy yellowish,
caespitose, branches contorted: S. Amer. R. subsigmoidea
Sp. 8-10·5 X4·5-5µ, aguttate: -5 cm. high (? more),
white tinged yellow: Malaya R. fiavo-alba
KEY TO THE SPECIES OF RAMARIA JNCERTAE SEDIS

Europe
Caespitose, pale ochraceous; sp. 7-10 X 6-7 µ, . olivaceous,
smooth: Germany ·. R. holsatica
White then reddish brown with fibrillose rhizomorphs, small:
sp. 2µ, round, smooth: in humus under Picea: Finland R. soluta
N. Amer.
11 cm. high, ochraceous, stem white: Ohio R. incurvata
Asia
7 cm., setulose, stem strigose: brown: sp. 10-12x4-4·5µ,
smooth: on the ground: Java . R. articulata
2 cm., yellowish: branches compressed, subfasciate: sp. 7-8 x 4µ,
smooth: on the ground: Java R. cornicu
larioides
Africa
Simple, white then flesh colour, 2·5 cm.: sp. ferruginous: on
Cyathea-trunks from S. Africa . R. Cyatheae
Branched
Egg-yellow, 6 cm.: sp. 9-r 1 X9µ, yellowish, smooth: S.
Thome . R. Henriquesii
Fuscous-ochraceous, 6 cm.: sp. 5-6µ (? 8-9), globose, smooth:
on branches or on the ground: Congo R. kisantuensis
Rufescent or ferruginous: in humus: S. Africa . R. Saccardoi
Trop. and S. Amer.
Sp. 15-19 X 7-9µ, smooth: 2·5 cm. high, terrestrial R. acutissima
554 RAMARIA
Sp. verrucose or echinulate
Sp. 3µ, subglobose: 2 cm., caespitose, slender; on rotting
fruits of Eucalyptus: Argentine . R. intricatissima
Sp. 8 X 6µ, echinulate: 8 cm., brownish: branches flattened:
on wood : Dominica R. cervicornis
Sp. ellipsoid, verruculose
Sp. 12 X 6 µ: branches slender: Guadeloupe R. guadelupensis
Sp. 8-10 X 4-5µ: 6 cm., brownish ochre: on the ground,
Guadeloupe R. leucoceras
Sp. 7-8 X 5 µ: 5 cm., greyish white: branches flat: on wood:
French Guiana R. albocinerea
Locality unknown
Slender, little branched, fawn-brown: on fern-trunks R. filicina
R. acutissima (B.) comb. nov.

Basinym: Clavaria acutissima B. in Mont., Gay FI. Chi!. 7, 1850, 386.
Synonyms: Lachnocladium acutissimum (B.) Sacc., Sy!!. 6, 1888, 740.
Clavaria carbonaria Mont., Guy. 1856, No. 425: Syll. Crypt. 1856, No. 597.
-2 · 5 cm. high, small, fuscescent (dried): stem tomcntose below, simple, soon very
much branched: primary branches subfasciculate, secondary dichotomous, fastigiate,
very acute and slender.
On the ground: S. Amer. (Guiana; Valdivia, Chile).
Spores 15-19 X 7-9 µ, yellow-brown, smooth, ellipsoid or subcylindric, obtuse,
2-3-guttate, apiculate.
Hyphae inflating somewhat, with clamps, thin-walled.
The type of this species in the Kew herbarium shows these remarkably large spores
so very abundantly in the hymenium that there can be no doubt of their origin. They
are the largest in the genus, together with those of R. apiahyana, but they are smooth
and blunt and suggest affinity with the Decolorans-group. Until the living fruit-bodies
have been described, however, it is impossible to place the species. The type-specimen
suggests a very stunted form of R. fiaccida.
The description of C. carbonaria seems identical, especially in the large characteristic
spores:
Blackening, fuscous-velutinate: stem short, dichotomous or fasciculate-ramose:
branches fastigiate, acute.
On burnt ground: Guyana (leg. Leprieur).
Spores 20µ long, yellowish ochraceous, amygdaloid.
R. albida (Pat.) comb. nov.

Basinym: Lachnocladiwn albidum Pat., Bull. Soc. Myc. Fr. r 898, 188.
5 cm. high, laxly caespitose, erect, coriaceous, ichitish: branches 7-8 mm. thick,
trifurcate, transversely sulcate-rugose, very shortly velutinate: branchlets short, rather
thick, cylindric, tips 3-4 dentate, obtuse: flesh fibrous, whitish: hymenium amphi
genous.
On trunks: Java.
Spores 13-16 X 8-roµ, ochraceous, ovoid, obtuse, apiculate at the base, set with
long spines.
The description refers, presumably, to dried specimens. The· colour and habit
suggest R. polypus, the spores are those of R. Zippelii and, if correctly ascribed, they
suggest an albino state of R. Zippelii.
RAMARIA SSS
R. albocinerea (Pat.) comb. nov.
Basinym: Lachnocladium albocinereum Pat., Joum. de Bot. 1889, 33.
4-5 cm. high, woody-rigid (? when dried), greyish white: stem 2 cm. X 5-7 mm.,
short, thickened at the base, villose and striate-canaliculate: branches short, broad,
flat, canaliculate, one to two times furcate, tips obtuse or truncate: hymenium uni
lateral, villous, whitish cinereous.
On rotten wood: French Guiana.
Spores 7-8x5 µ, brownish yellow, ovoid-apiculate, verruculose.
This may be another of Patouillard's descriptions of Scytinopogon, but the colour.
size, and shape of the spores indicate Ramaria aff. Stricta. Perhaps it is R. Moelleriana,
R. apiahyana (Speg.) c�mb. nov.

Basinym: Clavaria apiahyana Speg., Bo!. Ac. Nac. Cienc. Cordoba 23, 1919, extr.
p. 73: Sacc., Syll. 23, 1925, 487.
5-8 cm. high and wide, hemispherical: branches fleshy, ochraceous-white below,
orange above, tips intensely brick-red or nearly scarlet, subcylindric-canaliculate, smooth,
apices slightly thickened and shortly denticulate or subcristate: flesh white.
On the ground in woods: S. Amer. (Apiahy).
Spores 16-17 X 8-9 µ,ferruginous, densely and minutely verruculose, ellipsoid, apex
blunt, base unequally acute, often with one large gutta.
The description of this fungus is tantalizing. It suggests the most interesting species
of the genus and, yet, one cannot be certain. The colour suggests the Formosa-group:
the dark broad spores suggest the Echinospora-group: the warts on the spore also
suggest the Echinospora-group, but are they warts or spines? And does the spore
size include the warts or not? If related to C. Jormosa, it suggests a highly pigmented
ancestral form with echinulate or verrucose spores. If related to C. Jragillima or C.
Zippelii it suggests the connexion between the sombre, heavily pigmented Echinospora
group and the bright and lightly pigmented Formosa-group, cf. the orange-rufous
tips of R. nigrescens and R. Jragillima.
R. apiculata (Fr.) Donk

Rev. Nieder!. Homob. Aphyll. 2, 1933, 105.
Basinym: Clavaria apiculata Fr., Syst. Myc. 1, 1821, 470: Coker, Clav. U.S. Can ..
1923, 157, t. 39, 88: Lundell, Sv. Svamp. 3-4, 1936, 20.
Synonyms: Clavaria tsugina Pk., Bull. N.Y. St. Mus. 67, 1903, 27 (fide Coker).
Clavariella apiculata (Fr.) Karst., Hattsv. 2, 1882, 188.
? Clavaria Stillingeri Coker, Journ. El. Mitch. Sci. Soc. 42, 1927, 251, t. 47.
-7 cm. high, small or medium size, solitary or caespitose, light pinkish ochraceous or
creamy yellow, becoming deeper ochraceous brown, vinaceous-cinnamon or rufescent
alutaceous from the base upwards, tips whitish then paler concolorous, sometimes greenish:
stem 3-4 mm. thick, branched from or near the base, arising from an abundant
fibrillose mycelium, or a white tomentose myce!ial felt (when growing on wood):
branches rather elongate and flattened, numerous, rather lax, ending in 2-3 long acute
tips, with scurfy-tomentose sterile hymenium on the uppersides: flesh dense, tough,
slightly bitter, drying dark: smell practically none.
On dead wood, bark, and humus of coniferous trees: U.S.A. (common in the north
and south), Japan, Siberia, Europe.
Spores 6·5-1ox3·5-5µ,, dull ochraceous, minutely rough or minutely verruculose
to nearly smooth.
Basidia 7·5 fL wide, 4-spored: 36-45 X 7·5µ,, sterigmata 6-9µ, (Doty).
Hymenium -75µ, thick, thickening.
Hyphae 3-7µ, wide, the walls 1-2·5 µ, thick, often with occluded lumen, interwoven
and often 'knotty' in the central tissue.
556 RAMARIA
var. compacta Bourd. et Galz.

Hym. Fr. 1928, 98 (ut Clavaria dendroidea var. compacta).
3-5 cm. high, pale cream-ochraceous, tips whitish: branches very crowded, thick,
branchlets short, tuberculiform or subulate: flesh fibrous.
On a stump of Pinus: France (rare).
Spores 6--7(-9) X 4-4·5 µ., pale ochraceous, finely rough, oblong.
Basidia 45-80 X6-7·5µ.: hyphae 3-9µ. wide, clamped.
This is C. apiculata in the sense of Coker, with whom Lundell and Donk agree.
It is also C. apiculata sensu Karst., who gave the spores as 8-9 X4-5µ.. It is close to
C. stricta and differs in the pale tips, generally small size, laxer branching, tougher
and denser flesh, paler spores, habitat always on coniferous wood or humus (particu
larly Picea excelsa, according to Lundell) and, apparently, in the unchanging flesh,
though no author seems to have remarked on this absence of discoloration, nor to have
recorded its presence: the spores may also be smaller. Possibly, therefore, R. stricta var.
alba should belong to R. apiculata. Lundell notes that the tips of the branches are
tinged green when young; Coker was unable to detect the tint in American specimens,
but it is mentioned by Doty: it suggests, nevertheless, affinity with the little-known
R. testaceo-viridis.
Coker recognizes also the following three species: C. acris, C. pinicola, and C.
Stillingeri, which appear to me to be fully covered by his description of C. apiculata,
which, as a common fungus, is certainly variable:
Clavaria acris Pk., Rept. N.Y. St. Mus. 54, 1901, 155, t. H, f. 37-9: Coker, I.e., 162,
t. 89: Wehm., Pap. Mich. Ac. Sci. Arts' Lett. 20, 1935, 264: Doty, Clav. Pac. N.W.
1944, 39.
3·5-7·5 cm. high, nearly as wide, caespitose, reddish flesh-colour, tips whitish or
concolorous (Coker), buff pink to vinaceous cinnamon, tips paler (Wehm.): branching
from the base: branches repeatedly and subpalmately branched, sometimes com
pressed (Coker), fasciculate rather than dichotomous (Wehm.): flesh white, tough:
mycelium white: taste acrid (Coker), slightly and tardily acrid (Wehm.).
On rotten coniferous wood: U.S.A. (N.Y.), Canada (Nov. Scot., common; Wehm.).
Spores 6--7·5 X 4-5µ., pale ochraceous, rough (Coker): 6-7 X3·5-4·5µ., darker brown
than C. stricta (Wehm.): 5-7·5(-8) X3· 5-4·5µ., mostly 5·5-6·3 X3·5-4µ. in the type
(Doty).
This is said to differ from R. apiculata in its smaller spores and acrid taste, but there
is so much latitude in the spores of R. apiculata (see Table XXXIII) that it is difficult
to see what value the distinction can have. Wehmeyer's collection suggests also that
the acrid taste is not always pronounced. The spore-colours· given by Coker and
Wehmeyer are so different that one may doubt the identity of their collections:
Wehmeyer's plants suggest C. stricta var. concolor, and a peppery taste has been
recorded for C. stricta. Doty describes it as Capucine Buff with Capucine Orange tips,
and with acrid, not bitter, taste.
C. pinicola Burt, Ann. Mo. Bot. Gdn. 9, 1922, 25, t. 5, f. 32: Doty, Clav. Pac. N.W.
1944, 44.
Caespitose from a common white mycelium, slender, -3 cm. high (Doty), of rather
uniform diameter, sometimes simple, usually one to three times dichotomous: branches
cylindric, spreading, buff-pink to drab (Doty), drying huffy brown all over, tips acute:
slightly bitter (Doty). � .
On coniferous logs and bark: U.S.A. (Idaho, Oregon, California).
Spores 7·5-11 X4·2-5·5µ., pale ochraceous, smooth: 7·5-9·5 x4·5-5·5µ. (Doty).
Basidia 7-8µ. wide, 4-spored.
Hymenium thickening: hyp hae 3·5-6·5µ. wide, thick-walled.
Coker says this is merely a smooth and slightly larger-spored variety of R. apiculata,
hardly worthy of varietal rank. From Table XXXIII it can be seen that there is no
RAMARIA 557
difference from R. apiculata in spore-size, and Cok�r says that smooth spores occur
in most collections of R. apiculata. Doty maintains it as a species because the spores
are always smooth and broader than in R. apiculata.
C. Stillingeri: -2 cm. high, small, sparingly branched, solitary or gregarious, arising
from a dense white subiculum (without rhizomorphs), sparingly and openly branched,
dull brown (when dry) with whitish tips, smooth, rigid: sp. 8·5-12·5 x4·4-5·5µ,
ochraceous, smooth, ellipsoid: basidia 25-34X 7·5-9·5µ, with 4 sterigmata 7·5µ long:
on wood of Abies grandis, Elk River, Idaho: (leg. C. R. Stillinger, Lloyd's herb.).
This is said to be near Lentaria byssiseda and L. epichnoa, but it is exactly young
R. apicu/ata except for the slightly longer spores.
TABLE XXXIII. SPORE-SIZE IN RAMARIA APICULATA
Spore-size in µ. \. Authorit'.!:__ Country Remarks

7·5-9·8X3·7-4·2 Coke< U.S.A. Baker (Alabama)
6-9·7X3·8-4·5 I Sweden Friesian specimen (Kewj
8·5-9·5X4 C. tsugina (type)
7·5-9·3X4-5 " " Romell
8-9X4-5 I Karsten Finland
6·5-9·5X3·7-4·2 Coker U.S.A. N. Car. 2962
8·5-10(-12)X4-5(-6·4) I Doty
7·5-9·3X3·8-4·5 Coker N. Car. 2939
7·5-9·3X3·7-4 " " Burnham 114
8-9X4-5 Karsten Finland
7·3-8·3X4-4·6 Coker U.S.A. Price (Kentucky)
7-8·5X3·5 Washington
7·5-8X4·8-5"3_ Ellis (N.Y.)
7·5X4 Atkinson (N.Y.)
6·5-7·5X3·7-4·3 " " Holmes (N. Car.)
6-7(-9)X4-4·5 Bourdot France var. compacta
7·8-1 l X +·2-5·5 Coker U.S.A. C. pinicola: Burnham

7·5-10X4·2-5·5 " C. pinicola (type)
7·5-9X4·5-5 Burt
7·5-9·5X4·5-5·5 Doty
6-7·5X4-5 Coker " C. acris

6-7X3·5-4·5 Wehmeyer Canada
5-7·5(-8)X3'5-4·5 Doty U.S.A.
5·5-6·3X3·5-4 (type)
R. articulata (P. Henn.) comb. nov.

Basinym: Lachnocladium articulat11m P. Henn., Monsunia 1, 1900, 142.
5-7 cm. high, coriaceous, stalked, setulose, upright, pale brown (? dried): stem
1-1 ·5 cm. x 2-3 mm., terete, strigose: branches upright, crowded, repeatedly dichoto
mous or fasciculate, axils little compressed, subarticulatc, tips subulate, branchlets
atte11uate to each end.
On the ground: Java (Tjibodas, 1,500 m. alt.).
Spores 10-12 x 4-4·5µ, brown, subfusoid or ellipsoid, smooth, 1-guttulate.
If correctly described, the spores indicate Ramaria.
R. aurea (Fr.) Quel.

FI. Myc. 1888, 467.
Basinym: Clavaria aurea Fr., Epicr. 1838, 574: Coker, Clav. U.S. Can. 1923, 142,
t. 86: Bres., le. Myc., t. 1085: Gramberg, Pilze der Heimat 3, 1921, t. 31 (ut C. fiava):
Pilat, Bull. Soc. Myc. Fr. 49, 1933, 49.
558 RAMARIA
Synonyms: Clavariella aurea (Fr.) Karst., Hattsv. 2, 1882, 184.
Clavariafiava sens. Fr. 1838, 1874: sensu Lundell, in Krok et Almq. 5, ed. 2, 1932,
285: (fide Donk).
5-12 X 10-20 cm., massive, entirely ochraceous yellow, golden ochraceous or egg-yellow,
even somewhat orange or tawny, becoming wholly deep ochraceous with age, richly
coloured: stem 2-4 X 1-5 cm., massive, short, often bulbous, whitish at the base:
branches rather short, dense, thick, much branched, often cauliflower-like, the tips pale
when growing: flesh white, yellowish near the surface, unchanging, rather brittle, watery
marbled: edible.
On the ground in frondose or coniferous woods: Europe, U.S.A., Japan, Asia
Minor (Ilgaz-Dagh Mts., in woods of Abies Bornmulleriana, Pilat).'
Spores 8-15 X 3-6 µ, deep ochraceous, finely verruculose, rough or minutely rough,
sometimes almost smooth, oblong, one to several guttulate.
Basidia 45-75 X 6-8 µ. (Bourd. et Galz.), 35-45 X 6-9 µ. (Bres.), 2-4 spored.
Hymenium not thickening: subhymenium as in R. formosa.
Hyphae 3-15 µ. wide, thin-walled, cla.:nped: structure as in R. formosa.
(var. australis Coker, see below.)
This is the typical state of R. aurea as understood in Europe and, so it seems, in the
United States and Japan. I have found it in Switzerland and Austria, and I agree with
Coker that the spores are finely rough or nearly smooth, and thus they are distin
guishable from those of R. fiava with which R. aurea has been so much confused.
I find also that the spores of R. fiava are appreciably larger. Probably R. aurea has
been confused in Europe with the young, clear yellow, states of R. Mairei and with
the old ochraceous-yellow states of i?.. formo.sa, R.fiava, and R. Mairei, and hence the
verrucose spores ascribed to it by several authors. If I have correctly understood
R.fiava, it is readily distinguishable from R. aurea, though' Donk suggests they should
be united.
R. fiavo-brunnescens differs in the elongate branches, so that it has the form of
R. fiava and R.jormosa, and in the smaller spores which are often smooth. Neverthe
less it seems to merge into R. aurea, particularly in the United States, cf. R. fiavo
brunnescens, var. aurea.
C. aurea var. australis Coker, from the United States, seems to me a mixture of
abnormal specimens of R. sanguinea and R.fiavo-brunnescens or R. formosa. In all his
specimens the branches were elongate and the fruit-bodies comparatively slender, not
as in R. aurea. The description is:
C. aurea var. australis Coker, I.e., 144, t. 53-5, 87.
-11 cm. high and broad, branched from a pointed base, or caespitose, wholly rich
buffy orange, the base whitish and readily turning vinaceous-brown on handling: branches
rather elongate: flesh white, delicate, rather brittle: taste mildly bitter: smell musty,
faintly medicinal.
Spores u-13 X 4-4·5 µ, buffy ' ochraceous, minutely rough (verruculose in Coker,
I.e., t. 86, f. 2, 3):
On the ground in mixed woods: U.S.A. [N. Car., No. 2912 (type), No. 2913: N.Y.].
Form 1: creamy flesh, then creamy tan with only the tips orange and turning dull
brick-brown when bruised, base dull wine-brown when bruised: spores II-I 3 X
4-4·5 µ, buffy ochraceous, minutely rough: N. Car., No. 2882a.
Form 2: brownish tan tinged dull pink, the base deeper brown on br:uising: spores
7·5-II X3·7-5µ, ochraceous, minutely rough or smooth: N. Car., No. 2597.
R. Bataillei (Maire) comb. nov.
Basinym: Clavariella Bataillei Maire, Ann. Myc. II, 1913, 351, t. 18.
Synonym: Clavaria Bataillei (Maire) Sacc., Syll. 23, 1925, 488.
1
The records from Australia and S. America need confirmation. They may be R. ftavo
brunnescens var. aurea or R. ochraceo-salmonicolor.
RAMARIA 559
TABLE XXXIV. SPORE-SIZE IN RAMARIA AUREA
I
10-15X4-6
___ , ________
Spore-size in µ. Authority
Coker
,________
Italy
Country Remarks
rough (2 spec. in Herb.Bresad.)
u-14X4·5-5 ,, Sweden rough (Herb. Romell)
11-15·5X4-5·5 Corner Switzerland finely rough: Table XXXV
(Grisebach)
11-15(-19·5)X4·3-5·5 Austria finely rough
(Rekawinkl)
8-13X4-4·5 Bourd, et Galz. France puncticulate
9-12X4-5 Bresadola Italy verruculose
9-12X3-4 Rea Gt. Britain
8-11X4 Karsten Finland
9·3-11X4·5-6·5 Coker Sweden Friesian specimen in Herb
Curtis
9-13X4-5·5 Atkinson U.S.A. (Ithaca)
9·5-11X4·2-4·6 Coker ,, ( Albany) pale brownish yellow fruit
bodies'
11-13 X4-5 " (N.Y.)
9-12(-15)X4-4·5 Kauffman (Rockie (? = R. Jormosa) 2
Mt.)
II-13X4-4·5 Coker (N. Car.) minutely rough: var. australis
II-13X4-4·5 rough, mostly 12X4 µ.: var.
australis.
11x4·5 (N.Y.) var. australis
7·5-11X3·7-5 (N. Car.) mostly 9·5 µ. long, minutely
rough or smooth (no. 2597):
var. australis.
1
? old specimens of R. aurea or of R. fiavo-brunnescens.
2
.Size and habit of R. fiava: colour of branches apricot-yellow to buff yellow (Ridgway),
with lemon-yellow tips . ..under pine, spruce, fir (Pap. Mich. Ac.Sci. Arts Lett. 1, 1923, 123).
TABLE XXXV. SPORE-SIZE IN RAMARIA AUREA

(One fruit-body: Grisebach, Switzerland, Aug. I925)
Spore-size in µ. E' Number of records \I Spore-size in µ. E' Number of records

I
11·ox4·5 2·4 2 13·0X5·3 2·5 2

II'OX5·0 2·2 I 13·5X4·7 2·9 2
11·5X4·5 2·6 2 13·5X5·0 2·7 3
11·5X4·7 2·5 2 14·0X4•7 3·0 I
11·5 x5·0 2·5 3 14·ox5·0 2·8 4
12·0X4·0 3·0 I 14·0X5·3 2·6 2
12·0X4·5 2·7 3 14·5x5·0 2·9 2
12·0X4'7 2·5 I 14·5x5·5 2·6 I
12·0x 5·0 2·4 2 15·0X5·0 3·0 5
12·5X5·0 2·5 3 15·0X5·3 2·8 3
13·0X4·7 2·8 I 15·0X5·5 2·7 I
13·0X5·0 2·6 2 15·5x5·0 3·1 I
1
E = length -;- width.
Range: 11-15·5X4-5·5 µ., E = 2·2-3·1, for 50 spores.
Average of 10 spores 11-11·5 µ. long, E = 2·4
12-12·5 µ. E = 2·6
13-13·5 µ. E = 2·7
14-14·5 µ. E = 2·8
,, ,, 15-15·5 µ. E = 2·9
Average of 50 spores 13·3x4·9 µ., E = 2·7
566 RAMARIA
7-11 cm. high, rose-orange, the branches becoming greyish v,"olet or juscous-brozvnish
and spotted brown on touching, finally fulvescent from the spores: stem 2-5 cm. thick,
short and thick, dividing into many stout branches, persistently rose-orange, rather
pale, base white: branches erect, fastigiate, polychotomous then repeatedly dichoto
mous, rather elongate, subcylindric, sometimes slightly longitudinally rugulose, tips
subacute and golden ochraceous then fazvn ochraceous or subconcolorous: flesh zvhite,
quickly turning brownish violet on bruising or cutting: taste bitter, smell feeble.
On humus in coniferous forests: France trare).
Spores 12-15 X 4-5µ, ochraceous or pale cream ochre, ellipsoid-oblong, finely
verrucose-rough.
Basidia 70-85 x 7-8 µ, 2-4-sporeo.
Maire says this comes in the group of violaceous species (R. fennica, R. versatilis,
R. rufo-violacea, &c.) from which it differs in the pinkish orange stem and the dis
coloration of the flesh, and that in this colour-change it resembles R. testaceo-fiava.
To me, R. Bataillei appears like an old sordid specimen of R. Jormosa, from which it
differs only in the brownish or fuscous-fawn colour which the branches acquire with
age, and perhaps through incipient drying or through bruising by rain: thus R.
brunnea may be identical. R. testaceo-fiava also seems such a discoloured state of
R. formosa. In spore-characters, there seems no difference between the four species.
R. botrytis (Fr.) Ricken

Vademecum 1918, 253.
Basinym: Clavaria botrytis Fr., Syst. Myc. 1, 1821, 466: Bourd. et Galz., Bull. Soc.
Myc. Fr. 26, 1910, 213.: Maire, Bull. Soc. Myc. Fr. 27, 1911, 449 [ut Clavariella
botrytis (Fr.) Maire]: Rolland, Champ., t. 103: Gramberg, Pilze der Heimat 3rd ed. 2,
1921, t. 32: Bres., le. Myc., t. 1083: [haud sensu Coker = R. botrytoides (Pk.)].
Synonyms: Clavaria platyceras Viv., Fung. ltal. 1834, t. 54.
Clavaria purpurascens Paulet, le. Champ. 1855, 113, t. 194, f. 6.
Ramaria acroporphyr,ea Quel., FI. Myc. 1888, 466.
Clavaria sculpta Beck, Verh. k. Zool.-Bot. Ges. Wien, 39, 1889, 603 (teste Maire,
Cott. et Wakef.).
Clavaria Rielii Boud., Bull. Soc. Myc. Fr. 13, 1897, 14 (teste Maire, Cott. et
Wakef.}.
Clavariella botrytis (Fr.) Schroet., Krypt. FI. Schles. 3, 1888, 447.
Text-Fig. -248.
7-1s-x6-20 cm., massive, white then alutaceous, tan, or ochraceous with deep pink,
red, purple, or vinous tips: stem 3-4x1 ·5-6 cm., massive, white then yellowish, passing
into very numerous, crowded, thick, irregular branches: branches polychotomous,
stout below, very numerous, short and congested or somewhat elongate, much
branched, many abortive, often with crowded cauliflower-like tips: flesh white,
unchanging, moderately brittle, watery-marbled: smell pleasant, fruity: edible.
On the ground in frondose woods: Europe, U.S.A. (Oregon).
Spores r2-20X4-6 i-t, mostly 14-16x4·5-5·5µ, ochraceous, smooth but faintly
longitudinally striate (the striations often subreticulate, Cott. et Wakef.), r-3 guttulate,
oblong-elliptic.
Basidia 60-70 X 8-roµ: sterigmata 2-4.
Hyphae 3-12µ wide, clamped, the cells 50-180µ long.
forma parvula Bourd. et Galz.

Hym. Fr. 1928.
-2 cm. high, wholly rose-i,urple except the white base. France.
RAMARIA
var. alba Pearson
Tr. Brit. Myc. Soc. 29, 1946, 209.
Wholly white: spores pale ochraceous: England (Henley, Sussex).
This is one of the more striking and easily recognized of European fungi, yet there
has been much confusion over its identity. Fries placed it in the white-spared group
of branched Clavarias, where it remained until Schroeter transferred it to Clavariella.
From then onward there seems to have been no doubt concerning it among European
mycologists, except in so far as R. rufescens may or may not be synonymous. The
description which I give is that of the European species. But if the problem of its
original identity is simple, in other parts of the world there are varieties or allied species
which make its definition extremely difficult with our present insufficient knowledge.
It has been recorded from N. America, Japan, and Australia, yet, except for Doty's
record of true R. botrytis from Oregon, none of the others agree exactly with the
European species.
TABLE XXXVI. SPORES OF RAMARIA BOTRYTIS AND R. HOLORUBELLA
Spore-size in µ. Authority Country Remarks

C. botrytis
14-20X4·5-6 Corner Austria see Table XXXVII
12-18X4-6 Bresadola Italy
12-18X4-6 Donk Holland
12-16X4'5-6 Bourd. et Galz. France
12-16X4-5 Cott. et Wakef. England
12·5-16X4·5-5·5 Corner "
12�15X4-6 Rea "
II-16X4-5•5 Coker ? Italy Herb. Bresadola
(12-)14-16(-21) X(4·5-)5-6 Doty U.S.A. (Oregon)
C. holorubella
II-13X3-4·5 Atkinson U.S.A. (Ohio) Type: striate
10·5-13X4-5 Coker " " " "
12-13X4-4·5 Burt " " " "
11-13·5X3·7-4•5 Coker " (N. Car.) No. 2862 1
II-13X3-4-·3 " " " No. 563 : 1 mostly

n·5X3·5 µ
II-13X3·3-3·8 " " " No. 2845 :' striate
II-12·2X3·7-4 " " " 'No. 2884 1
10·2-12X3-3·7 " " " No. 531 :' smooth
9·3-12X4-4·8 " " " No. 57221
10-13(-15)X4-5 Kauffmann " (Colorado) as C. botrytis
10-13(-15)X5-6 " " (Mich.) "
12-15·5X3-4·5 Fawcett Australia as C. rufescens
1
As C. rufescens sensu Coker.
In N. America there appear to be two species macroscopically si.milar but with the
tips of the branches of slightly different colour and with shorter spores. One has
striate markings on the spores, like R. botrytis, and this was identified by Coker with
R. rufescens of Europe and by Kauffman with R. botrytis of Europe. I have called it
by a synonym, R. holorubella, because it does not seem the same as R. rufescens sensu
Bres. (p. 619) and because it has spores consistently shorter than the European R.
botrytis (see Table XXXVI). The other species has verruculose-rough or nearly
smooth spores and paler pink tips to the branches: it was called C. botrytis by Coker,
and Bresadola considered it the same as C. rufescens as understood by him. I have
called it by a synonym, R. botrytoides, because it does not agree exactly with either
5Il9 00
562 RAMARIA
European species. R. botrytoides belongs to the Formosa-group in the subbotrytis
series.
The Australian species identified as C. botrytis by Cleland and by Fawcett are
certainly R. botrytoides. So, too, appears to be the Japanese record, though true R.
botrytis and R. holorubella may occur in Japan.
TABLE XXXVII. SPORE-SIZE IN RAMARIA BOTRYTIS
(One fruit-body; Austria, Aug. I925)
Spore-size in µ.
14·0X4·7
_I_£_
3·0
Number of records -
--
l
Spore-size in µ.
16·0X5·3
E'
3·0
..
Number of records
4
14·0X5·0 2·8 l 16·0X6·o 2·7 I
14·5X4·5 3·2 l 16·5X6·o 2·8 I
14·5x4·7 3·1 l 17·0X4·5 3·8 2
14·5X5·0 2·9 4 17·0 X4·7 3·6 2
14·5X5·3 2·7 I I 17·0 X 5·0 3·4 2
I I
14·5x5·5 2·6 l 17·0X5·3 3·2 l
15·ox4·5 3·3 l 17·0x5·5 3·1 1
15·ox4·7 3·2 l 17·5 X 4·5 3·9 I
15·0X5·0 3·0 2 17·5 x6·o 2·9 I
15·0X5·3 2·8 2 18·0X5·0 3·6 I
15·0X5·5 2·7 I 18·0 X5·3 3·4 l
15·5X5·0 3·1 3 2o·ox5·0 4·0
I
l
16·ox4·5 3·6 l 20·0X5·5 3·6 l
16·0 x5·0 3·2 3
Range: i.1--20X4·5-6µ., E 2·6-4·0, for 44 spores.

Average of 10 spores 14-14·5 µ. long, E = 2·9
15-15·5 µ. E = 3·0
16-16·5 µ. E = 3·1
,, ,, 17-17·5 µ. E = 3·5
Average of 4 spores 18-20µ. ,, E = 3·7
Average, overall, 16·0X 5·1 µ., E = 3·1
R. botrytoides (Pk.) comb. nov.

Basinym: Clavaria botrytoides Pk., Bull. N.Y. St. Mus. 94, 1905, 21, 49, t. 93, f. 5-7:
Coker, Clav. U.S., Can. 1923, 1 II, t. 31, 32, 85 (ut C. botrytis): Burt, N. Am. Clav.
1922, 8: Fawcett, Proc. Roy. Soc. Viet. 51, 1939, 275 (ut C. botrytis).
7-16 x3-15 cm., massive, branching at the ground from a short, white, rooting base
set with numerous abortive pink-tipped branches, pallid cream, pinkish towards the
rose-pink tips, becoming tan or brownish tan with dull brick-brown tips with age: branches
curving upright, rather elongate, rugose, much branched distally: flesh firm, turgid,
brittle, concolorous: taste like green pea-hulls, smell similar.
On the ground in woods: U.S.A. (common throughout): Austral., Tasmania:
? Japan.
Spores 7-11·5x3·5-4·5/L, light buff yellow (Coker), rusty brown or cinnamon
(Peck), rough or nearly smooth, not striate, ellipsoid.
Basidia 8 ft wide, 4-sporcd: hymenium 30 fL thick.
Coker considered this species to be C. botrytis of Europe, but European mycologists
have always regarded C. botrytis as the more massive fungus with massive trunk,
darker purple tips, and much longer, smooth, but finely striate, spores. This short
spored American fungus, with the habit of R. formosa or R. Mairei, has not been
found in Europe. Hence I have revived Peck's name, on the authority of Coker's
identification. Kauffman also doubted Coker's interpretation of C. botrytis.
RAMARIA
The Australian fungi identified as C. botrytis by Fawcett agree almost exactly with
the American specimens, though their colour may be more inte.nsely pink when young
(tending towards R. subbotrytis). The descriptions of the Australian specimens
briefly is:
3-16 X 3-12 cm., mostly c. 8 cm. high, massive, with rather elongate branches:
wholly bright pink when young, becoming paler and subochraceous or salmon-buff (as the
spores ripen), with the tips persistently bright pink (rarely wholly white): flesh white,
crisp, brittle, soft, and pale buff with. age.
Spores 7·5-11·5(-12·5) X 3·8-4·5(-5·5)µ,, light yellow buff, minutely rough to nearly
smooth.
Common in all southern parts of Australia and Tasmania: edible.
On the other hand, the Australian specimens identified by Cleland and Cheel as
C. botrytis have distinctly wider spores (Table XXXVIII) and seem to lack the pink
colour, being wholly pale. yellowish, reddish fawn, yellowish fawn-buff, or pale fawn
brown. An unusual yellowish brown specimen with rod-like spores, 6-10 X 2-2·5 µ,,
was found on Mt. Lofty, N.S.W. Perhaps they should be referred to R. brunnea or
R. formosa, unless they indicate yet another species.
The Japanese specimens, referred to C. botrytis, must also belong here: the spores
were said to be almost smooth or faintly verruculose, not striate (Imai, 1931).
Bresadola considers C. botrytoides ( = C. botrytis sensu Coker) to be the species he
has identified as C. rufescens. Bresadola's illustration (le. Myc., t. 1084) shows a light
golden ochraceous fungus with rufescent (pale reddish-brown) tips and a v�ry massive
stem 8 X 3-3 ·5 cm., which does not agree with Coker's description of C. botrytoides,
though the spores are similar (8-10 X 3 ·5-4µ,, yellow, smooth). There is, unfortunately,
such confusion over C. rufescens, as I have mentioned under R. botrytis, that I prefer
to keep the European and American species apart, at least until they are better known.
R. subbotrytis is extremely close and seems distinguishable only by the more intense,
less evanescent, pink colour. R. secunda is also close but not, apparently, acrochroic.
TABLE XXXVIII. SPORE-DATA FOR RAMARIA BOTRYTOIDES
Spore-size in µ. Authority
I
I
Country Remarks
8·5-n·5x4·5 Coker U.S.A. Type (N.Y.): rough
7·5-10·3X3·3-4
7·5-10X3·8-4·2
»
»
I »
»
N. Car. 2628
" B 72: rough to nearly smooth
7'5-9·5 X3·7-4·5 " " " 344: minutely rough
6·6-9·5X3·5-4 " " " 2899: nearly smooth
7·5-9·3X3·8-4·5 " I » » 661: nearly smooth
7·7-9·3X3·8-4 » » » 2596: minutely rough
7·3-8·2 X3·3-7 » " » 2863
7·5-10X3·8-4·2 rawcett Australia minutely rough to almost smooth

11·5-12·5x5·5 Cleland, Chee! »
11X5•5 " "
10·5 X4·5 " "
10X4·5 » »
8·5-10·5 X4·5 " »
9x5·5 » »
R. Bourdotiana Maire
Fungi Catal. 2, 1937, 32.
Pseudonym: Clavaria stricta sem ..1 Bourd. et Galz., Hym. Fr. 1928, 98: sensu Bres.,
le. Myc., t. 1093 (excl. sporis) (fide Maire).
3-9 cm. high, pale yellowish, pale alutaceous or cream, then ochraceous, tinged
brownish or rufescent in age, not discoloring on bruising, tips white: stem 2-5 mm.
RAMARIA
wide, slender, cylindric, attached to white rhizomorphs: branches numerous, much
branched, erect, crowded, dichotomous, strict, acute: flesh firm: taste almost none.
vn dead wood, stumps, branches and roots of frondose trees (e.g. Ulmus, Acer):
France, Spain, Italy, N. Africa.
Spores 4-6 X 3-3·5µ. (5-6 X 2·5-3µ. (Maire) ), ochraceous, minutely rough, oblong.
Basidia 15-30 X 3 · 5-6µ.: sterigmata 2-4.
Hyphae 3-14µ. wide.
This species differs from R. stricta in the smaller spores, paler colour, white tips, and
unchanging flesh. Bourdot and Galzin described it as C. stricta which, in tum, they
called C. dendroidea and C. condensata. Maire recognized the fungus in N. Africa,
where it is frequent, and named it in honour of Bourdot. It may be a tropical species,
and R. polypus, which I have described from Malaya, is certainly close but 'with larger
spores and lax branching. The tropical R. luteo-fiaccida has similar, but distinctly
verruculose, spores and colouring, and it is humicolous with lax branching. R.
myceliosa, of N. America, has spores as small, but -verruculose or subechinulate, and
grows in coniferous humus.
R. brunnea (Zeller) comb. nov.
Basinym: Clavaria brunnea Zeller, Mycol. 27, 1935, 453, f. 2: Doty, Clav. Pac. N.W.
1944, 50, t. 7.
-15 cm. high and II cm. wide, massive, apricot-buff then fawn or darker brown,
drying pinkish cinnamon or cinnamon-buff to brown, somewhat vinaceous on bruising:
stem massive: branches elongate, erect, fastigiate, the larger ones hollow: flesh white,
bitter.
Under conifers: U.S.A. (Oregon).
Spores 10·5-12(-13) X 4·5-5 µ, ochraceous, verrucose, elongate ellipsoid.
Basidia 40-50 X 7·5-10·5 µ, densely granular, 4-spored.
'C. brunnea, one of Oregon's most beautiful species, is closely related to C.jormosa,
from which it may be separated by the narrower spores, darker colour and bitter
taste' (Doty). However, there is no distinction in the spore-size, but the absence of
yellow tips may be characteristic. Compare R. Bataillei and R. subspinulosa.
R. cacao (Coker) comb. nov.

Basinym: Clavaria cacao Coker, Clav. U.S. Can. 1923, 151, t. 58, 87: Mains, Pap.
Mich. Ac. Sci. Arts Lett. 20, 1935, 88-9.
7-8 X 5-6 cm., pale lavender-pink, at least the upper branches, becoming pale tan or
bufffrom below upwards, the tips pale lavender-pink for a long time, then shrivelling
brown: stem 1·5-2·5 cm. thick, short, firm, whitish, vinous on bruising: branches rather
open and -crooked, numerous, rather elongate, with crowded acute tips: flesh firm,
rigid, elastic, not brittle, pure white or slightly concolorous, not changing colour except at
the base: taste mild: smell noticeable on ageing or drying, medicinal-fragrant, like cacao
butter, stronger in dried plants and persistent for years.
On the ground in woods: U.S.A. (N. Car., Mich., rare).
Spores 13-16·5 X 3·5 µ, ochraceous, smooth, elongate ellipsoid, sway-backed.
Basidia 6-9·3 µ. wide, 4-spored.
Hymenium 65-95 µ. thick, with medium-sized crystals.
Hyphae 3-8 µ. wide, clamped.
This seems related to R. Mairei, but is distinguished by the smell, absence of yellow
colour, vinaceous discoloration of the stem, the firm flesh and smooth, sway-backed,
narrow spores. The last two features suggest Lentaria, but the general appearance of
the fruit-body like R. formosa, as shown by Coker's photograph, the vinaceous dis
coloration, the distinctly ochraceous spores, and the humicolous habitat without a
mycelial felt indicate Ramaria. It has the narrowest spores in the genus. Compare
R. subspinulosa with shorter, subverruculose spores and R. testaceoviolacea with
RAMARIA 565
reddish fruit-bodies and shorter, broader spores, but also with a strong sweet smell
when dried.
R. capitata (Lloyd) comb. nov.
Basinym: Clavaria capitata Lloyd, Myc. Notes 7, 1922, 1107: Fawcett, Proc. Roy.
Soc. Viet. 51, 1939, 276, t. 17, 20.
Massive, much branched (-10 cm. high, ? more), caespitose from a mass of mycelium,
maize-yellow to pale orange-yellow, the viscid capitate tops of the branches pale honey
yellow and semi-translucent: stems divided from the base, white: branches apparently
polychotornous, 8-15 mm. wide below, stout, curved-ascending with acute axils, with
or without a second set' of branches: tips becoming subglobose, then subcapitate, viscid,
semitranslucent, often coalescent laterally or somewhat in tiers: flesh white, crisp, brittle:
taste mild.
On the ground: Austral. (Victoria; Kinglake, Tourrorrong, Healesville, Cockatoo).
Spores 11-13 X 4·2-5 µ,, ochraceous, rough, ellipsoid, hyaline under the microscope.
Basidia with 2, less often 4, sterigmata: the hymeniurn on the dry sides of the
branches and on the viscid tops: clamps not seen.
This description is drawn from Fawcett'1: emendation of Lloyd's. It is a remarkable
fungus, to be imagined as a much-branched, yellow Leotia lubrica and it suggests some
unusual section of the genus. Possibly it presents an intermediate state from a
gelatinous fruit-body, as R. gelatinosa, with gelatinization restricted to the ultimate
tips, the capitate Leotia-like form of which seems to be caused by the gelatinization
of both the subhymeniurn and rnedullary tissue: the branches remain dry. Neverthe
less, Fawcett remarks that R. ochraceo-salmonicolor 'approaches rather close to this
species in the possession of a number of subglobose swellings at the tips, but these do
not differ in colour or texture from the rest of the plant'. If there are no clamps,
R. ignicolor may be related.
R. capucina (Pat.) comb. nov.
Basinyrn: Clavaria capucina Pat., Bull. Soc. Myc. Fr. 33, 1917, 50: Sacc., Syll.
Fung. 23, 1925, 487.
2-3 cm. high, yellow-ochraceous, fragile, fleshy: stern I cm. X 2-3 mm., brownish,
abruptly branched at the top: branches erect, slender, many times furcate, the tips
with 2-3 teeth: smell pleasant, slightly of aniseed: taste acrid,. almost salty.
On the ground near bamboos: lndo-China (Hanoi).
Spores 6 X 3 µ,, ochraceous, minutely verrucose, ellipsoid.
This is said to be near Lentaria byssiseda, but the resernbl�nce must be superficial.
It is doubtfully distinguishable, except by locality, from R. fiaccida or R. gracilis.
R. cervicornis (A. L. Smith) comb. nov.
Basinyrn: Clavaria cervicornis A. L. Smith, Joum. Linn. Soc. Bot. 35, 1901, 10.
-8 cm. high, flesh colour then cinnamon, velutinate-pruinose: stern -2 cm. high,
subterete: branches dichotomous, flattened, compressed, bushy upwards.
On rotten wood: Dominica.
Spores 8 X 6 µ,, yellow-brown, echinulate, ellipsoid.
This is said to be like a Thelephora in the flattened branches and brownish echinulate
spores, but the hyrnenium covers all sides of the branches. Flattened branching and
subglobose spores are exceptional in Ramaria and no other Echinosporae are ligni
colous. Until it is better known, therefore, its affinity cannot be guessed.
R. condensata (Fr.) Quel.
FI. Myc. 1888, 467.
Basinym: Clavaria condensata Fr., Ep. 1838, 575: Quel., Ass. Fr. 1880, 670: Bres.,
Fungi Trid. 1, p. 90, t. 101: le. Myc., t. 1092: (non Bourd. et Galz. = Ramaria
Bourdotiana et R. stricta).
566 RAMARIA
�ynonyms: Clavaria dendroidea Fr., Mon. Hym. Suec. 1857, 281: Lundell, Sv.
Svamp. 9-10, 1937, 12: (non Jungh. 1838 = Aphelaria dendroidea).
? Clavaria stricta (Fr.) Karst., Hattsv. 1882, 189: 1889, 467.
Clavariella condensata (Fr.) Karst., Hattsv. 1882, 184.
5-10 X 3-5 cm., alutaceous, tan-rufescent, or cafi-au-lait, becoming tawny or cinnamon
rufescent, tips clear yellow, old fruit-bodies reddish brown or ferruginous: stem
1-2 x 0·5-1 cm., generally indistinct, sometimes absent, soon divided into numerous
branches, pale alutaceous or ochraceous cream, whitish tomentose at the base and
attached to a white, floccose, mycelial felt: branches crowded, numerous, erect, parallel,
fastigiate, repeatedly dichotomous, tips acute: flesh white or paler concolorous, deep
yellow upward, unchanging, firm, tough: taste bitterish.
In humus of frondose (?) and coniferous woods: Europe (uncommon).
Spores 8-10(-12) x 4-5 µ., ochraceous, smooth, elliptic-amygdaloid, 1-guttate.
Basidia 30-40 X 4-5 µ., 4-spored.
This is C. condensata sensu Quel. and Bres. It seems to connect the Stricta-group
with the R. Strasseri-R. spinulosa series in the Decolorans-group.
Lundell's Swedish collections, identified as C. condensata, differ somewhat in these
respects:
Fruit-bodies as R. stricta, but not changing colour on bruising, arising from mycelial
strands -12 cm. long, adhering to bits of rotten wood: taste slightly acrid: smell rather
unpleasant, of aniseed and Tricholoma album: spores (6·5-)7-8·5(-<J) X (3·5-)4-4·5(-5) µ.,
pale ochraceous, indistinctly rough, oblong-ellipsoid, straighter and plumper than
typical R. stricta: basidia 18-22 X 6-7 µ.: 4 sterigmata, 4-5 µ.: near plants of Syringa.
This is the basis of his variety C. stricta var. condensata (Fr.). Now no other author
gives R. condensata as lignicolous like R. stricta, and R. condensata does not change
colour, apparently, like R. stricta, on bruising. Therefore, while I think that Lundell's
plants may be R. stricta, I disagree with this interpretation of C. condensata. R. stricta
has characteristically thick-walled hyphae and thickening hymenium: it has yet to be
shown that these features occur in R. condensata and, if not, it clearly belongs to the
R. spinulosa-alliance.
On the other hand, the Swedish specimens which Lundell identifies with C. den
droidea Fr. are certainly C. condensata sensu Bres., being humicolous, with smooth
spores 8-10 X 4-5 µ., unchanging flesh and yellow tips. The only difference of any
significance in Fries's descriptions of C. dendroidea and C. condensata is that the first
is placed among the 'truncigenae' and said to grow on branches and masses of leaves
in pine woods, and the second is said to grow on the ground in frondose woods. Thus,
C. dendroidea has often been considered as the yellow-tipped, lignicolous R. stricta,
e.g. C. dendroidea sensu Bourd. et Galz. The name C. dendroidea is, however, a later
homonym. Therefore I have retained C. condensata sensu Bres. for this humicolous
species. Karsten gave the spores of C. dendroidea as 8-12 x 4-5 µ..
C. spinulosa sensu Lundell (Sv. Svamp. 9-10, 1937, 13) also seems to me the same
as C. condensata. It is said to be dark reddish brown or smoky with obtuse then
subulate tips -3 cm. long, bright yellow when young, almost golden: spores
9-u(-12) X 4-4·5 µ., ochraceous, oblong-ellipsoid, nearly smooth, thick-walled: in
humus of coniferous woods, Sweden. For C. spinulosa Fries gives the tips as con
colorous, that is 'subcinnamomeis (fuligineo-spadiceis: Pers.)'. Lundell's fungus is not
C. spinulosa sensu Quel. or Rea ( = R. spinulosa).
R. conjuncta (Pk.) comb. nov.
Basinym: Clavaria conjuncta Pk., Bull. N.Y. St. Mus. 105, 1906, 16, 42, t. 102:
Kauffm., Pap. Mich. Ac. Sci. Arts Lett. u, 1930, 176: Doty, Clav. Pac. N.W. 1944, 56.
10-20 cm. high, cream colour to Naples yellow, deeper upward, young tips pinkish
yellow, soon yellow: stem usually buried in humus, dividing into a number of main
branches: rather fragile.
RAMARIA
On humus in coniferous forest: U.S.A. (Oregon, Wash.).
Spores 9-12X3·5-4µ,, pale yellowish under the microscope, minutely rough, sub-
cylindric: (8-10X3·5-4·5µ,, rough, Peck's type, teste Coker).
Basidia 80-85X5-6 µ,, 4-spored.
Cystidia none.
This is Kauffman's description of R. conjuncta which Coker reduced to R. botry
toides ( = C. botrytis sensu Coker). Kauffman and Doty agree with Peck that it is
allied with R. fiava.
R. conjuncta sensu Kauffm., R. divaricata, and R. subbotrytis are so closely related
as almost to form a continuous series. R. subbotrytis is pink (the pigment more or less
evanescent); R. conjuncta sensu Kauffm. is pale yellow with evanescent pink, and
R. divaricata is more persistently yellowish pink. R. conjuncta sensu Kauffm. and
R. divaricata have minutely rough spores and R. subbotrytis has nearly smooth shorter
spores, that is the juvenescent form of the series (as R. conjunctipes to R. formosa).
Therefore I have placed R. conjuncta sensu Kauffm. in Formosa. Nevertheless, the
problem arises whether the yellowish members of the R. subbotrytis-series, mentioned
on p. 140, are not related to the Flava-group rather than to Formosa.
R. conjunctipes (Coker) comb. nov.

Basinym: Clavaria conjunctipes Coker, Clav. U.S. Can. 1923, 131, t. 43, 91.
Synonym: ? C. Raveneliana Coker, Journ. El. Mitch. Sci. Soc. 63, 1947, 44, pl. 2, 12.
-7 cm. high, caespitose in gregarious clusters, salmon-yellow with lemon-yellow tips,
the old parts fading to cinnamon-buff: stems 1·5-2·5 mm. wide, slender, string-like,
fasciculate, not or scarcely thicker than the branches, arising from white rhizomorphs:
branches dichotomous, rather elongate, fastigiate, erect: flesh concolorous, flexible,
firm, rather tough: no smell or taste.
On the ground in frondose woods: U.S.A. (N. Car., Tenn., Conn.).
Spores 7-10 X4-5µ,, pale yellow, smooth or, perhaps, minutely rough.
var. odora Coker

i.e., I 32, t. 53, 86.
Stems -8 mm. thick, stouter: smell faint, fragrant: spores 5·5-6·3X4-4·8 µ,.
U.S.A. (N. Car.).
This resembles an attenuate state of R. formosa with the small spores of R. mb
botrytis but is without the hyphal property causing the chalkiness of the dried flesh.
Doty records a collection from Oregon with bright clear yellow fruit-bodies and
watery pinkish flesh: spores 7·5-8X 4·8 µ,, ochraceous (under the microscope), smooth:
in coniferous woods. Evidently the pink colour may be tardy in developing or wholly
absent. Compare, therefore, R. sinapicolor with similar habit.
There is a strange resemblance in smell, small size, colour, small spores, and
texture between R. gracilis and R. conjunctipes: careful comparison of them would be
instructive. C. Raveneliana seems to be young R. conjunctipes.
Coker's spore-data for R. conjunctipes are these:
N. Car. 5732. 7·4-8·5x4-4·5µ, (type)
5734 . 6·8-8·6X4·2-4·8µ,
,, 5768 . 6·5-8x4-4·8µ
Tenn. (Murrill) 7-8X 4·8-5 fL
Conn. (Coker 36) 7·4-10 X 3·8-5µ
R. cornicularioides (P. Henn.) comb. nov.

Basinym: Lachnocladium cornicularioides P. Henn., Monsunia 1, 1900, 143.
-2 cm. high, coriaceous, setulose, with a long root, pale yellow to brownish white
(dried), substipitate or caespitose-ramose from the base: branches widely compressed,
568 RAMARIA
subfasciate, short, repeatedly dichotomous, squarrose, with subulate or cristate
apices.
On the ground: Java (Tjibodas).
Spores 7-8 X 4µ,, brown, ovoid or ellipsoid, smooth, apiculate.
Very much branched, compact and small, with compressed branchlets. Said to
resemble superficially Cornicularia aculeata. If correct, the spores indicate Ramaria.
R. crispula (Fr.) Quel.
Fl. Myc. 1888, 464.
Basinym: Clavaria crispula Fr., Syst. Myc. 1, 1821, 470.
Synonym: Clavaria decurrens Pers., Myc. Eur. 1, 1822, 164 (fide Fries): (non sensu
Coker= Ramaria pusilla).
Clavaria muscigena Pers., Myc. Ei.ir. 1, 1822, 169 (fide.Fries).
Clavariella crispula (Fr.) Karst., Hattsv. 2, 1882, 189.
1·5-5 cm. high (? -7·5 cm.), tan then ochraceous, or ochraceous tinged alutaceous or
bistre-cream, tips concolorous or paler in old specimens, caespitose, much branched:
stem 1-3 mm. wide, slender, villous, with copious rhizomorphs at the base: branches
flexuous, multifid, divaricate, acute, often setaceous: flesh rather tough, unchanging.
On fallen trunks and logs of frondose trees, and on the surrounding earth: Europe
(not common), S. Austral. (Victoria). 1
Spores 5-7·5 X 3-4µ, (Bourd. et Galz.), 5-6 X 3µ, (Karst.), 6-7 X 3·2-4µ, (Fawcett),
ochraceous, laxly asperulate or finely rugulose.
Basidia 26-3 3 X 4·5-6µ,.
Hyphae 2-12µ, wide, thin-walled.
This is C. crispula sensu Karst. and Bourd. et Galz. It seems a lignicolous species
allied with R. stricta, but the thin-walled hyphae (if correctly given, for the flesh is
said to be tough) would place it with R. fiaccida. Bourdot and Galzin say that it never
grows on coniferous wood. Fawcett describes and illustrates it from Victoria as
growing in colonies among pure wood debris.
Coker identified C. decurrens with C. pusilla of the United States but, as shown on
page 618, there is no decisive evidence that Fries was wrong in reducing C. decurrens
to C. crispula. Coker gives the spores of Persoonian specimens of C. decurrens as
4·5-6 X 2·5-3·5µ,, and of Persoonian specimens of C. crispula as 4·5-6·5 X 2·5-:-4/L,
which agree well enough with Bourdot and Galzin's measurements.
R. cyanocephala (B. et C.) comb. nov.
Basinym: Clavaria cyanocephala B. et C., Journ. Linn. Soc. Bot. 10, 1869, 338: ...
Coker, Clav. U.S. Can. 1923, 191, t. 90.
Synonyms: Clavaria aeruginosa Pat., Bull. Soc. Myc. Fr. 14, 1898, 189: fide v.
Hohn, Sitzb. d. Kaiser!. Akad. Wiss. Wien 118, 1909, 15.
Lachnocladium giganteum Pat., Journ. de Bot. 3, 1889, 34, t. 1, f. 1 (an Ramaria
Zippelii?): Martin, Mycol. 31, 1939, 515.
L. insigne Pat., Journ. de Bot. 3, 1889, 34, t. 1, f. 2: (an R. Zippelii?).
Lachnocladium Englerianum P. Henn., Nat. Pfl. Fam. 1, 1897, 138, f. 73.
? Thelephora angustata Fr., Act. R. Soc. Sc. Upps. 3, 1, 1851, 108, fide Bres., Ann.
Myc. 14, 1916, 241:= Thelephora fide Burt, Ann. Mo. Bot. Gdn. 1, 1914, 199.Z
? Lachnocladium olivaceum P. Henn., Hedw. 1897, 196.
Pseudonym: Clavaria Zippelii auctt., non Lev. ( = Ramaria Zippelii): v. Ov., le.
Fung. Mai. 1, 1923, t. rn, 1b (ut Phaeoclavulina Zippelii).
1
The records from Brazil and New Zealand need confirmation. Cleland and Chee! (p. 869)
record it from New South Wales (Mt. Irvine) as pale orange, on a fallen trunk, spores 7 X 4·2 µ.,
white. Their fungus cannot be R. crispula.
2
If this is a Ramaria, it will be impossible to decide from a dried specimen whether it is
R. Zippe/ii or R. cyanocepha!a or R. {!mndis.
RAMARIA 569
6-30 X 5-15 cm., much branched, gregarious, rarely caespitose, fawn-colour,
brownish ochraceous or ochraceous cinnamon, then more or lessfuscous-umber, often tinged
cinnamon-ferrugihous from the spores, tips intense blue or greenish blue, all parts quickly
turning dingy vinous-purple on bruising: stem 0·6-6·5 X 0·3-1·6 cm., cylindric or slightly
TEx.T-FIG. 25 I. Ramaria cyanoceplzala: two immature spores on the left:

X r,ooo.
widened upward, relatively slender, concolorous, generally attenuate at the white base
subvillous with white mycelium: branches 5-15 .mm. wide below, generally regularly
dichotomous with successive branchings more or less at right angles, the internodes gradually
more elongate distally, short and obscure proximally, 1·5-2·5 mm. wide at the tips,
cylindric, erect, fastigiate, compact, crowded, sometimes spreading, sterile and sub
tomentose on the adaxial surfaces and on the uppersides of the axils, tips subacute,
other fruit-bodies with looser, more irregular branching, sometimes 3-4-chotomous
1;,elow: flesh whitish or pale yellowish, deeper yellow next the hymenium, quickly turning
dingy vinous-purple on exposure, rather dry but fibroso-brittle, breaking when bent at
90° , putrescent: taste slightly bitter and astringent: smell slight or none: mycelium
white, turning dingy purple on bruising.
TABLE XXXIX. SPORE-SIZE IN RAMARIA CYANOCEPHALA AND R. ZIPPELII
Spore-size in µ,
(spore-body) E' Authority Remarks
R. cyanoceplzala
12-17·5X7·5-,-10 1·7 Corner Pahang: see Table XLI
r2-16X7·5-8 I·8 Coker Jamaica
10-16X6-8 1·9 Petch Ceylon
II-IS X6-8 1·8 Corner Pahang: see Table XL
11·5-14·5 X7-8·5 1·6 " Penang: see Table XL
II-I4X5' 5-7·4 1·9 Coker Cuba (Wright No. 458, ? type)
9-15X6-7•5 1·8 " Porto Rico
10-13X5-6·5 2·0 Teng China
R. Zippelii
13-18.x7-9 1·9 Corner New Guinea (Carr No. 11201, Kanosia)
13-15X7-8 1·9 Bresadola Java
10-15X5-8 1·9 van Overeem Java
I1-14X5-7 2'1 Corner Malaya (4 collections)
12·5-17·5X7-10 1·8 Ito and Imai Bonin Islands (? = C. cyanocephala)
9-11x6-6·5 1·6 Bresadola Borneo (Lachnocladium echinosporum)
var. gracilis
8-10·5X5--6·5 1'6 Corner Johore (type): 2 collections: 2-3, mostly 3-, spared.
9-1oxs·s-6 1·7 Johore, Pahang: darker ferruginous brown fruit
bodies with white, unchanging flesh: 2-3,
mostly 3, spared..
1'6 Singapore (3-4-43): fawn ochraceous: flesh turn
ing vinous: 4-spored (rarely 2): no smell.

..
FIG. 252. FIG. 253.
TEXT-FIG. 252. Ramaria cyanocephala: thickening hymenium, X 500.

ToxT-FIG. 253. Ramaria cyanocephala: sterile hymenium on the upperside of an oblique
branch, x 500.
RAMARIA 57 1
On the ground in the forest: trop. Asia (Bonin Is!., Hainan, Ceylon to New Guinea),
trop. Amer. (West Indies, Panama), ? trop. Africa, S. Africa; apparently throughout
the tropics (? subtropics).
TABLE XL. SPORE-SIZE IN RAMARIA CYANOCEPHALA
I
One fruit-body, Penang (I940) One fruit-body, Pahang (4.n.30)
I
Spore-size in µ' E' Number of records Spore-size in µ�1-
�
Number of records
II·5X7·3 1·6 2 Il'OX6·5 1·7 2
11·5X7·5 1·6 11·ox7·0 1·6 2
I
5
11·5X7·7 1·5 I 11·5 x6·5 1·8 I
11·5x8·0 1·4 I 11'5Xio 1·6 I
I
11·5X8·5 1·4 r 11·5X7·3 1·6 2
12·ox7·5 1·6 3 11·5X7·5 1·6 2
12·0 X8·o r·5 2 12·0X6·0 2·0 I
12·0 X8·3 1·4 2 12·0X6·7 1·8 2
12·5X 8·3 I' 5 3 12·0X7·0 1·7 3
13·0X7·5 1·7 3 r2·5X7·0 I ·8 I
13·0 X7·7 1·7 I 12·5X7·3 1·7 l
13·ox8·0 r·6 I 12·5X7·5 1·6 2
13·0X7·0 1·9 I
13·5X7·0 r·9 I 13·0X7·5 1·7 2
13·5X6·5 2·r 2
13·5X7·5 1·8 I 13·5X6·7 2·0 !
r3·5X7·0 1·9 2
I
I
13·5XS·o r·7 2 r3·5X7·5 1·8 2
r4·0 X6·5 2·2 3
13·5X8·3 I·6 I 14·0X7·0 2·0 I
14·0 X7·3 1·9 I
14·0X7·7 1·8 I 14·0X7·5 1·9 4
14·5X7·0 2·r r
14·ox8·o 1·8 2 r5·0 x6·7 2·2 2
r4·0X8·3 r·7 I r5·0X7·0 2·r 2
r5·0X7·5 2·0 l
r4·0 X8·5 1 ·6 3 15·0X7·7 r·9 l
15·0X8·o r·9
r4·5X8·o 1·8 2
14·5X8·5 1·7 I
11·5-14·5X7-8·5 1·4-1·y 40 II-15X6-8 l ·6-2·2 50

11-11·5 long 1·6 Average of Io 11-n·5 long 1'6 A,·erage of 10
12-12·5 " I' 5 " 12-12·5 " 1·7 "
I3-13 '5 " 1·7 " 13-13·5 " 1·9 "
14-14·5 " 1·7 " q-14·5 " 2·0 "
12·8X7·9 1·6 Average of 40 15 long z·o "
13·2X7·1 I·8 Average of 50
I
1
Spore-body, excluding spines. 2
Spores 10-17·5 X 5·5-10µ (spore-body), fuscous ferruginous ochre, rather densely

echinulate with conical, acute, hyaline spines 1-3µ long, pip-shaped or amygdaliform
(mango-shaped), several guttulate: (at first colourless and smooth, then with coloured,
smooth wall becoming verrucose and, finally, echinulate).
Basidia 55-75 X 9-11 µ, subclavate, generally somewhat constricted below the
sterigmata, with a long narrow base 3µ wide, contents granular-guttulate and pale
ochraceous: sterigmata 2, occasionally 3 or 1, 6-8(-10)µ long.
Sterile basidia like the fertile basidia but never with spores, sterigmata often abortive
572 RAMARIA
or none, cytoplasm densely and rather darkly brownish granular-guttulate, the walls
often pale ochraceous but not thickened; abundant in the fertile and sterile hymenia.
Hymenium 60 µ. thick at first, gradually thickening to 600 µ. (showing macroscopically
as a fuscous umber line in section o( the fruit-body): subhymenium c. 30 µ. thick,
composed of 3-6 µ. wide hyphae, not or but slightly inflated, the cells 10-30 µ. long:
old basidia collapsing, but with many sterile basidia persistent in the hymenium, like
those of the sterile hymenium.
TABLE XLI. SPORE-SIZE IN RAMARIA CYANOCEPHALA
(One fruit-body; Pahang, I9JI)
Et E'
--
Spore-size in µ.
--
Number of records Spore-size in µ.
----
Number of records
u·ox8·o I'S 4 14·5X9·0 r·6 2
I2'0X8·5 r·4 I 14·5X9·3 1·6 I
12·5X7·5 r·6 I 14·5X9'5 1·5 I
12·5X7·7 1·6 I 15·0X8·o 1·9 I
12•5X8·o 1·8 I 15·0X8·3 1·8 2
12·5X8·3 1·5 I 15·0X8·5 1·8 2
12·5x8·5 1·5 I 15·0X9·0 r·7 2
13·0X8·5 I 'S I 15·0X9'5 1·6 l
13·0X8·7 1·5 I 15·5X8·3 1·9 I
13·5x7·5 1·8 I 15·5X8·5 r·8 I
13·5X8·o 1·7 2 16·ox8·0 2·0 l
13·5X8·3 1·6 I 16·0X8·7 1·8 I
13·5X8·5 1·6 2 16·0X9·0 1·8 I
13·5X9·0 1·5 I r6·ox9·3 r·7 I I
13·5X9·3 I '5 I r6·0X9·5 1·7 I
14·0X8·o r·8 I r6·5 X9·0 1·8 I
14·0X8·3 1·7 I 16·5X9·5 r·7 I
14·0X8·5 1·6 I 16·5X10·0 r·7 I
r4·0X9·0 r·6 2 r7·ox8·o 2·r I
14·5X8·3 r·7 I 17·5X9'7 I ·8 I I
Range: r2·0-r7·5x 7·5-ro·o µ., E = r·4-2·r, for 50 spores.

Average of ro spores r2-r2·5 µ. long, E = r·s
13-13·5 µ. ,, E = r·6
14-14·5µ. ,, E = 1·6
15-15·5 µ. ,, E = 1·8
,, ,, 16-17µ. ,, E = r·8
Average of 50 spores 14·3X8·6 µ., E = r ·7
E = length --:- width.
Sterile hymenium on the upper or adaxial parts of the branches and on the upper
part of the stem, composed of 3-6 µ. wide excrescent hyphae with numerous sterile
basidia, thickening but not so extensively as the fertile hymenium.
Hyphae 2·5-10µ. wide, -13µ. in places, mostly 3-6 µ. wide in the stem, clamped,
cells 50-200µ. long, cylindric, often rather unevenly inflated, with colourless or pale
yellow, thin or very slightly thickened walls (-0·5 µ.), smooth, compact, longitudinal,
with abrupt transition to the subhymenium, H-connexions frequent: hyphal tips at
the growing points 2-3 µ. wide, very gradually inflating.
Yellow-umber colour caused by the slightly ochraceous walls of the hyphae, the
pale yellowish oil-drops in many of the internal hyphae, and the brownish ochraceous
cytoplasm and guttulae of the sterile basidia: blue colour of the growing tips caused
by blue or greenish blue guttulae in the hyphae.
R. cyanocephala has been regarded as synonymous with R. Zippe/ii (Bresadola, v.
Hoehne!, Coker, van Overeem), but there is no proof that they intergrade, nor ha\'c
RAMARIA 573
the differences between them been clearly stated. I have found both species in Malaya,
R. cyanocephala being the commoner, indeed, one of the commonest Clavarioid fungi,
and it is always distinguishable by its brownish umber branches with blue, or greenish
blue, tips, so that it is one of the most striking species of the genus. R. Zippe/ii has
generally smaller fruit-bodies of a fuliginous ochraceous colour with white tips and
often, but perhaps not always, a stronger smell than R. cyanocephala: it may also have
rather smaller basidia, narrower hyphae (giving a tougher texture) and more often
polychotomous branching. The differences are slight and their spores are identical,
within wide limits. Nevertheless, I have seen no intergrades, though I have found
R. cyanocephala two or three times every year from 1929 to 1943. In Java, van Overeem
says that he never (or rarely?) found the blue-tipped species, but that R. Zippelii was
common: however, he illustrates both under the one name. In Ceylon, Petch found
only the blue-tipped species and not the white-tipped. R. Zippelii seems, further, to
vary into a smal!er-spored state which I have called var. gracilis and, thus, it approaches
the white-tipped temperate species, R. grandis, which indeed is doubtfully distinct.
To unite R. cyanocephala and R. Zippelii would, therefore, be confusing, and I have
called the white-tipped species R. Zippe/ii because in its original description there is
no mention of the blue tips clearly described for R. cyanocephala ('branches furcate,
intensely coeruleous above', B. et C.).
Coker gives Wright No. 458 (Cuba) as the type of R. cyanocephala, but in the
original description the only locality given is 'Bonin Islands', though it was described
in a paper on Cuban fungi. The exact type matters little, for the species is clearly
widespread throughout the tropics.
It is impossible to identify correctly, as R. cyanocephala or R. Zippe/ii, the tropical
American species of Lachnocladium described by Hennings and Patouillard with
brown echinulate spores. If R. Zippelii does not occur in tropical America, then they
must be R. cyanocephala. The colour of only the dried fruit-body has been described.
Compare also R. guyanense and R. leucoceras.
R. cyatheae (P. Henn.) comb. nov.
Basinym: Clavaria Cyatheae P. Henn., Sacc., Sy!!. 9, 1891, 250.
17-26 mm. high, simple, solitary or gregarious, spongy-fleshy, elongate-clavate,
acute, white then somezchat flesh-colour, head 7-11 X 1-1·5 mm.: stem 10-15 X I mm.,
glabrous, yellowish-white: spores subglobose, ferruginous.
On trunks of Cyathea Burh�i, from S. Africa, in Berlin Botanic Garden.
Very doubtful. Compare Laclmocladium Hisinieri, R. filicicola, and R. filicina.
R. cystidiophora (Kauffm.) comb. nov.

Basinym: Clamria cystidiophora Kauffm., Pap. Mich. Ac. Sci. Arts Lett. 8, 1928,
147: Doty, Clav. Pac. N.vV. 1944, 57.
ro-12 cm. high, caespitose, buff-yellon· to zcarm buff (Ridgway), tips citron-yellow,
whitish at the base: stem 5-8 X 1-1 ·2 cm., immersed below, sometimes tufted,
attenuate downward, ascending or decumbent: branches numerous, elongate, cylindric
or nearly so, 5-6 primary branches, branchlets 3-4 mm. wide: flesh white, moderately
fragile: taste mild: smell of aniseed.
In deep humus of woods of Douglas fir, cedar, and hemlock: U.S.A. (Lake Quiniault,
Washington).
Spores 5·5-7 X 3 ·5 µ,, tinged ochraceous-buff under the microscope, smooth or
obscurely punctate, oblong: 7·5-8 X 3·5 µ,, asperulate (Doty).
Basidia 75-80 X 5-6 f-L, 4-spored.
Cystidia 50-70 x 5-8 µ,, scattered, cylindric upwards, narrowed below, obtuse,
collapsing in old plants.
This may be only R. flm:o-brunnescens with unusually small spores. The cystidia
seem to be young excrescent basidia or possibly sterile basidia on the uppersides of the
574 RAMARIA
branches. Cystidia have not been found in any other species of the genus: nor has the
association of massive fruit-bodies and small spores.
Doty compares it with R. obtusissima with lighter colour and larger spores.
R. decolorans (Karst.) comb. nov.
Basinym: Clavariella decolorans Karst., Symb. Myc. Fenn. 32, 1893, 10.
Synonyms: Clavariellafennica Karst., Finl. Nat. o. Folk 50, 1891, 223: non Karst.
1882 = Ramariafennica.
Clavaria decolorans (Karst.) Sacc., Syll. 11, 1891, 135.
9 x6 cm., pallid white then yellowish or light yellow, tips clearer yellow, turning violet
on bruising: stem 1-2 cm. thick, rather stout, whitish at the base: branches numerous,
much branched, dichotomous or verticillate, -spreading, smooth or rugulose: branchlets
spreading or upright, often arcuate at the base, irregularly polychotomous or dichoto
mous: flesh pallid, rather tough, drying brownish or violet-alutaceous, bitter.
On the ground in woods: Finland.
Spores 10--20 X4--6µ, light yellowish (under the microscope), elongate-fusiform,
guttulate (? smooth).
Basidia 60 x 7--9µ.
This rare species appears to be the same as R. xanthosperma of N. America. They
indicate a northerly species with slightly pigmented fruit-bodies and the same colour-
change of the bruised tissue as in R. formosa.
R. divaricata (Pk.) comb.. nov.

