Вы находитесь на странице: 1из 354


Monograph No. 17
Rolf Singer

Cl _t, Of CANCER

^ - ---------



Organization for Flora Neotropica
Bronx, New York 10458

December 28, 1976


Published for
Organization for Flora Neotropica
The New York Botanical Garden

Ghillean T. Prance, Chairman
Enrique Forero
K. P. Dumont
Alvaro Femandez-Perez, Ex officio


Clark T. Rogerson, Editor

Richard A. Howard
Peter H. Raven
Paul C. Silva
Frans Stafleu
William L. Stern

Monograph No. 17 was edited by Clark T. Rogerson and K. P. Dumont


Founded and Conducted under the Auspices of UNESCO


1. Central America and Mexico ................................. Jerzy Rzedowski

2. West Indies ........... .......................... Richard A. Howard
3. Colombia-Ecuador ........................................ Enrique Forero
4. Peru-Bolivia ......................................... Ramn H. Ferreyra
5. Venezuela-Guiana ....... ............ ..... Leandro Aristeguieta
6. Brazil ........................................ Luis Guimaraes de Azavedo

MonographNo. 17
Rolf Singer

-Vk\ L t*'11





Organization for Flora Neotropica
Bronx, New York 10458

December 28, 1976

Published with the financial assistance of UNESCO.

Copyright ? 1976

Published by
The New York Botanical Garden
Bronx, New York 10458

Library of Congress Catalog Card Number 76-21378

International Standard Serial Number 0-89327-009-1
International Standard Book Number 0071-5794



Tricholomataceaetribe MarasmieaeFayod consists of severalgenera, some of which

are particularlywell representedby a large numberof species in the neotropics and the
tropics in general. These generahave formerly been combined into one large genus
MarasmiusFries which has often been confused with Marasmiellusand Collybia. Inso-
far as some of these generahave no hymenophore but have a smooth hymenial surface,
they have been termed "thelephoraceous"in the classicalwide sense, but agreementhas
graduallybeen reached that most of these do not belong in the Thelephoraceaeas this
family is now understood, but rather in agaricoid"cyphellaceous"genera. Some of them
are so closely related to Marasmiusand Chaetocalathusthat there can be no doubt that
they are "reduced"forms of the Marasmieaediffering only in smooth hymenial surface
and, in some cases, "cyphelloid" habit. They have been recognizedas genera different
from Marasmiusand Chaetocalathusin all those cases where additional charactersseparate
them from either of these lamellategenera, aside from the absence of a hymenophore
and/or the cyphelloid habit.
In the present monographI have excluded from considerationall those fungi which
do not belong in the subtribesCrinipellinaeor Marasmiinae;thus, the subtribeOudeman-
siellinaehas been omitted because in an earlierpaper (Singer, 1964a) I have given a mon-
ographof the South American species of Oudemansiellaand Mycenella which, with very
few additions or corrections, can serve as a monographof the neotropical species of
these genera. The third genus of subtribeOudemansiellinae,viz Physocystidium, con-
sists of a single species, P. cinnamomea (Dennis) Singer, which has been adequately
treated by Dennis (1951a) and Singer (1962a, 1973), whereasthe fourth one, Strobil-
urus, is not found in the neotropics proper, but only in the north temperatezone.
This leaves the Crinipellinae(with pseudoamyloidhairs or appendageson sterile
surfaces)and Marasmiinae(without such hairs). The morphologicalfeatures have been
amply discussed in previouspapers, particularlymy earliermonographof Marasmius
(Singer, 1965), Crinipellisand Chaetocalathus(Singer, 1942) and the cyphelloid Maras-
miinae (Singer, 1960). Aside from general data on the morphology and chemical
characters,the readerwill find in these papers indications on the best way to collect
and annotate specimensof these fungi. The delimitationsof the variousgenera in-
volved and the scope of all genera of the Marasmieaeon a world-widebasis are dis-
cussed in Singer (1974).
Whilepreliminarystudies on many genera of Marasmieaehave been published
by Singer (1942, 1965) in the past for the South Americancontinent or as a survey

Professor Titular Emerito, Facultad de Ciencias Exactas y Naturales, University of Buenos

Aires, Argentina: Visiting Professor, University of Illinois at Chicago Circle; Research Associate,
Field Museum of Natural History, Chicago, Illinois 60605.
2 Flora Neotropica

of the known species of the world flora, it has become obvious that there is a need
for a new treatmentincludingthe numerousCentraland North Americanspecieswhile
omitting the exclusively temperateones. At the same time there is a need to consider
new taxa, discoveredin regions where Marasmieaehave been collected before - and
especially in regions where mycologists have never been active until after 1965, such
as the Cordillerasof Colombia and the Amazonas region of Ecuador. Including,then,
the new discoveriesof the period between 1965 and 1973 but excluding the Oudeman-
siellinae, the Marasmieaeas representedin the present monographcontain a total of
12 genera and 322 species.2
These figures confirm the impression- not unexpected to those who have
collected fungi in the tropics that the Marasmieaeare among the most importantflor-
istic elements of that vast region, followed by the Collybieae(compare my monograph
of the neotropical species of MarasmiellusSinger 1973).


The species of Marasmieae,particularlythe Marasmiinae,are mostly wood- and

litter-inhabitingfungi, all of which are epigeous with exogenous spore production. None
of them are ectomycorrhizal. Most are saprophytic,but severalare decidedly, some even
exclusively, parasitic(e g Crinipellisperniciosa(Stahel) Singer,C. siparunaeSinger,Maras-
mius viegasii Singer,M. crinisequiMullerex Kalchbrenner),and it is these species which
are of interest to the tropical phytopathologist. In contrast with the genusMarasmiellus
and with Omphalinaand Gerronema,lichenizationhas never been observed in the
Marasmieae. Some of the saprophyticspecies seem to be specializedas far as the host
materialis concerned. While some species occur on a wide range of hosts, others are
limited to conifers, Pteridophyta,Monocotyledonesor Dicotyledones. Finally there is
a rather largegroup of species which appearto be restricted to a single host genus or
family. Some species are obviously restrictedto high-alpine(andine) zones reachingup
to abott 5000 m altitude; others are restrictedto subxerophyticand xerophytic regions.
There are also some species which are most common in destroyed vegetation or wherever
ecosystems are in disequilibriumcaused by human activities. The rest is restrictedto
the tropical and subtropicalwet forest. Many of the latter appearto grow only in the
subtropical-tropicalmontane forests while others are confined to the plains of the Ama-
zonas basin and/or the coastal forests of Brazil,and/or the Gulf area, and/or the Pacific
rain forests of Panamasouth to Ecuador. The determiningfactors for such area restric-
tion cannot, in most cases, be ascertained. Certainly,sometimes it is geographicalisola-
tion, sometimes restriction of the host species and sometimes climatic factors, particul-
arly the presence or absence (Pacific coastal forests) of a dry period. Edaphic factors
such as the physical and chemical compostion of the soils are obviously of minor, if any
direct influence since the majority of these fungi are litter-inhabitantsor inhabitantsof
dead fallen woody material,or else parasitizingor otherwise epiphytic on living plants.
The area of distribution,as can be seen from the precedingparagraph,varies from
very restricted island-likeareaswithin some of the major landscapesof the neotropics
to panneotropicalor even pantropical. Some of the pantropical
species such as Maras-
mius haematocephaluswill occasionally occur in the temperate zone of North

2Including Oudemansiellinae, there would be 16 genera and 330 species.

Introduction 3

where they are introduced with plant materialfrom the subtropicaland tropical zones.
Others frequently invade the greenhousesof the temperate zones e. g. Crinipellissipar-
unae. If I exclude the collections of adventitious species from the statistics, I may say
that of the approximately 300 well defined species of Marasmiusless than 10%are pan-
tropical and as many as 229 have been recognized as occurringin the neotropics. This
hlighpercentageis not only due to the relatively large, perhapsexceptionally large rep-
representationof Marasmiusin the neotropics but, to a certain degree, also to less exten-
sive collecting and relatively few monographicrevisions(Ceylon, Zaire) in the paleo-
tropics. A similarpreponderanceof neotropical species can be observed in other genera
such as Gloiocephala,Crinipellis,Chaetocalathus. The generaPhysocystidium,Manuripia,
Epictwphus,Hymnenogloeahave thus far been observed only in the neotropics and only
Palaeocephalaexclusively in the palaeotropics.
The statistics for the whole of the Marasmieaeshow that among the species of
Marasmieae(without Oudemansiellinae)treated in the present monographthere are
species belonging in the following four groups;
Group I. Species occurringexclusively in the subtropicsand/or tropics of the Americas
or in parts of it: 274 species or 85.9%
Group II. Species occurringaside from an area in the neotropics also in some parts of
the paleotropics,Oceania or Australia: 21 species or 6.6%
Group III. Species occurringaside from some parts of the neotropics also in the tem-
perate regions or parts of them: 21 species or 6.6%
Group IV. Species occurring in the neotropics but also in the paleotropics and in parts
of the temperate zones: 3 species or 0.9%
One may argue that as the palaeotropicsare graduallyexplored so that eventually
our knowledge of the Marasmieaeflora of the paleotropics will match the knowledge we
have now of the neotropical species, the percentageof the species belongingin group II
will turn out to be largerthan indicated above. But it may also be expected that as we
begin to discover further elements of the neotropic flora thus far unknown, we are likely
to deal mostly with species rare and restricted in their distribution,and therefore likely
to be strictly neotropical. Thus, the chances that the percentagesreported above will
be substantiallyaltered after future research,appear to be small.


Since ecologists - not so inexplicably, since it is difficult to obtain the collabora-

tion of a competent mycologist - have, in spite of their importantrole, omitted the
fungi from considerationin the description of tropical vegetation types and associations,
I am using the term mycosociology instead of phytosociology for the following data.
This had to be so because these observationswere not coordinated with a study of
phanerogamicor other cryptogamicvegetation. One might wish that scientists active
in elaboratingand defining plant communities in the neotropics as well as in other
regions had termed their work cormophyte-sociology(or some such word) since they
have not given due considerationto the fungi or protists.
Five localities with virgin,undisturbedvegetation of the Amazonas region whose
non-fungus vegetation can easily be checked, have been selected in order to see
whether comparableaspects (i e individualsfruiting duringthe main rainy season) of
the fruiting consociation are similar. These localities have been subdividedinto (1-3)
terra-firmevegetation of the Amazonian lowlands and (4-5) inundatedor inundable
4 Flora Neotropica

forests just after the retreat of the flooding. In these localities extensive and inten-
sive collecting of Marasmieaehas taken place (other collections have been added in
The results are interestinginsofar as surprisinglyfew species are common to the
five localities even within their groupingaccordingto inundability. This may mean
two things, one exclusive of the other: either Marasmieae(and obviously other agarics)
are, as a group of organisms,not characteristicfor the higher "taxa" of phytosociolog-
ically definable forest communities in the Amazonas region or else they are extremely
sensitive to minor differences in microclimate,host-distributionand geographicaliso-
lation. Only these extreme views can be taken, but which one of the two is difficult
to decide.
My own experience in other regions of the neotropics tends to favor the latter
alternative,since even a glance at the phytogeographicand ecological data in the keys
and species descriptionsshows that many species are selective as far as area and associ-
ation are concerned. On the other hand, the collections of parts of the vast Amazonas
region are not extensive enough to permit us to prove this hypothesis by the lists of
species indicated below (Tables I, II).
When flying low over the hylaea with obviously basically uniform vegetation -
and at a time when a given tree with spectacularaspect during its flowering period or
with recognizablefeatures (like Bertholletia excelsa Humboldt & Beaupland)can be
pointed out and is easily traced as far as its distribution in the forest is concerned -
one can often notice that many of these trees, obviously characteristicfor the partic-
ular forest type, are widely spaced, often with distances of over 1 km between indiv-
iduals. If it is assumed then, given the dependence of the heterotrophic organismson
their hosts, that fungus species may be equally scattered in the forest and that even
repeatedvisits and very intensive collecting will not registerevery population in fruit-
ing - species cannot be detected when in the vegetative state - it becomes clear that
the necessarilyreduced collecting area and the fungi observed in it, cannot by any
means be complete enough to produce fully comparablelists.
On the other hand, if the Marasmieaewere, although widely scattered in the
Amazonas Basin, neverthelessevenly distributedamong the communities making up
the hylaea, one would expect at least some species to be common to all three terra-
firme localities. This, however, does not apply, except in the case of Marasmiius
haematocephalus, which is so common and adaptable that there is no tropical or
subtropicalforest type in the neotropics where this fungus does not occur, and the
same species has also been observed in the palaeotropicsand, adventitiously, in the
temperate zone of North America.
Somewhat more similaritybetween localities 1, 2 and 3, and 4 and 5 respect-
ively would be apparentif we were able to add all collections from the Napo river
region and compare them to all localities visited in the Beni, Mamore and Madrede
Dios regions, or all collections of the lower Para and all gathered in inundated for-
ests of the upper Amazonasand tributaries. Yet, at present this would be meaning-
less since most other localities in these regions, i e, those not included in the lists
given on Tables I and II, show deteriorationto a variabledegree; they are far from
virgin forest, bit are either second growth or partly destroyed by fire, selective wood-
cutting or cultivation. As we have shown in previous papers(Singer in Hacskaylo,
1971; Singer & Moser, 1965; Singer in Piterbarg,1966; Singer & Morello, 1960) such
stands tend to give shelter to a large number of species otherwise rare or absent, fre-
Introduction 5

quently having a particularlywide area including some cosmopolitan species and

species forming cicatrizingmycorrhizae. Consequently, we have to rely on the data
given here or else we might obtain misleadingresults. The only way to have the
lists amplified is to revisit the five localities (and some additional ones) repeatedly.
Unfortunately this becomes more difficult every year since the virgin forest is des-
troyed at an alarmingrate not only in the Amazon region but also over the entire
neotropics. This is also the reason why the thus far somewhat inconclusiveresults
presented here are believed to merit publication.
As far as other regions of the neotropics are concerned, similarlists are now in
preparation. The data presented here (Table I-III) as well as others referringto the
mycosociological exploration of the neotropicalplant communities suggest,if nothing
else, that it is certainly worth the trouble to collect data on the fungus component of
each association wheneverecological work on neotropical forest communities is under-
taken. The present monographwill help to make identification of such collections

Amazonas, Terra Firme forest

1. May aspect (highest 2. March (main rainy season) 3. March (Main rainy season)
monthly average of rainfall) aspect in virgin forest, hylaea, aspect in virgin hylaea Bolivia:
in virgin hyalaea, Ecuador: Bolivia: Pando, Conquista Beni, Mamore River
Napo, Shushufindi

Marasmius puttemansii var Marasmius manuripiensis Marasmius guyanensis var

puttemansii M. corrugatus erythrocephalus
M. variabiliceps var derubricans M. haematocephalus M. hylaeicola
M. atrorubens M. helvolus M. pararotula
M. cladophyllus var clado- M. pseudocorrugatus M. megalospermus
phyllus M. pusio M. allocystis
M. haediniformis M. trinitatis M. bambusinus
M. haematocephalus var pur- M. griseoroseus M. bellus
pureomarginatus M. cohortalis M. chrysocephalus
M. helvolus M. niveus M. cladophyllus var clado-
M. hypophaeus M. silvicola phyllus
M. leoninus var leoninus Chaetocalathus columellifer M. ferrugineus
M. leoninus var aberrans Epicnaphus phalaropus M. haematocephalus var
M. neosessilis var neo- Manuripiabifida pseudotageticolor
sessilis [Calocybe cyanea] M. leoninus var orinocensis
M. neosessilis var monte- [Clitocybe podocarpi] M. montagneanus
piensis [Gerronema cyathiforme] M. pleuracanthus
Gloiocephala anastomosans [Pluteus amphicystis] M. trinitatis
Oudemansiella canarii [Pluteus pluvialis] [Marasmiellus pilosus]
[Campanella elongatispora] [Pluteus riberaltensis var [M. nigripes]
[Marasmiellus eburneus] conquistensis] [Hydropus nigrita var]
[Filoboletus gracilis] [Ramaria flaccida] [Mycena chlorinosma]
[Hydropus citrinus] [Lentaria surculus] [Panus rudis var strigellus]
[Panus rudis var strigellus] [Pistillaria linearis var [Lactarius sp]
[Russula aucarum ined.] macrospora] [Dacryopinax spathularia]
[Auricularia fuscosuccinea] [Favolaschia cinnabarina] [Lentaria surculus]
[Favolaschia rubra] [Favolaschia echinata] [Clavaria zollingeri var
[Favolaschia cinnabarina] [Favolaschia zenkeriana] microspora]
[Favolaschia sabalensis var [Pterula fluminensis]
[Simocybe amazonica]
6 Flora Neotropica


4. Marchaspect in inundableforests along 5. May-Juneaspect of the inundablerain
the lower Madrede Di6s up to Conquista. forest in the lower Amazonas. BRAZIL:
Bolivia:Pando Para',Belem
Marasmiustereticeps Marasmiusrotuloides
M. guyanensis M. cladophyllus
M. pandoanus M. leoninus
M. dodecaphyllus [Marasmiellusstenophylloides]
M. panerythrus [Marasmiellusdefibulatusvarstrictior]
Gloiocephalalongifimbriata [Marasmiellusbisporiger]
Crinipelliseggersiivar flavipes [Marasmiellusaporposeptus]
C. excentrica [Marasmiellusnigripes]
[Marasmielluspandoanus] [Hydropusparaensis]
[Trogiacantharelloides] [Hydropusdepauperatus]
[Pluteus neophlebophorus] [Hydropussemimarginellus]
[Inocybe matris-dei] [Gerronemastrombodes]
[Pterulapalmicolal [Mycenaaosma]
[Lepiota xanthophylla]


6. Tropicalrain forest, spring-summeraspect, 7. Tropicalrain forest, summeraspect,

Gulf Coast (Estaci6n Biologicade los Tuxtlas, PacificCoast low hills region near Tuxtla de
(nearly virgin) Gutierrez:Selva between Finca Sospiro and
El Pozo (nearlyvirgin)

Marasmiusleveilleanus Marasmiuschiapasensis
Marasmiuscarpenterianus Marasmiusatroincrustatusvar inodorosus
Marasmiusfloriceps Marasmiusbrunneocinctus
Marasmiushaematocephalus Marasmiusdictoyledoneus
Marasmiusrubroflavus Marasmiusxerampelinus
Marasmiustrinitatis Marasmiuschrysoblepharis
Marasmiusneosessilisvar montepiensis Marasmiushaematocephalus
[Many species of Agaricales,Aphyllophorales Marasmiuslongisporus
and Gasteromycetes,see Singer(to be pub- [Micromphaleseparatum]
lished,none of them observedin area7.1 [Mycenamaculatavar]


While I adhere in principle to the definition and limits of the area covered by
the neotropics as accepted by the Organizationfor Flora Neotropica, it may be useful
to point out here that the Marasmieaehave a mycelium protected by the host tissue
or substratumand the majority of species is found in spots with comparativelyfavor-
able microclimaticconditions. Consequently, many basically tropical species pass over
into the subtropical,a few even in the warm-temperatezone of both northernand
southern hemisphere. Because of such a relatively unalteredand gradualtransition of
the flora, it was believed to be preferableto admit a species to treatment in Flora Neo-
tropica even if it was found only in the subtropicalzone - provided its precise area
of distribution is insufficiently known - i e, occurrence in the intertropicalregion
Introduction 7

proper remains uncertain. However, in such cases a note is added stating that the par-
ticular species is marginalfor the neotropic mycoflora.


In the descriptivedata, colors are, unless otherwise indicated, those of Maerz&

Paul (1930), designatedeither by name given in quotation marksor by number. Other
terms are the ones commonly used in agarictaxonomy. Wheredefinitions or discuss-
ions of the terms are desirable,the reader is referredto Singer (1942, 1958b, 1965,
1974). The data on the types are given briefly only where precise data are indicated
under "Materialstudied." Data on the known distributioncan be found after descrip-
tion of the habitat or else they can be deduced from the material studied. The distrib-
ution can rarely be given in precise and detailed form so that a special paragraphon
distribution is not included.
Many taxa have been illustratedin the paperscited in the literature;these have
been cited at the end of each descriptionand discussionin the descriptivepart. The
originalillustrationsof the present paper correspondto the following magnifications:
carpophoresand carpophoreparts(pileus, etc) X 1/2; spores X 1000; other microscop-
ical details X 500. Exceptions are indicated in the explanation of figures.
If specimens cited in the text have been distributed,divided, or redetermined
under another name, the collector's numberis indicated in the Index of Exsiccatae
Herbariaare quoted accordingto the Index Herbariorum(Holmgren& Keuken
1974) with the correspondingabbreviations.


The present monographhas been preparedand edited with the help of two grants,
GB 7477, GB 35471, from the National Science Foundation, WashingtonD.C.
8 Flora Neotropica


Tricholomataceaetribe MarasmieaeFayod, Prodrome,Ann. Sci. Nat. Bot. VII. 9: 340;

1889 (as Marasm6s).

TYPE GENUS. Marasmius Fries.

Key to the Subtribes of Tribe Marasmieae

1. Pseudoamyloid or incrusted long hairs or hairs with apical flagella-like pseudoamyloid
appendages present on the surface of the pileus. Crinipellinae, p 8.
1. Pseudoamyloid long incrusted hairs and pseudoamyloid flagella-like appendages absent
on the surface of the pileus.
2. Spores rough to spinose or fungi combining the following characters: hymeniform or
subhymeniform epicutis present and elements of the epicutis smooth; hyphae without
clamp connections and inamyloid, or with clamp connections and with spores
broader than 7.5p; pseudorrhiza (georhiza) well developed; stipe central; lamellae well
developed. Oudemansiellinae [not treated here3]
2. Spores always smooth; epicutis not hymeniform or, if hymeniform, spores narrower
than 7.5u and hyphae with clamp connections or else pseudoamyloid; stipe institious
or with mycelial fibrillosity at the base; not or rarely developing a pseudorrhiza;
central, eccentric or absent; lamellae well developed or not.
3. Epicutis consisting of a trichodermial palisade of smooth, parallel or subparallel,
rather broad but elongated hyphal cells with intraparietzl pigment. or a non-
hymeniform layer of diverticulate hyphae and then base of stipe not insititious and
stipe central; hyphae inamyloid; with clamp connections; (see Oudeimansiellinae).
3. Epicutis different or else stipe insititious and/or not central; hyphae inamyloid or
pseudoamyloid; with or without clamp connections. Mlarasmiinae.p 61.

Tribe Marasmieae subtribe Oudemansiellinae Singer, Sydowia 15: 58. 1961 [1962]

Key to the Genera of Subtribe Oudemansiellinae

1. Spores smooth or grossly spinose; epicutis hymeniform or subhymeniform.
2. Clamp connections present. Oudemanisiella.
2. Clamp connections absent. Strobilurus.
1. Spores rough-asperulate or verruculose, more rarely smooth or spinulose; epicutis not
hymeniform but often with numerous erect, conspicuous dermatocystidia; elements of
the epicutis often diverticulate or else epicutis in form of a trichodermial palisade.
3. Epicutis not a trichodermial palisade. Mvscenella.
3. Epicutis a trichodermial palisade. PhYIsocYstidium.

Tribe Marasmieae subtribe Crinipellinae Singer, subtrib nov

Pileo crinibus plerumque pseudoamyloideis et crasse tunicatis, rarius inamyloideis
et incrustatis, semper longis dense tecto aut elementis externis epicuticularibus appendi-
culis pseudoamyloideis flagelliformibus praevisis. Hymenio lamelloso vel absente. Stipite
centrali vel excentrico aut pseudostipite praesente, rarius absente, et tunc carpophoris
cyphelliformibus. Hyphis inamyloideis.
Pileus densely covered with pseudoamyloid, thick-walled hairs, or rarely with
inamyloid and incrusted hairs, these hairs always long and unicellular or with secondary
septa, or else the external epicuticular elements covered with pseudoamyloid appendages.

3See Singer, R., (1964a and 1973).

Systematic Treatment 9

Stipe central or eccentric or else a pseudostipe present; carpophores,if neither a true

stipe nor a stipe rudiment present, often with smooth hymenial surface, in this case

TYPE GENUS.CrinipellisPatouillard.

Key to the Generaof SubtribeCrinipellinae

1. Long, conspicuous, mostly pseudoamyloid or else incrusted and thick-walled hairs
2. Stipe present, central or eccentric. 1. Crinipellis, p 9.
2. Stipe absent or rudimentary (functionless).
3. Lamellae present. 2. Chaetocalathus, p 153.
3. Lamellae absent, (hymenial surface smooth).
4. Hairs distinctly pseudoamyloid, thick-walled, long. 3. Lachnella, p. 58.
4. Hairs inamyloid or faintly pseudoamyloid, often thin-walled in part or entirely
but mostly incrusted. 5. Flagelloscypha, p. 60.
1. Long, conspicuous, pseudoamyloid or incrusted hairs absent, but the outermost elements
of the epicutis beset with pseudoamyloid flagella-like appendages. 4. Amyloflagellula, p 60.

1. CrinipellisPatouillard,Jour. de Bot. 3: 336. 1889.

TYPESPECIES. C. stipitaria(Fries) Patouillard.
Habit of the carpophorescollybioid, marasmioid,or rarely pleurotoid, but always
with a well developed true stipe: pileus and stipe covered with thick-walledhair-like
elements whose wall is partly or entirely pseudoamyloid;these hairs long and without
clamped septa, cyanophilous, rarely and slightly incrusted, most frequently without
incrustationand smooth. Hymenophorealways well developed, lamellate, lamellae
most frequently free or subfree;cheilocystidia present;cystidia on the sides of the
lamellae present or absent, not pseudoamyloid;spore print pure white or nearly so;
spores hyaline, 5-20 p long narrow to broad, thin- to thick-walledand eventually some-
times septate, inamyloid, more rarely pseudoamyloid,cyanophilous, smooth; basidia
without sideropliilousgranulosity,generally4-spored, sometimes somewhat deformed
(basidioles not sporulating);basidioles fusoid, rarely (in species with very voluminous
spores and basidia) more clavate-subcylindrical; stipe insititious, rarelywith a distinct
basal mycelium which is fibrillose or tomentose, or with white rhizomorphs;trama
inamyloid, clamp connections present. Saprophyticand parasiticon plants and plant
debris, stems, roots, fruits, woody material,etc.
Development of the carpophores:unknown; the youngest carpophoresseen have
the hymenial primordiumcovered by the hairs of the marginof the pileus and the
apex of the stipe.

Key to the Sections and Subsectionsof Crinipellis

1. Pileus purple, violet, lilac, crimson red etc., not changing to green or olive when treated
with NaOH or KOH (dried material) and hairs not greenish or greenish melleous in 5%
KOH. sect 4. Iopodinae, p 43.
2. Stipe elongated (more than three times as long as the diameter of the pileus), usually
straight, central, insititious; 20-26 mm long; spores fusoid to oblong and not over 4.7
p broad, or else ellipsoid and then not more than 4, broad (Q = about 2, or larger).
subsect Iopodinae, p 43.
2. Stipe different; spores as above or different. subsect Insignes, p 45.
1. Pileus pink or some shade of stramineous to brown, or pigment-less, frequently chestnut
to ferruginous-brown in wet, and more dull stramineous-tan in fresh-dry (faded) cond-
ition: if the pileus pink, turning green with a drop of 5% KOH.
10 Flora Neotropica

3. Pileus with KOH turning green or (green-) black and in the latter case the hairs and/or
the hypotrichium pale greenish or greenish-melleous or greenish gray rather than pale
ochraceous-brownish to dull ocher or hyaline; (pleuro-) cystidia usually present,
scarce to numerous. sect 3. Grisentinae, p 37.
3. Pileus with KOH merely turning darker (in the same color as found in the dried
pileus, often reverting to the color of the fresh wet specimens), or not reacting; hairs
of the pileus in 5% KOH pale ochraceous brownish to dull ocher, or hyaline; (pleuro-)
cystidia present or absent.
4. Stipe at first central but soon becoming eccentric, mostly short and curved, 2-7
mm long; pileus (at first) white or whitish (excepting sometimes the center); the
great majority of the spores with Q<2,< 10.5p long. sect 2. Excentricinae, p 34.
4. If the stipe less than 7 mm long, strictly central and straight and not white, but
more frequently the stipe longer than 7 mm and then also usually central, more or
less straight, more rarely inconstantly slightly eccentric or curved when mature; if
stipe eccentric, spores as above or different. sect I. Crinipellis, p 10.
5. Spores more than 9,u broad, subglobose. subsect Macrosphacrigerae, p 10.
5. Spores less than 9u broad.
6. Cheilocystidia present and mostly crowded, but pleurocystidia very scarce
and inconstant and mostly, if at all present, in form of cheilocystidia
slightly removed from the edge, rarely pleurocystidia well differentiated in
old carpophores, but lacking in younger ones. subsect Stipitarinac, p 11.
6. Cheilocystidia and pleurocystidia both differentiated, the pleurocystidia
usually different from the cheilocystidia. at least many of them.
subsect Heteromorphinae, p 32.

Crinipellissect Crinipellis

Pileus not brightly colored and not turning green or green-blackwith KOH or NaOH,
and the hairs not greenish or greenish-melleousin 5%KOH;stipe generally central and
pigmented, insititious. The type is that of the genus.

Crinipellissect Crinipellissubsect MacrosphaerigeraeSinger, subsect nov

Sporis subglobosis, 9 u latis vel latioribus.
TYPE SPECIES.C. macrosphaerigera Singer

1. Crinipellis macrosphaerigera Singer, Lilloa 26: 138. 1953.

TYPE. Sehnem 1917, from Brazil (LIL).
Pileus ochraceous straw-color with brownish shade when dry, finely pilose, with
appressed hairs, with a punctiform glabrous central dot, there black, convex later repand
8-16 mm broad. Lamellae (white?), dried almost concolorous with the pileus, medium
broad, distant, narrowly adnexed. - Stipe concolorous with the pileus, pilose, central,
equal, 10-25 X 1-1.3 mm. Context white.
Spores 12.3-13.8 X 9.5-11.7g , subglobose (varying from globose to very short
ellipsoid) with moderately thin wall which has a tendency to collapse whereby the
spores become seemingly angular, inamyloid, smooth. Hymenium: basidia 46-72 X
13.7- 16.5 u, 4-spored; basidioles narrowly clavate-subcylindrical; cystidia absent in
young carpophores but in old ones some scattered cystidioles are often present, these
e g 35 X 9.5u, ventricose, attenuate at the apex, hyaline; cheilocystidia 34-71 p long, of
variable diameter, dendroid, ramose either at the apex only, or else from the middle
portion upwards, branches erect, two to many, acute or more frequently cylindrical
with rounded tip, 4-15 p long, making the edge of the lamellae heteromorphous, or
subheteromorphous (if intermixed with a few basidioles and basidia). - Hyphae hyaline,
inamyloid, with clamp connections. Cortical layers: hairs of the margin of the pileus
4.27 p1broad, some hyaline or some brownish-melleous, strongly and entirely pseudoamy-
Crinipellis 11

loid, with the wall 1.4-2.8 , thick wall, not secondarilyseptate, very long, with rounded
On living tree in subtropicalforest
MATERIALSTUDIED. Known only from the type locality: BRAZIL. Rio Grande
do Sul: Estacao Sao Salvador, A. Sehnem 1917, 21 Jan 1946 (LIL: C 8339, type).

Crinipellissect Crinipellissubsect StipitarinaeSinger, Lilloa 8: 461. 1942.

Spores much smalleror much more elongated than in subsectionMacrosphaer-
igerae;(pleuro-) cystidia absent, or present only as cystidioles in old carpophores(or
where the lamellae have been bruisedin situ) or as cheilocystidia(scattered at a short
distance from the extreme edge); if as cystidioles, these are degeneratedbasidioles.
TYPESPECIES. C. stipitaria (Fries) Patouillard.

Key to the Species of Subsection Stipitarinae

1. Spores9-14.5(-20) X 3.5-6.3,u,often curvedQ 2.5-3.2, pileus up to 10 mm broad;stipe
more than three times as long as the diameter of the pileus. 2. carecomoeis.
1. Spores up to 13.5,/ long in 4-spored specimens, not curved, Q as above or more often
smaller; pileus 1-45 mm (mostly 1-35 mm) broad; stipe as above or relatively shorter.
2. Spores, at least many of them, reaching more than 9.5g, up to 13.5,u long; Q =
1.7-2.2. Growing mostly on monocotyledonous debris, dead culms, roots, leaves of
grasses, Araceae etc., often in xerophytic conditions.
3. Pileus 5-35 mm broad, obtuse or obtusely umbonate, but not papillate either in
youth or at maturity; stipe up to three times longer than the diameter of the
pileus; lamellae subdistant or distant; most hairs of the pileus either narrowed
obtuse or rounded at the tip.
4. Pileus 6-13 mm broad; spores (7-)8-11 X 3.5-5.5JL;cheilocystidia usually with
1-5 apicalfinger-likeappendages;rarelyentirepileuswhitish,dryingto gilvous
Tropical 4. C. urbica.
4. Pileus 8-25(-35) mm broad: spores 8.7-13 X 4-6.2p; cheilocystidia entire or
forked or with short mostly lateral knobs or outgrowths; pileus grayish, ochra-
ceous to cinnamon brown. 5. C. subtomentosa.
3. Pileus 1-10 mm broad, either with a glabrous central papilla or if not then most
stipe usually longer than three times the pileus diameter hairs of the pileus with
acute tips; lamellae close or subclose.
5. Pileus 1-2 mm broad, white with cinnamon ochraceous small disc which is
papillate and glabrous and naked. On large leaves of monocotyledonous forest
plants (Araceae). 3. C. gracilis.
5. Pileus 4-10 mm broad, brown ("spruce y" M&P ), obtuse. On grass. 6. C. megalospora.
2. Spores, at least in species growing on monotyledonous hosts and in xerophytic vege-
tation, not longer than 10.5p; Q variable.
6. On conifers(Podocarpus) 16. C. podocarpi.
6. On angiosperms.
7. On Monocotyledones, mostly Gramineae (but also other families).
8. Pileus 1-2-(3) mm broad; spores broad: 8-9.5 X 6-7.5,. Mostly on Bam-
buseae in Brazil (adventitious). 7. C. perpusilla.
8. Pileus larger; spores different.
9. Lamellae cream when fresh, close; spores 5.5-7.5 X 3.5-4.5g. Subtropi-
cal-montane, South-American. 10. C. catamarcensis.
9. Lamellae not cream color, or spores different.
10. Pileus 9-17 mm broad, "cattail," later somewhat pallescent; spores
broader than 4,. 12. C. atrobrunnea.
10. Pileus either smaller than 10 mm or spores narrower than 4,; pileus
rusty when fresh and wet, then pallescent but not particularly in
the center.
11. Pileus papillate or with distinct concentric walls and furrows
around the umbilicus; spores 6.3-8.5 X 3.1-3.8,; majority of
the hairs in the middle zone of the pileus with acute or sub-
acute tips, minority with obtuse tips. 8. C. bisulcata.

4M&P = Maerz & Paul (1930) - here as well as in all following keys and descriptions.
12 FloraNeotropica

11. Pileus with or without a small papilla or umbo, and not or in-
constantly and weakly concentrically walled-furrowed.
12. Spores 7-10 X 3.7-6p, Q 1.8-2. C. pseudostipitaria var rzesites.
12. Spores 7.7-10 X 3.5-4.5ui Q> 2.
9 C. pseudostipitariavarpseudostipitaria.
7. On dictoyledonous hosts.
13. Pileus about 40 mm broad; spores 7.5-9.3 X 5.4-6.8u; center of pileus less
or not pilose; surface more or less zoned. 13. C. missionenesis.
13. Pileus smaller, rarely reaching more than 20 mm in diameter; otherwise
as above or different.
14. Spores larger than 6.3 X 4.2p.
15. Spores up to 5.51 broad (or less broad); lamellae white or whitish
when fresh.
16. Growing on wood, woody sticks, branches or roots; cheilo-
cystidia appendiculate or not.
17. Hairs almost constantly with close secondary septa at least
in the great majority of the apical portions of the hairs of
the pileus (structure ladder-like), often with acute tips,
with walls 1.5-3, thick; cheilocystidia not crowned by
apical finger-like or thorn-like appendages, but often with
1-5 finger-like lateral and/or apical excrescenices; spores
6.5-12 X 3.5-6u, Q> 2 (rarely slightly < 2). 15. C. septotricha.
17. Hairs different, or spores and cheilocystidia different.
18. Pileus 9-19 mm broad, obtuse with depressed center;
stipe 8-16 mm long (i e about as long or not more than
twice as long as the diameter of the pileus), spores
(7-)9-10 X 4.5-5.3 p. Hygrophilous. 14. C. herrerae.
18. Pileus smaller or stipe (relatively) longer; spores as above
or different.
19. Cheilocystidia crowned with few to numerous apical
finger-like appendages or setulae, or with conspicuous
side branches or forked in lower portion, rarely a few
20. Pileus up to 7 mm broad; lamellae + lamellulae
12-20; spores 5.3u broad or broader. 17. C. stupparia.
20. Pileus somewhat larger; lamellae somewhat more
numerous; spores less than 5.4 p broad.
21. Spores (3-)4-5. broad.
22. Cystidia none. Hygrophilous species. 18. C. coroicae.
22. Pleprocystidia present; subxerophytic
species. 24. C. coinmixta.
21. Spores 2.7-3.5u broad. 19. C tenuipilosa.
19. Cheilocystidia entire or apically bifurcate, pileus
smaller than 6 mm, lamellae moderately close to sub-
distant, spores very variable: 7-10 X 3-6.5,. 20. C. mexicana.
16. Growing on fallen leaves or leaf petioles of dicotyledonous trees
and shrubs; at least some cheilocystidia with several apical ap-
23. Pileus 3-7mm broad; spores with a Q > 2 or = 2 (i e, spores
ellipsoid to oblong) tips of the hairs of the pileus acute
and rounded (both types present on the marginal zone of
the pileus); cheilocystidia generally short: 8-23p long. On
fallen leaves only. 21. C. foliicola.
23. Either pileus somewhat larger or spores with a Q < 2 tips of
the hairs on the margin of the pileus either all or almost
all obtuse or all or almost all acute; cheilocystidia 15-30
,v or longer. On fallen leaves and woody material.
24. Pileus 6-11 mm broad; hairs of pileus acute or subacute;
papilla none, or sometimes present and rather low;
spores (8-)9.3-10.5(-11.2) X (3.7-)4.2-5,4. On oak leaves.
22. C. phyllophila.
24. Pileus smaller or spores relatively broader. On leaves (ex-
cept oak) and woody material, (see "17" above).
Crinipellis 13

15. Spores often broaderthan 5.5,; lamellaewhite or creamto buff

when fresh.
25. Cheilocystidiawith mostly apical and rathernumerousappen-
dages;pileus not broaderthan 6 mm when dried(if the stipe is
central, see "17" above; if the stipe is eccentric, see section
25. Cheilocystidiaoften branchedbut not only apically;pileus as
aboveor larger.
26. Pileus small (to 6 mm), more tomentose and eventually
somewhat glabrescent around the papilla or glabrous on
disc; spores 6-9.8 X 4-7, (Q = 1.4-1.8 rarelyup to 2) (see
"19" above); if spores9-11 X 6-7.5u and pileus somewhat
largerat timies,compare. C. molfinoanap25.
26. Pileus about 18 mm broad, rather evenly pilose; spores
8.5-9.6 X 5.5-6.2u 23. C. dicotyledonum.
14. Sporesup to 6.3 X 4.2tp.not larger. 11. C schini.

Stirps Carecomoeis

What may be called stirps Carecomoeisis the group of obviously related species
no. 2-4 of this monograph.

2. Crinipelliscarecomoeis(Berkeley&Curtis) Singer, Lilloa 8: 487. 1942. Fig 1.

AgaricuscarecoimoeisBerkeley& Curtis,Jour. Linn. Soc. Bot. 10: 284. 1868.
Collybiacarecozmoeis(Berkeley& Curtis)Saccardo,Sylloge Fungorum5: 232. 1887.
TYPES:Wright,Cuba (FH, K)
Pileus ochraceous brown to rusty brown in the center when fresh, marginuniform-
ly white or white-rimosebetween curly strandsof hairs which are concolorous with the
center, radially plicate-pectinate-pilose,center tomentose-piloseand the hairs here also
mostly appressed,but less strongly pilose in the center than on the margin,with a few,
0.5 mm long brown bristles often present on the papilla, fading to "cinnamon buff" or
"tawny olive" (Ridgway) when dried, conic-campanulate,convex-campanulate,or con-
vex and papillate, often as high as broad, more often with than without a papilla, with
or without a concentric wall around the papilla, with fimbriate marginin most caps,
with a flat-bottom umbilicus if not papillate, 4-8.5 mm broad, 2.5-8 mm high. - Lam-
ellae white, dried at times assuminga pale orange buff shade, rather narrowto broad,
at first ascendant or subascendant,later horizontal or almost so, often ventricose, close
or subclose, free or subfree. Stipe white or pallid at apex, almost chestnut brown to
ferrugineousbelow, but fading to whitish to dirty light brown or pale argillaceousand
eventually unicolorously so, appressedlypilose all over excepting the base where the
rusty hairs are erect and hirsute when fresh, equal, hollow, insititious, 25-92 X 0.2-1
mm. Context white, thin, inodorous.
Spores (9-)13-14.5(-20) X (3.5-)4-5(-6.3) cylindric to subfusoid in frontal view,
frequently or mostly somewhat curved with a depressionon the middle portion of the
iner side or even sausage-shaped,hyaline, thin-walled,unicellular,overagedspores most-
ly yellowish, thick-walled,and sometimes with a cross-wallnear the hilar end, inamyloid,
sometimes a few slightly pseudoamyloid, smooth, cyanophilous. Hymenium:basidia
18-26 X 5.5-8 1, 4-spored, or 2-4 spored; basidiolesfusoid-subclavate;cystidia none, but
basidioles in old specimens often remainingsterile and transformedinto cystidioles which
are ampullaceousto ventricose-mucronate,29-33 X 10-14.5,, hyaline, thin-walled;
cheilocystidia numerous, 20-35 X 4-12 M,ampullaceous,subampullaceous,or ventricose
to ventricose-subclavate,not branchednor crowned with finger-likeappendagesor setae,
but sometimes with very short knotty outgrowths on the sides or a short obtuse forking
14 Flora Neotropica

or a mucro at the tip, thin-walledto very slightly thick-walled,or wall locally somewhat
thickened (to 1.5 u), hyaline to melleous-hyaline. Hyphae of the regularhymenophoral
tramawith clamp connections, hyaline, inamyloid. Coveringlayers: hairs of the pileus
3.5-11 p broad, with acute or rounded tips, with stramineousto melleous or (near mar-
gin also) hyaline 0.5-5.5 u thick (in KOH or NH40H), strongly to moderately strongly
(if strongly - purple) pseudoamyloid, some hairs with abrupt thickenings,all very long,
often bundled together with the longest hairs in the center of the strands,some hairs
with secondary septa but these scattered (and not ladder-like,crowded). Hypotrichium
of multi-septatehyphae which are broaderthan those of the trama proper (5-8 ,ubroad),
with hyaline, rather thin walls. Hairs of stipe much like the hairs of the marginof the
On fallen leaves of trees, on dead grasses(e g Paspalum),on dead herbaceous
leaves and stems (Dicotyledones), also in humus among Lycopodiales. solitary or greg-
arious, fruiting from April until August.
MATERIAL STUDIED. CUBA, Wright, both type and authentic material (FH). MEXICO.
Chiapas: between Ixtocomitan and Solusuchiapa, 1 Aug 1969, Singer M 8805 (F), il 8811 (F).
COLOMBIA. Valle: Buenaventura, San Joaquin, 19 Apr 1968, Singer B 6230 (F).
In the neotropics we find only var carecomoeis. The varietiessubelata and
litseae which may also be separatespecies of the same stirps are confined to the South
Pacific (Philippines,New Caledonia);they grow on fallen leathery leaves.
The collection Singer M 8805 is the only one without clamp connections, with
only bisporous basidia seen and with numerousand conspicuous cystidioles on the sides
of the older lamellae. This may be a bisporous-parthenogeneticform of the same spec-
ies, or possibly a related species; it was also the only one on Gramineae. The charac-
ters of this form have not been included in the general description of the species, but
the specimen is cited under "Materialstudied."

3. Crinipellisgracilis Singer, sp nov

TYPE.Singer B 3412 (F), from Pernambuco,Brazil
Pileo albo, centro cinnamomeo-ochraceo,minute pilosulo, centro subglabro,cam-
panulato-convexo,demum applanato, centro papillato, papilla interdum umbilico cen-
trali inserta, 1-2 mm lato. Lamellis candidis, subangustis,subarcuatis,confertis, liberis
vel subliberis. Stipite albo, mox atrocastaneoad basin, in siccis ex integro cinnamomeo,
albo-piloso, insititio, aequali, 7-8 X 0.1 mm. Contexto tenuissimo, in pileo albo, inodoro.
Sporis 9-13.5 X 5-6.5 (-7) ,, plerumque 10-11.5 X 5.5-6.5 u, subfusoideis,hyalinis,
inamyloideis, tenui-tunicatis,demum crasse tunicatis et nonnullis prope apicem septatis,
levibus. Hymenio: basidiis et basidiolis normalibus;cystidiis subnullisvel nullis; cheilo-
cystidiis subventricosis,excrescentiis nonnullis brevibusnodosis apicalibus instructis,
hyalinis. Hyphis in stipite parallelis,filamentosis, crasse tunicatis, brunneolo-tunicatis,
inamyloideis. Tegumentis: crinibuspilei hyalinis, crasse tunicatis, longis, 4-5 (-9)p latis,
ad apicem acutis vel rotundatis, saepe ad apicem tenuitunicatis,pseudoamyloideis,
parietibus 1.5-3 (-3.5)u crassis,septo secundariosubapicaliinterdum praesente. Crinibus
stipitis brevibusvel longis (brevibus subulatis), crasse tunicatis (parietibus 2-2.8 p crassis),
7-8.5 p diametro, omnibus vel plurimisacutis.
KOH in pileo sicco colorem haud mutat.
Ad folia Monocotyledonum (probabiliterAracearum),gregatim,
Pileus white with cinnamon-ochraceouscenter, minutely pilose with appressed,
somewhat curly hairs except for the naked papilla, but at the marginhairs projecting
so strongly that in young and dried specimens the lamellae may appear completely
Crinipellis 15

covered by them, convex, eventually flattened and often with umbilicus, with a constant
central small papilla, the latter often inserted in the umbilicus when carpophoresmature,
1-2 mm broad. Lamellae pure white, rather narrow, subarcuate,close, free or subfree.
Stipe white, soon becoming chestnut-fuliginousfrom the base upwardsand dried entire-
ly cinnamon, white-pilose, straight, insititious, equal, 7-8 X 0.1 mm. Context very thin,
in pileus white; odor none.
Spores 9-13.5 X 5-6.5(-7)p, mostly 10-11.5 X 5.5-6.5 A,subfusoid, hyaline,
smooth, thin-walled,becoming thick-walledwhen overmature,some with a septum
near the apex when overmature,inamyloid. Hymenium: basidia 18 X 7-8.8 u;
basidioles fusoid or subfusoid;cystidia few (like cheilocystidia) or none on the sides
of the lamellae; cheilocystidia subventricose,with some knob-like short excrescencies
at the apex, hyaline, few, size of basidia. Hyphae of the stipe parallelwith each
other, filamentous, thick-walled,brown-walled,inamyloid. Coveringlayers: epicutis
of pileus consisting of hairs, these very long, 4-5(-9) broad, hyaline, thick-walled,
rounded or acute at the tip which is often thin-walled,pseudoamyloid,wall 1.5-3(3.5)
; thick, some hairs with a subapicalsecondary septum. Hairs of the stipe short to
long (the short ones subulate), thick-walled(wall 2-2.81 thick), 7-8.5 u in diameter,
all or most with acute tip.
KOH on pileus negative.
On dead leaf of a monocotyledonous plant, probably Araceae.
MATERIALSTUDIED. BRAZIL.Pernambuco:Mun.Sao Lourenco,Matado Sao, Joao,
16 Jul 1960, SingerB 3412 (F, typus).
This is similar to, though still smaller than C. careconzoeis,but has slightly
smallerspores, less pigmentationespecially in youth, and a glabrouspapilla;it grows
on monocotyledonous dead leaves in the tropical forest.

4. Crinipellisurbica (Montagne)Dennis, Kew Bull. 1951: 409. 1951. Fig 2.

Agaricusurbicus Montagne,Ann. Sci. Nat. 1: 94. 1854.
Collibia urbica (Montagne)Saccardo,Sylloge Fungorum5: 218. 1887.
TYPE.Leprieur989, from French Guyana.
Pileus whitish when fresh, on drying somewhat gilvous (Hebelona-sinapizans
color), hirsute especially in the center or villous all over from minute pallid hairs, not
distinctly zonate, radiallysulcate, convex, soon with depressedcenter or entirely flat,
often obtusely umbonate, but without a distinct papilla, 6-13 mm broad. Lamellae
white, becoming somewhat brownish when dried, subdistant, with about 6 lamellulae,
broad, narrowly adnexed or subfree. Stipe brownish or argillaceous,pallid-hirsute,
straight,stuffed, insititious, equal, 17-30 X 1 mm. Context white.
Spores (7-)8-10(-11) X (3.5-)4-5(-5.5) u, with Q = + 2, ellipsoid, hyaline, smooth,
inamyloid. Hymenium: basidia 22-30 X 6-7 u; cystidia not seen; cheilocystidia21-30
X 6-8.5 ,. either entire, or more often with apical appendages,these finger-likeand
obtuse, 3-7 X 1.3-3 4, all hyaline, main body ventricose to cylindric-subventricose.
Hyphae of tramawith clamp connections, inamyloid. Coveringlayers: hairs of the
middle of the pileus mostly moderately long (to 270 u), 4-6.5 , broad, walls 1.5-2.5 p
thick, pseudoamyloid and with rounded or attenuate-obtusetips.
On sandy soil among grass,often on grassroots, often in dense groups.
Rio de Janeiro:Cabo Frias, Caqueiral,25 Jan 1967, Jezek 4 (BAFC).
This species has relatively narrowerspores, less pigmented pileus, less acute hairs
on the pileus, fewer simple and entire cheilocystidia and relatively shorter stipes in an
16 Flora Neotropica

averageof a population and a strictly neotropical distribution, than C. stipitaria. It

differs from C. pseudostipitariavar mesites in less pigmented surfaces,less close lam-
ellae, slightly larger spores and the absence of a papilla.

5. Crinipellissubtomentosa (Peck) Singer, Lilloa 8: 463. 1942.

Marasmius subtomentosus Peck, BulL Torrey Club 22: 487. 1895.
Marasmiuzstomentosuis Peck in Ellis Everhart, North American lFungi II 3403. 1903 (non
Marasmiiusdasypuis Spegazzini, Anal. Mus. Buenos Aires 19: 264. 1909.
Marasmius nolaneiformis Murrill, Bull. Torrey Club 67: 149. 1940.
TYPE. Bartholomew, KansasFungi 1735 (FH).
Pileus grayish ochraceous("walnut taffy" or "desert" M&P)to cinnamon brown
("Arab" with "stone" center, or with darker"Hindu"center and "leather br." around
it), becoming grayer and isabelline when fading or dried ( 11 C 4 with 11 D 6 center,
also paler, especially where cracked or glabrescent,or more grayish), subtomentose-
fibrillose from a hoary but thin felt-like covering, but becoming rivulose cracked over
much of the surface in older caps, often appearingzonate, often becoming coarsely
sulcate over the marginalportion, otherwise smooth, neither viscid nor radiallylong-
pilose, opaque, campanulate or convex, then applanateand often with depressed
center, or with a low, obtuse umbo, 8-25(-35) mm broad. Lamellaepale cream to
grayish ochraceous or sordid flesh color ("ahnong", "aloma", even reaching"army br.",
"tawny, birch" or between "fawn" and "Sandalwood" when old), not intervenosebut
often forked, irregularlyintermixed, moderately broad (to 2.7 mm), distant or sub-
distant, rounded and very narrowlyattached and eventually separating,or subfree to
quite free. Spore print pure white. Stipe concolorous with the pileus but when fresh
often more fulvous ("sorrell"M&P)sometimes paler or on the contrary, fuliginous-
fuscous when dried, hoary, fibrillose-pilose-rough,solid, or stuffed equal but mostly
with a basal bulb, immersedportion (if there is one) often brusquely broader, shorter
than three times as long as the pileus-diameter,25-55 X 1-3.5 mm (immersed portion
up to 6.5 mm broad in some specimens, bulb 3-4 mm in diameter);basal mycelium
none, or very scarce. Context white, fresh with a watery line above the lamellae, in-
odorous, mild to the taste.
Spores (8.7-)10-12.2(-13) X (4-)5.5-6(-6.2) , but on bisporous basidia 8.7-16
X 5.7-7p1,ellipsoid (Q variable,= 1.8-2.2), smooth, hyaline, inamyloid, thin-walled.
Hymeninum: basidia 1-2-3-4-spored,often the majority 2-spored, 31-40 X 7.5-12.3 P,
the 1-3 spored ones narrowerthan the 4-spored ones; cystidia none (or some cheilo-
cystidia somewhat away from edge); cheilocystidia 15-40 X 3.5-9 P, entire or forked
in upper portion (and then both forks cylindric or hook-like or club-shaped),besides
often with lateral knobs or hooks, but these always short, subcylindricto clavate. more
rarely swollen below, often irregular,bent or crooked, hyaline. Hyphae of the trama
hyaline, with clamp connections (even in the bisporous forms), inamyloid, not gelatin-
ous, thin-walledor in old carpophorespartly somewhat thick-walled;hymenophoral
trama regular,of filamentous, slightly interwoven hyphae; subhymeniumof small ele-
ments, some rectangularand in chains, some more elongated. Coveringlayers: hairs
of the pileus not very long in comparisonwith related species, usually not longer than
about 200 pu,somewhat irregularin diameter while others are equal, 3.5-7.5 p broad,
with walls strongly pseudoamyloid, 1.3-2.8 u thick, apex usually tapered and thinner-
walled but tips obtuse to rounded, rarely acute, many hyaline, others stramineousin
KOH;hypotrichium stramineous. Hairsof stipe similar.
On old grassroots and sometimes passing over to other plant debris, often sub-
Crinipellis 17

merged ones in sandy soil or earth, gregarious,fruiting from Januaryuntil July, in

North America perhapsalso later in the year. Always in open grass land or in semi-
xerophytic vegetation under scattered shrubs.
MATERIAL STUDIED. U.S.A.: Kansas: Rockport, Bartholomew, 1 Jul 1885; type,also
later collections (FH). Florida: Alachua Co., Gainesville, on open lawns, 31 May 1938, Murrill
(FLAS). ARGENTINA. Jujuy: Ledesma, Spegazzini, Jan 1906 (LPS); Tucuman: Capital, Parque
Urquiza, 8 Jan 1949, Singer (LIL). FRENCH EQUATORIAL AFRICA. Loango: Dybovsky,
1891, det. Patouillard (C. stipitaria) (FH).
The existence of the species in Africa might suggest that C. mauretanicaMaire
(1928) is the same species, but this has longer hairs, intervenoselamellae, filiform
cheilocystidia and grows on "buried plant debris" in Africa. It is A. chortophilus
ILLUSTRATION.Singer (1942) fig 2 d.

6. CrinipellismegalosporaSinger in Singer& Digilio, Liloa 25: 222. 1952. [1953]

Type. Singer T 1666 from Argentina(LIL, BAFC, MICH). Fig 3.
Pileus yellow brown ("spruce y" M&P)with deep fuliginous center, paler at the
marginbut not white, when faded "spruce yellow" appressedlyhairy on whitish ground
with the disc "camels hair," appressedlypilose all over and in dried condition the white
backgroundshowing between the hairs, convex with applanateto depresseddisc around
which there usually is a circularfurrow, not papillate, but marginstrongly declivitous,
4-10 mm broad, mostly 5-7 mm. Lamellaewhite or whitish, rathernarrow,close, sub-
free. Stipe ochraceous brown, light cinnamon or light grayishcinnamon when dry-faded,
grayishcinnamon near base, insititious, equal or subequal 25-30 X 0.6-1 mm, mostly
0.6-0.8 mm broad. Context white, inodorous.
Spores 9.5-11(-12.5) X (4.8-)5.5(-6.5) p, with Q = 1.8-2.3, ellipsoid to subfusoid
or oblong, hyaline, smooth, thin-walled,over-agedspores thick-walledand stramineous,
inamyloid. Hymenium: basidia 27-28 X 8-8.5 E,,4-spored;basidioles fusoid; cystidia
none excepting some cheilocystidia occasionally seen away from the edge; cheilocystidia
very numerous(always mixed with a few basidiaor basidioles), 19-50 X 4-8.3 u, or
broaderwhere branched,ventricose to clavate and with forked apex or with up to 5
apical finger-likeappendages,forks and main body also sometimes with lateral knobs
or short branchesor bulges, hyaline. Hyphae with clamp connections, hyaline, inamy-
loid. Coveringlayers: hairs of the pileus long, mostly more than 300 ,, 4.5-8.2 , broad,
with walls 1-4 1 thick, strongly pseudoamyloid, with tips needle-sharp,acute, in a minor-
ity attenuate-obtuseor even rounded, with or without cross septa or at least one cross-
septum underneaththe apex, but not consistently nor distinctly ladder-like,in KOH
melleous to hyaline, some fulvous in ammonia, those of the extreme marginsometimes
with some hook-like outgrowths or irregularities,otherwise those of the marginlike
those of the center. Hairsof the stipe similar.
On the base of tufts of grassand on grass roots in open places. South America,
1000-1200 m alt. Growinggregariously. Fruitingin the rainy season.
MATERIAL STUDIED. ARGENTINA. Tucuman, Anta Muerta, 26 Dec 1951, Singer T 1666
(LIL), holotype.

7. Crinipellisperpusilla(Spegazzini) Singer, Lilloa 25: 494. 1952.

Lentinus perpusillus Spegazzini, Bol. Acad. Nac. Ci. Cordoba 11: 406. 1889.
Crinipellis bamnbusaePatouillard, Bull. Soc. Mycol. France 24: 8. 1908.
Marasmius bambusae (Patouillard) Saccardo & Trotter, Sylloge Fungorum 21: 113. 1912.
Type. Puiggari1644, from Brazil.
Pileus whitish to pale tan or subrufescent,densely covered with minute almost
18 Flora Neotropica

silky appressedhairs, with smooth or somewhat sulcate margin,convex then applanate,

often umbonate, 1-2 mm broad. Lamellaewhite, subdistant, narrowlyadnexed. Stipe
whitish, faintly pilose-silky, but darkerin age where the hairs have been scraped off,
equal, central, 2-5 X 0.2-0.5 mm. Context white, very thin.
Spores 8-9.5 X 6-7.5 p1,short ellipsoid, thin-walledand hyaline, becoming some-
what thick-walled,inamyloid, smooth. Hymenium: basidia 35 X 7.5-8 ,, 2-4-spored;
cystidia not seen; cheilocystidia 27-35 X 5.5-10.5 p, rarely simple and entire, mostly
with 2-4 finger-likeapical branchesor appendagesor with short projections giving them
the profile of a fir tree, but often forked (branchesor appendages7.5-17.5 j, long).
Hyphae inamyloid, with clamp connections, hyaline. Coveringlayers: hairs of the
pileus (3-5)7 s, in diameter, thick-walled,with taperingbut obtuse apex, more rarely
broadly rounded at the tip, pseudoamyloid.
On Bambuseae,gregarious.
MATERIALSTUDIED. BRAZIL: Sao Paulo, Campinas,Noack 34 (FH) (type of C. bam-
busae). Apiai, Oct 1881 (LPS), type.
In this section, C. perpusillaand C gracilis are the smallest species of the Mono-
cotyledones-inhabitingforms, but they are easily distinguishablefrom each other, by
the size and shape of the spores.
ILLUSTRATION.Singer (1942) fig 2 e.

8. Crinipellisbisulcata (Patouillard& Gaillard)Patouillard,Jour. de Bot. 3: 336.

Collybia bisulcataPatouillard& Gaillard,Bull. Soc. Mycol. France4: 14. 1888.
Type. Gaillard136 (FH), from Venezuela.
Pileus when dried (fresh color not known) grayish straw color or very pale gray-
ish brown or between "tilleul buff" and "avellaneous"(Ridgway), with the disc "wood
brown" to almost black, apparentlythe deeper the umbilicus the darkerthe color of
the disc, strongly hairy except at the almost naked center, or merely less hairy if the
umbilicus is shallow, subglobose to convex but later flattened, with a very small papilla
in the umbilicus or with a shallow furrow around the papilla and with one or two add-
itional concentric furrows further outwards, 8-13 mm broad. Lamellaeinitially prob-
ably white but concolorous with the marginof the pileus when dried, moderately broad,
medium distant to rather distant, almost free or very narrowly adnexed. Stipe concolor-
ous with the pileus, strongly hairy, stuffed or more or less hollow, 12-55 X 1-2 mm.
Context whitish or white, but with a brown periphericline in the dried material.
Spores (6.3-)7-8.5 X 3-3.8 u Q=?2, hyaline, smooth, inamyloid. Hymenium:bas-
idia 35-41 X 6.5-8 ,; cystidia none seen; cheilocystidia 27-56 X 4-7.5 ,, simple or with
short protuberancesup to 6.5 ,ulong at the apex of the main body or all over, rarely
with 2-3 longer branchesup to 14,, never truly echinate of the Siccus-type. Hyphae
with clamp connections. Coveringlayers: hairs of the pileus 3.5-9.7 u broad, more often
acute or subacute than broadly rounded at the tip, with the apex of the rounded-tip-
ped hairs often slightly inflated and thin-walled,otherwise the walls up to 3.5 , thick
in the broadest hairs, hyaline to hyaline-stramineousin KOH, pseudoamyloid, some-
times septate below the apex by a secondary septum, not ladder-like.
On small stalks of Monocotyledones, apparentlyon grasses,fruiting from March
to April.
MATERIALSTUDIED. VENEZUELA:Orinoco-region,Atures,A. Gaillard136 (FH, type).
ECUADOR:Quito, Mar 1891, Lagerheim(FH, authenticJ.
This species is very close to the following one and to C. urbica. I have not been
able to re-coliect it.
Crinipellis 19

Patouillardand Gaillard(1888) P1 7, fig. 3.


9. CrinipellispseudostipitariaSinger, Lilloa 8: 470. 1942, subsp pseudostipitariavar

CrinipellispseudostipitariasubspoccidentalisSinger,Lilloa 8: 470. 1942.
TYPEDuss 492, from Guadeloupe(FH).
Pileus rusty fibrillose-woollywith fimbriate margin,dried tan color "cinnamon
buff' to "Sayal brown" (Ridgway), or "Mi ado brown." almost uniform in color, evenly
hairy all over, or the center slightly less hairy, with more appressedfibrillose-pilosecover-
ing, in dried material sometimes with concentric walls and furrows but these much less
distinct and less constant than in C. bisulcata. hemisphericalor almost conical when
young, then convex eventually applanate,with depressedcenter, sometimes distinctly
umbilicate, 3-13 mm broad. Lamellaewhite when fresh, close or subclose, rarely sub-
distant, sometimes some forked, subventricose,narrowto medium broad, attenuate and
narrowlyadnexed to subfree. Stipe concolorous with the pileus or more bister especially
where the hairs have been rubbed off, spiny strigose from agglutinatedhirsute hairs be-
low, more tomentose-pilose above, the apex eventually subglabrous,solid, central, equal,
insititious, 9-30 X 0.3-2 mm. Context white, inodorous.
Spores 7.7-10 X 3.5-4.5 ,, Q 2, ellipsoid to fusoid, hyaline, smooth, inamyloid.
Hymenium:basidia 16-27 X 5-7.7 p, 4-spored, Cystidia none; cheilocystidia 17-35 X
4-7 p, simple or forked, with one to mostly severalapical and/or lateral finger-likeappen-
dages 3-12 p long and sometimes spreadingto 12 p in breadth. Hyphae with clamp
connections, hyaline in KOH, inamyloid. Coveringlayers: hairs of the pileus 4-7 p
broad, light melleous, some hyaline, pseudoamyloid, with walls 1.2-2.5 juthick, with
rounded, more rarelyattenuate-obtusetips, without ladder-likesecondary septa (only
some hairs with scattered septal); hairs of stipe 5-7.7 p broad, with attenuate-obtusetip,
otherwise like those of the pileus.
On dead fallen grass debris and other monocotyledonous plants, sometimes passing
over onto other herbaceousstems or small sticks rotting on the ground nearby, gregar-
ious, fruiting in the rainy season. Known hosts: Panicurnmaximum.
MATERIALSTUDIED:CUBA: Wright855 (FH), probablybelonginghere. GUADELOUPE:
MorneHirondelle,Duss 1 765 (FH); CampSacal,Duss 1888 (FH, this and precedingcollection are
paratypes);Plateaudes Rivieres,Duss 492 (FH, type).BOLIVIA.La Paz: Nor-Yungas,Charobamba,
Singer B 720, 30 Jan 1956 (F).

9b. Crinipellispseudostipitariavar mesites Singer, Lilloa 8: 471. 1942. Fig 4.

TYPE.Duss 1877, from Martinique(FH).
Pileus ferruginous-chestnut,drying to ochraceousbuff with umber center, margin
sometimes pale stramineous,and fimbriatehairy-tomentoseall over, with a small and
ratherlow papilla, with almost sulcate margin,conic-convex to strongly convex, then
more applanate, 5-10 mm broad. Lamellaewhite, rathernarrow,close to almost sub-
distant, subfree. Spore print white. Stipe concolorous with pileus, tomentose-piloseto
strigose-pilose,insititious, equal, 9-35 X 0.3-1.5 mm. Context white, inodorous.
Spores 7-10 X 3.7-6 ,, with rathervariableQ (mostly about 1.8-2), ellipsoid, hyaline,
smooth, inamyloid, unicellular-aseptate.Hymenium:basidia 22.5-29.5 X 5-7.5 p, all
4-spored, or (2-3-)-4 spored;cystidia none; cheilocystidia 17-34-41 X 4-9.5 p, crowded
at the edges of the lamellae, few in the immediate neighborhoodof the edges, rarely
simple and entire and then mostly clavate, frequently forked or with three apical bran-
ches, but also apically appendiculatewith 3-5 branchesmaking the cheilocystidiumhand-
or Ramaria-shaped,very rarelywith strictly apical setula-likeappendages(en brosse),
20 Flora Neotropica

hyaline. Hyphae hyaline with clamp connections, inamyloid. Coveringlayers: hairs of

the pileus similar on marginand in center, 3.5-8 p broad, with occasional distant second-
ary septa, not ladder-like,with 1.5-2.2 p thick wall, pseudoamyloid,brownish in KOH,
mostly attenuate-obtuseor broadly rounded at the tip, rarely with an obtuse appendage
at the tip; Hypotrichiumconsisting of hyaline clamped hyphae with rather numerous
septa. hairs of the stipe 5-7.7 p broad, with 1.8-2 p thick pseudoamyloidwall, with
attenuate-obtuse,more rarely broadly rounded tip.
On dead grassesand Bambuseae,gregarious,fruiting in the rainy seasons. Known
hosts: Andropogon sp; Chusqueasp.
MATERIAL STUDIED. MARTINIQUE. Parc du College de St. Pierre, Aug 1900, Diiss 1877
(FH, type). BERMUDA. Brownr,Britton & Seaver 1540 (det. Murrill as C. stupparia), (NY, paratype).
ARGENTINA. Tucuman: Capital, 22 Jan 1955, Singer T 2071 (LIL). NIGERIA. Niger Valley Clhe-
alier (det. Patouillard as C. stipitaria), (FH).
This variety is intermediatebetween subsp pseudostipitariaand subsp orienltalis,
the latter having spores broaderthan 5 p (none narrower)growing exclusively in the

10. CrinipelliscatamarcensisSinger, Rev. Mycol. 18: 12. 1953.

TYPE. Singer T 1 745 (LIL) from Argentina.
Pileus rusty brown (between "Arab"and "burnt umber" M&P),hygrophanous,
faded about 12 D 8 M&P,long-hairywith undulatingappressedpilose covering all over
but near marginin dry condition the white trama of hypotrichium showing through
among the strands of hairs at least when mature, with a circularelevation around the
disc, or near the margin,showing a concentric circularslight depression,but at first
convex, later more repand and mostly obtusely low-umbonate, never abruptly papillate,
about 21 mm broad. Lamellaecream color (e g 9 C 3 M&P),about 2 mm broad, close,
free. Stipe nearly concolorous with the pileus, appearingfibrillose from numerous long
hairs, attenuate towards the insititious base, about 18 X 3.5 mm. Context white, thin,
Spores 5.5-7.5 X 3.54.5 ,, ellipsoid, broader than half their length or as broad
as half their length, hyaline, smooth, thin-walled,then moderately thickish walled, in-
amyloid but mostly a few pseudoamyloidones present, not septate. Hymenium:
basidia 26-27 X 6.5 u; cystidia none; cheilocystidia 20-25 X 6.5-7.5,, clavate or ven-
tricose, not appendiculateor with 1 to severalapical appendages1-14p long and then
often spreadingabove and broader than 7.5 u, rarely longer than 25 p (reaching43 ,
in some cells), all hyaline. Hyphae of the regularhymenophoral trama inamyloid
and with clamp connections. Coveringlayers: hairs of the pileus in KOH melleous,
fewer subhyaline, weakly but distinctly pseudoamyloid, the majority rounded-obtuse
at the tip, but in the center of the pileus also many acute or attenuate-obtuseones
present, on the marginnearly all rounded, 5.5-7(-8.2)u broad, long and without con-
spicuous cross-walls,with 2-2.8 , thick wall.
On roots of monocotyledonous plants in open montane vegetation, fruiting in
summer. In subxerophyticvegetation, often solitary. Known host: Cvnodon sp.
MATERIAL STUDIED. ARGENTINA. Catamarca: Valle del Rio Campo, NW of the Alto
de la Junta, Mesada, 17 Jan 1952, Singer T 1745 (LIL, types); Estancia Santa Rosa N of Rio Campo,
1650 m alt, 20 Jan 1952, Singer T 1782 (LIL, paratypes).

11. Crinipellisschini Singer, sp nov

TYPE. Argentina,Catamarca,Singer T 1 793 (LIL)
Pileo brunneo, appressepiloso, umbonato-papillato,8-19 mm lato. Lamellisalbis
Crinipellis 21

vel cremeis, liberis. Stipite subconcolori, appressepiloso, in parte inferiore hirsuto-rudi,

35-36 X 1.5-1.7 mm. Sporis exiguis 5.5-6.3 X 3-4.2 j, nonnullis pseudoamyloideis.
Cystidiis nullis; cheilocystidiis 15-34 X 6.8-8.2 p, nodulosis vel appendiculatis;pilis
pilei rotundato-obtusis. Ad stipulas ligneas at lignum putrescensSchini aliorumque
Dicotyledonum in zona subtropicalisubxerophyticamontana. Typus in LIL conservatur.
Pileus brown ("raw sienna" M&P,center eventually reaching"burnt umber" M&P),
not fading in dried condition or becoming "hazel" to "chipmunk,"appressedlypilose
all over, convex, later applanatein the wide marginalzone, soom depressedor flattened
around a large conspicuous papilla, this persistent, diameter 8-19 mm. Lamellaepure
white to pale cream, subclose, broad, subventricose,free. Stipe almost concolorous
with the pileus, appressedlypilose, but in lower portion hirsute-rough,insititious, equal
35-36 X 1.5-1.7 mm. Context white or whitish, inodorous.
Spores 5.5-6.3 X 3-4.2(-4.5) A, ellipsoid with a Q just barely above 2 to decidedly
below 2, mostly about 1.8-2, without suprahilardepression,hyaline, many pseudoamyl-
oid, smooth, Hymenium: basidia27.5 X 6.2 p, 4-spored;cystidia none; cheilocystidia
15-34 X 6.8-8.2 ,clavate or cylindricalor ventricose, most frequently clavate, with
nodules or short to long apical appendages(one to many, mostly 2-3); Hyphae thin-
walled, hyaline, inamyloid, with clamp connections. Coveringlayers: hairs of the pileus
(middle between center and margin)varyingfrom deep melleous to hyaline, thick-walled
with or more frequently without secondary septa, and these if present not forming a
ladder-likestructure, with broadly rounded, more rarely attenuate-obtuse,but never
acute tips, very long, pseudoamyloid, smooth.
On rotting fallen sticks and pieces of wood of dicotyledonous trees and shrubs
in the subxerophytic low woods of the subtropical,montane zone of northern Argen-
tina, gregarious,fruiting in the summer rainy season. Known host: Schinus sp.
MATERIALSTUDIED. ARGENTINA. Catamarca:Hill above the EstanciaSanta Rosa
near the Rfo Campo, 20 Jan 1952, Singer T 1793 (LIL, type); between Rio Chacrasand Rio Pot-
rero, 11 Feb 1952, Singer T 1855 (LIL).
The small pseudoamyloidspores remind one of C. catamarcensis,a species from
the same region but which grows on Monocotyledones ratherthan on Dicotyedones and
has a nonpapillatepileus and both absolutely and relatively shorter stipe. The lamellae
are paler in C. schini.

12. CrinipellisatrobrunneaPatouillard,Bull. Soc. Mycol. France 7: 308. 1891.

TYPES.Bon 4362 (FH), from Vietnam.
Pileus rather dark brown "Cattail, cafe noir" M&P,fading somewhat but still
relatively deep colored when dry on the margin,often fading much more in the center
where it becomes pale leather brown or light buff, with entire or somewhat crenulate
margin,strongly pilose all over, in old ones a flattened disc often surroundedby a
series of concentric furrows or one circulardepression,young convex, later more re-
pand and more or less umbilicate, 9-17 mm broad. Lamellaepure white when fresh
but tending to become brown in the herbarium,rather broad, moderately close or
distant, with 12-29 lamellae + lamellulae,rounded-freeor attenuate-subfree. Stipe
concolorous or "Cochin"M&P,strongly hairy with strandsof conspicuous hairs, equal
or subequal, insititious, 10-30 X 0.5-1 mm. Context white, very thin, inodorous.
Spores 6.5-9.5 X 3.5-5 L, with Q slightly below or slightly above 2, ellipsoid,
hyaline, smooth, inamyloid, Hymenium: basidia 23-30.5 X 6.5-8.5 ,, 4-spored;
cystidia none; cheilocystidia 19-48 X 3.5-8.5 p, versiform,entire to branchingwith
up to five usually more or less straightand obtuse branches,the common base or
22 Flora Neotropica

the entire cell cylindric to subfusoid or subclavate,if entire often strongly sinuose-
wavy or if branchedthe branchesmore rarely curved or crooked, and these rarely
subacute or acute, in dried material often strongly agglutinated,hyaline. Hyphae of
the trama filamentous, inamyloid, nongelatinizedwith clamp connections. Covering
layers: hairs of the pileus 4.3-10.5 broad, in KOH and ammonia stramineous-yellow-
ish to deep melleous, the paler ones in dried materialoften showing (necro-)pigment
precipitatedin brown granules,with distant secondary septa or without them, but
without ladder-likestructureeven near the mostly broadly rounded tip, very rarely
some with such a structrue,with 0.7-5.5 p thick wall, slightly but distinctly pseudo-
On grass culms and grass roots, often densely gregariouswith deeply immersed
stipes, in South America generally in gardens,especially seed beds, and apparently
adventitious. Fruiting in the rainy season, September and May in Asia, summer
(December to February)in South America.
MATERIALSTUDIED. ARGENTINA. Tucuman, gardenof the Instituto MiguelLillo,
Singer T 108 and 108a (LIL). NORTH VIETNAM. Hanoi,Bon 4362, 31 May 1890 (FH type):
Cai Kinhmassive: Bau Mau(Tonkin),Bouitan341b (FH, authentic).

13. CrinipellismissionensisSinger in Singer & Digilio, Lilloa 25:226. 1952.

TYPE. Singer & Digilio M 123 (LIL), from Argentina.
Pileus fuscous ("English oak," when younger "Java"but less reddish, with paler
margin,M&P),concentricallyzonate and at the extreme marginalmost white-pilose
with deeper colored hairs further inside, the hairs here more appressedlyand wavy-
tomentose but in the zone immediately surroundingthe papilla less hairy and often
almost naked, radially rugose in the intermediatezone, the papilla central, deepest
colored and nearly or quite glabrous,convex, the marginalregion becoming applanate.
with at first involute or strongly incurvedmargin,reachingabout 40 mm wlhenfully
mature. Lamellaewhitish when seen laterally, but with flesh colored to ochraceous
("sunset" M&P)shades, when seen from above, moderately broad, but very broad
near the stipe (4-4.5 mm broad there), subclose or moderately close, intermixed with
lamellulae,adnate to sinuate, not free. Stipe concolorous with the pileus, the apex
darkest, the middle zone usually lighter colored, entirely strigose, longitudinally
grooved-sulcate,solid, equal but at the apex and/or base mostly broadened,about
60 X 4 mm in fully mature specimens, insititious or almost so. Context white, fleshy,
but seemingly tough because of the hairy layers; odor of bitter almonds, but rather
Spores (7.5-)8.2(-9.3) X (5.4-)5.5(-6.8)u, ellipsoid, hyaline in ammonia, smooth,
some becoming slightly opaque and moderately thick walled, inamyloid, unicellular.
Hymenium: basidia 23-37 X 6-8.8 ,; cystidia none; cheilocystidia making the edge
heteromorphous,30-42 X 4.2-7.5 p, mostly entire and simple, versiform,often fusoid,
cylindrical, subclavate,often irregularin shape, mostly short and narrowlymucronate,
a minority of cheilocystidia laterally or apically branchedor forked at the apex, quite
or rather thin-walled,not or scarcely opaque, hyaline. Hyphae hyaline in the trama of
the pileus, inamyloid, with clamp connections, hymenophorallayer not gelatinized,
regular,hyaline. Coveringlayers;hairs of the pileus and stipe 2.7-6 , broad, some
densely fasciculate and agglutinated,in KOH melleous to hyaline, smooth, with 0.5-
1.3 p thick wall, strongly pseudoamyloid, attenuate with obtuse tip or broadly rounded
at the tip, those of the stipe slightly narrowerand with slightly thinner wall than those
of the pileus.
Crinipellis 23

On wood of Dicotyledones, growing from exposed roots of trees in ditches in

wet subtropicalforest, fruiting in spring(November, December)
MATERIAL STUDIED.Known only from the type locality: ARGENTINA. Misiones:
Cataratasdel Iguaz6 27 Nov 1949, Singer& Digilio M 123 (LIL).
This is easily the largestof the neotropical species of this section, and in some
aspects remindsone of Crinipelliszonata (Peck) Patouillard,from which it differs in
spore characters.
It is close to the African Crinipellisghanaensis Singer in Pegler but differs from
the latter in havingbroaderspores. Both the African and South Americanspecies
differ from C. zonata in the inamyloid spores, and in the absence of a characteristic

14. CrinipellisherreraeSinger, sp nov Fig 5.

TYPE: Singer 8125, from Veracruz, Mexico.
Pileo ferrugineo-brunneo,KOH ope haud virescente nec grisente, in disco tomen-
toso, circum discum piloso pilis centrum versus depressis,marginemversus sparsepiloso,
sulcato, convexo, centro depresso, obtuso, 9-19 mm lato. Lamellissubdistantibus,sub-
latis, subliberis. Stipite hyaline- vel pallide brunneo-piloso,subaequali8-16 X 1-1.5
mm. Sporis (7-)9-10 X (4-)4.5-5.3A,, ellipsoideis vel oblongo-subfusoideis. Basidiis
tetrasporis. Cystidiis nullis. Cheilocystidiisnumerosis,dimorphis. Hypotrichio
ex elementis latis ferrugineo-incrustatisefformato. Pilis haud multiseptatisneque
acutis, crasse tunicatis;eis stipitis similibus. Ad ramulosexiguos in silva pluviali,
typus in Mexico: Veracruz,Estacion Biologica de las Tuxtlas, SingerM 8125 (F).
Pileus rusty brown but ratherdeep colored and almost unchangedon drying,
ratherthinly beset with tough, long, brownish hairs which are denseraround the flat
disc where the hairs appear depressedinwards,on the small disc region few or no
hairs visible but here the surface tomentose (whereas in the marginalzone the surface
is glabrousamong the sparsehairs), sulcate on the margin,smooth in the center, con-
vex, soon with depressedcenter, neither umbilicate nor papillate or umbonate, 9-19 mm
broad. Lamellaebrown when collected, subdistant, not intervenose,medium broad to
broad, with white edge, narrowlyadnexed to subfree. Stipe with numeroushyaline to
melleous hyaline erect or suberect hairs on light brown to almost pallid ground,only
near base ground color brown, hirsute because of long (up to 0.5 mm) hairs in the
basal portion, the hairs somewhat shorter higher up, subequalor taperingdownwards,
insititious, 8-16 X 1-1.5 mm. Context white, inodorous.
Spores (7-)9-10 X (4-)4.5-5.3 u, with a Q 1.8-2, ellipsoid to oblong-subfusoid,
hyaline to brownish hyaline, inamyloid, smooth. Hymenium: basidia 23-27 X 5-7 ,,
4-spored;cystidia none; cheilocystidia numerous, making the edge heteromorphous,
dimorphic,(1) near marginof pileus 13-17 X 6-9 p, clavate to subventricoseand
broad, much like the cheilocystidia of most Inocybes, brownishhyaline, smooth and
entire, thin-walled,with an occasional hook, appendageor bulge at the apex, (2) over
most of the edge 28-38 X 6.5-9 u, with severalapical finger-likeappendages,and be-
cause of these apically spreadingto about 20 ,, main body cylindricalor subventricose
to clavate, often with ramificationsfrom the base or middle and thus assumingthe
shape of a carpophoreof Ranzaria,the branchesand appendagesusually with rounded
tips, rarely acute, sometimes secondarilyforked or secondarilylaterally or apically
appendiculate. Hyphae of the trama brownish-stramineous-hyaline, inamyloid, with
clamp connections, thin-walled,relatively broad but filamentous. Coveringlayers:
hairs of the pileus mostly more than 200 p long, not densely crowded, in KOH light
24 Flora Neotropica

melleous to tawny, without ladder-likesecondary septa, with the tip not acute, with
a diameter of 5-8 ,u,most frequently 5-6.5 ;,, with the wall 1.5-3 M,most frequently
1.5-2 Hthick, pseudoamyloid;hypotrichium consisting of repent broad (5-15 ,), rusty-
incrusted hyphae with clamp connections; hairs of the stipe very similar to those of the
marginof the pileus, stiff, more variableand less pigmented, pseudoamyloid but weaker
than the hairs of the pileus, 18-200 X 5-7 I in the apical, up to. 500 A,long in the lower
half, with hyaline wall 1.5-2 ,uthick, rarely with a short obtuse side-branch,otherwise
like the hairs of the pileus.
NaOH on dried pileus: More brightly ferruginous-brown,not green or gray.
On small woody twigs, apparentlyof dicotyledonous trees or vines, gregarious,
fruiting in the rainy season.
MATERIALSTUDIED. MEXICOVeracruz:7 km south of Montepio,Estacion Biologica
de las Tuxtlas,Herrera,21 June 1969, comm. SingerM 8125 (F, type).
The species is named for Dr. Teofilo Herrera,noted Mexican mycologist, collector
of the species.

15. Crinipellisseptotricha Singer, Lilloa 8: 468. 1942.

TYPE. Duss 513 (FH), from Guadeloupe.
Pileus rusty brown when fresh, but soon fading to avellaneousbrown, pale brown,
fulvous-buff, more or less strongly pilose, the hairs in curly strands radiallyappressed,
merely tomentose or somewhat less thickly pilose on the papilla, convex, but often with
a concentric wall or depressionaround the disc, more rarely alnost terraced, with a
moderately prominent, rarely inconspicuouspapilla visible in both fresh and dried
condition and located inside an umbilicus, or rising without an umbilicus, eventually
the whole middle zone flattened or depressedor at least becoming so in dried material,
3-16 mm broad mostly 6-12 mm. Lamellaepallid or white, then mostly becoming
brownish pallid especially in the edge zone, medium broad, subventricose,close or
subclose to medium close, not subdistantnor distant, emarginate-freeor subfree.
Stipe concolorous with the pileus, hairy, insititiOus, equal, 16-32 X 1-1.5 mm. Context
white, thin, inodorous.
Spores 6.5-12 X 3.5-6 p, ellipsoid, few slightly oblong, with or without a suprahilar
depressionor applanation,with Q > 2, rarely and ratherexceptionally very slightly
below 2, but often = 2, smooth, hyaline in KOH, with thin, eventually slightly thick-
ened wall, inamyloid. Hymenium: basidia 19-32 X 5.5-8.5 p, often 2-spored;basidioles
fusoid; cystidia none (or few cheilocystidia slightly away from the edge occurring);
cheilocystidia 25-37 X 4.2-7u versiform,often fusoid, ampullaceous,cylindrical, nar-
rowly subventricose,subclavate,the apex sometimes forked or more rarely subcapitate,
with or without (more often without) 1-5 apical finger-likeappendages,these 4-14 X
2.5-3 p, inserted at the same or at different levels, wall thin or somewhat thicker than
that of the basidia, hyaline to brownish hyaline, rarely with small wart-likeoutgrowths
all over their length, making the gill-edgeheteromorphous. Hyphae of the trama hyaline
thin-walled,inamyloid, with clamp connections. Coveringlayers: hairs of the pileus
(middle zone outside the central papilla) mostly light melleous-fuscidulous,some hyaline
in KOH, with a dense ladder-likestructurefrom numerous secondary septa at least
near the tip in most hairs, the septa at distances of between 5 and 11 , diameter of
the hairs 3-10.5 ,, walls mostly narrowedtowards the tip, but reaching,further below,
3(-4.5), thick, apex usually more or less attenuated, with acute, often hyaline, or
broadly rounded tips, some hairs with appendagesat the tip, pseudoamyloid;hypoth-
richium poorly individualized,hyphal cells here often with thicker walls than in the
Crinipellis 25

trama below, and broader;hairs of the stipe (middle of the stipe) simple and long,
rarely with a short lateral branch, the tips subacute to acute, with or more rarely
without ladder-structure,with 6-10 ,udiameter, with 1.5-2.7 , thick wall.
KOH on the dried pileus darkerbrown, but not tending to gray or greenish.
On dead wood of trees and shrubs, fruiting from December to August, often in
subxerophytic vegetation or in plantations. Known hosts: Coffea arabica,Achatocarpus
Fink (FH), probablybelonginghere. CUBA. Wright45, det. Berkeley& Curtis(A. stipitarius),
(FH, paratype). GUADELOUPE.Duss 484 (FH, paratypes). Duss 486 (FH, paratype):Basse-
Terre,Duss 513, 14 Jul 1901, (FH, typus). ARGENTINA. Tucuman:5 km S of Buruyacu,19
Feb 1957, Singer T 2989 (F); Posta de Lozano, 9 Feb 1966, Singer T 4995 (F).
Singer (1942) describedoriginallya "forma umbilicata ad int" differingfrom the
typical papillate form in the absence of a papilla in the umbilicus or on the disc, grow-
ing on smallersticks and twigs. Dennis (1951a) observed this form in the Arena For-
est on Trinidad(K) and described it with slightly longer spores (9-11 X 4-5 u),
This comes close to an extratropicalSouth Americanspecies which differs how-
ever in subglabrousto glabrous disc on the pileus and broaderspores. It is Crinipellis
mlolfinoalna (Speg.) Sing. which was described by Spegazzinias Marasmiusmolfinoanus
Speg., Bol. Acad. Nac. Ci. Cordoba29: 123, 1926, from dead roots of Fagaracoco. Only
the type (leg. Mlolfino212, LPS) is known. The spores are 9-11 X 6-7.5 u; the cheilo-
cystidia are entire or forked or with both apical and lateral short appendagesor diver-
ticula; the hairs are about 8 u broad and have the ladder-likestructureof C. septotricha,
with attenuate-obtuseto frequently needle-sharpacute tips, at least those at the mar-
gin of the pileus. It comes from the SierraGrande,provinciade Cordoba, at Yaci,
February 1912. This species, certainly different from C. septotricha, is mentioned
here because its host extends north into the subtropicalzone, together with the "Monte"
-vegetation characteristicfor the Sierrasde Cordoba.
ILLUSTRATION.Dennis (1951a), p1 20, fig 6.

16. Crinipellispodocarpi Singer in Singer & Digilio, Lilloa 25: 225, 1952. Fig 6.
TYPE. Singer T 833 (LIL), Argentina.
Pileus in the broad marginalzone light brown to ochraceous-ferrugineous (reach-
ing "Arab"M&Pfading to "terrapin"M&P),the entire disc region darkerchestnut to
fuliginous or the disc proper, a deep chestnut to ferruginous-fuliginousdot surrounded
by a narrowzone which is light rusty brown fresh and grayishpallid dried, and this
zone separatedfrom the marginalzone by a deeper colored (deep brown when dried)
concentric line, the extreme marginoften stramineous-pallid,hairy all over, but the
hairs long, often fasciculate in the broad marginalzone and frequently projectingbeyond
the margin,whereas in the disc zone the hairs are short and the surface appearsmore
tomentose to almost subglabrous,with some erect bristle-likehairs occasionally showing
on the papilla, the latter at first very prominent and reaching 1 mm in height, later
becoming inconspicuous and appearinglike a tiny umbo under a lens, convex with
applanatezone around the papilla or slightly depressedin the center in adult material,
the marginalzone strongly declivous for a long time and often radially sulcate, even-
tually more repandbut with circularcircumference,4-13 mm broad. Lamellaewhite
or whitish, broadertowards the stipe but still narrowto moderately broad there, close
to subdistant, mostly close or subclose, free. Stipe chestnut brown (e g 8 J 12, M&P)
below and somewhat paler (light brown) above, strongly pilose and hirsute below, less
hairy above, straight or curved, central very rarely slightly eccentric,equal or more
26 Flora Neotropica

rarely slightly broadenedat or near the insititious base, 9-28 X 0.8-1 mm. Context
white, thin, fleshy (but seemingly tough because of the hairy covering), inodorous.
Spores 7-8.2 X 5-7.2 u, with Q = 1.3-1.8, ellipsoid to subglobose, without sup-
rahilarapplanationor depression, hyaline, smooth, thin-walled,later often somewhat
thickish walled, inamyloid. Hymenium: basidia 19-36 X 7-8.8 ,, 4-spored;cystidia
none; cheilocystidia 12-35 X 6-8.5 p, with clavate, ventricose, or cylindricalmain
body and with four to nine erect or obliquely erect apical finger-likeappendages
which are 3.3-15 X 1-7 p,, mostly 3.5-5 X 1-2, , simple, more rarely forked, hyaline
(as the main body), inamyloid. Hyphae of the trama of the pileus and hymenophore
filamentous, non-gelatinized,thin-walled,hyaline, inamyloid, with clamp connections.
Coveringlayers: hairs of the pileus (marginalzone) long-filamentousor graduallytaper-
ing to a narrow-obtuseor broadly rounded tip, melleous in KOH, with thick (0.6-2 p)
wall which towards the apex often becomes thinner and more hyaline, with or without
sparseto numerous secondary septa (but these rather distant from each other, extra-
ordinarilyweakly and slowly pseudoamyloid, 3-8 , broad, mostly about 3.3-4.3 Abroad:
hypotrichium cutis-like, consisting of pale melleous to subhyaline short, rather broad
(2-8.3 u), often curved hyphal cells with clamp connections and with up to 1 Athick
walls, often with hyaline incrustationbut not pigment-incrusted.
On living and dead trunks and branchesof Podocarpusparlatorei, often ascend-
ing up to 3.5 m, gregarious,fruiting in the summerrainy season.
MATERIAL STUDIED.ARGENTINA. Jujuy:15 km NEof Postade Lozano,20 Feb 1966,
SingerT 5309 (F);Tucuman: Taficillo,6 Jan1950,SingerT 833 (UIL,types).
This species is often parasiticon Podocarplusin the subtropical-montaneto trop-
ical-montanezone of the Selva Tucumano-Boliviana. It is apparentlyclosest to C. stip-

17. Crinipellisstupparia(Berkeley & Curtis) Patouillard,Essai Tax. Hym. 143. 1900.

Fig 7.
AgaricusstuppariusBerkeley& Curtis,Jour. Linn. Soc. 10: 285, 1869.
CollybiastuppariaBerkeley& Curtis)Saccardo,Sylloge Fungorum5: 217. 1887.
TYPE. Wright(K, FH), from Cuba.
Pileus ochraceous brown in the broad marginalzone, fading to near "thrush"M&P,
with blackish brown disc which is often surroundedby a blackish line, sulcate over
about one third of the marginalradius, strongly and often coarsely pilose-floccose
with mostly fasciculate and agglutinatedhairs which form more or less curly-undulate
strandsbut graduallyless hairy and more tomentose and eventually glabrescentaround
the papilla, the latter mostly either naked or more often with a bunch of erect bristle-
like hairs crowning the tip of the papilla, convex or hemisphaerical,becoming applanate,
the center soon or eventually becoming depressedaround the usually prominent papilla,
1.5-7 mm broad. Lamellaewhite, in herbariumoften becoming brownish-wood-color,
medium broad to rather narrow,medium close to subdistant(12-20 lamellae and
lamellulae) free to narrowlyadnexed. Stipe concolorous with the pileus, pilose all
over, but at times hirsute hairs scattered, equal or somewhat attenuated upwards,
solid, then hollow, curved or straight,central, 3-25 X 0.5-1 mm. Context white, thin,
often with a brown periphericline or zone especially in the stipe, inodorous.
Spores 6-9.8 X 4-7 p, Q = 1.4-1.8(-2), ellipsoid to short ellipsoid, smooth, hyaline,
inamyloid, thin-walled,eventually often slightly thickish walled. Hymenium: basidia
19-26 X 5.5-10.5 p. 4-spored;cystidia none; cheilocystidia 15-38 X 3.5-11 pu,with
clavate, subventricoseor subcylindricalmain body and with four to twelve finger-
Crinipellis 27

like or forked apical appendageswhich are either erect or obliquely upright rarely
distributedon a cylindric elongation of the apex of the cheilocystidium and branch-
ing at different levels of the apex so that the apex looks like a fir tree, much more
commonly all at the tip, arisingat the same level, rarely the main body sending out
a long lateral filamentous branch, branchesand appendagesoften secondarily diverti-
culate by short wart- or knot-like protuberances,rarely the whole cheilocystidium
entire and smooth (in some preparationsno such cells found), the apical appendages
1.5-4.5 X 1-2 p and obtuse, the others 1-lOp long, the entire cell hyaline, often more
than 11 broad if appendagesor branchesare spreadingout above, not thick-walled,
inamyloid. Hyphae of the tramahyaline, inamyloid, with clamp connections. Cover-
ing layers: hairs of the pileus long and straight or flexuous, 4-11 broad, the walls
melleous-brownor melleous-fuscousto (more rarely) subhyaline, strongly pseudoamy-
loid and cyanophilous, walls 1.5-4s, thick, taperingat the apex to an obtuse tip, or
with a broadly rounded tip, without secondary septa, rarely with one or very few
scattered secondary septa, never with ladder-structure;hypotrichiumconsisting of
ratherbroad, cylindric, sometimes branching,almost thin- to moderately thickish-
walled hyphal cells which are inamyloid and melleous to brown pigmented, the
pipnent intraparietalbut also here and there incrusting the walls.
KOH on the dried pileus negative.
On sticks, branches,small twigs and fallen leaves of the evergreentough type,
always on dicotyledonous never on living host tissue, growingusually gregariously
in shaded ravinesand cloud forest vegetation but also in subtropicaland tropical
forest of the plains from Cuba to South Brazil.
MATERIALSTUDIED. CUBA. Wright(FH, type); Wright15 (part) (FH). VENEZUELA.
Dto,Federal:SierraCostera,El Junquito, + 1800 m alt, Deinnis1800, 10 Jun 1958 (K). COL-
UMBIA.Valle: Mun.Call: CerroLa Horqueta,+ 2000 -malt, 2 May 1968, R. SingerB 6903 (F).
BRAZIL. Rio Grandedo Sul: Sao Leopoldo, ex HerbarioTheissen,Fungi Austroamericani,Rick
213 (FH).
The Braziliancollection has been determined by Theissen as Marasmiusthwait-
esii Berk. & Br., but it is not identical with that species.