Basinym: Clavaria divaricata Pk., Bull. N. Y. St. Mus. 2, 1887, 11: Coker, Clav.
U.S. Can. 1923, 126, t. 40, 85.
Synonym: ? Clavaria fiava var. subtilis Coker, Le. 125, t. 85.
5-10 cm. high and wide, branched at the ground from a pointed base, yellowish flesh
colour, paler below, tips very pale, then concolorous, apparently the branches becoming
yellow towards the tips or wholly yellowish in some cases: branches rather open,
spreading, with many acute, rather divaricate tips: flesh concolorous, very fragile,
brittle: taste pleasant, nutty: smell slight.
Spores 9-12·5 X 3·7-5 µ., ochraceous buff, minutely rough.
Basidia 6--9· 5 µ wide: hymenium 50--60 µ. thick.
It is difficult to see how this differs from R. subbotrytis var. intermedia except in its
paler spores and yellowish pink colour: perhaps var. intermedia is a mixture.
C. fiava var. subtilis' seems to differ only in having the bJ.:anches pale clear yellow
upward and white flesh: it also resembles R.jormosa, but neither R. divaricata nor this
var. subtilis have the chalky dried flesh of R. formosa. Coker ascribes to R. divaricata
a collection (No. 1250) with wholly clear light-yellow, nearly citron-yellow, fruit
bodies, white rigid brittle flesh and pleasant nutty taste, which makes the description
of the species embrace that of var. subtilis.
Compare also R. obtusissima and R. secunda with smooth spores.
The following are the spore-measurements given by Coker:
No. 3063 9·7-11·5 X 3·7-4·5 µ. C. divaricata
No. 3037 9-11 x4-4·5µ
II·7-12·5 X4-4·7 µ. ,, (type)
No. 2843 10-II X4-4·5µ C. fiava var. subtilis (type)
Burnham No. 103 9·3-11·8 X3·8-4·6µ " subtilis
1
C. flava var. subtilis Coker: stem white, pointed, glabrous: branches rugose, pale fleshy
cream, clear pale yellow upward (near apricot-yellow): flesh nearly or quite white, drying
hard (not chalky or fragile): taste mild, acid-bitter: smell slight: spores 9·5-12 X 3·8-4·5 µ,
ochraceous-yellow, distinctly rough: basidia 7-9 µ wide, 4-spored: hymenium 55-70 µ thick:
hyphae 7-11 µ: on the ground in frondose woods (N. Car., N.Y.).
RAMARIA 575
R. durbana (van d. Bijl) comb. nov.
Basinym: Clavaria durbana van d. Bijl, S. Afr. Jour. Sci. 29, 11)3�, 322',. t. 3; f. 4':
-2·5 X 1·5 cm., caespitose, flesh-coloured, densely branched: stem 6 X 2 mm., short,
distinct: branches crowded, cylindric or compressed.
On dead wood: Natal (Durban).
Spores 6 X 3µ, pale ochraceous or subhyaline, distinctly rough, ellipsoid.
This is said to be like R. gracilis in colour and form, but with rougher spores. 1 think
it may be Scytinopogon. Compare R. ochracea.
R. eumorpha (Karst.) comb. nov.
Basinym: Clavariella eumorpha Karst., Symb. Myc. Fenn. 9, 55: Hattsv. 2, 1882,
185: Finl. Nat. o. Folk 48', 1889, 388.
Synonym: Clavaria eumorpha (Karst.) Sacc., Sy!!. Fung. 6, 1888, 701.
6-9 cm. high, pale ochre-yellow: stem 3 x 0·5 cm., elongate, narrow: branches:..
longitudinally rugose, very abundant, strict, crowded.
On the ground in woods (Acer): Esthonia (Mustjala).
Spores 7 X 4µ, ochre-yellow, verruculose, ellipsoid.
This may be only the well-developed, or less degenerate, state of�- fiacfidf.l•,.Rea
also describes R. fiaccida as 3-10 cm. high, but other authors emphasize its shlalli'size.
' .. ..
R. fennica (Karst.) Ricken · ·
' ., ·,.,, (� • I .. , ' •
· .
Vadem. 1920, 264.
Basinym: Clavariafennica Karst., Nat. Sallsk. Faun. et Flor. Fenn. 9, 1868, 372:
Bidr. Finl. Nat. o. Folk 48, 1889, 389: Bres., Fung. Trid. r, 24, t. 27: le. Myc., t. 1091:
FI. Bat. 27, 1928, t. 2135: (non C. fennica Karst. 1891 = R. decolorans Karst.: non
C. Jennica sensu Coker = R. Jumigata).
Synonym: Clavariellafennica Karst., Hattsv. 2, 1882, 184.
Text-Fig. 249.
6-12 x 7-8 cm.: stem 2-5 cm. thick, violaceous, white at the base: branches lurid olive,
Juliginous-yellow or yellowish olivaceous, then yellowish bistre, tips bright yellow, fuliginous
or blackening with age, very numerous, many arising at the top of the stem, cylindric or
subcompressed, longitudinally rugulose, erect: flesh white, rather firm, not fragile:
taste slightly bitter: smell none.
In humus of coniferous woods: Europe (Finland, Sweden, Germany, France,
Holland, Austria, northern Italy, Spain), rather rare.
Spores 8-12·5 X 4-5·5µ, ochraceous yellow or light ferruginous-ochraceous,
minutely rough, oblong ellipsoid, I-several guttate, apiculus 1-1·5µ long, conspicuous.
Basidia 40-50 X 6-7µ, 4-spored.
Hymenium not thickening, c. 45µ thick: subhymenium composed of 2-3·5µ wide,
interwoven hyphae.
Hyphae 2-8µ wide, thin-walled, clamped, as in R. formosa, rather densely inter
woven in the stem.
This species is close to the less uncommon R. fumigata, but differs in the olivaceous
branches, the tips of which are bright yellow when young and in full growth, though , ,. , '"' ·�
eventually they become concolorous. The spores of both seem identicaj, . . ,. ., ,, . "·• ....,,, .1 r : •
I have been able to study the microscopic structure of the fruit-body from dried'
specimens sent to me by Dr. Karl von Keissler from Austria (Pressbaum, 1925): they
were described as having violet stems and olive branches when fresh. The structure
is identical with that of R. Jormosa, from which the species differs only in the form of
the fruit-body with the many branches congested at the top of the stem, the flesh not
drying chalky-friable nor changing colour when fresh (so far as known), and in the
different colour of the fruit-body. The yellow tips of R. fennica clearly prove its
affinity with R. formosa, and the olivaceous colour of its branches is caused perhaps
by the clouding of the yellow pigment with the violaceous pigment characteristic of
llAMARIA
R. fumigata. I find the hyphae to be clamped at every septum, though Donk reports
them without clamps.
The following are the spore-measurements of R. fennica:
Spore-size in µ, Authority Country Spore-colour

8x4 Karsten (1889) Finland ochre-yellow
8-10X6-7 1 Bresadola N. Italy ochre-yellow
9·5-12·5X4-5 Donk Holland ochraceous
9-12X4·5-5·5 Comer Austria light ferruginous-ochre
1
? An error in width.
R. filicicola (Fawcett) comb. nov.

Basinym: Clavaria filicicola Fawcett, Proc. Roy. Soc. Viet. 52, 1940, 155, t. 6,
text-fig. I.
2-4 cm. high, small, variable in form, pure white when young, becoming maize
yellow to ochraceous buff: stem stout, soon divided, often villous from the mycelium,
arising from a dense white mycelium binding together the fibres of the Dicksonia-trunk:
branches stout below, dividihg into many crowded branchlets with subulate tips, or
sparingly branched: flesh solid, brittle, soft.and pliable with age: smell and taste none.
On trunks of Dicksonia antarctica, usually just below the crown of leaves: S.
Austral. (Victoria, Turton's Pass, Apollo Bay).
Spores 5·6-7 x 3·6-3·9µ, maize yello;w, rough or verruculose, shortly ellipsoid.
This belongs in the difficult group Verrucispora. It may be identical with R.
megalorhiza. Fawcett compares it with R. gracilis, of coniferous forests, but it lacks
the characteristic smell of aniseed. Compare R. filicina and R. luteo-fiaccida.
R. filicina (Sacc. et Syd.) comb. nov.

Basinym: Clavariafilicina Sacc .. et Syd., Syll. Fung. 14, 238.
Synonym: Clavaria cervina W. G. Smith, Joum. Bot. 1873, 66, t. 130, f. 9 (non
. C. c�ina B. et C. 1868, see Ramariopsis Kunzei).
Slender, little branched, fawn-colour· with brown base: branches obtuse, stuffe<;l ·
spores ochraceous..
On trunks of ferns in a hot-house: England (Royal Horticultural Society, South
Kensington).
Probably unrecognizable, but compare R. filicicola and Lachnocladium Hisingeri.
R. ftaccida (Fr.) Ricken

Vadem. 1918, 254.
Basinym: Clavariq fiaccida Fr., Syst. Myc. 1, 1821, 471: Cokei:, Clav. U.S. Can.
1923, 184, t. 70 (ut C. abietina 'small non-virescent form'): Wehrn., Pap. Mich. Ac.
Sci. Arts Lett. 20, 1935, 259: Chow, Bull. Fan Mem. Inst. Biol. 6 (Bot.), 1935, 162:
Fawcett, Proc. Roy, Soc. Viet. 51, 1939, 156 (ut C. abietina).
Synonyms: Clavariella fiaccida (Fr.) Karst., et var. dendroides Karst., Hattsv. 2,
1882, 186.
Clavaria corrugata Karst. (? ed.).
Clavariella corrugata Karst., Hattsv. 2, 1882, 186.
1·5-6 X 0·6-4 cm., tilutaceous or pale cream-ochraceous, becoming bright ochraceous,
brownish ochre or cinnamon (even light orange-yellow, Wehm.), slender,fiaccid, growing
from a white, floccose, mycelial felt: stem 0·5-1·5 cm. X 1-2·5 mm., or branched from
the base: branches numerous, erect, crowded, one to three times divided, incurving,
tips becoming acute, paler concolorous: flesh white, yellowish upward, unchanging,
tough and elastic, then flaccid: smell and taste inparticular, or slightly fragrant and
subacrid (Wehm.).
RAMARIA 577
On coniferous needles, rarely on frondose humus, chiefly unc;ler Picea, Abies, and
Tsuga: Europe (uncommon), U.S.A. (frequent in the nort:hfm States), Canada,
Japan, China, S. Austral.' (under Callitris propinqua and Pinus radiata), S. Africa
(Natal, Rhodesia, Boschberg), (? Dominica, teste Lorrain Smith, Linn. Soc. Bot.
Journ. 35, 1901, 10).
Spores 5-8 X 3-4µ, ochraceous, verruculose, ellipsoid or pip-shaped: (brownish,
minutely spinulose, Wehm.).
Basidia 20-30 X 4·5-7µ (-<)µ, Cott. et Wake£.), 4-spored.
Hymenium 45-60µ thick, thickening slightly (Coker).
This species is near R. ochraceo-virens, but differs in not turning green and in the
flaccid consistency. From R. gracilis it differs in the deeper ochraceous or brownish
colour and in the lack of the aniseed-smell. R. crispula is lignicolous, paler, and
sparingly branched. Nevertheless, the exact differences between the four need further
analysis, description, and illustration.
C.fiaccida sensu Rea appears to be another species with larger fruit-bodies -10 cm.
high, growing in woods and on heaths. It may be R. eumorpha, which seems to differ
only in the larger fruit-bodies.
C. corrugata, according to its description, differs in no way from typical small
R. fiaccida.
The following are the published spore-measurements of R.fiaccida.
TABLE XLII. SPORE-SIZE IN RAMARIA FLACCIDA
Spore-size in µ. Authority I Country Remarks

7-9 X 3·5-4·5 Chow China verruculose
6-8·5 X 3-4 Wehmeyer Canada minutely spinulose
6-8x,3-5 Cott., Wakef. Gt. Britain average 6-7 X 3 µ.
6-8 X 3-4 Karsten Finland as C. corrugata
5-8 x3-,..4 Bourd., Galz. France granular-rough
6-8x3 van der Bijl S. Africa
5·5-7·5 X 3·7 Cleland S. Austral.
6·2-8·1 X 3·1-4·3 Fawcett Victoria verruculose
5-'J X 3-3·5 Imai Japan "
4·8-7·7 X 2·5-4 Coker U.S.A.
5·5-7 X 3-3·7 " Sweden Friesian specimen at Uppsala
5-'J X 3·4-3·7 " " " " at Kew
5-6 X 2•8-3·7 " " " n in Herb. Curtis
5·5-6·5 X 2·5-3·3 " Finland Karsten's specimen of C.jlaccida
R. flava (Fr.) Quel.

FI. Myc. 1888, 466.
Basinym: Clavariafiava Fr., Syst. Myc. 1, 1821, 467, quoad ic. Schaeff. (non Fr.
1838, 1874 = R. aurea): Bres., le. Myc., t. 1087: Konr. et Maubl., le. Se!. Fung. 1926,
t. 491: Maire, Bull. Soc. Myc. Fr. 27, 190, 450: (non Coker= Ramaria fiavo
brunnescens).
Synonyms: Clavaria fiava var. divaricata Pers., Myc. Eur. 1, 1822, 162: Baria,
Champ. Nice, 1859, t. 40, f. 5.
Clavaria lutea Vitt., Descr. Fg. Mang. 1835, t. 29, f. 3 (fide Bres.).
Clavaria lutea Vent., Misc. Bresc. (1845-60), t. 41, f. 4 (fide Bres.).
Clavariella flava (Fr.) Schroet., Krypt. FI. Schles. 3, 1888, 447.
(Clavaria sanguinea Pers. Obs. 2, 1799, t. 3, f. 5: non Coker = Ramaria sanguinea).
Text-Fig. 248.
1
Massee's record from New Zealand needs verification.
5119
Pp
578 RAMARIA
10-20 X 7-15 cm., sulphur-yellow or lemon-yellow, ageing ochraceous: stem 5-8 X
4-5 cm., whitish at the base, often reddish or blood-red on bruising or with age: branches
crowded, compressed, fastigiate, polychotomous below, dichotomous above, rather
thin, with rather long internodes: flesh white or pale yellowish, brittle, fragile, watery
marbled, often reddening near the surface when bruised, particularly in the stem and
main branches (not drying chalky-friable): taste mild, edible.
TABLE XLIII. SPORE-DATA FOR RAMARIA FLAVA
Size in µ. Authority Country Remarks

I------- -------- -
12-20X4-6 Corner Austria I See Table XLIV: coarsely rough
11-20X 4·5-6·5 " " See Table XLV: coarsely rough
11-15 X4-5 Cott., Wakef. England granular
9-12 X4-5 Rea " verruculose
10-14X4-5 Bresadola Italy scarcely rough
9-12 X4-5 Konr., Maubl. France verruculose
9-14X4-5 Bourd., Galz. (flesh not reddening)
TABLE XLIV. SPORE-SIZE IN RAMAJUA FLAV.11

(One fruit-body,· Rekawinkl, Austria, Sept. I925)
Size in µ. E' Number of records Size in µ. E' Number of records
----
12·0X5·0 2·4 3 15·0X4·7 3·2 I
12·5X4'7 2·7 I 15·0X5·0 3·0 3
12'5X5·0 2·5 I 15·0X5'3 2·8 3
i3·0X4·0 3·3 I 15·0x 5·5 2·7 I
13·0X4·3 3·0 I 15·0X6·o 2·5 I
13•0X4·5 2·9 I 15·5x 5·0 3·1 I
13·0X4·7 2·8 I 15·5x 5·5 2·8 I
13•0X5·0 2·6 I ' 15·5X5·7 2·7 I
13•0X6·o 2'2 I 16·0 X4·0 4·0 I
13•5X4·7 2·9 3 16·0X4·5 3·6 I
13;5x 5·0 2·7 7 16·0 X4·7 3·4 2
13•5X5'3 2·6 2 16·ox 5·0 3·2 4
13·5x 5·5 2·5 I 16·0X5·3 3·0 2
13'5X6•o 2·3 I 16·0X5·5 2·9 I
14·0X4•0 3·5 I 16·5X4'7 3·5 2
14•0X4·5 3·1 2 16•5x 5·0 3'3 I
14•0X4•7 2·9 I 16·5X5'3 3·1 I
14•0X5•0 2·8 2 16·5x 5.5 3·0 I
14•0 X5'3 2·6 2 17·0 x 5·0 3'4 3·
14•5X4'5 3·2 2 17·5X5·0 3·5 I
14'5X6•o 2·4 I 2o·ox 5·5 3·6 I
15•0X4•5 3'3 I
Range: 12-20X4-6µ., E = 2·2-4·0, for 70 spores

Average ·of 5 spores 12-12·5 µ. long, E = 2·5
20 13-13·5 µ. " E= 2·7
14-14·5 I' ,, E= 2·9
15-15·5 JL " E = 2·9
II
13
16 16-16·5 JL " E = '3·2
4 17-17·5 µ. " E= 3'4
,, 1 ,, 20µ. long E= 3·6
Average of 70 spores 14·0x 5·0 µ., E= 2·8
1
E = length ...;- width.
RAMARIA 579
On the ground in frondose or coniferous woods: Europe.
Spores 11-18X 4-6·5 µ,, pale ochraceous, distinctly verruculose-rough, elongate
ellipsoid, 2-several guttulate.
Basidia and hyphae as in R. formosa: hymenium not thickening.
TABLE XLV. SPORE-SIZE IN RAMARIA FLAVA
(One fruit-body; Rekawinkl, Austria, Sept. I925: a different collection from
Table XLIV
Size in µ. E' Number of records Size in µ. E' 1
Number of records
I
u·oX4·5 2·4 I 15·5X5·3 3·0 I
u·oX5·0 2'2 I 15·5x5·5 2·8 2
11·5X5·0 2·3 I 16·0X5·0 3·2 I
12·0X4·7 2·6 3 16·o·x6·o 2·7 2
12·0X5·0 2·4 4 16·0X6·5 2·5 I
12·9X5·3 2·3 2 16·5X5·5 3·0 3
12·5X4·7 2·7 2 16·5X6·c 2·8 2
12·5X5·0 2·5 3 17·0X5·0 3·4 2
13·0X5·0 2·6 3 17·0X6·0 2·8 I
13·0X5·3 2·5 2 17·0 X 6·5 2·6 I
13·5x4·7 2·9 I 17·5X5·0 3·5 2
13·5X 5·0 2·7 2 17·5X5'7 3·1 I
13·5X5·3 2·6 I 17·5X6·o 2·9 I
14·0X4·5 3·1 I 18·0X5·5 3·3 .1
14·0X4·7 2·9 I 18·0X6·o 3·0 I
14·0X5·0 2·8 I 18·0X6·5 2·8 2
14·0X5·3 2·6 4 18·5X6·o 3·1 I
14·ox5·5 2·5 I 19·0X5·0 3·8 I
14·5x5·0 2·9 2 19·0X5·3 3·6 I
14·5X5·3 2·7 2 19·0X6·o 3·2 I
15·0X5·0 3·0 2 19·5X5·5 3·5 I
15·0X5'3 2·8 3 20·0X 5·5 3·6
I 2
I
15·ox5·5 2·7 2 20·0X6·o 3·3 I
15·0X6·o 2·5 2
Range: 11-20X4·5-6·5 µ., E = 2·2-3·8, for 79 spores

Average of 3 spores u-11·5 µ. long, E = 2·3
14 12-12·5 µ. E = 2·4
9 13-13·5 µ. E = 2·6
12 14-14·5 µ. E = 2·8
12 15-15·5 /J, E = 2·8
9 16-16·5 µ. E = 2·8
8 17-17·5 µ. E = 3·i5
5 18-18·5 µ. E = 3·0
4 19-19·5 µ. E = 3·5
,, 3 ,, 20µ.long, E=3·5
Average of 79 spores 15·0X5·2·µ., E = 2·9
1 E = length -;- width.
This is the generally accepted European interpretation of the species, which has not
been recognized in any other continent. As explained by Donk (1933), ·Fries described
C. fiava originally as white-spored, but cited Schaeffer's illustration: later (1838, 1874)
he confused it with C. aurea (and, probably, C. paludosa). The white-spored fungus
of Fries (1821) has never been found since and is probably an error.
I have received from Austria, through Dr. Karl von Keissler, two collections which
agree in colour, form, and red staining (on bruising) with the illustrations of Konrad
580 RAMARIA
and Maublanc and of Bresadola, but they have considerably larger and more verrucose
spores. If these characters are taken to define R. fiava, it is easily distinguished from
R. aurea, R. fiavo-brunnescens, R. formosa, and R. Mairei which have often been
mistaken for it. The records of R. fiava from N. America and S. Australia belong to
R. fiavo-brunnescens, but R. sanguinea from both these continents is very close and
differs chiefly in the shorter spores: it may be identical.
Var. divaricata is said to be a form with divaricate ultimate branches, but such
growth-forms need careful and extensive study before they can be accepted as more
than fortuitous.
R. xanthosperma approaches R. fiava in the reddening of the flesh and long spores,
but its colour is generally paler and the spores are smooth.
R. flavo-alba spec. nov.

4-5 cm. high (specimens not full grown), solitary or gregarious, white tinged buff on
the stem and branches,pale yellowish towards the tips of the branches: stem 1-3 X 0·8-2 cm.,
bluntly obconic, white at the shortly rooting base, with many stout main branches:
branches crowded, numerous, cylindric, curved ascending, longitudinally rugulose
from the lateral branches, polychotomous becoming dichotomous and elongate above,
with subulate, bifid, or somewhat incurved tips: flesh white, unchanging, brittle.
On earth on a bare bank in the forest: Malaya (Singapore, Reservoir Jungle,
22.8.39).
I,
TEXT-FIG. 25+. Ramaria fiavo-alba, :< 2: the polychotomous

branch-tips, :< 5.
RAMARIA 581
Spores 8-10·5 X4·5-5µ, pale ochraceous (pale brownish ochraceous under the
microscope), minutely rough and uneven or subverruculose (without distinct warts
or striations), contents cloudy granular, aguttate and not vacuolate.
Basidia 50-70 X 8-9µ, clavate, projecting-20µ: sterigmata 4, rarely 2-3, 4-5µ long,
rather short.
Hymenium c. 70µ thick, ? not thickening (but specimens immature), covering the
stem and branches except the basal 5-10 mm. of the stem: subhymenium 20µ thick,
composed of rather loosely interwoven, uninflated hyphae 2-3·5µ wide.
Hyphae 4-10µ wide, with slightly but distinctly thickened walls (not 0·5 µ. thick),
clamped, cells 30-150µ long: on the surface of the stem narrow, with scattered pro
cesses -20 X2-3µ.
Structure as R. formosa.
This has the habit of R. formosa but its spores resemble those of R. polypus, from
which it differs in the form of the fruit-body and the thin-walled hyphae. R. verna,
R. fiavo-alba, and R. pamparum suggest the derivation of the Stricta-group from
Decolorans.
R. flavo-brunnescens (Atk.) comb. nov.

Basinym: Clavariafiavobrunnescens Atk., Ann. Myc. 7, 1909, 367: Burt, N. Am.
Clav. 1922, 22: Coker, Clav. U.S. Can. 1923, 120, t. 36, 37, 39, 85 (ut C. fiava):
Kauffm., Pap. Mich. Ac. Sci. Arts Lett. 1, 1923, 124: Wehm., Pap. Mich. Ac. Sci.
Arts Lett. 20, 1935, 261: Fawcett, Proc. Roy. Soc. Viet. 52, 1940, 153, t. 6, 7 (ut C.
fiava): Doty, Clav. Pac. N.W. 1944, 54 (ut C.fiava).
Synonyms:? Ramaria Michaelis Huber, Zeitschr. f. Pilzk. 15, 1931, uo: 18, 1934,
34: Mich. Schulz Fi.ihr. f. Pilzfr., t. 315.
? Clavaria obtecta Britzm., Hym. Si.idb. 8, 1885, 288, f. 19.
7-20 X 7-18 cm., clear pale cream to light creamy yellow, Naples Yellow, deep prim
rose, antimony- or chrome-yellow, or ochraceous-orange, paler at maturity, tips con
colorous but turning brown on withering or bruising: stem short, attenuate downwa·rd,
rooting, branching at ground level and with abortive branches round the periphery:
branches numerous, rather irregular, internodes short or rather elongate, rugulose:
flesh concolorous, not reddening, very brittle, with age brownish and watery: smell and
taste pleasant, of green pea-nuts (Coker): taste none, smell faintly sweetish (Wehm.):
taste none or slightly bitter or nutty when dry, smell pleasant, Brassicaceous (Doty):
taste and odour mild, slightly fungal (Fawcett).
On the ground in frondose and coniferous woods: Europe (little known), N. Amer.
(widely distributed), China (Fukien), S. Austral.
Spores 7·5-12(-15) X 3-4·5(-5·5)µ, pale yellow to light ochraceous, minutely
roughened, or nearly smooth (Coker).
var. aurea Coker
Clav. U.S. Can. 1923, 124, t. 38, 85 (ut C.fiava var. aurea): Wehm., Pap. Mich. Ac.
Sci. Arts Lett. 20, 1935, 261 (ut C.fiava var. aurea): Fawcett, Proc. Roy. Soc. Viet. 52,
1940, 154, t. 7.
Synonyms: ? Ramaria elegans Huber, Zeitschr. f. Pilzk. 18, 1934, 34.
? Clavaria paludosa Lundell, in Krok et Alrnq. Sv. FI. 5th ed., pt. 2, 1932, 285.
-12 cm. high, ochraceous-buff to orange-buff or mikado-orange, clearer rich yellow or
chrome-yellow upicard, tinged salmon-orange or apricot downward, the tips turning
reddish vinaceous (Wehm.): stern tapered downward, white at the base: main branches
stout, dividing into numerous, rather elongate and open branches, mostly polychotornous:
flesh concolorous, rather brittle, unchanging, rather translucent (Fawcett): taste none:
smell unpleasant, slightly rancid.
On the ground in woods, frondose and coniferous: U.S.A. (N. Car., N.Y.), Canada
(Kov. Scot.), S. Austral., Europe (? unrecogniz5d).
582 RAMARIA
Spores 6·5-9·5 X 3-4(-4·5)µ, buff-yellow, ellipsoid-oblong, faintly roughened or
nearly or quite smooth.
(var. subtilis Coker, vide R. divaricata.)
Coker identified this species with the European C. fiava on the evidence of the short
spores which he found in European exsiccati. 1 Whether these specimens were cor
rectly named seems to me doubtful. They cannot correspond with R. fiava as
generally accepted by European mycologists and they may represent R. Mairei, as
mentioned on page 604. The N. American, S. Australian, and Chinese fungi, which
have been referred to C. fiava, agree in having shorter, less rough spores than in the
European R. fiava, more richly coloured fruit-bodies, and unchanging flesh. Hence,
I consider it advisable to separate them under Atkinson's name, which Coker gives as
a synonym.
The species comes, in colouring, between R. aurea and R. formosa, but it has the
stature of R. formosa and the texture of R. aurea, and the spores are smaller than in
either. Thus, I think R. Michaelis must belong to it, for it has light egg-yellow fruit
bodies with the stature of R. formosa and watery-marbled flesh as in R. aurea; its
main distinction, according to Huber, is that the yellow colour is not destroyed in
cooking, as it is in R. aurea and R. formosa, and it ranks as a good esculent: no spore
details were given, but it confirms Coker's findings that R. fiavo-brunnescens occurs,
unrecognized, in Europe. The various descriptions of the taste and smell of R. fiavo
brunnescens show that, without a very marked peculiarity, such have little diagnostic
value.
Var. aurea has smaller, more richly coloured fruit-bodies with even smaller spores.
TABLE XLVI. SPORE-DATA FOR RAMARIA FLAVO-BRUNNESCENS

10-12(-15)X (3-)4-4·5(-s-5) Doty Oregon, &c. lightly rough
10-14X 3·5-4·5 Kauffmann Colorado
11X4·5 Coker New York nearly smooth
10-11x4·5 " " rough
9·5-11X 3·5 Wehmeyer Nova Scotia smooth or minutely rough
7-11X 3·2-4·3 Fawcett Victoria rough
7-10·5X3·5-4·5 Teng Fukien minutely rough
7·8-10·3X 3-4 Coker N. Carolina No. 560
8-10X3·5-4·2 " " No. 2850
7·5-10X 3-4 " " No. 2855: minutely rough
" "
I
7·5-9·8X 3-4 No. 562
8-<)"7 X3-3·8 " " No. 795, mostly smooth
7-9·.(xn-4 " " No. 2922: nearly smooth
8,5-9.:3X4-4·5 " New York Burnham 108
7·3-9X3-3·7 " N. Carolina No. 2870
8·8X 3·2 " " No. 2895
8·5X 3'7 " I
Connecticut nearly or quite smooth
var. aurea
6·6-9·3X3-3·7 Coker N. Carolina No. 2851 (type)
7-9X 3·3-3·8 New York No. 13
6·6-8·2X3·3-3·7 " No. 14
6·6-8X 3-3·6 " N. Carolina No. 2893
7-8·5X 3·5 Wehmeyer Nova Scotia
7-9·5 X 3·5-4·5(-4·8) Fawcett Victoria
1
9-11x3·7-5µ., Herb. Pers.: 7·8-10X3·8-4·5µ., Karsten (Finland): 9-12X3·7-5µ.,
Thlimen (Bohemia): 7·5-10 :< 3·7-4·2 µ., Juel (Uppsala).
RAMARIA
Thus it approaches R. formosa but still lacks its peculiar texture on drying. For this
reason, I think R. elegans Huber belongs to var. aurea: it is described as being as large
as R. fiava, with orange-pink branches and watery-marbled flesh, and as edible (not
bitter as R. formosa): the spores were not described. Compare, also, the rich yellow
form of R. obtusissima.
..ffi?(9,�i�
r1<'""t lrr1·," �r
\"
;·.
TEXT-FIG. 255. Ramaria formosa: stages in the development of the

fruit-body, X 1.
C. paludosa seems, also, to belong to var. aurea, as a pale state, possibly as a habitat
form of swamp-localities. Lundell placed it next to R. fiava, but (in litt.) suggests
that the Swedish R. fiava, which does not have reddening flesh, is what European
mycologists generally identify as R. aurea-that is, R. fiavo-brunnescens var. aurea in
my sense, because of the elongate branches. The following is the original description
of C. paludosa, for the translation of which I am indebted to Lundell and Nannfeldt:
Similar to Clavariafiava (of Sweden), but usually of a higher and narrower growth,
10-22 x 7-12 cm., whitish pale, finally dirty pale, the tips in young specimens tinged
pale rose: stem usually indistinct, sometimes tapering to the base: main branches
straight, upright, somewhat stiff and fastigiate: secondary branches terete, often
RAMARIA
becoming rugulose, the tips usually rounded (variously obtuse or sub-bifid): in coni
ferous woods among mosses, especially in swamp-margins: autumn, edible: (hitherto
noticed only in the neighbourhood of Upsala, where it is rather common in certain
years; probably overlooked elsewhere).
C. obtecta Britzm. may be the earliest name for R. fiavo-brunnescens, if the spores
are correctly given. Its description is:
Yellowish: stem thick: branches short, thick, with unequal, acutely toothed, tips
of varying length.
In woods: Germany (Bavaria).
Spores 6-8 X 3-4 µ,, ochraceous.
TEXT-FIG. 256. Ramaria formosa, X 500.
R. formosa (Fr.) Quel.

FI. Myc. 1888, 466.
Basinym: Clavaria formosa Fr., Syst. Myc. 1, 1821, 466.
Synonyms: Cldvaria formosula Britz., Hym. Siidb. 8, 1885, 287.
C. densa Pk., Rept. N.Y. St. Mus. 41, 1888, 79 (fide Coker).
Clavariella formosa (Fr.) Karst., Hattsv. 2, 1882, 185.
Text-Figs. 248, 255, 256.
7-30 x 6-15 cm., gregarious or caespitose, pinkish buff, orange-rose, or pinkish
ochraceous, with lemon-yellow tips: stem 3-6 X 2·5-6 cm., soon breaking up into many
branches, often branched almost from the base and appearing as a cluster of separate
fruit-bodies, whitish at the base: branches many, polychotomous below, ultimately
bifid, internodes short or elongate, very variable, often longitudinally rugulose, tips
blunt or subacute: flesh white or subconcolorous, often turning vinous-brown then
blackish where cut or bruised, fragile, not hygrophanous, drying chalky friable; slightly
bitter (very bitter after cooking): poisonous, causing diarrhoea (Huber, Heim).
In humus in woods, chiefly frondose, arising from white rhizomorphs: Europe,
temp. N. Amer. and Asia (China, Japan), S. Austral.
RAMARIA 585
TABLE XLVII. S PORES OF RAMARIA FORMOSA
Spore-size in µ, Authority Country Remarks

13·7X5·7 (av.) Corner England II-17(-21)x5--'7 µ, (Table LI)
(2 collections)
13·2X 5·0 (av.) " England 11-15·5X4-6 fL (Table L)
13·2X4·1 (av.) " " 10-17x3·5-5 fL (Table XLIX)
12·3X 5·8 (av.) " Austria 10-14·5x5-6·7 µ (Table XLVIII)
8-13X4-6 Bourd., Galz. France
9·5-13X4-4·7 Donk Holland
8-12X 3·5-4·5 Bresadola Italy
9-11X5-6 Rea England
9-1 l X5 Cott., Wakef. "
8-11X4-5 Coker Europe Persoon's specimens
(8-10X 4-5 Britz. Germany C. formosula)
8-12·5X5-6 Teng China
9-12·2X4-6 Coker U.S.A. 22 collections
9·3-12·2X 4-5·3 Dearness Canada
9·5-1 lX4·5-5·5 Doty U.S.A. Oregon
8-11X4-6 Coker " 5 collections
8-9X 3·5-4·5 " " Shear (Alcove, N.Y.)
7·5-8·2X 4·5-4·8 " " C. densa (Type)
7-7·8X4-5 " " No. 2688 (N. Car.): typical chalky
flesh when dried, typical colour.
7-9(-11)X 4-5·5 Cleland, Chee! Australia
(7-8·5X 4·2 " " wholly pale yellow fruit-bodies)
TABLE XLVIII. SPORE-SIZE IN RAMARIA FORMOSA

(One fruit-body; Austria, Tulnerbach, Sept. 1925)
Spore-size in fL I E' I Number of records Spore-size in fL I£' Number of records
10·0X 5·0 2·0 l 12·0X6·3 I� 3
10·0 i< 5·3 1·9 2 12·0X6·5 l ·8 2
1o·ox5·5 1·8 l 12·5X6·5 1·9 l
10·5x5·0 2·1 l 13·0X5·3 2·5 2
10·5X5'3 2·0 l 13·0X5·5 2·4 l
10·5X5·5 1·9 2 13·0X6·3 2·1 2
10·5 X 5·7 1·8 2 13·0X6·5 2·0 2
I1·0X5·3 2·1 2 13·5X5·0 2·7 2
l l'OX5·7 1·9 l 13·5X6·3 2·1 l
11·ox6·0 l·8 2 14·0X5·0 2·8 l
11·0X6·3 l ·7 l 14·0 >'. 6·o 2·3 3
l l ·5 X5·3 2·2 2 q·o,'. 6·3 2'2 2
11·5 >'. 5·5 2· I I 14·0 / 6·7 2·1 l
r1·5x6·5 l ·8 I I 14·5 / 5·0 2·9 l
12·0>(5·5 2·2 2 14·5 / 5·3 2·7 l
12·ox5·7 2·1 l 14·5 i< 6·o 2·4 l
12·0 X 6·o 2·0 I I
I
Range, 10-q·5 ,< 5-6·7 fL, E = 1·7-2·9, for 50 spores
Average of 10 spores, 10-10·5 fL long, E = 1·9
11-11·5µ E=2·0
12-12·5 ,, E = 2·0
13-13·5 fL E = 2·3
,, " 14-14·5 µ, E = 2·4
Average of 50 spores, 12·3X5·8µ, E = 2·1
' E = length -c- width.

586 RAMARIA
Spores c. 8-15 x4-6 µ, very variable in size, ochraceous, oblong ellipsoid, coarsely
or finely rough, 1-3 guttulate:
Basidia 40-60X 7-10 µ: sterigmata 4, 5-7 µ .
Hymenium 50-55µ thick, not thickening: subhymenium 20-30µ thick, composed
of narrow, uninflated hyphae 2-3·5µ wide.
TABLE XLIX. SPORE-SIZE IN RAMARIA FORMOSA
(One fruit-body; England, Burnham Beeches, Bucks., I9.9.25) 1
Spore-size in µ. E' Number of records Spore-size in µ. EZ Number of records
-- --------
10·0X3·7 2·7 2 13·5X4·3 3·2 l
10·5X3·5 3·0 I 13·5X4·7 2·9 2
10·5X3'7 2·8 2 14·0X4·0 3·5 3
10·5X4·0 2·6 2 14·0X4·3 3·3 l
10·5X4·3 2·4 2 14·0x4·5 3·1 I
10·5X4·7 2'2 l 14·5x4·0 3·6 4
I l'OX3·5 3·1 2 14·5X4·3 3·4 I
II·oX3·7 2·9 4 15·0X3·5 4·2 l
11·0X4·0 2·8 I 15·0X4·0 3·8 2
11·0X4·3 2·6 l 15·0X4·3 3·5 I
11·5X3·5 3·3 I 15·0X4·5 3·3 3
11·5X4·0 2·9 I 15·5X4·0 3·9 I
12·0X3·5 3'4 4 15·5X4·3 3·6 l
12·0X3·7 3·2 l 15·5X4·7 3'3 I
12·0X4·0 3·0 3 16·0X3·5 4·6 l
12•5X3·7 3·4 l 16·0X4·5 3·6 l
12·5X4·0 3·1 I 16·0X4·7 3·4 3
13•0X3·7 3'5 I 16·5X4·7 3·5 I
13·0X4·0 3·3 3 17·0X4·5 3·8 l
13·0X4·3 3·0 l 17·0X4·7 3·6 2
13·0X4•7 2·8 I 17·0x5·0 3·4 i
13·5X4·0 3'4 I
Range, 10-17X3·5-5µ., E = 2·2-4·6, for 70 spores.

Average of 10 spores, 10-10·5µ. long, E = 2·8
II-11·5µ. E=2·9
12-12·5µ. E = 3·2
13-13·5µ. E = 3·2
14-14·5µ. E = 3·5
15-15·5µ. E = 3·6
,. ,. 16-17µ. ,. E = 3·7
Average of 70 spores, 13·2x4·1 µ., E = 3·3
1
This collection of several fruit-bodies had unusually narrow, l�ss verrucose spores, but
macroscopically it was typical.
2
Hyphae 3-13µ wide, cells 50-300µ long, with scattered ampullaeform swellings
11-16 µ wide, clamped, with firm but scarcely thickened walls, longitudinal, flexuous,
compact, with a few narrow laticiferous hyphae, not pseudoparenchymatous: in the
rhizomorphs 1-2µ wide.
This well-known fungus is one of the few common large species about the identity
of which there is no doubt. It is, moreover, widespread and, though variable, is
generally easy to recognize. It has not been found in the tropics but it probably occurs
there in the mountains.
Typically it has a moderately large, pinkish yellow fruit-body with massive trunk and
lemon-yellow tips to the crowded, erect branches. Typically, also, for the European
RAMARIA
specimens, the flesh turns vinous-brown on bruising, but the colour-change has
seldom been noted and it is not recorded by Coker, Wehmeyer, or Kauffman for the
N. American specimens: Coker, indeed, is emphatic that it does not occur, but Doty
describes it as 'often staining dark violet or black where bruised'. The normal colour
varies from yellow tinged pink to pinkish orange and, in age, pallid cinnamon-brown.
In shape it varies into forms with practically no stem, being then branched almost
from the base, or the stem may be very stout and massive or rather slender, in which
case the fruit-bodies, when subcaespitose, resemble R. conjunctipes. The branches
TABLE L. SPORE-SIZE IN RAMARIA FORMOSA
(One fruit-body; England, Beaconsfield, Bucks., Oct. I933)

Spore-size in µ, E'
--
Number of records Spore-size in µ, E'
--
Number of records
11·0X4·5 2·4 2 13·0X6·0 2·2 I
11·0X5·0 2·2 I 13·5X4·7 2·9 I
11·5X4·5 2·6 I 13·5X5·0 2·7 2
11·5X4·7 2·4 I 13·5X5·7 2·4 I
I1·5X5·0 2·3 5 14·0X4·7 3·0 I
12·0X4·0 3·0 I 14·ox5·0 2·8 3
12·0X4·5 2·7 I 14·0X5·3 2·6 3
12·0X4·7 2·6 I 14·0X5·7 2·5 1'
12·0x5·0 2·4 I 14·5X5·0 2·9 I
12·5X4·5 2·8 I 14·5X5·3 2·7 I
12·5X4·7 2·7 2 15·0X5·0 3·0 4
12·5X5·0 2·5 3 15·0X5·3 2·8 3
13·0X4·7 2·8 I 15·0X5·7 2·6 2
13·0X5·0 2·6 3 15·5x5·0 3·1 I
13·0X 5·3 2·5 I
Range, 11-15·5X4-6µ,, E = 2·2-3·1, for 50 spores.

Average of 10 spores, 11-II·5 µ, long, E = 2·4
12-12·5µ, E = 2·6
13-13·5µ, E = 2·6
14-14·5 µ, E = 2·7
,, ,, 15-15·5µ, ,, E = 2·9
Average of 50 spores, 13·2X5·0µ,, E = 2·6
1 E= length -:- width.
may be elongate and parallel, or short, thick, and cauliflower-like (especially in young
massive 1:pecimens), or few and laxly divaricate. Thus, macroscopically, R. formosa
may resemble R. aurea, R. fiava, R. fiavo-brunnescens, R..divaricata, R. conjunctipes,
and many other species, and I have no doubt that it has very often been mistaken and
that the discrepancies in the descriptions, particularly of the spores, given by various
authors for these similar species have arisen from misidentified specimens of R.
formosa. But there are, so far as known, two constant characters to distinguish it.
Firstly, the pinkish stem or branches have always yellow tips when actively growing:
secondly, as Coker has shown, the flesh of dried specimens becomes characteristically
chalky-friable; thus the species is readily distinguished both in the field and in the
herbarium. Only two other species, R. suecica and R. subspinulosa, have this con
sistency when dried and neither has yellow tips when fresh. Thus it has been possible
to advance the study of this species far beyond that of any other and to understand in
some measure its variability. But it is most desirable that the hyphal reason for the
peculiar texture of the dried flesh should be discovered in order that it may be used
in verifying the determination of fresh specimens. Huber emphasizes that in the
living state the flesh of R.formosa is not watery-marbled, as it is in R. aurea,fiava, &c.,
588 RAMARIA
but it is also not watery-marbled in R. Mairei, which is not chalky friable when dried.
Huber also emphasizes the bitterness of the flesh which becomes very pronounced on
cooking, but Doty gives it as tasteless or subacid with age or on drying, and remarks
that R. brunnea differs in its taste from R. jor'ITl()sa. Such a widespread species has,
doubtless, many local irregularities.
TABLE LI. SPORE-SIZE IN RAMARIA FORMOSA
(One fruit-body; England, Gt. Missenden, Bucks., Oct. I925) 1
Spore-size in µ. E• Number of records Spore-size in µ. E• Number of records
u·ox5·0 2'2 2 14·0X5·5 2·5 2
u·ox5·3 2'1 2 14·0X5•7 2·5 l
u·ox5·5 2'0 2 14·0X6·5 2·2 l
u·ox6·o 1·8 l 14·5X5·0 2·9 l
u·5X5·0 2·3 l
' 14·5X 5'3 2·7 l
II'5X5'3 2·2 2 14•5X6·o 2·4 J
12·ox5·0 2·4 l 15·0X5•0 3·0 1
12·0X5'3 2·3 l 15·0X5·3 2·8 2
12·0x5·5 2·2 3 15•0X5·7 2·6 I
12·0X6·0 2·0 3 15·0X6·0 2·5 3
12·5x5·5 2·3 l 15•0X6·3 2·4 l
12·5x6·o 2·1 l 15·5x6·0 2·6 l
13·0X5·0 2·6 /"I 15·5X6·5 2·4 l
13·0X5•3 2·5 2 16·0X6·0 2·7 4
13·0x5·5 2·4 l 16·0X6·3 2·5 l
13·0X6·0 2·2 l 16·0X6·5 2·5 l
13·0X6·5 2·0 l 17•0X6·o 2·8 l
13·5x5·5 2·5 3 ' 17·0X 6·5 2·6 l
13·5X 5·7 2·4 l 17·0X6•7 2·5 l
14•0X 5·3 2·6 3 17•0X7·0 2·4 l
Range, u-17X 5-'7 µ., E = 1·8-3·0, for 60 spores.

Average of 10 spores, u-u·5 µ. long, E = 2·1
12-12·5 µ. E = 2·2
13-13·5 µ. E = 2·4
14-14·5 µ. E = 2·6
15-15·5 µ. E = 2·6
,, ,, 16-17 µ. ,, E = 2·6
Average of 60 spqres, 13·7X5·7 µ., E = 2·4
1
Almost identical spore-variation occurred in a specimen from Rekawinkl, Austria (Sept.
1925), and in one from Nettlebed, England (Oct. 1926). This last had occasional giant spores
21X7 µ., which indicate the limiting width.
• E = length -;- width.
The spore-data are given in Tables XLVII-LI. I have studied the spe�es in detail
because of the very large number of reliable spore-records which exist. The spores
range 7-21 X 3·5-7 µ,, and this variation has led me to a simple law governing spore
and hyphal-growth (New Phyt. 46, 1947, 195-228).
R. fragillima (Sacc. et Syd.) comb. nov.

Basinym: Clavariafragillima Sacc. et Syd., Syll. Fung. 16, 1902, 206.
Synonyms: Clavaria echinospora P. Henn., Monsun. 1, 1900, 43, 141: (non B. et
Br., 1875 = Scytinopogon echinosporus).
Clavariellafragillima (Sacc. et Syd.) v. Ov., Bull. Jard. Bot. Buit. Ser. 3, S, 1925,
275: le. Fung. Mai. 1, 1923.
RAMARIA 589
Text-Figs. 2A and c, 250 a, 257, 258: Plate 12.
3-14 cm. high, solitary, brownish ochre to drab cinnamon-fawn, becoming darker
reddish brown or ochraceous-umber, light ochre towards the orange-ochre or rufescent
orange tips, dingy vinaceous on bruising: stem 1-4 cm. X 2·5-zo mm., often with a
rooting base -3·5 cm. long, flesh-colour to ochraceous-brown, villous with white
fttf
TEXT-FIG. 257. Ramaria fragillima: fruit-bodies, X 1: spores and basidia,

X 1,000.
mycelium at the base, sometimes divided from the base: branches rather stout and
polychotomous with short internodes below, becoming dichotomous with more
elongate internodes 1·5-2·5 mm. wide, numerous, crowded, cylindric, erect, parallel,
generally with regular, alternating branching, sometimes rather lax and irregular, tips
blunt or subacute: flesh white, all parts slowly turning dingy vinaceous when cut or
bruised, soft, brittle, sappy: smell strong, of Ganoderma (paracresol) like R. Zippelii:
mycelium white, purple when bruised.
590 RAMARIA
On the ground in the forest, or- under trees in gardens: trop. Asia (Java, Ceylon,
Malaya).
Spores 9-16 x 4-7·5 µ., ferruginous, echinulate with acute or subacute hyaline spines
1-2µ. long, elongate pip-shaped or clavate and slightly curved, variable in size and
shape, I-several guttulate.
Basidia 45-70 X 9-12µ., with yellowish brown, granular-vitreous, dense contents:
sterigmata 2-3-4, mostly 4, rarely 1, (3-)6-8 µ. long, often of very unequal length or
with 1-3 abortive and sterile.
Sterile basidia as in R. cyanocephala.
Hymenium 60-70µ. thick at first, gradually thickening to 250µ..
Sterile hymenium on the uppersides of the branches, as in R. cyanocephala.
Hyphae 4-12µ. wide, clamped, walls thin or very slightly thickened, exactly as in
R. Zippelii but wider.
Ochraceous-brown colour of the fruit-body caused by the
yellowish ochraceous guttulae of the subhymenial hyphae and
outer hyphae of the flesh, and by the brownish ochraceous
cytoplasm of the basidia: orange-ochre colour of the br;mch
tips caused by the reddish brown granular-guttulate contents
of the hyphal tips and young hymenial hyphae.
This is C. fragillima as understood by van Overeem.
Whether it is really C. echinospora P. Henn., the name of
which Saccardo changed to C. fragillima, I am doubtful
because Hennings described it as having very fragile,
uniformly yellow-brown fruit-bodies with somewhat curved
branchlets and small spores 7-8 X 4-5 µ.. I have found just
such a fungus in Malaya, which appears indistinguishable
from R. longicaulis (see p. 600), but it is impossible to know
whether Henning's description was accurate. I have therefore
followed van Overeem's identification of this fairly common
TEXT-FIG. 258. Ramaria species of tropical Asia. Petch has recognized it in Ceylon
fragillima, x 1. and I agree with his remark that the colour is generally
brighter than in van Overeem's painting.
In this serise, R. Jragillima is very near the north temperate, 2-spored R. nigrescens.
R. fragillima has slightly smaller spores (to be expected in tetraspores) and larger, not
blackening, fruit-bodies. R. nigrescens var. americana bridges the difference in the
spores, which are also tetraspores, but its flesh is unchanging. I hesitate to identify the
tropical with the temperate species, however, until it has been studied in greater
microscopical detail. R. fragillima is particularly interesting in showing not merely
the great variation in spore-size and shape in species of Ramaria but in showing a
transitional state between the 2-spored and the 4-spored, even tending to the 1-2-spored
state of R. Zippelii, and thus clearly showing how the 2-spored state is derived from the
4-spored.
The following are the published spore-measurements of R. Jragillima:
Spore-size in µ. I
(spore-body) E' Authority Remarks
II-15·5 X 5-7·5 2·1 Corner Malya: Table LIII
10-16 X 4·7-6·3 2·5 " " Table LII
10-16 X 5-6 2·4 " "
9-11 X 5-6 1·8 van Overeem Java
9-12x4-5 2·3 Petch Ceylon
1
A fruit-body, which I studied in 1929, grew in height at the average rate of 3 mm.
RAMARIA 591
per diem from 25 to 48 mm., when it withered during a spell of dry weather.
The total life of the fruit-body must be a month, or more.
TABLE LII. SPORE-SIZE IN RAMARIA FRAGILLIMA

(One fruit-body; Singapore, I930)
I Number of records
I_£_
I
Spore-size in µ. Spore-size in µ.
(spore-body)
-·---
(spore-body) E' Number of records
10·0 x4·7 2·1 13·5X5·0 2·7
10·0X5·0 2·0 5 13·5X5·3 2·6 I
10·5X4·7 2·2 l 13·5 x 5·5 2·5 2
10·5 x 5·0 2'1 2 14·0X5·0 2·8
10·5X5·3 2·0 14·0X5·3 2·6
11·0X4·7 2·3 2 14·0X5·5 2·5 3
11·0X5·0 2·2 3 14·0X5'7 2·4
11·5X5·0 2·3 4 14·0X6·o 2·3
II·5X5·3 2·2 14·5X5·3 2·7
12·0X4·7 2·6 14·5X6·o 2·4 2
12·0X5·0 2·4 3 15·0X5·0 3·0 3
12·0X5·3 2·3 2 15·0X5·3 2·8 l
12·0 x 5·5 2·2 l 15·0Y6·0 2·5 2
12·5X5·0 2·5 2 15·5X4·7 3'3
12·5 x5·5 2·3 15·5X6·0 2·6
13·0X5·0 2·6 5 15·5X6·3 2·5
13·0 )( 5·3 2·5 16·0 x 5·5 2·9
Range, 10-16X4·7-6·3 µ., E = 2·0-3·3, for 60 spores.

A\'erage of 10 spores, 10-10·5 µ. long, E =2·3
11-u·5µ. E'=2·3
12-12·5 µ. E = 2·4
13-13·5 µ. E =2·6
14-14·5 µ. E = 2·5
,, 15-16 µ. E =2·8
Average of 60 spores, 12·6X5·2 µ., E =2·5
1
E =length -::- width.
R. fumigata (Pk.) comb. nov.

Basinym: Clavariafumigata Pk., Rept. N.Y. St. Mus. 31, 1879, 38: Coker, Clav.
U.S. Can. 1923, 135, t. 46, 47, 48 (ut C. fennica Karst.): Fawcett, Proc. Roy. Soc.
Viet. 51, 1939, 273 (ut C. fennica Karst.).
Synonyms: Ramaria versatilis Quel., Ass. Fr. 22, 1893, 489.
Clm:aria versatilis (Quel.) Bourd. et Galz., Bull. Soc. Myc. Fr. 26, 1910, 214: Sacc.
et Trott., Sy!!. Fung. 21, 1912, 426� Pearson, Tr. Brit. Myc. Soc. 24, 1946, 209.
Clavariella versatilis (Quel.) Maire, Bull. Soc. Myc. Fr. 30, 1914, 218, t. 9.
? Clavariella cedretorum Maire, Bull. Soc. Myc. Fr. 1914, 217, t. 9.
? Clavaria cedretorum (Maire) Sacc., Sy!!. 23, 1925, 488.
5-12 cm. high, uholly lilac or violaceous, except the uhite base, the branches becoming
fuliginous-ochraceous, Juliginous-cinnamon, or bistre, the tips violaceousfor a long time:
stem 2·5-5 cm. wide, distinct, often massive or sub-bulbous, pale or clear lilac or
violaceous, the base white and attached by white mycelial strands: branches crowded,
often compressed, polychotomous below, eventually with dichotomous blunt tips,
main branches few, 2-3 mm. wide: flesh white (? rufescent on bruising), rather tough,
not fragile, not hygrophanous (Maire): taste slightly bitter, or slightly acrid and then
mild (Coker), indistinct and mild (Fawcett).
592 RAMARIA
On the ground in frondose woods: Europe (France, England, Spain, Sweden,
uncommon), U.S.A. (N.Y., Mass., Conn., N. Car., frequent), S. Austral. (Victoria).
Spores 8·5-12·5 X 3·7-5·5(-6)µ, ochraceous to cinnamon-buff, finely rough, oblong
subfusiform, 1-guttate.
TABLE LIII. SPORE-SIZE IN RAMARIA FRAGILLIMA
(One fruit-body; Singapore, I94I)
-
I I
Spore-size in µ Spore-size in µ
(spore-body) E' Number of records (spore-body) E' Number ofrecords
1I·oX6·o 1·8 3 13·5X6·5 2·1 2
1I·ox6·5 1·7 l 14·0X6·o 2·3 l
11·0X6·7 1·6 l 14·ox6·5 2·2 l
1I·s.xs·s 2·1 l 14·ox6·7 2·1 l
II"S X6·o 1·9 I 14·ox7·0 2·0 l
11·5x6·3 1·9 I 14·0X7·3 1·9 I
11·5X6·5 l ·8 1 14·0X7·5 l '9 2
1I·5x6·7 1·7 l 14·5xs·o 2·9 l
12·0X5·7 2·1 1 14·5X6·5 2·2 1
12·0X6·o 2·0 3 14·5X7·0 2·1 1
12·0X6·5 1·8 1 15·0X6·o 2·5 l
12·0X6·7 1·7 3 15·0 X 6·5 2·3 2
12·0X7·0 1·7 l 15·0X6·7 2·2 l
12·5X6·o 2·1 l 15·ox7·0 2·1 2
13·ox6·0 2·2 4 I'5·0X7'5 2·0 l
13·ox7·0 1·9 2 15·5X6·0 2·6 I
13·5X6·o 2·3 15·5X6·5 2·4
I
I I
13·5X6·3 2·1 1 15·5x7·0 2·2 l
Range, 11-15·5X5-7·5 µ, E = 1·6-2·9, for 50 spores.

Average of 10 spores, 1I-1I·sµ long, E = 1·8
12-12·5 µ E = 1·9
13-13·5µ E = 2·1
14-14·5µ E = 2·2
,, ,, 15-15·5µ, ,, E=2·3
Average of 50 spores, 13·2x6·5µ, E = 2·1

Basidia 30-50 X 6-8·5µ: sterigmata 2-4.
Hymenium 90-100µ thick (? including subhymenium): hyphae 3-8 µ wide (no
clamps, Coker), 4-15µ wide (Bourd.).
Coker considered this species the same as R.fennica, the colour of which is different
enough to distinguish it; but I agree with him that R. versatilis is identical with Peck's
species. In shape, the fruit-bodies resemble some states of R. for111osa, but the stem is
often rather bulbous and divides into a few stout main branches so that it appears as
if several fruit-bodies arise from a swollen base. In one collection Coker observed
that the flesh turned pink then slowly rose-red on bruising (No. 92 r, N. Car.), but
whether this is normal or abnormal is not known.
Bourdot's collection of C. fennica, from Sweden, ,vith the branches violet then grey
tinted yellow-bistre must also be referred here (Bull. Soc. Myc. Fr. 48, 1932, 207).
The S. Australian fungi referred to C. fennica sensu Coker by Fawcett are clearly
R. fumigata. Fawcett notes· that the old branches become fuliginous-green, thus
suggesting that there may be transitions to R. fennica, but' the ·characteristic yellow
tips of this species were not seen.
Regarding C. cedretorum, I can see no difference from R. fwnigata unlt>ss the
branches of C. cedretorum remain amethyst-lilac \\·hile those of R. funiigata discolour
RAMARIA 593
greyish or brownish, and the basidia of G. cedretorum are longer (70-80µ). The dis
coloration of R.fumigata may be only apparent and caused by the deposit of spores at
maturity, and the length of the basidia may include the subhymenial cells. The
description of C. cedretorum' shows, however, that there is a species with wholly
lilaceous or violaceous branches, without yellow tips, and therefore distinct from
R.fennica.
TABLE LIV. SPORE-DATA FOR RAMARIA FUMIGATA
Spore-size in µ, Authority i Country Remarks

7·7-9·3X3·7-4·4 Coker U.S.A. C. fumigata-type
8·3-9·2 X3·7 " " No. 779: cinnamon-buff
8·3-II"IX3·7-5·5 " " No. 806
9·3-IIX3·7-4·4 " " No. 21
9·8-II X3·7-4·4 " " No. 620: cinnamon-buff
IO-IIX 4-4·8 " " No. 921
7·6-11·9X4·2-5·5 " " No. 1282
8-10, ovoid- Quelet France as C. versatilis
elongate
9-10·5X3-4 Maire " "
9-12·5X4-5 Bourd., Galz. " "
9-11 X4·5-6 Bourdot Sweden as C. fennica
8-9X4·5-5 Pearson England minutely aculeate
9·8-1 IX3·7-4·4 Fawcett Victoria as C. Jennica
9-12 X 4·5-5 Maire Algiers as C. cedretorum
6-12X4-6 Werner Morocco as C. cedretorum var. marocana
R. gelatinosa (Coker) comb. nov.

Basinym: Clavaria gelatinosa Coker, Clav. U.S. Can. 1923, 137, t. 48-50, 86:
Doty, Clav. Pac. N.W. 1944, 49, t. 10.
7-13 X 5-14 cm., massive, caespitose, much branched, arising from a massive, short,
cre�m-white base (-6X 5 cm.), creamy white becoming flesh-colour, buff-pink, or
brownish cinnamon, darker flesh brown with age, the tips paler, drying pale ochraceous
or pale orange: branches very numerous, crowded, condensed or rather elongate, much
branched with pointed tips, with many abortive branches near the base, divided
eight to eleven times: flesh concolorous, gelatinnus, transparent, toughish-elastic, soft:
taste slightly acid, then bitterish: smell slight, of tobacco.
On the ground in woods: U.S.A. (N. Car., uncommon: Oregon, under mountain
hemlock and white fir).
Spores 7·5-1 ox 4·5-6 µ, ochraceous-buff, distinctly verrucose (not echinulate),
ellipsoid.
Basidia 8-1r µ wide: sterigmata 4, 7 µ long.
Hyphae 2-4 ·5 µ wide, rarely inflated in places, -r oµ wide at the septa or clamps,
with very gelatinous walls, widely spaced in the toughish gelatinous flesh, clamped.
Coker and Doty say this remarkable species is allied with R. formosa. At first sight
it suggests a new genus, according to the old principles of 'texture', for except the thin
surface-layer the flesh is wholly transparent and firmly gelatinous 'like a table-jelly'.
1
C. cedretorum: 8-10 X5-15 cm., solitary or gregarious: stem thick, short, solid, fleshy
compact, glabrous, white, soon divided: branches solid, subcylindric, sulcate-rugulose, short,
subdivaricate, densely crowded, fragile, beautifully amethyst-lilac, tips 2-5 tubercled and
concolorous: flesh white: somewhat bitter, smell slight: spores 9-1 z X4·5-5 µ,, ochraceous,
verrucose, ellipsoid-oblong, 1-several guttulate: basidia 70-80 X7-8 µ,, 2-4-spored: in woods
of Cedrus atlanticus, Algiers:
var. marocana: 3-5 cm. high: basidia 40-5ox8 µ,: spores 6-12X4-6µ,, not verrucose:
under Cedrus atlanticus, Morocco (Atlas Mts.): (Werner, Bull. Soc. Myc. Fr. 1934, 140).
SIIQ
Qq
594 RAMARIA
Coker says there is a slight tendency to gelatinous flesh in R. botrytoides. But the gap
between R. formosa and R. gelatinosa is bridged almost exactly by R. subgelatinosa
which I have found in Malaya. R. gelatinosa thus appears as a more gelatinous and
less pigmented, small-spored derivative of the R. formosa-group, R. subgelatinosa
being a more pigmented and less gelatinous, but equally small-spored derivative.
Neither has the vinaceous colour-change of R. Jormosa.
I have verified the presence of clamps in Coker's exsiccati in the Kew herbarium.
Compare, therefore, the �ubgelatinous European R. ignicolor, which has no clamps
but is otherwise very similar.
The following are the spore-data for R. gelatinosa:
7·5-9X4·5-5µ, mostly 8x4·6µ, verrucose, Coker No. 2413 (type).
8-9·8 X4·5-4·8µ, rough, Coker No. 2861.
7·7-9·2X4·5-4·8µ, Coker No. 2880.
8-9(-10·2)X 4·5-6 µ, 'reticulately ? roughened', Doty.
R. gracilis (Fr.) Quel.

FI. Myc. 1888, 463.
Basinym: Clavariagracilis Fr., Syst. Myc. 1, 1821, 475: Bres., Ann. Myc. 1, 1903,
161: Coker, Clav. U.S. Can. 1923, 169, t. 64, 88: Wehm., Pap. Mich. Ac. Sci. Arts
Lett. 20, 1935, 260: Lund., Sv. Svamp. 3-4, 1936, 19: Fawcett, Proc. Roy. Soc. Viet.
51, 1939, 17, t. 5.
Synonyms: Clavaria alutacea Lasch in Rabenh. Klotzs. herb. viv. myc., cent. 16,
1851, No. 1519: (fide Coker).
Clavariella gracilis (Fr.) Karst., Hattsv. 2, 1882, 188.
Clavaria fragrans E. & E., N. Am. Fungi, 2nd ser., No. 2023, 1888: Cooke, Grev.
17, 1889, 59: (fide Coker).
Clavariafragrantissima Atk., Ann. Myc. o, 1908, 57: (fide Coker).
Clavariafiavuloides Burt, Ann. Mo. Bot. Gdn. 9, 1922, 28, t. 5, f. 34: Kauffm., Pap.
Mich. Ac. Sci. Arts Lett. 5, 1925, 118: (fide Coker).
Clavaria byssiseda sensu Pat., Tab. An. n. 567.
Plate 13.
3-8X 1-4 cm., slender, sparingly or densely branched, mostly caespitose, white or
clay-colour, then ta11, alutaceous or pale buff, often tinged pinkish or flesh-colour, finally
pale cinnamon or pale ochraceous with age: stem -1·5 cm.X2-4 mm., occasionally
-2·5 cm. long, often decumbent, white or paler concolorous, with white,fioccose mycelial
strands or mycelial sheets at the base: branches generally numerous, slender, 1-2 mm.
wide below, 0·5-1 mm. above, erect or subflexuous and rather spreading, axils round,
sometimes irregular, tips slender, tapered, subacute or subcristate, white: flesh fibrous,
pliable, sappy and hygrophanous, not brittle: taste none or slightly bitter: smell slight
or distinct, of aniseed.
On the ground in coniferous woods, or under conifers: Europe (rare or uncommon),
U.S.A. (frequent in the north and south), Canada, S. Austral. (under Pinus radiata
and P. insignis; probably introduced).
Spores (4-)5-7 x 3-4µ, light yellow ochraceous to naples yellow (pale brown,
Wehmeyer), verruculose- to minutely rough (or nearly smooth, Coker, Fawcett),
oblong or ovoid, characteristically 1-2-guttate.
Basidia 25-45X4·5-7µ, 4-spored: 35-42X7µ, multiguttulate, with 4 sterigmata
6-7 µ long (Corner).
Hymenium thickening -120µ, absent from the axils and uppersides of the ma...
branches: subhymenial hyphae 2-3·5µ wide, interwoven.
Hyphae 3-9 µ wide, -15 µ in places, especially in clavate inflations near the septa,
clamped, thin-walled, long-celled, many of the hyphae in the lower part of the fruit-body
with thickened walls (0·5-1·5µ), with rather numerous abortive clavate branches.
RAMARIA 59 5
Colour in the hymenium and subhymenium, caused by the pale yellowish sap and
guttulae of the basidia.
Because of the thick-walled hyphae and indistinctly verrucose spores, I have classed
this species in the Stricta-group; where its terrestrial habit seems anomalous, though
the group has doubtless been derived from a terrestrial ancestor. The species is
generally classed with R. fiaccida (see p. 576) and R. ochraceo-virens, but it lacks their
deep ochraceous and brownish colours, and thus it resembles the coniferous-dwelling
R. condensata, which has the yellow colour evanescent at its tips. Indeed, R. gracilis,
like R. pol;ypus, suggests another connexion between the Stricta-group and Decolorans.
The characteristic smell suggests affinity with R. conjunctipes var. odora and R.
cystidiophora. Compare also R. capucina.
Very often the branching is more or less flattened, dichotomous or multifid, and
I have little doubt that R. palmata is only an extreme form of this kind.
Fawcett describes the species from S. Australia, where it has probably been intro
duced with Pious (? in soil or by air-blown spores). Its fruit-bodies reach 9 cm. high,
6 cm. wide, and have broader spores (-5µ wide).
TABLE LV. SPORES OF RAMARIA GRACILIS

5-6X3-3·6 Coker France Herb. Persoon: minutely rough
5-'r5x3-4 Bourd., Galz. " finely rough
5-6X3·5 Corner England " ,,
5-'7X3-4 Donk Holland slightly rough
5-6x3 Karsten Finland
5·5-6·5X3·5-4 Lundell Sweden verruculose
4•2-6·5X3-3·6 Coker " leg. Juel: barely rough
5·5-6·5X3-4 Bresadola Poland punctate rough
5·5-6·5X3-4 Coker Europe Herb. Bres.: leg. Eichler
5-6·5 X3•4-3·8 C. alutacea (Herb. Kew: Rab. Exs.
1519)
5·5x3·5 Cleland S. Australia
4·8-6·2X3·3-4·9 Fawcett " slightly rough to nearly smooth
6-7 long Coker N. Carolina Beardslee
5·2-7X3-3·3 Connecticut Underwood
4·8-7X3-3·5 N. Carolina Murrill, House; nearly smooth
4·8-7X2·5-4 New York Burnham 80
5·5-6·5X 3-3·7 · C. Jragrans (2 collections, Herb. New
York)
5-6·5X 3-3·5 N. Carolina 7069 : verruculose
4·5-6·5X 3-3·7 Virginia Holmes: minutely rough
4·3-6·5X2·8-3•4 New York Coker 18
4·8-6·3X 3 Ellis : smooth
4·8-6X 3-3·3 Coker, Burnham 107; rough
5-6X3-3·5 Coker 6
5·2X3 " Burnham 31 : nearly smooth
5·2x3 Michigan E. et E.: smooth
5x3 " New York Burnham 30: smooth
5-6 >< 3-3·5 Wehmeyer Nova Scotia pale brown spores
5·2-6X3-3·2 Doty Oregon roughened, yellow
R. grandis (Pk.) comb. nov.

Basinym: Clavariagrandis Pk., Bull. Torr. Bot. CL 29, 1902, 75: Coker, Clav. U.S.
Can. 1923, 192, t. 75, 76, 90.
Synonym: Clavaria spiculospora Atk., Ann. Myc. 7, 1909, 368: ( = C. spiculisperma
Atk., Sacc. Syll. 21, 1912).
6-18 X 2-12 cm., gregarious, uniformly deep brick-brown (antique-brown of Ridgway),
596 RAMARIA
tips abruptly white: stem 1-6 x 1-2 cm., stout, distinct, deeply rooted, white
tomentose below, quickly turning dull brownish lavender or purple on bruising:
branches numerous, upright, crowded, 2-4 times branched, rather stout, ending in stout,
abruptly rounded cusps: flesh pure white, immediately turning lavender-brown when cut
or bruised, firm but tender: taste mildly acid-bitterish, astringent after a few minutes.
In humus in frondose woods: U.S.A. (N. Car., Maryland): Japan.
Spores 11-13·5 X6·3-7 µ, antique brown (fuscous ferruginous), echinulate with
spines 1·5µ long. (Coker 953, u-13X6·3-7µ: Coker u86, 12X6·3µ: Schallert 38,
13·5X7 µ).
Basidia IOµ wide: sterigmata 2, 7µ long.
Hymenium c. 70µ thick, stratose (-4 layers) and thickening.
Hyphae 4-7·5µ wide, closely packed.
As mentioned on page 635, this will probably be found identical with R. Zippelii of
tropical Asia.
R. guadelupensis (Lev.) comb. nov.
Basinym: Merisma guadelupense Lev., Ann. Sci. Nat. Bot. Ser. 3, 5, 1846, 157.
Synonyms: Pterula guadelupensis (Lev.) Sacc., Syll. Fung. 6, 1888, 742.
Lachnocladium guadelupense (Lev.) Pat., Journ. de Bot. 3, 1889, 3, t. 1, f. 7: Burt,
Ann. Mo. Bot. Gdn. 6, 1919, 277.
Dendrocladium guadelupense (Lev.) Lloyd, Myc. Writ. 5, 1919, index 13.
With very short stem, hardly I cm. long, coriaceous, branched: branches very thin,
elongate, fastigiate, compressed, dichotomous, becoming fuscous: apices short, naked,
acute.
Guadeloupe.
Spores 12X6µ, brown, warted, apiculate at the base (Burt).
The description refers to a dried specimen, which has shrivelled so as to appear like
a Pterula. It is obviously a Ramaria, according to Burt's spore-data, related, perhaps,
to R. Zippelii or R. cyanocephala; but are the spores verrncose or echinulate?
R. guyanensis (Pat.) comb. nov.
Basinym: Lachnocladium guyanense Pat., Journ. de Bot. 3, 1889, 35, t. 1, f. 6.
8-IO cm. high, densely caespitose: stems 2-3 cm. x 1-2 mm., many, slender,
flexuous: branches slender, dichotomous, acute: hymenium unilateral, pulverulent,
on stems and branches.
On the ground in the forest: French Guiana.
Spores 6-8X4-5 µ, ochraceous-brown, echinulate, ovoid-mucronate.
The colour of the fruit-body is not given, but the species must be near R. longicaulis
and R. Zippelii var. gracilis. Compare also R. leucoceras, which appears to be a more
massive form.
R. Henriquesii (Bres. et Roum.) comb. nov.
Basinym: Clavaria Henriquesii Bres. et Roum., Rev. Myc. 1890, 36, t. 92, f. 5.
5-6X4-5 cm.: stem 2-3 x I cm., pale yellowish: branches short, subdichotomous,
terete or compressed, subrugulose, egg-yellow, glabrous: flesh white, without smell or
taste.
On the ground, S. Thome Isl., Guinea.
Spores 9-11 x 9 µ, yellowish, smooth, globose-ellipsoid.
The spores and compressed branches suggest Clavulina, in which a yellow colour
is unusual, though present in C. Leveillei. Compare R. kisantuensis.
R. holorubella (Atk.) comb. nov.
Basinym: Clavaria holorubella Atk., Ann. Myc. 6, 1908, 57.
Pseudonym: Clavaria rufescens sensu Coker, Clav. U.S. Can. 1923, 139, t. 51, 86:
Journ. El. Mitch. Sci. Soc. 1947: Fawcett, Proc. Roy. Soc. Viet. 51, 1939, 274, t. 20, 22.
RAMARIA 597
Clavaria botrytis sensu Kauffm., Pap. Mich. Ac. Sci. Arts Lett. 1, 1923, 123:
ibid., 8, 1928, 515 (haud sensu Coker = R. botrytoides).
C. australiana Clel., Trans. Roy. Soc. S. Austral. 55, 1931, 159: Toadst. S. Austral.
2, 1935, 266, f. 59: (fide Fawcett, I.e.).
8-18 X 8-12 cm. medium to large, light tan, the tips concolorous then soon pale rosy
vinaceous to darker vinaceous, finally deep dull brick-brown, sometimes wholly reddish
to madder-brown (Atkinson): stem short, stout, usually distinct, whitish then tan or
alutaceous, often stained deep dull vinaceous-rose colour (Coker), deeper. red than the
branches (Atkinson): branches delicate, very numerous, loosely crowded, spreading
on the sides, sometimes longitudinally rugulose, polychotomous, then dichotomous:
flesh white, or often stained vinaceous-rose below the surface-stains, reddish (Atkin
son), moderately brittle becoming pliable: taste rather bitter or subacid, or practically
none: smell none or very slight (Coker), like watercress (Atkinson).
On the ground in woods: U.S.A. (N. Car., Col., Mich., Ohio): S. Austral.
Spores (9-)u-13 X 3-5,u (Coker), 10-15(-16) X 4-6 ,u (Kauffm.), cinnamon-buff,
finely longitudinally striate, otherwise smooth, cylindric-ellipsoid, with curved apiculus:
(striations faint and often slow to appear, Kauffman).
Basidia 7·5-8 ,u wide, 4-spored: hymenium 55-65,u thick: hyphae 3-8 ,u wide,
clamped.
This is, in the main, Clavaria botrytis of N. America. Coker identified it with C.
rufescens of Europe, which is doubtfully identifiable and, as I have explained on page 619,
should be kept apart. Therefore I have used Atkinson's name on the authority of
Coker, who gives it as a synonym and says 'it is not to be distinguished from this
species by the description, and the spores are identical'.
R. holorubella is so close to R. botrytis that it may well be considered only a variety
with shorter, narrower spores and, perhaps, reddening flesh, for Coker emphasizes the
vinaceous staining of the stem.
The Australian fungi referred to C. rufescens sensu Coker by Fawcett seem also to
belong here. Their description, briefly, is:
10-12(-20) x 7(-15) cm., cinnamon-buff with vinaceous tips when young, the
purplish colour more pronounced at maturity, and extending along the upper parts of
the branches (Hydrangea red and dark vinaceous), brownish with age: stem white,
occasionally with picric-yellow stains, very massive: flesh white, soft, brittle in young
plants: taste not distinct, mild.
Spores 12-15 · 5 x 3-4· 5 µ,, ochraceous, roughened, longitudinally or obliquely striate.
S. Austral.: moderately common in Victoria.
The spore data for R. holorubella are given in Table XXXVI.
R. holsatica (P. Henn.) comb. nov.

Basinym: Clavariella holsatica P. Henn., Krypt. FI. Schles. 3, 1888, 240.
Synonym: Clavaria Holsatica (Henn.) Sacc., Syll. 11, 1895, 134.
Caespitose, rather tough, very much branched, pallid ochraceous, often white pruinose:
branches short, often flexuous, divaricate, crowded: branchlets concolorous, com
pressed, generally acute, dentate or laciniate, rarely thickened and obtuse.
In a garden: Germany (Bruhn, Schleswig Holstein).
Spores 1 o X 6-7,u or 7-8 X 6-6· 5 ,u, clear olivaceous, smooth, subglobose or ovoid.
The branching and subglobose spores suggest Clavulina.
R. ignicolor Bres. sp. nov.
4 cm. high, as a small flame-coloured R. formosa: stem stout, short, soon dividing
into a few main branches: flesh evidently subgelatinous.
On the ground in coniferous woods: Italy (Mendola, Aug. 1903, leg. Bres.).
Spores 7-8·5 X4-5µ,, pale yellow under the microscope, minutely rough or sub
verruculose, pip-shaped, rather broad, I-several guttulate.
RAMARIA
Hyphae 2-5 µ, wide, not inflated, without clamps, the walls firmly gelatinous.
I cannot find that this remarkable species was published by Bresadola. These notes
are taken from the light orange-ochraceous dried specimens in the Kew herbarium,
named Clavaria ignicolor Bres. It is the only species of Ramaria without clamps
(except, perhaps, R. capitata and R. luteo-fusca), yet its affinity seems clearly to be
with R. gelatinosa.
R. incurvata (Morgan) comb. nov.

Basinym: Clavaria incurvata Morgan, Cincinn. Soc. Nat. Hist. Journ. rr, 1888, 88,
t. 2, f. 2: Burt, N. Am. Clav. 1922, 62, t. rr, f. ror.
-rr cm. high, fragile, stem white, branches ochraceous: stem 3-4X 2·5 cm.:
branches dichotomous, much branched, flexuous, spreading, curving outward and
upward, apices dentate.
On the ground in woods: U.S.A. (Miami, Ohio).
Spores ochraceous.
Very doubtful, no specimen known. Coker suggests R. Strasseri, which Morgan's
figure is said to resemble.
R. intricatissima (Speg.) comb. nov.