18. Crinipelliscoroicae Singer, sp nov Fig 8.

TYPE.Singer B 558 (F), from Bolivia.
Pileo cupreo-ferrugineo,in siccis colore C. stipitariaegaudente, piloso, centro
tomentoso-piloso vel fibrilloso-tomentoso, convexo, dein applanato, obtuso vel centro
depresso, 9-11 mm lato. Lamellisalbis, confertis, liberis. Stipite pileo concolori, sed
ad apicem albo, glabro, basin versus piloso, aequali, 23-32 X 0.5-0.7 mm. Sporis 7.5-9
X 4-5.3,, ellipsoideis, inamyloideis. Basidiistetrasporis. Cystidiis nullis. Cheilocysti-
diis 16-28 X 5-7.5 j,, ventricosisvel clavatis et duobus vel six appendiculisapicalibus
coronatis. Hyphis inamyloideis, fibulatis. Pilis in pileo obtusis vel acutis, in stipite
Pileus copper-rustcolor, soon with whitish portion on the marginalpart, with
curly pilose covering evenly all over, or more tomentose-piloseto fibrillose-tomentose
in the center, there also often darkercolored at least in dried condition, dried fading
to the dull tan-strawcolor of C. stipitaria, with or without a small umbilicus when
seen in dried condition, and if with umbilicus, there are also concentric walls and
furrows around it, without papilla and only sometimes subumbonateconvex, then
applanateor with depressedcenter, 9-11 mm broad. Lamellaewhite, close, medium
broad to narrow, free. Stipe concolorous with the pileus but with white apex, pilose
28 Flora Neotropica

but with glabrousapex, insititious, equal, 23-32 X 0.5-0.7 mm. Context white, in-
Spores 7.5-9 X 4-5.3 , most frequently 8.5 X 4.5 p, ellipsoid, hyaline, some
pale melleous in KOH, inamyloid, smooth. Hymenium: basidia 17-22 X 6-7.5 p,
4-spored; basidioles fusoid, with basal clamp; cystidia none; cheilocystidia 16-28 X
5-7.5 p, ventricose to clavate and with two to six apical finger-likeappendageswhich
may all arise at the same level, but some also often at a deeper level, or cheilocystidia
forked near the base (where they are 2.5-3.5 p broad), some slightly constricted in the
middle, appendages2-13 p long and some secondarilyforked. Hyphae hyaline, more
or less interwoven in the trama, with clamp connections, inamyloid. Coveringlayers:
hairs of the pileus acute to broadly rounded, most frequently broadly rounded or
attenuate-obtuse,stramineousto melleous-brownishin KOH,psuedoamyloid, 3.5-6.2Ip
broad, with walls 1-21 thick, without ladder-structure;hypotrichium consisting of
rather thick-walled,mostly not swollen elements which are stramineous-hyalineor
hyaline, without any incrustation,or only with a pale melleous one; hairs of the
stipe like those of the pileus, but more often acute.
KOH on dried pileus darker,not green or gray.
On dicotyledonous twigs and branches(various species of trees and shrubs)
fallen to the ground, fruiting in the rainy season.
MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yunlas, Coroico, 26 Jan 1956. Singer B
558 (F, type).
This species differs from C. herreraein relatively longer stipe and in the lack of
the rusty incrustationof the hypotrichiumnit differs from the C. patoitillardiiof the
South Pacific in the lack of glabrouszones in the less differentiateddisc region of the
pileus, the more appendiculatecheilocystidia, and in the habitat. It differs from C.
stuppariain size of the carpophoresand shape of the spores, and from C co/iiiiiuta
in the absence of pleurocystidia.

19. Crinipellistenuipilosa Singer in Singer & Digilio, Lilloa 25: 228. 1952.
TYPES:Sitger T 1119 (LIL), from Tucuman, Argentina.
Pileus ochraceous-brown,almost ferruginous("gold brown" M&P),with "cinnamon
br." M&Pfascicles of hairs in the marginalzone, center more sparselypilose, on black
ground visible among the hairs under a lens in dried condition, whereasthe ground
color appearingunder the same circumstancesamong the hairs of the marginalzone is
white, but without any naked zones, nor distinctly concentricallyzonate, conic-appla-
nate, with a prominent papilla, not umbilicate, about 17 mm broad. Lamellaewhite,
subdistant, almost broad, free. Stipe brown below, concolorous with the pileus in the
middle portion, and white at the apex, flocculose-pilosebelow, appressedlvhairy in the
middle portion, and subglabrousat the apex, subequal, or slightly taperingdownwards,
34 X 2 mm. Context white; odor of Mlicroiimplale foeticlzdn, i e, of sauerkraut.
Spores 6.8-8.3 X 2.7-3.5 ; varying from as broad as 6.8 X 3.4p (Q=2) to as
narrowas 8.3 X 2.7 (Q=3), oblong or sub-fusoid, smooth, hyaline, inamyloid, thin-
walled, some over-agedones eventually thick-walled, 1-celled. Hymenium: basidia
4-spored;cystidia none, or extremely few near the edges and like the cheilocystidia;
cheilocystidia 16-30 p long, with two to severalapical cylindricalappendagesof 5.5-8.5
X 2 p1over a clavate main body, entirely hyaline. Hyphae with clamp connections,
inamyloid. Coveringlayers: hairs of the pileus agglutinatedin bunches, pseudoamyloid,
with broadly rounded tip, 4-8.3 p broad, wall 0.5-0.7 1 thick: hypotrichium of ramose
broad sometimes incrusted hyaline elements.
Crinipellis 29

On fallen dead branchesof dicotyledonous trees in sub-tropicalforest, solitary,

fruiting in the rainy season.
MATERIAL STUDIED.Only the type collection is known.
ARGENTINA. Tucuman:Quebrada de Lules,24 Jan1951,SingerT 1119 (LIL,type).
While there are sometimes some secondary septa in the hairs, there is no ladder
structure. This is the most important difference between this species and C. septotricha.
C. Iiirticeps(Peck) Sing., a temperate species of North America, differs in largersize
and somewhat largerspores.

20. Crinipellismexicana Singer, sp nov Fig 9.

TYPES. Murrill142 (NY), from Mexico.
Pileo papilla penicillatagaudente, 2-5 mm lato. Lamelliscremeo-albis,subconfertis
vel mediocriterconfertis. Stipite atrobrunneoin vegetis, centrali, curvato, 9-14 X 0.3-1
mm. Sporis 7-10 X 3-6.5 p; cystidiis raris,cheilocystidiis simillimis;cheilocystidiis
integris, interdumad apicem bifurcatis,haud multi-appendiculatis.Pilis pilei aut ro-
tundato-obtusisaut acutis. Ad caules ligneos Dicotyledonum in silvis tropicalibus
montanis. Typus in FH conservatus.
Pileus at first deep chestnut brown ("Mandalay"M&P),soon becoming deep
rusty-brown,both fresh and dried slightly lighter colored towards the margin, duller
and lighter colored when faded, reachinga dull stramineous-tan,strongly pilose, rad-
ially curly-rivulosein marginalportion (about two thirds of radius), radiallyappressedly
fibrillose and much less hairy in a disc zone but often with a brush-liketuft of bristle-
like hairs immediatelyaround or more often on the top of the central papilla, the
whole disc region occasionally glabrescent,convex, then applanatearound the umbo
or papilla, later more repandall over with the zone around the umbo or papilla be-
coming depressedand often umbiiicus-like,the papilla constant and abrupt, sometimes
on the tip of an umbo, sometimes prominent especially in young caps, 2-5 mm broad.
Lamellaecream white, tending to become pale brownish in dried material,narrow to
medium broad, subclose to moderately close, not ventricose, free. Stipe concolorous
with the pileus, densely and strongly to sparselyand weakly pilose all over, mostly
more strongly hirsute-pilosebelow and weakly pilose and glabrescentabove, central,
curved, insititious, solid, equal, 9-14 X 0.3-1 mm. Context white.
Spores 7-10 X 3-6.5 ,, most frequently 7.5-9 X 4.3-5.5 p, very variablein size
and shape but much more frequently with Q< 2 than with Q = 2 or > 2, hyaline,
smooth, inamyloid, thin-walled,when over-agedmore or less thick-walled. Hymenium:
basidia 20-35 X 4.5-7.5 p,,4-spored;basidioles fusoid; cystidia rarely observed, but if
present scarce and scattered, and much like the cheilocystidia;cheilocystidia making
the edge of the lamellae heteromorphousor almost so, 10-50 X 5.5-7 p, mostly 20-28
X 5.5-6 p, characteristicallyentire and not apically appendiculateor only rarely with
a forked apex or with tripartite(three finger-likeappendages)apex, clavate to vent-
ricose or subcylindrical,more rarelyampullaceous,but often constricted in the mid-
dle, rarely with an occasional lateraldiverticulumor apically crooked, hyaline. Hyphae
of the regularhymenophoraltramahyaline, inamyloid, with clamp connections, inter-
woven. Coveringlayers: hairs of the pileus long, 4-7.8 p broad, melleous to rusty brown-
ish in KOH, some subhyaline, some flexuous, some with rathernumerousthin-walled
secondary septa which are convex towards the apex, but rarely with a truly ladder-
like structure,walls 1-3 u thick, tip much more often attenuate-obtuseor rounded than
30 Flora Neotropica

KOH on dried pileus deeper dull brown, no greenish or gray reaction..

The smallest buttons seen have the hymenium entirely enclosed in the hairs cover-
ing the space between the marginof the pileus and the apex of the stipe.
On sticks and fallen twigs of Dicotyledones in montane tropical forest, gregarious.
Known host: Piperaceae,sp.
MATERIAL STUDIED.MEXICO.Veracruz: Jalapa,12-20Dec1909, W.A. & EdnaL.
Murrill142 NY, type),116 (FH,paratype).BOLIVIA.LaPaz: Nor-Yungas: Capillaria 22 Feb
1956, SingerB 1323 (F, paratype).
Singer(1942) originallybelieved this species to be the same as C patouillardii
Sing. which, in a restrictedsense, i e excluding the neotropical material, occurs only
in the South Pacific and grows on leaves, petioles etc. of Dicotyledones. The carpo-
phores in that species are slightly largerin an average,the pleurocystidiaslightly more
numerous, and the cheilocystidia slightly more frequently forked or appendiculate(with
1-5 apical appendages)and the hairs of the pileus more frequently acute than obtuse
or rounded. Undoubtedly the two species are closely related.

21. Crinipellisfoliicola Singer, Mycologia 47: 771. 1955. Fig 10.

TYPE.Singer F 444a (F), from Florida.
Pileus "MummyBrown" or between "MummyBrown" and "Prout'sBrown"
Ridgway, fading to "DresdenBrown" and "CinnamonBrown", sometimes more rusty
brown when fresh and then becoming sordid stramineouswhen dried, the disc often
darkerbut not blackish, wavy-appressedlypilose, but the disc-zone less hairy and appear-
ing merely tomentose, more rarely evenly hairy all over, if tomentose appearingmore
pallid-canescentwhen dried, except for the darkersmall umbo or papilla, in the margin-
al zone the white ground showing through the strandsof hairs in dried material, one
or two concentric furrows developing in dried material,and sometimes also visible in
fresh material,convex-campanulateor convex, then more or less applanatewith a
flattened or umbilicate depressedzone around the umbo or papilla, the marginalzone
often somewhat coarsely sulcate and occupying about two thirds of the radius or
somewhat more, 5-7.5 mm broad. Lamellaewhite to cream color, close to medium
close, with entire and at first always pure white edge, narrowto broad (up to 1.5 mm
broad), rounded free or attenuate free. Stipe concolorous with or slightly paler than
the marginalportion of the pileus and often slightly darker("Natal Brown" Ridgway)
at the base which is insititious, strongly pilose all over, even hirsute, solid, equal, 15-
48 X 0.2-0.9 mm. Context white, very thin, inodorous.
Spores (6.5-)7.2-10.7(-11.5) X (2.5-)3.2-4.8 p, with Q=1.9-2.5, fusoid to oblong,
or more rarely ellipsoid, hyaline, smooth, inamyloid, thin-walled,the over-matureones
usually thick-walledand sometimes with cross-septum. Hymenium: basidia 19-23 X
5.5-6.5 p, 4-spored;basidioles fusoid; cystidia none excepting, in some collections, some
cystidioles which differ from the basidioles only by being slightly broader(about 10 1O)
cheilocystidia (8-)10-22 3.5-8(-8.5)p, obconic or clavate, cylindrical or ventricose,
sometimes forked or with three branches,with, rarelywithout, finger-or sterigma-like
apical appendagesor hooks (1-7) which may again be somewhat diverticulateor forked,
1.5-6 p long, with rather thin wall, hyaline to yellow or melleous-hyaline,apparentlyat
first all hyaline, making the edge heteromorphous. Hyphae hyaline, inamyloid, with
clamp connections; hymenophoral trama regular. Coveringlayers: hairs of the pileus
in the marginalzone mostly long (about 350 ,), in the central zone often shorter, the
latter more often rounded at the tip, the former variableand both rounded to atten-
uate-obtuse as well as acute tips observed, all ratherpale melleous or more rarely sub-
Crinipellis 31

hyaline to hyaline in the apical portion, and there often with ladder-likeor at least
dense secondary septa, in the lower portion melleous to melleous-umberin KOH,
sometimes somewhat flexuous, rarely somewhat nodose, generally quite smooth and
non-incrusted,strongly pseudoamyloid, 2.7-7.5 p broad, but sometimes broadened to
up to 8 p diameter at the base, with walls, 0.5-1.8 p thick and towards the apex often
even thinner;hypotrichiumhyaline, pigment-less,more rarely pale melleous, multi-
septate with ratherbroad (12-40 X 6-16 ) hyphal cells and/or a single cell 4.5 X 4.5
p at the base of the hairs;hairs of the stipe 5-7 1broad, with walls 2-3 p thick, tips
rarelyrounded, most frequently acute, with or without ladder-structure.
KOH on dried pileus deeper rusty brown, not greeningor graying.
On fallen leaves, rarely passing over to small dead woody twigs, in subtropical
and tropical-montaneforests, gardensand plantations, always on Dicotyledones, soli-
tary or gregarious. Known hosts: Smilax sp., Quercussp., Coffea arabica.
MATERIALSTUDIED. U.S.A. Florida:HighlandsCo., HighlandsHammockState Park,
2 Sep 1942, SingerF 444a (F, type), 30 Aug 1942, SingerF 444 F. paratype). MEXICO. Oaxaca:
SierraMazateca:between Huautlade JimenezandTeotitlandel Camino,km 20-21, 11 Jul 1969,
SingerAl 8444 (F); Rancho del Cura 7 Jul 1969, SingerM 8288 (F); Road to Huautlade Jimenez,
CerroVerde, 6 Jul 1969, SingerM 8274 (F); Chiapas, 15 km W of San Cristobalde las Casas,2400
m alt, 2 Aug 1969, SingerM 8820 (F). VENEZUELA. Dto.Federal:CaracasBotanicalGarden19
Jun 1958, Dennis 1071A (K).
This is very close to C.phyllophila and C. dipterocarpi,the latter from Asia, the
former from Mexico. Crinipellisphyllophila differs in slightly largercarpophores,
slightly broaderspores, almost exclusively acute hairs on the margin of the pileus which
is more often obtuse than papillate and longer hyaline cheilocystidia. The extreme forms
of both species are easy to distinguish,but the more elongated forms of C foliicola
with longer spores - connected with the small-sporedtypical form by transitions- are
often difficult to distinguishfrom C phyllophila. It is not impossible that the latter
is merely an extreme form or variety of C foliicola, but our data on C. phyllophila are
somewhatincomplete with regardto the anatomy of the coveringlayers, and therefore,
for the time being, I describethe two as different species.

22. CrinipellisphyllophilaSinger,Sydowia 11: 358. 1958. Fig 11.

TYPE.SingerM 1507 (MICH),from Mexico.
Pileus in centralring-walland on the ribs between the sulcationsdeep ferruginous,
rich brown, but soon fadingto cinnamontan, paleron the marginbecausethe pale tan to
whitish groundcolor between the more scatteredhairsshows through,concolorous with
the marginor with the ring-wallin the disc-zonewhich is neither glabrousnor shining,but
likewise pilose, the central disc either flat or more rarelywith a centralpapilla,a circular
wall appearingaround the disc-zone, and outside the wall surfacecoarsely sulcate over
one third or one half of the radius,convex, very soon more applanateor even depressedin
the disc-zone, later flat-convexto applanateover all, 6-11 mm broad. Lamellaepure white,
not creamcolor when fresh, close to ratherclose, moderatelybroad or ratherbroad, free.
Spore print pure white. Stipe in upperportion concolorous with the center of the pileus,
in the lower portion concolorous with the margin,pilose, equal, insititious, 30-48 X 0.3-1.2
mm. Context white, thin, reviving,inodorous.
Spores(8-)9.3-10.5(-11.2) X (3.7-)4.3-5 p oblong, somewhatlonger than twice the
breadth,unicellulareven if overmature,hyaline, smooth, inamyloid,thin-walled,slightly
thick-walledwhen over-mature.Hymenium:basidia21-30 X 4.5-6 p, 4-spored;cystidia
none (excepting some occasionalcheilocystidiavery close to the edge proper);cheilocystidia
21-30 X 4.5-6 P crowned by two to six erect appendages2-5.5 X 1-1.2 ,, main body cla-
32 Flora Neotropica

vate to almost cylindric or somewhatventricose, all hyaline. Hyphae inamyloid, with

clamp connections, not gelatinizing. Coveringlayers: hairsof the pileus (marginalzone)
brown or melleous brownish,with the apicalportion usually hyaline or subhyaline,fila-
mentous but often somewhat flexuous, with needle-sharpor at least subacute tip,
never broadly rounded, smooth, 4.2-7 p broad, the walls 1-2.7 p thick, with very few
or no secondarysepta, pseudoamyloid.
KOHon the driedpileus not green or gray.
On fallen leaves of evergreenQuercusin a remainderof originaltropical-montane
mixed oak woods rotting in a thick layer of leafmold, fruitingin summer,gregarious.
MATERIALSTUDIED. Known only from the type locality; MEXICO.Oaxaca:Huautla
de Jimenez,below the Ranchodel Cura,about 1500 malt, 10 Jul 1957 Singer3 1507 MICH,type),
12 Jul 1957, SingerM 1528 (MICH, paratype).

23. Crinipellisdicotyledonum Singerin Singer& Digilio, Lilloa 25: 221. 1952.

TYPE. Singer T 1140 (LIL), from Argentina.
Pileusferruginous"Alamo"M&P,in the marginalzone partly lighter colored, evenly
pilose all over, eventuallyglabrescentespeciallyin the marginalzone, convex and papillate,
eventuallybecomingconvex with depressedcenter and a papillain the depression,about
18 mm broad. Lamellaenot white but buff (10 C 6,M&P),subclose, mediumbroad, emar-
ginate-free. Stipe concolorous with the pileus, pilose, graduallyand slightly attenuated
towardsthe abruptlybut slightly widened base which is insititious,about 19 mm long
and 1.5 mm broad. Context white, thin, with an odor ofMicroniphaleofetidlumor of
Spores 8.5-9.6 X 5.5-6.2 ,, ellipsoid, hyaline, inamyloid, thin-walled,the over-
matureones mostly thick-walled,smooth. Hymenium:basidia33-36 X 6.8-8.2 p, 4-spored;
cystidia none (excepting some cheilocystidiaoff the edge proper);cheilocystidiadensely
crowded on the heteromorphousedge of the lamellae,(20-)27-44(-58) X 5.5-7.5, either
simpleand entire and then cylindric-fusoid,or subulateor ramifiedin the basalor middle
portion of the cell, and then the branches4.8-5.5 pi broad, sometimes the main body
forked or tripartiteor somewhat nodose-unevenall over, with obtuse tips, rarelysubacute
slightly opaque, constantly hyaline. Hyphae in the regularhymenophoraltramahyaline,
with clamp connections, inamyloid. Coveringlayers:hairsof the pileus melleous-stramin-
eous in KOH,pseudoamyloid,smooth, 5.5-7 p broad,very slightly and graduallyattenuate
towardsthe apex, with walls 1.3-2.5 uthick, with roundedtip, without secondarysepta
or with few distant ones.
On dead branchesand small sticks of dicotyledonous woody plants in subtropical
forest, gregarious,fruitingin the summerrainy season.
MATERIAL STUDIED. ARGENTINA. Tucuman: Rio de los Sosas 27 Jan 1951, Singer T 1140
(LIL, type).

Crinipellissect Crinipellissubsect HeteromorphinaeSinger,Lilloa 8: 509. 1942.

Characterslike those of subsectionStipitarinae,but pleurocystidiadifferentiated.
TYPESPECIES. Crinipellis minutula (Hennings) Patouillard.

Key to the Species of SubsectionHeteromophinae

1. Sporesstronglyelongated,9-17.5(-20) X 3.5-6.3st(see subsectionStipitarinae).
1. Spores up to 9.7p long, or if 9-11 then 6-7.5 p broad.
2. Spores 9-11 X 6-7.5u (C. molfinoana, see note p 25).
2. Spores smaller (cf also C. mexicana and C. foliicola).
3. Spores with Q up to 2; stipe 13-30 X 0.7-1.5 mm 24a. C. commixta var commixta.
3. Spores with Q = 2 or larger;stip 10-17 X 0.5-0.7 mm. 24b. C. commixta varjunia.
Crinipellis 33

24. Crinipelliscommixta Singer in Singer & Digilio ex Singer, Rev. Mycol. 18: 11.
1953. Fig 12.

24a. Crinipelliscommixta var commixta

TYPE..Singer T 1 710 (LIL), from Catamarca,Argentina.
Pileus between "cochin" and "burnt umber"M&P(brown) when fresh and moist
but sometimes somewhat more fulvous in the disc-area,by dehydrationsoon fading to
"dogwood" or "butterscotch"M&P(tan color) with or without a more bister zone
around the central depression,the white ground color often showing between the
strandsof appressedcurly hairs of the wide marginalzone, and likewise appressedly
tomentose-piloseon the disc and with a few erect spinose hairs in the very center of
the depression,or on the central papilla or small umbo, but eventually sometimes
showing a ratherobtuse papilla or small umbo, later depressedaround the latter, and
eventually applanate to concave retainingthe papilla which becomes more evident when
dried, 4-11 mm broad. Lamellaepure white, close or subclose, about medium broad,
intermixed, free or subfree. - Stipe concolorous with the pileus (or reaching"suntan,
Merida"M&P),hairy, above more often appressedlytomentose-pilose,below more
often hirsute. insititious, equal or subequal or taping upwards, 13-30 X 0.5-2 mm.
Context white or whitish, inodorous.
Spores 6.7-9.5 X (3-)4-5.5 p, ellipsoid, few oblong, Q = 1.5-2, mostly below 2,
hyaline, smooth, occasionally a few weakly pseudoamyloid,unicellular,smooth. Hy-
menium: basidia 13-20 X 7-7.5 p, 4-spored;basidioles fusoid; cystidia present, 27-37
X 5-6 p, fusoid to clavate, or subcylindrical,rarelyventricose, entire or sometimes
forked, hyaline or subhyaline:cheilocystidia 17-29 X 4-7.5 1, like the pleurocystidia
or much more frequently with 2-6 short finger-likebranchesor apical appendages,
some branchingoff at the middle, more at the apex of the main body and of very
variablesize and shape, hyaline to pale stramineous,some somewhat opaque because
of slightly thickened wall, numerousand crowded, making the edge heteromorphous
or almost so. Hyphae hyaline, with clamp connections, mainly filamentous, inamyloid.
Hymenophoraltrama regular. Coveringlayers: hairs of the pileus in the wide margin-
al zone long (450 s and longer), melleous-brownishin KOH, some hyaline, all pseudo-
amyloid, smooth, 4-7 , broad, with 1.2-2.8 1,thick wall, with most frequently broadly
rounded or attenuate-obtuse,more rarely subacute to acute tip, many hairs flexuous,
some with few to rathernumerous secondary septa, but not showing a typical ladder-
structure:hairs of the lower portion of the stipe long, 5-7 p broad, pseudoamyloid,
slightly and graduallytaperingto an acute, more rarely obtuse or rounded tip, with
1.5-2.2 p thick wall.
KOH on pileus not provokinga green or gray color reaction.
On fallen woody twigs and branchesand on roots, always on Dicotyledones,
mostly gregarious,fruiting in the rainy seasons (summer or winter). Known hosts:
Acacia sp, Prosopis torquata,Dwialia breviflora.
MATERIALSTUDIED. ARGENTINA. Tucuman:San Pedrode Colalao, 23 Dec 1956,
Singer T 2859 (F); Tapia:6 Jan 1952, Singer T 1710 (LIL, type), T 1707 (LIL, paratype),15
Jun 1951, Singer T 1500 (LIL), probablybelonginghere; Catamarca:Valle de Suncho, Santa
Rosa, 1550 m alt, 18 Jan 1952, Singer T 1759 (LIL, paratype). CHILE.Colcnagua:Pumanque,
27 Jul 1967. W. Lazo PU-124 (SGO).

24b. Crinipelliscommixta var junia Singer, var nov

TYPE. Singer T 2791, from Argentina.
34 Flora Neotropica

A varietate comiimixtadiffert sporis angustioribus. Typus varietatisa R. Singer

Junio mense in Argentina,Tucuman, lectus et in LIL conservatus.
Pileus as in var comitixnta. Lamellaeas in var commixta. Stipe shorter: 10-17
X 0.7 mm.
Spores 6.7-9.5 X 3.5-4.2 p, Q = 2 or larger. Hymenium: cystidia nodose or not,
versiform, opaque, ratherscattered;cheilocystidia as pleurocystidiaor more frequently
with one apical appendageor bifurcate at the apex, sometimes with irregularlylocated
and sized branchlets,often ventricose in the middle or constricted. Hyphae and cover-
ing layers as in var. commtixta.
On dead dicotyledonous branches,fallen twigs, rotting roots etc. fruiting in the
cool, relatively drier winter season on soil with strong salt outcropping,at 400 m alt
approximately(in the plains) among subxerophyticvegetation elements, solitary, or
in small groups.
MATERIAL STUDIED.Known only from the type locality.
ARGENTINA. Tucuman:AroyoMista,15 Jun1955,SingerT 2791 (LIL,type).
In the type variety, I include a winter-fruitingform which may well be another
distinguishabletaxon, possibly on the subspecific level, in which the sporesare partly
pseudoamyloidwhen freshly dried, but relatively broad as in the summer-fruitingforms.
The narrow-sporedwinter-fruitingform which is here describedas var junitiadiffers in-
sofar as the spores are twice as long as broad, or more frequently less than twice as
broad. Also the stipe seems to be shorter. It should be pointed out that the winter-
fruiting Chilean form fruits at a time when there is a maximal humidity-evaporation
ratio, together with a majority of fungi in that region whereas east of the Andes in the
Chaco region (Chaco Santiagueno) winter humidity is considerablylower than summer-
humidity, and the great majority of fungi fruit in summer.
In order to prove that both PU-124 and var juntla,or at least the latter are sea-
sonal forms and as such should be called subspecies, it would be necessaryto gather
more collections of the winter-fruitingforms so that the constancy of the diagnostic
characterscan be re-checked.
Crinipelliscolninixta varjunia may be considereda form intermediatebetween
C. commixta and C n2inutula. It would seem that it is closer to C. conmmixtavar
C. minutula (Henn.) Pat. (Lentinus minuutulus Henn.) is still smaller than C. conm-
mixta varjunia and has still narrowerspores, more frequently a more typical ladder
structureand more often acute tips in the hairs of the pileus. It is a tropical African

Crinipellissect Excentricinae(Singer) Singer, stat nov

Crinipellissect Eu-CrinipellissubsectExcentricinaeSinger, Lilloa 8: 507. 1942.
Crinipellissect CrinipellissubsectExcentricae [sic] Singer,Agaricalesin modernTaxonomy,
2nd e. p 361. 1962.
TYPESPECIES.Crinipellisexcentrica (Patouillard& Gaillard)Patouillard.
Stipe at first central but soon becoming eccentric, mostly relatively short and
distinctly curved, 2-7 mm long; pileus and/or stipe tending to be white or whitish at
first (excepting sometimes the very center of the pileus). The great majority of the
spores smaller than or up to 10.5 p in length and half as broad as long or narrower.
Pleurocystidia at times present. Cheilocystidiaeither ventricose below with long and
relatively thin apical neck, or strongly apically branched-setulose. Surface of the
pileus not greeningor grayingwith KOH 5%;
Crinipellis 35

Key to the Species of Section Excentricinae

1. Pileus7-12mmbroad,withanabruptpapilla,lamellaecloseor subclose. 25. C.excentrica.
1. Pileus3-7mmbroad,withouta distinctpapilla,lamellaedistantor subdistant.
2. Cheilocystidia
ventricosewithwart-likeor shortrodshaped appendages or setulaein
theupperthird.Spores6-8.5 X 5.3-7.71;stipeappressedly villous-tomentose;
reniform.Onmyrtaceous branchlets. 26. C.myrti.
2. Cheilocystidia
ventricosebelow,witha rathernarrowandratherlongapicalneckand
thusampullaceous,the neckoften sparselydiverticulate. Spores8.2-10.5X 4.5-5.5,;
Noton Myrtaceae.
27. C. albipes.

25. Crinipellisexcentrica (Patouillard& Gaillard)Patouillard,Jour. de Bot. 3: 336.

1889. Fig 13.
CollybiaexcentricaPatouillard& Gaillard,Bull. Soc. Mycol. France4: 15. 1888.
TYPE.. Gaillard128 (FH), from Venezuela.
Pileus white with a brown to black punctiform papilla, sometimes with brown or
blackish dots around it, later sometimes somewhat brownish in the wide marginalzone,
silky-pilose to radiallyappressedstrongly hairy in the wide marginalzone as well as in
the center, but on the latter thinly silky and subglabrousor glabrescent,in the margin-
al region of the primordiawith fluffy-woolly, not radiallydepressedhairs, in age sul-
cate over one quarterto three quartersof the radius,convex and strongly papillate
when young, even later not depressedexcepting in dried material,and even then main-
taining a distinct, abrupt papilla, with reniform outline, not or rarely circular, 7-12 mm
broad. Lamellaewhite, close or subclose, narrow,intermixed, free or rounded-subfree
to narrowly adnexed. Stipe ochraceous tawny to cinnamon tawny, often fuscous when
dried, with more yellow-brown apex, short pubescent, later becoming subglabrous,in-
sititious, solid, curved and eccentric in adult caps, equal or taperingup- downwards,
3-6 X 0.3-0.5 mm. Context white, thin.
Spores 7-9.5 X 5-6(-7)u, ellipsoid, more rarely short ellipsoid, smooth, hyaline,
not pseudoamyloid. Hymenium: basidia 19.5-30 X 5-7 u, 4-spored;basidioles fusoid;
cystidia 26-38 X 6.5-8 u, entire or with one to four sterigmatoidoutgrowths at the
apex, opaque and somewhat thick-walled,projectingslightly beyond the basidia, fus-
oid, clavate, or otherwise elongated, hyaline, numerous;cheilocystidia 19-31 X 3.5-8.5
A, the branchesoften spreadingto up to 12 ,, most of them apical, some mediane,
with entire, bifurcate or sterigmatoid-setuloseapex and the setulae sometimes second-
arily knotty-diverticulate,the branchesalso often forked at the tip, most appendages
(of all kinds) obtuse or almost so and more or less finger-like,up to five, rarely only
one, appendagesand branchesup to 12 X 3 1 but usually much shorter and narrower.
Hyphae with clamp connections, inamyloid. Coveringlayers: hairs of the pileus hyaline,
2-5 p broad, with 1-2.5 uthick wall, with mostly rounded tip but some hairs with a
thin-walled,inamyloid or scarcely pseudoamyloid appendage and showing up to 4
clamplesscross-wallsthere, the appendageobtuse or acute, below the appendages
(if these are present) pseudoamyloidin young specimen but in older ones gradually
more and more weakly or moderately pseudoamyloid, sometimes wall appearingto
thicken in 5%KOH.
KOH on dried pileus negative (no color change).
On Bambuseae(and possibly other plants), dead branches,gregarious.
MATERIALSTUDIED. VENEZUELA. Atures (Orinocoregion),A. Gaillard128 (FH,
type). BOLIVIA. Pando: Madre de Dios, Rfo Madre de Dios, near Santa Teresa, 28 May 1956
SingerB 2295 (F).
ILLUSTRATION. Patouillard (1900), fig 70, Singer (1942), fig 1 f, fig 3 d.
36 Flora Neotropica

26. Crinipellismyrti Patouillardin Patouillard& Lagerheim,Bull. Soc. Mycol. France

9: 125. 1893.
Marasmiusmyrti (Patouillardin Pat. & Lag. Saccardo,Sylloge Fungorum11: 37. 1895.
TYPE.Lagerhleim(FH), from Ecuador.
Pileus pallid to pale brownish in dried condition, with appressedhairy squamules,
somewhat striate near margin,convex with incurvedmarginwhich is more villous than
squamulose,with depressedcenter when adult, (dried), but apparentlyat first convex.
3-6 mm broad, in dried condition. Lamellaewhite or whitish, distant, intermixed,
medium broad or rather narrow,almost free. Stipe dark chestnut brown when dried.
with appressedsilky fibrils, eccentric, 1.5-2.5 X 0.3 mm (when dried). Context white.
Spores 6-8.5 X 5.3-7.7 p,, broadly ellipsoid, thin-walledand tending to collapse
(and then often subangular),hyaline, smooth, inamyloid. Hymenium: basidia 40 X
12.3 p,;cystidia none seen; cheilocystidia 22 X 9 p, clavate to subventricose.in upper
third beset with short obtuse setulae, almost echinate ("Rotalis"-type). Hyphae with
clamp connections. Coveringlayers: hairs of the pileus hyaline, the thinner ones often
brownish, 4-8 , broad, with obtusely rounded, rarely acute tip, many with distinct
ladder structurebecause of numerous and moderately crowded secondary septa. only
slightly pseudoamyloid.
On fallen branchesand twigs of Mylrtus sp.
MATRRIAL STUDIED.Known only from the type locality. ECUADOR: Rfo
Pululahua,Lagerheim (FH,type).

27. Crinipellisalbipes Singer, sp nov Fig 14.

Types. Singer M18083 (F), from Mexico.
Pileo albo, cinnamomeo-ochraceo-maculato,lanato-tomentoso, 6-7 mm lato.
Lamellis albis, subdistantibus. Stipite albo, 3-5 X 0.5-0.8 mm, subexcentrico. Sporis
8.2-10.5 X 4.5-5.5 ,. Cystidiis nullis. Cheilocystidiis 25-53 X 4.5-9.5 u, ampullaceis,
apice 7-33 X 1.8-3 p, interdum laxe diverticulato. Pilis longis, obtusis, pseudoamyloideis.
Ad stipulas ligneas in silva pluviali gregatim,Mexico, Singer M 8083 (F).
Pileus white, with a small cinnamon-ochraceousdot above the attachment of the
stipe, finely and loosely wooly-tomentose all over, when wet shallowly sulculate over
three quartersof the radius, dried smooth or nearly so, convex, eventually with de-
pressed center, obtuse 6-7 mm broad. Lamellaewhite, subdistant, narrow, intermixed
didymous, with 12-13 through-lamellae,not intervenose,adnexed to rarely subfree.
Stipe white, finely long-pubescentfrom hyaline hairs from top to base,solid,curved,
subequal, slightly eccentric, more rarely nearly central, central, when quite young, in-
sititious, 3.5 X 0.5-0.8 mm. Context white, thin, inodorous.
Spores 8.2-10.5 X 4.5-5.5 p, ellipsoid to rarely oblong-subcylindrical,with a
slight suprahilardepression, rarely slightly bean-shapedin lateralview, with a Q =
1.7-2.1, more frequently < 2, hyaline, with one or more often two round oil droplets,
rarely without one, inamyloid. Hymenium: basidia 20-26 X 7.2-7.5(-9) p, 4-spored;
cystidia none; cheilocystidia making the edge heteromorphous,25-53 X 4.5-9.5 u,
ampullaceous,with a neck 7-33 X 1.8-3 4, straightor flexuous, eccentrically inserted
or central-apical(more commonly so), smooth or with scattered lateral or apical di-
verticula,sometimes at the apex bifurcate, hyaline, with rounded but sometimes narr-
owed tip. Hyphae hyaline, not gelatinized, 4-13, broad in the trama of the stipe,
with numerous large clamp connections, some partly slightly pseudoamyloid, others
inamyloid; with 1-1.5, rarely 2.5 u thick wall, more thin-wallednear subhymenium
which appearsto be subcellular. Hymenophoraltrama regular. Coveringlayers: hairs
Crinipellis 37

of the pileus long (e g 260 p long), 3-5.5 p broad, with 1.5-2.5 p thick wall, with mostly
slightly attenuated apex but with rounded-obtusetip (there diameter 2-3 1). Between
the hairs occasional (not dense) hyphal outgrowths of the hypotrichium are visible
(the hypotrichium otherwise not strongly differentiated from the trama of the pileus),
these outgrowths cylindrical,ventricose, or conical, 10-13 X 2-5 A; hairs of the stipe
not differentiated from those of the pileus, but still more scattered.
On woody stick (probably monocotyledonous), gregarious.
MATERIALSTUDIED. known only from the type locality. MEXICO.Veracruz:
Near Montepfo,Estacion Biologicade las Tuxtlas, 20 Jun 1969, SingerM 8083 (F, type).
Crinipellissect Grisentinae(Singer)Singer, stat nov
Crinipellissect Eu-CrinipellissubsectGrisentinaeSinger,Lilloa 8: 497. 1942.
TYPE. species. CrinipellismirabilisSinger.
Pileus either normally brown, rust-brown,chestnut, etc. and fading by de-
hydration, or else brightly pink to red, in the former case with pleurocystidia,in the
latter case without or with scarce pleurocystidia, in both cases distinctly turning gray
to green when a drop of 5%KOH or NaOH is applied to the dried surface of the
pileus and/or the hairs are greenish or green incrusted when seen under the micro-
scope in KOH or NaOH mounts.

Key to the Species of Section Grisentinae

1. Freshpileuspink,red or purple;on dicotyledonouswood or leaves.
2. Spores4-5 broad;stipe central.
3. Cheilocystidiacrownedwith severalapicalappendages;on wood. 33. C. rubida.
3. Cheilocystidiawith few if any appendages;on leaves. 32. C. dusenii.
2. Spores4.8-6.8u broad:stipe eccentricand not longerthan the diameterof the pileus.
34. C austrorubida.
1. Fresh Pileus neitherpink nor purple,but chestnut-brownto straw-ocher, rustybrownto
cinnamon-tan;pleurocystidiamoderatelyto quite numerous.
4. On wood or fallen leavesof Iresina,Solanum,Boehrneria,Metahybeand other dicoty-
ledonousvines,shrubsor trees.
5. Spores oblong, 3-3.5p broad:cheilocystidiacrownedby numerousapicalappen-
dages. 30. C. sapindacearum.
5. Spores ellipsoid, 4-6, broad, cheilocystidiaas above but also many (mostly the
majority)entireor simplyforkedor with scatteredbranches. 31. C. tucumanensis.
4. On grassroots, grass leaves,culms, or on other Monocotyledones,particularlyBam-
6. Pileusumbilicate,6-12 mm broad;cheilocystidiaentire or with rarelymore than
three apical appendagesor setulae; hairs partly deep green incrustedin KOH
mounts. 28. C. alcalivirens.
6. Pileus papillate,the papillaoften with an apicalbrush-likestrandof hairs;2-7 mm
broad;cheilocystidiain their majoritywith severalapicalappendages;hairstend-
ing to be greenish or greenish melleous in KOH mounts but not conspicuously
incrusted. 29. C. trichialis.

28. CrinipellisalcalivirensSinger, sp nov Fig 15.

TYPE.SingerB 3510 (F) from Colombia.
Pileo ferrugineo-velcastaneo-brunneo,desiccatione sordide corii-coloriet sub-
pallide crinito, 6-12 mm lato. Lamelliscandidis, confertis. Stipite pallidiusculo, umbri-
nescente, piloso, 12-14 X 0.6-0.8 mm. Sporis 7.5-8.5 X 4.5-5.7 ,. Cystidiis praesentibus.
Cheilocystidiisversiformibus. Pilis pilei Na OK vel KOH ope virescentribus,praesertim
incrustationisviridis causa. Ad Bambuseasin Columbia,Singer B 3510 (F), typus.
Pileus rusty to chestnut brown ("Arab" to "Mohawk"M&P),drying to dull tan
or a brown overlaidby almost pallid or quite pallid strandsof hairs, especially in the
center, pilose all over, with or without a small papilla but if there is one, it is covered
38 Flora Neotropica

by connivent-ascendanthairs, the large marginalzone with more or less appressed,and

the extreme margin with fimbriate-projectinghairs, adult caps, convex, later with a
shallow and small central depression,6-12 mm broad. Lamellaepure white, narrowor
rather narrow, close, mostly free, more rarelyvery narrowlyadnexed. Stipe at first
pallid or at least paler than the pileus, later with umber shades, tomentose-pilose
at the apex which is less pilose than the lower portion and remainspallid for a longer
time, mostly more or less curved, slightly eccentric when mature, insititious, 12-14 X
0.6-0.8 mm. Context white, thin, inodorous.
Spores 7.5-8.5 X 4.5-5.7 p, ellipsoid, Q = <2 hyaline, smooth, inamyloid. Hymen-
ium: basidia 20-22 X 7.5-8.5,, 4-spored;basidioles fusoid; cystidia 28-38 X 5.5-7.5 j,
narrowlyventricose, more rarely subfusoid or almost ampullaceous,opalescent-opaque,
with obtuse tip, with firm wall, without visible contents, numerous on sides and edges
of lamellae;cheilocystidia 20-35 X 6-9 1, similarto the pleurocystidiabut more versi-
form, numerous and crowded, often more clavate, often apically forked, or with 1-3
large apical appendages,these, if present, irregularlyshaped, obtuse or acute, thin to
moderately thickish walled. Hyphae of the tramahyaline not gelatinized, inamyloid,
with clamp connections. Hymenophoraltrama regular,of mostly filamentoushyphae,
Coveringlayers: hairs of the pileus long, 4-5 1 broad, light melleous brownish, but turn-
ing green in the pigmented portion from an incrustingmass of pigment which is deep
green in alkali (NaOH, KOH, 5 or 10%),with 1.5-2.7 p thick walls, with the apex thin-
ner-walledin many hairs, the tip acute, subacute or rounded, most frequently subacute,
secondary cross walls present in many hairs, the septa at a distance of at least 7 'u;
hypotrichium continuing the hairs downwardsbelow the first clamped septum, with
thinner (0.2-0.4 p) wall and graduallynarrowingdownwards,generallysubhyaline,not
dark incrusted from pigment unless the latter is greenishin KOH;hairs of the stipe
7-8.8 , broad, hyaline, subacute, more rarely rounded at the tip, walls 2.5-3.5 p thick,
generallywithout a typical ladder structureand without green pigment incrustation.
The green incrustation, where present, seems to dye the walls of the hyaline hairs pale
greenish in alkali.
KOH on the surface of the pileus: little reaction, in places vaguely greenish, in
others only slightly deeper brown.
On culms of Bambuseaein wet locality.
MATERIAL STUDIED.Known only from the type locality. COLOMBIA: Cundinamarca:
Saltode Tequendama, 24 Jul 1960,SingerB 3510 (F type).

29. Crinipellistrichialis(Leveill) Patouillard,Bull. Soc. Mycol. France 24: 8. 1908.

Fig 16.
AgaricustrichialisLeveille,Ann. Sci. Nat. III, 5: 113. 1846.
Naucoria? trichialis(L6veille)Saccardo,Sylloge Fungorum5: 853. 1887.
TYPE.Zollinger 2078 (FH) from Java.
Pileus fulvous brown to ochraceous brown("Alamo," "feuillemorte," "burnt umber"
mostly with some "Alamo" and the other shades mixed in), in age or dried somewhat
darkerand reachingfuliginous brown in the disc area, in the marginalhalf eventually
some white ground color showing through the strandsof hairs, occasionally the hairs
of the marginalzone whitish also, with entire or fimbriatemargin(from projecting
hairs), in the center more tomentose or short-pilosethan in the wide marginalregion,
and eventually glabrescent,but on the top of the central papilla often or always with a
penicillate fascicle of bristle-likeagglutinationsof long hairs, more or less sulcate on the
margin,campanulate-convexor convex, then flat, with a distinct (but sometimes only in
Crinipellis 39

dried condition and under a lens) dark and abrupt papilla, 2-7 mm broad. Lamellae
cream white to white, narrow,intermixed, subclose to moderately close, at first sub-
ascendant, soon horizontal, free. Stipe"russetbrown," the apex sometimes concolorous
with the center of the pileus or almost pallid, eventually almost bay, tomentose-pilose,
central or more rarely slightly eccentric, with widened but insititious base, 3-9 X 0.3-1
mm. Context white, in the region of the papilla relatively fleshy, otherwise extremely
thin, reviving,inodorous.
Spores 7.3-9.7(-10.2) X 4-6.7 j, mostly 8.5-9 X 4.7-5.7 A,,ellipsoid, few oblong, the
majority with a Q smaller than 2, with or without a suprahilarapplanation,thin- then
thick-walled(0.7 s) and if over-agedoften developinga central or eccentric septum,
smooth, hyaline, inamyloid. Hymenium: basidia 24-33 X 7-8 ,, 4-spored;cystidia 16-49
X 6.8-8.3 t, ventricose, fusoid or clavate, entire, with thin to moderately thickened wall
which is sometimes pale melleous but usually hyaline, entire or with (rather rarely) one
or two apical prongs, in some specimens incrusted by an amorphousincrustation;others
are cystidiole-like and resemble largerbasidioles, these often mucronate or with 1-2
apical prongs, hyaline, with mostly thickish wall, rarely solid. Cheilocystidianumerous
but often with mixed-in basidioles or basidia, 13-48 X 5.5-8.3 ,, varying from being
shaped like the pleurocystidiato more branchedor appendiculateand then ventricose
to clavate with one to severalapical, finger-likeappendages,others with lateral branch-
lets or knots, sometimes constricted, sometimes (rarely) capitate, sometimes incrusted.
Hyphae of the trama filamentous with clamp connections, inamyloid, thin-walled. Cov-
ering layers: hairs of the pileus golden melleous to subhyalineor hyaline, the pig-
mented ones often (but not all of them) turningolive brown, greenish or green in
KOH, but eventually, after long exposure to the alkaline medium becoming pale
greenish melleous, subhyaline or hyaline, not incrusted, 3.5-8(-11)u broad, with a
majority of acute or a majority of obtuse tips, but always both types present, with or
without a ladder-structurefrom secondary septa, the walls 1.3-2.8(-4), thick, often
flexuous, strongly pseudo-amyloid,some hairs at times intermixed with the others and
these more thin-walledand less strongly pseudoamyloid,more often obtuse. Hairsof
the stipe similar.
KOH on the surface of the dried pileus weakly or partiallygreenish,but not con-
stantly so, sometimes merely deeper brown, not black-brown,nor distinctly green or
On dead Monocotyledones, particularlyon culms of Bambuseae(Chusquea,Anrn-
dinaria), and on dead grass culms in subtropicaland tropical forests. Gregariousfruiting
in the rainy seasons.
MATERIALSTUDIED. VENEZUELA.Distrito Federal:Botanic Gardenof Caracas,19 Jun
1958, Deinnis1071 (K, F). BRAZIL.SSoPaulo: Cipo, 1 Sep 1964, Skvortzov& Alim 71393, (NY).
ARGENTINA. Tucuman;Las Lenguas,20 Feb 1951, SingerT1310 (LIL), Singer T 1291 (LIL),Sin-
ger T 1291a (LIL). INDONESIA. Java:Hohnel (FH), Zollinger2078 as "A. trichophorus"(FH, type).
The cystidia of this species are characteristicand generally numerous. In some
specimens true cystidia are formed, whereas in another collection a second, cystidiole-
like, type of pleurocystidiummay be predominant. This species is closely related to the
precedingone and should not be confused with such small species of section Crinipellis
as C. perpusilla,C. stupparia,or C. mexicata or C. commixta.

30. CrinipellissapindacearumSinger, sp nov

TYPE. Brazil:Singer B 3377 (F).
Pileo brunneolo, KOH ope nigricante,piloso, umbonato vel papillato, 3-5 mm lato.
40 Flora Neotropica

Lamellisconfertis. Stipite 9-14 X 0.4-0.5 mm. Sporis 7.5-8.2 X 3-3.5 , cystidiisnumero-

sis. Cheilocystidiis apicaliterappendiculatis. Hypotrichio viridi-incrustato. Ad folia cor-
iacea delapsa Sapindacearum. Typus Singer B 3377 (F).
Pileus brownish hairy (in the color of Panus crinitus), disc with a papilla or umbo
in an umbilicus, and here shorter and thinly fibrillose-pilosewith the papilla subglabrous
or glabrescent,with fimbriate-ciliatemargin,without a setiform fascicle of penicellate
bristle-likeagglutinatedhairs on the papilla, convex, with ratherlow papilla, 3-5 mm
broad. Lamellaewhitish or pallid, becoming brown in the herbarium,close, moderately
broad to narrow,subfree. Stipe concolorous with the pileus, with abundantcurly hairs
projectingup to 0.5 mm, insititious, approximatelythree times longer than the diameter
of the pileus, 9-14 X 0.4-0.5 mm. Context white, thin.
Spores 7.5-8.2 X 3-3.5 p, oblong, smooth, hyaline, inamyloid. Hymenium: basidia
15 X 5.5 u; cystidia numerous 18-24 X 5.3-9(-12)u, at distances of 7-18 ,, clavate or
subclavate,rarely with one or two slight constrictions, hyaline, opaque, entire, inamy-
loid; cheilocystidia making the edge heteromorphoussince they are crowded on the
edge and differentiated from the pleurocystidia,basidiomorphousbut with numerous
apical erect-subdivergentobtuse appendages2.7-7 X 1.2-2.5 p, the entire cell with the
appendagesincluded (7-)11-24 X 5-7.5 u. Hyphae with clamp connections, inamyloid,
those from which the cheilocystidia arise often greenish in KOH, otherwise hyaline.
Coveringlayers: hairs of the pileus long, 3-7.5 u broad, hyaline to light brownish, with
rounded or attenuate-obtusetips, with walls 1-3.5 u thick, pseudoamyloid, the hairs of
the middle of the broad marginalzone and those from the ciliate fibrils of the margin
not differentiated from each other; hypotrichium characteristicallygreen incrusted.
KOH on the dried pileus provokinga blackish reaction but with a greenish-mell-
eous or greenishbackground(from the hypotrichium.)
On dead fallen coriaceous leaves of Sapindaceaeand perhapsother dicotyledonous
families in low tropical rain forest, gregarious. Known host: MetahYlbesp.
MATERIALSTUDIED. Known from the type locality. BRAZIL. Paraibo:Jo5o Pessoa,
11 Jul 1960, SingerB 3377 (F, type).
This species is close to the South Pacific C. mirabilisSinger which has somewhat
largerspores, more abnormalhairs, on the pileus, more conical pileus and grows on dead
twigs. Crinipellissapindacearunm should not be confused with C. foliicola.

31. CrinipellistucumanensisSinger, sp nov

TYPE. Singer T 2158, from Argentina.
CharacteribusC. patouillardii,C. commixtae, C. corticalis gaudenssed KOH op6
oliveoviridans:apice stipitis castaneo-rubidocolore insignis;ad ramos et caules. Typus
in LIL conservatur.
Pileus fulvous-brownbetween "tarragona"and "Mohawk"or between "tarragona"
and "Alamo," between "rust, sorolla br." and "Arab," fading to "bure" or "Wigwam"
on the margin,in the center wet reachingas deep (but not as reddish) as "Mandalay"
(M&P)and fading there to "butterscotch,"the primordiastill entirely color of the adult
disc-region,densely and strongly pilose all over excepting the zone on the sides of the
papillaand around the papilla which is generally much less strongly hairy and tends to
become glabrescent,often with striate-sulcatemargin, often with one or two concentric
furrows around the papilla and there also often less hairy or almost glabrescent,convex,
then applanate, papillate when young and very strongly so when seen dried under a
lens, but in fresh materialthe papilla less striking,and in old material often so indistinct
that it can scarcely be distinguished,with long, agglutinatedbristle-likeerect hairs on
Crinipellis 41

the top of the papilla, but the papilla becoming naked in many cased in old specimens,
even at its tip, the papilla in older specimens often in the middle of a shallow umbilicus
or a slight depression,4-13 mm broad. Lamellaewhite, not cream, remainingwhitish in
age, intermixed-tridymous,almost narrowto (more frequently) broad, 1.2-1.8 mm broad,
close to subdistant, mostly subclose, mostly slightly ventricose, free, more rarelyvery
narrowlyadnexed. Stipe at first slightly paler than the pileus, later "kis kilim" to "co-
coa" or "leaf mold" or 15 E/H 12, M&P,but with a particularlybright reddishchestnut
color ( e g "chutney") at the apex, or even "Maracaibo"or "Java"there, sometimes at
the apex or entirely fading to "butterscotch"(M&P),with more or less appressedrather
dense hairs over most of the surface, but towards the base often with bristle-likehir-
sute hairiness,insititious,,equal, at the apex sometimes vertically furrowed, 13-36 X
0.3-1 mm. Context white, thin, inodorous.
Spores 5.5-8.5 X 4-6 ,, most frequently 7.5 X 4.8ju, ellipsoid, with Q < 2, hyaline,
thin-walled,when over-agedthick-walled,and averaging8.2 X 5.5 A1,inamyloid, smooth.
Hymenium: basidia 26-30 X 6.7-8.8 ,, 4-spored;cystidia moderately numerous,basidio-
morphous, well differentiatedfrom the basidia in phloxine-KOHpreparations,27-45 X
7-10.7 ,, optically "empty," hyaline, smooth and entire, thin-walled, clavate or clavate-
subvesiculose;cheilocystidia 18.7-40 X 5.3-11 sj, crowded at the edges of the lamellae,
simple, forked or branching,most frequently entire at the apex, and then ventricose or
ventricose-subcapitateor subclavate,very rarely crowned with more than two appendages,
but more frequently subvesiculosebelow with a mucro, mostly somewhat opaque, but
not thick-walled,sometimes many or all with a resinousmelleous-hyalineincrustation
or hyaline incrusted at the apex only. Hyphae of the trama hyaline, with clamp connect-
ions, inamyloid. Coveringlayers: hairs of the pileus between center and extreme margin
long and brown, a few among them subhyaline to hyaline, in 5%KOH turninggreenish
or greenish melleous, 4-6 ubroad, with acute, even needle-sharptip or with attenuate-
obtuse tip, hardly ever with broadly rounded tip, the walls 1.3-2.7 u thick, pseudoamyloid,
smooth, with or without secondary septa, but very rarely approachinga ladder-like
structure,occasionally some with acute apical appendage.
KOH on the dried pileus "olive green" M&P. NH4OH negative.
On fallen branches, sticks and woody vines, rotting logs, etc. of dicotyledonous
trees, shrubs or vines in subtropicalforests and in shady wet ravines,fruiting in the
summerrainy season (December until February). Known hosts: Iresina sp (Amaran-
thaceae);Boehmeeriacaudata (Urticaceae);Solatlni sp.
MATERIAL STUDIED. ARGENTINA. Tucuman: Yerba Buena, 12 Feb 1955, Singer T
2158 (LIL, type); Pie del Periquillo, 30 Dec 1951, Singer T 1691. (LIL); Quebrada de Lules, 24
Jan 1951, Singer T 1114 (LIL).
This species was formerly not distinguishedfrom C. patouillardiiand C. conmmrita.
However, the KOH reaction in C. tucumanensisis characteristicand the bright color at
the apex of the stipe has not been observed in C. commixta.
The EuropeanCrinipelliscorticalis (Desmazieres)Singer & Clemen9onis closely
related and differs from C. tucumanensisSinger in darkerKOH reaction, somewhat
largerspores, more scattered fusoid-subampullaceouscystidia, and the host range
(Caprifoliaceaeand Oleaceae).

32. Crinipellisdusenii (P. Hennings)Singer in Pegler, Kew Bull. 21: 518. 1968.
Fig 17.
Marasmniusdusenii Hennings, Bot. Jahrb. 22: 100. 1897.
TYPE.Dusen, from Cameroons(specimen lost).
42 Flora Neotropica

Pileus purple red or bright red ("Artillery"M&P)or so dotted on pallid ground,

but becoming brownish when dried, showing the white ground in the marginalzone, the
browning startingat the center, finely pilose-fuzzy there, with a more appressedlypilose
zone around a subglabrouscentral dot, the marginsomewhat more pilose than the center,
not zonate when fresh but slightly so when dried, sulcate, campanulate,then convex, of-
ten either umbonate or depressedin the center, 3-18 mm, mostly about 10 mm broad.
Lamellaewhite, narrow, more rarely medium broad, intermixed with lamellulae, round-
free or attenuate-subfree. Stipe umber or brown("WoodBrown"Ridgwayaccordingto
Pegler), but sometimes also purple or chestnut when fresh, pilose-hirsute,equal, insit-
itious, 14-40 X 0.5-1.5 mm. Context white, thin, inodorous; taste mild.
Spores 6.3-9 X 3.3-5 ,, mostly 7-8 X 4-5 p, ellipsoid, smooth, hyaline, inamyloid.
Hymenium: basidia 18-27.5 X 6-8 p,, 4-spored;basidioles fusoid: cystidia on edges and
sides of lamellae rathernumerous,(15-)27-37(-39) X (5.5-)6.5-9.5(13.5) ,, clavate, some-
times with one or two appendagesor forked, sometimes constricted in the upper part,
hyaline, inamyloid, opaque, with indistinct inner delimitation of the wall; cheilocystidia
rather numerous and often mixed with basidia, basidiolesand cystidia, i e, not always
making the edges heteromorphic, 15-32 X 3-8.5 p, versiform, often ventricose, fusoid,
cylindric, or subcapitate, mostly entire, more rarely forked or with one or two sterigma-
like appendages,thin-walled,hyaline or pale stramineous. Hyphae hyaline, with thin or
firm wall, with clamp connections, not gelatinized, inamyloid. Coveringlayers: epicutis
formed by hairs which are long and 2.5-7 lbroad, with up to 2.6 , thick wall (but wall
often much thinner in upper portion of hair), green in KOH, with rounded, rarely sub-
acute tip, smooth or vaguely incrusted in places, numerous, simple, rarelywith a secon-
dary septum in the upper portion of the hair, rarely laterally branched,no ladder-like
septation. Similarhairs on the surface of the stipe.
KOH on pileus (dried) glaucous to greenishgray, then green-black.
On fallen leaves of dicotyledonous trees, in Africa and South America.
MATERIAL STUDIED.ECUADOR.Tungurahua, Rio Verde,28 Apr1973,SingerB 7170
(F). GHANA.Tofo, 8 Apr1955,HoldenGC35 (K). ZAIRE.Binga,16 May1927,Goossens-
Fontana 605 (BR); Yangambi,18 May 1939, Louis 14862 (BR).
Pegler(1968),fig 6, la-lf.

33. Crinipellisrubida Patouillard& Heim, Ann. Crypt. Exot. 1: 273. 1928 ("rubidus").
Fig 18.
TYPE. Venezuela (PC).
Pileus bright pink when fresh, vinaceous when dry or dried, with a more dull col-
ored center, scarcely concentrically zonate, with the radialribs strigose clothed, almost
squamulose,with incised-denticulatemargin,convex, then applanateor depressed,with
a slight papilla in the center, reaching20 mm broad. Lamellaewhite, russet when dried.
Stipe concolorous with the pileus all over, strigose-hispid,central or equal, (5-)40-60 X
(0.7-)1 mm. Context white, thin, Odor none.
Spores 7.3-8 X 4-4.8 p, ellipsoid, smooth, hyaline, thin-walled,the over-agedones
thick-walledand with a central or eccentric septum so that the spores become two-
celled before germination,and then often rectangular,inamyloid. Hymenium: basidia
(l-)4-spored; cystidia none seen; cheilocystidia 20-28 X 6-6.7 p, clavate or ventricose,
with a crown of numerous sterigma-likeappendagesin the apical region, these erect or
somewhat oblique. Hyphae inamyloid. Coveringlayers: hairs of the pileus (marginal
zone) long, in bunches of subparallelhairs runningradiallyon the pileus, hyaline to
pale vinaceous in NH40H, 6.5-7 , broad, wall 1.5-2.7 , but thinner at the attenuated
Crinipellis 43

apex which has an obtuse tip.

KOH on the dried pileus green. NH4OH on the dried pileus deeper wine red.
On fallen twigs and branches,decayed wood. Known host: Sweetenia mahogani.
MATERIAL STUDIED. VENEZUELA. Territorio Amazonas: San Fernando de Ature, 23
Jul 1923, Grisol 31 (PC), type (lectotype).
This species has been tentatively identified with a collection by Dennis (1951a,
p430) from Trinidadbut, as mentioned before (Singer, 1955, p 396)this is a different
species whichlhas been named C trinitatis Dennis. This species, unknown to me,
appearsto belong in the section Iopodinae rather than the Grisentinae,and Dennis's
descriptionis repeated there.

34. CrinipellisaustrorubidaSinger, sp nov

TYPE. SinlgerT 917, from Argentina.
Pileo purpureo-rubro;crinibus epicuticularibusKOH ope olivacentibus. Typus in
Pileus purple red (between "canyon" and "rose ebony" M&P),deeper or paler
colored accordingto the density of the fibrils and hairs of the pileus), deeper colored
toward the more densely pilose center, convex, 5-6 mm broad. Lamellaewhite but the
interlamellarspaces pale purplish, distant (about 10 through-lamellaeand 10 lamellulae),
rathernarrow, adnate. Stipe concolorous with the pileus below but white above, fibril-
lose-pilose below, glabrousat the apex, curved, eccentric, about 5 X 0.7 mm. Context
white, inodorous.
Spores 7-8 X 4.8-6.8 1, ellipsoid to short ellipsoid, smooth, hyaline, inamyloid.
Hymenium: basidia 22-35 X 8-9 j,, 1-4-spored;basidioles filamentous;cystidia none;
cheilocystidia 40-48 X 8-9.7 u, versiform,often ampullaceous,some narrowlycylindr-
ical and capitate, etc., not making the edges heteromorphousin all parts. Hyphae inamy-
loid, not gelatinized, hyaline, filamentous, with clamp connections (so in pileus trama
and in the regularhymenophoraltrama). Corticallayer: epicutis of the pileus consis-
ting of hairs which are mostly strongly pseudoamyloid, few weakly so, with rounded tips,
some with an apical callus, with a partial pigment incrustationwhich is lilac in ammonia
but becomes olive green in KOH, with thick (1-2.5 p) wall, with a diameterof about
3-8.5 ,; hlypotrichiumof similarly incrusted,repent,relativelybroad hyphae with num-
erous clamped septa, 4-16.5 p in diameter.
On fallen twigs in subtropicalforest, Northern Argentina. Fruiting in January.
MATERIAL STUDIED. ARGENTINA. Tucuman, ROi de los Sosas, 18 Jan 1950 Singer
T 91 7 (LIL), Type (as C. eggersii).
This species belongs in the same section as the precedingspecies because of the
greeningof the pigment in KOH and the smallerspores, but is externally similarto
C. eggersii(section Iopodinae).

Crinipellissect Iopodinae (Singer) Singer, stat nov

Crinipellis sect Eti-Crinipellis subsect Iopodinae Singer, Lilloa 8: 498. 1942.
TYPE SPECIESCrinipellis iopus Singer.
Pileus bright colored: red, carmin-purple,crimson, lilac, violet, not greeningor
grayingwith KOH or NaOH, but sometimes becoming slowly slate blue with KOH.

Crinipellissect Iopodinae subsect Iopodinae (Singer) Singer, comb nov

(Yitnipellis sect Eu-Crinipellis subsect Iopodinae Singer, Lilloa 8: 498. 1942.
Pileus centrally stipitate and stipe mostly straight,insititious, elongated and more
than three times as long as the diameter of the pileus. Spores either long-fusoid to oblong
44 Flora Neotropica

and not broaderthan 4.7 p, or else ellipsoid and then not more than 4 p broad, Q = 2
or larger.
TYPESPECIES.Crinipellisiopus Singer.
Only neotropical species of the subsection:

35. CrinipellispurpureaSinger, sp nov Fig 19.

TYPE. Singer B 6151a (F), from Colombia.
Pileo purpureo,KOH ope purpureovel atropurpureo,piloso, 6-8 mm lato. Lam-
ellis albis vel subalbidis,confertis vel subconfertis,liberis. Stipite concolori dein brunneo,
insititio, 20-26 X 0.3-1 mm. Sporis 6.5-10.3 X 3-4.7 p. Cheilocystidiisechinatis. Pilis
variabilibus. Ad folia emortua delapsaDicotyledonum. Typus in F conservatur.
Pileus purple red, with dark purple to purple-blackcenter, about "Indian r." M&P
in the largermarginalzone but reaching"maroon" M&P,fading to light reddish brown
with fuliginous center or remainingpurple when dried,appressedlyradiallypilose around
a subglabrouscentral disc or papilla fibrillose-short-pilose,often slightly radiallyrimose
towards the margin when adult, hairs irregularprojectingover the marginand therefore
often ciliate, convex, finally with depressedcenter, in the depressionwith a low circu-
lar elevation around a small umbilicus, or instead of the umbilicus with a small to med-
ium more or less abrupt papilla, 6-8 mm broad. Lamellaewhite or whitish and often
with a slight pinkish gray shade, close to moderately close, moderately broad, didymous,
not collariate, free. Stipe concolorous with the pileus or vinaceous ("roan" M&P),even-
tually becoming deep umber brown from the base upwardsor not changingwith age,
hispid-piloseall over, insititious, equal or subequal, 20-26 X 0.3-1 mm. Context white
or whitish, rather thin, inodorous.
Spores 6.5-10.3 X 3-4.7 p, mostly + 8.3 X 3.8-4 p ellipsoid, oblong-subovoid,
fusoid, sometimes with concave inner side and therefore somewhat curved when seen
in lateral view, vith or without a suprahilarapplanationor depression,smooth, hyaline,
inamyloid, thin-walled, eventually sometimes thick-walledbut not septate. Hymenium:
basidia 16-23 X 5-7 p, (2-)4-spored;basidioles fusoid; cystidia none: cheilocystidia 9-26
X 2-10 ,, with a clavate, more rarely filamentous, cylindricalor ventricose main body,
echinate, sometimes only at the apex with finger-like,erect appendages,sometimes
with many oblique lateral echiniform or spinulose appendagesfrom about the middle
portion, rarely from the base of the main body up to the apex, appendages2-10 X
0.3-2(-2.5)p, with thin to very moderately thickened wall, hyaline, inamyloid; near
the marginof the pileus often only the main body without appendagespresent. Hy-
phae inamyloid, hyaline, non-gelatinized,with clamp connections. Hymenophoral
trama regular,of elongated, interwovenhyphae. Coveringlayers: hairs of the pileus
ratherlong, 2.5-6 p broad, with mostly attenuate-obtuse,but also some with acute or
broadly rounded tips, with 0.3-2 u thick wall which is usually even thinner near and on
the apex, in KOH mounts from purple caps purplishto hyaline-stramineous,in faded
caps pale fuscous or pale brownish, fewer hyaline to fulvous brown, distinctly pseu-
doamyloid, but the upper portion often inamyloid, some with secondary septa, smooth;
hypotrichium and spine-likeagglutinationsof smallerhairs fulvous-melleous,the latter
peg-like and with obtuse tips, the former consisting of melleous fulvous hyphae which
are often finely hyaline granular-incrusted; hairs of the stipe 12-240 X 5-7.5 1, a third
of them rounded at the tip, a third attenuate-obtuse,another third acute, often with
numerous secondary septa.
KOH on the dried pileus, even the faded one, purple to purplishblack, not
greeningnor graying.
Crinipellis 45

On fallen leaves and leaf petioles of dicotyledonous trees in tropical-montane

forests, fruiting in the rainy season. Known hosts: Lauraceae,sp; Melanostomataceae,
MATERIALSTUDIED. COLUMBIA.Valle, Mun.Cali, Saladito, 1800 m alt, 15 Apr 1968,
SingerB 6151a (F, type), BOLIVIA. La Paz: Nor-Yungas,Capellaria,22 Feb 1956, SingerB 1318
(F), Coroico, 7 Feb 1956, SingerB 958 (F), 6 Feb 1956, SingerB 935 (F), 4 Feb 1956, SingerB
860 (F).
Crinipellissect Iopodinae subsect Insignes Singer, subsect nov
Pileo laete colorato, stipite breviorediametro triplici pilei, plerumquecurvatovel
excentrico, insititio vel ad basin fibrilloso. Sporis eis subsectionis antecedentis similibus
vel latioribus. Typus subsectionis:Crinipellisinsignis Singer.
Key to the Species of SubsectionInsignes
1. Saprophytes.
2. Spores3.2-4,ubroad. 38. C. sublividia.
2. Sporesmore than 4,.
3. Cheilocystidiascarce,inconspicuous,3.7-5,ubroad. 36. C. insignis.
3. Cheilocystidiamakingthe edge of the lamellaeheteromorphous. 37. C. eggersii.
1. Parasites.
4. Pileuscrimsonred, "VandykeRed, AcajouRed," R., forming"krulloten"on Theo-
broma or growing from the base of living Vitex. Cheilocystidiasimple or forked;
sporesup to 9(-11), or smaller.
5. On Theobroma;spores7-9(-11) X 4-4.8(-5)u (if sporeslarger- see "3" above).
41. C. perniciosa.
5. On Vitex;spores5-7 X 3-4,. 40. C. trinitatis.
4. Pileus lilac: on other trees, usually fruitingon living,seeminglyhealthy, eventually
rotting cortex of standingbranchesof trees. Cheilocystidiamostly clavate,many or
almost all with 2-6 short apicalappendages;spores9-14 X 4.5-8,. 39. C. siparunae.

36. CrinipellisinsignisSinger, sp nov Fig 20.

TYPE. Singer B 2320 (F), from Bolivia.
Pileo purpureovel purpureo-roseo,radiatimrugoso, minute fibrilloso-pubescente
17-19 mm lato. Lamellisalbis, distantibus,subliberis. Stipite excentrico, usque ad 9
mm longo. Sporis 10.5-13.5 X 4.5-6.5 ,; cystidiis nullis; cheilocystidiis sparsis,angustis.
Pilis moderatim numerosis,debiliter pseudoamyloideis,obtusis. Ad ramulosligneos
emortuos. Typus in F conservatur.
Pileus purple ("India r. M&P")or purplishpink ("rose vale" M&P),the color
concentrated in the hairs which are densest in the center and therefore the center
darkerthan the margin, further towards the marginthese appressedfine hairs forming
a network, all appressedfibrillose-pubescent,on pallid or paler ground, radiallysulcate
along the through-lamellaebut less so on the marginproper, smooth or slightly uneven
on the disc, dry, with oval outline or slightly sinuate on the inner side, not circular,
convex, soon with applanate to shallowly depressedcenter, 17-19 mm broad. Lamellae
white, medium broad to broad, distant (9-10 through-lamellae)with very scattered
veined connections between the through lamellae or the ends of the lamellulaecon-
nected by veins with the nearestthrough-lamella,sometimes forked, young lamellae
without any intervenation,subfree, or very narrowlyadnexed. Stipe yellow and pur-
ple colored, becoming very dark when dried, with scanty hyaline pubescence,general-
ly seemingly glabrouswhen seen macroscopically,curved or straight,eccentric, stuffed,
then hollow, equal or slightly ventricose, seeminglyinsititious, up to 9 mm long, 1-1.3
mm broad; basal mycelium scanty, tomentose, often developed only on one side or
almost invisible macroscopically, but occasionally ascendant upon the lower half of the
stipe, pallid or dirty pallid. Context white, inodorous.
46 Flora Neotropica

Spores 10.5-13.5 X 4.5-6.51, ellipsoid to oblong, and sometimes somewhat

curved when seen in profile, but without suprahilardepression, smooth, hyaline, in-
amyloid, with walls up to 0.5 p thick. Hymenium: basidia 24.5-37 X 9-12 ,, 4-spored;
basidioles subfusoid, with basal clamps;cystidia none; cheilocystidia scattered and in-
conspicuous, 16-51 X 3.7-5 ,, fusoid or fusoid-subcapitateor wavy-filamentous,rarely
subclavate,sometimes with slightly nodulose apex, hyaline. Hyphae of the trama of
the pileus hyaline, inamyloid, with clamp connections. Hymenophoraltrama regular.
Hyphae of the stipe thick-walled,with clamp connections, appearingweakly pseu-
doamyloid in the rind-layer(but this may be due to the pigmentation). Covering
layers: hairs of the pileus relatively scattered, relatively moderately long, 5-8 Abroad
walls 0.5-1 u thick, with rounded tips, weakly pseudoamyloid. Hypotrichiumstrongly
developed around the hair-bases,but not all terminalhyphal cells giving rise to hairs
and then with rounded tips, hyaline, thin to thick (1 u) wall, (5-)9-12 u, broad, strong-
ly purple red incrusted (in NH40H).
KOH on the dried pileus causing darker(purple) spots, no greeningor graying.
On woody twigs in tropical forest.
MATERIALSTUDIED. Known only from the type locality. BOLIVIA. Beni: Vaca
Diez. Riberalta,29 Mar1956, SingerB 2320 (F. type).
This species is characterizedby some primitivecharacteristicssuch as the scat-
tered inconspicuouscheilocystidia, the weakly pseudoamyloidhairs, the distant lam-
ellae, and the slight development of the basal mycelium. Thus, it comes close to
Marasmielluswith which, however, it has no demonstrableaffinities. It is closest to
the following species.

37. CrinipelliseggersiiPatouillardin Patouillard& Lagerheim,Bull. Soc. Mycol.

France 9: 125. 1893. Fig 21.
Maraslius eggersii(Patouillardin Patouillard& Lagerheim)
Saccardo,Sylloge Fungorum11: 37. 1895.
Marasmiusvinosus Spegazzini,Anal. Mus.Nac. Buenos Aires 9: 264. 1909.
Type Eggers(FH), from Ecuador.

37a. Crinipelliseggersiivar eggersii

Pileus purple to violet purple ("India r.," "Aubusson,""canyon," "Anatolia,"
"rose ebony") sometimes with paler margin("Azalea" M&P),fibrillose-short-pilose,
in center often squamulose-fibrillose,becoming slightly to deeply sulcate over 1/3 to
3/4 of radius, convex, eventually often flattened or even with recurvedmargin,de-
pressed or umbilicate in the center, 3-32 mm broad. Lamellaewhite or gray ("cob-
web" M&P),the ground of the lamellae often pale violet ("Hydrangear." M&P)or
up to half-breadthof lamellae concolorous with the pileus, with pallid edges at first,
later sometimes part of edge faintly violet-purpleunder a lens, didymous or tridymous,
at first didymous, distant (with 10-14 through lamellae and > 10 lamellulae),rather
narrow,then broad, adnexed or adnate, and in the latter case often collariately sep-
arating,young without intervenation,old mostly more or less intervenose. Stipe con-
colorous with the pileus below and white at the apex, later deeper (8J 3, M&P),or
chestnut to reddishblack, finely and often sparselyvillous-pruinatewith subglabrous
apex, mostly curved or oblique, often eccentric or becoming so, sometimes subcentral
even in age but then shorter than pileus diameter, taperingupwardsor downwards,
sometimes with bulb, 2-15 X 0.4-3 mm; basal mycelium none or consisting of radiating,
white hyphae, if bulbous often densely strigose. Context white, thin, inodorous.
Crinipellis 47

Spores(7-)1-13(-14) X(4-)5.5-6.3(7.5) u, ellipsoid to oblong, on the inner side

often applanate, without a suprahilardepression,smooth, hyaline (but sometimes taking
up the violet pigment of the surfaces), inamyloid. Hymenium: basidia 22-45 X 8-12u,
(2-)4-spored or 1-2-3-4-spored,always mostly 4-spored;basidioles fusoid-filamentous;
cystidia none; cheilocystidia on and very near the edges numerousbut not everywhere
makinlgthe edges heteromorphous,rising from hyphae runningalong the edge, 22-58 X
2.8-17.3 u, mostly 25-42 X 8-9 u, versiform, most frequently ampullaceous,fusiform,
cylindric or clavate (and then often forked), obtuse or mucronate, sometimes with
claviculate, subcapitateor capitate apex, rarely filiform or subvesiculose,rarely with
1-2 short lateral or apical outgrowths, very rarely with severalfinger-likeoutgrowths,
but then these not all apical, hyaline to pale stramineous. Hyphae of the trama of
the pileus hyaline, inamyloid or few vaguely pseudoamyloid, with clamp connections,
not gelatinized, mostly filamentous. Hymenophoraltrama regular. Coveringlayers:
hairs of the pileus relatively moderately long but many up to 200 u long, 4.5-9(-12.5)
p broad, with rounded tips, with hyaline or stramineouswalls and at least in lower
portion often violet or purplisliincrusted by the pigment, pseudoamyloidbut some-
times slowly and weakly so, rather scattered to numerous but not very dense, with
1-2.5 p thick wall; hypotrichium consisting of repent fascicles of hyphal cells with
numerousclamp connections, strongly lilac or violet incrustedby the pigment when
seen in NH4OH mount, the pigment tending to turn greenishin H2SO4 and KOH
KOH reaction on dried pileus negative or becoming slate blue.
On dead branchesand sticks, coriaceous petioles and rotting logs in subtropical
forests, tropical-montaneforests, plantations and parks, always on Dicotyledones,
fruiting in the rainy seasons.
MATERIALSTUDIED.ECUADOR.Guayas,Balao, 26 Jan 1892, Eggers(FH, type); Napo,
Lago Agrio, 16 May 1973, SingerB 7504 (F). BOLIVIA. La Paz: Nor-Yungas:Rio Llolosa,
31 Jan 1956. SingerB 779 (BAFC);Rio Yariza,16 Feb 1956, SingerB 1181 (BAFC). ARGEN-
TINA. Tucuman,Quebradade Lules, 18 Dec 1951, Singer T 1647 (LIL);Capital,ParqueRoca,
Spegazzini(LPS, type of Il vinosusSpeg.).
One of the Bolivian specimenshad gray lamellae but was otherwise not much
different from the rest.
The purple red color of the fresh pileus mostly changes to a deep violet on dry-
ing (for example 48 H 3 to 56 H 2, M&P). On the other hand, var epiphyllus and var
lilaceiceps are violet or lilac when fresh.

37b. Crinipelliseggersiivar flavipes Singer, var nov

TYPE. Singer B 2310 (LIL, BAFC), from Bolivia.
Stipite partim flavo, ceterum var eggersii similis. Typus in LIL conservatur.
Pileus purple when fresh,"Anatolia""Mirador"with "Domingo" margin(M&P),
very dark when dried, vaguely to distinctly sulcate, fibrillose, pilose, even hispid-pilose,
the hairs mostly subhyaline on the margin,convex, mostly with a small, central umbo,
often in a central depression,circularto oval in outline, 4-10 mm broad. Lamellae
white, often with purplish-lividground, ratherbroad, young not intervenose,later
intervenose, distant, with entire edge but under a lens somewhat fimbriate from the
cheilocystidia, didymous, with about 11-12 through-lamellae,subfree to adnate and
in the latter case soon collariate-septating. Stipe white or yellowish-whiteat the apex,
orangeyellow, yellow or yellow brown at the base, with scattered hyaline hairs or
above subglabrous,tomentose below when seen under a lens, subcentralto eccentric,
48 Flora Neotropica

seemingly insititious or not insititious, often curved, solid, subequal or with a basal
bulb (3-5)5-13 mm long, 0.8-2 mm broad; basal mycelium at times scarcely visible,
but sometimes distinct and strigose or partiallyascendant on the lower part of the
stipe and then tomentose-vellereousand pallid. Context inodorous.
Spores 10-13.7 X 5.5-6.2p1,ellipsoid, about twice as long as broad, with thin
to slightly thickened wall, without suprahilardepression, not curved, hyaline, inamy-
loid. Hymenium: basidia 18-28 X (7-)8-9E; basidioles subfusoid or subcapitate;
cystidia none; cheilocystidia 25-90 X 7-25 I,, mostly 30-50 X 12-14 p, ampullaceous
with an apex 25-50 X 2.5-7 1, cylindric or subulate and obtuse, often capitate (cap-
itulum up to 12 ,u broad), sometimes oblique or with slight constriction(s), from one
third to equal in length as compared with the ventricose base, hyaline, with thin to
thick wall, 0.3-4 I some with a colorless, granular,thin incrustation,inamyloid,
making the edge of the lamellae heteromorphous. Hyphae thin-walledwith clamp
connections, hyaline, inamyloid. Coveringlayers: hairs of the pileus scattered or in
bunches, moderately long (35)60-200 X 5.5-12.5(-17)lp, irregularlyshaped or filament-
ous, some subventricoseor capitate, but simple, with attenuated or equal apex and
obtuse to rounded tips, with 1-1.7 , thick walls, in the lower portion hyaline to pale
violet in NH40H, more hyaline towards the tip, slowly and moderately strongly
pseudoamyloid(pinkish cinnamon in the Melzer reagent), in the lower portion often
brown or hyaline incrusted;hypotrichium with a strong violet intraparietaland in-
crusting pigment.
KOH on the dried pileus no color change, eventually slate blue or black, H2S04
On dead cortex of standing trees and on fallen branches,always on Dicotyledones,
fruiting in the rainy season. In the inundated as well as high tropical rain forest, in
the inundated forest above the high-waterlevel (to 3 m above ground).
MATERIAL STUDIED. ECUADOR. Napo, Lago Agrio, 17 May 1973, Singer B 7527a
(F) BOLIVIA. Pando: Madre de Dios, 28 Mar 1956, Singer B 2310 (LIL, Holotype; BAFC,
isotype of variety); Beni: Vaca Diez, Riberalta 31 Mar 1956, 170 m alt, Singer B 2378 (F).
Crinipelliseggersiivarflavipes Sing. may be merely a form of the type variety
since it has once (B 7527a) been found side by side with specimens showing no yel-
low pigment on the stipe. On the other hand, the following varietiesappear to be
quite constant.

37c. Crinipelliseggersiivar epiphyllus Singer, var nov

TYPE. Singer B 747, from Bolivia.
Carpophorisepiphyllis;multis cheilocystidiis appendiculatis. Typus in LIL.
Pileus lilac, 3-5 mm broad. Lamellaewhite, distant. Stipe white, strigoseto-
mentose near and at base but subinsititious,subglabrousabove, curved, 2-4 X 0.4-0.6
mm. Context white, inodorous.
Spores 9-13.8 X 5.5-7.5 ,, ellipsoid to ellipsoid-oblong,smooth, without suprah-
ilar depressionbut inner side often entirely applanate,hyaline, inamyloid. Hymenium:
basidia(2-)4-spored, about 37 X 11 ;;cystidia none excepting some cheilocystidia
near the edge; cheilocystidia numerousand making the edge heteromorphicin mature
specimens, 27-58 X 2.8-10.3 ,, of two types, (1) ampullaceouswith often claviculate
"neck" (claviculate ones 2.7-6.2 p across at apex; constriction beneath 1.5-3.5 pacross,)
more rarely cylindrical,(2) versiform,most with finger-orknot-like excresences, both
apically and laterally, rarely without these and then somewhat smaller than type (1)
and neither cylindric nor ampullaceous. Hyphae of pileus trama and hymenophoral
Crinipellis 49

tramahyaline, thin-walled,inamyloid, with clamp connections, but many secondary

(clamp-less)septa present. Coveringlayer: epicutis of the pileus consisting of hairs
which are hyaline or slightly fuscidulous inside or in the wall, some with short lat-
eral excrescences, with obtuse tips, pseudoamyloud,accompaniedby some shorter
On dead fallen leaves and leaf petioles of Theobroma(cacao plantation) in
the Yungas, Bolivia.
MATERIALSTUDIED. BOLIVIA. La Paz, NorYungas,Charobamba30 Jan 1956, Singer
B 747 (LIL).

37d. Crinipelliseggersiivar lilaceiceps Singer, var nov

TYPE. Singer B 7316, from Ecuador.
Pileo violaceo vel lilaceo; multis cheilocystidiis appendiculatis. Ad ligna in Aequ-
toria. Typus in F conservatusest.
Pileus violet to lilac ("Corinth,"47 J 1, 48 L 2, M&P),appressedlyfibrillose and
very finely pilose, glabrescent,in age often with a reticulate appressedfibrillosity which
is lilac (on white ground), coarsely sulcate over 5/6 to 6/7 of the radiusof mature
specimens, convex, umbilicate, 13-27 mm broad. Lamellaewhitewith pallid edges and
mostly purplish-violetground color or purplishviolet in the upper half, rathernarrow,
then rather broad but not ventricose, distant (about 7 through-lamellaewhen mature),
intervenosewhen mature, adnexed to adnate, often collariate. Stipe chestnut with
white apex, macroscopicallyappearingglabrousbut finely pilose with scatteredhairs,
the apex often glabrous,subcentralto somewhat eccentric, mostly curved, subequal
or taperingupwards, 5-6 X 1-2 mm; basal mycelium present, consisting of minute
white radiatingfibrils. Context white, thin; odor none.
Spores 10-13 X 6-6.5 p ellipsoid, smooth, hyaline, inamyloid. Hymenium:bas-
idia 32 X 8.5-10.5 p, 4-spored;cystidia none; cheilocystidia rising from hyphae running
along the edge, these intermittent but numerous, 15-55 X 6-17j , either ventricose or
cylindricalor ampullaceousto ampullaceous-ventricoseand with a constriction under-
neath a clavate, subcapitate or capitate apex; apex, if ampullaceous,3.5-8 p across,
sometimes forked with two such narrowedapices, constriction, if present, 4-9 Mwide,
capitulum, if present, 7.5-12 j in diameter, sometimes with 1-5 lateral (rarely some
apical) short-finger-likeappendages,but the vast majority entire, sometimes rising from
a vesiculose basal cell 10 1 diam, hyaline, thin-walled. Hyphae inamyloid, with clamp
connections. Coveringlayers: epicutis of pileus consisting of sparseto rathernumer-
ous but not dense hairs, these long (some short) and equal, with about 1 p thick,
slowly pseudoamyloid wall, with rounded tip, 4.5-6.5 p in diameter;hypotrichiumcon-
sisting of numerousappressedhyphal cells which are catenulate and broad, cylindric
to ventricose, hyaline but incrusted by a strong purple pigment-incrustation,with
thin to firm wall, 5-15 broad. Stipe likewise covered with pseudoamyloidhairs.
KOH on dried pileus without reaction.
On dead, fallen branchesof dicotyledonous trees in hylaea.
MATERIALSTUDIED. ECUADOR.Napo, LagoAgrio,9 May 1973, SingerB 7316 (F) type.
This variety appearsto be intermediatebetween Crinipellissiparunaeand Crini-
pellis eggersii. It differs from the first of these species in the predominantlyentire,
smooth cheilocystidia which are ampullaceousto subcapitateor capitate and in its
habitat on dead wood (which does not necessarilymean that the mycelium can never
be parasitic). It differs from var epiphylla in its habitat and color and from var egger-
sii in its color and presence of some appendiculatecheilocystidia.
50 Flora Neotropica

38. CrinipellissublividaMurrill,North AmericanFlora 9: 287. 1915.

TYPE. W.A. & Edna L. Murrill(NY), from Mexico.
Pileus "Indian Red" to "Light Vinaceous Lilac" (Ridgway), paler towards the
margin"Pale Vinaceous Lilac" (Ridgway), slightly tomentose, convex, margin striate,
sulcate when dry, 20 mm broad. Lamellaepale lilac, rather narrow,distant, very
narrowlyadnexed or free. Stipe slightly paler than the pileus, hairy, white tomentose
at the base, and a white mycelium coating the substratum,not insititious, 20 X 1.5
mm. Context moderately thin.
Spores 8.7-11(-14) X 3.5-4(-5) ,, subcylindrical,some slightly curved, Q > 2,
smooth, hyaline. Hymenium: basidia 29-35 X 7.5 j,; cheilocystidia 21-29 X 8-10 1,
clavate or with irregularprojections all over or only at the apex, more rarelyampull-
aceous or capitate, often forked, often asymmetric, walls thicker than those of the
basidia, sometimes up to 1 jpthick. Hyphae thin-walled,with clamp connections.
Coveringlayers: hairs of the pileus in minute tufts, often short or irregularlythickened,
all pseudoamyloidand with rounded-obtusetips. Hypotrichiumof hyphae which are
incrustedby a (now) brown pigment.
On dead branches,fruiting in January in virgin forest.
MATERIALSTUDIED. Known only from the type locality. MEXICO.Veracruz:
Motzorongonear Cordoba,15 Jan 1910, I'. A. & Edna L. Murrill1047 (NY, type).
The above description is based on a study of the type, published first in Singer
This species differs from the precedingone in the relativelylonger stipe which
is less or not eccentric and has a distinctly tomentose basal mycelium and also in the
narrowerspores. It has not been re-collected in Mexico, probably because the virgin
forests of Motzorongo no longer exist.
Crinipellispalmarum,from Africa, differs in having subclose lamellae, insititious
stipe and slightly smallerspores.

39. CrinipellissiparunaeSinger, Lilloa 8: 505. 1942.

TYPE. Singer, from a greenhousein U.S.S.R.
Pileus lilac, brownishlilac, livid pink, at first densely covered by tufts of hairs
which later become sparserand eventually leave pallid spaces between them so that
old caps, as a whole, look much paler than young ones and the tufts of hair becoming
squamulose(aspect often of Lepiota), dried sometimes dirty brownish melleous or
olive melleous, or else livid pallid, but in some specimens maintainingthe original
color, the outer two thirds of the radiuscostate-sulcateto vaguelysulcate in age, con-
vex, umbilicate or umbonate or neither, but color and surface charactersratheruni-
form, eventually flattened in part or all over, 6-22 mm broad. Lamellaewhite, nar-
row, later broad, reaching4 mm broad in the largest carpophores,some forked or
indistinctly rugulose-anastomosingin some specimens, in old specimens usually inter-
venose, medium close to distant, up to 14 through-lamellaepresent, adult tridymous
with many lamellulae, rounded-adnexedbut frequently soon separatingwith a free
collar. Spore print pure white. Stipe at first white, later brown with white apex,
eventually often entirely brown, dry, opaque, finely tomentose to tomentose-squam-
ulose, with less pilose-tomentose apex (which may be glabrescent),solid, mostly cur-
ved, equal, 3-14 X 0.8-2 mm. Context white, in the center relatively thick; odor none;
taste mild.
Spores 9-14 X 4.5-8,, mostly 9.5-12 X 5.5-6.5 r, ellipsoid, Q = 2 or somewhat
less, hyaline, smooth, inamyloid. Hymenium: basidia 33-40 X 6.8-7.8 z, (2-)4-spored;
Crinipellis 51

cystidia none; Cheilocystidia 16-30 X 5.5-11 p, clavate, rarelyventricose or subcylin-

dric, apically beset with rod-shapedor phialid-shapedto spinose (most frequently
rod-shaped)short appendages,mostly 2-6 (and most frequently 5-6) of them, and
these 3-5.5 X 1.5-1.7 p. in certain places also similarelements without appendages
visible and then main body usually 4-6.7 p broad, all hyaline, or some melleous-hy-
aline, inamyloid. Hyphae of the trama 5-10 p broad, filamentous, inamyloid, with
clamp connections. Hymenophoraltramaregular,hyphae interwoven. Covering
layers: hairs of the pileus not continuously crowded, usually about 25-150 X 4-11 u,
pseudoamyloid,sometimes abruptly thickened or deformed, with rounded tip or
attenuate-subacute,with walls 1.7-4 p thick, smooth.
Mycelium(as generallyalso carpophores)on living trees, cortex of branches
and twigs, often high above ground, in tropical and subtropicalforests and in green-
houses. In the greenhousesfruiting in June. Known host: Siparunasp.
MATERIALSTUDIED. CUBA. Wright858, as Marasmiusscalpturatus(ined.), (FH).
BRAZIL. Rio Grandedo Sul: Nova Petropolis,1925, Rick 477 (FH). U.S.S.R.: Leningrad:
Orangeriesof the BotanicalGarden,10 Jun 1936, Singer (LE, type).
This differs from C. perniciosa in the shape of the cheilocystidia, the color of
the pileus and the fruiting on living, but not deformed hosts other than Theobroma;
also the stipe does not tend to become yellow and the spores are largerthan those
of C. perniciosa.
It differs from Crinipelliseggersiiand its varieties by growing generallyon
living host material,by apically multiappendiculatecheilocystidia which may or may
not be intermixed with entire ones.
ILLUSTRATION. Singer (1942) fig 3e.

40. Crinipellistrinitatis Dennis, Kew Bull. Add. Ser. 3. Fungus Flora of Venezuela
and Adjacent Countries,p 465, 1970.
TYPE.Deiinis 376 (K), from Trinidad.
Pileus "Vandyke Red" to "Acajou Red" (Ridgway), clothed with short, sub-
erect hairs, with undulatingand indistinctly striate margin,convex with a small um-
bilicus, then flattened and often depressedat the center, up to 25 mm broad. Lamellae
"vinaceousbuff" with reddish edges, ratherclose, didymous, adnate. Spore print
pure white. Stipe "Acajou Red" (Ridgway), darkerand brownerbelow, downy, cy-
lindric or slightly enlargedupwards,solid, (14-23 X 1.5-2.5 mm). Context "Vinaceous
Buff" (Ridgway), dark brown on the base of the stipe, thin, thicker and brownerat
the disc of the pileus.
Spores 5-7 X 3-4 1, ellipsoid, inamyloid, [smooth]. Hymenium: cheilo-
cystidia 30 X 8 1, cylindricalor irregularand forked, thin-walled,red. Coveringlayers:
hairs of the pileus about 300 p long, 5-6 broad, more or less hyaline, pseudoamyloid
(deep red in Melzer'sreagent) incrustedwith light red granuleswhich dissolve in 10%
KOH solution to give a bluish purple tint, distantly septate, undulating,ratherthin-
walled: hypotrichium formed by polygonal thick-walledcells, walls red in Melzer's
On the bark of living trees, gregariouson base of Vitexsp. trunk.
MATERIALSTUDIED. Known only from the type locality. TRINIDAD. Path to
Maracaswaterfall,25 Nov 1949, Dennis 376 (K, type), not studied.
This species appearsto belong in subsectionInsignes, and on account of the rel-
atively small spores, it is different from all other species includingC. rubida (sect.
Grisentinae),with which it was originally- i e by Dennis 1951a p 430 - thought to
52 Flora Neotropica

be identical, but after the latter species was revised by Singer(1955), it became clear
that "C. rbidus sensu Dennis (1951a) is obviously a different species" which was then
redescribedas C. trinitatis by Dennis (1970). Since I have not restudiedDennlis376,
the description given above is an adaptation of Dennis's (1951a) description(data
derived from the illustrationin brackets).
The cheilocystidia are obviously different not only from those of C. rubida but
from those of C. siparuiae and C. perniciosa as well.
ILLUSTRATION. Dennis (1970), pl 5, fig 4.

41. Crinipellisperniciosa(Stahel) Singer, Lilloa 8: 503. 1942.

TYPE.Stahel, from Surinam (collections in Paramaribo?), not seen.
Marasmius Stahel,Dept.Landbouw
perniciosus in SurinameBull 33: 16. 1915.
Pileus crimson red, generally faintly so accordingto Stahel, later bleachingto
whitish and dried materialoften whitish, with a red-blackspot in the center, and with
others which are radiallyarrangedand have the same color and are visible on dried
material, radiallygrooved, campanulate,later expanded and often with a depressed
center, 2-25 mm broad, mostly 5-15 mm in diameter. Lamellaewhite in dried mater-
ial, rather thick (0.2 mm) in mature material,medium broad to ratherbroad (1-2 mm)
distant, (8-20 through-lamellae,mostly + 15), collariate. Spore print pure white. Stipe
white except at the thickened subbulbousbase which is light lemon yellow, later lemon
yellow to white above and dark brown red below, almost naked, but white pubescent
from the mycelium at the base, holow or becoming so (lumen 0.3-0.4 mm broad), 5-10
X 0.4-0.7 mm. Context very thin (for example 30-40 u accordingto Stahel) in the
pileus, reviving.
Spores 7-9 X 4-4.8, (from print), but accordingto Stahel up to 11 X 5 ,, ellip-
soid, hyaline, smooth, inamyloid. Hymenium: basidia 31-32 X 7-(8.8),, 4-spored;cystid-
ia none; cheilocystidia 35-50 X 9-14p, rather regularlybottle-shaped. Hyphae with
clamp connections. Coveringlayers: hairs of the pileus not numerousand dense, most
numerous on the center portion of the pileus, fresh with a red, thick wall, hyaline when
dried or from old carpophores,always rounded-obtuseat the tip, pseudoamyloid, 80-150
X 4-12,; hypotrichium consisting of thick-walled,inamyloid hyphae.
The mycelium infecting the living plants of Theobroniabut forming the fruit
bodies mostly on dead "krulloten"(witch-broom deformations)and on pods and in-
durated fruits of this genus, the mycelium causing the formation of the krulloten.
Fruits during wet weather. This is pathogenic on cacao trees, apparentlyoriginally
only in northern South America, but the disease has spreadto whereverTheobrolia has
been planted, from Ecuador to the West Indies. Known hosts: Theobromacacao, T.
bicolor, T. speciostum(a wild species).
MATERIALSTUDIED. SURINAM. Paramaribo:Stahel (FH), authentic,additionalmater-
ial (FH). BRITISH GUIANA.Coil.& det.F. L. Stevens(FH).
Data on the infection and the effect on the host as well as suggestedcontrol
measuresshould be looked up in Stahel's originalpaper or in the short digest given by
Singer(1942). The fact that the fungus causing the "krulloten disease"is still fre-
quently quoted asMarasmiusperniciosus, 30 years after its transfer to Crinipellis,is a
good illustration of the "conservatism"of some phytopathologistsand their reluctance
to adopt the results of mycological work.
Chaetocalathus 53

ILLUSTRATIONS. Singer(1942), fig 2 a; see alsoStahel(1919) pl 1-8;(1915), pl 1-12;

Dennis(1970), pi 3, fig 4.

2. ChaetocalathusSinger, Lilloa 8: 518. 1942.

TYPESPECIES.C. craterellus(Durieu & L6veillV)Singer.
Habit pleurotoid; pileus with thick-walledpseudoamyloid(to almost amyloid)
hairs which are long and distinctly separatedfrom the trama by a hypotrichial layer;
hymenophore well developed, lamellar;spores hyaline, thin-walled,eventually often
becoming somewhat thick-walled,occasionally with a septum, smooth, inamyloid,
sometimes pseudoamyloid, but never amyloid, cyanophilic;basidia without siderophilous
granulosity,4-spored, or more rarely with an inconstant immixture of 1-3-sporedones;
cystidia on the sides of the lamellae often present and either entire (and then incrusted
by a crystalline incrustation)or variously divided or forked, very frequently pseudo-
amyloid; stipe rudimentaryor more rarely absent, not directly attached to the sub-
stratum;cheilocystidia always present; tramainamyloid; hyphae with numerousclamp
connections. On wood, bamboo stalks, bark, leaves and stems.

Key to the Sections, Subsections,and Species of Chaetocalathus

1. Pseudoamyloidseta-likeor metuloid sterile bodies in the lamellaeabsent;only inamyloid
cystidiaand/or cheilocystidiapresent.[Not representedin the neotropics]. sect Chaetocalathus.
1. Pseudoamyloidsetoid or metuloid bodies in the lamellaepresent,rarelyconfinedto the
edge of the lamellae(in young specimens)and intermixedwith inamyloidcheilocystidia.
2. Pseudoamyloidbodies relativelythin, seta-like,acute,branched. sect Meristocystis,p 53.
1. C. columellifer.
2. Pseudoamyloidbodies reachingmore than 6ji broad, metuloid, or restrictedto the
edge (at least in young specimens)and cystidioid, simpleor 2(-4) forkedat the apex
and reachingover 7p diam. sect Holocystis,p 54.
3. Pigmentnone, only old specimensmay assumea creamcoloredtinge.
4. Cystidiathickwalledmetuloid, pseudoamyloid,often thicklyincrusted,on the
side of the lamellae, numerous, all simple and entire; cheilocystidiaoften
apicallyforked.Commonneotropicalspecies. 2. C. liliputianus.
4. Cystidianot as above; cheilocystidia,often forked, but also entireand merely
appendiculateor muscronatealong the edge, later pervadingthe sides of the
lamellae. 3. C. niduliformis.
3. Pigmentpresent;pileus coloredfrom the beginning.
5. Sporesrelativelynot broad: 9.5-10.5 X 5-5.3. Pileusgrayish-leather-color. On
livingMurraia. 4. C. asperifolius.
5. Sporesrelativelybroader;pileuspurplered or orange.
6. Pileuspurplered. 5. C. carmelioruber.
6. Pileusdull orange. 6. C. aurantiacus.

Chaetocalathussect MeristocystisSinger, Lilloa 8: 522. 1942.

Charactersas indicated in the key above.
TYPESPECIES.C. africanus (Patouillard)Singer
Only one species is known from the neotropics.

1. Chaetocalathuscolumellifer(Berkeley) Singer, Sydowia 9: 398. 1955. Fig 23.

AgaricuscolumelliferBerkeley,Hook. Journ. Bot. Kew Misc.4: 161. 1852.
Marasmiuscolumellifer(Berkeley)Pilat, Trans.Brit. Mycol. Soc. 33: 219. 1950, invalid.
TYPE. Borneo (K).
Pileus white or more rarely yellowish white, strongly radiallypilose-woolly or
woolly-subsmoothwith substrigosehairs, which are up to 0.2 mm long, but become
applicate in older specimens, the margin woolly tomentose to slightly crenulate-fim-
54 Flora Neotropica

briate, dry, often with projecting sterile margin,convex, attached with some central
to sublateralareaof the sterile surfaceand appearingcup-shaped,circularto sub-
elliptical, sometimes with a short pseudostipe, 1-8 mm broad. Lamellaewhite, dry pale
buff-tan (to umber when dried), rather narrowto medium broad, intermixed with
lamellulae when mature, moderately close to distant, concurrent to the stipe rudiment
to which they are adnexed or adnate, sporeprint white. Stipe only rudimentaryand
function-less,columelliform conical to knob-shaped,often obsolete, tomentose, white,
up to 1 mm long. Context extremely thin, white. Odor and taste none.
Spores 7-10.2(10.5) X 4.5-6.8 u, ellipsoid, smooth, thin-walled,becoming thick-
walled and often also with one cross septum when over-aged,inamyloid or pseudoamy-
loid (at first inamyloid in fresh material),hyaline in NH40H. Hymenium: basidia 26-
26.5 X 8-10 , 4-spored;cheilocystidia long, almost hair- or seta-like, occupying the
edges of the lamellae, but (as pleuroscystidia)also here and there appearingon the
sides of the lamellae, but may be absent in some sections, 31-90, long, entire or forked,
especially near the apical region, narrowlyfusoid to needle-shapedor cylindric, but
often with a ventricose swelling below, tip and branch-tipsacute or subacute, hyaline,
thick-walled,wall 1.5-1.7 p thick and psuedoamyloid, diameter 2-4.7 p but in basal
swellings where these are present reachingup to 7.5 p across, often deep-rootingwith-
out a septum and then reachingup to more than 100 p in length. Hyphae of the reg-
ular hymenophore hyaline, inamyloid, with some inflated cells intermixed, with clamp
connections. Coveringof the pileus consisting of usually simple, rarely forked thick-
walled pseudoamyloid hairs with a diameter of 2.7-7 p, either slightly incrusted or non-
incrusted, thick-walled,but wall somewhat thinner near apex, if the latter is attenuate
or acute, but some hairs with obtuse tip, always dense and long. Tomentum of the
stipe rudiment similar,hairs intermediatebetween those of the pileus and the hymenial
On palm fragmentsand frondose twigs fallen to the ground, gregarious,fruiting
in the rainy seasons in the lowland forests.
MATERIALSTUDIED. PANAMA. BarroColoradoIsland, 30 Jun 23 Aug 1952, Martin
& Welden7279, 8562, 8580, 8682 (F, IA). BOLIVIA. Beni, Vaca Diez, Ivon, 3 Apr 1956, Singer
B 2481 (LIL). INDONESIA. Java;south coast, Perigi,Donk (L, 11554). BORNEO. (K), type.

Chaetocalathussect Holocystis Singer, Lilloa 8: 526. 1942.

Characters,see key above.
TYPE SPECIES. C. carnelioruber Singer,

2. Chaetocalathusliliputianus(Montagne)Singer, Lilloa 8: 527. 1942.

Fig 24, 25, 26.
AgaricusliliputianusMontagne,Ann. Sci. Nat. IV. 1: 99. 1854.
Pleurotusliliputianus(Montagne)Saccardo,Sylloge Fungorum5: 385. 1887.
Marasmiusnidulus Berkeley& Curtis,Jour. Linn. Soc. 10: 299. 1868.
Crinipellisnidulus (Berkeley& Curtis)Patouillard,Jour.de Bot. 3: 336.1889 ("nitidulus").
CrinipelliscalosporusPatouillardin Duss, Flore Cryptogamiquedes Antillesfrangaises,
Champignons,p 255. 1904.
Marasmiuscalosporus(Patoulliard)Saccardo& D. Saccardo,Sylloge Fungorum17: 47. 1905.
PleurotopsiscalosporaMurrill,North AmericanFlora 9: 238. 1915.
TYPE(of Agaricus liliputianus).Leprieur, from French Guyana (not seen).
Pileus white, floccose-pilose to silky-pubescent, with almost entire margin,which
is usually inflexed on drying, attached to the substratumby a central to sublateralarea
and appearingcupuliform, smooth or with sulcate margin, sometimes with a short
pseudostipe, convex to almost resupinate, 1.5-10 mm broad. Lamellaewhite, dry
Chaetocalathus 55

often yellowish white, narrowto broad (mostly about 1-2 mm broad), subdistantor
distant, intermixedwhen mature, concurrent to the rudiment of the stipe which
is either central or sublateral,and then often merged with the pseudostipe, the lamel-
lae attenuated and narrowlyadnexed or free. Spore print white. Stipe rudimentary
unless lateral and confluent with a pseudostipe, sometimes so poorly developed that
it is practicallyabsent but more frequently knob-like, low, white pubescent, snow-
white when seen under a lens. Context extremely thin to relativelythick, white;
odor none.
Spores 7.5-11.5(13) X 5-8 p, ratherbroadly ellipsoid, without a distinct sup-
rahilarapplanationor depression,with thin, later slightly thickened wall which is
inamyloid or in some specimens, especially when over-agedmore or less strongly
pseudoamyloid, smooth, hyaline in NH40H, often with small internal oil droplets,
cyanophilous in cresyl blue mounts. Hymenium:basidia 17-30 X 6-9 p, (2-)4-spored;
cystidia on edge and sides metuloid, 18-51 X 6-12.5 p, fusoid to ventricose, all simple
and entire or few with cristate apex or appendageslike the cheilocystidia, with pseu-
doamyloid wall 0.4-4.5 j which is in the upper portion or entirely encrusted by
coarse hyaline crystals, with mostly obtuse to subacute tip, but also often acute or
rounded;cheilocystidia similarbut more often with 1-4 apical appendagesor more
rarely effilate into an up to 100 p long, thin rostrate apex, some with a constriction
or vaguely ampullaceous,very rarely with a lateral appendagesor burge, but usually
the main body simple, ratherpolymorphous, the wall mostly inamyloid, more rarely
distinctly pseudoamyloid, thinner-walled,(wall 0.2-0.5 p thick) making the edges
heteromorphous. Hyphae: in the context of the pileus in a lower layer interwoven
and irregularlyshaped but mostly filamentous, in an upper layer more radiallyarranged
and more persistently filamentous, not gelatinized, inamyloid, hyaline, with clamp
connections. Coveringlayer of the pileus consisting of dense hairs 100-400 long
which have thick (3-5.5 p), hyaline walls which are pseudoamyloidand incrustedby
a thin, finely granularincrustation,or without it, with acute to obtuse tips; hypotri-
chium consisting of prolongationsof the hyphae of the trama where the last member
below the hair is slightly swollen and more or less cheilocystidium-likebut, instead
of the appendagebearingthe hair, divided from it by a clamped septum, inamyloid.
On fallen branchesand decaying sticks, also leaves, of mono- and dicotyledonous
plants in and near the forest and in plantations, from the coastal to the cloud forest
region. Fruiting the year around. Known hosts: Richeria grandis;Clibadiumerosum;
Chusquea sp.
MATERIAL STUDIED. USA. Florida: Dade Co., Matheson Hammock 17 Nov 1942, Singer
F 741 (F). CUBA. Wright 78 (paratype of Al. nidulus) (FH); Kotlaba (PR). GUADELOUPE.
Bains Jaunes, Duss 193, 445 (type of Crinipellis calosponrs). MEXICO. Veracruz: Estaci6n Biolo-
gica de las Tuxtlas, 7 km south of Monte Pio, 20 Jun 1969, Singer M 8066 (F); Oaxaca Huautla de
Jimenez, 12 Jul 1957, Singer M 1544 (MICH); Chiapas: above Soluchuchiapas, 1 Aug 1969, Singer
M 8793 (F), Between Ixtacomitan and Soluchuchiapas, 400 m alt; 1 Aug 1969, Singer M 8806 (F).
COLOMBIA. Valle: Mun. Cali, Horqueta, 2 May 1968, Singer B 6927 (F); Dagua Road, N above
km 25, 1900 m alt in the Cordillera Occidental, 28 Apr 1968, Singer B 6463 (F); Camino Cali-
Buenaventura, Cordillera Occidental, west slope at 1950 m alt, GuznaLn4524 (F); Saladito 15 Apr
1968, Singer B 6166 (F); Anchicaya, 26 Apr 1968, Singer B 6398 (F); Boyaca, Los Robles, 22 Jun
1960, Singer B 3489 (F); Cundinamarca, Salto, 24 Jul 1960, Singer B 3506 (F). VENEZUELA.
Dpto. Miranda between Caracas and Cumana, 5 Jul 1972, Dumont-VE 3903 (NY). Parque Nacional
El Avila, El Eden, 19 Jun 1971, Dumont-VE 234 (NY); 30 Jun 1972, Dumont etal VE 3859 (NY);
Dpto. Federal: El Junquito, 10 Jun 1958, 1870 m alt, Dennis 1165 (F). ECUADOR. Tungurahua,
Rio Topo, 30 Apr 1973, Singer B 7224 (F); Rio Verde 28 Apr 1973, Singer B 7175 (F); Pichincha,
Quito: Rev. L. Mlille 14 (Lloyd Herb. BPI); Lagerheim, Jul 1892 (Patouillard Herb., FH); Napo, Lago
Agrio, 8 May 1973, Singer B 7301 (F); B 7293 (F). BOLIVIA. La Paz: Nor-Yungas, Rio Yariza 23
Feb 1956, Singer B 1378 (LIL). ARGENTINA. Misiones: Iguazu 26 Nov 1949, Singer M 98 (LIL).
56 FloraNeotropica

3. Chaetocalathusniduliformis(Murrill)Singer, Lilloa 8: 521. 1942.

Pleurotusniduliformis Saccardo,SyllogeFungorum
(Murrill) 23: 117. 1925.
Pleurotopsis NorthAmerican
Murrill, Flora9: 328. 1915.
TYPE.Brown, Britton & Seaver 1350 from Bermuda(NY).
Pileus white, pilose-floccose, smooth, with straight to undulate, on drying inflexed
margin,orbicular,attached with part of the sterile surface,inverted-cupuliform,1-5 mm
broad. Lamellaewhite, subclose to subdistant or distant, medium broad, concurrent to
stipe rudiment, adnexed to or free from it. Stipe rudimentaryand not functional, pure
white, central or eccentrically located, a very small low button or cone, velutinous, often
indistinct. Context white, very thin.
Spores 6.5-9.5 X 4.8-7 u, ellipsoid to broadly ellipsoid, smooth, hyaline, pseudoa-
myloid. Hymenium: basidia 25-32 X 8.5-8.8u, 4-spored;cheilocystidia 29-43 X 4-7(11.3)
u, versiform,either fusiform, subfusoid or ventricose to ampullaceousor clavate with
one to three apical appendageswhich have obtuse apices, most frequently bifurcate at
apex, appendages2-25 X 2-4 ,, hyaline with thick walls and then mostly pseudoamyloid,
or with thin wall and then mostly inamyloid, mostly thinly incrusted, densely crowded
at the edges but in age graduallybecoming more numerous(but always sparse)outside
the broad edge region as pleurocystidia,but these not different from cheilocystidia.
Hyphae hyaline, with clamp connections, inamyloid. Hymenophoraltrama regular,in
places appearingsubirregularbecause of swelling and inflated hyphal ends (31-70 X 14-
19 u). Epicutis: hairs long but ordinarilynot reaching300 u, 2.8 , broad rarely partially
inflated and there reaching9.5 u in diam., with thick (0.7-2 ,) wall which is distinctly
pseudoamyloid,near apex often with scattered secondary septa, the tip rounded or
attenuated to an obtusate tip, more rarely acute or subacute but not needle-sharpacute.
On stipe rudiment pseudoamyloid smallerhairs.
On dead twigs of Juniperus barbadensis,accordingto African collectors (reported
by Pegler) also on Morindacitrifolia and Uvariachazae, solitary or gregarious. Bermudas,
also in WestAfrica (Sierra Leone, Ghana, Congo).
MATERIALSTUDIED. BERMUDA. Brown, Britton & Sealer 1350 (type, NY), 1352.
Zaire. Kwango:Kinanga,H. Vandervst,11 Apr 1910 (BR).
This species is somehow intermediatebetween C. collumellifer and C. liliputianuts
but closer to C. liliputianus from which it differs mainly by the cystidial characters.
ILLUSTRATION. Singer (1942), fig 5.

4. Chaetocalathusasperifolius(Patouillard)Singer, Lilloa 8: 530. 1942.

CrinipellisasperifoliaPatouillard,Jour. de Bot. 3: 336. 1889.
Marasmiusasperifolius(Patouillard)Saccardo,Sylloge Fungorum9: 70. 1891.
Pleurotopsisasperifolia(Patouillard)Murrill,North AmericanFlora 9: 239. 1915.
TYPE.Duss 35, from Martinique.
Pileus light grayish leather brown, very slightly tomentose but distinctly pilose
under a lens, slightly striate from the point of attachment (on sterile surface) towards
the margin, later upper part of pileus becoming reflexed, with reniform outline and
often with an incision on the concave side which reaches the stipe rudiment, 4-10 mm
broad. Lamellaeconcolorous or paler, medium broad, attenuate, medium distant to
subdistant, intermixed. Stipe rudimentary,very small, hemispheric,hirsute, concolor-
ous or paler to almost white, disappearingin old specimens. Context very thin.
Spores 9.5-10.5 X 5-5.3 uellipsoid to subfusoid, inamyloid, hyaline in NH40H,
smooth. Hymenium: cystidia 38-60 X 8.5-21 p,, projectingbeyond the basidia, fusoid
more rarely clavate, sometimes ampullaceous,rarely forked, acute or obtuse, hyaline
in NH40H, with thick, pseudoamyloid wall. Epicutis of pileus consisting of long
Chaetocalathus 57

hairs, cylindric, rarely of some other shape, thick-walledincrusted, strongly pseudoamy-

loid with rounded tips. 4-6.8,broad.
On bark of living Murrayaexotica, after heavy rains, gregarious.

5. ChaetocalathuscarnelioruberSinger, Lilloa 8: 526. 1942.

TYPE. Linder, from Surinam (FH).
Pileus "carnelianRed (R.) when fresh becoming a color between "PerillaPurple"
and "MineralRed" (R) when dried, with paler margin,subtomentose, strongly pilose
under a lens, appearingalmost cup-shaped,attached to the sterile surface and mostly
reflexed, reniform in circumferencewith a deep incision on the concave side, 2-7 mm
broad when dried. Lamellaecream buff when fresh, brownish when dried, very narrow
distant or moderately distant, attenuate towards the incision of the pileus. Stipe rudi-
mentary, almost absent in the young carpophoresbut under a lens visible as a minute,
verruciform,lilac-brown,hirsute, button-like, function-lessstructure. Context very thin.
Spores 11 X 7 p, ellipsoid-ovoid,with granularcontents, thin-walled,Hymenium:
basidia 34.8 X 8.8 ; cystidia 38-100 X 13-32p fusoid, hyaline or yellowish in NH40H,
pseudoamyloidor nearly amyloid, with very thick walls, which are polystratous and
5-8.5 1 thick in the largest cells, incrusted by finely granularmaterialin the upper por-
tion, all entire. Hyphae with clamp connections. Hairsof the surface of the pileus
3.5-5 p in diameter, with thick pseudoamyloid or almost amyloid wall, with obtuse
rounded tips.
On dead branches.
MATERIALSTUDIED. SURINAM,Bartica,9 Dec 1923, D. H. Linder (FH), type.
Singer (1942), fig 6 f i.

6. ChaetocalathusaurantiacusSinger, sp nov
TYPE.Singer B 2201, from Bolivia.
Pileo obtuse aurantiaco;ceterum C. carneliorubersimilis. Typus in F conser-
Pileus dull orange, finely pilose, finely long-pilosewhen seen under the dissecting
microscope dry, with hyaline or white hairs, sulcate, convex, attached to the substratum
by part of the sterile surface, up to 6.8 mm broad. Lamellaeorange-pallid,distant,
narrow,concurrent at the rear incision, not intervenose. Stipe none, but an indistinct
stipe rudiment in the rear sinus present as a tomentose patch.
Spores (8-)11-13.3 X (5.5-)6-7.5 u, ellipsoid to ovoid without suprahilardepress-
ion or applanation,smooth, hyaline, inamyloid. Basidia25-29 X 11-13 u; cystidia 32-
89 X 7-18 , of the Geopetaluhm-type, with 4-6 p thick-wall, mostly fusoid and acute to
subacute, rarely once forked, with a thick cystalline incrustation,strongly pseudoamy-
loid (exceptionally some inamyloid);cheilocystidia 30-32 X 4-6 u, narrowlyventricose
to subfusoid, often constricted in the middle, often appendiculateor with bi- to tri-
lobed apex, wall thin to slightly thickened (to 1 ,), inamyloid. Hyphae hyaline, not
gelatinized, inamyloid, with clamp connections. Corticallayer: hairs of the surface of
the pileus dense, remarkablythin (3-4 p broad) and long, wall 1-1.2 p thick, pseudoamy-
loid, tips obtuse or subacute.
On dead dicotyledonous branchesin hylaea.
MATERIALSTUDIED. BOLIVIA. Pando, Manuripi,Conquista,25 Mar1956, SingerB
2201 (F), types.
58 Flora Neotropica

3. LachnellaFries, Corpus FlorarumSueciae 1. Floram scanicam sceipsit, p 343.

Charactersas given in the keys (p ), see also Singer (1973), p
TYPESPECIES.LachnellaalboviolascensAlbertini et Schweinitz ex Fries

Key to the Subgeneraand Species of Lachnella

1. Cystidiametuloid.Subgenus MetuloidiferaSinger. 1. L. cecropiae.
1. Cystidianone.Subgenus Lachnella.
2. Sporeselongatedandalmostsickle-shaped, 15.5-17X 5-6.5p;carpophores
up to 0.25 mmhighand0.4 mmbroad. 2. L. subfalcispora.
2. Sporeslesselongatedor fruitingbodieslarger.
3. TropicalspeciesfromnorthernSouthAmerica; spores12-17X 5-8,. 3. L. caracassana.
3. Subtropicalspecies;spores 10-13.5 X 5-8.3p. 4. L. villosa.

Lachnellasubgen MetuloidiferaSinger,Darwiniana14: 13. 1966.

Charactersas given in the key above.
Fig 27.
1. Lachnellacecropiae Singer, Darwiniana14: 13. 1966.
TYPE.Singer B 24901, from Bolivia (LIL).
Pileus a sessile cup which has a free margin,white. silky-tomentose on the out-
side, with often reflexed margin, 1-5 mm broad. Lamellaenone, hymenial surface
smooth, white, not deeply excavated but convave. Stipe or pseudostipe none. Context
very thin, white, waxy, slightly harderwhen dried, inodorous.
Spores 11.8-17.2 X 8-11.8 u, mostly 14-16.5 X 8-8.5 p, ellipsoid, smooth, hyaline,
inamyloid. Hymenium: basidia easily collapsing;cystidia 85-132 X (9-)15-25 ,, metuloid
with stratified thick pseudoamyloidwall, entire, fusoid or fusoid-ventricose,subacute
or acute, rarely obtuse but always attenuate towards the tip, with colorless crystals
thickly incrusted. Hyphae hyaline, inamyloid, with clamp connections. Coveringlay-
ers: epicutis of the pileus consisting of very long hairs covering densely the sterile
surface;these hairs 2.7-7 ,broad, smooth but mostly slightly incrusted, with thick
(1-3-, rarely to 4 u) psuedoamyloid wall, more frequently acute than obtuse, the apex
sometimes separatedby a sedondary(clampless) septum and thin-walled,but generally
thick-walledall over and aseptate.
On leaves of Cecropiain the Amazonas region.
MATERIAL STUDIED.BOLIVIA.Beni:VacaDiez,Ivon3 Apr1956,SingerB 24901
(LIL, BAFC), type.

Charactersgiven in the key above.
TYPESPECIES.LachnellaalboviolascensAlbertini & Schweinitz ex Fries.

2. Lachnella subfalcisporaReid, Kew Bull. 15: 267. 1961.

TYPE.Dennis 1472, from Venezuela (K).
Pileus a small cup, white, becoming subglobose or turbinatewhen dry, densely
crowded, the sterile surface villous, 0.1-0.4 mm broad. Lamellaenone, hymenial sur-
face smooth, white. Stipe or pseudostipe none, but cup often with a slight, tubercle-
like swelling.
Spores 15.5-17 X 5-6.5-(7) u, subfalcate,hyaline, smooth, inamyloid. Hymenium:
basidiaup to 12 broad, 4-spored; basidioles fusoid; cystidia none. Hyphae densely
compacted, hyaline, narrow. Cortical layer: hairs at first arising from the base of the
Lachnella 59

carpophoremore or less appressedto the sterile surface and mostly overtoppingthe

margin,joined to the hyphae of the trama by a clamp connection, very long, (up to
195 ,) and aseptate or with a few secondary septa, 4-5 pbroad, with thickened hyaline
wall which thins out towardsthe apex which may be slightly swollen, pseudoamyloid,
incrusted by a granularsubstance which dissolves in KOH(in which medium the hairs
become locally swollen and distorted).
On twigs, associated with Cyphellopsissubglobispora.
MATERIAL STUDIED.VENEZUELA. Dto Federal,CaracasBotanicGarden,30 Jun 1958,
Dennis1472 (K),type (seen,not sectioned;description
ILLUSTRATION. Reid (1961), fig 2, 2; 3, 6.

3. LachnellacaracassanaReid, Kew Bull. 15: 267. 1961.

TYPE.Dennis 1876, from Venezuela (K).
Pileus in form of a cup, white, villous on sterile surface, subglobosewhen dry,
0.3-0.4 mm broad. Lamellaenone: hymenial surface smooth, white. Stipe none but
carpophoresnarrowedinto a very short, stipe-like base which is usually buried beneath
the superficiallayer of bark.
Spores 12-17 X 5-8 1, elongated, somewhatasymmetricallyobpiriform,hyaline,
smooth, inamyloid. Hymenium: consisting of basidia, no cystidia. Hyphae as in the
precedingspecies but in the basal region of the carpophoremore agglutinated. Cor-
tical layer: hairs sheathing the fructification much like those of the precedingspecies.
On twigs. Gregariousbut carpophoresscattered.
MATERIALSTUDIED. Venezuela. Dto Federal,Caracas BotanicGarden14 Jul 1958,
Deiinis 1876 (K), type (seen, not sectioned;descriptionadapted from Reid).
Reid 1961 fig 3, 4

4. Lachnellavillosa (Persoon ex Schweinitz) Gillet, Champignonsde France, Dis-

comycetes, p 80. 1881.
Peziza villosa Persoonex Schweinitz,Schriften,NaturforschendeGesellschaftLeipzig 1: 120.
1822 (non Chevelier1826).
Cyphlellavillosa (Persoonex Schweinitz)Crouan,Flore Finist4re,p 61. 1867.
villosa(Persoonex Schweinitz)
Trichopeziza des Nassauer
Fuckel,Jahresbericht Vereins23-24:
296. 1870.
Chaetocv'pa villosa (Persoonex Schweinitz) O. Kuntze, RevisioGenerumPlantarum2: 848.
C'plhelladochimiosporaBerkeley& Broome,Ann. Mag.Nat. Hist. IV, 11: 343. 1873.
Cliaetocvphadochiniospora(Berkeley& Broome)O. Kuntze, RevisioGenerumPlantarum2:
847. 1891.
TYPE. Persoonl,from Germany(L).
Pileus white, in form of a cup which has inrolled marginwhen dry, hairy-villous
on the sterile surface, nearly 0.5 mm broad. Lamellaenone, hymenial surface smooth
and white. Stipe none, but often with a very short pseudostipe attached to a central
point of the sterile surface. Context very thin, white, inodorous.
Spores about 12 X 5 p(according to Cooke), 12-13.5 X 7-8.2u, fusoid to obcon-
ical in other collections, hyaline, smooth, inamyloid. Hymenium:basidia 25-40 X 7-10
,, 4-spored;basidioles fusoid; cystidia none. Hyphae hyaline, filamentous, with clamp
connections, inamyloid. Cortical layers: hairs long and aseptate, obtuse, 5-8 ,broad,
with moderately thick to thick, pseudoamyloidwall, incrustedby hyaline small crystals.
On dead wood and leaves of Dicotyledones. Known host in South America:
Abius jorullensisvar spachii. Probably confined to the montane and subtropicalzone.
In the neotropics said to occur in Ecuador;also in Europe and North America,Australia.
60 Flora Neotropica

MATERIAL STUDIED.ARGENTINA. Tucuman,Tafi,Tafidel Valle1 May1949,Singer

T 516 (LIL). GERMANY.Konigstein,Krieger,FungiSaxonici1457 (FH).
The description of this species as given above is derived from the Argentine
material determined as Cyphella villosa by W. B. Cooke.
This is in fairly good agreementwith the data obtained by Donk (1959) from
the type preservedin the Persoon Herbarium.
ILLUSTRATION. Patouillard(1900), fig 38, 8.

4. AmyloflagellulaSinger, Darwiniana14: 14. 1966.

Characters,see key p 4. and Singer (1966), p 14.
Type species: Cyphellapulchra Berkeley & Broome
Only one species known in the neotropics.

1. Amyloflagellulapseudoarachnoidea(Dennis) Singer, Darwiniana14: 14. 1966.

Fig 28.
Marasnius Dennis,Trans.Brit.Mycol.Soc. 34: 425. 1951.
Crinipellispseudoarachnoidea(Dennis) Singer,Sydowia 15: 60. 1961.
TYPE.Dennis 137, from Trinidad(K).
Pileus white or pale buff, deeply sulcate, convex, 3-4 mm broad. Lamellaepale
buff, distant (four to six) narrow, adnate. Spore print white. Stipe white, smooth,
glabrous, often curved, not longer but often shorter than the diameter of the pileus,
rising from a white rhizomorph. Context white, very thin.
Spores 18-19 4,u, lanceolate, smooth, hyaline, with mostly applanate inner
side when seen in profile, inamyloid. Hyphae of the pileus hyaline, inamyloid, with
clamp connections. Cortical layers: epicutis of the pileus and surface layers of the
stipe as well as the rhizomorphsconsisting of relatively broad, erect or ascendant hy-
aline and inamyloid terminalcells of hyphae, many of these forked or twice forked,
their apices rounded-obtuseand with a disc-like strictly apical pseudoamyloidzone
from which 4-10 flagellum-likethin, apical appendagesarise, these strongly pseudoamy-
loid, 1.7-4.8 ulong.
On forest litter, solitary.
MATERIALSTUDIED. TRINIDAD. ArenaForest,5 Oct 1949, Dennis 137 (K),
The macroscopicaldescriptionis given accordingto Dennis (1951 a) who illustrated
this species (pl 19, fig 20); also Dennis (1970), pl. 2, fig 20.

5. FlagelloscyphaDonk in Singer,Lilloa 22: 312. 1951.

Characters:See key characters.
TYPESPECIES. CyphlellaminuttissinlaBurt.
Only one species thus far observedin the neotropics.

1. Flagelloscyphapolylepidis Reid, Kew Bull. 15: 269. 1961.

TYPE. Dennis 1809, from Venezuela(K).
Pileus in form of cups, white, villose, becomingglobularwhen dry, shallowly dis-
coid in profile, up to 0.7 mm broad. Lamellaenone; smooth hymenial surfacein the con-
cave portion of the cup white. Stipe or pseudostipenone; only a rudimentarystipe-like
base present.
Spores 8-10 X 5-6.5 ,, broadly ellipsoid to pin-shaped,smooth, inamyloid. Hymen-
ium: basidia23.3-41.6 X 5.5-6.5 I, 4-spored;cystidia none. Hyphae of the monomitic
tramathin-walled,2.5 pin diameter,with clamp connections. Corticallayer: epicutis
Flagelloscypha 61

consistingof hairs densely coveringthe sterile surfaceof the cups, up to 150 X 2.5-3 Iu,
densely covered by detersilerod-shapedcrystals, divergent,with slightly thickened wall,
thinningout towardsthe stronglytaperingwhip-likeapex and also towardsthe often
somewhatnarrowedbase, long, narrow,hyaline, continuous (non-septate).
On gall of Poll'lepis sericea.
MATERIALSTUDIED. VENEZUELA.Merida:Sierrade Santo Domingo,LagunaNegra,
3440 m alt 27 Jul 1958, Detnnis1809 (K), type (seen not sectioned;descriptionaboveaccordingto
type descriptionby Reid).
ILLUSTRATION.Reid (1961), fig 3, 1.

Tribe Marasmieae subtribe Marasmiinae Singer, Sydowia 15: 60. 1961.

TYPEGENUS. Marasmius Fries.
Characters: not combining the characters indicated under subtribus Oudemansielli-
nae and subtribus Crinipellinae i e often with epicuticular broom cells, without epicuticular
flagella-like pseudoamyloid appendages, without long, thick-walled hairs, the stipe, if
present, often insititious or else mostly not forming a distinct pseudorrhiza; hyphae
pseudoamyloid or inamyloid.

Key to the Genera of Subtribe Marasmiinae

1. Hymenophorelamellate,the lamellaeintervenoseor not.
2. Epicutisnot hymeniform{see Marasmnius, p 62).
2. Epicutishymeniform.
3. Epicutisconsistingof or containingmanybroomcells (seeMarasmiusp 62).
3. Epicutisconsistingof smooth, mostly entire cells.
4. Capitateand/or metuloid and/or oleo-cystidiapresent in the epicutis and/or
the hymenium;stipe central, eccentric, lateral or absent; small carpophores
(pileus < 15 mm, <5 mm if centrallystipitate),some instititious. Gloiocephala,p 284.
4. Capitate, metuloid and oleo-cystidiaabsent unlessthepileus is centrallystipi-
tate and risingfrom basalmyceliumand then pileusreachinglargersize.
Marasmius,p 62.
1. Hymenophorewith lamellae which are reduced to a few veins or so strongly anas-
tomosingthat they appearporoidor merulioid,or else hymenialsurfacesmooth.
5. Epicutisreduced since the pileus is head-likeand hollow or else the portion of the
pileus (head) aroundand near the apex of the pseudostipesterile;tramaalwaysvery
thin in the pileus (head). [Other types of pileusconfigurationof the "physalacrioid"
type not representedin the neotropics.] Physalacria,p 304.
5. Epicutison the sterile upper side of the solid pileus well developedand not reduced
to a sterilezone; tramathin to ratherthick.
6. Epicuticularcells in the form of broom cells (Rotalis-type);carpophoresminute,
pileusless than 5 mmbroad.
7. Hymenial cystidia none; basal mycelium present; hyphae at least partly
pseudoamyloid. Epicnaphus,p 312.
7. Hymenialcystidiapresent;basalmyceliumabsent;hyphaeall inamyloid.
8. Cystidiaabsent;stipe risingfrom a rhizomorph. Manuripia,p 312.
8. Cystidiapresent;stipe risingfrom the substratum(see "3" above).
6. Elementsof the epicutisnot containingbroom cells.
9. Carpophoreswith a central,eccentricor lateralstipe, more rarelysessile.Epi-
cutis of the pileushymeniform.
10. Carpophoresrather large with long central stipe and smooth hymenial
surface, with basal mycelium;pigmentsof all partsof carpophorebright
(red or yellow). Hymenogloeap 313.
10. Carpophoreswith pileus to 15 mm broad, more often less broad, and
never brightred or yellow if centrallystipitate;stipe often institious(see
"4" above).
9. Carpophoreswith a centralor eccentric,rarelylateralpseudostipe(attachedto
the side of the pileus opposite to the hymenialsurface).Epicutisof the pileus
hymeniformor not.
11. Capitatedermato-or oleo-cystidiapresent(see Physalacria,p 304).
11. Capitatedermato-and oleo-cystidiaabsent. Rimbachia,p 314.
62 Flora Neotropica

6. MarasmiusFries, GeneraHymenomycetum,p 9. 1836.

HeliomycesLUveille,Ann. Sci Nat. III. 2:177. 1884
Androsaceus(Persoon)Patouillard,HymenomycetesEuropaei,p 105, 1887.
Chanaeceras0. Kuntze,RevisioGenerumPlantarum3, 2: 454. 1898.
MycenitisEarle,BulLNew York Bot. Gard.5: 414. 1909.
Collybiopsis(Schroterin Cohn) Earle,Bull New York Bot. Gard.5: 415. 1909. Basionym:
MarasmiussubgenusCollybiopsisSchroterin Cohn, Kryptogamenflora von Schlesien3a: 559.
TephrophanaEarle,BulLNew YorkBot. Gard.5: 27. 1909.
ScorteusEarle,Bull. New York Bot. Gard.5: 415. 1909.
PolvinarasmiusMurrill,North AmericanFlora9: 286. 1915.
? Vanromburghia Holtermann,MycologischeUntersuchungenin den Tropen,p 104. 1898.
TYPESPECIESMarasmius rotula (Scopoli ex Fries) Fries
Habit collybioid (to almost mycenoid) or pleurotoid, mostly somewhat toughish
and revivingafter dryingout when remoistened;hymenophorelamellate, rarelysmooth
or venose, lamellaesometimes intervenoseor even anastomosingand appearingporoid:
epicutis either hymeniformor not hymeniform:if not hymeniformconsistingof broom
cells or at any rate strongly diverticulate-nodulosehyphal elements(and then either the
whole tramaor only that of the stipe more or less distinctly pseudoamyloidor at least
the stipe seta-likeand shining,blackish,insititious, or absent to smalland eccentric to
lateral);tramaof the pileus monomitic, pseudoamyloidor inamyloid, no gelatinous
layers present;clamp connections present, or, more rarely,absent; if absent,epicutis
with broom cells; black rhizomorphsoften present, sometimes stipe risingfrom these:
telepods often formed; if the basal mycelium present, the epicutis always hymeniform;
some species formingendotrophicmycorrhiza,none ectotrophs or lichens. Most fre-
quently on wood or leaves, dead or living, more rarelyon other kinds of litter, among
mosses or grasseson earth, parasiticor saprophytic. Developmentof the carpophores
hemiangiocarpousat least in some species(paravelangiocarpous accordingto Reijnders;
1963; pileostipitocarpousin the type species accordingto Reijnders). Almost cosmo-
politan, but much more numerousin carpophoreproductionand numberof species in
the tropical regionsthan in the temperateand frigidzones.

Key to the Sections and Subsectionsof Marasmius

1. Epicutishymeniform,rarelylatersubhymeniform.
2. Stipe insititious and central and lamellae with a collarium;epicutis hymeniform,
consistingof broom cells. sect Marasmius.p 91.
3. Epicuticularbroom cells of the Rotula type. subsectPararotulae,p 92.
3. Epicuticularbroom cells of anothertype.
4. Epicuticularbroom cells of the Siccus-type. subsectPenicillati.p 121.
4. Epicuticularbroom cells, even those of the marginalregionof the pileus,with
erect finger-like appendages interspersedwith some large setulae (setoid
bodies)arisingdirectlyfrom the subjacenthyphaeand undivided.
subsectPenicillati, p 121.
2. Stipe insititious and central, or not insititious or not central;lamellaewithout a
collarium,free or directly attachedto the apex of the stipe;epicutisof pileuswith or
without broom cells.
5. Stipe insititiousand central,if not central,epicutisof smooth cells exclusively.
6. Upperpart of stipe, lamellaeand largermarginalportionof the pileuswhite or
whitish-cream;lamellaesometimesabsent, or replacedby veinsor a merulioid
hymenophore;epicuticular cells and cheilocystidiasmooth (if diverticulate,
not with pigmentedsetulae). SectionEpiphylli,p 87.
7. Apex of stipe consistingof inamyloidhyphae. subsectEpihyllini,p 87.
7. Apex of stipe pseudoamyloid. subsectEufoliatini,p 90.
6. Pileus and stipe more stronglypigmented;lamellaewell formed;epicuticular
cells and/or cheilocystidia,at least some of them, appendiculateor beset with
Marasmius 63

8. Elementsof the epicutissmooth;cheilocystidiacorncob-like(not smooth).

sect Scotophysini,p 86.
8. Elements of the epicutis at least in part with appendagesor else with
setulaeand then of the Rotalis-type,or manybroom cells seriate.
9. Smallspecies;most epicuticularcells are broom cells of the Rotalis-type
or they areaggregated-seriate. sect Hygrometrici,p 79.
9. Mediumsized species; many epicuticularelements smooth, the others
with apicalor subapicalfinger-likeappendages(not of the Rotalis-type).
sect Leveilleani,p 85.
5. Stipe eithernot insititiousor not central(or neither).
10. Stipe centralor subcentral;basal myceliumgenerallystronglydevelopedand
often strigose;epicuticularelements either smooth or in form of broom cells
of the Siccus-type.
11. Elementsof the epicutisall smooth without setulae.
12. Hyphaeof the tramapseudoamyloid;sporescyanophilic. sect Globulares,p 269.
12. Hyphaeof the tramainamyloid;sporesacyanophilic. sect Alliacei,p 263.
11. Elementsof the epicutis partlysmooth and partlyin form of broomcells
or else all cells of the epicutisin form of broom cells of te Siccus-type.
sect Sicci, p 159.
13. Hyphaeof the tramainamyloid. subsectInaequales,p 159.
13. Hyphaeof the tramapseudoamyloid. subsectSiccini,p 159.
10. Stipe strongly eccentric to lateral or even rudimentaryto absent in mature
specimens,only very young carpophoresmay be centrallystipitate;epicuticu-
lar elements in the form of broom cells (which may be of the Rotalis-or
Siccus-type,in the lattercase setulaeoften forkedor diverticulate),but occa-
sionally many or even the majoritysmooth; base of stipe or point of attach-
ment of the pileuseither insititiousor carpophorerisingfroma tomentoseor
fibrillose,usuallywhite, smallpatchof mycelium.
14. Epicutis of the pileus merely subhymeniform;setulae of epicuticular
broom cells hyaline or light golden yellow, not brown, many terminal
cells of epicutis without setulae;pleurocystidiain form of fusiformentire
cystidia with ratherthin walls but resinouscolored incrustationpresent;
stipe presentand not insititious;most hyphaeof the pileus-tramapseudo-
amyloid (see "15" below).
14. Epicutis of pileus hymeniform or subhymeniform,in the latter case
cystidia either not incrustedor incrustedcystidia only on and very near
the edges, or cystidia absent (exceptingfor broom cells which may occur
at the edge of the lamellaeas cheilocystidia);stipe present and either
insititious or not, or pileus sessile: hyphae of the pileus trama pseudo-
amyloidor inamyloid. sect Neosessilesp 253.
1. Epicutis of the pileus definitely not hymeniformor subhymeniformbut in form of
interlacedhyphaeor terminalbodieswhichbearsetulae.
15. Stipe central, insititious or inserted directly in a dark rhizomorph;conspicuous
incrustedpleurocystidianone. sect Androsacei,p 63.
15. Stipe eccentric or sublateral,sometimes (rarely)none, rising from a small white
mycelial patch appressedto the substratum(wood); conspicuousincrustedpleuro-
cystidiaalwayspresent. sect Fusicystides,p 251.

Schematicdrawingsshowingthe principaldifferencesbetween the sections of the

genus Alarasmiiusmay be compared in Singer (1965), plates I-IV (except for section
Leveillea ni).

Marasmiussect AndrosaceiKilhner,Le Botaniste 25: 60. 1933 (as "Androsaceae").

TYPESPECIES.Marasmius androsaceus (L. ex Fr.) Fr.
Pileus usually small, pigmented or not, with an epicutis of the Rameales-type or
irregularly arranged broom cells; spores generally medium sized, ellipsoid to oblong;
lamellae not collariate or more or less imperfectly collariate; stipe central, insititious,
dark, pilose, or glabrous; hyphae with or without clamp connections, moderately strongly
pseudoamyloid or inamyloid. On dead vegetable matter.
This is the only section which might possibly be confused with Marasmiellus. Wher-
64 Flora Neotropica

ever the genericposition of the specimenunder investigationis doubtful, a monographof

the neotropicalMarasmiellishould also be consulted (Singer 1973).

Key to the Species of Section Androsacei

1. Clampconnectionspresent.
2. Stipe glabrousand smooth (evenundera good lens).
3. Combiningthe habitat on wood sticks of Alnus or other woody materialin
montane alneta with white, distant (through-lamellae7-11) lamellae; hypo-
dermium with inconspicuous or no incrustation;stipe less than 15 mm long;
spores5.5-9.5 X 2.5-3.5, most frequently7-8.8 X 2.5-3.2, 1. M. valae.
3. Not combiningthe abovecharacters;on leaves(if on woody sticks or roots see "5"
4. Dried pileus papillateor stronglyumbilicate,2-3.5 mm broad;stipe relatively
long: 25-75 mm; through-lamellaerelatively few (7-10); species growing on
Polylepisleavesin the fog forestand sub-paramo. 2. AI.pol)ylepidis.
4. Pileuseitherlarger,or not or weaklydepressedwhen dried;through-lamellae in
maturespecimensat least 11. 3. M. cyrillidis.
2. Stipe, at least when young and fresh, finely pubescent or pruinate,pilose, or finely
setulose, sometimesmacroscopicallyappearingglabrous,but underbinoculardissect-
ing microscopedistinctlycoveredby microscopicalhairs,setae, or dermatocystidia(if
stipe risingfrom rhizomorph,see "15" below).
5. Spores 10-12 X 2.7-3.7,. 4. M. bactrosporus.
5. Sporesless elongatedor broader,not strictlyrod-shaped.
6. Lamellaegray when fresh, nine; stipe tendingto be eccentric;on dead barkof
leguminoustrees. 5. M. legziminosarulm.
6. Lamellaenot gray when fresh, (7-)10 or more through-lamellae; on deadwood
or deadleaves.
7. Coveringof the stipe consistingof short,acute, subulate-spinosediverticula
which are numerous but not crowded, erect to ascendant, subhyaline,
3.3-12 X 1-4u. WestIndies. 6. Alearlei.
7. Coveringof the stipe different.
8. Growingon leaves.
9. Growing on (wood and) fallen leaves of Liquidambarin Mexico;
pileus vinaceous brown only in youth, soon with very pale margin
which is, when mature,radiallyrugoseor sulcateup to 6/7th of the
radius sulcate; hypodermiumonly in the disc-regionspadiceousin-
crusted,otherwiseincrustationon the hyphaeratherinconspicuous,
pigment where present mainly intraparietal;cystidioid elements of
the coveringof the stipe < 70. 7..M. liquidambari.
9. Growingon fallen leavesof variousdicotyledonoushosts;pileusand
hypodermiumas above or less sulcate and with deepercoloredand/
or more conspicuouslyincrustingmaterial;cystidioid hairs of the
coveringof the stipe as aboveor different.
10. Stipe densely beset with villous hairs especially in the upper
portion, these hairslong with thin (0.3p) to slightly thickened
hyaline to brownishhyaline wall, obtuse above,and sometimes
grossly diverticulated,in our material8-60 X 4.5-9.5u; stipe
color deep (chestnut)brown;spores7-9 X 3-4p;on fallenleaves
of fagacioustrees. 8. M. splachnoides.
10. Stipe finely setulose or pruinate,elementsof the stipe covering
different from those describedabove, or at least different ele-
ments present; stipe cinnamon, chestnut or black; spores
smalleror narrower;on fallen leaves of oak or more often on
other substratum.
11. Lamellae mediumclose to subdistant,more rarelyclose or
distant; stipe when seen under a dissecting microscopein
young and fresh conditiondistinctlycoveredby long, acute,
hyaline bristles which are often incrusted,but are not per-
sistent, often breakingoff but sometimeslater regenerated,
in the basal third of the stipe also long Crinipellis-likehairs
present, these thick-walled,hyaline to aple melleous, the
inner wall layer often pseudoamyloid,taperinggraduallyto
Marasmius 65

the apex, up to 150 or 170u long; stipe alwayswith setae or

pubescentin the lowerportion. 9. M. hakgalensis.
11. Lamellaeclose to mediumclose; coveringof the stipe often
consisting only of small elements, if there are large hairs,
these are characteristicallyeffinate; stipe pilose only at the
apex when young, or pubescent only when seen under a
12. Long, thinly effilate hairs present at the apex of the
stipe;on Quercusleavesin centralMexico. 10. M. cryptotrichus.
12. Hairsof stipe short,not effilate.
13. Lamellae fresh white; carpophore accompaniedby
thin, dark rhizomorphs;tramaof pileus and hyaline
hyphae of stipe inamyloidor very slightlyand slowly
pseudoamyloid;rain forest species of the lower alti-
tudes. 11. M. chiapasensis.
13. Lamellae fresh pale leather brown to buff-whitish;
carpophoresnot accompaniedby darkrhizomorphs;
some hyphae of the pileus trama weakly pseudo-
amyloid, many pseudoamyloidin the apex of the
stipe; subtropicaland tropical-montanespecies (see
"4" above).
8. Growingon wood.
14. On Lituqidambar in Mexico(see "9" above).
14. On roots and woody rootlets of other trees in South America;
lamellaeratherbroad;stipe not accompaniedby darkrhizomorphs.
12. M. radicellicola.
1. Clampconnectionsabsentin the hyphaeof the carpophores.
15. Stipe only rarelyrisingfrom a black rhizomorph,usuallylonger than the diameter
of the pileus.
16. Odorof Allitium sativuIm.
17. Sporesellipsoidto piriform,6.2-9 X 2.5-4.8u; hypodermialelementsroughish
from incrustingpigmentbut pigmentsubhyalineto pale melleousand there-
fore incrustationnot conspicuous. 13. M. dysodes.
17. Spores mostly oblong, (6.5-)7-8.5 X 2-4,; hypodermiumstrongly sepia-
incrusted. 14. M. atroincrustatus.
16. Odornone.
18. Basidia4-spored;on dicotyledonousleaves.
19. Stipe pigmented.
20. Stipe indistinctlypilose, beset with thick-walledhairsof the Crinipel-
lis-type.Inodorousformsof M. atroincrustatus.
20. Stipe with dense fibrillose-furfuraceous white coveringunderneaththe
spadiceous-sepiaand somewhat shiningsurfaceof the stipe rind, this
coveringconsistingof a tangleof mostly clavatedermatocystidiawith
walls 0.2-0.5 thick, few thick-walledand then wall sepia, some
hyphous,at timesappendiculate.Florida. 15. M. atlanticus.
19. Stipe white. 16. M. eorotula.
18. Basidia2-spored;on monocotyledonousleaves. 17. M. defibulatus.
15. Stipe alwaysor almostalwaysrisingfrom conspicuousrhizomorphs,its lengthabout
equal with the diameterof the pileus. 18. M. tomentellus.

1. Marasmiusyalae Singer, sp nov Fig 29.

TYPE. Singer T 5131, from Argentina.
Pileo albo vel griseo, 3-4 mm lato; lamellis distantibus; stipite glabro, rhizomorphis
destituto. Sporis 5.5-8.5 X 2.5-3.5X. Ad quisquilias alneas in Argentina. Typus in F
conservatus est.
Pileus white or gray, becoming pale cafe au lait or fuscous to brown on drying,
glabrous, sulcate, convex, papillate or umbonate to subpapillate to subumbonate, some-
times with a depressed circular sulcus around the central umbo, 3-4 mm broad. Lam-
ellae white, sometimes becoming pale leather color on drying, with seven to 11 through-
66 Flora Neotropica

lamellae, regularlyinserted with lamellulae or with only a few shorter ones, with entire
margin,rather broad, collariate or broadly adnate or adnexed directly to the apex of
the stipe. Stipe gray with white apex when fresh, tending to become chestnut-fuligin-
ous or fuscous when dried, glabrous,shiny-setose, equal, insititious, about 12-13 mm
long, 0.1-0.2 mm broad. No rhizomorphs. Context thin, almost pallid, inodorous.
Spores (5.5-)7-8.8(-9.5) X 2.5-3.2(3.5) ,, oblong-subfusiformto subcylindrical,
sometimes up to three times as long as broad, smooth, hyaline, inamyloid. Hymenium:
basidia 16-24 X 5-7.5 p, 4-spored, often with a few 2-spored ones intermixed;cystidia
none; cheilocystidia numerousbut discontinuous, 16-24 p long ventricose or hyphous
but at times forked and the majority diverticulate,with irregularlydistributeddiver-
ticula 1-3.5 1 long, or with apical appendagesup to 5 1 or more in length, the whole
up to 8.5 p,in width; where cheilocystidia discontinuous, the edge either of modified
cheilocystidia, hot entire, or by basidioles and basidia, or both. Hyphae hyaline, more
rarely pale melleous in part inamyloid, or with a few vaguely pseudoamyloidstrands
in the pileus, paralleland colored in the lower portion of the stipe and there weakly
pseudoamyloid or nearly inamyloid, all with clamp connections and not gelatinized.
Corticallayer: epicutis of pileus formed by branched-diverticulatehyphae as in M.
androsaceuswith the diverticula 1-5.5 ulong; these elements not hymeniformly arranged
and not or scarcely incrusted by pigment, but hyaline to melleous (by an intraparietal
pigment) both here and in the hypodermium;some of the epicuticularelements cystidi-
form, elongated and diverticulateon the upper side.
On dicotyledonous sticks and wood particles fallen to the ground in Alnetum,
solitary in the montane forests.
MATERIAL STUDIED.ARGENTINA. Jujuy: Lagunasde Yala, 1900-2300 m alt,
13 Feb 1966, Singer T 5131 (F), type, 11 Feb 1966, Singer T 5059 (F).

2. Marasmiuspolylepidis Dennis in Singer, Sydowia 18: 341. 1965.

TYPE.Dennis 173 (K), from Venezuela.
Pileus evenly coloredbetween "kermanshah"and "kaffa" M&P(fuscous), dried
gray brown, not shining but glabrous,smooth at first then finely sulcate-striatein
marginalhalf and there eventually plicate-furrowed,hemisphericalto convex, the center
umbilicatedepressedor not, not umbonate, 3-4 mm broad. Lamellaepaler than pileus
to sordid white with white or whitish edge, intermixed with some lamellulae, about 7-10-
(14) through-lamellae,ratherclose, adnexed with a collarium. Stipe purplish-brown-
black, shining when fresh, glabrous,not setose under a lens, insititious on the substra-
tim, not accompaniedby rhizomorphs, 16-33 X 0.1-0.2 mm. Context very thin, pallid
or concolorous with surface in part, inodorous.
Spores 5.8-8 X 2.8-4 A, ellipsoid, some almost ellipsoid-oblong,hyaline, smooth,
inamyloid. Hymenium: basidia 12.5-17 X 3.5-4i , accordingto Dennis to 20 X 6 u,
and interspersedwith fusiform cystidia; in Yungas materialonly fusiform basidioles
seen and cystidia none; nature basidia 4-spored;cheilocystidia about 6-7 Abroad, hy-
phous and often forked, some swollen to vesiculose and pedicellate, all with mainly
apical diverticulawhich are somewhat irregularlydistributedand short, much as the
epicuticularelements of certain Marasmielli,hyaline, numerous;base of some basidia
Marasmius 67

partly pseudoamyloidat times. Hyphae of hymenophoraltrama and marginof pileus

inamyloid: of interior of stipe and below disc of pileus pseudoamyloid, but intermixed
with some inamyloid ones. Cortical layers: epicutis of pileus consisting of a non-hymeni-
form layer of elements similarto those of the edge of the lamellae, also hyaline.
Hypodermiumconsisting of hyphae in radialarrangement,strongly incrusted by a
deep brown pigment which appearsbright brown in the Melzerreagent.
On dead fallen leaves of dicotyledonous trees in montane forest.
MATERIAL STUDIED. VENEZUELA. Merida: Sierra de Santo Domingo, Laguna Negra,
on fallen leaves of Polvlepis sericea, 3440 m alt, 27 Jul 1958, J. J. Buza & Dennis 173 (K) descr.
& ilL BOLIVIA. La Paz, Nor-Yungas, Mururata, 1400 m alt, 9 Feb 1956, Singer B 1048 (LIL).
ILLUSTRATION.Singer (1965) pl. 5, fig 4.
What appearsto be a geographicrace restrictedto the southern part of the Poly-
lepis area can be collected commonly underPolylepis australisin Argentina. This race
differs but slightly in having non-umbilicatepileus; it is mostly accompaniedby black
rhizomorphs. Characteristicmaterialof this southern form was collected by me in
Argentina:Jujuy, Lagunasde Yala, 16 Feb 1966, T 5202 (F).

3. Marasmiuscyrillidis Dennis, Kew Bull. 22: 76. 1968. Fig 30.

TYPE.DetnnisJ 40, from Jamaica(K).
Pileus cinnamon to dark reddish brown when fresh, rarelypaler and about gil-
vous, tending to become wood-brownishor gray-brownwhen dried (fresh e g between
"Kermanshah"and "kaffa" M&Pin the center, the marginusually paler), smooth at
first, later sulculate to long sulcate, glabrous,convex, sometimes with applanateapex
but neither umbilicate nor papillate, 3-7.5 mm broad, mostly only up to 4 mm broad.
Lamellaepale leather brown to buff-whitish(not pure white except at the edge when
fresh), medium distant, 12-14 through-lammellaeintermixed with some lamellulae,not
or scarcely intervenose, narrow,to ratherbroad, not collariate, broadly adnexted to
adnate. Stipe deep chestnut to blackish-brown,shining when fresh, glabrous,smooth,
insititious, equal or slightly taperingupwards, 15-45 X 0.15-0.50 mm; rhizomorphs
none associated with the stipes. Context very thin, inodorous.
Spores 5.7-8.5 X 2.5-4 p, ellipsoid, ovoid-ellipsoidor oblong-ellipsoid,to cylindric-
oblong, smooth, hyaline, inamyloid. Hymenium: basidia 12.5-22 X 3.5-6.3 p, 4-spored,
sterigmata2-6 A,long; basidioles fusoid: cystidia none; cheilocystidia 9-22 X 5.5-8.5 p,
clavate, some hyphous, and often forked, apically beset with obtuse numerous hyaline
diverticula 1-3.5 X 0.4-1.2 ,. Hyphae filamentous, hyaline, with clamp connections, not
gelatinized, inamyloid or few weakly pseudoamyloid in the trama of the pileus, many
pseudoamyloid in the apex of the stipe includingits interior, where the hyphae are
almost subhyalineand parallel with each other. Cortical layer: epicutis of the pileus
of interwovenversiformelements, not in hymeniform arrangement,these often swollen
to 7 p diameterand some like the cheilocystidia, most distinctly diverticulatebut ir-
regularlyso, fewor many here and almost all in the hypodermiumdistinctly incrusted
by a stramineousor fuscous pigment;hyphae of the hypodermiummainly radially
arranged:hairs of stipe none or inconspicuousat apex of stipes 10-35 X 5-7,u, hyaline
inamyloid, thin-walled,entire, obtuse, often with wavy outline or constrictions.
On leaves of dicotyledonous trees (Quercushumboldtii, Cyrillaracemosifolia,
Coffea arabica and others).
MATERIAL STUDIED. U.S.A. Florida: Highlands Co., Sebring, Highlands Hammock State
Park, 2 Aug 1937, McFarlin 56 (BPI). JAMAICA. High Peak, 22 Dec 1949, Dennis J 40 (K),
type. COLOMBIA. Cauca, Los Araoz, 12 Apr 1968, Singer B 6057 (F),MEXICO. Oaxaca: San
68 FloraNeotropica

Andr6sHidalgo9 Jul 1969,SingerM 8356 (F). BOLIVIA.La Paz:Nor-Yungas belowMlururata,

1400m alt, 9 Feb 1956,SingerB 1048 (LIL,BAFC).
I confused this species with M. polylepidis when first encountered, then separated
it under a herbariumname but did not publish the latter because I believe that M1.cvrill-
idis is the same species. In most of my own collections I find the spores somewhat
broaderthan indicated by Dennis, but young spores are often exactly as those of 1/.
cyrillidis. The incrustationof the hyphae of the hypodermiumis indeed less coarse
than that of M. androsaceusbut I have always found it present. In 1A.androsaceius
the hyphae even of the rest of the pileus and hymenophoraltramaare pigment-in-
crusted whereas in M. cyrillidis they are not.

4. MarasmiusbactrosporusSinger, Sydowia 18: 334. 1965.

TYPE. Singer T 3521, from Argentina.
Pileus beige in youth 1 l-B-4, later 11-C-6M&Pwith fuscous gray ("soapstone")
disc, glabrous, smooth, eventually sometimes more or less uneven, convex, later convex-
applanate, 2.5-3.5 mm broad. Lamellaewhitish, later light sordid gray ("almond, bis-
cuit" M&P),medium close, intermixed, not regularlydi- or tridymous, 12-17 through-
lamellae, medium broad, adnate, sometimes with brown filbrils at the plane of attach-
ment so that they appearcollariate (but not!), eventually somewhat decurrent. Stipe
fuscous, apex concolorous with the lamellae, macroscopicallyglabrousbut hyaline-
pilose under a good lens, smooth, insititious, equal, 11-16 X 0.1-0.7 mm; black rhizo-
morphs accompanying the stipe, thinner than stipe, concolorous with it, also extremely
finely pilose but only partially so, both stipe and rhizomorpheventually glabrescent.
Context subhyalinein pileus, thin, inodorous.
Spores 10-12 X 2.7-3.7 2, rod-shaped,some slightly subfusoid-cylindric,often
with minute internal oil droplets, thin-walled,with a slight suprahilardepression,hy-
aline, smooth, inamyloid. Hymenium: basidia 22-27 X 5.7-6.7u, 2-4-spored,(often
4-spored), clavate, hyaline; basidioles fusoid; cystidia not very conspicuous, thin-walled,
hyaline, saccate, 37-38 X 11-14, ; cheilocystidia versiformand variablein size, about
11-12.5 p broad, hyaline to brownishhyaline, few smooth, most with obtuse short
(rarely reaching2.5 u length) diverticulaall over upper portion, making the edge hetero-
morphous. Hyphae of the pileus and hymenophoral tramahyaline, thin-walled,fila-
mentous, not gelatinized, some inamyloid, some extremely weakly pseudoamyloid;
hyphae of stipe rind deep fuscous, those of apex and interior hyaline to subhyaline
and moderately strongly but distinctly pseudoamyloid, strictly parallel,all with clamp
connections. Cortical layers: epicutis of the pileus consisting of the same sort of
elements which make up the gill edge but intermixed with smallerand hyphous ele-
ments which are likewise diverticulate,hyaline or pale brownish hyaline, not hymeni-
form; pilosity of stipe consisting of scattered thin-walledhyaline hairs which are sub-
cylindric or subampullaceousor subulate, often with widened base, obtuse, rarely sub-
acute, not acute, very rarely forked, not diverticulate,erect, 4-40 X 3-8.5 ,, hypoder-
mium of pileus, a cutis of filamentous hyphae which are subparallelto parallelwith
each other, about 3.5 p broad, strongly incrusted by fuscous pigment; covering of stipe
directly inserted on the rind of the stipe but at plane of insertion of lamellae peg-like
brownish fibrils frequently appearingwhich are homologous to hypodermiumof pileus
and consist of bunches of parallel or subparallelhyphae which are 2.5-8 p thick and
incrusted by fuscous pigment.
On fallen dead leaves of Myrtaceaein subtropical-montaneforest.
Marasmius 69

MATERIALSTUDIED. ARGENTINA.Tucuman:San Javier,ravinenorthwestof CiudadUniv-

ersitariaat 1000 m alt, 9 Jan 1960, SingerT 3521 (LIL), Type.
Singer(1965) pi 4, fig 2.

5. MarasmiusleguminosarumSinger,Sydowia 18: 348. 1965. Fig 31.

TYPE.SingerB 3394, from Brazil.
Pileus brown("auburn"M&P),darkerin the umbilicus,paler on the margin,there
shining,driedmore grayish("suntan"or "tanbark"in the middle, "grain"towardsmargin,
M&P),sulcate, glabrous,convex with an eccentricumbilicus,(2-)5-7 mm broad. Lamellae
grayish,dried sordidisabelline-cinnamon(11 E 6 to "bran"M&P),narrow,distant(about
9 through-lamellae)adnate. Stipe black or blackish,macroscopicallyglabrousbut under a
lens finely pallid-pubescent,glabrescent,insititious, eccentric, about 3-5 X 0.2 mm. Con-
text of the pileus white, or whitish(dried), ratherthin, inodorous.
Spores 6.5-7.5 X 3.3-3.7 ,, ellipsoid to oblong, smooth, hyaline, inamyloid. Hymen-
ium: basidia31 X 6.5 ,, cystidia none; cheilocystidiafilamentousbut some here and there
partly widened, sometimesmoniliform or diverticulate,often ramose,hyaline. Hyphaeof
the pileus tramahyaline, pseudoamyloid,with clamp connections, not gelatinized. Cor-
tical layers: epicutis of the pileus not hymeniform,consistingof elements risingfrom the
hypodermiumand ascendant,often interwoven,similarto the cheilocystidia;hypoder-
mium characteristicallydarkincrustedby the pigment;superficialhyaline hairsboth on
pileus and stipe surface,on pileus rare,on stipe rathernumerous,there thin, mostly acute,
stiff, often incrusted.
On wood of dicotyledonous trees, mostly Leguminosae(fallen twigs, bark, etc).
Known host: Bowdichiavirgilioides.
MATERIALSTUDIED. VENEZUELA. Bolivar:Along Roi Sacaica,7 Aug 1972, Dumont
et at, VE 7288 (NY). BRAZIL. Pernambuco:Dois Irmaos,13 Jul 1960, Sizger B 3394 (BAFC),
The eccentric stipe is characteristic. The collection from Venezuela has no spores.

6. Marasmiusearlei Murrill,North AmericanFlora 9: 258. 1915.

TYPE. Earle 269, from Jamaica(NY).
Pileus whitish with brownish center, in herbariumalmost uniformly colored brown-
ish, glabrous, smooth, then weakly to distinctly-sulculateat least when dry, convex, then
shallowly but distinctly depressedin the center, but neither umbilicate nor papillate,
3-5.5 mm broad. Lamellaewhite, subclose to subdistant(10-24 through-lamellae)some
forked, not intervenose, ratherbroad, adnate, not collariate or subcollariatelyseparat-
ing from the apex of the stipe. Stipe bay above, blackish below, mostly dark bister in
the herbarium,subglabrous,smooth, insititious, subequal,about 4 X 0.2 mm.
Spores 7.8 X 3.7-4 u, oblong, smooth, hyaline, inamyloid. Hymenium: cystidia
none seen: cheilocystidia basidiomorphous,with sinuose outline or subdiverticulate,
about 29 X 7.8 ,. Hyphae of the pileus hyaline, filamentous, with thin to thick (up
to 1 s,) wall, not imbedded in a gelatinous mass, with clamp connections, inamyloid;
hyphae of the stipe parallel, in the interior of the stipe hyaline to stramineous,inamy-
loid. Cortical layer: epicutis of the pileus a weak to ratherstrong continuous Rameales-
structure,there and in the hypodermiumwith an ochraceous intraparietalpigment which
is not or scarcely incrusting;covering of the stipe consisting of small, thorn-likeascen-
dant or erect subhyaline excrescences which are numerousbut not crowded, rising from
the hyphae of the rind of the stipe.
On dead dicotyledonous wood. West Indies.
70 Flora Neotropica

MATERIAL STUDIED. JAMAICA. Castleton Gardens, 28 Oct 1902, Earle 269 (NY), type.

7. MarasmiusliquidambariSinger, sp nov Fig 32.

TYPE. Singer M 8418/8420, from Mexico (F).
Pileo vinaceo-brunneodein cinnamoneo, 5-11 mm lato; lamellis albidis vel cremeis:
stipite subtiliter subpruninato-subflocculoso,rhizomorphisnigris haud adhaeso. Sporis
5.5-8.5 X 2.2-4 j. Ad folia ramosqueLiquidambaristyraciflui in Mexico. Typus in F
Pileus vinaceous brown at first, soon becoming cinnamon with paler margin to
pallid with cinnamon center, glabrous,smooth, over 6/7th of the radius sulcate when
fully mature, conical, then convex, eventually shallowly umbilicate to depressedin the
middle, with or without a papilla in the center of the depression, 5-11 mm broad. Lam-
ellae whitish to cream, eventually avellaneous,close to subdistant,not intervenose,about
18 through-lamellaeand 0-10 lamellulae, narrow to rather broad, subcollariate-separating
or attenuate-subfreeto adnexed. Stipe chestnut to blackish in lower two thirds with
the apex concolorous with the lamellae, subglabrousbut under a lens finely white pruin-
ate subflocculose but eventually somewhat glabrescentand then more or less shining,
insititious, equal, 10-15 X 0.3-0.7 mm; black rhizomorphsthinner than the stipe, gla-
brescent or glabrous,accompanyingthe carpophores. Context white, inodorous.
Spores 5.5-8.5 X 2.2-4 u, mostly 6.5-7.8 X 3-3.3 p, oblong to fusoid, more rarely
ellipsoid, smooth, hyaline, inamyloid. Hymenium: basidia 21.5-23 X 5.5-6.5 ,, (2-)4-
spored;cystidia none; cheilocystidia versiform, e g 33 X 5Li, often fusoid with nodose
to subdiverticulateapex, hyaline, sometimes merely a continuation of the epicuticular
structure;scattered pale melleous incrustationsand other extracellularmaterialoften
found in preparations of the hymenium. Hyphae of the pileus-tramafilamentous with
firm to thick (0.3-1 j,) walls, with clamp connections, not gelatinized, hyaline, some with
fine granularhyaline to pale melleous incrustations,those of the regularhymenophoral
trama either thin-walledor as above; those of the stipe trama all parallel,hyaline or
pale melleous in the core and at first also in the apex, with firm to thick wall, in the
rind a neutral brown and almost all thick-walled,only some strandsweakly to moder-
ately strongly pseudoamyloid, others, especially in the pileus, inamyloid. Cortical lay-
ers: epicutis of the pileus consisting of not hymeniformly arrangedirregular,often
cystidiform elements, most of these distinctly diverticulate(diverticula2-11 X 0.2-1 Iu)
hyaline to pale melleous in the marginalregion by an intraparietalpigment, in young
specimens in the center showing a very thin and intermittent, not conspicuous incrust-
ing resinous pigment, no gelatinization;hypoldermiumlikewise brownish spadiceous
pigmented in a manner similarto that of the epicutis, of thick-walledelements; cover-
ing of the stipe consisting of numerous but isolated cystididioid hairs, these (7-)12-70
X 4-8(-13) M, thin-walled,entire, or with knots and short side-branches,cylindric or
slightly taperingupwardsor with swellings here and there, with rounded tip, hyaline
straightor bent at an angle of ? 90
On branchesas well as on leaves of Liquidambarstvraciflua.
MATERIAL STUDIED. MEXICO. Oaxaca: San Agost'n, about 1850 m alt, 10 Jul 1969,
Singer M 8418/8420 (F), type. Queretaro, Mun. Landa de Matamoros, 5 km W of Lobo, 25 Jun
1959, Guznman3666 (ENCB).

8. Marasmiussplachnoides(Fries) Fries, Epicrisis384. 1838 (sensu Quelet). Fig 33.

Agaricusspachnoides Fries, Systema Mycologicum 1: 137. 1821.
Androsaceus splachnoides (Fries) Rea, Basidiomycetes 531. 1922.
TYPE. Europe (not conserved ?).
Marasmius 71

Pileus whitish with ochraceous-brownishcenter when fresh, sulcate, convex, with

eventually shallowly depressedor subumbonatecenter, 5-7 mm broad. Lamellaewhite,
close to subdistant(10-11 through-lamellae,numerouslamellulae, often didymous), nar-
row to rather broad, not intervenosebut at times anastomosingor forked, adnexed to
adnate, with a tendency to separatefrom the apex of the stipe and becoming subcol-
lariate. Stipe deep chestnut or deep brown, pilose to glabrescent,insititious, equal, 20-
40 X 0.4-0.8 mm. Context thin and white in the pileus, inodorous.
Spores 7-9 X 3-4.2 ,, oblong, smooth, hyaline, inamyloid. Hymenium:basidia
21-22 X 6.5-7.6 u, 4-spored;cystidia none; cheilocystidia forming a dense and contin-
uous (although occasionally interruptedby basidia) fringe, much like the elements of
the epicutis of the pileus, strongly and densely diverticulate. Hyphae hyaline and in-
amyloid in the pileus, pseudoamyloidin the stipe, with clamp connections, mainly
filamentous. Cortical layers: hypodermiumwith thin shee-like, deep brown incrust-
ations in the central portion of the pileus, otherwise consisting of hyaline, inamyloid
elements like the epicutis the elements of which are densely diverticulated;covering
of the stipe consisting of numerousbut not crowded, versiformhairs which are 8-60
X 4.5-9.5 p, either entire, cylindric to subulate with obtuse tips, or elongated and on
one side or all around beset with obtuse broad knot-like diverticula,with hyaline to
brownish-hyalinethin (0.3 u) to slightly thickened (0.6 u) wall.
On fallen leaves of Quercus.
MATERIALSTUDIED. COLUMBIA.Cundinanarca:Cerrode las Mercedes,Mun. Boyaca,
18 Aug 1964, Gurznwn4630 (ENCB). FRANCE. Yerres(S&O),Sep 1933, Romagnesi(LIL).
In the sense of Kihner and Romagnesi,this species appearsto occur mainly on
oak leaves both in the temperatezone of the Northern Hemisphereand in the tropics
south to Colombia (southern limit of Quercus).
Fries, who had seen only dried materialwhen describingthe species, obviously
gave an incomplete but too inconclusive descriptionwhich was emended by Quelet
(1886): the stipe was describedas rufous, which I have not observed on the French
materialnor on the Colombianone, but this color may be present in younger, fresher
specimens. The description given above is exclusively on the basis of neotropical ma-

9. MarasmiushakgalensisPetch, Trans. Brit. Mycol. Soc. 31: 35. 1947.

MarasmiushyalinotrichusSinger,Sydowia 18: 337. 1965.
TYPE. Petch 3920, from Ceylon (K, holotype).
Pileus tawny-cinnamon("Tuscan tan." "Sonora," "Titian gold," "Army brown"
M&P)usually somewhat lighter colored in the marginalregion, glabrous,smooth, soon
slightly to moderately strongly sulcate, convex, with weakly umbonate or umbilicate
center in the flattened to shallowly depressedcentral third, 2-6 mm broad. Lamellae
pale cream to almost concolorous with the marginof the pileus, not pure white in
fresh condition, dried buff to light avellaneous,rathernarrowto ratherbroad, close to
distant (-14 through-lamellaeand inserted with a few lamellulae) often some forked or
even anastomosingbut not intervenose,with entire not discolorous or with whitish
edge, entire, horizontal adnate or collariate adnate; Stipe young colored like the cen-
ter of the pileus, graduallybecoming blackish from the base upwards,long pilose-
hirsute all over (at least when seen under a lens) eventually sometimes glabrescent,
equal, insititious, 7-30 X 0.2 mm black or brown rhizomorphsaccompanyingthe car-
pophores, also pilose or mycelioid, usually thinner at the stipe, long-creeping. Context
thin, inodorous.
72 FloraNeotropica

Spores 5-9.3 X 2-4 u, mostly 6.5-8 X 2.5-3.2 /, oblong, some ellipsoid, smooth,
hyaline, inamyloid. Hymenium: basidia 17-22 X (4-)5.5-7.5(-8) ,1, 4-spored;cystidia
none; cheilocystidia with a main body 13.5-19.5 X 4-6.5 p, versiformbut often clavate,
with diverticula(which are sometimes fasciculate on a finger-likeappendage)mostly
severalto numerous, lateral and apical, 1.2-5 , long, wall hyaline, thin to firm; some
occasionally smooth cheilocystidia with an apical mucro, some basidioles and basidia
occasionally intermixed with the numerouscheilocystidia. Hyphae of the pileus-trama
and the regualrhymenophoraltramahyaline, thin-walled,ratherloosely arrangedbut
not gelatinized, inamyloid or at times vaguely pseudoamyloid,hyphae of the stipe
parallel,here many thick-walledand melleous in KOH,pseudoamyloid or merely vag-
uely pseudoamyloid, all with clamp connections (but often numerous secondary septa
present). Corticallayers: epicutis of the pileus not hymeniform, its elements cylindric
to vesiculose, hyaline or somewhat fuscous in their lower portion laterally and/or ap-
ically diverticulatewith diverticula2-6.5 X 0.8-1.8 1, many of them pigmented by an
incrustingpigment but this incrustationthin, resinous-sheet-like,fuscous or fuscidu-
lous; the same slight to ratherstrong pigment incrustationalso on many elements of
the hypodermium,elements elongated but often branched;covering of the stipe char-
acteristic but somewhat variable,according to the state of development of the stipe,
of three types: (1) hairs 20-100 X 2.7-3.7,, with thin to thick + pseudoamyloidwall
and acute apex, entire and hyaline, fragile but rigid, incrustedby fine crystals, (2) hairs
12-33 X 5-15(-25) , cylindric, ventricose, clavate or vesiculose, thick-walled(wall up
to 1.2 j thick), hyaline to pale brownish, pseudoamyloid or not, (3) hairs(12-)50-170
X 4.2-9.5 /, long-filamentousor graduallytaperingto an obtuse tip, thick-walled,hy-
aline, the inner wall-layerpseudoamyloid,the outer wall layer and the interior inamy-
loid, sometimes incrusted by hyaline, amorphousmaterial,generally of the appearance
and structureof Crinipellishairs, wall 1-4 thick but sometimes much thinner, and
less pseudoamyloid in the apical portion, more numerousthan type (2) and (1) in
the lowest portion of the stipe.
On dead, dicotyledonous leaves, fallen to the ground in tropical forest as well
as the tropical-montaneforests, from Mexico to Brazil, also in tropical Africa and Asia.
MATERIALSTUDIED. MEXICO. Morelos:NWof Tepoztlanat 2,200 m alt, 27 Jun 1969,
SingerM 8208 (F). BOLIVIA. Beni: Vaca Diez: Guayaramerfn, 9 Mar1956, SingerB 1 776/III
(LIL); 9 Mar1956, SingerB 1779 (LIL), type of M. hyalotrichus;17 Mar1956, SingerB 2094
(LIL). BRAZIL. Pernambuco:Camaragibe,6 Jul 1960, SingerB 3117 (BAFC);B3111 (BAFC);
Paraibo:Joao Pessoa, 11 Mar1960, SingerB 3342 (BAFC). CEYLON. Hakgala,Jan 1914, Petch
3920 (FH), isotype. ZAIRE. Bas-Congo,Kisantu,1901, J. Gillet (BR).
When this species is growing on oak leaves, it is sometimesdifficult to disting-
uish from the precedingspecies unless the hairs of the stipe are studied. These are
always entire, smooth in M. hakgalensis;also, the spores are slightly narrowerin the
ILLUSTRATIONS. Singer(1965) pl 6, fig 8-9; Petch (1947), fig 12.

10. MarasmiuscryptotrichusSinger, sp nov Fig 34.

TYPE. Singer M 8250, from Mexico.
Pileo fusco-griseo, 5-6 mm lato; lamellis cremeo-albis,confertis; stipite ad apicem
pilosulo, minus pilosulo basin versus;odore nullo. Sporis 5.8-6.5 X 2.5-3.5 ;' hyphis
fibuligeris,in hypodermio inconspicue pigmento incrustatis. Ad folia dejecta quercina
in Mexico. Typus in F conservatusest.
Pileus fuscous gray, glabrous,sulcate over about half of the radius,convex, ob-
tuse, not umbilicate, 5-6 mm broad. Lamellaecream white, close (about 14 through-
Marasmius 73

lamellae, some lamellulae), with entire edge and not intervenose,adnexed, not collar-
iate. Stipe chestnut to deep chestnut below, with white apex, insititious, pilose at apex,
less or scarcely so below (hairs not visible in fresh condition without a lens), subequal
to slightly taperingupwards,insititious, about 40 X 0.3 mm; dark rhizomorphsaccomp-
anying the carpophores. Context thin, inodorous.
Spores 5.8-6.5 X 2.5-3.5 g, ellipsoid to oblong, hyaline, smooth, inamyloid. Hy-
menium: basidia 19-21 X (4-)5.5-6 i, 4-spored;cystidia none; cheilocystidia like the
elements of the epicutis. Hyphae in pileus-tramaand the regularhymenophoraltrama
mostly thin-walled,not gelatinized, hyaline, inamyloid, with clamp connections, in the
stipe parallel,some with thin, some thick (1.3 p) or intermediatewall, multiseptate,
varyingbetween hyaline and brown, inamyloid, the brown ones very indistinctly pseu-
doamyloid. Corticallayers: epicutis of the pileus consisting of not hymeniformly ar-
arrangedversiformelements, these mostly erect and often cylindric to clavate, inamy-
loid, 7-10 X 4-7 p (main body), sometimes branching,some hyphous, all or most divert-
iculate, the hyphous ones often arisingfrom the clavate ones, hyaline, setulae or divert-
icular equal or more often taperingupwards,to a narrowbut obtuse tips, 3-5.2 X 1-(1.2)
L;hypodermiumcontrastingwith the epicutis in its pigmented elements which are
rather irregularin shape, mainly radially-elongatedwith hyaline to brownish wall and
thinly incrusted by an inconspicuous sheet-like spadiceouspigment-incrustation;cover-
ing of the stipe consisting of at apex denser and more conspicuous, towards base less
conspicuous hairs of two types (1) long (100-180 X (2-)5-10l), the broadenedbase-
portion up to 25 , long, the apex thin and piliform, 2-3 Abroad, with obtuse tip, at
times with secondary septa here and there, all walls inamyloid, hyaline and thin; (2)
short elements about (9-)24-26 X 8-10 , with thin to moderately thickened wall which
is hyaline to melleous hyaline, correspondingto the basal portion of the long hairs,
rounded obtuse above and not effilate, also inamyloid, all entire.
On dead leaves of Quercus,solitary. Mexico.
MATERIALSTUDIED. MEXICO. Morelos:NWof Tepoztlan,27 Jun 1969. SingerM
8250 (F) type.
This differs from other species growing on oak leaves in the characteristicsof the
hairs of the stipe which are not thick-wallednor acute (as in M hakgalensis)nor long
and diverticulate(as inM. splachnoides)but at least many of them characterizedby
the thin-walled,effilate apex. The spores are rather small in comparisonwith those of
M splachnoidesand relatively broaderthan those of M. hakgalensis.
Marasmiuscryptotrichus differs from the following species (which is closest) in
the presence of effilate hairs on the stipe, the more fuscous-grayrather than tan, och-
raceous brown color of the pileus and the less diversifiedcheilocystidia as well as by
the host.

11. MarasmiuschiapasensisSinger, sp nov Fig 35.

TYPE. SingerM 8969, from Mexico.
Pileo brunneo, 4-5.5 mm lato; lamellis albis, subconfertis;stipite sub lente sub-
tiliter pruinoso vel pubescente. Sporis 6-7.2 X 2.5-4 ,; cheilocystidiis dimorphis;hyphis
plerumquefibulatis, in hypodermio fortiter pigmento incrustatis. Ad folia delapsa
dicotyledonea in Mexico. Typus in F conservatusest.
Pileus (ochraceous)-brown,with paler marginalregion (fresh about "oak" in cen-
ter, "papyrus"toward margin,M&P),dried varyingfrom pale tan to almost blackish
brown (in center), convex, eventually and on drying with depressedcenter and dried
sometimes with a slight umbo in the depression,somewhat ruguloseand obscurely
74 Flora Neotropica

sulcate, 4-5.5 mm broad. Lamellaewhite, subclose, with 11-16 through-lamellaeand

some lamellulae, medium broad, adnate or adnexed, not collariate. Stipe dark chest-
nut to fuliginous and somewhat shining, setiform, appearingglabrousbut with a fine
pallid pruina or pubescence when seen dried under a lens, insititious, stuffed, equal,
16-19 X 0.1-0.3 mm; accompaniedby thin, glabrous,black rhizomorphs. Context thin,
Spores 6-7.2 X 2.5-4 p, mostly 3-3.3 p broad, ellipsoid to oblong or subfusoid,
hyaline, smooth, inamyloid. Hymenium: basidia 15-18 X 4-6 p, 4-spored or 2-4-spored;
basidioles fusoid, soon becoming clavate, with basal clamp; cystidia none or a few
cystidioles present, these like the cheilocystidia of type III; cheilocystidia of three
types, (1) broadly ventricose-clavate,22-23 X 9 p, hyaline, scarce, (2) similar,but
beset with apical and subapicalsetulae, main body about 12 X 7 j, setulae 2-6 X 1-2
,u, (3) fusoid-mucronateto ampullaceous20330.5 X 5.5-7 u, ratherfew; often one or
another type missing. Hyphae of the regularhymenophoraltramaand the pileus-trama
hyaline, filamentous, with clamp connections (but many secondary septa present), with
thin to somewhat thickened wall, inamyloid, not gelatinized;hyphae of the stipe par-
allel, thick- or thin-walled,hyaline in the interior and the apex of the stipe, brownish
in the rind, pseudoamyloid in the rind, slightly pseudoamyloid to inamyloid in the
hyaline portions. Cortical layers: epicutis of the pileus not hymeniform, consisting o f
finely diverticulatedversiformelements, none or few of these cystidioid, hyaline, or
with very scattered pigment incrustationsbut in the hypodermiumwith a strong, sheet-
like spadiceous or fuscous pigment incrustation;covering of the stipe consisting of
cystidioid, smooth, entire hairs which are 9-39 X 6-12.5 p, cylindric or ventricose,
rarely subclavateor subvesiculose,with broadly rounded-obtusetip, without effilate
appendagesor setulae anywhere, with stramineous-ochraceous,pale melleous. or sub-
hyaline thin to sometimes somewhat thickened (0.2-1 1u,most frequently 0.4 p), in-
amyloid wall; acute, incrustedand long, thick-walledhairs absent.
On tough coriaceous leaves of dicotyledonous trees in the leafmold of tropical
forest. Mexico.
MATERIAL STUDIED. MEXICO. Chiapas, between Finca Sospiro and El Pozo. 4 Aug
1969, Singer M 8969 (F), type; M 8943 (F).

12. MarasmiusradicellicolaSinger, sp nov

TYPE Singer B 4078, from Brazil.
Pileo brunneo, 2.5-3.5 mm lato; lamellis sordide ochreis, pro ratione latis, sub-
confertis vel subdistantibus;stipite sub lente subtiliterpruinato vel subpubescente;
rhizomorphisnullis visis; odore nullo. Sporis 4-7.3 X 3-3.7 p; hyphis fibuligeris,in
hypodermio spadiceo vel sepiaceo pigmenti incrustationecausa. - Ad radicesarborum
in Brasilia. Typus in F conservatusest.
Pileus brown, at first with whitish margin, fresh and dried sulcate over one
fifth to one half of the radius,glabrous,convex, obtuse, rarely slightly papillate, even-
tually often slightly depressedin the center, 2.5-3.5 mm broad. Lamellaedirty ocher
(not white when fresh or dried), relativelyvery broad, subclose to mostly subdistant
(about 7-12 through-lamellaeand some lamellulae), rounded-adnateto emarginate-
adnexed. Stipe black, at first with chestnut to white apex, setose, macroscopically
glabrousbut under a lens finely pruinate to subpubescent,shining, smooth, insititious,
10-19 X 0.1-0.2 mm; black rhizomorphsnot accompanyingthe stipes. Context pallid,
very thin, inodorous.
Spores(4.5-)5-7.3 X (2.7-)3-3.7 p, ellipsoid, smooth, hyaline, inamyloid. Hymen-
Marasmius 75

ium; basidia 15-17 X 4.5-6 A, 4-spored, few 2-sporedat times; basidioles fusoid; cystidia
none; cheilocystidia versiform, many diverticulatelike the diverticulateelements of the
epicutis. Hyphae of the pileus-tramathin-walled,with clamp connections, hyaline, fila-
mentous, inanyloid to very slightly pseudoamyloid in places; hyphae of the stipe all
parallelwith each other, some dark spadiceous, some hyaline, some thick-walled,some
thin-walled,some pseudoamyloid, some inamyloid (even in the apex of the stipe), 1-6
, broad, with scattereddepth and some secondary septa. Corticallayers: epicutis of
the pileus consisting of diverticulatehyphae and broom cells of various shapes, with
diverticula2-10 mostly about 3 ,uhigh,hyaline; hypodermiumat least in the middle
of the pileus distinctly spadiceous to sepia incrustedby sheets of pigment, its elements
irregularlyinterwoven;covering of the stipe consisting of numerousbut mostly small
(4-29 X 5-11 I, rarely longer) obtuse hairs, those of the upper portion of the stipe
all thin-walledand hyaline to pale stramineous(wall 0.3-0.5 j thick), those of the
lower portion of the stipe as above, or, fewer, hyaline to brown and thick-walledbut
otherwise similar, wall here 1-3 p thick.
On dead roots and woody rootlets of trees (Ficus?), gregarious. Brazil.
MATERIALSTUDIED. BRAZIL. Pernambuco,Dois IrmSos,14 Apr 1964, SingerB 4078
(F), type.

13. Marasmiusdysodes Singer, Sydowia 18: 335. 1965.

TYPE. Singer B 1690, from Bolivia.
Pileus light beige-grayish(between "lariat"and "walnut taffy" M&P)when dry,
unicolorous, glabrous,,almost smooth, convex, not umbonate but slightly depressedin
the center, 6 mm broad. Lamellaewhite, becoming deep brown dotted and eventually
entirely dirty gray brown, ratherbroad, subdistant or moderately close (about 10
through-lamellae),intermixed with a few lamellulae,adnate, not collariate. Stipe black,
smooth, macroscopicallyglabrouswhen fresh but with small, erect, white hairs when
seen under a lens, pilose from apex to base, setose but slightly broaderin the middle
(0.2 mm), 19 mm long, insititious on the substratum;accompaniedby blackish-brown
rhizomorphswhich are smooth and glabrous. Context very thin, with a strong odor
of garlic.
Spores 6.2-9 X 3.5-4.8 A, ellipsoid to pear-shaped,hyaline, smooth, inamyloid.
Hymenium: basidia 18-26 X 5.5-7 p, clavate, hyaline, 2-4-spored,not clamped at the
base; cystidia none; cheilocystidia unevenly distributed,hyphous and nodose or clavate
to vesiculose and diverticulate,in the latter case 13-17 X 10-12 j and similarto the
Rotalis-type of epicuticularbroom cells, diverticularcolorless, some few terminalel-
ements of edge without any diverticular Hyphae without clamp connections, inamy-
loid or almost so, but the inner-mostwall-layerof the thick-walledelements of the
context often pseudoamyloid,excepting the few connecting hyphae, the others
thick-walled,subhyaline to melleous, and strictly parallelwith each other. Cortical
layers: epicutis of pileus consisting of versiform elements, varying from hyphous
to swollen in the manner of the cheilocystidia, diverticulate,diverticulamelleous hy-
aline to pale melleous, 1-3.5 X 1-1.5 p; hypodermiummore or less thick-filamentous,
its hyphae roughishfrom incrustingpigment which is not conspicuous since it is very
pale melleous, often rathervoluminous;hairy covering of the stipe made up by erect
seta-like, but hyaline to melleous-hyaline,stiff hairs which are thick-(1.5 A) walled,
6-10 X 5-7 p, cylindrical or taperingslightly towards the apex which is almost always
always broadly rounded and smooth and entire, the innermost wall layer more or less
76 Flora Neotropica

On dicotyledonous leaves fallen to the ground in rain forest.

MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez, Guayaramerin, 7 Mar 1956, Singer B
1690 (LIL) type.
Singer(1965) pl 5, fig 3.

14. MarasmiusatroincrustatusSinger sp nov

TYPE.SingerM 8320, from Mexico.
Pileo cinnamomeo-alutaceo,plerumque3-3.5 mm lato; lamellis pallide isabelli-
nis; stipite pallide pubescente, glabrescente.rhizomorphishaud adhaeso, 17-19 mm
longo; odore alliaceo. Sporis 7-8.5 X 2-4 p; hyphis defibulatis, in hypodermio pig-
mento sepiaceo fortiter incrustatis. Ad folia dicotyledonea delapsa in Mexico. Typus
in F conservatur.

14a. Marasmiusatroincrustatusvar atroincrustatus

TYPE. Singer M 8320, from Mexico.
Pileus cinnamon buff, sulculate on margin,glabrous,convex, eventually slightly
applanate or depressedin the center, and neither papillate nor umbilicate, but some-
times with a slight umbo in the depressionwhen dried, 3-3.5-(6) mm broad. Lamellae
not pure white but pale isabelline or dull cream, medium close to subdistant(e g 11
through-lamellaeand some lamellulae), not intervenose,adnexed, at times partly or
entirely subcollariate. Stipe chestnut, dried brown or nearly black, the apex at first
white or somewhat paler, later concolorous, pallid-pubescent,glabrescent,solid, in-
sititious, setiform, equal, 17-19 X 0.2-3 mm; associated with but not rising from dark
rhizomorphswhich are also glabrescent. Context thin, white, with a distinct garlic
Spores (6.5-)7-8.5 X 2-4 ,, mostly 2.7-3.5 p broad, ellipsoid, oblong or subcylin-
drical,hyaline, smooth, inamyloid. Hymenium: basidia 19.5-23 X 5.5-7 ,, 4-spored;
cystidia none; cheilocystidia versiformwith a main body 15-24 X 3.5-7 1, diverticul-
ate with diverticulaerect to obliquely ascendant or curved 2-12.5 X 0.7-2.5 p,,all
hyaline. Hyphae of the trama of the pileus and hymenophore hyaline and filament-
ous, not or scarcely pseudoamyloid, not gelatinized, those of the stipe parallel,sub-
hyaline to hyaline in the core and at apex of stipe, brown in the rind, all without
clamp connections, most thick-walled,weakly and incompletely but undoubtedly
partially pseudoamyloid, especially in the rind, multi-septate. Cortical layers: epicu-
tis of the pileus not hymeniform, its elements towards the upper surface beset with
hyaline diverticular2-7.5 X 1 u, these elements versiformand 3-4 p broad, hyaline,
but some, like most of the hypodermium,incrustedby a distinct spadiceouspigment
incrustation, some cystidiform and then much like the cheilocystidia; hypodermium
hyphous, the broaderor all elements with abundant dark pigment incrustation,with-
otu clamp connections; covering of the stipe ,consistingof Crinipellis-likehairs, these
29-90 jplong and 4-7 u broad, thick-(1-2 l)walled, cylindric or taperinggraduallyto an
obtusate tip, all entire and smooth, most of them distinctly pseudoamyloid.
On dicotyledonous, dead, fallen leaves in tropical-montaneforest. Mexico.
MATERIAL STUDIED. MEXICO. Oaxaca: Rancho del Cura, 7 Jul 1969, Singer M 8320
(F) type.

14b. Marasmiusvar inodorosus Singer,var nov Fig 36.

TYPE.SingerM 8816, from Mexico.
Odore nullo.
Marasmius 77

This differs from the type variety mainly in inodorous carpophoresand perhaps
slightly less elongated spores (6.5-7.5 X 3.5-4 j) and somewhat paler marginand stripes
along the ridges of the sulci of the pileus.
On oak leaves in Mexico.
MATERIAL STUDIED.MEXICO.Chiapas:15 kmWof SanCristobal de lasCasas,2400
m alt 2 Aug1969,SingerM 8816 (F), type of variety.