Basinym: Clavaria intricatissima Speg., An. Mus. Nac. Buenos Aires, 19, 1909, 279.
Caespitose in fragile, subglobose or obovate tufts r-2 cm. high and wide, colour?:
stems 2-5 x r-1 ·5 mm., arising from a spreading mycelium, abruptly branched above:
branches short, soon closely 2-3-chotomous, tips very short, acute, often fimbriate
denticulate.
On rotting fruits of Eucalyptus globulus: Argentine.
Spores 3µ, wide, rubiginous, verrucose-gibbous, subglobose, r-guttate.
Perhaps this is the conidial state of a Pyrenomycete.
R. lnvalii (Cott. et Wakef.) Donk

Rev. Nieder!. Homobas. Aphyll. 2, 1933, rr3.
Basinym: Clavaria Invalii Cott. et Wakef., Tr. Brit. Myc. Soc. 6, 1919, 176.
Synonyms: Clavaria abietina Fr., Syst. Myc. r, 1821, 469 (quoad descr., non quoad
Pers.): Coker, Clav. U.S. Can. 1923, 182, t. 69, ut 'non-virescent form of pines':
Journ. El. Mitch. Sci. Soc. 1947, 54: Lundell et Nannf., Sv. Svamp. 3-4, 1936, 18.
Clavaria fiaccida var. Invalii Korir. et Maubl., le. Se!. Fung., t. 488: (specimina
an R. stricta?).
Clavariella abietina (Fr.) Karst., Hattsv. 2, 1882, 184.
4-8 cm. high, solitary, gregarious or caespitose, deep ochre becoming yellowish cinna
mon then brown-ochre in age, tips concolorous or pallid: stem r-2·5 cm.X 3-15 mm.,
distinct, often tomentose with a white mycelial felt and slender rhizomorphs: branches
rather slender, numerous, rather irregularly branched, even somewhat flattened, rigid,
erect, fastigiate, cylindric, pointed, the main branches few, 3-5 mm. wide, sometimes
merely forked twice, soon dichotomous with elongate internodes 2-3 mm. wide, and
erect, parallel, tapered ends r-r·5 mm. wide: flesh white, slightly ochraceous on cutting,
spongy, dry, rather tough, brittle in the branchlets: taste slightly bitter: smell slightly
pungent or sourish.
Under Pinus (Coker, Corner, Donk, Lundell), also Picea (Cott. and Wakef.):
Europe (uncommon), U.S.A. (N.Y., N. Car.).
Spores 6-10·5X3·5-5(-6)µ,, ochraceous, closely echinulate with short, acute,
hyaline spines 0·5µ, long, or 0·5-rµ, long in other collections, elongate, pip-shaped.
Basidia 40-48X 9-10·3 µ, (30-45X6-9 µ,, auctt.), with yellowish guttulae: sterigmata
(2-)4, 7-9µ, long.
RAMARIA 599
Hymenium not thickening or very slightly (-'75 µ,, Coker), becoming rugulose
when old.
Hyphae 3-10µ, wide, -12µ, in places, clamped, thin-walled, the walls becoming
firm but scarcely thickened in the older tissue, with frequent abortive clavate
branchings.
Colour in the hymenium and subhymenium caused by pale yellowish cytoplasm
and the yellowish guttulae of the basidia.
The confusion between this species and R. ochraceo-virens ( = Ramaria abietina
Quel., C. abietina auctt.) is explained on page 612. R. Invalii s. str. is distinguished
by its uniform yellow ochre then brownish colour, unchanging flesh, larger size, and
rather small, shortly echinulate spores. It bridges the gap between Echinospora and
Verrucispora, e.g. between R. longicaulis and R. fiaccida. The spores have been
described as echinulate (Cott. et Wakef.), shortly spinulose (Coker), and finely verru
cose (Donk). I found in two collections from Mildenhall (near Cambridge, England)
that the spores had spines 0·5-1 µ, long, and that in another and otherwise identical
collection from Cambridge (England) the spines were 0·5 µ, long like acute warts: if
these spines were blunt, the spores would be practically identical with those of R.
fia.ccida or R. subfennica: as it is, they are like those of R. longicaulis. Indeed, R. Invalii
and R. subfennica appear as hybrids between R. fia.ccida and R. longicaulis. The
branching tends to become flattened.
TABLE LVI. SPORE-SIZE IN RAMARIA lNVALII
Spore-size in µ. E' Authority I Country

8-10·5 X4-5 2'I Donk Holland
7-10X4-6 1·7 Karsten Finland
8·5-9·5 X4-5 2·0 Corner England
7-9X4 2·0 Bourd., Galz. France
7-9X4 2·0 Cott., Wakef. England
7-8(-9) X 4-4·5 I ·6(-2) Lundell Sweden
6·5-8X3•5-4·7 I·8 Corner England
6·3-9X 3·5-4 2·0 Coker U.S.A. (N.Y.)
6-8X 3-4 2·0 Konrad, Maubl. France
6-8X3-4 2·0 Bourd., Galz. "
6-8X3·5-3·8 1·9 Coker U.S.A. (N. Car.)
7'5X4 1·9
I " I " (N.Y.)
1
E = length -:- width.
R. Karstenii (Sacc. et Syd.) comb. nov.
Basinym: Clavaria Karstenii Sacc. et Syd., Syll. Fung. 14, 1899, 238.
Synonyms: Clavariella divaricata Karst., Symb. Myc. Fenn. 32, 1893, 8.
Clavaria divaricata (Karst.) Sacc., Syll. 9, 1891, 135 (non C. divaricata Pk. 1887).
-9 X 6 cm., very much branched, rather tough: stem glabrous, pallid rufescent:
branches divaricate, dichotomous or subverticillate, subfastigiate, almost smooth,
obtuse, rufous, tips at first alutaceous.
In woods of Abies: Finland.
Spores 6-10 x 4-6 µ,, ochraceous, hyaline under the microscope (? smooth).
R. Karstenii suggests R. sanguinea and R. spinulosa, but until it has been re
discovered it must be regarded as doubtful. Conceivably it is the same as R.
divaricata (Pk.).
R. kisantuensis (Sacc.) comb. nov.
Basinym: Clavaria kisantuensis Sacc., Ann. Myc. 4, 1906, 73: Beeli, Bull. Soc.
Roy. Bot. Belg. 58, 2, 1926, 7.
5- 6 cm. high, wholly fuscous-ochraceous: stem very short, branching from the
600 RAMARIA
base: branches 1-1·5 mm. thick, cylindric, elongate, becoming ochraceous-powdered,
sparsely and erectly branched distally, tips furcate, acute, concolorous: drying waxy
fragile.
On fallen rotten branches (Sacc.), in groups on the ground in swampy forest (Beeli):
Belgian Congo.
Spores 5-6µ, wide, ochraceous, glabrous, smooth (Sacc.): 8-9µ, wide, greyish,
round, smooth (Beeli).
This is a doubtful species, probably Clavulinopsis or Clavulina, because round
spores are anomalous in Ramaria. Beeli's fungus is surely different. Compare R.
Henriquesii.
R. leucoceras (Pat.) comb. nov.

Basinym: Lachnocladium leucoceras Pat., Journ. de Bot. 3, 1889, 33.
5-6 cm. high, brown (when dried): stem short, thick, villous, tuberculiform:
branches simple for 2-3 cm. long, then 2-3-chotomous, flat, glabrous, sterile side
smooth, fertile side puberulent, ochraceous, rugulose, axils rounded, sometimes with
a short median blunt point, tips obtusely rounded, white, hymenium unilateral.
On the ground in the forest: Guadeloupe.
Spores 8-10 x 4-5µ,, brown-ochraceous, verruculose, ovoid.
This must be allied with R. Zippe/ii and R. longicaulis, and may be identical with
R. guyanensis.
R. longicaulis (Pk.) comb. nov.

Basinym: Clavaria lol'!gicaulis Pk., Bull. Torr. Bot. Club 25, 1898, 371: Coker, Clav.
U.S. Can. 1923, 187, t. 72-4, 89, 90.
2· 5-9 X 1 ·3'-4 cm. gregarious or subcaespitose, cinnamon-tawny then darker brown,
tips somewhat lighter (not abruptly white as in R. grandis): stem 1·3-4 cm. X 1·5-7 mm.,
whitish downward, arising from thread-like, white rhizomorphs, the mycelial strands
and white part of the stem turning pink when bruised: branches upright, closely appressed,
cylindric or somewhat rugose, rather elongate, irregularly branched, acute: flesh
whitish or cinnamon, in some cases turning dull vinaceous on bruising, not very brittle:
smell slight.
In humus in frondose woods: U.S.A. (N. Car., Alabama, Colorado, Mass.,? Oregon,
rare).
Spores 7·5-u X 4·5-7µ,, ferruginous ochraceous, rather bluntly echinulate, ovoid
or broadly pip-shaped.
Basidia 5·5-7·5µ, wide, 4-spored.
Hymenium 60-75µ, thick.
Hyphae 4µ, wide, clamped, closely packed.
This species appears to be distinguished by the 4-spored basidia and concolorous
tips from the R. Zippelii-grandis alliance, but, as mentioned on page 636, R. Zippelii
var. gracilis may be the same as R. longicaulis, for it may have 3- or 4-spored basidia.
R. megalorhiza, from Ceylon, is similar but paler, lignicolous, and with smaller spores.
I am not sure how R. longicaulis differs from R. Invalii except in its rufescence.
The following are Coker's spore-measurements:
North Carolina 2287 8-n X 4·8-5·3 µ, (E "= 1·9)
Beardslee 8·2-10·5 x 4·8-6·8µ, (E = 1·6)
4416 7·5-8·5x4·8-5·6µ, (E = 1·5)
Alabama Peck (type) 7·5-8·5 X 4·5-5·5µ, (E = q)
North Carolina 2762 7·5--1·7 x 5·5-7µ, (E = 1·2 ? correct)
Massachusetts Davis 7·5 X 5·3 µ, (E = 1·4)
Colombia Braendle 7·5 X 5µ, (E ;= q)
I collected once, in Malaya, a fungus which seems to be the same as R. longicaulis,
RAMARIA 601
although, as mentioned on page 590, it may well be the fungus intended as C. echino
spora P. Henn. ( = Ramaria fragillima). The following is its description:
-6 cm. high, scattered, wholly dingy cinnamon-yellow, tips concolorous: stem
-15 X 2-3 mm. dilated above into 4 main branches: branches 1·5 mm. wide, cylindric,
becoming dichotomous with rather elongate nodes, tips subacute: flesh whitish or pale
ochraceous, not changing colour: on the ground in the forest, Singapore (Reservoir
Jungle, 20. 10.30).
Spores 8-10 X 4·5-5µ,, ferruginous-yellow, shortly echinulate with many slender
spines 0·5-0·7µ, long, elongate pip-shaped, apiculate.
TEXT-FIG. 259. Ramaria longicaulis: fruit-body

from i\1Ialaya (Singapore, 20.10.30), X 1.
Basidia with 2-3-4, mostly 4, sterigmata 6-10µ, long.
Hymenium thickening (but young).
Hyphae 3-13µ, wide, clamped, thin-walled.
Structure as in R. Zippelii.
Fruit-body like R. fragillima, but tips not orange-ochre, spores smaller, flesh
unchanging. Spores and colour of fruit-body as in R. longicaulis, but with wide hyphae
and variable sterigmata as in R. fragillima: Text-Figs. 250, 259.
R. luteo-flaccida sp. nov.

-4· 5 cm. high, solitary or gregarious, small, yellowish drab to pale yellowish ochraceous,
tips clearer and lighter yellow, with white unchanging flesh: stem 6-16 X 2-3 mm.·,
rather slender, with a few main branches, pallid yellowish or ochraceous drab,
becoming dingy, whitish villous at the base with slender white mycelial strands:
branches 2-3 mm. wide below, 1-2 mm. above, 2-3-chotomous, slightly dilated at the
points of branching, cylindric, ascending-erect, lax or rather crowded,. rather spread
ing, tapering to the acute, bifid, or subulate tips: flesh white, unchanging, rather spongy
flaccid, not brittle: smell in particular: taste?
In humus in the forest: Sumatra (Brastagi, c. 1,400 m. alt., 12.9.31).
Spores 5-6·5 X 3-3·5µ,, ochraceous, verruculose (even subechinulate), the fine warts
less than o·5µ, long, ellipsoid, pip-shaped, blunt at the apex.
Basidia 45-55 X 6-7µ,, subclavate: sterigmata 4, rarely 2-3, 5-6µ, long.
602 RAMARIA
Hymenium c. 70 µ. thick, thickening slightly: subhymenium 20 µ. thick, composed
of rather loosely interwoven hyphae 2-3 · 5 µ. wide.
Hyphae 3-<J/L wide, -12µ. in places, clamped, mostly with slightly, but distinctly,
thickened walls (c. o· 5µ.), cells 30-250 µ. long, short next the subhymenium: clamp
connexions often inflated 6-12µ. wide, occasionally branched from the clamp: H-con
nexions frequent, sometimes one or both branches more or less inflated-pyriform,
6-II µ. wide: sometimes the cells bearing short, lateral clavate processes -40 x 7-12 µ.
with rather dense contents.
� :::::::
TEXT-FIG. 260. Rarnaria luteo-fiaccida: spores, X r,ooo: clamp-connexions and

clavate branches of the medullary hyphae, :< 500.
Structure as R. formosa: medullary hyphae often somewhat convoiute, the outer
hyphae of the flesh rather narrow and compact: surface of the stem with narrow
longitudinal hyphae giving off a few processes 2-3 µ. wide, -50 µ. long.
Hymenium covering most of the stem, except.the basal 2-6 mm., and on all sides
of the branches, but absent from the uppersides of some of the main branches.
Macroscopically this is like a small R.fiava, but the spores are those of R. ochraceo
virens and R.fiaccida. The hyphae, also, have the'same swellings on the cells, clamps,
and H-connexions as in R. ochraceo-virens. But, whereas it is not R. ochraceo-virens,
it might be regarded as a variety of R. fiaccida. Such was my intention until I decided
,.
that to do so might be confusing. R. fiaccida appears to be a species of the temperate
coniferous forests: it has not been reported from the tropics, though it grows under
conifers in the south temperate regions, and its colour appears never to be the pale
clear yellowish ochraceous of R. luteo-fiac:cida. In the forest where I found this species.
there were scattered conifers such as Dacrydium and Podocarpus, but in such mixed
RAMARIA 603
vegetation it is impossible to discover from a single collection with what tree, if any,
a terrestrial fungus may be connected.
Compare R. filicicola.
R. luteo-fusca Maire
Fungi Catal. 2, 1937, 32.
3-4 cm. high, generally caespitose, umber with vivid ochraceous-yellow tips: stem
-12 X 1 mm., slender, umber, more or less tomentose at the base with paler mycelium:
branches terete, dichotomous, lax: flesh whitish or subfuscous: taste bitter, smell
almost none.
On the ground in fields: Spain (Catalonia, Santa Pau).
Spores 8-10 X 3-3·5 µ., ochraceous, distinctly verrucose, oblong, rounded at the
apex, attenuate at the base, not blueing with iodine.
Basidia 38-42 X 6 µ., 2-spored, cylindric-clavate.
Subhymenium c. 7-8 µ. wide, thin.
Hyphae 5-10µ. wide, without clamps, thin-walled, parallel, strongly adherent
together.
Maire compares this with R. fiaccida and R. crispula. The yellow tips recall R.
condensata. Apart from the uncertain R. macrospora, it is the only 2-spored Ramaria
other than those with echinulate spores, and this feature together with the intense
coloration suggests that it is a link between the Echinospora-group and Verrucispora,
unless it has echinulate spores and belongs to Echinospora. Its habitat in fields is
exceptional, but there may have been relics of woodland on the site of the field.
The spores are remarkably narrow for a 2-spored Ramaria: they resemble the
narrow 4-spores of R. Murrillii.
R. macrospora (Brinkm.) comb. nov.

Basinym: Phaeoclavulina macrospora Brinkm., Jahresb. Westf. Prov. Ver. Wiss.
Kunst. Bot. Sect. 1896-7, 197.
3-6 cm. high: stem fleshy: branches thick, irregularly branched, erect, smooth or
longitudinally rugulose, sordid ochraceous or subolivaceous, tips obtuse, 2-4 tmncate,
·concolorous: flesh dirty white, turning reddish or greenish black on bruising.
In fields under Juniperus, on calcareous soil: Germany (Westphalia).
Spores 14-18 X 5-7 µ., yellowish, ellipsoid, 'granulose', rough.
Basidia 2-spored.
This species is the basis of Phaeoclavulina, made in imitation of Clavulina but
without its significance. Brinkmann then added C. nigrescens and van Overeem the
tropical 2-spored species with echinulate spores, but there is no evidence that R.
macrospora with 'granulose' spores is nearly related to them.
R. macrospora appears to belong to the R. decolorans-R. xanthosperma group as a
2-spored derivative with slightly wider spores. Its rediscovery will be of great interest,
when described in detail.
R. madagascariensis (P. Henn.) como. nov.

Basinym: Clavaria madagascariensis P. Henn., Eng. Jahrb. 17, 1893, 22.
-7 cm. high, fleshy, branched, ochraceous: stem -1 cm. thick: branches elongate,
erect, parallel, crowded, rugulose: branchlets repeatedly divided, parallel or divergent,
with obtuse or cristate, multifid tips.
On the ground in the forest: Madagascar.
Spores u-14 x 4-6 µ., pale yellowish under the microscope, granular or punctate
rough, oblong, curved.
This is said to be near R. aurea, but the shape suggests R. fiavo-brunnescens. There
are two collections in the Kew herbarium which seem to be the same species, one
RAMARIA
from Java and one from Malaya, but neither van Overeem nor myself found them in
the living state:
Clavaria javanensis B. ms.
-10 cm. high, in form as R. subgelatinosa, colour ?: spores 11-13X 5-7 /L, ochraceous,
rugulose-verrucose: Java(? locality, date, and collector).
Ridley No. u650 (Penang Hill, Malaya, 1902): spores 10-u·5X6·5-7·5 /L, ochra
ceous, rather broadly and bluntly pip-shaped, with small, flat, discrete warts or
patches 0·2-0·3 /L high.
R. Mairei Donk
Synonyms: Clavaria pallida Bres., Fg. Mang. 1906, u6: le. Myc., t. 1088:
Malenc,:on, Bull. Soc. Myc. Fr. 41, 1925, t. 3: Konr. et Maubl., le. Se!. Fung., t. 490:
(non C. pallida B. et C. 1869, vide Ramariopsis Kunzei): (C. pallida Schaeff., Fung.
Bav. 1770, t. 286): Huber, Zeitschr. f. Pilzk. 18, 1934, 34.
Ramaria pallida(Bres.) Ricken, Vadem., 1920, 263.
? Clavaria extensa Herpell, Hedw. 52, 1912, 391.
TABLE LVII. SPORE-DATA FOR RAMARIA MAIRE!

10-16 X4-5 Corner Austria see Table LVIII
10--15·5 X 5-6·7 " England see Table LIX
9-12X5-6 Bourd., Galz. France
9-12X4·5-5·5 Konr., Maubl. "
9-12X4·5-5 Rea England
9-12X4-5 Bresadola Italy
6-18X2-10 cm., pale to deep lemon-yellow or pale ochraceous-cream, the young

branches generally suffused pale lilac or pale purple towards the ends, becoming wholly pale
ochraceous-yellow: stem 2-8 X2-4 cm., thick, distinct, often tinged pale coffee-colour,
pallid white at the base: branches numerous, elongate, rather lax or fairly crowded,
longitudinally rugulose, obtuse, sometimes tinged flesh-colour: flesh white, unchanging,
brittle, not hygrophanous, not drying chalky: taste sweet then bitter(especially after
cooking): indigestible or dangerous (Huber).
On the ground in frondose or, mainly, coniferous woods: Europe (frequent),
Morocco (Atlas Mts.).
Spores 9-16X4-6·7µ, mostly II-13X4·5-5·5µ, pale ochraceous, finely rough,
oblong, ellipsoid, 1-3 guttulate.
Basidia 75--90 X8-9 /L(Rea), 45-80X6-9 /L(Bourd. et Galz.), 40-60X8-9 /L (Bres.):
sterigmata 2-4, 8--9µ long.
Hyphae 2·5-8 /L wide, clamped, as in' R. formosa.
The recognition of this species in Europe has helped very effectively to clear up the
confusing statements which have been made about R. Jormosa, R. fiava, and, perhaps,
R. aurea. I have collected the species in Austria and on several occasions in England,
and have found it to be somewhat variable. The lilac colour may develop early, when
the fruit-bodies are easily recognizable, for the distal half of the branches, not merely
their tips, are so coloured, but in some cases it develops late, the young fruit-bodies
being entirely pale clear yellow(the colour ot R. fiava or of the tips of R. formosa),
and occasionally it does not develop at all, such fruit-bodies remaining pale yellow in
a troop of normally coloured specimens: and, generally, as the fruit-bodies become
pale ochraceous from the spores, so the lilac colour fades. Thus old and young fruit
bodies may be undistinguished, and such have undoubtedly been mistaken for R.fiava
.RAMARIA 605
and R. formosa, to which several authors have attributed a wholly yellow form. Such
states of R. Mairei may have been the reason for the small spores described by several
authors for R. fiava and for the distinctly verruculose spores described for R. aurea.
The spores are indistinguishable from those of R. formosa, unless a trifle less verru
culose, and though the branches are rather more elongate and lax in R. Mairei than
in R. formosa, the two can be distinguished only by their difference in colour and the
unchanging flesh of R. Mairei. C. extensa appears to be the undistinguished state of
R. Mairei, unless it is R. fiavo-brunnescens.'
Bresadola records his C. pallida from N. America, Ceylon, Malaya, and Australia,
but the records need confirmation from fresh collections. Dried specimens, without
adequate field-notes, could not be identified certainly.
TABLE LVIII. SPORE-SIZE IN RAMARIA MAIRE]

(One fruit-body, Burgenland, Austria, Sept. I924)
Size in µ. E* Number of records Size in µ. E*. I Number of records
10·0X4·3 2·3 I 13·ox5·0 2·6 2
10·0X4·5 2·2 3 13·5X4·0 3·4 2
10·0X4·7 2·1 2 13·5X4·5 3·0 I
10·5X4·0 2·6 I 13·5x5·0 2·9 2
10·5X4·5 2·3 l 14·0X4·5 3·1 2
ro·5X4·7 2'2 2 14·0X4·7 3·0 2
r1·0X4·5 2·4 3 14·0X5·0 2·8 3
u·oX4·7 2·3 l 14·5X4·0 3·6 l
11'5X4·5 2·6 I 14·5X4·5 3·2 I
u·5X4·7 2·4 3 14·5X5·0 2·9 I
11·5X5·0 2·3 2 15·0X4·0 3·8 l
12·0X4·3 2·8 2 15·0X4·5 3·3 I
12·0X4·7 2·6 I 2 15·0X4·7 3·2 I
12·0X5·0 2·4 I 15·0X5·0 3·0
I
3 2
12·5X4·5 2·8 I 15·5X4·5 3·4 I
12·5X4·7 2·7 I 15·5X4·7 3·3 2
12·5x5·0 2·5 I 15·5x5·0 3·1 I
I
13·0X4·5 2·9 I 16·0X4·7 3·4 I
13·0X4·7 2·8 2
* E = length -:- width.
Range, 10-16X4-5 µ., E = 2·.1-3·8, for 60 spores.

Average of 10 spores, 10-10·5 µ. long, E = 2·2
11-u·5µ. E=2·4
12-12·5 µ. E = 2·6
13-13·5 µ. E = 2·9
14-14·5 µ. E = 3·0
,, ,, 15-16µ. ,, E = 3·3
Average of 60 spores, 12·4X4·6 µ., E = 2·7
C. megalorhiza (B. et Br.) comb. nov.

Basinym: Clavaria megalorhiza B. et Br., Journ. Linn. Soc. Bot. 14, 1875, 75:
Coker, Clav. U.S. Can. 1923, 189, t. 89: Petch, Ann. R. Bot. Gdn. Per. 9, 1925, 335.
Synonym: Clavaria intricata Ces., Myc. Born., 1879, II; Sacc., Sy!!. 6, 1888,
702 (fide Petch).
1
C. extensa Herpell: -10 cm. high, light yellow: stem 2·5X 1·5 cm. thick, short, fleshy:
branches stout, subcompressed, more or less divaricate, longitudinally sulcate, rugulose,
branchlets with dense, obtuse, botryoidal tips: flesh white, soft: spores 8-13 X 4-5 µ., dull
yellow, oblong(? rough, ? smooth): taste mild, slowly bitterish: in woods, Germany.
606 RAMARIA
-4·5 X 2·5 cm., white, often tinged purple, yellow-brown when old, bushy, arising from
a single, short, stout stem or with several stems united by a sheet ofmycelium, with lax
rhizomorphs -4 mm. wide: stem repeatedly divided, usually from near the base:
branches numerous, divided at an acute angle, the upper branches broadly expanded
below the axils, terminating in a cluster of erect processes a few millimetres long.
TABLE LIX. SPORE-SIZE IN RAMARIA MAIRE!
(One fruit-body, Nettlebed, England, Oct. I927)
Size in µ E* Number of records Size in µ I E* Number of records
10·0X5·0 2·0 I 12·5X5·7 2·2 I
10·0X5·3 1·9 2 13·0X5·3 2·5 I
10·0X 5·5 1·8 I 13·0X5·5 2·4 3
10·0X6·0 1·7 l 13·0X6·0 2·2 I
10·5X5·0 2·1 I 13·5X5·3 2·6 2
10·5X5'3 2·0 3 13·5X5·7 2·4 l
10·5x5·5 1·9 I 13·5X6·o 2·3 2
11·0X5·0 2'2 l 14·0X5·0 2·8 2
11·0X5·3 2·1 I 14·ox5·5 2·5 I
rr·5X5·0 2·3 l 14·0X6·o 2·3 3
11·5X5·3 2·2 3 14·0X6·5 2·2 l
11·5X5·5 2·1 2 14·5X6·o 2·4 l
11·5x5·7 2·0 l 14·5X6·3 2·3 2
rr·5x6·o 1·9 I 15·0X5·5 2·7 I
12·0x5·0 2·4 I 15·0X5·7 2·6 2
12'0X5'3 2·3 2 15·0X6·o 2·5 I
12·0X5·5 2'2 I 15·0X6·5 2·3 2
12·0X 5·7 2'1 l 15·5X6·o 2·6 l
12·5X5·0 2·5 I 15·5x6·3 2·5 I
12·5X5·3 2·4 2 15·5x6·5 2·4 l
12·5X5·5 2·3 I 15·5X6·7 2·3 I
• E = length -:- width.
Range, 10-15·5X5-6·7 µ, E = 1·7-2·8, for 60 spores.

Average of 10 spores, 10-10·5 µ long, E = 1·9
rr-rr·5 µ E = 2·1
12-12·5 µ E = 2·3
13-13·5 µ E = 2·4
14-14·5 µ E = 2·4
,, ,, 15-15·5 µ E = 2·5
Average of 60 spores, 12·8X5·6 µ, E = 2·3
Among humus and on rotten wood: Ceylon.

Spores 5-7 X 3-5 p, (5-7 X 3 ·5-3 ·8 p,, Coker), yellow-brown in the mass ( dark ochra
ceous under the microscope, Coker), verruculose with small distinct blunt warts (not
echinulate), ovoid or oblong ovoid.
I have placed this in the Verrucispora-group because of the verruculose spores, but
the lignicolous habitat suggests that it may have thick-walled hyphae as in the Stricta
group. Indeed, R. polypus of Malaya may be the same. Compare also R. filicicola.
R. Moelleriana (Bres. et Roum.) comb. nov.

Basinym: Lachnocladium Moellerianum Bres. et Roum.. Re'(, Myc. 1890, 36,
t. 92, f. 6.
Synonyms: Lachnocladium Warburgii P. Henn., Hedw. 1893, 218: Monsunia 1,
1900, 8: (teste Bres., Sacc. Sy!!. 23, 1925, 495).
L. Lauterbachii P. Henn., Engl. Jahrb. 18, 1894, Beibl. 25: (teste Bres., I.e.).
RAMARIA
L. strictu.m P. Henn., Engl. Jahrb. 22, 1895, 86: (teste Bres., Le.).
? L. brunneum Bres., Ann. Myc. 9, 1911, 272.
3-10 cm. high, caespitose, much branched, coriaceous, tough, yellow or ochraceous:
stem -2 cm. X 3-5 mm., subterete, slightly thickened and white tomentose at the base,
arising from a pallid white mycelium: branches repeatedly dichotomous, strict, erect,
axils. arcuate and flattened, or acute and scarcely flattened, apices subcristate or
elongate-subulate: hymenium unilateral, on the lowerside of the main branches.
On dead wood: trop. Africa, trop. Asia, Bonin Isl. (L. Warburgii), New Guinea.
Spores 6-8 X 3-5µ, (6-7 X 4-4·5µ, in the type), ochraceous, fuscidulous, or pallid,
smooth, ellipsoid: 7-9 X 4·5-5·5 µ, ochraceous, finely punctate-rough (in the type of
L. strictum).
Hyphae (in the type of L. strictum) 3-8µ, wide, long-celled, clamped, with slightly
or distinctly thickened walls, particularly in the narrower hyphae.
This species is known only from dried specimens. I have examined the part of the
type-collection of L. strictum in the Kew herbarium and, assuming that Bresadola's
identification is correct, there can be no doubt that L. Moelleriana is a Ramaria allied
with R. stricta. The species is not mentioned by Petch for Ceylon or by van Overeem
for Java, and I have not found it in Malaya. Possibly L. palmatifidum, L. pteruloides,
and L. ralumense belong here. How this tropical group differs from the temperate
R. stricta-R. apiculata remains to be discovered. Compare also Lentaria tjibodensis.
The description of L. brunneum is:
-8 X 5 cm., brownish ochraceous: stem 1·5 cm. high and wide, tuber-like, almost
subterranean, arising from a white mycelium: branches repeatedly dichotomous, axils
compressed, arcuate, fertile side smooth and brown, sterile side puberulous rugose and
lurid ochraceous, tips 1-3-chotomous, acute: flesh spongy-coriaceous, lurid straw
colour.
On the ground: Congo (Kinkosi).
Spores 7-8 X 3·5-4·5µ,, pinkish tawny(? under the microscope), ellipsoid(? smooth).
Hyphae 2-7µ, wide, 'fumosis'.
Bresadola compared it with Ramaria spinulosa, but I can see no difference in the
description from R. Moelleriana except in habitat: the fruit-bodies, however, may have
grown from buried wood.
R. Murrillii (Coker) comb. nov.
Basinym: Clavaria Murrillii Coker, Clav. U.S. Can. 1923, 190, t. 77, 91: Heim,
Fungi lberici, 1934, 49, f. 11.
1 ·5-9 cm. high, solitary or gregarious, slender, laxly branched, ferruginous,, tips
brownish: stem -5 cm. long, long and slender, dull flesh-colour, loosely covered through
out most of its length with long, pale, flexuous cottony threads(when dried): branches
few, elongate, ascending, branched one to three times, tips subacute.
In humus in mixed woods: U.S.A. (Tenn.), Europe (Spain).
Spores 8·5-9·5 x 3-4µ,, brown under the microscope, echinulate with slender, acute
spines 0·5-1 µ, long, elongate pip-shaped.
Basidia 5-5·5 µ, wide, 4-spored.
Hymenium 50-60 µ, thick.
Hyphae 3-5·5µ wide, clamped.
Coker says this is distinguished by its slender, lax form, pale cottony stem(? when
dry), and long narrow spores, and that it comes nearest to R. longicaulis. It seems to
me also like a lax form of R. lnvalii with narrow spores (E = 2·6).
Compare R. luteo-fusca (umber with yellow tips), which has spores as narrow.
R. myceliosa (Pk.) comb. nov.

Basinym: Clavaria myceliosa Pk., Bull. Torr. Bot. Cl. 31, 1904, 182: Coker, Clav.
U.S. Can. 1923, 178, t. 88: Doty, Clav. Pac. N.W. 1944, 41.
608 RAMARIA
-4 X 3 cm., rarely 7 X 5 cm., branched abruptly five to eight times from the slender
base, with copious white rhizomorphs, ochraceous-buff, honey-yellow, pale orange-yellow
or cinnamon-drab, not discolouring: stem slender or indistinct: branches slender,
-1 mm. wide, spreading: flesh white, taste bitter, smell slight.
In humus under conifers (Pseudotsuga, Tsuga, Abies, Sequoia): U.S.A. (California,
Oregon).
Spores (3·5-)4-5(-6) X(2-)2·5-3·5(-4)µ, ochraceous, acutely verruculose or sub-
echinulate, ovoid to subglobose.
Basidia 15-22 X4·5-6µ: sterigmata 4, rarely 2,-3, 2-5 µ.
Hymenium ? not thickening (25-30µ thick).
Doty has considerably enlarged the description of this species, which is common in
all coniferous forests of the Pacific north-west, though usually referred to R. fiaccida.
Compare the lignicolous R. Bourdotiana with equally small spores. The spores are
verruculose with acute warts, less than 0·5 µ long, rather than echinulate (as in
Echinospora).
R. nguelensis (P. Henn.) comb. nov.

Basinym: Clavaria nguelensis P. Henn., Engl. Jahrb. 28, 1901, 320.
4-6 cm. high, yellow-brownish, fleshy, strict: stem short, rather thick, white
byssoid: branches repeatedly dichotomous, erect, subterete, smooth, tips subulate or
cristate.
On rotten wood: E. Africa (Usambara).
Spores 5-6 X3·5-4µ, epispore pale fuscidulous, smooth, ellipsoid-ovoid.
Possibly R. Moelleriana.
R. nigrescens (Brinkm.) Donk

Rev. Niederl. Homob. Aphyll. 2, 1933, 104.
Basinym: Phaeoclavuliria nigrescens Brinkm.: (non Clavaria nigrescens Fr., vide
Ramaria Zippelii).
Synonyms: Clavaria Broomei Cott. et Wakef., Tr. Br. Myc. Soc. 6, 1919, 570 (fide
Donk).
Phaeoclavulina Broomei(Cott. et Wakef.) v. Ov., Bull. Jard. Bot. Buit. Ser. 3, 5,
1923, 277.
4-8 X2-4 cm., solitary or gregarious, ochraceous-orange, tips darker orange, quickly
turning brown on bruising: stem short, not swollen, white: pinkish on bruising: branches
numerous, irregular, suberect, tips flattened, the larger branches rugulose: flesh white,
becoming vinous on bruising, especially in the stem, blackening on drying, rather tough:
taste bitter: mycelium white, scant.
On the ground in woods: Europe (uncommon), Japan.
Spores 12·5-20 x 5-8 µ, av. 15-16 X6-7 µ (Cott. et Wakef.), deep ochraceous to
orange, markedly aculeate, fusiform or pip-shaped.
Basidia 40-50 x 8-<)µ: sterigmata 2.
Hyphae 3-6µ wide, with vesicular ends 10-12µ wide: many large crystals in the
tissue.
var. americana var. nov.

Synonym: Clavaria Broomei sensu Coker, Clav. U.S. Can. 1923, 186, t. 71, 89.
Flesh not changing colour, tasteless: spores 12·5-18·5 X4·5-6µ: basidia 4-spored.
3-7 cm. high, ochraceous-brown, reddish brown with age, often blackening in
places on drying: stem 2-5 cm. x 2-4 mm., long, distinct: flesh watery white, not very
brittle: without smell or taste: spores deep reddish ochraceous, papillate (shortly and
subacutely echinulate).
In humus in frondose woods, or frondose woods with Tsuga: U.S.A. (N.Y.).
RA.MARIA 609
This is very near the tropical R. fragillima, particularly the 4-spored state var
americana, but I hesitate to unite them while the temperate species is so little known
TABLE LX. SPORE-DATA FOR RAMARIA NIGRE'SCENS
Spo re-size in µ, . _E_*'_ __A_u_th_o_ ri_·ty_ _

_ _ _ o_ _u_nt_r_y__
C Rema _rk s
,I l _ _ ___
14-20 X6-8 2 ·4 Cott., vVakef. Gt. Britain C. Broomei (type): av.
15-16x6--7µ,
14-20 X 6--8 2·4 Imai Japan I as C. Broomei
12·5-19·5 X 5-7 2'7 Coker Gt. Britain C. Broomei (type)
14-19·5 X 5·5-7 2·6 Europe C. nigrescens: Herb. Bresadola
I 1-16 X5-6·5 I 2·4 Sweden Specimens from Romell
12·5-18·5 X4·8-6 2·9 U.S.A. (N.Y.) var. americana (Burnham 98)
13·5-16X4·5-5·5 13·0 var. ame ricana (Burnham 141)
13·5-15·8X4·8-5·6 2·8 var. americana (Burnham 74)
*' E = length -,- width.

R. obtusissima (Pk.) comb. nov.
Basinym: Clavaria obtusissima Pk., Bull. N.Y. St. Mus. 167, 1913; Coker, Clav.
U.S. Can. 1923, 146, t. 56, 87: Pova, Pap. Mich. Ac. Sci. Arts Lett. 20, 1935, 141:
Wehm., Pap. Mich. Ac. Sci. Arts Lett. 20, 1935, 262: Doty, Clav. Pac. N.W. 1944, 58.
Synonyms: Clavaria obtusissima var. minor Pk., Bull. N.Y. St. Mus. 167, 1913, 34
(teste Coker).
? Clavaria albida Pk., Rep. N.Y. St. Mus. 41, 1887, 79.
9-15 cm. high, -13 cm. wide, large, massive, whitish, pale cream or yellowish buff
then cream-tan or ochraceous, fading to reddish brown (Coker), warm buff, amber
yellow, light ochraceous-buff or buff-yellow, tips lighter yellow (baryta- to apricot-yellow)
(Wehm.): stem 2-5 cm. wide, short, massive, breaking up into stout main branches at
ground level: branches dense, 1-5 mm. wide above, upright, rather elongate,.repeatedly
divided, tips blunt: flesh white, rather brittle, fleshy, fibrous or somewhat elastic,
unchanging: taste rather bitter: smell mildly rancid or, in drying, like ham (Coker),
faintly sweetish (Wehm.), of aniseed (Doty): edible (Doty).
On the ground in frondose or coniferous woods: U.S.A. (frequent), Canada (Nov.
Scot.).
Spores 9-15 x 3-5 µ., ochraceous, smooth, subcylindric-ellipsoid, often bent or sway
backed (Coker): 10-13 X 3-3·5 µ., pale brown, smooth, ellipsoid-cylindric (Wehm.):
12 x 4·2µ., smooth (Doty).
Basidia 7· 5-9· 5 µ. wide, 4-spored (Coker), 70 X 9 µ., with 4 sterigmata, 6 µ. long (Doty):
hymenium 90-100µ. thick: hyphae 3-8 µ. wide, clamped.
It is difficult to understand this species in view of the many variations which have
been ascribed to it. I have, therefore, restricted the foregoing description to what
seems to be the essential idea, that is a yellow Ramaria like R. formosa but with smooth
spores and unchanging flesh. The variations, described below, leave only the smooth
spores (perhaps, also, flesh not chalky-friable when dried) as the sole distinction from
Rough-spared form: spores 9·5-15 X 3·7-5·5 µ., ochraceous-yellow, distinctly rough:

R. Jormosa.
-9 X 3-6 cm., pale cream-tan then cream-ochraceous, tips concolorous, moderately

brittle,taste sweetish,smell none: N. Car. No. 3792: (Coker): ?Smith 16258,Washington.
This seems referable to R. Mairei or R. divaricata.
Pink form: fruit-bodies flesh-coloured; spores II-14 X 3 ·7-4· 5 µ., ochraceous,
smooth; N.Y., Burnham 96; (Coker): fruit-bodies pink like R. formosa but without
yellow tips; spores 10-13 X 3-3·5 µ., smooth; Nov. Scot. (Wehm.): light ochraceous
buff to light ochraceous-salmon (Zeller's collections, Doty): very young yellow tips
vinaceous or rosy round the base (Doty).
5119 Rr
610 RAMARIA
This must resemble R. secunda or pale specimens of R. subbotrytis, the spores of
which, however, are shorter and more or less rough.
Vinescent form: tips and bruised parts very slowly turning dull cinereous-violet or
reddish, or, in another collection, the staining more marked; spores 10-13 X 3-3·5µ,
smooth; Nov. Scot. (Wehm.): upper branches violet when old (coll. MD 976) or
turning violet when cooked (coll. MD 3153); Oregon (Doty).
Such staining may be normal, as it is in the allied R. decolorans and R. xanthosperma:
it is also a point in common with R. formosa.
Rich yellow form: cadmium-yellow then deep chrome, whitish at base, tips con
colorous; flesh white or pinkish, unchanging, brittle; taste mild; smell slightly dis
agreeable; spores 9-10· 5 X 3·5-4·5 µ, pale ochraceous, smooth; Michigan (Pova).
This may be a form of R. fiavo-brunnescens with smooth spores, cf. var. aurea.
Coker gives C. albida Pk. as a synonym, in which case it provides the earliest specific
(if inappropriate) epithet, 1 but it is possibly R. xanthosperma. Its description is:
-5 cm. high, whitish: stem short, thick: branches few, short, thick, much divided
into crowded branchlets with short acute tips: flesh firm, whitish, acrid then bitter:
sp. 14-15 X 3·8-5 µ (Coker), 12-15 x 2µ, pale ochraceous (Pk.), smooth: on the ground
in woods, U.S.A.
From Table LXI it can be seen that R. obtusissima has spores as variable in size and
ellipticity as R.formosa. The measurements are, presumably, average samples, so that
the range in ellipticity, 2·7-3·6, is not so great as that of R.formosa (2·1-3·3, according
to my averages, which include extremes). It may be that there is a narrow-spared
variety, with spores 3-4µ wide, as in Wehmeyer's collections from Nova Scotia.
TABLE LXI. SPORE-MEASUREMENTS OF RAMARIA OBTUSISSIMA
Spore-size in µ,
---·----
E*' Authority Country Remarks
II-14·5X3-4 3·6 Coker N. Car. No. 2896
IO-I3X3-3·5 3'5 Wehmeyer Nova Scotia several collections
14-15X3·8-5 3·3 Coker N.Y. C. albida (type)
II-IS X3·5-4·5 3·3 " Mo. (Glatfelter)
II-14X3·7-4 3·2 " N. Car. Nos. 2860, 2870
10·5-13X3:3-4 3·2 " ,, No. 577
9·3-u·3X 3·2-3·5 3·1 ,, ,, No. 2919
II-I4X3·7-4·4 3·1 ,, " No. 2866
12-15X4-5 3·0 " N.Y. Burnham 96a
8·7-12X3·2-3·7 3·0 ,, N:Car. No. 2909
I2X4'2 2·9 Doty Oregon
II-14·8X4-5 2·9 Coker Mass. C. obtusissima (type)
9'3-II'I X3·3-4 2·8 ,, Alabama
10·5-13·5X3·8-5 2·7 ,, N. Car. No. 3200
10-12·2X 3·7-4·4 2·7 ,, ,, No. 2867
9-10·5X3·5-4·5 2·4 Pova Mich. 'rich yellow form'
9·5-15X3·7-5. 5 2·7 Coker N. Car. 'rough-spored form'
II-14X3·7-4·5 3·0 ,, N.Y. 'pink form' No. 96
• E = length --:- width.
R. ochracea (Bres.) comb. nov.

Basinym: Lachnocladium ochraceum Bres., Bull. Soc. Roy. Bot. Belg. 1899, 157,
t. B, f. 4-5.
5-6 cm. high, caespitose, very much branched from the base, without a stem, drying
fuscous: branches repeatedly dichotomous, solid, subangled, dichotomies semicircular,
1
Clavaria albida Schaeff. (1763) dates from C. albida Schaeff. ex Sacc. (1916) as a valid
name.
RAMARIA 6u
axils compressed, tips acuminate, 2-3 furcate: hymenium unilateral, ochraceous
pulverulent from the spores.
On rotten trunks in the forest: Congo.
Spores 5-5·5 X 3-3 · 5µ,, yellowish, asperulate, obovoid.
Possibly R. polypus, R. megalorhiza, R. nguelensis, or R. durbana, if they are not the
same. But compare Scytinopogon.
R. ochraceo-salmonicolor (Clel.) comb. nov.

Basinym: Clavaria ochraceo-salmonicolor Clel., Tr. Roy. Soc. S. Austral. 55, 1931,
160: Toadst. S. Austral. 2, 1935, 205: Fawcett, Proc. Roy. Soc. Viet. 52, 1940, 154,
t. 6, text-fig. 1.
4-15 X 4-14 cm., compact, much branched, light yellow to deep yellow, light ochra
ceous-salmon or pinkish orange, the tips generally slightly more intensely coloured or
yellower (Cleland): stem 2-4 cm. wide, massive, distinct; branches stout below,
-10 mm. wide, polychotomous, dividing three to five times, becoming sulcate or
longitudinally rugulose, tips expanding into subglobose knobs, cauliflower-like: flesh
whitish or pallid, brittle: smell and taste inparticular.
On the ground in forested or heathy places: S. Austral. (N .S.W., Viet.).
Spores 8-10·5 X 4-5·2 fL (Fawcett), 7·5-13 X 3·5-7µ,, mostly 9-10 X 4µ, (Clel.),
yellow-brown or ochraceous, oblong ellipsoi,d or subpyriforni, finely roughened.
Fawcett has narrowed Cleland's description, for it seemed to include R. fiavo
brunnescens and var. aurea, possibly also R. capitata. Thus defined with short spores,
it is very near R. fiavo-brunnescens var. aurea but differs in the expanded tips which,
however, do not fuse or become viscid as in R. capitata. It is possible, nevertheless,
that R. ochraceo-salmonicolor is intermediate between them.
R. ochraceo-virens (Jungh.) Donk

Basinym: Clavaria ochraceo-virens Jungh., Linn. 5, 1830, 407, t. 7, f. 3: Lund. et
Nannf., Sv. Svamp. 3-4, 1936, 19.
Synonyms: Clavaria abietina Pers., Myc. Eur. 1, 1822, 164: Coker, Clav. U.S. Can.
1923, 179, t. 68, 89: Konr. et Maub., le. Sel. Fung., t. 489: Wehm., Pap. Mich. Ac.
Sci. Arts Lett. 20, 1935, 259: (non C. abietina Fr. = Ramaria Invalii).
Clavaria abietina Fr., f. Persoonii Fr., Hym. Eur. 1874, 671.
Ramaria abietina Quel., FI. Myc. 1888, 467.
Clavaria abietina subsp. cyanescens Romell in Krok et Almq. Sv. Fl. 2nd ed., 19u.
Clavaria virescens Gramberg, Pilze der Heimat, 2nd ed., 3, 1921, 33.
Ramaria virescens (Gramb.) Br. Hennings in Michael. Sch. Fuhr. f. Pilzfr., 3, 1927,
text ad No. 320 (excl. fig.).
Clavaria cyanescens (Rom.) Lundell in Krok et Almq. Sv. ;FI. 5th ed., 2, 1932, 286.
Pseudonym: Clavaria abietina auctt. non Fr. (e.g. Bres., Cott. et Wakef., Rea,
Bourd. et Galz., Doty).
1·5-4·5 cm. high and wide, very compact, dull yellow, dull ochraceous or olive
ochraceotts, greenish or olive-green when bruised or weathered, caespitose from a floccose
white mycelium with rhizomorphs: stem 0·5-1·5 x 0·3-0·8 cm., short, soon branched,
white tomentose at the base: branches 1-2 mm. thick below, numerous, crowded,
erect, irregularly branched, often minutely, polychotomous then dichotomous, blunt:
flesh concolorous, greening, rather tough: taste bitterish: smell rather strong or none.
On humus in coniferous woods: Europe (common in woods of Picea and Pinus),
U.S.A. (in woods of Tsuga, common in the north and west, rare in the south).1
Spores (5-)6-9(-10) X 3-4·5(-5)µ,, ochraceous or brownish ochraceous, finely
1 Cooke's record from Australia is doubtful (Handb. Austral. Fungi, p. 200).
-
612 RAMARIA
verruculose or merely rough (not echinulate), ovoid pip-shaped, or oblong, often
clumped together.
Basidia 30-50 x 6-7·5 µ,, narrowly clavate, multiguttulate: sterigmata (2-)4, 4-5µ
long: (basidia 35-40 x 7-8 µ,, Cott. et Wakef.: 18-34 X 4-5 µ,, Bourd. et Galz.:
20-35 X 4-5·5 µ,, Donk).
Hymenium not thickening: subhymenial hyphae 2·5-3·5µ, wide, as in R.Jormosa.
Hyphae 3-9 µ, wide, -11µ, in places, often
of uneven thickness, clamped, thin-walled,
but with very slightly thickened walls
(-o·5µ, thick) in the stem, cells 40-400µ,
long, often swollen at the clamps and
occasionally with a short clavate branch
-30 X 12µ,: the wider hyphae mainly
longitudinal and sparingly branched, the
narrower hyphae with frequent inter
weaving laterals and often surrounding
the wider hyphae, but monomitic.
Stem sterile, with narrow superficial
hyphae and a few projecting hyphal ends
. (excrescent to form the villosity at the
base): hymenium beginning abmptly at
TEXT-FIG. 261. Ramaria ochraceo-viren$: the stem-apex.
spores, X 1,000: clamps and a clavate branch Yellow colour in the cytoplasm of the
of the medullary hyphae, X 1, 000. subhymenial hyphae and basidia.
It is regrettable, as Donk remarks, that this unfamiliar, though descriptive, name
must be revived for the common fungus which has been widely known as Clavaria
abietina. Donk and Lundell have shown, however, that C. abietina Fi. (1821) must
undoubtedly be Clavaria Invalii Cott. et Wakef., though C. abietina Pers. (1794),
which supplied the specific epithet, is C. ochraceo-virens. Fries persistently intended
a non-virescent Clavaria for C. abietina, as shown by his remark in 1874, 'Forma
Persoonii minor, contrita virens' (Hym. Eur., p. 671): and the common Clavaria at
Upsala, in the Friesian localities, is C. lnvalii, according to Donk and Lundell.
However, there can also be no doubt that C. lnvalii and the virescent 'C. abietina'
were generally confused until Cotton and Wakefield described C. lnvalii (1919).'
From then until 1932, when Donk explained the true interpretation of C. abietina Fr.,
is but a small span in the history of mycolog¥, so that C. lnvalii appears as a stepping
stone. As Ramaria lnvalii it indicates the non-virescent species of coniferous woods
with echinulate spores. R. abietina Quel. was based on Clavaria abietina Pers. non
Fr. and, thus, the epithet 'abietina' cannot be used in Ramaria for C. abietina Fr.
Bourdot has proposed the conservation of C. abietina Fr. for the virescent species, but
this is clearly untenable according to the Rules of Nomenclature (Bull. Soc. Myc. Fr.
48, 1932, 207).
The ochraceous colour, small spores, tough flesh, and copious mycelium suggest
that R. ochraceo-virens is allied with R. stricta. But the toughness of the flesh is caused
by the abundant narrow interweaving hyphae in R. ochraceo-virens, not by the general
thickening of the hyphal walls as in R. stricta, and, so far as I have observed, the
hymenium does not thicken as it does in R. stricta. The green colour-change indicates
a cyanescence on a yellow background which is different from the purple-brown
colour change of R.jormosa and its allies. R. fiaccida, however, is very similar but not
virescent, and suggests that the virescence of R. ochraceo-virens is merely a specific
detail. But by this feature, its habitat, and squat form R. ochraceo-·virens is easily
1
Quelet (Vosges et Jura, 1872), Karsten (Hattsv. 1882), and Saccardo (Syll. 6, 1888)
interpreted C. abietina Fr. in the correct sense of C. lnvalii, thou�h Greville, Berkeley, Cooke,
l\1assee, and Quelet (1888) confused them.
RAMARIA 613
recognized. Old specimens, powdered with spores, however, may not show the
virescence, for it is a feature of active young tissue.
TABLE LXII. SPORE-SIZE IN RAMARIA OCHRACEO-VIRENS

8-10·6X3·8-5·3 Coker U.S.A. Utah (Miss· Burrows), dried plants greenish:
? a variety with large spores (Coker)
8-10X3·5-5 Donk Holland
7-10X(3-)4-5 Bourd., Galz. France
7-10X3-5 Cott., Wakef. Gt. Britain
7-<)X3·4-4·2 Coker Sweden I Specimen from Juel
6-<)X3-4·5 Konr., Maubl. France
7·5-8·5X4-4·5 Coker U.S.A. N. Carolina (Schw.)
6·2-8·5X3-3·7 " " N.Y., Coker and Burnham 110
7-8·2x4 " Sweden Specimen from Romell
6·6-8·5X3·7-4•4 " " Herb. Curtis, Friesian specimen det. c.
abietina
6-8·5X3·5-4·5 Corner Scotland
5-8X3·5-4·5 Coker Europe Herb. Persoon
7·5-7·8X3·7-4 " U.S.A. · Colorado (Clements)
7-7·5X3·5-4 " Canada leg. Macoun
5-'7·5X3·5-3·7 " U.S.A. Tennessee (Murrill)
6·5-'7X3·5-3·8 " Italy Herb. Bresadola
6·5-7X{3-)3"5-4 Lundell Sweden
6-7 X3-4 Rea Gt. Britain
7x3 ! Wehmeyer Canada echinulate
R. palmata (Pers.) Quel.

FI. Myc. 188, 467.
Basinym: Clavariapalmata Pers., Myc. Eur. 1, 1822, 171: Quel., Ass. Fr. 1880, 670.
Synonym: Clavariella palmata (Pers.) Karst., Hattsv. 2, 1882.
2-7 X 2-4 cm., caespitose, alutaceous, cream-or:hraceous, tan or tawny, becoming paler:
stem thin, much branched almost from the base: branches palmate, 2-3-fid and dilated
at the ramifications, thin, the tips acuminate, white: flesh pale, firm, smell sweet;
pleasant.
On the ground in coniferous woods, ? also in frondose woods: Europe.
Spores 6-7·5 X 4-4·5 µ. (Bourd. et Galz.), 6-7 X 3-4 µ. (Rea), 6-8 µ. long, elongate
ellipsoid (Quel.).
Basidia 27-32X4·5-6 µ..
Hyphae 3-9µ. wide, thin-walled, without clamps (Bourd. et Galz.).
var. guttaurensis Bres.

Sacc. FI. It. Crypt. Hym. 1916, 1242.
7 >'. 5 cm., lurid flesh colour: stem -3 cm. thick, cylindric or deformed, pale, soon
breaking up into strong subverticillate branches: branches and branchlets many times
dichotomous, smooth or rugulose, with arcuate axils, tips shortly 2-4-fid: flesh white:
smell and taste practically none.
On the ground under Castanea, in woods: N. Italy.
Spores 8-<) x 5 µ., lurid ochre-white, subamygdaliform (? smooth or rough).
Basidia 25-30 X 8 µ..
var. pluridentata Karst.

Symb. Myc. Fenn. 30, 1891, 62.
-4 cm. high, very much branched, yellowish white, drying alutaceous or
614 RAMARIA
pallid ochraceous: stem I cm.X 2-3 mm., thin: branches divaricate, verticillately
branched, convergent, smooth, mostly subcanaliculate, thickened and subcompressed
upwards, tips white, pluridentate.
In pine-woods: Esthonia (Mustjala).
Spores 6-'7X 3-4µ, ochraceous, smooth, ellipsoid.
This is C. palmata according to Quelet, Bourdot, Rea, and Donk. From R. gracilis
it differs in the palmate branching and the thin-walled hyphae, but palmate branching
is exceptional in Ramaria and the description leads one to suppose that it is some
polychotomous abnormality rnther than true flattened branching. Hence, I am inclined
to agree with Coker, that R. palmata is, at most, a form of R. gracilis.
Var. guattarensis may be a small specimen of R. suecica. Coker found that Bresadola
had mixed specimens of R. suecica, R. gracilis, and 'Clavaria abietina' under the name
C. palmata in his herbarium.
Var. pluridentata suggests R. gracilis, except for the smooth spores, which may be
an error.
R. polypus sp. nov.

Text-Figs. 6c, 262-5.
2-8 cm. high, solitary, gregarious or caespitose, pallid-white becoming tinged yellowish
or flesh colour, wholly pale yellowish drab, pale yellowish flesh colour, pallid dingy ochra
ceous or pallid cream-tan, the tips white or whitish: stem o-12X 1-3·5 mm., often
branched almost from the base, rarely elongate, -5 cm.X 3-8 mm. (in specimens attached
to underground pieces of wood), often flattened, slightly villous at the base, generally
with a white or cream-white floccose mycelial felt shortly spreading over the substratum
and with short rhizomorphs 0·2-0· 5 mm. wide: branches abundant, much branched
but laxly, curved ascending, 1-1·5 mm. wide and 3-4-chotomous below, dichotomous
above, more or less flattened, 2-3 mm. wide, below the axils, not palmately branched,
sometimes fastigiate, often conjoint, sometimes tubercled, tapering to the slender acute
tips 0·5 mm. wide: hymenium waxy, the sterile uppersides of the branches minutely
subvillous: flesh white, or pallid concolorous, unchanging, floccose-fibrous, pliant, some
what fleshy, waxy near the hymenium, slightly tough in the stem and main branches:
smell inparticular: taste ?
On rotten wood, rarely from buried wood, on dead palm trunks, or on rotten leaves,
in the forest: Malaya (frequent in the lowlands), New Guinea (Carr u715, Veiya,
Papua).
Spores 6-8(-g)X 4-5(-5·5)µ, cinnamon-buff to cinnamon-drab, pale pinkish cream
under the microscope when fresh, 1 but pale brownish ochraceous to very pale ochra
ceous after drying, rugulose-rough or subverruculose (without distinct warts), broadly
ellipsoid, blunt, aguttate, the wall slightly thickened.
Basidia 26-45 X 8-9µ, 35-50 X 7-9µ, or 50-60 X 8-9µ (? lengthening in the thicken
ing hymenium), clavafe: sterigmata 4, rarely 2-3, 3·5-4µ, 4-5 µ, or 5-6 µ long (accord
ing to the size of the basidium).
Hymenium thickening -250µ, often only slightly thickened -90 µ in some specimens,
absent from the uppersides of the branches, and from the lower 1-3 mm. of the stem:
subhymenium c. 20µ t,hick, composed of 2-4µ wide, thin-walled, hyphae.
Hyphae 2-8 µ wide, -11 µ in some collections, inflated -13 µ in places, clamped, with
distinctly thickened, smooth, colourless walls 0·5-1·5µ thick, cells 40-250µ long,
shorter-celled (20-50 µ) and thinner-walled next the subhymenium: H-connexions
frequent, the clamps sometimes inflated or irregularly formed: stetn be.coming covered
with a dense mat of 2-4µ wide mycelial hyphae.
Sterile hymenium -50 µ thick, consisting of a more or less continuous layer of hyphal
tips 3-5 µ wide, like sterile basidia, some of the tips with a narrow excrescent process
1
Possibly varying to pale ochraceous in some collections.
RAMARIA 615
-30 X2-3 µ and appearing as subventricose cystidia with a filiform apex, the super
ficial layer seated on a very loose 'subhymenium' of narrow hyphae.
Mycelial hyphae 1·5-3 µ wide, clamped, with thin or slightly thickened walls,
densely interwoven in the sterile mycelial felt.
TEXT-FIG. 262. Ramaria polypus, x 2.
Colour of the fruit-body caused by the oleaginous-granular contents of the basidia,

and by the colour of the spore-walls.
Spore-measurements:
6-8(-<J) X4-5(-5·5) I-" Singapore (Mandai Road, 17.5.31): spores pale cream
colour in the mass, but the fruit-bodies identical in all
other respects.
6·5-8 X5-5·5µ Carr II715 (Papua, 13.3.35, sea-level),
6-7·5 X4·5-5µ Singapore (Botanic Gardens, 18.11.34).
6·5-7·5 X4-4·5 I-" Pahang (Tembeling, 8.u,30).
6·5-7·5 X4-5µ Pahang (Tembeling 25.5.31).
6-7 X4-5µ Pahang (Tembeling 10.11.30: 16.11.30).
TEXT-FIG. 263. Ramaria polypus, x 2.
TEXT-FIG. 264. Ramaria polypus, X 2.

RAMARIA
This is a common species in Malaya and it may be identical with R. megalorhiza in
Ceylon, butt he spores of R. megalorhiza are said to have distinct warts. It is also like
R. subsigmoidea of S. America. The same aguttate rugulose spore occurs in R. flavo
alba of Malaya, which has terrestrial fruit-bodies and thin-walled hyphae. R. gracilis
appears macroscopically similar but it differs in its fragrance, 1-guttate spores, and
coniferous habitat.
TExT-FIG. 265. Ramaria pol:ypus: sterile hymenium and 'cystidia' on

the upperside of a branch, spores, and an unusual clamp on a thick
walled hypha: x 1,000.
Compare Lachnocladium reticulatum from Brazil, but with skeletal hyphae, though
very similar in form and habit.
R. pusilla (Pk.) comb. nov.

Basinym: Clavaria pusilla Pk., Buff. Soc. Nat. Sci. 1, 1873, 62: Rept. N.Y. St.
Mus. 25, 1873, 83: (non C. pusilla Pers. = Pistillaria pusilla: non C. pusilla Quel. =
Nyctalis).
Pseudonym: Clavaria decurrens Pers., sensu Coker, Clav. U.S. Can. 1923, 174,
t. 66, 88: (non Pers. = Ramaria crispula).
1·5-5 X 1 ·5-3 cm., small, slender, wholly dull cream-white or pale yellowish, often with
pinkish stains on the stem, or light olive-drab with pinkish purple stem, turning deep
purplish pink on bruising, drying ochraceous-yellow or drab: 0·5-1·5 Xo·15-0·7 cm.,
glabrous, toughish: branches rather few, open, lax, mostly flattened and angular, tips
small, subacute, spreading: flesh white, deep purplish pink on bruising, pliable, rather
tough: no smell or taste.
In humus under Pinus, Sequoia, and Tsuga: U.S.A. (N.Y., Oregon), Canada
(Ottawa).
Spores 4-6 X 2·3-3 · 5 µ., deep buff-yellow, verruculose, pip-shaped or ellipsoid,
aguttate.
Basidia. 4-5 µ. wide, 4-spored: hymenium 35-40 µ. thick, ? thickening (many included
spores): hyphac 1·5-8 µ. wide, clamped.
618 RAMARIA
var. australis Coker
Clav. U.S. Can. 1923, 177, t. 50, 67, 88 (ut Clavaria decurrens var. australis).
4-10 X2-8·5 cm., larger, but with slender branches, light buff-yellow, becoming
darker honey-yellow or isabelline from below upwards, reddening or flesh-pink where
bruised, drying deep brown: stem 1-2 X0·3-1 ·2 cm., rather stout, arising from a
fibrillose white mycelium: branches spreading, the side-branches turning upwards and
rather crowded, branchlets very numerous, slender, acute: flesh paler concolorous,
flesh-pink when cut or bruised, rather tough: taste bitter: smell none.
On the ground in frondose woods: U.S.A. (N. Car., N.Y.).
Spores 5-6·5 X2·5-3·5µ,, as in the type.
This species is readily distinguished by the small verruculose spores, reddening
flesh, and pallid white or yellowish rnlour. Coker referred it to Clavaria decurrens
Pers. on the evidence of the spores and general appearance of a collection in Persoon's
herbarium. He found the spores of this collection to be 4·5-6 X2·5-3·5µ,, pip-shaped
and minutely rough. Now C. decurrens was reduced to C. crispula by Fries, and so it
has been regarded ever since in Europe. The spores of R. crispula, according to
Bourdot and Galzin, are 5-7·5 X 3-4µ, and laxly asperulate or finely rugulose, and
the spores of a Persoonian specimen of R. crispula (collected near Paris) were found
by Coker to be 4·5-6·5 X 2·5-4µ,. Coker refers this Persoonian specimen also to
C. decurrens. It will be seen that in spore-size there is no difference between C.
decurrens of Persoon, C. crispula of Persoon, and C. crispula of Bourdot and Galzin:
and Karsten also gives the spore of C. crispula as 5-6 x 3µ,. Yet C. crispula macro
scopically is certainly different from C. decurrens sensu Coker, being an ochraceous
lignicolous fungus with unchanging flesh, and related, most probably, with R. stricta.
I think too much reliance cannot be placed on the size of the spores as a means of
identifying the small-spored species of Ramaria, and when, on such grounds, Coker
suggests Ramaria curta may be the same as his C. decurrens, he is undoubtedly
mistaken. Moreover, C. decurrens sensu Coker has not been found in Europe. There
fore, I have used Clavaria pusilla Pk. as the basinym of this species, and I have
followed Fries in reducing C. decurrens to C. crispula. See also C. subdecurrens.
The spore-measurements given for R. pusilla are these:
4·5-5·5 X2·5-3µ, Clavaria pusilla, type: roughish.
4·5-6 X2·3-3µ, Coker and Burnham, 106, 120: verruculose.
4-5·5 X3-3·5µ, Macoun, Canada.
4-4·5 X2-2·5µ, Oregon, echinulate (Doty).
5-6·5 X2·5-3µ, var. australis, type, No. 2769: verruculose.
6-6·5 X3-3·5 µ, var. australis. Burnham 4: roughish.
R. rufescens (Fr.) comb. nov.

Basinym: Clavaria rufescens Fr., Epicr. 1838, 74: Bres., le. Myc., t. 1084: (non
Coker= R. holorubella): Schaeff., le. 1770, t. 288.
6-15 X 8-12 cm., massive, whitish then pale alutaceous or pale golden-ochraceous
with ru/ous or obscurely pinkish tips: stem 8 X3-3·5 cm., massive, columnar, pallid:
branches polychotomous below, short and massive, crowded, much branched, the
internodes somewhat elongate towards the very numerous, congested, cauliflower-like,
dichotomous tips: flesh white, unchanging, watery-marbled, compact, slightly tough:
taste subacid: edible, but old specimens purgative.
On the ground in woods: Europe (N. Italy).
Spores 8-10 X3·5-4µ,, yellow, smooth, oblong-amygdaloid.
Basidia 40-45 X6-8µ,, 4-spored.
var. frondosarum Bres.
Hedw. 24, 1885, 148.
Stem white: tips not red.
RAMARIA 619
The use of this name is uncertain. Cotton and Wakefield and other European
mycologists have considered C. rufescens to be a form of C. botrytis with rufescent,
instead of purple, tips. Maire united it with C. pallida ( = R. Mairei), and then with
R. botrytis. Quelet regarded it as a variety of R. aurea. Rea considered it a distinct,
rare species, 'white becoming yellow upward, the branchlets all blood red, becoming
paler: flesh white, reddish at the apex of the branches: spores ochraceous', and indicated
that he had seen a living specimen in England. Bourdot and Galzin did not recognize
it in the French flora. Bresadola, whose description I have given, seems to be the only
European mycologist who has proved that there i� a European fungus agreeing with
the original description of C. rufescens, yet distinct from C. botrytis (in the colour of
the branch-tips and the size of the spores). Huber has also acknowledged Bresadola's
species (Zeitschr. f. Pilzk. 1934, 34). It remains to rediscover this fungus in Europe
and to compare its spores with those of R. holorubella of North America. Rea's fungus
was probably the same, but he gave no spore-measurements. The description given
by Fries (Hym. Eur. 670) is: 'trunco crasso, elastico, alutaceo, ramosissimo, ramis
multifidis, stipatis, subfastigiatis, laevibus alutaceis, apice rubris. Schaeff. t. 288': and
he indicated that he had seen a living specimen. It is generally assumed that Schaeffer's
picture is the 'type', though it seems possible that Fries based his description on a
living specimen. The description agrees with Bresadola's, though his painting does
not show truly red tips to the branches.
Coker identified with C. rufescens a N. American fungus which is undoubtedly
similar macroscopically to R. rufescens sensu Bres., but it has striate spores as R.
botrytis (of European authors, not of Coker), though shorter. I have kept this species
apart as R. holorubella. It is common in the United States and it is not impossible
that it occurs in Europe, though modern European mycologists have not discovered it:
therefore, I think it is unlikely to have been C. rufescens of Fries or Schaeffer.
Bresadola considered C. botrytis sensu Coker the same as C. rufescens sensu Bres.
It is possible, but as I have explained under R. botrytis, it is preferable to keep them
apart at present to avoid possible confusion. Coker emphasizes the rose-pink tips of
his C. botrytis ( = R. botrytoides).
var. frondosamm may be R. fiavo-brunnescens.
R. rufo-violacea (Baria) Quel.

FI. Myc. 1888.
Basinym: Clavaria rufo-violacea Baria, Champ. Nice, 1859, t. 42, f. 3-13.
Small or medium-size: stem thin, cylindric, white at the base: branches lax, simple
or forked, obtuse, amethyst with rusty yellow apices: fragile.
Spores ferruginous, oblong.
Southern France.
In colour this comes between R. fennica and R. fumigata, but differs from both in
its slender, sparingly branched form. Compare R. testaceo-violacea.
R. Saccardoi (Syd.) comb. nov.

Basinym: Clavaria Saccardci Syd., Syll. Fung. 14, 1899, 238.
Synonym: Clavaria dichotoma Kalchbr., Grev. 10, 1882, 105: (non Godey = Clavu
linopsis dichotoma).
Clavaria Kalchbrenneri Sacc., Syll. Fung. 6, 1888, 710: (non Muell. = Ramariopsis
crocea).
Branched, pale then rufescent or ferruginous: branches thick, repeatedly forked,
tips slender, forked or fasciculate, acute.
In humus: S. Africa (Boschberg).
Said to be near R. gracilis and R. stricta.
620 RAMARIA
R. sanguinea (Coker) comb. nov.
Basinym: Clavaria sanguinea Coker ex Doty, Clav. Pac. N.W. 1944, 52: Coker,
Clav. U.S. Can. 1923, 118, t. 35, t. 38 (ut C. sanguinea Pers. 1799 = Ramaria fiava):
Fawcett, Proc. Roy. Soc. Viet. 51, 1939, 278, t. 21.
Synonym: ? Clavaria umbraticola Leuba, Champ. Com. 77, t. 42.
6-8 ·5 X4· 3--7 ·5 cm., wholly pallid cream-white with the tips tinged yellowish, becoming
deeper yellow all over, or light egg-yellow in the upper half and deeper-coloured towards
the tips, the branches sometimes tinged pinkish yellow, all parts turning blood-red, brownish
red, brick-red or vinaceous when bruised, the whole fruit-body becoming dull reddish
brown with age: stem 1·5-2 X1-1·8 cm., rooting, shortly expanding into 2 to 5 main
branches: branches numerous, erect, rather crowded, blunt, the fruit-body rather fiat
topped and cauliflower-like: flesh concolorous, reddening only at the surface: smell
none or faintly of aniseed.
On the ground in woods, frondose and coniferous: U.S.A. (N. Car., Mass., Oregon,
rare): Australia (Kinglake, Victoria).
Spores 6·5-12 X3-47 µ,, ochraceous, minutely rough or smooth, with several guttae.
Basidia 7 µ, wide, 4-spored.
Coker used Persoori's invalid name for this species. European authors are agreed
that C. sanguinea Pers. (Obs. 1799, t. 3, f. 5) is R. fiava. Whether Coker's species is
also R. fiava in the European sense (non sensu Coker = R. fiavo-brunnescens), I am
uncertain, for its spores are distinctly shorter and the fruit-bodies appear to be pinker
or redder. However, the spores may well be as variable as in R. formosa, with the
shorter spores almost or quite smooth, and it has not been ascertained whether the
redder colour of R. sanguinea (at least, when old) is a true pigmentation or merely a
general rufescence as a result of weathering. It is certainly remarkable that the
European R. fiava has not been recorded from N. America or S. Australia, while
R. sanguinea of both these continents has not been found in Europe. Fawcett's
description of R. sanguinea from Australia agrees closely, even in spore-size, with
R. fiava as described by Bresadola and Konrad and Maublanc. Doty, however, gives
the colour of specimens of R. sanguinea from Oregon as 'pallid yellow to maize-yellow
or pink throughout'.
Clavaria umbraticola answers well to old, weathered specimens of R. sanguinea. It
was described as wholly reddish rufescent, much branched from the base: flesh white,
then rufescent; in mountain fir-woods, Switzerland. One must await fuller study of
fresh specimens of R. fiava in Europe to know whetht;r such is its end also.
Fawcett and Doty find the spores larger than given by Coker and finely rough, thus
bridging the gap to R. fiava. The published data are:
Coker (N. Car. 2656) 7·5-8·5 X3-3·8 µ,, smooth.
,, (N. Car. 4394) 6·2-8·2 X3·7-4·5µ,, smooth.
,, (Mass., Murr. et Thomps.) 7·8-9·5 X3·5-4µ,, smooth or slightly rough.
Doty (Oregon) (9-)10·5-12 X3·5-4·7µ,, slightly rough.
Fawcett (Victoria) 9-11 X3-4·5 µ,, slightly rough.
R. secunda (B.) comb. nov.

Basinym: Clavaria secunda B., Grev. 2, 1873, 7: Coker, Clav. U.S. Can. 1923, 150,
t. 57, 85: Wehm., Pap. Mich. Ac. Sci. Arts Lett. 20, 1935, 262: Doty, Clav. Pac. N.W.
1944, 58.
Synonym: Clavaria crassipes Pk., Bull. N.Y. St. Mus. 67, 1903, 27 (fide Coker).
4-_12 cm. tall, cream-white to pale tan tinged pale pink or flesh colour (light ochraceous
salmon to buff-pink), tips concolorous or paler or pinkish, often turning reddish brown
or cinereous-vinaceous on bruising or with age (Wehm.): stem 2-3·5 cm. long, -4 cm.
wide, stout, short, often with many abortive branches, soft, glabrous, white: branches
numerous, rather lax, axils round, smooth, tips blunt or subacute: flesh white, fibrous,
RAMARIA 621
soft, spongy, not brittle: taste mild, sweetish: smell none, or sweetish (Doty, Wehm.),
pleasant and sweetish, of old ham, or rancid (Coker).
On the ground in frondose and coniferous woods: U.S.A. (N.Y., N. Car., Oregon),
Canada (Nov. Scot.).
Spores 8-12·5 x4-5·5µ, ochraceous, smooth or minutely rough, ellipsoid.
Basidia 35-45 X 7-8µ: hymenium evidently thickening, 60-100µ (Doty).
This is said by Coker and Wehmeyer to be like R. obtusissima but without its yellow
tints and with shorter, broader spores (though there is considerable overlapping). On
the other hand, it seems near to R. divaricata and R. subbotrytis both in colour and
spores. The colour change noted by Wehmeyer is that of R. formosa.
Wehmeyer found a collection (No. 1116) in Nova Scotia which had smaller spores,
7-8·5 X 3·5-4·5µ, and smaller fruit-bodies, 4-7 cm. high. He referred it to R. secunda,
and suggested it represented well-grown R. verna, which Coker might have described
only from young plants.
TABLE LXIII. SPORE-SIZE FOR RAMARIA SECUNDA
Spore-size in µ. I Authority Remarks

n·5-12·5X 5-5·5 Coker Peck (1917): as C. crassipes
l0-l2X4-5·5 C. crassipes (type)·
10-12·2X 4·5-5·5 ,, C. secunda (co-type in Herb. Curtis)
10-12X4-4· 5 Doty Oregon
8·5-11·5X4-4·5 Coker No. 2876: mostly 9·3X4·3 µ.
8·5-11X4-4· 5 Peck (1916): as C. crassipes
8-11X3·8-5·3 ,, Burnham 112: slightly rough
8·5-10X4-5 Wehmeyer Nova Scotia
7-8·5X3·5-4·5 No. 1II6, Nova Scotia:?= C. verna
R. sinapicolor (Clel.) comb. nov.

Basinym: Clavaria sinapicolor Cleland, Tr. Roy. Soc. S. Austral. 55, 1931, 159:
Toadst. S. Austral. 2, 1935, 264, f. 58: Fawcett, Proc. Roy. Soc. Viet. 51, 1939, 272,
t. 21, text-fig. r.
-12 X 9 cm. caespitose, rarely subsolitary, much branched, light yellow, mustard- or
straw-yellow, or light orange-yellow, brighter yellow towards the tips (Clel.): stems
rather slender, densely fasciculate into a compound trunk, branched from the ground.
with the primary branches much elongate: branches rather elongate, fastigiate, axils
rounded and somewhat flattened, often longitudinally rugulose from the axils: flesh
white or pale cream, solid: taste mild, smell sweet (of U)"!:.:-flower.,), strong with age.
On the ground (in Eucalyptus-forest, Clel.): S. Austral. (N.S.W., Victoria,
abundant).
Spores 6-10 X 3-4·5 µ (Fawcett), 5·5-8(-10) X 3·8-4·5(-5)µ (Clel.), yellowish to
ochraceous, ellipsoid or subpyriform, rough or nearly smooth (Clel.).
This is very near R. fiavo-brunnescens but differs in habit, having fasciculate slender
stems in place of a massive trunk. Thus it resembles R. conjunctipes, which stands in
the same relation to I-<. formosa. They are the nearest approach in the genus to the
simple, caespitose fruit-bodies of Clavaria and Clavulinopsis.
If R. sinapicolor were suffused with pink, it would be identical with R. conjunctipes,
and it is noteworthy that Coker has described a fragrant variety of R. conjunctipes and
Doty records a clear yellow collection. Compare also R. cystidiophora.
R. soluta (Karst.) comb. nov.