15. Marasmiusatlanticus Singer, sp nov Fig 37.

TYPE.Singer F 347, from Florida, U.S.A.
Pileo rubescente-brunneoin centro, albo in margine;lamellis albis, sat crassiusculis,
confertis; stipite spadiceo-sepiaceo,fibrilloso-furfuraceo,elongato. Sporis 7.5-8 X 3.3 u;
hyphis defibulatis, incrustationibusinconspicuis. Ad folia quercinain Florida. Typus
in F conservatusest.
Pileus reddishbrown in center, white on margin,grooved-sulcate,glabrous,size
and shape as in Marasmiusandrosaceus. Lamellaewhite, rather thick, somewhat ir-
regular,close, adnate. Stipe deep spadiceous-sepia,somewhat shining under a dense
white fibrillose-furfuraceouscovering, insititious, elongated.
Spores 7.5-8 X 3.3 g, oblong, hyaline, smooth, inamyloid. Hymenium:
basidia 16-22.5 X 6-7.3 ,u;basidioles fusoid; cystidia none; cheilocystidia very irregular,
in some parts swollen but not vesiculose, often clavate and entire, main body e g 16-27
X 7-8 ,, some diverticulateor with side branches,hyaline, wall up to 0.5 p,thick. Hy-
phae without clamp connections, hyaline and inamyloid in the trama of the pileus and
the regularhymenophoraltrama, deep brown, paralleland pseudoamyloidin the rind
of the stipe, there with thick walls. Corticallayers: epicutis of the pileus not hymen-
iform, a rather thin layer with terminalcells often but not consistently showing wavy
outline or irregulardiverticula,here and in the hypodermiumsome sheet-like bt., very
thin and inconspicuouspigment incrustationspresent, but these intermittent, dark
brown; coveringof the stipe consisting of cystidioid hairs which are dense and often
interwoven, somewhatirregular,1244 X 6.5-13 p, fewer hyphous and 3 1 broad, sub-
hyaline to hyaline, mostly ending in a clavate or ventricose apex which may be appen-
diculate by one oblique finger-likeappendage,few sepia-brownand thick-walled,most
thin- or firm-walled,0.2-0.5 p thick, some of these cystidioid hairs also present on the
epicutis of the pileus where they are rare to scattered.
On leaves of Quercuslaurifoliain subtropicalforest.
MATERIALSTUDIED. U.S.A. Florida:HighlandsCounty, HighlandsHammockState Park,
21 Aug 1942, Singer F 347 (F), type.
This differs from the other oak leaf inhabitingspecies in the absence of clamp

16. Marasmiuseorotula Singer, sp nov Fig 38.

TYPE.Singer B 6380, from Colombia.
Pileo albo, 2-7.5 mm lato; lamellis albis, plus minusvecollariatis;stipite albo, ad
basin interdum pallidissimestramineo, 4.5-25 mm longo; odore nullo. Sporis 6.5-8 X
25-3 1; lhyphisdefibulatis. Ad folia dicotyledonea delapsain Columbia. Typus in F
Pileus white, glabrousor slightly furfuraceousunder a strong lens, smooth or
with short-folded-furrowedmargin,convex, obtuse, often with flattened or somewhat
narrowly depressedcenter when old or dried, 2-7.5 mm broad. Lamellaewhite, med-
ium close to subdistant(about 12 through-lamellae),inserted with some irregularly
78 Flora Neotropica

disposed lamellulae,horizontal, moderately broad to ratherbroad, not intervenose,

adnexed but removed from the stipe and connected by a more or less distinct collar-
ium leaving a radially fibrillose, narrowzone between the collarium and the apex of
the stipe. Stipe white, sometimes becoming palest stramineousbelow, never dark at
base, glabrousmacroscopicallybut minutely flocculose or pruinate in the lower por-
tion or all over when seen dried under a lens, insititious and equal or with a vaguely
disc-like wideningat the base, hollow, 4.5-25 X 0.1-0.3 mm. Context white, thin,
Spores (5.5-)6.5-8 X 2.5-3 ,, ellipsoid to mostly oblong, hyaline, smooth, inamy-
loid. Hymenium: basidia(14-)17-22 X 4.5-7 p, all 4-spored, or with some 2-spored
ones intermized;cystidia none; cheilocystidia 14-28 X 4-16j, clavate to vesiculose,
with apical and subapicalas well as often also divergentlateral diverticulaor setulae,
these varying from finger-liketo verruculose,very variablein size but mostly 2-4 X
1E, the main body often forked or tripartiteand each fork beset with setulae, the
wall thin and hyaline. Hyphae in the pileus-tramahyaline, filamentous, non-gelatinized,
inamyloid or very slightly and slowly pseudoamyloid, without clamp connections, walls
more often thin than thick (0.2-1.2 p thick); hyphae of the stipe like those of the
pileus but at least some of them distinctly to strongly pseudoamyloid,all parallel.
Corticallayers; epicutis of the pileus in places almost subhymeniformor in palisade
but in others irregularlyarranged,consisting of terminal bodies which are more or
less erect ot ascendant, 14-38 X 6-27 ,, beset with numerous diverticulalike those of
the cheilocystidia, also often branchedor appendiculatewitihdiverticulatedbranches
or appendages,the appendagesof the ascendantcells restricted to their upper surface,
with thin, hyaline, inamyloid wall, without basal clamp, the main body clavate to
vesiculose and sometimes pedicellate, some diverticulate elements repent and hyphous:
hypodermiumhyaline, of elongated horizontal hyphae inter-mixedwith occasional
spherocystoid elements; covering of stipe consisting of elements which are 10-50 X
5-12.5 p, sometimes catenulate, very versiform, but mostly clavate or ventricose, cylin-
drical or ampullaceous,vesiculose or conic-obtuse, sometimes mucronate with thin or,
more rarely, up to 1 thick, hyaline to pale stramineouswall which may be thickened
by duplication or triplication, especially at the apex of those elements which are en-
tirely smooth and entire.
On fallen dicotyledonous coriaceous leaves in tropical rain forest near West Coast
of Colombia.
Campamento Pulpapel25 Apr
1968,SingerB 6380 (F), type. SanJoaquin,22 Apr 1968,SingerB 6290 (F); 19 Apr 1968,
SingerB 6234 (F).
This has the appearanceof a representativeof the section Marasmiusor of Mara-
smiellus sect Defibulati, but clearly differs from the former innon-hymeniformepi-
cutis and the irregularand variableshape of the setulae, and from the latter in the
distinctly pseudoamyloidhyphae of the stipe. It differs from all other species of
section Androsacei in the almost complete absence of pigment, in the carpophores.

17. Marasmiusdefibulatus Singer, Sydowia 12: 127. 1958.

TYPE.D. H. Linder, from Surinam.
Pileus whitish, glabrous, rugose-striate,convex, somewhat papillate, 2-4 mm
broad. Lamellae whitish with white edges, subclose to ratherdistant (10-12 through-
lamellae, intermixed with shorter ones), narrowto broad, mostly broad, collariately
adnexed to a narrowand sometimes not too distinct collarium. Stipe black, macros-
Marasmius 79

copically glabrous, shining, setiform, about 20 mm long and up to 0.5 mm broad,

insititious, accompaniedby but not attached to black rhizomorphs.
Spores 6-9 X 3.2-4.5,, mostly 7.8-8 X 4,, ellipsoid, smooth, hyaline, inamy-
loid. Hymenium: basidia 18-26 X 4.5-6 M,2-spored;basidioles fusoid; cystidia none;
cheilocystidia ventricose to subvesiculoselike the epicuticularbroom cells but hyaline.
Hyphae hyaline, not gelatinized, regularin the hymenophoraltrama, there 2-3.7 ,lbroad,
inamyloid. Corticallayers: epicutis of the pileus not hymeniform, consisting of ap-
pressed, multiseptate hyphal elements, the terminal ones mostly cystidioid and densely
and regularlybeset with setulae, main body vesiculose to ventricose, often constricted
in the middle but also hyphous, setulae also on the hyphal ends, directed mainly up-
wards, 2-3 X 0.8-1.2,, all hyaline, many terminalcells ascendant or erect, all inamy-
loid; covering of the stipe consisting of hairs 18-530 X 3.5-10,, hyaline, taperingup-
wards graduallybut with rounded tip, entire, thick-walled(wall 0.7-4 ,).
On moist, decaying palm fronds.
MATERIAL STUDIED.SURINAM.Coppename River,CalabackCreek,13 Nov 1923,D.
H. Linder (FH), type.
Singer (1958a), fig 38.

18. Marasmiustomentellus Berkeley & Curtis, Jour. Linn. Soc 10: 298. 1868.
Fig 39.
TYPE. Berkeley Herbarium(K), from Jamaica(not studied).
Pileus fulvous to vinaceous brown, subrugulose,not sulcate, later more or less
sulcate, glabrous,convex, obtuse or with a small and obtuse umbo, 1-3 mm broad.
Lamellae becoming concolorous with the pileus in the herbarium,few (? 10) distant,
more rarely subdistant, with few lamellulae(1-5), not intervenose, not collariate, ad-
nexed. Stipe badious to blackish, pubescent, curved, rising from a common black
rhizomorphs,solid, subequal with slightly to distinctly narrowedapex, 1-4 X 0.2-0.6
mm; rhizomorphsblack, long, as thick or thicker than the diameter of the stipe.
Spores 11-11.5 X 4-5 ,, cylindric to oblong with mostly recurvedbasal portion
and flattened inner outline when seen in profile, smooth, hyaline, inamyloid. Hymeni-
um: basidia 27 X 6.5-8.5 ,, 4-spored;cystidia scattered, 60 X 6.5-7.5 1, ampullaceous
or fusoid, thin-walled,hyaline. Hyphae filamentous, inamyloid except for the pseu-
doamyloid rind-zoneof the stipe, without clamp connections, hyaline except for the
hypodermialzone of the pileus and the rind-zone of the stipe, which are pigment-in-
crusted, not gelatinized, those of the pileus radiallyarrangedand those of the stipe
parallel. Corticallayers: epicutis of the pileus consisting of irregularlyarranged,ir-
regularlydiverticulatedelements which are hyaline to subhyaline, forming a very thin
layer; liypodermiumvery broad, consisting of interwoven hyphae with conspicuous
brown pigment incrustation, 2.5-8 , broad, with walls up to 0.8 , thick; covering of
the stipe consisting of few erect hairs (which are merely continuations of the hyphae
of the rind of the stipe, about 25 X 5 ,', entire, with the wall about 0.7 ,uthick) and
seta-like hairs 50-125 X 5-7 ,, with hyaline, thick (1 ,) pseudoamyloidwall.
On heaps of dead twigs in accumulatedlitter. Southern U.S.A. and West Indies.
MATERIALSTUDIED. U.S.A. Louisiana:La Salle parish,May 1936, FrankSmith, comm.
F. A. Wolf (FH).

Marasmius sect Hygrometrici Ktlhner, Le Botaniste 25: 60. 1938 (as Hygrometricae).
TYPE SPECIES. Marasmius hygrometricus (Briganti) Saccardo.
Pileus usually small and well pigmented with an epicutis formed by broom cells
(usually of the Rotalis-type) which, in portions of the pileus form a hymeniform lay-
80 Flora Neotropica

er, often some of the epicuticularelements in chains, often some of them smooth,
(but never all of them); spores usually medium-sized;lamellae free to adnate, not or
incompletely collariate, rarelyabsent; cystidia on the sides of the lamellae usually
present or cheilocystidia dimorphic. Hyphae with clamp connections, inamyloid.
Stipe central, always insititious. On dead vegetable matter.

Key to the Species of Section Hygrometrici

1. Pileusrough or spiny from chains of thick-walled,often ornamentedcells, accompanied
by singlebroom cells.
2. Spores8.7-9.3 X 3.5-4.3u1.OnMagnoliain Florida. 22. M. magnoliae.
2. Spores6-8.5 X 3.2-4u. On variousmonocotyledonousand dicotyledonousbranches,
petiolesand leaf bladesin tropicalAmericaand Africasouth of the Equator.
23. A. echinosphaerus.
1. Pileus smooth or merely radiallysulcate, not with chainedcells, only with broom cells
of the Rotalis-type,some of these occasionallysmooth.
3. Lamellaeabsentor hymenialsurfaceradiallyveined. 21. M. sphaerodermus.
3. Lamellaedistinct, 7-14 through-lamellae.
4. Sporesmore than 3.3, broad,at least manyof them.
5. Pileus brownish-yellowto yellow-brown, 1-7 mm broad; lamellae not pure
white when fresh; stipe macroscopicallynot glabrous(see note underM. ilicis:
M. corbariensis).
5. Pileussmaller(1-3.5 mm) and differentlycolored.
6. Pileus brownishwhite, drying fuscous-brown(see note underM. ilicis: M.
6. Pileuswith an orange-brownto chestnuttone. 19. M. ilicis.
4. Sporesup to 3.3, broad.
7. Pileus 2-5 mm broad, ochraceous brown to brownish ochraceous, 7-10
through-lamellae present. 20. M. crescelntiae.
7. Pileus 1-2 mm broad;colored as indicatedabove or more sepia-ferruginous to
deep ferruginousor grayish-badious;4-8 through-lamellaepresent (see note
underM. crescentiae:M. capillipes).

19. Marasmiusilicis Singer, Lilloa 26: 142. 1953.

Marasmiussphaerodermusvar majorSpegazzini,Anal. Soc. Ci Argentina10: 163. 1881.
TYPE. Singer B 88/88a, from Brasil.
Pileus orange brown to chestnut brown ("spice" 13-B-12 M&Pto deeper red-
dish brown) with pallid to light saffron orange (almost "saffrony" i e paler than 10-
K-8) marginbetween striae or uniformly colored, disc reaching"Titian gold" (deep
cinnamon brown) dried entirely cinnamon to sordid cinnamon, sulcate over the larger
portion of the radius, convex to campanulatebut tending to have truncate or at
least somewhat applanate disc, glabrous, 1-3.5 mm broad. Lamellaewhite with white
edge, seven to ten, in mature caps also some few lamellulae occasionally present, nei-
ther intervenose nor collariate, moderately broad to rather narrow,subfree to adnate,
rather distant to distant. Stipe young umber to fuscous or brown ("cowboy, new
bronze" to "burnt umber M&P)eventually black, glabrous, smooth, shining, young
with white apex, insititious with a very small ring, 5-40 X 0.1 mm. Context extreme-
ly thin, white, unchanginginodorous.
Spores 6.7-9.5 X 3.5-5 ,, ellipsoid to ellipsoid-fusoid,thin-walled,hyaline, smooth,
often granularinside, inamyloid. Hymenium: basidia 22 X 7.5 j,; basidioles fusoid;
cystidioles and cystidia fusoid like those of the Epiphylli, hyaline, "empty," with an
ampullaceousto subcapitateapex, smooth, projecting 20-27 X 4-7 A; cheilocystidia
scattered and similarto epicuticularbroom cells. Hyphae of the pileus hyaline, not
gelatinized, filamentous, inamyloid, with clamp connections, 2-6 p,broad, thin to firm-
(wall up to 0.4 , thick)walled. Hymenophoraltrama regular,made up by similarhy-
Marasmius 81

phae; hyphae of the stipe parallel,those of the rind pigmented by a subfulvous,intra-

parietal pigment, 2-7 u broad, some inflated to 11.5 p, inamyloid. Corticallayers:epi-
cutis of the pileus hymeniform, consisting of broom cells of the Rotalis-type which
are 8-29 X 6.5-15 ,, hyaline (often rare) to melleous, to deep spadiceous or chestnut,
setulae subhyaline to melleous or deep spadiceous,(0.7-)1-1.2-(4.5) X 0.5-1.2p, the
wall often thickened up to 3.5 , at the apex of the main body; covering of the stipe:
broom cells (e g 11 X 4.2 ,) brownish to brownish-hyaline,versiform, often clavate
or cylindric with sterigma-likeoutgrowths up to 2.5 pprojecting;dermatocystidialike
the cystidia of the hymenium also present in most collections, these hyaline, e g
23 X 8 ,, apex 7.2 X 1.8 ,; superficialhyphae of stipe often distantly diverticulateby
spinose diverticula.
On dead dicotyledonous leaves, petioles, stems, most frequently on fallen cor-
iaceous leaves and petioles of Ilex and other trees and shrubsin tropical, subtropical
and marginal(gallery) forests. Known hosts: Ilex, Ligustrum,Persea,Eryngium.
MATERIALSTUDIED. VENEZUELA. Sucre: Along Rio Media,15 Jul 1972 Dumont et
al VE 5124 (NY). BOLIVIA. La Paz: Nor-Yungas,Rio Llolosa,31 Jan 1956, SingerB 758 (LIL).
BRAZIL. Rio Grandedo Sul: Taimbesinho5 Nov 1951, SingerB 88, 88a (LIL, MICH),type.
ARGENTINA. Buenos Aires: ParquePereyra20 Jun 1962, Horak, comm. SingerS 201 (BAFC).
The Venezuelan collection, the northernmostseen, is the only one that grew on
herbaceous stems; it differs also by slightly broaderspores and less developed cystidia.
These small differences do not seem to justify a special taxon.
I have included in the key two related extralimitalspecies: Marasmiuscorbariensis
Roumeguere(MarasnziushygrometricusBriganti)and Marasmiusexustus Berkeley &
Curtis(Fig 40).
Since Olea is planted in some regions of the neotropics, it is quite possible that
MIarasmliuscorbariensishas been introduced to that region although, as far as I know,
it has never been collected there. Marasiius exustus Berkeley & Curtis has likewise
not been collected in the neotropics but was discoveredon the Bonin Islands;the
type (FH) is much like Mlarasmiusilicis microscopically(spores 5.5-9.2 X 3-4.5 p).
ILLUSTRATION. Singer (1965) pl 7, fig 14.

20. MarasmiuscrescentiaeMurrill,North American Flora 9: 259. 1915. Fig 40.

TYPE.Earle 184, from Cuba.
Pileus ochraceous to ochraceousbrown, center deep ochraceousbrown, dried
rusty ochraceous brown with dark brown center, long sulcate, glabrous,campanulate-
convex, then convex, obtuse, not depressedand not papillate, 2-5 mm broad. Lamellae
cream-whitethen pale dirty-cream,the edge only in the marginalzone narrowlybrown-
marginateor not at all, distant (7-10 through-lamellae,no lamellulae), not or eventually
weakly intervenose, narrow to broad, adnexed. Stipe chestnut to blackish-chestnut,
shining, macroscopicallyglabrous,setiform, hollow, equal and insititious with the base
slightly lobed and volva-likewidened, 10-40 X 0.1-0.2 mm. Context thin, inodorous.
Spores about 6.5 X 2.7,u, oblong, hyaline, smooth, inamyloid. Hymenium: basi-
dia clavate; basidioles fusoid; cystidia on both edges and sides numerous, 22-30 X 7-7
.5 ,, fusoid-ventricosebelow, ampullaceouswith a rathernarrowobtuse apex which is
sometimes subclaviculatebut mostly cylindric or graduallyattenuated upwards,about
2 , broad, smooth and entire; cheilocystidia of the broom cell type also found on the
edges but either very rare or all concentrated in the region near the margin of the
pileus, 18-23 X 7-11 jA, always intermittent and not making the edge heteromorphous,
hyaline below, stramineousto chestnut brown above and there thick (1-1.5 1) walled,
82 FloraNeotropica

the upper three quartersor two thirds covered with rod-like setulae (Rotalis-type).
Hyphae of pileus and hymenophoraltrama hyaline, filamentous, interwoven, with
clamp connections, moderately thick-walled to thin-walled,not gelatinized, inamyloid.
Hyphae of the stipe parallel, in cortex brown and moderately to rather strongly thick-
walled, the paler to subhyalinehyphae of the core thin-walledto moderately thick-
walled, inamyloid. Cortical layers: epicutis of the pileus hymeniform, consisting of
broom cells much like cheilocystidia, some hyaline, some chestnut brown to stram-
ineous, thick-walledabove if pigmented;there are also some smooth, thick-walled,
strongly pigmented bodies, similarto the broom cells in shape but rising from the
hypodermium;they do not reach the level of the epicutis and remain somewhat im-
bedded; covering of the stipe: in the middle of the stipe of an adult specimen con-
sisting merely of some external hyphae which are nodulose-verruculose,some incon-
spicuous brown excrescences which are smooth, very scattered;some dermatocystidia
like the hymenial cystidia, also smooth.
On dead, fallen, dicotyledonous leaves and fruits, known hosts: Crescentia,
MATERIAL STUDIED.MEXICO: Morelos,NWof Tepoztlan,2200 m alt, 27 Jun 1969,
SingerM 8251 (F). CUBA. 5 milesE of Santiagode lasVegas,1 Sep 1904,Earle184 (NY)type.
Another narrow-sporedspecies is Marasmtius capillipes Saccardo(= Miarasmtius
pyrinus Ellis) which differs in size, color and having fewer lamellae. All collections
seen by me come from the temperate regions of Europe and North America,always
on deciduous (not coriaceous) leaves (of rosaceous trees, Sryringa,Acer, Popuhlts,etc.):
neither the fungus nor the hosts have been found in the native vegetation of the
neotropical regions but this species is very close to AMarasnnius crescentiae and might
yet show up in the subtropicalbelts, introduced into gardensor parks. It was indicated
for Chile by Spegazzinibut his descriptivedata and illustration refer to Marasmnius ilicis.

21. MarasmiussphaerodermusSpegazzini, Anal. Soc. Ci. Argentina9: 163. 1881.

Fig 41.
TYPE. SpegazziniHerbarium,La Plata, from Argentina(partly destroyed).
Pileus orange buff to brownishorange ("Saratoga"M&P),scarcely changingwhen
dried, glabrous, smooth, slightly granular,hemispherical,subumbonate,0.3-0.7 mm broad.
Lamellaenone or replaced by few radialveins, hymenial surface cream white. Stipe
maroon, chestnut or brown-blackat first with cream-white,later with paler apex, even-
tually unicolorous dark, macroscopicallyglabrous,insititious, equal or with broadened
base, 3-15 X 0.08-0.12 mm. Context of pileus white, extremely thin, inodorous.
Spores 7-10 X 3.5-5 j, ellipsoid to more rarely ellipsoid-oblong,smooth, hyaline,
inamyloid. Hymenium: basidia 17-20 X 7-7.5 u, 4-spored, with basal clamp; basidioles
fusoid; cystidia 20-33 X 5.507.5 M, ampullaceous,the apex cylindric or slightly clavi-
culate and 2-3 , wide, hyaline, with obtuse tip; there are also, very scattered, some
broom cells in the hymenium, much like the epicuticularbroom cells, but these are
missing in some sections. Hyphae of the pileus hyaline, filamentous, thin-walled,with
clamp connections, not gelatinized, inamyloid. Hyphae of the stipe parallel, dark brown
to hyaline, elongated, the brown ones incrusted by a brown or porphyry brown pigment.
Cortical layers: epicutis of the pileus consisting of broom cells 10-26 X 6-15 u, in hymen-
iform arrangement,varying from hyaline to deep brown, many melleous-ochraceous,
darkest usually in the upper portion, the pigment being concentrated in the thick por-
tion of the wall at the apex (up to 2.5 u thick) and in the setulae which are divergent,
1-2.5 ,llong, Rotalis-type, inamyloid; some dermatocystidiaoccasionally found among
Marasmius 83

the broom cells, the dermatocystidiamuch like the hymenial cystidia; covering of the
stipe consisting of scattered dermatocystidia,much like those of the hymenium but
much smaller,graduallydisappearingin old specimens.
On dead leaves and twigs. Known hosts: Salix humrboldtiana, Spathodea campa-
MATERIAL STUDIED. HAWAII. Manoa Valley, Oahu, Jan 1969, R. D. Goos (F). AR-
GENTINA.BuenosAires:La PlataRiver(no morecarpophores presentin the type envelope),
(LPS),type.Tierradel Fuego:Ushuaia,authenticmaterialbut only somestipesleft (LPS).
This species is evidently of wide distributionbut has been rarely collected, prob-
ably because of its small size. However, Spegazzini'soriginal description(aside from
the spore measurementswhich are obviously wrong) fits the Hawaiianmaterial so well
and the lack of lamellae is so characteristicfor this one species in section Hygrometrici
that I have no doubt but that they are identical with each other.

22. MarasmiusmagnoliaeSinger, Mycologia 37: 435. 1945.

TYPE.Singer F 1906, from Florida, U.S.A.
Pileus deep brown, then "Amber Brown" Ridgway, with the depressionsof the
radiately sulcate-rimosemarginpale buff, eventually somewhat pallescent and the whole
marginalzone "Clay Color" (R) pilose when mature, eventually somewhat glabrescent,
the smooth disc about 0.5 mm broad, the extreme marginfimbriate-ciliateat first,
hemisphericwhen convex, flattened eventually and becoming subumbilicateor with a
slight depressionin the middle of which there may be a small papilla, up to 5.5 mm
broad. Lamellaewhite, distant, entire, equal, moderately broad (1 mm), subfree. Stipe
blackish brown, macroscopicallysubglabrousbut at least partially subfibrillosewhen
seen under a lens, opaque, rarely slightly shining, insititious, more or less flexuous,
slightly taperingat the apex, 10-40 X 0.2-0.5 mm. Context white, very thin, inodor-
Spores 8.7-9.3 X 3.5-4.3 u, most frequently 8.8-9 X 4-4.2 p, ellipsoid-fusoid,
hyaline, smooth, inamyloid. Hymenium: basidia 26 X 6 p; cystidia none seen; cheilo-
cystidia fusoid, about 4-7 v broad, acute, smooth, entire, hyaline. Hyphae of the pileus-
tramahyaline, filamentous, inamyloid, with clamp connections. Corticallayers: epicu-
tis of the pileus consisting of short, vesiculose elements seriate in chains forming
bristle-likehairs (the ornamentationof the pileus surface) these elements beset with
conical to rod-shapeddiverticula2.5 Mhigh; among these "hairs"there are also ascend-
ant clavicularelements also seriate to form long chains and arisingfrom each others
lower-apicalside or forming a chain of normal filamentoushyphae, rarely the hairs
consisting of smooth members of free erect hyphal chains, the individualelements in
all these cases 1140 C 7-17v, the shortest ones e g 27 X 23 ,, all thick-walled,most
with brown walls, the terminalmembers often with blackish brown apex.
On the petioles of fallen leaves of Magnoliagrandiflora,very rarelyalso on other
parts the tree, very gregariousin "bay heads," fruiting in spring.
MATERIALSTUDIED U.S.A. Florida:AlachuaCounty, Gainesville11 May 1943, Singer
F 1906 (FH), type; Newnan'sLake, 12 May 1943, SingerF 1906a (FH).
This species is marginal as far as the neotropics are concerned but since it has
also been found in the subtropical zone, it is here admitted.
ILLUSTRATION.Singer (1958a) fig 8.

23. Marasmius echinosphaerus Singer, Bull. Jard. Bot. 34: 325. 1964.
TYPE.Schmitz-Levecq169, from Zaire.
84 FloraNeotropica

Pileus blackish spinulose on yellowish ground, dried dark bay, reddish chestnut
(e g 15 E 12, M&P),strongly sulcate over half the radius, echinulate-spinuloseall
over, subglobose to hemispheric,often higher than broad, 1.5-3 mm broad, 1.5-5 mm
high. Lamellae white, with the edges concolorous with the pileus but when young
only pale ocher, subdistant(about 10-17 through-lamellae),about one fifth of the
diameter of the pileus broad, attenuate-subfreeto free. Stipe concolorous with the
pileus below but at first with whitish apex, later and dried almost uniformly chestnut
brown, macroscopicallyglabrousbut finely echinulate-flocculose-roughwhen seen under
a lens, insititious, equal, 10-15 X 0.2-0.5 mm. Context rather fleshy and relatively
thick in the middle of the pileus, thin at margin, inodorous.
Spores 6-8.5 X 3.2-4 p, ellipsoid to oblong, smooth, hyaline, inamyloid. Hymen-
ium: basidia 18.5-26 X 6-6.5 u; basidioles fusoid; cystidia both on the edges and in
the lower part of the sides of the lamellae of two types: (1) 14-34 X 2.5-5.5 , ampull-
aceous, lanceolate or acicular,subacute, acute, or somewhat obtusate, hyaline, thin-
walled, often or rarely covered by a finely granularhyaline mass, (2) broom cells of
the Rotalis-type, 10-34 X 4-9 p, vesiculose to subvesiculose,beset with divergentset-
ulae 0.5-3 p projecting, melleous-hyalineto brown above, mostly hyaline below, wall
0.2-0.5 p thick. Hyphae of the pileus-tramahyaline, filamentous, thin-to firm-walled,
not gelatinized, with clamp connections, inamyloid, few swollen to 6.7 p broad; those
of the stipe strictly different abruptly changingat the apex to parallel,hyaline in the
core, but incrusted by a coarse succineous brown pigment in the rind, all firm-to
thick-walled,inamyloid. Cortical layers: epicutis of the pileus consisting of a hymen-
iform layer composed of four kinds of elements (a) broom cells like the cystidia of
type (2), above these reachinga wall diameterat their apex of up to 5 P, often broadly
clavate ventricose of subvesiculoseand more pigmented (these dark, thick-walled,even-
ly distributedamong the others; (b) similarelements agglutinatedin parallel rows con-
sisting of seriate catenulate broom cells and making up the ornamentationof the pil-
eus, in the marginalzone often replaced by large, one-celled broom cells 17-81 X 10-
16 p; the catenulate cells are about 19-36 X 7-20 p, becoming shorter towards the
terminalcells, many of them without setulae or merely rough;(c) occasional dermato-
cystidia correspondingto the cystidia of type (1) or slightly smaller:(d) broom cells
of the Siccus-type or entire, 12-20 X 5 p, often ramose-forkedinto branchesabout 2 P
broad, or simple, with obtuse tips, main body cylindric to slightly ventricose, chestnut
colored, scarce and sometimes wanting;hypodermiuma palisade where the terminal
cells constitute the epicutis, but here less or not pigmented, less or not thick-walled
and occasionally widened into endocystidia of the shape and sixe of the hymenial
cystidia of type (1); covering of the stipe consisting of elements similarto the epicutis
elements of type (a) and (d), the latter scarce and of some versiform rough to sub-
smooth brown, thick-walledelements 18-23 X 3-5 M.
On fallen branches,sticks, leaves and petioles of dicotyledonous trees.
MATERIALSTUDIED. ARGENTINA. Jujuy: Posta de Lozano, 1500 m alt, 21 Feb 1966,
Singer T 5429 (F). ZAIRE. Haut-Katanga,Kipushi,Kipopo, 1250 m alt, Schmitz-Levecq169 (BR)
Singer (1964b), fig 14, E-H.
The African Marasmius kroumirensis (Patouillard) Saccardo & Sydow has less
ornamentedpileus, adnate and fewer lamellae, broadly umbonate pileus and shorter
stipe, according to Pegler's(1966) type analysis.
However, I have found a fungus in Ecuadorwhich is somewhat intermediatebe-
tween M. kroumirensisand M. echinosphaerus. The materialis too scanty to determine
Marasmius 85

whether it belongs to one or another of these species or whether it representsa third

form. It would differ from M. echinosphaerusin being smaller,echinulate-spinulose
only at the apex of the pileus and the spores perhapsbeing somewhat narrower;it
would differ from Al. kroumirensisin being echinulate on the pileus, having a longer
stipe and from both perhapsin the ornamentationof the stipe-hyphae and the occas-
ional insertion on black rhizomorphs. Description see below.

23A. ?Marasmiuskroumirensis(Patouillard)Saccardo& Sydow, Sylloge Fungorum 14:

105. 1899.
Pileus brown, in apical portion brown echinulate, otherwise glabrousand often
glabrescent,sulcate, campanulate,obtuse, about 1 mm broad and high. Lamellaewhite,
distant (about 8-11 through-lamellae),with partiallyeventually brownish edges, adnate.
Stipe black, glabrous,insititious on rhizomorphsor directly on the leaf without a ring
or broadening,5-18 X 0.1-0.2 mm; black rhizomorphsabout 0.2-0.3 mm thick, glabrous,
scarce, but many telepods present. Context very thin.
Spores (few seen) 6-9 X 3-3.5 ,, smooth, hyaline, inamyloid. Hymenium: basidia
12 X 6.5 p, 4-spored;basidioles short-fusoid-mucronate,some cystidioid (sterile); broom
cells on the side of the lamellae none, on the edge of the lamellae as cheilocystidia
numerousand much like the epicuticularbroom cells. Hyphae inamyloid with clamp
connections, stramineousin the stipe but more brown in the rind-zone,all parallel.
Cortical layers: epicutis of the pileus in the marginalzone consisting of broom cells of
the Rotalis-type exclusively, these in hymeniform arrangement,main body 8-15 X 4.5-
10 p, with entirely thin wall or the wall at apex thickened, with a brown, intraparietal
pigment, setulae thin and rod-shaped,projecting0.5-1.2 p, subhyalineas in the cheilo-
cystidia or concolorous with main body; covering of the stipe none seen (perhaps fug-
acious) but the outermost hyphae beset with numerous setulae like those of the epi-
cuticular broom cells; no broom cells proper seen.
On leaves and leaf petioles of a dicotyledonous tree in tropical forest, gregarious.
MATERIALSTUDIED. ECUADOR. Tungurahua,Rib Topo, at about 1450 m alt, 30 Apr
1973, SingerB 7208 (F).

Marasmiussect LeveilleaniSinger, Bull. Jard. Bot. Bruxelles 34: 362. 1964.

TYPESPECIES. Marasmiusleveilleanus(Berkeley) Patouillard.
Pileus medium sized to ratherlarge (9-39 mm broad), usually well pigmented,
with an epicutis of smooth, ventricose to clavate elenients, some of which show finger-
like, mostly apical outgrowths or low warty processes;spores medium sized; lamellae
free but not collariate;hyphae with clamp connections, stipe central, insititious on dead
vegetable matter; cystidia none on the sides of the lamellae.

24. Marasmiusleveilleanus(Berkeley) Patouillard,Bull. Soc. Mycol. France 33: 55.

1917. Fig 42.
HeliomlycesleveilleanusBerkeley,London Jour. Bot. 6: 490. 1847.
umbraculumBerkeley& Broome,Jour. Linn. Soc. 14: 36. 1873.
TYPE. Gardner,from Ceylon (k) (not studied).
Pileus a deep and bright chestnut-rustcolor, eventually slightly lighter colored,
glabrous,sulcate to sulcate plicate on the margin,more rugose to venose-reticulate-
venose in the central pottion of the pileus, convex or convex-campanulatethen appla-
nate around a distinct umbo but eventually depressedin the center and subumbonate,
7-39 mm broad. Lamellaewhite to pale cream with not discolorous edge, sometimes
86 FloraNeotropica

anastomosing,mostly intervenose when quite mature, subclose to (mostly) subdistant,

narrow then medium broad, with few lamellulaeand sometimes all entire, narrowly
adnexed to remote, free, not collariate. Stipe bister to chestnut, shining, hollow, gla-
brous, smooth, equal or subequal, insititious, 35-60 X 0.5-1.7 mm; often accompanied
by (but not rising from) black, long rhizomorphs. Context white in pileus, stipe and
core of rhizomorphs,unchanging,relatively thick in the pileus center, inodorous.
Spores 5.7-9.7 X 3-4.5 u, mostly about 7-8.7 X 3.3-3.8 p, ellipsoid, oblong to
ovoid-oblong or subamygdaliform,hyaline, smooth, inamyloid. Hymenium: basidia
19-28 X 6-9 p, 4-spored, rarely (3-)4-spored;basidioles fusoid; cystidia none; cheilo-
cystidia much like the epicuticularbroom cells but all except about 1%hyaline, with
thin to more rarely slightly thickened wall 0.3-0.8, rarely to 2 p thick. Hyphae of the
pileus-tramaparallel or subparallel,inamyloid or pseudoamyloid, with clamp connect-
ions, with thin to moderately thick wall (0.2-0.4 p) of variablediameter(some inflated
ones to 12 , broad but not dimorphic). Cortical layers: epicutis of the pileus consist-
ing of broom cells in hymeniform arrangementbut many cells smooth and entire, ven-
tricose to claviform, fewer cylindrical, often with eccentric bulge or central mucro,
rarely forked, with thin to moderately thick wall, with apical or subapicalfinger-like
appendageswhich are straightand erect to somewhat oblique, 2-10 X 1.7-3-(5)Pu,en-
tire cell hyaline to stramineousor tawny-succineous(in the upper portion or all over),
15-40 X 5-14 ,; covering layer of stipe not differentiated.
On wood (fallen pieces of wood, dead fallen branches)of dicotyledonous trees,
often on Leguminoseae. Pantropicalbut rare in the neotropics.
MATERIALSTUDIED. MEXICO. Veracruz,3 km S of Montepio, 22 Jun 1969, SingerM1
8137 (F). ZAIRE. Yangambi,J. Louis 11010 (BR).
The type was studied by Patouillardand is obviously identical with the collections
described above.
ILLUSTRATION. Singer (1946b), fig 15, A-C.

Marasmiussect Scotophysini Singer, Sydowia 18: 342. 1965.

TYPESPECIES. Marasmiusscotophysinus Singer.
Pileus small, well pigmented, with hymeniform epicutis consisting of smooth, glo-
bose elements which are pedicellate; spores oblong to subcylindric,medium-sized;lamel-
lae adnexed to adnate, not collariate, minutely white fringed from corncob-shapedcheil-
ocystidia; hyphae with clamp connections, inamyloid; stipe central, attached to leaves
by an insititious base; cystidia on the sides of lamellae, none; stipe without conspicuous

25. Marasmiusscotophysinus Singer, Sydowia 18: 342. 1965.

TYPE.Singer B 1444, from Bolivia.
Pileus blackish livid purple (8-E-3) striate on sordid pallid to purple ground, sul-
cate to disc, glabrous,not unpolished, convex-applanate,3.5-5 mm broad. Lamellae
sordid pallid, tridymous, close, minutely white fringed at edges, rounded adnexed to
(mostly) adnate, almost narrow to moderately broad. Stipe black, shining, glabrous,
smooth or with slight minute, longitudinal furrows, insititious, 28-33 X 0.1-0.2 mm at
apex, graduallywidening below to 0.2-0.4 mm. Context inodorous.
Spores 7.5-9.7 X 2.5-3.7 A, oblong to subcylindric,thin-walled,hyaline, smooth,
inamyloid. Hymenium: basidia 21 X 5 p, 4-spored;cystidia none; cheilocystidia 15-30
X 9-13 p, consisting of a long, 3.5 p thick pedicel and an upper half which is corncob-
like or cauliflower-likediverticulatein its upper portion, diverticulashort and dense,
Marasmius 87

obtuse, about 1.5 X 1.5 p, both main body and diverticulahyaline, the upper portion
often forked. Hyphae: hymenophoral trama regular,consisting of filamentous(1.3-2,u
diam.) hyphae which are not gelatinized, inamyloid, those of the pileus radiallyarranged,
otherwise similar,also inamyloid but having pseudoamyloidcontents at times, opaque
and long, often irregularlybranched;all hyphae with clamp connections, those of the
stipe black incrusted, also inamyloid. Cortical layer: epicutis of pileus consisting of
globose, pedicellate cells which are either hyaline or with a porphyry sap, aside from
that, wall colored deep sepia by an intraparietalpigment, globose part (without ped-
icel) 6-14 I in diameter,pedicel up to 13 , long, these elements in hymeniform arrange-
ment with the contents partly pseudoamyloid;hypodermiumconsisting of filamentous
hyphae with membranapigment.
On dead dicotyledonous leaf in cloud forest (2200 m).
MATERIAL STUDIED.BOLIVIA. La Paz: Nor-Yungas, Cerro Uchumachi, 25 Feb 1956,
Singer B 1444 (LIL), type.
This species differs from the Hygrometriciin the smooth epicuticularelements,
from the Epiphylli in the pigment and the absence of pleurocystidia,from Pararotulae
in inamyloid hyphae and in non-collariatelamellae, and finally from the genus Gloio-
cephala in the absence of long hairs or oleocystidia and from all these groups in the
characteristic,almost unique cheilocystidia. It seems thereforejustified to erect a
special section for this species even though the section has thus far only a single rep-
ILLUSTRATION.Singer (1965), pl 6, fig 11.

Marasmiussect Epiphylli Kthner, Le Botaniste 25: 60. 1933 (as Epiphylleae).

TYPE SPECIES.M. epiphyllus (Persoon ex Fries) Fries.
Pileus small, white or whitish (or with pale buff disc) with hymeniform epicutis
consisting of smooth, globose of clavate, more rarely diverticulateelements and/or de-
nnatocystidia (in the latter case dermatocystidianeither pseudoamyloid nor capitate)
without long, bristle-likehairs; spores narrowlycylindric to oblong or ellipsoid, small
to large;lamellae subfree to subdecurrent,not collariate, sometimes absent; hyphae
with or without clamp connections, inamyloid or pseudoamyloid(but if hyphae
pseudoamyloid,epicuticularhyphae always smooth); stipe central or eccentric, attach-
ed to leaves, more rarely to wood by an insititious base, cystidia on the sides of the
lamellae present but sometimes very scattered and inconspicuous(because similarto
fusoid basidioles), mostly fusoid to ampullaceous,rarelyvesiculose, generallythin-
walled and hyaline, but may have pigmented resinous incrustation;stipe with conspic-
uous hairs or dermatocystidia.

Marasmiussect Epiphylli subsect Epiphyllini Singer, Sydowia 18: 134. 1965.

TYPESPECIES. A1.epiphyllus (Persoon ex Fries) Fries.
Tramalhyphae at least in the apex of the stipe inamyloid, epicuticularcells
weakly diverticulateor smooth; hymenophore veined radially,merulioid or lamellate,
sometimes absent.

Key to the Species of SubsectionEpiphyllini

1. Epicuticular cells all smooth; hyphae with or without clamp connections.
2. Hyphae with clamp connections.
3. Hymenophoral configuration merulioid or lamellate; odor often pleasant; cystidia
88 Flora Neotropica

4. Hymenophore merulioid; spores 3.6-4.3 X 2-2.2 p. 26. M. euosmus.

4. Hymenophore lamellate; spores 5.5-7 X 2-3 p (see M. subagricola; M. epiphyllus
if spores still larger5).
3. Hymenophore none; hymenial surface smooth. 27. M. nmartini.
2. Hyphae without clamp connections. 28. M, carpenterianus.
1. Epicuticular cells in their majority weakly diverticulated; hyphae with clamp con-
nections. 29. M. munyozii.

26. Marasmiuseuosmus Singer, Sydowia 12: 69. 1958.

TYPE. Martin & Welden 8702, from Panama.
Pileus white, glabrous,convex, then irregularlyflattened, membranous,3-8.5 mm
broad. Lamellae white, narrow,with obtuse edges, almost venose, with anastomosing
cross-veinsas high as the through-lamellaeforming a poroid-merulioidhymenophore,
adnexed. Spore print white. Stipe white, long-filiform,pruinate, equal to slightly
taperingupwards,insititious, 8-13 X 0.3-0.5 mnm.Context white, thin, odor of flowers,
Spores 6.6-4.3 X 2-2.2 u, hyaline, cylindrical, smooth. Hymenium: cystidia vesi-
culose or ventricose or clavate, hyaline, not refringent. Hyphae: hymenophoraltrama
regular,hyaline, all hyphae hyaline with clamp connections, inamyloid. Cortical layers:
epicutis of pileus cellular, consisting of smooth, hyaline, subisodiametricor isodiametric
cells; covering layer of stipe with numerous to crowded dermatocystidia,these thin-
walled or almost so, usually clavate but also capitate or ventricose-ampullaceous,erect,
28-40 X 8.7-11.5 u.
On a log, gregarious.
MATERIAL STUDIED. PANAMA. Canal Zone, Barro Colorado Island, 24 Aug 1952, Mar-
tinl & Welden 8702 (IA), type.

27. Marasmiusmartini Singer, Sydowia 12: 70. 1958.

Cyrnatella longipes Martin, Lloydia 7: 79. 1944.
Pileus white, dried yellow-brown,glabrous, smooth, cupulate to broadly funnel-
shaped, 1-2 mm broad. Lamellaenone; hymenial surface inferior and smooth, white.
Stipe white at the apex, abruptly yellow-brown below and shading to nearly black at
the base, glabrous, filamentous, with a socle-like base which is insititious, up to 160
X 0.2 mm. Context said to be subgelatinous(macroscopically,fresh), almost cart-
ilaginous when dried, very thin, white in the pileus and the upper part of the stipe;
no odor recorded.
Spores 9-11 X 4p , ellipsoid to oblong, inner side often applanate, smooth, hy-
aline, thin-walled,inamyloid. Hymenium: basidia 30-35 X 7 , 4-spored;cystidia 30-
40 X ? 9 ,, ventricose-cylindricalor apiculate, hyaline, little or not projecting, not
conspicuous, smooth. Hyphae filamentous, hyaline, inamyloid, with clamp connect-
ions. Cortical layer: epicutis of the pileus consisting of a layer of vesiculose, hyaline
smooth cells, forming a hymeniform structure.
On the bark of a fallen branch.
MATERIAL STUDIED. COLOMBIA. Sierra Nevada de Santa Marta, 1250-1500 m alt, 17
Dec 1935, Martin 3469 (IA), type.
ILLUSTRATION.Martin (1944), fig 11.

28. MarasmiuscarpenterianusSinger, Sydowia 12: 70. 1958.

TYPE. Carpenter 136, from Peru.
Neither M. subagricola nor M. epiphyllus have been found in the neotropics but do occur in northern
Florida. As for M. subagricola see also p 269.
Marasmius 89

Pileus whitish, smooth, convex, then convex-applanate,obtuse, not umbilicate,

2-6 mm broad. Lamellaewhite, ratherdistant (10-12 through lamellae, of those some
simple, the others simply or doubly forked), with some lamellulae,rather narrowbut
not venose except in immaturestate, broadly adnate. Stipe brown, at first with white
apex, setose, insititious under binocularlens pruinose, increasinglypruinose-pubescent
toward base, longer than the diameter of the pileus, 0.1-0.2 mm broad. Context white,
thin, inodorous.
Spores 6-7.7 X 2-4 p, ellipsoid with suprahilarapplanationor a small and weak
depression,hyaline, smooth, inamyloid. Hymenium: basidia 17-20 X 5.5-6.5 u, (2-)
4-spored;cystidia 25-37 X 6-10.5 1, ampullaceouswith the widest diameter in the
middle, hyaline, smooth, thin-walled,rather numerous. Hyphae: subhymeniumappear-
ing subcellular. Trama of pileus of hyphae which are parallelwith each other, slightly
thick-walled,not metachromaticin cresyl blue mounts, inamyloid, without clamp
connections. Cortical layers: epicitis of pileus consisting of subisodiametricor isodia-
metric cells of various diameters,e g 1 1.5-16 X 6.5-11.5 u, smooth, thin-walled;surface
of stipe with dermatocystidiawhich are hyaline, or more often melleous, with moder-
ately thin walls, very versiform,usually rounded above, ampullaceousor subulate or
cylindrical,etc, sometimes with a septum in the lower portion (5-)20-30 X (3.5-)7-10.
On fallen green leaves of a dicotyledonous plant in tropical forest.
MATERIAL STUDIED. PERU. Huanaco:TingoMaria,Cuevade los Pavos700 m alt, J. B.
Carpelter136 (LIL),type. MEXICO.Veracruz: Estaci6nBiologicade lasTuxtlas,21 Jun1969,
Singer M 8115 (F).
ILLUSTRATION. Singer (1958a), fig 2.

29. Marasmiusmunyozii Singer, Beih. Nova Hedwigia29: 100. 1969.

TYPE. Singer M 6517, from Chile.
Pileus white, with a flush or very pale cinnamon-ochraceousin the center, short
and slightly striate at the margin,glabrous,convex, 5-5.5 mm broad. Lamellaewhite,
distant (7-13 through-lamellae,some lamellulae)not anastomosing,narrow,later
medium broad, horizontal, finely flocculose at edge when seen under a lens, adnexed.
Stipe white, towards base chestnut, pruinose or pubescent, filamentous-equal,more
finely pubescent at base but insititious, 10-11 X 0.3-0.4 mm. Context white, pale
brownish in the lower part of the stipe, very thin, inodorous.
Spored 6-7.8 X 2.2-3 A, fusoid, hyaline, inamyloid. Hymenium: basidia 15-21 X
6.5 p, (2-)4-spored;cystidia inconspicuous,scarce;cheilocystidia making the gill edge
heteromorphouslike the epicuticularcells. Hyphae filamentous, with clamp connect-
ions, inamyloid, not gelatinized. Cortical layers: epicutis of the pileus hymeniform or
subhymeniform,consisting of elements which are 15-25 X 12.55-15 p,, vesiculose, in
their upper portion mostly beset with setulose appendages,these sparse or ratherdense,
short cylindrical, up to 2.5 p projecting,divergent,the whole cell hyaline and thin-walled,
often pedicellate; covering of the stipe consisting of dermatocystidia,these up to 165 ,
long, in their lower portion up to 8.5 broad, towards the apex long effilate (1.5 , dia-
meter) and/or in the lower portion lobed-appendiculatewith thin to moderately thick
wall, hyaline or subhyaline.
On grass leaves in the coastal fog forest.
MATERIAL STUDIED. CHILE. Coquimbo: Fray Jorge, 16 Mar 1967, Singer M 6517
(SGO), type.
This species differs from another species known to have lacerate or appendiculate
epicuticularelements, the EuropeanMarasamiusepiphylloides Rea, by its much smaller
90 Flora Neotropica

spores and habitat on grassesratherthan Hedera. It is a somewhat marginalspecies in

a vegetation of predominantlysouthern origin, but comes from the subtropicalzone of
North Chile.

Marasmiussect Epiphylli subsect Eufoliatini Singer, subsect nov

TYPESPECIES. M. eufoliatus Klhner.
Hyphis partim manifeste pseudoamyloideis,fibulatis aut defibulatis.
Hyphae the carpophoreor at least the stipe distinctly pseudoamyloid;hyphae
with or without (in the neotropics without) clamps.

Key to the Species of SubsectionEufoliatini

1. Epicutis made up of cystidioid elements; spores 3.8-4.5p broad, ellipsoid to ellipsoid
oblong. 30. M. sanctixaverii.
1. Epicutis made up of claviformcells 12-23 X 7-14p, no dermatocystidiapresent;spores
2.5-3.2u broad,long-fusoid. 31. M. caliensis.

30. MarasmiussanctixaveriiSinger, Sydowia 18: 342. 1965.

TYPE. Singer T 3156, from Argentina.
Pileus white, glabrous,smooth, convex with depressedcenter, 7-8 mm broad.
Lamellaewhite, distant, about 12 through-lamellae,not very broad, at maturity becom-
ing distinctly forked-anastomosingintervenose, even young ones forked but then not
intervenose, adnate-subdecurrent.Stipe white at apex, chestnut brown below, macrosco-
pically subglabrousbut under a lens distinctly pubescent, equal, insititious, 15-17 X 0.3-
0.4 mm. Context inodorous.
Spores 8.2-9.7 X 3.8-4.5 u, ellipsoid to (mostly) ellipsoid-oblong,without sup-
rahilardepressionor applanation,hyaline, smooth, inamyloid. Hymenium: basidia 22 X
6.2 ,, 4-spored, hyaline, clavate;cystidia on edge and sides moderately numerous, 27-43
X 8-10 , often incrusted at the tip, thin-walled,hyaline, fusiform. Hyphae of hymeno-
phoral trama and pileus rather irregularlyarrangedalthough with axial respectively
radialtrend, filamentous but not parallelwith each other, those of subhymeniumnot
to weakly pseudoamyloid, those of pileus, stipe and hymenophoraltrama distinctly
pseudoamyloid;all hyphae without clamp connections. Cortical layers: epicutis of
pileus consisting exclusively of dermatocystidiaforminga hymeniform layer, many of
the dermatocystidiaalmost globose below but only seemingly cellular since even these
are mucronate and cystidioid, all hyaline, thin-walled,crowded, inamyloid, versiform,
varying from ventricose-clavateto ventricose-fusoidampullaceousor constricted, most
frequently pear-shaped,the largermore cystidioid ones 23-32 X 6.8-10.5 ,, the smaller
ones e g 20 X 6.8 , but with all sorts of transititonsand the larger ones not distinctly
or consistently projecting;Hypodermiumhyphous, scarcely pseudoamyloid;surface of
stipe beset with numerous dermatocystidiawhich form its pubescence, even more vari-
able than the elements of the epicutis of the pileus, of three main types (1) ventricose-
ampullaceous,e g 28 X 9 ,, mostly curved at base, hyaline, extremely numerous(2)
ascendinginto a subglobose lead (3) filiform to cylindrical,hyaline, 27-54 X 4.7 u.
On dead, fallen leaves (probably of Allophylus edulis) rolled around the pupa
of a lepidopterousinsect, gregarious.
MATERIALSTUDIED. ARGENTINA. Tucuman:CiudadUniversitariaat 1100 malt,
Singer T 3156 (LIL) type.
ILLUSTRATION.Singer (1965), pi 6, fig 12.
Marasmius 91

31. Marasmiuscaliensis Singer, sp nov Fig 43.

TYPE.Singer B 6149, from Colombia.
Pileo albo, 2.2-7 mm lato; lamellis albis, distantibus, demum intervenosis;stipite
dilute cinnarmomeo-corricolore vel castanescentein parte inferiore, albo vel albido ad
apicem; odore nullo. Sporis 6.5-10.5 X 2.5-3.5 j,, fusiformibus;hyphis stipitis defi-
bulatis, pseudoamyloideis;epicute hymeniformi ex elementis 12-27 X 7-14 p clavatis,
vesiculosis, ventricoso-subelongatisefformata, dermatocystidiisnullis. Ad folia dico-
tyledonea in Columbiaet Venezuela. Typus in F conservatusest.
Pileus white, smooth, later rugose or short-sulcate, glabrous,convex, 2.2-7 mm
broad. Lamellaewhite, distant (up to 12 through-lamellaewhich are sometimes forked,
few lamellulae) in age distinctly intervenose, narrowbut not veniform, or only a few
so, adnexed, adnate or subdecurrent. Stipe white or whitish above, light cinnamon
tawny to chestnut below, macroscopicallysubglabrousbut pruinose-pubescentunder
a lens, curving near apex if growing on the undersideof the leaves, insititious, equal,
3-12 X 0.1-0.3 mm. Context white, thin, inodorous.
Spores (6-)6.5-10.5 X 2.5-3.5 u, fusoid to long-fusoid, smooth, hyaline, inamyloid.
Hymenium: Basidia 16-22 X 5-7.5 ,, 4-spored;basidioles fusoid; cystidia more numer-
ous on edges than on sides of lamellae, 22-37 X 4-9 p, ampullaceousor fusoid with
ratherthin and short neck, rarely subcylindric,rarely the neck widened in the middle,
hyaline, sometimes slightly incrusted(hyaline), at edge sometimes partly replacedby
cells similarto those of the epicutis, especially near the marginof the pileus, all
cystidia thin-walled,inamyloid. Hyphae of the pileus-tramafilamentous, 2-6 pbroad,
thin-walled,weakly pseudoamyloidor inamyloid, without clamp connections; hyphae
of the stipe parallelwith each other, hyaline at the apex and strongly pseudoamyloid,
filamentous, multiseptate, without clamp connections, 2.5-5 p broad. Corticallayers;
epicutis of the pileus hymeniform, its elements 12-27 X 7-14,, clavate, vesiculose or
ventricose-subelongated,hyaline, smooth, sometimes a few mucronate, rarely bi-lobed
at the apex, thin- to firm-walled,no dermatocystidiadifferentiated;covering of the
stipe consisting of dermatocystidiaand short, hair-likeexcrescenceswhich are 3-30
X 3-9 p, versiform,cylindric, conical, filiform, hemisphericalor ventricose, sometimes
bi-lobed but smooth, not appendiculate,thin-walled(wall 0.1-0.3 p) and hyaline, en-
tire, with rounded tip, numerous.
On dicotyledonous leaves, singly but gregarious,in tropical forest.
MATERIAL STUDIED.VENEZUELA. Dto Federal:7 km Southof Chichiriviche,
Jul 1972,Dumontet al VE6707 (NY);betweenColoniaTovarandEl Tigre,29 Jul 1972,Dumont
et al VE6555 (NY). COLOMBIA. Valle:Cali,Saladito,15 Apr1968,SingerB 6149 (F), type.

Marasmiussect Marasmius
RotulaeFries,Epicrisisp 384. 1838.
CollariatiBataille,FloreMonographiqueMarasmiusd'Europep 26. 1919.
TYPESPECIES. M. rotula (Scopoli ex Fries) Fries.
Pileus generally small, white or pigmented and often with a knob or papilla in
the middle which in mature specimensis often located in an umbilicus and surrounded
by a pallid ring zone, with hymeniform epicutis which eventually may become slightly
disorganized,consisting of broom cells which at their apex or the entire upper portion
are covered by minute to ratherlong erect or divergentsetulae without long conspic-
uous hairs;spores small to large, ellipsoid to oblong, mostly (or always) acyanophilic;
lamellae subcollariateto mostly distinctly collariate;hyphae with clamp connections,
mostly pseudoamyloid at least at the apex of the stipe, more rarely all inamyloid;
stipe centrally attached to leaves or wood, sometimes to black rhizomorphswhich are
92 FloraNeotropica

attached, on leaves or wood, always distinctly insititious; cystidia none, rarely present
but inconspicuous;stipe and rhizomorphsmostly glabrous,in a few species pilose.

Marasmiussect Marasmiussubsect Pararotulae(Singer) Singer, stat nov

Marasmiussect PararotulaeSinger,Sydowia 18: 336. 1965.
TYPE SPECIES. Marasmius pararotula Singer.
Epicuticular broom cells of the Rotalis-type (with divergent setulae)

Key to the Species of Subsection Pararotulae

1. Pileus in mature specimensessentially white or whitish (excepting the centralknob or
papilla which may be white or pigmented)on drying sometimes becoming very pale
argillaceousor gilvous.
2. All or the greatmajorityof the stipesattachednot to the substratumbut to conspicu-
ous blackrhizomorphs.
3. Lamellae9-16, mostly 12-14; pileus 6 mm broad;Gulf area. 32. M. multiceps.
3. Lamellae7-9; pileus 0.5-2 mm broad:Amazonasregion. 33. M. cupressiformis.
2. All or the great majorityof the stipesattachedto the substratumand accompaniedor
not by blackrhizomorphs.
4. Centraldot absent or light colored or brownishand soon disappearing; sporesup
to 8.5, long.
5. Spores small, 5-7 X 2.2-3.5 ; centralknob or dot gilvousor cork-coloror pale
flesh; lamellae13-17;pileus 10 mm broador broader;on leaves. 34. M. rotuloides.
5. Spores as small as above or larger;centraldot, knob or umbo cork-color,or
white; lamellae as above or fewer or more numerous;pileus less than 9 mm
broad;on wood or on leaves.
6. On wood; centralumto white. 35. Al.panamenisis.
6. On leaves.
7. Sporesreachingmore than 4.8p broad;pileus 1-1.8 mm broad;lamellae
5-8. 36. MI.pararotula.
7. Sporesup to 4.8 Mbroad;lamellae9-17.
8. Speciesof the Amazonregion, Colombiaor Venezuela;darkcentral
dot absentin freshpileus.
9. Region of the central dot of the pileus not elevated but flat:
spores5.5-8.5 X 2.5-4 p lamellae11-17; tropical-montanespecies.
37. M. vergeliensis.
9. Region of the centraldot somewhatto stronglyelevated,show-
ing at least in the dry stage a centralknob or a small umbo or
papilla in the umbilicusor in the smooth central apex of the
pileus; spores as above or different; lamellaeas above or more
numerousor fewer.
10. Spores 6.8-10 X 2.8-4.8p; pileus 1-5 mm broad;umbo sur-
rounded by a concolorous zone which may appear paler
because of finely cracked(undera lens) surfacewhen dried.
11. Lamellae9-12. 38. M. apateliuls.
11. Lamellae13-17. 39. M. rotalis.
10. Spores 5-6 X 2.5u; pileus 5 mm broad or rarely broader;
umbo surroundedby a conspicuouswhite zone (dried).
40. M. castellanoi.
8. Species of the Southern Hemisphere.Yungas to Selva Boliviano-
Tucumana(see "12" below).
4. Centraldot either fuliginous, deep ferruginous,deep brown or dark gray or, if
paler,sporeslargerthan 10 long.
12. Spores14.5-16.5 X 3.7-4.3p. 41. M. pandoanus
12. Sporessmaller.
13. Spores10-14 X 3-4 ; lamellae10-16. 42. M. oaxacanus.
13. Sporessmalleror broaderor lamellaefewer.
14. Lamellae6-9; spores7.5-11 X 4.5 p. 43. M. manuripiensis.
14. Lamellaemore numerousor spores either largeror smallerbut
alwaysnarroweror broaderthan indicatedabove.
15. Pileus pale flesh pink; spores 8.5-9 X 3.5-4 p (see "24"
Marasmius 93

15. Pileus white or whitish when fresh or mostly so, but never
flesh pink ever;sporesdifferent.
16. Pileuswith a ratherpale or pallidumbo;spores12.5-14.2
X 7.5-9,u;lamellaeabout seven. 44. M. psychotriophilus.
16. Pileus with a dark umbo; spores 5.5-7.8 X 2-3(-4)u;
lamellae14-20 (if fewer compare"6" above).
17. Pileusabout 6 mm broad;lamellaeabout 20; on oak
leaves. 45. M. cundinamarcae.
17. Pileus up to 5.5 mm broad;lamellae11-17; on dead
leavesother than oak. SouthernSouth America(if in
NorthernNorth America.compare"11" above).
46. M. leucorotalis.
I. Pileus in mature specimensnot white in the largermarginalregion (if only old, dried
pale avellaneous,pale ocher or pale argillaceouspilei seen, compare also "2" above).
18. Pileus sepia or mummybrown, 2-3 mm broad;lamellaeon edges discolorous(con-
colorouswith the pileus);spores11-14 X 3.5-5 ; on bark. 47. M. arimanus.
18. Not combiningthese characters.
19. Pileusgray to beige (betweenblackishgrayto pale cinereouswhen fresh).
20. Sporeslargerthan 9,ulongand 4.5g broad,(see also subsectionPenicillati).
48. M. griseofuscescens.
20. Sporessmaller.
21. Lamellaewith cinereousto darkgray edge; spores7-9 X 3.5-4.7p; cheilo-
cystidiawith mostly sepiacoloredsetulaeand hyalinemainbody. Mexico.
49. M. scototephrodes.
21. Lamellae with white edge which may become brownish after drying;
spores 5.5-8 X 2.5-4ui;cheilocystidiaeither entirely hyaline or if setulae
arebrown, mainbody with hyalinewall but fuscous cell sap.Colombia.
50. M. leucozonitiformis.
19. Pileusnot so colored.
22. Pileus when fresh purplishred or red or pale flesh pink with dark colored
centraldot on papilla;lamellaeeitherwith discolorousedge and collariumor
evenly flesh-pinkall over.
23. Spores 8-11 X 4.5-5u;edge of lamellaeand collariumred;stipe 15-35 mm
long. 54. M. carminis.
23. Sporesnarrower;edge of lamellaenot discolorous;stipe sometimeslonger.
24. Pileuspale flesh pink;lamellaeconcolorouswith pileus. 53. M. carneotinctus.
24. Pileusdifferently colored; lamellaewhite with white edges but collar-
ium + discolorous(see "25" below).
22. Pileus neither red nor pale flesh pink but either brown or ferruginous or
orange or violet-lilac (if gilvous or pale argillaceous, see "2" above).
25. Spores over 10p long and up to 3.5g broad; pileus white when young but
soon becoming brown with a paler zone around the central dark dot and
on the margin. 51. 1M.tanyspermus.
25. Spores less narrow.
26. Pileus violet-lilac; lamellae ? 12; spores 6.2-9.5 X 2.5-3w; on dicotyle-
donous leaves in tropical-montane forest. 52. M. violeorotalis.
26. Pileus differently colored. Key II.

Key II
1. Spores voluminous: 8.5-13 X (5-)5.7-9(-10.3),(; pileus orange ferruginous, small, with a
pale, avellaneous-cinnamon zone around a central, black dot, conical-truncate when
fresh, barrel-shaped when dried; collarium ochraceous. Cloud forest species. 55. M. buzae.
1. Spores less voluminous or pileus and collarium different.
2. On dicotyledonous wood (if on woody stems of vines compare M. variabiliceps var
separatus) in tropical forest and montane woods.
3. Stipe rising from a common rhizomorphic strand, yellowish; pileus brown, less
than 1 mm broad (if broader and stipe black, see subsect Penicillati); spores not
reaching 4 p broad. 56. M. populiformis.
3. Stipe often accompanied by rhizomorphs; although not attached to them but
insititious on the substratum.
4. On slender twigs, thin vines(or occasionally on small pieces of Aextoxicum).
5. Pileus Brussels Brown (R) 1.5-2 mm broad; spores 6-7 X 2-2.6p; lamellae
8-10. 57. M. minimus.
94 FloraNeotropica

5. Pileus not Brussels brown, larger; spores broader; lamellac more numerous
(see "9" below).
4. On rotten logs and their fragments, on fallen branches or twigs, never on
6. Rain forest species with setulae of the epicuticular broom cells not reaching
1.5 p in length; lamellae about 12. 58. M. dodecaphyllus.
6. Growing in subtropical or tropical-montane forests; many setulae of the
epicuticular broom cells longer than 1.5 p; lamellae often more than 12.
7. Pileus beige-brownish; spores 4.3-5.7,/ broad. Southern South America.
59. M. platysperinus.
7. Pileus deep ferruginous to cocoa color; spores narrower. Northern and
Equatorial South America.
8. Spores 6.2-7.5 X 3.2-3.7p; cheilocystidia mostly setulose. 60. M. nebularum.
8. Spores shorter; cheilocystidia mostly smooth. 61. M. aequatorialis.
2. On dead fallen leaves and their petioles, rarely on vines or thin branches.
9. In the coastal cloud forests of Northern Chile, also in South Chile and Juan
Fernandez, on Nothomyrcia and other Myrtaceae and on Aextoxicum.
62. A1.nothomivrciae.
9. Neither on Myrtaceae nor on Aextoxicaceae.
10. Spores 5.5-7.5 X 2-3.5 u; pileus with a pallid or white zone around the central
dark do or papilla, >1.5 mm broad.
11. Pileus (fresh and dried) brown "sorrel" to "Mohawk M&P," 3-5mm
broad; lamellae 15-17; spores up to and mostly 7.5 X 3.5u. Rainforest of
the plains (Amazon). 63. M. h!laeicola.
11. Pileus ochraceous, beige or deep orange-ferrugineous or light brown;
lamellae often fewer or more numerous.
12. Pileus clay color or light brown, 1.5-5 mm broad; lamellae 10-16,
edges eventually concolorous with the pileus, collarium pallid (com-
pare M. rotuloides and related species, also - Key I, "5"). Tropical
montane and subtropical zone. 46. M. letlcorotalis var leucozonites.
12. Pileus deep orange-ferrugineous (see "19" below).
10. Spores generally somewhat larger; pileus with or without a white or pallid
zone around the central dot, of variable size.
13. Lamellae (17-)20(-22); collarium often brown; pileus broader than
14. Pileus brown both fresh and dried, not ferrugineous nor chestnut
to mahogany on drying or dried; stipe up to 56 mm long.
15. Spores 7.2-10 X 4.5-5,. 64. M. vigintifolius
15. Spores 6.5-8 X 2.8-3.8. 65. M. idroboi.
14. Pileus russet brown when fresh but becoming deep ferruginous
brown, chestnut or mahogany when dried; stipe 58-100 mm long
(see "18" below).
13. Lamellae fewer (up to 17) or collarium not brown.
16. Stipe rising from a common rhizomorphic strand.
17. Spores 7.5-10.3 X 4.3-6p. 66. M. eucladopus.
17. Spores 10-11.7 X 4-4.2u (see "19" below).
16. Stipe rising from the substratum.
18. Lamellae about nine, not discolorous at edges or collarium;
pileus 1.5-2 mm broad. 67. AM.tetrachrous.
18. Lamellae more numerous or pileus <1.5 mm broad.
19. Lamellae about 12, extraordinarily broad; edges and collar-
ium concolorous with the sides of the lamellae; hyphae of
pileus not or scarcely pseudoamyloid; spores 8.2-11.2 X
3.5-4p1;pileus 2-2.5 X 2 mm with vertical sides. Rain forest
species. 68. M. tereticeps.
19. Lamellae about 9-14 and the collarium and often also edges
of lamellae discolorous, ochraceous, orange or brown or else
lamellae more numerous (16-21); hyphae of pileus weakly
to distinctly pseudoamyloid; spores and pileus as above or
20. Pileus 1.5 mm broad or broader. 69. Al. variabiliceps.
[Cf. also M. griseofulscescenss!
20. Pileus <1 mm high and broad; lamellae 9-11; spores 6-8
X 3.5-41i. Lowland rain forest species. 70. M. baeocephalus.
Marasmius 95

32. Marasmiusmulticeps Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 298. 1869.
muLlticeps(Berkeley& Curtis)Murrill,North AmericanFlora 9: 286. 1915.
Polymarasmiussubnulticeps Murrill,North AmericanFlora 9: 286. 1915.
Marasmiussubimulticeps(Murrill)Saccardo& Trotter, Sylloge Fungorum23: 155. 1925.
TYPE. Wright160 (K), from Cuba.
Pileus white but in age and on drying sometimes tending to become brownish ful-
vous, with chestnut-fuliginousto black umbo both fresh and dried, sulcate, but the
umbo and papilla smooth, hemisphericor convex, umbilicate, mostly with a distinct
papilla or umbo in the umbilicus or at least a somewhat elevated, dark dot present,
1.5-6 mm broad. Lamellaewhite, distant, with (9-)11-14(-16) through-lamellae,all
entire or 1-3 lamellulaepresent, ratherbroad, shallowly but distinctly collariate. Stipe
black (at first with pallid or brown apex), glabrous,insititious with a ring-jointand
risingfrom black rhizomorphs,equal or taperingupwards, 5-25 X 0.2-0.5 mm . Black
rhizomorphsglabrous,creepingor arbuscular-ascendant, their main stem 0.6 mm thick.
Context of the pileus white, thin.
Spores 6.2-8.5 X 2.2-3.7,, ellipsoid to ellipsoid-oblong,smooth, hyaline, inamy-
loid. Hymenium: basidia4-spored, basidioles fusoid, some remainingsterile (cystidioles);
cystidia none; cheilocystidia with main body hyaline, 12-13 X 6.8-7 p, beset with div-
ergent setulae 4-5 X 0.8-1 ,u,also hyaline, attenuated to a subacute tip. Hyphae hyaline
in the pileus-trama,chestnut-fuliginousin the stipe, there paralleland thin-to thick-walled,
many distinctly pseudoamyloid but in the pileus-tramaand the hymenophoraltrama
(which is regular)mostly thin-walledand only slowly and vaguely pseudoamyloid,those
of the rind of stipe and rhizomorphsstrongly forking-interlaced,those of the core of
the stipe and rhizomorphsoften granular-incrusted by a hyaline incrustation,all with
clamp connections and filamentous but some swollen to 10 p. Cortical layers: epicutis
of the pileus hymeniform or subhymeniform,consisting of broom cells 12-17 X 9-12.5
It, of the Rotalis-type, hyaline, with numeroussetulae which are either rod-shapedor
conical or even rounded-verruculose,0.8-4 X 0.6-1.9 u, the main body often laterally
extended or else vesiculose, inamyloid.
On dead logs and trunks. Cuba, Belize, Guatemala.
MATERIALSTUDIED. CUBA. Wright160 (FH, K), type; Fungi CubensesWrightiani132.
(FH). GUATEMALA. PlateauaboveChiatla,18 Nov 1959,Degener,comm.Petrak,II (BAFC).
Dennis (1951c) data on the type of M. submulticepsshow that it is conspecific
with A. multiceps.
ILLUSTRATIONS. Dennis (1951 b) fig 11; (1951 c) fig 28.

33. MarasmiuscupressiformisBerkeley, Hook. Jour. Bot. 8: 140. 1856.

TYPE.Spruce 75, from Brazil.
Pileus white, whitish, beige to brownish in the herbarium,with a narrow. fuli-
ginous-chestnutcolored center, glabrous,sulcate, convex, at first conical then convex
with a prominent papilla in the umbilicus, 0.5-2 mm broad. Lamellae white, browning
in the herbarium,distant (7-9 through-lamellae,all equal), broad, collariate. Stipe
chestnut-black,glabrous,smooth, setiform, equal or taperingupwards,insititious on
rhizomorphs,1.2-4 X 0.1-0.4 mm. Rhizomorphsblack, long, often ramifiedand as-
cendant, often thicker than the stipes, smooth and glabrous. Context very thin, white
in pileus.
Spores (5.8-)6.7-8.5 X (3-)3.5-4.5(-5.5) p, ellipsoid, smooth, hyaline, inamyloid.
Hymenium: basidia 13.5-14 X 5-5.5 yu;cystidia none; cheilocystidia similarto the broom
cells of the epicutis. Hyphae of the pileus tramahyaline, not gelatinized, thin-walled,
96 Flora Neotropica

not or vaguely pseudoamyloid;those of the stipe and the region above its insertion
distinctly pseudoamyloid,here parallel, all hyphae with clamp connections. Cortical
layer: epicutis of the pileus nearly hymeniform, consisting of two types of epicuti-
cular broom cells, (1) of the Rotalis-type, with the main body 11-14 X 7-11 U, cla-
vate to vesiculose, sometimes 2-3-forked, setulae 1-2 X 0.5-0.7 p, divergentor obliquely
erect in the subapicularregion, hyaline, (2) of the Siccus-type, 20-34 X 6-13 p,, the
latter type dense and numerous in the center, widely dispersedto rare in the margin
al region of the pileus, pigmented, rarely subhyaline, with walls 1-4.5 u thick, setulae
apical and erect or sometimes in form of side-branches,finger-likeor obtusely coni-
cal, 2-15 X 1.5-4,, some of the "broom cells" without appendagesor setulae,
Over dead wood and leafmold in tropical rain forest. Pantropical(?), at least
in the Amazonas and the Congo regions.
MATERIALSTUDIED. BRAZIL. Amazonas:"Panure"(=Sao Jer6nimo),Spruce 75 (K),
type. ZAIRE. J. Ghesquiere1355 (BR).
Singer (1964b), fig 15, D, E, F.; Berkeley (1856) pl 5. fig 3.

34. Marasmiusrotuloides Dennis, Trans. Brit. Mycol. Soc. 34: 415. 1951.
TYPE.Dennis 232, from Trinidad.
Pileus white or whitish in tile marginalzone, towards the center mostly flushed
flesh-ocher to yellow-beige or light ochraceous to pallid, with a white or paler ring
zone around the central dot which varies from almost flat to low-umbonate, dried
more beige to pale grayishbeige, the central dot or umbo concolorous with the zone
around the white zone, not chestnut either fresh or dried (the flesh-ocherzone fresh
about 12 C 6/7 or "yellow beige" M&P),long and deeply sulcate, glabrous,strongly
convex with an umbilicus and with or without a low umbo in the umbilicus, 9-17 mm
broad. Lamellaewhite, distant (13-16 through-lamellae,no lamellulae), broad, distinct-
ly collariate. Spore print white. Stipe dark red-brownto usually black, at first with
white apex, glabrous, smooth, insititious or widened into a very narrow, insititious
disc, equal, not attached to rhizomorphs,20-37 X 0.2-0.9 mm. Context in pileus
white, thin, inodorous.
Spores 5-7 X 2-3.5 ,, ellipsoid to fusoid-oblong, smooth, hyaline, inamyloid.
Hymenium: basidia 17-19 X 4-5,u, basidioles fusoid; cystidia none; cheilocystidia hy-
aline thin to firm-walled,otherwise like the epicuticularbroom-cellsin shape and size.
Hyphae with thin to often thick walls, filamentous, pseudoamyloid, with clamp con-
nections, hyaline in pileus-tramaand the regularhymenophoraltrama, not gelatinized.
Cortical layers: epicutis of pileus hymeniform, with broom-cellsof the Rotalis-type,
main body 12-27 X 8-26 ,, obpiriform to vesiculose, some laterally extended and even
hyphous at times, hyaline, inamyloid, with a wall thin- to 1 p thick, over most of the
surface beset with divergentsetulae which are rod-shaped,hyaline to light stramineous,
1-2 X 0.88.
On dead, fallen, dicotyledonous leaves in tropical forest.
MATERIALSTUDIED. TRINIDAD.Cerrode Aripo, 23 Oct 1949, Deinnis232 (K), type.
BRAZIL. Para,EstanciaPirelli,7 Jun 1966, SingerB 4231/I (BAFC), B 4224 (BAFC);Belem,
10 Jun 1966, SingerB 4252 (BAFC);B 4269 (BAFC).
Dennis (1951a) pl. 19, fig 14.

35. MarasmiuspanamensisSinger, sp nov

TYPE. Martin& Welden8733, from Panama.
Pileo albo; lamellis albis, distantibus,latissimis,collariatis;stipite nigro apice albo,
Marasmius 97

rhizomorphisnigris praesentibussed stipiti haud adhaesis. Sporis4.5-7.2 X 2.5-3.6M;

elementis epicuticularibuseis Marasmiirotalis analogis. Ad lignum in insula BarroCo-
lorado. Typus in F conservatusest.
Pileus white, glabrous,sulcate, barrel-shapedto convex, umbonate in an umbili-
cus, about 3 mm broad. Lamellaewhite, distant (15-17 through-lamellae,no lamell-
ulae), very broad, distinctly collariate. Stipe black with whitish apex, insititious, equal
about 15 X 0.2 mm, accompaniedby (but not rising from) black rhizomorphswhich
are thinner than the stipe and glabrous. Context of the pileus white, thin.
Spores 4.5-7.2 X 2.5-3.6 u , ellipsoid, smooth, hyaline, inamyloid. Hymenium:
basidia 4-spored;basidioles fusoid; cystidia none; cheilocystidia hyaline, like the epic-
uticular broom cells. Hyphae of the regulargill trama hyaline, inamyloid, with clamp
connections. Hyphae of the apex of the stipe parallel,strongly pseudoamyloid. Cor-
tical layers: epicutis of the pileus consisting of broom cells in hymeniform arrangement,
broom cells of the Rotalis-type, main body 15-23 X 11-16 1, vesiculose, hyaline, few
yellowish-fulvousat times, setulae all over the cell, divergent.
On wood, Panama.
MATERIALSTUDIED. PANAMA. BarroColoradoIsland, 15/16 Aug 1952, Martin&
lelden 8733 (F), type.

36. MarasmiuspararotulaSinger, Sydowia 18: 340. 1965.

TYPE. Singer B 1886, from Bolivia.
Pileus white, occasionally with a central or eccentric gray spot, subglabrousor
glabrescent,under a lens aspersedwith sugaryparticles which later disappear,campanu-
late, some convex, more or less distinctly umbilicate, especially when dried, sulcate
when dried, 1-1.8 mm broad. Lamellaewhite, the edges with sugary fringe, under a
lens appearingmore or less distinctly collariate, ratherbroad, all entire, distant to very
distant (five to eight), mostly five when young and seven when mature. Stipe white
at apex, deep umber toward base, glabrousand smooth, insititious without a basal
ring, solid, filamentous, 5-12 X 0.1 mm. Context almost none, hyaline, inodorous.
Spores 7.5-8.3 X 4.8-5.5,, ellipsoid, ratherbroad, smooth, hyaline, inamyloid.
Hymenium: basidia 2-4-spored, 16-17 X 7.3-7.7 u;cystidia none; cheilocystidia caus-
ing the sugaryappearanceof the edge of the lamellae, hyaline similarto the epicuti-
cular elements. Hyphae: subhymeniumof very short, small elements; hymenophoral
trama regular,not bilateral,hyphae of pileus trama ratherirregularin size and shape,
subhyaline to hyaline in ammonia, not voluminous as in Mycena but some occasional
cells ratherbroad and short although generallyfilamentous or subcylindric,interwoven,
hyphae of stipe with slightly thickened walls and strictly parallel,hyaline only in upper
portion of the stipe, filamentous;all hyphae pseudoamyloidand with clamp connect-
ions. Cortical layers: epicutis of pileus consisting of spherocysts which form patches
of hymeniform covering, besides some free cells present, all these elements 19-36 X
12-28 A, either smooth (rarely) or vaguely rugose or (the majority) with small setuloid
diverticula,these hyaline as the main body and irregularlydistributedover the upper
portion of the surface of the cell, ratherfew at times, 0.7-5 X 1-2 A;where the trama
or hypodermiumis exposed, a Rameales-structureof nodose, irregularelements can be
observed. No covering layer on the stipe.
On dead fallen leaves of dicotyledonous plants in tropical rain forest.
MATERIALSTUDIED. BOLIVIA. Beni: Vaca Diez, Guayaramerfn,3 km below the
villagein virginforest, 12 Feb 1956, SingerB 1886 (LIL), type.
ILLUSTRATION:Singer (1965) pl 7, fig 13.
98 Flora Neotropica

37. MarasmiusvergeliensisSinger, sp nov Fig 44V.

TYPE.Singer B 6426, from Colombia.
Pileo albo, 3-4 mm lato; lamellis albis, subdistantibusvel distantibus,collariatis;
stipite nigo apice albo; rhizomorphispraesentibussed stipiti haud adhaesis. Sporis 5.5
-8.5 X 2.5-4p; elementis epicuticularibuseis Marasmiirotalis analogis. Ad folia de-
lapsa dicotyledonea in Columbia. Typus in F conservatusest.
Pileus white, with a light brownish to white, smooth dot (not or scarcely ele-
vated) in the umbilicus and even if brownish, becoming pallescent in age and/or on
drying, leaving the whole pileus surface white, but dried in some caps very pale ocher
or champagnecolor over the marginalzone, the latter strongly sulcate, convex, with
a narrow,umbilicate depressionin the center, the depressionflat-bottomed and even-
tually often slightly tomentose under a lens, 3-4 mm broad and up to 3 mm high.
Lamellaewhite, subdistant to distant (11-17 through lamellae), all equal, broad,
collariate, surface inside collarium smooth, narrow. Stipe black with white apex,
sometimes a gray zone between the apex and the rest, drying to pale umber or smoke-
fuscous-bisteror remainingblack, shiny, setiform, glabrous,equal, insititious, accom-
panied by, but not connected with black rhizomorphs,11-22 X 0.1-0.5 mm. Context
white in the pileus, very thin, inodorous (perhaps the mycelium with a fishy odor.)
Spores 5.5-8.5 X 2.5-4 p, mostly 7-8 X 2.8-3.5 P, fusoid, smooth, hyaline, in-
amyloid. Hymenium: basidia 14-15 X 5.3 u, 4-spored:basidioles fusoid; cystidia none;
cheilocystidia numerous, like the epicuticularbroom cells. Hyphae of the hymenophoral
trama regularlyarranged,almost or quite inamyloid, those of the pileus-tramaweakly, of
the stipe-tramadistinctly pseudoarnlyoid,thin-walledin the pileus, with clamp connect-
ions. Cortical layer: epicutis of the pileus hymeniform, consisting of broom cells of the
Rotalis-type, main body 13-18.5 X 5.5-17 p, entirely hyaline, clavate to globose, thin-
walled, inamyloid, setulae hyaline or subhyaline, 1-2 X 0.4-0.8-(1) p.
On fallen dicotyledonous leaves in tropical-montaneforest. Colombia.
MATERIAL STUDIED.COLOMBIA. Valle,Mun.Dagua,Corr.Vergel,27 Apr1968,Singer
B 6426 (F), type; 14 Apr 1968, SingerB 6119 (F).
This species is closely related to MarasmiuscapillarisEllis from North America
which grows on oak and other leaves; common in the Midwest. The latter differs from
Mlarasmiusvergeliensisin the host, largerspores (7.5-10 X 3.8-4.3 p) and slightly larger

38. Marasmiusapatelius Singer, Bull. Jard. Bot. Bruxelles 24: 332. 1964. Fig 44A.
TYPE.Vanderyst695, from Zaire.
Pileus at first white (?), dried beige to dull pale brownish or beige-ochraceouswith
paler margin,long sulcate, glabrous,strongly convex, umbilicate, with a small umbo or
papilla in the umbilicus, 1-3 mm broad. Lamellaewhite, distant (9-12 through-lamellae,
no lamellulae), broad, some forked, distinctly collariate. Stipe bister to blackish with
white apex, glabrous,setiform, smooth, insititious, some not developingpilei (telepodia),
some associated with very thin, black rhizomorphsbut not attached to them, 10-20 X
0.1-0.2 mm. Context of the pileus white, very thin.
Spores 6.8-10 X 2.8-4.8,, ellipsoid, fusoid or oblong, some cylindrical, smooth,
hyaline, inamyloid. Hymenium: basidia 18 X 5.7 u; cystidia none; cheilocystidia like
the epicuticularbroom cells. Hyphae of the pileus trama densely packed and inter-
woven, basically runningradially,parallelin the stipe, pseudoamyloid, with clamp con-
nections. Cortical layer: epicutis of the pileus hymeniform, consisting of hyaline or
subhyaline broom cells of the Rotalis-type, main body 12-24(-35) X 10-19(-25) , ves-
Marasmius 99

iculose, often pedicellate to 10 u deep, setulae hyaline to pale melleous 1-2.5 X 0.5-1.5 p.
On coriaceous, fallen, dicotyledonous leaves, in tropical forest. South America
and tropical Africa.
MATERIALSTUDIED. VENEZUELA. Sucre:alongRio Media,15 Jul 1972,Dumont
et al VE 5145 (NY). ZAIRE. Bas-Congo,Kisantu,Vanderiyst695 (BR), type.
Singer (1964b) fig 16, H- I.

39. Marasmiusrotalis Berkeley & Broome, Jour. Linn. Soc. Bot. 14: 40. 1873.
TYPE. Twaites 810, from Ceylon.
Pileus whitish to pale cream, soon (even before being dried) turning dirty beige or
brownish(e g "oakwood," "wigwam"),center "coffee" M&P,grayishbeige or concol-
orous with the margin, sulcate, glabrousor sometimes rivulose-subscabrousin a zone
around the disc (and this zone consequently more pallid even in dried materialbut not
constantly so) the umbo itself not discolorous-darkbut pallid to light avellaneous-argill-
aceous, convex, umbilicate, with a small, low umbo or an obtuse papilla in the umbilicus,
1.5-5 mm broad. Lamellaewhite to pale cream, with white to (dried) slightlydiscolorous
edge (concolorous with the marginof the pileus), distant (13-17 through-lamellae,no
lamellulae or more rarely 1-2 lamellulaepresent) broad, collariate, the collariumwhen
dried also concolorous with the gill edges. Stipe black, blackish bister, or very dark
chestnut, with white apex, glabrous,smooth, insititious, equal, usually accompaniedby
thin, dark rhizomorphsbut never attached to them. Context very thin, inodorous.
Spores 6.8-8.8 X 3-4 p, oblong, smooth, hyaline, inamyloid. Hymenium:basidia
19 X 5.5 u; cystidia none; cheilocystidia like the epicuticularbroom cells. Hyphae of
pileus-tramainamyloid, with clamp connections. Cortical layers: epicutis of pileus
hymeniform, broom cells 10-23 X 5-15 s,, of the Rotalis-type,hyaline, with thin wall
and brown-fuscoussetulae which are divergent, 1.3-3.5 X 0.6-2.5 1, mostly 1.8-2.8 X
1.2-1.4 g, rod-shaped,obtuse.
On dead coriaceous leaves of dicotyledonous trees, pantropical. Known host:
MATERIAL STUDIED.COLOMBIA. Boyaca.:Tunja,Arcabuco,28 Jul 1960,SingerB
3560 (BAFC).CEYLON.Peradiniya, Thwaites810 (K),type. ZAIRE.Yangambi, ReserveIsa-
lowe,470 m alt Louis14934 (BR).
This species is close to Marasmiuscundinamrarcae, M. apatelius, and M. leuco-
rotalii In the Colombianoak woods it may most easily be confused with M. apatelius
which has fewer lamellae and M. cundinanarcae with more numerouslamellae and
ILLUSTRATION. Singer (1964b), fig 16, E-G.

40. Marasmiuscastellanoi Singer, sp nov

TYPE.Singer B 4199, from Brazil.
Pileo albo vel albido-pallido,umbone exiguo subericolorein umbilico centraliin-
serto; lamellis albis, distantibus,collariatis;stipite primumnigricanteapice albo. Sporis
5-6 X 2.5 :;hyphis pilei plerumquepseudoamyloideis,fibulatis; elementis epicuticulari-
bus typi Marasmiirotalis praesentibus. Ad folia emortua dicotyledonea in Brasilia.
Typus in BAFC conservatusest.
Pileus fresh white to whitish-pallid,on ridges becoming "cork"(M&P) when dry,
white around a "cork" colored umbo, glabrous,deeply sulcate, convex, umbilicate and
umbonate in the umbilicus, 5(-16) mm broad. Lamellaewhite with white edges, med-
ium broad, distant (9-11 through-lamellae,no lamellulae)distinctly collariate. Stipe
100 Flora Neotropica,

blackish at first with white apex, glabrous, equal, insititious, about 11 X 0.1-0.2 mm.
Context of pileus white, thin, inodorous.
Spores 5-6 X 2.5 p, ellipsoid, smooth, inamyloid. Hymenium: basidia 5-5.5 p
broad; cystidia none; cheilocystidia like the epicuticularbroom cells. Hyphae of the
hymenophoraltrama and the pileus-tramahyaline, many of these distinctly pseudoamy-
loid, with clamp connections. Cortical layer: epicutis of the pileus hymeniform, con-
sisting of broom cells of the Rotalis-type, main body 17-20 X 12 u, hyaline, setulae
short and divergent.
On dead dicotyledonous leaf in tropical rain forest.
MATERIALSTUDIED. BRAZIL:Para, EstanciaPirelli,7 Jun 1966, SingerB 4199 (BAFC),

41. MarasmiuspandoanusSinger, Sydowia 18: 339. 1965.

TYPE.Singer B 2145, from Bolivia.
Pileus white with a blackish dot in the center, the latter becoming pale fuscous in
dried material, outside the disc strongly sulcate, convex, umbilicate, about 5 mm broad.
Lamellaewhite, edges concolorous with sides, broad, distant (11), collariate with a very
distant collarium. Stipe black with white apex, glabrousand smooth, setose, insititious,
30 X 0.1 mm. Context very thin, inodorous.
Spores 14.5-16.5 X 3.7-4.3 p, ventricose-oblongwith suprahilardepressionand a
conspicuous oblique hilar appendage,hyaline, smooth. Hymenium:consisting of rather
short basidia and basidioles, hyaline; cheilocystidia 18-27 X 6-14.5 u, of the Rotalis-type,
with setulae divergingand short (1-2.8 p) and sometimes (those on summit of cells)
branching-forked,entirely subhyaline, clavate or subventricoseand taperingto an obtuse
mucro or vesiculose, rarely hyphous. Hyphae of the pileus more or less pseudoamyloid
with clamp connections; those of the stipe strictly parallel,3.3-4 , in diameter,with
somewhat thickened wall, in central hyaline strand strongly pseudoamyloid. Cortical
layers: epicutis of pileus basically hymeniformly arrangedbut slightly disarrangedin
places consisting of the same elements that occur on the edges of the lamellae but
more consistently vesiculose and then the upper portion often horizontally elongated
so that they appearalmost short-hyphous(as in Marasmzius ntulticeps);surface of stipe
with some minute outgrowths similarto the setulae of the cheilocystidia.
On fallen leaves of various species of dicotyledonous trees, (including Theobroila)
in inundation zone of rain forest.
MATERIAL STUDIED.BOLIVIA.Pando:Manuripi, SantaRosaon RibMladre de Dios,22
Mar1956,SingerB 2145 (LIL),type.
ILLUSTRATION: Singer (1965) pl 7, fig 15.

42. Marasmiusoaxacanus Singer, Sydowia 12: 232. 1958.

TYPE.Singer M 1519, from Mexico.
Pileus white, becoming whitish-argillaceousor argillaceous-pallidwhen dried, with
a white zone around a pale fuscous to fuscous-subfuliginousdot which becomes light
gray, pale fuscous or whitish when dried, smooth in the umbilicus, otherwise long sul-
cate, glabrous,convex, with deep umbilicus without an umbo or papilla inside but ap-
pearingtomentose when seen under a lens (dried), 3-4 mm broad. Lamellaewhite, sub-
close at first, subdistant to distant (10-16 through-lamellaeno lamellulae),sometimes
forked, very broad, distinctly collariate with a free collarium. Stipe deep brown to
black with at first white apex, glabrous, smooth, setiform, insititious, not seen accom-
Marasmius 101

panied by black rhizomorphs,20-30 X 0.2-0.4 mm. Context of pileus white, thin,

Spores 10-14 X 3-4 p, fusoid, smooth, hyaline, inamyloid. Hymenium: basidia
16-22 X 6.3-7.5 1, 4-spored;cystidia none; cheilocystidia like the epicuticularbroom
cells but always entirely hyaline. Hyphae of the pileus tramahyaline, thin-walled,
with clamp connections, distinctly pseudoamyloidin pileus and stipe. Corticallayers:
epicutis of the pileus hymeniform or subhymeniform,consisting of broom cells of the
Rotalis-type, these entirely hyaline or some melleous-hyalineexcept those of the area
of the dark, central dot of the pileus, either subhyalineor pale grayishfuscidulous,
main body 15-24 X 15-22 m,vesiculose to vesiculose-subcylindric,oftenglobose, se-
tulae thin-rodshaped,1.3-1.5 M projecting.
On fallen leaves of Inga, CentralMexico.
MATERIALSTUDIED. MEXICO:Oaxaca,Huautlade Jimenez, 1660 m alt, 10 Jul 1957,
SingerM 1519 (LIL) type; 8 Jul 1969, SingerM 8330 (F).
Young carpophoresof Marasmiustanyspermusshould not be mistaken for Maras-
mius oaxacanus
ILLUSTRATION. Singer (1965), pl 11, fig 54.

43. MarasmiusmanuripiensisSinger, Sydowia 18: 338. 1965.

TYPE.Singer B 2227, from Bolivia.
Pileus white to cream color with a minute ochraceousto pale fuscous cen-
tral dot, dried pale ochraceouscinnamon or light caf6 au lait with concolorous or most-
ly cinnamon fuscous central dot, localized in a small umbo inside an umbilicus, sulcate,
convex, 2-5 mm broad. Lamellaewhite, equal, 6-9, rathernarrow,collariate to a nar-
row and moderately distinct collarium,distant. Stipe black with white apex, remaining
so in dried condition, glabrousand smooth, setose, insititious, 4-10 X 0.2-0.5 mm.
Spores 7.5-11 X 4-5 A,ellipsoid, smooth, hyaline, with ratherthin, inamyloid wall.
Hymenium:basidia 19 X 7.2 ,; basidioles fusoid, often with mucro; cystidia none; cheilo-
cystidia like the epicuticularelements. Hyphae of the pileus and hymenophoraltrama
partly inamyloid, partly weakly pseudoamlyoid,inamyloid in subhymenium;all hyphae
with clamp connections. Corticallayers: epicutis of pileus consisting of elements of the
Rotalis-type, hymeniformlyarranged,vesiculose, hyaline with hyaline to yellowish-sub-
hyaline setulae, the main body 20-25 X 15-18 p, setulae divergentover the upper por-
tion of cell but sometimes somewhat transient towards the Siccus-type, small, mostly
cylindric and obtuse, 1.3-2.2 X 0.4-0.9 w.
On dead dicotyledonous twigs and petioles, fallen from a tree of the rain forest.
MATERIALSTUDIED. BOLIVIA. Pando:Manuripi,Conquista,25 Mar1956, SingerB 2227
(LIL), type.
Singer(1965), pl 7, fig 16.

44. MarasmiuspsychotriophilusSinger, Sydowia 18: 350. 1965. Fig 45P.

TYPE. Singer B 4029, from Brazil.
Pileus whitish with white center, eventually entirely whitish or with grayish-cream
center, old and dried often entirely grayish-cream,glabrous,sulcate, convex, somewhat
depressedaround a central umbo, 1-1.5 mm broad. Lamellaewhite with white edge,
distant (seven through-lamellaeonly), broad, distinctly collariate. Stipe white, later
black from the base upwards,glabrous,insititious, 10-15 X 0.1 mm. Context white in
the pileus, thin, inodorous.
Spores 12.5-14.2 X 7.5-9,, ellipsoid-fusoid,broad, smooth, hyaline, inamyloid.
102 Flora Neotropica

Hymenium: basidia 24.5-26.8 X 12.5-13 , 4-spored;cystidia none; cheilocystidia 15-20

X 12.5-18.5 ,, vesiculose, hyaline, of the Rotalis-type, setulae 1.2-1.5 X 0.2-0.9 ,. Hy-
phae of the pileus tramathin-walled,filamentous, inamyloid. Corticallayers: epicutis
of the pileus hymeniform, consisting of elements much like the cheilocystidia but set-
ulae mostly pale avellaneous(from dried materialin KOH).
On fallen leaves of Psychotriac
MATERIAL STUDIED.BRAZIL: Rio de Janeiro,SerradosOrgaos,24 Oct 1961,SingerB
4029 (BAFC)type.

45. MarasmiuscundinamarcaeSinger, sp nov Fig 45C.

TYPE. Singer B 6009, from Colombia.
Pileo albo vel pallide alutaceo-albo,umbone exiguo atro praedito, 5-6 mm lato;
lamellis cremeo-albis,moderatim distantibus,sat latis, collariatis;stipite fuligineo apice
albo; rhizomorphisnullis. Sporis 5.5-7.2 X 2-3 A; elementis epicuticularibustypi Mara-
smii rotalis. Ad folia delapsaquercinain Columbia. Typus in F conservaturest.
Pileus white to pale buff-white on dehydrationand in the herbariumbecoming
pale beige around a white circularzone surroundinga fuliginous dot, sulcate except in
the center, glabrous,convex, moderately deeply umbilicate with a dark, low umbo or
papilla in the middle of the depression,5-6 mm broad. Lamellaecream white with
white edge, moderately distant (+ 20 through-lamellae;no lamellulae)ratherbroad,
collariate. Stipe fuliginous with white apex, glabrous,insititious, not accompaniedby
black rhizomorphs,equal or slightly taperingupwards, 28-30 X 0.3-0.8 mm. Context
white, thin, inodorous.
Spores 5.5-7.2 X 2-3 l, ellipsoid to oblong or fusoid, smooth, hyaline, inamyloid.
Hymenium: basidia4-spored;basidioles fusoid; cystidia none; cheilocystidia like the
epicuticularbroom cells, all hyaline. Hyphae in pileus and stipe pseudoamyloidwith
clamp connections. Corticallayer: epicutis hymeniform, of broom cells of the Rotalis-
type, 10-21 X 9-18 M,vesiculose or subvesiculose,with setulae 1-1.2 X 0.5-1 ,u,all hy-
aline (fresh or recently dried in KOH).
On fallen oak leaves in oak woods. Colombia.
MATERIAL STUDIED.COLOMBIA. Cundinamarca:Bosquede la Mercedes, Bojaca,8
Apr 1968, SingerB 6009 (F), type.

46. MarasmiusleucorotalisSinger, Sydowia 18: 337. 1965.

46a. Marasmiusleucorotalisvar leucoratalis

TYPE.Singer B 1414, from Bolivia.
Pileus white, with a fuliginous or black dot in the center, eventually tending to
become pale beige in the marginalzone on drying but leavinga white zone around the
black dot, glabrous,sulcate, convex, umbilicate, the bottom of the umbilicus either
flat or with a low umbo, 3-4 mm broad. Lamellaewhite with white edge or cream
white, subdistantor medium distant (14-15 through-lamellae,no lamellulae)rather
broad, distinctly collariate. Stipe black or fuliginous with white apex, glabrous,smooth,
equal or slightly taperingupward, setiform, insititious, 15-17 X 0.2-0.3 mm; no black
rhizomorphsobserved. Context of the pileus white, thin, inodorous.
Spores 7-7.8 X 2.7-3 pi,ellipsoid to oblong or fusoid, smooth, hyaline, inamyloid.
Hymenium:basidia 19-20 X 4.5-5 u, 4-spored;cystidia none; cheilocystidia like the
epicuticularbroom cells, hyaline. Hyphae filamentous,hyaline, pseudoamyloidboth
in the pileus and stipe, with clamp connections. Corticallayer: epicutis of the pileus
Marasmius 103

hymeniform, consisting of broom cells which are 12-35 X 10-33 ,, of the Rotalis-type,
vesiculose and sometimes broaderthan long, sometimes longer than broad, sometimes
pedicellate, setulae dow to the middle (or lower) of the cell, rod-shaped1-2 X 0.3-1 p,
all hyaline.
On fallen dicotyledonous leaves in tropical-montaneforest. Bolivia.
MATERIALSTUDIED. BOLIVIA. La Paz; Nor-Yungas,Rio Yariza,23 Feb 1956, Singer
B 1414 (LIL) type.
Singer(1965) pi 7, fig 17.

46b. MarasmiusleucorotalisvardiscopallescensSinger,var nov

TYPE. Singer T 3772, from Argentina.
Maculo centrali pilei pallescente lamellisquemagis distantibusdiffert.
Pileus white, with fuscous-blackto light fuscous central dot but this in dried
condition pallescent to whitish or gray and surroundedby a white ring even when the
marginalzone has assumed a pale beige discolorationin the herbarium,more often
without such a white ring-zone,glabrous, sulcate, hemisphericalto convex, umbilicate,
with or without a papilla in the depression, 1-3 mm broad. Lamellaewhite with white
edges, distant (10-11 through-lamellae,no lamellulae),broad, distinctly collariate with
white collarium. Stipe black with white apex, dried fuscous bister with white apex,
somewhat shining, glabrous, smooth, insititious, not accompaniedby black rhizomorphs,
10-15 X 0.1 mm. Context of pileus white, thin, inodorous.
Spores 5.5-7 X 2.8-3.5 p(Q = ? 2), ellipsoid, smooth, hyaline, inamyloid. Hy-
menium: basidia 4-spored;basidioles fusoid; cystidia none; cheilocystidia like the epi-
cuticularbroom cells, all hyaline. Hyphae with clamp connections. Corticallayer:
epicutis hymeniform, broom cells voluminous, with pale brownish to hyaline setulae
2 uhigh and rod-shaped,divergent.
On fallen leaves of Phoebe porphyria in subtropicalforest.
MATERIALSTUDIED. ARGENTINA:Tucumain,Rio Cochunaat 1350 m alt 27 Feb 1952,
Singer T 3772 (LIL), type.

46c. Marasmiusleucorotalisvar leucozonites (Singer) Singer,comb nov

Marasmiusleucozonites Singer,Sydowia 18: 337. 1965.
MarasmiuspluvialisSinger,Sydowia 18: 341. 1965.
TYPE.Singer T 2018, from Argentina.
Pileus white to dull ochraceous(fresh), elevations between sulci paler or white,
depressionaround dark central spot white and remainingso but marginalzone tending
to become pale sordid ochraceous to pale argillaceous-beigein the herbarium,glabrous,
sulcate, deeply umbilicate, not papillate in the umbilicus when fresh but on drying
tending to become slightly umbonate, 3-5.5 mm broad. Lamellaewhite, with either
white or pale dirty ochraceous to pale beige edges when dried, often near the margin
of the pileus discoloring, subclose when young, subdistant to distant when old (11-16
through-lamellae,no lamellulae)broad, collariate. Stipe dark umber with white apex,
later almost black all over, glabrous,smooth, setiform, shining, insititious, not associ-
ated with black rhizomorphs,equal, 25-32 X 0.1-0.3 mm. Context of pileus white,
thin, inodorous.
Spores 6.7-7.5 X 2.8-3(-4) p, oblong to fusoid, smooth, hyaline, inamyloid. Hy-
mneium: basidia 4-spored;basidioles fusoid; cystidia none; cheilocystidia like epicuti-
cular broom cells (some with pale melleous, brownish setulae). Hyphae hyaline in the
pileus, thin-walled,weakly pseudoamyloid,with clamp connections; hyphae of the apex
of the stipe parallel,hyaline, strongly pseudoamyloid. Corticallayer: epicutis hymeni-
104 Flora Neotropica

form, broom cells of the Rotalis-type e g 28 X 16 p,, beset with hyaline to (in herbar-
ium materialmostly) pale melleous brownish setulae 202.7 plong, divergent.
On leaves of dicotyledonous trees, frequently (but not exclusively) on Phoebe
MATERIALSTUDIED. ARGENTINA. Tucuman,Sierrade San Javier,CiudadUniversit-
aria, 1000 m alt, 15 Jan 1955, Singer T 2018 (LIL), type.
The precedingvariety differs from this in being less pigmented, smallerand with
fewer lamellae and the central dark dot as well as the stipe becoming paler on drying,
also in slightly smallerand relatively broaderspores. Both differ from the type variety
by a few rather secondary characters;the type variety appearsto be intermediatebe-
tween the other two varieties.
The binomialMarasmiuspluvialis was intended for a species I have now redes-
cribed as Marasmiushvlaeicola Singer (no. 63). Because of an unfortunate printer's
error, the description of Marasmiusleucozoniiteswith its type designation(T 2018) has
been repeated under the headingMarasmiuspluvialis Singer (1965, p 163) so that
formally it becomes a synonym of Marasmiusleucozonites.
Young specimens of Marasmiusplatvspermusmight be looked for here or under
M cundinamarcae. It differs however in habitat and broaderspores.

47. MarasmiusarimanusDennis, Trans. Brit. Mycol. Soc. 34: 418. 1951 (as Xi. ari-
TYPE. Dennis 391, from Trinidad.
Pileus sepia or mummy brown, striate-sulcateto the center, convex with a
shallow umbilicus, 2-3 mm broad. Lamellaepale buff, distant (7), moderately broad,
attenuated to each end, attached to a shallow collarium forming little more than a
ridgeround the apex of the stipe, with sepia colored edges. Stipe almost black, slen-
der, wiry, smooth and polished throughout. Context thin, white, membranous.
Spores 11-14 X 3.5-4 ,, tear-shaped,inamyloid. Hymenium: cheilocystidia like
the epicuticularbroom cells, 8 lpbroad. Coveringlayers: epicutis of pileus formed by
pear-shapedelements with reddish brown processes.
On bark of dead logs, Trinidad:Arima, MissDora Kong, 28 Nov 1949, comm.
Dennis 391 (K), type.
I have studied the material superficially;it consists of a fragmentwhich I did
not dare section. It is not fully clear whether the epicuticularbroom cells and cheil-
ocystidia are of the Rotalis-type or the Siccus-type. The species seems to be close
to MarasmiusfuligineorotulaSinger. The description is adopted from Dennis' original
account. It would suggest that even if the broom cells were considered as being of
the Siccus-type, the spores would be more elongated and the carpophoresmuch
smaller in Dennis' species.
ILLUSTRATION. Dennis (1951a) pl 23, fig 5.

48. MarasmiusgriseofuscescensSinger, sp nov

TYPE.Singer B 7361, from Ecuador
Pileo griseo, siccando fusco, 1.5-2.2 mm lato; lamellis albis, siccando subcinereis,
distantibus,collariatis;stipite fuligineo vel atrocastaneoapice albopallido. Sporis 9-11
X 4-5.5 u; hyphis fibulatis, in tramate pilei inamyloideis;elementis epicuticularibustypi
Marasmiirotalis. Ad folia dicotyledonea delapsa.
Pileus at first almost uniformly gray, later gray striate on white ground, dried
fuscous, with a flat or slightly convex pale grayish-pallidto cinereous ground of the
Marasmius 105

non-papillateumbilicatus,glabrous,sulcate, convex, about 1.5-2.2 mm broad. Lamellae

white, dried palest cinerous or cinereous pallid, with white edge, broad, distant (10-1
through-lamellae,no lamellulae)not intervenose,collariate. Stipe fuliginous to dark
chestnut with white-pallidapex, glabrous,setose, equal, insititious, (5-)20-25 X 0.2-0.3
mm. Context white, thin.
Spores 9-11 X 4-5.5 ,, ellipsoid, smooth, hyaline, inamyloid. Hymenium: bas-
idia 21-26.5 X 7-9 ,, 4-spored;basidioles fusoid; cystidia none; cheilocystidia like the
epicuticularbroom cells but all hyaline. Hyphae filamentous, some swollen to 15 p,
hyaline in NH40H, with clamp connections, in the trama of the pileus and hymeno-
phore inamyloid. Cortical layer: epicutis of the pileus hymeniform, consisting of
broom cells of the Rotalis-type, main body vesiculose or subvesiculose,thin-walled,
12-20 X 9-15,, hyaline, setulae melleous to brownish, rod-shaped,1.5-2.2 X 0.8-1 .
On dicotyledonous leaves fallen to the ground in tropical rain forest, perhaps
also in subtropicalforest.
MATERIAL STUDIED. ECUADOR. Napo, Lago Agrio, 11 May 1973, Singer B 7361 (F)
This species is characteristicbecause of the change of color from gray to fuscous
when dried, the white edges of the lamellae and the medium sized spores. These are
scanty in the type; consequently, the measurementsmay not cover the entire breadth
of variation.
It is probable that the specimens registeredby Singer(1965, p 173) as Marasmius
aff. rotalis are identical with IM.griseofuscescens. These were collected by Singer& Dig-
ilio (ll 26, M .3) in Argentina,Misiones,Cataratasdel lguazfi, 22/24 Nov 1949 (LIL)
and are perhaps specimens which have passed the originalgray state.
ILLUSTRATION.Singer (1958a) fig 42.

49. Marasmiusscototephrodes Singer, sp nov Fig 46.

TYPE.Singer 8299, from Mexico.
Pileo cinereo, maculo centrali griseo zona alba vel concolori cingulato, umbilicato,
4-6 mm lato: lamellis albis acie atrogriseis,distantibus,griseo-collariatis;stipite nigro;
rhizomorphistenuissimisnigris praesentibussed stipiti haud adhaesis. Sporis 7-9 X 3.5-
4.7 u; cheilocystidiis sepiaceo-setulosis;hyphis fibulatis, pseudoamyloideis;elementis
epicuticularibustypi Marasmiirotalis. Ad folia, petiolos, frustulaDicotyledonum in
Mexico. Typus in F conservatusest.
Pileus ash gray with a white or concolorous zone around a central light gray to
dark gray dot, in the intermediatezone and the center smooth but surface tending to
become cracked, marginalzone long-sulcate,glabrous,convex, umbilicate, not papil-
late, 4-6 mm broad. Lamellaewhite with cinereous to dark gray edge, distant (10-12
through-lamellae,no lamellulae), broad, distinctly collariate, collariumgray. Stipe
black, shining, equal, setiform insititious, 20-25 X 0.1-0.2 mm, accompaniedby very
thin black rhizomorphsbut never attached to them: some telepodia present. Context
thin, inodorous.
Spores 7-9 X 3.5-4.7,, ellipsoid, some oblong ellipsoid, few subcylindric(but
broad), smooth, hyaline, inamyloid, with a slight suprahilardepression. Hymenium:
basidia 19-21.5 X 5.5-6.8 ,; basidioles fusoid; cystidia none; cheilocystidia like the
epicuticularbroom cells but mostly with sepia, rarely subhyaline setulae and hyaline
main body. Hyphae of the center of the pileus and the apex of the stipe weakly to
distinctly pseudoamyloid, with numerousconnections. Cortical layer: epicutis of the
pileus hymeniform, consisting of broom cells of the Rotalis-type, main body 10-20 X
106 FloraNeotropica

7-16.5 u, ventricose or ventriocse-subclavate,hyaline or fuscidulous to sepia, with up

to 0.5 , thick wall, setulae 1.5-5 X 0.7-1.3 ,u, rod-shapedand divergent,fewer hyaline
than sepia.
On dicotyledonous leaves and small sticks, petioles, etc. In Mexico.
MATERIAL STUDIED. MEXICO. Oaxaca, Rancho del Cura (Sierra Mazateca) 7 Jul 1969,
1580 m alt, Singer M 8299 (F), type.

50. MarasmiusleucozonitiformisSinger sp nov

TYPE. Singer B 6105 A, from Colombia.
Pileo griseo vel pallide griseo, zona albida circum maculam centralematrum prae-
dito, 1.5-5.5 mm lato; lamellis albidis acie frequenterdiscoloribus,distantibus,collari-
atis; stipite nigro acie albo. Sporis 5.5-8 X 2.5-4 /; elementis epicuticularibustypi
Marasmiirotalis. Ad folia dicotyledonea in Columbia. Tupus in F conservatur.
Pileus gray, pale gray, often turningbeige on drying, retaininghowever a white
or whitish zone around the dark dot in the center of the umbilicus, this zone whitish
because minutely rivulosecracked(visible undera lens), showingthe trama,the
dark spot subfuliginousbut often pallescent to pale gray or whitish on drying, macro-
scopically glabrous,sulcate except for the umbilicus, convex, at first shallowly later
deeply umbilicate, without an umbo or papilla in the depression, 1.5-9 mm broad.
Lamellaewhite but on drying often showing a discolorous(brownish) edge and a
brownish collarium, distant (8-12 through-lamellae,no lamellulae), sometimes forked,
medium broad to very broad, distinctly collariate. Stipe black with white apex, later
entirely black, glabrous,setiform, insititious, with numerousblack rhizomorphsaccom-
panying (but not connected with) the carpophores,3.5-18 X 0.1-0.2 mm. Context of
pileus white, thin, inodorous.
Spores 5.5-8(-9.3) X 2.5-4 1, ellipsoid to more often oblong to fusoid, smooth,
hyaline, inamyloid, Hymenium: basidia 21-24 X 5 ,, 4-spored, sometimes some 2-spored
ones intermixed;basidioles fusoid, subacute;cystidia none; cheilocystidia like the epi-
cuticular broom cells but many or most hyaline. Hyphae of the trama of the pileus
hyaline, filamentous, with clamp connections, inamyloid;hyphae of the apex of the
stipe hyaline, later pigmented, parallel,weakly pseudoamyloid. Cortical layer: epicutis
of the pileus hymeniform, consisting of broom cells of the Rotalis-type, main body
12-26 X 6-18 p, with thin, rarely thick (to 1 j,) subhyaline or hyaline wall and with
often fuscidulous cell sap, with mostly brown setulae 1-2.5 I projecting, divergent.
On leathery, dead fallen leaves of dicotyledonous trees in tropical-montane
rain forest in Colombia, CordilleraOccidental.
MATERIAL STUDIED. COLOMBIA. Valle, Mun. Dagua, Corr. Vergel, Finca La Araucana,
14 Apr 1968, Singer B 6105 A (F) type; Singer B 6105 (F); up to 1900 m alt above km 25 of
Dagua-road, Mun. Dagua, 28 Apr 1968, Singer B 6444 (F); Buenaventura, Finca La Flora, 23 Apr
1968, SingerB 6316 (F).
Among the species with originally white pileus, this pale gray or gray species is
closest to Marasmiusleucorotalis from which it differs, aside from the colors, in less
pseudoamyloid tramal hyphae and the association with black rhizomorphs;from Mara-
smius rotalis it differs in having fewer lamellae.

51. MarasmiustanyspermusSinger, Sydowia 18: 343. 1965.

TYPE,Singer B 2052, from Bolivia.
Pileus at first whitish, then brown ("eldorado" M&P)with paler margin,a zone
concolorous with the marginalone surroundingthe central dark dot which becomes
pale sordid gray in dried material, sulcate except in the umbilicus and with a small
Marasmius 107

umbo correspondingin extension to the blackish dot inside the umbilicus, convex
otherwise, 1.5-3.2 mm broad. Lamellaepale cream, 10-11, ratherbroad, with white
edge, collariate. Stipe umber with white apex, glabrous,insititious, 20 X 0.1 mm. No
Spores 10.3-15 X 2-3.5 p, hyaline, smooth, inamyloid. Hymenium: basidia 18-
19.3 X 5.5-6 p; cystidia none seen; cheilocystidia like epicuticularelements. Hyphae
of pileus, hymenophoraltrama hyaline, with clamp connections, in stipe (apex and
hyaline interior of lower portion) moderately strongly pseudoamyloid,somewhat thick-
walled, otherwise inamyloid to very weakly pseudoamyloid. Coveringlayers: epicutis
of pileus hymeniform, consisting of cells of the Rotalis-type, hyaline, 9-37 s,broad,
beset with hyaline or (in places) pale melleous setulae, these small (1.2-1.8 X 0.8-1 a),
On dead dicotyledonous leaves in tropical rain forest.
MATERIALSTUDIED. BOLIVIA. Beni: Vaca Diez, Guayaramerin,16 Mar1956, Singer
B 2052 (LIL), type.
When seen in the white stage, the carpophoresmay be mistaken for those of
Marasmiuspanldoalus or Marasmiusoaxacanus which do not become brown when
mature and have broaderspores.

52. Marasmiuscarneotinctus Singer, sp nov Fig 47.

TYPE. GuzmnOn 4580, from Colombia.
Pileo carneo-roseo,atrobrunneo-papillato,3-4 mm lato; lamellis carneo-roseis,sub-
confertis, collario saepe incompleto; stipite atrobrunneoapice albido. Sporis 8.5-9 X
3.5-4 A; elementis epicuticularibustypi Marasmiirotalis. In silva tropicali-montana
Columbiae. Typus in ENCBconservatur.
Pileus flesh-pinkwith deep brown papilla, glabrous, sulcate, towards marginfine-
ly venose under a lens, convex, deeply umbilicate with a low papilla in the umbilicus,
34 mm broad. Lamellaeflesh-pink,subclose (17 through-lamellae,no lamellulae)
with an often poorly developed collarium. Stipe blackish-brownwith whitish apex,
glabrous,insititious, about 40 X 0.4-0.5 mm. Context thin, inodorous.
Spores 8.5-9 X 3.5-4 ,, oblong, smooth, hyaline, inamyloid. Hymenium: basi-
dia clavate; basidioles fusoid; cystidia none; cheilocystidia scattered, 24-36 X 3.8-6.5 ,u,
basidiole-like,subcylindric-fusoid,with wavy-uneven,somewhat lobed or entire apex,
in the latter case obtuse and sometimes subcapitate,hyaline. Hyphae of the pileus
trama remarkablybroad, hyaline, with firm to somewhat thick or firm wall, with clamp
connections, pseudoamyloid. Cortical layer: epicutis of the pileus hymeniform, broom
cells of the Rotalis-type, 18-32 X 15-31 s, vesiculose, most with somewhat thickened
or firm walls, setulae low and obtuse, melleous-hyalineto pale melleous, divergent.
In tropical-montaneforest, Colombia, west slope of the CordilleraOccidental.
MATERIALSTUDIED. COLOMBIA.Valle, along road to Buenaventura.Pacific slope,
1950 m alt, 26 Jul 1964. Guzman4580 (ENCB)type.

53. Marasmiusvioleorotalis Singer, sp nov Fig 48.

TYPE.Singer B 6450, from Colombia.
Pileo violaceo-lilaceo;lamellis lilaceo-pallidis,collariatis;stipite nigro. Sporis 6.2-
9.5 X 2.5-3 ,; elementis epicuticularibustypi Marasmiirotalis. Ad folia dicotyledonea
emortua in Columbia. Typus in F conservatusest.
Pileus violet-lilac with an almost pallid zone around the fulvous-brownpapilla,
this zone not distinctly white but appearingpallid because of the rivulose-cracked
108 Flora Neotropica

visible (under a lens) surface, showing the trama, glabrous,sulcate, excepting the cen-
ter, strongly convex, umbilicate, with a low papilla in the umbilicus, about 5 mm broad.
Lamellaelilac-pallidwith pale lilac edge and pale lilac ground, distant (+ 12 through-
lamellae, no lamellulae), broad, collariate. Stipe black, glabrous, flexuous, insititious,
55-58 X 0.2 mm. Context inodorous.
Spores 6.2-9.5 X 2.5-3 p, mostly 7.5-8.5 X 2.5 j,, fusoid to subcylindrical,smooth,
hyaline, inamyloid. Hymenium: basidia 14-15 X 4.5-5.5 p; basidioles fusoid; cystidia
none; cheilocystidia like the epicuticularbroom cells but more often subhyaline. Hy-
phae in the pileus hyaline (KOH), inamyloid, with clamp connections. Cortical layer:
epicutis of the pileus hymeniform, broom cells of the Rotalis-type, 13-21 X 6-14 ,,
vesiculose, more rarely clavate, setulae mostly pale melleous, short, rod-like, 1-1.5-(2)
x 0.4-0.8-(1) 1,
On dead dicotyledonous leaves in tropical-montaneforest, CordilleraOccidental,
MATERIALSTUDIED. COLOMBIA.Valle, near Daguaroad above km 25, at 1900-2000
n alt, 28 Apr 1968, SingerB 6450 (F) type.

54. MarasmiuscarminisSinger, sp nov

MarasmiuscarminisSinger,Sydowia 18: 156. 1965 (without Latin).
TYPE. Singer B 1484, from Bolivia.
Pileo purpureo-rubro,circum maculam nigramcentralem albo vel rubro, 0.5-2 mm
lato et alto; lamellis albis, rubromarginatis,12-13 integris tantum; stipite 15-35 mm longo,
rhizomorphisnullis. Sporis 7-11 X 4-5.5; ;cheilocystidiis typi Marasmiirotalis. Ad fo-
lia emortua dicotyledonea in silva tropicali-montana. Typus in LIL conservatusest.
Pileus deep purple red, in age and particularlyon drying fading to red-brown
("chutney"), sulcate, white around a central black dot or concolorous there, umbilicate,
in dried materialgray to gray-blackin umbilicus, bottom of umbilicus somewhat con-
vex very young caps convex and umbonate, later convex-umlbilicatebut remaining rather
high, 0.5-2 mm broad, 0.5-2 mm high, sulcate purple zone almost vertical. Lamellae
white with discolorous edge near the marginof the pileus and likewise colored on the
edge of the collarium, about 12-13, broad. Stipe buff brown to blackish brown with
white apex, glabrousand smooth, shining, setose, insititious, 15-35 X 0.1-0.2 mm. No
Spores 7-11 X 4-5.5 u, mostly 8.5-10.5 X 4.5-5 p, hyaline, smooth. Hymenium:
basidia 16.5 X 6.2 p, but also largerelements present which may be basidioles(up to
27 X 9.7 ,); cystidioles optically "empty," e g 28 X 9 u, thin-walled,ventricose, broadly
rounded above or with broadly rounded mucro; cheilocystidia like epicuticularbroom
cells but hyaline where edge is concolorous with sides of lamellae. Hyphae in pileus
trama moderately strongly but rapidly reacting: pseudoamyloid,with clamp connect-
ions. Coveringlayer: epicutis of pileus hymeniform, consisting of cells of the Rotalis-
type, although some form a transition towards the Siccus-type, diameter of the main
body 5-25 u, vesiculose-pedicellateor vesiculose and sessile, rarely forked or laterally
elongated, hyaline to pale brownishand beset with numerous rod-shapedto conic-spin-
ulose-obtuse setulae 2-3 X 1.5-2 p.
On dead dicotyledonous leaves in tropical montane forest.
MATERIALSTUDIED. BOLIVIA.La Paz: Nor-Yungas,CarmenPampa,26 Feb 1956, at
2000 m alt, SingerB 1484 (LIL), type; B 1477 (LIL), paratype.
ILLUSTRATION.Singer (1965), p1. 7, fig. 18.
Marasmius 109

55. Marasmiusbuzae Dennis, Kew Bull. 15: 98. 1961.

TYPE.Dennis 1775, from Venezuela.
Pileus a rather saturatedorange ochraceous to deep orange ferruginousbut be-
tween striae and around a central black dot paler to pallid, otherwise "tarragona"or
5-B12 M&Ppaler regions "sundown"or paler, little changingwhen dry (between "rust,
sorolla" and "Arab"M&P),sulcate over most of the declivous zone, with a flat-bottomed
umbilicus, truncate-conicto terete-cylindricor barrel-shaped,about 1.5 mm broad and
high. Lamellaewhitish with an ochraceous to ocher brown border near the marginof
the pileus and on the collarium, subdistant to distant, 7-11, very broad, collariate.
Stipe umber to umber-blackor black, glabrousand smooth, shining, insititious, 17-24
X 0.1 mm, rising from the substratumbut occasionally accompaniedby scanty, thick,
black rhizomorphs.
Spores voluminous, 8.5-13 X (5-)5.7-9(-10.3) , hyaline, smooth. Hymenium:
basidiavoluminous, 29-40 X 9.5-11.7 ,, 4-spored, sometimes a few 2-spored ones pre-
sent; cystidia none observed;cheilocystidia like epicuticularelements but with the hy-
aline ones predominatingat least over wide stretches of the edge of the lamellae. Hy-
phae rapidly weakly pseudoamyloid(but distinctly so) in trama of pileus, with clamp
connections. Coveringlayers: epicutis of pileus hymeniform, consisting of elements
of the Rotalis-type, 12-35 X 11-29 u, vesiculose to clavate or cylindrical, setulae div-
erging, short and rathercoarse, 1.3-2.3 X 1.3-1.6 p, often coveringthe whole upper
half of the cell, in some cells hyaline but mostly brown to brownish ochraceous.
On dead fallen dicotyledonous leaves in lower marginalzone of cloud forest.
MATERIALSTUDIED. BOLIVIA. La Paz: Nor-Yungas,CerroUchumachiat 2200 m alt
25 Feb 1956, SingerB 1433 (LIL);Cataratas"San Juan" 2400 m alt, 28 Jan 1956, SingerB 656
(LIL). VENEZUELA. LagunaNegra,Sierrade Santo Domingo,Estado Merida,1 Aug 1958, Dennis
1775 (K), type.
ILLUSTRATIONDennis (1961), fig 51.

56. MarasmiuspopuliformisBerkeley, Hook. Jour. Bot. 8: 140. 1856.

TYPE. Spruce, from Brazil.
Pileus fuscous or dark brown, sulcate over the convex surface up to a shallow
umbilicus with low dark umbo, less than I mm broad. Lamellae whitish, edges con-
colorous with pileus, 5-6 according to Berkeley, about 10 according to Dennis, collar-
iate, equal. Stipe yellowish, smooth, up to 3 mm long, hollow, arising from a thicker
golden yellow (yellowish when dry) smooth, polished rhizomorph, many stipes on a
rhizomorphic strand about 100 mm long, 0.1 mm thick.
Spores 7 X 3 ,, elliptical. Hymenium: cheilocystidia like epicuticular elements.
Cortical layers: epicuticularelements about 8 uin diameter, with short, reddish-brown
processes, apparentlyof the Rotalis-type,
On dead branches.
MATERIALSTUDIED. BRAZIL. Amazonas,Ipanure("Panure")on Rio Negro,Spruce
(K), type.
I have not sectioned the type since it had been studied previously by Dennis
(1951b) whose findings are incorporated in the description given above. The species
has never been recollected.
ILLUSTRATION.Dennis (1951b) fig 14; Berkeley (1856) pl 5. fig 2.

57. Marasmius minimus Dennis, Kew Bull. 15: 98. 1961.

TYPE. Denniis 1068, from Venezuela.
110 Flora Neotropica

Pileus Brusselsbrown (Ridgway), radially sulcate from all inconspicuous central,

darkerarea, convex, 1.5-2 mm broad. Lamellaecream, edge concolorous, equal, 8-10,
broad, collariate. Stipe dark brown with paler apex, hair-like,smooth, 4-6 mm long,
Spores 6-7 X 2-2.5 p, elliptical. Hymenium: cystidia none; cheilocystidia similar
to epicuticularelements. Hyphae of trama inamyloid. Coveringlayer of pileus: epi-
cutis formed by subglobosecells 10-17 y, in diameter, with thin brown walls bearingon
their outer surface numerousminute hemisphericalprocesses up to 1 ,projecting.
On fallen, slender twigs.
MATERIALSTUDIED. VENEZUELA. Dto. Federal,CaracasBotanicalGarden,950 m
alt, 19 Jun 1958, Dennis 1068 (K), type.
Singer(1965) pl 7, fig 20, Dennis (1961), fig 52.
58. MarasmiusdodecaphyllusSinger,Sydowia 18: 335. 1965.
TYPE.Singer B 2151, from Bolivia.
Pileus ochraceous-avellaneouswhen fresh, helvolous (much like dried pilei of
M. helvolus) when dried, with a central, deep gray flat dot, the dot remainingdeep
gray and flat in dried material,around it somewhat rivulose under a good lens, other-
wise quite glabrous,sulcate or slightly sulcate in marginalhalf, not pallid or white
around central dot, convex with depressedcenter, about 9 mm broad. Lamellaewhite
with white edges, buff when dried, moderately broad, 12 through-lamellaeand 2 lam-
ellulae, collariate. Stipe umber, deep fuscous when dried, shining, smooth, insititious,
15 X 0.2 mm.
Spores (4-)7.5-8.2 X 3-4.5 p, but perhapsmore variable(few seen in type). smooth,
hyaline, ellipsoid, inamyloid. Hymenium: basidia 23-24 X 7.5 ,: cystidia none: chei-
locystidialike epicuticularelements but on an averagesmallerand hyaline to subhyaline.
Hyphae of pileus and hymenophoraltrama hyaline, with clamp connections, although
some hyphae clampless, inamyloid. Cortical layers: epicutis of pileus hymeniform, its
elements balloon-shaped,of the Rotalis-type, hyaline, subhyaline or pape melleous with
pale melleous to brown, more rarely subhyaline divergentsetulae, main body 19-27 X 22
;A,setulae 1-1.3 , long.
On a fallen piece of rotting dicotyledonouswood in tropicalrain forest.
MATERIALSTUDIED. BOLIVIA.Pando: Manuripi,Santa Rosa on Rio Madrede Dios, 22
Mar1956, SingerB 2151 (LIL), type.
This species differs from M. nanuiripiensisin a different (deeper) color, also being
differently colored in the herbarium,furthermorein a largernumber of lamellae, flat
central dark spot, and large diameter of the pileus.
Among the Europeanspecies, this comes close to M. wettsteinii (a species often
considered conspecific with Ml.bulliardii)on coniferous needles which has 10-16 lame-
llae but smallerpileus and longer stipe M wettsteinzii is constantly accompaniedby
blackish, thin rhizomorphs.
ILLUSTRATION. Singer (1965) pl 7, fig 21.

59. MarasmiusplatyspermusSinger in Singer & Digilio, Lilloa 25: 215. 1952.

59a. Marasmiusplatyspermusforma platyspermus

TYPE.Singer T 203, from Argentina.
Pileus at first whitish with gray-brownto dark gray central dot, soon becoming
"Yellow beige" (13-H-7) to "wigwam"(14-G-10) or "bure"(13-H-8), dried about "but-
terscotch" (14-F-9 M&P)generally remainingwhitish in the umbilicus around the dark
Marasmius 111

dot and sometimes between elevations of sulcate margin,with or without a central

papilla coinciding with the dark dot, convex, 2-12 mm broad. Lamellaewhite, even-
tually avellaneouspallid, distant or subdistant, 13-20, all equal, broad, sometimes dis-
tinctly anastomosingin old large caps, collariate. Spore print white. Stipe black with
the apex at first white, glabrous,smooth, somewhat shining, setose, sometimes slightly
attenuate toward the apex, insititious on the substratumbut sometimes observedto
arise from a rhizomorph,12-75 X 0.2-0.5 mm, accompaniedby dark rhizomorphs,
sterile stipes (lelopodia) up to 175 mm long. Context very thin, white, inodorous.
Spores 7-10 X 4.3-5.7 1, ellipsoid, smooth, hyaline, wall inamyloid, thin to
moderately thin with oblique, thin, hilar appendage. Hymenium: basidia 22-32.5 X
6.5-8.3 ,, clavate, hyaline, 4-spored;cystidia none seen; cheilocystidia like hyaline
epicuticularelements. Hyphae of pileus occasionally weakly and slowly pseudoamy-
loid or inamyloid, those of hymenophoraltrama inamyloid, those of white apex of
stipe weakly pseudoamyloid, all with clamp connections, with thin to moderately
thickened walls, hyaline except in black part of stipe. Cortical layers: epicutis of
pileus hymeniform or subhymeniform,consisting of elements of the Rotalis-type but
in many of them the setulae of the sides of the cells obliquely ascendingand on the
tip of the cells more elongate than on the sides, main body 13-39 X 9-13 u, clavate
to vesiculose or subglobose, often pedicellate, hyaline or upper portion melleous,
setulae mostly melleous, at places hyaline, 1.5-2.2 .llong, on tip of cells sometimes
On trunks, logs, fallen branches,roots, rotting in subtropicalforest and less humid
montane woods, also in the montane Alnetum, lower zone, gregarious.Known hosts:
Boelznleria,Schilus, Allophylus. Altitudinal range: 800-1600 m.
MATERIALSTUDIED. ARGENTINA. Tucuman:Sierrade San Javier,ParqueAconquija,
12 Mar1949, Singer T 203 (LIL) type; 5 Mar1949, Sinzger& Digilio T 195 (LIL) paratype;24
Mar1949, Singer259 (LIL); Anta Muerta,24 Apr 1949, Singer & Digilio T 501 (LIL); 26 Dec
1951, Singer T 1669 (LIL); paratype;CiudadUniversitaria,20 Mar1951, Singer T 1408 (LIL);
Mar1957; Singer T 3049 (LIL);San Agustfn,1380 m alt, 15 Jan 1957, Singer T 2902 (LIL);Cat-
amarca:Andalgala,above EstanciaSanta Rosa (Suncho), 18 Jan 1952, Singer T 1 755 (LIL).
Singer(1958a) fig 40.

59b. Marasmiusplatyspermusforma scandens Singer,forma nov

TYPE. Singer T 3591, from Argentina.
Pileo pallidioreet habitatione in arboreviva differt. Typus in LIL conservatur.
Pileus white, then pale avellaneoustan in age and when dried, with a flat central
black dot in a depression,otherwise convex, grooved-sulcate,glabrous, 5-9 mm broad.
Lamellaewhite, broad, with the edge concolorous with the sides, without lamellulae,
19-21, widely collariate. Stipe at first white, later becoming black from the base up-
wards, glabrous,shining, smooth, equal or slightly taperingupwards,insititious on the
substratum,up to 16 mm long and up to 0.7 mm thick; black rhizomorphs,none seen.
Context inodorous.
Spores 7.2-9 X 4-5.3 uA,hyaline, smooth, ellipsoid, with suprahilardepressionor
applanation,thin-walled. Hymenium: cheilocystidia like epicuticularbroom cells but
all hyaline, with shorter(1.2-1.8 ,) setulae, 15-27 X 9-15 ,. Hyphae of pileus thin-
walled, filamentous, with clamp connections. Coveringlayers: epicutis of pileus hy-
meniform consisting of broom cell elements of the Rotalis type, in the upper portion
of the main body and the setulae melleous, the latter cylindric to spinulose and 2-2.5
,Along,sometimes longer near the tip of the broom cell, shorter below, with ratherthin
112 Flora Neotropica

wall in the main body, with hyaline pedicel, main body about 14-26 X 8.5-16.5m.
Growing on the bark of Duranta serratifolia,living tree, about 3 m above the
ground, gregariousbut not cespitose.
MATERIALSTUDIED. ARGENTINA. Tucuman:DepartanentoTaff, CiudadUniversit-
aria, Sierrade San Javierat 1200 malt. 24 Mar1960, Singer T 3591 (LIL), type.
This seems to be a habitat form with somewhat paler pileus and very slightly
smallerspores, rare. It should not be confused with AL.leucorotalis which differs in
having fewer lamellae and narrowerspores.
This commonspecies of the ArgentineNorthwest is confined to the lower range
of the mountains. The precedingspecies (no. 58) differs in less numerousand there-
fore more distant lamellae, slightly different colors, generallyshorter and thinner stipe
and more consistently acanthophysoid(equally and always divergentlyshort setuloid)
epicuticularelements. The rhizomorphsare either less or not developed in M. dode-
Among the holarctic species, MI.wettsteinii differs in growing on conifer needles,
slightly smaller,especially narrowerspores and a slight differencein colors. M bul-
liardiidiffers in growing on frondose leaves, fewer lamellae and even narrowerspores.

60. MarasmiusnebularumSinger, Sydowia 18: 350. 1965. Fig 49.

TYPE. Singer B 3637, from Colombia.
Pileus dark-ferruginous(between "sierra"and "caldera"M&P),cocoa brown when
dried ("cocoa" M&P),uniformly colored, glabrous,cylindrical, umbilicate, with a small
papilla in the umbilicus, 4 mm broad, 3 mm high. Lamellaewhite with white edge,
subdistant(16 through-lamellae,no lamellulae), broad, white-collariate. Stipe chestnut
with white apex, amooth, glabrous, insititious, 7-0.2 mm. Context thin, inodorous.
Spores 6.2-7.5 X 3.2-3.7 ,, ellipsoid or oblong (Q = + 2), smooth, hyaline, in-
amyloid. Hymenium: basidia clavate, basidioles fusoid; cystidia none; cheilocystidia
hyaline, otherwise like the epicuticularbroom cells. Hyphae of the pileus-tramahy-
aline, with clamp connections, inamyloid. Cortical layer: epicutis of the pileus hymen-
iform, broom cells of the Rotalis-type, about 16 X 14 ,, vesiculose or subvesiculose,
hyaline or more frequently rusty-brown,setulae 1.2-3.7 X 1-1.5 p.
On small woody twigs in fog forest, Colombia.
MaTERIALSTUDIED. COLOMBIA.Boyaca,between RamiriquiandRond6n, 30 Jul 1960,
SingerB 3637 (BAFC), type.

61. MarasmiusaequatorialisSinger, sp nov

TYPE.Singer B 7538, from Ecuador.
Pileo laete ferrugineo-aurantiaco vel aurantiaco,pallescente, umbilico albo mac-
ulo atriore praedito;lamellis albis, demum cremeis, acie haud discoloratis, subconfertis,
collariatis;stipite nigro. Sporis 5.5-6.5 X 4-4.5 p; hyphis tramatispilei inamyloideis,
frequenterlatissimis, fibulatis;elementis epicuticularibustypi Marasmiirotalis, hyalinis,
succineo-setulosis. Ad ramos radicesqueLeguminosarumin regione andina Aequatoriae.
Typus in F conservatusest.
Pileus bright rusty orange or orange when fresh, sometimes fading to whitish, the
low and flat-bottomed umbilicus white with a central darkerdot, this however rarely
in form of an extremely minute papilla and the umbilicusarea brown-scurfy-cracked
on white context ground, the orange portion becoming brown and paler striped on dry-
ing, sulcate, glabrous,convex and very steep towards margin, 5-12 mm broad. Lam-
ellae white, finally cream, with pallid edge, broad, subclose (20-22 through-lamellae)
Marasmius 113

entire, collariate. Stipe black, glabrous,setose, insititious, 7-17 X 0.2-0.3 mm. Context
very thin, white in the pileus, inodorous.
Spores 5.5-6.5 X 4-4.5 p, ellipsoid, smooth, hyaline, inamyloid. Hymenium:
basidia 17-23 X 5-6.7 1, 4-spored;cystidia none; cheilocystidia in mature specimens
mostly smooth, vesiculose, 21-30 X 8.5-18 j, thin-walled,hyaline, some, a minority
broom cells, these with a main body as above and apical diverginghyaline setulae,
1.5-2.5 s long. Hyphae of the pileus trama hyaline, with clamp connections, inamy-
loid, often very broad (3-19 p diameter in mature specimens);thin- to firm-walled;hy-
phac of the stipe dark pigmented, parallelwith each other, vaguely pseudoamyloid.
Cortical layer: epicutis of the pileus hymeniform, consisting of broom cells of the
Rotalis-type, main body 10-35 X 8-19 , often (the longest) long-pedicellate,thin-
walled. hlyaline,setulae succineous-brown,rodshaped, 1-2.2 p long, divergingand
coveringa wide area of the cell, smooth cells very scattered or absent.
On dead standing branchesor roots of leguminousshrubsand trees in the Andean
re,ionl, gregarious.
MATERIALSTUDIED. ECUADOR.Pichincha,Rio Machaingaro,
2600 m alt, 20 May 1973,
Singer B 7538 (1), type.
This differs fronmM. icnbularumt in the not uniformly colored pileus, broaderyet
shorter spores and, as from mlost species in this group, in the fact that the fully mature
specimenllshows a large majority of smooth cheilocystidia.

62. MarasmiusnothomyrciaeSinger, Arkiv Botanik 4: 384. 1959.

TYPI:.SparreF 56, from Juan Fernandez(Chile).
Pileus cinnamon to topaz color (12 C 8 to "topaz" M&P)with the zone around
the center orange ("paloma" M&P),dried light orange cinnamon to tawny (between
"tawny" and "cookie" M&P)or between "Sayal Br." and "Butterscotch"M&P(brown),
either whitish or "topaz" when fresh in the umbilicus(there under a lens scurfy show-
ing the white pileus-trama),dried here deep-grayto buff-grayor whitish to light cin-
inamonto tawny, in the middle of the umbilicus with a central dot which is at first
(fresh) conlcolorous with the rest of the umbilicus, later more gray to grayish black,
dried grayisshblack to black, glabrous,excepting the scurfy zone, where present, sul-
cate except in the umnbilicus, half-barrel-shaped,then strongly convex to convex, al-
ways abruptly umbilicate, 2-7 mm broad. Lamellaewhite with weakly to decididly
discolored("paloma" M&P)edge and collarium, the discolorationnot always complete
and ofteii visible only with a lens in dried condition, ratherdistant (11-15 through-
lamellae, no lamellulae)broad, widely and distinctly collariate. Stipe chestnut-brown-
black to black, shining, with white apex as long as the central dot of the pileus has
not become gray or black, later uniformly black, glabrous, almost constantly accompan-
ied by (but not attached to) numerourblack rhizomorphs,insititious, 15-50 X 0.1-0.3
mm, often remiainingsterile ("telepodia"). Context very thin in the pileus, inodorous.
Spores (6-)6.5-10 X 3.5-4.5(-5) p, fusoid, oblong-fusoidor ellipsoid, smooth, hy-
aline, inamyloid. Hymenium: basidiaabout 23 X 6.5 p, 4-spored;basidioles fusoid;
cystidia lnone:cheilocystidia like epicuticularbroom cells but setulae either golden to
golden hyaline or hyaline. Hyphae of the pileus-tramahyaline, thin-walled,with clamp
connections, moderately but rapidly pseudoamyloid. Cortical layer: epicutis of the
pileus hymeniforim,broom cells of the Rotalis type, 10-23.5 X 7.5-17.5 p, hyaline but
with deep golden melleous to tawny-ferruginoussetulae covering the apex or the upper
114 Flora Neotropica

two thirds of the cell, setulae 1.2-2.7 X 0.9-1.3 ji, cylindric or verruculose,obtuse or
On leaves fallen to the ground, rarely on dead wood rotting nearby or on stems
and small pieces of woody vines, on Dicotyledones, rarely on dung. Known hosts: Notho-
myrcia, Asara, Aextoxicum.
MATERIAL STUDIED.CHILE.JuanFernandez: pathto El Camote,about400
m alt, 18 Mar1955, SparreF 56 (S) type; Coquimbo:Fray Jorge, 16 Apr 1967, SingerM 6515
(SGO). As for CentralandSouthChileancollectionsstudiedsee Singer(1969, p. 103).
This species is somewhat marginalas far as the neotropics are concerned since
the only locality known in the subtropicalbelt is a fog forest in North Chile.

63. Marasmiushylaeicola Singer, sp nov Fig 50.

TYPE.Singer B 7298, from Ecuador.
Pileo brunneo, zona alba vel pallidiorecircum maculum centralem nigrumornanto,
3-5 mm lato; lamellis albis, collariatis;stipite brunneolo apice albo vel pallido; rhizomor-
phis nullis visis. Sporis 5.5-7.5 X 2-3.5,; elementis epicuticularibustypi Marasmiirot-
alis. Ad folia delapsa arborumdicotyledonearumin hylaea. Typus in F conservatusest.
Pileus brown, "gold pheasant" to "sorrel"or between "copper br." and "Mohawk"
(M&P)when fresh, with white (because of cracking)or paler, narrow or ratherwide
zone around a black or blackish dot, dried becoming "kis kilim" to "cattail" (M&P)in
marginalzone and black or gray in center with the pallid zone remaining+ distinct, sul-
cate, glabrous,convex, umbilicate, with a flat bottom or dark dot slightly elevated, 3-5
mm broad. Lamellaewhite, with white edge or with an inconsistent and very narrow
discoloration on the edge (lens!) which is concolorous with the marginof the pileus,
eventually or on drying tending to become wood brown all over, subdistant to distant
(15-17 through lamellae, no lamellulae), broad, collariate. Stipe brownish, almost sepia
with white or pallid apex, smooth, glabrous,insititious, 26-42 X 0.1-0.2 mm; no black
rhizomorphsseen. Context thin, inodorous.
Spores 5.5-7.5 X 2-3.5 p. ellipsoid to oblong, smooth, hyaline, inamyloid. Hymen-
ium: basidia 15-18 X 4.5-6.3 u, 4-spored;basidioles fusoid; cystidia none; cheilocystidia
like epicuticularbroom cells but many or all entirely hyaline, often scarce. Hyphae of
the pileus-tramahyaline, weakly to distinctly pseudoamyloid, with clamp connections,
strongly pseudoamyloid and parallel in the apex of the stipe. Corticallayer: epicutis
of the pileus hymeniform, broom cells 9-30(-40) X 6.8-20(-27) , of the Rotalis-type,
main body brownish hyaline to hyaline, some obliquely pedicellate; setulae divergent,
1.2-2.7 X 0.6-1.2 p, rod-shaped,obtuse, mostly brownish to deep chestnut brown.
On dead fallen leaves of dicotyledonous trees in virgin tropical rain forest.
MATERIALSTUDIED. ECUADOR. Napo. Lago Agrio, 8 May 1973, SingerB 7298 (F),
type. BOLIVIA.Beni,VacaDiez, 3 km belowGuayaramerin
12 Mar1956,SingerB 1873 (BAFC);
B 1874 (BAFC).
Marasmiushylaeicola is characterizedby its often narrow,pallid zone around a
dark dot, the basically brown color of the pileus (from the beginning), the small spores
and the foliicolous habitat. It differs from Maramiusletcorotalis in its color and dis-
tribution and from Marasmiusminimus Dennis in the presence of a pallid zone and its
habitat on dead leaves ratherthan twigs, more numerouslamellae and largersize. It
also differs from all forms of the Marasmiusvariabilicepscomplex in having smaller
ILLUSTRATION. Singer (1965) pl. 7, fig 22.
Marasmius 115

64. Marasmiusvigintifolius Singer, Sydowia 18: 345. 1965.

TYPE.Singer B 871, from Bolivia.
Pileus "tuscan tan, sauterne"(ochraceous tan) or between "Cognac"and "russet
br." or 15-J-12, 13-J-10 to 14-G-1 ("tortoise shell") (M&P),i e various shades of
brown to fuscous without a red tinge, sometimes "clove" (fuscous) and gradually
"suntan"(light sordid tan) to extreme marginwhen fresh, in dried condition between
"kis kilim" and "burnt umber" or "clove" to 8-J-12, (M&P)not deep rusty chestnut
color, in fresh and dried condition mature caps with a narrowto broad and conspicu-
ous white zone around the central dark dot which is black to blackish but may become
concolorous with marginin dried material, glabrous,sulcate over one half to two thirds
of the radius, campanulateto truncate-conicalwhen young, mostly becoming convex
when mature, with a ratherbroad and deep umbilicus, with or without a minute umbo
in the umbilicus, 2-10 mm broad. Lamellaewhite, edges and collariumusually distinc-
tly brown or at least partiallyso, 17-22, mostly 20, subclose to subdistant,rather broad
to broad (1.5 mm), collariumnarrowor wide. Stipe umber black to black with initially
white apex, dried fuliginous or deep chestnut to nearly black, shining,glabrousand
smooth, insititious, setose, or very slightly taperingtowards apex, 14-56 X 0.1-0.3 mm;
no rhizomorphsor with scanty dark rhizomorphsunconnected with base of stipe. Con-
text thin, inodorous.
Spores 7.2-10 X 4.5-5 g, mostly 8.2-9 X 4.2-4.5 ,, ellipsoid, smooth, hyaline,
inamyloid. Hymenium:basidia 16-26 X 5-6.3-9 2, 4-spored;cystidioles rare, broad
vesiculose or vesiculose-clavate,hyaline, smooth, e g 8-9 p in diameterwhich are a
special type of basidiole;true pleurocystidianone; cheilocystidialike epicuticular
broom cells. Hyphae hyaline, filamentous, slowly and weakly pseudoamyloidin trama
of pileus. Corticallayers: epicutis of pileus hymeniform or subhymeniform,its elemen-
ts vesiculose or vesiculose-pedicellate,sometimes in adult specimenshorizontally exten-
ded, main body 13.5-20 X 19-23.5 A,very few almost hyphous, of the Rotalis-type,
hyaline, beset with divergingshort, rod-like setulae which are melleous brownish, deep
ochraceousbrown or deep sepia, 2-27 X 1-1.3,.
On blade and petioles of fallen dicotyledonous leaves in montane tropicalforest.
MATERIALSTUDIED. BOLIVIA. La Paz: Nor-Yungas,Coroico,4 Feb 1956, SingerB 871
(LIL), type; B 873 (LIL), paratype;CarmenPampa1 Feb 1956, B 800 (LIL), paratype;17 Feb 1956,
B 1245 (LIL), paratype;19 Feb 1956, B 1956 (LIL), paratype. BRAZIL. Guanabara,Florestade
Tijuca 12 Oct 1961, SingerB 4006 (BAFC).
Singer(1965) pl 8, fig 23.

65. Marasmiusidroboi Singer, sp nov

TYPE. SingerB 6925, from Colombia.
Pileo brunneo, zona alba vel alutaceo-pallidacircum maculambrunneamvel fus-
cum centralempraedito, 4-8 mm lato; lamellis albis, collario pallide coriicolori praeditis,
subconfertis;stipite nigro; rhizomorphisnullis. Sporis 6.5-8 X 2.8-3.8 p; hyphis non-
nullis tramatispilei pseudoamyloideis,fibulatis;elementis epicuticularibus typi Marasmii
rotalis. Ad folia dicotyledonea delapsain Columbia. Typus in F conservatusest.
Pileus brown, fresh ("oak M&P)and dried ("Cochin"M&P)with paler radial
stripes at least when fresh, with a white to buff-pallidzone around a central brown to
fuscous dot which bleaches to buff-pallidor gray when dried, sulcate, convex with
steeply declivous marginand abrupt umbilicus which is flattened on the ground, 4-8
mm broad and up to 6 mm high. Lamellaewhite with pale tan collariumand some-
times also the edges finely and weakly discolored brown to pale tan under a lens, sub-
116 FloraNeotropica

close (20-23 through-lamellae,no lamellulae),broad, distinctly,collariate. Stipe black

(perhapsinitially with white apex), glabrous,insititious, not accompaniedby rhizo-
morphs, 22-25 X 0.1-0.2 mm. Context thin, inodorous.
Spores 6.5-8 X 2.8-3.8,, ellipsoid to slightly oblong, smooth, hyaline, inamyloid.
Hymenium:basidia 16-19 X 5-6.5 M,4-spored;basidiolesfusoid; cystidia none; cheil-
ocystidia like the epicuticularbroom cells, those on the very edge more or less pigmen-
ted with some hyaline one intermixed, those accompanyingthem further towards the
side of the lamellae, all hyaline. Hyphae of the pileus-tramahyaline, thin-walled,some
thick-walled,some distinctly pseudoamyloid,some weakly pseudoamyloid,all with clamp
connections; some cleiferoushyphae present. Corticallayers: epicutis of the pileus
hymeniform,broom cells of the Rotalis type, main body either entirely hyaline or hy-
aline in the lower part and brownish in the upper, 17-36 X 8-22 ;, vesiculose, setulae
1.3-2.5 X 0.3-1.5,, verruculoseor rod-shaped,mostly brown, fewer subhyaline.
On dead fallen dicotyledonous (ericaceous and rubiaceous)leaves. Colombia.
MATERIALSTUDIED. COLOMBIA.Boyaca:Tunja,Arcubuco,Finca de Luis Antonio
Porras,29 Jul 1960, SingerB 3599 (BAFC);Valle, Cali, CerroHorqueta,2000 m alt, 2 May 1968,
SingerB 6925 (F), type.
This species is closely related to Marasmiusvigintifolius but differs in the narrow-
er spores and the radiatelystriped pileus.

66. MarasmiuseucladopusSinger, Sydowia 18: 336. 1965.

TYPE. Singer,B 1478, from Bolivia.
Pileus somewhat more ochraceousthan "copper br." M&P(rusty), dried turning
deep red-brown("Mandalay"M&P),dried with a pallid center, marginedby a wall, with
an insignificantumbo in the pallid zone but umbo not black at least in dried material,
sulcate on the large, colored, declivous portion, convex to truncate-conicalwith a cen-
tral depressionin which the umbo is inserted, 1-1.2 mm broad, 0.8-1 mm high. Lamel-
lae whitish, edge concolorous with the sides or with the pileus, with brown margined
collarium,at least in dreid materialcollariumwide and distinct, distant (9-10), very
broad. Stipe light dirty umber, smooth and glabrouswith white, thinner apex, inserted
without any ring or rings into a central common rhizomorphicstipe-like strand, indiv-
idual stipe 5-11 X 0.1 mm; stipe-bearingrhizomorphsblack, above more dirty umber,
smooth and glabrous,producingseveralcarpophores,insititious.
Spores 8.8-9 X 5.8 p, ellipsoid, smooth, hyaline (few seen). Hymenium:basidia
16-17 X 8 ,; cystidia none; cheilocystidialike epicuticularelements but mostly hyaline.
Hyphae of pileus-tramahyaline, practicallyinamyloid, yellowish even after 30 minutes
exposure to Melzer's,reagent,with clamp connections. Corticallayers: epicutis of pil-
eus hymeniform, its elements of the Rotalis-type, the majority covered down to the
middle by divergingshort setulae, or reminiscentof the Siccus-type(with the setulae
although short and divergingall concentrated on the apical portion of the cells main
body 16-27 X 8.8-20p , clavate,vesiculose, even globose, setulae brown, 1.3-4.2 X 1.2-
1.8 ,.
On dead dicotyledonous leaves fallen to the ground in montane tropical forest.
MATERIALSTUDIED. BOLIVIA. La Paz: Nor-Yungas,CarmenPampa,26 Feb 1956, Singer,
B 1478 (LIL), type.
The idea that this might be M. populiformis, not rediscoveredsince the time of
Spruce, is very tempting. However, the color of the stipe, the size of the fruiting bodies,
the color of the pileus, the habitat, the spores as indicated by Dennis (195 lb) are all
ILLUSTRATION.Singer (1965) pl 8, fig 27.
Marasmius 117

67. MarasmiustetrachrousSinger Sydowia 18: 343. 1965 (p 171 ex errore tetrachroinus).

TYPE.Singer B 1528, from Bolivia.
Pileus young ferruginous(between "feuille morte" and "rust sorolla" M&P),then
fresh "Windsor tan" or between "rust sorolla" and "Arab"or "Alamo" to "Arab"(still
rusty brown) and eventually sometimes buffy alutaceous (1 -F-7 M&P),dried deep ferru-
ginous brown to ferruginouschestnut e g "chutney," fresh showing a rather narrowwhite
ring around a central black dot but this latter usually disappearingon drying and not
always clearly marked,very young caps with a prominent whitish or sordid pallid to
blackish papilla, later white zone with a very low umbo or flat, sulcate in colored zone,
campanulateor conical, then almost cylindrical-drumshaped but with narrowerdisc which
is truncate at the tip or even slightly depressed, 1.5-2 mm broad and as high. Lamellae
white with white edges and collarium, about nine, very broad, collariate, mostly all equal,
very rarely with very few lamellulae. Stipe blackish brown to chestnut brown, smooth and
glabrous, shining, setose, insititious, 9-15 X 0.1 mm: no rhizmorphsobserved.
Spores 7-10.5 X 3.5-5(-6.5), mostly 9.5-10.5 X ? 5p, ellipsoid, smooth, hyaline, in-
amyloid. Hymenium: basidia broad and short, 16.5-19.5 X 10-11u cystidia none; cheilo-
cystidia of the Rotalis-type, vesiculose, 16-17p broad, hyaline. Hyphae of the pileus
tramaweakly pseudoamyloid. Coveringlayers: epicutis of the pileus hymeniform, with
elements of the Rotalis-type, these 20-27 X 18-19l, hyaline to pale fuscous, setulae rod-
shaped, ochraceous brown, 2.5-4 X 1.3p.
On fallen dicotyledonous leaves in montane tropical forest, and on leaf petioles,
at about 2000 m altitude.
MATERIALSTUDIED. BOLIVIA. La Paz: Nor Yungas,CarmenPampa,26 Feb 1956,
SingerB 1528 (LIL) type: B 1485 (LIL) paratype:B 1530 (LIL) paratype.
The rhizomorphs,if at all formed, are scanty and inconspicuous. Their lack of
prominence, the small size of the carpophores,the high pileus, the small number of
lamellae with their white edges and collariumand the medium-sizedellipsoid spores are
ILLUSTRATION. Singer (1965) pl 8, fig 29.

68. Marasmiustereticeps Singer, Sydowia 18: 343.1965.

TYPE. Singer B 2299, from Bolivia.
Pileus cinnamon ochraceous, not becoming deep ferruginouschestnut when dried,
the upper flat portion white with a darkerdot in the middle of the white field which is
surroundedby a wall-like convex zone concolorous with the vertical marginalzone, cen-
tral dark dot becoming rather indistinct on drying, cylindric-drum-shaped and terete,
sulcate over vertical sides, about as high or higher than broad, 2 mm in diameter. Lam-
ellae white, dried buff, extraordinarilybroad (broader than the radius of the pileus),
twelve, not intermixed, simple, distinctly collariate. Stipe blackish with white apex,
dried sordid light umber much as in M. pallipes, glabrousshining, with a minute pallid
disc at the insititious base, setose, 10-11 X 0.1 mm; no rhizomorphs.
Spores 8.2-11.2 X 3.5-4,, cylindric to ellipsoid-oblong,hyaline, smooth, inamyloid.
Hymenium: basidia 19-22 X 7.5;; basidioles fusoid, often with mucro; cystidia none;
cheilocystidia like epicuticularelements but with hyaline or subhyaline setulae. Hyphae:
in pileus and hymenophoraltrama regularbut very thin, hyphae hyaiine, inamyloid, with
clamp connections. Cortical layer: epicutis of pileus hymeniform, of Rotalis-type ele-
ments with the main body 18-30 X 12-21u, obpiriform,ovoid, clavate, vesiculose etc,
hyaline with melleous to brown, short, obtuse setulae of 2 X 1.3 , (so on the vertical
outside of the pileus).
118 FloraNeotropica

On dicotyledonous leaves fallen to the ground in tropical rain forest.

MATERIAL STUDIED.BOLIVIA.Pando:Madrede Dios,SantaTeresa,28 Mar1956,
SingerB 2299 (LIL)type.
ILLUSTRATION. Singer (1965) pl 8, fig 28.

69. MarasmiusvariabilicepsSinger, Sydowia 18: 344. 1965.

69a. Marasmiusvariabilicepsvar variabiliceps

TYPE.Singer B 1531, from Bolivia.
Pileus varyingbetween the color of M. platyspermusand Al. vigintifbliuswhen fresh
but on partial dehydration becoming deep rust brown, fresh e g "russet br," partially
dried between "Rust sorolla" and "Arab" and in the herbariumbetween "chutney" and
"Java"M&P(red-brownto brown-red,a deep shade), with a ratherbroad white zone
around a central dark dot which is black or blackish when fresh and gray to palest sor-
did brown when dried, glabrous,sulcate, convex, with a central depressionand not or
scarcely umbonate in the depression5-9 mm broad. Lamellaepallid to white, with brown
collariumwith white edges, broad, moderately distant to subclose, 20 or around 20.
Stipe black with white apex, dried deep chestnut to chestnut black, glabrous and smooth
slightly taperingupwardsor equal, setose, insititious, 58-100 X 0.2-0.4 mm; rhizomorphs
scanty, inconspicuous, independent of stipe.
Spores rathervariablein shape and size, 8.3-11.5 X 3.5-5.5,, often relatively broad
ellipsoid, hyaline, smooth. Hymenium: basidiabroad as in i1. vigintijolius;cystidia none;
cheilocystidia like epicuticularelements but all hyaline. Cortical layer: epicutis of pil-
eus hymeniform or subhymeniform,of the Rotalis-type, very few epicuticularelements
approachingsomewhat the Siccus type, 24-29 X 18p, beset with ochraceous ferruginous
to brown setulae which are rod-shaped,3-4.8 X 1.3-1.5~,.
On dead fallen dicotyledonous leaves in montane tropical forest.
MATERIAL STUDIED.BOLIVIA.LaPaz:Nor-Yungas, CarmenPampa,26 Feb 1956,
SingerB 1531 (LIL),type;1 Feb 1956,B 800a (LIL),paratype.
This species is very close to M. vigiztifblius from which it differs mainly in the
color of the dried pileus and the longer setulae of the epicuticularelements, also the
longer stipe and more variablespores.
ILLUSTRATION. Singer (1965) pl 8, fig 24.
There is a number of forms, all characteristicof the Yungas region, which are dif-
ficult to separatefrom the type form as independent species and which appear to be de-
pauperateforms, yet by their spore and lamellae charactersvaryingwithin certain defin-
ite limits they are apparentlyhereditaryraces. They are neither geographicalraces nor
mere forms. However, I have listed them as "varieties"(Singer 1941). It will not be
possible to determine the correct taxon applicableas long as the infertility between these
forms has not been confirmed experimentally. It may be that we deal here with sub-
stratum adaptations(mycoecotypes), but it is prematureto guess because it was impos-
sible to determine the leaves on which every form occurred inasmuchas the tropical
montane forest of the Yungas is immensely rich in species and most host species were
neither flowering nor fruiting at the peak of the rainy season. One of the varietieshere
included was found in the continuation of the Yungas towards the south in Argentine
territory, in subtropicalforest.

69b. Marasmiusvariabilicepsvar mesites Singer, Sydowia 18: 344. 1965.

TYPE. Singer B 1210, from Bolivia.
Marasmius 119

Pileus rusty "feuille morte" M&Pwith marginpale cinnamon ("golden wheat"),

dried with a white zone around a black dot, dried "russet br." with "rust sorolla" M&P
center, campanulate,sulcate, 5.5 X 5.5 mm. Lamellaepale cream with broad brown-
marginedcollarium, at least in dried condition, subdistant(17) all equal. Stipe black
witli whitish apex, simply attached to a black rhisomorph,30 X 0.6 mm.
Spores 10-11.7 X 4-4.8,, mostly 10.3-11 X 4-4.2p, ellipsoid-subfusoid,smooth,
hyaline. Cortical layers: epicutis of pileus with elements of the Rotalis-type, setulae
short and ochraceous brown.
In tropical montane forest, on leaf of dicotyledonous tree.
MATERIALSTUDIED. BOLIVIA. La Paz: Nor-Yungas,Rio Yariza, 16 Feb 1956,
SingerB 1210 (LIL), type.
ILLUSTRATION.Singer (1965) p1. 8, fig. 26.

69c. Marasmiusvariabilicepsvar tucumanensisSinger, Sydowia 18: 344. 1965.

TYPE. Singer T 1290, from Argentina.
Pileus "cognac," "cookie" M&Pon marginalportion, dried between "clove" and
"Mandalay"M&P(deep brown with chestnut-ferruginoustinge) with a strikingwhite
zone around a central small blackish dot, the dot appearingpapillate gray in dried ma-
terial, glabrousand smooth in the middle, sulcate in the pigmented zone of the margin,
convex, depressed in center, 2-8 mm broad. Lamellaewhite with edge and collarium
either brown or almost concolorous with sides, 13-17, ratherdistant, broad, collariate.
Stipe black, glabrousand smooth, insititious, shining, equal, setose, 18-35 X 0.1-0.4
mm; long dark rhizomorphsassociated with carpophoresbut stipe never rising from
them. Context very thin, inodorous.
Spores 7.5-8.8 X 4-4.8,u, smooth, hyaline. Hymenium: basidia clavate, 4-spored;
basidioles fusoid; cystidia none seen; cheilocystidia somewhat like epicuticularelements.
Hyphae somewhat pseudoamylod in trama of pileus, with clamp connections. Cortical
layers: epicutis of pileus hymeniform, consisting of elements of the Rotalis-type, some
somewhat transient towards the Siccus-type or at least as in 1. platyspermus(those on
the tip of the cells tending to be longer, those on the sides somewhat oblique), main
body hyaline, setulae brownish melleous, 1.2-3.3 X 1-2,.
On leaves of Phoebe porphyria, in subtropicalforest, 1000 m altitude.
MATERIALSTUDIED. ARGENTINA. Tucuman:Chicligasta:Rio Cochuna,20 Feb 1951,
Singer T 1290 (LIL) type; Quebradade los Sosas, 1100 m alt, 12 Apr 1961, Singer T 3705 (BAFC).
ILLUSTRATION. Singer (1958a) fig 44.

69d. Marasmiusvariabilicepsvar separatusSinger, Sydowia 18: 344. 1965.

TYPE. Singer B 1547, from Bolivia.
Pileus ochraceous brown, brighter(rustier) than "tortoise" and "Hispano,"almost
"gold pheasant,"dried red-brown("Mohawk")with a whitish -zone around the black
central dot which in dried condition maintainsitself as a flat gray disc with pallid star-
like fibrils around it, sulcate in the colored zone, convex, white zone and central dot de-
pressed, 1.5-4 mm broad. Lamellaewhite, edges brown, collariumusually at least brown
spotted, rarely even edges, not distinctly discolorous, subdistant(11-14, mostly 14 in
largermature specimens) broad. Stipe blackish chestnut to black, smooth and glabrous,
shining, insititious directly on the substratum,up to 25 X 0.2 mm. Context thin, off
white, inodorous.
Spores 8.3-10.3 X 4.8-7u, ellipsoid, smooth, hyaline. Hymenium: cheilocystidia
like epicuticular broom cells, also with brownish setulae. Cortical layers: epicutis of
120 Flora Neotropica

pileus with Rotalis-type elements, setulae ochraceous brown, short, obtuse.

Otherwise like type variety.
On stem of dead vine in montane tropical forest.
MATERIALSTUDIED. BOLIVIA. La Paz: Nor-Yungas,CarmenPampa,26 Feb 1956, Sing-
er B 1547 (LIL) type.

69e. Marasmiusvariabilicepsvar heptachroinusSinger, Sydowia 18: 344. 1965.

TYPE.Singer B 1529, from Bolivia.
Pileus a saturatedorange ferruginous("Ginger,Kaiserbr. M&P")on drying be-
coming deep brown-red,ferruginous-chestnut("Java"), sulcate, white around a central
black dot which on drying becomes sordid gray, barrel-shapedand vaguely umbonate in
truncate top, 2 mm broad and high. Lamellaewhite, edges usually not discolorous
but in dried specimens at least the collariumdistinctly brown, distant (12), broad.
Stipe fuscous, glabrous, shining, insititious, 25 X 0.1-0.2 mm.
Spores 7-8.3 X 3.6-4.8u, ellipsoid, smooth, hyaline. Hymenium: basidia as short
and broad as in M. tetrachroinus;cheilocystidia like epicuticularelements but mostly
hyaline. Hyphae somewhat pseudoamyloid in trama of pileus. Cortical layers: epicuti-
cular elements of the Rotalis-type, main body e g 20 X 1l , but also larger,setulae
ochraceous brown to deep rusty ochraceous brown, obtuse, rod-like, short, but longer
than in M. tetrachrous,3-4.3 X 1.3-1.5,.
On dead fallen dicotyledonous tree leaf in montane tropical forest.
Otherwiselike type variety.
MATERIALSTUDIED. BOLIVIA. La Paz: Nor-Yungas,CarmenPampa,26 Feb 1956,
SingerB 1529 (LIL), type.
ILLUSTRATION.Singer (1965), p1 8, fig 25.

69f. Marasmiusvariabilicepsvar nothotereticeps Singer, Sydowia 18: 344. 1965.

TYPE. Singer B 1411, from Bolivia.
Pileus "Kobe" M&P(deep rufous) with depressedwhite umbilical zone and a dark
dot in the center, dried between "cocoa" and "Mohawk"(deep red brown) sulcate to
white zone, subcylindricor barrel-shaped,1.2 X 1.2 mm. Lamellaewith a brown-rufous
marginedcollariumbut with pallid edges, widely collariate, subdistant(12) broad. Stipe
much like that of A. tereticeps, much paler brown above and terete at apex when dried,
fresh with white apex, somewhat compressedbelow when dried, insititious, 12 X 0.1 mm;
no rhizomorphs.
Spores 7-9.5 X 2.7-3,, smooth, inamyloid. Hymenium: basidia 15-20 X 4.5-6p,
4-spored; cystidia, not seen; cheilocystidia like epicuticularbroom cells but more often
hyaline. Hyphae of trama of pileus very weakly pseudoamyloidat first but after about
10 minutes some strongly pseudoamyloid, all with clamp connections. Cortical layers:
epicutis of the pileus hymeniform, elements of Rotalis type, e g 22 X 16.5S, more or
less pseudoamyloid, upper portion with divergingshort (to 2 X 1j) brown setulae.
On tender dicotyledonous leaf fallen to the ground in montane tropical forest.
MATERIALSTUDIED. BOLIVIA. La Paz: Nor-Yungas,Rio Yariza,23 Feb 1956, Singer
B 1411 (LIL), type.

69g. Marasmiusvariabilicepsvar derubricansSinger, Sydowia 18: 344. 1965. Fig 51.

TYPE. Singer B 1395, from Bolivia.
Pileus ferruginous("feuille morte") to 12-B-11 M&P(light orange cinnamon) when
very young, later when quite fresh more reddish, when dehydrated in situ "Agate" M&P
Marasmius 121

(saturated deep ferruginous)or brown, dried between "tortoise shell" and "cognac" M&P
(brown without a reddish tinge) glabrous, sulcate to a narrowwhite or paler often indis-
tinct zone in umbilicus, the white zone surroundinga black spot which is flat and becomes
dark gray on drying, barrel-shapedand practicallyas high as broad, 2-4 mm broad. Lam-
ellae white with white to slightly browned (dried) edges and brown-marginedcollarium,
narrowly collariate, subclose to subdistant,(10-13), broad. Stipe black with at first
white apex, glabrous, insititious setose, often long, 15-60 X 0.1-0.3 mm; no rhizomorphs.
Spores 9-10.7 X 2.7-4,u, rarely as short as 8.5, and as broad as 4.2,. Hymenium:
cheilocystidia like epicuticularelements. Hyphae of pileus pseudoamyloid. Corticallay-
ers: epicutis of pileus with Rotalis type elements, main body 17-26 X 15-22A, vesiculose;
setulae rod-shaped,2-3.5 X Il,.
On dead dicotyledonous leaves fallen to the ground in tropical forest.
MATERIAL STUDIED. ECUADOR. Napo. Shushufindi, 15 May 1973, Singer B 7460 (F).
BOLIVIA. La Paz: Nor-Yungas, Rio Yariza, 23 Feb 1956, Singer B 1395 (LIL) type.

70. MarasmiusbaeocephalusSinger, sp nov Fig 52B.

TYPE.Singer B 7311, from Ecuador.
Pileo brunneo, albo-zonato, atro-maculato,minuto; lamellis brunneo-marginatis,
collariatis;stipite atrocastaneoapice pallido; rhizomorphisnullis. Sporis 6-8 X 3.5-4,;
elementis epicuticularibustypi Marasmiirotalis. Ad folia dicotyledonea delapsa in Aequa-
toria. Typus in F conservatusest.
Pileus brown when dry, (dried near "Sudan br." M&P)with a white or whitish
zone in the umbilicus surroundinga dark spot which is gray in dried condition, and not
or scarcely elevated, sulcate outside the umbilicus, glabrous, truncate-conic,0.7-0.8 mm
broad, 0.6-0.8 mm high. Lamellaewhite with brownish ochraceousedge and collarium
at least when dried, subdistant(9-11 through-lamellae,no lamellulae),extremely broad
in relation to the length, collariate. Stipe deep chestnut to chocolate with pallid apex,
glabrous, shining, setose, insititious, 8.5-10 X 0.1 mm; not accompaniedby rhizomorphs.
Context extremely thin in the pileus.
Spores 6-8 X 3.5-4p, ellipsoid, smooth, hyaline, inamyloid. Hymenium: basidia 15
X 6.5-7 , 4-spored;basidioles fusoid, acute; cystidia none; cheilocystidia like the epicut-
icular broom cells but all with hyaline main body and somewhat lighter colored (than in
the epicutis) setulae. Hyphae of the tramaof the pileus 2-8s, broad, subhyaline, pseudoa-
myloid, with clamp connections; hymenophoraltrama regular,thin, consisting of hyphae
1.8-3.2 P broad, rather dense and interwoven;no gelatinous zones present. Hyphae of the
stipe parallel,pseudoamyloid. Cortical layer: epicutis of the pileus hymeniform, consis-
ting of broom cells of the Rotalis-type, main body 12-18 X 7-20p,, thin-walled,hyaline
or subhyaline, vesiculose; setulae about 1-1.5, high, light gold brown or melleous, rod-
On leathery dicotyledonous leaf fallen to the ground in the hylaea.
MATERIAL STUDIED. ECUADOR: Napo, 8 May 1973, Singer B 7311 (F), type.
This is easily the smallest species of this section. Only one collection with two
carpophores has been made thus far but the fructifications are easily overlooked and the
species may be much more common.

Marasmius sect Marasmius subsect Penicillati Singer, subsect nov

Elementis epicuticularibustypi Marasmiisicci.
TYPE SPECIES. Marasmius graminum (Libert) Berkeley.
122 FloraNeotropica

Key to the Species of SubsectionPenicillati

1. Pileus whitish, with dark central dot, reaching 2 mm; spores 5.8-6.8 X 1.8-2.8(3.3), (if
spores and pileus larger, see "19" below). 71. Mf.peckii
1. Pileus differently colored or larger or spores larger.
2. Spores less than three times as long as broad, ellipsoid, fusoid, ovoid or oblong.
3. Pileus white when fresh (sometimes becoming pale umber or pale argillaceous
when drying or dried), with or without a dark central dot or papilla.
4. Epicuticular broom cells with few shallow and always quite obtuse lobes, the
upper portion of the cell wall pseudoamyloid; central dark spot umbonate,
pileus reaching 2 mm in diameter; spores 8-9(-10) X (3-)3.5-4.2 u; on fallen
coriaceous leaves, cespitose. 72. M.1chr?sochaetes
4. Epicuticular broom cells with narrower, less obtuse and generally more numer-
ous setulae at least in the marginal region of the pileus (outside the umbilicus),
setulae inamyloid, more rarely pseudoamyloid; central dark spot or umbo
present or absent; spores as above or different; on leaves or wood.
5. Pileus without a central pigmented spot in both fresh and dried condition;
spores 8-11 X 4.5-5.5,; on bamboo. 73. M. aspilocephalts.
5. Pileus either in fresh or in dried condition with a central dark spot or a
small papilla.
6. Central pigmented spot flat, dot-like or lamellae 12-16, see "11" below.
6. Central pigmented spot covering a prominent papilla; lamellae about
7. Many stipes rising from black rhizomorphs,others directly from the
substratum (wood); rhizomorphs often very long and often ramose.
abundant and conspicuous; pileus 1-2 mm broad. 74. A. pallenticeps.
7. All stipes rise from the dead monocotyledonous or dicotyledonous
leaves which are the constant habitat of this widely distributed
species of the tropical lowlands: rhizomorphs few and thin or none;
pileus 1-6 mm broad. 75. M. conicopapillatus.
3. Pileus well pigmented even outside the central dark spot (if this exists).
8. Pileus gray (violet gray, blackish gray, cinereous. smoke-gray) not clay color,
brown, rufescent to ferruginous or orange and not purple or red.
9. Spores more than 8 p long, pileus mostly 4 mm broad or broader.
10. On monocotyledonous trash; spores 7.2-10.5 X 3.7-5.5t,. 76. M1.nigrobninneus.
10. On dicotyledonous litter; spores 10.2-12.3 X 4.5-6u.s, 77. M. fildigineorotula.
(Note: If spores 11-14 X 3.5-4p. compare Pararotulae: M,. rimanus).
9. Spores less than 8ulong; pileus smaller than 4 mm (colnpare also "11"
below). 78. AI.magnisetulosu
8. Pileus not gray.
11. Pileus in no part or zone orange rusty, orange-flame-color. orange yellow
or salmon orange nor with purple red, blood red. brown-red or pink tinge
unless the pigment is between brown and rusty ochraceous or ocher to
argillaceous and becomes rufescent or deep chestnut ferruginous on dry-
ing (take alternative only if pileus is somewhat red or orange or rufous in
fresh condition).
12. Many or all stipes arising from a dark rhizomorph (if the latter is
hirsute, see no. 80).
13. All epicuticular elements thick-walled and strongly pseudoamyloid
with coarse, obtuse lobed diverticula which are often scarce, be-
tween cinnamon and vinaceous brown but intermixed with hyaline
cells; pileus at first light brown then cinnamon-fuscous; lamellae
seven; spores 7.5-9 X 4-4.5 u. 79. M. misionensis.
13. Epicuticular elements different or lamellae numerous.
14. Pileus white with dull bay or chestnut umbo becoming brown
all over when dried; lamellae about 16. Central American
species (compare subsection Pararotulae, key I, "3").
14. Not so colored or lamellae fewer (or both). South American
15. Pileus neither fresh nor dried umbilicate, umbonate or papil-
late, 5-6 mm broad; on fallen leaves (see key II below,
15. Pileus strongly papillate or stipe insititious on wood or on
rhizomorphs growing on wood.
Marasmius 123

16. Pileus 0.5-1 mm broad: spores 7-8 X 4-5/, or 8.2-9 X

2.2-3.5(-?5) (see key III below, "3").
16. Pleus larger;spores6.3-7.7 X 2-2.5(-3),u(see key II "3").
12. All stipes arisingdirectlyfrom the substratum.
17. Rhizomorphsin part pilose (pilosity caused by thick-walled,ob-
tuse, brownishhairs);pileus light cinnamonwhen fresh becoming
ratherdeep rusty-chestnutwhen dried;spores9.3-9.7 X 4-4.5u; on
dead leaves and petioles in tropical rain forest of the Amazon
region. 81. M. conquistensis.
17. Rhizomorphs,if present,glabrous;fungi not combiningthe charac-
ters indicatedabove.
18. Stipe shortand blackor almostblack;pileusavellaneous-brown-
ish but white around a central black dot (papilla), becoming
brownish-orange to light sordidorange-brownon drying;spores
9.5-13.2 X 4.5-6.2,i; rhizomorphsabundant; lamellae 9-10,
edgesmostly concolorouswith the pileus;on leaves. 82. M. boliviae.
18. Not combining these characters.Stipe over 20 mm long, not
deeper colored than the marginof the pileus or at least not
darkerthan fulvous or medium brown;pileus cinnamonbuff,
ochraceousbuff, light brown,clay color, fulvousor white;with-
out a white zone aroundthe centraldot or papilla(unless en-
tirely white excepting the papilla);spores as above or smaller;
rhizomorphsless abundant or absent (but "telepodes"often
present); lamellae 9-14; edges and collariumnot concolorous
with the pileus unless the latter is white on woody materialof
both Monocotyledonesand Dicotyledones.
19. Spores(4-)4.5-6.2j broad. 83. AMpallipes.
19. Spores7.5-9.5 X 4-4.2 . 84. M. aciculaeformis.
11. Pileus (fresh) partly or entirely brightercolored, orange,rufous, rusty-
orange,orange-yellow,salmon-orange, red, pink, purple,red-brown. Key II.
2. Sporesthree times longerthanbroador still narrower. Key II.

Key II
1. Pileusred to brown-redor red-brownor purplewhen fresh, either so coloredwhen wet
or becomingso when dryingout in situ.
2. Sporesthree timesas long as broador narrower(Q73).
3. Sporessmallerthan 7.7 X 3,; risingfrom rhizomorphs. 85. M. polycladus.
3. Spores largeror stipe not rising from rhizomorphsbut directly from the sub-
4. Sporesin theirgreatmajorityover 14p long.
5. On leaves and litter of all kinds, tropical; spores (9-)14-18.5 X
(1.5-)2.5-3.3/; lamellaedistant (11-15); edge of the lamellaepink or con-
colorous with the pileus, the latter tendingto be a rich deep brownwhen
dried. 86. M. rubromarginatus.
5. On wood in subtropicalvegetationof South America;spores still more
voluminousor lamellaesubdistant(16-17).
6. Lamellae >14. 87. M. marthae.
6. Lamellae<14. 88. M. sanguirotalis.
4. Sporesnot reaching14/uin length(see "10" below).
2. Sporesless than three times longerthan broad(Q < 3).
7. Stipes risingfrom rhizomorphs;both stipe and rhizomorphwith seta-likehairs(if
stipeglabroussee no. 80, p 131).
8. Pileus pink without a white zone around the dark purplelow umbo; spores
7.5-8 X 2.8-3U. 89. M. microdendron.
8. Pileusred with a white zone arounda white to blackishcentralpapilla;spores
7-9 X 3.5-4.2,u. 90. M. hippiochaetes.
7. Stipe either rising directly from the substratumor from rhizomorpns;in either
caseboth stipes and rhizomorphs(if present)glabrous.
9. On wood; stipe risingdirectlyfrom the substratum;pileuspurplered, 3-10 mm
broad;spores4-7,ubroad;context of pileus and hymenophoraltramapseudo-
amyloid, collariumoften discolorous(brown). 91. M. xerampelinus.
9. On fallen leavesor smalltwigs or stems; stipe risingdirectly from substratum
or from dark rhizomorphs;pileus as above or smaller;hymenophoraltrama
124 Flora Ncotropica

and pilcus-trama weakly or not pseudoamlyloid or distinctly pscudoaliyloid;

collariuin not brown.
10. Spores reaching more tllan 9pt in length; eitller only the papilla of tile
young pileus is red or the pileus-trama is inamiyloid.
II. Only the papilla of the young pileus is red, spores more than 4.5
broad. On Bromeliaceae. 96. M. hrolmeliacearni.
II. Margin of pileus also red in sonmestage or condition; spores less than
4.7, broad; pileus with a promiennt papilla wlhich may become less
prominent in age and is never surrounded by a white or pallid zone (if
such a zone is present, comipare M. giyanensis var crvlthrocephallis!)
trama of pileus and hymenophore inamyloid; setulae of broom cells
not red. 92. M. hiorami.
10. Spores up to 9u long; setulac of broom cells, if pigmented. at least par-
tially (upper part) red but the red pigment im0ostly clianging to brown or
imeleolus after a few minutes in alkaline ImediumI; hyphae of tile pilcus-
trama varying from very slowly and weakly to strongly pscudoainyloid
(reaction unknown in Al. ari)poensis).
12. Pileus imahogany red without a distilnct wlite or pallid zone around
the prominently papillate center; lamellae pale buff' witliout discolor-
otis edge or with white edge; spores 6-8 X 3p (Dennis); lamellae 8;
stipe arising from the substratumt. 93. M. arilpoensis.
12. Pileus usually more purple red or pinkish-red or becoming so when
drying in situ, generally with a distinct wliite or pallid zone around the
center whlich may be papillate or not; lamellae with strongly discolor-
ous (concolorous with the margin of the pileus) edge or with intermit-
tently to very weakly discolorous edge; stipe rising from the sub-
stratum or from rhizomorphs.
13. Edge of lamellae red; pileus 1-3.5 nimmbroad, disc umbonate,
blood-red to bright pinkish red. 95. M. edvalliaius.
13. Edge of lamellae eitlier not discolorous or miore often very nar-
rowly and intermittently concolorous with the pileus; pileus 2-6
nmmbroad, umbonate-papillate or not; collariumi often indistinct.
94. A1.lputtemansii.
1. Pileus orange, orange-rufescent, salmon-orange, golden yellow, yellow ochlr, orange-fer-
ruginous or brown to terra cotta but not becoming purple red when drying out in situ
nor with red papilla wheh young.
14. Spores three times longer than broad or still narrower.
15. Spores 16.5-19.5 X 4-4.2p . 97. M. megalospermus.
15. Spores only up to 16, long or shorter.
16. On wood, rarely on leaves; black rhizomiorphs present; spores larger than
11.7u (if on fern stems and grass see "18" below: if pileus sepia to mumimy
colored. See subsection Pararotulae, p 92).
17. Pileus golden yellow; rain forest species. 98. AM.chlrvsocephalts.
17. Pileus red-brown ("Kobe" M&P);subxerophytic species. 99. M. xcro)ph,'ticus.
16. On leaves of Dicotyledones and Monocotylcdones; rhizonorphs rarely
18. Pileus pale stramineous, gradually reaching light golden yellow ("mellow
glow" M&P) towards the center; spores 10-13.2 X 3.8-4.2u.
105. l. rhizomorphogeton.
18. Pileus differently colored; spores 2.5-4p broad. 100. M. guvanensis.
14. Spores less than three times longer than broad (Q <3).
15. Stipes glabrous, arising from a hirsute rhizomorph. 80. M. schultesii
15. Stipes if arising from a rhizomorph, then rhizomorph glabrous. Key III.

Key II1
1. Stipes rising from rhizomorphs (some may rise directly from the substratum).
2. On wood.
3. Spores 7.3-11 X 3.5-5.3p or somewhat larger.
4. Pileus 1.5-2 mm broad; stipe 3-4 mm long; all or most stipes rising from
rhizomorphs (see "6" below).
4. Pileus 1-8 mm broad; stipe 12-40 mm long; most stipes rising directly from the
substratum (see "11" below).
3. Spores 8.2-9 X 2.2-3.5(-5)u or 7-8 X 4-5 . 101. M. trichorhizus.
Marasmius 125

2. On leavesand monocotyledonousculms.
5. Spores9.5-13.2 X 4.5-6.2u; edge of lamellaediscolorous(see Key I above, "18").
5. Sporesless broad.
6. Pileus orange-ferruginous,drying brown ("cocoa" M&P);edge of lamellae
white and cheilocystidiawith hyaline setulae; spores 8.5-11 X 3.5-4.1M.On
dicotyledonousleavesin the Amazonasregion. 103. M. robertsonii.
6. Pileus orange with deeper colored papilla,sometimessomewhatbleachingin
the marginalregion; edge of the lamellae intermittentlyor indistinglydis-
colorous, some cheilocystidia with pigmented setulae; spores 7.5-11(-13) X
3.5-5 p. Mostlyon monocotyledonousleavesand culms. 102. M. crinisequi.
1. All stipesrisingfrom the substratum.
7. On wood, generallyof Dicotyledones.
8. Edgeof lamellaenot discolorous.
9. Carpophoresaccompaniedby black rhizomorphs.pileusoften ratherpale dirty
orange to pale cinnamon buff when wet, without a white zone around the
deep browncentralpapilla.
10. Spores9.5-12.3u long. 104. M. dicotyledoneus.
10. Sporesusuallyshorter(see "11" and "12" below).
9. Carpophoresnot accompaniedby black rhizomorphs.pileus orange with or
without a white zone around a darkerpapilla or umbilicus;spores 7.5-10l
long, about twice as long as broador slightlybroader(if sporesmoreelongated
compareno. 105 (M.rhizomorphogeton).
106. M. graminumvarschiniand varsubalpinus.
8. Edgeof lamellaediscolorous.
11. Spores4.7-5.3, broad.SouthernSouth America. 107. M. praecox.
11. Spores3.5-4.3,ubroad.Floridato Amazonas. 108. M. ruforotula.
7. On dicotyledonousand monocotyledonousleaves,on culmsof Bambuseaeand twigs
of other Monocotyledones,not on woody materialof dicotyledonous trees and
shrubs,rarelyon stemsand frondsof Pteridophyta.
12. Edgeof lamellaenot discolorous;on monocotyledonoushosts.
13. Spores relatively elongated, 10-13.2 X 3.8-4.2p; pileus ratherpale and dull
orangeocher, dried deeper colored (rufescent).Amazonasregion (see key II
above "18").
13. Sporeseither smalleror broader;pileuswell pigmentedorangeto ferruginous
orange or orangered, sometimespallescentin age. Subtropical,tropicaland
subtropical-montane zone of both hemispheres.
106. M. graminumn vargraminumand varculmisedus.
(Note: If on Bromeliaceae, compare A bromeliacearum, Key II, "11" no.
12. Edge of lamellae discolorous (concolorous with the margin of the pileus) or else
on dicotyledonous hosts.
14. Edge of lamellae not discolorous; on fallen leaves of dicotyledonous trees and
15. Spores 9-11.8 X 4.8-6.2 or 9-11.5 X 3.5-4.3 ; pileus bright "ferrugin-
ous" (M&P), becoming brown ("Arab" or "hazel" to 15 J 10, M&P)with
a pallid zone around a low umbo; hyphae of the trama of the pileus
inamyloid;rhizomorphseither none or scarceand inconspicuous;lamellae
12-13; collariumwhite. 110. M. pallidocinctus.
15. Spores 6.8-8.5 X 3.5-42; pileus deep rufous to rufescent-orange, on drying
becoming brown ("kis kilim" M&P) with a light brown zone around a
deep brown central dot, exumbonate; hyphae of the trama of the pileus
pseudoamyloid; inside of collarium tending to be brown; lamellae about
10; rhizomorphs accompanying the carpophores (if papillate and without
rhizomorphs, see "16" below). 109. M. brunneocinctus.
14. Edge of the lamellae discolorous at least at full maturity.
16. Spores 9.5-1 3.2 X 4.5-6.2/p (see key I above, "18").
16. Sporessmaller(cf also M. praecoxno. 107, see "11" above).
17. Spores 6.5-10(-10.2) X (2.5-)3-4.5 ; at least the dry pileus with a low
umbo or papilla;lamellae7-11. On dicotyledonousleaves.
111. M. rufomarginatus.
17. Spores 7.3-10 X 4-5u, i e relatively broader; central dark dot of the
pileus generally not elevated; lamellae 10-13. More frequently on
monocotyledonous than on dicotyledonous leaves. 112. M. foliicola.
126 Flora Neotropica

71. Marasmiuspeckii Murrill,North AmericanFlora 9: 254. 1915. Fig 52 P.

TYPE.M. E. Peck, from Belize.
Pileus whitish, sometimes discoloringto gray but mostly remainingwhitish when
dried,with blackishor dark chestnut-fuliginouspapilla,glabrous,sulcate, hemispherical,
unbilicatewith a dark, ratherlow papillain the umbilious,up to 2 mm broad. Lamellae
white, distant, (8-10 through-lamellae,no lamellulae),very broad, distinctly collariate.
Stipe brown or blackish,glabrous,shining,setiform, instititious, not accompaniedby
dark rhizomorphs,20-30 mm long; some sterile stipes (telepodia) present.
Spores 5.8-6.8 X 1.8-2.8p, oblong to fusoid, smooth, hyaline, inamyloid. Hymen-
ium: basidia 15.3-16.7 X 5.5-6.5., 4-spored;cystidia none; cheilocystidialike the epicu-
ticularbroom cells. Hyphaewith clamp connections, parallel and stronglypseudoamy-
loid in the tramaof the stipe. Corticallayer: epicutis of the pileus hymeniform,broom
cells of the Siccus type, main body 5.5-10 X 5.5-7.8,u, mostly clavate,hyaline, setulae
apical 3.3-9 X 0.5-0.9- (2.2)p, spinose, subacuteto obtusate, hyaline.
On dead dicotyledonous leaves in BritishHonduras.
MATERIALSTUDIED. BELIZE(British Honduras),1906, M. E. Peck (NY), type.

72. Marasmiuschrysochaetes(Berkeley & Curtis),Jour. Linn. Soc. Bot. 10: 297. 1868.
Crinipellischrysochaetes(Berkeley& Curtis)Singer,Lilloa8: 461. 1942.
TYPE.Wright162, from Cuba.
Pileuswhite, dried tending to "pinkishbuff' Ridgway,and developingsometimes
a brown umbo, sulcate, glabrous,convex, umbilicate,with a small umbo in the um-
bilious, 1.2-2 mm broad. Lamellaewhite, distant (about ten throughlamellae,no
lamellulae),ratherbroad, distinctly collariate. Stipe fulvous or light brown, shining,
insititious,glabrous,smooth, 22-25 X 0.2 mm (dried), some telepodes. Context ex-
tremely thin.
Spores 8-9 X 3.5-4.2u, ellipsoid-oblong,smooth, hyaline, inamyloid. Hymenium:
basidia4-spored,basidiolesfusoid; cystidia none; cheilocystidialike the epicuticular
broom cells but generallyhyaline and inamyloid. Hyphae of pileus-tramaand hymen-
ophoral tramahyaline, thin, ratherthin-walled,inamyloidwith clamp connections; those
of the stipe parallel,pseudoamyloid. Corticallayers:epicutis hymeniformor subhymen-
iform, broom cells of the Siccus type, 14-28 X 7-9u, with apical obtuse appendages
which are often finger-likeand 2-3/ubroad, entire wall firm to thick (to 3.5 ,)- walled,
Subcespitoseon dead leaves of the evergreenkind. Cuba.
I have not studied part of the type deposited at Kew but Dennis indicates
similarcharacteristicsas given above althoughhe finds the sporesslightly narrower
(9-10 X 3-3.5g).
ILLUSTRATIONS: Singer(1942)fig 1,g. Dennis(195lb) fig3.

73. MarasmiusaspilocephalusSinger,Sydowia 18: 334. 1965.

TYPE.SingerB 2214, from Bolivia.
Pileus white, in umbilicuspale buff or pale cinnamon,weakly to distinctly
sulcate where white, glabrouswithout a darkdot in the center, not papillate,convex,
umbilicate,4-7 mm. broad. Lamellaewhite, with white edges, distant (7-9), equal or
with 1-2 lamellulaeintermixed,broad, collariate. Stipe black with white apex,
glabrous,instititious, setose, 5-6 X 0.1-0.2 mm.
Spores 8-11 X 4.5-5.5 ,, smooth, hyaline, inamyloid. Hymenium:cystidia,
Marasmius 127

not seen; cheilocystidialike epicuticularbroom cells. Hyphaeof pileus tramahyaline,

with clamp connections, inamyloid or almost so. Corticallayers:epicutis of pileus
hymeniform,its elements of the Siccus-type,hyaline with hyaline apicalsetulae, few
with golden melleous setulae, main body e g 9-11 X 8-1 1 , setulae up to 5 A.long,
On leavesand small culms of Bambusasp.
MATERIALSTUDIED.BOLIVIA.Pando:Conquista,25 Mar1956. SingerB 2214 (LIL)
Singer(1965) pl 9, fig 32

74. MarasmiuspallenticepsSinger sp nov

Marasmius trichorhizus var pallenticeps Singer. Sydowia 18: 178. 1965 (nomen nudum).
TYPE.SingerM 1123, from Argentina.
Pileo albo vel pallido, juventute papilla atrobrunneaprominente ornato; lamellis
albis, collariatis;stipite aut e substratoaut e rhizomophisnato. Sporis 7-8.5 X 2.7-4.5,;
elementis epicuticularibustypi Marasmiichrysochaetispraesentibus. Ad ramos emor-
tuos in silva subtropicaliargentinensi. Typus in LIL conservatusest.
Pileus white or pallid, in dried condition stramineouswhitish, only a few caps
now brown, glabrous,weakly sulcate, with a darkbrown prominentpapillain young
specimens, later more repandand maintainingthe darkpapillaor papillabecoming
paler, 1-2 mm broad, 0.5-1.2 mm high. Lamellaewhite with apparentlypallidedges,
distant (7), collariate. Stipe copper brown, reddishbrown, glabrous,smooth, shining,
insititious, or pallid and risingfrom a copper brown shiningrhizomorph,the rhizomor-
phogenousstipes about 3-5 X 0.05-0.1 mm, the other 4-10 X 0.1-0.2 mm; rhizomorphs
often very long and often ramose,smooth and glabrous,insititiouswhere insertedin
the substratum. Context white, inodorous.
Spores 7-8.5 X 2.7-4.5 ,, ellipsoid, smooth, hyaline, inamyloid. Hymenium:
not studied. Hyphae of pileus tramathin-walledwith clamp connections, inamyloid
or almost so; of stipe strictly parallel, thick-walled,strongly pseudoamyloid,also the
hyphae of the rhizomorphs. Corticallayers:epicutis of pileus hymeniformor sub-
hymeniform,consistingof broom cells with erect or somewhatspreading-subdiverging
strictly apical setulae, e g 8 X 5.5 p,, setulae 2.5-4 X 0.3-1 ,, main body hyaline, setulae
hyaline to pale golden melleous;these elements are intermixedwith coarseelements of
the Chrysochaetes-type,i e thick-walledand not setulose but with coarsebroadver-
rucose to finger-likeprocesses,or nodose, versiform,hyaline and strongly pseudoam-
loid, but strongly colored brown in center (papilla),often forked.
On dead branchesin subtropicalforest, gregarious.
MATERIALSTUDIED. ARGENTINA.Misiones:Frontera,GeneralManuelBelgranonear
San Antonio, 19 Apr 1957, SingerM 1123 (LIL), type.
Singer (1965) pl 9, fig 33.