Basinym: Clavaria soluta Karst., Med. Soc. Faun. et Flor. Fenn. 5, 1879, 30.
Synonyms: Clavaria crispula subsp. soluta Karst., Finl. Nat. o. Folk. 37, -1882, 189·.
RAMARIA
Clavariella soluta Karst., Finl. Nat. o. Folk. 48, 1889, 390: Acta. Soc. Fenn. 18,
1891, 109, t. 78.
3-4 cm. high, whitish then reddish brown, with decumbent, branching, fil:irillose
rhizomorphs; longitudinally rugulose: stem narrow: branches few, projecting, divari
cate, with subulate tips.
Investing roots and cones of Picea excelsa: Finland.
Spores 2µ. wide, ochraceous, round, smooth.
This is doubtful. The cotype (Karsten 1308) in the Kew herbarium is poorly
developed Lentaria byssiseda with typical spores (14-18 X 4 µ.) and slightly thick-walled,
clamped hyphae.
R. spinulosa (Fr.) Quel.
FI. Myc. 1888, 468.
Basinym: Clavaria spinulosa Fr., Syst. Myc. 1, 1821, 468.
Synonym: Clavariella spinulosa (Fr.) Karst., Hattsv. 2, 1882.
5-8 X 3-6 cm., somewhat cinnamon or brownish umber, tips concolorous: stem 2-4 X
2-3 cm., rather massive, pallid then yellowish, much branched: branches elongate,
crowded, tense, straight, subulate, spinulose: flesh white, firm, tough.
On the ground in woods and heaths: Europe (Sweden, France, Gt. Britain, rare).
Spores 8--<J X 4-5µ. (Rea), 10-12 X 5-6·5 µ. (Bourd. et Galz.), ochraceous, (? smooth),
ellipsoid or obovoid, incurved.
This appears to be a rare species. I have given the description of it as understood
by Quelet, Bresadola, Rea, and Bourdot and Galzin. Lundell has recently ascribed a
Swedish collection to it, but his fungus seems to me identical with R. condensata, as
I have mentioned on page 566. Fries emphasized the concolorous tips of C. spinulosa
and this feature is repeated by Quelet and Rea, so that it would appear characteristic
and not merely the effect of ageing on a fruit-body with yellow tips when young. In
form and colour R. spinulosa resembles R. Invalii, but it differs in its smooth spores
and, possibly, it does not grow in coniferous woods. Its colouring and spores
conform with those of R. Strasseri.
R. Strasseri (Bres.) comb. nov.
Basinym: Clavaria Strasseri Bres., Strasser Pilzfl. Sonntagsberg 2, 1900, 3: le. Myc.,
t. 1089: Coker, Clav. U.S. Can. 1923, 149, t. 87.
7-13 X 4-12 cm., pale whitish alutaceous or cream-tan, then pale brownish alutaceous,
finally cinnamon-umber, tips concolorous: stem 2-5 x 3-6 cm., obovate, massive, with
several spreading branches at the top, white, or stained tan: branches strong, com
pressed, polychotomous then dichotomous, rather flexuous and lax, with arcuate axils,
tips obtuse or subacute: flesh white, soft, fibrous, not very brittle, watery-marbled:
taste rather bitter: smell faint, of old ham (Coker): edible.
TABLE LXIV. SPORE-DATA FOR RAMARIA STRASSER[

- ·- --
12-16 X4-6 Bresadola Italy, Austria
n·8-15·5 X5-6·6 Coker " Herb. Bres.
13-15 X3·8-5 " " "
I
14-18·5 X4·8-6·7 ,, N. Carolina No. 2897
14-17 X5·5-6·3 ,, " No. 2927
On the ground in coniferous or frondose woods: Europe, tN. Italy, Austria, rare):
U.S.A. (N. Car.).
Spores 12-16 X 4-6µ., ochraceous (Bres.), 14-18·5 X 5-7 µ. yellow-buff (Coker):
smooth, amygdaloid-oblong, guttulate.
Basidia 60-70 X 8-10µ., 4-spored.
RAMARIA 623
Hymenium 90-110µ thick (? including subhymenium): hyphae 4-10µ, clamped
(Coker).
This species is distinguished by the pale brownish alutaceous colour, rather ungainly
form, large smooth spores, and unchanging white flesh. It seems to be related to
R. cacao, R. obtusissima, R. condensata, and R. spinulosa, having the brownish pigment
of the last two without any yellow, and with larger spores and fruit-bodies. The
American specimens have slightly larger spores than the European.
R. stricta (Fr.) Quel.
FI. Myc. 1888, 464.
Basinym: Clavaria stricta Fr., Syst. Myc. 1, 1821, 468: Konr. et Maubl., Jc. Se!.
Fung., t. 487: (non Bres., non Bourd. et Galz. = Ramaria Bourdotiana).
Synonyms: Clavaria Syringarum Pers., Myc. Eur. 1, 1822, 164 (teste Lundell).
Clavaria pruinella Ces., Rabenh. Fungi Eur. Exs. Cent. 5, 1861, No. 14: Bot. Zeit.
1862, 198: Sacc. Sy!!. 21, 1912, 427: ibid. 23, 1925, 489.
Clavaria kewensis Mass., Journ. Bot. 34, 1896, 153: (fide Cott. et Wakef., Coker).
Clavaria stricta var. condensata Lundell, Sv. Svamp. 5-6, 1936, 26.
(C. dendroidea et C. condensata sensu Bourd. et Galz.).
Clavariella stricta (Fr.) Karst., Hattsv. 2, 1882, 184.
4-10 X 3-8 cm., often caespitose, pallid-yellow or alutaceous to ochraceous, sometimes
tinged flesh colour, becoming brownish ochraceous or fuscous-cinnamon, tips clear yellow
( concolorous with age), all parts becoming fuscous or vinous on bruising: stem 1-6 cm. X
3-10 mm., distinct, pale, arising from a white mycelial felt or thread-like rhizomorphs:
branches numerous, much branched, dichotomous, erect, fastigiate, elongate, tense,
straight, subparallel, acute: flesh white or yellowish, tough, pliant, the hymenium waxy:
taste bitter or slightly peppery: smell faint or, often, pleasant, somewhat aromatic or
of aniseed.
On dead wood, stumps, trunks, branches of frondose and coniferous trees: Europe
(common), Japan. 1
Spores 7-10 x 4-5µ, rusty ochraceous or cinnamon-ochraceous, minutely rough or
almost smooth (without distinct warts), oblong or pip-shaped.
Basidia 30-40 X 7-9µ: sterigmata (2-)4.
Hymenium thickening to 100 µ, at least, absent from the uppersides of the branches:
subhymenium composed of 2-3 · 5 µ-wide, thin-walled hyphae.
Hyphae 3-10µ wide, clamped, cells 30-200µ long, the walls more or less stro1l/dY
thickened (0·5-2 µ), the thin-walled hyphae more branched than the thick-walled, often
of uneven width, the thick-walled hyphae often kinked, strictly longitudinal near the
subhymenium, looser, thicker-walled, and often much entangled in the central part.
Mycelial hyphae 2-4µ wide, with slightly thickened walls; in the mycelial felt at the
base of the stem 1citlt many narrow, thick-walled, unbranched, and aseptate hyphae,
1 · 5-3µ wide (walls o· 5-1µ thick), like skeletal hyphae.
var. alba Cott. et Wakef.

Tr. Br. Myc. Soc. 6, 1918, 174.
Wholly cream-white: spores typical.
On the ground among fallen leaves, twigs, &c.: Scotland (Drumnadrochit).
var. concolor var. nov.
Synonyms: Clavaria stricta sensu Coker, Clav. U.S. Can. 1923, 162, t. 61, 62, 88:
Wehm., Pap. Mich. Ac. Sci. Arts Lett. 20, 1935, 264: Teng, Sinensia 7, 1936, 256:
Fawcett, Proc. Roy. Soc. Viet. 51, 1939, 18, t. 5: Doty, Clav. Pac. N.W. 1944, 48.
1
The record from New Caledonia needs confirmation: that from Ceylon is R. megalorhiza
according to Petch.
RAMARIA
C. leucotephra B. et C., Grev. 2, 1873, 7: (fide Coker).
? Lachnocladium Atkinsonii Bres., Journ. Myc. 8, 1902, II9: (fide Coker, forsan
R. apiculata).
C. densissima Pk., Bull. Torr. Bot. Cl. 30, 1903, 98: (fide Coker).
Lachnocladium odoratum Atk., Ann. Myc. 6, 1908, 58: (fide Coker).
? Lachnocladium erectum Burt, Ann. Mo. Bot. Gdn. 6, 1919, 276, t. 5.
Pale fleshy tan, cinnamon-brown at maturity, deeper brown or reddish brown on
bruising (Coker): light buff to pinkish buff, vinaceous where bruised (Wehm.): pinkish
buff to light ochraceous-buff (Teng): isabelline, honey-yellow, or ochraceous-buff
(Fawcett): tips concolorous, or creamy (Fawcett): smell often fragrant, of aniseed, or
strong and of radishes.
On wood of frondose trees (also coniferous, Doty): U.S.A., Canada, China, S.
Austral.
Spores 6-10· 5X3· 5-5µ,, cinnamon-buff, or ochraceous (Doty, Fawcett), finely
rough or nearly smooth, distinctly rough (Fawcett).
Hymenium thickening: hyp hae thick-walled.
var. fumida Pk.

Rept. N.Y. St. Mus. 41, 1888, 86.
Dingy fuliginous-brown, tips concolorous.
U.S.A.
var. violaceo-tincta Bourd. et Galz.

Hym. Fr. 1928, 98 (ut C. condensata var. violaceo-tincta).
Stem tinged violet: hyphae 2-3 (-6) µ, wide: spores 7-9 X4-5µ,.
On a stump of Alnus: France (rare).
This is the common lignicolous Clavaria of Europe, which forms beautiful waxy
looking masses round old stumps. Karsten, Cotton and Wakefield, Rea, Konrad and
Maublanc, Donk, and Lundell agree in their interpretation of C. stricta, the important
points being the strict, subparallel branches with yellow tips, the discoloration on
bruising, the tough flesh, the minutely rough spores, and the lignicolous habit: to these
I would add the thick-walled hyphae, the dimitic construction of the mycelial felt
round the base of the stem, and the thickening hymenium.
::. stricta sensu Bourd. et Galz., and Bresadola, is a different species with smaller
spores, namely R. Bourdotiana.
The peculiarity of the N. American, Chinese, and S. Australian fungi, described as
C. stricta, is the absence of the clear yellow tips which distinguish the European
species, For this reason I have made var. concolor, the tips of which are either con
colorous or pallid whitish or cream. According to Coker, var. concolor grows only on
frondose wood, but Doty finds it on both frondose and coniferous wood, just as R.
stricta in Europe. Thus the main distinction between C. stricta and C. apiculata
drawn by Coker disappears, and I am not at all sure that var. concolor is different
from R. apiculata, unless there is no vinescence on bruising in R. apiculata.
Fawcett's description of C. stricta from S. Australia suggests a more ochraceous
yellow fungus with rougher spores than the N. American, and there is no mention of
any discoloration on bruising. Several tropical species resemble it, but they are, as
yet, too little known for certain identification.
As mentioned under R. condensata, C. stricta var. condensata Lundell is probably
R. stricta and not R. condensata. The cotype of C. pruinella in the Kew herbarium is
typically R. stricta, with finely rough, pale ochraceous spores 8-10X4-5µ,.
The spore-data for var. concolor are these:
6 -10·5X3·5-5µ, (Coker), 7-8·5X3-4µ, (Wehm.), 7-8·5X3·5-4µ, (Teng), 9-10·5X
4-5µ, (Doty), 6·3- 9X3·5-4·5µ, (Fawcett).
RAMARIA 625
R. subbotrytis (Coker) comb. nov.
Basinym: Clavaria subbotrytis Coker, Clav. U.S. Can. 1923, 116, t. 28, 33, 85:
Kauffm., Pap. Mich. Ac. Sci. Arts Lett. 8, 1928, 150: Doty, Clav. Pac. N.W. 1944, 55.
TEXT-FIG. 266. Ramaria stricta: medullary hyphae of the fruit-body and, on the right, skeletal
and generative hyphae from the mycelial subiculum: X 1,000.
4-10 x 5·5-9 cm., wholly coral-pink when young,fading cream-ochraceous in the lower
part (begonia-rose to geranium-pink of Ridgway, fading in wind or sun to rose
incarnate or salmon-pink: Kauffm.): stem 1-3 cm. high and thick, tapered downwards,
clean, glabrous, persistently pink at the base, or white: branches 5-10 mm. thick and
polychotomous below, 2-4 mm. thick and dichotomous above, numerous, fairly com
pact, upright or slightly divaricately curved, tips blunt 1-1·5 mm. wide: flesh con
colorous and remaining pink longer than the surface, unchanging, rather brittle, fragile,
not chalky when dry: smell and taste slight (as in R. botrytoides) or none (Kauffm.).
In humus in woods (Quercus and coniferous), rather rare: U.S.A. (N. Car., Pa.,
Mich., Wash., Md., Oregon).
Spores 7-9· 5 X 3-3 ·7 µ, cinnamon-ochraceous, pinkish in thick deposit (Coker),
5119 S S
626 RAMARIA
pinkish buff in the mass and tinged pink under the microscope (Kauffm.), ellipsoid;
finely and rather sparsely verrucose (Coker), smooth (Kauffm.).
Basidia 6·5-9· 5 µ. wide, 4-spored.
Hymenium 55-65 µ. thick: cystidia none (Kauffm.).
Hyphae without clamps (Coker).
var. intermedia Coker
J.c. l 17, t. 34, 85.
Stem rather massive, with abortive branches, often rugulose.
Spores 8·5-II x 3·5-4· 5 µ., cinnamon-ochraceous, nearly smooth.
From R. formosa this species seems to differ primarily in the texture of the flesh,
which does not dry chalky. Secondarily, its colour is more intensely pink (the pigment
pervading the flesh), the tips are not yellow, the flesh does not change colour, and the
spores are browner, narrower, and smoother. But I write secondarily because none of
these characters seems decisive, especially when var. intermedia is considered. Thus,
for the colour, Coker says some collections are delicate pinkish salmon all over, with
tints of light ochraceous in maturing parts, and for another he gives 'pink all over
except for the creamy tips': for No. 2824 (var. intermedia) he gives the tips pale lemon
yellow and for No. 2825 (var. intermedia) the tips only slightly lighter than the flesh.
In other words, it seems that the yellow colour may not be entirely absent. Regarding
the absence of colour-change of the flesh, R. subbotrytis agrees with the American
specimens of R. formosa but not with the European. And regarding the spores, in
Nos. 2847 and 3055 (see Table LXV) they are the same·as those of R.formosa in size and
marking, while those of No. 4316 are the same in colour: moreover, some American
collections of R.formosa have spores as small as those of R. subbotrytis (see Table LXV).
The pink form of R. obtusissima is also extremely close and seems to differ main!� in
the longer, smooth spores.
One must conclude that R. subbotrytis covers several spt:c1es, or varieties, in the
Formosa-group which differ from the yellow-pink R. formosa-series in the deficient
yellow colour, the smoother, narrower spores, and in the flesh not being vinescent or
drying chalky, viz. R. botrytoides, R. conjuncta, R. divaricata, R. secunda, R. subbotrytis,
and, perhaps, R. testaceo-violacea.
TABLE LXV. SPORE-DATA FOR RAMARIA SUBBOTRYTIS
Spore-size in µ, Authority Remarks

7·5-9 X 3-3·7 Coker No. 4679, nearly smooth
7-9·3X3·3-3,7 No. 2621, minutely rough
7·7-9·3X3·3-3·7 " No. 3297
7-9(-10)X3-3'"5 Kauffmann smooth
var. intermedia
9·5-11X3·5-4·5 Coker No. 2847: type: nearly smooth
9·7-1 l X 3·8-4 No. 112, nearly smooth
8·5-II X3·7-5 No. 3055, slightly rough
8·5-11X3·8-4 No. 2853, minutely rough
9-10·5 X 3·7-4·2 No. 2824 (tips pale lemon yellow)
8·5-10·5X3·5-4 No. 2825 (tips slightly paler than flesh)
8·3-9·5X3"3-3·7 No. 781, smooth
7-9·3X2·8-3·7 No. 4316, spores light ochraceous, tinged salmon
R. subdecurrens (Coker) comb. nov.

Basinym: Clavaria subdecurrens Coker, Clav. U.S. Can. 1923, 172, t. 65, 89: Doty,
Clav. Pac. N.W. 1944, 43.
RAMARIA
-3·5 X2·5 cm., pale cream-ochraceous then somewhat deeper, but pallid, ochraceous,
tips persistently lavender (pale to deep pinkish lavender): stern short or none: branches
compact, cylindric, often flattened and angled, tips acute, many, often cristate: flesh
pliable, tender, not changing colour: taste acid or acid-bitter: smell none.
In humus under Tsuga and Abies: U.S.A. (N.Y., Oregon, Wash., rare).
Spores 5·5-7·7 X2·5-3·3µ, deep ochraceous, verruculose, narrowly ellipsoid:
8--<J·5 x4·5µ, rough (Doty).
Basidia 4·5-5µ wide, 4-spored: -40µ long (Doty).
Hymenium 50-55 µ thick, thickening slightly.
Coker says this is near R. flaccida and R. pusilla and that on each of the two occasions
when he found it, it was growing near, but not mixed with, R. flaccida. It differs from
both in the colour, especially of the tips, and in the narrow spores. The colour recalls
that of R. Mairei, R. subspinulosa, and R. cacao, with which species i.t may be allied.
R. subspinulosa, also, has a thickening hymenium.
The spore-measurements given by Coker are these:
6·5-77 X3-3·3µ New York, Burnham 81.
6-7x2·5-3µ ,, Coker & Burnham 101 (type).
5·5-6-3 X2·5-3µ House 1447
R. subfennica (Coker) comb. nov.
Basinym: Clavaria subfennica Coker, Journ. El. Mitch. Sci. Soc. 63, 1947, 45, pl. II
(upper figure), pl. 12 (figs. 5, 6).
-17 cm. high overall, the rooting stem-like base 3·5-6·5 cm. long, deep dull brown
(argus brown, Ridg.), the tips concolorous, the rooting base white becoming sordid
when handled: branches erect, fastigiate, divided four to six times: flesh soft, rather
flexible, cracking when bent to 45 °, straw-colour: smell none, taste inparticular.
In humus in deciduous woods: U.S.A. (N. Car., Fodderstack Mountain, Coker
12879, type: Coker 13378).
Spores 8·5-11 X3·8-4·5µ, chrome yellow (light orange yellow, Ridg.), minutely
verrucose, rather oblong ellipsoid.
Basidia 7-10 µ wide, 4-spored.
Hymenium thickening?
According to Coker, this has the spores of R. fennica, but the habit and colour of
R. longicaulis. It seems to me very close to R. Invalii which, however, is a species of
coniferous woods. Compare R. flaccida and R. eumorpha.
R. subgelatinosa sp. nov.
-15 cm. high, solitary or 3-4 tufted, beautifully apricot-pink with horn-yelloiv tips:
stem -4x1 cm., abrupt, buff-yellow, w,hite at the base: branches 4-8 mm. thick, and
polychotomous below, cylindric, dichotomous upwards and gradually narrowed to the
blunt, simple or bifid tips, c. 1 mm. wide, somewhat fastigiate, smooth or longitudinally
rugulose below the branchings, especially towards the tips, the outer branches rather
spreading: flesh paler concolorous or pallid-white, not discolouring, slightly but firmly
gelatinous, rather firm, not brittle, drying cartilaginous and membranous horny.
On the ground in the forest: Malaya (Pahang, Kuala Teku, 9.9.37, lowland).
Spores 8·5-10·5 X 4·5-5µ, yellow-brown under the microscope, verruculose-rough,
subcylindric-ellipsoid, 1-3 guttate or multiguttulate.
Basidia 50-70 X8-11µ, clavate, densely granular-guttulate: sterigmata (2-)4,
5-7 fl, long, straight.
Hymenium 70µ thick, not thickening or only very slightly; absent from the upper
sides of the inclined branches or as a short sterile rudimentary hymenium: sub
hymenium c. 20µ thick, composed of 2-4µ wide, interwoven, uninflated hyphae.
Hyphae 3-12µ wide, cells 30-120µ long, shorter (20-50µ long) near the
628 RAMARIA
subhymenium, clamped, with slightly thickened submucilaginous walls (not watery
gelatinous), longitudinal and entwined without interweaving hyphae: on the surface
of the stem 3-5 µ, wide, with a few shortly projecting hyphal ends.
TEXT-FIG. 267. Ramaria subgelatinosa, X 1.

Hyphal tips in the growing-points 2-3·5 µ, wide, the subterminal cells 15-25 µ, long,
on delimitation.
This species differs from R. formosa in its subgelatinous flesh, without colour
change, and in its small spores. It thus connects R. formosa with R. gelatinosa, as
I have explained under that species. R. subgelatinosa shows what important mycological
discoveries are yet to be made in the big forested areas of the tropics, and how difficult
they may be to find, for I have seen it only once in thirteen years.
R. subsigmoidea (Sacc. et Syd.) comb. nov.

Basinym: Clavaria subsigmoidea Sacc. et Syd., Sy!!. Fung. 16, 1902, 204.
Synonyms: Clavaria pampaeana Speg., Fungi Arg. Nov. 1899, 182, n. 279: (non
Speg. 1881 = Clavulina pampaeana).
RAMARIA 629
Clavaria pamparum Speg., Ann. Mus. Nat. Buenos Aires 19, 1909, 280.
2-3 x 1-5 cm., caespitose, dingy waxy-yellow or greyish yellow, delicate, somewhat
fleshy: stems fasciculate, solid, arising from a fibrillose-tubercular white mycelium:
branches 1-2 cm. long, solid, much crowded, often and repeatedly sigmoid-contorted,
often very shortly tubercular-subramose and anastomosing, tips entire or shortly bifurcate,
rather obtuse.
On the ground among roots of Carduus: Argentine (Arrecifes, Maj.: near La Plata).
Spores 6-7 X 4-4·5 µ,, ochraceous, laxly rugulose, contents somewhat tawny,
aguttate, or with 1-2 guttulae.
The small size and rugulose, aguttate spores suggest affinity with R. polypus of the
Stricta-group and R. fiavo-afba of Decolorans.
R. subspinulosa (Coker) comb. nov.

Basinym: Clavaria subspinulosa Coker, Clav. U.S. Can. 1923, t. 44, 45, 86.
6-10·5 X 3·5-7 cm., much branched, cinnamon-brown below, passing through light
cinnamon to pale lavender-pink above, the upper parts darkening to pinkish cinnamon
with age and finally wholly dark fleshy brown, the tips remaining pale lavender-pink or
lilac longer than the branches: stem 2-3 X 1-3 cm., stout, distinct, roughly felted with
fibrous rhizoids at the base, pale cinnamon-buff, sometimes lavender above: branches
numerous, elongate or rather stumpy, dichotomous, often longitudinally rugulose
above: flesh concolorous, moderately brittle, drying whitish and chalky-friable (as in
R. formosa): taste acid-bitterish: smell faint, musty.
On the ground in mixed and pine-woods: U.S.A. (S. Car., Virginia, not rare).
Spores 7· 5-12 X 4-5·5 µ,, ochraceous, elliptic, sparsely verruculose or nearly smooth.
Basidia 7-8·3 µ, wide: hymenium 90 µ, thick, thickening with many embedded spores.
This species has unusual interest because it combines the characters of many others
and yet cannot be placed exactly with any of them. Thus it has the shape and con
sistency, when dried, of R. formosa and R. suecica, the colour of pale R. cacao, sug
gestive also of R. Mairei, and the spores of R. brunnea or R. secunda. Coker compares
it with R. spinulosa, but the European interpretation of this species suggests that the
resemblance is superficial. The thickening hymenium is interesting because it occurs
in very few large fleshy species of Eu-Ramaria, and seems to clinch the affinity with
R. subdecurrens.
TABLE LXVI. SPORE-DATA FOR RAMARIA SUBSPINULOSA
Spore-size in µ, (Coker)
-----Remarks -----
9·5-11(-12·5) X4·8-5·5(-6·5) Murrill 421, Virginia
9-12·2 X3·7-4·8 Coker 2689: slightly rough
7·5-1 l X3·8-4·8 2649
7· 5-10·8 X4·5-5·5 2653: mostly 9·3 X4·8 µ,
9-10·5 X4-5 2664 (type), 2390
7·5-10 X3·8-5 2635
9-9·5 X 4·5-5·5 2724
8·5-9·5 X4' 5-5 2856
R. suecica (Fr.) Donk

Basinym: Clavaria suecica Fr., Syst. Myc. 1, 1821, 469: Coker, Clav. U.S. Can.
1923, t. 63, 89: Bourd.. Bull. Soc. Myc. Fr. 49, 1932, 207: Wehm., Pap. Mich. Ac.
Sci. Arts Lett. 20, 1935, 259: Lundell, Sv. Svamp. 9-10, 1937, 13.
Synonyms: Clavaria circinans Pk., Rept. N.Y. St. Mus. 39, 1886, 43, t. 1 (fide Coker).
C. flavula Atk., Ann. Myc. 6, 1908, 56: (fide Coker).
-
630 RAMARIA
Clavariella suecica(Fr.), Karst., Hattsv. 2, 1882, 184.
2-10 cm. high, pale whitish flesh-colour, pinkish buff, pinkish tan or rose-pink,
deepening upwards with age, then fading cream-ochraceous, alutaceous, tan or pale
cinnamon-buff, tips concolorous or whitish: stem 4-20 mm. wide, thick or slender,
arising from a white tomentose base, with white rhizomorphs, sometimes branched from
the base: branches rather upright, branched one to three times about the same levels, with
a brush-like mat of pointed tips, somewhat divergent or erect on one side of the stem
and outcurved on the other, main branches few, lower axils with orange-brown
scurfy areas (Wehm.): flesh pinkish or nearly white, coriaceous-soft, pliable, toughish,
often very soft, drying brittle and chalky: taste bitter, slightly or strongly: smell musty
or rancid (Wehm.), agreeable(Lundell) or none (Coker).
In coniferous humus, oftea in troops and in lines(? mycorrhizal), also under hard
woods (Wehm.): N. temp.(northern Europe, China, Canada, northern U.S.A.).
Spores 7-11·5 X 2·5-5µ,, mostly 7-9 X 3-4, ochraceous (Karst., Lundell, Bourd.,
Coker: pale brown, Wehm.), rough or minutely verrucose, oblong to fusiform.
Basidia 30-50 X 5-7" 5 µ,, 4-spored(Bourd.).
Hymenium 50-70µ, thick, not thickening(Coker: ? including subhymenium).
Hyphae 3-6 µ, wide, in places -9µ, wide, clamped, thin-walled (Coker).
Because of its tough, though soft, flesh, this species is usually placed in the Stricta
group. In all other points it agrees with R. formosa, particularly in the texture of the
dried flesh. R. conjz�nctipes, which is near to R. formosa, is also said to have rather
tough flesh. I have, therefore, placed R. suecica in the Formosa-group. It is a northerly,
subarctic or mountain, species which seems readily identifiable, to judge from the
general agreement in descriptions. For this reason, Doty's specimens from Oregon
(1944, p. 49), seem hardly to belong: they are described as yellow without a trace of
the characteristic pink which pervades R. suecica, at least when in full vigour, and as
similar to R. stricta, which hardly seems possible. They may represent R. stricta as
it usually occurs in Europe.
R. palmata var. guttaurensis may be R. suecica; perhaps, also, Wehmeyer No. n6
referred to R. secunda.
TABLE LXVII. SPORE-DATA FOR RAlvIARIA SUECJCA
Spore-size in µ Authority Country I----- ----------·

Remarks
9-n·5X4-4·5 Bourdot Sweden finely rough
7-8 X3-4 Karsten Finland
7·5-8·5X3-.:4 Lundell Sweden finely rough
7-9 X3·8-4·2 Coker " slightly rough (Herb. Fries.)
IOX5 Dank ?
7-8·5X2·5-4 Wehmeyer Canada finely rough
7·8-8X3-4 Coker U.S.A. Burnham 57 (N.Y.)
7·5-8·2X3·7-4·2 " " Coker 19 (N.Y.)
6·7-8·2X3·5-4 " " Farlow (New Hamp.)
7-9·3X3·4-4 " " C. fiavula (type)"
" Not 9-12X3-3·5 µ, as given by Atkinson.
R. testaceo-flava (Bres.) comb. nov.

Basinym: Clavaria testaceo-flava Bres., Fungi. Trid. 1, 1881, 61, t. 69: le. Myc.,
t. 1090.
Synonym: Clavaria oblectanea Britzm.: Hym. Sildb. 10, 1891, 149, f. 87: Rev.
Hym., 1909, 222.
3-5 cm. high: stem 1-2 cm. X 5-6 mm., or none, pale rufescent: branches little
branched, the main branches rather narrowly spathulate with furcate tips, polychoto
mous below, becoming dichotomous, cinnamon-rufescent or pale testaceous with egg-
RAMARIA
yellow, obtuse, multifid tips: flesh white, turning vinaceous on exposure to the air: taste a
little bitter, smell none.
In coniferous woods, on humus: Europe (N. Italy, Bavaria).
Spores 10-14X4-5µ, (11-15X4-5µ,, Coker), yellow under the microscope, granular
rough, oblong.
Basidia 35-45X4-5 fL·
This species is generally regarded as allied with R. Invalii and R. condensata. To
· me it is a diminutive or undeveloped, faded or dilutely and dingily coloured state of
R. formosa, like R. Bataillei only smaller and paler; it has the characteristic colour
change of R. f ormosa and identical spores.
R. testaceo-violacea (Doty) comb. nov.
Basinym: Clavaria testaceoviolacea Doty, Clav. Pac. N.W. 1944, 46, t. 4, f. 28, t. 6A.
-3 cm. high, irregularly branched two to three times, blood-red-brown to light reddish
salmon, often becoming nearly pale lilac or violaceous: stem indistinct: branches some
what flattened with rounded axils: flesh white, unchanging, bitter, without smell.
Among grass under American elms: U.S.A. (Oregon, Nov., Dec.).
Spores (8-)9-11(-12) X3 ·8-4· 5(-5)µ,, light ochraceous, verruculose-rough to nearly
smooth, ellipsoid, often 1-guttate.
Hymenium -uoµ, thick (? thickening): basidia 7-8·5µ, wide: stcrigmata 4, 7-10µ,
long.
This seems to combine the most vivid reddish pink of the Subbotrytis-series with
the lilaceous and violaceous tint of R. Mairei. Compare, also, R. rufo-violacea in
which the coloration appears inverted (rufous tips and violaceous branches). It shows
how extraordinarily complicated the interrelations of these large Ramarias are to our
present understanding.
R. testaceo-viridis (Doty) comb. nov.
Basinym: Clavaria testaceoviridis Doty, Clav. Pac. N.W. 1944, 47, t. 8.
Synonym: C. testaceo-fiava var. testaceo-viridis Atk., Ann. Myc. 6, 1908, 58.
-5X4 cm., clustered or subcaespitose, stout and compact, alutaceous to light pinkish
cinnamon, umber-brown where bruised, (Doty), pale drab (Atk.), tips pale yellowish green
to olive-green, drying pinkish buff to snuff-brown without a trace of green: stem short,
soon divided into prominent main branches 0·5 cm. long: branches divided about
three times, tips pluridentate, sometimes hollow.
In coniferous woods and on rotten fir-�ood in pastures: U.S.A. (N. Car., Oregon).
Spores 10-12X4-4:5µ, (type, Atk., Doty): 9-ux3·2-;·8 µ,, mo.;tly 10·5X3·75µ,
(Doty), pale yellow-orange (Doty), rough or indistinctly verrucose.
Hymenium 45-60µ, thick: basidia 4-spored.
This seems the nearest ally yet discovered of R. ochraceo-virens, but the indistinctly
roughened spores belong rather to R. fennica-R. Jormosa. The habit is that of R.
ochraceo-virens, if with the main branches a little more elongate.
R. tubulosa (Fr.) comb. nov.
Basinym: Clavaria tubulosa Fr., Elench. 1828, 229: (Berk., Hook. Journ. Bot. 1856,
275: doubtful identification).
Synonym: Lachnocladium tubulosum (Fr.) Lev., Ann. Sci. Nat. Bot. Ser. 3, 5, 1846,
159 (ut Eriocladus fistulosus err.): Pat., Journ. de Bot. 3, 1889, 33,. f. 8.
Erect, densely caespitose, very much branched, intensely yellow: branches hollow,
subfastigiate, strict, acuminate, velutinate, very fragile (dried): stems immersed.
On sandy ground: Brazil (leg. Bornemann): also (fide Pat.) French Guiana (leg,
Leprieur) and Guadeloupe (leg. L'Herminier).
Spores 10-12X5-6 µ,, brown, aculeolate (Pat.).
It is to be assumed that Patouillard examined the Friesian type-collection m
632 RAMARIA
identifying the later collections. The species must therefore belong in the Echino
spora-group, and its resemblance to Lachnocladium s. str. is superficial.
R. verna (Coker) comb. nov.

Basinym: Clavaria verna Coker, Clav. U.S. Can. 1923, 141, t. 52, 86.
-10X9 cm., wholly pale tan or buff or pale flesh-tan: stem 1-3 cm. high and wide,
stout, short, distinct, white: branches numerous, open, lax, slightly rugose upwards,
rather blunt, tips concolorous: flesh white, sometimes ochraceous when old or bruised,
not reddening, soft, not brittle: smell and taste none.
On the ground in woods: U.S.A. (N. Car., in May).
Spores 6·5--<;·5(-10·5)X3-4µ, maize-yellow to pale tan-flesh colour, verruculose,
minutely rough or nearly smooth.
Basidia 5-6·5µ wide, 4-spored.
Hymenium 50-60µ thick: hyphae 4·5µ wide.
Wehmeyer suggests this is only a vernal and slightly small-spored state of R.
secunda. He found a collection in Nova Scotia which appeared intermediate between
them (see p. 621). On the other hand it is very similar to R. flavo-alba of Malaya.
Coker's spore-measurements are these:
No. 3068 7·5--<;·5(-10·5) X 3·3-3·7µ, mostly 8-8·5X3·5µ
,, 3053 7"5-8 X3·7-4µ
,, 3067 7-7·7 X3·7 µ
,, 3039 6·5-7·5X3-3·3µ, type.
R. xanthosperma (Pk.) comb. nov.

Basinym: Clavaria xanthosperma Pk., Bull. N.Y. St. Mus. 94, 1905, 21: Coker,
Clav. U.S. Can. 1923, 141, t. 86: Wehm., Pap. Mich. Ac. Sci. Arts Lett. 20, 1935, 263.
3--<J X4-6 cm., white then yellowish or cream colour, sometimes red on bruising
(Peck), ivory yellow, cream or naples-yellow to cream-buff, tips somewhat brighter yellow,
with dull vinaceous tints where bruised (Wehm.): stem 1-1·5 cm. wide, stout, breaking
up into stout branches 6-12 mm. wide: branches many, compact, erect, with a broad
curving angle, slightly lacunose, irregularly branched, tapered: flesh white, somewhat
gelatinous-watery, fibrous-tough.
On the ground in woods: U.S.A. (N.Y.), Canada (Nov. Scot.), rare.
Spores 13-15X4-5µ, pale yellow, oblong (Peck): 12-15·5X4-5µ, nearly smooth
or very slightly verruculose (Coker, for Peck's type): 12-14X2·5-3·5µ, light yellow
brown t smooth, oblong-ellipsoid, sinuous, bent at one end, apiculate at the other
(Wehm.).
Basidia 7-8µ wide.
Hymenium filled with gelatinous, granular, gloecystidium-like elements, possibly
young basidia (Wehm.).
This is a rare species distingui$hed by the pale ivory colour, vinaceous discoloration,
and long, smooth spores. Wehmeyer says it is near C. albida Pk. (see R. obtusissima)
and C. crassipes Pk. (see R. secunda). I think it is identical with R. decolorans of Fin
land, but I refrain from uniting such rare and little-known fungi. If R. xanthosperma
is found to have the same spore-range as R. decolorans, it can then hardly be doubted
that they are conspecific, but Wehmeyer's measurements suggest a narrower spore for
the N. American species.
R. Zippelii (Lev.) comb. nov.

Basinym: Clavaria Zippe/ii Lev., Ann. Sci. Nat. Ser. 3, 2, 1844, 215: Bres., Ann.
Myc. 5, 1907, 239.
Synonyms: Clavaria Dozei Lev., l.c. 216; (fide v. Ov.).
? Clavaria nigrescens Fr., Nov. Symb. Myc. 1851, u6.
RAMARIA 633
C. Thwaitesii B. et Br.,Joum. Linn. Soc. Bot. 14, 1874, 75; (fide Petch: 'with
fuscous tips').
? C. phaeocladia Pat.,Bull. Soc. Myc. Fr. 1898, 189.
Lachnocladium Hookeri B.,Joum. Linn. Soc. Bot. 14, 1875, 75.
L. echinosporum Bres.,Ann. Myc. 9,1911,551.
Phaeoclavulina Zippelii (Lev.) v. Ov., Bull. Jard. Bot. Buit. Se:. 3, 5, 1923, 276:
le. Fung. Mal. 1, 1923,t. 1,f. I.
(Lachnocladium neglectum Mass., ? ined.)
Text-Figs. 2B and D,250d, 268,269,271,272: Plate 14.
FIG. 268.
TEXT-FIG. 268. Ramaria Zippelii, X r.
TEXT-FIG. 269. Ramaria Zippelii, young fruit-bodies, X 1: branch-tips, X 5.
-13 cm. high,solitary,pale alutaceous, pale ochraceous or dirty ochraceous, then pale
fuliginous-tan, fuliginous-ochraceous, or cinnamon-subfuscous, dingy, dusted with the
brownish ochraceous spores, tips white, all parts quickly turning dingy rufescent
vinaceous or purple-brown when bruised: stem 1·5-0· 5 cm. X 5-10 mm., cylindric,
rather long and slender, thickened upwards, and concolorous,base whitish: branches
4-6 mm. wide below, 1·5-2 mm. above,numerous,polychotomous at first (3-8 teeth),
becoming 2-3-chotomous, rather congested and stiffly upright, cylindric, tips rather
bluntly tapered, simple or bifid with two short and slightly incurved, sublunate tips,
outer branches often curved ascending: uppersides of the branches and axils more or
RAMARIA
less sterile, subtomentose, pale alutaceous: flesh white, quickly changing colour, rather
fibrous and tough: smell strong, of Ganoderma (paracresol).
On the ground in the forest: India, Ceylon, Java, Malaya, Borneo, New Guinea,
New Caledonia.
Spores 10-15X5-8 µ, (spore-body), sometimes 13-18 X7--9 µ,,1 deep ochraceous to
Jerruginous-ochraceous, echinulate with acute colourless spines 1-2µ, long, elongate
pip-shaped, 1-2-guttate, or with several guttulae.
T'ExT-FIG. 270. Ramaria Zippelii var. gracilis, X 1!:

branchlets, x 4.
Basidia 40-50X7--9µ,, subclavate, projecting -15µ,: sterigmata I-2, 6-10µ, long:
spores often more or less reversed on the sterigma with the apiculus pointing abaxially
or laterally ..
Sterile basidia like the fertile basidia but without spores and with more or less
aborted sterigmata, with brownish yellow guttulate contents, abundant in the fertil,
and sterile hymenia.
Hymenium at first 50µ, thick, gradually thickening to 200µ,: subhymenial hyphae
2-3·5µ, wide.
Sterile hymenium on the lower part of the stem and on the uppersides of the
branches 50-70µ, thick.
Hyphae 3-7µ, wide, narrower, slightly inflated, clamped, cells 40-350 µ, long, walls
smooth, thin or slightly thickened (less than 0·5µ,}, occasionally with short, clavate,
thin-walled vacuolate branches 20-30 x 7-1oµ,: construction as in R. cyanocephala.
1
Probably monospores.
RAMARIA 035
Colour caused by yellowish or brownish guttulae in the hyphae ofthe suohymenium
tttt •• ',
and sterile hymenium,and by the large yellowish guttulae and brownish-ochre cyto
plasm of the fertile and sterile basidia: hyphae of the flesh and growing tips colourless.
TEXT-FIG. 271. Ramaria Zippe/ii (left), normal and abnormal basidia: var. gracilis (right):
X 1,000.
var. gracilis var. nov.

Text-Figs. 270,271: Plates 12,13.
2-8 cm. high,smaller and more slender, sometimes darker brown; stem 0·5-4 cm. X
2-5 mm., slender: branches 1-2 mm. wide below, generally dichotomous from the
first,generally rather lax and spreading,the bifurcate tips with semilunar axils: flesh
white, turning purplish vinaceous on bruising (unchanging in some collections),smell
sometimes absent.
On the ground in the forest: Malaya (Johore, Gunong Pulai, Oct. 1941, type;
Kluang, Nov. 1941); (Singapore, Bukit Timah, 3-4-43; Botanic Gardens, 31.12.19,
No. 5484); (Pahang, Tembeling, 20.11.30).
Spores 7-10 X 4·5-6·5 µ., spines 0·5-1 µ. long,smaller with shorter spines.
Basidia 40-65 X 8-11 µ.: sterigmata 2-3-4,mostly 3 or 4,6-7 µ. long.
Hyphae and general structure as in the typical state, but the sterile hymenium
feebly developed as abortive clusters of sterile basidia,or even absent from the then
naked uppersides of the branches.
My reasons for distinguishing R. cyanocephala have been given on page 573.
R. Zippelii thus stands very near R. grandis of temperate N. America and Japan.
Indeed,R. grandis may be identical,because the only difference that I can detect in
its description is a richer brown colour. Similarly,var. gracilis may be the 4-spored
R. longicaulis of N. America which is also variable in the colour-change of its flesh:
but Coker remarks that the tips of R. longicaulis are paler concolorous with the
RAMARIA
branches, not abruptly white as in R. grandis (and as in var. gracilis). Until the
variation of these fungi is better known, to combine them may only be confusing.
Concerning var. gracilis, I have possibly piaced more than one variety under
this name, for the Singapore collection has 4-spored basidia while the Johore and
Pahang collections have 3-, occasionally 2-,
spored basidia, in which respect they connect
the 2-spored R. cyanocephala, R. Zippelii, and
R. grandis with the 4-spored R. longicaulis.
R. megalorhiza may be a related species with
smaller, verruculose spores, paler colour, and
unchanging flesh, but it is inadequately known.
The 4-spored basidia of var. gracilis have
the same size as the 2-spored basidia of typi
. cal R. Zippelii, and the spores of var. gracilis
have roughly two-thirds the linear measure
of those of typical R. Zippelii. It would seem,
therefore, that the large size of the spores of
typical R. Zippelii is caused merely through
reduction to the 2-spored state. But that the
relation is not so simple is shown by the
small-spored, 2-spored collections of var.
gracilis.
R. Zippelii may be pantropic, as R. cyano
cephala.
L. echinosporum Bres., as described, is
indistinguishable from R. Zippelii. Its spores
approach those of var. gracilis, but it is des
cribed as a stout fungus, and the spores are
probably not a fair sample. As a dried speci
men it cannot be otherwise identified.
C. nigrescens is likewise doubtful. Its
description is :
TExT-FIG. 272. Ramaria Zippelii:
medullary hyphae, X 500. Cinnamon (fresh), divided to the base,
drying black, woody, and sulcate-lacunose,
branches solid, thick, obtuse: Madeira (leg. Kamphovener).
C. phaeocladia was said to have smooth, colourless spores 13 X 6-8µ, but as an
entirely brown, terrestrial, branched Clavaria (8-10 cm. high) from Java it agrees well
with ]?.. Zippelii: the spores may have been immature. Compare, however, Lentaria
surculus. Possibly R. albida also belongs here.
The type of L. Hookeri (Khasya Hills, India) in the Kew herbarium has the typical
spores of R. Zippelii (u·5-14·5x6-7µ, echinulate, spines 1µ long). It might, of
course, have been R. cyanocephala, for there is no field-note on the colour. Similarly,
the type of L. neglectum (Sarasii1173, La Foa, New Caledonia) in the Kew herbarium
·s
has spores 17-18 x 8-9µ, spines 1-1 µ, on 2-spored basidia.
RAMARIOPSIS Donk emend.

Rev. Nied. Homob. Aphyll. 2, 1933, 89 (ut subgenus).
Fruit-bodies branched, rarely simple, variously coloured, waxy-fleshy, brittle or
rather tough: stem generally distinct and rather slender, often white villous: branches
radial, polychotomous or dichotomous, cylindric.
Terrestrial, rarely lignicolous: 10 spp., mostly N. temp., 2 in S. Austral., 1 in trop.
Amer.
RAMARIOPSIS 637
Spores white, ellipsoid or subglobose, finely verruculose or echinulate (spines c. o·5µ
long or less), 1-guttate.
Basidia clavate, small to medium size: sterigmata 4, occasionally 2, straight or
slightly curved.
Cystidia none.
Hymenium thickening or not: subhymenial hyphae narrow.
Hyphae monomitic, inflating, clamped, not secondarily septate, with thin or slightly
thickened walls, generally rather narrow 4-10 µ wide (-20 µ in R. asterospora), with or
without narrow interweaving hyphae.
Type-species: R. Kunzei.
This genus is ClltVUlinopsis with very finely echinulate small spores. The species
are, with one exception (R. asperulospora), branched. They connect, therefore, with
the branched species of Clavulinopsis and appear as a distinct, monophyletic group.
The spines are much smaller, more numerous, and more regular than the relatively
stout processes on the spores of Clavulinopsis subgen. Acularia. Care must be taken
to distinguish Scytinopogon, which has angular, as well as echinulate, spores and
flattened branching.
Clavulinopsis luteo-ochracea may belong to Ramariopsis. Compare also Clavaria
gracillima sensu Wehm., from Nova Scotia (p. 227, under Clavaria argillacea), which
may be a new, simple species of Ramariopsis.
KEY TO THE SPECIES OF RAMARIOPSIS

Sp. ellipsoid, 6 µ or more long
Sp. 5-7 X 2·8-3·5µ, verruculose: -5 cm. high, straw-colour,
somewhat rugulous, rather sparingly and irregularly bi:anched:
Spain R. clavuligera
Sp. 6-8 X 4-5µ, echinulate-rough: Austral. R. lorithamnus
(Simple: see Clavulinopsis subgen. Acularia)
Sp. subglobose or less than 6µ long
Simple, wood-brown: sp. 5-7 µ, globose: U.S.A. R. asperulospora
Branched
Lilac or violet, stem often reddish or yellow, -2·5 cm. high,
slender: sp. 2·5-4µ: Europe, N. Amer. R. pulchella
White
Wholly white or pallid: N. temp. . R. Kunzei
(Stem grey: -15 cm. high: on rotten wood: U.S.A. C. lentofragilis,
p. 643)
(White to alutaceous, pinkish or yellpwish brown towards
the tips: U.S.A. C. arborea,
p. 643)
Yellowish, orange, or greenish
4-7 cm. high, ochraceous, much branched: U.S.A. R. asterella
-3·5 cm. high, pale yellowish drab, slender, sparingly
branched: England . R. tenuiramosa
-1·5 cm. high, greenish yellow, much branched from the
base: Sweden . R. curta
Golden to orange, branches few, -2·5 cm. high
Stem smooth or minutely scurfy: N. temp., S. Austral.. R. crocea
Stem tomentose, rough: N. Amer. R. vestitipes
RAMARIOPSIS
(-5 cm. high, simple or sparingly branched, yellowish: stem
scurfy downwards, becoming rufous ochraceous: N. temp. Clavulinopsis
luteo-ochracea)
R. asperulospora (Atk.) comb. nov.

Basinym: Clavaria asperulospora Atk., Ann. Myc. 6, 1908, 55: Coker, Clav. U.S.
Can. 1923, 63, t. 81.
4-7 cm. X 2-3 mm., simple, clustered, cylindric, blunt, wood brown.
On the ground: U.S.A. (N.Y.)
Spores 5-7µ, white, finely and shortly echinulate as in R. Kunzei, globose.
Basidia 3ox 10-12µ: sterigmata 4.
Coker says this is a good species, not recently collected. Not to be confused with
Clavaria purpurea with ellipsoid spor':'s. It is the only species known in the genus
with simple fruit-bodies: the size of the spores suggests Clavulinopsis subgen. Acularia,
cf. C. helvola ( = Clavaria dissipabilis).
R. asterella (Atk.) comb. nov.

Basinym: Clavaria asterella Atk., Ann. Myc. 6, 1908, 55: Burt, Ann. Mo. Bot. Gdn.
9, 1922, t. 7, f. 52: Coker, Clav. U.S. Can. 1923, 105, t. 84.
5-7 cm. high, branched, ochraceous, narrow: stem short: primary branches open,
divaricate, axils rounded, upper branches fastigiate: soft, flexible.
On humus: U.S.A. (N. Car.).
Spores 4-5x 2·5-3µ, white, finely and shortly echinulate as in R. Kunzei, subglobose
or obovoid, r-guttate.
Coker suggests this is a colour form of R. Kunzei. R. tenuiramosa may be related.
R. clavuligera (Heim) comb. nov.

Basinym: Clavaria clavuligera Heim, Fungi lberici 1934, 46, f. ro.
2-5 cm. high, rather sparingly and irregularly branched, wholly straw-colour: stem
-10 X 2-6 mm., very short, often concrescent with others, with slender rhizoids at the
base: branches few (5-12), as thick as the stem, cylindric or somewhat flattened, at first
verticillate, claviform, obtuse, shortly bifid or trifid, ultimately slender, unequal, some
times strongly incurved, sulcate, rugose: tips simple or bifid, obtuse or abruptly·
narrowed: fleshy tough, pliant.
On the ground among grass: Spain.
Spores 5-7 X 2·8-3·6µ, white, obovate-cylindric, set with minute, conical, spaced
warts.
Basidia 6-7 µ wide: sterigmata 4, 5 µ long.
Hyphae 4-<J µ wide.
This is like a malformed, pale yellowish R. Kunzei with ellipsoid spores, perhaps
related to R. asterella and R. tenuiramosa with shorter spores. On the other hand, it
may be a form of Scytinopogon angulisporus: the absence of tomentum at the base of
the stem, the presence of rhizoids, and the ellipsoid spores suggest that genus which
may well grow in southern Europe as in the southern United States: the inflated
hyphae and spaced warts on the spore indicate, more certainly, Ramariopsis, as R.
lorithamnus.
R. crocea (Fr.) comb. nov.

Basinym: Clavaria crocea Fr., Syst. Myc. r, 1821, 472: sensu Coker, Clav. U.S.
Can. 1923, 107, t. 28, 84: Wehm., Pap. Mich. Ac. Sci. Arts Lett. 20, 1935, 256: Clel.,
Toadstools . .. S. Austral. 2, 1935, 267, f. 60: Fawcett, Proc. Roy. Soc.Viet. 51, 1939,
14, t. 4, f. 3.
Synonyms: Clavariella crocea (Fr.) Harter, Krypt. FI. Mark Brand. 6, 1910, 164.
RAMARIOPSIS 639
Clavaria Kalchbrenneri Muell., Proc. Linn. Soc. N.S. Wales 1882, 105: (haud
C. Kalchbrenneri Sacc. 1888 = Ramaria Kalchbrenneri).
-5 cm. high, branched, solitary or 2-4 together, wholly golden-yellow to rich chrome
orange, occasionally greenish when bruised (Coker): stem -1 cm. long, t-! the height
of the fruit-body, distinct, minutely furfuraceous: branches laxly dichotomous two to
four times, cylindric, axils lunate, tips acute or blunt: tender but elastic, not brittle:
no smell or taste (Coker, Doty), taste bad but no smell (Wehm.).
On the ground among grass in woods: Europe (France, Germany, Gt. Britain,
Holland, rare), U.S.A. (Conn., N.Y., Pa., N. Car.), S. Austral., Japan.
Spores 3-4 X 2-3·5 µ., white, obscurely asperulate, subglobose, 1-guttate.
Basidia 4-spored.
Hymenium 20 µ. thick (? not thickening).
Hyphae 2·5-5·5µ., clamped.
This is Coker's interpretation of Clavaria crocea Fr., which appears to be very rare
in Europe, yet in his sense it seems to be a widespread temperate or subtropical
species. Superficially it is like a small, delicate, and more brilliantly coloured Clavu
linopsis corniculata: microscopically it has spores like Ramariopsis Kunzei, and parti
cularly R. pulchella, which may be, as Coker suggests, only a differently coloured state
of R. crocea. According to Coker the only correct figure of the European plant is
Persoon's (le. et Descr. 2, 1798, t. 9, f. 6), Britzelmayr's being erroneous. R. vestitipes
seems identical. According to Coker and Doty there may be pale yellow and white
forms.
There appears to be some variation in the size of the spores, as shown in the accom
panying Table. The measurements given by Massee and Cooke (6-7 X 2-4 µ.) are clearly
wrong.
TABLE LXVIII. SPORE-DATA IN RAMARIOPSIS CROCEA
Authority Spore-size in µ. Remarks

Coker 2·5, subglobose No. 4660 (N. Car.)
3-3·3 X 2·5-2·7 No. 21
3-3·7 X 1·8-2·5 No. 5599 (N. Car.)
3-3·6 X 2·2-2·8 No. 4805 (N. Car.)
" 3·4-3·7 No. 4843 (N. Car.)
Atkinson 4x3 No. 23376 (N.Y.)
Wehmeyer 2·5-3·5, globose U.S.A.
Doty 3-4·5 X 3-4 "
Cleland 4 x 3·5, pip-shaped S. Austral.
Imai 2·5-3·5, globose Japan
R. curta (Fr.) comb. nov.

Basinym: Clavaria curta Fr., Monogr. Hym. Suec. 2, 1857, 281: Cke., Grev. 2,
1874, 136: ibid. 4, 1875, 67.
1-1·5 cm. high, very much branched, greenish yellow, compressed: stem none:
branches obtuse, crowded.
Among grass and fir-needles: Europe (Sweden, ? Gt. Britain).
Spores 4-5 X 3-3 ·5 µ., white (? yellowish), minutely rough, subglobose (Coker, for
Friesian specimen in the Kew herb.): 3·8-5 X 2·5-3·5 µ. (Coker for specimen in Herb.
Bresad. det. C. curta).
This rare species may be only a form of R. crocea. Coker suggests that, since the
spores of Friesian specimens of C. curta appear yellowish, it may be Ramaria decurrens:
macroscopically there is little agreement, and the spores are typical of Ramariopsis.
The English records are doubtful.
RAMARIOPSIS
R. Kunzei (Fr.) Dank

Rev. Nied. Homob. Aphyll. 2, 1933, 89.
Basinym: Clavaria Kunzei Fr., Syst. Myc. 1, 1821, 474.
Synonyms: C. chionea Pers., Myc. Eur. 1, 1822, 161 (teste Bourd. et Galz., Cott. et
Wakef., Coker, Dank).
C. subcorticalis Schw., Tr. Am. Phil. Soc. 2, 4, 1832, 182: (teste Coker).
? C. setosa Balb. et Nace., Fl. Tice. 2, 345, t. 22, f. 2: Sacc. Mich. 1, 1878, 548.
C. velutina E. et E., N. Am. Fung. Ser. 2, 1888, No. 2024: (teste Coker).
? C. arborea Atk., Ann. Myc. 6, 1908, 56: Burt, Ann. Mo. Bot. Gdn. 9, 1922, 30,
t. 6, f. 38.
C. asperula Atk., Ann. Myc. 6, 1908, 54: (teste Coker).
C. asperulans Atk., ibid., 55: (teste Coker).
? C. lentofragilis Atk., ibid., 57: Coker, Clav. U.S. Can. 1923, 102, t. 84: Burt,
Ann. Mo. Bot. Gdn. 9, 1922, 37, t. 7, f. 54.
C. elongata Britz., Rev. Hym. 1909, 221: Hym. Tab. 472, f. 50.
C. subcaespitosa Pk., Bull. N.Y. St. Mus. 167, 1913, 39: Coker, Clav. U.S. Can.
1923, 101, t. 84: Coker, Journ. El. Mitch. Sci. Soc. SS, 1939, 385, t. 43, 44.
Lachnodadium subsimile B. et C., Grev. 1, 1873, 161: (teste Coker).
Lachnocladium subcorticale (Schw.) Burt, Ann. Mo. Bot. Gdn. 9, 1922, 66:
(= Clavaria subcorticalis Schw.).
Clavulina Kunzei (Fr.) Schroet., Krypt. Fl. Schles. Pilze r�88, 442: (quoad nomen,
haud quoad descript. = Clavulina cristata, fide Dank).
Ramaria Kunzei (Fr.) Quel., Fl. Myc. 1888, 464.
? Clavaria dealbata B., Hook. Journ. Bot. 1856, 275.
l C. cervina B. et C., Journ. Linn. Soc. Bot. 10, 1868, 338.
? C. pallida, ibid., 338: (= C. cervina, teste Burt).
? Lachnocladium cervinum (B. et C.) Pat., Journ. de Bot. 3, 1886, 26: Burt, Ann. Mo.
Bot. Gdn. 6, 1919, 273, t. 5, f. 9, Text-Fig. 12: Overholts, Sci. Surv. P.R. and Virg. Isl.
8, 1926, 148: Martin, Lloydia 4, 1941, 266: (= C. cervina B. et C.).
? L. pallidum (B. et C.) Pat., Journ. de Bot. 3, 1886, 26: Burt, Ann. Mo. Bot. Gdn. 6,
1919, 273: (= C. pallida B. et C.).
? L. dealbatum (B.) Cke., Grev. 20, 1901, ro: Burt, Ann. Mo. Bot. Gdn. 7, 1922, 72,
t. 11, f. II4: (= c: dealbata B.: non v. d. Bijl = Scytinopogon angulisporus).
Pseudonym: Clavaria subtilis sensu Bres., le. Myc., t. 1099: (non Fr. = Clavu
linopsis subtilis).
2-12 cm. high, snow-white, ivory-white, or cream-white, rarely tinged pink or flesh
colour, solitary, gregarious or caespitose, much branched or with few branches, rather
pliant and elastic, varying more or less brittle, stem and bases of the main branches shortly
villoso-tomentose: stem 0·5-2·5 cm.X2·5-6 mm., sometimes absent, generally distinct,
sometimes becoming yellow or pink at the base: branches in large specimens 2-5 mm.
thick and 3-5-chotomous below, becoming narrow (0·5-2 mm. thick) and dichotomous
above, erect, outcurving round the sides of the fruit-body, fastigiate, crowded or loose,
generally cylindric, sometimes flattened, axils rounded and narrow, in small specimens
slender and repeatedly dichotomous, sometimes with great regularity, tips acute or blunt,
never cristate: smell and taste none, or slight and inparticular.
In woods and pastures, mostly terrestrial, occasionally on decayed wood: N. temp.,
rather common (Europe, N. Amer., Japan):? trap. Amer. (Jamaica, Cuba, Brazil):
? Natal: Queensland.
Spores 3-5·5X2·3-4·5µ, white, minutely echinulate, verruculose or merely asperulate,
broadly ellipsoid to nearly globose, with a small apiculus, the spines to 0·5µ long and
acute, or shorter and blunt, the wall slightly thickened, 1-guttate.
Basidia 25-40X5-6µ, clavate: sterigmata 4, occasionally 2, 4-4·5µ long.
RAMARIOPSIS
Cystidia none..
Hymenium 30-40 µ thick at first, gradually thickening to 100 µ, absent from the
trunk and upper villous sides of the main branches: subhymenium well-developed,
composed of narrow, 2-3 µ wide, closely interwoven hyphae, not inflating.
TFXT-FIG. 273. Rarnariopsis Kunzei, X 1.
@ ·�oo··· G)·· Ar.·.:;i

i�I..
(.".: t:;:,
\;;' ,,• ,.�-. t:.�
TEXT-FIG. 274. Rarnariopsis Kunzei: spores, X r,ooo.
Hyphae monomitic, with clamp-connexions, 3-10µ wide, the cells 40-320µ long,
thin-walled, smooth: in the tomentum on the stem, 2-4µ wide, loosely interwoven,
clamped, with smooth, thin walls.
var. deformis var. nov.

Synonym: Clavaria Krombholzii Fr., Epicr. 1838, 572, pr. p. Krombh., t. 54,
f. 15-16 (non f. 18-20 = Clavulina rugosa var. alcyonaria): Bres., le. Myc., t. 1097:
Donk, Rev. Nied. Homob. Aphyll. 2, 1933, 90: Lundell et Nannf., Sv. Svamp. 1-2,
1934, 17.
3-6 cm. broad and high, usually caespitose, short, squat, compact; branches Jew,
compressed or clavate, often deformed, blunt.
Europe: with the typical state.
Spores typical.
var. Favreae var. nov.

Synonyms: Ramaria Favreae Quel., Ass. Fr. 22, 1893, 489, t. 3, f. 13.
Clavaria Favreae (Quel.) Sacc. et Trav., Syll. 21, 1912, 429: Bourd. et Galz., Hym.
Fr. 1928.
As var. deformis, but the branches much flattened, spathulate-flabelliform or
laminate (as a rudimentary Sparassis), often curved, the ultimate branches horn-like
with dentate or subcristate tips, white, fragile.
Spores 3-5 x 3-4µ (Bourd. et Galz.), 6µ ellipsoid (Quel.).
5119 Tt
RAMARIOPSIS
In coniferous woods: France (rare).
Coker and Konrad and Maublanc reduce this to C. coralloides, which they also reduce
to Clavulina cristata. The spores of C. Favreae, however, are those of R. Kunzei, and
it is obviously related to var. deformis or the 'Krombholzii-form' of R. Kunzei.
var. subasperata var. nov.

Synonym: Clavaria minutula Bourd. et Galz. f. subasperata Bourd. et Galz., Hym.
Fr. 1928, 105.
1-2 cm. nigh: branches few, forked or bidentate.
Terrestrial: France.
Spores 4-4·5 X 3-4µ., very laxly and vaguely asperulate.
Basidia 18-24 X 4-4·5 µ..
Hyphae 2-6 µ. wide.
Variability. This widespread and not uncommon fungus is extremely variable, as
the list of synonyms implies. Its limits are imperfectly known. Careful descriptions
of separate colonies of fruit-bodies are needed to determine whether varieties can be
distinguished by the size, colour, branching, and texture of the fruit-body, the size and
asperulation of the spores, habitat, and so on. The branching is usually described as
dichotomous, which is true only of slender specimens (e.g. Quel., Champ. Jura et
Vosges 3, 1875, t. 2, f. 11). Large specimens are polychotomous at first. Wehmeyer
has described small specimens with flattened, almost digitate, branching, which may
belong, however, to Scytinopogon (Pap. Mich. Ac. Sci. Arts Lett. 20, 1935, 258, t. 52).
The specimens which I have called var. deformis have irregular, misshapen branches,
but whether they are a normal growth, or merely young states of large fruit-bodies, or
abnormalities caused by frost or some other agent is not known. The texture is usually
rather brittle, toughening on drying (so that dried, exotic specimens were referred to
Lachnocladium by Patouillard), but Coker considers it to be rather tough 'in the
United States and distinguishes Clavaria subcaespitosa by its very brittle, translucent
white branches: such would be considered typical R. Kunzei in Europe. The habitat
is usually terrestrial, or humicolous, but Wehmeyer records it on much-decayed wood
in Nova Scotia: C. lentofragilis, which seems identical, was found on rotten wood in
a sphagnum bog at Long Island: C. cervina, also seemingly identical, was found on
dead wood in Cuba. It appears never to have been found on wood in Europe, where
the typical state is well known. The distribution is clearly north temperate. It has
not been recorded certainly from south temperate regions: the Natal record is con
sidered doubtful by van der Bijl, and may well refer to Scytinopogon; but Cooke's
record from Queensland has been confirmed by Fawcett from a specimen named by
Cooke in the Melbourne herbarium (sp. 3·5-4·5 X 2·5-3·5 µ., rough). It has not been
recorded from tropical Africa or tropical Asia but, as discussed later under C. cervina
and C. dealbata, it seems to occur in the American tropics. Coker records it from
Jamaica at an altitude of 5,000 ft., where the climate may resemble that of a temperate
summer.
The spore-variation, which is extremely slight, is shown in Table LXIX. Var. sub
asperata, referred by Bourdot and Galzin to Clavulinopsis minutula, has larger spores
than typical C. minutula and seems to be very small specimens of R. Kunzei. The
American collections seem often to have slightly smaller spores than the European,
though C. subcaespitosa agrees with the European collections, as it does macro
scopically.
Synonyms. In Europe there are two, if not three, synonyms. C. chwnea is generally
acknowledged to be R. Kunzei and Coker found the typical spores on Persoon's
specimens of C. chionea (Rea's interpretation of C. chionea may be Clavulinopsis
subtilis, which has the !,lnooth, 1-guttate spore of Clavulinopsis). C. elongata is, at
most, a form with elongate internodes. C. setosa is indistinguishable from the descrip
tion (spores 3·5-4µ.). In the United States there appear to be eight synonyms. The
RAMARIOPSIS
species seems to be more variable there, and to Coker we owe the elucidation and
correct definition of the species with its characteristic spore: in Europe the spore had
previously been described as smooth, and the species had been confused with Clavulina
cristata and Clavulinopsis subtili.�. Coker admits C. subcorticalis, C. velutina, C.
asperula, C. asperulans, and L. subsimile as synonyms, but keeps C. arborea, C. lento
fragilis, and C. subcaespitosa apart. He regards R. Kunzei as a small Clavaria 3-6 cm.
high, rarely 9 cm., with rather tough flesh. C. subcaespitosa he considers to be larger
(-12 ctn.), very fragile, translucent white, with larger spores (4· 5 X 3-4µ.): it agrees
well, however, with European R. Kunzei, as already noted. C. lentofragilis he con
siders as larger still with greyish stem and C. arborea as, perhaps, a colour form with
the branches becoming alutaceous and, towards the tips, rose-pink or yellowish brown.
Their descriptions are:
C. arborea Atk.
Very much branched dichotomously, curved and sometimes· deformed, white to
alutaceous, terminal branches rose-pink, or yellowish brown probably when old:
spores 3-4 x 2-3µ., white, obovoid, asperulate: basidia 4-spored: on the ground in
woods, U.S.A. (N.Y.).
G. lentofragilis Atk.
-x5 cm. high, 12 cm. wide: trunk 2-4X2-3 cm., grey: branches white, repeatedly
dichotomous, with soft fragile conic tips, taste and smell inparticular: spores 4-6 µ.,
white, ovoid to subglobose, asperulate: on rotten wood in sphagnum swamp, U .S.A.
(LoIJ.g Is!., N.Y.).
In reducing C. Krombholzii to a variety of R. Kunzei I have given it a new name to

avoid confusion. Originally C. Krombholzii covered this misshapen state of R. Kunzei
and also Clavaria grossa ( = Clavulina rugosa var. alcyonaria), as shown by Cotton
and Wakefield, and by Donk. C. Krombholzii has been variously taken for one or other
species by different mycologists. Donk and Lundell agree that there is such a mis
shapen fungus with the spores of R. Kunzei in Sweden, and this is what I intend by
var. deformis. C. Krombholzii and C. grossa are both nomina ambigua.
Coker mentions a white form of R. crocea collected by Jaap in Sweden: it has spores
2·5-3·7 X 2-3µ., obscurely rough. Possibly it is R. Kunzei var. subasperata. Bresadola
had named it Clavaria subtilis, which was his misnomer for R. Kunzei.
C. cervina, C. dealbata, C. pallida. In the American tropics there appears to be
either typical R. Kunzei or a very similar species which has been called Clavaria
dealbata and Clavaria cervina. As Coker first showed, their spores are identical with
those of R. Kunzei, but the fmit-body of the type of C. dealbaia resenibles that of
Scytinopogon, and C. cervina is said to be lignicolous. I think that when better known
they will be found to be a separate species, though the dried material affords no
reliable difference. Martin's recent Panama collection, referred to Lachnocladium
cervinum, supports this view, and it is certainly odd that R. Kunzei, so common and
widespread in the north temperate region, has not been reported from other parts of
the tropics. The descriptions are these:
C. cervina B. et C.
Synonym: Clavaria pallida B. et C. (teste Burt, Coker).
-4 cm. high, much branched from a stout trunk (tawny-olive or pale fawn when
dried) : trunk shortly tomentose, short: branches repeatedly dichotomous, slender,
with very acute tips, even subfiliform (hairy with hyaline hairs 1 · 5 µ. wide projecting
10µ. beyond the basidia, and longer on the stem: Burt): coriaceous: spores 3·7-4·5µ.,
white, subglobose, echinulate (Burt): spores of C. pallida, 3-3·7µ., verruculose as in
RAMARIOPSIS
R. Kunzei (Coker): on dead wood, Cuba (also Mexico and Guadeloupe, fide Pat.:
Puerto Rico, fide Overholts).
Fruit-bodies dull pinkish with white tips: distinctly fetid: spores pure white in
the mass, 3·5-4 X 2·5-3µ. (excluding the short spines): Barro Colorado Is!., Panama:
(Martin, G. W., in Lloydia 4, 1941, 266).
C. dealbata B.
-5 cm. high, white, opaque, as if whitewashed, caespitose, fastigiate: trunk short,
cylindric, 1·5-2 mm. thick, forked five to six times so as to make a tree-like tuft:
branches somewhat divaricate, broader than the stem, strongly compressed when
dried, the axils acute below, round above, tips acute: rather tough: spores 3-:r4µ.
(Coker), 3·5-4 X 2·5-3µ. (Burt), 3-4 X 2-3µ. (Corner, type in the Kew herb.) white,
verruculose or shortly echinulate as in R. Kunzei: on the ground, Brazil.
TABLE LXIX. SPORE-SIZE IN RAMARIOPSIS KUNZEi
Size in µ, Country Authority

3-5X3-4 France Bourdot, Galzin
3·5-4·5, globose Gt. Britain Cotton, Wakefield
4·5-5·5X3·5-4·5 " Corner
4-4·8, subglobose Holland Donk
4·8 x4
3·8-4·8X3·5-4·0
Italy
Sweden I Coker (herb. Bres., as C. suhtilis)
" (herb. Romell)
3·7, globose Europe " (C. chionea-type)
3-6X3-5 France Bourdot (C. Krombholzii)
4X3 Gt. Britain Rea (C. Krombholzii)
4-6X4-5 Italy Bresadola (C. Krombholzii)
4-4·5X3-4 France Bourdot (var. subasperata)
3-4, globose Germany Britzelmayr (C. elongata)
Average: 4·3X3·9 Europe
---·--
3·5-4·5, globose Japan Imai
3-5·2X2·2-4 U.S.A., Canada Coker (20 collections)
2·5-3, globose Canada Wehmeyer
3·5-4·5X2·3-3·5 U.S.A. Coker (C. asperula-type)
3·0, globose " " (C. subcorticalis-type)
2·5-3·2, globose " " (C. velutina-type)
3-4X2-3 " I Atkinson (C. arborea)
Average: 3·6X2·9 - U.S.A., Canada (25 collections)
4-6X3-4·2 U.S.A. Coker (C. subcaespitosa)
4-6, subglobose " " (C. lentofragilis)
I
3-4x2·0-3·5 Brazil Coker, Burt, Corner (C. dealbata)
3-4·5, subglobose Cuba Coker, Burt (C. cervina)
R. lorithamnus (B.) comb. nov.

Basinym: Clavaria lorithamnus B., Journ. Linn. Soc. Bot. 13, 1873, 169, no. 146:
Cke., Handb. Austral. Fungi 1892, 201.
Text-Fig. 275.
ODO 0-'
-4 cm. high, branched, pallid umber: branches straight,
apices shortly bifid and rather acute.
·.:· ;! On the ground: Austral. (Victoria, Dandenong).
�;.
Spores 6-8 X 4-5 µ., white, minutely echinulately rough,
TEXT-FIG. 275. Ramariop often rather sparsely, elliptic pip-shaped, not angular, the
sis lorithanmus. X r ,ooo. wall slightly thickened.
RAMARIOPSIS
Hyphae -10µ. wide, long-celled, clamped.
I have examined the type at Kew, and found these distinctive spores very abundant.
The species thus belongs to the same alliance as R. clavuligera.
. The record of C. lorithamnus from S. Africa is, according to van der Bijl, Ramaria
fiaccida.
R. pulchella (Boud.) comb. nov.
Basinym: Clavaria pulchella Boud., Bull. Soc. Myc. Fr. 3, 1887, t. 13, f. 2: Bourd.
et Galz.; Hym. Fr. 1928 (ut var. pulchella): Coker, Journ. El. Mitch. Sci. Soc. 63,
1947, 60, pl. l, 12.
Synonyms: C. tenuissima Sacc., Michelia 1, 1878, 436: (non C. tenuissima Lev.,
1846).
C. Bresadolae Cav., Myc. Lomb., p. 8, t. 26, f. 6-8.
C. Bizzozeriana Sacc., Syll. Fung. 6, 1888, 693.
C. exigua Pk., Rept. N.Y. St. Mus. 54, 1901, 155 (teste Coker).
C. conchyliata Allen, Tr. Brit. Myc. Soc. 3, 1908, 92.
C. pulchella Boud. var. Bizzozeriana Bourd. et Galz., Hym. Fr. 1928.
C. pulchella Boud. var. asperula Bourd. et Galz., ibid.
C. Cavarae Sacc. et Trott., Syll. 21, 1912, 436: (= C. Bresadolae Cav., non Quel.).
Text-Fig. 276: Plate ro.
1-2 cm. high, sparingly dichotomously branched one to three times, very slender,
violet, solitary or gregarious: stem short, generally distinct, white, yellow or reddish
yellow, base white tomentose: branches few, slender, filiform, divaricate: waxy-firm,
without smell.
On bare soil in woods: Europe (Gt. Britain, France, Italy, Switzerland), U.S.A.
(N.Y., N. Car.): uncommon.
Spores 3-4·5 X (2-)2·5-3·5µ., white, minutely verrucose (warts less than 0·5µ. high),
subglobose pip-shaped, slightly thick-walled, 1-guttate.
Basidia 20-30 X 6-7µ. (12-22 X 3-4·5µ., Bourd. et Galz.): sterigmata (2-)4, 4-5µ. long.
Hymenium thickening slightly -50µ. (or more?): subhymenium narrow; composed
of densely interwoven hyphae 1·5-3µ. wide: cystidia none.
Hyphae 2-7µ. wide, clamped, thin-walled or becoming slightly thick-walled, cells
40-200µ. long, some narrow hyphae with very elongate cells, with scattered small
crystals, 2-8µ. wide, between the. hyphae: the longitudinal hyphae occasionally bearing
short, clavate or fusiform, lateral processes, -30 x 8-15µ., filled with dense granular
cytoplasm, or with short, swollen, ampullaeform segments, 20-30 X 8-15µ., with
similar contents.
Hyphae of the tomentum at the base of the stem 1·5-2·5µ. wide, thin or slightly
thick-walled.
This beautiful little fungus is easy to recognize from its stature, colour, and small
spores, yet I think it has been mistaken for small states of Clavaria Zollingeri, which
differs in its brittle texture, larger aguttate spores, and secondarily septate hyphae.
Var. pulchella, which Bourdot and Galzin distinguish as the typical state of Clavaria
pulchella, is said to have smooth spores 4-5 X 2·5-3µ., which suggest C. Zollingeri.
Hence I am doubtful of the correct interpretation of the species, for which there is
probably no type. I have followed Coker in considering C. pulchella to represent the
violet fungus with asperulate spores, so nearly allied with Ramariopsis crocea. Clavaria
conchyliata (of England) has such spores and so does a specimen of Bresadola's, named
by him as C. pulchella, according to Coker (Baria, Nice, spores 2·2-3µ., minutely
rough).
Bourdot and Galzin distinguish C. pulchella var. Bizzozeriana (1 cm. high; spores
2-3·5µ., smooth: basidia 15-18 X 3-4µ.) from var. asperula (1-2 cm. high, branches
dichotomous; spores 2·8-4µ., asperulate: basidia 12-22 x 4-4·5µ.). I doubt if there is
any difference save, perhaps, in the size of the spores. I find the spores of English
RAMARIOPSIS
specimens to be 4-4· 5 X 3-3·5µ, with basidia 20-30 X 6-7µ, and Bresadola gives the
spores as 4 X 2·5-3 µ and the basidia as 25 X 5-6µ. Rea gives the spores as 2·5-3 µ,
subglobose, and the basidia as 6-7 µ wide. Cotton and Wakefield give the same spores
and the basidia as 15-18 x 3-4µ. Coker gives the spores of C. exigua as 2·5-3·5µ,
).
subglobose, and the basidia as 15-18 X 3-4µ; for the co-type of C. tenuissima Sacc.,
at Kew, he gives the spores as 2·5-3·5 µ. These measurements agree with those of
Bourdot and Galzin. They differ from mine in referring, probably, to the width of
the spores, which are distinctly longer than broad, and to immature basidia.
TEXT-FIG. 276. Ramariopsis pulchella: spores, basidiurn,

clavate branches, and crystals of ,the medullary hyphae,
X I,O<?O,
C.· Cavarae (spores 4-5µ) is indistinguishable in its description from this charac
teristic species.
Fawcett's record of C. Bizzozeriana from Victoria may be a small state of Clavaria
Zollingeri with unusually small spores. 1
R. tenuiramosa spec. nov.
Text-Fig. 277.
-3·5 cm. high, wholly pale yellowish drab or pale straw-colour, slender, sparingly
branched, solitary or gregarious, slightly
tough and pliant.
Stem 3-10 X 1 mm., generally distinct,
slightly white tomentose at the extreme
base: branches o·5-1 mm. wide, slender,
few, 2-3-chotomous below, dichotomous
above, branched one to three times, rather
elongate, ascending, subdivaricate, terete,
subacute, axils round: without smell or
taste.
Among pine-needles under Pteris in a
wood of Pinus sylvestris: England (High
TEXT-FIG. 277. Ramariopsis tenuiramosa, X r.
Spores, X 1,coo. Wycombe, Bucks., Sept. 1924).
Spores 3·5-4·5 X 3-3·5µ, white, minutely
verruculose or subechinulate, subglobose, 1-guttate.
Basidia and hyphae as in R. Kunzei.
Hymenium not thickening: subhymenium very slightly developed. )
1
-3·5 cm. high, solitary or a few together, slate-violet or bluish, becoming greyish lavender
and avellaneous with age: stem slender, often white pruinose below: branches dichotomous,
open, cylindric, relatively long, sparingly divided, becoming slightly incurved and prong-like,
axils rounded: flesh concolorous and pliable: spores 2· 5-3·5 µ, white, smooth, globose, minutely
apiculate: on bare ground or among moss, Victoria. (Proc. Roy. Soc. Viet. 51, 1939, 12,
t. 3, f. r.)
RAMARIOPSIS
This species differs from R. Kunzei in its colour, which is never white, in its small
size, scarcely tomentose trunk, and simple hymenium, and from R. clavuligera in its
smaller, subglobose spores. It suggests a pale, slender state of R. asterella of the
United States. Possibly R. Kunzei forma subasperata is identical.
R. vestitipes (Pk.) comb. nov.

Basinym: Clavaria vestitipes Pk., Bull. N.Y. St. Mus. II6, 1907, 34.
Synonyms: Clavaria bicolor Pk., ibid. 54, 1902, 954 (haud C. bicolor Mass. 1901 =
Clavaria Zollingeri).
C. Peckii Sacc. et D. Sacc., Sy!!. Fung. 17, 1905, 196 (haud- C. Peckii Sacc.,
1891 = Clavulinopsis corniculata).
Lachnocladium bicolor (Pk.) Burt, Ann. Mo. Bot. Gard. 6, 1919, 274, t. 5, f. 6.
Lachnocladium vestitipes (Pk.) Burt, ibid. 9, 1922, 67, t. 11, f. 104 (ut L. vestipes).
-2·5 cm. high, branched, gregarious, orange-yellow: stem 1-2 mm. thick, slender, pale
yellow, tomentose, divided into 2 or more short compressed branches themselves
dichotomous once or twice: tips acute, concolorous: flesh rather tough in the stem.
On the ground under Pinus: U.S.A. (N.Y., rare).
Spores 3·5-4·5 X 3-3·7 µ,, white, minutely asperulate, subglobose or ovoid.
Basidia 4 · 5-5 · 5 µ, wide: sterigmata 4: hymenium 20 µ, thick.
This seems identical with R. crocea, but Coker says it can be distinguished by the
tomentose tough stem, larger size, and slightly larger spores. Coker gives the stem of
R. crocea as minutely furfuraceous. Wehmeyer gives the fruit-bodies of R. crocea as
reaching 2·5 cm. high (1·5 cm., Coker). Coker gives the flesh of R. crocea as tender
but elastic, not brittle. And the difference in spore-size is about o· 5 µ,.
SCYTINOPOGON Singer
Lloydia 8, 19j..5, 139.
Fruit-body with flattened branching in one plane, becoming confused by twisting of
the branches: branches flattened, more or less broadly dilating before dividing, becoming
elongate and ligulate, polychotomous below, dichotomous above (or wholly dichoto
mous in slender specimens): hymenium inferior, the uppersides of the branches and
axils with a sterile hymenium: flesh rather tough, fibrous, pliant, more or less coria
ceous: white, cream, alutaceous, tan, or tinged pink or purple.
Terrestrial: 2 spp., tropical and ip. the southern states of U.S.A.
Spores white, or pale tan or yellowish, echinulate or verruculose, angularly ellipsoid,
the wall slightly thickened, aguttate, non-amyloid.
Basidia small, narrow: sterigmata (2-3-)4.
Cystidia none.
Hyphae monomitic, clamped, not inflating or very irregularly, often becoming thick
walled, colourless.
Type-species: S. angulisporus (Pat.).
This genus was made for Pterula pallescens Bres., which is a synonym of Clavaria
angulispora Pat. The peculiar branching and the angular-echinulate spores separate
it from Ramariopsis, which it resembles superficially. Its alliance seems with Thele
phora s. str. Aphelaria has smooth spores, branching in all directions, and, typically,
no clamps. If the branches of Scytinopogon are carefully untwisted and flattened out,
it can be seen that they have arisen from the repeated expansion and subdivision of
the original plate of tissue forming the stem and that, if they did not twist by unequal
(? residual) inflation of the tissue, they would form a flabelliform frond as in many of
-
the Rhodophyceae.
SCYTINOPOGON
KEY TO THE SPECIES OF SCYTINOPOGON
More or less suffused lilac or pale purple:
Hyphae variously inflated -12µ, wide: trop. Asia S. echinosporus
Chalk white, often tinged cream or pale tan:
Hyphae mostly 2-4µ, wide, rarely -10µ, m places: tropics
generally and N. Amer. . S. angulisporus
S. angulisporus (Pat.) comb. nov.

Basinym: Clavaria angulispora Pat., Bull. Soc. Myc. Fr. 4, 1888, 41, t. 13, f. 4:
Coker, Clav. U.S. Can. 1923, 103, t. 30, 84.
TEXT-FIG. 278. Scyti11opogo11 angulisporus: stout,

typical form, deeply immersed in the ground (black),
the branches still young and 'Thelephoroid', 'x 1.
Synonyms: ? Clavaria guarapiensis Speg., Fung. Guar. Pug. 1, 1883, n. 83.
? Clavaria Spegazzinii Sacc., Sy!!. Fung. 6, 1888, 707 ( = C. cladonia Speg., Fung.
Guar. Pug. 1, 1883, n. 84, non Kalchbr.).
SCYTINOPOGON
? Lachnocladium cirratum Pat., Journ. Bot. 1889, 167.
? Lachnocladium Schweinfurthianum P. Henn., Engl. Jahrb, 17, 1893, 21.
Pterula pallescens Bres., Bull. Soc. Bot. Belg. 1899, 157: Killerm., Nat. Pfl. Fam.
2nd ed. 6, 1928, 157, f. 102j.
Lachnocladium Molleri P. Henn., Hedw. 1897, 196.
Lachnocladium Schwackei P. Henn., ibid. 196.
Lachnocladium Zenkeri P. Henn., Engl. Jahrb. 30, 1901, 42.
Lachnocladium usambarense P. Henn., ibid. 34, 1905, 44.
TEXT-FIG. 279. Scytinopogon angulisporus: fruit-bodies with the branches

displayed, x 1.
Lachnocladium dubiosum Bres. ap. Rick, Brot. 5, 1906, 13, t. 6, f. 3 (without diagnosis):
Ann. Myc. 18, 1920, 50.
Lachnocladium chartaceum Pat., ibid. 5, 1907, 365.
Clavaria nodulosperma Atk., ibid. 7, 1909, 368.
Lachnocladium pallescens (Bres.) Bres., Hedw. 56, 1915, 304.
? Thelephora lactea Pat., Bull. Soc. Myc. Fr. 39, 1923, 47.
Lachnocladium echinosporum Bres. var. microsporum Beeli, Bull. Soc. Roy. Bot. Belg.
58, 1926, 7.
650 SCYTINOPOGON
Scytinopogon pallescens (Bres.) Singer, Lloydia 8, 1945, 139.
[Clavaria Baileyi Mass. ms., in Herb. Kew.]
[Clavaria Sarasinii Cotton ms., in Herb. Kew.]
Lachnocladium geniculatum Lev. (Ann. Sci. Nat. 1846, 154) sensu Lloyd, Myc.
Notes 65, 1921, 1050, t. 1949 (an verum ?): [ = Clavaria geniculata (Lev.) Sacc., Syll.
6, 1888, 708].
Lachnocladium dealbatum sensu van der Bijl, S. Afr. Journ. Sci. 29, 1932, 320,
t. 3, f. 2.
Text-Figs. 5, 278-80, 282, 283.
-20 X 14 cm., solitary, gregarious, or, generally, densely caespitose, palmately and
fastigiately branched from a flattened trunk-like base immersed in the groun(i,, branching
in one plane but twisted, branches and branchlets flattened and narrowly spathulate or
ligulate, the internodes longer distally, the uppersides of the branches and axils sterile
and minutely submentose, the undersides covered by the even waxy hymenium,
chalk-white becoming cream or light cream-tan, the tips white: flesh cream-white, slightly
coriaceous or fibrous-subcoriaceous, pliant, flaccid, tough at the base of the stem: some
times smelling of old ham (Coker), or iodoform (Singer).
Stem 1-5 cm. X 3-4 mm. thick, arising from a tangled mass of thin white mycelial
strands 0·3-2 mm. wide in the soil, dividing palmately before reaching the surface,
sometimes branched from the base, often many stems fastigiate and irregularly connate,
dilated and flattened 1-2·5 cm. wide below the points of branching.
Lower branches 3-6 mm. wide at ground-level, 2-3·5 mm. above, 5-12-chotomous
below, flattened and fanning out 4-15 mm. wide below the points of branching,
twisted, slightly rugulose below the branchings: upper branches 1-2 mm. wide, often
connate laterally, narrowly ligulate, 2-4-chotomous becoming dichotomous, the distal
internodes gradually more elongate 2-5 cm. X 1-1·5 mm.: tips of young active branches
acute, becoming flattened-cuneate or truncate-flabellate before branching, ultimately
subulate or subterete and filiform, o· 5 mm. wide.
On the ground in the forest (perhaps also lignicolous): apparently throughout the
tropics, U.S.A. (N. Car.): Malaya, not infrequent in the lowlands and mountains to
r,300 m. alt.: Uganda, common (as Lachnocladium Zenkeri): Mauritius.
Spores (4·5-)5·5-7 X (2·5-)3·5-4(-4·5)µ, white (? pale tan-ochraceous), ellipsoid,
slightly thick-walled, finely verruculose or echinulate with acute warts or spines 0·3-0·5µ
long, slightly angular (? aguttate).
Basidia 20-35 X 5·5-7µ, short, clavate, projecting -roµ, finely granular-vacuolate:
sterigmata 2-3-4, mostly 3-4, 4-5µ long.
Cystidia none.
Hymenium thickening -300µ, not layered, the old basidia collapsed, developing
1-4 mm. behind the growing tips, covering undersides of the branches to within
3-10 mm. from the ground: subhymenium originally 20-25µ thick, composed of loosely
interwoven hyphae 2·5-4µ wide, thin-walled and not inflated.
Sterile hymenium on the uppersides of the branches and axils 30-50µ thick, loosely
interwoven with scattered sterile basidia, or merely a tangle of loosely interwoven
hyphae 2-4µ wide, without modified ends, and forming a layer 50-200µ thick,
becoming disrupted and appearing subtomentose.
Stem with a thin.or fairly thick felt of 2-4µ wide, loosely interwoven hyphae on the
surface and often, in the upper part, with a more or less collapsed sterile hymenium.
Hyphae monomitic, 2-4µ wide, rarely -6µ wide, occasionally -ro µ wide in parts
of the cells at the base of the stem, clamped, not inflated, with thin or slightly thickened
walls o·5-1·5µ thick at the base of the stem, the cells -250 µ, long, cylindric or of
variable diameter, the clamps sometimes slightly swollen, occasionally branching
from the clamps: the hyphae strictly longitudinal, entwined, compact, without inter
weaving hyphae.
Growing tips of normal construction, the hymenium developing very loosely and
SCYTINOPOGON 651
gradually by the out-turning of the tips of the superficial hyphae: subterminal cells of
the hyphae of the flesh 40-170 µ long on delimitation.
var. curtus var. nov.
Text-Fig. 280.
2-5 · 5 cm. high, much branched but short: stem short, breaking up into several main
branches at ground-level: branches narrow, 1-1·5 mm. wide, flattened 2-5 mm. wide
below the axils, densely palmate-fastigiate: pure white or drab white, yellowish with
age, tips chalk-white.
TEXT-FIG. 280. Scytinopogon angulisporus: A, var. parvus: B, D, var. curtus: c, var. gracilis:
:.< I.
On the ground in the forest: Malaya (Pahang, Fraser's Hill, 1,300 m. alt., 25.11.30:
Kota Glanggi, low alt., 5.12.30).
This is a miniature form.
var. parvus var. nov.

Text-Fig. 280.
-9 cm. high: branches 1-2 mm. wide, flattened to 3-7 mm. wide below the axils,
narrower than typical: white becoming pale cream, tips chalk-white.
On the ground in the forest: Malaya (Pahang, Tembeling 8.11.30, 3.12.30).
This is intermediate between var. curtus and the typical state.

-10 X 2-4 cm., solitary, or subcaespitose, slender: stem 0·5-3 cm. X 2 mm.,
rather slender and elongate, not breaking up into many branches at ground level but
erect and above-ground: branches 1-2 mm. wide, flattened 3-7 mm. wide below the
axils, rather sparingly and laxly branched, sometimes wholly dichotomous, somewhat
spreading, the tips thin and acute: pallid white or drap, becoming cream-white, or
pale flesh-colour downward.
652 SCYTINOPOGON
On the ground in humus, in the forest: Malaya (Pahang, Tembeling 5.11.30,
17.11.30).
This is one of the widespread, variable, and commoner tropical fungi which has
been described as a new species from nearly every country where it has been collected.
It is known from tropical America, Africa, Asia, and Australasia (Queensland, New
Caledonia), and it extends into the south temperate parts of N. America. Coker first
recognized it in the living state and his American species probably belong to var.
gracilis. I have taken the best and most characteristically developed specimens as
typical, though they may not be the nomenclatural type. Either this typical state or
var. curtus occurs as well in the United States, for Coker remarks of his collection 4393
(N. Car.) that the young fruit-bodies widen suddenly into a plate, breaking up along
the margin into a fringe of branches, the plate and fringe being 2 · 5 cm. wide. Perhaps,
also, the flattened-palmate specimens of Ramariopsis Kunzei, collected by Wehmeyer
in Nova Scotia. belong here.
TEXT-FIG. 281. Scytinopogon angulisporus var. gracilis, x r.
Coker reduced C. nodulosperma Atk. and L. dubios11m Bres. as synonyms. P.

pallescens, as described by Bresadola, and later by Singer as Scytinopogon, has the
characteristic spores-by which the genus can at once be recognized-the small
basidia, narrow uninflated hyphae, and flattened branching, so that it must be synony
mous, perhaps, with var. gracilis. Bresadola gives as synonyms L. usambarense,
L. Molleri, and L. Schwackei. The types of L. Zenkeri, C. Baileyi ms., and C. Sarazinii
ms. I have studied at Kew. L. Schweinfurthianum, from Central Africa, is the typical
state of the species with fan-like axils, according to the description: likewise L. zand
baiense, from Java, with rough spores may also be var. gracilis, but as it is said to be
'with setulae', I have kept it in Lachnocladium. L. chartaceum (Brazil, lignicolous) is
indistinguishable from var. parvus: Coker gives the spores of the type as 6 X 4µ,,
asperulate. Similarly L. echinosporum var. microsporum (Belgian Congo, lignicolous),
is a slender form of S. angulisporus: it has typical echinulate, white spores 5-6 X 3-3·5µ,:
(L. echinosporum, itself, is Ramaria Zippelii).
Thelephora lactea (Indo-China) is clearly a young state of this species. The larger
size of the spores may refer to overall measurement of the spines, and the statement
SCYTINOPOGON 653
'very fragile' to the condition of the dried specimen which becomes very brittle. The
description is:
Thelephora lactea Pat.: very fragile, entirely white, coriaceous, bushy, erect,
5-10 cm. high, with a common trunk divided several times into flattened, thong-like,
triangular lobes; hymenium inferior, smooth: spores 8 X 6 µ, white or pale straw,
angular, echinulate: basidia 18-20 X 10 µ, clavate, with 4 short sterigmata: var.
obscura Pat., 2-3 cm. high, with brownish tips: on sandy paths, Cambodia: 'Analogue
a Phylacteria se rattache a Thelephora par sa trame blanche et ses spores non colorees.'
0 G.. . @·...:.. o· u��} �, �

(,:.'
,
'' "' � ,•l
\.•
/;;:.t
TEXT-FIG. 282. Scytinopogon angulisporus: spores, incipient hymenial hyphae, and old
excrescent hymenial hyphae, X 1 ,coo.
L. geniculatum sensu Lloyd (Philippine Is!.) is also S. angulisporus, possibly var.

parvus, but whether it is true L. geniculaturn Lev. (S. Amer.) can be decided only by
reference to the type. If so, it supplies the earliest specific epithet. Leveille's descrip
tion is:
5-6 cm. high, branched from the base, branches somewhat terete, dichotomous,
divaricate, light yellow, axils compressed, tips acute, crowded: on the ground, Para
meribo (leg. Kegel).
L. cirratum Pat. is perhaps a variety with long slender branches. Its spores, how
ever, are said to be subangular and not echinulate or verrucose: they may l:,e immature.
Its description is:
8-15 x 8-10 cm., caespitose: stems 3-5 x 1-2 mm., divided .almost from the base
into many, slender, subcylindric branches, repeatedly divided with long acute tips
(circinate-intricate, ? the effect of drying): branches white and pulverulent on one
side, ochraceous and glabrous on the other: spores 4 x 3 µ, subhyaline, ovoid-rotund,
smooth or subangular ('non echinulatis, nee verrucosis'), 1-guttate: on the ground,
French Guiana.
There are no descriptions of the spores for Spegazzini's two species, which must be
compared with Clavaria sparassoides (p. 267). Their main points are:
C. guarapiensis: white (drying cinereous): arising from a fibrous floccose mycelium
pervading and embracing particles of the substratum: stem simple or 3�4 partite from
the base: primary and secondary branches 2 cm.X3 mm., 4-13 partite: ultimate
branches with bifid tips: in humus in woods, near Guarapi, S. Amer.
C. spegazzinii: white (drying greyish white), arising from a fibrilloso-floccose
mycelial nodule, ramoso-partite from the base: primary branches 3 cm. X I mm.,
SCYTINOPOGON
straight or somewhat flexuous: branchlets laciniately divided, the tips 2-3 X 0·3-0·4 mm.:
on the ground in woods near Guarapi, S. Amer.: said to resemble Cladonia rangiferina.
The earliest collection of S. angulisporus seems to be 'Pterula compressa Kl.' (ined.),
of which there is a co-type from Mauritius in the Kew herbarium.
I'
/·:···
c·
!
A A
TEXT-FIG. 283. Scytinopogon angulisporus: A, thick-walled
hyphae from the base of the stem: the remainder being ordinary
uninflated medullary hyphae, X 500.
Spores. In preserved specimens, dried or in alcohol-formalin, the spores may be

hyaline-white or with very pale-yellowish or pale-tan walls, as seen under the micro
scope, but I have not seen a fresh spore-print. Singer gives them as yellow, but again
it is not clear if the colour refers to fresh or dried spores, and it is well known that
white spores may become yellowish on drying, as in Clavulina, Lachnocladium, and
Clavariadelphus. On the other hand, Clavaria durbana v. d. Bijl · (Natal) may be
synonymous and it is said to have pale-ochraceous spores: (I have kept the species,
SCYTINOPOGON 655
therefore, in Ramaria). There are also considerable variations in the size and markings
of the spores in different collections. Coker gives the range as 4·2 X 2·5 to 7·5 X 5µ, for
N. American specimens. The Malayan collections are more regular, 5·5-7 x 3·5-4·5µ.
In the African collections of L. Zenkeri at Kew they are mostly 5-6 x 3µ. Some
collections have very minute spines which appear as little more than roughenings of
the epispore: others are distinctly echinulate.
S. echinosporus (B. and Br.) comb. nov.

Basinym: Clavaria echinospora B. and Br., Journ. Linn. Soc. Bot. 14, 1875, 75
(haud Boud. et Pat. = Clavulinopsis helvola): emend. Petch, Ann. R. Bot. Gard. Per.
9, 1925, 336.
Synonym: Clavaria implexa Lev. sensu v. Ov., Bull. Jard. Bot. Buit. Ser. 3, 5, 1923,
272: le. Fung. Mai. 1, 1923 (haud Lev.).
-10 X 7 cm., branched, in more or less depressed globose tufts arising from a common
stem or cluster of stems, pallid or brownish below, pale purple or lilac above (Petch),
pale dingyflesh-colour to dingy whitish, even slightly violet, tips dark violet-brown (v. Ov.),
black on drying (Petch): stem -10 X 3-5 mm., short, cylindric or subcompressed
(? from mutual pressure), arising from numerous white mycelial strands: branches
slender, 2·5 mm. thick below, 0·5-1 mm. above, repeatedly dichotomous, the outer
curved, ascending with wide, rounded axils, the inner with acute axils, crowded or
rather loose, ultimate branchlets long, cylindric, tapering, simple and subacute, bifid
with semilunar axils (v. Ov.), or even flattened and cristate (Petch): flesh white, rather
tough, subcoriaceous (v. Ov.), cartilaginous (Petch).
On the ground among grass: Ceylon, Java.
Spores 4-7 X 3-4µ (Petch), 5-6·5 X 2·8-4µ (v. Ov.), white, coarsely echinulate with
short broad-based spines or coarsely tuberculate nodular, ovoid (Petch); finely
verrucose-rough, ellipsoid (v. Ov.): angular and echinulate with spines 0·2-0·5 µ long
(Corner).
Basidia 22-24 >< 5·5 µ: sterigmata 4, short (v. Ov.).
Hyphae irregularly inflated in places -12 µ wide, mostly rather narrow, thin-walled,
clamped, some with oily contents.
I have not found this in Malaya, though it almost certainly occurs, but I have
examined the type and a later collection of Petch's at Kew. As Petch suggested,
C. implexa sensu v. Ov. is certainly the same as the Ceylon fungus and I agree that
Leveille's original description is too vague for accurate identification.
S. echinosporus is very near the more slender forms of S. angulisporus, but the fruit
bodies seem more distinctly coloured and the more inflated hyphae, often with oily
contents, seem different. Their spores, however, are identical.
TYPHULA Fr., emend. Karst.

Fries, Syst. Myc. 1, 1821, 494: Karst., Finl. Nat. o. Folk 37, 1882, xvii et 181:
Killerm., Zeitschr. f. Pilzk. 18, 1934, 98-108, 137-9: Remsberg, Mycologia 32, 1940,
52-96: Vang, Medd. Plantepat. Afd. Kg!. Vet. Landb. Koh. 28, 1945, 1-45.
Synonyms: Typhula subgen. Phacorrhizae Fr., sensu auctt. (non Leptorrhizae
Fr. = Pistillaria).
Phacorrhiza Pers., Myc. Eur. 1, 1822, 192: Greville, Scot. Crypt. FI. 1, 1823, t. 43.
Pistillaria Fr. sensu Donk, Rev. Nieder!. Homobas. Aphyll. 2, 1933, 100: (non
sensu auctt.).
Fruit-bodies 1-100 mm. high, simple, spuriously branched in a few species, very
small to elongate, slender, with a filiform sterile stem, arising from a small sclerotium, and
a subglobose, clavate, cylindric, or elongate filiform fertile head: texture rather horny or
cartilaginous, especially in the stem, to waxy-firm or soft, especially in the head.
656 TYPHULA
On dead plant-remains, not humicolous or terricolous (? not truly lignicolous,
though corticolous), saprophytic or parasitic: N. temp.(? S. temp.), absent from the
tropics: c. 40 species .
. , ',
I,
l'.' i
' ·. . �
1, 1,
a.
TExT-FIG. 284. Fruit-bodies of Typhula: a, T. variabilis: b, T. incarnata (= T. Itoana):

c, T. intermedia. X 5: (from Remsberg).
Spores white, smooth, ellipsoid, thin-walled, aguttate, or with 1-2 minute guttulae.
Basidia with 2-8, mostly 4, sterigmata.
Cystidia present only in T. cystidiophora (so far as known).
Hymenium not thickening, covering the head on all sides or absent from the
growing apex: subhymenium varying well-developed to almost absent.
TEXT-FIG. 285. Fruit-bodies of Typhula: a, T. gyrans: b, T. umbrina: c, T. pertenuis: d, T.
subulata: e, T. latissima: f, T. idahoensis: g, T. virgata: h, T. 1Jibumi: i, T. sphaeroidea:
j, T. athyrii. X 5: (from Remsberg).
5119
uu
658 TYPHULA
Caulocystidia generally present, scattered.
Hyphae monomitic, inflating, primarily clamped, though the clamps absent from the
fruit-bodies in some species, often incrusted with crystals especially in the sclerotium,
on the surface of the stem, and in the subhymenium, not secondarily septate, occa
sional uninflated hyphae with resinous oleaginous contents: hyphal walls often more or
less firmly or toughly mucilaginous, the superficial hyphae of the stem with thickened
agglutinated walls forming a firm peripheral cylinder I-2 hyphae thick.
\r c..
TEXT-FIG. 286. Surface views of sclerotia: a,

T. idahoensis: b, T. incarnata (= T. Itoana): c,
T. athyrii: d, T. variabilis: e, T. umbrina: J, T.
viburni: g, T. phacorrhiza: h, T. gyrans: i, T.
intermedia. X c. 250: (from Remsberg).
Sclerotium 0·5-6 mm. wide, small, globose or more or less flattened, generally
yellow, brown, or black, with a cortex of agglutinated hyphae and a dark-coloured
cuticle formed from the outer hyphae walls, or with an epidermis of narrow hyphae:
medulla composed of free or agglutinated hyphae.
Lecto-type: T. phacorrhiza.
The association of the sclerotium-bearing species in Typhula gives a natural and
interesting assemblage, as it is the one specialization of Clavarioid fungi: indeed, it
seems the only new idea in the family other than the universal degeneration of the
fruit-body. Practically, Typhula becomes important because several species attack
crop-plants and, now that they have been grown in artificial culture, they may become
subjects of physiological, cytological, and genetical research. It is one of the few
genera of higher fungi which impinges on laboratory science, so it behoves systematists
to assist with accurate classification. Unfortunately, except for Remsberg's researches
in the United States, the genus has been much neglected and its systematy is super
ficial and inexact. Already, misnomers are creeping into plant pathology and con-
TYPHULA
fusing the issue of long investigations, than which there can be nothing more aggravat
ing or disheartening: and to avoid this fate, I would advise investigators to describe
in full the species which they study, for the extra labour will dispel the shadow of
doubt. Little now can be gathered about the genus from the limbo of the past, but
from living material there will be plenty. No species has been recorded from the
south temperate regions, or from the vast 'temperate' area,s of tropical and subtropical
mountains, though from the tropical lowlands the genus seems truly to be absent.
In the account which follows, I have drawn greatly on Miss Remsberg's paper,
which may not be accessible to many mycologists, and, if I have altered or criticized,
it is for consistency in general description or for clarification of doubtful matters:
I have studied her work as the beginning of the scientific organization of the genus.
My own researches have been limited to a few European species which I collected
mainly between 1925 and 1928, namely T. phacorrhiza, T. erythropus, T. graminum,
T. gyrans, T. sclerotioides, and T. quisquiliaris ( = Pistillaria quisquiliaris). But of all
genera of fungi, I find this the happiest, for it recalls in the wearying tropics most
vividly the crisp smell of autumn afternoons and sodden leaves thronged with little
fungi.
NATURAL CLASSIFICATION OF TYPHULA1

Subgenus Phacorrhizae Fr.
Sclerotial hyphae wholly agglutinated, (he medulla horny-cartilaginous (occasionally
with free hyphae in the centre): longitudinal hyphae of the fruit-body rather toughly
mucilaginous with more or less agglutinated walls at least in the stem, the texture
horny-cartilaginous (especially in the stem) to waxy-horny or submucilaginous
(especially in the head).
T. euphorbiae, T. gyrans, T. incarnata, T. phacorrhiza, T. sphaeroidea, T. viburni.
Subgenus Subcarnosae subgen. nov.
Medullary hyphae of the sclerotium not agglutinated, often thick-walled or incrusted,
the texture floccose or spongy-coriaceous: cortex of sclerotium with cuticle but no
epidermis: hyphae of the fruit-body agglutinated only in the superficial layer on the
stem, the inner hyphae of the stem with dry or thinly mucilaginous walls: texture of
the head waxy-soft.
T. athyrii, T. erythropus, T. graminum, T. idahoensis, T. intermedia, T. latissima,
T. pertenuis, T. sclerotioides, T. subulata, T. trifolii, T. umbrina, T. variabilis, T.
virgata.
Phacorrhizae
Head filiform-cylindric, elongate, the tip often persistently sterile:
subhyrnenium slight (? in T. incarnata): sclerotium with a
cuticle, without epidermis
2 · 5-10 cm. high, white, then cream, pale ochraceous, or brownish:
sp. r r-20 X 4-8 µ,: sclerotium superficial, attached by a minute
red-brown stalk, pyriform or flattened on both sides: N. temp. T. phacorrhiza
1-4 cm. high, white then pinkish with white stem: sp. 8-15 X
3-8 µ,: sclerotium erumpent, subglobose or convex above,
flattened or concave beneath: N. temp. T. incarnata
Head shortly cylindric, clavate or subglobose, soon fertile over the
apex: subhymenium well developed: sclerotium without cuticle
but with an epidermis of 2-5 layers of hyphae with thick brown
ish walls (? in T. euphorbiae and T. quisquiliaris)
1
Only those species in which the hyphal construction of the_.sclerotiun1,. as well as the
fruit-body, is adequately known can as yet be placed in this arrangement. An Artificial Key
is gin•n on p. 662.
660 TYPHULA
Head shortly elongate, 1-6 x 0·3-1·5 mm.: spores mostly less
than 11µ, long
Rather stout: head 0·5-1·5 mm. wide: stem 0·1-1·2 mm. thick,
white then brownish at the base: sp. 8·5-11·5 x4-6 µ,:
sclerotium 0·5-3 mm. convex-plane, brown to black, the
centre often not agglutinated: on dead leaves and stems:
Europe, N. Amer. T. gyrans
Head shortly clavate, ovoid or subglobose, 0·8-3Xo·5-1·5 mm.
(? with well-developed cortex)
Sp. 7·5-10X3·5-5·5µ,
Head I mm. wide, subglobose: stem 4 mm. long, brown:
sclerotium lenticular, yellow: on stems of Euphorbia:
Europe . T. euphoroiae
Head 1·2-2X0·5-1 mm., ellipsoid to ovoid, white to pale
brown: stem 4 Xo·1-0·2 mm., white, brownish at the
base: sclerotium 0·5-1 mm., convexo-plane, rich brown:
on leaves of Viburnum: U.S.A. . T. viburni
Sp. 11µ, or more long
Head 1-3Xo·4-1·2 mm., ovoid, clavate, transparent white:
stem 1-10Xo·1-0·3 mm., white: sp. 11-15 x5-6 µ,:
sclerotia 1-3X0·3-0·4 mm., oblong fusoid then flattened,
immersed, ochraceous to dark brown: on Pteris petioles:
Europe . T. quisquiliaris
Head 0·8-1·5Xo·5-1·5 mm., ovoid to subglobose, chalk
white (? opaque): stem 1-2 x0·3-0·5 mm., white, base
reddish brown: sp. 11·5-13·5 x6-9µ,: sclerotium 0·5-1
mm., convexo-plane, brown then black, somewhat rough:
on stems of Rubus: U.S.A. T. sphaeroidea
(Compare T. elegans; sp. 10-15 X5µ,, on Viburnum: U.S.A.)
Subcarnosae
Cortex of sclerotium composed of several layers of agglutinated
hyphae: sp. 5-9 X2-4·5 (? -5) fL
Head 1·5-5(-10)X0·2-0·4 mm., white: stem 5-35 X0·1-0·3 mm.,
white: sclerotium 0·5-5 mm., ochra.ceous then brown or
blackish, the cuticle 5-8µ, thick: on dead herbaceous remains :
Europe, N. Amer. . T. sclerotioides
Cortex composed of a single layer of agglutinated hyphae (? 1-3 in
T. umbrina): spores mostly 9-16 X4-8 µ (smaller in T. erythropus)
Head elongate-cylindric or subulate, generally more than 8 mm.
long, the apex often persistently sterile: sclerotia 1-6 mm.
wide, cuticle 10-15µ, thick (7-10µ, in T. pertenuis)
Fruit-body white, then fuliginous, often yellowish or oliva-
ceous, especially the stem: head 4-28 x0·4-3 mm., often
becoming hollow: stem 5-65 x0·2-1 mm., long: sclerotia
1-6 mm.: on dead herbaceous remains: Europe, N. Amer. T. variabilis
Fruit-body drab white, the stem becoming brownish: head
8-13Xo·5-2 mm., conic-subulate: stem 3-5Xo·r-0·2 mm.:
short: sclerotia 2-3 mm.: on dead leaves and grasses: Canada T. subulata
TYPHULA 661
Fruit-body white
Head 5-20 X 0·5-2 mm.: stem 5-15 X 0·3-1 mm., weak,
rarely erect: sclerotia 1·5-3·5 xo·5-1 mm., ri,ch brown
then black, drying rough and dull (with microscopic
rosettes of tubercles on the cuticle): on dead leaves and
stems: Europe, U.S.A. T. intermedia
Head 8-15 X 0·5-3 mm.: stem 4-10 X 0·5-1·5 mm.: sclerotia
2-3 mm., bistre to blackish: on bare soil: U.S.A. T. pertenuis
(Sclerotia o·5-0·8 mm.: head 5-11 mm.: T. virgata, below.)
Head shortly cylindric to subclavate 0·5-8 mm. long (-11 mm.

in T. virgata), the apex soon fertile: sclerotia mostly small
Sp. 6-9(-11) X 2·5-3·5(-4)µ.
Stem reddish brown or black, head white: sclerotia
0·5-2 X 0·4-1 mm., reddish brown or black, lenticular: on
dead leaves, twigs, stems, &c.: Europe T. erythropus
Sp. 9-16 X 3·5-8µ: stem not reddish brown
Fruit-body brown, or white becoming brownish: sclerotial
cuticle 7-20 µ thick
Fruit-body brown with darker stem: head 4-7 Xo·5-1·5
mm.: stem 2-5 X 0·1-0·5 mm.: sclerotium 0·5-2 mm.,
the rind 5-20µ thick: basidia with 4-6-8 sterigmata:
parasitic on grasses (Triticum, Agropyrum): U.S.A. . T. idahoensis
Fruit-body white then brownish: sclerotium 2-4 mm.
wide: head 3-8 Xo·8-2 mm.: stem 3-10 Xo·1-0·5 mm.:
basidia with 4 sterigmata
Head regular, scarcely flatt�ned: stem becoming brown
ish or blackish at the base: cuticle of sclerotium
7-12µ thick: on turnips (Brassica) and leaves and
rhizomes of Iris: Canada T. umbrina
Head often flattened and irregular, broadly elliptic to
clavate: sclerotial cuticle 10-15µ thick: on dead
stems of Typha latifolia: U.S.A. . T. latissima
Fruit-body white: s�lerotial cuticle 5-10µ thick (? .in T.
trifolii)
Sclerotium tawny then black, oblong, 0·5-1 mm., im
mersed: head o·5-1 mm. long, fusiform ellipsoid: stem
2-4 mm. long, capillary: sp. 10 X 3·5µ: on dead grasses:
Finland, England . T. graminum
Sclerotium 0·5-0·8 mm. wide: head 5-11 X 1-1·5 mm.,
cylindric: stem 2-6 xo· 1-0·2 mm.: on dead herbaceous
stems: U.S.A. T. virgata
Sclerotium 0·5-1·5 mm. wide: head 3-7 Xo·5-2·5 mm.,
subclavate, often branched: stem 2-5 X o·5-r mm.:
on wintering stocks of Athyrium angustum: U.S.A. T. athyrii
Sclerotium 1-2 mm., rough (? medullary hyphae thin
walled): head 2-7 mm. long: stem 3-10 mm.: on
various leguminous herbs : Europe T. trijtJ[ii
662 TYPHULA
ARTIFICIAL KEY TO THE SPECIES OF TYPHULA
Head elongate, cylindric, filiform or conic-subulate, often per-
sistently sterile at the apex, 10 mm. or more long . A
Head shortly cylindric, ellipsoid or subclavate, 0·5-10 mm. long . B
Head subglobose, obovoid, or shortly and broadly ellipsoid,
0·5-3 mm. long and nearly as wide C
A. (Head elongate)
Sp. less than 8 µ. long
With cystidia: head 20-50)< 1-1·5 mm., white or yellowish:
stem 20-25Xo· 5-1 mm.: sp. 5-6· 5 X 2· 5-311.: Oregon . T. cystidiophora
No cystidia
Europe: on roots and leaves of Beta: sp. 6-8x2·5-3·5µ T. betae
(Argentine: on Bignonia-leaves: ? no stem: sclerotium very
small (?): sp. 6-8X3-4µ Pterula
tenerrima)
Sp. larger
Pale ochraceous, 2·5-10 mm. high: N. temp T. phacorrhiza
Pink with white stem: N. temp. T. incarnata
Greyish: N. temp. T. variabilis
Drab white, with brownish stem: conic-subulate: Canada T. subulata
White
Sclerotium rough with microscopic rosettes of tubercles:
weak, rarely erect: sp. 12-16x4-8µ: N. temp. T. intermedia
Sclerotium without microscopic rosettes
Sp. 8-1ox5-6 (or 8-14X2-5)µ: Europe T. sclerotiicola
Sp. 8-12·5X4-6µ: U.S.A. T. pertenuis
Sp. 11X5 µ(?):head white, then fuscescent to 23 mm. long:
stem -35 mm. long: on Rubus: Germany T. Laschii
B. (Head intermediate)
Sp. 9-15 µ. long
Yellow: sp. 9-12X3-4µ: on petioles of Fraxinus: France T. lutescens
Greyish: head 2-3 mm. long: sp. 9-10X4µ: France T. semen sensu
Bourd. et
Galz. (p. 687)
Brown, reddish brown, or white, becoming brown
Sterigmata 4-8: brown with darker stem: on grasses: U.S.A. T. idahoensis
Sterigmata 2 -
4
Reddish brown: on leaves of Brassica: Japan . T. tochinaiana
White then brownish
Head regular, scarcely flattened:stem becoming brownish
or blackish at the base: on Brassica and Iris: Canada T. umbrina
Head often flattened and irregular, broadly ellipsoid to
clavate: on Typha: U.S.A. T. latissima
White
On Athyrium: U.S.A. T. athyrii
TYPHULA 663
On Leguminosae: Europe T. trifolii
On various herbaceous stems or grass-leaves
Head 5-II X 1-1·5 mm., cylindric: U.S.A. T. virgata
Head 2-5 X 0·5-1·3 mm.: N. temp. T. gyrans
Head 0·5-1 X 0·15-0·2 mm.: on grasses: Europe T. graminum
Sp. smaller
Sp. 3-3·5 X 1-2µ,: white, branched: on bark: France T. ramealis
Sp. 4-9µ, long (-IIµ, in T. erythropus)
Stem becoming brown or blackish
3 mm. high: sp. 4-5 X 2 µ,: France T. Patouillardii
Larger: sp. 5--<) µ, long
Stem pinkish red, then reddish black: Europe T. erythropus
Stem violaceous black, 15-20 mm. long: France T. neglecta
Stem white, then pale rufescent, 3-5 mm. long: head
ochraceous: on leaves of Olea: N. Africa . T. pallens
Stem fuscous below, 10-16 mm. long: head very small,
white: on leaves of Buxus: Spain T. buxi
Wholly white
Fruit-bodies over 10 mm. high
10-12 fruit-bodies per sclerotium: 2 sterigmata: on bark
of Castanea: France T. caespitosa var.
coacervata
Fruit-bodies solitary: 4 sterigmata: on twigs and her-
baceous remains, N. temp. T. sclerotioides
Fruit-bodies smaller, -5 mm. high: sterigmata 2
Simple or two to three times forked: on stems of Mentha:
France T. corallina
Simple: stem minutely bulbous: sclerotia often with
minute, discoid conidiophores: on stems of Eupati>-
rium: France T. bulbosa
Sp. unknown
Azure blue, 6-12 mm. high, many per sclerotium: on leaves of
Juglans: Italy T. caespitosa
Reddish purple to bay brown, -5 mm. high: on stems. of Gen-
tiana: France T. Muelleri
Stem blackish brown: head acuminate, linear, white or pink:
-15 mm. high: on leaves of Fraxinus: Germany . T. crassipes
White
Head oblong, blunt: stem very short: 4 mm. high: on leaves of
Hedera: Italy T. hederaecola
C. (Head subglobose or ovoid)

Sp. more than 10 µ, long
1 mm. high: head 0·6Xo·5 mm.: on twigs of Viburnum: U.S.A. T. elegans
Larger
Head 1-3 mm. long: on Pteris petioles: Europe . T. quisquiliaris
Head 0·8-1·5 mm.: on stems of Rubus: U.S.A . . T. sphaeroidea
TYPHULA
Sp. mostly less than 10 µ, long
On twigs of Abies and Pinus: yellowish: Europe T. abietina
On stems of Euphorbia: stem brown: Europe T. euphorbiae
On leaves of Populus: white: Europe T. ovata
On leaves of Viburnum: white, then brownish: U.S.A. T. viburni
Sp. unknown: white
On leaves of Carex: Finland T. caricina
On stems of Osmunda: U.S.A. T. filicina
T. abietina (Fckl.) comb. nov.

Basinym: Pistillaria abietina Fckl., Symb. Myc. 1, suppt., 1871, 4: Bres., Ann.
Myc. 1, 1903, II3: Killerm., Zeitschr. f. Pilzk. 18, 1934, 106, f. B.
Head 1-1·5 Xo·4-0·5 mm., yellowish then straw-colour (brownish, drying white:
Killerm.), obovoid or compressed-spathulate: stem 1-4 X0·2-0·35 mm., filiform,
diaphanous, glabrous.
d.
h.
TEXT-FIG. 287. Sections of the sclerotial wall: a, T. phacorrhiza: b, T. gyrans: c, T. incarnata

(= T. Itoana): d, T. sphaeroidea: e, T. variabilis:f, T. umbrina: g, T. viburni: h, T. intermedia
(the rind gelatinous-tough). X c. 150: (from Remsberg).
On twigs of Pinus and Abies: Europe (Germany, Poland, France).
Spores (6·5-)8-10·5 X 4·5-5 µ,, white, smooth (Donk, for type-collection); 9-11 X
4-6 µ, (Bres.); 7-8 X4µ,, 1-guttulate (Killerm.).
Basidia 32-40 X5-6·5µ,, with 2-3 sterigrnata 5µ, long (Donk): 30-35 X6-8µ, (Bres.):
2op, long, with 2 sterigmata 1op, long (Killerm.).
Hyphae 3-5 p, wide, thin-walled (Donk, Bres.).
Sclerotium 2-3·5 mm. wide, flattened, smooth, irregularly lobed,. brownish, under
the bark (Donk): 2-5 mm. wide (Bres.).
Possibly this is not uncommon. Donk has supplied me with his notes on the type
collection (Fckl. 24, in Herb. Oudem.). The descriptions of Bresadola and Killermann
agree well but suggest considerable range in spore-size, dependent perhaps on the
number of spores on the basidium.
TYPHULA 665
Compare Pistil/aria lignicola, which may be the sclerotium-free form or an allied
species.
T. athyrii Remsberg
Mycol. 32, 1940, 91, f. 27, 28, 31.
Text-Figs. 285j, 286 c.
4-15 mm. high, white, yellowish with age or on drying, clavate, erect, straight or
slightly curved, blunt, simple or frequently much branched, 1-several per sclerotium:
head 3-7 xo·5-2·5 mm., cylindric, straight or curved, solid, apex fertile: stem
2-5 X 0·5-1 mm., puberulous, gradually thickening into the club.
On overwintering stocks of Athyrium angustum, fruiting in autumn: U.S.A.
Spores 8-13X4-6 µ., av. II x4 µ., fusiform-ellipsoid, apiculate.
Basidia 30X 8 µ.: sterigmata 4.
Hyphae of head and hymenium incrusted, not those of the stem.
Sclerotium 0·5-1·5 x0·5-0·7 mm., globose or elongate, convex above, flat or concave
below, argillaceous to fulvous olive or brownish, erumpent, simple or (in culture)
coalescing: cuticle 7-10 µ. thick, golden yellow, rough: cortex composed of one layer
of hyphae with wide lumen: medulla composed of free, interwoven, incrusted hyphae,
loose in the centre.
Temperature-range 0-25° C., opt. 18-21 ° C.: sclerotia after 5-10 days in culture:
sterile fruit-bodies not seen in culture.
T. betae Rostr.
Orn. Landbrugets Kulturpl. 2,Q881, 92: Lind, Dan. Fungi 1913, 365: Sacc., Syll.
Fung. 23, 1925, 497= Vang, I.e. (p. 655).
Misidentifications: = T. variabilis sensu Rambousek, Zeitschr. f. Zilkerind. 49,
1925, 197 (R.A.M. 4, 1925, 521, abstract).
= T. variabilis sensu Schmidt, Die Deutsche Ziickerind. 58, 1933, 17 (R.A.M. 12,
1933, 416, abstract).
? = T. variabilis sensu Voglino, La Difesa delle Piante, Torino, 6, 1929, 1 (R.A.M.
8, 1929, 597, abstract).
2-5 cm. high, simple, white: head 1--3 cm. long(? 0·2--0·3 mm. wide), fusoid-
elongate, attenuate into the hairy stem.
On roots and leaves of Beta: Europe (Denmark, Germany, Spain), Azores.
Spores 6-7X2·3-3µ. (Rambousek): (6-7x2·5-3·5µ., Voglino).
Basidia with 4 sterigmata (Rambousek).
Sclerotium black, internally white (Rostrup): 1 ·25-2 mm., globose or ovoid, hyaline
then brown (Rambousek): (2-5 mm., yellow then orange, finally red-brown, globose
or oblong: Voglino).
There appears to be confusion over this species, probably because of the vague
original description. As a parasite on sugar-beet it should be recognizable, and for
this reason I have referred here the species called T. variabilis by Rambousek and
Schmidt. They describe a Typhula parasitic on sugar-beet, macroscopically like a
white T. variabilis, but with small spores like T. trifolii: such a fungus cannot be
T. variabilis, which has more or less fuliginous fruit-bodies and large spores. Schroeter
ascribed small spores, 6-7X 2·5-3 µ., to T. variabilis, so that his fungus may also have
been T. betae. Voglino's fungus was said to grow on potato, asparagus, and sugar
beet: the spores agree with those of Rambousek's fungus, but the sclerotia are con
siderably larger. If my surmise is correct, T. betae and T. cystidiophora are the only
Typhulas known to have the elongate head of unlimited growth and small spures.
As a parasite, T. betae attacks the underground stems and roots, particularly in
low-lying and damp or heavily manured ground. Once established, it quickly rots the
underground parts of the host, covering them with white mycelium and rhizomorphs:
it then attacks the wilted leaves, eventually rotting the whole plant and forming
666 TYPHULA
sclerotia on the surface. Rambousek says, however, that it may be only a saprophyte
on injured tissue and that it damages the host by accelerating decay and attracting
insect pests (see Postscriptum, p. 688).
T. bulbosa (Pat.) comb. nov.

Basinym: Pistillaria bulbosa Pat., Bull. Soc. Bot. Fr. 1885, 45:
Tab. An. n. 473.
Text-Fig. 288.
4-5 mm. high, white, simple: head linear-fusiform, acute:
stem minutely bulbous, pellucidly puberulous: sclerotium brownish,
embossed.
On dead stems of Eupatorium: France.
Spores 6-7X2 µ (5X2 · 5 µ, Donk), white, ovoid, incurved or
TEXT-FIG. 288. Typh attenuate.
ula bulbosa: sclerotia Basidia c. 16X6-8 µ, 2-4 spored (Donk).
with fruit-bodies and Hyphae 2-5 µ wide (Donk).
discoid conidiophores: Conidia 8-10 x 2 µ, cylindric, white, borne on discoid bodies,
(from Pat., Tab. An. 1-2 mm. wide, on the sclerotia.
n. 473).
I have added Donk's ms. notes on the type-specimens.
Compare T. corallina.
T. buxi Maire
Fungi Catalaun. 1933, 28, f. 4.
-18 mm. high, simple: head 1-1·5 mm. long, very short and small, cylindric or
oblong-cylindric, obtuse, shortly attenuate at the base, white then pallid: stem long,
filiform, c. 0·1 mm. wide, cylindric, with long sparse hairs throughout, pallid above,
fuscous at the base, pellucid at the apex.
On fallen leaves of Buxus sempervirens: Spain (in the mountains, Catalaunia).
Spores 6--'7X3-3·5 µ, white, smooth, oblong, pale greyish blue with iodine solution.
Basidia 15-18X5-6µ clavate: sterigmata 4, 3-4µ long.
Caulocystidia -70 µ long, or more, c. 1 µ wide, slightly swollen at the base.
Hyphae of stem 4µ wide, parallel, longitudinal, rather thick walled, clamped,
scarcely incrusted with crystals (absent also from the head).
Sclerotium 0·5-1·2 mm. wide, subglobose or lenticular, smooth, dirty citrine,
erumpent.
This species is distinguished by the yellowish sclerotium, laxly pilose, filiform stem,
small white head, small spores, and absence of crystalline incrustation. T. pallens has
similar spores and hairs, but a darker sclerotium and stem, and abundant crystals.
T. caespitosa Ces.
Bot. Zeit. 1855, 77: Sacc., Syll. Fung. 6, 748.
6-12 mm. high, caespitose, many from a sclerotium, clear azure blue, subclavate, a
little curved, obtuse, sclerotium oblong, convex, brown, erumpent.
On petioles and nerves of Juglans: Italy.
var. coacervata Bourd. et Galz.

Hym. Fr. 1927, 130.
-25 mm. high, Io-20 clubs per sclerotium, subcylindric, obtuse, often slightly thickened
at the apex, simple or with 1-2 branches, white then a little glaucous, drying brown.
On bark of Castanea sativa: France.
Spores 7-9X4-4·5 µ, white, smooth, oblong or elliptic, aguttate.
Basidia 18-27X5-7µ: sterigmata 2, 4-5 µ long.
Hymenium covering the whole head including the apex.
TYPHULA
Hyphae 2-6µ, wide, thin walled.
Sclerotium 2 mm. wide, subglobose, rough, brown.
The variety may not be conspecific. The shape and length of the head is not given.
It is remarkable that so many fruit-bodies can grow from so small a sclerotium.
T. caricina Karst.
Finl. Nat. o Folk 37, 1882, 184: Act. Soc. Sci. Fenn. 15, 1888, 185, t. 6.
Head I mm. long, obovoid, oblong, v.:hite, obtuse: stem 4-5 mm. long, puberulous,
white.
On dead leaves of Carex vesicaria: Finland.
Sclerotium 0·5-1 mm. wide, globose or flattened, white then yellowish, persistently
subepidermal.
This little-known species is said to differ from T. graminum in the thicker, shorter,
obtuse head and the smaller, paler sclerotium always covered by the epidermis of the
leaf. The habitat should enable it to be rediscovered, but I doubt if it differs from
T. graminum.
T. corallina Quel.
Ass. Fr. 1883, 505, t. 6, f. 15.
Synonym: T. nivea Pat., Tab. An. 1883, 23, n. 42 (tide Sacc., Sy!!. 6, 746).
3-5 mm. high, white, simple or shortly and bluntly bifid, flexuous, filiform, blunt,
attenuate to the indistinct stem, glabrous.
On dead stems of Mentha: France (Jura).
Spores 6µ,, white, ovoid-pruniform (Quel.): 5 x 2·5 µ, (Pat.).
Basidia 2-4-spored.
Sclerotium c. 1 x0·5 mm., brown, round or oblong.
'Allied to gracilis and furcata' (Quel.). Compare T. bulbosa and T. sclerotioides..
T. crassipes Fckl.
Symb. Myc., 1869, 32.
-15 mm. high: head acuminate, linear, zchite or pale pink: stem blackish brown,
thickened at the base.
On dead leaves of Fraxinus: Germany.
Spores ovate, hyaline.
Sclerotium globose or elongate, fuscous black.
Compare T. erythropus and T. neglecta.
T. cystidiophora Kauffm.
Pap. Mich. Ac. Sci. Arts Lett. 5, 1925, 119.
Simple: head 2-5 cm. X 1-1·5 mm., cylindric, obtuse, white or pale yellowish,
soft, fleshy, subfragile: stem 2-2·5 cm. X 0·5-1 mm., concolorous, slender, tough,
puberulous.
On the soil among moss in conifer forest, subgregarious: U.S.A. (Oregon, Mt.
Hood).
Spores 5-6·5 X 2·5-3µ,, white or tinged ochraceous, smooth, narrowly elliptic
subovoid, guttate.
Basidia 45-50 X 4-5µ,: sterigmata 4.
Cystidia 50-75 x8-12µ,, lanceolate to subcylindric below with slender stalk, sub
acute to subcapitate above, hyaline, thin walled, abundant.
Sclerotium 4-6 mm., depressed globose, fuscous.
The large fruit-bodies, large basidia, srri.aJI guttate spores, and the cystidia render
this a very unusual Typhula, if, indeed, it is such. It may be the most primitive
668 TYPHULA
member of the genus or it may be an independent sclerotioid derivative of some other
Clavarioid stock.
T. elegans (B. et C.) comb. nov.
Basinym: Pistillaria elegans B. et C., Grev. 2, 1873, 18.
1 mm. high: head subglobose, c. 600 X500 µ, orange (? dried): stem c. 600-700 x
135 µ, short, cylindric, glabrous.
On dead, slender twigs of Viburnum opulus: U.S.A. (S. Car.: No. 2780).
Spores 10·5-15 X4·7-5·3µ, ellipsoid pip-shaped (Donk).
Basidia 25-35 x6-9µ (Donk).
Hyphae 3-7·5 µ wide (Donk).
Sclerotium compressed, dark brown, c. 0·7 X0·4 mm. (Donk).
Donk has given me his ms. notes on the co-type in the Paris herbarium. The species
comes very near T. viburni but has even smaller· fruit-bodies, though with larger
spores. Compare also T. sphaeroidea with wider spores. Possibly they are all varieties
of one species, closely related with T. quisquiliaris.
T. erythropus Fr.
Syst. Myc. 1, 1821, 495.
Synonyms: Clavaria erythropus Pers., Myc. Eur. 1, 1822, 19 I.
Phacorrhiza erythropus (Fr.) Grev., Scot. Crypt. FI. 1, 1823, t. 43.
Cnazonaria erythropus (Fr.) Donk, Rev. Nieder!. Homobas. Aphyll. 2, 1933, 97.
Text-Fig. 289.
5-25 mm. high, slender, singly or rarely 2-3 per sclerotium: head 1-6 X 0·3-0·5 mm.,
cylindric or clavate obtuse, often slightly curved, becoming wholly fertile, white: stem
1-20 Xo· 1-0·3 mm., filiform, puberulous, pinkish red then reddish black and horny.
On various dead leaves, sticks, stems, Pteris petioles: Europe (common: autumnal).
Spores 5-9 X2·5-3·5µ (-9 X4µ Bourd. et Galz.), (7-11 X3-4µ, Donk), white,
smooth, oblong, pip-shaped, aguttate.
Basidia 18-30 X5-7 µ: sterigmata 2-4, 4-6 µ long.
Subhymenium c. 30 µ thick, the hyphae 2-6 µ wide, short-celled, closely interwoven.
Caulocystidia -60 X4-8 µ (below), conical, apex acute or obtuse, walls thickening,
and becoming reddish brown, the tips generally colourless and thin walled.
Hyphae 2-20µ wide, the cells 100-300µ long, some septa with clamps, others with
out: with scattered crystals: rather toughly agglutinated in the stem, at the surface
narrow, firmly agglutinated and with reddish-brown walls: rather softly agglutinated
in the head.
Sclerotium o·5-2 x0·4-1 mm., flattened-ellipsoid or shortly oblong, smooth, red
brown to blackish, more or less erumpent: medulla composed of rather closely inter
woven, free hyphae 3-10µ wide with thin or thickened walls, some heavily incrusted
with granules: cortex as a single layer of cells, 7-10µ wide, with wide lumen: cuticle
1-3µ thick, red-brown: in surface view with irregularly oblong cells, 10-30 X5-12µ,
the walls thickened, reddish brown, undulate, with a few peg-like processes into the
lumina.
Tulasne says that this species fruits in autumn from petioles of Alnus glutinosa
which have been rotting on the ground for a year. It is, perhaps, the commonest
Typhula in Europe.
" T. erythropus is near T. graminum and T. sclerotioides. From both it differs in the
red-brown stem, which is the colour of the sclerotial epidermis conveyed into the
agglutinated walls of the superficial hyphae of the stem. Its sclerotium has the same
simple construction as in T. graminum, but some of the medullary hyphae are heavily
incrusted as in T. sclerotioides. But for the red-brown colour of the stem and sclero
tium it would hardly be distinguishable from T. sclerotioides. T. neglecta may be
identical.
TYPHULA 669
T. euphorbiae (Fckl.) Fr.
Hym. Eur. 1874, 684.
Basinym: Pistillaria Euphorbiae Fckl., Bot. Zeit. 1861, 100: (lapsu calami Claviceps
Euphorbiae): Exs. No. 1287.
TExT-FIG. 289. Typhula erythropus: fruit bodies, X 5: sclerotium surface, x 500: spores,
basidia, caulocystidia, and clamps, x 1,ooo.
Head 1 mm. wide, white, clavate subglobose, very small: stem 4 mm. long, distinct,
puberulous, fuscous (125-150 µ. wide, Donk).
On decorticated stems of Euphorbia: Germany.
Spores 8·5-10 X 4·5-5 µ., pip-shaped (Donk).
Basidia 28-33 X 8 µ. (Donk).
Hyphae 3·5-5 µ. (Donk).
Sclerotium lenticular, diaphanous, yellow: medulla wholly agglutinated (de Bary).
f. virescens Rab.
Fg. Eur. No. 2351.
On rotting leaves of Juglans: Germany.
TYPHULA
I have supplemented the meagre description of this little-known species from Donk's
ms. notes. It seems very near T. viburni.
T. filicina Pk.
Rept. N.Y. St. Mus. 27, 100, t. 1, f. 29-30.
White: head thickened, obovoid or ellipsoid, obtuse, erect: stem firm, short,
pubescent, slightly thickened at the base.
On dead stems of Osmunda cinnamomea: U.S.A. (Forestburgh, N.Y.).
Sclerotium chestnut brown.
Said to be near T. gyrans. Compare T. quisquiliaris.
T. graminum Karst.
Nat. Soc. Fenn. 9, 373: Finl. Nat. o Folk 37, 1882, 183.
Text-Fig. 290.
2-6 mm. high, scattered, white: head o·5-1 Xo·15-2 mm., opaque, ellipsoid to sub
cylindric, blunt or subacute: stem 1·5-5 Xo·o5-0·08 mm., capillary, puberulous and
translucent white.
On dead grass leaves: Finland (sclerotia common, fruit-bodies rare), England (on
Molinia, fruit-bodies, Sept. 1946).
Spores 9·7-10·5 X3·3-3·7 µ,, white, smooth, ellipsoid-elongate, subacute, cloudy
vacuolate or, on discharge, 1-2 guttulate,. apiculus very small.
Basidia 25-33 X7-8·3 µ,, subclavate: sterigmata 4, 4-5 µ, long.
Caulocystidia as sterile basidia, often with slightly thickened wall, mostly prolonged
into a short or long, filiform aseptate hair -100 X1-2 µ,, sometimes bifid with some
what recurved arms.
Hyphae 3-8 µ, wide, thin-walled, without clamps, longitudinal, mostly 2-5 µ, in the
stem, the outermost agglutinated into a thin firm sheath 1-2 hyphae thick, not
agglutinated in the head: crystals absent, or a few in the head.
Subhymenium 10-15 µ, thick, 2-3 cells deep, slight.
Sclerotium 0·5-1 X0·2-0·4 mm., very small, persistently subepidermal, tawny, then
blackish, smooth, ellipsoid: medulla of free, but compact, interwoven hyphae 3-6 µ,
wide, thin walled, smooth (not incrusted) with dense, cloudy contents: cortex as a
single layer of agglutinated hyphae with wide lumen, the cells in surface view with
irregular interlocking processes, c. 10-30 X 5-10µ, (overall); cuticle red-brown, 5-6 µ,
thick, smooth.
var. filicina Karst.
Finl. Nat. o Folk 37, 1882, 183.
Fruit-bodies larger: on Polystichum spinulosum, Finland.
This species was thought to be the cause of Typhula blight of cereals until Imai
proved that it was a different species, I. Itoana
(see T. incarnata). Remsberg agrees with Imai
and has shown that the sclerotium of T.
graminum has a different structure (on the
evidence of a specimen from Karsten's type
collection).
In September 1946 I found a number of
fruit-bodies on dead Molinia leaves at
Chippenham Fen (Cambridgeshire). The
very small fruit-bodies agreed with Karsten's
meagre, if sufficient, description (no spores
TExT-Frc. 290. Typhula graminum: given), and with Remsberg's description of
fruit-bodies, x5: spores, xr,ooo: the sclerotial structure, which agreement
caulocystidia, x 500. clinches the identification. The description
TYPHULA
which I have given is similar to that of T. sclerotioides, but there is no 'cortex' between
the longitudinal medullary hyphae of the head and the subhymenium. The absence
of clamps in both species is noteworthy.
The fruit-bodies greatly resemble slender specimens of Pistillaria setipes, with
which I compared them at the time of finding. P. setipes has a stouter stem and head,
wider hyphae with clamps, and ha� no sclerotium. T. graminum is probably not
uncommon,but very small and inconspicuous.
Compare T. trifolii and T. caricina. Var. filicina may not be conspecific.
T. gyrans Fr.
Syst. Myc. 1,1821,495: de Bary,Pilze 1866,44: Quel.,FI. Myc. 1888,454: Rems
berg, Mycol. 32, 1940, 74, f. 12, 36, 45, 57: (non Pat., Wehm., Macdonald = T.
sclerotioides).
Synonyms: Clavaria gyrans Pers.,Mvc. Eur. 1,1822,191.
Clavaria juncea Lev., Ann. Sci. Nat. Ser. 2, 20, 1843, 230, t. 7, f. 1: (non Fr.
= Clavariadelphusjunceus).
Clavaria complanata de Bary,Pilze 1866, 44: Comp. Morph. Fungi 1887, 42.
Typhula complanata (de Bary), Schroet. Pilz. Schles. 1,1889,438.
(Sclerotiumjunceum Tode,S. complanatum Tode).
Text-Figs. 47, 285a,286h, 287b,291,292.
7-25 mm. high,simple, one or more from a sclerotium,rather stout, white or pale
yellowish white, becoming brownish at the base of the stem: head 2-5 xo·5-1·5 mm.,
clavate to subcylindric, obtuse, sometimes becoming hollow: stem 5-20 X o·1-1·25 mm.,
distinct,finely puberulous,twisting spirally on drying (causing the head to gyrate).
On dead leaves,petioles,herbaceous stems: Europe,N. Amer.
Spores 8·5-11·5 X 4-6µ, white, smooth, ellipsoid or obovoid, flattened adaxially,
apiculus slight.
Basidia 20-28 X 6-8 µ: sterigmata 4: cystidia none.
Subhymenium rather broad, well developed, the cells 8-16 x 4-6µ,radiating and
interwoven,the innermost cells joining with the longitudinal hyphae, short and almost
forming a 'cortex'.
Caulocystidia -60 X 4-6µ,slightly conical,attenuate to the blunt or subacute apex,
slightly thick walled in the proximal part,scattered,more crowded towards the base
of the stem.
Hyphae 3-8 µ wide,the cells 100-400 µ long in the stem,shorter in the head,with
softly mucilaginous,agglutinated walls: sparingly incrusted with crystals,more densely
in the subhymenium and stem,especially towards the base: clamps absent from the
head and basidia,occasional on some stem-hyphae.
Hyphae agglutinated firmly at the surface of the stem, the inner hyphae often
cramped and spirally contorted.
Sclerotium 0·5-3 mm. wide, convex-plane, smooth, drying rough, shrinking with
age or on drying,light brown becoming blackish when old, internally white,subepidermal,
single or (in culture) coalesced in irregular masses: medulla wholly agglutinated,
without cortex or cuticle,occasionally filamentous in the centre: epidermis 7· 5µ thick,
composed of several layers of hyphae 'winding round the medulla', with thick, golden
to dark-brown agglutinated walls: in surface view with irregularly oblong cells, the
walls thickened and slightly wavy, and with a few loose hyphae outside them: the
outer layers of the epidermis weathering away.
Temperature-range 0-25° C., opt. 12-18 ° C.: sclerotia after 7-14 days in culture:
sterile sporophores not seen in culture: sclerotia sometimes producing stolons from
which fruit-bodies arise.
There has been confusion over this species. Patouillard and Wehmeyer assigned to
it a species with small spores,5-6 X 2µ. Remsberg has assigned to it a species with large
spores. I have been fortunate to find both species. The large-spored one has the
TYPHULA
agglutinated sclerotia described for T. gyrans by de Bary and Remsberg: the small
spored one has free medullary hyphae. Therefore I conclude that the small-spored
one is not Clavaria cornplanata ( = T. gyrans) as understood by de Bary, and that
Remsberg has correctly followed de Bary's interpretation. The small-spored species
I have identified with T. sclerotioides. Fries and other early mycologists probably
confused both species, for macroscopically they are very similar, and I consider that
de Bary first defined T. gyrans microscopically in describing its sclerotial structure.
There is probably no authentic type-specimen, unless as a 'Friesian specimen', and
therefore it is better to accept de Bary's identification.
(l: J''\
�
FIG. 291. FIG. 292.
TEXT-FIG. 291. Typhula gyrans, x 5.

TEXT-FIG. 292. Typhula gyrans: spores, X 1,000: caulocystidia, X 500.
Remsberg gives T. stolonifera as a synonym, but Quelet described it with small

spores, so it must belong to T. sclerotioides. She also gives Pistillaria setipes ( = Clavaria
setipes = Cnazonaria setipes) as a synonym, but there is no evidence that it grows
from a sclerotium, and I have retained it under Pistillaria.
Young fruit-bodies show clearly Oltmann's 'spring-brunnen' type of multifila
mentous soma.
The specimens which I have studied were collected at 1,900 m. alt. near Gletzsh in
Switzerland, 27.7.25, and they must represent a spring fruiting. Remsberg records
the sclerotia for autumn, but does not mention the season of fruiting.
The following collections, named T. brassicae Donk ms., would appear to belong
to T. gyrans, perhaps as a 2-spored variety (cf. T. quisquiliaris, 2- or 4-spored), but
compare also T. umbrina. The description is taken from Donk's notes:
4-10 mm. high: head shortly cylindric, blunt, gradually tapered into the pubescent
or glabrous stalk: sclerotium 1-1·5 mm. wide, round, smooth: spores 9-14X4-5·5µ,,
ellipsoid pip-shaped: basidia 25-32X 5·5-7·5 µ,, with 2 widely divergent sterigmata 7 µ,
long: hyphae 4-6 µ, wide, thin walled: with abundant oxalate crystals on the stem: on
rotten stems of Brassica: Holland. (Valkenberg, leg. J. Rick: Naaldwijk. leg. v. d.
Tappen).
TYPHULA
T. hederaecola (Ces.) comb. nov.
Basinym: Pistillaria hederaecola Ces.,Bot. Zeit. 1855,77: Sacc.,Syll. 6,757.
3-4 mm. high,pale white: head oblong,blunt,not attenuate at the base: stem very
short,filiform: sclerotium lenticular, small,pale yellow to chestnut-brown.
On the underside of dead leaves of Hedera: Italy,France.
Little known.
T. idahoensis Remsberg
Mycol. 32,1940,89,f. 1,18,20,29.
Synonym: ? T. borealis Ekstrand,Vaxtskyddnotiser Vaxskyddsanst,Stockh. I,I937,
3,f. 1-3 (R.A.M. 16,1937,802,abstr.): Nilsson, Sverig. Utsad. Tidskr. 1,1940, 4
(R.A.M. 20,1941,251).
Text-Figs. 49,285f, 286a.
5-10 mm. high, bright warm brown to wood-brown, the head paler than the stalk,
clavate, erect, straight or slightly curved, simple, rarely branched, I-several per
sclerotium,or arising directly from the mycelium or from stolons from the mycelium
and growing on the ground: head 4-7Xo·5-1·5 mm.,vinaceous-brown,hinnuleous,
or wood-brown, apex darker, straight, elongate-fusiform, often thickened at the apex,
or cylindric, the apex sterile or becoming fertile: stem 2-5Xo·1-0·5 mm., bistre,
umber,or dark brown,glabrous or puberulous at the base.
On leaves and stems of Triticum vulgare and Agropyrum cristatum, parasitic: sclerotia
with the melting of the snow,fruit-bodies in autumn: U.S.A. (Idaho,Montana).
Spores 8-13·5 X4-8µ,av. 10·5-4·5µ,ovate-ellipsoid,apiculate.
Basidia 27-32 X6-8µ,clamped at the base: sterigmata 4-6-8.
Hyphae incrusted,clamped.
Sclerotium 0·5-2X0·5-0·9 mm.,globose or subglobose,flat below,convex above,
glabrous, shining then dull, erumpent or superficial, chestnut-brown to blackish,
never coalescent in culture: cortex composed of a single layer of thick-walled,
agglutinated hyphae: cuticle 5-20 µ thick,deep reddish brown: medulla of free inter
woven hyphae,not agglutinated,loose in the centre.
Temperature-range 0-18 ° C.,opt. 9-12 ° C.: sclerotia after 5-10 days in culture:
sterile brown sporophores abundant in culture.
This species causes the same kind of disease as T. incarnata, but the sclerotia are
easily distinguished by their colour. T. borealis is very similar, if not identical;
I have not seen its description: it is said to grow parasitically on grasses,rye,wheat,
and cruciferous weeds in Sweden.
T. idahoensis comes near T. umbrina and T. latissima.
T. incarnata Lasch ex Fr.