75. MarasmiusconicopapillatusHennings,EnglerBot. Jahrb. 22: 100. 1895.

TYPE.Dusen, from Cameroons(type lost).
Pileus white, eventually and mostly in the herbariumbecoming light beige to
brown (e g "Arab"M&P),with the center from the beginningbrown to black,
glabrous, sulcate, center smooth, papillate, convex, later papilla in an umbilicus,
1-6 mm broad. Lamellaewhite or whitish with white or whitish edge, distant, (6-12
through-lamellae,0-2 lamellulae), simple, ratherbroad to broad, collariate. Stipe
black, at first with white apex, glabrous, smooth, setiform, simple and insititious
128 FloraNeotropica

on the substratum,mostly accompaniedby black rhizomorphs,9-38 X 0.1-0.2 mm.

Context thin, inodorous.
Spores 6.8-12 X 3.2-4.5,, ellipsoid to oblong or subcylindrical,smooth,
hyaline, inamyloid. Hymenium: basidia 11.5-18 X 5-7.3s, 4-spored; cystidia none;
cheilocystidia like the epicuticularbroom cells. Hypae of the pileus trama and the
hymenophoral trama hyaline, with clamp connections, partly and weakly pseudoamy-
loid, many remaininginamyloid in Melzer'sreagent. Cortical layer: epicutis of pileus
hymeniform, broom cells of the Siccus-type, main body hyaline, vesiculose to
clavate, 4.5-17.5 X 4.5-12.31, sometimes lobed, thin-walled,setulae 2-5.3 X 0.6-1.7M,
conical or rod-shaped,obtuse or subacute, hyaline.
On dead leaves of both Monocotyledones and Dicotyledones in tropical rain
MATERIALSTUDIED. BOLIVIA. Pando:Madrede Dios, Las Piedras,5 Apr 1956,
109 (BR);Yangambi,Louis 14935 (BR).
SingerB 2528 (LIL). ZAIRE. Eala,Goossens-Fon2tana
If indeed the American collection is identical with the African collection, the
spores must be very variable. It is possible that the Americanspecies with spores
8.2-12 X 4-4.5 u (slightly larger)is a constantly larger-sporedrace (subspecies) but
too few collections have been made to separatethem under a subspecific name.
IILUSTRATION. Singer (1964b) fig 18, D-E; (1965), pl 8, fig 31.

76. Marasmiusnigrobrunneus(Patouillard)Saccardo,Sylloge Fungorum 11: 37. 1895.

AndrosaceusnigrobrunneusPatouillard,Jour. de Bot. 5: 308. 1891.
MarasmiusgriseoviolaceusPetch, Trans.Brit. Mycol. Soc. 31: 42. 1947.

TYPE. Bon 4397, from Vietnam.

Pileus gray (between "Deauvillesand" and "turtledove," "smoke brown")
becoming "limestone" M&P,sometimes whitish cinereous, glabrous,sulcate to deep
sulcate-groovedto the umbilicus, with a low black papilla or central dot in the um-
bilicus or a shallow depression, 2-9 mm. broad. Lamellaewhite, the edges white or
concolorous with the pileus, equal, or with 1(-2) lamellulae,8-13, medium broad to
broad, very narrowly but distinctly collariate, distant. Stipe black or deep umber
with white apex, unpolished to somewhat shining, glabrousmacroscopically,but
sometimes distinctly scabrous-spinuloseunder a lens in the lower part, smooth,
insititious either (rarely in the neotropics) on the rhizomorphs,or more commonly
directly on the substratum, 13-73 X 0.1-0.5 mm. Context white, thin, inodorous.
Spores (6.2-)7.5-10.5 X 3.5-5.5,, mostly about 9 X 4-5A, or in some collections
slightly smalleron an average,ellipsoid to oblong-ellipsoid,smooth, hyaline, inamyloid.
Hymenium: basidia 24-28 X 6-7.8M, 4-spored; cystidia none; cheilocystidia like the
epicuticularbroom cells but in some specimens rather scattered or all hyaline (and
then lamellae with white edges). Hyphae of pileus hyaline, weakly and slowly but
undoubtedly pseudoamyloid, those of apex of stipe strongly pseudoamyloid, all with
clamp connections. Corticallayers: epicutis of pileus hymeniform (but sometimes deter-
sile) and consisting of broom cells of the Siccus-type, although often rather coarsely
diverticulateat apex, main body hyaline and thin-walledor at least in upper portion
sordid gray to deep sepia-fuliginousor umber, some without setulae, some more or less
thick-walled,9-20 X 4.5-12A, mostly vesiculose-clavate,setulae mostly apical and
erect, hyaline to sepia-fuligineousor umber, 2-5.3 X 0.7-2.81, generally ratherbroad
and almost always rounded-obtuseat tip; covering layer of stipe inconspicuous, con-
sisting of small hairs, sometimes ascendingat the apex and the base and then sausage
Marasmius 129

shaped, in other cases erect and cylindric, blackish fuscous, thick-walled,setose, but
obtuse, e g 11-14 X 5.5,.
On dead leaves and culms of monocotyledonous plants, especially Bambuseae
(Banibusa,Chusquea),but also frequently on Bromeliaceae(Aechmea) and palms.;
smaller carpophoresoften on grasses.
MATERIALSTUDIED. TRINIDAD. St. Joseph, 30 Nov 1949, Deinnis409 (K). BOLIVIA.
La Paz: Nor-YungasCarmenPampa,2000 m alt, 26 Feb 1956, SingerB 1544 (LIL);B 1545 (LIL).
ARGENTINA. Tucuman:Capital,22 Jan 1955, Singer T 2077 (LIL); 10 Feb 1955, T 2149 (LIL);
T 2150 (LIL); Tafi del Valle, 17 Feb 1956, Singer T 4081 (F); Jujuy: Lagunasde Yala, 14 Feb
1966, Singer T 5127 (F). VIETNAM. Hanoi, Kes6, 31 May 1890, Bon 4397 (FH), type; Demange
6 7 (H). INDIA. Gurdespur,JaggatPur, 15 Aug 1941, SultanAhrmad509 (FH). CEYLON. Bo-
tanic Gardens,Peradiniya,1 Nov 1914, Petch 4238 (K, FH). typus of M. griseoviolaceus.
ILLUSTRATION.Singer (1958a), fig 48 [as AI. griseoviolaceus]; Dennis (1951a) 19,
fig 18 [as MI.griseoviolaceus/.

77. MarasmiusfuligineorotulaSinger, Sydowia 18: 336. 1956.

TYPE.SinlgerB 1193, from Bolivia.
Pileus gray black ("Cordovan"),becoming "Tuscan tan" (M&P)(isabelline-argilla-
ceous) in age. without a distinct black dot or papilla in the center, campanulate,then
convex, umbilicate, primordiaonly papillate, sulcate when mature, 4-11 mm broad.
Lamellaewhite, 7-10 equal, collariate. Stipe black with white apex, glabrous and
smooth, insititious. 5-10 X 0.1-0.2 mm; black rhizomorphspresent, but scattered and
Spores (7.5-)10.2-12.3 X 4.5-6u, somewhat broaderthan half their length or about
half their length in width, more rarely narrowerthan half their length, ellipsoid-sub-
fusoid, thin-walled,smooth, hyaline, inamyloid. Hymenium: basidia 16-29 X 5.5-7.2u,
mostly 4-spored, some few at times 2-spored;cystidia not observed;cheilocystidia like
epicuticularbroom cells, but very few with other then hyaline setulae and setulae more
clearly of the Rotalis-type. Hyphae of the pileus tramapartly weakly pseudoamyloid,
with clamp connections. Corticallayers: epicutis of puleus hymeniform, consisting of
elements varying in shape and appearancebetween the Siccus-type and the Rotalis-
type, the majority of the Siccus-type, but setulae covering never more than one third
of the cells, main body hyaline, often with thickened walls (wall up to 1.5 u), clavate
to vesiculose-pedicellateor subglobose-pedicellate,10-24 X 4-10.5 u, some cells smooth
without setulae, the majority with hyaline or spadiceous to blackish-spadiceoussetulae
of extremely variableshape and size, some forked, some acute, some obtuse.
On dicotyledonous stickes in montane tropical forest.
MATERIALSTUDIED. BOLIVIA. La Paz: Nor-Yungas,Rio Yariza, 16 Feb 1956,
SingerB 1193 (LIL), type.
This species is intermediatebetween the species with Rotalis-type epicuticular
elements and those with Siccus-type.
ILLUSTRATION. Singer (1965) p 8, fig 30.

78. MarasmiusmagnisetulosusSinger, sp nov Fig 53.

TYPE. Dumonit et al VE 6206, from Venezuela.
Pileo griseo vel brunneolo-griseoin siccis, sub lente velutino, 0.3-1 mm lato, 0.2-
0.5 mm alto; lamellis albidis vel grisello-albis,acie haud discoloribus,distantibus,colla-
riatis;stipite atrobrunneoapice albido-pallido;rhizomorphisabsentibus. Sporis 5.5-6.5
X 3-3.5,; hyphis pilei inamyloideis, fibulatis;elementis epicuticularibuscystidiiformi-
130 Flora Neotropica

bus dimorphis, nonnullis integris. Ad folia dicotyledonea emortua in Venezuela. Ty-

pus in NY conservatusest.
Pileus gray or brownish gray (dried), long-sulcate,macroscopicallyglabrous, under
a lens velvety, semiorbicularto campanulateor subcylindrical,with an apical flattening
in the middle of which there is a conspicuous papilla, 0.3-1 X 0.2-0.5 mm. Lamellae
whitish or grayish white, with not discolorous edge, distant (seven through-lamellae,no
lamellulae), so short and broad that the interlamellarspaces appearlike pores, collari-
ate. Stipe deep brown with pallid apex, glabrous, smooth, setiform, insititious, about
7-8 mm long, 800 p broad; no rhizomorphs. Context very thin.
Spores 5.5-6.5 X 3-3.5up,ellipsoid to fusoid-ellipsoid,smooth, hyaline, inamyloid.
Hymenium: basidia 18 X 5.5 1, 4-spored; cystidia none; cheilocystidia like the elements
of the epicutis sometimes differentiated. Hyphae in pileus trama hyaline, thin-walled,
not gelatinized, inamyloid, 2-5p broad, mostly filamentous, some inflated up to 9p
broad, with clamp connections. Hyphae of the stipe hyaline in inner, tawny in rind-
layer, all with clamp connections (but some secondary septa present), 2-6p broad,
parallel,weakly pseudoamyloid. Hymenophoraltrama regular,of subparallel,hyaline
hyphae 2-34ibroad. Cortical layers: epicutis hymeniform, its elements consisting of
upright, cystidiform cells of two types (1) 10-18 X 4.5-7,u, very numerous, with finger-
like, mostly erect to slightly oblique appendageswhich are il their majority apical (two
or four) but some lateral, basically of the Siccus-type but more clavarioidor reminding
one of the cheilocystidia of Marasmiellus,definitely not of the Rotalis-type. hyaline be-
low, brown above, some entirely brown, some merely mellous, with walls 0.2-0.5, thick,
appendages3-15 X 2-61 , with obtuse tip, (2) 13-35 X 5-7p, rather numerous,entire,
not branchedor a few with one very short branch near the base, cylindricalampullace-
ous or clavate, some finely granularincrusted, otherwise like type (1). Surface of stipe
without vestiment, but some occasional hyaline hyphae present, these thin and appressed:
neither these nor the elements of the epicutis pseudoamyloid.
On coriaceous dead dicotyledonous leaves, gregarious. Venezuela.
MATERIAL STUDIED. VENEZUELA. Dto Federal: Parque Nacional El Avila, 27 Feb 1972,
Dumont et al VE 6206 (NY), type.
The unusual structureof the epicutis and the shape of the pileus, small number of
lamellae and small spores are characteristic.

79. MarasmiusmisionensisSydowia 18: 338. 1965.

TYPE. Singer M 1047, from Argentina.
Pileus at first light brown, then cinnamon fuscous, with paler stripes in the sulci.
dried uniformly "auburn,"sulcate, glabrous, convex, with a small, insignificant,concolor-
ous umbo in a central depression, 1.7-2 mm broad. Lamellaewhite or whitish, with
white or whitish edges, broad, distant (7), equal, collariate. Stipe brown, then black,
dried umber, shining, glabrousand smooth, risingfrom a slightly thicker rhizomorph,
14 X 0.1-0.2 mm.
Spores 7.5-9 X 44.5,4, ellipsoid, hyaline, smooth, inamyloid. Hymenium: basi-
dia 18 X 5.5,L; basidioles fusiform; cystidia none; cheilocystidia like epicuticularbroom
cells but not or not strongly pigmented. Hyphae of pileus trama and hymenophoral
tramahyaline, with clamp connections, interwoven, thin-walled,some somewhat thick-
walled, slightly but distinctly although unevenly pseudoamyloid. Cortical layer: epicutis
of pileus hymeniform, consisting of broom cells of the Siccus-type but transient to the
Chrysochaetes-type,12-22 X 6-13A1,most with setulae but some entire, characteristic-
ally colored between cinnamon and vinaceous brown, with a few hyaline ones inter-
Marasmius 131

mixed, concolorous in main body and setulae, all the pigmented ones and many of the
hyaline ones thick-walled(wall 1-22,thick). setulae coarse, obtuse, up to 5.5,i long and
2-2.5k, broad, mostly cylindric, where they are missing the cells are fusoid e g 17 X
On forest debris.
MATERIALSTUDIED. ARGENTINA. Misiones:Iguazfu,Refugio Yaguarete,16 Apr 1957,
Singer M 1047 (LIL) type.
ILLUSTRATION.Singer (1965) p1 10, fig 4.

80. Marasmiusschultesii Singer, sp nov

TYPE. Schultes & Cabrera12869, from Colombia (F).
Pileo brunneo, unicolori, papillato, 0.5 mm lato in siccis; lamellis ad marginem
aciemque haud discoloribus 6 integris, collariatis;stipite rubido-brunneo,glabro, e
rhizomorphishirsuto-pilosisnato. Sporis 7.5-9 X 4-5.5,u; cheilocystidiis hyalinis vel
subhyalinis,ceterum elementis epicuticularibussimillimis;hyphis tramatispilei inamy-
loideis; elementis epicuticularibuspilei aurantiacisvel aurantiaco-luteis,rariussubhya-
linis, crasse tunicatis. Ad radicemin silvatropicali. Typus in F conservatusest.
Pileus brown, dried cinnamon, glabrous, sulcate, with a central papilla which is
concolorous and not surroundedby a pallid zone, later small-papillatein a central de-
pression, dried up to 0.5 mm broad. Lamellaebuff-pallidwhen dried, not discolorous
at the edge, about six through-lamellaeand no lamellulae,broad, collariate. Stipe red-
dish brown, glabrous, attached to the much thicker rhizomorphby a node and some-
times also with a mediane node, rarely branched,equal, about 3 mm long and up to
0.1 mm broad; rhizomorphsconcolorous, hirsute-piloseall over, about 65-120 X + 0.3
mm, insititious. Context pallid, very thin. Spores 7.5-9 X 4-5.5j, ellipsoid to more
rarely oblong, smooth, hyaline, inamyloid, tending to germinatefrom the distal point
in situ. Hymenium: basidia about 12 X 5p; cystidia none seen; cheilocystidia like the
epicuticularbroom cells but mostly hyaline or subhyaline. Hyphae of the trama of the
pileus and the hymenophoral tramahyaline or subhyaline, filamentous, inamyloid, with
clamp connections. Cortical layers: epicutis of the pileus hymeniform, consisting of
broom cells of the Siccus-type, main body 10-15(-18) X (4-)4.5-8.5A, orange to orange-
yellow in NH40H, more rarely subhyaline, with 1.2-1.5Mthick wall, more rarely with
thin wall, with few to numerousapical setulae, these obtuse, concolorous with the
Imainbody, 3.5-7.5(-13) X 1-1.8(-2.5)), often forked, covering of the stipe consisting
of very scattered setiform hairs which are hyaline or subhyaline and acute, about 35
X 5.5L,, with about lp thick wall.
On a root, saprophytic, in tropical forest ("highlandforest" of the Amazon ba-
MATERIALSTUDIED. COLOMBIA.Amazonas,Soratama,Rio Apaporis,between Rio
Pacoa and Rio Kananarfabout 250 m altitude 3 Jul 1951, R. E. Schultes & I. Cabrera12869
(BPI, F), type.
The color of the pileus as indicated by the collectors as well as the spore data
and thick-walledbroom cells suggest affinity with Ml.misionentsis(no. 79); the hairy
rhizomorphsuggestsaffinity with M. microdendronand M. hippiochaetes. If the
scarce setiform hairs on the stipe should mean that the stipe is initially hairy as the
rhizomorphs,M. schultesii would be different from M. microdendronbecause of the
much broaderspores, and from M. hippiochaetes by the absence of a white zone
around the central papilla; it differs from both in the brown color of the marginof
the pileus.
132 Flora Neotropica

81. Marasmiusconquistensis Singer, Sydowia 18: 334. 1965.

TYPE.Singer B 2190, from Bolivia.
Pileus light cinnamon when fresh, on drying becoming rather deep rust brown
to ferruginous-reddish-brown (near "chutney" when dried but slightly less red and
more brown), young caps usually having a distinct pallid zone around a black papilla,
later the pallid zone disappearingand a small black papilla remainingin a central um-
bilicus or depressionwhen mature, otherwise convex when mature, sulcate over the
convex part of the pileus, 2-11 mm broad, 0.7-1.2 mm high. Lamellaewhite with
discolorous edge and collarium, distant, 9-10, rarely intermixed with one lamellula,
with well formed collarium. Stipe black with white apex, later entirely black, gla-
brous and smooth, shining, insititious with a simple very minute ring directly on the
substratum,about 7-13 X 0.1-0.2 mm; black rhizomorphspresent, accompanyingthe
carpophoresbut not connected with the stipes, in part more or less pilose.
Spores 9.3-9.7 X 4-4.5,i, hyaline, smooth. Hymenium: basidia 21.5 X 7p, cheilo-
cystidia like epicuticularbroom cells. Hyphae of pileus trama and hymenophoral trama
hyaline, thin-walled,with clamp connections, slowly and extremely weakly pseudoamv-
loid, practically inamyloid. Cortical layers: epicutis of pileus consisting of broom cells
of the Siccus-type, main body hyaline or only upper portion somewhat melleous to br-
ownish, 11-13.8 X 5.3-7 ,, rather thin-walled,cylindric to clavate or subventricose,set-
ulae 3-5.5 X 1.2-1.6p, either hyaline or deep reddish rust brown, obtuse; covering of
the rhizomorphsconsisting of numerous setae which are 10-100 X 2.5-5s cylindric, or
sometimes with a ventricose portion up to 9i, broad, or sometimes capitate at apex
(apex 9-13 j diameter), with thick (1.3-2.2,u) wall, brownish, more often obtuse than
acute, much like the epicuticularelements of Crinipellis,not incrusted.
On dead fallen dicotyledonous leaves and their petioles in rain forest.
MATERIAL STUDIED.BOLIVIA.Pando:Manuripi, Conquista, 25 Mar1956.SinlgerB
2190 (LIL),type.
ILLUSTRATION. Singer (1965) pl 10, fig 43.

82. Marasmiusboliviae Singer, Sydowia 18: 334, 1965.

TYPE. Singer B 1098, from Bolivia.
Pileus avellaneous-brownish ("toast" M&P),white around a black central dot,
papillate where the black dot is located, at least in dried condition, dried brownish or-
ange light sordid orange brown (1 l-C-9, further inwards "caramel"M&P).sulcate to
disc, convex with depressedcenter, 2.5-3.5 mm broad. Lamellaewhite, with edges con-
colorous with the pileus but less so in some specimens near the collarium. distant, 9-10
(no lamellulae), broad (0.5 mm), collariate. Stipe black with white apex, smooth and
glabrous, simple, very rarely forked, shining, setose, insititious, 10 X 0.1 mm; black
rhizomorphsassociated with the carpophores,also insititious where arisingfrom sub-
stratum, glabrous.
Spores 9.5-13.2 X 4.5-6.2 u, ellipsoid, smooth, some slightly pseudoamyloid, most
inamyloid, hyaline. Hymenium: basidia 30 X 8.2,, 4-spored, inamyloid, a few slightly
pseudoamyloid, clavate; cystidia none; cheilocystidia like epicuticularelements. Hyphae
of pileus and hymenophoral trama hyaline, filamentous, some with somewhat thickened
walls, many pseudoamyloid, others inamyloid, all with clamp connectioIls. Cortical lay-
ers: epicutis of pileus consisting of a hymeniform layer of broom cells of the Siccus-
type, mottled by regularlydistributed cells without pigment, with pale golden yellow
pigment or entirely deep golden melleous, main body with cinnamon to hyaline cell
sap, 5.5-16.5 X 5.5-10s, mostly vesiculose, rathervariablein volume, setulae coarse
Marasmius 133

and at times few, numberingfrom one forked setular up to 12 setulae, 4-12 X 1.3-1.7
,, rarely reachingup to 2.8s diameter at base, either taperingupwardsto a subacute
or obtuse tip, or more frequently cylindric with broadly rounded tip.
On the thick central nerve of dicotyledonous leaves (Inga ? edulis).
MATERIALSTUDIED. BOLIVIA. La Paz: Nor-Yungas,Coroico, 12 Feb 1956, SingerB
1098 (LIL), type.
This species, in dried condition, has the appearanceof Marasmiusdicotyledoneus,
I. gramfiumtvar schinii,etc. Only few spores were found in the type collection and
it may be expected that the maximum-minimumrange is still wider than indicated
Marasmiiusboliviae differs from the Argentine forms mentioned above, in the
more voluminous spores, the discolored edge of the lamellae and the sometimes scanty,
sometimes very long setulae of the broom cells, also by the habitat on dicotyledoneous
leaves, and most easily, by the color of the fresh pileus.
ILLUSTRATION. Singer(1965), pl 10, fig 46.

83. Marasmiuspallipes Spegazzini, Anal. Soc. Ci. Argentina 16: 272. 1883.
TYPE. Balansa3379, from Paraguay.
Pileus cinnamon buff, ochraceous buff, light brown or clay color, at times slightly
zonate in the declivous portion, or graduallypaler towards the extreme margin,dried
between "gold br." and "Bombay M&P,"occasionally with somewhat more ochraceous
extreme margin("chipmunk") and deeper regions in center ("coffee"), without a white
or pallid zone around a black central dot or papilla, the central dot well differentiatedwit]
an extraordinarilysmall papilla in its center, in younger stages papilla often prominent
and deep chestnut to blackish chestnut, later slightly convex to flattened, sulcate except
in the central umbilicus containing the black dot, otherwise convex and glabrous,
2-8 mm broad. Lamellaewhite, edges and sides white, eventually or dried becoming
somewhat paler than the pileus, broad, distant, equal or with 1-2 lamellulaeor 1-2
forked ones when mature, 9-13 through-lamellae,collariate. Stipe paler than the
pileus, often reachinglight chestnut below and pale stramineousabove, dried usually
almost uniformly pale cinnamon or pale reddish fuscous, or sordid stramineous,
smooth and glabrous,shining, insititious to the substratumwith a very minute basal
disc, at apex often white in young specimens, setose-wiry,equal 18-40 X 0.1-0.7 mm;
rhizomorphsconcolorous with stipe or black, not regularlyobserved, accompaniedby
"telepodes," all glabrous,often branched,mostly scanty. Context thin, white, in-
Spores (7.5-) 8.2-14 X (4-)4.5-6.2p, in some collections relatively small with the
most common size being 9.7 X 4.5k or 10-11 X 4-4.8 uellipsoid to ellipsoid-oblongor
even slightly subfusoid, smooth, hyaline, inamyloid, tending to germinateon the lam-
ellae. Hymenium: basidia(19-)23-32 X 6.8-8.10.5, , 4-spored;basidioles fusoid; cyst-
idia none: cheilocystidia like epicuticularbroom cells but predominantlyhyaline to
pale melleous with melleous to melleous golden setulae. Hyphae: trama of pileus and
hymenophoral tramaweakly pseudoamyloid;that of stipe somewhat more strongly but
not deeply or rapidly pseudoamyloid;all hyphae with clamp connections. Cortical
layers: epicutis of pileus hymeniform, consisting of broom cells of the Siccus-type,
main body 11-22 X 6.5-12,L, often hand-shapedor Clavariella-shaped, some slightly
thick-walled(0.6gi), brownish hyaline to pale melleous, some quite hyaline, setulae
apical and erect but sometimes slightly spreading-divergent-subapicular, 2-5.5 X ? 1.3j
134 Flora Neotropica

(at base), deep golden melleous, melleous, or brown, sometimes coarse, spinulose to
subcylindric,more often obtuse than subacute.
On bark,roots, woody monocotyledonous and dicotyledonous materialin forest
humus, dead twigs, mossy logs in tropical and subtropicalforest, densely gregariousto
MATERIALSTUDIED. BRAZIL.Santa Catharina:Sao Canisiode Porto Novo, 1928, J. Rick
642 (FH). PARAGUAY. Guarapf,Mar 1880, Balansa3379 (LPS), type. ARGENTINA. Salta: Orin,
3 km west from Rio Blanco bridge, 17 Mar 1955, Singer T 2302 (LIL);Misiones:Iguazu, Punto Canoa
29 Feb 1960, R. T. Guerrero118 (LIL). BOLIVIA. Beni: Vaca Diez, Ivon, 3 Apr 1956, SingerB
2480 (LIL).
This occurs also in Trinidadon Ficus benjaminasince materialdescribed and ill-
ustrated by Dennis (195 la) under the name of M. aciculaefbrimisB & C. appears to be
Spegazzini apparentlydescribed the materialexclusively from dried material,and
it is obvious that he misdescribedthe colors.
The Salta and Bolivian materialdiffers from the other materialsin having smaller
spores; these are in the former only 9.7-10 X 4.8-5.4s, but some spores reach 11.2 X
5.8,u, in the latter 9.7 X 4.5 more rarely 8.2-1 1.7 X 4-4.8,u. It is this material that
has a distinct papilla when young, becomes graduallypaler colored (from chestnut to
ochraceous buff and light brown and eventually pallescent to pallid), maintaininga
deep chestnut central dot (umbo), and becoming between "gold br." and "Bombay"
M&Pwhen dried. The Brazilianmaterialbecame more gray-brownwhen dried and
showed less of a darkercentral umbo. It was also less pseudoamyloid. The mater-
ial from Misiones has very short and broad basidia(19-20 X 8-10.5,).
Whetherthese small differences are significant can be determined only after
the recollection of more material from more regions.
ILLUSTRATION. Singer(1965) pi 10, fig 43; Dennis (195 a) pl 20, fig 5.

84. MarasmiusaciculaeformisBerkeley & Curtis, Jour. Linn. Soc. Bot. 10: 297. 1868.

84a. Marasmiusaciculaeformisvar aciculaeformis

TYPE. Wright,from Cuba.
Pileus fulvous, scarcely sulcate, convex, 4 mm broad. Lamellaewhitish, few,
collariate. Stipe almost fulvous, shining, insititious, glabrous,40 X 0.2 mim.
Spores 7.5-9.5 X 4-4.2,4, ellipsoid to oblong, smooth, hyaline, inamyloid. Hymen-
ium: basidia 25-27 X 7,; basidioles fusoid; cystidia none; cheilocystidia like epicuticular
broom cells. Hyphae hyaline, inamyloid in pileus-trama. Cortical layer: epicutis of pi-
leus hymeniform, broom cells of the Siccus-type, hyaline or subhyaline, in superposition
yellowish, with coarse apical conic to conic-obtuse setulae.
On rotten wood densely cespitose, Cuba.
MATERIALSTUDIED. CUBA. Wright121 (K, FH), type.

84b. Marasmiusaciculaeformisvar albus Dennis, Kew Bull. 15: 98. 1961.

Pileus white, sulcate, with a chestnut-brownsmall papilla in the center, 5-6.5 mm
broad. Lamellaewhite, subdistant(12-14 lamellae), collariate. Stipe brown with white
apex, glabrous, insititious, 25-30 X 0.2-0.5 mm.
Spores 7-9 X 4,p (Dennis).
Otherwise apparentlylike Marasm7ius pallipes Spegazzini(no. 82 above).
On dead wood, cespitose.
Marasmius 135

MATERIAL STUDIED.Colorsketchandnotesby Dennison the type of the variety.VEN-

BotanicGarden,6 Sep 1959,Dennis1023 (K).
EZUELA.Dto Federal:Caracas

85. Marasmiuspolycladus Montagne,Ann. Sci. Nat. Bot. IV. 1: 113. 1854. Fig 54.
TYPE.Leprieur, from French Guyana.
Pileus brownish purple to blood red, brown when dried, glabrous,sulcate, hemispher-
ical to convex, with a strong papilla later in an umbilicus, 4-8 mm broad. Lamellaefuscou
(dried), with not discolorous edge, distant, broad, collariate. Stipe spadiceous-black,
short, about 0.25 mm broad, inserted in a sheet- to volva-likenode of the broad black,
shining, glabrousrhizomorphs.
Spores 6.3-7.7 X 2-2.5(-3) u, ellipsoid-oblong,smooth, hyaline, inamyloid. Hymen-
ium: basidioles fusoid; cystidia none; cheilocystidia like epicuticularbroom cells but
more often hyaline. Hyphae weakly pseudoamyloid, almost inamyloid in hymenophoral
trama and the interior of the rhizomorphs. Cortical layer: epicutis of pileus hymeniform
broom cells of the Siccus-type, main body 11-26 X 7-8c, clavate, with hyaline to mell-
eous thin to thick (1.5 ) wall, with melleous to reddishbrown setulae, these acicularto
rod-like,acute to obtuse, 4.5-6.5 X 1-2.
On and around dead branchesof trees, French Guyana.
MATERIAL STUDIED. FRENCH GUYANA.Cayenne,Leprieur(K),partof type.

86. MarasmiusrubromarginatusDennis, Trans. Brit. Myco. Soc. 34: 415. 1951.

TYPE. Deznnis 339, from Venezuela.
Pileus carmine-purpleto brown-red("caldera,"M/P or more red), dried bay-mahog-
any ("falcon, muskrat,""leafmold" "Rembrandt"M&Por even deeper on disc), without
a white zone or with a very narrowone or with a purple-blackzone around the center,
sulcate, glabrous, convex, with umbilicus showing a flat bottom, or with a small papilla
or even a prominent one in the umbilicus (which may be absent in very young material),
2-12 mm broad. Lamellaewhitish, eventually pale buff, or cream color from the be-
ginning,with red, eventuallyoften brown edge, distant (11-15 through-lamellae,no lame-
llulae), broad, collariate(collarium not discolorous). Stipe umber, dark gray, brown, or
black, setiform, glabrous,smooth, insititious on the substratum, 13-40 X 0.1-0.3 mm;
thin black rhizomorphsmostly accompanyingthe stipes but not constantly so, and
stipe never rising from rhizomorphs. Context white, thin, inodorous.
Spores (9-)12.5-18.5 X (1.5-)2.5-3.3,, claviform-fusoidto lanceolate, somewhat
curved at times, smooth, hyaline, inamyloid. Hymenium: basidia 17-28.5 X 5-9,u, (2-)
4-spored;basidioles fusoid; cystidia none; cheilocystidia like epicuticularbroom cells
but sometimes smallerand narroweron the averageand more hyaline ones intermixed.
Hyphae of the pileus tramahyaline, 2-20, broad, partly weakly to strongly pseudoamy-
loid, pseudoamyloid in the apex of the stipe, there parallel,with clamp connections.
Cortical layer: epicutis of pileus hymeniform, consisting of broom cells, these of the
Siccus-type, main body 9-19 X 6-13,, clavate or subvesiculose,often bilobed, few hy-
aline, most often reddish ochraceous to ochraceous or pale golden to deep melleous,
with thin to slightly thickened (up to 1 ,) wall which is inamyloid or pseudoamyloid,
with setulae concolorous or brown rarely hyaline, 2-6.5 X 0.5-1.81u,conic, fewer rod-
shaped, subotuse to subacute, rarely few or even none, but mostly many; hypodermium
subhyaline to golden melleous, often more or less pseudoamyloid. Stipe without tegu-
On fallen leaves of dicotyledonous trees in forest, known host: Ficus doliaria.
MATERIAL STUDIED. VENEZUELA. Maracay: Rancho Grande, Mt. Periquito, 22 Nov
136 Flora Neotropica

1949, Dennis 339 (K) type. ECUADOR. Napo, Lago Agrio, 16 May 1973, SingerB 7512 (F).
BOLIVIA. Beni: Vaca Diez, 22 km east of Riberalta,near Arroyo Florida, 7 Apr 1956, SingerB
2565 (LIL). BRAZIL. Pernambuco:Camaragibe,7 Jul 1960, SinlgerB 3282 (BAFC);B 3250
(BAFC);6 Jul 1960, SingerB 3125 (BAFC).
There is a puzzling inconsistency in the reaction of the hyphae with the Melzer's
reagent. Collections B 3250 and B 7512 showed distinctly pseudoamyloid pileus-trama,
B 3282 only in the zone underneaththe epicutis. Marasmiuns martlae has slightly broad-
er spores and more lamellae;it is lignicolous.
ILLUSTRATION. Singer (1965) pl 9, fig 35; Dennis (1951a) pl 19, fig 4.

87. MarasmiusmarthaeSinger, Sydowia 12: 137. 1958.

TYPE. Singer T 1730, from Argentina.
Pileus deep purple red ("maroon" to "carbuncle,"almost "Piccadily"M&P)deep
sulcate, glabrous, deeply umbilicate, sometimes purple black in the umbilicus but neither
black dotted nor papillate, and without a white or pallid zone around the central por-
tion when fresh, disc blackeningin dried condition, convex, 4-8 mm broad. Lamellae
pure white, with deep purple edge, subdistant(16-17), equal, broad, ventricose, collar-
iate. Stipe pure white when very young, soon becoming black from base upwards,all
black in mature dried material,smooth and glabrous, shining, setose, equal, rarely
slightly broaderat base, insititious on the substratum,20-35 X 0.2-0.3 mnm.Context
very little, concolorous under the epicutis, otherwise white in the pileus, at first white
in stipe.
Spores (8.3-)14.5-16 X 4(-4.3) c,, thin-walled, smooth, hyaline, inamyloid. Hv-
menium: basidia 20-34 X 5.5-7/u, 4-spored;basidioles fusoid; cystidia, none: cheilocy-
stidia like epicuticularbroom cells. Hyphae with clamp connections, in pileus moder-
ately pseudoamyloid, in stipe strongly pseudoamyloid. Cortical layers: epicutis of
pileus hymeniform, consisting of broom cells of the Siccus-type, 13.5-29 X 6.8-17.2
p, hyaline, often forked or divided in the mannerof Clavariclla,beset with apical
erect setulae which are obtuse, melleous or melleous-brown,2.5-5.5/ long.
On pieces of wood of dicotyledonous trees in garden in subtropicalzone.
MATERIAL STUDIED. ARGENTINA. Tucuman: Capital, Garden of Institute Migucl Lillo,
14 Jan 1952, Martha Sinlger & Singer T 1730 (LIL), type.

88. MarasmiussanguirotalisSinger in Singer & Digilio, Lilloa 25: 19.1952.

88a. Marasmiussanguirotalisvar sanguirotalis

TYPE. Sitlger, T 1240, from Argentina
Pileus dark purple with purplish black or black center, glabrous,sulcate, convex,
2-5 mm broad. Lamellaewhitish or cream white, edges concolorous with sides or
slightly purplish in the immediate neightborhood of the marginof the pileus, distant
(11), broad collariate. Stipe umber with whitish apex, smooth and glabrous, shining,
insititious, 15-32 X 0.1-0.2 mm. Context very thin, inodorous.
Spores 14.3-20 X 3-4 ,, thin-walled,hyaline, smooth, inamyloid. Hymenium:
basidia clavate: cystidia none; cheilocystidia mostly hyaline, fewer colored as the epic-
uticular broom cells, otherwise like these. Hyphae of trama of pileus with clamp con-
nections, inamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of broom
cells of the Siccus-type, with main body 12-26 X 6.8-12, and yellowish melleous to
melleous brownish, with erect apical setulae 2.7-4.2(-7) , long and brownish, obtuse.
On rotting fallen small branches and on wood in marginal and subxerophytic
woods, solitary.
Marasmius 137

MATERIALSTUDIED.ARGENTINA.Tucuman:Quebradade Lules,18 Feb 1951,SingerT1240

(LIL,MICH),type (LIL-portionfragmentaryanddeteriorated);NearTapia,T 1233 (LIL,paratype.
In some specimens, under a good dissecting microscope, there are traces of dis-
colored (red) gill edge in parts of severallamellae while others are free of them. The
difference between this (type) variety and the following variety lies in this character
as well as in the inamyloid pileus-tramaof the former and in the habitat.

88b. Marasmiussanguirotalis var paucilamellatus(Singer) Singer, comb nov

M1arasmiusnmarthaevar paucilamellatus Singer, Sydowia 12: 138. 1958.
TYPE..Singer T 2190, from Argentina
Pileus deep Venetian red to blackish purple ("tanagra,""carbuncle"with "chippen-
dale" center, later "couldron," "Egyptianr.," or between "tanagra"and "mascara"
to "copperleaf"M&P),with a black central dot and dried sometimes black papillate,
or black umbonate (small low umbo) 2.5-10 mm broad, otherwise as type variety.
Lamellaewhitish to ocher, more or less strongly purple margined,about 2.5 mm.
broad, distant (11-13), equal, collariate. Stipe gray when very young, then umber to
blackish, 25-38 X 0.2 mm, otherwise like type variety.
Spores 16-21.5 X 3-4 p, mostly about 17 7 long, otherwise like type variety.
Hymenium and covering layers like in type variety. Hyphae of trama of pileus weak-
ly but distinctly pseudoamyloid.
On roots of Eugenia sp, and on bark and other woody matter in subtropical
forest and in gardensin the subtropicalzone.
MATERIALSTUDIED. ARGENTINA.Tucuman:Capital,20 Feb 1955, Singer T 2190
(LIL), type; SierraSan Javier,ParqueAconquija,17 Feb 1955, T 2179 (LIL), paratypes;Ciudad
Universitaria,15 Dec 1956, T 2853 (LIL); 1 Jan 1960, T 3471 (LIL).
ILLUSTRATION. Singer (1965), pl 9, fig 34.

89. MarasmiusmicrodendronSinger, Sydowia 18: 349. 1965. Fig 55.

TYPE.Singer B 3448, from Brazil
Pileus pink, dried less brightly purplish-pink(5 J 7, M&P),glabrous,sulcate, convex,
with (concolorous) umbilicus or central depression,with a low dark purple papilla in the
umbilicus, 0.5-1 mm broad. Lamellaewhitish, with whitish edge, distant (8-9 through-
laiellae, no lamellulae), ratherbroad, collariate. Stipe stramineous-umber,ratherlight
colored, insititious and without nodes rising from rhizomorphs,with sparsehyaline hairs,
1-2.5 X 0.05-0.1 mm; rhizomorphsdendroid-ascendantand somewhat thicker than stipes,
30-35 mm long, otherwise like stipes but eventually glabrescent. Context very thin, in-
Spores 7.5-8 X 2.8-3 p, elongated, smooth, hyaline, inamyloid. Hymenium: cyst-
idia none; cheilocystidia 11-21 X 6-10, hyaline, otherwise like the epicuticularbroom
cells, type (1). Hyphae weakly but distinctly pseudoamyloid in pileus-trama,more
strongly so in the stipe. Cortical layer: epicutis of pileus hymeniform, broom cells in
size like the cheilocystidia, also of the Siccus-type, of two types, (1) vesiculose to vesi-
culose-clavate,sometimes forked with obtuse forks, with moderately thick wall, pinkish
or pink with subhyaline, obtuse setulae (1.2-2.8 X 1-2 p), (2) vesiculose-ventricoseto
broadly clavate, usually entire, with thicker (up to 1.7 p) melleous wall, with apical muc-
ro or with severalcoarse apical appendages,but more frequently without appendages
and broadly rounded above.
On dead dicotyledonous leaves in rain forest.
MATERIALSTUDIED. BRAZIL. Amazonas,30 km N of Manaus,19 Jul 1960, SingerB 3448
138 Flora Neotropica

90. Marasmiushippiochaetes Berkeley, Hook. Jour. Bot. 8: 139. 1856.

TYPE.Spruce, from Brazil.
Pileus bright red ("Sultan" M&P),dried blood red to purple red ("vineyard,"M&P
4 K 7, etc.), glabrous,hemisphericto almost cubic then convex with the disc applanate
or shallowly umbilicate and about half as high as broad, with a minute papilla in the ui-
bilicus, umbilicus usually white with the papilla white or light gray in dried material,
sometimes blackish in fresh material, the white umbilicus sometimes becoming sordid,
sulcate to disc, 0.8-3.5 mm broad. Lamellaewhite with red edges and red collarium bor-
der, more rarely either collarium or edges not red-bordered,distant (6-12), broad (half
as broad as radius of pileus), collariate. Stipe pale umber to succineous-umberwhen
fresh, dried sometimes paler (almost stramineous),shining, but not glabrousin well pre-
served material, except sometimes at the originallywhite apex, with minute setae pilose,
glabrescentand smooth, arising from a minute disc-like node (from a rhizomorph)or
more rarely from the substratum,usually severalcarpophoresbranchingoff a rhizomorph
in botryoid arrangementor unilaterally(the latter if rhizomorphis creeping or ascending)
3-120 X 0.1-0.2 mm; rhizomorphslike the stipes, also pilose, glabrescentramose, often
very long, often markedat intervalswith a series of nodes (not so in Bolivian material!),
numerous,generally as thick or thicker than the stipe. Context inodorous, very little.
Spores (5-)7-9 X 3.5-4.2 y, ellipsoid to ellipsoid-oblong,smooth, hyaline with mod-
erately thin walls and not collapsing once they have reached full maturity (and then 7.5-
9 X 3.7-4.2 ), inamyloid. Hymenium: basidia 4-spored;basidioles fusoid; cystidia none;
cheilocystidia like epicuticularbroom cells. Hyphae in pileus trama filamentous, thin-
walled, slowly distinctly pseudoamyloid, in stipe apex strongly pseudoamyloidand par-
allel. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of the
Siccus-type, main body cylindric to clavate or vesiculose-oval,lobate-setuloseat apex
12-25 X 6-13 u, most pinkish purple red when studied in ammonia mounts of dried
material,very few hyaline or subhyaline, thick-walled(wall about 1.3 p thick), rufous
in Melzer'sreagent, setulae coarse, very rarely none, or few, and then cells appearing
seta-like cylindric and obtuse at their tips, 2-8.5-(14.5) X 1.5-4 p, concolorous with main
body; Seta-like hairs of the stipe and the rhizomorph 100-200 X 5.5 filiform, gradually
somewhat tapering towards the obtuse tip, pale melleous, smooth, thick-walled(wall
about 1.5 p thick).
On leaves and their petioles fallen from dicotyledonous trees to the ground in
forest. South Americanrain forests.
MATERIALSTUDIED.SURINAM.Bartica,20 Aug 1937, Sandwith1580 (K). VENEZUELA.
Bolivar,Chimautamassif, 1100-1700 m alt, 30/31 May 1953, Stev'ermark15605 (NY). BRAZIL.
Amazonas:Rio Uaupes, 30/31 May 1853, Spnice (K), type. BOLIVIA. Beni: Vaca Diez, Guayar-
amerin,9 Mar 1956, SingerB 1762 (LIL); 14 Mar1956, SingerB 1996 (LIL); 17 Mar 1956, Singer
B 2069 (LIL).
ILLUSTRATION.Dennis (1951b), fig 9 (above).

91. MarasmiusxerampelinusSinger, sp nov Fig 56.

TYPE.SingerM 8961, from Mexico.
Pileo purpureovel purpureo-rubro,umbilicato, in umbilico saepe papillato,3-10 mm
lato; lamellisalbis, ad aciem albisvel prope marginempilei hoc subconcoloribus,distantibus,
latissimis,collariatis,stipite nigrovel nigro-brunneo,acie albo demumambrino, glabro;rhiz-
omorphispraesentibussed stipiti haud affixis. Sporis 8.5-11 X 5.5-7.5 ,; hyphis tramatis
pilei et hymenophoripseudoamyloideis,fibulatis;elementis epicuticularibuseis Marasmii
sicci analogis,nonnulliscrassitunicatis,nonnulispigmentatis. Ad lignum emortuumin
Marasmius 139

Mexico. Typus in F conservatusest.

Pileus "India red" to "vineyard"or "chianti" to "akhbar,"M&P,i e purple to
purple red, blackish purple when dried, with concolorous umbilicus or the latter somewhat
darker,glabrous, sulcate, convex, umbilicate, with or without a small papilla in the umbil-
icus, 3-10 mm broad. Lamellaewhite, with white edges or the edges somewhat concolor-
ous with the pileus near the marginof the pileus, dried sometimes partly brown, on the
inside of the collarium partly or entirely brown, distant (about 11 through-lamellae),
very broad, collariate. Stipe black or very deep brown with white, later umber brown
apex, setiform, glabrous,shining, insititious on the substratumbut accompaniedby some-
what thinner (than stipe) black rhizomorphscreeping on the substratum(10-19 X 0.3-
0.4 mm). Context white, inodorous.
Spores (6-)8.5-11 X (4-)5.5-7.5 p,, ellipsoid to short ellipsoid, smooth, hyaline, in-
amyloid. Hymenium:basidia 26-33 X 9-11 p, 2-4-spored,rarely one or another 5-spored;
basidiolesfusoid; cystidia none but there are occasionally some cheilocystidia(broom
cells) away from the edge proper;cheilocystidia not makingthe edge strictly heteromor-
phous but intermixed with basidia and basidioles, near marginof pileus mostly like the
epicuticularbroom cells, otherwise similarbut hyaline. Hyphae of the pileus-tramaand
the regularhymenophoraltrama hyaline, with clamp connections, not gelatinized, strong-
ly pseudoamyloid, those of the stipe parallel,also strongly pseudoamyloid. Corticallay-
er: epicutis of the pileus hymeniform, broom cells of the Siccus-type, main body 10-23
(-27.5) X 7-12 p, mostly clavate, inamyloid or pseudoamyloidwith thin to thick wall
(0.2-0.9 ,) hyaline to reddish yellow or golden cinnamon, the pigmented ones distribu-
ted in such a pattern that the scalp preparationappearsmottled, with hyaline to con-
colorous setulae, these 2.8-6 X 1-2.7 1, rod-shapedor conical and subacute.
On dead wood in tropical forest. Mexico.
MATERIALSTUDIED. MEXICO.Chiapas,between Finca Sospiroand El Pozo, 4 Mar1969,
Singer 118961 (F), type; M 8962 (F).

92. MarasmiushioramiMurrill,North Amer. Flora 9: 256. 1915.

92a. Marasmiushioramivar hiorami

TYPE.Hioram, comm. Johnston, from Puerto Rico.
Pileus fresh red with darkerumbilicus zone, without a white or cream colored or
at all paler zone around the central papilla, glabrous, sulcate over half the radiusof the
pileus, convex to campanulate-cylindric(more the latter shape when young), drying (and
in the herbarium)a deep rust color with dark brown center, 2-3(-4) mm broad. Lame-
llae apparentlywhite or whitish when fresh but darkeningsomewhat in the herbarium,
with not discolorous edge, distant (about 12 through-lamellae),ratherbroad, adnate to
an indistinct or narrowcollarium. Stipe fuliginous to blackish, at first with white apex,
setiform, smooth, glabrous,insititous on the substratumaccompaniedby but not attach-
ed to black rhizomorphsand sterile stipes ("telepodes") 20-30 X 0.1-0.2 mm. Context
Spores (8.5-)10-13.5 X (3.3-)3.5-4.2(-4.5) M, oblong, smooth, hyaline, inamyloid.
Hymenium:basidia 23 X 6.3-7.8 M' cystidia none; cheilocystidia like epicuticularbroom
cells but less pigmented. Hyphae of the pileus trama inamyloid or vaguely pseudoamy-
loid in parts; those of the apex of the stipe deep purple, pseudoamyloid. Corticallay-
er: epicutis of the pileus hymeniform, broom cells of the Siccus-type, main body 9-15
.2 X 6.8-11.3 u, hyaline, later some assumingthe same golden melleous to succineous
140 Flora Neotropica

pigment as the setulae; setulae 1-3.2 X 0.5-1.2 , golden melleous to succineous, brown
in the Melzer'sreagent, not initially red; obtuse, sometimes forking;on the umbo of the
pileus there are some irregularlyclavate broom cells with thick walls turningvinaceous
in Melzer's,with few setulae or these replaced by rounded, lobulate protuberancesor
simply uneven above.
On dead tough leaves of dicotyledonous trees and shrubs. Puerto Rico.
comm. Johnston
Nov 1912 (NY), type.
ILLUSTRATION. Dennis (1951c), fig 8.
The color of the pileus was annotated as red by Johnston who says that he has
collected the same (?) fungus also on Polypodialesand sugarcane, dicotyledonous
branches,etc. The following variety agrees with the type in almost all characters,
certainly in all important charactersseparatingthis species from the following related
species exceptingthe spore length.

92b. Marasmiushiorami var austroamericanusSinger, var nov

TYPE. Singer B 2499, from Bolivia.
Sporis minoribusa Marasmiohiorami var hiorami differt. Typus in LIL conserva-
Pileus purple red ("Algerianr.") or pale purple red at first or even orange red
("chinook" M&P)or reddish brown ("cocoa" M&P)when wet but when drying in situ
becoming purple red ("chianti" to "Kazak"M&P)when dried in the herbariumtend-
ing to become deep ferruginousor red-brown("Mandalay"M&Por rustier), the center
at first paler or concolorous, later much darkerin most specimens,glabrous,sulcate
over half the radiusor somewhat wider, convex, soon umbilicate. with a prominent
papilla which is soon the center of the umbilicus, eventually sometimes relatively small
(papilla appearingas a small umbo), 2-3 mm broad. Lamellaewhite, later discolored
buffy or brownish, with the edge concolorous or near marginred to brown or vaguely
intermittently red (dried) mostly not at all discolorous (edge at first white), subclose
to distant (7-13 through-lamellae),broad, narrowlyto widely and distinctly collariate.
Stipe succineous to chestnut-fuliginousor black, at first with white apex, shining, seti-
form, glabrous,smooth, insititious on the substratum,accompaniedby but not attached
to "telepodes" and/or black rhizomorphswhich are thinner than the stipe, stipe 12-35
X 0.1-0.2 mm, mostly about 17-20 mm long. Context hyaline-white,very thin, inodor-
Spores 7-11 X 3.5-4.5 p, ellipsoid-oblong,smooth, hyaline, inamyloid. Hymenium:
basidia 22 X 7.2 A, 4-spored, sometimes some with pseudoamyloidcontents: basidioles
fusoid; cystidia none but some cystidioles like sterile basidioles and occasionally forked
sometimes present;cheilocystidia like the epicuticularbroom cells but most of them hy-
aline (pigmented only in the neighborhoodof the marginof the pileus or on scarce pig-
menteddots along the edge where this discolorationis present). Hyphaeof the pileus-
tramahyaline, with clamp connections, inamyloid, here and in the regularhymenophoral
tramafilamentous and thin-walled,hyphae of the stipe parallelmore or less pseudoamy-
loid. Corticallayer: epicutis of the pileus hymeniform, broom cells of the Siccus-type,
main body 9-25 X 6-12 ,, clavate, vesiculose, ventricose or broad above and pedicellate,
often shaped like Ramariacarpophores,hyaline to brown (melleous, ochraceousbrown),
near marginof the pileus thin-walled,with setulae apical, erect (but in some cells with
somewhat divergentsetulae making transition to the Rotalis-type) 2.7-5.5 X 0.8-2 p, rod-
Marasmius 141

like and mostly obtuse, more rarely attenuated upwards,sometimes forked, all or most
(with a few hyaline) pigmented melleous to rather deep warm brown but not red (not
even partly and initially); among these, particularlyin the center of the pileus some
thick-walledand more strongly pigmented(but not red) broom cells with variableshape
and fewer setulae, even no setulae at all, at the apex.
On dead stems and mostly leaves of dicotyledonous trees fallen to the ground but
also on monocotyledonous (e g musaceous) leaves. South America.
MATERIALSTUDIED. BRAZIL. Guapore,Guajaramirim, 10 Mar1956, SingerB 1820
(LIL). BOLIVIA. Pando: Madrede Dios. Las Piedras,5 Apr 1956, SingerB 2499 (LIL), type;
B 24995 (LTL);Beni: Vaca Diez, Guayaramer'n,8 Mar1956, SingerB 1704 (LIL);La Paz: Nor-
Yungas,CarmenPampa,26 Feb 1956, SingerB 1483 (LIL).
ILLUSTRATION.Singer(1965) pl 10, fig 40 (as AI.aripoensis).

93. Marasmiusaripoensis(Dennis) Singer, Sydowia 18: 188. 1965.

trichorhizusvararipoensisDennis,Trans. Brit. Mycol. Soc. 34: 417. 1951.
TYPE. Deinnis231, from Trinidad.
Pileus mahoganyred (illustration correspondsto "mohawk"M&P)with prominent
black umbo which is later in a shallow depression, otherwise convex, about 2 mm broad.
Lamellaepale buff with not discolorous edge, distant (about 8 through-lamellae),broad,
collariate. Stipe black and polished throughout, about 27 X 0.1 mm. Context thin,
Spores 6.8 X 3 #, elliptical (appearingto be oblong) smooth, hyaline. Cortical
layer: epicutis of pileus consisting of pear-shapedbroom cells about 7 u across, with short
dark red processes.
On fallen leaves of Inga and other trees in forest. Trinidad.
MATERIALSTUDIED.(seen but not sectioned). TRINIDAD. Cerrode Aripo about
2000 feet alt, 23 Oct 1949, Dennis 231 (K), type.
No black rhizomorphsare mentioned and the dark red setulae of the epicuticular
broom cells as well as the much smaller spores would differentiate this from the preced-
ing species with which Singer(1965) identified it earlier.It differs fromMarasmius
edwallianusformasimplex in the colors of both pileus and lamellaedges. It appears
to be closest to Marasmiusputtemansii(the type of which was seen by Dennis) but
differs in the absence of a creamor white zone aroundthe papillawhich is also more
prominenthere than in that species. Furthermore,the sporesappearto be different
from those of all forms of M. puttemansiiknown to me and the absenceof black
rhizomorphswould also distinguishthis species.
It is difficult to judge whether the color of the marginof the pileus is similarto
that of Ml.putteiiansii since a purplishred stage in fresh condition, characteristicfor
both Ml. hioraiii and M. puttenmansii,is not mentioned by Dennis. On the other hand,
Dennis (1951a) drawsa pallid line below the papilla (pl 19, fig 16)which may or may
not indicate the presence of a pallid zone in that stage. At present anlduntil further
fresh materialhas been collected and more spores been measured,1 maintainM. ari-
poeilsis as a species independentof M. puttemanssiibut cannot exclude the possibility
that it might be a variety of the latter.
ILLUSTRATION. Dennis (1951a) pl 19, fig 16.

94. Marasmiusputtemansii Hennings,Hedwigia43: 203. 1904.

oligocladusSinger,Sydowia 18: 190. 1965 ad interim(not validlypublished).
142 FloraNeotropica

94a. Marasmiusputtemansiivar puttemansii

puttemlansiiHenningsvar subtricliorlhizus
Marasmiuls Singer,Sydowia 18: 352. 1965.
TYPE. Puttemans 860, from Brazil.
Pileus "briarwood"M&P(purple-redto red-brown)when dryingin situ often be-
comingbrown with a whitish to creamzone originatingin maturespecimensbecause of
crackingof the surfacelayer arounda gray to blackishdot (papilla)but this pallid
zone often only on one side of the papilla or with minute "briarwood" spots or ob-
solete in the herbariumbecoming reddish brown or bay at least in part, sulcate, convex,
soon umbilicate with or without a small umbo or ratherlow to medium sized papilla
which is eventually in the middle of the umbilicus, 2-5 mm broad. Lamellaewhite or
becoming cream, with white edges but often with indistinct, intermittent, discolorous
(color of the pileus) spots on the edges becoming more distinct when seen in dried
condition under a lens, distant (10-13 through-lamellae,no lamellulae),broad, collar-
iate but the collarium sometimes indistinct, narrowand (dried) appressedto the stipe.
Stipe dark brown to brown-blackwith white apex, glabrous, smooth, insititious either
on the black rhizomorphswhich often accompany the carpophoresor directly on the
substratum,insititious with or without a basal sheet or node, 8-80 X 0.2 mm. Con-
text white, thin, inodorous.
Spores 6.3-9 X 2.3-3 p, mostly about 7-8 X 3 1, oblong or tear-shaped,smooth,
hyaline, inamyloid. Hymenium:basidia 17-18.5 X 5-6 ;' cystidia none; cheilocystidia
like the epicuticularbroom cells but often most of them hyaline. Hyphae of the pi-
leus-tramahyaline, with clamp connections, from slowly and weakly to distinctly pseu-
doamyloid; those of the stipe parallel,pseudoamyloid. Cortical layer: epicutis of the
pileus hymeniform, broom cells of the Siccus type, either entirely brown or red (after
a while becoming melleous in alkali) or only the upper portion of the main body tus-
kan red or else only the setulae brown or red and rarely some cells entirely hyaline,
main body 6-19 X 5-11 $,, vesiculose, clavate, rarely ampullaceous,setulae 2.5-8 X
0.5-1.6 u, conic-obtuse or cylindric-rodshaped,more rarely subacute.
On dead leaves and petioles or small twigs and stems of dicotyledonous trees
(especially Clusiaceaeand Myrtaceae,in Brazil) west to the upper Amazonas.
MATERIALSTUDIED. BRAZIL. SSo Paulo, Serrada Cantareira,21 Mar1904, Puttemnans
860 (RBR) part of the type collection; Pernambuco:Camaragibe,7 Jul 1960, Guanabara,Rio de
Janeiro,JaraimBotanico, 28 Jan 1961. SingerC 3165 (F). ECUADOR. Napo, Shushufindi,13
May 1973, SingerB 7394 (F).
There is a photograph of the type specimen at USDA herbarium in Beltsville (BPI).

94b. Marasmiusputtemansiivar oligocladus Singer, var nov

oligocladusSinger,Sydowia 18: 190. 1965 ad interim.
TYPE. Singer B 2330, from Bolivia.
Sporis latioribusa typo specieis differt. Typus in LIL conservatusest.
Pileus reddishbrown to rust ("copper br." to "Alamo" M&P)when wet, drying to
purple red ("Kazak" M&P),dried deeper (e g "Maracaibo"M&P),with sordid pallid
uneven center, with or without a blackish dot in the center, sulcate, glabrous,convex
with at first steeply declivous margin,later sometimes applanate,usually becoming um-
bilicate in age or umbilicate already in the young state, with or without a very low
papilla or small umbo, 2-6 mm broad. Lamellaewhite or whitish, drying cream, white
or with a very thin pigmented (concolorous with margin of pileus) line, often intermit-
tently so along the edges, distant (8-1 1 through-lamellae, no lamellulae), broad, collar-
iate but collarium at times indistinct, (narrow and partly appressed to the stipe). Stipe
black, smooth, glabrous, shining, insititious, either rising from a black rhizomorph or ris-
Marasmius 143

ing insititiously directly from the substratum,in the first case attached to a disc-like
node or ring, 15-50 X 0.1-0.2 mm. Context white, thin, inodorous.
Spores 6.5-8.5 X 3.3-4.5 s, mostly 7-7.7 X 3.5-(4) ,, ellipsoid to ellipsoid-oblong,
smooth, hyaline, inamyloid. Hymenium:basidia 17-25.5 X 5.5-6 u, 4-spored;cystidia
none; cheilocystidia like the epicuticularbroom cells but mostly hyaline. Hyphae of
the pileus-tramahyaline, with clamp connections, with thin or ratherthin wall, dis-
tinctly pseudoamyloid;hyphae of the stipe parallel,pseudoamyloid. Corticallayer:
epicutis of the pileus hymeniform, broom cells of the Siccus-type, main body clavate
and sometimes above broadenedlaterally, more rarelyventricoseof ampullaceous9-16
X 3.5-7.5-(14) , with often slightly thickened wall (0.5-1 A), hyaline to cinnamon and
often red at the apex (but becoming melleous after a while in alkali medium), with
setulae 1.8-5.5 X 0.7-1.8 ,u, equal or attenuated to an obtuse tip, cinnamon to purple
or brownish purple (but becoming melleous in alkali), in some cells yellowish to hyaline.
On dead leaves of dicotyledonous trees (e g "Pau Ferro"). Brazil.
MATERIALSTUDIED. BOLIVIA. Beni, Vaca Diez, Riberalta,29 Mar1956, SingerB
2330 (LIL), type. BRAZIL. Pernambuco:Camaragibe,7 Jun 1960, SingerB 3251 (BAFC).
In both varieties(which differ mainly in the breadth of the spores), the number
of stipes attached to a rhizomorphsratherthan the substratumvaries from zero to
the majority but there are always many stipes directly attached to the substratum;in
some collections only the latter can be found. In view of the inconstancy of this
characterin this group, I now combine var subtrichorhizuswith var puttenansii.
ILLUSTRATION. Singer (1965), pl 10, fig 42.

95. MarasmiusedwallianusHennings,Hedwigia39: (135). 1900.

TYPE. Edwall, from Brazil(holotypus lost, syntypus see below).
Pileus blood-red, remainingso or fading to brown-redin the herbarium,glabrous,
sulcate, cubic to hemisphericalor campanulate,papillate, often flattened around the
papilla, 0.8-2.5 mm broad. Lamellaewhite with red ege, distant, (about 11 through-
lamellae, no lamellulae), broad, collariate. Stipe pale brownish to bister, shining, in-
sititious either on rhizomorphsor directly on the substratum("forma simplex"), and
then relatively long, otherwise 3-6 X 0.1-0.2 mm; rhizomorphsglabrous. Context
white, very thin.
Spores 7-8 X 3-4.2,, ellipsoid, some oblong, smooth, hyaline, inamyloid. Hy-
menium: basidia 17-21 X 7.2 u, fusoid; cystidia none; cheilocystidia like the epicut-
icular broom cells. Hyphae pseudoamyloid. Cortical layer: epicutis of pileus hymen-
iform, broom cells of the Siccus-type, main body 9-19 X 6.7-13.5 L, hyaline to dark
purple red, often thick-walled,setulae rod-shapedto slightly attenuate to a subotuse
or obtuse tip, purple red to brown-red, 2.3-3.3(-14.5) X 0.6-1.1 ,, mostly 3-3.5 p long.
On fallen dicotyledonous leaves in the forest. Brazil.
MATERIALSTUDIED. BRAZIL. Bahia, Torrend56 (BPI);SSo Paulo, Alto de Serra,
Edwall(FH) syntype;Rio Grandedo Sul, Theissen(FH), the latteroften with white lamella-edge
(W),thus doubtfuL

96. MarasmiusbromeliacearumSinger in Singer& Digilio, Lilloa 25: 212. 1952.

TYPE. Singer T 1457, from Argentina.
Pileus pinkish cinnamon to ferruginous("Maya,""Eldorado"M&P)on the elevated
portion (along the ribs between the sulci or in all the convex portion leavinga pallid zone
around the dark spot in the umbilicus), but dried pileus virtuallyunicolorous ("hazel,"
"harvest"or 13 K 10 M&P)excepting the more ochraceouszone toward papilla(in some
144 FloraNeotropica'`

specimens) and the small central spot or umbo which is brown, the umbo being red when
young and black or blackish when mature in fresh condition, sulcate over the convex
portion, otherwise smooth, glabrous,4-5.6 mm broad. Lamellaewhite or whitish, dis-
tant (9-11 through lamellae), ratherbroad with white or whitish edge and collarium.
Stipe sordid ochraceous brown (13 K 10 to "tortoise shell") to sordid umber, with pal-
lid apex when young, and with deeper colored base when old, in dried condition whole
stipe light brownish, pale cinnamon, or cinnamon, shining, smooth, and glabrous, 26-60
X 0.2-0.3 mm, directly on substratuminsititious; no rhizomorphs. Context extremely
thin, white, inodorous.
Spores 9-11.8 X 4.8-6.2 p, ellipsoid, more rarely ellipsoid oblong, or somewhat
clavate-ellipsoid,smooth, hyaline, inamyloid. Hymenium:basidia 19-19.5 X 6-6.5 ,:
cystidia none; cheilocystidia similarto epicuticularbroom cells, but often few and scat-
tered. Hyphae of pileus filamentous, hyaline, varying from inamyloid to distinctly
pseudoamyloid,with clamp connections. Corticallayers: epicutis of pileus consisting
of elements of the Siccus-type, main body hyaline, subhyalineor golden melleous, some-
times forked at tip and often hand-shaped,clavateto subcylindricif entire. 12-20 X 6-7.
5 P; setulae about 2.5-5.7 u long and 1.5 u broad at base, hyaline to golden, mostly golden
spinulose, sometimes forked at tip; intermixed with these broom cells there are sometimes
some smooth hyaline or golden yellow to melleous vesiculose bodies.
On fallen epiphytic Bromeliaceaein subtropicalforest.
MATERIAL STUDIED. ARGENTINA. Tucuman: Sierra de San Javier. Parque Aconquija at
800 m, Singer T 1457 (LIL), type.
ILLUSTRATION.Singer (1965) pi 10, fig 45.

97. MarasmiusmegalospermusSinger,Sydowia 18: 338. 1965.

TYPE.Singer B 1927, from Bolivia.
Pileus brown to terracotta(13 A 12 to "terracotta"),with a very small and low,
deep colored or gray umbo or dot in a sharply umbilicate center, campanulateto con-
vex with steeply declivous sulcate marginand ratherhigh, 4-10 mm broad. Lamellae
cream color to deep ochraceous,often about 13, with the edges not discolorous and
not quite even, broad, collariate. Stipe yellow succineous to pale sordid umber, smooth
and glabrous,setose, insititious in the substratum,17-22 X 0.1 mm.
Spores 16.5-19.5 X 4-4.2 p, fusoid. hyaline, smooth, inamyloid. Hymenium: con-
sisting of basidia;cystidia none seen. Hyphae of pileus often slightly thick-walled,mod-
erately strongly but distinctly pseudoamyloid,with clamp connections. Corticallayers:
epicutis of pileus hymeniform, broom cells of the Siccus-type, but sometimes slightly
transient towards the Rotalis-type, main body cylindric, clavate or vesiculose, some-
times with geniculate pedicel, hayline or brown, generally thin-walled, 10-14 X 7-9 u,
setulae apical but sometimes obliquely-erectimmediately below apex, brown, 2-4 X
On dead dicotyledonous leaves, in virgin forest.
MATERIAL STUDIED. BOLIVIA. Beni, Vaca Diez, 3 km below Guayarmerin, 12 Mar
1956, Singer B 1927 (LIL), type; B 1928 (LIL), paratype.
ILLUSTRATION. Singer (1965), pi 9, fig 36.

98. MarasmiuschrysocephalusSinger, sp nov

TYPE. Singer B 1936, from Bolivia.
Pileo aureo-flavo,siccando ochrascente,primumconico, papillato,dein in umbilico
minute umbonato, papillaumbonequenigro, 2-4 mm lato; lemellis albis, distantibus,co-
Marasmius 145

llariatis;stipite nigricantevel fuligineo, primumad apicemalbo; rhizomorphispraesenti-

bus sed stipiti haud affixis. Sporis 11.7-15.2 X 3.2-4.2u, elementis epicuticularibuseis
Marasmiisicci analogis. Ad folia delapsanec non ad ramulosminoresin hylaea boliviana.
Typus in LIL conservatusest.
Pileus "Florida gold" M&P(golden yellow), dried ochraceous or ochraceousbuff,
glabrous,sulcate, at first conical with a prominent black papilla,later with a small
black umbo in a small umbilicus, hemisphericto convex, 2-4 mm broad. Lamellae
white, edge not visibly discolorous, distant, 7-12, collariate, ratherbroad. Stipe black-
ish or fuliginous, at first with white apex, smooth and glabrous,insititious on the sub-
stratum, rarely on a black rhizomorph,20-30 X 0.1-0.2 mm; rhizomorphssimilarto
stipe, accompanyingthe fruiting bodies.
Spores 11.7-15.2 X 3.2-4.2 u, fusoid, inaequilateralin profile, smooth, hyaline,
inamyloid. Hymenium:basidia 14-18 X 4.5-5 p,;cystidia not seen; cheilocystidia like
epicuticularbroom cells but a largernumber of hyaline ones and often scattered over
certain distances. Hyphae hyaline to pale melleous, in hymenophoraltramaalmost
inamyloid but slowly and weakly pseudoamyloidin the trama of the pileus. Cortical
layers: epicutis of the pileus hymeniform, consisting of broom cells which are either
hyaline with hyaline or melleous setulae, or entirely deep golden melleous and then
often with slightly to distinctly thick-walledmain body, main body 8-17 X 6-9 M,
clavate, setulae 2.5-4.5 X 1-1.3 ,, obtuse or sometimes acute, generallyquite distinctly
of the Siccus-type but at times slightly divergenton and very near the apex as in M.
On fallen leaves and small twigs fallen to the ground in tropical forest.
MATERIALSTUDIED. BOLIVIA. Beni: Vaca Diez 3 km below Guayaramerin,in virgin
rain forest, 12 Mar1956, SingerB 1936 (LIL), type.
This was formerly(Singer 1965) interpretedas identicalwith Marasmiushiorami
but after our own type studies on the latter it has been recognizedas an independent
ILLUSTRATION. Singer(1965) pl 9, fig 37.

99. Marasmiusxerophyticus Singer, Sydowia 18: 345. 1965.

TYPE.Singer T 2948, from Argentina.
Pileus ferruginous-red-brown ("Kobe" M&P),convex, minutely umbilicate with a
minute umbo inside the umbilicus, the umbo macroscopicallyappearingas a minute black
central dot in fresh material,glabrous,sulcate, without a white or pallid zone around the
black dot, 2 mm broad. Lamellaewhite, edge concolorous with the sides of the lamellae
distant (11), collariate. Stipe gray and paler at apex when fresh, light brown with
almost fulvous apex when dried, glabrousand smooth, insititious on the substratum,13-14
X 0.2 mm, associated with black rhizomorphs.
Spores 13.5-15.8 X 3.5(-4) , accicular-fusoid,smooth, hyaline, iiiamyloid. Hymenium:
basidia 26-30 X 6-7 ,; cystidia none. Hyphae with clamp connections, inamyloid in trama
of pileus. Corticallayers: epicutis of pileus consisting of three types of broom cells of
the Siccus-type and of irregularelements similarto those of the Rameales-type(Maras-
miellus); the latter elongated and erect or curved, with rounded-obtuseirregularnodulose
excrescences, all brownish, without typical setulae, sometimes somewhat thick-walled,of
variablesize; the Siccus-type broom cells 9.5-25 X 4-18 u with erect apical setulae 2-4.5
X 0.7-1.3 1, either (1) all hyaline or (2) hyaline with light golden melleous setulae, or
(3) entirely brownish melleous.
On rotting dicotyledonous wood in subxerophyticto xerophytic woods.
146 Flora Neotropica

MATERIAL STUDIED.ARGENTINA. Salta:Gicmes,lowerpartof Quebrada

de Yaquiasme,
7 Feb 1957,SingerT 2948 (LIL)type.
ILLUSTRATION. Singer(1965), pl 9, fig 38.

100. Marasmiusguyanensis Montagne,Ann. Sci. Nat. Bot. IV. 1: 114. 1854.

00a. Marasmiusguyanensisvarguyanensis
TYPE. Leprieur,from French Guyana.
Pileus yellow buff, yellow ocher, chrome yellow or cadmium yellow, bronze yellow,
("nugget, bronze y.," "daffodil," Spanish y.," "deep chrome," "cadmiumy." M&P)when
dry (in situ) turning orange-goldbrown to ferruginous("Punljab,""Arabianbr.," "ferrug-
inous," "feuille morte" M&P),often collected in this shade when not quite freshly grown,
dried a color between "Punjab"and "feuille morte" or near "Arabianbr." M&Por yellow-
ish brown and sometimes if poorly preparedreachinga sordid brown color in the herb-
arium, fresh with a central black dot which in dried materialappearsas a deep brown to
blackish papilla or small umbo prominent in young materialbut later hidden in a small
umbilicus,without white zone around black dot, but sometimes in fresh condition with
a paler yellow ring around it, glabrous,sulcate outside umbilicus when mature, 1.2-6 mm,
mostly 2-3.5 mm broad. Lamellaewhite, in age whitish to cream, with the edges con-
colorous with sides, in dried materialeventually sometimes entirely subconcolorouswith
pileus, medium broad to broad, mostly broad, distant, 8-11, collariate. Stipe succineous
brown to umber, later black, at first entire stipe white (in buttons when quite fresh),
later only apex whitish to concolorous with pileus, and eventually apex concolorous
with lower portion of stipe, smooth and glabrous,rarelyminutely pruinate(?) at base,
insititious directly on the substratum, 10-40 X 0.1 mm, black rhizomorphsnot accom-
panying the carpophores.
Spores 8.5-14 X 2.5-4 ,, clavate or comma-shaped,smooth, thin-walled,eventually
often collapsing, hyaline, inamyloid. Hymenium: basidia 19-21 X 6-8 , 4-spored, cla-
vate, hyaline: basidioles fusoid; cystidia none; cheilocystidia, broom cells of the Siccus-
type, hyaline to yellowish hyaline, otherwise like the epicuticularbroom cells. Hyphae
of subhymenium,hymenophoraltramaand trama of pileus hyaline in ammonia, with
clamp connections, inamyloid in subhymenium,inamyloid to weakly pseudoamyloidin
hymenophoraltrama and more distinctly (though weakly) pseudoamyloidin pileus trama
accompaniedby some oleiferous hyphae; tramaof pileus a very thin layer, pseudoamy-
loidity noticeable only in accumulationsof hyphae; hyphae of stipe in strictly parallel
arrangement,with numerousclampedsepta, weakly pseudoamyloid. Corticallayers:epic-
utis of pileus hymeniform, consisting of broom cells generally of the Siccus-type but
some with slightly divergentsetulae which descend somewhat from the extreme tip of
the cell and thus vaguely remindone of some epicuticularelements of the Rotalis-
type, clavate, or vesiculose-ventricoseand always broadly rounded, even applanate,often
forked or cauliflower-likedivided in upper portion, 10-25 X 4.8-17 p, either entirely
hyaline or hyaline in lower portion or entirely brownishyellow or melleous to deep
orange, generally thin-walledbut often some thick-walled,setulae 1.5-4 X 0.7-1.5 s,
i e ratheithin at base, rathercoarse,acute, subacuteor obtuse, simple,varyingfrom
hyaline to light yellow to deep golden yellow or at times even brownish yellow. Cells
of the black-dot-regionof the center similarbut either without setulae or these replaced
by broadenedrounded conidia-likeappendages,chestnut brown and thick-walled,form-
ing a palisaderatherthan a hymeniform layer. Stipe surface without a coveringlayer.
Marasmius 147

On dicotyledonous leaves and leaf petioles, more rarely on monocotyledonous

leaves and stems, rarely on small woody sticks of dicotyledons, in tropical rain forest.
Host once determinedas Carapaguianensis (sec. Dennis).
MATERIALSTUDIED. TRINIDAD. ArenaForest, 30 Oct 1949, Dennis 255 (K). FRENCH
GUYANA(K), authentic type ? BRAZIL. Acre: Sena Madureira,7 Oct 1968, Pranceet al 7904 (NY).
VENEZUELA. Sucre: Rio Media,15 Jul 1972, Dumonttet al VE 5174 (NY). BOLIVIA. Pando:
Madrede Dios, Santa Teresa,28 Mar1956, SingerB 2305 (LIL); Las Piedras,5 Apr 1956, B 2527
(LIL); Beni: Vaca Diez, Guayaramer'n,16-17 Mar 1956, SingerB 2040, B 2101 (LIL); 3 km below
Guayaramerfn in virginrain forest, 12 Mar1956, SingerB 1935 (LIL).
It was at first thought that Dennis's interpretationof the species is slightly differ-
ent from the originalconcept. But after collecting a large number of specimens,it was
found that there is enough variationin the shape of the epicuticularbroom cells and the
Melzerreaction of the trama of the pileus to consider the Trinidadand the French Guy-
ana materialextremes of the same species and even variety. The color (umber) of the
pileus as indicated by Montagneand copied by others is probably the color of old herb-
arium specimens, not characteristicfor living or well preparedspecimens.
ILLUSTRATIONS. Singer(1965), pl 9, fig 39; Dennis (1951a) pl 19, fig 17.

10Ob.Marasmiusguyanensisvar erythrocephalusSinger,Sydowia 18: 336. 1965.

TYPE. Singer B 1876, from Bolivia.
Differing from the type variety in the color of the pileus: Pileus orange red ("to-
tem" to "Brazilr" M&P)when quite fresh and bleachinglittle (in age "Nasturitium"M&P)
pallid around a central black dot but pallid zone very narrowand difficult to observe in
dried material,dried pileus fuscous brown. Lamellaewhite. Stipe black with white apex.
Spores 8.3-13 X 2.7-4 u. Epicuticular broom cells with main body 8-17 X 4.5-10 s,
thin-walledor almost so, hyaline to melleous, setulae 2-5.5 X 1-1.3 u, rarelyhyaline, most-
ly light golden melleous attenuate upwardsbut tip itself obtuse in most cases, rarely
Otherwiselike the type variety.
On dead dicotyledonous leaf in virgin rain forest.
12 Mar1956,
MATERIALSTUDIED. BOLIVIA. Beni, Vaca Diez, 3 km below Guayaramerrn,
Singer B 1876 (LIL), type.
This is a rare color variantof the species, apparentlydue to the slight microscopical
differences evident in the structureof the epicutis.
ILLUSTRATION.Singer(1965) pl 9, fig 39E.

lOOc.Marasmiusguyanensisvar guayarensisSinger, Sydowia 18: 336. 1965.

TYPE. Singer B 2014, from Bolivia.
Macroscopicallydiffering in the fresh color of the pileus which is a deep but pure
orange("bittersweet o." M&P). Microscopicallydiffering in somewhat smaller spores:
8.5-11.5 X 2.7-4 A. On monocotyledonous branch in tropical rain forest. Otherwise
generallylike type variety.
MATERIALSTUDIED. BOLIVIA. Beni, Vaca Diez, Guayaramerfn,16 Mar1956, SingerB
2014 (LIL), type.
This was found only once. It differs more decidedly from the type variety showing
a somewhat different fresh color, and at the same time having smallerspores.

101. MarasmiustrichorhizusSpegazzini,Anal. Soc. Ci. Argentina 16: 173. 1883.

TYPE. Caa-Guazfi,Paraguay(no type materialfound).
Pileus testaceous or rufescent, glabrous,hemispherical,0.5-1 mm broad. Lamellae
148 FloraNeotropica

concolorous, pliciform, few, ?collariate. Stipe dark, somewhat shining, erect, glabrous,
10-20 X 0.15-0.18 mm risingfrom thicker rhizomorphswhich are creeping and form a
very extended (200-500 mm), very loose "rhizomorphoidmycelium," which forms
annularjoints where the stipes are inserted (description adapted from Spagazzini, 1883).
Microscopicalcharacterssee below.
On bark of rotting logs in virgin forest Caa-Guazfi,Paraguay.
Spegazzinihimself says that his materialconsisted of old, somewhat obliterated
specimens which should be recollected and restudied. He indicated later that this species
was recollected in Paraguay(Fiebrig, February 1920 on trunks of Pithecollobium scalare)
but reduced his species to synonymy with Marasmiusequicrinis(M. crinis-equiMueller
ex Kalchbrenner);this species is strongly papillate. Paraguayanmaterial(Piragu,July
1879) is conservedat LPS and does indeeed show a papilla but no sporescould be re-
covered by me or by Dennis (1951a); but Spegazzinifound them to be 7-8 X 4-5 p. It
is indeed probable that the Paraguayancollections refer to Marasmiuscrinisequi. There
is however a further collection determinedMarasmiustrichorhizusby Spegazzini. AR-
GENTINA. Formosa: Dec 1900, Kermes 723 (LPS). This materialhas spores 8.2-9
X 2.2-3.5,, thus narrowerthan indicated by Spegazzini. It has epicuticularbroom cells
of the Siccus-type, 12.3-19.2 X 8.2-11 p, setulae 2.5-3.7 p long, main body pale yellow-
ish, setulae brown. Hyphae of the pileus inamyloid,with clamp connections. The color
of the pileus is now brown but its originalcolor is not indicated.
There are two possibilities. Either the original description is taken at face value
and the later Paraguayancollection only is taken as identical with the type; in this
caseMarasmiustrichorhizuswould become a (lignicolous?)form of Marasmiuscrinis-
equi Or else the Formosanmaterialis taken as authentic and Spegazzini'sfailureto
indicate the presence of a prominent papilla and normal(not fold-like) lamellae is con-
sidered erroneous or exceptional. In this case Marasmiustrichorhizuswould be a good
species, not fully and partially wrongly described. A decision as to which approachto
take, should, I believe, be postponed until an attempt has been made to recollect the
species at the type locality, inasmuchas Spegazzini'sspore measurementssuggest that
the type might have been close to our no. 80 (M. schultesii) if the hairinessof the
rhizomorphshas disappearedor was overlooked.

102. Marasmiuscrinis-equiMuellerex Kalchbrenner,Grevillea8: 153. 1880.

Marasimiusequicrinis Mueller in Berkeley, Jour. Linn. Soc. Bot. 18: 383. 1881.
Marasmiiusgramzintum(Libert) Berkeley & Broome var equicrinis (Berkeley) Dennis, Trans. Brit.
Mycol. Soc. 34: 416. 1951.
? Marasmius repens Hennings, Engler Bot. Jahrb. 23: 548. 1896.

102a. Marasmiuscrinis-equivar monocotyledonum Singer, var nov

TYPE. Staner C 325, from Zaire.
Sporis usque ad llu longis. Ad monocotyledones. Typus in BR.
Pileus orange with deeper colored papilla, somewhat bleachingoccasionally, in the
marginalregion, glabrous, sulcate, convex, papillate, 1.5-2 mm broad. Lamellaepale
orange-yellowwith deeper colored punctations or stretches along the edge and paler on
the sides, distant, few, moderately broad, collariate. Stipe chestnut to chestnut black,
then black, sometimes at first white at the apex, shining, glabrous,rising from black
rhizomorphs,3-4 X 0.1 mm; rhizomorphsabundant,long, smooth, glabrous. Context
white, very thin, inodorous.
Spores 7.5-11 X 3.7-5 p, ellipsoid to ellipsoid-oblong,few tear-shapedor fusoid,
Marasmius 149

smooth, hyaline, inamyloid. Hymenium:basidia 24.6 X 9,u; cystidia none; cheilocys-

tidia like the epicuticularbroom cells. Hyphae of the trama of the pileus hyaline, fil-
amentous, with clamp connections, inamyloid; those of the stipe + pseudoamyloid.
Corticallayer: epicutis of the pileus hymeniform, broom cells of the Siccus-type, main
body 8-20 X 6.5-9.5 , versiform,hyaline to pale golden or gold brown to pale cinna-
mon, with thin walls, with apical setulae, 1.5-4.3 X 0.5-2.3 ,, hyaline to more often
golden yellow to yellow-brown or cinnamon.
On monocotyledonous stems and culms, leaves, etc, in the tropics of the Wes-
tern and Easternhemispheres.
MATERIALSTUDIED. ZAIRE. Bamania,Eala,P. StanzerC 325 (BR); Angodia,J. Lebrun
3009 (BR),type.
Materialsimilarto the African materialcited above has been collected by Petch
in Ceylon (K) and by Dennis on Trinidad(K) but these collections have been checked
only macroscopicallyby me. They appear to be identical with the African material.
On the other hand, the type collection of M. crinisequi(var crinisequi)also at K has
been revised by Pegler(1965). It differs from the variety describedabove in longer
spores (9-13 X 3.5-5 p) and habitat on myrtaceous leaves and twigs.
ILLUSTRATION. Singer (1964), fig 18, A-C.

103. Marasmiusrobertsonii Singer, Sydowia 18: 342. 1965.

TYPE. Singer B 2349, from Bolivia.
Pileus "terracotta"M&P(orange-ferruginous)dried "cocoa" and towards margin
often dulled to "gold br." i e neither as deep colored as most other rhizomorphogeneous
species nor orange or pallid, with a black dot in the umbilicus when fresh but in dried
condition unicolorous all over the disc, sulcate in a broad marginalzone, at least when
mature, at first campanulateto convex and distinctly papillate, later with a small umbo
in the umbilicus, otherwise convex, 3-6 mm. broad. Lamellaewhite, dried buff with
white edge (dried buff), broad, simple, equal, distant, nine, collariate. Stipe black,
smooth and glabrous,insititious with a minute disc-like node either, only a few cases,
directly on the substratumor in the majority of a population, inserted on a black
rhizomorph,7-20 X 0.1-0.2 mm; rhizomorphsblack, glabrous,smooth or with occas-
ional (1-2) nodes and node-bearingramifications,sometimes insititious in the substra-
tum, sometimes long. Context inodorous.
Spores 10 X 3.5-4.1 ,, ellipsoid-oblongor cylindric, with a slight suprahilar
applanationor depression,hyaline, smooth, inamyloid, occasionally with small wart-
like processes which are the beginningsof germinationon the lamellae. Hymenium:
cystidia none; cheilocystidia hyaline, otherwise like the epicuticularbroom cells.
Hyphae of the trama of the pileus inamyloid, with clamp connections. Corticallay-
ers: epicutis of the pileus consisting of broom cells of the Siccus-type of which there
are two kinds, (a) the majority with thin walls and hyaline to yellowish, with fine
cylindric to spinulose setulae, (b) the minority with thickened walls and yellowish to
cinnamon, with coarser(to 7 X 1.5 , or shorter) setulae.
On dead fallen dicotyledonous leaves in rain forest.
MATERIAL STUDIED. BOLIVIA. Beni, Vaca Diez, Riberalta, 30 Mar 1956, Singer B
2349 (LIL), type.
This species might easily be mistaken for Al. trichorhizusfrom which it apparently
differs in the color of the pileus, and certainly in the habitat (leaves instead of wood),
but also in the spore size which does not coincide with either this author's measurements
on authentic materialof Al. trichorhizus,nor with Spegazzini's.
150 Flora Neotropica

Marasmiusrobertsonii differs from the type variety of Marasmiuscrinis-equiin

smaller spores and colors.
ILLUSTRATION.Singer (1965) pl 10, fig 48.

104. Marasmiusdicotyledoneus (Singer) Singer,comb nov

Marasmiusequicrinisvar dicotyledoneusSinger,Sydowia 18: 336. 1965.
TYPE. Singer T 1854, from Argentina.
Pileus fresh pale cinnamon buff in middle portion, white at margin,dried "cara-
mel" except for a minute deep brown central papilla, 1.3-3 mm broad. Lamellaewhite
with white edge, equal 9-10, collariate. Stipe umber to black, 4-6 X 0.1-0.2 mm, accom-
panied by small, inconspicuous, scanty and thin ihizomorphs, but all stipes insititious
directly on the substratum.
Spores 9.5-12.3 X 3.5-4.7 p. Hymenium: basidia 22 X 7 p, 4-spored;cystidia none.
Hyphae: trama of pileus hyaline, of thin-walledhyphae with clamp connections, weakly
partly pseudoamyloid,hyphae of hymenophoraltrama inamyloid, of apex of stipe
pseudoamyloid. Corticallayers: epicutis of pileus consisting of hymeniform layer of
broom cells of the Siccus-type, main body often branched-cellular,or clavate, hyaline,
setulae in most cells melleous, in few hyaline or chestnut, 2.5-7 p long, (mostly ? 3 p
On dead twigs of Compositaeand Solanaceae(Baccharis?,Solanum?)in sub-
tropical-montane,ratherdry zone.
MATERIAL STUDIED. ARGENTINA. Catamarca, Dpto. Andalgala, Rio Potrero, 1400 m
alt, 10 Feb 1952, Singer T 1854 (LIL), type.

105. Marasmiusrhizomorphogeton(Singer) Singer, comb nov

Marasmius equicrinis var rhizomorphogeton Singer, Sydowia 18: 335. 1965.
TYPE. Singer B 2531, from Bolivia.
Pileus dull orange ochraceous to light golden yellow (11 H 6 to "mellow glow"
M&P)around a minute flat central brown to black dot (not papillate), and gradually
more and more pallid stramineoustowards the marginof the pileus, sulcate except in
the umbilicate center, but in some weakly, irregularlyor shallowly sulcate, in others
strongly regularlysulcate-furrowed,dried pallid with a slight yellowish tinge in many
towards the disc, or between "Saratoga"and 13 K 10 M&P(brownish), glabrous,3-7
mm broad. Lamellaecream, with cream to pallid edge, distant (8-11), broad or med-
ium broad. Stipe umber to succineous, reachingblackish in fresh condition, but dried
mostly umber-succineous,glabrousand smooth, insititious on the substratum,simple,
equal, setose, 19-32 X 0.1-0.5 mm; accompaniedby dark rhizomorphsof the same
charactersas the stipe, however often with one or two nodes and often branching,the
repent ones at times with fibrils, often very long, where arisingfrom substratum,in-
sititious. Context thin, white, inodorous.
Spores 10-13.2 X 3.8-4.2 p, clavate-ellipsoidto oblong, sometimes subcylindric,
thin-walled,smooth, hyaline, inamyloid. Hymenium: basidioles fusoid; cystidia none.
Hyphae of pileus varying from weakly pseudoamyloidto mostly inamyloid with some
strandsof hyphae appearingvery weakly pseudoamyloid in accumulations. Cortical
layers: epicutis of pileus consisting of broom cells of the Siccus-type with some few
slightly transientto the Rotalis-type in occasional collections, i e with setulae slightly
divergingalthough only apical and subapical,main body 12-20 X 5.5-11 u, hyaline,
clavate, ventricose, or vesiculose-pedicellate,setulae 3-3.5 X 0.6-0.8 j,, conic, obtuse
or subobtuse, or subacute, pale golden hyaline to deep golden melleous.
Marasmius 151

On grassculms, sometimes also on tree fern rachis, gregarious,particularlyalong

rain forest brooks but not seen in inundatedforest.
MATERIALSTUDIED. BOLIVIA. Pando: Madrede Dios, Las Piedras,5 Apr 1956, Singer
B 2531 (LIL), type.

105b.Marasmiusrhizomorphogetonforma arrhizomorphicusSinger, f nov

Rhizomorphisnullis. Differs from the type form in the absence of rhizomorphic
MATERIALSTUDIED. BOLIVIA:Pando, Las Piedras,5 Apr 1956, SingerB 2518 (LIL), type.
The rhizomorphlessform is paler than the type form of the variety and it is this
form that was found both on grassculms and ferns. The type form is known only from
ILLUSTRATION. Singer(1965) pl 10, fig 47.

106. Marasmiusgraminum(Libert) Berkeley & Broome in Berkeley, Outlines of Bri-

tish Fungology p 222. 1860.
AgariciusgraminumLibert,PlantaeCryptogamicaeArduennaeII no. 119. 1832.
MarasmiuspruinatusBerkeley& Curtis,Ann. Mag.Nat. Hist. III. 4: 295. 1859.
curreyi Berkeley& Broome,Ann. Mag.Nat. Hist. V. 3: 209. 1879.
tritici Young, Phytopathology25: 116. 1925.
culmisedusSingerin Singer& Digilio, Lilloa 25: 193. 1952.

106a. Marasmiusgraminum var graminum

MIarasmius var brevisporusDennis,Trans.Brit. Mycol. Soc. 34: 416. 1951.
TYPE. Libert 119, from Belgium.
Pileus light orange to rufescent "terracotta"to "carrotr." or 1l-C-l1 M&Pand
fading to ochraceous brown or pallescent in old caps and in the herbarium,with darker,
lateritious, often almost black center and under a lens some spots of the same color as
the center around the latter visible, but without a pallid zone around it, glabrous,long-
sulcate, even plicate, campanulateor convex, papillate or obtuse, soon convex and flat-
tened around the central zone and becoming shallowly umbilicate, 1.5-15 mm broad, in
the neotropics mostly about 2-7 mm. Lamellaewhite to cream, with white edges, dis-
tant (8-15 through-lamellae)broad, but sometimes strongly narrowedor evanescentat
margin,collariate but the collariumoccasionallyvery narrowor adnate to the stipe, in
other carpophoresof the same population broad and leaving a somewhat fibrillose, white
space between the collariumand the apex of the stipe. Stipe reddishbrown or brown,
often becoming black from base upwards,mostly at first white at the apex, shining,gla-
brous, insititious on the substratumand generallynot accompaniedby black rhizomorphs,
equal, 10-30 X 0.1-0.3 mm. Context white, thin, inodorous.
Spores when mature typically of two sizes (1) 6-9 X 3-4.5 y, ellipsoid, (2) 9-12
X 4-5 ,, ellipsoid to ellipsoid-oblongbut frequently one or the other size dominating
or almost exclusive, smooth, hyaline, inamyloid. Hymenium: basidia 18-23 X 7.2-8.3 y,
4-spored;cystidia none; cheilocystidia like the epicuticularbroom cells but mostly all
or almost all entirely hyaline. Hyphae of the pileus-tramamostly filamentous, hyaline,
not gelatinized, slowly and not strongly but undoubtedly at least in their majority pseu-
doamyloid, with clamp connections; those of the apex of the stipe parallel,distinctly
and strongly pseudoamyloid. Corticallayer: epicutis of pileus hymeniform, broom cells
of the Siccus type, main body 11.5-16 X 5.2-111, mostly hyaline to pale golden and
thin to firm-walled,regularlyintermixed with more distinctly pigmentedand thicker-
152 FloraNeotropica

walled cells, all with apical setulae 2-5.5 X 1-1.5 i, these spinulose-obtuse,often some-
what curved, hyaline to more frequently golden melleous.
On grassculms, both living and dead, also on grass roots and leaves, solitary or in
small groups. Known hosts: Triticuni,Agroprymum, Cynodon, Bambuseae,possibly also
on small twigs of other Monocotyledones.
MATERIAL STUDIED. TRINIDAD. St. Joseph, 30 Nov 1949, Dennis 411 (K), type of var
brevisporus; St. Augustin, 16 Jun 1949, Dennis 101 B (K); 101 A (K). MEXICO. Chiapas, between
Finca Sospiro and El Pozo, 4 Aug 1969, Singer M 8956 (F). HONDURAS. Atlantida: Telas, 14 Dec
1927, 15 Mar 1928, P. Standley, det. V. K. Charles (BPI). ARGENTINA. Catamarca: Suncho, Santa
Rosa, 20 Jan 1952, Singer T 1787 (LIL); Rio Potreros at 1400 m alt, 10 Feb 1952, Singer T 1816
(LIL). U.S.A. New England, Sprague (FH), type of M. pniinatus; Florida: Highlands County, High-
lands Hammock State Park, 25 Aug 1942, Singer F 395 (F); Illinois: Prairie W of Highlands Park,
9 Jul 1973, Singer N 4670 (F); Abington, 12 Jul 1924, Young (BPI) type of M. tritici, Minnesota:
Ramsay County, St. Paul, 20 Jul 1928, Christensen (BPI). U.S.S.R. Leningrad, 19-26 Aug 1938,
Singer (LE). FRANCE. Mesnil-Val, Aug 1931, Madamiefi FPraniois (LIL). BELGIUM. Ardennes,
Libert 119 (BPI, LE), type.
The spore size alone cannot serve to- distinguishvarietiesof taxonomic significance
since in the majority of the collections both types of spores are present, even in the
type of var brevisporus Dennis we find a small number of spores up to 11.5 , long.
ILLUSTRATION. Singer (1958a), fig 50.

106b.Marasmiusgraminumvar culmisedus(Singer ini Singer & Digilio) Singer, Sydowia

12: 142. 1958.
Marasmius culmisedus Singer & Digilio, Lilloa 25: 193. 195?.
TYPE. Singer T 313, from Argentina.
Pileus varying from ferruginous("Ginger"to 13-D-12 M&P)to ferruginousrufous
("henna" to "Kobe"), sometimes bleachingin age or at least in the furrows or in spots,
with tan colored to blackish central umbilicus with a knob-like papilla or elevation in-
side the umbilicus, glabrous,sulcate outside the umbilicus, convex, 3-15 mm broad.
Lamellaewhite in age and on drying reachingpale buff, distant, equal, narrowto rather
broad, sometimes not reachingthe margin,moderately distant to distant (9-18), collar-
iate but collarium often not well developed or not fully free. Stipe black with white
to pallid apex, glabrousand smooth, insititious directly on the substratum,central,
rarely slightly eccentric, 6-36 X 0.1-0.5 mm, not associated with rhizomorphs. Context
very thin, inodorous.
Spores(8-)9-12.3 X (4-)4.2-5.3(-7)u, ellipsoid to ovoidor subfusoid, smooth, hy-
aline, inamyloid. Hymenium: basidia 22-35 X 7-9 p, 4-spored;cystidia none; cheilocyst-
idia like epicuticularbroom cells but more hyaline ones present, often irregularand with
few or no setulae. Hyphae in tramaof pileus varyingfrom weakly and only partially
pseudoamyloid( i e some hyphae inamyloid) to inamyloid, hyaline, with clamp connect-
ions. Those of stipe apex strongly pseudoamyloid. Cortical layer: epicutis of pileus
hymeniform, consisting of broom cells of the Siccus type, main body 13-23 X 5.8-111,
obpiriform,piriform or cylindric to clavate, sometimes divided like a Clavariellacarpop-
hore, hyaline or orange-melleousand then thicker-walledthan most cells, setulae 2-12
X 1-3(-4.5)u, mostly around 3-7 , long, mostly all erect, melleous or brownish or golden,
fewer hyaline or subhyaline, obtuse, often irregularor forked, some entire (without
On grassleaves and culms, usually gregarious. SubtropicalnorthwesternArgentina
and north to the Amazonas region.
MATERIAL STUDIED. ECUADOR. Napo, Francisco de Orellana, 12 May 1973, Singer B
7378 (F); lago Agrio, 9 May 1973, Singer B 7317 (F). ARGENTINA. Tucuman, San Pablo, 29
Mar 1949, Singer T 313 (LIL), type.
Marasmius 153

This variety differs from the type variety in less or not pseudoamyloidpileus-
trama, slightly largerspores and the more rusty color.
ILLUSTRATION. Singer (1958a), fig 51.