Epicr. 1838, 585: Donk, Rev. Nieder!. Homobas. Aphyll. 2, 1933, 95: Rabenh.
Fungi Eur.,No. 1313.
Synonyms: ? Clavaria filata Pers.,Myc. Eur. 1,1822,87 (fide Fr.).
Typhula Itoana Imai,Trans. Sapporo Nat. Hist. Soc. II,1929,39-44: Jap. Journ.
Bot. 8, 1936, 5: Remsberg, Mycol. 32, 1940, 70, f. 19, 30, 42, 52, 56: Wexham,
Phytopath.,31,1931,940: Vang,I.e. (see p. 655).
T. graminum auctt. non Karst.: Volk, Zeitschr. f. Pfl. Krankh. 47, 1937, 339:
Tasugi,Journ. Agr. Exp. Sta. Tokyo,1,1929,41.
Text-Figs. 284b,286b, 287c.
3·4-30 mm. high, solitary or in small groups from the sclerotium, simple,rarely
with a branch: head 1-20X0·4-2 mm.,filiform, cylindric,ellipsoid-oblong when short,
becoming hollow,acute then blunt,often subarcuate,whitish then flesh colour or rose
pink: stem 5-20Xo·5-1 mm.,translucent white or whitish cream, then greyish or dingy
brownish,puberulous.
5119 XX
TYPHULA
On dead culms and leaves of cereals and grasses, and on herbaceous stems and
leaves, often parasitic on cereals: sclerotia in spring, fruit-bodies in autumn: N. temp.
(Europe, Asia, N. Amer.).
Spores 7·5-10·5X4·5-5·5µ (Donk), 4-14x2-6µ (Tasugi; mostly 7-10X3-5µ),
6-13 X 3-6µ(Imai), 11-15 X4 8 -µ(Remsberg: av. 11 ·7 X6µ), white, smooth, ellipsoid
or ovoid, flattened adaxially: 12 x 6 µ (Vang).
Basidia 20-28X5-6µ (Donk), 15-40X3-7µ (Tasugi; mostly 20-30X4-6µ),
27-34X5-8 µ (Imai, Remsberg): sterigmata 4, rarely: 27-35 X5-8 µ (Vang).
Hyphae 3 ·5-6µ wide, thin walled, clamped, with a few gloeocystidium-like hyphae,
more or less agglutinated throughout the hymenium; incrusted with crystals in the
subhymenium and sparsely in the stem (Remsberg).
Sclerotia 0·5-4·5 X0·5-2 mm., subglobose, more or less flattened, smooth, drying
or ageing rough, sometimes irregular, pinkish orange when immature, then tawny to
reddish brown or dark brown, erumpent, often falling off, singly or(in culture) coalescent:
medulla wholly agglutinated, often filamentous in the centre, hollow in large sclerotia:
cuticle 8µ thick, golden to reddish brown.
Temperature-range 0-25 ° C. (Tasugi, Imai, Volk), 0-18 ° C.(Remsberg), opt. 8 ° C.
(Volk), 9-12 ° C. (Remsberg): sclerotia formed after 5-10 days in culture.
Donk agrees with me that the Typhula blight of cereals known as T. ltoana is
identical with T. incarnata. There is no difference between Donk's revised descrip
tion of T. incarnata, based on authentic European material, and lmai's description of
T. ltoana in Japan. Remsberg gives slightly longer spores for T. ltoana in the United
States, but describes the sclerotium with a structure very similar to that of T. phacor
rhiza, to which T. incarnata is very closely allied. I think that such variation in spore
size, particularly length, will be found to be common in Typhula species. If C. filata
is truly identical it will supply the earliest specific epithet, but it was said to have no
sclerotium. From T. phacorrhiza the species differs in having a distinct stem(? better
developed subhymenium to mark off the wider head) and shorter, pink fruit-body.
Greville's figure of Phacorrhiza filiformis (as a synonym of T. phacorrhiza) is T.
incarnata according to Fries, but Donk considers it to be T. phacorrhiza.
Imai first distinguished T. ltoana from T. graminum as the cause of Typhula blight
of cereals, and Remsberg confirmed this conclusion by examining the sclerotial struc
ture of Karsten's type-collection of T. graminum: its sclerotium has a filamentous, not
agglutinated, medulla though, macroscopically, both sclerotia are very similar. Rems
berg also found that Sclerotiumfulvum Fr. (in Roumeg. Fungi. Gallici 1400) was the
sclerotium of T. ltoana, but in the absence of a type-collection one may assume that
the sclerotia of both T. graminum and T. incarnata passed, for Fries, as S. fulvum.
T. incarnata closely resembles Pistillaria elegantula, which is said to have a short,
subclavate head and to lack a sclerotium, but its hyphal construction is not known.
Descriptions of the Typhula blight 1 of cereals caused by T. incarnata have been
given by Tasugi, Imai, and Volk (under the names of T. ltoana and T. graminum). It
attacks low-temperature cereals and wild grasses of all kinds, without host-specializa
tion, and it can grow saprophytically on their remains as well as on other herbaceous
plants. The disease appears to be the most virulent of any caused by Typhula. Because
of its low optimum temperature, the fungus thrives when the host-plants are becoming
numbed. Thus, in Japan, it attacks young cereals in late autumn and continues to
invade them during the winter under the snow, on the melting of which the diseased
culms and leaves of the host dry up and die, giving the plants a scorched appearance,
and sclerotia form on them: fruit-bodies are produced in autumn. In the summer
the growth of the fungus is retarded, and the host is not killed. "In Germany, it is wet
conditions and waterlogged soiLwhich favour the disease, and doubtless have the same
1
This suitable name for Typhula-diseases was proposed by Imai. That caused by T.
incarnata is also known as snow-rot, snow-scald, winter-rot, frost-injury, frost-rot, sclerotium
blight, and seedling disease. Thus vernacular names .
•
TYPHULA
effect in lowering the vitality of the host as that of the parasite increases. The
sclerotia are the main source of infection, for they may produce hyphae or fine white
rhizomorphs on germination which can infect the host, at least in its seedling state, by
passing straight through its cell-walls (Imai): similarly with the germ-tubes of the
hasidiospores (Tasugi). The fungus destroys the roots and outer leaves, and, if the
entire young plant is not killed, the mycelium may kill the secondary haulms or impede
their emergence by binding together the leaf primordia (Volk). Wheat is the most
resistant cereal (Imai). Slight infection often leads the way for Ophiobolus graminis
(Volk). Sclerotia remain viable for 25 months, according to Volk, or 16 months for
sclerotia from artificial culture.
Special conditions are needed for the production of the fruit-bodies. They are low
temperature, 4-10 ° C. (Imai), and full sunlight acting on the sclerotia (Imai, Volk), or
ultra-violet light of wave-length 2,700-3,250 A (Remsberg). But it is difficult to see
why the fruit-bodies are not then formed in late spring, immediately after the sclerotia:
it seems that a period of dormancy must intervene. In cold, dark places, filiform
dendroid, sterile fruit-bodies are produced (Tasugi).
T. intermedia Appel. et Laubert.
Arb. k. Biol. Anst. f. Land. u. Forst-Wirtsch. 5, 1905, 153, t. 3: Remsberg, Mycol.
32, 1940, 85, f. 16, 37, 41.
Text-Figs. 284c, 2861, 287h.
18-40 mm. high, white, clavate, erect, flexuous or even prostrate, simple (or sterile,
filiform, and branched), one or more per sclerotium: head 5-20 X 0·5-2 mm., cylindric
tapering, straight or slightly curved, becoming hollow, eventually fertile over the apex:
stem 5-15 X 0·3-1 mm., weak, rarely erect, usually flexuous or prostrate from the
weight of the head, puberulous.
On fallen overwintered leaves, petioles, and herbaceous stems, and on stems of
Vitis vinifera: sclerotia in spring, fruit-bodies in autumn (Remsberg, U.S.A.):
sclerotia overwintering, fruit-bodies in spring (Appel and Laubert, Germany):
Europe, U.S.A.
Spores 11·5-16 X 4-8µ,, av. 14 X 6µ,, ellipsoid, flattened adaxially, apiculate: with
1 large gutta and several small ones (Appel and Laubert).
Basidia 25-30 X 9-10µ, (Remsberg): sterigmata 4, 7·5-10 X 3µ, (Appel and Laubert).
Hyphae incrusted with crystals.
Sclerotium 1-4 X 0·5-1 mm., subglobose (? flattened), reddish brown or chestnut to
nearly black, superficial or erumpent, smooth and g-listening when fresh, drying rough
and dull, falling off from the substratum at maturity: cortex composed of a single
layer of agglutinated hyphae: cuticle II-15µ, thick, very dark reddish brown, golden
towards the medulla, rough and with small tubercles set in rosettes: medulla of free inter
woven, not agglutinated, hyphae, loose in the centre.
Temperature-range 0-21 ° C., opt. 12-15 ° C.: sclerotia after 7-14 days in culture:
sterile, branching sporophores in culture at 6-15 ° C. frequent, especially with ultra
violet radiation.
This is very near T. variabilis, from which it differs in the white fruit-bodies and
narrow lumina of the cortical hyphae of the sclerotium. T. lactea, with milk-white
fruit-bodies, is generally referred to T. variabilis, but there is nothing to distinguish it
from T. intermedia. Compare T. sclerotiicola.
T. Laschii Rabenh.
Bot. Zeit. 1849.
Filiform: head -23 mm. long, cylindric, white, then fuscescent: stem -34 mm. 1ong,
pruinose then glabrous, white.
On dead stems of Rubus caesius: Germany.
(Spores 11 X 5µ,, ellipsoid: Donk.)
TYPHULA
Sclerotia subcompressed, rugulose, blackish.
This is said to resemble T. phacorrhiza but to differ in form (rather as a very
elongate T. erythropus). The spore data are from a single measurement in Donk's ms.
notes on Rabenhorst's collection.
T. latissima Remsberg
Mycol. 32, 1940, 93, f. 13.
Text-Fig. 285e.
6-r8 mm. high, white or pale olive-brown, yellowish with age or on drying, erect,
straight or curved, simple r-several per sclerotium: head 4-8 X0·8-2 mm., cylindric
clavate, subglobose, or flattened, often irregular, blunt or rounded, apex fertile, solid,
blackening with age: stem 3-ro Xo·r---0·5 mm., very slender, filiform, straight or
flexuous, puberulous, thickened at the base, solid.
On stems of Typha latifolia; fruit-bodies in autumn: U.S.A.
Spores ro-15 X4-8µ, av. ro·5 X4µ, ovate, apiculate.
Basidia 30 X8 µ: sterigmata 4.
Hyphae incrusted in stem and head.
Sclerotium 2·5-3 X1·5-2·5 mm., subglobose or flattened and unequal-sided,
erumpent, bone-brown, simple or (in culture) coalescent: cortex composed of a single
layer of agglutinated hyphae with wide lumen: cuticle ro-15µ thick, deep reddish
brown, rough; medulla composed of free interwoven hyphae, loose in the centre.
no sterile sporophores in culture.
This seems to me almost the same as T. umbrina. The photograph shows what
appear to be malformed heads scarcely answering the description.
T. lutescens Boud.
Bull. Soc. Myc. Fr. 16, 1900, 197, t. 9, f. 2.
Head 5-8 X1·5-2 mm., yellow: stem whitish, sparsely pilose, two to three times as
long as head: sclerotium tawny, oblong, 3 X1·5-3 mm.
Spores 9-12 X3-4µ, white, smooth, oblong-cylindric.
On petioles of Fraxinus: France.
Little known. Possibly a form of T. gyrans.
T. Muelleri (Saut.) comb. nov.

Basinym: Pistillaria Muelleri Saut., Flora 1852, 579.
Synonyms: Pistillaria sclerotioides Fr., Syst. Myc. 1, 1821, 497.
Typhula sclerotioides Karst., Finl. Nat. o Folk, 37, 1882, 181 (non T. sclerotioides Fr.
1838).
Head 2-3 mm. high, blunt, clavate, reddish or purplish, then bay-brown: stem 2 mm.
long (? concolorous), set with radiating fibrils at the base: sclerotium bistre, tuber
culoid.
On dried stems of Gentiana: France, Norway.
The colour is so striking that the species should be easily recognized though it is
clearly rare. T. tochinaiana seems identical. In making the new combination T.
Muelleri, I have accepted P. Muelleri as a synonym as given by Fries (H ym. Eur. 686)
and Saccardo (Syll. 6, 752): thus the new name is, as Lloyd would have said, a juggle,
though necessary.
T. neglecta Pat.
Rev. Myc. 1885, 152: Tab. An. n. 471.
8-15 mm. high: head 2-<J X0·2-0·4 mm., linear, cylindric, straight or a little curved,
white: stem 15-20 X0·09-0·2 mm., capillary, rigid, violaceous black, brown puberulous.
TYPHULA
On dead petioles of Populus, Juglans, &c., on heaped stems of Helianthu;, tuberosus,
and on the ground: France.
Spores 5-7 X2-3 µ,, 9-10 X2 µ, (Tab. An.), white, smooth, cylindric, or slightly
curved.
Basidia 16-21 X3-4µ,: sterigmata 2-3-4.
Caulocystidia 15-22 X4µ,, cystidium-like.
Hyphae 6-19µ, wide.
Sclerotium I mm. wide, globose, blackish brown, erumpent or in the ground.
This seems to be the same as T. erythropus.
T. ovata Karst.
Finl. Nat. o Folk 37, 1882, 183: (non Schroet. 1888 = Pistillaria pusilla).
Synonyms: ? Phacorrhiza Amansii Brond., Ree. Pl. Crypt. 1, 1828, 10, t. r.
? Pistillaria Amansii (Brond.) Sacc., Syll. 11, 1895, 141.
1 ·5-5 mm. high, white: head enlarged, ovoid: stem short, glabrous: sclerotium
2-5 mm. wide, lenticular, pale brown then darker brown to blackish.
Compare Pistillaria pusilla, which lacks the sclerotium.
The description of P. Amansii is:
Gregarious, cylindric, white, fasciculate, arising from an ochraceous, sinuate-rugose
tubercle: on fallen le.aves of Populus nigra, France (S. Amans). If identical with
T. ovata, it is the earlier name.
T. pallens Maire
Bull. Soc. Myc. Fr. 44, 1928, 53.
5-<J mm. long, solitary or gregarious, fleshy-fibrous, taste mild, smell faint: head
2-4 X 0·5 mm., clavate or cylindric-clavate, rounded or obtuse, ochre-white then
ochraceous-rufescent when old: stem �-5 X0·25 mm., filiform, with short spreading
hairs, white, then rufescent at the base and rufescent-brown upwards.
On dead leaves of Olea europea: N. Africa (Mauretania).
Spores 6-7 X3 µ,, white, smooth, oblong, blunt, guttulate.
Basidia 22-25 X5µ,, cylindric clavate, sterigmata 3, 3-5 µ, long.
Caulocystidia filiform, slender, hyaline, aseptate.
Hyphae 4-6 fL wide, much incrusted with crystals on the surface of
the stem.
Sclerotium smooth, round, more or less compressed, glabrous,
rather fuscous, internally white, immersed.
This is said to be near T. erythropus and T. Laschii. Compare
also T. Buxi.
T. Patouillardii (Quel.) comb. nov.

Basinym: Pistillaria Patouillardii Quel., Ass. Fr. 1883, 507:
Patouillard, Tab. An. 1883, n. 48.
Text-Fig. 293.
2-3 mm. high (Pat.): head filiform, white, acute, pruinose: stem
2-3 mm. high (Quel.), 0·5 mm. (Pat.), verv short (Bourd. et Galz.),
distinct, fawn-brown or tawny, with crystab among the hyphae.
On dead stems of Carduus (Quel.), on dead leaves (Pat., Bourd.):
France.
Spores? 8 µ, long, ovoid-oblong (Quel.): 4-5 X 2µ,, obovoid, (Pat., TEXT-FIG. 293.
T. Patouillardi£:
Bourd.).
fruit-bodies with
Basidia 2-spored (Quel.), 4-spored (Pat.). and without sclc
Sclerotium small, brown, round or compressed, rough, bearing rotia: (from Pat.
1-3 fruit-bodies, or the fruit-bodies without sclerotia. Tab. An. n. .;8).
Possibly there are two fungi confused under this name by Quelet
TYPHULA
and Patouillard. As understood by Bourdot and Galzin, who make no mention of
a sclerotium, Pistillaria Patouillardii differs from P. albobrunnea (p. 476) only in the
filiform head, which may be an expression of age. Quelet's description suggests a
Typhula related to T. erythropus and T. o'vata.
Pistillaria Patouillardii sensu Killerm. may be Pistillaria epiphylla (p. 480).
[The name is not to be confused with Pistillina Patouillardii = Pistillaria capitata
(Pat.) Sacc.]
T. pertenuis Remsberg
Mycol. 32, 1940, 83, f. 22.
Text-Fig. 285c.
6-30 mm. high, white, then avellaneous when old or on rlrying, erect or occasionally
prostrate, simple, one per sclerotium, or arising from the mycelium (or, in culture,
from stromatic crusts): head 8-15X0·5-3 mm., elongate-clavate, obtuse, solid, the
apex becoming fertile (or ? persistently sterile in some cases): stem 4-10X0·5-1 mm.
puberulous.
On soil, fruiting in autumn: U.S.A.
Spores 8-12·5x4-6 µ., av. 9X5 µ., ovoid, apiculate.
Basidia 25X8 µ.: sterigmata 4.
Hyphae more or less incrusted.
Sclerotia 2-3 mm. wide, subglobose, smooth, then rough, bistre to blackish, single
or (in culture) coalescent: cortex composed of a single layer of agglutinated hyphae
with wide lumen: cuticle 7-10 µ. thick, golden-brown, rough: medulla composed of
free interwoven hyphae, loose in the centre, with wide lumen next the cortex.
Temperature-range 0-21 ° C., opt. 15-18 ° C.: sclerotia formed after 5-7 days in
culture: sterile sporophores not seen in culture.
Compare T. virgata.
T. phacorrhiza Fr.
Syst. Myc. 1, 1821, 192.
Synonyms: Clavaria phacorrhiza Pers., Myc. Eur. 1, 1822, 192.
? C. minor Lev., Ann. Sci. Nat. 1843, t. 7, f. 2.
C. scutellata de Bary, Pilze 1866, 44: Comp. Morph. Fungi 1887, 42.
Phacorrhiza filiformis Grev., Scot. Crypt. Fl. 2, 1824, t. 93: ( = Typhula incarnata
teste Fries, Hym. Eur.: non T. filiformis Fr. = Clavariadelphus junceus).
Typhula subphacorrhiza Britz., Hymen, Siidbay. 8, 1891, 15, f. 77-8.
? T. fiavescens Saut., Flora 1, 1847, 317. ..
Text-Figs. 48, 286g, 287a.
2·5-10(-14) cm. high, filiform, white, then cream, brownish, yellow-ochraceous or pale
ochraceous-brown, singly or, occasionally, 2-3 per sclerotium: head 10-20X0·5-1 mm.
(? longer), filiform-cylindric, with acute sterile tip, scarcely distinguishable from the stem:
stem 0·3-0·8 mm. wide, very variable in height, flexuous, filiform, sparsely puberulous,
at least near the base: texture rather horny cartilaginous.
On dead leaves, petioles, herbaceous stems and grasses: sclerotia in spring, fruit
bodies in autumn: N. temp. (Europe, Asia, N. Amer.).
Spores (9-)II-15(-20)X4-8 µ., very variable, white, smooth, ellipsoid or sub
cylindric, blunt, flattened adaxially, thin-walled, aguttate.
Basidia 20-33X9-12 µ., clavate, without clamps: sterigmata 2-4,. 6-8 µ. long.
Subhymenium practically none, the basidia arising directly from the longitudinal
hyphae: cystidia none.
Caulocystidia -80X4-6 µ., slightly tapered or subcylindric, slightly thick-walled at
the base, colourless, smooth, aseptate, reduced in the upper part of the stem to short
subclavate processes like sterile basidia.
Hyphae 3-16 µ. wide, with long or short cells, slightly constricted at the septa,
TYPHULA
without clamps except on some of the narrow superficial hyphae of the stem, with
agglutinated walls, firmly mucilaginous in the stem, rather softly in the head: with
abundant coarse, dark-brown crystals among the superficial hyphae of the stem and
head (? always): the superficial hyphae of the stem slightly thick walled and toughly
agglutinated.
Sclerotium r-6(-8) X 1 ·5-3 ·5 mm., flattened, especially on drying, sometimes lobed
and irregular, pallid then tawny, russet or orange-brown to dark brown, superficial,
adherent by a minute dark-brown or reddish-brown stalk, smooth then rugose on drying,
very cartilaginous, singly or (in culture) coalescent: medulla wholly and toughly
agglutinated, sometimes filamentous in the centre or hollow in large sclerotia: cuticle
c. -ro µ, thick, yellow to reddish brown: without epidermis (Sclerotium scutellatum).
Temperature-range 0-21 ° C., opt. 12-15 ° C.: sclerotia after 7-12 days in culture.
This species has been made the lecto-type of the genus by Donk and Remsberg.
It has the largest fruit-bodies, with unlimited apical growth, the least-developed sub
hymenium, the most agglutinated tissue (at least, in the sclerotium), and the most
variable spores. It greatly resembles Clavariadelphusjunceus in its fruit-bodies, spores,
basidia, and occasional gloeocystidia-like hyphae, but C. junceus never has a sclerotium
and its hyphae are agglutinated only on the surface of the stem. The colour of the
fruit-body in T. phacorrhiza seems to be caused mainly by the brown crystals among
the superficial hyphae. For details of spore-variation, see Corner (1947).
The description of Clavaria minor is:
'gregaria, pumila, simplex, capillaris, levis, apice acuminata, tuberculoso subglobose
flavo innata: ad terram, pr. Paris, Galliae' (Sacc.).
The description of T. fiavescens is:
Up to 25 mm. high, yellowish, head cylindric: stem slender, glabrous; sclerotium
globose: on the ground among moss, Germany.
T. quisquiliaris (Fr.) comb. nov.
Basinym: Pistillaria quisquiliaris Fr., Syst. Myc. 1, 1821, 497: Pat., Tab. An.,
No. 687: Donk, Rev. Nieder!. Homobas. Aphyll. z, 1933: Josserand, Bull. Soc. Myc.
Fr. 53, 1937, 225, f. 17.
Synonyms: Clavaria obtusa Pers., Myc. Eur. 1, 1822, 190 (pr. p.: alt. p. = Pistillaria
uncialis).
Clavaria quisquiliaris (Fr.) Karst., Finl. Nat. o Folk, 37, 1882, 178: var. infiata
Karst., ibid.
Pistillaria puberula B., Out!. 1860, 286.
3-9 mm. high, simple, rarely bifid, white, one per sclerotium: head 1 ·3 X
0·4-1 ·2 mm., cylindric, then ovoid, pyriform, or subglobose, entirely fertile, sometimes
flattened: stem 1-6(-10) X 0·1-0·3 mm., finely puberulous, especially near the base, or
nearly glabrous: texture soft, then rigid or rather waxy-horny, drying cartilaginous.
On dead petioles of Pteris aquilina (? other ferns): Europe (common).
Spores 11-15 X 4-6 µ,, white, smooth, subcylindric or subamygdaliform, thin-walled,
aguttate; the contents granular (Donk).
Basidia 30-40 X 8-9 µ, (Josserand), X 5 · 5-6·5 µ, (Donk), X 6-7 µ, (Lundell); with
4 sterigmata (Josserand), 2 sterigmata 5-6µ long (Donk, Lundell).
Subhymenium well developed as in T. gyrans: cystidia none.
Caulocystidia -120 X 4-8 µ, at the base, 1·5-3 µ, at the tip, narrowly conical tapering,
acute or obtuse, generally slightly thick walled to the tip, smooth, colourless, aseptate.
Hyphae 2-7 µ, wide in tht> stem, long celled, most septa with clamps, with softly
agglutinated walls, the narrow superficial hyphae of the stem toughly agglutinated
and slightly thick walled.
Sclerotium 1-3 X 0·3-0·4 mm., oblong (like an ant pupa), smooth, ochraceous,
immersed: structure apparently as in T. phacorrhiza.
680 TYPHULA
I have transferred this species to Typhula because it has a sclerotium and because
it seems to me to have the same agglutinated hyphal structure as T. gyrans and T.
phacorrhiza, from which it differs mainly in the very limited apical growth. The
sclerotium, which is always deeply immersed below the hard 'cortex' of the bracken
petiole, was overlooked by early mycologists: it was first found by Tulasne, and its
presence has been established by Donk and Josserand. Donk has made it the lecto-type
of Pistillaria s. str., because of its small fruit-body and waxy flesh drying horny. I do
not find that these are adequate criteria for a generic separation from T. phacorrhiza,
as mentioned on page 147, and the two extremes are bridged by T. incarnata and
T. gyrans.
-·· · ·-
TEXT-FIG. 294. Typhula quisquiliaris, X 5:

the sclerotia beneath the surface of the
Pteris petiole.
TEXT-FIG. 295. Typhula quisquiliaris: spores, caulocystidia, and clamps, X 1,000.
Pistillaria puberula is merely a form with more hairy stem (Donk, Josserand). The
hairiness is variable and the stem may even be microscopically glabrous.
Bayliss-Elliott has described the fungus in culture, but, as no sclerotia were formed,
I am doubtful if it was rightly identified with P. quisquiliaris (Proc. Birm. Nat. Hist.
Phil. Soc. 16, 1933, 98). In culture, it formed zones of fruit-bodies alternating with
zones of sterile mycelium, both fruit-bodies and mycelium discolouring yellowish or
fuscous with age. What were taken to be normal fruit-bodies in culture had short
stems 2 mm. long and oval heads 1-2 mm. Jong, but the heads were often absent, and
some fruit-bodies (on bread- and potato-agar) had stems 2-4 cm. long, often branched,
TYPHULA 681
sometimes palmately or cristately with or without small fertile heads at the top. The
spores were 8-11 X4-5 µ: basidia 4-spored. The fungus was isolated from soil
cultures.
T. ramealis Speg. et :floum.
Rev. Myc. 2, 1880, 15.
Synonym: Pistillaria ramealis Cke., Grev. 8, 1880, 81.
(Pistillaria ramealis Lib. in herb.)
Branched, white, pubescent, with subfastigiate, obtuse, white clubs: sclerotium
black, small 'Sphaeria-like'.
On bark of various trees (Syringa, Rubus): Belgium.
Spores 3-3·5 X1-2µ (Speg. et Roum.).
Little known. Perhaps the same as T. caespitosa var. coacervata.
T. sclerotiicola (Allesch.) comb. nov.
Basinym: Clavaria sclerotiicola A. Allesch. Verz., d. Sudb. Pilz., p. 65: Sacc., Syll.
II, 1895, 135.
Filiform subulate, often with a few irregular branches, acute, glabrous, white, base
not narrowed: sclerotium flat, ellipsoid or fusoid, often irregular.
On rotten stems of Lunaria rediviva and Urtica dioica: Germany (Oberammergau).
Spores 8-10 X5-6µ, aguttate, or 8-14 X2-5µ, 2-guttate, ovoid-pyriform, hyaline.
This may be Typhula intermedia.
T. sclerotioides (Pers.) Fr.

Epicr. 1838, 585: Bourd. et Galz., Hym. Fr. 1927, 695: (non Karst. 1882 = Typhula
Muelleri).
Basinym: Phacorrhiza sclerotioides Pers., Myc. Eur. 1, 1822, t. 11, f. 1-2: Bail,
Hedw. 1, 1853,_93.
Synonyms: T. Friesii Karst., Finl. Nat. o Folk 37, 1882, 182.
T. stolonifera Quel., Ass. Fr. 1883, 506, t. 6, f. 17: Pat., Tab. An., No. 264.
T. asphodeli Pat., Journ. de Bot. 1894, 20.
T. stricta Appel, Arb. k. Biol. Anst. f. Land-Forst-Wirtsch. 5, 1905, 3, 151.
T. gyrans sensu Pat., Tab. An., No. 262: Bourd. et Galz., Hym. Fr. 1928: Wehm.,
Pap. Mich. Ac. Sci. Ari:s Lett. 20, 1935, 264: Macdonald, Ann. Appl. Biol. 21, 1934,
590: (non Fr.).
T. mucor sensu Kauffm., Pap. Mich. Ac. Sci. Arts Lett. 5, 1925, 120: (non Pat., see
Pistillaria setipes).
Text-Figs. 46, 51, 296-8.
Fruit-bodies -30 mm. high, slender, one per sclerotium: head 1·5-6(-10) X
0·2-0·4 mm., linear subulate then subclavate and blunt, white then pallid yellowish,
opaque, waxy, wholly fertile: stem, 5-25 Xo· 1-0·3 mm., filiform, slightly attenuate
upward (apex 90-200µ wide), very variable in length, pellucid white, very finely or
sparsely puberulous or appearing glabrous, sometimes brownish at the base.
On dead herbaceous stems and leaves, and on fallen leaves and small twigs of trees:
fruiting in autumn: Europe, N. Amer. (Oregon, Manit., Nov. Scot.).
Spores 6-9·5 X2-3·5(-4)µ, white, smooth, cylindric, blunt, flattened adaxially or
subarcuate, aguttate.
Basidia 20-30 X5-7 µ: sterigmata 4, 4-5 µ long (2 sterigmata, Bourd. et Galz.).
Subhymenium c. 20µ thick, composed of cells 6-12 X2·5-4µ wide and connected
to the longitudinal hyphae by a 'cortex' 20-30µ wide, composed of similar but larger
cells 15-25 X6-12µ: cystidia none.
Caulocystidia -75 X4-8 µ at the base, conical, slightly thick-walled in the proximal
part, tapered to an acute thin-walled apex, very scattered, even at the base of the stem
(? very reduced or absent in some specimens).
682 TYPHULA
Hyphae 5-20µ wide, the longitudinal hyphae with long fusiform cells (-500µ long),
with or without clamps, shorter celled and more inflated at the base of the stem (cells
25-200 X 10-30µ), with slightly and rather softly mucilaginous walls, the narrow
hyphae (3-9 µ wide) at the surface of the stem firmly agglutinated and slightly thick
walled (at the apex of the stem generally thin walled and not agglutinated, and with
scattered sterile basidia): a few narrow hyphae 3-5µ wide, with resinous-oleaginous
contents.
Sclerotium 0·5-5 mm. wide, very variable in size, subglobose, becoming flattened or
concave on one side, pale ochraceous then brownish and finally
blackish, smooth, erumpent: cortex 40-60 µ thick, composed of
many ·agglutinated hyphae 3-8µ wide, not in distinct layers:
cuticle 5-8 µ thick, yellow brown: medulla white, composed of
free, loosely interwoven hyphae, mostly 5-9 µ wide, thick walled
(walls 1-2µ) and densely incrusted with small colourless crystals
1-3 Xo·5-2µ, a few hyphae 3-5µ wide and thin walled: in
surface view, with irregular oblong cells, the lumina 12-20 X
4-8µ, separated by rather wavy brown walls 2-3µ thick: fruit
bodies erumpent from the medulla below the cortex.
I first found this fungus with unattached sclerotia among dead
herbaceous remains in Wicken Fen (Cambridgeshire, England,
22.8.26). I took it for T. gyrans until I discovered that the
slender fruit-bodies, small spores, and sclerotial structure did
not agree with T. gyrans described by Remsberg. whose
interpretation, as I have explained under that spedes, I consider
to be correct. I then found that my fungus was certainly the
same· as that described by Wehmeyer and Patouillard as T.
gyrans. and by Kauffmann as T. mucor, and I thought it a new
species until I came across Bourdot and Galzin's interpretation
of T. sclerotioides Fr., which is placed among the corrections and
additions at the end of their work. Theirs is the first microscopic
interpretation of T. sclerotioides, and as it agrees in all points,
except the width of the spores and the number of sterigmata, with
the fungi found by Patouillard, Kauffmann, Wehmeyer, and
myself, it seems advisable to follow their lead. I have since
found it commonly in the fens of East Anglia (Sept. 1946).
The point which Persoon made about Phacorrhiza sclerotioides
was the endogenous origin of the fruit-body, which he imagined
developed in the sclerotium like an Amanita in its volva. My
TEXT-FIG. 296.
fungus from Wicken Fen had also endogenous fruit-bodies, Typhula sclerotioides,
though it was only the primordial shaft which formed in the X 5.
medulla, and, clearly, it is the unusually thick cortex of the
sclerotium which shows so well in this species what is probably the normal method of
development of the fruit-body in Typhula, the thin cortex of most sclerotia scarcely
revealing the slit where the fruit-body emerges. Thus I am convinced that T. sclero
tioides has at length been satisfactorily rediscovered in spite, as Bourdot and Galzin
remark, of the criticism of Fries. The species is closely allied with T. erythropus.
T. stolonifera appears to be only an unusual form with the fruit-body developed
from a stolon, as may happen in T. gyrans: but the small spores 7 µ long (5 X 2 µ in
Quelet's figure) prevent it from being T. gyrans. T. stricta has slightly wider (1 µ
wider) spores than T. sclerotioides, which scarcely seems important. T. trifolii is very
similar macroscopically, but it has much larger spores and, as its sclerotial structure
is not known, I have kept it apart. T. betae appears related, but has an elongate head.
As I understand, T. sclerotioides is a white Typhula like T. gyrans but with more
slender fruit-body, small spores, and less agglutinated hyphae. The collections which
TYPHULA 683
I refer to it vary in their spores, caulocystidia, and sclerotial size, which do not appear
to be specific. Thus:
Spores 5-5·2 X 2 µ. (Wehm.), 5-6 X 2 µ. (Pat.), 5-7 X 3 µ. (T. Asphodeli), 6-8 X 3-4·5 µ.
(Appel, Bourd. et Galz.), 7 µ. pruniform (Quel.), 7-9·5 X 2·5-3 µ. (Corner, Kauffm.),
5·5-8 X 3-3·5 µ. (Corner).
.Caulocystidia sparse (Corner), stem glabrous (Kauffm., Wehm.): stem closely pilose
(Quel., for T. stolonifera).
0000000
TEXT-FIG. 297. Typhula sclerotioides:
spores, X 1 ,ooo.
TEXT-FIG. 298. T;,plmla sclerotioides: sclerotial surface,

> 1,000.
Hyphae with clamps in some collections (Corner, 1946), without clamps in other
collections (Bourd., Corner, 1926).
Sclerotium 0·5-1 mm., pale brown then black (Wehm.): minute, dark brown
(Kauffm.): 1-1·5 X 0·5-0·8 X 0·3-0·5 mm., pale yellow-ochraceo11s to pal'.! ochraceous
brown, then fuscous (Corner): 1-4 mm. blackish, becoming cupular and dentate
(Bourd. et Galz.): -5 mm. shining black, sometimes umbilicate (Appel): globular,
black (Quel.).
Habitat on Mulgedium alpi11um and Cacalia (Bourd. et Galz.), on Solanum tuberosum
(Appel), on leaves of frondose trees (Kauffm., Quel.), on rotting leaves of Asphodelus
(Tunis, T. Asphodeli), on fallen leaves and slender twigs of Alnus and Salix (Corner).
In this place, also, I put the following manuscript species, T. polymorpha Donk
ined.: its slightly wider spores suggest as already noted, T. stricta:
-12 mm. high, white: head -2 mm. wide, cylindric-subclavate, narrowed to the
apex or blunt, gradually attenuate into the glabrous stem (t-t the total height, very
variable): sclerotium -1·5 mm. wide, round, smooth, light to dark brown, internally
whitish: spores 5-7"5 X 4-4·5 µ., ovoid pip-shaped: basidia 20-35 X 5-6 µ., with (3-)4
sterigmata 5-6 µ. long: hyphae inflated in places 20-30 µ. wide, with scattered oxalate
crystals: on dead herbaceous stems: Holland (Katwijk-Binnen, leg. S. J. Scholte,
Nov. 1926).
The species described by Macdonald (Le.) as T. gyrans also belongs here. The
spores are given as 5·6x4·4 µ. (:iv. of 50), and they suggest that T. sclerotioides may
TYPHULA
have two varieties, one with narrow spores, the other with wider spores as in the pre
ceding T. polymorpha Donk and, perhaps, T. stricta. Macdonald investigated the
species in culture and found that it was not parasitic on cabbages in Scotland, though
growing on their rotting remains. Mycelial growth occurred between 0-25 ° C., with
an optimum at 15-17 ° C.: that for the production of fruit-bodies was 17-20 ° C., and
for sclerotia 13-15 ° C. The sclerotia took 9-10 days to develop and gave out drops of
water in maturing. In abnormal cases minute sclerotia developed on the ends of
rudimentary, sterile fruit-bodies. The sclerotia germinated at any time from 8 days
to 5 months after their formation. The fruit-bodies were occasionally branched, even
with as many as seven antler-like branches. Sometimes, in place of normal fruit-bodies,
there developed in artificial culture elongate, filiform-flexuous bodies, up to 12 cm.
long, as overgrown stalks, which eventually became fertile (cf. Clavariadelphus
junceus). Four nuclei were formed in the basidium and the spores were uninucleate.
The species was heterothallic. (But see Postscriptum, p. 688.)
T. sphaeroidea Remsberg
Mycol. 32, 1940, 74, f. 23, 24, 43, 44, 49, 54.
Synonym: ? T. rubicola B. et C., Grev. 2, 1873, 18.
Text-Figs. 49, 2851, 287d.
1-3 mm. high, erect, simple, generally one per sclerotium or arising directly from
the mycelium (? in culture only): head 0·8-1 ·5 X 0·5-1·5 mm., subglobose or pyriform,
chalk-white, drying milk-white, wholly fertile: stem 1-2 X 0·3-0·5 mm., puberulous,
white, reddish brown at the thickened base.
On dead stems of Rubus, fruiting in August: U.S.A.
Spores 11·5-13 ·6 X 6-9 µ., av. 12 X 7 µ., white, smooth, ovate, truncate, apiculate.
Basidia 20-45 X 9·5-12 µ., breaking away from the subhymenial hyphae: sterigmata 4.
Hyphae of stem and subhymenium incrusted.
Sclerotium 0·5-1 mm., convexo-plane, wood-brown then almost black, somewhat
rough, immersed, solitary or (in culture) clustered: medulla wholly agglutinated, or
with the hyphae free and loosely interwoven, sometimes with thick-walled hyphae in
the centre: epidermis 15-30 µ. thick, composed of 3-4 layers of hyphae with thick,
golden or red-brown, agglutinated walls, unevenly delimited from the medulla:
cuticle none.
Temperature-range 0-25 ° C., opt. 12-18 ° C.: fruit-bodies develop abundantly
from the mycelium at 12-18 ° C. in culture: sterile abnormal sporophores at 9-12 ° C.
This species seems close to T. quisquiliaris. The description of T. rubicola is:
8 mm. high, filiform, opaque, cylindric, with clavate white apex, the base inserted
. in the matrix: on dead Rubus: U.S.A. (Pa., Michener No. 4332).
T. subulata Remsberg
Mycol. 32, 1940, 94, f. 14.
Text-Fig. 285d.
9-18 mm. high, simple, straight or curved, 1-3 per sclerotium: head 8-13 x
2-2·5 mm., subulate, attenuate to the very acute sterile tip, pallid-white: stem
3-5 xo·1-0·2 mm., short, filiform, glabrous, brownish (darker than the head).
On dead leaves and grass: fruiting in autumn: Canada (Man.).
Spores 12·5-15·5 X 7-8 µ., av. 14 X 7·5 µ., white, ellipsoid, apiculate.
Basidia 30 X 8 µ., 4-spored.
Hyphae more or less incrusted.
Sclerotium 2-2·8 mm., subglobose or elongate, convexo-plane, glabrous, drying
rough, brown, erumpent, simple or (in culture) coalescent: cortex composed of a single
layer of thick-walled agglutinated hyphae: <yticle 10-15 µ. thick, golden-brown, rough:
medulla composed of interwoven free hyphae.
TYPHULA 685
° °
Temperature-range 0-21 C., opt. 12-15 C.: sclerotia after 5-7 days in culture:
sterile brown fruit-bodies frequent in culture from sclerotia or mycelial mats.
This species is close to T. variabilis, but differs in the subulate-acute head on a short,
very slender stalk, in the paler colour of the fruit-body, and in the cortical hyphae of
the sclerotia having a narrow lumen: perhaps also in the absence of caulocystidia.
T. tochinaiana (Imai) comb. nov.

Basinym: Clavaria tochinaiana Imai, Tr. Sapp. Nat. Hist. Soc. II, 1930, 72: 12,
1931, 19.
Simple, gregarious or solitary, medium-size, reddish brown, arising from a depressed
globose or cupular, light-reddish-brown sclerotium.
Spores 12·5-15 x6-7·5µ, white.
On rotting leaves of Brassica oleracea: sclerotia in spring, fruit-bodies in autumn:
Japan.
This suggests the little-known T. Muelleri.
T. trifolii Rostr.
Ug. Ld. 35, 1890, 72: Lind, Dan. Fungi 1913, 365: Sacc., Syll. 23, 1925, 497:
Noble, Ann. Bot. New Ser. 1, 1937, 67: Tabajdy (see abstr. R.A.M. 15, 1936, 725).
Synonym: ? Pistillaria clavulata Ellis, Bull. Torr. Bot. Cl. 1877, 133.
8-17 mm. high, white or yellowish white: head shortly cylindric to rather elongate,
blunt or subacute: stem puberulous at the base.
On stems and leaves of Trifolium incarnatum, T. pratense, T. repens, Anthyllis
vulneraria, Medicago luputina, saprophytic and parasitic: Europe (continental).
Spores 8·5-12·5 X3·8-6·5µ, av. II X5p., white, smooth,
Basidia 4-spored.
Hyphae without clamps in the fruit-body, the mycelial hyphae clamped.
Sclerotia 1-2 mm. wide, subglobose, hard, rough, black or dark brown: cortex thin:
medulla of free, interwoven, thin-walled hyphae, loose in the centre, or the large
sclerotia hollow.
This species is a facultative parasite of similar behaviour to T. incarnata. Systemati
cally it seems to come between T. betae and T. sclerotioides in the size and shape of the
head, but it ha.& considerably larger spores: a comparison of the structure of the three
will be interesting and informative.
Noble has studied the species in culture. The following is a resume:
Four kinds of mycelium in culture, namely haploid submerged, haploid superficial,
diploid submerged, and diploid superficial. The submerged mycelia spread more
quickly than the superficial, the hyphae of which have shorter and more abundant
branches: many of these branches have no nucleus.
(a) Haploid Mycelium. Laxly branched at a wide angle, th<! apical cell occasionally
branching. Cells typically I-nucleate, but some with 2-3 nuclei, especially near the
apex of the hypha. Anastomoses betw}'!en branches of the same myr.elium frequent,
the nucleus of one cell passing into the other cell. Oidia 3·6 Xo·8µ, white, ellipsoid
cylindric, I-nucleate, formed on short, subcylindric projections from the cells.
· (b) Diploid Mycelium. Cells 2-nucleate, the septa with clamps, often branched from
the clamp. Some cells 3-4-nucleate, occasionally 1-nucleate, and apparently nuclear
division not always conjugate. Anastomoses between hyphae of the same mycelium
frequent, the 2 nuclei from I cell passing into the other, and the nuclei of the invaded
cell disintegrating. Sometimes a branch has only I nucleus and grows into a haploid
hypha with oidia.
In both superficial and submerged mycelia the nucleus (or paired nuclei) does not
enter a branch until it is four to six times as long as wide.
Clamps are formed mostly by the backward growth of the hook which may be met
terminally or laterally by a peg from the penultimate cell. The clamp forms about the
686 TYPHULA
middle of the terminal cell but at the base of a branch, at its junction with the parent
hypha.
Sclerotia form at 13 ° C. abundantly, but none above 23 ° C.
Fruit-bodies appear immediately after the sclerotia in culture but, if the sclerotia
are removed and planted in sand, a resting period intervenes before they develop
fruit-bodies.
The species is heterothallic, but it is not certain whether bipolar or tetrapolar. The
conjugate nuclei fuse in the basidium: the fusion nucleus undergoes two divisions and
1 daughter nucleus enters each spore which remains uninucleate.
Haploid Fruit-bodies. As the diploid but much smaller, 1·5-'7 mm. long, sometimes
arising from the mycelium instead of from the sclerotia, and such with the lower part
of the stalk dark brown and its surface composed of thick-walled, dark, agglutinated
hyphae like those of the sclerotial cortex. Sporing vigorously with haploid spores
5 X 2·5 µ (half the size of the diploid, but of the same shape). Sclerotia as the diploid,
but smaller. Nuclear fusion in the basidium exactly as in the diploid fruit-bodies.
Retaining their vigour through several generations (? indefinitely).
The disease caused by T. trifolii has been called 'Crimson Clover Rot', but it would
seem desirable merely to call it 'Typhula blight of clover'. Infection is said to occur in
autumn, when the host-plants wilt and" die back, especially in wet weather. Snow
does not hinder the disease. The conditions which favour it are early August sowing
of clover, dense autumn stands, excessive nitrogenous manure, and lack of crop
rotation. Trifolium incarnatum is said to be the most susceptible host. Thus the disease
seems almost identical in its course with the Typhula blight of cereals caused by
T. Itoana.
Pistillaria clavulata; 2 mm. high, white, abruptly clavate: stem somewhat curved,
tomentose-pubescen-t: sclerotium orbicular-depressed, subrufescent, subcuticular:
spores c. 10 µ long, oblong or elliptic: basidia bicuspidate: on dead stems of Des
modium: U.S.A. (S. Jersey). If identical with T. trifolii it supplies an earlier specific
epithet, but compare T. gyram.
T. umbrina Remsberg
Mycol. 32, 1940, 77, f. 15, 33, 39.
Text-Figs. 285b, 286e, 287/.
8-15 mm. high, erect, straight or slightly curved, simple, I-several per sclerotium:
head 3-8 x 0·8-1·5 mm., cylindric-subclavate, blunt, apex fertile, solid, white or
avellaneous, stem 3-4 X 0·2-0·5 mm., minutely puberulous, especially at the base,
rufescent-brown to wood-brown, base often blackish.
On roots of Brassica campestris, and leaves and rhizomes of Iris: Canada.
Spores u·5-15·5 x4-8µ, av. 12·5 x5·5µ, ovate, apiculus slight.
Basidia 31-39 X 6-8µ, 4-spored.
Hyphae of stem and head incrusted.
Sclerotium 0·5-4 mm. wide, convexo-plane, smooth or slightly rough, erumpent or
superficial, dark brown to chestnut, drying blackish, solitary (in culture never coalescent):
cortex composed of a single or double layer of agglutinated hyphae with fairly wide
lumen: cuticle 7-12µ thick, reddish brown: medulla composed of free, interwoven,
incrusted hyphae.
Temperature-range 0-18 ° C., opt. 6-12° C.: sclerotia after 5-7 days in culture:
no sterile fruit-bodies seen in culture.
This species is said to cause a mild necrosis of the host. It seems close to T. latissima
which has a flattened and irregular head and slightly different cortical structure in the
sclerotium. The cortex of T. umbrina seems to have I or 2 layers of agglutinated
hyphae and thus approaches the thicker cortex of T. sclerotioides..
Compare T. gyrans, which also seems to occur on Brassica stems in Europe.
TYPHULA
T. variabilis Riess
Hedw. 1, 1853, 21, t. 3, f. a-b: Remsberg, Mycol. 32, 1940, 79, f. 7, 8, 9, 50, 51, 55.
Synonyms: Sclerotium semen Fr., Syst. Myc. 2, 1822, 249.
Typhula semen Quel., Bull. Soc. Bot. Fr. 24, 1877, 330, t. 6, f. 2 (teste Karst.,
Remsberg).
? T. lactea Tul., Sel. Fung. Carp. 1, 1861, 106 (teste Remsberg).
Text-Figs. 284a, 286d, 287e.
2-7 cm. tall, greyish white or pale greyish bistre to fuliginous or olive-buff, darker at
the apex and base, brownish on drying, simple or occasionally branched, I-several per
sclerotium: head 4-28 X 0·4-3 mm., cylindric-tapered, the apex sterile, becoming
flattened and hollow, sometimes proliferating short spur-like branches; stem 5-65 X
0·3-1 mm., puberulous, pilose towards the base, paler than the head, often pellucid
white at the apex.
On dead stems, leaves, petioles, grasses, and vegetables in cold storage: sclerotia in
spring, fruit-bodies in autumn: N. temp. (Europe, N. Amer., ? Asia).
Spores 9-15·5 X4-6µ, white, ellipsoid or subfusiform, flattened adaxially, smooth,
aguttate, apiculate.
Basidia 27-35 X 6-8µ, clamped: sterigmata 2-4 (2·5-3· 5µ long: ? correct).
Hyphae 3-15µ wide, clamped (Donk): incrusted in the subhymenium, less so in
the stem (Remsberg).
Sclerotia 5-6 X 3-5 mm. (Remsberg), 1-2 mm. when dry (Donk), subglobose to
lenticular, like large Brassica seeds, ochraceous then chestnut-brown or blackish, smooth
then rough and furrowed, superficial, singly or (in culture) coalesced, indented at the
point of attachment to a minute stalk: cortex composed of a single layer of agglutinated
hyphae with wide lumen: cuticle 10-15 µ thick, dark reddish brown, rough: medulla
composed of free, interwoven hyphae, often loose in the centre.
Temperature-range 0-21 ° C., opt. 12-15 ° C.: sclerotia after 7-14 days in culture,
occasionally formed on the surface of old fruit-bodies under wet conditions: sterile
white sporophores occasionally produced in culture.
This species has been confused with the small-spored T. betae, as I have explained
under it, perhaps also with the white T. intermedia. Therefore I am doubtful of the
records that it is a parasite, except for Remsberg's observation that it is weakly patho
genic on stored celery in Canada and U.S.A. Macroscopically it resembles T. phacor
rhiza, but the head is wider, much more distinct, and becomes hollow because the
tissue is only slightly agglutinated: the sclerotial structure is also different. Donk says
that T. variabilis sensu Bourd. et Galz. is T. phacorrhiza. T. semen st:nsu Bourd. et
Galz. seems to be the true T. variabilis but with small fruit-body.' In its typical state,
with unlimited growth, well-developed subhymenium, slight agglutination, and large
spores, T. variabilis seems nearer the primitive Typhula than any other species.
Quelet says (for T. semen) that the sclerotium develops in winter in vegetable heaps:
it is then spherical, glutinous, and white. In summer it lies in the humus and turns
hard, rough, and black. In autumn it fruits.
Tulasne says (for T. variabilis) that the head is simple or variously branched, the
branches occasionally fasciculate, and that it is white at first, becoming cinereous with
age. The fruit-bodies take 15-20 days to reach full size, but others, growing more
slowly, may take over a month. During the day the growing clubs become flaccid and
wilt because of the drier air, but they recover during the night: thus, their growth is
periodically inhibited during the day-time.
T. lactea is said to have milk-white fruit-bodies, often bluish at the tips, and to grow
on dead stems of Zea. Unfortunately there are no microscopical details, and, though
1
Head 2-3 mm. long, not fistular or fluted: stem pilose, arising from a globular, brownish
black sclerotium, grey inside: spores 9-10 x 4 µ, oblong: basidia 30-35 X 6-7 µ,, 2-spored: on
the ground: France.
688 TYPHULA
it has been referred to T. variabilis, it may be T. intermedia. Tulasne observed that
its sclerotia, collected in autumn and winter and placed in moist sand the next spring,
did not fruit until the very end of August. Tulasne's illustration shows. short branches
as occur in T. variabilis.
T. viburni Remsberg
Mycol. 32, 1940, 76, f. 25, 26, 34, 45.
Text-Figs. 285h, 286/, 287g.
2-5 mm. high, simple, singly from the sclerotium: head 1 ·2-2X0·5-1 mm., cylindric,
fusiform ellipsoid or ovoid, white then pale brown, solid: stem -4·5Xo· 1-0·2 mm.,
filiform, puberulous, white, brown at the base.
On dead leaves of Viburnum cassinoides, fruiting in August: U.S.A.
Spores 7·5-10X3·5-5µ., av. 8·5 X4·5µ., ovoid, apiculate.
Basidia 20-24X7-8 µ., 4-spored.
Hyphae incrusted in the head and s·;em.
Sclerotia 0·5-1 mm. wide, convexo-plane, smooth or slightly rough, erumpent,
cinnamon-brown to chestnut-brown, solitary or (in culture) clustered, not coalescent:
medulla wholly agglutinated but the hyphal outlines visible: epidermis 12-20 µ. thick,
consisting of several layers of narrow hyphae with thick, dark-brown, agglutinated
walls, the outer hyphae disrupting as the sclerotium enlarges.
sterile fruit-bodies frequent in culture.
This may be identical with T. elegans. It seems close to T. gyra11s, but with smaller
fruit-body and spores.
T. virgata Remsberg
Mycol. 32, 1940, 81, f. 1, 21.
Text-Fig. 285g, 287g.
4-12 mm. high, white, cylindric, simple, singly from the sclerotium or from the myce
lium (or from stromatic crusts in culture): head 5-11X1·5 mm., cylindric or thickened
upwards, blunt, solid, apex fertile: stem 2-6 x o·1-0·2 mm., 'indistinct', puberulous.
On dead herbaceous stems, fruiting in autumn: U.S.A.
Spores 9-15X4-6 µ., av. 12X4·5µ., ellipsoid, apiculate.
Basidia 25-30X6-8 µ.: sterigmata 2-4.
Hyphae more or less incrusted in the head and stem.
Sclerotia 0·5-0·8 mm. wide, glabrous or flattened, brown, erumpent, simple or (in
culture) coalescent: cortex composed of a single layer of agglutinated hyphae with wide
lumen: cuticle 5-10µ. thick, brown: medulla composed of free, interwoven hyphae,
loose in the centre.
sterile fruit-bodies not seen in culture.
The description suggests a relatively stout, cylindric, or subclavate head and very
slender stem, but the figure shows only a wholly subcylindric fruit-body.
Postscriptum. Vang (1945, see p. 703) has redescribed T. betae as T. brassicae

(Berg. ex Fr.) comb. nov., based on Sclerotium brassicae Berg. ex Fr., Syst. Myc. 2,
1823, 259, but this name refers to the imperfect state and does not invalidate Rostrup's
based on the perfect. According to Vang, T. betae belongs to subgen. Subcarnosae,
having the fruit-body and spores of T. sclerotioides and the sclerotia of T. variabilis.
T. gyrans sensu MacDonald (see p. 683) may be T. betae. Vang's data are: fruit-body
-2(-3) cm. high, head 4-7X0·5-0-·75 mm.: spores 6-9X3-4µ.: basidia 4-spored:
sclerotium 0·5-2 mm., white, then brown to black, rind 9-15 µ. thick, eortex with
a single layer of hyphae with wide lumen, medulla of free interwoven hyphae: sapro
phytic on Beta and Brassica.
CLAVARIOID GENERA
SYNONYMS AND TYPE-SPECIES
Genus Synonyms Type-species

Aphelaria Corner A. dendroides (Jungh.) Comer
Araeocoryne Corner A. elegans Corner
Caripia O. K. Hypolyssus B. (non Pers.) C. Montagnei (B.) O.K.
Perona Fr. (non Pers.) ,, ,,
Ceratellopsis Konr. et Maub. Ceratella Pat. (non Hook.) C. aculeata (Pat.) Corner
Chaetotyphula Corner C. hyalina (Jungh.) Corner
Clavaria Fr. s. str. C. vermicularis Fr.
Holocoryne Bon. C. acu.ta Fr.
Stichoclavaria Ulbr.
Clavariachaete Corner C. rubiginosum (Cke.) Corner
Clavariadelphus Donk C. pistillaris (Fr.) Donk
Clavicorona Doty C. taxophila (Thom) Doty
Clavulina Schroet. Stichoramaria Ulbr. C. cristata (Fr.) Schroet.
Clavulinopsis v. Ov. C. miniata (B.) Corner
Cornicu.laria Bon. C. corniculata Fr.
Defiexula co·rner D. fascicularis (Bres. et Pat.)
Corner
Dimorphocystis Corner D. laevis Corner
Hormomitaria Corner H. su.lphu.rea Corner
Lachnocladium Lev. Eriocladus Lev.
Stelligera Heim ex Doty S. rnembranacea Heim ex Doty
Lentaria Corner L. su.rculu.s (B.) Corner
Mucronella Fr. Mu.cronia Fr. M. ·aggregata Fr.
Myxomycidium Mass. l'vl. pendulum Mass.
Physalacria Pk. Eoagaricus Krieger P. infiata (Schw.) Pk.
Baumanniella P. Henn. P. togoensis (P. Henn.) Corner
Pistillaria Fr.
Cnazonaria Cda. P. setipes Grev.
Scleromitra Cda. P. micans Fr.
Sphaerula Pat. P. capitata (Pat.) Sacc.
Gliocoryne Maire P. uncialis (Grev.) Const. et Duf.
Pistillina Quel. P. hyalina Quel.
Pterula Fr. P. su.b11/ata Fr.
Penicillaria Chev. P. multifida Fr.
Phaeopterula P. Henn. P. hirsu.ta P. Henn.
Pterulicium Corner P. xylogenum (B. et Br.) Corner
Ramaria S. F. Gray R. botrytis (Fr.) Ricken
Ramaria Bon., Quel.
Clavariella Karst.
Cladaria Ritg.
Phaeoclavu.lina Brinkm. R. macrospora (Brinkm.) Corner
Dendrocladium Lloyd
Ramariopsis Donk R. Kunzei (Fr.) Donk
Scytinopogon Singer S. angulisporus (Pat.) Corner
Typlmla Fr. T. phacorrhiza Fr.
Phacorrlziza Pers. T. sclerotioides (Pers.) Fr.
Pistillaria sensu Donk T. quisquiliaris (Fr.) Corner
5119 Yy
NOVITATES
Aphelaria gen. nov.
Receptacula clavarioidea applanato-ramosa, rarius subsimplicia, coriacea vel
gelatinosa, alba vel pallida, plerumque ad terram, raro ad lignum: sporis albis, levibus,
subglobosis vel ellipsoideis, aguttatis vel minute guttulatis: hymenio spissato, cystidiis
deficientibus: hyphis hyalinis, haud inflatis, plus minus crasse tunicatis, fibulis
praeditis aut typice deficientibus.
A. incarnata sp. nov.
Ree. ad :z cm. alta, parum ramosa, alba dein pallide ochracea vel incarnata, coriacea :
stipite glabro vel basi strigoso: sporis 11-15 X 5-6 /J-, elongato-ovoideis: sterigmatibus
:z(-4): hyphis ut in A. spiculosa: ad terram in silvis, Singapore.
A. spiculosa sp. nov.

Ree. ad 3 cm. alta, !axe ramosa, ramulis attenuatis, filiformibus, alba dein sub
incarnata, lenta, coriacea: stipite plerumque ex integro spiculoso-strigoso: sporis
8-10·5 X 5·5-7 /J-, ellipsoideis, aguttatis: sterigmatibus (:z-)4: hyphis :z-3 ·5 /J- latis,
fibulis deficientibus: ad terram in silvis, Malaya, ins. Philippin.
Araeocoryne gen. nov.

Receptacula simplicia, erecta, parva, gracilia, apice subtruncato sterili, stlp1te
distincto, carne molli vel ceracea, ad ramulos et folia dejecta: sporis albis, ievibus,
aguttatis, ellipsoideis: hymenio spissato, cystidiis deficientibus: hyphis inflatis, fibulis
praeditis, tenuiter tunicatis.
A. elegans sp. nov.
Ree. ad 15 mm. alta, alba, stipite disco byssoideo affixo: sporis 10-1:zx6-8 f-,L:
Malaya (Johore).
Chaetotyphula gen. nov.

Receptacula pusilla, clavulata, st1p1tata, simplicia, ut in Pistillaria sed hymenio
basim versus evoluto: ad lignum, folia dejecta, et truncos: sporis albis, levibus, aguttatis
vel 1-:z-guttulatis: basidiis 1-:z sterigmatibus praeditis: cystidiis crasse tunicatis: hyphis
septatis, inflatis, fibulis deficientibus.
C. hyalina (Jungh.) Corner var. microcystis var. nov.
Cystidiis minoribus 30-50 X 5-7 /J-, paucis: ad folia dejecta arborum et palmarum,
Malaya (Singapore).
Clavaria Fr. emend.

Receptacula radiato-ramosa vel simplicia, ceracea, rarius subleni:a, ad terram: sporis
hyalinis, albis, vel incarnatis, levibus, saepius aguttatis vel minute guttulatis: hyphis
septis ordinis secundi praeditis, fibulis deficientibus vel solum ad basim basidiorum.
Subgen. Syncoryne Fr. emend.

Basidiis fibulis deficientibus: sporis saepius aguttatis: hyphis plerumque ordine
secundo septatis: receptacula ramosa vel simplicia: C. vermiculari typica.
Subgen. Holocoryne Fr. emend.
Basidiis fibulis !axis praeditis: sporis minute guttulatis: hyphis septis ordinis
secundi saepe deficientibus: rec. simplicia: C. acuta typica.
C. argillacea var. brevispora var. nov.
Sporis brevioribus, 6-9 X 4-5 /J-: Gallia.
NOVITATES
C. argillacea var. gracillima var. nov.
Ree. -2·5 cm. alta, gracilia, flavida, stipite laete flavo: sporis 7·5-10 X 3·5-4µ,
minoribus: Am. bor.
C. argillacea var. sphagnicola var. nov.

Ree. ad II cm. X 2-7 mm., majora, saepe compressa: sporis 9-11·5 X 5·5-7µ,
latioribus: inter Sphagna, Gallia.
C. atrobadia nom. nov.

Ree. ad 10 cm. alta, simplicia, caespitosa, rufo-badia dein nigricantia, fragilia, odore
exigue farinoso: sporis 5 X 2-2 · 5 µ: inter gramina, Europa.
C. atro-umbrina nom. nov.

Ree. ad 7 cm. alta, simplicia, solitaria vel subcaespitosa, umbrina dein fuligineo
brunnea, hymenio mox longitud. ruguloso, came molli elastica, inodora: sporis
4·5-6·5 X 2·2-3"2µ: inter gramina et muscos, Am. bor.
C. filiola sp. nov.

Ree. ad 15 mm. alta, pusilla, alba, simplicia, stip1te distincto, ceracea, gregaria:
sporis 4 ·5-5 X 4µ, subglobosis, aguttatis: basidiis fibulis deficientibus: hymenio non
spissato: hyphis 4-8µ latis, inflatis, sparsim ordine secundo septatis: ad terram in
silvis, Singapore.
C. fossicola sp. nov.

Ree. ad 12 mm. alta, pusilla, simplicia vel rarius subramosa, alba dein saepe cremea
vel isabellina, subulata dein fusiformia et obtusa, tenacia, subcartilaginea, inodora:
sporis 5-6·5 X 2-2·7µ, aguttatis vel 1-2-guttulatis: hymenio spissato: basidiis fibulis
deficientibus: hyphis 3-10µ latis, non vel raro ordine secundo septatis: ad terram
inter Protococcoideas, Malaya.
C. Gibbseae var. tenuis var. nov.

Ree. ad 5·5 cm. x 1-2·5 mm., solitaria vel gregaria, graciliora: sporis 8·5-10 X 7-8µ:
ad terram, Malaya.
C. Gibbseae var. tenuis f. microspora form. nov.

*
Sporis 6-7·5 X 5·5-7µ, minoribus, subglobosis: ad terram, Malaya.
C. Gibbseae var. megaspora var. nov.

Sporis 9-13 X 6-8µ, late ellipsoideis, majoribus: ad terram, Malaya.
C. rosea var. subglobosa var. nov.

Sporis 5-8 X 4-6µ, subglobosis vel late ellipsoideis: ad terram, Germania, Anglia.
C. vermicularis var. singaporensis var. nov.

Sporis 7-8·5 X 5-6µ, majoribus, ovoideis, 1-guttatis: ad terram, Singapore: rec. ad
_4 cm. alta, gregaria.
Clavariachaete gen. nov.

Receptacula ramosa, ochraceo-ferruginea, coriacea, lignicola: hymenio amphigeno
non spissato, setulis tunicis brunneis incrassatis praedito, gloeocystidiis et dicho
physibus deficientibus: sporis?: hyphis non inflatis, tunicis tenuibus vel leniter
incrassatis, saepius flavido-brunneolis, fibulis deficientibus: Am. austr.
692 NOVITATES
Clavariadelphus Donk
Subgen. nov. Eu-clavariadelphus
Receptacula ampla, apice fertili, clavata v. ligulata, came spongiosa, hyphis multo
intertextis.
Typus: C. pistillaris (Fr.) Donk
Subgen. nov. Cantherellopsis
Ree. ampla, turbinata v. subpileata, apice sterili, carne spongiosa, saepe cava, hyphis
multo intertextis.
Typus: C. truncatus (Quel.) Donk
Subgen. nov. Typhulopsis
Ree. plus minus gracilia, subclavata v. filiformia, apice fertili, came rigido-succosa,
hyphis longitudinalibus, stipitis superficie agglutinatis: saepe ramicola, lignicola v.
epiphylla.
Typus: C. junceus (Fr.).
C. pistillaris (Fr.) Donk
var. americanus var. nov.
Ree. saepe incarnata v. rosea, dein ochracea v. alutacea, saepe complanata v.
ligulata, nonnunquam breviter ramosa cervicornia, solitaria v. caespitulosa: sporis
minoribus 7-12·5 X 3·7-7·5µ, av. 9·7 X 5·4µ.: in silvis frondosis, raro pinetis, Am. bor.
var. atrobrunneus var. nov.
Ree. brunneo-umbrina v. atrobrunnea, basim versus dilutiora, stipite flavido
cinnamomeo, came alba: sporis 10-12 X 6-8µ.: in pinetis, China.
Clavulina Schroet.
Subgen. nov. Fusco-clavulina
Hyphae tunicis leviter incrassatis, brunneolis, vix inflatae: basidiis brevibus,
20-40µ. longis.
Typus: C. ornatipes (Pk.) Corner
Subgen. nov. Eu-clavulina
Hyphae incoloratae, plus minus inflatae: basidiis saepe longioribus.
Typus: C. cristata (Fr.) Schroet.
C. cavipes sp. nov.
Ree. ad 7'5 cm. alta, sparsim ramosa, bifurcata vel palmata, nonnunquam simplicia,
subcaespitosa vel solitaria, alba, dein ramis ramulisque carneo-griseis, vinaceis, vel
incarnatis, stipite saepe cavo: sporis 8-10 X 7-8·5 µ., subglobosis, 1-guttatis: hymenio
incrassato: hyphis ut in C. cristata: ad terram et lignum putridum in silvis, Malaya.
Subsp. nov. Eucristata
Ramulis plus minus cristatis, non cinerascentibus.
var. incarnata var. nov.
Receptacula alba, dein incarnata, aetate brunneola vel griseola, ramis cristatis dein
elongatis subulatis: sporis 8-10·5 X 7-9µ., v. 7-9 X 6-7·5 µ.: in locis humidis, hymenio
cystidiato: in locis paludosis, inter humum v. ad folia graminea putrida, Anglia.
Subsp. nov. Coralloides
Ramulis vix v. non cristatis, non cinerascentibus.
NOVITATES
var. coralloides var. nov.
Ree. ad 10 cm. alta: ram1s v1x cristatis, elongatis: Europa: ( = Clavaria coral
loides Fr.).
var. subrugosa var. nov.
Ree. sparsim ramosa: ramis crassis, obtusis, non cristatis, simplicibus v. semel
bifurcatis: sporis 7-10x6-8·5 µ,: Europa.
Subsp. nov. Cinerascens
Ramulis phi.s minus cristatis, cinerascentibus.
C. decipiens sp. nov.
Ree. ad 5 cm. alta, caespitosa v. solitaria, pallido-cinnamomea, Pteruliformia,
fibrosa: stipite saepe ph,1s minus strigoso-spiculoso: ramulis paucis, gracilibus, fasti
giatis, subpalmatis v. bifurcatis, apicibus subulatis, breviter filiformibus: sporis
8-10X7-9µ,, tunica brunneola, subglobosis, 1-guttatis: basidiis 20-40X 5·5-7µ,:
hymenio incrassato, unilaterali: hyphis 3-7µ, latis, vix inflatis, tunicis brunneolis,
fibulatis, ut in C. ornatipes: ad terram in silvis, Malaya.
C. gallica nom. nov.
Synonym: Clavaria alba sensu Bourd. et Galz., Bull. Soc. Myc. Fr. 26, 1910, 21 I.
C. gigartinoides sp. nov.
Ree. ad 15X9 cm., caespitosa, valde ramosa, ramis palmato-divisis, saepe connatis,
ramulis saepe cristatis, pallide flavidula, apicibus cremeis, dein pallide cervina v.
subfuliginea: came tactu subrufescenti, subfragili: hymenio ramis inferioribus uni
laterali, subcervino, incrassato: sporis 7-8·5X 6·5-7"5µ,, subglobosis, 1-guttatis:
basidiis 38-55 x 6--7µ,: hyphis ut in C. cristata: ad terram in silvis, Malaya.
C. gracilis sp. nov.
Ree. ad. 6 cm.X 1-2 mm., simplicia, solitaria v. gregaria, cylindrica dein subclavata,
gracilia, ceracea, breviter stipitata, inodora: hymenio levi, incrassato: sporis 8-10µ,,
globosis, 1-guttatis: basidiis 40-60X8-10µ,: hyphis ut in C. cristata: ad terram in
silvis, Malaya.
C. ingrata sp. nov.
Ree. ad 9 cm.X 2-3 ·5 mm., vel compressa ad 6 mm. lata, solitaria v. caespitulosa,
simplicia, fusiformia, alba dein flavidula, ceracea, odore forti ut in Trigonella, sapore
amaro ingrato: sporis 10-12X9·5-10·5µ,, subglobosis, 1-guttatis: basidiis 60-75X
7-9µ,: hyphis ut in C. cristata: ad terram in silvis, Malaya.
var. atricha var. nov.
Cystidiis deficientibus: ad terram, in silvis, Malaya.
var. alcyonaria var. nov.
Ree. ramosa, ramis paucis simplicibus v. 1-3 furcatis, obtusis, elongatis, non
cristatis: sporis 9-12X7-9µ,: Europa.
var. canaliculata var. nov.
Ree. ad 20 cm. alta, longitud. sulcata, non rugulosa, tandem cava: sporis 8-12X
7-9µ,: Europa.
var. macrospora var. nov.
Sporis 12-14x8-10µ,: Europa.
694 NOVITATES
Clavulinopsis v. Ov. emend.
Receptacula radiato-ramosa vel simplicia, ceracea vel sublenta, saepe laete colorata,
ad terram, raro ad lignum vel ad fragmenta vegetabilia emortua: sporis albis, levibus
vel valde aculeato, saepius 1-guttatis: hyphis inflatis, fibulis praeditis, non ordine
secundo septatis.
Subgen. Acularia subgen. nov.
Sporis valde aculeatis: C. helvola typica.
Subgen. Eu-Clavulinopsis subgen. nov.
Sporis levibus: C. miniata typica.
C. arenicola sp. nov.
Ree. ad 2·5 cm. alta, simplicia, sparsa v. gregaria, compressa, ligulata, fusiformia
v. obtusa, alba dein pallide ochracea e basi apicem versus, odore forti (ut Agarico
campestri): sporis 7-8 X 5-6µ, piriformibus v. lacrimiformibus, 1-guttatis: hymenio
spissato: hyphis 3-12µ latis: ad terram arenosam, Malaya.
C. brevipes sp. nov.
Ree. ad 7 cm. alta, simplicia, sparsa, gregaria v. subfasciculata, subclavata, alba dein
cremea, stipite distincto, sapore amaro ingrato: sporis 5-7·5 X 4-5·7µ, ovoideis,
1-guttatis: hymenio spissato: hyphis 6-18µ latis: ad terram in silvis, Singapore.
C. filipes nom. nov.
Ree. ad 2·5 cm. alta, simplicia, solitaria v. gregaria, cylindrica, filiformia v. sub
clavata, pallide albo-cremea vel subvirescentia, stipite distincto subincarnato, ceracea:
sporis 5�7"5 X 3·5-4·5µ: Am. bor.
C. Jruticula sp. nov.
Ree. ad 2 cm. alta, ramosa, solitaria v. gregaria, alba dein pallide ochracea, tandem
pallide brunneo-ochracea e basi usque ad ramos majores, ceracea, inodora: sporis
4-5 X 2·5-3·5µ, ovoideis, 1-guttatis: hymenio spissato: hyphis 2-15µ latis: ad terram
umbrosam, Malaya,
C. hastula sp. nov.
Ree. ad 7 mm. alta, pusilla, spathulata vel lanceolata, compressa, tenuissima, rarius
apici 2-3-fido, pallide incarnata v. cremea, stipite brevi, came molli subgelatinosa:
sporis 5--7 X 2·5-3µ, ellipsoideis: basidiis pyriformibus sterigmatibus 4-6 praeditis:
hymenio spissato: hyphis 2-2·5µ latis, hie inde 5-12µ inflatis, tunicis gelatinosis: ad
terram umbrosam inter Cyanophyceas, Malaya.
C. helvola var. geoglossoides var. nov.
Ree. compressa, canaliculata: sporis 6-9 X 4-8µ, vel 6-8µ subglobosis: inter
gramina, Gallia.
C. luteo-alba var. latispora var. nov.
Sporis. 6-10 X 4-6µ, latioribus: Europa.
C. luteo-alba var. longispora var. nov.
Sporis 10-12(-15) X 5-6(-8) µ, longioribus: Germania, Italia.
C. miniata var. rosacea var. nov.
Receptacula carnea vel rosacea, minus colorata: Asia trop.
C. miniata var. sanguinea var. nov.
Receptacula parva, ad 7·5 cm. alta, rubra, punicea vel cruenta: ad terram, lignum
putridum et folia dejecta: Asia et Australasia trop.
NOVITATES 695
C. ochracea nom. nov.
Ree. ad. 2·6 cm. alta, simplicia, caespitosa, ochracea: sporis 7-9x4·5µ, ovoideis,
1-guttatis: ad terram, Africa austr.
C. similis sp. nov.

Ree. ad 4 cm. alta, simplicia vel saepius semel v. bis dichotoma, solitaria v. sub
caespitosa, alba dein flavescentia, ceracea, ramulis elongatis acutis: sporis 4 ·s,-6x
2·5-3µ, ellipsoideis: hymenio? non spissato: hyphis 2·5-8µ latis: ad terram in silvis,
Malaya.
Deflexula gen. nov.

Receptacula parva, ad 25 mm. longa, ut spinae subulatae caespitulosae, deflexae,
simplices v. ramosae, ceraceo-coriacea, spinis e basi spinae primae proliferanti
bus: sporis albis, levibus, globosis v. ellipsoideis, magnis, aguttatis: basidiis magnis,
sterigmatibus 4: cystidiis deficientibus: hymenio incrassato: hyphis biformibus, aut
skeletalibus aseptatis, tunica incrassata, haud ramosis, aut ramosis, tunica tenui,
septatis fibulatis: ad lignum: genus Pterulae affinis, superficie Hydnaceus: typus, D.
Jascicularis (Bres. et Pat.).
D. lilaceo-brunnea sp. nov.

Ree. parva, ad 3·5 X0·2-0·5 mm., 2-20-fasciculata, spinis subulatis simplicibus,
alba dein flavida, brunneola, lilacina, v. subviolacea: sporis 12-14µ latis, globosis:
basidiis 50-75 X15-18µ: hyphis skeletalibus 3-5(-7)µ latis: ad ramos emortuos in
silvis, Malaya, Sumatra.
Dimorphocystis gen. nov.

Ut Pterula sed receptaculis pusillis, 0·5-20 mm. altis, simplicibus, filiformibus vel
capitulatis, albis: sporis albis, levibus, ellipsoideis, aguttatis: cystidiis biformibus, aut
fusoideis, conicis, elongatis, aut brevioribus apicibus digitatolobulatis, plus minus
crasse tunicatis: hymenio non incrassato: hyphis biformibus, ut in Pterula: ad folia
emortua in silvis, Malaya.
D. capitatus sp. nov.

Ree. ad o·6 mm. alta, perpusilla, capitulata, erecta v. inversa: stipite -500x40-50 µ,
glabro, hyalino: capitulo 140-180µ lato, subgloboso, hispidulo: sporis 6-7 X2·5-3·5µ:
basidiis 4-sporig.: cystidiis fusoideis 40-95x8-15µ, tunicis 3-6µ crassis, rugulosis
v. leviter incrustatis: cystidiis lobulatis 18-40 X5-8µ, lobulis incrustatis: hyphis
fibulatis: amphigena, ad folia dejecta, Singapore.
D. laevis sp. nov.

Ree. ad 18 mm. alta, 150-200µ lata, filiformia, erecta: stipite 2-5 mm. X200-280µ:
sporis u-14(-16) X4-5µ: basidiis 2-sporig.: cystidiis fusoideis 40-75 X10-18µ,
tunicis 0·5-2µ crassis, levibus: cystidiis lobulatis 20-35 X5-12µ, levibus: hyphis non
fibulatis: hypergena, ad folia dejecta Eugeniae cerinae, Singapore.
D. subcapitatus sp. nov.

Ree. o·5-4 mm. alta, stipitata, subglobosa v. subclavata: stipite 200-800 X30-70µ,
subinde ad 3·5 mm. longo, puberulo: capitulo 90-350 X130-250µ, subgloboso v.
subclavato, puberulo: sporis 11-13 X3-4 µ: basidiis 2-sporig.: cystidiis fusoideis
30--75 X 9-15µ, tunicis 1-4µ crassis, leviter rugulosis, non incrustatis: cystidiis
lobulatis 18-35 X5-9µ, leviter incrustatis: hyphis non fibulatis: hypogena, ad folia
dejecta Eugeniae cerinae cum D. laevi, Singapore.
NOVITATES
Hormomitaria gen. nov.
Reeeptaeula simplieia, pusilla, eaespitosa, inversa, pendentia, st1pite brevi, clavula
eonieo-aeuminata elongata, ad basim clavulae sterilia, eeraceo-subgelatinosa, plena,
lignicola: sporis hyalinis, levibus, ellipsoideis, aguttatis: hymenio oleoeystidiis im
mersis praedito, non spissato: hyphis inflatis, in stipite saepius ordine secundo septatis,
in clavula ordine secundo moniliforme septatis, tunicis subgelatinosis: caulocystidiis
cylindrieis v. clavatis.
H. sulphurea sp. nov.
Ree. ad 12 mm. longa, clavula sulphurea: stipite 0·3-0·8Xo·15-0·3 mm., alba:
sporis 5-6X3-3·5µ.: Malaya.
Lachnocladium Lev. emend.

Reeeptaeula applanato-ramosa, tenacia, subcoriacea, flavida, oehracea, ferruginea
vel cinnamomea: sporis albis, levibus, aguttatis, parvis: hymenio saepe unilaterali et
incrassato, gloeoeystidiis praedito: hyphis non inflatis, fibulis deficientibus, dicho
physibus praeditis: came ae medulla hyphis longitudinalibus, cortice dichophysibus,
construeta.
L. divaricatum var. cinnamomeum var. nov.
Ree. ad 10 X 8 cm., cinnamomeo-ochracea, ramulis laete oehraeeis, odore cimicario:
sporis 3-3·5 X2·5µ.: hymenio spissato: medulla in stipite ramisque 100-350µ. lata,
cortice 200-400µ. crassis: ad humum et lignum putridum in silvis, Malaya.
L. divaricatum var. sordidum var. nov.
Ree. ad 15 cm. alta, stricta, fastigiata, sordide cervina, ramulis flavidis, odore acerbo
(ut in Ganodermati): sporis 3-3·.5X2·5-3µ.: hymenio,.medulla, et cortice ut in var.
cinnamomeo: ad truneos in silvis, Malaya.
L. fiavidum sp. nov.
Ree. ad 7 cm. alta, dense ramosa, fastigiata, sordide flavida, apicibus flavo-albis,
? inodora: sporis 3-3·5 X2·5µ.: hymenio non spissato: medulla 100-250µ. latis: ad
lignum putridum in silvis, Malaya.
L. fulvum sp. nov.
Ree. ad 9 cm. alta, dense ramosa, fulvo-oehracea, ramulis laete flavis, apicibus albis,
odore acerbo: sporis 3·5-4·3x3-3·5µ., apiculo conspicuo 1·5µ. longo: hymenio non
spissato: medulla 150-350µ. lata: ad humum et ramulos emortuos in silvis, Malaya.
L. molle sp. nov.
Ree. ad 7·5 cm. alta, dense ramosa, ramulis divaricatis, !axe aseendentibus, sordide
ochraceo, carne molli: sporis 3-3·5X2·5µ.: hymenio spissata: medulla m ram1s
600-800µ. lata, eortiee 100-200µ. crasso: ad humum in silvis, Malaya.
L. zonatum sp. nov.
Ree. ad 5· s cm. alta, dense ramosa, divarieata, cervino-ochraeea, basim versus sub
ferruginea: sporis 3-3·5 X2·5-3·3µ.: hymenio non spissato: eortiee zonato-spissato,
zona utraque basidiis in strato simplici instructis: medulla 100-300µ. lata: ad terram
et ramulos emortuos in silvis, Singapore.
Lentaria gen. nov.