106c. Marasmiusgraminumvar schini Singer,Sydowia 18: 336. 1965.

TYPE. Singer T 1788, from Argentina.
Pileus "terracotta"(M&P)to salmon orange, old more or less pallescent, glabrous,
sulcate, campanulate,umbonate, then convex and umbilicate with a small papilla in the
umbilicus, 3.5-6 mm broad. Lamellaewhite with white edge and collarium, distant (10-
14 through lamellae, 0-1 lamellulae),broad, collariate. Stipe umber to black with white
apex, glabrous,insititious on the substratum,not associated with rhizomorphs,equal,
4-10 X 0.1 mm. Context white, thin, inodorous.
Spores 7.5-10 X 3.5-5 n, most frequently 8.2-9.7 X 4-4.5 p, nearly twice as long as
broad, ellipsoid to ellipsoid-oblong,smooth, hyaline, inamyloid. Hymenium:basidia4-
spored; cystidia none; cheilocystidia like the epicuticularbroom cells but all hyaline or a
few pale stramineousto golden stramineous. Hyphae of the trama of the pileus hyaline,
mostly filamentous,varying from extremely slowly and weakly pseudoamyloidto inamy-
loid, always some decidedly inamyloid, with clamp connections; hyphae of the apex of
the stipe parallel,moderately strongly pseudoamyloid. Cortical layer: epicutis of the
pileus hymeniform, broom cells of the Siccus type, much like those of var culmisedus.
On roots, branchesand stems of dicotyledonous plants in NorthwesternArgen-
MATERIAL STUDIED. ARGENTINA. Jujuy: Posta de Lozano, 9 Feb 1966, Singer T
5003 (F); Catamarca: Suncho, Santa Rosa, 20 Jan 1952, Singer T 1788 (LIL), type.

106d.Marasmiusgraminumvar subalpinusSinger, Sydowia 18: 336. 1965.

TYPE. Silger S 159, from Argentina.
Pileus "burnt sienna," umbo in an umbilicus and surroundinghighest wall of
pileus deep brown with a pallid zone between the disc and the margin,soon becoming
salmon orange(1 l-E-10) with central deep brown to black dot, at first almost barrel-
shaped and papillate, later relativelylower, convex with central depressionor umbili-
cus, sulcate, 3-5 mm broad. Lamellaewhite with white edge, 8-9, distinctly collariate.
Stipe fuliginous to black, 9-10 X 0.1-0.2 mm; no rhizomorphs. Context thin, inodor-
Spores 7.5-9 X 4.5-4.7 ,, ellipsoid, smooth, hyaline, inamyloid. Hymenium: cheil-
ocystidia like epicuticularbroom cells but hyaline. Hyphae of pileus partly slowly weak-
ly pseudoamyloid,partly inamyloid, hyaline, inequal, with clamp connections. Cortical
layers: epicutis of pileus hymeniform, with broom cells of the Siccus-type, main body
hyaline, 11-20 X 6-10 ;,, setulae broadly cylindric and obtuse or spinulose and mostly
subacute and thinner, 2.5-4.2 X 1-1.5 ,, in some cells hyaline, in others golden stramin-
On dead wood of Baccharissp.
MATERIAL STUDIED. ARGENTINA: Tucuman: Tafi del Valle, Quebradita del Mastil, at 2300
m alt, 7 Feb 1959, Singer S 159 (LIL), type.
This agrees in spores with var brevisporusDennis but differs in the pallid zone of
young specimens, the shorter stipe, fewer lamellae and the habitat.
ILLUSTRATION. Singer (1965), pl 10, fig 50.
154 Flora Neotropica

107. Marasmiuspraecox Singer, Sydowia 12: 140. 1959.

TYPE. Singer T 2033, from Argentina.
Pileus varying from rusty orange to orange rufous ("burnt sienna," "feuille morte,"
4 G/H 12; 5 E 12; 4 A 12; "Monterey,""Nasturtium,""Mosque,""terracotta" M&P
very rarely bleaching somewhat in age) becoming deep reddish orange to brown red when
dry in situ ("Monterey," "henna," "Indian red," "caldera"M&P),dried (completely de-
hydrated to air-dry,so in herbarium)reaching"copper brown" to "Mohawk"with the
extreme margin sometimes nearer "chipmunk"or "Arabianbr.," glabrous, sulcate to
disc, sometimes with a paler (but not pallid or white) zone around the central fuscous
or black dot which in dry caps appearsas a brown, fuscous, concolorous, or blackish
chestnut small knot or wart, located in an umbilicus, otherwise convex, but young cam-
panulate to convex with a prominent papilla, 1-8 mm broad. Lamellaewhite, whitish,
cream white or cream yellow, with the edge more or less tinged by the color of the
pileus (on lower lamellae and away from marginof pileus sometimes scarcelyvisible!),
broad, but not ventricose, distant at maturity, equal or more rarely with 1(-2) lamellu-
lae, 8-13, distinctly collariate. Stipe light umber to black with pallid to white apex in
youth, glabrousand smooth, 12-40 X 0.1-0.2 mm, insititious on the substratumbut
accompaniedby black branchedrhizomorphsor rhizomorphsof the color of the stipe
which are glabrous and often provided with ring-nodes,insititious on the substratum,
rarely stipe insititious on the rhizomorphbut not severalstipes branchingoff a single
rhizomorph. Context very thin, white in pileus, inodorous.
Spores (7-)7.3-9.5(-11) X (4-)4.7-5.3(-6.2)p, smooth hyaline, thin-walled,often
with a central oil droplet, ellipsoid, inamyloid, tending to germinateon lamellae. Hy-
menium: basidia 21-24 X 5-6.2 u, clavate, 4-spored;basidioles fusoid; cystidia none;
cheilocystidia like epicuticularbroom cells. Hyphae: in hymenophoraltrama inamy-
loid or almost so, in tramaof pileus varying from inamyloid to very weakly pseudo-
amyloid, in center of pileus and apex of stipe distinctly pseudoamyloid, in pileus at
maturity varying from hyaline to pale yellowish, of variablediameter,clamp connect-
ions present. Cortical layers: epicutis of pileus hymeniform, consisting of broom
cells of the Siccus-type, main body rarelygolden orange to golden melleous and then
often thick-walled,more numerousbeing the thin-walledcells which are hyaline or
subhyaline, or pale golden, hand-shaped. Clavariella-shaped, or clavate to vesiculose
or cylindric-broom-shaped,9-26 X 4.5-11 p, setulae erect or suberect, apical, golden
orange to golden melleous or golden yellow 2-8.2 X 1-2 (at base).
On dead wood, (roots, logs, fallen branches,woody sticks of all kinds, spines,
bark, etc.) and even on dead fern stems and various leavesattached to the main sub-
stratum. Identified host: Eugenia. Generallygregarious. Fruiting early in the rainy
season: December until mid-February. Basal subtropicalforest, here common.

107a. Marasmiuspraecox forma praecox

Pileus rusty orange when quite fresh, or even reddish orange or rufous.
MATERIAL STUDIED. ARGENTINA. Tucuman: Dpto. Tafi. Tafi Viejo, Quebrada de la
Toma, 16 Dec 1951, Helberger, comm. Singer T 1645 (LIL); NE of Yerba Buena, 20 Jan 1955.
Singer T 2033 (LIL), type; 12 Feb 1955, T 2161 (LIL). topotype; 20 Feb 1955, T 2200 (LIL),
topotype; Parque Aconquija, 17 Feb 1955, Singer T 2182 (LIL), paratype; Cuidad Universitaria,
15 Dec 1956, Singer T 2852 (LIL); Anta Muerta, 23 Feb 1949, Singer T 143 (LIL).

107b.Marasmiuspraecox forma aurantiicolorSinger, Sydowia 18: 340. 1965.

Differing from the type form in a more cinnamon orange pileus (11 A/C 10/11 to 12
Marasmius 155

12 B 11), dehydratedpartially to between "feuille morte" and pl. 13 A 12, i e more

orange when quite fresh and young than when partially dehydrated. Center of pileus
with or without a black dot (papilla).
MATERIAL STUDIED. ARGENTINA. Tucuman: Dpto. Taff, Parque Aconquija 17 Feb
1955, Singer T 2180 (LIL), type; T 2178 (LIL); Ciudad Universitaria, 1 Jan 1960, Singer T 3473

107c Marasmiuspraecox forma rubroumbonatusSinger, Sydowia 18: 340. 1965.

Differing from the type form in a more orange cinnamon pileus which becomes
reddishorange when half dry, with a red (drying brown) small umbo, dried (in herb-
arium) "Mohawk"(brown red) with "Arabianbrown" marginamd umbo "Mohawk"
MATERIAL STUDIED. ARGENTINA. Tucuman: Dpto. Taff, Parque Aconquija, 12 Mar
1949, Singer T 208 (LIL). type.
The spores are generallynot largerthan 9.7 X 5.3 ,. but in collection T 2852,
otherwise typical (f praecox), I observedspores slightly larger:9-11 X 5-6.2 p. I do not
think it possible to separatethis form from the others in view of the fact that some
collections have intermediatespores e g T 2178: (8-)8.7-10.7 X 4.5-5.3 ,.
The few cases of branchingstipe observedmay lead to the misconception that
this is Al. trichorhizus. This latter species should therefore be compared. Marasmius
praecox differs from 1M.robertsoniiin being more orange in dried, especially half-de-
hydrated condition than in the fresh state, and the lamellae edges being concolorous
with the pileus rather than the sides (faces) of the lamellae. There is also a difference in
Vlarasmiuspraecox is so close to AI. nfbrotula of southern Florida that it was at
first confused with that species by this author. However, the spores are consistently and
considerablynarrowerin the latter than in the former.
ILLUSTRATION. Singer (1965), pi 10, fig 49.

108. MarasmiusruforotulaSinger, Sydowia 2: 34. 1948.

TYPE. Singer F 1456, from Florida(USA).
Pileus rufous-red("rufous," "vinaceousrufous," "ocher red," more rarely between
"MarsYellow" and "Amber Brown" [Ridgway], "rust, sorollarbr" or "chrysanthemum"
with 4 G 12 margin,M&P),with deep brown to dark bister center, with sometimes cren-
ate margin,sulcate-plicate,convex, somewhat umbilicate, the zone around the papilla
sometimes minutely crackingrivulose, otherwise all glabrous,4-9 mm broad. Lamellae
whitish to cream, the edge concolorous with the sides or narrowlydiscolorous(con-
colorous with the marginof the pileus), distant (e g 8-14 through lamellae and 0-2 often
venose lamellulae), broad, sometimes somewhat intervenose,collariate but collarium some-
times low and narrow. Stipe cinnamon-fulvousto (rarely) succineous with or without an
olive tinge, or almost orange to between "Honey Color" and "Clay Color" (Ridgway),
often becoming "Bister"(Ridgway) or nearly black, shining, smooth, glabrous,setiform,
insititious to the substratum,equal, 12-35 X 0.1-0.3 mm. Context white, very thin, in-
Spores 7.2-10.2 X 3.5-4.3E, most frequently 7.5-9.3 X 3.7-4 p, ellipsoid to oblong,
more often oblong-subfusoid,smooth, hyaline, inamyloid. Hymenium: basidia 16-27 X
4-7.3,, 4-spored;cystidia none; cheilocystidia like the epicuticularbroom cells. Hyphae
of the pileus-tramahyaline, with clamp connections, most filamentous but many broad
(reachingabout 12 in diameter), some inamyloid, some pseudoamyloid. Hyphae of
156 Flora Neotropica

the stipe parallel, pseudoamyloid. Cortical layer: epicutis of the pileus hymeniform,
broom cells of the Siccus-type, main body 8-29 X 4-12 p, clavate or cylindrical, with
ratherthin wall, golden melleous to (few) subhyaline, with apical coarse setulae 3-8
X 1-2 u, deeper colored than main body, acute or obtuse.
On both mono-cotyledonous and dicotyledonous hosts (mostly dead dicotyledon-
ous wood) but also on living roots etc, sometimes among mosses, gregariousin subtrop-
ical and tropical forests, from Florida to Ecuador. Known hosts: Coccolobis, Carya,
Quercus,Ficus, Sabal.
MATERIAL STUDIED. U.S.A. Florida: Dade County, Matheson Hammock, 3 Nov 1942.
Singer F 1456 (FH) type, and various other collections from Florida (F, FH). MEXICO. Veracruz:
Estacion Biologica de las Tuxtlas, 19 Jun 1969, Singer M 8020 (F); M 8043 (F). ECUADOR. Napo:
Lago Agrio, 16 May 1973, Singer B 7506 (F).

109. MarasmiusbrunneocinctusSinger, sp nov Fig 57.

TYPE. SingerM 8982, from Mexico.
Pileo atrorufo, rufescente-aurantiaco,maculo atrobrunneozona brunneolo-brunnea
cincto, 7-8 mm lato; lamellis cremeis acie haud discolori, collario cremeo, distantibus;
stipite nigricanteapice albo; rhizomorphispraesentibussed stipiti haud adhaerentibus
Sporis 6.5-8.5 X 3.5-4 u; hyphis pilei pseudoamyloideis;elementis epicuticularibuscras-
situnicatis, typi Marasmiisicci. Ad folia dicotyledonea emortua in Mexico. Typus in
F conservatusest.
Pileus deep rufous, rufescent-orange,dried between "kis kilim" and 8 J 12 (M&P),
with a paler zone (brown spot visible on brownish pallid ground under a lens) around
the deep brown, round spot in the center, but the paler zone poorly differentiated,
glabrous,sulcate, with a smooth, flat-bottomedumbilicus and a convex mediane zone,
eventually often recurvedat extreme margin 7-8 mm broad. Lamellaecream with not
discolorous edge, distant (10 through-lamellae),all equal and simple, broad but some-
times strongly narrowedat marginof pileus and not reachingthe latter, collariate with
a cream colored collarium, leaving in dried material between the rim of the collarium
and the apex of the stipe a brown, sterile zone. Stipe blackish with white apex, seti-
form, shining, glabrous,insititious on the substratumbut accompaniedby thin, black
rhizomorphs,equal, 23 X 0.2 mm; rhizomorphsthin, not very abundant. Context
white, thin, inodorous.
Spores 6.5-8.5 X 3.5-4 p, ellipsoid, smooth, hyaline, inamyloid. Hymeniurn:basid-
ia 21-26 X 6.2-7.2 p,,4-spored;cystidia none; cheilocystidia none seen (possibly eroded).
Hyphae in the hymenophoraland pileus-tramaas well as in the apex of the stipe hyaline,
firm- but not thick-walled,with clamp connections (but many secondary septa observed)
distinctly pseudoamyloid. Corticallayer: epicutis of the pileus hymeniform, broom cells
of the Siccus-type, more or less strongly pigmented cells evenly distributedand surface
in scalp therefore appearingmottled, main body 8.25 X 6-13.5 u, thickwalled and yellow
with deep yellow to golden yellow setulae, or thin-walledand hyaline to yellowish hy-
aline with subhyaline to yellow setulae, the latter apica!, taperingupwards,3-5.5 X 0.8-
1.3-(2) ,.
On dicotyledonous dead leaves and leaf petioles in tropical rain forest. Mexico.
MATERIAL STUDIED. MEXICO. Chiapas: between Finca Sospiro and El Pozo, 4 Aug 1969,
Singer M 8982 (F), type.

110. MarasmiuspallidocinctusSinger sp nov

Marasmius 157

11Oa.Marasmiuspallidocinctusvar pallidocinctus
TYPE.Dumont et al VE 701, from Venezuela.
Pileo ferrugineo-aurantiaco,zona pallida circum maculum centralem fumoso-ful-
igineum ornato, 3-6 mm lato; lamellis albis, subdistantibus,collariatis, stipite atrocast-
aneo; rhizomorphisnullis. Sporis 9-10.5 X 3.5-4.3 1; hyphis pilei debiliterlentoque
pseudoamyloideis,fibulatis;elementis epicuticularibustypi Marasmiisicci. Ad folia
dicotyledonea delapsa in Venezuela. Typus in NY conservatusest.
Pileus (dried) rusty orange with a pallid zone around a central fumosous-fuliginous
spot, glabrous,sulcate-plicate,convex, deeply depressedin the center with a low papilla
becoming visible in dried material, 3-6 mm broad. Lamellaewhite with white edge and
collarium, subdistant(about 12-23 through-lamellae),broad, collariate. Stipe deep chest-
nut, glabrous,equal, insititious on the substratum,not rising from nor accompaniedby
black rhizomorphs,long: 75-80 X 0.3 mm; "telepodes" often formed. Context whitish,
thin, inodorous.
Spores 9-10.5 X 3.5-4.3 p, ellipsoid-oblong,smooth, hyaline, inamyloid. Hymenium:
basidia4-spored;basidioles fusoid; cystidia none; cheilocystidia like the epicuticularbroom
cells but all hyaline. Hyphae of the pileus-tramayellowish-hyalineto hyaline, weakly and
slowly but undoubtedly pseudoamyloid,with clamp connections, filamentous, 4-5 u broad,
thin-walled,not gelatinized;hyphae of the stipe parallelpigmented, pseudoamyloid. Cor-
tical layer: epicutis of pileus hymeniform, somewhat mottled because of alternatinglypaler
and darkerbroom cells, these of the Siccus-type, main body 11-15 X 4.5-6 u, thin-walled,
yellowish hyaline, with setulae 2-5 X 0.5-1.5 ,, acute or obtuse, simple or forked, yellow-
ish hyaline to golden ochraceous.
On dicotyledonous leaves fallen to the ground. Venezuela.
MATERIAL STUDIED.VENEZUELA. Dto Federal, Mt. Naiguata, above La Planta, 25 Jun
1971. Dumont et al VE 701 (NY).

1lOb.Marasmiuspallidocinctusvar latisporusSinger,var nov Fig 58.

TYPE. Singer B 535, from Bolivia.
A var. pallidocincto differt sporis 9-11.8 X 4.8-6.2 j. Typus in LIL conservatur.
Pileus "ferruginous"M&P,pallid around a black, central spot which in dried con-
dition becomes brown with a paler but not pallid zone around it, marginalzone in dried
condition "hazel" to 13 J 10, sulcate there, glabrous,center smooth, convex, umbilicate,
when dried showing a low and obtuse (brown) umbo in the umbilicus, 3 mm broad.
Lamellaewhite or whitish with white or whitish edges and collarium, distant (12-13
through-lamellae,no lamellulae),collariate. Stipe sordid ochraceousbrown (13 K 10 to
"tortoise shell" M&P),deeper colored at the base, with at first white apex, glabrous,
smooth, insititious on the substratum,not accompaniedby rhizomorphs,equal, 60 X
0.2 mm. Context thin, inodorous.
Spores 9-11.8 X4.8-6.2 p, ellipsoid, smooth, hyaline, inamyloid.Hymenium:bas-
idia 19-19.5 X 6-6.5 u; cystidia none; cheilocystidia ratherscattered,like epicuticular
broom cells but less pigmented. Hyphae of the pileus-tramahyaline, mostly filamentous,
inamyloid or almost so. Cortical layer: epicutis of pileus hymeniform, broom cells of
the Siccus-type, main body 12-20 X 6-7.5 1, hyaline to golden melleous, setulae 2.5-5.7
X 1.5 1, spinulose, sometimes forked, mostly golden, fewer hyaline.
On dead, fallen, dicotyledonous leaf, gregarious. Yungas.
MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Coroico, 29 Jan 1956, Singer B 535
(LIL), type.
158 Flora Neotropica

The different size of the spores seems to be the only important difference between
this and the type variety.

111. MarasmiusrufomarginatusSinger, Sydowia 12: 134. 1958.

TYPE. Singer F 1219, from Florida (U.S.A.).
Pileus deep orange red to orange brown or salmon orange("English Red," "Orange
rusous" or "MarsOrange"(Ridgway) becoming "Salmon Orange"or "MarsOrange"or
"chinook' (M&P)in age), except for the center, which is "Burnt Sienna" (Ridgway) or
chestnut in most specimens, more orange brown when dried, slightly paler or concolor-
ous around the brown center, not showing a white zone or, if such a zone is present, it
is due to crackingwhereby the white trama of the pileus is partly exposed (use lens!),
glabrous,sulcate to sulcate-plicatewith smooth center, at first campanulate-papillate,then
hemisphericalto convex, soon rather deeply and often narrowlyumbilicate and at least
when dry showing a small umbo or papilla in the umbilicus, 0.8-6.5 mm broad, mostly
2-5 mm broad. Lamellaewhite, eventually often becoming cream color, with the edge
at least in the marginalhalf discolorous(concolorous with the marginof the pileus)
distant (7-11, more often 10-11 than 7-9 through-lamellaeand 0-2 lamellulae), simple
or one to severalforked near stipe, broad (1-2 mm), collariate but the collariumat
first low, later often incomplete and sometimes partly adheringto the apex of the
stipe. Stipe chestnut to blackish-brownor black, with at first white apex, shining,
glabrous,setiform, insititious on the substratum,8-28 X 0.1-0.2 mm: black rhizomorphs
not seen; "telepodes" often formed. Context hyaline-white,thin, inodorous.
Spores 6.5-10(-10.2) X (2.5-)3-4-5 p, mostly 7.8-8.7 X 3.7-4 p, ellipsoid-claviculate
to oblong, rarely ellipsoid Q = (1.7-)2.1-2.5, with the hilar end often stronglyelongated
and acute in overmature spores, smooth, hyaline, inamyloid. Hymenium: basidia 30 X
8-10.2 ,, 4-spored;cystidia none; cheilocystidia like the epicuticularbroom cells or more
subhyaline to golden orange. Hyphae of the pileus-tramahyaline, weakly and slowly but
distinctly (purple) pseudoamyloid, in the hymenophoraltrama which is regular,often
almost to quite inamyloid, in the apex of the stipe parallel,pseudoamyloid,all hyphae
with clamp connections. Cortical layers: epicutis of the pileus hymeniform, broom cells
of the Siccus-type, main body 6-22.7 X 3-9.3 p, varying from hyaline to rusty-orange,
some thick-walled;setulae 2-7 X 0.8-2 p, rod-shapedto conical, subacute to obtuse, gold-
en yellow to deep rusty orange, some hyaline, erect at the apex and often oblique just
below the apex.
On leaves of dicotyledonous trees, fallen to the ground in subtropicaland tropical
to tropical-montaneforest. Florida to Mexico, south to Colombia. Known host: Nec-
MATERIALSTUDIED. U.S.A. Florida:Dade County, Miami,SimpsonPark,23 Oct 1942.
SingerF 1219 (FH), type; MathesonHammock,30 Oct 1942, SingerF 1219a (FH); 22 Oct 1942,
F 1196 and 24 Oct 1942, F 1196a (FH). MEXICO.Chiapas:Tuxtla de Gutierrez,BotanicalGarden.
3 Aug 1969, SingerM 8921 (F). COLOMBIA.Valle: Corr.Vergel, Finca La Araucana,14 Apr 1968,
SingerB 6126 (F). ECUADOR. Tungurahua,Rio Verde, 28 Apr 1973, SingerB 7172 (F).

112. Marasmiusfoliicola Singer, sp nov

Marasmius ruforotula var foliicola Singer, Sydowia 12: 136-137. 1958, nomen nudum.
Marasmiusfoliicola (Singer)Singer,Sydowia 18: 205. 1965 "ad interim."
TYPE.SingerT 2147, from Argentina.
Pileo aurantiaco,in umbilico nigro-maculato,4-7 mm lato; lamellis albis, acie aur-
antiacis, distantibus,collariatis;stipite atrocastaneovel nigro, ad apicem primum saepe
albo, 15-75 X 0.1-0.5 mm; rhizomorphisnullis. Sporis 7.3-10 X 4-5 p; elementis epicut-
Marasmius 159

icularibustypi Marasmiisicci. Ad folia monocotyledonea et dicotyledonea delapsain

Argentina. Typus in LIL conservatus est.
Pileus orange (shading from "bittersweeto." to "Tokyo" M&P),and margineven-
tually bleachingin many specimens to light salmon orange(9-F-8), deeply sulcate to
disc, with a black dot (and sometimes very small planetary dots) in an umbilicus, the
dot becoming blackish brown to pale rufous brown, or disappearingaltogether when
drying out, convex around the umbilicus, 4-7 mm. Lamellaewhite with orange edge
(this discolorationat times discontinuousand not very conspicuous or in some caps
limited to the region near the marginof the pileus or to the lamellae edges reachinga
higher level, i e broaderlamellae, than others, but always more or less distinct), flat
to arcuate, distant, 10-13, collariate, rarely incompletely collariate. Stipe in very young
and fresh materialdeep brownish orange with white upper portion, soon dark chestnut
or black with or without white apex, glabrousand smooth, insititious directly on the
substratum,15-75 X 0.1-0.5 mm, not accompaniedby rhizomorphs. Context very thin,
white, inodorous.
Spores 7.3-10 X 4-5 j,, ellipsoid, smooth, hyaline, inamyloid, often germinating
on the lamellae. Hymenium without cystidia; cheilocystidia like the epicuticular broom
cells but some hyaline ones (both setulae and main body) present. Hyphae of pileus-
tramaweakly but undoubtedly pseudoamyloid, with clamp connections. Coveringlay-
ers: epicutis of pileus consisting of Siccus-type broom cells 13-23 X 5.5-9(-11.7)A, some-
times spreadingto 16 u diameter,golden to orange with paler colored main body, very
few broom cells outside the disc area entirely melleous brown.
On dead or fallen leaves in subtropicaland tropical zone, more frequently on
monocotyledonous e g (Aech,mea,Bambusa) than on dicotyledonous leaves.
MATERIAL STUDIED. ARGENTINA. Salta: Oran, 18 Mar 1955, Singer T 2311 (LIL);
Tucuman: Capital, Garden of the Instituto Miguel Lillo, 10 Feb 1955 Singer T 2147, (LIL) TYPUS.
This species must occur throughout South America, since illustrationsof a Mara-
smtiusfound by Dennis on dead Bauhiniaand on dead Mlusaleaves respectively,in
Caracasand in Cortadade Guayabo (Miranda),Venezuela, with spores 7-9 X 3.5-5 ,,
show exactly the same characters,Dennis 1020, 1157 (both K, v.i.).
Mjarasmius plicatus Wakker,ZiektenSuikerrietJava 1 p 195, Leiden 1898, is
possibly this. I have not seen the type.
MIarasmlius fbliicola might possibly be a synonym of Marasmiusgordipes Saccardo&
Paoletti, Micetes Malaccenses,Atti R. Istituto Veneto Sciencie, Lettere ed Arti VI. 6:
387428, pl. 5, fig. 2, 19. I have not located the type of the latter. Nevertheless,the
colors of both these Asiatic fungi are describeddifferently and the number of through-
lamellae is smaller. I believe therefore that they are different and accept Mlarasmius
foliicola as an autonomous species.
Marasmiusexustus Berkeley & Curtis from the Bonin Islandshas been mentioned
as a possible synonym of Marasmiusgraminurmor M. foliicola. Indeed, WesternHemi-
sphere materialdeterminedas suchprobablybelongs to Marasiniusfoliicola but the type
(FH) of A. exustus has been studied by me and belongs in section Hygrometrici.
ILLUSTRATION. Singer (1965) pi 10, fig 51, 52.

Marasmiussect Sicci Singer, Mycologia 50: 106. 1958.

TYPESPECIES. Maraslius siccus (Schweinitz) Fries.
Pileus rathersmall to large, white or pigmented, with hymeniform epicutis con-
sisting partly or more often entirely of broom cells of the Siccus-type spores small to
large;lamellae not collariate;hyphae with clamp connections, mostly (except in Mara-
160 FloraNeotropica

smius inaequalis) strongly pseudoamyloidin pileus and stipe; stipe centrally attached,
growingfrom leaves or wood etc, always with basal mycelium; black rhizomorphsabsent.

Key to the Subsections of Section Sicci

1. Tramaof the pileusinamyloid. subsection
1. Tramaof the pileuspseudoamyloid. subsection

Marasmiussect Sicci subsect InaequalesSinger,Sydowia 12: 97. 1959.

TYPESPECIES.MarasmiusinaequalisBerkeley & Curtis.
Hyphae of the carpophoreinamyloid.

113. MarasmiusinaequalisBerkeley & Curtis, Jour. Linn. Soc. Bot. 10: 297. 1869.
TYPE. WVright, from Cuba.
Pileus white, center brown ("cookie" M&P),dried entirely like illarasmiutshelvolus
in color, glabrous,sulcate, campanulateto convex, 2-5 mm broad. Lamellaewhite with
white edge, distant or subdistant(10-12 through-lamellae),rathernarrow,subconcolorous
with the pileus when dried, narrowlyadnexed to free, definitely not collariate. Stipe
chestnut brown with pallid or white apex, dried chestnut-amber,smooth, glabrous,shin-
ing, 4-32 X 0.1-0.2 mm; basal mycelium tomentose or strigose, white, accompaniedby
whitish mycelial patches and rhizomorphsin many populations. Context very thin, in-
Spores 11-15 X(2-)3-3.3 ,, guttiform-subclavateto cylindric-subfusoid,often miore
or less curved or at least the hilar appendagestrongly curved inwards,smooth, hyaline,
inamyloid. Hymenium: basidia 20-21 X 5.5-6.5 p, 4-spored;cystidia none; cheilocystidia
like the epicuticularbroom cells but here all thin-walled,hyaline to melleous. Hyphae
of the trama of the pileus with clamp connections, inamyloid. Cortical layers: epicutis
of the pileus hymeniform, broom cells of the Siccus-type, of two kinds, (1) thin-walled,
hyaline to melleous, sometimes all hyaline, main body 8.5-17 X 5.5-10u, setulae acute
or subacute, 3.3-4.5 X + 1 ,, (2) as above but thick-walled,wall up to 1.7 p thick.
On rotting dicotyledonous wood (sticks, chips, roots, etc.), gregarious.
MATERIALSTUDIED.CUBA. Santiagode Cuba, 1857. Wright114.4 (FH), authentic.
BOLIVIA. La Paz, Nor-Yungas,Coroico, 30 Jan 1956, SingerB 736 (LIL).
Singer(1958a) fig 18.

Marasmiussect Sicci subsect Siccini Singer, Sydowia 18: 343. 1965.

TYPESPECIES.Marasmiussiccus (Schweinitz) Fries.
Hyphae of the carpophorepseudoamyloid.

Key to the Series of Subsection Siccini

1. Setae in pileus-epicutisand hymenophore none; stipe glabrous except for the basal
mycelium(tomentoseor strigose),morerarelyveryfinelypruinate.
2. Cystidiaon the sidesof the lamellaeabsentor not clearlydifferentiated. seriesLeonini, p 160.
2. Cystidiaon the sidesof the lamellaepresent. seriesHaematocephali,p 201.
1. Setae in pileus-epicutisand/or hymenophorepresent,or stipe distinctly pruinate,pube-
scent or hirsutefrom dermatocystidioidhairsor setae. seriesActinopodes,p 236.

Marasmiussect Sicci subsect Siccini ser Leonini Singer, ser nov

Cystidiis ad latera lamellarumvix a cheilocystidiis distinctis nec typo cystidiorum
Marasmiisicci - Typus: Marasmiusleoninus Berkeley.
Cystidia and setoid hairs absent.
Marasmius 161

Key to the Species of SeriesLeonili

1. Pileus white over most of its surface when young (in part pale cream or cork color at
2. Spores larger than 14,. 114. M. subrotula.
2. Spores not reaching 14p in length.
3. Lamellae medium distant to distant; spores up to 11 p long or longer.
4. Pileus up to 9 mm broad.
5. Spores (7-)12.5 p long; cystidia present but scarce (see series Haematocephali).
5. Spores 6-9.2(-1 l)p long; cystidia none. 133. M. proletarius.
4. Pleus more than 10 mm broad.
6. Pileus neither in age or on drying becoming yellow; spores 11-13.5 Aulong;
setulae of epicuticular broom cells hyaline. 132. M. haediniformis.
6. Pileus tending to cream or yellow; spores 8-12.7u long; setulae of the
epicuticular broom cells at least in accumulation and in parts of the pileus
distinctly yellowish to chrome yellow. 141. M. bellus.
3. Lamellae crowded, close or subclose; spores up to 11 u long.
7. Pileus 6-11 mmnbroad, pallid. 134. M. microhaedinus.
7. Pileus pigmented, eventually often bleached, more than 12 mm in diameter
when quite mature (see "8" below).
1. Pileus not predominantly white.
8. Pileus red or purple at least when young and fresh.
9. Pileus bright red (orange red, scarlet red); spores 8.2-9.7, long, lamellae orange
or reddish. 140. M. ruber.
9. Pileus more purple red or purple, sometimes radially striped or dotted and stripes
white, buff or ochraceous; spores larger.
10. Pileus 42-72 mm broad (largest of the group); pileus with buff colored dots;
spores 12-21 X 3.5-4.8 M,oblong or fusoid. 115. M. amazonicus.
10. Pileus smaller.
11. Pileus not radially striped or not conspicuously so; basal mycelium brown;
spores up to 14,i long. 116. M. tucumanus.
11. Spores 12-20.3u1 long; basal mycelium (fresh) white to pale brown or ful-
vous; pileus conspicuously radially striped. 117. M. tageticolor.
8. Pileus never red or purple when fresh or dried, excepting in some rare cases when the
pileus is partially a deep orange red or reddish brown to bay color.
12. Spores 13.8-22 X 3.3-5.5 A.
13. Carpophores rather small, pileus generally campanulate for a long time before
expanding and not over 17(-20) mm broad, rarely broader, habit of
Marasmiuzshaematocephalus or M. anomalus.
14. Pileus conspicuously striped white or yellowish and red-brown or brown.
15. Edges of the lamellae concolorous with the sides; on dicotyledonous
leaves. 118. M. poecilus.
15. Edges of the lamellae concolorous with the pileus, not white; on bark
and wood. 119. M. phaeus.
14. Pileus not striped radially or, if striped, stripes not very conspicuous or
not white or yellow.
16. On fallen dicotyledonous leaves. 120. M. banbusiniformis.
16. On wood.
17. Edge of lamellae showing a broad zone of strongly pigmented
cheilocystidia; pileus about 17 mm broad. 121. M. mazatecus.
17. Edge of lamellae white or whitish, few or not pigmented cheilocys-
tidia present, more rarely there is a thin line of discoloration at the
very edge which becomes distinct only when dried material is ob-
served under a lens; pileus generally not broader than 10 mm.
18. Fresh pileus cork color or cinnamon-isabelline (see "B" above).
18. Fresh pileus gold brown, raw sienna, ochraceous tawny or
orange to orange-rufescent.
19. Pileus gold brown or raw sienna, then ochraceous tawny to
ferrugineous, 4-10 mm broad; pileus conic to campanulate,
often papillate. In tropical and tropical montane forests up
to the subparamo, not in subxerophytic vegetation.
122. M. longisporus.
162 Flora Neotropica

19. Pileus orange to orange rufescent, 5-6 mm broad, hemi-

spheric and obtuse. In xerophytic and subxerophytic vege-
tation. 123. M. onoticus.
13. Carpophores 7-37 mm broad, pileus soon becoming convex-repand.
20. Spores 4-5.5u broad and up to 18, long; pileus red-brown in the center.
cinnamon on the margin; lamellae close. On wood. 124. M. rubricosuls.
20. Spores narrower; pileus differently colored (more yellow and orange)
when fresh; on dead fallen leaves. 125. M. rlhabarbarimts.
12. Spores somewhat to considerably smaller, reaching in some species 16.2(-17)j. in
length, but not longer, many mature spores less than 13.8, in length (if only few,
possibly young spores were observed, compare also "13" above).
21. Pileus with some olive shade or entirely olive colored, sometimes nearly
22. Spores 6-9 X (3-)3.5-4.5(-5)u, lamellae subclose to medium close, pileus
5-19 mm.
23. Dried lamellae gray with fuliginous edge. 139. M. ater.
23. Dried lamellae pallid to grayish, edges concolorous witli sides or very
much paler than fuliginous. 138. M. digilioi.
22. Spores larger or lamellae more distant or pileus broader.
24. Spores larger than 8.5 p. From Florida to northern South America.
25. Edge of lamellae in places vaguely pale orange discolorous or else
concolorous with the sides of the lamellae. 135. M. trinitatis.
25. Edge of lamellae discolorous, olive. 136. M. olirascenticeps.
24. Spores smaller than 8.5 . Amazonas region. 1 37. M. epelaeus.
21. Pileus without any olive shade in fresh condition and not nearly black in
dried condition. Key II.

Key II
1. Spores 11-16.2(-17) X 2.7-6.3u.
2. Lamellae subdistant to distant.
3. Dried pileus "cocoa," "leather or" M&P often radially striped in different shades.
of brown; fresh pileus not orange or orange-ferrugineous.
4. Spores 11-15.2 X 2.8-4u; pileus 9-31 mm broad; cn woody twigs and vines and
on dicotyledonous leaves; cystidia usually present but often few or in-
conspicuous. 174. Mi.helholus.
4. Spores 12-16 X 2.5-3, (i e relatively narrower); pileus 27-42 mm broad; on
wood in Pacific rain forest. 130. M. helioloides.
3. Dried pileus differently colored, e g "Ginger," "persimmon," tending to "Alamo,"
"terracotta," "ferruginous," "caldera" in the center, young pilei often entirely
"raw sienna" i e more brightly ochraceous brown, brown-red or orange-rust when
dried, fresh pileus mostly orange or ferruginous or bay to ferruginous.
5. Lamellae with discolorous edges; spores 12-13.8 X 3.3-3.7,u; pileus 4-9 mmi
broad, orange, rusty orange; mostly growing on grass leaves, more rarely passing
over unto Populus or Taxodtium leaves. Southern subtropics. 131. Al. graminizcola.
5. Lamellae discolorous only in a narrow intermittent line (under a lens. \when
dried) or concolorous with the sides of the lamellae; pileus as small as above or
larger; spores as above or smaller or larger; not with the above habitat and
more widely distributed; not combining the characters indicated above.
6. Spores only up to 13 long; lamellae only medium distant to subdistant;
pileus small to medium (up to 20 mm broad); species of the northern part
of South America and reaching Mexico and the whole Caribbean area (see
"19" below).
6. Spores, at least many of them, reaching more than 12.5 p in length, often
up to 16(-17)p long. Widely distributed all over the tropics and subtropics
of the Americas. 126. M. berteroi.
2. Lamellae close or subclose.
7. Edge of lamellae distinctly and constantly discolorous; spores 11-15 X 3-4 g pileus
6-9 mm broad; lamellae subclose. On leaves of Dicotyledones. 129. M. neglectus.
7. Edge of lamellae concolorous with the sides or, if slightly discolorous, with dif-
ferent spores or not combining the above characters.
8. Spores 12.5-16 X (2.5-)3-4u; pileus on margin subsmooth or sulcate, in the
center smooth or convoluted; stipe covered with subhyaline, clavate, entire
dermatocystidia, or without any covering.
Marasmius 163

9. Stipe covered with clavate, entire dermatocystidia; pileus on margin sub-

smooth, in the center convoluted. 127. M. convoluticeps.
9. Stipe without dermatocystidia and broom cells; pileus deeply sulcate, not
convolute (see "4" above).
8. Spores (9.7-)11-15 X 4-6.3 (relatively broader); pileus with sulcate margin
and a rugose zone further inward; stipe covered by broom cells. 128. M. matrisdei.
1. Spores somewhat to considerably smaller than 11-16.2(-17) X 2.7-6.3 U, generally not
over 12(-12.5) long.
10. Hymenophore appearing poroid because of anastomoses between the crowded to
close lamellae. 150. M. cladophyllus.
10. Hymenophore strictly lamellate but often intervenose or forked, yet never offering a
pore-like aspect.
11. Pileus more than 7 mm broad when mature.
12. Pileus cream color to yellow, lamellae subdistant to distant; spores 8-12.7 X
34.8 A 141. M. bellus.
12. Pileus neither cream colored nor yellow except in combination with other
colors: brownish, oclraceous-brown, red-brown, orange, orange-salmon,
orange-ferruginous, bright rusty brown.
13. Pileus when dried dull brown, ochraceous buff when fresh; edges of lamel-
lae indistinctly concolorous with the pileus; stipe without broom cells;
epicuticular broom cells not accompanied by smooth vesiculose bodies;
spores small, 7-8.2 X 2.7-3,; lamellae subclose to subdistant (not close
nor distant). Rain forest species. 142. A. beniensis.
13. Pileus, if brown when dried or ochraceous buff when fresh, showing a
broadly discolorous gill edge (concolorous with the pileus) or a consider-
able number of smooth elements in the epicutis especially in older pilei;
spores as above or somewhat larger; lamellae as above or crowded to close.
14. Edge of the lamellae distinctly broadly discolorous; smooth epicuti-
cular cells none or rare and very scattered. 143. M. bezerrae.
14. Either the lamellae are not or only slightly discolorous or the epicutis
contains a larger number of smooth cells aside from broom cells in
adult specimens.
15. Epicutis of adult specimens constantly consisting of two types of
elements: (a) broom cells, (b) smooth cells like the main bodies of
the broom cells but without setulae.
16. Numerous normal broom cells on the stipe, these varying from
scattered to rather dense; pileus light brown to cinnamon-
brownish, furrowed-sulcate almost to the center, not rugose-
venose or radially rugose in the center; spores 6.2-9 X 5-3.711.
147. M. cuatrecasasii.
16. Broom cells varying from rather numerous to absent on the
stipe; pileus colored as above or more orange, often bleaching in
age, not or slightly sulcate or striate on the margin or becoming
densely long-sulculate, center (or sometimes almost the whole
pileus) at first often smooth but later becoming rugose-scrobic-
ulate or radially venose-rugose. 148. M. corrugatus.
15. Epicutis of pileus consisting exclusively or almost exclusively of
typical broom cells; pileus usually long-sulcate and with weakly or
mostly not rugose center.
17. Pileus orange, rust colored or red-brown, Q of spores = + 3.
18. On wood or leaves; lamellae subclose to distant.

19. Pileus 15-42 mm broad; stipe 1.2-4 mm broad; lamellae

free to adnate; pileus obtuse or with a low broad umbo
or subumbonate, often broadly depressed in age, more
rarely with a wall-like circular depression around an um-
bilicus; broom cells scattered over the surface of the
stipe but sometimes without these.
145. M. leoninus.
19. Pileus 9-20 mm broad; stipe equal and up to 1.2 mm
broad, or slightly attenuated upwards with the base up
to 1.5 mm broad; pileus mostly distinctly small-
umbonate or papillate, more rarely with an abrupt cen-
tral umbilicus; lamellae free to subfree; surface of stipe
164 Flora Neotropica

without broom cells, or with setulae devoid of a main

body (if wit'i broom cells see "16" above).
146. M. floriceps.
18. On wood and needles of Araucaria; lamellae subclose to
crowded. 149. M. araucariae.
17. Pileus as above or brown; spoies less elongated (Q = 1.7-2.7),
7-9.5 X 3.5-4.5 uor 7.5-11 X 3-6,.
20. Pileus brown "cocoa" with "seminole" margin (M&P) or
"chutney;" stipe finely pruinate at apex, pileus up to 60
mm broad. 144. M. napoensis.
20. Pileus dull orange, often with pallescent margin, 10-30 mm
broad; stipe glabrous. 145a. M. leoninus var aberrans.
11. Pileus 2-8 mm broad. 151. M. pusio.

114. MarasmiussubrotulaMurrill,North AmericanFlora 9: 259. 1915.

Marasmtiusimelinocephalus Singer, Sydowia 18: 338. 1965.
TYPE. Earle 501, from Jamaica.
Pileus entirely "cork" color or center "wild honey" (cinnamon-isabelline)and then
often with whitish or paler (than "cork") margin,or on the contrary, "wild honey" all
over, dried "terrapin"in center and "bran"on margin,or between "Arab"and "tortoise'
M&P,when young entirely smooth but soon strongly sulcate almost all over, glabrous,
campanulate,becoming almost applanateand then umbonate, 4-15 mm broad. Lamellae
white, not collariate, all equal, distant, subventricose,free to adnexed. Stipe at first
hyaline with succineous or chestnut colored base, later chestnut to chestnut black or
umber with white apex, eventually unicolorously chestnut black or deep umber,glabrous,
smooth, setose, 20-55 X 0.2-0.5 mm; basal mycelium sparseto well visible, tomentose,
white. Context very thin, inodorous.
Spores (14-)15-23 X (2.7-)3.5-5.5 p, fusoid-clavate,thin-walled,hyaline, smooth.
Hymenium: basidiaup to 31 X 8.2-13.7 ,, 4-spored;cystidia often present but few and
poorly or not differentiated,basidiomorphous(like basidioles, fusoid, or basidiaclavate),
neither projecting nor deeper rooting, not appendiculatenor distinctly opaque, but some-
times with a very thin, weak incrustationon apex; cheilocystidia like the majority of
the epicuticularbroom cells, all hyaline or subhyaline. Hyphae hyaline, pseudoamyloid,
with clamp connections. Cortical layers: epicutis of pileus hymeniform, consisting of
broom cells, 6-21 X 5-13 A, of two types, the majority (type 1) consisting of hyaline,
thin-walledmain body and hyaline to pale brownishmelleous setulae 1.5-4(-10) X 0.8 p;
regularlyscattered among these and giving a scalp preparationa mottled appearance,
are broom cells of type II with a main body whose wall graduallythickens towards the
apex and is concolorous with the setulae, and setulae deep melleous to spadiceous-ful-
iginous, some broom cells type II with few or no setulae.
On earth or, more often, dung-coveredfallen dead branchesof dicotyledonous
trees and shrubs, particularlyAchatocarpusnigricansin marginaland subxerophytic for-
est in the subtropicalzone, densely gregariousto cespitose, sometimes directly and ex-
clusively on horse dung, sometimes on dead dicotyledonous twigs.
MATERIAL STUDIED. JAMAICA. Hope Gardens, 16 Nov 1902, Earle 501 (NY) type.
BOLIVIA. La Paz, Nor-Yungas, Charobamba, 30 Jan 1956, Singer B 755 (LIL). ARGENTINA.
Salta, Rfo Piedras between Metan and Gilemes, 25 Mar 1951, Singer T 1428 (LIL); Tucuman:
south of Tapia, 17 Feb 1951, Singer T 1221 (LIL); 5 km S of Burruyacui, 19 Feb 1957, Singer T
2997 (LIL), type of M. melinocephalus.
ILLUSTRATION: Singer(1965) pl 15, fig 82 T; (1958a), fig 32.
Marasmius 165

115. MarasmiusamazonicusHennings,Hedwigia43: 183. 1904.

TYPE. Lost (see below).
Pileus deep purple to deep lilac violet, the disc blackishpurple ( e g 56-J/L - 5
M&P)near center with numerousorangebrown ("talavera"),near marginpale buffy
orange("capucin buff') or everywherebuff dots which are oval to round and 2-4 mm
in diameter,glabrous,deeply sulcate almost to the disc, campanulate-convexto convex
applanate,with umbilicate disc, 42-72 mm broad. Lamellaewhite or buff-cream,broad
(6-16 mm broad), ventricose, intermixedwith 0-1 lamellulae,not intervenose,with en-
tire, not discolorousedge, free to removed from the apex of the stipe and then leavingun-
covered a white, sulcate disc around the extreme apex of the stipe, distant (19-22 lam-
ellae). Stipe deep chestnut to black, paler brown at apex when young, slightly longi-
tudinally striate when dried, otherwise smooth, glabrous,shining,hollow, equal or
graduallyand very slightly taperingupwards,terete and sometimes somewhat compress-
ed, sometimes arisingfrom a small socle which is covered by the basal mycelium, 80-
145 X 2.5-3.5 mm; basal mycelium tomentose, "cocoa" color. Context thin, white,
toughish, inodorous.
Spores 12-21 X 3.5-4.8,u, oblong to fusoid, in profile often somewhat curved,
attenuate towards the base, hyaline, thin-walled,smooth, inamyloid. Hymenium:
basidiahyaline, 4-spored (?); cystidia none; cheilocystidia like epicuticularelements,
but all hyaline. Hyphae hyaline, distinctly pseudoamyloid. Corticallayers: epicutis
of pileus consisting of broom cells which are hymeniformly arranged;these hyaline to
subhyalineseen individually(NH40H) but in thicker layer deep porphyry brown, in-
dividualsetulae mostly hyaline or subhyalinebut here and there some purple black
setulae present, total measurements15-18 X 9-10,, main body often indistinct and
reduced, setulae spinulose and subacute, taperingfrom a 2-2.7,ubroad base, 5-9A
long, entirely inamyloid.
On trunks and on very rotten wood among leafmold, in small groups, not ces-
MATERIALSTUDIED. BOLIVIA. Beni: Vaca Diez, Guayaramerin,8 Mar1956, Singer
B 1706 (LIL); 14 Mar1956, B 1946 (LIL).
The species was originallydescribedfrom Jurua miry, Amazonas,Brazil, collec-
ted by Ule. The type was conservedin Berlin where it was destroyed during the last
war. The descriptionhowever, leaves not the slightest doubt as to the identity of the
material. This is the most spectacularspecies of South Americaas far as external
appearanceis concerned. A similaror analogousspecies from Cameroon,M. superbus
Henn., differs in umbonate pileus with rugolose-venoseand not dotted surfaceand,
accordingto Hennings,collariatelamellae, and broaderstipe. It belongs in section
ILLUSTRATION. Singer(1965), pl 14, fig 72.

116. MarasmiustucumanusSinger in Singer& Digilio, Lilloa 25: 206. 1952.

TYPE. Singer T 1460, from Argentina.
Pileus intensely pink (54-K-3 M&P)with dark purple ("wild raspberry,"6-E-6)
center, deeply sulcate to the center, glabrous,indistinctly papillate at least on drying,
6-11 mm broad. Lamellaepink, soon becoming white in the edge half, 1.5 mm broad,
not collariate, distant, 10-17 through-lamellaeand 0-6 lamellulae,free, later adnexed
to adnate. Stipe at first entirely purple-redthen black towards the base, glabrous,
smooth and shining, with a minute disc or socle at base which is bearingthe basal
166 FloraNeotropica

mycelium 33 X 1.5 mm, base reaching0.5 mm, basal mycelium tomentose, brown,
with alternatinghyaline fibers even in fresh material. Context very thin, in cortex of
stipe almost purple, then blackening,fragile, inodorous.
Spores 10.3-15 X 3.3-4.8A, mostly about 11.5-13.7 X 3.5-4,, with strongly
attenuatedhilar portion which is either straightor curved towards the inside or out-
side and then assuminga sickle or tilda shape, otherwise ventricose, hyaline, smooth,
inamyloid. Hymenium:basidia 25-30 X 5.5-7,, clavate, 4-spored;basidioles fusoid;
cystidia none or few and then not differentiated(cystidioles), 30-44 X 7.5-8.5jA,
neither metachromaticnor opaque. Hyphae pseudoamyloidwith clamp connections.
Corticallayers: epicutis of the pileus hymeniform, consisting of broom cells which are
hyaline to reddishbrown or melleous, 17.5-20.5 X 7-7.5., mottled in scalp view, set-
ulae 5.5-7.5Mlong obtuse.
On dicotyledonous, dead, fallen twigs and branchletsof the humid forest in the
subtropicalzone, solitary or in small groups in the shade.
MATERIALSTUDIED. ARGENTINA.Tucuman,Taff: ParqueAconquija,13 Mar 1951,
Singer T 1460 (LIL) type; T 1477 (LIL) paratype(cf Singer1965, p 258, footnote!).

117. Marasmiustageticolor Berkeley, Hook. Jour. Bot. 8: 136. 1856.

TYPE. Spruce 37, from Brazil.
Pileus red (deep blood red, purple red or carminered, dried "Macracaibo"M&P)
with white or ochraceous-whitishor (dried) ocher radialstripes, glabrous,smooth on
the disc, sulcate on the margin,campanulate-convex,later convex-repandand often um-
bilicate, 4.5-18 mm broad. Lamellaewhite with white edges, distant, not or scarcely
intervenose,narrowlyadnexed to free. Stipe at first often red, later umber, smooth,
glabrous,equal or subequal,hollow, 20-50 X 0.2-0.7 mm; basal mycelium abundant,
pure white to pale fulvous. Context white, thin, inodorous.
Spores 15-20 X 3-4,, mostly 17.5-19 X 3.5,, acicular to subclavate,smooth,
hyaline, inamyloid. Hymenium:basidia 16.5-21.5 X 6.5-8,A, 4-spored;cystidia none;
cheilocystidialike the epicuticularbroom cells but all hyaline to subhyaline. Hyphae
hyaline in the pileus, with thin to barely firm wall, with clamp connections, pseudoa-
myloid. Corticallayer: epicutis of the pileus hymeniform, broom cells of the Siccus-
type, main body 17-17.5 X 8-91u,hyaline, with setulae erect and apical, some some-
times slightly spreading,acute or subacute, pale melleous to subhyaline,mostly hy-
aline in the pale-stripedzones, deep melleous in the red-stripedzones, 5.5-9 X 1.2-2.5u.
On twigs as well as on coriaceousleaves, fallen to the ground, in tropical forest.
MATERIALSTUDIED. MEXICO:Chiapas,between Ixtacomitanand Solusuchiapas,1 Aug
1969, SingerB 8804 (F). VENEZUELA. Sucre:NWof Irapa,8 Aug 1972, Dumont et al VE4300
(NY). BRAZIL. Amazonas:"Panure"(Ipanur6= Sao Jeronimo),Spruce37 (K) type.
Singer(1965), pi 14, fig 77; Berkeley, (1856) pl 5, fig 1.

118. Marasmiuspoecilus Berkeley, Hook. Jour. Bot. 8: 137. 1856.

TYPE. Spruce 139 (part), from Brazil.
Pileus purplishbrown to fulvous brown (dried almost spadiceous)with radialwhite
to yellow stripes, deeply sulcate to the umbilicate center, glabrous,campanulate-convex
and eventually with uplifted margin,8-25 mm broad. Lamellaedeep cream with not dis-
colorous edges, medium broad, distant, free to adnexed. Stipe fulvous below, paler above
or almost entirely light umber, glabrous,equal or subequal,35-80 X 0.2-0.7 mm; basal
mycelium dirty pallid to brownish, dried mostly fulvous, varying from scanty to abund-
ant and strigose. Context white, thin, inodorous.
Marasmius 167

Spores 12-20.3 X 3.3-4.2 p, smooth, hyaline, inamyloid. Hymenium: basidia 20-22

X 7 p, 4-spored;cystidia none or basidiomorphous,in form of cystididoles, 23-34 X 7-11
p1,not projecting,hyaline to yellowish hyaline, very slightly yellowish resinous incrusted,
without contents, not metachromaticin cresyl blue mounts; cheilocystidia like the epi-
cuticular broom cells and mostly hyaline with hyaline to pale fulvous setulae. Hyphae
and cortical layer as in the precedingspecies.
On dead coriaceous dicotyledonous leaves fallen to the ground in tropical forest.
MATERIAL STUDIED. VENEZUELA. Amazonas, Alto Rio Orinoco near Guiaka, 175 m al-
titude, 30 Mar 1953, Maguire & Wurdack 34796 (NY). BRAZIL. Amazonas, "Panure" (Ipanure =
Sao Jer6nimo), Febr 1853, Spruce 139, part (K), type. BOLIVIA. Beni, Guayaramerin, 14 Feb 1956,
Singer B 1981 (LIL).
This species is extremely close to the precedingone which I did not separatespec-
ifically in the past. However, the colors of the stipe and pileus are different in these
species, the stipe of I. poecilus perhapssomewhat longer, and the spores slightly broader
in relation to length. Since the color of the fresh specimens is also different, the brown
color of the stripes not appearingthrough drying, and the collector (Spruce) collected
them as different species under different numbers,I prefer to describe them separately.
I believe the type of Marasmiuspoecilus to be a mixed collection. The leaf with
five good and three imperfect carpophoreswhich I consider the holyotypes, undoubtedly
bears the species describedabove as M. poecilus. Berkeley indicated the stipe as insit-
itious which it is not. In some carpophoresof the type the stipe has a poorly develop-
ed basal mycelium but this is always visible under a lens.

119. Marasmiusphaeus Berkeley & Curtis, Jour. Linn. Soc. Bot.

TYPE. Wright18, from Cuba.
Pileus dark reddish brown (slightly deeper and rustier than "mohawk"when fresh)
with white radialstripes, glabrous,sulcate to the narrow(sometimes subumbonate)disc,
campanulateto conic-applanate,6-9 mm broad. Lamellaeon edges concolorous with
pileus, sides fresh white, distant, about ten, ascendant, eventually more horizontal, nar-
row to broad, adnexed. Stipe umber or light date brown when fresh, often darkerin
the herbarium,glabrousand smooth, shining, setose, subequal, 12-30 X 0.2-0.8 mm;
basal mycelium scanty to well developed, strigose, white or whitish, dried sordid gray-
ish white or whitish. Context very thin, inodorous.
Spores 16-22 X 3.8-4.2 p, lanceolate-claviform,hyaline, smooth, inamyloid. Hy-
menium: basidia 25 X 5 (according to Dennis 40 X 5-6 - ?); cystidia none; cheilo-
cystidia exactly like the epicuticularbroom cells. Hyphae pseudoamyloidwith clamp
connections. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells
of the Siccus-type, 15-16.5 X 6-8.5 but if setulae divergesomewhat or main body
forked spreadingto 9-11 p wide, setulae 5.5-8.5 X 1.2-1.8 (at base)p, gold brown to
succineous, on few cells hyaline (except where the stripes are: here all hyaline), obtuse
to acute, spinulose or rod-shaped.
On dead bark and on wood of standinglive dicotyledonous trees in rain forest,
MATERIAL STUDIED. CUBA: Wright 18 (FH, K) type. BOLIVIA. Beni: Vaca Diez, Ivon,
3 Apr 1956, Singer B 2428 (LIL).
ILLUSTRATION:Singer (1965) pl. 14, fig. 79.

120. MarasmiusbambusiniformisSinger, sp nov Fig 59.

TYPE.Singer B 7480, from Ecuador.
168 FloraNeotropica

Pileo ferrugineo,4.5-7 mm lato; lamellis candidis;stipite 18-30 mm longo. Sporis

15-18.5 X 3.7-5 ,; cystidiolis frequenterpraesentibus. Ad folia delapsamono- et dico-
tyledonea. Typus in hylaea aequatorianalectus et in F conservatus.
Pileus ferruginous"chinook," dried "rust sorolla" to "Gypsy" M&P,not or slightly
striped (between sulci) orange (11 H 9, M&P)glabrous,sulcate, campanulate-convex,
about 4.5-7 mm broad. Lamellaepure white, often becoming brownish white when dried,
with edges concolorous with the pileus, narrowto broad, few. Stipe chestnut or ferruginous,
at first with white apex, smooth, glabrous,with slightly thickened base which is pubes-
cent from the scanty basal mycelium, 18-30 X 0.2 mm, base 0.3 mm across. Context
very thin, inodorous.
Spores 15-18.5 X 3.7-5 p, fusoid, smooth, hyaline, inamyloid. Hymenium:basidia
13-20 X 5-102, 4-spored, basidioles fusoid; some cystidiole-like sterile cells often present,
these 20-24 X 5-9 p, versiform,usually clavate with or without lateral or apical appendage,
hyaline, thin-walled,arisingfrom the same level as the basidioles;cheilocystidia like the
epicuticularbroom cells. Hyphae in the trama of the pileus hyaline and with clamp
connections, thin-walled,pseudoamyloid. Corticallayers: epicutis of the pileus consis-
ting of a hymenial layer of broom cells with hyaline main body and golden melleous to
melleous setulae, in some cells main body golden melleous or melleous, in some setulae
hyaline, main body 7-15 X 4.5-10 p, setulae 2-6 X 0.2-1.5 ,.
On fallen leaves of Mono- and Dicotyledones, and on grassculms.
MATERIAL STUDIED. BRAZIL. Pernambuco: Camaragibe, 7 Jul 1960, Singer B 3281
(BAFC). ECUADOR. Napo, Lago Agrio, 16 May 1973, Singer B 7480 (F), type.

121. MarasmiusmazatecusSinger, sp nov Fig 60.

TYPE. SingerM 8368, from Mexico.
Pileo aurantiaco-ferrugineo,sulcato, 17 mm circiter lato; lamellis albis, ad aciem
ferrugineo-aurantiacis, distantibus,haud collariatis,stipite castaneo, 20 X 0.8 mm, basi
strigoso. Sporis 17-21 X 3.5-4.5 ,y; hyphis pseudoamyloideis. Ad lignum dicotyledoneum
in Mexico. Typus in F conservatusest.
Pileus orange ferruginous,deeper rusty-reddishin the center, dried "Caldera"(M&P)
in center and striae, "Monterrey"(M&P)between striae, deeply sulcate, glabrous,convex,
mostly with a small umbilicus when dried, eventually with uplifted margin, about 17 mm
broad. Lamellae white with rusty orange edge, distant (9 through-lamellae,some narrow
and short lamellulae), rathernarrow (2 mm), leaving an orange-ferruginousband at the
apex of the stipe from which they are free when dried. Stipe chestnut, dried brown,
fresh with white apex underneaththe orange-ferruginousband appearingin dried and
dry material, quite glabrousexcept for the moderately developed strigose-woollyfulvous
basal mycelium, about 20 X 0.8 mm. Context white in the pileus, inodorous.
Spores 17-21 X 3.5-4.5 p, fusoid, hyaline, smooth, inamyloid. Hymenium: con-
sisting entirely of basidiaand basidioles;cheilocystidia making the edge heteromorphous,
in shape like the epicuticularbroom cells. Hyphae of the pileus tramahyaline, with
clamp connections pseudoamyloid. Cortical layer: epicuticularbroom cells in hymen-
ial arrangement,main body hyaline or orange, with thin to moderately thick wall,
10-28 X 6-8 p, cylindric to clavate, more rarelyventricose, setulae 2-13 X 1-1.8 1, or-
ange-brown,orange-yellow,yellow, in very rarecases absent.
On dicotyledonous wood.
MATERIAL STUDIED. MEXICO. Oaxaca: San Andres Hidalgo, Sierra Mazateca, 8 Jul 1969,
Singer M 8368 (F), type.
Marasmius 169

122. Marasmiuslongisporus(Patouillard& Gaillard)Saccardo,Sylloge Fungorum9: 68.

AndrosaceuslongispontsPatouillard& Gaillard,Bull. Soc. Mycol. France4: 20. 1888.
TYPE. Gaillard 21, from Venezuela.
Pileus at first between "mast color" and "raw sienna," soon fresh and dried och-
raceous tawny to deep ferruginous("henna" with the marginbetween "henna"and
"chutney" or between "feuille morte" or "rust sorolla" and "Peruvianbr" or pl. 13
A 12, M&P),the marginoften paler, not or dried indistinctly narrowlyradiallystriped
(ochraceous) but distinctly long-sulcate,glabrous,conic to campanulate,often ? um-
bonate, 4-10 mm broad, 5-6 mm high. Lamellaewhite, subdistantto distant, when
dried with a narrowly discolorous(color of the pileus) edge, at least when seen under
a lens, rather narrow,at first ascendant,adnexed to free. Stipe at first white, soon
succineous or yellowish umber, tending to blackish chestnut at base, but mostly re-
maining white at apex, smooth, macroscopicallyglabrous,under a strong lens at first
finely pruinate but soon glabrescent,filamentous-setose,shining, 27-60 X 0.2-0.6 mm;
basal mycelium abundant, tomentose, sordid whitish tending to orange when dried.
Context white, thin, inodorous.
Spores (12-)13.5-19 X 3-5 1,, mostly over 14.5 p long and under 4,Ebroad, cla-
vate to fusoid, smooth, hyaline, inamyloid. Hymenium: basidia 20-28 X 8.5-11 -,
4-spored:cystidia none, but some of the basidioles often remainingsterile, fusoid,
ampullaceousor constricted, thin-walled,few; cheilocystidia like the epicuticular
broom cells or entirely hyaline. Hyphae of the trama of the pileus hyaline, thin-walled,
with clamp connections, pseudoamyloid. Cortical layer: epicutis of the pileus hymeni-
form, consisting of broom cells, these with a main body 5-19 X 3-9(-14)u, hyaline,
but in upper portion mostly pale melleous to melleous, with a thin or ratherthin (up
to 0.5 p) wall, setulae constantly present, 2.5-10 X 0.5-2 ,, golden yellow, golden
ferruginous,melleous brownish or ferrugineous-melleous,conical, acute or subacute;
covering of young stipe consisting of scattered small broom cells.
On wood and bark of dicotyledonous trees and on very decayed wood in the
leafmold in tropical and tropical-montane(to subparamo)forests and scrub.
MATERIAL STUDIED. MEXICO. Veracruz: 7 km south of Montepio, 19 Jun 1969, Singer
.1 8027 (F); Chiapas, between Finca Sospiro and El Pozo, 4 Aug 1969. Singer M 8948 (F). VEN-
EZUELA. Bolivar: Puerto Zamuro (Rio Orinoco) 25 May 1887 Gaillard 21 (FH), type. ECUADOR.
Pichincha. Quito. Quebrada de Miraflores at 2950 m alt, 24 Apr 1973, Singer B 7120 (F).
This is a reviseddescription since my earlierdescriptionwas based only on the type
and fresh material from Bolivia. The latter, however, later turned out to be 1. bert-
eroi var berteroi, whereas newly collected Mexicanand Ecuadorianmaterialis obviously
identical with the Venezuelan type.
ILLUSTRATION. Singer (1965) pl 14, fig 78.

123. Marasmiusonoticus Singer, sp nov

Marasmliu s onoticus Singer, Sydowia 18: 252. 1965, nom nud
Type. Singer T 2995, from Argentina
Pileo aurantiacovel aurantiaco-rufescente,hemisphaerico,5-6 mm lato; lamellis
albis, distantibus;stipite castaneo, 30 X 0.5 mm. Sporis 13.8-18 X 3.5-4.8 j; cheilo-
cystidiis et cellulis epicuticularibustypi Marasmiisicci. Ad lignum dicotyledoneum in
silvis subxerophyticis. Typus in LIL conservatusest.
Pileus orange to orangerufescent ("talavera",margin"doubloon"), deeply sulcate,
glabrous,unicolorous, hemispheric,5-6 mm broad. Lamellaewhite, distant, ten, rather
170 Flora Neotropica

narrow, free. Stipe deep chestnut below and slightly paler above, glabrous,smooth,
shining, equal, setose, 30 X 0.5 mm; basal mycelium white, not extensive. Context
very thin, inodorous.
Spores 13.8-18 X 3.5-4.8p, hyaline, smooth. Hymenium: cystidia, none seen;
cheilocystidia like epicuticularbroom cells; hyphae hyaline, psuedoamyloid. Cortical
layer: epicutis of pileus hymeniform, consisting of broom cells, 15-16.5 X 7-8.5 u,
sometimes spreadingbecause of divergenceof setulae to a diameterof 9-11 p, setulae
5.5-8.5 X 1.2-1.8u (at base), golden succineous, few hyaline, obtuse orsubacute, spin-
ulose or rod-shaped.
On dead dicotyledonous wood in subxerophytic woods.
MATERIAL STUDIED. ARGENTINA: Tucumnan:Burruyacuf, 5 kim south of town, 19 Feb
1957, Singer T 2995 (LIL) holotype.
It is possible that an additional collection (Argentina:Santiago del Estero, El Sal-
vador, July 1962, D. Foguelman, BAFC) also belongs here but this is only 15 mm high
and 1.5 mm broad and may be a nanism of M. berteroi.
Illustration. Singer (1965) pl 14, fig 76.

124. MarasmiusrubricosusMontagne,Ann. Sci. Nat. Bot. IV. 1: 110. 1854.

TYPE. French Guyana (not found).
Pileus deep brown ("Mandalay")in center when fresh, with light cinnamon (12-
D-8) margin,in dried condition deep ferruginousin center and deep chestnut brown on
margin,glabrous,densely striate, sulcate on margin,applanatewith a very obtuse umbo,
dried more convex since margin tends to incurve broadly on drying, about 37 mm broad.
Lamellaesordid whitish, drying to light brown, narrow, fresh free and moderately broad,
drying narrow and subfree, close with numerous lamellae of various lengths. Stipe chest-
nut, glabrous or subglabrous,about 50 X 1.5 mm; basal mycelium very abundant, to-
mentose, forming a socle-like broadeningof the base, white with a slight fulvous shade
wlien fresh, pale fulvous when dried. Context inodorous.
Spores 15-18 X 4-5.5 ,, hyaline, smooth. Hymenium: basidia 22 X 4.8 M,4-spored,
clavate;cystidia not numerousand not distinctly differentiatedbut occasional cystidioles
occurring(ampullaceousto clavate, 26-27 X 10-11 p); cheilocystidia like epicuticular
broom cells. Hyphae pseudoamyloid. Cortical layers: epicutis hymeniform, consisting
of broom cells with a main body, clavate or ventricose, hyaline to melleous hyaline and
setulae either brown or hyaline (therefore surface in scalp view mottled), 3-7 X 1-1.8 ,
spinulose the entire broom cell 12-25 X 5-9 ,.
On dead wood.
MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez: Guayaramerfn, 14 Mar 1956, Singer B
1984 (LIL).
Only one collection is referredhere on the strength of Dennis's remarks(1951a, p
420) which would fit our collection perfectly. These remarksrefer to authentic mater-
ial sent to Berkeley by Montagne.
ILLUSTRATION. Singer (1965) pl 14, fig 75.

125. MarasmiusrhabarberinusBerkeley, Hook. Jour. Bot. 8: 135. 1856.

TYPE. Spruce 140, from Brazil.
Pileus rhubarbyellow, orange (e.gr. 1 l-D-11 M&P)dried bright orange fulvous (e
g "spice"), smooth or reticulate-rugose,distinctly sulcate-furrowedin marginaltwo-thirds
when mature or dried, glabrous,convex, with or more rarely without a small umbo, 7-31
mm broad. Lamellaepallid, equal or with some very short lamellulae intermixed, not
Marasmius 171

intervenose, rathernarrow to ratherbroad, (up to 4.5 mm broad), free or even remote-

free, distant. more rarely sub-distant(15 through-lamellaeor more). Stipe chestnut,
glabrous,smooth, equal, 18-60 X 0.2-1.4 mm; basal mycelium well developed, strigose,
white or becoming pale fulvous. Context thin, white, inodorous.
Spores 14-21 X 2.3-3.8,, cylindric-clavateto cylindric-fusoid,straightor very
slightly curved in lower portion when seen in profile or at least more convex on outer
than on inner side, hyaline, smooth, inamyloid. Hymenium: basidia narrowlyclavate,
35-36 X 6-6.8 p, 4-spored;basidioles fusoid; cystidia none; cheilocystidia hyaline to
pale melleous in upper portion, otherwise like epicuticularelements, making the edge
of the lamellae heteromorphousor nearly so. Hyphae of pileus and hymenophoraltrama
hyaline, thin-walledand filamentous, with clamp connections, pseudoamyloid. Cortical
layers: epicutis of pileus consisting of broom cells of the Siccus-type, these either all
hyaline or hyaline with golden melleous, erect setulae, both color types mixed in a
definite pattern and therefore epicutis marbled-dottedin scalp preparations. Stipe with-
out any broom cells or hairs.
On dead fallen leaves in tropical and subtropicalforest.
MATERIALSTUDIED. BRAZIL. Amazonas:Rio Purua(Purus),Trail 161 (K), authentic;
"Panure"(Ipanure= Sao Jeronimo), Spnuce140 (K), holotype (FH), isotype (K). ARGENTINA.
Misiones:Arroyo PirayGuazi, 24 Feb 1960, R. T. Guerrero73 (LIL).
It is the medium-sizedcarpophoreof the holotype collection which yielded the
spores which are identical with those found in the Argentinematerial.A Colombian
collection which differs from the other collections in more chestnut and rusty orange
pileus, yellowish, adnexed lamellae, and slightly largersize of the carpophores,was re-
ferred to AMarasmius berteroi var major in spite of the fact that the spores were some-
whlatlargerthan usual so that it may have keyed out here (with Al. rhabarberinlus).
ILLUSTRATIONS.Singer (1965) pi 13, fig 71; (1958a) fig 23.

126. Marasmiusberteroi (Leveille) Murrill,North AmericanFlora 9: 267. 1915.

berteroiLeveille,Ann. Sci. Nat. III. 2: 177. 1844.

126a. Marasmiusberteroivar berteroi

TYPE. Bertero, from Puerto Rico.
Pileus broadly shallowly sulcate-furrowed,campanulate-convex,8-15 mm broad,
otherwise like the type variety. Lamellaewhite to cream, not intervenose,subdistant
to (mostly) distant, edge concolorous with sides, eleven to sixteen, ascendantor sub-
horizontal(oblique), moderately broad to broad, subventricoseto ventricose. free.
Stipe 20-57 X 0.1-1 mm, otherwise like that of the var najor.
Spores 11.3-16.2 X 3-4 . Hymeniumlike in var major, cystidia occasionally
observed but inconspicuous,mostly not reachinglevel of sterigmataand basidiomorphous
or like basidioles and not differentiatedin cresyl blue or phoxine mounts but some
elements of the hymenium apparentlyremainingsterile and showing slightly thicker
walls and slight incrustations(cystidioles?); cheilozystidiaall hyaline, otherwise like
epicuticularbroom cells. Hyphae clearlymetachromaticin cresyl blue mounts in the sense
that most hyphae are are vinous red as contrastingwith the bluishhymenialelements, hy-
aline, pseudoamyloid,not gelatinized, with clamp connections, some slightly thick-
walled. Cortical layers: epicutis of pileus hymeniform,consisting of broom cells,
their main body 10-22 X 5-9y, clavate, mostly hyaline to subhyalinebut in some
broom cells concolorous with setulae and strongly pigmented and then slightly thick-
walled, and strongly contrastingin cresyl blue mounts where they are violet (this
172 FloraNeotropica

type of elements evenly distributedamong the other paler broom cells and giving
the scalp preparationsa mottled appearance),setulae hyaline to golden melleous,
3.5-3.8 X 0.7-1.5 (at base),, subacute to acute.
On forest litter adheringto mixtures of sticks and fallen leaves of various tree
species rotting in the woods and forests, singly or in closely joined pairs.
MATERIALSTUDIED. PUERTORICO. "Agaricusfuillus Bertero,"Bertero
(HerbariumPersoon (L) Type. BOLIVIA. La Paz: Nor-Yungas,Charobambaat 1200-1300 m
alt, 30 Jan 1956, SingerB 725 (LIL). ARGENTINA. Salta: Oran, 3.5 km west of Rib Blanco
bridge, 17 Mar 1955, Singer T 2310 (LIL);Tucuman:CiudadUniversitaria.1000 m alt, 26 May
1957, Singer T 3080 (LIL);Taficillo, 1500 m alt, 23 Feb 1955, Singer T 2219 (LIL);Yerba
Buena,600 m alt, 12 Feb 1955, Singer T 2159 (BAFC).
This variety differs from the following variety in habit and size. Mlarasnmius
flavofuscus Berkeley & Curtis from the Bonin Islandsis close. The type (FH) differs
mainly in the color of the pileus as described.
ILLUSTRATION. Singer (1965) p 14, fig 74.

126b.Marasmiusberteroi var major Singer, Sydowia 18: 334. 1965 (ex errorep 249
"var macrus").
TYPE. Singer T 2886, from Argentina.
Pileus bright orange fulvous, orange, orange red, rufous ferruginousor intensely
ferruginous(between "feuille morte" and "Eldorado"or between "feuille morte" and
"Peruvianbr.", "Agate," "Kobe", "henna",4-A-i2, towards marginsometimes "sun-
burst";"Ponce de Leon," ll-L-10, or "Saratoga"(M&P)on center and ribs, sometimes
reaching"Maracaibo"or deeperand richer than "caldera";color often reachingtones
deeper and richer than those of Maerz& Paul (such as Villalobos SO, 12 , 8), dried
similarto the fresh color, or "gold pheasant"or 12-F-9, glabrous,not viscid, at first
over half, soon over 4/5 to 5/6 of radius,sulcate to deeply furrowed, otherwise (on
disc) either smooth or rugose-venose,campanulateor conic-truncatewith flattened
disc, later convex and often umbonate, eventually convex-applanateor with uplifted
marginand tending to become obtuse, frequently even with depressedcenter, 10-56
mm broad. Lamellaecream, pale cream, cream whitish, yellow white, milk white,
often becoming pale orangeyellow when dried, usually more yellow if older, the
edges usually concolorous, rarely few lamellaemostly not intervenose,narrowlydis-
colorous, more rarely sparselyto densely intervenose,all entire or with few lamell-
ulae (through-lamellae9-19; lamellulae0-18), ventricose or not, narrowto broad,
3-12 mm broad, sometimes not quite reachingthe margin,narrowlyadnexed to
free, not collariate, spore print pure white. - Stipe reddishbrown to chestnut be-
low shading deeper and deeper towards the base, less deep and sometimes almost
umber further up (below "mohawk," "Mandalay,""Hindu," above "broncho"),
apex generally concolorous with lamellae (whitish to cream) but sometimes yellow,
especially when young, ("lemon yellow," 10-I/L-2/3 M&P),base often almost black,
glabrous,smooth, shining or opaque, at first taperingupwardslater sometimes equal,
narrowlyhollow (with a thin, rigid cortex and a tubulose cavity) 30-80 C 1-3 mm,
basal mycelium well developed, tomentose, white, more rarely buffy white.
Context white, in youth often lemon yellow in the apex of the stipe, rather
tough, even hard in stipe when old, inodorous.
Spores (8-)9.5-15.3 X 2.7-4 p, fusoid-ellipsoid, hyaline, smooth, inamyloid.
Hymenium: basidia 20-30 X 5.5-7 ,, clavate, 4-spored, hyaline, some opaque;
basidioles fusoid; cystidia (if present) not morphologicallyor chemically differen-
tiated from the basidioles;cheilocystidia like the epicuticularbroom cells but some-
Marasmius 173

times rather scattered and rare, with obtuse, sometimes long, erect setulae, more
rarely rathernumerousbut not making the edge heteromerous(intermixed with
basidia and basidioles). Hyphae, all thin-walledin pileus, everywherepseudoamy-
loid and with clamp connections, in old specimensmany hyphae, especially in stipe,
with thick walls. Cortical layers: epicutis of the pileus hymeniform, consisting of
broom cells, these 23-26 X 6-8.2 1,, of two types, (1) with hyaline and thin-walled
main body and hyaline to melleous setulae, (2) with the main body melleous to
bright ochraceousferruginousat least in upper portion which also may be thick-
walled; setulae apical and erect, often ratherlong, especially in type 2 some rather
coarse, e g about 1.7 , thick below, but in type 1 usually much more slender,pro-
jecting 5-11 .; the two types regularlydistributedand scalp appearancemottled.
On branchesand trunks fallen to the ground(dicotyledonous trees) and on leaf-
mold of dicotyledons, monocotyledons, conifers, in montane, subtropical,tropical-mon-
tane forest, solitary or gregariousbut scarcelycespitose (Podocarpus,Gramineaespp,
Alnus, Phoebe, etc).
MATERIALSTUDIED.BELIZE. El Cayo, Valentin, 28 Jul 1936, E. B. Mains3660 (MICH).
COLOMBIA.Valle: Buenaventura,Calima,22 Apr 1968, SingerB 6295 (F), a large-sporedform
see annotationbelow. ECUADOR. Napo: Lago Agrio, 17 May 1973, SingerB 7518 (F); Tungurah-
ua, Rio Topo, 30 Apr 1973, SingerB 7225 (F). BOLIVIA. La Paz: Nor-Yungas,Rio suapi, 1250-
1300 in alt, 9 Feb 1956, SingerB 1043 (LIL);Rio Yariza,23 Feb 1956, SingerB 1394 (LIL);Char-
obamba,30 Jan 1956, Singer B 693 (LIL). PERU. Cusco, Paucartambo,Atalaya,Carbon,700 m
alt, Vargas13519 (LIL). BRAZIL. SantaCatarina:Porto Novo, 1928, J. Rick (FH). ARGENTINA.
Tucuman:Anta Muerta,1100 m alt, 10 Apr 1949, Singer& Digilio T 418 (LIL);La Angostura,1800
m alt, 1 May 1949, Singer T 540 (LIL);Taficillo, 1700 m alt, 4 Mar 1951, Singer T 1368 (LIL);
Rio de los Sosas, 900 m, 4 Feb 1955, Singer T 2126 (LIL);CiudadUniversitaria,1100 m alt, 14
Jan 1957, Singer T 2886 (LIL) type; T 2887 (LIL);23 Jan 1957, T 2879 (LIL);26 Jan 1957, T
2913 (LIL); 1 May 1957, P. Wygodzinsky,det. R. SingerT 3073 (LIL);Misiones:Iguazu,Ref. "Yag-
uarete ," 29 Feb 1960, R. T. Guerrero133 (LIL).
The collection from Colombia (B 6295) agreesmacroscopicallywith var major but
may key out with Marasmiusrhabarberinusrather than M. berteroi because the spores
were 12-17 X 3 p, thus largerthan usual in M. berteroi var major. Since this collection
differs in severalaspects includingthe habitat on wood ratherthan dead leaves, from M.
rhabarberinus,it has been considered a large-sporedform of M. berteroi but it may re-
quire a special subspecific taxon.
The collection from Ecuador(B 7518) agreeswith var major except for the dis-
colorous or partly discolorous(concolorous with pileus) edge of the lamellae and may
also be a variety in its own right or subvarietyof var major.
ILLUSTRATION. Singer(1965) pl 14, fig 73.

127. Marasmiusconvoluticeps Singer, sp nov Fig 61.

TYPE: G. & L. T. Eiten & G. M. Felipe, from Brazil.
Pileo dilute brunneo, siccando laete ferrugineo,superficieconvoluta gaudente, us-
que ad 15 mm lato; lamellis ad aciem haud discoloribus,confertis vel subconfertis,lib-
eris vel subliberis;stipite atrobrunneo,ad basin strigoso. Sporis 12.5-16 X 3-4,; cys-
tidiis nullis. Ad lignum emortuum in Brasilia. Typus in F conservatur.
Pileus fresh light bright brown becoming bright ferruginous("rust, sorolla br" to
"gypsy" M&P)when dried, not or scarcely sulcate when fresh or dry but with con-
voluted gyrose venose-rugosesurface almost all over, glabrous,broadly and obtusely
conic-campanulate (much like M. haematocephalus in shape and size) sometimes umbon-
ate, eventually sometimes more repand, up to 15 mm broad. Lamellae cream with a
purplish tinge, the edge not discolorous, close to medium close, medium broad, free
174 FloraNeotropica

to subfree. Stipe dark brown, subglabrous,under a strong lens very finely pubescent
or subpruinose,smooth, setose-equal,up to 25 mm long, 0.3-0.6 mm broad; basal
mycelium strigose, tending to be or to become fulvous. Context thin, white, inodor-
Spores 12.5-16 X (2.5-)3-4 u, fusoid, on apex and base sometimes slightly re-
curved towards the same side, smooth, hyaline, thin-walled,with severalsmall oil
droplets,inamyloid. Hymenium: basidia4-spored;basidioles fusoid; cystidia none;
cheilocystidia like the epicuticularbroom cells but subhyaline throughout. Hyphae
of pileus tramahyaline, of stipe brown, parallel, thick-walled. Corticallayers: epic-
utis of pileus consisting of broom cells with a main body 10-12 X 5.5-7 p, thin-walled,
brownish to hyaline, claviform;setulae 4.5-12 X 1-1.8 p, pale golden-ferruginous,acute
or subacute, spinose-conical;covering of the stipe consisting of widely spaced (long
stretches of surface layer without any covering), dermatocystidioidsmall hairs which
are 9-27 X 3.8-7.2 j, hyaline or subhyaline,claviform,more rarelycylindrical,with
thin to more rarely thick wall, entire or occasionally with one short inconspicuous
apical apiculus.
On dead wood (wet rotting logs).
MATERIAL STUDIED. BRAZIL. Amazonas: Municipalidad Manaus, Itacoatiara road in
virgin "terra firme forest," 30 Jan 1963, G. & L. T. Eiten & Gil M. Felipe 5254 (F), type.
This is close to M. matrisdeiwhich differs, especially with regardto the type of
coveringon the stipe.

128. MarasmiusmatrisdeiSinger, Sydowia 18: 338. 1965.

TYPE. Singer B 2523, from Bolivia.
Pileus when quite fresh beautifully light orange(10-H-7) to ochraceousorange
(11-L-9) in the central portion with light ochraceous orangemargin( l-G-9), soon
fading to melleous brown ("mast color" to "Punjab")in the center, dried "Yucatan"
on marginand "Sudan brown" in center, slightly hygrophanousand bleaching-fading
parts becoming pale ochraceouswhen dry (not dried), sulcate on marginover one
fifth to one third of the radiusboth in young and in old specimens,between sulcate
zone and center usually rugose and frequently striped, glabrous,at first with incurved
margin,obtuse, or more often subumbonateto umbonate, campanulate,soon repand,
becoming flat-convex, or flat conical, 7-17 mm broad. Lamellaepale cream, close,
narrow, adnate. Spore print pure white. Stipe at first entirely pallid white, then
ochraceousbrown with white apex, entirely very finely pruinate, sometimes with a
small socle-like base, usually taperingupwards,about 25 mm long, 0.7-1.5 mm broad;
basal mycelium white. Context white or whitish, unchanging,very thin in pileus,
Spores (9.7-)11-15 X 4-6.3 u oblong (not lanceolate), thin-walled,easily collapsing,
hyaline, smooth, inamyloid. Hymenium: basidia 27.5-31 X 6.8-7 p, 4-spored;cystidia none;
cheilocystidia like the broom cells of the epicutis of the pileus. Hyphae: subhymenium
hyaline, consisting of narrowelements which are irregularlyarrangedand hyphous but
multiseptate (septa close); hymenophoraltramaand trama of pileus consisting of hyaline
hyphae which are all homogeneous but tend to be yellowish in the upper part of the
pileus. All hyphae with clamp connections and pseudoamyloid. Corticallayers: epicut-
is of pileus hymeniform, consisting of broom cells of the Siccus-type,main body mostly
clavate, more rarely cylindric or irregular,setulae about 6.5 p long, and alternatingly
(groups of broom cells all hyaline, others all golden melleous) pigmented or not (thus
surface mottled in scalp view), taperingupwardsbut mostly obtuse, more rarely acute,
Marasmius 175

few elongated to 18 plength, total measurementsof broom cells of pileus 24-31 X 7.5-9
p; surface of stipe with numerousbroom cells but the main body often strongly reduced
and often nothing but setulae (sometimes elongated up to 23 p and then setiform!) visible.
On fallen stems and branchletsof both mono- and dicotyledonous plants in trop-
ical rain forest.
MATERIAL STUDIED. BOLIVIA. Pando: Madre de Dios, Las Piedras, 5 Apr 1956, Singer
B 2523 (LIL), type.
ILLUSTRATION. Singer (1965) p 14, fig 80.

129. Marasmiusneglectus Singer, sp nov Fig 62.

TYPE. Singer B 6381, from Colombia.
Pileo atroferrugineo-rubido,magis fulvidus interstrias,papillato, 6-9 mm lato; lam-
ellis dilute grisello-ochraceis,acie atroferrugineis,subconfertis,haud intervenosis;stipite
ad basin castaneo, strigose. Sporis 11-15 X 3-4 p; cystidiis perpauciset vix distinctis,
cystidioliformibus. Ad folia delapsa dicotyledonea in hylaea columbiana. Typus in F
Pileus deep-rusty-red-brown,deeper on center and striae, more rusty-orange-ful-
vous between striae, dried ratherdeep ferruginous-fulvous,occasionally slightly striped
when dried, with eventually slightly lobulate margin,glabrous,sulcate, conic to campan-
ulate, papillate, 6-9 mm broad. Lamellaelight grayish-ochraceouswith deep ferruginous
(discolorous) edge, broad, subclose (about 16 lamellae and many lamellulae),not inter-
venose, rounded-subfree. Stipe chestnut below, light grayish ochraceousat the apex,
glabrousbut not shining, subequal, 16-18 X 0.3 mm; basal mycelium light grayishoch-
raceous becoming fulvous on drying, radiallystrigose. Context thin, whitish in the
Spores 11-15 X 3-4 p (but few seen), fusoid, smooth, hyaline, inamyloid. Hymen-
ium: basidia 19 X 7.5 ,; basidioles fusiform;cystidia very few and poorly differentiated
from the basidioles, somewhat opaque but not deep-rooting(probably sterile basidioles-
cystidioles); cheilocystidia like the epicuticularbroom cells but always with hyaline
main body. Hyphae of the pileus tramahyaline, thin-walled,with clamp connections,
strongly pseudoamyloid. Corticallayers: epicutis of the pileus hymeniform, consisting
of broom cells, main body 11.5-15 X 6-9 u, hyaline to palest fulvous, thin-walled,vesiculose
to obconic or clavate; setulae 6-7.6 X 1-1.2 1, melleous-ferruginous,subacute to acute,
On the mid-nerveof dicotyledonous leaves fallen to the ground in tropical rain
MATERIAL STUDIED. COLOMBIA. Valle: Buenaventura, along Juanchaco road, 25 Apr
1968, Singer B 6381 (F), type.
This species has the aspect of the M. ferrugineus-groupbut lacks true cystidia.
Since only few spores were seen in the type, it may be that the spores reach somewhat
largersize in more abundantlysporulatingmaterial.

130. Marasmiushelvoloides Singer, sp nov Fig 63.

TYPE. SingerB 6298 a, from Colombia.
Pileo cinnamomeo,debiliterochraceo-radiato,magis brunneo siccando, sulcato,
convexo, centro umbilicato vel depresso, 2742 mm lato; lamellis albis vel albidis, sub-
confertis vel distantibus,haud intervenosis,anguste adnexis; stipite atrocastraneoapice
pallidiore,ad basin strigoso. Sporis 12-15 X 2.5-3 ,; cystidiisnullusvel inconspicuis
176 FloraNeotropica

rarisqueet cystidioliformibus. Ad lignum putridumin hylaea columbiana. Typus in

F conservatusest.
Pileus cinnamon, somewhat striped ochraceousin the depressionof the sulci,
dried more like 1. helvolus i e near "Sudan br." or "cocoa" with the stripes "nugget"
to "Antique br" (M&P),folded sulcate, glabrous,convex, often flattening on one side,
with slightly to strongly umbilicate or narrowlydepressedcenter, 27-42 mm broad.
Lamellaewhite or whitish, edges not discolorous, subclose to distant, not intervenose,
rathernarrow, narrowlyadnexed. Stipe dark chestnut or deep chocolate with paler
apex (pale cinnamon e g), glabrous to indistinctly pruinateunder a strong lens, equal,
48-72 X 1-2 mm; basal mycelium abundant, radiating-strigose,tending to whitish-
fulvous. Context of pileus white.
Spores 12-15 X 2.5-3 u, fusoid, smooth, hyaline, inamyloid. Hymenium:basidia
16.5-21 X 4.5-6.5 1; cystidia none or very inconspicuousand rare, cystidiole-like;
cheilocystidia with hyaline main body and pale ochraceousorange to hyaline setulae,
otherwise like the epicuticularbroom cells. Hyphae hyaline and pseudoamyloidin the
trama of the pileus. Corticallayer: epicutis of the pileus hymeniform consisting of
broom cells with a main body about 17 X 6-10 p, subhyalineto palest melleous, cla-
vate, setulae 3-6.5 X 1-1.5 ,, acute, narrowlyconical, subhyalineto deep ochraceous;
Coveringof the stipe scarcely developed, no broom cells present.
On very rotten wood in tropical rain forest.
MATERIAL STUDIED. COLOMBIA. Valle: Buenaventura, Calima, 22 Apr 1968, Singer B
6298a (F), type; B 6298b (F).
This is one of the largerspecies with very variablenumber of lamellae and should
not be confused with Marasmiusberteroi var major which differs in colors and no 174,
M. helvolus (compare description there).

131. MarasmiusgraminicolaSpegazzini,Anal. Mus. Nac. Buenos Aires 6 111. 1899.

TYPE. Spegazzini, from Argentina.
Pileus orange to deep rufous-orange(e g 10 G 10 to 11 F 11, M&P),sulcate, gla-
brous, sometimes umbilicate, at first obtusely conical to campanulate,later campanulate-
convex, 4-9 mm broad, 2.5-5 mm high. Lamellaewhite, with finely or intermittently
orange edges, distant (8-9 through-lamellaein most adult specimens), ratherbroad, all
equal, ratherbroad to broad, not intervenose, adnexed to subfree. Stipe umber to
bronze, succineous-umberor rufescent-bister,glabrousand shining, smooth, with at
first white apex, inserted to the pileus with an orange band, subequal, 20-40 X 0.2-
0.7 mm, basal mycelium moderately conspicuous, consisting of velutinous covering
or radiatingthin filaments, white. Context white in the pileus, thin, inodorous.
Spores (10-)12-13.8 X 3.3-3.7 ,, claviform-boatshapedor subfusoid, smooth,
hyaline, inamyloid. Hymenium consisting exclusively of basidia and basidioles;cyst-
idia none; cheilocystidia like the epicuticularbroom cells (but mostly with hyaline
main body and thin walls. Hyphae of the context of the pileus hyaline, thin-walled,
with clamp connections, pseudoamyloid. Cortical layer: epicutis of pileus hymeni-
form, consisting of broom cells with the main body 10-19 X 4-6.3pu, with thin or
somewhat thickened wall, hyaline or tawny orange to orange brown, the setulae 2.5-5.7
X 0.5-1 ., golden yellow to tawny orange, orangebrown, melleous, rarelyhyaline, con-
stantly present, subacute, rod-shapedto thin-spinulose.
On dead fallen leaves and stems of trees (e g Populus, Taxodium, Gramineaeand
other monocotyledonous plants, probably always basically on Gramineaebut passing
Marasmius 177

over to other materialnearby, fruiting in springand summer).

MATERIALSTUDIED. MEXICO.Veracruz,3 km South of Montepio,22 Jun 1969, Sin-
ger 41 8142 (F). ARGENTINA. Buenos Aires:El Cazador(Rio Parana)19 Dec 1965, SingerS
590 (BAFC):La Plata,Spegazzini(LPS), type.

132. MarasmiushaediniformisSinger, Bull. Jard. Bot. Bruxelles34: 363. 1964.

TYPE.de Witte 9432, from Zaire.
Pileus white, dried whitish (slightly radiallystriate with dirty pallid) with con-
colorous center, neither in age nor on drying becoming yellow, glabrous,sulcate, (mod-
erately deeply so) to the center, the center often venose, campanulate-convexthen con-
ves and sometimes with slightly depressedcenter in age, 18-37 mm broad. Lamellae
white with white edge, narrow,distant (10-17 through-lamellae,some lamellulae), often
not reachingthe extreme marginor reduced to veins there, not intervenose,subfree
to adnate. Stipe chestnut to bister, with white apex, glabrous,hollow, equal or taper-
ing downwards,25-45 X 0.4-0.8 mm, towards apex often as broad as 1.5 mm; basal
mycelium well developed, white. Context white, in pileus, thin to extremely thin (at
marginof pileus); odor none.
Spores 11-13.5 X 3-4 u, mostly 12.5-13.5 X 3-3.5 p, fusoid, some more claviform,
smooth, hyaline, inamyloid. Hymenium:basidia 18-19 X 7.5-7.7 1,, 4-spored:basidioles
fusoid: cystidioles few to rathernumerouswith the shape of the basidiolesbut sometimes
somewhat irregular,inconspicuousand thin-walled,not or scarcelyprojecting, 18-24 X
5-6.7 ,; true cystidia none; cheilocystidia exceptionally without setulae but normally
all or almost all in form of broom cells like the epicuticularbroom cells. Hyphae of the
pileus trama hyaline, filamentous, not gelatinized,thin-walled,with clamp connections,
pseudoamyloid;those of the lower portion of the stipe pigmented and parallelwith each
other. Cortical layers. Epicutis of the pileus hymeniform, consisting of broom cells
which are entirely hyaline (including the setulae), the main body thin- or firm-, but not
thick-walled,mostly clavate, sometimes cylindrical, 6-18 X 4.5-8(-11) ,, setulae 1.3-6 X
0.5-0.7 ,, subacute or acute, apical, some slightly oblique.
On fallen leaves of dicotyledonous trees in the litter of tropical forests.
MATERIALSTUDIED. ECUADOR. Napo, Shushufindi,240 m alt, 15 May 1973, Singer
B 7446 (F). ZAIRE. ParcNationalAlbert, aroundHoysha, 1050 m alt, de Witte9432 (BR), type.
Singer(1964b) fig 24, B-E.

133. MarasmiusproletariusBerkeley & Curtis, Jour. Linn. Soc. Bot. 10: 296. 1869.
Fig 64.
TYPE. Wright, from Cuba.
Pileus white when fresh but soon becoming cream white to pale cream, on drying
gilvous to pale ochraceouscinnamon in the center and gilvous-whiteon the margin,gla-
brous, with sulcate margin,dull convex, eventually applanate,2-5.5 mm broad. Lam-
ellae white, eventually cream, dried gilvous-whitish,with edges concolorous with the
sides, distant to medium distant, tridymous, with about 12 through-lamellae and many
lamellulae,not intervenose, rathernarrow,to broad, adnate, more rarely adnexed.
Stipe pallid-white,mostly avellaneousto chestnut from the base upwards,pallescent
on drying if chestnut and becoming pale stramineous or pale bister in the herbarium,
glabrous to very finely pruinate (under a lens) smooth, equal, hollow-tubulose, 6-18 X
0.2-0.3 mm; basal mycelium radiating,white, cottony-tomentose and sometimes some-
what ascendant on the stipe; white rhizomorphsoften present, tomentose. Context
white, very thin, inodorous.
178 Flora Neotropica

Spores 6-9.2(-11) X 2.5-4(-4.8)U, narrowlyfusoid to ellipsoid-oblong,or ellipsoid,

smooth, hyaline, inamyloid. Hymenium:basidia 15-18 X 5.8-7 u, 4-spored;basidioles
fusoid; cystidia none; cheilocystidia like epicuticularbroom cells. Hyphae of the trama of
the pileus hyaline, with clamp connections, pseudoamyloid, filamentous, but in places
swollen, especially near the forks, thin-walled:those of the stipe parallel, with pale
golden to cinnamon wall in the pigmented part and there thick-walled,hyaline in the
core and there thick-or thin-walled,2-7 p broad, pseudoamyloid. Corticallayers: epic-
utis of pileus hymeniform or subhymeniform,sometimes slightly irregular,broom cells
of the Siccus type, few transient to the Rotalis type, main body 10-24 X 5-14 p, hy-
aline, vesiculose, clavate or ventricose, sometimes some forked or lobed, thin-walled,
with setulae 1.3-5 X 0.7-1 u, erect or oblique or curved or forked, but mostly simple,
hyaline or some golden melleous, or pale ochraceous to golden; covering of the stipe
consisting of two types of elements, (1) pilose nodose, remindingone of the elements
found in Marasmiellussection Rameales, often forked, about 3 p broad, hyaline and
thin-walled,(2) broom cells like those of the epicutis of the pileus, but these scattered
and often relatively long.
On small stems and leaves of Monocotyledones and Dicotyledones, sometimes on
Gramineae. Cuba and Mexico.
MATERIALSTUDIED. CUBA. Wright(K, FH), type. MEXICO.Tabasco:Villahermosa,
Parquede la Venta, 1 Aug 1969, SingerM 8783 (F); MA
8785 (F).

134. MarasmiusmicrohaedinusSinger, Sydowia 18: 338. 1965.

TYPE. Singer B 25 78, from Bolivia.
Pileus hyaline, hygrophanous,dry-freshwhite, dried pale ochraceousbuff, glabrous,
smooth, obtuse, convex, then becoming concave, 6-11 mm broad. Lamellaewhite, close,
narrow, with some lamellulaeintermixed, not intervenoseor rarely(when stipe eccentri-
cally attached) somewhat forked-anastomosing,subfree. Stipe chestnut with white apex.
macroscopicallyglabrousto very finely subpruinate(under a lens), equal to very slightly
taperingupwards,26-28 X 0.7-1.3 mm; basal mycelium consistingof a strigosity which
radiatesfrom the base over the substratum,the appressedfibers becoming light fulvous
when dried, the others remainingwhite. Context thin, white, inodorous.
Spores 7-9 X 2.8-4 , ellipsoid to ellipsoid-oblongor fusoid, hyaline, smooth, in-
amyloid. Hymenium:basidia 21-22 X 7.2-7.5 p, clavate, hyaline, 4-spored:basidioles
clavate-fusiform(broadest above but acute or with narrow, small mucro); cystidia none;
cheilocystidia like the broom cells of the epicutis of the pileus but often smaller. Hy-
phae of pileus and hymenophoraltrama hyaline, with clamp connections, stroniglypseu-
doamyloid. Cortical layers: epicutis of pileus consisting of broom cells, hymeniform,
broom cells of the Siccus type, 16-20 X 8-11 p, setulae 3.5-4.2 X 1 p, subacute to acute,
both main body and setulae completely hyaline.
On dead stems of shrubsor herbaceousplants mixed with leaves (of a dicotyled-
onous tree) fallen to the ground in rain forest and on dead, rotting branchesof Legum-
MATERIALSTUDIED. COLOMBIA.Valle: Rfo Pance nearCali, I May 1968, SingerB
6512 (F). BOLIVIA. Vaca Diez, Riberalta.8 Apr 1956, SingerB 2578 (LIL), type.
Singer (1965) pl 15, fig 81

135. Marasmiustrinitatis Dennis, Trans. Brit. Mycol. Soc. 34: 425. 1951.
Marasmius 179

135a. Marasmiustrinitatis var trinitatis

TYPE. Dennzis158, from Trinidad.
Pileus olive fuscous e g 15-E-12 to "partridge"to fuliginous("elk" M&P)or deep
olive, ("willow" M&P),often with tawny to rusty center or intermediatezone (e g "rust,
sorolla br.") or with deep brown-redmargin,when very young often entirely "caldera"
(brown red), but later becoming golden cinnamon to ochraceousbrown (13-B-11 to "Arab")
with or without a sordid olive brown ("oak") center, dried mixed "cocoa" and "Arab"
or unicolorous one of these shades, or between "tarragona"and "Mohawk"(generally
losing the olive tinge on drying), glabrous,at first smooth, at maturity becoming long
sulcate-striate,center sometimes venose-rugose,but more frequently smooth, obtuse
or subumbonate,or somewhat to distinctly depressedin the center, campanulate-convex,
then convex-subapplanate,9-34 mm broad.Lamellaeyoung often cinerousor sordid
white, soon white, buffish white or pale flesh cream, with the edges concolorous with
sides, close to subdistant,narrow,eventually sometimes becoming broad in the larger
caps (up to 4-5 mm broad), rounded adnexed to subfree or free, not intervenose,more
rarelymoderately strongly intervenosein older specimens. Spore print pure white.
Stipe at first whitish with pale fulvous base, soon becoming chestnut with white apex,
glabrous,smooth, shining, equal or thickened towards the base, hollow, 27-50 X 1-2
nmm;basal mycelium strigose, white or whitish or pale fulvous, dried white, or pallid
with an occasional fiber of fibrils turned light fulvous, or entriely light fulvous. Con-
text thin, inodorous.
Spores 8.3-12.5 X 2.7-4,, fusoid-oblong,oblong, or sometimes subcylindric,
often applanateto subconcaveon inner, always convex on outer side, hyaline, smooth,
inamyloid. Hymenium: basidia 24-25 X 5-5.5 p, hyaline, clavate, 4-spored;cystidia
none; cheilocystidia present, like epicuticularelements but entirely hyaline. Hyphae
very broad and irregularin the trama of the pileus, some thickwalled,with clamp con-
nections, pseudoamyloid. Cortical layers: epicutis of pileus hymeniform, consisting
of broom cells of the Siccus-type, hyaline with golden melleous setulae, setulae often
very long (but not so in all collections).
On rotten sticks and leaves, bark, fruits and trunks of monocotyledonous and
dicotyledonous trees in rain forest and its margin,usually gregarious,sometimes den-
sely gregarious.
MATERIALSTUDIED. U.S.A. Florida: HighlandsCounty, HighlandsHammockState
Park,30 Mar1942 SingerF 449 (F). MEXICO.Veracruz,7 km S of Montepio,EstacionBio-
logica de los Tuxtlas 21 Jul 1969, SingerM 8104 (F). TRINIDAD. Non-Pareil,SangreGrande,
1 Oct 1949, Deinnis158 (K), type. BRAZIL. Guapore,Guajaramirim, 10 Mar1956, SingerB
1821 (LIL). BOLIVIA. Pando: Manuripi,Conquista,25 Mar 1956, SingerB 2225 (LIL); 24 Mar
1956, B 2166 (LIL), (?).
Singer (1965) pl 15, fig 86 T.

135b.Marasmiustrinitatis var immarginatusSinger, Sydowia 18: 344. 1965.

TYPE. Singer B 1883, from Bolivia.
Pileus, lamellae, and spore print as in var trinitatis. Stipe ochraceousbrown to
chestnut brown with white apex, at first very finely pruinate but glabrescent,other-
wise like type variety. Spores as in type variety, sometimes sausage-shaped.Hymen-
ium as in type variety but without any cheilocystidia! Hyphae as in type variety.
Coveringlayers: epicutis as in type variety; surface of stipe beset with broom cells
which rise from a reduced (small) main body (e g 7 X 4.8 c,), setulae divergent,spin-
180 Flora Neotropica

ulose and acute, many as long as main body or slightly longer, others very long and
reaching 18 , in length.
On bark and rotten wood of dicotyledonous hosts in virgin rain forest.
MATERIAL STUDIED. COLOMBIA. Valle: Buenaventura, San Joaquin. 19 Apr 1968,
Singer B 6253 (F). BOLIVIA. Beni: Vaca Diez, 3 km below Guayaramerin, 12 Mar 1956, Singer
B 1883 (LIL), type: B 1893 (LIL); 22 km east of Riberalta, 7 Apr 1956, Singer B 2554 (LIL),
probably here.
ILLUSTRATION.Singer (1965), pl 15, fig 86. 1.

136. MarasmiusolivascenticepsSinger, sp nov

Marasinius leoninuzs var olivascenticeps Singer, Sydowia 12: 105. 1958 "ad interim."
TYPE. Singer F 596, from Florida, U.S.A.
Pileo roseolo-brunneoad marginem,fuligineo-olivaceoin centro, sulcato, usque
ad 20 mm lato; lamellis albidis, olivaceo-marginatis;stipite castaneo, mycelio basali
fulvo vel brunneolo. Sporis 8.5 X 3.5-4.5 ;, cystidiis nullis. Ad folia delapsa nec non
ad lignum putridum in Florida. Typus in F conservatusest.
Pileus'fresh pinkish brown on the marginbut fuliginous-olivein the center, dried
fulvous-brownishwith an olive tinge in the center, with sulcate margin,glabrous,smooth
in the center, convex, about 20 mm broad or slightly less. Lamellaewhitish with olive
colored (discolorous) edge, moderately broad, moderately distant or subdistant. free to
adnexed. Stipe chestnut, chestnut-blackbelow when old, much paler towards the apex,
the apex itself usually white when fresh, smooth, slightly white pruinose above the
base and the apex often seemingly thinly pruinose under a lens, otherwise glabrous.40-
50 X 1.5-2.5 mm; basal mycelium tomentose or strigose, fulvous or brownish. Context
white, rather thin excepting the center of the pileus.
Spores 8.5 X 3.5-4.5 A;fusoid, smooth, hyaline, inamyloid. Hymeniurn:basidia
4-spored;cystidia none; cheilocystidia like the epicuticularbroom cells, also occurring
somewhat away from the edge proper, 20-33 X 5.7-6.5 , or spreadingabove. hyaline,
the setulae brownish melleous, somewhat coarse or long-conical,3.5-5.7 X 1.5/ . Hy-
phae of the pileus trama hyaline, mostly thin-walled,with clamp connections, in the
regularhymenophoral tramaall thin-walled. Cortical layers: epicutis of the pileus hy-
meniform, consisting of broom cells, appearingmottled when seen in scalp preparation
because some cells are either melleous hyaline, or golden melleous to brownishmelleous
because of the pigment of the setulae: main body 25-28 X 5-7.8 u, setulae 4.3-7.8 X
Both on foliage and rotten wood, Florida.
MATERIAL STUDIED. U.S.A. Florida, Highlands Haimmock State Park. 5 Sept 1942.
Singer F 596 (F), type.
The discolorousedge and the medium sized sporescharacterizethis species among
those with olive tinges (no. 138-141). None of the olive species is truly cystidiate in our
neotropical collections. In Africa, however, a species, 11. elaeocephalusSinger. with olive
pileus and distinct cystidia occurs.

137. Marasmiusepelaeus Singer, Sydowia 18: 335. 1965.

TYPE. Singer B 3047, from Bolivia.
Pileus olive brown to metallic olive ("syrup" to "bronze lustre" M&P)glabrous.
smooth or little sulcate, not venose-rugoseeither, convex with a shallow umbilicus, 17-
25 mm broad. Lamellaewhite to buffish white, subdistant,moderately broad, free,
sometimes adnexed. Spore print white. Stipe chestnut with white apex, equal, hollow,
Marasmius 181

25-54 X 1.5-2 mm; basal mycelium white. Context inodorous, extremely thin in pil-
Spores 7.5-8.2 X 3.5 p, ellipsoid-oblong,hyaline, smooth. Hymenium: basidia4-
spored;cystidia none: cheilocystidia like epicuticularbroom cells, but all hyaline. Hy-
phae of pileus and hymenophoraltrama strongly pseudoamyloid. Cortical layers: epic-
utis of pileus hymeniform, consisting of broom cells of the Siccus-type, main body
hyaline, rarely concolorous with the setulae, 10-14 X 7p, setulae golden yellow, 4.8-
5.5 X 1 p.
On dead (mainly dicotyledonous) leaves in rain forest.
MATERIAL STUDIED.BOLIVIA.Beni:VacaDiez,ArroyoFlorida,22 kmwestof Rib-
eralta,7 Apr 1956, SingerB 3047 (LIL), type.
This is intermediatebetween M. trinitatis var trinitatis (from which it differs in
the smallerspores and subdistantlamellae) and M. digilioi (which differs in its smaller
carpophoresand closer lamellae).
ILLUSTRATION. Singer (1965) pl 15, fig 87.

138. Marasmiusdigilioi Singer in Singer & Digilio, Lilloa 25: 201. 1952.
TYPE. Digilio & Singer T 488, from Argentina.
Pileus olive gray, olive brown, olivaceous-spadiceous("buffalo" "olive brown,"
14-J-6, 15-J-7/8, 15-L-8/9 M&P)sometimes with "cattail" or "biskra"in center), gla-
brous, smooth to distantly furrowed-subsmooth,eventually often becoming long-sulcate,
otherwise smooth, or more rarely rugulose in the center, often undulate, with a slight
depressionor/and a papilla in the center, otherwise convex, but eventually applanate
to irregular,5-16 mm broad. Lamellae(olive-brown)whitish or grayishpallid when
young, later between pure white and "rubber,"with the edges concolorouswith sides or
more rarely with pileus, subclose to medium close, narrow to broad, ventricose or not,
rounded-freeto rounded adnexed, Spore print pure white. Stipe rusty ochraceousbrown
to deep chestnut below and graduallypaler above with white apex ("Java," "alamo,"
"chutney"), glabrous, smooth, shining, filiform hollow, 15-40 X 0.4-1.5 mm; basal
mycelium tomentose or strigose, fresh usually white, dried often pale fulvous. Context
thin, white in pileus, inodorous.
Spores 6-9 X (3-)3.5-4.5(-5),, ellipsoid with suprahilardepression,hyaline, smooth.
Hymenium: basidia 20-28 X 6.8-9,u, clavate, 4-spored;basidioles fusoid; cystidia none or
cystidiole-like near edge; cheilocystidia either all like the epicuticularbroom cells or dif-
fering in being all hyaline, or else these broom cells intermixed with ampullaceousto ven-
tricose elements which have sometimes a slight knoblike swelling at the tip, at times
branchedbut not setulae-bearing,20-27.5 X 5.8-6.5 p, i e cystidiole-likeand in this
form also occasionally occurringnear the edge. Hyphae with clamp connections, pseu-
doamyloid. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells of
the Siccus-type, usually mottled in scalp view (hyaline-melleous)broom cells 14-30 X
5.8-11.7 u, main body hyaline, cylindric, clavate or vesiculous setulae erect, acute, some-
times relatively long, most melleous to bright golden melleous but on some cells all
hyaline e g 5-8 p long.
On rotting wood, more rarely on dead foliage mixed with other detritus in mon-
tane and subtropicalmontane, also subxerophyticpremontaneand tropical montane
MATERIAL STUDIED. BOLIVIA. La Paz: Nor-Yungas, Coroico, 29 Jan 1956, Singer B
668 (LIL). ARGENTINA. Tucuman, Sierra de San Javier, Anta Muerta, 24 Apr 1949, Digilio &
Singer T 488 (LIL), type; Parque Aconquija, Singer & Digilio T 278 (LIL); Tafi del Valle, Que-
182 FloraNeotropica

braditadel Mastil,15 Jan1950,SingerT 881 (LIL);Tapia,15 Feb 1951,SingerT 1488 (LIL);

Taficillo, 23 Feb 1955, Singer T 2218 (LIL).

139. Marasmiusater Singer, sp nov

TYPE.Dumont et al VE 5980, from Venezuela.
Pileo atro, parvo;lamellis griseis, ad aciem fuligineo-punctatis,confertis; stipite
subatro, apicem versus pallidiore. Sporis 6.5-7.5 X 2.8-3.7 p; cystidiis ad latera lam-
ellarumnullis; cheilocystidiis et elementis epicuticularibusfrequenterpigmento intra-
parietalifuligineo instructis, setulis atrosepiaceisvel fuligineisapicaliterappendiculatis.
Ad lignumiin Venezuela. Typus in NY conservatusest.
Pileus when dried black, indistinctly sulcate, sometimes somewhat rugulose, 3-4.5
mm broad when dried. Lamellaegray with fuliginous punctations along the edge, close,
moderately broad, not intervenose, subfree. Stipe blackish with paler apex, glabrous,
somewhat shiny, dried 14-25 X 0.2-0.3 mm; basal mycelium pale tawny-fulvous. Con-
text very thin.
Spores 6.5-7.5 X 2.8-3.7,u, oblong to subfusoid, smooth, hyaline, inamyloid. Hy-
menium: basidia 16-25 X 6-6.5 ,, 4-spored;basidioles fusoid; cystidia none on the
sides of the lamellae, but a few indistinct ones in some sections seen among the cheil-
ocystidia: cheilocystidia numerous but often intermittent, like the epicuticularbroom
cells, but most of them slightly narrower. Hyphae of the trama of the pileus hyaline,
thin-walled,with clamp connections, pseudoamyloid. Cortical layer: epicutis of the
pileus hymeniform, consisting of broom cells, main body sometimes hyaline, but in
the majority of the broom cells with a fuliginous intraparietalpigment, 8-15 X 4.5-9
AI,with firm but not thick wall, with dark sepia to fuliginous setulae, these 3.5-7-(9.
5) p long and rod-shapedto conical.
On decorticated, dicotyledonous wood.
MATERIALSTUDIED. VENEZUELA. Dto Federal,El Avila, 24 Jul 1972. Dumont et
al VE 5980 (NY), type.

140. Marasmiusruber Singer, Sydowia 18: 342. 1965.

TYPE. Singer B 1638, from Bolivia.
Pileus of one of the following orange or red colors or mixed: "tile red," "pi-
rate," "mandarinered," "burnt orange," "cadmiumorange," "sunkiss,""deep chrome
yellow," "Ponce de Leon," "Indian orange," "orangepeel," " Cadmiumyellow," 2-A-
12 M&Pthe general sequence being from golden yellow or orange to red as the spec-
imen matures and becomes less humid, at first smooth for a long time, then sulcate,
sometimes strongly and long sulcate and then tending to be striped red and orange,
or deep orange and light orange, or orange and golden yellow, glabrous,obtuse, or
slightly papillate, sometimes with depressedor umbilicate center, otherwise campan-
ulate, then regularlyto irregularlyconvex and eventually often applanate, 11-50 mm
broad. Lamellae "pirate" or "apricot yellow" or "Egyptian red" near edges, sides
concolorous with edges or "cadmiumorange"or 10-L-9, generallyat first more
orange and on drying or maturingmore red, narrow,tridymous or even tetradymous,
not intervenoseor with interspacesslightly rugose to slightly intervenose,appearing
close where there are many lamellulaeand distant or subdistant behind (about 20
through-lamellaein a medium sized specimen of 17 mm diameter),varying from
attenuate-freeto adnate. Spore print white. Stipe yellowish cream to flesh color
all cream when quite young, soon becoming some deep red brown ("Java," "Mara-
caibo") below and gradually(through "gold pheasant") paler upwards,at apex re-
Marasmius 183

maining "sunburst",11-E-7, glabrousto subglabrous,not pilose except at base,

equal or slightly taperingupwards,sometimes apex again thickened, 15-48 X 0.8-2
mm; basal mycelium abundant,strigose,varying from whitish to light fulvous when
fresh and often mixed "raw sienna" and pallid "Inca gold" and white. Context
thin, odorless.
Spores 8.2-9.7 X 2.8-4 , fusoid to oblong, thin-walled,hyaline, smooth, in-
amyloid. Hymenium: basidia 17.5-19.5 X 4.8-7.5 A, 4-spored;cystidia none; cheilo-
cystidia hyaline, in shape and structurelike the epicuticularbroom cells. Hyphae
of pileus trama and hymenophorehyaline but becoming pale orangeall over from
a diffusing pigment, thin-walled,sometimes many thick-walledones intermixed,
filamentousbut at times very irregularin shape at places, with clamp connections,
pseudoamyloid. Corticallayers: epicutis of pileus hymeniform, consisting of broom
cells of the Siccus-type, the whole epicuticularlayer developinga cadmium yellow
pigment soluble in NH4OHand KOH preparationsand diffusing and disappearing
eventually by pressure,main body of broom cells cadmiumyellow or hyaline, mostly
hyaline, e g 16-17 X 7 1, setulae hyaline or more often yellowish to golden yellow,
e g 3.5-5 j1long and 1-1.3 1diameterat base, spinulose and acute or rod-shapedand
obtuse, scalp preparationwith a mottled appearance,
On rotting dicotyledonous sticks, leaves and wood, and on dead monocotyledonous
leaves, in rain forest.
MATERIAL STUDIED. TRINIDAD. Arena Forest, 30 Oct 1949, Dennis 246 A (K); 2 Aug
1947, R. E. D. Baker, det. Dennis (as M. haematocephalus), (K). BOLIVIA. Beni: Vaca Diez, Guay-
aramerfn, 7 Mar 1956, Singer B 1638 (LIL), type; 8 Mar 1956, B 1712 (LIL), paratype; 14 Mar 1956,
B 1982 (LIL), paratype; 16 Mar 1956, B 2027 (LIL), paratype; 3 km below Guayaramerfn in virgin
forest, 12 Mar 1956, Singer B 1868 (LIL), paratype; Riberalta, 1 Apr 1956, Singer B 2413, (LIL),
Singer(1965) pi 15, fig 22.

141. Marasmiusbellus Berkeley, Hook. Jour. Bot 8: 139. 1856.

TYPE. Spruce, from Brazil.
Pileus cream color, then yellow ("sunriseyellow", 9-G-4 10-F-6 M&P),perhaps
slightly hygrophanousat times, paler between striae and sometimes pallid in elevated
parts, dried sometimes "honey yellow" (Ridgway) accordingto Dennis, in our specimens
varyingfrom 12-F-9 to as brownish as 13-I-10 or a mixture of "cognac" and "topaz,"
glabrous,varying from short to long and moderately to deeply sulcate, center smooth
to rugulose,campanulatethen convex, eventually applanate,with or without a small
umbo, 13-35 mm broad. Lamellaewhite to subconcolorouswith pileus (e g lamellar
groundpaler than the pileus, lamellarsides pale cream 9-C-2), subdistantto distant,
at times ratherstrongly intervenosebut mostly not intervenose,simple or with some
forked ones intermixed, narrow to medium broad, up to 3.5 mm broad, sinuate to
rounded and adnexed to adnate to a widened fertile apex of the stipe. Stipe dark
chestnut brown below, at least in age, otherwise lighter brown and with the apex con-
colorous with the lamellae, later brown or chestnut, glabrous,smooth, 35-54 X 1-1.2
(-4) mm; basal mycelium fresh, varyingfrom white to light fulvous (10-H-6) dried
generallyfulvous or partly fulvous (mixed with white fibers), often accompanied
by white to fulvous white byssuslikemycelium covering the surface of the substrat-
um. Context thin, white or whitish-pallidin pileus, with a weak odor or more
often odorless.
Spores 8-12.7 X 3-4.81, oblong or fusoid, often with applanateinner side, smooth,
hyaline, inamyloid. Hymenium:basidia clavate;basidiolesfusoid, or clavate;cystidia
184 FloraNeotropica

few and not clearly differentiated(cystidioles?), or entirely absent; cheilocystidia us-

ually confined to the extreme edges, of the lamellae, rarely scattered ones found near
the edges, like the epicuticularbroom cells in all regardsexcept that they have at
times smallersetulae than the latter. Hyphae and often lower portion of main body
of epicuticularelements pseudoamyloid,those of pileus and stipe very strongly pseu-
doamyloid. Corticallayers: epicutis of pileus hymeniform, consisting of broom cells
of the Siccus-type, 13-32 X 5.5-9 ,, ventricose to vesiculose, generallythin-walled,
with apical and often slightly divergingsetulae, the latter spinulose, ratherthin, sub-
acute, the entire broom cells appearinghyaline or subhyalinewhen seen individually
but in recently collected relatively well pigmentedmaterialthey are yellowish to
chrome yellow when seen in accumulations.
On forest and plantation litter of all kind, such as grass,herbaceousstems, mono-
cotyledonous and dicotyledonous leaves, sometimes attached to small pieces of wood,
in coastal montane tropical forest, Amazonasrain forest, etc.
MATERIALSTUDIED. BRAZIL. Amazonas:"Panure"(=Ipanure=Sao Jeronimo),Spruce
(K) type; Pernambuco:Recife, Dois Irmaos,SeverinoJose da Silva, 10 Apr 1955, comm. Chaves
Batista2224 (LIL). BOLIVIA. Beni: Vaca Diez, virginforest 3 km below Guayaramerin,12 Mar
1956, SingerB 1898 (LIL);La Paz: Nor-Yungas,Coroico, 25 Jan 1956, SingerB 669 (LIL).
Singer(1965), pl 15, fig 84; (1958a) fig 21; see also Dennis
(1961) fig 43.

142. Marasmiusbeniensis Singer, Sydowia 18: 334. 1965.

TYPE. Singer B 2000, from Bolivia.
Pileus ochraceous-alutaceous(11-G-7), dried dull brown (14-F-11) sulcate-rugose
to the disc which is only slightly rugose and umbilicate, glabrous,flat when mature,
24 mm but much smallerwhen dried. Lamellaewhite, edge indistinctly concolorous
with the pileus, subdistant to subclose (not close or crowded) intermixedwith lame-
llulae which are often crisp or forked, otherwise not intervenose,adnate. Stipe "Al-
amo" at base and graduallyless deeply rust-ochraceousbrown towards the apex which,
itself, is white, glabrous,about 40 X 1 mm; basal mycelium white, abundant,dried
mixed pale fulvous and white. Context inodorous.
Spores 7-8.2 X 2.7-3 , not larger. Hymenium:basidia 15-20 X 4.8 p, 4-spored;
cystidia none; cheilocystidialike epicuticularbroom cells. Hyphae of pileus very
strongly interwovenand variablein shape and thickness of the wall, pseudoamyloid.
Corticallayers: epicutis of the pileus hymeniform,consisting of broom cells of the
Siccus-type(none of them smooth!) main body e g 14 X 7 u, vesiculose to clavate,
golden brownish to golden melleous, not very deeply so but consistently so and the
scalp preparationnot showing hyaline and colored zones (not mottled!), setulae con-
colorous with main body, 5.5-7 , long, thin and acute or spinulose and obtuse; cover-
ing of stipe almost absent, consisting of inconspicuous,minute, smooth, conic excres-
cencies which are very scattered and reach 10-17 X 4-5.5 (at base)l, neither hairs
nor setae or broom cells present.
On small sticks in rain forest.
MATERIALSTUDIED. BOLIVIA. Beni: Guayaramerfn, 14 Mar 1956, SingerB 2000
(LIL), type. PANAMA. GatumLake, 10 Aug 1940, Lorentz 3035 (BPI).
The latter collection is somewhat doubtful since the spores are slightly larger
but the type specimen may not have been fully mature.
ILLUSTRATION: Singer(1965), pi 15, fig 83.
6Holotype, the largercarpophoreof this collection.
Marasmius 185

143. Marasmiusbezerrae Singer, sp nov Fig 65.

143a. Marasmiusbezerraevar bezerrae

TYPE.SingerB 3409, from Brazil.
Pileo dilute brunneo vel ferrugineo,sulcato, 7-17 mm lato; lamellis albis vel cremeis
aurantio-vel brunneo-marginatis,confertis vel subdistantibus;stipite 15-50 X 0.6-1 mm.
Sporis 9-11 X 3.2-5.2 1; cystidiis veris nullis. Ad frustula arborumdicotyledonearumde-
lapsa in silva tropicali Brasiliae. Typus in F conservatusest.
Pileus light brown to ferruginous(e g 12 J 9) and "rust, sorolla br" or ferrugin-
ous (M&P)on margin,reaching"Maracaibo"in the center, dried between "rust, sorolla
br" and "cocoa" or near "caldera"(M&P),strongly sulcate in the marginalportion, con-
vex, obtuse or broadly umbonate, sometimes more applanatein age, 7-17 mm broad.
Lamellaepallid to cream, with a broadly and distinctly discolorous(orange to concolor-
ous with some part of the pileus) edge, medium close to subdistant,narrow to moder-
ately broad, not intervenoseor in age slightly intervenose,often some forked, adnate,
more rarely rounded-adnate. Stipe deep cinnamon brown or chestnut or deep chestnut
brown, graduallymore pallid at the apex at first, equal, perfectly glabrous,hollow, 15-50
X 0.6-1 mm; basal mycelium moderately abundantbut distinct, strigose, pale dirty
fulvous at least when dried. Context of pileus white, thin, inodorous.
Spores (8-)9-11 X 3.2-5.2 ,, most frequently around 4 , broad, ellipsoid to fusoid-
oblong, smooth, inamyloid. Hymenium: basidia 16-27 X 4.8-10 ,, 4-spored, perhaps
sometimes some 1-3-sporedones intermixed;basidiolesfusoid; cystidia none; rarelya
few cystidioid bodies little differentiatedfrom the basidiolespresentat the gill edge;
cheilocystidia like the epicuticularbroom cells. Hyphae of pileus hyaline, all firm- or
thin- to thick-walled,moderately strongly but distinctly pseudoamyloid,not dimorphic,
radiallyarranged,interwoven, with clamp connections. Corticallayers: epicutis of the
pileus hymeniform, consisting of broom cells, with the main body 9-15.5 X 5.5-10.5 u,
hyaline to pale orange or golden orange or light melleous, thin-walled,setulae 3-7.3 X
0.4-22, obtuse, spinulose, deep brownish melleous, golden melleous or orange, in other
cells subhyaline, the scalp view of the epicutis therefore appearingmottled. Surface
of the stipe showing a few scattered inconspicuousversiformhairs but no broom cell
dermatocystidiaor setuloid excrescencies.
On dicotyledonous sticks fallen to the ground in tropical forest.
MATERIALSTUDIED. BRAZIL. Pernambuco,Caramagibe,14 Jul 1960, Singer& Bezerra,
SingerB 3409 (F), type.

143b.Marasmiusbezerraevar chitepecensisSinger,var nov Fig 66.

TYPE. Guzmzbnz 2800 B, from Mexico.
Varietasfoliicola lamellis confertis et pileo sublevi insignis.
Differs from the type variety only in close lamellae and not or slightly sulcate
marginand growth on dicotyledonous leaves.
MATERIALSTUDIED. MEXICO.Chitepec, 30 Oct 1960, Guzman2800 B (F) type.
The relatively broad spores and the strongly discolorousedge of the lamellae
are characteristic.

143c. Marasmiusbezerraevar griseoferrugineus(Singer) Singer, comb nov

leoninusvargriseoferrugineusSinger,Sydowia 18: 337. 1965.
TYPE. Singer B 1919, from Bolivia.
Differs from the precedingvarieties in gray lamellae which are subclose but appear
subdistantwhen dried, and grayishpallid apex of stipe.
186 Flora Neotropica

On dead dicotyledonous leaves.

MATERIALSTUDIED. BOLIVIA. Beni, Vaca Diez, 3 km below Guayaramerini
in virgin
tropicalrainforest,12 Mar1956,SingerB 1919 (LIL),type.
A complete description of this variety has been given by Singer (1965, p 277 as
M. leoninus var griseoferrugineus).

144. Marasmiusnapoensis Singer, sp nov Fig 67.

TYPE. Singer B 7348, from Ecuador.
Pileo brunneo, longe profundequesulcato, usque ad 60 mm lato; lamellis albidis,
aciebus vix discoloribus,distantibus;stipite castameo, 60-77 X 2 mm. Sporis 7.5-11
X 3-6 ,, ellipsoideis;cystidiis nullis. Ad folia dicotyledonea ligno putrido immixta in
Aequatoria. Typus in F conservatusest.
Pileus "cocoa" with "Seminole" (M&P)marginor "chutney" (M&P)i e brown, with
the center fresh deeper and rustier than "Mandalay"(M&P),dried a very deep rusty
brown with the center almost blackish-purplish-ferruginous, long and deeply sulcate on
the margin,the center smooth to venose, glabrous,convex, obtuse, with depressedcen-
ter when mature, up to 60 mm broad. Lamellaewhitish, then (in situ) discolored sor-
did brownish with the edges not discolorous(excepting in dry condition slightly brown
near stipe and/or edge), distant, scarcely to distinctly intervenose,broad, attenuate-
subfree or rounded-subfreeto adnexed. Stipe deep chestnut below, at first with whit-
ish apex, macroscopicallyglabrousbut under a lens finely pruinateat apex, subequal,
hollow, 60-77 X 2 mm; basal mycelium abundant, ocher-whitish. Context of pileus
thin, white, inodorous.
Spores 7.5-11 X 3-6 u, (Q = 1.7-2.7) characteristicallyellipsoid, some oblong,
ovoid or broadly fusoid, mostly with a slight suprahilardepression,smooth, hyaline,
inamyloid. Hymenium: basidia 26-32 X 7-7.5 p, 4-spored, basidioles fusoid; cystidia
none; cheilocystidia extremely versiform, mostly in form of broom cells which are
entirely pale brownish or with hyaline main body and hyaline to pale golden yellow
setulae, some (few) without setulae. Hyphae of the pileus tramahyaline, with clamp
connections, with thin wall, pseudoamyloid. Cortical layers: epicutis of the pileus
hymeniform, consisting of broom cells, these with a main body 8-20 X 4-12,, versi-
form, ochraceous to brown, fewer hyaline or subhyaline, with thin or thick wall: set-
ulae 4.7-11 X 1-1.5 u, acute, all erect or somewhat spreadingbut all apical, deep
succineous brown or ochraceousbrown; covering of the stipe (pruina) consisting of
broom cells which are like the cheilocystidia or have a more reduced main body and
hyaline to pale golden setulae up to 21 u long, often forked.
On dead dicotyledonous leaves intermixed with some rotten pieces of wood.
MATERIAL STUDIED. ECUADOR. Napo, Lago Agrio, 17 May 1973. Singer B 7519 (F).
type; Sacha 4 (Norte), 10 May 1973, Singer F 7348 (F).

145. Marasmiusleoninus Berkeley, Hook. Jour. 8: 135. 1856.

Androsaceus orinocensis Patouillard & Gaillard, Bull. Soc. Mycol. France 4: 21, 1888.
Marasmius orinocensis (Patouillard & Gaillard) Saccardo, Sylloge Fungorum 9: 68. 1891.

145a. Marasmiusleoninus var leoninus

TYPE. Spruce 112, from Brazil
Pileus cinnamon, cinnamon fulvous, light to rich orange, fulvous-orange,orange-
ferrugineous,almost brick-red,orange-chestnut,orange ochraceous(10-H-8, 11-F/I-10,
12-C-11, 13-B-ll, "talavera,""Saratoga"to "cornhusk,""spice," "cinnamon,"
Marasmius 187

"gold pheasant," "feuille morte," "autumn glory," "coromandel"M&P(but rustier)

marginoften somewhat paler (e g "Titian," "ferruginous,""Punjab"),dried either still
deeper colored ("caldera,""mohawk,""Java,""casserole,""gypsy," "auburn"M&P)or
in parts as yellow as "Yucatan," following patterns establishedby subhygrophanous
zones at times present in the fresh pileus, glabroussulcate to deeply furrowed-sulcate
(not merely short transparentlystriate), leavinga small to large disc either smooth or
more rarely weakly rugulose-verruculose, convex, then convex with rathernarrowly
depressedcenter, often with recurved-applanate to even uplifting marginat maturity,
or, on the contrary, with very declivous marginand an applanatezone around an um-
bilicus or an umbilicate small umbo, eventually mostly entirely subapplanateto con-
cave, 15-42 mm broad. Lamellaepure white, cream white, or light cream, with the
edges concolorous with sides, or at least not conspicuously discolorous, narrowto
broad (and if broad 3-4.5 mm broad), subclose to distant, mostly moderately close
to subdistant, not or very slightly intervenose, free, subfree, rounded-adnexed,adnex-
ed to adnate to a widened apex of stipe (very variablein gill-attachment),intermixed
with lamellulaeand sometimes regularlytridymous with entire edge average 12-16
through lamellae. Spore print pure white. Stipe fulvous to very deep chestnut or
ochraceousbrown, apex at first white pallid or concolorous with the lamellae, later
concolorous with base or merely somewhat lighter colored (Aesculus-seed-color,or
brownishochraceous),glabrousor subglabrous,smooth, shining to opaque equal or
taperingupwards,hollow, 38-78 X 0.8-3 mm; basal mycelium tomentose or/and
strigose, fulvous, pale orange fulvous ("gold leaf," "melon glow," "daffoldil"):fulvous
white, or pure white, often mixed white and fulvous, sometimes white and becoming
somewhat fulvous on drying, whitish rhizomorphsoften observed. Context mostly
relatively fleshy in center, thin in margin,tough in stipe, unchanging,taste mild;
odor like that of Collybia dryophila or practicallyabsent.
Spores 7-12.3 X 2-4.7 p, the most common length measurementsrangingfrom
8.7-10.7 to 10.5-11 P, fusoid to oblong-ellipsoid,thin-walled,inamyloid, hyaline,
smooth. Hymenium: basidia 15.5-29 X 4.3-7.5 p, (2-?) 4-spored, hyaline; cystidia,
none, or cystidiole like and not differentiatedfrom basidiaand basidiolesinasmuch
as the latter vary from transparentto subopaque-refringent; cheilocystidiapresent or
absent, present like the epicuticularbroom cells. Hyphae either all equal and thin-
walled, filamentous, strongly pseudoamyloid, with clamp connections, or similarbut
of two kinds, some thick-walledand even resemblingat times a Crinipellis-hairwhen
seen in the Melzer'sreagent,very variablein size and shape, the others less strongly
pseudoamlyoid,thin-walledand more regularlyfilamentous, not to very slightly gel-
atinized where the trama(of the pileus) is dimorphic,hyphae of stipe thick-walled,
all strictly parallel. Corticallayers: epicutis of the pileus hymeniform,consisting
of broom cells of the Siccus-type,main body mostly hyaline or all hyaline, in some
collections a minority of main bodies concolorous with the pigmented setulae, these
often moderately thick-walled,otherwise thin-walled,subcylindric,ventricose, vesi-
culose-pedicellateor clavate, 15-25 X 5.5-9(-18.5 if spreading-broadened or ramified)
,: setulae 3.5-9.5(-18) X 0.7-2 (base!) u acute, subacute, or obtuse, sometimes all
cells with golden hyaline to hyaline setulae (and main bodies) and sometimes more
non-pigmentedthan pigmented broom cells present, but more frequently most cells
with melleous to succineous-melleoussetulae, and a minority with hyaline ones, more
rarelysome chestnut-brownones also present;smooth cells generallyvery few or none;
coveringlayer of stipe either entirely absent, or else consisting of scattered broom cells,
often with reducedmain bodies and elongated setulae.
188 Flora Neotropica

On woody substratum(both monocotyledonous and dicotyledonous plants),

but also mixtures of small twigs, rotting leaves and other trash, even sawdust, palm
fruits, etc., solitary or in dense groups, but not cespitose.
MATERIAL STUDIED. U.S.A. Florida: Highland Co., Highland Hammock State Park. 4
Sep 1942, Singer F 570 (F); Dade Co.: Simpson Park, Miami, 14 Sep 1942, Singer F 693 (F.FH).
VENEZUELA. Amazonas: Atures, Jul 1887, Gaillard 116 (FH), type of M. oritnocensis; Sucre'
NW of Irapa, 11 Jul 1972, Duniont et al VE 4676 (NY). ECUADOR. Napo, Shushufindi, 13
May 1973, Singer B 7416 (F). BRAZIL. Amazonas: "Panure" (Ipanure=Sao Jer6nimo), Mar
1853, - Spruce 112 (K) type of M. leoninus; Rio Grande do Sul: Sao Leopoldo. 1933, Rick (as
M. fulviceps) (FH). BOLIVIA. Beni: Vaca Diez, Guayaramerin, 8 Mar 1956, Singer B 1718, B
1722, B 1722a, (LIL); 14 Mar 1956, B 1990 (LIL); 11 Mar 1956, B 1830/1(LIL); 3 km. below
Guayaramerin in virgin forest, 12 Mar 1956, Singer B 1888, B 1896 (LIL); La Paz: Nor-Yungas.
Coroico, 1800 m alt, 15 Feb 1956, Singer B 1168 (LIL). ARGENTINA. Misiones: Posadas, 5
Apr 1953, Martinez 89 (LIL); Corrientes: Saladas, 29 Apr 1950, Pedro Goretta 92 (LIL).
This species, of common occurence in many regions between South Florida and
Northeast Argentina,is variablein severalrespects (1) in the pruinosity of the stipe,
(2) structureof the trama of the pileus, (3) the presenceor absence of cheilocystidia.
Of the specimens enumeratedfor this species above, most had the stipe annotated
as glabrous and, indeed, under the binocularit appearedstill glabrous,and under the
microscope no broom cells were discoveredaway from the apex. However, in B 1 722,
1722a, 1830, we have three collections, all from the Bolivian Amazonas region (Oriente)
where the stipe is pruinose to subpruinoseand scattered broom cells can be found. The
colors of the three collections are similarbut not identical, the lamellaevary in density
from specimen to specimen of the same collection. Since we cannot find any correlated
characterwe do not think this form to be worth of a varietal name at the present
In Singer (1958a), I attempted to separate,describingthem separately,a form
with dimorphic and a form with homogeneous pileus trama. The form with the dim-
orphic trama was our Florida collection F 693. I thought at the time that this char-
acter was perhapscorrelatedwith white basal mycelium and presence of cheilocystidia
but I have found all transitionsfrom white to fulvous basal mycelium, although in
other collections of the dinorphic form (e g B 1896, 1718 from the "Oriente"),1
found cheilocystidia constantly present. There are undoubtedly transitionalcases be-
tween dimorphicand homogeneous trama(as in B 1 718) which I interpretas an in-
dication of incomplete maturity, but the homogeneous forms are often providedwith
numerouscheilocystidia (B 1831 and F 570). It seems that the dimorphic-cheilocyst-
idiate form is also representedby the type of M. orinocensis and one of the Argentine
specimens. In the former we also find very slightly pruinose stipe (s. 1.) which may
mean that the cheilocystidiate and dimorphicform is also the one with broom cells
on the stipe, but this latter correlation we have not been able to confirm in all cases.
If such a correlation exists and a name is needed, it would be M. orinocensis which
ought to supply the epithet desired. In the type of A. leotninuscheilocystidia "en
brosse" are present and the trama is not or not distinctly dimorphic. This is also the
case in a collection from the Yungas(B 1168) which has the same basal mycelium and
cheilocystidia as the type of M. leoninus, even seems to grow on the same substratum
(dicotyledonous leaves, here of Melanostomaceae);the trama is likewise not distinctly
This means that among the collections which show cheilocystidia, not all have
dimorphictrama and those that do not, either have colored basal mycelium and grow
on leaves (apparently the most typical M. leoiinus) or have white mycelium and grow
on palm (M. aff. orinocensis Sing. 1958a).
Marasmius 189

The absence of cheilocystidia is not always easy to demonstratesince in some

specimensthe edge of the lamellae has been damaged. I find cheilocystidia of the
broom cell type in the great majority of all collections; where they are absent, as in
the Argentine collection from Corrientes,I find dimorphictrama and glabrousstipe.
It is uncertain whether this correlationholds any better than others, but I have
studied materialwhere the cheilocystidia are scanty which may be interpretedas a
transitionalform (B 1722); here the stipe is somewhat pruinose. The typical cheilo-
cystidiate form was described by Singer (1958a, p 107, 108).
Consequently, I am not certain whether the degree of pruinosity and the pres-
ence or absence of cheilocystidia are useful charactersfor the infraspecifictaxonomy
of the Marasmius-leoniiius-complex but as far as the hyphal structureof the trama is
concerned, I have thus far found a constant correlationbetween monomorphic(thin-
to firm-walledhyphae only) tramalhyphae and the substratumon fallen leaves on one
hand and dimorphic(many thick-walledand a variablenumber of thin-walledhyphae)
hyphal structurewith lignicolous habit. Thus I divided the species tentatively into two
Var leoninlus. Hyphal structureof the pileus trama monomorphic,thick-walled
liyphae here exceptional or absent. Foliicolous.
Var orinocensis (Patouillard& Gaillard)Singer ad interim. Many strictly thick-
walled hyphae (in adult specimens) in the pileus-tramapresent. Lignicolous.
ILLUSTRATION. Singer(1965) pl 15, fig 90.
The following variety is obviously very close to both the precedingbut differs
in the fact that this correlationis absent (here both thin- and thick-walledhyphae in
the pileus-tramaand habitat on dicotyledonous fallen leaves as well as on wood); also
the spores are shorter than usual in Marasmiusleoninus and the edge of the lamellae
is tending to become somewhat discolorouswhen dried:

145b.Marasmiusleoninus var aberransSinger, var nov Fig 68.

TYPE. Singer M 8188, from Mexico.
Hyphis et crassi-et tenuitunicatis in tramatepilei, sporis brevioribus,acie lam-
ellarum subdiscoloriin siccis nec non habitatione ad folia dicotyledonea delapsadif-
fert. Typus varietatisin F conservatusest.
Pileus dull orange, (ll-E-10 to 12 E 12) dried 11 I 9 (M&P)with paler, even
pallescent marginalzone, glabrous,coarsely deeply rugose-sulcateup to near the center,
convex, often somewhat umbonate or slightly umbilicate when dried, 10-30 mm broad.
Lamellaewhitish, fresh with concolorous (whitish) edge, but on drying edge variably
either concolorous or partly to almost entirely pale orangy, slightly ventricose, not in-
tervenose, subdistant, subfree to adnate. Stipe deep chestnut below with white apex
or chestnut above, blackish below, the apex at first always whitish, glabrous,smooth,
subequal, 25-45 X 0.8-1 mm; basal mycelium abundant,strigose, fresh whitish, on
drying tending to become fulvous at least in parts. Context of the pileus white, in-
Spores 7-9.5 X 3.5-4.5,, ellipsoid, fewer oblong, without suprahilardepressions,
smooth, hyaline, inamyloid. Hymenium: basidia 19-26 X 6-7 u, 4-spored;basidioles
fusoid; cystidia none; cheilocystidia like the epicuticularbroom cells but relativelymore
of them, sometimes all, hyaline. Hyphae of the pileus-tramawith thick (reaching 1 ,)
to mostly thin to firm wall, not gelatinized, with clamp connections, pseudoamyloid.
Corticallayer: epicutis of the pileus hymeniform, consisting of broom cells, with hy-
aline wall but most filled with a dissolved intracellularpale orangy pigment, clavate,
190 Flora Neotropica

cylindricalor ventricose, 9-19 X 4-12 g, setulae 2-16 X 0.7-2 u, conical, acute or sub-
acute, palest golden to golden hyaline or quite hyaline, or with a more conspicuous in-
traparietalmelleous golden pigment.
On dicotyledonous leaves fallen to the ground as well as on nearby dicotyledonous
wood, gregarious.
MATERIAL STUDIED. MEXICO. Veracruz, 3 km west of Fortin de las Flores, 24 Jan 1969,
Singer M 8188 (F), type. ECUADOR. Napo, Shushufindi, 13 May 1973, Singer B 7414 (F).
This variety approachesM1.iiapoesis because of its broader spores but differs in
havingthe colors of M. leoninus, glabrousstipe and non-interwovenlamellae.

146. Marasmiusfloriceps Berkeley & Curtis, Jour. Linn. Soc. Bot. 10: 298. 1869.
Fig 69.
TYPE. Wright31, from Cuba.
Pileus bright orange-rusty,bright orange or bright red-brown,sometimes orange
with some inconspicuousbrown radial stripes and/or orange brown or orange yellow
marginalband (e g "terracotta"with "Nasturtium"center or "toltec" with center 11
C 10 to 12 B 10 or "doubloon" with "Punjab"center or "golden wheat" with "rose
amber" center, dried e g 3 A 12, the orangebrown becoming "copper br" M&P),
glabrous,sulcate excepting the disc, rarely short-sulcatewhen mature with, more rarely
without rugose or venose center, campanulateto convex or subhemispherical,later
often with either more declivous or upturned margin,the center often papillate but
also with umbilicate center, 9-20 mm broad. Lamellaewhite to orange-cream,subclose
to subdistant, the edges not discolorous or only near margin pale orange when dried,
narrowto ratherbroad, at first somewhat ascendant, then horizontal or almost so,
free to rounded-adnexed,intermixed with many lamellulae. Stipe brown to deep chest-
nut with white to pale cream apex at least at first, glabrous,equal or subequal, hollow,
23-58 X 0.5-1 mm; basal mycelium scanty to more often abundantand strigose. white
or fulvous to tawny when fresh and dried. Context of the pileus white, very thin,
Spores(8-)8.5-13 X (2.7-)3-4.2(-4.8),, fusoid to fusoid-oblongbut some some-
times relatively broad and often curved in lower portion if narrow, smooth, hyaline, in-
amyloid. Hymenium: basidia 16-24 X 6-8 ,, 4-spored, basidioles fusoid; cystidia none
except for a few inconspicuoussubulate and hyaline ones at the gill edges among the
cheilocystidia which are broom cells like the epicuticularbroom cells but usually all
hyaline or with golden hyaline or orange-hyalinesetulae. Hyphae of the trama of the
pileus hyaline with firm but not distinctly thickened wall although often slightly swell-
ing in alkali but not dimorphic, filamentous, not gelatinized, with clamp connections,
but often many secondarysepta present, pseudoamyloid,those of the stipe parallel,+
thickwalled, deep yellow. Subhymeniumcellular. Hymenophoraltrama regular,con-
sisting of filamentous but sometimes relatively broad, thin-walledelements. Cortical
layers: epicutis of the pileus hymeniform, consisting of broom cells, their main body
9-13 X 8-11 u, hyaline to ochraceous, scalp preparationsappearingmottled because the
setulae are either hyaline or golden to deep golden melleous, acute to subacute, (2-)3-10
M,long; covering of the stipe none, except for scarce, minute excrescencieslooking like
setulae without main body.
On fallen dicotyledonous leaves, stems and twigs of Leguminosae,rotten wood
of all kinds, but also on small monocotyledonous stems.
MATERIAL STUDIED. CUBA. Wright 31 (K, FH), type. MEXICO. Veracruz: Estaci6n
Biologica de las Tuxtlas, 7 km S of Montepfo, 21 Jun 1969, Singer M 8116 (F); 29 Jul 1969,
Marasmius 191

SingerM 8741 (F); 20 Jun 1969, SingerM 8088 (F). COLOMBIA.Valle, MunicipalidadCali,
Rio Prance,1 May 1968, SingerB 6511 (F); BuenaVentura,Calima,18 Apr 1968, SingerB 6206
This species is often difficult to distinguishfrom M. berteroi var berteroi and
I. corrugatusvar auranltiacusbut the charactersemphasizedin the key should make
it possible to come to an identification in all cases if the distributionof the three
species is taken into considerationand the cortical layers of M. corrugatusare duly
ILLUSTRATION. Dennis (1951b), fig 7 (type).

147. MarasmiuscuatrecasasiiSinger, sp nov Fig 70.

TYPE. Singer B 6517, from Colombia.
Pileo dilute ochraceo vel cinnamomeo-brunneoloin centro, pallescentein mar-
gine hygrophano,longe sulcato, 20-31 mm lato; lamellis pallidis, satis angustis, subcon-
fertis vel subdistantibus;stipite cinnamomeovel castaneo, sub lente subtiliter pruino-
so. Sporis 6.2-9 X 2.5-3.7 p; cystidiis nullis vel admodum inconspicuispaucisque. Ad
lignum in Columbia. Typus in F conservatusest. Species de scientia botanica colom-
biana meritissimocl. J. Cuatrecasasdedicata.
Pileus light ochraceousbrown or cinnamon brownish in the center, bleachingto
light cinnamon brownish or pallid on the margin,hygrophanous,often partly white
when dry, but dried "topaz" to "hazel" (M&P),distinctly long (almost to the center)
sulcate to sulcate-furrowed,glabrous,convex, 20-31 mm broad. Lamellaepallid,
rathernarrow, subclose or in mature specimensmostly subdistant,narrowlyadnexed
to rounded-adnateor adnate. Stipe dull cinnamon to dark chestnut or dark cinnamon
spotted at the apex, macroscopicallyglabrousappearingsubvelutinouswhen fresh,
very finely and densely pruinate when seen under a lens in dried condition, hollow,
equal or subequal, 22-23 X 1-2 mm; basal mycelium finely and slightly tomentose or
strigose, mostly rather scanty. Context white, thin, odor none or slight (of HCN).
Spores 6.2-9 X 2.5-3.7 p,, oblong-fusoid, smooth, hyaline, inamyloid. Hymenium:
basidia 16-24 X 5-7 ,, 4-spored;cystidia none or very inconspicuousand few (as in
AM.helvolus); cheilocystidia forming a band of sterile basidioles(cystidioles, 20-21 X
4-5 , varying from clavate to vesiculose, mucronate or more rarely obtuse, hyaline,
inamyloid); besides hyaline broom cells, many or all without setulae. Hyphae very
strongly interwoven and variablein the thickness of the wall as well as the shape of
the hyphal cells but basically thin-walledand radiallyarrangedin the pileus, subpara-
llel in the stipe, strongly pseudoamyloidwith clamp connections. Corticallayers:
epicutis of pileus hymeniform, broom cells of the Siccus type, main body 10-15 X 6-10
$,, often hand-shapedor subcylindricalto clavate, hyaline, many with and many with-
out setulae, these, where present, rarely hyaline, mostly melleous-ochraceous,obtuse,
4-11 X 1-1.7 1, the broom cells alternatingsingly or in groups with the smooth cells;
surface of the stipe densely beset with broom cells like those of the epicutis of the
pileus, some of these smooth (without setulae).
On wood. Colombia.
MATERIALSTUDIED. COLOMBIA.Valle: Buenaventura,Calima,"Pulpapel,"20 Apr
1968, SingerB 6258 (F); Rio Pance nearCali, 1 May 1968, SingerB 6517 (F), type.

148. Marasmiuscorrugatus(Patouillard)Saccardo& Sydow, Sylloge Fungorum16:

AndrosaceuiscorrugatusPatouillard,Bull. Soc. Mycol. 16: 175. 1900.
192 Flora Neotropica

148a. Marasmiuscorrugatusvar corrugatus

TYPE. Duss, from Guadeloupe.
Pileus pinkish orange to reddishcinnamon, (e g 10-C-10M&Pto "Saratoga,"
dried "caramel"mixed with "Alamo" M&P,a narrowdisc often with a tan or "burnt
umber" shade in both fresh and dried state but this discolorationoften missing and,
instead, center sometimes more reddishorange than otherwise, smooth to finely striate
on margin, otherwise more or less strongly venose or radiallyrugose, glabrous,convex,
often with depressedcenter in age, 10-47 mm broad. Lamellaewhite to whitish, nar-
row, crowded or close, rarely medium close, not intervenose,adnexed, some almost
subfree, others almost adnate, in the herbariumusually darkeningto red-brown(be-
tween "cocoa" and "Arabianbrown"). Stipe in primordiumwhite, then fulvous to
chestnut with orange ochraceous upper portion, smooth and glabrous,opaque to
shining, terete or in largerspecimens often canaliculate-double,equal, hollow, 20-33 X
1-2.5 mm, basal mycelium tomentose or woolly-strigose,brownish or white or pale
fulvous when dried. Context ratherthin, white, unchanging,inodorous.
Spores 7.5-11 X 3-4.5 ,. Hymenium: basidia 20 X 5.5 p, 4-spored, some 2-
spored in some preparations,clavate, hyaline; basidioles fusoid; cystidia none or
cystidiole-like (and doubtful); cheilocystidia like epicuticularbroom cells but all
hyaline or stramineous-hyalineor with few cells with colored setulae, the latter often
somewhat divergent-spreading,sometimes few to none, not making the edge hetero-
morphous but mixed with basidiaand basidioles at least near the stipe. Hyphae of
pileus and hymenophoral tramahyaline, thin-walled,sometimes some thick-walledbut
often very broad, in stipe parallel,all hyphae strongly pseudoamyloidand with clamp
connections. Cortical layers: epicutis of pileus hymeniform, consisting of broom cells,
the latter mixed with smooth cells (which are like broom cells consisting entirely of
a main body, no setulae), main body about 12.5 , long, either entirely hyaline or
partly melleous, very rarely entirely melleous, thin-walled,rarely with moderately
thickened wall, clavate or vesiculose, setulae rather irregularin shape and size, 2-8.5
X 1.3-2 p (at base) but not unfrequently some strongly elongated and then reaching
up to 22 X 3.5 j,, these setulae varying form golden yellow melleous to orange ful-
vous, spinulose, rod-shapedor even fusoid, sometimes forked, mostly simple, acute
to obtuse, scalp preparationsshow mottled surface because of even distributionamong
the colored setulae and smooth cells of some broom cells with completely hyaline
setulae; in age the percentageof the smooth epicuticularcells grows in relation to the
broom cells but not conspicuously so, and no conspicuous color change of the pileus
has been correlated to the phenomenon. Surface of the stipe without broom cells but
in some specimens some minute conic excrescences emergingfrom the parallelsurface
hyphae of the cortical layer; these excrescencesinconspicuous,small and scattered,
brownish, occasionally fasciculate and appearingto belong to a broom cell with reduced
main body; also some scattered dermatocystidia(e g clavate, 30 X 6.5 ,, thin-walled)
mostly present.
On decayed wood but also on mixtures of vegetable debris (herbaceous stems,
branchlets,rotten leaves of Gramineae,Solanum, Schinus, Pteridophyta)in tropical
rain forest and coastal forest, in montane forests of various types, usually gregarious
in small groups, not cespitose.
re, NWof Irapa,7 Jul 1972, Dumont et al VE 4194 (NY). BOLIVIA. Pando: Manuripi,Conquista,
24 Mar 1956, Singer B 21 74 (LIL); Beni: Vaca Diez, Guayaramerin,. 12 Mar 1956, Singer B 1899
(LIL) ARGENTINA. Catamarca:Andalgala. Above west bank of Rfo Chacras("Las Estancias")
11 Feb 1952, 1500m alt,Singer T 1822 (LIL).
Marasmius 193

The Argentine collection T 1822 both geographicallyand in habitat widely

separatedfrom the type may not be simply synonymous; I found neither hairs nor
broom cells on the stipe. This collection is only temporarilyinsertedhere.
ILLUSTRATION.Singer (1965), pl 15, fig 89.

148b.Marasmiuscorrugatusvar aurantiacus(Murrill)Singer, comb nov

Marasmiutsaurantiacus(Murrill)Singer,Sydowia 18: 268. 1965.
aurantiacusMurrill,Bull. TorreyClub 66: 157. 1939.
TYPE. Murrill,from Florida (U.S.A.).
Pileus orange (4 A 12 M&P)then becoming more ferruginous("copper br" M&P,
"Burnt Sienna" Ridgway), subhygrophanousand bleachingin spots or areas on drying
and in age, center at times more reddish orange or on the contrary, duller (tan) dried,
often as deep as orange-chestnutor ochraceousbrown, glabrous,smooth or slightly fine-
ly striate at maturity or in dried caps, but in many specimensmore or less distinctly
corrugated(radiate-venose-rugose)in the center, conic or campanulate-convex,then
applanateand often with uplifted marginand center gibbous, 8-22 mm broad. Lamellae
white or whitish to cream with frequently orange ground, close or crowded to medium
close at maturity, subvenose, then narrow(3 mm) to medium broad, adnexed, adnate
or subdecurrent,not intervenoseor very slightly so, about 22 or more through-lamellae.
Stipe at first white with ochraceous-pallidor pale orange ochraceousbase, later concol-
orous with the pileus with white apex, becoming ferruginousbrown, often canaliculate,
otherwise smooth, glabrousor subglabrousopaque and when seen under a lens the
young surface is often covered by a thin, scattered pruinosity, glabrescent,equal or very
slightly taperingupwards,hollow, 20-45 X 1-2.5(-3.5) mm; basal mycelium strigose,
white, more rarely partially fulvous or entirely orange-fulvous. Context thin, white,
inodorous or with a slight odor of sauerkraut.
Spores(6.3-)6.8-11 X 3-3.7 ,, in the typical form mostly 6.8-10 long, hyaline,
smooth, inamyloid, oblong. Hymenium: basidia4-spored;basidioles fusoid; cystidia
none: cheilocystidia like epicuticularbroom cells but all hyaline to pale melleous and
with normally subacute setulae on all cells (smooth cells on edges, few); occasionally
(southermost collections) most cheilocystidia with golden setulae as in the epicuticular
broom cells. Hyphae in trama of pileus, hymenophore and stipe hyaline, strongly pseu-
doamyloid, in trama of stipe and basal mycelium frequently thick-walled,all hyphae
with clamp connections. Corticallayers: epicutis of pileus hymeniform, containing
broom cells of the Siccus-type, main body 7-31.5 X 5.5-12.8 g, cylindric to clavate,
sometimes subvesiculose,sometimes forked, hyaline to colored (the pigmented ones
fewer and often very scattered, almost regularlydistributed)in the latter case usually
pale melleous below and deeper melleous brown above and then wall often thick, more
rarely pigmented main bodies absent; setulae 3-12.8, rarely up to 22 Allong,at base
0.5-1.5(-4),, characteristicallyvariablein length and also varying from acute and spin-
ulose to rodshapedand obtuse, golden, golden melleous or deep melleous to melleous
brown, sometimes intermixed with cells with subhyaline or hyaline setulae and with
cells without any setulae (but these generallya small minority). Coveringof the stipe
consisting of very scattered to rathernumerous broom cells similarto those of the
pileus but often narrowerwith long hyaline to golden melleous setulae, those often
divergentspreading,sometimes arisingalmost directly from the surfacehyphae of the
cortical layer of the stipe; entire cell (if main body not reduced) e g 20 X 8.7 g, spin-
ules e g 6.3-10.7 , long. Underlyingcortical hyphae strictly parallel. Setae, conical
excrescencies,etc absent on stipe and even broom cells tending to disappearin old
194 Flora Neotropica

specimens, but thin-walled,minute, hyaline excrescenciesof variableshape often accom-

panying the broom cells.
On dead wood, generally on very rotten wood of dicotyledonous trees often cov-
ered by leafmold or on rotting chips of wood in leafmold, solitary to densely gregarious,
almost cespitose; in warm-temperate,subtropical,tropical and tropical-montaneforest
in the northern hemisphereand in tropical forest in the southern hemisphere. Type
on Quercus.
MATERIALSTUDIED. U.S.A. Florida:AlachuaCounty, PlaneraHammock,16 Jul 1938,
lMuirrill(FLAS), type; HighlandsCounty, HighlandsHammockState Park,5 Sep 1942, SingerF 596
(F). COLOMBIA.Valle, Rio Pance, 1600-1800 m alt, SingerB 6945 (F). BRAZIL. Pernambuco:
Dois Irmaos,5 Jun 1960, SingerB 3072 (BAFC). ECUADOR. Napo, Lago Agrio, 8 May 1973,
SingerB 7303 (F). BOLIVIA. La Paz. Nor-Yungas,Coroico, 27 Jan 1956, SingerB 676 (LIL).
ARGENTINA. Misiones,Caminode San Ignacioa Posadas,km 105, R. T. Guerrero50, 23 Feb
1960 (LIL).
This variety is similarto lMarasmniuscorrugatusvar corrugatuswhich is usually
somewhat larger, with slightly broaderspores, not growingin cespitose groups and hav-
ing no broom cells with well developed main body on the surface of the stipe.
ILLUSTRATION. Singer (1965) pl 15, fig 88.

148c. Marasmiuscorrugatusvar lacustrisSinger, var nov Fig 71.

A var aurantiacocui ceterum simillima haec varietas differt pileo maiore, derma-
tocystidiis stipitis dimorphis. Typus in F conservatusest.
Pileus when quite young ochraceous brown or orange but soon, before maturity,
turningpartly hyaline or pale orange, hygrophanous,white or whitish to leather-brown-
ish when dry, glabrous,smooth or slightly sulcate to sulcate on margin, not scrobicul-
ate rugose on the disc but sometimes venose almost all over, convex to conic convex,
sometimes somewhat more repand with recurvedmargin,with or without a shallow
depressionor subumbonate, 25-57 mm broad. Lamellaewhite, sometimes with orangy
ground, edges white, close to crowded, narrowor very narrow,with entire edge, some-
times interruptedwith numerouslamellulae,eventually distinctly if not strongly inter-
venose, when young only rugose in the lamellae-ground,rounded-adnateto adnate or
subfree. Stipe tawny to chestnut below, at first white in upper half or third, partly
or completely glabrous,equal or slightly attenuated upwards,hollow, 50-100 X 1.2-2
mm; basal mycelium present, woolly, white or sordid pallid. Context white in pileus.
thin, inodorous.
Spores (6-)7.5(-10.5) X 2.8-3.5 ju,ellipsoid-oblongto fusoid, smooth, hyaline, in-
amyloid. Hymenium: basidia 18-22 X 5.5-7 ,, 4-spored:cystidia none: cheilocystidia
dimorphic, some in form of broom cells, entirely hyaline, few with golden setulae, 17-
27 X 7-24k, with setulae 2-7.5 X 0.5-1.2 u, acute; aside from the broom cells there are
two types of smooth cells: (1) like the epicuticularsmooth cells, 15-18 X 7-9.5 , (2)
basidiole-likebut sterile, e g 20-21 X 4-5 1. Hyphae of the pileus-tramaboth thin- and
thick-walled,hyaline, pseudoamyloid, with clamp connections. Corticallayer: epicutis
of pileus hymeniform, consisting of (1) broom cells with setulae and (2) broom-cell-like
cells but without setulae, the former with a hyaline, rarely pale golden melleous main
body 11-24 X 6-12 and setulae hyaline to stramineousor golden melleous, acute or ob-
tuse, 1.8-7 X 0.4-1.3 u, rarely some setulae up to 22 X 2 p, the latter 11-24 X 11-15 ,,
smooth, vesiculose, hyaline, in younger pilei scattered, in older ones often very numer-
ous, or else very numerous from the beginning;covering of the stipe consisting of very
scatteredminute dermatocystidia or hairs, these of two types (1) broom cells much
like those of the epicutis, (2) entire thin-walled,hyaline, erect cells which are 8-27 X
Marasmius 195

3-6.5,, ventricose, subcapitateor subcylindrical,obtuse; at the apex of stipe type 2

more numerous,at middle and below broom cells more numerous.
On woody dicotyledonous sticks and stumps of trees, but passing over onto
rotting leaves, gregariousbut not caespitose. Colombia to Ecuador.
MATERIALSTUDIED. ECUADOR. Napo, Lago Agrio, 7 May 1973, SingerB 7262 (F),
SingerB 7422 (F), 11 May 1973, B 7366 (F).

148d.Marasmius corrugatus var portonovensis Singer, var nov

Marasmniusportonovensis Singer, Sydowia 18: 272. 1965 nom subnud "ad interim."
TYPE. Rick 576, from Brazil.
A var corrugatodiffert stipite pro ratione magis elongato, pileo minus intense
colorato, cellulis penicillatis epicuticularibuset cheilocystidialbusminus numerosis;
hyphis tramatishymenophoralismanifeste dimorphicis,sporis paullum minoribus. Ty-
pus in FH conservatusest.
Pileus "Light PinkishCinnamon"and "PinkishCinnamon"(Ridgway), between
"hazel" and "cigarette"M&Pwhen dried, old fresh specimens flesh color (near "Peach
Blow," 1 -B-5 M&P),central portion or entire pileus radiate-rugose,slightly paler be-
tween rugositiesand marginfaintly sulcate in dried material,convex, dried 7-32 mm
broad. Lamellaewhite, brownish pallid with white edge when dried, narrow,close to
subdistant, intermixed with lamellulae,horizontal, adnexed to almost adnate. Stipe
chestnut brown, opaque to slightly shining but not pruinose, equal, hollow, 20-62 X
0.7-2.3 mm; basal mycelium strongly strigose, white to pale fulvous, extensive. Con-
text thin, white.
Spores 6.2-7.5 X 2.7-4 p, hyaline, smooth, ellipsoid to oblong, sometimes tilda-
shaped, inamyloid. Hymenium: basidiaclavate, 18-19 X 7 c,, 4-spored;basidioles fus-
oid; cystidia none; cheilocystidia dimorphic,most like the smooth, epicuticularelements,
fewer like the epicuticularbroom cells and entirely hyaline, some irregular,sometimes
forked, often nodulose or even diverticulatewhere the subhymenialelements form the
external layer of the edge, these hyaline, remindingone of the cheilocystidia of Maras-
miellzs. Hyphae: in stipe all hyphae brown and parallel,filamentous, in pileus and
hymenophoraltrama irregularin shape, not all filamentous, characteristicallydimor-
phic when mature, some thin-walledand moderately pseudoamyloid,others thick-
walled like hairs of Crinipellisand very strongly pseudoamyloid,clamp connections
present. Cortical layers: epicutis of pileus characteristicallyconsisting of dimorphic
elements, type I being clavate-vesiculoseand hyaline without setulae and only with
occasionalasperitiesor nodulose surface, 13.5-31 X 7-13 p, type II typical broom
cells of the Siccus-type (although setulae somewhat spreading),hyaline or in upper
portion slightly golden, as far as the main body is concerned, the latter of the same
size as the elements of type I, setulae golden hyaline to golden melleous, extremely
variablein length, (1-)3-8(-35) X 1-2(-3.5),, sometimes forked, generallyrod-shaped
and obtuse or subacute, some broom cells thick-walled;on the stipe no covering
elements except some inconstant and very scattered short, erect, thin-walled,versi-
form hyphal ends, broom cells on stipe absent.
On small branches and large stems in forest litter, also on rotten logs.
MATERIALSTUDIED. U.S.A. Florida:HighlandsHammockState Park,27 Aug 1942,
SingerF 415 (F). BRAZIL. SantaCatarina:Porto Novo, Rick 576 (FH), type. ARGENTINA.
Misiones:BernardoIrigoyen,27 Feb 1960, R. T. Guerrero102 (LIL);Iguazu,Refugio "Yaguar-
et6," 29 Feb 1960, R. T. Guerrero 114 (LIL).
This is thinner and more elongate, also apparently less intensely orange-chest-
nut than the type of M. corrugatus and differs from the latter likewise in the rela-
196 FloraNeotropica

tively small number of broom cells among the setula-lesscells in the epicutis and gill
edge, in the strongly dimorphic characterof the hyphae of the hymenophoraltrama
and in somewhat smallerspores. It differs from M1.aurantiacuswhich it resembles
in size, by the scarcity of broom cells on edges of the lamellae and surface of the
stipe and likewise by the absence of setulae on the majority of the epicuticularele-
ments of the pileus, and in the color of the pileus.
ILLUSTRATION. Singer (1958a) fig 25.
Stirps Cladophyllus
This stirps is intermediatebetween seriesLeonini and Actinopodes; it is charac-
terized by the type of stipe-covering,crowded to close and often anastomosinglamellae
and often glabrescentstipe; there are two subgroups,one with, one without distinct

149. MarasmiusaraucariaeSinger, Sydowia 18: 333. 1965.

TYPE.Singer M 1147, from Argentina.
Pileus castaneous-ferrugineous(so deep and rich that nothing comparablewas
found in Maerz& Paul) or ferruginousor paler flesh color in age (3-A-11), dried about
"rust, sorolla br." or somewhat deeper, glabrousto subvelutinous,slightly radiallyrugose
and not sulcate or striate at first, later becoming more or less sulcate on margin,convex
with umbonate center, more rarely obtuse or even with slightly depressedcenter, later
always convex with depressedcenter and eventually at timcs irregularlyflattened, 15-
30 mm broad. Lamellaepallid-whiteto pallid-cream,subclose to crowded, in age gen-
erally subclose, intermixed with numerous, sometimes crisp lamellulae,occasionally
some forked or with transverseribs on lamellae-sidesbut not distinctly intervenoseeven
in old specimens, narrow to moderately broad, free to rounded-adnexed. Stipe cinnamon
below, cream white above, macroscopicallyappearingglabrousbut under a lens white
pruinate, only in young specimens pruinosityvisible without the help of a lens. smooth,
equal or taperingupwards, 29-70 X 1.5-2.5 mm; basal mycelium fulvous-whitish. Con-
text ratherthin, inodorous.
Spores 9-12.5 X 2.7-4.5, , oblong, inner side almost applanate,hyaline, thin-walled,
inamyloid. Hymenium: basidia hyaline, clavate, (2-)4-spored, 24-26.5 X 6.5-7.5 ,; cystidia
none; cheilocystidia like epicuticularelements but mostly all hyaline or pale melleous.
Hyphae of trama of pileus and lamellae pseudoamyloid,with clamp coInnections, in age
up to 22 j, broad and walls up to 1.3 thick, hyaline. Corticallayers: epicutis of pileus
hymeniform, consisting of broom cells of the Siccus-type, main body 9-16 X 4.5-7.2 u,
hyaline, or in upper portion melleous or entirely golden melleous, clavate, setulae 3.5-7
X 1-1.7 /,, erect, golden melleous, acute or obtuse. Hairsof the stipe surface, e g 50 X
5.5 p, mostly smaller,versiform,usually subcylindricwith rounded tip, septate or not,
thin-walled,hyaline, numerous.
On very rotten wood and needles of Araucariabrasilienisisin mixed forests of the
subtropical-montanetype, perhapsalso on other substrata.
MATERIALSTUDIED. ARGENTINA. Misiones.Frontera,Gral.ManuelBelgranco,20 Apr
1957, Singer M 1147 (LIL) type; 19 Apr 1957, 1 1082 (LIL) paratype; Bernardo Irigoyen, R. T.
Guerrero 92 (LIL) paratype.
ILLUSTRATION:Singer (1965) pi 22, fig 116.

150. Marasmiuscladophyllus Berkeley, Hook. Jour. Bot 8: 138. 1856.

Marasmiiuspolyporoides Murrill, North American Flora 9: 266. 1915.
Favolaschia mnagnificaHennings, Hedwigia 36: 203. 1897.
Marasmius 197

Saccardo& Sydowin Saccardo,

Laschiamagnifica(Hennings) 14: 199. 1899.
magnificus Singer,Lloydia8: 218. 1945.

150a. Marasmiuscladophyllusvar cladophyllus

TYPE.Spruce 89, from Brazil.
Pileus orange fulvous with deep rusty fulvous center or orange ferruginousor a
similarshade (12-I-11 to 4-A/E-12, "feuille morte," "mosque," "terracotta,""burnt
sienna, M&P"between "zinc orange"(Ridgway) and "tawny" (Ridgway) to "amber
brown" (Ridgway) the center and its neighborhoodoften "Persimmon"M&P)rarely
brown (between "tawny" and "Pekinese"M&P),sometimes "Kis kilim" with "Hispano"
M&Pmargin(intermediatezone 12-D-ll), glabrous,smooth or slightly rugose or reticu-
late, generally not deeply radiallysulcate in the marginalzone unless with cross veins
at the same time, more rarely with rugose center and slightly sulcate margin,umbilicate
or with more broadly depressedcenter in age, campanulateto convex, eventually flat,
15-60 mm broad. Lamellaecream, sordid buff to buffish-pallid(9-G-6, 10-D-4, 11-C-
4, "blush," "mellow mauve"), often with the edges concolorous with the pileus, but
not in all carpophores,in some carpophoresthe whole edge third or half subconcolor-
ous with pileus but paler and the rest of the lamellae sordid (buffish-) pallid, mostly
strictly horizontal-linearwhen mature, but varyingfrom (mostly) narrowto (very
rarely) altnost broad, strongly forking and merulioid-anastomosingor at least distinct-
ly intervenose,often appearingporoid, intermixed but lamellulaeanastomosingwith
through-lamellaeand often not clearly differentiated,close to (more rarely) subdistant,
adnexed to adnate-subdecurrent.Spore print pure white. Stipe pale brownish with
brownish white apex or, especially later, becoming dark chestnut below and deep ferru-
ginous further upwards("tarragona"upwards,base "Rembrandt"to "caldera"or "Mar-
acaibo"), or entirely reddishbrown ("cocoa" to "trotteur tan"), often assuminga blue
gray shine from the pubescence which covers it from apex to base, at least when seen
under a lens but often glabrescentin age or pubescence becoming obsolete after hand-
ling, equal or taperingupwards,30-112 X 1-4 mm; basal mycelium coarsely tomentose-
strigose, mostly fulvous or sordid brown, etc ("Windortan," "Titian,""alamo"), some-
times with some white fibrils immixed, rarely entirely white. Context white, unchang-
ing, inodorous, rarely witli a slight odor of Collybia dryophila or sauerkraut.
Spores 7.8-12.7 X 2.3-3.8 p, most frequently around 9-10 X 3-4.5 A, ellipsoid-
oblong, hyaline, smooth, inamyloid. Hymenium: basidia 4-spored;cystidia none;cheil-
ocystidia like epicuticularbroom cells but some or more cells with hyaline setulae,
often not occupying the entire edge and replacedby hyphal ends and versiformcheil-
ocystidia like those in varglaberripesand var intermedius. Hyphae of hymenophoral
and pileus tramahyaline, smooth, in age some with thin, some with thick walls, pseu-
doamyloid, with clamp connections. Corticallayers: epicutis of pileus hymeniform,
consisting of broom cells of the Siccus-type,15-25(-32) u long, main body either hy-
aline or concolorous with the setulae, setulae either all uniformly orange tan or on
some cells hyaline to subhyaline, on others melleous to brownishmelleous and some-
times on others even deeper colored, in the latter case scalp preparationsgiving a
mottled appearance,setulae spinulose 3.3-12(-24) X 1.3-1.8(-3.6)ui. Pubescence of
stipe consisting of hyaline, thin-walledhairs, these versiformor contorted, often cy-
lindric or ventricose, with broadly rounded apex or forked at apex in a few cells,
sometimes even with wavy-nodulesoutline or diverticula,remindingone of a Ram-
eales-structure(Marasmiellus),septate or unicellular,35-50 X 4-9.8 A(6.8-9.8 Ain
198 FloraNeotropica

middle of stipe of mature specimen), often accompaniedby ventricose cells with or

without mucro or over some areas replaced by these.
On monocotyledonous and dicotyledonous (but apparentlyalways Angiospermae)
wood and leaves, fruits and sticks, solitary or in small groups, not cespitose, in tropical
rain forest and in subtropicalto tropical forests along the Gulf and Atlantic Coast but
not in tropical-montane(Yungas type) forests and not known from northwesternArgen-
tina. Known hosts: Leguminosae,Palmaeand Heliconia sp.
MATERIALSTUDIED. U.S.A. Florida:HighlandsCounty, HighlandsHammockState Park,
3 Sep 1942, SingerF 521 (F); 26 Aug 1942, F 406 (F). BELIZE:El Cayo, Valentin, 24 Jun 1936,
E. B. Mains3628 (MICH),probablytopotype of M. polyporoides;October 1906, Morton E. Peck
(NY) type of M. polyporoides. PANAMA. CanalZone: BarroColoradoIsland, 13 Jul 1952, Mar-
tin & Welden7673 (LA); Corundu,13 Aug 1952, Martin8360 (IA); Rio Sardinilla,10 Jul 1952,
Martin,Welden& Bouchl 7552 (IA). ECUADOR. Napo, Lago Agrio, 16 May 1973, SingerB 7490
(F). BRAZIL. Para,Belem, 10 Jun 1966, SingerB 4248 (BAFC);Amazonas:Rio Jurua,Bom Fim,
Nov 1900, Ule, HerbariumBrasiliense2699 (FH); Panure(Ipanure(Ipanure= Slo Jeronimo),Sprnce
89 (K) type; Rio Javary,Rematedos Males, 1927, P. Occhioni 15, RB 5, 2, 17 (LIL); Pernambuco:
Camaragibe,6 Jul 1960, SingerB 3109 (LIL); Rio de Janeiro:Angrados Reis. 30 Sep 1952, Singer
B 436 (F); Rio Grandedo Sul: Serro Azul 1928, Rick 422 (FH). BOLIVIA. Beni, Vaca Diez, Guay-
aramerin,6 Mar1956, SingerB 15979 (LIL), 8 Mar1956, B 1727 (LIL),B 1759 (LIL), 11 Mar1956,
B 1829 9LIL), 14 Mar 1956, B 1978 (LIL), 16 Mar1956, B 2016 (LIL), 3 km below Guayaramerin
in virginforest, 12 Mar1956, SingerB 1884 (LIL). ARGENTINA. Misiones:Arroyo PirayGuazli,
21 Feb 1960, R. T. Guerrero74 (LIL); BernardoIrigoyen 27 Feb 1960, R. T. Guerrero68 (LIL).
Singer(1965) p1 22, fig 117.

150b.Marasmiuscladophyllusvar glaberripesSinger, var nov

TYPE.Singer B 7160, from Ecuador.
Stipite glaberrimorecedit. Typus in F conservatusest.
This differs from the type variety in glabrous stipe, absence of broom cells on the
edge of the lamellae, slightly different color and largersize.
Pileus yellow brown to cinnamon ("Yucatan" to "caramel"M&P)often with much
paler("harvest" M&P)margin,with a deeper brown (between "Kis kilim" and "Sudan
br" M&P)central dot, smooth, glabrous,convex, then undulated-applanate,young often
somewhat umbonate, 37-64 mm broad. Lamellaeradiatingbut so densely anastomosing
that they appear pore-like, pallid with orange ground, later sometimes with partially
ferruginousspots, narrow(1 mm), the pores almost isodiametric,adnate. Stipe chestnut,
with pallid apex, glabrous(even under a lens), hollow, taperingupwards67-76 X 1.5-2
mm; basal mycelium strigose, abundant, pallid. Context of pileus white, thin, membran-
ous at margin,inodorous.
Spores (7.8-)8.5-9.8 X 2.5 ,, narrowlyoblong-fusoid, smooth, hyaline, inamyloid.
Hymenium: basidia 18 X 5.5 p, 4-spored;cystidia none; cheilocystidia consisting of pseu-
doamyloid hyphal ends which are conspicuously contorted and often forked, about 3.5 ,
broad, no broom cells. Hyphae of the pileus trama hyaline, thick-walled,few thin-walled,
strongly pseudoamyloid, with clamp connections. Cortical layers: epicutis of the pileus
hymeniform, consisting of broom cells only, these variegatedas in the type variety, the
main body 8-18 X 5-8.5 ,, mostly clavate, setulae acute, rather thin, 4-7 Mlong; covering
of stipe none.
On dead leaves and small sticks of dicotyledonous trees and shrubs,gregarious.
East slope of the Andes.
MATERIAL STUDIED. ECUADOR. Tungurahua, Rio Verde, S of Rfo Pastaza at 1600 m
alt, 28 Apr 197 3, Singer B 7160 (F), type.
Marasmius 199

150c. Marasmiuscladophyllusvar intermediusSinger, var nov

TYPE. Singer B 7343, from Ecuador.
Differt pileo minore stipiteque subglabro. Typus in F conservatusest.
This is intermediatebetween var cladophyllus and varglaberripes. The pileus is
smallerthan in var glaberripes,the stipe is subglabrousand externally beset by minute
hypha-likeexcrescences which are occasionally like hyaline setulae (but singly on a
strongly reduced or absent main body); only few broaderhairs like those of the type
variety are present. The lamellae are sordid white and often have a partially rusty edge
when seen in dried condition but the cheilocystidia are scattered, thin-walled,of two
types (a) fusoid and long beaked, 23-26 X 5-6 ,, hyaline, and (b) vesiculose, often muc-
ronate, hyaline or browned, 12-27 X 6-18 p, inamyloid;the hyphae are dimorphic, some
strongly pseudoamyloidand thick-walled,others inamyloid to vaguely pseudoamyloid
and thin-walled.
On rotting dicotyledonous wood and rotting fallen dicotyledonous leaves in hylaea.
MATERIAL STUDIED. ECUADOR. Napo, Sacha 4, 10 May 1973, Singer B 7343 (F), type;
Rio Coca, Francisco de Orcllana, 12 May 1973, Singer B 7379 (F).
I have not revised the entire materialcited for the type variety. It is possible that
some of these materialsbelong to either varglaberripesor var intermedius I have form-
erly inserted this species with the species of seriesActinopodes but the existence of the
varietiesdescribedhere suggests that its place is ratherin series Leonini.

151. Marasmiuspusio Berkeley & Curtis,Ann. Mag. Nat. Hist. 12: 427. 1853.
lMarasmius guatopoensis Dennis, Kew Bull 15: 95. 1961 (a variety).

151a. Marasmiuspusio var pusio

TYPE. Ravenel 816, from U.S.A.
Pileus orange, orange cinnamon, salmon orange, deep brownish orange, ochraceous
yellow, when fresh ("Moresco,""apricot," "Ponce de Leon," 10-D-9, 10-H-7), the young-
er, the deeper orange eventually fading to almost pallid in many collections, especially in
the marginalzone, dried about light melleous (ll-G/H-6), glabrous,smooth, eventually be-
coming weakly shallowly and coarsely sulcate and on drying often short-sulcateon margin,
not viscid, convex, not umbilicate but in some collections occasionallyumbonate, sooner
or later becoming applanate, 2-8 mm broad. Lamellaepure white to cream, all equal or
more commonly rather regularlydidymous or tridymous, rathernarrowto very broad,
arcuate to horizontal, never ascendant, close to subdistant(the smallerthe carpophore
the more distant appearingthe lamellae and through-lamellaerelativelynot very numer-
ous: 8-12), free to adnate to a fleshy disc, edges concolorous with sides. Stipe entirely
white when quite young, later becoming "Windsortan" with white or cream to light
yellowish (concolorous with lamellae) apex, and eventually slightly browner than pileus
to umber or almost chestnut all over, extreme apex of stipe sometimes discoid, con-
colorous with lamellae or pileus, macroscopicallyseeminglyglabrouswhen fresh but
under a strong lens finely pruinate although soon or eventually glabrescent,the pilosity
often becoming denser towards the base where it merges with the basal mycelium, us-
ually central, more rarely somewhat eccentric, equal, curved or geniculate, 4-26 X 0.1-
0.5 mm, basal mycelium abundantalthough often indistinct when wet, strigose or to-
mentose and accompaniedby thin white rhizomorphsin some collections, pure white
or partly orange-pallid,dried white or partly fulvous-white. Context white, extremely
thin, inodorous.
200 FloraNeotropica

Spores 6-10.3 X 2.5-4.5(-5.5) p, very variable,most frequently around 7.5-9.5 X

3-4 p, oblong-ellipsoidto oblong cylindrical, at times with a spur-likebulge on inner
side near hilar appendageor with a bulge on the outer side like Canpanella alba, rarely
curved, generally with a slight suprahilardepression,thin-walled,hyaline, smooth, in-
amyloid. Hymenium:basidia 18-20 X 6-6.5 p, clavate, hyaline, 4-spored;basidioles
fusoid or fusoid-mucronate;cystidia none, or in some collections present but cystidiole-
like, not differentiated from basidiaand basidioles and only exceptionally tending to be
slightly broaderand with firmer wall than basidia, fusoid to clavate, e g 30-40 X 6.5-8 p;
cheilocystidia like the epicuticularbroom cells, at places crowded, at places more scat-
tered. Hyphae of tramaof pileus and hymenophore thin-walledand thin-filamentous,
hyaline, varyingfrom moderately strongly but distinctly to very strongly pseudoamyloid,
all with clamp connections, hyphae of stipe parallelwith each other, always strongly
pseudoamyloid. Corticallayers: epicutis of pileus subhymeniformor hymeniform, con-
sisting of broom cells of the Siccus-type, main body 6-35 X 5.5-11 1, hyaline or yellow-
ish-hyaline,simple or branchingRanaria-like, often pedicellate, clavate, vesiculose cylin-
dric-ventricoseor mushroom-shaped,pedicel 4-4.5 u thick, setulae either likewise all hy-
aline to yellowish hyaline or cells intermixed with some with thicker walls and with or
without setulae and then these setulae deep golden and with other cells with thin walls
and golden yellow setulae or only with the latter type of cells, setulae 5.5(-19) X 0.5-1.3
(-2.5) p, mostly simple but also often branchingonce or twice, mostly subacute or
acute but also often obtuse, spinulose or rod-shaped;pruinosity of stipe consisting of
numerous to scattered broom cells with ventricose main body which is often reduced
and then the cells seemingly consisting of a few fascicles of setulae, these setulae at
times taking the shape of setae (when they are solitary) and then hyaline to subhyaline,
needle sharp to obtuse, simple or branching,1.3-2.5 p thick, at times accompaniedby
diverticulatesuperficialhyphae, more numeroustoward the base, very scattered at apex.
On dead fallen twigs, bark, leaf petioles and on rotten wood of dicotyledonous
trees and shrubs.
MATERIAL STUDIED. U.S.A. South Carolina: Santee Canal, Ravenel 816 (FH) type.
ECUADOR. Napo: Lago Agrio, 12 May 1973, Singer B 7363 (F). BOLIVIA. Beni: Vaca Diez,
Arroyo Florida 22 km east of Riberalta, 7 Apr 1956, Singer B 2566 (LIL); Pando: Manuripi, Con-
quista, 24 Mar 1956, Singer B 2172 (LIL).
ILLUSTRATION.Singer (1965) pl 16, fig 92.

151b.Marasmiuspusio var guatopoensis (Dennis) Singer, comb nov

MarasmiusguatopoensisDennis, Kew Bull. 15: 95. 1961.
TYPE.Dennis 1103, from Venezuela.
Differs from the type variety in broaderspores (7.3-11 X 3.3-4.8 u); on fern rachis
and other Pteridophytaremnantsbut sometimes passingover onto dicotyledonous stems
and sticks, usually growing in rows. The stipe is generallyshort (not or not much longer
than the diameter of the pileus which reaches up to 5 mm in diameter).
MATERIAL STUDIED. VENEZUELA. Miranda: Guatopo, 25 Jun 1953, Deinnis 1103 (K).
ARGENTINA. Tucuman, Yerba Buena, 12 Feb 1955, Singer T 2155 (LIL).
ILLUSTRATION. Dennis (1961) fig 46.

151c. Marasmiuspusio var poliobasis Singer, Sydowia 18: 343. 1965.

TYPE. Singer B 722, from Bolivia.
Differs from the type variety in the color of the base of the stipe which shows a
caesious tint or the basal mycelium is partly or entirely gray. The variety has been fully
described in Sydowia 18: 342.
Marasmius 201

On dead dicot wood in tropical rain forest and tropical montane forest, arising
from rhizomorphsor directly from the substratum.
MATERIAL STUDIED. BOLIVIA. Beni: Vaca Diez, Guayaramer'n, 18 Mar 1956, Singer B
2122 (LIL); La Paz: Nor-Yungas, Charobamba, 30 Jan 1956, Singer B 722 (LIL) type.
ILLUSTRATION. Singer (1965) pl 16, fig 92.

151d.Marasmiuspusio var graminivorusSinger, var nov

TYPE. Singer T 4058, from Argentina.
Haec varietas graminicola differt sporis 8-9.5 X 4.3-5,u stipiteque pallidiore.
This variety differs from variety guatopoensis with which it shares the spore shape
(7-)8-9.5 X (4-)4.3-5 ,, ellipsoid to broadly fusoid, in somewhat eccentric, orange pallid
stipe and habitat on grasses.
MATERIAL STUDIED. ARGENTINA. Jujuy: Lagunas de Yala at 2300-2400 m alt, 13 Feb
1965, Singer T 4058 (F), type.
Marasmius tener Berkeley & Curtis (Proc. Am. Acad. Sci. 41: 121. 1860) is certain-
ly similar but is said to be white.

Marasmius sect Sicci subsect Siccini ser Haematocephali Singer, ser nov
Cystidiis ad latera lamellarum bene evolutis. - Typus: Marasmius haematocephalus
(Montagne) Fries.
Pleurocystidia present.

Key to the Species of Series Haematocephali

Key I
1. All pleurocystidia in form of broom cells.
2. Broom cells of the sides of the lamellae restricted to a narrow region near the edge
proper(see series:Leonini).
2. Broom cells all over the sides of the lamellae.
3. Lamellae distant to very distant. 152. 31. setulosifolius.
3. Lamellae close to subclose. 153. M. pleuracanthus.
1. Broom cells restricted to the edges of the lamellae or their immediate neighborhood;
pleurocystidia metuloid or gloeocystidioid. not beset by apical setulae but sometimes
with one or two versiform appendages or outgrowths.
4. Pleurocystidia characteristically filled with distinct coarse oily contents; spores
11.5-20.5 X 2.7-4.5S.
5. Pileus broader than 15 mm; stipe > 1 mm broad; edges of lamellae not discolor-
ous or at least never brown (see "8" below).
5. Pileus 6-12 mm broad; stipe up to 0.8 mm broad; edges of lamellae often brown.
6. Through-lamellae 17-20; growing in tropical-montane forest. 156. M. oleiger.
6. Through-lamellae up to 15(-16).
7. Spores 16.8-20.5 X 3.2-4.5u; dried pileus between "tawny" and "cookie"
(MI&P);West Indies. 154. M. imusicola.
7. Spores 13.7-17.8 X 3.5-4.5,; dried pileus near "cocoa," ".Mohawk" or
"Mandalay" M&P; South America. 155. M. allocystis.
4. Pleurocystidia without coarse oleaginous contents (there may be minute oily droplets
in the contents which may make the cystidium appear very finely granular or else
contents honrogeneous); spores as above, or smaller.
8. Pileus white or whitish.
9. Lamellae mostly distant; pileus > 10 mm broad; cystidia conspicuous and gen-
erally numerous. 161. M. haedinus.
9. Lamellae medium close to distant; pileus 2-7 mm broad; cystidia few or mod-
erately conspicuous. 162. M. splitgerberi.
8. Pileus neither white nor whitish.
10. Pileus pink, red, purple, at least when young, not merely reddish brown,
reddish-orange-ferruginous) but it may or may not dry out to a dull brown
or cinnamon, orange-ferruginous or purplish brown.
202 FloraNeotropica

11. Pileus pale pinkish lilac, 5-7 mm broad; spores 11-17 X 3.2-4.5;' cystidia
broadly clavate or broadly clavate-ventricose. 32-43 X 10-15.2p; through-
lamellae 10-11. On dicotyledonous leaves and petioles. 158. Af.pallescens.
11. Pileus often deeper colored or broader, or else lamellae less or more
numerous or cystidia more variable. Both on leaves and other substrata.
12. Lamellae subclose to subdistant (about 17-20 through-lamellae);
spores 13-14.7 X 3.5-4.2u (if smaller see key II below). 150. 11.paner)ythnls.
12. Lamellae fewer; spores larger (14-22 long). 160. Ml.haematocephalus.
10. Pileus when young and fresh without a distinct red or purple tinge ( but may
be pinkish cinnamon). Key I1.

Key II
1. Lamellae subdistant to distant.
2. Spores larger than 13.2u or at least many of them larger.
3. Pileus over 15 mm broad or at least mostly reaching larger size when fully mature.
4. Edge of lamellae discolorous, interlamellar spaces not intervenose; spores
14.5-19 X 3-5.3p. 163. 1. h.vpophaeus.
4. Edge of lamellae not or scarcely discolorous.
5. Lamellae intervenose; spores 13.5-16 X 3-4.5 u; on rotten wood in montane
forests in Mexico. 164. M. guzmanianus.
5. Lamellae not or only slightly intervenose w hen mature or else not growing
on wood; spores as above or larger.
6. Spores 12-16 X 2.5-4,; cystidia poorly differentiated; on rotten wood
(see "16" below, but compare series A).
6. Spores generally reaching larger size (either longer or broader or both).
7. Growing on dead monocotyledonous and dicotyledonous leaves:
cystidia sometimes poorly differentiated (but numerous).
165. .1 montagneanus.
7. Growing on wood; cystidia always well differentiated and numerous.
8. Spores 28-37.5 X 3.5-5.5 p. 157. M. megistosporns.
8. Spores 14.5-19.5 X 3.4u.