Receptacula radiato-ramosa, rarius simplicia, lenta, cartilaginea vel coriaeea, alba,
alutaeea, ochraeea, incarnata, vel brunneola, saepissime mycelio byssoideo copioso
exorientia, sapore saepe amaro, lignicola: sporis albis (vel suboehraceis ?), ellipsoideis,
NOVITATES
saepius elongatis, aguttatis: hymenio saepe spissato: cystidiis deficientibus: hyphis
plus minus inflatis, tunicis incrassatis, fibulis praeditis.
Subgen. Eu-Lentaria subgen. nov.
Sporis elongatis, ultra 7 µ longis: hyphis tunicis valde incrassatis: receptaculis dense
ramosis: L. surculus typica.
Subgen. Lentariopsis subgen. nov.
Sporis parvis, 4-7µ longis: hyphis tunicis leniter incrassatis: receptaculi� parvis,
saepe paulum ramosis, vel simplicibus: L. epichnoa typica.
Mucronella Fr.
M. pusilla sp. nov.
Receptacula ad 500 X 50-130µ, albo-flavidula v. aetate sordide ochracea, conica,
apice sterili, stipite brevi (10-75 X 30-50 µ), dense gregaria sed discreta, subiculo
nullo: sporis 4-4·7 X 2·7-3·3 µ, ellipsoideis, aguttatis, levibus: basidiis II-15 X 4-5 µ,
4-sporig.: cystidiis nullis: hyphis uniformibus, 2-7 µ latis, fibulatis, tunicis flavidulis:
ad lignum emortuum ramosque dejectos, Malaya.
Physalacria Pk.
P. australiensis sp. nov.
Synonym: P. infiata (Schw.) Pk. sensu Fawcett, Proc. Roy. Soc. Viet. 52, 1940, 175,
t. 7, f. 2-3, Text-fig. 1 A, B, c.
Receptacula ad 10 X (-13) mm. alta, solitaria v. 2-3 gregaria, rarius caespitosa,
albida dein fl.avidula v. brunneola: stipite 3-6 X 1-1·5 mm., puberulo: clavula 3-5 mm.
lata, irreg. globosa, basim versus saepe plicata: sporis 3-4 X 2-3µ: basidiis 2-sporigeris:
cystidiis subventricosis, tenuiter tunicatis, resino-incrustatis: ad lignum, Victoria.
P. corticola sp. nov.
Ree. ad 1·2 mm. alta, pusilla, gregaria v. solitaria, alba: stipite 200-500 X 80-150µ,
sparsim puberulo: clavula 0·3-0·7 mm., lata, subglobosa dein compressa v. subplicata,
basi subumbilicata: sporis 5·5-6·5 X 2·5-3µ, levibus: hymenio ex integro fertili: oleo
cystidiis 35-50 X 8-17µ: caulocystidiis ad 30 X 3-7µ, ut basidiis sterilibus v. oleo
cystidiis parvis: ad corticem emortuum et lignum, Singapore.
P. tropica sp. nov.
Ree. ad 22 mm. alta, caespitosa, alba, stipite e basi brunnescenti: stipite 3-12 x
0·2-0·6 mm., pruinoso-puberulo: clavula 4-10 mm. lata, subglobosa, mox compressa
v. irregulariter plicata, saepe subcerebriformi, basi umbilicato et saepe lacunis 3-6
ovalibus circumcincto: sporis 4·5-5·5 X 2·5-3µ, levibus: hymenio basi clavulae sterili,
cystidiis fusiformibus 30-70 X 7-16 _µ, levibus, tenuiter tunicatis praedito: hymenio
fertili oleocystidiis plus minus immersis 35-65 X 10-18 µ praedito: caulocystidiis ad
80 X 3-7(-13)µ, cylindricis, clavatis, v. subventricosis, saepius flexuosis, tunicis leniter
incrassatis et flavescentibus: ad ligna et truncos, Malaya.
Pistillaria Fr.
P. rhodocionides sp. nov.
Receptacula 1-2 mm. alta, simplicia, roseo-incamata, raro albida, gregaria: capitulo
150-250 µ lato, subfusiformi v. subclavato, farto: stipite 100-600 X 100-140 µ, glabro,
concolori: sporis 7-9·5 x4·3-5·3µ, albis, ellipsoideis, levibus, aguttatis: basidiis
21-33 x7·5-9µ: sterigmatibus 4, 10-12µ longis: hyphis uniformibus, 3-7µ latis,
tenuiter tunicatis, non fibulatis, paulum infl.atis, intus pallide roseis: ad legumina
emortua Cassiae alatae, Singapore.
698 NOVITATES
P. subuncialis sp. nov.
Receptacula ad 15X1 mm., simplicia, raro furcata, cylindricea v. vix subclavata,
stipite indistincto, sordide flavidula: sporis 10-13X5-6 µ,, albis, levibus, elongato
ellipsoideis, aguttatis: basidiis 20-30X7-10 µ,: sterigmatibus 4, 7-10 µ, longis:
cystidiis -35X17 µ, ventricosis, obtusis, ut basidiis sterilibus inflatis: ad culmos
emortuos Junci, Anglia.
Pterula Fr.
P. brunneola sp. nov.
Receptacula ad 9X12 cm., valde ramosa, caespitosa, pallide cervina, basim versus
subfuliginea, ramulis brunneolis aoicibus albidis, exigue pisciodora: stipite 3-25X
1·5-2 mm.: ramis 1 mm. latis, 2-3-4-furcatis v. palmato-divisis: ramulis gracilibus
furcatis, apicibus attenuatis sed vix filiformibus: sporis 6-7 X3-3·5 µ,, albis, levibus,
ellipsoideis: basidiis 20-30X 6-7µ, 4-sporig.: cystidiis nullis: hymenio ad 100µ
incrassato, ad ramos majores hypogeno: hyphis skeletalibus 2-5 µ, latis, tunicis
0·5-1 µ, crassis, alteris fibulatis: ad humum in silvis, Malaya.
P. debilis sp. nov.
Ree. ad 5 cm. alta, saepe multo breviora, paulum ramosa, ramulis saep1ss1me
indivisis, flaccida, gracilia, tenacia, alba dein e basi brunneolescentia: ramulis spuriis
100-200(-300)µ Iatis, filiformibus, !axe v. subfastigiate dispositis: sporis 7-9·5 x
3·5-4 µ,, albis, levibus, ellipsoideis, aguttatis v. 1-2 minute guttulatis: basidiis
19-26X6-7µ, 4-sporig.: cystidiis 30-40X5-8 µ,, sparsis, tenuiter tunicatis, subven
tricosis: hymenio non incrassato: hyphis biformibus, skeletalibus 3-5 µ, latis, tunicis
0·5-1 µ, crassis, alteris fibulatis: ad culmos emortuos Junci, Anglia.
P. epiphylla sp. nov.
Ree. ad 18 mm.X120-250µ, simplicia, filiformia, erecta, alba dein flavidula:
stipite 2-4X0·2-0·3 mm., fuscescenti, erumpenti, puberulo: sporis 9-13(-14)X4-5 µ,,
albis, Ievibus, aguttatis: basidiis 22-28X8-10µ, 4-sporig.: cystidiis 30-50X5-11µ,
conicis v. subventricosis, tenuiter tunicatis, obtusis: hymenio non incrassato: caulo
cystidiis -45X 4-11µ, tunicis brunneolis subincrassatis: hyphis biformibus, skele
talibus 3-5 µ, latis, tunicis 0·5-1·5 µ, crassis, alteris fibulatis, saepe e fibula ramosis:
stipite non epithelio vestito: ad folia dejecta (lxonanthes reticulata, Linacearum),
Singapore.
P. rigida Donk sp. nov.
Ree. ad 3·8 x 0·4-0·7 mm., Iinearia, apicem versus attenuata, hie illic uno v. pluribus
ramis ascendentibus, candida v. flavescentia, rigida, in sicco subcorneola, stipite nullo,
e subiculo circumscripto subvilloso nascentia, solitaria v. 2-4 gregaria: sporis
(13-)15-:17X(6-)6·5-7·5 µ, albis, levibus, ellipsoideis: hyphis 2-3·5 µ, latis, plus minus
crasse tunicatis: ad lignum mucidum, Hollandia.
P. subtyphuloides sp. nov.
Ree. ad 20 mm. alta, ut in P. typhuloides var. minor: sporis longioribus 10-13X4-5 µ,:
hymenio -120µ incrassato: cystidiis nullis: stipite epithelio 15-30µ crasso vestito, ex
hyphis 2-4µ latis, dense pseudoparenchymaticis, tunicis exterioribus fuscidulis, com
positis: hyphis saepe fibulatis: ad folia et caules Costi (Zingiberacearum), Singapore.
P. typhuloides sp. nov.
Ree. ad 35 mm. alta, 0·5-1·3 mm.1ata, solitaria v. gregaria, simplicia v. ramis 1-2,
filiformia, subulata, alba dein flavescentia v. subochracea: stipite 2-4 mm. Iongo,
fusco, puberulo: sporis 9-10X4-5 µ,, albis, levibus, ellipsoideis, aguttatis: basidiis
24-30 X7-9 µ,, 4-sporig.: cystidiis nullis: hymenio non incrassato; caulocystidiis
NOVITATES 699
-50 X5-8µ, comc1s v. subcylindraceis, saepe acutis, tumc1s brunneolis, paulum
incrassatis: stipite epithelio tenui vestito, ex hyphis 4-8µ latis, pseudoparenchy
maticis, tunicis exterioribus fuscidulis, compositis: hyphis biformibus, skeletalibus
3-5 µ latis, tunicis 0·5-1 ·s µ incrassatis, alteris non fibulatis: ad folia emortua et
inflorescentias dejectas in silvis, Singapore.
var. minor var. nov.

Ree. ad 11 X0·1-0·25 mm., graciliora: sporis 8-10 X3·5-4·5µ: h yphis raro fibulatis:
ad gramina emortua, Singapore.
P. verticillata sp. nov.

Ree. ad 6 cm. alta, valde ramosa, caespitosa v. gregaria, pallide sordido-incarnata,
flavidula, v. pallide cervina, apicibus albis: stipite 3-15 X1-2·5 mm., plus minus
strigoso: ramis 3-4 verticillatis, 0·5-1 mm. latis: ramulis 0·2-0·4 mm. latis, furcatis,
!axis, elongatis, apicibus filiformibus: sporis 6-7 X3-3·5µ, albis, levibus: basidiis
15-26 X5-7 µ, 4-sporig.: cystidiis nullis: hymenio ad 170µ incrassato: hyphis biformi
bus, skeletalibus 3·5-8(-10)µ latis, tunicis flavidulis 0·5-1·5µ crassis, alteris fibulatis:
ad humum, folia, et ramulos dejectos in silvis, Malaya.
P. vinacea sp. nov.

Ree. ad 7 X6 cm., ut in P. brunneola sed vinacea v. sordide purpurea, ramulis
flavido-incamatis, ramis saepe subverticillatis, ramulis furcatis, filiformibus: sporis
7-8 X3-3·s µ: ad humum in silvis, Malaya.
Pterulicium gen. nov.

Ut Pterula sed receptaculo resupinato Corticioideo e basi st1p1t1s evoluto: rec.
erecta ramosa ex hyphis biformibus instructa: rec. resupinata ex hyp his uniformibus
instructa, hyphis skeletalibus deficientibus: 1 sp., ad lignum emortuum, culmos et
. caules bambusarum palmarumque, Asia trop.
Pterulicium xylogenum (B. et Br.) comb. nov. = Pterula xylogena B. et Br.
Ramaria S. F. Gray emend. Donk

R. fiavo-alba sp. nov.
Receptacula 4-5 cm. alta, ramosa, solitaria v. gregaria, alba, dein subochracea,
apices versus flavidula: stipite 1-3 X0·8-2 cm.: ramis numerosis, congestis, rugulosis,
ramulis subulatis: came alba, fragili: sporis 8-10·5 X4·5-5µ, pallide ochraceis,
asperulosis v. subverruculosis, aguttatis: basidiis 50-70 X8-9µ: sterigmatibus 4,
4-5µ longi: hyphis 4-10 µ latis, fibulatis: ad terram in silvis, Singapore.
R. ignicolor sp. nov.

Receptacula ad 4 cm. alta, ramosa, ut in R. formosa, ignicoloria, ? subgelatinosa:
sporis 7-8·5 x 4-5µ, ellipsoideis, asperulosis v. subverruculosis, guttatis: hyphis 2-5µ
latis, vix v. non inflatis, fibulis deficientibus, tunicis subgelatinosis: ad terram in
pinetis, Italia (pr. Mendola, leg. Bresadola, det. Clavaria ignicolor, Bres., ? ined.).
R. luteo-fiaccida sp. nov.

Receptacula ad 4·5 cm. alta, ramosa, solitaria v. gregaria, pallide luteo-ochracea,
apicibus laete flavidulis: stipite 6-16 x 2-3 mm., gracili: ramis 2-3 mm. latis, ramulis
1-2 mm., bifidis, numerosis, subdivaricatis, apicibus subulatis: came alba, inodora:
sporis 5-6·5 x 3-3 ·5 µ, ochraceis, verruculosis: basidiis 45-55 X6-7 µ: sterigmatibus 4,
5-6 µ longis: hyphis 3-9(-18) µ latis, fibulatis: ad terram in silvis, Sumatra (pr.
Brastagi, c. 1,500 m. altitudine, Sept. 1931).
700 NOVITATES
var. americana var. nov.
Carne nee vinescenti nee nigrescenti, insipida: sporis angustioribus, 12·5-18·5 X
4·5-6µ: basidiis 4-sporigeris: ad humum in silvis frondosis, Am. bor.
R. polypus sp. nov.
Receptacula 2-8 cm. alta, ramosa, gracilia, gregaria v. caespitosa, albida dein
flavidula, subincarnata, ochracea v. alutacea, apicibus albis: stipite -12 X1-3·5 mm.,
e mycelio byssoideo cremeo-albo orienti: ramulis !axis, numerosis, arcuato-ascendenti
bus, plus minus applanatis, apicibus acutis: carne albida, floccoso-fibrosa, lenta,
inodora: sporis 6-8(-9) X4-5(-5·5)µ, cinnamomeo-ochracea, rugulosis, aguttatis:
basidiis 26-45 X8-9-µ, v. 50-60 X8-9,.,, in hymenio incrassato: sterigmatibus 4,
3·5-6µ longis: hymenio ad 250µ incrassato: hyp his 2-n(-13)µ latis, fibulatis, tunicis
0·5-1·5 µ crassis: ad lignum putridum et truncos dejectos palmarum, raro ad humum,
in silvis, Malaya, New Guinea.
var. concolor var. nov.
Pallide ochracea, subincarnata, v. alutacea, dein cinnamomea, apicibus concoloribus
(non laete flavis): sporis 6-10·5 X3·5-5µ, asperulosis v. sublevibus: Am. bor.,
Australia.
R. subgelatinosa sp. nov.
Receptacula ad 15 cm. alta, solitaria v. subcaespitosa, valde ramosa, robusta, ut in
R. formosa, roseo-armeniaca, apicibus luteis: stipite -4 X1 cm.: ramis 4-8 mm. latis,
ramulis 1-2 mm., numerosis: came subconcolori v. pallida, subgelatinosa, sicco
cartilaginea: sporis 8·5-10·5 X4·5-5µ, ellipsoideis, verruculosis: basidiis 50-70 X
8-·uµ: sterigmatibus 4, 5-7µ longis: hymenio non v. vix incrassato: hyphis 3-12µ
latis, fibulatis, tunicis subgelatinosis: ad humum in silvis, Malaya (Pahang).
R. Zippelii (Lev.) comb. nov.
Receptacula parva, gracilia, 2-8 cm. alta: sporis minoribus, 7-10 X4·5-6·5µ, aculeis
brevioribus 0·5-1µ longis: basidiis saepissime 3-4-sporigeris: ad humum in silvis,
Malaya.
Ramariopsis Donk, gen. emend.

Ut Clavulinopsis, sed differt sporis minute echinulatis, receptaculis plerumque
ramosis.
R. Kunzei var. deformis var. nov.
Ree. ad 6 cm. alta, brevia, compacta, ramis ramulisque paucis compressis v. clavatis,
obtusis, saepe deformia: Europa.
R. Kunzei var. Favreae var. nov.
Ut var. deformis, sed ramis valde compressis spathulato-flabelliformibus v. laminatis,
saepe curvatis, ramulis corniformibus dentatis cristatis: in pinetis, Gallia.
R. Kunzei v.ar. subasperata var. nov.
Ree. ad 2 cm. alta, pusilla, ramulis paucis: sporis !axe asperulatis; ad terram, Gallia.
R. tenuiramosa sp. nov.

Ree. ad 3·5 cm. alta, sordide flavida v. straminea, gracilia, stip1te vix tomentoso,
ramulis paucis elongatis subdivaricatis !axis: sporis 3·5-4·5 X3-3·5µ, subglobosis,
minute echinulatis: hymenio non spissato: hyphis ut in R. Kunzei: in pinetis, Anglia.
NOVITATES 701
Scytinopogon Singer
Scytinopogon angulisporus var. curtus var. nov.
Ree. ad 5·5 cm. alta, breviora, ramis ramulisque angustioribus: ad terram in silvis,
Malaya.
S. angulisporus var. parvus var. nov.
Ree. ad 9 cm. alta, graciliora, ramis ramulisque angustioribus: ad terram in silvis,
Malaya.
S. angulisporus var. gracilis var. nov.
Ree. ad ro cm. alta, stipite elongate gracili 2 mm. lato, erecto, non in solo immerse,
non flabellato-diviso, ramis ramulisque gracilibus, plerumque dichotomis, !axe et
saepe parce ramosa: ad terram in silvis, Malaya.
Typhula Fr.
Subgen. nov. Subcarnosae
Hyphae sclerotiorum medullares non agglutinatae, saepe crasse tunicatae et in
crustatae: hyphae receptaculorum solae in cortice stipitis agglutinatae: capitula ceracea.
Typus: T. sclerotioides (Pers.) Fr.
BAKER, GLADYS E. Studies in the genus Physalacria. Bull. Torr. Bot. Cl. 68, 1941,
265-88.
BAUCH, R. Untersuchungen uber zweisporige Hymenomyceten II. Kern-degenera
tion bei einigen Clavaria-arten. Arch. f. Protistenk. 58, 1927, 285-99.
VAN DER BIJL, P. A.. Oor enige Suid-Afrikaanse Clavaria- soorte of Knotsswamme.
South Afr. Journ. Sci. 29, 1932, 317-23.
BISBY, G. R., BULLER, A. H. R., DEARNESS, J. Fungi of Manitoba, 1929.
BouRDOT, H., and GALZIN, A. Hymenomycetes de France 1, 1928.
BRESADOLA, J. lconographia Mycologica, 1927-33.
BULLER, A. H. R. Spore-discharge in the Clavarieae. Res. Fungi 2, 1922, 179.
-- The Diploid Cell and the Diploidisation Process in Plants and Animals, with
special reference to the Higher Fungi. Bot. Rev. 7, 1941, 335-431.
BURT, E. A. The Thelephoraceae of North America 1. Ann. Mo. Bot. Gdn. 1, 1914,
185-228.
--The North American species of Clavaria. Ibid. 9, 1922, 1-78.
CLELAND, J. B., and CHEEL, E. Records of Australian Fungi 1. Proc. Linn. Soc.
N.S. W. 41, 1916, 866-70.
CLELAND, J. B. Toadstools and Mushrooms and other Larger Fungi of South
Australia, Pt. 2, 1935.
COKER, W. C. The Clavarias of the United States and Canada, 1923.
-- Further Notes on Clavarias. Journ. El. Mitch. Sci. Soc. 63, 1947, 43-67, pl. 1-14.
Coorrn, M. C. Handbook ofAustralian Fungi, 1892.
CORNER, E. J. H. Variation in the size and shape of spores, basidia and. cystidia in
Basidiomycetes. New Phyt. 46, 1947, 10,.
-- Studies in the basidium. 1. The Ampoule effect. Ibid. 47, 1948, 22.
COTTON, A. D. Notes on British Clavariae. Tr. Brit. Myc. Soc. 1906-11.
-- and WAKEFIELD, E. M. A Revision of the British Clavariae. Ibid. 6, 1919,
164-98.
DoNK, M.A. Revision der Niederlandischen Homobasidiomycetae-Aphyllophoraceae
II. Med. Bot. Mus. Herb. R. Univ. Utrecht, No. 9, 1933.
DOTY, M. S. Clavaria, the Species Known from Oregon and the Pacific North West.
Oreg. St. Col. 1944.
--A preliminary key to the genera of Clavarioid fungi. Bull. Chic. Ac. Sci. 8, 1948,
173-8.
FAWCETT, STELLA G. M. Studies on Australian Clavariaceae, Pt. I, Proc. Roy. Soc.
Viet. 51, 1938, 1-20: Pt. 11, ibid. 1939, 265-80: Pt. III, ibid. 52, 1940, 153-63.
FRIES, E. Systema Mycologicum 1, 1821: Elenchus Fungorum 1828: Epicrisis
Systematis Mycologici 1838 (Clavariaceae), Hymenomycetes Europaei sive
Epicriseos Systematis Mycologici, 2nd ed., 1874.
GILBERT, E. J. La Spore des Champignons superieurs. Paris, 1927.
HARPER, E. T. The Clavaria fistulosa group. Mycol. 10, 1918, 53-7.
HEIM, R. Fungi lberici: Publ. de la Junta d. Ciencies Naturales d. Barcelona 1S, 1934,
44-52 (Clavariaceae).
HEMMI, T., and KuRATA, S. Miscellaneous notes on Japanese Fungi. Acta Phytotax.
Geobot. 2, 1933, 109.
IMAI, SANSHI. On the Clavariaceae of Ja pan, I, Tr. Sapporo Nat. Hist. Soc. 11, 1929,
38-44: II, ibid. 1930, 70-7: III, ibid. 12, 1931, 9-29: IV, ibid. 13, 1934, 385-7.
ITO, S., and IMAI, S. Fungi of Bonin Islands. Ibid. 15, 1937, 53.
JUEL, D. Cytologische Pilzstudien I: d. Basidien d. Gattung Cantharellus, Craterellus
u. Clavaria. Nov. Act. Reg. Soc. Scie11t. Upsal. 4, 1916, 3-34.
KARSTEN, P.A. Hattsvampar Rysslands, Finlands o. d. Skandinavska Halfons, Bijdr.
t. Kann., Finl. Nat. o. Folk 37, 1882, Finlands Basidsv., ibid. 48, 1889, and 50, I,891,
KAUFFMAN, C. H. The Mycological Flora of the Higher Rockies of Colorado. Pap.
Mich. Ac. Sci. Arts Lett. 1, 1921, 101-50. The fungous flora of Mt. Hood, with
some new species, ibid. 5, 1925, u5-48. Cystidia in the genus Clavaria and some
undescribed species, ibid. 8, 1927, 141-5I. A study of the fungous flora of Lake
Superior region of Mich., with some new species, ibid. 9, 1929, 169-218. The fun
gous flora of the Siskiyou Mountains in Southern Oregon, ibid. II, 1930, 151-209.
KILLERMANN, S. Die Natilrlichen Pflanzenfamilien, 2nd ed., 6, 1928.
KONRAD, P., and MAUBLANC, A. !cones Selectae Fungorum, 1935-7.
KRIEGER, L. C. C. The Mushroom Handbook, N.Y. 1936.
LINDER, D. H. The genus Myxomycidium. Mycol. 26, 1934, 332-43.
LUNDELL, S., and NANNFELDT, J. A. Svenska Svampar . . . 1935-47.
MAIRE, R. Recherches cytologiques et taxonomiques sur les Basidiomycetes, .Annexe
du Bull. Soc. Myc. Fr. 18, 1902, 1-210. Fungi Catalaunici, Publ. de la Junta
d. Ciencies Naturales de Barcelona 15, 1933, 28-31 (Clavariaceae): ibid. 3, 1937,
30-3 (Clavariaceae).
MARTIN, G. W. Some tropical American 'clavarias'. Lilloa 5, 1940, 191-6.
MASSEE, G. British Fungus Flora I, 1892.
-- The Fungus Flora of New Zealand, Pt. II. Trans. N. Zeal. Inst. 39, 1906, I.
MOELLER, E. H. Fungi of the Faeroes, Pt. I, 1945 (Copenhagen).
VAN OVEREEM, C. Bull. Jard. Bot. Buit. Ser. 3, 5, 1923.
-- and WEESE, J. !cones Fungorum Malayensium I-IV, 1923 (Vienna).
OVERHOLTS, L. 0., in Mycological Explorations of Venezuela. Monogr. Univ.,Puerto
Rico, Ser. B., No. 2, 1934.
PATOUILLARD, N. Tabulae Analyticae Fungorum 1883-9. Les Hymenomycetes
d'Europe, 1887: Essai Taxonomique sur les families et les genres des Hymeno
mycetes, 1900.
PERSOON, C. H. Mycologia Europaea I, 1822.
PETCH, T. Notes on Ceylon Clavarieae. Ann. R. Bot. Gdn. Perad. 9, 1925, 329.
QuELET, L. Flore Mycologique de la France, 1888. Les Champignons du Jura et des
Vosges, 1869-75.
REA, C. British Basidiomycetae, 1922 (Cambridge).
REMSBERG, RUTH E. Studies in the genus Typhula. Mycol. 32, 1940, 52-96.
RENOUF, H. Recherches botaniques et chimiques sur les Clavaires rameuses. These
1923, 1-133 (Strasbourg, M. Colin) [not seen].
ROGERS, D. P. The basidium. Univ. of Iowa, Stud. Nat. Hist. 16. 1934, 160-82.
Index Botanicus sistens omnes fungorum species in C. H. Persoonii Mycologia
Europaea, 1942 (Cambridge, Mass.).
SACCARDO, P. Sylloge Fungorum VI-XXIII, 1888-1925.
SCHROETER, J. Die Pilze Schlesiens, in Cohn's Kryptogamen-Flora von Schlesien 3,
1889, 442-3.
SPEGAZZINI, C. Fungi Argentini. An. Mus. Nac. Buenos Aires, 1880-99.
TENG, S. C. Fungi of Nanking. Contrib. Biol. Lab. Sci. Soc. China, Bot. 7, 1932, 105.
Fungi of Chekiang. Ibid. 8, 1932, 62. Additional Fungi from China. Sinensia 7,
1936, 212.
TEODORO, N. G. An enumeration of Philippine Fungi. Dept. Agr. Comm. Manila,
Techn. Bull. 4, 1937.
VANG, J. Typhula Species on Agricultural Plants in Denmark. Medd. Plantepat.
Afd. Kgl. Vet. Landb., Copenhagen, 28, 1945, 1-45.
WAKEFIELD, E. M. Fungi from New Caledonia. Journ. Linn. Soc. Bot. 46, 1922, 88.
INDEX TO GENERA, SPECIES, AND SYNONYMS
(Synonyms in italics)
Aleurodiscus helveolus Bres. (1925) = Clavariadelphus fistulosus var. contortus
I
Aphelaria Comer
A. aurantiaca (P. Henn.) Comer
A. brunneola (B. et C.) Comer
A. deflectens (Bres.) Comer
A. dendroides (Jungh.) Corner
A. flabellata (Wakef.) Comer
A. incarnata Corner
A. pusio (B.) Corner
A. spiculosa Corner
A. tasmanica (Lloyd) Comer
A. trachodes (B.) Corner
A. tropica (Mont.) Corner
A. tuberosa (Grev.) Corner
Araeocoryne Corner
A. elegans Corner
Baumanniella P. Henn. = Physalacria Pk.
B. brasiliensis Rick = Phys"alacria brasiliensis (Rick) Corner
B. togoensis P. Henn. = Physalacria togoensis (P. Henn.) Corner
Calocera albipes (Mont.) B. et Br., see Lentaria mucida (Fr.) Comer
C. dilatata Mont. = Pterula dilatata (Mont.) 'Corner
C. tuberosa Fr. = Clavariadelphus fistulosus (Fr.) Corner
Caripia 0.K.
C. Montagnei (B.) O.K.
Ceratella Pat. = Ceratellopsis Konr. et Maubl.
C. aculeata Pat. = Ceratellopsis aculeata (Pat.) Corner
C. aculina Pat. = Pterula gracilis (B. et Desm.) Corner
C. acuminata (Fckl.) Pat. = Ceratellopsis acuminata (Fckl.) Corner
C. Ferryi Quel. = Pterula gracilis (B. et Desm.) Corner
C. Helenae Pat. = Ceratellopsis Helenae (Pat.) Corner
C. macrospora Pat. = Pterula macrospora (Pat.) Corner
C. Queletii Pat. = Ceratellopsis Queletii (Pat.) Konr. et Maubl.
C. uncialis (Grev.) = Pistillaria uncialis Grev.
Ceratellopsis Konr. et Maubl.
C. aciculata (Dur. et Lev.) Corner
C. aculeata (Pat.) Corner
C. acuminata (Fckl.) Corner
C. asphodeli Corner
C. Brondaei (Quel.) Corner
C. caespitulosa (Sacc.) Corner
C. Carestiae (Sacc.) Corner
C. dryopteridis (Imai) Comer
C. equiseticola (Boud.) Corner
C. graminicola (Bourd. et Galz.) Corner
C. Helenae (Pat.) Corner
C. mucedinacea (Boud.) Corner
C. mucosa (B. et C.) Corner
INDEX TO GENERA, SPECIES, AND SYNONYMS 705
C. Queletii (Pat.) Konr. et Maubl.
C: Rickii (Oud.) Corner
C. rosella (Fr.) Corner
var. ramosa Pat.
C. sagittaeformis (Pat.) Corner
C. Sydowii (Bres.) Corner
C. thujicola (Kauffm.) Corner
C. tremula (SaC£:.) Corner
Chaetotyphula Corner
C. actiniceps (Petch) Corner
C. hyalina (Jungh.) Corner
var. microcystis Corner
Cladaria Ritg. = Ramaria S. F. Gray
Clavaria Fr.
C. abietina Fr. = Ramaria Invalii (Cott. et Wakef.) Donk
subsp. cyanescens Rom. = Ramaria ochraceo-virens (Jungh.) Donk
f. Persoonii Fr. = Ramaria ochraceo-virens (Jungh.) Donk
C. abietina Pers. = Ramaria ochraceo-virens (Jungh.) Donk
sensu Coker 'small virescent form' = Ramaria ochraceo-virens (Jungh.) Donk
sensu Coker 'small non-virescent form' = Ramaria flaccida (Fr.) Ricken
C. acris Pk., see Ramaria apiculata (Fr.) Donk
C. aculeata Blonski
C. aculina Quel. = Pterula gracilis (B. et Desm.) Corner
C. acuta Fr.
C. acutissima B. = Ramaria acutissima (B.) Corner
C. acutissima Mont. (ined.) = Aphelaria tropica (Mont.) Corner
C. adustipes Speg. = Pterula adustipes (Speg.) Corner
C. aeruginosa Pat. = Ramaria cyanocephala (B. et C.) Corner
C. affinis Pat. et Doass.
C. affiata Lagger. = Lentaria affi.ata (Lagger.) Corner
C. alba sensu Bourd. et Galz. = Clavulina gallica Corner
C. alba Lloyd = Lentaria mucida (Fr.) Corner
C. alba Pers. = Clavaria vermicularis Fr.
C. alba Pers. = Clavulina cristata var. coralloides Corner
C. albida Pk., see Ramaria obtusissima (Pk.) Corner
C. albida Sacc. = Clavulina cristata var. nivea Bourd. et Galz.
C. albidula Lasch = nomen nudum
C. albipes Mont., see Lentaria mucida (Fr.) Corner
C. alcicornis Lev. = Clavulina Leveillei (Sacc.) v. Ov.
C. alcicornis Zoll. et Mor. = Clavulinopsis alcicornis (Zoll. et Mor.) Corner
C. alliacea Corner
C. alnea Schulz = Clavariadelphus fistulosus (Fr.) Corner
C. alpina Saut. = ? Clavulina cristata var. Jappa Karst.
C. alutacea Lasch = Ramaria gracilis (Fr.) Quel.
C. amethystea Pers. = Clavulina amethystina (Fr.) Donk
C. amethystina Fr. = Clavulina amethystina (Fr.) Donk
var. purpurea Bourd. et Galz. = Clavulina amethystina var. lilacina Quel.
C. amethystina sensu Cott. et vVakef. = Clavaria Zollingeri Lev.
C. amethystina Zoll. = Clavaria Zollingeri Lev.
C. amethystinoides Pk. = Clavulina amethystinoides (Pk.) Corner
C. amoena Zoll. et Mor. = Clavulinopsis amoena (Zoll. et Mor.) Corner
C. angulispora Pat. = Scytinopogon angulisporus (Pat.) Corner
C. angustata Fr. = Clavulinopsis helvola (Fr.) Corner
C. anomala Fr.
5119 zz
706 INDEX TO GENERA, SPECIES, AND SYNONYMS
C. apiahyana Speg. = Ramaria apiahyana (Speg.) Corner
C. apiculata Fr.= Ramaria apiculata (Fr.) Donk
C. appalachiensis Coker= Clavulinopsis appalachiensi& (Coker) Corner
C. arborea Atk., see Ramariopsis Kunzei (Fr.) Donk
C. arborescens B. = Clavaria Zollingeri Lev.
C. Archeri B. = Clavulinopsis Archeri (B.) Corner
C. arctata Britz. = Clavulina rugosa var. alcyonaria Corner
C. ardenia Fr. = Clavariadelphus fistulosus (Fr.) Corner
C. argillacea Fr.
f. citrina Bourd. et Galz.
var. brevispora Corner
var. dispar Pers.
var. gracillima Corner
var. obtusata Favre
var. sphagnicola Corner
C. argillosa Britz., see Clavaria argillacea Fr.
C. asperula Atk.= Ramariopsis Kunzei (Fr.) Donk
C. asperulans Atk.= Ramariopsis Kunzei (Fr.) Donk
C. asperulospora Atk. = Ramariopsis asperulospora (Atk.) Corner
C. asterella Atk.= Ramariopsis asterella (Atk.) Corner
C. asterospora Pat. = Clavulinopsis asterospora (Pat.) Corner
C. atrobadia Corner
C. atroumbrina Corner
C. aurantia Cke. et Mass. = Clavulinopsis Archeri (B.) Corner
C. aurantia sensu Clel. = Clavulinopsis amoena (Zoll. et Mor.) Corner
C. aurantia Pers. ex Karst.= Clavulinopsis luteo-alba var. latispora Corner
C. aurantiaca Fr.= Clavulinopsis helvola (Fr.) Corner
C. aurantio-cinnabarina Schw.= Clavulinopsis aurantio-cinnabarina (Schw.) Corner
C. aurea Fr. = Ramaria aurea (Fr.) Quel.
C. austera Britz., see Clavulinopsis corniculata (Fr.) Corner f. simplex Donk
C. australiana Clel. = Ramaria holorubella (Atk.) Corner
C. avellaneo-nigrescens Imai
C. Baileyi Mass.= Scytinopogon angulisporus (Pat.) Corner
C. Barlae Bres.
C. Bataillei (Maire) Sacc. = Ramaria Bataillei (Maire) Corner
C. Berkleyi Mont.
C. Bessonii Pat.= Clavulina Bessonii (Pat.) Corner
C. bicolor Mass.= Clavaria Zollingeri Lev.
C. bicolor Pk. = Ramariopsis vestitipes (Pk.) Corner
C. bifida Chev., see Clavulinopsis fusiformis (Fr.) Corner
C. biformis Atk.= Clavulinopsis biformis (Atk.) Corner
C. bifurca S. F. Gray = Clavulinopsis fusiformis var. ceranoides W. G. Smith
C. Bizzozeriana Sacc. = Ramariopsis pulchella var. Bizzozeriana Bourd. et Galz.
C. boninensis Ito et Imai= Clavulinopsis boninensis (Ito et Imai) Corner
C. botrytis Fr. = Ramaria botrytis (Fr.) Ricken
sensu Coker= Ramaria botrytoides (Pk.) Corner
sensu Kauffm. = Ramaria holorubella (Atk.) Corner
C. botrytoides Pk. = Ramaria botrytoides (Pk.) Corner
C. Bourdotii Bres.= Hydnaceae (see p. 265)
C. brachiata Fr. = Clavariadelphus fistulpsus var. contortus Corner
C. Braunii P. Henn., see Lachnocladium aurantiacum B. et Br.
C. Bresadolae Cav.= C. Cavarae Sacc. et Trott. = Ramariopsis pulchella (Boud.)
Corner
C. Bresadolae Quel. = Hydnaceae (seep. 265)
C. Brondaei Quel.= Ceratellopsis Brondaei (Quel.) Comer
C. Broomei Cott. et Wakef.= Ramaria nigrescens (Brinkm.) Donk
C. Brunaudii Quel.= Pterula gracilis (B. et Desm.) Comer
C. brunnea Zeller= Ramaria brunnea (Zeller) Corner
C. brunneola B.= Aphelaria brunneola (B.) Comer
C. bulbosa Pers.= ? Clavariadelphus fistulosus var. contortus Corner
C. bulbosa Wallr., see Clavulinopsis pulchra f. coccineo-basalis Josser.
C. byssiseda auctt.= Lentaria byssiseda Comer
C. byssiseda Fr.= C. himantia (Schw.) Fr. (seep. 265)
C. byssiseda sensu Pat.= Ramaria gracilis (Fr.) Quel.
C. cacao Coker= Ramaria cacao (Coker) Cotner
C. caepicolorosa Fawcett= Clavulinopsis miniata (B.) Corner
C. calocera Martin= Clavulinopsis calocera (Martin) Corner
C. caloceriformis Oudem.= Clavulinopsis corniculata (Fr.) Comer
C. canaliculata Fr.= Clavulina rugosa var. canaliculata Corner
C. candelabrum Mass.= Clavicorona candelabrum (Mass.) Corner
C. candida Weinm.= Clavulinopsis candida (Weinm.) Comer
C. candidula Karst.= Pistillaria setipes Grev.
C. capitata Lloyd= Ramaria capitata (Lloyd) Corner
C. capucina Pat.= Ramaria capucina (Pat.) Corner
C. carbonaria Mont.= Ramaria acutissima (B.) Comer
C. cardinalis Boud. et Pat.= Clavulinopsis miniata var. sanguinea Corner
C. carnea Wallr., see Pterula mulfida Fr.
C. cartilaginea B. et C.= Clavulina cartilaginea (B. et C.) Comer
C. Cavarae Sacc. et Trott.= Ramarioosis pulchella (Boud.) Corner
C. cedretorum (Maire) Sacc., see Ramaria fumigata (Pk.) Corner
C. ceranoides Fr.= Clavulinopsis fusiformis (Fr.) Corner
C. cervicornis A. L. Smith= Ramaria cervicornis (A. L. Smith) Corner
C. cervina B. et C., see Ramariopsis Kunzei (Fr.) Donk
C. cervina W. G. Smith= Ramaria filicina (Sacc. et Syd.) Corner
C. chionea Pers.= Ramariopsis Kunzei (Fr.) Donk
C. chlorina B. et Br.= fungus imperfectus (fide Petch, Ann. R. B. Gard. Per. 9, 1924,
145).
C. chondroides B.= Clavulina chondroides (B.) Corner
C. chordostyla Pers.= Pistillaria Todei (Fr.) Corner
C. cinerea Fr.= Clavulina cinerea (Fr.) Schroet.
C. cinereo-atra Rick
C. cineroides Atk.= Clavulinopsis cineroides (Atlc.) Corner
C. cinnamomea Fawcett= Clavulinopsis cinnamomea (Fawcett) Corner
C. circinans Pk.= Ramaria suecica (Fr,) Donk
C. cirrhata B.= Clavulina cirrhata (B.) Corner
. C. citriceps Atk.
C. citrina sensu Hemmi et Kurata, see Clavulinopsis helvola Corner
C. citrina Quel.= Clavaria argillacea f. citrina Bourd. et Galz.
C. citrino-alba Moller= Clavulinopsis citrino-alba (Moeller) Corner
C. cladonia Speg.= C. Spegazzinii Sacc., see Scytinopogon angulisporus (Pat.) Corner
C. cladoniae Kalchbr.
C. clara B. et C., see Clavulinopsis amoena (Zoll. et Mor.) Corner
C. clavaeformis Britz.= Clavulina rugosa {Fr.) Schroet.
C. clavata Pk.= Clavulinopsis vernalis (Schw.) Corner
C. clavuligera Heim.= Ramariopsis clavuligera (Heim.) Corner
C. cochleariformis Pers.= ? Clavulinopsis fusiformis Fr.
C. coerulescens Karst., see Clavulina amethystina (Fr.) Donk
C. Colensoi B.= Clavicorona Colensoi (B.) Corner
C. coliformis Boud. = Clavulinopsis coliformis (Boud.) Comer
C. comosa Pat., see Clavulina cristata var. subcinerea Donk
C. complanata Clel. = Clavulina complanata Comer
C. complanata de Bary= Typhula gyrans Fr.
C. compressa B., see Lentaria surculus (B.) Corner
C. compressa Schrot.
C. compressa Schw.= Clavulinopsis fusiformis (Fr.) Corner
C. conchyliata Allen= Ramariopsis puchella var. Bizzozeriana Bourd. et Galz.
C. condensata sensu Bourd. et Galz. = Ramaria stricta (Fr.) Quel.
C. condensata Fr.= Ramaria condensata (Fr.) Quel.
C. condensata var. violaceo-tincta Bourd. et Galz.= Ramaria stricta var. violaceo-
tincta (Bourd. et Galz.) Corner
C. conjuncta Pk.= Ramaria conjuncta (Pk.) Corner
C. conjunctipes Pk.= Ramaria conjunctipes (Pk.) Corner
C. connata B.= Clavulina connata (B.) Corner
C. constans Coker= Clavulinopsis constans (Coker) Corner
C. contorta Fr.= Clavariadelphus fistulosus var. contortus Corner
C. corallino-rosacea Clel.= Clavulinopsis corallino-rosacea (Clel.)
C. coralloides Fr.= Clavulina cristata var. coralloides Corner
C. Corbierei Bourd. et Galz.
C. coriaria Pers.
C. cornea Fr.= Calocera cornea Fr.
C. corniculata Fr. = Clavulinopsis corniculata (Fr.) Corner
C. coronata Schw.= Clavicorona pyxidata (Fr.) Doty
C. coronata Zipp.= Clavicorona javanica (Sacc. et Syd.) Corner
C. coronilla Martin = Lentaria coronilla (Martin) Corner
C. corrugata Karst.= Ramaria flaccida (Fr.) Ricken
C. corticola Quel. = Lentaria corticola (Quel.) Corner
C. corynoides Pk.= Clavaria vermicularis Fr.
C. crassa Britz., see Clavulina cinerea f. sublilascens Bourd. et Galz.
C. crassipes Pk.= Ramaria secunda (B.) Corner
C. cretacea Coker
C. crispa Pers.= Sparassis crispa Fr.
C. crispula Fr. = Ramaria crispula (Fr.) Quel.
C. crispula subsp. soluta Karst. = Ramaria soluta (Karst.) Corner
C. cristata Fr.= Clavulina cristata (Fr.) Schroet.
var. fuligineo-cinerascens Bourd. et Galz., see Clavulina cristata var. bicolor Donk
C. cristata J ungh.= Thelephora cristata Fr.= Sebacina incrustans (Pers.) Tul.
C. cristatula P. Henn. et Nym.= Clavulina Leveillei (Sacc.) v. Ov.
C. crocea Fr. = Ramariopsis crocea (Fr.) Corner
C. Crosslandii Cott.
C. culmigena (Fr.) Karst. = Pistillaria culmigena Fr.
C. curta Fr. = Ramariopsis curta (Fr.) Corner
C. cyanescens (Rom.) Lundell = Ramaria ochraceo-virens (Jungh.) Donk
C. cyanocephala B. et C.= Ramaria cyanocephala (B. et C.) Corner
C. Cyatheae P. Henn.= Ramaria cyatheae (P. Henn.) Corner
C. cylindrica Chev.= Clavulinopsis helvola (Fr.) Corner
C. cylindrica S. F. Gray= Clavaria vermicularis Fr.
C. cystidiophora Kauffm.= Ramaria cystidiophora (Kauffm.) Corner
C. Daigremontiana Boud.= Clavulinopsis Daigremontiana (Boud.) Corner
C. Daulnoyae Quel.
C. dealbata B., see Ramariopsis Kunzei (Fr.) Donk
C. decolor B. et Br.
C. decolorans (Karst.) Sacc. = Ramaria decolorans (Karst.) Corner
INDEX TO GENERA, SPECIES, AND SYNONYMS 7 09
c;. decurrens Pers.= Ramaria crispula (Fr.) Quel.
sensu Coker= Ram.aria pusilla (Pk.) Comer
C. delicata Fr.= Lentaria delicata (Fr.) Comer
C. delicia B.= Clavulina delicia (B.) Comer
C. dendroidea sensu Bourd. et Galz., et Bres.= Ramaria stricta (Fr.) Quel.
var. compacta Bourd. et Galz.= Ramaria apiculata var. compacta (Bom:d. et Galz.)
Corner
C. dendroidea Fr.= Ramaria condensata (Fr.) Quel.
C. dendroides Jungh.= Aphelaria dendroides (Jungh.) Comer
C. densa Pk.= Ramaria formosa (Fr.) Quel.
C. densissima Pk.= Ramaria stricta var. concolor Corner
C. depokensis v. Ov.= Clavulinopsis depokensis (v. Ov.) Corner
C. diaphana (Fr.) Karst.= Pistillaria diaphana Fr.
C. dichotoma God.= Clavulinopsis dichotoma (God.) Corner
C. dichotoma Kalchbr.= Ramaria Saccardoi (Syd.) Comer
C. digitata Pers.= Xylaria digitata Grev.
C. dilatata Mont.= Pterula dilatata (Mont.) Comer
C. dissipabilis Britz.= Clavulinopsis helvola (Fr.) Corner
C. distincta Britz.= Clavulinopsis corniculata f. simplex Donk
C. divaricata Karst.= Ramaria Karstenii (Sacc. et Syd.) Corner
C. divaricata Pk.= Ramaria divaricata (Pk.) Corner
C. Dozyi Lev.= Ramaria Zipelii (Lev.) Corner
C. ? dubiosa Speg.= ? Pyrenomycete, see p. 266
C. durbana van d. Bijl= Ramaria durbana (van d. Bijl) Comer
C. eburnea Pers.= Clavaria vermicularis Fr.
C. echinospora B. et Br.= Scytinopogon echinosporus (B. et Br.) Corner
C. echinospora Boud. et Pat.= Clavulinopsis helvola (Fr.) Comer
C. echinospora P. Henn.= Ramaria fragillima (Sacc. et Syd.) Comer
C. elegantula Karst. = Pistillaria elegantula (Karst.) Corner
C. elongata Britz.= Ramariopsis Kunzei (Fr.) Donk
C. epichnoa Fr.= Lentaria epichnoa (Fr.) Corner
C. epiphylla Quel.= Pistillaria epiphylla (Quel.) Corner
C. ericetorum Pers.= Clavaria argillacea Fr.
C. erythropus Pers.= Typhula erythropus Fr.
C. eumorpha (Karst.) Sacc.= Ramaria eumorpha (Karst.) Corner
C. exigua Pk.= Ramariopsis pulchella (Boud.) Corner
C. exilis Pers.
C. extensa Herpell= ? Ramaria Mairei Donk
C. falcata Fr.= Clavaria acuta Fr.
var. citronipes Quel., see Clavaria acuta Fr.
var. tenuissima Weinm., see Clavaria acuta Fr.
C. falcatula P. Henn.= Pterula falcatula (P. Henn.) Corner
C. fallax Pers.= Clavulina cristata (Fr.) Schroet.
C. fasciculata Pers.= Clavulinopsis fusiformis (Fr.) Corner
C. fastigiata Fr. = Clavulinopsis corniculata (Fr.) Corner
C. Favreae (Quel.) Sacc. et Ti:ott. = Ramariopsis Kunzei var. Favreae Corner
C. fellea Pk.= Clavulinopsis corniculata (Fr.) Corner
C. Jennica Karst. (1882) = Ramana fennica (Karst.) Ricken
C. fennica Karst. (1891) = Ramaria decolorans (Karst.) Corner
C. fennica sensu Coker= Ramaria fumigata (Pk.) Corner
C. filaris Kalchbr.= Pterula filaris (Kalchbr.) Corner
C. filata Pers., see Typhula incarnata Fr.
C. filicicola Fawcett = Ramaria filicicola (Fawcett) Corner
C. filicina Pers.= Pistillaria Todei (Fr.) Corner
C. filicina Sacc. et Syd.= Ramaria filicina (Sacc. et Syd.) Corner
C. filiformis (Fr.) Karst.= Clavariadelphus junceus (Fr.) Corner
C. filiformis P. Henn. et Nym.= Clavulinopsis spiralis (Jungh.) Corner
C. filiola Corner
C. filipes B. et Rav.= Clavaria acuta Fr.
C. filipes sensu Coker= Clavulinopsis filipes Corner
C. fimbriata Quel.
C. fistulosa Fr.= Clavariadelphus fistulosus (Fr.) Corner
C. flabellata Wakef.= Aphelaria flabellata (Wakef.) Corner
C. flaccid.a Fr.= Ramaria flaccida (Fr.) Ricken
var. Invalii Konr. et Maubl.= Ramaria lnvalii (Cott. et Wakef.) Donk
C. flagelliformis B.= Aphelaria dendroides (Jungh.) Corner
C. flammans B.= Clavulinopsis helvola (Fr.) Corner
C. flava Fr.= Ramaria flava (Fr.) Quel.
C. flava sensu Coker= Ramaria flavo-brunnescens (Atk.) Corne!'
var. subtilis Coker= Ramaria divaricata (Pk.) Corner
C. flavella B. et C.= Clavulinopsis flavella (B. et C.) Corner
C. flavipes Fr.= Clavaria argillacea Fr.
C. flavo-brunnescens Atk.= Ramaria flavo-brunnescens (Atk.) Corner
C. flavula Atk.= Ramaria suecica (Fr.) Donk
C. flavuloides Burt= Ramaria gracilis (Fr.) Quel.
C. Fleisch�iana P. Henn.= Clavulinopsis Fleischeriana (P. Henn.) Corner
C. flexuosa Lasch= nomen nudum
C. floridana Singer= Clavulina floridana (Singer) Corner
C. foetid<t Atk.= Clavaria fuscata Oudem.
C. formosa Fr.= Ramaria formosa (Fr.) Quel.
C. formosula Britz.= Ramaria formosa (Fr.) Quel.
C. fossicola Corner
C. fragilis Fr. = C. vermicularis Fr.
C. fragillima Sacc. et Syd.= Ramaria fragillima (Sacc. et Syd.) Corner
C. Jruticum Karst.= Pistillariafruticum Karst. see P. pusilla Fr.
C. fuegiana S_\)eg.= Pistillaria fuegiana (Speg.) Corner
C. fuliginea Pers.= Clavulina cinerea (Fr.) Schroet.
C. fumigata Pk.= Ramaria fumigata (Pk.) Corner
C. fumosa Fr.
C. fumosoides Kauffm.= Clavaria purpurea Fr.
C. furcata P. Henn. et Nym.= Clavulinopsis umbrina (Lev.) Corner
C. Jurcellata Fr.= Lachnocladium furcellatum (Fr.) Lev.
C. fuscata Oudem.
C. fuscescens Fr.
C:/uscipes Pers.= Pistillaria fuscipes (Pers.) Corner
C. Jusco-lilacina B.= ? Pyrenomycete
C. fuscp-lilacina sensu v. Ov. = Clavulina Leveillei (Sacc.) v. Ov.
C. fusiformis Fr.= Clavulinopsis fusiformis (Fr.) Corner
var. /Intillarum Pat., see Clavulinopsis miniata var. sanguinea Corner
var. ceranoides W. G. Smith
var. congoensis Beeli
C. gelatinosa Coker= Ramaria gelatinosa (Coker) Corner
C. geniculata (Lev.) Sacc., see Scytinopogon angulisporus (Pat.) Corner
G. geoglossoides Boud. et Pat.= Clavulinopsis helvola var. geoglossoides Corner
C. Gibbsiae Ramsb.
- var. megaspora Corner
var. tenuis Corner
f. microspora Corner
C. gigantea Schw. = Acurtis gigantea (Schw.) Fr. (? an abortive agaric).
C. gigantula Britz. = Clavulina rugosa (Fr.) Schri:it.
C. gigaspora Cott. et Wakef. = Aphelaria tuberosa (Gr_ev.) Corner
C. globospora Kauffm. = Clavulinopsis appalachiensis (Coker) Corner
C. Gollani P. Henn.
C. Gordius Speg. = Pterula Gordius (Speg.) Corner
C. gracilior Britz. = Clavaria vermicularis Fr.
C. gracilis (B.) Karst. = Pterula gracilis (B. et Desm.) Corner
sensu Karst. = Pistillaria anceps (Karst.) Corner
C. gracilis Fr. = Ramaria gracilis (Fr.) Quel.
C. gracilis (Sow.) auctt. = Clavaria vermicularis var. gracilis Bour<l. et Galz.
C. gracillima Pk. = Clavaria argillacea var. gracillima Corner
C. gracillima Wakker = C. Wakkeri Sacc. et Syd., see Pterula Oryzae
C. grandis Pk. = Ramaria grandis (Pk.) Corner
C. granulata Pers. = ? Pistillaria granulata Pat.
C. gregalis Britz. = Clavulina rugosa var. alcyonaria Corner
C. Greletii Boud.
C. grisea Fr. = Clavulina cinerea (Fr.) Schroet.
f. petricola Bourd. et Galz. = Clavulina cinerea (Fr.) Schroet.
C. griseola Rea = Clavulinopsis griseola (Rea) Corner
C. grossa auctt., see Clavulina cristata var. subrugosa Corner, and Clavulina rugosa
var. alcyonaria Corner
C. guarapiensis Speg., see Scytinopogon angulisporus (Pat.) Corner
C. Guilleminii Bourd. et Galz.
C. gyrans Pers. = Typhula gyrans Fr.
C. helicoides Pat.
C. helveola sensu Coker = Clavulinopsis luteo-alba (Rea) Corner
C. helvola Fr. = Clavulinopsis helvola (Fr.) Corner
var. dispar Pers. = Clavulinopsis corniculata (Fr.) Corner
C. Henriquesii Bres. et Roum. = Ramaria Henriquesii (Bres. et Roum.) Comer
C. herculeana S. F. Gray = Clavariadelphus pistillaris (Fr.) Donk
C. Herveyi Pk. = Clavulina rugosa (Fr.) Schroet.
C. himantia (Schw.) Fr. = Hydnaceae (see p. 265)
C. hirta Pers. = Clavariadelphus junceus (Fr.) Corner
C. Holmskjoldii Oud. = Clavulinopsis Holmskjoldii (Oud.) Corner
C. holorubella Atk. = Ramaria holorubella (Atk.) Corner
C. Holsatica (P. Henn.) Sacc. = Ramaria holsatica (P. Henn.) Corner
C. humilis Cke. = Clavulina humilis (Cke.) Corner
C. ignea Pers.
C. ignicolor Bres., see Ramaria ignicolor Bres.
C. implexa Lev. = Lachnocladium implexum (Lev.) P. Henn.
sensu van Ov. = Scytinopogon echinosporus (B. et Br.) Corner
C. inaequalis Fr. nomen ambiguum (see p. 343).
var. aurantia Bres. = Clavulinopsis pulchra (Pk.) Corner
var. aurantia sensu Bourd. et Galz. see Clavulinopsis amoena
C. incarnata Weinm.
C. incrassata Chev. = Clavulina rugosa var. alcyonaria Corner
C. incurvata Morgan = Ramaria incurvata (Morgan) Corner
C. ingrica Weinm. = Clavulina rugosa var. canaliculata Corner
C. intricata Ces. = Ramaria megalorhiza (B. et Br.) Corner
C. intricatissima Speg. = Ramaria intricatissima (Speg.) Corner
C. Invalii Cott. et Wakef. = Ramaria lnvalii (Cott. et Wakef.) Donk
C. isabellina Bres.
C. Jacquemontii Lev.
C. Janseniana Holterrn. (C. Janseiana) = Pterula.Janseniana (Holterrn.) Corner
C. javanensis B. ms., see Ramaria madagascariensis (P. Henn.) Corner
C. javanica Sacc. et Syd. = Clavicorona javanica (Sacc. et Syd.) Corner
C. juncea Fr. = Clavariadelphus junceus (Fr.) Corner
C. juncea Lev. = Typ hula gyrans Fr.
C. Kalchbrenneri Mull. = Ramariopsis crocea (Fr.) Corner
C. Kalchbrenneri Sacc. = Ramaria Saccardoi (Syd.) Corner
C. Karstenii Sacc. et Syd. = Ramaria Karstenii (Sacc. et Syd.) Corner
C. kewensis Mass. = Ramaria stricta (Fr.) Quel.
C. kisantuensis Sacc. = Ramaria kisantuensis (Sacc.) Corner
C. Klotzschii Lasch = Clavariadelphus junceus (Fr.) Corner
C. Krombholzii Fr. = Ramariopsis Kunzei var. deforrnis Corner
C. Kunzei Fr. = Ramariopsis Kunzei (Fr.) Donk
C. laciniata Pers. = Clavulina cristata var. fimbriata Corner
C. laeta B. et Br. = Clavulinopsis miniata (B.) Corner
C. laeta sensu Lloyd = Clavulinopsis amoena (Zoll. et Mor.) Corner
C. laeticolor B. et C., see Clavulinopsis amoena (Zoll. et Mor.) Corner
C. laetissima B. = Clavicorona pyxidata (Fr.) Doty
C. lanceolata Imai = Clavaria argillacea Fr.
C. Lauri Bory= a diseased, woody excrescence of Laurus nobilis (f. Tulasne, Se!. Fung.
Carp. 1, 1861, 125).
C. lavendula Pk. = Clavaria Zollingeri Lev.
C. Lecomtei Pat.
C. Ledermanni Bres. = Clavulinopsis Lederrnanni (Bres.) Corner
C. lentofragilis Atk., see Ramariopsis Kunzei (Fr.) Donk
C. leucotephra B. et C. = Ramaria stricta var. concolor Corner
C. Leveillei Sacc. = Clavulina Leveillei (Sacc.) v. Ov.
C. ligata Britz., see Clavulinopsis fusiforrnis (Fr.) Corner, and Clavulina amethy-
stinoides (Pk.) Corner
C. ligula Fr. = Clavariadelphus ligula (Fr.) Donk
C. liguloides P. Henn. et Nym. = Clavulinopsis liguloides (P. Henn. et Nym.) Corner
C. lilacina Fr. = Clavulina amethystina var. lilacina Quel.
C. lilacina Jungh., see Clavaria Zollingeri Lev.
C. longicaulis Pk. = Ramaria longicaulis (Pk.) Corner
C. longipes Karst., see Pistillaria setipes Grev.
C. lorithamnus B. = Ramariopsis lorithamnus (B.) Corner
C. lurida Kalchbr. = Aphelaria dendroides (Jungh.) Corner
C. lutea Vent. et Vitt. = Ramaria flava (Fr.) Quel.
C. luteo-alba Rea = Clavulinopsis luteo-alba (Rea) Corner
C. luteola Pers. = Clavariadelphus ligula (Fr.) Donk
C. luteo-ochracea Cav. = Clavulinopsis luteo-ochracea (Cav.) Corner
C. luteostirpata Fawcett = Clavulinopsis luteostirpata (Fawcett) Corner
C. luteo-tenerrima v. Ov. = Clavulinopsis luteo-tenerrima (v. Ov.) Corner
C. luticola Fr. = Clavulinopsis luticola (Fr.) Corner
C. Macounii Pk. = Clavulinopsis Macounii (Pk.) Corner
C. macropus Fr. = Clavulinopsis subtilis (Fr.) Corner
C. macrorrhiza Fr. = Clavariadelphus fistulosus f. macrorrhiza Bourd. et Galz.
C. macrospora Brinkm. = Ramaria macrospora (Brinkm.) Corner
C. macrospora Britz. = Clavulina rugosa var. macrospora Britz.
C. madagascariensis P. Henn. = Ramaria madagascariensis (P. Henn.) Corner
C. Mairei Battetta = Clavicorona Mairei (Battetta) Corner
C. maricola Kauffm. = Clavariadelphus maricolus (Kauffm.) Corner
C. meakanensis Imai = Clavaria argillacea Fr.
C. megalorhiza B. et Br. = Rarilaria megalorhiza (B. et Br.) Corner
C. merismatoides Schw. = Tremellodendron
C. micans (Fr.) Karst. = Pistillaria micans Fr.
C. Michelii Rea = Clavulinopsis Michelii (Rea) Corner
C. microscopica Malb. et Sacc. = ? Ceratellopsis aculeata (Pat.) Corner
var. Asphodeli Pat. = Ceratellopsis asphodeli Corner
C. microspora Josser. = Clavulinopsis rufipes (Atk.) Corner
C. miltina B. = Clavulinopsis miniata var. sanguinea Corner
C. miniata B. = Clavulinopsis miniata (B.) Corner
C. miniata sensu Petch = Clavaria helicoides Pat. et Dern.
C. minor Lev., see Typhula phacorrhiza Fr.
C. minuta Pers., see Pistillaria fulgida Fr.
C. minutula Bourd. et Galz. = Clavulinopsis minutula (Bourd. et Galz.) Corner
f. subasperata Bourd. et Galz. = Ramariopsis Kunzei var. subasperata Corner
C. mira Pat. = Clavariadelphus mirus (Pat.) Corner
C. misella B. et C. = Pistillaria misella (B. et C.) Corner
C. Miyabeana Imai = Clavulinopsis aurantia-cinnabarina (Schw.) Corner
C. molaris B. (1878) = ? Corticium pezizoideum (fide Coker: no type)
C. mucida Fr. = Lentaria mucida (Fr.) Corner
C. mucronella Bres. = Mucronella brasiliensis Corner
C. Mulleri B. = Clavaria vermicularis Fr.
C. Murrillii Coker = Ramaria Murrillii (Coker) Corner
C. muscicola Pers. = Eocronartium muscicolum (Pers.) Fitzp.
C. muscigena Karst. = Eocronartium muscicolum (Pers.) Fitzp.
C. muscigena Pers. = Ramaria crispula (Fr.) Quel.
C. muscoides Fr. = Clavulinopsis corniculata (Fr.) Corner
C. muscorum Cum. (1805) ex Bres. = nomen invalidum
C. muscorum Karst. = ? Pistillaria tenera Corner
C. mutans Burt = Clavulina cristata var. mutans Moeller
C. myceliosa Pk. = Ramaria myceliosa (Pk.) Corner
C. nebulosa Pk. = Clavaria purpurea Fr.
C. nebulosoides Kauffm.
C. nguelensis P. Henn. = Ramaria nguelensis (P. Henn.) Corner
C. nigra Lev. = ? Pyrenomycete
C. nigrescens Brinkm. = Ramaria nigrescens (Brinkm.) Donk
C. nigrescens Fr., see Ramaria Zippelii (Lev.) Corner
C. nigricans Lloyd
C. nigrita sensu Bresadola = Clavaria atrobadia Corner
C. nigrita sensu Coker = Clavaria atroumbrina Corner
C. nigrita Fr. = Geoglossum nigritum (Fr.) Cke.
C. nivea Quel. = Clavaria vermicularis Fr.
C. nodulosperma Atk. = Scytinopogon angulisporus (Pat.) Corner
C. Nymaniana P. Henn. = Clavaria Zollingeri Lev.
C. oblectanea Britz. = Ramaria testaceo-flava (Bres.) Corner
C. obtecta Britz., see Ramaria flavo-brunnescens (Atk.) Corner
C. obtusa Pers., see Pistillaria uncialis and Typhula quisquiliaris
C. obtusata Boud. = Clavaria argillacea var. obtusata Favre
C. obtusissima Pk. = Ramaria obtusissima (Pk.) Corner
C. obtusiuscula Britz. = Clavulina cristata (Fr.) Schroet.
C. occidentalis Zeller = C. purpurea Fr.
C. ochraceo-salmonicolor Clel. = Ramaria ochraceo-salmonicolor (Clel.) Corner
C. ochraceo-virens Jungh. = Ramaria ochraceo-virens (Jungh.) Donk
C. odorata (Karst.) Sacc. = Clavulina odorata Karst.
C. ornatipes Pk. = Clavulina ornatipes (Pk.) Corner
C. ornithopoda Mass. = Aphelaria dendroides (Jungh.) Corner
C. ovata (Fr.) Karst. = Pistillaria pusilla Fr.
C. ovata Pers. = Pistillaria pusilla Fr.
C. pallescens Pk. = Clavaria argillacea Fr.
C. pallida B. et C., see Ramariopsis Kunzei (Fr.) Donk
C. pallida Bres. = Ramaria Mairei Donk
C. pallidorosea Fawcett, see Clavulinopsis amoena (Zoll. et Mor.)
C. palmata Pers. = Ramaria palmata (Pers.) Quel.
C. paludicola Lib. = ? Clavulinopsis vernalis (Schw.) Corner
C. paludosa Lundell, see Ramaria flavo-brunnescens var. aurea Coker
C. pampaeana Speg. (1881) = Clavulina pampaeana (Speg.) Corner
C. pampaeana Speg. (1899) = Ramaria subsigmoidea (Sacc. et Syd.) Corner
C. pamparum Speg. = Ramaria subsigmoidea (Sacc. et Syd.) Corner
C. panurensis B. = Clavulina panurensis (B.) Corner
C. paradoxa Karst. = Pistillaria paradoxa (Karst.) Corner
C. patagonica Speg. = Clavulinopsis patagonica (Speg.) Corner
C. Patouillardii Bres. = Lentaria Patouillardii (Bres.) Corner
C. paupercula B. et C. = nom. nud., see Pistillaria misella
C. Peckii Sacc. (1891) = Clavulinopsis corniculata (Fr.) Corner
C. Peckii Sacc. et Syd. (1905) = Ramariopsis vestitipes (Pk.) Corner
C. pellucidula Britz., see Clavulinopsis falcata (Fr.) Corner
C. peronata Pers. = Pistillaria peronata (Pers.) Corner
C. persimilis Cott. = Clavulinopsis pulchra (Pk.) Corner
C. Petersii B. et C. = Clavicorona pyxidata (Fr.) Doty
C. phacorrhiza Pers. = Typhula phacorrhiza Fr.
C. phaeocladia Pat. = Ramaria Zippelii (Lev.) Corner
C. phoenicea Zoll. et Mor. = Clavulinopsis miniata (B.) Corner
C. phyllophila Mcalp. = Pterula phyllophila (Mcalp.) Corner
C. pilosa Pers. = Clavariade.lphus fistulosus (Fr.) Corner
C. pilosa Burt = Clavulina pilosa (Burt) Corner
C. pinicola Burt, see Ramaria apiculata (Fr.) Donk
C. pinophila Pk. = Lentaria byssiseda Corner
C. piperata· Kauffm., see Clavicorona Colensoi (B.) Corner
C. pistillaris Fr. = Clavariadelphus pistillaris (Fr.) Donk
f. unicolor Coker = Clavariadelphus unicolor (B. et Rav.) Corner
C. pistilliforma Pers. = Clavaria vermicularis Fr.
C. platyceras Viv. = Ramaria botrytis (Fr.) Ricken
C. plebeia Fr.
C. plumosa Schw. = Pterula plumosa (Schw.) Fr.
C. pogonati Coker = Clavulinopsis pogonati (Coker) Corner
C. polita Fr.
C. portentosa B. et Br.
C. praetervisa Britz. = Clavulinopsis helvola (Fr.) Corner
C. pratensis Fr. = Clavulinopsis corniculata (Fr.) Corner
C. propera Bourd. et Galz. = Clavulinopsis propera (Bourd. et Galz.) Corner
C. pruinella Ces, = Ramaria stricta (Fr.) Quel.
C. pseudofiava Britz. = Clavulina rugosa (Fr.) Schroet.
C. pteruloides Pat., see Clavulina cirrhata (B.) Corner
C. Puiggarii Speg. = Clavulinopsis Puiggarii (Speg.) Corner
C. pulchella Boud. = Ramariopsis pulchella (Boud.) Corner
C. pulchra Pk. = Clavulinopsis pulchra (Pk.) Corner
C. pulchra sensu van d. Bijl = Clavulinopsis ochracea Corner
C. Pullei Donk
C. purpurascens Paulet = Ramaria botrytis (Fr.) Quel.
C. purpurea Fr., et var. australis Coker
C. pusilla (Fr.) Karst. = Pistillaria pusilla Fr.
C. pusilla Pk. = Ramaria pusilla (Pk.) Comer
C. pusilla Quel. = Nyctalis
C. pusio B. = Aphelaria pusio (B.) Comer
C. pyxidata Fr. = Clavicorona pyxidata (Fr.) Doty
C. quercicola Imai, see Pistillaria albobrunnea Pat.
C. quisquiliaris (Fr.) Karst. = Typhula quisquiliaris (Fr.) Comer
C. radiata Lev., see Clavicorona, p. 294.
C. ramentacea (Fr.) Karst. = Clavariadelphus junceus (Fr.) Corner
C. Raveneliana Coker = Ramaria conjunctipes (Coker) Corner
C. regularis Herpell
C. rhizomorpha B., see Lachnocladium aurantiacum (B. et Br.) Petch
C. rhodochroa Mont
C. Ridleyi Mass. = ? Pyrenomycete
C. Rielii Boud. = Ramaria botrytis (Fr.) Quel.
C. rivalis Britz., see Aphelaria tuberosa (Grev.) Comer
C. rosacea P. Henn. = Clavulinopsis miniata var. rosacea Corner
C. rosalana Petch = Clavaria Zollingeri Lev.
C. rosea Fr.
var. attenuata Cke.
var. subglobosa Corner
C. rosella (Fr.) Karst. = Ceratellopsis rosella (Fr.) Comer
C. rosella sensu Bourd. et Galz. = Clavaria rosea Fr.
C. rubella Pers. ap. Quel., see Clavaria purpurea Fr.
C. rubescens (Quel.) Sacc., see Clavulina rugosa (Fr.) Schroet.
C. rubra B. et Cke. (1876) = Calocera rubra B. et Cke.
C. rufa Fr., see Clavulinopsis helvola (Fr.) Corner
C. rufescens Fr. = Ramaria rufescens (Fr.) Corner
serisu Coker = Ramaria holorubella (Atk.) Corner
C. rufipes Atk. = Clavulinopsis rufipes (Atk.) Corner
C. rufobrunnea Coker
C. rufo-violacea Baria = Ramaria rufo-violacea (Baria) Quel.
0
C. rugosa Fr. -� Clavulina rugosa (Fr.) Schroet.

sensu Cleland, see Clavulina ingrata Corner
C. Saccardoi Syd. = Ramaria Saccardoi (Syd.) Comer
C. sachalinensis Imai = Clavariadelphus sachalinensis (Imai) Corner
C. sanguinea Coker = Ramaria sanguinea (Coker) Corner
C. sanguineo-acuta v. Ov. = Clavulinopsis miniata var. sanguinea Comer
C. Sarasinii Cotton = Scytinopogon angulisporus (Pat.) Corner
C. scabra B.
C. Schaefferi Sacc. = Clavulina amethystina var. lilacina Quel.
C. Schimadai Imai = Clavulina aurantio-cinnabarina (Schw.) Comer
C. Schroteri P. Henn. = C. compressa Schroet.
C. Schweinfurthiana P. Henn.
C. sclerotiicola Allesch. = Typhula sclerotiicola (Allesch.) Corner
C. sculpta Beck = Ramaria botrytis (Fr.) Quel.
C. scutellata de Bary = Typhula phacorrhiza Fr.
C. secunda B. = Ramaria secunda (B.) Corner
C. secundiramea Lev., see Pterula Commersonii (Lev.) Lloyd
C. seminicola P. Henn.
C. semivestita B. et Br. = Clavulinopsis semivestita (B. et Br.) Comer
C. setacea Kalchbr. = Pterula setacea (Kalchbr.) Comer
C. setipes Grev. = Pistillaria setipes Grev.
C. setosa Balb. et Nocc. = Ramariopsis Kunzei (Fr.) Donk
C. setulosa B. = Lachnocladium setulosum (B.) Lev.
C. sibutiana Har. et Pat.
C. similis Baud. et Pat. = Clavulinopsis helvola (Fr.) Corner
C. similis Pk. = Clavulinopsis corniculata (Fr.) Corner
C. simplex Retz. ap. Karst. = Clavaria vermicularis Fr.
C. simulans B., see Lachnocladium furcellatum Lev.
C. sinapicolor Clel. = Ramaria sinapicolor (Clel.) Corner
C. solida S. F. Gray = Clavaria vermicularis Fr.
C. soluta Karst. = Ramaria soluta (Karst.) Corner
C. sparassoides Speg. = Polyporaceae, seep. 267.
C. spathulata Chev. = Clavariadelphus pistillaris (Fr.) Dank
C. spathulata Pk. = Pistillaria spathulata Corner
C. spathuliformis Bres. = Clavulinopsis spathuliformis (Bres.) Corner
C. Spegazzinii Sacc., see Scytinopogon angulisporus (Pat.) Corner
C. sphaerospora E. et E. = Clavulina cinerea (Fr.) Schroet.
C. sphagnicola Baud. = Clavaria argillacea var. sphagnicola Corner
C. spiculospora Atk. (C. spiculisperma Atk.) = Ramaria grandis (Pk.) Corner
C. spinulosa Fr. = Ramaria spinulosa (Fr.) Quel.
C. spiralis Jungh. = Clavulinopsis spiralis (Jungh.) Corner
C. Sprucei B. = Clavulina Sprucei (B.) Corner
C. Stillingeri Coker, see Ramaria apiculata (Fr.) Dank
C. straminea Cott.
C. Strasseri Bres. = Ramaria Strasseri (Bres.) Corner
C. striata Fr.
C. stricta Fr. = Ramaria stricta (Fr.) Quel.
sensu Coker = Ramaria stricta var. concolor Corner
sensu Bres., et Bourd. et Galz. = Ramaria Bourdotiana Maire
var. condensata Lund., see Ramaria condensata (Fr.) Quel.
C. strigosa Pers. = Clavariadelphus fistulosus (Fr.) Corner
C. strigosa P. Henn. et Nym. = Clavulinopsis amoena (Zoll. et Mor.) Corner
C. subacuta Imai
C. subargillacea Ito et Imai = Clavulinopsis amoena (Zoll. et Mor.) Corner
C. subaurantiaca P. Henn. et Nym. = Clavulinopsis amoena (Zoll. et Mor.) Corner
C. subbotrytis Coker = Ramaria subbotrvtis (Coker) Corner
C. subcaespitosa Pk. = Ramariopsis Kunzei (Fr.) Dank
C. subcorticalis Schw. = Ramariopsis Kunzei (Fr.) Dank
C. su,bdecurrens Coker = Ramaria subdecurrens (Coker) Corner
C. subfalcata Atk., see Clavaria argillacea Fr.
C. subfalcata Karst. = Pistillaria typhuloides (Pk.) Burt
C. subfastigiata Britz. = Clavulinopsis subfastigiata (Britz.) Corner
C. subfennica Coker = Ramaria subfennica (Coker) Corner
C. subfistulosa .P. Henn.
C. subfiava Britz. = Clavulinopsis subflava (Britz.) Corner
C. sublilacina Karst. = Clavulina amethystina var. lilacina Quel.
C. subrugosa Clel. = Clavulina subrugosa (Clel.) Corner
C. subsigmoidea (Sacc. et Syd.) = Ramaria subsigmoidea (Sacc. et Syd.) Corner
C. subspinulosa Coker = Ramaria subspinulosa (Coker) Corner
C. subtilis Fr. = Clavulinopsis subtilis (Fr.) Corner
C. subulata B. et Cke. = Pterula Gordius (Speg.) Corner
C. subumbrinella Imai = Clavulinopsis subumbrinella (Imai) Corner
C. suecica Fr. = Ramaria suecica (Fr.) Dank
C. sulphurascens Schw. = Clavulinopsis sulphurascens (Schw.) Corner
C. surculus B. = Lentaria surculus (B.) Corner
C. Sydowii Bres. = Ceratellopsis Sydowii (Bres.) Corner
C. syringarum Pers. = Ramaria stricta (Fr.) Quel.
C. ·tasmanica B. = Clavulina tasmanica (B.) Corner
C. taxophila (Thom) Lloyd = Clavicorona taxophila (Thom) Doty
C. tenacella Fr.
C. tenacella Pers. sensu Boud. = Clavulina cinerea f. sublilascens Bourd. et Galz.
C. tenax Lev. = Pterula tenax Lev.
C. tenax Schw. = Tremellodendron tenax (Schw.) Burt
C. tenella Boud. = Clavulinopsis tenella (Boud.) Corner
C. tenerrima Ma!IS. et Cross!. = Clavulinopsis tenerima (Mass. et Cross!.) Corner
C. tenuicula Bourd. et Galz. = Clavulinopsis tenuicula (Bourd. et Galz.) Corner
C. tenuipes B. et Br.
C. tenuis Schw. = Pistillaria tenera Corner
C. tenuissima Lev. = ? Pterula capillaris (Lev.) Sacc.
C. tenuissima Sacc. = Ramariopsis pulchella (Boud.) Corner
C. testaceo-fiava Bres. = Ramaria testaceo-flava (Bres.) Corner
var. testaceo-viridis Atk. = Ramaria testaceo-viridis (Doty) Corner
C. testaceo-violacea Doty = Ramaria testaceo-violacea (Doty) Corner
C. testaceo-viridis Doty = Ramaria testaceo-viridis (Doty) Corner
C. tetragona Schw. = Clavulinopsis tetragona (Schw.) Corner
C. Thwaitesii B. et Br. = Ramaria Zippelii (Lev.) Corner
C. tjibodensis P. Henn. = Lentaria tjibodensis (P. Henn.) Corner
C. tochinaiana Imai = Typhula tochinaiana (Imai) Comer
C. Todei (Fr.) Karst. = Pistillaria Todei (Fr.) Corner
C. tonkinensis Pat.
C. tortilis Pers. = Clavariadelphus junceus (Fr.) Corner
C. trichoclada Sacc. et Pao!. = Lentaria surculus (B.) Comer
C. trichomorpha Schw. = Pistillaria trichomorpha (Schw.) Corner
C. trichopus Grev. = Pistillaria setipes Grev.
C. trichopus Pers. = Clavulina cristata (Fr.) Schroet.
C. trichotoma Lev., see Lentaria surculus (B.) Corner
C. truncata Lovejoy = Clavariadelphus truncata (Quel.) Donk
C. truncata Quel. = Clavariadelphus truncatus (Quel.) Donk
C. tsugina Pk. = Ramaria apiculata (Fr.) Donk
C. tuberosa Fr. = Clavariadelphus fistulosus (Fr.) Corner
C. tubulosa Fr. = Ramaria tubulosa (Fr.) Comer
C. typhoidea P. Henn., see Lachnocladium aurantiacum (B. et Br.) Petch
C. typhuloides Pk. = Pistillaria typhuloides (Pk.) Burt
C. umbraticola Leuba, see Ramaria sanguinea (Coker) Corner
C. umbrina B. = Clavulinopsis umbrinella (Sacc.) Corner
C. umbrina Lev. = Clavulinopsis umbrina (Lev.) Corner
C. umbrinella Sacc. c·, Clavulinopsis umbrinella (Sacc.) Corner
C. uncialis Grev. c.c Pistillaria uncialis (Grev.) Const. et Dufour
C. unicolor (B. et Rav.) Doty 0
-,Clavariadelphus unicolor (B. et Rav.) Corner
C. unistirpis Britz. = Clavulina rugosa var. alcyonaria Corner
C. velutina E. et E. = Ramariopsis Kunzei (Fr.) Donk
C. vermicularis Fr.
f. fasciata Bourd. et Galz.
var. gracilis Bourd. et Galz.
var. singaporensis Corner
var. sphaerospora Bourd. et Galz.
C. vermiculata auctt. = C. vermicularis Fr.
C. verna Coker = Ramaria verna (Coker) Corner
C. vernalis Schw. = Clavulinopsis vernalis (Schw.) Corner
C. verpiformis B. et Cke.
C. versatilis (Quel.) Bourd. et Galz. = Ramaria fumigata (Pk.) Corner
C. vestitipes Pk. = Ramariopsis vestitipes (Pk.) Corner
C. vinaceo-cervina Clel. = Clavulina vinaceo-cervina (Clel.) Corner
C. violacea Petch = Clavaria Zollingeri Lev.
C. virescens Gramberg = Ramaria ochraceo-virens (Jungh.) Donk
C. virgata Fr. = Lentaria virgata (Fr.) Corner
C. virgultorum Pers. = Clavariadelphus junceus (Fr.) Corner
C. viscosa Pers. = Calocera viscosa Fr.
C. vitellina Pers. = Clavulinopsis corniculata (Fr.) Corner
C. xanthosperma Pk. = Ramaria xanthosperma (Pk.) Corner
C. xylarioides Petch
C. xylogena B. et Br. = Pterulicium xylogenum (B. et Br.) Corner
C. Zippelii Lev. = Ramaria Zippelii (Lev.) Corner
C. Zollingeri Lev.
Clavariachaete Corner
C. Peckoltii (Lloyd) Corner
C. rubiginosum (Cke.) Corner
Clavariadelphus Donk
C. fistulosus (Fr.) Corner
f. macrorrhizus Bourd. et Galz.
var. contortus Corner
C. junceus (Fr.) Corner
C. ligula (Fr.) Donk
C. maricolus (Kauffm.) Corner
C. mirus (Pat.) Corner
C. pistillaris (Fr.) Donk
var. americanus Corner
C. sachalinensis (Imai) Corner
var. atrobrunneus Corner
C. unicolor (B. et Rav.) Corner
Clavariella Karst. = Rarnaria S. F. Gray emend. Donk
C. abietina (Fr.) Karst. = Ramaria lnvalii (Cott. et Wakef.) Donk
C. apiculata (Fr.) Karst. = Ramaria apiculata (Fr.) Donk
C. aurea (Fr.) Karst. = Ramaria aurea (Fr.) Quel.
C. Bataillei Maire = Ramaria Bataillei (Maire) Corner
C. botrytis (Fr.) Schroet. = Ramaria botrytis (Fr.) Ricken
C. byssiseda (Fr.) Karst. = Lentaria byssiseda Corner
C. cedretorum Maire, see Ramaria fumigata (Pk.) Corner
C. condensata (Fr.) Karst. = Ramaria condensata (Fr.) Quel.
C. corrugata Karst. = Ramaria flaccida (Fr.) Ricken
C. crispula (Fr.) Karst. = Ramaria crispula (Fr.) Quel.
C. cristata (Fr.) Karst. = Clavulina cristata (Fr.) Schroet.
C. decolorans Karst. = Ramaria decolorans (Karst.) Corner
C. dendroidea (Fr.) Karst. = Ramaria condensata (Fr.) Quel.
C. divaricata Karst. = Ramaria Karstenii (Sacc. et Syd.) Corner
C. eumorpha Karst. = Ramaria eumorpha (Karst.) Corner
C. fennica Karst. (1882) = Ramaria fennica (Karst.) Ricken
C. fennica Karst. (1891) = Ramaria dec_olorans (Karst.) Corner
C. fistulosa (Fr.) Karst. = Clavariadelphus fistulosus (Fr.) Corner
C. fiaccida (Fr.) Karst. = Ramaria flaccida (Fr.) Ricken
C. fiava (Fr.) Schroet. = Ramaria flava (Fr.) Quel.
C. formosa (Fr.) Karst. = Ramaria formosa (Fr.) Quel.
C. fragillima (Sacc. et Syd.) v. Ov. = Ramaria fragillima (Sacc. et Syd.) Corner
•
C. gracilis (Fr.) Karst. = R�maria gracilis (Fr.) Quel.
C. grisea (Fr.) Karst. = Clavulina cinerea (Fr.) Schroet.
C. holsatica P. Henn. = Ramaria holsatica (P. Henn.) Corner
C. ligula (Fr.) Karst. = Clavariadelphus ligula (Fr.) Donk
C. palmata (Pers.) Karst. = Ramaria palmata (Pers.) Quel.
C. paradoxa Karst. = Pistillaria paradoxa (Karst.) Corner
C. pistillaris (Fr.) Karst. = Clavariadelphus pistillaris (Fr.) Do�k
C. soluta Karst. = Ramaria soluta (Karst.) Corner
C. spinulosa (Fr.) Karst. = Ramaria spinulosa (Fr.) Quel.
C. stricta (Fr.) Karst. = Ramaria stricta (Fr.) Quel.
C. suecica (Fr.) Karst. = Ramaria suecica (Fr.) Donk
C. trichopus (Pers.) Karst. = Ramaria suecica (Fr.) Donk
C. versatilis (Quel.) Maire = Ramaria fumigata (Pk.) Corner
Clavicorona Doty
C. candelabrum (Mass.) Corner
C. Colensoi (B.) Corner
C. coronata (Schw.) Doty, see C. pyxidata (Fr.) Doty
C. cristata (Kauffm.) Doty
C. javanica (Sacc. et Syd.) Corner
C. Mairei (Battetta) Corner
C. pyxidata (Fr.) Doty
var. asperospora Fawcett
C. taxophila (Thom) Doty
C. tuba (Heim) Corner
Clavulina Schroet.
C. amethystina (Fr.) Donk
var. lilacina Quel.
C. amethystinoides (Pk.) Corner
C. Bessonii (Pat.) Corner
C. cartilaginea (B. et C.) Corner
C. castaneipes (Atk.) Corner
C. cavipes Corner
C. chondro1des (B.) Corner
C. cinerea (Fr.) Schroet.
f. subcristata Bourd. et Galz.
f. sublilascens Bourd. et Galz.
var. gracilis Rea
var. odorata Bourd. et Galz.
C. cirrhata (B.) Corner
C. complanata Corner
C. connata (B.) Corner
var. bicolor Donk
var. coralloides Corner
var. curta Jungh.
var. fimbriata Fr.
var. flexosa Jungh.
var. incarnata Corner
var. lappa Karst.
var. mutans Moell.
var. nivea Bourd. et Galz.
var. subcinerea Donk
var. subrugosa Corner
C. decipiens Corner
C. delicia (B.) Corner
C. floridana (Singer) Corner
C. gallica Corner
C. gigartinoides Corner
C. gracilis Corner
C. humilis (Cke.) Corner
C. ingrata Corner
var. atricha Corner
C. odorata Karst.
C. ornatipes (Pk.) Corner
C. pampaeana (Speg.) Corner
C. panurensis (B.) Corner
C. pilosa Corner
f. mitruloides Bourd. et Galz.
var. alcyonaria Corner
var. canaliculata Corner
var. fuliginea Fr.
var. grisea Fr.
var. hercynica Fr.
var. macrospora Corner
C. Sprucei (B.) Corner
C. subrugosa (Clel.) Corner
C. tasmanica (B.) Corner
C. Vimderystii (Bres.) Corner
C. vinaceo-cervina (Clel.) Corner
Clavulinopsis v. Ov.
C. alcicornis (Zoll. et Mor.) Corner
C. amoena (Zoll. et Mor.) Corner
C. appalachiensis (Coker) Corner
C. Archeri (B.) Corner
C. arenicola Corner
C. asterospora (Pat.) Corner
C. aurantio-cinnabarina (Schw.) Corner
C. biformis (Atk.) Corner
C. boninensis (Ito et Imai) Corner
C. brevipe� Corner
C. calocera (Martin) Corner
C. candida (Weinm.) Corner
C. cineroides (Atk.) Corner
C. cinnamomea (Fawcett) Corner
C. citrino-alba (Moel!.) Corner
C. coliformis (Boud.) Corner
C. constans (Coker) Corner
C. corallino-rosacea (Clel.) Corner
C. corniculata (Fr.) Corner
f. simplex Donk
C. Daigremontiana (Boud.) Corner
C. depokensis (v. Ov.) Corner
C. dichotoma (God.) Corner
C. filipes Corner
C. flavella (B. et C.) Corner
C. Fleischeriana (P. Henn.) Corner
INDEX TO GENERA, SPECIES, AND SYNONYMS r2.r
C. fruticula Comer
C. fusiformis (Fr.) Comer
var. ceranoides W. G. Smith
C. griseola (Rea) Comer
C. hastula Comer
C. helvola (Fr.) Corner
var. geoglossoides Corner
C. Holmskjoldii (Oud.) Corner
C. Ledermanni (Bres.) Corner
C. liguloides (P. Henn. et Nym.) Comer
C. luteo-alba (Rea) Comer
f. montana Coker
var. latispora Corner
var. longispora Comer
C. luteo-ochracea (Cav.) Comer
C. luteo-stirpata (Fawcett) Comer
C. luteo-tenerrima (v. Ov.) Comer
C. luticola (Fr.) Comer
C. Macounii (Pk.) Corner
C. Michelii (Rea) Corner
C. miniata (B.) Corner
var. rosacea Corner
var. sanguinea Corner
C. minutula (Bourd. et Galz.) Corner
C. ochracea Corner
C. patagonica (Speg.) Corner
C. pogonati (Coker) Corner
C. propera (Bourd.) Comer
C. Puiggarii (Speg.) Corner
C. pulchra (Pk.) Corner
f. coccineo-basalis J osserand
C. pusilla (Coker) Corner
C. rufipes (Atk.) Comer
C. semivestita (B. et Br.) Comer
C. similis Comer
C. spathuliformis (Bres.) Corner
C. spiralis (Jungh.) Comer
C. subfastigiata (Britz.) Comer
C. subflava (Britz.) Comer
C. subtilis (Fr.) Comer
C. subumbrinella (Imai) Corner
C. sulcata v. Ov. ,,;, C. miniata (B.) Comer
C. sulphurascens (Schw.) Comer
C. tenella (Boud.) Comer
C. tenerrima (Mass. et Cross!.) Comer
C. tenuicula (Bourd. et Galz.) Comer
C. tetragona (Schw.) Comer
C. umbrina (Lev.) Corner
C. umbrinella (Sacc.) Comer
C. vernalis (Schw.) Comer
Cnazonaria Cda. = Pistillaria Fr.
C. aculina (Quel.) Donk = Pterula gracilis (B. et Desm.) Comer
C. erythropus (Fr.) Donk = Typhula erythropus Fr.
C. inaequalis (Fr.) Donk = Pistillaria setipes Grev.
5119
3A
C. Rickii (Oud.) Donk= Ceratellopsis Rickii (Oud.) Corner
C. setipes (Grev.) Cda. = Pistillaria setipes Grev.
Cornicularia Bon. = Clavulinopsis v. Ov.
Craterellus corrugis Pk. = Clavariadelphus unicolor (B. et" Br.) Corner
C. cristatus Kauffm.= Clavicorona cristata (Kauffm.) Doty
C. pistillaris Fr. = Clavariadelphus truncatus (Quel.) Donk
C. taxophilus Thom = Clavicorona taxophila (Thom) Doty
C. unicolor B. et Br.= Clavariadelphus unicolor (B. et Br.) Corner
Crinula paradoxa B. et C. = Cronartium quercuum Schroet.
Deflexula Corner
D. fascicularis (Bres. et Pat.) Corner
D. lilaceo-brunnea Corner
D. nivea (Pat.) Corner
D. pacifica (Kobayashi) Corner
D. ulmi (Pk.) Corner
Dendrocladium Lloyd= Ramaria S. F. Gray emend. Donk
D. Archeri (B.) Lloyd= Thelephora Archeri B.
D. congestum (B.) Lloyci= Thelephora congesta B.
D. fructicolum Lloyd, see p. 534
D. furcellatum (Fr.) Lloyd= Lachnocladium furcellatum (Fr.) Lev.
D. guadelupense (Lev.) Lloyd = Ramaria guadelupensis (Lev.) Corner
D. hirsutum (P. Henn.) Lloyd = Pterula hirsuta P. Henn.
D. juruense (P. Henn.) Lloyd= Pterula juruensis P. Henn.
D. Peckoltii Lloyd = Clavariachaete Peckoltii (Lloyd) Corner
Dimorphocystis Corner
D. capitatus Corner
D. laevis Corner
D. subcapitatus Corner
Eoagaricus Krieger= Physalacria Pk.

E. infiatus (Schw.) Krieger = P. inflata (Schw.) Pk.
Eriocladus Lev. = Lachnocladium Lev.
E. acicularis Lev.= Aphelaria dendroides (Jungh.) Corner
E. brasiliensis Lev. = Lachnocladium brasiliense Lev.
E. compressus (B.) Lev., see Lentaria surculus (B.) Comer
E. funalis Lev. = Aphelaria dendroides (Jungh.) Corner
E. furcellatus (Fr.) Lev. = Lachnocladium furcellatum (Fr.) Lev.
E. scoparius Lev. = Aphelaria dendroides (Jungh.) Corner
E. setulosus (B.) Lev. = Lachnocladium setulosum (B.) Lev.
E. tubulosus (Fr.) Lev.= Ramaria tubulosa (Fr.) Corner
Gliocoryne Maire = Pistillaria Fr.

G. uncialis (Grev.) Maire = Pistillaria uncialis Grev.
Gliocoryne auctt.= Gliocoryne Maire = Pistillaria Fr.
Heringia Schw., see Caripia O.K.

Holocoryne Bon. = Clavaria Fr.
Hormomitaria Comer
H. sulphurea Corner
Hydnum himantia Schw., see Clavaria himantia (Schw.) Fr., p. 265
H. sclerodontium B. et Mont. see Pterula sclerodontia (B.) Lloyd, p. 536
H. Sprucei Mont., see Pterula subplumosa P. Henn.
Hypolyssus B.= Caripia O.K.
H. clavarioides (Torrend) Lloyd (1915) = Lachnocladium moniliforme P. Henn.
H. foetidus Mass. = Caripia Montagnei (B.) O.K.
H. Montagnei B. = Caripia Montagnei (B.) 0.K.
H. Sprucei Mass. = Caripia Montagnei (B.) O.K.
Isaria aurantiaca B. et Br. = Lachnocladium aurantiacum (B. et Br.) Petch
I. calva Fr. = Mucronella calva Fr.
Lachnocladium Lev.
L. aciculata Lev., see Aphelaria dendroides (Jungh.) Corner
L. acutissimum (B.) Sacc. = Ramaria acutissima (B.) Corner
L. albidum Pat. = Ramaria albida (Pat.) Corner
L. albocinereum Pat. = Ramaria albocinerea (Pat.) Corner
L. ambiginosum Cke. = L. brasiliense Lev.
L. Archeri (B.) Bres. = Thelephora Archeri B.
L. articulatum P. Henn. = Ramaria articulata (P. Henn.) Corner
L. asterosetosum P. Henn. = L. divaricatum Pat.
L. Atkinsonii Bres., see Lentaria Micheneri (B. et C.) Corner
L. aurantiacum (B. et Br.) Petch
L. bicolor (Pk.) Burt = Ramariopsis vestitipes (Pk.) Corner
L. brasiliense Lev.
L. brunneum Bres., see Ramaria Moelleriana (Bres. et Roum.) Corner
L. cartilagineum B. et C. = Clavulina cartilaginea (B. et C.) Corner
L. cervino-album P. Henn. = L. divaricatum Pat.
L. cervinum (B. et C.) Pat., see Ramariopsis Kunzei (Fr.) Donk
L. chartaceum Pat. = Scytinopogon angulisporus (Pat.) Corner
L. cirratum Pat., see Scytinopogon angulisporus (Pat.) Corner
L. cladonioides P. Henn.
L. clavarioideum Pat.
L. compressum (B.) Lev., see Lentaria surculus (B.) Corner
L. congestum (B.) Lloyd = Thelephora congesta B.
L. cornicularioides P. Henn. = Ramaria cornicularioides (P. Henn.) Comer
L. cristatum Lloyd
L. dealbatum (B.) Cke:, see Ramariopsis Kunzei (Fr.) Donk
L. defiectens Bres. = Aphelaria deflectens (Bres.) Corner
L. delicia (B.) Cke. = Clavulina delicia (B.) Corner
L. dendroides (Jungh.) Sacc. et Syd. = Aphelaria dendroides (Jungh.) Corner
L. dilatatum (Mont.) = Calocera (?) dilatata Mont.
L. divaricatum Pat.
var. cinnamomeum Corner
var. sordidum Corner
L. dubiosum Bres. = Scytinopogon angulisporus (Pat.) Corner
L. Dusenii P. Henn.
L. echinosporum Bres. = Ramaria Zippelii (Lev.) Corner
var. microsporum Beeli = Scytinopogon angulisporus (Pat.) Corner
L. Englerianum P. Henn. = Ramaria cyanocephala (B. et C.) Corner
L. erectum Burt = Ramaria stricta var. concolor Corner
L. fiagelliforme (ll.) Cke. = Aphelaria dendroides (Jungh.) Corner
L. flavidum Corner
L. fulvum Corner
L. funale (Lev.) Sacc. = Aphelaria dendroides (Jungh.) Corner
L. furcellarioides P. Henn., see Clavicorona Colensoi (B.) Corner
L. furcellatum (Fr.) Lev.
L. galaxauroides P. Henn.
L. geniculatum Lev., see Scytinopogon angulisporus (Pat.) Corner
5119 3A2
L. giganteum Pat. = Ramaria cyanocephala (B. et C.) Corner
L. gu.adelupense (Lev.) Pat. = Ramaria guadelupensis (Lev.) Corner
L. guyanensis Pat. = Ramaria guyanensis (Pat.) Corner
L. hamatum P. Henn., see Clavulina cirrhata (B.) Corner
L. himalayense Mass.
L. Hisingeri (Karst.) Corner
L. Hoffmanni (P. Henn.) = L. reticulatum B. et C.
L. Hookeri B. = Ramaria Zippelii (Lev.) Corner
L. implexum (Lev.) P. Henn.
L. insigne Pat. = Ramaria cyanocephala (B. et C.) Corner
L. Janseanum (Holterm.) P. Henn. = Pterula Janseniana (Holterm.) Corner
L. Jansenianum (Holterm.) P. Henn. = Pterula Janseniana (Holterm.) Corner
L. Kurzii Cke. = Aphelaria dendroides (Jungh.) Corner
L. Lauterbachii P. Henn. = Ramaria Moelleriana (Bres. et Roum.) Corner
L. leucoceras Pat. = Ramaria leucoceras (Pat.) Corner
L. madeirense P. Henn. = L. divaricatum Pat.
L. manaosense P. Henn.
L. Micheneri B. et C. = Lentaria Micheneri (B. et C.) Corner
L. Moelleri P. Henn. = Scytinopogon angulisporus (Pat.) Corner
L. Moellerianum Bres. et Roum. = Ramaria Moelleriana (Bres. et Roum.) Corner
L. molle Corner
L. moniliforme P. Henn.
L. mussooriense P. Henn.
L. neglectum Mass. = Ramaria Zippelii (Lev.) Corner
L. oehraceum Bres. = Ramaria ochracea (Bres.) Corner
L. odoratum Atk. = Ramaria stricta (Fr.) Quel. sensu Coker
L. olivaceum P. Henn., see Ramaria cyanocephala (B. et C.) Corner
L. ornatipes (Pk.) Burt = C!avulina ornatipes (Pk.) Corner
L. pallens Bres. = Scytinopogon angulisporus (Pat.) Corner
L. pallescens Bres. = Scytinopogon angulisporus (Pat.) Corner
L. pallidum (B. et C.) Pat., see Ramariopsis Kunzei (Fr.) Donk
L. palmatifidum P. Henn.
L. palmatum P. Henn.
L. pteruliforme P. Henn.
L. pteruloides P. Henn.
L. pungens Lev. = Pterula pungens (Lev.) Sacc.
L. pusillum Coker = Clavulinopsis pusilla (Coker) Corner
L. quangense P. Henn.
L. ralumense P. Henn.
L. rameale B. et Br. = Septobasidium pteruloides (Mont.) Pat.
L. ramalinoides P. Henn.
L. reticulatum B. et C.
L. rubiginosum Cke. = Clavariacliaete rubiginosum (Cke.) Corner
L. samoense P. Henn. = L. brasiliense Lev.
L. Sarasinii P. Henn.
L. Schwackei P. Henn. = Scytinopogon angulisporus (Pat.) Corner
L. Schweinfurthianum P. Henn. = Scytinopogon angulisporus (Pat.) Corner
L. scoparium (Lev.) Sacc., see Aphelaria dendroides (Jungh.) Corner
L. semivestitum B. et Br. = Aphelaria tuberosa (Grev.) Corner
L. setosum (Pk.) Sacc. (1888) = Hirsutella setosa (Pk.) Pat.
L. setulosum (B.) Lev.
L. simplex P. Henn., see Clavulinopsis miniata (B.) Corner
L. simulans B. et Br. = Thelephora simulans (B. et Br.) Corner
L. strictissimum P. Henn., see Clavulina cirrhata (B.) Corner
L. strictum P. Henn., = Ramaria Moellcriana (Bres. et Raum.) Corner
L. subarticulatum P. Henn.
L. subcorticale (Schw.) Burt = Ramariopsis Kunzei (Fr.) Dank
L. subochraceum Sacc.
L. subpteruloides P. Henn. (1898)
L. subpteruloides P. Henn. (1900) = L. pteruliforme P. Henn.
L. subsimile B. et C. = Ramariopsis Kunzei (Fr.) Donk
L. taxiforme (Mont.) Sacc. = Pterula taxiformis Mont.
L. tonkinense Pat., see Lentaria surculus (B.) Corner
L. tubulosum (Fr.) Lev. = Ramaria tubulosa (Fr.) Corner
L. Ulei P. Henn.
L. usa�barense P. Henn. = Scytinopogon angulisporus (Pat.) Corner
L. Vanderystii Bres. = C!avulina Vanderystii (Bres.) Corner
L. vestitipes (Pk.) Burt = Ramariopsis vestitipes (Pk.) Corner
L. victoriense P. Henn.
L. violaceum Pat.
L. vitellinum Pat. = Pterula vitellina (Pat.) Corner
L. Warburgii P. Henn. = Ramaria Moelleriana (Bres. et Raum.) Corner
L. zandbaiense P. Henn. = Scytinopogon angulisporus (Pat.) Corner
L. Zenkeri P. Henn. = Scytinopogon angulisporus (Pat.) Corner
L. zonatum Corner
Lentaria Corner
L. affiata (Lagger.) Corner
L. byssiseda Corner
L. coronilla (Martin) Corner
L. corticola (Quel.) Corner
L. delicata (Fr.) Corner
L. epichnoa (Fr.) Corner
L. Micheneri (B. et C.) Corner
L. mucida (Fr.) Corner
L. Patouillardii (Bres.) Corner
L. surculus (B.) Corner
L. tjibodensis (P. Henn.) Corner
L. virgata (Fr.) Corner
Merisma aczculare Lev. = Aphelaria dendroides (Jungh.) Corner

M. capillaris Lev. = Pterula capillaris (Lev.) Sacc.
M. dendroides (Jungh.) Lev. = Aphelaria dendroides (Jungh.) Corner
M. guadelupense Lev. = Ramaria guadelupensis (Lev.) Corner
M. implexa Lev. = Lachnocladium implexum (Lev.) P. Henn.
M. pungens Lev. = Pterula pungens (Lev.) Sacc.
M. tuberosa Grev. = Aphelaria tuberosa (Grev.) Corner
Mitrula infiata (Schw.) Cke. = Physalacria inflata (Schw.) Pk.
Mucronella Fr.
M. abnormis P. Henn. = M. aggregata Fr.
M. aggregata Fr.
M. alba Lloyd
M. argentina Speg.
M. brasiliensis Corner
M. calva Fr.
M. fascicularis Fr. (1874) = Protohydnum fasciculare (Fr.) Bres.
M. fascicularis Fr. sensu Lloyd = Deflexula fascicularis (Bres. et Pat.) Corner
M. minutissima Pk. = M. aggregata Fr.
M. pacifica Kobayashi = Deflexula pacifica (Koh.) Corner
M. pusilla Corner
M. ramosa Lloyd = M. aggregata Fr.
M. Rickii Oud. = Ceratellopsis Rickii (Oud.) Corner
M. subtilis Karst.
M. tenuipes Lloyd
M. togoensis P. Henn.
M. ulmi Pk. = Deflexula ulmi (Pk.) Corner
M. viticola Pass. et Beltr., see Pistillaria micans Fr.
Mucronia Fr. = Mucronella Fr.
M. aggregata Fr. = Mucronella aggregata Fr.
M. calva Fr. = Mucronella calva Fr.
Myxomycidium Mass.
M. guianense Linder
M. nodosum Linder
M. pendulum Mass.
Penicillaria Chev. = Pterula Fr.

P. multifida Chev. = Pterula multifida Fr.
Perona Fr. = Caripia O.K
P. Montagnei (B.) Fr. = Caripia Montagnei (B.) 0.K.
Phacorrhiza Pers. = Typhula Fr.
P. Amansii Brond., see Typhula ovata Karst.
P. erythropus (Fr.) Grev. = Typhula erythropus Fr.
P. filiformis Grev. = Typhula phacorrhiza Fr.
P. sclerotioides Pers. = Typhula sclerotioides (Pers.) Fr.
Phaeoclavulina Brinkm. = Ramaria S. F. Gray
P. Broomei (Cott. et Wakef.) v. Ov. = Ramaria nigrescens (Brinkm.) Donk
P. macrospora Brinkm. = Ramaria macrospora (Brinkm.) Corner
P. nigrescens Brinkm. = Ramaria nigrescens (Brinkm.) Donk
P. Zippelii (Lev.) v. Ov. = Ramaria Zippelii (Lev.) Corner
Phaeopterula P. Henn. = Pterula Fr.
P. hirsuta P. Henn. = Pterula hirsuta P. Henn.
P. juruensis P. Henn. = Pterula juruensis P. Henn.
Physalacria Pk.
P. aggregata Martin et Baker
P. andina Pat.
P. australiensis Corner
P. bambusae v. Hoehn.
P. brasiliensis (Rick) Corner
P. changensis Rostr.
P. clusiae Syd.
P. concinna Syd.
P. corticola Corner
P. Decaryi Pat.
P. inflata (Schw.) Pk.
P. Langloisii E. et E.
P. orinocensis Pat.
var. andina Pat. et Gaill. = P. andina Pat.
P. rugosa Rick
P. Sanctae-Martae Martin et Baker
P. sasae Imai
P. stilboidea (Cke.) Sacc.
P. tenera Syd. = P. andina Pat.
P. togoensis (P. Henn.) Corner
P. tropica Corner
P. tuba Heim = Clavicorona tuba (Heim) Corner
P. villosa Petch
Pistillaria Fr.
P. abietina (Fckl.) = Typhula abietina (Fckl.) Corner
P. acicula Bourd. et Galz. = Pterula gracilis (B. et Desm.) Corner
P. aciculata Dur. et Lev. = Ceratellopsis aciculata (Dur. et Lev.) Corner
P. actiniceps Petch = Chaetotyphula actiniceps (Petch) Corner
P. aculeata Pat. = Ceratellopsis aculeata (Pat.) Corner
P. aculina Quel. = Pterula gracilis (B. et Desm.) Corner
P. acuminata Fckl. = Ceratellopsis acuminata (Fckl.) Corner
sensu Bourd. et Galz. = ? Ceratellopsis graminicola (Bourd. et Galz.) Corner
P. albobrunnea Pat.
P. alnicola Pk.
P. Amansii (Brond.) Sacc., see Typhula ovata Karst.
P. ampelina Bourd. et Galz.
P. anceps (Karst.) Corner
P. attenuata Syd. = Pterula gracilis (B. et Desm.) Corner
P. Bartholomaei E. et E.
P. Batesii Pk.
P. bellunensis Speg.
P. bipindiensis (P. Henn.) Corner
P. Boudieri Pat.
P. brunneola (Pat.) Sacc. = Pistillina brunneola Pat.
P. bulbosa Pat. = Typhula bulbosa (Pat.) Corner
P. caespitulosa Sacc. = Ceratellopsis caespitulosa (Sacc.) Corner
P. candidula Karst. = P. setipes Grev.
P. capitata (Pat.) Sacc.
P. cardiospora (Quel.) = P. culmigena Mont. et Fr.
P. Carestiae Sacc. = Ceratellopsis Carestiae (Sacc.) Corner
P. carnea Preuss. = P. micans Fr.
P. cinereo-alba (Bon.) Sacc. (1891) = fungus imperfectus
P. clavulata Ellis, see Typhula trifolii Rostr.
P. cylindracea Karst. = P. setipes Grev.
P. culmigena Mont. et Fr.
sensu Karst., Quel., Schroet. = Ceratellopsis aculeata (Pat.) Corner
P. cupressiformis B. et C.
P. cylindrospora (Atk.) Corner
P. cytisi Pat.
P. diaphana Fr.
var. albobrunnea Quel. = P. albobrunnea Pat.
P. dryopteridis Imai = Ceratellopsis dryopteridis (Imai) Corner
P. elegans B. et C. = Typhula elegans (B. et C.) Corner
P. elegantula (Karst.) Corner
P. epiphylla (Quel.) Corner
P. equiseticola Baud. = Ceratellopsis equiseticola (Boud.) Corner
P. euphorbiae Fckl. = Typhula euphorbiae (Fckl.) Karst.
P. Ferryi (Quel.) Sacc. = Pterula gracilis (B. et Desm.) Corner
P. fruticum Karst. = P. pusilla Fr.
P. fuegiana (Speg.) Corner
P. fulgida Fr.
P. fulvida Karst. = P. fulgida Fr.
P. jurcata W. G. Smith = Clavaria tenuipes B. et Br.
P. fuscipes (Pers.) Corner
P. fusiformis Kauffm.
P. gilva (Lasch) Corner
P. glandulosa (Preuss) Corner
P. gracilis (B. et Desm.) Pat. = Pterula gracilis (B. et Desm.) Corner
sensu Pat. = ? Ceratellopsis aculeata (Pat.) Corner
P. graminicola Bourd. et Galz. = Ceratellopsis graminicola (Bourd. et Galz.) Corner
P. granulata Pat.
P. hederaecola Ces. = Typhula hederaecola (Ces.) Corner
P. Helenae Pat. = Ceratellopsis Helenae (Pat.) Corner
P. hyalina (Quel.) Sacc. = Pistillina hyalina Quel.
P. inaequalis Lasch, see P. setipes Grev.
sensu Pat. see P. albobrunnea Pat.
P. incarnata Desm. = P. micans Fr.
P. Johnsonii Mass. = fungus imperfectus, seep. 497
P. juncicola Bourd. et Galz. = Pterula gracilis (B. et Desm.) Corner
P. lignicola (Killerm.) Corner
P. limicola (Saut.) Corner
P. lividula (Roll.) Corner
P. macrospora (Pat.) Sacc. = Pterula macrospora (Pat.) Corner
P. maculaecola Fckl.
P. micans Fr.
var. Sedi Karst.
P. misella (B. et C.) Corner
P. montevidensis Speg.
(P. mucedinacea Boud. = P. mucedinea Boud.)
P. mucedinea Boud. = Ceratellopsis mucedinea (Boud.) Comer
P. mucoroides Sacc. = Ceratellopsis mucedinea (Boud.) Comer
P. Muelleri Saut. = Typhula Muelleri (Saut.) Corner
P. muscicola Fr. = Eocronartium muscicolum (Fr.) Fitzp.
P. mycophila (Fckl.) Comer
P. oleae (Maire) Corner
P. oryzae Remsb. = Pterula oryzae (Remsb.) Corner
P. ovata Fr. = P. pusilla Fr.
(P. paradoxa B. et C. nom. nud. = Cronartium quercuum Schroet.)
P. paradoxa (Karst.) Comer
P. Patouillardii Quel. = Typhula Patouillardii (Quel.) Corner
P. penniseti P. Henn. = Pterula penniseti (P. Henn.) Corner
P. peronata (Pers.) Comer
P. phaeosperma (P. Henn.) Corner
P. puberula B. = Typhula quisquiliaris (Fr.) Corner
var. viscidula Karst. = P. typhuloides (Pk.) Burt
P. purpurea W. G. Smith
P. pusilla Fr.
var. lanceolata Quel. = Ceratellopsis sagittaeformis (Pat.) Corner
P. Quiletii Pat. = Ceratellopsis Queletii (Pat.) Konr. et Maubl.
P. quercus Oud., see P. pusilla Fr.
P. quisquiliaris Fr. = Typhula quisquiliaris (Fr.) Comer
P. ramealis Cke. = Typhula ramealis (Lib.) Bourd. et Galz.
P. rhodocionides Corner
P. rosella Fr. = Ceratellopsis rosella (Fr.) Comer
P. rubra (Fautr. et Ferry) Sacc. = Pistillina rubra Fautr. et Ferry
P. sagittaeformis Pat. = Ceratellopsis sagittaeformis (Pat.) Corner
var. rubi Killerm., see P. pusilla Fr.
P. sclerotioides Fr. = Typhula Muelleri (Saut.) Comer
�- sedi Karst. = P. micans var. sedi Karst.
P. setipes Grev.
P. spathulata Corner
P. subfalcata Karst. = P. typhuloides (Pk.) Burt
P. subfasciculata (E. et E.) Corner
P. subpellucida B. et C.
P. subuncialis Corner
P. syringae Fckl.
P. tenera Corner
P. tenuipes (B. et Br.) Mass. = Clavaria tenuipes B. et Br.
P. tenuis (Fr.) Corner
P. Thaxteri Burt = Pistillina Thaxteri (Burt) Corner
P. thujicola Kauffm. = Ceratellopsis thujicola (Kauffm.) Corner
P. Todei (Fr.) Corner
P. tremula Sacc. = Ceratellopsis tremula (:::iacc,) Corner
P. trichomorpha (Schw.) Corner
P. typhicola Bourd. et Galz.
P. typhuloides (Pk.) Burt
P. uliginosa Crouan
var. albo-Iutea Keissler
P. uncialis Grev.
P. viticola Pk., see P. setipes Grev.
Pistillina Quel.
P. bnmneola Pat.
P. hyalina Quel.
P. paradoxa (B. et C.) Cke., nom. nud. = Cronartium quercuum Schroet.
P. Patouillardii Quel. = Pistillaria capitata (Pat.) Sacc.
P. rubra Fautr. et Ferry
P. stilboidea Cke. = Physalacria stilboidea (Cke.) Sacc.
P. Thaxteri (Burt) Corner
Podostrumbium Kz. ap. Lev., see Caripia
Polyozus Karst. see Aphelaria Corner
P. contortus Karst. = A. tuberosa (Grev.) Corner
P. Hisingeri Karst. = Lachnocladium Hisingeri (Karst.) Corner
Pterula Fr.
P. abietis Lloyd
P. aciculaeformis Lloyd
P. actinaeformis (B.) Lloyd = Aphelaria trachodes (B.) Corner
P. acutissima Mont. (ined.) = Aphelaria tropica (Mont.) Corner
P. adustipes (Speg.) Corner
P. angustata (Lev.) Sacc.
var. fodinaria Sacc.
P. arbuscula Bres., see P. capillaris (Lev.) Sacc.
P. aurantiaca P. Henn. = Aphelaria aurantiaca (P. Henn.) Corner
P. Bresadoleana P. Henn.
P. brunneola Corner
P. capillaris (Lev.) Sacc.
P. Commersonii (Lev.) Lloyd
P. debilis Corner
P. Decaryi Pat. (ined.) = Aphelaria pusio (B.) Corner
P. dendroides (Jungh.) Fr. = Aphelaria dendroides (Jungh.) Corner
P. densissima B. et C.
P. dichotoma Sauter
P. dilatata (Mont.) Corner
7 30 INDEX TO GENERA, SPECIES, AND SYNONYMS
P. divaricata Lev., see Lachnocladium divaricatum (B.) Pat.
P. divaricata Pk. = P. plumosa (Schw.) Fr.
P. epiphylla Corner
P. exserta Lloyd
P. falcatula (P. Henn.) Corner
P. Jascicularis Bres. et Pat. = Deflexula fascicularis (Bres. et Pat.) Corner
P. filaris (Kalchbr.) Corner
P. fructicola Bres.
P. fruticum Lloyd
P. fulvescens Bres.
P. fusispora Yasuda
P. Gordius (Speg.) Corner
P. gracilis (B. et Desm.) Corner
P. grandis Syd.
P. guadelupensis (Lev.) Sacc. = Ramaria guadelupensis (Lev.) Corner
P. himalayensis (Mass.) Lloyd = Lachnocladium himalayense Mass.
P. hirsuta P. Henn.
P. humilis Speg., see P. taxiformis Mont.
var. tucumanensis Speg.
P. importata P. Henn. = Pterulicium xy:logenum (B. et Br.) Corner
P. incarnata Pat.
P. incisa Lloyd
P. Janseniana (Holterm.) Corner
P. juruensis (P. Henn.) Corner
P. Landolphiae Lloyd
P. laxa Pat.
P. lutea Pruess (1851) = Anthina sp.
P. luzonensis Lloyd
P. macrospora (Pat.) Corner
P. Mannii Lloyd
P. merismatoides (Schw.) Sacc. = Tremellodendron
P. multifida Fr.
P. nana Pat.
P. nivea Pat. = Deflexula nivea (Pat.) Corner
P. oryzae (Remsb.) Corner
P. pallescens Bres. = Scytinopogon angulisporus (Pat.) Corner
P. penicillata Lloyd
. P. pennata P. Henn. = P. subplumosa P. Henn.
P. penniseti (P. Henn.) Corner
P. phyllophila (Mcalp.) Corner
P. plumosa (Schw.) Fr.
P. p1mgens (Lev.) Sacc.
P. pusilla Bres.
P. pusio (B.) Bres. = Aphelaria pusio (B.) Corner
P. rigida Donk
P. sclerodontia (B. et Mont.) Lloyd
P. secundiramea Speg.
P. setacea (Kalchbr.) Corner
P. setosa Pk. (1875) = Hirsutella setosa (Pk.) Pat.
P. simplex Sacc. et Pao!. = Pterulicium xylogenum (B. et Br.) Corner
P. squarrosa P. Henn., see Aphelaria aurantiaca (P. Henn.) Corner
P. subaquatica Bres. et Roum. = Pterulicium xylogenum (B. et Br.) Corner
P. subplumosa P. Henn.
P. subsimplex P. Henn.
INDEX TO GENERA, SPECIES, AND SYNONYMS 7 31
P. subtyphuloides Corner
P. subulaeformis (B. et C.) Corner
P. subulata Fr.
P. tahitensis Reichardt
P. tasmanica Lloyd = Aphelaria tasmanica (Lloyd) Corner
P. taxiformis Mont.
P. tenax (Schw.) Sacc.= Tremellodendron
P. tenerrima (P. Henn.) Corner
P. tenuissima (Curt.) Corner
P. tenuissirna Lev., see P. taxiformis Mont.
P. timorensis Torr.
P. togoensis P. Henn.
P. trachodes (B.) Lloyd = Aphelaria trachodes (B.) Corner
P. tropica Mont. = Aphelaria tropica (Mont.) Corner
P. typhuloides Corner
var. minor Corner
P. Uleana P. Henn.
P. Vanderystii P. Henn.
P. verticillata Corner
P. vinacea Corner
P. vitellina (Pat.) Corner
P. Winkleriana P. Henn.
P. xylogena (B. et Br.) Petch = Pterulicium xylogenum (B. et Br.) Corner
Pterulicium Corner
P. xylogenum (B. et Br.) Corner
Ramaria S. F. Gray emend. Donk

R. abietina Quel.= R. ochraceo-virens (Jungh.) Donk
R. acroporphyria Quel. R. botrytis (Fr.) Rick.
R. acutissima (B.) Corner
R. alba (Pers.) Quel. = Clavulina cristata var. coralloidcs Cornet
R. albida (Pat.) Comer
R. albocinerea (Pat.) Corner
R. amethystea S. F. Gray= Clavulina amethystina (Fr.) Donk
R. amethystina (Fr.) Quel. = Clavulina amethystina (Fr.) Donk
R. apiahyana (Speg.) Corner
R. apiculata (Fr.) Donk
var. compacta Bourd. et Galz.
R. articulata (P. Henn.) Comer
R. aurea (Fr.) Quel.
R. Bataillei (Maire) Comer
R. botrytis (Fr.) Rick.
f. parvula Bourd. et Galz.
var. alba Pearson
R. botrytoides (Pk.) Corner
R. Bourdotiana Maire
R. brunnea (Zeller) Corner
R. byssiseda (Fr.) Quel. �� Lentaria byssiseda Corner
R. cacao (Coker) Corner
R. capitata (Lloyd) Corner
R. capucina (Pat.) Corner
R. ceranoides S. F. Gray---� Clavulinopsis fusiformis var. ceranoides
R. cervicornis (A. L. Smith) Corner
R. cinerea S. F. Gray, et (Fr.) Quel. = Clavulina cinerea (Fr.) Schroet.
R. condensata (Fr.) Quel.
R. conjuncta (Pk.) Corner
R. conjunctipes (Coker) Corner
var. odora Coker
R. coralloides (Fr.) Quel.= Clavulina cristata var. coralloide,; Corner
R. coriacea S. F. Gray= Clavaria coriaria Pers.
R. cornicularioides (P. Henn.) Corner
R. corniculata S. F. Gray, et (Fr.) Quel.= Clavulinopsis corniculata (Fr.) Corner
R. crispula (Fr.) Quel.
R. cristata (Fr.) Quel.= Clavulina cristata (Fr.) Schroet.
R. cyanocephala (B. et C.) Corner
R. cyatheae (P. Henn.) Corner
R. cystidiophora (Kauffm.) Corner
R. decolorans (Karst.) Corner
R. divaricata (Pk.) Corner
R. durbana (van d. Bijl) Corner
R. elegans Huber, see R. flavo-brunnescens var. aurea Coker
R. eumorpha (Karst.) Corner
R. fastigiata (Fr.) Quel.= Clavulinopsis corniculata (Fr.) Corner
R. Favreae Quel. = Ramariopsis Kunzei var. Favreae Corner
R. fennica (Karst.) Rick.
R. filicicola (Fawcett) Corner
R. filicina (Sacc. et Syd.) Corner
R. flaccida (Fr.) Rick.
R. flava (Fr.) Quel.
R. flavo-alba Corner
R. flavo-brunnescens (Atk.) Corner
var. aurea Coker
R. formosa (Fr.) Quel.
R. fragillima (Sacc. et Syd.) Corner
R. fumigata (Pk.) Corner
R. gelatinosa (Coker) Corner
R. gracilis (Fr.) Quel.
R. grandis (Pk.) Corner
R. grossa (Fr.) Quel. = Clavulina rugosa var. alcyonaria Corner
R. guadelupensis (Lev.) Corner
R. guyanensis (Pat.) Corner
R. Henriquesii (Bres. et Roum.) Corner
R. holorubella (Atk.) Corner
R. holsatica (P. Henn.) Corner
R. ignicolor Bres.
R. incurvata (Morgan) Corner
R. intricatissima (Speg.) Corner
R. lnvalii (Cott. et Wakef.) Donk
R. Karstenii (Sacc. et Syd.) Corner
R. kisantuensis (Sacc.) Corner
R. Kunzei (Fr.) Quel. = Ramariopsis Kunzei (Fr.) Donk
R. leucoceras (Pat.) Corner
R. lilacina (Fr.) Quel. = Clavulina amethystina var. lilacina Quel.
R. longicaulis (Pk.) Corner
R. luteo-flaccida Corner
R. luteo-fusca Maire
R. macrospora (Brinkm.) Corner
R. madagascariensis (P. Henn.) Corner
R. Mairei Donk
R. megalorhiza (B. et Br.) Corner
R. Michaelis Huber, see R. flavo-brunnescens (Atk.) Corner
R. Moelleriana (Bres. et Roum.) Corner
R. Murrillii (Coker) Corner
R. myceliosa (Pk.) Gomer
R. nguelensis (P. Henn.) Corner
var. americana Corner
R. obtusissima (Pk.) Corner
R. ochracea (Bres.) Corner
R. ochraceo-salmonicolor (Clel.) Corner
R. ochraceo-virens (Jungh.) Donk
R. pallida (Bres.) Rick. = R. Mairei Donk
R. palmata (Pers.) Quel.
var. guttaurensis Bres.
var. pluridentata Karst.
R. polypus Corner
R. pratensis S. F. Gray = Clavulinopsis corniculata (Fr.) Corner
R. pusilla (Pk.) Corner
var. australis Coker
R. rubescens Quel., see Clavulina rugosa (Fr.) Schroet.
R. rufescens (Fr.) Corner
var. frondosarum Bres.
R. rufo-violacea (Baria) Quel.
R. rugosa S. F. Gray, et (Fr.) Quel. = Clavulina rugosa (Fr.) Schroet.
R. Saccardoi (Syd.) Corner
R. sanguinea (Coker) Corner
R. secunda (B.) Corner
R. sinapicolor (Clel.) Corner
R. soluta (Karst.) Corner
R. spinulosa (Fr.) Quel.
R. Strasseri (Bres.) Corner
var. alba Cott. et Wakef.
var. concolor Corner
var. fumida Pk.
var. violaceo-tincta Bourd. et Galz.
R. subbotrytis (Coker) Corner
var. intermedia Coker
R. subdecurrens (Coker) Corner
R. subfennica (Coker) Corner
R. subgelatinosa Corner
R. subsigmoidea (Sacc. et Syd.) Corner
R. subspinulosa (Coker) Corner
R. subtilis (Fr.) Quel. = Clavulinopsis subtilis (Fr.) Corner
R. suecica (Fr.) Donk
R. testaceo-flava (Bres.) Corner
R. testaceo-violacea (Doty) Corner
R. testaceo-viridis (Doty) Corner
R. tuberosa S. F. Gray = Clavaria tuberosa Fr.
R. tubulosa (Fr.) Corner
R. verna (Coker) Corner
R. versatilis Quel. = R. fumigata (Pk.) Comer
R. virescens (Gramb.) Br. Henn. = R. ochraceo-virens (Jungh.) Donlc
R. xanthospenna (Pk.) Corner
R. Zippelii (Lev.) Corner
var. gracilis Corner
Ramariopsis Donk
R. asperulospora (Atk.) Corner
R. asterella (Atk.) Corner
R. clavuligera (Heim) Corner
R. crocea (Fr.) Corner
R. curta (Fr.) Corner
R. Kunzei (Fr.) Donlc
var. deformis Corner
var. Favreae Corner
var. subasperata Corner
R. lorithamnus (B.) Corner
R. pulchella (Boud.) Corner
R. tenuiramosa Corner
R. vestitipes (Pk.) Corner
Scleromitra Cda. = Pistillaria Fr.

S. coccinea Cda. = Pistillaria micans Fr.
Sc(erotium complanatum Tode = Typhula gyrans Fr.
S. Junceum Tode = Typhula gyrans Fr.
S. semen Fr. = Typhula variabilis Riess
Scytinopogon Singer
S. angulisporus (Pat.) Corner
S. echinosporus (B. et Br.) Corner
S. pallescens (Bres.) Singer = S. angulisporus (Pat.) Corner
Sphaerula Pat. = Pistillaria Fr.
S. capitata Pat. = P. capitata (Pat.) Sacc.
Stelligera Heim ex Doty = Lachnocladium Lev.
S. membranacea Heim ex Doty = ? L. divaricatum Pat.
Stereum tuberosum (Grev.) Mass. = Aphelaria tuberosa (Grev.) Corner
Stichoclavaria Ulbr. = Clavaria Fr.
Stichoramaria Ulbr. = Clavulina Schroet.
Thelephora acanthacea Lev. = Aphelaria dendroides (Jungh.) Corner

T. actinaeformis B. = Aphelaria trachodes (B.) Corner
T. angustata Fr., see Ramaria cyanocephala (B. et C.) Corner
T. bidentata Pat. = Aphelaria dendroides (Pat.) Corner
T. clavarioides Torr. = Lachnocladium moniliforme P. Henn.
T. Commersonii Lev. = Pterula Commersonii (Lev.) Lloyd
T. contorta Karst. = Aphelaria tuberosa (Grev.) Corner
T. dendroides (Jungh.) Lev. = Aphelaria dendroides (Jungh.) Corner
T. funalis Lev. = Aphelaria dendroides (Jungh.) Corner
T. lactea Pat. = Scytinopogon angulisporus (Pat.) Corner
T. oubanguiensis Har. et Pat. = Aphelaria dendroides (Jungh.) Corner
T. scoparia Lev. = Aphelaria dendroides (Jungh.) Corner
T. trachodes B. = Aphelaria trachodes (B.) Corner
T. tuberosa (Grev.) Fr. = Aphelaria tuberosa (Grev.) Corner
Typhula Fr.
T. abietina (Fckl.) Corner
T. anceps Karst. = Pistillaria anceps (Karst.) Corner
T. asP,hodeli Pat. = T. sclerotioides (Pers.) Fr.
T. athyrii Remsb.
T. betae Rostr.
T. bipindiensis P. Henn. = Pistillaria bipindiensis (P. Henn.) Comer
T. borealis Ekstr., see T. idahoensis Remsb.
T. Bresadolae Sacc. et Dalla Costa = Eocronartium muscicolum (Fr.) Fitzp.
T. Brunaudii Quel. = Pterula gracilis (B. et Desm.) Comer
T. bulbosa (Pat.) Comer
T. buxi Maire
T. caespitosa Ces.
var. coacervata Bourd. et Galz.
T. candida Fr. = Pistillaria setipes Grev.
var. fruticum Bourd. et Galz., see Pistillaria pusilla Fr.
T. caricina Karst.
T. castaneipes Atk. = Clavulina castaneipes (Atk.) Comer
T. complanata (de B.) Schroet. = Typhula gyrans Fr.
T. corallina Quel.
T. crassipes Fckl.
T. culmigena (Fr.) Schroet. = Pistillaria culmigena Fr.
T. cylindrospora Atk. = Pistillaria cylindrospora (Atk.) Corner
T. cystidiophora Kauffm.
T. elegans (B. et C.) Corner
T. elegantula Karst. = Pistillaria elegantula (Karst.) Corner
'f. erythropus Fr.
T. euphorbiae (Fckl.) Fr.
f. virescens Rab.
T. falcata Karst. = Pistillaria typhuloides (Pk.) Burt
T. filicina Pk.
T. filiformis Fr. = Clavariadelphus junceus (Fr.) Corner
T. flavescens Saut., see T. phacorrhiza Fr.
T. Friesii Karst. = T. sclerotioides (Pers.) Fr.
T. fruticum Karst., see P. pusilla Fr.
T. fuscipes (Pers.) Fr. = Pistillaria fuscipes (Pers.) Corner
T. gilva Lasch = Pistillaria gilva (Lasch) Corner
T. glandulosa Preuss = Pistillaria glandulosa (Preuss) Corner
T. gracilis B. et Desm. = Pterula gracilis (B. et Desm.) Corner
T. gracilis sensu Bourd. et Galz. = Pistillaria typhicola Bourd. et Galz.
T. gracillima White = Pistillaria uncialis Grev.
T. graminum Karst.
var. filicina Karst.
T. graminum auctt. = T. incarnata Fr.
T. Grevillei Fr. = Pistillaria setipes Grev.
T. gyrans Fr.
T. gyrans auctt. = T. sclerotioides (Pers.) Fr. et T. betae Rostr.
T. hederaecola (Ces.) Corner
T. hirsuta Cke., see Pistillaria setipes Grev.
T. hyalina Jungh. = Chaetotyphula hyalina (Jungh.) Corner
T. idahoensis Remsb.
T. incarnata Fr.
T. intermedia Appel
T. Itoana Imai = T. incarnata Fr.
T. juncea (Fr.) Karst. = Clavariadelphus junceus (Fr.) Corner
T. lactea Tu!., see T. variabilis Riess
T. Laschii Rab.
T. latissima Remsb.
T. lignicola Killerm. = Pistillaria lignicola (Killerm.) Corner
T. limicola Saut. = Pistillaria limicola (Saut.) Corner
T. lividula Roll. = Pistillaria lividula (Roll.) Corner
T. longipes Karst., see Pistillaria setipes Grev.
T. lutescens Boud.
T. mucor Pat. see Pistillaria setipes Grev.
..
T. mucosa B. et C. = Ceratellopsis mucosa (B. et C.) Corner
T. Muelleri (Saut.) Corner
T. muscicola Fr. (1838) = Eocronartium muscicolum (Fr.) Fitzp.
T. mycophila Fckl. = Pistillaria mycophila (Fckl.) Pat.
T. neglecta Pat.
T. nivea Pat. = T. corallina Quel.
T. oleae Maire = Pistillaria oleae (Maire) Corner
T. ovata Karst.
T. ovata (Fr.) Schroet. = Pistillarii pusilla Fr.
T. pallens Maire
T. Patouillardii (Quel.) Corner
T. peronata (Pers.) Fr. = Pistillaria peronata (Pers.) Corner
T. pertenuis Remsb.
T. phacorrhiza Fr.
T. phaeosperma P. Henn. = Pistillaria phaeosperma (P. Henn.) Corner
T. pusilla (Fr.) Schroet. = Pistillaria pusilla Fr.
T. quisquiliaris (Fr.) Corner
T. ramealis Speg. et Roum.
T. ramentacea Fr. 0= Clavariadelphus junceus (Fr.) Corner
T. rubicola B. et C., see T. sphaeroidea Remsb.
T. sclerotiicola (Allesch.) Corner
T. sclerotioides (Pers.) Fr.
T. sclerotioides Karst. = T. Muelleri (Saut.) Corner
T. semen Quel. = T. variabilis Riess.
T. sphaeroidea Remsb.
T. stolonifera Quel. = T. sclerotioides (Pers.) Fr.
T. stricta Appel. = T. sclerotioides (Pers.) Fr.
T. subfasciculata E. et E. = Pistillaria subfasciculata (E. et E.) Corner
T. subphacorrhiza Britzm. = T. phacorrhiza Fr.
T. subulaeformis B. et C. = Pterula subulaeformis (B. et C.) Corner
T. subulata Remsb.
T. tenerrima P. Henn. = Pterula tenerrima (P. Henn.) Corner
T. tenerrima Speg., see Pterula tenerrima (P. Henn.) Corner
T. tenuis Fr. = Pistillaria tenuis (Fr.) Corner
T. tenuissima C. = Pistillaria tenuissima (C.) Corner
T. tochinaiana (Imai) Corner
T. Todei Fr. = Pistillaria Todei (Fr.) Corner
T. Traillii B. et C. = Tremellales
T. translucens B. et Br;, 'Not a fungus' (Massee)
T. trifolii Rostr.
T. tucumanensis Speg., see Pterula tenerrima (P. Henn.) Corner
T. uleana P. Henn., see Pterula Gordius (Speg.) Corner
T. umbrina Remsb.
T. variabilis Riess
T. variabilis auctt., see T. betae Rostr.
T. viburni Remsb.
T. villosa Fr. = ? Isaria chrysopoda Bres. (fide Killerm. 1934)
T. virgata Remsb.
GLOSSARY
Abaxial, describing the surface of the basidiospore away from the long axis of the
basidium.
Acerose, needle-shaped.
Achroic, without pigmentation (p. 21).
Acrochroic, especially coloured in the hyphal tips of the growing point (p. 22).
Acuminate, tapered to a point.
Adaxial, describing the surface of the basidiospore next to the long axis of the basidium:
usually that with the apiculus.
Adventitious, referring to branches of the fruit-body not developed from the growing
point or margin (p. 5).
Agglutinated, with the hyphal walls stuck together (p. 17).
Aguttate; describing spores and other cells not containing oil-drops.
Allantoid, sausage-shaped.
Alutaceous, the colour of buff leather.
Amyloid, giving a blue reaction with iodine.
Apiculus, the peg-like projection of the basidiospore attaching it to the sterigma.
Appressed, flattened on to the surface.
Arcuate, bent like a bow.
Attenuate, with diminishing substance (p. 7).
Axil, the distal side of a branch-junction.
Basinym, the earliest valid name of a species employing the same specific epithet as
the current legitimate binomial (Furtado, Gardens' Bull., S.S., 1937, 9, 233).
Bifid, divided half-way into two.
Binding hyphae, thick-walled, much-branched, interwoven and, often, coralloid hyphae.
Caespitose, growing in tufts.

Capitate, having a head.
Cartilaginous, with a horny homogeneous appearance, as of cartilage.
Caulocystidia, hyphal ends on the stem-surface resembling cystidia or sterile basidia.
Cerebriform, with an irregular brain-like appearance.
Clamps, clamp-connexions, see p. 19.
Clavarioid, having the appearance of Clavaria.
Clavate, club-shaped, or thickened distally.
Concrescent, referring to hyphae, growing tugether in a tissue.
Coriaceous, with a tough or leathery texture.
Cortex, the layer of diveqrent hyphae surrounding, in some fruit-bodies, the medulla
of longitudinal hyphae, and comparable with a sterile hymenium.
Cottony, with fine hyphae or fibrils, as strands of cotton.
Cristate, with small pointed conical, or tooth-like, projections (p. 7).
Cyanescent, turning blue.
Cystidia, sterile and, usually, enlarged or modified hymenial elements, either project
ing or immersed in the hymenium (p. 19).
Cystochroic, with pigment in the cell-vacuoles (p. 22).
Cytochroic, with pigment in the cytoplasm (p. 22).
Dichophyses, the apical cell, usually of a short lateral hypha, modified into a stalk and
repeatedly dichotomous, attenuate branches of limited growth (p. 85).
Dichotomous, divided into two branches.
7 38 GLOSSARY
Dimitic, with two systems of hyphae, usually skeletal and generative (see footnote,
p. 13).
Direct, referring to the growth of fruit-bodies in which cell-enlargement occurs at the
same time as cell-division.
Divaricate, extremely divergent, spread asunder.
E, the ratio of length to width (spores, basidia, &c.).

Echinulate, referring to spores set with acute spines.
Ectochroic, with pigment on the outside of the hypha (p. 22).
Endochroic, with pigment inside the cell (p. 22).
Epichroic, discolouring on injury (p. 21).
Epihymenium, a thin layer of interwoven hyphae developing on the surface of the
hymenium.
Epiphytic (applied to saprophytes and parasites), growing on definite plant-remains as
opposed to humus or soil (p. 2 I).
Euchroic, with true pigmentation, as opposed to epichroic (p. 21).
Fasciate, the condition of a stem when several have coalesced.

Fastigiate, clustered and erect, as in bundles.
Ferruginous, rust-coloured.
Fibrillose, composed of longitudinal fibrils, or appearing as if traversed by fine fibrils.
Filijorm, thread-shaped.
Flabellate, flabelliform, fan-shaped.
Flattened branching, with the apices of the branches flattening and dilating before
dividing (p. 4).
Fleshy, sappy, soft, putrescent.
Floccose, set with flecks of matted fibrils (flocci).
Furcate, forked.
Furfuraceous, scurfy, or with small, soft scales.
Fuscescent, becoming fuscous.
Fuscous, dusky, too brown for a grey.
Fusiform, thick, but tapering towards each end.
Generative hyphae, thin-walled septate branching hyphae with or without clamps, in

fruit-bodies with other kinds of hyphae (skeletal and binding).
Gloeocystidia, cystidia with dense, usually oleaginous, contents, thin- or slightly thick
walled, often long and serpentine, more or less deeply embedded in the hymenium
(p. 20).
Granular, referring to spore-contents, with finely divided cytoplasmic inclusions.
Gregarious, in colonies or troops, not caespitose.
Guttate, guttulate, referring to spore-contents, with a large oil-drop (gutta) or many
oil-droplets (guttulae).
Gymnocarpic, the fruit-body developing in naked state without enveloping tissue.
Hymenium, the layer of fertile basidia.

Hysterochroic, referring to fruit-bodies becoming slowly discoloured from base to apex
in age (p. 21).
Incrusted, referring to hyphae with matter excreted on the walls.

Indirect, referring to the growth of fruit-bodies in which cell-enlargement occurs
mainly after the period of cell-division.
Inflated, inflation, referring to hyphae with enlarged cells (p. 14)..
Juvenescence, the process of maturing at a normally immature stage of development.

GLOSSARY 739
Ligulate, strap-shaped.
Lipochroic, with pigment in oil-drops (p. 22).
Medulla, the part of the fruit-body composed mainly, or entirely, of longitudinal

hyphae (whether corticated or not).
Mesochroic, with pigment in the hyphal wall (p. 22).
Metachroic, changing colour through the appearance of a new pigment in maturer
tissue (p. 22).
Monaxial, with undivided stem or axis.
Monomitic, with one system of hyphae (see footnote, p. 13).
Monopodial, referring to a multiaxial fruit-body with the original axis persistent and
predominant.
Multiaxial, with the stem or axis divided into many branches.
Multifid, divided into many segments.
Multiguttulate, containing many oil-droplets (guttulae).
Obovoid, referring to spores widest in the distal half (away from the apiculus).
Ochraceous, of a dull yellow colour.
Oleocystidia, cystidia with an oily resinous exudate (p. 20).
Ovoid, referring to spores widest in the proximal half (next to the apiculus).
Perforate, referring to fruit-bodies with a hole extending through the apex or top into
the stem.
Phycophilous, growing with algae (p. 21).
Polychotomous, with the apex dividing simultaneously into more than two branches.
Primordial shaft, the initial monaxial, conical or conico-c:ylindric, state of the develop-
ing fruit-body.
Pruinose, powdered.
Pteruloid, having the characters of the Pterula fruit-body.
Puberulous, set with short hairs.
Pubescent, set with rather long hairs.
Radial branching, referring to branching in more than one plane, without flattening of
the apex previous to branching (p. 4).
Reticulate, referring to spores with a network of ridges on the surface.
Rhizomorph, a root-like mycelial strand of concrescent hyphae.
Rough, referring to spores with uneven surface with indistinct markings.
Rufescent, turning reddish.
Rugose, set with ridges: rugulose, with short or faint ridges.
Sclerotioid, having the appearance of a sclerotium.

Sclerotium, an aggregation of mycelial hyphae, forming a discrete body usually with a
coloured resistant skin (p. 160).
Secondarily septate, referring to hyphae in which the cells develop additional septa
(without clamps) subdividing them into rows of secondary cells (p. 14).
Secondary mycelium, the mycelium developed from the base of the fruit-body.
Setae, thick-walled, often coloured, rigid cystidia (p. 20).
Setose, setulose, bearing setae (or small setae, setulose): or macroscopically set with
small spines.
Simple fruit-body, unbranched.
Skeletal hyphae, thick-walled, mostly unbranched and aseptate, hyphae in dimitic and
trimitic systems (p. 16).
Spathulate, with a gradually widened and flattened blunt end, as a spatula.
Spinulose, set with small or short spines.
740 GLOSSARY
Squarrose, with spreading and outstanding processes.
Stolon, a rhizomorph ending in a sclerotium or fruit-body (p. 149).
Striate, marked with lines.
Strigose, set with long stiffish hairs or bristles.
Subacerose, referring to immature basidia, lanceolate-subacute.
Subcaespitose, tending to be caespitose, or in small tufts.
Subhymenium, the layer of interwoven hyphae between the hymenium and medulla or
trama, giving rise to the basidia.
Subiculum, a mycelial felt producing fruit-bodies.
Successive branching, referring to clavarioid fruit-bodies in which branching is not
synchronous.
Sulcate, grooved or furrowed.
Suppressed branching, referring to branches initiated but undeveloped (p. 7).
Synonym, a valid but illegitimate name, or an alternative name for a species in another
generic combination.
Terete, cylindrical, but usually tap,:'!ring.

Thickening hymenium, referring to a hymenium which thickens by outgrowth of new
basidia overtopping the previous ones (p. u).
Tomentose, with shaggy matted hairs.
Torsion, referring to twisted branches (p. 8).
Trimitic, with three systems of hyphae (see footnote, p. 13).
Troop, a group of fruit-bodies from one mycelium.
Truncate, with the end flattened and appearing as if cut off.
Turbinate, top-shaped, or turnip-shaped.
Uninfiated, referring to hyphae in which the cells do not expand (p. 14).
Velutinate, with a finely velvety appearance.

Verrucose, referring to spores set with blunt warts or short blunt spines: verruculose,
minutely warted.
Villous, shaggy with long hairs (not matted, as in tomentose).
Vinescent, turning wine-red.
Virescent, turning green.
Waxy, with the consistency or appearance of wax.

I
PLATE I
:! 5
r, 2, Clavaria Zollingeri, from Malaya (r, Form B, from Penang: 2, Form A,

from Singapore): 3, Clavaria straminea, from England (from a painting in
Cotton's notes): 4, Clavaria helicoides, from Johore: 5, 6, Clavaria incarnata, from
England: X j.
PLATE 2
--
r, Clavaria Gibbsiae var. megaspora, from Singapore: 3, var. tenuis, from

Singapore: 2, Clavaria argillacea, from England: 4, Clavaria acuta, from Eng
land: 5, Clavulinopsis spiralis, from Singapore: x l
PLATE 3
/\
' :--,ii,.
fl !
Clavariadelphus pistillaris, from England: X I.

PLATE 4
1, Clavulina cinerea and 2, var. gracilis, from England: 3, f. sublilascens, from

England: 4, C. cristata var. incarnata, from England: X 1.
PLATE 5
1, 2, Clavulina cartilaginea, from Singapore: 3, Lentaria surculus, from Singa

pore: X !,
\
A, Clavulina gracilis, from Singapore: B, Clavulinopsis arenicola, from Maiaya:

the remainder Clavulina Leveillei, from Singapore: x J.
PLATE 7
- -
I, Clavulinopsis alcicornis, from Malaya: 4, 7, Clavulinopsis miniata, from Singa
pore: 5, 6, var. rosacea: 2, j, var. sanguinea, from Singapore (on leaf) and Kedah
(on wood): X !.
PLATE 8
1, .2, Clavulinopsis luteo-alba, from England: 3, 6, C. amoena, orange form, from

Malaya: 4, S, C. pulchra, from England: X j.
PLATE 9
Clavulinopsis luteo-tenerrima (upper and middle row,s), from Singapore: C.

amoena (lower row), branched, caespitose, and fistulose forms from Malaya: X !.
PLATE IO
.
I
1, Clavulinopsis corniculata, from England: 2, C. helvola, from England: 3, C.

umbrinella, from England: 4, Ramariopsis pulchella, from England: s, Clavulin
opsis fruticula, from Singapore: 6, 7, C. spiralis, from Singapore: X !.
PLATE I I
I, 2, Lachnocladium zonatum, from Singapore: 3, Defiexula fascicularis, from

Singapore: 4, Lachnocladium julvum, from New Guinea, and S, from Singa
pore: X I.
PLATE I2
Ramariafragillima (upper left), from Singapore: R. Zippe/ii var. gracilis (upper

right), from Johore: R. Invalii (below), from England: X 1.
vI �
iJ 1
•
f
.
( .
3
H
(.;.)
· Ramaria ochraceo-virens, from England: 3, R. Zippelii var. gracilis, from

I, 2,
Johore: 4, R. luteo-fiaccida, from Sumatra: 5, R. gracilis, from England: X I.
PLATE 14
Ramaria Zippe/ii (upper row), from Singapor-e: R. cyanocephala (lower row),

from Singapore: Aphelaria dendroides (middle), from Singapore: · X !.
PLATE 15
1, Clavarzafossicola, from Singapore: 2, Hormomitaria sulphurea, from Kedah:

3, Clavulina ornatipes, from Singapore: 4, Defiexula lilaceo-brunnea, from Singa
pore: S, Clavicorona candelabrum, from Singapore: X I.
PLATE 16
1I ) ,j Ii
.!. I ,\,
:1 ,\
I
I, Pterula typhuloides, from Singapore, X 1!: 2. var. minor, from Singapore, X r!:
3, Chaetotyphula hyalina, from Singapore, X r: 4, 5, Pterulicium xylogenu.m from
Singapore, X r,
,..
Bishen Singh Mahendra Pal Singh

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23-A, New Connaught Place, P.O. Box 137,

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J

BISHEN SINGH MAHENDRA PAL SINGH

OXFORD UNIVERSITY PRESS

Affinity of Clavarioid genera 22

TAXONOMIC AND MORPHOLOGICAL NOTES ON GENERA 32

INDEX TO GENERA, SPECIES, AND SYNONYMS

polychotomous state. As the process is repeated the branchlets become

TllxT-FIG. 3. Diagrams of successive stages in the

'growing-point' so formed new growing-points appear at regular intervals

TEXT-FIG. 5. Flattened branching in one plane in Scytinopogon angulisporus, the

to the expanding, but attenuate, dying-away apices and so helps to produce

Thus the branched Clavarioid fruit-body consists of a series of growth-

TEXT-FIG. 7. Apex of a branch of Pterulicium xylogenum,

and Lachnocladit1..m may also be the absence of thickening in the former,

walled, more or less unbranched, colourless, skeletal hyphae. 1 Among the

TEXT-FIG. 9. Basidia and clamps of Clavulinopsis fusiformis, X 1 ,ooo.

TExT-FIG. 10. Basidia and hyphae, without clamps, of Clavaria Zollingeri,

TEXT-FIG. 12. Hyphae (uninflated) and hymenium of Aphelaria dendroides:

TEXT-FIG. 13. Dimitic hyphae of Pterula verticillata: normal skeletal-hyphae,

DESCRIPTION OF THE CLAVARIOID FRUIT-BODY 19

Lachnocladium and Clavicorona have gloeocystidia, which are somewhat

The small fruit-bodies of Typhula, Pistillaria, Ceratellopsis, &c., in the

PARTICULAR KINDS OF ENDOCHROISM

AFFINITY OF CLAVARIOID GENERA

COLLECTION AND STUDY

ARAEOCORYNE gen. nov.

I have not seen young primordia of Araeocoryne, but there can be no

. i!l \Jll\lljl l\l\l1 \l l1\ litf1 f1\ \1 11\ll'JI l�i

, i\ Jililij1)/li !1l/j1/J 1rII 11 1

ij� 1 \ i\1l1i1 1 t\1 :1 i1 1 1 :i1 :it11 /1 1tl

according to Berkeley and Fries, they are agarics parasitized by Hypocrea. In

CERATELLOPSIS Konr. et Maubl.

CLAVARIA Fr. s. str.

STRUCTURE AND DEVELOPMENT OF CLA VARIA ZOLLINGERI

CLAVARIACHAETE gen. nov.

DEVELOPMENT AND STRUCTURE OF C. PISTILLARIS

ToxT-FIG. 19. Clavicorona candelabrum: full-grown and developing fruit-bodies, x 2.

TEXT-FIG. 21. Clavicoronajavanica: polychotomous tips, X 15.

Dimitic State. Skeletal hyphae are typically modified longitudinal hyphae

THE FRUIT-BODY OF C. CANDELABRUM

l\'�� \-)' \ f ( 1-j;lfljp

Structure of the Secondary Mycelium. At the base of the fruit-body branches

TEXT-Frc. 24. Clavulina ornatipes: hyphae from the incipient hymenium

by the upgrowth of narrow hyphae on the surface and, when a sufficient

..... ........ .,..,_.,.---------"'""'""�-"'""�--------A..-

TEXT-FIG. 27. Clavulina cristata: diagrams of the structure of a cristate

�ii11 U; i/fl il\����l .

��:�:::: �o ;i:;:::::r:�e 'fn��:i:

j branches are thickened by the up-

;�l;/l;l 111 \\�1\\1tr:

2. C. ornatipes and C. decipiens with brown-walled, little-inflated hyphae,

TABLE II. SPORE-SIZE IN CLAVULINA CRISTATA, C. CINEREA, C. RUGOSA,

7-fJX6-8 France Bourdot and Galzin C. cristata

7-1ox6-8 France Bourdot and Galzin C. amethystina

7·5-8·5 X 6-7·5 U.S.A.(S.Car.) Coker C. amethystinoides

10·6X8·2 Average C. rugosa

CLAVULINOPSIS van Ov.

STRUCTURE OF THE FRUIT-BODY

In C. luteo-tenerrima it does not thicken; possibly, also, in C. similis, but I

DEVELOPMENT OF THE FRUIT-BODY OF CLAVULINOPSIS AMOENA

ToxT-FIG. 3 I. Lachnocladium zonatum: full-grown and developing dichophyses, X 500.

TEXT-FIG. 34. Lachnocladium divaricatum

,. which undergoes no subsequent elongation or thickening because the cells do

�ii11 U; i/fl il\��l .

Pterulicium has a similar sterile hymenium without caulocystidia. In De

(Sp. ochraceous, 6 X 4/L: fruit-body with dichotomous branch